The Peach

Botany, Production and Uses

This page intentionally left blank
 The Peach
Botany, Production and Uses

Edited by

Desmond R. Layne

Department of Horticulture, Clemson University, Clemson, South Carolina, USA

and

Daniele Bassi

Dipartimento di Produzione Vegetale, University of Milan, Milan, Italy

 CABI is a trading name of CAB International

CABI Head Office CABI North American Office

Nosworthy Way 875 Massachusetts Avenue
Wallingford 7th Floor
Oxfordshire OX10 8DE Cambridge, MA 02139
UK USA

Tel: +44 (0)1491 832111 Tel: +1 617 395 4056

Fax: +44 (0)1491 833508 Fax: +1 617 354 6875
E-mail: cabi@cabi.org E-mail: cabi-nao@cabi.org
Website: www.cabi.org

© CAB International 2008. All rights reserved. No part of this publication

may be reproduced in any form or by any means, electronically,
mechanically, by photocopying, recording or otherwise, without
the prior permission of the copyright owners.

A catalogue record for this book is available from the British Library,
London, UK.

Library of Congress Cataloging-in-Publication Data

The peach : botany, production and uses /edited by Desmond R. Layne

and Daniele Bassi.
p. cm.
Includes index.
ISBN 978-1-84593-386-9
1. Peach--Breeding. 2. Peach--Diseases and pests. 3. Peach--
Harvesting. I. Layne, Desmond R. II. Bassi, Daniele.

SB371.P42 2008
634’.25--dc22 2007045093
ISBN: 978 1 84593 386 9

Typeset by AMA Dataset Ltd, UK.

Printed and bound in the UK by Biddles, King’s Lynn.
 Contents

Dedication vii

Contributors x

Preface xiii

Acknowledgements xvi

1. Botany and Taxonomy 1

D. Bassi and R. Monet

2. History of Cultivation and Trends in China 37

H. Huang, Z. Cheng, Z. Zhang and Y. Wang

3. Classical Genetics and Breeding 61

R. Monet and D. Bassi

4. Genetic Engineering and Genomics 85

A.G. Abbott, P. Arús and R. Scorza

5. Low-chill Cultivar Development 106

B.L. Topp, W.B. Sherman and M.C.B. Raseira

6. Fresh Market Cultivar Development 139

W.R. Okie, T. Bacon and D. Bassi

7. Processing Peach Cultivar Development 175

T.M. Gradziel and J.P. McCaa

8. Rootstock Development 193

G.L. Reighard and F. Loreti

v
vi

9. Propagation Techniques 221

F. Loreti and S. Morini

10. Carbon Assimilation, Partitioning and Budget Modelling 244

T.M. DeJong and A. Moing

11. Orchard Planting Systems 264

L. Corelli-Grappadelli and R.P. Marini

12. Crop Load Management 289

R.P. Marini and G.L. Reighard

13. Nutrient and Water Requirements of Peach Trees 303

R.S. Johnson

14. Orchard Floor Management Systems 332

T.J. Tworkoski and D.M. Glenn

15. Diseases of Peach Caused by Fungi and Fungal-like Organisms: Biology,

Epidemiology and Management 352
J.E. Adaskaveg, G. Schnabel and H. Förster

16. Diseases Caused by Prokaryotes – Bacteria and Phytoplasmas 407

D.F. Ritchie, M. Barba and M.C. Pagani

17. Viruses and Viroids of Peach Trees 435

M. Cambra, R. Flores, V. Pallás, P. Gentit and T. Candresse

18. Insects and Mites 467

D.L. Horton, J. Fuest and P. Cravedi

19. Nematodes 505

A.P. Nyczepir and D. Esmenjaud

20. Preharvest Factors Affecting Peach Quality 536

C.H. Crisosto and G. Costa

21. Ripening, Nutrition and Postharvest Physiology 550

A. Ramina, P. Tonutti and B. McGlasson

22. Harvesting and Postharvest Handling of Peaches for the Fresh Market 575
C.H. Crisosto and D. Valero

Index 597

Colour plates 1–73 can be found following p. 144.

Colour plates 74–163 can be found following p. 336.
Colour plates 164–245 can be found following p. 496.
 Dedication

Richard E.C. Layne

Richard E.C. Layne served as Research
Scientist (1963–1996) and directed the
fruit breeding programme (1969–1996)
at the Agriculture and Agri-Food
Canada Research Centre at Harrow,
Ontario. While at Harrow, his scien-
tific research, new cultivar develop-
ment and outreach efforts significantly
impacted the Canadian tree fruit
industry and many other temperate,
fruit-growing regions around the
world. After growing up on the tropi-
cal island of St Vincent, West Indies, he
attended McGill University, where he
earned his BSc degree in Agronomy.
Next, he earned MS and PhD degrees
in Agronomy from the University, of
Wisconsin. His first research experi-
ence with fruit crops occurred when
he began his new job with Agriculture
Canada at Harrow in 1963.
During his tenure at Harrow he
was responsible for the introduction of
36 new fruit cultivars, ornamentals
and rootstocks. Specifically, these
included 15 peaches (ten cultivars,
three ornamentals and two rootstocks),
13 apricots (12 cultivars and one root-
stock), four nectarines and four pear
cultivars. In addition to Ontario, these cultivars play significant roles in the fruit crop indus-
tries of Michigan, New York, Pennsylvania, New Jersey and various European countries. This

vii
viii

is due, in part, to their improved bud and wood cold hardiness, superior disease resistance,
attractive colour, good eating quality and consistency of production.
In Ontario, his peach cultivars span the entire ripening season. Those that are currently
most widely planted include ‘Harrow Diamond’, ‘Harrow Beauty’, ‘Harson’, ‘Harrow Dawn’,
‘Harrow Fair’ and ‘Harblaze’ (nectarine). His apricots also span the ripening season in Ontario,
with ‘Harcot’ and ‘Harogem’ being those cultivars most widely planted. ‘Haroblush’, ‘Harojoy’,
‘Harostar’, ‘Hargrand’ and ‘Harval’ are becoming more widely planted now. His ‘Harlayne’
apricot with natural resistance to Plum pox virus has become a donor of this resistance trait in
conventional breeding programmes around the world. ‘Harrow Delight’, ‘Harrow Sweet’ and
‘AC Harrow Delicious’ pears all offer improved tolerance to fireblight.
In addition to his cultivar development work, his pioneering work on cold hardiness
physiology and applied work on scion/rootstock relationships, peach orchard management
and integrated production systems including irrigation have added significantly to the pomo-
logical scientific literature. This includes more than 130 publications and six book chapters.
He has received numerous research awards and served in important leadership roles during
his career. These include the Wilder Silver Medal (1996) awarded by the American Pomological
Society (APS) and Outstanding Researcher (1993) by the American Society for Horticultural
Science (ASHS). In 1992 he was elected a Fellow of ASHS. He received the Carroll R. Miller
award from ASHS in 1977, 1978, 1982 and 1985 for ‘excellence in research dealing with the
improved production and utilization of peaches’. He received the Paul Howard Shepard Award
for the best paper published by the APS in 1967 and 1982. He was President (1991/2) of the
APS and he was also President of the Canadian Society for Horticultural Science (1976/7). He
has been an international consultant to the USA, Brazil and China, and participated in an
exchange fellowship with INRA, France.
As a young boy, I fondly remember going with my dad to his research orchards and seeing
and tasting the results of his labours. I had the privilege of learning much about other cultures
and the importance of scientific collaboration and exchange while sitting at the dinner table at
our home as we hosted scientists visiting my dad from around the world. While in college
assisting him in his breeding and orchard management programmes at Harrow, I developed a
passion for pomological research and extension that has been the focus of my career. My dad
has been and continues to be a great friend, mentor and inspiration; and I fondly dedicate this
book to him.
Desmond R. Layne

Silviero Sansavini
Silviero Sansavini has been Professor of Pomology at the University of Bologna, Italy, since
1974. After growing up on a small fruit farm in the heart of one of the major fruit-growing areas
in northern Italy (the Romagna part of the Po Valley), he attended the University of Bologna,
where he earned a degree in Agricultural Science. His early career involved a stint as an exten-
sion phytopathologist, before being hired at the same university, where he went through the
ranks to Full Professor, and served as Chairman of the Department of Fruit and Woody Plant
Sciences almost continuously from 1977 to the end of 2007.
A ‘man with a mission’, he has promoted the art and science of fruit growing throughout
his career by leading many nationally funded research projects and participating in many col-
laborative international ones, including several funded by the EU. As a natural consequence of
this commitment and activity, he has been convener and/or organizer of well over 100 scien-
tific meetings or extension events at the international and national levels. He was elected
Chairman of the International Society for Horticultural Science in 1994 and in his 4-year tenure
he organized a signal event for horticulture: The World Congress on Horticultural Research
(Rome, 1998), the first joint venture carried out by the International Society for Horticultural
Science and the American Society for Horticultural Science (ASHS), which marked the first
 ix

time that horticulturists from all over

the globe had convened to discuss polit-
ical and economic aspects related to
worldwide horticultural research.
He has received many awards rec-
ognizing his dedication to horticulture:
from the National Canners Association
Award (assigned by the ASHS for his
work on ‘ripening of nectarines and can-
ning peaches’) to the Wilder Medal 2000
awarded by the American Pomological
Society, the Gold Veitch Memorial Medal
from the Royal Horticultural Society
(London, UK) and an Honorary Degree
in Horticultural Sciences from the Uni-
versity of Budapest, Hungary. At the
national level, he has been recognized
by the Italian Ministry of Higher Educa-
tion. He was named ASHS Fellow in
1995.
His editorial activity is tireless. He
has been Editor since 1986 of the monthly
Italian journal Rivista di Frutticoltura,
and is an editorial board member of the
following journals: Tree Fruit Production
(Binghamton, New York, USA), Fruits
(CIRAD, Montpellier, France),
L’Arboriculture Fruitière and the Interna-
tional Journal of Agronomy, Agricultural
Ecosystem and Plant Genetics (INRA,
Montfavet, France). Throughout his career he has edited the proceedings of innumerable sym-
posia and meetings at the international and national levels.
He is also Member of several academies: the Italian National Academy of Agriculture, the
Italian Academy of Science, a Corresponding Member of the Academie d’Agriculture of France,
the Italian Academy of Grape and Wine and the prestigious Academy of ‘Georgofili’ (Florence,
Italy).
He is author, co-author or editor of more than 500 papers, proceedings and books on bio-
logical, physiological and genetic aspects of pome and stone fruits. His in-depth command of
very diverse facets of horticultural research is witness to his passionate commitment to horti-
culture as a whole, his scientific curiosity and his hard-working and demanding attitude,
which are the unique traits that have attracted hundreds of students from Italy and abroad to
work with him. His students, many of whom have followed his path in research, value the
privilege of working with one of the most dedicated, open-minded and challenging personali-
ties of the international fruit science community.
Daniele Bassi
Luca Corelli-Grappadelli
 Contributors

Abbott, Albert G., Department of Genetics and Biochemistry, 116 Jordan Hall, Clemson University,
Clemson, SC 29634, USA.
Adaskaveg, James E., Department of Plant Pathology and Microbiology, University of California –
Riverside, 242 Fawcett Laboratory, Riverside, CA 92521, USA.
Arús, Pere, Institut de Recerca i Tecnologia Agroalimentàries (IRTA), E-08348 Cabrils (Barcelona),
Spain.
Bacon, Terry, Sun World International, Inc., 16350 Driver Road, Bakersfield, CA 93308, USA.
Barba, Marina, Instituto per la Patologia Vegetale, Via Bertero 22, I-00156 Rome, Italy.
Bassi, Daniele, Dipartimento di Produzione Vegetale, University of Milan, Via Celoria 2, I-20133
Milan, Italy.
Cambra, Mariano, Departamento Protección Vegetal y Biotecnología, Instituto Valenciano de Investiga-
ciones Agrarias (IVIA), Carretera Moncada – Náquera Km 4.5, Apartado Oficial, E-46113 Moncada
(Valencia), Spain.
Candresse, Thierry, Plant–Virus Interactions, UMR–GDPP–IBVM, INRA Bordeaux-Aquitaine, BP
81, F-33883 Villenave d’Ornon Cedex, France.
Cheng, Zhongping, Wuhan Institute of Botany, The Chinese Academy of Sciences, Hubei 430074,
People’s Republic of China.
Corelli-Grappadelli, Luca, Dipartimento di Colture Arboree, University of Bologna, Via Fanin 46,
I-40127 Bologna, Italy.
Costa, Guglielmo, Dipartimento di Colture Arboree, University of Bologna, Via Fanin 46, I-40127
Bologna, Italy.
Cravedi, Piero, Instituto di Entomologia e Patologia Vegetale, Piacenza Campus, Università Cattolica
del Sacro Cuore, Via Emilia Parmense 84, I-29100 Piacenza, Italy.
Crisosto, Carlos H., Department of Plant Sciences, University of California, Kearney Agricultural
Center, 9240 South Riverbend, Parlier, CA 93648, USA.
DeJong, Theodore M., Department of Plant Sciences, 3037 Wickson Hall, One Shields Avenue,
University of California, Davis, CA 95616-8683, USA.
Esmenjaud, Daniel, INRA, UMR ‘Interactions Biotiques et Santé Végétale’ (IBSV), 400 Route des
Chappes, F-06560 Sophia-Antipolis Cedex, France.
Flores, Ricardo, Instituto de Biología Molecular y Celular de Plantas, CSIC/Universidad Politécnica
de Valencia, Avenida de los Naranjos s/n, E-46022 Valencia, Spain.

x
 xi

Förster, Helga, Department of Plant Pathology, University of California, One Shields Avenue, Davis,
CA 95616, USA.
Fuest, Jaime, Department of Entomology, University of Georgia, 463 Biological Sciences Building, 120
Cedar Street, Athens, GA 30602-2603, USA.
Gentit, Pascal, Virology Laboratory, CTIFL, BP21 Lanxade, F-24130 La Force, France.
Glenn, D. Michael, USDA–ARS Appalachian Fruit Research Station, 2217 Wiltshire Road,
Kearneysville, WV 25430, USA.
Gradziel, Tom M., Department of Plant Sciences, 2055 Wickson Hall, One Shields Avenue, University
of California, Davis, CA 95616-8683, USA.
Horton, Dan L., Department of Entomology, University of Georgia, 463 Biological Sciences Building,
120 Cedar Street, Athens, GA 30602-2603, USA.
Huang, Hongwen, Wuhan Institute of Botany/Wuhan Botanical Garden, The Chinese Academy of
Sciences, Hubei 430074, People’s Republic of China and South China Institute of Botany/South
China Botanical Garden, Guangzhou 510650, People’s Republic of China.
Johnson, R. Scott, Department of Plant Sciences, University of California, Kearney Agricultural
Center, 9240 South Riverbend, Parlier, CA 93648, USA.
Layne, Desmond R., Department of Horticulture, Clemson University, 50 Cherry Road, Clemson, SC
29634-0319, USA.
Loreti, Filiberto, Dipartimento di Coltivazione e Difesa delle Specie Legnose ‘G. Scaramuzzi’, Sezione
di Coltivazioni Arboree, Facoltà di Agraria, University of Pisa, Via del Borghetto 80, I-56124 Pisa,
Italy.
McCaa, James P. (‘Pat’), Del Monte Foods, PO Box 30190, 2716 East Miner Avenue, Stockton, CA
95213, USA.
McGlasson, William (‘Barry’), Center for Plant and Food Science (PAFS), University of Western
Sydney, Hawkesbury Campus, Locked Bag 1797, South Penrith Distribution Centre, NSW 1797,
Australia.
Marini, Richard P., Department of Horticulture, The Pennsylvania State University, 119 Tyson Building,
University Park, PA 16802, USA.
Moing, Annick, Unite de Physiologia Vegetable, INRA Domaine de la Grande Ferrade, 71 avenue
Edouard Bourlauz, BP 81, F-33883 Villenave d’Ornon Cedex, France.
Monet, René (retired), National Institute for Agronomical Research (INRA), Bordeaux Research
Centre, Bordeaux, France.
Morini, Stefano, Dipartimento di Coltivazione e Difesa delle Specie Legnose ‘G. Scaramuzzi’, Sezione
di Coltivazioni Arboree, Facoltà di Agraria, University of Pisa, Via del Borghetto 80, I-56124 Pisa,
Italy.
Nyczepir, Andy, USDA–ARS Southeastern Fruit and Tree Nut Research Laboratory, 21 Dunbar
Road, Byron, GA 31008, USA.
Okie, William R. (‘Dick’), USDA-ARS Southeastern Fruit and Tree Nut Research Laboratory, 21
Dunbar Road, Byron, GA 31008, USA.
Pagani, Maria Cristina, BASF Corporation, 26 Davis Drive, Research Triangle Park, NC 27709,
USA.
Pallás, Vincente, Instituto de Biología Molecular y Celular de Plantas, CSIC/Universidad Politécnica
de Valencia, Avenida de los Naranjos s/n, E-46022 Valencia, Spain.
Ramina, Angelo, Dipartimento di Agronomia Ambientale e Produzioni Vegetali, University of Padova –
Agripolis, Viale dell’Università 16, I-35020 Legnaro (PD), Italy.
Raseira, Maria do Carmo Bassols, Empressa Brasileira de Pesquisa Agropecuaria/Embrapa Clima
Temperado, Caixa Postal 403, 96001-970 Pelotas, Rio Grande do Sul, Brazil.
Reighard, Gregory L., Department of Horticulture, Clemson University, 50 Cherry Road, Clemson,
SC 29634-0319, USA.
Ritchie, David F., Department of Plant Pathology, North Carolina State University, 2406 Gardner
Hall, Campus Box 7616, Raleigh, NC 27695-7616, USA.
xii

Schnabel, Guido, Department of Entomology, Soils and Plant Sciences, Clemson University,
Clemson, SC 29634-0315, USA.
Scorza, Ralph, USDA–ARS Appalachian Fruit Research Station, 2217 Wiltshire Road, Kearneysville,
WV 25430, USA.
Sherman, Wayne B., Department of Horticultural Sciences, 717 Hull Road, 2133 Fifield Hall, PO Box
110690, University of Florida, Gainesville, FL 32611-0690, USA.
Tonutti, Pietro, Scuola Superiore Sant’Anna, Piazza Martiri della Libertà 33, I-56127 Pisa, Italy.
Topp, Bruce L., Department of Primary Industries and Fisheries, Agency for Food and Fibre Services,
Queensland Horticulture Institute, Maroochy Research Station, PO Box 5083 SCMC, Nambour,
QLD 4560, Australia.
Tworkoski, Thomas J., USDA-ARS Appalachian Fruit Research Station, 2217 Wiltshire Road,
Kearneysville, WV 25430, USA.
Valero, Daniel, Department of Food Technology, Head of Post-Harvest and Quality, University
Miguel Hernández, Ctra. Beniel – km. 3,2, E-03312 Orihuela, Alicante, Spain.
Wang, Ying, Wuhan Institute of Botany, The Chinese Academy of Sciences, Hubei 430074, People’s
Republic of China.
Zhang, Zhonghui, Wuhan Institute of Botany, The Chinese Academy of Sciences, Hubei 430074,
People’s Republic of China.
 Preface

Before the 19th century, many non-Asians believed that the peach originated from Persia
(modern-day Iran). Peach probably came to Persia from China along the silk trading routes in
the 2nd or 3rd century BC. The Persian origin hypothesis was due, in part, to the fact that
peaches were brought from Persia to Europe by the Roman army in the 1st century BC. Alphonse
De Candolle in his ‘Origin of Cultivated Plants’ (1885, Appleton and Co., New York, pp. 221–229)
contended that peach originated in China. In his tome, ‘The Peaches of New York’ (1917, J.B. Lyon
Co., Albany), U.P. Hedrick made the same assertion. In fact, Chinese literature refers to peach
more than 1000 years before it first appeared in any European writings. There is documented
evidence of peach cultivation in China for more than 3000 years ago.
The peach is a symbol of immortality in Taoist mythology. The Queen Mother (goddess)
of the West (Xi Wang Mu) had a jade palace that was surrounded by a beautiful garden con-
taining the peach trees of immortality. In the classic Chinese novel, ‘The Journey to the West’
(Wu, Ch’eng-en ~1590 AD, translated by Anthony C. Yu), Sun Wukong, or the Monkey King
(picture inset), attained immortality as a result of a memorable visit to this garden:

‘I have been authorized by the Jade

Emperor,’ said the Monkey King, ‘to look
after the Garden of Immortal Peaches.’ The
local spirit hurriedly saluted him and led
him inside. The Monkey King then asked
the local spirit, ‘How many trees are there?’
‘There are three thousand six hundred,’ said
the local spirit. ‘In the front are one
thousand two hundred trees with little
flowers and small fruits. These ripen once
every three thousand years, and after one
taste of them a man will become immortal
with healthy limbs and a lightweight body.
In the middle are one thousand two
hundred trees of layered flowers and sweet
fruits. They ripen once every six thousand
years. If a man eats them, he will ascend to
Heaven with the mist and never grow old.

xiii
xiv

At the back are one thousand two hundred trees with fruits of purple veins and pale yellow pits.
These ripen once every nine thousand years and, if eaten, will make a man’s age equal to that of
Heaven and Earth, the sun and the moon.’ Highly pleased by these words, the Monkey King made
thorough inspection of the trees and a list of the arbors and pavilions before returning to his
residence. One day he saw that more than half of the peaches on the branches of the older trees had
ripened, and he wanted very much to eat one and sample its novel taste. Closely followed, however,
by the local spirit of the garden, the stewards, and the divine attendants, he found it inconvenient to
do so. He therefore devised a plan on the spur of the moment and said to them, ‘Why don’t you all
wait for me outside and let me rest a while in this arbor?’ The various immortals withdrew
accordingly. The Monkey King then took off his cap and robe and climbed up onto a big tree. He
selected the large peaches that were thoroughly ripened and plucking many of them, ate to his
heart’s content right on the branches.

Peach is revered as a delicious and healthy summer fruit in most temperate regions of the
world. It is highly perishable but, depending on market demands and fruit availability, it can
be a very profitable fruit crop for the careful farmer. Today it is a major fruit crop of commerce
in China, Italy, Spain, the USA, and Greece, the top five producing countries, respectively.
Currently, there are nearly 1.5 million ha of peaches in production worldwide with the vast
majority planted in China (approx. 46%).
Tremendous diversity exists within the cultivated peach germplasm for tree size, growth
habit, flower size and colour, chill hour requirement, fruit size, shape, flesh texture, flesh colour,
flesh acidity, stone adherence to flesh, etc. As a result, many hundreds of cultivars of peaches
are grown successfully from climatic and geographic regions as diverse as southern Canada to
the highlands of Thailand.
Because of the small genome size (about twice that of Arabidopsis), peach has been selected
as a model species for studying genomics in the Rosaceae. An extensive physical map/genetic
map has been developed and vast genetic information is available through an online database.
Marker-assisted selection in conjunction with conventional breeding techniques will undoubt-
edly lead to new cultivars with enhanced pest resistance, nutritional value and other novel
traits. Until a reliable transformation and regeneration protocol can be developed, however,
some of the novel genetic advances that have occurred in other fruits and crop plants remain
to be fully realized.
Management of light interception, careful pruning and training, orchard floor, soil fertility,
water availability and crop load are just a few of the many complexities of producing peaches
profitably. Combining these factors with the reality that peach is subject to many difficult to
manage pests makes it even more of a challenge. Breeding for pest resistance has resulted in
some improvements in tolerance to a few diseases and insects, but if trees are left unattended
in most locations, they will die prematurely. Thus, careful rootstock and cultivar selection,
combined with proper site selection and pest management (monitoring, forecasting, thresh-
olds, cultural and physical controls, biological and chemical control), are vital for success.
This comprehensive treatise by 49 research scientists from eight countries summarizes the
current state of knowledge in topics ranging from botany and taxonomy to breeding and genet-
ics of cultivars and rootstocks, propagation, physiology and planting systems, crop and pest
management and postharvest physiology. The goal was to provide research scientists, exten-
sion personnel, students, professional fruit growers and others with a vital resource on peach
and its culture.
Desmond R. Layne
Daniele Bassi
 xv

Des Layne Daniele Bassi

 Acknowledgements

The Editors would like to acknowledge the chapter authors and the following individuals for
critical external peer review of chapter manuscripts in this text: T. Beckman, W. Bentley, D.
Byrne, J. Clark, J. Cline, K.R. Day, R. Ebel, L. Georgi, J. Girona, R. Gucci, F. Hale, A. Iezzoni,
R.E.C. Layne, S. Lewis, A. Liverani, N.W. Miles, A. Naor, M.L. Parker, J.C. Pech, J. Rushing, M.
Rieger, E. Sanchez, S. Scott, W. Shane, H. Sherm, C. Xiloyannis and J. Walgenbach.
The Editors would also like to acknowledge the following individuals and organizations
that helped underwrite the expenses for the colour plate section of the book. In the USA, these
included Dr Norman F. Childers, National Peach Council (USA), Al Pearson, Adams County
Nursery, the Burchell Nursery, Inc., South Carolina Agricultural Experiment Station, South
Carolina Peach Council, Titan Peach Farms and Jerrold A. Watson & Sons. In Italy, these
included Apofruit, Apoconerpo, Orogel Fresco, Pempacorer, the Growers Consortium for Crop
Development (Newplant), the Crop Research Center (CRPV), the Apulian Nursery Consor-
tium (COVIP) and the Italian Society for Horticultural Science (SOI).

xvi
 1 Botany and Taxonomy

Daniele Bassi1 and René Monet2

1University
of Milan, Milan, Italy
2National Institute for Agronomical Research (INRA), Bordeaux Research Centre,
France (retired)

1.1 Introduction: Origin and Dissemination of Peach 1

1.2 Species Systematics: Cultivated Peach and Wild Relatives 2
1.3 Peach Morphology: Description and Variability of the Main Organs 3
Tree 3
Leaf 6
Flower and fruit development 11
Fruit appearance and composition 16
Endocarp (stone, pit) and seed 25
1.4 Peach Biology and Phenology 25
Floral biology and fruit set 25
Chilling and heat requirements 26
Phenological phases 27
Time of flowering 27
Time of ripening (fruit development period) 27
1.5 Cultivar Classification 29
Peach description sheet 29
Morphological and commercial classifications 30
Phenological classification 30

1.1 Introduction: Origin and
Dissemination of Peach
The botanical name of peach (Prunus persica
(L.) Batsch) refers to the putative country of
origin, Persia (actual Iran), and Linné (1758)
first named the species based on this opinion
(Amygdalus persica). Only in the 19th century
was the Far East geographical origin (western
China) finally acknowledged (De Candolle,
1883; Hedrick, 1917; Vavilov, 1951); written
records and archaeological evidence date peach
domestication at least as far back as 3000 BC
(Li, 1984).
Faust and Timon (1995) gave a detailed
account of the possible genetic and geo-
graphic origin and dissemination of the peach
and related species. Taking into consider-
ation the long history of cultivation of this
species and its growing role in several coun-
tries over the centuries, many scholars have
attempted the classification of the species

© CAB International 2008. The Peach: Botany, Production and Uses

(eds D.R. Layne and D. Bassi) 1
2 D. Bassi and R. Monet
and its related forms. Early writings classi-
fied almond and peach under the same spe-
cies, but later they were classified as separate
entities (De Candolle, 1883) although possi-
bly sharing the same putative ancestor. The
close geographic origin may account for this
hypothesis: while peach is native to the
Tarim basin north of the Kun Lun moun-
tains, almond is native to the south of the
same mountains (beside the northern borders
of Afghanistan and Pakistan).
The westward movement of peach could
have brought it into Persia (actual Iran) in the
2nd to 1st century BC, shortly before the
arrival of the Roman army. Early Latin schol-
ars mentioned peaches in Italy in the 1st cen-
tury BC. The ‘Gallic’ peaches, described of
French origin, may account for a possible sec-
ond, independent arrival of the tree to Europe.
Thus, peach could have reached France through
the Balkan route almost simultaneously to its
arrival in Italy, along the Danube river and
the Black Sea region (Werneck, 1956). Addi-
tionally, in the Middle Ages, France became
probably the second major point of origin
after China, according to many records. Peach
was brought to the American continent in two
different waves. The first was led by the Span-
iards, starting in Central America from the
first half of the 16th century. In northern
America, peaches were later cultivated by the
native peoples and propagated by seeds. This
early introduction in the Americas has given
rise to many landraces, most featuring the
non-melting flesh. Some are still cultivated
for local fresh markets or even exploited as
valuable sources for interesting traits such as
‘evergreen’ (Diaz, 1974) and disease resistance,
e.g. powdery mildew or rust (Rodriguez and
Sherman, 1990; Pérez et al., 1993). Interestingly,
even today, non-melting local cultivars for the
fresh market are very popular in southern
Europe (Greece, southern Italy and Spain).
The second wave of peach introduction
in the USA was a direct import from China in
the mid-1850s, when the well-known ‘Chinese
Cling’ was grown at the Delaware Experi-
mental Station. This tree originated ‘Elberta’,
one of the main ancestors of the modern culti-
vars grown in the USA and elsewhere in the
most important peach-growing countries
(Okie et al., 1985; Scorza et al., 1985).
1.2 Species Systematics: Cultivated
Peach and Wild Relatives
The argument about the scientific name of
peach was finally set by the classification of
Bailey (1927) that grouped all stone fruits
under the Prunus genus. Hedrick (1917) stated
as not rational a classification of peach
species based almost exclusively on the mor-
phological traits of the fruits. Consequently,
the species classification based on fruit traits
(shape, glabrous skin) or tree growth habit is
to be regarded as an early attempt, before
Mendel’s laws enlightened the mere intra-
specific nature of those singly inherited
traits. The following classification is summa-
rized after Knight (1969), Watkins (1976) and
Rehder (1990). Peach is included in the
Euamygdalus Schneid. section of the Amygdalus
subgenus, and can be distinguished from
almond (Prunus dulcis (Mill.) D.A. Webb)
because the mesocarp of the latter becomes
dry and splits at maturity, while the leaves
are serrulate.
P. persica (L.) Batsch is a diploid species
(2n = 16) with a medium tree height (up to 8
m); the leaves are lanceolate, glabrous and
serrate, broadest near the middle, with a glan-
dular petiole; the flowers are generally pink,
but also white or red; the fruit is pubescent or
glabrous, fleshy and the mesocarp does not
split; the stony endocarp is very deeply pitted,
furrowed and very hard.
The main species related to peach are
listed below; all of them show fruits with very
poor eating quality, although some could be
interesting for disease resistance traits or as
rootstocks.
Prunus davidiana (Carr.) Franch. is a wild
species native to north-eastern China, where
it is used as a seedling rootstock, given its tol-
erance to drought, although it is very sensi-
tive to nematodes; the tree is tall (up to 10 m),
with a reddish-brown bark; the leaves are
long and glabrous, ovate-lanceolate, broadest
near the base; the flower is white or light pink;
the pit is small and pitted; the flesh is freestone.
Accessions of this species have been hybrid-
ized with peach to improve disease resistance
on scion cultivars to plum pox, powdery
mildew, leaf curl, etc. (Moing et al., 2003) or to
 Botany and Taxonomy
breed interspecific rootstocks adaptable to
marginal soils or to replant problems (Pisani
and Roselli, 1983; Roselli et al., 1985; Edin and
Garcin, 1994).
Prunus ferganensis (Kost. and Rjab) Kov.
and Kost. is a wild form found in western
China classified as a subspecies of P. persica; a
wide variability in terms of fruit types can be
found (yellow- and white-fleshed peaches, nec-
tarines, etc.); the leaves have parallel veins and
there are parallel grooves in the stone, both sin-
gle Mendelian traits (Okie and Rieger, 2003).
The seed can be cyanogenic glycoside-free (not
bitter). It has resistance to powdery mildew.
Prunus kansuensis Rehd. is a wild species
found in north-east China, also used as a
seedling rootstock in China; it is a bushy tree
with glabrous winter buds, early-blooming,
even so the flowers are considered to be rather
resistant to frost (Meader and Blake, 1939);
the leaves are villous along the midrib near
the base, broadest below the middle; the style
is longer than the stamens. The fruit quality is
very poor (astringent); the stone is furrowed
(with parallel grooves) but not pitted.
Prunus mira Koehne is a wild species
found in far-west China (eastern Tibet); the
tree is tall (up to 20 m) and long-living, up to
1000 years (Wang, 1985); the leaves are lance-
olate, villous along the midrib beneath,
rounded at base; the flowers are white. The
fruit is very variable in shape, colour and size;
the stone is smooth, although in some types it
resembles P. persica. Some forms are culti-
vated in Tibet and it is also used as a seedling
rootstock in some regions of India. It is
reputed as an ancestor of P. persica, having
been spread south and east from the Himala-
yan mountains (Yoshida, 1987).
1.3 Peach Morphology: Description and
Variability of the Main Organs
Tree
The trunk is straight and smooth, with a red-
dish-greenish bark in the first year, later
becoming dark grey-silver.
The root system develops within 50–60
cm in depth, depending on soil type; roots are
3
orange-white while young, turning dark orange
when older with large lenticels. The tree can
live for 20–30 years, but in commercial plant-
ings the average duration is limited to 12–15
years at most, because of either the cultivar
becoming obsolete or the loss of productivity.
Fruit production begins from the second or
third year.
Fruiting wood and buds
One-year-old shoots are reddish-green, turn-
ing dark grey-silver when older. Buds are
found at the base of the leaves. Each node nor-
mally displays three buds: two lateral flower
buds and one vegetative bud in the middle,
but up to four or five flower buds can be found
and sometimes even more, particularly in the
ornamental types. Sometimes only one flower
bud is present beside the vegetative, or even
only three vegetative buds. There is variability
in setting flower buds, where the genotypes
with high bud set transmit the character to the
progeny with high heritability (Weinberger,
1944); the resulting seedlings tend to be preco-
cious (Rodriguez and Sherman, 1986).
Different types of shoots can be found,
with different contribution to yield and fruit
quality.
● One-year-old shoot: wood of good vigour
(50–100 cm) with flower buds along its
axis, which ends with a vegetative bud
(Fig. 1.1/Plate 1). It is the most important
wood in commercial cultivars, since in
peach fruit size is positively related to
shoot vigour and total current-season
wood available (Manaresi and Draghetti,
1915; Marini and Sowers, 1994). Flower
bud set may be uneven along the shoot
and cultivars may be classified according
to basal, middle or apical distribution
(Bellini and Scaramuzzi, 1976).
● Brindle: a 1-year-old fruiting shoot of weak
vigour (about 10–25 cm) with flower buds
along its axis, which ends with a vegetative
bud. This is the most important wood
(‘hangers’) for canning peaches, where me-
dium to small-sized fruits are most prized.
● Feather: side shoot arising from a bud in
the same year as that when the bud was
formed.
4 D. Bassi and R. Monet
Fig. 1.1. One-year-old fruiting shoots at bloom.
● Spur (or dard): a very short (about 1–3
cm), 1-year-old shoot that terminates in a
vegetative bud (vegetative spur), even-
tually surrounded by one or more flower
buds (fruiting spur).
● Water sprout: wood of high (excessive)
vigour, with no flower buds.
● Branch: a limb older than 4–5 years.
Okie (1998) describes a ‘collar’ trait as a swol-
len collar growing where the shoot attaches to
the trunk, resulting in knobs when the shoot
is eventually eliminated. The trait seems
monogenic and recessive.
INTERNODE LENGTH AND DWARFISM. Internode
length is influenced by both quantitative and
qualitative Mendelian traits. Tree size, however,
is predominantly influenced by qualitative
traits, independently of the growth habit (see
section on ‘Tree growth habit’ below), particu-
larly in the case of the dwarf phenotypes, where
internodes are less than 10 mm in length (Fig.
1.2/Plate 2). The regular dwarf tree also features
a dense canopy due to larger and thicker leaves.
The first dwarf phenotype was described by
Lammerts (1945) from the ‘Chinese Dwarf’
peach as recessive and monogenic (Dw/dw). He
also described bushy trees obtained in a ratio of
15:1 by self-pollinating the ‘Babcock’ peach as
recessive and controlled by two genes (Bu1/bu1,
Bu2/bu2).
Hansche (1988) described a more size-
reducing dwarf trait than Dw/dw in a prog-
eny from ‘Redcal’ peach (heterozygous for
that trait) and reported it as monogenic and
recessive (Dw2/dw2). Chaparro et al. (1994)
described an even smaller dwarf (Dw3/dw3),
with a very thin stem and willow-like leaves.
Monet and Salesses (1975) described one
more dwarf mutant, smaller than Dw/dw.
Heterozygous individuals are semi-dwarf; by
self-pollination, these individuals give three
phenotypes: normal, semi-dwarf (like the F1
parent) and dwarf with Mendelian propor-
tions 1:2:1. It is a monogenic trait with incom-
plete dominance (N/n).
Gradziel and Beres (1993) reported a
similar semi-dwarf as above, featuring
internode length half of the standard size,
with thicker shoots and greener leaves. The
tree is upright but open and with 1-year-old
shoots from 30 to 50% shorter than standards.
When crossed to standard growth habit geno-
types it shows an intermediate heritability
(1:1).
 Botany and Taxonomy
Fig. 1.2. Dwarf peaches in a commercial orchard.
Okie (1998) described ‘mini-pillar’, a tree
form growing upright with wavy twigs where
the internode is about half the normal length.
TREE GROWTH HABIT. The tree growth habit
(or tree form) can be defined as the overall ap-
pearance of a tree’s canopy, i.e. the whole set
of a tree’s vigour, which could be expressed
as the total amount of dry matter or by the
rates of shoot growth (quantitative traits) and
structural (type of fruiting shoots and their
distribution through the canopy, their angle
and internode length) traits that determine its
natural architecture (Bassi, 2003). For a better
understanding of the canopy architecture,
two types of branch angle should be taken in
consideration: (i) the ‘crotch’ angle, formed
by the 1-year-old fruiting shoot (basal part)
and the limb from which it grows; and (ii) the
‘extension’ angle, formed by the reference
axis (trunk or branch from which the 1-year-
old fruiting shoot grows) and the juncture
line between the shoot’s point of origin and
its apex at full growth (Fig. 1.3/Plate 3). This
latter parameter makes it possible to account
for the direction of the shoot’s distal part,
which, regardless of the crotch angle, can be
vertical, upright, outwards (deviating more or
less from the vertical), weeping or procumbent
5
(Scorza, 1984, 2002; Scorza et al., 1989, 2002;
Bassi et al., 1994; Bassi, 2003).
Several growth habits are known in
peach (see a comprehensive list in Chapter 3,
Table 3.1).
● Arching: similar to the upright (see be-
low), but with a distinct curvature in the
1-year-old shoots (Werner and Chaparro,
2005).
● Columnar: marked by branches and
shoots growing vertically at a notably nar-
row crotch angle (no more than 35–40°)
that makes the tree look like a column (or
‘pillar’).
● Compact: marked by a large number of
lateral shoots and relatively short inter-
nodes that produces a dense canopy. The
overall shape is semi-spherical for a
bush-like appearance. A genetic curiosity,
‘corky triangle’, has been observed in a
progeny resulting from a self-pollination
of ‘Redhaven’ peach (Monet and Bastard,
1982). A triangular ‘corky’ (necrotic) zone
develops close to most buds on the epi-
dermis of the shoot. This character is as-
sociated with a lower size of the tree due
to a short internode. The ‘Compact Red-
haven’ peach studied by Van Well (1974),
6 D. Bassi and R. Monet
Fig. 1.3. Relationship between crotch (α) and
extension angles (β). (From Bassi, 2003.)
Fideghelli et al. (1979) and Mehlenbacher
and Scorza (1986) shows this character.
It is possible that the ‘corky triangle’ re-
cessive and monogenic (T/t) trait and
‘compact’ tree characters are under pleio-
trophic control of the same mutation.
● Open: overall tree shape varying from
goblet to slightly flat (transversal diame-
ter more developed than the vertical).
The 1-year-old fruiting shoots are marked
by a wide crotch angle (about 60–70°).
● Spreading: intermediate between open
and weeping.
● Spur: marked by the prevalence of spurs;
the internodes can be shorter than nor-
mal, in which case the canopy size could
be semi-dwarf or dwarf.
● Standard: the prevailing growth habit in
commercial cultivars; it is marked by me-
dium crotch angle (40–60°) and internode
length; canopy shape is generally semi-
spherical or slightly upright (the cano-
py’s vertical diameter is more developed
than the cross-section’s).
● Twister: a unique branching pattern where
after 15–30 cm of growth of the shoot, the
phyllotaxy increases from normal 135° to
about 180°, with buds and leaves aligned
at the opposite sides of the stem, on the
same plane; after 15–20 nodes the shoot
stops growing and normal shoots break
from buds below, repeating the same pat-
tern, i.e. normal first, then twisted; the trait
seems monogenic and recessive (Okie,
1998).
● Upright: the canopy is more developed
in height than in width and is larger (in
diameter) than the columnar. One-year-
old shoots and branches are marked by a
relatively narrow crotch angle (about
50°) and vertical growth.
● Weeping: marked by shoots of medium
or wide crotch (larger than 70°) and ex-
tended (more than 90°) angles, bending
downward (positive geotropic growth),
the result being a profile that is goblet-
like or more developed in width (Monet
et al., 1988).
The main growth habits are sketched in
Fig. 1.4 and summarized in Table 1.1.
Leaf
For peach, new shoots and leaves form follow-
ing anthesis. Two temporary lateral stipules
are found at the petiole base; they abscise when
the leaf is fully developed (Fig. 1.5/Plate 4).
 Botany and Taxonomy
Fig. 1.4. Main growth habit in peach: (a) standard; (b) columnar; (c) upright; (d) compact;
(e) weeping; (f) open (from Bassi, 2003).
The leaf blade may be flat or wavy. The
wavy blade results from a differential growth
of the leaf margins and this is recessive and
monogenic (Wa/wa; Scott and Cullinan, 1942);
it also has very sharply serrated margins. This
leaf character was reported as often associated
to a dwarf tree growth habit, with poor set.
A narrow-width leaf (willow-like shape)
with very serrated margins was reported by
Chaparro et al. (1994). This phenotype, desig-
nated as Wa2/wa2 and genetically linked to
the dwarf gene Dw3, does not have the
wavy leaf edges as the original homozygous
wa/wa trait, other than showing a narrow
width.
One more narrow-leaf trait was reported
by Okie and Scorza (2002), partially dominant
over standard shape (probably several genes are
involved), the tree size being sometimes smaller
and weaker than standard (Fig. 1.6/Plate 5).
Progenies segregate for variable width:length
ratio from 10 (narrow) to 0.25% (standard
7
shape). Narrow leaves show higher water use
efficiency than standard (Glenn et al., 2000).
There are glands at the margin of the leaf,
located at the blade base and at the petiole;
three phenotypes are known that show Men-
delian inheritance with incomplete dominance,
the absence being recessive (E/e; Connors,
1921): (i) reniform (homozygous dominant);
(ii) globose (heterozygous); and (iii) eglandu-
lar (homozygous recessive) (Fig. 1.7/Plate 6).
The leaves with reniform (kidney-shaped)
glands have crenate margins, varying in num-
ber from two to eight or more, those located
on the margin being smaller than those on the
petiole.
The leaves with globose glands have
serrate-crenate margins, but glandless leaves
with serrate margins are frequently found on
the same tree; the glands are round, smaller
than reniform, varying in number from none
to over eight, those on the petiole being
stalked.
 8
Table 1.1. Characteristics of main growth habits in peach. (Data from a comparative trial in Italy, trees not pruned; from Bassi, 2003.)

Canopy Angle (degrees from vertical)

Tree form Tree size Trunk cross-sectional Internode

D. Bassi and R. Monet

(by crotch angle) (height) area (cm2) Height (m) Width (m) Crotch (°) Extended (°) length (mm)

Columnar Very tall 154 ± 16 5.0 ± 0.5 1.5 ± 0.33 36.8 ± 7.9 30.8 ± 11.5 19.1 ± 2.9
Upright Very tall 183 ± 12 5.0 ± 0.4 2.5 ± 0.3 49.6 ± 6.7 41.6 ± 6.5 25.1 ± 2.9
Standard Medium (regular) 180 ± 3.9 3.6 ± 0.3 2.8 ± 0.2 59.4 ± 9.3 53.9 ± 13.8 28.0 ± 3.0
Semi-dwarf 76 ± 3.2 2.8 ± 0.1 2.3 ± 0.4 58.9 ± 8.9 55.2 ± 14.1 19.0 ± 3.0
Dwarf 59 ± 2.9 1.8 ± 0.4 2.7 ± 0.3 58.2 ± 13.8 60.5 ± 10.2 7.5 ± 0.9
Open Small 230 ± 18 2.7 ± 0.2 3.0 ± 0.25 65.7 ± 12.0 70.0 ± 19.0 23.0 ± 3.0
Weeping Small 205 ± 25 2.5 ± 0.2 4.2 ± 0.3 73.0 ± 12.0 120.0 ± 13.0 26.0 ± 0.3
Compact Small 170 ± 16 2.5 ± 0.3 3.8 ± 0.33 66.7 ± 9.3 71.6 ± 15.5 20.0 ± 3.0
 Botany and Taxonomy 9

Fig. 1.5. Leaf stipules at petiole base.

Fig. 1.6. Narrow leaves (left) compared with normal sized. Scale in centimetres.

Marginal reniform glands are pale green,
while marginal globose glands are yellow.
The eglandular phenotype is associated
with a strong susceptibility to powdery mil-
dew (Podosphaera pannosa (Wallr.:Fr.) Braun &
Takamatsu), and it is systematically elimi-
nated in breeding operations, although this
phenotype could show a good degree of resis-
tance to leaf curl, Taphrina deformans (Berk.)
Tul. (Wickson, 1889; Monet, 1983). Cultivars
10 D. Bassi and R. Monet
Fig. 1.7. Leaf glands: (a) reniform; (b) globose; (c) eglandular (note close-up, below).
with globose glands are generally more sus-
ceptible to powdery mildew than those with
reniform glands (Saunier, 1973).
The leaf blade is darker on the adaxial
side and the colour of the main veins is related
to the flesh colour: yellowish (in yellow-
fleshed fruits) or greenish-white (in white-
fleshed fruits).
REDLEAF. The ‘redleaf’ trait, first described in
a US feral population named ‘Tennessee Nat-
ural’, a seedling rootstock (Hedrick, 1917),
shows purple-red epidermis, particularly on
the young leaves and fruits, while flowers are
dark pink (Fig. 1.8/Plate 7). The lamina tends
to recover the regular green colour late in the
season, but the leaf veins remain purple.
Blake (1937) showed that the character is in-
completely dominant and monogenic: Gr/gr.
At the heterozygous level, the leaves appear
copper-red or copper-red green even at the
young stage, while when homozygous they
are darker red.
From observations on ‘Flordaguard’ (homo-
zygous ‘redleaf’) seedlings, two additional
‘redleaf’ phenotypes have been described,
suggesting the presence of at least one addi-
tional gene (Okie, 1998). In one phenotype the
red fades slowly to a bronze colour. In a sec-
ond phenotype the red fades quickly and
completely (‘quick fade’) to green before
leaves attain full size, except the main vein
and petiole for some time. In a third pheno-
type the leaves fade at an intermediate rate,
with a red-green mosaic on fading full-size
leaves, as in the dominant homozygous. At
the end of the season all types fade to com-
pletely green. The ‘quick fade’ is reputed as
monogenic and recessive.
ANTHOCYANIN DEFICIENCY. The character was
first observed on the old US peach cultivar
‘Summer Heath’. The shoots remain green af-
ter lignification. Flowers have light pink pet-
als, sepals are green and the fruit skin is fee-
bly pink. The overall anthocyanin content is
very low. The character is recessive and mo-
nogenic: An/an (Monet, 1967).
ANTHOCYANINLESS. These trees bear white
flowers; the receptacle and the sepals are
green while the stamens are yellow (instead
of reddish). The young shoots remain green
even after lignification. The fruits are yel-
low at maturity without any trace of red
pigments (Fig. 1.9/Plate 8). The character was
described as recessive and monogenic: W/w
(Lammerts, 1945). ‘Anthocyaninless’ is re-
ported epistatic to ‘redleaf’ (Chaparro et al.,
1995).
 Botany and Taxonomy
Fig. 1.8. Redleaf (left) and greenleaf (right) peach trees.
GREEN APHID RESISTANCE. Resistance to green
peach aphid (Myzus persicae Sulzer) was found
on a weeping tree (‘Weeping Flower Peach’)
and on a seedling rootstock (‘Rubira’) by Mas-
sonié et al. (1982). It is a resistance based on a
hypersensitivity reaction and the aphid makes
only a testing probe on young shoots or leaves.
Around the testing site, within a few days, a
typical necrotic zone develops (Fig. 1.10/
Plate 9). The trait is monogenic and the resis-
tance is dominant over the sensitivity (Rm1/
rm1; Monet and Massonié, 1994).
Flower and fruit development
Peach has hermaphroditic, perigynous flowers.
The reddish-green calyx is gamosepalous and
falls (‘split-jacket’ or ‘shuck-split’ stage) after
the initial swelling of the fruitlet. The inner
surface of the calyx, where the nectaries are
located, is white-greenish in white-fleshed and
yellow to deep orange in yellow-fleshed fruits.
The petals are separated and two shapes
of corolla are known: (i) showy (rose-shaped)
with large petals (Fig. 1.11/Plate 10); and
(ii) non-showy (bell-shaped) with small
petals, where the anthers emerge from the
corolla before full anthesis (Fig. 1.12/Plate
11
11). Connors (1920) and Bailey and French
(1949) studied the inheritance of the flower
shape and the non-showy was found to be
dominant (Sh/sh), the large-sized showy
being dominant over the small-sized showy
(L/l).
Normally there are five petals, from pure
white to dark red, although most cultivars
display a pale to dark pink. In ornamental
forms pure white and red colours are com-
monly found, and chrysanthemum-like pet-
als have also been reported (Yoshida et al.,
2000) (Fig. 1.13/Plate 12).
The number of petals changes according
to the semi-double and the double flower
traits. In the former phenotype the petal num-
ber varies from 12 to 24, the number of stamens
transformed in petals being very low. In the
latter trait the majority of stamens are trans-
formed in petals arranged in two concentric
circles of petals, the inner being darker than
the outer, thus conferring the impression of a
double flower.
The inheritance of double flower is not
known but Lammerts (1945) observed that
three genes are involved in the semi-double
trait: a recessive conditional (d1) and two that
modify the number of petals in excess (dm1
and dm2). Examination of the phenotypic
12 D. Bassi and R. Monet

Fig. 1.9. Fruit skin colour variability. From top centre going clockwise to the right: a yellow nectarine
(anthocyaninless); a white peach (anthocyaninless); a yellow peach (100% blush); a white peach;
a non-melting peach; a yellow flat nectarine; a white flat peach; a white nectarine; a yellow nectarine.

Fig. 1.10. Hypersensitivity reaction on a resistant peach after a proof bite by a green aphid.
 Botany and Taxonomy 13

Fig. 1.11. Flower type: showy.

Fig. 1.12. Flower type:

non-showy.

frequencies observed by Lammerts (1945) homeotic: their mutation transforms one

suggests a simpler explanation with one organ to another.
recessive gene responsible for the presence of As many as 20 to 30 stamens are attached
supernumerary petals (D1/d1). The genes that to the calyx. The anthers are reddish unless
regulate the organs’ morphogenesis are male sterility (anthers without pollen) is
14 D. Bassi and R. Monet

White, simple White, semi-double Variegated, white and red

Chrysanthemum-like, red, semi-

Red, simple Red, semi-double double

Pink, semi-double

Fig. 1.13. Flower diversity in ornamental peach cultivars. (Courtesy of M. Yoshida, Japan.)

present: sterile anthers look pale yellow
instead of reddish before dehiscence. Follow-
ing self-pollination, some progenies may be
male sterile but individuals bearing this trait
are typically eliminated from selection. Nev-
ertheless, some male sterile cultivars were
largely cultivated, the most famous being
‘J.H. Hale’ because of the outstanding quality
of its fruits. Scott and Weinberger (1944) listed
a series of cultivars heterozygous for the trait.
Male sterility was finally described as mono-
genic and recessive (Ps/ps; Bailey and French,
1949). A new nuclear male sterility trait was
found in ‘White Glory’ (a weeping, ornamen-
tal cultivar) which is different from the gene
of ‘J.H. Hale’: Ps2/ps2 (Chaparro et al., 1994;
Werner and Creller, 1997).
Microsporogenesis begins in winter
(Knowlton, 1924; Draczynski, 1958) and is
followed by meiosis close to bud swell. The
gynoecium is superior and it is obviously
glabrous in nectarines (see section on ‘Skin
adherence and pubescence’ below). Abnor-
mally high temperatures during flower bud
 Botany and Taxonomy
initiation may lead to double or triple gynoe-
cium (and pistils), which in turn may eventu-
ally give rise to double or triple fruits
(discarded during thinning because they are
of no commercial value; Fig. 1.14/Plate 13).
The eight-nucleate stage of the megagameto-
phyte happens a few days before full anthe-
sis, when the two polar nuclei migrate into
the middle of the embryo sac, which elon-
gates after union of the polar nuclei and dou-
bles its length at the time of fertilization. The
ovary contains two ovules, but normally only
one is fertilized. Time from pollination to fer-
tilization depends on temperature, varying
from 24–48 h (Toyama, 1980; Baipo et al., 1989)
to 12 days (Herrero and Arbeloa, 1989). The
endosperm becomes multinucleate about 10
days after fertilization and cellular after 5–6
weeks (Harrold, 1935; Lilien-Kipnis and Lavee,
1971). Around 8 weeks after full bloom the
integuments reach the maximum size (about
20 mm). The first division of the zygote occurs
about 2 weeks after ovule fertilization (Harr-
old, 1935). The embryo fills the testa, absorb-
ing the nucellus and the endosperm in about
100–110 days from full bloom. Thereafter it
completes its growth by dry matter accumu-
lation: starch, protein and lipids (about 50%).
The ovary (fruit) undergoes four main stages
Fig. 1.14. Flat fruit, from a double ovary.
15
of growth. The first, rather rapid stage (stage
I) is marked by cell division (the length of this
phase is nearly the same no matter the fruit
development period, FDP). This is followed
by a slower stage (stage II) where most of the
dry matter is employed in pit hardening and
seed and embryo growth (this period is very
short in very early-ripening genotypes, where
at fruit maturity the stone is incompletely lig-
nified and the embryo does not reach full
maturity, in vitro rescue being necessary to
recover germination and plant development).
The third stage (stage III) is more rapid
because cell enlargement and elongation is
according to FDP. The last stage (stage IV) is
the ripening phase (Lilleland, 1933; Tukey,
1933; Harrold, 1935; Gage and Stutte, 1991).
The fruit peduncle remains attached to
the shoot after fruit senescence and natural
abscission.
Fruit appearance and composition
Shape and size (weight)
The peach fruit is a drupe. Almost all com-
mercial cultivars share round (globose) or
elongated (either oval or more or less oblong)
16 D. Bassi and R. Monet
fruits (Fig. 1.15, left and right, respectively/
Plate 14), elongated shapes being dominant
over round (Blake, 1932).
The flat (or ‘saucer’ or ‘pan-tao’, from the
Chinese words ‘flat peach’) fruit, introduced
from China, has long been a botanic curiosity
in Western countries (Fig. 1.15, centre/Plate
14). The fruit is flattened at opposite poles
and the shape affects not only the fruit but
also the pit, which is flattened too and rather
small. The seed is spherical and has poor ger-
mination. The trait is monogenic and domi-
nant over round/elongated: S/s (Lesley,
1940). The homozygous genotype is lethal.
Fruit weight varies from less than 50 g
in wild forms to 80–110 g for the very, very
early-ripening genotypes to over 680 g (Li,
1984), although commercial standards require
from 180 to 230 g, depending on FDP and final
use.
Fruit weight follows a quantitative inheri-
tance. Connors (1922) reported that prepotency
in fruit size inheritance depended on parent
(Fig. 1.16/Plate 15). Lesley (1957), working on
self-pollination up to the seventh generation
on several lines, observed an inbreeding effect
on fruit size for only one line. Similar findings
are reported by Monet et al. (1996), who had no
inbreeding effect after three selfing cycles and
Fig. 1.15. Main fruit shapes in commercial cultivars: globose (left), flat (centre), oblong (right).
reported a bias towards large size in assessing
progenies from parents differing in fruit size.
Skin adherence and pubescence
Skin adherence to the mesocarp was reported
as recessive to non-adherence, even if influ-
ence from flesh texture may occur (Lesley,
1957).
Two major phenotypes are known about
epidermis surface: (i) the standard, fuzzy peach;
and (ii) the nectarine, with a glabrous skin.
The nature of the nectarine skin character was
described as recessive (Rivers, 1906; Blake,
1932; Blake and Connors, 1936) and mono-
genic: G/g (Bailey and French, 1949). Faust
and Timon (1995) indicated that the nectarine
mutation probably first appeared in the most
north-western part of China, in the Tarin
basin (part of the Turkistan region). As early
as the 14th century, nectarines were noted in
Europe (Gallesio, 2003).
From studies of the fuzzless skin trait, three
different types of heritable skin surface (at a
microscopic level) were described (Fogle and
Faust, 1975). One more phenotype, intermedi-
ate between nectarine and peach, with a rough
surface, was also described. It has a pleiotrophic
effect on lack of pubescence (or trichomes) on
 Botany and Taxonomy
Fig. 1.16. Fruit size gain in F1 progeny from distant-size parents.
dormant buds, which look shiny and smooth
(Okie and Prince, 1982). The trait was described
as monogenic and recessive: Rs/rs (Okie, 1998).
The locus controlling the nectarine trait
is probably closely linked to or has a pleiotrophic
effect on several other traits. Several nectarine
sports, compared with their peach progeni-
tors, often show smaller and rounder fruits
(reduced cell number), greater specific gravity,
higher soluble solids and organic acids. Other
traits may also be involved in the mutation,
e.g. the FDP and the chill unit requirement
(Wen et al., 1995).
The smooth skin of nectarines makes
them more susceptible to russeting, mechani-
cal bruising and pest damage (e.g. thrips)
than peaches but exploitation of the nectarine
trait has led to tremendous advances in peach
breeding. During the period from the 1970s to
the 1990s, nectarine cultivars became so pop-
ular that their profits overtook that of standard
peaches. Presently, in some countries such as
Italy, their prices have reached a steady level,
comparable to the standard peaches.
Nectarine features are so distinct that
sometimes they are regarded by marketers as
an independent species. Indeed, the bright
17
and glossy over-colour and sometimes dis-
tinct flavour play significant roles in driving
consumer demand in most markets.
Skin and flesh colour
Skin (ground) and flesh colour, white or yellow,
are among the most popular and commercial
criteria for peach cultivar classification, due
to the peculiar features of these phenotypes
(Figs 1.9 and 1.17/Plates 8 and 16). ‘White’
peaches are revered for their distinct flavour
and/or aroma (Robertson et al., 1990), although
most of them are either too soft or too suscep-
tible to skin bruising and flesh browning and
thus not competitive with the ‘yellow’ types,
where carotenoids (the orange pigments) could
mask oxidation from bruising or other blem-
ishes. Xanthophylls, the yellow pigments, are
synthesized via hydroxylation from carote-
noids: lutein from b-carotene, zeaxanthin,
antheraxanthin and violaxanthin from b-
carotene (Demmig-Adams and Adams, 2002).
Carotenoids are photosynthetically active
pigments, while xanthophylls dissipate the
light excess that can disrupt photosynthesis.
Carotenoids and xanthophylls are localized
18 D. Bassi and R. Monet

in chloroplasts (chromoplasts) and are found
in very small amounts in white peaches. Among
carotenoids, b-carotene and b-cryptoxanthin
are the primary provitamin A factors, their
concentrations reaching around 2000 mg/kg
of fresh weight (FW) for the former and up to
3400 mg/kg FW for the latter (Tourjie et al.,
1998). Connors (1920) showed that white flesh
is dominant to yellow flesh. Bailey and French
(1949) suggested the symbols Y/y for the two
alleles.
Anthocyanins, the glycoside derivatives
of the anthocyanidins, are responsible for all
colours from blue to red and are localized
within the cell vacuoles, in either the epider-
mis or the flesh. They are synthesized from
flavonoids via phenylalanine.
The presence of anthocyanins is indepen-
dent of the skin ground colour, either yellow
or white, and can be of quantitative or quali-
tative origin. The quantitative trait is posi-
tively influenced by light exposure and the
pigments reach the maximum concentration
at full ripening, while the qualitative trait,
expressed only in the epidermis, is not related
to light nor to ripening and the localization is
limited to the skin. There are two indepen-
dent genes regulating the phenotype of quali-
tative origin. One is the dominant ‘redleaf’
trait (see section on ‘Redleaf’ above); the sec-
ond is recessive and is expressed in the fruits’
epidermis only, which looks much brighter
than in ‘redleaf’, and was designated as Fr/fr
(Beckman and Sherman, 2003).
A qualitative suppressive trait for fruit
red skin, dubbed ‘highlighter’ (H/h; Beckman
et al., 2005), was recently reported. When
homozygous recessive it completely sup-
presses the red pigments, but only in the fruit,
whereas the w/w trait is characterized by the
absence of anthocyanins in any plant tissue.
While carotenoids are rather heat-stable,
anthocyanins are very labile and subject to
browning in canning operations; this has led
to the selection for flesh of canning peaches
that is anthocyanin-free. Since localization of
anthocyanins in skin is independent from
that in flesh, commercial canning peaches
may or may not develop a red over-colour,
given the fruit is peeled before canning.
When anthocyanins are present in the
flesh, their localization is mainly under the
skin and/or close to the pit, the latter reported
as dominant over no red (Lesley, 1957). The

Fig. 1.17. Flesh colour variability. From left, top row: non-melting flesh, two yellows (greenish and
bright yellow), white; bottom row: red flesh (‘blood’), a yellow melting (anthocyaninless), a yellow and
a white melting (flat shape), a white stony-hard (anthocyaninless).
 Botany and Taxonomy
amount and distribution of anthocyanins
throughout the mesocarp depends on the cul-
tivar (quantitative trait) and may be variably
expressed among the fruits on a given tree.
Red (‘blood’) flesh peaches
One exception to the erratic distribution of
the red pigments within the flesh are the ‘red
flesh’ (or ‘blood’ flesh) peaches (Fig. 1.18/
Plate 17) and nectarines (Fig. 1.19/Plate 18),
where almost all of the flesh is heavily stained
by anthocyanins (Gallesio, 1817–1839) inde-
pendently of the basic flesh colour, either
white or yellow (Fig. 1.20/Plate 19). In both
types, when the ‘red flesh’ trait is present, the
skin has a distinct purple, dull finish. The trait
was first described as dominant by Blake
(1932), and confirmed by Werner et al. (1998).
Flesh compounds influencing flavour
and aroma
Several compounds contribute to the overall
flavour of the flesh: aromatics (volatiles), organic
acids, phenolics and sugars are among the
most known.
Fig. 1.18. Red ‘blood’ flesh in peach. (Courtesy of A. Liverani, Forli, Italy.)
19
Among the aromatics or volatiles (alco-
hols, aldehydes, esters, etc.) that contribute
significantly to the typical peach aroma, pecu-
liar distribution patterns of hexanal (up to
about 740 ppm), trans-2-hexanal (up to about
120 ppm), linalool (up to about 250 ppm),
g- and d-decalactone (up to about 130 and
25 ppm, respectively) between white- and
yellow-fleshed cultivars were found, with the
former showing higher total amounts (Rob-
ertson et al., 1990).
The main organic acid is malic (over 50%
of the total), followed by citric, quinic and
succinic acids. When individually analysed,
total acids, expressed as malic acid, may
range from 0.9 up to about 1.6% FW.
Ascorbic acid (vitamin C) content in peach
fruits is generally low (below 10 mg/100 g FW),
but in some cases it may be threefold higher
(Liverani and D’Alessandro, 1999).
Sugar content is generally based on
assessing the soluble solids content (SSC) by
refractometer. This value may reach up to
20% or more, although the average values
found in commercial cultivars range from 9 to
15% (Byrne et al., 1991; Crisosto et al., 1998).
When individually analysed, total sugars
20 D. Bassi and R. Monet

Fig. 1.19. Red ‘blood’ flesh in nectarine. (Courtesy of A. Liverani, Forli, Italy.)

Fig. 1.20. Genetic variation for red ‘blood’ flesh trait in peach. (Courtesy of W.R. Okie, Byron,
Georgia, USA.)

may reach up to 16% FW; the main sugar
being sucrose, ranging from 45 to above 80%
of total sugars, followed by either glucose or
fructose (Bassi and Selli, 1990; Byrne et al.,
1991) and sorbitol; other alcohol-soluble
sugars are in low or barely detectable
amounts: inositol, mannose, xylitol and
xylose. Low-quality peaches may exhibit a
fourfold lower amount of fructose (the sweet-
est sugar in peaches) and a threefold higher
 Botany and Taxonomy
amount of sorbitol but similar SSC compared
with high-quality peaches (Robertson et al.,
1988).
Some genotypes show a distinct low
acidity level, resulting in the so-called ‘low-
acid’ (LA) trait. In early times, this character
was described as an attribute of the ‘honey
peaches’ group (Reimer, 1906), easily distin-
guishable from the ‘acidic’ peaches by a
remarkably sweet taste. The trait is well
known and regarded in the Far East countries
(e.g. China, Japan, Korea), where very sweet
fruits are particularly appreciated. From prog-
enies obtained by self-pollination of ‘Red-
wing’ and ‘Robin’ white peaches heterozygous
for the trait, Monet (1979) described the low
or sub-acid as a dominant and monogenic
character (D/d). The LA phenotype shows
mainly citric and malic (about 50%), but also
quinic (about 20%) acids in lower concentra-
tions than standard phenotypes, while total
acidity is from two- to fourfold lower (0.4 ver-
sus 1.4: average values from pooled LA and
standard cultivars, respectively). The pH of
the LA ranges above 4.0, while in standard
phenotypes the pH is below 3.9. The ratio
between SSC and titratable acidity (TA) is
almost four times higher in the LA pheno-
types (Yoshida, 1970; Monet, 1979; Ventura
et al., 1995; Moing et al., 1998; Liverani et al.,
2003). SSC is comparable to that in the stan-
dard phenotype, although LA types show
more sucrose and less glucose (Liverani et al.,
2003). For a better taste of the LA peaches,
SSC above 12% is suggested, in order to over-
come the bland feeling of a low acidity cou-
pled with too low sugar content (Delgado,
1998). Nevertheless, the TA of the best-tasting
LA cultivars is at least twice that in most of
the other LA cultivars (Liverani et al., 2003).
Phenolic compounds may play a signifi-
cant role in flavour because they are responsi-
ble for the ‘astringent’ taste. When comparing
low- and high-quality white-fleshed cultivars,
Robertson et al. (1988) noted that for those
rated as unacceptable, there were comparable
amounts of sugars and acids but seven times
more phenolics than in the high-quality culti-
vars (120–140 mg/100 g FW). Oddly, even
higher phenolic contents (up to 150 mg) found
in commercial white peaches of local origin in
Italy were rated as acceptable (Bassi and Selli,
21
1990). The discrepancy between these reports
may be due to the differences in the aromatic
profiles of the cultivars evaluated. It is possi-
ble that the very strong aroma of the Italian
white peaches partially masked the taste of
the phenolics present.
Polyphenolic compounds are also
responsible for the browning resulting from
their oxidation by the enzyme polyphenylox-
idase (PPO). Mechanical damage of the cells
(skin, flesh) results in the rupture of the vacu-
oles where the polyphenolic compounds are
stored, thus exposing them to enzymatic oxi-
dation by PPO. As a result, quinones are
formed that polymerize to brown-coloured
pigments. Since these traits are of a quantita-
tive nature, their heritability can be estimated
(Hansche and Boynton, 1986) so that cultivars
with low susceptibility can be selected.
Flesh becomes more astringent in many
cultivars under cool summer temperatures
(D. Bassi, personal observation).
Flesh texture
The composition of the cell wall strongly
affects flesh texture. So far at least three main
distinct phenotypes are known, even if not all
is understood in terms of genetic determina-
tion and biochemical pathways during the
final ripening stages. The first two pheno-
types, described by Bailey and French (1932),
are the melting (M) and the non-melting (NM).
The M texture shows a prominent softening
in the last stage (stage IV) of ripening, until a
complete melting. Variability in firmness (or
rate of softening) is found within this pheno-
type and Yoshida (1976) distinguished between
soft, medium and firm. The ‘firm’ type (FM)
softens rather slowly and is less susceptible to
bruising during handling, thus allowing an
easier management of harvest timing and
other grading and shipping operations, and
displays a longer shelf life. In addition, it
shows a rather high amount of water-insolu-
ble pectins and of Ca bound to the cell wall
(Mignani et al., 2006).
The NM phenotype (the so-called ‘can-
ning peach’) shows a firm texture when
fully mature, softens slowly when overripe
but never melts. Rather, it becomes rubbery
(because the loss of water) during the
22 D. Bassi and R. Monet
senescence stage, when most cultivars could
display a peculiar off-flavour (Sherman et al.,
1990). However, Beckman and Sherman (1996)
noted that it is possible to select for the absence
of off-flavours within breeding progenies.
The lack of softening in the NM pheno-
type is related to the loss of endopolygalactu-
ronase (endoPGase) activity, the enzyme
responsible for cleaving pectins (polygalac-
turonic acid chains) from the cell wall in the
M fruits (Lester et al., 1996). Morgutti et al.
(2006) detected a mutation in the endoPG
gene, exploitable for early marker-assisted
selection of the trait, even when heterozy-
gous. The melting trait was described to be
dominant over the non-melting (M/m; Bailey
and French, 1932). See the section on ‘Flesh
adherence to endocarp (stone, pit)’ for more
details about the inheritance of this trait.
M and NM phenotypes develop a rather
high amount of ethylene between stage III
and stage IV, often more abundantly in NM
types (Mignani et al., 2006). The NM flesh is
also much less susceptible to mealiness, a
rather common storage chilling injury disor-
der (Brovelli et al., 1998). Recently, a large QTL
(quantitative trait locus: a DNA zone contain-
ing several genes responsible for a given quan-
titative trait) for mealiness was detected for the
Fig. 1.21. Stony-hard fruit: white (anthocyaninless) flesh.
endoPG locus, confirming the previous obser-
vation that this disorder occurs particularly in
M freestone phenotypes (Peace et al., 2005b).
The third flesh texture phenotype was
first described by Yoshida (1976). He classified
a very firm and crispy, ‘stony-hard’ (SH) flesh
type as belonging to the M family. However,
this type never melts, as in ‘Yumyeong’, a
white-fleshed peach from Korea. Its fruit
resembles an NM phenotype, becoming rub-
bery when senescent, and can be either white-
or yellow-fleshed (Fig. 1.21/Plate 20, Fig.
1.22/Plate 21), the only remarkable difference
from NM being the almost complete lack of
ethylene production (Goffreda, 1992; Haji
et al., 2001, 2003; Tatsuki et al., 2006), although
it can be stress-induced, as from storage
below 10°C (Tatsuki et al., 2006; Gamberini,
2007; Begheldo et al., 2008). The recessive gene
was named Hd (Scorza and Sherman, 1996).
Water-insoluble pectins in cell walls are rather
high, as Ca is bound to them, although its
content is variable (Bassi et al., 1998). Stony-
hard flesh is often LA, but when progenies
are obtained for breeding purposes, and this
firm texture segregates with the acidic (non-
low acid) character, a prominent sour taste
prevails (J.C. Goffreda, New Jersey, 1997, per-
sonal communication).
 Botany and Taxonomy
From the biochemical point of view, the
lack of ethylene evolution is due to the tran-
scription suppression (and not to its muta-
tion) of the 1-aminocyclopropane-1-carboxylic
acid synthase isogene (Pp-ACS1), a key gene
of the ethylene enzymatic pathway (Tatsuki
et al., 2006, 2007).
From the genetic point of view, the inde-
pendent inheritance of the SH flesh from M
and NM has been demonstrated, also sug-
gesting an epistatic effect of SH, since when
exogenous ethylene is applied the SH/M
(hdhd/f–) phenotype is induced to melt, while
the SH/NM (hdhd/f1f1) keeps firmer, as a
standard NM flesh (Haji et al., 2005; see also
Table 3.1).
From the practical point of view, SH
fruits are often very difficult to distinguish
from NM or very firm, unripe, M phenotypes.
Therefore, evaluation of progenies from con-
trolled crosses segregating for SH is puzzling.
SH flesh identification on the tree is very time-
consuming to determine (several passes are
required in order to check firmness evolution)
and sensory evaluation (by tasting) is not
always reliable. So far, the only sound method
for SH texture phenotyping is to measure eth-
ylene production (Goffreda, 1992).
Fig. 1.22. Stony-hard fruit: yellow flesh.
23
Furthermore, there is a possible fourth
flesh texture trait (‘very, very firm’), whose
phenotype resembles very much the SH
flesh in firmness and crispiness, but when
fully ripe becomes melting and develops
ethylene, although in an unpredictable
fashion from year to year (I. Mignani, Italy,
2007, personal communication). This flesh
texture, firmer than the ‘firm’ M types, is
found in many recently developed new cul-
tivars (both nectarines, e.g. ‘Big Top’, and
standard peaches, e.g. ‘Rich Lady’ and
‘Diamond Princess’). It was first commer-
cially introduced from private breeders
from California, and is sometimes (e.g. ‘Big
Top’ and others) associated with the low-
acid trait.
Its remarkable keeping quality, particu-
larly on the tree, is a rewarding character
for both growers and consumers. However,
this flesh type is very difficult to assess, and
the same problems as described above for SH
flesh phenotyping are experienced. Work is in
progress to fully understand this trait, from
both the biochemical (physiological) and
genetic points of view, since it seems to be a
dominant Mendelian trait over the standard
M type.
24 D. Bassi and R. Monet

A tentative classification of peach fruit However, since in the studied families

flesh phenotypes is presented in Table 1.2.
Flesh adherence to endocarp (stone, pit)
The flesh may or may not adhere to the
endocarp. According to Bailey and French
(1932) this trait is controlled by the ‘freestone’
locus, where the freestone (F_) allele is domi-
nant over the clingstone (ff). Intermediate
behaviour (semi-freestone or semi-clingstone)
with varying degrees of adhesion is observed,
particularly in early-ripening genotypes
(FDP less than 100 days) (Weinberger, 1950).
Rapid flesh maturation, due to early ripening,
delays the appearance of the freestone
phenotype, thus the semi-freestone pheno-
types should be regarded as ‘physiologically
clingstone’ but ‘genetically freestone’ (Beck-
man and Sherman, 1996). Since the intensity
of these characters changes during fruit
ripening, flesh adherence to the endocarp in
early genotypes should be assessed at full
ripening or even at the early stage of
senescence.
Bailey and French (1932) suggested that
the flesh adherence to the pit gene and the
flesh texture gene were linked on the same
chromosome. This was the first published
linkage in peach genetics.
one phenotype was missing (freestone-NM,
as expected by recombination), the Bailey and
French interpretation was reconsidered,
although semi-freestone peaches with NM
flesh have been observed where a thin layer
of flesh adheres to the stone (Beckman and
Sherman, 1996). The genetic nature of this lat-
ter phenotype is not clarified yet, and the
semi-freestone trait in NM flesh genotypes
should be regarded as of different nature than
in M peaches (T.G. Beckman, Georgia, 2005,
personal communication). Using the data of
Bailey and French (1932), Monet (1989) sug-
gested the three-alleles-one-locus theory. The
three following phenotypes resulted: free-
stone-M (FF, Ff or Ff1), clingstone-M (ff or ff1)
and clingstone-NM (f1f1), with dominance
from left to right. From recent studies on
progenies segregating for endocarp adher-
ence and flesh texture (M and NM), four
alleles for the endoPGase enzyme were found
responsible for the same three flesh pheno-
types as above at a single locus (F): freestone-
and clingstone-M and clingstone-NM; the
fourth allele is a null-allele (absence) that has
the same phenotypic effect as the f1 allele, a
mutation that nullifies the enzyme function
(Peace et al., 2005a). A diagnostic PCR test
was made available to detect all four alleles.

Table 1.2. Tentative classification of peach fruit flesh phenotypes from chemical analysis and sensory
evaluation at physiological maturity. (From Yoshida, 1976; Bassi et al., 1998; Haji et al., 2005; Mignani
et al., 2006.)

Pectinsa

Flesh texture Firmness Soluble Insoluble Calciuma Ethyleneb

Melting
Soft Low +++ + ++ ++
Firm High +++ ++ +++ ++
Very, very firmc Very high ++ ++ ++ +
Stony-hard Very high + +++ ++ –/(+)
Non-melting Very high +++ +++ +++ +++
aFlesh content; no clear-cut threshold between different phenotypes.
bAmount produced by whole fruits.
cSimilar to the ‘stony-hard’, but produces ethylene and becomes melting in the very last stages of ripening

(e.g. ‘Big Top’ nectarine).

Number of + represents relative content within columns, respectively; – means absence; (+) means
traces.
 Botany and Taxonomy
Thus, the Monet (1989) hypothesis of three
alleles at the same locus controlling both
flesh texture and endocarp adherence was
confirmed and the previous theory of Bailey
and French (1932), suggesting a linkage
between flesh texture and its adherence to the
pit, should be rejected.
SH cultivars known so far are all cling-
stone (A. Liverani, Italy, 2005, personal com-
munication).
Fruit ‘keeping’ quality and resistance
to manipulation (bruising)
‘Keeping’ quality is related to resistance of the
fruit to blemishes caused by excessive soften-
ing and refers mainly to firmness of the flesh.
Bruising depends mainly on skin resis-
tance to browning; thus a rather firm fruit
(flesh) could also be prone to bruises if the
skin is too delicate.
Endocarp (stone, pit) and seed
The endocarp is lignified, the outer surface
being deeply furrowed and pitted. In very,
very early-ripening genotypes (FDP less than
55–60 days) lignification is limited and the
endocarp may be rather soft, thus allowing
consumption of the entire fruit, when the
seed is not bitter. Two examples in Italy are
‘Borgia’ and ‘Lucrezia’ (Bassi and Rizzo,
1995). A more or less pronounced ridge is
present in the ventral suture, and a needle
(very acute tip) could be present at the apex.
The latter trait is undesirable in canned peaches
because its fragments are difficult to eliminate
from processed fruits. Endocarp splitting (at
the carpel suture) or shattering (radial frac-
tures) may affect either early- or late-ripening
cultivars. Both are commercially undesirable
and the former a consumer hazard in eating
the stone fragments. While it is possible to
select against these undesirable traits, it is
reported that cultural practices to improve
fruit size (supplemental irrigation, girdling,
etc.) may also increase the incidence of endo-
carp splitting or shattering.
The stone shape changes according to the
fruit shape, from globose (in round fruit),
25
elliptic (in ovate or elliptic fruit) to round-
oblate (in flat fruits). It contains one
(exceptionally two) seed(s), whose cyanidic
glucoside content makes them taste very bit-
ter. This latter trait is monogenic and domi-
nant over the non-bitter phenotype (Sk/sk;
Werner and Creller, 1997) that can be found in
some nectarines, e.g. ‘Fantasia’, Early Sun-
grand’, etc. Although the locus for the non-
bitter seed is very close to the nectarine trait
(12 cM), it could be recombined with pubes-
cent skin via cross-breeding.
Seeds are highly germinable after
stratification, their chilling requirement being
related to the chilling requirement of the
mother tree (Pérez, 1990; Pérez et al., 1993).
Viability is hampered in early-ripening geno-
types (FDP less than 100–120 days). For these,
aseptic embryo culture is needed to ensure
germination, allowing rescue of immature
embryos as early as 50 days after full bloom
(Fig. 1.23/Plate 22) (Tukey, 1935; Ramming,
1985).
1.4 Peach Biology and Phenology
Floral biology and fruit set
Peach is an insect-pollinated species and it is
self-fertile. Even if some genotypes show a
low fruit set, no evidence has ever been reported
of self-incompatibility as happens in most
other Prunus species. Flower fertilization
from self-pollination is generally high (rang-
ing from 10 to 90% of fruit set), resulting in a
high number of fruitlets (Szabò and Nyéki,
1999); thus crop reduction by fruit removal,
i.e. thinning, is required in order to gain com-
mercial fruit size. Even if cross-contamination
in closely planted trees may reach around
14–25% (Fogle and Dermen, 1969; Fogle, 1977),
cross-pollination under normal conditions is
lower than 5% (Hesse, 1975; Fogle, 1977). The
only character affecting yield is male sterility,
but this trait has been eliminated in present-
day commercial cultivars.
Some genotypes, mainly nectarines,
could be affected by a continuous fruit drop,
well after the ‘June drop’, even leading to
substantial crop losses.
26 D. Bassi and R. Monet
Chilling and heat requirements
Chilling requirement is the amount of cold
(temperature below a given threshold)
required by flower and leaf buds in order
to complete morphological development
(particularly for reproductive organs) and
rest. Several methods have been proposed so
far to measure this physiological requirement.
The simplest method is that of Weinberger
(1950), where the number of hours below 7°C
is taken in account. This method is popular
worldwide, but it does have some limitations.
Richardson et al. (1974), with their ‘Utah
model’, defined chilling units (CU), giving
specific weight to different temperatures,
understanding the role of negation of rest
above a given threshold (16°C) before rest
completion. A better evaluation of the effect
Fig. 1.23. Comparison between
mature (left) and immature (right)
embryos taken from ‘Spring Crest’
peach at two different stages: the
immature embryos fail to germinate
under standard stratification procedures
and need to be rescued in vitro as the
very early-ripening genotypes.
of fluctuating temperatures and their role in
negation of rest in the low-chill areas was
given by the ‘dynamic model’ of Erez et al.
(1990). While CU can be measured by artifi-
cial methods (Dennis, 2003), a simple and
cheaper method is to utilize standard cultivar
bloom times as an indication of the chilling
requirement of unknown genotypes (Scorza
and Sherman, 1996). See Chapter 5 for further
details.
Ranking known genotypes for CU, the
lowest level may be around 50 CU (‘Florda-
Grande’), up to more than 1500 CU.
Analogous to chilling, heat requirements
refer to the amount of heat (temperature above
a given threshold) after endo-dormancy is ful-
filled to achieve organ development, from
bloom (Richardson et al., 1974; Citadin et al.,
2001) and foliation to fruit maturation.
 Botany and Taxonomy
The ‘evergreen’ trait has been described,
where the terminal growth never stops (no
terminal bud formation) unless killed by frost
(Lammerts, 1945; Diaz, 1974), while lateral
buds show about 450 CU. The trait, probably
due to lack of phytochrome response and
resulting from a deletion of the wild type
(Bielenberg et al., 2004), was found to be
monogenic and recessive: Evg/evg (Rodri-
guez et al., 1994). In subtropical regions, it
allows the yielding of two crops per year.
The trait is a candidate model system to
study winter dormancy in woody plants
(Wang et al., 2003).
Phenological phases
The peculiar stages of morphological develop-
ment of the main organs (bud, flower, leaf and
fruit) from bud break to leaf fall are termed
phenological phases (Fig. 1.24/Plate 23). The
occurrence of the single stages (e.g. bud break,
full bloom, split-jacket, etc.) plays a key a role
in determining specific orchard operations
(e.g. the spray schedule against pests and dis-
eases) or in cultivar assessment for the evalua-
tion of their environment adaptability.
Time of flowering
Time of flowering depends on the CUs neces-
sary to fulfil rest and on the growing degree
hour (GDH) accumulation in order to reach
full bloom. Even if the two traits are genetically
distinct, it is not simple to separate the two
components in selection breeding, particularly
because the threshold temperatures needed
for their fulfilment have not been determined
(Scorza and Okie, 1991). Since bloom date is
quantitatively inherited and heritability is
rather high, breeding could be addressed based
on the parents’ behaviour in a given environ-
ment, where up to 40 days from the earliest- to
the latest-blooming cultivar has been recorded.
For further details see Chapter 5.
Time of ripening (fruit development period)
There is no relationship between flowering
and ripening time.
27
While wild peaches exhibit from medium
to late ripening, i.e. an FDP from 120 to 210
days from full bloom to the onset of ripening,
the FDP of commercial cultivars may range
from as early as 55 (Ramming and Tanner,
1987) and 60 days (Bassi and Rizzo, 1995) to
as late as 270 days, e.g. some local cultivars
from Sicily, Italy (Caruso et al., 1992; Caruso
and Sottile, 1999).
An interesting ripening mutation (‘slow
ripening’) described in progenies of ‘Fantasia’
nectarine hinders completion of ripening (Fig.
1.25/Plate 24). Fruit development is appar-
ently halted before the end of the cell expan-
sion phase (stage III) and the flesh either
never softens or softens very slowly, while it
keeps a crispy texture (but not of the non-
melting type). The skin ground colour and
flesh are greenish and the flavour is very poor,
despite lower acidity and higher pH and sol-
uble solids (but similar total sugars) than
‘Fantasia’ (Brecht and Kader, 1984; Brecht
et al., 1984). Ethylene and carbon dioxide pro-
duction are very low, no aroma is developed
and the fruits remain firm on the tree even
after leaves abscise in autumn. The fruit of
this mutant is susceptible to internal break-
down. After ripening-inducing treatments
using propylene gas, the fruit eventually
becomes soft and advances the onset but not
the level of ethylene, without improving the
poor texture and flavour. The ripening behav-
iour seems intermediate between climateric
and non-climateric classes of fruits, suggesting
a basic similarity between those two categories.
The trait was classified as monogenic and
recessive (Sr/sr), ‘Fantasia’, ‘Flamekist’ and
‘Fairlane’ nectarines being heterozygous for
this trait (Ramming, 1991).
Although the trait regulating FDP is
clearly quantitative, the presence of major
genes can be clearly presumed by two pieces
of evidence. First, when measuring the time
of ripening in large progenies from distant (in
terms of FDP) parents, grouping of offspring
in bimodal or trimodal distribution is often
observed. Almost all of the offspring ripen
within the parental dates, with some seed-
lings ripening earlier or later (Yamaguchi
et al., 1984; Bassi et al., 1988). Second, bud
sports, known mutants from commercial cul-
tivars, show FDP that are roughly separated
28 D. Bassi and R. Monet

(a) (b) (c)

-

Dormant bud Bud swell Pink stage

(d) (e) (f)

Early bloom Full bloom Late bloom

(g) (h) (i)

Petal fall Split-jacket Fruit set

(j) (k) (l)

Small fruitlets (cell division) Pit hardening Final swell

(m) (n) (o)

Fruit veraison Commercial ripening Physiological ripening

Fig. 1.24. Main phenological stages in peach. (Courtesy of E. Bellini, Florence University, Italy.)
 Botany and Taxonomy
Fig. 1.25. Slow-ripening nectarine trees after leaf fall (dormant season).
by weekly intervals, as in ‘Red Gold’ nectar-
ine (Bassi et al., 2004): this opens an interest-
ing insight for genomics in search of QTLs.
1.5 Cultivar Classification
Several cultivars, local types and landraces
have been described in peach. Due to the
rather high number of morphological Mende-
lian traits, the cultivar classification could be
addressed under several keys. For the com-
prehensive list of these traits please refer to
Chapter 3.
In addition, physiological and quantita-
tive traits of economic importance also play a
significant role in horticultural peach taxon-
omy (phenology).
Peach description sheet
After Zielinski (1955), Bellini and Scaramuzzi
(1976) and Bellini et al. (2007), for a compre-
hensive characterization of a given cultivar
the following organs should be described.
● Tree: see section on ‘Tree growth habit’
above.
29
● Shoot (on at least 30 one-year-old fruit-
ing shoots, after leaf abscission): length,
internode length, number of flower buds/
nodes, flower bud distribution along the
stem, bark colour.
● Flowers (on at least 30 items, at full bloom):
type, colour, petal size (length and width),
length of the pistil towards the stamens;
colour of calyx (inner and outer).
● Leaf (on at least 30 items collected from
the middle section of fruiting shoots, ex-
cluding the petiole): colour of the blade
(green, purple) and main veins (greenish
or yellowish); size (blade length and width),
shape (length:width ratio and position of
the broadest width referred to the mid-
dle); surface (flat, wavy); apical and basal
angle of the blade; margin shape (crenate
or serrated); glands (eglandular, globose,
reniform).
● Fruit (on at least 30 items): weight; size
(length, diameters: suture and cheeks);
shape (two sections: along the suture and
equatorial); base cavity depth and width;
apex shape and tip (or beak, if present);
suture (line (no cavity), deep, medium
or shallow). Skin: pubescense (absent,
light, coarse); red blush (per cent coverage
and pattern at physiological ripe stage:
30 D. Bassi and R. Monet

uniform, dotted, striped, etc.). Flesh: firm- FLESH COLOUR

ness (by penetrometer); colour (yellow, 1. White
white); red flesh (‘blood’ trait: present, ab- 2. Yellow
sent); anthocyanins distribution (under
skin, close to the pit, in the middle); fla- FLESH TEXTURE
vour (by taste assessment); browning po- 1. Melting
tential; texture (melting, non-melting, 2. Non-melting
stony-hard: for better evaluation of the 3. Stony-hard
latter, ethylene production measurement
is suggested); fibrousness (fine, coarse, FLESH ACIDITY
medium); measurement of: sugars (total 1. Acidic
soluble solids); titrable acidity; pH. Stone: 2. Low-acid
adherence to flesh (air-free, free, semi-cling,
cling); size (length, width and breadth);
shape; colour; surface (rough, smooth);
ridge; grooves and pits; propensity to split Phenological classification
or shatter.
CHILLING REQUIREMENT
1. Evergreen (no dormancy under tropical
Morphological and commercial classifications or subtropical climates).
2. From very low (less than 100 CU) to very
Cultivars could be classified under several high (over 1000–1200 CU); most commercial
criteria, depending on scope of evaluation. cultivars ranging from 650 to 900 CU.

TREE USE BLOOM DATE

1. Fruit production Could be reported either as the calendar date
2. Ornamental (flowers, leaves, growth habit) referred as to that particular place, or as the
amount of CU and GDH to accomplish rest
FRUIT TYPE (COMMERCIAL) and to start blooming.
1. Peach (pubescent skin)
2. Nectarine (glabrous skin) RIPE DATE
3. Canning peach (non-melting flesh) As above, it could be recorded as a calendar
date, when the very first fruits (5–10%) accom-
FRUIT SHAPE plish physiological ripening (i.e. they become
1. Round/elongated palatable), or as the number of days from full
2. Flat bloom to the onset of ripening (FDP).

References

Bailey, L.H. (1927) The Standard Cyclopedia of Horticulture. Macmillan, New York.
Bailey, J.S. and French, A.P. (1932) The inheritance of certain characters in the peach. Proceedings of the
American Society for Horticultural Science 29, 127–130.
Bailey, J.S. and French, A.P. (1949) The inheritance of certain fruit and foliage characters in the peach.
Massachusetts Agricultural Experiment Station Bulletin No. 452.
Baipo, W., Yincai, Q., Yongfa, Z. and Hua, C. (1989) The effect of meteorological factors on the pollination,
fertilization and fruit setting of the peach. Acta Horticulturae Sinica 16, 11–15.
Bassi, D. (ed.) (2003) Growth Habits in Stone Fruit Trees. Il Divulgatore, Bologna, Italy.
Bassi, D. and Rizzo, M. (1995) ‘Borgia’ e ‘Lucrezia’, nuove pesche extraprecoci ottenute all’Università di
Bologna. Rivista di Frutticoltura e di Ortofloricoltura 2, 73.
 Botany and Taxonomy 31

Bassi, D. and Selli, R. (1990) Evaluation of fruit quality in peach and apricot. Advances in Horticultural
Science 2, 107–112.
Bassi, D., Gambardella, M. and Negri, P. (1988) Date of ripening and two morphological fruit traits in peach
progenies. Acta Horticulturae 254, 59–66.
Bassi, D., Dima, A. and Scorza, R. (1994) Tree structure and pruning response of six peach growth forms.
Journal of the American Society for Horticultural Science 119, 378–382.
Bassi, D., Mignani, I. and Rizzo, M. (1998) Calcium and pectin influence on peach flesh texture. Acta Horti-
culturae 465/2, 433–438.
Bassi, D., Rizzo, M. and Bassi, L. (2004) Tardigold, mutazione tardiva di Stark Redgold. L’Informatore Agrario
37, 65–66.
Beckman, T.G. and Sherman, W.B. (1996) The non-melting semi-freestone peach. Fruit Varieties Journal 50,
189–193.
Beckman, T.G. and Sherman, W.B. (2003) Probable qualitative inheritance of full red skin color in peach.
HortScience 38, 1184–1185.
Beckman, T.G., Rodriguez Alcazar, J., Sherman, W.B. and Werner, D.J. (2005) Evidence for qualitative
suppression of red skin color in peach. HortScience 40, 523–524.
Begheldo, M., Manganaris, G.A., Bonghi, C. and Tonutti, P. (2008) Different postharvest conditions modulate
ripening and ethylene biosynthetic and signal transduction pathways in stony hard peaches. Postharvest
Biology and Technology 48, 84–91.
Bellini, E. and Scaramuzzi, F. (1976) Monografia delle principali cultivar di pesco, Vol. II. Consiglio Nazionale
delle Ricerche, Florence, Italy.
Bellini, E., Giordani, E., Gianelli, G. and Picardi, E. (2007) The fruit woody species. Descriptor list. Agenzia
Regionale Sviluppo Innovazione Settore Agricolo-Forestale, Florence, Italy.
Bielenberg, D.G., Wang, Y., Fan, S., Reighard, G.L., Scorza, R. and Abbott, A.G. (2004) A deletion affecting
several gene candidates is present in the Evergrowing peach mutant. Journal of Heredity 5, 436–444.
Blake, M.A. (1932) The J.H. Hale as a parent in peach crosses. Proceedings of the American Society for Hor-
ticultural Science 29, 131–136.
Blake, M.A. (1937) Progress in peach breeding. Proceedings of the American Society for Horticultural Science
35, 49–53.
Blake, M.A. and Connors, C.H. (1936) Early results of peach breeding in New Jersey. New Jersey Agricultural
Experiment Station Bulletin No. 599.
Brecht, J.K. and Kader, A.A. (1984) Ethylene production by fruit of some slow-ripening nectarine genotypes.
Journal of the American Society for Horticultural Science 109, 763–767.
Brecht, J.K., Kader, A.A. and Ramming, D.W. (1984) Description and postharvest physiology of some slow-
ripening nectarine genotypes. Journal of the American Society of Horticultural Science 109, 596–
600.
Brovelli, E.A., Brecht, J.K., Sherman, W.B. and Sims, C.A. (1998) Anatomical and physiological responses of
melting-flesh and nonmelting-flesh peaches to postharvest chilling. Journal of the American Society for
Horticultural Science 123, 668–674.
Byrne, D.H., Nikolic, A.N. and Burns, E.E. (1991) Variability in sugars, acids, firmness, and color
characteristics of 12 peach genotypes. Journal of the American Society for Horticultural Science 116,
1004–1006.
Caruso, T. and Sottile, F. (1999) La peschicoltura autunnale in Sicilia: aspetti ambientali, varietali e colturali.
Rivista di Frutticoltura ed Ortofloricoltura 2, 39–46.
Caruso, T., Di Lorenzo, R. and Barone, E. (1992) Il germoplasma del pesco in Sicilia: aspetti genetici e bioa-
gronomici. In: Proceedings of the Symposium ‘Germoplasma Frutticolo’: salvaguardia e valorizzazione
delle risorse genetiche. Consiglio Nazionale delle Ricerche, Alghero, Italy, pp. 285–293.
Chaparro, J.X., Werner, D.J., O’Malley, D. and Sederoff, R.R. (1994) Targeted mapping and linkage analysis of
morphological, isozyme, and RAPD markers in peach. Theoretical and Applied Genetics 87, 805–815.
Chaparro, J.X., Werner, D.J., Whetten, R.W. and O’Malley, D.M. (1995) Inheritance, genetic interaction, and
biochemical characterization of anthocyanin phenotypes in peach. Journal of Heredity 86, 32–38.
Citadin, I., Raseira, M.C.B., Herter, F.G. and Baptista da Silva, J. (2001) Heat requirement for blooming and
leafing in peach. HortScience 36, 305–307.
Connors, C.H. (1920) Some notes on the inheritance of unit characters in the peach. Proceedings of the
American Society for Horticultural Science 16, 24–36.
Connors, C.H. (1921) Inheritance of foliar glands of the peach. Proceedings of the American Society for Hor-
ticultural Science 18, 20–26.
32 D. Bassi and R. Monet

Connors, C.H. (1922) Peach breeding. A summary of results. Proceedings of the American Society for Horti-
cultural Science 19, 108–115.
Crisosto, G.M., Crisosto, C.H. and Watkins, M. (1998) Chemical and organoleptic description of white flesh
nectarines and peaches. Acta Horticulturae 465, 497–505.
De Candolle, A. (1883) L’origine delle piante coltivate. Fratelli Dumolard, Milan, Italy.
Delgado, M. (1998) Plus de sucres pour les douces. L’Arboriculture Fruitiére 519, 21–23.
Demmig-Adams, B. and Adams, W.W. (2002) Antioxidants in photosynthesis and human nutrition. Science
5601, 2149–2153.
Dennis, F.G. (2003). Problems in standardizing methods for evaluating the chilling requirements for the break-
ing of dormancy in buds of woody plants. HortScience 38, 347–350.
Diaz, M.D. (1974) Vegetative and reproductive growth habits of evergreen peach trees in Mexico. Proceedings
of the XIX International Horticultural Congress 18, 525.
Draczynski, M. (1958) The course of pollen differentiation in almond, peach and apricot and the influence of
bud temperature on these processes. Gartenbauwissenshaft 23, 327–341.
Edin, M. and Garcin, A. (1994) Un noveau porte-greffe du pecher Cadaman Avimag. L’Arboriculture Fruitière
475, 20–23.
Erez, A., Fishman, S., Linsley-Noakes, G.C. and Allan, P. (1990) The dynamic model for rest completion in
peach buds. Acta Horticulturae 276, 165–173.
Faust, M. and Timon, B. (1995) Origin and dissemination of peach. Horticultural Reviews 17, 331–379.
Fideghelli, C., Della Strada, G., Quarta, R. and Rosati, P. (1979) Genetic semi-dwarf peach selections. In:
Proceedings of Eucarpia Fruit Section Symposium, Tree Fruit Breeding. INRA, Angers, France, pp. 3–7.
Fogle, H.W. (1977) Self-pollination and its implications in peach improvement. Fruit Varieties Journal 31,
74–75.
Fogle, H.W. and Dermen, H. (1969) Genetic and chimeral constitution of three leaf variegation in the peach.
Journal of Heredity 60, 323–328.
Fogle, H.W. and Faust, M. (1975) Ultrastructure of nectarine fruit surfaces. Proceedings of the American Soci-
ety for Horticultural Science 100, 434–439.
Gage, J. and Stutte, G. (1991) Developmental indices of peach: an anatomical framework. HortScience 26,
459–463.
Gallesio, G. (1817–1839) Pomona Italiana, ossia Trattato degli alberi fruttiferi, Vol. 1. Niccolò Capurro, Pisa, Italy.
Gallesio, G. (2003) Il trattato del pesco di Giorgio Gallesio. In: Baldini, E. (ed.) Gli inediti trattati del pesco e
del ciliegio. Complementi scientifici della ‘Pomona Italiana’ di Giorgio Gallesio. Accademia dei Geor-
gofili, Florence, Italy, pp. 9–146.
Gamberini, A. (2007) Molecular markers and controlling genes of peach flesh texture. PhD thesis, University
of Bologna, Bologna, Italy.
Glenn, D.M., Scorza, R. and Basset, C. (2000) Physiological and morphological traits associated with increased
water use efficiency in the narrow-leaf peach. HortScience 35, 1241–1243.
Goffreda, J.C. (1992) Stony hard gene of peach alters ethylene biosynthesis, respiration and other ripening-
related characters. HortScience 6, 610.
Gradziel, T.M. and Beres, W. (1993) Semidwarf growth habit in clingstone peach with desirable tree and fruit
qualities. HortScience 28, 1045–1047.
Haji, T., Yaegaki, H. and Yamaguchi, M. (2001) Changes in ethylene production and flesh firmness of melting,
nonmelting and stony hard peaches after harvest. Journal of the Japanese Society for Horticultural Sci-
ence 70, 458–459.
Haji, T., Yaegaki, H. and Yamaguchi, M. (2003) Softening of stony hard peach by ethylene and the induction
of endogenous ethylene by 1-aminocyclopropane-1-carboxylic acid (ACC). Journal of the Japanese
Society for Horticultural Science 72, 212–217.
Haji, T., Yaegaki, H. and Yamaguchi, M. (2005) Inheritance and expression of fruit texture melting, non-
melting and stony hard in peach. Scientia Horticulturae 105, 241–248.
Hansche, P.E. (1988) Two genes that induce brachytic dwarfism in peach. HortScience 23, 604–606.
Hansche, P.E. and Boynton, B. (1986) Heritability of enzymatic browning in peaches. HortScience 21, 1195–
1197.
Harrold, T.J. (1935) Comparative study of developing and aborting fruits of Prunus persica. Botanical Gazette
96, 505–520.
Hedrick, U.P. (1917) The Peaches of New York. J.B. Lyon Company Printers, Albany, New York.
Herrero, M. and Arbeloa, A. (1989) Influence of the pistil on pollen tube kinetics in peach (Prunus persica).
American Journal of Botany 76, 1441–1447.
 Botany and Taxonomy 33

Hesse, C.O. (1975) Peaches. In: Janick, J. and Moore, J.N. (eds) Advances in Fruit Breeding. Purdue Univer-
sity Press, West Lafayette, Indiana, pp. 285–335.
Knight, R.L. (1969) Abstract Bibliography of Fruit Breeding and Genetics to 1965. Prunus. Eastern Press, London.
Knowlton, H.E. (1924) Pollen abortion in peach. Proceedings of the American Society for Horticultural
Science 21, 67–69.
Lammerts, W.E. (1945) The breeding of ornamental edible peaches for mild climates. I. Inheritance of tree and
flower characters. American Journal of Botany 32, 53–61.
Lesley, J.W. (1940) A genetic study of saucer fruit shape and other characters in the peach. Proceedings of the
American Society for Horticultural Science 37, 218–222.
Lesley, J.W. (1957) A genetic study of inbreeding and of crossing inbred lines in peaches. Proceedings of the
American Society for Horticultural Science 70, 93–103.
Lester, D.R., Sherman, W.B. and Atwell, B.J. (1996) Endopolygalacturonase and the melting flesh (M) locus in
peach. Journal of the American Society for Horticultural Science 121, 231–235.
Li, Z. (1984) Peach germplasm and breeding in China. HortScience 19, 348–351.
Lilien-Kipnis, H. and Lavee, S. (1971) Anatomical changes during the development of ‘Ventura’ peach fruits.
Journal of Horticultural Science 46, 103–110.
Lilleland, O. (1933) Growth study of the peach fruit. Proceedings of the American Society for Horticultural
Science 29, 8–12.
Linné (Linnaei), C. (1758) Systema Naturae, 10th edn. Laurentii Salvii, Holmae, Stockholm.
Liverani, A. and D’Alessandro, D. (1999) La qualità gustative dei frutti nell’attività di miglioramento genetico
del pesco presso l’ISF di Forlì. Rivista di Frutticoltura ed Ortofloricoltura 2, 30–37.
Liverani, A., Giovannini, D., Brandi, F. and Merli, M. (2003) Le pesche subacide. L’Informatore Agrario 31,
43–48.
Manaresi, A. and Draghetti, A. (1915) Influenza del germoglio ascellare sullo sviluppo e sulla composizione
del frutto del pesco. Bollettino Associazione Orticola Professionale Italiana 1, 1–4.
Marini, R.P. and Sowers, D.L. (1994) Peach fruit weight is influenced by crop density and fruiting shoot
length but not position on the shoot. Journal of the American Society for Horticultural Science 119,
180–184.
Massonié, G., Maison, P., Monet, R. and Grasselly, G. (1982) Resistance au puceron vert du pêcher Myzus
persicae Sulzer (Homoptera Aphididae) chez Prunus persicae (L.) Batsch et d’autres espèces de Prunus.
Agronomie 2, 63–70.
Meader, E.M. and Blake, M.A. (1939) Some plant characteristics of the second generation progeny of Prunus
persica and Prunus kansuensis crosses. Proceedings of the American Society for Horticultural Science
37, 223–231.
Mehlenbacher, S.A. and Scorza, R. (1986) Inheritance of growth habit in progenies of ‘Com-Pact Redhaven’
peach. HortScience 21, 124–126.
Mignani, I., Ortugno, C. and Bassi, D. (2006) Biochemical parameters for the evaluation of different peach
flesh types. Acta Horticulturae 713, 441–448.
Moing, A., Svanella, L., Monet, R., Rothan, C., Diakou, P., Gaudillere, J.P. and Rolin, D. (1998) Organic acid
metabolism during the fruit development of two peach cultivars. Acta Horticulturae 465, 425–432.
Moing, A., Pöessel, J.L., Svanella-Dumas, L., Loonis, M. and Kervella, J. (2003) Biochemical basis of low
fruit quality of Prunus davidiana, a pest and disease donor for peach breeding. Journal of the American
Society for Horticultural Science 128, 55–62.
Monet, R. (1967) Contribution à l’étude génétique du pêcher. Annales de l’Amélioration des Plantes 17, 5–11.
Monet, R. (1979) Transmission génétique du caractér ‘fruit doux’ chez le pêcher. Incidence sur la sélection
pour la qualiter. In : Proceedings of Eucarpia Fruit Section: Tree Fruit Breeding. INRA, Angers, France,
pp. 273–279.
Monet, R. (1983) Le pêcher. Genetique et physiolgie. Masson, Paris, France.
Monet, R. (1989) Peach genetics: past, present and future. Acta Horticulturae 254, 49–57.
Monet, R. and Bastard, Y. (1982) Une anomalie du fonctionnement de l’apex à hérédité mendélienne chez le
pêcher (Prunus persica (L.) Batsch). Agronomie 2, 103–106.
Monet, R. and Massonié, G. (1994) Déterminisme génétique de la résistance au puceron vert (Myzus persi-
cae) chez le pêcher. Résultats complémentaires. Agronomie 2, 177–182.
Monet, R. and Salesses, G. (1975) Un nouveau mutant de nanisme chez le pecher. Annales des Amélioration
des Plantes 25, 353–359.
Monet, R., Bastard, Y. and Gibault, B. (1988) Etude génétique du caractère ‘port pleureur’ chez le pêcher.
Agronomie 8, 127–132.
34 D. Bassi and R. Monet

Monet, R., Guye, A. and Roy, M. (1996) Effect of inbreeding and crossing inbred lines on the weight of peach
fruit. Acta Horticulturae 374, 77–82.
Morgutti, N., Negrini, F.F., Nocito, A., Ghiani, D., Bassi, D. and Cocucci, M. (2006) Changes in endo-
polygalacturonase levels and characterization of a putative endo-PG gene during fruit softening in
peach genotypes with nonmelting flesh and melting flesh fruit phenotypes. New Phytologist 171, 315–
328.
Okie, W.R. (1998) Preliminary descriptions of five new peach genes. Acta Horticulturae 465, 107–110.
Okie, W.R. and Prince, V.E. (1982) Surface features of a novel peach × nectarine hybrid. HortScience 17,
66–67.
Okie, W.R. and Rieger, M. (2003) Inheritance of venation pattern in Prunus ferganensis × persica hybrids. Acta
Horticulturae 622, 261–264.
Okie, W.R. and Scorza, R. (2002) Breeding peach for narrow leaf width. Acta Horticulturae 592, 137–141.
Okie, W.R., Ramming, D.W. and Scorza, R. (1985) Peach, nectarine, and other stone fruit breeding by the
USDA in the last two decades. HortScience 20, 633–641.
Peace, C.P., Crisosto, C.H. and Gradziel, T.M. (2005a) Endopolygalacturonase: a candidate gene for freestone
and melting flesh in peach. Molecular Breeding 16, 21–31.
Peace, C.P., Ahmad, R., Gradziel, T.M., Dundekar, A.M and Crisosto, C.H. (2005b) The use of
molecular genetics to improve peach and nectarine post-storage quality. Acta Horticulturae 682,
403–409.
Pérez, G.S. (1990) Relationship between parental blossom season and speed of seed germination. Hort-
Science 25, 958–960.
Pérez, S., Montes, S. and Mejìa C. (1993) Analysis of peach germplasm in Mexico. Journal of the American
Society for Horticultural Science 118, 519–524.
Pisani, P.L. and Roselli, G. (1983) Interspecific hybridization of Prunus persica × P. davidiana to obtain new
peach rootstocks. Genetica Agraria 1/2, 197–198.
Ramming, D.W. (1985) In ovulo embryo culture of early maturing Prunus. HortScience 20, 419–420.
Ramming, D.W. (1991) Genetic control of a slow-ripening fruit trait in nectarine. Canadian Journal of Plant
Science 71, 601–603.
Ramming, D.W. and Tanner, O. (1987) Goldcrest peach. Fruit Varieties Journal 41, 52–53.
Rehder, A. (1990) Manual of Cultivated Trees and Shrubs Hardy in North America. Dioscorides Press, Port-
land, Oregon.
Reimer, F.S. (1906) The honey peach group. Florida Agricultural Experiment Station Bulletin No. 73.
Richardson, E.A., Seeley, S.D. and Walker, D.R. (1974) A model for estimating the completion of rest for Red-
haven and Elberta peach trees. HortScience 9, 331–332.
Rivers, S. (1906) The cross-breeding of peaches and nectarines. Report on Third International Conference on
Genetics. Royal Horticultural Society, London, pp. 463–467.
Robertson, J.A., Meredith, F.I. and Scorza, R. (1988) Characteristics of fruit from high and low-quality peach
cultivars. HortScience 23, 1032–1034.
Robertson, J.A., Horvat, R.J., Lyon, B.G., Meredith, F.I., Senter, S.D. and Okie, W.R. (1990) Comparison of
quality characteristics of selected yellow and white-fleshed peach cultivars. Journal of Food Science 55,
1308–1311.
Rodriguez, A.J., Sherman, W.B., Scorza, R. and Wisniewski, M. (1994) ‘Evergreen’ peach, its inheritance and
dormant behavior. Journal of the American Society for Horticultural Science 119, 789–792.
Rodriguez, G.A. and Sherman, W.B. (1990) ‘Oro A’ peach germplasm. HortScience 25, 128.
Rodriguez, J. and Sherman, W.B. (1986) Relationship between parental flower bud set and seedling preco-
ciousness in peach and nectarine, Prunus persica (L.) Batsch. Fruit Varieties Journal 40, 8–12.
Roselli, G., Pisani, P.L. and Cerulli, M. (1985) Observations on the performance of various Prunus davidi-
ana × Prunus persica hybrids as rootstocks for peach. L’Informatore Agrario 12, 75–81.
Saunier, R. (1973) Contribution a l’étude des relations existant entre certains caractères a déterminisme géné-
tique simple chez le pêcher et la sensibilité a l’oidium, Sphaeroteca pannosa (Wallr) Lev. des cultivars
de cette espéce. Annales des Amélioration des Plantes 23, 235–243.
Scorza, R. (1984) Characterization of four distinct peach tree growth types. Journal of the American Society
for Horticultural Science 109, 455–457.
Scorza, R. and Okie, W.R. (1991) Peaches (Prunus). In: Moore, J.N. and Ballington, J.R. Jr (eds) Genetic
resources of temperate fruit and nut crops. Acta Horticulturae 290, 177–231.
Scorza, R. and Sherman, W.B. (1996) Peaches. In: Janick, J. and Moore, J.N. (eds) Fruit Breeding. Vol. I. Tree
and Tropical Fruits. Wiley, New York, pp. 325–440.
 Botany and Taxonomy 35

Scorza, R., Mehlenbacher, S.A. and Lightner, G.W. (1985) Inbreeding and coancestry of freestone peach cul-
tivars of the eastern United States and implications for peach germplasm improvement. Journal of the
American Society for Horticultural Science 110, 547–552.
Scorza, R., Lightner, G.W. and Liverani, A. (1989) The Pillar peach tree and growth habit analysis of com-
pact × Pillar progeny. Journal of the American Society for Horticultural Science 114, 991–995.
Scorza, R., Bassi, D. and Liverani, A. (2002) Genetic interaction of pillar (columnar), compact and dwarf
peach tree genotypes. Journal of the American Society for Horticultural Science 127, 254–261.
Scott, D. and Cullinan, F. (1942) The inheritance of wavy leaf character in the peach. Journal of Heredity 33,
293–295.
Scott, D.H. and Weinberger, J.H. (1944) Inheritance of pollen sterility in some peach varieties. Proceedings of
the American Society for Horticultural Science 45, 229–232.
Sherman, W.B., Topp, B.L. and Lyrene, P.M. (1990) Non melting flesh for fresh market peaches. Proceedings
of the Florida State Horticultural Society 103, 293–294.
Szabò, Z. and Nyéki, J. (1999) Self-pollination in peach. International Journal of Horticultural Science 5,
76–78.
Tatsuki, M., Haji, T. and Yamaguchi, M. (2006) The involvement of 1-aminocyclopropane-1-carboxylic acid
synthase isogene, Pp-ACS1, in peach fruit softening. Journal of Experimental Botany 57, 1281–1289.
Tatsuki, M., Haji, T. and Yamaguchi, M. (2007) The peach 1-aminocyclopropane-1-carboxylic acid synthase
isogene, Pp-ACS1, is required for fruit softening. In: Ramina, A., Chang, C., Giovannoni, J., Klee, H., Pera-
ta, P. and Woltering, E. (eds) Advances in Plant Ethylene Research. Proceedings of the 7th International
Symposium on the Plant Hormone Ethylene. Springer, Wageningen, The Netherlands, pp. 175–180.
Tourjie, K.R., Barret, D.M., Romero, M.V. and Gradziel, T.M. (1998) Measuring flesh colour variability among
processing clingstone peach genotypes differing in carotenoid composition. Journal of the American
Society for Horticultural Science 123, 433–437.
Toyama, T.K. (1980) The pollen receptivity period and its relation to fruit setting in the stone fruits. Fruit Variet-
ies Journal 34, 2–4.
Tukey, H.B. (1933) Artificial culture of sweet cherry embryos. Journal of Heredity 24, 7–12.
Tukey, H.B. (1935) Artificial culture methods for isolated embryos of deciduous fruits. Proceedings of the
American Society for Horticultural Science 32, 313–322.
Van Well, R.G. (1974) Com-Pact Redhaven. Fruit Varieties Journal 28, 37.
Vavilov, N.I. (1951) The Origin, Variation, Immunity and Breeding of Cultivated Plants. Selected Writings of
N.I. Vavilov. Chronica Botanica Company, Waltham, Massachusetts.
Ventura, M., Courvoisier, V. and Sansavini, S. (1995) Relazione tra scambi gassosi e qualità di pesche e net-
tarine durante la maturazione. In: Proceedings of ‘XXII Convegno Peschicolo’. Chambers of Commerce
of Forlì-Cesena and Ravenna, Cesena, Italy, pp. 71–80.
Wang, Y.L. (1985) Peach growing and germplasm in China. Acta Horticulturae 173, 51–55.
Wang, Y., Reighard, G.L., Scorza, R., Jenkins, T.C. and Line, M.J. (2003) Characterizing the evergrowing phe-
notype in peach. Acta Horticulturae 618, 455–462.
Watkins, R. (1976) Cherry, plum, peach, apricot and almond. In: Simmonds, N.W. (ed.) Evolution of Crop
Plants. Longman, London, pp. 242–247.
Weinberger, J.H. (1944) Characteristics of the progeny of certain peach varieties. Proceedings of the American
Society for Horticultural Science 45, 233–238.
Weinberger, J.H. (1950) Chilling requirement of peach varieties. Proceedings of the American Society for
Horticultural Science 56, 122–128.
Wen, I.C., Koch, K.E. and Sherman, W.B. (1995) Comparing fruit and tree characteristics of two peaches and
their nectarine mutants. Journal of the American Society for Horticultural Science 120, 101–106.
Werneck, H.L. (1956) Romischer und vorromischer Wein- und Obstbau in Ostereichischer Donauraum. Ver-
handlungen der Zoologisch-Botanischen-Gesellschaft, Wien.
Werner, D.J. and Chaparro, J.X. (2005) Genetic interaction of pillar and weeping peach genotypes. Hort-
Science 40, 18–20.
Werner, D.J. and Creller, M.A. (1997) Inheritance of sweet kernel and male sterility. Journal of American Soci-
ety for Horticultural Science 122, 215–217.
Werner, D.J., Creller, M.A. and Chaparro, J.X. (1998) Inheritance of the blood-flesh trait in peach. HortScience
33, 1243–1246.
Wickson, E.J. (1889) California Fruits 308.
Yamaguchi, M., Yoshida, M. and Kyotani, H. (1984) Studies on the distribution of ripening time of certain
progenies in peach breeding. Bulletin of the Fruit Tree Research Station, Series A 11, 15–33.
36 D. Bassi and R. Monet

Yoshida, M. (1970) Genetical studies on the fruit quality of peach varieties. 1. Acidity. Bulletin of the Tree
Research Station Series A 9, 1–15.
Yoshida, M. (1976) Genetical studies on the fruit quality of peach varieties. 3. Texture and keeping quality.
Bulletin of the Tree Research Station, Series A 3, 1–16.
Yoshida, M. (1987) Peach germplasm. Kajitsu Nippon 42, 70–74.
Yoshida, M., Yamane, K., Ijiro, Y. and Fujishige, N. (2000) Studies on ornamental peach cultivars. Bulletin of
the College of Agriculture, Utsunomiya University, Japan 17, 1–14.
Zielinski, Q.B. (1955) Modern Systematic Pomology. W.M.C. Brown, Dubuque, Iowa.
2 History of Cultivation and Trends in China

Hongwen Huang,1,2 Zhongping Cheng,1 Zhonghui Zhang1 and

Ying Wang1
1Wuhan Botanical Garden/Wuhan Institute of Botany, Chinese Academy of Sciences,
People’s Republic of China
2South China Botanical Garden, Guangzhou, Chinese Academy of Sciences, People’s

Republic of China

2.1 Origin of the Peach 37

2.2 History of Peach Cultivation in China 38
Pre-Qin Dynasty (1100–221 BC) 38
Eastern and Western Han Dynasty (222 BC–220 AD) 40
From Wei–Jin Dynasty to Sui–Tang Dynasty and Five Dynasties period (221–960 AD) 40
Song, Yuan, Ming, Qing Dynasties and Republican period (961–1948 AD) 42
2.3 Current Status of Chinese Peach Production and Trends in China 44
Peach germplasm collection, repositories, evaluation and utilization in China 44
Main peach cultivars, peach production and growing regions in China 45
Rapid development of greenhouse production 48
Peach postharvesting, processing and marketing in China 49
Problems faced by the Chinese peach industry 51
Current trends of Chinese peach production 52
2.4 Summary/Conclusion 57

2.1 Origin of the Peach (Chen, 1994). A similar archaeological finding

The peach originated in China (Wang and
Zhuang, 2001), where it is a symbol of long
life (Fig. 2.1/Plate 25). Numerous pieces of
evidence have revealed that China has the
longest history of peach cultivation in the
world. One discovery demonstrated that peach
growing in China dates back to Neolithic
times. When a Neolithic village site was dis-
covered in Hemudu village, Yujao city, Zheji-
ang province in 1973, finds included wild
peach stones dating back to 6000–7000 BC
in Taixi village of Gaochen city, Hebei province,
at the site of ruins from the Shang Dynasty
(1600–1100 BC), revealed two peach stones
measuring 1.6 cm × 1.0 cm and 2.0 cm × 1.2
cm. Their shape, size and surface groove pat-
terns were almost the same as those of current
peach cultivars in China. An expedition con-
ducted by the Chinese Academy of Sciences
during 1973–1976 discovered tremendously
diverse genetic resources of wild peach that
are still widely grown in large areas of China,
including Tibet, Gansu, eastern Shaanxi,

© CAB International 2008. The Peach: Botany, Production and Uses

(eds D.R. Layne and D. Bassi) 37
38 H. Huang et al.
Fig. 2.1. A Chinese painting by Ma Tai (1885–
1935), telling a story that an old man with white
hair but a young, boyish face always steals
peaches. People make fun of this long-lived man,
but he explains ‘Peach is good for my health’.
southern Henan, south-western Sichuan and
western Yunnan provinces (Qu and Sun,
1990). Tibetan peaches (Prunus mira Koehnes)
were found in Jiacha and Lang counties of
south-eastern Tibet. Some trees having trunk
circumferences of about 6 m appeared to be
more than 300 years old. A fruit tree survey
team of the Tibet crop resource expedition
during 1981–1982 also found an ancient
Tibetan peach tree with a height of 21 m and a
trunk circumference of 10 m, which was
believed to be more than 1000 years old. This
tree was located in Changdu region of Tibet,
where the Hengduan Mountains–Sanjiang
(Three Rivers) area was being geo-botanically
investigated (Duan et al., 1983). Obviously,
Tibet and Gansu province, where P. mira and
Prunus kansuensis Rehd. are native, should be
regarded as one of the original native centres
for peach. The peach has a remarkably exten-
sive distribution throughout China: from Tai-
wan and southern Guangdong provinces in
the subtropics, to cold temperate regions as
far north as in Yanbian, Jilin province (Wang
and Zhuang, 2001); from the west and south-
west regions as far as Xinjiang and Tibet auton-
omous districts, to east regions as far as all
coastal provinces in China. However, commer-
cial peach cultivation is limited within the lati-
tude range of 23–50°N (Fig. 2.2/Plate 26).
2.2 History of Peach Cultivation in China
The peach is one of the most ancient domesti-
cated fruits in China. As early as 4000 years
ago, the value of peach had been recognized
and exploited by the Chinese with extensive
efforts on natural selection and domestica-
tion. Several unique historical phases of Chi-
nese peach domestication and cultivation are
summarized in the following.
Pre-Qin Dynasty (1100–221 BC)
There may be even earlier descriptions of
peach in ancient Chinese texts. ‘桃 pro-
nounced Tao’ (Chinese name for the outdated
genus Amygdalus) refers to peach and is
described in the ShiJing (The Book of Odes or
The Book of Songs), written between 1100 and
600 BC (Anon., 11th–6th century BC). The tao
is described in the ShiJing – Weifeng chapter as
‘Peach growing in garden, its fruit for eating’,
indicating that peach was cultivated in China
 Cultivation and Trends in China 39

Fig. 2.2. Chinese peach production regions: (I) north-west drought region; (II) northern China
plain region; (III) Changjiang River humid region; (IV) Yunnan–Guizhou high plateau cold region;
(V) Qinghai–Tibet plateau cold peach region; (VI) north-eastern China cold region; (VII) southern
China subtropical region.

3000 years ago. In the ShiJing – ZhouNan chap-
ter, the biological characteristics of peach are
concisely described as ‘桃之夭夭，灼灼其
华。桃之夭夭，其叶蓁蓁。桃之夭夭，有贲其
实’, illustrating beautiful fire-like flowers
when blooming, flourishing foliage and dense
fruiting, symbolizing a family’s prosperity,
happiness and luck. The ShiJing –DaYa chap-
ter describes peach as ‘投我以桃，报之以李。
园中有桃，有贲其实’, clearly evidencing that
peach was widely cultivated and fruit were
produced plentifully during that time. Later,
a historiography book LüShiChunQiu (300 BC)
depicts peach blooming in spring as ‘仲春之
月桃始华’ (Lü, 300 BC). The earliest illustra-
tion of peach cultivation and ecological
requirements in Chinese ancient literature
probably occurs during the Zhanguo period
(Warring States, 500–300 BC). In the encyclo-
paedia GuanZi – DiYuan volume (Guan, 5th
century BC), it is written ‘五沃之土，宜彼群
木，其桃其李’, explaining that peach cultiva-
tion requires good soils and peach responds
to high fertilization. It concisely illustrates a
40 H. Huang et al.
relationship between peach growth and soil
conditions. In addition, HanFeiZhi – WaiChu
volume 33 (Han, 280–233 BC) describes ‘子产
治郑，桃李之荫于街者莫援也’, which explains
that besides use as edible fruits, the peach
was also used for landscaping as a shade tree
in the current Xinzheng area, Henan prov-
ince. During this pre-Qin Dynasty period,
peach was widely grown in southern China.
According to the Chinese Agricultural Archa-
eological Plate Collection, peach stones have
been discovered at many archaeological
locations in southern China, from the East-
ern Zhou Dynasty (770 BC) at Jiangling, Hubei
province and from the Zhanguo period
(Warring States, 475–221 BC) at Pujiang,
Sichuan province and at Hezhang, Guizhou
province.
Eastern and Western Han Dynasty
(222 BC–220 AD)
During this historical period, cultivar selec-
tion and cultural techniques were developed,
resulting in a new era of peach domestication
in China. Fifteen records related to peach are
identified in ShiJi (a history annals, 100 Bc)
(Sima, 1st century BC), 19 records in HanShu
(history annals of the Han Dynasty, 1st cen-
tury AD) (Ban, 39–92 AD) and 14 in Post-Han-
Shu (history annals of the late Han Dynasty,
3rd century AD). These records encompass a
variety of peach-related delineations from
town and street names, mountain and river
names and even official titles. In ErYa (ancient
dictionary for terms and names) – Shimu
chapter (notes in trees) (Anon., 2nd century
BC), there are records of species, varieties and
cultivars for peaches described as ‘旄，冬桃’
and ‘榹，山桃’, indicating the two most impor-
tant species of Prunus persica and Prunus davi-
diana of modern peach production. In XiJingZaJi
(a miscellanea) edited by Ge Hong (Ge, 1st
century AD), the fruit trees in the emperor’s
garden (current Xian, Shaanxi province) are
described, including many peach varieties:
‘Qin Tao’ (Shaanxi peach), ‘Si Tao’ (peach),
‘Xiang He Tao’ (locally named peach), ‘Shuang
Tao’ (frost peach), ‘Jin Cheng Tao’ (locally
named peach), ‘Qi Di Tao’ (fancy petiole
peach) and ‘Zi Wen Tao’ (purple paper peach).
Thus, many locally selected peach varieties
existed in China more than 2000 years ago.
The relationship of peach blooming to
local climate is written in Pre-HanShu (dynasty
annals) – GouXu Records as ‘桃方华时，即有
雨水，. . . . . ，谓之桃花水耳’,
. indicating a
very similar rainy season during P. persica
blooming as normally occurs at the present
time in south and south-central China. Peach
culture techniques were also developed dur-
ing this period. SiMinYueLing (farming calen-
dar) written by Cui Shi during the Eastern Han
Dynasty (Cui, 2nd century AD) provides ‘正
月…自朔及晦可移诸树，. . . 唯有果实者及望而
止’, indicating that peach trees should be
planted or transplanted in January in the Lunar
calendar, but fruiting trees should not be trans-
planted. It clearly suggests that peach planting
should be done in February and good peach
production requires fertile soils. Wang Bao
even records a high-density planting of peach
in TongYue from the Western Han Dynasty
(Wang, 1st century AD), stating ‘种植桃、
李. . . . . ，三丈一树，八尺为行，果类相似
. ，纵
横相当’ that suggests a 6.93 m × 1.85 m spacing.
From Wei–Jin Dynasty to Sui–Tang Dynasty
and Five Dynasties period (221–960 AD)
The traditional Chinese peach culture was
established during these centuries. New pro-
duction techniques were advanced and new
uses for peaches were further exploited. First,
peach cultivar development by seedling selec-
tion and domestication of wild trees resulted
in a large number of new cultivar releases. For
example, GuangZhi written by Guo Yigong in
the Eastern Jin period (Guo, 4th century AD)
records ‘there are winter peach, summer white
peach, autumn white peach and many local
peaches cultivated in low reach region of
HuangHe (Yellow River), among many beauti-
ful peaches having an outstanding autumn
dark-red peach’. YeZhongJi written by Lu Hui
in the Western Jin period (Lu, 3rd century AD)
describes ‘a hook-nose peach weighting one
kilo growing at ShiHu garden’ (currently Lin-
Zhang region, Hebei province). During the
Northern Wei period (386–534 AD), a very
famous book QiMinYaoShu (encyclopaedia
 Cultivation and Trends in China
for living) written by Jia Sixie (Jia, 533–544 AD)
lists more than ten cultivars previously
described and also adds a new variety, ‘Nai
Tao’. In addition, from the Wei–Jin period,
records of high-quality peach cultivars are
found in TaoFu (peach poetry) by Fu Xuan (Fu,
2nd century AD) in Western Jin, such as ‘early
summer ripening’, suggesting an existence of
early cultivars, and ‘sweet and crispy’ for non-
melting peaches. Furthermore, selection and
cultivation for ornamental and yellow-fleshed
peaches are recorded during the Tang Dynasty
(618–907 AD); for example, KuaiYuanTianBaoYi-
Shi (a historiography book) describes a ‘new
ornamental peach with thousand-leaf like flow-
ers was introduced into the Emperor’s garden’.
TangShu (history annals of the Tang Dynasty)
records: ‘In ZhenGuan 21 [Tang Dynasty calen-
dar, 647 AD], western KangJu country [now the
north-west Xinjiang autonomous region] pays
tribute to a goose-egg sized and gold coloured
peach, called “golden peach”’.
Second, knowledge about tree biology
and propagation was extensively improved,
setting the foundation for the traditional sys-
tem of Chinese peach cultivation. QiMin-
YaoShu (encyclopaedia for living; Jia, 533–544
AD) summarizes a cultivated peach tree life
cycle that is very similar to modern peach
production. ‘桃性早实，三岁结子，七八年便
老，老则子细，十年则死’ illustrates that
peach is precocious, fruiting at age 3 years,
production declines at age 7–8 years when
fruits are getting smaller, and trees die in 10
years. For seed germination and seedling
propagation, seed stratification was well
understood and traditional methods were
well developed at this time. These are illus-
trated in QiMinYaoShu: ‘桃熟时，于墙南阳中
暖处，深宽为坑。选取好桃数十枚，劈其核，
即内牛粪中，头向上，取好烂粪和土厚覆之，
令厚尺余。至春桃始幼时，徐徐拨去粪土，皆
应生芽，合取核种之，万不一失。其余以熟粪
粪之，则益桃味’, indicating that seed stratifi-
cation was crucial to embryo development
for seed germination and that fermented
manures are good both for seed stratification
and for improvement of peach fruit flavour
and overall quality. In addition, a layering
propagation method is also developed and
summarized in the book, detailing shoot
types and soils that should be used.
41
Third, planting and transplanting are
precisely summarized in QiMinYaoShu as ‘以
锹合土掘移之’, encouraging planting and
transplanting seedlings with soil balls hold-
ing the roots. Various peach pruning methods
had also been developed and are documented
in the book, such as the application of girdling
or mechanical injury to the tree trunk to sup-
press vegetative growth, increase fruiting and
raise production. Some of these methods are
concisely illustrated in QiMinYaoShu as ‘桃性
皮急，四年以上，宜以刀竖 其皮’, explaining
that the ring bark method should not be used
until trees are 4 years old and cutting should
be careful on the bark. This demonstrates that
ancient Chinese peach growers had a good
understanding of tree reproduction biology and
management practices in peach production.
Finally, frost protection and pest control
techniques were developed along with prac-
tices in peach orchard management. This
overall frost protection method for all fruit
trees including peach is well documented in
QiMinYaoShu (Jia, 533–544 AD) as ‘凡五果（包
括桃树）花盛时遇霜，则无子。常预于园中，
往往贮恶草生粪，天雨新晴，北风寒切，是夜
必霜。此时放火作熅，少得烟气，则免于霜
矣。’, illustrating that all five kinds of species
(including peach) are vulnerable to frost dam-
age during blooming. Frost damage to flow-
ers will cause production failure. To prevent
frost damage, orchards need to be stored with
straws and manures. Whenever a halt in rain
occurs with a clean sky and cold wind from
the north, it signals an overnight frost. Burn-
ing straw mixed with manures and releasing
smog will prevent frost damage. This smok-
ing method for preventing flower frost is still
one of the major applications widely used in
peach orchards in northern China. Applica-
tion of pest control measures also began dur-
ing these centuries. As documented in
QiMinYaoShu: ‘凡五果及桑正月一日鸡鸣时，
把火遍照其下，则无虫灾’, saying that in culti-
vation of the five kinds of species and mulberry,
lighting up the overall orchard using torches in
the Chinese New Year will prevent insect dam-
age. In fact, this idea and method underline the
principle that the current application of ultravi-
olet light for insect control is based on.
The above ancient Chinese literature
during the time of the Wei–Jin to Sui–Tang
42 H. Huang et al.
dynasties and the Five Dynasties period
probably sheds light only on the tip of the
iceberg of a rich Chinese peach cultivation
history, abundant natural resources and well-
developed understanding of cultural technol-
ogy. Some historical knowledge about
germplasm resources, tree biology, propaga-
tion and orchard management is still worthy
of further study.
Song, Yuan, Ming, Qing Dynasties and
Republican period (961–1948 AD)
During these 1000 years, domestic and inter-
national exchanges in agriculture played an
important role in Chinese peach production
and development. Peach production regions
were continuously expanding and many new
production regions were developed. As infor-
mation and technology was spread domesti-
cally, and to some extent abroad, peach
culture developed further during this time.
Over many years, Chinese peach growers
systematically domesticated and continu-
ously selected improved germplasm. This
resulted in substantial improvements of peach
culture and production in China. For example,
LuoYangHuaMuJi (LuoYang flower and tree
collections) written by Zhou ShiHou (1081 AD)
records 30 peach varieties: ‘Xiao Tao’ (small
peach), ‘Shiyue Tao’ (October peach), ‘Dong
Tao’ (winter peach), ‘Pan Tao’ (flat peach),
‘Qianye Tao’ (thousand-leaf peach), ‘Chan
Tao’ (twisted peach), ‘Erse Tao’ (double-
coloured peach), ‘Hehuan Erse Tao’ (dual-
colour happiness peach), ‘Qiangyefei Tao’
(thousand-leaf pink peach), ‘Dayu Tao’ (big
royal peach), ‘Baiyu Tao’ (white royal peach),
‘Jin Tao’ (golden peach), ‘Yin Tao’ (silver
peach), ‘Bai Tao’ (white peach), ‘Kunlun Tao’
(Kunlun Mt. peach), ‘Hanli Tao’ (big peach),
‘Yanzhi Tao’ (crimson peach), ‘Zao Tao’ (early
peach), ‘You Tao’ (smooth and waxy skin
peach), ‘Renmian Tao’ (people face peach),
‘Mi Tao’ (honey peach), ‘Pingding Tao’ (non-
tip peach), ‘Pang Tao’ (fat peach), ‘Ziye DaTao’
(purple-leaved big peach), ‘Li Tao’ (gift peach),
‘Fang Tao’ (square peach), ‘Fenzhou Tao’
(local township named), ‘Putian Tao’ (local
township named), ‘Hongrang Tao’ (red-fleshed
peach) and ‘Guang Tao’ (non-pubescence peach).
The honey peach (typical southern, melting,
low-acid type) and the non-tip peach are prob-
ably the progenitor varieties that all southern
Chinese peaches are derived from. The Yuan
Dynasty’s WangZhen’s Farming Book written
by Wang Zhen in the 14th century (Wang,
1313 AD) lists two additional early peach vari-
eties, ‘Luosi white’ (fine white) and a late
variety ‘Guoyan red’ (passing wild goose
red). The most famous Chinese pharmaco-
poeia is the BenCao GangMu written by Li Shi-
zhen during the Ming Dynasty (Li, 1578 AD),
which classifies peach varieties by colours,
fruit shapes and maturity date into different
categories, saying ‘桃品种甚多，其花有红、
紫、白、千叶二色，其实有红桃、绯桃、碧
桃、缃桃、白桃、乌桃、金桃、银桃、胭脂
桃，皆以色名者也；有锦桃、油桃、御桃、方
桃、匾桃、偏核桃，皆以形名者也；有五月
桃、十月冬桃、秋桃、霜桃，皆以时名者也’,
meaning that there are many varieties of
peaches. The flower colours vary from red,
purple, white to double colours. The fruit
colour variations range from red to pure
white. Varieties are named after their colours
(including pink peach, light-pink peach, crim-
son peach, red peach, dark-red peach, purple
peach, golden peach, silver peach and white
peach); also, they are named after their shapes
and appearance (including tip peach, smooth
peach, royal-type peach, square peach and
flat peach); in addition, some are named after
their maturity (including May peach, October
peach, autumn peach, winter peach, etc.).
Slightly later at the end of the Ming Dynasty,
QunFangPu (Florilegium) written by Wang
Xiangji (Wang, 1621 AD) gives detailed descrip-
tions for some known cultivars, besides list-
ing previous cultivars; for example, flat peach
is described as ‘shape like a cake, taste sweet’.
It also records details about Shanghai honey
(melting) peach as ‘it is native to Shanghai,
but best peaches are produced from the GuSh-
angBaoXi garden with sweet taste little less
than lychee’. ShuiMiTaoPu (melting honey
peach register) by Chu Hua in the Qing
Dynasty (Chu, 1813 AD) records: ‘Melting peach
was from the Gu’s fragrance garden in Ming
Dynasty, it is juicy and tastes sweet, so called
melting peach’. The origin of this type of
peach was unknown, but probably derived
from peaches from Beijing or Kaifeng.
 Cultivation and Trends in China
Peach production in the Qing Dynasty
was further expanded and some major pro-
duction regions included Taiyuan, Shanxi
province; Kaifeng, Luoyang, Shangqiu, Henan
province; Chengdu, Sichuan province; Hang-
zhou, Zhejiang province; Hejian, Shenxian,
Suning, Hebei province; Feicheng, Shandong
province; and Nanhui, Baoshan, Shanghai
city. Until the Republican period (1920s), new
peach production regions were developing
very rapidly, extending into many new areas
including Wuxian, Wuxi, Yangzhou, Zhenji-
ang, Jiangsu province (eastern China); Suzhou,
Dangshan, Anhui province (central eastern
China); Ningling, Yanshi, Henan province
(central northern China); Fenghua, Zhejiang
province (eastern China); Zibo, Shandong
province (northern China); Taigu, Shanxi prov-
ince (north-western China); and central Shaanxi
and eastern Gansu province (western China).
During these ten centuries, peach culture
technology had also developed to a new level.
The documentations recorded in the Chinese
literature are also more detailed. For example,
NongSangYiShiZuoYao (farming, clothing,
dieting abstracts) written by Lu Mingshan in
the Yuan Dynasty (Lu, 1314 AD) describes how
peach trees should be planted as ‘桃树栽时提
根与地平，使侧根舒畅易活’, explaining that
when planting a tree, holding the tree and
making its lateral roots spread will improve
the survivability. In ZhiBenTiGang written in
the Qing Dynasty, Yang Shen (Yang, 1774 AD)
illustrates more precisely that successful
planting of peaches depends on a good under-
standing of root–shoot balance: ‘Planting: if a
tree has more shoots and less roots, shoots
need to be thinned; or if a tree has less shoots
and more roots, then roots need to be thinned’.
This documented how to maintain a balance
between shoots and roots. ‘The peach seed-
lings should be planted in spring time when
they are still in dormancy, otherwise, trees
should be planted in autumn when their
leaves fall off. Planting pits need to be deep
and wide for root spreading and develop-
ment, watering planted trees sufficiently and
returning the surface soil on to the roots
slowly will ensure soil to be settled on roots
well’. This is not surprising because large-
scale peach planting and production growth
occurred during this time. Evidently, ancient
43
Chinese peach growers gained rich experi-
ences from these 1000 years of cultivation and
they greatly improved their understanding of
peach tree physiology. They summarized the
relationship between successful planting of
peach trees and the growth balance of root
and shoots. During the same time, propaga-
tion and orchard management techniques
had been greatly developed, noted in
ShuiMiTaoPu (juicy peach register) as ‘桃树生
二，三年可接，多在春分前，秋分后，高树根
二，三尺许锯去，以快刀修光，使不沁水，又
向靠皮带膜处（韧皮部与形成层接合部） ，以
上切下一寸余，却以水蜜桃东南北枝两边削成
马耳状者，在口中含热插下，用纸封固，外涂
以泥，在加匿叶护之’, saying that the peach
can be grafted on 1- or 2-year-old seedlings,
usually before vernal equinox or after autum-
nal equinox. The seedling top is cut off at
60–90 cm from the ground using a sharp knife
and a cut between bark and wood about 3–5
cm is made. The scion-wood should be col-
lected from the east or south side of the can-
opy of the mature ‘juicy peach’. A mouse-ear
shaped piece is cut from buds of the scion-
wood and inserted into the cut of the seedling
(the scion piece can put in the mouth for
warmth and moisture), and then wrapped
with paper and soil slurry. The grafting tech-
niques for peach propagation were well
developed during this time. Also, orchard
management in southern China was well
understood during this period as described in
this book: ‘the peach growing in the south
usually suffers waterlogging that resulted in
root rot and orchard failures during rainy sea-
son. Deep ditches are needed for good drain-
age and fruit quality. The peach is drought
tolerant but sensitive to waterlogging, well-
drained orchards produce large and high
quality fruits’. The characteristics of peach
loving full sun and good drainage and
drought tolerance are well documented.
Good orchard practice for high yields and
high quality had been developed in almost all
major peach production regions in China dur-
ing this time.
Throughout the 4000-year history of
peach cultivation, Chinese peach growers
have made significant contributions to peach
domestication and peach industry develop-
ment through exploring wild germplasm,
44 H. Huang et al.
selection and cultivar development, develop-
ment of thorough understanding of tree
physiology and development of many impor-
tant cultural practices. China’s rich heritage
of peach cultivation is worthy of great appre-
ciation worldwide and of study by modern
pomologists.
2.3 Current Status of Chinese Peach
Production and Trends in China
Peach is one of the top five most important
fruit tree crops in China. Since China became
the largest peach producer in the world in
1994, peach planting and production have
increased steadily up to 1.4 million ha and
4.38 million t in 2003. Of this production
approximately 80% was in white-fleshed melt-
ing peach cultivars and 20% in other cultivars.
The total area planted in China has remained
static at around half the world total of 2.2 million
ha in recent years. The massive production of
Chinese apples and pears is having a marked
effect on world supplies and trade in pip fruit
and fruit juices. The increase in peach produc-
tion could likewise have a similar impact on
the international market.
Peach germplasm collection, repositories,
evaluation and utilization in China
The national peach germplasm survey con-
ducted during the 1950s has played an impor-
tant role in current germplasm collections
and repositories. Five species and 16 varieties
and forms were identified within subgenus
Amygdalus of genus Prunus. In addition, 800
traditional landraces and cultivars were doc-
umented (Wang et al., 1989; Guan and Wang,
1993). These efforts have resulted in the estab-
lishment of three national peach germplasm
repositories in Beijing, Zhengzhou and Nan-
jing. Local efforts in peach germplasm collec-
tion also occurred simultaneously in
Shanghai, Dalian and Shanxi. Altogether, more
than 1000 germplasm accessions have been
collected and maintained to safeguard against
genetic erosion or complete loss due to recent
rapid changes in the Chinese economy and
society (Wang et al., 1989). The efforts, together
with active programmes in screening and
evaluation, continued throughout the 1990s.
In addition, a number of foreign cultivars
were introduced to China, such as ‘Kanto 5’
and ‘Myoujou’ from Japan, and ‘NJN’ and
‘Babygold’ from the USA. In China, the peach
national repositories also serve as breeding
centres. Many new varieties have been devel-
oped from these three national and other local
repositories. The repositories also conduct
joint efforts in evaluation of the existing germ-
plasm, particularly some unique genotypes. For
example, the Fruit Germplasm Checklist (edited
by the Fruit Research Institute of the Chinese
Academy of Agricultural Science, 1993, 1998)
lists 648 good peach genotypes (landraces)
with detailed information on place of origin,
fruit maturity, fruit size, flesh characteristics,
free- or clingstone, soluble solids, soluble sug-
ars and acid, vitamin C, pollen viability, and
other unique characteristics and uses.
Screening and evaluation efforts at the
repositories have resulted in a marked
increase in the understanding of special geno-
types and the effective use of germplasm.
First, some commercial cultivars have been
selected directly from those superior geno-
types and used in peach production, such as
the northern variety ‘Feicheng Tao’ with big
fruit size and good transportability and storage
quality, as well as ‘Shenzhoushuimi’ (Sheng-
zhou melting honey), ‘Hanlumi’ (Cold dew
honey), ‘Huayumi’ (Flower pure honey) and
‘Baihua’ (White flower). These traditional
varieties are widely used in Chinese peach
production and have generated tremendous
benefit to both peach growers and the local
agricultural economy. Second, the germplasm
resources have been used as breeding materi-
als in conventional breeding programmes for
cultivar improvement. For example, ‘Shang-
haishuimi’ (Shanghai melting honey) has
played an essential role in newly released cul-
tivars both domestically and in foreign coun-
tries. In fact, many new cultivars are derived
from ‘Shanghaishuimi’, such as ‘Okubo’,
‘Hakuho’, ‘Yuhualu’ and ‘Zhaohui’. Mean-
while, new introduced nectarines and their
pollens have been used for hybridization with
traditional Chinese varieties, resulting in a num-
ber of new Chinese nectarine varieties. This has
greatly improved the nectarine varieties
 Cultivation and Trends in China
available and expanded nectarine production
in China. Third, progress has been made for
selection of pest-resistant genotypes that have
proved very useful in breeding programmes.
The results of germplasm evaluation have pro-
vided valuable resistance genetic materials that
have been directly or indirectly used in both
conventional breeding and new genetic engi-
neering programmes. This includes ‘Gansu
Tao-1’ (P. kansuensis Rehd.) and ‘Shouxing Tao-1’
(dwarf peach, P. persica var. densa Makino) with
proven root-knot nematode resistance.
Main peach cultivars, peach production
and growing regions in China
A great number of peach cultivars have been
developed in China with many different fruit
characteristics, adaptability and market val-
ues (Wang, 1990; Wang and Zhuang, 2001).
Consequently, the cultivars in Chinese peach
production vary in different geographic
regions and even in different provinces. The
China Fruit Plant Monograph – Peach Flora
(Wang and Zhuang, 2001) registers 495 culti-
vars with detailed information about cultivar
characteristics and production. This resulted
from an extensive evaluation of more than
1000 germplasm accessions conducted by the
two main national peach germplasm reposi-
tories at Beijing and Zhengzhou. Below, we
briefly list the main high-performance culti-
vars widely used in current Chinese peach
production by their ripening date (Wang,
1990; Liu et al., 1999; Guo et al., 2000; Zhu
et al., 2000; Wang and Zhuang, 2001). Although
many varieties have been selected and devel-
oped in China and are used in current peach
production, including melting peach, nectar-
ines, flat peach and ornamental peach, the
majority of the Chinese cultivars are white-
fleshed and melting type, accounting for more
than 80% of the total cultivars in current
Chinese peach production (Zhu et al., 2003).
Very early cultivars (less than 65 days from
full bloom to harvest)
● Common (white flesh, melting) peach (P.
persica Sieb et Zucc.): ‘Chunlei’ (Spring
45
bud), ‘Chunhua’ (Spring flower), ‘Zaoxi-
alu’ (Early morning glow dew), ‘Huiyu-
lu’ (Sunshine rain dew).
● Flat peach (P. persica f. compressa (Loud.)
Rehd.): ‘Zaoshoumi’ (Early big honey),
‘Zaolupantao’ (Early dew flat peach).
● Nectarine: ‘Shuguang’ (Dawn), ‘Huaguang’
(China glory).
Early cultivars (66–90 days from full
bloom to harvest)
● Common peach: ‘Baixianglu’ (White frag-
ment dew), ‘Yuhualu’ (Rain flower dew),
‘Yulu’ (Pure dew), ‘Yinhualu’ (Silver flow-
er dew), ‘Beinong-2’ (Beijing Agricultural
University – 2), ‘Zhaoxiang’ (Morning
glow), ‘Xianghui-1’ (Glow ray-1), ‘Beinong
Zaoyan’ (Beijing Agricultural University –
Early beauty), ‘Sunagawase’ (Japanese
cultivar), ‘Kurakato’ (Japanese cultivar).
● Yellow-fleshed peach: ‘Flavorlate’, ‘Fer-
tilia Morettini’.
● Flat peach: ‘Zaokuimitao’ (Early chief
flat peach), ‘Zaohuangpantao’ (Early yel-
low flat peach), ‘Wuyuexuanbiangang’
(May fresh flat peach), ‘Zaolupantao’
(Early dew flat peach).
● Nectarine: ‘Armking’, ‘Ruiguang-2’ (Lucky
ray-2), ‘Ruiguang-3’ (Lucky ray-3), ‘Yan-
guang’ (Beauty ray).
Mid-season cultivars (91–120 days
from full bloom to harvest)
● Common peach: ‘Zhaohui’ (Morning
sunshine), ‘Baifeng-2’ (White phoenix-2),
‘Zaoxiangyu’ (Early fragment jade), ‘Dat-
uanmilu’ (Big honey dew), ‘Japan-89’.
● Yellow-fleshed peach: ‘NJC88’, ‘Cullinan’,
‘Lianhuang’ (Even yellow), ‘Chengxiang’
(Orange fragment), ‘Myoujou’, ‘Redhaven’
(USA), ‘Babygold-5’, ‘Babygold-6’.
● Flat peach: ‘Sahuahongpantao’ (Splash
flower red flat), ‘Baimangpantao’ (White
awn flat), ‘Changshengpantao’ (Longev-
ity flat), ‘Fenghuapantao’ (Fenghua flat),
‘Chenpupantao’ (Chenpu flat), ‘Yulupan-
tao’ (Pure dew flat).
● Nectarine: ‘Zhongyoupantao’ (Mid nectar-
ine flat), ‘Zaohongzhu’ (Early red bead).
46 H. Huang et al.

Late-season cultivars (121–150 days ● Yellow-fleshed peach: ‘Bositao’ (Persian

from full bloom to harvest)
● Common peach: ‘Baihua’ (White flower),
‘Xinbaihua’ (New white flower), ‘Shenzhou’
(Shenzhou white honey), ‘Shenzhouhong-
mi’ (Shenzhou red honey), ‘Wanshuomi’
(Late big honey), ‘Feichengtao’ (Feicheng
peach), ‘Jingmi’ (Beijing honey), ‘Jingyu’
(Beijing jade), ‘Longhuashuimi’ (Longhua
melting honey), ‘Early red-2’.
● Yellow-fleshed peach: ‘Elberta’ (USA),
‘Fillips’, ‘Jincheng’ (Golden orange), ‘Jinxiu’
(Splendid), ‘Long 1-2-4’ (Dragon 1-2-4),
‘Xizhuang-1’ (West village – 1).
● Flat peach: ‘Huangroupantao’ (Yellow
flesh flat), ‘Jiaqingpantao’ (Jiaqing flat),
‘Lihepantao’ (freestone flat).
Very late season (more than 150 days
from full bloom to harvest)
● Common peach: ‘Dunhuadongtao’
(Dunhuang winter peach), ‘Qingzhou-
baipimitao’ (Qingzhou white skin honey
peach), ‘Yexiandongtao’ (Yexian winter
peach), ‘Zhonghuashoutao’ (China lon-
gevity peach).
peach).
● Nectarine: ‘Hongliguang’ (Red plum
shine).
Peach production
The total land area devoted to peaches and
nectarines in China has seen a more than
fourfold expansion during the period 1984–
2006 (141,351 to 652,700 ha, respectively)
(Fig. 2.3). More than 45% of the total land area
devoted to peaches in the world is in China.
Moreover, the total production (tonnes) has
followed a similar increase. By 1993, China
produced more peaches than any other coun-
try in the world (Fig. 2.4). Each year since
then China’s production has increased, so
that, by 2006, China produced 7.5 million t.
This represented 44% of the total supply. The
top five producing countries in 2006 were
China, Italy, Spain, the USA and Greece, pro-
ducing 44%, 10%, 7%, 5% and 5% of the world
total, respectively. For the last 30 years, aver-
age yield (kg/ha) has been less in China than
in the other top producing countries (Fig. 2.5).
However, since 1990, Chinese average yield
has increased each year such that in 2006 it

700,000
650,000
600,000
550,000
500,000
450,000
Total area (ha)

400,000 China
Greece
350,000 Italy
300,000 Spain
USA
250,000
200,000
150,000
100,000
50,000
0
19 1
19 3
65

19 7
19 9
71

19 3
75

19 7
19 9
19 1
19 3
19 5
19 7
89

19 1
19 3
19 5
19 7
99

20 1
20 3
05
6
6

6
6

7
7
8
8
8
8

9
9
9
9

0
0
19

19

19

19

19

20

Year

Fig. 2.3. Total peach and nectarine area (hectares) in the top five producing countries during the
period 1961–2006. (Source: http://faostat.fao.org, accessed August 2007.)
 Cultivation and Trends in China 47

8,000,000

7,000,000

6,000,000
Total production (t)

5,000,000
China
Greece
4,000,000 Italy
Spain
USA
3,000,000

2,000,000

1,000,000

0
19 1
19 3
19 5
19 7
69

19 1
73

19 5
19 7
19 9
19 1
19 3
19 5
19 7
19 9
19 1
19 3
19 5
19 7
20 9
20 1
20 3
05
6
6
6
6

7
7
7
8
8
8
8
8
9
9
9
9
9
0
0
19

19

19

Year

Fig. 2.4. Total peach and nectarine production (tonnes) of the top five producing countries during
the period 1961–2006. (Source: http://faostat.fao.org, accessed August 2007.)

24,000
22,000
20,000
18,000
Average yield (kg/ha)

16,000
14,000 China
Greece
12,000
Italy
10,000 Spain
USA
8,000
6,000
4,000
2,000
0
19 1
63

19 5
67

19 9
19 1
73

19 5
77

19 9
19 1
19 3
19 5
19 7
19 9
91

19 3
19 5
97

20 9
20 1
03
05
6

6
7

7
8
8
8
8
8

9
9

9
0
19

19

19

19

19

19

19

20

Year

Fig. 2.5. Average peach and nectarine yield (kilograms per hectare) in the top five producing
countries during the period 1961–2006. (Source: http://faostat.fao.org, accessed August 2007.)
48 H. Huang et al.
was just slightly less than the USA (11,506
versus 12,731 kg/ha, respectively).
The five top peach-producing provinces
are Shangdong, Hebei, Henan, Hubei and
Jiangsu (Wang, 2003). With current changes in
the agricultural industry in China, peach
acreage seems to be increasing in Sichuan and
Hunan provinces where citrus was overpro-
duced. The same trend is also occurring in
Yunnan, Guizhou, Fujian, Jiangxi and Guangxi
provinces where peach production is devel-
oping in higher elevation areas. In addition,
peach greenhouse production and high-
density cultivation have recently emerged in
limited areas of northern China.
Peach-growing regions
The natural range of wild peach is spread
widely over much of China. However, com-
mercial peach production is limited within
the latitude range 25–45°N (Fig. 2.2/Plate 26).
It is largely concentrated in northern, central
to eastern and north-western China. In general,
peach production in China can be divided
into seven regions based on regional climate
and ecological differences (Wang and Zhuang,
2001). Peach cultivars in these main production
regions are divided into regional groups that
are significantly different from one another.
1. North-west drought peach region in-
cludes Xinjiang and Ningxia autonomous re-
gion, Shaanxi and Gansu provinces.
2. Northern China plain region, the most
important traditional and current peach pro-
duction area in China. The northernmost
boundary region corresponds to the Qinling
Mountains and the Huai He River, including
Beijing, Tianjing, Hebei province, and southern
Liaoning, Shangdong, Shanxi, most of Henan,
Jiangsu and northern Anhui provinces.
3. Changjiang River humid region, having
a large area in central and eastern China, includ-
ing southern Jiangsu, Zhejiang, Shanghai,
southern Anhui, Jiangxi and Hubei and both
Chengdu and Hanzhong plain areas.
4. Yunnan–Guizhou high plateau cold re-
gion, a small-restricted area including Yunnan,
Guizhou and south-west Sichuan provinces.
5. Qinghai–Tibet plateau cold region, a lim-
ited area in the Tibet autonomous region, most
of Qinghai and western Sichuan provinces.
6. North-eastern China cold region, this is
the northernmost region of Chinese peach
production, further than 41°N latitude, includ-
ing Jilin and part of Heilongjiang provinces.
7. Southern China subtropical region, with
south limit at 23°N latitude, including a large
area to the south side of the Changjiang River
of Fujian, Jiangxi, southern Hunan, north
Guangdong, north Guangxi and Taiwan
provinces.
The first five regions are suitable peach pro-
duction regions, while the last two are mar-
ginal regions, as shown in Fig. 2.2/Plate 26.
Rapid development of greenhouse production
Protected cultivation of peach was successful
as early as 1995 in Shandong Agricultural
University (Gao et al., 2004) (Fig. 2.6/Plate
27). Greenhouse cultivation of peach has
greatly extended the peach marketing season
in China. Very early peach cultivars with low
chilling requirement and late- or very late-
season cultivars have been promoted for the
extreme early and late seasons, respectively,
with fivefold higher market price than regu-
lar orchard-produced peaches. A systematic
greenhouse cultivation technique has been
developed, including applying plant growth
regulation chemicals, reducing the size of foli-
age, summer pruning, girdling in the autumn,
artificial application of drought stress, root
pruning, and use of dwarfing rootstocks and
dwarfing or semi-dwarfing cultivars.
The microclimate in the greenhouse is
adjusted to maximize peach production by
controlling light, water, temperature, humid-
ity and CO2:O2 ratio. In addition to applying
extra artificial illumination, films with better
transparency have been used for covering
materials, and reflective films have been used
on the ground and in the air for regulating
light. Ripening stimulation is usually regu-
lated by controlling temperature during
bloom time between 5°C (night) and 22°C
(day) and during fruit ripening at 25–30°C,
which accelerates the ripening by 10 to 50
days. Large differences between night tem-
perature and day temperature can improve
the fruit quality. Delaying ripening is more
 Cultivation and Trends in China
Fig. 2.6. Modern greenhouse peach production.
often practised in the northern areas. Improved
insulation in greenhouses plus the frozen
ground of the semi-ground greenhouses in
northern China can be effective to hold a low
temperature (< 7°C) in the greenhouse for 50
days during spring, when temperature is ris-
ing in March to May. This can effectively
delay bud break in the spring and prolong
fruit ripening for 10–30 days. The humidity
and water content in the soil are usually reg-
ulated according to the growing season by
irrigation.
Other techniques are also used in the
greenhouse, which include breaking dor-
mancy with chemicals (hydrogen cyanamide)
and applying different types of fertilizers
based on the cultivars and growth season.
Typical spacing in the greenhouse is 1 m × 2
m. This high planting density usually requires
a special summer pruning method called PCR
pruning (postharvest canopy removal) for
controlling tree size. Trees before PCR are
shown in Fig. 2.7/Plate 28. The method
includes pruning off all current shoots as soon
as fruits are harvested, followed by several
summer tippings of new growing shoots for
enhancing flower bud formation. Trees after
49
PCR are shown in Fig. 2.8/Plate 29. Foliar dis-
ease control could be a problem due to high
humidity in the greenhouse production sys-
tem, but greenhouse management such as
ventilation and irrigation controls usually
regulates humidity.
The greenhouse peach production sys-
tem usually remains productive for about 10
years. Intercropping systems for greenhouse
peach cultivation are also being developed
for maximizing the output of greenhouse
productivity, such as strawberry intercropped
between rows in a peach greenhouse. Cur-
rently, greenhouse peach production has
reached about 14,000 ha (Li et al., 1995; Wang
et al., 1995; Zhu and Wang 1997; Wang and
Niu, 1998; Zhang and Yu, 2002).
Peach postharvesting, processing
and marketing in China
Nearly all fresh peaches produced in China are
marketed within the country. Marketing is as
for all other fresh fruits and vegetables: farms
or farming cooperatives send peaches to distri-
bution centres (large trading centres organized
50 H. Huang et al.

Fig. 2.7. Modern greenhouse peach production before postharvest canopy removal.

Fig. 2.8. Modern greenhouse peach production after postharvest canopy removal.
 Cultivation and Trends in China
by local governments), and from there the
fruit are sent to fruit stores or wholesale cen-
tres in cities. Most peach production for local
fresh markets requires little packing. For dis-
tant provincial markets, packing and trans-
portation are necessary. Fruits are usually
harvested before completely ripe (70–80%
ripe) and packed in cardboard boxes (50
cm × 40 cm × 20 cm; about 10 kg of fruit per
box) for storage and shipping. For prolonged
supplies to the markets several storage tech-
niques are usually used, mostly by cold stor-
age, although controlled-atmosphere storage
and low-pressure storage are used to a certain
extent (Du et al., 2000). Cold storage usually
applies 0–1°C temperature and 85–90% rela-
tive humidity, while controlled-atmosphere
storage is created with 0–1°C temperature,
5% CO2 and 1–3% O2. The processed peach
products in China are mostly canned, dry
fruit and sliced dry products. Recently a new
series of processed products has been devel-
oped with advances of modern industrial
technology, including peach juice, peach tea
drinks, beer, fruit jelly and peach candies
(Zheng, 1995; Zheng et al., 2001).
Approximately 80% of Chinese peach
production is for the domestic fresh market.
Small quantities of fresh peach, mostly white-
fleshed melting peach, have been exported to
South-east Asian countries since the mid-
1950s. There is a trend for increased fresh
peach exports to neighbouring countries in
recent years. Substantial amounts of processed
products are also exported to European and
American markets (Wang and Zhuang, 2001).
Problems faced by the Chinese peach industry
Although peach production has become an
important rapidly developing industry, the
lack of overall industry organization and
long-term strategic planning presents barri-
ers for future profitability (Jiang, 2000; Zhu
et al., 2003). Some of the problems that need
to be overcome include the following.
Cultivars in production
For a healthy industry, China needs a balanced
proportion of different ripening peaches for a
51
prolonged market supply. Particularly, the
overproduced early-season peaches have
greatly hampered industry development for
market supply. Also, an unbalanced ratio of
the white-fleshed melting peach does not meet
consumer demand for a diversity of peach
fruit types (Chen and Liu, 1999; Wang, 2000).
Orchard management
Poor orchard management is responsible for
low yields per hectare, and poor fruit quality.
A large percentage of peach orchards might
be currently operated in extensive (low-care)
cultivation. For example, high-density
orchards without appropriate canopy man-
agement have caused production to decline
rapidly and shortened peach tree life. Driven
by short-term profit return, large-dose appli-
cation of chemical fertilizers without or with
less organic manures has overridden recom-
mended management practices of balanced
fertilization based on orchard age, yields, soil
types and climate difference. Under these
orchard practices most good cultivars cannot
reach their highest yields and quality for mar-
keting (Chen, 2002; Wang, 2000).
Postharvest issues
Lack of postharvesting facilities and expenses
associated with handling fruit have ham-
pered quality control (Du et al., 2000; Zhao
and Chen, 2004). Protocols for peach clean-
ing, sorting and packing have not been well
developed and are necessary for a well-regu-
lated marketing system. Also, a lack of stor-
age facilities and poor coordination between
storage and transportation systems have
caused tremendous losses during storage and
transportation in some high-yielding years
and reduced profit returns to peach farmers
(Zhao and Chen, 2004). A well-organized pro-
vincial or regional marketing system is
urgently needed.
Small family farms
Most Chinese peach production is on family-
based small farms. This small-scale and locally
operated production and marketing is proba-
bly responsible for the poor coordination of
52 H. Huang et al.
orchard production, postharvest handling
and storage and shipping (Zhu et al., 2003).
Consequently, it is difficult to standardize the
production protocols and regulate quality
controls. The current system must change, or
the Chinese peach industry will remain less
competitive within the world markets.
Applied research and extension education
Variety trials and an extension education sys-
tem are not well established. Consequently,
new cultivar propagation increases and sales
are not regulated and some cultivars are over-
planted before they are adequately tested
(Wang, 2000).
Current trends of Chinese peach production
Germplasm collection and evaluation
Germplasm collection and evaluation needs
to be a more scientifically based, long-term
commitment for further industry develop-
ment. It has been recognized among Chinese
botanists and peach researchers that the lack
of investigations in peach native centres and
the lack of genetic diversity assessment across
China’s diverse geographic and climatic
regions are hampering a more scientifically
based coverage of the existing gene pool of
native peach germplasm in China (Zhao et al.,
2000). More research efforts will need to be
devoted towards documentation and inven-
tory assessment of wild germplasm and
genetic differentiation across native centres in
different Chinese geographic regions. New
molecular fingerprinting techniques will need
to be widely used to establish the authenticity
of Chinese traditional cultivars and landraces
and to enhance management in Chinese
peach repositories. Furthermore, molecular
investigations into the taxonomic uncertain-
ties of subspecies, varieties and forms within
the genus should be considered. Evaluating
and discovering new genes regulating novel
and important traits, such as compact tree
forms, weeping habits, double petals and dif-
ferent flower colours, sweet kernels and pest
resistance, etc., will be incorporated into
peach breeding programmes for a variety of
purposes to enhance the applications of unique
Chinese peach resources (Wang and Zhuang,
2001). These new strategic goals will contribute
greatly to new peach cultivar improvement
and benefit the world peach industry.
Cultivars/market
Chinese peach production will be readjusted
according to market needs and new cultivar
releases will be accelerated in the future.
Based on peach production zones in
China, the peach cultivars used in the different
peach-growing regions will be more market-
oriented and readjusted to market changes.
For a prolonged market supply, the peach
cultivars in different ripening dates may be
structured as the ratio of 5:35:30:25:5 for very
early cultivars (<65 days after full bloom, this
category includes protected cultivation of
peaches and nectarines), early-season culti-
vars (66–90 days after full bloom), mid-season
cultivars (91–120 days after full bloom), late-
season cultivars (121–150 days after full
bloom) and very late-season cultivars (>151
days after full bloom), respectively (Zhu et al.,
2003). The peach market tends to have greater
demand for early cultivars with larger fruit
size and good flavour; for mid–late cultivars
with larger size, good appearance and stor-
age quality; and for very late cultivars with
medium size, no fruit cracking, good appear-
ance and stress tolerance (Jiang, 2000).
New peach cultivar development will
focus on more novel characteristics, such as
high content of carotene and flesh browning
tolerance for yellow-fleshed cultivars; new
nectarines with wide adaptability, larger size,
high fruit quality and light red or golden-
coloured skin; and new flat peach cultivars
with larger size, good appearance and rich
flavour and aroma, and less fruit cracking.
New novel cultivars will be promoted to
develop niche markets. New breeding pro-
grammes for flat peach cultivars will have a
greater emphasis on dwarf or semi-dwarf and
compact tree forms for high-density plantings
and ornamental flower types. Low-chilling
genetic resources of both Chinese and foreign
origin will be used to develop <500 chill-hour
peach cultivars for growing in southern China
(Wang et al., 2000). With targeted exploration
 Cultivation and Trends in China
of wild peach genetic germplasm and new
biotech applications in peach breeding, culti-
vars with stress tolerance and pest resistance
may be developed and used in the Chinese
peach industry. Cultivar replacement time
will be reduced from the current 20–30 years
to a 10–15-year period with 80% newly
released cultivars in the production. The ratio
of melting peach versus non-melting peach
will be changed from the current 9:1 to 7:3 or
6:4 in the future (Jiang, 2000).
Production systems
Production systems are moving towards
large-scale, intensive and industrialized orga-
nizations or cooperatives. Current production
systems based on family operations should
be gradually reformed to a more industrial-
ized system of professional farm cooperatives
comprising farms with large centralized pack-
ing and shipping or processing facilities (Yi,
2003). This will help the Chinese peach pro-
duction and industrialization system to be
upgraded to a more competitive, world stan-
dard, highly commercialized and efficient
system. The new development of intensive
Fig. 2.9. Conventional peach orchard.
53
production and marketing organizations will
enhance China’s efforts to fully participate in
the world market as a new member of the
World Trade Organization.
Orchard system management
The peach orchard system will probably be
changed from currently intensive tillage to
less tillage or free tillage or even biodegrad-
ing films for weed control and for improving
fruit quality and yield. Fertilization in peach
orchards should reconsidered using tradi-
tional organic fertilizers for high-quality fruit.
Irrigation systems will need to be more water-
efficient with wide application of dripping or
microsprinkler irrigations (Shi et al., 1990).
The traditional low-density orchards
with large tree size (Fig. 2.9/Plate 30) have
been changing to current higher-density orchard
systems (Figs 2.10 and 2.11/Plates 31 and 32).
New rootstocks for compact or semi-dwarf
tree forms have played an important role in
high-density orchard development. Recently,
in addition to using semi-dwarf peach cultivars,
dense-pubescent cherry (Prunus polytricha
Koehne) and Chinese dwarf cherry (Prunus
54 H. Huang et al.

Fig. 2.10. Modern high-density peach orchard.

Fig. 2.11. Modern high-density peach orchard (dormant).

humilis Bunge) are being widely used as root-
stocks to reduce tree size (Wang and Zhuang,
2001). Rootstock breeding for high-density
orchards has resulted in specific selections
such as ‘Ailihong’ (Dwarf bright red, P. persica
var. densa) that has short internodes and is
suitable for high-density plantings (Gao et al.,
1992). In application of available dwarf root-
stocks and other dwarfing resources from dif-
ferent Prunus species, further exploitation and
development of rootstocks will be more focused
on new rootstocks with dwarf characteristics
 Cultivation and Trends in China
combined with the potential of improving fruit
quality and yields in high-density orchards.
Planting space will be more optimized towards
the specific requirements of different cultivars
and rootstock types to improve sunlight pene-
tration and fruit quality (Zhu et al., 2003).
Peach pruning systems tend to be more
simplified and less labour-intensive. A vari-
ety of different pruning methods are used in
Chinese peach orchards, such as vase figura-
tive, modified vase, multi-scaffold, natural open
centre and modified central leader forms (Wang
et al., 1989). Recent development of simplified
training and management systems has been
aimed towards the demand of high-density
orchards or less labour-intensive management
systems (Wang et al., 1999). A single leader form
has been developed for high-density orchards,
while the ‘V’ shape and two-scaffold ‘Y’ forms
are widely used in newly planted orchards
(Liu, 1997; Wu et al., 1998; Xu et al., 1998). The
tree training process has also been simplified
to increase earlier fruit production with the aid
of summer pruning applications.
Greenhouse cultivation
Greenhouse cultivation has been showing
great market potential in recent years. Further
efforts are needed towards the selection and
evaluation of suitable candidates of either
early cultivars with low chilling requirement
or very late cultivars with high quality. This
development should significantly prolong
the market window of fresh fruit supplies
(Wang et al., 2002). Greenhouse cultivation
technology will be improved into a central
computerized system for managing fertiliz-
ers, water, light, temperature and gas (CO2) in
the greenhouse (Wang and Zong, 1996).
Biotechnology applications
In vitro propagation has been successfully
developed for peach, which could be used in
the production of virus-free rootstock and
increase propagation efficiency (Wu et al., 2003).
Marker-assisted selection can dramatically
reduce the time, labour and space cost during
the selection processes for peach breeding
programmes. High-throughput genotyping
using microsatellite markers can even be easily
55
used in regular biological laboratories. With
more research efforts devoted to the genetic
transformation of Prunus species, geneti-
cally modified peach and nectarines might
be available for peach production in the
future. This may include environmentally
friendly, genetically engineered cultivars
with a wide spectrum of pest resistance,
frost tolerance, compact tree forms, semi-
dwarf or dwarf varieties, as well as new
cultivars with novel genes regulating ripen-
ing and postharvest storage and shipping
characteristics (Liu et al., 1991; Jiang et al.,
1993; Wei and Timon, 1994; Ma and Li, 1999;
Zhang et al., 2001).
New pest control approaches
Many common peach diseases and insects
occur in China, including brown rot (Monilinia
fructicola (Wint.) Rehm), leaf curl (Taphrina
deformans (Berk) Tul.), anthracnose (Gloeospo-
rium laeticolor Berk), bacterial leaf spot (Xan-
thomonas pruni (Smith) Dowson), canker (Valsa
leucostoma (Pers.) Fr.) and gummosis, as well as
insects including the pyralid moth (Dichocro-
cis punctiferalis Guen.), aphids (Myzus persicae
Sulzer), leaf moth (Thosea sinensis Walker),
trunk borer (Aromia bungii Fald.) and root-
knot nematodes (Meloidogyne spp.) (Wang
et al., 1989). Different pest control approaches
such as chemical, agricultural, physical and
bio-controls are being utilized for Chinese
peach production. Future improvement in
peach pest controls should be geared
towards the bio-control approaches to meet
the growing market demand for organic
fruits. Other more sophisticated integrated
pest management systems will also be devel-
oped for achieving fruits free of pesticide
residues (Chen, 2002; Ma et al., 2002; Ma and
Jia, 2003).
Ornamental peach
Peach has become one of the most important
ornamental trees for early spring bloom in
China. Due to the rich culture and history of
peach cultivation, more exotic and ornamental
peach varieties have been used for recreation
tourism, shade trees and cut flowers (note
ornamental ‘chrysanthemum’ peach, Fig. 2.12/
56 H. Huang et al.

Fig. 2.12. Ornamental ‘chrysanthemum’ peach.

Fig. 2.13. Ornamental ‘longevity’ peach. (From Wang and Zhuang, 2001.)
 Cultivation and Trends in China 57

Plate 33; and ‘longevity’ peach, Fig. 2.13/
Plate 34). Many peach flower gardens have
been established all over China, including Peach
Origins in Hunan, Broken Bridge near West
Lake, Peach Peak in the Yellow Mountains,
Dragon Spring Recreation Site in Chengdu,
Sichun, Xiangshan in Beijing, Leting in Hebei,
etc. In Hong Kong, Macau, Guangzhou and
the Zhujiang triangle region, the peach flower
is called ‘the Christmas tree in China’, since
the peach flower is one of the most important
cut flowers for the Chinese Spring Festival in
south-eastern provinces. Chinese people in
southern Asia, especially Singapore, are very
fond of peach flowers. Recently, peach flow-
ers have become more popular in big cities,
including Beijing, Shanghai and Dalian, for
cut flowers during the Chinese Spring Festi-
val holiday (Jiang, 2000).
germplasm, selection and cultivar develop-
ment, development of a thorough under-
standing of tree physiology and development
of many important cultural practices. Tre-
mendously diverse genetic resources of wild
peaches are still widely grown in large areas
of China and will provide valuable breeding
materials for cultivar improvement and for
supporting the sustainable peach industry.
China is now the largest peach producer in
the world, with a total planted area of 2.2 mil-
lion ha and an annual production of about 6
million t, which accounts for half of the total
world acreage and nearly half of the total
world production. Although many problems
remain, China’s peach industry will continue
to improve and facilitate many new develop-
ments for the future.

2.4 Summary/Conclusion
China has played a crucial role in peach
domestication for world peach cultivation.
Throughout the 4000-year history of peach
cultivation, Chinese peach growers have
made a rich heritage for the world of
peach cultivation through exploring wild
Acknowledgements
We would like to thank Chen Xuzhong,
research associate of Wuhan Botanical Gar-
den, for translating the Chinese references
and Qing Lin, Professor of Beijing Agricul-
tural College, for providing photographs.
This work was supported by CAS key project
KSCX2-SW-104.

References

Anon. (11th–6th century BC) ShiJing (The Book of Songs). This book compiles the works from the 11th century
BC to the 6th century BC, 305 volumes in total; the poems being divided into three categories: Feng (Bal-
lads), Ya (Festal Odes) and Song (Sacrificial Songs).
Anon. (2nd century BC) ErYa. This is one of the earliest reference books serving as a dictionary for names and
terms, and was probably written (author unknown) between the Qin Dynasty and the Han Dynasty; 20
chapters in total, but only 19 chapters remain. The descriptions about plant classification in the ErYa
were reprinted in 1962 in the Collected Papers of Science History, issue 4.
Ban, G. (39–92 AD) HanShu. Ban Gu was a historian of the Western Han Dynasty, HanShu is the historical
annals; it describes the development of philosophy, economy and literature for a period of 200 years in
the Western Han Dynasty.
Chen, P. and Liu, J.-X. (1999) Present situation of Chinese peach new variety selection. Hebei Fruits 3,
8–9.
Chen, W.-H. (1994) Illustrated Handbook of Ancient Relics of Chinese Agriculture. Jiangxi Technology Press,
Nanchang, China, pp. 99–100.
Chen, W.-Y. (2002) Technical standards for organic cultivation of ‘Feicheng’ peach. Hebei Fruits 4, 20–21.
Chu, H. (1813 AD) ShuiMiTaoPu (melting honey peach register). ShuiMiTaoPu is a monograph on the cultiva-
tion of honey peach in Shanghai region, including history, cultivar description, propagation method,
culture technique and orchard management. Reprinted in Guangxu 26 (1900 AD) in the Journal of Agron-
omy October, No. 124, 1–4.
58 H. Huang et al.

Cui, S. (2nd century AD) SiMingYueLing – Vol. 69. Cui Shi was an agronomist of the Eastern Han Dynasty and
SiMingYueLing is the earliest farming calendar in China; it describes and records all activities of army,
farming, industry, business and the way of living.
Du, J.-C., Ma, K. and Wang, X.-N. (2000) Research advances in storage and handling technology for peach.
Northern Fruits 6, 1–3.
Duan, S.-L., Zong, X.-P., Liu, X.-Y., Zuo, Y.-Z. and Duan, Y.-C. (1983) Preliminary report on germplasm re-
sources of fruit trees in Xizang. Acta Horticulturea Sinica 4, 217–223.
Fruit Research Institute, China Academy of Agricultural Science (ed.) (1993) Fruit Germplasm Checklist. China
Agricultural Press, Beijing.
Fruit Research Institute, China Academy of Agricultural Science (ed.) (1998) Fruit Germplasm Checklist. China
Agricultural Press, Beijing.
Fu, X. (2nd century AD) Peach Ode.
Gao, H., Wang, S. and Wang, J. (2004) Fruit protected cultivation in China. Acta Horticulturae 633,
59–66.
Gao, M.-X., Liu, P.-Z. and Ren, J.-X. (1992) Relationship between sunlight distribution and yield and fruit
quality of different tree training systems. China Fruits 4, 10–13.
Ge, H. (1st century AD) XiJingZaJi. Ge Hong was a Taoistic theorist and doctor; the book XiJingZaJi is an ad-
versaria about Taoism and agricultural activities in XiJin region (now Xi’an city, Shanxi province).
Guan, H.-R. and Wang, X.-S. (1993) Present situation of peach resource and development trends in China.
Land and Natural Resources Research 2, 62–65.
Guan, Z. (5th century BC) GuanZi-DiYuan. Translated and interpreted by Xia, W.-Y. (1958), Zhonghua Press,
Beijing, pp. 59–60.
Guo, H., Zhou, J.-T., Zhao, M.-Z., Yu, M.-L. and Ma, J.-J. (2000) Brief description of good peach cultivars.
Journal of Fruit Science 17(Suppl.), 72–74.
Guo, Y.-G. (4th century AD) GuangZhi. This is a chorography about geography, climate, local culture and ag-
riculture.
Han, F.-Z. (280–233 BC) HanFeiZi. Han was a philosopher in the late Warring States Period; the book after his
name compiles his works, 33 volumes in total.
Jia, S.-X. (533–544 AD) QiMingYaoShu. QiMingYaoShu is an encyclopaedia for living; 92 chapters in total,
divided into ten volumes. The book records crops, fruit tree cultivation and breeding techniques, etc.
Reprinted in 1963 by Agricultural Press, Beijing, pp. 49–52.
Jiang, H.-J., Pan, J.-S. and Meng, X.-F. (1993) The study of meristem-tip culture in vitro for peach. Acta Agri-
culturae Universitatis Pekinensis 19, 49–52.
Jiang, Q. (2000) The status and development trend of China’s peach production. Beijing Agricultural Science
18, 35–38.
Li, S.-Z. (1578 AD) BenCaoGangMu. Li Shizhen was a traditional Chinese medicine doctor in the Ming Dy-
nasty; BenCaoGangMu is one of the most important Chinese pharmacopoeias. Reprinted in 1957 by
People’s Medicine Publishing House, Beijing, 1256 pp.
Li, Y.-W., Wang, X.-H. and Zuo, Q.-Y. (1995) Greenhouse peach cultivation. Shanxi Fruits 2, 14–15.
Liu, H.-C. (1997) Two pruning methods for Y form peach tree training in greenhouse. Beijing Agriculturea 8, 18.
Liu, M., Wu, Z., Ding, G.-Q., Wang, F.-T. and Liu, Y.-H. (1999) A new very late mature peach variety – Zhon-
ghuashoutao. China Fruits 3, 7–8.
Liu, Y.-S., Hu, N.-Y. and Lu, G.-M. (1991) Research on ovule culture in vitro for early-mature peach. Acta
Botanica Boreali-Occidentalia Sinica 19, 37–43.
Lü, B.-W. (300 BC) LüShiChunQiu (Lü’s spring and autumn annals).
Lu, H. (3rd century AD) YeZhongJi. YeZhongJi is an adversaria at an ancient place, Yezhong in the Western Jin
Dynasty (currently Lin-Zhang region, Hebei province).
Lu, M.-S. (1314 AD) NongSangYiShiZuoYao (farming, clothing, dieting abstracts). Reprinted in 1962 by China
Agricultural Press, Beijing, pp. 26–27.
Ma, F.-W. and Li, J.-R. (1999) Callus formation from cultured protoplasts of Prunus davidiana. Acta Agricultu-
rae Boreali-Occidentalis Sinica 8, 73–76.
Ma, Z.-J., Huang, X.-H., Yong, W., Du, Y.-Q. and Wei, W.-D. (2002) Field trial of Kurarinone natural pesticide
efficiency against red mite and Carposina niponensis Walsingham. Journal of Ningxia Agricultural
College 23, 75–76.
Ma, Z.-S. and Jia, Y.-Y. (2003) New techniques for disease and pest control in organic peach production.
Hebei Fruits 4, 24–25.
 Cultivation and Trends in China 59

Qu, Z.-Z and Sun, Y.-W. (1990) Systematic Pomology. China Agriculture Press, Beijing, p. 69.
Shi, Y.-Z., Liu, Y.-R. and Liang, Y.-W. (1990) Study on drip-irrigation in peach orchard. Journal of Fruit Science
7, 105–108.
Sima, Q. (1st century BC) ShiJi (a history annals, 100 BC). Sima was a historian, literateur and thinker in the
Western Han Dynasty; ShiJi is the first general history in China.
Wang, B. (1st century AD) TongYue.
Wang, J.-Y. (2003) China’s fruit production and development suggestions. China Fruits 6, 42–44.
Wang, L.-R. (2000) Reviewing present situation of US peach and nectarine production and development of
Chinese peach industry. China Fruits 3, 44–46.
Wang, L.-R., Zhu, G.-R. and Zuo, Q.-Y. (1995) Choosing peach variety for protected cultivation. China Fruits
4, 34–35.
Wang, L.-R., Zhu, G.-R. and Zuo, Q.-Y. (2000) Advance in breeding of low chilling peaches and nectarines.
Journal of Fruit Science 17, 57–62.
Wang, X.-J. (1621 AD) QunFangPu (Florilegium). QunFangPu is a farming book with literary quotations, art
and culture; it illustrates each plant’s morphological traits and cultivation method (including fruit trees).
Reprinted in 1957 by Zhonghua Press, Beijing.
Wang, Y.-P., Yang, F.-K., Luo, C.-X., Hu, X. and Mu, C.-G. (2002) Technique of delaying maturity for green-
house peach cultivation in cold zone. Shanxi Fruits 2, 16–18.
Wang, Z. (1313 AD) WangZhen’s Farming Book. This is a monograph about crop cultivations (including fruit
and vegetable). Reprinted in 1981 by China Agricultural Press, Beijing, pp. 127–128.
Wang, Z.-H. (1990) Peach Cultivars. China Agricultural Press, Beijing.
Wang, Z.-H. and Zhuang, E.-J. (2001) China Fruit Monograph – Peach Flora. China Forestry Press, Beijing,
pp. 42–51.
Wang, Z.-H., Lu, Z.-Y., Hu, Z.-L. and Zhang, K.-B. (1989) Peach. In: China Agricultural Science and Technol-
ogy Achievements in Forty Years. China Agricultural Press, Beijing, pp. 179–184.
Wang, Z.-Q. and Niu, L. (1998) Present situation and research priorities for China’s protected fruit production.
Economical Forest Research 16, 62–65.
Wang, Z.-Q. and Zong, X.-P. (1996) Research and development in Chinese peach breeding and cultivation
technology for 21 century. Economical Forest Research 14(Suppl.), 20–23.
Wang, Z.-Q., Liu, S.-W., Niu, L., Fan, W. and Liu, H.-C. (1999) Study on the training and pruning system for
nectarines in protected cultivation. Journal of Fruit Science 16, 185–191.
Wei, S. and Timon, B. (1994) Improvement in propagating virus-free peach (P. persica) plants in vitro. Jiangsu
Journal of Agricultural Science 10, 1–4.
Wu, Y.-J., Zhang, S.-L., Zhang, L.-L., Shen, J.-Q. and Wu, D.-J. (2003) Shoot regeneration from immature
cotyledons of peach. Journal of Zhejiang University (Series – Agricultural & Life Sciences) 29(1),
93–96.
Wu, Y.-L., Xu, Z.-N. and Zhuang, E.-J. (1998) Technique for central leader pruning system in high density
peach cultivation. China Fruits 3, 22–23.
Xu, Z.-N., Wu, Y.-L. and Zhuang, E.-J. (1998) Field experiments with central leader training and pruning sys-
tem for the high-density peach orchard. Journal of Fruit Science 15, 317–321.
Yang, S. (1774 AD) ZhiBenTiGang. ZhiBenTiGang is an agricultural textbook for field crop, fruit tree and mul-
berry. Reprinted in 1957 by Zhonghua Press, Beijing, pp. 67–69.
Yi, F.-H. (2003) Present situation, prospects and strategies for Chinese fruit industry development. Abstracts of
Chinese Horticulture 6, 4–5.
Zhang, Y.-Q. and Yu, D.-Q. (2002) Present situation of development and countermeasure for protected horti-
culture. World Agriculture 11, 41–43.
Zhang, Y.-Q., Chen, D.-M., Jin, Y.-F. and Zhang, S.-L. (2001) Regeneration of peach plantlets from callus de-
rived from explant. Acta Horticulturae Sinica 28, 342–344.
Zhao, G.-X. and Chen, X. (2004) Fruit storage technology. Northern Fruits 1, 69–71.
Zhao, M.-J., Guo, H., Yu, M.-L., Zhou, J.-T. and Ma, R.-J. (2000) Research advance in peach genetic resource.
Journal of Fruit Science 17(Suppl.), 46–49.
Zheng, F.-G. (1995) A series of processed products from ‘honey’ peach. Food Science 16, 66–67.
Zheng, H.-Y., Yin, Y.-N., Yu, C.-G. and Xu, J.-H. (2001) Processing technique for series of nectarine products.
Jiangsu Food and Fermentation 4, 35–37.
Zhou, S.-H. (1018 AD) LuoYangHuaMuJi; hand-written copy in the Qing Dynasty. This is a monograph describ-
ing flower and fruit tree cultivation in present-day Luoyang region, Henan province.
60 H. Huang et al.

Zhu, G.-R. and Wang, L.-R. (1997) Protected peach cultivation and key techniques. Deciduous Fruits 3,
42–43.
Zhu, G.-R., Wang, L.-R., Zuo, Q.-Y. and Fang, W.-C. (2000) Good peach cultivar resources. China Seed Indus-
try 2, 16–19.
Zhu, G.-R., Wang, L.-R. and Fang, W.-C. (2003) The status of peach production in China and development
strategies. Deciduous Fruits 4, 14–16.
 3 Classical Genetics and Breeding

Rene Monet1 and Daniele Bassi2

1National Institute for Agronomical Research (INRA), Bordeaux Research Centre,
France (retired)
2University of Milan, Milan, Italy

3.1 Classical Peach Genetics 61

Qualitative (single) characters and their inheritance 61
Quantitative characters 66
Ploidy 66
3.2 Peach Breeding 68
First steps 68
Objectives 69
Floral biology and selection cycles 72
Methodology 72
Technique 74

3.1 Classical Peach Genetics
With the rediscovery of Mendel’s laws at the
beginning of the 20th century, breeders, who
had worked empirically before, became
interested in character inheritance and tried
to verify if these laws had general applica-
tion. First results came from the observation
of progenies obtained for species improve-
ment rather than from specific genetic
studies.
For fruit trees, where mutated individu-
als are scarce and reproductive cycles slow,
genetic studies were neglected for a long time.
Nevertheless, peach was an exception. Breed-
ers and researchers have collected many data
on character inheritance, so this species can
be regarded as a model for genetic knowledge
advancement. Here, we review results that
are useful for a breeder as well as those in
relation to single (Mendelian) or quantitative
characters.
Qualitative (single) characters and their
inheritance
The most important single traits found in peach
are described and discussed in Chapter 1.
They are grouped in Table 3.1 with their mode
of inheritance and citation of the first author(s)
who studied them.
The small number of available mutants,
the long time needed to create F2 generations
and the land occupied by trees did not permit
the location of the genes which determine the
characters described above on the eight linkage
groups expected in peach. Only molecular

© CAB International 2008. The Peach: Botany, Production and Uses

(eds D.R. Layne and D. Bassi) 61
 62
Table 3.1. Qualitative, Mendelian traits in peach and association to specific genomic linkage groups. (Updated from Sansavini et al., 2006.)

Linkage
Phenotype and symbol Genotype groupa Note Reference

Tree
Broomy (columnar or pillar) (Br) br/br 2 Incomplete dominance; phenotype is upright Scorza et al. (1989, 2002); Yamazaki et al.
when Br is heterozygous with the alleles for (1987); Chaparro et al. (1994)
the standard, dwarf, compact or weeping
growth habits
Upright (Up) Br/br See: Columnar Scorza et al. (1989)

R. Monet and D. Bassi

br/pl
br/dw
br/Ct
Arching (Ar) Brbr/plpl Upright weeping; similar to the Up, but with a Werner and Chaparro (2005)
distinct curvature of the 1-year-old shoots; from
F2 or backcross progenies of columnar (Br) ×
weeping (Pl) crosses; Br is epistatic to Pl
Bushy (Bu) bu1/bu1 Bu1 and Bu2 are independent Lammerts (1945)
bu2/bu2
Compact (Ct) Ct/– Mehlenbacher and Scorza (1986)
Dwarf (Dw) dw/dw 6 Short internode (<10 mm) Lammerts (1945)
dw2/dw2 Very dwarf Hansche (1988)
dw3/dw3 Extremely dwarf, thin stem Chaparro et al. (1994)
Semi-dwarf (N) n/n Incomplete dominance Monet and Salesses (1975)
Weeping (Pl) pl/pl Incomplete dominance, featuring open, Monet et al. (1988)
intermediate canopy when heterozygous
(Pl/pl) (Bassi and Rizzo, 2000). Pl from pleureur
(to weep, in French)
Standard Dw/– This growth habit results from the allelic status
Br/Br of any of these known genotypes
Pl/Pl
ct/ct
Graft incompatibility with
Damas 1869 plum (I)
Corky triangle (T)
Evergreen (Evg)
Anthocyanin deficiency (An)
Anthocyaninless (W)
I1/– Two dominant genes; incompatibility found only Salesses and Al-Kai (1985)
I2/– in some nectarines
t/t Epidermic suberification at bud base in 1-year- Monet and Bastard (1982)
old shoot
evg/evg 1 Also called evergrowing (Bielenberg et al., 2004): Lammerts (1945); Rodriguez et al. (1994)
terminal buds do not go dormant
an/an Pale pink flowers Monet (1967)
w/w White flowers; no red anywhere Lammerts (1945)
wv/wv Wv is unstable and produces variegated Lammerts (1945); Chaparro et al. (1995)
flowers (peppermint)

Resistance to root-knot nematode, Mj1/– 2 Mj1 and Mj2 are independent Lownsberry and Thomson (1959);
Meloidogyne javanica (Mj) Sharpe et al. (1970); Lu et al. (2000)

Mj2/–

Genetics and Breeding

Resistance to root-knot nematode, Mi/– 22 Weinberger et al. (1943); Lu et al. (2000)
Meloidogyne incognita (Mi)
Resistant to both species Mij/
Green aphid resistance (Rm) Rm1/– Monet and Massonié (1994)

Leaf
Foliar glands (E) 7 Incomplete dominance Connors (1921)
Reniform E/E
Globose E/e
Eglandular e/e High susceptibility to powdery mildew; serrate leaf
margin
Redleaf (Gr) Gr/– 6–8 Red incompletely dominant over green in leaves Blake (1937)
and fruit skin ground colour
Albinism (C) c/c Plant does not survive Bailey and French (1932)
Wavy-leaf (Wa) wa/wa Scott and Cullinan (1942)
Crinkle leaf (CL) cl/cl Associated with very oblate fruit shape Ledbetter (1996)
Willow-leaf (Wa2) wa2/wa2 Chaparro et al. (1994)

(Continued)

63
 64
Table 3.1. continued

Linkage
Phenotype and symbol Genotype groupa Note Reference

Flower
Non-showy (Sh) Sh/– 1 Connors (1920); Bailey and French (1942);
Lammerts (1945)
Large size (L) L/- Connors (1920); Lammerts (1945);
Bailey and French (1949)
Double (D1) d1/d1 2 More than five petals; often incompletely Lammerts (1945)

R. Monet and D. Bassi

dominant; number of extra petals controlled
by one or two recessive genes
Fewer extra petals (Dm1, Dm2) dm1/dm1 Dm1 and Dm2 are independent and additive Lammerts (1945); Yamazaki et al. (1987)
dm2/dm2
Dark pink petal (P) P/– Lammerts (1945)
Red petal (R) r/r Lammerts (1945); Chaparro et al. (1995)
Male sterility (from ‘J.H. Hale’) (Ps) ps/ps 6 Sometimes has some viable pollen Connors (1926); Blake and Connors
(1936); Scott and Weinberger (1944)
Male sterility (from ‘White Glory’) (Ps2) ps2/ps2 Blake (1932); Werner and Creller (1997)

Fruit
Slow ripening (Sr) sr/sr Ramming (1991)
Saucer (flat) shape (S) S/– 6 S/S is lethal (Guo et al., 2002) Lesley (1940)
Aborting fruit (Af)b af/af 6 Dirlewanger et al. (2006)
Blood red flesh (Bf) Bf/– Pigment appears in immature fruit and main Blake (1932); Werner et al. (1998)
leaf vein; often smaller trees
Rough skin (Rs) rs/rs Matte skin surface; glabrous flower buds Okie and Prince (1982); Okie (1988b)
Glabrous skin (nectarine) (G) g/g 5 Fuzzless Blake (1932); Blake and Connors (1936)
Full red skin (Fr) fr/fr Only on fruit Beckman and Sherman (2003)
Highlighter (H) h/h Red colour suppression on fruit skin Beckman et al. (2005)
White flesh (Y) Y/– 1 Also affects calyx cup and leaf colour Connors (1920)
Flesh texture and pit adherence (F)c
Melting freestone F/– 4 Bailey and French (1932, 1949);
Monet (1989); Peace et al. (2005)
Melting clingstone f/f Peace et al. (2005)
f/f1
f/n
Non-melting clingstone f1/f1 Peace et al. (2005)
f1/n
n/n
Stony hard flesh (Hd)d hd/hd Yoshida (1976); Scorza and
Sherman (1996)
hdhd/F Stony hard, melting Haji et al. (2005)
hdhd/f1f1 Stony hard, non-melting Haji et al. (2005)
Low-acid flesh (D) D/– 7 D for douce (sweet, in French) Monet (1979)
Sweet kernel (Sk) Sk/sk Werner and Creller (1997)

Genetics and Breeding

aDirlewanger et al. (2004).
bThe ‘aborting fruit’ has been reported as a recessive trait causing the abortion of all fruits within 2 months after full bloom; since it co-segregates with flat fruit
shape at the homozygous level (S/S), it is still not clear whether the ‘aborting fruit’ phenotype is regulated from the same locus or from a novel gene (Af).
cFour alleles at the same locus controlling both flesh texture (endopolygalacturonase enzyme expression) and pit adherence; the fourth, null allele (n), has the

same effect as the f1 allele (non-melting clingstone) (Peace et al., 2005).

dThe independent inheritance of this trait was demonstrated, also suggesting an epistatic influence on the F locus, since the stony-hard, melting (hdhd/f–)

phenotype is induced to soften when exogenous ethylene is applied (Haji et al., 2005).

65
66 R. Monet and D. Bassi
markers have permitted establishment of
these groups and the position of some of these
genes on them (see Chapter 4).
Nevertheless, the observation of characters
in F2 families allowed determination in some
cases of independent disjunction or linkage
(Table 3.2) (Bailey and French, 1949; Monet,
1967; Monet and Bastard, 1972, 1983; Monet
et al., 1985, 1988; Chaparro et al., 1995).
Quantitative characters
Characters with quantitative inheritance have
great importance in selection, because they
are often related to the economic value of the
harvested fruit. Traits such as fruit size, fruit
colour, firmness and taste were considered at
the beginning of peach classical breeding. These
traits have polygenic control and are influenced
by environment changes. Within a population,
their variation is not discrete but continuous.
Thus, it is difficult to know precisely the number
of minor genes involved that control expres-
sion of the character.
Quantitative genetics, which takes into
account all of these specificities, is essentially
mathematics. It works on population distri-
bution graphs from which are compared the
characteristics of central values (means) and
the characteristics of dispersion (variances). It
evaluates, for one character, the share that is
attributable to gene function and that due to
environmental effect. It also tries to predict
the effect of one reproductive cycle on the
character evolution. Finally, it provides a
mathematical basis for selection to help breed-
ers. Heritability (h) is one of them; it is defined
by the ratio of the variance attributable to
additive effects of genes on the character (VA)
to the phenotypic variance (VP), i.e. heritabil-
ity in the narrow sense:
h2 = VA/VP.
In some situations, the heritability of a
character can be estimated by the coefficient
of regression between offspring to mid-
parent:
h2 = bPE.
In this case, if S is the applied level of selec-
tion on a quantitative character, the response
R is proportional to the regression coefficient
and then to heritability:
R = bPE S = h2S.
If heritability is small, near zero, there will be
no response. If it is near one, the response will
be strong. Thus, for a breeder, the knowledge
of the heritability of one character under
selection is of major importance because it
indicates if selection will be efficient or not.
For peach, Hansche et al. (1972) and Hansche
and Boyton (1986a,b) estimated heritability of
the main agronomic characters. The values
they obtained are reported in Table 3.3.
Ploidy
Peach is a diploid plant with 2n = 16 chromo-
somes. Sometimes in this species one can find
monoploids (or haploids) in progenies that
have received only the basic chromosome
number (n = 8). This set comes from the female
parent, from which a cell of its embryo sac
(oospore, generally) divides without fertiliza-
tion and gives an embryo.
Hesse (1971) made a detailed description
of two monoploids obtained, by chance, in a
progeny resulting from a cross between a nec-
tarine and a canning clingstone peach. These
trees were dwarf and possessed all the recessive
characters of the female parent. Moreover,
shoots were slender, with short internodes;
leaves were narrow and long; flowers and
fruits were small. In ageing, the shoot cam-
bium developed outgrowths which gave them
a knotty appearance. Hesse (1971) also observed
the pollen grain germination of these mono-
ploids. The percentage of grains able to ger-
minate was low (less than 7%). These viable
grains would have the normal basic chromo-
some number.
Toyama (1974) searched systematically
for monoploids in peach seedlings, trying to
isolate them before their germination. He focu-
sed on seeds with two embryos (one of which
may have proceeded from a monoploid non-
fertilized cell of the embryo sac). He obtained
16 monoploids, five of them from seeds with
two embryos. For this result, he examined
20,053 seedlings which gave a relatively high
Table 3.2. Linkages and independent disjunctions in peach.

Corky Red Anthocyanin Antho- Foliar Non-showy/ Male White flesh/ Melting flesh/ Flat Low-acid Sweet Myzus
Weeping triangle leaves deficicient cyaninless glands showy flower sterility Nectarine yellow flesh non-melting flesh fruit flesh kernel resistance

Weeping

Corky Indt
triangle
Red leaves ?? ??

Anthocyanin ?? ?? ??
deficient
Antho- ?? ?? Indt ??
cyaninless

Genetics and Breeding

Foliar glands ?? ?? ?? ?? ??

Non-showy/ Indt ?? ?? Indt ?? Indt

showy flower
Male sterility ?? ?? ?? ?? ?? Indt ??

Nectarine ?? ?? ?? ?? ?? ?? Indt ??

White flesh/ Indt Indt ?? ?? ?? Indt Indt Indt Indt

yellow flesh
Melting flesh/ ?? ?? ?? Indt ?? Indt Indt ?? ?? Indt
non-melting
flesh
Flat fruit ?? ?? ?? ?? ?? ?? ?? Indt ?? Indt ??

Low-acid ?? ?? ?? ?? ?? ?? ?? Indt ?? Indt ?? Link

flesh 30 cM
Sweet kernel ?? ?? ?? ?? ?? ?? ?? ?? Link 12 cM ?? ?? ?? ??

Myzus Indt Indt ?? ?? ?? ?? Indt ?? ?? Indt ?? ?? ?? ??

resistance

Indt, independent; Link, linkage; ??, not observed.

67
68 R. Monet and D. Bassi
Table 3.3. Estimates of heritability of major quantitative traits in peach.
Trait
Full bloom
Amount of ripening
Ripening date
Crop
Fruit length
Fruit cheek
Fruit suture
Fruit firmness
Fruit acidity
Soluble solids
Juvenility (flower number)
Intensity of browning
ratio (approximately 8 out of 10,000). Toyama
obtained pure lines by doubling the chromo-
some number of some of these monoploids
with colchicine.
Pooler and Scorza (1995) observed the
pollen grains of monoploid peach trees. Some
of them, not reduced, were diploids and capa-
ble of germination. Thus it is possible to
obtain triploid peach trees.
We observed (R. Monet, unpublished
results) that monoploid peach trees can give
some fruits (Fig. 3.1/Plate 35). The seeds asso-
ciated with these fruits were able to germi-
nate and some were triploids; others were
diploids and aneuploids. Triploids were vig-
orous but with little fertility; diploids, if they
resulted from self-pollination (the monoploid
parent was maintained in a greenhouse dur-
ing flowering but without protection), were
pure lines; aneuploids, which have a chromo-
some in excess or in deficit, were small plants
with a very slow rate of development.
3.2 Peach Breeding
First steps
The first peach cultivars to be developed
resulted from chance seedlings that were iso-
lated and propagated by grafting. In this way,
each country has constituted a cultivar pool
Heritability Reference
0.39 Hansche et al. (1972)
0.38 Hansche et al. (1972)
0.84 Hansche et al. (1972)
0.08 Hansche et al. (1972)
0.31 Hansche et al. (1972)
0.26 Hansche et al. (1972)
0.29 Hansche et al. (1972)
0.13 Hansche et al. (1972)
0.19 Hansche et al. (1972)
0.01 Hansche et al. (1972)
0.16 Hansche and Boyton (1986a)
0.35 Hansche and Boyton (1986b)
whose richness has increased continually with
new discoveries and germplasm exchanges.
By the 18th century, ‘physiocrats’ of a
French economics school considered agricul-
ture as a unique source of ‘net product’. They
originated important literature to describe
the best agricultural practices and make
inventories of the best productive genetic
material. The first pomologies were compila-
tions of all the available fruit cultivars with
their main characters, their agronomical inter-
est and their organoleptic qualities. Each coun-
try with an orcharding tradition has established
cultivar records whose content has increased
continually. In France, the Duhamel du Mon-
ceau (1768) pomology book described 42
peach cultivars; one century later, the one of
Leroy (1879) described 143 cultivars.
Attempts at classification based on the
main botanical characters were also made.
For example, De Mortillet (1865) made a
dichotomous classification of peach cultivars
depending on whether they had epidermal
fuzz or not, whether they were freestone or
clingstone, whether their flowers were large,
medium or small and whether their foliar
glands were reniform, circular or absent.
In the USA, Downing’s pomology (1866)
described 136 cultivars. It mentioned, probably
for the first time in fruit culture, hybridization
as a new method to obtain varieties:
Cross-breeding is then nothing more than
removing out of the blossom of the fruit tree
 Genetics and Breeding
Fig. 3.1. A monoploid peach with some fruits.
the stamens, or male parent, and bringing
those of another, and different variety of fruit,
and dusting the pistil of female parent with
them, – a process sufficiently simple, but
which has the most marked effect on the
seeds produced. It is only within about fifty
years that cross breeding has being practiced.
Hedrick’s pomology (1917) describes culti-
vars such as ‘J.H. Hale’, ‘Elberta’ and ‘Chinese
Cling’, which were extensively used in the
first peach breeding programmes.
Pomological records provide the first step
in the search and inventory of the best fruit
genetic material, but they do not have inno-
vative capacity. When Hedrick published his
pomology in the USA, the first peach breed-
ing programmes were already initiated (Con-
nors, 1917), and their contribution significantly
modified the available cultivar assortment
worldwide.
Objectives
One can criticize the first generation of breed-
ers for their emphasis on improvement of
commercial characteristics of the fruit such as
69
colour, firmness and attractiveness while not
focusing on taste, hardiness or their adapta-
tion to economically efficient growing systems.
The improvement of commercial characteris-
tics was nevertheless necessary, because old
cultivars were not very attractive, unsuitable
for handling and shipping and did not satisfy
commercial requirements. Today one can
consider these defects almost completely
corrected.
Some 30 years ago, breeders developed
objectives which had interest for consumers
and growers. These innovative objectives are
presented below (Childers, 1975; Monet, 1995;
see also Chapters 5, 6 and 7).
Reduction of production costs
A cultivar has low production costs if it is
well adapted to the place where one culti-
vates it. If it is not adapted, conditions favour-
able to its culture must be obtained artificially,
and that will increase costs. Adaptation to
climatic factors has been and continues to be
an objective of some breeding programmes,
like obtaining hardiness of dormant buds for
northern-limit zones of peach culture as in
70 R. Monet and D. Bassi
the Great Lakes region in Canada (Layne,
1982) and Michigan and New York in the USA
(Callahan et al., 1991), or creating low-chill
cultivars for subtropical regions as in Florida,
Texas (Sherman and Rodriguez-Alcazar, 1987)
and Georgia (Krewer et al., 1997) in the USA
and Rio Grande do Sul in Brazil (Nakasu
et al., 1981).
Also interesting are the projects whose
goal is to improve disease and pest resistance.
The reduction of spray applications is desir-
able because they reduce production costs and
environmental contamination. Indeed genetic
engineering could bring, in the near future,
original and rapid solutions for problems of
pest and disease resistance. However, consum-
ers are still reluctant in using food products
derived from transformed plants, the so-called
‘genetically modified organisms’.
Using a classical approach, many breed-
ing programmes have attempted to improve
peach pest and disease resistance.
● Bacterial leaf spot (Xanthomonas campes-
tris) in the USA (Werner et al., 1986).
● Brown rot (Monilinia fruticola Aderh. et
Ruhl. (Honey)) in the USA (Gradziel et al.,
Fig. 3.2. A weeping tree trained in central axis.
1997) and Europe (Pascal et al., 1994;
Bassi et al., 1998).
● Cytospora canker (Leucostoma personii and
Leucostoma cincta) in the USA (Chang et al.,
1989) and Italy (Liverani and Fideghelli,
1993).
● Green peach aphid (Myzus persicae) in
France (Monet et al., 1997; Pascal et al.,
1997) and Italy (Liverani and Fideghelli,
1993; Bassi et al., 1994).
● Leaf curl (Taphrina deformans) in the USA
(Scorza, 1992) and Italy (Bellini et al.,
1993).
● Powdery mildew (Spherotheca pannosa) in
France (Pascal, 1990) and Italy (Liverani
and Fideghelli, 1993).
One can reduce production costs with the use
of cultivars whose tree size and shape are
naturally adapted to the growing system.
Thus, breeding efforts to develop dwarf trees
for high-density plantings (Fideghelli et al.,
1979; Hansche, 1988; Liverani and Fideghelli,
1993), ‘pillar’ (columnar) trees for tree–wall
systems (Scorza et al., 1989) and weeping trees
(Fig. 3.2/Plate 36) for central leader pruning
(Monet et al., 1988) have been explored.
 Genetics and Breeding
Diversification for the consumer
Consumers like variation in their food
choices. To bring diversity, peach breeders
draw on Mendelian genetic variability. In
the past, white- or yellow-fleshed peaches
only could be found in markets but now
both are available. Then, from the 1960s,
came nectarines. New Mendelian characters
that modify other fruit attributes and taste
are being introduced. The flat peach charac-
ter has been the object of several breeding
programmes, e.g. in the People’s Republic of
China, France (Monet et al., 1985), Italy and
the USA (Sherman and Lyrene, 2001). Blood-
fleshed peaches and nectarines are being
developed in France (T. Pascal, Avignon, 1995,
personal communication) and in the USA
(Okie, 1988a). Conversely, anthocyaninless
71
nectarines have been created in the USA
(Okie, 1988a). Canning clingstone anthocya-
ninless nectarines are also the objective of a
breeding programme in Italy (Bellini et al.,
1994) and the USA.
All of these Mendelian characters can be
recombined. For example, the ‘honey’ character
(low-acid), very popular only in the Far East
countries, has been introduced also in breed-
ing programmes in Europe and the USA. It
permits the harvest of fruit earlier without
adversely affecting the taste. Combining flat,
low-acid and nectarine, a sweet, flat nectarine
(‘platerine’) has been obtained (Fig. 3.3/Plate
37). In the same way, the introduction of the non-
melting flesh character in cultivars for the
fresh market gives the possibility of harvest-
ing the fruits near maturity at their optimal
Fig. 3.3. Flat nectarines (‘platerines’
in French).
72 R. Monet and D. Bassi
quality and taste with greater durability for
handling and transport to distant markets.
Finally, these numerous projects are likely
to bring innovations in peach culture with
more interest for growers and consumers. This
represents a change in emphasis compared
with the breeding for commercial and horti-
cultural objectives only.
Floral biology and selection cycles
Although peach is preferentially autogamous,
there is no impediment to intraspecific cross-
ing. The breeder can include in a selection
scheme self-pollination and intraspecific
crosses. This is an ideal situation; the sole
break of progression is the 3-year sexual cycle.
Seedlings bear their first harvest from 2 to 3
years after obtaining the seeds, depending on
climatic and cultivation conditions. For fur-
ther details also see Chapter 1.
Methodology
Peach cultivars are clones, propagated by
grafting. Thus, the simplest selection meth-
ods can give results as good as the more
complex. An exceptional recombination
may occur by chance, resulting in a tree that
possesses the required commercial attri-
butes. This tree then can become the mother
tree, from which one can cut scions neces-
sary for its multiplication to be perpetuated
in space and time.
To obtain this mother tree, one can utilize
mutagenesis, polyploidization or character
recombination by inter- or intraspecific sexual
reproduction. Mutagenesis, polyploidization
and interspecific sexual reproduction have
been used very little in peach breeding; results
obtained by these methods have been largely
unsatisfactory. Alternatively, recombinations
obtained by intraspecific cross-breeding have
allowed noteworthy improvements of the
species and continue to supply most of the
replacement cultivars. The general breeding
schemes (see Monet, 1995; Scorza and Sherman,
1996) are more or less complicated and are
presented below.
Parental choice: phenotypic
This very simple method is always used in
private breeding programmes. It consists of
crossing two individuals which present com-
plementary phenotypic characters. For exam-
ple, one can cross a cultivar with soft-fleshed
fruit but whose epidermis is very coloured
with a cultivar with firm-fleshed fruit but
whose epidermis is little coloured in the hope
of obtaining, in the progeny, individuals with
firm flesh and very coloured skin. This
method has led to improvement of most of
the fruit characters, especially those in rela-
tion to their commercial attributes. Some
breeders also cross the best cultivars available
at the time to obtain better recombinations.
This method has several advantages.
● It is simple and fast, since it does not use
long-term schemes.
● The majority of characters of interest
have quantitative inheritance with high
heritability.
● Working with large families increases one’s
chance to obtain favourable combinations.
Nevertheless, it has some disadvantages.
● The repeated use of the best cultivars leads
to phenotypic homogeneity (e.g. in the
market, peaches become more and more
similar). This is the result of their geno-
typic homogeneity, a consequence of the
close parentage of their progenitors.
● A phenotype results from the additive ef-
fect of genes plus the genetic interactions
(dominance and epistasis effects), so that
the choice of two parents on their pheno-
typic value alone can result in progenies
without interest. While individual breed-
ers may gain empirical experience after
the observation of many crosses, they
may not like to share this information
with other breeders (particularly in the
private sector).
Parental choice: genotypic
To really know the genetic value of a parent, it
is necessary to observe its progeny. In peach,
the possibility to do self-pollination permits an
accurate evaluation of this value. It provides
 Genetics and Breeding 73

valuable information on its heterozygous state
(the more the progeny is heterogeneous, the
more the parent is heterozygous). It informs
also on its capacity to transmit important hor-
ticultural characters. To conduct self-pollina-
tion of progenitors before crossing could
permit a better choice of parents. In fact this is
not often done because it lengthens the breed-
ing process by a new sexual cycle.
Monet (1995) tried to obtain pure lines
with peach using a genotypic choice of the
parents. The method is as follows.
● Many cultivars of great interest are self-
pollinated simultaneously; the observa-
tion of the obtained families permits the
elimination, between these parents, of
those with insufficient value.
● In each of the remaining progenies, two
of the best individuals are chosen (or more,
but it increases the work load), which are
self-pollinated; the comparison of the
two obtained families permits an elimi-
nation of the less interesting individual
and its family.
● In the family resulting from the best-
performing individual, the two best in-
dividuals are self-pollinated to choose
the parents of the next generation, and so
on.
This process (Fig. 3.4/Plate 38) has been fol-
lowed during six generations for one parent
and less (three to five) for eight others. After
six self-pollination cycles, the resemblance
between individuals is more and more strong
and shows that we are near the pure line. The
inbreeding depression is light probably because
peach is preferentially an autogamous spe-
cies. The cross between individuals taken from

Starting parents A B C

Self-pollination

G1

Parent eliminated Family treated like A

Self-pollination Self-pollination to obtain a new
pure line

G2

Parent eliminated

Self-pollination Self-pollination

G3

Self-pollination Self-pollination Parent eliminated

G4
Fig. 3.4. Scheme based on genotypic choice of parents to obtain pure lines. In each generation,
parents giving progenies with low horticultural value are eliminated.
74 R. Monet and D. Bassi
families with three self-pollination cycles gave
families with good agronomic performances
and this process led to the elimination of indi-
viduals without agronomic interest. However,
intercrossing pure lines obtained by repeated
self-pollination (Lesley, 1957) or pure lines
obtained by doubling the chromosome num-
ber of monoploids (Scorza and Pooler, 1993)
shows that there are no important heterosis
effects with peaches. Therefore, the use of pure
lines can be reserved for theoretical research
but has little agronomic interest.
Character of Mendelian inheritance
The goal of many current breeding program-
mes is to create cultivars which possess a par-
ticular character with Mendelian inheritance.
These characters do not change from one gen-
eration to the next. If they are dominant they
are present in each generation; if they are reces-
sive, they can disappear by cross-breeding, but
they will reappear after self-pollination. The
breeder is in a particularly comfortable situa-
tion. The main difficulty in these programmes
results from the fact that the parent that brings
the Mendelian character could be near the
wild state and produce fruits that are neither
edible nor marketable and also can present
important horticultural defects. It is only after
several recurrent crosses with an improved
cultivar that these defects will be eliminated.
Self-pollination exploits recombinations and
recessive characters, while intra- and interspe-
cific crossing permits the introduction of new
characters within the genetic pool under
selection. Figure 3.5 shows a breeding scheme
used to introduce the resistance to green
aphid using a monogenic dominant resistance
found in a weeping ornamental peach. It
employs alternation of self-pollination and
crossing. Five generations were required to
obtain the first marketable selections, more
than 20 years after the beginning of the pro-
gramme.
Technique
Breeding techniques have been described in
detail by Hesse (1975); below we recall only
the main operations followed in peach.
Operations during flowering
CROSSING PROCEDURES. Peach flowers, being
hermaphroditic and self-fertile, require emas-
culation of the flowers of the female parent
before pollination. Emasculation takes place
at balloon stage, i.e. some days before flower
opening (for non-showy flowers, because an-
thers emerge beyond the petals, it should be
ensured that none of them are open before
emasculation is initiated). With tweezers, the
part of the floral receptacle that bears the an-
thers, sepals and petals is eliminated (Fig.
3.6/Plate 39). After emasculation, all that re-
mains is the pistil and a little piece of the flo-
ral receptacle. An emasculated flower does
not attract insects; thus out-crossing is rare
and usually it is not necessary to protect a tree
where all flowers are emasculated. However,
for genetic studies, to be sure that there are no
accidental pollinations by wind or insects (i.e.
honeybees; Fig. 3.7/Plate 40), a cage with a
fine grid can be placed over the tree, which
will be maintained during flowering of the
neighbouring trees.
The male parent will provide the desired
pollen. Flowers are harvested at the balloon
stage (when anthers are not dehiscent). With
tweezers the flowers are opened; the anthers
are removed and placed in a Petri dish which
is maintained in a dry place or in a desiccator
with anhydrous calcium sulfate to obtain
dehiscence. Dry pollen can be stored for up to
2 years in a refrigerator at –30°C or for many
years at –80°C.
Pollination of the female parent can be
done 24–48 h after emasculation. Pollen, col-
lected from the anthers of the male parent
(Fig. 3.8/Plate 41), is brought with a brush (or
by fingertip) (Fig. 3.9/Plate 42) and applied to
the stigmas of emasculated flowers. To save
pollen, glass rods could be used to apply pol-
len on to pistils.
SELF-POLLINATION. This results from the natu-
ral transfer of pollen from dehiscent anthers to
the stigma of the same flower. It is not necessary
to intervene by hand to obtain self-pollination,
even if it could improve pollen transport on
to the stigma and then fruit yield. Before
flowering, branches can be wrapped with wa-
terproof paper bags (Fig. 3.10/Plate 43) or the
 Genetics and Breeding 75

Starting parents Redhaven Weeping ornamental peach

clone S1161 clone S2678

Self-pollination Self-pollination

F0 Family (1971) S1161: (167 trees) S2678: (55 trees)

Parents of 1st cycle S1161: 12 S2678: 47

Cross-pollination

F1 Family (1975) S(1161:12 x 2678:47) (2 trees)

Parent of 2nd cycle S(1161:12 x 2678:47)1

Self-pollination

F2 Family (1981) S(1161:12 x 2678:47)1: (55 trees)

Early Sungrand
Parents of 3rd cycle S(1161:12 x 2678:47)1: 55 clone S3965

Cross-pollination

F3 Family (1987) S[(1161:12 x 2678:47)1: 55 x 3965] (64 trees)

Parent of 4th cycle S[(1161:12 x 2678:47)1: 55 x 3965] 56

Self-pollination

F4 Family (1991) S[(1161:12 x 2678:47)1: 55 x 3965] 56: (64 trees)

In 1994, two selections (trees nr. 5 and nr. 14) were retained in this F4 family for evaluation

Fig. 3.5. Scheme used for the development of cultivars resistant to green aphid, Myzus persicae.
(From Monet, 1995.)

whole tree can be protected by an insect-proof
cage. These barriers against accidental pol-
linations are removed when flowering is
completed.
Raising seedlings
Pollinated flowers give fruits and seeds. When
these seeds have a normal development, as in
medium- and late-ripening trees, germination
can be easily attained without difficulty by
humid stratification at 0–4°C for 3 months, in
order to overcome dormancy. Seeds are placed
(better without the pit, to improve and speed
germination) in moist sand, poured into bags
(alternatively, into sealed Petri dishes with
moist paper) and stored in a cold room at
0–4°C. After 3–5 months, germination occurs,
76 R. Monet and D. Bassi

Fig. 3.6. Emasculated flower (on left). (Courtesy of W.R. Okie, Byron, Georgia, USA.)

Fig. 3.7. Pollination by honeybee. (Courtesy of D.R. Layne, Clemson, South Carolina, USA.)
 Genetics and Breeding 77

Fig. 3.8. Dehydrated stamens ready for artificial pollination: anthers and filaments render the pollen
less sticky.

Fig. 3.9. Emasculated flower pollinated by finger, a fast way for cross-breeding.
78 R. Monet and D. Bassi
Fig. 3.10. Self-pollination with waterproof paper bags.
a young root appears out of the seeds and
the germinated seeds are transplanted in the
greenhouse.
When seeds are insufficiently developed
(for seeds obtained from mother parents which
ripen their fruits less than about 110–120 days
after flowering), embryos of these seeds have
little reserves and are too weak to germinate in
normal stratification; embryo culture should be
used to germinate them. Embryo culture in vitro
avoids bacterial or fungal contamination, pre-
vents rapid dehydration, and provides essen-
tial nutrients to the embryo. In practice, the
different steps of this technique are as follows.
● Fruits are harvested at the veraison stage
(before physiological ripening), their flesh
is removed, seeds are removed from the
pits and their integuments are sterilized
in a solution of calcium hypochlorite (8 g
calcium hypochlorite in 20 ml water, 20
min of stirring, filtration) for 10 min and
then washed in sterile water.
● Inside a sterile room the embryo is ex-
tracted from the seed; with a sterile pair of
tweezers, the integuments of the seed are
removed and the embryo is separated from
the endosperm, then it is placed in a flask
or test-tube with the nutritive medium.
● The flasks containing the embryos are
placed in a cold room at 0°C for 1 month
to eliminate dormancy and then taken to
a growing room (at 22–24°C).
● When their development is sufficient
(Fig. 3.11/Plate 44), they are transplanted
in a greenhouse.
A comprehensive review on embryo cul-
ture in fruit breeding was given by Ramming
(1990). Obtained by classical stratification or
by embryo culture, seedlings continue their
growth in a greenhouse during winter. At the
end of the dormant season, they can be trans-
planted in the orchard. Plantation distances
vary from one breeder to another. Some pre-
fer high densities of 0.5 m × 1 m, others prefer
a wider spacing of 2 m × 5 m. High density
induces competition between trees and
reduces fruit quality, making it more difficult
for the breeder to choose the best individuals
(Fig. 3.12/Plate 45).
Seedling evaluation and selection
Sexual reproduction results in the genetic
recombination of characters. Thus, between
individuals from the same family issued from
a cross or a self-pollination, there is a variability
 Genetics and Breeding 79

Fig. 3.11. Young seedling at the end

of embryo culture.

Fig. 3.12. Seedling orchard at first flowering.

80 R. Monet and D. Bassi
associated with genetic effects and, for quan-
titative characters, there is also a variability
associated with environmental effects. The
problem for the breeder is how to choose the
best recombinant.
For Mendelian characters, selection in a
progeny is straightforward; the breeder has
two simple options: presence or absence of
the character.
For quantitative characters, the choice is
more difficult, and a good definition of selec-
tion criteria is fundamental. Measuring the
character is, in this case, a great advantage
because it suppresses all subjective influences.
Finally, for pest and disease resistance,
there is a need for careful evaluation of tree
contamination and damage assessment that
is specific for each pest and disease.
The selection criteria depend on the breed-
er’s objectives but some must be considered
intangible. A cultivar has no chance to have
great success if it does not respond to them.
They can be of horticultural importance like
the degree of fertility (yield). Overall, commer-
cial attributes are essential: fruits should be
attractive, firm and have good taste. To these
basic objectives are associated specific objec-
tives (introduction of a new character, a dis-
ease resistance, etc.) that justify the breeding
programme.
For individuals of families under selec-
tion, it is essential to record the value for each
important character. The simplest method is
recording in a notebook; but, today, many
electronic devices are available that permit
transfer of data to the computer for data
processing.
A breeder has the tendency to focus on
some characters more than others and may
forget that a cultivar is a compromise between
horticultural and commercial attributes. To
avoid this drawback, a selection index (I)
could be established. The simplest is to calcu-
late, for each tree of the family, the sum of
products of the value given to each character
(C) included in the index by a coefficient (n)
reflecting its importance:
I = C1n1 + C2n2 + . . .
Trees with the highest index will be selected
for in the next generation (or eventually
assessed as potential new cultivars).
Release of a new cultivar
When the level of improvement is sufficient,
the breeder possesses within his families indi-
viduals with the potential to become new cul-
tivars. Trees can be propagated by grafting
for a comparative test with cultivars of the
same period of maturity. This test is essential
to avoid mistakes because a judgement based
on one individual within a family is not the
same as when this individual is grafted. The
comparative test must be simple; a statistical
plan is not necessary. For example, two grafted
trees from each advanced selection will be
placed side by side with two grafted trees of
the reference cultivar. The new advanced selec-
tion can be easily compared with the refer-
ence trees.
At the present time, it is also important
to evaluate the sanitary state of the future
new cultivar. Bud sticks or leaves should be
taken from the original tree and used for labo-
ratory tests to detect viruses and phytoplas-
mas that can be transmitted by grafting. For
peach, the tests may search specially for: Plum
pox virus, the family of Isometric Labile Ring-
spot virus, Nematode polyhedric virus and Euro-
pean stone fruit yellows phytoplasma. If the plant
is healthy, it is recommended to keep some
copies of it inside an insect-proof repository
to preserve a source of buds with their origi-
nal qualities. If it is contaminated, it is better
to eliminate it because the success of virus or
phytoplasma therapies is not assured, even
though for selections of particular interest
efforts could be justified in order to obtain a
healthy tree. However, the basis of vegetative
propagation of the new cultivar will come
from healthy plants.
To create a new variety is expensive and
justifies a return on the investment. This
needs legal protection. In the past, commer-
cial patenting was done only by private
breeders. Today, cultivars from public pro-
grammes are also being patented. The condi-
tions of patenting are different from one
country to another. In the USA patenting a
cultivar has the same value as patenting an
industrial process. In Europe, a new cultivar
receives a certificate (Certificat d’Obtention
Varietale) that has approximately the same
value as a patent. To be patented a cultivar
 Genetics and Breeding 81

must be original (not a copy of another exist-
ing cultivar) and healthy. The legal protection
lasts 20 years; it covers its phenotype only but
there is a strong demand for an extension to
the genotype.
The release of a new cultivar is always a
point of satisfaction for the breeder because it
gives evidence of the success of the project.
However, the commercial success of a new
cultivar depends on many factors: its agro-
nomic value can be negated by a defect that
may become evident only in the orchard.
New peach cultivars have a short lifespan: 10
to 20 years approximately. If we compare this
duration with that needed to create a truly
innovative cultivar (20 years on average), one
can say that it is an ungrateful job. However,
some cultivars retain their commercial use-
fulness for many years, e.g. ‘Redhaven’, but
they tend to be the exception rather than the
rule. The problem lies in the fact that the
breeder is often aiming at a moving target.
While the new cultivar may have successfully
combined the desired characters that were
sought when the programme was initiated,
the cultivar requirements of the market may
have changed during the 15–20-year period
in which the cultivar was being developed.
Thus, the new cultivar may not meet the exist-
ing market requirements when released. This
is an inherent risk in peach breeding, but the
evidence of genetic advancement in new
peach cultivars compared with older ones
shows that it is a risk well worth taking.

References

Bailey, J.S. and French, A.P. (1932) The inheritance of certain characters in the peach. Proceedings of the
American Society for Horticultural Science 29, 127–130.
Bailey, J.S. and French, A.P. (1942) The inheritance of blossom type and blossom size in the peach. Proceed-
ings of the American Society for Horticultural Science 40, 248–250.
Bailey, J.S. and French, A.P. (1949) The inheritance of certain fruit and foliage characters in the peach.
Massachusetts Agricultural Experiment Station Bulletin 452.
Bassi, D. and Rizzo, M. (2000) Peach breeding for growth habit. Acta Horticulturae 538, 411–414.
Bassi, D., Liverani, A. and Rizzo, M. (1994) Il miglioramento genetico del pesco in Emilia Romagna: risultati
e prospettive. Rivista di Frutticoltura 1, 11–23.
Bassi, D., Rizzo, M. and Cantoni, L. (1998) Assaying brown rot (Monilinia laxa Aderh. et Ruhl. (Honey))
susceptibility in peach cultivars and progeny. Acta Horticolturae 465/2, 715–721.
Beckman, T.G. and Sherman, W.B. (2003) Probable qualitative inheritance of full red skin color in peach.
HortScience 38, 1184–1185.
Beckman, T.G., Rodriguez Alcazar, J., Sherman, W.B. and Werner, D.J. (2005) Evidence for qualitative sup-
pression of red skin color in peach. HortScience 40, 523–524.
Bellini, E., Surico, G., Mugnai, L., Natarelli, L. and Nencetti, V. (1993) Osservazioni su una progenie di pesco
resistente a Taphrina deformans (Berck. Tul.). Italus Hortus 1, 11–13.
Bellini, E., Giannelli, G., Giordani, E. and Sabbatini, I. (1994) Costituzione di nettarine a polpa bianca e di
percoche a buccia liscia. Rivista di Frutticoltura 4, 23–37.
Bielenberg, D.G., Wang, Y., Fan, S., Reighard, G.L., Scorza, R. and Abbott, A.G. (2004) A deletion affecting
several gene candidates is present in the evergrowing peach mutant. Journal of Heredity 95, 436–444.
Blake, M.A. (1932) The J.H. Hale peach as a parent in peach crosses. Proceedings of the American Society for
Horticultural Science 29, 131–136.
Blake, M.A. (1937) Progress in peach breeding. Proceedings of the American Society for Horticultural Science
35, 49–53.
Blake, M.A. and Connors, C.H. (1936) Early results of peach breeding in New Jersey. New Jersey Agricultural
Experiment Station Bulletin 599.
Callahan, A., Scorza, R., Morgens, P., Mante, S., Cordts, J. and Cohen, R. (1991) Breeding for cold hardiness:
searching for genes to improve fruit quality in cold-hardy peach germplasm. HortScience 26, 522–526.
Chang, L.S., Lezzoni, A., Adams, G. and Howell, G.S. (1989) Leucostoma personii, tolerance and cold hardiness
among diverse peach genotypes. Journal of the American Society for Horticultural Science 114, 482–485.
Chaparro, J.X., Werner, D.J., O’Malley, D. and Sederoff, R.R. (1994) Targeted mapping and linkage analysis of
morphological, isozyme, and RAPD markers in peach. Theoretical and Applied Genetics 87, 805–815.
82 R. Monet and D. Bassi

Chaparro, J.X., Werner, D.J., Whetten, R.W. and O’Malley, D.M. (1995) Inheritance, genetic interaction, and
biochemical characterisation of anthocyanin phenotypes in peach. Journal of Heredity 86, 32–38.
Childers, N.F. (1975) The Peach. Horticultural Publications, New Brunswick, New Jersey, 659 pp.
Connors, C.H. (1917) Methods in breeding peaches. Proceedings of the American Society for Horticultural
Science 14, 126–127.
Connors, C.H. (1920) Some notes on the inheritance of unit characters in the peach. Proceedings of the
American Society for Horticultural Science 1919 16, 24–36.
Connors, C.H. (1921) Inheritance of foliar glands of the peach. Proceedings of the American Society for
Horticultural Science 18, 20–26.
Connors, C.H. (1926) The sterility of ‘J.H. Hale’. New Jersey Agriculture Experimental Station Annual Report
(1925) 46, 90–91.
De Mortillet, M.P. (1865) Les meilleurs fruits. Le pêcher. Prudhomme et Giroud, Grenoble, France.
Dirlewanger, E., Graziano, E., Joobeur, T., Garriga-Caldere, F., Cosson, P., Howad, W. and Arùs, P. (2004)
Comparative mapping and marker-assisted selection in Rosaceae fruit crops. Proceedings of the
National Academy of Sciences USA 101, 9891–9896.
Dirlewanger, E., Cosson P., Boudehri, K., Renaud, C., Capdeville, G., Tauzin, Y., Laigret, F. and Moing, A. (2006)
Development of a second generation genetic linkage map for peach [Prunus persica (L.) Batsch.] and char-
acterization of morphological traits affecting flower and fruit. Tree Genetics & Genome 1, 1–13.
Downing, C. (1866) The Fruits and Fruit Trees of America. Wiley, New York.
Duhamel du Monceau, M. (1768) Traité des arbres fruitiers. Tome 2. Le pêcher. Saillant, Paris.
Fideghelli, C., Della Strada, G., Quarta, R. and Rosati, P. (1979) Genetic semi-dwarf peach selections. In:
Proceedings of Eucarpia Fruit Section Symposium, Tree Fruit Breeding. INRA, Angers, France, pp. 3–7.
Gradziel, T.M., Thorpe, M.A., Bostock, R.M. and Wilcox, S. (1997) Breeding for brown rot (Monilinia fructi-
cola) resistance in clingstone peach with emphasis on the role of fruit phenolics. Acta Horticulturae 465,
161–170.
Guo, J., Jang, Q., Zhang, K., Zhao, J. and Yang, Y. (2002) Screening for molecular marker linked to saucer gene
of peach fruit shape. Acta Horticulturae 592, 267–271.
Haji, T., Yaegaki, H. and Yamaguchi, M. (2005) Inheritance and expression of fruit texture melting, non-melting
and stony hard in peach. Scientia Horticulturae 105, 241–248.
Hansche, P.E. (1988) Two genes that induce brachytic dwarfism in peach. HortScience 23, 604–606.
Hansche, P.E. and Boynton, B. (1986a) Heritability of juvenility in peach. HortScience 21, 1195–1197.
Hansche, P.E. and Boynton, B. (1986b) Heritability of enzymatic browning in peaches. HortScience 21, 1197–
1198.
Hansche, P.E., Hesse, C.O. and Beres, V. (1972) Estimate of genetic and environmental effects on several traits
in peach. Journal of the American Society for Horticultural Science 97, 9–12.
Hedrick, U.P. (1917) The Peaches of New York. J.B. Lyon Company, Albany, New York.
Hesse, C.O. (1971) Monoploid peaches, Prunus persica L. Batsch: description and meiotic analysis. Journal
of the American Society for Horticultural Science 96, 326–330.
Hesse, C.O. (1975) Peaches. In: Janick, J.J. and Moore, J.N. (eds) Advances in Fruit Breeding. Purdue University
Press, West Lafayette, Indiana, pp. 285–335.
Krewer, G., Beckman, T.G. and Sherman, W.B. (1997) Moderate chilling peach and nectarine breeding and
evaluation program at Attapulgus, Georgia. Acta Horticulturae 465, 171–175.
Lammerts, W.E. (1945) The breeding of ornamental edible peaches for mild climates. I. Inheritance of tree and
flower characters. American Journal of Botany 32, 53–61.
Layne, R.E.C. (1982) Cold hardiness of peaches and nectarines following a test winter. Fruit Varieties Journal
36, 90–98.
Ledbetter, C.A. (1996) Characterization and inheritance of the crinkle-leaf trait in peach and peach–almond
hybrids. Journal of Genetics and Breeding 50, 5–60.
Leroy, A. (1879) Dictionnaire de pomologie. Tome VI. Fruits à noyau. Goin, Paris.
Lesley, J.W. (1940) A genetic study of saucer fruit shape and other characters in the peach. Proceedings of the
American Society for Horticultural Science 37, 218–222.
Lesley, J.W. (1957) A genetic study of inbreeding and of crossing inbred lines in peaches. Proceedings of the
American Society for Horticultural Science 70, 93–103.
Liverani, A. and Fideghelli, C. (1993) Il miglioramento genetico del pesco: risultati e prospettive. Rivista di
Frutticoltura 5, 11–21.
Lownsberry, B.F. and Thomson, I.J. (1959) Progress in nematology related to horticulture. Proceedings of the
American Society for Horticultural Science 74, 730–746.
 Genetics and Breeding 83

Lu, Z.X., Reighard, G.L., Nyczepir, A.P., Beckman, T.G. and Ramming, D.W. (2000) Inheritance of resistance
to root-knot nematodes (Meloidogyne sp.) in Prunus rootstocks. HortScience 35, 1344–1346.
Mehlenbacher, S.A. and Scorza, R. (1986) Inheritance of growth habit in progenies of ‘Com-Pact Redhaven’
peach. HortScience 21, 124–126.
Monet, R. (1967) Contribution à l’étude génétique du pêcher. Annales de l’Amélioration des Plantes 17, 5–11.
Monet, R. (1979) Transmission génétique du caractère ‘fruit doux’ chez le pêcher. Incidence sur la sélection
pour la qualité. In: Proceeedings of Eucarpia Fruit Section Symposium. Tree Fruit Breeding. INRA, Angers,
France, pp. 273–276.
Monet, R. (1989) Peach genetics: past, present and future. Acta Horticulturae 254, 49–57.
Monet, R. (1995) Il miglioramento genetico del pesco. In: Bellini, E. (ed.) Proceeding of the International
Symposium ‘State of the Art and Perspectives of World Genetic Improvement of Fruit Tree Species’.
ERSO, Faenza, Italy, pp. 13–27.
Monet, R. and Bastard, Y. (1972) Contribution à l’étude du contrôle génétique de quelques caractères
morphologiques chez le pêcher. Annales de l’Amélioration des Plantes 22, 399–403.
Monet, R. and Bastard, Y. (1982) Une anomalie du fonctionnement de l’apex à hérédité mendélienne chez le
pêcher (Prunus persica (L.) Batsch). Agronomie 2, 103–106.
Monet, R. and Bastard, Y. (1983) Nouveaux cas de ségrégation indépendante de caractères mendéliens chez
le pêcher. Agronomie 3, 387–390.
Monet, R. and Massonié, G. (1994) Déterminisme génétique de la résistance au puceron vert (Myzus persicae)
chez le pêcher. Résultats complémentaires. Agronomie 2, 177–182.
Monet, R. and Salesses, G. (1975) Un nouveau mutant de nanisme chez le pecher. Annales de l’Amélioration
des Plantes 25, 353–359.
Monet, R., Bastard, Y. and Gibault, B. (1985) Etude génétique et amélioration des pêches plates. Agronomie
5, 727–731.
Monet, R., Bastard, Y. and Gibault, B. (1988) Etude génétique du caractère ‘port pleureur’ chez le pêcher.
Agronomie 8, 127–132.
Monet, R., Guye, A. and Massonié, G. (1997) Breeding for resistance to green aphid, Myzus persicae Sulzer.
Acta Horticulturae 465, 171–175.
Nakasu, B., Bassols, M. and Feliciano, A. (1981) Temperate fruit breeding in Brazil. Fruit Varieties Journal 35,
114–122.
Okie, W.R. (1988a) USDA peach and nectarine breeding at Byron, Georgia. In: Childers, N. and Sherman, W.
(eds) The Peach. Horticultural Publications, Gainesville, Florida, pp. 51–56.
Okie, W. R. (1988b) Preliminary descriptions of five new peach genes. Acta Horticulturae 465, 107–110.
Okie, W.R. and Prince, V.E. (1982) Surface features of a novel peach × nectarine hybrid. HortScience 17,
66–67.
Pascal, T. (1990) Etude de la résistance à l’oïdium (Spherotheca pannosa) chez le pêcher (Prunus persica L.
Batsch). MSc. thesis, University of Montpellier, Montpellier, France.
Pascal, T., Levigneron, A., Kervella, J. and Nguyen-The, C. (1994) Evaluation of two screening methods for
resistance of apricot, plum and peach to Monilinia laxa. Euphytica 77, 19–23.
Pascal, T., Kervella, J., Pfeiffer, F.G., Sauge, M.H. and Esmenjaud, D. (1997) Evaluation of the interspecific
progeny Prunus persica cv. Summergrand × Prunus davidiana for disease resistance and some agro-
nomic features. Acta Horticulturae 465, 185–191.
Peace, C.P., Crisosto, C.H. and Gradziel, T.M. (2005) Endopolygalacturonase: a candidate gene for freestone
and melting flesh in peach. Molecular Breeding 16, 21–31.
Pooler, M.R. and Scorza, R. (1995) Occurrence of viable eggs in haploid peach. Fruit Varieties Journal 49,
239–241.
Ramming, D.W. (1990) The use of embryo culture in fruit breeding. HortScience 25, 393–398.
Ramming, D.W. (1991) Genetic control of a slow-ripening fruit trait in nectarine. Canadian Journal of Plant
Science 71, 601–603.
Rodriguez, A.J., Sherman, W.B., Scorza, R. and Wisniewski, M. (1994) ‘Evergreen’ peach, its inheritance and
dormant behavior. Journal of the American Society for Horticultural Science 119, 789–792.
Salesses, G. and Al-Kai, N. (1985) Simply inherited grafting incompatibility in peach. Acta Horticulturae 173,
57–62.
Sansavini, S., Bassi, D. and Gamberini, A. (2006) Peach breeding, genetic and new cultivar trends. Acta Hor-
ticulturae 713, 23–52.
Scorza, R. (1992) Evaluation of foreign peach and nectarine introductions in the US for the resistance of leaf
curl (Taphrina deformans Berck. Tul.). Fruit Varieties Journal 46, 141–145.
84 R. Monet and D. Bassi

Scorza, R. and Pooler, M. (1993) Development and testing of F1 hybrid peaches as an alternative peach produc-
tion strategy. HortScience 28, 455.
Scorza, R. and Sherman, W.B. (1996) Peaches. In: Janick, J. and Moore, J.N. (eds). Fruit Breeding. Vol. I. Tree
and Tropical Fruits. Wiley, New York, pp. 325–440.
Scorza, R., Lightner, G.W. and Liverani, A. (1989) The Pillar peach tree and growth habit analysis of
compact × Pillar progeny. Journal of the American Society for Horticultural Science 114, 991–995.
Scorza, R., Bassi, D. and Liverani, A. (2002) Genetic interaction of pillar (columnar), compact and dwarf
peach tree genotypes. Journal of the American Society for Horticultural Science 127, 254–261.
Scott, D. and Cullinan, F. (1942) The inheritance of wavy leaf character in the peach. Journal of Heredity 33,
293–295.
Scott, D.H. and Weinberger, J.H. (1944) Inheritance of pollen sterility in some peach varieties. Proceedings of
the American Society for Horticultural Science 45, 229–232.
Sharpe, R.H., Hesse, C.O., Lownsberry, B.F., Perry, V.G. and Hansen, C.J. (1970) Breeding peaches for root-knot
nematode resistance. Journal of the American Society for Horticultural Science 94, 209–212.
Sherman, W.B. and Lyrene, P.M. (2001) ‘UFO’, a saucer or donut peach. Journal of the American Pomological
Society 55, 2–3.
Sherman, W.B. and Rodriguez-Alcazar, J. (1987) Breeding of low chill peach and nectarines for mild winters.
HortScience 22, 1233–1236.
Toyama, T.K. (1974) Haploidy in peach. HortScience 9, 187–188.
Weinberger, J.H., Math, P.C. and Scott, D.H. (1943) Inheritance study of root-knot nematode resistance in
certain peach varieties. Proceedings of the American Society for Horticultural Science 42, 321–325.
Werner, D.J. and Chaparro, J.X. (2005) Genetic interaction of pillar and weeping peach genotypes. Hort-
Science 40, 18–20.
Werner, D.J. and Creller, M.A. (1997) Inheritance of sweet kernel and male sterility. Journal of the American
Society for Horticultural Science 122, 215–217.
Werner, D.J., Creller, M.A. and Chaparro, J.X. (1998) Inheritance of the blood-flesh trait in peach. HortScience
33, 1243–1246.
Werner, D., Ritchie, D., Cain, D. and Zehr, E. (1986) Susceptibility of peaches and nectarines, plant introduc-
tions, and other Prunus species to bacterial leaf spot. HortScience 21, 127–130.
Yamazaki, K., Okabe, M. and Takahashi, E. (1987) Inheritance of some characteristics and breeding of new
hybrids in flowering peaches. Bulletin of Kanagawa Horticultural Experimental Station 34, 46–53.
Yoshida, M. (1976) Genetical studies on the fruit quality of peach varieties. III. Texture and keeping quality.
Bulletin of the Fruit Tree Research Station, Series A 3, 1–16.
 4 Genetic Engineering and Genomics

A.G. Abbott,1 P. Arús2 and R. Scorza3

1Clemson University, Clemson, South Carolina, USA
2Institut de Recerca i Tecnologia Agroalimentàries (IRTA), Cabrils, Spain
3USDA-AFRS, Kearneysville, West Virginia, USA

4.1 Peach Molecular Genetics 85

Brief description of peach genetics 85
Application of molecular genetic mapping approaches to peach genetics 86
Comparative mapping of peach and other Prunus species 88
Comparative mapping of peach to Arabidopsis 92
4.2 Peach Genomics and Gene Discovery 92
Construction of the peach physical map and its use in gene discovery 93
Summary of current peach expressed sequence tags projects and data 94
Comparative physical mapping of peach and other model genome species 96
The peach genome database and gene discovery 97
Rosaceae genome database and peach as a model genome species 97
4.3 Transformation in Peach 97
4.4 Conclusions and Perspectives 100

4.1 Peach Molecular Genetics detection of each chromosome individually

Brief description of peach genetics
The cultivated peach is a diploid species (2n =
2x = 16) that belongs to the Rosaceae family,
subfamily Prunoideae, genus Prunus and sub-
genus Amygdalus. The peach karyotype has
been studied during meiosis (Jelenkovic and
Harrington, 1972) and mitosis (Salesses and
Mouras, 1977) and consists of a clearly identi-
fiable, large submetacentric chromosome and
seven more chromosomes of smaller size, two
of them acrocentric. Recent results with fluo-
rescence in situ hybridization in the closely
related almond (Prunus dulcis) have enabled
based on chromosome length and the posi-
tions of the rDNA genes (Corredor et al.,
2004). The size of the Prunus genome is one of
the smallest among cultivated species, with
an estimated length of 290 Mbp in peach (Baird
et al., 1994), approximately twice the size of
Arabidopsis.
Most of the crosses between peach and
species included in the subgenus Amygdalus
are possible and produce fertile hybrids, like
those obtained with the peach-like Prunus
ferganensis, Prunus mira, Prunus davidiana and
Prunus kansuensis, or the cultivated almond.
Crosses with species of other subgenera (Pruno-
phora and Cerasus) such as apricot (Prunus

© CAB International 2008. The Peach: Botany, Production and Uses

(eds D.R. Layne and D. Bassi) 85
86 A.G. Abbott et al.
armeniaca), Myrobalan plum (Prunus cerasifera),
European plum (Prunus domestica), Japanese
plum (Prunus salicina) or sour cherry (Prunus
cerasus) are also possible, but fertile hybrids
are produced only occasionally (Scorza and
Sherman, 1996). In all, these results suggest
that there is an enormous gene pool available
for the development of improved cultivars,
although this variability has so far been
underexploited.
One of the distinctive characteristics of
peach is its self-compatible mating behaviour,
unlike the majority of its congeneric species,
which have a gametophytic self-incompati-
bility system. Selfing (Miller et al., 1989), plus
important bottlenecks in its recent breeding
history (Scorza et al., 1985), have resulted in a
lower level of genetic variability in peach
compared with the other Prunus crops (Byrne,
1990). The high economic interest of peach, its
self-compatible nature that allows the devel-
opment of F2 progenies and the possibility of
shortening the juvenile period to 1–2 years
after planting (Scorza and Sherman, 1996)
make peach a more adequate organism for
genetic analysis than other Prunus crops,
resulting in more detailed studies. A total of
42 morphological characters of simple Men-
delian inheritance were discovered during
the last century (Dirlewanger and Arús, 2008)
but, until the recent development of molecular
marker maps, only a few linkage relationships
among them could be established. Five link-
age groups involving 11 major genes were
reported by Monet et al. (1996).
Application of molecular genetic mapping
approaches to peach genetics
The first map constructed in fruit trees was
reported by Chaparro et al. (1994) using 83
random amplified polymorphic DNA (RAPD)
markers, one isozyme and four morphologi-
cal characters in a peach intraspecific F2 prog-
eny. Two more maps based on restriction
fragment length polymorphism (RFLP) mark-
ers were published shortly thereafter; the first
constructed in a peach × peach F2 progeny
(Rajapakse et al., 1995) and the second in a
peach × almond F2 progeny (Foolad et al.,
1995). The peach maps that followed inte-
grated dominant RAPD and amplified frag-
ment length polymorphism (AFLP) markers
with co-dominant (RFLP) and morphological
markers (Dirlewanger et al., 1998) or were
constructed almost entirely with AFLP mark-
ers (Lu et al., 1998). These maps can be consid-
ered incomplete, as they generally detected a
number of linkage groups different from the
eight expected, had a low average marker
density (4.5–8.5 cM/marker), included large
gaps without markers and many of the mark-
ers identified (8–28%) were unlinked.
The first saturated linkage map, con-
structed exclusively with transferable mark-
ers (11 isozymes and 226 RFLPs, most of them
detected with Rosaceae DNA probes) in a
‘Texas’ almond × ‘Earlygold’ peach F2 popu-
lation, was published by a European consor-
tium (Joobeur et al., 1998). All these markers
were distributed into eight linkage groups
with a total distance of 491 cM, representing
an average density of 2.0 cM/marker, and
with all gaps between markers shorter than
12 cM. This map (abbreviated as the T × E
map) has recently been improved by the addi-
tion of 176 simple sequence repeat (SSR)
markers (Aranzana et al., 2003; Dirlewanger
et al., 2004) and 123 RFLPs (Dominguez et al.,
2003), most of them obtained with Arabidopsis
DNA probes. From the 536 markers currently
placed on the T × E map, 466 (87%) are based
on known, publicly available DNA sequences,
with 198 (37%) of these sequences corre-
sponding to a putative protein.
The Prunus scientific community has
adopted the T × E map as the reference map
for the genus. It provides a set of transferable
markers that can be used as anchors for map
construction in other progenies, a common
linkage group terminology and marker order
within each linkage group, and a highly poly-
morphic population that allows mapping
markers that would not segregate in most
peach intraspecific crosses. Several inter- or
intraspecific peach maps anchored with the
T × E map have been obtained and their char-
acteristics are summarized in Table 4.1.
The network of maps interconnected
with T × E constitutes a resource from which
many markers can be found to saturate spe-
cific genomic regions of any progeny or to
Table 4.1. Peach linkage maps and anchor markers with the ‘Texas’ × ‘Earlygold’ (T × E) reference map.

No. of Anchors No. of linkage

Population Species Type markers with T × E groups Referencea

‘Texas’ × ‘Earlygold’ Almond × peach F2 536 536 8 Joobeur et al. (1998); Aranzana et al.

Genetic Engineering and Genomics

(2003); Dominguez et al. (2003);
Dirlewanger et al. (2004)
NC174RL × ‘Pillar’ Peach F2 84 0 15 Chaparro et al. (1994)
‘N.J. Pillar’ × KV77119 Peach F2 47 2 8 Rajapakse et al. (1995)
‘Padre’ × ‘54P455’ Almond × peach F2 161 38 8 Foolad et al. (1995); Bliss et al. (2002)
‘Ferjalou Jalousia®’ × ‘Fantasia’ Peach F2 69 48 8 Dirlewanger et al. (1998);
Etienne et al. (2002)
‘Lovell’ × ‘Nemared’ Peach F2 153 0 15 Lu et al. (1998)
‘Garfi’ × ‘Nemared’ Almond × peach F2 51 50 7b Jáuregui et al. (2001)
IF731 × Prunus ferganensis Peach × P. ferganensis BC1 141 32 10 Dettori et al. (2001)
‘Akame’ × ‘Juseitou’ Peach F2 120 56 7b Yamamoto et al. (2001, 2005)
‘Summergrand’ × P1908 Peach × Prunus F2 153 62 8 Foulongne et al. (2003a)
davidiana

aWhere more than one reference is given the data presented are either from the most recent publication or from the combination of the data from all publications.
bLinkage groups 6 and 8 of these maps were mapped as a single group due to the effects of a reciprocal translocation.

87
88 A.G. Abbott et al.
search for markers covering the whole gen-
ome adequate for quantitative trait locus
(QTL) or other genetic analyses. Given that
peach has a low level of intraspecific variation,
a very dense ‘consensus’ map with highly
polymorphic markers well distributed in all
genomic regions is necessary to ensure that
segregating markers are found where needed
in the population of interest. To reach this
goal, a supplementary effort will be required
to increase the number of SSRs mapped
(Aranzana et al., 2003) in parallel with tar-
geted strategies to fill regions with low SSR
density (Wang et al., 2001, 2002; Georgi et al.,
2002).
The existence of a single reference map
has made it possible to locate the major genes
and QTLs that segregated in different popula-
tions (Table 4.2). In total, 21 major genes have
been assigned to specific positions on the
T × E map, 18 of these major genes described
in peach plus three more affecting flower or
kernel characteristics that segregated in
almond × peach crosses. Joint analysis between
markers and quantitatively inherited charac-
ters has also been undertaken for bloom and
maturity date, fruit quality, tree architecture
and disease resistance (Abbott et al., 1998;
Viruel et al., 1998; Dirlewanger et al., 1999;
Etienne et al., 2002; Verde et al., 2002; Fou-
longne et al., 2003b). In addition, QTLs with
major effects or consistently found in differ-
ent years have been detected for all of the
above characters, enabling the approximate
positioning of 28 QTLs on the T × E map.
Most of the major genes located on the
Prunus general map can be selected with
neighbouring markers. Other strategies for
gene tagging that do not require knowledge
of the map position, such as bulked segregant
analysis (Michelmore et al., 1991), have also
been used successfully in peach (Chaparro et al.,
1994; Warburton et al., 1996; Lu et al., 1998). In
spite of this information being available, the
use of markers for commercial breeding is
still in its infancy. Marker-assisted selection is
currently used in a rootstock breeding pro-
gramme to pyramid a root-knot nematode
(Meloidogyne spp.) resistance gene coming
from ‘Nemared’ peach (Lu et al., 1998; Yama-
moto and Hayashi, 2002; Arús et al., 2004)
with another independent root-knot nema-
tode resistance gene coming from Myrobalan
plum (Claverie et al., 2004). However, selections
using markers of other well-characterized
genes affecting fruit characters (i.e. flesh colour,
skin pubescence, fruit shape, fruit sweetness)
have not been reported. This is undoubtedly
due to the fact that the variability of major
traits of interest to breeders (i.e. ripening time,
fruit quality and other characters) is quantita-
tively inherited. There is information on some
of these traits (Dirlewanger et al., 1999; Eti-
enne et al., 2002), but a more detailed knowl-
edge of the number, effects and map positions
of the QTLs affecting them is necessary before
QTL-associated markers can be routinely
integrated in selection programmes.
Additional candidates for marker-assisted
selection in peach are genes or QTLs that can
be introgressed into peach from other wild or
cultivated species, such as disease or pest
resistance. Examples include resistance to
sharka (Plum pox virus) from apricot, mapped
by Vilanova et al. (2003) in linkage group 1 of
Prunus, and other disease resistance QTLs
(mildew, leaf curl, aphids, sharka) from P.
davidiana identified by Viruel et al. (1998) and
Foulongne (2002). Introgression from wild
species is facilitated with markers using whole
genome selection approaches (Tanksley et al.,
1989) that reduce the number of generations
needed for recovery of the genome of the cul-
tivated species or elite genotype.
Comparative mapping of peach and other
Prunus species
The transferable markers (RFLPs, SSRs and
isozymes) mapped in the T × E population
have been used for the construction of link-
age maps in other Prunus species. Detailed
comparisons can be made between this map
and those of almond (Joobeur et al., 2000),
apricot (Lambert et al., 2004), P. davidiana
(Foulongne et al., 2003a), cherry (Dirlewanger
et al., 2003) and P. cerasifera (E. Dirlewanger,
INRA Bordeaux, 2004, personal communica-
tion). The order and distribution of the mark-
ers into the eight linkage groups was generally
identical between species, suggesting a high
degree of synteny (see Fig. 4.1/Plate 46 below).
Occasional marker position discrepancies
Table 4.2. Description of the major genes (21) and quantitative trait loci (QTLs; 28) affecting morphological or agronomic characters in peach that can be
placed on the Prunus reference map.

Linkage
Character groupa Symbolb Population Reference

Flesh colour (white/yellow) G1 Y ‘Padre’ × ‘54P455’ Warburton et al. (1996); Bliss et al. (2002)
Evergrowing G1 Evg ‘Empress op op dwarf’ × PI442380 Wang et al. (2002)
Internode length G1 QTL (Prunus ferganensis × ‘IF310828’)BC1 Verde et al. (2002)
Powdery mildew resistance G1 QTL ‘Summergrand’ × P1908 Foulongne et al. (2003b)
‘Garfi’ × ‘Nemared’

Genetic Engineering and Genomics

Flower colour G1 B Jáuregui (1998)
Root-knot nematode G2 Mi c ‘P.2175’ × ‘GN22’, ‘Akame’ × ‘Juseitou’, Jáuregui (1998); Lu et al. (1998); Yamamoto
resistance ‘Lowell’ × ‘Nemared’, ‘Garfi’ × ‘Nemared’, et al. (2001); Bliss et al. (2002); Claverie et al.
‘Padre’ × ‘54P45’ (2004)
Ripening time, fruit skin G2 QTL (P. ferganensis × ‘IF310828’) BC1 Verde et al. (2002)
colour, soluble solids content
Double flower G2 Dl ‘NC174RL’ × ‘PI’ Chaparro et al. (1994)
Broomy (or pillar) growth habit G2 Br Various progenies Scorza et al. (2002)
Flesh colour around the stone G3 Cs ‘Akame’ × ‘Jusetou’ Yamamoto et al. (2001)
Anther colour (yellow/anthocyanic) G3 Ag ‘Texas’ × ‘Earlygold’ Joobeur (1998)
Leaf curl resistance G3 QTL ‘Summergrand’ × P1908 Viruel et al. (1998)
Fruit weight, fruit diameter, G3 QTL ‘Suncrest’ × ‘Bailey’ Abbott et al. (1998)
glucose content
Polycarpel G3 Pcp ‘Padre’ × ‘54P455’ Bliss et al. (2002)
Flower colour G3 Fc ‘Akame’ × ‘Jusetou’ Yamamoto et al. (2001)
Blooming time, ripening time, G4 QTL ‘Ferjalou Jalousia®’ × ‘Fantasia’, Etienne et al. (2002); Verde et al. (2002)
fruit development period (P. ferganensis × ‘IF310828’)BC1
Soluble solids content, fructose, G4 QTL ‘Ferjalou Jalousia®’ × ‘Fantasia’ Etienne et al. (2002)
glucose
Flesh adhesion (clingstone/ G4 F (P. ferganensis × ‘IF310828’) BC1, Dettori et al. (2001); Yamamoto et al. (2001);
freestone) ‘Akame’ × ‘Juseitou’ Verde et al. (2002)
Non-acid fruit G5 D ‘Ferjalou Jalousia®’ × ‘Fantasia’ Dirlewanger et al. (1998, 1999); Etienne et al.
(2002)

89
(Continued)
 90
Table 4.2. Continued

Linkage
Character groupa Symbolb Population Reference

Sucrose, malate, titrable acidity, G5 QTL ‘Ferjalou Jalousia®’ × ‘Fantasia’ Etienne et al. (2002)
pH, sucrose
Skin hairiness (nectarine/peach) G5 G ‘Ferjalou Jalousia®’ × ‘Fantasia’; Dirlewanger et al. (1998, 1999); Bliss et al.
‘Padre’ × ‘54P455’ (2002)
Kernel taste (bitter/sweet) G5 Sk ‘Padre’ × ‘54P455’ Bliss et al. (2002)
Ripening time, fruit skin G6 QTL (P. ferganensis × ‘IF310828’)BC1 Verde et al. (2002)
colour, soluble solids content

A.G. Abbott et al.

Plant height (normal/dwarf) G6 Dw ‘Akame’ × ‘Juseitou’ Yamamoto et al. (2001)
Leaf shape (narrow/wide) G6 Nl ‘Akame’ × ‘Juseitou’ Yamamoto et al. (2001)
Male sterility G6 Ps ‘Ferjalou Jalousia®’ × ‘Fantasia’ Dirlewanger et al. (1998)
Powdery mildew resistance G6 QTL ‘Summergrand’ × P1908 Foulongne et al. (2003b)
Leaf curl resistance G6 QTL ‘Summergrand’ × P1908 Viruel et al. (1998)
Fruit shape (flat/round) G6 S* ‘Ferjalou Jalousia®’ × ‘Fantasia’ Dirlewanger et al. (1998, 1999)
Leaf colour (red/yellow) G6–G8 Gr ‘Garfi’ × ‘Nemared’, ‘Akame’ × ‘Juseitou’ Jáuregui (1998); Yamamoto et al. (2001)
Fruit skin colour G6–G8 Sc ‘Akame’ × ‘Juseitou’ Yamamoto et al. (2001)
Leaf gland (reniform/globose/ G7 E (P. ferganensis × ‘IF310828’)BC1 Dettori et al. (2001)
eglandular)
Resistance to mildew G7 QTL (P. ferganensis × ‘IF310828’)BC1 Verde et al. (2002)
Powdery mildew resistance G8 QTL ‘Summergrand’ × P1908 Foulongne et al. (2003b)
Quinase G8 QTL ‘Ferjalou Jalousia®’ × ‘Fantasia’ Etienne et al. (2002)
aG6–G8 genes located close to the translocation breakpoint between these two linkage groups.
bQTLs are included if they have been consistently found (at least in two independent measurements) in the indicated populations.
cOne or two genes of nematode resistance with different notations and one QTL with ? have been described in this linkage group.
 Genetic Engineering and Genomics 91

Chromosome 1

Contigs Frameworks

2.5: AG53
–1 –2 4.3: AC24, FG81, FG83
5.1: AG109
–3 6.8: EPPCU3062
–4 8.7: AG102
–5 10.8: HDL
–6 12.6: AG51
16.2: AG116

–7 –8 25.8: AC32, AC47

27.9: AG29

–9 35.7: FG28

11 –10 40.5: FG5, PC7

13 –12 41.3: AC7, AG113
42.8: EPPCU1589, EPPCU3088, pchgme56
48.0: AG105
–14 49.6: AG44
49.8: TSA2

–15 –16 65.1: FG36, FG228

65.7: AC18, AC23
72.9: PC32A
77.4: BPPCT028

–17 87.0: AG36

[17 contigs] [31 markers]

FPC file date: 00:02 Wed 23 May 2007

Fig. 4.1. Initial physical map of linkage group 1 of peach depicted in the Prunus genome web site
(http://www.bioinfo.wsu.edu/gdr/, accessed August 2007).

among species maps are attributed to the
mapping of different duplicated loci detected
by the same RFLP probe or SSR primer pair.
These results strongly indicate that the group
of Prunus species studied to date shares a
nearly identical genome. Therefore, the infor-
mation on gene sequence and position obtained
in one Prunus species would be generally use-
ful for the rest.
An exception to the full collinearity obser-
ved within Prunus was reported by Jáuregui
et al. (2001), who demonstrated the presence
of a reciprocal translocation between linkage
groups 6 and 8 in an F2 progeny of ‘Garfi’
almond × ‘Nemared’ peach, and established
the approximate position of the translocation
breakpoint. Although it was not possible to
determine the chromosomal configuration of
the parents of this cross, the fact that previous
maps involving peach, almond or both species
never detected the translocation suggested
that this mutation occurred only in part of the
germplasm of either peach or almond. The
same translocation was found in the F2 prog-
eny derived from a cross of the peach cultivars
‘Akame’ and ‘Juseitou’ (Yamamoto et al., 2001,
92 A.G. Abbott et al.
2005), suggesting that it occurs within peach.
Since one of the parents of each cross, ‘Nem-
ared’ and ‘Juseitou’, is a red-leaved cultivar,
and the translocation breakpoint is located in
the same chromosomal region as the gene
that determines red versus green leaf colour
(Gr/gr), perhaps there is a relationship
between the translocation and the leaf colour
trait. This observation is worthy of further
study.
Comparative mapping of peach to
Arabidopsis
The Prunus map and the Arabidopsis thaliana
genome sequence have been compared using
a set of RFLP markers mapped in T × E obtained
either with probes of different species (mainly
Prunus and apple) that had a high level of
sequence conservation with Arabidopsis
(TBLASTX values lower than 10−15) or with
Arabidopsis probes that hybridized well to
Prunus DNA (Dominguez et al., 2003). The posi-
tion of 227 Prunus loci (map average density
of 2.6 cM/marker) could be compared with
that of 703 Arabidopsis homologous sequen-
ces. The criterion for declaring a syntenic
region was that three or more homologous
markers had to be located within 1% of the
Prunus map distance (6 cM) and within 1%
of the Arabidopsis genome (1.2 Mb). In addi-
tion, blocks with gaps longer than 1% of
either genome were rejected. With these
stringent criteria it was possible to detect 37
syntenic regions, covering 23% and 17% of
the Prunus and Arabidopsis genomes, respec-
tively. The longest of these regions included
13 markers for a distance of 25 cM in linkage
group 2 of Prunus and 16 homologous
sequences spanning 5.4 Mb in chromosome
5 of Arabidopsis.
Dominguez et al. (2003) used the same
approach to compare Arabidopsis with the
genomes of other dicotyledonous species
belonging to three different families (sugar-
beet, sunflower and potato). These results
indicated a similar level of synteny to that
found between Prunus and Arabidopsis. Although
map comparisons between the four crop spe-
cies were not directly possible, common syn-
tenic regions with Arabidopsis were frequent:
20 of the 37 syntenic Prunus–Arabidopsis
blocks were also common to two or three of
the other species, indicating that synteny
between these species is important. Interest-
ingly, three regions of Arabidopsis (in chromo-
somes 1, 2 and 3) were syntenic to all species
studied, suggesting that they are of evolu-
tionary significance.
The sequence of peach bacterial artificial
chromosomes (BACs) and BAC ends located
in the proximity of the region around the ever-
growing gene in linkage group 1 was com-
pared with that of Arabidopsis by Georgi et al.
(2003). Predicted genes in these sequences were
homologous to genes scattered along the five
chromosomes of Arabidopsis, although some
of them were placed in close positions in both
genomes. Macro- and microsynteny results
coincide in detecting a fragmentary but still
existing synteny between these two genomes.
This indicates that the sequence of Arabidopsis
is a resource that may be useful for character-
izing specific regions of the Prunus genome to
search for genes of interest or markers for
their selection.
4.2 Peach Genomics and Gene Discovery
Although Prunus is an economically and bio-
logically important genus, little was known
about the genome structure and organization
of its members up until the advent of DNA
marker technologies. With respect to the
application of DNA marker technologies to
the problem of developing genetic resources
in trees, peach has distinct advantages that
make it suitable as a model species for struc-
tural and functional genomics studies. It is
diploid with a small genome and a large
number of genes controlling fundamentally
important traits that have been genetically
described (Moore and Janick, 1975; Mowrey
et al., 1990). These include genes controlling
flower development, fruit development, tree
growth habit, dormancy, cold hardiness, and
disease and pest resistance. As described
above, extensive and detailed molecular genetic
mapping efforts have positioned many of
these traits (both single gene and QTL) on
saturated genetic linkage maps.
 Genetic Engineering and Genomics
In order to bridge the gap between the
map position of important traits and candidate
gene identification, the International Rosa-
ceae Genome Consortium (IRGC) was created
to promote the development of the peach as an
index genome for the identification, charac-
terization and cloning of important rosaceous
species genes using structural and functional
genomics technologies. The current status of
this programme is outlined below.
Construction of the peach physical map and
its use in gene discovery
Large-insert libraries and physical maps are
important tools for map-based cloning of
Mendelian loci (Arondel et al., 1992) and QTLs
(Frary et al., 2000). In cooperation with the
Clemson University Genomics Institute, sep-
arate peach BAC libraries were constructed
for ‘Nemared’ rootstock and a haploid of
‘Lovell’. The restriction enzymes used were
HindIII and Sau3A1, respectively. The ‘Nem-
ared’ library consists of approximately 40,000
clones with average inserts approximately
60 kb in size. The theoretical coverage of the
genome is eight- to tenfold but in practice it is
approximately four- to fivefold. The haploid
‘Lovell’ library consists of approximately
35,000 clones with an approximate average
insert size around 80 kb, yielding a theoretical
tenfold coverage of the genome.
Utilizing these BAC library resources the
IRGC is constructing a complete physical
map of the peach genome anchored on the
general Prunus genetic map (Joobeur et al.,
1998), essentially following strategies utilized
to develop the Drosophila physical map and
others (Marra et al., 1999; Hoskins et al., 2000;
Tao et al., 2001; Cone et al., 2002). The approach
utilizes a combination of hybridizing mapped
markers, BAC fingerprinting and in our case
hybridizing expressed sequence tag (EST)
sequences. With the current Prunus molecular
marker map resources, 210 low-copy mapped
RFLP markers, 3700 peach fruit ESTs, 80 resis-
tance gene analogues, 200 specific cDNAs
and numerous specific AFLP markers have
been hybridized to the BAC libraries. We
completed BAC fingerprinting approximately
93
20,000 BACs (15,000 from the ‘Nemared’
library and 5000 from the haploid ‘Lovell’
library), from which approximately 15,000
have been used to construct an initial physi-
cal map (see map specifics in Table 4.3 and
Fig. 4.1/Plate 46).
Physical mapping software (FPC (Fin-
gerPrinted Contigs) version 4.7; Soderlund
et al., 2000) was used to construct an initial
physical map of the peach genome following
strategies employed by others (Marra et al.,
1999, Tao et al., 2001) to construct physical
maps in other crops. Initially, the map was
constructed at a cut-off from e−10 to e−12 and
tolerance of 5 to obtain all high-confidence
overlapping BAC inserts (contigs). These
were then merged by testing end clones at a
cut-off value ranging from e−8 to e−11. Since
there was a significant amount of hybridiza-
tion data, merges were often achieved based
on common hybridization of BACs in differ-
ent contigs. However, if only BAC fingerprint
data existed, we noted the merge points for
further testing. Presently, the framework map
is composed of about 1000 contigs containing
approximately 8000 clones (see Table 4.3 and
Fig. 4.1). Based on estimates of an average
BAC insert size of 60 kb and an average of
60% degree of overlap in contigs, 80% or bet-
ter of the peach genome should be in high-
confidence contigs. Currently adding in
orphan singleton BACs (approximately 7000,
not in contigs from initial map construction)
and merging contigs at lower cut-off scores is
under way to finalize the initial peach physi-
cal map. Preliminary estimates from trial
merges of contigs suggest that the initial map
will consist of 800–900 contigs with an aver-
age of 12 clones per contig upon completion
of the analysis. Since the map includes marker
hybridization data from the general Prunus
genetic map, the developing physical map is
directly anchored to the genetic map. From
initial analysis of the integrated genetic/
physical map, there is already evidence for
duplication of some regions of the peach
genome. The developing physical map is
located at the Prunus genome web site within
the Genome Database for Rosaceae (GDR)
(www.bioinfo.wsu.edu/gdr). This database
is under ongoing development (for details see
below).
94 A.G. Abbott et al.

Table 4.3. Peach physical map data.

BACs fingerprinting information

No. of bands per fingerprint

Class Contig Single Avg.a <5 <15 <25 <35 <45 <55 ≥55

BAC 8360 6419 18.6 0.0b 33.5 45.9 16.9 3.2 0.4 0.1

Contig information

Size distribution

max >200 200:100 100:50 50:25 25:10 9:3 2 Total

Contig 296 1 0 0 8 232 726 303 1270

Anchored 43 0 0 0 1 74 91 0 176

Marker information

No. of contigs No. of clones

Class Total Avg. 0 1 2 >2 Avg. 1 <5 <10 ≥10

Probe 863 1.2 207 429 129 98 2.9 322 389 124 28

BAC, bacterial artificial chromosome.

aAverage number of bands analysed per fingerprint.
bPercentage of clones containing the specified number of fragments.

Summary of current peach expressed
sequence tag projects and data
Peach expressed sequence tag functional
genomics database development
With the support of the US Department of
Agriculture (USDA), the IRGC initiated a
peach EST project with the central goal of
developing the unique expressed gene set
(unigene set) for peach. The current efforts
are centred on sequencing 30,000–40,000 cDNAs
from libraries of developing fruit, shoot and
seeds. Original expectations were that these
would resolve into 3000–4000 unigenes; how-
ever, this number was obtained from the first
15,000 sequences finished. The data summary
for the completed analysis of 15,000 cDNAs
from developing peach fruit and almond seed
libraries is available at www.bioinfo.wsu.
edu/gdr/. Sequencing of developing shoot
and root cDNAs is in progress.
We have also begun mapping peach ESTs
on the developing physical/genetic peach
map and have determined that a significant
portion of ESTs (11%) hybridized on our BAC
libraries are placed directly on genetically
mapped anchored contigs in the physical map
(Fig. 4.2/Plate 47 and Table 4.3). From the cur-
rent 15,000 sequences, a peach/almond unigene
set has been initiated. This unigene set consists
of 3842 putative unique genes.
Transcript map
One hundred and eighty ESTs (11%) have
been localized in 86 locations (involving 80
core markers) on the general Prunus genetic
map by common hybridization with RFLP
markers to BACs in the ‘Nemared’ library.
This EST resource will provide candidate
genes for marked regions of the Prunus maps
containing traits of interest and will be avail-
able online through the Prunus genome data-
base noted before. From the initial fruit
unigene set, we have completed hybridizing
in excess of 1700 ESTs on to the ‘Nemared’
BAC library. From this set, 184 ESTs have
been directly located on the general Prunus
genome map through common hybridization
 Peach – Transcript map (180 ESTs, 86 locations – 12-01-03)
G1 G2 1 G3 G4 38 G5 3 G6 G7 1 G8
1 1
2
0 MC013 0 AC33A+ 4 0 CC127, MC115, 1 0 AC55A 2 0 FG26A 4 1
2 AG53 1
1
2 2 AC31+, AC10+, 4 4 2
MC028A,
Idh-2, AG56 1
0
1
TSA4
CC135C
2
1 0
1
AC44
AC52A, 1 } 1 0 AG2A 5
2 PC104A
FG81A, 4 5 3 AG101A 6
2 AG43A+, AC37A+
14 3 3 AG54, AG13, 3 1
5 FG83 3 5 AC13+ 3 3 5 PLG35 1 FG54A
PLG26C 4 Ext1 1
6 AC24
AG109
5 } 2 6 FG13
5 7 AG8B 9
5
FG215, AG40 1 4
AC43B+ 17 PLG59 , CC11A 7 AG112A 1
4 8 1
1 9 CC47+, AC41A 9 9 PLG26B 2
10 AG102 5 10 AG7, Me-1 1 4
4
1 11 CC129 8 38 3 10 MC023B
12 Pij1+ 12 AG63B
4 1 13 4 5 11 2
13 Mdl1 2
5
13 AG21A+
FG10 (JxF) 3 } 1
14 PC70, CC124 14
AG21B, AC9
AC55B 2 AC42 (F) 7
13 FG84A, 2 13 LY29,
15
16
AG51 3
CC23, PC78 3
14 CC131B+ 7 15 AG32A 6
1 16 PLG17
15
AC49, AG114 2 3 15
1 AC50, 9 } 3 MC003B
B4G3 1 15
FG230A
CC131A
1
18
AC45 (T, F)
AG116A
15
18 AG20A+ 12
2
18 B4A9 9
AC53 (D) 4 } 2 FG1A 5 Ma1 (Myrobalan)
5 19 6 18 PC12 1 2 18 AC22A 3
19 PLG39A
20
AG35+, AG107A+
20 CC116, LY5A 20 AG25A 12 5 2
20 PC102 PC5+ AG58 (T,F)
22 FG220+ 2 22 Tip1 8 21 PLG39D FG49 (PxF) 2 } 1
1
1
23 AG52A+, PC9A+ 3 23 AG57A+ 2 1 1 23 PLG2 22 PC101+
1
1 24 PC14 2 23 CC63, MC225 6 23 AG4A+, 11
25 PC26 1 25 CC2+
26 AG6, FG205, 2 7 6Pgd-1
25 AG104 4 25
TubA2 10
26 AC27A+, 2 26 CC69+ 26 Pyk1
27 AG32C 6 MC030A 1
CC135B 1
7
} 5 CC3A

Genetic Engineering and Genomics

28 AC32 28
29 AC47, PC30 7 29 CC12A+, AG110+ 1 3
1 30 MC007+, AG1A+ 3 30 CC59, AG62 5
29 AC8 1
30 Sdh-1
31 AG23 (T) 6
}2 7 31 PC34A
AG29A 30 2 32 PC67A +, CC8+ 1 3 4
AG25B (T,JxF) 12 }2 34 MC004A 34 B7A5A+, PC13+, 15
1
33
AG26A,
PC29A
3
5
32 FG37 4

35 CC6A 35 CC115+, AC19+, 4 2 35 Aco-2+, AG50A, 2 35 FG202, 4 35 AG39B

PC85, FG16, 1 2 CC11B 1
2 1
37 AG22A, AG36A 5
37 MC045, PC32B
2 37 AG106+, Pgd-2+ 3 4 37 FG3 2
1
AG46 1 FG53 (PxF, S)
} 1 36 AG60A
38
FG9A, LY5B, 4
AC21B+, Ole1+
5 } 2 39
FG38+
FG22A, AG45, 9 6 2 3
FG78 (PxF) 3
38 AC48A+ 2
2 40 CC56 40 CC133B
MC044, FG28 40 Aco-1,
40 Pgm-1+ 1 1
5 PLG39B, AC51 MC003A 41 FG119A
42 MC001, PC35A 41 FG201A+ 5
43 PC15
10
1 evg 43 CC138+,
9 3 1
44 PC7 AT/CTC2 Omt1+ CC135A, AG8A
AC7A, 3 (SCxB) 6 1 3 45 FG4 3 2 1
46 AG37 5 46 AC43A 18 46 TSA3+ 2
47 AG113, FG5 1 3 47 47 CC133A 3 47 AG61A, 47
48 Pgm-2 CC125+
4 3 11 AG49 2
AG108A 3 FG104 (PxF)
49 AG47 , AG30
50 PC6A+, FG6 50 AG33, 13
49 Prp1 9 AG10 (D, T, PxF) 2 } 1
51 LY21 MC011A PC21 8
3 2 51
FG14
51 FG42+
2 52 AC26 2
3 54 PC1, CC52 1
Pru1, 2
55
56
AG105A
AG44
7
8
} 7 55 FG27 5
54
AG14A, 5
57 MC024A 2 Adf1,
58 TSA2 58 AG12B, 12 57 AG63A 8

60 Lap-2+
7
AG16B 2 } 1
59 AG17A 4 59
MC043
PC36+
6
FG98, FG106 (PxF) 4 } 3 61 PC73
62 CC132
63 LY37+, MC022+ 3
1 1
65 Lap-1+, LY16A+ 65 PC28A 10
AC3, B6H11 (T)
2
3
} 1 1 3 67 PC60, 2
66 FG24 1

69 FG228+, FG36 PC105A

5 6
71 AC23+, AC18 1 1 6
73 Pgl1 5 1
1 74 Ltp2 1
77 AG8C, PC32A 9 2
6
78 FG8A
1 79 PLG39C, FG209A
18
CC110, PLG26A, 80 AG111A
81
TubA3 10
PCsl104 (PxA) 4
} 3

85 AG36B 9

Fig. 4.2. General Prunus genetic map (Joobeur et al., 1998) and developing peach physical/expressed sequence tag (EST) map. Markers depicted in bold
have been used to develop contigs of peach bacterial artificial chromosomes (BACs), which are depicted as light-coloured circles with the number of BACs
detected by each marker inscribed in the circle. Green circles denote two adjacent markers detecting the same BACs. Black flags depict peach and tomato

95
EST positions, with the number of ESTs at this location posted above the flags.
96 A.G. Abbott et al.
Table 4.4. Current peach EST database.
BACs Average no. of
EST probes identified BACs detected
1552 (PP_LE) 5920 3.8
68 (LF) 209 3.1
Total: 1620 6129 3.8
of mapped molecular markers and ESTs.
BACs have been identified in the ‘Nemared’
library for nearly 85% of these ESTs. Initial
hybridizations on the haploid ‘Lovell’ BAC
library of about 100 ESTs failing to detect
BACs in the ‘Nemared’ library have been 60%
successful. Thus, upon completion of the
physical map, virtually all unigene EST loca-
tions will be identified.
In cooperation with Dr V. Decroocq (INRA,
Bordeaux, France), we are also mapping resis-
tance gene analogues (RGAs) and resistance-
associated genes (RAGs). We have completed
hybridizing over 80 different RGAs/RAGs and
are currently positioning these on the physical
map. We have already positioned one RAG in
a contig that maps to a putative resistance gene
location for Plum pox virus (V. Decroocq and
D. Abernathy, unpublished results), demon-
strating the potential for the transcript map of
peach to provide candidate genes.
The structural and functional genomics
databases of peach serve as tools for microsyn-
teny analysis of regions of interest and for
gene cloning investigations. With the integra-
tion of sequenced cDNA loci (EST loci), the
physical map database immediately provides
candidate genes located in the genetically
marked intervals containing traits of interest.
These associations provide the potential to
greatly speed the process of gene discovery
and characterization.
Comparative physical mapping of peach and
other model genome species
One of the most important contributions of
DNA marker technology to fundamental stud-
ies in plant biology is the ability to rapidly
compare genome organization in closely
related as well as diverse species. Compara-
tive mapping studies can identify highly con-
served genome blocks and regions of lesser
conservation. Identification and molecular
dissection of these evolutionarily conserved
regions may uncover genetic associations
that, by virtue of their preservation, are impli-
cated as important for plant development. In
addition, comparative mapping information
can serve as a starting point for initial map-
ping and gene cloning investigations in poorly
characterized species.
The comparative genome sequence orga-
nization of plant genomes has not been exam-
ined as extensively as chromosomal mapping
level studies; however, some reports suggest
that within families there is a significant pres-
ervation of gene repertoire and order among
plants with quite different genome sizes (Dun-
ford et al., 1995; Bennetzen et al., 1996; Chen
et al., 1997; Kilian et al., 1997; Avramova et al.,
1998). Initial comparative sequencing studies
between Arabidopsis and rice have revealed
some conservation of genomic structure in
defined regions. The data suggest, however,
that genes are being dispersed into and out of
regions by mechanisms such as transposition,
thus obscuring microsynteny across great evo-
lutionary distances (van Dodeweerd et al.,
1999). Future research is necessary to examine
the degree of microsynteny within and among
plant families.
As discussed in the genetic mapping sec-
tion above, limited comparative mapping
between peach and other model genome spe-
cies was done utilizing molecular marker
technologies (Dominguez et al., 2003). This lack
of comparative data is also evident at the high-
resolution level; however, there are several
reports suggesting that specific regions of the
peach genome maintain a very limited micro-
synteny with the Arabidopsis genome (Georgi
et al., 2002). These initial studies demonstrate
that substantial genome rearrangements have
occurred, thus limiting the value of interfam-
ily comparative genomics as a tool for gene
discovery. However, within the Prunus genus,
the high level of genome preservation at the
low-resolution scale suggests that utilization
of the peach genome as an anchor for identifi-
cation of important genes in other species is
more promising. Initial high-resolution
comparative studies of peach with plum and
 Genetic Engineering and Genomics
apricot suggest that the peach genome data-
base will serve as an excellent source of can-
didate genes for traits in these species (M.
Badenes, IVIA, Valencia, Spain, 2004, personal
communication; D. Esmenjaud, INRA Antibes,
France, 2004, personal communication).
The peach genome database and
gene discovery
As peach genomic resources are generated,
utilization of these resources is now occur-
ring. Peach genome resources are being used
for identification of genes important to decid-
uous tree life history traits.
A non-dormant genotype (‘Evergrowing’,
USDA PI442380) of peach was previously
identified in southern Mexico, where killing
frosts do not occur. In Mexico, terminal growth
on ‘evergrowing’ trees is continuous under
favourable environmental conditions and
leaves are retained until they are lost due to
drought and/or disease (Diaz, 1974). At more
northern latitudes, the ‘Evergrowing’ peach
does not appear to respond to winter dormancy
cues, exhibiting persistence of shoot growth
and a lack of leaf abscission in response to short
days and low temperatures in the autumn until
these tissues are killed by freezing temperatures
(Rodriguez et al., 1994). A number of crosses
with evergreen (non-dormant) trees using dif-
ferent deciduous trees were made and all nine
of the F2 progenies fit a 3:1 (deciduous:non-
dormant) ratio, suggesting that the ‘Evergrow-
ing’ phenotype is controlled by a single recessive
gene (Rodriguez et al., 1994). Thus, ‘Evergrow-
ing’ trees provide a model system to examine
the genetic control of winter dormancy through
the cloning and functional analysis of a major
gene known to be required for dormancy.
In order to map the evergrowing gene locus
in peach, crosses were performed between the
peach cultivar ‘Empress’ and the non-dormant
peach selection PI442380 (Werner and Okie,
1998). Two F1 trees from this cross were self-
pollinated to produce an F2 population. F2
seeds were germinated in the greenhouse in
winter and transplanted to the field in June.
The growth characteristics of the progeny were
evaluated at the USDA-ARS Appalachian Fruit
Research Station in Kearneysville, WV. One
97
hundred and nine of 314 F2 trees exhibiting
consistent phenotypes in 1997 and 1998 were
randomly selected for the genetic mapping of
the recessive evg gene (Wang et al., 2002).
Bulk segregant AFLP analysis was per-
formed to identify AFLP markers linked to
the evg gene, and a local molecular linkage
map covering a total genetic distance of 79.3
cM was constructed (Wang et al., 2002). Four
flanking AFLP markers were cloned, sequenced
and converted into sequence tagged site (STS)
markers (Wang et al., 2002). A localized physi-
cal map of the evergrowing region was initiated
from the closest STS marker utilizing the peach
physical map resources. A chromosomal walk
in both directions was initiated from the BAC
PpN18F12 (Prunus persica ‘Nemared’ 18F12),
which contained the closest STS marker EAT/
MCAC. Identification of markers from the
contig developed from this walk suggested
that the evergrowing region was contained
within a limited number of BACs, which were
then subjected to BAC sequencing (Bielen-
berg et al., 2004). Gene candidates were iden-
tified of the MADS box transcription factor
class. Southern hybridization analysis of this
region demonstrated a deletion spanning the
putative evergrowing region in the evg mutant.
Candidate genes in this region and their
homologues in other tree systems (poplar,
plum) are currently under characterization.
Rosaceae genome database and peach as a
model genome species
We have taken a worldwide cooperative
approach to develop the peach as a reference
genome for fruiting trees. All the structural
and functional genomics resources are incor-
porated in the Genome Database for Rosaceae
(GDR) to serve as the repository for Rosaceae
genomics data worldwide. The Prunus genom-
ics data set, as well as data of other important
Rosaceae species, are included in the GDR
(http://www.bioinfo.wsu.edu/gdr).
4.3 Transformation in Peach
Peach (P. persica) and its smooth-skinned sport,
nectarine, is one of the most commercially
98 A.G. Abbott et al.
important stone fruit species (a group that
also includes apricot, cherry and plum).
World production of peach and nectarine is
about 17 million t (FAO, 2007). Biotic and abi-
otic stress factors such as pests, diseases,
drought and postharvest losses reduce stone
fruit production worldwide. New improved
cultivars have been released but many pro-
duction problems have yet to be solved by
conventional plant breeding (Scorza and
Sherman, 1996; Srinivasan and Scorza, 1999).
Genetic transformation is an alternative method
of stone fruit improvement that may be par-
ticularly useful to increase biotic and abiotic
stress resistance and fruit quality (Scorza,
1991, 2001; Scorza et al., 1995a; Srinivasan
et al., 2004). Plant genetic transformation gen-
erally involves the transfer of DNA with the
desired gene(s) into cells and the regenera-
tion of transgenic plants from the transformed
cells through in vitro culture.
While genetic transformation is an
important tool for peach improvement, a reli-
able and reproducible transformation and
regeneration system from somatic tissue has
yet to be developed. The following summa-
rizes the reports of work in peach transforma-
tion and regeneration.
Although induction of somatic embryo-
genesis has been reported for peach, conver-
sion of these somatic embryos into plants is
far from routine (Scorza, 2001). Raj Bhansali
et al. (1990) induced somatic embryos from
1–3 mm long immature zygotic embryos of
peaches and nectarines. To induce embryogenic
callus, the zygotic embryos were cultured on
Murashige and Skoog (MS) basal medium
(Murashige and Skoog, 1962) containing glu-
tamine, myo-inositol and casein hydrolysate
(each 500 mg/l), 2,4-dichlorophenoxyacetic
acid (2,4-D) (5 mg/l), kinetin (2 mg/l) and
6-benzylaminopurine (BAP) (2 mg/l). The
zygotic embryo cultures were initially incu-
bated for 10 days in the dark and then exposed
to continuous light for 20 days at 23°C. To
produce somatic embryos, these calli were
transferred on to MS basal medium contain-
ing glutamine, myo-inositol and casein hydro-
lysate (each 500 mg/l), MES (2-(N-morpholino)
ethanesulfonic acid) (976 mg/l) and activated
charcoal (2.5 g/l). No growth regulators were
added to the medium. Somatic embryos were
produced on this medium after three to five
transfers at 3-week intervals. Plant develop-
ment from embryos was on basal medium sup-
plemented with BAP (1.0 mg/l), a-naphthalene
acetic acid (NAA) (0.1 mg/l) and 0.03% acti-
vated charcoal. Efforts to produce long-term
embryogenic callus yielded embryos with
abnormal morphologies.
Guohua and Yu (2002) produced embryo-
genic callus from immature cotyledons of
four Chinese peach cultivars using a two-step
process. Culture on MS-based basal medium
supplemented with BAP (0.1 mg/l) and 2,4-D
(1.0 mg/l) for 6 weeks (transferred to fresh
medium after 3 weeks) was followed by cul-
ture on the same basal medium with the addi-
tion of BAP (0.01 mg/l) and 2,4-D (0.1 mg/l).
This procedure induced up to 95% of the
immature embryos to produce callus with up
to eight somatic embryos per explant. Up to
75% of these somatic embryos produced
shoots. While BAP has been generally used
for peach regeneration, Guohua and Yu (2002)
found thidiazuron (TDZ) (0.1–2.5 mg/l), in
combination with NAA (0.01 mg/l), to more
effectively induce adventitious shoots directly
from immature cotyledons of ‘Jingyan’ peach
when compared with BAP, kinetin, zeatin and
CPPU (N-(2-chloro-4-pyridinyl)-N′-phenylurea).
Scorza et al. (1990a) produced somatic
embryogenic cultures from immature (45–50
days post bloom) embryos. Following a
6-month culture period on the media of Ham-
merschlag et al. (1985), these cultures became
growth regulator-independent (habituated)
and continually produced somatic embryos
for up to 4 years. These embryogenic cultures
only rarely germinated to produce viable
shoots even when exposed to a number of
treatments including cold treatment and vari-
ous growth regulators.
Direct adventitious shoot regeneration
without intervening somatic embryo produc-
tion has been induced from callus derived
from immature zygotic peach embryos (Ham-
merschlag et al., 1985). To produce a white
nodular shoot regenerative callus, immature
zygotic embryos were excised from 70-day-
old ‘Sunhigh’ and ‘Suncrest’ peach fruits. These
were first cultured in MS basal medium sup-
plemented with 2.4 mM 2,4-D and 0.44 mM BAP
and then transferred to medium containing
 Genetic Engineering and Genomics
0.27 mM NAA and 2.2 mM BAP. Shoots were
regenerated from these calli after transferring
to a medium containing 2.2 mM BAP and 1.35
mM NAA. Regenerated shoots were rooted in
the dark on a medium containing 28.5 mM
indole-3-butyric acid (IBA). Direct shoot organo-
genesis has also been induced from immature
cotyledons excised from 70-day-old fruits of
peach cultivars ‘Bailey’, ‘Boone County’, ‘Sun-
crest’ and ‘Hann’ after 4–6 weeks’ culture on
MS medium containing 2.5 mM IBA and 7.5 mM
TDZ (Mante et al., 1989). Roots were produced
in 50–70% of the regenerated shoots cultured
under light on half-strength MS medium with
2.5 or 5 mM IBA.
The use of immature zygotic explants
limits source material availability to only a
few months out of the year. Pooler and Scorza
(1995) demonstrated adventitious shoot pro-
duction from mature cotyledons of peach
rootstock (‘Nemaguard’, ‘Flordaguard’ and
‘Nemared’) seeds that had been cold-stored at
4°C for 1–3 years. Cotyledons were cultured
in the dark on MS medium containing 1.25,
2.5, 6.25 or 12.5 mM IBA. The regenerated shoots
were maintained on medium containing 0.1
mM IBA and 1.0 mM BAP and shoots were
rooted in half-strength MS basal medium sup-
plemented with 5 mM IBA. This procedure
provides a relatively simple method for peach
regeneration that can be used year-round on
responsive genotypes. As with all peach
regeneration systems developed to date,
successful regeneration is highly genotype-
dependent.
Most of the preceding reports of regen-
eration from peach have focused on the use of
zygotic tissues, and most from immature
zygotic embryos. In contrast, Gentile et al.
(2002) reported adventitious shoot regenera-
tion from callus cultures of young leaves (1–2
mm long) from in vitro-grown peach shoots in
a medium containing 9 mM BAP and 0.54 mM
NAA. Regeneration rates of 13–28% were
obtained using three cultivars from diverse
origins and two seedling selections. Most
regeneration was obtained from leaf petioles.
Preconditioning the in vitro shoots that were
the sources of leaf explants in medium con-
taining 2 mg BAP/l and ≤0.2 mg NAA/l for
up to 4 months was a critical step in the regen-
eration process.
99
Clearly, it is possible to regenerate peach
plants in vitro. This has been achieved for the
most part by using zygotic tissues. These
explant sources have generally not been
favoured for tree fruit transformation because
the ability to improve established cultivars is
lost. Each seed-derived genotype is unique
and not a clone of the parent. Transformation
of zygotic tissues would be useful for provid-
ing unique and useful genes to breeding pro-
grammes where they could be incorporated
into new germplasm. Given the facts that: (i)
the generation cycle for peach is approxi-
mately 3 years (a short cycle compared with
most tree fruit species; Sherman and Lyrene,
1983); (ii) most new peach cultivars are pro-
duced by breeding programmes versus the
selection of sports of established cultivars;
and (iii) peach varieties are continually replaced
at a fairly rapid pace (10–12 years or less in
some areas), the efficient transformation of
peach germplasm can be of great benefit to
the genetic improvement of this species.
While the production of transgenic Prunus
currently depends largely on the efficiency of
regeneration of plants from transformed cells,
the efficiency of transformation itself is also
an important factor, one that takes on an even
greater level of importance in the case of low
regeneration rates. Several reviews have been
published on transformation of Prunus spe-
cies, including peach (Scorza and Hammer-
schlag, 1992; Scorza et al., 1995a; Rugini and
Gutierrez-Pesce, 1999; Srinivasan and Scorza,
1999; Srinivasan et al., 2004). Transformation
efficiency is affected by many factors including
the method of transformation (e.g. Agrobacte-
rium tumefaciens or biolistics), the transforma-
tion environment and the antibiotic selection
pressure. In most published reports, A. tume-
faciens has been used to transfer the DNA
plasmids carrying the gene(s) of interest to
peach cells. Neomycin phosphotransferase II
(NPTII) has been used as the selectable marker
and, in some cases, b-glucuronidase (GUS) as
a visual marker of transformation.
Although peach is infected by wild A.
tumefaciens and crown gall disease is common
in Prunus (Scorza and Sherman, 1996), the
transformation efficiency of peach cells in
vitro with disarmed A. tumefaciens is relatively
low. To date, there is only one report of the
100 A.G. Abbott et al.
development of transgenic peach plants
(Smigocki and Hammerschlag, 1991). In that
study, transgenic ‘Redhaven’ peach plants
expressing the A. tumefaciens ipt (isopentenyl
phosphotransferase) gene were regenerated
from immature zygotic embryogenic calli.
This ‘Redhaven’ peach was transformed with
a shooty mutant strain of A. tumefaciens,
tms328::Tn5, which carries an octopine type
Ti plasmid with a functional cytokinin gene
and a mutated auxin gene. The use of this
cytokinin-producing shooty-mutant strain of
A. tumefaciens may have been responsible for
the successful regeneration of transformed
shoots. The levels of endogenous cytokinins
in the few regenerated shoots that were pro-
duced were reported to be 50-fold higher than
the levels in untransformed controls. The
transgenic plants showed alterations in growth
habit compared with untransformed controls.
These included dwarf stature, increased
branching and delayed leaf senescence (Ham-
merschlag et al., 1997; Hammerschlag and
Smigocki, 1998). These growth characteristics
are indicative of high cytokinin levels and
were apparently due to the expression of the
ipt gene in the transgenic peach lines.
Scorza et al. (1990b) reported the trans-
formation of peach leaf segments, immature
embryos and long-term embryonic callus
using A. tumefaciens strain A281 carrying
plasmid pGA472 with the NPTII selectable
marker. Transformation rates of 5% of imma-
ture embryos and up to 64% of leaf segments
were observed. These explant sources did not
undergo organogensis, thus no transgenic
shoots were obtained from that work.
In contrast to A. tumefaciens-based trans-
formation studies, particle bombardment
(biolistics) has also been used to produce sta-
bly transformed embryogenic peach callus
(Ye et al., 1994). Embryogenic callus derived
from immature embryos was used as the
starting material. A high frequency of subcul-
ture (3 days to 2 weeks) prior to bombard-
ment was considered important in order to
maintain actively dividing cells, which tend
to be most receptive to biolistic transforma-
tion. From 114 callus lines, 65 were putatively
transformed. Seven lines were confirmed as
transformants by PCR and GUS histological
assays. No regeneration was obtained from
the transformed embryogenic callus pro-
duced in the study. Transient expression tests
of biolistic transformation of embryogenic
callus, embryonic axes, cotyledons, immature
embryos, leaf discs and shoot tips demon-
strated high levels of transformation efficiency.
However, shoot tip explant transformation
showed efficiency (number of GUS-positive
spots/sample) that was approximately 80%
lower than the average of the other explants.
The ability to transform peach embryogenic
callus, embryonic axes and cotyledons was
considered to be significant because regenera-
tion from these tissues had been previously
reported.
Peach is not unique in the Prunus in its
recalcitrance to transformation and regenera-
tion. There are few reports of the successful
production of transgenic Prunus species. Those
species that have been transformed include
apricot (P. armeniaca) (Laimer da Camara
Machado et al., 1992), sweet cherry (Prunus
avium) (Brasileiro et al., 1991), sweet × sour
cherry (Dolgov and Firsov, 1999), almond
(Prunus amygdalus) (Miguel and Oliveira,
1999), P. avium × Prunus pseudocerasus cv. Colt
(Gutierrez-Pesce et al., 1998), Prunus subhir-
tella autumno rosa (da Câmara Machado et al.,
1995) and P. domestica (European or prune plum)
(Mante et al., 1991). For most of these species
there exists a single report of the development
of only a few transgenic plants. The P. domes-
tica system, which uses mature seeds as the
explant source, has been used repeatedly to
develop transgenic trees (Mante et al., 1991;
Scorza et al., 1994, 1995b; Padilla et al., 2003)
and presents what can be considered a reli-
able and routine system. It is such a system
that remains a goal for peach and one that
will advance the utilization of gene transfer
for peach improvement.
4.4 Conclusions and Perspectives
Significant progress has been made in recent
years to understand the genome organization
in peach and its close relatives. The genome
organization of Prunus species is highly col-
linear; there is significant progress towards
the completion of a physical map/genetic
 Genetic Engineering and Genomics 101

map resource and significant numbers of genes
have been identified through EST and genomic
sequencing projects in peach. These gene
sequences are mapped on the physical/
genetic map database and this information is
publicly available in the GDR. This work
paves the way for future identification and
cloning of important genes in cultivation of
peach and other Prunus species. Recent
reports utilizing these resources have demon-
strated the importance of this database for
identification and study of important fruit
tree genes. Manipulation of these genes in
peach awaits the development of a reliable
transformation system for peach.
Future work will focus on the utilization
of this gene information and marker systems
for manipulation of important characters in
the breeding schemes. Integration of the
molecular genetic resources for peach with
traditional breeding programmes promises to
streamline the breeding process and provide
new and improved varieties for the global
market.

References

Abbott, A.G., Rajapakse, S., Sosinski, B., Lu, Z.X., Sossey-Alaoui, K., Gannavarapu, M., Reighard, G., Ballard,
R.E., Baird, W.V., Scorza, R. and Callahan, A. (1998) Construction of saturated linkage maps of peach
crosses segregating for characters controlling fruit quality, tree architecture and pest resistance. Acta
Horticulturae 465, 41–49.
Aranzana, M.J., Pineda, A., Cosson, P., Dirlewanger, E., Ascasibar, J., Cipriani, G., Ryder, C.D., Testolin, R.,
Abbott, A., King, G.J., Iezzoni, A.F. and Arús, P. (2003) A set of simple-sequence repeat (SSR) markers
covering the Prunus genome. Theoretical and Applied Genetics 106, 819–825.
Arondel, V., Lemieux, B., Hwang, I., Gibson, S., Goodman, H.M. and Somerville, C.R. (1992) Map-based
cloning of a gene controlling omega-3 fatty acid desaturation in Arabidopsis. Science 258, 1353–1355.
Arús, P., Mnejja, M., Esmenjaud, D., Bosselut, N. and Dirlewanger, E. (2004) High marker density around the
peach nematode resistance genes. Acta Horticulturae 658, 567–571.
Avramova, Z., Tikhonov, A., Chen, M. and Bennetzen, J.L. (1998) Matrix attachment regions and structural
colinearity in the genomes of two grass species. Nucleic Acids Research 26, 761–767.
Baird, W.V., Estager, A.S. and Wells, J. (1994) Estimating nuclear DNA content in peach and related diploid
species using laser flow cytometry and DNA hybridization. Journal of the American Society of Horticul-
tural Science 199, 1312–1316.
Bennetzen, J.L., SanMiguel, P., Liu, C.N., Chen, M., Tikhonov, A., Costa de Oliveira, A., Jin, Y.K., Avramova,
Z., Woo, S.S., Zhang, H. and Wing, R.A. (1996) The Hybaid Lecture. Microcollinearity and segmental
duplication in the evolution of grass nuclear genomes. Symposium of the Society of Experimental
Biology 50, 1–3.
Bielenberg, D.G., Wang, Y., Fan, S., Reighard, G.L., Scorza, R. and Abbott, A.G. (2004) A deletion affecting
several gene candidates is present in the peach Evergrowing mutant. Journal of Heredity 95, 436–444.
Bliss, F.A., Arulsekar, S., Foolad, M.R., Becerra, V., Gillen, A.M., Warburton, M.L., Dandekar, A.M., Kocsisne,
G.M. and Mydin, K.K. (2002) An expanded genetic linkage map of Prunus based on an interspecific
cross between almond and peach. Genome 45, 520–529.
Brasileiro, A.C., Leple, J.C., Muzzin, J., Ounnoughi, D., Michael, M.F. and Jouanin, L. (1991) An alternative approach
for gene transfer in trees using wild-type Agrobacterium strains. Plant Molecular Biology 17, 441–452.
Byrne, D.H. (1990) Isozyme variability in four diploid stone fruits compared with other woody perennial
plants. Journal of Heredity 81, 68–71.
Chaparro, J.X., Werner, D.J., O’Malley, D. and Sederoff, R.R. (1994) Targeted mapping and linkage analysis of
morphological isozyme, and RAPD markers in peach. Theoretical and Applied Genetics 87, 805–815.
Chen, M., SanMiguel, P., de Oliveira, A.C., Woo, S.S., Zhang, H., Wing, R.A. and Bennetzen, J.L. (1997) Mi-
crocolinearity in sh2-homologous regions of the maize, rice, and sorghum genomes. Proceedings of the
National Academy of Sciences USA 94, 3431–3435.
Claverie, M., Bosselut, N., Lecouls, A.C., Voisin, R., Lafargue, B., Poizat, C., Kleinhentz, M., Laigret, F., Dirl-
ewanger, E. and Esmenjaud, D. (2004) Location of independent root-knot nematode resistance genes in
plum and peach. Theoretical and Applied Genetics 108, 765–773.
Cone, K.C., McMullen, M.D., Bi, I.V., Davis, G.L., Yim, Y., Gardiner, J.M., Polacco, M.L., Sanchez-Villeda, H.,
Fang, Z., Schroeder, S.G., Havermann, S.A., Bowers, J.E., Paterson, A.H., Soderlund, C.A., Engler, F.W.,
102 A.G. Abbott et al.

Wing, R.A. and Coe, E.H. Jr. (2002) Genetic, physical, and informatics resources for maize. On the road
to an integrated map. Plant Physiology 130, 1598–1605.
Corredor, E., Román, M., García, E., Perera, E., Arús, P. and Naranjo, T. (2004) Physical mapping of rDNA
genes enables to establish the karyotype of almond. Annals of Applied Biology 144, 219–222.
da Câmara Machado, A., Puschmann, M., Puringer, H., Kremen, R., Katinger, H. and Laimer da Câmara
Machado, M. (1995) Somatic embryogenesis of Prunus subhirtella autumno rosa and regeneration of
transgenic plants after Agrobacterium-mediated transformation. Plant Cell Reports 14, 335–340.
Dettori, M.T., Quarta, R. and Verde, I. (2001) A peach linkage map integrating RFLPs, SSRs, RAPDs, and mor-
phological markers. Genome 44, 783–790.
Diaz, M.D. (1974) Vegetative and reproductive growth habits of evergreen peach trees in Mexico. Proceedings
of the XIX International Horticultural Congress 18, 525.
Dirlewanger, E. and Arús, P. (2008) Markers in fruit tree breeding: improvement of peach. In: Lörz, H. and
Wenzel, G. (eds) Molecular Marker Systems in Plant Breeding and Crop Improvement. Springer, Berlin,
pp. 279–304.
Dirlewanger, E., Pronier, V., Parvery, C., Rothan, C., Guye, A. and Monet, R. (1998) Genetic linkage map of
peach (Prunus persica (L.) Batsch) using morphological and molecular markers. Theoretical and Applied
Genetics 97, 888–895.
Dirlewanger, E., Moing, A., Rothan, C., Svanella, L., Pronier, V., Guye, A., Plomion, C. and Monet, R. (1999)
Mapping QTL controlling fruit quality in peach (Prunus persica (L.) Batsch). Theoretical and Applied
Genetics 98, 18–31.
Dirlewanger, E., Cosson, P., Poizat, C., Laigret, F., Aranzana, M.J., Arús, P., Dettori, M.T., Verde, I. and Quarta,
R. (2003) Synteny within the Prunus genomes detected by molecular markers. Acta Horticulturae 622,
177–187.
Dirlewanger, E., Graziano E., Joobeur, T., Garriga-Calderé, F., Cosson P., Howad, W. and Arús, P. (2004) Com-
parative mapping and marker assisted selection in Rosaceae fruit crops. Proceedings of the National
Academy of Sciences USA 101, 9891–9896.
Dolgov, S.V. and Firsov, A.P. (1999) Regeneration and agrobacterial transformation of sour cherry leaf disks.
Acta Horticulturae 484, 577–580.
Dominguez, I., Graziano, E., Gebhardt, C., Barakat, A., Berry, S., Arús, P., Delseny, M. and Barnes, S. (2003)
Plant genome archaeology: evidence for conserved ancestral chromosome segments in dicotyledonous
plant species. Plant Biotechnology Journal 1, 91–99.
Dunford, R.P., Kurata, N., Laurie, D.A., Money, T.A., Minobe, Y. and Moore, G. (1995) Conservation of fine-
scale DNA marker order in the genomes of rice and the Triticeae. Nucleic Acids Research 23, 2724–
2728.
Etienne, C., Rothan, C., Moing, A., Plomion, C., Bodenes, C., Dumas, L.S., Cosson, P., Pronier, V., Monet, R.
and Dirlewanger, E. (2002) Candidate genes and QTL for sugar and organic acid content in peach (Pru-
nus persica (L.) Batsch). Theoretical and Applied Genetics 105, 145–159.
FAO (2007) FAOSTAT Database. http://faostat.fao.org (accessed August 2007).
Foolad, M.R., Arulsekar, S., Becerra, V. and Bliss, F.A. (1995) A genetic map of Prunus based on an interspe-
cific cross between peach and almond. Theoretical and Applied Genetics 91, 262–269.
Foulongne, M. (2002) Introduction d’une résistence polygénique à l’oïdium chez le pêcher Prunus persica à
partir d’une espèce sauvage Prunus davidiana. PhD thesis, Université de la Mediterranée, Faculté de
Sciences de Marseille–Luminy, France.
Foulongne, M., Pascal, T., Arús, P. and Kervella, J. (2003a) The potential of Prunus davidiana for introgression
into peach [Prunus persica (L.) Batsch] assessed by comparative mapping. Theoretical and Applied Ge-
netics 107, 227–238.
Foulongne, M., Pascal, T., Pfeiffer, F. and Kervella, J. (2003b) QTL for powdery mildew resistance in
peach × Prunus davidiana crosses: consistency across generations and environments. Molecular Breed-
ing 12, 33–50.
Frary, A., Nesbitt, T.C., Frary, A., Grandillo, S., van der Knaap, E., Cong, B., Liu, J., Meller, J., Elber, R., Alpert,
K.B. and Tanksley, S.D. (2000) fw2.2: a quantitative trait locus key to the evolution of tomato fruit size.
Science 289, 85–88.
Gentile, A., Monticelli, S. and Damiano, C. (2002) Adventitious shoot regeneration in peach [Prunus persica
(L.) Batsch]. Plant Cell Reports 20, 1011–1016.
Georgi, L.L., Wang, Y., Yvergniaux, D., Ormsbee, T., Iñigo, M., Reighard, G. and Abbott, A.G. (2002) Construc-
tion of a BAC library and its application to the identification of simple sequence repeats in peach (Prunus
persica (L.) Batsch). Theoretical and Applied Genetics 105, 1151–1158.
 Genetic Engineering and Genomics 103

Georgi, L.L., Wang, L., Reighard, G.L., Mao, L., Wing, R.A. and Abbott, A.G. (2003) Comparison of peach and
Arabidopsis genomic sequences: fragmentary conservation of gene neighborhoods. Genome 46, 268–
276.
Guohua, Y. and Yu, Z. (2002) Plant regeneration from excised immature embryos of peach (Prunus persica L.).
Acta Horticulturae Sinica 29, 480–482.
Gutierrez-Pesce, P., Taylor, K., Muleo, R. and Rugini, E. (1998) Somatic embryogenesis and shoot regeneration
from transgenic roots of the cherry rootstock Colt (Prunus avium × Prunus pseudocerasus) mediated by
pRi 1855 T-DNA of Agrobacterium rhizogenes. Plant Cell Reports 17, 574–580.
Hammerschlag, F.A. and Smigocki, A.C. (1998) Growth and in vitro propagation of peach plants transformed
with the shooty mutant strain of Agrobacterium tumefaciens. HortScience 33, 897–899.
Hammerschlag, F.A., Bauchan, G. and Scorza, R. (1985) Regeneration of peach plants from callus derived
from immature embryos. Theoretical and Applied Genetics 70, 248–251.
Hammerschlag, F.A., McCanna, I.J. and Smigocki, A.C. (1997) Characterization of transgenic peach plants
containing a cytokinin biosynthesis gene. Acta Horticulturae 447, 569–574.
Hoskins, R.A., Nelson, C.R., Berman, B.P., Laverty, T.R., George, R.A., Ciesiolka, L., Naeemuddin, M.,
Arenson, A.D., Durbin, J., David, R.G., Tabor, P.E., Bailey, M.R., DeShazo, D.R., Catanese, J., Mammoser,
A., Osoegawa, K., de Jong, P.J., Celniker, S.E., Gibbs, R.A., Rubin, G.M. and Scherer, S.E. (2000) BAC-
based physical map of the major autosomes of Drosophila melanogaster. Science 287, 2271–2274.
Jáuregui, B. (1998) Localización de marcadores moleculares ligados a caracteres agronómicos en un cruzamien-
to interespecífico almendro × melocotonero. PhD thesis, Universitat de Barcelona, Barcelona, Spain.
Jáuregui, B., de Vicente, M.C., Messeguer, R., Felipe, A., Bonnet, A., Salesses, G. and Arús, P. (2001) A recipro-
cal translocation between ‘Garfi’ almond and ‘Nemared’ peach. Theoretical and Applied Genetics 102,
1169–1176.
Jelenkovic, G. and Harrington, E. (1972) Morphology of the pachytene chromosomes in Prunus persica.
Canadian Journal of Genetics and Cytology 14, 317–324.
Joobeur, T. (1998) Construcción de un mapa de marcadores moleculares y análisis genético de caracteres
agronómicos en Prunus. PhD thesis, Universitat de Lleida, Lleida, Spain.
Joobeur, T., Viruel, M.A., de Vicente, M.C., Jáuregui, B., Ballester, J., Dettori, M.T., Verde, I., Truco, M.J.,
Messeguer, R., Batlle, I., Quarta, R., Dirlewanger, E. and Arús, P. (1998) Construction of a saturated link-
age map for Prunus using an almond × peach F2 progeny. Theoretical and Applied Genetics 97, 1034–
1041.
Joobeur, T., Periam, N., de Vicente, M.C., King, G. and Arús, P. (2000) Development of a second generation
linkage map for almond using RAPD and SSR markers. Genome 43, 649–655.
Kilian, A., Chen, J., Han, F., Steffenson, B. and Kleinhofs, A. (1997) Towards map-based cloning of the barley
stem rust resistance genes Rpg1 and rpg4 using rice as an intergenomic cloning vehicle. Plant Molecular
Biology 35, 187–195.
Laimer da Camara Machado, M., da Camara Machado, A., Hanzer, V., Weiss, H., Regner, F., Steinkellner, H.,
Mattanovich, D., Plail, R., Knap, E., Kalthoff, B. and Katinger, H. (1992) Regeneration of transgenic plants
of Prunus armeniaca containing the coat protein gene of plum pox virus. Plant Cell Reports 11, 25–29.
Lambert, P., Hagen, L.S., Arús, P. and Audergon, J.M. (2004) Genetic linkage maps of two apricot cultivars
(Prunus armeniaca L.) compared with the almond ‘Texas’ × peach ‘Earlygold’ reference map for Prunus.
Theoretical and Applied Genetics 108, 1120–1130.
Lu, Z.X., Sosinski, B., Reighard, G.L., Baird, W.V. and Abbott, A.G. (1998) Construction of a genetic linkage
map and identification of AFLP markers for resistance to root-knot nematodes in peach rootstocks. Ge-
nome 41, 199–207.
Mante, S., Scorza, R. and Cordts, J. (1989) Plant regeneration from cotyledons of Prunus persica, Prunus do-
mestica, and Prunus cerasus. Plant Cell, Tissue and Organ Culture 19, 1–11.
Mante, S., Morgens, P., Scorza, R., Cordts, J. and Callahan, A. (1991) Agrobacterium-mediated transformation
of plum (Prunus domestica) hypocotyl slices and regeneration of transgenic plants. Biotechnology 9,
853–857.
Marra, M., Kucaba, T., Sekhon, M., Hillier, L., Martienssen, R., Chinwalla, A., Crokett, J., Fedele, J., Grover, H.,
Gund, C., McCombie, W.R., McDonald, K., McPherson, J., Mudd, N., Parnell, L., Schein, J., Seim, R.,
Shelby, P., Waterston, R. and Wilson, R. (1999). A map for sequence analysis of the Arabidopsis thaliana
genome. Nature Genetics 22, 269–270.
Michelmore, R.W., Paran, I. and Kesseli, R.V. (1991) Identification of markers linked to disease-resistance
genes by bulked segregant analysis: a rapid method to detect markers in specific genomic regions by
using segregating populations. Proceedings of the National Academy of Sciences USA 88, 1212–1217.
104 A.G. Abbott et al.

Miguel, C. and Oliveira, M.M. (1999) Transgenic almond (Prunus dulcis Mill.) plants obtained by Agrobacte-
rium-mediated transformation of leaf explants. Plant Cell Reports 18, 387–393.
Miller, P.J., Parfitt, D.E. and Weinbaum, S.A. (1989) Outcrossing in peach. HortScience 24, 359–360.
Monet, R., Guye, A., Roy, M. and Dachary, N. (1996) Peach Mendelian genetics: a short review and new
results. Agronomie 16, 321–329.
Moore, J. and Janick, J. (eds) (1975) Advances in Fruit Breeding. Purdue University Press, West Lafayette,
Indiana.
Mowrey, B.D., Werner, D.J. and Byrne, D.H. (1990) Inheritance of isocitrate dehydrogenase, malate dehydro-
genase, and shikimate dehydrogenase in peach and peach × almond hybrids. Journal of the American
Society of Horticultural Science 115, 312–319.
Murashige, T. and Skoog, F. (1962) A revised medium for rapid growth and bioassays with tobacco tissue
cultures. Physiologia Plantarum 15, 473–497.
Padilla, I.M.G., Webb, K. and Scorza, R. (2003) Early antibiotic selection and efficient rooting and acclimati-
zation improve the production of transgenic plum plants (Prunus domestica L.). Plant Cell Reports 22,
38–45.
Pooler, M.R. and Scorza, R. (1995) Regeneration of peach [Prunus persica (L.) Batsch] rootstock cultivars from
cotyledons of mature stored seed. HortScience 30, 355–356.
Rajapakse, S., Bethoff, L.E., He, G., Estager, A.E., Scorza, R., Verde, I., Ballard, R.E., Baird, W.V., Callahan, A.,
Monet, R. and Abbott, A.G. (1995) Genetic linkage mapping in peach using morphological, RFLP and
RAPD markers. Theoretical and Applied Genetics 90, 503–510.
Raj Bhansali, R., Driver, J.A.A. and Durzan, D.J. (1990) Rapid multiplication of adventitious somatic embryos
in peach and nectarine by secondary embryogenesis. Plant Cell Reports 9, 280–284.
Rodriguez, J., Sherman, W.B., Scorza, R., Wisniewski, M. and Okie, W.R. (1994) Evergreen peach, its inheri-
tance and dormant behavior. Journal of the American Society for Horticultural Science 119, 789–792.
Rugini, E. and Gutierrez-Pesce, P. (1999) Transgenic Prunus fruit species (almond, apricot, cherry rootstocks,
sour cherry and sweet cherry, peach and plum). In: Bajaj, Y.P.S. (ed.) Biotechnology in Agriculture and
Forestry. Vol. 44. Transgenic Trees. Springer, Berlin, pp. 245–262.
Salesses, G. and Mouras, A. (1977) Tentative d’utilisation des protoplastes pour l’etude des chromosomes
chez les Prunus. Annals de l’Amelioration des Plantes 27, 363–368.
Scorza, R. (1991) Gene transfer for the genetic improvement of perennial fruit and nut crops. HortScience 26,
1033–1035.
Scorza, R. (2001) Progress in tree fruit improvement through molecular genetics. HortScience 36, 855–857.
Scorza, R. and Hammerschlag, F. (1992) Emerging technologies for the genetic improvement of stone fruits.
In: Hammerschlag, F.A. and Litz, R.E. (eds) Biotechnology of Perennial Fruit Crops. CAB International,
Wallingford, UK, pp. 277–302.
Scorza, R. and Sherman, W.B. (1996) Peaches. In: Janick, J. and Moore, J.N. (eds) Fruit Breeding. Vol. I. Tree
and Tropical Fruit. Wiley, New York, pp. 325–440.
Scorza, R., Mehlenbacher, S.A. and Lightner, G.W. (1985) Inbreeding and co-ancestry of freestone peach
cultivars of the eastern United States and implications for peach germplasm improvement. Journal of the
American Society of Horticultural Science 110, 547–552.
Scorza, R., Cordts, J. and Mante, S. (1990a) Long-term somatic embryo production and plant regeneration
from embryo-derived peach callus. Acta Horticulturae 280, 183–190.
Scorza, R., Morgens, P., Cordts, J.M., Mante, S. and Callahan, A. (1990b) Agrobacterium-mediated transfor-
mation of peach [Prunus persica (L.) Batsch] leaf segments, immature embryos, and long-term embryo-
genic callus. In vitro Cellular and Developmental Biology 26, 829–834.
Scorza, R., Ravelonandro, M., Callahan, A., Cordts, J.M., Fuchs, M., Dunez, J. and Gonsalves, D. (1994)
Transgenic plums (Prunus domestica L.) express the plum pox virus coat protein gene. Plant Cell Reports
14, 18–22.
Scorza, R., Hammerschlag, F.A. and Zimmerman, T.W. (1995a) Genetic transformation in Prunus persica
(peach) and Prunus domestica (plum). In: Bajaj, Y.P.S. (ed.) Biotechnology in Agriculture and Forestry,
Vol. 34. Springer, Berlin, pp. 255–268.
Scorza, R., Levy, L., Damsteegt, V., Yepes, M., Cordts, J., Hadidi, A., Slightom, J. and Gonsalves, D. (1995b)
Transformation of European plum (Prunus domestica L.) with the papaya ringspot virus coat protein gene
and reaction of transgenic plants to inoculation with plum pox virus. Journal of the American Society of
Horticultural Science 120, 943–952.
Scorza, R., Melnicenco, L., Dang, P. and Abbott, A.G. (2002) Testing a microsatellite marker for selection of
columnar growth habit in peach (Prunus persica (L.) Batsch). Acta Horticulturae 592, 285–289.
 Genetic Engineering and Genomics 105

Sherman, W.B. and Lyrene, P.M. (1983) Handling seedling populations. In: Moore, J.N. and Janick, J. (eds)
Methods in Fruit Breeding. Purdue University Press, West Lafayette, Indiana, pp. 66–73.
Smigocki, A.C. and Hammerschlag, F.A. (1991) Regeneration of plants from peach embryo cells infected with
shooty mutant strain of Agrobacterium. Journal of the American Society of Horticultural Science 116,
1092–1097.
Soderlund, C., Humphray, S., Dunham, A. and French, L. (2000) Contigs built with fingerprints, markers, and
FPC V4.7. Genome Research 10, 1772–1787.
Srinivasan, C. and Scorza, R. (1999) Transformation of somatic embryos of trees and grapevine. In: Jain, S.M.,
Gupta, P.K. and Newton, N.J. (eds) Somatic Embryogenesis in Woody Plants, Vol. 5. Kluwer Academic
Press, London, pp. 313–330.
Srinivasan, C., Padilla, I.M.G. and Scorza, R. (2004) Prunus species. In: Litz, R. (ed.) Biotechnology of Fruit
and Nut Crops. CAB International, Wallingford, UK, pp. 512–542.
Tanksley, S.D., Young, N.D., Paterson, A.H. and Bonierbale, M.W. (1989) RFLP mapping in plant breeding:
new tools for an old science. Biotechnology 7, 257–263.
Tao, Q., Chang, Y., Wang, J., Chen, H., Islam-Faridi, M.N., Scheuring, C., Wang, B., Stelly, D.M. and Zhang,
H. (2001) Bacterial artificial chromosome-based physical map of the rice genome constructed by restric-
tion fingerprint analysis. Genetics 158, 1711–1724.
van Dodeweerd, A.-M., Hall, C.R., Bent, E.G., Johnson, S.J., Bevan, M.W. and Bancroft, I. (1999) Identifica-
tion and analysis of homoeologous segments of the genomes of rice and Arabidopsis thaliana. Genome
42, 887–892.
Verde, I., Quarta, R., Cerdrola, C. and Dettori, M.T. (2002) QTL analysis of agronomic traits in a BC1 peach
population. Acta Horticulturae 592, 291–297.
Vilanova, S., Romero, C., Abbott, A.G., Llácer, G. and Badenes, M.L. (2003) An apricot (Prunus armeniaca L.)
F2 progeny linkage map based on SSR and AFLP markers, mapping plum pox virus resistance and self-
incompatibility traits. Theoretical and Applied Genetics 107, 239–247.
Viruel, M.A., Madur, D., Dirlewanger, E., Pascal, T. and Kervella, J. (1998) Mapping quantitative trait loci
controlling peach leaf curl resistance. Acta Horticulturae 465, 79–87.
Wang, Y., Georgi, L.L., Zhebentyayeva, T.N., Reighard, G.L., Scorza, R. and Abbott, A.G. (2001) High through-
put targeted SSR marker development in peach (Prunus persica (L.) Batsch). Genome 45, 319–328.
Wang, Y., Georgi, L.L., Reighard, G.L., Scorza, R. and Abbott, A.G. (2002) Genetic mapping of the evergrowing
gene in peach (Prunus persica (L.) Batsch). Journal of Heredity 93, 352–358.
Warburton, M.L., Becerra-Velasquez, V.L., Goffreda, J.C. and Bliss, F.A. (1996) Utility of RAPD markers in
identifying genetic linkages to genes of economic interest in peach. Theoretical and Applied Genetics
93, 920–925.
Werner, D.J. and Okie, W.R. (1998) A history and description of the Prunus persica plant introduction collection.
HortScience 33, 787–793.
Yamamoto, T. and Hayashi, T. (2002) New root-knot nematode resistance genes and their STS markers in
peach. Scientia Horticulturae 96, 81–90.
Yamamoto, T., Shimada, T., Imai, T., Yaegaki, H., Haji, T., Matsuta, N., Yamaguchi, M. and Hayashi, T. (2001)
Characterization of morphological traits based on a genetic linkage map in peach. Breeding Science 51,
271–278.
Yamamoto, Y., Yamaguchi, M. and Hayashi, T. (2005) An integrated genetic linkage map of peach by SSR, STS,
AFLP and RAPD. Journal of the Japanese Society for Horticultural Science 74, 204–213.
Ye, X., Brown, S.K., Scorza, R., Cordts, J.M. and Sanford, J.C. (1994) Genetic transformation of peach tissues
by particle bombardment. Journal of the American Society of Horticultural Science 119, 367–373.
 5 Low-chill Cultivar Development

B.L. Topp,1 W.B. Sherman2 and M.C.B. Raseira3

1Queensland Department of Primary Industries & Fisheries, Nambour,
Queensland, Australia
2Horticultural Science Department, University of Florida, Gainesville, Florida, USA
3Empresa Brasileira de Pesquisa Agropecuaria/Embrapa Clima Temperado, Pelotas,

Rio Grande do Sul, Brazil

5.1 Introduction 107

Reasons for breeding low-chill cultivars 107
History of low-chill cultivar development 108
5.2 Current Low-chill Breeding Programmes 110
Australia, New South Wales 113
Australia, Queensland 113
Brazil, Pelotas 117
Brazil, Sao Paulo 118
Mexico, Chapingo 118
Mexico, Queretaro 118
South Africa, Stellenbosch 119
Taiwan, Taichung 119
Thailand, Angkhang 119
USA, California, Sun World 119
USA, California, Zaiger Genetics 120
USA, Florida, University of Florida 120
USA, Texas, Texas A&M University 121
5.3 Common Objectives in Low-chill Peach Breeding 121
Low chilling requirement 121
Time of flowering 122
Flower bud density 123
Blind nodes and bud drop 123
Fruit development period 123
Fruit shape 125
Fruit firmness 126
Disease resistance 127
5.4 Management of Low-chill Breeding Populations 129
Hybrid seed production 129
Seedling populations 132
Testing advanced selections 133
5.5 Conclusions 133
106 © CAB International 2008. The Peach: Botany, Production and Uses
(eds D.R. Layne and D. Bassi)
 Low-chill Cultivar Development
5.1 Introduction
When Professor R.E. Smith commenced
planning for pathological work in southern
California about 1904, one of the troubles to
be studied and, if possible, prevented, was a
rather vague disease of peaches, in which
there was delayed blooming and foliation,
with loss of crop. . . . Among other things
undertaken in connection with the peach
problem was a breeding project in which the
adaptability of certain South China peaches
to the climate of southern California was to
be combined with the desired commercial
qualities of ordinary varieties.
(Horne et al., 1926)
So began the first attempts in US public
breeding aimed at developing low-chill peach
cultivars.
The term ‘low-chill’, used to describe peach
cultivars, is a relative and somewhat artificial
term. Chilling refers to the amount of low
temperatures required by leaf and flower
buds during winter to break dormancy and
commence normal growth and development
each growing season. Chilling has tradition-
ally been quantified as chill-hours, measuring
the number of hours below 7°C (Weinberger,
1950), and more recently as chill units (CU),
which allow for partial chill-hour accumula-
tion and chill negation (Dennis, 2003). The
distinction of cultivars as low-, medium- and
high-chill provides broad categories rather than
exact definitions, as there is actually a contin-
uum of chilling requirements. As a general
rule, low-chill cultivars are those adapted to
subtropical environments, high-chill cultivars
are those adapted to temperate environments
and medium-chill cultivars are intermediate
in adaptation. In this chapter we focus on the
breeding of low-chill peach cultivars adapted
to the subtropics, even though they may be
grown in temperate environments (in the
absence of spring freezes that kill the bloom)
or in tropical highlands.
Reasons for breeding low-chill cultivars
Low-chill peach cultivars are economically
important for a number of reasons. In sub-
tropical regions, they provide a fresh supply
107
of fruit for local markets and the potential to
transport fruit to more distant markets not
producing peaches at that time. The low chill-
ing requirement of these cultivars means that
the trees flower in mid- to late winter. In
warm environments the fruit develop rapidly,
resulting in harvests that are up to 2 months
earlier than cultivars grown in cooler, temper-
ate regions. Fruit prices are usually high at
this time.
Globalization and reduction in trade
barriers between northern and southern
hemisphere countries will impact on low-
chill cultivar development. Old markets that
supplied low-chill cultivars because they
were early, but lacked fruit quality, will be
rendered obsolete by the importation of late-
season, high-quality, high-chill fruit from
opposite hemispheres. In Australia, the only
peaches on the market in September are from
the low-chill cultivar ‘Flordaprince’, but
California has applied to export peaches to
Australia. If this occurs, then late-season
Californian cultivars such as ‘August Red’,
‘Autumn Flame’ and ‘September Sun’ will be
on Australian markets in September and may
out-compete ‘Flordaprince’ fruit in terms of
size, colour, firmness and flavour. New mar-
kets are also likely to develop for low-chill
cultivars, but this will require improvements
in flesh firmness, transportability and specific
fruit quality characteristics tailored to new
markets.
Finally, low-chill cultivars may become
more widely planted if global warming trends
continue. The Intergovernmental Panel on
Climate Change reported projected increases
in global average surface temperatures of
0.8–2.6°C by 2050, with minimums increasing
at twice the rate of maximums (Watson et al.,
2001). Increases of this magnitude would
result in less winter chilling in many loca-
tions. Hennessy and Clayton-Greene (1995)
modelled an increase of 1.5–2.0°C and found
it would increase the risk of prolonged dor-
mancy for stone fruits at many temperate
fruit-growing regions in southern Australia.
Using a modified Utah chill model, chilling at
key Australian fruit-growing regions was
shown to decrease (Table 5.1). Under these
conditions, the demand for low-chill cultivars
would increase considerably.
108 B.L. Topp et al.

Table 5.1. Annual average chilling (chill units, CU) accumulated in 1995 and predicted for 2030 at sites
in Australian peach-growing regions. The range of values for 2030 represents the effect of a low and high
global warming scenario. (Adapted from Hennessy and Clayton-Greene, 1995.)

Location Average chilling (CU), 1995 Predicted chilling (CU), 2030

Bridgetown, Western Australia 1409 671–1159

Renmark, South Australia 1187 529–983
Griffith, New South Wales 1486 944–1316
Stanthorpe, Queensland 1316 888–1185
Swan Hill, Victoria 1608 898–1397

History of low-chill cultivar development
From its original home in China, the peach
has been widely distributed through Europe,
Asia, South Africa, Australia and the Ameri-
cas, and over the centuries there has been
selection of locally adapted cultivars (Culli-
nan, 1937). The honey and peento peaches
from southern China are adapted to the sub-
tropics and were the original source of low-
chill adaptation used in early low-chill peach
breeding in southern California (Cullinan,
1937). The European or Persian peaches were
introduced from China to the Mediterranean,
west Asia, Italy, Spain, Iran and Turkey (Li,
1984). This germplasm, including non-melt-
ing, yellow-fleshed clingstones, was intro-
duced to the New World by Spanish and
Portuguese explorers in the 16th century. This
peach group provided source material used
in low-chill peach development in South and
Central America.
While the original source of low-chill
requirement is from southern China, it reached
the subtropics by varied routes. In Australia,
subtropical peaches are thought to have been
introduced by Chinese immigrants in the
1800s and gave rise to cultivars such as ‘China
Flat’, ‘Bell’s November’, ‘Beauty of Boorood-
abin’, ‘Watt’s Early Champion’ and strains of
‘Dwarf Peach’ (Ward, 1952). The ‘China Flat’
was imported as seed into the USA from Aus-
tralia in 1869 and named ‘Australian Saucer’
(Hume, 1902; Price, 1905; Lammerts, 1941).
The heterozygous nature of the flat or peento
shape (because homozygous peento is lethal)
would have ensured segregation for both
regular and flat-shaped peaches from this
imported seed. Hume (1902) reported that 24
varieties belong to this peento group includ-
ing ‘Angel’, ‘Jewel’, ‘Bidwell Early’, ‘Waldo’
and ‘Peen-to’ and that they were adapted to
subtropical conditions, especially in Florida.
In the USA, low-chill breeding began in
1907 at the University of California Citrus
Experiment Station at Riverside, and not long
after at the Chaffee Junior College. Low-chill
adaptation was obtained from cultivars from
the southern Chinese and peento groups of
peaches including ‘Peento’ and ‘Lukens Honey’.
These were intercrossed and also combined
with high-chill cultivars such as ‘Elberta’ and
‘Mayflower’ to produce a range of white- and
yellow-fleshed, melting and non-melting cul-
tivars. In 1933, these programmes jointly
released ‘Babcock’, a 350 CU peach described
as an early-ripening, white-fleshed freestone
of fair size and good quality (Cullinan, 1937).
‘Babcock’ and its derivates ‘Giant Babcock’
and ‘Early Babcock’ were still being grown in
California in the 1990s and exported to Asia
where low-acid types are popular. Between
1933 and 1958 these breeding programmes
released 23 cultivars adapted to the mild win-
ters of southern California with 350 to 450 CU
required (Table 5.2). ‘Babcock’ was also used
extensively in later breeding programmes as
a source of low-chill adaptation.
In the late 1940s and early 1950s, low-
chill breeding commenced in the USA at the
University of Florida (UF) and in Brazil at Sao
Paulo and Pelotas. All three programmes used
the south China-derived ‘Peento’ and ‘Hawai-
ian’ as founding parents for low-chill adapta-
tion (Byrne, 2003). In addition, the Brazilian
programmes used a number of local cultivars
Table 5.2. Characteristics and parentage of peach cultivars released from the University of California, Riverside (1907–1961) and Chaffey Junior College
(1919–1947) breeding programmes in southern California. (From Okie, 1998.)

Cultivar Year of release Chill (CU) Flesh coloura Origin Female parent Male parent

‘Babcock’ 1933 350 W UC–Riverside ‘Strawberry’ × ‘Peento’ F2

‘C.O. Smith’ 1933 450 W UC–Riverside ‘Strawberry’ × ‘Peento’ F2
‘Chaffey’ 1939 450 W Chaffey Jr Coll. ‘Lukens Honey’ ‘Elberta’
‘Fontana’ 1939 450 Y Chaffey Jr Coll. ‘Sims’ ‘Feicheng’ × ‘Bolivian Cling’

Low-chill Cultivar Development

‘Golden State’ 1939 450 Y UC–Riverside ‘Paragon’ ‘BH7-7-4’
‘Hermosa’ 1939 450 W UC–Riverside ‘J.H. Hale’ ‘Babcock’
‘Prenda’ 1939 350 W UC–Riverside ‘J.H. Hale’ ‘11-May’
‘Ramona’ 1939 450 Y UC–Riverside ‘7-2’ ‘OP’
‘Weldon’ 1939 450 Y Chaffey Jr Coll. ‘Babcock’ ‘Elberta’
‘Bonita’ 1943 450 Y UC–Riverside ‘Prenda’ ‘Sunglow’
‘Banquet’ 1945 100 Y UC–Riverside ‘Babcock’ ‘Quetta’
‘Babdon’ 1948 450 W Chaffey Jr Coll. ‘Babcock’ ‘Weldon’
‘Chadon’ 1948 450 W Chaffey Jr Coll. ‘Chaffey’ ‘Weldon’
‘Gloribloom’ 1948 450 Y Chaffey Jr Coll. ‘Babcock’ ‘Elberta’
‘Honeyberta’ 1948 450 Y Chaffey Jr Coll. ‘Lukens Honey’ ‘Elberta’
‘Impon’ 1948 450 Y Chaffey Jr Coll. ‘Imperial’ ‘Paragon’
‘Maydon’ 1948 450 W Chaffey Jr Coll. ‘Mayflower’ ‘Weldon’
‘Maywel’ 1948 450 W Chaffey Jr Coll. ‘Mayflower’ ‘Weldon’
‘Welberta’ 1948 450 Y Chaffey Jr Coll. ‘Weldon’ ‘Elberta’
‘Rubidoux’ 1949 350 Y UC–Riverside Unknown Unknown
‘Ventura’ 1951 450 Y UC–Riverside ‘Hermosa’ ‘71-9’
‘Rochon’ 1958 450 Y UC–Riverside ‘Peento Hybrid’ (‘Rochester’ × ‘Peento’)
‘Tejon’ 1958 350 Y UC–Riverside Unknown Unknown
aFlesh colour: W, white; Y, yellow.

109
110 B.L. Topp et al.
imported by the Portuguese in the 16th cen-
tury (Byrne and Bacon, 1999).
Byrne and Bacon (1999) analysed pedi-
grees of cultivars released from 1976 to 1997
to determine the founding parents of low-
chill breeding programmes in Brazil, Mexico
and the USA. ‘Peento’ was identified as a pri-
mary source in all breeding programmes and
is in the pedigree of the UF cultivars ‘Sunred’,
‘Sunlite’ and ‘Maravilha’. Other important
founding clones, as sources of the low-chill-
ing trait, are ‘Okinawa’ and ‘Hawaiian’. All
three cultivars produce fruit with a long fruit
development period (FDP; days from flower-
ing to harvest) that is white-fleshed, soft and
poor quality, but have provided a source of
adaptation to subtropical environments.
Lack of genetic diversity in breeding
populations may limit cultivar improvement
and prevent development of novel gene com-
binations for future progress. High levels of
inbreeding and co-ancestry have been reported
for high-chill peaches (Scorza et al., 1985).
While all low-chill breeding programmes have
used high-chill germplasm for high fruit qual-
ity, the inclusion of native low-chill parents in
the breeding means they are genetically dis-
tinct. Scorza et al. (1988) studied 32 cultivars
released by UF and found that the level of
inbreeding was 0.039, assuming open pollina-
tion equated to outcrossing by unrelated males,
but 0.286 with an assumption of open pollina-
tion equating to selfing. The second case is a
relatively high level of inbreeding, consider-
ing that the inbreeding coefficient will be
0.125 for half-sib mating and 0.250 for full-sib
mating, and is comparable with the levels of
inbreeding in eastern US high-chill peaches.
Scorza et al. (1988) concluded that although
unique and unrelated germplasm had been
incorporated into the UF cultivars, a certain
level of inbreeding had been required to con-
centrate the tree and fruit quality traits neces-
sary for commercialization.
5.2 Current Low-chill Breeding
Programmes
Eleven public and two private breeding pro-
grammes across seven countries are currently
developing new low-chill peach and nectar-
ine cultivars (Table 5.3). Most of these pro-
grammes have a primary focus on breeding
low-chill cultivars although some, such as
Zaiger Genetics, Inc., breed predominately
for high-chill but have released several low-
chill cultivars.
Low-chill breeding programmes are
located where the average temperature of the
coldest month ranges from 12 to 16°C (Table
5.3). This corresponds to an average winter
chilling of 250 to 650 CU using the Sharpe–
Weinberger model (Sherman and Rodriquez-
Alcazar, 1987; Sharpe et al., 1990). Low-chill
breeding programmes are located between
latitudes of 19° and 30°. Programmes located
at the lower tropical latitudes are at high ele-
vation, thus providing winter chill (Table 5.3).
Low-chill breeding programmes are
established in environments ranging from
dry deserts, humid subtropics and tropical
highlands to temperate locations (Table 5.4).
Cultivars released from these programmes
provide orchardists with adaptation to vary-
ing climatic, disease and pest pressures. The
oldest of the current programmes are in the
USA (Gainesville), Brazil (Pelotas and Campi-
nas) and South Africa (Stellenbosch). The
number of programmes has increased in the
past 25 years with five programmes com-
mencing in the 1980s and another three com-
mencing in the 1990s (Table 5.4).
It is interesting to speculate on the rea-
sons for the increased effort in low-chill
breeding over the past 25 years. Part of the
increase is probably due to demand for culti-
vars adapted to local environments and local
markets that were not being filled by imported
cultivars. For example, the breeding pro-
grammes in Mexico select specifically for
non-melting, yellow-fleshed peaches that
have a small amount of red blush. Mexican
consumers associate this type of fruit with the
high eating quality of native criollo peaches
and so the Mexican breeders have market
forces directing their breeding efforts in this
direction; this is quite different to the objec-
tives of many other breeding programmes
that concentrate on highly coloured, melting-
flesh types. Increased activity is probably also
due to the influence of the UF breeding pro-
gramme. New UF cultivars have been spread
Table 5.3. Location, latitude, altitude, coldest month mean temperature and winter chilling of 13 breeding programmes that have released low-chill peach and
nectarine cultivars.

Mean temperature Mean winter

Country Institution or company Latitude (°) Altitude (m) of coldest month (°C) chilling (CU)a

Australia Department of Primary Industries and Fisheries, 26 25 15.0 200

Nambour, Queensland
Australia University of Western Sydney, Richmond, New South Wales 33 113 10.2 540

Low-chill Cultivar Development

Brazil Empressa Brasileira de Pesquisa Agropecuaria/Centro 32 60 12.3 370
Nacional de Pesquisa de Fruteiras de Clima Temperado
(EMBRAPA/CPACT), Pelotas, Rio Grande do Sul
Brazil Instituto Agronomico (IAC), Campinas, Sao Paulo 23 669 17.0 100
Mexico Centro de Fruiticultura, Colegio de Postgraduidos, Chapingo, 19 2245 12.9 320
Mexico State
Mexico Universidad Autonoma de Queretaro, Area Agricola, 20 1876 14.8 210
Queretaro, Queretaro
South Africa Agricultural Research Council Infruitec–Nietvoorbij, 33 91 12.4 360
Stellenbosch
Taiwan Taiwan Agricultural Research Institute, Wufeng, Taichung 24 85 15.7 160
Thailand Royal Project Foundation, Kasetsart University, Angkhang 19 1400 13.0 320
Royal Agricultural Station, Angkhang
USA Sun World International, Inc., Coachella, California 33 −22 12.3 370
USA Zaiger Genetics, Inc., Modesto, California 37 27 7.5 870
USA University of Florida, Gainesville, Florida 29 29 12.7 340
USA Texas A&M University, Weslaco, Texas 26 22 14.2 240
aWinter chill units (CU) of each location calculated using mean temperature of coldest month as per George and Nissen (1998).

111
112 B.L. Topp et al.

Table 5.4. Low-chill peach breeding programmesa arranged by starting year and climate type.

Climate type

Starting year Dry Humid subtropics Temperate Tropical highland

1937–1953 Pelotas, Brazil Stellenbosch,

Sao Paulo, Brazil South Africa
Gainesville, USA
1980–1990 Weslaco, USA Taichung, Taiwan Chapingo, Mexico
Coachella, USA Queretaro, Mexico
1990–2000 Nambour, Australia Richmond, Australia Angkhang, Thailand
aCountry, town, institution/company responsible for the breeding programme: Australia, Nambour,
Department of Primary Industries and Fisheries; Australia, Richmond, University of Western
Sydney; Brazil, Sao Paulo, Instituto Agronomico (IAC); Brazil, Pelotas, Empressa Brasileira de Pesquisa
Agropecuaria/Centro Nacional de Pesquisa de Fruteiras de Clima Temperado (EMBRAPA/CPACT);
Mexico, Chapingo, Colegio de Postgraduados; Mexico, Queretaro, Instituto Nacional de Investigaciones
Forestales y Agropecunarias; South Africa, Stellenbosch, Agricultural Research Council Infruitec; Taiwan,
Taichung, Taiwan Agricultural Research Institute; Thailand, Angkhang, Royal Foundation and Kasetsart
University; USA, Coachella, Sun World International, Inc.; USA, Gainesville, University of Florida; USA,
Weslaco, Texas A&M University.

far and wide, particularly up until 1992, when
all material was in the public domain. UF cul-
tivars are grown commercially and are the
mainstay of low-chill industries in many
countries. In Australia, for example, although
there was a small low-chill industry prior to
1980, the industry only expanded with the
introduction of UF cultivars. The industry has
now moved on to the next phase of develop-
ment where it requires new cultivars that are
tailored for the local environment and mar-
ket. However, without the importation of the
UF cultivars, the local industry would not
have developed and be in a position to spon-
sor new breeding.
Of the 842 new peach and nectarine cul-
tivar descriptions published internationally
between 1980 and 1992, approximately 16%
(135 cultivars) were low-chill (Della Strada et al.,
1996). In the next decade (1991–2001) 1126 new
peach and nectarine cultivar descriptions
were published, of which 71 (6%) were classi-
fied as low-chill (Della Strada and Fideghelli,
2003). The majority of low-chill cultivar releases
are from the three longest-running public breed-
ing programmes located in the USA (Gaines-
ville, Florida) and Brazil (Pelotas and Sao
Paulo) (Fig. 5.1/Plate 48). These programmes
have been operating for about 50 years and
released over 100 cultivars from 1980 to 2004
(Table 5.5). They have the common objective
of breeding for adaptation to subtropical
environments, but differ in other ways. The
Brazilian programmes are supplying culti-
vars for the full stone fruit season and so
release cultivars with FDP up to 180 days. In
the USA, the bulk of peach production is from
temperate high-chill locations with fruit avail-
able from late May; hence the Florida pro-
gramme aims to produce only short FDP
cultivars that are harvested before this time.
In the following sections low-chill breed-
ing programmes are described in terms of
objectives and outputs. There are sources of
low-chill cultivars other than the breeding
programmes described here. Some high-chill
breeding programmes develop low-chill culti-
vars for specific purposes. In California, private
breeding programmes located in temperate
environments select low-chill cultivars for
early production in the southern part of the
Central Valley. In Japan, low-chill peaches are
being created for use in protected culture (I.
Kataoka and K. Beppu, Japan, 2004, personal
communication). Traditional Japanese high-
chill cultivars such as ‘Hakuho’ are hybrid-
ized with low-chill cultivars ‘Flordaprince’,
‘Flordaglo’, ‘TropicSnow’ and ‘Earligrande’.
 Low-chill Cultivar Development
70
60
No. of cultivar releases
50
40
30
20
10
0
Brazil
Fig. 5.1. Numbers of new low-chill peach and nectarine cultivars released from four countries,
1980–1992. These countries accounted for 94% of low-chill cultivars released in this period. (Adapted
from Della Strada et al., 1996.)
Low-chill cultivars can be forced to flower in
heated plastic tunnels earlier than high-chill
cultivars and so ripen earlier. In China, there
is breeding for low-chill ornamental peach
cultivars with attractive flowers that can be
used for peach flower festivals during Chi-
nese New Year (Wang et al., 2002) but there is
only limited breeding for low-chill cultivars
for fresh fruit (L. Wang, China, 2004, personal
communication).
Australia, New South Wales
In Australia, low-chill cultivars are used to
extend the early season. Currently the early-
ripening UF cultivars are the mainstay of this
industry. Tree sales from the Australian Nurs-
erymen’s Fruit Improvement Company in
2001–2003 ranked ‘TropicBeauty’, ‘Florda-
prince’ and ‘UFGold’ as the most planted peach
cultivars and ‘SunWright’ and ‘SunBest’ as
the most planted nectarine cultivars (G. Por-
ter, Australia, 2004, personal communication).
The University of Western Sydney began
breeding low- and medium-chill peaches in
the early 1990s. The programme is located at
Richmond, New South Wales, in a temperate
location that receives annual winter chilling
of 400–600 CU. New selections are protected
and commercialized through the Phytonova
113
Canning peach
Nectarine
Peach
Mexico South Africa USA
company (McPhee, 2003). One of the selection
strategies for obtaining high eating quality is
to develop cultivars with FDP of 120 days.
The programme has released ten peach and
nectarine selections in the 300–400 CU range.
Australia, Queensland
The Queensland Department of Primary
Industries and Fisheries commenced breed-
ing low-chill peaches at Maroochy Research
Station at Nambour in the mid-1990s. The
aim of the programme is to produce high-
quality peach and nectarine cultivars requir-
ing 100–400 CU but with emphasis in the
100–150 CU range. Breeding is partly spon-
sored by grower organizations and has devel-
oped in response to their requirements for
new cultivars with local adaptation and with
their input into the breeding objectives and
commercialization of the new cultivars.
Domestically, it is important that quality
fruit are produced by the low-chill sector of the
industry in order to start consumers enjoying
their spring purchases of peaches. There is
also the prospect of expanding Australia’s
export to South-east Asia from September to
December as there is a marked reduction in
peach volume in Asia at this time (Nissen
et al., 2000; Wei, 2001). Initial hybridizations
114 B.L. Topp et al.

Table 5.5. Low-chill (<400 CU) peach and nectarine cultivars released from 1980 to 2004. Cultivars are
arranged alphabetically by the breeding programme from which they were released and then by cultivar name.

Year of Chillb Flesh Flesh FDPe

Cultivar introduction Cropa (CU) typec colourd (days) Purposef

Colegio de Postgraduados, Chapingo, Mexico

‘Diamante Mejorado’ 1995 P 250 NM Y 110 FR
‘Oro Mex’ 1995 P 350 NM Y 130 FR
‘Oro-Azteca’ 1992 P 275 NM Y 130 FR
‘Oro-B’ 1995 P 275 NM Y 105 FR
Empressa Brasileira de Pesquisa Agropecuaria (EMBRAPA), Pelotas, Rio Grande do Sul, Brazil
‘Ametista’ 1994 P 400 NM Y 131 PR
‘Anita’ 1999 N 375 NM W 97 FR
‘Atenas’ 2004 P 250 NM Y 125 DU
‘Bolinha’ 1986 P 400 NM Y 151 PR
‘BR-2’ 1981 P 300 NM O–Y 160 PR
‘BR-6’ 1981 P 350 NM Y 151 PR
‘Branca’ 1989 N 350 M W 90 FR
‘Charme’ 2000 P 350 M W 105 FR
‘Chimarrita’ 2000 P 350 M W 115 FR
‘Chinoca’ 1987 P 275 M W 106 FR
‘Chirua’ 1995 P 250 M W 113 FR
‘Chula’ 1995 P 400 M W 122 FR
‘Della Nona’ 1992 P >300 M W 121 FR
‘Dulce’ 1992 N 400 M W 103 FR
‘Eldorado’ 1989 P 300 NM Y 143 DU
‘Esmeralda’ 1987 P 350 NM Y 126 PR
‘Granada’ 1995 P 300 NM Y 98 PR
‘Granito’ 1993 P 400 NM O–Y 112 PR
‘Guaiaca’ 1993 P 250 M Y 99 FR
‘Jade’ 1987 P 300 NM Y 124 PR
‘Jubileu’ 1998 P 300 NM Y 127 PR
‘Leonense’ 1998 P 275 NM Y 128 PR
‘Linda’ 1989 N 400 M Y 95 FR
‘Maciel’ 1992 P <300 NM Y 136 DU
‘Marfim’ 1999 P 350 M W 120 FR
‘Marli’ 1984 P 300 M W 108 FR
‘Olimpia’ 2004 P 300 NM Y 150 DU
‘Onix’ 1985 P 300 NM Y 122 PR
‘Pampeano’ 1993 P 175 M W 86 FR
‘Pepita’ 2000 P 150 NM Y 91 PR
‘Pilcha’ 1985 P 400 M Y 116 FR
‘Planalto’ 1992 P 400 NM W 101 FR
‘Precocinho’ 1981 P 150 NM Y 101 PR
‘Riograndense’ 1991 P <300 NM Y 113 DU
‘Santa Aurea’ 2004 P 400 NM Y 122 DU
‘Sensacao’ 2004 P 200–300 NM Y 107 DU
‘Sentinela’ 1985 P 150 M W 98 FR
‘Turmalina’ 1999 P 350 NM Y 111 PR
‘Vanguarda’ 1989 P <150 NM Y 102 PR
Infruitec, Stellenbosch, South Africa
‘Alpine’ (p)g 1997 N L M Y Late Nov FR
‘Classic’ (p) 1994 P M NM Y Early Dec FR
‘Crimson Giant’ (p) 1995 N L M Y Mid-Dec FR

(Continued)
 Low-chill Cultivar Development 115

Table 5.5. continued

Year of Chillb Flesh Flesh FDPe

Cultivar introduction Cropa (CU) typec colourd (days) Purposef

‘Donnarine’ (p) 1987 N L M Y Mid-Dec FR

‘Elandia’ (p) 1998 P L-M M Y Late Jan FR
‘Escellence’ (p) 1995 P L-M M Y Late Dec FR
‘Marg. Pride’ (p) 1991 N L M Y Mid-Dec FR
‘Novadonna’ (p) 1990 P L-M NM Y Late Nov FR
‘Olympia’ (p) 1985 N M M Y Mid-Dec DU
‘Oribi’ (p) 1993 P M M Y Late Feb DU
‘Summersun’ (p) 1995 P L NM Y Late Nov DU
‘Transvalia’ (p) 1987 P L NM O-Y Mid-Nov FR
‘Unico’ (p) 1996 N L M Y Mid-Nov FR
‘Western Cling’ (p) 1994 P L NM Y Early Dec FR
Instituto Agronomico (IAC), Sao Paulo, Brazil
‘Arlequim’ 1981 P <50 M W 180 FR
‘Aurora-1’ 1987 P <50 NM Y 110 FR
‘Aurora-2’ 1987 P <50 NM Y 118 FR
‘Branca De Guapiara’ 1984 N <50 M W 115 FR
‘Canario’ 1982 P <50 M Y 128 FR
‘Catita’ 1982 P <50 M W 150 FR
‘Catuiba’ 1982 P <50 M Y 125 FR
‘Centenaria’ 1987 N <50 M Y 108 FR
‘Centenario’ 1987 P <50 M Y 115 FR
‘Delioso Precoce’ 1987 P <100 M W 107 FR
‘Docura’ 1980 P <50 M W 135 FR
‘Docura-2’ 1983 P <50 M W 115 FR
‘Docura-3’ 1983 P <50 M W 120 FR
‘Docura-4’ 1987 P <50 M W 120 FR
‘Douradão’ 1998 P <100 M Y 105 FR
‘Dourado-1’ 1985 P <50 M Y 108 FR
‘Dourado-2’ 1985 P <50 M Y 115 FR
‘Joia-1’ 1983 P <50 M W 100 FR
‘Joia-2’ 1983 P <50 M W 105 FR
‘Joia-3’ 1987 P <50 M W 110 FR
‘Joia-4’ 1987 P <50 M W 103 FR
‘Joia-5’ 1987 P <50 M W 105 FR
‘Josefina’ 1986 N <50 M W 112 FR
‘Momo’ 1981 P <50 NM Y 185 DU
‘Ouromel-2’ 1983 P <50 M Y 110 FR
‘Ouromel-3’ 1983 P <50 M Y 108 FR
‘Ouromel-4’ 1983 P <50 M Y 112 FR
‘Perola De Mairinque’ 1980 P <50 M W 160 FR
‘Petisco-2’ 1983 P <50 M Y 145 FR
‘Precoce De Itupeva’ 1984 N <50 M W 115 FR
‘Regis’ 1987 N <50 M Y 95 DU
‘Rosalina’ 1984 N <50 M Y 120 DU
‘Sol Do Vale’ 1982 P <50 M W 145 FR
‘Somel’ 1984 N <50 M W 120 FR
‘Tropical’ 1989 P <50 M Y 80 FR
‘Tropical-2’ 2002 P <50 M Y 78 FR
Instituto Nacional de Investigaciones Forestales y Agropecuarias (INIFAP), Mexico
‘Cengua-Guia’ 1990 P 250 M W 160 RO
‘Comonfort’ 1992 P 250 NM Y 130 FR

(Continued)
116 B.L. Topp et al.

Table 5.5. continued

Year of Chillb Flesh Flesh FDPe

Cultivar introduction Cropa (CU) typec colourd (days) Purposef

‘Dorado’ 1992 P 250 NM Y 110 FR

‘Regio’ 1992 P 200 NM Y 130 FR
‘Rendidor’ 1992 P 300 NM Y 160 FR
‘Seleccion 165’ 1980 P 300 NM Y 150 GE
Sun World International, Inc., Bakersfield, California, USA
‘Supechfifteen’ (p) 2002 P 150 M Y 5 Apr FR
‘Supechthirteen’ (p) 2002 P 150 M Y 1 Apr FR
Taiwan Agricultural Research Institute, Taichung, Taiwan
‘SpringHoney’ 2003 P 180 M W 82 FR
Texas A&M University, Weslaco, Texas, USA
‘Earligrande’ 1979 P 250 M Y 75 FR
‘TropicPrince’ (p) 2001 P 150 M Y <90 FR
‘ValleGrande’ 1990 P 250 M Y 85 FR
University of Florida, Gainesville, Florida, USA
‘Carolina’ 1990 N 325 M Y 90 FR
‘Desertred’ 1983 P 250 M Y 90 FR
‘Diamante Especial’ 1995 P 250 NM Y 120 FR
‘Fla.85-1’ 1995 P 400 M Y 67 GE
‘Flordadawn’ 1989 P 250 M Y 60 FR
‘Flordagem’ 1983 P 250 M Y 85 FR
‘Flordaglo’ 1988 P 150 M W 85 FR
‘Flordaguard’ 1991 P 300 M Y 130 RO
‘FlordaMex 1’ 1989 P 400 M Y 95 FR
‘Flordaprince’ 1982 P 150 M Y 80 FR
‘FlordaRio’ 1994 P 400 M Y 92 FR
‘Flordastar’ 1988 P 225 M Y 72 FR
‘Forestgold’ 1991 P 350 M Y 95 FR
‘Hermosillo’ 1984 P 300 M Y 108 FR
‘Newbelle’ 1984 P 150 M Y 105 FR
‘Opedepe’ 1982 P 150 M Y 85 FR
‘Oro A’ 1989 P 250 NM Y 83 FR
‘Rayon’ 1982 P 175 M Y 105 FR
‘Sherman’s Red’ 1985 P 300 M Y 75 FR
‘SunBest’ (p) 2002 N 225 M Y 90 FR
‘Sunblaze’ 1985 N 250 M Y 90 FR
‘Sunbob’ 1989 N 200 M Y 100 FR
‘Suncoast’ 1994 N 400 M Y 77 FR
‘Sundollar’ 1989 N 400 M Y 70 FR
‘Sundowner’ 1987 N 250 M Y 90 FR
‘Sunhome’ 1985 N 300 M Y 88 FR
‘Sunmist’ 1994 N 300 M W 80 FR
‘Sunraycer’ 1994 N 275 M Y 85 FR
‘Sunsnow’ 1990 N 250 M W 90 FR
‘Sunsplash’ 1993 N 400 M Y 74 FR
‘SunWright’ 1991 N 200 M Y 80 FR
‘UF2000’ (p) 2000 P 300 NM Y 95 FR
‘UFBeauty’ (p) 2003 P 200 NM Y 83 FR
‘UFCharm’ (p) 2000 P 250 NM Y 80 FR
‘UFGold’ (p) 1996 P 225 NM Y 80 FR
‘UFO’ (p) 2000 P 250 NM Y 95 FR

(Continued)
 Low-chill Cultivar Development 117

Table 5.5. continued

Year of Chillb Flesh Flesh FDPe

Cultivar introduction Cropa (CU) typec colourd (days) Purposef

‘UFQueen’ (p) 1998 N 250 NM Y 97 FR

‘UFSun’ (p) 2003 P 100 NM Y 103 FR
‘White Opal’ 1998 P 200 M W 75 FR
‘White Satin’ 2000 P 250 M W 75 FR
‘Zorrito’ 1985 P 275 M Y 95 OR
University of Florida and Texas A&M University, USA
‘FlordaGrande’ 1985 P 50 M Y 105 FR
‘Tropic Blush’ 1990 P 200 M Y 90 FR
‘TropicBeauty’ 1988 P 150 M Y 85 FR
‘TropicSnow’ 1988 P 250 M W 93 FR
‘TropicSweet’ 1986 P 175 M Y 90 FR
University of Georgia, US Department of Agriculture and University of Florida, USA
‘Gulfking’ (p) 2004 P 350 NM Y 73–80 FR
‘Gulfprince’ (p) 2000 P 375 NM Y 94 FR
University of Western Sydney, Richmond, New South Wales, Australia
‘Dawn Gold’ (p) 2003 N 200–300 M Y 109 FR
‘December Ice’ (p) 2003 N 350 M W 129 FR
‘Hail’ (p) 2003 N 200–300 M W 106 FR
‘Honey Ice’ (p) 2003 N 400 M W 128 FR
‘Pale Ice’ (p) 2003 N 250–300 M W 113 FR
Zaiger Genetics, Inc., Modesto, California, USA
‘April Glo’ (p) 1990 N 150 M Y <90 FR
‘Earliglo’ (p) 1990 N 150 M Y <90 FR
‘Earlitreat’ (p) 1997 P 300 M Y <90 FR
‘Evas Pride’ (p) 1991 P 250 M Y <90 FR
‘Mayglo’ (p) 1984 N 200 M Y <90 FR
‘Red Roy’ (p) 2001 N 300 M Y <90 FR
‘Snow Angel’ (p) 2004 P 250 M W <90 FR
‘Zee Fire’ (p) 2003 N 250 M Y <90 FR
aCrop: P, peach; N, nectarine.
bCultivar chilling requirement in chill units (CU); or L, low-chill; M, moderate-chill.
cFlesh type: M, melting; NM, non-melting.
dFlesh colour: O, orange; W, white; Y, yellow.
eFDP: fruit development period in days; or harvest date at place of origin.
fPurpose: DU, dual purpose of fresh and processing; FR, fresh; GE, germplasm; OR, ornamental;

PR, processing; RO, rootstock.

g(p), patented.

have drawn heavily on parents from UF but
have also incorporated local subtropical peaches
(Ward, 1952) and germplasm from Brazil, Mex-
ico, California, Georgia, Vietnam and China.
Brazil, Pelotas
The breeding programme of the Brazilian Agri-
cultural Research Enterprise (Embrapa Clima
Temperado) is located at the Agriculture
Research Centre of Temperate Crops (CPACT)
at Pelotas, Rio Grande do Sul. It began in 1953
and has released 21 fresh market peach, six
nectarine and 25 dual-purpose (fresh and pro-
cessing) peach cultivars requiring 150–500
CU. The peach harvest in Rio Grande do Sul
has been extended from 15 days in 1963, when
there were only two cultivars (‘Aldrighi’, a
seed-propagated clingstone and ‘Delicioso’, a
118 B.L. Topp et al.
white-fleshed peach) grown, to over 100 days
with the current releases (Raseira et al., 1992).
The main objectives are for adaptation to 150–
500 CU but with emphasis on 200–300 CU,
productivity, resistance to brown rot, bacterial
spot and frost, and acceptable fruit size, shape,
firmness and flavour (Raseira et al., 2003).
Temperatures during the winter bloom
period are highly variable in southern Brazil
and this contributes to variation in yield. Cul-
tivars such as ‘Esmeralda’ consistently pro-
duce heavy crops despite these conditions;
the reason for this cultivar’s specific yield sta-
bility is an area of ongoing research.
Brazil, Sao Paulo
This programme began in 1950 at the Instituto
Agronomico (IAC) in Sao Paulo. The aim of
the programme is to produce peaches requir-
ing 50–150 CU. Initial germplasm included
‘Jewel’, ‘Suber’, ‘Hall’s Yellow’, ‘Lake City’
and ‘Angel’ imported from the USA, plus a
number of locally selected seedlings probably
derived from Portuguese introductions from
the 16th century (Barbosa et al., 1997). These
were crossed with medium-chill North Amer-
ican germplasm. From 1970 to 2000 the best
IAC cultivars were intercrossed with selec-
tions from UF. Emphasis has been on fresh
market fruit or fruit that can serve the dual
purpose of canning and fresh market.
Fifty-eight cultivars have been released,
all needing 50–200 CU, and account for about
1.2 million of the 1.9 million trees grown in
the state of Sao Paulo. The remaining culti-
vars are releases from UF and EMBRAPA.
IAC cultivars cover the full harvest season
with FDP of 80–180 days. Future breeding
will aim to produce compact tree structure,
ultra-early ripening, extra-large fruit size,
very attractive and firm fruit, improved dis-
ease and pest resistance and tolerance to
warmer climates.
Mexico, Chapingo
The breeding programme of the Centro de
Fruiticultura, Colegio de Postgraduidos at
Chapingo started in 1985 with the main objec-
tive being to develop low-chill peaches over a
wide range of FDP with resistance to pow-
dery mildew. Breeders at UF and Chapingo
have cooperated closely, sharing and testing
germplasm and jointly releasing the full
yellow-skin, non-melting flesh peach ‘Oro A’
in 1989 (this cultivar has found a niche in
Australia, where it is marketed as a ‘peachcot’
owing to its colour, size and low fuzz). The
Chapingo programme almost exclusively
produces non-melting peaches for the fresh
market. Mexican consumers associate high
sugar and good flavour with mostly yellow-
skinned criollo peaches that have only a small
amount of red blush and so the breeding pro-
gramme has selected for this appearance. A
series of mildew-resistant peaches has been
released and now the programme is focusing
on developing resistance to Monilinia (partic-
ularly the blossom blight phase of the dis-
ease). Adaptation is sought mainly in the
150–450 CU range, but since 1993 there has
been some breeding for 500–700 CU geno-
types for the peach industry in the Aguascali-
entes and Zacatecas regions.
Mexico, Queretaro
Salvador Perez at the Universidad Autonoma
de Queretaro has studied the phenology and
adaptation of feral Mexican and imported
peach seedling populations across Mexico in
order to define breeding strategies for the
subtropics (Perez, 1989, 2003, 2004; Perez
et al., 1993; Perez-Gonzalez, 2001, 2002). Sev-
eral cultivars were released from the Instituto
Nacional de Investigaciones Forestales y Agr-
opecuarias (INIFAP) breeding programme
(Table 5.5) with emphasis on developing resis-
tance to powdery mildew in non-melting flesh
peach needing 50–450 CU. The focus now is
on fruit quality in non-melting flesh types for
the fresh market, in both yellow and white
flesh. Fruit must have low to medium acidity
with <0.6 mEq malic acid, high sugar with >15%
total soluble solids (TSS), medium size (120–180
g), round to flat shape and from bright orange
to red skin covering 80% of the surface.
Seedlings are grown on commercial
orchards rather than at a research station, as
 Low-chill Cultivar Development
occurs with most other programmes. This
was done because the university research plot
was located far from the commercial sites and
it allows screening of higher-chilling seedling
populations at high-elevation orchards. From
100 to 2000 seedlings are produced each
year. Advanced selections are tested for 5
years prior to public release. The most recent
release is ‘Fred’ (‘Lucero’ × ‘Springcrest’) in
2000, which is a 450 CU, non-melting peach
with excellent quality and resistance to flesh
browning.
South Africa, Stellenbosch
The Agricultural Research Council (ARC)
Infruitec–Nietvoorbij breeding programme
started in 1937 and caters for the South Afri-
can canning industry, dried fruit industry and
fresh market, both domestic and export. Inad-
equate chilling of traditional high-chill peach
cultivars is a problem in the early-maturing
region in South Africa and so breeding has
concentrated on developing cultivars with a
chilling requirement of <400 CU. Specific
breeding objectives are high skin over-colour,
development of >80% red blush, round fruit
with no tip or suture, high flesh firmness,
high sugar to acid ratio, >13% TSS, fine tex-
ture, absence of split pits and a minimum
cold storage life of 4 weeks.
Approximately 3000 hybrid seedlings
are produced each year, with about two-thirds
being low-chill (<400 CU) early types. Selec-
tion is for peach and nectarine, yellow and
white flesh and both melting and non-melting
flesh. Advanced selections are planted in
semi-commercial trials prior to release. This
programme accounts for 95% of all processed
peach, 90% of fresh market peach and 60% of
fresh market nectarine cultivars grown in
South Africa.
Taiwan, Taichung
The Taiwan Agricultural Research Institute
began breeding in 1980 for peach, nectarine
and now peento cultivars for the local mar-
ket. High-chill peach cultivars from Japan
119
were introduced to the highlands (>1500 m
above sea level) in the 1950s, but soil erosion
on the steep slopes led the Taiwan govern-
ment to commence breeding for low-chill cul-
tivars that could be grown at lower elevations
(locations receiving <250 CU). New cultivars
must be white-fleshed, low- or sub-acid, with
high juiciness and high sugar level. The stony-
hard flesh ‘Ing-go’ is being used as a standard
for firmness as are the non-melting flesh
genotypes from Brazil and Florida (Wen and
Sherman, 2002). ‘Tainung No. 1’ peach was
selected from a cross between the Brazilian
peach ‘Premier’ and the Florida peach ‘Flordak-
ing’ and named in 2001. More recently
‘SpringHoney’ peach was released to fill the
need for a lowland, subtropical cultivar that
produces early-ripening, low-acid, white,
melting-flesh fruit (Ou and Wen, 2003). It was
derived from a population of ‘Premier’ ×
‘Flordabelle’.
Thailand, Angkhang
Low-chill peach breeding by the Royal Proj-
ect Foundation and Kasetsart University
commenced in 1997. Test plots are located in
the mountainous regions of northern Thai-
land at the Royal Angkhang Research Station
and also at Khunwang. The tropical highland
peach-growing regions are characterized by
long growing season, high rainfall and mild
winters. Objectives are for 100–300 CU culti-
vars, ripening during the dry season from
March to June, and fruit with bright red skin,
round shape, white and yellow flesh and
intense flavour including both low- and high-
acid types. The breeder works closely with
and is testing advanced selections from the
Texas A&M University (TAMU) programme.
USA, California, Sun World
Sun World International, Inc. has been actively
involved in low-chill peach breeding since
1987 when the first large-scale seedling popu-
lations were planted in the Coachella Valley
of southern California. In this region there is
often less than 300 CU and trees are exposed
120 B.L. Topp et al.
to prolonged periods above 38°C during much
of summer and into autumn. The goal was to
develop cultivars adapted to these conditions
that ripen in the first half of April, before the
lower San Joaquin Valley harvest begins. The
programme initially used UF selections as
parents both in combination with commercial
Californian cultivars and mated inter se. About
3000 seedlings from low-chill × low-chill crosses
are planted annually in the Coachella Valley
and about 2500 seedlings annually from low-
chill × medium-chill crosses are planted near
Bakersfield. Sun World also has an active
breeding programme for low-chill plums and
a small effort for low-chill apricot and root-
stocks.
High temperatures in spring in the
Coachella Valley result in FDPs up to 20 days
less than for Bakersfield in the San Joaquin
Valley. The short FDP and high temperatures
in Coachella mean that small fruit size, low
fruit firmness, prominent tips and low or
inconsistent bloom require high selection
pressure in the breeding programme.
New cultivars are patented if they are to
be grown commercially in the USA or distrib-
uted to other countries where they are licensed
to growers on a production–royalty basis.
‘Supechthirteen’ and ‘Supechfifteen’ are pat-
ented peach cultivars grown commercially by
Sun World in the Coachella Valley.
USA, California, Zaiger Genetics
Zaiger Genetics, Inc. has released a number of
low-chill peach and nectarine cultivars, with
‘Mayglo’ nectarine in 1984 being one of their
first. Low-chill seedlings are selected at high-
chill test plots at Modesto, California. The pri-
mary goal of these selections is for very early
ripening in California. Creating earlier ripen-
ing through earlier flowering (lower chill) is
possible in temperate climates such as Cali-
fornia where spring frosts do not occur.
Low-chill selections made in this type of
environment require testing in the subtropics
to screen for problems associated with grow-
ing peaches in these locations (blind nodes,
pointy fruit shape often accompanied by suture
bulges, uneven ripening and bud drop). All
cultivars from this private breeding progra-
mme are protected by plant patent.
USA, Florida, University of Florida
Since its inception in 1953, the programme
has released over 40 cultivars of peach and
nectarine that are being tested in 76 countries
and territories and grown commercially in 23
countries. As this programme has been active
for the past 50 years, many of the cultivars
have been replaced by newer ones. Advanced
germplasm from the UF programme has been
used as parental material in most low-chill
peach breeding programmes. In addition,
breeders from Australia, Mexico, Spain, Tai-
wan and the USA have studied at the UF pro-
gramme and developed breeding philosophies
and techniques from this association. Breed-
ers from many other low-chill programmes
have benefited from close interaction with the
UF programme. The original objectives at UF
were to produce a series of peach and nectar-
ine cultivars adapted to Florida and bearing
high-quality, melting-flesh fruit with short
FDP of <100 days (Sherman and Rodriquez-
Alcazar, 1987). The ‘Florda’ series of peaches
and ‘Sun’ series of nectarines comprise this
group of cultivars.
More recently, the UF programme has
changed to breeding non-melting peaches
and nectarines in order to improve flavour
(Sherman et al., 1990). The idea is that with
firmer, non-melting flesh, growers can leave
fruit on the tree closer to its physiological full-
ripe stage and still have sufficiently firm fruit
to market. After incorporating the non-melt-
ing flesh gene from Mexican, North Carolina
and Brazilian germplasm, the breeders have
selected heavily to remove undesirable traits
of long FDP, high chill, lack of red skin, off-
flavours in over-ripe fruit (Karakurt et al.,
2000) and low acidity. The non-melting flesh
series of peach and nectarine cultivars are
prefixed with ‘UF’ (‘UFSun’, ‘UFQueen’, etc.).
Main selection criteria are for either white or
yellow non-melting flesh, more than 11% TSS,
higher red skin over-colour, higher aroma
and flavour, less twiggy (selecting for thicker
and fewer fruiting laterals) tree structure with
high flower bud density, few blind nodes, low
 Low-chill Cultivar Development
bud drop, and fruit set with night tempera-
ture above 14°C. Since 1992, UF has acquired
plant breeding rights on all new cultivars.
Additionally, UF, the University of Geor-
gia and the US Department of Agriculture
cooperate in a breeding programme at Atta-
pulgus, Georgia to produce mid-chill cultivars
requiring 400 to 650 CU but with some in the
250 to 400 CU range (Beckman et al., 1995;
Krewer et al., 1998). Cultivars released from
this programme are named with the prefix
‘Gulf’, for example ‘Gulfking’ and ‘Gulfprince’.
USA, Texas, Texas A&M University
The TAMU stone fruit breeding programme
commenced at College Station in 1939 and is
selecting both medium- (350–650 CU) and
low-chill (<350 CU) cultivars. In the 1980s,
collaborative work with UF resulted in joint
releases of ‘Flordagrande’, ‘TropicBeauty’,
‘TropicSnow’ and ‘TropicSweet’. Low-chill
breeding continues with evaluation at the
South Texas Research and Extension Center
at Weslaco and at sites in Florida, California,
Mexico, Thailand and Uruguay. An initial objec-
tive of this work was to develop peach culti-
vars ripening after ‘Flordaprince’ and before
‘TropicBeauty’. The first release is ‘Tropic-
Prince’, a yellow-fleshed, melting peach,
ripening after ‘Flordaprince’.
During the 1990s, TAMU began to
develop business collaborations with Burchell
Nursery in the USA and several organizations
in Spain, Australia and Uruguay to better uti-
lize diverse genotypes in breeding and to
generate funding. These collaborations have
allowed the TAMU programme to expand the
breeding objectives beyond yellow-fleshed
peaches to include work on non-melting,
white-fleshed, low-acid, high soluble solids
nectarines and peento materials.
5.3 Common Objectives in Low-chill
Peach Breeding
Objectives of low-chill peach breeding over-
lap to a large extent with those for high-chill
peach. Successful new cultivars must combine
121
characteristics that make the trees well adap-
ted and productive, and the fruit desired by
consumers. Many of the selection traits that
are common in both high- and low-chill breed-
ing programmes have been described previ-
ously (Scorza and Okie, 1990; Scorza and
Sherman, 1996; Byrne et al., 2000). The follow-
ing discussion concentrates on those traits
that require particular selection emphasis as a
consequence of working with low-chill geno-
types.
Low chilling requirement
The effects of chilling in breaking leaf and
flower bud endo-dormancy are not well
understood and this is reflected by the variety
of methods used for measuring chilling
requirement. Weinberger (1950) defined 1
chill-hour as 1 h at or below 7.2°C (45°F); a
peach cultivar with a chilling requirement of
800 chill-hours would need 800 h below 7.2°C
to break dormancy. Richardson et al. (1974)
presented the ‘Utah’ model which ascribes
units of chill and allowed for negation of
chilling by high temperatures. In this model,
1 chill unit is the maximum amount of chill-
ing that can occur at the optimum tempera-
ture with partial chilling accumulated at less
optimum temperatures. Erez et al. (1990) pro-
posed a dynamic model that allows for nega-
tion of chilling but only by high temperatures
occurring immediately after the chilling. In
low-chill regions, where the daytime winter
temperatures are often in the chill negation
range, the dynamic model appears to be more
effective than the Utah or chill-hour model
(Allan et al., 1995). The Weinberger–Sharpe
model (Sharpe et al., 1990) uses average monthly
winter temperatures to predict how cold the
winter was and thus the relative amount of
winter chilling accumulated. This model is
useful for predicting location chilling accumu-
lation in order to choose cultivars with similar
chilling requirements for cultivar testing.
Breeders require a method of measuring
the chilling requirement of genotypes. One
method is to use controlled screening tests
whereby whole trees, rooted cuttings, excised
shoots or single nodes are forced to flower in
122 B.L. Topp et al.
warm glasshouses after exposure to known
amounts of chilling temperatures (Dennis,
2003). These tests are expensive and it is not
feasible to incorporate them into a breeding
programme that requires evaluation of thou-
sands of genotypes. In practice, peach breed-
ers use bloom date as an indicator of chilling
requirement with the lowest-chill genotypes
flowering first (Scorza and Sherman, 1996).
Standard cultivars are used as comparators.
In the UF programme the standards used are
‘Okinawa’ (150 CU), ‘Sunred’ (250 CU), ‘Early
Amber’ (350 CU) and ‘Sunlite’ (450 CU) (Sher-
man and Lyrene, 1998).
Several authors have reported on proge-
nies in which the mean chilling requirement
of the seedlings was statistically the same as
the mid-parent mean (Lesley, 1944; Lammerts,
1945; Sharpe, 1961; Bassols, 1973). The con-
clusion is that chilling requirement is a quan-
titatively inherited character with genes
having a cumulative, similar effect. However,
there is some evidence for one or a few genes
with major effects (Scorza and Sherman,
1996), skewing the distribution curve of the
progeny towards the lower-chilling parent.
In the Brazilian breeding programme it
has been possible to produce seedlings with
good adaptation to mild winters in the first
generation of crossing between low-chill Bra-
zilian seed parents and high-chill Canadian
and US pollen parents. The low-chill seed
parents that have been efficient in transfer-
ring lower chilling requirement to their prog-
eny are ‘Maciel’, ‘Precocinho’ and ‘Diamante’,
and particularly ‘Cerrito’ and ‘Sunred’ (M.C.B.
Raseira, Brazil, 2004, personal communication).
Time of flowering
Time of flowering depends on two factors: the
CU necessary to complete rest and the grow-
ing degree-hours (GDH) required after endo-
dormancy to reach full bloom (Richardson
et al., 1974; Citadin et al., 2001). Breeders may
wish to select for late bloom to reduce the risk
of spring frost damage. If there was variabil-
ity for both CU and GDH in peach then it
should be possible to produce late-flowering
cultivars by increasing the CU requirement or
the GDH requirement or both. In low-chill
locations, the CU requirement cannot be
increased a great deal without reduction in
productivity that occurs with inadequate
chilling and loss of adaptation.
It is difficult and expensive to measure
the separate components of CU and GDH,
and there is still much to understand in terms
of the critical threshold temperatures that are
needed for their accumulation (Felker and
Robitaille, 1985). A pragmatic solution is to
select for late bloom and productivity at the
same time. In this way, the selected genotypes
are not late blooming due to an increased CU
requirement that makes them poorly adapted.
Selection for late-blooming genotypes in a
medium- to high-chill environment that reli-
ably set high crop loads was examined by
Souza et al. (2000) in a population of low- and
medium-chill peaches. The cultivars ‘Sun-
land’ and ‘Gaschina Novembre’ had high
breeding values for late bloom and high fruit
density, indicating they would be suitable
parents for making genetic progress for both
traits.
Working with 1311 low-chill genotypes
belonging to 30 different progenies, Quezada
et al. (2000) found extremely high values for
both broad sense and narrow sense heritabil-
ity in blossom date (close to 1), and concluded
that predictable progress could be achieved
based on parents’ behaviour. Souza et al.
(1998a) also reported a high heritability of
0.78 for full bloom date in a medium-chill
peach population at the TAMU programme.
In this population, selection of the 5% of gen-
otypes with earliest bloom and inter-mating
for one generation would decrease the aver-
age bloom date of the population by 14 days.
Most late-flowering peach genotypes are
not productive when grown in low-chill loca-
tions due in large part to inadequate chilling,
but possibly also due to failure to set under
high night temperatures (Edwards, 1987;
Rouse and Sherman, 2002a). There are reports
of low-chill genotypes that bloom late such as
some Mexican germplasm from Aguascalien-
ties (Scorza and Okie, 1990) and the cultivars
‘BR-1’, ‘Delicioso’ and ‘Della Nona’ from Bra-
zil (Citadin et al., 2001). There also appear to
be some cultivars, such as ‘Flordaprince’ and
‘TropicBeauty’, that are able to set crops under
 Low-chill Cultivar Development 123

high (16–18°C) night temperatures (Rouse reduced productivity. Formation of blind nodes
and Sherman, 2002a). There may be scope for was observed to occur most frequently when
selecting for late bloom using this type of ger- temperatures were high and tree growth was
mplasm that has high heat requirements to low (Boonprakob and Byrne, 2003). There is
flower and ability to set later in the flowering great genotypic variability for this trait (Boon-
season when nights are warmer. prakob et al., 1994; Richards et al., 1994) and
several cultivars exist with low blind node
propensity (Table 5.6).
Flower bud density Bud drop in peaches has high genetic
variability and is usually associated with
highly fluctuating winter temperatures or
Flower bud density is of importance in regions
high mean minimum mid-winter tempera-
subject to spring frost damage. High flower
tures. In a study of 13 peach cultivars over 10
bud density can provide insurance against
years at Fresno, California, Weinberger (1967)
crop loss in that buds of different develop-
concluded that a mid-winter mean minimum
mental stages have varying threshold mortal-
above 4.4°C was critical for appearance of
ity temperatures (Proebsting and Mills, 1978).
bud drop in susceptible cultivars. Temperate
Genotypes with high flower bud density will
peach cultivars bred in uniformly cool winter
have a range of undeveloped buds at a given
climates often transmit bud drop when
date when frost occurs.
hybridized into a low-chill programme, thus
The medium-chill peach ‘Texstar’ (450
selection must be practised where winters are
CU) sets heavy crops under spring freeze
not uniformly cool.
conditions. It produces ten times the number
of flower buds necessary for a commercial
crop and loses about 70% during bloom in
years with and without freezes (Byrne, 1986). Fruit development period
This loss was greater than other clones that
failed to set commercial crops during the
Almost all low-chill breeding programmes
freeze, but was compensated by the very high
have short FDP as a major objective in order
flower bud density of ‘Texstar’.
to produce early-ripening cultivars. In many
Peach and nectarine cultivars from the UF
subtropical locations, early ripening is
breeding programme generally have high
required to ensure the crop is harvested prior
flower bud density, which was inherited from
to the onset of the summer rainy season and
‘Okinawa’, one of the original low-chill parents
the associated problems with disease and
(Sherman and Lyrene, 2003). The high flower
fruit decay. At Gainesville, Florida, the rainy
bud density is necessary to ensure regular
season generally commences in the second
crops after Florida’s frequent spring freezes. In
week of June, which means that all the
locations such as coastal northern New South
adapted 350 CU genotypes have FDP of <100
Wales and Queensland in Australia, spring
days (Sherman and Rodriguez-Alcazar, 1987).
freezes seldom occur and the high flower bud
Independent of this reason, early ripening is
density of UF cultivars is an expensive burden
of economic importance because of the high
to orchardists because of the high cost of hand-
fruit prices for early-season fruit.
thinning fruitlets. Flower density (0.41) and
Length of FDP and time of ripening are
node density (0.48) were reported to have
highly heritable traits in low-chill and medium-
moderate heritability, which would allow mani-
chill peach germplasm (Souza et al., 1998b)
pulation of either trait (Souza et al., 1998a).
and are readily altered by selection. The
occurrence of bud sports ripening 7–10 days
earlier than the original clone and the pres-
Blind nodes and bud drop ence of redleaf markers (Fig. 5.2/Plate 49) in
early-ripening cultivars (Sherman et al., 1972)
Blind nodes are nodes that lack floral and indicate there may be genes with a major
vegetative buds and are therefore a cause of effect on ripening time. The earliest-ripening
124 B.L. Topp et al.

Table 5.6. Blind node propensity of peach and nectarine clones tested at Gainesville, Floridaa and
College Station, Texasb.

Clonea Blind nodes (%) Cloneb Blind nodes (%)

‘Fla.4-4’ 10 ‘Fla.1-8’ 11
‘Rayon’ 28 ‘Desertred’ 16
‘TropicSweet’ 31 ‘Flordaking’ 33
‘Sunred’ 32 ‘Gulfpride’ 36
‘Sunhome’ 41 ‘Earligrande’ 37
‘Sunlite’ 46 ‘Loring’ 38
‘Sundollar’ 50 ‘Goldcrest’ 39
‘Newbelle’ 64 ‘Sunhome’ 40
‘Oro A’ 69 ‘P51-2’ 41
‘Fla.90-44C’ 84 ‘Sentinel’ 42
‘BY3-1197’ 44
‘Junegold’ 45
‘Sunland’ 45
‘Elberta’ 47
‘Juneprince’ 51
‘BY4-7124’ 56
‘BY5-938’ 59
‘BY3-600’ 60
‘Springcrest’ 66
‘Cherrygold’ 81
aFrom Richards et al. (1994).
bFrom Boonprakob et al. (1994).

Fig. 5.2. Regular green leaf peach seedlings in foreground and peach seedling displaying redleaf
character associated with short fruit development period (FDP) in centre at Nambour, Queensland,
Australia. This trait can be used for early identification of short FDP seedlings in late summer. (From
Sherman et al., 1972.)
 Low-chill Cultivar Development 125

low-chill cultivars have FDP of the order of 60
to 70 days (Table 5.7) and so can still be decreased
significantly when compared with high-chill
cultivars such as ‘Goldcrest’ that have FDP of 55
days (Ramming and Tanner, 1987).
Fruit shape
Symmetrical and rounded fruit are desirable
because they result in less damage during
harvest and handling. Production of fruits
with large stylar tips and/or suture bulges is
a common problem in mild winter regions
(Fig. 5.3/Plate 50). For a single cultivar, the tip
size and suture bulge may vary across loca-
tions and years, with larger tips and bulges
more frequent in warmer locations (Topp and
Sherman, 1989). Some genotypes that are round
in cool locations become pointed when grown
in warmer regions (Fig. 5.4/Plate 51) but others
such as ‘Sunlite’ nectarine remain relatively
tip-free at all locations. Heritability estimates

Table 5.7. Low-chill peach and nectarine cultivars with short fruit development period (FDP)a.
(From Okie, 1998.)

Cultivar Crop Chilling (CU) FDP (days)

‘Flordadawn’ Peach 250 60

‘Sherman’s Early’ Peach 425 60
‘Flordaglobe’ Peach 475 62
‘Fla.85-1’ Peach 400 67
‘Flordaking’ Peach 400 69
‘Sundollar’ Nectarine 400 70
‘Flordastar’ Peach 225 72
aDays from flowering to harvest.

Fig. 5.3. Peach fruit with prominent stylar tips that are commonly observed in cultivars with poor
adaptation to mild winter locations and are selected against in low-chill breeding.
126 B.L. Topp et al.

Fig. 5.4. Influence of location on fruit shape. This medium-chill (450 CU) peach selection
produced: (a) round fruit at the high-chill (900 CU) location of Stanthorpe, Queensland, Australia
and (b) pointed fruit at the low-chill (200 CU) location of Nambour, Queensland, Australia.

of 0.45 and 0.43 for fruit tip and fruit shape
rating, respectively, indicate that genetic
advance for these traits is possible (Souza
et al., 1998b). Some recent cultivars from Flor-
ida, Texas and southern Brazil are examples
of the achieved goal for round fruit shape.
Fruit firmness
In traditional melting-flesh genotypes, firmness
is mostly a measure of evenness of ripening
within a fruit at good red over-colour and
yellow ground colour development. Thus, to
 Low-chill Cultivar Development
increase firmness, especially in genotypes with
short FDP (<100 days) where firmness is lack-
ing compared with genotypes with FDP of
120–180 days, breeders have selected for
advanced red skin and yellow ground colour
while the fruit is physiologically immature,
resulting in low-brix fruit with low flavour
or aroma. Two flesh types, non-melting and
stony-hard, offer potential to overcome this
problem by allowing the fruit to mature (tree
ripen) with higher brix and aroma while
retaining firmness for shelf-life.
Both the non-melting and stony-hard
flesh traits are controlled by single recessive
genes (Scorza and Sherman, 1996). Non-melting
flesh is characterized by absence of the endo-
polygalacturonase enzyme that is responsible
for flesh softening in melting peaches (Lester
et al., 1996). Mature fruit with non-melting
flesh soften at a slower rate than those with
melting flesh. Stony-hard flesh fruit produce
no ethylene and so stay firm unless exposed
to exogenous ethylene (Haji et al., 2003). It
will be important to study consumer reaction
to both these flesh types in developing firm
peaches that are suited to particular markets
(Brovelli et al., 1999; Williamson and Sargent,
1999).
The non-melting flesh trait is present in
all the recent cultivar releases in the low-chill
breeding programmes at Chapingo and Que-
retaro in Mexico and at UF in the USA. Since
the 1980s, Brazil has released dual-purpose
low-chill cultivars with non-melting yellow
flesh. The stony-hard flesh trait is present in
old low-chill peach cultivars in China and
Taiwan which generally produce fruit that are
white-fleshed, small and with long FDP. Ou
and Wen (2003) use one such Taiwanese peach
cultivar, ‘In-ge-taur’, as a comparator in a
release note for a new melting-flesh cultivar.
Several low-chill breeding programmes are
experimenting with the use of stony-hard
flesh but no cultivars have been released.
Disease resistance
Increased pressure from disease and insect
pests is encountered in subtropical compared
with temperate growing regions. There is
127
often more active development and multiple
generations of the pest due to higher spring
and summer temperatures, increased rainfall,
increased length of the growing season and
warmer winters which allow carryover of
pests from one season to the next. Breeding
for resistance to these pests is therefore impor-
tant in low-chill peach breeding (Topp and
Sherman, 2000). Scorza and Okie (1990) list 22
fungal and bacterial pathogens, 11 viruses,
four nematodes and 28 insects as problems in
peach culture. Of these, only resistance to
nematodes, bacterial spot, brown rot, pow-
dery mildew, rust and gummosis are actively
being selected for in low-chill breeding pro-
grammes (Table 5.8). Genetic resources avail-
able for Prunus resistance breeding are well
documented by Scorza and Okie (1990) and
Byrne et al. (2000).
Brown rot is a major disease of peach,
causing blossom blight and fruit infection
(Ogawa et al., 1995). Many low-chill peach-
growing regions aim to finish production
before the rainy season because of the diffi-
culty and expense of controlling this disease.
The Brazilian peach cultivar ‘Bolinha’ was
reported to be resistant to brown rot as indi-
cated by reduced rate of lesion development
and sporulation and low incidence of infected
fruit in the field (Feliciano et al., 1987). This
resistance occurs in the fruit epidermis and is
associated with high levels of phenolic com-
pounds in the fruit flesh and epidermis
(Gradziel et al., 1998). Unfortunately the phe-
nolic compounds, chlorogenic acid and caf-
feic acid, which are associated with resistance
are also associated with development of flesh
browning. ‘Bolinha’ has been used as a parent
in the EMBRAPA breeding programme and
brown rot resistance has been incorporated in
the recently released peach ‘Jubileu’ (Raseira
and Nakasu, 2000a). ‘Pepita’ also has low lev-
els of brown rot but this is considered a result
of early ripening and thus due to infection
avoidance (Raseira and Nakasu, 2000b).
Leaf rust (Tranzchelia discolor (Fuckel)
Tranzchel & Litvinov) is a major disease in
peaches grown in subtropical climates with
high summer rainfall. The extent and onset of
defoliation is related to the time of initial
infection (Bertrand, 1995). In temperate regions,
leaf rust seldom appears before late August
128 B.L. Topp et al.

Table 5.8. Pests and diseases being selected for in low-chill peach cultivar development. (Data from
Scorza and Okie, 1990; Okie and Pusey, 1996; Brooks and Olmo, 1997; Byrne et al., 2000; W. Sherman,
USA, 2004, personal communication.)

Breeding Genotypes with some level

Disease Pathogen programme of resistance

Nematodes Meloidogyne javanica UF ‘Okinawa’ and ‘Flordaguard’ rootstocks

Meloidogyne incognita ‘Flordahome’ ornamental
Bacterial spot Xanthomonas campestris UF ‘Flordastar’, ‘Flordacrest’, ‘Sunblaze’,
pv. pruni (Smith) Dye ‘Suncoast’, ‘Sunhome’, ‘Sunraycer’,
‘FlordaMex 1’
Brown rot Monilinia fructicola (Wint.) EMBRAPA ‘Bolinha’, ‘Pepita’, feral Mexican
Honey seedlings
Monilinia laxa (Aderh. UF ‘Sungold’
& Ruhl.) Honey
Powdery Sphaerotheca pannosa UF ‘Flordagold’, ‘Flordagrande’,
mildew (Wallr.:Fr.) Lev. ‘Flordahome’, ‘Flordaking’, ‘Okinawa’
Mexico ‘Aztecgold’, ‘Hermosillo’, ‘Oro A’
Brazil ‘Diamante’
Rust Tranzchelia discolor UF ‘UF2000’, ‘UFQueen’, ‘SunBest’
(Fuckel) Tranzchel &
Litvinov
Gummosis Botrysphaeria dothidea UF ‘Sundowner’
(Moug.:Fr.) Ces. & De Not. USDA (Byron, PI65821
GA)

UF, University of Florida; EMBRAPA, Brazilian Agricultural Research Enterprise; USDA, US Department
of Agriculture.

(northern hemisphere) and so does not cause
significant early defoliation. However, in the
subtropics, rust appears in June (northern
hemisphere) and the subsequent premature
defoliation results in early leafing and flower-
ing. Early defoliation was shown to affect
depth of dormancy, growth ability of buds
and bud development (Lloyd and Firth, 1990).
In locations where winter freezes occur, the
return bloom is killed and subsequent yield
potential is reduced.
Perez et al. (1993) reported that local
Mexican seedling selections were more resis-
tant to rust than introduced cultivars. Highest
levels of resistance were found in the evergreen
types, with resistance partially correlated with
late ripening (r = 0.62). For the peach cultivar
‘UF2000’, it was reported that ‘leaves did not
drop readily when infected with rust as on
most varieties’ (Sherman and Lyrene, 2000).
Following this observation it was found that
12 genotypes (including the cultivars ‘UFQueen’,
‘SunBest’ and ‘UF2000’) from the UF breeding
programme had resistance to rust-induced
defoliation (Rouse and Sherman, 2002b). The
12 resistant genotypes averaged 5% defolia-
tion compared with 90% defoliation for the
remaining 181 susceptible genotypes. Resis-
tant selections had less rust than susceptible
selections at Imokalee in 2001, averaging two
and 61 lesions per leaf, respectively. The
authors speculated that the mode of inheri-
tance involved recessive genes for resistance
because the resistant genotypes had suscep-
tible parents.
Emphasis on breeding for resistance to
disease and pests is likely to increase in the
future. Currently, orchardists will plant sus-
ceptible cultivars and bear the cost of
increased plant protection if the fruit quality
(size, colour, firmness, shape and flavour) is
high enough. In the future, deregistration of
fungicides and insecticides is likely to increase
due to public pressure. Under these condi-
tions, genetic resistance combined with alter-
native management techniques (bait spraying,
 Low-chill Cultivar Development 129

pheromone disruption, protective canopies, selection management this can be increased

improved sanitation, etc.) will become more
important in subtropical peach culture.
5.4 Management of Low-chill Breeding
Populations
Low-chill breeders use specific management
techniques to take advantage of, and over-
come problems associated with, the subtropi-
cal environment. Long growing seasons allow
trees to reach large sizes; pests and diseases
go through many breeding cycles; high and
fluctuating winter temperatures reduce effec-
tive chilling; genotypes flowering in late win-
ter and early spring are prone to frost damage;
and flowering and harvest periods are often
prolonged. The shortest possible time for one
cycle of a recurrent mass selection programme,
from pollination to cultivar release, is 8 years
(Table 5.9). With poor seedling and advanced
up to 15 years. Some of the problems in pro-
ducing and managing breeding populations
in the subtropics are discussed in the follow-
ing section.
Hybrid seed production
Hybrid seed in temperate fruit breeding pro-
grammes is harvested in early to late summer,
stratified for 2–3 months and germinated in
autumn. In the subtropics, the seed is har-
vested in late spring, stratified for 4–6 weeks
and germinated in summer when maximum
temperatures are often above 30°C. Rosetting
of seedlings can be a major problem under
these conditions (Fig. 5.5/Plate 52). At the UF
breeding programme, germinating seed is
placed close to evaporative coolers in the
greenhouse for the first 10–14 days after ger-
mination to reduce ambient temperatures and

Table 5.9. Calendar of events in one cycle of low-chill breeding.

Year Activity Numbers

0 Identification, collection and propagation of parental 20 to 50 parents

material
1 Hybridization of parents; harvest, stratify and germinate 20,000 to 40,000 pollinations to
hybrid seed; grow-on seedlings and field plants produce 4000 seeds
2 1st year of field growth for seedling population 2,000 to 3,000 seedlings
3 50% of seedlings produce fruit in 2nd leaf; evaluate fruit 1,000 to 1,500 fruiting seedlings
and tree characteristics; select outstanding genotypes
Propagate selected genotypes on to virus-tested Propagate 20 selections
rootstock and plant at test sites
4 Propagated selections undergo first season of growth Test all 20 selections at two
at test sites sites
Evaluate remainder of seedling population; select Propagate 20 selections
outstanding genotypes
Propagate selected genotypes on to virus-tested Test about five elite selections
rootstock and plant at test sites at six sites
5 Evaluate 1st crop of superior selections
6 Evaluate 2nd crop of superior selections
Screen promising selections for pollen-borne viruses
7 Evaluate 3rd crop of superior selections
Make decision to release
Increase budwood supply
Complete Plant Breeders Rights
8 Name and release cultivar Release cultivar for commercial
planting
130 B.L. Topp et al.
decrease rosetting. Rosetting is also cultivar-
dependent and was found to be high in seed-
lings from genotypes with FDP of <110 days
but dropped rapidly as FDP increased (Bacon
and Byrne, 1995). Experience at UF with root-
stock seed indicates that genotypes with FDP
of >120 days can be successfully germinated
with no rosetting by drying and then re-imbibing
Fig. 5.5. (a) Rosetting of peach
seedling terminal bud can be a
major source of seedling loss in
the glasshouse or during trans-
planting to the field. (b) Normal
shoots developing from below
the rosetted terminal.
the seed prior to stratification. Seeley et al. (1998)
used this method to germinate ‘Johnson Elberta’
seed and reported that rosetting decreased
with increasing duration of stratification over
a range of stratification temperatures. Byrne
et al. (2000) noted that stratifying endocarp-
removed seed at 3–5°C rather than at 7–10°C
decreased rosetting. Thus, both stratification
 Low-chill Cultivar Development
temperature and duration influence rosetting.
Pinching out the rosetted terminal bud of the
rosetted seedling can force lower buds to
sprout and so partly alleviates the problem
(Barbosa et al., 1989). This is not 100% effective
and is not possible in plants that have no basal
buds. Application of giberellic acid to rosetted
peach × almond rootstock plantlets derived
from tissue culture was effective in forcing
new shoots (Tsipouridis and Thomidis, 2003).
Many low-chill breeding programmes
have early ripening as an objective and so
short FDP seed must be germinated. Seed
from genotypes with FDP of <90 days is con-
sidered immature and generally requires
embryo culture for adequate seed germina-
tion, while those with FDP of <70 days require
in ovulo embryo culture to allow the embryo
to develop inside the seedcoat prior to germi-
nation. There is an intermediate stage of FDP
of 80–90 days when embryos can be germi-
nated with standard stratification, but with
varying results depending on the genotype
and environment during fruit development.
Generally, trees with heavy crop loads will
ripen fruit later and so give improved seed
germination (W.B. Sherman, USA, 2004, per-
sonal communication).
Low-chill peach breeders sometimes
wish to combine high-chill and low-chill par-
ents in order to introduce specific tree or fruit
characteristics from temperate cultivars into
their low-chill breeding populations. Usually,
low-chill pollen is collected from the subtrop-
ical location and used the same season for
pollinating the high-chill parent in a temper-
ate location. Alternatively, the high-chill pol-
len can be collected the previous year, stored
and then used in the subtropical location. It is
difficult to decide the best location for grow-
ing out these hybrid populations. Chilling
requirement of these populations is skewed
towards the low-chill parent, with the medi-
um-chill seedlings the largest class, but there
will be transgressive segregants (seedlings
with chilling outside the range of the two par-
ents). If the aim is solely to select low-chill
genotypes in this first generation then it is
possible to grow the population in the sub-
tropical environment.
Selection for flower bud chilling require-
ment on the basis of seed chilling requirement
131
would be useful in allowing the separation of
genotypes at the time of germination into dif-
ferent chill categories. Observations have fre-
quently been made that seed from low-chill
genotypes requires less time in stratification
to germinate than seed from high-chill geno-
types. Rodriguez-Alcazar and Sherman (1985)
studied peach germplasm with chilling
requirements of 200 to 450 CU and found a
significant but low correlation (r = 0.21) between
individual seed chilling requirement and the
flower bud chilling requirement of the result-
ing seedling tree. Perez-Gonzalez (1990) stud-
ied feral Mexican and introduced peach
genotypes that had a wide range of chilling
requirements, from ‘Okinawa’ (150 CU) to
‘Elegant Lady’ (750 CU). There was a strong
correlation between the time of bloom of indi-
vidual trees and time to germinate for the
resulting selfed seed, for both the Mexican
(r = 0.71) and the introduced (r = 0.87) germ-
plasm. In hybrid combinations between early-,
medium- and late-blooming genotypes, there
was also a correlation between the chilling
requirement of the parents and the stratifica-
tion period required to germinate the result-
ing seed. It seems that where the parents do
not differ greatly in chilling requirement it is
not practical to pre-select for chilling require-
ment on the basis of length of seed stratifica-
tion; however, it should be possible to use the
relative mean stratification time of seed lots
from divergent chilling germplasm as an
indicator of broad chilling requirement to
allow selection of appropriate sites for evalu-
ation of the resulting seedlings.
An interesting side note of this work
(Perez-Gonzalez, 1990) was the marked dif-
ference between the selfed populations of
local Mexican feral selections and introduced
selections in terms of flowering time and seed
germination. While there was a strong corre-
lation between parental flowering time and
resulting seed germination time for both sets
of material, it was noted that the Mexican
peaches flowered much later for the same
relative seed germination time. Perez con-
cluded that the feral Mexican germplasm,
with natural selection occurring for genera-
tions in a spring frost environment, had a
high heat requirement for flowering or seed
germination (Perez-Gonzalez, 1990).
132 B.L. Topp et al.
Seedling populations
An ideal seedling management system will
minimize the juvenile period to accelerate
genetic gain, provide a uniform environment
for all seedlings to minimize non-genetic
variation, allow accurate prediction of the
seedling’s worth as a potential cultivar and
do all this at the lowest possible cost (Rodri-
guez-Alcazar et al., 1986). Techniques adopted
in low-chill breeding programmes to facilitate
these objectives include the use of high-den-
sity plantings, protective canopies, growth
regulators and standard orchard practices to
encourage rapid tree growth (i.e. irrigation,
fertilization, pest control).
High-density planting of seedlings increa-
ses breeding efficiency by reducing costs of
land, soil fumigation, irrigation and weed
control (Sherman and Lyrene, 1983). Sherman
et al. (1973) described a high-density fruiting
nursery where the trees were planted at prop-
agation nursery densities (1.0 m × 0.2 m), but
left until fruiting rather than transplanted
after one season (Fig. 5.6/Plate 53). This sys-
tem promotes rapid growth in height and so
Fig. 5.6. High-density fruiting nursery at Gainesville, Florida, showing the size of second leaf peach
seedlings grown in the subtropics.
enhances precocious fruiting by increasing
the distance between roots and apical buds
(Zimmerman, 1972).
Rodriquez-Alcazar et al. (1986) studied
the performance of seedlings in a high-den-
sity peach nursery at Gainesville, Florida and
the subsequent performance of grafted selec-
tions from this nursery. They found a signifi-
cant correlation between the two sets of
material for ratings of FDP, chilling require-
ment, fruit weight and fruit colour, but not for
crop load, fruit shape or firmness. It was
noted that poor correlations may indicate the
limited range of variability in the selected
population rather being than an indication
that the selection procedure was not efficient
in discarding those with low expectations or
in identifying superior genotypes. In forest
tree breeding studies the heritability of vari-
ous traits such as diameter at breast height
has been estimated at differing tree ages to
allow determination of the most efficient
stage of measurement. Similar studies using a
full population of all stage 1 and stage 2 seed-
lings would be needed for an assessment in
peach.
 Low-chill Cultivar Development
Reducing generation length is a major
objective in seedling management because
the rate of response to selection is inversely
proportional to the length of the selection
cycle (Hansche, 1983). Selection for precocity
and implementation of high levels of man-
agement to encourage rapid seedling growth
in high-density nurseries have reduced peach
breeding cycles to 3 or 4 years (Scorza and
Sherman, 1996). In current commercial low-
chill breeding germplasm it is not uncommon
to have 20–75% of the seedlings fruit 2 years
from seed. Juvenility may be further reduced
by increasing the proportion of the popula-
tion that fruits in the second growing season
or by inducing a proportion of the seedlings
to fruit 1 year from seed. Repeated cycles of
the fruiting nursery system have appeared to
increase the precocity and the percentage of
seedlings fruiting in 2 years.
With the long growing seasons and warm
temperatures in the subtropics it is common
for seedlings to be 2–3 m tall in the first year.
In the second year almost all the seedlings
will be at a height when they have reached
the juvenile–adult phase change. Applica-
tions of growth regulators such as paclobutra-
zol have been reported to reduce the length of
the juvenile period in citrus seedlings (Snow-
ball et al., 1994) and similar methods should
be tested in peach.
Avoiding stratification of spring-har-
vested low-chill seed by direct germination
methods (Taylor, 1957) followed by early
summer planting of resulting seedlings would
allow production of trees that were 1–2 m in
height by the end of that subtropical growing
season. These seedlings may then be induced
to flower the following spring, thus reducing
the generation time to 1 year.
Testing advanced selections
Methods for testing advanced selections vary
across low-chill breeding programmes. They
include the common elements of rapid, impar-
tial testing over multiple sites and years, com-
parison with standard cultivars and use of
commercial agronomic systems.
At the UF breeding programme, the min-
imum amount of data required prior to release
133
is three commercial crops obtained from two
to five budded trees at one representative
location. In practice, there is usually more
data because advanced selections are concur-
rently tested at numerous sites. In other pro-
grammes, such as ARC Infruitec in South
Africa, there is a phase of semi-commercial
planting prior to release.
Although most of the low-chill breeding
programmes exist at public institutions, there
has been increasing pressure to obtain exter-
nal funding sources to assist in cultivar devel-
opment. Funding partners are usually involved
in the testing of advanced selections and pro-
vide valuable feedback to the breeder on the
worth of the new selections and the direction
of the breeding programme. These partners
may be nurseries, fruit-growing companies,
packinghouses, market agents, retailers, grower
cooperatives or individual orchardists.
5.5 Conclusions
The quality of low-chill peach cultivars has
increased significantly since the mid-1960s.
Many industries started with cultivars such
as ‘Flordasun’ and ‘Sunred’ that allowed pro-
duction of peach and nectarine in new loca-
tions, and at harvest times that had previously
not occurred. The long-running breeding pro-
gramme at UF in the USA has had a signifi-
cant impact on world peach production with
its cultivars planted extensively around the
world, while the programmes at EMBRAPA
and IAC in Brazil and ARC Infruitec in South
Africa have been successful in supplying
cultivars for their local industries. More
recently, low-chill breeding has started in
desert environments (California and south-
ern Texas), tropical highlands (Mexico and
Thailand) and in the humid subtropics (Aus-
tralia, Taiwan).
The driving force for developing low-
chill peach cultivars is the production of ear-
ly-ripening fruit to extend the peach season.
In some countries such as Brazil and Thailand,
where only low-chill peaches are produced, a
full range of low-chill ripening times is required;
but for most locations, low-chill peaches
provide an early supplement to the high-chill
134 B.L. Topp et al.

season. If forecasts of global warming are
correct, low-chill cultivars will be required in
more locations.
Low-chill peach breeding has received
far less resources than high-chill breeding, in
keeping with the smaller size of the low-chill
peach industry. Combined with the subopti-
mal environmental conditions that exist in
most low-chill production regions, this means
that the quality of low-chill cultivars has often
been secondary to high-chill cultivars. Most
of the ideal peach-growing locations (Mediter-
ranean environments) require medium- and
high-chill cultivars and in these environments
breeders have been able to select strongly for
fruit quality characteristics without diluting
selection intensity for environmental adap-
tation. All of the current low-chill peach
breeding programmes are incorporating the
significant high-chill peach breeding gains
into their programmes by the introgression of
high-chill germplasm.
With increased pressure on public breed-
ing programmes to obtain external funding,
there has been a trend of increasing protec-
tion of new cultivars with Plant Breeders
Rights and Plant Patents. This has resulted in
less sharing of germplasm. With increased
consumer demand for consistently high fruit
quality, breeders have shifted emphasis in
selection from productivity to eating quality.
This has only been possible because of the
earlier breeding work that provided germ-
plasm with good adaptation to many of the
problems encountered in low-chill peach-
growing locations. The emphasis in breeding
for quality will continue, and there will be
increased emphasis on novel fruit types for
consumers and increased disease resistance
and altered tree architecture for producers.

Contributors

Bacon, Terry, Sun World International, Inc., PO Box 80298, Bakersfield, CA 93380-0298, USA.
Barbosa, Wilson, Instituto Agronomico (IAC), Caixa Postal 28, 13001-970 Campinas, Sao Paulo, Brazil.
Beppu, Kenji, Faculty of Agriculture, Kagawa University, Miki, Kagawa 761-0795, Japan.
Boonprakob, Unaroj, Department of Horticulture, Kasetsart University, Mampangsaen, Nakhonpathorn
73140, Thailand.
Burger, Stella, Stargrow South Africa, PO Box 12536, Die Boord 7613, South Africa.
Byrne, David, Texas A&M University, Department Horticultural Science, College Station, TX 77843-2133,
USA.
Darmody, Liz, Zee Sweet Pty Ltd, PO Box 21 Monbulk, VIC 3793, Australia.
Kataoka, Ikuo, Faculty of Agriculture, Kagawa University, Miki, Kagawa 761-0795, Japan.
Perez, Salvador, Prol. Zaragoza 408, Jardines de la Hacienda, Queretaro, Qro 76180, Mexico.
Porter, Gavin, ANFIC, George Street, Bathurst, NSW 2795, Australia.
Richards, Graeme, University of Western Sydney, Richmond, NSW 2753, Australia.
Rodriguez-Alcazar, Jorge, Centro de Fruitcultura, Colegio de Postgraduados, Chapingo 56230, Mexico.
Russell, Dougal, Department of Primary Industries, PO Box 501, Stanthorpe, QLD 4380, Australia.
Smith, Chris, ARC Infruitec–Nietvoorbij, Private Bag X5013, Stellenbosch 7599, South Africa.
Wang, Lirong, Zhengzhou Fruit Research Institute, Chinese Academy of Science, Zhengzhou, Henan 450009,
People’s Republic of China.
Wen, Ien Chie, TARI, 189 Chungcheng Road, Wufeng, 413 Taichung, Taiwan, Republic of China.

References

Allan, P., Rufus, G., Linsley-Noakes, G.C. and Matthee, G.W. (1995) Winter chill models in a mild subtropical
area and effects of constant 6°C chilling on peach budbreak. Acta Horticulturae 409, 9–17.
Bacon, T.A. and Byrne, D.H. (1995) Relationships of fruit development period, seed germination, seedling
survival, and percent dry weight of ovule in peach. HortScience 30, 833.
Barbosa, W., Campo-Dall’Orto, F.A. and Ojima, M. (1989) Eliminacao de anomalias fisiologicas, in vitro, de
plantulas de pessegueiro. Bragantia 48, 13–19.
 Low-chill Cultivar Development 135

Barbosa, W., Ojima, M., Campo-Dall’Orto, F.A., Rigitano, O., Martins, F.P., Santos, R.R. and Castro, J.L.
(1997) Melhoramento do pessegueiro para regioes de clima subtropical-temperado: realizacoes do
Instituto Agronomico no periodo de 1950–1990. Documentos IAC 52. Instituto Agronomico, Campinas,
Brazil.
Bassols, M.C.B. (1973) Phenotypic segregation within an hybrid seedling population of peach, Prunus persica
(L.) Batsch. Masters thesis, University of Arkansas, Fayetteville, Arkansas.
Beckman, T.G., Krewer, G., Sherman, W.B. and Okie, W.R. (1995) Breeding moderate chill peaches for the
lower coastal plain. Proceedings of the Florida State Horticultural Society 108, 345–348.
Bertrand, P.F. (1995) Rust. In: Ogawa, J.M., Zehr, E.J., Bird, G.W., Ritchie, D.F., Uriu, K. and Uyemoto, J.K.
(eds) Compendium of Stone Fruit Diseases. APS Press, St. Paul, Minnesota, pp. 23–24.
Boonprakob, U. and Byrne, D.H. (2003) Temperature influences blind node development in peach. Acta
Horticulturae 618, 463–467.
Boonprakob, U., Byrne, D.H. and Rouse, R.E. (1994) A method for blind node evaluation. Fruit Varieties
Journal 48, 101–103.
Brooks, R.M. and Olmo, H.P. (1997) Register of Fruit & Nut Varieties, 3rd edn. ASHS Press, Alexandria, Virginia.
Brovelli, E.A., Brecht, J.K., Sherman, W.B., Sims, C.A. and Harrison, J.M. (1999) Sensory and compositional
attributes of melting- and non-melting-flesh peaches for the fresh market. Journal of the Science of Food
and Agriculture 79, 707–712.
Byrne, D.H. (1986) Mechanisms of spring freeze injury avoidance in peach. HortScience 21, 1235–1236.
Byrne, D.H. (2003) Founding clones of low chilling fresh market peach germplasm developed in the USA and
Brazil. Acta Horticulturae 606, 17–21.
Byrne, D.H. and Bacon, T.A. (1999) Founding clones of low-chill fresh market peach germplasm. Fruit Varieties
Journal 53, 162–171.
Byrne, D.H., Sherman, W.B. and Bacon, T.A. (2000) Stone fruit genetic pool and its exploitation for growing
under warm winter conditions. In: Erez, A. (ed.) Temperate Fruit Crops in Warm Climates. Kluwer
Academic Publishers, Dordrecht, The Netherlands, pp. 157–230.
Citadin, I., Raseira, M.C.B., Herter, F.G. and Baptista da Silva, J. (2001) Heat requirement for blooming and
leafing in peach. HortScience 36, 305–307.
Cullinan, F.P. (1937) Improvement of stone fruits. In: United States Department of Agriculture Yearbook of
Agriculture. United States Government Printing Office, Washington, DC, pp. 665–748.
Della Strada, G. and Fideghelli, C. (2003) Le cultivar di drupacee introdotte dal 1991 al 2001. L’Informatore
Agrario 41, 65–70.
Della Strada, G., Fideghelli, C. and Grassi, F. (1996) Peach and nectarine cultivars introduced in the world
from 1980 to 1992. Acta Horticulturae 374, 43–51.
Dennis, F.G. (2003) Problems in standardizing methods for evaluating the chilling requirements for the break-
ing of dormancy in buds of woody plants. HortScience 38, 347–350.
Edwards, G.R. (1987) Temperature in relation to peach culture in the tropics. Acta Horticulturae 199, 61–62.
Erez, A., Fishman, S., Linsley-Noakes, G.C. and Allan, P. (1990) The dynamic model for rest completion in
peach buds. Acta Horticulturae 276, 165–173.
Feliciano, A., Feliciano, A.J. and Ogawa, J.M. (1987) Monolinia fructicola resistance in the peach cultivar
Bolinha. Phytopathology 77, 776–780.
Felker, F.C. and Robitaille, H.A. (1985) Chilling accumulation and rest of sour cherry flower buds. Journal of
the American Society for Horticultural Science 110, 227–232.
George, A. and Nissen, B. (1998) Key issues: determining chilling units. In: Vock, N.T. (ed.) Low Chill Stone-
fruit Information Kit. Queensland Government, Brisbane, Australia, pp. 25–28.
Gradziel, T.M., Thorpe, M.A., Bostock, R.M. and Wilcox, S. (1998) Breeding for brown rot (Monilinia fructi-
cola) resistance in clingstone peach with emphasis on the role of fruit phenolics. Acta Horticulturae 465,
161–170.
Haji, T., Yaegaki, H. and Yamaguchi, M. (2003) Softening of stony hard peach by ethylene and the induction
of endogenous ethylene by 1-aminocyclopropane-1-carboxylic acid (ACC). Journal of the Japanese
Society for Horticultural Science 72, 212–217.
Hansche, P.E. (1983) Response to selection. In: Moore, J.N. and Janick, J. (eds) Methods in Fruit Breeding.
Purdue University, West Lafayette, Indiana, pp. 154–171.
Hennessy, K.J. and Clayton-Greene, K. (1995) Greenhouse warming and vernalisation of high-chill fruit in
southern Australia. Climatic Change 30, 327–428.
Horne, W.T., Weldon, G.P. and Babcock, E.B. (1926) Resistance of peach hybrids to an obscure disease in
Southern California. Journal of Heredity 17, 98–104.
136 B.L. Topp et al.

Hume, H.H. (1902) The Peen-to peach group. Bulletin Florida Agricultural Experiment Station 62, 505–519.
Cited in: Knight, R.L. (ed.) (1969) Abstract Bibliography of Fruit Breeding and Genetics to 1965. Prunus.
CAB, London.
Karakurt, Y., Huber, D.J. and Sherman, W.B. (2000) Development of off-flavours in non-melting flesh peach
genotypes. Journal of the Science of Food and Agriculture 80, 1841–1847.
Krewer, G., Beckman, T.G. and Sherman, W.B. (1998) Moderate chilling peach and nectarine breeding and
evaluation program at Attapulgus, Georgia. Acta Horticulturae 465, 155–160.
Lammerts, W.E. (1941) An evaluation of peach and nectarine varieties in terms of winter chilling requirements
and breeding possibilities. Proceedings of the American Society for Horticultural Science 39, 205–211.
Lammerts, W.E. (1945) The breeding of ornamental edible peaches for mild climates. I. Inheritance of tree and
flower characters. American Journal of Botany 32, 53–61.
Lesley, J.W. (1944) Peach breeding in relation to winter chilling requirements. Proceedings of the American
Society for Horticultural Science 45, 243–250.
Lester, D.R., Sherman, W.B. and Atwell, B.J. (1996) Endopolygalacturonase and the melting flesh (M) locus in
peach. Journal of the American Society for Horticultural Science 121, 231–235.
Li, Z.L. (1984) Peach germplasm and breeding in China. HortScience 19, 348–351.
Lloyd, J. and Firth, D. (1990) Effect of defoliation time on depth of dormancy and bloom time for low-chill
peaches. HortScience 25, 1575–1578.
McPhee, G. (2003) Phytonova launched. Summerfruit Australia Quarterly 5, 20–21.
Nissen, R.J., George, A.P. and Ward, G. (2000) Stonefruit export marketing opportunities. Australian Fresh
Stone Fruit Quarterly 2, 12–16.
Ogawa, J.M., Zehr, E.I. and Biggs, A.R. (1995) Brown rot. In: Ogawa, J.M., Zehr, E.I., Bird, G.W., Ritchie, D.F., Uriu,
K. and Uyemoto, J.K. (eds) Compendium of Stone Fruit Diseases. APS Press, St. Paul, Minnesota, pp. 7–10.
Okie, W.R. (1998) Handbook of Peach and Nectarine Varieties. USDA Agricultural Handbook No. 714. US
Government Printing Office, Washington, DC.
Okie, W.R. and Pusey, P.L. (1996) USDA peach breeding in Georgia: current status and breeding for resistance
to Botryosphaeria. Acta Horticulturae 374, 151–158.
Ou, S.K. and Wen, I.C. (2003) ‘SpringHoney’ peach. HortScience 38, 633–634.
Perez, S. (1989) Characterization of Mexican peach populations from tropical and subtropical regions. Acta
Horticulturae 254, 139–144.
Perez, S. (2003) Improved peach rootstocks and nursery management practices for subtropical climates.
Scientia Horticulturae 98, 149–156.
Perez, S. (2004) Yield stability of peach germplasm differing in dormancy and blooming season in the Mexican
subtropics. Scientia Horticulturae 100, 15–21.
Perez, S., Montes, S. and Mejia, C. (1993) Analysis of peach germplasm in Mexico. Journal of the American
Society for Horticultural Science 118, 519–524.
Perez-Gonzalez, S. (1990) Relationship between parental blossom season and speed of seed germination in
peach. HortScience 25, 958–960.
Perez-Gonzalez, S. (2001) Importance of Brazilian peach germplasm for the Mexican subtropics. Acta Horti-
culturae 565, 75–78.
Perez-Gonzalez, S. (2002) Variables associated with evolution and adaptation of peach seedlings to subtropical
environments. Acta Horticulturae 592, 143–148.
Price, R.H. (1905) Classification of the peach according to races. In: The Cherry, Peach, Pear, Plum, Small
Fruits, Special Report of the American Pomological Society, pp. 68–71. Cited in: Knight, R.L. (ed.) (1969)
Abstract Bibliography of Fruit Breeding and Genetics to 1965. Prunus. CAB, London.
Proebsting, E.L. and Mills, H.H. (1978) Low temperature resistance of developing flower buds of six deciduous
fruit species. Journal of the American Society for Horticultural Science 103, 192–198.
Quezada, A.C., Raseira, M.C.B., Baptista da Silva, J. and Citadin, I. (2000) Herdabilidade da epoca de floracao
no pessegueiro (Prunus persica L. Batsch). Agropecuaria Clima Temperado, Pelotas 3, 75–84.
Ramming, D.W. and Tanner, O. (1987) Goldcrest peach. Fruit Varieties Journal 41, 52–53.
Raseira, M.C.B. and Nakasu, B.H. (2000a) EMBRAPA-CPACT Prunus breeding program. In: Silveira, E. (ed.)
Prunus Breeders Meeting, 2000, Pelotas. Summaries. Embrapa Clima Temperado Documentos No. 75.
Pelotas, RS, Brazil, pp. 73–77.
Raseira, M.C.B. and Nakasu, B.H. (2000b) Pepita. A peach cultivar for canning. Agropecuária Clima
Temperado, Pelotas 3, 283–285.
Raseira, M.C.B., Nakasu, B.H., Santos, A.M., Fortes, J.F., Martins, O.M., Raseira, A. and Bernardi, J. (1992) The
CNPFT/EMBRAPA fruit breeding program in Brazil. HortScience 27, 1154–1157.
 Low-chill Cultivar Development 137

Raseira, M.C.B., Herter, F. and Posser, C.A.S. (2003) The EMBRAPA/Clima Temperado peach breeding program
and adaptation to subtropical regions. Acta Horticulturae 606, 45–50.
Richards, G.D., Porter, G.W., Rodriguez-Alcazar, J. and Sherman, W.B. (1994) Incidence of blind nodes in
low-chill peach and nectarine germplasm. Fruit Varieties Journal 48, 199–202.
Richardson, E.A., Seeley, S.D. and Walker, D.R. (1974) A model for estimating the completion of rest for
Redhaven and Elberta peach trees. HortScience 9, 331–332.
Rodriguez-Alcazar, J. and Sherman, W.B. (1985) Relationships between parental, seed, and seedling chilling
requirement in peach and nectarine. Journal of the American Society for Horticultural Science 110,
627–630.
Rodriguez-Alcazar, J., Sherman, W.B. and Lyrene, P.M. (1986) High density nursery system for breeding peach
and nectarine: a 10-year analysis. Journal of the American Society for Horticultural Science 111, 311–
315.
Rouse, R.E. and Sherman, W.B. (2002a) High night temperatures during bloom affect fruit set in peach. Pro-
ceedings of the Florida State Horticultural Society 115, 96–97.
Rouse, R.E. and Sherman, W.B. (2002b) Foliar rust resistance in low-chill peaches. Proceedings of the Florida
State Horticultural Society 115, 98–100.
Scorza, R. and Okie, W.R. (1990) Peaches (Prunus). In: Moore, J.N. and Ballington, J.R. Jr (eds) Genetic
resources of temperate fruit and nut crops. Acta Horticulturae 290, 177–231.
Scorza, R. and Sherman, W.B. (1996) Peaches. In: Janick, J. and Moore, J.N. (eds) Fruit Breeding. Vol. I. Tree
and Tropical Fruit. Wiley, New York, pp. 325–440.
Scorza, R., Mehlenbacher, S.A. and Lightner, G.W. (1985) Inbreeding and coancestry of freestone peach
cultivars of the eastern United States and implications for peach germplasm improvement. Journal of the
American Society for Horticultural Science 110, 547–552.
Scorza, R., Sherman, W.B. and Lightner, G.W. (1988) Inbreeding and co-ancestry of low chill short fruit devel-
opment period freestone peaches and nectarines produced by the University of Florida breeding pro-
gram. Fruit Varieties Journal 42, 79–85.
Seeley, S.D., Ayanoglu, H. and Frisby, J.W. (1998) Peach seedling emergence and growth in response to
isothermal and cycled stratification treatments reveal two dormancy components. Journal of the American
Society for Horticultural Science 123, 776–780.
Sharpe, R.H. (1961) Developing new peach varieties for Florida. Proceedings of the Florida State Horticul-
tural Society 74, 348–352.
Sharpe, R.H., Sherman, W.B. and Martsolf, J.D. (1990) Peach cultivars in Florida and their chilling require-
ments. Acta Horticulturae 279, 191–197.
Sherman, W.B. and Lyrene, P.M. (1983) Handling seedling populations. In: Moore, J.N. and Janick, J. (eds)
Methods In Fruit Breeding. Purdue University, West Lafayette, Indiana, pp. 66–73.
Sherman, W.B. and Lyrene, P. (1998) Bloom time in low-chill peaches. Fruit Varieties Journal 52, 226–228.
Sherman, W.B. and Lyrene, P.M. (2000) ‘UF2000’ peach. Journal of the American Pomological Society 54,
48.
Sherman, W.B. and Lyrene, P.M. (2003) Low chill breeding of deciduous fruit at the University of Florida. Acta
Horticulturae 622, 599–605.
Sherman, W.B. and Rodriguez-Alcazar, J. (1987) Breeding of low-chill peach and nectarine for mild winters.
HortScience 22, 1233–1236.
Sherman, W.B., Sharpe, R.H. and Prince, V.E. (1972) Two red leaf characters associated with early ripening in
peaches. HortScience 7, 502–503.
Sherman, W.B., Sharpe, R.H. and Janick, J. (1973) The fruiting nursery: ultrahigh density for evaluation of
blueberry and peach seedlings. HortScience 8, 170–172.
Sherman, W.B., Topp, B.L. and Lyrene, P.M. (1990) Non-melting flesh for fresh market peaches. Proceedings
of the Florida State Horticultural Society 103, 293–294.
Snowball, A.M., Warrington, I.J., Halligan, E.A. and Mullins, M.G. (1994) Phase change in citrus: the effects
of main stem node number, branch habit and paclobutrazol application on flowering in citrus seedlings.
Journal of Horticultural Science 69, 149–160.
Souza, V.A.B., Byrne, D.H. and Taylor, J.F. (1998a) Heritability, genetic and phenotypic correlations, and
predicted selection response of quantitative traits in peach. I. An analysis of several reproductive traits.
Journal of the American Society for Horticultural Science 123, 598–603.
Souza, V.A.B., Byrne, D.H. and Taylor, J.F. (1998b) Heritability, genetic and phenotypic correlations, and
predicted selection response of quantitative traits in peach. II. An analysis of several fruit traits. Journal of
the American Society for Horticultural Science 123, 604–611.
138 B.L. Topp et al.

Souza, V.A.B., Byrne, D.H. and Taylor, J.F. (2000) Predicted breeding values for nine plant and fruit character-
istics of 28 peach genotypes. Journal of the American Society for Horticultural Science 125, 460–465.
Taylor, J.W. (1957) Growth of non-stratified peach embryos. Proceedings of the American Society for Horti-
cultural Science 69, 148–151.
Topp, B.L. and Sherman, W.B. (1989) Location influences on fruit traits of low-chill peaches in Australia.
Proceedings of the Florida State Horticultural Society 102, 195–199.
Topp, B.L. and Sherman, W.B. (2000) Breeding strategies for developing temperate fruits for the subtropics,
with particular reference to Prunus. Acta Horticulturae 522, 235–240.
Tsipouridis, C. and Thomidis, T. (2003) Methods to improve the in vitro culture of GF677 (peach × almond)
peach rootstock. New Zealand Journal of Crop and Horticultural Science 31, 361–364.
Wang, L., Zhu, G. and Fang, W. (2002) Peach germplasm and breeding programs at Zhengzhou in China. Acta
Horticulturae 592, 177–182.
Ward, K.M. (1952) The peach. Queensland Agricultural Journal 74, 323–334.
Watson, T.D., Albritton, L., Barker, T. et al. (2001) Climate Change 2001: Synthesis Report. Summary for Poli-
cymakers. An Assessment of the Intergovernmental Panel on Climate Change. http://www.ipcc.ch/pdf/
climate-changes-2001/synthesis-spm/synthesis-spm-en.pdf (accessed January 2008).
Wei, S. (2001) Singapore & Hong Kong market research for early season stone fruit. Australian Fresh Stone
Fruit Quarterly 3, 8–12.
Weinberger, J.H. (1950) Chilling requirement of peach varieties. Proceedings of the American Society for
Horticultural Science 56, 122–128.
Weinberger, J.H. (1967) Studies on bud drop in peaches. Proceedings of the American Society for Horticul-
tural Science 91, 78–83.
Wen, I.C. and Sherman, W.B. (2002) Evaluation and breeding of peaches and nectarines for subtropical
Taiwan. Acta Horticulturae 592, 191–196.
Williamson, J.G. and Sargent, S.A. (1999) Postharvest characteristics and consumer acceptance of non-melting
peaches. Proceedings of the Florida State Horticultural Society 112, 241–242.
Zimmerman, R.H. (1972) Juvenility and flowering in woody plants: a review. HortScience 7, 447–455.
 6 Fresh Market Cultivar Development

W.R. Okie,1 T. Bacon2 and D. Bassi3

1USDA-ARS Southeastern Fruit and Tree Nut Research Laboratory, Byron, Georgia, USA
2Sun World International, Inc., Bakersfield, California, USA
3University of Milan, Milan, Italy

6.1 Introduction 140

6.2 Early Cultivation 140
6.3 Early US Breeding Programmes 141
North-eastern USA and Canada 142
South-eastern USA 144
California 146
6.4 Modern US Breeding Programmes 150
North-eastern USA and Canada 151
South-eastern USA 153
California 154
6.5 European Programmes 161
Bulgaria 161
France 161
Greece 163
Hungary 163
Italy 163
Poland 165
Romania 166
Serbia 167
Spain 167
Ukraine and former Soviet states 167
6.6 Pacific and African Programmes 168
China 168
Japan 168
Korea 169
New Zealand 169
South Africa 169
6.7 Conclusions 169

© CAB International 2008. The Peach: Botany, Production and Uses

(eds D.R. Layne and D. Bassi) 139
140 W.R. Okie et al.
6.1 Introduction
Successful peach production depends on suit-
able cultivars, regardless of where the indus-
try is located. Peaches have a short shelf-life
compared with many crops and each cultivar
has a short production season. As a result,
multiple cultivars are needed to provide fresh
fruit from April to September (October to
March in the southern hemisphere) across
many regions and climates. Few consumers
recognize specific peach cultivars in the mar-
ket and few cultivars are marketed by name.
Such anonymity allows the rapid acceptance
of new cultivars, thus encouraging additional
cultivars to be developed.
In general, peaches for fresh market
consumption must suit the intended market.
In most cases preferred fruit are large (6–8 cm
diameter), predominantly red with yellow
ground colour and with short pubescence or
glabrous (nectarine), firm enough to be
transported (perhaps for several weeks for
exporting industries), of regular round shape
and of good eating quality (taste and tex-
ture). Flesh colour in the USA has tradition-
ally been yellow, but white is acceptable or
preferred in some markets. Most fresh mar-
ket peaches are freestone, except for the
early-season cultivars. Important excep-
tions are those sold in countries preferring
non-melting or canning clingstones (often
with minimal external red blush), such as
southern Italy, Spain, Mexico and other
countries with Spanish influence. Many of
the later-season nectarines are also cling-
stone, apparently because of higher eating
quality compared with freestone nectarines
in that season. In recent years the market has
diversified in terms of flesh colour, acidity
(normal or low), texture (melting, non-melting
and stony-hard) and shape (round or flat).
Changes are also coming in tree architecture
to provide cultivars ranging from dwarf
through standard to pillar and weeping,
although commercial adoption of non-stan-
dard trees remains to be seen. Most of the
breeding for low-chill regions, as well as that
for canning peaches, is discussed in other
chapters in this book (see Chapters 5 and 7,
respectively).
6.2 Early Cultivation
As peaches spread around the world, most
orchards consisted of seedling trees. For
centuries, farmers selected superior trees
from those grown from seed. Over time dis-
tinct strains arose in the various regions
where peaches were well adapted. In many
areas these local strains are still grown, at
least as a source for rootstock seed. Exam-
ples are the vineyard peaches of the former
Yugoslavia, ‘Criollo’ in Argentina, ‘San
Miguel’ in Spain and locally grown cling
peaches in central America. Fruit growers
later wished to preserve the trees that pos-
sessed useful or superior characteristics
and began to vegetatively propagate these
individuals.
Early Spanish explorers and colonists
brought the first peaches to North America as
seed. This new fruit was also spread across
the south-eastern and south-western states by
Native Americans to the extent that some
later European explorers thought peaches
were native to the USA. Descendants of these
seedlings became known as ‘Indian’ peaches
and live on in types such as ‘Tennessee Natu-
ral’ and ‘Indian Blood’. Later, mostly white-
fleshed cultivars from England and France
were brought into the eastern USA. Begin-
ning in the early 1800s, these European culti-
vars and the best local seedlings began to be
propagated by budding. Hedrick’s (1917)
classic work, The Peaches of New York, listed
2181 cultivars known at that time. A similar,
briefer work from Europe which included
nectarines included many more European
cultivars (Jouin, 1913). Many of the cultivars
in these books were of unknown seedling ori-
gin. For nearly all the rest only the seed par-
ent was known. In most areas where peaches
were adapted, the industries were based on
local cultivars well into the 20th century. As
late as 1937, a listing of important US culti-
vars was all seedling selections (Cullinan,
1937). Up until this time the nectarine was
primarily a novelty crop grown in drier cli-
mates for the purpose of drying. The small
size and a marked tendency for the skin to
crack in rainy weather limited its commercial
production elsewhere.
 Fresh Market Cultivar Development
T. Rivers of Hertfordshire, England attem-
pted to improve peaches and nectarines by
systematically growing seedlings of better-
adapted cultivars and selections beginning in
the 1820s, making him perhaps the first peach
breeder. Without knowledge of modern
genetics and control of the pollen parent, he
had only limited success despite releasing
over 30 peach cultivars (Rivers, 1866; Hoare,
1950). The rediscovery of Mendel’s work and
the development of modern genetics impacted
peach breeding as it was initiated by public
institutions. Rivers’ grandson published one
of the first reports of intentional hybridiza-
tion in 1906 (Rivers, 1906), in which he dis-
cussed inheritance of glabrous skin (nectarine),
flower type and gland type. His assessment
of the task of peach breeding was prophetic
(and a good pun): ‘It is a labor of Sisyphus,
but the stone occasionally lodges on the top
of the hill’.
Although peaches have been improved
in most areas in which they were grown by
selection of adapted types, the dramatic
improvement of the last century has taken
place mostly in North America.
6.3. Early US Breeding Programmes
In 1850 the ‘Chinese Cling’ (‘Shanghai Shui
Mi’) was introduced into the USA as a potted
tree. Seeds of this peach planted in Marshall-
ville, Georgia by S.H. Rumph produced the
cultivars ‘Georgia Belle’ and ‘Elberta’, which
formed the basis of the early industries across
the USA (Cullinan, 1937; Myers et al., 1989).
At that time most peaches were white-fleshed
English and French cultivars or their descen-
dants. ‘Elberta’ was found to be much firmer
and widely adapted, such that it was grown
from the south-east to the north-east, provid-
ing a continuous supply of fruit of this cultivar
all season long. Yellow-fleshed peaches showed
bruising less than white-fleshed peaches and
gradually yellow came to predominate, as
breeders released mostly yellow-fleshed cul-
tivars. ‘Elberta’ is now grown in only a few
places, but the name remains one of the few
that the public recognizes. ‘Elberta’ is found
in the parentage of most commercial peaches
141
developed in the USA. Today, peach cultivars
are less celebrated and are viewed as one of
many aspects of production. It is unlikely that
any new cultivar will become as widely known
among the general public as did ‘Elberta’.
Several amateur breeders produced early
selections in the USA (Cullinan, 1937; Okie,
1998). J.W. Kerr of Denton, Maryland crossed
‘Elberta’ with ‘Early Beauty’, ‘Rivers’ and
‘Mountain Rose’ in 1888, resulting in ‘Den-
ton’, ‘Elriv’ and ‘Elrose’. It is not clear if these
were hand pollinations, but they appear to be
the first recorded hybrids of peach. J.W. Steu-
benrauch of Mexia, Texas selected a number
of popular cultivars from seedlings. In about
1900, he budded a tree with both ‘Elberta’ and
‘Belle October’ to obtain seedlings combining
characters of both parents. ‘Frank’ was the
most successful result of this crossing.
Around 1900, the peach grower J.H. Hale
found an off-type tree on his farm in Connecti-
cut, which also performed well on his Georgia
farm. Introduced in 1912 as ‘J.H. Hale’, this
peach was widely planted, in part due to its
superior firmness, and often used in breeding
in later years. It was thought to have been a
seedling of ‘Elberta’ as well.
Over the next century, the cultivar pic-
ture changed from a time when most new cul-
tivars were once chance seedlings found by
peach growers (Table 6.1). During the 20th
century, numerous state and federal breeding
programmes were undertaken in the USA.
Although peaches and nectarines were grown
in many countries at that time, little breeding
was done outside the USA, a result perhaps
of the maturity of the industries elsewhere as
well as the lack of structured research capa-
bilities reflected in the land grant universities
of the USA. In any case, peach breeding flour-
ished in the USA in the 20th century; progress
was so dramatic that new, improved US culti-
vars spread around the world, often replacing
existing local cultivars and selections.
Breeding in the USA can be divided into
four segments – north, south, deep south and
west. In the northern tier of states, as one
might expect, bud and tree hardiness are crit-
ical issues in breeding. Tolerance to rain and
humid conditions and the resulting diseases
is important, which is true across the entire
eastern USA. Similar cultivars are used from
142 W.R. Okie et al.

Table 6.1. Historically important peach and nectarine (*) cultivars and their date and locale of origin.
(From Cullinan, 1937.)

Cultivar Year Origin

‘Late Crawford’ 1815 New Jersey, USA

‘Early Crawford’ 1820 New Jersey, USA
‘Chinese Cling’ 1850 Shanghai, China
‘St John’ 1860 ?, USA
‘Georgia Belle’ 1870 Georgia, USA
‘Elberta’ 1870 Georgia, USA
‘Phillips’ 1880 California, USA
‘Champion’ 1880 Illinois, USA
‘Lovell’ 1882 California, USA
‘Hiley’ 1886 Georgia, USA
‘Admiral Dewey’ 1899 Georgia, USA
‘Mayflower’ 1909? North Carolina, USA
‘J.H. Hale’ 1912 Connecticut, USA
‘Paloro’ 1912 California, USA
‘Fay Elberta’ 1915 California, USA
‘Golden Jubilee’ 1926 New Jersey, USA
‘Halehaven’ 1932 Michigan, USA
‘Sunhigh’ 1938 New Jersey, USA
‘Redhaven’ 1940 Michigan, USA
‘Quetta’* Pre-1906 Pakistan
‘Lippiatt’s’* Pre-1916 New Zealand
‘Le Grand’* 1942 California, USA

the north down to the uplands of the south-
ern states. The main peach industry in the
southern states runs across the region between
the uplands and the coastal plain, and is sub-
ject to years when chilling is inadequate for
some peaches. In this zone mid-winter tem-
peratures are rarely an issue, but it is critical
to have cultivars requiring 650–850 h of chill-
ing below 7°C. As one moves towards the
coast where the annual chilling is even less, a
different set of cultivars is required (see Chap-
ter 5). Generally, as a given peach cultivar is
cultivated further south, its fruit tends to have
more prominent tips and sutures, and to
retain a green ground colour closer to soften-
ing time. Finally, in California, chilling and
hardiness are less important due to the more
moderate climate. However, high summer
temperatures tend to reduce red skin colour
in comparison to the same cultivar grown in
the eastern states. California peaches often
have fewer flower buds, which reduces thin-
ning costs and enhances fruit size, but which
results in light crops when grown in colder
climates.
North-eastern USA and Canada
New York
The first formal institutional breeding pro-
gramme was established in 1895 in Geneva,
New York. S.A. Beach planted open-pollinated
seeds of ‘Elberta’. In 1910 U.P. Hedrick made
the first crosses there. Hedrick’s major contri-
bution was to assemble The Peaches of New
York, an excellent reference on peach cultivars
and culture (Hedrick, 1917). As with most of
these early programmes, populations were
modest and progress was slow. Only one cul-
tivar was released in the first 50 years of the
programme. In recent years breeding at
Geneva has stopped and only testing of new
cultivars and selections continues.
Iowa and Illinois
Programmes in Iowa and Illinois began in 1905
and 1907, respectively, to develop more cold-
hardy peaches for the mid-west USA. Iowa
breeders S.A. Beach and T.J. Maney were
 Fresh Market Cultivar Development
innovative in their use of cold-hardy germ-
plasm such as ‘Bailey’ and Prunus davidiana,
but the result was a step back in terms of fruit
size and quality. The only release from this
work in Iowa was ‘Polly’ in 1932. Breeding in
Illinois produced no cultivars until the ‘Prairie’
series in 1946 (Mowry, 1960). J.E. Markham, a
private breeder, in 1932 released ‘Hal-Berta
Giant’, which was the first patented peach
(US Plant Patent 7). Breeding is hampered in
climates where cropping is unreliable. As
with most of the northern programmes, these
are now closed.
Ontario
Canada represents the northern limits of com-
mercial peach production (Layne, 1997). Pro-
tected sites along the Great Lakes in Ontario
are suitable for cold-hardy cultivars. In 1911
peach breeding was begun by the Ontario
Department of Agriculture at Vineland, with
the goals of producing cultivars to extend the
‘Elberta’ season as well as to develop culti-
vars for canning. Fourteen cultivars had been
released by 1964, the most important of which
were ‘Veteran’, ‘Valiant’ and ‘Vedette’. Since
then the emphasis has shifted to development
of canning clings (see Chapter 7). Meanwhile
a second programme was started at Harrow
by Agriculture Canada. G.M. Weaver and later
R.E.C. Layne developed cold-hardy peaches
and nectarines that thrived in their climate
and had suitable fruit characteristics. This pro-
gramme pioneered testing selections methodi-
cally by use of controlled freezing chambers
to gauge cold hardiness of wood and buds.
Many of their cultivars were and still are widely
planted in Ontario and the northern USA,
especially ‘Canadian Harmony’, ‘Harbrite’,
‘Harson’, ‘Harrow Diamond’ and ‘Harrow
Beauty’. Nectarines ‘Hardired’ and ‘Harblaze’
have also been important.
New Jersey
Breeding in New Jersey began at Rutgers Uni-
versity in 1914 with C.H. Connors and later
M.A. Blake. At that time the main early culti-
vars were ‘Greensboro’, ‘Waddell’, ‘Connetts’,
‘Lola’ and ‘Carman’, followed in ripening by
‘Hiley’, ‘Belle’ and ‘Elberta’. The objectives of
143
the programme were to study peach genetics
and develop a replacement for ‘Carman’. The
first releases from this early breeding came in
1925 with the release of 15 new cultivars,
which represent the first cultivars released by
an institutional breeding programme in the
USA (Blake and Connors, 1936). By 1937 three
of these ranked in the top ten cultivars grown
in the state, with ‘Golden Jubilee’ the most
successful of the group. This programme was
perhaps the most diverse in the country, devel-
oping not only fresh market peaches, but also
nectarines and canning clings. In later years
flat and low-acid types were also included.
M.A. Blake, and then later L.F. Hough and
C.H. Bailey, were the first to extensively col-
lect and utilize germplasm from around the
world, in order to provide new characters
useful to the industry (Blake and Edgerton,
1946). Over the 20th century New Jersey
released more new cultivars than any other
peach breeding programme.
Virginia
Despite an early start to breeding in Virginia
(1914), little progress was made until mid-
century, when G.D. Oberle released a series of
peaches and nectarines. He selected strongly
for climatic adaptation and his releases such
as ‘Jefferson’, ‘Monroe’ and ‘Washington’ were
quite cold-hardy and resistant or tolerant to
bacterial spot and brown rot. Oberle also
released nectarines, ‘Cavalier’, ‘Redbud’, ‘Cher-
okee’ and others, that were better able than
most to tolerate the rainy climate of the eastern
USA (Oberle, 1971). Oberle was not replaced
and the programme ended in the 1970s.
Massachusetts and New Hampshire
Peach breeding in New England began in
Massachusetts in 1918. Breeders J.K. Shaw,
J.S. Bailey and A.P. French grew only a few
thousand seedlings, none of which were
deemed worthy of release. However, they
made very close observations of their popula-
tions and were able to determine the inheri-
tance of many characters. Their 1949 bulletin
summarized the then current state of peach
genetics and clarified the inheritance of most
important characters (Bailey and French, 1949).
144 W.R. Okie et al.
Breeding work there terminated shortly there-
after, but continued for 15 more years at New
Hampshire under A.F. Yeager and E.M.
Meader. In 1964 Meader released ‘Reliance’,
which is one of the most cold-hardy peaches
known, although the fruit quality is not up to
commercial standards.
Michigan
Although a latecomer to the breeding busi-
ness, Michigan stands out in impact on the
industry. Stanley Johnston was the son of an
agricultural extension agent in Michigan. He
was hired as Superintendent of the South
Haven Experiment Station in 1920, a position
he held until his death in 1969. He recognized
that the local peach industry grew too much
‘Elberta’, so there was potential for earlier,
firm, red peaches. He saved the seeds from
pollination studies done to prove ‘J.H. Hale’
needed a pollinator and from these made his
first selections, resulting in the release of
‘Halehaven’ (1932) and ‘Kalhaven’ (1938).
Although he was not trained as a geneticist,
he had a keen eye for what the farmer needed.
A cross of ‘Halehaven’ by ‘Kalhaven’ resulted
in ‘Redhaven’, released in 1940 (Iezzoni,
1987). When initially released it was the first
good freestone on the market. It quickly came
to prominence for higher-chill regions because
of its productivity, firmness and appearance,
and at one time was the most widely grown
peach in the world. Even now it is still widely
grown, although it is being replaced in some
areas. Later Michigan releases from a lifetime
total of 21,000 seedlings included ‘Fairhaven’,
‘Sunhaven’, ‘Richhaven’, ‘Glohaven’ and
‘Cresthaven’ (Kessler, 1969).
Maryland
Breeders at University of Maryland began work
in 1929, but the first release was ‘Redskin’ in
1944. It is still grown in some areas but is
declining in popularity. Breeding efforts soon
ended, but advanced selections were carried
on with the intent of developing late-ripening
cultivars with better red colour and fruit
quality. Later releases included ‘Marhigh’,
‘Marsun’ and ‘Marqueen’, which are little-
grown now. The US Department of Agriculture
(USDA) initiated a breeding programme in 1936
at its research facility in Beltsville, Maryland to
develop high-quality, cold-resistant peaches
(Havis et al., 1947; Okie et al., 1985). Breeder F.P.
Cullinan (1937) wrote a classic chapter on stone
fruit breeding up to that time for the 1937 USDA
Yearbook of Agriculture. He was followed by
breeders A.L. Havis and H.W. Fogle. Although
the programme had only a limited number of
releases, including ‘Ranger’ (1952) and ‘Redg-
lobe’ (1954), important contributions were
made to peach genetics. ‘Redglobe’ is the still
widely grown in the south-eastern USA, mak-
ing it one of the oldest cultivars still important
there. In 1980 the programme was moved to a
new facility in Kearneysville, West Virginia.
South-eastern USA
Texas
Texas has a wider range of chilling than most
states, so it is difficult for a single programme
to cover the entire gamut. Although Texas
had a long history of private breeding, public
breeding at Texas A&M University in College
Station only started in 1935. Chilling accumu-
lation at College Station is moderate most years,
so more effort went into developing low- to
moderate-chill cultivars, a chill range which
also had fewer breeders working to improve
it. No cultivars were named until 30 years later,
with the release of ‘Sam Houston’, followed
by ‘TAMU Denman’ and ‘Milam’. As the pro-
gramme under D.H. Byrne has shifted more
towards low-chill development in recent
years, it is discussed further in Chapter 5.
Georgia
USDA stone fruit breeding in Georgia began
in 1937 at the Horticultural Fruit Laboratory
in Fort Valley, in the centre of the main peach
production area (Okie et al., 1985). J.H. Wein-
berger was the peach breeder until 1954,
when he transferred to Fresno, California to
begin the peach breeding there. Weinberger
published seminal papers about the chilling
requirements of peach cultivars in addition
to naming several important peach cultivars.
He developed cultivars such as ‘Cardinal’
1

Plate 1. One-year-old fruiting shoots at bloom.

Plate 2. Dwarf peaches in a commercial orchard.
Plate 3. Relationship between crotch (α) and extension (β) angles.
Plate 4. Leaf stipules at petiole base.
Plate 5. Narrow leaves (left) compared to normal sized (right). Scale in centimetres.
6A B C

7 8

9 10

Plate 6. Leaf glands: (A) reniform; (B) globose; (C) eglandular (note close-up, below).
Plate 7. Red leaf (left) and green leaf (right) peach trees.
Plate 8. Fruit skin colour variability. From top centre going clockwise: a yellow nectarine (anthocyaninless), a
white peach (anthocyaninless), a yellow peach (100% blush), a white peach, a non-melting peach, a yellow flat
nectarine, a white flat peach, a white nectarine and a yellow nectarine.
Plate 9. Hyper-sensitivity reaction on a resistant peach after a proof bite by a green aphid.
Plate 10. Flower type: showy.
11 13

12

White, simple White, semi-double Variegated, white and red

Red, simple Red, semi-double

Chrysanthemum-like, red,
semi-double Pink, semi-double

Plate 11. Flower type: non-showy.

Plate 12. Flower diversity in ornamental peach cultivars (courtesy of M. Yoshida, Japan).
Plate 13. Flat fruit, from a double ovary.
14 15

16 17

18 19

Plate 14. Main fruit shapes in commercial cultivars: globose (left), flat (centre), oblong (right).
Plate 15. Fruit size gain in F1 progeny from distant-size parents.
Plate 16. Flesh colour variability. From left, top row: non-melting flesh, two yellows (greenish and bright yellow), a
white; bottom row: red flesh (‘blood’), a yellow melting (anthocyaninless), a yellow and a white melting (flat
shape), a white stony-hard (anthocyaninless).
Plate 17. Red 'blood' flesh in peach (courtesy of A. Liverani, Forli, Italy).
Plate 18. Red 'blood' flesh in nectarine (courtesy of A. Liverani, Forli, Italy).
Plate 19. Genetic variation for red 'blood' flesh trait in peach (courtesy of W.R. Okie, Byron, Georgia, USA).
20

21

22

24

Plate 20. Stony-hard fruit: white (anthocyaninless) flesh.

Plate 21. Stony-hard fruit: yellow flesh.
Plate 22. Comparison between mature (left) and immature (right) embryos taken from ‘Spring Crest’ peach at two
different stages: the immature embryos fail to germinate under standard stratification procedures and need to be
rescued in vitro as the very early ripening genotypes.
Plate 24. Slow ripening nectarine trees after leaf fall (dormant season).
23

Plate 23. Main phenological stages in peach (courtesy of E. Bellini, Florence University, Italy).
25
26

27

28

Plate 25. A Chinese painting by Ma Tai (1885–1935), telling a story that an old man with white hair but a young,
boyish face always steals peaches. People make fun of this long-lived man, but he explains ‘Peach is good for my
health’.
Plate 26. Chinese peach production regions: (I) north-west drought region; (II) northern China plain region; (III)
Changjiang River humid region; (IV) Yunnan–Guizhou high plateau cold region; (V) Qinghai–Tibet plateau cold
peach region; (VI) north-eastern China cold region; (VII) southern China subtropical region.
Plate 27. Modern greenhouse production.
Plate 28. Modern greenhouse production before postharvest canopy removal.
29 30

31 32

33 34

Plate 29. Modern greenhouse production after postharvest canopy removal.

Plate 30. Conventional orchard.
Plate 31. Modern high density orchard.
Plate 32. Modern high density orchard (dormant).
Plate 33. Ornamental 'chrysanthemum' peach.
Plate 34. Ornamental 'longevity' peach (from Wang and Zhuang, 2001).
35 36

37 38

39 40

Plate 35. A monoploid peach with some fruits.

Plate 36. A weeping tree trained in central axis.
Plate 37. Flat nectarines (‘platerines’ in French).
Plate 38. Scheme based on genotypic choice of parents to obtain pure lines. In each generation, parents giving
progenies with low horticultural value are eliminated.
Plate 39. Emasculated flower (on left) (courtesy of W.R. Okie, Byron, Georgia, USA).
Plate 40. Pollination by honeybee (courtesy of D.R. Layne, Clemson, South Carolina, USA).
41 42

44

43

46
45

Plate 41. Dehydrated stamens ready for artificial pollination: anthers and filaments render the pollen less sticky.
Plate 42. Emasculated flower pollinated by finger, a fast way for cross-breeding.
Plate 43. Self-pollination with waterproof paper bags.
Plate 44. Young seedling at the end of embryo culture.
Plate 45. Seedling orchard at first flowering.
Plate 46. Initial physical map of linkage group 1 of peach depicted in the Prunus genome website
(http://www.bioinfo.wsu.edu/gdr/).
47

48
Plate 47. General Prunus genetic map (Joobeur et al., 1998) and developing peach physical/EST map. Markers depicted in bold have been used to develop contigs of
peach BACs, which are depicted as the light-coloured circles with the number of BACs detected by each marker inscribed in the circle. Green circles denote two adjacent
markers detecting the same BACs. Black flags denote peach and tomato EST positions with the number of ESTs at this location posted above the flags.
Plate 48. Numbers of new low-chill peach and nectarine cultivars released from four countries, 1980–1992. These countries accounted for 94% of low-chill cultivars
released in this period (adapted from Della Strada et al., 1996).
49 50

52A 51A

B
B

53

Plate 49. Regular green leaf peach seedlings in foreground and peach seedling displaying red leaf character as-
sociated with short fruit development period (FDP) in centre at Nambour, Queensland. This trait can be used for
early identification of short FDP seedlings in late summer (Sherman et al., 1972).
Plate 50. Peach fruit with prominent stylar tips that are commonly observed in cultivars with poor adaptation to
mild winter locations and are selected against in low-chill breeding.
Plate 51. Influence of location on fruit shape. This medium-chill (450 CU) peach selection produced: (A) round
fruit at the high-chill (900 CU) location of Stanthorpe Queensland and (B) pointed fruit at the low-chill (200 CU)
location of Nambour Queensland.
Plate 52. (A) Rosetting of peach seedling terminal bud can be a major source of seedling loss in the glasshouse
or during transplanting to the field. (B) Normal shoots developing from below the rosetted terminal.
Plate 53. High density fruiting nursery at Gainesville, Florida showing size of second leaf peach seedlings grown
in subtropics.
54 55

58
56 Loadel(1) 0
Stanislaus(2) 3
Crosses for
Carson(3) 5 genetic
Dee-Six (3) 5 Cultivar Select parents recombination
release for desired
Goodwin(3) 14 traits (1 year)
Bowen(1) 16 (1–5 years)
Fay Elberta(1) 17
Andross(3) 18
Arakelian(1) 19
Peak(1) 21
Klamp(3) 21
Tuolume(2) 22
Andora(1) 23
Select among
Ross(3) Evaluate
23 progeny
performance at
Rizzi(3) 28 multiple sites
(2–5 years)
and years
Dr. Davis(3) 30
Carolyn(3) 31 (8–16 years)
Monaco(1) 33
Lilleland(3) 35
Halford(1) 36
Everts(1) 37
Wiser(1) 38
Riegels(3) 38
Stam(1) 39
Hesse(3) 40
Sullivan #4(1) 41
Corona(3) 44
0 5 10 15 20 25 30 35 40 45 50

57 Corona Hesse

Late
4 Ross
Carolyn Dr.Davis
Everts Rizzi
Ripe period

Andora Lilleland
3 Ross Riegels
Klamt

Bowen
Fortuna Andross
2
Goodwin
Jungerman Tufts
Carson Dee-Six
1
1940 1946 1952 1958 1964 1970 1976 1982 1988 1994 2000
1943 1949 1955 1961 1967 1973 1979 1985 1991 1997
Year

Plate 54. Ripe fruit of ‘Rubyprince’ peach, a typical modern peach with extensive red blush and short
pubescence.
Plate 55. Processing clingstone peach showing the uniformly yellow, firm, non-melting flesh and associated
clingstone type stone-to-flesh adhesion.
Plate 56. Harvest sequence of California processing peach cultivars using the fresh market freestone ‘Fay
Elberta’ as a reference cultivar (1 – grower selection; 2 – private breeder release; none – released by public
breeding programme).
Plate 57. History of processing clingstone cultivar release by public breeding programmes in California showing
punctuated release at approximately 20-year intervals.
Plate 58. Basic components of a processing peach breeding programme with estimated duration in years.
59 20

62
15
Number of seedings

10

0
5 15 25 35 45 55 65 75 85
Days after 1 June

60 61

64
63

Plate 59. Fruit ripe date for progeny from self-pollination of the cultivar ‘Carson’ showing unusual bi-modal
distribution. (‘Carson’ normally ripens approximately 40 days after 1 June).
Plate 60. Poor lye peeling of a clingstone genotype with unacceptably thick epidermis.
Plate 61. Halved section of overripe clingstone peach demonstrating the common pattern of vascular strands
radiating from endocarp to outer mesocarp.
Plate 62. Rooted hardwood cuttings of peach.
Plate 63. Mist propagation facility for rooting of semi-hardwood cuttings.
Plate 64. Rooted semi-hardwood cuttings that were treated with auxins.
66

65 67

68

Plate 65. Hundreds of micropropagated plantlets in commercial facility.

Plate 66. The main steps of the micropropagation cycle.
Plate 67. Acclimatization of young micropropagated plantlets.
Plate 68. Excised embyros in aseptic culture.
69 70

71 72

73

Plate 69. Plantlets generated from aseptic embryo culture.

Plate 70. Traditional open vase tree form.
Plate 71. Palmette tree form.
Plate 72. Central leader tree form.
Plate 73. Fusetto orchard.
 Fresh Market Cultivar Development 145

and ‘Dixired’ that shifted the earliest ripening
date 2 weeks earlier than before. V.E. Prince
continued the breeding in 1954. In 1964, the
programme moved 20 miles east to the newly
opened South-eastern Fruit and Tree Nut
Research Laboratory in Byron. Prince also
developed many important peaches, in par-
ticular ‘Springcrest’, which became the most
widely grown peach since ‘Redhaven’. ‘Spring-
crest’, along with ‘Springbrite’ and ‘Spring-
old’, moved the earliest ripe date several
weeks even earlier, albeit at the expense of
fruit size and eating quality. Together Prince
and Weinberger released 21 peaches, one nec-
tarine and ‘Nemaguard’ rootstock. Only a
few are still being heavily planted although
‘Nemaguard’ is still the major peach rootstock
in California. One of Weinberger’s selections,
FV89-14, spawned an amazing range of descen-
dants although it was too susceptible to bacte-
rial spot for release on its own. Its east and
west coast progeny include ‘Springcrest’ (and
all its descendants) and many others (Table
6.2). ‘Springcrest’ at one time dominated early
peach production in California and Europe
(Okie and Myers, 1991).
North Carolina
F.E. Correll Jr was hired to start a peach breed-
ing programme at North Carolina State in
1955. About 10 years earlier C. Clayton had
been hired as a fruit pathologist. Clayton had
a good horticulture background and was very
active in the efforts to help growers. It was
apparent to him that the peach industry
needed cultivars resistant to bacterial spot
disease if it was going to thrive on the state’s
sandy soils. After Correll was hired the two
of them cooperated in one of the first cross-
discipline team efforts to develop disease-
resistant tree fruits. Although their cultivars
were not widely grown outside North Caro-
lina, the project was successful and their close

Table 6.2. Peach and nectarine cultivars descended from US Department of Agriculture selection
FV89-14, listed by most immediate parent. Parents are shown in bold and bud mutations in italics.
First-generation offspring are underlined. (From Okie and Myers, 1991.)

FV89-14 ‘Springcrest’ ‘Maycrest’ Other parenta

‘Autumn Red’ ‘Cristelle’ (=‘Primecrest’) ‘Early Maycrest ’ ‘Crimson Lady’

‘Camden’ ‘Earlicrest’ ‘Michaelian (Ra-2)’ —'Crimson Princess'
‘Fayette’ ‘Early Crest ’ (=‘San Isidoro’) ‘Queencrest ’ —‘Snow Duchess’
‘Flamecrest’ ‘Early Maycrest ’
‘Flavorcrest’ ‘Firecrest ’ ‘Earlitreat’ ‘Crown Princess’
‘Goldcrest’ ‘Maycrest’ ‘Gayla Rich’ —'Golden Princess'
‘Goldprince’ ‘Morning Sun’ ‘Kay Glo’ —‘Candy Red’
‘Gulfcrest’ ‘Queencrest’ ‘Klondike White’ —‘Ivory Queen’
‘Gulfking’ ‘Ray Crest ’ ‘May Sweet’ —‘Ivory Princess’
‘Spring Baby’ ‘Ruby May ’ ‘Polar Light’
‘Springcrest’ ‘Starcrest ’ (=‘Chastar ’) ‘Rich May’ ‘Fayette’
‘Spring Gem’ ‘Siesta Gem’ —‘June Crest’
‘Springold’ ‘Ambercrest’ ‘Spring Treat’ —‘PP16,179’
‘Starlite’ ‘Crimson Lady’ ‘Snow Dance’ —‘Super Lady’
‘Sunprince’ ‘Crown Princess’ ‘Snow Kist’ —‘Topcrest’
‘TexKing’ ‘Golden Crest’ ‘Snow Peak’
‘Honey Bee’ ‘Sunlit Snow’ ‘Topcrest’
‘Snow Duchess’ ‘Super Rich’ —‘Bev’s Red’
‘Springprince’ ‘Sweet Alice’ —‘Snow Prince’
‘Sugar Time’ ‘Sweet Crest’
‘Supecheight’ ‘Vista Snow’
‘Supechnine’ ‘Zee Diamond’
‘Zee Fire’
aIndented cultivars are descended from parent immediately above.
146 W.R. Okie et al.
cooperation was a model for others coming
after them. Correll also put great emphasis on
low flesh browning as a project goal. The peach
breeding was successful in developing a series
of bacterial spot-resistant cultivars, the most
resistant of which were ‘Clayton’ and ‘Candor’.
Other popular cultivars were ‘Norman’, ‘Win-
blo’ and ‘Biscoe’ (Werner and Ritchie, 1982).
In recent years the programme under
D.J. Werner emphasized late bloom and abil-
ity to crop despite spring frosts. ‘Contender’
has become popular for this reason, and the
newest releases ‘Intrepid’ and ‘Challenger’
are also quite bud hardy. Werner also released
a series of columnar (‘pillar’) shaped orna-
mentals with double flowers: ‘Corinthian
Pink’, ‘Corinthian Rose’ and ‘Corinthian White’,
along with ‘White Glory’, a weeping white-
flowered nectarine. The North Carolina pro-
gramme continues evaluations but is no
longer making new crosses.
Louisiana
Louisiana began peach breeding in 1951 under
P.L. Hawthorne and later J.C. Taylor, although
the Calhoun Research Station had been involved
in cultivar testing since 1889. The emphasis
was on disease resistance, mainly to bacterial
spot, reliable cropping and the fruit attributes
of size, attractiveness, flavour and shipping
quality. Some of the most important cultivars
released were ‘La Premiere’ (1965), ‘Surecrop’
(1973), ‘Harvester’ (1973) and ‘Majestic’ (1979).
‘Harvester’ has been one of the primary
south-eastern US cultivars since its release
(Graham, 1999). A secondary programme at
Idlewild, Louisiana developed moderate-chill
peaches for local markets, including low-acid
types popular among certain local consumers.
California
As early as 1860 there were already over a
million peach trees in California. By 1886 Cal-
ifornia was shipping boxes of ‘Alexander’,
‘Early Crawford’, ‘Late Crawford’ and ‘Hales
Early’ to the eastern USA. At about this same
time the drying industry began to expand.
‘Lovell’ and ‘Muir’, discovered as chance seed-
lings, were found to be well suited to drying.
By 1910 dried production was over 20,000 t
(Butterfield, 1938). After ‘Elberta’ and ‘J.H.
Hale’ were introduced, they dominated the
shipping industry until about 1940. Burbank’s
‘July Elberta’ (probably no relation to ‘Elberta’),
released in 1930, was the only cultivar of his to
be commercially important and the second
peach to be patented (US Plant Patent 15). But-
terfield (1938) lists 61 mostly chance seedling
peach cultivars that had originated in Califor-
nia by that time along with this comment:
‘With all of the many California peach culti-
vars shown in the accompanying list, it would
seem that nothing else would be desired’.
University of California
Public peach breeding in California had begun
at University of California at Riverside in
1907 to develop low-chill peaches suited for
southern California. A similar programme was
begun in 1919 at Chaffee Junior College in
Ontario, California. In the process of develop-
ing lower-chill peaches, W.E. Lammerts also
did the most thorough study of inheritance of
ornamental features such as multiple petals
(double flowers) (Lammerts, 1945). The most
important release from the low-chill work
was ‘Babcock’, a crisp (maybe it is a stony-hard
flesh?), low-acid, white-fleshed peach that was
the precursor to the low-acid peaches that are
currently in vogue, and in fact was grown for
many years as a niche market (Weldon and
Lesley, 1933; Butterfield, 1938). It was also
apparently a source, through its offspring ‘Giant
Babcock’, of the very firm texture of some
modern low-acid whites such as ‘White Lady’.
University breeders had little impact on free-
stone peach breeding, although University of
California at Davis did release several culti-
vars in 1977 such as ‘Firered’ and ‘Calred’.
P.E. Hansche also put substantial effort into
developing dwarf peaches with commercial
fruit quality, releasing several cultivars includ-
ing ‘Valley Red’ in 1989 (Hansche and Beres,
1989). So far dwarf peaches have not had
commercial impact due to management issues,
but breeding continues in Italy.
As the shipping industry developed,
there was a need for new cultivars over a
wider season that could be picked more firm,
with better colour and shipped over greater
 Fresh Market Cultivar Development 147

distances. Up to that time most freestone cul-
tivar development was conducted by federal
and state research programmes in the eastern
USA. Two private breeders, G. Merrill and
F.W. Anderson, initiated programmes in Cali-
fornia to breed cultivars especially for the
developing California industry. These two
pioneers remained friends throughout their
long careers and it is reported that, early in
their work, they established a ‘gentleman’s
agreement’ that Merrill would focus on peach
breeding and Anderson would concentrate
on nectarines (J. Slaughter, California, 2004,
personal communication).
Merrill
‘Gemfree’, ‘Merrill Gemfree’, ‘June Lady’, ‘Red
Lady’, ‘Halloween’, ‘Franciscan’, ‘Forty Niner’,
‘Sundance’ and others, many of which are still
being grown worldwide. During the 1970s and
early 1980s, up to 40% of all peaches shipped
in California were Merrill cultivars (Zaiger,
1988). Moreover, three Merrill peaches, ‘Ele-
gant Lady’, ‘Angelus’ and ‘O’Henry’, are still
among the top planted cultivars in California
today (Tables 6.3 and 6.4). Merrill worked with
various nurseries in California who assessed
tree royalty charges to commercial growers.
Merrill remained active in cultivar develop-
ment until shortly before his death in 1973
and his work helped to establish the next gen-
eration of breeders (Stark, 1974).

G. Merrill (1899–1973) worked mainly with Anderson

peaches, introducing important cultivars such
as ‘O’Henry’, ‘Early O’Henry’, ‘Elegant Lady’, Nectarines were a novelty crop in California
‘Spring Lady’, ‘Angelus’, ‘Sparkle’, ‘Parade’, in the middle of the last century, as the fruit

Table 6.3. Leading California peach cultivars in order of production averaged over 2002–2003.
(From CTFA, 2003.)

Cultivar Harvest starts: day/month/year Production (t)

‘O’Henry’ 29/7/2003 23,099

‘Elegant Lady’ 4/7/2003 21,137
‘Rich Lady’ 7/6/2003 10,001
‘Autumn Flame’ 11/8/2003 9,290
‘Summer Lady’ 21/7/2003 9,012
‘Zee Lady’ 15/7/2003 8,624
‘Ryan Sun’ 7/8/2003 7,890
‘Crimson Lady’ 23/5/2003 7,880
‘September Sun’ 23/8/2003 7,846
‘Snow Giant’ 31/7/2003 6,621
‘Summer Sweet’ 30/6/2003 6,068
‘Crown Princess’ 27/5/2003 5,660
‘Brittney Lane’ 29/5/2003 5,404
‘Fancy Lady’ 13/6/2003 5,341
‘Flavorcrest’ 11/6/2003 5,080
‘Diamond Princess’ 30/6/2003 4,705
‘September Snow’ 16/8/2003 4,628
‘Queencrest’ 13/5/2003 4,535
‘Spring Snow’ (40GH121) 14/5/2003 4,325
‘Snow King’ 23/7/2003 4,182
‘Autumn Snow’ (‘Yukon King’) 29/7/2003 3,890
‘Sweet Scarlet’ 30/5/2003 3,776
‘August Lady’ 5/8/2003 3,772
‘Super Rich’ 6/5/2003 3,364
‘Saturn’ 5/6/2003 3,061
‘Ivory Princess’ 3/6/2003 3,044
148 W.R. Okie et al.

Table 6.4. Peach tree sales reported by California nurseries, 1999–2002 (three planting years), arranged by season of
ripening. (From CTFA, 2003.)

Cultivar Season Ripe Flesha Acidityb Released Origin Trees Per cent

Early
‘Burpeachfourteen’ V. early 8 May Y M 2003 Burchell 69,000 4.0
‘Super Rich’ V. early 10 May Y M 1997 Zaiger 142,406 8.2
‘Queencrest’ V. early 12 May Y M 1987 L. Balakian 35,489 2.0
‘Burpeachone’ Early 19 May Y M 2001 Burchell 83,100 4.8
‘Spring Snow’ Early 24 May W L 1997 Zaiger 67,713 3.9
‘Crimson Lady’ Early 25 May Y M 1992 Bradford 27,878 1.6
‘Brittney Lane’ Early 29 May Y M 1998 Zaiger 71,493 4.1
‘Ivory Princess’ Early 6 Jun W L 2000 Bradford 41,754 2.4
‘Earlirich’ Early 7 Jun Y M 1994 Zaiger 39,237 2.3
‘Rich Lady’ Early 10 Jun Y M 1990 Zaiger 22,570 1.3
Others 146,608 8.4
Total 747,248 43.1

Mid-season
‘Vista’ Mid 15 Jun Y M 1996 Zaiger 25,387 1.5
‘Country Sweet’ Mid 18 Jun Y L 1999 Zaiger 71,950 4.1
‘Burpeachfive’ Mid 1 Jul Y M 2002 Burchell 56,632 3.3
‘Summer Sweet’ Mid 7 Jul W L 1992 Zaiger 24,532 1.4
‘Elegant Lady’ Mid 19 Jul Y M 1979 Merrill 57,513 3.3
‘Sweet Dream’ Mid 20 Jul Y L 1998 Zaiger 42,792 2.5
‘Burpeachsix’ Mid 21 Jul Y M 2002 Burchell 45,415 2.6
‘Angelus’ Mid 25 Jul Y M 1966 Merrill 23,348 1.3
‘Zee Lady’ Mid 31 Jul Y M 1986 Zaiger 26,100 1.5
‘O’Henry’ Mid 5 Aug Y M 1970 Merrill 35,273 2.0
Others 268,633 15.5
Total 677,575 39.0

Late
‘Autumn Snow’ Late 15 Aug W L 1997 Zaiger 19,311 1.1
‘Ryan Sun’ Late 20 Aug Y M 1983 Chamberlin 30,729 1.8
‘September Snow’ Late 28 Aug W L 1992 Zaiger 26,927 1.6
‘Burpeachfour’ Late 31 Aug Y M 2002 Burchell 29,000 1.7
‘Burpeachthree’ Late 1 Sep Y M 2002 Burchell 17,000 1.0
‘September Sun’ Late 2 Sep Y L 1987 Chamberlin 13,697 0.8
‘Fairtime’ Late 5 Sep Y M 1968 USDA 44,682 2.6
‘Autumn Flame’ Late 7 Sep Y M 1996 J. Doyle 55,689 3.2
‘Snowfall’ Late 8 Sep W L 2000 Zaiger 12,239 0.7
‘Sweet September’ Late 14 Sep Y L 1997 Zaiger 42,887 2.5
Others 18,261 1.1
Total 310,422 17.9

TOTAL PEACHES 1,735,245

USDA, US Department of Agriculture.

aFlesh colour: Y, yellow; W, white.
bAcidity type: M, standard; L, sub-acid.

was small, with little red skin colour, unat-
tractive green ground colour, and a tendency
for the skin to crack (Ramming, 1988). ‘Stan-
wick’, ‘Gower’ and ‘Quetta’ (Werner and
Okie, 1998) dominated California production.
F.W. Anderson, a private breeder in Merced,
California, began intercrossing of nectarines
such as ‘Quetta’ (from Pakistan) and ‘Lippi-
att’s’ (from New Zealand) with large peaches.
The release in 1942 of ‘Le Grand’ nectarine,
 Fresh Market Cultivar Development 149

which was the first large, attractive nectarine,
was the basis of the nectarine industry world-
wide. Most of our current nectarines can be
traced back to ‘Le Grand’ or his other releases.
As a result Anderson is widely known as the
‘father of the modern nectarine industry’. He
introduced many other important cultivars
including ‘Sun Grand’, ‘Red Grand’, ‘Grand
Haven’, ‘May Grand’, ‘Early Sun Grand’, ‘Sum-
mer Grand’, ‘Spring Grand’, ‘Red June’, ‘June
Grand’, ‘Aurelio Grand’, ‘Red Diamond’, ‘Sum-
mer Beaut’, ‘Spring Red’ and ‘Autumn Grand’.
Anderson’s nectarines dominated markets
throughout the world for many years, and
during the 1970s over 90% of all nectarines
shipped in California were his introductions
(Zaiger, 1988). Currently only ‘Red Diamond’
remains a major cultivar (Table 6.5). Anderson
worked with commercial nurseries who asses-
sed a per-tree royalty to commercial growers
and with Stark Brothers Nursery for sales of
backyard trees (L.G. Bradford, California,
2004, personal communication).
Not only was Anderson influential in the
fruit industry, he had an important influence
in establishing the next generation of California
breeders as well. N.G. Bradford, who went on
to establish Bradford Genetics, started working
with Anderson as a farm hand in 1941, became
general foreman in 1950, and continued until
Anderson’s death in 1982. C.F. Zaiger also
worked with Anderson during 1956–1957 and
went on to establish Zaiger Genetics, one of
the most important breeding programmes
today. Anderson remained active in breeding
late into his 80s and sold his programme to
N.G. Bradford shortly before his death in 1982
(Bradford, 1988).

Table 6.5. Leading California nectarine cultivars in order of production averaged over 2002–2003.
(From CTFA, 2003.)

Harvest starts:
Cultivar day/month/year Production (t)

‘Spring Bright’ 10/6/2003 21,998

‘Summer Fire’ 19/7/2003 15,812
‘Summer Bright’ 4/7/2003 15,372
‘August Red’ 12/8/2003 11,641
‘Rose Diamond’ 23/5/2003 11,343
‘September Red’ 22/8/2003 8,558
‘Ruby Diamond’ 23/6/2003 8,512
‘Diamond Bright’ 2/6/2003 7,948
‘Arctic Snow’/‘White Jewel’ 2/6/2003 7,947
‘Red Jim’ 7/8/2003 6,598
‘Fire Pearl’ 12/7/2003 6,179
‘Bright Pearl’ 5/7/2003 5,800
‘Arctic Pride’ 8/8/2003 5,631
‘Diamond Ray’ 30/6/2003 5,188
‘Honey Blaze’ 7/6/2003 4,925
‘Grand Pearl’ 30/6/2003 4,889
‘Red Diamond’ 25/6/2003 4,425
‘Mayglo’ 13/5/2003 4,322
‘Royal Glo’ 21/5/2003 4,252
‘Arctic Sweet’ 6/6/2003 4,163
‘Honey Kist’ 12/6/2003 3,779
‘Arctic Mist’ 6/9/2003 3,723
‘Fire Sweet’ 15/7/2003 3,610
‘Arctic Star’ 24/5/2003 3,461
‘Summer Blush’ 12/8/2003 3,437
‘Mayfire’ 8/5/2003 3,147
150 W.R. Okie et al.
US Department of Agriculture
USDA began peach breeding in California in
1920 at Palo Alto and later Davis, primarily to
develop better canning and drying peaches.
Later the cling breeding was shifted to Uni-
versity of California at Davis and the fresh
market breeding was expanded by J.H. Wein-
berger, who had worked in fruit breeding
with the USDA in Fort Valley, Georgia from
1937 and moved to Fresno, California in 1954
to continue his breeding work (Okie et al.,
1985). Although Weinberger grew relatively
few seedlings (about 4000 total stone fruits per
year in Fresno, which included plums and
apricots), he had clear goals in mind and a
discerning eye to find the best in his material.
His PhD training in plant physiology helped
him to ‘know his organism’ and he published
important papers on chilling and adaptation.
Extensive on-farm testing ensured the named
cultivars were likely to be successful. During
a period in the 1980s Weinberger’s nectarines,
including the leading cultivar at the time,
‘Fantasia’, were responsible for over 40% of
total nectarine production in the USA. Other
important Weinberger nectarines at the time
included ‘Fairlane’, ‘Firebrite’, ‘Flamekist’,
‘Flavortop’ and ‘Independence’. Weinberger
peach cultivars comprised over 25% of the
peaches shipped in California at this time,
including ‘Coronet’, ‘Desert Gold’, ‘Fairtime’,
‘Fayette’, ‘Flamecrest’, ‘Flavorcrest’, ‘Redtop’,
‘Regina’, ‘Summerset’ and ‘Suncrest’. ‘Flavor-
crest’ and to a lesser extent ‘Springcrest’ are the
only cultivars of his that are still significant in
production (Table 6.3). He also developed the
leading nematode-resistant rootstocks, ‘Nem-
aguard’ and ‘Nemared’, on which over 90%
of stone fruit are grown in California (Childers
and Sherman, 1988). Weinberger’s prolific
success, and the fact that USDA cultivars are
available worldwide with no royalty charge,
caused uncertainty with private breeders
during the 1970s and 1980s, and prompted
C.F. Zaiger in 1988 to refer to the private
breeder as ‘a dying breed’ (Zaiger, 1988). After
his retirement from the USDA in 1973, Wein-
berger consulted with Superior Farming
Company (now Sun World International, Inc.)
to help establish their breeding programme
near Bakersfield, California and remained
active in the fruit industry until shortly before
his death in 2000.
6.4. Modern US Breeding Programmes
Most breeding for fresh market peaches fol-
lows similar protocols. Advanced selections
and cultivars from in-house as well as other
programmes are evaluated to find superior
parents. Parents with superior characteristics
are crossed by hand with the objective of pro-
ducing seedlings that combine the desirable
qualities of the parents. Seed from the cross-
ings (or open-pollinated seed of previous
seedlings) is usually artificially stratified.
Seedlings grown in a greenhouse can be
planted the spring following harvest, saving
time, or moved to a nursery for a year. Small
seedlings are transplanted in the spring to
seedling rows, usually with trees spaced 0.5–1 m
apart. At this spacing trees become crowded
after several years so must be evaluated and
quickly rogued. Seedling trees usually bear
fruit in the third summer from planting, at
which time the best are selected for further
testing or propagation and the rest removed.
Seedling trees can be readily evaluated for
most visual fruit characters, but fruit usually
is smaller than on a commercially grown tree.
Eating quality is the most difficult aspect to
judge, as a taste test of a single fruit is often
not representative of the entire tree. In cli-
mates that are highly variable from year to
year, more years of observations are needed
to get a true picture of the selection’s adapta-
tion because many characters such as crop,
size, shape and quality can vary greatly from
year to year. One means to reduce the time
factor for such areas is to plant budded trees
in several widespread locations. A few breed-
ers do complete evaluations of every seed-
ling, but most find this too time-consuming
except for selected populations. Most public
breeders grow from 1000 to 5000 seedlings
per year, which is all their resources of land
and labour will allow. Some of the private
programmes may grow ten times this quan-
tity, which naturally allows for more rapid
progress. The timeframe from cross to release
is usually 10–15 years for a public programme,
 Fresh Market Cultivar Development
but frequently private breeders do less testing
and thus have a shorter turnover time.
In the last several decades, public peach
breeding in the USA has been reduced, par-
ticularly in marginal climates where peach
production is more challenging; industries
have shrunk and industry support has waned.
State programmes have also been hurt by
budget cuts (at both the state and local levels)
since tree fruit breeding is expensive, long term
and less dramatic than some other research
areas. Many of the programmes never had
much impact, a result of inadequate resources
of time and money, as well as climatic prob-
lems. It is difficult to make progress when
crosses and seedling crops are often killed by
frost. In contrast to the recent US situation,
countries in Europe and Asia have recognized
the value of developing cultivars specifically
adapted for their climate and industry. These
new programmes have a big advantage in
having much better parents to choose from
due to the last century of improvement in the
USA. For example, obtaining selections with
high red skin colour is facilitated if a breeder
uses the newer full-red peaches such as
‘O’Henry’ and ‘Rich May’ as parents.
North-eastern USA and Canada
Major cultivars grown in the north-eastern
USA and Michigan are shown in Table 6.6.
Most of these are from northern breeding pro-
grammes, or are mutations of northern culti-
vars. In general the cultivars are less recently
released compared with what is grown in
California, a reflection of the breeding effort
that has been available to produce new
improved cultivars.
New Jersey
The New Jersey breeding programme has con-
tinued under J.C. Goffreda and A. Voordeckers
with a dual emphasis. Recent selections include
standard yellow- and white-fleshed peaches
with resistance to bacterial spot and constric-
tion canker (Fusicoccum or Phomposis canker). In
addition they have emphasized niche markets,
including flat (peento or pantao) peaches,
low-acid and stony-hard (crisp) flesh, also
151
with disease resistance. ‘Saturn’ (NJF-2),
probably the first bred flat peach, was named
by Starks Nursery in 1985 but languished
until the late 1990s when California growers
began marketing it as a specialty item (Table
6.3). The programme plans on releasing ten or
so cultivars in the next couple of years, includ-
ing a series of stony-hard peaches, at least
three flat peaches, one nectarine and several
melting white- and yellow-fleshed peaches.
Rutgers also released a cold-hardy, late-
blooming ornamental, ‘Jerseypink’.
US Department of Agriculture–Agricultural
Research Service
The Beltsville breeding work (now USDA–
Agricultural Research Service or USDA-ARS)
has been continued at Kearneysville, West
Virginia, by R. Scorza. Releases ‘Bounty’ and
‘Sentry’ have become important north-eastern
USA cultivars (Table 6.6). ‘Earliscarlet’ pro-
vides a brightly coloured, adapted nectarine
for eastern US growers. Non-standard tree
growth habits have been a focus in recent
years, and much has been done to understand
the different forms and patterns of growth.
Advanced selections include those with semi-
dwarf and compact growth habits, which
reduce tree size and pruning while maintain-
ing a manageable size. Two new releases are
the first fruit type cultivars using the columnar
(‘pillar’) gene. ‘Crimson Rocket’ is homozy-
gous for the gene and is columnar in habit;
‘Sweet-N-Up’ is heterozygous, which gives it
an intermediate upright architecture. These tree
shapes are conducive to more dense plantings.
Michigan
The Michigan State University peach breed-
ing programme since 1992 has been under the
direction of W. Shane who has expanded work
initiated by A.F. Iezzoni, now primarily a
cherry breeder. The traditional breeding pro-
gramme is developing yellow (mostly) and
white, melting-flesh cultivars in the post-
‘Redhaven’ season with better skin colour, size,
firmness, productivity and disease resistance.
Particular emphasis is placed on mid-winter
cold hardiness and resistance to Leucostoma
canker, which causes serious limb decline in
152 W.R. Okie et al.

Table 6.6. Major fresh market peach cultivars in the north-eastern and south-eastern USA, listed by ripening
date, with source and release date. (In part from J. Frecon, USA, 2005, personal communication.)

Cultivar Origin Year

North-eastern USA
‘Harrow Diamond’ Ontario 1984
‘Harrow Dawn’ Ontario 1996
‘Sentry’ USDA-ARS, West Virginia 1980
‘Garnet Beauty’ (=‘Redhaven’ mutation) Ontario 1958
‘Redhaven’ Michigan AES 1940
‘JohnBoy’ (=‘Loring’ mutation) New Jersey AES 1988
‘Harrow Beauty’ Ontario 1983
‘Flamin Fury PF17’ Michigan – Friday 1993
‘Bounty’ USDA-ARS, West Virginia 1988
‘Loring’ Missouri AES 1946
‘Flamin Fury PF23’ Michigan – Friday 1993
‘Blake’ New Jersey AES 1953
‘Harcrest’ Ontario 1983
‘Cresthaven’ Michigan AES 1963
‘Jerseyqueen’ New Jersey AES 1964
‘Encore’ New Jersey AES 1980
‘Laurol’ (=‘Jerseyqueen’ mutation) New Jersey 1992
‘Flamin Fury PF24-007’ Michigan – Friday 1996
‘Redskin’ Maryland AES 1944
South-eastern USA
‘Springprince’ USDA-ARS, Georgia 1998
‘Sunbrite’ USDA-ARS, Georgia 1976
‘Goldprince’ USDA-ARS, Georgia 1989
‘Rubyprince’ USDA-ARS, Georgia 1997
‘Summerprince’ USDA-ARS, Georgia 1992
‘Juneprince’ USDA-ARS, Georgia 1985
‘GaLa’ USDA, Louisiana 1992
‘Harvester’ Louisiana AES 1973
‘Cary Mac’ (=‘Loring’ mutation) South Carolina 1976
‘Redglobe’ USDA-ARS, Maryland 1954
‘Majestic’ Louisiana AES 1979
‘Summergold’ USDA-ARS, Georgia 1970
‘Fireprince’ USDA-ARS, Georgia 1985
‘Contender’ North Carolina AES 1987
‘Cresthaven’ Michigan AES 1963
‘Sunprince’ USDA-ARS, Georgia 1981
‘O’Henry’ California – Merrill 1970
‘Flameprince’ USDA-ARS, Georgia 1993
‘Big Red’ (unreleased) USDA-ARS, California ~1980
‘Autumnprince’ USDA-ARS, Georgia 1998

USDA-ARS, US Department of Agriculture–Agricultural Research Service; AES, Agricultural Experiment

Station.

colder climates. In the early 1990s Iezzoni and were then used as parents to move the resistance
others initiated a long-term project to develop into high-quality peaches. The programme has
canker-tolerant peaches. They identified peach generated 2000 to 3000 seedlings per year,
selections with resistance to canker, which resulting in the naming of ‘Beaumont’ in 2004.
 Fresh Market Cultivar Development
Fruit Acres
The scarcity of new cultivars adapted to the
north-east in the last 20 years prompted a
resurgence in private breeding in Michigan,
home to two breeding programmes started by
growers, who run them in conjunction with
commercial peach orchards. The ‘Fruit Acres’
programme was started by J.E. Friday and is
now run by his daughter and son-in-law, A.
and R. Bjorge, located in Coloma. They are
marketing the ‘Stellar’ series of peaches. Pop-
ular releases in more northern states include
‘Coralstar’, ‘Glowingstar’ and ‘Redstar’.
‘Flamin Fury’
A second Michigan breeding programme was
started by cousin P.J. Friday, also in Coloma.
He has developed the ‘Flamin Fury’ or ‘PF-’
series of peaches. These cultivars have provided
a needed set of redder, bacterial spot-resistant
new peaches for the region. Some such as
PF12A, PF23 and PF17 have been well
accepted commercially, primarily in the
northern USA. More recently PF24-007 has
been widely planted due to its very large size.
Ontario
Breeding in Ontario has been consolidated
into a single programme at Vineland, with the
station now under the University of Guelph
rather than the provincial government. With
the retirement of N.W. Miles, breeding for late-
season canning peaches continues as an
emphasis under J. Subramanian. Unfortu-
nately the programme at Harrow was a victim
of budget cutting and in 1996 it was closed.
Advanced Harrow selections are at Vineland
for final testing. Subramanian will also be
aiming to develop mid- and late-season white-
and yellow-fleshed freestones with improved
bacterial spot resistance. It is unclear if nec-
tarine development will continue.
South-eastern USA
Major southern cultivars are listed in Table 6.6,
excluding moderate- and low-chill cultivars
which are discussed in Chapter 5. As with the
153
north-eastern USA, most are publicly bred
cultivars from within the region.
Arkansas
The University of Arkansas peach breeding
programme was begun in the mid-1960s by
J.N. Moore with strong cooperation with R.
Rom. The main focus of the programme was
canning cling peach breeding, primarily for
baby food use. The contribution from F.
Hough and Rutgers University was very sub-
stantial in the early years (see Chapter 7). The
programme focus shifted in the late 1990s
under J.R. Clark to emphasize fresh market
peaches and to continue the nectarine pro-
gramme that had been ongoing for some
years. ‘White River’ (2002) was the first fresh
market peach from the programme, and was
followed in 2004 by low-acid types ‘White
County’ and ‘White Rock’ (Clark et al., 2005).
Nectarines released from the programme are
‘Westbrook’, ‘Arrington’ and ‘Bradley’ (all in
2000). The latter two nectarines are non-melt-
ing flesh types. The programme objectives
include the continued improvement of nec-
tarines, with emphasis on white- and yellow-
fleshed types with melting and non-melting
flesh. Fresh market peach breeding focuses
primarily on white-fleshed peaches with non-
melting flesh. Sub-acid and standard acidity
types are included in the programme as are
free- and clingstones. A small project to
improve flat peaches is also under way. Bacte-
rial spot resistance is a major priority in the
programme, as the programme is operated in
an area with high selection pressure for this
trait. The germplasm used in the programme
has been largely US cultivars and breeding
selections from other programmes. A series of
four ornamentals with dwarf (‘Bonfire’, ‘Lep-
rechaun’) or weeping (‘Crimson Cascade’,
‘Pink Cascade’) habits was released in 1994.
Louisiana
Peach breeding in Louisiana ebbed in the
1980s after a series of difficult winters. Under
C.E. Johnson and later C.J. Graham, the pro-
gramme expanded to develop adapted
peaches with high productivity and disease
resistance. Unfortunately budget cuts resulted
154 W.R. Okie et al.
in the termination of the programme in 2004.
Recent releases from the moderate-chill pro-
gramme at Clinton now under C.E. Johnson
include ‘LaBelle’, ‘LaRouge’ and ‘LaSweet’.
North Carolina
In recent years the programme under D.J.
Werner emphasized late bloom and ability to
crop despite spring frosts. ‘Contender’ has
become popular for this reason and the newest
releases ‘Intrepid’ and ‘Challenger’ are also
quite bud hardy. ‘China Pearl’ was released to
provide a late-blooming, white-fleshed peach.
Werner also released a series of pillar-shaped
ornamentals with double flowers: ‘Corinthian
Pink’, ‘Corinthian Rose’ and ‘Corinthian White’,
along with ‘White Glory’, a weeping white-
flowered nectarine. Other recent releases
include ‘Carolina Gold’, a yellow, melting-flesh,
late-season cultivar, and ‘Galactica’, a large,
low-acid, white peento. The North Carolina
programme continues evaluations but is no
longer making crosses.
US Department of Agriculture–Agricultural
Research Service
The south-eastern US peach industry is plagued
by variability in winter climate, which makes
it difficult to select cultivars that get adequate
chill every year but do not bloom too early
and suffer from frost damage. The USDA-
ARS peach breeding programme at Byron,
Georgia is now the primary source of new
commercial cultivars for the south-eastern
USA, given the reduction in programmes in
adjacent states. Peach growers need a
sequence of cultivars ripening from 60 days
before, to 30 days after, ‘Elberta’ (mid-July at
Byron). The cultivar must be consistently pro-
ductive, giving large-sized, firm and well-
blushed fruit. Critical time slots lacking
suitable cultivars are the period before ‘Spring-
crest’ and the seasons 1–2 weeks before and
after ‘Elberta’. New releases ‘Scarletprince’,
‘Julyprince’, ‘Early Augustprince’ and ‘August-
prince’ may fill the late-season niche. Most of
the 4000 seedlings planted annually in recent
years have been aimed at developing mid-
and late-season peaches, rather than very
early. Since 1980 all yellow-fleshed peaches
have used ‘-prince’ as part of the name, to hon-
our V.E. Prince’s efforts in establishing the
Byron facility and to provide a ‘brand’ name
for growers. The most successful releases have
been ‘Juneprince’, ‘Goldprince’, ‘Sunprince’,
‘Flameprince’, ‘Rubyprince’ (Fig. 6.1/Plate 54)
and ‘Springprince’ (Okie, 1997). White-fleshed
peaches ‘Scarletpearl’ and ‘Southern Pearl’,
as well as white-fleshed nectarine ‘Roseprin-
cess’, have been useful for local markets. Aux-
iliary interests are in chilling as it relates to
the possibility of developing a low-chill but
high-heat requirement peach, narrow-leaf
trees, and novelty fruits including blood-flesh
and flat shape.
California
The California Tree Fruit Agreement (CTFA)
reports shipments of over 170 different peach
and nectarine cultivars that provide fresh
fruit from April through October (Tables 6.3
and 6.5). Private breeders dominate the field
of cultivar development in California due to
reduced funding of government breeding
programmes and to improvements in their
ability to work globally and enforce intellec-
tual property (IP) rights. The main private
California breeding programmes in terms of
size are Zaiger Genetics, Inc., Bradford Genet-
ics, Inc., Burchell Nursery, Inc. and Sun World
International, Inc. In order to remain compet-
itive, all these programmes have developed
strong international programmes and may
derive as much or more revenue from inter-
national sources as from domestic sources.
Private breeders derive revenue from IP
licensing, often in the form of per-tree royalty
assessments collected by licensed nurseries.
There has also been a recent trend to develop
‘variety clubs’ and other production royalty
programmes with managed plantings and
development of ‘branded’ stone fruit series.
Many large growers and marketers world-
wide are turning to managed programmes
because of the perception that unmanaged
plantings lead to unhealthy overproduction
and reduced fruit quality. In the USA, the
Zaiger, Bradford and Burchell programmes
are linked closely to nursery companies that
 Fresh Market Cultivar Development
work with the programmes to test and pro-
mote new cultivars, and have exclusive rights
to tree sales of most of the programmes’ releases.
The result of these relationships is evident in
recent tree sales in California, where 76% of
peach trees and 84% of nectarine trees sold
during 1999–2002 were cultivars from these
three programmes (Tables 6.4 and 6.7).
US Department of Agriculture–Agricultural
Research Service
D.W. Ramming has continued peach breed-
ing at USDA in Fresno (now at Parlier) from
1974, emphasizing the very early and very
late seasons, as well as improved eating qual-
ity. His research on embryo culture as a means
to produce seedlings of early × early seedlings,
155
Fig. 6.1. Ripe fruit of ‘Rubyprince’
peach, a typical modern peach
with extensive red blush and short
pubescence.
which normally have low-viability seed, has
enabled the programme to develop much
improved ultra-early cultivars such as ‘May-
fire’ and ‘Crimson Baby’ nectarines as well as
‘Spring Baby’ and ‘Spring Gem’ peaches.
‘Spring Baby’ was one of the first non-melting
flesh peaches for the shipping market. Since
all early peaches are clingstone for physiolog-
ical reasons, the clingstone character associ-
ated with the non-melting flesh was not a
problem, and it could hang on the tree longer
without premature softening. The embryo-
rescue technique has since been adopted on a
larger scale by the private programmes, with
impressive results. ‘Autumn Red’ was released
to provide a high-quality, well-coloured, late-
ripening peach. ‘September Free’ is a high-
quality, freestone, late-ripening nectarine, as
 156
Table 6.7. Nectarine tree sales reported by California nurseries, 1999–2002 (3 planting years), arranged by season of ripening. (From CTFA, 2003.)

Cultivar Season Ripe Flesha Acidityb Released Origin Trees Per cent

Early
‘Mayfire’ V. early 7 May Y M 1983 USDA 34,083 2.9
‘Burnecten’ V. early 7 May Y M 2003 Burchell 23,677 2.0
‘Crimson Baby’ Early 23 May Y M 1999 USDA 27,258 2.3
‘Red Roy’ Early 27 May Y M 2001 Zaiger 30,700 2.6
‘Kay Sweet’ Early 30 May Y L 1999 Bradford 41,538 3.6
‘Royal Glo’ Early 1 Jun Y M 1993 Zaiger 25,902 2.2
‘Arctic Star’ Early 1 Jun W L 1995 Zaiger 23,590 2.0

W.R. Okie et al.

‘Diamond Bright’ Early 9 Jun Y M 1996 Bradford 59,197 5.1
‘Honey Blaze’ Early 11 Jun Y L 1998 Zaiger 64,480 5.6
‘Arctic Sweet’ Early 14 Jun W L 1996 Zaiger 18,863 1.6
Others 63,830 5.5
Total 413,118 35.6

Mid-season
‘June Pearl’ Mid 16 Jun W L 1995 Bradford 21,193 1.8
‘Spring Bright’ Mid 24 Jun Y M 1991 Bradford 30,449 2.6
‘Kay Pearl’ Mid 27 Jun W L 1999 Bradford 29,065 2.5
‘Ruby Sweet’ Mid 28 Jun Y L 1997 Bradford 30,806 2.7
‘Diamond Ray’ Mid 4 Jul Y M 1994 Bradford 29,315 2.5
‘Arctic Jay’ Mid 9 Jul W L 1997 Zaiger 22,159 1.9
‘Grand Pearl’ Mid 10 Jul W L 1997 Bradford 32,227 2.8
‘Honey Kist’ Mid 19 Jul Y L 1995 Zaiger 23,118 2.0
‘Fire Sweet’ Mid 27 Jul Y L 1997 Bradford 27,049 2.3
‘August Pearl’ Mid 13 Aug W L 1999 Bradford 35,095 3.0
Others 174,300 15.0
Total 454,776 39.2
Late
‘Regal Pearl’ Late 19 Aug W L 2000 Bradford 5,180 0.4
‘August Fire’ Late 20 Aug Y M 2000 N. Waldner 111,546 9.6
‘Arctic Pride’ Late 27 Aug W L 1993 Zaiger 20,058 1.7
‘August Red’ Late 31 Aug Y M 1988 Bradford 10,596 0.9
‘Arctic Snow’ Late 3 Sep W L 1992 Zaiger 41,330 3.6
‘September Bright’ Late 3 Sep Y M 2003 Bradford 7,146 0.6
‘September Free’ Late 5 Sep Y M 1999 USDA 11,696 1.0
‘Burnectfour’ Late 11 Sep Y M 2003 Burchell 20,600 1.8
‘Arctic Mist’ Late 13 Sep W L 1999 Zaiger 36,243 3.1

Fresh Market Cultivar Development

‘September Red’ V. late 19 Sep Y L 1986 Bradford 28,320 2.4
Others 0 0.0
Total 292,715 25.2

TOTAL NECTARINES 1,160,609

aFlesh colour: Y, yellow; W, white.
bAcidity type: M, standard; L, sub-acid.

157
158 W.R. Okie et al.
an alternative to the many late clings on the
market. ‘Mayfire’, ‘Crimson Baby’ and Septem-
ber Free’ have been widely planted in recent
years (Table 6.7). Most recently, ‘Galaxy’ was
released to provide a large flat peach as an
adjunct to ‘Saturn’. It is the largest flat peach
currently available. Although ARS cultivars
had a reputation for high quality and success
due to extensive on-farm testing, industry
impatience with the pace of public breeding
in comparison to the much larger private pro-
grammes and their many new cultivars is
resulting in de-emphasizing of peach breed-
ing relative to the past.
Zaiger Genetics, Inc.
Zaiger Genetics is a family-owned breeding
organization that was founded in Modesto by
C.F. (Floyd) Zaiger and his wife in 1958 after
he had worked for two seasons as an appren-
tice with fruit breeding pioneer F.W. Ander-
son. Zaiger patented his first cultivars, ‘Royal
Gold’ peach and ‘Crimson Gold’ nectarine, in
the 1960s, and today the company holds over
200 US plant patents. Floyd has been the prin-
cipal breeder through much of the history of
the programme, but today he works together
with his children to direct the programme. The
Zaigers have achieved global prominence for
a robust and comprehensive breeding progra-
mme that includes peaches, nectarines, plums,
apricots, cherries and Prunus rootstocks.
Through the years Zaiger has been quick
to recognize opportunities and respond to
industry needs throughout the world. It started
Table 6.8. Series names used by private breeders for different types of peaches and nectarines.
Yellow
Fruit/breeder Acid Low-acid
Peach
Bradford ‘Princess’ ‘Candy’
Burchell ‘Flame’ ‘Sweet Flame’
Zaiger ‘Sweet’
Nectarine
Bradford ‘Bright’ ‘Sweet’
Burchell ‘Flare’ ‘Sweet Flare’
Zaiger ‘Glo’ ‘Honey’
relatively early with white-fleshed peach
development, beginning in the late 1960s to
develop white-fleshed cultivars for European
growers. In the 1990s, when California grow-
ers became interested in white-fleshed fruit
for Asian markets, Zaiger was ahead of other
programmes. Zaiger has also led other breed-
ers in the development of low-acidity, yellow-
fleshed cultivars, and was an early developer
of lower-chilling cultivars for California. In
the USA, Zaiger works with Dave Wilson
Nursery, which tests new cultivars in its own
commercial test block and sponsors fruit-
tasting panels. In order for growers to distin-
guish fruit types, Zaiger groups cultivars into
general series names (Table 6.8). Selected low-
acidity peaches and nectarines with premium
flavour are also denoted as members of the
‘Zee Sweet®’ managed production programme
and are licensed with production royalties in
certain parts of the world.
With peaches, 50% of recent tree plant-
ings in California have been Zaiger cultivars,
including the most heavily planted peach, the
early-season cultivar ‘Super Rich’, with 8.2%
of peach tree sales (Table 6.4). Two other early
peaches, ‘Brittney Lane’ (4.1%) and ‘Spring
Snow’ (3.9%), were heavily planted, as well as
the mid-season, low-acidity, yellow-fleshed
‘Country Sweet’ (4.1%). Zaiger peaches are
well distributed through the ripening season
with five of the top ten planted early- and
mid-season cultivars, and four of the top ten
late-season cultivars. With nectarines, 32%
of recent tree plantings in California have
been Zaiger cultivars, led by the low-acidity,
White
Acid Low-acid High quality
‘Ivory’, ‘Ice’, ‘Snow’ ‘Candy’
‘Snow Flame’ ‘Snow Flame’
‘Snow’
‘Pearl’ ‘Pearl’ ‘Candy’
‘Snow Flare’ ‘Snow Flare’
‘Arctic’
 Fresh Market Cultivar Development
yellow-fleshed ‘Honey Blaze’ cultivar with
5.6% of nectarine tree sales (Table 6.7). Other
important Zaiger nectarines include ‘Arctic
Snow’ (3.6%), ‘Arctic Mist’ (3.1%) and ‘Red
Roy’ (2.6%). Zaiger’s success in nectarines is
most evident in early-season cultivars which
account for five of the top ten cultivars planted
recently. Zaiger’s success in early-ripening
nectarines is largely due to its relatively
advanced embryo-rescue facilities. Addition-
ally, two of the top ten planted mid-season
nectarines and three of the top ten late-season
nectarines are Zaiger releases (Table 6.7).
Bradford Genetics, Inc.
Bradford Genetics is a division of Bradford
Farms, a family-owned organization in Le
Grand that was founded by N.G. Bradford,
who worked for the fruit breeding pioneer
F.W. Anderson from the early 1940s until
1981, when he purchased the Anderson pro-
gramme and breeding materials about a year
before Anderson’s death. N.G. Bradford was
the principal breeder for much of the history
of the Bradford programme and continues to
work with his son, L.G. Bradford, who directs
the programme today.
The Bradford programme started with
mostly mid-season, yellow-fleshed nectarines
with traditional acidity and a relatively high
chilling requirement. For the first 10 years,
Bradford built on Anderson’s success, aided
by superior breeding parents such as ‘Red
Diamond’ nectarine, which established a new
standard for firmness and colour when
released in 1972. Bradford nectarines contin-
ued to dominate the industry throughout the
1990s; however, increased competition from
other programmes and changes in the indus-
try encouraged expansion into other areas.
During the 1990s, Bradford made a major
shift towards development of white-fleshed
peaches and nectarines. The initial emphasis
was to develop acidic, white-fleshed cultivars
that were preferred by growers in France. How-
ever, many of the white-fleshed selections
turned out to be low-acidity types, which for-
tunately coincided with the sudden growth of
export markets in Asia, where low-acidity
types are preferred. The resulting ‘Pearl’ series
of low-acid, white-fleshed nectarines has been
159
very popular with growers in California and
in the southern hemisphere. The white-fleshed
peaches ‘Ivory Princess’, ‘Ice Princess’ and
‘Snow Princess’ have also been popular culti-
vars worldwide. Additionally, to remain com-
petitive, Bradford has made a shift towards
development of earlier-ripening cultivars and
cultivars with lower chilling requirement,
although a lack of high-capacity embryo-res-
cue facilities has slowed progress.
Today Bradford works closely with
Bright Brothers Nursery in California, which
tests new cultivars in commercial orchards
and has exclusive rights to most Bradford cul-
tivar tree sales in California. Bradford also
has a strong presence in most fruit-growing
regions in the world and L.G. Bradford reports
that today more Bradford trees are sold over-
seas than in the USA. In order for growers to
distinguish fruit types, Bradford groups culti-
vars into general series names (Table 6.8).
With nectarines, Bradford continues to
be a strong competitor and 45% of recent nec-
tarine tree sales were Bradford cultivars, led
by the early-ripening ‘Diamond Bright’ nec-
tarine with 5.1% of nectarine tree sales
(Table 6.7). Other important Bradford nectar-
ines include ‘Kay Sweet’ (3.6%), ‘August
Pearl’ (3%), ‘Grand Pearl’ (2.8%) and ‘Ruby
Sweet’ (2.7%). It is noteworthy that seven of the
top ten Bradford nectarines planted recently
are low-acidity types. Bradford continues to
be strongest in mid-season nectarines, with
eight of the top ten planted cultivars in Cali-
fornia. It is also strong in late-season cultivars,
with four of the top ten cultivars. However,
with early-ripening nectarines, only two of the
top ten planted cultivars are Bradford releases.
In spite of considerable progress with peaches,
only 6% of recent peach tree plantings in Cal-
ifornia were Bradford cultivars, including
‘Ivory Princess’ with 2.4% of tree sales and
‘Crimson Lady’ with 1.6% (Table 6.4). How-
ever, the new generation of premium-flavour
peaches in the ‘Candy’ series may increase
Bradford’s share in the future as growers
begin to plant the new cultivars.
Burchell Nursery, Inc.
Burchell Nursery is a family-owned nursery
company in Oakdale and Fowler, founded in
160 W.R. Okie et al.
1942 by I. Burchell, and led today by his son,
W.T. Burchell, and his grandson, T.W. Burchell.
The breeding programme was established so
that Burchell could remain competitive with
other large nurseries that had exclusive
arrangements with breeders. In 1985, Burchell
enlisted the aid of J.F. Doyle from the Univer-
sity of California to help establish a breeding
programme in Fowler, California. However,
by 1989, Doyle needed to return to a full-time
commitment with the University of Califor-
nia, and J. Slaughter along with T. Gerdts,
both Burchell field representatives, stepped
in to continue the programme. The release of
‘Autumn Flame’ peach, one of the top ten
planted peaches today, was a result of Doyle’s
contribution to the Burchell programme
(Tables 6.3 and 6.4).
To date more than 40 plant patents have
been filed. Burchell also works cooperatively
with D.H. Byrne at Texas A&M University, to
develop cultivars with low chilling require-
ment. New cultivar patents are filed with a
varietal name that has a prefix, ‘Burpeach’
(peaches) or ‘Burnect’ (nectarines), and a
sequential numerical suffix, resulting in a
name such as ‘Burpeachone’. In addition to a
varietal name, Burchell groups cultivars into
trade names so that growers and marketers
can distinguish fruit types (Table 6.8).
The Burchell programme is unique in
being wholly owned by a nursery company,
and breeders Slaughter and Gerdts also act as
salesmen, field representatives and interna-
tional licensing agents. They work closely
with growers to test new cultivars in com-
mercial orchards and offer technical support
for the cultivars they develop. With peaches,
20% of recent tree plantings in California have
been Burchell cultivars, led by the early-sea-
son peaches ‘Burpeachone’ (4.8% of peach
tree sales) and ‘Burpeachfourteen’ (4%), and
the mid-season peach ‘Burpeachfive’ (3.3%).
Burchell cultivars are well distributed through-
out the ripening season, with two of the top
ten planted cultivars in each of the early-,
mid- and late-season categories (Table 6.4).
Thus far all of the top-planted Burchell
peaches are acidic, yellow-fleshed types. With
nectarines 7% of recent tree plantings in Cali-
fornia have been Burchell cultivars, including
the early-season ‘Burnecten’ (2% of nectarine
tree sales) and the late-season ‘Burnectfour’
(1.8%), both acidic, yellow-fleshed types
(Table 6.7).
Sun World International, Inc.
After fruit breeding pioneer J.W. Weinberger
retired from the USDA-ARS in 1973, he con-
sulted with Superior Farming Company to
help establish a breeding programme near
Bakersfield, California to develop stone fruit
and table grape cultivars for the company’s
use. In the late 1980s, two fruit breeders joined
Superior Farming to continue the programme,
D.W. Cain for table grapes and C. Fear for
stone fruit breeding. When Sun World Inter-
national, Inc., a grower and marketer of fruits
and vegetables, purchased Superior Farming
in 1989, Fear left the company. Stone fruit
breeding, however, continued, first under
B.D. Mowery and later D.W. Cain, until 2000
when T.A. Bacon joined Sun World. Today
Bacon conducts a peach breeding programme
entirely focused on early-ripening cultivars,
primarily at Bakersfield. Sun World has high-
capacity embryo-rescue facilities and green-
houses required for breeding early-ripening
stone fruit. It also has one of the most robust
low-chill (200 chill units or less) breeding pro-
grammes in the world in California’s Coachella
Valley.
New Sun World cultivar patents are filed
with a name that has a prefix ‘Supech’ (peaches)
or ‘Sunect’ (nectarines) and a sequential
numerical suffix, resulting in a varietal name
such as ‘Supechsix’. Sun World groups culti-
vars as to fruit type and markets them under
umbrella brand names such as AMBER
CREST® brand peaches. Individual cultivars
are selected for the brands to provide a con-
tinuous series through the season, and can be
replaced with improved cultivars without
affecting brand name consistency. Currently
the AMBER CREST® brand peach series con-
sists of six yellow-fleshed, traditional-acidity
cultivars, beginning with the low-chill culti-
vars ‘Supechthirteen’ and ‘Supechfifteen’,
which are harvested in the Coachella Valley
during the first half of April. ‘Supechsix’ and
other cultivars grown further north in the
San Joaquin Valley follow on in May. With
nectarines, Sun World is evaluating several
 Fresh Market Cultivar Development
low-chill cultivars that ripen during April in
the Coachella Valley, and promotes ‘Sunec-
twentyone’, which ripens in the first week of
May in the San Joaquin Valley. The Sun World
stone fruit programme has until recently
developed new cultivars exclusively for Sun
World plantings in the USA, but now inter-
national licensing is also available.
6.5 European Programmes
For most of the 20th century, peach breeding
in countries outside the USA was limited. In
many places, cultivars from US breeding pro-
grammes were imported to replace local cul-
tivars that were no longer profitable to grow.
Through the latter half of the 20th century
there was limited breeding in Italy (notably
by A. Morettini in Florence), France (primarily
by R. Monet at Bordeaux), Australia (at Tatura
for canning peaches) and other countries. Many
national breeding programmes have been
initiated or expanded since the mid-1980s.
Bulgaria
The modern fruit industry in Bulgaria dates
back to the early 1950s. After 1960 it grew rap-
idly, particularly in terms of horticultural
advancements and new cultivars introduced,
thanks to the role played by breeding pro-
grammes. Peaches spread to vast areas of the
District of Sliven, well known in Bulgaria as
the ‘Valley of the Peaches’, because they were
well adapted there.
The first introduced cultivars in modern
orchards were ‘J.H. Hale’ and ‘Elberta’, which
were grown along with the locally well-
known, late-ripening ‘Dupnishka’. The same
cultivars were the starting point of breeding
programmes from the mid-1950s that resulted
in the introduction of ‘Zlatna Krichimka’ (bud
mutation of ‘J.H. Hale’) and ‘Septemvriiska’
(‘Elberta’ × ‘Karman’). Later, V. Velkov, Director
of the Fruit Growing Institute in Plovdiv from
1962 to 1972, introduced ‘Rumyana’, ‘Asenova
Krepost’, ‘Nay-Ranna Zhalta’ and ‘Velkova’.
The same institute became the leader of
an intense effort on peach breeding carried
161
out at the experimental stations in Sliven,
Pomorie and Petrich. Peach breeder Y. Grigo-
rov introduced for the fresh market ‘Bulgarska
Ranna’, ‘Yulska Edra’ and ‘Petrichka’, later
followed by ‘Tundzha 1’ and ‘Tundzha 2’ for
canning. At the same time another breeder, A.
Arangelov, carried out a number of genetic
studies on peach while also developing ‘Zlatna
Krichimka’, ‘Chervena Kurtovka’, ‘Elinpelin-
ska’ and ‘Zlatka’. By artificial mutagenesis A.
Angelov developed ‘Plovdiv 1’, ‘Plovdiv 2’,
‘Plovdiv 6’ and ‘Yoneta’.
Particularly important was the work of
S. D’bov (1974, 1975), who hybridized peach
with Prunus ferganensis for three decades, intro-
ducing cultivars resistant to powdery mildew
(Podosphaera pannosa (Wallr.:Fr.) Braun & Taka-
matsu): ‘Aheloy’ and ‘Remil’ for fresh market
and ‘Malo Konare’ and ‘Stoyka’ for canning.
Breeding programmes for peach were re-
started at Plovdiv in 1989 by A. Zhivondov
and his group. They are based on the progress
made in Bulgaria and worldwide but are
focused on the new horticultural and market
requirements. The major breeding objectives
are: developing cultivars resistant to powdery
mildew (with P. ferganensis as the source of
resistance) and leaf curl (Taphrina deformans
(Berk.) Tul.); widening the ripening season by
the introduction of early and late cultivars;
and establishing cultivars that are easy to
train with size-controlled trees (e.g. columnar,
dwarf, weeping).
France
The introduction of the first peach trees in
France is ascribed to the Romans under the
name of ‘malum persicum’ even though,
according to early Latin writers, the Gallic
people already knew a freestone fruit called
‘galicum’, larger than the ‘persicum’ intro-
duced by the Romans. Peach was truly wide-
spread in all of France during the Middle
Ages. Charlemagne recommended that all his
officers plant peach trees in their gardens.
Grafting of peaches has been practised since
the 16th century. The adoption of the ‘espalier’
(tree training on trellis or hedgerow) between
1635 and 1650 contributed to the improve-
ment and the expansion of its culture. The old
162 W.R. Okie et al.
peach cultivars were grown in the gardens
close to the large cities (e.g. Paris, Lyon),
where the fruits could be easily sold. Selec-
tions from open-pollinated seedlings in the
16th century gave rise to cultivars adapted to
various environments, such as ‘Hasty Ferri-
ere’ in the Paris area. In the orchard of the
‘Sun King’ in Versailles, there were at least 40
different cultivars, whose names often evoked
the female charms: ‘Nipple of Venus’, ‘Beauti-
ful of Chevreuse’, ‘Large Mignonne’, etc. By
the 18th century peaches were mainly white-
fleshed, the yellow ones being called either
‘alberge’ or ‘peach-apricot’. An inventory with
descriptions of peach cultivars was carried
out in the 19th century. In 1856, the railway
connecting Paris, Lyon and Marseilles allowed
the rise of new areas of peach production in
southern France, in particular in Ardèche and
Drôme provinces. At the turn of the century,
Jouin (1913) catalogued over 300 peaches and
nectarines based on leaf, fruit and flower
characteristics. In the 1930s the intensification
of peach production was made possible by
the ‘invasion’ of the US cultivars coming from
controlled-hybridization programmes. Fol-
lowing the Second World War the peach
industry was strongly reshaped and nowa-
days the major areas of production are located
in southern France, with Rhône-Alpes,
Languedoc-Roussillon, Midi-Pyrénées and
Provence-Alpes-Côte d’Azur being the most
important provinces.
Modern peach breeding in France may
be traced back to the early 1960s (R. Monet,
France, 2004, personal communication). After
an earlier release of a white peach series from
Hugard and Saunier (known as ‘Genadix’),
R. Monet at the Institut National pour la
Réchèrche Agronomique (INRA) started a
formal programme to address the lack of
knowledge of heritability of important traits
(Monet, 1967, 1977, 1983; Monet and Bastard,
1982). In addition to traditional crossing and
selection, he made inbred lines, pursuing the
obtaining of ‘hybrid’ progenies as in maize or
other vegetable crops. Although he reached
the sixth inbred generation, he never attained
his final goal. He also described some novel
traits, among them weeping tree habit and
the resistances to green aphid and leaf curl
(Massonie et al., 1982; Monet, 1984; Monet
et al., 1988). Some of his selections may still be
useful as breeding material, particularly for
the weeping habit and aphid resistance. He
also released a flat peach ‘Platina’ as well as a
flat nectarine, ‘Mesembrine’.
At the end of the 1990s the programme
was moved from Bordeaux to the station in
Avignon. The present programme aims to
breed for improved fruit quality as well as
resistance to powdery mildew, leaf curl, green
peach aphid, Plum pox virus (PPV) and nema-
todes (Pascal et al., 1998; Moing et al., 2003).
This programme is focused on the introgres-
sion of resistance genes by repeated back-
crosses to cultivars of commercial value
(Sauge et al., 1998). The breeding programme
is based on ‘Malo Konare’ and ‘Pamirskij 5’ as
resistance sources to powdery mildew, and
on ‘Weeping Flower Peach’ and ‘Rubira®’ for
green peach aphid resistance. Both traits are
controlled by major dominant genes (Pascal
et al., 2002b). Early selection procedures in
order to reduce selection cycles and field
evaluations against powdery mildew were
carried out (Dirlewanger et al., 1996). Intro-
gression of quantitative resistance from the
wild related species P. davidiana (clone P1908),
as a donor of resistance to several pests and
diseases including PPV, has been explored
and could lead to the acquisition of durable
resistances (Pascal et al., 2002a). Qualitative
trait loci (QTLs) involving both resistance and
fruit quality traits were identified, with the
aim to develop marker-assisted selection as
the first step of the selection process (Fou-
longne et al., 2003). One of the major problems
of the interspecific crossing is the poor fruit
characteristics of the wild parent, since even
two generations of backcrossing were not
enough to regain the commercial appearance
of the fruits (Kervella et al., 2002; Quilot et al.,
2004b). To overcome this constraint, a new
approach was undertaken in order to improve
the fruit quality, to understand the genetic
variability observed in the interspecific prog-
eny by combining QTL analysis and ecophys-
iological models (Quilot et al., 2004a). Tree
growth habit is also under study, with the aim
to obtain easy-to-train trees with short and
thick branches, in order to assure large and
uniform size of fruits (Genard et al., 1994;
Kervella et al., 1994, 1998).
 Fresh Market Cultivar Development
Partnership with private breeders (Pascal
and Monteux-Caillet, 1998) led to the devel-
opment of white (‘Caprice’, ‘Surprise’ and
‘Elise’) and yellow (‘Conquise’) peaches. More
recently, several private enterprises have been
involved, e.g. Meynaud, Valla and especially
Maillard, whose introductions like ‘Big Bang®’
and ‘Sweetcap®’ are beginning to be widely
planted in France as well as Spain and Italy.
Some private growers are also involved in
breeding for new cultivars, e.g. Buffat, Dumont
and Monteux-Caillet are working with INRA.
‘Zephir®’ (‘Zephyr’ in the USA), ‘Emeraude®’,
‘Jade®’ and ‘Topaze®’ are the most widely
known introductions from this cooperative
programme, and make up about 5% of the
white nectarine orchards in France. Also,
some white-fleshed peaches like ‘Opale®’ and
‘Melina®’ are frequently planted in all dis-
tricts of peach cultivation. The main goal of
these private breeding programmes is to
develop new outstanding cultivars to meet
the needs of the modern industry (size, firm-
ness, productivity), of good taste (high sugar
content, diversity of flavours), with some
emphasis on unusual traits like flat-shaped
fruits (particularly for white peaches) and
‘red flesh’ nectarines (Zambujo, 2004). Today
French cultivars represent about 30% of the
peach industry in France. New INRA releases
are nationally distributed by the association
of nurseries ‘CEP Innovations’.
Greece
Although the peach reached Greece some
centuries before the Christian era, it was
propagated by seed until the beginning of the
20th century. Currently the peach industry is
based on foreign cultivars, mainly from the
USA. A breeding programme aimed at improv-
ing the field resistance to PPV by crossing
virus-susceptible, high-quality local acces-
sions to some field-tolerant US cultivars, e.g.
‘Elegant Lady’, ‘July Lady’ and ‘May Crest’,
was begun and then stopped because of lim-
ited results.
A programme was recently started at the
Naoussa Pomology Institute in order to obtain
both scion cultivars and rootstocks well adapted
to the Greek environments (occurrence of
163
spring frost, calcareous soils, etc.). The main
goals of the programme are disease resistance
(PPV, brown rot, Pseudomonas spp.) of non-
melting peaches, both yellow and white, using
local (white-fleshed ‘Flagar’), US (‘Andross’,
‘Fortuna’, etc.) or Japanese (white, low-acid
peach ‘Akatsuki’) parents. More recently, a
clonal selection project for old local cultivars
has been started.
Hungary
Hungary is characterized by a strong conti-
nental climate, but there are opportunities for
small-scale cultivation in some regions: close
to Budapest, in southern Hungary (Szeged-
Szatymaz) and in the milder region of the
large Balaton Lake. Although there is no for-
mal breeding work in the country, amateurs
and scholars have collected local germplasm
or made occasional crosses. Worthy of men-
tion are the works of P. Tóth (active from 1930
to 1965) and I. Tamássy (1950s–1960s). Among
the more notable Hungarian cultivars are
‘Mariska’ (white flesh), ‘Piroska’ (white flesh),
‘Vérbarack’ (red flesh), all of local origin; and
‘Aranycsillag’ (yellow flesh) and ‘Remény’
(flat fruit, white flesh) from controlled crosses.
Overall, the peach industry is still based on
old US cultivars, e.g. ‘Redhaven’, ‘Suncrest’,
‘Champion’, etc. (Szabó, 2007).
Italy
Italy, with a long history of peach cultivation,
is one of the first peach industries in the
world. Peach germplasm importation from
Asia dates back to the first centuries BC accord-
ing to the earliest-known written records from
Latin scholars. Because of its very early estab-
lishment in Italy, and given its easy propaga-
tion by seed, peach germplasm developed a
wealth of variability in terms of fruit type and
season of ripening. With the predominance of
self-pollination (Hesse, 1975; Fogle, 1977),
repeated seed propagation led to the establish-
ment of rather uniform populations used as
local cultivars and only the most outstanding
were propagated by grafting. Local preferences
164 W.R. Okie et al.
and natural pressure were crucial in guiding
the selection of non-melting, yellow-fleshed
types in central and southern Italy, and of
freestone, white-fleshed types at northern
latitudes, that were more adapted to the con-
tinental-like climate (Gallesio, 2003). This pat-
tern was the same for many centuries and as
recently as the mid-1960s in northern Italy
about 50% of the commercial production was
of white-fleshed, local cultivars (Branzanti
and Sansavini, 1965). They were appreciated
by growers for their hardiness and yield reli-
ability and by consumers for their distinctive
fruit flavour and fragrance. Market pressures
suddenly reduced white peach production to
6% at the beginning of the 1980s in Emilia-
Romagna, the major peach-growing district
in northern Italy (Bassi et al., 1980).
Nectarines were first reported in Italy
between the 14th and 15th century, possibly
imported into Spain and southern Italy by
the Arabs. They were called ‘pesche-noci’
(walnut-peach) for their small, greenish-
skinned fruit and, owing to this very poor
fruit appearance, were regarded only as a
curiosity (Gallesio, 2003) until the massive
introduction of improved cultivars from
California in the mid-1970s. The first com-
mercial orchards were started around the end
of the 19th century (Zago, 1923; Ghinassi,
1927; Rosi, 1965; Baldini, 2001) using local
germplasm, mainly white-fleshed cultivars.
As the first yellow-fleshed cultivars were
made available from the USA, they gradually
replaced the local types.
Modern breeding programmes began in
the late 1920s, with several amateurs as well
as scientists pioneering the work: C. Cappucci,
P. Martinis, A. Pieri, A. Pirovano and A.
Ragionieri, among others. It was A. Morettini
at the University of Florence who started a
true large-scale breeding programme, lasting
for over 30 years, until the 1970s (Fideghelli
and Sartori, 1998). Neglecting the Italian ger-
mplasm and resorting only to introductions
from the USA for his breeding stock, particu-
larly ‘J.H. Hale’ (the ideotype at that time),
Morettini developed about 20 cultivars
(Pirovano, 1953; Morettini, 1961). They were both
white- and yellow-fleshed, standard fresh
market peaches, covering a ripening season
from mid-June to August, a rather large span
for that time. Other scientists in Rome (Piro-
vano, 1936) and Bologna (Capucci, 1950)
developed mainly white peaches from local
germplasm. None of these early introductions
have commercial importance at present.
Near the end of the 1960s, three public
programmes were started almost simultane-
ously at the Institute of Fruitculture of the
Ministry of Agriculture (operated in Rome
and Forlì, northern Italy) and at the universi-
ties of Bologna and Florence. Mostly these
programmes are aimed at the introduction of
cultivars better adapted to the Italian (north-
ern) environments with improved hardiness
and yield reliability, since most of the com-
mercial cultivars imported from California
showed poor crop reliability. However, the
booming of the peach industry in southern
districts has driven new breeding for lower-
chilling requirements.
Much effort is being devoted to pest and
disease resistance, particularly to brown rot
(Monilinia spp.), leaf curl (T. deformans (Berk.)
Tul.), powdery mildew (P. pannosa (Wallr.:Fr.)
Braun & Takamatsu) and green aphids (Myzus
persicae Sulzer). For brown rot, promising
selections showing good field tolerance are
under advanced evaluation in the Bologna
and Forlì programmes. Parents for resistance
come from foreign germplasm, e.g. ‘Con-
tender’ peach from the eastern USA (Liverani
et al., 1997; Bassi et al., 1998b). For powdery
mildew, a disease affecting all of the culti-
vated accessions (Roselli and Bellini, 1976),
foreign germplasm such as ‘Oro A’ (a seed-
ling of ‘Diamante’, a non-melting Brazilian
cultivar; Rodriguez and Sherman, 1990) is
being used. Although this trait seems quanti-
tatively inherited, there are reports of major
dominant genes (D’bov, 1974, 1975) and it is
possible to select for a good level of resistance
in the F1 populations (Liverani et al., 2003).
For leaf curl, an interesting hypersensitivity
resistance has been found in Italian local
white peaches, e.g. ‘Cesarini’ and chance
seedling ‘Exoascus RM’ (Bellini et al., 1998;
Liverani et al., 2003). Promising selections are
under evaluation.
Fruit quality improvement has been
addressed by different approaches. Given the
rather good level of size and external appear-
ance, efforts are being concentrated towards
 Fresh Market Cultivar Development
inner components, e.g. flesh texture and flavour
(Bassi et al., 1998a; Liverani and D’Alessandro,
1999; Giovannini et al., 2004). Traits like low-
acid taste, stony-hard flesh, canning nectar-
ines (anthocyanin-free fruit) and increased
ascorbic acid are also of interest (Yoshida,
1970, 1976; Bellini, 1994; Bellini et al., 1996;
Liverani et al., 2002).
Genetic modification of tree architecture
could lead to the development of an easy-to-
train tree. The topic has been addressed since
the end of the 1970s; so far, commercial qual-
ity genotypes have been achieved for the
dwarf, columnar and upright phenotypes
(Fideghelli et al., 1979, 1992; Bassi et al., 1994;
Liverani et al., 2004). The weeping type
(Monet et al., 1988) has also been investigated
(Bassi and Rizzo, 2000).
In the large Italian peach collections
(about 2400 accessions), more than 20% are of
Italian origin (Bellini et al., 1990). Neverthe-
less, the pedigrees of the more than 200 Italian
cultivars released in the last 25 years show a
predominance of US cultivars, particularly
from California. The most interesting recent
results exploiting the local white-fleshed peach
germplasm are ‘Aliblanca’ and ‘Alirosada’
(from Forlì), ‘Maria Bianca’ (from Florence)
and ‘Rubia’ and ‘Rubisco’ (from Bologna).
Although it was terminated at the end of the
1980s, the programme of the former Institute
of Fruitculture in Verona (north-eastern Italy)
had the foresight to exploit the locally grown
white peaches with the aim of improving
their firmness and shelf-life while keeping
their hardiness and outstanding flavour. ‘Ata-
lanta’ and ‘Ione’ were the best achievements
from this notable breeding programme.
More recently, a programme was started
at the University of Palermo in Sicily, with the
goal to introduce cultivars with different
chilling requirements adapted to the diverse
environments of the island. Desired adapta-
tions include either medium- to low-chill at
sea level or medium- to high-chill in the inner
valleys and hills. Desired traits are late and
very late (October) ripening season, non-
melting flesh for fresh market with improved
flavour (over 15°Brix) and shelf-life, and
altered growth habit (either columnar or
brachytic dwarf). Parents are chosen from the
rich Sicilian germplasm (Marchese et al., 2005)
165
for the late-ripening and non-melting, high-
quality fruits (e.g. ‘Imera’, ‘Xirbi’, ‘Tudia’,
‘Tardiva di Leonforte’, etc.) and from Florida
cultivars for the low-chilling requirements
(‘Flordastar’, ‘Flordacrest’, etc.).
Private programmes
Several amateurs were actively involved in
developing new cultivars at the onset of the
modern Italian peach industry, often starting
with local germplasm crossed to introduc-
tions from the USA (Branzanti and Sansavini,
1965). Almost all of this early work is lost, with
a few exceptions. The white-fleshed peach ‘Iris
Rosso’, bred in the early 1950s by amateur
P. Martinis, is still grown today.
The following active private programmes
are noteworthy: Bubani, CIV nurseries, Min-
guzzi, Morsiani, Montanari and Ossani. They
are all located in northern Italy (Emilia-
Romagna region). Their goals are strictly
commercial and based on the exploitation of
US cultivars, particularly nectarines, either
via open and self-pollination or crossing with
other commercial introductions, e.g. ‘Elegant
Lady’, ‘Fantasia’, ‘Big Top’, etc. Ossani (1987)
first introduced several white-fleshed nectar-
ines starting from ‘Snow Queen’ and ‘Red-
gold’, some of them being very popular today
(‘Caldesi’ and ‘Silver’ series). The private
breeders have contributed more than 50% of
the Italian peach introductions in the last 25
years (Fideghelli and Sartori, 1998). Finally,
credit should be given to (Stark) ‘Redgold’: it
is not only still the most widely grown nectar-
ine in Italy, but also the most popular parent
used in nectarine breeding programmes in
Italy. Among the most noteworthy of its off-
spring are ‘Ambra’, ‘Caldesi’ white-flesh series,
‘Orion’, ‘Maria Aurelia’ and ‘Venus’ among
the nectarines, and ‘Rome Star’ among the
peaches. Leading cultivars in northern Italy
are shown in Table 6.9.
Poland
A limited breeding programme was estab-
lished at the Pomology Research Institute at
Skierniewice in the mid-1960s, aimed at the
introduction of cold-resistant peaches for the
166 W.R. Okie et al.

Table 6.9. The most important peach and nectarine cultivars in Italian orchards from a survey
conducted in main northern Italy peach districts, representing around 60% of the regional production.
Percentages are relative to the total number of trees in that age bracket (column). Cultivars from Italian
programmes are in italics. (Data from CSO, Ferrara, Italy.)

>4 years 1–3 years >4 years 1–3 years

Peaches (%) (%) Nectarines (%) (%)

‘Springbelle’a 15 8 ‘Stark Redgold’ 14 8

‘Fayette’ 9 – ‘Venus’ 9 3
‘Elegant Lady’ 7 – ‘Big Top’ 9 10
‘Springcrest’ 6 2 ‘Sweet Lady’ 5 9
‘Royal Glory’ 6 10 ‘Sweet Red’ 5 8
‘Redhaven’ 4 5 ‘Maria Carla’ 5 7
‘Royal Gem’ 4 4 ‘Nectaross’ 4 2
‘Glohaven’ 4 2 ‘Maria Aurelia’ 4 2
‘Rich Lady’ 4 8 ‘Maria Laura’ 4 –
‘May Crest’ 4 – ‘Caldesi 2000’c 3 –
‘Flavorcrest’ 3 – ‘Supercrimson Gold’ 3 –
‘Maria Marta’ 3 7 ‘Ambra’ 3 4
‘Suncrest’ 3 4 ‘Guerriera’ 3 –
‘Sinphonie’ 1 8 ‘Amiga’ – 3
‘Rosa del West’b,c 1 4 ‘Laura’ 2 3
‘Kaweah’ – 4 ‘Lady Erica’ – 3
‘Rome Star’ 1 4 ‘Morsiani 60’ – 3
aBud sport from ‘Springcrest’.
bChance seedling.
cWhite-fleshed.

Polish continental environment (Jakubowski,
1998). US cultivars were crossed to Chinese
germplasm (‘Winter Peach’, ‘Manchurian
Peach’) from north-west China, known for its
cold hardiness. The programme was extended
to the F2 stage and two cultivars were released
from the resulting progeny, showing a tree
hardiness similar (‘Inka’) or superior (‘Iskra’)
to ‘Redhaven’. The programme was then dis-
continued. Several of their Chinese accessions
were later imported into Canada (by R.E.C.
Layne) and later into the USA for use as cold-
hardy rootstocks, including ‘Tzim Pee Tao’,
‘Chui Lum Tao’, ‘Hui Hun Tao’ and ‘Siberian C’.
Romania
Peach breeding in Romania dates back to the
end of the 1950s (Ivascu and Balan, 1998).
The breeding stock has been developed
from foreign cultivars, mainly from the USA
(‘J.H. Hale’, ‘Elberta’ and others), Harrow, Can-
ada (for cold tolerance), from western European
countries (France, Italy) and, lately, from
China. The first wave of breeding effort was
aimed at obtaining cultivars adapted to the
harsh continental environment of the country
(fluctuating temperatures in mid-winter,
dropping as low as −35°C), particularly for
the late season, owing to the sporadic crop-
ping of the early-ripening cultivars due to the
cold temperatures in spring. After 1985, more
attention was paid to disease resistance (brown
rot, leaf curl, powdery mildew, stem canker
and PPV) owing to the heavy rains common
in the peach-growing areas and the scarcity of
effective chemicals for spray programmes
because of the limited financial resources of
the country (Ivascu, 2002). After 1990, more
diversified goals were added, concerning fruit
types and quality (non-melting for canning
and nectarines), extension of the ripening
season and tree architecture (particularly for
the brachytic dwarf) (Stanica et al., 2002a,b).
 Fresh Market Cultivar Development
Progress has been made at the Research Sta-
tions for Fruit Tree Growing at Baneasa and
Constanza in breeding for cold resistance
(‘Splendid’, ‘Superba de Toamna’, ‘Triumf’ and
‘Victoria’ peaches), improved nectarines (‘Cora’,
‘Delta’, ‘Mihaela’, ‘Romamer 2’ and ‘Tina’),
dwarf tree (‘Cecilia’ and ‘Puiu’ peaches,
‘Livia’ and ‘Melania’ nectarines), ornamental
(‘Dan’ nectarine and ‘Paul’ peach) and disease
resistance (‘Victoria’ peach, highly resistant
to both powdery mildew and stem canker;
‘Flacara’ peach, resistant to powdery mildew;
several selections tolerant to leaf curl, brown
rot or Pseudomonas spp.). Presently, about
two-thirds of the most widely grown culti-
vars are releases from the Romanian breeding
programmes.
Serbia
Peach breeding in Serbia (former Yugoslavia)
started in the early 1950s at the Fruit Research
Institute in Čačak (Ogasanovič, 1998), fol-
lowed by programmes at Beograd and Novi
Sad. Breeders began with US germplasm in
order to obtain cultivars adapted to the coun-
try’s continental environment. More recently,
cultivars from European countries, mainly
Italy, were added as parents. Several new cul-
tivars have been released, all selected from
progenies derived from foreign germplasm:
‘Maja’, ‘Vesna’ and ‘Gročanka’ (all from self-
pollination of ‘Glohaven’), as well as ‘Dora’,
‘Čačak’, ‘Goca’ and ‘Julija’, among others.
One of the peculiarities of the region is
the wealth of peach landraces called ‘vineyard
peaches’, traditionally propagated by seed,
and also the most common source of peach
rootstocks in the country (Vujanic′-Varga and
Ognjanov, 1992). These peaches show a wide
range of fruit types and some valuable horti-
cultural traits (cold resistance, drought toler-
ance, disease resistance e.g. to powdery
mildew and leaf curl), but so far have been
scarcely used in breeding programmes.
Spain
Spain is known worldwide for its wealth of
local germplasm, particularly of non-melting,
167
yellow-fleshed local accessions. In the late
Middle Ages they were introduced into Cen-
tral and South America, where they became
landraces (Badenes et al., 1998b). Non-melting
flesh peaches are the most valuable in the
Spanish fresh market and much work has
been done on evaluation and clonal selection
of the best types (Badenes et al., 1998a). At
present, there are several ongoing breeding
programmes, three in public institutions (since
1993, at Saragoza and Valencia) and at least
seven in private enterprises (M.L. Badenes,
Spain, 2004, personal communication). Other
than specifically improving yellow clingstone
peaches, efforts are aimed at improving fruit
quality, disease resistance and broadening the
ripening season. A minor goal is to improve
flat-shaped peaches, a favourite fruit in some
markets. Some of the private programmes are
focused on improving very early-ripening,
melting-flesh nectarines and peaches, best
suited for the southern regions of the country,
often resorting to foreign breeding stocks in
agreement with US breeders.
Ukraine and former Soviet states
Ukraine has a climate favourable for peach
production, more so than most of the former
Soviet Union, which is too cold. Ivan N. Ryabov
at the Nikita Botanical Garden near Yalta began
the programme over 50 years ago by collecting
a great deal of peach germplasm in Central
Asia and the Trans-Caucasus. Over the years
dozens of cultivars of peaches and nectarines
have been released that are more disease-re-
sistant (particularly powdery mildew and
leaf curl), cold-hardy and productive in their
climate (Yezhov et al., 2005). Important peaches
include ‘Krymsky Feyerverk’, ‘Sagdiets’,
‘Startovy’, ‘Dostoyny’, ‘Otlichnik’ and ‘Posol
Mira’. Top nectarine releases include ‘Ametist’,
‘Evpatoriysky’, ‘Ishunsky’ and ‘Rubinovy 8’.
The Uzbek Research Institute of Plant
Industry, located in Tashkent Province of the
Republic of Uzbekistan, was established in
1924 by the well-known Russian plant explorer
and breeder Nikolai I. Vavilov. A modest peach
breeding programme there released these
adapted fresh market cultivars: peaches ‘Rannii
168 W.R. Okie et al.
Belii VIRa’, ‘Gulnoz’, ‘Zolotoi Ubilei’, ‘Chim-
gan’ and ‘Lyuchak’; and nectarine ‘Uchkun’
(Mavlyanova et al., 2005). The latter two peaches
have been used for drying and canning as well.
Breeding in Latvia traces back to before
1950 at the Horticultural Plant Breeding Exper-
imental Station in Dobele and 1938 at the
Botanical Gardens of the University of Latvia
(Kaufmane and Lacis, 2004). To be grown suc-
cessfully, peach cultivars need to be extremely
cold-hardy and have good spring bud hardi-
ness. Releases include ‘Ziemelu persiks’, ‘Zelda’,
‘Rita’, ‘Maira’ and ‘Viktors’. All are white-
fleshed except ‘Rita’; the latter two were
released in 2004.
6.6 Pacific and African Programmes
China
Although China is the ancestral home of the
peach, and the introduction of ‘Chinese Cling’
led to the development of improved peaches
in the USA, breeding programmes are rela-
tively new there. From the late 1950s until the
1990s there were canning cling programmes
in several places. Fresh market breeding
flourished in the 1970s and 1980s but the 20 or
so original programmes have been reduced to
only six currently, mostly developing low-
acid, white-fleshed freestone peaches (L. Wang,
Zhengzhou, 2007, personal communication).
In general, the programmes have intercrossed
Chinese and Japanese cultivars with US culti-
vars. There are no traditional nectarine cultivars
in China and available Japanese germplasm
was relatively poor in quality, so mostly US
cultivars have been used for breeding.
Breeding at Dalian Institute of Agricultural
Sciences began with yellow canning clings, and
releases ‘Fenghuang’ and ‘Lianhuang’ formed
the basis of the Chinese canning industry. More
recently focus has been on nectarines as well
as late-ripening, freestone peaches. Beijing
Academy of Agriculture and Forest Sciences
has one of the older programmes, which began
in the 1960s. ‘Qingfeng’ and ‘Maixiang’ are
important releases. Flat peach cultivars include
‘Ruipan 1’ and ‘Ruipan 4’. Current efforts are
directed at nectarines and late-ripening flat
peaches. At the Beijing Forest Institute, there
is a programme to develop low-chilling orna-
mental peaches. The programme at the Depart-
ment of Horticulture, Jiangshu Academy of
Agricultural Sciences in Nanjing, released the
prominent early peach ‘Yuhualu’ in 1972 as
well as other early peaches. Current efforts
are aimed at late-ripening, freestone peaches
with resistance to fungal gummosis. Nectarines
tolerant of rainy climates are also being devel-
oped. Zhengzhou Fruit Institute in Henan is
developing low- to medium-chill peaches and
nectarines, as well as flat nectarines. It also
maintains a peach germplasm repository and
has an interest in ornamental peaches. The
Department of Horticulture of Shanghai
Academy of Agricultural Sciences is develop-
ing early-ripening peaches and nectarines
primarily; releases include ‘Chunlei’ and
‘Chunhua’ (Li, 1984; Wang and Lu, 1992). For
more information on China, see Chapter 2.
Japan
Many of the peaches grown in Japan origi-
nated as grower selections and mutations of
traditional cultivars like ‘Hakuto’ and
‘Hakuho’ (Kozaki et al., 1996). ‘Hakuto’ is
apparently descended from ‘Shanhai Suimit-
suto’ (Yamamoto et al., 2003), which is thought
to be synonymous with ‘Chinese Cling’. Con-
sumers prefer high-quality, low-acid fruit
with red blush. Nearly all peaches currently
grown are white-fleshed clings, but recently
yellow-fleshed peaches, such as ‘Ogonto’ and
‘Golden Peach’, have been increasingly grown
for fresh consumption. Early-season produc-
tion once declined due to problems produc-
ing consistently high-quality fruit, but the
market is now recovering because of new
early cultivars with relatively high quality
(such as ‘Hikawa Hakuho’ from Yamanashi,
and ‘Chiyohime’). The most important breed-
ing programme has been carried on by the
National Institute of Fruit Tree Science in Tsu-
kuba. Under the direction of M. Yoshida and
more recently M. Yamaguchi, it has released a
series of peaches including ‘Chiyohime’,
‘Akatsuki’, ‘Natsuotome’, ‘Yuzora’ and ‘Aki-
zora’. Local research stations located in the
 Fresh Market Cultivar Development
major production area of peach are also active
in breeding, including Yamanashi (released
‘Yumeshizuku’, a very large peach), Nagano
(released ‘Natsuki’), Fukushima (released
‘Fukuotome’ and ‘Hatsuotome’, very early
peaches) and Okayama.
Nectarine is not very popular because of
the high acidity of available cultivars and the
tendency for fruit cracking, but that may
change with the release of low-acid nectarine
cultivars from the breeding programme at the
Yamanashi Prefectural Fruit Tree Experimen-
tal Station. ‘Sweet Nectarine Reimei’ and
‘Sweet Nectarine Reiou’ are yellow-fleshed,
while ‘Sweet Nectarine Shoku’ and ‘Sweet
Nectarine Shogyoku’ are white-fleshed. Orna-
mental peaches have been traditional favou-
rites in Japan and many flower types and tree
growth habits are available.
Korea
Peach culture in Korea dates back nearly 2000
years, but modern commercial production
began 100 years ago with the introduction of
better Japanese cultivars by scientists at what
is now the National Horticultural Research
Institute in Suwon. Although minimal breeding
started in 1957, the emphasis was increased in
the mid-1980s. Goals are to develop firmer,
white-fleshed, low-acid peaches that retain
the high quality consumers prefer, but also
have resistance to bacterial spot and other
diseases. The most successful releases have
been ‘Cheonhong’ nectarine and ‘Yumyeong’,
a large, white, stony-hard cling peach con-
sumed fresh (Chung et al., 1998). ‘Yumyeong’,
a selection from Japanese parentage, has been
used as a parent in other countries such as
Italy and New Zealand because of its size and
firmness.
New Zealand
Peach breeding by HortResearch in Havelock
North, New Zealand began as an effort to
improve canning clings. In the mid-1980s
work was expanded to develop fruit suitable
for export, particularly for the Asian market,
169
which prefers low-acid, white-fleshed peaches.
Cultivars must be adapted to the maritime
climate with disease resistance and frost tol-
erance, along with having high-quality, ship-
pable fruit. P.G. Glucina and currently M.T.
Malone have released ‘Coconut Ice’ and ‘Scar-
let O’Hara’ derived from ‘Yumyeong’ from
Korea as well as ‘Havelock Pearl’ derived from
‘Hakuto’ from Japan. Breeding is also under
way to develop adapted low-acid, white
nectarines.
South Africa
Early efforts in peach breeding in South Africa
were for canning clingstones and lower-chill
peaches. In recent years efforts have been
made to develop highly blushed, adapted,
yellow-fleshed peaches and nectarines that
will retain good quality after several weeks in
transit. An Agricultural Research Council
Infruitec–Nietvoorbij early dessert nectarine,
‘Alpine’ (released in 1997), has performed
exceptionally well on European markets.
Numerous other nectarine (‘Nectar’, ‘Royal
Gem’, ‘Sunburst’, ‘Crimson Giant’) and peach
(‘Excellence’, ‘Witzenberg’) cultivars have
been named and other selections are still
under evaluation for commercial use in the
export market. Most are low-chilling except
for ‘Royal Gem’, which is medium-chill. For
more information see Chapter 5.
6.7 Conclusions
Great progress has been made by peach
breeders in the last century. Many modern
cultivars are large-fruited and highly produc-
tive, with extensive red skin colour, little
pubescence, round shape, and very firm,
slow-softening, yellow or white flesh. Unfor-
tunately some of these gains have come at the
expense of eating quality, for both genetic and
cultural reasons. As a result, most programmes,
and particularly the private breeders, are now
putting increased emphasis on this aspect. In
more difficult climates, meaning outside Cali-
fornia, the process has been slower to incor-
porate these advances into peaches that also
170 W.R. Okie et al.
have tolerance to problems associated with
cold, rain and disease. In some cases indus-
tries have declined faster than breeders were
able to develop suitable cultivars, thus taking
the breeding programmes out as well.
The trend towards patenting most new
cultivars is changing the industries as well as
the breeding programmes. Traditionally, pub-
lic breeders shared and cross-tested breeding
stock and advanced selections. Now that most
public breeders are patenting new releases,
less testing is done across sites in order to
protect the selections. At the time of release
growers may have less information available
on which to base a decision to plant the culti-
var. Private breeders are able to grow larger
numbers of seedlings than public breeders and
hence make bigger gains in improvement.
Over time these improvements tend to be
incorporated into other programmes. In some
industries public breeders are focusing on ger-
mplasm screening and enhancement for char-
acters such as disease resistance, while their
private counterparts focus on cultivar devel-
opment. In California, private breeders pro-
vide the bulk of freestone peach cultivars. This
has not occurred in the eastern USA with the
exception of the private programmes in Michi-
gan. The success of the private Michigan pro-
grammes may relate to a void in new cultivars
developed for the northern climates, but it is
not clear if they will be self-sustaining. It seems
unlikely that private programmes will be as
successful in the fragmented eastern indus-
tries, which have more challenging climatic
and disease situations relative to California.
The increase in breeding programmes
outside the USA is a welcome trend given the
decline of public breeding programmes in the
USA. It may reduce the overseas market for
privately bred US cultivars, but in the long
run will likely be of greater benefit to the local
industries. Few countries have a climate as
ideal as that of California for growing peaches
and nectarines. Locally selected cultivars can
References
Anon. (1997) The Brooks and Olmo Register of Fruit and Nut Varieties, 3rd edn. ASHS Press, Alexandria, Vir-
ginia.
be better adapted and reduce the risk of crop
loss in years of high disease pressure.
Acknowledgements
Thanks are due to the following scholars for
providing information related to their coun-
tries: A. Zhivondov (Bulgaria); R. Monet and
T. Pascal (France); C. Tsipouridis (Greece); Z.
Szabó (Hungary); G. Baroni, E. Bellini, T. Car-
uso, A. Liverani, A. Martinelli (CIV), V. Mazzotti
(CSO) and V. Ossani (Italy); A. Ivascu (Roma-
nia); V. Ognjanov (Serbia); M.L. Badenes (Spain);
L. Wang (China); and M. Yamaguchi (Japan).
A note about references
The American Pomological Society published
brief descriptive lists of peaches and other
fruit from 1920 to 1950. In 1942, R.M. Brooks
and H.P. Olmo began a Register of New Fruit
and Nut Varieties. The new register consisted
of many lists published in the Proceedings of
the American Society for Horticultural Science
from 1944 to 1970 and later in HortScience.
Periodically it was consolidated into book
form (Anon., 1997). The most comprehensive
single listing of cultivars is USDA Agricul-
tural Handbook 714, which has descriptions
of over 700 cultivars from the USA as well as
a similar number from other countries, plus
an index of 6000 names and synonyms (Okie,
1998). Unfortunately there is no single listing
of the newest US patented cultivars other
than the patents themselves, which can be
viewed at http://www.uspto.gov/patft/index.
html. Several excellent books with complete
descriptions and photographs have been pub-
lished in Europe, particularly France (most
recently by Hilaire and Giauque, 1994) and
Italy (most recently by Della Strada et al.,
1984, 1986; Conte et al., 1994).
 Fresh Market Cultivar Development 171

Badenes, M.L., Martínez-Calvo, J. and Llácer, G. (1998a) Analysis of peach germplasm from Spain. Acta
Horticulturae 465, 243–250.
Badenes, M.L., Werner, D.J., Martínez-Calvo, J., Lorente, M. and Llácer, G. (1998b) An overview of the peach
industry of Spain. Fruit Varieties Journal 52, 11–17.
Bailey, J.S. and French, A.P. (1949) The inheritance of certain fruit and foliage characters in the peach. Mas-
sachusetts Agricultural Experiment Station Bulletin 452.
Baldini, E. (2001) Luci ed ombre della peschicoltura romagnola. Rivista di Frutticoltura e di Ortofloricoltura
7/8, 87–93.
Bassi, D. and Rizzo, M. (2000) Peach breeding for growth habit. Acta Horticulturae 538, 411–414.
Bassi, D., Sansavini, S., Marangoni, B. and Bordini, R. (1980) Recupero delle pesche bianche: prove agrono-
miche comparative di vecchie cultivar e selezioni locali della Romagna. In: Proceedings of ‘XV Con-
vegno Peschicolo’. Chambers of Commerce of Forlì and Ravenna, Ravenna, Italy, pp. 153–180.
Bassi, D., Dima, A. and Scorza, R. (1994) Tree structure and pruning response of six peach growth forms.
Journal of the American Society for Horticultural Science 119, 378–382.
Bassi, D., Mignani, I. and Rizzo, M. (1998a) Calcium and pectin influence on peach flesh texture. Acta
Horticulturae 465, 433–438.
Bassi, D., Rizzo, M. and Cantoni, L. (1998b) Assaying brown rot (Monilinia laxa Aderh. et Ruhl. (Honey))
susceptibility in peach cultivars and progeny. Acta Horticulturae 465, 715–721.
Bellini, E. (1994) Maria Dolce: nuova cultivar di nettarina a maturazione tardiva pregevole per consistenza e
sapore della polpa. In: Proceedings of ‘2nd Giornate Scientifiche SOI’. Società Orticola Italiana, Flor-
ence, Italy, pp. 171–172.
Bellini, E., Giannelli, G., Giordani, E. and Picardi, E. (1990) Reperimento e difesa delle risorse genetiche del
pesco in Italia. L’Informatore Agrario 9, 181–191.
Bellini, E., Giannelli, G., Giordani, E. and Sabbatini, I. (1996) Peach genetic improvement: breeding program
carried on at Florence to obtain canning nectarines. Acta Horticulturae 374, 21–32.
Bellini, E., Giordani, E., Nencetti, V. and Paffetti, D. (1998) Indagine della variabilità nell’espressione genica
in una progenie di pesco presunta resistente a Taphrina deformans (Berk.) Tuls. In: Proceedings of ‘4th
Giornate Scientifiche SOI’. Società Orticola Italiana, Sanremo, Italy, pp. 3–4.
Blake, M.A. and Connors, C.H. (1936) Early results of peach breeding in New Jersey. New Jersey Agricultural
Experiment Station Bulletin 599.
Blake, M.A. and Edgerton, L.J. (1946) Standards for classifying peach characters. New Jersey Agricultural
Experiment Station Bulletin 728.
Bradford, N. (1988) The Fred Anderson/Norman Bradford private nectarine breeding program. In: Childers,
N.F. and Sherman, W.B. (eds) The Peach. Horticultural Publications, Gainesville, Florida, pp. 106–108.
Branzanti, E.C. and Sansavini, S. (1965) Le cultivar di pesco. Edizioni Agricole, Bologna, Italy.
Butterfield, H.M. (1938) History of deciduous fruits in California. Pioneer days in California’s peach industry.
The Blue Anchor 15, 14–18.
Capucci, C. (1950) Le selezioni di pesco ‘C. Capucci’. Rivista della Ortoflorofrutticoltura Italiana 75, 167–176.
Childers, N.F. and Sherman, W.B. (1988) John Howard Weinberger. In: Childers, N.F. and Sherman, W.B. (eds)
The Peach. Horticultural Publications, Gainesville, Florida, p. II.
Chung, K.H., Kang, S.J., Jun, J.H. and Okie, W.R. (1998) Stone fruit production and breeding in Korea. Fruit
Varieties Journal 52, 184–189.
Clark, J.R., Moore, J.N. and Perkins-Veazie, P. (2005) ‘White Rock’ and ‘White County’ peaches. HortScience
40, 1561–1565.
Conte, L., Della Strada, G., Fideghelli, C., Insero, O., Liverani, A., Moser, L. and Nicotra, A. (1994) Monogra-
fia di cultivar di pesco, nettarine, percoche. Istituto Sperimentale per la Frutticoltura, Rome.
CTFA (2003) California Tree Fruit Agreement 2002 Annual Report. California Tree Fruit Agreement, Sacra-
mento, California.
Cullinan, F.P. (1937) Improvement of stone fruits. In: Better Plants and Animals – II. 1937 Yearbook of Agricul-
ture. US Department of Agriculture, Washington, DC, pp. 665–748.
D’bov, S. (1974) Inheritance of resistance to powdery mildew in peach. I. Resistance of vegetative organs in
freestone varieties with pubescent fruit. Genetika i Selektsiya 7, 281–291.
D’bov, S. (1975) Inheritance of resistance to powdery mildew in peach. II. Resistance of vegetative organs in
the F1 from crosses between freestone and clingstone varieties with pubescent fruit. Genetika i Selektsiya
8, 267–271.
Della Strada, G., Fideghelli, C., Liverani, A., Monastra, F. and Rivalta, L. (1984) Monografia di cultivar di
pesco da consumo fresco, Vol. 1. Istituto Sperimentale per la Frutticoltura, Rome.
172 W.R. Okie et al.

Della Strada, G., Fideghelli, C., Insero, O., Liverani, A., Monastra, F. and Rivalta, L. (1986) Monografia di
cultivar di pesco da consumo fresco, Vol. 2. Istituto Sperimentale per la Frutticoltura, Rome.
Dirlewanger, E., Pascal, T., Zuger, C. and Kervella, J. (1996) Analysis of molecular markers associated with
powdery mildew resistance genes in peach (Prunus persica (L.) Batsch) × Prunus davidiana hybrids.
Theoretical and Applied Genetics 93, 909–919.
Fideghelli, C. and Sartori, A. (1998) Peach. In: Scarascia Mugnozza, G.T. and Pagnotta, M.A. (eds) Italian
Contribution to Plant Genetics and Breeding. University of Tuscia, Viterbo, Italy, pp. 643–649.
Fideghelli, C., Della Strada, G., Quarta, R. and Rosati, P. (1979) Genetic semi-dwarf peach selections. In:
Proceedings of the Symposium on ‘Tree Fruit Breeding’. Eucarpia, Angers, France, pp. 11–18.
Fideghelli, C., Della Strada, G. and Grassi, F. (1992) Calipso e Circe, due nuove cultivar nane di pesco. In:
Proceedings of ‘Giornate Scientifiche SOI’. Società Orticola Italiana, Florence, Italy, pp. 414–415.
Fogle, H.W. (1977) Self-pollination and its implications in peach improvement. Fruit Varieties Journal 31,
74–75.
Foulongne, M., Pascal, T., Pfeiffer, F. and Kervella, J. (2003) QTLs for powdery mildew resistance in peach × Prunus
davidiana crosses: consistency across generations and environments. Molecular Breeding 12, 33–50.
Gallesio, G. (2003) Il trattato del pesco di Giorgio Gallesio. In: Baldini, E. (ed.) Gli inediti trattati del pesco e del
ciliegio. Complementi scientifici della ‘Pomona Italiana’ di Giorgio Gallesio. Accademia dei Georgofili,
Florence, Italy, pp. 9–146.
Genard, M., Pages, L. and Kervella, J. (1994). Relationship between sylleptic branching and components of
parent shoot development in the peach tree. Annals of Botany 74, 465–470.
Ghinassi, P. (1927) I progressi dell’agricoltura romagnola nell’ultimo venticinquennio. L’Italia Agricola 12,
682–705.
Giovannini, D., Liverani, A., Merli, M. and Brandi, F. (2004) Strategie del miglioramento genetico per miglio-
rare alcuni aspetti della qualità nel pesco. Rivista di Frutticoltura e di Ortofloricoltura 7/8, 38–43.
Graham, C.J. (1999) ‘Harvester’ peach. Fruit Varieties Journal 53, 130–132.
Hansche, P.E. and Beres, W. (1989) Three brachytic peach cultivars: Valley Gem, Valley Red, and Valley Sun.
HortScience 24, 707–709.
Havis, L., Weinberger, J.H. and Hesse, C.O. (1947) Better peaches are coming. In: Stefferud, A. (ed.) Science
in Farming – The Yearbook of Agriculture 1943–1947. US Department of Agriculture, Washington, DC,
pp. 304–311.
Hedrick, U.P. (1917) The Peaches of New York. J.B. Lyon Company Printers, Albany, New York.
Hesse, C.O. (1975) Peaches. In: Janick, J. and Moore, J.N. (eds) Advances in Fruit Breeding. Purdue University
Press, West Lafayette, Indiana, pp. 285–335.
Hilaire, C. and Giauque, P. (1994) Pêche – Les Variétés & Leur Conduite. Centre Technique Interprofessionnel
des Fruits et Légumes, Paris.
Hoare, A.H. (1950) The peach in England. Journal of the Ministry of Agriculture 57, 27–29.
Iezzoni, A.F. (1987) The Redhaven peach. Fruit Varieties Journal 41, 50–52.
Ivascu, A. (2002) New trends of peach tree growing in Romania. Acta Horticulturae 592, 93–97.
Ivascu, A. and Balan, V. (1998) Peach breeding programme at the Research Station for Fruit Tree Growing
Baneasa, Romania. Acta Horticulturae 465, 129–136.
Jakubowski, T. (1998) Breeding of peach cultivars in Poland. Acta Horticulturae 465, 125–127.
Jouin, J. (1913) Classification des peches et nectarines. Imprimerie Paul Legendre, Lyon, France.
Kaufmane, E. and Lacis, G. (2004) Winter-hardy apricots and peaches with good fruit quality in Latvia. Journal
of Fruit and Ornamental Plant Research Special Edition 12, 321–329.
Kervella, J., Pages, L. and Genard, M. (1994) Genotypic differences in the length–diameter relationship of
branches of one-year-old peach and nectarine trees. Journal of the American Society for Horticultural
Science 119, 616–619.
Kervella, J., Pfeiffer, F., Pagès, L., Génard, M. and Serra, V. (1998) Taking into account relationships between
ecophysiological processes in the breeding for peach tree characteristics. Acta Horticulturae 465, 145–
154.
Kervella, J., Foulongne, M., Kraif, R., Pascal, T., Pfeiffer, F., Pradier, M., Genard, M., Quilot, B., Lescourret, F.,
Reick, M., Moing, A., Renaud, C., Etienne, C. and Rothan, C. (2002) Prunus davidiana and derived
progenies: a valuable material for fruit quality studies. Acta Horticulturae 592, 109–115.
Kessler, G.M. (1969) Stanley Johnston. Fruit Varieties and Horticultural Digest 23, 37–38.
Kozaki, I., Ueno, I., Tsuchiya, S. and Kajiura, I. (1996) The Fruit in Japan. Yokendo, Tokyo.
Lammerts, W.E. (1945) The breeding of ornamental edible peaches for mild climates. 1. Inheritance of tree
and flower characters. American Journal of Botany 32, 53–61.
 Fresh Market Cultivar Development 173

Layne, R.E.C. (1997) Peach and nectarine breeding in Canada: 1911 to 1995. Fruit Varieties Journal 51,
218–228.
Li, Z.-L. (1984) Peach germplasm and breeding in China. HortScience 19, 348–351.
Liverani, A. and D’Alessandro, D. (1999). La qualità gustative dei frutti nell’attività di miglioramento genetico
del pesco presso l’ISF di Forlì. Rivista di Frutticoltura e di Ortofloricoltura 2, 30–37.
Liverani, A., Calisesi, F. and Bendandi, C. (1997) Il miglioramento genetico del pesco per la resistenza a
Monilia. In: Proceedings of ‘XXII Convegno Peschicolo’. Società Orticola Italiana, Cesena, Italy, pp.
103–106.
Liverani, A., Giovannini, D. and Brandi, F. (2002) Increasing fruit quality of peaches and nectarines: the main
goals of ISF-FO (Italy). Acta Horticulturae 592, 507–514.
Liverani, A., Giovannini, D., Brandi, F. and Merli, M. (2003) Miglioramento genetico del pesco per la resist-
enza alle principali malattie crittogamiche. In: Proceedings of ‘Presentazione dei risultati ottenuti dal
progetto finalizzato Mipaf-Frutticoltura’, Vol. 1. MiPAF (Ministero delle Politiche Agricole e Forestali),
Cesena, Italy, pp. 33–35.
Liverani, A., Giovannini, D., Brandi, F. and Merli, M. (2004) Development of new peach varieties with colum-
nar and upright growth habit. Acta Horticulturae 663, 381–386.
Marchese, A., Tobutt, K.R. and Caruso, T. (2005) Molecular characterisation of Sicilian Prunus persica cultivars
using microsatellites. Journal of Horticultural Science and Biotechnology 80, 121–129.
Massonie, G., Maison, P., Monet, R. and Grasselly, C. (1982) Résistance au puceron vert du pêcher, Myzus
persicae Sulzer (Homoptera Aphididae) chez Prunus persica (L.) Batsch et d’autres espèces de Prunus.
Agronomie 2, 63–70.
Mavlyanova, R.F., Abdullaev, F.K., Khodjiev, P., Zaurov, D.E., Molnar, T.J., Goffreda, J.C., Orton, T.J. and Funk,
C.R. (2005) Plant genetic resources and scientific activities of the Uzbek Research Institute of Plant
Industry. HortScience 40, 10–14.
Moing, A., Pöessel, J.L., Svanella-Dumas, L., Loonis, M. and Kervella, J. (2003) Biochemical basis of low fruit
quality of Prunus davidiana, a pest and disease donor for peach breeding. Journal of the American Society
for Horticultural Science 128, 55–62.
Monet, R. (1967) Contribution à l’étude génètique du pêcher. Annales de l’Amélioration des Plantes 17, 5–11.
Monet, R. (1977) Amélioration du pêcher par voie sexuée, exposé d’une méthode. Annales de l’Amélioration
des Plantes 27, 223–234.
Monet, R. (1983) Le pecher, génétique et physiologie. Masson, Paris.
Monet, R. (1984) Heredity of the resistance to leaf curl (Taphrina deformans) and green aphid (Myzus persi-
cae) in the peach. Acta Horticulturae 173, 21–23.
Monet, R. and Bastard, Y. (1982) Estimation du coefficient de régression enfant/parent de quelques caractéres
du pecher dans le cas de familles issues d’autofécondations. Agronomie 2, 347–358.
Monet, R., Bastard, Y. and Gibault, B. (1988) Etude génétique du caractère ‘port pleureur’ chez le pêcher.
Agronomie 8, 127–132.
Morettini, A. (1961) Le nuove cultivar Morettini. Consiglio Nazionale delle Ricerche, Florence, Italy.
Mowry, J.B. (1960) Selecting peach varieties for new plantings. Transactions of the Illinois State Horticulture
Society and Illinois Fruit Council 94, 97–109.
Myers, S.C., Okie, W.R. and Lightner, G. (1989) The Elberta peach. Fruit Varieties Journal 43, 130–138.
Oberle, G.D. (1971) What’s new in peach varieties at VPI and the mid-Atlantic states. In: Proceedings of the
30th Annual Convention. National Peach Council, Atlanta, Georgia, pp. 116–119.
Ogasanovič, D. (1998) Dora, a medium late, good quality, peach cultivar. Acta Horticulturae 465, 193–196.
Okie, W.R. (1997) USDA stone fruit breeding in the southeastern United States. Fruit Varieties Journal 51,
211–217.
Okie, W.R. (1998) Handbook of Peach and Nectarine Varieties. US Department of Agriculture Handbook No.
714. US Government Printing Office, Washington, DC.
Okie, W.R. and Myers, S.C. (1991) Springcrest peach. Fruit Varieties Journal 45, 190–192.
Okie, W.R., Ramming, D.W. and Scorza, R. (1985) Peach, nectarine and other stone fruit breeding by the
USDA in the last two decades. HortScience 20, 633–641.
Ossani, V. (1987) Tre nuove nettarine bianche: Caldesi 2000, Caldesi 2010, Caldesi 2020. L’Informatore
Agrario 22, 52–56.
Pascal, T. and Monteux-Caillet, R. (1998) Peach breeding in France. Acta Horticulturae 465, 117–123.
Pascal, T., Kervella, J., Pfeiffer, F.G., Sauge, M.H. and Esmenjaud, D. (1998) Evaluation of the interspecific
progeny Prunus persica cv. Summergrand × Prunus davidiana for disease resistance and some agro-
nomic features. Acta Horticulturae 465, 185–191.
174 W.R. Okie et al.

Pascal, T., Pfeiffer, F. and Kervella, J. (2002a) Preliminary observations on the resistance to sharka in peach.
Acta Horticulturae 592, 699–706.
Pascal, T., Pfeiffer, F., Kervella, J., Lacroze, J.P. and Sauge, M. (2002b) Inheritance of green peach aphid resis-
tance in the peach cultivar ‘Rubira’. Plant Breeding 121, 459–461.
Pirovano, A. (1936) Miglioramento genetico del pesco. L’Italia Agricola 10, 711–722.
Pirovano, A. (1953) Le nuove pesche italiane. Istituto di Frutticoltura ed Elettrogenetica, Rome.
Quilot, B., Genard, M., Kervella, J. and Lescourret, F. (2004a) Analysis of genotypic variation in fruit flesh total
sugar content via an ecophysiological model applied to peach. Theoretical and Applied Genetics 109,
440–449.
Quilot, B., Wu, B.H., Kervella, J., Genard, M., Foulongne, M. and Moreau, K. (2004b) QTL analysis of quality
traits in an advanced backcross between Prunus persica cultivars and the wild relative species P. davidiana.
Theoretical and Applied Genetics 109, 884–897.
Ramming, D. (1988) California peach and nectarine breeding objectives, cultivars. In: Childers, N.F. and
Sherman, W.B. (eds) The Peach. Horticultural Publications, Gainesville, Florida, pp. 89–96.
Rivers, H.S. (1906) The cross-breeding of peaches and nectarines. In: Report of the Third International Confer-
ence on Genetics. Royal Horticultural Society, London, pp. 463–467.
Rivers, T. (1866) On raising peaches, nectarines, and other stone fruits from seed. In: Report of Proceedings of
International Horticultural Exhibition and Botanical Congress. Truscott, Son, & Simmons, London, pp.
148–155.
Rodriguez, J.A. and Sherman, W.B. (1990). ‘Oro A’ peach germplasm. HortScience 25, 128.
Roselli, G. and Bellini, E. (1976) Indagine sulla suscettibilità all’oidio Sphaerotheca pannosa [Vallr] Lev. in
una collezione di cultivar di pesco. Rivista della Ortoflorofrutticoltura Italiana 60, 40–55.
Rosi, M. (1965) Frutticoltura. In: Calzecchi-Onesti, A. and Marinucci, M. (eds) Enciclopedia agraria italiana,
Vol. 5. Ramo Editoriale degli Agricoltori, Rome, pp. 6–36.
Sauge, M.H., Kervella, J. and Rahbe, Y. (1998). Probing behaviour of the green peach aphid Myzus persicae
on resistant Prunus genotypes. Entomologia Experimentalis et Applicata 89, 223–232.
Stanica, F., Cepoiu, N. and Dumitru, L.M. (2002a) New dwarf peach and nectarine tree varieties registered in
2000 by the Fruit Research Station Constanta, Romania. Acta Horticulturae 592, 161–163.
Stanica, F., Cepoiu, N., Dumitru, L.M. and Caretu, G. (2002b) New ornamental dwarf peach tree varieties in
Romania. Acta Horticulturae 592, 165–167.
Stark, P. Jr (1974) A tribute to Grant Merrill. Fruit Varieties Journal 28, 12–14.
Szabó, Z. (2007) O”szibarack (Peach). In: Soltész, M. (ed.) Magyar gyümölcsfajták (Hungarian Fruit Varieties).
Mezo”gazda Kiadó, Budapest.
Vujanic’-Varga, D. and Ognjanov, V. (1992) Conservation of vineyard peach populations in Yugoslavia. FAO/
IBPGR Plant Genetic Resources Newsletter 90, 28–30.
Wang, Z.-H. and Lu, Z.-X. (1992) Advances in peach breeding in China. HortScience 27, 729–732.
Weldon, G.P. and Lesley, J.W. (1933). The Babcock peach. University of California Agricultural Experiment
Station Circular 328.
Werner, D.J. and Okie, W.R. (1998) A history and description of the Prunus persica Plant Introduction collec-
tion. HortScience 33, 787–793.
Werner, D.J. and Ritchie, D.F. (1982) Peach cultivars introduced by the North Carolina Agricultural Research
Service 1965 to 1981. North Carolina Agricultural Research Service Station Bulletin 464.
Yamamoto, T., Mochida, K. and Hayashi, T. (2003) Shanhai Suimitsuto, one of the origins of Japanese peach
cultivars. Journal of the Japanese Society for Horticultural Science 72, 116–121.
Yezhov, V.N., Smykov, A.V., Smykov, V.K., Khokhlov, S.Y., Zaurov, D.E., Mehlenbacher, S.A., Molnar, T.J., Gof-
freda, J.C. and Funk, C.R. (2005) Genetic resources of temperate and subtropical fruit and nut species at
the Nikita Botanical Gardens. HortScience 40, 5–9.
Yoshida, M. (1970) Genetical studies on the fruit quality of peach varieties. 1. Acidity. Bulletin of the Tree
Research Station, Series A 9, 1–15.
Yoshida, M. (1976) Genetical studies on the fruit quality of peach varieties. 3. Texture and keeping quality.
Bulletin of the Tree Research Station, Series A 3, 1–16.
Zago, F. (1923) Istruzione pomologica. L’Italia Agricola 1, 1–17.
Zaiger, F. (1988) Objectives, experiences of a California private peach breeder. In: Childers, N.F. and
Sherman, W.B. (eds) The Peach. Horticultural Publications, Gainesville, Florida, pp. 100–105.
Zambujo, C. (2004) Nectavigne. Une démarche qui redonne le sourire à la vallée du Rhône. Arboriculture
Fruitière 586, 15–16.
 7 Processing Peach Cultivar Development

T.M. Gradziel1 and J.P. McCaa2

1Department of Plant Sciences, University of California, Davis, California, USA
2Del Monte Corporation, Stockton, California, USA

7.1 Introduction 175

7.2 Harvest 176
7.3 Raw Product Supply, Transport and Storage 181
7.4 Grading 182
7.5 Fruit Halving and Pitting 184
7.6 Peeling and Blanching 186
7.7 Sorting and Canning 187
7.8 Pasteurization 188
7.9 Marketing 188

7.1 Introduction 504,486 t for the USA as a whole (Tables 7.2

The production of peaches for processing is
an important industry in many temperate
growing regions of the world (Table 7.1). In
Europe, Spain and Greece are major producers
with combined production often exceeding
500,000 t (canned net weight). Approximately
400,000 t of peaches are canned in North
America, with most occurring in California
(CCPA, 2003). In South America, Argentina
and Chile are major canned peach exporters
with combined production of approximately
112,500 t. Other important southern hemi-
sphere producers include South Africa with
approximately 100,000 t and Australia with
44,000 t. In California, the total 2002 processed
peach (canned, purée, juice, frozen and dried)
production of 587,506 t (USDA, 2004) sur-
passed the total fresh market production of
and 7.3). Total US production of processed
peach was 617,021 t in 2002 or 55% of total
fresh market and processed utilized peach
production.
In the major processing peach produc-
tion regions, cultivars are developed within
or alongside breeding programmes for fresh
market fruit since the genetics and breeding
methodologies are identical (Nakasu et al.,
1981; Bellini, 1985; Moore, 1985; Fideghelli,
1987; Zhang et al., 1996; Layne, 1997; Todorovic
et al., 1998; Mishich et al., 1998; Etienne et al.,
2002). Information on peach genetics and
genetic recombination strategies presented in
Chapter 3 are thus applicable to the breeding
of processing peach cultivars. The cultivar
improvement programmes differ primarily
in the breeding objectives and consequently
the selection criteria, as fresh market and

© CAB International 2008. The Peach: Botany, Production and Uses

(eds D.R. Layne and D. Bassi) 175
176 T.M. Gradziel and J.P. McCaa

Table 7.1. Estimated 2003/4 world production of processing cultivar ideotypes differ in impor-
canned peaches and change in production from tant and sometimes dramatic ways.
the previous year. Southern hemisphere figures A major difference between processing
are for 2002/3 marketing year. (From CCPA, and fresh market cultivars is that processing
2003.)
peaches are largely handled as a bulk com-
Country Production (t) Change (%)
modity, requiring greater fruit durability and
uniformity for bulk handling and processing
USA 380,270 −5 and greater yields to compensate for the gen-
Spain 154,000 −8 erally lower price for the raw product. In
South Africa 98,000 +27 addition, since most flavour volatiles are lost
Argentina 67,000 +43 during processing, processing peach breed-
Greecea 52,000 −82 ing programmes must emphasize other com-
Chile 45,500 +17 ponents of peach flavour (Sistrunk et al., 1979;
Australia 44,000 +8 Gonzalez et al., 1992; Mishich et al., 1998; Blanc
All other 17,500 N/A and Arregui, 1999). While some fresh market
Total 858,270 −24 freestone cultivars are processed (Tables 7.2
aGreek productivity is down from a typical and 7.3), their soft melting flesh and typical
production of 400,000 t following two consecutive high levels of water-soluble anthocyanin pig-
years of severe frost damage. ments result in poor processed quality unless
processing is by rapid freezing or drying.
Approximately 83% of all processed peach
cultivars utilize the clingstone, non-melting
flesh trait (Table 7.3) since the firm flesh of
Table 7.2. Total 2002 utilized processing peach
such fruit (Fig. 7.1/Plate 55) is more resistant
production in the USA. (From USDA, 2004.)
to physical damage during bulk fruit harvest,
Use and type State Production (t) transport, processing and fruit slicing (Carles,
1984).
Canned Canning (including purée and juice)
Clingstone California 464,025 accounts for approximately 83% of all pro-
Others 17,418 cessed peach tonnage in the USA with other
Frozen
products being frozen slices and diced prod-
Freestone California 82,645
ucts, dried peaches and baby food (Table 7.2).
Others 9,570
Dried Common processed forms include peach
Freestone California 12,882 halves, sliced or diced peach, purée and juice.
Purée, juice, etc. While the final product and even the method
Clingstone California 27,954 of preparation may vary, most processing
Freestone All 2,527 involves a common sequence of basic events
Total California 590,033 as generalized in Fig. 7.2 (Dauthy, 1995; Bur-
USA 617,021 rows, 2001). To be successful, a cultivar must
be compatible with the nuances of every com-
ponent of the processing pathway. For this
reason it is wise to consider cultivar selection
Table 7.3. Comparison of 2002 utilized fresh
criteria within the framework of this path-
market and processing peach production for the
way.
USA. (From USDA, 2004.)

Utilization Production (t)

Total fresh market 504,486 7.2 Harvest

Total processing freestone 107,624
Total processing clingstone 509,397
Peaches for processing are generally harvested
Total processing 617,021
near the tree-ripe stage to maximize fruit size,
Total fresh market and processing 1,121,507
yield and quality (Souty, 1972; Elkins, 1979;
 Processing Peach Cultivar Development
Fig. 7.1. Processing clingstone peach showing the uniformly yellow, firm, non-melting flesh and
associated clingstone type stone-to-flesh adhesion.
Sistrunk et al., 1979; Carles, 1984; Delwiche
et al., 1987; Brooks et al., 1993; Wang et al.,
1993; Mishich et al., 1998; Blanc and Arregui,
1999). Most freestone peaches, because of their
characteristic soft, melting flesh at maturity,
are harvested at an earlier ripening stage
prior to final flesh softening (see Chapter 5).
The non-melting clingstone peach (Fig. 7.1/
Plate 55) not only allows harvest at full matu-
rity, but also the more rapid, bulk harvest
required to compensate for the generally
lower farm-gate value of processing cling-
stone peaches (Bourne, 1974; Pressey and
Avants, 1978; Sistrunk et al., 1979). Machine
harvest is sometimes used but can result in
considerable fruit damage if tree, fruit and
equipment attributes are not optimized (Ray
and Morris, 1974; Zocca, 1975; Zocca and
Fridley, 1977). Besides the non-melting cling-
stone fruit type, other traits which optimize
harvest efficiency include open and accessible
tree architecture, ease of fruit removal from
peduncle and uniform ripening (Sistrunk
et al., 1979; Egea et al., 1989; Grossman and
DeJong, 1998).
A wide range of tree architectures is pos-
sible in peach because of the genetic variations
177
available (Fideghelli et al., 1979; Scorza and
Lightner, 1989; Gradziel and Beres, 1993; see
also Chapters 3 and 6) and the common use of
training and pruning practices to modify tree
growth (Scorza and Sherman, 1996; Gross-
man and DeJong, 1998; DeJong et al., 1999).
The open vase and, under higher densities,
the perpendicular V system presently enjoy a
wide popularity due to their efficient light
interception and easier maintenance (Gross-
man and DeJong, 1998; Mishich et al., 1998;
DeJong et al., 1999). Differences in time to fruit
maturity occur even within the more open
tree architectures because of differences in
fruit position within the scaffold and within
the tree (Ray and Morris, 1974; Zocca, 1975;
Zocca and Fridley, 1977; DeJong et al., 1999).
Less fruit thinning is applied to processing
peach, resulting in yields that can be up to
5 t/ha higher than fresh market freestone
varieties (USDA, 2004), but which acts to fur-
ther prolong the ripening period. Uneven rip-
ening results in the need for inefficient multiple
hand pickings or a high percentage of low-
grade fruit if once-over or mechanical harvest-
ing is employed (Ray and Morris, 1974; Zocca,
1975; Zocca and Fridley, 1977). Peach cultivars
178 T.M. Gradziel and J.P. McCaa

contribute to uneven ripening and respond

Harvest
Transport
Grading
Fruit halving and pitting
Peeling and blanching
Sorting and canning
Pasteurization
Marketing
Fig. 7.2. A flow diagram for processing peach.
differ in the duration of both the flowering
period and the fruit ripening period (Sistrunk
et al., 1979; Monastra et al., 1984; Fideghelli,
1987; Egea et al., 1989; Lopez-Rivares and
Suarez-Garcia, 1994; Paunovic et al., 1996;
Layne, 1997; Berman et al., 1998; Okie, 1998;
Todorovic et al., 1998; Etienne et al., 2002).
Flower and fruit ripening period both
favourably to selection (Hansche et al., 1972;
see also Chapter 3). Even the most uniform-
flowering cultivars will show differences in
ripening time resulting from differences in
sunlight exposure due to fruit position within
the canopy. Within these physiological limita-
tions, multiple harvest can still be minimized
through selection for precocious fruit flesh
ripening (Gradziel, 1994) or through the selec-
tion of genotypes which resist rapid fruit qual-
ity deterioration and fruit abscission once full
fruit maturity has been achieved, thus allow-
ing delayed harvests (Sistrunk et al., 1979).
The complete and rapid formation of a fruit
abscission zone is also required for efficient
harvest but the cultivar should not be prone
to preharvest fruit drop.
Harvest yield remains the single most
important determinant of cultivar success
and one of the most difficult for developing
efficient selection indices. Yield performance
must be considered both within season and
over the productive life of the orchard (DeJong
et al., 1999). A typical processing peach orchard
in California is expected to be productive for
20–30 years. Similarly, cultivar turnover is
lower in processing peach compared with
fresh market cultivars, which are more fre-
quently replaced due to changing market
preference (Scorza and Sherman, 1996). Many
of the major processing peaches in world pro-
duction, including ‘Loadel’, ‘Carson’, ‘Andross’
and ‘Halford’, were discovered over 50 years
ago, with others such as ‘Sullivan #4’ released
in 1929 (Anon., 1997; Okie, 1998) (Figs 7.3 and
7.4/Plates 56 and 57). Processing cultivars
also need to be adaptable to a greater range in
growing locations and environmental condi-
tions. Extensive field-testing is currently the
only dependable indicator of ultimate culti-
var performance (Monastra et al., 1984; Egea
et al., 1989; Lopez-Rivares and Suarez-Garcia,
1994; Layne, 1997). While controlled hybrid-
izations and the subsequent selection among
resultant seedlings may consume 4–10 years
of a breeding cycle, replicated regional trials
of resulting selections typically consume an
additional 10–16 years (Fig. 7.5/Plate 58).
Breeding cycles exceeding 20 years from the
initial hybridization to cultivar release are not
unusual in processing cultivar development
 Processing Peach Cultivar Development 179

Loadel(1) 0
Stanislaus(2) 3
Carson(3) 5
Dee-Six (3) 5
Goodwin(3) 14
Bowen(1) 16
Fay Elberta(1) 17
Andross(3) 18
Arakelian(1) 19
Peak(1) 21
Klamp(3) 21
Tuolume(2) 22
Andora(1) 23
Ross(3) 23
Rizzi(3) 28
Dr. Davis(3) 30
Carolyn(3) 31
Monaco(1) 33
Lilleland(3) 35
Halford(1) 36
Everts(1) 37
Wiser(1) 38
Riegels(3) 38
Stam(1) 39
Hesse(3) 40
Sullivan #4(1) 41
Corona(3) 44
0 5 10 15 20 25 30 35 40 45 50

Fig. 7.3. Harvest sequence days of California processing peach cultivars using the fresh market free-
stone ‘Fay Elberta’ as a reference cultivar (1, grower selection; 2, private breeder release; 3, released by
public breeding programme).

programmes (Fig. 7.4/Plate 57). This need for
more extensive testing has made processing
peach breeding less profitable for private breed-
ing programmes, with most processing peach
cultivars being released by public breeding
programmes or as chance seedlings selected
by growers (Fig. 7.3/Plate 56).
Estimates of yield heritability based on
current selection indices are typically less
than 10% (Hansche et al., 1972). Emerging
strategies of marker-assisted selection pro-
vide opportunities for more efficient genetic
selection for factors controlling yield (Bliss
et al., 2002; Etienne et al., 2002; Aranzana et al.,
2003; see also Chapter 4). The effective inter-
pretation of molecular marker data, however,
will require an improved understanding of
the developmental/physiological determinants
of crop yield. At the most basic level, peach
yield is a function of crop load and fruit size
(Berman et al., 1998). Despite the high costs of
subsequent fruit thinning (DeJong et al., 1999),
initial high flower/fruit densities are desir-
able as a buffer against subsequent detrimen-
tal environments such as insufficient winter
chilling, frost or storm damage, which can
greatly reduced crop potential. Recent prog-
ress in characterizing fruit sizing potential has
180 T.M. Gradziel and J.P. McCaa

Corona Hesse

Late
4 Ross
Carolyn Dr.Davis
Everts Rizzi
Ripe period

Andora Lilleland
3 Ross Riegels
Klamt

Bowen
Fortuna Andross
2
Goodwin
Jungerman Tufts
Carson Dee-Six
1
1940 1946 1952 1958 1964 1970 1976 1982 1988 1994 2000
1943 1949 1955 1961 1967 1973 1979 1985 1991 1997
Year

Fig. 7.4. History of processing clingstone cultivar release by public breeding programmes in
California showing punctuated release at approximately 20-year intervals (ripe period: 1, extra early;
to 4, extra late).

Crosses for
genetic
Cultivar Select parents recombination
release for desired
traits (1 year)

(1–5 years)

Select among
Evaluate
progeny
performance at
multiple sites
(2–5 years)
and years

(8–16 years)

Fig. 7.5. Basic components of a processing peach breeding programme with estimated duration in
years.
 Processing Peach Cultivar Development
been made with the development of increas-
ingly accurate models of vegetative and
reproductive growth in clingstone peach (Ber-
man et al., 1998; DeJong et al., 1999) and the
use of these models in characterizing the rela-
tionships between cultivar yield and accumu-
lated heat units (Berman et al., 1998). Theoretical
maximum yields predicted by these models
approach 70 t/ha, approximately 20 t/ha
above the current industry average, but this is
comparable to yields obtained in controlled
field studies (Berman et al., 1998).
7.3 Raw Product Supply, Transport
and Storage
Efficient fruit processing is typically a
‘supply-on-demand’ enterprise, as posthar-
vest storage is generally too expensive to be
used routinely. Due to the high financial
investment in equipment, however, fruit pro-
cessing facilities need to run at high capacity
over the entire duration of the harvest season
in order to maximize financial return. To
ensure a uniform supply of raw product for
such ‘just-in-time’ delivery, processing plant
fieldsmen utilize fruit from different locations
and, when necessary, from cold storage (Ray
and Morris, 1974; Brecht et al., 1982; Kader
et al., 1982; Kader, 1997). The preferred method
of securing a constant supply of raw product,
however, is through the planting of cultivars
with sequential ripening times (Fig. 7.3/Plate
56). Ripening time in peach, while apparently
controlled by many genes, has heritability as
high as 0.84 (Hansche et al., 1972; Hansche
and Boynton, 1986a). Such high response to
selection has been confirmed in many breed-
ing programmes where targeted maturity
time has generally been achieved by selecting
parents having ripe dates which bracket the
desired maturity time. While such breeding
strategies often result in a large proportion of
progeny ripening within the desired harvest
period, occasionally the targeted harvest time
will defy this parental averaging approach.
The consequent gap in raw product supply
can become a serious impediment to process-
ing efficiency as shown by the 10-day produc-
tion gap between the cultivars ‘Carson’ and
181
‘Bowen’ (Fig. 7.3/Plate 56). The genetic nature
of such aberrant maturity gaps is demon-
strated in the distribution of progeny obtained
by selfing the ‘Carson’ cultivar (Fig. 7.6/Plate
59). Rather than the expected normal distri-
bution of ripening times centred near that of
the selfed parent (or the parental mean in
cross-pollinations), a bimodal distribution is
observed where the ripening gap is similar to
that observed in Fig. 7.3/Plate 56. The nature
of such production gaps suggests the presence
of inherent fruit developmental thresholds.
For example, if the threshold event is achieved
within the required time (perhaps relative
to vegetative or endocarp growth), normal
development will continue. For genotypes
that have not yet achieved the developmental
threshold, however, further development will
be delayed. This problem is often overcome
by bringing in outside, unrelated germplasm
for hybridization with locally adapted breed-
ing lines.
Raw product availability will also limit
the duration of the processing season. Mid-
season cultivars tend to show the highest per
hectare yields, while yields in later-maturing
cultivars decline. With increasingly earlier-
maturing cultivars, the reduction of available
solar heat/light units becomes a major limita-
tion to potential yield. Berman et al. (1998)
have estimated yield penalties as high as 1.8
t/ha per day for genotypes ripening earlier
than the ‘Loadel’ ripening period as charted
in Fig. 7.3/Plate 56.
Yield reductions from postharvest losses
occur primarily as physical and disease
damage during transport. O’Brien et al. (1978)
found that only 56% of harvested fruit
remained unbruised following typical trans-
port to the processing plant. Physical protu-
berances, including peduncles remaining
attached to the fruit and excessive ‘beaking’
of the fruit stylar end, will damage adjacent
fruit in the bulk bins and so should be avoided.
Damaged fruit is subsequently much more
susceptible to disease. While processing
peach cultivars are generally susceptible to
the same diseases as fresh market peaches,
the greater physical contact between fruit
during harvest, transport and storage makes
processing peach much more vulnerable to
postharvest losses from fungal pathogens
182
20
15
Number of seedings
10
5
0
5 15 25 35
Days after 1 June
such as Monilinia, Alternaria and Botrytis spp.,
which spread rapidly when fruit are in con-
tact (Chapter 15). Losses during postharvest
storage are primarily from such rapidly
spreading fungal diseases and from the dete-
rioration in fruit quality (Brecht et al., 1982).
Selection for resistance to postharvest disease
(Nakasu et al., 1981; Gradziel and Wang, 1993;
Gradziel, 1994) and physiological deteriora-
tion (Kader et al., 1982; Robertson et al., 1992;
Gradziel et al., 1993a,b; Kader, 1997) has been
successful, although the underlying mecha-
nisms are only now being characterized
(Crisosto and Labavitch, 2002). The use of
modified atmospheres to prolong fruit qual-
ity has shown only limited success for pro-
cessing peach (Brecht et al., 1982; Kader, 1997)
although cold storage in commercial facilities
capable of manipulating storage atmospheres
is routinely practised in California.
7.4 Grading
Prior to processing, fruit is typically evaluated
either at grading stations near the orchard or at
the processing plant. The grading process,
which often determines the price paid for the
T.M. Gradziel and J.P. McCaa
Fig. 7.6. Fruit ripe date for
progeny from self-pollination
of the cultivar ‘Carson’
showing unusual bimodal
distribution (‘Carson’ normally
45 55 65 75 85 ripens approximately 40 days
after 1 June).
shipment, evaluates fruit size, maturity, firm-
ness and defects. Fruits smaller than 6 cm in
diameter are often not purchased because
they are difficult to process and result in an
inferior product. Fruits larger than approxi-
mately 9 cm are similarly difficult to process
in equipment adjusted for more standard
sizes, and the higher levels of fruit thinning
needed to achieve the larger size contributes
to reduced yields. For this reason, the highest-
yielding cultivars produce large numbers of
uniform size fruit of approximately 6–9 cm in
diameter with undersized fruit typically rep-
resenting less than 5% of total production.
Fruit maturity is characterized primarily
by flesh colour after removing approximately
1 cm of the epidermis and underlying meso-
carp tissue (Delwiche et al., 1987). Internal
colour is compared against a colour standard,
with fruit failing to achieve minimum colour
levels being considered ‘green’ fruit and often
not purchased. Flesh colour is determined
primarily by the carotenoid pigments b-caro-
tene and b-cryptoxanthin (Tourjee et al., 1998).
Although the carotenoid biosynthesis path-
way appears to be influenced by several genes
(Chapter 3), selection for fruit flesh colours
ranging from golden-yellow to yellow-orange
is readily achieved. Selection for high levels
 Processing Peach Cultivar Development
of b-carotene may result in fruit with an unde-
sirable dull orange flesh colour and a ‘melon-
like’ aftertaste. Higher b-cryptoxanthin levels
result in a more desirable yellow-gold flesh
colour without aftertaste or loss of provita-
min A carotenoid content (Tourjee et al., 1998).
Red pigmentation of fruit flesh is strongly
selected against in processing peach cultivars
because, unlike the water-insoluble and heat-
stable carotenoid pigments, the water-soluble
red anthocyanin pigments break down dur-
ing the cooking process, resulting in an unde-
sirable brown staining of fruit flesh and syrup.
Since the fruit skin or epidermis is removed
during processing, some red pigmentation or
‘blush’ is tolerated in processing cultivars
(Fig. 7.1/Plate 55). Excessive red skin pig-
mentation, however, can lead to staining of
the underlying tissue with associated fruit
peeling problems (Fig. 7.7/Plate 60). Areas of
red skin pigmentation are also associated
with higher levels of bird damage in the field
(T.M. Gradziel, personal observation).
Some recent processing peach cultivars,
such as ‘Maria Serena’ (Bellini, 1983) and
‘Maria Dorata’ (Bellini, 2000) from Italy, show
an apparent complete suppression of red antho-
cyanin pigmentation in the fruit. Chapparo
Fig. 7.7. Poor lye peeling of a clingstone genotype with unacceptably thick epidermis.
183
et al. (1995) have characterized the inheri-
tance, genetic interaction and biochemistry of
known anthocyanin phenotypes in peach,
and studies have evaluated the genetic con-
trol of the anthocyanin-free trait (see Chapter 1)
as well as its consequence on fruit sensory
quality (T. Beckman, USA, 2004, personal
communication).
Peach flesh firmness is commonly mea-
sured with hand-held penetrometer-type pres-
sure testers because of their portability and
ease of operation (Kader et al., 1982; Kader,
1997). Clingstone fruit flesh tends to be het-
erogeneous, often being more fibrous near the
endocarp and more homogeneous near the
epidermis (Fig. 7.8/Plate 61). Although more
sophisticated instruments exist to measure
the change in texture throughout the tissue,
providing more precise information, they are
more cumbersome to use for routine sam-
pling (Bourne, 1974). Peaches with flesh punc-
ture pressures of less than 3 kg (after skin
removal, measured with an 8 mm flat probe)
are often damaged during processing. Flesh
pressures greater than 6 kg, however, are asso-
ciated with excessive adhesion between the
stony endocarp and fleshy mesocarp, leading
to problems in pit removal and, more seriously,
184 T.M. Gradziel and J.P. McCaa

Fig. 7.8. Halved section of an overripe clingstone peach demonstrating the common pattern of
vascular strands radiating from endocarp to outer mesocarp.

the retention of broken pit shards in pro-
cessed fruit, making it hazardous to con-
sumers. Endocarp cracking early in fruit
development can also lead to large shards of
the fractured pit being embedded in the fleshy
mesocarp. These early-split pits may lead to
later splitting of the mesocarp or flesh along
the suture line, encouraging disease and insect
contamination. Invisible split pits, in which
the mesocarp remains intact, can often be
identified by abnormally large fruit size and
greater fruit asymmetry resulting from inter-
nal endocarp splitting and wound tissue
development. Although such fruit can be sal-
vaged by hand pitting, the extra cost of this
special treatment makes split pits and geno-
types prone to this problem undesirable.
Even in high-quality processing culti-
vars, the flesh colour and firmness can vary
considerably with maturation from unripe to
overripe fruit (Kader et al., 1982; Gradziel,
1994; Kader, 1997). Precise indices of fruit
maturity are thus essential for accurate fruit
grading. Despite a general selection for uni-
form fruit flesh colour, commercial processing
cultivars can also vary considerably in their
final flesh colour, which can confound fruit
grading (Delwiche et al., 1987). For this rea-
son, it is common to use separate colour stan-
dards to grade fruit maturity of different
cultivar groups and/or maturity periods.
7.5 Fruit Halving and Pitting
Fruit halving and pitting can be done by hand
or by machine. Hand cutting, while more
expensive, can utilize a range of different fruit
sizes, textures and pit qualities, including
split and broken pits. Hand pitting is typi-
cally achieved by sliding a sharp, concave-
shaped knife under the pit to remove it from
the flesh. This cutting action results in an
undesirable ‘scooped-out’ appearance of the
fruit and most hand-pitted or irregularly cut
fruit are diverted to sliced and diced products,
where the irregularities are less apparent, or,
if unsuitable for this, used in purées and pie
fillings (Burrows, 2001). High-quality fruit
 Processing Peach Cultivar Development
may also be puréed directly to produce pre-
mium processed products such as baby foods.
The common method of mechanical pit-
ting uses a rotary torque pitter, where the
individual fruit and pit is clasped by sharp
knives from opposite sides at the suture line
while counter-rotating rubber cups on either
side of the fruit torque the cut halves away
from the partially immobilized pit. The proper
alignment of the peach suture with the pitting
machine cutting knives is done mechanically
by using the asymmetry of the peduncle cav-
ity to determine the suture orientation (Del-
wiche, 1989). Consequently, peach cultivars
developed for mechanical pitting need to have
a well-defined peduncle asymmetry, need to
harvest free from peduncle and need to have a
sufficiently long peduncle. Very short pedun-
cle attaching the fruit to the tree branch will
result in depressions at the peduncle area
where the fruit grows around the branch. The
resulting branch imprint can mimic the
peduncle cavity, resulting in misalignment of
fruit going into the mechanical pitter.
Pitting machines are very efficient but
need to be adjusted for specific fruit shapes,
textures and fruit and pit sizes. Different fruit
sizes are handled by first sorting incoming
fruit and directing them to the appropriately
adjusted mechanical pitter. Variations in fruit
shape and firmness and pit size and integrity
(split, fragmented, etc.) are more difficult to
manage and are common causes for mechani-
cal pitter failures and consequent shut-down
of associated processing lines. Uniformity for
these characteristics is thus a prerequisite for
processing cultivars. The genetic control of
these traits is still poorly understood (Han-
sche et al., 1972; Scorza and Lightner, 1989; see
also Chapter 1). While it is thus initially diffi-
cult to predict progeny performance from
parental genotypes, experience with different
crossing combinations will identify geno-
types more frequently associated with infe-
rior progeny in one or more of these traits,
and such genotypes should be avoided. Gen-
otypes which normally develop good fruit
symmetry can occasionally develop excessive
fruit elongation with ‘beaking’ at the stylar
end, particularly following warm winter/early
spring conditions. While often associated with
low winter-chill conditions, this elongated fruit
185
growth may be a consequence of the warmer
temperatures during early spring growth. For
this reason, breeding selections showing erratic
performance under growing conditions likely
to be encountered during commercial pro-
duction should be rouged-out.
A serious problem with mechanical pit-
ting is the breakage of fragments from the
ridged pit that then become embedded in the
processed product. While certain genotypes
are more prone to this problem, the non-melting
flesh canning clingstone peach is inherently
predisposed to embedded fragments due to
the clingstone or high stone-to-flesh adhesion
associated with the non-melting flesh trait.
Although recent genetic studies suggest that
the two traits may be controlled by separate
genes which are highly linked, over 50 years
of intense breeding efforts have failed to
develop non-melting flesh freestone peach
types suitable for canning. The combination
of melting flesh with clingstone pit adhesion
is common in very early-ripening fresh mar-
ket peaches, but here the clingstone nature is
a consequence of rapid fruit development
and the true freestone genotype can be read-
ily determined by crossing to later-maturing
tester lines. Gradziel (2002) has reported the
recovery of good-quality, firm-flesh freestone
peaches in backcrosses from peach × almond
(Prunus dulcis) interspecific hybrids. While
the controlling genes appear distinct from both
freestone and clingstone, the trait appears
unstable in certain environments, resulting in
rapid flesh softening in overripe fruit. A sec-
ond strategy to breeding firm-flesh, freestone
peach involves the incorporation of the ‘stony-
hard’ trait into high-quality freestone types.
Stony hard also appears to be genetically dis-
tinct from the freestone/clingstone trait and
acts by suppressing the normal ethylene-in-
duced fruit softening at ripening (Haji et al.,
2003). Breeding lines combining the stony-
hard and freestone trait have demonstrated
greater flesh firmness in ripe fruit, but the
level of firmness (typically about 3 kg) was
insufficient to avoid extensive damage dur-
ing processing and was also associated with
undesirable textural changes in processed
samples. Beckman and Sherman (1996) and
Van Der Heyden et al. (1997) have described a
non-melting, semi-freestone trait which, while
186 T.M. Gradziel and J.P. McCaa
not yet fully genetically characterized, appears
to be readily recovered in certain crossing com-
binations (T.M. Gradziel, personal observation).
Unfortunately, the ripe fruit remains softer than
clingstone fruit and the semi-freestone trait
appears to be strongly associated with a pro-
nounced red anthocyanin pigmentation of
the fruit pit cavity, making it unacceptable for
canning (see also earlier evaluation of this
trait by Robertson et al., 1992).
Unless the association between melting
flesh and freestone habit can be altered, free-
stone cultivars will remain unsuitable for
most processing uses. Breaking this undesirable
association might be achieved through break-
age of the putative genetic linkage between
these traits or, alternatively, the incorporation of
independent genes promoting a more free-
stone tendency in non-melting genetic back-
grounds. Pressey and Avants (1978) have
reported that, unlike melting-flesh freestones,
non-melting clingstone fruit lack endopolyga-
lacturonase activity. Callahan et al. (2004) dem-
onstrated a correlation with the non-melting
flesh texture and observed deletions in the
endopolygalacturonase gene cluster. These
findings suggest that opportunities for geneti-
cally engineering a suppression of the putative
flesh-softening endopolygalacturonase gene
in freestone genotypes though pleiotrophic
effects such as unstable mesocarp integrity
may ultimately confound this approach.
The incorporation of the clingstone trait
into fresh market varieties, however, is com-
mon in many parts of the world, particularly
Europe and South America, because the asso-
ciated non-melting firm flesh facilitates ship-
ping and handling, allowing higher-quality
fruit to be harvested closer to the tree-ripe
stage (see Chapter 5). This greater versatility
of the clingstone/non-melting trait has encour-
aged the development of dual-purpose peaches
for both processing and fresh market (Wilson
and Boudreaux, 1986; Raseira et al., 1998; Bellini,
2000), thus allowing greater marketing flexi-
bility. The flavour of fresh clingstone peaches
is distinct from freestone peaches with their
more pronounced aromatic components.
Shortly after the full-ripe stage of maturity,
clingstone peaches often also develop a char-
acteristic off-flavour which appears to be
associated with alcohol metabolism.
7.6 Peeling and Blanching
While clingstone are usually distinguished from
freestone peaches by their firmer, non-melting
flesh and continued adhesion between the
endocarp and inner mesocarp in ripe fruit, they
also show greater adhesion between the exo-
carp (skin) and outer mesocarp tissue (Rotaru,
1994). In full-ripe, melting-flesh, freestone culti-
vars, a breakdown in the adhesion between the
skin and underlying flesh often allows the skin
to be readily peeled from the fruit, particularly
after blanching. The more pronounced attach-
ment in clingstone cultivars necessitates more
rigorous methods for peeling. In bulk process-
ing, clingstone peach halves are frequently
peeled by immersion in or spraying with hot
caustic solutions such as 1–2.5% lye for 30–60 s
(Dauthy, 1995; Burrows, 2001). The loosened
skin is then physically removed by passing the
peach halves under powerful water sprays
which also wash off remnants of the caustic
solution. Although caustic solutions are recy-
cled, their eventual disposal is becoming
increasingly costly and other methods of skin
removal are being evaluated. Steaming fruit to
loosen the skin followed by skin removal by
rotating rubber or abrasive rollers can be effec-
tive if the cultivar has a thinner epidermis or
less pronounced epidermis-to-mesocarp adhe-
sion. Both epidermis thickness and adhesion
are strongly affected by environmental condi-
tions, making this approach less dependable.
Poor peeling can occur in either approach for
excessively thick epidermis tissue, which
should be avoided in new cultivars (Fig. 7.7/
Plate 60). Intense red blushing of the skin should
also be avoided as it can lead to staining and
peeling problems of the underlying flesh tissue.
Carroad et al. (1980) have determined that steam
generation for use in peeling, blanching and
cooking accounts for 98% of all energy con-
sumed in a clingstone peach cannery.
Because the peel is removed in process-
ing, greater variations for epidermis charac-
teristics are tolerated in processing cultivars,
which can be exploited for improved disease
and pest resistance. More compact epidermal
cell layers, and higher levels of phenolics,
pectin and cutin, have been incorporated in pro-
cessing varieties to improve fungal resistance.
Similarly, high trichome densities with their
 Processing Peach Cultivar Development
associated pest resistance (Massonié et al.,
1982; Monet and Massonié, 1994) and peach
‘fuzziness’ are acceptable in processing culti-
vars since they are removed during process-
ing. Removal of the epidermis and underlying
tissue during processing also acts to remove
most residues from pre- and postharvest pesti-
cide treatments (Lentza, 1995; Lentza and
Chitzanidis, 1995, 1996). The range of breed-
ing options allowed by the removal of the
epidermis at processing is demonstrated by
the development of processing peach selec-
tions in which the epidermis retains its imma-
ture green colour and associated high levels
of fungal resistance at full maturity as deter-
mined by flesh colour (Gradziel, 1994).
Following peeling, peaches are blanched
for 1–2 min in steam or water at 80°C to inacti-
vate the oxidizing enzymes which otherwise
turn exposed flesh brown. Blanching also sup-
presses the oxidation and associated degenera-
tion of fruit phytonutrients, thus preserving
their availability to the consumer (Elkins, 1979).
Canning results in significant initial phytonu-
trient losses from the heat treatment but the
remaining phytonutrients are more stable dur-
ing later storage, whereas freezing will show
less initial loss but increasing loss with storage
time (Chang et al., 2000). Previous bruising of
flesh that occurred during harvest and trans-
port is not controlled by blanching. Control is
primarily by selecting against high levels of
phenolics and polyphenyloxidases which
catalyse the flesh-browning reactions. Sizeable
genetic differences in both phenolic and poly-
phenyloxidase levels have been demonstrated
in processing peach genotypes (Gradziel and
Wang, 1993; Chang et al., 2000). Hansche and
Boynton (1986b) have estimated the heritabil-
ity for fruit enzymatic browning at 0.35 for
freestone selections. A high vulnerability to
flesh bruising is also a major limitation to the
processing of white-fleshed clingstone peaches,
though white-fleshed processing cultivars
have been released (Lu et al., 1995).
7.7 Sorting and Canning
Following peeling, the halved peaches are
sorted and graded. The best-quality halves
187
are canned as Fancy USDA grade. The smaller,
less perfect fruit are designated, in diminish-
ing order, as Choice, Standard, Second and
Pie grades (Burrows, 2001). Sliced and diced
peaches can be prepared from damaged or
otherwise irregular halves. After cutting, the
fruit is inspected to remove any damaged or
blemished sections. Fruit is typically placed
into cans, then hot syrup is added and the can
is closed using steam-flow closure (Dauthy,
1995; Burrows, 2001). Container size can
range from multi-litre to single-serving tinned
metal and plastic containers or to glass packs
of generally intermediate volumes. The tin in
the metal cans act as a catalyst to preserve
fruit colour brightness (Blanc and Arregui,
1999; Burrows, 2001). The mixing of cultivars
having different flesh colours or colour bright-
ness is avoided because such differences in
fruit colour give the consumer the impression
that some of the fruit from an individual con-
tainer is either immature or over-mature.
Differences in individual fruit flavour and
sweetness tend to be masked when canned in
the traditional 30°Brix syrup or natural fruit
juice. Raw fruit flavour and sugar level become
more important when fruit is packed in
lighter syrup or water. Kader et al. (1982) and
Kader (1997) have reported that fresh fruit
flavour is positively correlated to processed
flavour when established cultivars are evalu-
ated. However, the fruit volatiles normally
associated with flavour in fresh peaches are
lost in processing and cultivar flavour is
largely determined by sugar content and still
poorly understood non-volatile flavour com-
ponents such as organic acids (Gonzalez et al.,
1992). Genetic variability in both fruit sugar
and organic acid content has been character-
ized in processing peach (Gonzalez et al.,
1992; Brooks et al., 1993; Wang et al., 1993; Eti-
enne et al., 2002) though it remains largely
unexploited, partly due to low heritability
(Etienne et al., 2002). Hansche et al. (1972) and
Hansche and Boynton (1986a) estimated heri-
tability of 0.17 and 0.35, respectively, for free-
stone selections. While reporting low heritability
for the individual processing clingstone sug-
ars sucrose, glucose, fructose and sorbitol,
Brooks et al. (1993) reported heritability of 0.72
or higher for total soluble solids, acidity and
sugar:acid ratios.
188 T.M. Gradziel and J.P. McCaa
In the preliminary evaluations of breed-
ing selections, fresh flavour, with the excep-
tion of soluble solids content, is generally not
a good indicator of processed product flavour
since significant changes can occur with pro-
cessing. The nectarine cultivars, for example,
with their typical high levels of acids, sugars
and aromatic volatiles, can develop ‘off-fla-
vours’ when processed. Consequently, while
clingstone nectarines exist (see Bellini, 2000),
they are typically not used for processing.
The processing of samples under commercial
conditions remains the only effective way to
evaluate processed quality of new selections.
Fruit texture, phytonutrient content and colour
can also change with processing, sometimes
in unanticipated ways (Tourjee et al., 1998;
Chang et al., 2000).
Poor fruit quality, including damaged,
blemished and undersized fruit, reduces case
yield efficiency (the proportion of the initial
raw product that can be successfully pro-
cessed). Large pit size and poor pit quality
leading to split and/or fragmented pits are
major causes of case yield reductions. A red
anthocyanin staining and/or brown, corky,
imprinting of the fruit pit cavity is often asso-
ciated with greater pit fragmentation and
may be related to modified lignin synthesis
since the development pathways are physio-
logically related. Because anthocyanins are
heat-labile and water-soluble, cultivars express-
ing this trait are undesirable as the red antho-
cyanins tend to oxidize to a red-brown colour
when processed, leading to staining of the
syrup in addition to the peach flesh.
7.8 Pasteurization
In the final stage of processing, the sealed and
canned fruit are pasteurized in hot water or
atmospheric steam followed by rapid cool-
ing. The cooking time will depend on the con-
tainer volume, the type of fruit product and
fruit pH. At a pH of 4.6 or higher, Clostridium
spp. and related bacteria can become a con-
tamination problem if not cooked for longer
periods. Increased cooking time, however,
results in decreased firmness of the processed
product. Ca additives may be added to pre-
serve textural firmness but are not commonly
utilized (Dauthy, 1995; Burrows, 2001). Lower
fruit pH levels reduce the required cooking
time and so the associated fruit softening, but
also alter the sugar:acid ratio, which is often
associated with improved flavour perception
(Sistrunk et al., 1979; Gonzalez et al., 1992;
Wang et al., 1993). The raw fruit pH of process-
ing cultivars typically ranges from 3.5 to 4.5
and while lower pH selections are possible,
they are not widely utilized in processing.
Citric acid may also be added to acidify the
fruit. At pH values of less than 4, bacteria will
not multiply and shorter pasteurization treat-
ment is allowed (Burrows, 2001). Aseptic pro-
cessing, using rapid sterilization techniques
which are less damaging to fruit quality, are
routinely used for purées and juices (Bettison,
2001; Burrows, 2001). Technological advances
to allow aseptic processing of whole or
sectioned fruit offer the promise of better pres-
ervation of fruit flavour, texture and phyto-
nutrient content (Leonard et al., 1983).
7.9 Marketing
Market globalization of the processing food
industries is imposing greater uniformity and
standardization on the final processed prod-
uct. Greater market flexibility, including single-
serving packaging size, greater phytonutrient
content and new fruit mixtures (cocktails),
are moving the processed peach ideotype to
more compact sizes and more yellow-gold
flesh colour. Both processors and consumers
are demanding reduced pesticide residues in
the product. Global competition among both
processed and fresh fruits has promoted the
processing of a premium product, competi-
tive with fresh market flavour and eating
quality, yet with the convenience and versa-
tility of convenient packaging, storability and
phytonutrient enhancement. This continuing
emphasis on processed product quality is
shifting the measure of industry productivity
from orchard or production yield to case yield
(the proportion of the initial raw product that
can be processed to final products meeting
the raised quality standards). To achieve these
higher case yields, new cultivars will be required
 Processing Peach Cultivar Development 189

to produce a higher-quality product with
reduced agrochemical inputs yet without loss
in orchard productivity. While genetic options
within the traditional peach germplasm
appear too limited to achieve these larger
breeding goals, a rich source of new germ-
plasm has recently been characterized within
wild peach (Pascal et al., 1997) and almond
relatives (Martínez-Gómez et al., 2003a,b). Suc-
cessfully introgressed traits include improved
fruit flavour and textural quality, pest resis-
tance and ripening habits allowing once-over
harvesting (Gradziel, 2002). Unlike seed-
propagated annual crops, processing peach
enjoys several advantages for such novel gene
introgression. Progeny combining the best traits
from both parents, while rare, are propagated
vegetatively, effectively capturing the elite
genotype. The extensive cultural management
typical for peach allows considerable flexibil-
ity in adapting to atypical tree characteristics,
which can often be managed by modified cul-
tural practices. Technologies utilized within
the processing stream (Fig. 7.2) can also be
adapted to handle novel fruit types as long as
case yield efficiencies are maintained. For
example, the use of high-speed colour sorters
allows the concurrent processing of standard
peach types along with newer orange-gold
flesh types with their higher provitamin A con-
tents. Similarly, modifications of the standard
mechanical pitter used for freestone peaches
allow their use with firm, non-melting flesh,
freestone types. Concurrently, the develop-
ment of new technologies for rapid pasteuri-
zation and aseptic packaging will make
available minimally processed, tree-ripe fruit
in consumer-convenient packaging allowing
extended storage at ambient temperatures. In
effect, processed fruit will become less an
alternative to fresh fruit and more an aug-
mented or ‘value-added’ form of fresh fruit.

References

Anon. (1997) The Brooks and Olmo Register of New Fruit and Nut Varieties. ASHS Press, Alexandria, Virginia.
Aranzana, M.J., Pineda, A., Cosson, P., Dirlewanger, E., Ascasibar, J., Cipriani, G., Ryder, C.D., Testolin, R.,
Abbott, A., King, G.J., Iezzoni, A.F. and Arús, P. (2003) A set of simple-sequence repeat (SSR) markers
covering the Prunus genome. Theoretical & Applied Genetics 106, 819–825.
Beckman, T.G. and Sherman, W.B. (1996) The non melting semi-freestone peach. Fruit Varieties Journal 50,
189–193.
Bellini, E. (1983) Maria Serena, a new early maturing peach for processing. Rivista di Frutticoltura e di Orto-
floricoltura 45, 9–10.
Bellini, E. (1985) Recent changes in peach growing in Italy. Ponte del Concordato Italiano Grandine 52, 52–70.
Bellini, E. (2000) Maria Dorata, an anthocyanin free nectarine for processing and fresh consumption.
L’Informatore Agrario 56, 63–64.
Berman, M.E., Rosati, A., Pace, L., Grossman, Y.L. and DeJong, T.M. (1998) Using simulation modeling to es-
timate the relationship between date of fruit maturity and yield potential in peach. Fruit Varieties Journal
52, 229–235.
Bettison, J. (2001) Packaging for fruit products. In: Arthey, D. and Ashurst, P.R. (eds) Fruit Processing, Nutrition,
Products, and Quality Management, 2nd edn. Aspen Publishers, Gaithersburg, Maryland, pp. 205–224.
Blanc, P. and Arregui, M. (1999) Tinned peaches: the clingstone peach reviewed for today’s taste. Arboricul-
ture Fruitière 533, 33–36.
Bliss, F.A., Arulsekar, S., Foolad, M.R., Becerra, V., Gillen, A.M., Warburton, M.L., Dandekar, A.M., Kocsisne,
G.M. and Mydin, K.K. (2002) An expanded genetic linkage map of Prunus based on an interspecific
cross between almond and peach. Genome 45, 520–529.
Bourne, M.C. (1974) Textural changes in ripening peaches. Canadian Institute of Food Science and Technol-
ogy Journal 7, 11–15.
Brecht, J.K., Kader, A.A., Heintz, C.M. and Norona, R.C. (1982) Controlled atmosphere and ethylene effects on
quality of California canning apricots and clingstone peaches. Journal of Food Science 47, 432–436.
Brooks, S.J., Moore, J.N. and Murphy, J.B. (1993) Quantitative and qualitative changes in sugar content of
peach genotypes (Prunus persica (L.) Batsch.). Journal of the American Society for Horticultural Science
118, 97–100.
190 T.M. Gradziel and J.P. McCaa

Burrows, G. (2001) Production of thermally processed and frozen fruit. In: Arthey, D. and Ashurst, P.R. (eds) Fruit
Processing, Nutrition, Products, and Quality Management, 2nd edn. Aspen Publishers, Gaithersburg,
Maryland, pp. 149–176.
Callahan, A.M., Scorza, R.M., Bassett, C., Nickerson, M. and Abeles, F.B. (2004) Deletions in an endopoly-
galacuronase gene cluster correlate with non-melting flesh texture in peach. Functional Plant Biology
31, 159–168.
Carles, L. (1984) Clingstone peaches, fruits specially adapted for the manufacture of fruits in syrup. Arboriculture
Fruitière 31, 47–48.
Carroad, P.A., Singh, R.P., Chinnan, M.S., Jacob, N.L. and Rose, W.W. (1980) Peach processing. Journal of
Food Science 45, 723–725.
CCPA (2003) USDA: world canned peach supply falls. Cling Peach Review 39, 8.
Chang, S., Tan, C., Frankel, E.N. and Barrett, D.M. (2000) Low density lipoprotein antioxidant activity of
phenolic compounds and polyphenol oxidase activity in selected clingstone peach cultivars. Journal
of Agricultural and Food Chemistry 48, 147–151.
Chapparo, J.X., Werner, D.J., Whetten, R.W. and O’Malley, D.M. (1995) Inheritance, genetic interaction, and
biochemical characterisation of anthocyanin phenotypes in peach. Journal of Heredity 86, 32–38.
Crisosto, C.H. and Labavitch, J.M. (2002) Developing a quantitative method to evaluate peach (Prunus persica)
flesh mealiness. Postharvest Biology and Technology 25, 151–158.
Dauthy, M.E. (1995) Fruit and Vegetable Processing. FAO Agricultural Services Bulletin No. 119. Food and
Agriculture Organization of the United Nations, Rome.
DeJong, T.M., Tsuji, W., Doyle, J.F. and Grossman, Y.L. (1999) Comparative economic efficiency of four peach
production systems in California. HortScience 34, 73–78.
Delwiche, M.J. (1989) Alignment detection of clingstone peaches during pitting. American Society of Agricul-
tural Engineers No. 89-6021.
Delwiche, M.J., Tang, S. and Rumsey, J.W. (1987) Maturity evaluation of clingstone peaches by optical mea-
surement. American Society of Agricultural Engineers No. 87-6018.
Egea, L., Egea, J., Garcia, J.E. and Berenguer, T. (1989) A comparative trial of clingstone peach cultivars. ITEA
Produccion Vegetal 20, 35–43.
Elkins, E.R. (1979) Nutrient content of raw and canned green beans, peaches, and sweet potatoes. Food
Technology 33, 66–70.
Etienne, C., Rothan, C., Moing, A., Plomion, C., Bodenes, C., Dumas, L.S., Cosson, P., Pronier, V., Monet, R.
and Dirlewanger, E. (2002) Candidate genes and QTLs for sugar and organic acid content in peach (Pru-
nus persica (L.) Batsch). Theoretical & Applied Genetics 105, 145–159.
Fideghelli, C. (1987) Recent fruit cultivars from Fruit Culture Institute, Rome. Genetica Agraria 41, 201–213.
Fideghelli, C., Della Strada, G., Quarta, R. and Rosati, P. (1979) Genetic semi-dwarf peach selections. In:
Proceedings of Eucarpia Fruit Section Symposium, Tree Fruit Breeding. INRA, Angers, France, pp. 3–7.
Gonzalez, A.R., Mauromoustakos, A., Prokakis, G. and Aselage, J. (1992) Sugars and acidity of processing
peaches in Arkansas. Arkansas Farm Research 41, 11–12.
Gradziel, T.M. (1994) Changes in susceptibility to brown rot with ripening in three clingstone peach geno-
types. Journal of the American Society for Horticultural Science 119, 101–105.
Gradziel, T.M. (2002) Almond species as a source of new genes for peach improvement. Acta Horticulturae
592, 81–88.
Gradziel, T.M. and Beres, W. (1993) Semidwarf growth habit in clingstone peach with desirable tree and fruit
qualities. HortScience 28, 1045–1047.
Gradziel, T.M. and Wang, D. (1993) Evaluation of brown rot resistance and its relation to enzymatic browning
in clingstone peach germplasm. Journal of the American Society for Horticultural Science 118, 675–679.
Gradziel, T.M., Beres, W., Doyle, J. and Weeks, C. (1993a) ‘Rizzi’: a processing clingstone peach with extended
postharvest storage potential. HortScience 28, 230.
Gradziel, T.M., Beres, W. and Pelletreau, K. (1993b) Inbreeding in California canning clingstone peach culti-
vars. Fruit Varieties Journal 47, 160–168.
Grossman, Y.L. and DeJong, T.M. (1998) Training and pruning system effects on vegetative growth potential,
light interception, and cropping efficiency in peach trees. Journal of the American Society for Horticul-
tural Science 123, 1058–1064.
Haji, T., Yaegaki, H. and Yamaguchi, M. (2003) Softening in stony hard peach by ethylene and induction of
endogenous ethylene by 1-aminocyclopropane-1-carboxylic acid (ACC). Journal of the Japanese Society
of Horticultural Science 72, 212–217.
Hansche, P.E. and Boynton, B. (1986a) Heritability of juvenility in peach. HortScience 21, 1195–1197.
 Processing Peach Cultivar Development 191

Hansche, P.E. and Boynton, B. (1986b) Heritability of enzymatic browning in peaches. HortScience 21, 1197–
1198.
Hansche, P.E., Hesse, C.O. and Beres, V. (1972) Estimate of genetic and environmental effects on several traits
in peach. Journal of American Society for Horticultural Science 97, 9–12.
Kader, A.A. (1997) A summary of CA requirements and recommendations for fruits other than apples and
pears. In: Proceedings of the Seventh International Controlled Atmosphere Research Conference (CA
1997). Vol. 3. Fruits other than Apples and Pears. UC Davis Press, Davis, California, pp. 217–221.
Kader, A.A., Heintz, C.M. and Chordas, A. (1982) Postharvest quality of fresh and canned clingstone peaches
as influenced by genotypes and maturity at harvest. Journal of the American Society for Horticultural
Science 107, 947–951.
Layne, R.E.C. (1997) Peach and nectarine breeding in Canada: 1911 to 1995. Fruit Varieties Journal 51,
218–228
Lentza, R.C. (1995) Residues of iprodione in fresh and canned peaches after pre- and postharvest treatment.
Journal of Agricultural and Food Chemistry 43, 1357–1360.
Lentza, R.C. and Chitzanidis, A. (1995) Residues of benomyl on and in fresh and canned peaches following
treatments to control storage decay. Annales de l’Institut Phytopathologique Benaki 2, 121–130.
Lentza, R.C. and Chitzanidis, A. (1996) Residues of dicloran in clingstone peaches after pre- and postharvest
application. Bulletin of Environmental Contamination and Toxicology 56, 231–239.
Leonard, S.J., Heil, J.R., Carroad, P.A., Merson, R.L. and Wolcott, T.K. (1983) High vacuum flame sterilized
fruits storage study on sliced clingstone peaches, sliced Bartlett pears and diced fruit. Journal of Food
Science 48, 1484–1491.
Lopez-Rivares, E.P. and Suarez-Garcia, M.P. (1994) Agronomic performance of 19 clingstone peach cultivars
in the Middle Valley of Guadalquivir. ITEA Produccion Vegetal 90, 6–17.
Lu, G.L., Su, J.Y., Chen, Z.F., Wang, J.W. and Li, J. (1995) A promising, canning, white-fleshed peach variety
Yubai. Journal of Fruit Science 12, 51–53.
Martínez-Gómez, P., Arulsekar, S., Potter, D. and Gradziel, T.M. (2003a) An extended interspecific gene pool
available to peach and almond breeding as characterized using simple sequence repeat (SSR) markers.
Euphytica 131, 313–322.
Martínez-Gómez, P., Arulsekar, S., Potter, D. and Gradziel, T.M. (2003b). Relationships among peach, almond
and related species as detected by SSR markers. Journal of American Society for Horticultural Science
128, 667–671.
Massonié, G., Maison, P., Monet, R. and Grasselly, G. (1982) Resistance au puceron vert du pêcher Myzus
persicae Sulzer (Homoptera aphididae) chez Prunus persicae (L.) Batsch et d’autres espèces de Prunus.
Agronomie 2, 63–70.
Mishich, P., Todorovich, R., Zec, G., Misic, P. and Todorovic, R. (1998) Breeding peach cultivars for fresh use
and canning. Rasteniev dni Nauki 35, 575–577.
Monastra, F., Proto, D., Fideghelli, C., Grassi, G., Della Strada, G., Magliano, V. and Pennone, F. (1984)
Cultivars for processing: agronomic and technological data sheets. Clingstone peaches. L’Informatore
Agrario 40, 51–86.
Monet, R. and Massonié, G. (1994) Déterminisme génétique de la résistance au puceron vert (Myzus persicae)
chez le pêcher. Résultats complémentaires. Agronomie 2, 177–182.
Moore, J.N. (1985) Four new fruit cultivars for the South. HortScience 20, 651.
Nakasu, B., Bassols, M. and Feliciano, A. (1981) Temperate fruit breeding in Brazil. Fruit Varieties Journal 35,
114–122.
O’Brien, M., Fridley, R.B. and Claypool, L.L. (1978) Food losses in harvest and handling systems for fruits and
vegetables. Transactions of the ASAE 21, 386–390.
Okie, W.R. (1998) Handbook of Peach and Nectarine Varieties. US Department of Agriculture Agricultural
Handbook No. 714. US Government Printing Office, Washington, DC.
Pascal, T., Kervella, J., Pfeiffer, F.G., Sauge, M.H. and Esmenjaud, D. (1997) Evaluation of the interspecific
progeny Prunus persica cv. Summergrand × Prunus davidiana for disease resistance and some agronom-
ic features. Acta Horticulturae 465, 185–191.
Paunovic, S.A., Paunovic, A.S. and Fideghelli, C. (1996) Investigation of peach germplasm (Prunus persica sp.
vulgaris = vineyard peach) in situ in Yugoslavia. Acta Horticulturae 374, 201–207.
Pressey, R. and Avants, J.K. (1978) Difference in polygalacturonase composition of clingstone and freestone
peaches. Journal of Food Science 43, 1415–1417.
Raseira, M.C.B., Nakasu, B.H., Fortes, J. and Martins, O.M. (1998) ‘Leonense’, a dual purpose peach cultivar,
adapted to southern Brazil. Fruit Varieties Journal 52, 89–91.
192 T.M. Gradziel and J.P. McCaa

Ray, L. and Morris, J.R. (1974) Post-harvest behavior of once-over harvested peaches. Arkansas Farm Research
23, 2.
Robertson, J.A., Meredith, F.I., Lyon, B.G., Chapman, G.W. and Sherman, W.B. (1992) Ripening and cold stor-
age changes in the quality characteristics of nonmelting clingstone peaches (FLA 9-20C). Journal of Food
Science 57, 462–465.
Rotaru, G. (1994) Comparative anatomy of the pericarp in the fruit of plum, apricot and peach. Buletinul
Academiei de Stiinte a Republicii Moldova. Stiinte Biologice si Chimice 3, 11–18.
Scorza, R. and Lightner, G.W. (1989) The pillar peach tree and growth habit, analysis of compact × pillar
progeny. Journal of American Society for Horticultural Science 114, 33–38.
Scorza, R. and Sherman, W.B. (1996) Peaches. In: Janick, J. and Moore, J.N. (eds) Fruit Breeding. Vol. 1. Tree
and Tropical Fruits. Wiley, New York, pp. 325–440.
Sistrunk, W.A., Rom, R.C., Moore, J.N., Junek, J. and Brown, S.A. (1979) Quality parameters for evaluating
clingstone peach selections. Arkansas Farm Research 28, 2.
Souty, M. (1972) Vitamin C in peaches for canning. Qualitas Plantarum et Materiae Vegetabiles 21, 223–228.
Todorovic, R.R., Misic, P.D., Zec, G.N. and Monet, R. (1998) Twenty five years of peach breeding in the
research institute INI ‘PKB Agroekonomik’, Beograd. Acta Horticulturae 465, 137–140.
Tourjee, K.R., Barrett, D.M., Romero, M.V. and Gradziel, T.M. (1998) Measuring flesh color variability among
processing clingstone peach genotypes differing in carotenoid composition. Journal of the American
Society for Horticultural Science 123, 433–437.
USDA (2004) USDA – National Agricultural Statistics Service, Noncitrus Fruits and Nuts. http://ffas.usda.gov/
htp/Hort_Circular/2006/01-6/canned%20peach%20feature%20Nov%202005.pdf (accessed November
2004).
Van Der Heyden, C.R., Holford, P. and Richards, G.D. (1997) A new source of peach germplasm containing
semi-freestone nonmelting flesh types. HortScience 32, 288–289.
Wang, T., Gonzalez, A.R., Gbur, E.E. and Aselage, J.M. (1993) Organic acid changes during ripening of pro-
cessing peaches. Journal of Food Science 58, 631–632.
Wilson, P.W. and Boudreaux, J.E. (1986) Developing a dual purpose peach in Louisiana. Fruit Varieties Journal
40, 93–96.
Zhang, G.R., Zong, X.P., Shen, Y.S., Wang, Z.Q., Zuo, Q.Y. and Zhu, G.R. (1996) Zhenghuang 5, a new late
canning peach cultivar. Journal of Fruit Science 13, 130–131.
Zocca, A. (1975) Further trials on the mechanical harvesting of clingstone peaches. In: Atti Incontro Frutticolo
su Raccolta Meccanica e Sistemi di Allevamento per la Frutticoltura da Industria. Società Orticola Italiana,
Bologna, Italy, pp. 53–56.
Zocca, A. and Fridley, R.B. (1977) Mechanical harvesting of clingstone peaches. Journal of Agricultural Engi-
neering Research 22, 247–257.
 8 Rootstock Development

Gregory L. Reighard1 and Filiberto Loreti2

1Department of Horticulture, Clemson University, Clemson, South Carolina, USA
2Department of Fruit Science and Crop Protection, University of Pisa, Pisa, Italy

8.1 Introduction 193

8.2 Selection Criteria for Peach Rootstocks 194
Graft compatibility 194
Ease of sexual or asexual propagation 195
Resistance to abiotic stresses 195
Resistance to soil pests and diseases 196
Increased yield and fruit quality 198
Tree size control and anchorage 199
8.3 Characteristics of Peach Rootstocks 200
Section Euamygdalus species cultivars 200
Sections Euprunus and Microcerasus species cultivars 205
Interspecific hybrid cultivars 207
8.4 Peach Rootstock Breeding Programmes 211
Research institutions and objectives 212
Rootstocks in development 214
8.5 Outlook 215

8.1 Introduction
New rootstock cultivars possessing diverse
horticultural traits for stone fruit crops includ-
ing peach (Prunus persica (L) Batsch) are being
developed by both public and private pro-
grammes for testing and release to fruit grow-
ers worldwide (Renaud et al., 1988; Felipe et al.,
1997; Loreti, 1997; Loreti and Massai, 2002a;
Reighard, 2002; Moreno, 2004). Rootstocks
provide a cultural tool for peach growers to
increase productivity and improve efficiency
via better tree survival, controlled tree vigour
and increased fruit size, yield and quality. Thus,
the choice of rootstock becomes as economically
important as the scion cultivar whenever peach
trees must be grown on soils having high bulk
density, coarse texture (sand), parasitic nema-
todes, root rot fungal pathogens, high pH or
other orchard replant problems. If one or
more of these conditions are present, peach
tree survival and growth can be improved
significantly by selecting the appropriate
rootstock for each soil or site situation.
Historically, peach cultivars that were
grown in Australia, Brazil, Canada, Chile,

© CAB International 2008. The Peach: Botany, Production and Uses

(eds D.R. Layne and D. Bassi) 193
194 G.L. Reighard and F. Loreti
China, France, Greece, Italy, Mexico, New
Zealand, South Africa, Spain and the USA
had been grafted to peach seedling rootstocks
that inherently lacked traits such as scion
dwarfing and tolerance to soil waterlogging,
drought, calcareous soils, ectoparasitic nema-
todes, crown gall (Agrobacterium tumefaciens),
soil-borne fungal pathogens and orchard
replant problems (Layne, 1987). However, as
production costs increased and chemical con-
trol practices became cost-prohibitive or
unavailable, new interspecific Prunus L. root-
stocks were developed to overcome these
biotic and abiotic soil obstacles, which had
been simply circumvented in years past by
orchard relocation, tile drainage or chemical
fumigation (Loreti, 1984, 1997). This chapter
discusses traditional and new rootstock culti-
vars for peach and their potential for solving
some of the specific soil and site problems
that peach growers are experiencing world-
wide. In addition, selection criteria, horticul-
tural characteristics and breeding programmes
for these rootstocks are discussed.
8.2 Selection Criteria for Peach
Rootstocks
Graft compatibility
Peach is partially to completely graft-compati-
ble with several species within its taxonomic
Section Euamygdalus Schneid, which includes
P. persica, Prunus dulcis (Mill.) D.A. Webb and
Prunus davidiana (Carr.) Franch. Peach and espe-
cially nectarine are much less graft-compatible
with other Prunus spp. from Sections Euprunus,
Prunocerasus and Microcerasus. Breeding new
rootstocks for peach from intra- and interspe-
cific crosses requires extensive nursery- and
field-testing of numerous budded peach and
nectarine scion cultivars to ascertain good
graft compatibility for tree health and survival
under normal orchard conditions (Yamaguchi
et al., 2004; Zarrouk et al., 2006). Thus, breed-
ers must understand the types and potential
causes of incompatibility between different
rootstock and scion genotypes.
Peach is compatible with itself and
peach × almond hybrids. Compatibility is
much less with almond (P. dulcis), but some
almond genotypes have been selected for
compatibility with peach for use as peach
rootstocks in Hungary, Moldova, Egypt and
Algeria (J. Anderson, Utah, 2005, personal
communication; F. Loreti, unpublished
results; Z. Szabo, Hungary, 2005, personal
communication). Peach is also compatible
with P. davidiana and its hybrids with peach.
Other peach-like species such as Prunus mira
Koehne, Prunus ferganensis (Kostov & Rjabov)
Kovalev & Kostov and Prunus kansuensis Reh-
der have been successfully used as rootstocks
for peach in China (Wang et al., 2002).
Peach has been budded with many spe-
cies from Section Euprunus. Compatibility has
been good with some rootstock selections and
cultivars from Prunus insititia L. (damson
plums), Prunus spinosa L. (sloe plums), Pru-
nus domestica L. (European plums), Prunus
salicina Lindl. (Japanese plums) and Prunus
cerasifera Ehrh. (Myrobalan or cherry plums).
Myrobalan plums are often more compatible
when they are first hybridized with other
plums. Salesses and Bonnet (1992) reported
the existence of two types of genetic incom-
patibility of the Myrobalan and sloe plums
with peach. In the case of ‘Damas GF 1869’ (a
pentaploid rootstock, probably P. domestica ×
P. spinosa), at least two dominant alleles are
responsible for the incompatibility of peach
cultivars grafted on this rootstock (Salesses
and Alkai, 1985). However, this is not the case
for Myrobalan, which may have another type
of genetic control. Incompatibility between
Myrobalan and peach appeared to be a trans-
located type, which is evident by abnormal
scion behaviour such as leaf yellowing and
reduction in vigour. Translocated incompati-
bility appeared to result from impaired
phloem transport from the shoots to the roots
due to the degeneration and/or decrease in
the number of sieve tubes at the graft union,
which reduced carbohydrate transport to the
roots and eventually starved them after 1 or 2
years (Moing and Gaudillere, 1992; Moreno
et al., 1994a). McClintock (1948) also observed
this for peach cultivars budded to ‘Marianna
2624’, a P. cerasifera hybrid. However, incom-
patibility between sloes and peach appeared
to be localized by having weak graft unions
characterized by necrosis and absence of
 Rootstock Development
lignified tissues in the graft union (Salesses
et al., 1988).
Peach has been propagated on several
species from Sections Prunocerasus and Micro-
cerasus with limited success. The best examples
are some commercially available selections of
Prunus americana Marshall (e.g. Bailey Nurs-
eries, Newport, Minnesota) and Prunus pumila
L. (‘Pumiselect®’) that appear to be somewhat
to very compatible with many peach culti-
vars. Some other species of these sections that
have been tried as peach rootstocks or inter-
stocks are Prunus subcordata Benth. (Roberts
and Westwood, 1981), Prunus angustifolia
Marshall and Prunus hortulana L.H. Bailey
(Johnson, 1938; Graham, 2002), Prunus besseyi
L.H. Bailey (Funt and Goulart, 1981; Indreias
et al., 2004), Prunus japonica Thunb. and Prunus
tomentosa Thunb. (Funt and Goulart, 1981; Wang
et al., 2002). All of these species tend to dwarf
peach scion cultivars.
Ease of sexual or asexual propagation
All rootstocks developed for commercial release
should be nursery- or micropropagation-
friendly to allow for adequate production of
trees to sell at a market-driven price. The past
use of peach seedling rootstocks for peach by
the nursery industry was partially driven by
the availability of inexpensive seed from can-
nery cultivars, the ease of seedling propaga-
tion and budding, and the lack of proven
substitute cultivars, both seedling and clonal.
The last half of the 20th century brought an
increase in rootstock breeding and selection,
resulting in nematode-resistant peach seedling
rootstocks and Prunus L. hybrids resistant to
nematodes in addition to tolerance to differ-
ent soil textures and chemistries (Pinochet
et al., 1999, 2002).
New clonal hybrid rootstocks necessitated
technological innovations before propagators
could generate sufficient liners for tree fruit
nurseries. Propagation methods for hardwood
and softwood cuttings were refined and then
replaced by tissue culture and micropropaga-
tion systems to mass-produce thousands of
select single genotypes of P. insititia and P.
domestica as well as hybrid Prunus L. rootstocks
195
for peach (Zuccherelli, 1979; Martinelli, 1985;
Fiorino and Loreti, 1987; Andreu and Marin,
2004). One of the first examples was ‘GF 677’,
which progressed from propagation by root-
ing softwood cuttings under mist to tissue
culture of explants for the micropropagation
of millions of transplants annually (Loreti
and Morini, 1983). Therefore, fruit tree root-
stocks should be screened first for ease of
propagation by either seed or asexual propa-
gation, before they are released to commercial
fruit nurseries and the growers.
Resistance to abiotic stresses
Calcareous and high bulk density soils
Peach seedling rootstocks are not adapted to
poorly drained, heavy clay soil or to calcare-
ous soil where pH is above 7.5. When peach
cultivars budded to peach rootstocks are
planted in high pH or alkaline soil, trees become
weak, unproductive and iron-deficient. A
number of new hybrid rootstocks developed
in Europe were selected for these calcareous
conditions. These include: (i) the French root-
stocks ‘Jaspi®’ ((P. domestica × P. salicina) × P.
spinosa L.), ‘Julior®’ (P. insititia × P. domestica),
‘GF 677’ (a natural peach–almond hybrid)
and ‘Cadaman’ (P. persica × P. davidiana); (ii)
the Italian rootstocks ‘Barrier 1’ (P. persica × P.
davidiana), ‘Mr.S. 2/5’ (P. cerasifera × P.
spinosa?), ‘Mr.S. 2/8’ (P. cerasifera) and ‘Sirio’
(P. persica × P. dulcis); and (iii) the Spanish
rootstocks ‘Adesoto 101’ (P. insititia), ‘Montizo’
(P. insititia) and the peach–almond hybrids
‘Adafuel’, ‘Adarcias’, ‘Garnem’, ‘Felinem’
and ‘Monegro’ (Cambra, 1990; Moreno et al.,
1994b, 1995a; Felipe et al., 1997; Loreti, 1997).
Compatibility with peach is fair to excellent
with these rootstocks except for ‘Jaspi’ (Loreti
and Massai, 2002a), and high pH problems
are lessened when these rootstocks are used
instead of peach seedlings for many peach
and some nectarine cultivars.
Waterlogging
Peach grown on peach seedling rootstocks
and planted in poorly drained or seasonally
waterlogged soils eventually declines or dies.
196 G.L. Reighard and F. Loreti
Many European Prunus L. rootstocks devel-
oped in the past 20 years are listed as tolerant
of waterlogging. Rootstocks labelled as toler-
ant to waterlogged soils and ‘compatible’ with
peach include ‘Jaspi®’, ‘Julior®’, ‘Penta’ (P.
domestica), ‘Tetra’ (P. domestica), ‘Mr.S. 2/5’,
‘Barrier 1’, ‘Adesoto 101’, ‘Montizo’ and
‘Krymsk® 1’ (P. tomentosa × P. cerasifera). In
South Africa, ‘SAPO 778’, which is ‘Siberian
C’ peach crossed with a plum–almond hybrid
by Zaiger Genetics, Inc. (Modesto, Califor-
nia), is also moderately tolerant of wet soils
(Lötze, 1997). Many of these rootstocks were
developed in Mediterranean climates that
receive their rainfall in the winter. However,
in other peach-growing areas of the world
such as eastern North America, waterlogging
can occur during the growing season. There-
fore, in regions outside where these rootstocks
were developed, it is uncertain whether these
rootstocks are tolerant of wet soil conditions
in both the dormant season and the growing
season. Only future testing will answer these
questions.
Winter temperatures
Winter cold hardiness of peach root systems
varies considerably among rootstock culti-
vars. The absence of snow cover and some
orchard floor management practices can
increase the susceptibility of peach seedling
rootstocks to cold injury. Rootstocks that are
inherently cold-hardy or de-acclimatize at a
slower rate after warm temperatures are nec-
essary to grow peaches in cold regions. Many
of the available cold-tolerant peach rootstocks
originated from a now defunct Canadian
breeding programme (Layne, 1987) and more
recently from the Vavilov Research Institute
of Plant Industry (VRI) near Krymsk, Russia.
Releases of the cold-hardy peach seedling
rootstocks ‘Siberian C’, ‘Harrow Blood’, ‘Tzim
Pee Tao’ and ‘Chui Lum Tao’ have improved
cold hardiness of peach scion cultivars
(Weaver, 1967a,b; Layne and Jui, 1994) but
have had some other deficiencies such as sus-
ceptibility to nematodes or fungal root rots.
Therefore, their commercial utility is limited.
The recent introduction of three Russian
rootstocks, ‘Krymsk® 1’, ‘Krymsk® 2’ (P. incana
(Pall.) Batsch × P. tomentosa) and ‘Krymsk®
86’ (P. cerasifera × P. persica), may offer more
cold hardiness than the Canadian peach root-
stocks since they were developed from Prunus
species from colder regions that are marginal
for peach production due to very cold winters
and spring freezes. Another option may be
selected clones of P. americana, which is cold-
hardy. Currently, open-pollinated seedlings
from P. americana genotypes selected for com-
patibility with peach are being used as peach
rootstocks by several mail order nurseries for
the fruit hobbyist market in the northern USA.
Cool soil temperatures up to 60 days
after peach bloom in low-chill areas of Aus-
tralia are thought to be responsible for ‘spring
shock syndrome’, which significantly reduces
yields and is influenced by rootstock (Malcolm
et al., 1999). Low-chill adapted rootstocks
such as ‘Okinawa’ and ‘Flordaguard’ as well
as other cultivars and selections are being
tested in low-chill production areas of Aus-
tralia to ameliorate this syndrome.
Resistance to soil pests and diseases
Parasitic nematodes
Many nematode species successfully parasit-
ize peach roots and reduce peach tree growth
and productivity. Four kinds of nematodes
are recognized as harmful to peach tree roots
(Nyczepir and Becker, 1998). They are the
ring (Mesocriconema xenoplax (Raski) Loof &
de Grisse), root-knot (Meloidogyne incognita
(Kofoid & White) Chitwood, Meloidogyne
javanica (Treub) Chitwood, Meloidogyne are-
naria (Neal) Chitwood, Meloidogyne floridensis
Handoo et al. and Meloidogyne hapla Chit-
wood), root-lesion (Pratylenchus vulnus Allen
& Jensen and Pratylenchus penetrans (Cobb)
Chitwood & Oteifa) and dagger (Xiphinema
americanum Cobb) nematodes. Rootstocks
often are categorized as immune, resistant,
tolerant or susceptible to nematodes. Tolerant
rootstocks are fair to good hosts for a specific
nematode, but nematode reproduction and
feeding does not significantly alter the root-
stock’s ability to supply the scion’s mineral,
hormonal and water requirements to survive,
grow and bear fruit. Rootstocks susceptible
to a specific nematode are good hosts for
 Rootstock Development
nematode reproduction and are negatively
affected by nematode feeding in terms of tree
survival, growth and fruiting (Nyczepir and
Becker, 1998).
Ring nematode, an ectoparasitic nema-
tode, has been linked directly to both the
onset of peach tree short life syndrome (PTSL)
in the south-eastern USA (Zehr et al., 1976;
Nyczepir et al., 1983) and the bacterial canker
complex in California (Davis and English,
1969; McKenry, 1989). Many different peach
rootstocks including the new hybrid releases
have been tested for reaction to ring nema-
tode. However, most were good hosts for ring
nematode (Westcott et al., 1994) and were sus-
ceptible to PTSL (Okie et al., 1994a). Only one
rootstock, the peach seedling ‘Guardian®’,
has had acceptable survival in field tests in
South Carolina and Georgia (Okie et al., 1994b;
Beckman et al., 1997a; Reighard et al., 1997)
even though it is a good host for ring nema-
tode (Nyczepir et al., 1992). In California,
‘Lovell’ peach seedlings that are sold as clones
from Duarte Nursery (Hughson, California)
have shown a level of ring protection that is
similar or slightly superior to ‘Guardian®’,
although ‘Guardian®’ provides superior root-
knot nematode protection over ‘Lovell’ (M.
McHenry, California, 2006, personal commu-
nication). However, finding resistance to ring
nematode in a Prunus L. rootstock that is also
compatible with peach has been unsuccessful
thus far.
Root-knot nematodes significantly reduce
peach tree growth. There are five known spe-
cies of root-knot nematode (M. arenaria, M.
incognita, M. javanica, M. floridensis, M. hapla)
as well as a number of races within each spe-
cies that feed on peach. M. incognita and M.
javanica are the most common on peach in the
USA. Many peach rootstocks were introduced
to the USA in the 20th century because of their
root-knot nematode resistance (Day, 1953).
These included ‘Shalil’, ‘Yunnan’ and ‘Oki-
nawa’. These rootstocks either were not resis-
tant to M. javanica or had other problems and
were eventually replaced via selective breed-
ing, which led to the root-knot resistant
rootstocks ‘Nemaguard’, ‘Nemared’, ‘Florda-
guard’ and ‘Guardian®’ (Brooks and Olmo,
1961; Ramming and Tanner, 1983; Sherman
et al., 1991; Okie et al., 1994b).
197
During this same time, in other peach-
growing countries, breeders also selected for
root-knot resistance. Plum rootstocks tended to
be immune to Meloidogyne spp., while other
rootstocks tended to have different levels of
resistance. Root-knot resistant rootstocks relea-
sed include the Italian rootstocks ‘Barrier 1’,
‘Penta’ and ‘Tetra’; the Spanish rootstocks
‘Adesoto 101’, ‘Garnem’, ‘Felinem’ and ‘Mone-
gro’; and the French rootstocks ‘Myran®’ (a
cross of ‘Belsiana’ (P. cerasifera × P. salicina) and
‘Yunnan’ peach) and ‘Ishtara®’ (a cross of ‘Bel-
siana’ and a (P. persica × P. cerasifera) hybrid)
(Fernández et al., 1994; Moreno et al., 1995a;
Esmenjaud et al., 1997; Felipe et al., 1997, Nicotra
and Moser, 1997; Gomez-Aparisi et al., 2001).
Root-lesion (P. vulnus and P. penetrans)
and dagger (X. americanum) nematodes are
two other ectoparasitic nematodes that feed
on and damage peach roots. Root-lesion nem-
atodes can reduce tree growth and fruit pro-
duction if not controlled. P. vulnus is a problem
in the southern USA and California, while P.
penetrans occurs in northern areas. Peach root-
stocks listed as tolerant to P. vulnus in Europe
are ‘Rubira®’, ‘GF 305’, ‘Penta’, ‘Tetra’ and
‘P.S.B2’ (P. persica) (Alcañiz et al., 1996). Some
new rootstocks showing P. vulnus tolerance in
greenhouse studies in Spain are ‘Krymsk® 86’
and to a lesser extent ‘Krymsk® 1’ and
‘Krymsk® 2’ (Pinochet et al., 2000). These same
three rootstocks in California have shown some
resistance to root-lesion nematode in the
greenhouse and/or 2-year-old field studies,
but they have been susceptible to root-knot
nematodes (M. McKenry, California, 2006,
personal communication). In Canada, testing
of ‘GF 305’ by McFadden-Smith et al. (1998)
showed that this rootstock was quite suscep-
tible to P. penetrans and that the Canadian
peach seedling rootstock ‘Chui Lum Tao’ was
more tolerant in greenhouse studies. In addi-
tion, ‘Bailey’, ‘Higama®’ and ‘Guardian®’
were less susceptible to P. penetrans than many
of the European rootstocks tested, while in
California the level of susceptibility in ‘Guard-
ian®’ to P. vulnus was similar to ‘Nemaguard’
(M. McKenry, California, 2006, personal com-
munication). However, multiple nematode
species and races create a significant obstacle
to finding a broadly adapted, nematode-
resistant rootstock.
198 G.L. Reighard and F. Loreti
Dagger nematodes damage peach trees
by serving as the vector for Tomato ringspot
virus, which causes stem pitting. Since many
weed species such as dandelions (Taraxacum
officinale Weber) are hosts for this virus, dag-
ger nematode resistance or virus tolerance in
rootstocks is probably the best way to prevent
infection and/or tree injury. Dagger nema-
tode species are vectors for other nepoviruses
throughout the world. Peach seedling root-
stocks are not resistant to dagger nematodes,
and therefore, non-P. persica rootstocks need
to be evaluated for resistance to the nematode
or the virus. Some cherry plum P. cerasifera
genotypes are less sensitive to Tomato ringspot
virus (Hoy and Mircetich, 1984; Halbrendt et al.,
1994). Therefore, rootstocks such as the clonal
rootstocks ‘Mr.S. 2/5’ and ‘Mr.S. 2/8’ from
Italy, ‘Krymsk® 1’ from Russia and ‘Adara’ (P.
cerasifera) from Spain may offer some resis-
tance, but they still remain untested for these
viruses, and ‘Adara’ has limited compatibil-
ity with peach (Moreno et al., 1995b).
Pathogenic soil fungi
On fine-textured, high bulk density or poorly
drained soils, peach seedling rootstocks are at
risk of becoming infected with Phytophthora
spp. de Bary, which causes crown rot (Elena
and Tsipouridis, 2000). Similarly, all root-
stocks of P. persica are susceptible to the oak
root rot fungus (Armillaria mellea (Vahl:Fr.)
P. Kumm. and Armillaria tabescens (Scop.)
Dennis et al.) regardless of soil texture or
drainage. Both fungi are difficult to control or
eradicate; therefore, genetic resistance to them
is highly desirable in rootstocks.
Many hybrid rootstocks released by
European breeders are listed as tolerant to
replant sites and soil diseases. However, spe-
cific soil diseases usually are not identified.
‘Ishtara®’ and ‘Myran®’ were reported by
Renaud et al. (1988) to be resistant or tolerant
to oak root rot (A. mellea) in France, but Beck-
man and Pusey (2001) in the south-eastern
USA found these rootstocks were susceptible
to A. tabescens. This suggests that after these
exported rootstocks emerge from quarantine
facilities in regions outside Europe, they will
still require additional field-testing to evalu-
ate traits such as ‘disease resistance’ before
they are commercially produced in large num-
bers for local growers.
Increased yield and fruit quality
Numerous studies have found that some
peach rootstocks increase yield, fruit size and
fruit quality of commercial peach cultivars
(Layne, 1994; Moreno et al., 1994b; Guidoni
et al., 1998; Loreti and Massai, 2002b; Massai
and Loreti, 2004; Reighard et al., 2004). Under
conventional training systems, yield increases
reported for selected rootstocks have been
primarily due either to increased tree vigour
and precocity or to increased tree survival
and longevity. Fruit quality improvements
such as size, skin colour and soluble solids
content (SSC) have largely been observed
from peach cultivars on plum and interspe-
cific hybrid rootstocks (De Salvador et al.,
2002; G. Reighard, unpublished results).
On typical peach orchard sites (i.e. well-
drained soils), peach seedling rootstocks have
performed as well as any peach-compatible
rootstocks composed of other Prunus L. spe-
cies or interspecific hybrids. Long-term test-
ing (i.e. NC-140 project) of primarily peach
seedling rootstocks throughout North Amer-
ica (Perry et al., 2000; Reighard et al., 2004)
showed that rootstocks that induced the high-
est scion productivity were those that induced
the best scion growth or survival at individual
locations. In a former NC-140 test, ‘Redhaven’
on all five of the peach seedling rootstocks
plus ‘GF 677’, a peach–almond hybrid, grew
the largest and had the highest yields com-
pared with several interspecific plum hybrid
rootstocks (Perry et al., 2000). Although ‘GF
677’ was the most vigorous rootstock, it also
had the lowest yield efficiency compared
with the peach seedling rootstocks.
In another 20-location NC-140 test,
‘Redhaven’ on peach seedling rootstocks
‘Lovell’, ‘GF 305’, ‘Montclar®’ and ‘Guard-
ian®’ had tree heights and crown widths that
were significantly greater than on 15 other
rootstocks (Reighard et al., 2004). ‘Redhaven’
cumulative fruit yields were also highest on
these four rootstocks. Fruit production appea-
red to be directly affected by the number of
bearing shoots supported by the canopy.
 Rootstock Development
However, cumulative yield efficiency (kg/cm2
TCSA) was highest on the lower-yielding but
slightly less vigorous peach rootstocks ‘Bai-
ley’ and ‘Tennessee Natural 281-1’.
On problem orchard sites, rootstocks that
impart a survival advantage to peach cultivars
generally will out-yield precocious and vigor-
ous rootstocks that do not survive or grow
well due to negative biotic or abiotic factors.
Examples of these problems are calcareous
soils, fine-textured soils, drought, pathogenic
fungi, bacteria and nematodes, cold climates
and replant sites. In many of these situations,
peach seedling rootstocks are often not the best
option. Thus, testing and selection of appro-
priate rootstocks for peach growth and sur-
vival are necessary for each production area.
Data from the NC-140 peach trials suggested
that rootstock cultivar productivity, in relation
to all rootstocks tested, tended to be consistent
across time. Therefore, since relative rankings
of the rootstock cultivar yields changed little
during the prime bearing years, the yield
potential for a rootstock compared with other
rootstocks at each test location could be deter-
mined as soon as the trees reached full produc-
tion at approximately 4 or 5 years.
Fruit weight can sometimes be signifi-
cantly influenced by rootstock, but cultural
(pruning and thinning) and environmental
factors (water and sunlight) often obscure or
negate these rootstock-imparted differences,
especially on peach seedling rootstocks. Hybrid
rootstocks such as ‘Ishtara®’ (semi-dwarf) and
‘Myran®’ (vigorous) have typically produced
larger and smaller fruit, respectively, in the
NC-140 trials in the USA (G. Reighard, per-
sonal observation). However, these differences
have been very small (<10 g). Other rootstock
trials ongoing and in progress have reported
increased fruit size with semi-dwarf and dwar-
fing hybrid rootstocks (R.S. Johnson, California,
2006, personal communication). Dwarf-type
rootstocks require high-density planting sys-
tems; and therefore different parameters (i.e.
spacing, canopy volume) are used to measure
yield and production efficiency.
Besides fruit size, data documenting
other effects induced by the rootstock on fruit
quality such as SSC and skin colour are limited
(Albás et al., 2004). Smaller canopy size due to
size-controlling rootstocks may increase
199
sunlight penetration and thus indirectly
improve fruit skin coloration and SSC. Dwarf-
ing rootstocks such as ‘Krymsk® 1’, ‘Krymsk®
2’ and ‘Mr.S. 2/5’ have increased fruit SSC in
some peach cultivars in South Carolina (G.
Reighard, unpublished results). In Italy,
‘Ishtara®’, ‘Mr.S. 2/5’ and ‘Barrier 1’ increased
fruit size and SSC (De Salvador and Monas-
tra, 1996; Remorini et al., 2005).
Tree size control and anchorage
Peach seedling rootstocks including brachytic
dwarfs rarely reduce scion vigour by more than
10–15%. Until the past 10–20 years, size control
in peaches through the use of rootstocks of other
Prunus species has not been achieved satisfacto-
rily owing to incompatibility or poor tree vigour.
Without graft-compatible, size-controlling root-
stocks such as in apple, increases in peach
orchard productivity via intensive training sys-
tems are probably unattainable. Fortunately,
new dwarfing rootstocks for peaches are being
selected for reduced vigour, graft compatibility
and sustained fruit production without reduc-
tions in fruit size and quality.
European rootstocks considered mildly
dwarfing (approximate percentage of peach
standard) include ‘Ishtara®’ (70%), ‘Julior®’
(70%), ‘Rubira®’ (90%), ‘Tetra’ (90%), ‘Mr.S.
2/5’ (90%), ‘Adara’ (80%), ‘Adarcias’ (70%)
and the Italian peach seedlings ‘P.S.A5’ (80%)
and ‘P.S.B2’ (90%). Semi-dwarfing rootstocks
are ‘Pumiselect®’ (60%), a P. pumila L. selection
developed in Germany (Jacob, 1992), ‘Sirio’
(60%) (Loreti and Massai, 1998), ‘Adesoto 101’
(60–80%) and ‘Krymsk® 1’ (formerly ‘VVA-1’)
(50%) and ‘Krymsk® 2’ (formerly ‘VSV-1’)
(40%) (Devyatov, 1996). In addition, in Cali-
fornia a new release ‘Controller 5’ (formerly
‘K-146-43’) for peach (DeJong et al., 2004a,
2005) is now in advanced testing and has
reportedly maintained yield efficiency and
fruit size despite significant tree dwarfing.
Other dwarfing rootstocks for peach include
the use of P. japonica and P. tomentosa in Japan
(Mizutani et al., 1985; Murase et al., 1986),
although they may be too dwarfing, and new
selections coming from the breeding pro-
grammes in Italy (e.g. ‘I.S.’ series, Pisa) and
Spain (CSIC, Zaragoza). The degree of
200 G.L. Reighard and F. Loreti
dwarfing of these rootstocks will vary with
the climate, soil type and site history. There-
fore, until there is extensive geographic test-
ing, it is unknown how these rootstocks will
perform as size-controlling rootstocks in other
peach production areas.
8.3 Characteristics of Peach Rootstocks
Several factors have recently prompted inter-
national research work to focus on the devel-
opment of new rootstocks suitable for the
needs of modern peach growing. These fac-
tors can be summarized as: (i) expansion of
peach growing into non-optimal and low-
chill areas; (ii) the need for rootstocks that can
reduce canopy vigour and increase planting
densities; (iii) the search for greater resistance
to diseases and soil or environmental prob-
lems; and (iv) as alternatives to use of soil
fumigants. Thus, the range of rootstocks now
available for peach worldwide has increased
dramatically over the last few years.
Traditionally, three groups can be identi-
fied, based on their genetic origin.
1. Euamygdalus species cultivars (P. persica,
Persica vulgaris).
2. Euprunus species cultivars (P. domestica,
P. insititia, P. cerasifera).
3. Interspecific hybrid cultivars (from con-
trolled or natural hybridization of a combina-
tion of Prunus species including P. persica, P.
dulcis, P. domestica, P. cerasifera, P. insititia, P.
davidiana, P. spinosa, P. salicina).
Until the last decade or so, only a few root-
stocks were commonly used in peach orchards.
Among those recently developed, some have
been found to perform well in some peach-
growing areas, while for other rootstocks the
results are still inconclusive (Table 8.1). Below
we summarize the most commonly known
rootstocks and those offering more interest-
ing prospects for the future.
Section Euamygdalus species cultivars
Open-pollinated peach seedlings are still the
most widely used rootstock for peach world-
wide. Nursery peach seed usually originates
from one of three sources: (i) local feral peach
trees or naturalized populations; (ii) commer-
cial processing or drying cultivars; or (iii)
peach rootstock cultivars planted in virus-
indexed seed orchards. Naturalized or wild-
type peaches continue to be used in many
countries, where in some cases they represent
the most frequently used rootstock for peach.
Seeds from P. persica are still used in some
countries of the Mediterranean basin, giving
rise to what is called the ‘Slavic seedling’,
which is often referred to as P. vulgaris P. Mill.
This rootstock, originating from the Balkan
peninsula, has small stones (pits) which pro-
duce a high yield of nursery seedlings. How-
ever, the problem with seeds from wild-type
peaches is their genetic variability and lack of
uniformity in the nursery and the orchard. In
addition, these peach seedlings can have viral
infection rates ranging between 5 and 10%.
Overall, feral peach seedlings – and the ‘Slavic
seedling’ in particular – can be considered
good rootstocks, but their primary disadvan-
tage compared with peach rootstock cultivars
is the tendency to produce very heteroge-
neous plants, due to the fact that commercial
nursery seeds are often derived from differ-
ent genetic sources (Loreti, 1984).
In the important peach-growing countries,
peach seedling rootstocks are now almost
exclusively cloned peach cultivars that are
maintained in virus-indexed orchards. A survey
recently conducted in these countries indi-
cated that peach seedling rootstocks are still
the most commonly used rootstocks for peach,
especially those obtained from commercial
cultivars (Loreti and Massai, 2006a). Some of
the more commonly used peach rootstock culti-
vars are ‘Halford’, ‘Lovell’, ‘Bailey’, ‘Siberian
C’, ‘Guardian®’, ‘Nemared’ and ‘Nemaguard’
in the USA and Canada; ‘Jerez’, ‘Jalacingo’
and ‘Tetela’ in Mexico; ‘Cuaresmillo’ in Argen-
tina; ‘Chucho Picudo’ in Chile; ‘Aldrighi’ and
‘Capdeboscq’ in Brazil; ‘Elberta’ and ‘Golden
Queen’ in Australia and New Zealand; ‘Ohatsu’
(‘Ohatsumomo’), ‘Tsukuba#1’ and ‘Tsukuba#4’
in Japan; ‘Baladi’ in Israel; ‘Mit Gharmr’ in
Egypt; ‘Missour’ in Algeria; ‘Halge’ in Iran;
‘Kakamas’ in South Africa; ‘I.D. 20/Ag1’ in
Greece; ‘De Balc’ in Romania; ‘B-VA-2’ and
‘Lesiberian’ in the Czech Republic; ‘CEPE’
and ‘C.2630’ in Hungary; and ‘Mandzurska’
Table 8.1. Commercial peach rootstock cultivars and their reported horticultural characteristics.

Nematode resistance
Rootstock Origin Species Vigour Cold Tolerance to Alkaline soil
cultivara (country)b codec ratingd hardinesse Mi/Mjf Ppvg Mxh wet soilsi tolerance

‘GF 305’ France 1 2 No 3 3 3 2 No

‘Montclar®’ France 1 2 No 3 3 3 2 Some
‘Rubira®’ France 1 3 No 3 2 2 2 Maybe
‘P.S.B2’ Italy 1 2 No 1 2 ? 2 No
‘Lovell’, ‘Halford’ USA 1 2 No 3 2 2 2 No

Rootstock Development
‘Nemaguard’ USA 1 1 No 1 2 3 2 No
‘Guardian®’ USA 1 1 No 1 2 2 2 No
‘Bailey’ USA 1 3 Yes 3 2 3 2 No
‘Siberian C’ Canada 1 3 Yes 3 3 3 3 No
‘GF 677’ France 2 1 No 3 3 3 3 Yes
‘Adafuel’ Spain 2 1 No 3 3 ? 3 Yes
‘Garnem’ Spain 2 1 No 1 3 3 3 Yes
‘Felinem’ Spain 2 1 No 1 2 ? 3 Yes
‘Adarcias’ Spain 2 2 No 3 3 ? 3 Yes
‘Sirio’ Italy 2 3 No 3 3 ? 3 Yes
‘Castore’, ‘Polluce’ Italy 2 2 No ? ? ? 3 Yes
‘Hansen 2168’, USA 2 1 No 1 2 3 3 Yes
‘Hansen 536’
‘Nickels’ USA 2 1 No 1 1 3 3 Yes
‘Penta’ Italy 3 1 No 1 2 2 1 Yes
‘Tetra’ Italy 3 2 No 1 3 3 1 Yes
‘Mr.S. 2/5’ Italy 4 3 No 1 3 3 1 Yes
‘Krymsk® 86’ Russia 5 2 Yes 3/1 1 ? 1 Yes

(Continued)

201
 202
Table 8.1. continued

Nematode resistance
Rootstock Origin Species Vigour Cold Tolerance to Alkaline soil
cultivara (country)b codec ratingd hardinesse Mi/Mjf Ppvg Mxh wet soilsi tolerance

‘Krymsk® 1’ Russia 6 3 Yes 3 2 3 1 Yes

‘Krymsk® 2’ Russia 7 4 Yes 3 2 ? 2 Some
‘Adesoto 101’ Spain 8 3 No 1 3 3 1 Yes
‘Montizo’ Spain 8 3 No 1 3 3 1 Yes
‘Julior®’ France 9 3 No 1 3 3 1 Some
‘Pumiselect®’ Germany 10 3 Yes 1 3 2 3 No

G.L. Reighard and F. Loreti

‘Barrier 1’ Italy 11 1 No 1 2 3 1 Yes
‘Cadaman®’ France 11 2 No 1 3 3 1 Yes
‘Ishtara®’ France 12 3 No 1 3 3 2 No
‘Myran®’ France 12 1 No 1 ? 3 1 No
‘Controller 5’ USA 13 4 No 3 3 3 3 No
‘Viking’ USA 14 1 No 1 1 2 1 Yes

aAdditional testing of compatibility with peach cultivars is advised for ‘Ishtara®’, ‘Krymsk® 1’ and ‘Krymsk® 2’. Excessive suckering may occur with ‘Adesoto 101’

and ‘Julior®’.
bCountry of origin and/or initial testing.
cSpecies type: 1, Prunus persica; 2, Prunus dulcis × P. persica; 3, Prunus domestica; 4, Prunus cerasifera; 5, P. cerasifera × P. persica; 6, Prunus tomentosa ×

P. cerasifera; 7, Prunus incana × P. tomentosa; 8, Prunus insititia; 9, P. insititia × P. domestica; 10, Prunus pumila; 11, P. persica × Prunus davidiana; 12,
(P. cerasifera × Prunus salicina) × P. persica; 13, P. salicina × P. persica; 14, unknown interspecific cross.
d1, vigour similar to ‘GF 677’ or ‘Nemaguard’; 2, vigour similar to ‘Lovell’; 3, vigour 5–25% less than ‘Lovell’; 4. vigour at least 30% less than ‘Lovell’.
eRootstock is considered to have better cold hardiness than ‘Lovell’.
fResistance to root-knot nematodes (Meloidogyne incognita, Meloidogyne javanica): 1, immune or resistant; 2, moderately resistant or some tolerance; 3,

susceptible; ?, unknown.
gResistance to root-lesion nematode (Pratylenchus penetrans or Pratylenchus vulnus): 1, immune or resistant; 2, moderately resistant or some tolerance; 3,

susceptible; ?, unknown.
hResistance to ring nematode (Mesocriconema xenoplax): 1, immune or resistant; 2, moderately resistant or some tolerance; 3, susceptible; ?, unknown.
iTolerance of fine-textured soils when waterlogged: 1, good; 2, fair; 3, poor.
 Rootstock Development
and ‘Rakoniewicha’ in Poland. The seedlings
obtained from these cultivars are preferable
because they are largely virus-free and more
genetically uniform than those from feral or
naturalized peach trees.
Among peach seedling rootstocks, culti-
vars showing certain distinct bioagronomic
characteristics and different degrees of sus-
ceptibility to biotic and abiotic stresses have
been selected in recent decades. Some of these
selections are now seldom used, such as ‘GF
305’, while others are gaining in popularity.
For instance, the ‘P.S.’ series from the Univer-
sity of Pisa, namely ‘P.S.A5’, ‘P.S.A7’ and
‘P.S.B2’ (Loreti and Massai, 2002a), and the
selections developed in France, such as ‘Mont-
clar®’ and to a much lesser extent ‘Rubira®’
and ‘Higama®’, are being planted where peach
seedling rootstocks grow well.
Peach rootstocks in the USA have almost
exclusively been and still are open-pollinated
seedlings of P. persica. Currently these peach
seedlings, derived primarily from the root-
stock cultivars ‘Lovell’, ‘Halford’, ‘Nema-
guard’, ‘Nemared’, ‘Bailey’ and ‘Guardian®’,
constitute greater than 95% of the peach root-
stocks used in the USA. Of these six root-
stocks, ‘Lovell’ and ‘Halford’ are thought to
be siblings, and ‘Nemared’ and ‘Guardian®’
have ‘Nemaguard’ in their pedigree. ‘Bailey’
is not related by parentage to the other five
seedling rootstocks. In field-testing, these
rootstocks differ in vigour, root-knot nema-
tode resistance and bacterial canker (i.e. PTSL)
tolerance, but have similar effects on many
other horticultural traits including economi-
cally important ones such as fruit size and
yield of scion cultivars. All of these peach
seedling rootstocks are susceptible to the
same soil diseases and conditions, which limit
their productivity and longevity in many oth-
erwise good production sites.
‘GF 305’
‘GF 305’ was selected at the National Institute
of Agronomic Research (INRA) in Pont-de-la-
Maye, France in 1945 for its vigorous growth
and uniformity in the nursery (Grasselly,
1983). This rootstock is compatible with all
peach and nectarine cultivars and has
good growth and productivity in the field.
203
However, ‘GF 305’ is very susceptible to
Agrobacterium, Phythophtora, root-knot and
root-lesion nematodes and some viruses, and
now is being planted less. It is still used by
virologists as a virus indicator in peach.
‘P.S.’ series
The ‘P.S.’ series originated at the Department
of Fruit Science and Crop Protection of Pisa
University (Scaramuzzi et al., 1976). The most
interesting selections were ‘P.S.A5’, ‘P.S.A6’,
‘P.S.A7’ and ‘P.S.B2’, which were selected for
their uniform growth and the different degree
of vigour induced in the scion. ‘P.S.A6’ is the
most vigorous (similar to Balkan peach seed-
lings) while ‘P.S.B2’, ‘P.S.A7’ and ‘P.S.A5’ are
10–15%, 15–20% and 20–25% less vigorous,
respectively, than ‘P.S.A6’ (Loreti and Massai,
1995; Pellegrino et al., 1997).
‘P.S.A5’ is a peach seedling rootstock that
induces less vigour in the scion than do other
peach seedlings and encourages uniform
growth, precocity, good crop efficiency and
improved fruit quality in scion cultivars
(Loreti and Massai, 1988). Furthermore, it has
a slightly higher resistance to waterlogging
than most other peach seedlings, while its
alkalinity tolerance is similar to other seed-
ling rootstocks. It is resistant to Verticillium
wilt (Cirulli et al., 2001). ‘P.S.A5’ is particu-
larly suitable for the free spindle or delayed
vase training system and is excellent for fer-
tile soils, vigorous or early-ripening cultivars,
and medium–high density planting systems.
‘P.S.A6’ is uniform and rapid growing in
the nursery and can be propagated via hard-
wood cuttings. It induces high vigour in
grafted trees, though slightly less than ‘GF
677’, and performs well in poor soils. Under
these conditions and without irrigation, it still
is highly productive by virtue of its well-
developed root system.
‘P.S.A7’ also induces very uniform vigour
and rapid growth of seedlings in the nursery.
Like other peach seedlings, it is not well adapted
to wet, heavy and poorly drained soils, but
appears better adapted to loam soils with
medium or high fertility. Furthermore, while
maintaining good productivity and fruit qual-
ity, it induces 15–20% less vigour in the scion
than does a standard seedling. This allows for
204 G.L. Reighard and F. Loreti
slightly narrower tree spacing in fertile soils
and with vigorous cultivars.
‘P.S.B2’ is characterized by uniform and
rapid growth of seedlings in the nursery and
can be propagated by hardwood cuttings, but
has some susceptibility to powdery mildew.
Resistance of ‘P.S.B2’ to root waterlogging
and calcareous soils is similar to that observed
in other peach seedlings. This rootstock is less
sensitive to replant problems, probably on
account of its greater resistance to P. vulnus. It
also performs well even in heavy soils, pro-
vided that waterlogging does not occur. ‘P.S.B2’
induces medium–high vigour in grafted trees,
but at least 10–15% less than Slavic seedlings,
as well as high productivity, early ripening
and good fruit quality.
‘Rubira®’
‘Rubira®’ was selected from peach seedlings
grown at INRA from a Californian seedlot
imported in 1960 and owes its name to its red-
coloured leaves. ‘Rubira®’ seedlings have uni-
form and vigorous growth in the nursery, but
are slightly susceptible to powdery mildew
(Loreti, 1984) and resistant to the green peach
aphid, Myzus persicae (Sulzer) (Massonie and
Paison, 1979). ‘Rubira®’ induces 15–20%
lower vigour than ‘GF 677’ and is more preco-
cious with good productivity. It is therefore
recommended for vigorous cultivars that are
slow to enter into production and for higher
planting densities. It is also less susceptible to
crown gall (A. tumefaciens) and P. vulnus than
other peach seedlings, but is susceptible to M.
incognita and M. arenaria nematodes (Loreti,
1984).
‘Higama®’
‘Higama®’ was selected from peach seedlings
grown from another seedlot imported in 1960
from Japan at INRA (Grasselly, 1983). This
rootstock induces high vigour in the scion
and is suitable for early-season cultivars. In
addition, ‘Higama®’ has a higher sensitivity
to iron chlorosis (J.-L. Poëssel, France, 2001,
personal communication) and crown gall than
other peach seedling rootstocks, but has a fair
degree of resistance to M. javanica and M. incog-
nita nematodes. However, it had relatively
low yields and survival in several trials (De
Salvador and Monastra, 1996; Reighard et al.,
2004).
‘Montclar® Chanturgue’
‘Montclar®’ was selected at INRA in 1960 and
has similar characteristics to ‘Higama®’ with
regard to high seed production, uniform
seedling growth and vigour in the nursery,
and increased vigour in scion cultivars (Gras-
selly, 1983). It differs from the other French
seedling rootstocks in exhibiting greater resis-
tance to iron-induced chlorosis and better
uptake of iron and magnesium from the soil.
It is a popular peach rootstock in Europe.
‘Nemaguard’
‘Nemaguard’ rootstock was selected from
seedlings from a seedlot received in 1949 by
the US Department of Agriculture (USDA),
which was labelled P. davidiana and was even-
tually released as ‘FV 234-1’ in 1959 (Brooks
and Olmo, 1961). Thought to be a putative
hybrid of P. persica × P. davidiana, field obser-
vations (Okie, 1998) and molecular studies
(Lu et al., 1996) indicate that it is primarily P.
persica. ‘Nemaguard’ seedlings are uniform
and vigorous, compatible with peach and
nectarine cultivars, and impart excellent scion
vigour and productivity. It has good resis-
tance to M. incognita, M. javanica and M. are-
naria, but recent research has confirmed that a
new root-knot species (M. floridensis) (Handoo
et al., 2004) can reproduce in the roots of
‘Nemaguard’ (Nyczepir and Beckman, 2000).
‘Nemaguard’ is fairly tolerant of crown gall,
but is sensitive to P. vulnus, fungal root rots,
Verticillium, iron chlorosis and root waterlog-
ging and may reduce winter hardiness of
scion cultivars in cold climates. ‘Nemaguard’
suckers extensively and is very sensitive to
ring nematode (M. xenoplax) feeding, which
leads to tree injury and death from bacterial
canker (Pseudomonas syringae pv. syringae van
Hall) and PTSL (Zehr et al., 1976; Nyczepir
et al., 1983). Despite these limitations, ‘Nema-
guard’ is one of the most widely planted stone
fruit rootstocks in California and South America.
For peach production, however, it performs
best on fumigated ring nematode-infested
 Rootstock Development
soils, virgin peach sites, and soils that have
root-knot nematode problems.
‘Nemared’
‘Nemared’ is a redleaf selection released in
1983 by the USDA that has ‘Nemaguard’ in its
lineage (Ramming and Tanner, 1983). ‘Nem-
ared’ in the nursery is similar to ‘Nemaguard’
but produces seedlings with less lateral branches
thus facilitating budding. Field characteristics
of ‘Nemared’ are also similar to ‘Nemaguard’,
except ‘Nemared’ produces a slightly more
vigorous tree with equal or better root-knot
resistance, but has increased susceptibility to
bacterial canker (M. McKenry, California,
2006, personal communication).
‘Guardian® BY520-9’
‘Guardian®’ rootstock traces its lineage back
four generations to a ‘Nemaguard’ cross in
1954 and was released in 1993 jointly by USDA
and Clemson University (Okie et al., 1994b).
‘Guardian®’ has many traits similar to ‘Nema-
guard’. The primary differences are that ‘Guard-
ian®’ has lower seed germination, slightly less
root-knot nematode resistance (Nyczepir et al.,
1999, 2006), supports fewer M. xenoplax, and
exhibits significantly higher tolerance to ring
nematode, bacterial canker and PTSL (Beck-
man et al., 1997a; Reighard et al., 1997). The
latter trait has made it the most popular root-
stock, especially for nematode-infested replant
sites in the south-eastern USA, even though it
is susceptible to Armillaria root rot.
‘Lovell’ and ‘Halford’
‘Lovell’ and ‘Halford’ are old drying and pro-
cessing peach cultivars that were selected as
seedlings from California orchards around
1882 and 1921, respectively (Okie, 1998). ‘Lovell’
has high seed germination of uniform seed-
lings that are compatible with all peach and
nectarine cultivars. Scion vigour is slightly
less than on ‘Nemaguard’. ‘Lovell’ does not
sucker and is susceptible to root-knot and
root-lesion nematodes, but has better toler-
ance to ring nematodes, bacterial canker and
PTSL than ‘Nemaguard’. ‘Lovell’ is suscepti-
ble to waterlogging, crown gall, Phytophthora
spp. and Armillaria spp. ‘Halford’ is similar to
205
‘Lovell’ in nursery and field characteristics
and is even thought to be a ‘Lovell’ seedling.
Since ‘Lovell’ is no longer used as a drying
cultivar, seed can only be obtained from nurs-
ery seed orchards. Thus, ‘Halford’ has gradu-
ally supplanted ‘Lovell’ in the nursery trade.
However, since ‘Halford’ is usually sold in
bulk as cannery pits, questions about whether
it is true to type occasionally arise.
‘Bailey’
‘Bailey’ is a naturalized peach selection from
Iowa, circa 1836, that produces uniform seed-
lings with good vigour (Okie, 1998). It has
good cold hardiness for a peach and is rated
only slightly less cold-hardy than ‘Siberian
C’. It has fair tolerance to root-lesion nema-
tode and is a popular rootstock on sandy soils
in more northern climates. It will usually pro-
duce a slightly smaller tree than ‘Lovell’, but
is very productive. It is susceptible to root-
knot nematodes, waterlogging, fungal root
rots and PTSL in the southern USA.
‘Siberian C’
‘Siberian C’ was selected by the Agriculture
Canada Research Station at Harrow, Ontario
(Canada) in 1967 (Weaver, 1967b; Layne, 1980;
Okie, 1998). The seedlings are uniform with
good cold hardiness. Cultivars grafted to this
rootstock show medium or medium–low
vigour and good precocity. Productivity and
crop efficiency are satisfactory. ‘Siberian C’
has the additional advantage of inducing
slightly earlier ripening time, as well as having
a lower susceptibility to Leucostoma or Valsa
canker (Leucostoma cincta (Sacc.) Hohn.), but it
is not tolerant of root waterlogging, nema-
todes, bacterial canker or crown gall. ‘Siberian
C’ tends to de-harden the scion cultivar before
spring in moderate climates, and therefore is
mostly planted in colder regions.
Sections Euprunus and Microcerasus
species cultivars
Species of Section Euprunus cover a wide range
of rootstocks, but only a few have become
widely used for peach. Rootstocks from these
206 G.L. Reighard and F. Loreti
species are more resistant to waterlogging, cal-
careous soil, replant problems and specific soil
pathogens than Section Euamygdalus species.
However, these rootstocks have other problems
with peach such as nutritional deficiencies,
non-uniform growth, excessive suckering and
frequent graft incompatibility (Loreti, 1984).
These negative characteristics were present
to various degrees in ‘historical’ rootstocks
such as ‘Akerman’, ‘Brussel’, ‘Common Plum’,
‘Common Mussel’, ‘Pershore’ and others, which
now have mostly disappeared by virtue of
genetic improvements by breeders within these
species. Therefore, only commonly known root-
stocks that have promising traits are described.
Prunus insititia cultivars
‘St. Julien A’ is a selection from a plum popu-
lation derived from P. insititia, which includes
genotypes that have morphological and likely
genetic characteristics (Casas et al., 1999) sim-
ilar to P. domestica. Seedlings from this geneti-
cally complex population exhibit extreme
morphological variability, which also leads to
heterogeneous size control in budded scions.
Therefore, a number of clones (e.g. A, B, C, G,
J, K) were selected at East Malling, UK, among
which ‘St. Julien A’ was the most interesting
and widely used (Fiorino, 1967). This clone is
propagated fairly easily by layering, and also
by hardwood cuttings. ‘St. Julien A’ is com-
patible with commercial peach cultivars and
is considered cold-hardy. However, it now is
used infrequently because of its sensitivity to
calcareous soil and pathogens such as P. syringae
(Reighard, 1994).
‘St. Julien GF 655/2’ was selected by INRA
from open-pollinated seedlings of ‘St. Julien
d’Orleans’ and is propagated by layering, hard-
wood cuttings and micropropagation (Loreti,
1984; Zacchini and Morini, 1995). ‘GF 655/2’
has a rather shallow root system that does not
adapt well to droughty soils, but it is fairly tol-
erant of calcareous, heavy, waterlogged soils
and replant sickness, albeit to a lesser degree
than ‘Damas GF 1869’ (P. domestica × P. spinosa)
(Salesses, 1977). It also induces medium–low
vigour, precocity and satisfactory scion pro-
ductivity. Consequently, this rootstock is suit-
able for medium- to high-density planting
systems even in highly fertile soils. ‘GF 655/2’
suckers profusely (even more so if propa-
gated in vitro) but less than ‘GF 1869’ and has
good resistance to crown gall (Bliss et al.,
1999) and the silver leaf fungus (Stereum pur-
pureum).
‘Adesoto 101’ (‘Empyrean® 101’ in the
USA) was selected by the Experimental Sta-
tion of Aula Dei (ESAD-CSIC) in Zaragoza
(Spain) from a population of open-pollinated
seedlings of ‘Pollizo de Murcia’ (Moreno et al.,
1995a). This clonal rootstock can be used for
peach, apricot, plum and almond and has
good compatibility with all peach and nectar-
ine cultivars tested in Spain (M. Moreno,
Spain, 2006, personal communication). How-
ever, poor scion growth and/or survival on
‘Adesoto 101’ have been observed in two
peach cultivars at several test locations in the
USA (G. Reighard, unpublished results). It
propagates easily by both hardwood cuttings
and in vitro techniques (preferred nursery
method) and also adapts satisfactorily to
calcareous (up to 10–11% limestone), fine-
textured and waterlogged soils. Peach culti-
vars budded on this rootstock show lower
vigour than trees budded on peach seedling
or ‘Damas GF 1869’ and the fruits ripen about
3–7 days earlier. ‘Adesoto 101’ is resistant or
immune to M. arenaria, M. incognita and M.
javanica and is moderately tolerant of P. vul-
nus (Pinochet et al., 1999) but has been highly
susceptible to bacterial canker and PTSL in
South Carolina (Reighard et al., 2006).
Prunus domestica cultivars
‘Damas C’, derived from P. domestica, was exten-
sively used in the past as a peach rootstock.
‘Damas C’ was selected from ‘Black Damas’
(P. domestica) at the former East Malling
Research Station, UK (Hatton, 1921; Taylor,
1949; Tydeman, 1956). This rootstock is easily
propagated by layering, but not by hardwood
cuttings. ‘Damas C’ induces good vigour in
budded trees, but its shallow root system
leads to weak anchorage. It is suitable for wet,
cold soils that are not very deep. However, it
suckers profusely, which is why it has fallen
out of favour with commercial producers.
‘GF 43’ rootstock was selected in 1950 by
INRA from a ‘Prune d’Ente’ seedling. It is
readily propagated by micropropagation,
 Rootstock Development
though propagation by stooling and hard-
wood cuttings has been unsatisfactory. Having
a deep and well-developed root system, this
rootstock exhibits good tolerance of slightly
calcareous, wet and heavy soils and is inter-
mediately resistant to root waterlogging
(Grasselly, 1987). It induces vigour in budded
trees similar to peach seedlings and has good
graft compatibility with all cultivars tested.
However, trees on ‘GF 43’ are non-precocious
with low production efficiency. ‘GF 43’ pro-
duces few or no suckers and is somewhat
sensitive to replant sickness and susceptible
to Chlorotic leaf spot virus, which can cause
graft incompatibility.
‘Tetra’ (‘Empyrean® 3’ in the USA) root-
stock originated from open-pollinated seedlings
of ‘Regina Claudia Verde’ at the Experimental
Institute of Fruit Growing, Rome, Italy. It
readily propagates by either micropropaga-
tion or hardwood cuttings. Trees are vigorous
and uniform in the nursery and easily bud-
ded or grafted. ‘Tetra’ has a uniformly devel-
oped root system, which has good anchorage.
In addition, it adapts to many soil types and
has resistance to calcareous soils comparable
to that of ‘GF 677’ but with good tolerance of
root waterlogging. ‘Tetra’ has excellent com-
patibility with peach, nectarine, apricot and
plums and rarely suckers. Flowering can be
delayed 5–6 days compared with ‘GF 677’.
Tree vigour is 15–20% less on ‘Tetra’ than on
‘GF 677’, and fruit ripens 3–4 days earlier.
‘Tetra’ is mostly resistant to root-knot nema-
todes (Meloidogyne spp.), highly susceptible
to P. vulnus, and preliminary trials show resis-
tance to Phytophthora cinnamomi and tolerance
to A. mellea (Nicotra and Moser, 1997; M.
McKenry, California, 2006, personal commu-
nication).
‘Penta’ (‘Empyrean® 2’ in the USA) root-
stock originated from open-pollinated seedlings
of ‘Imperial Epineuse’ at the Experimental Insti-
tute of Fruit Growing, Rome, Italy. Its character-
istics are similar to ‘Tetra’, except it differs in
its ability to induce higher vigour, similar to
‘GF 677’, and it does not induce early fruit
ripening (Nicotra and Moser, 1997). In Cali-
fornia field trials, it was a very good host for
M. xenoplax and was similar to ‘Nemaguard’
in its host status to P. vulnus (M. McKenry,
California, 2006, personal communication).
207
Prunus pumila cultivars
‘Pumiselect®’ (‘Rhenus 2’), a P. pumila selec-
tion from Germany (Jacob, 1992), is compati-
ble with many peach cultivars. It is resistant
to M. javanica and tolerant of sandy soils and
drought but is susceptible to waterlogging, A.
tabescens and iron chlorosis, and sometimes
exhibits uneven anchorage, leading to lean-
ing. Peach fruit size on ‘Pumiselect®’ root-
stocks was reported to be smaller than on
peach rootstocks (Reighard et al., 2007). Selec-
tions of P. pumila such as ‘Mando’ have been
poor hosts for ring nematodes (M. xenoplax)
(Westcott et al., 1994). However, in Californian
trials, ring nematode host status for ‘Pumise-
lect®’ was less than ‘Nemaguard’ but higher
than ‘Lovell’ seedlings (M. McKenry, Califor-
nia, 2006, personal communication).
Interspecific hybrid cultivars
The most interesting hybrids of this group are
derived from peach × almond and peach × P.
davidiana, which have helped remedy serious
problems caused by nematodes and iron chlo-
rosis, the latter being extremely common in
many European peach-growing areas. Fur-
thermore, since interspecific hybrids are less
susceptible to certain pathogens, they are a
useful tool for overcoming abnormalities of
the soil environment such as root waterlogging,
droughty soils, salinity or replant problems.
Among these rootstocks, the peach–almond
‘GF 677’ has become the most widespread
rootstock in the European peach-growing
areas. Research in the last few years has
reported several similar new hybrids possess-
ing superior horticulture traits that are still
under field trials (Salesses et al., 1998; Moreno,
2004; Massai and Loreti, 2004).
New peach rootstock cultivars have been
imported into the USA during the past 10–15
years with the majority of them being com-
plex Prunus L. hybrids that must be propa-
gated vegetatively. The pedigrees of these
hybrids contain several different Prunus L.
species which confer traits that P. persica lacks,
such as adaptation or tolerance to heavy soils,
waterlogging, alkalinity, drought, vigour con-
trol and soil fungal diseases (Reighard, 2000).
208 G.L. Reighard and F. Loreti
Within the USA, a few clonal rootstocks
are being used in commercial peach produc-
tion on a limited scale. ‘Atlas’ and ‘Viking’
from Zaiger Genetics, Inc. in California are
interspecific Prunus L. hybrids used on replant
sites due to their replant tolerance, root-knot
resistance, protection against ring nematode
and tree vigour (www.davewilson.com),
although ‘Viking’ grows off much better the
first year if grown as a container tree rather
than as a bare-root seedling. An old (i.e. 1957)
USDA-released cultivar, ‘Hiawatha’ (P. besseyi ×
P. salicina), is now being evaluated for vigour
control as a rootstock. Several other promis-
ing size-controlling clonal rootstocks were
released in 2004 from California. These root-
stocks are P. salicina × P. persica hybrids and
are named ‘Controller 5’ (formerly ‘K146-43’)
and ‘Controller 9’ (formerly ‘P30-135’) (DeJong
et al., 2005). Both these rootstocks are good
hosts to M. incognita and P. vulnus nema-
todes (M. McKenry, California, 2006, personal
communication).
Peach × almond hybrids
‘GF 677’ (‘Paramount®’ in the USA) is a natu-
ral hybrid selected by INRA. It may be propa-
gated by softwood and hardwood cuttings,
but now is exclusively propagated in vitro
(micropropagation). It is a very vigorous root-
stock (10–15% more vigorous than peach
seedlings) with a well-developed root system
that ensures good anchorage. It is adapted to
infertile and droughty soils as long as they
are permeable and well-drained. Its resistance
to iron chlorosis is high, having good produc-
tivity even in soils with 10–12% limestone. In
addition, replant problems are ameliorated
because of its vigour. These characteristics, along
with good graft compatibility with commer-
cial peach and nectarine cultivars, have made
this rootstock extremely popular in Italy and
other Mediterranean basin countries, where it
is almost as widespread as the standard seed-
ling rootstocks (Loreti and Massai, 1995).
Despite these favourable traits, ‘GF 677’
often induces excessive scion vigour, result-
ing in delayed precocity, low yields in the first
few years, smaller fruit size and poor fruit
colour. However, these defects disappear when
the trees achieve a vegetative–reproductive
balance and enter into full production. Still,
this rootstock is not recommended for very
fertile soils or high planting densities. ‘GF
677’ also adapts poorly to heavy and water-
logged soils since it is sensitive to root water-
logging. Furthermore, it is susceptible to
A. mellea, M. incognita, A. tumefaciens, Phyto-
phthora cactorum and S. purpureum (Loreti and
Massai, 1995), and has a lesser degree of
susceptibility to Verticillium albo-atrum, some-
where between that of almond and peach
(Loreti and Massai, 2006a).
‘I.S.’ SERIES.The ‘I.S.’ series of clones were de-
rived from seedlings obtained by open polli-
nation of the peach–almond hybrid ‘GF 557’,
which was selected by the Department of
Fruit Science and Crop Protection at the Uni-
versity of Pisa (Loreti and Massai, 1998). This
series has many characteristics in common
with ‘GF 677’, but they differ by the different
degrees of vigour induced in scion cultivars.
The most interesting clones are, in increasing
order of vigour, ‘Sirio’, ‘Castore’ and ‘Polluce’
(Loreti and Massai, 1998, 2006b).
‘Sirio’ (formerly ‘I.S. 5/22’) has poor root
induction ability and is difficult to propagate
by stoolbeds and layering, but can be propa-
gated by cuttings and even more efficiently
with in vitro micropropagation (Loreti and
Massai, 1994). ‘Sirio’ produces a good root
system and is adapted to fertile and perme-
able soils, but still exerts control over vegeta-
tive growth. Its resistance to iron chlorosis is
good, though slightly less than that of ‘GF
677’. Compared with trees grafted on to ‘GF
677’, trees budded on ‘Sirio’ are about 40%
smaller, yield earlier and have better crop effi-
ciency, larger fruit size and improved fruit
colour. It is compatible with the main com-
mercial cultivars. ‘Sirio’ is suitable for high-
density planting systems on fertile and
chlorosis-inducing soils where ‘GF 677’ can-
not be used, due to its vigour. It is unknown if
‘Sirio’ is suitable for replant sickness sites
(Loreti and Massai, 1998).
‘Castore’ (formerly ‘I.S. 5/19’) also has
poor root induction ability, but can be micro-
propagated in vitro. ‘Castore’ prefers fertile
and permeable soils, whereas it is unsuitable
for non-tiled heavy and waterlogged soils.
It is a semi-dwarfing rootstock, reducing
 Rootstock Development
vegetative growth to about 30% less than ‘GF
677’. Production is good, and ‘Castore’ achieves
high crop efficiency based on fruit yield ver-
sus vegetative growth parameters. Fruit qual-
ity is improved with higher SSC, favourable
sugar:acid ratio and intense fruit colour.
Moreover, this stock also is adapted to infer-
tile soil. ‘Castore’ is very suitable for fertile
soils and high-density planting systems (Loreti
and Massai, 2006b).
‘Polluce’ (formerly ‘I.S. 5/8’) is similar to
‘Castore’ in difficulty to propagate, but it
can be micropropagated. Like other peach–
almond hybrids, it is not suitable for wet,
heavy and inadequately drained soils, but is
adapted to permeable soils with medium or
high fertility. It induces about 20% less vigour
to the scion than ‘GF 677’, and produces good
yields with high yield efficiency and improved
fruit quality. ‘Polluce’ offers an alternative to
‘GF 677’ in medium- to high-fertility soils,
where it allows closer tree spacing in orchards
and easier tree maintenance. In poor soils, it
can be semi-dwarfing but still achieves good
production (Loreti and Massai, 2006b).
‘HANSEN 2168’ AND ‘HANSEN 536’. The Hansen
clones were selected by the University of Cal-
ifornia (USA) from a seedling population ob-
tained by a peach × almond cross (Kester and
Asay, 1986). They are propagated by hard-
wood cuttings, but also can be micropropa-
gated. Both of these rootstocks induce greater
vigour than peach seedlings, but have pro-
ductivity similar to peach and ‘GF 677’. In
addition, both have tolerance to calcareous
soils and resistance to root-knot nematodes
(M. incognita, M. javanica) and have tolerance
to drought and saline soils. ‘Hansen 2168’
(= ‘Hansen 2’) is moderately tolerant of Phy-
tophthora but both clones are very sensitive to
crown gall, with ‘Hansen 536’ (= ‘Hansen 5’)
slightly more susceptible. They also are more
susceptible than peach to Verticillium wilt,
waterlogging, and perhaps bacterial canker.
Since these rootstocks have only recently been
introduced in Europe, they still need time to
be evaluated; although preliminary observa-
tions are not promising because they have not
been as tolerant to calcareous soils as ‘GF 677’
or other new European rootstocks (M. More-
no, Spain, 2006, personal communication).
209
‘ADAFUEL’. ‘Adafuel’, an almond–peach hy-
brid, was selected at ESAD-CSIC (Zaragoza,
Spain) from a seedling population obtained
by open pollination of the ‘Marcona’ almond
cultivar (Cambra, 1990). It propagates easily
by hardwood cuttings, giving a better rooting
percentage than ‘GF 677’. ‘Adafuel’ is extreme-
ly vigorous and is suitable for calcareous and
loam soils provided they are well-drained. It
is graft-compatible with most commercial
peach and almond cultivars and is similar to
‘GF 677’ in its growth and yield characteristics
as a rootstock (Moreno et al., 1994b). ‘Adafuel’
is resistant to powdery mildew (Sphaerotheca
pannosa), plum rust (Tranzschelia pruni-spinosae)
and shot hole (Corineum beijerinckii), but is
very susceptible to Meloidoigyne spp., which
limits its commercial utility.
‘ADARCIAS’. ‘Adarcias’, like ‘Adafuel’, is an
almond–peach hybrid selected from an open-
pollinated seedling population at ESAD-
CSIC. It readily propagates by hardwood cut-
tings, but can be micropropagated in vitro
(Moreno and Cambra, 1994). It is a rootstock
that performs well in calcareous and loam
soils if they are well-drained. ‘Adarcias’ in-
duces lower vigour than ‘Adafuel’ and ‘GF
677’, but has greater crop efficiency and high-
er fruit SSC (Albás et al., 2004). Therefore it
reduces excessive tree growth, and thus limits
management costs. It is graft-compatible with
the several peach and nectarine cultivars test-
ed thus far. Nursery trials indicate that ‘Adar-
cias’ is resistant to C. beijerinckii Oud. and
T. pruni-spinosae (Pers.) Diet.
Peach × Prunus davidiana hybrids
‘Cadaman® Avimag’ rootstock was selected
in Hungary from an interspecific hybrid of P.
persica × P. davidiana, and then introduced in
France by INRA (Edin and Garcin, 1996).
‘Cadaman®’ can be propagated by softwood or
semi-hardwood cuttings. The vigour induced
in the scion is initially comparable to that of ‘GF
677’, but vigour tends to decrease 4 or 5 years
after orchard establishment. Furthermore, ‘Cada-
man®’ induces earlier precocity, comparable
productivity and larger fruit size compared
with ‘GF 677’. ‘Cadaman®’ shows good resis-
tance to waterlogging, unlike peach–almond
210 G.L. Reighard and F. Loreti
hybrids, and is tolerant to iron chlorosis and
replant sickness. It is also resistant to M. incog-
nita and M. javanica, but is as good a host to
M. xenoplax as ‘Nemaguard’. Owing to its
high vigour, ‘Cadaman®’ is not an appropriate
rootstock for high-density planting systems.
‘Barrier 1’ (‘Primo’ or ‘Empyrean® 1’ in the
USA) is another peach × P. davidiana hybrid
selected at the former Institute for the Propa-
gation of Woody Species, Florence, Italy. The
vigour induced by ‘Barrier 1’ is similar to or
greater than that by ‘GF 677’, and ‘Barrier 1’
has an extensive, deep root system that ensures
good anchorage. It propagates easily by hard-
wood cuttings or micropropagation. It is well
adapted to different pedologic environments
such as calcareous soil, waterlogging or replant
sickness, and has good resistance to root-knot
nematodes (Roselli, 1998; Loreti and Massai,
2002a). Furthermore, it induces higher pro-
ductivity and larger fruit size than does ‘GF
677’. Therefore, this rootstock is adapted to
replant soils that have waterlogging problems
where peach–almond hybrids cannot be used.
Prunus cerasifera hybrids
The rootstocks ‘Mr.S. 2/5’ and ‘Mr.S. 2/8’
were selected by the Department of Fruit Sci-
ence and Crop Protection of the University of
Pisa, Italy from an open-pollinated Myrobalan
(P. cerasifera) population. The origin of ‘Mr.S.
2/5’ is uncertain, but since it is a pentaploid
hybrid (2n = 40) it probably is a spontaneous
P. cerasifera × P. spinosa hybrid (Loreti et al.,
1988). It also could be a P. domestica × P. spinosa
hybrid like ‘Damas GF 1869’ (also 2n = 40),
since its morphological characteristics are
similar to those of P. domestica.
The two ‘Mr.S.’ clones have many char-
acteristics in common, such as easy propaga-
tion by layering, hardwood cuttings and
micropropagation. For commercial propaga-
tion, the in vitro micropropagation method is
used. Budding operations must begin a few
days earlier than in other rootstocks, because
the vegetative growth in the nursery ceases
rather early (Loreti and Massai, 1995). In
addition, ‘Mr.S. 2/5’ is sensitive to rust (T.
pruni-spinosae (Pers.:Pers.) Diet.), but less so
to Phytophthora spp. and Armillaria spp. Graft
compatibility has been good with all tested
peach and nectarine cultivars, and scion vigour
is reduced by 10–15% compared with that of
peach seedlings and by 25–30% compared
with ‘GF 677’. Both rootstocks are tolerant of
replant disease. Suckering has been limited in
Italy but excessive suckering in ‘Mr.S. 2/5’ has
been observed in the USA (G. Reighard, per-
sonal observation). Peach production on these
rootstocks has been good with good fruit size,
increased SSC and good fruit colour compared
with ‘GF 677’ (De Salvador and Monastra,
1996; Loreti and Massai, 1999). Fruit also ripen
a few days earlier on ‘Mr.S. 2/5’. Differences
between the two ‘Mr.S.’ clones are that ‘Mr.S.
2/5’ has greater resistance to crown gall (Zoina
and Raio, 1999), calcareous soil and root water-
logging, and can survive extended periods of
low soil oxygen content.
‘Myran® Yumir’ or ‘Myran®’ is an inter-
specific hybrid of ‘Belsiana’ plum (probably
P. cerasifera × P. salicina) and ‘Yunnan’ peach
selected by INRA in 1950 and propagates by
hardwood and semi-hardwood cuttings (Renaud
et al., 1988). ‘Myran®’ induces medium–high
vigour in grafted peach cultivars and has
good graft compatibility. Scion vigour of
mature trees has been less than ‘GF 305’ peach
in Europe, but in the USA scion vigour is
10–15% greater than for peach rootstocks
(Reighard et al., 2004). Fruit ripening, fruit
size and colour are similar to trees grafted on
to the peach seedling, but yield efficiency has
been low in the USA. ‘Myran®’ is free from
suckering, tolerant of root waterlogging, toler-
ant to A. mellea (i.e. France not USA) and Melo-
idogyne spp. nematodes, but it is sensitive to
calcareous soil and susceptible to bacterial
canker and PTSL (Grasselley, 1987; Renaud
et al., 1988; Reighard et al., 1997).
‘Ishtara® Ferciana’ or ‘Ishtara®’ is a com-
plex interspecific hybrid of ‘Belsiana’ plum
(probably P. cerasifera × P. salicina) and a natu-
ral hybrid of P. cerasifera × P. persica selected
by INRA (Grasselly, 1987; Renaud et al., 1988).
It is a multiple species-compatible rootstock,
which induces medium vigour and can be
used for peach, apricot and plum cultivars.
‘Ishtara®’ has good graft compatibility with
peach, rarely suckers, and has shown good
resistance to the peach tree borer (Synanthedon
exitiosa (Say)) in the USA (Reighard et al., 2004).
It adapts well to different soil and climatic
 Rootstock Development
conditions, although sensitivity to water-
logged and calcareous soils has been observed
in Europe. In Italy, ‘Ishtara®’ has induced ear-
lier fruit maturity by several days in ‘Sun-
crest’ (Loreti and Massai, 2002b), but no
phenology differences in peach have been
observed in the USA (Reighard et al., 2004).
Productivity is good on loams and fertile
soils, where it reduces peach tree growth by
up to 30% and improves fruit size. However,
fruit yields have been low in the USA, and
peach cultivars on ‘Ishtara®’ are extremely
susceptible to bacterial canker and PTSL
(Reighard et al., 1997).
European plum hybrids
‘Julior® Ferdor’ or ‘Julior®’ is an interspecific
hybrid of P. insititia × P. domestica selected in
1965 by INRA. It is graft-compatible with
peach, plum and apricot cultivars and is
propagated by hardwood cuttings and micro-
propagation. ‘Julior®’ induces low to medium
vigour, albeit greater than ‘GF 655/2’, and is a
dwarfing rootstock for peach cultivars even
in very fertile soils (Loreti and Massai, 2002a).
It has good resistance to root waterlogging, but
is sensitive to calcareous soil above pH 8.2
(Grasselly, 1988). It is quite variable in growth
in different pedoclimatic environments, lacks
precocity and suckers profusely soon after
orchard establishment. ‘Redtop’ peach on
‘Julior®’ rootstock in South Carolina has had
poor yields, low yield efficiency, early fruit
maturity, improved fruit quality and high
mortality from bacterial canker (G. Reighard,
unpublished results). In California, ‘Julior®’
supports four times the ring nematode num-
bers of that for ‘Nemaguard’ (M. McKenry,
California, 2006, personal communication).
‘Jaspi® Fereley’ or ‘Jaspi®’ rootstock was
selected by INRA from a cross of P. salicina × P.
spinosa (Renaud and Salesses, 1990). It is a
multiple species-compatible rootstock, but is
more suitable for apricot and plum than
peach as compatibility with peach is limited.
‘Jaspi®’ induces 20% less vigour than peach
seedlings in Europe, but is very dwarfing
(~60% of standard) in trials in the USA. It is
adapted to difficult soil environments due to
its tolerance to waterlogging, calcareous soil
and replant sickness, but peach cultivars on
211
‘Jaspi®’ are extremely susceptible to bacterial
canker (Reighard et al., 2006). Unfortunately,
‘Jaspi®’ is not recommended in the USA because
it is probably incompatible with many peach
cultivars.
8.4 Peach Rootstock Breeding
Programmes
The adoption of rootstocks more suited to the
requirements of technologically advanced
fruit growing began in Europe in the 1950s,
but it was not until the development of inten-
sive fruit growing that it assumed greater
importance. For pome fruits such as apple
and pear, the introduction of new rootstocks
occurred rapidly, so that in recent years there
is almost exclusive use of clonal rootstocks.
However, for stone fruits like peach, the demand
for non-traditional rootstocks emerged some-
what later, when peach began to be planted
extensively into areas having soil and climatic
conditions that were not fully suited to this
species such as waterlogged, compacted, sub-
calcareous or alkaline soils, and environ-
ments subject to frequent low temperatures
in winter and, in particular, spring frosts
(Loreti, 1988).
It is now generally accepted that consid-
erable technical and economic advantages
can be achieved by using rootstocks selected
for their genetic characteristics and environ-
mental adaptation. Such rootstocks have
become widely available in the nursery trade,
the demand is increasing and the market is
insisting on new rootstocks with enhanced
qualities. This has prompted a number of sci-
entific institutions, public organizations and
private breeders to develop new rootstocks
that more closely match the requirements of
modern fruit growing. In Europe, research
has been particularly active in countries such
as France, Spain and Italy, where extensive
work on genetic improvement of fruit tree
rootstocks has been undertaken for several
decades with special attention now being
focused on new rootstocks for peach. Table
8.1 lists many standard and recently released
rootstocks being used for commercial peach
production throughout the world.
212 G.L. Reighard and F. Loreti
Research institutions and objectives
France
In France, genetic improvement and evalua-
tion of peach rootstocks by INRA began in the
1940s and has focused over the last 30 years
on both the selection of new lines and clones
from common rootstocks and on the develop-
ment of multiple species-compatible, inter-
specific hybrids. In addition to evaluation of
the main bioagronomic parameters, attention
has focused on determination of sensitivity to
the main biotic and abiotic sources of stress in
order to select the most tolerant or resistant
rootstocks. To that end, numerous trials were
conducted to assess seed germination, chill-
ing requirements, seedling morphology and
uniformity of progeny from selected mother
tree genotypes or ‘cultivars’.
Significant genotypes were obtained by
virtue of the selection programme designed
first by J. Souty at Pont-de-la-Maye and then
subsequently by R. Bernhard, C. Grasselly
and G. Salesses. The programme was based
primarily on the interspecific peach–almond
cross, with more recent experiments focusing
on developing new peach × P. davidiana
hybrids. Concurrently, an interspecific hybrid-
ization programme using different Prunus L.
species was initiated. The objectives were to
create genotypes with greater resistance to
root waterlogging, nematodes, Phytophthora
and Armillaria, and to improve graft compat-
ibility and scion dwarfing traits (Salesses
et al., 1998).
The vast and complex genetic improve-
ment work carried out by INRA for over half
a century has resulted in the development of
numerous peach rootstocks, some of which
are already widely adopted in Mediterranean
basin countries. Currently a new genetic
improvement programme is under way using
P. cerasifera and almond hybrids with the aim
of selecting new rootstocks resistant to root
waterlogging, drought, iron chlorosis and
Meloidogyne spp. nematodes (Dirlewanger
et al., 2002). This programme has successfully
used biotechnology via marker-assisted selec-
tion to pyramid resistance genes from three
species into one rootstock. Progeny from this
programme are still under development and
testing, but they hold tremendous promise
for ‘designer’ rootstocks in the future.
Italy
In Italy, rootstock genetic improvement was
begun in the late 1950s by Professor F. Scara-
muzzi. Attention focused on assessment of
the bioagronomic characteristics of a very
extensive range of rootstocks belonging to
different species obtained either by crosses or
by selection of seedlings and wild types col-
lected in different areas of Italy or other Med-
iterranean countries (Scaramuzzi et al., 1976).
This work was continued in more depth by F.
Loreti, R. Guerriero and R. Massai, first at the
former Istituto di Coltivazioni Arboree in Pisa
and then, since 1986, at the Department of
Fruit Science and Crop Protection of the Uni-
versity of Pisa.
Research carried out since the late 1950s
has mainly involved genetic improvement
programmes for pear, plum, apricot and
particularly peach rootstocks. The programme
objectives for the development of peach root-
stocks were: (i) uniform growth in the nurs-
ery; (ii) use of seed propagation; (iii) graft
compatibility with most peach and nectarine
cultivars; (iv) different degrees of dwarfing of
the scion to facilitate adaptation to different
soil fertility and orchard systems; (v) increased
production efficiency and improvement of
fruit quality; and (vi) resistance or reduced
susceptibility to biotic and abiotic stresses,
with special attention to root waterlogging,
salinity, drought, calcareous iron chlorosis
and replant problems. This programme has
successfully produced tolerant peach seed-
ling and interspecific hybrid rootstocks with
differing levels of scion dwarfing.
Another Italian institution that devoted
considerable resources to genetic improve-
ment of peach rootstocks is the Experimental
Institute of Fruit Growing in Rome. Research
began in the late 1970s, and primarily focused
on rootstocks tolerant to waterlogging and
calcareous soil. Trials were conducted on
seedlings obtained by open pollination from
about 100 plum cultivars. At the same time, a
programme for selection of rootstocks resis-
tant to M. incognita, M. javanica and P. vulnus
nematodes was undertaken, using P. persica
 Rootstock Development
and peach–almond hybrids (Nicotra and Moser,
1997, 1998).
Since 1995, work in Italy on genetic
improvement and bioagronomic evaluation
of peach rootstocks has been coordinated and
funded by the Ministry of Agricultural and
Forestry Policies through two national proj-
ects. The first of these, ‘Genetic Improvement
of Rootstocks’, focuses on development of
rootstocks through crossing and selection
programmes, and more recently through bio-
technology methods. The second project,
‘Evaluation of Rootstocks’, evaluates the most
widely used rootstocks available in the Italian
nursery market. Fourteen ‘Operative Units’,
composed of members of university institutes,
experimental stations and producer associa-
tions, are involved in work on this project and
have established experimental field trials in
the most important Italian peach-growing
areas (Loreti, 1997). This project observes the
bioagronomic behaviour of proposed peach
rootstocks prior to their release for commer-
cial use. Information is then made available to
fruit growers through the publication of peri-
odic lists in which rootstocks are classified into
one of three categories: recommended, prom-
ising or non-recommended (Fideghelli and
Nicotra, 2002).
Spain
In Spain, selection of peach rootstocks was
initiated in the 1960s with the identification of
P. insititia clones that could be easily propa-
gated by hardwood cuttings. The first studies
were performed at EEAD-CSIC in Zaragoza
using ‘Pollizo de Murcia’ germplasm, which
is a P. insititia population that was widely
used in local nurseries due to its ease of prop-
agation by suckers and tolerance to iron chlo-
rosis, root asphyxia and salinity (Cambra,
1970).
Attention focused initially on graft compati-
bility of the test material with peach, apricot,
plum and almond cultivars and its adapta-
tion to compacted and calcareous soils.
Efforts were subsequently directed towards
identifying clones with naturally high root-
ing potential, in order to limit commercial
propagation by suckers to avoid virus dis-
eases. Additional ‘Pollizo’ clone selection
213
programmes were undertaken in the follow-
ing years by EEAD-CSIC and Unidad de
Fruticultura del Centro de Investigación y
Tecnologia Agroalimentaria de Aragón (CITA,
formerly SIA-DGA) of Zaragoza and the Cen-
tro Regional de Investigaciones Agrarias of
La Alberca, Murcia.
On the other hand, at the EEAD-CSIC,
selection of peach × almond hybrids began in
1970 with the identification and collection of
spontaneous hybrids or open-pollinated seed-
ling populations from throughout Spain.
Work basically focused on ease of vegetative
propagation, graft compatibility with peach
and almond, and tolerance to iron chlorosis.
Later, the best clones were evaluated for vigour
and production characteristics in grafted scion
cultivars (Cambra, 1990; Moreno and Cambra,
1994).
Recently, these research institutes and
the private nursery Agromillora Catalana SA
have moved towards developing rootstocks
with greater tolerance to soils with high active
limestone content, prolonged summer drought
and poor soil fertility – all of which represent
characteristics found in the most important
peach-growing areas in Spain (Aragón, Cata-
lonia and Murcia) – as well as resistance to
root-knot and root-lesion nematodes (Pinochet
et al., 2002). Therefore, controlled interspecific
crosses have been made with the purpose of
bringing together the desirable traits of plum
species (P. cerasifera, P. insititia), P. davidiana,
almond and peach into new rootstocks (Moreno,
2004).
USA and Canada
Peach rootstock development in the USA
evolved in the early 20th century and was
carried out by scientists from the USDA and
public universities. This work has now shifted
during the past 20 years to mostly private
nurseries and breeders. Initially, much of the
first rootstock research for peach occurred in
the 1930s and 1940s to find cultivars or selec-
tions that were tolerant to the known biotic
and abiotic stresses on peaches (Tufts, 1929;
Hutchins, 1936; Long and Whitehouse, 1943;
Weinberger et al., 1943; Havis et al., 1946; Day,
1953). This work largely screened new germ-
plasm and existing cultivars from P. persica
214 G.L. Reighard and F. Loreti
and other Prunus L. species for desired traits.
Later, the first breeding work was initiated to
incorporate root-knot nematode resistance
from sources such as ‘Shalil’, ‘Yunnan’, ‘S-37’,
‘Okinawa’ and ‘Nemaguard’. This work pro-
gressed from the 1950s and 1960s (Hansen
et al., 1956; Sharpe, 1957; Sharpe et al., 1969)
to the 1980s and 1990s (Sherman et al., 1981,
1991; Ramming and Tanner, 1983; Okie et al.,
1994b), where a number of selections were
released (e.g. ‘Nemared’, ‘Flordaguard’ and
‘Guardian®’).
Since at least the mid-1980s, peach–
almond rootstocks for calcareous soil have
been developed at the University of Califor-
nia (UC) at Davis by D. Kester (e.g. ‘Hansen
2168’ and ‘Hansen 536’, ‘Nickels’), Burchell
Nursery, Inc. (Oakdale, California) (e.g. ‘SLAP’ =
‘Cornerstone’) and Bright’s Nursery, Inc.
(Reedley, California) (e.g. ‘Bright’s Hybrid®’),
and dwarfing rootstocks by USDA and UC-
Davis (DeJong et al., 2004a,b). Widely adapted,
interspecific rootstocks (e.g. ‘Citation’, ‘Viking’
and ‘Atlas’) have been released by Zaiger
Genetics, Inc. for multiple types of soil and
replant problems. In the southern USA, root-
stocks that are resistant to bacterial canker
and Armillaria spp. are currently under evalu-
ation at USDA in Byron, Georgia (Beckman
et al., 1997b) and for tolerance to salinity and
calcareous soils in Texas (Ottman and Byrne,
1988; Shi and Byrne, 1995). Most all other
peach rootstock development and evaluation
in the USA are of imported rootstocks licensed
by commercial nurseries. Many of the above
rootstocks are tested in North America in the
NC-140 national rootstock trials (www.nc140.
org) prior to or shortly after being available to
fruit growers.
In Canada, ‘Harrow Blood’ and ‘Siberian
C’ rootstocks were released in 1967 by Agri-
culture Canada at Harrow, Ontario (Layne,
1980; Okie, 1998). Later, the Harrow breeding
programme under R.E.C. Layne selected sev-
eral genotypes from ‘Bailey’ × ‘Siberian C’
crosses. These selections plus two cultivars,
‘Chui Lum Tao’ and ‘Tzim Pee Tao’, from
northern China are being evaluated for cold
hardiness, P. penetrans tolerance and L. cinta
resistance. Selections ‘H7338013’ and
‘H7338019’ have yielded as well as standard
peach rootstocks but have not been named or
released (Layne and Jui, 1994; Reighard et al.,
2004).
Rootstocks in development
Work on genetic improvement of stone fruit
rootstocks continues to produce innovations
that expand the range of rootstocks available
on the world market (Beckman and Lang,
2003). Research on high-vigour almond–
peach hybrids conducted in Spain has recently
led to the selection of several clones of par-
ticular interest: ‘Monegro’, ‘Garnem’ and
‘Felinem’, all obtained by the Unidad de Fru-
ticultura SIA-DGA of Zaragoza from the
almond ‘Garfi’ × ‘Nemared’ cross. These root
well with high vigour (greater than ‘GF 677’),
have red-coloured leaves, and are tolerant to
calcareous soil and resistant to root-knot
nematodes (Felipe et al., 1997; Gomez-Aparisi
et al., 2001; Iglesias et al., 2004; Moreno, 2004).
The Centro de Investigación y Desarrollo
Agroalimentario from Murcia has also
recently developed an almond–peach root-
stock, ‘Mayor®’, which has high vigour and
performs satisfactorily on poor, calcareous
and droughty soils, but is highly susceptible
to root-knot nematode (Pinochet et al., 2002).
A large group of new rootstocks has been
developed over the past few years in Eastern
Europe. In particular, genetic improvement
work conducted in Romania (Indreias et al.,
2004) has led to the development of a number
of rootstocks derived from peach seedling
selections (e.g. ‘Tomis 1’, ‘Tomis 79’, ‘T16’, ‘P1s’,
‘Oradea 1’, ‘De Balc’) or from selections of inter-
specific crosses of Prunus spp. (e.g. ‘Adaptabil’,
‘Miroper’). In Hungary, a number of peach–
almond selections are being investigated.
These include ‘O.R.T. (XXXI) 50’, ‘O.R.T. (XXXI)
51’, ‘C 410’ and ‘PeMa’ (Z. Szabo, Hungary,
2004, personal communication). In Serbia,
vineyard peach and Myrobalan seedling
selections are being tested as peach rootstocks
(Ognjanov et al., 2004). In addition, cold-hardy
rootstocks have been developed for peach in
Russia by G. Eremin from interspecific crosses
of Prunus spp. The selections ‘VVA-1’, ‘VSV-1’
and ‘Kuban 86’ have been licensed for testing
and trademarked as ‘Krymsk® 1’, ‘Krymsk®
2’ and ‘Krymsk® 86’, respectively.
 Rootstock Development 215

In Asia, China has begun to select for
rootstocks for its diverse geography and
endemic soil and pest problems (Wang et al.,
2002). However, no selections have yet been
released outside China. In Japan, the ‘Tsu-
kuba’ series (P. persica), the ‘Yusuraume’ series
(P. tomentosa) and P. japonica selections have
been developed (Yoshida and Seike, 1981; F.
Loreti, unpublished results). All are root-knot
resistant and semi-dwarfing except for the
‘Tsukuba’ series, which has a wide range of
tree vigour.
8.5 Outlook
New and better peach rootstocks from France,
Italy, Spain and other countries will soon be
available to replace older commercial root-
stocks. Almost all of these rootstocks are species
hybrids that must be propagated vegetatively,
such that in vitro micropropagation is being
or will be used to mass-produce these unique
hybrid rootstocks. As new technology permits
efficient mass propagation of these valuable
genotypes, proprietary issues (i.e. ownership,
patent royalties) abound. This is due, in part,
to the decrease in public research funding and
the considerable resources and time invested
to develop and test fruit tree rootstocks. Other
factors also may complicate the commercial
release of new rootstocks within and outside
their country of origin, such as patent laws
and licensing agreements that must be nego-
tiated between government agencies, nurser-
ies and grower groups. Despite these potential
roadblocks, new rootstocks are gradually filter-
ing through to fruit growers. These improved
cultivars together with the continued advan-
ces in biotechnology and the genome map-
ping of the Rosaceae family (www.rosaceae.
org; Arús et al., 2006) will lead to even more
dynamic rootstocks for peach within the next
20 years.

References

Albás, E.S., Jiménez, S., Aparicio, J., Betrán, J.A. and Moreno, M.A. (2004) Effect of several peach × almond
hybrid rootstocks on fruit quality of peaches. Acta Horticulturae 658, 321–326.
Alcañiz, E., Pinochet, J., Fernández, C., Esmenjaud, D. and Felipe, A. (1996) Evaluation of Prunus rootstocks
for root-lesion nematode resistance. HortScience 31, 1013–1016.
Andreu, P. and Marin, J.A. (2004) Micropropagation enhances in vitro establishment and multiplication of new culti-
vars from field grown plants of Adesoto 101 (Prunus insititia) rootstock. Acta Horticulturae 658, 605–609.
Arús, P., Yamamoto, T., Dirlewanger, E. and Abbott, A.G. (2006) Synteny in the Rosaceae. Plant Breeding Re-
views 27, 175–211.
Beckman, T.G. and Lang, G.A. (2003) Rootstock breeding for stonefruits. Acta Horticulturae 622, 531–551.
Beckman, T.G. and Pusey, P.L. (2001) Field testing peach rootstocks for resistance to Armillaria root rot. Hort-
Science 36, 101–103.
Beckman, T.G., Reighard, G.L., Okie, W.R., Nyczepir, A.P., Zehr, E.I. and Newall, W.C. (1997a) History, cur-
rent status and future potential of Guardian™ peach rootstock. Acta Horticulturae 451, 251–258.
Beckman, T.G., Nyczepir, A.P. and Okie, W.R. (1997b) The USDA-ARS stone fruit rootstock development
program at Byron, Georgia. Acta Horticulturae 451, 237–241.
Bliss, F.A., Almehdi, A.A., Dandekar, A.M., Schuerman, P.L. and Bellaloui, N. (1999) Crown gall resistance in
accessions of 20 Prunus species. HortScience 34, 326–330.
Brooks, R.M. and Olmo, H.P. (1961) Register of new fruit and nut varieties: Nemaguard peach. Proceedings
of the American Society of Horticultural Science 78, 634–635.
Cambra, R. (1970) Selección de ‘Pollizos de Murcia’ y otros ciruelos locales españoles. ITEA Producción
Vegetal 1, 115–126.
Cambra, R. (1990) ‘Adafuel’, an almond × peach hybrid rootstock. HortScience 25, 584.
Casas, A.M., Igartua, E., Balaguer, G. and Moreno, M.A. (1999) Genetic diversity of Prunus rootstocks ana-
lyzed by RAPD markers. Euphytica 110, 139–149.
Cirulli, M., Amenduni, M., Colella, C., El-Shaer, E. and D’Amico, M. (2001) Ricerca di portinnesti resistenti
alla tracheoverticilliosi. Italus Hortus 8, 50–52.
Davis, J.R. and English, H. (1969) Factors related to the development of bacterial canker in peach. Phytopa-
thology 59, 588–595.
216 G.L. Reighard and F. Loreti

Day, L.H. (1953) Rootstocks for stone fruits. California Agricultural Experiment Station Bulletin 736.
DeJong, T., Johnson, R.S., Doyle, J.F., Weibel, A., Solari, L., Basile, B., Marsal, J., Ramming, D. and Bryla, D.
(2004a) Growth, yield and physiological behaviour of size-controlling peach rootstocks developed in
California. Acta Horticulturae 658, 449–455.
DeJong, T., Almehdi, A., Johnson, S. and Day, K. (2004b) Improved rootstocks for peach and nectarine. In:
California Tree Fruit Agreement 2004 Annual Research Report. California Tree Fruit Agreement, Reedley,
California, pp. 98–107.
DeJong, T., Johnson, R.S., Doyle, J. and Ramming, D. (2005) Research yields size-controlling rootstocks for
peach production. California Agriculture 59, 80–83.
De Salvador, F.R. and Monastra, F. (1996) Agronomic evaluation of different peach rootstocks. Acta Horticul-
turae 374, 195–200.
De Salvador, F.R., Ondradu, G. and Scalas, B. (2002) Horticultural behaviour of different species and hybrids
as rootstocks for peach. Acta Horticulturae 592, 317–322.
Devyatov, A.S. (1996) Root system of plum trees on standard and dwarfing rootstocks. Fruit Varieties Journal
50, 229–235.
Dirlewanger, E., Salesses, G., Bonnet, A., Kleinhentz, M., Esmenjaud, D., Bosselut, N., Voisin, R., Bergougnoux,
V., Lecouls, A.C., Poessel, J.L., Faurobert, M., Arus, P., Gomez-Aparisi, J., Xiloyannis, C. and Di Vito, M.
(2002) Breeding for Prunus rootstocks cumulating resistance to root-knot nematodes and favorable agro-
nomic traits under Mediterranean environments: a European project. Acta Horticulturae 592, 61–67.
Edin, M. and Garcin, A. (1996) Un nuovo portinnesto ibrido per il pesco: Cadaman® Avimag. Frutticoltura 7/8,
33–35.
Elena, K. and Tsipouridis, K. (2000) Evaluation of resistance of stone fruit rootstocks to Phytophthora crown
rot. Journal of Phytopathology 148, 365–369.
Esmenjaud, D., Minot, J.C., Voisin, R., Pinochet, J., Simard, M.H. and Salesses, G. (1997) Differential response
to root-knot nematodes in Prunus species and correlative genetic implications. Journal of Nematology
29, 370–380.
Felipe, A.J., Gomez-Aparisi, J., Socias i Company, R. and Carrera, M. (1997) The almond × peach hybrid root-
stocks breeding program at Zaragoza (Spain). Acta Horticulturae 451, 259–262.
Fernández, C., Pinochet, J., Esmenjaud, D., Salesses, G. and Felipe, A. (1994) Resistance among new Prunus
rootstocks and selections to root-knot nematodes in Spain and France. HortScience 29, 1064–1067.
Fideghelli, C. and Nicotra, A. (2002) The Italian national peach cultivar and rootstock trial. Acta Horticulturae
592, 331–334.
Fiorino, P. (1967) I susini portinnesti del pesco. Rivista Ortoflorofrutticoltura Italiana 5, 355–385.
Fiorino, P. and Loreti, F. (1987) Propagation of fruit trees by tissue culture in Italy. HortScience 22, 353–358.
Funt, R.C. and Goulart, B.L. (1981) Performance of several peach cultivars on Prunus tomentosa and Prunus
besseyi in Maryland. Fruit Varieties Journal 35, 20–23.
Gomez-Aparisi, J., Carrera, M., Felipe, A.J. and Socias i Company, R. (2001) ‘Garnem’, ‘Monegro’ and ‘Felinem’:
new almond × peach hybrid rootstocks, nematode resistant and red leaved for stone fruits. ITEA Produc-
ción Vegetal 97, 282–288.
Graham, C.J. (2002) Rootstock test for perpendicular V training system. Acta Horticulturae 592, 351–355.
Grasselly, C. (1983) Nouvelles obtentions INRA de pechers port-greffes: multiplies par semences.
L’Arboriculture Fruitière 357, 50–55.
Grasselly, C. (1987) New French stone fruit rootstocks. Fruit Varieties Journal 41, 65–67.
Grasselly, C. (1988) Les porte-greffe du pecher. L’Arboriculture Fruitière 409, 29–34.
Guidoni, S., Lovisolo, C., Ferrandino, A. and Pellegrino, S. (1998) Influenza di nuovi portinnesti sullo sviluppo
fogliare e sulle prime fruttificazioni del pesco cv. ‘Suncrest’. Frutticoltura 4, 77–81.
Halbrendt, J.M., Podleckis, E.V., Hadidi, A., Scorza, R. and Welliver, R. (1994) A rapid protocol for evaluating
Prunus germplasm for tomato ringspot virus resistance. HortScience 29, 1068–1070.
Handoo, Z.A., Nyczepir, A.P., Esmenjaud, D., Van Der Beek, J.G., Castagnone-Sereno, P., Carta, L.K., Skantar,
A.M. and Higgins, J.A. (2004) Morphological, molecular, and differential-host characterization of Melo-
idogyne floridensis n. sp. (Nematoda: Meloidogynidae), a root-knot nematode parasitizing peach in
Florida. Journal of Nematology 36, 20–35.
Hansen, C.J., Lownsbery, B.F. and Hesse, C.O. (1956) Nematode resistance in peaches. California Agriculture
10(9), 5, 11.
Hatton, R.G. (1921) Stocks for stone fruits. Journal of Pomology 2, 209–245.
Havis, L., Marth, P.C. and Gardner, F.E. (1946) Orchard performance of peach variety seedlings as rootstocks
for peaches. Proceedings of the American Society of Horticultural Science 48, 115–120.
 Rootstock Development 217

Hoy, J.W. and Mircetich, S.M. (1984) Prune brownline disease: susceptibility of prune rootstocks and tomato
ringspot virus detection. Phytopathology 74, 272–276.
Hutchins, L.M. (1936) Nematode-resistant peach rootstocks of superior vigor. Proceedings of the American
Society of Horticultural Science 34, 330–338.
Iglesias, I., Montserrat, R., Carbo, J., Bonany, J. and Casals, M. (2004) Evaluation of agronomical performance
of several peach rootstocks in Lleida and Girona (Catalonia, NE-Spain). Acta Horticulturae 658, 341–
344.
Indreias, A., Dutu, I. and Stefan, I. (2004) Peach rootstocks created and used in Romania. Acta Horticulturae
658, 505–508.
Jacob, H. (1992) Prunus pumila L., eine geeignete schwachwachsende Pfirsichuntererlage. Erwerbsobstbau
34, 144–146.
Johnson, S. (1938) Prunus mexicana and Prunus hortulana as rootstocks for peaches. Michigan Agriculture
Experiment Station Quarterly Bulletin 21, 17–18.
Kester, D.E. and Asay, R.N. (1986) ‘Hansen 2168’ and ‘Hansen 536’: two new Prunus rootstock clones. Hort-
Science 21, 331–332.
Layne, R.E.C. (1980) Prospects of new hardy peach rootstocks and cultivars for the 1980’s. Compact Fruit
Trees 13, 117–122.
Layne, R.E.C. (1987) Peach rootstocks. In: Rom, R.C. and Carlson, R.F. (eds) Rootstocks for Fruit Crops. Wiley,
New York, pp. 185–216.
Layne, R.E.C. (1994) Prunus rootstocks affect long-term orchard performance of ‘Redhaven’ peach on Brook-
stone clay loam. HortScience 29, 167–171.
Layne, R.E.C. and Jui, P.Y. (1994) Genetically diverse peach seedling rootstocks affect long-term performance
of ‘Redhaven’ peach on Fox sand. Journal of the American Society of Horticultural Science 119, 1303–
1311.
Long, J.C. and Whitehouse, W.E. (1943) Variations in root-knot nematode infection of various lines of peach prog-
enies at Chico, California. Proceedings of the American Society of Horticultural Science 43, 119–123.
Loreti, F. (1984) Attuali conoscenze sui principali portinnesti degli alberi da frutta. Frutticoltura 9, 9–60.
Loreti, F. (1988) Presente e futuro dei portinnesti degli alberi da frutto. Frutticoltura 1–2, 77–86.
Loreti, F. (1997) Bioagronomic evaluation of the main fruit tree rootstocks in Italy. Acta Horticulturae 451,
201–208.
Loreti, F. and Massai, R. (1988) Il contributo dell’Universita di Pisa al miglioramento genetico dei portinnesti.
Frutticoltura 4, 9–14.
Loreti, F. and Massai, R. (1994) Sirio: nuovo portinnesto ibrido pesco × mandorlo. L’Informatore Agrario
(supplemento) 28, 47–49.
Loreti, F. and Massai, R. (1995) Orientamenti per la scelta dei portinnesti del pesco. L’Informatore Agrario
(supplemento) 32, 37–42.
Loreti, F. and Massai, R. (1998) Sirio: new peach × almond hybrid rootstock for peach. Acta Horticulturae
465, 229–236.
Loreti, F. and Massai, R. (1999) I portinnesti del pesco. L’Informatore Agrario (supplemento) 6, 39–44.
Loreti, F. and Massai, R. (2002a) I portinnesti del pesco. L’Informatore Agrario 51, 36–42.
Loreti, F. and Massai, R. (2002b) MiPAF targeted project for evaluation of peach rootstocks in Italy: results of
six years of observations. Acta Horticulturae 592, 117–124.
Loreti, F. and Massai, R. (2006a) State of arts on peach rootstocks and orchard systems. Acta Horticulturae
713, 253–268.
Loreti, F. and Massai, R. (2006b) ‘Castore’ and ‘Polluce’: two new hybrid rootstocks for peach. Acta Horticul-
turae 713, 275–278.
Loreti, F. and Morini, S. (1983) Mass propagation of fruit trees in Italy by tissue culture: present status and
perspectives. The International Plant Propagators’ Society 32, 283–291.
Loreti, F., Guerriero, R. and Massai, R. (1988) Una nuova ed interessante selezione di susino portinnesto:
‘Mr.S. 2/5’. In: Atti Convegno Nazionale ‘I portinnesti delle piante da frutto’, Ferrara, Italy, pp. 45–50.
Lötze, G. (1997) Quest for better rootstocks continues. Deciduous Fruit Grower March, 92–93.
Lu, Z.X., Reighard, G.L., Baird, W.V., Abbott, A.G. and Rajapakse, S. (1996) Identification of peach rootstock
cultivars by RAPD markers. HortScience 31, 127–129.
McClintock, J.A. (1948) A study of uncongeniality between peaches and the Marianna plum as a stock. Journal
of Agricultural Research 77, 253–260.
McFadden-Smith, W., Miles, N.W. and Potter, J.W. (1998) Greenhouse evaluation of Prunus rootstocks for resis-
tance or tolerance to the root-lesion nematode (Pratylenchus penetrans). Acta Horticulturae 465, 723–730.
218 G.L. Reighard and F. Loreti

McKenry, M.V. (1989) Nematodes. In: LaRue, J.H. and Johnson, R.S. (eds) Peaches, Plums, and Nectarines:
Growing and Handling for Fresh Market. University of California, Oakland, California, pp. 139–147.
Malcolm, P., Holford, B., McGlasson, B., Newman, S., Richards, G. and Topp, B. (1999) Growing low chill
peaches and nectarines on high chill rootstocks causes spring shock syndrome. Australian Fresh Stone
Fruit Quarterly 1(1), 11–12.
Martinelli, A. (1985) Factors affecting in vitro propagation of the peach–almond hybrids ‘Hansen 2128’ and
‘Hansen 536’. Acta Horticulturae 173, 237–244.
Massai, R. and Loreti, F. (2004) Preliminary observations on nine peach rootstocks grown in a replant soil.
Acta Horticulturae 658, 185–192.
Massonie, G. and Paison, P. (1979) Resistance de deux variétés de Prunus persica L. Batsch à Myzus persicae
Suiz. et à Myzus varians Davids: études préliminaire des mécanismes de la résistence. Annales de Zo-
ologie Ecologie Animale 11, 479–485.
Mizutani, F., Yamada, M., Taniguchi, T., Koizumi, K., Sigiura, A. and Tomana, T. (1985) Dwarfing effect of P.
japonica and P. tomentosa rootstocks on ‘Ohkubo’ peach trees. Journal of Japanese Society of Horticul-
tural Science 54, 327–335.
Moing, A. and Gaudillere, J.P. (1992) Carbon and nitrogen partitioning in peach/plum grafts. Tree Physiology
10, 81–92.
Moreno, M. (2004) Breeding and selection of Prunus rootstocks at the Aula Dei Experimental Station, Zara-
goza, Spain. Acta Horticulturae 658, 519–528.
Moreno, M. and Cambra, R. (1994) Adarcias: an almond × peach hybrid rootstock. HortScience 29, 925.
Moreno, M., Gaudillere, J.P. and Moing, A. (1994a) Protein and amino acid content in compatible and incom-
patible peach/plum grafts. Journal of Horticultural Science 69, 955–962.
Moreno, M., Tabuenca, M.C. and Cambra, R. (1994b) Performance of Adafuel and Adarcias as peach root-
stocks. HortScience 29, 1271–1273.
Moreno, M., Tabuenca, M. and Cambra, R. (1995a) Adesoto 101, a plum rootstock for peaches and other
stone fruit. HortScience 30, 1314–1315.
Moreno, M., Tabuenca, M. and Cambra, R. (1995b) Adara, a plum rootstock for cherries and other stone fruit
species. HortScience 30, 1316–1317.
Murase, S., Suzuki, K. and Yamazaki, T. (1986) Studies on dwarfing rootstocks of peach in Japan. Bulletin of
the Fruit Tree Research Station, Series A 13, 31–49.
Nicotra, A. and Moser, L. (1997) Two new rootstocks for peach and nectarines: Penta and Tetra. Acta Horticul-
turae 451, 269–271.
Nicotra, N. and Moser, L. (1998) Costituzione di nuovi portinnesti all’Istituto Sperimentale per la Frutticoltura
di Roma. Frutticoltura 4, 14–15.
Nyczepir, A.P. and Becker, J.O. (1998) Fruit and citrus trees. In: Plant and Nematode Interactions. American
Society of Agronomy Monograph No. 36. American Society of Agronomy, Madison, Wisconsin, pp.
637–684.
Nyczepir, A.P. and Beckman, T.G. (2000) Host status of Guardian peach rootstock to Meloidogyne incognita
and M. javanica. HortScience 35, 772.
Nyczepir, A.P., Zehr, E.I., Lewis, S.A. and Harshman, D.C. (1983) Short life of peach trees induced by Cricone-
mella xenoplax. Plant Disease 67, 507–508.
Nyczepir, A.P., Okie, W.R. and Beckman, T.G. (1992) Evaluating Prunus genotypes for resistance/tolerance to
Criconemella xenoplax. In: Proceedings of the 5th Stone Fruit Tree Decline Workshop, Biglerville, Penn-
sylvania, 24–26 September 1990. ARS-USDA, Byron, Georgia, pp. 37–42.
Nyczepir, A.P., Beckman, T.G. and Reighard, G.L. (1999) Reproduction and development of Meloidogyne sp.
and M. javanica on Guardian peach rootstock. Journal of Nematology 31, 334–340.
Nyczepir, A.P., Beckman, T.G. and Reighard, G.L. (2006) Field evaluation of ‘Guardian®’ peach rootstock to
different root-knot nematode species. Acta Horticulturae 713, 303–309.
Ognjanov, V., Cerovic, S., Bozovic, D., Ninic-Todorovic, J. and Golosin, B. (2004) Selection of vineyard peach
and myrobalan seedling rootstocks. In: 8th International Symposium on Integrating Canopy, Rootstock
and Environmental Physiology in Orchard Systems. International Society for Horticultural Science, Leu-
ven, Belgium, p. 44.
Okie, W.R. (1998) Handbook of Peach and Nectarine Varieties: Performance in the Southeastern United
States and Index of Names. USDA Agriculture Handbook No. 714. US Government Printing Office,
Washington, DC.
Okie, W.R., Reighard, G.L., Beckman, T.G., Nyczepir, A.P., Reilly, C.C., Zehr, E.I. and Newall, W.C. Jr (1994a)
Field-screening Prunus for longevity in the southeastern United States. HortScience 29, 673–677.
 Rootstock Development 219

Okie, W.R., Beckman, T.G., Nyczepir, A.P., Reighard, G.L., Newall, W.C. Jr and Zehr, E.I. (1994b) BY520-9, a
peach rootstock for the southeastern United States that increases scion longevity. HortScience 29, 705–706.
Ottman, Y. and Byrne, D.H. (1988) Screening rootstocks of Prunus for relative salt tolerance. HortScience 23,
375–378.
Pellegrino, S., Massai, R., Loreti, F. and Strocco, S. (1997) Comportamento bio-agronomico di alcune selezioni
di pesco Franco innestate con la cv. ‘Maria Bianca’. Frutticoltura 11, 47–50.
Perry, R., Reighard, G., Ferree, D., Barden, J., Beckman, T., Brown, G., Cummins, J., Durner, E., Greene, G.,
Johnson, S., Layne, R., Morrison, F., Myers, S., Okie, W., Rom, C., Rom, R., Taylor, B., Walker, D., Warmund, M.
and Yu, K. (2000) Performance of the 1984 NC-140 cooperative peach rootstock planting. Journal of the
American Pomological Society 54, 6–10.
Pinochet, J., Calvet, C., Herenandez-Dorrego, A., Bonet, A., Felipe, A. and Moreno, M. (1999) Resistance of
peach and plum rootstocks from Spain, France, and Italy to root-knot nematode Meloidogyne javanica.
HortScience 34, 1259–1262.
Pinochet, J., Fernandez, C., Calvet, C., Hernandez-Dorrego, A. and Felipe, A. (2000) Selection against Praty-
lenchus vulnus populations attacking Prunus rootstocks. HortScience 35, 1333–1337.
Pinochet, J., Fernandez, C., Cunill, M., Torrents, J., Felipe, A., Lopez, M.M., Lastra, B. and Penyalver, R. (2002)
Response of new interspecific hybrids for peach to root-knot and lesion nematodes, and crown gall. Acta
Horticulturae 592, 707–716.
Ramming, D.W. and Tanner, O. (1983) ‘Nemared’ peach rootstock. HortScience 18, 376.
Reighard, G.L. (1994) Field performance of 28 Prunus rootstocks and interstems in South Carolina. Hort-
Science 29, 476.
Reighard, G.L. (2000) Peach rootstocks for the United States: are foreign rootstocks the answer? HortTechnology
10, 714–718.
Reighard, G.L. (2002) Current directions of peach rootstock programs worldwide. Acta Horticulturae 592,
421–428.
Reighard, G.L., Newall, W.C., Beckman, T.G., Okie, W.R., Zehr, E.I. and Nyczepir, A.P. (1997) Field perfor-
mance of Prunus rootstock cultivars and selections on replant soils in South Carolina. Acta Horticulturae
451, 243–250.
Reighard, G., Andersen, R., Anderson, J., Autio, W., Beckman, T., Baker, T., Belding, R., Brown, G., Byers, R.,
Cowgill, W., Deyton, D., Durner, E., Erb, A., Ferree, D., Gaus, A., Godin, R., Hayden, R., Hirst, P., Kadir,
S., Kaps, M., Larsen, H., Lindstrom, T., Miles, N., Morrison, F., Myers, S., Ouellette, D., Rom, C., Shane,
W., Taylor, B., Taylor, K., Walsh, C. and Warmund, M. (2004) Growth and yield of Redhaven peach on 19
rootstocks at 20 North American locations. Journal of the American Pomological Society 58, 174–202.
Reighard, G.L., Ouellette, D.R. and Brock, K.H. (2006) Growth and survival of 20 peach rootstocks and selec-
tions in South Carolina. Acta Horticulturae 713, 269–274.
Reighard, G.L., Ouellette, D.R. and Brock, K.H. (2007) Survival, growth and yield for Carogem peach on an
interstem and two dwarfing rootstocks. Acta Horticulturae 732, 303–306.
Remorini, D., Massai, R. and Loreti, F. (2005) Effetto del portinnesto e della gestione della chioma sulla qual-
ità dei frutti di pesco. Frutticoltura 12, 43–48.
Renaud, R. and Salesses, G. (1990) Prunier/pecher: deux nouveaux port-greffes. Fruits et Legumes 73, 22–23.
Renaud, R., Bernhard, R., Grasselly, C. and Dosba, F. (1988) Diploid plum × peach hybrid rootstocks for stone
fruit trees. HortScience 23, 115–117.
Roberts, A.N. and Westwood, M.N. (1981) Rootstock studies with peach and Prunus subcordata Benth. Fruit
Varieties Journal 35, 12–20.
Roselli, G. (1998) Miglioramento genetico dei portinnesti presso il CNR di Firenze. Frutticoltura 4, 20–22.
Salesses, G. (1977) Research about the origin of two Prunus rootstocks, natural interspecific hybrids: an il-
lustration of a cytological study carried out in order to create new Prunus rootstocks. Annales de
L’Amelioration des Plantes 27, 235–243.
Salesses, G. and Alkai, N. (1985) Simply inherited grafting incompatibility in peach. Acta Horticulturae 173,
57–62.
Salesses, G. and Bonnet, A. (1992) Some physiological and genetic aspects of peach/plum graft incompatibility.
Acta Horticulturae 315, 177–186.
Salesses, G., Renaud, R. and Bonnet, A. (1988) Creation de port-greffe par hybridation interspecifique au sein
des pruniers. In: Proceedings of 8th Colloque sur les Recherches Frutieres. INRA-CTIFL, Bordeaux,
France, pp. 151–159.
Salesses, G., Dirlewanger, E., Bonnet, A., Lecouls, A.C. and Esmenjaud, D. (1998) Interspecific hybridization
and rootstock breeding for peach. Acta Horticulturae 465, 209–217.
220 G.L. Reighard and F. Loreti

Scaramuzzi, F., Loreti, F. and Guerriero, R. (1976) La selezione di nuovi portinnesti seguiti dall’Istituto di
Coltivazioni Arboree di Pisa. In: Atti incontro frutticolo SOI ‘I portinnesti degli alberi da frutto’. SOI-ICA,
Pisa, Italy, pp. 15–26.
Sharpe, R.H. (1957) ‘Okinawa’ peach resists root-knot nematodes. Florida Agriculture Research Report Janu-
ary, 18.
Sharpe, R.H., Hesse, C.O., Lownsbery, B.F., Perry, V.G. and Hansen, C.J. (1969) Breeding peaches for root-
knot nematode resistance. Journal of the American Society of Horticultural Science 94, 209–212.
Sherman, W.B., Lyrene, P.M. and Hansche, P.E. (1981) Breeding peach rootstocks resistant to rootknot nema-
todes. HortScience 16, 523–524.
Sherman, W.B., Lyrene, P.M. and Sharpe, R.H. (1991) Flordaguard peach rootstock. HortScience 26, 427–
428.
Shi, Y. and Byrne, D.H. (1995) Tolerance of Prunus rootstocks to potassium carbonate-induced chlorosis.
Journal of the American Society of Horticultural Science 120, 283–285.
Taylor, H.V. (1949) The Plums of England. Crosby, Lockwood and Son, Ltd, London.
Tufts, W.P. (1929) Nematode resistance of certain peach seedlings. Proceedings of the American Society of
Horticultural Science 26, 98–100.
Tydeman, H.M. (1956) A description and classification of certain plum rootstocks. Report of the East Malling
Research Station 1956.
Wang, L., Zhu, G. and Fang, W. (2002) Peach germplasm and breeding programs at Zhengzhou in China. Acta
Horticulturae 592, 177–182.
Weaver, G.M. (1967a) Harrow Blood peach. In: Canadian Horticultural Council Report. Canadian Horticul-
tural Council, Ottawa, p. 187.
Weaver, G.M. (1967b) Siberian C peach. In: Canadian Horticultural Council Report. Canadian Horticultural
Council, Ottawa, pp. 187–188.
Weinberger, J.H., Marth, P.C. and Scott, D.H. (1943) Inheritance study of root-knot nematode resistance in
certain peach lines. Proceedings of the American Society of Horticultural Science 42, 321–325.
Westcott, S.W. III, Zehr, E.I., Newall, W.C. Jr and Cain, D.W. (1994) Suitability of Prunus selections as hosts
for the ring nematode (Criconemella xenoplax). Journal of the American Society of Horticultural Science
119, 920–924.
Yamaguchi, M., Haji, T., Yaegaki, H. and Nakano, M. (2004) Screening of graft-compatibility between ‘Akat-
suki’ and several interstocks of related species and interspecific hybrids grafted on peach seedlings.
Bulletin of National Institute of Fruit Tree Science 3, 67–76.
Yoshida, M. and Seike, K. (1981) Breeding of peach rootstocks resistant to root-knot nematode. II. Breeding of
resistant rootstocks by hybridization. Bulletin of the Fruit Tree Research Station, Series A 8, 31–45.
Zacchini, M. and Morini, S. (1995) The effect of photoperiod reduction on growth of Prunus insititia GF 655/2
cultures during multiplication and rooting. Plant Propagator 7, 14–16.
Zarrouk, O., Aparicio, J., Gogorcena, Y. and Moreno, M.A. (2006) Graft compatibility for new peach root-
stocks in nursery. Acta Horticulturae 713, 327–329.
Zehr, E.I., Miller, R.W. and Smith, F.H. (1976) Soil fumigation and peach rootstocks for protection against
peach tree short life. Phytopathology 66, 689–694.
Zoina, A. and Raio, A. (1999) Susceptibility of some peach rootstocks to crown gall. Journal of Plant Pathol-
ogy 81, 181–187.
Zuccherelli, G. (1979) Moltiplicazione in vitro dei portinnesti clonali di pesco. Frutticoltura 41, 15–20.
 9 Propagation Techniques

F. Loreti and S. Morini

Department of Fruit Science and Crop Protection, University of Pisa, Pisa, Italy

9.1 Introduction 221

9.2 Nursery Seedling Propagation 222
Nursery site and soil 222
Propagation from seed 222
9.3 Nursery Hardwood Cutting 224
9.4 Greenhouse Semi-hardwood Cutting 227
9.5 Micropropagation 230
Preparation of the propagation material 230
Explant disinfection and setting up aseptic culture 231
Shoot proliferation 232
Rooting and preparation for in vivo growth 233
Transfer and acclimatization of young plants 233
Culture medium 234
Problems connected to in vitro propagation 235
9.6 Embryo Culture 236
9.7 Stoolbeds and Layering 237
9.8 Scion Propagation 237
Budding and grafting 237
Micrografting 239
Herbaceous grafting 240
9.9 Outlook 240

9.1 Introduction its quality is poor, it can invalidate the other

Crop efficiency of a peach orchard during its
commercial life is influenced by several factors
such as planting system, cultural practices,
environmental and soil conditions. The nursery
quality of the maiden trees used to plant the
orchard also plays a prominent role because, if
factors. Tree quality is represented by a num-
ber of bioagronomic, genetic and sanitary
characters, which collectively make the orchard
capable of reaching its higher cropping poten-
tial. Tree quality depends on the right choice of
plant material, propagating techniques and
cultural management in the nursery.

© CAB International 2008. The Peach: Botany, Production and Uses

(eds D.R. Layne and D. Bassi) 221
222 F. Loreti and S. Morini
Peach cultivars, like the majority of fruit
tree species, are genetically heterozygous and
therefore propagated exclusively by asexual
methods. This is considered necessary because
reproduction from seed produces individuals
that differ appreciably from one another and from
the parent tree. Thus, although current propaga-
tion techniques make it possible to obtain self-
rooted varieties, grafting desirable scions on to
suitable rootstocks still represents the most
common propagation technique for peach.
Grafting is also favoured by the recent
development of a wide range of rootstocks
capable of overcoming the problems related
to biotic and abiotic stress that could not be
satisfactorily addressed with self-rooted cul-
tivars. In addition, scion vigour can more eas-
ily be controlled by the use of appropriate
rootstocks, which limit canopy growth and
consequently also reduce management costs.
Further advantages of medium–low vigour
rootstocks are their ability to induce early
cropping of the scion and enhance its produc-
tion efficiency; they also adapt well to the cur-
rent trend towards increased planting density
even in highly fertile soils.
Budding and grafting represent the
worldwide method for scion cultivar propa-
gation. For rootstocks, on the other hand,
both sexual and asexual propagation meth-
ods can be adopted.
9.2 Nursery Seedling Propagation
Nursery site and soil
For establishment of nursery activities, there
are several general principles that must be
observed. Of particular importance is the
choice of nursery site and soil. Both of these
aspects are fundamental for economically
sound nursery management.
In choosing an appropriate site, the nurs-
ery should be established on a topographi-
cally level site to facilitate the various cultural
operations, with easy access to irrigation water
and in full sun. Ideally, areas that may be
affected by late spring frost should be avoided,
as young growing shoots could suffer freezing
injury during the growth phase, resulting in
stunted maiden trees that are too small for
commercial sale. This is of particular impor-
tance if the June budding technique is to be
adopted. Grafting operations can be conducted
in June only if the rootstock has achieved a suf-
ficient diameter by the end of spring.
Windy sites should be avoided because
they require the additional cost of protection
by an artificial or living windbreak. Proximity
to the ocean coast should likewise be avoided
because winds carrying salt can cause foliar
damage leading to fairly extensive necrosis
especially of young leaves and shoot tips.
Choice of soil type is important. Soil tex-
tures that are too heavy, too light or poorly
drained should be avoided. To obtain good-
quality rootstocks it is necessary to choose
medium-texture, well-drained soils with access
to irrigation water. A further requirement is
that soils previously used for peach growing
should be avoided in order to preclude the
risk of replant disorders. Nurserymen who do
not have sufficient land available to ensure
proper rotation of the nursery plants are
obliged to treat the soil with fumigants. Methyl
bromide is the most commonly used fumi-
gant, especially for the control of nematodes.
However, it should not be overlooked that
this fumigant not only creates problems of a
microbiological nature, but also its use through-
out the European Union was prohibited after
2005. As an alternative, good results have
been obtained by soil solarization (Katan and
DeVay, 1991), which has been adopted for
replant problems in peach nurseries (Di Vaio
et al., 2001). In addition, it is essential to
acquire information on soil nutritional status
and pH prior to nursery establishment. A
deficiency of macro- or microelements could
be corrected by suitable fertilization. It is more
difficult, on the other hand, to correct pH when
active limestone results in a pH of 8–10,
causing severe symptoms of leaf chlorosis.
Finally, soils contaminated with crown
gall (Agrobacterium tumefaciens) should never
be used for nursery production.
Propagation from seed
Propagation from seed still represents the
most widely adopted method for nursery
 Propagation Techniques
production of peach seedling rootstocks.
Although the availability of clonal rootstocks
for peach has over the last few years led to
progressively decreasing use of seedlings, there
are a number of reasons why seedlings are
still the preferred propagation method today.
The first reason is economic given the lower
cost of obtaining pits (stones) and producing
seedlings in the nursery as compared with
clonal rootstocks, the latter requiring royalty
fees. Another reason consists of the easy avail-
ability of seedlings to the nursery market, as
the material utilized is obtained from the seed
of commercial varieties extracted during
industrial processing, from Prunus sylvestris
(Slavic seedling) produced at low cost in the
Balkan countries or, in some cases, from wild-
type peaches, although the latter are now
used only in countries with less advanced
fruit-growing practices. A wide selection of
interesting peach seedlings is now available,
carefully selected for specific genetically
homogeneous characters transmitted to the
progenies by sexual reproduction.
If selected seedlings are to be used, the
pits are produced by setting up special fields
of seed-bearing mother trees. Care is taken to
isolate them from any peach orchards in the
vicinity, to prevent unwanted cross-pollination.
Examples of such seedling lines are ‘Rubirà’,
‘Higamà’ and ‘Montclar’ from France; ‘P.S.A5’,
‘P.S.A6’ and ‘P.S.A7’ from Italy; ‘Siberian C’,
‘Bailey’ and ‘Harrow Blood’ from Canada; and
‘Nemaguard’, ‘Nemared’ and ‘Guardian’ from
the USA. Not only are these seedlings homo-
geneous within their line, but they have also
been selected for their different degrees of
vigour, frost resistance induction of the scion,
or reduced susceptibility or resistance to
nematodes.
Seed provenance and characteristics are
important considerations for seed propaga-
tion. Seeds should be obtained from areas
having a climate similar to the planned grow-
ing area, in order to avoid any difficulty of cli-
matic adaptation that could result in stunted
growth. This danger is particularly likely to
arise if seedlings are grown in a colder climate
than the area of seed provenance (Hartmann
et al., 2002).
When pits are supplied by the processing
industry, it is preferable to make use of varieties
223
that have already been tested as rootstocks.
Many commercial varieties commonly used
for processing are important seed sources for
rootstocks. Such is the case of ‘Lovell’, ‘Halford’,
‘Bailey’ and ‘Suncling’ in the USA, Canada and
Mexico; ‘Polara’ and ‘Sims’ in Argentina;
‘Algrighi’ and ‘Capdeboscq’ in Brazil; and
‘Golden Queen’ and ‘Elberta’ in Australia,
etc. It is clear that such varieties produce root-
stocks whose characteristics are less uniform
than in genetically selected lines of peach, but
they are still more satisfactory than wild-type
peach.
A useful precaution is to avoid collecting
seeds from very early-ripening varieties, as
these have poor germination ability due to
the presence of aborted or immature embryos.
It is also important to separate the pits
promptly from the fruit flesh, preferably as
soon as the fruit has been picked, after which
they should be washed and dried and placed in
storage rooms until needed for stratification.
Knowledge of seed dormancy constitutes
another crucial factor in nursery practice.
Dormancy is due partly to the presence of the
stony endocarp, which prevents moisture
penetration, with the result that seed imbibi-
tion and embryo development cannot take
place. But it is also caused by the presence of
inhibiting substances such as abscisic acid,
which is considered to be an important factor
preventing seed germination and inducing
embryo dormancy in other stone fruits as
well (Lang, 1996).
Dormancy breaking can be achieved by
stratifying pits in sand or other material that
is kept moist and well aerated. Stratification
is maintained at a temperature of 3–5°C for
10–12 weeks of vernalization but with the
possibility of varying both the temperature
and the duration as a function of the root-
stock’s specific chilling requirements. It is
generally observed that by stratifying the pits
at 5°C, the highest germination percentage is
obtained after 60 days for ‘P.S.A6’ and after 74
days for ‘P.S.B2’. Higher temperatures or insuf-
ficient chilling could lead to development of
seedlings with normal roots but rosetted epi-
cotyls. Excessive duration of cold stratifica-
tion could induce a portion of seeds to enter
secondary dormancy (Guerriero and Scala-
brelli, 1984).
224 F. Loreti and S. Morini
Other methods are also available for
breaking dormancy, such as pit scarification
or hormone treatment of seeds. In the former
case, the aim is to crack the woody endocarp
in order to make it permeable to moisture and
oxygen, but without damaging the seed. For
hormone treatment, gibberellic acid (GA3)
may be used to overcome physiological dor-
mancy and stimulate germination in seeds with
dormant embryos (Koornneef et al., 2002).
It may also be possible to allow chilling
to occur directly in the field in the autumn
(direct seedbed sowing or even row sowing
in mounds), although a brief period of strati-
fication may still be beneficial. While this
technique may be advantageous in order to
perform budding in June (on account of more
rapid seedling growth), it may cause a poorly
branched root system and consequently be
more affected by transplantation once the
rootstock liners are planted out.
The row spacing adopted in the nursery
is important. While inter-row spacing may to
some extent depend on the implements uti-
lized for normal cultural techniques, it is
advisable not to adopt excessively close spac-
ing within the row, in contrast to the tendency
often observed in common nursery practice.
Adequate spacing within the row is particu-
larly important when it is necessary to obtain
maiden trees that are abundantly feathered
between 40 and 60 cm from the ground. It is
likewise crucial for certain training systems,
such as the sprint palmette, free spindle and
delayed open centre, in which these feathers
are used to build the framework of the first
branches.
9.3 Nursery Hardwood Cutting
Propagation from cuttings is based on the
ability of plant cells to resume meristematic
activity and produce adventitious roots in tis-
sues that have reached a fairly advanced stage
of differentiation. Since the cutting is repre-
sented by a portion of an organ (growing shoot,
dormant shoot, leaf or root) that is separated
from the mother plant, it may suffer from the
influence of environmental conditions. Thus in
order to promote rooting, metabolic activity
must be shielded from the adverse effects of
stress that could impair rhizogenesis. Only
species characterized by a naturally high root-
ing potential or endowed with burr knots can
be propagated by setting the cuttings to root
directly in the nursery. Numerous studies on
the behaviour of self-rooted peach cultivars
have shown that the results are poor. How-
ever, this propagation method is adopted
exclusively for some clonal rootstocks.
Among the various types of cuttings (soft-
wood, semi-hardwood and hardwood), hard-
wood and to a lesser extent semi-hardwood
cuttings are the only ones used for commercial
peach propagation. Hardwood cuttings collec-
ted from the mother trees during the autumn–
winter period are generally 25–30 cm long
and are taken from the median or apical por-
tion of 1-year-old shoots. Since they are com-
posed of a number of distinct tissues, hardwood
cuttings generally possess nutritional, energy
and hormone substances capable of inducing
root initiation and the consequent development
of adventitious roots. But when such substances
are present in insufficient quantity to promote
rhizogenesis, appropriate treatment or propa-
gation techniques must be undertaken to
enhance the rooting process. It should also be
noted that the rhizogenic potential of a given
peach rootstock or cultivar is dependent upon
a number of factors, including the characteris-
tics of the mother tree, the time of harvesting
of the cuttings, rhizogenic treatment and the
propagation technique adopted.
It has been extensively documented for
many species that there are distinct advan-
tages in taking cuttings from trees reared
exclusively for the production of propagation
material. This not only guarantees that the
cuttings will be genetically true to type and of
suitable health status, but it also allows better
control over the factors influencing their root-
ing potential. Thus it is well known in nurs-
ery practice that the management techniques
applied to mother trees can exert positive
effects on the rooting potential of cuttings,
although no documented studies on this aspect
in peach are available. For example, the type of
training can influence rooting by modifying
the vigour of branches from which cuttings
are taken: if trees are severely pruned to
encourage the formation of more vigorous
 Propagation Techniques
branches, a favourable effect on rooting is
generally observed (Bartolini and Fiorino,
1978). It was also found that the application
of specific fertilizer dressings may modify the
tissue nutritional balance, leading to more
effective induction and growth of adventi-
tious roots. In this regard, some minerals such
as P, K, Ca and N have been shown to be sig-
nificantly involved in the rooting process
(Blazich, 1988). Furthermore, the use of stock
hedges maintained for harvesting cuttings
not only allows implementation of manage-
ment techniques specifically designed to
ensure good-quality cuttings (fungal and hor-
mone treatment to enhance the rooting poten-
tial of cuttings), but also facilitates the planning
and harvesting of propagation material at
times most suitable for rhizogenesis (Barto-
lini and Fiorino, 1978).
In this context, one important observa-
tion is that the time of harvesting of cuttings
has a noticeable effect on adventitious root
formation, which shows marked variation in
response during the autumn–winter period.
Some studies (Loreti et al., 1985) have reported
that peach cuttings taken between October
and January generally exhibit a more satisfac-
tory rooting response in the northern hemi-
sphere. Some plum rootstocks used for peach,
which either are now very infrequent or have
been completely abandoned, such as ‘Bromp-
ton’, ‘S. Giuliano A’, ‘GF 1869’, ‘GF 43’ and
‘GF 655/2’, were long propagated from cut-
tings harvested during this period. More
recently, other rootstocks have shown fairly
high rooting percentages when cuttings are
harvested during the autumn period. Among
the different genotypes, a certain variability
of response in relation to different time peri-
ods is observed. Consequently, the outcome
of rooting may not always be fully satisfac-
tory. A number of physiological markers of
rooting competence (e.g. content of auxins
and phenolic compounds, enzymes, etc.)
appeared to be related to the rooting of cut-
tings (Fadl and Hartmann, 1967; Bassuk and
Howard, 1981; Bhattacharya, 1988), but they
were not always reliable among the several
species to predict rooting response. Recently,
another physiological parameter not directly
involved in the rooting process (i.e. redox
status), has provided interesting results as a
225
rooting marker (Sorce et al., 2004). Moreover,
only for a few species (Lorenzi and Ceccarelli,
1978) are reliable and easily recognizable
morphological rooting markers known to
identify the most suitable moment for taking
cuttings. In most cases to date, the time for
harvesting cuttings is established empirically
on the basis of the operator’s experience
regarding the behaviour of a given genotype.
The duration of the period in which cut-
tings have the greatest rooting potential is
another highly important factor for the success
or failure of the rooting process. Some root-
stocks, such as the peach–almond hybrid ‘GF
677’, have only a limited period during which
cuttings can be taken, namely from December
to January. Other peach genotypes, such as
‘P.S.B2’, have a much longer period, ranging
from September to January (Loreti et al., 1985).
However, if cuttings of this rootstock are har-
vested towards the end of winter, rooting
shows no appreciable difference whereas bud
take in the field is often drastically reduced.
This highlights the crucial role of the time of
harvesting on successful propagation.
Despite this awareness, the phenomena
that cause rhizogenesis to vary with varia-
tions in time of harvesting remain to be deter-
mined. While considerable importance is
attributed to tissue physiological conditions
and to the balance between different hormones
that act as inhibitors or promoters of rooting, a
multiplicity of factors can directly or indi-
rectly influence the rooting process and some
of these are yet to be defined (Loreti and
Pisani, 1982).
In general, with hardwood cuttings, a
substantial improvement in rooting has been
obtained by applying auxins. Auxin applied
to cuttings from rootstocks with good natural
rooting potential was able to further enhance
rooting performance when the cuttings were
allowed to root directly in the nursery. But the
most widespread and increasingly popular
utilization of auxins in nursery practice con-
cerns rootstocks that show poor rhizogenic
activity, which in the past suffered from a
very low success rate for propagation from
cuttings.
Many auxin compounds are available,
among which IBA (indole-3-butyric acid) is
the most frequently used. It can be applied at
226 F. Loreti and S. Morini
concentrations ranging from 10–20 ppm (dilute
solutions) up to 1000–3000 ppm (concentrated
solutions). In the former case, treatment con-
sists of dipping the base of the cuttings into
the solution for 12–24 h, while a quick dip of
just a few seconds is sufficient with concen-
trated solutions. An alternative method uses
a powder preparation, which is more practi-
cal to handle but has proved to be less effec-
tive. The solutions generally used for peach
hardwood rootstock cutting propagation are
water–alcohol solutions with IBA concentra-
tions ranging from 1000 ppm, to promote
rooting of ‘P.S.A5’ or ‘P.S.B2’, to up to 2000
ppm, for the rootstocks ‘Tetra’ and ‘Penta’
(Nicotra and Moser, 1996).
Additional procedures are sometimes
adopted to further enhance rooting. In certain
cases the cutting material is etiolated prior to
IBA treatment, while at other times the cuttings
may be subjected to washing for 24 h in running
water before auxin is supplied. Rooting of cut-
tings treated with auxin can be enhanced by
bottom heating. In fact, an appreciable improve-
ment in both rooting and transplant take of
November–January-harvested cuttings was
observed with this technique (Erez and Yablow-
itz, 1981; Scalabrelli and Couvillon, 1986). This
method exploits the positive effect of tempera-
ture in promoting tissue metabolic activity,
whereby heat applied in conjunction with auxin
treatment enhances the root induction process
of cuttings harvested during the winter rest
period (Fiorino and Loreti, 1965).
Generally, in order to trigger formation
of root initials, it may be sufficient to main-
tain a temperature of 18–20°C for a few days
around the basal portion of the cuttings, tak-
ing care to ensure that the upper part remains
exposed to the cooler ambient temperature so
that buds remain in the state of rest. The
equipment required to set up bottom heating
is simple and inexpensive: all that is needed
is a heated bin supplied with a heat source at
the base and a thermostat to set and maintain
a constant temperature. Cuttings are planted
in an inert medium, generally composed of
perlite or perlite plus peat (3:1), which allows
good aeration and suitable humidity.
Cuttings rooted by this method usually
exhibit a good root system with buds that are
still quiescent, favouring successful transplant
take in the nursery (Fig. 9.1/Plate 62). Best
results are obtained when a hardening phase is
included prior to transplanting the rooted cut-
tings. This involves gradually decreasing the
bin temperature until ambient temperature is
reached, enabling root tissues to achieve a
higher degree of differentiation and greater
stability, with the twofold advantage of reduc-
ing the risk of transplantation injury and
encouraging more rapid resumption of water
and nutrient uptake from soil.
However, in peach one important feature
that should not be overlooked is that even if
the rooting and transplanting stages are car-
ried out with the utmost attention in order to
preclude any damage to the roots, not all cut-
tings will survive in the nursery. Also, a certain
degree of cutting mortality is noted during the
rooting stage as well, regardless of whether
rooting took place in the heated bin or directly
in the nursery. Research into the underlying
causes of this problem has shown that cut-
tings undergo a collapse that starts from the
basal tissues, leading to the formation of large
lysigenous areas which in turn cause the
detachment of cortical tissue from the xylem
(Fabbri, 1977; Biricolti et al., 1990). The cutting
collapse seems to be independent of propaga-
tion techniques or the various treatments,
while it is probably related to prunasin, a cya-
nogenic glycoside that decreases during root-
ing (Fiorino and Mattii, 1992). An increase,
albeit still somewhat limited, in the cutting
survival rate could possibly be obtained by
subjecting the basal tissues of the cutting to
washing for 24 h, both before auxin treatment
and before transplanting rooted cuttings to the
open field (Bartolini and Fabbri, 1982). Washing
would appear to have the effect of favouring
removal of the above-mentioned cyanogenic
glycosides or of some intermediate decompo-
sition products such as cyanidric acid.
Overall, bottom heating continues to offer
a useful method for propagation of peach root-
stocks, hardwood cuttings being taken from
the end of autumn to the beginning of winter
and then treated with IBA. As noted in the
‘Nursery Seedling Propagation’ section above,
numerous rootstocks are propagated using
this method including ‘Penta’, ‘Tetra’, ‘Ades-
oto 101’, ‘GF 677’, ‘Adafuel’, ‘Adarcia’, ‘Myran
Yumir’, ‘Citation’, ‘Hansen 2168’, ‘Hansen 536’
 Propagation Techniques
and ‘Yulior’, giving results that can vary from
satisfactory to very good. But it should be
kept in mind that these are now almost exclu-
sively propagated in vitro.
9.4 Greenhouse Semi-hardwood Cutting
As an alternative to propagation by hard-
wood cutting, semi-hardwood cuttings are
227
Fig. 9.1. Rooted hardwood cuttings
of peach.
sometimes collected to propagate peach cul-
tivars and rootstocks. Characterized by tis-
sues that are still undergoing differentiation,
semi-hardwood cuttings are harvested in the
northern hemisphere during mid-July to
mid-August, from non-lignified shoots.
Since shoots have a partially woody
consistency and generally have two leaves,
semi-hardwood cuttings can only be used
in conjunction with mist propagation
228 F. Loreti and S. Morini
(Fig. 9.2/Plate 63). This technique enables
elevated relative humidity to be maintained
inside the rooting environment (glasshouse,
tunnel or special open-air facility) by means
of a fine mist spray. This leaves a thin film of
water on the leaves, thereby reducing evapo-
transpiration and the overall temperature of
the cutting, with the latter continuing to exer-
cise its metabolic activity. However, even with
the most sophisticated devices for delivery of
carefully measured quantities of water, total
control of leaf evapotranspiration cannot eas-
ily be achieved on account of the variations in
temperature and light intensity at different
times of the day. Therefore the risk that cut-
tings may suffer water stress can never be
excluded. If results are unsatisfactory when
only a mist spray is applied, a cooling system
and shade cover may provide a means of
decreasing the temperature and light inten-
sity within the glasshouse.
In addition to the need for a suitable
environment to avert the risk of water stress,
semi-hardwood cuttings, like hardwood cut-
tings, show a better rooting performance when
treated with auxins (Fig. 9.3/Plate 64). IBA is
the most extensively used, but in some cases
Fig. 9.2. Mist propagation facility for rooting of semi-hardwood cuttings.
good results have also been achieved with
NAA (naphthalene acetic acid) or mixtures of
the two hormones. Ideally concentrations
should be low, and whenever possible it is
preferable to use the water-soluble IBA potas-
sium salt, rather than requiring a water–alcohol
solution. Total auxin concentrations range
between 100 and 500 ppm up to a maximum
of 1000–3000 ppm. The base of the cutting is
dipped in the solution for just a few seconds.
Higher concentrations can lead to tissue
dehydration due to the elevated alcohol con-
tent. This could lead to disintegration of cortical
tissues at the base of the cuttings which may be
predisposed to fungal and bacterial attack.
Potential damage can be averted by using
porous media, such as perlite, which allows
good aeration and facilitates water drainage.
Interesting rooting results have also been
obtained by spraying cutting leaves with
auxin (IBA or NAA at 25–50 ppm) in addition
to, or instead of, the basal treatment (Fiorino
and Vitagliano, 1968). Thus basal applications
of 1000 ppm IBA combined with leaf spraying
of 50 ppm IBA on ‘Okinawa’ cuttings resulted
in 90% rooting, with more roots per cutting.
Other promising treatments that enhanced
 Propagation Techniques
rooting include spraying leaves with nutrient
solutions at the end of each daily water-mist-
ing cycle. Treatment with boron, potassium
and manganese was likewise found to increase
the rooting percentage of ‘Okinawa’ cuttings,
while manganese and iron were observed to
be particularly active, achieving a noticeable
increase in rooting by preventing early leaf
senescence (Fiorino and Vitagliano, 1968).
Although the rooting treatment clearly
plays a crucial role in the rhizogenic process,
it is by no means the only factor. The particu-
lar shoot portion from which the cutting is
taken also has major importance, with apical
cuttings giving a better rooting response than
basal cuttings in some genotypes, while the
opposite appears to be true in other geno-
types (Marini, 1983). Many other aspects must
also be taken into consideration, and nursery-
men may often be guided by their own expe-
rience acquired in nursery practice.
One vital aspect, however, common to
all semi-hardwood cuttings, is the need for
229
Fig. 9.3. Rooted semi-hardwood
cuttings that were treated with auxins.
extreme care through acclimatization prior to
transfer to the open field. This is because the
leaf stomatal closure mechanism on new
shoots developed during the rooting stage is
slowed down because of the elevated relative
humidity in the misting glasshouse. Thus,
water loss from tissues can be severe once the
cutting is transferred to the external environ-
ment. Therefore the cuttings must undergo a
period of gradual reduction of relative humid-
ity by progressively increasing the time
intervals between misting operations. Alter-
natively, cuttings can be placed under shad-
ing facilities equipped with irrigation, until
the delicate tissues have become accustomed
to external environmental conditions and
developed a sufficient cuticle.
A further consideration is that a consider-
ably longer period of time is required to obtain
a rootstock that can be budded with the scion.
Since the cuttings are harvested during the
summer and time must be allowed for root
formation and acclimatization, trees will not
230 F. Loreti and S. Morini
be ready for grafting until the end of summer
the following year. This means an extra bur-
den of management costs in addition to the
expense incurred in setting up the misting
glasshouse, so that overall costs for semi-
hardwood cuttings are likely to be noticeably
higher than for hardwood cuttings.
Assessment of the costs and benefits of
the different techniques must be conducted
by the individual nurseryman based on spe-
cific nursery requirements. Economics aside,
it can be stated that propagation by semi-
hardwood cutting with the aid of the misting
technique can be performed for a fairly wide
range of clonal peach rootstocks, as described
in the previous section.
9.5 Micropropagation
Micropropagation is a method of vegetative
propagation performed in the laboratory by
means of in vitro tissue culturing. It is based
on the concept of cell totipotency, first hypoth-
esized by Wichow in 1858 (Vasil and Vasil,
1972). Many studies with in-depth research
were undertaken before it became possible to
propose this technique as a commercial prop-
agation method. A fundamental step in this
developmental process was made by Morel
(1964), who was the first to succeed in clonal
propagation of orchids from shoot tips.
Since then notable progress has been
achieved with horticultural species. Research
on fruit trees began somewhat later, but intense
experimental work has made it possible to
move within just a few years from laboratory
studies to commercial nursery production
(Fiorino and Loreti, 1987). At present, micro-
propagation is adopted for a considerable num-
ber of fruit tree species, including varieties – but
above all rootstocks – which have displayed
severe limitations with conventional propa-
gation methods. Micropropagation represents
the most widely adopted method for propa-
gation of numerous peach clonal rootstocks.
With micropropagation many plants can
be obtained with a small amount of starting
material, within a very short period of time
and in an extremely small space (Fig. 9.4/
Plate 65). As numerous new plantlets can be
obtained from a starting single bud or shoot
tip, even one mother tree can be a sufficient
explant source. In addition, this technique
makes it possible to propagate species (culti-
vars and rootstocks) characterized by poor
rooting ability and therefore difficult to mul-
tiply with traditional techniques. As the entire
propagation cycle takes place in the labora-
tory, greater flexibility is achieved in planning
the production, which is no longer dependent
on environmental and cultural conditions. It
is also worth noting that in vitro-propagated
plants not only are easily transportable, but
are also subjected to less rigorous controls by
customs, health authorities and quarantine
services since the plantlets are free from
pathogenic microorganisms at the end of the
production cycle.
In comparison with traditional propaga-
tion techniques, however, micropropagation
presents several disadvantages. High costs
are incurred in setting up the laboratories and
purchasing the necessary consumables. Spe-
cialized scientific personnel as well as highly
skilled operators are needed. Furthermore,
somaclonal variation may arise in new adven-
titious shoots when the material is maintained
in vitro too long. Plants with juvenile charac-
teristics may also appear. Most of these nega-
tive aspects can be overcome by adopting
suitable remedies at specific stages of micro-
propagation.
Overall, the process leading from prepa-
ration of the mother trees to acclimatization
of the new plantlets has been subdivided into
five stages, representing the main steps of the
micropropagation cycle (Fig. 9.5/Plate 66).
Preparation of the propagation material
This step involves preparing the mother trees
(or portions of these) from which explants will
be collected for later culturing. The mother
trees must not only possess the genetic charac-
teristics of the cultivars and rootstocks to be
obtained, but must also satisfy specific health
requirements. In particular, they must be virus-
free and have the lowest possible microbial
load on external tissues, in order to ensure
effective explant sterilization. Subjecting the
 Propagation Techniques
mother trees to one or two fungicide applica-
tions some days prior to explant harvesting
has been found to facilitate achievement of
the appropriate health status.
A highly effective technique to sterilize
the starting material consists of rearing the
plants after having satisfied their chilling
requirement. This is performed in a controlled
environment with strict monitoring of light,
temperature and hygiene conditions, where
the buds will be induced to open and form
new shoots. Given their low microbial load,
such shoots can easily be sterilized in large
quantities. Similar results can be obtained
using robust and well-lignified 1-year-old
branches.
It is worth pointing out that the opera-
tions noted below and those carried out in the
231
Fig. 9.4. Hundreds of micropropagated
plantlets in a commercial facility.
following rooting stage should be conducted
in a sterile environment, using a laminar flow
hood and taking care to sterilize working tools
each time.
Explant disinfection and setting up
aseptic culture
This step consists of decontaminating the
explants from any infectious agents that may
be present. The success of this operation
depends on the products utilized, as well as
explant type and size. For peach, a single-node
cutting is taken from actively growing shoots.
The axillary bud is left intact but the axillary
leaf is removed. Alternatively, after removing
232 F. Loreti and S. Morini

Culture establishment

Shoot tip collection

Acclimatized plants

Shoot proliferation

Plantlet acclimatization

Shoot rooting

Fig. 9.5. The main steps of the micropropagation cycle.

larger leaves, a shoot tip explant no longer
than about 1 cm may be used. Explants are
washed under running water for 30 min and
then sterilized with 20% sodium hypochlorite
for 20 min or 0.15% mercury chloride for 8
min. These treatments have proved to be effec-
tive without causing appreciable tissue dam-
age. Following sterilization, explants are rinsed
three or four times in sterile distilled water and
after renewing the explant basal cut they are
transferred to proliferation medium.
Shoot proliferation
The aim of this stage is to promote shoot for-
mation from the axillary buds of each growing
explant. To induce this process, growth regu-
lators (cytokinins) are added to the culture
medium in order to reduce apical dominance.
For peach, the most frequently used growth
regulators are benzylaminopurine (BAP) and
kinetin, varying from a minimum of 0.5–0.6
mg/1 to a maximum of 1–1.2 mg/1 in rela-
tion to the genotype and type of explant. Thus
while it has been found sufficient to add BAP
0.6 mg/1 to MS basal medium (Murashige
and Skoog, 1962) in order to induce prolifera-
tion of ‘Hansen 2168’ and ‘Hansen 536’ root-
stocks (Martinelli, 1985), for ‘P.S.B2’ the best
results are obtained by adding BAP 1.2 mg/1
to WPM medium (Lloyd and McCown, 1980).
N has also been shown to improve both initial
growth and the number of axillary buds
(Loreti et al., 1988).
 Propagation Techniques
Proliferation can also be favourably influ-
enced by the growth cabinet light conditions.
A 4 h light/2 h dark cycle stimulates in vitro
shoot formation and growth more success-
fully than a 16 h/8 h cycle (Morini et al., 1990).
Other factors found to influence growth of
cultures include the photoperiod, light quality
(Loreti et al., 1991), agar type and concentration
(Debergh, 1983) and culture vessel relative
humidity (Sciutti and Morini, 1993). Once the
optimal conditions for proliferation have been
identified, the axillary buds are separated from
each propagule. These are then transferred to
fresh medium (subculture) every 15–20 days.
The proliferation rate may at times be
fairly low during the first subcultures, but
increases noticeably in subsequent subcul-
tures. In peach rootstocks the rate is fairly high,
ranging from a minimum of 10–15 shoots in
some peach × almond clones (Loreti et al.,
1985) to a maximum of 30–35, as in the case of
‘GF 677’ and plum ‘Mr.S. 2/5’.
Rooting and preparation for in vivo growth
Shoots obtained from the proliferation stage
are separated and allowed to root on an aux-
in-enriched medium. Prior to rooting, it may
be necessary to transfer poorly developed
shoots to a medium containing cytokinin at
low concentration in order to stimulate shoot
extension. This operation results in more
robust, homogeneous shoots that root better
and are easier to handle.
The auxins most widely utilized to pro-
mote rooting are IBA, IAA (3-indoleacetic acid)
and NAA, supplied singly or in combination
with one another. Concentration varies from
0.5 to 1–1.5 mg/1. It should be noted that root-
ing shoots may sometimes suffer pronounced
stress due to a sudden change in the type of
hormone used. During root induction in peach,
an increased auxin concentration frequently
causes leaf yellowing although rooting is not
affected. Supplementing the substrate with
polyamines leads to a marked improvement
in quality of the rooted shoots, which thus
develop good roots and normal leaves. The
auxin concentration is of crucial importance.
With excessive auxin, growth of the shoot tip
may be slowed or arrested. This would have a
233
negative effect during the acclimatization
stage, in which the plantlets would no longer
be capable of resuming rapid growth.
Many other factors can influence the
rooting stage, such as donor plant age and
physiological status, juvenile status acquired
through the various subcultures, tempera-
ture, light intensity and light/dark cycles in
the growth chamber, in addition to the vari-
ous components that make up the culture
media (Debergh and Read, 1991).
For most microcuttings, root induction
takes place in vitro. However, after rhizogenic
treatment the microcuttings could be allowed to
root directly in vivo in a protected environment
supplied with fogging or misting equipment.
Generally initiation occurs in the presence of
auxins, but some times they do not appear to be
effective on rooting. Other substances, such as
phenols and gibberellins, would seem to improve
rooting ability. Some authors (Mosella Chancel
et al., 1980) have suggested adopting a three-
phase procedure for in vitro rooting of a peach
rootstock, modifying the hormone composition
of the culture medium. In such an approach,
induction occurs in the presence of auxins (IAA
or NAA) and phenolic substances (phlorizin);
initiation then takes place in the presence of the
same auxins, phenolic substances (quercetin,
rutin) and gibberellin (GA3), while root elonga-
tion is achieved in the absence of auxins and
cytokinins but with gibberellin (GA3).
In day-to-day practice, most micropropa-
gation laboratories have developed their own
experimental protocols. Their procedures are
based on the fundamental principles of micro-
propagation and the most recent acquisitions
of scientific research, but are designed also to
reflect the laboratory’s specific experience in
working on propagation of the different spe-
cies, cultivars or clones.
Transfer and acclimatization of young plants
Young plantlets are next transferred from the
culture vessel to the external environment. The
transfer stage is an extremely delicate part of
the micropropagation technique, as in vitro-
grown plantlets frequently exhibit profound
changes in the structure and functioning of their
organs, potentially leading to stress during
234 F. Loreti and S. Morini
acclimatization. The extent of stress varies as a
function of the species, cultivar or clone and
also the methodology adopted.
After the plantlets have been removed
from the culture vessel, the root system is care-
fully washed and the plantlets are transplanted
to the new medium. They will generally take
more successfully if placed in a greenhouse
equipped with a double tunnel, in similar tem-
perature and humidity conditions as in the
culture vessel. Once vegetative activity has
resumed, temperature and humidity are grad-
ually reduced until external environmental
conditions are reached (Fig. 9.6/Plate 67).
While duration of the acclimatization
phase may vary depending on the genotype,
it is normally around 20–30 days, after which
the plantlets have grown to a height of 15–20
cm and achieved a sufficiently elevated degree
of differentiation to allow transfer to the open
field (Morini and Barbieri, 1986). Planting out
in the field should be conducted in suitable
climatic conditions, to avoid exposing the
plantlets to excessive stress.
Culture medium
The success of micropropagation depends to
a large extent on the type of culture medium
Fig. 9.6. Acclimatization of young micropropagated plantlets.
utilized. The medium is composed of macro-
and microelements, vitamins, growth regula-
tors such as cytokinins, gibberellin and auxins,
and a gelling agent (agar) to ensure that it is
sufficiently compact to maintain the cultures
in a vertical position. Furthermore, since in
vitro cultures have only limited photosyn-
thetic ability on account of the scant available
carbon dioxide and low light intensity, the
medium is also enriched with sucrose as an
energy source.
It is difficult to state exactly what the best
substrate for each rootstock is, as the response
of a culture depends on a multiplicity of fac-
tors that may differ from one laboratory to
another. This means that a given medium
may need to be adjusted in various ways in
relation to the response obtained from a given
rootstock. In general, however, MS medium is
the most widely used and has been exten-
sively modified to adapt it to the various spe-
cies. This medium has given excellent results
with plum ‘Mr.S. 2/5’, ‘GF 677’, ‘GF
43’, ‘Damasco 1869’, ‘San Giuliano 655/2’
(Zuccherelli, 1979), ‘Nemaguard’ (Miller et al.,
1982) and with some peach cultivars (Ham-
merschlag, 1982). Some clones of the ‘I.S.’
series (peach × almond hybrids) have given a
better response with DKW medium (Driver
and Kuniyuki, 1984).
 Propagation Techniques
Problems connected to in vitro propagation
A number of problems arising while the
explants are undergoing in vitro culturing or
during the acclimatization stage may affect
the outcome of this technique. Some of these
difficulties are fairly easy to solve, while oth-
ers are more complex and at times inseparable
at the current state of scientific knowledge.
Problems arising during in vitro culturing
It is not unusual, especially during the spring,
for culture vessels to be contaminated by fungi
and/or bacteria. The cause of contamination
frequently resides in inappropriate handling
by the operator within the laminar flow hood
or inadequate sterilization of tweezers and
scalpels. Such aspects are easily identified
and can be solved simply by greater care in
execution of the manoeuvres. Insufficient
sterilization of the culture substrate may con-
stitute another possible cause. Other contami-
nations of a bacterial nature may appear at
the base of the explant in cultures that have
undergone prolonged in vitro culturing. In
such cases, contamination may conceivably
have arisen from bacteria that are endogenous
to the tissue itself and happen to have found
ideal growth conditions in the particular
medium, although it can be difficult to make
a definite diagnosis.
In seeking to attenuate the damage and
restore explant viability, it is important to
ascertain the intensity of contamination and
the culturing stage during which it is mani-
fested. During the shoot proliferation stage, if
contamination is not severe, a successful rem-
edy is to adopt PPM™ (Plant Preservative
Medium), a heat-stable preservative/biocide
which can be used to effectively prevent or
reduce microbial contamination in plant tis-
sue culture. This compound can slow down
or halt bacterial development and enable a
certain number of subcultures to be per-
formed without appreciable effect on growth.
Alternatively, rooting may be induced early
in an attempt to salvage the greatest possible
number of plantlets.
Tissue vitrification can also represent a
troublesome problem, appearing with vari-
able incidence in relation to the genotype.
235
Vitrification results from excessive water
uptake by tissues and the symptoms are
unmistakeable. If a culture has a predisposi-
tion to this condition, the symptoms will
become more prominent with increasing cul-
ture growth. Attempts to alleviate or eliminate
this problem should therefore focus on reduc-
ing the cytokinin concentration, or using a less
biologically active cytokinin, increased agar
and/or sucrose concentration, or lower cul-
ture vessel relative humidity. Since some
peach rootstocks have notable susceptibility
to vitrification, this phenomenon must be
carefully monitored and kept under control
to avoid excessive loss of cultures.
Finally, explant tissues respond to the
stimulus of the basal cut by producing paren-
chyma cells which appear as small or medi-
um-sized masses of callus. However, the
extent of the response varies with the geno-
type. Explant callus tissue is also stimulated
by the presence of auxin in the proliferation
substrate, even when the auxin concentration
is very low. In general, the presence of callus
at the explant base is undesirable, especially
if a considerably large mass is produced.
Under the stimulation of cytokinin, such cal-
lus could give rise to the differentiation of
adventitious shoots of uncertain genetic char-
acterization. The presence of callus during
the shoot rooting stage likewise represents an
undesirable phenomenon, because it may
adversely affect subsequent plantlet acclima-
tization by favouring the development of
microorganisms present in the transplanta-
tion substrate. In addition, it may undergo
laceration, allowing the onset of rotting at the
rupture site.
Problems arising during plantlet
acclimatization
Shoot cultures may undergo pronounced
changes in tissue structure and growth mech-
anisms as a result of elevated and constant
relative humidity in the culture vessel and
low light intensity in the growth cabinet. Thus,
when rooted shoots are to be removed from
the culture vessel and transferred to the exter-
nal environment, it is essential to allow tis-
sues to adapt gradually in order to avert stress
during the acclimatization stage. First, it is
236 F. Loreti and S. Morini
vital that the roots are placed on a medium
capable of promoting rapid and effective
resumption of growth. This involves careful
choice of the type of peat (Montalti, 1979),
nutrient concentration and environmental
conditions. Relative humidity is the most
important factor that can influence plantlet
acclimatization and it is maintained at high
values by the means of double tunnels, where
relative humidity is better controlled. Resump-
tion of shoot tip activity indicates that the root
system is viable. Shoot acclimatization can
then be initiated by opening the tunnel for
progressively longer periods of time. Great
care should be taken in this operation as the
plantlets are particularly susceptible to water
loss through the leaves, whose anatomic
structure presents incomplete palisade tissue,
larger spaces in the spongiform tissue, lesser
formation of cuticular wax on the leaf blade
and completely open stomata that take a long
time to restore the closure mechanism. But
the greatest danger of all is the onset of severe
water stress that would compromise plantlet
growth or even survival. For species very sus-
ceptible to water stress, it would be advisable
to submit the shoots to an acclimatization
pretreatment a few days before the end of
rooting. At this time, the partial opening of
the in vitro culture vessel enables the plantlets
to become accustomed earlier to the lower
relative humidity of the external environ-
ment. With this procedure, possible leaf water
stresses could be counteracted by water
absorption from the roots, which still are in
the rooting medium.
Other potential problems should not be
overlooked. For example, the plantlets could
be susceptible to microorganisms for which
the warm steamy conditions of the acclimati-
zation glasshouse could represent an ideal
incubator. Therefore, it is essential to ensure
constant monitoring of plantlet health, so that
any preventive treatment that may be required
can be promptly undertaken with the aim
of attenuating or preferably precluding the
onset of such problems.
Over the past few years a procedure has
been developed which has been found highly
effective during the plantlet acclimatization
stage. It consists of the use of mycorrhizae
at the moment of transfer from the culture
vessel to transplantation medium, as mycor-
rhizal symbiosis not only acts as a biofertilizer
but also induces morphological, physiologi-
cal and biochemical modifications in the
plant. Mycorrhizal fungi generally have posi-
tive effects on root system growth, in terms of
better water and nutrient uptake, greater resis-
tance to various types of stress and sometimes
also protection of roots against soil pathogens.
The application of mycorrhizae to microprop-
agation has recently been reviewed (Morini and
Giovannetti, 2004) and suitable nursery proce-
dures should be devised as soon as possible.
9.6 Embryo Culture
Embryo culture is a technique by which an
immature embryo is excised from a seed and
cultured in aseptic culture (Kester and Hesse,
1955, Hartmann et al., 2002) (Fig. 9.7/Plate
68). This technique is very useful to improve
breeding programmes of early-maturing peach
cultivars. In these cultivars, fruit mature before
the embryo has completed its development. In
natural conditions the embryo can abort, but it
can be collected before fruit ripening and
aseptically cultivated on an artificial growth
medium to permit it to germinate and grow
into a plant (Fig. 9.8/Plate 69). The success of
the procedure depends upon the size of the
embryo at collecting time and on the ripening
time of the mother cultivar. Embryos of about
10 mm can be easily cultured on SBH medium
(Smith et al., 1969) and the smaller embryos
(5–10 mm) on MS medium (Ramming, 1990).
Embryos in the globular stage and smaller are
more difficult to culture successfully. For
embryos of 1–5 mm, the sucrose concentra-
tion in the growth medium was found to be
more critical to embryo development than the
hormone content (Ramming, 1990). Ovule
culture, first used in 1981, helped to better
overcome the difficulty encountered with
very small embryos. It is a procedure by
which the ovules are cultured in Stewart and
Hsu medium (Stewart and Hsu, 1977), in the
presence of charcoal and a sucrose and fruc-
tose sugar source, for 4 weeks in the dark, at
25°C. At the end of this period, the embryos
are excised and cultivated on WPM medium
 Propagation Techniques
Fig. 9.7. Excised embryos in aseptic culture.
and 2% sucrose. After a cold treatment at 4°C
for 10 weeks, the embryos are moved to light
and 18°C to induce embryo growth (Ander-
son et al., 2006). Fruit storage before ovule cul-
ture is detrimental to both embryo growth
and survival.
9.7 Stoolbeds and Layering
Stoolbeds and layering are infrequent meth-
ods of vegetative propagation for clonal peach
rootstock multiplication. Although not iden-
tical, both methods are based on the same
principle: promoting vigorous juvenile shoots
that are then partially covered with soil, saw-
dust or other material so that the basal zone is
allowed to develop in the absence of light.
Shoot rejuvenation and etiolation, together
with the special humidity and aeration condi-
tions that occur around the base of the shoots,
favour root induction and development of
adventitious roots. Etiolation may cause starch
and phenolic compounds to accumulate in
shoots and exert a positive influence on rhizo-
genesis (Maynard, 1991; Biricolti et al., 1994).
The two techniques, described in detail
by Hartmann et al. (2002), are widely used for
propagation of clonal rootstocks of apple and
pear, but have been only sporadically adopted
for peach rootstocks. These techniques are
used for the propagation of some rootstocks
such as ‘St. Julian A’, ‘Damas C’ and ‘GF 655/2’,
rootstocks that are no longer used for peach.
237
In addition to the poor results obtained
so far with peach rootstocks (Fiorino, 1968,
1972), these two propagation methods suffer
from a further severe limitation. In an extremely
dynamic nursery market in which the demand
for trees varies from year to year, neither the
stoolbed nor the layering technique allows
production to be planned in relation to mar-
ket requirements. A nursery establishment
designed for production of rooted shoots
obtained with these methods has an average
duration of 12–15 years, which inevitably
implies that there will be years of insufficient
or excess production, depending on the fluc-
tuation of market demand.
9.8 Scion Propagation
Budding and grafting
It can be stated that grafting still constitutes
virtually the only technique adopted for
propagation of peach cultivars. A number of
attempts have been made in recent years to
utilize ‘self-rooted’ varieties of peach and
nectarine, but the use of selected rootstocks is,
in most cases, crucial.
While many different grafting techniques
are available, ‘budding’ is the one most
widely adopted for peach. Wedge, splice-side
and saddle grafts, frequent in the past for
regrafting of adult trees, have now been largely
abandoned. Chip budding, on the other hand,
238 F. Loreti and S. Morini
which is already widely utilized in the USA, is
attracting growing interest among European
nurserymen as well. But regardless of the
type of graft, successful take of the graft
depends fundamentally on three conditions.
1. Graft compatibility of scion and root-
stock, ensuring a solid, durable and efficient
union.
Fig. 9.8. Plantlets generated from
aseptic embryo culture.
2. Correspondence and intimate contact be-
tween scion and rootstock cambial zones.
3. Resumption of cambial activity of both
rootstock (slightly earlier) and scion and begin-
ning of the fusion process immediately after
grafting operations have been performed.
In addition, success of the graft and subse-
quent development of the maiden tree also
 Propagation Techniques
depends on proper choice of the grafting
period and care of the young trees in the nurs-
ery. Moreover, grafting can be used not merely
for normal propagation of the cultivars but
also to promote the recovery of trees affected
by virus infection or mycoplasmas (micro-
grafting).
Budding is the technique that has become
most popular in common nursery practice, on
account of its simplicity and low cost. The
technique consists of the insertion of a vegeta-
tive scion bud into the T-cut stem of the root-
stock. It can be carried out at the end of the
summer (‘dormant bud’), in spring (‘vegeta-
tive bud’) or in mid-June (‘June budding’) in
the northern hemisphere.
The dormant bud graft is performed
starting from the last 10-day period of July up
to mid-September on clonal seedlings or root-
stocks that have reached a stem diameter of
roughly 8–10 mm, when the young trees are
in active growth. It has been noted that some
rootstocks, such as ‘Mr.S. 2/5’, suffer early
cessation of growth. In such cases, in order to
ensure good bud take it is advisable to bring
the budding operations forward by about
10–15 days compared to the seedling, as the
crucial period of cambial activity and thus of
lifting of the bark from the central stem is
shorter (Loreti and Massai, 1995).
The buds to be used for budding must be
taken from virus-free mother tree shoots. Care
must be taken to use those situated in the cen-
tral part of the shoot, which are usually better
shaped and riper. Budding is carried out at
10–15 cm from the ground, after removing the
feathers. The scion should be shortened above
the graft union, in spring, immediately before
bud break. While they remain in the nursery,
the budded trees should be managed accord-
ing to normal cultural practices (removal of
basal feathers, fertilization, irrigation and pes-
ticide application) in order to obtain maiden
trees about 1.2–1.4 m tall.
The vegetative bud graft is less widely
used than budding. It is mainly adopted in
cases when the dormant bud inserted in the
rootstock during the summer failed to take.
When this occurs, the vegetative bud graft is
performed in the following spring.
In this case the buds used for budding
should be taken from portions of shoots
239
removed from the mother trees during the
winter and stored in a moist cool environ-
ment (2–4°C) until required for budding. The
graft is then performed according to the same
technique as the dormant bud graft.
The third type of budding, June budding,
can be adopted only in areas characterized by
a very mild spring, allowing early growth of
the nursery trees. The technique described by
Hartmann et al. (2002) consists of grafting peach
seedlings which, due to appropriate manage-
ment practices, have achieved a suitable size
for grafting by mid-June. The buds, taken
from actively growing shoots, open immedi-
ately after the grafting operation, giving rise
to maiden trees that reach a height of 60–80
cm by October.
One of the main advantages of June bud-
ding is that it allows peach maiden trees to be
obtained within a single year, giving consid-
erable savings on cost compared with the
other two types of bud graft. Furthermore,
the trees possess a more fasciculate root sys-
tem, making them less susceptible to failure
at transplanting.
Another type of graft that is attracting
increasing attention is chip budding. Although
known for many years, it has only recently
been adopted by European nurserymen. In
addition to allowing greater flexibility in time
of execution, in many cases it is preferred to
the dormant bud graft (T-budding) in order
to supply trees characterized by both a stron-
ger union at the point of the graft and also a
more balanced and uniform development of
the maiden trees.
Finally, wedge, splice-side and saddle
grafts that were widely used in the past are sel-
dom used today. Previously they were per-
formed to repeat grafts in cases when the tree
did not correspond to the desired variety, or to
rapidly convert an existing orchard to one with
newer scion cultivars. Although this operation
is technically valid, it is justifiable only if the
cost of converting the orchard is lower than that
of purchasing and establishing new trees.
Micrografting
This is a grafting technique performed in
vitro, which has recently been introduced to
240 F. Loreti and S. Morini
favour the recovery of certain fruit tree spe-
cies from virus infection without causing any
impairment of genetic and varietal character-
istics. Adopted for the first time to restore
virus-affected citrus trees to health, it was
subsequently applied to some peach (Barba
et al., 1995), almond and cherry cultivars sen-
sitive to prolonged heat treatment. In peach,
satisfactory recovery percentages have been
obtained for some viral diseases such as Chlo-
rotic leaf spot virus and Prunus necrotic ring spot
virus, while in the case of Prune dwarf virus
results varied according to the viral strain
(Navarro et al., 1982). The technique consists
of decapitating an in vitro-grown plantlet,
after which a 0.2–0.3 mm portion of the tip of
the cultivar to be grafted is placed on it. The
micrografted plant is maintained in culture
for 5–6 weeks prior to transplantation. The
success of this technique depends on numer-
ous important factors including the pheno-
logical status of the tip donor mother plant,
the rootstock and substrate utilized, the
micrograft growth conditions, and plantlet
acclimatization to the external environment.
Herbaceous grafting
This is a technique in which a shoot tip col-
lected from in vitro culture is grafted on to a
very young seedling rootstock (only a few
millimetres in diameter). Cuttings may be
produced by traditional techniques or from
micropropagation and are raised in plastic
containers (Preka and Cherubini, 2001). This
type of grafting is performed ex vitro and is
much simpler to execute than micrografting.
Thus it is a technique that has higher propa-
gation efficiency and in Italy some micro-
propagation laboratories have found it suitable
for mass propagation. The rootstock is cut back
at about 10 cm from the soil, incised longitudi-
nally, and top-grafted with a scion that has the
basal end cut to a V-shaped wedge. The scion
is then inserted in the stock and tied with
Parafìlm. Grafting establishment will occur in
a greenhouse, at high relative humidity and
controlled temperature. The major advantage
of this type of grafting is that the new plant
can be obtained in 1 year.
9.9 Outlook
As reported above, the evolution of peach
propagation techniques has produced signifi-
cant changes over time. Today, plants of high
genetic, sanitary and agronomic quality are
available. Nevertheless, a dynamic nursery
industry exists that is aimed particularly at
advanced propagation systems to improve
the economic and production efficiency and
to provide plants with superior characters. At
present, it is very difficult to predict further
advancements in nursery techniques. Micro-
propagation was certainly the greatest inno-
vation of the last few decades but it also
showed some limitations with a number of
genotypes that even today are propagated by
traditional techniques. Better application of
micropropagation for peach rootstocks is
strictly related to the possibility of setting up
more efficient, less expensive and more func-
tional procedures.
A new stimulus to peach rootstock prop-
agation could occur by somatic embryogene-
sis. This technology is based on the capacity
of differentiated cells to resume meristematic
activity under specific growth conditions
controlled by bioreactors and to produce
somatic embryos with structures similar to
zygotic embryos. Some species propagated
by this technique have shown the ability to
produce large numbers of new plants, in less
time and space than can be attained by micro-
propagation. The major problems currently
are: the scarce response to somatic embryo-
inducing treatments shown by most woody
fruit species; the difficulty of somatic embryos
to convert into plants; and the rarity of
somatic embryo-producing plants that are
true to type. In the case of peach rootstocks,
somatic embryogenesis could develop into a
valid alternative to traditional techniques,
but it is not possible to evaluate its applicabil-
ity until the above-mentioned aspects are
clearly defined.
Thanks to the uninterrupted advances of
molecular biology, research on genes control-
ling root induction may facilitate the develop-
ment of transgenic plants by genetic engineering
or transformation by A. tumefaciens (Radchuk
and Korkhovoy, 2005).
 Propagation Techniques 241

Future research on mechanisms of adven- physiological, genetic and molecular aspects

titious rooting may enable the use of cuttings
for propagating woody species as a suitable
alternative to other techniques. Cutting prop-
agation is actually easy to apply and needs
relatively simple facilities and equipment that
most nursery operations already have.
Today an intense research activity is
progressing to determine the underlying
involved. Thus, we may expect that deeper
knowledge on rooting mechanisms will be
acquired in a reasonably short time. This
should allow further improvements of the
vegetative propagation techniques currently
applied, producing new and more advanced
propagation systems.

References

Anderson, N., Byrne, D. and Ramming, D. (2006) In ovule culture success as affected by sugar source and
fruit storage duration in nectarine. Acta Horticulturae 713, 89–92.
Barba, M., Cupidi, A., Loreti, S., Faggioli, F. and Martino, L. (1995) In vitro micrografting: a technique to
eliminate peach latent mosaic viroid from peach. Acta Horticulturae 386, 531–535.
Bartolini, G. and Fabbri, A. (1982) Una doppia bagnatura per aumentare la sopravvivenza delle barbatelle di
pesco. Rivista Ortoflorofrutticoltura Italiana 4, 323–329.
Bartolini, G. and Fiorino, P. (1978) Gli interventi sulle piante madri per migliorare la radicazione delle talee.
In: Seminario sul vivaismo e controllo della rizogenesi mediante fitoregolatori. Editrice Tecnico Scienti-
fica, Pisa, Italy, pp. 9–25.
Bassuk, N.L. and Howard, B.H. (1981) A positive correlation between endogenous root-inducing cofactor
activity in vacuum-extracted sap and seasonal changes in rooting of M 26 winter apple cuttings. Journal
of Horticultural Science 56, 301–312.
Bhattacharya, N.C. (1988) Enzyme activity during adventitious rooting. In: Davis, T.D., Haissig, B.E. and
Sankhla, N. (eds) Adventitious Root Formation in Cuttings. Dioscorides Press, Portland, Oregon, pp.
88–97.
Biricolti, S., Mariotti, P. and Mattii, G.B. (1990) Anatomical studies on the collapse of rooted cuttings and
grafts in peach. Advances in Horticultural Science 3, 159–162.
Biricolti, S., Fabbri, A., Ferrini, F. and Pisani, P.L. (1994) Adventitious rooting in chestnut: an anatomical inves-
tigation. Scientia Horticulturae 59, 197–205.
Blazich, F.A. (1988) Mineral nutrition and adventitious rooting. In: Davis, T.D., Haissig, B.E. and Sankhla, N.
(eds) Adventitious Root Formation in Cuttings. Dioscorides Press, Portland, Oregon, pp. 61–69.
Debergh, P.C. (1983) Effect of agar brand and concentration on the tissue culture medium. Physiologia Plan-
tarum 59, 270–276.
Debergh, P.C. and Read, P.E. (1991) Micropropagation. In: Debergh, P.C. and Zimmermann, R.H. (eds) Micro-
propagation: Technology and Application. Kluwer Academic Publishers, Dordrecht, The Netherlands,
pp. 1–14.
Di Vaio, C., Buccheri, M. and Cirillo, C. (2001) Impiego della solarizzazione e della bromurazione del terreno
nel reimpianto del pesco: effetto sullo sviluppo vegetativo di astoni in vivaio. Italus Hortus 8, 39–40.
Driver, J.A. and Kuniyuki, A.H. (1984) In vitro propagation of Paradox walnut rootstock. HortScience 19,
507–509.
Erez, A. and Yablowitz, Z. (1981) Rooting of peach hardwood cuttings for the meadow orchard. Scientia Hor-
ticulturae 15, 137–144.
Fabbri, A. (1977) La propagazione del pesco per talea di ramo. Rivista Ortoflorofrutticoltura Italiana 1, 126–136.
Fadl, M.S. and Hartmann, H.T. (1967) Relationship between seasonal changes in endogenous promoters and
inhibitors in pear buds and cutting bases and the rooting of pear hardwood cuttings. Proceedings of the
American Society for Horticultural Science 91, 96–112.
Fiorino, P. (1968) Nuove tecniche per ottenere barbatelle di pesco. II – Ricerche sulla ‘propaggine per trincea’.
Rivista Ortoflorofrutticoltura Italiana 52, 205–212.
Fiorino, P. (1972) Nuove tecniche per ottenere barbatelle di pesco. IV – Ricerche sulla ‘margotta di ceppaia’.
Rivista Ortoflorofrutticoltura Italiana 56, 100–104.
Fiorino, P. and Loreti, F. (1965) La propagazione del pesco per talea legnosa. In: Atti del Congresso del Pesco.
Tipo-lito degli Stimmatini, Verona, Italy, pp. 483–495.
242 F. Loreti and S. Morini

Fiorino, P. and Loreti, F. (1987) Propagation of fruit trees by tissue culture in Italy. HortScience 22, 353–358.
Fiorino, P. and Mattii, G.B. (1992) The role of prunasin in ‘collapse’ of rooted peach cutting. Advances in
Horticultural Science 1, 11–14.
Fiorino, P. and Vitagliano, C. (1968) Nuove tecniche per ottenere barbatelle di pesco: III ‘Ulteriori ricerche
sulla nebulizzazione’. Rivista Ortoflorofrutticoltura Italiana 6, 779–795.
Guerriero, R. and Scalabrelli, G. (1984) Effect of stratification duration on seed germination of several peach
line rootstocks. Acta Horticulturae 173, 185–190.
Hammerschlag, F. (1982) Factors affecting establishment and growth of peach shoots in vitro. HortScience 17,
85–86.
Hartmann, H.T., Kester, D.E., Davies, F.T. Jr and Geneve, R.L. (2002) Plant Propagation. Principles and Prac-
tices, 7th edn. Prentice-Hall, Upper Saddle River, New Jersey.
Katan, J. and DeVay, J.E. (1991) Soil solarization: historical perspectives, principles, and uses. In: Katan, J. and
DeVay, J.E. (eds) Soil Solarization. CRC Press Inc., Boca Raton, Florida, pp. 23–39.
Kester, D.E. and Hesse, C.O. (1955) Embryo culture of peach varieties in relation to season of ripening. Pro-
ceedings of the American Society for Horticultural Science 65, 265–273.
Koornneef, M., Bentsinka, L. and Hilhorstb, H. (2002) Seed dormancy and germination. Current Opinion in
Plant Biology 51, 33–36.
Lang, G.A. (1996) Plant Dormancy: Physiology, Biochemistry, and Molecular Biology. CAB International,
Wallingford, UK.
Lloyd, G.B. and McCown, B.H. (1980) Commercially feasible micropropagation of mountain laurel, Kalmia
latifoglia, by use of shoot tip culture. Combined Proceedings of the International Plant Propagators Society
30, 421–427.
Lorenzi, R. and Ceccarelli, N. (1978) Variazioni stagionali del potenziale rizogenp delle talee e sue modifica-
zioni con fitoregolatori. In: Seminario sul vivaismo e controllo della rizogenesi mediante fitoregolatori.
Editrice Tecnico Scientifica, Pisa, Italy, pp. 27–36.
Loreti, F. and Massai, R. (1995) Orientamenti per la scelta dei portinnesti del pesco. L’Informatore Agrario 32,
37–42.
Loreti, F. and Pisani, P.L. (1982) Physiological and technical factors affecting rooting in woody species. In:
Proceedings of the XXIst International Horticultural Congress. International Society for Horticultural
Science, Wageningen, The Netherlands, pp. 295–309.
Loreti, F., Morini, S. and Grilli, A. (1985) Rooting response of BS B2 and G.F. 677 rootstocks cutting. Acta
Horticulturae 173, 261–269.
Loreti, F., Morini, S. and Concetti, S. (1988) Effect of potassium and nitrogen concentration on growth of
peach shoots cultured in vitro. Acta Horticulturae 227, 311–317.
Loreti, F., Muleo, R. and Morini, S. (1991) Effect of light quality on growth of in vitro cultured organs and
tissues. The International Plant Propagation Society Combined Proceedings 40, 615–623.
Marini, R.P. (1983) Rooting of semihardwood peach cuttings as affected by shoot position and thickness.
HortScience 18, 718–719.
Martinelli, A. (1985) Factors affecting in vitro propagation of the peach–almond hybrids ‘Hansen 2128’ and
‘Hansen 536’. Acta Horticulturae 173, 237–244.
Maynard, B.K. (1991) Stock plant etiolation and stem banding effect on the auxin dose–response of rooting in
stem cuttings of Carpinus betulus L. ‘Fastigiata’. Journal Plant Growth Regulation 10, 305–311.
Miller, G.A., Coston, D.C., Denny, E.G. and Romeo, M.E. (1982) In vitro propagation of Nemaguard peach
rootstock. HortScience 17, 194.
Montalti, G. (1979) Influenza del substrato nell’ambientamento di alcune piante da frutto derivate da coltura
in vitro. In: Atti dell’Incontro su: Tecniche di Colture in vitro per la propagazione su vasta scala delle
specie ortoflorofrutticole. F&F Parretti Grafiche, Florence, Italy, pp. 119–126.
Morel, G.M. (1964) Tissue culture. A new means for clonal propagation of orchids. American Orchid Society
Bullettin 33.
Morini, S. and Barbieri, C. (1986) La propagazione in vitro: fattori che condizionano l’acclimatazione delle
piantine. L’Informatore Agrario 2, 67–72.
Morini, S., Fortuna, P., Sciutti, R. and Muleo, R. (1990) Effect of different light–dark cycles on growth of fruit
tree shoots cultured in vitro. Advances in Horticultural Science 4, 163–166.
Morini, S. and Giovannetti, M. (2004) La micorrizazione, una biotecnologia per la produzione in vivaio di
piante arboree da frutto di elevata qualita. Frutticoltura 12, 43–46.
 Propagation Techniques 243

Mosella Chancel, L., Macheix, J.J. and Jonard, R. (1980) Les conditions du microbuturage in vitro su pecher
(Prunus persica Batsch): influences combinées des substances de croissance et de divers compose phé-
nolique. Physiologie Vegetal 18, 597–608.
Murashige, T. and Skoog, F. (1962) A revised medium for rapid growth and bioassays with tobacco tissue
cultures. Physiologia Plantarum 15, 473–497.
Navarro, L., Llacer, G., Cambra, M., Arregui, J.M. and Juarez, J. (1982) Shoot-tip grafting in vitro for elimina-
tion of viruses in peach plants (Prunus persica Batsch). Acta Horticulturae 130, 185–192.
Nicotra, A. and Moser, L. (1996) Two new plum rootstocks for peach and nectarines: penta and tetra. Acta
Horticulturae 451, 269–271.
Preka, P. and Cherubini, S. (2001) Tecniche di innesto erbaceo per la propagazione di piante arboree da
frutto. Frutticoltura 5, 39–41.
Radchuk, V.V. and Korkhovoy, V.I. (2005) The rolB gene promotes rooting in vitro and increases fresh root
weight in vivo of transformed apple scion cultivar ‘Florina’. Plant Cell Tissue and Organ Culture 81,
203–212.
Ramming, D.W. (1990) The use of embryo culture in fruit breeding. HortScience 25, 393–398.
Scalabrelli, G. and Couvillon, G.A. (1986) The interaction between IBA treatment and other factors in rooting
and establishment of peach hardwood cuttings. Acta Horticulturae 179, 855–862.
Sciutti, R. and Morini, S. (1993) Effect of relative humidity in in vitro culture on some growth characteristics
of a plum rootstock during shoot proliferation and on plantlet survival. Advances in Horticultural
Science 4, 153–156.
Smith, C.A., Bailey, C.H. and Hough, L.F. (1969) Methods for germinating seeds of some fruit species with
special reference to growing seedlings from immature embryos. New Jersey Agricultural Experiment
Station Bulletin 823.
Sorce, C., Paolicchi, F., Ceccarelli, N., Lorenzi, R. and Picciarelli, P. (2004) Relation between cell redox status
and adventitious rooting in cuttings of Cupressus sempervirens. In: SIFV-SIGA Joint Congress, Lecce, Ita-
ly, 15–18 September, vol. XLIII Annual Congress, p. 34.
Stewart, J.M. and Hsu, C.L. (1977) In-ovulo embryo culture and seedling development of cotton (Gossypium
hirsutum L.). Planta 137, 113–117.
Vasil, I.K. and Vasil, V. (1972) Totipotency and embryogenesis in plant cell and tissue cultures. In vitro Cellular
and Developmental Biology – Plant 8, 117–125.
Zuccherelli, G. (1979) Moltiplicazione in vitro dei portinnesti clonali di pesco. Rivista di Frutticoltura 41,
15–20.
 10 Carbon Assimilation, Partitioning and
Budget Modelling

T.M. DeJong1 and A. Moing2

1Department of Plant Sciences, University of California, Davis, California, USA
2Unite de Physiologia Vegetable, INRA, Bordeaux, France

10.1 Introduction 244

10.2 Carbon Assimilation 245
Leaf photosynthetic characteristics 245
Fruit photosynthesis 246
Canopy carbon assimilation 247
10.3 Carbon Partitioning 247
Carbon partitioning within source leaves 247
Carbon transport from the leaves 248
Carbon partitioning within fruit 248
Carbon partitioning within vegetative growing sink organs 250
Carbon partitioning and mobilization in perennial parts 250
Carbohydrate partitioning at the whole-plant level 251
Effect of biotic stress 254
10.4 Modelling the Carbon Economy of Peach 254
Models of fruit growth and carbon economy 254
Modelling tree growth and carbon economy 255
10.5 Future Directions 256

10.1 Introduction assimilation and the efficiency and effective-

Virtually all of the energy and nearly all of the
dry matter that a plant obtains to support
growth and development comes from solar
energy and the process of photosynthesis.
Ultimately, crop productivity is dependent on
plant photosynthetic efficiency and the effi-
ciency with which photosynthates are parti-
tioned. Peach production is no exception, and
peach tree growth and productivity are
strongly dependent on the efficiency of carbon
ness of the distribution and use of carbohy-
drates for tree growth maintenance and the
production of quality fruit. In fact, most of the
cultural operations that a fruit grower does
are in some way connected to influencing
either the assimilation of carbon or the distri-
bution and use of carbohydrates in the plant.
Therefore, it is important to have a basic
understanding of the processes involved in
carbon assimilation and the distribution and
use of carbohydrates in peach trees.

244 © CAB International 2008. The Peach: Botany, Production and Uses
(eds D.R. Layne and D. Bassi)
 Carbon Assimilation, Partitioning and Budget Modelling
Research specific to the leaf photosyn-
thetic processes in peach is not as plentiful as
for many other crops. However, there is ade-
quate information to know the general behav-
iour of peach leaves with respect to most of
the environmental and cultural factors that
are known to influence leaf and canopy pho-
tosynthesis of crop plants. Interestingly, while
research on peach leaf photosynthesis is not
as plentiful as for some other crops, there is
probably more research reported on the details
of carbohydrate partitioning/distribution than
for any other fruit tree species.
Carbon partitioning and utilization by
the various organs (sinks) of the plant has been
extensively studied at several levels. There
have been numerous studies concerned with
the import and metabolism of specific carbo-
hydrates in various sink tissues while others
have focused on the total amounts of carbon
that get distributed to the various parts and/
or functions of the plant and the factors that
control that distribution. In this review we
consider the uptake and metabolism of carbon
of specific sink tissues and then look at the
broader aspects of carbon partitioning and
utilization at the whole-plant level.
10.2 Carbon Assimilation
Leaf photosynthetic characteristics
Like virtually all other temperate deciduous
tree fruit species, peach leaves exhibit the
classic characteristics of leaves having the
C3 biosynthetic pathway of photosynthesis
(Brown, 1994). Maximal reported rates of leaf
CO2 assimilation are around 20–22 µmol/
m2/s (DeJong et al., 1989) while many
researchers have reported lower values (Flore
and Lakso, 1989). Leaf carboxylation efficiency
appears to saturate well beyond intercellular
CO2 concentrations of 300 ml/l (DeJong, 1983;
Rosati et al., 1999) and estimated intercellular
CO2 concentrations under normal conditions
have been reported to be between 180 and
250 ml/l (DeJong, 1983, 1986a). Individual
leaf CO2 assimilation appears to approach
light saturation at 700–1000 mmol/m2/s
depending on the leaf position in the canopy
245
(DeJong, 1983; Kappel and Flore, 1983; Rosati
et al., 1999). Walcroft et al. (2002) reported
effects of changes in absorbed photosyntheti-
cally active radiation (PAR) on peach leaf
morphological and physiological properties.
Leaf mass per unit area and leaf N concentra-
tion were non-linearly related to absorbed
PAR, and there was a weak linear relationship
between leaf N concentration per unit mass
and absorbed PAR. The leaf ribulose bis-
phosphate carboxylase (RUBISCO) content
increases with the amount of N applied to
young trees (Nii et al., 1997). In the canopy,
spatial variability in photosynthetic capacity
results from acclimatization to varying absorbed
PAR as the crown develops, acclimatization
being driven principally by changes in leaf
mass per unit area rather than the amount or
partitioning of leaf N (Rosati et al., 2000; Wal-
croft et al., 2002). At the molecular level, the
RNA abundance of a light-harvesting type II
chlorophyll a/b-binding protein is lower in
mature shaded leaves compared with sun-
exposed leaves (Bassett and Callahan, 2003).
The leaf photosynthesis apparatus of peach
appears to be fairly insensitive to changes in
temperature between 20°C and 32°C but can
be substantially inhibited by temperatures
above and below this range (Crews et al.,
1975; Reyes-Lopez, 1984; Girona et al., 1993).
Leaf stomatal conductance has been
reported to be fairly closely coupled to leaf pho-
tosynthesis in peach and the coupling increases
with exposure to water stress (Reyes-Lopez,
1984; Girona et al., 1993). Although peach is a
mesophytic species, leaf CO2 assimilation
appears to be fairly tolerant of water stress in
the field and leaves can maintain more than
50% of their photosynthetic capacity down to
leaf water potentials as low as –2.0 MPa (Girona
et al., 1993). However, there have been reports
of sensitivity to drought treatments in potted-
plant experiments (Rieger and Duemmel,
1992) and this sensitivity is not always related
to leaf water potential, suggesting the involve-
ment of non-hydraulic signalling (Rieger et al.,
2003). Peach leaves also appear to be relatively
tolerant to ozone air pollution in comparison
with several other temperate deciduous fruit
tree species (Retzlaff et al., 1991).
Although it is widely accepted that leaf age
strongly affects the photosynthetic capacity
246 T.M. DeJong and A. Moing
of individual leaves of most species (Flore
and Lakso, 1989), there is very little data
available specifically for peach. Peach tree
canopies grow quite rapidly and peach leaf
CO2 assimilation capacity is strongly influ-
enced by internal canopy shading (Marini
and Marini, 1983; DeJong and Doyle, 1985;
DeJong et al., 1989; Rosati et al., 1999; LeRoux
et al., 2001). Therefore it is difficult to separate
leaf age effects from changes in leaf light
exposure in naturally growing field canopies.
It is apparent that leaves in the most exposed
parts of the tree canopies tend to maintain
relatively constant photosynthetic capacities
in the absence of major stress during the mid-
dle part of the growing season from May to
September (in the northern hemisphere)
(DeJong, 1986a).
Probably the most controversial reported
influence on leaf CO2 assimilation rate in
peach is the factor of crop load or proximity
of fruit to leaves. Early work by Crews et al.
(1975) reported approximately 20% higher
rates of photosynthesis in leaves in close
proximity to fruits compared with leaves
measured further down the same branch and
that leaf photosynthesis peaked during the
period of peak fruit growth (i.e. maximum
fruit sink strength). CO2 exchange measure-
ments in these experiments used cut branches
brought in from the field. It is unclear what
influence cutting a branch might have on sink
effects on photosynthesis with this experi-
mental set-up. At about the same time, Chal-
mers et al. (1975) reported that peach leaf
photosynthesis can vary by as much as 50%
due to sink demand corresponding to fruit
development patterns in the field. However,
their conclusions are questionable because it
is likely that their measurements were light-
limited and influenced by spreading of the
tree due to crop load (Westwood and Gerber,
1958). Subsequent research has reported
11–15% higher leaf CO2 assimilation rates in
fruited versus defruited mature peach trees
during the peak period of fruit growth
(DeJong, 1986a). The higher CO2 assimilation
rates were attributed to increases in leaf sto-
matal conductance rather than mesophyll
conductance or other leaf photosynthetic
properties. Other research reported similar
crop load effects on leaf stomatal conductance
with nectarines (DeJong, 1986b). Leaf photo-
synthesis can be strongly reduced within 48 h
of fruit removal, indicating that sudden loss
of fruit sink strength has a greater effect on
CO2 assimilation than long-term presence of
the fruit (Mandre et al., 1995). The relatively
modest fruit effects on photosynthesis reported
in these studies on peach in comparison with
apple (Hansen, 1970; Avery, 1975) are prob-
ably due to the fact that peach is grown on
vigorous rootstocks and there are many alter-
native vegetative sinks for carbohydrates in
most situations (Flore and Lakso, 1989).
There have been dramatic changes in
peach genetic characteristics through domes-
tication and breeding over the past couple of
centuries. However, crop improvement efforts
in peach have primarily focused on fruit qual-
ity traits and timing of fruit maturity. The
resulting changes have apparently had little
effect on leaf photosynthetic characteristics
of peach (Quilot et al., 2004a), although there
appears to be substantial variability in the
related Prunus species, which are primarily related
to tree canopy density and distribution of leaf
N (DeJong, 1983; Rieger and Duemmel, 1992).
Fruit photosynthesis
In other rosaceous species such as apple,
Vemmos and Goldwin (1994) suggested that
flower photosynthesis might play an impor-
tant role in flower growth and fruit set. How-
ever, no data are available for peach flowers.
Green peach fruit are capable of photosynthe-
sis but the majority of CO2 assimilation that
occurs in fruit even when they are small
merely serves to offset a portion of the CO2
produced by respiration (Pavel and DeJong,
1993b). However, peach fruit photosynthesis
is not insignificant and has been estimated to
account for between 5% and 9% of the total
carbohydrate budget of developing peach
fruit, depending on the position of the fruit in
the canopy (Pavel and DeJong, 1993b). Although
these estimates of the photosynthetic contri-
bution of peach fruits are lower than has been
estimated for other species (Bazzaz et al., 1979;
Kappes, 1985; Birkhold et al., 1992), the contri-
bution of fruit photosynthesis to the overall
carbon economy of peach fruit production
 Carbon Assimilation, Partitioning and Budget Modelling
should not be ignored. Phosphoenolpyruvate
carboxylase, involved in C4 and CAM (crassu-
lacean acid metabolism) plant photosynthesis,
is also involved in non-photosynthetic pro-
cesses in all plant species and may contribute to
re-assimilation of respired CO2 in peach fruit
tissues as in apple (Blanke and Lenz, 1989).
Canopy carbon assimilation
Like with all crops (Charles-Edwards, 1982),
canopy photosynthesis depends on the amount
of light intercepted by the plant as well as
individual leaf photosynthetic capacity. Light
interception can be strongly influenced by
training system (Giuliani et al., 1998, Gross-
man and DeJong, 1998). Whole-canopy CO2
assimilation in peach has been measured
directly (Giuliani et al., 1998) and calculated
through modelling techniques (Grossman
and DeJong, 1994b; Giuliani and Magnanini,
2002). Whole-canopy assimilation rates of peach
trees can be more than 25 g CHO/m2/day
during peak periods of the growing season.
This is similar to values reported for herba-
ceous species (Ng and Loomis, 1984). These
rates are possible, in spite of the relatively mod-
est maximum leaf photosynthetic potential of
peach (20–22 mmol/m2/s) compared with
other C3 crop species (Brown, 1994), because
of the high leaf area index of peach trees
(Chalmers and van den Ende, 1975; Gross-
man and DeJong, 1998). Peach canopies gen-
erally have a high leaf area index after the
initial spring growth flush (6–8 m2 of leaf area
per square metre of ground area) and indi-
vidual leaf photosynthetic capacity varies
greatly within the canopy. This variation has
been shown to correspond to leaf N content
per unit leaf area, leaf specific weight and leaf
light exposure (DeJong, 1982; Kappel and Flore,
1983; Rosati et al., 2000; LeRoux et al., 2001).
Rosati et al. (2002) have shown that there is
also a linear relationship between calculated
daily net CO2 assimilation and daily leaf light
interception of individual peach leaves. Fur-
thermore they demonstrated that daily canopy
photosynthetic radiation use efficiency of crop
canopies can be estimated from measurements
of the photosynthetic light response of leaves
at the top of the canopy and the incident PAR
247
above the canopy. These relationships should
make estimates of canopy photosynthesis of
peach trees under non-stressed field condi-
tions much easier than they were previously
thought to be.
10.3 Carbon Partitioning
Carbon partitioning within source leaves
Like most tree species in the Rosaceae, the pri-
mary products of photosynthesis in mature
peach leaves are sorbitol, sucrose and starch
with generally lesser amounts of glucose and
fructose (Bieleski, 1982; Moing et al., 1992).
Although there is significant variation in the
ratio of sorbitol to sucrose content in leaves of
different peach cultivars, the leaf sorbitol con-
tent of all cultivars is higher than sucrose. The
sorbitol:sucrose ratio of most cultivars exam-
ined is about 2–3:1 (Escobar-Gutiérrez and
Gaudillère, 1994). In mature leaves sorbitol
biosynthesis is accomplished by an NADPH-
dependent aldose-6-phosphate reductase that
catalyses the conversion of the photosynthetic
product glucose-6-phosphate to sorbitol-6-
phosphate (Hirai, 1981). A cDNA clone encod-
ing this enzyme was first isolated from apple
(Kanayama et al., 1992). The sorbitol-6-phosphate
is converted into sorbitol by a specific sorbitol-
6-phosphate phosphatase (Grant and ap Rees,
1981). The expression of aldose-6-phosphate
reductase was studied in terms of protein and
RNA in developing peach leaves, confirming
its association with the source function
(Sakanishi et al., 1998). Aldose-6-phosphate
reductase activity and protein level in leaves
also varies seasonally. In apple, aldose-6-
phosphate reductase activity appears to be
regulated by inorganic phosphate and diva-
lent cations (Zhou et al., 2003a). Since sorbitol-
6-phosphate phosphatase activity is inhibited
by inorganic phosphate and sorbitol (Zhou
et al., 2003b), it may be a regulatory step in
sorbitol biosynthesis. It appears that the rela-
tive concentrations of sorbitol, sucrose, glu-
cose and fructose are fairly closely regulated
in leaves photosynthesizing under constant
steady-state laboratory conditions while starch
increases steadily during the photoperiod
248 T.M. DeJong and A. Moing
(Moing et al., 1992). In leaf discs, the sorbitol
content and the level of mRNA encoding
putative aldose-6-phosphate reductase were
higher under low temperature (Deguchi et al.,
2002). The control mechanisms of cellular car-
bon partitioning into sorbitol versus sucrose
in rosaceous species are unknown. However,
there is some evidence that sucrose content is
more tightly regulated than sorbitol in peach
leaves (Moing et al., 1994) and that sucrose
can be used for conversion into sorbitol within
the leaf (Moing et al., 1992). Recently, in apple,
the relationship between sucrose synthesis and
sorbitol metabolism has begun to be unrav-
elled. It has been shown that leaf sucrose phos-
phate synthase, catalysing one of the final
steps of sucrose synthesis in the cytosol of
photosynthetic cells, is inhibited by sorbitol-
6-phosphate (Zhou et al., 2002).
Early studies in apple showed that source
leaves have minimal capacity for sorbitol
degradation (Grant and ap Rees, 1981; Loescher
et al., 1982). Therefore, the fate of sorbitol in
source leaves is storage or export through
phloem. However, this conclusion should be
re-examined since one isoform of NAD-
dependent sorbitol dehydrogenase involved
in sorbitol breakdown was recently shown to
be expressed in source leaves of apple (Park
et al., 2002). As with other sugar alcohols, sor-
bitol can be stored temporarily in cell vacu-
oles of source leaves or in sinks, suggesting
the presence of a tonoplast-bound alditol car-
rier as noted for mannitol-synthesizing spe-
cies (Greutert et al., 1998).
Environmental factors can influence the
relative amounts of the various carbohydrates
in the leaves. Factors that favour higher rates
of photosynthesis, such as increasing light
exposure, tend to enhance the flux of all three
major carbohydrates in peach leaves, with
starch increasing the most at the highest light
levels (Escobar-Gutiérrez and Gaudillère,
1997). Studies with apple and peach indicate
that factors causing sudden increases in pho-
tosynthate supply relative to demand, such as
lengthening photoperiod, CO2 enrichment or
fruit removal, increase leaf sorbitol and starch
content while sucrose is less affected (Wang
et al., 1998, 1999; Ali and Nii, 1999). Drought
stress has been reported to enhance the produc-
tion of sorbitol relative to sucrose and starch in
both apple (Wang et al., 1996) and peach
(Escobar-Gutiérrez et al., 1998; Rieger et al.,
2003). However, this behaviour does not appear
to contribute to increased osmotic adjustment
in peach (Escobar-Gutiérrez et al., 1998).
Carbon transport from the leaves
The rate of carbohydrate turnover in peach
leaves appears to be substantially less than
reported for herbaceous species and this
appears to correspond with a lower carbon
export rate (Moing et al., 1992). Although minor
concentrations of sorbitol, glucose, sucrose
and fructose can be extracted from xylem of
petioles of mature leaves and subtending
shoots, the majority of carbohydrate export
from leaves occurs in the phloem as sorbitol
and sucrose (Moing et al., 1997). The sorbitol:
sucrose ratio in the phloem sap generally cor-
responds to the ratio found in leaf tissue, i.e.
about 2–3:1. Although the possibility of sym-
plastic phloem loading of sorbitol has not
been ruled out, it is likely that apoplastic
phloem loading through an active transport
pathway is the most important means of
exporting carbohydrates from mature leaves
in peach (Moing et al., 1997). H+-sucrose sym-
porters, which are involved in active uptake
of sucrose across the plasma membrane, have
an essential role in phloem loading of sucrose-
transporting species (Lemoine, 2000). H+-
polyol symporters certainly play an additional
role in phloem loading in sorbitol-transporting
species, as shown for mannitol symporters in
celery (Noiraud et al., 2001). Later, phloem
unloading in young leaves or fruit probably
involves another apoplastic step with sorbitol
transporters involved in partitioning and
accumulation of sorbitol in sink tissues as
reported in sour cherry (Gao et al., 2003).
Carbon partitioning within fruit
Early in fruit development, glucose and fruc-
tose are the most prominent sugars but they
decline gradually while sucrose concentration
increases rapidly and is dominant during
stage III or the final swell period (Moriguchi
et al., 1990, 1992; Pavel and DeJong, 1993a;
 Carbon Assimilation, Partitioning and Budget Modelling
Genard and Souty, 1996; Vizzotto et al., 1996;
Lo Bianco et al., 1999a; Genard et al., 2003).
Sorbitol and starch concentrations change less
and remain relatively low throughout fruit
growth. These findings are consistent with
the concept that early spring growth in peach
trees is largely dependent on stored carbohy-
drates that are mobilized after the release
from dormancy and transported to the growing
sinks through the xylem (as hexoses) (Loescher
et al., 1990). Subsequently, phloem is reacti-
vated and differentiated (Evert, 1960) and new
photosynthates are transported primarily as
sorbitol and sucrose (Moing et al., 1994).
Peach domestication and hybridization
appear to have resulted in large increases in
fruit size (potential fruit growth; Quilot et al.,
2002) and soluble sugar concentration as shown
by comparative studies between wild species
and commercial cultivars (Robertson et al.,
1988; Moing et al., 2003). Concerning com-
mercial cultivars, one study reporting genetic
variation in sugar content of clingstone peach
cultivars showed that sucrose was the domi-
nant sugar even in the early stages of fruit
growth (Brooks et al., 1993). It would be inter-
esting to determine if the differences that have
been reported for early fruit sugar concentra-
tions between the freestone (Moriguchi et al.,
1990, 1992; Pavel and DeJong, 1993a; Genard
and Souty, 1996; Vizzotto et al., 1996; Lo Bianco
et al., 1999a; Genard et al., 2003) and clingstone
(Brooks et al., 1993) cultivars are consistent
characteristics for the two fruit types.
Our understanding of carbohydrate
metabolism during later stages of peach fruit
growth is less clear and there is confusion
about whether peach fruit preferentially use
sorbitol or sucrose as the primary source of
transported carbohydrate. Several groups of
researchers have attempted to follow the pat-
tern of enzymes involved in peach fruit car-
bohydrate metabolism to try to discern the
major source of carbon supplying fruit growth
and to understand the mechanism of the rapid
rise in sucrose concentration during peach
development. Moriguchi et al. (1990) attrib-
uted the rapid rise of sucrose concentrations
in peach fruits to increased sucrose synthase
activity (operating in the synthesis direction;
Moriguchi and Yamaki, 1988). They reported
very low levels of sucrose phosphate synthase
249
activity and did not find decreasing acid
invertase activity while sucrose accumulated,
so they concluded that neither of these enzymes
could account for the changes in sucrose
concentrations. Moriguchi et al. (1990) also
reported that sorbitol appeared to be con-
verted to glucose by sorbitol oxidase and that
both sucrose and sorbitol could be used to
support fruit growth. Hubbard et al. (1991)
reported similar enzyme activities for peach
fruit, but in addition to increases in sucrose
synthase activity they also detected sucrose
phosphate synthase activity that could help
account for the rise in sucrose concentrations.
Subsequent research has reported decreasing
sucrose synthase and invertase enzyme activ-
ities and fairly constant sucrose phosphate
synthase activity while sucrose accumulates
in the peach fruit (Vizzotto et al., 1996). The
latter authors also conducted glucose and
sucrose uptake studies with fruit mesocarp
tissue and reported higher uptake rates for
glucose than sucrose. They concluded that
their results were consistent with the possibil-
ity that sorbitol and sucrose could be taken
up by peach mesocarp tissue by either an
apoplastic or symplastic route of phloem
unloading. However, the recent cloning of
sorbitol transporters in sour cherry fruit (Gao
et al., 2003) is in favour of an apoplastic step.
The mode of phloem unloading in peach fruit
may depend on the fruit development stage
as suggested in tomato (Ho, 1992).
Recent research on the carbohydrate
metabolism of fruit and vegetative sinks in
peach has increased confusion over the role of
sorbitol as a source of carbohydrate for sup-
plying fruit growth. In contrast to Moriguchi
et al. (1990), Lo Bianco et al. (1999a) reported
that they could detect no sorbitol oxidase in
peach mesocarp tissue during any stage of
development and sorbitol dehydrogenase
activity was present only during the last stage
of fruit growth and then only in small amounts.
Similar to reports of previous researchers, Lo
Bianco et al. (1999a) noted that sucrose syn-
thase and invertase activities were high early
in the fruit development period and were low
during stage III of fruit growth. However,
data on general enzyme activity should be
considered with care due to the possible pres-
ence of several enzyme isoforms. For instance,
250 T.M. DeJong and A. Moing
in apple fruit, four cDNAs encoding isoforms
of NAD-dependent sorbitol dehydrogenase
have been cloned (Park et al., 2002). The four
corresponding genes were highly expressed
at the late stage of apple fruit development.
Moreover, post-translational regulation of
invertases and sucrose synthases may be
involved as suggested in other fruits (Tanase
et al., 2002; Zhou et al., 2003c). On the other
hand, Yamada et al. (2001), who cloned an NAD-
dependent sorbitol dehydrogenase from peach
fruit and studied its expression during devel-
opment, proposed that the activity of NAD-
dependent sorbitol dehydrogenase is regulated
by the transcription of the gene.
Genard et al. (2003) used an entirely dif-
ferent approach for analysing the changes in
peach fruit sugar concentrations in response
to assimilate supply, metabolism and dilu-
tion. They used the sugar model of Genard
and Souty (1996), which predicts carbon par-
titioning into sucrose, sorbitol, glucose and
fructose in peach mesocarp tissue by calculat-
ing rates of sugar transformation and com-
puting concentrations of the various sugars,
and compared the results with measured con-
centrations of carbohydrates in fruits. Based
on their analysis, fruit carbohydrates differed
in sensitivity to assimilate supply, metabo-
lism and dilution (fruit volume). Sucrose was
highly sensitive to assimilate supply and
dilution effects but not subject to much meta-
bolic transformation. Sorbitol was calculated
to be the most important carbohydrate involved
in peach fruit metabolism and consequently
its concentration was generally low in spite of
its high proportion in the assimilate supply.
Glucose and fructose represented a transitory
storage/metabolic pool. Their assessment
appears to be in conflict with the hypothesis
of Lo Bianco et al. (1999a) regarding the role of
sorbitol as a carbon source for fruit growth
but provides an insightful synthesis that is in
agreement with most of the other literature
on peach fruit carbohydrate metabolism.
Carbon partitioning within vegetative
growing sink organs
Meristematic tissues in vegetative peach buds
appear to have higher uptake capacity for
glucose than sucrose or sorbitol (Maurel et al.,
2004). However, Lo Bianco et al. (1999a) found
sorbitol dehydrogenase activities in active
root tips to be high and in shoot tips they
were positively correlated with shoot growth
rates. This factor and the corresponding peri-
ods of peak sink and enzyme activities led
these researchers to hypothesize that, in peach
trees, sucrose is the major form of carbohy-
drate used for fruit growth whereas sorbitol
has a predominant role in supporting vegeta-
tive growth. There seems to be general agree-
ment that sorbitol is a primary source of
carbohydrates for vegetative sinks such as root
tips, growing shoots (Lo Bianco et al., 1999b)
and immature leaves (Moing et al., 1992).
These findings appear to agree with similar
studies with apple (Loescher et al., 1982;
Yamaki and Ishikawa, 1986).
Carbon partitioning and mobilization in
perennial parts
It is well known that the majority of the over-
wintering storage of carbohydrates in fruit
trees is in the form of starch that is found in
the bark and wood of stems, trunk and roots
(Stassen et al., 1981, 1982; Tromp, 1983; Oliveira
and Priestley, 1988). Raffinose and stachyose
are also stored in Prunus perennial parts dur-
ing winter (Gaudillère et al., 1992). Although
the concentration of starch and soluble carbo-
hydrates is often higher in the bark, storage in
woody tissue accounts for the greatest pro-
portion of stored carbohydrates in the tree
(Stassen et al., 1982). There is little informa-
tion specific for peach regarding partitioning
to long-term storage or starch metabolism
within sink organs. Descriptive studies have
shown strong seasonal patterns in starch con-
centration with the highest levels occurring in
the dormant season (Stassen et al., 1981; Jor-
dan and Habib, 1996; Yano et al., 2000). Starch
storage that occurs during the growing sea-
son is often considered a residual sink for car-
bohydrates that is active after all the growing
sinks are satisfied (Grossman and DeJong,
1994b). However, there must be specific
mechanisms that regulate starch metabolism
in storage tissue and certainly this activity
 Carbon Assimilation, Partitioning and Budget Modelling
should be considered to be as much of a sink
as the growth of specific organs. This is true
especially when one considers that winter
storage and spring mobilization of carbohy-
drates is essential for tree survival, flowering
and initial organ growth.
Although the essential nature of mobili-
zation of stored carbon is readily apparent
when one considers the annual cycle of a peach
tree, the relative contribution of this part of
the carbon budget to the functioning of the
whole plant is generally underappreciated. In
the stem of peach, 50% of starch was hydroly-
sed between October and December (Marquat
et al., 1999). The early study of Petrov and
Manolov (1973) using 14CO2 labelling in
autumn demonstrated accumulation and
mobilization of carbon reserves in peach. 13C
is used nowadays as a tracer to study the dis-
tribution of 13C-labelled assimilates during
dormancy following assimilation of 13CO2
during the preceding autumn. It is also used
to follow mobilization and translocation of
13C- labelled assimilates in the following spring
but there has been limited use of this tech-
nique with peach. During the winter dormant
season, phloem transport in temperate decidu-
ous trees is thought to become non-functional
(Evert, 1960) and whatever long-distance trans-
port of carbohydrate occurs is apparently
through the xylem (Loescher et al., 1990). Sor-
bitol has been reported to be the primary car-
bohydrate present in the xylem of peach trees
during the winter months but it declines in
the spring when the hexoses, glucose and
fructose, become more prevalent (Loescher
et al., 1990; Maurel et al., 2004). As noted pre-
viously, meristematic tissues in peach buds
and young fruit appear to have higher uptake
capacity for glucose than sucrose or sorbitol
(Vizzotto et al., 1996; Maurel et al., 2004). It has
been reported that sorbitol is the primary car-
bohydrate source for the growth of young
leaves (Moing et al., 1992) and shoot and root
tips (Lo Bianco et al., 1999b). It would be inter-
esting to know if this is true for the early
spring flush of growth as well, since sorbitol
is apparently not found in large amounts in the
early spring xylem transport stream. It is gen-
erally recognized that the first several days of
spring growth is dependent on carbohydrates
mobilized from storage and thereafter there is
251
thought to be a transition to dependence on
current assimilates (Priestley, 1970).
Carbohydrate partitioning at the
whole-plant level
Over the past couple of decades the concept
that carbohydrate partitioning at the whole-
plant level is primarily driven by growth and
development of individual organs has become
widely accepted (Gifford and Evans, 1981;
Watson and Casper, 1984; Ho, 1988; Marcelis,
1994; Weinstein and Yanai, 1994; Lacointe,
2000). Grossman and DeJong (1994a) used
this concept in the development of the PEACH
model and later DeJong (1999) outlined four
principles for applying this concept to logi-
cally understand carbon partitioning in peach
(and other fruit) trees.
The first principle is that a tree is a collec-
tion of semi-autonomous organs and each organ
has a genetically determined, organ-specific devel-
opment pattern and growth potential. Although
much emphasis is often placed on consider-
ing plants as highly integrated organisms, the
concept of semi-autonomy among organs is
widely recognized (White, 1979; Harper, 1980;
Watson and Casper, 1984; Sprugel et al., 1991).
Indeed, the primary morphological features
that distinguish one species or cultivar (in the
case of peach) from another are at the organ
or sub-organ level (i.e. fruit or leaf shape and
size, floral characteristics, etc.), not at the whole-
plant level. Furthermore, although variation
exists, the developmental patterns and growth
rates of individual organs under specific envi-
ronmental conditions are generally predict-
able and have been modelled. Models have
been developed for the growth of peach fruit
(DeJong and Goudriaan, 1989; Pavel and
DeJong, 1993c; Grossman and DeJong, 1995b;
Genard and Huguet, 1996; Genard and Souty,
1996), shoots and branches (Costes et al., 1993;
Grossman and DeJong, 1995a; Genard et al.,
1998; Lescourret et al., 1998) and roots (Bidel
et al., 2000). Although tree pruning and train-
ing can drastically alter the shape of peach
trees, they generally have very little effect on
individual organ characteristics other than
those explained by changes in the local
microenvironment of the organs or changes
252 T.M. DeJong and A. Moing
in the availability of carbohydrates due to the
proximity of other sinks.
The fact that there appears to be some
level of branch autonomy (Sprugel et al., 1991)
in peach trees further reinforces this first prin-
ciple. Branch autonomy tends to functionally
isolate some sinks from sources of carbohy-
drates. When sinks are manipulated through
pruning or fruit thinning to create an appar-
ent abundance of photosynthate in one part
of the tree and an undersupply somewhere
else, the carbon does not freely move to the
location of greatest demand. When one scaf-
fold of Y-shaped peach trees was defruited
the remaining fruit on the fruited scaffold
benefited very little from the carbon that
should have been available for fruit growth
from the defruited scaffold (Marsal et al.,
2003). Interestingly, scaffold diameter growth
appeared to be one of the sinks that benefited
most from the removal of fruit, while root
growth was only marginally affected. There is
much to be learned about the movement of
carbohydrates within the context of the whole
tree. The role of branch autonomy in the early
spring, when much of the carbon used for
growth is mobilized from storage in the root,
trunk and major branches and is presumably
transported in the xylem, is also very poorly
understood.
Carbon partitioning at the branch level
has been studied in peach explicitly with
radioactive tracer studies (Corelli Grappadelli
et al., 1996) and by manipulating leaf number
and fruit load in isolated branches (Genard
et al., 1998). Implicit conclusions about carbon
partitioning within shoots have also been
drawn from fruit thinning studies to deter-
mine optimal fruit positioning for fruit size
(Spencer and Couvillon, 1975; Marini and
Sowers, 1994). These studies support the idea
that fruit are strong sinks for carbon within
shoots but their influence on where recently
fixed carbon goes varies substantially within
the local context of the stem unit.
The second principle is that the genetic
potential of an organ is activated or deactivated by
organ-specific, endogenous and/or environmental
signals. The semi-autonomous nature of indi-
vidual organs is further demonstrated by the
fact that individual organs on a tree can be
experimentally activated by manipulating
factors that stimulate the growth of specific
organs independently from processes occur-
ring in organs elsewhere on the tree. For
instance, exposing individual buds on a branch
to rest-breaking treatments can induce bud
break in those buds while similar buds on
other parts of the tree remain inactive (Chan-
dler, 1942). Similarly, grafting multiple culti-
vars with differing chilling requirements on
to one trunk will not influence the inherent
chilling exposure required for activation by
the branch of each specific cultivar. Also,
removing the apical meristem on a shoot will
promote the activation of growth of lateral
buds on the remaining part of the shoot while
buds on other shoots are unaffected (Harris,
1983). Although the exact mechanisms of the
environment and/or endogenous signals that
activate growth are not fully understood, the
primary site of activation is clearly at the
organ or sub-organ level. This is certainly one
area where hormones play key roles in influ-
encing carbon partitioning at the whole-tree
level, as suggested by data on hormone con-
centrations in xylem sap (Sorce et al., 2002).
The third principle is that after an organ is
activated, current environmental conditions and
genetic growth potential interact to determine
conditional organ growth capacity. Although
often overlooked, ambient temperature is
probably the single most important environ-
mental factor influencing organ growth. The
importance of temperature is related to the
strong dependence of respiration on tempera-
ture. All real plant organ growth is dependent
on metabolic activity and enzyme function,
and these processes are linked to respiration.
Plant respiration generally has a temperature
response quotient (Q10) of about 2 (doubles
for every 10°C increase in temperature between
5°C and 35°C; Amthor, 1989) and peach is no
exception (Pavel and DeJong, 1993c; Gross-
man and DeJong, 1994a). Therefore, condi-
tional growth capacity of any organ is highly
dependent on ambient temperature. The con-
ditional growth capacity of peach fruits grow-
ing under near-optimal field conditions has
been modelled for several peach cultivars
using relative growth rate functions (DeJong
and Goudriaan, 1989; Pavel and DeJong,
1993c; Grossman and DeJong, 1995a; Berman
et al., 1998). That other environmental factors
 Carbon Assimilation, Partitioning and Budget Modelling
such as water status can also have a substan-
tial effect on organ growth is well documented
(Bradford and Hsiao, 1982). Extension growth
of peach shoots has been successfully mod-
elled by considering temperature and dynamic
changes in shoot water status (Berman and
DeJong, 1997a; Basile et al., 2003). Although
peach fruit growth is quite sensitive to water
stress it is important to distinguish between
growth in fresh versus dry matter since the
former is much more sensitive than the latter
(Berman and DeJong, 1997b; Girona et al.,
2004). Nutrient availability also can strongly
influence conditional organ growth capacity
because certain nutrients are required as con-
stituents for the growing organs. Accordingly,
Saenz et al. (1997) have demonstrated that N
availability can influence developmental pat-
terns of peach fruit.
Finally, realized organ growth is a consequence
of conditional organ growth capacity, resource avail-
ability (assimilate and nutrient supply) and inter-
organ competition for those resources. When
conditional growth capacity of an organ is
set, organ growth should proceed at a rate
equal to the conditional growth capacity as
long as transport is not limited and enough
resources (carbohydrates) are available to
support that organ’s growth and the growth
of all other competing organs. However, if the
tree does not have enough carbohydrate to
support the conditional growth capacity of all
organs or carbohydrate transport within the
tree is limited, then the growth of an individual
organ will be a function of its ability to compete
for available carbohydrates with other organs.
Since flowering and pollination are not major
problems in the more productive peach-
growing regions, most peach cultivars set
very heavy fruit loads. Therefore lack of avail-
able assimilates and inter-fruit competition
for carbohydrates is generally the primary
factor limiting realized fruit growth in mature
peach trees and fruit thinning is essential to
manage this competition (Dorsey and McMunn,
1928; Cain and Mehlenbacher, 1956; Johnson
and Handley, 1989; Pavel and DeJong, 1993b;
DeJong and Grossman, 1995; Grossman and
DeJong, 1995b; Costa and Vizzotto, 2000).
Certainly there are some limitations to carbo-
hydrate transport within the tree (DeJong and
Grossman, 1995; Marsal et al., 2003) but these
253
are difficult to quantify specifically. There is
substantial evidence that peach fruit growth
can compete effectively for carbohydrates with
shoot, trunk and root growth when the crop
loads are high and all fruit are considered as a
collective sink (Proebsting, 1958; Grossman
and DeJong, 1995a; Marsal et al., 2003). But
there is some evidence to the contrary when
pruning systems stimulate excess vegetative
shoot growth (Grossman and DeJong, 1998).
There is also clear documentation of the capac-
ity of individual fruit organs to compete with
each other and/or vegetative sinks at the local
branch level (Genard et al., 1998). To compli-
cate things further, the competitive ability of
fruit for carbohydrates appears to vary with
the stage of fruit development (DeJong and
Grossman, 1995).
Upon examining these four principles
for understanding carbon partitioning it
becomes apparent that phenological patterns
of organ growth are the principal determi-
nants of carbon partitioning in peach trees.
When experiments are conducted involving
different crop load treatments or some other
treatment that dramatically favours the growth
of one type of organ over others, biomass data
collected at the end of the season appear to
indicate that some organs are in direct compe-
tition with others (Proebsting, 1958; Chalmers
and van den Ende, 1975). However, when
seasonal patterns of growth are analysed, it is
apparent that direct competition between dif-
ferent organ types is limited by temporal sep-
aration of growth activities (DeJong et al.,
1987; Miller and Walsh, 1988; Rufat and DeJong,
2001; Berman and DeJong, 2003). Generally,
in late-maturing peach cultivars shoot and
root growth is the dominant sink shortly after
bud break in the spring. This period is fol-
lowed by a peak of fruit growth and then
there is a resurgence of root growth (L. Pace,
unpublished results) and shoot diameter
growth after harvest (Grossman and DeJong,
1995a; Berman and DeJong, 2003). It is inter-
esting that breeding efforts to create cultivars
with early fruit ripening times has apparently
interfered with the natural temporal separa-
tion of dominant sink activities in peach trees.
The dominant period of fruit growth of early-
maturing peach cultivars often coincides
directly with the early peak of shoot growth.
254 T.M. DeJong and A. Moing
This has increased the competition between
fruit and shoot growth, resulting in decreased
yield potential (DeJong et al., 1987; Grossman
and DeJong, 1995a). There is also some evi-
dence that selection for early-maturing culti-
vars has involved selection for decreases in
total fruit growth potential and dry matter
content and these factors also account for dif-
ferences in yield potential between early- and
late-maturing cultivars (Berman et al., 1998).
The selection for early-maturing fruit has also
increased the competition for carbohydrates
between organs within the fruit such that
seed development corresponds with the
period of flesh enlargement (Pavel and
DeJong, 1993a) as well as increasing the indi-
vidual fruit relative growth rates, so that the
tree cannot support as many fruits at one time
(Grossman and DeJong, 1995a,b).
Effect of biotic stress
Biotic stress may also modify carbon parti-
tioning through effects on sources and/or
sinks. Photosynthesis is drastically reduced
in powdery mildew-affected trees (Toma
et al., 2003). Leaf injuries caused by silver mite
decrease photosynthetic rate and have effects
on fruit quality in the current year and in the
next year (Kondo and Hiramatsu, 1999).
Increased partitioning of dry weight and car-
bohydrate fractions from shoots to roots and
shifts in proportional composition of individ-
ual sugars occurred in ‘Nemaguard’ rootstocks
infested with nematodes (Olien et al., 1995). If
parasitism was severe, reduced levels of car-
bohydrates in the above-ground part of the
tree could result in tree injury or death at lev-
els of environmental and biological stresses
that do not injure healthy trees.
10.4 Modelling the Carbon
Economy of Peach
A host of environmental (biotic and abiotic)
and endogenous factors can simultaneously
influence carbon assimilation and partition-
ing within leaves, transport processes between
sources and sinks, activation and conditional
growth capacity of individual organs, and the
relative ability of organs to compete with each
other for carbon. The only way to function-
ally understand and quantify the importance
of these factors in an integrated system is to
develop computer simulation models of that
system. The work in this area with peach has
been focused principally in two areas: models
of fruit growth and of the whole tree.
Models of fruit growth and carbon economy
Conners (1919) first described the double-
sigmoid growth curve of peach fruit almost a
century ago. In 1989, DeJong and Goudriaan
demonstrated that the double-sigmoid growth
curve could be modelled with relative growth
rate functions and developed a quantitative
computer simulation model of the carbohy-
drate requirements for peach fruit growth.
This work estimated the respiratory growth
coefficient for peaches to be 0.309 mg CO2/
mg fruit dry weight and allowed for subse-
quent models to calculate total carbohydrate
requirements for peach fruit growth and dev-
elopment (DeJong and Walton, 1989). Fruit
respiration accounts for 16–20% of the carbo-
hydrate required for producing a peach fruit.
The carbohydrate demand for an individual
peach fruit peaks just before harvest and can
be as high as 1.3 g CHO per fruit per day.
DeJong and co-workers focused on the devel-
opment of simulation models for estimating
the carbohydrate demand of the cohort of
larger-sized fruit on a tree (thus quantifying
the fruit potential carbohydrate demands).
Meanwhile, Genard and co-workers con-
centrated on models that take into account
the great variation in fruit growth among the
whole population of fruit in a typical peach
canopy. In 1993, Genard and Bruchou pre-
sented a multivariate approach to modelling
peach fruit growth that investigated the effect
of leaf:fruit ratio and between- and within-
tree variability in growth of individual fruits.
Interestingly, their model showed that size
differences among fruits within shoots accoun-
ted for more than half of the total variability
in fruit size while leaf:fruit ratio was not a
major factor. However, subsequent models
documented the importance of leaf:fruit ratio
 Carbon Assimilation, Partitioning and Budget Modelling
in determining the size of fruits on well-
illuminated girdled shoots (Genard et al.,
1998; Lescourret et al., 1998). Research by
Genard and Souty (1996) produced a very
interesting model of mesocarp sugar contents
in relation to peach fruit growth. Using fruit
flesh dry weight data and a set of literature-
based conceptual assumptions for carbon
partitioning into various sugars, these authors
successfully modelled fruit flesh sucrose, glu-
cose, fructose and sorbitol contents and con-
centrations at different crop loads. They also
used the model to estimate the contribution
of each sugar to relative sweetness during the
last 50 days of fruit development. Their model
indicates that sucrose is the most important
sugar in determining fruit sweetness at fruit
maturity but that fructose is more important
earlier in fruit development. Subsequently
this model was linked to a model of fresh
mass growth of peaches to simulate the effect
of water supply on fruit sugar concentrations
(Genard and Huguet, 1997) and to a model of
carbon acquisition and partitioning for simu-
lating the effects of light interception and fruit
thinning on peach sweetness (Genard et al.,
1999). Most recently this model was used to
further distinguish the effects of assimilate
supply, dilution and metabolism on fruit sugar
concentrations (Genard et al., 2003). These
models substantiate that sorbitol is the most
important carbohydrate used for metabolism
in peach fruit, while sucrose does not undergo
metabolic transformation but is subject to a
dilution effect during fruit development. Glu-
cose and fructose constitute a transitory stor-
age pool and their concentrations are closely
related to metabolism. Collectively these mod-
els provide a substantial resource for under-
standing the influence of crop load and various
environmental factors such as light exposure,
water and temperature on the dynamics of
growth and quality of peach fruit.
Modelling tree growth and carbon economy
The fruit growth model of DeJong and Goud-
riaan (1989) was based on the premise that
fruit growth potential is genetically controlled
and can be approximated by modelling the
255
relative growth rates of fruit on lightly cropped
(sink-limited) trees. This led to the develop-
ment of a carbon-based simulation model of
reproductive and vegetative growth of whole
peach trees. By conducting a series of field
experiments to collect fruit and vegetative
growth and respiration data under varying
crop load scenarios, Grossman and DeJong
(1994a, 1995a,b,c) quantified the seasonal
interactions between the various organ types,
which were used to develop sub-models of
seasonal carbohydrate demand by the fruit,
shoots, major branches and trunks of mature
peach trees. The sub-models of carbohydrate
demand were linked with a sub-model of
canopy photosynthesis to estimate carbohy-
drate supply and the result was a whole-tree
model that predicted the daily partitioning of
carbon into various organ types (Grossman
and DeJong, 1994b). The PEACH model esti-
mated that for the high-density central leader
planting where the data were collected, an
individual tree assimilated 33 kg CHO between
bloom in early March and the beginning of
leaf-fall in mid-October. This was equivalent
to 4.1 kg CHO/m2/year. For a normally
cropped, August-ripening cultivar (‘Cal Red’),
the model estimated that approximately one-
third of the annual carbohydrate budget went
to maintenance respiration while two-thirds
went to biomass growth and growth respira-
tion. Simulated fruit growth, above-ground
vegetative growth and root activity each con-
sumed about one-third of the carbohydrate
used for growth. However, the model also
clearly showed that these patterns of carbohy-
drate use are dependent on crop management
practices such as fruit thinning. Subsequently
the PEACH model was used to estimate the
effect of differences in seasonal air temperature
patterns on carbon assimilation, crop growth
and respiration and yield (DeJong et al., 1996).
This work showed that it is likely that whole-
tree canopy photosynthesis may be enhanced
in very warm climates but these conditions
may not lead to increased yields since higher
maintenance respiration rates may actually
reduce the amount of carbohydrate available
for growth. Berman et al. (1998) have also
used this model to estimate the effect of fruit
maturity date on yield potential of process-
ing peaches grown in California. This work
256 T.M. DeJong and A. Moing
predicted that the yield potential of cultivars
developed with fruit maturity dates earlier
than the second week in July (the harvest
period of the current earliest processing cling-
stone peaches in California) will be reduced
by approximately 1.8 t/ha for each day that
harvest is advanced unless the physiological
characteristics of the trees that govern carbon
partitioning are also substantially altered.
The original PEACH model had two major
limitations. The first was that carbohydrate
partitioning to root growth and development
was not explicitly modelled and this problem
still persists. However, the general patterns
and quantities of dry matter partitioning to
root growth and respiration that the model
predicts are in general agreement with data
on dry matter distribution in peach trees
(Chalmers and van den Ende, 1975; Miller
and Walsh, 1988; Grossman 1993; Rufat and
DeJong, 2001). All of these reports indicate
that root biomass of peach trees accounts for
about one-quarter of the total tree biomass,
depending on crop load and other manage-
ment factors. Trees on size-controlling root-
stocks appear to allocate a slightly higher
proportion of their dry mass to roots (Salvati-
erra et al., 1998; Basile et al., 2003) but very
limited data are available on this question.
Kubota et al. (1990) monitored the movement
of 13C-labelled photosynthates into peach
roots and found a dwarfing rootstock (Prunus
tomentosa) received a greater proportion of
labelled carbon 120 h after feeding than did a
vigorous rootstock (Prunus persica). This label-
ling pattern fits biomass distribution data;
however, it may be unwise to draw too strong
conclusions since there were limited data and
P. tomentosa dwarfing appears to be a result of
partial scion incompatibility.
The second major limitation of the cur-
rent PEACH model is that all the individual
organs of each type in a tree are modelled
together as a mean organ of that type. Thus
the proximity of individual organs to sources
or sinks of carbohydrate or location-depen-
dent environmental stimuli such as light are
not accounted for. Allen et al. (2002) have
begun developing a new, context-sensitive,
visual graphics-based approach to explicitly
model carbon transport and partitioning
within peach tree canopies. This model takes
advantage of L-systems technology (Prusinkie-
wicz, 1998) to keep track of every organ that
develops on a simulated graphic representa-
tion of a tree. Algorithms are being developed
to govern the movement of carbohydrate into
and out of each organ as the tree develops
through several growing seasons. This research
is still in the formative stage but it holds sub-
stantial promise as a platform for beginning
to understand the integrated processes involved
in carbohydrate partitioning to individual
organs within the context of the whole tree. It
should also provide a framework for incorpo-
rating the quality modelling of peach fruits
(Genard et al., 2003) into integrated whole-
tree models of carbohydrate partitioning in
peach.
10.5 Future Directions
There are two areas of scientific investigation
that are likely to have major impacts on our
understanding of carbon assimilation, parti-
tioning and budget modelling of peach trees
in the next decade. These are the techniques
of molecular genetics and physiology and the
rapid advances in information technology.
Peach is considered as the model species for
structural and functional genomics of Rosa-
ceae (Abbott et al., 2002; http://www.genome.
clemson.edu/gdr/; see Chapter 4). Genomic
resources including expressed sequence tags
and a physical genetic map are being devel-
oped. Molecular physiology is just beginning
to enlighten scientists about the regulation of:
(i) enzyme activity within cells of source and
sink tissues; and (ii) transport between source
and sink tissues. This work will provide a more
thorough understanding of the regulation of
source and sink activities at the cellular level
and unlock the current mysteries regarding the
role of the various carbohydrates in the over-
all scheme of the carbon budget. One very
interesting question in this regard is the pur-
pose of having two major transport carbohy-
drates (sucrose and sorbitol) in the rosaceous
species.
The potential for application of the recent
developments in computer technology has yet
to be efficiently used in efforts to synthesize
 Carbon Assimilation, Partitioning and Budget Modelling 257

the knowledge about numerous aspects of
carbon assimilation and partitioning into an
integrated model of peach tree growth and
carbon economy. Much work can be done at
several levels of organization by just using
current knowledge, let alone incorporating
the new information that is being developed
using new molecular techniques. New sensor
technology is now available to monitor
hundreds of facets of growth and environ-
mental parameters simultaneously, but this
work will require an integrated intellectual
framework to make it efficient. Advancing
efforts with functional/structural models to
couple plant architectural models with physi-
ological models should make that feasible in
the near future (LeRoux et al., 2001). More-
over, the combination of genetic and ecophys-
iological models should also help to unravel
the mechanisms of carbon partitioning
involved in the elaboration of fruit quality
(Quilot et al., 2002, 2004b).

References

Abbott, A.G., Lecouls, A.C., Wang, L., Georgi, L., Scorza, R. and Reighard, G. (2002) Peach: the model genome
for Rosaceae genomics. Acta Horticulturae 592, 199–203.
Ali, K. and Nii, N. (1999) Fruiting effects on diurnal changes in sorbitol and starch contents in peach leaves.
Journal of the Japanese Society of Horticultural Science 68, 739–745.
Allen, M.T., DeJong, T.M. and Prusinkiewicz, P. (2002) Using L-systems to model carbon transport and parti-
tioning in developing peach trees. Acta Horticulturae 584, 29–35.
Amthor, J.S. (1989) Respiration and Crop Productivity. Springer, New York.
Avery, D.J. (1975) Effects of fruits on photosynthetic efficiency. In: Pereira, H.C. (ed.) Climate and the Orchard.
Commonwealth Bureau of Horticulture and Plantation Crops, Research Review No. 5. Commonwealth
Agricultural Bureaux, Farnham Royal, UK, pp. 110–112.
Basile, B., Marsal, J. and DeJong, T.M. (2003) Daily shoot extension growth of peach trees growing on root-
stocks that reduce scion growth is related to daily dynamics of stem water potential. Tree Physiology 23,
695–704.
Bassett, C.L. and Callahan, A.M. (2003) Characterization of a type II chlorophyll a/b-binding protein gene
(Lhcb2*Pp1) in peach. II. mRNA abundance in developing leaves exposed to sun or shade. Tree Physiology
23, 473–480.
Bazzaz, F.A., Carlson, R.W. and Harper, J.L. (1979) Contribution to reproductive effort by photosynthesis of
flowers and fruits. Nature 279, 554–555.
Berman, M.E. and DeJong, T.M. (1997a) Crop load and water stress effects on daily stem growth in peach
(Prunus persica). Tree Physiology 17, 467–472.
Berman, M.E. and DeJong, T.M. (1997b) Diurnal patterns of stem extension growth in peach (Prunus persica):
temperature and fluctuations in water status determine growth rate. Physiologia Plantarum 100, 361–370.
Berman, M.E. and DeJong, T.M. (2003) Seasonal patterns of vegetative growth and competition with reproduc-
tive sinks in peach (Prunus persica). Journal of Horticultural Science & Biotechnology 78, 303–309.
Berman, M.E., Rosati, A., Pace, L., Grossman, Y. and DeJong, T.M. (1998) Using simulation modeling to esti-
mate the relationship between date of fruit maturity and yield potential in peach. Fruit Varieties Journal
52, 229–235.
Bidel, L.P.R., Pages, L., Rivère, L.M., Pelloux, G. and Lorendeau, J.Y. (2000) Mass Flow Dyn I: a carbon trans-
port and partitioning model for root system architecture. Annals of Botany 85, 869–886.
Bieleski, R.L. (1982) Sugar alcohols. In: Loewus, F.A. and Tanner, W. (eds) Encyclopedia of Plant Physiology.
Vol 13A. Plant Carbohydrates I. Intracellular Carbohydrates. Springer, Berlin, pp. 158–192.
Birkhold, K.T., Koch, K.E. and Darnell, R.L. (1992) Carbon and nitrogen economy of developing rabbiteye
blueberry fruit. Journal of the American Society for Horticultural Science 117, 139–145.
Blanke, M.M. and Lenz, F. (1989) Fruit photosynthesis – a review. Plant, Cell & Environment 12, 31–46.
Bradford, K.J. and Hsiao, T.C. (1982) Physiological responses to moderate water stress. In: Lange, O.L., Nobel,
P.S., Osmond, C.B. and Ziegler, H. (eds) Physiological Plant Ecology II. Water Relations and Carbon
Assimilation. Springer, Berlin, pp. 263–324.
Brooks, S.J., Moore, J.N. and Murphy, J.B. (1993) Quantitative and qualitative changes in sugar content of
peach genotypes [Prunus persica (L.) Batsch.] Journal of the American Society for Horticultural Science
118, 97–100.
258 T.M. DeJong and A. Moing

Brown, R.H. (1994) The conservative nature of crop photosynthesis and the implications of carbon fixation
pathways. In: Boote, K.J., Bennett, J.M., Sinclair, T.R. and Paulsen, G.M. (eds) Physiology and Determina-
tion of Crop Yield. American Society of Agronomy, Inc., Madison, Wisconsin, pp. 211–219.
Cain, J.C. and Mehlenbacher, R.J. (1956) Effects of nitrogen and pruning on trunk growth in peaches. Proceed-
ing of the American Society for Horticultural Science 67, 139–143.
Chalmers, D.J. and van den Ende, B. (1975) Productivity of peach trees: factors affecting dry-weight distribu-
tion during tree growth. Annals of Botany 39, 423–432.
Chalmers, D.J., Canterford, R.L., Jerie, P.H., Jones, T.R. and Ugalde, T.D. (1975) Photosynthesis in relation to
growth and distribution of fruit in peach trees. Australian Journal of Plant Physiology 2, 635–645.
Chandler, W.H. (1942) Deciduous Orchards. Lea and Febiger, Philadelphia, Pennsylvania.
Charles-Edwards, D.A. (1982) Physiological Determinants of Crop Growth. Academic Press, Sydney, Australia.
Conners, C.H. (1919) Growth of fruits of peach. New Jersey Agricultural Experiment Station Annual Report
40, 82–88.
Corelli Grappadelli, L., Ravaglia, G. and Asirelli, A. (1996) Shoot type and light exposure influence carbon
partitioning in peach cv. Elegant Lady. Journal of Horticultural Science 71, 533–543.
Costa, G. and Vizzotto, G. (2000) Fruit thinning of peach trees. Plant Growth Regulation 31, 113–119.
Costes, E., Lanri, P.E., Guédon, Y. and de Reffye, P. (1993) Modelling growth of peach trees using the renewal
theory. Acta Horticulturae 349, 253–258.
Crews, C.E., Williams, S.L. and Vines, H.M. (1975) Characteristics of photosynthesis in peach leaves. Planta
126, 97–104.
Deguchi, M., Saeki, H., Ohkawa, W., Kanahama, K. and Kanayama, Y. (2002) Effects of low temperature on
sorbitol biosynthesis in peach leaves. Journal of the Japanese Society for Horticultural Science 71, 446–
448.
DeJong, T.M. (1982) Leaf nitrogen content and CO2 assimilation capacity in peach. Journal of the American
Society for Horticultural Science 107, 955–959.
DeJong, T.M. (1983) CO2 assimilation characteristics of five Prunus tree fruit species. Journal of the American
Society for Horticultural Science 108, 303–307.
DeJong, T.M. (1986a) Fruit effects on photosynthesis in Prunus persica. Physiologia Plantarum 66, 149–153.
DeJong, T.M. (1986b) Effects of reproductive and vegetative sink activity on leaf conductance and water
potential in Prunus persica L. Batsch. Scientia Horticulturae 29, 131–137.
DeJong, T.M. (1999) Developmental and environmental control of dry-matter partitioning in peach. Hort-
Science 34, 1037–1040.
DeJong, T.M. and Doyle, J.F. (1985) Seasonal relationships between leaf nitrogen content (photosynthetic
capacity) and leaf canopy light exposure in peach Prunus persica. Plant, Cell & Environment 8, 701–706.
DeJong, T.M. and Goudriaan, J. (1989) Modeling peach fruit growth and carbohydrate requirements: reevalu-
ation of the double-sigmoid growth pattern. Journal of the American Society for Horticultural Science
114, 800–804.
DeJong, T.M. and Grossman, Y.L. (1995) Quantifying sink and source limitations on dry matter partitioning to
fruit growth in peach trees. Physiologia Plantarum 95, 437–443.
DeJong, T.M. and Walton, E.F. (1989) Carbohydrate requirements of peach fruit growth and respiration. Tree
Physiology 5, 329–335.
DeJong, T.M., Doyle, J.F. and Day, K.R. (1987) Seasonal patterns of reproductive and vegetative sink activity
in early and late maturing peach (Prunus persica) cultivars. Physiologia Plantarum 71, 83–88.
DeJong, T.M., Day, K.R. and Johnson, R.S. (1989) Partitioning of leaf nitrogen with respect to within canopy light
exposure and nitrogen availability in peach (Prunus persica). Trees: Structure and Function 3, 89–95.
DeJong, T.M., Grossman, Y.L., Vosburg, S.F. and Pace, L.S. (1996) PEACH: a user friendly peach tree growth
and yield simulation model for research and education. Acta Horticulturae 416, 199–206.
Dorsey, M.J. and McMunn, R.L. (1928) The third report on the Illinois thinning investigations. Proceedings of
the American Society for Horticultural Science 25, 269–276.
Escobar-Gutiérrez, A.J. and Gaudillére, J.-P. (1994) Variability in sorbitol:sucrose ratios in mature leaves of
different peach cultivars. Journal of the American Society for Horticultural Science 119, 321–324.
Escobar-Gutiérrez, A.J. and Gaudillère, J.-P. (1997) Carbon partitioning in source leaves of peach, a sorbitol-
synthesizing species is modified by photosynthetic rate. Physiologia Plantarum 100, 353–360.
Escobar-Gutiérrez, A.J., Moing, A. and Gaudillère, J.P. (1998) Time course of carbohydrates concentration in
mature leaves of peach seedlings (Prunus persica L. Batsch). Acta Horticulturae 465, 337–344.
Evert, R.F. (1960) Phloem structure in Pyrus communis L. and its seasonal changes. University of California
Publications in Botany 32, 127–196.
 Carbon Assimilation, Partitioning and Budget Modelling 259

Flore, J.A. and Lakso, A.N. (1989) Environmental and physiological regulation of photosynthesis in fruit crops.
Horticultural Reviews 11, 111–157.
Gao, Z.F., Maurousset, L., Lemoine, R., Yoo, S.D., van Nocker, S. and Loescher, W. (2003) Cloning, expression,
and characterization of sorbitol transporters from developing sour cherry fruit and leaf sink tissues. Plant
Physiology 131, 1566–1575.
Gaudillère, J.P., Moing, A. and Carbonne, F. (1992) Vigour and non-structural carbohydrates in young prune
trees. Scientia Horticulturae 51, 197–211.
Genard, M. and Bruchou, C. (1993) A functional and exploratory approach to studying growth: the example
of the peach fruit. Journal of the American Society for Horticultural Science 118, 317–323.
Genard, M. and Huguet, J.G. (1996) Modeling the response of peach fruit growth to water stress. Tree Physiology
16, 407–415.
Genard, M. and Huguet, J.G. (1997) Modeling the effect of irrigation on peach fruit quality. Acta Horticulturae
449, 161–168.
Genard, M. and Souty, M. (1996) Modeling the peach sugar contents in relation to fruit growth. Journal of the
American Society for Horticultural Science 121, 1122–1131.
Genard, M., Lescourret, F., Ben Mimoun, M., Besset, M.J. and Bussi, C. (1998) A simulation model of growth
at the shoot-bearing fruit level. II. Test and effect of source and sink factors in the case of peach.
European Journal of Agronomy 9, 189–202.
Genard, M., Lescourret, F. and Ben Mimoun, M. (1999) Simulation of the effect of fruit thinning on peach quality.
Acta Horticulturae 499, 61–68.
Genard, M., Lescourret, F., Gomez, L. and Habib, R. (2003) Changes in fruit sugar concentrations in response
to assimilate supply, metabolism and dilution: a modeling approach applied to peach fruit. Tree Physiology
23, 373–385.
Gifford, R.M. and Evans, L.T. (1981) Photosynthesis, carbon partitioning and yield. Annual Review of Plant
Physiology 32, 485–509.
Girona, J., Mata, M., Goldhamer, D.A., Johnson, R.S. and DeJong, T.M. (1993) Patterns of soil and tree water
status and leaf functioning during regulated deficit irrigation scheduling in peach. Journal of the American
Society for Horticultural Science 118, 580–586.
Girona, J., Marsal, J., Mata, M., Arbones, A. and DeJong, T.M. (2004) A comparison of the combined effect of
water stress and crop load on fruit growth during different phenological stages in young peach trees.
Journal of Horticultural Science & Biotechnology 79, 308–315.
Giuliani, R. and Magnanini, E. (2002) A systematic approach to model radiation interception and gas
exchange in whole-tree peach canopies. Acta Horticulturae 584, 101–105.
Giuliani, R., Magnanini, E. and Corelli Grappadelli, L. (1998) Whole canopy gas exchanges and light inter-
ception of three peach training systems. Acta Horticulturae 465, 309–317.
Grant, C.R. and ap Rees, T. (1981) Sorbitol metabolism by apple seedlings. Phytochemistry 20, 1505–1511.
Greutert, H., Martinola, E. and Keller, F. (1998) Mannitol transport by vacuoles of storage parenchyma of
celery petioles operates by facilitated diffusion. Journal of Plant Physiology 153, 91–96.
Grossman, Y.L. (1993) The carbon economy of reproductive and vegetative growth of a woody perennial,
peach (Prunus persica (L.) Batsch): growth potentials, respiratory demand and a simulation model. PhD
dissertation, University of California, Davis, California.
Grossman, Y.L. and DeJong, T.M. (1994a) Carbohydrate requirements for dark respiration by peach vegetative
organs. Tree Physiology 14, 37–48.
Grossman, Y.L. and DeJong, T.M. (1994b) PEACH: a simulation model of reproductive and vegetative growth
in peach trees. Tree Physiology 14, 329–345.
Grossman, Y.L. and DeJong, T.M. (1995a) Maximum fruit growth potential and seasonal patterns of resource
dynamics during peach growth. Annals of Botany 75, 553–560.
Grossman, Y.L. and DeJong, T.M. (1995b) Maximum fruit growth potential following resource limitation dur-
ing peach growth. Annals of Botany 75, 561–567.
Grossman, Y.L. and DeJong, T.M. (1995c) Maximum vegetative growth potential and seasonal patterns of
resource dynamics during peach growth. Annals of Botany 76, 473–482.
Grossman, Y.L. and DeJong, T.M. (1998) Training and pruning system effects on vegetative growth potential,
light interception, and cropping efficiency in peach trees. Journal of the American Society for Horticul-
tural Science 123, 1058–1064.
Hansen, P. (1970) 14C studies on apple trees. VI. Influence of the fruit on the photosynthesis of the leaves, and
the relative photosynthetic yields of fruits and leaves. Physiologia Plantarum 23, 805–810.
Harper, J.L. (1980) Plant demography and ecological theory. Oikos 35, 244–253.
260 T.M. DeJong and A. Moing

Harris, R.W. (1983) Arboriculture. Prentice-Hall, Englewood Cliffs, New Jersey.

Hirai, M. (1981) Purification and characteristics of sorbitol-6-phosphate dehydrogenase from loquat leaves.
Plant Physiology 67, 221–224.
Ho, L.C. (1988) Metabolism and compartmentation of imported sugars in sink organs in relation to sink
strength. Annual Review of Plant Physiology 39, 355–378.
Ho, L.C. (1992) Fruit growth and sink strength. In: Marshall, C. and Grace, J. (eds) Fruit and Seed Production.
Aspects of Development, Environmental Physiology and Ecology. Cambridge University Press, Cambridge,
UK, pp. 101–124.
Hubbard, N.L., Pharr, D.M. and Huber, S.C. (1991) Sucrose phosphate synthase and other sucrose metaboliz-
ing enzymes in fruits of various species. Physiologia Plantarum 82, 191–196.
Johnson, R.S. and Handley, D.F. (1989) Thinning response of early, mid-, and late-season peaches. Journal of
the American Society for Horticultural Science 11, 852–855.
Jordan, M.O. and Habib, R. (1996) Mobilizable carbon reserves in young peach trees as evidenced by trunk
girdling experiments. Journal of Experimental Botany 47, 79–87.
Kanayama, Y., Mori, H., Imaseki, H. and Yamaki, S. (1992) Nucleotide sequence of a cDNA encoding NADP-
sorbitol-6-phosphate dehydrogenase from apple. Plant Physiology 100, 1607–1608.
Kappel, F. and Flore, J.A. (1983) Effect of shade on photosynthesis, specific leaf weight, leaf chlorophyll content,
and morphology of young peach trees. Journal of the American Society for Horticultural Science 108,
541–544.
Kappes, E.M. (1985) Carbohydrate production, balance and translocation in leaves, shoots and fruits of ‘Mont-
morency’ sour cherry. PhD dissertation, Michigan State University, East Lansing, Michigan.
Kondo, A. and Hiramatsu, T. (1999) Analysis of peach tree damage caused by peach silver mite, Aculus fock-
eui (Nalepa et Trouessart) (Acari: Eriophyidae). Japanese Journal of Applied Entomology and Zoology 43,
189–193.
Kubota, N., Kohno, A. and Shimamura, K. (1990) Translocation and distribution of 13C-photosynthates in
‘Sanyo Suimitsu’ peach trees as affected by different rootstocks. Journal of the Japanese Society for
Horticultural Science 59, 319–324.
Lacointe, A. (2000) Carbon allocation among tree organs: a review of basic processes and representation in
functional–structural models. Annals of Forest Science 57, 521–534.
Lemoine, R. (2000) Sucrose transporters in plants: update on function and structure. Biochimica et Biophysica
Acta – Biomembranes 1465, 246–262.
LeRoux, X., Walcroft, A.S., Daudet, F.A., Sinoquet, H., Chaves, M.M., Rodrignes, A. and Osorio, L. (2001)
Photosynthetic light acclimation in peach leaves: importance of changes in mass:area ratio, nitrogen
concentration, and leaf nitrogen partitioning. Tree Physiology 21, 377–386.
Lescourret, F., Ben Mimoun, M. and Genard, M. (1998) A simulation model of growth at the shoot-bearing
fruit level. I. Description and parameterization for peach. European Journal of Agronomy 9, 173–188.
Lo Bianco, R., Rieger, M. and Sung, S.-J.S. (1999a) Carbohydrate metabolism of vegetative and reproductive
sinks in the late-maturing peach cultivar ‘Encore’. Tree Physiology 19, 103–109.
Lo Bianco, R., Rieger, M. and Sung, S-.J.S. (1999b) Activities of sucrose and sorbitol metabolizing enzymes in
vegetative sinks of peach and correlation with sink growth rate. Journal of the American Society for
Horticultural Science 124, 381–388.
Loescher, W.H., Marlow, G.C. and Kennedy, R.A. (1982) Sorbitol metabolism and sink–source interconver-
sions in developing apple leaves. Plant Physiology 20, 335–339.
Loescher, W.H., McCamant, T. and Keller, J.D. (1990) Carbohydrate reserves, translocation and storage in
woody plant roots. HortScience 25, 274–281.
Mandre, O., Rieger, M., Myers, S.C., Seversen, R. and Regnard, J.-L. (1995) Interaction of root confinement
and fruiting in peach. Journal of the American Society for Horticultural Science 120, 228–234.
Marcelis, L.F.M. (1994) A simulation model for dry matter partitioning in cucumber. Annals of Botany 74,
43–52.
Marini, R.P. and Marini, M.C. (1983) Seasonal changes in specific leaf weight, net photosynthesis, and chlo-
rophyll content of peach leaves as affected by light penetration and canopy position. Journal of the
American Society for Horticultural Science 108, 600–605.
Marini, R.P. and Sowers, D. (1994) Peach fruit weight is influenced by crop density and fruiting shoot length
but not position on the shoot. Journal of the American Society for Horticultural Science 119, 180–184.
Marquat, C., Vandamme, M., Gendraud, M. and Petel, G. (1999) Dormancy in vegetative buds of peach: rela-
tion between carbohydrate absorption potentials and carbohydrate concentration in the bud during
dormancy and its release. Scientia Horticulturae 79, 151–162.
 Carbon Assimilation, Partitioning and Budget Modelling 261

Marsal, J., Basile, B., Solari, L. and DeJong, T.M. (2003) Influence of branch autonomy on fruit, scaffold, trunk
and root growth during Stage III of peach fruit development. Tree Physiology 23, 313–323.
Maurel, K., Leite, G.B., Bonhomme, M., Guillot, A., Rageau, R., Pétel, G. and Sakr, S. (2004) Trophic control of
bud break in the peach tree (Prunus persica): a possible role of hexoses. Tree Physiology 24, 579–588.
Miller, A.N. and Walsh, C.S. (1988) Growth and seasonal partitioning of dry matter in eight-year-old ‘Loring’
peach trees. Journal of the American Society for Horticultural Science 113, 309–314.
Moing, A., Carbonne, F., Rashad, M.H. and Gaudillere, J.P. (1992) Carbon fluxes in mature peach leaves. Plant
Physiology 100, 1878–1884.
Moing, A., Escobar-Gutiérrez, A. and Gaudillère, J.P. (1994) Modeling carbon export out of mature peach
leaves. Plant Physiology 106, 591–600.
Moing, A., Carbonne, F., Rashad, M.H. and Gaudillère, J.P. (1997) Phloem loading in peach: symplastic or
apoplastic? Physiologia Plantarum 101, 489–496.
Moing, A., Poëssel, J.L., Svanella-Dumas, L., Loonis, M. and Kervella, J. (2003) Biochemical basis of low fruit
quality of Prunus davidiana, a pest and disease resistance donor for peach breeding. Journal of the
American Society for Horticultural Science 128, 55–62.
Moriguchi, T. and Yamaki, S. (1988) Purification and characterization of sucrose synthase from peach (Prunus
persica) fruit. Plant & Cell Physiology 29, 1361–1366.
Moriguchi, T., Sanada, T. and Yamaki, S. (1990) Seasonal fluctuations in some enzymes relating to sucrose and
sorbitol metabolism in peach fruit. Journal of the American Society for Horticultural Science 115, 278–281.
Moriguchi, T., Abe, K., Sanada, T. and Yamaki, S. (1992) Levels and role of sucrose synthase, sucrose-phosphate
synthase, and acid invertase in sucrose accumulation in fruit of Asian pear. Journal of the American
Society for Horticultural Science 117, 274–278.
Ng, E. and Loomis, R.S. (1984) Simulation of Growth and Yield of the Potato Crop. Pudoc, Wageningen, The
Netherlands.
Nii, N., Yamaguchi, K. and Nishimura, M. (1997) Changes in carbohydrate and ribulose bisphosphate carboxylase-
oxygenase contents in peach leaves after applications of different amounts of nitrogen fertilizer. Journal
of the Japanese Society for Horticultural Science 66, 505–511.
Noiraud, N., Maurousset, L. and Lemoine, R. (2001) Transport of polyols in higher plants. Plant Physiology
and Biochemistry 39, 717–728.
Olien, W.C., Graham, C.J., Hardin, M.E. and Bridges, W.C. (1995) Peach rootstock differences in ring nema-
tode tolerance related to effects on tree dry weight, carbohydrate and prunasin contents. Physiologia
Plantarum 94, 117–123.
Oliveira, C.M. and Priestley, C.A. (1988) Carbohydrate reserves in deciduous fruit trees. Horticultural Review
10, 403–430.
Park, S.W., Song, K.J., Kim, M.Y., Hwang, J.H., Shin, Y.U., Kim, W.C. and Chung, W.I. (2002) Molecular cloning
and characterization of four cDNAs encoding the isoforms of NAD-dependent sorbitol dehydrogenase
from Fuji apple. Plant Science 162, 513–519.
Pavel, E.W. and DeJong, T.M. (1993a) Relative growth rate and its relationship to compositional changes of
non structural carbohydrates in the mesocarp of developing peach fruits. Journal of the American Society
for Horticultural Science 118, 503–508.
Pavel, E.W. and DeJong, T.M. (1993b) Seasonal CO2 exchange patterns of developing peach (Prunus persica)
fruits in response to temperature, light and CO2 concentration. Physiologia Plantarum 88, 322–330.
Pavel, E.W. and DeJong, T.M. (1993c) Estimating the photosynthetic contribution of developing peach (Prunus persica)
fruits to their growth and maintenance carbohydrate requirements. Physiologia Plantarum 88, 331–338.
Petrov, A.A. and Manolov, P. (1973) Autumn accumulation of reserve 14C-labelled assimilates and their spring
mobilization in young peach trees. Comptes Rendus de l’Académie Agricole Georgi Dimitrov 6, 91–102.
Priestley, C.A. (1970) Carbohydrate storage and utilization. In: Luckwill, L.C. and Cutting, C.V. (eds) Physiol-
ogy of Tree Crops. Academic Press, London, pp. 113–126.
Proebsting, E.L. (1958) A quantitative evaluation of the effect of fruiting on growth of Elberta peach trees.
Proceedings of the American Society for Horticultural Science 71, 103–109.
Prusinkiewicz, P. (1998) Modeling of spatial structure and development of plants: a review. Scientia Horticul-
turae 74, 113–149.
Quilot, B., Genard, M., Kervella, J. and Lescourret, F. (2002) Ecophysiological analysis of genotypic variation
in peach fruit growth. Journal of Experimental Botany 53, 1613–1625.
Quilot, B., Génard, M. and Kervella, J. (2004a) Leaf light saturated photosynthesis for wild and cultivated
peach genotypes and their hybrids: a simple mathematical modeling analysis. Journal of Horticultural
Science & Biotechnology 79, 546–553.
262 T.M. DeJong and A. Moing

Quilot, B., Genard, M., Kervella, J. and Lescourret, F. (2004b) Analysis of genotypic variation in fruit flesh
total sugar content via an ecophysiological model applied to peach. Theoretical and Applied Genetics
109, 440–449.
Retzlaff, W.A., Williams, L.E. and DeJong, T.M. (1991) The effect of different atmospheric ozone partial pres-
sures on photosynthesis and growth of nine fruit and nut tree species. Tree Physiology 8, 93–105.
Reyes-Lopez, A. (1984) Effect of water stress on photosynthesis and leaf conductance in pistachio (Pistacia
vera L.) and peach (Prunus persica Batch.). PhD dissertation, University of California, Davis, California
Rieger, M. and Duemmel, M.J. (1992) Comparison of drought resistance among Prunus species from divergent
habitats. Tree Physiology 11, 369–380.
Rieger, M., Lo Bianco, R. and Okie, W.R. (2003) Response of Prunus ferganensis, Prunus persica and two in-
terspecific hybrids to moderate drought stress. Tree Physiology 23, 51–58.
Robertson, J.A., Meredith, F.I. and Scorza, R. (1988) Characteristics of fruit from high- and low-quality peach
cultivars. HortScience 23, 1032–1034.
Rosati, A., Esparza, G., DeJong, T.M. and Pearcy, R.W. (1999) Influence of canopy light environment and
nitrogen availability on leaf photosynthetic characteristics and photosynthetic nitrogen-use efficiency of
field-grown nectarine trees. Tree Physiology 19, 173–180.
Rosati, A., Day, K.R. and DeJong, T.M (2000) Distribution of leaf mass per unit area and leaf nitrogen concen-
tration determine partitioning of leaf nitrogen within tree canopies. Tree Physiology 20, 271–276.
Rosati, A., DeJong, T.M. and Esparza, G. (2002) Physiological basis for light use efficiency models. Acta
Horticulturae 584, 89–94.
Rufat, J. and DeJong, T.M. (2001) Estimating seasonal nitrogen dynamics in peach trees in response to nitrogen
availability. Tree Physiology 21, 1133–1140.
Saenz, J.L., DeJong, T.M. and Weinbaum, S.A. (1997) Nitrogen stimulated increases in peach yields are
associated with extended fruit development period and increased fruit sink capacity. Journal of the
American Society for Horticultural Science 122, 772–777.
Sakanishi, K., Kanayama,Y., Mori, H., Yamada, K. and Yamaki, S. (1998) Expression of the gene for NADP-
dependent sorbitol-6-phosphate dehydrogenase in peach leaves of various developmental stages. Plant
& Cell Physiology 39, 1372–1374.
Salvatierra, M.A., Gemma, H. and Iwahori, S. (1998) Partitioning of carbohydrates and development of tissues
in the graft union of peaches grafted on Prunus tomentosa Thunb. rootstock. Journal of the Japanese
Society for Horticultural Science 67, 475–482.
Sorce, C., Massai, R., Picciarelli, P. and Lorenzi, R. (2002) Hormonal relationships in xylem sap of grafted and
ungrafted Prunus rootstocks. Scientia Horticulturae 93, 333–342.
Spencer, S. and Couvillon, G.A. (1975) The relationship of node position to bloom date, fruit size and
endosperm development of the peach, Prunus persica (L.) Batsch cv. Sullivan’s Elberta. Journal of the
American Society for Horticultural Science 100, 242–244.
Sprugel, D.G., Hinkley, T.M. and Schaap, W. (1991) The theory and practice of branch autonomy. Annual
Review of the Ecology System 22, 309–334.
Stassen P.J.C., Strydom, D.K. and Stindt, H.W. (1981) Seasonal changes in carbohydrate fractions of young
Kakamas peach trees. Agroplantae 13, 47–53.
Stassen, P.J.C., Bergh, O., Bester, C.W.L. and DuPreez, M.M. (1982) Reserves in full-bearing peach trees –
carbohydrate reserves and their implications to orchard practices. The Deciduous Fruit Grower 32,
424–430.
Tanase, K., Shiratake, K., Mori, H. and Yamaki, S. (2002) Changes in the phosphorylation state of sucrose
synthase during development of Japanese pear fruit. Physiologia Plantarum 114, 21–26.
Toma, S., Ivascu, A., Balan, V., Delian, E. and Oprea, M. (2003) Evaluation of genetic resources of peach and
nectarine for powdery mildew resistance by physiological parameters. Acta Horticulturae 623, 291–
298.
Tromp, J. (1983) Nutrient reserves in roots of fruit trees, in particular carbohydrates and nitrogen. Plant and
Soil 71, 401–413.
Vemmos, S.N. and Goldwin, G.K. (1994) The photosynthetic activity of Cox’s Orange Pippin apple flowers in
relation to fruit setting. Annals of Botany 73, 385–391.
Vizzotto, G., Pinton, R., Varanini, Z. and Costa, G. (1996) Sucrose accumulation in developing peach fruit.
Physiologia Plantarum 96, 225–230.
Walcroft, A., Le Roux, X., Diaz-Espejo, A., Dones, N. and Sinoquet, H. (2002) Effects of crown development
on leaf irradiance, leaf morphology and photosynthetic capacity in a peach tree. Tree Physiology 13,
929–938.
 Carbon Assimilation, Partitioning and Budget Modelling 263

Wang, Z., Quebedeaux, B. and Stutte, G.W. (1996) Partitioning of [14C]glucose into sorbitol and other carbo-
hydrates in apple under stress. Australian Journal of Plant Physiology 23, 245–251.
Wang, Z., Yuan, Z. and Quebedeaux, B. (1998) Photoperiod alters partitioning of newly-fixed 14C and reserve
carbon into sorbitiol, sucrose and starch in apple leaves, stem and roots. Australian Journal of Plant
Physiology 25, 503–506.
Wang, Z., Pan, Q. and Quebedeaux, B. (1999) Carbon partitioning into sorbitol, sucrose and starch in source
and sink apple leaves as affected by elevated CO2. Environmental and Experimental Botany 41, 39–46.
Watson, M.A. and Casper, B.B. (1984) Morphogenetic constraints on patterns of carbon distribution in plants.
Annual Review of the Ecology System 15, 233–250.
Weinstein, D.A. and Yanai, R.D. (1994) Integrating the effects of simultaneous multiple stresses on plants using
the simulation model TREGRO. Journal of Environmental Quality 23, 418–428.
Westwood, M.N. and Gerber, R.K. (1958) Seasonal light intensity and fruit quality factors as related to
the method of pruning peach trees. Proceedings of the American Society for Horticultural Science 72,
85–91.
White, J. (1979) The plant as a metapopulation. Annual Review of the Ecology System 10, 109–145.
Yamada, K., Niwa, N., Shiratake, K. and Yamaki, S. (2001) cDNA cloning of NAD-dependent sorbitol dehy-
drogenase from peach fruit and its expression during fruit development. Journal of Horticultural Science
& Biotechnology 76, 581–587.
Yamaki, S. and Ishikawa, K. (1986) Roles of four sorbitol related enzymes and invertase in the seasonal
alteration of sugar metabolism in apple tissue. Journal of the American Society for Horticultural Science
111, 134–137.
Yano, T., Shinkai, S., Inoue, H. and Moriguchi, K. (2000) Seasonal changes in starch and soluble sugar con-
tents of two peach cultivars on Prunus tomentosa rootstock showing different degree of tree decline.
Journal of the Japanese Society for Horticultural Science 69, 711–717.
Zhou, R., Sicher, R.C. and Quebedeaux, B. (2002) Apple leaf sucrose-phosphate synthase is inhibited by
sorbitol-6-phosphate. Functional Plant Biology 29, 569–574.
Zhou, L.L., Chen, C.C., Ming, R., Christopher, D.A. and Paull, R.E. (2003a) Apoplastic invertase and its
enhanced expression and post-translation control during fruit maturation and ripening. Journal of the
American Society for Horticultural Science 128, 628–635.
Zhou, R., Cheng, L.L. and Wayne, R. (2003b) Purification and characterization of sorbitol-6-phosphate phos-
phatase from apple leaves. Plant Science 165, 227–232.
Zhou, R., Sicher, R.C., Cheng, L.L. and Quebedeaux, B. (2003c) Regulation of apple leaf aldose-6-phosphate
reductase activity by inorganic phosphate and divalent cations. Functional Plant Biology 30, 1037–
1043.
 11 Orchard Planting Systems

L. Corelli-Grappadelli1 and R.P. Marini2

1Dipartimento di Colture Arboree, University of Bologna, Bologna, Italy
2Department of Horticulture, The Pennsylvania State University, University Park,
Pennsylvania, USA

11.1 Introduction 264

11.2 Evolution of Training and Pruning Principles 265
Minimal pruning 265
Summer pruning 266
Cropping as a tool in training 266
Intensification of peach orchard systems 266
11.3 Light and Tree Physiology 267
Photosynthetic characteristics 267
Light interception and distribution 267
Fruit quality 269
Flower bud differentiation 270
11.4 Modern Orchard Systems 270
Open vase 271
Palmette 272
Central leader 274
Y-shaped tree 276
Meadow orchards 278
Protected cultivation 279
11.5 Effects of Training System and Planting Density on Yield 280
Tree form effect on yield 280
Effect of planting density on yield 281
Effect of tree density plus tree form on yield 282
11.6 Future Trends 283

11.1 Introduction since 1974, with China accounting for most of

Peach is one of the most important temperate
fruit tree crops, with a world production in 2003
of about 14.8 million t. Total peach plantings
and output worldwide have nearly doubled
this change (FAOSTAT, 2004). Over the same
period of time, output per hectare has increased
from 9.7 to 10.4 t; in the most advanced coun-
tries, output generally exceeds 15 t/ha. This
difference certainly reflects vast differences in

264 © CAB International 2008. The Peach: Botany, Production and Uses
(eds D.R. Layne and D. Bassi)
 Orchard Planting Systems
production techniques, including the choice
and management of the orchard design (the
combination of cultivar, rootstock, training
system, spacing, tree size). The orchard designs
adopted worldwide vary widely; it is known
that different training systems can have quite
different productive potentials. A large num-
ber of studies have been undertaken globally
since the mid-1950s to elucidate several aspects
of tree performance in peach, and these have
been reviewed recently (Marini and Corelli
Grappadelli, 2006). This past half century has
seen a broad change in the forms peach trees
are trained to and in the way a given form is
achieved, and all along constant increases in
productivity and efficiency have accompa-
nied these changes. The thrust behind this
evolution has mostly been the economics of
the operation. For peach growing to remain
economically viable in the more advanced
countries, particularly those in which agricul-
tural land is at a premium, it is necessary to
increase orchard efficiency and productivity.
In this chapter the evolution of the training
and pruning techniques that have accompa-
nied and fostered the evolution of orchard
training systems, as well as the scientific bases
underpinning these technical achievements,
are reviewed. Then follow a detailed discus-
sion of the most widely adopted training sys-
tems and a forecast of future trends in this
important part of peach cultivation.
11.2 Evolution of Training and
Pruning Principles
In modern orchards, shortening the initial
unproductive period is emphasized: the sooner
trees come into bearing, the sooner the eco-
nomic burden on the orchard begins to be
alleviated. Instrumental to this are high plant-
ing densities: even if each individual tree does
not bear a large amount of fruit, the higher
number of trees per hectare allows sizeable
initial yields. Later on in the life of the orchard,
the differences in yield between high- and low-
density orchard designs tend to disappear
because at this stage most orchards will inter-
cept similar amounts of light, which translates
into similar yields. The pruning techniques
265
adopted in the training stage of the tree are
the key to hastening bearing. Today, there is
a consensus that the most important aspects
include reducing the number of pruning cuts
in the early years, avoiding heading cuts
wherever possible, using summer rather than
winter pruning to train the trees, and allow-
ing cropping in the early years as a tool to
reduce vegetative growth since this species
lacks vigour-controlling rootstocks.
Minimal pruning
In general, the fewer pruning cuts made to
the tree in the first 2 to 3 years from planting,
the quicker the tree will come into bearing.
The minimal amount of pruning favours the
onset of the reproductive phase of the tree.
This concept was first promoted by extension
agents and researchers in Italy in the early
1950s (Baldassari, 1950), who demonstrated
the advantages of the so-called ‘potatura a
tutta cima’ (no-heading-cuts pruning). Bal-
dassari argued against excessive cuts in the
early years because this would alter the equi-
librium of the tree towards a vegetative
response, but he also showed the advantage
of leaving the leader of the tree (and of the
branches) intact, in order to hasten the repro-
ductive stage. Therefore, only thinning cuts
removing entire shoots or branches should be
made, whereas all heading cuts on 1-year-old
wood (widely used to obtain regular branch
formation in the open vase or regular pal-
mette) were abandoned. This type of pruning
approach quite effectively shortened the time
needed for the transition of the tree from veg-
etative to reproductive growth.
Following this strategy is best accom-
plished by planting well-feathered maiden
trees: the best feathers can be selected to form
the permanent tree structure. However, pro-
duction of good maiden trees can be difficult,
as peach trees require high light to develop
laterals and in the nursery peaches are often
planted too closely. As a result, peach trees
tend to grow quite tall (particularly on vigor-
ous rootstocks) and may have only a few
short laterals, often with narrow crotch angles
or too high on the tree to start the first tier of
266 L. Corelli-Grappadelli and R.P. Marini
branches. Thinning out excessive feathers
and cutting them back to the first or second
node may induce regrowth of laterals that can
be used to start the permanent structure of
the tree without resorting to heading back the
leader. If there are no good laterals the leader
could still be maintained, but summer prun-
ing is required to check the growth of the
higher current-season laterals by pinching.
This needs to be repeated in the summer, so
that good laterals are formed by lower buds
during the first year. This is not optimal
because the first season in the field is spent
trying to overcome the problem, rather than
training the tree, but it can be preferable to
cutting back the leader. Heading of the tree in
the first year should be performed only in the
case of trees devoid of or with only poor later-
als, or when the training system requires it
(such as in the Y-trellis). When heading is nec-
essary it should be done at planting, so that
the tree may develop new laterals and have
the entire season to regrow. When the newly
formed laterals are long enough (around
20–30 cm) shoot selection is necessary to
choose the shoot that will reform the tree.
Summer pruning
Another important tool for shortening the
unproductive period is summer pruning.
This is mainly because summer pruning facil-
itates controlling the development of the tree
in a ‘softer’ way: very often a cut can be avoided
if the shoot can instead be bent, trained,
twisted, pinched, etc. All these operations are
far less ‘traumatic’ for the tree and cause less
reaction than dormant pruning. As a result,
the tree is less prone to vegetative growth
reactions which can be the result of heavy
cuts made during the winter. Another advan-
tage is that the tree is not investing too many
resources in structures that are not permanent.
Summer pruning is a practice most important
in the training phase of the orchard, when the
trees are young and must reach their adult
stage as quickly as possible. However, it is
also important for some high-planting den-
sity systems where heavy shading can cause a
loss of bearing wood in the lower parts of the
canopy. Because summer pruning removes
foliage, this impacts not only current-season
fruit growth but also the reserves for early
growth in the next season. In general fruit
colour formation can be improved by summer
pruning, but fruit size and internal quality
may suffer from the reduction in available
assimilates. It is difficult to devise a success-
ful summer pruning strategy because of the
many processes occurring simultaneously
which depend on light, i.e. fruit growth, shoot
growth, flower bud differentiation and reserve
accumulation.
Cropping as a tool in training
Allowing the presence of fruit on the tree is
increasingly used to train trees in a ‘soft’ man-
ner. Traditionally, all fruit would be removed
from young trees to improve the growth and
establishment of the tree structure. This was
done because the goal was to establish a
durable, robust tree, which would last many
years. As the goal has shifted towards less
durable trees, cropping has been regarded as
a natural means of containing tree vigour
with minimal pruning. Peach trees can feather
quite normally in the nursery, provided suf-
ficient light is available during the growing
season (which may not always be the case, as
indicated above). As a consequence, the
potential exists for a certain number of fruit to
be borne on the tree in the year of planting.
The number of fruits to be left, however, must
be small: the overriding goal is to control tree
growth, not to produce high early yields. Up
to five or six fruits per tree may be left in the
year of planting. Cropping acts by altering
partitioning of resources towards the grow-
ing fruit and away from vegetative growth.
Intensification of peach orchard systems
Work with apple demonstrated the feasibility
of increasing early yields by increasing the
number of trees per hectare. The high-density
orchards, based on smaller trees planted closer
together, reach their full production potential
more quickly than ordinary orchards; their
 Orchard Planting Systems
early yields, however, are also substantial
because of the high number of trees, which
acts as a powerful multiplier of even small
yields per tree. The increasing costs facing
growers in some peach-producing countries
have been a strong incentive for the change
towards more intensive orchard systems
(Sansavini, 1974; Bandoli, 1980). Since dwarfing
rootstocks for peach do not currently exist, the
trend towards high tree densities has never
reached the levels that are commonly observed
for apple, which is very frequently planted at
above 2000–3000 and up to 7000–8000 trees/ha.
However, the benefits of high-density plant-
ings can be reaped with peach as well. Well-
feathered trees from the nursery which are
minimally pruned, for example, can produce a
profitable crop in the year of planting. In
addition, as indicated above, growers can use
cropping as a powerful tool to control tree
vigour. It is interesting that the most commonly
used rootstock in Italy for almost two decades
has been the vigorous ‘GF 677’ peach × almond
hybrid. The reason for this apparent contradic-
tion is that, as peach wood dies quickly under
conditions of low light, the vigorous rootstock
provides the potential for regrowth from shaded
inner sections of the canopy when light condi-
tions are improved by pruning.
11.3 Light and Tree Physiology
Light is the driving factor for all plants on the
planet. Dry mass production of a plant depends
on the amount of light intercepted (Monteith,
1977), but in fruit trees the relationships between
light intercepted and yield are fairly complex
because they depend on the photosynthetic
characteristics of a species and on the amount
of light intercepted and distributed within a
canopy. It is therefore necessary to review
these aspects briefly before discussing peach
orchard systems.
Photosynthetic characteristics
The peach exhibits a C3 photosynthetic car-
bon fixation pathway; the carbon exchange
rate (CER) of peach leaves saturates at about
800 µmol/m2 per second of photosyntheti-
267
cally active radiation (PAR; 400 to 700 nm)
(Marini and Marini, 1983; DeJong and Doyle,
1985). The only report of whole-tree CER
shows saturation at about 1600 µmol/m2 per
second of PAR (Giuliani et al., 1998). This may
be due to the fact that whole-canopy CER
takes into account respiring organs such as
wood and fruits in addition to leaves, and
also reflects the various net photosynthesis
rates attained by leaves that are at different
light levels. Leaf mass per area (LMA; the
ratio of leaf dry weight to leaf area) is lower
for shaded than for non-shaded leaves and
was found to be a biological integrator of
cumulative light exposure for a leaf until mid-
season (Marini and Barden, 1981). LMA of
leaves exposed to a range of light levels for 18
days was linearly related to light level, and
CER measured under saturating light was
related to LMA. The CER and LMA of leaves
shaded for 18 days and then exposed to full
sun for 26 and 4 days, respectively, were simi-
lar to those of non-shaded leaves (Marini and
Sowers, 1990). Leaf nitrogen content was related
to the light environment of a leaf, and leaf
nitrogen can be reallocated from shaded leaves
to more exposed leaves of the tree canopy
(DeJong and Doyle, 1985). These data indicate
that the redistribution of leaf nitrogen is a means
for maximizing whole-tree carbon gain.
Light interception and distribution
Mathematical models indicate that the amount
of light intercepted by a tree sets its maximum
potential for yield, as the latter cannot exceed
the former; however, the actual yield pro-
duced by a tree will also reflect the influence
of temperature, the tree’s vegetative/repro-
ductive equilibrium, its hormonal balance,
and its nutritional and water status. From the
standpoint of light alone, orchard productiv-
ity is influenced by the relationships between
orchard design (i.e. tree form, tree spacing
and row orientation) and available light, which
depends on average sun position. The goal is
to ensure concurrent high light interception
and good distribution throughout the canopy.
Modelling work in apple (Cain, 1972; Jackson
and Palmer, 1972; Jackson and Middleton,
1988) has indicated that two ways exist to
268 L. Corelli-Grappadelli and R.P. Marini
increase the amount of light intercepted by a
tree: either by increasing the density of the
canopy (which causes poor distribution within
the canopy) or by increasing the orchard’s
leaf area index (LAI; ratio of leaf area to unit
of land area) by planting a greater amount of
trees of smaller size (which allows for good
transmission of the light to all the parts of a
tree). This work has laid the theoretical foun-
dation for high-density plantings of apple
and other crops worldwide, including peach.
Lakso (1994), drawing upon results from
many studies in different parts of the world,
showed apple yields to be linearly related to
the percentage available light intercepted by
the tree. Interestingly, above 50% light inter-
ception the relationship appears less tight.
One of the reasons this could be attributed to
illustrates the effect of vegetative growth ver-
sus planting density. Low-cropping trees often
partition more resources to vegetative growth,
which increases their light interception with-
out a concurrent increase in yield. On the other
hand, orchard designs based on many small
trees, capable of high light interception and
distribution within their smaller canopies, may
retain greater crop loads (Robinson, 1997).
The amount of light intercepted and its
distribution depend not only on the amount
available at the particular latitude, but also on
the type of weather prevailing during the sum-
mer. In the north-eastern USA, for example, or
in northern Europe, conditions prevailing in
the summer include many hours of haze-
cloudiness which cause a shift in the propor-
tion of diffuse/direct light available. This in
turn has an effect on the amount of light that
can reach the internal positions of the canopy.
Lakso and Musselmann (1976) reported for
upstate New York that maximum light pene-
tration to the interior canopy occurs under a
mix of direct and diffuse radiation rather than
under maximum direct light conditions. On
the other hand, Mediterranean climates such
as Italy, California and Spain normally have
very high amounts of direct light and this may
largely alter these relationships. Fruit quality
parameters for apples developing in a cloud-
less, high direct radiation environment of
Washington State showed different relation-
ships with light levels than fruits from the mid-
Atlantic states (Campbell and Marini, 1992).
Reported light distribution patterns vary
between different fruit tree forms. In general,
the outer periphery of the canopy intercepts
and reflects a high proportion of the incoming
radiation, and this causes different light distri-
bution profiles for different training systems.
Light was distributed in a U-shaped pattern
along a horizontal cross-section of the canopy
for central leader trees (Porpiglia and Barden,
1980; Marini and Barden, 1982) and in a
W-shaped pattern in open vase type trees
(Marini and Marini, 1983). These studies
showed that in all systems light penetration
declines rapidly from the tree periphery
towards the tree centre. Most often the zone
which receives adequate light for high fruit
quality is within about 1.5 m from the outside
of the tree (Marini and Barden, 1982; Kappel
et al., 1983). Relatively high light levels were
measured in the centre of open vase trees
(Marini and Marini, 1983) and following prun-
ing or shearing of hedgerow systems in the
summer (Kappel et al., 1983; Marini, 1985).
The light levels within the canopy vary
also with height from the ground, as the upper
layers of leaves will project a shadow on those
below, and with time of the day. Génard and
Baret (1994) showed the variations in the
amounts of light transmitted to shoots in
open vase peach trees to be a function of posi-
tion in the canopy and of time of day. Well-
exposed shoots were mostly located at the top
of the tree and were relatively erect. Shoots
located in the outer parts of the canopy were
slightly but significantly more sunlit than oth-
ers. Some shoots were exposed to light almost
all day while other shoots were in sunlight
very little. About 30% of the shoots received
<30% of the incoming light. Light transmitted
to the shoots did not depend on shoot com-
pass direction. Drooping shoots were prefer-
entially shaded by horizontal and erect shoots
above and next to their position on the branch.
Virtually all orchard systems far from
intercept all of the total light available to them.
Among the reasons for this are the need to
allow room between rows for equipment
access and the need to set adequate spacing to
accommodate the height and girth of the
trees: if trees were very closely spaced together,
only the upper layers of their canopies would
receive any light. In apple, experimental
 Orchard Planting Systems 269

orchards have been reported to achieve
greater than 80% light interception (several
authors, cited in Robinson, 2003), but an aver-
age for the industry is certainly much less. In
many cases, well-managed orchards may
approach 50% interception of available light.
In peach, Grossman and DeJong (1998) mea-
sured the daily patterns of intercepted photo-
synthetic photon flux (PPF) in four peach
training systems in California, which varied
in planting density. The intercepted PPF was
relatively constant during mid-day hours
(from 09.00 to 15.00 h) for all training systems.
The high-density Kearney Agricultural Cen-
ter V (KAC-V; 1196 trees/ha) system and the
cordon system (919 trees/ha) both intercepted
more light than the low-density KAC-V (919
trees/ha). The open vase (299 trees/ha) sys-
tem intercepted the least light. The high-density
KAC-V and cordon systems intercepted nearly
twice as much light as the open vase system,
but no system intercepted much more than
50% of the available light. Robinson and Lakso
(1991) also found that Y-shaped apple trees
intercepted more light than conical-shaped
trees (70% versus 55% available light). This
training system is also capable of high light
interception in peach. Corelli Grappadelli et al.
(unpublished results) have measured light
interception of three training systems (pal-
mette, delayed vasette and Y-trellis) at stan-
dard commercial densities. The Y-trellis had
50% greater light interception from the second
season, and in some seasons has reached about
80% interception. The delayed vasette and the
palmette were always lower, in particular the
palmette, which was normally below 50%
(Fig. 11.1). More information is needed to
determine the relationship between light
interception and peach yield per hectare.
Only then can orchard systems be developed
that capture the optimum amount of light.
Fruit quality
Peach fruit quality depends on its position
within the canopy, because this influences the
amount of light intercepted by the fruit and
the leaves near the fruit. The sensitivity of the

100
Light interception (% available)

80

60

40

Delayed vasette
20 Palmette
Y-trellis

0
0 2 4 6 8 10
Year of planting

Fig. 11.1. Light interception pattern for a 9-year-old ‘Red Gold’ orchard, grafted on clonal seedlings,
trained in three different forms: delayed vasette (5.0 m × 3.7 m; 545 trees/ha), palmette (5.0 m × 2.8 m;
727 trees/ha) and Y-trellis (5.0 m × 1.6 m; 1274 trees/ha). The common spacing between rows in this
trial was dictated by a drain tile system which had spacing between tiles of 5 m. Light interception was
measured once a year, around summer solstice. The slight decreases for all systems after the seventh
year are due to reform pruning applied to keep the trees from expanding excessively. (From Corelli-
Grappadelli et al., unpublished results.)
270 L. Corelli-Grappadelli and R.P. Marini

peach to light varies during the season: shad-
ing portions of trees during the final 6 weeks
before harvest (Marini and Sowers, 1990;
Marini et al., 1991) and measuring light inter-
ception by fruit during the final swell (Lewal-
len and Marini, 2003) allowed the identification
of critical light levels for various aspects of
peach fruit quality. Flore and Kesner (1982)
also described critical light levels for various
aspects of peach tree growth and fruit quality.
These critical light levels are presented in
Table 11.1.
Flower bud differentiation
As discussed above, at high light interception
levels canopies may tend to become too dense.
If this is the case, the light-driven process of
flower bud differentiation may be hampered
with a consequent negative impact on fruit
quality, as well as yield. Levels of interception
greater than 70% of available light have been
reported for apple orchards, but at such levels
light distribution may be inadequate for con-
tinued high yields of quality fruit (Jackson,
1980). Spurs of pome fruits require at least
30% of light to remain fruitful (Jackson, 1967),
but leafy peach shoots must be exposed to
adequate light to remain alive. Heavily shaded
portions of peach canopies are devoid of fruit-
ing shoots because they die during the late
summer. Flower bud density was also posi-
tively related to light levels from about 50 to
100 days after bloom (Marini and Sowers,
1990). Therefore, to maintain high produc-
tion, light distribution in peach orchards is
more important than in apple orchards.
11.4 Modern Orchard Systems
In each fruit-growing area of the world, fairly
specific environmental, technical, organiza-
tional and economic constraints may limit the
choice of training system and orchard design
that growers can adopt. These constraints
affect the choice of cultivar, rootstock, tree
training system and tree spacing, which results
in what is normally defined as an orchard
system. The successful grower is one who can
adapt the various techniques for training and
managing the tree to their local conditions.
Because of this variability, while it can be said
that peach training systems are derived from
essentially four basic tree shapes, a great
many variations of each of these shapes exist,
which depend on the type of nursery tree
(maiden, dormant budded, trees grafted in
situ), the orchard site (particularly soil and

Table 11.1. A summary of critical light levels (percentage of full sun) for various aspects of peach leaf
function and fruit quality.

Light threshold (% of full sun)

Flore and Marini and Sowers (1990);

Parameter Response to light Kesner (1982) Marini et al. (1991)

Leaf size Negative 36 40

Leaf thickness Positive 36 40
Specific leaf weight Positive 36 40
Maximum CER Positive 36 35
Cold hardiness Positive 21 –
Fruit size Positive – 10
Fruit colour Positive 36 30
Soluble solids Positive – 50
Flesh firmness None – No response
Flower densitya Positive – 23

CER, carbon exchange rate.

aFlower bud formation was most affected by shade during the period 50 to 100 days after bloom.
 Orchard Planting Systems
temperature conditions), the skill of the
labour, etc. The shift towards higher densities
in peach has required adaptations and changes
in training systems, as each training system is
ideally suited to a specific density. Therefore,
if one scored training systems according to
density of planting for example, almost a con-
tinuum could be found in terms of trees per
hectare but with widely different training sys-
tems: from the very low-density open centre,
to smaller vases, to the palmette, to various
Y-trellis, V-form and slender spindle (fusetto)
systems, up to meadow orchards. As only the
most widely adopted training systems are
described in this chapter, the reader should
be aware that, for each of them, many vari-
ants can be found in the literature.
One common feature of many intensive
peach orchard systems is that they strive to
bring the orchard into production as early as
possible, for obvious economic reasons. If well-
feathered trees are planted, it is not uncom-
mon to pick peaches in the year of planting
(Sansavini and Neri, 2005). This is not only a
way to reduce the unproductive period at the
beginning of orchard life, but also serves the
purpose of controlling tree vigour, thus help-
ing in the training stage when peach trees tend
to grow excessively. On the other hand, early
bearing normally shortens the productive life
of an orchard. In the intensive peach industries
of southern European countries (Italy, Spain,
France) this is not considered a disadvantage
because it is desirable to renew orchards with
cultivars with improved fruit quality and that
expand the harvest season (Borras Escriba,
2001; Sansavini and Neri, 2005).
Open vase
Open vase or open-centre trees have been most
common in commercial orchards for more
than 150 years (Cole, 1849). Worldwide, this
system probably is the most adopted one
occurring in the largest acreage. Open vase
trees in California, USA usually have three
primary scaffold branches, each with two
secondary scaffolds; mature trees are fairly
upright and are about 4 to 5 m tall (Rizzi, 1975).
In eastern North America open vase and
271
modified leader trees have three to six primary
scaffold branches and bench cuts are used to
develop low (2.2 to 3.2 m tall) spreading trees
with tree densities of 220 to 550 trees/ha
(Fig. 11.2/Plate 70).
The open vase, with about 500 trees/ha,
is the most important training system in Spain
(Royo Díaz and Martínez Lopez, 1992), France
(Hugard, 1986; Hilaire, 2003) and Greece. In
Spain the variants of the open vase include
the ‘vaso Italiano’, ‘vaso de plataformas’ and
‘vaso Californiano’. The main difference is in
the angle of the branches and the hierarchy of
the secondary branches. Trees of the Italian vase
are spaced more widely because the branches
are trained to be more horizontal. The ‘vaso
de plataformas’ has more upright scaffold
branches (35 to 40° from the vertical) and each
carries three tiers of secondary branches; up
to four in the first tier and decreasing to two
in the top tier. The height of the tree is about
3 m. The ‘vaso Californiano’ features three
fairly upright scaffold branches (15 to 20° from
vertical), each with two secondary scaffolds
(Royo Díaz and Martínez Lopez, 1992).
In Italy, the open vase has long been
among the most popular training systems in
both the northern and southern growing dis-
tricts (Giovannini and Monastra, 1995;
Sansavini et al., 2000). The traditional open-
centre tree is no longer adopted as its density
is too low (400 trees/ha and less) and the
yields are consequently low, particularly in
the early years. In addition, that system
requires detailed pruning, which is too expen-
sive and delays bearing (Sansavini and Neri,
2005). A variant of the open centre which is
adopted in some parts of Italy is the delayed
vase. This tree is based on the natural basitony
of the young peach tree, and adopts all the
modern principles of pruning: minimal prun-
ing in the early years, no heading cuts, crop-
ping as a tool to control tree vigour.
Maintaining the tree leader for 3–4 years
helps open up the lower branches, which nat-
urally expand outwards, towards the light. At
the same time, by producing fruit, the leader
contributes to those important early yields.
When a well-feathered tree from the nursery
is used (feathers should not lack in the first
50–70 cm from the ground, where the first
whorl of branches is formed), this tree requires
272 L. Corelli-Grappadelli and R.P. Marini
Fig. 11.2. Traditional open vase tree form.
very little pruning in the first and second
seasons. Summer pruning is preferentially
adopted during training, and must be per-
formed throughout the life of the orchard to
maintain bearing wood inside the canopy
(Mascanzoni, 1998). This is important to
maintain the low bearing zone of the tree,
which is one of the main advantages of this
form. The leader is removed only after the
third, or sometimes the fourth, season, thus
the name ‘delayed’. The number of branches
retained may vary from four to six, arranged
in two whorls. Tree density is about 550 trees/
ha. In vigorous soils this vase can be difficult
to maintain low, within 2.2 to 2.5 m of the
ground, while it does very well with low-
vigour cultivar/rootstock combinations or in
low-vigour environments. A variant of the
vase that is sometimes used in Italy and Spain
is the so-called ‘forma libera’ or free shape
(Sansavini, 1980; Royo Díaz and Martínez
Lopez, 1992; Sansavini and Neri, 2005). The
tree is totally unpruned in the first 3 to 4 years,
which makes it very productive early on, but
in severe need of reform pruning after this
initial period. Because the tree develops a
bushy appearance, very often it is pruned
back to form a vase, which reduces yield in
the years immediately following this reform
pruning. This is the main reason for its lim-
ited adoption.
In general, the advantages of open vase
and open-centre trees are in the limited num-
ber of trees per hectare (which helps reducing
planting costs), the lack of a trellis system
and, at least for some of them, the low height,
which makes it possible to prune, thin and
pick from the ground without using ladders
(they do not lend to platforms at all). The dis-
advantages include low early yields (if the
trees are headed back and severely pruned
during training), late frosts since the short
trees are more prone to freezing damage, and
greater difficulty performing some cultural
practices, such as herbicide applications and
mowing, because of the spread of the tree
(Mascanzoni, 1998).
Palmette
Hedgerow training systems from which the
palmette was derived were in use in French
gardens in the 1700s (Sansavini and Neri,
 Orchard Planting Systems 273

2005). Hedgerows were introduced in com-
mercial orchards in Italy in the early 1900s,
but the modern palmette training system
was introduced by Baldassari (1950), an
innovative grower in Ferrara (northern Italy),
who switched to this training system from
the traditional vase- or pyramid-shaped fruit
trees to improve orchard efficiency and prof-
itability. This system proved to be more pro-
ductive and less expensive than the traditional
vase (Fideghelli, 1969) and ideally suited to
the use of platforms for pruning, thinning
and picking, which greatly improved labour
efficiency over the use of ladders needed for
the other systems. The palmette found its
greatest diffusion in Italian peach orchards
and to a lesser degree in other countries, espe-
cially Spain and France. In some important
peach districts of Italy it is still the leading
training system for peach (Sansavini et al.,
2000).
The palmette is normally formed from a
well-feathered tree, which is planted without
heading cuts and with some thinning cuts in
case there are too many laterals or they are
too crowded. The tree is allowed to grow
freely in the first year after planting; if it
grows fast and develops sufficiently, the first
tier of branches can be selected during the
first summer, or else the choice is postponed
to the second growing season. The following
tiers are chosen during the second/third
growing seasons, depending on the speed of
tree development. The branches are selected
according to height (no less than 80–100 cm
from the ground or the nearest branch) and
crotch angle, without complying with any
geometric scheme. Selecting the branches in
the summer has the advantage that they are
easier to work with (if they have not already
hardened) and will generally result in wider
crotch angles, which are desirable from a
mechanical as well as a growing (reduced
vigour) point of view. Branch selection and
positioning may require considerable time
(Corelli Grappadelli et al., 1986) and repre-
sents one of the major drawbacks of this
system compared with those based on a
free structure without trellising (e.g. fusetto,
delayed vasette, free shape). The branches are
tied to the trellis, which provides support for
the tree and a frame to form the structure
of the hedge. The number of branches is
reduced, sometimes to one tier, which is
grown out and up to occupy the space
between adjacent trees. Palmette trees are not
slower to come into bearing compared with
other systems (e.g. fusetto). If not headed in
the first year, canopy development will be the
same as for any other training system (Table
11.2). When the tree is established, pruning
the palmette is as rapid as other systems
(Sansavini et al., 1980). The narrow canopy
allows good light penetration (Corelli Grap-
padelli and Sansavini, 1989), the trellis helps
maintain the shape, and throughout the life of
the orchard the tree normally requires less
summer pruning to retain bearing wood in
the lower parts of the canopy. Typical densi-
ties for the palmette are between 600 and 900
trees/ha, with spacing varying between 4.5
m × 3.5 m and 4.0 m × 2.5 m, and a tree height
of 3.5–4.5 m, depending on the cultivar/
rootstock combination and soil fertility.

Table 11.2. Annual and cumulative yield per tree and per hectare of ‘Flavorcrest’ peach trees trained
as palmette, delayed vasette and free shape, planted at the density of 727 trees/ha, over the first four
growing seasons. (From Corelli Grappadelli et al., 1986.)

Yield per tree (kg) Yield per hectare (t)

Training system 2nd year 3rd year 4th year Sum of 2nd–4th years Sum of 2nd–4th years

Palmette 5.3a 7.3b 39.0a 51.6b 375a

Delayed vasette 6.3b 4.8a 35.4a 46.5a 338a
Free shape 6.8b 9.7c 50.6b 67.1c 486a
a,b,cMeans within columns with unlike superscript letters were significantly different by the least significant

difference test, a = 0.05.

274 L. Corelli-Grappadelli and R.P. Marini
The suitability of the palmette for use
with platforms helped it win over the taller
and wider trees in the orchard of the 1960s
and 1970s; but the capital investments required
for platforms and for trellising became its weak-
ness in the 1980s and 1990s, except for areas of
high-vigour conditions where it is impossible
to keep trees short and with the fruiting zone
near the ground (Fig. 11.3/Plate 71).
Central leader
While peach trees can be trained according to
this system, for a number of reasons it is not
well suited to peach: the trees become very
large, they lose bearing wood in the low and
inner canopy, their shape is not easy to main-
tain, and it can make the use of platforms dif-
ficult. Therefore, peach trees are rarely trained
to central leader (Fig. 11.4/Plate 72). Other
systems have been proposed, derived from
the central leader. The adaptation of the ‘axis
central’ to peach in France (Hugard, 1981;
Belluau and Lemaire, 1986) and the introduc-
tion of fusetto or free spindle in Italy (Bar-
gioni et al., 1983) were attempts to increase
peach tree densities to be similar, if not equal,
to those of apple (1000–2000 trees/ha). How-
Fig. 11.3. Palmette tree form.
ever, both systems tend to suffer from exces-
sive shading and loss of bearing wood in the
lower part of the canopy, particularly after
4–5 years from planting and at the higher
densities (Belluau and Lemaire, 1986; Loreti
et al., 1989). The trees have a single vertical
leader, are conical in shape and are about
3–3.5 m tall (Bargioni et al., 1983). The trees
create a hedgerow (Fig. 11.5/Plate 73), well
suited to picking platforms. Both systems fea-
ture a leader, with no permanent branches.
Fruiting wood is retained on short branches,
which are renewed every few years (Fig. 11.6/
Plate 74). Since central leader trees are closer
to the natural growth habit of peach than the
palmette, these training systems require less
pruning during training (if the tree is ade-
quately feathered) and their training is much
easier. As a result, early yields are normally
fairly high, which is their greatest advantage.
However, summer pruning is required
throughout the life of the orchard, as the
lower fruiting zone tends to become bare
because of excessive shading from the top of
the tree. The difficulty in maintaining the
fusetto within the limits set by the spacing
and the loss of productivity when the trees
become too shaded has limited its adoption.
Today this system is only marginally adopted
in Italy (Sansavini and Neri, 2005). A variant
 Orchard Planting Systems 275

Fig. 11.4. Central leader tree form.

Fig. 11.5. Fusetto orchard.

276 L. Corelli-Grappadelli and R.P. Marini
of the fusetto has been proposed for southern
Italy by Caruso et al. (1997) for early-ripening
peaches in a long-season Sicilian growing dis-
trict. The ‘dwarfed fusetto’ (as it is called by
proponents) is a shorter tree than fusetto, and
it is planted to lower density (about 700 trees/
ha) with reduced height (2.5–2.8 m). Pruning
is done immediately after harvest, when fruit-
bearing wood is removed and headed back to
the lowest current-season lateral. After har-
vest, the canopy is thus formed by short
‘stubs’ carrying current-season growth which
will become next year’s bearing wood. The
long vegetative season and very early (low-
chill) cultivars are very important features of
Fig. 11.6. Fusetto tree form.
this system, which relies on having sufficient
time in the season after harvest to develop the
laterals that will carry next year’s crop. In
their comparison of this system with a variant
of the tatura trellis (free-standing tatura), how-
ever, the authors found that the dwarfed fusetto
was not as satisfactory as the other system, since
its per-hectare yields were about half without
any gain in fruit quality (Caruso et al., 1997).
Y-shaped tree
To form Y-shaped trees, the leader is removed
and only two primary scaffold branches are
 Orchard Planting Systems
retained, which are trained to grow perpen-
dicular to the row axis (Fig. 11.7/Plate 75).
This system is suited to high densities, up to
2000 trees/ha, but most commonly densities
range between 900 and 1500 trees/ha, with
spacing between 4.0–4.5 m by 1.2–1.5 m (Chal-
mers et al., 1978; Corelli Grappadelli et al.,
1986; Caruso et al., 1997, 2003). This tree form
was first adopted many years ago (Baldassari,
1950), but interest in it has been renewed by
its potential for very high light interception
(above 70%; Nuzzo et al., 2000). This system
intercepts twice as much radiation as the pal-
mette and the delayed vase in the second leaf
(40% versus 20% incoming radiation) and main-
tains greater levels throughout the life of the
orchard (65% versus 55% and 42% for Y-trellis,
delayed vase and free palmette at the ninth
leaf, respectively; Fig. 11.1). A variant of this
system is the so-called ‘V’, where trees are
planted at an angle from the horizontal, alter-
nated along the row, so as to form a V-shaped
canopy (Costa et al., 1989) perpendicular to the
277
row axis. Densities for this system can be quite
high, reaching 2000 trees/ha (4 m × 1.25 m).
The tatura trellis (Chalmers et al., 1978) was
developed for mechanical harvest of cling
peaches and scaffold limbs are supported
with a wire trellis. The Kearney Agricultural
Center perpendicular-V (KAC-V) was devel-
oped for hand harvest and is not supported
(DeJong et al., 1994). Typical tree densities are
900–1200 trees/ha and trees are about 5.5 m
tall. The MIA trellis is a modification of the
V-shaped tree. The A-shaped canopy is devel-
oped by orienting the leaders at 60° from verti-
cal and leaning trees in adjacent rows towards
each other. The Y-trellis developed in south-
ern Italy (Caruso et al., 2003) features a wider
angle between the branches (45°, compared
with 35° for the tatura trellis), which are allowed
to converge and touch each other in the mid-
dle of the alley row, giving this system light
interception in excess of 70% (Nuzzo et al.,
2000) and very high productivity (Caruso
et al., 2003). A particular version of the Y-trellis
Fig. 11.7. Y-shaped tree form.
(Courtesy of D.R. Layne, Clemson,
South Carolina, USA.)
278 L. Corelli-Grappadelli and R.P. Marini
has been proposed by Caruso et al. (1997) for
low-chill, early-season cultivars in a variant
of the meadow orchard concept proposed by
Erez (1976, 1978). The goal is to restrict growth
and retain tree fertility, despite the smaller
tree size, which is trained without a trellis.
This free-standing tatura was dubbed the
‘fsTatura’ by the authors, who adopt a prun-
ing technique calling for the heading of all
1-year-old wood above the most proximal,
current-season lateral shoot. This is done
immediately after harvest; with very early-
season cultivars, the long season remaining
allows for the differentiation of flower buds
on these shoots. The next spring, the shoots
bloom and bear fruit. By adopting a vigorous
rootstock (‘GF 677’) this cycle can be main-
tained. Yields in this system can be high,
because of the high tree density (2000 tree/
ha). The average yield for the first three har-
vest seasons was 18 t/ha, but the yield in the
first harvest (the third year from planting)
was 14 t/ha (Caruso et al., 1997). The Y-trellis
is the system used in protected cultivation in
southern Italy because this shape is well
adapted to the tunnels used, where the sup-
port arches are set along the row, and the
vegetation expands out towards the dome of
the tunnel. Tree spacing is reduced (4.5 m × 1.5
m) and tree densities are fairly high (about
1500 trees/ha), favouring precocity and high
yields. For ‘Armking’ nectarine on seedling
rootstock in the Basilicata region of southern
Italy, Fideghelli et al. (1988) reported yields of
5.0 t/ha in the second season and up to about
30 t/ha at full production.
Meadow orchards
The meadow orchard, originally developed
by Hudson (1971), was an ultra-high-density
(about 100,000 trees/ha) full field-cover orchard
for apples intended for mechanical harvest by
mowing the trees with their fruit as grass in a
meadow. Production was based on a biennial
cycle starting with a vegetative flush followed
by harvest the next season, after which the
trees were cut back to a stump, to restart the
cycle. Theoretically, the two primary advan-
tages for the meadow orchard are that the
yields of young orchards would be high
because the orchard space is filled quickly,
and the fruiting zone would remain near the
ground because the tree top would be peri-
odically removed. The apple meadow orchard
was not economical because apple trees
cropped only every other year.
Peaches seem better suited to a meadow
orchard system for two reasons: (i) establish-
ment costs are relatively low because peach
trees can be produced from rooted cuttings
(Couvillon and Erez, 1980); and (ii) peaches
fruit on 1-year-old wood and may crop annu-
ally. Erez (1988) evaluated two variations of
the meadow orchard in Israel: the ‘mechanized
system’, developed for mechanized harvesting
(Erez, 1976, 1978), and the ‘intensive system’
(Erez, 1988). The former system involved a
2-year cycle similar to apple, whereas in the
intensive system the tree was trained to two
main shoots and each winter one of the two
shoots was headed back to a short stump to
allow regeneration of new growth and flower
bud formation during the growing season.
Therefore, each side of the tree fruited every
second year. The second system appeared
better for flower bud differentiation, fruit set
and yields, and suitable for late-season culti-
vars as well, whereas the mechanized system
required very long growing seasons and was
thus limited to early-maturing cultivars. Opti-
mum tree spacing for the mechanized system
is probably about 1.5 to 1.8 m between rows
and 0.6 m between trees within the row. For
the intensive system, trees should be spaced
about 1.5 m × 0.5 m. A modified mechanized
system was tested in Georgia, USA (Couvil-
lon and Erez, 1982) with 9810 or 3924 trees/
ha. Early-maturing, but not late-maturing, cul-
tivars regenerated enough flower buds for
annual production. Trees became deficient in
several elements and constant fertigation was
needed to alleviate the problem. Although
yields were high for a young orchard, fruit
size was too small; the percentage of fruit >5.7
cm in diameter was only 12%, 46% and 7% for
‘Redhaven’, ‘Loring’ and ‘Blake’, respectively.
In Florida, USA, Crocker et al. (1988)
tested a meadow orchard with 3333 trees/ha,
where trees were topped after harvest at
0.75 m above ground to leave the basal por-
tion of scaffold branches. This system’s annual
cropping and yields in young orchards were
 Orchard Planting Systems
considerably higher than for traditional peach
orchards. In Sicily, Caruso et al. (1997) have
proposed a similar approach, which tries to
exploit the peach vegetative vigour for early-
season cultivars: immediately after harvest,
all the fruiting wood is shortened, cut above
the most basal lateral, current-season growth.
This produces a tree which only has short
stumps with fruiting wood at the start of the
new season, similar to some of the grape
training systems. Testing with fusetto and
modified tatura has indicated the latter to be
better suited to this type of management.
Another alternative to encourage annual crop-
ping is to cut alternate trees in the row after
harvest (Evert, 1988). In Sicily, a study was con-
ducted on a commercial farm where meadow
orchard trees were grown under greenhouse
cultivation (Bellini et al., 2000). Tree densities
were 5000 trees/ha for the small vase trees
and 3300 trees/ha for Y-trellis. ‘Maravilha’, a
low-chill cultivar, was used. Trees were highly
productive: average annual yields during the
first four seasons were 25 and 33 t/ha for trees
trained as Y-trellis and small vase, respec-
tively. The authors indicated that trees could
be successfully grown at these very high den-
sities, although maintaining a productive can-
opy required summer pruning several times
per season, particularly with the denser small
vase. Despite the potential, however, virtually
no commercial meadow orchards exist, because
of problems with tree survival and the lack of
suitable, low-cost mechanical equipment that
would be needed for their management.
Protected cultivation
This type of peach cultivation has been widely
practised in southern Italy over the last few
decades (and now also in China; see Chapter
2 of this volume), where climatic conditions
allow the use of PVC tunnels (without heat-
ing) to advance ripening, particularly of early
cultivars. Fruit can bring high prices because
they reach the market when no other peaches
are available. The first studies in protected
cultivation were reported in the early 1970s
(Sansavini, 1974). In the early 1980s several
research trials were established (Caruso et al.,
1989; Bellini et al., 1997, 2000), dealing with
279
different solutions regarding the type of
greenhouse, cultivars and training systems.
PVC-covered greenhouses were trialled, as
well as tunnels. Today, most of this industry
is under tunnels with the support arches
placed in the row, and trees are trained to
Y-trellis at about 1500 trees/ha (Fideghelli
et al., 1988). Early-season cultivars are used,
such as ‘Armking’, along with low-chilling
requirement cultivars, because these allow
the earliest pickings. A typically grown culti-
var, according to these criteria, is ‘Maravilha’.
Low-chill cultivars tend to have very high
fertility index and percentage fruit set (Car-
uso et al., 1989; Bellini et al., 1997, 2000), which
is not frequently the case with high-chill ones.
There is a requirement for careful summer
pruning of these trees, in order to maintain
good flower bud differentiation and good
fruit quality traits; however, no authors report
increased difficulties in management of the
trees due to the enclosure, as far as pests and
fungi are concerned. The vast majority of
commercial orchards are based on the Y-trellis,
although other systems have been trialled, for
example the modified meadow orchard or
low central axis (MMO and LCA, respec-
tively; Caruso et al., 1989). The MMO is the
same as described by Erez (1976), where the
fruiting shoot is cut back after harvest and a
new sprout is selected and trained for next
year’s crop. The LCA, on the other hand, has
a central axis where all fruiting shoots are
inserted directly on the leader and they are
pruned back to short stubs immediately after
harvest, in order to stimulate the emission of
a new fruiting shoot from below the cut made.
This shoot will bear next year’s crop before
being removed and the cycle restarted. In
general, these studies have indicated that it is
difficult to retain high productivity with the
MMO, since the continuous removal and for-
mation of a new canopy rapidly depletes the
tree of its bearing potential, and crops decline
in a few years. Overall, there is agreement
over the Y-trellis being a better performer (a
fact certainly not lost to the industry) because
of its early, high crops, sustained over the
years, at a density much lower than its coun-
terparts. An additional benefit is that it does
not require summer pruning to be performed
immediately after harvest (such as is the case
280 L. Corelli-Grappadelli and R.P. Marini
with the other ones), since here summer prun-
ing is only needed to improve flower bud dif-
ferentiation on existing shoots by allowing
more light inside the canopy. In addition to
these considerations, this training system lends
itself quite well to the structure of the tunnel,
because the supporting arches can be placed
within the tree rows and the inter-row is free for
movement, an important consideration when
operating equipment in otherwise fairly
restricted space. The tunnels are covered after
completion of the chill requirement, which is
normally during January in southern Italy. Fol-
lowing closure of the tunnel, flowering occurs
in early February and, depending on the culti-
var, harvest commences in mid- to late April.
Thus fruit can be harvested about a month ear-
lier than in traditional orchards. Yields are sat-
isfactory, especially considering the premium
prices received for early fruit. Fideghelli et al.
(1988) reported an average yield of 20.9 t/ha
over the first 4 years for ‘Armking’.
Studies were conducted in Nova Scotia
and Ontario, Canada to assess the feasibility
of growing peaches in heated greenhouses,
with trees grown in large containers and in
the soil in the two areas, respectively (Crowe
et al., 1987; Miles and Leuty, 1988). In both
areas, growing peaches outdoors is restricted
by low winter temperatures and short grow-
ing seasons. In Ontario, three training sys-
tems were evaluated for 4 years and the tatura
trellis had the highest yields. Compared with
similar trees in the field, yields were 40–80%
higher in the greenhouse. Establishment costs
were more than three times higher for pro-
tected culture, but due to higher yields and
higher prices for early-season fruit, the pro-
tected culture system was more profitable than
the standard system. Given today’s energy
prices, however, it is unlikely that the eco-
nomics of a heated greenhouse peach orchard
would justify its adoption.
The study in Nova Scotia, Canada was
carried out with early-season peach cultivars
grown in large containers and overwintered
in heated poly-covered houses (Crowe et al.,
1987). Average annual yield was about 16 to
19 kg/tree from the third to the seventh year
after planting. Although these yields were less
than 30% of what would be expected of field-
grown trees in traditional peach-growing
regions, the system was considered to be prof-
itable in regions where traditionally grown
peaches are normally not profitable.
Protected cultivation is a commercial
practice in southern Italy, where low-cost, non-
heated, PVC-covered tunnels can be used
because the climatic conditions allow for suf-
ficient natural heating of the greenhouses. If
this were not the case, the cost of heating
would totally offset the economics of this type
of cultivation in that district. In areas where
heating is required, the economic feasibility
of protected peach culture may depend on
the price of fuel to maintain trees at appropri-
ate temperatures. Interest rates on the capital
required to build protective structures may
also impact the profitability of these systems.
11.5 Effects of Training System and
Planting Density on Yield
Tree form effect on yield
Comparison of different shapes is important
to evaluate tree performance, especially a sys-
tem’s capacity for early fruit production, a
vital trait in highly intensive orchards. Often
the training systems requiring the least initial
pruning are most precocious and tend to have
higher cumulative yields over the first four to
six seasons. Many trials have demonstrated
this advantage for more ‘natural’ systems.
The free shape, which is nearly non-pruned
initially, the fusetto and sometimes the delayed
vasette all require less early pruning and are
more precocious than systems such as the
palmette or the Y-trellis (Table 11.2) (Sansavini
et al., 1980, 1985; Bargioni et al., 1985; Bassi et al.,
1985; Corelli Grappadelli et al., 1986; Blay
Coll, 1988; Toribio Mancebo, 1993). The higher
yields of these systems are normally restricted
to the first 4 to 6 years; as orchards age yields
tend to level off and become similar between
these systems (Bassi et al., 1985).
In other studies, the KAC-V and the
cordon systems planted at the same density
produced similar yields per hectare (Gross-
man and DeJong, 1998). When planted at low
densities, open vase trees had higher yields
per tree, but lower yield per unit of land area
 Orchard Planting Systems
under the canopy or per unit of canopy vol-
ume than did central leader trees (Marini et al.,
1995). In another experiment central leader
and open vase forms were compared at low
density (370 trees/ha) and at moderate den-
sity (740 trees/ha) (Marini and Sowers, 2000).
The interaction between tree form and tree
density was not significant, and tree form had
little effect on cumulative yield. Allison and
Overcash (1987) compared central leader trees
with a palmette trellis system planted at 1292
trees/ha. Training system did not influence
cumulative yield during the first 4 years.
Menzies (1988) compared a palmette hedge-
row with the Lincoln canopy, both at 1000
trees/ha: after 6 years cumulative yield was
about 25% greater for the palmette system. Tay-
lor (1988) compared six different training sys-
tems, all with 1680 trees/ha. Low temperatures
eliminated the crop in the third year. Yields in
the fourth year ranked from highest to lowest
as follows: open vase > modified Belgian
fence > tatura trellis > perpendicular fan >
parallel fan > central leader. These results gen-
erally indicate that training system has little
impact on peach yield.
In apple, Robinson (1997) showed that
yield differences between Y-trellis, slender
spindle and palmette were related to light
interception, because yield per unit of inter-
cepted light energy was similar. In agreement
with this, whole-canopy photosynthesis of
peach trees was linearly related to the amount
of light intercepted by the tree (Giuliani et al.,
1998), irrespective of training system (Y-trellis,
sprint palmette and delayed vase). The Y-trellis
intercepted more light, had the highest pho-
tosynthetic rates and the highest yields over
nine cropping seasons (L. Corelli Grappadelli,
unpublished results). Therefore, when train-
ing systems planted at the same density do
not show much difference in yield over the
life of the orchard, this should not be surpris-
ing if they efficiently fill the allotted space
and intercept similar amounts of light.
Effect of planting density on yield
Conventional orchards are planned to fit the
dimensions of mature bearing trees. As a
281
result, trees require 2 years or more to fill their
allotted space and for the orchard to attain
maximum yields, even if the trees are not
pruned to hasten their development. Several
experiments were performed where tree den-
sity, but not tree form, was varied. While
some studies showed a positive response of
yield to density (Reeder et al., 1980; Miles
et al., 1999), several authors found that yield
per hectare increased less than proportionally
to tree density (Giulivo et al., 1984; DeJong
et al., 1999; Marini and Sowers, 2000). Hutton
et al. (1987) compared three training systems,
each at three tree densities, and showed that
the effect of tree density on yield varied with
the training system, and in some cases increas-
ing density caused a decrease in cumulative
yield. Working with fusetto at varying tree
densities (1250, 1665 and 2500 trees/ha) in
Italy, Bargioni et al. (1985) concluded that 1665
trees/ha was optimum for their conditions.
The density of 2500 trees/ha had slightly
higher yield and was not justified from an
economic point of view. As the cost of the
trees is very often quite high, adding up to a
substantial portion of the total planting costs
in high-density orchards (particularly so for
patented cultivars), it is easy to see that den-
sity does have an important impact on the
financial aspects of peach growing.
Since maximum orchard productivity
cannot be attained until trees fill their allotted
space, the larger that trees are allowed to
grow the further apart they must be planted,
and the longer it will take to reach full pro-
duction. Planting small trees at higher tree
densities combined with minimal pruning
encourages earlier yields, but once trees fill
the allotted space, the capacity for yield will
be limited by the amount of light that can be
intercepted. That is why in virtually all the
trials there is a poor relationship between the
cumulative yield increase and tree density
increase after the first few seasons.
Results with apples are similar. Palmer
et al. (1989) compared systems with densities
ranging from 2667 to 8889 trees/ha, trained as
single-, double- and triple-row spindles and
six-row beds of full field mini-trees. They found
after six cropping seasons that the highest-
density system, with 233% more trees/ha,
produced only 38% more fruit. However, in
282 L. Corelli-Grappadelli and R.P. Marini
the year after planting these trees yielded 14
to 20 t/ha, compared with 5 to 11 t/ha for the
other systems. From the fourth season on,
when the orchards were at full bearing, there
were no differences in yield per year across
all training systems.
Training systems with small trees are bet-
ter suited to high densities and in general
they tend to have high early yields. Later on
this advantage is lost and the risk of losing
production due to excessive competition
becomes a factor to be carefully evaluated.
Because of the interaction between tree behav-
iour with site environmental conditions, the
ideal tree density will vary with site and train-
ing system. Regardless of training system, in
most of the studies with reported increases in
yield, these were usually relatively minor as
tree densities increased above 1000 trees/ha.
Effect of tree density plus tree form on yield
As discussed above, the choice of training
system dictates the spacing that can be used
and often it also defines the training strategy,
i.e. the sequence of summer and winter prun-
ing events that will lead to a fully developed
tree. As a result, yield may be dramatically
affected by the combination of these two fac-
tors, but not always in favour of the higher
densities. Often the yields in the early years
are in favour of the higher densities, irrespec-
tive of the system (Phillips and Weaver, 1975;
Hayden and Emerson, 1988), but later on
these differences may disappear or be reversed.
Leuty and Pree (1980) continued the trial of
Phillips and Weaver (1975) for another 4 years
and found that cumulative yields were related
to tree density only during the first three fruit-
ing years. After 9 years, increasing the tree
population from 397 to 1157 trees/ha (290%)
resulted in only a 12% increase in cumulative
yield per hectare. Pruning and thinning costs
were also related to tree density. They con-
cluded that the benefit of high density was
mostly in the early years, but high density
offered little economic advantage over the life
of the orchard. Kappel et al. (1983) found fairly
similar cumulative yields, over a 10-year period,
for oblique fan (567 trees/ha), modified central
leader (606 trees/ha), canted oblique fan (969
trees/ha) and open centre (381 trees/ha) (125,
111, 104 and 92 t/ha, respectively).
Increasing tree density can have benefi-
cial effects at very low densities, because at
these wide spacings the time needed to fill the
space allotted to the tree is much longer and
therefore the performance of higher-density
orchards is superior for longer periods of
time. In their study, Layne et al. (1981) found
that yield increased by up to 96% as tree den-
sity increased from 266 to 536 trees/ha. Hut-
ton et al. (1987) compared three systems and
found that the optimum tree density appeared
to vary with tree form and was 1732 trees/ha
for palmette hedgerow, 1234 trees/ha for tat-
ura trellis and 1110 trees/ha for MIA trellis.
Menzies (1988) reported higher cumulative
yields for 6 years for palmette hedgerow (1000
trees/ha) and tatura trellis (2500 trees/ha)
and lowest for open vase (500 trees/ha), with
intermediate yields for Lincoln canopy (1000
trees/ha) and central leader (666 trees/ha). A
similar advantage of high-density over low-
density systems was reported by Grossman
and DeJong (1998) working with trees trained
as KAC-V, cordon (1196, 919 trees/ha) and
open vase (299 trees/ha). Similar results were
reported in a different study by DeJong et al.
(1992). This study concluded that the practi-
cal advantages and disadvantages of the
high-density systems are probably less related
to crop yield than to orchard management
considerations such as tree structural strength,
uniformity, access to ladder work and sim-
plicity of cultural operations.
The primary advantage of high-density
over low-density systems is that the orchard
space is filled quickly and yields of young
orchards are improved. DeJong et al. (1994)
compared the KAC-V (909 trees/ha) with the
traditional open vase system (298 trees/ha).
Cumulative yields for the first 3 years were
about 44 t/ha for the KAC-V and 24 t/ha for
the open vase, but after 8 years yields were
224 and 190 t/ha, respectively. Therefore, the
increased yield due to high tree populations
is temporary and the economic feasibility of
such systems may depend on the cost of trees
and the value of new cultivars where supply
has not satisfied demand.
The training strategy, which is a conse-
quence of the choice of training system, can
 Orchard Planting Systems
have a large impact on productivity, especially
in the critical early years. The no-heading-cuts
strategy demonstrates this. Open vase trees
are normally headed in their first season, and
this causes delayed and low initial yields. The
delayed vasette is not headed at planting, and
its early yields can be substantial. Delayed
vasette planted at 727 trees/ha had similar
yield per tree in the second season to fusetto
at the same density. Fusetto and Y-trellis planted
at 1454 trees/ha had lower yields. The study
was terminated after 4 years, due to tree mor-
tality following a winter freeze, but through
the third cropping season the delayed vasette
still had high yields per tree. However, because
of the planting density, this form had one of
the lowest cumulative yields for the three
seasons. Cumulative yields per hectare were
52.9, 64.4 and 50.1 t for the delayed vasette,
fusetto and Y-trellis with 727 trees/ha, respec-
tively. The fusetto and Y-trellis at 1454 trees/
ha had 96.1 and 77.3 t/ha, respectively (Corelli
Grappadelli et al., 1986).
High tree density, almost irrespective of
training system, appears to be advantageous
for yields in the early years. This is the result
of the training techniques used, which stimu-
late early production, and the high number of
trees per hectare. This advantage may be lost
over the life of the orchard, particularly if the
orchard is planted at intermediate densities
and if it is maintained for more than 20 years.
For this reason the choice of training system
may have less to do with tree performance
than with the equipment on the farm, labour
availability, proficiency in training the tree
into a given form, the cost of the trees and of
establishing and maintaining various systems,
and their impact on fruit size and quality.
11.6 Future Trends
Researchers and commercial producers have
extensively evaluated peach orchard systems.
Unless new cultivars, rootstocks or plant growth
regulators become available for controlling
tree vegetative growth, future orchards will
consist of low or moderate tree densities. The
most common tree training systems (i.e. the
open vase and its derivatives in the western
283
hemisphere and much of Europe, the palmette
in some European countries, the Y-trellis) will
continue to be adopted, because radical depar-
tures from these systems are not foreseeable.
Certainly, training and management strategies
will continue to evolve to address the continu-
ing need to improve early crops, to alleviate the
economic burden of peach growing in those
countries. It can be expected that peach orchard
systems will evolve differently in different parts
of the world, reflecting the economic and tech-
nological conditions in the different areas.
Factors other than strictly horticultural
or physiological will be of prime importance
in determining the choice of the tree form.
The previous experience with a given system,
the cost and availability of land and skilled
labour, and ownership of previously pur-
chased materials and equipment are certainly
equally important determinants of these deci-
sions. Where land is available and relatively
inexpensive, simply planting more hectares
can enhance farm income, even if low-density,
extensive (as opposed to intensive) systems
are adopted. Where new orchard land is very
expensive and largely unavailable, on the
other hand, the pressure will be on increasing
the farm income by increasing returns per
unit of land area or per unit of input.
Most developing countries with peach
industries are located in the southern hemi-
sphere and farmers obtain high wholesale
prices for fresh fruit exported to developed
countries during their off-season. Both land
and labour are available and inexpensive, but
labour is unskilled. Tree uniformity within an
orchard will facilitate standardizing orchard
operations performed by non-skilled labour.
Orchard systems will evolve slowly unless
economic conditions change.
Small orchards dominate the peach indus-
try in most of Europe and orchard owners do
much of the routine work; farmers must per-
form most of the orchard operations because
labour is very expensive. Although land and
labour are already being used efficiently, there
will be increasing economic pressure to
improve efficiencies. Intensive orchard prac-
tices, performed by highly skilled labour, will
be required to improve early bearing and
yields per unit of land area. Minimal pruning,
summer pruning and cropping in the early
284 L. Corelli-Grappadelli and R.P. Marini
seasons are just examples of the shift in the
techniques employed to obtain early produc-
tion. These orchards, heavily exploited from
early on, will be less durable than traditional
ones; this is not perceived as a problem since
it allows for cultivar renovation, although it
poses the increasing problem of replant sites
(release of allelopathic compounds, increases
in soil-borne diseases, depletion of soil nat-
ural resources, etc.), which do lower tree per-
formance.
Peach growers in western North America
grow mostly fairly tall (4.0 to 5.0 m) open vase
trees that are planted at about 300 trees/ha.
North American orchards are relatively large
and workers are unskilled. Therefore, trees
within an orchard will have to be uniform to
facilitate orchard worker training. The cost of
labour is increasing rapidly and California
growers are starting to reduce tree height. As
land and labour become increasingly scarce
and expensive, peach orchard systems probably
will evolve towards those found in Europe.
Some type of hedgerow system that allows use
of platforms (Fig. 11.8/Plate 76) may domi-
nate the peach industry and, where platforms
Fig. 11.8. Modern picking platform.
are not used, tree height will be limited to less
than 3.0 m. At higher latitudes, where trees
grow slowly and winter injury results in early
tree mortality, tree densities will increase to
allow rapid filling of orchard space.
Modifications of existing orchard systems
and development of new systems will require
research breakthroughs. Rootstocks that pro-
vide a range of tree vigour and enhance carbon
partitioning, such as those available for apple,
may allow development of novel orchard sys-
tems. Plant growth regulators to control flow-
ering and vegetative growth, crop load or fruit
ripening could be utilized in new orchard sys-
tems. Growers need trees that produce high
yields of uniformly high-quality fruit that can
be harvested tree-ripe. Improvements in other
aspects of fruit quality such as fruit size, colour,
soluble solids concentration, flesh firmness
and high concentrations of health-promoting
compounds may improve peach consump-
tion and lead to increased prices. There is a
need for long-term studies to determine the
profitability of various orchard practices and
orchard systems. More information on funda-
mental relationships in peach tree physiology
 Orchard Planting Systems 285

is needed to develop new systems or to modify
systems. The relationship between light inter-
ception and yield per unit land area has not
been elucidated in peach. The scant data exis-
ting are relative to single-season, short-term
observations; no long-term studies have been
carried out comparing light interception and
marketable yields of the leading training
systems. Currently peach growers do not
have fruit thinning strategies, nor practical
methods for assessing in real time the prog-
ress of the crop and the performance of the
orchard. A better understanding of partition-
ing is needed to develop better strategies for
hand thinning and pruning that may alter
carbon partitioning.

References

Allison, M.L. and Overcash, J.P. (1987) Factors affecting hedgerow peach orchard establishment. Journal of
the American Society for Horticultural Science 112, 62–66.
Baldassari, T. (1950) La potatura dei fruttiferi. In: Zucchini, M. (ed.) Proceedings of II Convegno Provinciale
Frutticolo. SATE, Ferrara, Italy, pp. 71–76.
Bandoli, A. (1980) Untitled. In: Sansavini, S. (ed.) Proceedings of XV Convegno Peschicolo. Litografica Faenza,
Faenza, Italy, pp. 312–313.
Bargioni, G., Loreti, F. and Pisani, P.L. (1983) Performance of peach and nectarine in a high density system in
Italy. HortScience 18, 143–146.
Bargioni, G., Pisani, P.L. and Loreti, F. (1985) Ten years of research on peach and nectarine in a high density
system in the Verona area. Acta Horticulturae 173, 299–310.
Bassi, D., Brighenti, G. and Nardi, V. (1985) Training systems of Klamt cling peach: performance after 8 years – IV
Contribution. Acta Horticulturae 173, 339–348.
Bellini, E., Falqui, D. and Musso, O. (1997) La coltura protetta del pesco. L’Informatore Agrario 31, 41–50.
Bellini, E., Falqui, D. and Musso, O. (2000) Comparison between two training systems in peach protected
culture in Sicily. Acta Horticulturae 513, 427–433.
Belluau, E. and Lemaire, A. (1986) Evaluation of nine years experimenting on high-density peach orchard in
south east of France. Acta Horticulturae 160, 327–340.
Blay Coll, J. (1988) Formación del melocotonero en un solo eje principal (fuseto). Fruticultura Profesional
15(5/6), 3–17.
Borras Escriba, F. (2001) La peschicoltura spagnola nel mercato globale. In: Sansavini, S. and Lugli, S. (eds)
Proceedings of XXIV Convegno Peschicolo. Grafiche MDM, Forlì, Italy, pp. 31–34.
Cain, J.C. (1972) Hedgerow orchard design for most efficient interception of solar radiation. Effects of tree
size, shade, spacing and row direction. Search Agriculture 2, 1–14.
Campbell, R.C. and Marini, R.P. (1992) Light environment and time of harvest affect ‘Delicious’ apple fruit
quality characteristics. Journal of the American Society for Horticultural Science 117, 551–557.
Caruso, T., Di Marco, L., Giovannini, D. and Motisi, A. (1989) Trials on training systems for greenhouse
cultivated peach trees. Acta Horticulturae 243, 345–352.
Caruso, T., Giovannini, D., Marra, F.P. and Sottile, F. (1997) Two new planting systems for early ripening
peaches (Prunus persica L. Batsch): yield and fruit quality in four low-chill cultivars. Journal of Horticultural
Science 72, 873–883.
Caruso, T., Di Vaio, C., Guarino, F., Motisi, A. and Nuzzo, V. (2003) Le tipologie d’impianto per la peschicoltura
intensiva nell’Italia meridionale. In: Marra, F.P. and Sottile, F. (eds) Proceedings of IV Convegno Nazion-
ale sulla Peschicoltura Meridionale. Panuzzo Prontostampa, Caltanissetta, Italy, pp. 44–51.
Chalmers, D.J., van den Ende, B. and van Heek, L. (1978) Productivity and mechanization of the ‘Tatura
Trellis’ orchard. HortScience 13, 517–521.
Cole, S.W. (1849) The American Fruit Book. John P. Jewett, New York.
Corelli Grappadelli, L. and Sansavini, S. (1989) Light interception and photosynthesis related to planting
density and canopy management in apple. Acta Horticulturae 243, 159–174.
Corelli Grappadelli, L., Brighenti, G., Palara, U., Ravaioli, F. and Sansavini, S. (1986) Esperienze su forme di
allevamento del pesco per medie ed alte densità di impianto. Rivista di Frutticoltura 12, 55–60.
Costa, G., Miserocchi, O., Succi, F. and Martelli, S. (1989) Forme di allevamento del pesco per impianti ad
elevata densità di piantagione. Rivista di Frutticoltura 1, 63–66.
286 L. Corelli-Grappadelli and R.P. Marini

Couvillon, G.A. and Erez, A. (1980) Rooting, survival, and development of several peach cultivars propagated
from semi hardwood cuttings. HortScience 15, 41–43.
Couvillon, G.A. and Erez, A. (1982) Production potential and tree regeneration in a peach meadow orchard
in southeastern United States. Compact Fruit Tree 15, 166–169.
Crocker, T.E., Sherman, W.B., Young, M.J. and McDermont, L.G. (1988) An experimental high-density system
for low-chill peaches and nectarines. In: Childers, N. and Sherman, W.B. (eds) The Peach. Horticultural
Publishers, Gainesville, Florida, pp. 432–433.
Crowe, A.D., Henniger, D.R., Craig, W.E. and O’Regan, R.T. (1987) Containerized peach orchards are profit-
able and avoid winter injury. Compact Fruit Tree 20, 90–94.
DeJong, T.M. and Doyle, J.F. (1985) Seasonal relationships between leaf nitrogen content (photosynthetic ca-
pacity) and leaf canopy light exposure in peach (Prunus persica). Plant, Cell & Environment 8, 701–706.
DeJong, T.M., Day, K.R. and Doyle, J.F. (1992) Evaluation of training/pruning systems for peach, plum and
nectarine trees in California. Acta Horticulturae 322, 99–105.
DeJong, T.M., Day, K.R., Doyle, J.F. and Johnson, R.S. (1994) The Kearney Agricultural Center perpendicular
‘V’ (KAC-V) orchard system for peaches and nectarines. HortTechnology 4, 362–367.
DeJong, T.M., Tsuji, W., Doyle, J.F. and Grossman, Y.L. (1999) Comparative economic efficiency of four peach
production systems in California. HortScience 34, 73–78.
Erez, A. (1976) Meadow orchard for the peach. Scientia Horticulturae 5, 43–48.
Erez, A. (1978) Adaptation of the peach to the meadow orchard system. Acta Horticulturae 65, 245–250.
Erez, A. (1988) Peach meadow orchard: two feasible systems. In: Childers, N. and Sherman, W.B. (eds) The
Peach. Horticultural Publishers, Gainesville, Florida, pp. 424–431.
Evert, D.R. (1988) A modified meadow system. In: Childers, N. and Sherman, W.B. (eds) The Peach. Horticul-
tural Publishers, Gainesville, Florida, p. 434.
FAOSTAT (2004) ProdStat: Crops. http://faostat.fao.org/site/567/default.aspx (accessed October 2004).
Fideghelli, C. (1969) Confronto tra i sistemi di allevamento a vaso ed a palmetta del pesco. In: Baldini, E. (ed.)
Proceedings of Convegno Giornate di studio sulla potatura degli alberi da frutto. Firenze, pp. 133–147.
Fideghelli, C., Monastra, F. and De Salvador, R.F. (1988) Evoluzione delle forme d’allevamento e delle
densità d’impianto in peschicoltura. In: Sansavini, S. (ed.) Proceedings of XVIII Convegno Peschicolo.
Grafiche MDM, Forlì, Italy, pp. 63–81.
Flore, J.A. and Kesner, C. (1982) Orchard design for stone fruit based on light interception. Compact Fruit Tree
5, 159–165.
Génard, M. and Baret, F. (1994) Spatial and temporal variation of light inside peach trees. Journal of the
American Society for Horticultural Science 119, 669–677.
Giovannini, D. and Monastra, F. (1995) Tipologie d’impianto e forme di allevamento per la peschicoltura
meridionale. In: Pavia, R., Della Strada, G. and Grassi, F. (eds) Proceedings of Convegno su ‘Ricerca e
innovazione per la peschicoltura meridionale. Interstampa, Rome, pp. 195–211.
Giuliani, R., Magnanini, E. and Corelli Grappadelli, L. (1998) Whole canopy gas exchange and light intercep-
tion of three peach training systems. Acta Horticulturae 465, 309–317.
Giulivo, C., Ramina, A. and Costa, G. (1984) Effects of planting density on peach and nectarine productivity.
Journal of the American Society for Horticultural Science 109, 287–290.
Grossman, Y.L. and DeJong, T.M. (1998) Training and pruning system effects on vegetative growth potential,
light interception, and cropping efficiency in peach trees. Journal of the American Society for Horticul-
tural Science 123, 1058–1064.
Hayden, R.A. and Emerson, F.H. (1988) High density plantings for peaches. In: Childers, N. and Sherman,
W.B. (eds) The Peach. Horticultural Publishers, Gainesville, Florida, pp. 404–411.
Hilaire, C. (2003) The peach industry in France: state of the art, research and development. In: Marra, F.P.
and Sottile, F. (eds) Proceedings of the First Mediterranean Peach Symposium. Panuzzo Prontostampa,
Caltanissetta, Italy, pp. 27–34.
Hudson, J.P. (1971) Meadow orchards. Agriculture 78, 157–160.
Hugard, J. (1981) High density plantings in French orchards: development and current achievements. Acta
Horticulturae 114, 300–307.
Hugard, J. (1986) Main features of fruit production in the French Mediterranean area. Acta Horticulturae 160,
15–22.
Hutton, R.J., McFadyen, L.M. and Warwick, J.L. (1987) Relative productivity and yield efficiency of canning
peach trees in three intensive growing systems. HortScience 22, 552–560.
Jackson, J.E. (1967) Variability in fruit size and colour within individual trees. Annual Report of East Malling
Research Station for 1966, 110–115.
 Orchard Planting Systems 287

Jackson, J.E. (1980) Light interception and utilization by orchards. Horticultural Reviews 2, 208–267.
Jackson, J.E. and Middleton, S.G. (1988) Progettazione del frutteto per la massima produttività e qualità. In:
Youssef, J. (ed.) Proceedings of Conference ‘Coltura del melo verso gli anni ‘90’. Chiandetti, Udine, Italy,
pp. 309–320.
Jackson, J.E. and Palmer, J.W. (1972) Interception of light by model hedgerow orchards in relation to latitude,
time of year and hedgerow configuration and orientation. Journal of Applied Ecology 9, 341–357.
Kappel, F., Flore, J.A. and Layne, R.E.C. (1983) Characterization of the light microclimate in four peach hedge-
row canopies. Journal of the American Society for Horticultural Science 108, 102–105.
Lakso, A.N. (1994) Apple. In: Schaffer, B.S. and Anderson, P.C. (eds) Handbook of Environmental Physiology
of Fruit Crops. Vol. I. Temperate Crops. CRC Press, Boca Raton, Florida, pp. 3–42.
Lakso, A.N. and Musselmann, R.C. (1976) The effects of cloudiness on interior diffuse light in apple trees.
Journal of the American Society for Horticultural Science 101, 642–644.
Layne, R.E.C., Tan, C.S. and Fulton, J.M. (1981) Effect of irrigation and tree density on peach production.
Journal of the American Society for Horticultural Science 106, 151–156.
Leuty, S.J. and Pree, D.J. (1980) The influence of tree population and summer pruning on productivity, growth,
and quality of peaches. Journal of the American Society for Horticultural Science 105, 702–705.
Lewallen, K.S. and Marini, R.P. (2003) Flesh firmness and ground color in peach as influenced by light and
canopy position. Journal of the American Society for Horticultural Science 128, 163–170.
Loreti, F., Massai, R. and Morini, S. (1989) Further observations on high-density nectarine plantings. Acta
Horticulturae 243, 353–360.
Marini, R.P. (1985) Vegetative growth, yield, and fruit quality of peach as influenced by dormant pruning,
summer pruning, and summer topping. Journal of the American Society for Horticultural Science 110,
133–139.
Marini, R.P. and Barden, J.A. (1981) Seasonal correlations of specific leaf weight to net photosynthesis and
dark respiration of apple leaves. Photosynthetic Research 2, 251–258.
Marini, R.P. and Barden, J.A. (1982) Light penetration on overcast and clear days, and specific leaf weight in
apple trees as affected by summer or dormant pruning. Journal of the American Society for Horticultural
Science 107, 39–43.
Marini, R.P. and Corelli Grappadelli, L. (2006) Peach orchard systems. Horticultural Reviews 32, 63–110.
Marini, R.P. and Marini, M.C. (1983) Seasonal changes in specific leaf weight, net photosynthesis, and chlo-
rophyll content of peach leaves as affected by light penetration and canopy position. Journal of the
American Society for Horticultural Science 108, 600–605.
Marini, R.P. and Sowers, D.S. (1990) Net photosynthesis, specific leaf weight, and flowering of peach as
influenced by shade. HortScience 25, 331–334.
Marini, R.P. and Sowers, D.S. (2000) Peach tree growth, yield, and profitability as influenced by tree form and
tree density. HortScience 35, 837–842.
Marini, R.P., Sowers, D.S. and Marini, M.C. (1991) Peach fruit quality is affected by shade during final swell
of fruit growth. Journal of the American Society for Horticultural Science 116, 383–389.
Marini, R.P., Sowers, D.S. and Marini, M.C. (1995) Tree form and heading height at planting affect peach tree
yield and crop value. HortScience 30, 1196–1201.
Mascanzoni, G. (1998) Evoluzione e problematiche del ‘vaso ritardato’ nel pesco. In: Sansavini, S. and Errani,
A. (eds) Frutticoltura ad alta densità. Edagricole, Bologna, Italy, pp. 237–245.
Menzies, A.R. (1988) Evolution of peach tree forms in New South Wales, Australia. In: Childers, N. and Sherman, W.B.
(eds) The Peach. Horticultural Publishers, Gainesville, Florida, pp. 446–465.
Miles, N.W. and Leuty, S.J. (1988) Protected peach culture. In: Childers, N. and Sherman, W.B. (eds) The
Peach. Horticultural Publishers, Gainesville, Florida, pp. 380–386.
Miles, N.W., Guarnaccia, R. and Slingerland, K. (1999) High density peach production in Ontario. New York
Fruit Quarterly 7(4), 1–5.
Monteith, J.L. (1977) Climate and the efficiency of crop production in Britain. Philosophical Transactions
Royal Society of London Bulletin 281, 277–294.
Nuzzo, V., Dichio, B., Palese, A.M. and Xiloyannis, C. (2000) Sviluppo della chioma ed intercettazione radia-
tiva in piante di pesco allevate ad Y trasversale ed a vaso ritardato nei primi tre anni dall’impianto. In:
Failla, O. and Piagnani, I. (eds) Proceedings of V giornate Scientifiche SOI. Edizioni Tecnos, Milan, Italy,
pp. 319–320.
Palmer, J.W., Bünemann, G., Sansavini, S., Wagenmakers, P.S. and Winter, F. (1989) The international planting
systems trial. Acta Horticulturae 243, 231–241.
Phillips, J.H.H. and Weaver, G.M. (1975) A high-density peach orchard. HortScience 10, 580–582.
288 L. Corelli-Grappadelli and R.P. Marini

Porpiglia, P.J. and Barden, J.A. (1980) Seasonal trends in net photosynthetic potential, dark respiration, and
specific leaf weight of apple leaves as affected by canopy position. Journal of the American Society for
Horticultural Science 105, 920–923.
Reeder, B.D., Bowen, H.H. and Aldred, W.H. (1980) Peach tree training and spacing. HortScience 15, 580–
581.
Rizzi, A.D. (1975) Training and modifying peach trees in California for mechanical handling. In: Childers, N.
and Sherman, W.B. (eds) The Peach. Horticultural Publishers, Gainesville, Florida, pp. 214–225.
Robinson, T.L. (1997) Interaction of tree form and rootstock on light interception, yield and efficiency of ‘Empire’,
‘Delicious’ and ‘Jonagold’ apple trees trained to different systems. Acta Horticulturae 451, 427–436.
Robinson, T.L. (2003) Apple-orchard planting systems. In: Ferree, D.C. and Warrington, I. (eds) Apples: Botany,
Production and Uses. CAB International, Wallingford, UK, pp. 345–407.
Robinson, T.L. and Lakso, A.N. (1991) Bases of yield and production efficiency in apple orchard systems.
Journal of the American Society for Horticultural Science 116, 188–194.
Royo Díaz, J.B. and Martínez Lopez, T. (1992) Sistemas de formación en melocotonero. Fruticultura Profe-
sional 46, 22–31.
Sansavini, S. (1974). Indirizzi tecnico-agronomici della peschicoltura romagnola. L’Italia Agricola 1–2,
1–47.
Sansavini, S. (1980) Impianti e allevamento del pesco: analisi e prospettive delle tendenze in atto. In: Sansavi-
ni, S. (ed.) Proceedings of XV Convegno Peschicolo. Litografica Faenza, Faenza, Italy, pp. 63–116.
Sansavini, S. and Neri, D. (2005) Forme di allevamento e potatura del pesco. In: Sansavini, S. and Fideghelli,
C. (eds) Manuale di Peschicoltura. Edagricole, Bologna, Italy, pp. 115–143.
Sansavini, S., Bassi, D. and Giunchi, L. (1980) Prove comparative di allevamento del pesco da industria. In:
Sansavini, S. (ed.) Proceedings of XV Convegno Peschicolo. Litografica Faenza, Faenza, Italy, pp. 267–
278.
Sansavini, S., Corelli Grappadelli, L. and Giunchi, L. (1985) Peach yield efficiency as related to tree shape.
Acta Horticulturae 173, 139–158.
Sansavini, S., Corelli Grappadelli, L., Costa, G., Lugli, S., Marangoni, B., Tagliavini, M., Ventura, M., Abeti, D.,
Ferali, S., Marani, G., Mascanzoni, G., Molducci, S., Proni, R., Sama, A., Spada, G., Vitali, S., Turroni, P.,
Minguzzi, A. and Randi, M. (2000) Ricostituzione degli impianti e nuovi indirizzi produttivi della pe-
schicoltura romagnola. In: Sansavini, S. and Lugli, S. (eds) Proceedings of XXIII Convegno Peschicolo.
Grafiche MDM, Forlì, Italy, pp. 62–74.
Taylor, B.H. (1988) Promising high density peach systems in Illinois. In: Childers, N. and Sherman, W.B. (eds)
The Peach. Horticultural Publishers, Gainesville, Florida, pp. 491–498.
Toribio Mancebo, F. (1993) Juntos, pero bien formados. Albear (1), 20–27.
 12 Crop Load Management

R.P. Marini1 and G.L. Reighard2

1Department of Horticulture, The Pennsylvania State University, University Park,
Pennsylvania, USA
2Department of Horticulture, Clemson University, Clemson, South Carolina, USA

12.1 Introduction 289

12.2 Fruit Growth and Development 290
12.3 Quantifying Crop Load 290
12.4 Modifying Crop Load 291
Pre-bloom 291
Bloom 293
Post-bloom 294
12.5 Crop Load Interactions with Stresses 297
Tree density 297
Water stress 297
Leaf-feeding arthropods 297
High temperatures 298
Low temperatures 298
12.6 Future Directions 298

12.1 Introduction were productive for only a few years and

Crop load management is an important aspect
of peach production. Most peach trees pro-
duce thousands of flowers and, if conditions
are favourable, may set several thousand fruit
per tree. If all these fruit are allowed to develop,
the weight of the fruit will break branches
and the fruit will be small and have low sugar
concentrations. To avoid overcropping, the
number of fruit per tree must be managed.
In descriptions of North American peach
production during the 18th and 19th centu-
ries there is no mention of fruit thinning (Cole,
1849; Fitz, 1872), but accounts indicate trees
orchards were replanted frequently. The rea-
sons for early tree mortality were probably
tree-boring insects, disease and winter damage,
as well as limb breakage due to overcropping.
Fruit thinning is now a standard com-
mercial practice, and fruit are most commonly
removed by hand. Hand removal of excess
fruit is the most expensive preharvest activity
in peach production, but it is absolutely nec-
essary to produce a saleable crop. During the
last half-century the peach industry has been
searching for less expensive ways to remove
unwanted fruit. This chapter provides a dis-
cussion of peach tree physiology, which is

© CAB International 2008. The Peach: Botany, Production and Uses

(eds D.R. Layne and D. Bassi) 289
290 R.P. Marini and G.L. Reighard
needed to fully understand the concept of
fruit thinning, and the literature leading to
the recommendation for fruit thinning is
reviewed. We also review the various meth-
ods that have been developed to thin peach
trees and some of the factors influencing the
efficacy of these practices, and we provide
some suggestions for future research.
12.2 Fruit Growth and Development
During the late season, especially after fruit
harvest, photoassimilate is translocated from
leaves to the woody portions of the tree, where
it is stored as starch. Spring growth of roots,
cambium, shoots and fruit all depend on the
reserve carbohydrates. Unlike some other deci-
duous fruit trees, peach trees bloom before
there is much leaf development. Whole-tree
photosynthesis is insufficient to support early-
season growth of the tree and fruit. Most of
the energy for early-season fruit growth is
derived from reserve carbohydrates, and
fruits must compete with other parts of the
tree and with each other for carbohydrates.
Peach fruit growth is traditionally divided
into three stages (Tukey, 1933; Chalmers and
van den Ende, 1975; Zucconi, 1986; Gage and
Stutte, 1991). The first stage is a period of rapid
fruit growth from bloom to about 50 days
after bloom, when the stony endocarp (pit)
hardens. The length of the first stage depends
on cultivar and temperature. Fruit growth dur-
ing this stage is primarily due to cell division,
but some cell expansion and intercellular space
formation occurs during the second half of
this stage. The duration of the second stage
depends on the maturity date of the cultivar.
Stage II may last a few days in very early-
maturing cultivars or up to 2 months in late-
maturing cultivars. During this stage there is
little increase in fruit size, but fruit dry weight
continues to increase due to endocarp devel-
opment, which requires a considerable amount
of assimilate. The third stage is a period of
rapid fruit size increase as the mesocarp cells
expand. This final stage is often called the ‘final
swell’, lasts several weeks before harvest and
is the time of rapid sugar accumulation.
Fruits compete with other growing parts of
the tree for assimilates. Grossman and DeJong
(1995a,b) identified periods of resource-limited
fruit growth. For an early-maturing cultivar,
resource limitation began about 38 days after
bloom and continued through harvest, which
was 84 days following bloom. For the late-
maturing cultivar, the first period was similar
to the early-season cultivar but a second period
occurred during the last 28 days of the final
swell, 126 to 154 days after bloom. Suppressed
fruit growth caused by resource limitations
was only temporary and fruit growth resumed
normally when resources became available.
The primary objective of crop load man-
agement is to minimize resource limitations
for fruit growth in proper balance with vegeta-
tive development. Because fruit size is related
to the number of cells per fruit and resources
are limited during the second half of stage I, it
is most important to remove excess fruit dur-
ing the first half of stage I to encourage cell
division of the remaining fruit. It is also
important to minimize stresses during the
final swell of late-season cultivars to ensure
maximum increases in fruit size.
12.3 Quantifying Crop Load
Peach thinning to improve fruit size has been
practised for many years. Early in the 20th
century it was known that fruit size was
related to the number of leaves per fruit, but
the nature of this relationship was not known.
Beach (1903; cited by Overholser and Claypool,
1931) reported a correlation between size of
peach leaves and fruit. Small-fruited cultivars
had smaller leaves than large-fruited culti-
vars. Based on apple thinning research in the
1920s, several researchers performed experi-
ments during the 1930s to determine the pre-
cise relationship between leaf area or leaf
number and size and quality of peach fruit.
Weinberger (1931) removed varying numbers
of fruit from ringed branches on peach trees
to develop eight ratios of leaf to fruit in early
July in Maryland. He reported a curvilinear
response, where fruit size increased at a
decreasing rate as leaves per fruit increased
from 10 to 75. From 30 to 75 leaves per fruit,
the response was nearly linear; the increasing
fruit volume per leaf was about 0.75 cm3 for
‘Elberta’ and 0.17 cm3 for ‘Early Crawford’.
 Crop Load Management
Sugars and acidity also increased with increas-
ing numbers of leaves per fruit. Based on this
study, 30 to 40 leaves per fruit seemed
adequate for producing large, high-quality
peaches. A similar experiment performed the
next season produced similar results for fruit
development; and shoots on limbs with high
leaf: fruit ratio produced more flower buds
and had higher starch levels in the wood
(Weinberger and Cullinan, 1932). Similar
results were reported for ringed limbs in
North Carolina, where fruit size and quality
increased as the number of leaves per fruit
was increased from 10 to 45 (Jones, 1932).
In Washington State, Overholser and Clay-
pool (1931) thinned branches to varying num-
bers of leaves per fruit in mid-June on ringed
and non-ringed limbs. There was a gradual
increase in fruit size as leaves per fruit increased
from 25 to about 80, but there was little increase
in fruit size with more than 85 leaves per fruit.
The increase in fruit size was greater and more
consistent for girdled limbs. These experi-
ments show that there is a positive relation-
ship between number of leaves per fruit and
fruit size, but there are problems with the meth-
odology and the leaf area per fruit required to
produce large fruit is still unknown. These
often-cited leaf:fruit ratios may not be very
accurate because there was no consideration
of the efficiency of a leaf and whether or not it
was in a high light environment. The leaf:fruit
ratios were established fairly early in the sea-
son. Normally leaf number per tree increases
until late season, so the number of leaves
early in the season is low but increases as the
season progresses. Most of these experiments
were performed with girdled limbs, and gir-
dling is known to improve fruit size and to
advance maturity. The crop load on non-treated
limbs was not reported and the non-treated
limbs may have influenced fruit on treat-
ment limbs. The authors usually did not indi-
cate if the fruit were all harvested on the same
date, or if they were harvested on the basis of
some maturity factor such as ground colour.
Fruit on heavy-cropped trees usually mature
later than fruit on light-cropped trees. Because
fruit continue to grow every day until harvest,
it is important to harvest fruit at the proper
stage of maturity to compare treatments in
peach experiments.
291
Shoemaker (1933) was the first to show
experimentally that early thinning and heavy
thinning to a spacing of 15 to 20 cm between
fruit was more profitable than later thinning
and less severe thinning. The increased fruit
size resulted in a higher crop value, which
offset the increased cost of thinning and the
slight loss of yield.
12.4 Modifying Crop Load
Numerous methods to reduce crop load have
been evaluated (Byers et al., 2003). Gould
(1918) summarized peach-growing practices
across the USA at the turn of the 19th century
and reported that the need for peach thinning
was a ‘complete oneness of opinion’ but ‘not . . .
in all cases with regard to practice’. No matter
which thinning method is employed, the pri-
mary objective is to reduce the crop to levels
that promote adequate fruit size at a reasonable
cost. Under commercial growing conditions,
maximizing fruit size can be accomplished by
selecting appropriate cultivars and by using
various cultural practices before bloom, dur-
ing bloom or after bloom.
Pre-bloom
Several factors occurring before bloom can
influence crop load and fruit size at harvest.
These factors include environmental condi-
tions and several cultural practices. In some
years low winter temperatures may kill some
of the flower buds, resulting in low numbers
of fruit per tree. A frost that kills a portion of
the flower buds or flowers during bloom will
result in improved fruit size of the remaining
fruit. However, peach growers try to minimize
the impact of low temperatures because freezes
and frosts often reduce crop loads below prof-
itable levels, but there are several methods to
reduce the number of blossoms to lower thin-
ning costs and improve fruit size at harvest.
Genetics
An important consideration when choosing a
cultivar is the influence of genetics on fruit
size. Commercial peach producers are aware
292 R.P. Marini and G.L. Reighard
that some cultivars produce larger-sized fruits
than others.
Scorza et al. (1991) identified the factors
associated with the inherent size of different
cultivars. They found that the number of cells
per fruit was higher for large-fruited cultivars
than for small-fruited cultivars. Differences in
fruit size were due to cell numbers rather than
cell size, and these differences were apparent
in the flower buds during the autumn before
bloom. Flower buds of large-fruited cultivars
had 2.5 times as many cells in mid-October
than small-fruited cultivars and the difference
in cell numbers per fruit remained consistent
until harvest. Before planting a new cultivar,
growers should learn as much as possible
about the inherent ability of a cultivar to pro-
duce fruit of marketable size with yields that
are profitable.
Growers must also realize that the ideal
crop load will vary for different cultivars.
Johnson and Handley (1989) thinned an early-
season, a mid-season and a late-season culti-
var to varying numbers of fruit per tree and
estimated average fruit weight at harvest.
They found that average fruit weight was lin-
early related to number of fruit per tree and
the decrease in fruit size for each additional
fruit per tree (slope of the line) was similar for
all three cultivars. In general, the average
fruit weight was reduced by 0.49 to 0.58 g for
each additional fruit that was left on the tree.
Slopes reported for other experiments were
similar (Cain and Mehlenbacher, 1956; West-
wood and Batjer, 1958), indicating that this
relationship is fairly robust for different culti-
vars, seasons and locations. For a given tree
size and crop load, average fruit weight is
determined by the inherent characteristics of
the cultivar. For example, with 600 fruit per tree,
average fruit weight was 100 g, 150 g and 220 g
per fruit for ‘May Crest’, ‘June Lady’ and ‘Ele-
gant Lady’, respectively. To obtain an average
fruit weight that is acceptable for fresh fruit
sales (130 g), ‘May Crest’ trees must be thinned
to only 190 fruit, ‘June Lady’ can be thinned to
1000 fruit per tree and ‘Elegant Lady’ can sup-
port 2000 fruit per tree. These results indicate
that potential marketable yield per tree, which
is a function of number of fruit per tree and
fruit size, will be higher for large-fruited cul-
tivars. However, potential marketable yield
per unit land area will likely depend on the
number of fruit per canopy volume per unit
land area, which is related to tree density and
tree size. This is an area that needs additional
work and should also include a comparison
of orchard training systems.
Cultivars also vary in their ability to pro-
duce flower buds. For a given cultivar, early-
season fruit set and the need for fruit thinning
are positively related to the number of flow-
ers per tree, and fruit set varies from year to
year depending on a number of factors. Peach
cultivars differ inherently in flower bud num-
ber (Blake, 1943; Werner et al., 1988). In a
peach cultivar trial in New Jersey, USA, the
number of flowers per metre of shoot length
varied from 30 for ‘Slaybaugh Special’ to 54
for ‘Springold’ (L. Miller, New Jersey, 1983,
personal communication). Flower density for
the same cultivar was also quite variable from
year to year. During a 5-year period the num-
ber of flowers per metre of shoot length for
‘Garnet Beauty’ ranged from three to 45.
These differences in flower density were due
to the tendency of a cultivar to produce flower
buds and the ability of buds of a cultivar to
survive the winter. Flower bud density is also
negatively related to the previous season’s
crop load. Non-thinned shoots, especially at
the basal nodes, had flower bud densities 25%
lower than hand-thinned trees (Byers et al.,
1990). Fruit set is also quite variable from year
to year depending on weather conditions.
Fruit set per 100 flowers was about 35 (Byers
and Lyons, 1985; Byers and Marini, 1994), 54
(Marini, 1985) and 60 (Byers et al., 1990) in
three different experiments. Due to differ-
ences in bloom density, some cultivars have
lower fruit densities than others.
Plant growth regulators
The bloom density inherent in a given culti-
var may also be modified with applications of
plant growth regulators. Late-summer and
autumn application of gibberellic acid (GA3)
killed flower buds (Stembridge and LaRue,
1969), but applications during early to mid-
summer inhibited flower bud initiation at
the lower nodes in peach (Edgerton, 1966;
Byers et al., 1990) and nectarine (Garcia-Pallas
et al., 2001). A commercial formulation of GA3,
 Crop Load Management 293

Release LC®, reduced flower bud formation by 50% reduced the need for hand thinning
and improved fruit size in commercial peach without reducing fruit size (Marini, 2002).
orchards (Southwick et al., 1998a,b). It was Pruning to remove more than 50% of the
registered for commercial use in California by length of a shoot resulted in reduced fruit
Abbott Laboratories, Inc. and later as Ralex® size. Retaining only one-fifth as many shoots
by Valent BioSciences Corp. for several years. as the number of fruits desired per tree, and
However, the registration was withdrawn thinning fruit so that five fruit were retained
due to inconsistent results and discouraging per shoot after hand thinning, reduced the
grower experiences. need for hand thinning, improved fruit size
Dormant-season application of vegetable- and increased profit more than normal prun-
based oils (e.g. soybean) has been used safely ing and hand thinning (Marini, 2003).
and effectively to pre-bloom thin peach flower
buds (Myers et al., 1996; Moran et al., 2000)
but has produced inconsistent results when Bloom
applied under commercial orchard conditions
(Reighard et al., 2003b; Andris et al., 2004). During bloom, peach producers have an oppor-
Flower buds on peach cultivars have been pre- tunity to remove blossoms by physical means
bloom thinned by 15 to 40% at rates of 6%, 7% or they can reduce fruit set with chemicals.
and 8% (v:v) of soybean oil–water in grower
orchards in South Carolina (Muriu Njoroge, Physical removal
2002; Reighard et al., 2003a). The 8% rate has
Many commercial peach producers reduce
been the most cost-effective for the majority of
early-season crop load by removing flowers
cultivars. However, differences in cultivar sen-
during bloom. When flower buds are in the
sitivity and annual environmental effects (i.e.
pink stage until shuck split, buds, flowers
winter temperatures) have lessened the effi-
and young fruit can be removed with fingers
cacy in some years, and thus commercial accep-
or by running a stiff brush along one side of
tance of this practice is limited at this time.
the shoot. The object is to remove about half of
Other chemicals have also been applied
the flowers. Baugher et al. (1988) demonstrated
during the dormant season to thin flower buds
that dragging curtains of large-diameter ropes
but sometimes led to over-thinning because
over the trees during bloom physically removed
they are dependent on concentration and tem-
enough blossoms to reduce hand-thinning
perature. These include ethephon (e.g. Eth-
costs and improved fruit size enough to be cost-
rel), which is absorbed by the buds and breaks
effective. Mechanical rope-thinning attachments
down into ethylene but can cause excessive
for tractors are available from commercial ven-
bud mortality if applied during unseasonably
dors and are routinely used by peach growers.
warm temperatures especially in autumn
However, trees need to be pruned carefully to
(Williams, 1989). This potential risk may be
a spreading open-vase shape to allow the
one of the reasons why it has not been labelled
ropes to fall freely through the tree to obtain
for peach in the USA. Hydrogen cyanamide
adequate thinning throughout the canopy.
(e.g. Dormex), which is labelled for dormant-
High-pressure water spray systems to
season application in some fruits to augment
remove blossoms were investigated by R. Byers
inadequate chilling, has also been examined
(1995, Virginia, personal communication) for
experimentally as a thinner. It can effectively
blossom thinning. This approach was never
thin flower buds, but the timing and rate are
adopted to a large extent, in part due to the
critical in peach or else over-thinning can
large volume of water required and some con-
occur (Fallahi et al., 1990; Powell, 1994).
cern about damage to scaffolds by the water.

Pruning Chemicals

Another method of reducing the number of Several strategies for modifying numbers of
flower buds per tree is by pruning off shoots flowers per tree at bloom have been investi-
with flower buds. Heading all fruiting shoots gated. Spraying trees during bloom with
294 R.P. Marini and G.L. Reighard
chemicals that kill flower parts and prevent
fertilization is relatively inexpensive, but
results are often inconsistent (Byers and Lyons,
1982, 1985; Byers et al., 2003). A number of
caustic chemicals, herbicides and surfactants
have been examined as bloom thinners. Some
growers used sodium dinitro-o-cresylate
(DNOC) but results were mixed. The material
was often applied when about 20% of the pis-
tils were showing and again when the final
20% of the flowers were open, in an attempt
to eliminate about 40% of the crop. Although,
with experience, some large growers produced
acceptable results with 1-aminomethanamide
dihydrogen tetraoxosulfate, the registration
on this material has never been approved on
peach and was removed on apple. Wilthin®, a
sulfcarbamide, is another caustic chemical
that is labelled for use as a thinner (Myers et al.,
1993), but it has performed inconsistently some
years (Greene et al., 2001) and requires close
attention to the handling and application pro-
cedures listed on the label. Research with
Wilthin® (R. Byers, Virginia, 1997, personal
communication) indicated that dilute appli-
cations should be made with an air-blast
sprayer (1000–2000 l/ha, depending on tree
size and spacing) at the rate of 7.5–15 l
Wilthin®/ha when about 90% of the flowers
have opened and about 10% of the buds are in
the pink stage of development. Proper timing
is critical for consistent results in the east, but
in California the timing may be less critical.
Ammonium thiosulfate (ATS), a liquid
fertilizer, has been widely researched as a
means of ‘burning’ or desiccating flower parts
to reduce flower numbers and fruit set (Johnson,
1998; Byers, 1999; Greene et al., 2001). How-
ever, there is no ATS label to thin peach flow-
ers and its somewhat inconsistent performance
probably accounts for corporate disinterest in
pursuit of a label. In spite of the lack of inter-
est by agrichemical companies, ATS research
as a bloom thinner continues in many tree
fruit regions in the USA because of its avail-
ability, low cost, user safety and general effec-
tiveness.
Other chemicals, especially surfactants,
have also been tested for thinning properties.
The surfactant ArmoThin® (N,N-bis-2-(w-
hydroxypolyoxyethylene/polyoxypropylene)
ethyl alkylamine) was tested in the 1990s as a
bloom thinner, but was never used commer-
cially (Southwick et al., 1998b). Another sur-
factant still under investigation is a dodecyl
ether of polyethylene glycol (i.e. Tetgitol™
TMN-6, also known as Surfactant WK or
DuPont WK). It has shown promise in thin-
ning flowers in orchard trials in the south-
eastern USA, but it has not been registered for
thinning peaches (Ebel et al., 1999; Wilkins
et al., 2004). It should be kept in mind that the
effectiveness of air-blast application of any
potential thinning agent is expected to vary
with the amount of water applied, air tem-
perature, humidity, surfactants, stage of flower/
fruit development, orchard training system
or other factors. Many of these products are
commercially available, but growers should
not use them for peach thinning unless they
are registered for peach thinning because
the results are inconsistent and trees may be
injured.
Post-bloom
Although thinning before or during bloom
has the greatest impact on fruit size, post-
bloom thinning allows one to first assess
fruit set and minimize the potential for over-
thinning.
Hand
Hand thinning has been the most common
method of reducing the crop load on peach
trees for many years. The general rule of thumb
has been to space fruit about 12, 15 and 20 cm
apart for large-, medium- and small-fruit cul-
tivars, respectively. In some regions distance
between fruit is not considered, rather fruit
number per shoot is varied depending on shoot
location in the canopy or season of harvest.
The number of fruit retained per shoot can be
two or three in the bottom of the tree, three or
four in the middle of the tree, and five in the
top of the tree. Fruit retained per shoot may
also vary from three for early-season culti-
vars, to four to six for late-season cultivars.
Spacing fruit to a given distance or to a given
number per shoot requires adequate pruning
to remove excess fruiting shoots. An alternative
approach may be to retain a certain number
 Crop Load Management
of fruit per hectare; the desired number would
vary with the cultivar and with the desired
fruit size at harvest. Based on data published
by Johnson and Handley (1989) in California,
to obtain fruit that are 6.3 cm in diameter trees
should be thinned to 70,000, 345,000 and 690,000
fruit/ha for small-, medium- and large-sized
cultivars. If 7.0 cm fruit are desired, then the
crop load should be reduced to about 70,000
and 140,000 fruit/ha for medium- and large-
fruited cultivars, respectively. These fruit num-
bers should probably be reduced by about
30% in the eastern USA, where tree canopies
are smaller and there is less available light
than in California. Small-fruited cultivars do
not have the potential to produce 7.0 cm diam-
eter fruit, even at very low crop loads. The
small fruit size associated with early-season
cultivars may be due to cell numbers or the
short duration of fruit growth.
The earlier trees are thinned, the greater
the effect will be on fruit size. Havis (1962)
demonstrated that reducing the crop load
before or during bloom maximizes fruit size,
and hand thinning becomes less effective for
each day after bloom that thinning is delayed.
In general, early-season cultivars are small-
fruited and require earlier and more severe
thinning than late-season cultivars. This is
why so much emphasis has been placed on
methods to reduce flower density at bloom. It
is difficult to find orchard workers who do a
good job of hand thinning. Hand thinning
requires a large labour force with good super-
vision and can cost from $375 to $1875/ha at
$9/h depending on tree size and fruit set (G.
Reighard, 2006, South Carolina, personal
communication). Estimates for California are
about $1514 (Hasey et al., 2004) to $2423 per
hectare (Day et al., 2004). Therefore research-
ers and growers have been looking for ways
to reduce thinning costs. Rather than remov-
ing small fruit solely by hand, some growers
use rubber hoses, plastic baseball bats or pad-
ded sticks to knock fruit off the tree, followed
by final hand thinning. This method is less
expensive than hand thinning alone, but it
does not facilitate the selective removal of
insect-injured and small fruit. Fruit removal
with bats or sticks is most efficient if delayed
until fruits are large enough to be shaken off
when the limb is struck.
295
The effect of thinning on fruit size is
related to the time of thinning. Generally the
earlier the thinning is accomplished, the
greater the effect on final fruit size. However,
Dorsey and McMunn (1935) demonstrated that
even thinning later in the season is beneficial.
They thinned trees at various times during
the season starting 64 days after bloom when
the pits were hardening. They found that thin-
ning up to 4 weeks before harvest increased
the percentage and the yield of large fruit.
During a drought, commercial peach produc-
ers without irrigation sometimes remove a
portion of the crop during the final swell to
improve the size of the remaining fruit.
The typical instructions for thinning
peaches are to space the fruit relatively evenly
along the fruiting shoot. However, the dis-
tance between fruit and the distribution of
fruit along the shoot do not affect fruit size
(Corelli Grappadelli and Coston, 1991; Marini
and Sowers, 1994; Ben Mimoun and DeJong,
1999). Marini and Sowers (1994) reported that
fruit on long shoots tended to be larger than
fruit on short shoots and fruit on shoots with
lateral shoots tended to be larger than fruit on
shoots with only a terminal shoot. Heavily
cropped branches on lightly cropped trees
also had larger fruit than heavily cropped
branches on heavily cropped trees. Therefore,
the number of fruit per tree is more important
than the distribution of fruit on the tree,
because fruit size depends on photosynthate
from leaves in the immediate vicinity of the
fruit as well as photosynthate from more dis-
tant parts of the tree.
Mechanical
Some large peach growers use mechanical tree
shakers or branch shakers to remove excess
fruit, but smaller growers cannot justify the
cost of a shaker. For this method to be suc-
cessful, trees must be properly trained. About
60 cm of space between the ground and the
lowest scaffold branches is required to attach
the clamp to the trunk. Trees with no more
than four scaffold branches and without wil-
lowy branches or many sub-scaffolds are most
conducive to shaking. Shakers can injure the
bark and predispose the tissue to Cytospora
canker infection. Hand thinning is usually
296 R.P. Marini and G.L. Reighard
required for touch-up. Attempts to do all of
the thinning with a shaker may result in over-
thinning, and there is a tendency to remove
the largest fruit. A good shaker operator, work-
ing with well-trained trees, can remove 50–80%
of the fruit, while the rest must be removed by
hand. As with baseball batting, small fruit are
difficult to remove by shaking and so shakers
must be used towards the later stages of thin-
ning, which may be too late for early-season or
small-fruited cultivars (Kamas et al., 1998).
Chemicals
Successful post-bloom apple thinning with
chemicals led to a search to identify similar
materials for peaches. Since the 1940s research-
ers have tested many apple thinners on peach,
including naphthalene acetic acid (NAA),
naphthylacetamide, carbaryl, and others. Early
attempts to thin peach trees with NAA at
bloom or shortly after bloom were somewhat
successful, and Murneek and Hibbard (1947)
showed that NAA was most effective when
applied about 30 days after full bloom. Kelly
(1955) applied NAA at concentrations of 30 to
50 ppm and found that NAA at 30 or 40 ppm
thinned peaches when applied at about 14
days after shuck off in two of three experi-
ments. Edgerton and Hoffman (1952) reported
that NAA applied a month after bloom caused
more thinning than DNOC at bloom, but the
concentration of NAA that provided adequate
thinning varied from 20 to 40 ppm depending
on the cultivar. Research results with NAA
were so inconsistent that interest shifted to
the carbamate compounds. Horsfall and Moore
(1956) evaluated eight formulations of carbam-
ates as post-bloom peach thinners, and chloro
IPC showed the most promise. Although the
level of thinning was sometimes acceptable,
results varied with season and cultivar, and
some shoots were not thinned well, causing
fruit clustering. None of these materials proved
to be consistently effective post-bloom thin-
ners for commercial use.
During the 1970s and 1980s ethylene-re-
leasing compounds were tested. Ethephon
caused fruit thinning, but also often caused
partial defoliation and tree injury. Another
ethylene- releasing agent, CGA-15281 (2-chlor-
ethylmethyl-bis(phenylmethoxy)silane),
provided fairly consistent fruit thinning with
little leaf drop in the south-eastern USA (Gam-
brell and Sims, 1983), but in the mid-Atlantic
states defoliation was often a problem. The
product was never developed because the
company responsible went out of business.
Research on apples, and to a lesser extent
on peach, indicated that natural or chemically
induced fruit abscission might be accentuated
by limiting photosynthesis. Therefore, Byers
et al. (1984) shaded peach trees and also
sprayed trees with photosynthesis-inhibiting
herbicides, such as terbacil. Covering the trees
with 92% shade cloth from 31 to 41 days after
bloom, but not earlier, reduced fruit set
severely and improved fruit size slightly. Ter-
bacil applied at the rate of 500 ppm and at 35
days after bloom caused over-thinning with
no defoliation. Although this approach was
quite innovative and showed great promise,
no chemical company was willing to register
a herbicide for peach thinning.
Pruning
Harmon (1933) was the first to demonstrate
that pruning could influence fruit size and
quality. In a 4-year experiment he compared
non-pruned trees with trees that were pruned,
and then the number of shoots per tree was
varied by cutting back the shoots by varying
amounts. As pruning became more severe,
there were fewer fruits set per tree and fruit
size increased, but yield per tree was highest
for moderately pruned trees. The author indi-
cated that from an economic standpoint the
least expensive thinning of fruit is done by
pruning. Hand thinning is a necessary accom-
paniment to adjust the number of fruits to the
leaf area of the tree. Marini (2002, 2003) fol-
lowed up on this pruning work. Heading all
fruiting shoots on a tree by 50% while dormant,
pruning reduced fruit set and hand-thinning
costs, and sometimes improved fruit size,
compared with not heading shoots. Heading
to remove more than 50% of each shoot fur-
ther reduced fruit set, but negatively affected
fruit size. Pruning to retain varying numbers
of shoots per tree also had an impact on fruit
size. As the number of shoots retained per
tree declined, fruit set and the need for hand
thinning was reduced and fruit size increased.
 Crop Load Management
When only about 100 shoots were retained per
tree, the combination of reduced thinning
costs and improved fruit size resulted in a net
profit $6000/ha greater than for trees with
250 shoots. This practice is commonly followed
in California (K. Day, California, 2000, personal
communication), but the concept is new for
growers in the eastern USA. Following more
research to identify the appropriate crop loads
for different cultivars, it should be possible to
prune trees to a given number of shoots per
hectare and then thin the trees to a given
number of fruit per shoot to obtain the desired
number of fruit per hectare. The V-shaped
tree would be particularly well suited to this
approach because the simple tree structure
facilitates pruning to a specific number of
shoots per tree (DeJong et al., 1994).
12.5 Crop Load Interactions
with Stresses
A number of tree stresses can have a negative
impact on fruit size. If these stresses can be
predicted early in the season, crop load can be
reduced to minimize the effects of stress.
Tree density
To take advantage of high production during
the early years after orchard establishment,
some peach growers have started to increase
the number of trees per hectare. Standard tree
densities are about 300 to 350 trees/ha, but
newer plantings may have as many as 1300
trees/ha. Giulivo et al. (1984) planted peach
and nectarine trees at densities ranging from
1250 to 2000 trees/ha. Normally fruit size is
negatively related to fruit density (number of
fruit/cm2 trunk cross-sectional area). How-
ever, they found that both fruit size and fruit
density were negatively related to tree den-
sity. The reduction in fruit size was related to
a reduction in fruit relative growth rate dur-
ing stage I of fruit development. In a study of
peach orchard systems, Grossman and DeJong
(1998) reported that the dry weight of indi-
vidual fruits was inversely related to tree
density, but it may have been a function of
297
yield per hectare. Their data indicate that tree
form as well as tree density may influence
fruit size. In another study, fruit size and the
percentage of marketable yield were inversely
related to tree density (Marini and Sowers,
2000). Even when adjusted for number of
fruits per hectare, fruit weight was higher for
low-density trees than for high-density trees.
The reason for reduced fruit weight may be
increased competition for water and nutrients
between trees or increased shading in close
plantings. Whatever the reason, these data
indicate that high-density peach plantings
should be dormant-pruned or thinned more
aggressively than low-density plantings.
Water stress
Both crop load and water availability affect
peach fruit size, but there is little information
available concerning the combined effects of
water stress and crop load. Both Morris et al.
(1962) and Girona et al. (2002) showed that the
effects of crop load and irrigation on fruit size
were additive, rather than interactive; how-
ever, at heavy crop loads only a small per-
centage of harvested fruit from non-irrigated
trees were of marketable size. Berman and
DeJong (1996) performed an experiment with
widely differing crop loads to study the
effects of water stress on fruit growth. Fresh
weight, which is analogous to fruit size, for
all crop loads was reduced by water stress to
a similar extent. However, the reduction in
fruit dry weight by water stress was greatest
for high crop loads. The reduction in dry
weight on heavy-cropped trees was probably
due to resource-limited conditions resulting
from the combination of a high carbon demand
and water-reduced photosynthesis. These
data indicate that irrigated orchards can sup-
port heavier crop loads than non-irrigated
orchards.
Leaf-feeding arthropods
Arthropods that feed on peach foliage would
theoretically reduce photosynthesis and fruit
size and quality. In apple, the negative effect
298 R.P. Marini and G.L. Reighard
of indirect pests increased as population levels
increased (Marini et al., 1994). For non-fruiting
sour cherry trees, Layne and Flore (1992)
showed that trees could compensate for up to
20% defoliation with minor effects on carbon
assimilation. However, much less informa-
tion is available for peach. Seasonal accumu-
lations of 8900 cumulative European red
mite-days did not influence fruit size or qual-
ity of ‘Cresthaven’ peaches (McClernan and
Marini, 1986), but more information is needed
on the effects of other indirect pests, such as
Japanese beetle and two-spotted spider mite,
and of diseases such as bacterial leaf spot, on
peach fruit size and quality.
High temperatures
Unusually high temperatures during applica-
tions of soybean oil in January (>28°C) in
South Carolina gave significantly higher
flower bud thinning and increased fruit size
at harvest (Lennon, 2004). In addition, tem-
peratures from bloom to 30 days after bloom
affect fruit development rates and can be used
to predict harvest maturity date (Ben Mimoun
and DeJong, 1999). In California during 2004,
abnormally high temperatures at and after
bloom advanced fruit maturity by 10–14 days.
However, fruit size was decreased by one to
two sizes below normal. The high tempera-
tures advanced fruit development much
quicker than normal so the trees’ carbohy-
drate reserves had probably already become
limiting before growers initiated thinning
(DeJong, 2005). This dilution of reserves to
the rapidly developing fruit crop reduced
potential fruit growth early in stage I, and the
fruit never recovered by harvest time. High
temperatures (>38°C) during flower bud ini-
tiation after harvest in late summer can also
negatively affect next year’s flower buds.
Postharvest reduction of irrigation in August
was shown by Handley and Johnson (2000)
not only to increase water stress and leaf tem-
perature, but also increased fruit doubles and
deep sutures the following year. In South Car-
olina, mean 24 h temperatures during August
1999 averaged 27.7°C, or 2°C above normal,
in the Ridge peach region, which was one of
the warmest Augusts recorded at the South-
east Regional Climate Center, Columbia,
South Carolina since 1900. In late July and into
August, growers irrigated primarily only late-
season cultivars, which were still unpicked
even though August rainfall was 5.3 cm below
normal and trees were showing heat stress. In
the subsequent growing season (i.e. 2000), an
abnormally high incidence of fruit doubling
occurred in some of the non-irrigated, early-
and mid-season cultivars (C.R. Carr III and
L.F. Holmes III, South Carolina, 2000, personal
communication).
Low temperatures
Thinning during bloom may increase shoot
growth and flower bud density for next year’s
crop and also increase the cold tolerance of
buds (Byers and Marini, 1994). Thus, thinning
during bloom may be more important for
northern regions that are prone to low-
temperature injury during the winter and for
cultivars that naturally produce low flower
bud densities.
12.6 Future Directions
During the 20th century the need for peach
crop load management was clearly demon-
strated and fruit thinning became a standard
commercial practice. Researchers have des-
cribed in detail the physiology of fruit growth
and development, and this basic information
was used to develop fruit thinning pro-
grammes. Future market demands will ulti-
mately influence orchard practices such as
crop load management. Assuming the
demand for large fruit continues, fruit size
should be an important criterion for peach
breeders. Small and medium-sized cultivars
are rapidly becoming non-profitable for some
markets. The demand for large fruit will
increase the need for early and aggressive
fruit thinning. Growers will continue to
search for fruit thinning methods that are less
expensive than hand thinning. More research
is needed to develop chemicals that can be
applied pre- and post-bloom to reduce final
 Crop Load Management 299

fruit set. This will require a better understand-
ing of how the efficacy of these materials
interacts with climatic factors, the condition
of the tree and the physiological development
of the fruit. We also need a better understand-
ing of the relationship between fruit size and
number of fruit per unit of land area or unit of
canopy volume for different cultivars, differ-
ent orchard systems, and irrigated and non-
irrigated orchards. After obtaining these types
of information, researchers and growers can
develop fruit thinning strategies to obtain the
most profitable crop load for specific situa-
tions and for different markets.

References

Andris, H.L., Johnson, R.S., Klassen, K. and Day, K. (2004) Stone fruit thinning with soybean oil. 2004 California
Tree Fruit Agreement Annual Research Report. California Tree Fruit Agreement, Reedley, California, pp.
47–51.
Baugher, T.A., Elliot, K.C., Blizzard, S.H., Walter, S.I. and Keiser, T.A. (1988). Mechanical bloom thinning of
peach. HortScience 23, 981–983.
Beach, S.A. (1903) Correlation between different parts of the plant in form, color, size, and other characteris-
tics. In: Proceedings of the International Conference on Plant Breeding and Hybridization Memoirs, Vol.
1. Horticultural Society of New York, New York, p. 63.
Ben Mimoun, M. and DeJong, T.M. (1999) Using the relation between growing degree hours and harvest date
to estimate run-times for PEACH: a tree growth and yield simulation model. Acta Horticulturae 499,
107–114.
Berman, M.E. and DeJong, T.M. (1996) Water stress and crop load effects on fruit fresh and dry weights in
peach (Prunus persica). Tree Physiology 16, 859–864.
Blake, M.A. (1943) Classification of fruit bud development on peaches and nectarines and its significance in
cultural practice. New Jersey Agricultural Experiment Station Bulletin No. 706.
Byers, R.E. (1999) Effects of bloom-thinning chemicals on peach fruit set. Journal of Tree Fruit Production 2,
59–78.
Byers, R.E. and Lyons, C.G. Jr (1982) Flower bud removal with surfactants for peach thinning. HortScience 17,
377–378.
Byers, R.E. and Lyons, C.G. Jr (1985) Peach flower thinning and possible sites of action of desiccating chemi-
cals. Journal of the American Society for Horticultural Science 110, 662–667.
Byers, R.E. and Marini, R.P. (1994) Influence of blossom and fruit thinning on peach flower bud tolerance to
an early spring freeze. HortScience 29, 146–148.
Byers, R.E., Lyons, C.G. Jr, Del Valle, T.B., Barden, J.A. and Young, R.W. (1984) Peach fruit abscission by shading
and photosynthetic inhibition. HortScience 19, 649–651.
Byers, R.E., Carbaugh, D.H. and Presley, C.N. (1990) The influence of bloom thinning and GA3 sprays on
flower bud numbers and distribution in peach trees. Journal of Horticultural Science 65, 143–150.
Byers, R.E., Costa, G. and Vizzotto, G. (2003) Flower and fruit thinning of peach and other Prunus. In: Janick,
J. (ed.) Horticultural Reviews, Vol. 28. Wiley, New York, pp. 351–392.
Cain, J.C. and Mehlenbacher, J.R. (1956) Effects of nitrogen and pruning on trunk growth in peaches. Proceed-
ings of the American Society for Horticultural Science 67, 139–143.
Chalmers, D.J. and van den Ende, B. (1975) A reappraisal of the growth and development of peach fruit.
Australian Journal of Plant Physiology 2, 623–634.
Cole, S.W. (1849) The American Fruit Book. John P. Jewett, New York.
Corelli Grappadelli, L. and Coston, D.C. (1991) Thinning pattern and light environment in peach tree canopies
influence fruit quality. HortScience 26, 1464–1466.
Day, K.R., Andris, H.L., Klonsky, K.M. and DeMoura, R.L. (2004) Sample costs to establish and produce
peaches. http://coststudies.ucdavis.edu/files/peachesjv2004.pdf (accessed July 2005).
DeJong, T.M. (2005) Using physiological concepts to understand early spring temperature effects on fruit
growth and anticipating fruit size problems at harvest. Summerfruit Autumn, 10–13.
DeJong, T.M., Day, K.R., Doyle, J.F. and Johnson, R.S. (1994) The Kearney Agricultural Center perpendicular
‘V’ (KAC-V) orchard system for peaches and nectarines. HortTechnology 4, 362–367.
Dorsey, M.J. and McMunn, R.L. (1935) Peach thinning investigations V: a study of late thinning. Proceedings
of the American Society for Horticultural Science 33, 280–283.
300 R.P. Marini and G.L. Reighard

Ebel, R.C., Caylor, A., Pitts, J. and Himelrick, D.G. (1999) ‘Surfactant WK’ for thinning peach blossoms. Fruit
Varieties Journal 53, 184–188.
Edgerton, L.J. (1966) Some effects of gibberellin and growth retardants on bud development and cold hardi-
ness of peach. Proceedings of the American Society for Horticultural Science 88, 197–203.
Edgerton, L.J. and Hoffman, M.B. (1952) The effect of thinning peaches with bloom and post-bloom sprays on
the cold hardiness of the fruit buds. Proceedings of the American Society for Horticultural Science 60,
155–159.
Fallahi, E., Kilby, M. and Moon, J.W. (1990) Effects of various chemicals on dormancy, maturity and thinning
of peaches. In: Deciduous Fruit and Nut. University of Arizona, Series P-83. University of Arizona,
Tucson, Arizona, pp. 121–128.
Fitz, J. (1872) The Southern Apple and Peach Culturist. J.W. Randolf and English, Richmond, Virginia.
Gage, J. and Stutte, G. (1991) Developmental indices of peach: an anatomical framework. HortScience 26,
459–463.
Gambrell, C.E. and Sims, E.T. (1983) Results of eight years with CGA-15281 as a postbloom thinner for
peaches. Journal of the American Society for Horticultural Science 108, 605–608.
Garcia-Pallas, I., Val, J. and Blanco, A. (2001) The inhibition of flower bud differentiation in ‘Crimson gold’
nectarine with GA3 as an alternative to hand thinning. Scientia Horticulturae 90, 265–278.
Girona, J., Marsal, J., Mata, M., Arbones, A. and Mata, A. (2002) The combined effect of fruit load and water
stress in different peach fruit growth stages (Prunus persica L). Acta Horticulturae 584, 149–152.
Giulivo, C., Ramina, A. and Costa, G. (1984) Effects of planting density on peach and nectarine productivity.
Journal of the American Society for Horticultural Science 109, 287–290.
Gould, H.P. (1918) Peach Growing. The Macmillan Company, New York.
Greene, D.W., Hauschild, K.I. and Krupa, J. (2001) Effect of blossom thinners on fruit set and fruit size of
peaches. HortTechnology 11, 179–183.
Grossman, Y.L. and DeJong, T.M. (1995a) Maximum fruit growth potential and seasonal patterns of resource
dynamics during peach growth. Annals of Botany 75, 553–560.
Grossman, Y.L. and DeJong, T.M. (1995b) Maximum fruit growth potential following resource limitation
during peach growth. Annals of Botany 75, 561–567.
Grossman, Y.L. and DeJong, T.M. (1998) Training and pruning systems effects on vegetative growth potential,
light interception, and cropping efficiency in peach trees. Journal of the American Society for Horticul-
tural Science 123, 1058–1064.
Handley, D.F. and Johnson, R.S. (2000) Late summer irrigation of water-stressed peach trees reduces fruit
doubles and deep sutures. HortScience 35, 771.
Harmon, F.N. (1933) Relation of pruning and thinning to fruit size and yield of Paloro peaches. Proceedings
of the American Society for Horticultural Science 30, 219–222.
Hasey, J., Duncan, R., Norton, M., Konsky, K.M. and Livingston, P. (2004) Sample costs to produce cling
peaches: Sacramento and San Joaquin valleys, extra-early harvested varieties. http://coststudies.ucdavis.
edu/files/peachessacsjv2004.pdf (accessed July 2005).
Havis, A.L. (1962) Effects of time of fruit thinning of Redhaven peach. Proceedings of the American Society
for Horticultural Science 80, 172–176.
Horsfall, F. and Moore, R.C. (1956) Isopropyl N-(3 chlorophenyl)-carbamates and other carbamates as fruit
thinning sprays for Halehaven, Ambergem, and Elberta peaches. Proceedings of the American Society
for Horticultural Science 68, 63–69.
Johnson, R.S. (1998) ATS works well as bloom thinner on stone fruits. Good Fruit Grower 49, 14–15.
Johnson, R.S. and Handley D.F. (1989) Thinning response of early, mid-, and late-season peaches. Journal of
the American Society for Horticultural Science 114, 852–855.
Jones, I.D. (1932) Further observations on influence of leaf area on fruit growth and quality in the peach.
Proceedings of the American Society for Horticultural Science 29, 34–38.
Kamas, J., McEachem, G., Stein, L. and Roe, N. (1998) Peach Growing in Texas. http://aggie-horticulture.
tamu.edu/extension/peach/peach.html (accessed July 2005).
Kelly, V.W. (1955) Time of application of naphthaleneacetic acid for fruit thinning of the peach in relation to
the June drop. Proceedings of the American Society for Horticultural Science 66, 70–72.
Layne, D.R. and Flore, J.A. (1992) Photosynthetic compensation to partial leaf area reduction in sour cherry.
Journal of the American Society for Horticultural Science 117, 279–286.
Lennon, S.F. (2004) Integrated pest management strategies for reducing pesticide-related risks and increas-
ing production efficiency in South Carolina peaches. MSc thesis, Clemson University, Clemson, South
Carolina.
 Crop Load Management 301

McClernan, W.A. and Marini, R.P. (1986) European red mite on yield, fruit quality, and growth of peach trees.
HortScience 21, 244–246.
Marini, R.P. (1985) Vegetative growth, yield, and fruit quality of peach as influenced by dormant pruning,
summer pruning, and summer topping. Journal of the American Society for Horticultural Science 110,
133–139.
Marini, R.P. (2002) Heading fruiting shoots before bloom is equally effective as blossom removal in peach
crop load management. HortScience 37, 642–646.
Marini, R.P. (2003) Peach fruit weight, yield, and crop value are affected by number of fruiting shoots per tree.
HortScience 38, 512–514.
Marini, R.P. and Sowers, D.L. (1994) Peach fruit weight is influenced by crop density and fruiting shoot length
but not position on the shoot. Journal of the American Society for Horticultural Science 119, 180–184.
Marini, R.P. and Sowers, D.L. (2000) Peach tree growth, yield, and profitability as influenced by tree form and
tree density. HortScience 35, 837–842.
Marini, R.P., Pfeiffer, D.G. and Sowers, D.S. (1994) Influence of European red mite (Acari: Tetranychidae) and
crop density on fruit size and quality and on crop value of ‘Delicious’ apples. Journal of Economic En-
tomology 87, 1302–1311.
Moran, R.E., Deyton, D.E., Sams, C.E. and Cummins, J.C. (2000) Applying soybean oil to dormant peach trees
thins flower buds. HortScience 35, 615–619.
Morris, J.R., Kattan, A.A. and Arrington, E.H. (1962) Response of Elberta peaches to the interactive effects of
irrigation, pruning, and thinning. Proceedings of the American Society for Horticultural Science 80,
177–189.
Muriu Njoroge, S. (2002) Crop load manipulation in peach: comparison of soybean oil, ammonium thiosulfate
and hand-thinning strategies. MSc. thesis, Clemson University, Clemson, South Carolina.
Murneek, A.E. and Hibbard, A.D. (1947) Investigations on thinning of peaches by means of caustic and
hormone sprays. Proceedings of the American Society for Horticultural Science 50, 206.
Myers, R.E., Deyton, D.E. and Sams, C.E. (1996) Applying soybean oil to dormant peach trees alters internal
atmosphere, reduces respiration, delays boom, and thins flower buds. Journal of the American Society
for Horticultural Science 121, 96–100.
Myers, S.C., King, A. and Savelle, A.T. (1993) Bloom thinning of ‘Winblo’ peach and ‘Fantasia’ nectarine with
monocarbamide dihydrogensulfate. HortScience 28, 616–617.
Overholser, E.L. and Claypool, L.L. (1931) The relation of leaf area per peach to physical properties and
chemical composition. Proceedings of the American Society for Horticultural Science 28, 5–17.
Powell, A.A. (1994) Action Program for Dormex Application on Peaches. Alabama Cooperative Extension
Service Agriculture & Natural Resources Timely Information FN-94-1. Auburn University, Auburn, Alabama.
Reighard, G.L., Ouellette, D.R. and Brock, K.H. (2003a) Economic analysis of soybean oil application to delay
peach bloom and pre-bloom thin peach flowers. In: Reighard, G. (ed.) Annual Peach Research Report
Vol. III. South Carolina Peach Council, Columbia, South Carolina, pp. 104–113.
Reighard, G.L., Njoroge, S.M., Lennon, S., Ouellette, D. and Brock, K. (2003b) Reducing peach (Prunus persica)
flower bud numbers with a mid-winter application of soybean oil. In: Proceedings of the 30th Annual
Conference of the Plant Growth Regulation Society of America. The Plant Growth Regulation Society of
America, LaGrange, Georgia, p. 28.
Scorza, R.L., May, G., Purnell, B. and Upchurch, B. (1991) Differences in number and area of mesocarp cells
between small- and large-fruited peach cultivars. Journal of the American Society for Horticultural
Science 116, 861–864.
Shoemaker, J.S. (1933) Certain advantages of early thinning of Elberta. Proceedings of the American Society
for Horticultural Science 30, 223–224.
Southwick, S.M., Weis, K.G., Yeager, J.T., Rupert, M.E. and Hasey, J.K. (1998a) Chemical thinning of cling
peach. Acta Horticulturae 465, 647–654
Southwick, S.M., Weis, K.G., Yeagor, J.T., Hasey, J.K. and Rupert, M.E. (1998b) Bloom thinning of ‘Loadel’
cling peach with a surfactant: effects of concentration, carrier volume, and differential applications
within the canopy. HortTechnology 8, 55–58.
Stembridge, G.E. and LaRue, J.H. (1969) The effect of potassium gibberellate on flower bud development in
the ‘Redskin’ peach. Journal of the American Society for Horticultural Science 94, 492–495.
Tukey, H.B. (1933) Growth of peach embryo in relation to growth of fruit and season of ripening. Proceedings
of the American Society for Horticultural Science 30, 209–218.
Weinberger, J.H. (1931) The relation of leaf area to size and quality of peaches. Proceedings of the American
Society for Horticultural Science 28, 18–22.
302 R.P. Marini and G.L. Reighard

Weinberger, J.H. and Cullinan, F.P. (1932) Further studies on the relation between leaf area and size of fruit,
chemical composition, and fruit bud formation in Elberta peaches. Proceedings of the American Society
for Horticultural Science 29, 23–27.
Werner, D.J., Mowrey, B.D. and Chaparro, J.X. (1988) Variability in flower bud number among peach and
nectarine cultivars. HortScience 23, 578–580.
Westwood, M.N. and Batjer, L.P. (1958) Size of Elberta and J.H. Hale peaches during the thinning period as
related to size at harvest. Proceedings of the American Society for Horticultural Science 72, 102–105.
Wilkins, B.S., Ebel, R.C., Dozier, W.A., Pitts, J. and Boozer, R. (2004) Tergitol TMN-6 for thinning peach blos-
soms. HortScience 39, 1611–1613.
Williams, K.M. (1989) Peach bloom delay using fall applications of Ethrel and Pro-Gibb. Acta Horticulturae
254, 151–154.
Zucconi, F. (1986) Peach. In: Monselise, S.P. (ed.) Handbook of Fruit Set and Development. CRC Press, Boca
Raton, Florida, pp. 303–321.
 13 Nutrient and Water Requirements
of Peach Trees

R.S. Johnson
Department of Plant Sciences, University of California, Davis,
California, USA

13.1 Introduction 303

13.2 Nutrient Requirements 304
Diagnostic methods 304
Nitrogen 305
Phosphorus 308
Potassium 308
Calcium 310
Magnesium 311
Sulfur 311
Zinc 312
Boron 314
Iron 316
Manganese 317
Copper 318
13.3 Peach Water Requirements 318
Water stress 321

13.1 Introduction of water. Thus, information has been devel-

When grown on fertile soils well supplied
with nutrients and provided with adequate
amounts of water, peach trees are very vigor-
ous and productive. Many locations around
the world where peaches are grown enjoy
these conditions. However, there are also situ-
ations of peach orchards growing in more
challenging soils (infertile, high pH, low in
micronutrients, etc.) or with insufficient supplies
© CAB International 2008. The Peach: Botany, Production and Uses
(eds D.R. Layne and D. Bassi)
oped to help orchard managers deal with these
suboptimal conditions. This chapter deals with
the management of nutrients and water under
both optimal and suboptimal conditions. As
there has been increasing concern in recent
years over environmental protection, the chal-
lenge is to supply adequate amounts of nutri-
ents and water for good production, but guard
against excessive amounts that might lead to
environmental contamination.
303
304 R.S. Johnson

13.2 Nutrient Requirements can be used as a diagnostic tool, but often

deficiency symptoms do not develop until
As with all other plants, peach trees require at serious damage to the tree or crop has already
least 16 essential elements for optimum occurred. Also, loss in productivity can occur
growth and productivity. A deficiency of any without any leaf symptoms for some nutri-
one of these nutrients can lead to problems ents. Therefore, a more predictive method
with production, fruit quality, vegetative would be desirable.
growth or tree health. Some deficiencies are The standard method that has been
extremely rare in field-grown peach trees and developed for peaches, as well as most other
information is limited. However, in general, plants, is a mid-summer sample of mature,
the deficiency threshold for each nutrient has mid-shoot leaves. Batjer and Westwood (1958)
been determined (see Table 13.1 below), as sampled ‘Elberta’ peach leaves throughout
well as a standard method of sampling. In the whole growing season and found relatively
recent years, research has focused on improved constant values of most nutrients during the
sampling methodologies in an effort to better period from 100 to 125 days after bloom
diagnose and predict nutritional problems. (Fig. 13.1/Plate 77). They recommended this
Also, there has been much research effort as a stable period for leaf sampling, when con-
directed towards improved nutrient uptake sistent values could be obtained from year to
efficiency so that environmental contamina- year. Researchers from many different peach-
tion can be minimized and orchard profitabil- growing areas around the world followed the
ity maximized. This chapter presents the same general protocol and came up with
standard information developed over many remarkably similar values for most nutrients
years as well as recent research efforts. in healthy orchards (Leece et al., 1971). Thus,
this has become the standard procedure for
peach orchards and will be referred to for
each nutrient throughout this chapter. How-
Diagnostic methods
ever, the method does have drawbacks and
considerable efforts have been made in recent
There is a need to sample nutrient levels in years to develop alternative approaches.
the tree so corrective measures can be imple- The biggest drawback is one of timing:
mented in a timely manner. Leaf symptoms 100 to 125 days after bloom is generally too
late in the season to apply any corrective mea-
sures for yield and fruit quality in that year.
Table 13.1. Peach deficiency thresholds and An early-season or even dormant sampling
sufficiency ranges that have been set by various would be preferable. Flower sampling for Fe
researchers. Values for mid-shoot leaves collected content showed promise as a predictor of iron
in mid-summer and expressed on a dry weight chlorosis, although there was variability
basis.
among sites (Sanz et al., 1997a) and the abso-
Deficiency Sufficiency
lute values changed considerably from year
Nutrient threshold range to year (Sanz et al., 1997b). Deficiency thresh-
olds have been proposed for most nutrients
N (%) 2.2–2.4 2.6–3.5 but have not been validated in commercial
P (%) 0.09–0.12 0.14–0.40 orchards (Sanz et al., 1995). Sanz and Monta-
K (%) 0.75–1.0 2.0–3.0 ñes (1995) correlated flower nutrient levels
Ca (%) 1.0 1.5–3.0 with leaf nutrient levels at both 60 and 120
Mg (%) 0.10–0.30 0.30–0.80 days after bloom and reported significant cor-
S (%) 0.09 0.14–0.40 relations with most nutrients. Similar results
Zn (ppm) 10–20 20–50 were also obtained by correlating the two
B (ppm) 15–30 30–70
dates with each other (Montañes and Sanz,
Fe (ppm) – 80–250
1994). They also found that leaf nutrients at
Mn (ppm) 20 40–200
Cu (ppm) 3 5–16
60 days after bloom tended to correlate better
with yield than later dates (Sanz et al., 1992b).
 Nutrient and Water Requirements
N, P, Zn
Mn, Mg, B, CI
Ca
Apr May June July Aug Sept
All these relationships still need to be more
widely tested before they can be used reliably.
There is also the possibility of using dormant
shoots or roots as an early predictor of tree
nutrient status (Johnson et al., 2006).
Researchers have also been evaluating
more sophisticated methods of analysing
nutrient data. Both the diagnosis and recom-
mendation integrated system (DRIS) and the
deviation from optimum percentage (DOP)
method have been applied to peach trees with
some success. The DRIS approach uses nutri-
ent ratios and products to come up with a
series of indices that theoretically stay con-
stant over time (Walworth and Sumner, 1987).
The DOP approach simply quantifies the
deviation of a given sample from a theoretical
optimum (Montañes et al., 1993). Generally,
the DOP method has been found to be more
useful because its results are unambiguous
and it provides some guidelines for how
much fertilizer may be needed to correct a
deficiency (Monge et al., 1995; Sanz, 1999).
Before any new protocol or approach can be
widely adopted, it will be important to relate
it to the various components of yield and fruit
quality so there is a physiological basis for its
implementation. In summary, good progress
has been made with various approaches to
sampling nutrients in peach trees. However,
none has yet shown consistent enough results
to replace the standard method of sampling
mature leaves in mid-summer.
305
Fig. 13.1. Generalized shapes
of concentration curves of
mineral nutrients in the leaf
during growing season. Curves
Oct show trends, not actual levels,
for the northern hemisphere.
Nitrogen
N is a critical element for plant life since it is
found in many important compounds includ-
ing amino acids, proteins, enzymes, nucleic
acids (component of DNA) and chlorophyll.
It is the one nutrient that needs to be applied
universally to peach orchards throughout the
world. Although annual applications may not
always be necessary, continued productivity
can only be maintained with regular applica-
tions of N fertilizers. Peach trees respond very
dramatically to N. With heavy fertilization,
and under favourable environmental condi-
tions, extreme vegetative growth and a very
dense canopy will result (much more than for
most other fruit trees). Therefore, care must
be taken to prevent both over- and under-
fertilization.
In recent years, many growers have cut
back substantially on the amount of N applied
to their peach orchards. At one time, recom-
mended rates of 200 kg N/ha or more were
common (Daane et al., 1995; Tagliavini et al.,
1996). Current recommendations are often
around 100 kg N/ha or even less (Tagliavini
and Marangoni, 2002). There are at least three
explanations for this change in practice. First,
growers have become more environmentally
aware and are striving to minimize nitrate
pollution in groundwater and runoff. Second,
the subtle effects of over-fertilization, which
have generally been recognized in the past,
306 R.S. Johnson
have only recently been clearly documented
(Weinbaum et al., 1992; Daane et al., 1995).
Finally, improvements in N application effi-
ciency have recently been demonstrated and
are being adopted by many growers.
N deficiency is identified by vegetative
growth with light green or yellow leaves. A
characteristic red coloration generally devel-
ops on shoots and leaf blades (Fig. 13.2/Plate
78). Often this discoloration takes on the form
of red or brown spots on the leaves. Fruit pro-
duced under low N are smaller but have
increased red colour compared with fruit pro-
duced with high N (Fig. 13.3/Plate 79) (Ball-
inger et al., 1966; Johnson and Uriu, 1989).
Even though flower density and fruit set are
generally not affected by N, overall yields are
decreased because of reduced fruit size and
less fruiting sites due to shorter shoots. With
severe deficiency, fruit are more astringent
and fibrous and of inferior eating quality
(Proebsting et al., 1957; Ballinger et al., 1966).
When N levels are too high, the most
obvious problems arise from excessive vege-
tative growth (Lobit et al., 2001). The resultant
shading causes less red coloration on the fruit,
a delay in maturity and can lead to extensive
Fig. 13.2. Red coloration on the leaves and stems of a nitrogen-deficient peach shoot.
dying out of fruiting shoots on the inside and
bottom of the canopy (Ballinger et al., 1966).
Furthermore, pruning costs are more and it
has been demonstrated that yields and fruit
size are not increased compared with more
moderate fertilization rates (Daane et al., 1995).
Also, many pest and disease problems tend to
be exacerbated at high N levels (Daines et al.,
1958; Jafar, 1958; Daane et al., 1995). Despite
the widespread belief that high N harms fruit
quality and storability, there are few scientific
studies to support this claim (Claypool, 1975).
Greater water loss due to a thinner cuticle has
been reported, but often researchers have
found no differences in firmness, soluble sol-
ids content, acidity, bruising potential or
internal breakdown (Daane et al., 1995).
Because of the increasing concern with N
pollution in the environment much attention
has been paid to tactics for improving N
uptake efficiency in peach trees. One approach
that has shown promise is the use of foliar
urea. Early studies showed potential for many
fruit trees such as apple, but it did not appear
to work as efficiently for peach (Weinberger
et al., 1949; Norton and Childers, 1954; Leece
and Kenworthy, 1971; Swietlik and Faust, 1984).
 Nutrient and Water Requirements
Fig. 13.3. Smoother finish and more red coloration on fruit from low-nitrogen trees (top row)
compared with high-nitrogen trees (bottom row).
Recently, using more precise techniques (such
as 15N), it has been demonstrated to be an
effective method of supplying N to peach
trees, especially when applied in the autumn
before leaf senescence (Rosecrance et al.,
1998a; Furuya and Umemiya, 2002). Estimates
of 60–70% uptake of foliar urea have been
reported as long as the application is not
made too late in the season (Rosecrance et al.,
1998b; Tagliavini et al., 1998). Much of that N
is then mobilized out of the leaf and stored in
perennial structures of the tree for use by new
growth the next spring. Many of the early
studies used low concentrations of urea solu-
tions (generally 1% or less) in order to avoid
phytotoxicity. Only a small percentage of total
N needs could be supplied in this way and
may be part of the reason for apparent failure
of these studies. Recent approaches have used
higher concentrations (5–10%) or multiple
sprays in the autumn when leaf phytotoxicity
is not a major concern and have shown sub-
stantial increases in stored N. Attempts to
supply total N needs through foliar urea led
to a decrease in fruit size, but the approach of
replacing half of soil-applied N with autumn
foliar urea showed no loss of productivity or
fruit quality (Johnson et al., 2001). Therefore,
307
this is one strategy available to growers for
reducing the potential of environmental pol-
lution by substantially reducing soil-applied
fertilizers, without risking a loss of produc-
tion. Furthermore, it has been demonstrated
that the urea spray can be combined with
other materials commonly applied in the
autumn such as Zn, thus making it a practical
strategy as well (Johnson and Andris, 2001).
Some recommendations now include autum-
nal foliar urea sprays as a supplement to the
standard soil applications (Rombola et al.,
1995; Tagliavini and Marangoni, 2002).
Other approaches to improving N utili-
zation efficiency involve more precise timing,
placement and amounts of soil-applied N fer-
tilizers. Applying fertilizers through a low-
volume irrigation system (fertigation) has
generally demonstrated that considerably
less fertilizer (sometimes about half) can be
used compared with broadcast fertilization
and still have the same growth and leaf N
response (Smith et al., 1979; Neilsen et al.,
1999). Even when using broadcast fertiliza-
tion, splitting the applications into several
smaller amounts increases efficiency. N uptake
by mature peach orchards has been estimated
to be between 0.5 and 1.0 kg N/ha/day
308 R.S. Johnson
during the period from rapid shoot growth
until early autumn (Rufat and DeJong, 2001).
Furthermore, this fairly constant uptake rate
appears to be similar for early- and late-
maturing varieties (Policarpo et al., 2002). The
uptake rate is much lower early in the spring
and late in the autumn, especially after leaves
begin to senesce. This information could be
used as a guide to help meter out small amounts
of fertilizer based on tree demand throughout
the season. Tagliavini et al. (1996) developed a
fertilization strategy based on this concept
combined with soil sampling at three differ-
ent periods to estimate N available to trees.
The method was used in 15 different orchards
and the resulting fertilizer usage was 50%
that of other orchards in the area with no
noticeable reduction in yield or growth. In the
future, these types of approaches will be
increasingly important as more environmen-
tally sound practices will be required.
Severe deficiency is generally found at
mid-summer leaf N concentrations below 2.2
to 2.4% (Table 13.1) (Leece et al., 1971; Johnson
and Uriu, 1989; Weir and Cresswell, 1993; Rob-
inson et al., 1997). In the past, adequate or nor-
mal levels have usually been set at 3.0 to 3.5%
(Leece et al., 1971; Weir and Cresswell, 1993;
Robinson et al., 1997). However, with increased
concerns over environmental contamination,
and with recent studies documenting the neg-
ative aspects of high N, it is advisable in most
situations to keep leaf N concentrations
slightly below 3.0% (Johnson and Uriu, 1989).
Phosphorus
P is the key factor in compounds that store,
transfer and utilize energy in plants. It is also
found in nucleic acids, the building blocks for
DNA. There have only been a few reports of P
deficiency in field-grown peach trees around
the world (Lilleland, 1932; Veerhoff, 1948)
and fertilization with this nutrient has often
been ineffective (Proebsting and Kinman,
1933; Ballinger et al., 1966). Even in soils where
field crops have dramatically responded to P
fertilization, peach trees (and other fruit and
nut trees) have shown no response (Lilleland
et al., 1942). Nevertheless, there are soils very
low in P where peach trees have shown a
growth response to added P (Lilleland and
Brown, 1940). In addition, there are a few doc-
umented situations of peach trees responding
to P even when soil or tree levels have indi-
cated adequate amounts (Taylor, 1975; Neilsen
et al., 1990). Generally the response has been
one of increased growth, and untreated trees
in these situations have shown no leaf defi-
ciency symptoms. Thus, although many consi-
der it unnecessary, some local recommendations
call for P fertilization as a standard practice,
especially at planting (Van Niekerk and
Pienaar, 1967; Sánchez, 1999; Gil Salaya, 2000).
P deficiency has been induced in young
trees growing in sand culture. One of the first
symptoms is a reduction in growth with no
abnormal leaf symptoms (Weinberger and
Cullinan, 1936; Lilleland and Brown, 1940).
With more severe deficiency, leaves develop a
dark purplish-green colour (Fig. 13.4/Plate
80) and are narrow, leathery and flat (Cullinan
and Batjer, 1943). Fruit ripen earlier and have
a greenish ground colour with poor eating
quality (Shear and Faust, 1980). A mid-sum-
mer leaf P level below 0.09 to 0.11% is gener-
ally considered deficient (Table 13.1) (Lilleland
and Brown, 1940; Leece et al., 1971; Shear and
Faust, 1980; Robinson et al., 1997). We have
found a reduction in growth below 0.12% P in
bearing peach trees growing in sand culture
(R.S. Johnson, unpublished results). Healthy
trees are usually between 0.14 and 0.25% P
(Robinson et al., 1997), with some of the most
productive orchards often in the lower part of
this range (Lilleland and Brown, 1942). Prob-
lems of excess P can be encountered at leaf
levels above about 0.40% (Leece et al., 1971;
Robinson et al., 1997). High P can exacerbate a
Zn deficiency situation (Ballinger et al., 1966).
When needed, any P-containing fertilizer
is effective at raising the P level in trees. Taylor
and Issell (1976) found soil applications to be
more effective than foliar applications at
improving the P status of peaches.
Potassium
K is very mobile within plants and can move
readily in and out of cells. Because of this
characteristic, K plays an important role in
maintaining cell turgor and in the opening
 Nutrient and Water Requirements
and closing of stomata. It is also the activator
of many different enzyme systems. At one time,
K deficiency was widely reported throughout
many peach-growing areas of the world
(Scott, 1939; Boynton, 1944; Lilleland et al.,
1962; Ballinger et al., 1966). Although it can
still be a problem in some stone fruit orchards
(such as prunes in California), it is generally
not a major concern in modern peach orchards
(Johnson and Uriu, 1989; Weir and Cresswell,
1993). The deficiency can be easily corrected
with K-containing fertilizers, which are often
recommended as a standard practice (Taglia-
vini and Marangoni, 2002). The amount of K
in peach fruits is relatively large so K removal
from the orchard can be substantial with
highly productive trees.
The most characteristic symptom of K
deficiency is a pale green leaf colour and leaf
rolling that appears in mid-summer (Fig.
13.5/Plate 81) (Dunbar and Anthony, 1937;
Lilleland et al., 1962; Weir and Cresswell,
1993). Eventually necrosis can occur at the
309
Fig. 13.4. Purple coloration and
leathery texture of a phosphorus-
deficient peach leaf.
leaf margins, producing a ‘scorched’ appear-
ance. With severe deficiency, shoot growth is
inhibited, leading to a small, unthrifty-looking
tree. The shoots are thin and reddish and few
flower buds are initiated (Dunbar and Anthony,
1937). The fruit produced are smaller (Lille-
land et al., 1962), advanced in maturity and
have poor colour. There is some controversy
over the effect of K on fruit quality. Several
studies have shown a positive correlation
between leaf K and fruit acidity (Kwong and
Fisher, 1962; Cummings and Reeves, 1971) or
between leaf K and fruit colour (Cummings,
1965), but no relationship with fruit soluble
solids content. Others have shown no effect of
K on firmness, keeping quality or acidity
(Weinberger, 1929). Tagliavini and Marangoni
(2002) have suggested that peach trees might
benefit from more efficient K fertilization
using fertigation. They provide evidence that
fruit size, per cent soluble solids content and
colour can all be improved even when the
trees are not deficient in K (1.35–1.6% K).
310
Fig. 13.5. Pale colour and leaf rolling caused by potassium deficiency in peach leaves.
A mid-summer leaf K value below 0.75 to
1.0% (Table 13.1) indicates a definite deficiency
and K fertilization will generally provide a
positive response (Boynton, 1944; Ballinger
et al., 1966; Shear and Faust, 1980). Values
between 1.0 and 2.0% are considered low or
marginal (Leece et al., 1971; Weir and Cresswell,
1993; Robinson et al., 1997) and increasing the
level may be beneficial in some situations.
Most normal, productive peach trees are
between 2.0 and 3.0% K. Correction of K defi-
ciency is accomplished with soil applications
of various formulations of K-containing fer-
tilizers. Generally the recommendation is to
make a single application every 3 to 4 years
(Weir and Cresswell, 1993). Fertigation has
been shown to be a more efficient method of
supplying K to trees (Johnson and Uriu, 1989).
Care should be taken to prevent over-applica-
tion of K to peach trees. Although K toxicity
has not been reported, an excess can easily
lead to cation imbalance in the soil and cause
R.S. Johnson
deficiencies of other nutrients such as Mg
(Weir and Cresswell, 1993).
Calcium
Ca is a major constituent of cell walls and
membranes and plays a role in their proper
functioning. It is also involved with pollen
germination, cell division, environmental sig-
nalling and protecting cells from toxins. Ca
deficiency is generally not a problem with
peach trees. Even though dozens of Ca-related
disorders have been identified in other horti-
cultural species (Shear, 1975), none have been
reported in the scientific literature for peaches.
Likewise, Ca deficiency in field-grown trees
has not been documented (Shear and Faust,
1980; Johnson and Uriu, 1989; Weir and Cress-
well, 1993).
Symptoms of Ca deficiency have been
induced in hydroponically grown peach
 Nutrient and Water Requirements
seedlings (Edwards and Horton, 1979) and in
small fruiting trees grown in sand culture
(Abdalla and Childers, 1973). One of the first
symptoms to appear is a reduction in root
growth. Subsequent roots that develop are
often swollen and stubby. Early leaf symp-
toms include marginal leaf chlorosis, which
develops into necrosis and eventually leads
to defoliation. Some shoot tips die back. Fruits
on Ca-deficient trees are smaller, lower in sugar,
firmer and have poorer colour and flavour.
Deficiency thresholds for peach leaves have
generally been set at 1.0% (Table 13.1) (Shear
and Faust, 1980; Weir and Cresswell, 1993;
Robinson et al., 1997) even though this has
never been confirmed in the field. Abdalla
and Childers (1973) found optimum fruit
quality in sand culture trees at leaf values
greater than 2.0%.
Because Ca is found in cell walls, it has
been associated with cell rigidity and fruit
firmness. Addition of Ca has successfully
delayed softening of various fruits (Pooviah
et al., 1988) and reduced decay (Conway, 1982).
Results with peaches have not been very con-
sistent. There have been several reports that
preharvest applications of Ca sprays have
reduced fruit rot, maintained firmness and
improved flavour, aroma and appearance of
peaches (Bhullar et al., 1981; Robson et al., 1989;
Adaskaveg et al., 1992; Biggs et al., 1997). Oth-
ers, however, have measured no benefit from
as many as 10 to 12 sprays of Ca throughout
the season (Conway et al., 1987; Crisosto et al.,
2000). Conway et al. (1987) were able to dra-
matically increase (two- to fourfold) the Ca
content of fruit using pressure infiltration,
which reduced decay by 40 to 60%. However,
the treatment caused injury to the surface of
the fruit. Likewise, Wills and Mahendra (1989)
used a similar procedure and increased fruit
storage life from 11.1 to 14.4 days, but induced
the same skin damage. Therefore the proce-
dure is unlikely to have commercial value.
Magnesium
Mg is a part of the chlorophyll molecule and
also acts as an activator of many different
enzymes. In most peach-growing areas of the
world Mg deficiency is not considered a major
311
problem. The disorder can occur in some
locations with very sandy acidic soils, espe-
cially if there is extensive leaching by high
rainfall or excessive irrigation (Beyers and
Terblanche, 1971d; Weir and Cresswell, 1993).
Also, Mg deficiency can be induced by heavy
applications of K- and Ca-containing fertiliz-
ers. These nutrients compete with Mg for cat-
ion exchange sites in the soil. In fruit-growing
areas where Mg deficiency occurs, peach is
considered to be less affected than other fruit
trees and vines (Beyers and Terblanche, 1971d).
Mg deficiency symptoms start as a pale
green discoloration at the tips and margins of
older leaves (Fig. 13.6/Plate 82). The affected
area develops into a bright yellow chlorosis and
eventually some marginal necrosis may occur.
Reddish brown areas may develop within the
chlorotic areas. A triangular area at the base of
the leaf remains green (Beyers and Terblanche,
1971d; Weir and Cresswell, 1993). Additional
symptoms have been described as interveinal
necrotic spotting which starts as areas on the
leaves that appear to be water-soaked
(McClung, 1953). Some defoliation can occur,
starting with basal leaves. Unless the disorder
becomes severe, normal extension growth con-
tinues and tree productivity is not reduced.
With severe deficiency shoot growth stops,
flower bud formation is inhibited and yield is
greatly decreased. The fruit that develop are
small, have poor colour and often fail to mature.
Deficiency symptoms develop at mid-
summer leaf levels below 0.20 to 0.25% Mg
(McClung, 1953; Shear and Faust, 1980; Johnson
and Uriu, 1989; Weir and Cresswell, 1993;
Robinson et al., 1997) although some research-
ers have set the deficiency threshold as low as
0.10% or as high as 0.30% (Table 13.1) (Leece
et al., 1971; Sánchez, 1999). If correction is neces-
sary, soil-applied fertilizers or foliar sprays con-
taining Mg have proved effective. Fertilizers
applied to the soil may take 2 to 3 years to com-
pletely eliminate symptoms, while foliar appli-
cations act faster but provide a more temporary
response (Beyers and Terblanche, 1971d).
Sulfur
S is a part of two amino acids and thus is
a component of many proteins and enzymes.
312 R.S. Johnson
It can be supplied to peach trees from many
sources including fertilizers, soil amend-
ments, pesticides, irrigation water, acid rain
and even atmospheric pollutants. Therefore,
S deficiency has seldom been encountered in
peach-growing areas and often is not included
in nutrient deficiency threshold tables. A few
researchers have estimated the optimum
range to be about 0.17 to 0.40% (Leece et al.,
1971; Robinson et al., 1997). An early study,
working with young potted trees, set the defi-
ciency threshold at 0.01% (100 ppm) (Benson
et al., 1963). Recently, a more thorough experi-
ment determined the threshold to be much
higher (Finch et al., 1997). These researchers
set the threshold at 0.09% (Table 13.1) with
adequacy levels ranging from 0.14 to 0.25%.
There have been a few sketchy reports of
S deficiency in the field (Shear and Faust,
1980; Johnson, 1993) and some evidence of
positive growth responses to S applications
(Powell et al., 1995; Finch et al., 1997). Descrip-
tion of deficiency symptoms has mainly come
Fig. 13.6. Discoloration on leaf
margins and tip caused by magnesium
deficiency.
from potted tree experiments (Benson et al.,
1963; Finch et al., 1997). Symptoms of S defi-
ciency are very similar to N deficiency, with
reduced growth and smaller, yellow leaves.
The main difference is that S deficiency tends
to start with young leaves at the tip of the
shoot. Also, with severe S deficiency, necrosis
can develop along leaf margins.
Zinc
Zn is involved in many functions in plants as
it has been found in over 80 proteins. One of
its roles is in the formation of the plant hor-
mone auxin. A lack of auxin leads to stunting
of leaves and shoots, one of the characteristic
symptoms of Zn deficiency. The importance
of Zn in peach trees was first discovered in
California with the identification of a ‘little
leaf’ disorder (Chandler et al., 1931). This dis-
order was subsequently associated with Zn
deficiency (Chandler, 1937) and has since
 Nutrient and Water Requirements
been found in almost all peach-growing areas
on many different soil types around the world
(Takkar and Walker, 1993; Swietlik, 1999). It is
considered a major problem in Australia (Weir
and Cresswell, 1993) and many locations in
North America (Bell and Childers, 1954).
Peach trees are more sensitive to this disorder
than many other crops (Swietlik, 1999). Initial
symptoms of the deficiency include interveinal
chlorosis (Fig. 13.7/Plate 83) which is hard to
distinguish from Mn deficiency (Bollard, 1953;
Woodbridge, 1954). As the disorder becomes
more severe, shortened internodes and nar-
row, pointed leaves at the end of shoots give
the characteristic rosetting or little leaf symp-
toms (Fig. 13.8/Plate 84). Often leaves have a
wavy margin. Eventually defoliation of older
leaves and twig dieback can occur and fruit
production is greatly reduced. What fruit are
produced have a tendency to be smaller, more
flattened and break down earlier than normal
(Chandler, 1937). Zn toxicity is rare but can be
induced by over-fertilization with Zn chelates
(Wallace et al., 1983).
Early surveys of Zn-deficient peach
orchards generally found mid-summer leaf
Fig. 13.7. Zinc deficiency symptoms of interveinal chlorosis on a peach leaf.
313
Zn levels to be below 10 to 15 ppm. However,
sometimes leaf levels as high as 25 ppm were
reported (McClung, 1954) and some healthy
orchards without symptoms tested as low as 6
to 10 ppm (Bollard, 1953; McClung, 1954;
Woodbridge, 1954). Therefore, there has not
been general agreement on establishing a defi-
ciency threshold. In reviewing the literature
for peach, Shear and Faust (1980) set the
threshold at 12 ppm. However, others have
often used higher values of 15 ppm (Leece
et al., 1971; Johnson and Uriu, 1989; Robinson
et al., 1997) or even 20 ppm (Sánchez, 1999; Gil
Salaya, 2000) (Table 13.1). One problem with
setting the threshold too high is that unneces-
sary corrective measures might be imposed.
Sánchez and Righetti (2002) found many high-
yielding fruit orchards had Zn levels between
12 and 16 ppm, and concluded that either the
generally accepted threshold of 18 to 20 ppm is
too high or mid-summer leaf sampling does
not adequately reflect the Zn status of the tree.
Generally, the recommended treatment
for Zn deficiency is foliar applications of
ZnSO4. However, suggested rates and timing
vary considerably (Bell and Childers, 1954;
314
Fig. 13.8. Rosetting or ‘little leaf’ symptoms of zinc deficiency in peaches.
McClung, 1954; Johnson and Uriu, 1989; Weir
and Cresswell, 1993; Neilsen and Neilsen, 1994).
Soil applications have also been effective but
require much higher rates (Bell and Childers,
1954; Mann et al., 1986). Soil-applied Zn che-
lates have shown efficacy but are generally
considered less cost-effective than ZnSO4
(Arce et al., 1992). Perhaps a better under-
standing of mycorrhizal association with
peach roots could help improve Zn uptake
efficiency. Gilmore (1971) showed that some
cultures of mycorrhizae were more effective
than others at alleviating Zn deficiency symp-
toms in peach seedlings.
In the past, it has been widely accepted
that Zn is not very mobile within the plant
and that foliar treatments may never correct a
root deficiency (Swietlik, 2002a). However,
recent research has demonstrated the move-
ment of Zn into roots of peach trees from a
foliar application in autumn (Sánchez et al.,
2006). Clearly, more research is needed and
should focus on critical periods for Zn supply
in the tree (Swietlik, 2002b) and the behaviour
of Zn in the soil and plant, so that effective
treatments can be made with a minimal
impact on the environment.
R.S. Johnson
Boron
B is an essential nutrient for plant growth and
development, but its exact physiological role
is still poorly understood. When deficient it
can affect many plant processes including
pollen tube growth, sugar transport, meri-
stem growth, cell wall synthesis, hormone
production and membrane integrity. B defi-
ciency is widespread around the world, hav-
ing been reported in at least 132 crops from
over 80 countries (Shorrocks, 1997), and is
often considered to be a major problem. How-
ever, peach is much less sensitive than most
other plants and only a few cases have been
documented in the field (McLarty and Wood-
bridge, 1950; McClung and Clayton, 1956;
Ballinger et al., 1966). More recent research
indicates that B is very mobile in sorbitol-rich
plants such as peach and apple (Brown and
Hu, 1996). An easily translocated compound
is formed from B and sorbitol (and other
sugar alcohols) that accounts for this mobility
(Hu et al., 1997). This provides at least part of
the explanation for why peach is less sensi-
tive than many other plants to B deficiency.
However, peach is generally considered to be
 Nutrient and Water Requirements
even less sensitive than many other sorbitol-
producing fruit trees such as apple and pear
(Ballinger et al., 1966; Weir and Cresswell,
1993), so additional factors may be involved.
The high mobility of B in peach also
makes it more sensitive to toxicity than many
other crops (Gupta et al., 1985). Although
there are few reports of B toxicity in the field
for peach orchards, care needs to be taken to
prevent it from happening when applying B
fertilizers, as there appears to be a rather nar-
row range between deficiency and toxicity.
The reports of B toxicity in the literature have
been associated with excessive soil applica-
tions (Williams and Veerhoff, 1948; McLarty
and Woodbridge, 1950; Cibes et al., 1955; Dye
et al., 1984). Furthermore, once B toxicity has
been induced, it can take several years to alle-
viate the problem (Hernandez and Childers,
1956). Even though B can be leached from the
soil fairly easily, it appears that this nutrient
can be stored in relatively large amounts in
peach trees (Dye et al., 1984), thus carrying on
the problem from one year to the next.
The first symptom of B deficiency is the
dying back of shoots and twigs in the spring
(McLarty and Woodbridge, 1950; Ballinger
et al., 1966; Kamali and Childers, 1970). These
shoots apparently grew normally the year
before. In fact, there appears to be no warning
of the disorder beforehand such as gumming
or necrosis. The first sign of a problem occurs
when buds on the shoots fail to break in the
spring. These buds slowly turn brown and
die and the stem tissue may continue to look
normal for another couple of weeks before
drying up. This dieback can occur over a large
portion or even the whole tree. Vegetative
buds that do survive often produce normal
growth, although the first leaves to develop
may be narrow and small, with edges rolled
inward. McClung and Clayton (1956) described
somewhat different symptoms. They did not
see shoot dieback initially. Instead, the vegeta-
tive buds grew but produced highly distorted
shoots with small leaves and shortened inter-
nodes. The leaves were often asymmetrical
with irregular and chlorotic margins. In
extreme cases these shoots would die gradu-
ally, leading to dieback by early summer. In
less severe cases the shoots eventually pro-
duced normal growth by mid-summer.
315
Fruit set is often greatly reduced by B defi-
ciency (McClung and Clayton, 1956) as is gen-
erally seen for all fruit trees (Shear and Faust,
1980). The fruit that do develop can appear
normal on the outside, but necrotic areas may
occur around the pit and the fruit may ripen
early. Fruit set and fruit disorders often occur
without any leaf symptoms, indicating the
particular sensitivity of reproductive processes
to B deficiency (Shear and Faust, 1980).
B toxicity has symptoms that can easily
be confused with B deficiency (Dye et al.,
1984; Neilsen et al., 1985). As with the defi-
ciency, the first sign of toxicity is shoot die-
back in the spring or early summer (McLarty
and Woodbridge, 1950; Cibes et al., 1955; Dye
et al., 1984). Leaves produced can be small
and crinkled. Sometimes they have small
necrotic areas near the midrib which eventu-
ally drop out, leaving a perforated appear-
ance. In sand culture, leaves became wrinkled
and necrotic along the entire margin (Haas,
1929). Other symptoms include cankers along
the shoots, profuse gumming and rough and
cracked bark. Often fruit that develop are
malformed, showing lopsided growth, crack-
ing and prominent sutures.
Using mid-summer mature leaf samples
to establish thresholds for B deficiency and
toxicity has not been very straightforward.
First, there is not always a good correlation
between symptom severity and leaf B content
(McClung and Clayton, 1956). Second, the
effects of B deficiency on fruit set and fruit
disorders are sometimes evident without any
apparent leaf symptoms (McClung and Clay-
ton, 1956). Williams and Veerhoff (1948) found
improvements in fruit size by applying B to
peach trees showing no deficiency symptoms.
Thus, it may be difficult to determine the effect
of B on more subtle processes in the plant.
Nevertheless, researchers have generally
established deficiency thresholds at about 15
to 20 ppm (McLarty and Woodbridge, 1950;
Ballinger et al., 1966; Leece et al., 1971; Shear and
Faust, 1980; Weir and Cresswell, 1993; Robinson
et al., 1997; Gil Salaya, 2000), with some as high
as 30 ppm (Table 13.1) (Sánchez, 1999). For
toxicity, the threshold is generally set to about
80 to 100 ppm (McLarty and Woodbridge,
1950; Leece et al., 1971; Robinson et al., 1997),
but others have suggested damage may occur
316 R.S. Johnson
as low as 50 ppm (Neilsen et al., 1985). Improve-
ments in the sampling procedure could be
very helpful, since there appears to be such a
narrow range of B for optimum performance.
Dye et al. (1984) found spring leaves to be
more diagnostic of B toxicity than mid-
summer leaves. For most field crops, newly
emerged leaves are generally used for B deter-
mination (Bell et al., 2002). Alternatively, fruit
tissue could be useful for diagnosis since B
accumulates there in B-mobile plants such as
peach (Hernandez and Childers, 1956; Dye
et al., 1984; Shu et al., 1994). This approach has
proved effective in another Prunus species,
almond (Nyomora et al., 1997).
Correction of B deficiency is best achieved
by foliar applications. Soil applications can
also be effective, but many factors such as soil
pH and texture can affect B availability, mak-
ing this method less predictable. Further-
more, due to the extreme sensitivity of peach
to B toxicity, damage has been reported even
at recommended rates of soil application.
Recommended rates vary from 10 to 30 kg/ha,
but often only every third to fifth year. Recom-
mended foliar rates range from 0.5 to 2 kg/ha.
Although studies with labelled B have shown
leaf and fruit uptake to be less than 0.5% (Shu
et al., 1993, 1994, 1997), this rate appears to be
enough to correct B deficiency. For correcting
B toxicity it is necessary to leach excess B
from the soil. This may take three times more
water than is needed for other salts because B
can be adsorbed by soil particles (Gupta et al.,
1985).
Iron
Fe in plant cells can easily be transformed
from one oxidation state to another, giving up
(or capturing) energy in the process. Thus, its
main role is in transferring energy during the
processes of photosynthesis and respiration.
It is very rare for Fe to be deficient in orchard
soils. However, under calcareous soil condi-
tions with high pH (7.5 to 8.5), Fe can become
immobilized within the tree, leading to a con-
dition called lime-induced or iron chlorosis.
Peach trees are particularly susceptible to this
type of Fe deficiency and it is a major problem
in many peach-growing areas around the world
(Ballinger et al., 1966; Razeto, 1982; Byrne, 1988).
In some locations it affects more than half the
orchards and is considered the main nutritional
disorder for fruit trees (Sanz et al., 1992a; Taglia-
vini et al., 2000). Peach trees are considered more
susceptible than most other fruit species to Fe
deficiency (Razeto, 1982).
The characteristic symptoms of iron chloro-
sis are a ‘netting’ appearance on the leaves, where
the veins remain green but all areas in between
turn yellow (Fig. 13.9/Plate 85) (Johnson and
Uriu, 1989; Weir and Cresswell, 1993). It starts
with young leaves but can eventually spread to
the whole tree. Severely affected leaves become
bleached as they lose chlorophyll. Eventually
these leaves develop burnt spots and extensive
shoot dieback can occur. With severe deficiency,
shoot growth, flowering, fruit production and
fruit size are greatly decreased (Beyers and Ter-
blanche, 1971c; Sanz et al., 1997b).
This disorder does not appear to be a
straightforward Fe deficiency as the total Fe
level in mid-summer leaves does not corre-
late well with deficiency symptoms (Shear
and Faust, 1980; Abadia et al., 2000). Appar-
ently, various factors such as pH and bicar-
bonate levels can cause the Fe in the plant to
become unavailable (Köseoglu, 1995; Morales
et al., 1998). Attempts to correlate available Fe
with symptoms have shown promise (Abadia
et al., 1985) but have not always been success-
ful (Rashid et al., 1990). Therefore, it has been
a challenge to develop a diagnostic tool for
quantifying the problem. There has been
some limited success with using leaf Fe val-
ues from leaves collected in the spring rather
than mid-summer. Recently, flower analysis
has been proposed and good correlations
have been shown between flower Fe values
and mid-summer symptoms (Sanz et al.,
1997a; Abadia et al., 2000). However, absolute
levels vary considerably from year to year, so
recommendations cannot yet be made. Oth-
ers have had less success with this approach
(Toselli et al., 2000). Perhaps other nutrients
are involved in the disorder and a multiple
regression formula will need to be developed.
Igartua et al. (2000) have proposed using the
K:Zn ratio in addition to flower Fe for diag-
nosing iron chlorosis.
Correction of iron chlorosis has also
proved to be challenging. Foliar applications
 Nutrient and Water Requirements
Fig. 13.9. ‘Netting’ symptom of iron chlorosis on a peach leaf; veins remain green while the rest of
the leaf has turned yellow.
with various materials have given variable
results and, when effective, provide only tem-
porary correction of symptoms (Razeto, 1982;
Reed et al., 1988; Vizzotto and Costa, 1995).
Trunk injection treatments have tended to
work a little better, but are much more labour-
intensive and still do not correct the problem
permanently (Ballinger et al., 1966; Yoshikawa,
1988). Soil application of chelated materials
such as Fe EDDHA (ethylenediaminedi(o-hy-
droxyphenyl)acetic acid) has been one of the
more effective treatments, but it is very expen-
sive (Beyers and Terblanche, 1971c; Reed et al.,
1988; Sanz et al., 1992a). Apparently, lower
rates can be used if application is made through
a low-volume irrigation system (Razeto, 1982).
Ultimately, the best solution is to lower soil
pH with acidifying amendments. This can be
prohibitively expensive for the whole soil
mass, but also works if only a small portion of
the soil is acidified (Razeto, 1982; Johnson
and Uriu, 1989). Selection of rootstock can
make a big difference as some stocks are more
resistant to iron chlorosis (Kester and Asay,
1986; Stylinanides et al., 1989). ‘GF 677’ and
317
other peach–almond hybrids have performed
well on sites prone to this disorder (Syrgian-
nidis, 1985; Byrne, 1988).
Manganese
Mn plays an important role in photosynthesis
and also activates a number of enzyme sys-
tems. Mn deficiency has been documented in
most peach-growing areas around the world.
Generally, the symptoms observed have been
quite minor, and are not considered a serious
problem since yield and vegetative growth
are not noticeably affected (Epstein and Lille-
land, 1942; Boynton et al., 1951; Woodbridge
and McLarty, 1951; Beyers and Terblanche,
1971a; Rogers et al., 1974; Johnson and Uriu,
1989; Weir and Cresswell, 1993; Gil Salaya,
2000). Severe deficiency has also been reported
(Beyers and Terblanche, 1971a), but is rare.
Symptoms start as small, irregularly shaped,
light green spots in the interveinal and mar-
ginal areas of older leaves. Eventually these
318 R.S. Johnson
spots grow together while the area along the
veins and midribs remains green, giving the
characteristic ‘herringbone’ pattern of Mn
deficiency (Fig. 13.10/Plate 86). Generally fruit
size, leaf size and shoot growth are not affected.
Severe symptoms include dieback of termi-
nals and premature defoliation with a marked
reduction in flowering and fruit set. Almost
all nutrient deficiency tables indicate 20 ppm
in mid-summer leaves to be the threshold for
peaches (Table 13.1) (Leece et al., 1971; Shear
and Faust, 1980; Johnson and Uriu, 1989; Weir
and Cresswell, 1993; Robinson et al., 1997).
Most researchers have found that a foliar spray
of MnSO4 in the spring is the most effective
method of alleviating symptoms (Boynton
et al., 1951; Woodbridge and McLarty, 1951;
Beyers and Terblanche, 1971a; Gil Salaya, 2000).
Copper
The main role of Cu in plants is energy transfer
during photosynthesis, but it is also involved
in several other plant processes. Despite a very
old report of severe Cu deficiency in peach and
Fig. 13.10. Manganese deficiency symptoms on a peach leaf; bands along the main veins remain
green.
other fruit trees (Anderssen, 1932), this nutri-
ent never seems to create major problems in
modern peach orchards. Perhaps this is due to
the widespread use of Cu as an effective and
inexpensive broad-spectrum pesticide (Epstein
and Bassein, 2001). Where Cu deficiency has
been reported, it was more of a problem on
other fruit trees such as apple, pear and citrus
(Beyers and Terblanche, 1971b). Deficiency
symptoms start with a pale green to bright yel-
low discoloration between the veins of leaves
in the spring. Eventually growing shoot tips
wither and die back, and multiple bud devel-
opment below leads to abnormal branching
and a bushy appearance. Extensive dieback of
branches can occur. Affected trees often have a
light or no crop. The deficiency threshold has
generally been set at 3 ppm Cu in mid-summer
leaves (Table 13.1) (Shear and Faust, 1980; Weir
and Cresswell, 1993; Robinson et al., 1997).
13.3 Peach Water Requirements
Peach trees tend to have high water requirements
compared with other fruit trees. Most experts
 Nutrient and Water Requirements
agree that China is the location where peaches
evolved (Faust and Timon, 1995). The areas of
China with heavy concentrations of native
trees are often quite humid. Thus, it seems
likely that peaches evolved under conditions
of abundant water and are therefore not very
water-conservative or resistant to water stress
(Proebsting and Middleton, 1980). From a
practical perspective, orchard managers have
often found supplemental irrigation to be
beneficial, even in locations with high rainfall
(Feldstein and Childers, 1965; Reeder et al.,
1979; Horton et al., 1981; Layne et al., 1981;
Layne and Tan, 1984). One of the biggest
benefits has been an increase in fruit size
(Daniell, 1982). Other benefits have included
more uniform fruit ripening (Feldstein and
Childers, 1965) and less winter injury and tree
disease (Layne and Tan, 1984). In colder
climates, if supplemental irrigation is contin-
ued too late in the season so the tree does not
harden off as well, flower bud hardiness and
tree survival can be reduced (Layne et al.,
1994). Supplemental irrigation has also shown
substantial benefits with young trees (Black
et al., 1977; Daniell, 1982; Renquist, 1987),
even when there was no subsequent advan-
tage once the trees matured (Layne et al., 1981,
1994). Generally, the trees grew considerably
better and thus came into production much
earlier. One study has shown no benefit
from irrigation in a humid climate, but the
experimental site had a shallow water table
(1–2 m) and wetter than normal conditions
during the 4 years of the experiment (Layne
et al., 1996).
Too much water can lead to problems
with peach trees as they are considered one of
the most sensitive fruit species to waterlog-
ging and anaerobic soil conditions (Andersen
et al., 1984; Alvino et al., 1986; Schaffer et al.,
1992). Therefore, in heavier soils or those with
an impermeable hardpan layer, irrigation
management is much more critical. Even in
lighter-textured soils, overirrigation can lead
to iron chlorosis symptoms (Johnson and
Uriu, 1989) and greater probability of diseases
such as Phytophthora root rot (Teviotdale et al.,
1989).
In drier peach-growing areas around the
world, irrigation is a standard practice. Much
research has been conducted to determine the
319
amount of water to apply to an orchard of
peach trees. Orchards lose water through soil
evaporation and through transpiration from
the leaves. The combination of these two is
termed evapotranspiration (ET) and can be
affected by several factors. First, weather con-
ditions can have dramatic effects on orchard
ET. In a hot, sunny and dry climate, trees will
use much more water than under cool, cloudy
and humid conditions. The combined effect
of weather conditions can be estimated by
evaporation from an open body of water (pan
evaporation) or by measuring water loss from
a reference crop, typically mown grass. The
latter is termed ET0 and is derived from
weather station parameters. When orchard
ET is compared with ET0 (or pan evapora-
tion), a fairly constant ratio is obtained and is
called a crop coefficient, Kc. Using various
measures of tree water use such as drainage
lysimeters or root zone soil moisture deple-
tion, Kc values well over 1.0 (Chalmers et al.,
1983; Natali et al., 1985b; Tormann and Heyns,
1988) and even as great as 1.5 (Boland et al.,
1993) have been reported for peach trees.
Maximum Kc values less than 1.0 have also
been measured, but often under less than
optimum conditions such as under fertilized
or isolated trees (Worthington et al., 1984;
Garnier et al., 1986), or under conditions of
high rainfall (Strabbioli, 1992a). Using a
weighing lysimeter (Phene et al., 1991), which
provides the most accurate measure of tree
water use, mid-summer Kc values of 1.1 to 1.2
(based on ET0) have been reported for
unstressed trees (Ayars et al., 2003; Johnson
et al., 2005). Thus, healthy peach orchards use
a large amount of water. Supplemental irriga-
tion of more than 1000 mm may be required
in hot, dry areas (Ayars et al., 2003).
Another factor affecting orchard ET is
leaf area development, as it has been shown
that tree water use is directly proportional to
light interception by the canopy (Johnson et al.,
2000; Goodwin et al., 2006) (Fig. 13.11). Since
peach trees are pruned hard, it takes several
months for the canopy to develop fully. There-
fore, the seasonal Kc pattern for peach starts
low, about 0.1 to 0.2, early in the season
(Ayars et al., 2003). Kc values then increase
steadily as leaf area and therefore canopy light
interception continue to expand (Fig. 13.12).
320 R.S. Johnson

1.6
1990 1991 1992 1993 1994
1.4

1.2

1.0
Kc

0.8

0.6

0.4

0.2

0
0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8
Proportion of midday light interception

Fig. 13.11. Mature peach tree crop coefficients (Kc) as a function of the proportion of available light
intercepted by the canopy at midday. Equation of the regression line is y = 1.59x + 0.082, R2 = 0.86.
(Adapted from Ayars et al., 2003 with permission of Springer Science Publishers, The Netherlands.)

1.4

1.2
Crop Coefficient (Kc)

1.0 y = 0.0087x – 0.4263

R2 = 0.9835
0.8

0.6

0.4

0.2

0.0
0 50 100 150 200 250 300
Day of year

Fig. 13.12. Seasonal pattern of crop coefficients (Kc) for mature peach trees (average from 2001 to
2003) as calculated from a weighing lysimeter. Each point is a weekly average. Regression line is for
data collected from bud break to early July (about day 180). Equation of the regression line is
y = 0.0087x – 0.4263, R2 = 0.9835. (Adapted from Johnson et al., 2005 with permission from
California Agriculture, Oakland, California, copyright 2005 U.C. Regents.)

In contrast, values for other deciduous trees
that are not pruned as heavily have generally
been reported to be much higher early in the
season (Allen et al., 1998). Some have reported
seasonal tree water use to follow the double-
sigmoid pattern of fruit growth (Chalmers
et al., 1983; Klein, 1983; Boland et al., 1993) but
this is not supported by weighing lysimeter
data. Both a late-season cultivar (Ayars et al.,
2003) and an early-season cultivar (Johnson
et al., 2005) had similar seasonal patterns of
water use that reflected canopy growth and not
fruit growth. A couple of approaches to model-
ling peach tree water use have been proposed.
One is based on light interception of the canopy
as it develops (Johnson et al., 2002, 2004).
 Nutrient and Water Requirements
The other approach is based on the stages of
fruit growth of various varieties (Klein, 1993).
Irrigation needs for young trees depend
strongly on the irrigation delivery system as
soil evaporation can become a major compo-
nent of ET. The transpiration component is
proportional to canopy light interception (Fig.
13.13), similar to mature trees (see Fig. 13.11).
However, the soil evaporation component
depends on how much of the soil surface is
wetted, and can be very large under flood or
furrow irrigation systems. A drip system can
save substantial amounts of applied water on
young trees (Fereres et al., 1982). Generally,
with efficient irrigation systems (drip or
microsprinklers), researchers have found
young tree ET to be about 50% greater than
the value for transpiration alone shown in
Fig. 13.13 (Black et al., 1977; Fereres et al., 1982;
Renquist, 1987). In other words, a canopy
shading 20% of the orchard floor would have
a Kc value of about 0.45 (0.30 + 50% = 0.45).
Water stress
Water stress can have many negative effects
on the growth and productivity of peach
trees. With severe stress, photosynthesis and
vegetative growth are greatly reduced, lead-
ing to diminished fruit production. Xylem
cavitation becomes increasingly severe as
water stress progresses. Ameglio et al. (1998)
reported 100% embolism at a xylem water
0.6
0.5
0.4
Kcb
0.3
0.2
0.1
0
0 5 10 15
Per cent light interception
Fig. 13.13. Crop coefficients of transpiration alone (no soil evaporation) or basal crop coefficients
(Kcb) of a 1-year-old peach tree as predicted by per cent light interception of the canopy at solar noon.
R2 of the regression line is 0.987. (Adapted from Johnson et al., 2002 with permission from the Interna-
tional Society of Horticultural Science, Belgium.)
321
potential of −3.0 MPa. In arid regions without
supplemental irrigation, peach trees tend to
die faster than other fruit trees unless they are
pruned severely to reduce leaf area and crop
load (Proebsting and Middleton, 1980). Heavy
crop loads have been shown to lead to greater
water stress (Berman and DeJong, 1996; Naor
et al., 1999) due to higher stomatal opening in
fruited versus non-fruited trees (DeJong,
1986). Fruit quality is also affected by water
stress as fruit size is reduced and abnormal
flower buds can develop, leading to such
disorders as fruit doubles and deep sutures
(Patten et al., 1989; Handley and Johnson, 2000;
Naor et al., 2005). Even normal-appearing
fruit are often astringent and lacking in red
coloration under severe water stress (Proebst-
ing and Middleton, 1980). Severe postharvest
water stress can reduce yield in the subse-
quent season (Naor et al., 2005).
If a more moderate water stress condi-
tion is imposed and carefully managed, some
beneficial results can be achieved. Since peach
trees are inherently very vigorous, heavy prun-
ing is required every year. Therefore, moderate
reductions in vegetative growth induced by
water stress would generally be considered
beneficial since they could reduce pruning and
improve light penetration though the canopy.
Furthermore, flowering and fruit set have
been reported to increase (Chalmers et al.,
1985; Larson et al., 1988) or remain unchanged
(Besset et al., 2001) with moderate water stress,
so continued productivity would not be a
problem. As stomatal conductance decreases
20 25 30 35
322 R.S. Johnson
with stress, some have reported no reduction
in carbon assimilation, thus leading to greater
water use efficiency (Cheng et al., 1996). Theo-
retically, under moderate stress, irrigation
water savings could be achieved without
reducing productivity. However, fruit fresh
weight is very sensitive to water stress, so it
can be reduced while fruit dry weight is not
affected (Girona et al., 2004). With a light or
moderate crop load, Berman and DeJong
(1996) found water stress reduced fruit fresh
weight but not dry weight. With a heavy crop
load, both were reduced. An increase in fruit
sugar concentration has generally been asso-
ciated with moderately stressed peach trees
(Gelly et al., 2004). Often this is simply a matter
of concentrating the sugars as fruit water con-
tent decreases. Thus, increased sugars come at
the expense of decreased fruit fresh weight
(Crisosto et al., 1994; Besset et al., 2001). How-
ever, it has been reported that greater sugar
concentration can occur without a loss of fruit
size (Kobashi et al., 1997). Kobashi et al. (2000)
documented an increase in sorbitol, sucrose
and total sugars with moderate but not severe
stress. Increased fruit growth could occur
under these conditions due to lower osmotic
and therefore greater turgor potential. This
appears to be the basis for a few reports of
improved fruit size with imposed stress, even
though most studies have shown decreased
fruit size with any level of water stress.
Strategies for imposing water stress
The potential benefits that can be achieved
with moderate water stress have led to a cou-
ple of strategies for imposing water stress in
commercial peach orchards. One approach is
to focus on the period after harvest of early-
maturing varieties. With no fruit on the tree,
the strategy is to save water while minimiz-
ing problems with productivity and fruit
quality in the following season and to prevent
permanent tree damage. A second approach
is to impose stress during the growth of the
fruit in such a way that no decrease (or even
an increase) in fruit size occurs. This strategy
has proved difficult to implement and may
only be successful under certain conditions.
Imposing water stress after harvest of an
early June-maturing peach cultivar has been
achieved for four successive years in California
(Johnson et al., 1992). The orchard was on a
deep sandy loam soil under flood irrigation.
By the end of the season, measures of water
status (stomatal conductance, leaf water
potential, trunk growth and diurnal shrink-
age) indicated substantial stress, but prema-
ture defoliation was minimal and subsequent
yields were not reduced (Larson et al., 1988).
Pruning weights were only slightly reduced
since the majority of vegetative growth occurs
early in the season. The biggest drawback to
this practice was a substantial increase in fruit
doubles the following year in the most severe
water-stressed treatment. By relieving late-
summer water stress during the carpel differ-
entiation period, fruit doubles and the
associated defect of deep sutures were reduced
to levels of the well-watered control (Handley
and Johnson, 2000; Naor et al., 2005). Thus, sig-
nificant irrigation water savings can be
achieved without sacrificing yield, fruit qual-
ity or tree health. The level of stress can become
severe enough to cause defoliation and reduce
yield in the following year (Naor et al., 2005). A
threshold of −2.0 MPa for midday stem water
potential in late summer (August in Israel
and California) has been proposed to mini-
mize productivity and fruit quality problems
(Naor et al., 2005; Naor, 2006).
The second approach to imposing water
stress has generally been referred to in the lit-
erature as regulated deficit irrigation (RDI). It
involves deficit irrigation during the slow
growth phase (or stage II) of a late-season
peach and also corresponds to the period of
rapid shoot growth. During the final, rapid
stage of fruit growth (stage III), full irrigation
(or greater) is applied through harvest. The
theory, which is not well substantiated, states
that the deficit irrigation will inhibit vegetative
growth to a greater extent than fruit growth.
Perhaps osmotic adjustment in the fruit will
also occur during this time. Then, once the
stress is alleviated during stage III, fruit will
grow at an increased rate due to less competi-
tion from shoots or greater turgor pressure
(Behboudian and Mills, 1997). Studies from
Australia reported positive results using RDI,
with fruit size increased by as much as 30%
compared with fully irrigated controls (Chal-
mers et al., 1981; Mitchell and Chalmers, 1982).
 Nutrient and Water Requirements 323

Other researchers have not had the same suc-
cess, as fruit size either was reduced by RDI
(Girona, 1989) or showed no significant increase
(Li et al., 1989; Strabbioli, 1992b; Boland et al.,
1993). Nevertheless, the water savings of 35
to 40% were substantial in these studies. Thus,
RDI offers a viable strategy for water conser-
vation, especially in locations where water is
scarce or very expensive.
The RDI strategy needs refinement before
it can be applied widely. There are still many
questions about the details of how to imple-
ment this approach under different conditions.
For example, the timing of stress imposition
and alleviation are not clear. Li et al. (1989)
have shown evidence that better results were
obtained by starting the stress earlier than
stage II. Also, alleviation of stress may need to
be started well ahead of stage III in situations
where roots are deep or water penetration into
the soil is poor. Girona et al. (1993) reported
that a period of 3 weeks after resumption of
full irrigation was needed before midday leaf
water potential recovered completely. Research
is also needed on the factors influencing
osmotic adjustment in the fruit. It appears the
degree of stress may have some influence
(Kobashi et al., 2000). The rate at which stress
develops may also be important (Natali et al.,
1985a; Olien and Flore, 1990). It has been sug-
gested that the success of RDI by researchers
in Australia may be due to the ability to
quickly impose and relieve stress under the
shallow soil conditions of their experiment
(Johnson and Handley, 2000).
Most of the RDI studies did not report
measurements of plant water status. This
makes it difficult to transfer information from
one location or soil type to another and expect
the same results. For future experiments it
will be important to assess the stress level
being experienced by the tree. The measure-
ment that has probably shown the best con-
sistency and correlation with tree performance
has been midday stem water potential (Garnier
and Berger, 1985; McCutchan and Shackel,
1992). Not only is it a good indicator of the
stress level in the tree, but it also correlates
well with midday stomatal closure (Marsal
and Girona, 1997), tree water use (Johnson
et al., 2005) and fruit size (Naor et al., 1999,
2001). It is proving to be a useful indicator of
stress in a wide range of fruit and nut trees
(Shackel et al., 1997; Naor, 2006).

References

Abadia, J., Nishio, J.N., Monge, E., Montañes, L. and Heras, L. (1985) Mineral composition of peach leaves
affected by iron chlorosis. Journal of Plant Nutrition 8, 965–975.
Abadia, J., Tagliavini, M., Grasa, R., Belkhodja, R., Abadia, A., Sanz, M., Faria, E.A., Tsipouridis, C. and
Marangoni, B. (2000) Using the flower Fe concentration for estimating chlorosis status in fruit tree or-
chards: a summary report. Journal of Plant Nutrition 23, 2023–2033.
Abdalla, D.A. and Childers, N.F. (1973) Calcium nutrition of peach and prune relative to growth, fruiting, and
fruit quality. Journal of the American Society for Horticultural Science 98, 517–522.
Adaskaveg, J.E., Ogawa, J.M. and Feliciano, A.J. (1992) Comparisons of calcium-based and film-forming
materials for control of brown rot of peach caused by Monilinia fructicola. Phytopathology 82, 1158.
Allen, R.A., Pereira, L.S., Raes, D. and Smith, M. (1998) Crop Evapotranspiration: Guidelines for Computing
Crop Water Requirements. FAO Irrigation and Drainage Paper No. 56. Food and Agriculture Organiza-
tion of the United Nations, Rome.
Alvino, A., Magliulo, V. and Zerbi, G. (1986) Problems of peach (Prunus persica) tolerance to anaerobic con-
ditions due to excess soil water. Rivista Ortoflorofrutticoltura Italiana 70, 263–270.
Ameglio, T., Cochard, H., Picon, C. and Cohen, M. (1998) Water relations and hydraulic architecture of peach
trees under drought conditions. Acta Horticulturae 465, 355–362.
Andersen, P.C., Lombard, P.B. and Westwood, M.N. (1984) Leaf conductance, growth, and survival of willow
and deciduous fruit tree species under flooded soil conditions. Journal of the American Society for
Horticultural Science 109, 132–138.
Anderssen, F.G. (1932) Chlorosis of deciduous fruit trees due to a copper deficiency. Journal of Pomology and
Horticulture Science 10, 130–146.
324 R.S. Johnson

Arce, J.P., Storey, J.B. and Lyons, C.G. (1992) Effectiveness of three different zinc fertilizers and two methods
of application for the control of ‘Little-Leaf’ in peach trees in south Texas. Communications in Soil Science
and Plant Analyses 23, 1945–1962.
Ayars, J.E., Johnson, R.S., Phene, C.J., Trout, T.J., Clark, D.A. and Mead, R.M. (2003) Water use by drip
irrigated late season peaches. Irrigation Science 22, 187–194.
Ballinger, W.E., Bell, H.K. and Childers, N.F. (1966) Peach nutrition. In: Childers, N.F. (ed.) Nutrition of Fruit
Crops, Temperate, Sub-tropical, Tropical. Somerset Press, Sommerville, New Jersey, pp. 276–390.
Batjer, L.P. and Westwood, M.N. (1958) Seasonal trend of several nutrient elements in leaves and fruits of
Elberta peach. Proceedings of the American Society for Horticultural Science 71, 116–126.
Behboudian, M.H. and Mills, T.M. (1997) Deficit irrigation in deciduous orchards. Horticultural Reviews 21,
105–131.
Bell, H.K. and Childers, N.F. (1954) Peach nutrition. In: Childers, N.F. (ed.) Fruit Nutrition. Somerset Press,
Sommerville, New Jersey, pp. 495–641.
Bell, R.W., Dell, B. and Huang, L. (2002) Boron requirement of plants. In: Goldbach, H.E., Rerkasem, B.,
Wimmer, M.A., Brown, P.H., Thellier, M. and Bell, R.W. (eds) Boron in Plant and Animal Nutrition.
Kluwer Academic/Plenum Publishers, New York, pp. 63–85.
Benson, N.R., Degman, E.S., Chmelir, I.C. and Chennault, W. (1963) Sulfur deficiency in deciduous tree fruits.
Proceedings of the American Society for Horticultural Science 83, 55–62.
Berman, M.E. and DeJong, T.M. (1996) Water stress and crop load effects on fruit fresh and dry weights in
peach (Prunus persica). Tree Physiology 16, 859–864.
Besset, J., Genard, M., Girard, T., Serra, V. and Bussi, C. (2001) Effect of water stress applied during the final
stage of rapid growth on peach trees (cv. Big-Top). Scientia Horticulturae 91, 289–303.
Beyers, E. and Terblanche, J.H. (1971a) Identification and control of trace element deficiencies. II. Manganese
deficiency and toxicity. The Deciduous Fruit Grower 21, 167–171.
Beyers, E. and Terblanche, J.H. (1971b) Identification and control of trace element deficiencies. III. Copper
deficiency. The Deciduous Fruit Grower 21, 199–202.
Beyers, E. and Terblanche, J.H. (1971c) Identification and control of trace element deficiencies. V. Iron
deficiency. The Deciduous Fruit Grower 21, 265–268.
Beyers, E. and Terblanche, J.H. (1971d) Identification and control of trace element deficiencies. VI. Magnesium
deficiency. The Deciduous Fruit Grower 21, 305–309.
Bhullar, J.S., Dhillon, B.S. and Randhawa, J.S. (1981) Effect of pre-harvest calcium nitrate sprays on the ambient
storage of Flordasun peach fruits. Journal of Research Punjab Agricultural University 18, 282–286.
Biggs, A.R., El-Kholi, M.M., El-Neshawy, S. and Nickerson, R. (1997) Effects of calcium salts on growth, poly-
galacturonase activity, and infection of peach fruit by Monilinia fructicola. Plant Disease 81, 399–403.
Black, J.D.F., Mitchell, P.D. and Newgreen, P.N. (1977) Optimum irrigation rates for young trickle irrigated
peach trees. Australian Journal of Experimental Agriculture and Animal Husbandry 17, 342–345.
Boland, A.-M., Mitchell, P.D., Jerie, P.H. and Goodwin, I. (1993) The effect of regulated deficit irrigation on
tree water use and growth of peach. Journal of Horticultural Science 68, 261–274.
Bollard, E.G. (1953) Zinc deficiency in peaches and nectarines. New Zealand Journal of Science and Technology
35A, 15–18.
Boynton, D. (1944) Responses of young Elberta peach and Montmorency cherry trees to potassium fertilization
in New York. Proceedings of the American Society for Horticultural Science 44, 31–33.
Boynton, D., Krochmal, A. and Konecny, J. (1951) Leaf and soil analyses for manganese in relation to interveinal
leaf chlorosis in some sour cherry, peach and apple trees of New York. Proceedings of the American Society
for Horticultural Science 57, 1–8.
Brown, P.H. and Hu, H. (1996) Phloem mobility of boron is species dependent: evidence for phloem mobility
in sorbitol-rich species. Annals of Botany 77, 497–505.
Byrne, D.H. (1988) Comparative growth of two peach seedling rootstocks under alkaline soil conditions.
Journal of Plant Nutrition 11, 1663–1669.
Chalmers, D.J., Mitchell, P.D. and van Heek, L. (1981) Control of peach tree growth and productivity by regu-
lated water supply, tree density, and summer pruning. Journal of the American Society for Horticultural
Science 106, 307–312.
Chalmers, D.J., Olsson, K.A. and Jones, T.R. (1983) Water relations of peach trees and orchards. In: Kozlowski,
T.T. (ed.) Water Deficits and Plant Growth. Academic Press, New York, pp. 197–232.
Chalmers, D.J., Mitchell, P.D. and Jerie, P.H. (1985) The relation between irrigation, growth and productivity
of peach trees. Acta Horticulturae 173, 283–288.
Chandler, W.H. (1937) Zinc as a nutrient for plants. Botanical Gazette 98, 625–646.
 Nutrient and Water Requirements 325

Chandler, W.H., Hoagland, D.R. and Hibbard, P.L. (1931) Little-leaf or rosette in fruit trees. Proceedings of the
American Society for Horticultural Science 28, 556–560.
Cheng, L., Cheng, S., Shu, H. and Luo, X. (1996) Effect of mild water stress on CO2 assimilation and water use
efficiency of field-grown peach trees. Acta Horticulturae 374, 121–125.
Cibes, H.R., Hernandez, E. and Childers, N.F. (1955) Boron toxicity induced in a New Jersey peach orchard.
Part I. Proceedings of the American Society for Horticultural Science 66, 13–20.
Claypool, L.L. (1975) Plant nutrition and deciduous fruit crop quality. HortScience 10, 45–47.
Conway, W.S. (1982) Effect of postharvest calcium treatment on decay of ‘Delicious’ apples. Plant Disease 66,
402–403.
Conway, W.S., Greene, G.M. II and Hickey, K.D. (1987) Effects of preharvest and postharvest calcium treat-
ments of peaches on decay caused by Monilinia fructicola. Plant Disease 71, 1084–1086.
Crisosto, C.H., Johnson, R.S., Luza, J.G. and Crisosto, G.M. (1994) Irrigation regimes affect fruit soluble solids
concentration and rate of water loss of ‘O’Henry’ peaches. HortScience 29, 1169–1171.
Crisosto, C.H., Day, K.R., Johnson, R.S. and Garner, D. (2000) Influence of in-season foliar calcium sprays on
fruit quality and surface discoloration incidence of peaches and nectarines. Journal of the American
Pomological Society 54, 118–122.
Cullinan, F.P. and Batjer, L.P. (1943) Nitrogen, phosphorus, and potassium inter-relationships in young peach
and apple trees. Soil Science 55, 49–60.
Cummings, G.A. (1965) Effect of potassium and magnesium fertilization on the yield, size, maturity,
and color of Elberta peaches. Proceedings of the American Society for Horticultural Science 85,
133–140.
Cummings, G.A. and Reeves, J. (1971) Factors influencing chemical characteristics of peaches. Journal of the
American Society for Horticultural Science 96, 320–322.
Daane, K.M., Johnson, R.S., Michailides, T.J., Crisosto, C.H., Dlott, J.W., Ramirez, H.T., Yokota, G.Y. and
Morgan, D.P. (1995) Excess nitrogen raises nectarine susceptibility to disease and insects. California
Agriculture 49, 13–18.
Daines, R.H., Cohoon, D.F., Leone, I. and Brennan, E. (1958) Control of fusicoccum canker of peach by nutrition,
defoliation, and protective fungicides. Phytopathology 48, 400–407.
Daniell, J.W. (1982) Effect of trickle irrigation on the growth and yield of ‘Loring’ peach trees. Journal of
Horticultural Science 57, 393–399.
DeJong, T.M. (1986) Fruit effects on photosynthesis in Prunus persica. Physiologia Plantarum 66, 149–153.
Dunbar, C.O. and Anthony, R.D. (1937) Two cases of potassium deficiency in peach orchards in South Central
Pennsylvania. Proceedings of the American Society for Horticultural Science 35, 320–325.
Dye, M.H., Buchanan, L., Dorofaeff, F.D. and Beecroft, F.G. (1984) Boron toxicity in peach and nectarine trees
in Otago. New Zealand Journal of Experimental Agriculture 12, 303–313.
Edwards, J.H. and Horton, B.D. (1979) Response of peach seedlings to calcium concentration in nutrient solu-
tion. Journal of the American Society for Horticultural Science 104, 97–99.
Epstein, E. and Lilleland, O. (1942) A preliminary study of the manganese content of the leaves of some
deciduous fruit trees. Proceedings of the American Society for Horticultural Science 41, 11–18.
Epstein, L. and Bassein, S. (2001) Pesticide applications of copper on perennial crops in California, 1993 to
1998. Journal of Environmental Quality 30, 1844–1847.
Faust, M. and Timon, B. (1995) Origin and dissemination of peach. Horticultural Reviews 17, 331–379.
Feldstein, J. and Childers, N.F. (1965) Effects of irrigation on peaches in Pennsylvania. Proceedings of the
American Society for Horticultural Science 87, 145–153.
Fereres, E., Martinich, D.A., Aldrich, T.M., Castel, J.R., Holzapfel, E. and Schulbach, H. (1982) Drip irrigation
saves money in young almond orchards. California Agriculture 36, 12–13.
Finch, C.R., Byrne, D.H., Lyons, C.G. and Pennington, H.D. (1997) Sulfur nutrition requirements of peach
trees. Journal of Plant Nutrition 20, 1711–1721.
Furuya, S. and Umemiya, Y. (2002) The influence of chemical forms on foliar-applied nitrogen absorption for
peach trees. Acta Horticulturae 594, 97–103.
Garnier, E. and Berger, A. (1985) Testing water potential in peach trees as an indicator of water stress. Journal
Horticultural Science 60, 47–56.
Garnier, E., Berger, A. and Rambal, S. (1986) Water balance and pattern of soil water uptake in a peach
orchard. Agricultural Water Management 11, 145–158.
Gelly, M., Recasens, I., Girona, J., Mata, M., Arbones, A., Rufat, J. and Marsal, J. (2004) Effects of stage II and
postharvest deficit irrigation on peach quality during maturation and after cold storage. Journal of the
Science of Food and Agriculture 84, 561–568.
326 R.S. Johnson

Gilmore, A.E. (1971) The influence of endotrophic mycorrhizae on the growth of peach seedlings. Journal of
the American Society for Horticultural Science 96, 35–37.
Gil Salaya, G.F. (2000) FrutiCultura la produccion de fruta. Fruta de climas templado y subtropical y uva de
vino. Coleccion En Agricultura, Facultad De Agronomia E Ingenieria Forestal. Ediciones Universidad
Catolica de Chile, Santiago.
Girona, J. (1989) Physiological, growth and production responses of late maturing peach (Prunus persica L.
Batsch) to controlled deficit irrigation. MS thesis, University of California, Davis, California.
Girona, J., Mata, M., Goldhamer, D.A., Johnson, R.S. and DeJong, T.M. (1993) Patterns of soil and tree water
status and leaf functioning during regulated deficit irrigation scheduling in peach. Journal of the Ameri-
can Society for Horticultural Science 118, 580–586.
Girona, J., Marsal, J., Mata, M., Arbones, A. and DeJong, T.M. (2004) A comparison of the combined effect of
water stress and crop load on fruit growth during different phenology stages in young peach trees. Jour-
nal of Horticultural Science & Biotechnology 79, 308–315.
Goodwin, I., Whitfield, D.M. and Connor, D.J. (2006) Effects of tree size on water use of peach (Prunus per-
sica L. Batsch). Irrigation Science 24, 59–68.
Gupta, U.C., Jame, Y.W., Campbell, C.A., Leyshon, A.J. and Nicholaichuk, W. (1985) Boron toxicity and defi-
ciency: a review. Canadian Journal of Soil Science 65, 381–409.
Haas, A.R.C. (1929) Toxic effect of boron on fruit trees. The Botanical Gazette 88, 113–131.
Handley, D.F. and Johnson, R.S. (2000) Late summer irrigation of water-stressed peach trees reduces fruit
doubles and deep sutures. HortScience 35, 771.
Hernandez, E. and Childers, N.F. (1956) Boron toxicity induced in a New Jersey peach orchard. Part II. Pro-
ceedings of the American Society for Horticultural Science 67, 121–129.
Horton, B.D., Wehunt, E.J., Edwards, J.H., Bruce, R.R. and Chesnee, J.L. (1981) The effects of drip irrigation
and soil fumigation on ‘Redglobe’ peach yields and growth. Journal of the American Society for Horti-
cultural Science 106, 438–443.
Hu, H., Penn, S.G., Lebrilla, C.B. and Brown, P.H. (1997) Isolation and characterization of soluble boron
complexes in higher plants. The mechanism of phloem mobility of boron. Plant Physiology 113, 649–
655.
Igartua, E., Grasa, R., Sanz, M., Abadia, A. and Abadia, J. (2000) Prognosis of iron chlorosis from the mineral
composition of flowers in peach. Journal of Horticultural Science & Biotechnology 75, 111–118.
Jafar, H. (1958) Investigation on the control of peach rust (Tranzschelia pruni-spinosae Pers.). New Zealand
Journal of Agricultural Research 1, 660–664.
Johnson, R.S. (1993) Stone fruit: peaches and nectarines. In: Bennett, W.F. (ed.) Nutrient Deficiencies and
Toxicities in Crop Plants. APS Press, St. Paul, Minnesota, pp. 171–176.
Johnson, R.S. and Andris, H.L. (2001) Combining low biuret urea with foliar zinc sulfate sprays to fertilize
peach and nectarine trees in the fall. Acta Horticulturae 564, 321–327.
Johnson, R.S. and Handley, D.F. (2000) Using water stress to control vegetative growth and productivity of
temperate fruit trees. HortScience 35, 1048–1050.
Johnson, R.S. and Uriu, K. (1989) Mineral nutrition. In: LaRue, J.H. and Johnson, R.S. (eds) Peaches, Plums,
and Nectarines: Growing and Handling for Fresh Market. University of California Division of Agricul-
ture and Natural Resources, Publication No. 3331. University of California, Oakland, California, pp.
68–81.
Johnson, R.S., Handley, D.F. and DeJong, T.M. (1992) Long-term response of early maturing peach trees to
postharvest water deficits. Journal of the American Society for Horticultural Science 117, 881–886.
Johnson, R.S., Ayars, J., Trout, T., Mead, R. and Phene, C. (2000) Crop coefficients for mature peach trees are
well correlated with midday canopy light interception. Acta Horticulturae 537, 455–460.
Johnson, R.S., Rosecrance, R., Weinbaum, S., Andris, H. and Wang, J. (2001) Can we approach complete
dependence on foliar-applied urea nitrogen in an early-maturing peach? Journal of the American Society
for Horticultural Science 126, 364–370.
Johnson, R.S., Ayars, J. and Hsiao, T. (2002) Modeling young peach tree evapotranspiration. Acta Horticultu-
rae 584, 107–113.
Johnson, R.S., Ayars, J. and Hsiao, T. (2004) Improving a model for predicting peach tree evapotranspiration.
Acta Horticulturae 664, 341–346.
Johnson, R.S., Williams, L.E., Ayars, J.E. and Trout, T.J. (2005) Weighing lysimeters for studying tree and vine
water relations. California Agriculture 59, 133–136.
Johnson, R.S., Andris, H., Day, K. and Beede, B. (2006) Using dormant shoots to determine the nutritional
status of peach trees. Acta Horticulturae 721, 285–290.
 Nutrient and Water Requirements 327

Kamali, A.R. and Childers, N.F. (1970) Growth and fruiting of peach in sand culture as affected by boron and
a fritted form of trace elements. Journal of the American Society for Horticultural Science 95, 652–656.
Kester, D.E. and Asay, R.N. (1986) ‘Hansen 2168’ and ‘Hansen 536’: two new Prunus rootstock clones. Hort-
Science 21, 331–332.
Klein, I. (1983) Drip irrigation based on soil matric potential conserves water in peach and grape. HortScience
18, 942–944.
Klein, I. (1993) Irrigation modeling of peach and nectarine. Acta Horticulturae 349, 225–228.
Kobashi, K., Gemma, H. and Iwahori, S. (1997) Effect of water stress on fruit quality and endogenous abscisic
acid (ABA) content in peach fruit. Environmental Control Biology 35, 267–274.
Kobashi, K., Gemma, H. and Iwahore, S. (2000) Abscisic acid content and sugar metabolism of peaches
grown under water stress. Journal of the American Society for Horticultural Science 125, 425–428.
Köseoglu, A.T. (1995) Investigations of relationships between iron status of peach leaves and soil properties.
Journal of Plant Nutrition 18, 1845–1859.
Kwong, S.S. and Fisher, E.G. (1962) Potassium effects on titratable acidity and the soluble nitrogenous compounds
of ‘Jerseyland’ peach. Proceedings of the American Society for Horticultural Science 81, 168–171.
Larson, K.D., DeJong, T.M. and Johnson, R.S. (1988) Physiological and growth responses of mature peach trees
to postharvest water stress. Journal of the American Society for Horticultural Science 113, 296–300.
Layne, R.E.C. and Tan, C.S. (1984) Long-term influence of irrigation and tree density on growth, survival and
production of peach. Journal of the American Society for Horticultural Science 109, 795–799.
Layne, R.E.C., Tan, C.S. and Fulton, J.M. (1981) Effect of irrigation and tree density on peach production.
Journal of the American Society for Horticultural Science 106, 151–156.
Layne, R.E.C., Tan, C.S. and Hunter, D.M. (1994) Cultivar, ground-cover, and irrigation treatments and their
interactions affect long-term performance of peach trees. Journal of the American Society for Horticul-
tural Science 119, 12–19.
Layne, R.E.C., Tan, C.S. and Hunter, D.M. (1996) Irrigation and fertilizer application methods affect perfor-
mance of high-density peach orchards. HortScience 31, 370–375.
Leece, D.R. and Kenworthy, A.L. (1971) Effect of potassium nitrate foliar sprays on leaf nitrogen concentration
and growth of peach trees. HortScience 6, 171–173.
Leece, D.R., Cradock, F.W. and Carter, O.G. (1971) Development of leaf nutrient concentration standards for
peach trees in New South Wales. Journal of Horticultural Science 46, 163–175.
Li, S.-H., Huguet, J.-G., Schoch, P.G. and Orlando, P. (1989) Response of peach tree growth and cropping to
soil water deficit at various phenological stages of fruit development. Journal of Horticultural Science 64,
541–552.
Lilleland, O. (1932) Experiments in K and P deficiencies with fruit trees in the field. Proceedings of the
American Society for Horticultural Science 29, 272–276.
Lilleland, O. and Brown, J.G. (1940) The phosphate nutrition of fruit trees. II. Continued response to phosphate
applied at the time of planting. Proceedings of the American Society for Horticultural Science 37, 53–57.
Lilleland, O. and Brown, J.G. (1942) The phosphate nutrition of fruit trees. IV. The phosphate content of peach
leaves from 130 orchards in California and some factors which may influence it. Proceedings of the
American Society for Horticultural Science 41, 1–10.
Lilleland, O., Brown, J.G. and Conrad, J. P. (1942) The phosphate nutrition of fruit trees. III. Comparison of fruit
tree and field crop responses on a phosphate deficient soil. Proceedings of the American Society for
Horticultural Science 40, 1–7.
Lilleland, O., Uriu, K., Muroaka, T. and Pearson, J. (1962) The relationship of potassium in the peach leaf to
fruit growth and size at harvest. Proceedings of the American Society for Horticultural Science 81,
162–167.
Lobit, P., Soing, S., Genard, M. and Habib, R. (2001) Effects of timing of nitrogen fertilization on shoot devel-
opment in peach (Prunus persica) trees. Tree Physiology 20, 35–42.
McClung, A.C. (1953) Magnesium deficiency in North Carolina peach orchards. Proceedings of the American
Society for Horticultural Science 62, 123–130.
McClung, A.C. (1954) The occurrence and correction of zinc deficiency in North Carolina peach orchards.
Proceedings of the American Society for Horticultural Science 64, 75–80.
McClung, A.C. and Clayton, C.N. (1956) Boron in relation to foliar and fruiting abnormalities of peach. Plant
Disease Reporter 40, 542–548.
McCutchan, H. and Shackel, K.A. (1992) Stem-water potential as a sensitive indicator of water stress in prune
trees (Prunus domestica L. cv. French). Journal of the American Society for Horticultural Science 117,
607–611.
328 R.S. Johnson

McLarty, H.R. and Woodbridge, C.G. (1950) Boron in relation to the culture of the peach tree. Scientific
Agriculture 30, 392–395.
Mann, M.S., Sidhu, B.S., Chahil, B.S. and Mann, S.S. (1986) Effect of different rates of zinc applied to soil and
foliage of peach (Prunus persica Batsch) on zinc concentration in leaves, fruit yield and quality. In:
Chadha, T.R., Bhutani, V.P. and Kaul, J.L. (eds) Advances in Research on Temperate Fruits. Dr Y.S. Parmar
University of Horticulture and Forestry, Solan, India, pp. 189–191.
Marsal, J. and Girona, J. (1997) Relationship between leaf water potential and gas exchange activity at differ-
ent phenological stages and fruit loads in peach trees. Journal of the American Society for Horticultural
Science 122, 415–421.
Mitchell, P.D. and Chalmers, D.J. (1982) The effect of reduced water supply on peach tree growth and yields.
Journal of the American Society for Horticultural Science 107, 853–856.
Monge, E., Montañes, L., Val, J. and Sanz, M. (1995) A comparative study of the DOP and the DRIS methods,
for evaluating the nutritional status of peach trees. Acta Horticulturae 383, 191–199.
Montañes, L. and Sanz, M. (1994) Prediction of reference values for early leaf analysis for peach trees. Journal
of Plant Nutrition 17, 1647–1657.
Montañes, L., Herar, L., Abadia, J. and Sanz, M. (1993) Plant analysis interpretation based on a new index:
deviation from optimum percentage (DOP). Journal of Plant Nutrition 16, 1289–1308.
Morales, F., Grasa, R., Abadia, A. and Abadia, J. (1998) Iron chlorosis paradox in fruit trees. Journal of Plant
Nutrition 21, 815–825.
Naor, A. (2006) Irrigation scheduling and evaluation of tree water status in deciduous orchards. Horticultural
Reviews 32, 111–165.
Naor, A., Klein, I., Hupert, H., Grinblat, Y., Peres, M. and Kaufman, A. (1999) Water stress and crop level in-
teractions in relation to nectarine yield, fruit size distribution, and water potentials. Journal of the Amer-
ican Society for Horticultural Science 124, 189–193.
Naor, A., Hupert, H., Greenblat, Y., Peres, M., Kaufman, A. and Klein, I. (2001) The response of nectarine fruit
size and midday stem water potential to irrigation level in stage III and crop load. Journal of the Ameri-
can Society for Horticultural Science 126, 140–143.
Naor, A., Stern, R., Peres, M., Greenblat, Y., Gal, Y. and Flaishman, M. (2005) Timing and severity of posthar-
vest water stress affect following-year productivity and fruit quality of field-grown ‘Snow Queen’ nectar-
ine. Journal of the American Society for Horticultural Science 130, 806–812.
Natali, S., Xiloyannis, C. and Pezzarossa, B. (1985a) Relationship between soil water content, leaf water potential
and fruit growth during different fruit growing phases of peach trees. Acta Horticulturae 171, 167–180.
Natali, S., Xiloyannis, C. and Mugano, M. (1985b) Water consumption in high density peach trees. Acta
Horticulturae 173, 413–420.
Neilsen, G.H. and Neilsen, D. (1994) Tree fruit zinc nutrition In: Peterson, A.B. and Stevens, R.G. (eds) Tree
Fruit Nutrition (Short Course Proceedings). Good Fruit Grower, Washington State Fruit Commission,
Yakima, Washington, pp. 85–93.
Neilsen, G.H., Yorstan, J., van Lierop, W. and Hoyt, P.B. (1985) Relationships between leaf and soil boron and
boron toxicity of peaches in British Columbia. Canadian Journal of Soil Science 65, 213–217.
Neilsen, G.H., Hogue, E.J. and Yorston, J. (1990) Response of fruit trees to phosphorus fertilization. Acta
Horticulturae 274, 347–359.
Neilsen, G.H., Neilsen, D. and Peryea, F. (1999) Response of soil and irrigated fruit trees to fertigation or
broadcast application of nitrogen, phosphorus, and potassium. HortTechnology 9, 393–401.
Norton, R.A. and Childers, N.F. (1954) Experiments with urea sprays on the peach. Proceedings of the American
Society for Horticultural Science 63, 23–31.
Nyomora, A.M.S., Brown, P.H. and Freeman, M. (1997) Fall foliar-applied boron increases tissue boron concen-
tration and nut set of almond. Journal of the American Society for Horticultural Science 122, 405–410.
Olien, M.E. and Flore, J.A. (1990) Effect of a rapid water stress and a slow water stress on the growth of ‘Red-
haven’ peach trees. Fruit Varieties Journal 44, 4–11.
Patten, K., Nimr, G. and Neuendorff, E. (1989) Fruit doubling of peaches as affected by water stress. Acta
Horticulturae 254, 319–321.
Phene, C.J., Hoffman, G.J., Howell, T.A., Clark, D.A., Mead, R.M., Johnson, R.S. and Williams, L.E. (1991)
Automated lysimeter for irrigation and drainage control. In: Lysimeters for Evapotranspiration and Envi-
ronmental Measurements. IR Div/ASCE, Honolulu, Hawaii, pp. 28–36.
Policarpo, M., Di Marco, L., Caruso, T., Gioacchini, P. and Tagliavini, M. (2002) Dynamics of nitrogen uptake
and partitioning in early and late fruit ripening peach (Prunus persica) tree genotypes under a Mediter-
ranean climate. Plant and Soil 239, 207–214.
 Nutrient and Water Requirements 329

Pooviah, B.W., Glenn, G.M. and Reddy, A.S.N. (1988) Calcium and fruit softening: physiology and biochem-
istry. Horticultural Reviews 10, 107–152.
Powell, J.C., Lyons, C.G. and Haby, V.A. (1995) Effects of copper, zinc, and sulfur application to peach trees
on coastal plain soil. Communications in Soil Science and Plant Analysis 26, 1637–1648.
Proebsting, E.L. and Kinman, C.F. (1933) Orchard trials of nitrogen and phosphorus. Proceedings of the American
Society for Horticultural Science 30, 426–430.
Proebsting, E.L. Jr and Middleton, J.E. (1980) The behavior of peach and pear trees under extreme drought
stress. Journal of the American Society for Horticultural Science 105, 380–385.
Proebsting, E.L. Jr, Carter, G.H., Ingalsbe, D.W. and Neubert, A.M. (1957) Relationship between leaf nitrogen and can-
ning quality of Elberta peaches. Proceedings of the American Society for Horticultural Science 69, 131–140.
Rashid, A., Couvillon, G.A. and Jones, J.B. (1990) Assessment of Fe status of peach rootstocks by techniques
used to distinguish chlorotic and non-chlorotic leaves. Journal of Plant Nutrition 13, 285–307.
Razeto, B. (1982) Treatments for iron chlorosis in peach trees. Journal of Plant Nutrition 5, 917–922.
Reed, D.W., Lyons, C.G. Jr and McEachern, G.R. (1988) Field evaluation of inorganic and chelated iron fertil-
izers as foliar sprays and soil application. Journal of Plant Nutrition 11, 1369–1378.
Reeder, B.D., Newman, J.S. and Worthington, J.E. (1979) Effect of trickle irrigation on peach trees. Hort-
Science 14, 36–37.
Renquist, R. (1987) Evapotranspiration calculations for young peach trees and growth responses to irrigation
amount and frequency. HortScience 22, 221–223.
Robinson, J.B., Treeby, M.T. and Stephenson, R.A. (1997) Fruits, vines and nuts. In: Reuter, D.J. and Robinson,
J.B. (eds) Plant Analysis, An Interpretation Manual, 2nd edn. CSIRO Publishing, Collingwood, Australia,
pp. 349–382.
Robson, M.G., Hopfinger, J.A. and Eck, P. (1989) Postharvest sensory evaluation of calcium treated peach fruit.
Acta Horticulturae 254, 173–176.
Rogers, E., Johnson, G. and Johnson, D. (1974) Iron-induced manganese deficiency in ‘Sungold’ peach and its effects
on fruit composition and quality. Journal of the American Society for Horticultural Science 99, 242–244.
Rombola, A.D., Quartieri, M., Tagliavini, M., Iannone, C., Zavalloni, C. and Marangoni, B. (1995) Slow-
release N fertilizers and foliar application of urea in the peach orchard. In: Atti del XXII Convegno Pe-
schicolo. Società Orticola Italiana, Cesena, pp. 195–201.
Rosecrance, R.C., Johnson, R.S. and Weinbaum, S.A. (1998a) The effect of timing of post-harvest foliar urea
sprays on nitrogen absorption and partitioning in peach and nectarine trees. Journal of Horticultural
Science & Biotechnology 73, 856–861.
Rosecrance, R.C., Johnson, R.S. and Weinbaum, S.A. (1998b) Foliar uptake of urea-N by nectarine leaves: a
reassessment. HortScience 33, 158.
Rufat, J. and DeJong, T.M. (2001) Estimating seasonal nitrogen dynamics in peach trees in response to nitrogen
availability. Tree Physiology 21, 1133–1140.
Sánchez, E.E. (1999) Nutricion mineral de frutales de pepita y carozo. Publicacion del Instituto Nacional de
Tecnologia Agropecuaria. Estacion Experimental Alto Valle de Rio Negro, Macrorregion Patagonia Norte,
Argentina.
Sánchez, E.E. and Righetti, T.L. (2002) Misleading zinc deficiency diagnosis in pome fruit and inappropriate
use of foliar zinc sprays. Acta Horticulturae 594, 363–368.
Sánchez, E.E., Weinbaum, S.A. and Johnson, R.S. (2006) Comparative movement of labeled nitrogen and zinc
in 1-year-old peach [Prunus persica (L.) Batsch] trees following late-season foliar application. Journal of
Horticultural Science & Biotechnology 81, 839–844.
Sanz, M. (1999) Evaluation of interpretation of DRIS system during growing season of the peach tree: com-
parison with DOP method. Communications in Soil Science and Plant Analysis 30, 1025–1036.
Sanz, M. and Montañes, L. (1995) Flower analysis as a new approach to diagnosing the nutritional status of
the peach tree. Journal of Plant Nutrition 18, 1667–1675.
Sanz, M., Cavero, J. and Abadia, J. (1992a) Iron chlorosis in the Ebro River Basin, Spain. Journal of Plant Nutri-
tion 15, 1971–1981.
Sanz, M., Heras, L. and Montañes, L. (1992b) Relationships between yield and leaf nutrient contents in peach
trees: early nutritional status diagnosis. Journal of Plant Nutrition 15, 1457–1466.
Sanz, M., Val, J., Monge, E. and Montañes, L. (1995) Is it possible to diagnose the nutritional status of peach
trees by chemical analysis of their flowers? Acta Horticulturae 383, 159–163.
Sanz, M., Belkhodja, R., Toselli, M., Montañes, L., Abadia, A., Tagliavini, M., Marangoni, B. and Abadia, J.
(1997a) Floral analysis as a possible tool for the prognosis of iron deficiency in peach. Acta Horticultu-
rae 448, 241–245.
330 R.S. Johnson

Sanz, M., Pascual, J. and Machin, J. (1997b) Prognosis and correction of iron chlorosis in peach trees: influ-
ence on fruit quality. Journal of Plant Nutrition 20, 1567–1572.
Schaffer, B., Andersen, P.C. and Ploetz, R.C. (1992) Responses of fruit crops to flooding. Horticultural Reviews
13, 257–301.
Scott, L.E. (1939) Response of peach trees to potassium and phosphorus fertilizers in the Sandhill Area of the
Southeast. Proceedings of the American Society for Horticultural Science 36, 56–60.
Shackel, K.A., Ahmadi, H., Biasi, W., Buchner, R., Goldhamer, D., Gurusinghe, S., Hasey, J., Kester, D., Krue-
ger, B., Lampinen, B., McGourty, G., Micke, W., Mitcham, E., Olson, B., Pelletrau, K., Philips, H., Ramos,
D., Schwankl, L., Sibbett, S., Southwick, S., Stevenson, M., Thorpe, M., Weinbaum, S. and Yeager, J. (1997)
Plant water status as an index of irrigation need in deciduous fruit trees. HortTechnology 7, 23–29.
Shear, C.B. (1975) Calcium-related disorders of fruits and vegetables. HortScience 10, 361–365.
Shear, C.B. and Faust, M. (1980) Nutritional ranges in deciduous tree fruits and nuts. Horticultural Reviews 2,
142–163.
Shorrocks, V.M. (1997) The occurrence and correction of boron deficiency. Plant and Soil 193, 121–148.
Shu, Z.-H., Oberly, G.H. and Cary, E.E. (1993) Time course study on the mobility and pattern of distribution
of foliar-applied boron in peaches. Journal of Plant Nutrition 16, 1661–1673.
Shu, Z.-H., Oberly, G.H., Cary, E.E. and Rutzke, M. (1994) Absorption and translocation of boron applied to
aerial tissues of fruiting ‘Reliance’ peach trees. HortScience 29, 25–27.
Shu, Z.-H., Oberly, G.H. and Cary, E.E. (1997) Absorption, movement and distribution of boron applied to
peach (Prunus persica L. Batsch) fruits. In: Bell, R.W. and Rerkasem, B. (eds) Boron in Soils and Plants.
Kluwer Academic Publishers, Dordrecht, The Netherlands, pp. 209–212.
Smith, M.W., Kenworthy, A.L. and Bedford, C.L. (1979) The response of fruit trees to injection of nitrogen through
a trickle irrigation system. Journal of the American Society for Horticultural Science 104, 311–313.
Strabbioli, G. (1992a) Peach water requirements in Central Valley. Acta Horticulturae 315, 203–210.
Strabbioli, G. (1992b) The influence of regulated deficit irrigation (RDI) on the growth and productivity of
peach trees. Acta Horticulturae 315, 211–217.
Stylinanides, D.C., Tsipourides, C.G. and Michailides, Z.S. (1989) Resistance to iron deficiency of five peach
rootstocks. Acta Horticulturae 254, 185–187.
Swietlik, D. (1999) Zinc nutrition in horticultural crops. Horticultural Reviews 23, 110–178.
Swietlik, D. (2002a) Zinc nutrition of fruit crops. HortTechnology 12, 45–50.
Swietlik, D. (2002b) Zinc nutrition of fruit trees by foliar sprays. Acta Horticulturae 594, 123–129.
Swietlik, D. and Faust, M. (1984) Foliar nutrition of fruit crops. Horticultural Reviews 6, 287–355.
Syrgiannidis, G. (1985) Control of iron chlorosis and replant diseases in peach by using the GF 677 rootstock.
Acta Horticulturae 173, 383–388.
Tagliavini, M. and Marangoni, B. (2002) Major nutritional issues in deciduous fruit orchards of Northern Italy.
HortTechnology 12, 26–31.
Tagliavini, M., Scudellari, D., Marangoni, B. and Toselli, M. (1996) Nitrogen fertilization management in or-
chards to reconcile productivity and environmental aspects. Fertilizer Research 43, 93–102.
Tagliavini, M., Millard, P. and Quartieri, M. (1998) Storage of foliar-absorbed nitrogen and remobilization for
spring growth in young nectarine (Prunus persica var. nectarine) trees. Tree Physiology 18, 203–207.
Tagliavini, M., Abadia, J., Rombola, A.D., Abadia, A., Tsipouridis, C. and Marangoni, B. (2000) Agronomic
means for the control of iron deficiency chlorosis in deciduous fruit trees. Journal of Plant Nutrition 23,
2007–2022.
Takkar, P.N. and Walker, C.D. (1993) The distribution and correction of zinc deficiency. In: Robson, A.D. (ed.)
Proceedings of the International Symposium on Zinc in Soils and Plants. Kluwer Academic Publishers,
Boston, Massachusetts, pp. 151–164.
Taylor, B.K. (1975) Response of newly planted peach and apple trees to superphosphate. Australian Journal of
Agricultural Research 26, 521–528.
Taylor, B.K. and Issell, L.G. (1976) Comparative effects of foliar- and root-applied phosphorus on one-year-old
peach trees. Australian Journal of Experimental Agriculture and Animal Husbandry 16, 596–599.
Teviotdale, B.T., Ogawa, J.M., Nyland, G. and Kirkpatrick, B.C. (1989) Diseases. In: LaRue, J.H. and Johnson,
R.S. (eds) Peaches, Plums, and Nectarines: Growing and Handling for Fresh Market. University of Cali-
fornia Division of Agriculture and Natural Resources, Publication No. 3331. University of California,
Oakland, California, pp. 118–132.
Tormann, H. and Heyns, D.J. (1988) Effect of progressive soil water tension on soil water availability, vegetative
growth and crop yield of ‘Independence’ nectarines grown in lysimeters. The Deciduous Fruit Grower 38,
430–434.
 Nutrient and Water Requirements 331

Toselli, M., Marangoni, B. and Tagliavini, M. (2000) Iron content in vegetative and reproductive organs of
nectarine trees in calcareous soils during the development of chlorosis. European Journal of Agronomy
13, 279–286.
Van Niekerk, R., Le, P.E. and Pienaar, W.J. (1967) Fertilization programme for fruit trees and table grape vines
in the winter rainfall area. The Deciduous Fruit Grower 17, 141–148.
Veerhoff, O. (1948) Phosphorus deficiency of peach trees in the Sandhills area of North Carolina. Proceedings
of the American Society for Horticultural Science 50, 209–218.
Vizzotto, G. and Costa, G. (1995) Chemical methods to overcome iron-chlorosis in peach trees. Acta
Horticulturae 383, 429–436.
Wallace, A., Wallace, G.A. and Samman, Y. (1983) Zinc and manganese chelate toxicity on nursery and seed-
ling peach trees. Journal of Plant Nutrition 6, 473–489.
Walworth, J.L. and Sumner, M.E. (1987) The diagnosis and recommendation integrated system (DRIS).
Advances in Soil Science 6, 149–185.
Weinbaum, S.A., Johnson, R.S. and DeJong, T.M. (1992) Causes and consequences of overfertilization in
orchards. HortTechnology 2, 112–121.
Weinberger, J.H. (1929) The effect of various potash fertilizers on the firmness and keeping quality of fruits.
Proceedings of the American Society for Horticultural Science 26, 174–179.
Weinberger, J.H. and Cullinan, F.P. (1936) Symptoms of some mineral deficiencies in one-year ‘Elberta’ peach
trees. Proceedings of the American Society for Horticultural Science 34, 249–254.
Weinberger, J.H., Prince, V.E. and Havis, L. (1949) Tests on foliar fertilization of peach trees with urea.
Proceedings of the American Society for Horticultural Science 53, 26–28.
Weir, R.G. and Cresswell, G.C. (1993) Plant Nutrient Disorders 1. Temperate and Subtropical Fruit and Nut
Crops. Inkarta Press, Melbourne, Australia.
Williams, C.F. and Veerhoff, O. (1948) Response of peach trees to boron. Proceedings of the American Society
for Horticultural Science 52, 88–96.
Wills, R.B.H. and Mahendra, M.S. (1989) Effect of postharvest application of calcium on ripening of peach.
Australian Journal of Experimental Agriculture 29, 751–753.
Woodbridge, C.G. (1954) Zinc deficiency in fruit trees in the Okanagan Valley in British Columbia. Canadian
Journal of Agricultural Science 34, 545–551.
Woodbridge, C.G. and McLarty, H.R. (1951) Manganese deficiency in peach and apple in British Columbia.
Scientific Agriculture 31, 435–438.
Worthington, J.W., McFarland, M.J. and Rodrigue, P. (1984) Water requirements of peach as recorded by
weighing lysimeters. HortScience 19, 90–91
Yoshikawa, F.T. (1988) Correcting iron deficiency of peach trees. Journal of Plant Nutrition 11, 1387–1396.
 14 Orchard Floor Management Systems

T.J. Tworkoski and D.M. Glenn

USDA-ARS, Appalachian Fruit Research Station, Kearneysville, West Virginia, USA

14.1 Introduction 332

14.2 Components 332
Orchard floor preparation before planting 332
Weed effects 334
Weed management 335
Orchard floor management effects on insects and small mammals 339
Management of ground covers, row middles and drive alleys 340
Ground cover interactions with irrigation and fertilization 341
14.3 Management Systems 342
Year-round vegetation-free orchard floor 342
Vegetation-free tree rows with vegetated drive alleys 342
Permanent vegetation management by mowing 345
14.4 Orchard Floor Influences 346
Effects of management systems on the environment 346
Orchard floor management interactions with the peach tree 346

14.1 Introduction 14.2 Components

The orchard floor is the soil and understorey
vegetation of an orchard ecosystem. Orchard
floor management decisions can affect the pre-
valence of weeds, insects, small mammals,
disease, soil fertility, water availability, and
potential for erosion and pollution. Appropriate
management of the orchard floor is important
for economic success of the grower and sustain-
ability of the orchard environment. This chapter
considers individual system management
components, their integration and their influ-
ences when managing a peach orchard floor.
332 © CAB International 2008. The Peach: Botany, Production and Uses
Orchard floor preparation before planting
An orchard floor management programme
should begin before tree planting with an under-
standing of potential biological pests and of abi-
otic soil conditions. Site preparation can avoid
or reduce problems associated with the orchard
floor of young peach orchards. Particular atten-
tion should be addressed to weed flora, water
availability, soil pH and structure, the presence of
hard pans and long-term nutrient needs, which
have been discussed elsewhere in this book.
(eds D.R. Layne and D. Bassi)
 Orchard Floor Management
Pre-plant management of weeds and soils
Weed competition can significantly reduce the
growth rate of young peach trees and delay
time to first cropping year. Prior to planting,
weeds that can adversely affect peach trees
should be identified and targeted for control.
Perennial weeds are problematic because
many reproduce vegetatively and spread in
undisturbed soils beneath peach trees. The
management of perennial weeds like john-
songrass (Sorghum halepense (L.) Perse.), poi-
son ivy (Toxicodenron radicans (L.) Kutze) and
Virginia creeper (Parthenocissus quinquefolia
(L.) Planch.) can require two or more herbi-
cide or cultivation treatments (Tworkoski and
Young, 1990). In addition, weeds such as yel-
low nutsedge (Cyperus esculentus L.) are not
controlled by herbicides labelled for bearing
peach trees. Repeated cultivations or herbi-
cide applications prior to planting can avoid
injury to newly planted peach trees.
Annual weeds also compete with young
peach trees and harbour insect and disease
pests that injure peach trees and fruit (Duffus,
1971; Weller et al., 1985; Skroch and Shribbs,
1986). Establishing a grass sod in the year or
two prior to planting can help reduce peren-
nial weed and weed seed banks. The weed
seed bank in the soil can be reduced if weeds
are not allowed to flower. Mowing in conjunc-
tion with competition from the grass cover
selects against many broadleaved weeds while
favouring grass. Grass sod also improves the
organic matter, soil structure and other soil
properties, compared with bare or tilled soil.
The species of grass used as a pre-planting
cover should be adapted to the planting envi-
ronment and subsequent sod management
issues must be considered. Planting fruit trees
in soil prepared with ‘K-31’ tall fescue (Fes-
tuca arundinacea Schreber) sod, killed before
tree planting, reduced subsurface leaching of
nitrate-N and also reduced the amount of
herbicide used in young orchards (Biggs et al.,
1997). However, if left as a ground cover, ‘K-31’
fescue can become competitive with peach
trees (Welker and Glenn, 1988). Other ground
covers such as common bermudagrass (Cyn-
odon dactylon L.) may inhibit the growth of
newly planted peach trees by competition
and allelopathic effects (Weller et al., 1985).
333
Less competitive but well-adapted grasses can
also be sown where future tree rows are to be
planted (Butler, 1986; Willmott et al., 2000).
Several months before planting peach
trees, the sod should be killed with a non-
selective contact herbicide. In this killed sod,
grass residue acts as a mulch to increase soil
water availability, suppress weeds and enable
water penetration into the soil (Welker and
Glenn, 1988; Glenn and Welker, 1989b). The
dead sod can break down after it has been
killed but it may be removed where trees are
planted to reduce habitat for rodents near tree
trunks. A planting hole that has been dug
with an auger may suffice. Cultivation may
still be necessary in narrow strips within the
killed sod into which trees are planted. How-
ever, soil cultivation should be minimized to
reduce bringing weed seed to the soil surface,
where it may germinate. Additional weed
and soil-borne disease control can be obtained
by soil solarization in the vegetation-free strips.
Clear, polyethylene plastic can be installed on
the even surface of moist soil to elevate soil
temperatures for 6 to 8 weeks during the
warmest times of the growing season prior to
planting. Soil sterilization has greatest prom-
ise in warm areas such as the south-east USA
and California, where the required steriliza-
tion time can decrease as soil temperatures
increase above 37°C. Improved plastic sheet-
ing may enable soil sterilization to be used in
cooler regions (Katan and DeVay, 1991).
Raised beds
Poor drainage can be mitigated with raised
planting beds. In Ohio, yield of peach trees
improved by 56% over 5 years when the trees
were grown in raised beds rather than flat
areas (Funt et al., 1997). The improved yield
may have been associated with reduced water-
logging problems that may stimulate shallow
rooting, and greater root regeneration may
have enabled the trees to exploit water from
deeper soil when the shallow soil dried. Raised
beds did not increase yield of young peach
trees grown in sandy soil in New Jersey (Beld-
ing et al., 2003). Trees grown in raised beds in
sandy soils may require supplemental irriga-
tion but, in heavy soils, raised beds appear to
benefit trees by increasing gas exchange to
334 T.J. Tworkoski and D.M. Glenn
the root system. The impact of raised beds on
mature tree growth and susceptibility to wind
throw requires further study.
Weed effects
Yield loss due to weeds in
established orchards
Weeds can reduce yield in 3-year-old peach
trees by 94% (Welker, 1984). Older peach trees
were less susceptible to weed competition,
but the competitive impact of weeds is likely
to vary with weed species and the availability
of potentially limiting resources (Layne et al.,
1981; Glenn and Welker, 1989b; Tworkoski
and Glenn, 2001). For example, peach yield is
likely to decrease with weed competition under
dry conditions. Yield reductions due to weed
competition have been demonstrated in other
temperate tree crops. In Alabama, cumulative
yield of pecans over nine seasons increased
by 358% in trees grown with weed control
compared with trees grown without weed
control (Foshee et al., 1997). However, the costs
of complete weed control were not recovered
from gains in yield until the eighth season
after establishment. In New York, yield and
growth of young apple trees increased as
duration of weed control increased during
the growing season (Merwin and Ray, 1997).
Significant benefits were achieved by initiat-
ing weed control early in the growing season
(e.g. May instead of June). When the orchard
floor was maintained weed-free from bloom
until 12 weeks after bloom, MacRae et al. (2007)
found that fruit size, number and total yield
of peach were greater than when the weed-
free interval was shorter.
Winter annual weeds, such as chickweed
(Stellaria spp.), may not compete significantly
with established peach trees but they may
threaten the economic viability of the crop
by attracting or providing an overwintering
habitat to insects (Lygus spp. and stink bugs)
(Atanassov et al., 2002; Parker, 2003). These
piercing–sucking insects can move from the
weed and cause cat-facing damage to the
exterior of fruit (Killian and Meyer, 1984). In
California, mustard (Brassica spp.), wild
radish (Raphanus raphanistrum L.) and vetch
(Vicia spp.) hosted Lygus hesperus, Lygus elisus
and Calocoris norvegicus, which moved to trees
and caused fruit damage (Pickel et al., 2002).
Clean cultivation and effective control of
broadleaved weeds can suppress these pests
(Atanassov et al., 2002). Programmes that
include cultivation for pest management must
weigh possible adverse effects of cultivation
on the fine, shallow roots of peach trees. Flow-
ering weeds also can disrupt the pollination
of peach trees during bloom by attracting
bees and other wild insects.
Dandelions (Taraxacum officinale Wigg.)
and chickweed may serve as alternative hosts
for viruses such as Tomato ringspot virus,
which can be transmitted to peach trees by
nematodes and adversely affect peach trees
(Powell and Forer, 1982; Skroch and Shribbs,
1986). Other weed species can support high
populations of insects (e.g. aphids and leaf-
hoppers) and nematodes that transmit viruses
(Duffus, 1971). Nematodes such as root-knot
nematode and dagger nematode can cause
direct damage to peaches as well as vectoring
disease. Based on Duffus’ (1971) observations,
the general reduction of weeds will help
reduce virus-induced diseases because nema-
todes and nematode-transmitted viruses have
a diverse range of weed hosts.
Benefits of weed-like vegetation
Negative impacts of weeds are well docu-
mented but some invading plants that are
characterized as weeds may have value. A
potentially confounding effect of understorey
vegetation is that although broadleaved
weeds and grass can both deplete soil water,
water deficits may be ameliorated by the
greater moisture penetration into soil covered
with grass (Atkinson and White, 1981; Glenn
and Welker, 1989b). Often understorey vege-
tation is classified as ‘weeds’ and ‘grasses’.
This classification can be relevant because
grasses can compete with peach trees but they
generally do not cause problems as disease
and insect hosts. In addition, grouping broad-
leaved weeds together simplifies manage-
ment decisions but this form of general
categorization should be scrutinized. There is
evidence that some broadleaved vegetation
may be beneficial by providing habitat to
 Orchard Floor Management
predatory insects that feed on herbivorous
insects (see references within Haynes, 1980
and Atkinson and White, 1981; Wooldridge
and Botha, 1991). Brown (2001) was able to
increase biological control of insect pests on
peach with buckwheat (Fagopyrum esculen-
tum), dill (Anethum graveolens), tansy (Phacelia
tanacetifolia) and a wildflower mix without
increasing the damage by plant bugs or stink
bugs (Brown, 2002). Some, perhaps many,
broadleaved plants that are classified as weeds
may have little impact on peach production
or may even be viable ground covers. Improved
knowledge of the impact of specific weed
populations can contribute to efforts to man-
age weeds in the understorey community.
Weed management
In established orchards, identification and
control of problem weeds should be based on
impact potential. Some weeds pose signifi-
cant threats of competition (e.g. johnsongrass)
whereas others do not (e.g. Whitlow grass
(Draba verna L.) and nimblewill grass (Muhlen-
bergia schreberi J.F. Gmel.)) (Parker and Meyer,
1996). Weeds posing a significant threat are
often best managed while they are seedlings
since large weeds can be highly competitive,
become significant seed sources and be diffi-
cult to kill. The location of weeds in an orchard
will also influence management strategies. In
many established peach orchards the orchard
floor is partitioned into areas below the trees
(the tree row) and between the tree rows
(drive alleys). It is possible that a plant which
is desirable in the drive alley (e.g. fescue) may
be a weed in the adjacent tree row. A widely
used weed control technology in the USA is
with synthetic chemical herbicides, but other
techniques are available and are of increasing
interest. Combinations of cultivation, flaming,
mulching and ‘natural product herbicides’ may
be acceptable for use in ‘organic’ management.
Herbicides
Two general categories of herbicide include
pre-emergence and post-emergence herbicides.
The pre-emergence herbicides are applied to
335
the soil prior to weed seed germination and
are absorbed by the emerging weed seedling.
Pre-emergence herbicides kill weed seedlings
by different modes of action. Post-emergence
herbicides kill weeds on contact and these
also have different modes of action. Post-
emergence herbicides are useful for ‘spot’
applications to localized infestations of large
or troublesome weeds such as perennial weed
escapes. Pre-emergence and post-emergence
herbicides can be applied together or sepa-
rately in rotation to manage weeds. Numer-
ous herbicides may be used to manage weeds
in peach orchards but their use must comply
with label instructions (Table 14.1). It is vital
to distinguish between herbicides used in
non-bearing and bearing peach trees because
some herbicides will damage young trees
with green bark.
Decisions to apply herbicides should be
based on scouting results and site history with
the goal of maintaining weed populations to
acceptable threshold levels. Thresholds can
be based on the potential economic loss due
to decreased yield or fruit quality or threat to
labour operations associated with increased
density of weed populations. As noted ear-
lier, some winter annual weeds (e.g. chick-
weed) and ground covers (e.g. clover) harbour
cat-facing insects such as stink bugs that pose
an unacceptable risk to fruit quality. In south-
ern locations of the USA these broadleaved
plants can be managed with 2,4-D applica-
tions 8 weeks prior to bloom without damag-
ing grass in drive alleys. It is generally known
that yields decrease due to weed competition
and that young peach trees are more suscep-
tible than mature peach trees (Weller et al.,
1985). However, little is known about economic
threshold levels of weeds related to insect injury
and yield loss for peach trees and there is a
need for additional research in this area.
Consecutive applications of the same
herbicide or of different herbicides with the
same mode of action may lead to the devel-
opment of weed populations that are resis-
tant to the class of herbicides being used
(Welker, 1984). Combinations of herbicides
with different modes of action have been
found to be more effective in weed control
than using a single herbicide (Welker, 1984).
However, even repeat applications of the
336 T.J. Tworkoski and D.M. Glenn

Table 14.1. Herbicidesa currently recommended for peach culture in the USA.

Herbicide and active ingredient Application

Pre-emergence
Dichlobenil (Casoron) Established trees for control of broadleaved weeds,
2,6-dichlorobenzonitrile quackgrass and fescue
Diuron (Karmex) Trees established at least 3 years in the orchard for
3-(3,4-dichlorophenyl)-1,1-dimethylurea control of annual broadleaved and grass weeds
Isoxaben (Gallery) Control of broadleaved weeds in non-bearing trees
N-[3-(1-ethyl-1-methylpropyl)-5-isoxazolyl]
-2,6-dimethoxybenzamide
Napropamide (Devrinol) Established bearing and non-bearing trees for
N,N-diethyl-2-(1-naphthalenyloxy)propionamide control of annual grasses and small-seeded
broadleaved weeds
Norflurazon (Solicam) Control of annual grasses and small-seeded
4-chloro-5-(methylamino)-2-(α,α,α-trifluoro- broadleaved weeds
m-tolyl)-3(2H)-pyridazinone
Oryzalin (Surflan) Bearing and non-bearing trees for control of annual
3,5-dinitro-N4,N4-dipropylsulfanilamide grasses and small-seeded broadleaved weeds
Oxyflurofen (Goal) Control of broadleaved weeds in established trees
2-chloro-1-(3-ethoxy-4-nitrophenoxy)-
4-(trifluoromethyl)benzene
Pendimethalin (Prowl) Non-bearing established trees for control of annual
N-(1-ethylpropyl)-3,4-dimethyl-2,6- grasses and small-seeded broadleaved weeds
dinitrobenzenamine
Pronamide (Kerb) Control of grass and small-seeded broadleaved
3,5-dichloro-N-(1,1-dimethyl-2-propynyl) weeds in established trees
benzamide
Simazine (Princep and generics) Trees established at least 1 year in the orchard for
2-chloro-4,6-bis(ethylamino)-S-triazine control of annual broadleaved weeds
Terbacil (Sinbar) Established trees for control of grass and
3-tert-butyl-5-chloro-6-methyluracil broadleaved weeds

Post-emergence
2,4-D amine (generics) Control of annual and perennial broadleaved weeds
(2,4-dicholorophenoxy)acetic acid in established trees
Fluazifop (Fusilade) Control of annual and perennial grasses in newly
(R)-2-[4-[[5-(trifluoromethyl)-2-pyridinyl]oxy] planted and established trees
phenoxy]propanoate
Glyphosate (Roundup/Touchdown) Control of broadleaved and grass weeds
N-(phosphonomethyl)glycine
MSMA (MSMA Arsonate) Selective control of annual and perennial weeds in
monosodium acid methanearsonate non-bearing trees
Paraquat (Gramoxone) Control of broadleaved and small grass weeds
1,1′-dimethyl-4,4′-bipyridinium dichloride
Scythe Non-selective burn-down of weeds
pelargonic acid
Sethoxydim (Poast) Control of annual and perennial grasses in
2-[1-(ethoxyimino)butyl]-5-[2-(ethylthio)propyl]- established trees
3-hydroxy-2-cyclohexen-1-one
aNotendorsed by the US Department of Agriculture. Labels and state extension recommendations
should be followed.
74 75

76

77

78

Plate 74. Fusetto tree form.

Plate 75. Y-shaped tree form (courtesy of D.R. Layne, Clemson, South Carolina, USA).
Plate 76. Modern picking platform.
Plate 77. Generalized shapes of concentration curves of mineral nutrients in the leaf during growing season.
Curves show trends, not actual levels for the northern hemisphere.
Plate 78. Red coloration on leaves and stems of nitrogen-deficient peach shoot.
79 80

82

81

83 84

Plate 79. Smoother finish and more red coloration on fruit from low nitrogen trees (top row) compared to high
nitrogen trees (bottom row).
Plate 80. Purple coloration and leathery texture of phosphorus-deficient peach leaf.
Plate 81. Pale colour and leaf rolling caused by potassium deficiency in peach leaves.
Plate 82. Discoloration on leaf margins and tip caused by magnesium deficiency.
Plate 83. Zinc deficiency symptoms of interveinal chlorosis on a peach leaf.
Plate 84. Rosetting or ‘little leaf’ symptoms of zinc deficiency in peaches.
85 86

87 88

89 90

91

Plate 85. ‘Netting’ symptom of iron chlorosis on a peach leaf. Veins remain green while the rest of the leaf has
turned yellow.
Plate 86. Manganese deficiency symptoms on a peach leaf. Bands along the main veins remain green.
Plate 87. Ten-year-old ‘Sunhigh’ peach trees, originally planted in 2.4 m vegetation-free areas with composted
poultry litter placed beneath some trees at a rate of 11.6 kg litter/m2 to a depth of 10 cm (1.1 kg N per tree). Pho-
tograph was taken the second season after application when some weeds had begun to grow through the mulch.
Plate 88. Trees planted in cultivated (left) and killed-sod (right) strips. Killed sod provides weed suppression and
improves water penetration.
Plate 89. ‘Jersey Dawn’ and ‘Redskin’ peach tree, the same age (approximately 3 years after planting) but some
planted and grown in 0.6 m (small trees in foreground) and others grown in 2.4 m (large trees) vegetation-free
areas with K-31 fescue in drive alleys.
Plate 90. Brown rot blossom blight of peach caused by Monilinia fructicola. Infected blossom with gumming and
canker development at the base of the peduncle.
Plate 91. Brown rot fruit rot of peach caused by Monilinia fructicola.
92 94

93

Plate 92. Jacket rot of peach caused by Botrytis cinerea. The jacket (or shuck) is infected, but the immature fruit is still disease-free.
Plate 93. Green fruit rot of peach caused by Monilinia fructicola. The decay spreads to neighbouring healthy fruit by contact.
Plate 94. Apothecia of Sclerotinia sclerotiorum are produced on overwintering sclerotia (bottom). Ascospores are forcibly discharged in a spore cloud (reprinted, with
permission, from Strand, 1999).
95 96

97
98

99
100

Plate 95. Lesion of the shothole fungus Wilsonomyces carpophilus on peach twig where the pathogen
overwinters. Sporodochia of the fungus in the centre of a stem lesion.
Plate 96. Symptoms of shothole caused by Wilsonomyces carpophilus on peach fruit and leaf.
Plate 97. Symptoms of scab caused by Fusicladosporium carpophilum on peach fruit.
Plate 98. Sporulating stem lesion of peach rust caused by Tranzschelia discolor.
Plate 99. Symptoms of peach rust caused by Tranzschelia discolor on peach leaves.
Plate 100. Leaf lesions with uredinia of peach rust caused by Tranzschelia discolor.
101

103

102

104

Plate 101. Symptoms of peach rust caused by Tranzschelia discolor on peach fruit.
Plate 102. Peach leaf curl caused by Taphrina deformans.
Plate 103. Advanced symptoms of peach leaf curl caused by Taphrina deformans with leaf deformation,
discoloration, necrosis and a subtle white layer of sexually produced asci containing ascospores.
Plate 104. Symptoms of peach leaf curl caused by Taphrina deformans on developing peach fruit.
105 106

107
108

109
110

Plate 105. Peach twig with overwintering mycelium and embedded chasmothecia of powdery mildew caused by
Podosphaera pannosa.
Plate 106. Powdery mildew caused by Podosphaera pannosa on peach leaves.
Plate 107. Powdery mildew caused by Podosphaera pannosa on developing peach fruit.
Plate 108. Powdery mildew caused by Podosphaera pannosa on mature peach fruit.
Plate 109. Powdery mildew caused by Podosphaera pannosa on developing nectarine fruit.
Plate 110. Close-up of chasmothecia of powdery mildew caused by Podosphaera pannosa on peach twig.
111 112

113

114

116

115

Plate 111. Symptoms of silver leaf disease caused by Chondrostereum purpureum on peach leaves. Left:
diseased leaf. Right: healthy leaf.
Plate 112. Cross-section through scaffold branch of peach infected by Chondrostereum purpureum.
Plate 113. Peach tree with Leucostoma (Cytospora) or perennial cankers on scaffold branches (reprinted from
Ogawa et al., 1995, with permission of APS).
Plate 114. Leucostoma or perennial cankers caused by Leucostoma cincta (Cytospora cincta) around nodes of
peach shoot (reprinted from Ogawa et al., 1995, with permission of APS).
Plate 115. Pycnidia of Leucostoma cincta (Cytospora cincta) on cherry twig. Spore masses are exuded by some
pycnidia on the upper part of the twig.
Plate 116. Fungal gummosis caused by Botryosphaeria dothidea. Blisters develop around lenticels on young bark
that develop in the second or third growing season of the tree.
117 118

119

120 122

121

Plate 117. Fungal gummosis caused by Botryosphaeria dothidea. Gumming blisters and necrotic tissue under
lenticels that develop under the bark.
Plate 118. Fungal gummosis caused by Botryosphaeria dothidea. Lesions may coalesce and form extensive
cankers with excessive gumming on lower peach tree trunks.
Plate 119. Constriction canker caused by Phomopsis amygdali. One-year-old peach twigs with infections around
buds. The lower twig shows the zonations around the infection site.
Plate 120. Constriction canker caused by Phomopsis amygdali. Flagging and withering of blighted peach twigs
distal to twig cankers.
Plate 121. Cirri (tendrils) exuding from pycnidia of Phomopsis amygdali. Each cirrus is composed of abundant
conidia of the pathogen.
Plate 122. Dieback of peach tree in foreground affected by Phytophthora root and crown rot (reprinted, with
permission, from Strand, 1999).
123 124

125 126

127 128

Plate 123. Phytophthora crown rot on peach tree (reprinted, with permission, from Strand, 1999).
Plate 124. Peach tree with an aerial infection of a Phytophthora sp. (reprinted, with permission, from Strand,
1999).
Plate 125. Leaf symptoms of peach tree affected by Armillaria root rot.
Plate 126. Armillaria-infected peach trees showing dieback.
Plate 127. Mycelial fans of Armillaria sp. under the bark of infected peach tree.
Plate 128. Rhizomorph of Armillaria mellea (top) and healthy root (bottom) (reprinted, with permission, from
Strand, 1999).
129 130

131 133

132

Plate 129. Basidiomes of Armillaria sp. at the base of infected peach tree.
Plate 130. Verticillium-infected peach tree with wilting of branches (reprinted, with permission, from Strand,
1999).
Plate 131. Cross-section through branch of Verticillium-infected peach tree with discoloured outer xylem
(reprinted, with permission, from Strand, 1999).
Plate 132. Orchard with trees affected by peach tree short life.
Plate 133. Discoloured tissue under the bark of peach tree short life-affected tree.
134 135

136 137

138

139

Plate 134. Rootstock sprouting from peach tree killed by peach tree short life.
Plate 135. Postharvest brown rot of peach fruit caused by Monilinia fructicola.
Plate 136. Postharvest grey mould of nectarine fruit caused by Botrytis cinerea.
Plate 137. Postharvest Rhizopus rot of peach fruit caused by Rhizopus stolonifer with decay spreading by contact
to healthy fruit (nesting).
Plate 138. Postharvest Gilbertella rot of peach fruit caused by Gilbertella persicaria.
Plate 139. Sour rot caused by Geotrichum candidum initiated in wounds of the skin.
140 141

142 143

145
144

Plate 140. Anthracnose of peach caused by Colletotrichum acutatum showing circular rings where spores of the
fungus are produced.
Plate 141. Electron micrograph of phloem sieve tube occluded with cells of phytoplasma.
Plate 142. Leaves of periwinkle (Catharanthus roseus), which can serve as an indicator plant for phytoplasmas,
colonized by dodder (Cuscuta sp.), which serves as a transmission bridge.
Plate 143. Dead peach buds and twig canker associated with bacterial canker.
Plate 144. Branch death and sucker growth from the rootstock associated with bacterial canker.
Plate 145. Wood with discoloured streaks and the collapse and wilt of newly emerged growth associated with
bacterial canker.
146 147

148

149

151
150

Plate 146. Small branch killed by Pseudomonas syringae pv. syringae with the canker extending into a larger limb.
Plate 147. Water-soaked bark and lightly discoloured wood in early spring associated with bacterial canker.
Plate 148. Discoloured wood approximately a week after occurrence of cold damage, with the bark easily sepa-
rating and appearing undamaged.
Plate 149. Non-fluorescent and fluorescent bacteria associated with the bacterial canker complex cultivated on
King’s medium B.
Plate 150. Chlorotic peach leaves with lesions caused by Xanthomonas arboricola pv. pruni.
Plate 151. Newly formed, water-soaked, greyish, angular bacterial spot lesions on peach leaf caused by
Xanthomonas arboricola pv. pruni.
152 153

154
155

156 157

Plate 152. Bacteria streaming from leaf lesion caused by Xanthomonas arboricola pv. pruni.
Plate 153. Newly formed, water-soaked bacterial spot lesions on peach fruit near the growth stage of pit hardening.
Plate 154. Bacterial spot lesions on fruit at harvest confined to the fruit surface that developed from infections
occurring after pit hardening.
Plate 155. Bacterial spot lesions on fruit at harvest caused by early infections occurring soon after shuck split and
before pit hardening.
Plate 156. Bacterial spot spring canker (Xanthomonas arboricola pv. pruni) with a black, greasy appearance.
Plate 157. Terminal dieback and black tip canker, summer cankers on current year’s twigs, lesions on leaves,
and nodes where leaves have defoliated caused by Xanthomonas arboricola pv. pruni.
158 159

160
161

162 163

Plate 158. Discoloured bark tissue of spring canker Xanthomonas arboricola pv. pruni.
Plate 159. Crown gall caused by Agrobacterium tumefaciens on peach trees recently dug from a nursery.
Plate 160. Peach tree expressing phony peach disease symptoms. Leaves are greener and denser and internodes
are shortened giving the tree a compact, flat canopy with an ‘umbrella-like’ appearance (from R.F. Mizell, III).
Plate 161. X-disease phytoplasma-infected peach with necrotic lesions non-uniformly distributed on rolled and
pale green leaves.
Plate 162. X-disease phytoplasma-infected trees exhibit a loss of vigour, leaves have a pale green colour and
severely infected branches die.
Plate 163. Tree infected with peach yellows phytoplasma showing a bushy appearance on ends of branches and
premature colouring and ripening of fruit (from A. Ragozzino).
 Orchard Floor Management
same combination of herbicides can result in
population shifts and new weed problems
(Tworkoski et al., 2000b). Long-term applica-
tions of the same herbicide may also contrib-
ute to herbicide residue carry-over, which can
adversely affect the growth of newly planted
peach trees (Tworkoski et al., 2000a; Tworko-
ski and Miller, 2001). Repeated control of
weeds with pre-emergence herbicides and
mechanical tillage may reduce soil structure,
fertility and orchard productivity compared
with ‘living’ and straw–hay mulches (Merwin
et al., 1994) and killed-sod systems (Glenn
and Welker, 1989b). Productivity loss may be
associated with reduced organic matter and
water infiltration, and with elevated soil tem-
peratures in non-mulched sites. At the time of
flowering, heat absorbed during the day by
vegetation-free orchard floors will radiate
from the soil at night, warm the air and pos-
sibly reduce cold injury to peach blossoms.
Thus, soil that is continuously bare may con-
tribute to reduced long-term orchard produc-
tivity but time intervals without vegetation
cover can protect current-year cropping.
Herbicides which may be acceptable for
organic growers have been developed. In gen-
eral they are contact-active and may require
repeated applications when established plants
are being controlled. These herbicides include
pelargonic acid (Scythe; Mycogen Corp., San
Diego, California), vinegar (BurnOut; St Gabriels
Laboratory, Orange, Virginia) and essential
oil of clove (Matran; EcoSmart Technologies,
Inc., Franklin, Tennessee). Essential oils of cin-
namon, clove, summer savory and red thyme
have herbicidal activity and may be useful as
‘natural product herbicides’ (Tworkoski, 2002).
However, these herbicides can be expensive
and may have limited efficacy against some
weeds. These herbicides appear to be most
effective when applied to small weeds, gener-
ally early in the growing season.
Mechanical and flaming techniques
Mechanical weed control devices such as discs
or cultivators can control ground vegetation
in areas where the orchard floor is not main-
tained with ground covers, such as in tree
rows. Where drive alleys are maintained in
permanent ground cover, cultivation can be
337
performed beneath trees but care is necessary
to avoid damaging low branches or tree trunks.
Where drive alleys are maintained in winter
ground covers, shallow disking in early spring
can eliminate or reduce competition and pro-
vide residual mulch. Shallow disturbance of
the soil is necessary to minimize bringing new
weed seed to the soil surface and to avoid
damaging shallow peach roots. Mechanical
cultivators have been developed that can till
to the tree and reduce or avoid damaging the
trunk (Weed Badger, Marion, North Dakota;
The Green Hoe Co., Portland, New York).
Flame weeding eliminates weeds by
searing, not burning, the vegetation (Ames and
Kuepper, 2004). Torches fuelled by kerosene
or propane are pulled behind a tractor at a
speed that will wilt vegetation. Drawbacks to
flaming include fire and fuel hazards, poten-
tial tree injury, water and fuel requirements,
and lack of uniform weed kill. Flaming is an
alternative weed management technology that
may be useful in organic systems. Hand and
tractor-mounted flame weeding equipment is
commercially available (Flame Engineering,
Inc., LaCrosse, Kansas; Thermal Weed Control
Systems, Inc., Neillsville, Wisconsin).
Mulching
Organic mulches, such as straw, sawdust and
composted animal waste, can be applied
beneath fruit trees to suppress weeds (Fig. 14.1/
Plate 87). Composted poultry litter applied to
a depth of 10 cm beneath peach trees sup-
pressed soil-germinating weeds but additional
control was necessary for weeds germinating
in the mulch (Preusch and Tworkoski, 2003).
Composted mulch that is applied in too deep
a layer may have the undesirable effect of
releasing significant P to the soil (Preusch et al.,
2002). A layer of newspaper or cardboard could
be applied beneath the mulch to increase
weed suppression (Ames and Kuepper, 2004)
but moisture penetration may be impeded
and, as this lower layer degrades, weeds can
grow through the mulch (T.J. Tworkoski, per-
sonal observation). If the mulch is raked aside,
the lower newspaper layer replaced and the
mulch raked back, then weeds can be suppres-
sed for a longer time. These are labour-intensive
practices which may not be economically
338 T.J. Tworkoski and D.M. Glenn

Fig. 14.1. Ten-year-old ‘Sunhigh’ peach trees, originally planted in 2.4 m vegetation-free areas with
composted poultry litter placed beneath some trees at a rate of 11.6 kg litter/m2 to a depth of 10 cm
(1.1 kg N/tree). Photograph was taken the second season after application when some weeds had
begun to grow through the mulch.

viable for large-scale commercial operations.
Other beneficial effects of mulch include slow
release of some nutrients, increased soil organic
matter and improved soil structure (Haynes,
1980). Soil moisture retention is improved
when mulch is applied to tree rows (Skroch
and Shribbs, 1986). Compost mulch has also
been shown to inhibit growth of the brown rot
fungus, Monilinia fructicola (G. Wint) Honey, in
the laboratory (Brown and Tworkoski, 2004).
High microbial biodiversity in compost may
increase competition for resources and reduce
production of inoculum. Pine straw or hay
applied beneath trees to a depth of 12 cm can
provide some mulching benefits; however,
they may pose a fire hazard.
Organic material that can be used for
mulch may not be widely available and trans-
portation costs of the large quantities neces-
sary for mulch may prohibit its use. These
problems can be ameliorated if mulch can be
obtained from ‘on-site’ activities such as
growing sorghum sudangrass (Sorghum ×
drummondii (Steudel) Millsp. & Chase) for
mulch (Ames and Kuepper, 2004). Cover crops
between trees or in fields near the orchard can
be a source of organic mulch. Sickle bar mow-
ers can cut ground covers, which can then be
raked or blown beneath trees. To prevent the
ground cover from becoming a weed problem
in the tree rows, the ground cover in the drive
alley should be mown before it produces
seed. In addition, mulch should be kept at
least 20–30 cm from the trunk to avoid rodent
and collar rot injury to peach tree trunks
(Ames and Kuepper, 2004).
Inorganic mulches such as black plastic
and geotextile sheets can effectively suppress
weeds. Their use requires a greater initial invest-
ment than most organic mulches. Another
drawback is that the waste fabric must be
removed and disposed of, but this problem
may be less significant if longer-lived mulch
is used. In the south-eastern USA, growers
using black plastic and raised beds can achieve
weed control for up to 2 years, provided weed
seed has been killed with methyl bromide.
Drip irrigation beneath the plastic is necessary
in this system.
Biological control of weeds in peach
orchards has been attempted by planting a
short-lived cover plant which grows quickly
 Orchard Floor Management
and dies, resulting in weed suppression by
competition and residue which acts as a mulch.
An example of such smother crops is Brassica
campestris, which was planted in May and
suppressed early-season weeds while it was
actively growing (Halbrendt, 1993). However,
by July weed growth was similar to untreated
plots. For success, smother crops require weed
suppression for longer periods without com-
petition that reduces yield of peach trees.
Orchard floor management effects
on insects and small mammals
Cover crops and weeds can provide food and
habitat for rodents, insects, nematodes, microbes
and viruses (Norris, 1986). Management of the
orchard floor can influence the balance between
organisms that are beneficial and those that
are harmful to peach production. Research
has provided insight into orchard floor mani-
pulation effects on insect populations and
behaviour in apple and pear orchards but less
is known in peach orchards. The potential
impact of such pests or the benefits of other
insects should be considered in the selection
of an orchard floor management system.
Insect populations are directly and indi-
rectly affected by the composition and abun-
dance of flora on the orchard floor (Alston,
1994). In peach, control of many annual broad-
leaved weeds and legumes is necessary because
they provide habitat for tarnished plant bugs
and stink bugs (LaRue and Johnson, 1989; Ata-
nassov et al., 2002; Ames and Kuepper, 2004). In
apple orchards, broadleaved weeds such as
common mallow (Malva neglecta Wallr.), field
bindweed (Convolvulus arvensis L.), knotweed
(Polygonum spp.), morning glory (Ipomoea spp.),
prickly lettuce (Lactuca serriola L.), puncture vine
(Tribulus terrestris L.) and white sweetclover
(Melilotus alba Medikus) enhanced phytopha-
gous mites that are detrimental to tree produc-
tivity (Alston, 1994). Phytophagous mites were
managed to economically acceptable levels by
reducing these host broadleaved weed species
to less than 12% of ground cover while main-
taining ground cover of at least 50% of other
ground cover species (orchardgrass (Dactylis
glomerata L), red fescue (Festuca rubra L.) and
339
lucerne (Medicago sativa L.)) which harbour
predatory mites.
Research has demonstrated that ground
covers such as mustards (Brassica spp.), buck-
wheat (Fagopyrum spp.), dwarf sorghum
(Sorghum spp.) and various members of the
Apiaceae (Umbelliferae) and Asteraceae (Composi-
tae) families can attract more beneficial insects
than pests. In addition to floristic effects, man-
agement of the orchard floor can influence
insect behaviour and population demograph-
ics (Brown et al., 1997; Brown and Glenn, 1999).
Mowing frequency can affect the population
size of phytophagous and predacious insects,
with increases of both groups resulting from
decreased mowing in pear orchards in the
north-western USA (Horton et al., 2003). There
is a possibility of managing movement of nat-
ural enemies of insects into fruit tree canopies
with cultural manipulation of the orchard
floor, such as mowing. In California it is rec-
ommended that lucerne growing near a fruit
crop not be mowed if there are Calocoris bugs
feeding on the legume, since mowing may
induce movement of Calocoris bugs into the
tree canopy, where they may damage fruit
(Ames and Kuepper, 2004). Increased research
is needed to understand and improve the use
of understorey vegetation as a tool for insect
pest management in peach orchards.
In addition to cover crops and weeds,
mulches and herbicides that are part of the
orchard floor management system can influ-
ence insect populations. Modification of the
orchard floor with composted mulch enhanced
ground-foraging generalist predators, and
predator activity may be enhanced when the
orchard floor is disturbed or treated with her-
bicide (Brown and Tworkoski, 2004; Mathews
et al., 2004). Mulch increased prey resources
to support predator populations and herbi-
cides enhanced habitat for predators with
improved physical cover and microclimate
effects. However, herbicide use can also decrease
predatory insect populations. Integrated pest
management (IPM) programmes that target
control of the two-spotted spider mite (Tetrany-
chus urticae Koch) with the predacious mite
(Neoseiulus (Typhlodromus) fallacies (Garman))
should avoid use of 2,4-D amine, gramoxone
and terbacil because these herbicides were
more toxic to the predator and were likely to
340 T.J. Tworkoski and D.M. Glenn
differentially decrease predator populations
(Rock and Yeargan, 1973).
Ground cover and mulch near tree trunks
can lead to rodent damage in fruit orchards.
In New York, meadow vole (Microtus pennsyl-
vanicus Ord) density increased and young apple
trees were injured in orchards with crown vetch
(Coronilla varia L.), hay–straw mulch and red
fescue (F. rubra L.) sod below trees (Merwin
et al., 1999). Tree damage was controlled by
reducing vegetation, trapping, and using tree
guards and vole predators. In British Colum-
bia, Canada, intensive weed control within an
orchard reduced montane vole (Microtus mon-
tanus) abundance but a compensatory increase
in other small mammal populations (deer
mouse (Peromyscus maniculatus) and north-
western chipmunk (Eutamias amoenus)) was
observed (Sullivan et al., 1998). Montane vole
damage to apple trees was reduced with scent
mixtures from ermine (Mustela erminea) (Sul-
livan et al., 1990). Such chemical repellants
may have application to rodent control in peach
orchards.
Management of ground covers,
row middles and drive alleys
The entire orchard floor, or a part of it, can be
mulched, cultivated, treated with herbicide
or maintained with ground covers. The tree
rows can be maintained vegetation-free while
the drive alleys between tree rows are man-
aged with ground covers. Ideal ground cov-
ers should help control erosion, stand up to
traffic, require low maintenance, suppress
weeds, improve soil quality and not compete
with the peach trees. However, ground covers
and mulch can reduce flood and furrow irri-
gation efficiency and may not be used during
the growing season, particularly under dry
conditions in areas that include parts of Cali-
fornia. Ground covers must adapt to climatic
and edaphic conditions. In addition, manage-
ment of drive alleys must be coordinated with
tree rows. For example, ground covers in
drive alleys may be cut to provide organic
mulches beneath tree rows and they may also
provide habitat for beneficial insects such as
predatory mites and spiders.
Generally, grass is more beneficial to soil
flora and fauna than clean cultivation (Haynes,
1980). Grass ground covers with potential for
use in drive alleys have been evaluated in
terms of rate of grass establishment, height
and spreading characteristics, root and water
use traits, and tolerance to drought, heat,
shade, cold and traffic (Butler, 1986).
Many growers in the eastern USA plant
vigorous grasses on the orchard floor such as
Kentucky bluegrass (Poa pratensis L.), fescue
(Festuca elatior L.) and orchardgrass (D. glom-
erata L.) with herbicide strips in the tree row
(Skroch and Shribbs, 1986). These cool-season
grasses may not be the best choice as an
orchard floor cover because they may require
frequent mowing and can spread aggressively
from drive alleys into tree rows (Willmott et al.,
2000). In New Jersey, perennial ryegrass (Lolium
perenne L.) and creeping red fescue (F. rubra
L.) may succumb to infectious diseases or
environmental stress and should not be used.
A number of grass cultivars of tall, hard and
chewings fescues (F. arundinacea Schreber, Fes-
tuca longifolia Thuill. and F. rubra, respectively)
that require lower maintenance have been
recommended (Willmott et al., 2000). In North
Carolina, Parker and Meyer (1996) deter-
mined that peach tree growth was greater
when grown with nimblewill grass (M. schreberi
J.F. (Gmel.)) than in plots with weeds, centi-
pedegrass (Eremochloa ophiuroides (Munro)
Hack.) or bahiagrass (Paspalum notatum Flugge).
Peach roots grew deeper and in greater number
and lateral distribution in nimblewill grass or
bare ground than in other grass treatments. In
the Pacific Northwest, most orchards use
perennial grass with shallow roots as ground
cover between tree rows and vegetation-free
strips beneath trees (Granatstein, 2002).
In the southern USA winter legumes
such as vetch (Vicia spp.) and subterranean
clover (Trifolium subterraneum L.) have been
used as cover crops to improve soil physical
and nutrient properties that can benefit fruit
trees (Hoyt and Hargrove, 1986). Subterra-
nean clover had excellent reseeding ability,
and may contribute N and organic matter to
the orchard floor. However, the timing of
nutrient release from legumes must support
peach production without adversely affecting
fruit quality or vegetative tree growth that
 Orchard Floor Management
can result from excessive N late in the season.
In addition, some warm-season legumes may
compete with trees for water and harbour
phytophagous insects. None the less, after
weighing costs and benefits, legumes may
play a role in sustainable practices for peach
orchard floor management. Subterranean clo-
ver has been proposed as a useful ground
cover for drive alleys in peach orchards in
warm locations where winter temperatures
do not drop below –18°C (Ames and Kuepper,
2004). The subterranean clover reseeds in
early summer and dies in the heat of late sum-
mer to produce a weed-suppressive mulch in
Arkansas and California. The clover has also
provided a habitat for beneficial insects.
Ground covers with different species
composition, including broadleaved weeds,
favour diverse arthropod communities that
have been useful for IPM in orchards (Wool-
dridge and Botha, 1991). Such broadleaved
plants may serve as alternative food sources
for predatory mites that feed on thrips, aphids
and other mite species. However, ground
cover flora must be managed to reduce injury
from phytophagous insects. Cat-facing injury
by stink bugs (Pentatomidae) and tarnished
plant bugs (Lygus lineolaris) was correlated with
the presence of legumes such as vetch (Vicia
spp.), clover (Trifolium spp.) and annuals such
as chickweed, pepperweed (Lepidium spp.)
and henbit (Lamium amplexicaule L.) (Killian
and Meyer, 1984; Meyer, 1984). Some species
of plants, if kept succulent, may be used as a
trap crop to attract stink bugs away from the
peaches (Brown, 2002). Spider mite (Tetrany-
chus spp.) populations may also increase in
these ground covers and migrate to peach trees
when ground covers begin to senesce (Meagher
and Meyer, 1990). Ring nematode (Mesocri-
conema xenoplax) may move from legumes as
well as from weeds such as dandelion and
purslane to peach roots (Zehr et al., 1986,
1990).
Ground cover management is not res-
tricted to vegetation management. Synthetic
ground covers have been used to modify the
orchard environment to enhance fruit quality.
Reflective mulches have been applied in drive
alleys to increase light intensity in orchards
and to increase red colour of peaches. Reflect-
ing films can modify the composition of
341
anthocyanins, flavonoids, chlorophyll and
carotenoids in apples (Ju et al., 1999). Layne
et al. (2001) found increased red surface colour
of peaches when metallized reflective film
was placed beneath peach trees 2 to 4 weeks
before harvest. Peaches in the tree lower can-
opy were redder and overall price could
increase by $1 per 11-kg box with increased
colour. Estimated costs for the metallized
mulch was $220/ha.
Ground cover interactions with
irrigation and fertilization
Irrigation combined with managed competi-
tion may increase peach tree productivity
while reducing excess vegetative growth. In
Australia, vegetative vigour was suppressed
and yield was increased by intraspecific root
competition from a high-density orchard
planting in combination with restricted irri-
gation (Chalmers et al., 1981). Interspecific
weed competition from ground cover can also
dwarf young peach trees without reducing
yield expressed on a trunk cross-sectional
area basis (Glenn and Welker, 1996). Reduc-
ing tree size by manipulation of the orchard
floor ground cover requires close manage-
ment of fertilizer and water inputs. In older,
8-year-old peach trees, grass sod competition
reduced total yield and yield of large fruit
(>65 mm) in West Virginia (Glenn and Welker,
1996). In Ontario, Canada, permanent drive
alleys with creeping red fescue combined with
trickle irrigation increased total yield and
yield of large fruit (Layne and Tan, 1988). Yield
decreased when peach trees were grown with
grassed drive alleys but without irrigation.
Yield and the fruit in large size classes increased
with irrigation when ground covers were
present but the economic balance of irrigation
costs and yield benefits should be analysed.
It is likely that supplemental fertilizer
will be needed in productive orchards, regard-
less of the ground cover used, and fertilizer
applications must be prescribed on the basis
of appropriate soil and leaf analyses. In addi-
tion to fertilizer applications, soil nutrients
can be manipulated by partial or complete
kill of ground covers or by addition of organic
342 T.J. Tworkoski and D.M. Glenn
mulches. In Europe, a permanent ground
cover of white clover (Trifolium repens) was
mowed regularly under older trees or winter
rye was sown under young trees in late sum-
mer, followed by mechanical removal in spring
(Bloksma, 2000; Bloksma and Jansonius, 2002).
The permanent ground cover and late sum-
mer sowing provided a means of transferring
soil N from late summer/autumn to the fol-
lowing spring. Organic mulches of bark, grass
and composted waste may also provide a
slow release of nutrients while conserving
water and suppressing weeds throughout the
year. However, organic mulches can increase
the balance of C to N in the soil and immobi-
lize nutrients from soil-applied fertilizer.
14.3 Management Systems
The primary goal of orchard floor manage-
ment systems is to control understorey vege-
tation and manage resources to ensure the
economic success and sustainability of the
orchard. Selection of the type of ground cover,
its spatial and temporal distribution, and the
method of control are critical aspects that
must be integrated for orchard floor manage-
ment. The ground may be completely covered
with permanent vegetation or be controlled
by cultivation, mulch or herbicides so that
some portion of the ground cover remains
(Hogue and Neilsen, 1987). This section con-
siders the integration of several of the previ-
ously described management components in
management systems.
Year-round vegetation-free orchard floor
Benefits and disadvantages of different orchard
floor management systems have been reviewed
(Haynes, 1980; Skroch and Shribbs, 1986; Hogue
and Neilsen, 1987). Continuous, clean cultiva-
tion of the orchard floor aerates the soil and
eliminates competition but loss of organic
matter, breakdown of soil structure, increased
potential for erosion and destruction of shal-
low tree roots will occur. In dry areas where irri-
gation is the primary source of water for peach
crops, year-round vegetation-free conditions
have been used. Year-round tillage is used in
California to control weeds, save water and
provide ditches for furrow and flood irriga-
tion (Vossen and Ingals, 2002). Soil compac-
tion resulting from repeated cultivations can
occur. Rip cultivation may then be necessary
with shanks deep enough to break through
the cultivated layer and allow water to pene-
trate the soil profile. However, deep cultiva-
tion can require heavy equipment, which may
contribute to further soil compaction.
East of the Rocky Mountains, the practice
of clean cultivation in bearing peach orchards
declined during the middle part of the 20th
century (Fogle et al., 1965). However, in South
Carolina and Georgia, some large peach pro-
ducers returned to a herbicide-maintained
bare orchard floor system (D.R. Layne, South
Carolina, 2004, personal communication). In
other eastern locations, in place of complete
removal of vegetation, ground covers were
grown as permanent or temporary compo-
nents of the orchard floor. Ground cover veg-
etation increases soil organic matter, structure
and water penetration, but ground covers
must be managed to control competition and
reduce pests that are associated with them.
Vegetation-free tree rows with
vegetated drive alleys
In the USA the orchard floor beneath peach
trees is often maintained free of weeds with
herbicides and drive alleys may contain tem-
porary or permanent ground covers (Elmore
et al., 1997). Management decisions regarding
the orchard floor before and shortly after plant-
ing will affect the composition and manage-
ment of the orchard floor in a mature orchard.
Establishment of the orchard floor
In preparation for planting, a ground cover of
grass sod should be installed as a fallow crop
for at least 2 years to adjust soil pH and to
decrease numbers of nematodes, weed seeds
and soil pathogens. The goal of a 2-year fal-
low period should be pursued but growers may
reduce time to replant based on economic
pressures on the available land. Several months
before planting (e.g. September before an
April planting), tree rows are laid out and sod
 Orchard Floor Management
is killed with a post-emergence, non-selective
and non-residue herbicide (e.g. glyphosate).
The killed sod can improve growth of newly
planted peach trees due to increased organic
matter and water penetration, while acting as
a mulch to suppress weeds (Welker and
Glenn, 1988, 1990; Glenn and Welker, 1989a)
(Fig. 14.2/Plate 88). Planting peach trees in
killed sod increased growth by 120% and
fruit yield by 160% during the first 3 years
after planting, compared with vegetation-free
strips maintained by cultivation or herbicides
(Glenn and Welker, 1989a). Often sod is killed
only where trees are to be planted so that liv-
ing grass remains as the foundation for drive
alleys. The width of the killed-sod area in tree
rows will strongly affect peach tree growth
and branch angle (Welker and Glenn, 1989,
1991). Tree size decreases as killed-sod width
is reduced below 2 m.
Proximity of living sod to the planted
peach tree can regulate competition and sub-
sequently affect tree size and yield (Welker
and Glenn, 1989, 1991). Newly planted peach
trees were significantly dwarfed by 1 m and
not by 3 m vegetation-free strips. Dwarfed
trees were as efficient (i.e. yield per unit trunk
cross-sectional area) as large trees. ‘Cultural
Fig. 14.2. Trees planted in cultivated (left) and killed-sod (right) strips. Killed sod provides weed
suppression and improves water penetration.
343
dwarfing’ of peach trees by sod competition
may enable growers to increase tree density
and yield per hectare with trees that require
less pruning. Yield efficiency of peach trees
increased as the size of vegetation-free ground
area increased to 9 m2 and then yield efficiency
remained constant, suggesting that closer tree
spacing within a row can increase yield per
hectare (Welker and Glenn, 1989). Reduced
vegetation-free ground area has reduced sprout
growth and pruning needed to maintain tree
size (Glenn and Welker, 1996). All ground
covers are not equally competitive and selection
of a ground cover will influence the dwarfing
effect on planted peach trees. Planting peach
trees into subterranean clover (T. subterraneum
L.) resulted in reduced growth and leaf N, P
and K compared with trees planted in a
herbicide-treated strip. However, tree growth
recovery was observed in the second year
after planting (Stasiak and Rom, 1991).
Young peach orchards may require up to
three seasons until they bear fruit. During this
establishment time, crops such as potatoes,
strawberries and other vegetables can be inter-
planted within row middles to provide income
and offset initial expenditures. For example,
up to four seasons of horticultural crops were
344 T.J. Tworkoski and D.M. Glenn
harvested prior to the first year of commercial
peach production (Leuty, 2003). Intercrop-
ping requires careful management, is labour-
intensive and is not amenable to moderate- or
high-density plantings that are commonly
used. In addition, pesticide drift from one
crop to another that it is not registered for is a
potential problem. Finally, the tree crop can be
inadvertently damaged while managing or har-
vesting the crop planted in the row middle.
Orchard floor composition of the
established orchard
Grass is often used as permanent ground
cover in drive alleys because it is amenable to
management and harbours fewer pests than
broadleaved ground covers. Although grass
competition severely inhibits growth of newly
planted peach trees, permanent sod in drive
alleys of established trees is often less debili-
tating (Hill, 1962). The amount of competition
can be managed based on the species of grass
used, the size of the vegetation-free area within
a tree row, suppressive treatment of the grass,
irrigation and fertilization.
Some grass cultivars (e.g. ‘K-31’ tall fes-
cue) are highly competitive with peach trees
and less competitive cultivars have been rec-
ommended for orchard floor cover. Used as a
permanent and complete ground cover, orchard
grass reduced peach yield by up to 37% but
‘Linn’ perennial ryegrass did not reduce yield
in 8-year-old peach trees (Tworkoski and
Glenn, 2001). Other non-competitive grasses
have been recommended as suitable ground
covers (see section on ‘Management of ground
covers, row middles and drive alleys’ above;
Willmott et al., 2000). Ground covers other
than grass have been recommended for apple
(Vossen and Ingals, 2002) but more research is
needed to determine benefits of forbs (herba-
ceous plants, excluding grasses) ground cover
in peach. For example, common vetch has
extrafloral nectaries on the stipules, which
may provide nectar to beneficial insects.
Legumes and weeds are often controlled
with herbicides although other techniques
have been used (see ‘Weed effects’ above). Site
conditions influence availability of nutrients
and water and pest threats. These environ-
mental conditions can be used to construct
models that determine action thresholds for
vegetation control. Weed suppression was
more critical early in the season rather than
late in the season for apple production (Merwin
and Ray, 1997). In North Carolina, MacRae
et al. (2007) noted that when the orchard floor
was kept weed-free with paraquat during the
first 12 weeks after bloom for peach, fruit
number, size and total yield were greater than
for weed-free periods of shorter duration.
Vegetation-free tree rows in summer with
control of cover crops
Ground cover can positively affect orchard pro-
ductivity and sustainability. As previously dis-
cussed, ground covers may provide a storage
pool of nutrients that can be carried from one
growing season to the next. In tree rows,
early-season herbicide applications allowed
N mineralization to begin. Grass cover that
incorporated herbicide application for no-till
control of ground covers may also be used to
increase availability of Ca, Mg, K and P (Haynes
and Goh, 1980). Grass cover can reduce leaching
of Ca, Mg, K, P, NH+4 and NO –3 and management
of grass, by frequent mowing, can contribute
significantly to mineral cycling and nutrient
availability within an orchard (Haynes, 1980).
Ground covers in drive alleys can be mowed,
possibly chopped, and transferred to tree rows
as mulch for weed suppression. However, large
amounts of biomass may be needed to control
weeds successfully (Elmore et al., 1993).
One ground cover management strategy
is to plant seed mixtures in late summer or
autumn with one or two mowings in late winter
or spring to control ground cover height.
Ground covers are then killed by mowing
close to the ground (e.g. legumes) and/or by
tilling and incorporating ground covers into
the soil the following spring. In Europe, late-
season ground covers included fodder radish
(Raphanus sativus v. olieferus), turnip (Brassica
rapa v. rapa), Phacelia (Phacelia tenacetifolia)
and winter rye (Secale cereale) (Bloksma and
Jansonius, 2002). In California, legume and
legume/grass blends have also been sown in
late summer, grown in autumn and winter, and
mechanically killed with shallow soil disking
or close mowing in the next growing season
(Vossen and Ingals, 2002). The planted ground
 Orchard Floor Management
covers can be used as ‘green manure’ to sup-
press winter weeds and add N and organic
matter to soil. If the mowed ground cover is
not incorporated into soil it can be used as a
mulch to suppress weeds in tree rows but
fewer nutrients are added to soil in the short
term. Total ground cover may require early
spring cultivation in orchards that are furrow,
flood or sprinkler irrigated.
In a review of orchard floor management
systems, Hogue and Neilsen (1987) deter-
mined that organic mulching in tree rows
combined with managed grass in drive alleys
would provide the most benefit for cropping
and for soil properties. However, costs, rodent
control and mulch availability must be consid-
ered. Significant increase in consumer demand
for reduced inputs of synthetic chemicals and
for increased organic production presents new
challenges for orchard floor management in
peach orchards. Research is needed to discover
and develop components for orchard floor
management that can serve organic production
systems. Components such as mechanical till-
age and non-competitive ground covers that
have been discussed will be useful, but addi-
tional tools are needed. Compatible cultural
and genetic components can be integrated in
the orchard ecosystem. Emergent traits of an
ecosystem, such as productivity, nutrient
cycling and nutrient loss to ground and sur-
face water, can be optimized in orchards under
organic and conventional management. Broad
comparisons between systems, including
energy inputs and extrinsic costs (e.g. CO2
emissions and topsoil loss), will eventually be
needed to critically compare management sys-
tems. In the near term, improved understand-
ing of nutrient dynamics and the relationships
among antagonistic or synergistic organisms,
and the manipulation of the orchard ecosystem
to achieve grower goals will be likely areas for
future progress in orchard floor management.
Vegetation-free tree rows year-round
In this management scheme drive alleys, not
tree rows, are vegetated in winter. Tree rows
may be kept vegetation-free year-round to
reduce competition and eliminate pest habitat.
Pre-emergence herbicide applied early in the
growing season effectively controls weeds.
345
Post-emergence herbicides, cultivation and
flaming may also be used effectively (see ‘Weed
management’ above; Table 14.1). Vegetation in
drive alleys can act as filter strips which foster
vertical infiltration of surface water, which
reduces pesticide runoff (Watanabe and Gris-
mer, 2001). Another benefit is that night-time
air temperatures can be increased in early
spring by radiant energy release from bare soil
to reduce chances for frost damage. Disadvan-
tages include surface compaction of soil, which
can reduce water infiltration and break down
soil structure in the absence of ground cover.
Permanent vegetation management
by mowing
Permanent vegetation beneath peach trees
requires management to prevent tall plants
from growing into the peach canopy and dis-
rupting orchard worker operations. The con-
cept is to maintain low vegetation by planting
ground covers with genetically based low
stature or by reducing ground cover height
by mowing. As young peach trees are very
susceptible to competition, this approach
seems more viable with older trees and irriga-
tion may be necessary. Allelopathic interac-
tions between ground cover and peach trees
must also be monitored (Weller et al., 1985). In
addition, pest problems and tree damage
from mowing can result. In New Zealand,
research identified dichondra, hard fescue
and creeping red fescue as shallow-rooted, low-
growing ground covers that were dense and
could suppress new weed growth beneath
fruit crops (Harrington et al., 2000a,b). Addi-
tional weed control, such as mowing or
selective herbicides, was needed to control
perennial weeds that could grow through
these ground covers. Benefits of improved soil
quality and reduced herbicide use may justify
permanent vegetation beneath peach trees in
some applications. Also, permanent sod may
recycle NO –3 near the soil surface and reduce
NO –3 leaching that can pollute ground water
(Wiedenfeld et al., 1999). White clover (Trifo-
lium repens) mixed with sod may contribute N
to soil after mowing and subsequent mineral-
ization of organic matter (Bloksma, 2000).
Supplemental fertilizer may be needed in early
346 T.J. Tworkoski and D.M. Glenn
spring and summer if organic matter pools
are small and the dynamics of nutrient move-
ment in this system have yet to be clarified.
Shallow-rooted grasses such as Kentucky
bluegrass (P. pratensis L.), annual bluegrass (Poa
annua L.), fescue (F. elatior L.) and orchardgrass
(D. glomerata L.) deplete less moisture from an
orchard than deep-rooted sods (Skroch and
Shribbs, 1986; Hogue and Neilsen, 1987). A
variety of grasses and legumes can be used in
California apple orchards that manage the
understorey with mowing or cultivation
(Vossen and Ingals, 2002). Similar species may
have use in peach orchards. Combination of
low ground cover or mulches with weed con-
trol (e.g. flaming, mowing, natural product her-
bicides) merits closer scrutiny for organically
acceptable practices (Weibel and Haseli, 2003).
14.4 Orchard Floor Influences
Effects of management systems on the
environment
Ground cover can affect heat transfer and
energy relationships in orchards (Snyder and
Connell, 1993). Reducing vegetation allows
greater absorption of solar energy by soil and
increases radiant heating from the soil in the
night-time. The resulting increase can help
prevent freeze injury of flowers in spring. The
range and fluctuation in soil temperature can
affect root growth rates and frost injury to
roots. Permanent sod cover of creeping red
fescue in row middles reduced fluctuations in
daily soil temperatures and provided some
frost protection compared with a system of
clean cultivation in summer and temporary
ground cover in winter (Tan and Layne, 1993).
Irrigation lowered soil temperature in sum-
mer with evaporative cooling, and elevated
soil moisture persisted to ameliorate soil tem-
peratures in winter.
Soil water availability to peach trees was
increased when cultivated for two or more
years compared with trees grown with per-
manent sod cover (Kenworthy, 1953). How-
ever, in the long run (25 years) sod cover
improved soil water penetration and holding
capacity compared with long-term cultivation.
Research demonstrated that this long-term
effect was in part due to increased rainfall
capture (less runoff) and improved soil prop-
erties (increased aggregate stability, macro-
porosity and microbial respiration) (Welker
and Glenn, 1988; Glenn and Welker, 1989b).
Orchard floor management interactions
with the peach tree
Orchard floor management will affect peach
tree size. Obviously, nutrient availability will
affect growth and yield but it can also influ-
ence growth in more subtle ways. Root den-
sity of young peach trees will decrease in sod
(Glenn and Welker, 1991; Parker and Meyer,
1996). When maintained in weed-free alleys,
trees exploit more of the grassed areas as they
age (Atkinson, 1980). Competition might be
manipulated, perhaps to the growers’ advan-
tage, to affect branch angle, excessive vegeta-
tive growth and possibly tree architecture.
Reduced peach root growth resulting from
competing ground cover may alter root-
produced signals, which can affect shoot
development and tree architecture. Grass
competition altered both dry weight and N
partitioning within branches of a 3-year-old
peach tree; the proportion of the N and mass
partitioned into fruit decreased as the size of
the vegetation-free area decreased (Tworkoski
et al., 1997). In contrast, the proportion of N
and mass partitioned into stem and leaves
increased or were unaffected as the size of the
vegetation-free area decreased. The implica-
tion of these findings is that peach yield may
be more sensitive than vegetative shoot
growth to increased grass competition.
Young peach trees can be dwarfed and
growth of mature peach trees reduced when
grown for several years with grass competi-
tion (Tworkoski, 2000; Tworkoski and Glenn,
2001) (Fig. 14.3/Plate 89). However, peach
tree size (mass, trunk diameter and crown size)
influenced shoot regrowth following pruning
more than grass competition (Tworkoski, 2000).
In this case, ground covers appeared to deprive
peach trees of soil nutrients by exploiting the
upper soil. Fruit yield was reduced when
peach trees are dwarfed by competition from
sod in drive alleys but the yield efficiency based
 Orchard Floor Management 347

Fig. 14.3. ‘Jersey Dawn’ and ‘Redskin’ peach trees, the same age (approximately 3 years after plant-
ing) but some planted and grown in 0.6 m (small trees in foreground) and others grown in 2.4 m (large
trees) vegetation-free areas with ‘K-31’ fescue in drive alleys.

on tree size (kg yield/cm2 trunk area) and include less evident gains from soil and water
water use (kg yield/cm water use plus pre- conservation and from biological regulation
cipitation) was not changed (Glenn and Welker, of pest populations. Such economic and envi-
1996). High-density plantings might, there- ronmental benefits should be incorporated
fore, be attained by dwarfing peach trees with into future models that assist with orchard
grass competition. The cumulative yield of floor management decisions.
such plantings has not yet been determined.
Orchard floor management of peach has
undergone a significant change from complete Disclaimer
mechanical control of vegetation to manage-
ment of temporary and permanent vegetation
Mention of trade names or commercial
with mechanical and chemical techniques.
products in this book chapter is solely for the
The future may incorporate ground covers
purpose of providing specific information and
that can be managed with environmentally
does not imply recommendation or endorse-
appropriate techniques for weed control that
ment by the US Department of Agriculture.
enhance soil fertility and stability. These tech-
niques will almost certainly be incorporated
with information-based technologies and with
computer models to help establish economic Acknowledgement
action thresholds for weed, water and nutri-
ent management decisions. Management The authors thank Drs Fumiomi Takeda, Mark
decisions are often based on increases in Brown and Mike Parker for critical reviews
yield and cosmetic quality, but decisions can that improved this book chapter.
348 T.J. Tworkoski and D.M. Glenn

References

Alston, D.G. (1994) Effect of apple orchard floor vegetation on density and dispersal of phytophagous and
predaceous mites in Utah. Agriculture, Ecosystems, and Environment 50, 73–84.
Ames, G.K. and Kuepper, G. (2004) Tree fruits: organic production overview. Horticulture Systems Guide.
http://www.attra.org/attra-pub/fruitover.html (accessed August 2007).
Atanassov, A., Shearer, P.W., Hamilton, G. and Polk, D. (2002) Development and implementation of a reduced
risk peach arthropod management program in New Jersey. Horticultural Entomology 95, 803–812.
Atkinson, D. (1980) The distribution and effectiveness of the roots of tree crops. Horticultural Reviews 2,
424–490.
Atkinson, D. and White, G.C. (1981) The effects of weeds and weed control on temperate fruit orchards and
their environment. In: Thresh, J.M. (ed.) Pests, Pathogens, and Vegetation. Pitman Publishing, Marshfield,
Massachusetts, pp. 415–428.
Belding, R.D., Majek, B.A., Lokaj, G.R.W., Hammerstedt, J. and Ayeni, A.O. (2003) Orchard floor preparation
did not affect early peach tree performance on Aura sandy loam soil. HortTechnology 13, 321–324.
Biggs, A.R., Baugher, T.A., Collins, A.R., Hogmire, H.W., Kotcon, J.B., Glenn, D.M., Sexstone, A.J. and Byers,
R.E. (1997) Growth of apple trees, nitrate mobility and pest populations following a corn versus fescue
crop rotation. American Journal of Alternative Agriculture 12, 162–172.
Bloksma, J. (2000) Soil management in organic fruit growing. Symposium organic fruit HRI-EMRA-ADAS-Wye
College, Ashford (Kent, UK). http://www.louisbolk.nl (accessed January 2004).
Bloksma, J. and Jansonius, P. (2002) Undergrowth of late summer sowings at the tree strip. In: Proceedings of
the 10th International Conference on Cultivation Technique and Phytopathological Problems in Organic
Fruit-Growing and Viticulture. Louis Bolk Instituut, Driebergen, The Netherlands, pp. 185–186.
Brown, M.W. (2001) Flowering ground cover plants for pest management in peach and apple orchards. Inte-
grated Fruit Protection 24, 379–382.
Brown, M.W. (2002) Are flowering plants taboo in peach orchards? Acta Horticulturae 592, 659–662.
Brown, M.W. and Glenn, D.M. (1999) Ground cover plants and selective insecticides as pest management
tools in apple orchards. Horticultural Entomology 92, 889–905.
Brown, M.W. and Tworkoski, T. (2004) Pest management benefits of compost mulch in apple orchards. Agri-
culture, Ecosystems, and Environment 103, 465–472.
Brown, M.W., Niemczyk, E., Baicu, T., Balazs, K., Jarosik, V., Jenser, G., Kocourek, F., Olszak, R., Serboiu, A.
and van der Zwet, T. (1997) Enhanced biological control in apple orchards using ground covers and se-
lective insecticides: an international study. Zahradnictvi 24, 35–37.
Butler, J.D. (1986) Grass interplanting in horticulture cropping systems. HortScience 21, 394–396.
Chalmers, D.J., Mitchell, P.D. and van Heek, L. (1981) Control of peach tree growth and productivity by regu-
lated water supply, tree density, and summer pruning. Journal of the American Society for Horticultural
Science 106, 307–312.
Duffus, J.E. (1971) Role of weeds in the incidence of virus diseases. Annual Review of Phytopathology 9,
319–340.
Elmore, C.L., Smith, R.J., Weber, E. and Miller, R. (1993) Use of cover crops for weed management in grape
and tree fruits. HortScience 28, 495.
Elmore, C.L., Merwin, I. and Cudney, D. (1997) Weed management in tree fruit, nuts, citrus and vine crops.
In: McGiffen, M.E. (ed.) Weed Management in Horticultural Crops. ASHS Press, Alexandria, Virginia, pp.
17–29.
Fogle, H.W., Keil, H.L., Redit, W.H., Cochran, L.C. and Baker, H. (1965) Peach Production East of the Rocky
Mountains. USDA-ARS Agriculture Handbook No. 280. US Department of Agriculture, Washington,
DC.
Foshee, W.G., Goodman, R.W., Patterson, M.G., Goff, W.D. and Dozier, W.A. (1997) Weed control increases
yield and economic returns from young ‘Desirable’ pecan trees. Journal of the American Society for
Horticultural Science 122, 588–593.
Funt, R.C., Schmittgen, M.C. and Schwab, G.O. (1997) Raised beds and microirrigation influence peach
production. HortScience 32, 677–682.
Glenn, D.M. and Welker, W.V. (1989a) Cultural practices for enhanced growth of young peach trees. Journal
of Alternative Agriculture 4, 8–11.
Glenn, D.M. and Welker, W.V. (1989b) Orchard soil management systems influence rainfall infiltration. Journal
of the American Society for Horticultural Science 114, 10–14.
 Orchard Floor Management 349

Glenn, D.M. and Welker, W.V. (1991) Soil management affects shoot and root growth, nutrient availability,
and water uptake of young peach trees. Journal of the American Society for Horticultural Science 116,
238–241.
Glenn, D.M. and Welker, W.V. (1996) Sod competition in peach production: II. Establishment beneath mature
trees. Journal of the American Society for Horticultural Science 121, 670–675.
Granatstein, D. (2002) Tree Fruit Production with Organic Farming Methods. http://organic.tfrec.wsu.edu/
OrganicIFP/OrganicFruitProduction/OrganicMgt.PDF (accessed August 2007).
Halbrendt, J.M. (1993) Research report: evaluation of a ‘Smother crop’ for orchard weed control. Pennsylvania
Fruit News 73, 38–39.
Harrington, K., Rahman, A., Hartley, J. and James, T. (2000a) Ground covers for orchards – past research. The
Orchardist October, 54–57.
Harrington, K., Rahman, A., Hartley, J. and James, T. (2000b) Ground covers for orchards – present research.
The Orchardist November, 66–68.
Haynes, R.J. (1980) Influence of soil management practice on the orchard agro-ecosystem. Agro-Ecosystems
6, 3–32.
Haynes, R.J. and Goh, K.M. (1980) Seasonal levels of available nutrients under grassed-down, cultivated and
zero-tilled orchard soil management practices. Australian Journal of Soil Research 18, 363–373.
Hill, R.G. (1962) The effect of sod as a soil management practice upon the growth and yield of the peach.
Ohio Agricultural Experiment Station Research Bulletin No. 903.
Hogue, E.J. and Neilsen, G.H. (1987) Orchard floor vegetation management. Horticultural Reviews 9, 377–430.
Horton, D.R., Broers, D.A., Lewis, R.R., Granatstein, D., Zack, R.S., Unruh, T.R., Moldenke, A.R. and Brown,
J.J. (2003) Effects of mowing frequency on densities of natural enemies in three Pacific Northwest pear
orchards. Entomologia Experimentalis et Applicata 106, 135–145.
Hoyt, G.D. and Hargrove, W.L. (1986) Legume cover crops for improving crop and soil management in the
southern United States. HortScience 21, 397–402.
Ju, Z., Duan, Y. and Ju, Z. (1999) Effects of covering the orchard floor with reflecting films on pigment accu-
mulation and fruit coloration in ‘Fuji’ apples. Scientia Horticulturae 82, 47–52.
Katan, J. and DeVay, J.E. (eds) (1991) Soil Solarization. CRC Press Inc., Boston, Massachusetts.
Kenworthy, A.L. (1953) Moisture in orchard soils as influenced by age of sod and clean cultivation. Michigan
Quarterly Bulletin 35, 454–459.
Killian, J.C. and Meyer, J.R. (1984) Effect of weed management on catfacing damage to peaches in North
Carolina. Journal of Economic Entomology 77, 1596–1600.
LaRue, J.H. and Johnson, R.S. (1989) Peaches, Plums, and Nectarines: Growing and Handling for Fresh
Market. University of California Division of Agriculture and Natural Resources, Publication No. 3331.
University of California, Oakland, California, p. 105.
Layne, D.R., Jiang, Z. and Rushing, J.W. (2001) Tree fruit reflective film improves red skin coloration and
advances maturity in peach. HortTechnology 11, 234–242.
Layne, R.E.C. and Tan, C.S. (1988) Influence of cultivars, ground covers, and trickle irrigation on early growth,
yield, and cold hardiness of peaches on Fox sand. Journal of the American Society for Horticultural
Science 113, 518–525.
Layne, R.E.C., Tan, C.S. and Fulton, J.M. (1981) Effect of irrigation and tree density on peach production.
Journal of the American Society for Horticultural Science 106, 151–156.
Leuty, T. (2003) Intercropping trees and annual crops. Ontario Ministry of Agriculture and Food. http://www.
omafra.gov.on.ca/english/crops/facts/info_intercropping.htm (accessed August 2007).
MacRae, A.W., Mitchem, W.E., Monks, D.W., Parker, M.L. and Galloway, R.K. (2007) Tree growth, fruit size,
and yield response of mature peach to weed-free intervals. Weed Technology 21, 102–105.
Mathews, C.R., Bottrell, D.G. and Brown, M.W. (2004) Habitat manipulation of the apple orchard floor to
increase ground-dwelling predators and predation of Cydia pomonella (L.) (Lepidoptera: Tortricidae).
Biological Control 30, 265–273.
Meagher, R.L. and Meyer, J.R. (1990) Influence of ground cover and herbicide treatments on Tetranychus
urticae populations in peach orchards. Experimental and Applied Acarology 9, 149–158.
Merwin, I.A. and Ray, J.A. (1997) Spatial and temporal factors in weed interference with newly planted apple
trees. HortScience 32, 633–637.
Merwin, I.A., Stiles, W.C. and van Es, H.M. (1994) Orchard groundcover management impacts on soil physical
properties. Journal of the American Society for Horticultural Science 119, 216–222.
Merwin, I.A., Ray, J.A. and Curtis, P.D. (1999) Orchard groundcover management systems affect meadow vole
populations and damage to apple trees. HortScience 34, 271–274.
350 T.J. Tworkoski and D.M. Glenn

Meyer, J.R. (1984) Catfacing in peaches: effects of ground cover and surrounding vegetation. In: Proceedings
of the 43rd Annual Convention of the National Peach Council. National Peach Council, Columbia,
South Carolina, pp. 5–11.
Norris, R.F. (1986) Weeds and integrated pest management systems. HortScience 21, 402–410.
Parker, M.L. (2003) Growing peaches in North Carolina. http://www.ces.ncsu.edu/depts/hort/hil/ag30.html
(accessed August 2007).
Parker, M.L. and Meyer, J.R. (1996) Peach tree vegetative and root growth respond to orchard floor manage-
ment. HortScience 31, 330–333.
Pickel, C., Bentley, W.J., Hasey, J.K. and Day, K.R. (2002) Peach plant bugs. In: UC IPM Pest Management
Guidelines: Peach. UC ANR Publication 3454. http://www.ipm.ucdavis.edu/PMG/r602300511.html
(accessed August 2007).
Powell, C.A. and Forer, L.B. (1982) Reservoirs of tomato ringspot virus in fruit orchards. Plant Disease 66, 583–584.
Preusch, P.L. and Tworkoski, T.J. (2003) Nitrogen and phosphorus availability and weed suppression from
composted poultry litter applied as mulch in a peach orchard. HortScience 38, 1108–1111.
Preusch, P.L., Adler, P.R., Sikora, L.J. and Tworkoski, T.J. (2002) Nitrogen and phosphorus availability in com-
posted and uncomposted poultry litter. Journal of Environmental Quality 31, 2051–2057.
Rock, G.C. and Yeargan, D.R. (1973) Toxicity of apple orchard herbicides and growth-regulating chemicals to
Neoseiulus fallacies and twospotted spider mite. Journal of Economic Entomology 66, 1342–1343.
Skroch, W.A. and Shribbs, J.M. (1986) Orchard floor management: an overview. HortScience 21, 390–394.
Snyder, R.L. and Connell, J.H. (1993) Ground cover height affects pre-dawn orchard floor temperature.
California Agriculture 47, 9–12.
Stasiak, M.J. and Rom, R.C. (1991) Subterranean clover (Trifolium subterraneum L.) ground cover affects on
growth and foliar nutrients status of young peach (Prunus persica (L.) Batsch). HortScience 26, 77.
Sullivan, T.P., Crump, D.R., Wieser, H. and Dixon, E.A. (1990) Comparison of release devices for stoat (Mustela
erminea) semiochemicals used as montane vole (Microtus montanus) repellents. Journal of Chemical
Ecology 16, 951–957.
Sullivan, T.P., Sullivan, D.S., Hogue, E.J., Lautenschlager, R.A. and Wagner, R.G. (1998) Population dynamics
of small mammals in relation to vegetation management in orchard agroecosystems: compensatory re-
sponses in abundance and biomass. Crop Protection 17, 1–11.
Tan, C.S. and Layne, R.E.C. (1993) Irrigation and ground cover management effect on soil temperature in a
mature peach orchard. Canadian Journal of Plant Science 73, 857–870.
Tworkoski, T. (2000) Response of potted peach trees to pruning and grass competition. HortScience 35, 1209–1212.
Tworkoski, T. (2002) Herbicide effects of essential oils. Weed Science 50, 425–431.
Tworkoski, T.J. and Glenn, D.M. (2001) Yield, shoot and root growth, and physiological responses of mature
peach trees to grass competition. HortScience 36, 1214–1218.
Tworkoski, T. and Miller, S. (2001) Apple and peach orchard establishment following multi-year use of diuron,
simazine, and terbacil. HortScience 36, 1211–1213.
Tworkoski, T.J. and Young, R.S. (1990) Rate and time of triclopyr application to control Virginia creeper in a
peach orchard. HortScience 25, 443–445.
Tworkoski, T.J., Glenn, D.M. and Welker, W.V. (1997) Carbohydrate and nitrogen partitioning within one-year
shoots of young peach trees grown with grass competition. HortScience 32, 1174–1177.
Tworkoski, T.J., Welker, W.V. and Vass, G.D. (2000a) Soil residues following repeat applications of diuron,
simazine, and terbacil. Weed Technology 14, 191–196.
Tworkoski, T.J., Welker, W.V. and Vass, G.D. (2000b) Weed community changes following diuron, simazine,
or terbacil application. Weed Technology 14, 197–203.
Vossen, P. and Ingals, C. (2002) Orchard Floor Management. http://cesonoma.ucdavis.edu/HORTIC/orchard_
floor.pdf (accessed August 2007).
Watanabe, H. and Grismer, M.E. (2001) Diazinon transport through inter-row vegetative filter strips: micro-
ecosystem modeling. Journal of Hydrology 247, 183–199.
Weibel, F. and Haseli, A. (2003) Organic apple production – with emphasis on European experiences. In: Ferree,
D.C. and Warrington, I.J. (eds) Apples: Botany, Production and Uses. CAB International, Cambridge,
Massachusetts, pp. 551–583.
Welker, W.V. (1984) The effects of oryzalin alone and in combination with diuron and simazine on young
peach trees. HortScience 19, 824–826.
Welker, W.V. and Glenn, D.M. (1988) Growth responses of young peach trees and changes in soil character-
istics with sod and conventional planting systems. Journal of the American Society for Horticultural Sci-
ence 113, 652–656.
 Orchard Floor Management 351

Welker, W.V. and Glenn, D.M. (1989) Sod proximity influences the growth and yield of young peach trees.
Journal of the American Society for Horticultural Science 114, 856–859.
Welker, W.V. and Glenn, D.M. (1990) Peach tree growth as influenced by grass species used in a killed-sod
planting system. HortScience 25, 514–515.
Welker, W.V. and Glenn, D.M. (1991) Growth and response of young peach trees to distribution pattern of
vegetation-free area. HortScience 26, 1141–1142.
Weller, S.C., Skroch, W.A. and Monaco, T.J. (1985) Common bermudagrass (Cynodon dactylon) interference
in newly planted peach (Prunus persica) trees. Weed Science 33, 50–56.
Wiedenfeld, B., Fenn, L.B., Miyamoto, S., Swietlik, D. and Marlene, C. (1999) Using sod to manage nitrogen
in orchard floors. Communications in Soil Science and Plant Analysis 30, 353–363.
Willmott, J., Frecon, J. and Cowgill, W. (2000) Turfgrass for Orchard and Nursery Floor Management. Rutgers
Cooperative Extension Fact Sheet FS319. Rutgers University, New Brunswick, New Jersey.
Wooldridge, J. and Botha, J.H. (1991) Observations on orchard floor management practices: implications for
integrated pest management. Deciduous Fruit Grower 41, 296–298.
Zehr, E.I., Lewis, S.A. and Bonner, M.J. (1986) Some herbaceous hosts of the ring nematode (Criconemella
xenoplax). Plant Disease 70, 1066–1069.
Zehr, E.I., Aitken, J.B., Scott, J.M. and Meyer, J.R. (1990) Additional hosts for the ring nematode, Criconemella
xenoplax. Journal of Nematology 22, 86–89.
 15 Diseases of Peach Caused by Fungi and
Fungal-like Organisms: Biology, Epidemiology
and Management

J.E. Adaskaveg1, G. Schnabel2 and H. Förster3

1Department
of Plant Pathology, University of California at Riverside, Riverside,
California, USA
2Department of Entomology, Soils, and Plant Sciences, Clemson University, Clemson,

South Carolina, USA

3Department of Plant Pathology, University of California, Davis, California, USA

15.1 Introduction 353

15.2 Blossom, Foliage and Fruit Diseases 353
Brown rot 353
Jacket rot and green fruit rot 360
Shot hole 361
Scab 363
Rust 365
Peach leaf curl 368
Powdery mildew 368
15.3 Trunk and Scaffold Diseases 373
Silver leaf disease 373
Leucostoma (Cytospora) canker 375
Fungal gummosis 378
Constriction canker 380
15.4 Root and Crown Diseases 383
Phytophthora root and crown rots 383
Armillaria root rot 386
Verticillium wilt 392
Peach tree short life 393
15.5 Postharvest Diseases 396
Brown rot and grey mould 396
Rhizopus rot, Gilbertella and Mucor decays 396
Sour rot 398
Management of postharvest decays 400
15.6 Other Diseases of Peach and Nectarine 401
15.7 Concluding Remarks 401

352 © CAB International 2008. The Peach: Botany, Production and Uses
(eds D.R. Layne and D. Bassi)
 Diseases Caused by Fungi
15.1 Introduction
A large number of fungal pathogens attack
the blossoms, foliage, fruit, branches, trunks
and roots of peach and nectarine trees. The
diseases covered in this chapter are consid-
ered important diseases worldwide and are
also found on other stone fruit crops. Most of
them occur preharvest in the orchard and
determine the overall productivity and fruit
quality of the orchard during its lifespan,
whereas others are major postharvest decays
that have a tremendous annual economic
impact after season-long inputs for produc-
tion. Of the preharvest diseases discussed,
some consistently cause annual losses in
production. Others occur more sporadically
and develop under specific climatic conditions.
All of the diseases included in this discus-
sion are caused by microorganisms that are
heterotrophic and require water for growth.
The organisms belong to two kingdoms, the
True Fungi and the Stramenopila (Chrom-
ista). Both kingdoms previously were grouped
together as ‘Fungi’ because of similarities in
growth and mode of nutrient uptake, and
because members of both kingdoms repro-
duce by spores. Members of the True Fungi
discussed here belong to the phyla Zygomycota,
Basidiomycota and, most importantly, Ascomy-
cota. The phyla are characterized by the specific
ways that sexual spores (zygospores, basidi-
ospores and ascospores, respectively) are pro-
duced. These spores may be formed in fruiting
structures called basidiomes (i.e. Basidiomy-
cota) or ascoma (i.e. Ascomycota). Ascoma of the
Ascomycota range from cup-shaped (apothe-
cia) to flask-shaped (perithecia) to spherical
(cleistothecia, chasmothecia) bodies or are in
cavities embedded in fungal tissue (ascostroma).
In addition, asexual spores (sporangiospores
in the Zygomycota, conidia in the Ascomycota
and sometimes in the Basidiomycota, or somatic
spores called chlamydospores for any group)
may be formed. Dense survival structures
formed by fungal mycelium (i.e. sclerotia) or
by mycelium and host tissue (i.e. pseudoscle-
rotia) may also be present. Commonly within
the Ascomycota as well as other phyla, fungi
have two different names, one for the asexual
(anamorph) stage and another one for the
353
sexual (teleomorph) stage. Within the Strameno-
pila, the phylum Oomycota contains important
plant pathogens in the genus Phytophthora.
These organisms generally are soil-borne and
are adapted to wet environments, and thus
are less of a problem when soil water content
is managed. Organisms in the Oomycota pro-
duce asexual motile zoospores that are wall-less
and flagellate and are capable of swimming
towards roots using chemotaxis. Zoospores
form in a sac-like cell called a sporangium.
Oospores are sexually produced spores that
are surrounded by a thick wall. These cells,
in addition to asexually produced chlamy-
dospores, are survival structures.
Pathogens that infect roots are commonly
referred to as soil-borne organisms. Those
that occur above ground commonly have air-
borne propagules that have adapted to rain-
splash or wind dispersal. The purpose of this
chapter is to provide the peach and nectarine
horticulturalist with an overview of the major
fungal diseases including a description of
their distribution, the symptoms (description
of disease) and signs (structures of the patho-
gen) involved with each disease, critical eco-
logical and epidemiological information, and
important management concepts and prac-
tices (Table 15.1). Also included are represen-
tative disease cycles for the major fungal
groups, which visualize host–pathogen rela-
tionships over the growing season and illus-
trate the importance of the different fungal
infection propagules. More detailed informa-
tion on each of the diseases discussed can be
found in the references provided, including
some extensive book contributions (Snowdon,
1990; Ogawa and English, 1991; Ogawa et al.,
1995; Strand, 1999).
15.2 Blossom, Foliage and Fruit Diseases
Brown rot
Brown rot is a major fungal disease of all com-
mercially grown Prunus spp. in most regions
of the world and can result in extensive crop
losses (Batra, 1991). In areas with high rain-
fall, severe epidemics may occur in most years.
In more arid locations, losses may be serious

Table 15.1. Common fungal diseases of peach and nectarine and important information on the biology, epidemiology and management of each disease.
Disease Pathogen
Blossom, foliage
and fruit diseases
Brown rot Monilinia fructicola
Monilinia laxa
Monilinia fructigena
Jacket rot and Botrytis cinerea
green fruit rot Sclerotinia sclerotiorum
M. fructicola
M. laxa
Peach leaf curl Taphrina deformans
Powdery mildew Podosphaera pannosa
(= Sphaerotheca pannosa)
Podosphaera leucotricha
Podosphaera clandestina
Rust Tranzschelia discolor
Tranzschelia pruni-spinosae
Scab Fusicladosporium
carpophilum (= Cladosporium
carpophilum)
Shot hole Wilsonomyces carpophilus Conidia
Trunk and scaffold
diseases
Leucostoma Leucostoma cincta
(Cytospora) canker (Cytospora cincta)
(syn. perennial canker) Leucostoma persoonii
(Cytospora leucostoma)
Silver leaf disease Chondrostereum
purpureum
Important reproductive Symptoms of economic
stages(s) for infectiona significance
Conidia, ascospores Fruit decay
Conidia, ascospores? Fruit decay
Conidia, ascospores Fruit decay
Conidia Fruit decay
Ascospores Fruit decay
Conidia, ascospores Fruit decay
Conidia, ascospores? Fruit decay
Bud conidia Defoliation, fruit off-grades
Conidia, ascospores Defoliation, fruit off-grades
Conidia Fruit off-grades
Conidia, ascospores Defoliation
Urediniospores Defoliation, fruit off-grades
(aeciospores?)
Urediniospores Defoliation, fruit off-grades
(aeciospores?)
Conidia, ascospores? Fruit off-grades
Defoliation, fruit off-grades
Conidia (ascospores?) Branch dieback, tree decline
and death
Conidia (ascospores?) Branch dieback, tree decline
and death
Basidiospores Branch dieback, tree decline
and death
354
Primary managementb
Sanitation, fungicides
Sanitation, fungicides
Sanitation, fungicides
Sanitation, fungicides
Orchard floor management, fungicides
Sanitation, fungicides
Sanitation, fungicides
J.E. Adaskaveg et al.
Fungicides
Cultivar selection, fungicides
Cultivar selection, fungicides
Cultivar selection, fungicides
Fungicides (removal of alternative host?)
Fungicides (removal of alternative host?)
Fungicides
Fungicides
Sanitation, pruning practices
Sanitation, pruning practices
Sanitation, pruning practices
Fungal gummosis Botryosphaeria dothidea Ascospores and conidia Branch dieback, tree decline Sanitation and winter pruning
(syn. peach blister (Botryosphaeria obtusa and and death (removal of alternative host?)
canker or ibokawa byo) Botryosphaeria rhodinia)
Constriction canker Phomopsis amygdali Conidia Branch dieback, tree decline Cultivar selection, sanitation and winter
and death pruning, use of low-N fertilization and
broad-spectrum fungicides

Root and crown

diseases
Armillaria root rot Armillaria mellea Rhizomorphs, mycelium Root and crown rot, Site selection (avoidance), sanitation,
tree decline and death soil fumigation
Armillaria ostoyae Rhizomorphs, mycelium Root and crown rot, Site selection (avoidance), sanitation,
tree decline and death soil fumigation
Armillaria tabescens Mycelium (rhizomorphs?) Root and crown rot, Site selection (avoidance), sanitation,

Diseases Caused by Fungi

tree decline and death soil fumigation
Peach tree short life Pseudomonas spp. Bacterial cells Tree decline and death Site selection (avoidance),
soil fumigation, rootstocks
Cytospora spp. Conidia (ascospores) Branch dieback, Sanitation, pruning practices
tree decline and death
Phytophthora root Phytophthora cactorum Zoospores, Root and crown rot, Irrigation management (e.g. raised beds,
and crown rots chlamydospores? tree decline and death sprinkler shields to prevent trunk
wetness), rootstocks, fungicides
Phytophthora cambivora Zoospores, Root and crown rot, Irrigation management, rootstocks,
chlamydospores? tree decline and death fungicides
Phytophthora cinnamomi Zoospores, Root and crown rot, Irrigation management, rootstocks,
chlamydospores tree decline and death fungicides
Phytophthora citricola Zoospores Root and crown rot, Irrigation management, rootstocks,
tree decline and death fungicides
Phytophthora citrophthora Zoospores, Root and crown rot, Irrigation management, rootstocks,
chlamydospores? tree decline and death fungicides
Phytophthora cryptogea Zoospores Root and crown rot, Irrigation management, rootstocks,
tree decline and death fungicides
Phytophthora drechsleri Zoospores, Root and crown rot, Irrigation management, rootstocks,
chlamydospores tree decline and death fungicides

(Continued)

355
 356
Table 15.1. continued

Important reproductive Symptoms of economic

Disease Pathogen stages(s) for infectiona significance Primary managementb

Phytophthora parasitica
Phytophthora syringae
Verticillium wilt Verticillium dahliae
Zoospores, Root and crown rot, Irrigation management, rootstocks,
chlamydospores tree decline and death fungicides
Zoospores Root and crown rot, Irrigation management, rootstocks,
tree decline and death fungicides
Microsclerotia Wilt Site selection (avoidance), soil fumigation

Postharvest diseases

J.E. Adaskaveg et al.

Brown rot M. fructicola Conidia Fruit decay Temperature management, sanitation,
fungicides
M. laxa Conidia Fruit decay Temperature management, sanitation,
fungicides
M. fructigena Conidia Fruit decay Temperature management, sanitation,
fungicides
Grey mould B. cinerea Conidia Fruit decay Temperature management, sanitation,
fungicides
Gilbertella decay Gilbertella persicaria Sporangiospores Fruit decay Temperature management, sanitation,
fungicides
Mucor decay Mucor piriformis Sporangiospores Fruit decay Temperature management, sanitation,
Mucor spp. fungicides
Sour rot Geotrichum candidum Arthroconidia Fruit decay Temperature management, sanitation
aPropagules with a question mark have a limited or unknown role in the disease cycle.
bManagement practices with a question mark have a limited or poorly defined role in controlling the disease.
 Diseases Caused by Fungi
when environmental conditions are condu-
cive for the pathogen to produce blossom
blight and quiescent infections on developing
fruit or brown rot of mature fruit when rains
occur at harvest time. The disease is caused
by three species of the genus Monilinia that
can be differentiated by their cultural charac-
teristics (Byrde and Willetts, 1977). Monilinia
fructicola Winter (Honey), the cause of North
American brown rot, is the main pathogen on
peaches and other stone fruits in most regions,
except in Europe, where it is not found (Batra,
1991). Monilinia laxa (Aderh. & Ruhl.) Honey
causes European brown rot on peaches and
other stone fruits worldwide, but generally is
less important on peaches in areas where
M. fructicola is also present. The third species,
Monilinia fructigena Honey in Whetzel, is not
present in North America. In Europe, its pri-
mary distribution area, it mainly occurs on
pome crops and only occasionally on peach
and nectarine.
The brown rot disease cycle includes
blossom and twig blights and, most economi-
cally important, pre- and postharvest fruit
decay (Fig. 15.1). Primary inoculum sources in
the spring are overwintering brown rot fruit
mummies on the tree, which produce asexual
conidia in sporodochia, and fruit mummies
on the orchard floor, which produce sexual
fruiting structures (apothecia) and spores
(ascospores) (Biggs and Northover, 1985).
Twig cankers and infected fruit peduncles can
be additional sources of inoculum (Sutton
and Clayton, 1972; Biggs and Northover,
1985). Blossom blight rarely reduces the crop
load, but blighted blossoms and infected
shoots provide secondary inoculum for fruit
infections later in the season. Infections can
occur over a wide temperature range with an
optimum of 22.5–25°C (Biggs and Northover,
1988a; Tamm and Flückiger, 1993). Minimum
wetness durations of 3–5 h are required at
20°C, whereas 18 h are needed at 10°C.
The first symptoms of blossom blight
(Fig. 15.2/Plate 90) are necrosis of the anthers
and browning of the filaments that proceeds
to the floral tube, ovary and peduncle. Infec-
tions may extend into the twig, which may be
girdled. As infected flowers wilt and turn
brown, they generally firmly stick to the twig
in a gummy mass. In wet weather, infected
357
flowers become covered with greyish to tan
sporodochia. Blossom infections by M. laxa,
and less commonly by M. fructicola, may
result in twig cankers that often show exten-
sive gum formation at the advancing margin.
The canker may girdle the twig, resulting in
blight of the distal twig with leaves turning
tan to brown and remaining attached. If the
branch is not girdled, surrounding healthy
tissue will produce callus. On large cankers,
sporulation may continue for several years.
Infection of fruit occurs by direct pene-
tration through the cuticle, through suture
cracks or other injuries, or indirectly through
stomata. Susceptibility of fruit to infection by
Monilinia spp. is high during the early stages
of fruit development, decreases during green
fruit stages and then increases again as fruit
mature and ripen (Biggs and Northover,
1988b; Gradziel, 1994). On mature or ripening
fruit, brown rot typically develops as a rap-
idly spreading, firm, brown decay (Fig. 15.3/
Plate 91). Under optimum conditions, decay
of ripe peaches infected by M. fructicola may
be visible within 48 h of infection. Quiescent
infections that occur on developing fruitlets
and on ripening fruit in less favourable envi-
ronments (arid and semi-arid climates) may
become active when fruit mature prior to or
after harvest. Molecular techniques have been
developed for the detection of quiescent fruit
infections of stone fruit and for species identi-
fication (Förster and Adaskaveg, 2000; Hughes
et al., 2000; Boehm et al., 2001; Côte et al., 2004).
Infections on green fruit that are injured by
frost or insects or are dropped to the orchard
floor, especially during late thinning, may
sporulate and provide additional inoculum.
Orchard sanitation practices that include
removal of mummified fruit and infected twigs,
as well as twigs with cankers, from trees are
important components of an integrated man-
agement approach. In some peach production
areas, removal of alternative hosts such as wild
Prunus spp. may also be an effective strategy.
No brown rot-resistant peach cultivars are
available, but there are considerable differ-
ences in susceptibility among peach (cling
and freestone) cultivars (Gradziel and Wang,
1993; Bassi et al., 1998). Protective fungicide
treatments provide the best control for both
blossom blight and fruit rot. The proper use
358 J.E. Adaskaveg et al.

Twig blight and Overwintering

rot of immature mummy on
and mature fruit ground

Conidia

Conidia

Gumming Overwintering
Blossom mummy on tree
blight

Ascus and Apothecia

ascospores

Fig. 15.1. Disease cycle of brown rot of peach caused by Monilinia fructicola. The fungal pathogen
overwinters in fruit mummies either on the tree or on the orchard floor. In spring, asexual conidia
produced from mummies on the tree and sexual ascospores produced in asci that are formed in
fruiting bodies (apothecia) on the ground are the primary inoculum. These spores are wind-dispersed
to susceptible host tissues (e.g. blossoms) and germinate under favourable wetness and temperature
conditions. Infections of blossom tissue can occur within 6–12 h at 15–20°C. After 3 to 5 days, blos-
soms become blighted. Diseased blossoms typically remain attached and the infection spreads into
the peduncle and down into the twig. The infection continues with the formation of a twig canker
that often develops a gumdrop as a host response. Conidia form on infected tissue and serve as
secondary inoculum for infection of immature (green) and mature fruit. Fruit infections may also
result in twig blight during severe outbreaks. (Drawing by J.E. Adaskaveg.)
 Diseases Caused by Fungi 359

Fig. 15.2. Brown rot blossom blight

of peach caused by Monilinia fructicola.
Infected blossom with gumming and
canker development at the base of the
peduncle.

Fig. 15.3. Brown rot fruit rot of peach caused by Monilinia fructicola.

of fungicides with local systemic activity pro- Blossom applications have to be done as sus-
tects flowers and fruit, reduces the amount of ceptible flower parts are exposed and before
sporulation formed on the infected tissue, and the occurrence of periods with conducive
reduces sources of overwintering inoculum. wetness and temperatures. Fungicides need
360 J.E. Adaskaveg et al.
not be applied to immature fruit unless wet-
ness conditions are especially favourable for
infection or injury caused by insects or cold
has increased the likelihood of disease. Thus,
insect control during this period may be an
important consideration. Management of
brown rot and several other pathogens is
heavily dependent on fungicides and, thus,
fungicide resistance management practices
are essential to prevent the development of
insensitive pathogen populations using the
newer single-site mode of action materials. In
some peach production areas, fungicide resis-
tance has developed in populations of M.
fructicola against the benzimidazoles and crop
losses have occurred (Ogawa et al., 1988).
More recently extensive shifts in effective
concentrations (EC50 values) of the demethy-
lation-inhibiting fungicides have also been
observed in the field (Zehr et al., 1999; Schna-
bel et al., 2004).
Jacket rot and green fruit rot
Decay of flower parts resulting in jacket rot
and subsequent green fruit rot can be a serious
problem in wet years or in foggy production
Fig. 15.4. Jacket rot of peach caused by Botrytis cinerea. The jacket (or shuck) is infected, but the
immature fruit is still disease-free.
areas with prolonged wetness periods during
bloom. This disease can affect all stone fruit
crops including peach and nectarine that are
the least susceptible. One or several fungi
including Monilinia spp., Botrytis cinerea Pers.:
Fr. (teleomorph Botryotinia fuckeliana (de Bary)
Whetzel) and Sclerotinia sclerotiorum (Lib.) de
Bary commonly cause jacket rot and green
fruit rot, depending on the geographical loca-
tion and the presence of the pathogens. All of
these fungi have a cosmopolitan distribution
and thus this disease presumably occurs
throughout temperate fruit-growing regions
of the world wherever wetness occurs during
the bloom period.
The disease begins with the pathogens
attacking the calyx (floral cup or later the
‘jacket’ or ‘shuck’ around the developing fruit),
petals and other flower parts, causing them to
wither (Fig. 15.4/Plate 92). As fruit develop
and come in contact with diseased blossom
parts, the pathogens can grow into the healthy
fruit. A brown lesion develops and spreads
quickly into the small or immature fruit (Fig.
15.5/Plate 93). B. cinerea is characterized by
greyish tufts of conidia on infected senescent
or dying plant tissues or on sclerotia on the
orchard floor. Germination of conidia occurs
 Diseases Caused by Fungi
Fig. 15.5. Green fruit rot of peach caused by Monilinia fructicola. The decay spreads to neighbouring
healthy fruit by contact.
with wetness over a wide temperature range
with an optimum of 15–20°C. Wetness is not
required if relative humidity is above 98%.
With S. sclerotiorum, white mycelium devel-
ops on infected fruit and blossom tissues; no
asexual spores are produced. Sexual spores of
S. sclerotiorum are produced from apothecia
that develop from sclerotia on the orchard
floor (Fig. 15.6/Plate 94), especially under
vegetation of cover crops. Ascospores are
forcibly discharged from the apothecia fol-
lowing changes in relative humidity or by
physical disturbance. Infection of floral parts
by ascospores requires 48–72 h of wetness
with temperature seldom being limiting. Scle-
rotia of this fungus only form after infected
structures fall to the ground. Monilinia species
can also cause green fruit rot after partial col-
onization of blossom tissue or after causing
brown rot blossom blight. Spores of the jacket
rot and green fruit rot fungi are wind-borne
or are disseminated by splashing rain and
germinate to infect the senescent blossom tis-
sues. From infections of blossom tissues the
pathogens can move into fruit tissue, pro-
vided that cool, wet weather occurs, which
361
delays the jacket from separating from the
developing fruit and allows fungal growth.
Removal of weeds and senescent plant
tissues from the orchard floor is probably
beneficial in reducing inoculum levels for
jacket rot and green fruit rot. Fungicide appli-
cations with materials effective against all
green fruit rot pathogens during bloom, espe-
cially full bloom, are suggested to prevent
losses.
Shot hole
Shot hole disease of stone fruits occurs world-
wide but is especially important in the peach-
producing areas of the western USA. It is
most important on peaches, nectarines and
apricots. Outbreaks of the disease on peach
occur following favourable environments
and when no protective fungicide treatments
were applied during the dormant season.
Shot hole is caused by the imperfect fungus
Wilsonomyces carpophilus (Lév.) Adaskaveg,
Ogawa & Butler, which was previously
362 J.E. Adaskaveg et al.
classified in the imperfect genera Coryneum,
Clasterosporium and Stigmina.
Symptoms of shot hole occur on twigs,
leaves and fruit (Wilson, 1937). The primary
damage to peach is the killing of twigs and
buds. In the absence of management prac-
tices, unsightly superficial fruit infections
render the crop unmarketable. On twigs,
where the fungus overwinters, lesions first
appear as purplish spots 2–3 mm in diameter
that enlarge up to 10 mm in diameter, turn
brown and are slightly sunken (Fig. 15.7/
Plate 95). In the light tan centre of the lesions,
asexual conidia of the fungus are produced in
sporodochia in the spring. These conidia are
the primary inoculum for new infections;
they are rain-splash dispersed to developing
blossoms and leaves. Conidia may remain
Fig. 15.6. Apothecia of Sclerotinia
sclerotiorum are produced on
over wintering sclerotia (bottom).
Ascospores are forcibly discharged
in a spore cloud. (Reprinted, with
permission, from Strand, 1999.)
viable for several months when kept dry.
Infected buds are another overwintering
stage of the fungus where primary inoculum
is produced in the spring and often through-
out the season. Scales of infected buds turn
dark and are sometimes covered with gummy
exudates. Lesions on leaves and fruit also
start out as small purplish spots that turn
brown and expand up to 10 mm in diameter.
During warm, dry weather, lesions on leaves
abscise to produce the typical shot hole symp-
toms. Early dehiscence mostly eliminates the
formation of sporodochia on shot hole lesions.
In cool, wet environments, however, lesions
remain attached to leaves and sporodochia
commonly develop in the centre of each
lesion. Infected leaves may drop but early
defoliation of peach trees is rare. On fruit,
 Diseases Caused by Fungi
lesions turn into raised, corky areas that do
not extend into the mesocarp tissues (Fig. 15.8/
Plate 96).
Conidia of the fungus germinate over a
wide temperature range. Twig infections
require at least 24 h of continuous wetness,
whereas shorter wetness durations are
required for leaf infections. Thus, at 20–25°C,
leaf infections occur after 8–12 h of wetness.
Epidemics of shot hole occur when high rain-
fall commonly occurs. For management of the
disease in semi-arid climates like in Califor-
nia, dormant fungicide sprays to protect
against twig and bud infections are applied in
the autumn after leaf drop and before winter
rains begin. In wetter climates, protective
fungicides also have to be applied at leaf
emergence and through fruit set.
363
Fig. 15.7. Lesion of the shot hole
fungus Wilsonomyces carpophilus on
peach twig, where the pathogen
overwinters. Sporodochia of the
fungus can be seen in the centre of
a stem lesion.
Scab
Scab is an important fungal disease of peaches,
nectarines and other Prunus species in warm
areas with high rainfall, such as the south-
eastern USA, or when orchards are irrigated
by overhead sprinkler. The disease also has
caused epidemics on peach in semi-arid areas
like California in years with unusually high
rainfall. Scab is caused by the fungus Venturia
carpophila E.E. Fisher. The sexual stage has
rarely been observed but the asexual stage,
Fusicladosporium carpophilum (Thüm.) Par-
tridge & Morgan-Jones (formerly Cladospo-
rium carpophilum Thüm.), is common.
Infections can occur on twigs, leaves and
fruit, but symptoms are most noticeable on fruit.
On twigs, where the fungus overwinters,
364 J.E. Adaskaveg et al.
superficial circular to oval lesions that are
slightly raised develop on new succulent
growth in the spring (Keitt, 1917; Bensaude
and Keitt, 1928). These lesions initially appear
as water-soaked spots, which darken with age
and become brown and later purple to dark
brown and have a raised border. At the end of
the season, lesions are oval in shape and
3 mm × 5 mm to 5 mm × 8 mm in size. They
persist throughout the host dormant period.
In the following spring, during periods with
relative humidity between 70 and 100%, the
fungus produces abundant primary inocu-
lum in the form of asexual conidia from oliva-
ceous tufts of mycelium within the lesion
(Lawrence and Zehr, 1982; Gottwald, 1983)
especially after petal fall until mid-spring.
Conidia are dispersed by wind and rain. They
germinate between 15 and 30°C with an opti-
mum temperature of 25–30°C (Lawrence and
Zehr, 1982). Maximum conidial germination
occurs in free water, but conidia can also ger-
minate at 94–100% relative humidity. The
Fig. 15.8. Symptoms of shot hole
caused by Wilsonomyces carpophilus on
peach fruit and leaf.
fungus does not survive in the twig lesions
for more than a second season due to bark
formation during twig growth. With unusu-
ally severe infections, new shoots of the host
may die back. On leaves, symptoms may
occur in early summer on the lower surfaces
and are visible as irregular, blotchy lesions
slightly darker in colour than the surround-
ing healthy tissue. Lesions turn olive green
once sporulation of the fungus occurs. Infec-
tions of leaves are rarely severe enough to
cause serious defoliation of trees. On fruits,
small circular spots develop mainly on the
upper, exposed surfaces (Fig. 15.9/Plate 97).
They are 5–10 mm in diameter, are first green
and turn olive to black once the fungus spo-
rulates and sometimes have a green to yellow
halo. When fruit are severely infected, lesions
coalesce and form large, superficial blotches
that make the fruit unmarketable. Although
scab on peach and nectarine fruit may develop
similar to bacterial spot caused by Xanthomo-
nas arboricola pv. pruni (Smith) Vauterin, in
 Diseases Caused by Fungi
most early-season infections bacterial spot on
fruit is visible as small, irregular, water-
soaked brown spots that may occur anywhere
on the fruit surface and later develop into
deep, cavernous-appearing lesions.
Management of scab is mainly accom-
plished with the use of fungicidal sprays that
are applied at shuck split and every 2 weeks
thereafter through early summer. In semi-arid
climates, effective management can also be
obtained by reducing inoculum dispersal and
leaf wetness from foliar sprinkler irrigation.
Sanitation by removing infected twigs is
impractical because large numbers of over-
wintering lesions persist on fruiting wood.
Pruning trees to allow adequate sunlight pen-
etration and unimpeded air movement may
improve scab management by facilitating
rapid drying and good fungicide coverage.
Rust
Peach rust occurs wherever stone fruit are
grown. Incidence of the disease in different
years and different peach production regions
is highly variable and epidemics occur in years
with excessive wetness during the growing
365
Fig. 15.9. Symptoms of scab caused
by Fusicladosporium carpophilum on
peach fruit.
season. Rust is caused by two species of the
biotrophic genus Tranzschelia, Tranzschelia dis-
color Tranz. & Lit. and Tranzschelia pruni-spino-
sae Pers., that differ in morphology of their
teliospores. T. discolor is found worldwide,
whereas T. pruni-spinosae is mainly found in
Europe and the central and eastern USA
(Dunegan, 1938). For T. discolor, different for-
mae speciales have been described on different
stone fruit hosts with strains on peach being
designated as T. discolor f. sp. persicae. Although
cross-infection of formae speciales among host
species occurs, virulence is generally reduced.
Tranzschelia spp. are macrocyclic, heteroecious
rusts that alternate between species of Prunus
and genera in the Ranunculaceae (buttercup
family). In mild climates, the fungus can sur-
vive on Prunus spp. without infecting the
alternative host. Because the uredinial stage
is capable of repeated infections on Prunus
spp., this host can be seriously damaged,
causing economic losses.
On peach, in addition to leaves and fruit,
the fungus can infect stem tissues, which are
important sources of primary inoculum in the
spring (Fig. 15.10/Plate 98). Leaf infections
develop as angular, yellow lesions with rusty-
brown urediniospore-producing pustules (ure-
dinia) on the lower leaf surfaces (Goldsworthy
366 J.E. Adaskaveg et al.

Fig. 15.10. Sporulating stem lesion of peach rust caused by Tranzschelia discolor.

Fig. 15.11. Symptoms of peach rust caused by Tranzschelia discolor on peach leaves.

and Smith, 1931) (Figs 15.11 and 15.12/Plates result in premature defoliation of the tree
99 and 100). Late in the growing season, ured- during autumn and stimulate flowering,
inia develop into telia that produce dark- which may reduce tree vigour or productivity
coloured teliospores. Heavy leaf infection can in the subsequent season. Symptoms on
 Diseases Caused by Fungi
immature fruit are green, circular lesions 2–3
mm in diameter (Fig. 15.13/Plate 101). On
mature fruit, lesions are sunken with yellow
halos and the mesocarp below the lesion is
discoloured. Current-year peach stems are
infected during outbreaks of the disease in
367
Fig. 15.12. Leaf lesions with uredinia
of peach rust caused by Tranzschelia
discolor.
Fig. 15.13. Symptoms of peach rust
caused by Tranzschelia discolor on
peach fruit.
late autumn and the fungus survives as myce-
lium in symptomless stems during the win-
ter. In early spring, infections are first visible
as water-soaked lesions. The epidermis then
becomes raised and ruptures with the devel-
opment of uredinia and urediniospores that
368 J.E. Adaskaveg et al.
function as primary inoculum for leaf infec-
tions. Later, as the stems grow in circumfer-
ence, lesions split open lengthwise along the
stems. Lesions will not continue to produce
spores in the following season because infec-
tions are then delimited by a wound perid-
erm. Stem lesions are superficial and are not
considered directly damaging to the tree.
On peach, both species of Tranzschelia
function as asexual fungi. Stages of the fun-
gus that occur on the alternative host have
not been observed in many locations and the
aecial stage generally is not considered an
important inoculum source. As described
above, the fungi overwinter as mycelium in
twigs, but may also survive as urediniospores
on twigs or as uredinia on non-abscised
leaves. Urediniospores germinate over a wide
temperature range (8–38°C; 13–26°C is opti-
mum) and require wetness or a saturated
atmosphere. At 20°C, 18 h of wetness are
required for adequate leaf infection. Thus, for
the development of epidemics, the presence
of viable spores and conducive periods of
wetness are needed.
For management of peach rust, preven-
tive applications of fungicides are done before
rain events during the spring. The emergence
of sporulating stem lesions as indicated dur-
ing monitoring programmes in the spring has
been used as a starting date for fungicide
applications (Soto-Estrada et al., 2003).
Peach leaf curl
Peach leaf curl is a cosmopolitan disease and
occurs wherever peaches and nectarines are
grown. Economic losses have been reported
historically, but crop losses in orchards treated
with fungicides are now negligible. In mild
climates the disease can consistently be
observed, but its occurrence is more erratic
in most other production areas.
Peach leaf curl is caused by Taphrina
deformans (Berk.) Tul. Symptoms occur mainly
on newly formed leaves in the spring (Fig.
15.14/Plate 102). Leaves first develop dis-
coloured areas that thicken and then become
wrinkled and puckered, causing the leaves
to curl. Infected leaves can have a range of
colours from light green to yellow, red and
purple (Fig. 15.15/Plate 103). They may be
covered with a subtle white layer of sexually
produced spore sacs (asci containing ascospores)
and then turn brown and generally abscise.
Defoliated trees will leaf out again, but fruit
set will be sparse in the current year and the
following season. Fruit infections are less
common. They are characterized by irregu-
lar, raised green (initially) or reddish lesions
(Fig. 15.16/Plate 104).
The fungus overwinters in the asexual,
yeast-like bud-conidia stage, which contami-
nates twigs and buds of the tree. Emerging
leaves are infected when the fungus changes
to a mycelial parasitic phase. Naked asci are
then produced on the leaf surface and the
sexual ascospores are forcibly discharged.
These spores germinate and form more bud-
conidia that contaminate twig tissues. Infec-
tions occur at temperatures of 10–21°C, but
ascospores and bud-conidia can survive hot,
dry conditions for several months. Periods of
cool, wet weather during early bud develop-
ment favour leaf curl disease. When tempera-
tures at early leaf development are high,
infections rarely become established.
Peach leaf curl can be managed with one
well-timed preventive fungicide application,
either in late autumn after 90% of the leaves
have fallen or in spring before bud swell.
Treatments after infection or symptom devel-
opment are ineffective. Sanitation and cul-
tural practices do not provide control against
this disease. Most peach and nectarine culti-
vars are susceptible to the disease, but there is
a wide range of susceptibilities. Vigour of
infected trees should be maintained by
irrigation and N fertilization management, as
well as reducing stress from crop load by
extra thinning.
Powdery mildew
Powdery mildew of peach and nectarine occurs
worldwide, but is most damaging in semi-
arid growing areas. The disease can be caused
by several different species of fungi that com-
monly occur on rosaceous plants (Yarwood,
1939). Three species have been reported on
 Diseases Caused by Fungi
peach, with Podosphaera pannosa (Wallr.:Fr.)
Braun & Takamatsu (formerly Sphaerotheca pan-
nosa (Wallr.:Fr.) Lév.) being the most important.
Podosphaera leucotricha (Ellis & Everh.) E.S.
Salmon is less common, and Podosphaera clan-
destina (Wallr.:Fr.) Lév. has been reported on
peach seedlings in the eastern USA, whereas P.
tridactyla has been observed on peach in south-
ern California (J.E. Adaskaveg and H. Förster,
unpublished). Fruit infections caused by P. pan-
nosa and P. leucotricha cause the most economic
damage, but leaf infections are important
sources of inoculum. In nurseries, powdery
mildew leaf infections can cause significant
damage to seedlings and small trees.
The susceptibility of peach and other stone
fruit crops varies greatly among cultivars. The
eglandular (without glands at the leaf base)
peach cultivars are more susceptible than the
glandular ones. Furthermore, in some cultivars,
tissues also vary in their susceptibility with
fruit being more or less susceptible than leaves,
369
Fig. 15.14. Peach leaf curl caused by
Taphrina deformans.
depending on the mildew species involved and
maturity of host tissue. Leaves, buds, green
shoots and fruit are commonly attacked, but
flower infections are rare. On twigs, mildews
are white and felt-like, even in the winter in
mild climates (Fig. 15.17/Plate 105). The first
symptoms on peach leaves infected by P. pan-
nosa are small, circular, white, web-like colo-
nies that become powdery once masses of
asexual conidia are produced in chains (Fig.
15.18/Plate 106). Leaves may then curl or
become stunted. Older infections commonly
result in leaf chlorosis and necrosis. Severe
infections may result in defoliation.
Fruit are susceptible from the early
stages of development until pit hardening on
peach, nectarine and plum, but not other Pru-
nus spp. (Weinhold, 1961). White circular spots
may enlarge, coalesce and cover large areas of
the fruit (Figs 15.19–15.21/Plates 107–109).
Infections usually result in some deformation
of the fruit surface with depressed or slightly
370 J.E. Adaskaveg et al.

Fig. 15.15. Advanced symptoms of peach

leaf curl caused by Taphrina deformans
with leaf deformation, discoloration,
necrosis and a subtle white layer of sexually
produced asci containing ascospores.

Fig. 15.16. Symptoms of peach leaf curl caused by Taphrina deformans on developing peach fruit.
 Diseases Caused by Fungi
Fig. 15.17. Peach twig with overwintering
mycelium and embedded chasmothecia of
powdery mildew caused by Podosphaera
pannosa.
raised areas. Infections on peach fruit become
necrotic after pit hardening, whereas on nec-
tarine and occasionally also on peach the tis-
sue remains green. Any fruit with blemishes
caused by powdery mildew are generally
unmarketable.
Based on indirect evidence, P. leucotricha
(mainly an apple pathogen) presumably is
involved in causing another powdery mildew
symptom on peach fruit known as ‘rusty spot’
(Daines and Trout, 1977, Ries and Royse,
371
1978). With this disease, small, circular, orange-
rusty lesions develop on the fruit, which
enlarge and may cover the entire fruit. No
symptoms occur on leaves and stems. Lesion
development has been related to rapid fruit
growth. Incidence of the disease increases from
shuck-fall stage of fruit development until 60
days after full bloom and epidemics typically
last from 17 to 30 days (Furman et al., 2003a).
P. pannosa overwinters as mycelium in
shoots and in the inner bud scales (Weinhold,
1961). In milder climates, young twigs may be
covered with dense mats of mycelium. Dur-
ing a recent winter in California we found for
the first time on peach the sexual fruiting bod-
ies (chasmothecia, formerly referred to as ery-
siphaceous cleistothecia or perithecia) of the
fungus, which were embedded in these myce-
lial mats (Fig. 15.22/Plate 110). In the spring,
newly developing leaves become diseased as
they emerge from infected buds. When chas-
mothecia are present, ascospores are released
that also serve as primary inoculum. Because
roses are an important host for the pathogen
and disease is not always managed on this
host, diseased roses can be major contributors
to the development of epidemics of peach
powdery mildew. Secondary infections by the
wind-disseminated, asexual conidia occur
throughout the growing season. Conidia ger-
minate between 2°C and 37°C, with an opti-
mum of 21°C. Conidia can germinate in free
water and at relative humidity of 43–100%.
Excessive durations of wetness will kill
conidia of powdery mildew fungi. During
periods with warm, humid conditions the
disease can quickly develop into an epidemic.
Management of powdery mildew is by
cultural practices and the use of protective
fungicide treatments. Less-susceptible culti-
vars should be planted in areas that commonly
have a high incidence of disease. To reduce
the relative humidity in the orchard, the fre-
quency of irrigation periods should be mini-
mized and low-angle sprinklers should be
used to keep foliage dry. Fungicide applica-
tions are done from calyx (i.e. shuck) fall until
the pit-hardening stage of fruit development
for peach and nectarine. Adequate manage-
ment of rusty spot was achieved with three to
five fungicide applications (Furman et al.,
2003b).
372 J.E. Adaskaveg et al.

Fig. 15.18. Powdery mildew caused by Podosphaera pannosa on peach leaves.

Fig. 15.19. Powdery mildew caused

by Podosphaera pannosa on developing
peach fruit.
 Diseases Caused by Fungi 373

Fig. 15.20. Powdery mildew caused by

Podosphaera pannosa on mature peach
fruit.

Fig. 15.21. Powdery mildew caused by Podosphaera pannosa on developing nectarine fruit.

15.3 Trunk and Scaffold Diseases
Silver leaf disease
Silver leaf disease of peach is caused by the
fungus Chondrostereum purpureum (Pers.:Fr.)
Pouz. The disease also affects a wide range of
other cultivated and non-cultivated hardwood
tree species, particularly willow and poplar.
Silver leaf disease has been reported from
most temperate-zone stone fruit production
areas and can also be a problem in nurseries.
374 J.E. Adaskaveg et al.
Fig. 15.22. Close-up of chasmothecia of powdery mildew caused by Podosphaera pannosa on peach
twig.
The fungus can also grow saprophytically on
tree logs and prunings.
Leaves of silver leaf-diseased trees become
silvery in appearance (Fig. 15.23/Plate 111),
which is most noticeable on new growth in
the spring when damage by other diseases
and pests does not obscure the symptoms.
These symptoms occur soon after the patho-
gen invades the woody tissues. Silver leaf
symptoms result from a toxin produced by
the pathogen that causes the upper epidermis
to separate from the palisade mesophyll layer
of the leaf. The separated layers then reflect
light differently from healthy tissue, giving
leaves the silvery appearance. Infected leaves
may then become necrotic and abscise. In the
wood of infected trees, two symptoms occur,
wood discoloration and white rot. Wood dis-
coloration in branches is often angular to pie-
shaped in cross-section, whereas in the trunk
the discoloration occurs in the older secondary
xylem (Fig. 15.24/Plate 112). In white rot, the
decayed wood becomes mottled to bleached
white and eventually spongy soft. Substantial
decay of the trunk may occur and extend into
the scaffold branches and into the roots.
Leathery fruiting bodies of the fungus may
develop in bracket-like clusters on tree trunks
and scaffold branches of living trees and on dead
wood. They produce sexual basidiospores
that are the only known propagules of the fun-
gus. They are wind-disseminated and function
as inoculum in disseminating the organism.
On peaches, fruiting bodies are not easily
found, although they are thought to persist for
up to 2 years, producing spores during warm,
moist environments at any time of the year.
Like most wood decay fungi, C. purpureum
needs a fresh wound that exposes wood of
the branch or trunk. Wind-disseminated spores
of the fungus are deposited on the moist wood,
germinate and quickly invade the xylem tis-
sues of the wood. The fungus grows over a
wide temperature range with an optimum of
25°C. Wood-exposing wounds are most sus-
ceptible in the first week after injury. Wound
healing, which prevents infection by the fun-
gus, progresses at different rates in different
climates and at different times of the year.
A wide range of perennial hosts, inocu-
lum production over a long period, the impos-
sibility of protecting all wounded surfaces
and the inability to eradicate established
infections from tree trunks make silver leaf a
difficult disease to control. Management prac-
tices should therefore focus on starting with
clean nursery stock and on preventing the
establishment of the fungus by minimizing
large wood-exposing wounds and using proper
pruning practices. Sanitation measures to
 Diseases Caused by Fungi
reduce inoculum include removing and burn-
ing infected wood. The most effective pruning
wound treatments are biocontrol agents such
as Trichoderma spp., which act by pre-colonizing
the wood and site-excluding the pathogen.
Leucostoma (Cytospora) canker
Leucostoma (Cytospora) or perennial canker is
an important disease of stone fruit crops
including peach and nectarine, especially in
colder climates. It is also associated with the
peach tree short life syndrome in the south-
eastern USA (see below) and is part of the
apoplexy disease complex on stone fruits in
Europe. The disease also occurs in the Pacific
Northwest of the USA, as well as in Canada,
South America and Japan. Leucostoma canker
375
Fig. 15.23. Symptoms of silver leaf
disease caused by Chondrostereum
purpureum on peach leaves: diseased
leaf (left) and healthy leaf (right).
reduces the number of bearing branches, kills
twigs and shortens tree life.
Leucostoma canker is caused by Leucos-
toma cincta (Fr.:Fr.) Höhn. (syn. Valsa cincta
(Fr.:Fr.) Fr.) (anamorph Cytospora cincta Sacc.)
and Leucostoma persoonii Höhn. (syn. Valsa leu-
costoma (Pers.:Fr.) Fr.) (anamorph Cytospora
leucostoma (Pers.:Fr.) Fr. These fungi produce
a compound fruiting body that consists of a
central pycnidium that produces hyaline
asexual conidia. These conidia ooze from the
pinhead-sized, black pycnidia to form long
orangish tendrils (cirri). Perithecia develop in
or under older stroma around the pycnidium.
In the perithecia, sexual spores (ascospores)
are formed.
The pathogens infect small and large
branches of trees (Fig. 15.25/Plate 113). Infec-
tions result in dieback that continues from
376 J.E. Adaskaveg et al.

Fig. 15.24. Cross-section through scaffold branch of peach infected by Chondrostereum purpureum.

Fig. 15.25. Peach tree with Leucostoma (Cytospora) or perennial cankers on scaffold branches.
(Reprinted from Ogawa et al., 1995, with permission of APS.)
 Diseases Caused by Fungi
apical branches to the main scaffold branches
and trunk. On small branches, lesions appear
as sunken, zonate, discoloured areas that
develop around dead buds or previous years’
leaf scars and are observed 2 to 4 weeks after
bud break (Fig. 15.26/Plate 114). Amber gum
may ooze from these lesions as they age and
darken unless the branch dies. On large
branches, scaffolds and trunks, conspicuous
elliptical cankers develop. Copious amounts
of gum may exude from the cankers and, with
age, the bark dries and cracks form, exposing
the blackened tissue beneath. If infections
follow winter injury or develop on branches
weakened from infections by other patho-
gens, gumming will not be associated with the
cankers. Cankers typically develop concen-
tric rings from alternation between canker
377
extension and callus formation during dor-
mant and growing seasons, respectively. Pyc-
nidia erupt directly through the bark within
the branch or trunk canker and give the sur-
face of the bark a pimpled appearance (Fig.
15.27/Plate 115). Thus, pycnidia are not asso-
ciated with bark lenticels. Discoloured wood,
wilting and chlorotic and dehiscent leaves are
other symptoms associated with Leucostoma
canker.
Conidia are the primary inoculum for the
disease. They are most abundantly produced
under cool, moist conditions of late autumn
and early spring, but are present throughout
the year if rainfall is sufficient. Conidia are
dispersed by rain, wind and possibly birds
and wood-boring beetles. Germination requires
the presence of free water or 100% relative
Fig. 15.26. Leucostoma or perennial
cankers caused by Leucostoma cincta
(Cytospora cincta) around nodes of
peach shoot. (Reprinted from Ogawa
et al., 1995, with permission of APS.)
378 J.E. Adaskaveg et al.
Fig. 15.27. Pycnidia of Leucostoma cincta (Cytospora cincta) on cherry twig. Spore masses are exuded
by some pycnidia on the upper part of the twig.
humidity and a carbon source. Most infections
occur on wood injured by sunburn, pruning,
insects or rodents. Trees stressed by freezing,
nutrient deficiency and infections by ring
nematodes and bacterial canker are predis-
posed to the disease. Young vigorous trees are
less susceptible. On older weakened trees
many new infections may occur at the nodes
of 1-year-old shoots.
Because Leucostoma canker develops on
weakened trees, requires injuries for infec-
tions and usually follows winter injury in
cold climates and sunburn in warm climates,
the management of Leucostoma canker requires
an integrated approach of cultural and pest
management practices. In both cold and
warm climates, the disease can be kept to a
minimum with good cultural practices that
ensure tree vigour and hardiness. Painting
trunks white with 100% acrylic latex paint
prevents sunburn (‘south-west’) injuries and
helps reduce Leucostoma canker. Trees should
not be planted on heavy clay or shallow soils,
where moisture and nutrient stress may occur.
Fungicides have not been effective. In sum-
mary, management of Leucostoma canker is
based on preventive measures that minimize
improper pruning cuts (Biggs, 1989), winter
injury, sunburn and insect damage, promote
optimum tree health, and facilitate rapid
wound healing.
Fungal gummosis
Fungal gummosis of peach trees was first
described in the 1960s almost concurrently
in the south-eastern USA and in Japan, but
has since been reported in Australia and
China. The disease is caused by Botryosphaeria
dothidea (Moug.:Fr.) Ces. & De Not (anamorph
Fusicoccum aesculi Corda). This fungal species
has one of the largest host ranges of plant
pathogenic fungi, but one physiological race
of the pathogen is specific to peach. Other spe-
cies of Botryosphaeria, Botryosphaeria rhodina
(Cooke) Arx and Botryosphaeria obtusa (Schein.)
Shoemaker, have also been reported to cause
gummosis, but these organisms are primarily
wound invaders that are not known to cause
lenticel infections of the bark like B. dothidea
(Pusey et al., 1995).
Lenticel infections by B. dothidea occur
primarily in the summer months, whereas
wound infections by all three species may
occur at other times during the growing sea-
son with wet weather and when inoculum is
available. Characteristic symptoms of the dis-
ease are first evident as blisters around lenti-
cels on young bark (Fig. 15.28/Plate 116) and
develop in the second or third growing sea-
son of the tree. Subsequently, blisters enlarge
and necrotic lesions 5–15 mm in diameter
develop around the lenticels, which later are
 Diseases Caused by Fungi
sunken and exhibit excessive gum exudation
(Fig. 15.29/Plate 117). Lesions may coalesce
and form extensive cankers (Fig. 15.30/Plate
118). On young branches blisters 1–6 mm in
diameter develop around lenticels, but there
is no gumming. Symptoms generally first
occur on the trunk and then on scaffold
branches or even on smaller fruiting branches.
Trees in general good health will form a peri-
derm around infection sites and diseased tis-
sue is walled off from healthy tissue. Fungal
gummosis can significantly suppress tree
growth and fruit yield on susceptible peach
cultivars (Beckman et al., 2003). Branches and
entire trees in poorly managed orchards,
however, may die.
B. dothidea overwinters in diseased bark
and woody tissues. The pathogen produces
379
Fig. 15.28. Fungal gummosis caused
by Botryosphaeria dothidea. Blisters
develop around lenticels on young
bark, which develop in the second or
third growing season of the tree.
asexual conidia in pycnidia and sexual asco-
spores in ascostroma perithecia in branch
and trunk cankers. Under wet conditions
abundant conidia may be produced, which
are disseminated by splashing irrigation
water or rain.
For management of the disease, dead
trees and infected wood should be removed
or destroyed (e.g. by burning, shredding or
flail-mowing) to reduce inoculum sources.
Winter pruning is encouraged to avoid peri-
ods when inoculum is present. During the
summer, inoculum is commonly present and
pruning at this time may result in rapid colo-
nization of wounds by any of the three spe-
cies of Botryosphaeria. Fungicide applications
may prevent infections, but may not be cost-
effective in many areas.
380 J.E. Adaskaveg et al.
Constriction canker
Peach constriction canker, also known as Fusi-
coccum canker, is a disease caused by the ana-
morphic fungus Phomopsis amygdali (Del.)
Tuset & Portilla (previously known as Fusic-
occum amygdali Del). The disease has been
reported from the mid-Atlantic region of the
USA, European countries (the UK, France,
Italy, Portugal, Bulgaria), North Africa (Tuni-
sia), South America (Argentina, Brazil) and
Japan. Historically, constriction canker was
devastating to peach production in the mid-
Atlantic states in the mid-1900s. Some peach
cultivars such as ‘Golden Jubilee’ were so
severely affected that the disease prevented
its cultivation in New Jersey. Oddly, in the
Fig. 15.29. Fungal gummosis caused
by Botryosphaeria dothidea. Gumming
blisters and necrotic tissue under
lenticels, which develop under the
bark.
1970s, the disease virtually disappeared in the
USA but it caused extensive tree decline and
death in Italy. Recently, constriction canker
has reappeared in southern New York and New
Jersey. The pathogen has also been reported
on almond in Europe and in California.
Symptoms develop primarily on twigs
and leaves. Typically, stem infections develop
on 1-year-old shoots as reddish-brown or pale
brown, sunken, elliptical lesions around buds
or nodes (Fig. 15.31/Plate 119). This is in
contrast to cankers caused by brown rot
blossom blight, which develop from infected
blossoms. The sunken lesions of constriction
canker enlarge (2–8 cm) to form cankers with
alternating light and dark brown rings or
zones. Subsequently, the twigs are girdled (i.e.
 Diseases Caused by Fungi
constriction canker) and develop a blighted
appearance. Lesions of constriction canker
also exude some gum and sometimes can be
confused with brown rot infections, but the
latter disease causes more profuse gumming
from infections. The fungus produces a toxin
(fusicoccin) that is distally translocated from
twig cankers and contributes to leaf wilting
and yellowing on blighted twigs (Fig. 15.32/
Plate 120). On leaves, the disease develops as
brown, zonate, circular to irregular spots that
have black asexual pycnidia in the centre. The
pathogen is restricted to leaf lesions in the
summer but may continue to develop into the
vascular tissue in the autumn. Brown, firm
fruit decay has also been reported but both
leaf and fruit infections are considered of
minor importance.
381
Fig. 15.30. Fungal gummosis caused
by Botryosphaeria dothidea. Lesions may
coalesce and form extensive cankers
with excessive gumming on lower
peach tree trunks.
Infections may develop throughout the
growing season with unusually wet weather.
Lesions commonly develop in spring and
autumn, but under favourable environmental
conditions (i.e. wet weather) lesion numbers
may continue to increase into summer. High N
levels within trees also favour the disease. The
pathogen produces numerous conidia in long
tendrils or cirri from pycnidia (Fig. 15.33/Plate
121) during wet weather. The conidia are rain-
splash dispersed and germinate over a wide
range of temperatures (5–36°C). Although the
optimal temperature range for fungal growth
is 29–30°C, successful host infection occurs
mainly at 5–15°C. In autumn, infections occur
through leaf scars and during the growing sea-
son through buds, scars on bud scales, stipules
and fruits, or directly through young shoots.
382 J.E. Adaskaveg et al.

Fig. 15.31. Constriction canker caused by Phomopsis amygdali. One-year-old peach twigs with
infections around buds. The lower twig shows the zonations around the infection site.

Fig. 15.32. Constriction canker caused by Phomopsis amygdali. Flagging and withering of blighted
peach twigs distal to twig cankers.

Peach cultivars differ widely in their sus-
ceptibility to constriction canker. Thus, plant-
ing of less-susceptible cultivars is an important
consideration in establishing an orchard in
areas where the disease is a problem. Fertil-
ization programmes that carefully regulate N
to low or moderate levels decrease host sus-
ceptibility. Judicious pruning of infected
branches and careful removal of prunings
from the orchard help reduce inoculum and
prevent the spread of infections, but this may
not be economically feasible and results may
be inconsistent (Lalancette and Robison, 2002).
The use of fungicides (e.g. benzimidazoles,
captan, captafol, chlorothalonil) applied before
bud break and in the autumn is effective for
disease management (Lalancette and Robison,
2002). The extensive use of benzimidazoles
 Diseases Caused by Fungi
Fig. 15.33. Cirri (tendrils) exuding from pycnidia of Phomopsis amygdali. Each cirrus is composed of
abundant conidia of the pathogen.
and captan for management of brown rot and
scab during the growing season has probably
contributed to the reduced occurrence of con-
striction canker in the past. The increased use
of newer fungicides that have a lower persis-
tence in the orchard environment may be
responsible for the recent increase of the dis-
ease. In addition, fungicides are not com-
monly applied in late summer and autumn,
when a significant amount of infections may
occur during wet weather.
15.4 Root and Crown Diseases
Phytophthora root and crown rots
Phytophthora root and crown rots are destruc-
tive soil-borne diseases of stone fruits that
occur worldwide in nurseries and orchards.
Trees can be affected at all ages and infections
often result in tree death. The identification of
diseases caused by Phytophthora spp. has
improved over past decades due to better
detection methods including the development
of selective media. The incidence of disease
has increased because orchards are estab-
lished in less than optimal locations that have
poor soil drainage or use improper irrigation
practices. Phytophthora root and crown rots
383
can be caused by a number of species in the
genus Phytophthora (Wilcox and Ellis, 1989),
fungal-like organisms that are not closely
related to the ‘true’ fungi and are classified in
a separate kingdom (Chromista or Strameno-
pila). Species include Phytophthora cactorum
(Lebert & Cohn) J. Schröt., Phytophthora cambiv-
ora (Petri) Buisman, Phytophthora cinnamomi
Rands, Phytophthora citricola Sawada, Phy-
tophthora citrophthora (R.E. Sm. & E.H. Sm.)
Leonian, Phytophthora cryptogea Pethybr. &
Lafferty, Phytophthora drechsleri Tucker, Phy-
tophthora megasperma Drechs., Phytophthora
parasitica Dastur and Phytophthora syringae
(Kleb.) Kleb. In a particular location, several
species of Phytophthora may be present ende-
mically; other species may be introduced by
infested irrigation water, soil or plant mater-
ial. When free water is present, all species
commonly produce sporangia that release
numerous asexual zoospores and they may
also form resistant chlamydospores and sexual
oospores. Ecologically, the different species
differ in their geographic distribution, their
characteristic temperature optimum, their
capability to cause root and/or crown rots, their
seasonal occurrence and in their virulence.
Trees infected by Phytophthora spp. show
poor terminal growth and leaves are small,
chlorotic and sparse (Fig. 15.34/Plate 122).
Fruit may be undersized, highly coloured and
384 J.E. Adaskaveg et al.

Fig. 15.34. Dieback of peach tree in foreground affected by Phytophthora root and crown rot.
(Reprinted, with permission, from Strand, 1999.)

sunburned. Trees either show dieback and
decline progressively over several seasons or
die suddenly in late spring or summer fol-
lowing years with excessive wet weather.
These symptoms are often not very character-
istic and may be confused with other diseases
and disorders. Mild symptoms of root and
crown rot are not always noticeable and yet
yield is reduced. Crown rots develop at the
root crown and/or at the base of the trunk
(Fig. 15.35/Plate 123). The bark is killed and
discoloration may extend into the outer lay-
ers of the xylem tissues. As the decay expands,
a canker develops that is often accompanied
by the production of copious amounts of amber
to brown gum. Cankers have distinct borders
between diseased and healthy tissues. If the
canker encircles the trunk, the tree is girdled
and dies. If the lesion ceases expansion or the
pathogen dies, callus tissue will grow into the
dead areas of the bark in an attempt to heal
the wound. Cankers are usually limited to the
crown tissues, but sometimes may extend up
the trunk and occasionally into scaffold bran-
ches. Aerial cankers develop when inoculum
is disseminated by rain or sprinkler water to
upper parts of the tree, especially the scaffold
crotches, where water tends to accumulate
 Diseases Caused by Fungi
(Fig. 15.36/Plate 124). Infected roots exhibit
decay, sometimes extending to the crown. On
feeder roots, portions of the outer cortical tis-
sue are disintegrated, leaving only the white,
hair-like stele protruding from the decaying
outer tissue.
Phytophthora spp. are mainly soil-borne
organisms (Fig. 15.37). They survive as chla-
mydospores, hyphae or oospores in root
debris in the soil. At high soil moisture levels,
the fungi produce abundant sporangia that
release numerous zoospores. Zoospores are
motile and are attracted by root exudates to
host plant roots, where they encyst and infect
mainly the fine feeder roots. After each rain or
irrigation new generations of sporangia are
produced, which will restart the disease cycle.
Zoospores are considered the main infective
propagules, but other structures (i.e. chlamy-
dospores, oospores) may also cause infections.
385
Fig. 15.35. Phytophthora crown rot
on peach tree. (Reprinted, with
permission, from Strand, 1999.)
Although Phytophthora spp. are pathogenic
on their own, in many cases Phytophthora root
rots are increased with root injuries caused by
nematodes or other organisms.
Phytophthora root and crown rots are
managed by the use of resistant rootstocks,
careful soil water management and by fungi-
cides. A considerable degree of resistance
among stone fruit rootstocks is available, but
none is immune. Because the zoospores are
dependent on water for movement, the dis-
ease is greatly affected by soil water manage-
ment. Ideally, orchards should be established
on well-drained soils that are never water-
saturated for more than 24 h. In soils with
poor drainage, trees should be planted on
ridges or raised beds. Flood or furrow irriga-
tion increases the disease, whereas irrigating
with micro-sprinklers and allowing the sur-
face soil to dry between irrigations will reduce
386 J.E. Adaskaveg et al.
root rot. In addition, sprinklers should never
be directed towards tree trunks and crotches.
Control of root and crown rots may also be
achieved with the use of systemic fungicides
that are applied to the soil, injected, or sprayed
or painted on gummosis lesions. In problem-
atic areas, fungicides together with nemati-
cides or a soil fumigant should be used to
protect replants during the first 2 years of
growth.
Armillaria root rot
Armillaria root rot (syn. shoestring root rot,
oak root rot) is an important disease of
peaches and other stone fruits worldwide,
but losses have been greatest in production
Fig. 15.36. Peach tree with an aerial
infection of a Phytophthora spp.
(Reprinted, with permission, from
Strand, 1999.)
areas in North America. Many native, orna-
mental and agricultural woody plants are
affected by Armillaria root rot. Trees can be
killed at all ages. Severe outbreaks are often
observed on land planted successively with
peaches where inoculum was allowed to
build up gradually (Savage et al., 1953) or
where forest trees were recently removed. In
contrast to Phytophthora root rot, Armillaria
root rot is also common on well-drained soils.
Early symptoms of Armillaria root rot
include poor terminal growth and under-
sized, curled leaves on all major limbs (Fig.
15.38/Plate 125). Infected peach trees may
collapse suddenly during summer months with
a majority of leaves still attached (Fig. 15.39/
Plate 126). In older orchards with widely
spaced trees, the disease may expand in a cir-
cular fashion, but in high-density orchards
 Diseases Caused by Fungi 387

Symptoms of Phytophthora root

and crown rot include gumming
and necrosis under the bark of the
lower trunk and necrotic roots
Feeder roots are
infected by zoospores
and possibly
chlamydospores

Gumming
and necrosis

Symptoms of
aerial
Phytophthora
cankers include Gumming Dying tree
gumming and
necrosis under the
bark

Sporangia form from mycelium or

Oospores and chlamydospores are
germinating chlamydospores or produced by some species of
oospores and release zoospores Phytophthora in infected roots

Fig. 15.37. Disease cycle of Phytophthora root and crown rot of peach. Phytophthora spp. are
mainly soil-borne organisms that survive as chlamydospores, mycelium (hyphae) or oospores in root
debris in the soil. At high soil moisture levels, the fungi produce abundant sporangia with numer-
ous motile zoospores that are attracted by exudates of host plant roots. Zoospores encyst on root
surfaces and infect fine feeder roots. Tree crowns that are subjected to prolonged wetness from
flood irrigation or heavy rain often develop crown rot. After each rain or irrigation new generations
of sporangia are produced, which will restart the disease cycle. Zoospores are considered the main
infective propagules, but other structures (i.e. chlamydospores, oospores) may also cause infec-
tions. Propagules of Phytophthora spp. are carried to aerial portions of trees by dust, soil particles and
splashing water, where they can cause aerial Phytophthora cankers. Trees infected by Phytophthora
spp. show decreased growth and may ultimately die. (Drawing by J.E. Adaskaveg and H. Förster.)
388 J.E. Adaskaveg et al.

Fig. 15.38. Leaf symptoms of peach tree affected by Armillaria root rot.

Fig. 15.39. Armillaria-infected peach trees showing dieback.

 Diseases Caused by Fungi
the disease often progresses along tree rows.
White mycelial fans that develop in the cam-
bium between the bark and wood on the
crown are highly diagnostic signs of the dis-
ease (Fig. 15.40/Plate 127). In advanced stages
of colonization, white rot wood decay occurs
and the wood is soft, spongy and bleached
whitish in colour (Morrison et al., 1991). The
decay is a result of the degradation of cellu-
lose, hemicellulose and lignin. Roots may
have black, shoestring-like mycelial strands
called rhizomorphs attached to the surface
(Fig. 15.41/Plate 128). Armillaria tabescens,
however, does not form such rhizomorphs
under field conditions (although they have
been induced in the laboratory). Typically in
late summer to early autumn clusters of brown
mushrooms develop at the base of infected
trees (Fig. 15.42/Plate 129).
Until the late 1970s, Armillaria mellea
(Vahl:Fr.) Kummer was considered the causal
organism of Armillaria root rot. Based on com-
patibility assays, it was subdivided into ten
biological species (Anderson and Ullrich,
1979) that have since been shown to be well-
defined morphologically and genetically dis-
Fig. 15.40. Mycelial fans of Armillaria sp. under the bark of infected peach tree.
389
tinct. Three species of Armillaria are currently
associated with peach. In North America, A.
mellea sensu stricto (Vahl:Fr.) P. Kumm., Armil-
laria ostoyae (Romagn.) Herink and A. tabescens
(Scop.) Dennis, Orton & Hora (syn. Clitocybe
tabescens) have been found in California,
Michigan and in the south-eastern USA,
respectively. A. tabescens was also identified
on peach in Japan (Fujii and Hatamoto, 1974).
A. mellea and A. ostoyae are the predominant
species in Europe. The species can be easily
differentiated when fruiting bodies are avail-
able. In their absence, labour-intensive com-
patibility tests (Korhonen, 1978; Anderson and
Ullrich, 1979) and, more recently, DNA diag-
nostics are being used for species identifica-
tion (Harrington and Wingfield, 1995; White
et al., 1998; Sierra et al., 1999). Previous DNA
methods were limited in their usefulness for
distinguishing A. tabescens from other species.
Based on ITS1 sequence data, a PCR-based
identification technique was recently devel-
oped to distinguish A. tabescens from A. mellea
(Schnabel et al., 2005).
The disease cycle for Armillaria root rot
is shown in Fig. 15.43. Armillaria spp. persist
390 J.E. Adaskaveg et al.

Fig. 15.41. Rhizomorph of Armillaria mellea (top) and healthy root (bottom). (Reprinted, with
permission, from Strand, 1999.)

Fig. 15.42. Basidiomes of Armillaria sp. at the base of infected peach tree.
 Diseases Caused by Fungi 391

Root-to-root contact and Rhizomorph

spread of disease in infection
centres

Healthy root

Rhizomorphs growing from

diseased to healthy tree (note:
some species do not form rhizomorphs)

White rot
of wood

Dying
Possibly, basidiospores
tree
infect wounded
roots or exposed
wood Basidium and Mycelial fan
basidiospores under bark and
rhizomorph
on root

Clusters of honey-coloured Immature and mature

basidiomes with white spores basidiomes (mushrooms)
and annulus on stipe (note: some at tree base
species do not possess an annulus)

Fig. 15.43. Disease cycle of Armillaria root rot of peach caused by Armillaria spp. The pathogen
persists in the soil as mycelium in infected roots or as rhizomorphs that are associated with tree
roots, depending on the species of Armillaria involved. New infections of healthy roots originate from
infected root segments in the soil, from root-to-root contact, or from contact with rhizomorphs.
Basidiospores that are produced on the fruiting bodies are not considered major infection
propagules in fruit orchards. Early symptoms of Armillaria root rot include poor tree growth. Infected
peach trees may collapse suddenly during summer months with a majority of leaves still attached.
White mycelial fans that develop in the cambium between the bark and wood on the crown are highly
diagnostic signs of the disease. In advanced stages of colonization, white rot wood decay occurs
and the wood is soft, spongy and bleached whitish in colour. Roots may have black, shoestring-like
mycelial strands called rhizomorphs attached to the surface. Typically in late summer to early autumn
clusters of brown mushrooms develop at the base of infected trees. (Drawing by J.E. Adaskaveg.)

in the soil for many years as mycelium in saprophytically. Thus, very young trees are
infected plant tissues or as rhizomorphs. The often killed on replant sites or older trees after
pathogens are most successful in invading they have grown for 5 years or more. Infec-
healthy tissue if they first become established tions of healthy roots originate from infected
392 J.E. Adaskaveg et al.

root segments in the soil, from root-to-root
contact, or from contact with rhizomorphs.
Spread by root-to-root contact or rhizomorphs
was estimated to be 0.8–3.2 m/year (Kable,
1974). Basidiospores that are produced on the
fruiting bodies are not considered major
infection propagules in fruit orchards (Rizzo
et al., 1998; Termorshuizen, 2000).
A strategy to prevent the spread of Armil-
laria root rot disease from an infection centre
is presented in Fig. 15.44. If tree rows are sep-
arated by a sod-middle, roots of trees are
mostly confined to the herbicide-treated strip
and it is sufficient to remove trees adjacent
(dotted circles) to the infected tree within a
row. If trees are grown on bare ground, roots
are likely to spread evenly in all directions
and all trees surrounding (dotted and waved
circles) the infected tree should be removed.
Management strategies for Armillaria
root rot are not very effective. If sites are seri-
ously infested they are usually abandoned.
The most effective strategy is to avoid plant-
ing new orchards on infested soil. Options to
manage the disease on replant sites are lim-
ited and often only marginally effective.
Chemical control options with moderate effi-
cacy include pre- and post-planting fumiga-
tions to reduce inoculum in the soil (Munnecke
et al., 1970; Savage et al., 1974; Adaskaveg
et al., 1999); however, many fumigants face
increased regulation and, in the case of methyl
bromide for example, are scheduled for immi-
nent phasing out. Cultural management prac-
tices include raking out old roots before
replanting, planting trees on raised beds and
removal of trees surrounding an infection
centre. Susceptibility differences in fruit tree
rootstocks to Armillaria infection have been
reported (Beckman et al., 1998), but none of the
currently available rootstocks is immune.
Verticillium wilt
Verticillium wilt of peach and other stone fruit
crops caused by Verticillium dahliae Kleb.
occurs in many parts of the world and can
cause serious economic losses, although the
disease has been recently more of sporadic
occurrence. Many plants including tree and
annual vegetable crops have been reported as
hosts of this fungal pathogen. Peach and nec-
tarine trees sometimes develop Verticillium
wilt when planted in locations where highly
susceptible crops such as cotton, tomato or
peppers were grown for a number of years.
The disease is most serious on young trees
although some peach cultivars may recover
with age. Mature trees are most susceptible in
cooler climates.
The soil-borne V. dahliae produces resis-
tant structures (microsclerotia) that can sur-
vive in the soil for many years in the absence
of hosts when soil temperatures are between
5 and 15°C and soil moisture-holding capacity is
between 50 and 75%. Conidia are short-lived
and are considered to have a minor role in
infection and dissemination of the fungus

Tree
row

Infected
tree

Fig. 15.44. Strategy to prevent the spread of Armillaria root rot disease from an infection centre. If
tree rows are separated by a sod-middle, roots of trees are mostly confined to the herbicide-treated
strip and it is sufficient to remove trees adjacent (dotted circles) to the infected tree within a row.
If trees are grown on bare ground, roots are likely to spread evenly in all directions and all trees
surrounding (dotted and waved circles) the infected tree should be removed.
 Diseases Caused by Fungi
from tree to tree. The fungus is found in the
highest concentration at a soil depth of 15–30 cm
but can be found as deep as 105 cm. The
pathogen infects the roots of host plants in the
spring and invades the vascular system, moving
up through the xylem as mycelium and asex-
ual conidia, interfering with water transport in
the xylem. The first symptom of Verticillium
wilt is a sudden wilting of leaves on one or
more branches during hot weather in the
summer (Fig. 15.45/Plate 130). Subsequently,
leaves yellow, brown and curl up. Sometimes
older, lower leaves wither and fall before
younger, upper leaves on the terminal ends of
branches. Thus, young trees can be killed, but
the primary symptoms on several-year-old
trees are poor growth and low productivity.
Longitudinally cut branches of diseased trees
show brown to black streaks in the sapwood.
In cross-sections, a portion of the outer xylem
will be stained dark brown to black (Fig. 15.46/
Plate 131). In hot weather, the pathogen dies
in the upper part of infected trees and the tree
can recover with new growth the following
season. The disease cycle, however, can recur
each growing season.
The most effective management practice
for Verticillium wilt is to avoid planting orchards
in soils where inoculum of the fungal pathogen
Fig. 15.45. Verticillium-infected peach tree with wilting of branches. (Reprinted, with permission,
from Strand, 1999.)
393
has increased on other susceptible crops that
have been grown for a number of seasons.
Inoculum of the pathogen can be reduced
using chemical fumigation, solarization,
flooding fallow fields, growing grass crops
for several seasons, or any combination of
these treatments. Rootstocks resistant to V.
dahliae are unknown. Minimizing tree stress
through maintenance of soil fertility and soil
moisture will help trees tolerate the disease
and encourage their recovery.
Peach tree short life
Peach tree short life (PTSL) is a disease com-
plex characterized by sudden wilt and col-
lapse of new growth and death of all aerial
portions of the tree (see also Chapters 16 and
19). The disease is part of general replant dis-
orders, which refer to problems that diminish
tree growth and productivity. PTSL also
affects other stone fruits and has been a seri-
ous threat to commercial peach growers in
the south-eastern USA for more than 100
years (Chandler, 1969). The number of peach
trees lost to PTSL in South Carolina roughly
tripled when the soil fumigant DBCP (1,2-
dibromo-3-chloropropane) was no longer
394 J.E. Adaskaveg et al.
registered for use. The disease is similar to the
bacterial canker complex in California with
the exception of cold injury, which kills many
trees with PTSL. Bacterial and nematode
aspects are similar for the two diseases.
The symptoms defining the PTSL syn-
drome were established in the early 1970s.
The canopy of a peach tree suddenly collapses
before, during or just after bloom, usually 3–6
years after planting (Fig. 15.47/Plate 132). Tree
sap often oozes out of scaffold limbs and trunk.
On many limbs, the tissue just beneath the
bark is discoloured (Fig. 15.48/Plate 133) and
has a sour odour. Peripheral parts of the tree,
although dying, may still reveal healthy tissue
underneath the bark. Trees are killed only to
the soil line. Later in the season, shoots from
the living rootstock often emerge (Ritchie and
Clayton, 1981) (Fig. 15.49/Plate 134). This is in
contrast to trees infected with Armillaria root
Fig. 15.46. Cross-section through
branch of Verticillium-infected peach
tree with discoloured outer xylem.
(Reprinted, with permission, from
Strand, 1999.)
rot, where root suckers do not emerge from the
rootstock because it may be completely dead.
PTSL is a complex disease with biotic
and abiotic factors that may contribute to the
disease directly or indirectly. The immediate
causes of death can be cold injury (Nesmith
and Dowler, 1976), bacterial canker caused by
Pseudomonas syringae Van Hall, a combination
of the two, and perhaps Cytospora canker.
Cold injury in the south-eastern USA occurs
in late winter after the rest period has been
broken and physiological activity resumes. In
a normal season, sufficient chill-hours have
accumulated by late January and peach trees
can lose hardiness after periods of warm
weather in the dormant season. Like frost injury,
bacterial canker can affect all parts of a peach
tree if the tree is stressed. Indirect factors that
predispose trees to PTSL include the com-
monly found ring nematode (Mesocriconema
 Diseases Caused by Fungi
Fig. 15.47. Orchard with trees affected by peach tree short life.
xenoplax (Raski) Loof and de Grisse) (Ritchie
and Clayton, 1981), improper rootstock selec-
tion (Zehr et al., 1976) and time of pruning
(Nesmith and Dowler, 1976), root injury, phys-
ical characteristics of the orchard site such as
pH and soil structure, and planting on land
previously used for peach production.
The disease is most common in sandy
soils. Losses are more frequent and severe in
orchards where peaches or other stone fruits
have been grown previously than in locations
where peaches have not been grown before.
In clay soils less difficulty is encountered when
peach orchard sites are replanted. Ideally, peach
plantings should be established on new ‘vir-
gin’ land with no recent history of growing
stone fruit. Site selection in any given region,
however, is often compromised by other factors
(e.g. urban encroachment, unsuitable micro-
climate, proximity to packinghouses).
The complexity of the PTSL syndrome
complicates attempts to effectively manage
the disease in commercial orchards. A 10-Point
Programme that emphasizes practices to
enhance the health and vigour of peach trees
was developed (Ritchie and Clayton, 1981).
The programme consists of the following rec-
395
ommendations: (i) adjust pH to at least 6.5
before planting; (ii) cultivate subsoil before
planting; (iii) fumigate on replant sites with
sandy soil and other nematode-infested soils
(see Chapter 19); (iv) continue to fumigate such
soils at approximately 2-year intervals or as
indicated by nematode populations; (v) select
rootstocks tolerant or resistant to nematodes;
(vi) purchase rootstocks that are certified to
be free of nematodes or have been grown on
fumigated soil; (vii) apply nutrients and lime
as needed; (viii) delay pruning to late winter
(February and March); (ix) use recommended
herbicides for weed control; and (x) remove
and burn all dead or dying trees. Alternatives
to methyl bromide, such as chloropicrin and
methyl iodide, are currently being explored
as soil fumigants. Pre-plant fumigation is
expensive and does not provide long-term con-
trol of nematodes. Currently, one of the most
effective management strategies is the use of
rootstocks such as the highly PTSL-tolerant
‘Guardian’ (BY520-9 line) (Okie et al., 1994),
which is rapidly replacing other rootstocks in
the south-eastern USA. This rootstock is also
very resistant to the ring nematode. Unfortu-
nately, the BY520-9 lines are genetically highly
396 J.E. Adaskaveg et al.
Fig. 15.48. Discoloured tissue under the bark
of peach tree short life-affected tree.
diverse and research is ongoing to further
select lines with horticultural characteristics
similar or superior to commercial rootstocks
(Wilkins et al., 2002).
15.5 Postharvest Diseases
Brown rot and grey mould
Worldwide, brown rot caused by M. fructicola,
M. laxa and M. fructigena is the most impor-
tant postharvest decay of peach and other
stone fruits and causes losses every year.
Losses are especially high in areas where the
more aggressive M. fructicola is present and,
without management, large amounts of the
crop may be destroyed. Grey mould, caused
by B. cinerea, also occurs every year, but dam-
age is generally less serious. Preharvest
aspects of the brown rot and grey mould
pathogens are discussed above. Fungal inoc-
ulum starts building up in the orchard when
blossom blight and jacket rot or green fruit rot
are not managed. At harvest time, when air-
borne and surface-deposited conidia gain
entry through small wounds on the fruit,
postharvest decay can result. Conidia germi-
nate and start growing into the fruit within
4 to 6 h at 20–25°C. Some of the postharvest
brown rot and grey mould decay originates
from quiescent infections that occur during
early fruit development and become actively
growing when the fruit is maturing. Both
decays are firm, brownish and develop rap-
idly (Figs 15.50 and 15.51/Plates 135 and 136).
Fruit may be completely rotted after 3–4 days
at 20°C. Both decays are not easily differenti-
ated at the earlier stages, but decay caused by
B. cinerea is of a slightly lighter brown to tan
colour than the one caused by Monilinia spp.
At advanced decay stages the fruit surface is
covered with cottony fungal mycelium and
conidia. Fungal structures of Monilinia spp.
are of a light brown colour, whereas those of
B. cinerea are grey. Brown rot decay on fruit in
cold storage often appears black with little or
no sporulation. The black-coloured areas
(pseudosclerotia) consist of both fungal and
host tissues.
Rhizopus rot, Gilbertella and Mucor decays
Three fungal species in the Zygomycetes (order
Mucorales) that also can cause significant post-
harvest losses on peach and nectarine are Rhizo-
pus stolonifer (Ehrenb.:Fr.) Vuill., Gilbertella
persicaria (E.D. Eddy) Hesselt., Mucor piriformis
E. Fisch and other species of Mucor. The three
fungi are rarely a problem preharvest, except in
warm, wet climates, where infections may
already occur through injuries or cracks on
ripening fruit on the tree. The spores of these
 Diseases Caused by Fungi 397

Fig. 15.49. Rootstock sprouting from peach tree killed by peach tree short life.

Fig. 15.50. Postharvest brown rot of peach fruit caused by Monilinia fructicola.
398 J.E. Adaskaveg et al.
fungi are ubiquitous in soils and are easily
wind-disseminated. Infections occur on mature
fruit that are injured or bruised, especially
during harvest and postharvest handling.
Rhizopus-infected fruit are first covered with a
thick cottony mycelial layer that rapidly starts
sporulating, forming tiny, black, terminal
sporangia that contain large numbers of spores
(Fig. 15.52/Plate 137). Fruit infected by R.
stolonifer decay very rapidly and an entire
fruit may turn into a soft, watery rot within
1–2 days at optimum temperatures. Likewise,
a cottony mycelial mat, which is shorter than
that of R. stolonifer, first covers Gilbertella-
infected fruit. Small, black, glistening sporan-
gia are produced terminally on the mycelium
(Fig. 15.53/Plate 138). Mucor-infected fruit first
look similar to Rhizopus rot, but sporangio-
phores are often longer and sporangia are tan
to brown in colour. Fruit infections by these
pathogens may spread quickly by hyphal con-
tact to healthy fruit (nesting), destroying large
numbers of fruit in a basket or box. R. stolonifer
and G. persicaria do not grow at temperatures
below 4°C. The optimum temperature range
for growth of R. stolonifer is 21–27°C with a
maximum of 33°C, whereas for G. persicaria
the optimum range is 30–33°C with a maxi-
mum of 39°C. In contrast, the optimum range
Fig. 15.51. Postharvest grey mould
of nectarine fruit caused by Botrytis
cinerea.
for growth of M. piriformis is between 10 and
15°C and the fungus can grow at temperatures
near freezing, but not at or above 27°C.
Sour rot
Sour rot of peach, caused by Geotrichum
candidum Link (teleomorph Galactomyces
geotrichum (E.E. Butler and L.J. Petersen) Red-
head and Malloch), has only been infrequently
reported to cause problems of traditionally
handled and marketed fruit (e.g. fruit picked,
hydro-cooled, transported at low temperatures
and displayed at markets). Fruit that are tree-
ripened or pre-conditioned (fruit ripened after
harvest for 1–2 days or until a pre-selected
firmness is reached and then refrigerated) are
more prone to the disease. Sour rot-like infec-
tions may also be caused by other yeasts and
possibly other organisms that are not well
characterized. Sour rot occurs mainly on ripe
fruit but may also occur on severely injured
immature fruit. Symptoms include a watery,
soft decay with a thin layer of white mycelial
growth on the fruit surface (Fig. 15.54/Plate
139). The decay may reach the pit and
consume the entire fruit. Rotted fruit have a
 Diseases Caused by Fungi 399

Fig. 15.52. Postharvest Rhizopus rot of peach fruit caused by Rhizopus stolonifer with decay spreading
by contact to healthy fruit (nesting).

Fig. 15.53. Postharvest Gilbertella rot of peach fruit caused by Gilbertella persicaria.

characteristic yeasty to vinegary odour; how- pathogen that decays fruit after spores are
ever, other odours may develop with bacte- deposited into injuries. The organism is wide-
rial contamination that commonly occurs in spread on organic material in the soil and is
the watery decay. G. candidum is a wound commonly found in dust or dirt on fruit
400 J.E. Adaskaveg et al.
Fig. 15.54. Sour rot caused by Geotrichum candidum initiated in wounds of the skin.
surfaces. During harvest micro-wounds occur
on the fruit and these injuries may function as
infection sites. The minimum temperature for
spore germination, growth and infection of
the fungus is about 2°C, the optimum is
25–27°C, and the maximum is 38°C.
Management of postharvest decays
Management of postharvest decays requires an
integrated approach with pre- and postharvest
components that focus on maintaining a
healthy crop, delaying fruit senescence,
avoiding injuries to fruit, sanitation practices
and fungicide use (Adaskaveg et al., 2002).
Preharvest cultural practices include cultivar
selection, fertilization, pruning, weed control,
irrigation and sanitation (e.g. removal of dis-
eased plant material such as fruit mummies).
For brown rot, nectarines are more suscepti-
ble than peaches, and fresh market (i.e. free-
stone) peaches are typically more susceptible
than canning (i.e. clingstone) peaches. Cultivar
differences also exist, although no cultivar is
completely resistant. Excess N fertilization is
known to increase susceptibility to brown rot
and other diseases. Orchard planting designs,
pruning practices and orchard floor (includ-
ing ground covers) management can signifi-
cantly alter the microclimate. Dense tree
canopies can restrict air movement, increase
relative humidity, prolong leaf wetness dura-
tion and decrease pesticide efficacy. Irrigation
systems can also affect the incidence of dis-
ease. High-angle sprinklers should be avoided
to prevent canopies and tree trunks from get-
ting wet. Preharvest fungicide treatments are
successfully used for management of brown
rot and grey mould. Treatments that are
applied 14 to 0 days before harvest help to
protect wounds from infections that occur at
harvest time. Fungicides with locally systemic
action also can inactivate some of the brown rot
and grey mould quiescent infections that only
penetrate the fruit for several cell layers. After
harvest it is important to remove all fruit from
the trees and from the orchard floor. Additional
sanitation practices include the removal of tree
prunings and cleaning of harvest equipment,
including bins, pick bags and trailers.
Postharvest handling and marketing
practices that minimize fruit injuries, utilize
sanitation of fruit and equipment, and employ
temperature management of harvested fruit
are essential in any management programme.
For minimizing injuries, shock-absorbing
foam pads are used in most harvesting, han-
dling and packing equipment. Pre-washing
and hydro-cooling treatments that contain
sodium hypochlorite or other oxidizing mate-
rials (e.g. ozone, chlorine dioxide) are used
for fruit sanitation. Harvest bins are usually
high-pressure washed or steam-treated. Other
 Diseases Caused by Fungi
sanitizing materials for equipment, but not
fruit, include quaternary ammonium com-
pounds. Temperature management is critical
to all postharvest handling systems for stone
fruit crops. Stone fruits are best stored at 0°C.
Low temperatures maximize the potential life
of the fruit and slow fungal development. At
0°C, some decay fungi such as R. stolonifer and
G. candidum will not grow, whereas others such
as Monilinia spp. and B. cinerea are greatly
reduced in their growth. Finally, postharvest
fungicide treatments are used to ensure the pre-
vention of decay from latent and active infec-
tions. These treatments also prevent the spread
of decay between adjacent fruit (i.e. nesting) in
packing containers. Without postharvest fungi-
cide use, fruit usually cannot be stored safely at
storage and display-shelf temperatures after
long-distance transportation and marketing
(Adaskaveg et al., 2002). New, safer postharvest
fungicides are currently being registered in the
USA that are highly effective against brown rot,
grey mould and Rhizopus rot.
15.6 Other Diseases of Peach and
Nectarine
A number of additional diseases affecting dif-
ferent parts of peach and nectarine trees have
been reported. Some of these diseases are lim-
ited in their geographic distribution, whereas
others are widespread but do not limit peach
and nectarine production. In favourable envi-
ronments, however, many of these diseases
can be economically important and limit crop
production. Examples of these diseases are
listed in Table 15.2.
Anthracnose caused by Colletotrichum acu-
tatum J.H. Simmonds and Colletotrichum gloeo-
sporioides (Penz.) Penz. & Sacc. in Penz. occurs
in areas and seasons with high rainfall and
warm temperatures during fruit ripening. Cir-
cular lesions that are brown, firm and slightly
sunken expand slowly over the fruit surface
(Fig. 15.55/Plate 140). Another diagnostic fea-
ture is that the lesion in cross-section is cone-
shaped, hardened and is easily separated from
the healthy mesocarp. Asexual fruiting struc-
tures of the pathogens (acervuli) are produced
in concentric rings on the fruit surface, bearing
401
masses of conidia that are disseminated by
splashing rain. Fungicide sprays can be used
to manage anthracnose where it is a problem.
Other diseases listed in Table 15.2 include
black knot caused by Apiosorina morbosa
(Schwein.:Fr.) Arx, Botryosphaeria fruit rot
caused by species of Botryosphaeria discussed
above, Diplodina fruit rot caused by Diplodina
persicae Horn & Hawthorne, frosty mildew
caused by Mycosphaerella pruni-persicae Deigh-
ton, Leucotelium white rust caused by Sorataea
pruni-persicae Tranzschel (formerly Leucote-
lium pruni-persicae (Hori) Tranzschel), Phyma-
totrichum root rot caused by Phymatotrichopsis
omnivore (Duggar) Hennebert, Rosellinia root
rot caused by Rosellinia necatrix Prill., Sclero-
tium stem rot caused by Sclerotium rolfsii Sacc.,
target leaf spot caused by Phyllosticta persicae
Sacc., violet root rot caused by Helicobasidium
mompa Tanaka, sour pit caused by Candida
inconspicua (Lodder & Kreger-Van Rij) Meyer
& Yarrow, and wood decay caused by a num-
ber of fungal species (over 56 species in North
America) in the phylum Basidiomycota
(Adaskaveg et al., 1993).
15.7 Concluding Remarks
Numerous fungal and fungal-like organisms are
pathogens of peach and nectarine. Many of
these organisms have a worldwide distribu-
tion, whereas others occur only locally to
regionally. Disease incidence is largely depen-
dent on climatic conditions and the distribu-
tion of the causal microorganism in peach
production areas, but also on the peach geno-
type. Many of the diseases are limiting to crop
production if they are not managed effec-
tively. Knowledge about peach diseases has
increased over the years. New pathogens
have been identified, described and studied
at the biological, epidemiological and molec-
ular levels. In most cases, they can be
effectively managed.
With worldwide trade and increased
travel, the potential for diseases to spread and
become limiting factors to peach production
has also increased. In the immediate future,
peach production will be dependent on the
success of programmes that prevent patho-
gen movement and introduction to new
Table 15.2. Other fungal diseases with variable importance in the production of peach and nectarine, their geographical occurrence, primary symptoms and
management practices.
Disease
Blossom, foliage and
fruit diseases
Anthracnose
Diplodina fruit rot
Frosty mildew
Leucotelium white rust
Target leaf spot
Trunk, scaffold and
branch diseases
Black knot
Sclerotium stem rot
Wood decay fungi
Root and crown diseases
Phymatotrichum root rot Phymatotrichopsis omnivora
Rosellinia root rot
Violet root rot
Postharvest diseases
Botryosphaeria fruit rot
Sour pit
aManagement practices with a question mark have a limited or poorly defined role in controlling the disease.
Pathogen
Colletotrichum acutatum and
Colletotrichum gloeosporioides
Diplodina persicae
Mycosphaerella pruni-persicae Widespread but infrequent
Leucotelium pruni-persicae
Phyllosticta persicae
Apiosorina morbosa
Sclerotium rolfsii
Species in the Basidiomycota
Rosellinia necatrix
Helicobasidium mompa
Botryosphaeria dothidea,
Botryosphaeria obtusa and
Botryosphaeria rhodinia
Torulopsis inconspicua
Geographical distribution Primary symptoms
Widespread but infrequent Sunken, firm fruit lesions
with concentric rings of
sporulation zones
South-eastern USA Leaf and fruit spots
White mildew on leaves
Japan, Korea, China Angular leaf lesions with rusty
brown uredinia or white telia
Italy, India, USA Leaf spot (target appearance)
Eastern USA Black, elongated swellings on
twigs
Nursery disease, widespread Stem cankers, wilting
in warm areas
Widespread in older orchards Wood rot – white and brown
knots
South-western USA and Mexico Dead roots are colonized by
golden mycelial strands with
cruciform hyphae
Widespread in warm areas, Root rot, tree decline
sporadic in the USA
Japan, Korea, China Infected roots are colonized by
purplish mycelium
South-eastern USA Fruit decay (sunken
soft lesions, no sporulation)
Widespread Fruit decay
402
Primary management practicesa
Fungicides (removal
of alternate host?)
Fungicides
None
Elimination of alternate host
None
J.E. Adaskaveg et al.
Elimination of alternate hosts and judicial
use of fungicides during active shoot growth
Sanitation, fungicides
Time of pruning, sanitation, removal of
infected branches
Integrated approaches – plant in disease
free areas, deep soil fumigation
Integrated approaches – soil fumigation,
fungicides
Integrated approaches – plant in disease
free areas, soil fumigation, fungicides
Preharvest disease management,
sanitation, fungicides
None available
 Diseases Caused by Fungi 403

Fig. 15.55. Anthracnose of peach

caused by Colletotrichum acutatum
showing circular rings where spores
of the fungus are produced.

geographical areas. Quarantines that com-
pletely restrict plant movement, as well as
controlled plant movement (e.g. the federal
Inter-Regional No. 2 or IR-2 programme in
the USA) and production certification pro-
grammes for breeders and nurseries poten-
tially will prevent or minimize harmful
microorganisms from being spread while
allowing exchange of diverse genetic plant
material. Furthermore, new control practices
based on biological and genetic information
of host and pathogen, as well as on novel
antimicrobial and host defence chemistries
are continuing to be developed. Ultimately,
with the promise of biotechnology, peach
may be genetically modified similar to some
other crops to fit the demands for new varie-
ties that are more tolerant of pests and dis-
eases. Thus, although plant pathological
aspects in peach production may become
more complex, the challenges will likely be
met by an array of traditional and new host
defence strategies.
Acknowledgements
Special thanks to Drs N. Lalancette and L.
Pusey for providing information and images
on constriction canker and fungal gummosis,
respectively. We also thank the University
of California, Agricultural and Natural
Resources (ANR) for providing images of
Sclerotinia sclerotiorum, shot hole infections of
peach twigs, Phytophthora root and crown
rots, Verticillium wilt and rhizomorphs of
Armillaria mellea and the American Phyto-
pathological Society (APS) for providing
images of Leucostoma canker.

References

Adaskaveg, J.E., Miller, R.W. and Gilbertson, R.L. (1993) Wood decay, lignicolous fungi, and decline of peach
trees in South Carolina. Plant Disease 77, 707–711.
Adaskaveg, J.E., Förster, H., Wade, L., Thompson, D.F. and Connell, J.H. (1999) Efficacy of sodium tetrathio-
carbonate and propiconazole in managing Armillaria root rot of almond on peach rootstock. Plant Disease
83, 240–246.
Adaskaveg, J.E., Förster, H. and Sommer, N.F. (2002) Principles of postharvest pathology and management of
decays of edible horticultural crops. In: Kader, A. (ed.) Postharvest Technology of Horticultural Crops.
University of California Agricultural and Natural Resources, Publication No. 3311. University of California,
Oakland, California, pp. 163–195.
404 J.E. Adaskaveg et al.

Anderson, J.B. and Ullrich, R.C. (1979) Biological species of Armillaria mellea in North America. Mycologia
71, 402–414.
Bassi, D., Rizzo, M. and Cantoni, L. (1998) Assaying brown rot [Monilinia laxa Aderh. et Ruhl. (Honey)] sus-
ceptibility in peach cultivars and progeny. In: Monet, R. (ed.) Proceedings of the Fourth International
Peach Symposium. Acta Horticulturae 465, 715–721.
Batra, L.R. (1991) World Species of Monilinia (Fungi): Their Ecology, Biosystematics and Control. Mycologia
Memoir No. 16. J. Cramer, Berlin.
Beckman, T.G., Okie, W.R., Nyczepir, A.P., Pusey, P.L. and Reilly, C.C. (1998) Relative susceptibility of peach
and plum germplasm to Armillaria root rot. HortScience 33, 1062–1065.
Beckman, T.G., Pusey, P.L. and Bertrand, P.F. (2003) Impact of fungal gummosis on peach trees. HortScience
38, 1141–1143.
Bensaude, M. and Keitt, G.W. (1928) Comparative studies of certain Cladosporium diseases of stone fruits.
Phytopathology 18, 313–329.
Biggs, A.R. (1989) Effect of pruning technique on Leucostoma infection and callus formation over wounds in
peach trees. Plant Disease 73, 771–773.
Biggs, A.R. and Northover, J. (1985) Inoculum sources for Monilinia fructicola in Ontario peach orchards.
Canadian Journal of Plant Pathology 7, 302–307.
Biggs, A.R. and Northover, J. (1988a) Influence of temperature and wetness duration on infection of peach and
sweet cherry fruits by Monilinia fructicola. Phytopathology 78, 1352–1356.
Biggs, A.R. and Northover, J. (1988b) Early and late-season susceptibility of peach fruits to Monilinia fructi-
cola. Plant Disease 72, 1070–1074.
Boehm, E.W.A., Ma, Z. and Michailides, T.J. (2001) Species-specific detection of Monilinia fructicola from
California stone fruits and flowers. Phytopathology 91, 428–439.
Byrde, R.J.W. and Willetts, H.J. (1977) The Brown Rot Fungi of Fruit: Their Biology and Control. Pergamon
Press, New York.
Chandler, W.A. (1969) Reduction in mortality of peach trees following preplant soil fumigation. Plant Disease
Reporter 53, 49–53.
Côté, M.-J., Tardiff, M.-C. and Meldrum, A.J. (2004) Identification of Monilinia fructigena, M. fructicola,
M. laxa, and Monilia polystroma on inoculated and naturally infected fruit using multiplex PCR. Plant
Disease 88, 1219–1225.
Daines, R.H. and Trout, J.R. (1977) Incidence of rusty spot of peach as influenced by proximity to apple trees.
Plant Disease Reporter 61, 835–836.
Dunegan, J.C. (1938) The rust of stone fruits. Phytopathology 28, 411–427.
Förster, H. and Adaskaveg, J.E. (2000) Early brown rot infections in sweet cherry fruit are detected by Monilinia-
specific DNA primers. Phytopathology 90, 171–178.
Fujii, S. and Hatamoto, M. (1974) Peach withering disease caused by Armillaria tabescens. Shokubutsu Boeki
28, 1–4.
Furman, L.A., Lalancette, N. and White, J.F. (2003a) Peach rusty spot epidemics: temporal analysis and rela-
tionship to fruit growth. Plant Disease 87, 366–374.
Furman, L.A., Lalancette, N. and White, J.F. (2003b) Peach rusty spot epidemics: management with
fungicide, effect on fruit growth, and the incidence–lesion density relationship. Plant Disease 87,
1477–1486.
Goldsworthy, M.C. and Smith, R.E. (1931) Studies on a rust of clingstone peaches in California. Phytopathology
21, 133–168.
Gottwald, T.R. (1983) Factors affecting spore liberation by Cladosporium carpophilum. Phytopathology 73,
1500–1505.
Gradziel, T.M. (1994) Changes in susceptibility to brown rot with ripening in three clingstone peach geno-
types. Journal of the American Society for Horticultural Science 119, 101–105.
Gradziel, T.M. and Wang, D. (1993) Evaluation of brown rot resistance and its relation to enzymatic browning
in clingstone peach germplasm. Journal of the American Society for Horticultural Science 118, 675–
679.
Harrington, T.C. and Wingfield, B.D. (1995) A PCR based identification method for species of Armillaria. My-
cologia 87, 280–288.
Hughes, K.J.D., Fulton, C.E., McReynolds, D. and Lane, C.R. (2000) Development of new PCR primers for
identification of Monilinia species. OEPP/EPPO Bulletin 30, 507–511.
Kable, P.F. (1974) Spread of Armillariella sp. in a peach orchard. Transactions of the British Mycological Society
62, 89–98.
 Diseases Caused by Fungi 405

Keitt, G.W. (1917) Peach Scab and Its Control. USDA Bulletin No. 395. US Department of Agriculture,
Washington, DC.
Korhonen, K. (1978) Interfertility and clonal size in the Armillaria mellea complex. Karstenia 18, 31–42.
Lalancette, N. and Robison, D.M. (2002) Effect of fungicides, application timing, and canker removal on
incidence and severity of constriction canker of peach. Plant Disease 86, 721–728.
Lawrence, E.G. and Zehr, E.I. (1982) Environmental effects on the development and dissemination of
Cladosporium carpophilum on peach. Phytopathology 72, 773–776.
Morrison, D.E., Williams, R.E. and Whitney, R.D. (1991) Infection, disease development, diagnosis, and detec-
tion. In: Shaw, C.G. and Kile, G. (eds) Armillaria Root Disease. USDA Forest Service Agricultural Handbook
No. 691. US Department of Agriculture Forest Service, Washington, DC, pp. 62–75.
Munnecke, D.E., Wilbur, W.D. and Kolbezen, M.J. (1970) Dosage response of Armillaria mellea to methyl
bromide. Phytopathology 60, 992–993.
Nesmith, W.C. and Dowler, W.M. (1976) Cultural practices affect cold hardiness and peach tree short life.
Journal of the American Society for Horticultural Science 101, 116–119.
Ogawa, J.M. and English, H. (1991) Diseases of Temperature Zone Tree Fruit and Nut Crops. University of
California Division of Agriculture and Natural Resources, Publication No. 3345. University of California,
Oakland, California.
Ogawa, J.M., Manji, B.T., Adaskaveg, J.E. and Michailides, T.J. (1988) Population dynamics of benzimidazole-
resistant Monilinia species on stone fruit trees in California. In: Delp, C.J. (ed.) Fungicide Resistance in
North America. American Phytopathology Society Press, St. Paul, Minnesota, pp. 36–39.
Ogawa, J.M., Zehr, E.I., Bird, G.W., Ritchie, D.F., Uriu, K. and Uyemoto, J.K. (1995) Compendium of Stone
Fruit Diseases. American Phytopathology Society Press, St. Paul, Minnesota.
Okie, W.R., Beckman, T.G., Nyczepir, A.P., Reighard, G.L., Newall, W.C. and Zehr, E.I. (1994) BY5209, a peach
rootstock for the southeastern United States that increases scion longevity. HortScience 29, 705–706.
Pusey, P.L., Kitajima, H. and Wu, Y. (1995) Fungal gummosis. In: Ogawa, J.M., Zehr, E.I., Bird, G.W., Ritchie,
D.F., Uriu, K. and Uyemoto, J.K. (eds) Compendium of Stone Fruit Diseases. American Phytopathology
Society Press, St. Paul, Minnesota, pp. 33–34.
Ries, S.M. and Royse, D.J. (1978) Peach rusty spot epidemiology: incidence as affected by distance from a
powdery mildew-infected apple orchard. Phytopathology 68, 896–899.
Ritchie, D.F. and Clayton, C.N. (1981) Peach tree short life: a complex of interacting factors. Plant Disease 65,
462–469.
Rizzo, D.M., Whiting, E.C. and Elkins, R.B. (1998) Spatial distribution of Armillaria mellea in pear orchards.
Plant Disease 82, 1226–1231.
Savage, E.F., Weinberger, J.H., Luttrell, E.S. and Rhoads, A.S. (1953) Clitocybe root rot – a disease of eco-
nomic importance in Georgia peach orchards. Plant Disease Reporter 37, 269–270.
Savage, E.F., Hayden, R.A. and Futral, J.G. (1974) Effect of soil fumigants on growth, yield and longevity of
Dixired peach trees. University of Georgia, Research Bulletin 148, 3–22.
Schnabel, G., Bryson, P.K., Bridges, W.C. and Brannen, P.M. (2004) Reduced sensitivity in Monilinia fructicola to
propiconazole in Georgia and implications for disease management. Plant Disease 88, 1000–1004.
Schnabel, G., Ash, J.S. and Bryson, P.K. (2005) Identification and characterization of Armillaria tabescens from
the southeastern United States. Mycological Research 109, 1208–1222.
Sierra, A.P., Whitehead, D.S. and Whitehead, M.P. (1999) Investigation of a PCR-based method for the routine
identification of British Armillaria species. Mycological Research 103, 1631–1636.
Snowdon, A.L. (1990) A Color Atlas of Post-Harvest Diseases and Disorders of Fruit and Vegetables. Vol. 1.
General Introduction and Fruits. Wolfe Scientific Ltd, London.
Soto-Estrada, A., Förster, H. and Adaskaveg, J.E. (2003) New fungicides and inoculum-precipitation based
application strategies for managing peach rust in California. Plant Disease 87, 1094–1101.
Strand, L. (1999) Integrated Pest Management for Stone Fruits. University of California Statewide Integrated
Pest Management Program. University of California Agricultural and Natural Resources, Publication No.
3389. University of California, Oakland, California.
Sutton, T.B. and Clayton, C.N. (1972) Role and survival of Monilinia fructicola in blighted peach branches.
Phytopathology 62, 1369–1373.
Tamm, L. and Flückiger, W. (1993) Influence of temperature and moisture on growth, spore production and
conidial germination of Monilinia laxa. Phytopathology 83, 1321–1326.
Termorshuizen, A.J. (2000) Ecology and epidemiology of Armillaria. In: Fox, R.T.V. (ed.) Armillaria Root Rot:
Biology and Control of Honey Fungus. Intercept, Andover, UK, pp. 45–64.
Weinhold, A.R. (1961) The orchard development of peach powdery mildew. Phytopathology 51, 478–481.
406 J.E. Adaskaveg et al.

White, E.E., Dubertz, C.P., Cruickshank, M.G. and Morrison, D.J. (1998) DNA diagnostic for Armillaria species
in British Columbia: within and between species variation in the IGS-1 and IGS-2 regions. Mycologia 90,
125–131.
Wilcox, W.F. and Ellis, M.A. (1989) Phytophthora root and crown rot of peach in the eastern Great Lakes
region. Plant Disease 73, 794–798.
Wilkins, B.S., Ebel, R.C., Dozier, W.A., Pitts, J., Eakes, D.J., Himelrick, D.G., Beckman, T. and Nyczepir, A.P.
(2002) Field performance of Guardian (TM) peach rootstock selections. HortScience 37, 1049–1052.
Wilson, E.E. (1937) The shot-hole disease of stone-fruit trees. California University Agriculture Experiment
Station Bulletin 608, 3–40.
Yarwood, C.E. (1939) Powdery mildews of peach and rose. Phytopathology 29, 282–284.
Zehr, E.I., Miller, R.W. and Smith, F.H. (1976) Soil fumigation and peach rootstocks for protection against
peach tree short life. Phytopathology 66, 689–694.
Zehr, E.I., Luszcz, L.A., Olien, W.C., Newall, W.C. and Toler, J.E. (1999) Reduced sensitivity in Monilinia fructi-
cola to propiconazole following prolonged exposure in peach orchards. Plant Disease 83, 913–916.
 16 Diseases Caused by Prokaryotes –
Bacteria and Phytoplasmas

D.F. Ritchie,1 M. Barba2 and M.C. Pagani3

1Department of Plant Pathology, North Carolina State University, Raleigh,
North Carolina, USA
2Istituto per la Patologia Vegetale, Rome, Italy
3BASF Corporation, Research Triangle Park, North Carolina, USA

16.1 Introduction 407

16.2 Bacterial Diseases 411
Bacterial canker complex 411
Bacterial spot 416
Crown gall 418
Phony peach disease 423
16.3 Phytoplasma Diseases 424
Peach X-disease 424
Peach yellows 426
Peach rosette 427
Peach yellow leaf roll 428
European stone fruit yellows 429
16.4 Summary 431

16.1 Introduction Mollicutes, order Mycoplasmatales. Unlike myco-

Bacteria and phytoplasmas are prokaryotic
microorganisms. Prokaryotes lack a nucleus
and other membrane-enclosed organelles.
Both bacteria and phytoplasmas are single-
celled or capable of living as single cells. The
cellular membrane of bacteria is surrounded
by a rigid cell wall whereas phytoplasmas
lack the cell wall, being enclosed only by a
cellular membrane and thus varying in shape
(i.e. pleomorphic). Phytoplasmas were previ-
ously known as ‘mycoplasma-like organisms’
and taxonomically are placed in the class
plasmas of humans and animals, phytoplasmas
have not yet been cultured on synthetic media.
In their plant hosts they are localized in phloem
sieve tubes, which they can occlude completely
(Fig. 16.1/Plate 141). Multiple years are usu-
ally required for entire colonization of woody
plant hosts. Thus, at an early stage of infection,
trees often express symptoms only on a single
branch. Most phytoplasmas overwinter in the
roots of the diseased plant.
The phytoplasma environment is either
inside the phloem vessels or in their phloem-
feeding leafhopper or psyllid insect vectors.

© CAB International 2008. The Peach: Botany, Production and Uses

(eds D.R. Layne and D. Bassi) 407
408 D.F. Ritchie et al.
In contrast, the bacterial pathogens of peach
can all be cultured on synthetic media and
except for Xylella fastidiosa, the phony peach
pathogen, do not exhibit a high degree of host
tissue specificity. Some bacterial pathogens
can also exist on plant surfaces as epiphytes
(Cross, 1966). Bacteria multiply through binary
fission, resulting in large populations within
days or even hours. The multiplication process
for phytoplasmas is not totally understood,
but may result from cellular fragmentation.
Presently, there are four bacterial-caused
diseases (Table 16.1) and five phytoplasma-
caused diseases (Table 16.2) described on
peaches. Phony peach is limited to the south-
eastern USA, but bacterial canker, bacterial
spot and crown gall occur worldwide where
favourable environments exist. The bacterial
pathogens cause leaf and fruit spots, cankers,
twig dieback, galls and tree death. Bacterial
Fig. 16.1. Electron micrograph of
phloem sieve tube occluded with cells
of phytoplasma.
canker and bacterial spot cause the greatest
economic loss but their occurrence is sporadic
and highly influenced by weather conditions.
Phytoplasma-caused diseases are potentially
devastating because once infection occurs
there is no cure, fruit production and quality
are reduced, and trees usually die within a
few years after infection. Management of bac-
teria- and phytoplasma-caused diseases relies
primarily upon pathogen exclusion using quar-
antines, certified pathogen-free plant material,
host resistance if available, production site
selection and vector control. Chemicals are
limited mostly to Cu compounds, which can
be phytotoxic to peach foliage, and the antibiotic
oxytetracycline, which is used in some regions.
Phytoplasmas cause vegetative and fruit
disorders that are expressed throughout the
entire tree or only in specific plant organs.
Flowers and vegetative growth may emerge
 Diseases Caused by Prokaryotes 409

Table 16.1. Diseases caused by bacteria, the pathogens, typical symptoms and primary control
methods.

Disease Pathogen Typical symptoms Primary control method(s)

Bacterial Pseudomonas Cankers, shoot dieback, Exclusion, cultural practices

canker complex syringae pv. syringae, leaf and flower bud death, that minimize tree stress
P. syringae pv. persicae root injury, tree death
Bacterial spot Xanthomonas Leaf and fruit lesions, Host resistance,
arboricola (syn. leaf chlorosis and bactericides, planting
campestris) pv. pruni defoliation, twig cankers site selection
Crown gall Agrobacterium Galls on lower stem/trunk Sanitation, exclusion,
tumefaciens near soil line and on roots biocontrol
Phony peach Xylella fastidiosa Flattened, compact canopy, Removal of diseased
shortened internodes, tree trees, eradication of
dwarfing, reduced fruit size secondary hosts
and production

prematurely in the spring, the internodes may
be shortened, lateral buds may proliferate,
and rosetting or premature defoliation may
occur. Fruit number and size may be reduced,
ripening may be non-uniform, fruit may drop
prematurely and taste bland or bitter. Terminal
dieback is common in the advanced stages of
phytoplasma-caused disease and tissues have
increased sensitivity to winter freezes. Trees
exhibit stunting and a general decline and often
die within 2–3 years after symptom onset.
Phytoplasmas are transmitted by phloem-
feeding insects such as leafhoppers and psyl-
lids. These insects acquire the pathogen while
feeding on infected plants. Insects generally
remain infectious for life although trans-ovar-
ian transmission (i.e. to offspring) has not
been shown. Phytoplasmas can be transmit-
ted by dodder (Cuscuta spp.) (Fig. 16.2/ Plate
142), but not mechanically in sap. They can be
transmitted through use of infected propaga-
tive material but are not known to be trans-
mitted in seeds and pollen.
Historically, phytoplasmas were detected
using indexing with indicator host plants such
as periwinkle (Catharanthus roseus) (Fig. 16.2/
Plate 142), electron microscopy, or DAPI
(4′,6′-diamidino-2-phenylindole) staining and
epifluorescence microscopy. Serological meth-
ods improved detection, but major advances
occurred with the use of genomic-based hybrid-
izations, PCR and nucleotide sequencing. All
diagnostic protocols, however, must take into
account that phytoplasmas generally occur in
relatively low titres, are unevenly distributed
within the tree, and their presence in different
parts of a tree varies with plant growth stage.
Diagnosis of phytoplasma-caused diseases
solely on symptomatology can result in incor-
rect conclusions. Different phytoplasmas can
cause similar symptoms. Environmental con-
ditions, host plant nutrition and host genetic
abnormalities also can produce symptoms
similar to those caused by phytoplasmas.
Phytoplasmas are classified into 15 groups
comprising more than 40 subgroups (Lee et al.,
2000; Montano et al., 2001). Those infecting peach
belong to two very distinct groups. The first is
the apple proliferation (AP) group (group 16Sr
X), which includes European stone fruit yellows
(ESFY) and peach yellow leaf roll (PYLR). ESFY
also belongs to subgroup B, whereas PYLR
belongs to subgroup C. The second is the X-dis-
ease group (group 16Sr III), which includes
peach X, peach yellows and peach rosette.
Management and control strategies for
all phytoplasma-caused diseases of peaches
are similar. They begin with preventive mea-
sures including use of certified pathogen-free
propagative material and the enforcement of
quarantines for the pathogen and vectors.
Where the disease is endemic, control of local
vector populations and elimination of woody
and herbaceous host plants that serve as res-
ervoirs for both the phytoplasmas and the
vector are recommended.
 410
Table 16.2. Diseases caused by phytoplasmas, the synonyms, classification, vector(s), geographical distribution and other woody, non-peach hosts.

Geographical Hosts other

Disease Synonyms Taxonomic classification Vector(s) distribution than peach

Peach X Peach western X, Group 16Sr III-A of Leafhoppers: Colladonus Canada, USA Sour and sweet cherry,
peach yellow Lee et al. (2000) geminatus, Colladonus Prunus virginiana
leaf roll montanus, Fieberiella florii,

D.F. Ritchie et al.

Paraphlepsius irroratus
Peach yellows Little peach, Related to, but distinct Leafhopper: Macropsis Canada, USA Almond, apricot,
peach red suture from, peach X-disease trimaculata Prunus salicina
phytoplasma
Peach rosette (PR) None Related to, but distinct Unknown USA P. salicina, almond,
from, peach X-disease apricot, cherry,
phytoplasma ornamental and
wild Prunus spp.
Peach yellow leaf roll None Group 16Sr X-C of Psyllid: Cacopsylla USA None known
(PYLR) Lee et al. (2000) pyricola
European stone Peach yellows, Group 16Sr X-B of Psyllid: Cacopsylla Europe Almond, apricot,
fruit yellows (ESFY) European peach Lee et al. (2000) pruni plum, Japanese plum,
yellows, peach Prunus serrulata
decline
 Diseases Caused by Prokaryotes 411

Fig. 16.2. Leaves of periwinkle (Catharanthus roseus), which can serve as an indicator plant for
phytoplasmas, colonized by dodder (Cuscuta spp.), which serves as a transmission bridge.

16.2 Bacterial Diseases Symptoms usually are first noticed in early

Bacterial canker complex
Bacterial canker and a decline syndrome
affect peaches and other stone fruits world-
wide. The disease is also known as bacterial
gummosis, sour sap or blast, and has been
recognized on stone fruit trees since the early
1900s (Cross, 1966). On peach, this disease is
associated with bud, shoot and branch die-
back and tree death (Davis and English, 1969).
Bacterial canker is a component of the peach
tree short life complex in the south-eastern
USA (Peterson and Dowler, 1965; Ritchie and
Clayton, 1981). The disease is also part of a
decline syndrome in France and New Zealand
(Prunier et al., 1970; Young, 1988). For infection
and disease development, usually trees must
be predisposed by adverse environmental or
biological factors (Davis and English, 1969;
Ritchie and Clayton, 1981; Cao et al., 2005).
Symptoms are variable within individual
trees and across the orchard. Healthy-appearing
trees can exist next to dead or dying trees.
spring as trees emerge from dormancy. Fruit
and leaf buds may fail to open (Fig. 16.3/
Plate 143), be delayed in opening, or open
and then cease growth, wilt and die back.
This may occur on individual branches or the
entire tree (Fig. 16.4/Plate 144). An elliptical
canker may be observed at the base of killed
buds (Fig. 16.3/Plate 143). Removal of the
outer bark commonly reveals reddish-brown,
water-soaked streaks in the wood (Fig. 16.5/
Plate 145). The discoloured tissue may extend
a few centimetres or encompass an entire
branch and extend into the trunk (Fig. 16.6/
Plate 146). There is an associated sour sap
odour with the diseased tissue. Water-soaked
areas may develop in the bark with the dis-
coloured tissue beneath the surface of the
outer bark (Fig. 16.7/Plate 147). Leaf and fruit
symptoms have not been observed on peaches.
Trees in their second to sixth growing season
are more prone to the bacterial canker com-
plex than are older trees. Injury from freezing
temperatures produces symptoms similar to
bacterial canker and may favour the infection
412 D.F. Ritchie et al.

Fig. 16.3. Dead peach buds and twig

canker associated with bacterial canker.

Fig. 16.4. Branch death and sucker growth from the rootstock associated with bacterial canker.
 Diseases Caused by Prokaryotes 413

Fig. 16.5. Wood with discoloured streaks and the collapse and wilt of newly emerged growth
associated with bacterial canker.

Fig. 16.6. Small branch killed by

Pseudomonas syringae pv. syringae with
the canker extending into a larger limb.
414 D.F. Ritchie et al.
process. Cold injury causes the cambial area
to develop a brown discoloration, with the bark
often splitting and separating easily from the
wood (Fig. 16.8/Plate 148). This characteristic
contrasts with bacterial canker, where the
diseased bark remains attached to the wood
(Fig. 16.5/Plate 145). Symptoms of bacterial
canker and cold injury merge as the season
progresses and isolation of the bacterial patho-
gen becomes difficult (Endert-Kirkpatrick
and Ritchie, 1988). Where Pseudomonas syrin-
gae pv. syringae is involved, roots remain func-
tional and often produce suckers (Fig. 16.4/
Plate 144). In contrast, for Pseudomonas syrin-
gae pv. persicae, roots also are attacked by the
pathogen (Prunier et al., 1970; Young, 1988).
Bacteria associated with bacterial canker
and decline were first investigated in Germany
in 1907 as a disease of cherry trees. In 1911, a
similar disease was reported in North America
(Cross, 1966). Investigations of a bacterial dis-
ease of stone fruits in England in the 1920s
Fig. 16.7. Water-soaked bark and
lightly discoloured wood in early
spring associated with bacterial canker.
led to the description of two additional spe-
cies, Pseudomonas prunicola and Pseudomonas
mors-prunorum. It was concluded that, except
for P. mors-prunorum, all such species were
strains of the lilac pathogen P. syringae van
Hall 1902 (Cross, 1966). Peterson and Dowler
(1965) associated P. syringae with peach tree
deaths in the south-eastern USA. Prunier et al.
(1970) reported that a non-fluorescent bacte-
rium was associated with a peach decline
syndrome in France and named the pathogen
P. syringae pv. persicae. In 1967, a similar bacte-
rial pathogen was associated with a decline
syndrome of peaches, nectarines and Japa-
nese plums in New Zealand (Young, 1988).
Thus, at least two pathovars of P. syringae
are associated with bacterial canker of peach
trees: (i) P. syringae pv. syringae van Hall
causes disease on many commercially grown
Prunus spp. in addition to peach; and (ii)
P. syringae pv. persicae is mostly limited to and
possibly more aggressive on peach, but also
 Diseases Caused by Prokaryotes 415

Fig. 16.8. Discoloured wood

approximately a week after the
occurrence of cold damage, with the
bark separating easily and appearing
undamaged.

Fig. 16.9. Non-fluorescent and fluorescent bacteria associated with the bacterial canker complex
cultivated on King’s medium B.

occurs on nectarine and plum. P. syringae pv. toxin syringomycin, while P. syringae pv. per-
syringae utilizes inositol and lactate, fluoresces sicae does not utilize these two carbohydrates,
on King’s medium B (Fig. 16.9/Plate 149) is non-fluorescent and produces the toxin
(Young and Triggs, 1994) and produces the persicomycin (Barzic, 1999).
416 D.F. Ritchie et al.
P. syringae consists of a ubiquitous group
of bacteria that can exist internally as plant
pathogens and externally as epiphytes (Cross,
1966; Dowler and Weaver, 1975). The patho-
gen can be disseminated by splashing rain,
possibly through pruning, and with contami-
nated nursery stock. Infections of the woody
portions of the tree are presumed to occur in
autumn through leaf scars, with symptom
expression in the following early spring as
trees emerge from dormancy. Infection and
disease development are favoured by wet and
cool environmental conditions. Widespread
infection and canker enlargement occur in trees
with high water content (Vigouroux, 1999).
Chemical sprays targeted at the patho-
gen have not adequately controlled bacterial
canker on peaches. The best success has been
achieved when management practices focus
on alleviating predisposing factors (Davis
and English, 1969; Ritchie and Clayton, 1981;
Cao et al., 2005). These include selection of
planting sites suited for peaches, proper root-
stock and cultivar selection adapted to the
growing region, reduction of nematode stress
through the use of pre-plant soil fumigation
and appropriate resistant rootstocks, and other
cultural practices that minimize tree stress
such as delaying pruning until late winter.
Bacterial spot
Bacterial spot, also known as bacteriosis, bac-
terial leaf spot and bacterial shot hole, infects
most Prunus species but causes significant
economic losses on peaches, nectarines and
plums. Bacterial spot was originally described
on Japanese plum (Prunus salicina Lindl.)
from Michigan in 1903 (Smith, 1903). The dis-
ease occurs in most countries where peaches
are grown including Argentina, Australia, Brazil,
Canada, France, Italy, Japan, New Zealand,
South Africa, the USA and Uruguay. Bacterial
spot is most common and severe where
peaches are grown in light, sandy soils and
moist, warm environment conditions occur.
Bacterial spot is caused by Xanthomonas arbori-
cola (syn. campestris) pv. pruni (Smith) (Vauterin
et al., 1995). The bacterium is a Gram-negative,
strict aerobe having an optimal growth
temperature of 24–30°C. Growth on YDC
medium (yeast–dextrose–calcium carbonate–
agar) or SPA medium (sucrose–peptone–agar)
for 48–72 h results in light yellow, lemon-
coloured, mucoid colonies.
Bacterial spot affects leaves, fruits and
twigs. Leaf lesions and chlorosis are usually
the first and most obvious symptoms observed
(Fig. 16.10/Plate 150). Lesions start as grey-
ish, angular areas approximately 1–3 mm in
width (Fig. 16.11/Plate 151). Lesion centres
become purple and necrotic and, if centres
abscise, leaves develop a shot-hole, tattered
appearance. Leaves may become chlorotic
and abscise. Lesions contain large numbers of
bacteria and bacterial streaming can be
observed when lesions are placed in a droplet
of water and viewed at 100–200× under a
microscope (Fig. 16.12/Plate 152). This is a
quick diagnostic method to differentiate bac-
terial-caused lesions from those caused by
other entities such as Cu or other pesticides.
Lesions from pesticide injury are almost
always circular, non-water soaked and not
angular when compared with bacterial spot
lesions. On highly susceptible varieties, mul-
tiple years of severe premature defoliation
result in reduced numbers of fruit buds,
reduced fruit crops and weakened trees.
Fruit symptoms are first visible as small,
angular, water-soaked lesions 3–5 weeks after
petal fall (Fig. 16.13/Plate 153). These early
infections can develop into deep, cavernous-
appearing lesions (Fig. 16.14/Plate 155).
Lesions that develop from later infections,
especially those after pit hardening, tend to
be confined more to the fruit surface (Fig.
16.15/Plate 154). Bacterial spot lesions on
fruit may be confused with peach scab lesions,
which are darker in colour, circular and usu-
ally more restricted to the surface of the peach
skin. Also, no bacterial streaming should be
detected from peach scab.
Spring cankers are first visible near time
of bloom and develop on the previous sea-
son’s twig growth, usually centred on a leaf
scar (Fig. 16.16/ Plate 156). Leaf and flower
buds fail to open or the canker extends down-
ward from the terminal bud, which fails to
open, resulting in a ‘black tip’ canker (Fig.
16.17/Plate 157). Cankers may extend several
centimetres from the dead buds and have a
 Diseases Caused by Prokaryotes
black, greasy, wet appearance. Cutting the
outer bark reveals brownish, water-soaked
bark that usually does not extend into the
wood (Fig. 16.18/Plate 158), differentiating it
from bacterial canker. During the growing
season, cankers can develop on the current
season’s twig growth; these are termed ‘sum-
mer cankers’ (Fig. 16.17/Plate 157).
The pathogen overwinters on peach trees
in association with buds, in protected areas
on the woody tree surface, and in leaf scars
that become infected during defoliation the
previous season (Shepard and Zehr, 1994).
Leaf-scar infections can result in spring
cankers that become active as new leaf tissue
emerges the following season, serving as a
major source of primary inoculum. Moisture
such as splashing or wind-blown rain or drip-
ping of dew is essential for bacterial dissemi-
nation to newly emerging leaves and fruit.
Fruit can become infected as they emerge
from the shuck if there is adequate moisture.
Frequent periods of rainfall and water conges-
tion of plant tissues are important for leaf
infection (Zehr and Shepard, 1996; Battilani
417
Fig. 16.10. Chlorotic peach leaves
with lesions caused by Xanthomonas
arboricola pv. pruni.
et al., 1999). Severe fruit infections are most
common when rainfall or extended periods of
water congestion occur during shuck split to
near pit hardening (Ritchie, 1993). Bacterial
spot is more severe where peaches are grown
in sandy than in heavier soils. This may be
associated with increased leaf and fruit injury
from wind-blown sand, from physiological
factors related to water potential, predisposi-
tion by nematodes, or a combination of these
(Zehr and Shepard, 1996). Leaf infections
occur as long as new leaf growth continues.
The time from infection until symptom
expression is influenced by temperature.
Under optimal bacterial growth conditions,
symptom expression may occur in 3–5 days
but under cool conditions this may take up to
14 days.
Peach cultivars vary greatly in susceptibil-
ity and the most effective control is through the
use of host plant resistance (Werner and Ritchie,
1986). Unfortunately, many resistant varieties
lack specific desirable fruit and marketing
characteristics (Okie, 1998). Most varieties
developed in dry regions where the disease
418 D.F. Ritchie et al.
Fig. 16.11. Newly formed, water-soaked, greyish,
angular bacterial spot lesions on peach leaf
caused by Xanthomonas arboricola pv. pruni.
does not occur are highly susceptible and,
when planted in regions where environmen-
tal conditions favour bacterial spot and the
pathogen is present or is introduced, fruit loss
can be severe. Once bacterial spot is estab-
lished in an orchard, disease control can be
very difficult (Ritchie, 1999). Injury from
blowing sand may be minimized by planting
and maintaining appropriate ground covers
in and near the orchard. Windbreaks appro-
priately placed to blunt the damaging effects
of strong winds, while still allowing for air
movement, may aid in reducing bacterial
spot.
Chemical controls have shown limited
efficacy but are most effective when manage-
ment programmes are started before the
infection of newly emerged leaves and fruits
occurs, with success depending on weather
conditions and disease pressure. Foliar-applied
chemicals consist primarily of Cu-containing
materials and in the USA also oxytetracycline
(Ritchie, 1999). Peach foliage is very sensitive
to Cu and severe phytotoxicity can result if
misused. Cu use focuses on three to five
applications from start of bud break through
shuck split, when only a limited amount of
foliage is present (Ritchie, 1999; Brannen et al.,
2007). The rate of elemental Cu is reduced
each subsequent application (Brannen et al.,
2007). After shuck split, very low rates of ele-
mental Cu or other materials registered for
use in the region have shown some success.
Sprays are usually applied at 7–14-day inter-
vals, depending upon rainfall and disease
pressure.
Crown gall
The greatest economic impact of this disease
occurs in the nursery from culling of symp-
tomatic trees. Galls commonly develop in the
lower stem or trunk region at the root crown
just below the soil line (Fig. 16.19/Plate 159),
but can occur on roots and occasionally aerial
plant parts. Galls may develop to several cen-
timetres in diameter. Tissue of newly formed
and expanding galls is soft and spongy. Young
trees may be stunted and older trees may
express symptoms typical of root problems.
On mature trees, galls often are not discov-
ered until the roots are exposed when trees
are removed. As galls age, they become
woody and usually are colonized by other
microbes, resulting in decay.
Agrobacterium belongs to the family of
nitrogen-fixing bacteria, Rhizobiaceae, and
consists of several species of bacteria that
induce hyperplastic plant growths. Agrobacte-
rium tumefaciens, also known as biovar 2, is
the species that induces galls in peach. Sev-
eral selective media have been developed to
aid in isolation and differentiation of the spe-
cies (biovars) (Brisbane and Kerr, 1983) and
 Diseases Caused by Prokaryotes 419

Fig. 16.12. Bacteria streaming from leaf lesion caused by Xanthomonas arboricola pv. pruni.

Fig. 16.13. Newly formed,

water-soaked, bacterial spot lesions
on peach fruit near the growth stage
of pit hardening.

PCR-based primers also have been developed the Ti-plasmid. A segment of the Ti-plasmid
(Haas et al., 1995). (the T-DNA) is transferred to the plant cell via
The tumourigenic ability is due to a conjugation-like process and incorporated
genes on a large bacterial plasmid termed into the plant genome. Loss of the Ti-plasmid
420 D.F. Ritchie et al.
renders the A. tumefaciens strains non-
tumourigenic. Such strains are classified as
Agrobacterium radiobacter.
Infection requires wounds and is favoured
by moist conditions and poorly drained soils.
Although the Ti-plasmid DNA transfer is
favoured by temperatures just below 20°C,
gall development occurs more rapidly at tem-
peratures greater than 20°C with galls visible
Fig. 16.14. Bacterial spot lesions on
fruit at harvest caused by early
infections occurring soon after shuck
split and before pit hardening.
Fig. 16.15. Bacterial spot lesions on
fruit at harvest which are confined to
the fruit surface and developed from
infections occurring after pit harden-
ing.
2–4 weeks after infection (Fullner et al., 1996).
Crown gall is more common in temperate
than tropical regions. Once plant cells are
transformed, bacterial presence no longer is
necessary for tumour development since the
T-DNA replicates with the host DNA. Gene
expression causes the plant to produce excess
amounts of indoleacetic acid and cytokinin,
which stimulate hyperplastic growth.
 Diseases Caused by Prokaryotes 421

Fig. 16.16. Bacterial spot spring canker (Xanthomonas arboricola pv. pruni) with a black, greasy
appearance.

Fig. 16.17. Terminal dieback and black tip canker, summer cankers on current year’s twigs, lesions
on leaves, and nodes where leaves have defoliated caused by Xanthomonas arboricola pv. pruni.
422 D.F. Ritchie et al.

Fig. 16.18. Discoloured bark tissue of spring canker Xanthomonas arboricola pv. pruni.

Fig. 16.19. Crown gall caused by

Agrobacterium tumefaciens on peach
trees recently dug from a nursery.
 Diseases Caused by Prokaryotes 423

Species of Agrobacterium are ubiquitous Phony peach disease

in the soil and can be isolated from agricul-
tural and non-agricultural soils (Bouzar and
Moore, 1987). The ability to exist in latent
infections and in soil adhering to root surfaces
allows for pathogen dissemination with nurs-
ery stock and through the movement of soil
and water (Moore, 1976). Under experimental
conditions, agrobacteria have survived in soil
for more than 2 years, with survival negatively
affected by soil temperatures greater than 34°C
and acidic conditions.
Crown gall is best controlled by preven-
tion of infections. This starts with a pathogen-
free nursery site, inspection of trees as
removed from the nursery and the planting of
quality, gall-free trees. Seed used for root-
stocks can be treated using the appropriate
biocontrol strain of A. radiobacter. Roots of
trees also can be treated with this agent at
planting time (Jones and Kerr, 1989). Removal
of the galls does not necessarily prevent galls
from occurring later, thus trees having galls
should be destroyed. Once trees have been
planted, infections can be minimized by avoid-
ing practices that cause injuries (e.g. tillage) to
the lower trunk and roots.
Phony (phoney) peach disease (PPD) reduces
fruit quality and the economic life of trees.
This disease is endemic in the south-eastern
USA, where symptoms were first described
in 1890. Sporadic epidemics have occurred
several times since then. Tree losses have
occurred from eastern Texas to the south-
eastern Atlantic states, being most severe in
Georgia, northern Florida and southern Ala-
bama (Wells et al., 1983; Mizell et al., 2003).
Diseased trees appear dwarfed compared
with non-diseased trees. Dark green leaves on
branches with shortened internodes give the
tree a compact, flattened, umbrella-like can-
opy (Fig. 16.20/Plate 160). Infected trees tend
to bloom several days earlier than healthy
trees and retain their leaves longer in autumn.
Another typical symptom occurring 3–5 years
after infection is the production of progres-
sively smaller, more colourful, but non-
marketable fruit (Evert and Smittle, 1989).
PPD is caused by the fastidious xylem-
limited, Gram-negative bacterium, Xylella fas-
tidiosa Wells et al. (Wells et al., 1983). Detection
of X. fastidiosa was hampered because of

Fig. 16.20. Peach tree expressing phony peach disease symptoms. Leaves are greener and denser
and internodes are shortened, giving the tree a compact, flat canopy with an ‘umbrella-like’
appearance. (Courtesy of R.F. Mizell III, University of Florida, Quincy, Florida, USA.)
424 D.F. Ritchie et al.
difficulty in culturing the pathogen. In 1973,
PPD was first associated with a rickettsia-like
bacterium before finally being cultured in 1983.
PCR primers have proved useful to identify
naturally infected field samples (Rodrigues
et al., 2003). Genetically distinct strains of X.
fastidiosa are responsible for a variety of eco-
nomically important plant diseases. Strains
of X. fastidiosa cause citrus variegated chlo-
rosis (CVC), Pierce’s disease, PPD, plum leaf
scald (PLS) and leaf scorch diseases in elms,
almonds, maples, oleander, pear, oaks, syca-
mores and coffee (Nunes et al., 2003). A phylo-
genetic analysis based on partial sequences of
the gyrB gene placed X. fastidiosa strains into
three clusters: (i) grapevine-associated strains;
(ii) citrus–coffee strains; and (iii) a cluster
composed of all other strains (Rodrigues et al.,
2003). A common aetiology between strains
that cause PPD on peaches and PLS on plums
was shown (Wells et al., 1981; Abrahams and
Norton, 1994).
Plants of more than 30 families can serve
as natural reservoirs and secondary hosts for
the pathogen. The most important for PPD
are wild Prunus species and grasses such as
johnsongrass (Sorghum halapense) (Mizell et al.,
2003). X. fastidiosa enters the peach xylem tis-
sue via the feeding of sharpshooter leafhop-
per vectors, primarily Homalodisca coagulata
and Oncometopia nigricans. The polyphagous
H. coagulata has a moderate feeding prefer-
ence for plums and a low preference for
peaches. Insect numbers also are reduced on
trees that express PPD symptoms. The xylem
fluid chemistry of the scion/rootstock combi-
nation affects the feeding behaviour and per-
formance of PPD vectors (Andersen et al.,
1994). X. fastidiosa multiplies and spreads sys-
temically throughout the xylem, occluding
the vessels and restricting water movement
(Mizell et al., 2003). X. fastidiosa is usually
much more abundant in peach roots than in
stems or leaves, which contrasts with plums,
where it is homogeneously distributed through-
out the tree (Evert and Smittle, 1989). This is
considered important as it results in lack of
secondary spread within peach orchards and,
thus, peach may be a ‘dead-end’ host.
There is no cure for PPD and so control is
based on preventing spread through early
detection and removal of diseased trees. Wild
plums within approximately 400 m should be
removed and weeds controlled in and close to
peach orchards to reduce reservoir hosts for
the pathogen and the insect vectors. Severe
summer pruning should be avoided since
intense regrowth is attractive to leafhoppers.
Control of leafhoppers with routine spraying
has not proved effective. New orchards
should not be established close to infected
Prunus trees before removal and destruction
of such trees (Mizell et al., 2003).
16.3 Phytoplasma Diseases
Peach X-disease
Peach X-disease was described in California
in 1931. It also occurs in eastern fruit-growing
areas of North America, where it continues to
cause economic losses to stone fruits, but has
not been reported outside North America
(EPPO/CABI, 1997b). In addition to peach,
the disease is economically important on tart
and sweet cherries, Japanese plum (P. salic-
ina), almonds and apricots, and occurs on
some wild Prunus species (e.g. Prunus virgini-
ana), which serve as important pathogen res-
ervoirs (Kirkpatrick et al., 1995). Eastern and
western strains of the pathogen are similar
and the term X-disease is currently used to
describe this group of yellows-inducing Pru-
nus pathogens (Kirkpatrick et al., 1995).
During the early stages of infection, trees
show very irregularly distributed symptoms.
The phytoplasma pathogen requires more
than 2 years to colonize a mature tree com-
pletely (Kirkpatrick et al., 1995). Symptoms
occur on the leaves as irregularly distributed
yellow or necrotic spots (Fig. 16.21/Plate 161).
The number of lesions increases during the
season and lesion centres may abscise, creat-
ing a shot-hole appearance. Leaves are rolled
and have a pale green colour, conferring an
appearance of reduced tree vigour. Often
infected leaves at the base of the shoots
abscise, inducing a terminal rosetting (Fig.
16.22/Plate 162). Symptomatic branches are
more sensitive to freezes and young trees can
die within 1–3 years after infection (Gilmer
and Blodgett, 1976). Fruit size and yield are
 Diseases Caused by Prokaryotes 425

Fig. 16.21. X-disease phytoplasma-

infected peach with necrotic lesions
non-uniformly distributed on rolled
and pale green leaves.

Fig. 16.22. X-disease phytoplasma-infected trees exhibit a loss of vigour, leaves have a pale green
colour and severely infected branches die.
426 D.F. Ritchie et al.
greatly reduced; remaining fruit ripen several
days later than normal and often have an
insipid flavour.
Grannet and Gilmer (1971) demonstrated
the association of a phytoplasma with
X-disease in various Prunus species. This
phytoplasma belongs to the X-disease group
(group 16Sr III) (Lee et al., 2000). Analysis of
16S rDNA of phytoplasmas associated with
individual trees affected by little peach, red
suture and rosette strongly suggests that
these phytoplasmas are closely related,
exhibiting greater than 99% similarity (Scott
and Zimmerman, 2001), but the diseases are
considered to be different because of epide-
miological properties including geographical
distribution, host range and insect vectors.
The greatest potential for infection occurs
during late spring and summer. Peach
X-disease has been transmitted by grafting to
several Prunus spp. and through dodder (Cus-
cuta spp.) to periwinkle and other herbaceous
plants such as celery, carrot, parsley, tobacco
and tomato (Gilmer and Blodgett, 1976). The
primary method of field dissemination is by
leafhoppers, with Colladonus montanus, Colla-
donus geminatus, Fieberiella florii and Paraphlep-
sius irroratus considered the most important.
Each vector has its own ecological niche, which
characterizes the natural spread to peach
orchards. For C. montanus, the probability to
infect peach orchards is greater in areas where
sugarbeet or other dicotyledonous plants are
present because this vector overwinters and
multiplies on these plants. Wild cherries, such
as bitter cherry (Prunus emarginata) and
chokecherry (P. virginiana), serve as reservoirs
of this phytoplasma in the eastern USA where
the vector is the cherry leafhopper, F. florii,
which is very efficient at transmitting the
phytoplasma from wild cherries to peaches.
In areas where P. irroratus is the primary vec-
tor, it feeds on infected monocotyledonous
plants during the day and at night moves to
peach or other woody hosts (Rosenberger and
Jones, 1977; Kirkpatrick et al., 1995).
Diseased peach trees should be removed
and destroyed as soon as detected. Cultural
practices that reduce the risk of disease spread
include control of the leafhopper vector pop-
ulation and elimination of woody and herba-
ceous plants that serve as reservoirs for the
phytoplasma or that allow the vector to over-
winter and multiply. In the eastern USA,
removal of wild chokecherry within 200 m of
orchards reduced disease incidence (Rosen-
berger and Jones, 1977).
Peach yellows
Peach yellows, little peach and red suture are
caused by the same phytoplasma (Kirkpatrick,
1995b; EPPO/CABI, 1997c), although there are
epidemiological differences (Larsen and Water-
worth, 1995). Peach yellows epidemics with
economic losses occurred during the 19th and
early 20th centuries along the eastern US coastal
states from Maryland to South Carolina. The
disease has not been reported in far western
and southern states (south of South Carolina)
or outside the USA (EPPO/CABI, 1997c).
Peach red suture, however, has been obser-
ved in eastern Canada, Russia and possibly
France and Israel (Larsen and Waterworth,
1995). All cultivars of peach and nectarine are
susceptible. Almonds, apricots and Japanese
plum (P. salicina) can serve as reservoirs for
the pathogen (EPPO/CABI, 1997c).
Disease symptoms are as follows.
● Peach yellows: Flowers and foliage of in-
fected trees emerge 2–4 weeks earlier
than on healthy trees. Leaves are small and
narrow, inwardly rolled with a down-
ward droop, chlorotic and may develop
red spots. Latent-infected buds initiate
new growth by producing multiple shoots,
especially on the internal portion of the
canopy, giving the tree a bushy appear-
ance. Terminal dieback of twigs and
branches is common in advanced stages
of the disease. Severely diseased trees die
within 2–4 years. Fruits ripen and colour
several days earlier than normal (Fig. 16.23/
Plate 163) and, although of normal or
larger size, have a bland or bitter flavour
(Pine and Gilmer, 1976).
● Little peach: Symptoms are similar as for
peach yellows but differ in two distinc-
tive characteristics. Young leaves initially
are greener than normal and later become
yellow. Fruits are reduced in size and
ripen several days to 3 weeks later than
normal (Pine and Gilmer, 1976).
 Diseases Caused by Prokaryotes
● Red suture: Leaves develop a yellowish-
green appearance soon after petal fall
and become greenish-bronze just before
harvest. The most distinctive symptom
occurs on fruits as a premature ripening
and softening in the suture area, while
the other part of the fruit remains green
and hard. The suture area is prominent,
bumpy, swollen, accompanied by an in-
tense dark red to purple colour for red
cultivars and a deep yellow for yellow cul-
tivars, and an intense internal colour along
the suture (Klos, 1976; Larsen and Water-
worth, 1995). Fruit flavour is often insipid.
Jones et al. (1974) demonstrated that a phyto-
plasma is responsible for the peach yellows
symptoms. This phytoplasma belongs to the
X-disease group (group 16Sr III) (Lee et al.,
2000), but is distinct from the peach X-disease
phytoplasma (Kirkpatrick, 1995b; EPPO/
CABI, 1997c). Phytoplasmas associated with
little peach, red suture and western X-disease
are more than 99% similar (Scott and Zim-
merman, 2001). These phytoplasmas also are
closely related to those causing peach rosette.
Peach yellows was transmitted experimen-
tally by grafting to several Prunus spp. and
through dodder (Cuscuta spp.) to periwinkle
427
Fig. 16.23. Tree infected with peach
yellows phytoplasma showing a bushy
appearance on ends of branches and
premature colouring and ripening of
fruit. (Courtesy of A. Ragozzino.)
(Jones et al., 1974). Symptom expression is
influenced by the site of inoculation. This
phytoplasma moves downward more rapidly
than upward. Hence, infection near the apical
portion of the tree causes symptom expres-
sion much earlier than does infection nearer
the soil (Pine and Gilmer, 1976). Peach yel-
lows and little peach are transmitted by
the plum leafhopper Macropsis trimaculata,
whereas the vector of red suture is unknown
(EEOP/CABI, 1997c). Because plums are the
preferred host of M. trimaculata, latently
infected plums may serve as reservoirs for
transmission to peaches. This plum leafhop-
per vector does not occur in Europe (EEOP/
CABI, 1997c).
Diseased peach trees should be removed
and destroyed as soon as detected. Plums
located near peach orchards should be care-
fully monitored and removed if they test pos-
itive for peach yellows.
Peach rosette
Peach rosette (PR) was first described in the
state of Georgia, USA, in 1881, and later
reported in other south-eastern states as far
428 D.F. Ritchie et al.
west as Texas (Kenknight, 1976). Initially, it
was considered a southern strain of peach
yellows. The disease has not been reported
from western states or outside the USA
(Kirkpatrick, 1995a). Kirkpatrick et al. (1975)
demonstrated experimentally that a phyto-
plasma is responsible for the symptoms. The
disease is sporadic in occurrence and cur-
rently considered of minor importance.
The defining symptom of PR is the short-
ening of internodes on new shoots, creating
tight clusters of leaves that cause a rosette or
witches’ broom appearance. In early summer,
premature defoliation occurs at the base of
the shoots, leaving tufts of young leaves near
the shoot tip. The interior tree canopy fails
to develop secondary shoots, resulting in a
sparse appearance. Leaves are normal in size
but may become chlorotic. Blossom and fruit
production are reduced and the few fruits
produced ripen irregularly and abscise pre-
maturely. Infected trees generally survive the
season that the infection occurred, but in
severe cases die within a year after onset of
symptoms (Kenknight, 1976; Kirkpatrick,
1995a). These two characteristics (sudden
reduction of fruit production and tree death)
are different from peach yellows. The PR phy-
toplasma belongs to the X-disease group
(group 16Sr III) (Lee et al., 2000), but is dis-
tinct from the peach X-disease phytoplasma
(Kirkpatrick, 1995a; EPPO/CABI, 1997a).
Peach, Japanese plum (P. salicina) and
wild plum species (Prunus augustifolia, Prunus
hortulana, Prunus munsoniana) are important
native hosts of PR (Kenknight, 1976). Rosette
symptoms occurred within 4–11 months after
experimental transmission to peach, plum,
almond and cherry seedlings by grafting
(Kirkpatrick et al., 1975). The PR phytoplasma
has been transmitted through dodder to peri-
winkle, tomato and tobacco (Kenknight, 1976).
Field surveys indicate spread from tree
to tree, but infected peach trees apparently
are not the primary reservoir of the phyto-
plasma since such trees generally die within a
year after infection. Diseased peach trees are
usually first detected at the border of the
orchard, suggesting that the pathogen is
introduced from wild plums growing near
the orchard. Infected plums survive for years
and serve as reservoirs of the phytoplasma
(Kenknight, 1976; Scott and Zimmerman,
2001). No insect vector(s) has been identified.
Control measures include removal and
destruction of diseased trees as soon as
detected. Wild plums should not be allowed
to grow near peach orchards.
Peach yellow leaf roll
Peach yellow leaf roll (PYLR) and western
X-disease are the two major phytoplasma-
caused diseases of peaches in California
(Blomquist and Kirkpatrick, 2002). Nyland and
Schlocker (1951) first reported PYLR in 1951. It
apparently is restricted to a few northern Cali-
fornia counties where it periodically has caused
severe tree losses (Nyland and Schlocker, 1951;
Purcell et al., 1981; Blomquist and Kirkpatrick,
2002). Until recently, PYLR was considered a
different manifestation of peach X-disease.
PYLR has been observed only on peach
trees. Leaves of infected trees have normal
size, but are yellow and roll downward with
a tendency to curve toward the stem. The leaf
midribs and lateral veins are enlarged and
affected leaves abscise prematurely. Fruit pro-
duction decreases, a rapid tree decline occurs,
and the tree can die within 3 years after the
onset of symptoms. Late-season leaf symp-
toms sometimes result in shot holes, thus
resembling peach X-disease (Blomquist and
Kirkpatrick, 2002).
The PYLR phytoplasma is a member of
the AP (apple proliferation) group (group
16Sr X) (Kison et al., 1997), which includes
phytoplasmas infecting stone fruits in Europe
(European stone fruit yellows, ESFY), pears
(pear decline, PD) and apples (AP). Restric-
tion fragment length polymorphism analysis
of PCR-amplified rDNA differentiated the
PYLR phytoplasma from the AP and ESFY
phytoplasmas, but not from the PD phyto-
plasma (Kison et al., 1997). Worldwide, peach
and pear orchards can be found adjacent yet
PYLR occurs only in California, suggesting
that the PYLR sub-strain of the PD phyto-
plasma evolved in California (Blomquist and
Kirkpatrick, 2002).
PYLR is most evident in peach orchards
adjacent to pears, thus pear orchards are
considered the primary reservoir for PYLR
 Diseases Caused by Prokaryotes 429

(Purcell et al., 1981; Blomquist and Kirkpatrick,
2002). Psylla insects (Homoptera: Psyllidae)
transmit phytoplasmas that belong to the AP
group, which include the PD and the ESFY
phytoplasmas (Carraro et al., 1998; Frisinghelli
et al., 2000). PYLR phytoplasma also is trans-
mitted by psylla, which is considered the pri-
mary vector (Blomquist and Kirkpatrick, 2002).
In late summer to early autumn, psylla move
from pears to the surrounding vegetation and
thus transmit the PYLR phytoplasma to other
hosts. The use of effective insecticides in pear
orchards after fruit harvest for reducing late-
season psylla populations, thus restricting
psylla movement to peach orchards, has effec-
tively reduced PYLR in northern California
(Blomquist and Kirkpatrick, 2002).
European stone fruit yellows
Decline diseases of apricot and Japanese plum
were observed in France and Italy in the early
1900s and named apricot chlorotic leaf roll
(ACLR) and plum leptonecrosis (PLN) or plum
decline. Similar diseases were later described
on peach and almond in Germany and Spain.
Today, all these decline diseases are referred to
as ESFY (European stone fruit yellows) (Lorenz
et al., 1994). The disease occurs on Prunus spe-
cies in all Mediterranean countries and as far
north as Germany. It is economically the most
important phytoplasma-caused disease of stone
fruits in Europe (Poggi-Pollini et al., 2001).
Symptoms that develop soon after infec-
tion are irregularly distributed on the branches
inoculated by the insect vector. Two or more
years are required to completely colonize a
mature tree. The greatest potential for infection
occurs in late summer (August–September)
when symptoms are particularly evident on
the foliage. Leaves of infected branches are
small in size, roll longitudinally upward, are
yellow to red in colour and thick and brittle
(Fig. 16.24/Plate 164). Leaf midribs and lat-
eral veins are enlarged and affected leaves
may become necrotic and abscise prematurely
(Fig. 16.25/Plate 165). Diseased trees have
reduced vigour (Fig. 16.26/Plate 166), are
more sensitive to winter freezes and decline
gradually, with reduced fruit productivity
(Poggi-Pollini et al., 2001). Early emergence of
flowers and leaves may occur during the win-
ter, allowing for easy differentiation from

Fig. 16.24. Leaves on branch infected with European stone fruit yellows phytoplasma (left) are
small in size, rolled upward longitudinally and have a pale yellow colour compared with leaves on
a non-infected branch (right). (Courtesy of A. Ragozzino.)
430 D.F. Ritchie et al.
healthy trees. Symptom expression is influ-
enced by the virulence of individual phyto-
plasma strains and the susceptibility of the
rootstock and scion. Some strains are very
aggressive, especially when susceptible root-
stocks (e.g. peach seedlings, ‘GF 677’, ‘Rubira’,
‘Montclar’, ‘Rutgers Red Leaf’) are used.
These aggressive strains also cause high mor-
tality when scions are grafted on to tolerant
rootstocks (Kison and Seemüller, 2001).
ESFY is caused by a phytoplasma belong-
ing to the AP group (group 16Sr X) (Lee et al.,
2000). It differs from the PYLR and X-disease
phytoplasmas, which occur in North America
(Dosba et al., 1991; Ahrens et al., 1993; Seemüller
et al., 1998). The ESFY phytoplasma persists in
the aerial portion of peach trees during the
dormant season and can be dormant-bud
Fig. 16.25. Peach tree infected
with European stone fruit yellows
phytoplasma with leaf midrib and
lateral veins enlarged and necrotic.
(Courtesy of L. Giunchedi.)
transmitted by grafting (Seemüller et al., 1998;
Jarausch et al., 1999). Cross-inoculation experi-
ments with phytoplasma isolates infecting vari-
ous Prunus species suggest that ACLR, PLN
and yellowing and decline of almond and peach
have a common aetiology (Dosba et al., 1991).
Ahrens et al. (1993) confirmed that the phyto-
plasmas associated with naturally infected apri-
cot, plum, almond and peach are closely related.
The psylla, Cacopsylla pruni, is the major
natural vector. The phytoplasma is acquired
when the psylla feed on infected trees for 2–4
days and they remain infective until their
death. A latent period of 2–3 weeks between
acquisition and transmission may occur
(Carraro et al., 1998). Several wild Prunus spe-
cies (Prunus spinosa, Prunus cerasifera, Prunus
domestica) serve as reservoirs for the ESFY
 Diseases Caused by Prokaryotes 431

Fig. 16.26. Two trees of the same age of cultivar ‘Baby Gold 7’. The tree in foreground is infected
with the European stone fruit yellows phytoplasma (exhibiting reduced vigour); compare with the
non-infected tree in the background. (Courtesy of A. Ragozzino.)

phytoplasma and as hosts for the insect vec-
tor (Carraro et al., 2002).
Although ESFY is present in many Euro-
pean countries, spread currently is reduced
through phytosanitary regulations. Other
management practices include removal and
destruction of diseased trees and eradication
of wild Prunus spp. when identified, and the
control of psylla populations.
16.3 Summary
Although there are relatively few diseases of
peach caused by bacteria and phytoplasmas,
these can cause serious economic fruit losses,
render trees non-productive and shorten tree
life. Diagnosis based on symptoms ranges
from relatively easy for bacterial spot to diffi-
cult or unreliable for the phytoplasma-caused
diseases, for which confirmation must include
serological or genomic methods. Disease
management is based on prevention and
starts with pathogen exclusion from a region
or individual orchard by use of quarantines
and certified plant material. Antibacterial
chemicals are few, have limited efficacy and
may produce unacceptable phytotoxicity. Host
plant resistance, when available in adapted
cultivars, is very effective.

References

Abrahams, B. and Norton, J. (1994) Transmission of plum leaf scald or phony peach disease, Xylella fastidiosa
Wells, by two budding methods in peach and plum. HortScience 29, 736.
Ahrens, U., Lorenz, K.H. and Seemüller, E. (1993) Genetic diversity among mycoplasma-like organisms
associated with stone fruit diseases. Molecular Plant–Microbe Interactions 6, 686–691.
432 D.F. Ritchie et al.

Andersen, P., Brodbeck, B. and Mizell, R. (1994) Influence of xylem fluid chemistry of Prunus spp. on the
abundance and performance of adult Homalodisca coagulata. HortScience 29, 493.
Barzic, M.R. (1999) Persicomycin production by strains of Pseudomonas syringae pv. persicae. Physiological
and Molecular Plant Pathology 55, 243–250.
Battilani, P., Rossi, V. and Saccardi, A. (1999) Development of Xanthomonas arboricola pv. pruni epidemics
on peaches. Journal of Plant Pathology 81, 161–171.
Blomquist, C.L. and Kirkpatrick, B.C. (2002) Identification of phytoplasma taxa and insect vectors of peach
yellow leaf roll disease in California. Plant Disease 86, 759–763.
Bouzar, H. and Moore, L.W. (1987) Isolation of different Agrobacterium biovars from natural oak savanna and
tall grass prairie. Applied and Environmental Microbiology 53, 717–721.
Brannen, P., Horton, D., Bellinger, B. and Ritchie, D. (2007) Southeastern Peach, Nectarine and Plum Pest Manage-
ment and Cultural Guide. Georgia Extension Bulletin No. 1171 (revised annually). University of Georgia
Cooperative Extension Service, College of Agricultural and Environmental Sciences, Athens, Georgia.
Brisbane, P.G. and Kerr, A. (1983) Selective media for three biovars of Agrobacterium. Journal of Applied
Bacteriology 54, 425–431.
Cao, T., Duncan, R.A., McKenry, M.V., Shachel, K.A., DeJong, T.M. and Kirkpatrick, B.C. (2005) Interaction
between nitrogen-fertilized peach trees and expression of syrB, a gene involved in syringomycin produc-
tion in Pseudomonas syrinage pv. syringae. Phytopathology 95, 581–586.
Carraro, L., Osler, R., Loi, N., Ermacora, P. and Refatti, E. (1998) Transmission of European stone fruit yellows
phytoplasma by Cacopsylla pruni. Journal of Plant Pathology 80, 233–239.
Carraro, L., Ferrini, F., Ermacora, P. and Loi, N. (2002) Role of wild Prunus species in the epidemiology of
European stone fruit yellows. Plant Pathology 51, 513–517.
Cross, J.E. (1966) Epidemiological relations of the pseudomonad pathogens of deciduous fruit trees. Annual
Review of Phytopathology 4, 291–310.
Davis, J.R. and English, H. (1969) Factors related to the development of bacterial canker in peach. Phytopa-
thology 59, 588–595.
Dosba, F., Lansac, M., Mazy, K., Garnier, M. and Eyquard, J.P. (1991) Incidence of different diseases associ-
ated with mycoplasmalike organisms in different species of Prunus. Acta Horticulturae 283, 311–320.
Dowler, W.M. and Weaver, D.J. (1975) Isolation and characterization of fluorescent pseudomonads from ap-
parently healthy peach trees. Phytopathology 65, 233–236.
Endert-Kirkpatrick, E. and Ritchie, D.F. (1988) Involvement of pH in the competition between Cytospora
cincta and Pseudomonas syringae pv. syringae. Phytopathology 78, 619–624.
EPPO/CABI (1997a) Peach rosette phytoplasma. In: Smith, I.M., McNamara, D.G., Scott, P.R. and Holderness,
M. (eds) Quarantine Pests for Europe, 2nd edn. CAB International, Wallingford, UK, pp. 1036–1038.
EPPO/CABI (1997b) Peach X-disease phytoplasma. In: Smith, I.M., McNamara, D.G., Scott, P.R. and Holderness,
M. (eds) Quarantine Pests for Europe, 2nd edn. CAB International, Wallingford, UK, pp. 1039–1043.
EPPO/CABI (1997c) Peach yellows phytoplasma. In: Smith, I.M., McNamara, D.G., Scott, P.R. and Holderness,
M. (eds) Quarantine Pests for Europe, 2nd edn. CAB International, Wallingford, UK, pp. 1044–1047.
Evert, D. and Smittle, D. (1989) Phony disease influences peach leaf characteristics. HortScience 24, 1000–
1002.
Frisinghelli, C., Delaiti, L., Grando, M.S., Forti, D. and Vindimian, M.E. (2000) Cacopsylla costalis (Flor 1861),
as a vector of apple proliferation in Trentino. Journal of Phytopathology 148, 425–431.
Fullner, K.J., Lara, J.C. and Nester, E.W. (1996) Pilus assembly by Agrobacterium T-DNA transfers genes. Sci-
ence 273, 1107–1109.
Gilmer, R.M. and Blodgett, E.C. (1976) X-disease. In: Virus Diseases and Noninfectious Disorders of Stone
Fruits in North America. USDA Agriculture Handbook No. 437. US Department of Agriculture–Agricul-
tural Research Service, Washington, DC, pp. 145–155.
Grannet, A.L. and Gilmer, R.M. (1971) Mycoplasma associated with X-disease in various Prunus species.
Phytopathology 61, 1036–1037.
Haas, J.H., Moore, L.W., Ream, W. and Manulis, S. (1995) Universal PCR primers for detection of phytopatho-
genic Agrobacterium strains. Applied and Environmental Microbiology 61, 2879–2884.
Jarausch, W., Lansac, M. and Dosba, F. (1999) Seasonal colonization pattern of European stone fruit yellows
phytoplasmas in different Prunus species by specific PCR. Journal of Phytopathology 147, 47–54.
Jones, A.L., Hooper, G.R., Rosenberger, D.A. and Chevalier, J. (1974) Mycoplasma-like bodies associated with
peach and periwinkle exhibiting symptoms of peach yellows. Phytopathology 64, 1154–1156.
Jones, D.A. and Kerr, A. (1989) Agrobacterium radiobacter strain K1026, a genetically engineered derivative
of strain K84, for biocontrol of crown gall. Plant Disease 73, 15–18.
 Diseases Caused by Prokaryotes 433

Kenknight, G. (1976) Peach rosette. In: Virus Diseases and Noninfectious Disorders of Stone Fruits in North
America. USDA Agriculture Handbook No. 437. US Department of Agriculture–Agricultural Research
Service, Washington, DC, pp. 73–76.
Kirkpatrick, B.C. (1995a) Peach rosette. In: Ogawa, J.M., Zehr, E.I., Bird, G.W., Ritchie, D.F., Uriu, K. and
Uyemoto, J.K. (eds) Compendium of Stone Fruit Diseases. APS Press, St. Paul, Minnesota, pp. 56–57.
Kirkpatrick, B.C. (1995b) Peach yellows. In: Ogawa, J.M., Zehr, E.I., Bird, G.W., Ritchie, D.F., Uriu, K. and
Uyemoto, J.K. (eds) Compendium of Stone Fruit Diseases. APS Press, St. Paul, Minnesota, p. 57.
Kirkpatrick, B.C., Uyemoto, J.K. and Purcell, A.H. (1995) X-disease. In: Ogawa, J.M., Zehr, E.I., Bird, G.W.,
Ritchie, D.F., Uriu, K. and Uyemoto, J.K. (eds) Compendium of Stone Fruit Diseases. APS Press, St. Paul,
Minnesota, pp. 57–59.
Kirkpatrick, H.C., Lowe, S.K. and Nyland, G. (1975) Peach rosette: the morphology of an associated myco-
plasmalike organism and the chemotherapy of the disease. Phytopathology 65, 864–870.
Kison, H. and Seemüller, E. (2001) Differences in strain virulence of the European stone fruit yellows phyto-
plasma and susceptibility of stone fruit trees on various rootstocks to this pathogen. Journal of Phytopa-
thology 149, 533–541.
Kison, H., Kirkpatrick, B.C. and Seemüller, E. (1997) Genetic comparison of the peach yellow leaf roll agent
with European fruit tree phytoplasmas of the apple proliferation group. Plant Pathology 46, 538–544.
Klos, E.J. (1976) Red suture. In: Virus Diseases and Noninfectious Disorders of Stone Fruits in North America.
USDA Agriculture Handbook No. 437. US Department of Agriculture–Agricultural Research Service,
Washington, DC, pp. 133–134.
Larsen, H.J. and Waterworth, H.E. (1995) Peach red suture. In: Ogawa, J.M., Zehr, E.I., Bird, G.W., Ritchie,
D.F., Uriu, K. and Uyemoto, J.K. (eds) Compendium of Stone Fruit Diseases. APS Press, St. Paul, Min-
nesota, pp. 55–56.
Lee, I.M., Davis, R.E., Gundersen, E. and Rindal, D.E. (2000) Phytoplasmas: phytopathogenic Mollicutes. An-
nual Review of Microbiology 54, 221–255.
Lorenz, K.H., Dosba, F., Poggi-Pollini, C., Llacer, G. and Seemüller, E. (1994) Phytoplasma diseases on Prunus
species in Europe are caused by genetically similar organisms. Zeitschrift für Pflanzenkrankheiten und
Pflanzenschutz 101, 567–575.
Mizell, R., Andersen, P., Tipping, C. and Brodbeck, B. (2003) Xylella fastidiosa diseases and their leafhopper
vectors. University of Florida Cooperative Extension Service, ENY-683. http://edis.ifas.ufl.edu/BODY_
IN174 (accessed January 2008).
Montano, H.G., Davis, R.E., Dally, E.L., Hogenhout, S., Pimental, J.P. and Brioso Paulo, S.T. (2001) Candidatus
Phytoplasma brasiliense, a new phytoplasma taxon associated with hibiscus witches’ broom disease.
International Journal of Systematic and Evolutionary Microbiology 51, 1109–1118.
Moore, L.W. (1976) Latent infections and seasonal variability of crown gall development in seedlings of three
Prunus species. Phytopathology 66, 1097–1101.
Nunes, L., Rosato, Y., Muto, N., Yanai, G., da Silva, V., Leite, D., Goncalves, E., de Souza, A., Coletta-Filho, H.,
Machado, M., Lopes, S. and Costa, R. (2003) Microarray analyses of Xylella fastidiosa provide evidence of
coordinated transcription control of laterally transferred elements. Genome Research 13, 570–578.
Nyland, G. and Schlocker, A. (1951) Yellow leaf roll of peach. Plant Disease Reporter 35, 33.
Okie, W.R. (1998) Handbook of Peach and Nectarine Varieties. USDA Agriculture Handbook No. 714. US
Government Printing Office, Washington, DC.
Peterson, D.H. and Dowler, W.M. (1965) Bacterial canker of stone fruits in the southeastern states. Plant Dis-
ease Reporter 49, 701–702.
Pine, T.S. and Gilmer, R.M. (1976) Peach yellows. In: Virus Diseases and Noninfectious Disorders of Stone
Fruits in North America. USDA Agriculture Handbook No. 437. US Department of Agriculture–Agricul-
tural Research Service, Washington, DC, pp. 91–95.
Poggi-Pollini, C., Bissani, R. and Giunchedi, L. (2001) Occurrence of European stone fruit yellows phyto-
plasma (ESFYP) infection in peach orchards in Northern-Central Italy. Journal of Phytopathology 149,
725–730.
Prunier, J.P., Luisetti, J. and Gardan, L. (1970) Etudes sur les bacterioses des arbres fruitiers. II. Caracterisation
d’un Pseudomonas non fluorescent, agent d’une bacteriose nouvelle chez le pecher. Annales de Phyto-
pathologie 2, 168–197.
Purcell, A.H., Nyland, G., Raju, B.C. and Heringer, M.R. (1981) Peach yellow leaf roll epidemic in northern
California: effects of peach cultivar, tree age, and proximity to pear orchards. Plant Disease 65, 365–368.
Ritchie, D.F. (1993) Time of peach fruit infection by Xanthomonas campestris pv. pruni. Phytopathology
83, 1376.
434 D.F. Ritchie et al.

Ritchie, D.F. (1999) Sprays for control of bacterial spot of peach cultivars having different levels of disease
susceptibility, 1998. Fungicide & Nematicide Tests 54, 63–64.
Ritchie, D.F. and Clayton, C.N. (1981) Peach tree short life: a complex of interacting factors. Plant Disease 65,
462–469.
Rodrigues, J., Silva-Stenico, M., Gomes, J., Lopes, J. and Tsai, S. (2003) Detection and diversity assessment of
Xylella fastidiosa in field-collected plant and insect samples by using 16S rRNA and gyrB sequences.
Applied and Environmental Microbiology 69, 4249–4255.
Rosenberger, D.A. and Jones, A.L. (1977) Spread of X-disease in Michigan peach orchards. Plant Disease
Reporter 61, 830–834.
Scott, S.W. and Zimmerman, M.T. (2001) Peach rosette, little peach, and red suture are diseases induced by a
phytoplasma closely related to western X-disease. Acta Horticulturae 550, 351–354.
Seemüller, E., Stolz, H. and Kison, H. (1998) Persistence of the European stone fruit yellows phytoplasma in
aerial parts of Prunus taxa during the dormant season. Journal of Phytopathology 146, 407–410.
Shepard, D.P. and Zehr, E.I. (1994) Epiphytic persistence of Xanthomonas campestris pv. pruni on peach and
plum. Plant Disease 78, 627–629.
Smith, E.F. (1903) Observations on a hitherto unreported bacterial disease, the cause of which enters the plant
through ordinary stomata. Science (USA) 17, 456–457.
Vauterin, L., Hoste, B., Kesters, K. and Swings, J. (1995) Reclassification of Xanthomonas. International Jour-
nal of Systematic Bacteriology 45, 472–489.
Vigouroux, A. (1999) Bacterial canker of peach: effect of tree winter water content on the spread of infection
through frost-related water soaking in stems. Journal of Phytopathology 147, 553–559.
Wells, J., Raju, J., Thompson, J. and Lowe, S. (1981) Etiology of phony peach and plum leaf scald diseases.
Phytopathology 71, 1156–1161.
Wells, J., Raju, B. and Nyland, G. (1983) Isolation, culture and pathogenicity of the bacterium causing phony
disease of peach. Phytopathology 73, 859–862.
Werner, D.J. and Ritchie, D.F. (1986) Susceptibility of peaches and nectarines, plant introductions, and other
Prunus species to bacterial spot. HortScience 21, 127–130.
Young, J.M. (1988) Pseudomonas syringae pv. persicae from nectarine, peach, and Japanese plum in New
Zealand. OEPP/EPPO Bulletin 18, 141–151.
Young, J.M. and Triggs, C.M. (1994) Evaluation of determinative tests for pathovars of Pseudomonas syringae
van Hall 1902. Journal of Applied Bacteriology 77, 195–207.
Zehr, E.I. and Shepard, D.P. (1996) Bacterial spot of peach as influenced by water congestion, leaf wetness
duration, and temperature. Plant Disease 80, 339–341.
 17 Viruses and Viroids of Peach Trees

M. Cambra,1 R. Flores,2 V. Pallás,2 P. Gentit3 and T. Candresse4

1InstitutoValenciano de Investigaciones Agrarias (IVIA), Department of Plant
Protection and Biotechnology, Valencia, Spain
2Instituto de Biología Molecular y Celular de Plantas (IBMCP), Universidad Politécnica

de Valencia–CSIC, Valencia, Spain

3Centre Technique Interprofessionnel des Fruits et Lègumes de Lanxade (CTIFL),

La Force, France
4Institut National de la Recherche Agronomique, UMR GDPP–IBVM Equipe de

Virologie, Villenave d’Ornon, France

17.1 Introduction 436

17.2 Sharka or Plum Pox 437
Economic importance and geographic distribution 437
Symptoms 437
Causal agents and diagnosis 440
Epidemiology 442
Control 442
17.3 Ilarviruses 443
Economic importance and geographic distribution 443
Symptoms 443
Causal agents and diagnosis 446
Epidemiology 447
Control 447
17.4 Tomato Ringspot Virus 447
Economic importance and geographic distribution 447
Symptoms 448
Causal agent and diagnosis 448
Epidemiology 449
Control 449
17.5 Strawberry Latent Ringspot Virus 449
Economic importance and geographic distribution 449
Symptoms 449
Causal agent and diagnosis 449
Epidemiology 450
Control 450
17.6 Peach Rosette Mosaic Virus 451
Economic importance and geographic distribution 451
Symptoms 451
© CAB International 2008. The Peach: Botany, Production and Uses
(eds D.R. Layne and D. Bassi) 435
436 M. Cambra et al.

Causal agent and diagnosis 451

Epidemiology 451
Control 451
17.7 Peach Latent Mosaic 451
Economic importance and geographic distribution 451
Symptoms 451
Causal agent and diagnosis 452
Epidemiology 454
Control 454
17.8 Peach Dapple 454
Economic importance and geographic distribution 454
Symptoms 454
Causal agent and diagnosis 455
Epidemiology 455
Control 455
17.9 Peach Mosaic 455
Economic importance and geographic distribution 455
Symptoms 455
Causal agent and diagnosis 455
Epidemiology 456
Control 456
17.10 Diseases of Unknown Aetiology and Experimental Infections in Peach 456
Apricot latent virus, Peach sooty ringspot virus and Peach asteroid spot virus 456
Apple chlorotic leaf spot virus 456
Nepoviruses 457
17.11 Concluding Remarks and Perspectives 457

17.1 Introduction over long distances is the main cause of the

Viruses and viroids of peach trees incite over
20 different diseases, including some that
result from mixed infections of different
viruses. Agents belonging to these pathogen
groups most likely are also involved in other
diseases of uncertain aetiology. Some of the
diseases caused by these graft-transmissible
pathogens are economically important, partic-
ularly when they affect fruit quality or induce
severe tree decline. On the other hand, some
viruses and viroids that produce conspicuous
symptoms on specific peach cultivars or other
Prunus spp. do not cause phenotypic altera-
tions on other peach cultivars or hosts.
Most peach viruses and viroids do not
spread naturally, although some viruses may
be spread by aphid and nematode vectors or
through pollen. The propagation of infected
sources of plant material and their exchange
distribution of viruses and viroids in the
peach industry worldwide. The use of symp-
tomless infected trees for grafting purposes
considerably contributes to the spread of
viruses and viroids.
Reliable techniques based on serological
and molecular methods are available today
for the detection of most peach viruses and
viroids. These are gradually and successfully
replacing the conventional biological meth-
ods (indexing) and are contributing directly
to the control of the peach diseases based on
the exclusive use of virus- and viroid-free
propagation materials.
This chapter on diseases caused by
viruses and viroids includes 11 characterized
agents that are responsible for the main viral
problems in peach. The chapter also contains
comments on diseases of unknown aetiology
and on experimental infections of peach.
 Viruses and Viroids
17.2 Sharka or Plum Pox
Economic importance and
geographic distribution
Sharka (pox in Slavic), or plum pox disease, is
considered one of the most devastating dis-
eases of stone fruit in terms of agronomic
impact and economic importance (Dunez and
Sutic, 1988; Németh, 1994). The disease is
very detrimental in apricot, peach and plum
trees because it reduces fruit quality and can
result in premature drop of fruit. The estimated
costs associated with sharka disease man-
agement in the last 30 years exceeded 10,000
million worldwide (Cambra et al., 2006). It is
caused by Plum pox virus (PPV).
The PPV epidemic originated in Eastern
Europe: the disease was described for the first
time in about 1917 on plums and in 1933 on
apricots, both growing in Bulgaria (Atanasoff,
1932, 1935). Since then, the virus has progres-
sively spread to a large part of the European
continent, around the Mediterranean basin
and to the near and Middle East (Roy and
Smith, 1994). It has been found in South and
North America (Argentina, Chile, USA,
Canada) (Roy and Smith, 1994; Levy et al.,
2000; Thompson et al., 2001; Dal Zotto et al.,
Fig. 17.1. Discoloration symptoms of Plum pox virus type M on flowers of peach cultivar ‘Baby Gold
6’. (Courtesy of J.C. Desvignes, La Force, France.)
437
2006) and in Asia (Kazakhstan and China)
(Spiegel et al., 2004; Navrátil et al., 2005). An
update of PPV and sharka disease has been
published (García and Cambra, 2007).
Symptoms
In peach, symptoms may appear on petals,
leaves and fruits. Discoloration of petals
(colour breaking) can occur on the flowers of
some peach varieties (Figs 17.1 and 17.2/
Plates 167 and 168). Symptoms on leaves
include mild light green discoloration, chlo-
rotic spots, bands or rings, vein clearing or
yellowing, and even leaf deformation (Figs
17.3 and 17.4, Plates 169 and 170), and are
particularly evident in spring.
Infected fruit show chlorotic spots or
underpigmented yellow rings or line patterns
(Figs 17.5, 17.6 and 17.7/Plates 171, 172 and
173). Fruit may become deformed or irregular
in shape and develop either small brown or
necrotic areas. Diseased fruit may show inter-
nal browning of the flesh and have a reduced
quality. In some cases the diseased fruit drop
prematurely from the tree. The development
of fruit in early varieties (May to June) is coin-
cidental with the highest concentration of
438 M. Cambra et al.

Fig. 17.2. Colour break symptoms of Plum pox virus type M on flower petals of peach cultivar
‘Gladys’. (Courtesy of M.A. Cambra, Zaragoza, Spain.)

Fig. 17.3. Leaf symptoms of Plum pox

virus on ‘GF 305’ peach seedlings used
as indicator plant.

PPV in the tree in Mediterranean climates.
Viral titre decreases later in summer (July to
September), when the fruit of late varieties
are developing. Consequently, there is a broad
general trend for early-ripening varieties to
be much more sensitive and to show more
severe symptoms on fruit than later-maturing
varieties.
Plum pox disease symptoms in peach
vary considerably with the cultivar, age of the
Viruses and Viroids 439

Fig. 17.4. Leaf symptoms of Plum pox

virus type M on peach cultivar ‘Royal
Gem’.

Fig. 17.5. Fruit symptoms (under-

pigmented yellow rings) caused by
Plum pox virus in peach cultivar
‘Springcrest’.
440 M. Cambra et al.

Fig. 17.6. Chlorotic rings and line

pattern caused by Plum pox virus in
fruit of nectarine cultivar ‘Arm King’.

Fig. 17.7. Chlorotic spots and underpigmented yellow rings caused by Plum pox virus in fruits of
peach cultivar ‘Catherine’. (Courtesy of M.A. Cambra, Zaragoza, Spain.)

plant, climatic conditions (temperature) and
PPV type.
Causal agents and diagnosis
PPV is a member of the viral genus Potyvirus
in the family Potyviridae (López-Moya and
García, 1999). PPV particles are flexuous rods
about 700 nm in length and 11 nm in width.
They are composed of a single-stranded RNA
molecule of about 10,000 nucleotides in length
coated by up to 2000 subunits of a single coat
protein. The virus’s expression strategy, typi-
cal of potyviruses, includes translation of a
unique, long, open reading frame into a large
polyprotein that is processed after translation
by three viral proteinases to yield nine or ten
mature viral proteins. In recent years, know-
ledge of the molecular biology of potyviruses
 Viruses and Viroids
in general, and of PPV in particular, has
increased significantly (Riechmann et al., 1992;
Shukla et al., 1994; Revers et al., 1999).
The numerous PPV isolates differ in both
biological and epidemiological properties
such as aggressiveness, aphid transmissibil-
ity and symptomatology. Two main groups or
types, Dideron (D) and Marcus (M), were dis-
tinguished serologically by Kerlan and Dunez
(1976). With the advent of molecular tech-
niques, complete or partial genomic sequences
have been obtained for a large number of PPV
isolates. Analysis of these sequences has con-
firmed the existence of major groups of PPV
isolates which correspond to the initial D and
M groups. Two additional minor groups, repre-
sented by the geographically limited El Amar
isolate from Egypt (PPV-EA; Wetzel et al., 1991a)
and by the cherry-adapted isolates (PPV-C;
Nemchinov and Hadidi, 1996; Nemchinov
et al., 1996), have been identified. An unusual
PPV isolate reported from Canada (PPV-W;
James et al., 2003) probably represents a dis-
tinct fifth PPV group. In addition, the wide
distribution of natural recombinants between
the D and M types of PPV has been recently
demonstrated, leading to the suggestion to
consider these recombinant isolates as consti-
tuting a new group (PPV-Rec; Glasa et al.,
2004). Consequently, PPV isolates can currently
be clustered into six clearly different groups.
Given the variability of PPV, all techniques
other than sequencing or some PCR-based
assays (see below) may provide erroneous
answers on the typing of a small percentage
of isolates (Candresse et al., 1998a). However,
the discrimination between D and M groups
of PPV isolates is possible for most isolates
using a variety of techniques. These include:
(i) different serological pattern or reaction
with D- or M-specific monoclonal antibodies
(Cambra et al., 1994; Boscia et al., 1997); (ii)
electrophoretic mobility of the viral coat pro-
tein as assessed by Western blot (Bousalem
et al., 1994; Pasquini and Barba, 1994); (iii)
sequence analysis of PCR fragments corre-
sponding to the C-terminal region of the PPV
coat protein gene or RsaI restriction fragment
length polymorphism analysis of this region
(Wetzel et al., 1991b; Bousalem et al., 1994;
Candresse et al., 1994); and (iv) different vari-
ants of PCR, heminested-PCR, nested-PCR
441
and cooperational PCR (Co-PCR), using spe-
cific primers (Candresse et al., 1994; Olmos
et al., 1997, 1999, 2002, 2003) including colori-
metric detection of the amplicons with D- or
M-specific probes. In addition, real-time PCR
(RT-PCR) assays using either SYBR Green or
TaqMan chemistries have recently been devel-
oped that allow discrimination between D
and M PPV types (Varga and James, 2005;
Capote et al., 2006).
Detection of all PPV isolates can be
achieved by using either monoclonal antibody
5B-IVIA (Cambra et al., 1994) or polyclonal
antibodies in a double-antibody sandwich
indirect (DASI; also named triple-antibody
sandwich, TAS) or a double-antibody sand-
wich (DAS) ELISA assay, respectively. Specific
detection of PPV-D isolates (Cambra et al.,
1994), PPV-M (Boscia et al., 1997), PPV-C
(Myrta et al., 2000) and PPV-EA (Myrta et al.,
1998) is possible by using available ELISA kits.
Molecular hybridization techniques (Varveri
et al., 1988) and different PCR-based assays
have been developed for the detection
(Korschineck et al., 1991; Wetzel et al., 1991b,
1992; Candresse et al., 1994, 1995; Levy et al.,
1994; Olmos et al., 1996) or for the simultane-
ous detection and typing of PPV isolates
(Olmos et al., 1997; Candresse et al., 1998a). Dif-
ferent systems of viral target preparation prior
to PCR have been developed based on immu-
nocapture (Wetzel et al., 1992) or, without the
need for extract preparation, on print or squash
capture (Olmos et al., 1996). The use of immo-
bilized targets on paper (Cambra et al., 1997)
allowed the detection of PPV in single aphids
(Olmos et al., 1997) by squash capture-PCR.
Nested-PCR in a single closed tube (Olmos
et al., 1999) has been applied for the sensitive
detection of PPV targets in plant material and
in single aphids. A Co-PCR system using a
universal probe for hybridization (Olmos et al.,
2002) has been described, affording sensitivity
similar to that of nested-PCR. Real-time RT-
PCR assays have been developed to detect and
quantify PPV targets in plant material and
individual aphids (Schneider et al., 2004; Olmos
et al., 2005; Varga and James, 2005; Capote et al.,
2006) with sensitivity higher than obtained by
the previously described methods and, in some
cases, without the need for RNA purification
(Olmos et al., 2005). Generally, serological and
442 M. Cambra et al.
molecular characterization of PPV isolates cor-
relate very well, although serological typing
may prove in error for a few unusual isolates
(Candresse et al., 1998a).
A European protocol for the detection
and characterization of PPV has been devel-
oped. The recommended methods include
indexing (graft inoculation of ‘GF 305’ peach
seedlings or Prunus tomentosa), serological
and molecular tests (EPPO, 2004).
Epidemiology
The movement of infected plant propagation
material is considered to be the most impor-
tant means of long-distance spread of PPV. In
addition, the virus is non-persistently trans-
mitted by a number of aphid species existing
in each region (Kunze and Krczal, 1971;
Labonne et al., 1995). Non-aphid transmissi-
ble isolates have also been described (Maiss
et al., 1989; López-Moya et al., 1995).
Under natural conditions, PPV easily
infects fruit tree species of the genus Prunus
used as commercial varieties or rootstocks: apri-
cot (Prunus armeniaca), European plum (Prunus
domestica), Japanese plum (Prunus salicina),
peach (Prunus persica), Myrobalan flowering
plum (Prunus cerasifera) and Mariana plum (Pru-
nus mariana). Sour (Prunus cerasus) and sweet
(Prunus avium) cherries and almond (Prunus dul-
cis) trees may be infected occasionally. The virus
also infects most wild or ornamental Prunus
species such as Western sand cherry (Prunus
besseyi), purple-leaved sand cherry (Prunus cis-
tena), damson plum (Prunus insititia), Nanking
cherry (P. tomentosa), flowering almond (Prunus
triloba) and blackthorn (Prunus spinosa).
The virus can be mechanically transmit-
ted to numerous Prunus species, service tree
(Sorbus domestica) and several herbaceous
plants. Nicotiana benthamiana, Nicotiana gluti-
nosa, Nicotiana clevelandii, garden pea (Pisum
sativum) and Chenopodium foetidum are fre-
quently used as experimental host plants.
The PPV isolates belonging to the D or M
groups appear to show different epidemiologi-
cal behaviours, at least under western Euro-
pean conditions. Generally, the M isolates
tend to be spread more readily by aphids to
peach trees than the D isolates. M isolates
cause faster epidemics and more severe symp-
toms in peach flowers, leaves and fruit than D
isolates. The D isolates are able to spread nat-
urally in apricot and plum orchards but spread
much more rarely from these hosts to peach
trees. These are, however, broad discrimina-
tions that may not apply to some individual
isolates or under different epidemiological
conditions.
Control
As is also the case for potyviruses infecting
other host crops, fruit trees cannot be effi-
ciently protected from PPV infection by the
use of aphicides. Potyviruses are transmitted
in a non-persistent manner; PPV can be inoc-
ulated by a viruliferous aphid during very
short probes so that aphicides have no direct
effect on the process. Aphid vector popula-
tions do not, for the most part, reside or
develop on the fruit tree crops. Thus applica-
tions of insecticides to a growing orchard
have only limited effects.
In this context, control measures against
PPV are of two kinds: (i) efforts at prophy-
laxis designed to reduce or eliminate the viral
load in the environment (quarantine mea-
sures, eradication programmes, use of certi-
fied virus-tested planting material, etc.); and
(ii) efforts at breeding for resistance. It should
also be mentioned that in countries with
endemic infection a third strategy, relying on
the deployment of varieties with a reduced
susceptibility (in particular with reduced
symptoms on fruit), is widely used, despite
the fact that it provides no real disease control
(quite the contrary in fact) but allows instead
for continued production under endemic
infection conditions.
In countries where infection levels are
still low, strategies based on a strict control of
the virus and on prophylaxis are generally used.
PPV was recognized early as a major pathogen
on stone fruit crops and was therefore included
on quarantine lists. In Europe, for example, it
was recognized as a quarantine pathogen in
1973 (Dosba, 1992). Currently, the European
quarantine status of PPV is referred in the
EPPO (European and Mediterranean Plant
Protection Organization) Quarantine List A2
 Viruses and Viroids
and EU Annex designation IIAII. Similarly, in
many countries plum pox is subject to strict
control measures, including eradication
schemes. In practice, there are, however, very
few cases in which PPV was actually eradi-
cated from a country (only Switzerland and
the Netherlands). To be successful, these
strategies must be undertaken early after the
introduction of the disease in a country or
region and must be very vigorously enforced.
Generally, eradication schemes are labour-in-
tensive and based on either visual inspection
of symptoms or ELISA testing. Such approaches
also necessitate a collective action which is
often difficult to obtain from fruit growers
unless effective compensatory measures are
implemented. Even if not completely success-
ful, these eradication strategies have frequently
allowed limitation of the rate of spread of the
disease and of its impact to a low and man-
ageable percentage of infection. Two different
programmes are being conducted in the USA
and Canada, since the year 2000, with the aim
of eradicating PPV. The success of these pro-
grammes will be assessed in the near future.
A complement to quarantine and eradi-
cation measures is the wide use of certified,
virus-tested or virus-free planting material.
Sanitation techniques, detection and sam-
pling techniques, information on the protec-
tion of nurseries, etc. are available today,
enabling many countries to develop efficient
certification programmes.
In parallel with these control measures,
efforts aimed at the development of PPV-
resistant Prunus varieties have been undertaken
in many countries. These programmes have
explored both classical breeding approaches
(screening of germplasm to identify resistance
sources) and biotechnological approaches. In
this respect, peach has so far proved a more
complicated target than apricot and plum.
Extensive screening of germplasm has failed to
identify sources of resistance within the peach
species, so that current efforts are aimed at the
exploitation of resistance identified in the
related Chinese wild peach (Prunus davidiana)
by introgression of the trait through interspe-
cific hybridization (Pascal et al., 1997; Escalettes
et al., 1998; Bassi, 2006). By contrast, in apricot
and plum, resistance sources have been identi-
fied within the target species, so breeding
443
efforts are clearly more advanced (Dosba et al.,
1989; 1991; Dicenta and Audergon, 1998; Karayi-
annis et al., 1999; Vilanova et al., 2003).
Genetic transformation to create PPV-
resistant transgenic plants, through the use of
coat protein-mediated protection, has been
successful in both apricot and plum (Laimer
da Camara Machado et al., 1992; Scorza et al.,
1994, 2001; Ravelonandro et al., 1997). Effi-
cient genetic constructions are therefore avail-
able, some of which have been successfully
validated in field tests (Ravelonandro et al.,
2000, 2002; Hily et al., 2004). In addition, novel
approaches to immunomodulate host–PPV
pathogen interactions by expression of anti-
body genes in plants have emerged (Esteban
et al., 2003). All these possibilities have, how-
ever, not been transferred to peach because
this species has so far largely remained refrac-
tory to genetic transformation. If a system for
transformation/regeneration of peach was to
become more broadly available, biotechnol-
ogy could offer a new and innovative strategy
to fight the sharka disease.
17.3 Ilarviruses
Economic importance and geographic
distribution
Ilarviruses that infect peach crops are Prunus
necrotic ringspot virus (PNRSV), Prune dwarf
virus (PDV), Apple mosaic virus (ApMV) and
American plum line pattern virus (APLPV).
PNRSV, PDV and ApMV are found world-
wide, infecting a wide range of Prunus species.
Until very recently APLPV was only reported
in North America in Japanese plum (P. salicina),
peach (P. persica) and flowering cherry (Prunus
serrulata) (Fulton, 1982). APLPV is included in
the EPPO Quarantine List A1 of plant patho-
gens (Smith et al., 1992) and was recently
reported in Palestine, Italy, Albania and Tuni-
sia (Myrta et al., 2002; Alayasa et al., 2003).
Symptoms
Symptoms associated with PNRSV infec-
tion in peaches include foliar chlorotic rings
444 M. Cambra et al.
(Fig. 17.8/Plate 174), necrotic spots (Fig. 17.9/
Plate 175) or deformation, and bark necrosis,
pitting, splitting or girding. Infected trees
tend to have smaller trunk diameter than
uninfected ones and the yield of infected trees
is usually reduced (Saunier, 1972; Pusey and
Yadava, 1991; Scott et al., 2001). In addition,
fruit maturity is affected depending on the
rootstock. Besides necrotic leaf spot, colour
break on flower petals has been observed on
the mature trees of some peach varieties (e.g.
‘Rio Oso Gem’). Some cultivars do not show
overt symptoms.
Symptoms caused by PDV in peach are
less obvious and the virus produces only mild
stunting in most peach cultivars and essen-
tially no leaf symptoms. In some peach culti-
vars, however, PDV infection produces a
dense canopy due to shortening of shoot
internodes (Fig. 17.10/Plate 176) and typical
ring spots.
Mixed infection with PNRSV plus PDV
results in a disease known in Australia as
peach rosette and decline (Smith and Challen,
1977) and in California as peach stunt disease
(Asai and Uyemoto, 1991). Affected trees
show premature defoliation at the end of the
first growing season following inoculation.
Infected trees display bark splitting and
increased water sprout (sucker) production,
and yield is reduced by up to 60%. Reduc-
tions in shoot growth, trunk diameter and the
number of leaves produced were observed
and a delay in bloom date has also been noted
(Uyemoto et al., 1992).
ApMV is not very frequent in peach. The
virus induces a bright yellow mosaic and/or
spots (Fig. 17.11/Plate 177), so nurserymen
Fig. 17.8. Leaf symptoms (chlorotic
rings and deformation) caused by
Prunus necrotic ringspot virus in peach.
 Viruses and Viroids 445

Fig. 17.9. Necrotic spots caused by Prunus necrotic ringspot virus in peach leaves.

Fig. 17.10. Dense canopy due to

shortening of shoot internodes caused
by Prune dwarf virus in peach trees.
446 M. Cambra et al.
generally avoid propagating the infected
trees. Finally, in spring APLPV causes a thin
reticulation or yellowing of the little veins on
the leaves of the infected trees.
Causal agents and diagnosis
PNRSV, PDV, ApMV and APLPV belong to
the genus Ilarvirus, whose members are char-
acterized by a tripartite genome and quasi-
isometric particles (van Regenmortel et al.,
2000). Ilarviruses affecting stone fruit are closely
related phylogenetically (Sánchez-Navarro and
Pallás, 1997) and show at least 65% sequence
similarity at the coat protein level.
Historically, serological procedures have
extensively used assays for the detection of
these four viruses (e.g. Halk et al., 1984; Mink
Fig. 17.11. Leaf symptoms (bright
yellow mosaic and spots) caused by
Apple mosaic virus in peach.
and Aichele, 1984; Scott et al., 1989; Uyemoto
et al., 1989). However, erratic ELISA results
have been reported for shoot-leaf samples
collected in early May when day tempera-
tures exceeded 38°C over a 12-day period
(Uyemoto et al., 1989). In addition, Scott et al.
(1992) have described that when using ELISA,
PNRSV was detectable in peach tissues only
until stem elongation ceased.
Molecular hybridization using radiola-
belled probes allows the discrimination
between healthy and virus-infected material
throughout the growing season (Scott et al.,
1992) and is suitable for detecting PNRSV
serotypes that react poorly in ELISA (Crosslin
et al., 1992). Nevertheless, the use of radiola-
belling renders this approach unsuitable for
routine diagnosis. Non-isotopic RNA probes
have been used to monitor PNRSV infection
after in vitro micrografting (Heuss-La Rosa
 Viruses and Viroids
et al., 1995) and to detect these viruses in field
conditions (Pallás et al., 1998; Sánchez-Navarro
et al., 1998; Saade et al., 2000).
In addition, several variations of the PCR
technique have been applied for the detection
of these viruses (Parakh et al., 1995; Rowhani
et al., 1995; Spiegel et al., 1996; Sánchez-Navarro
et al., 1998; Helguera et al., 2001; Scott and
Zimmerman, 2001).
Both molecular hybridization-based meth-
ods and PCR-based methods detect these
viruses even when they are only present in low
concentration in the plant samples being tested.
As a step towards the development of more
rapid methods of detection, attempts have been
made to simultaneously detect at least two of
the ilarviruses. This has been easily achieved
by using mixtures of several non-isotopic
probes in molecular hybridization, for instance
for the detection of PNRSV, PDV and ApMV
(Saade et al., 2000). Detection of two ilarvi-
ruses or one ilarvirus together with other
important stone fruit virus, e.g. PNRSV and
PPV (Kölber et al., 1998), in a single multiplex
RT-PCR with the same sensitivity as that
observed during individual RT-PCR analyses
has been reported. Additionally, a PCR-ELISA
procedure has been described for the simulta-
neous detection of ApMV and PNRSV (Can-
dresse et al., 1998b) but, in this case, an extra
probe-capture step was required to differenti-
ate between the two viruses. PNRSV, PDV
and ApMV were simultaneously detected in
a single assay by RT-PCR (Saade et al., 2000).
More recently, a multiplex RT-PCR method
has been developed to detect eight stone fruit
viruses, including the four ilarviruses, in a
single assay (Aparicio et al., 2003).
Epidemiology
PNRSV and PDV are pollen- and seed-borne
(Cole et al., 1982; Hamilton et al., 1984; Kelley
and Cameron, 1986; Aparicio et al., 1999), which
contributes to their rapid spread in stone fruit
trees (Mink, 1992; Uyemoto et al., 1992).
Infected pollen has been shown to play a key
role in both seed and plant-to-plant transmis-
sion of these viruses. Recent evidence suggests,
however, that certain biological agents such
447
as thrips or other flower-dwelling arthropods
may facilitate plant-to-plant transmission
through pollen (Greber et al., 1992; Mink,
1992; Johansen et al., 1994). PNRSV has been
detected in seedlings grown from seeds pro-
duced on healthy plants of several Prunus
species after hand pollination with pollen
from infected plants, strongly suggesting that
fertilization could also be a contaminating
event (reviewed by Mink, 1992). However,
contradictory results have been reported con-
cerning the location of PNRSV on or in pollen
grains from infected cherry trees. Cole et al.
(1982) showed that PNRSV antigens were
easily removed from intact pollen by wash-
ing, indicating that most if not all the virus
was on the pollen surface. Results obtained
by Kelley and Cameron (1986) and Digiaro
et al. (1992) suggested that PNRSV was located
both externally and internally. In situ hybrid-
ization experiments confirmed that the virus
is located within the pollen grains (Aparicio
et al., 1999).
Control
The most important method for controlling
the diseases caused by ilarviruses affecting
peach crops is the use of virus-tested certified
plants. Effective management requires prompt
removal and destruction of diseased plants.
Nurseries producing basic propagation
material need to be separated from commer-
cial orchards by an appropriate distance to
prevent or limit contaminating pollen flow.
Thermotherapy (24–32 days at 38°C) and api-
cal meristem culture have been used to elimi-
nate these viruses. Cross-protection with mild
PNRSV isolates has been used occasionally.
17.4 Tomato Ringspot Virus
Economic importance and geographic
distribution
First described in North America, where it is
endemic, Tomato ringspot virus (ToRSV) has been
associated with various diseases in different
Prunus hosts. On peach, two diseases have
448 M. Cambra et al.
been described: (i) the peach yellow bud mosaic
disease, which is restricted to North America
and is associated with the distribution of the
nematode vector (Schlocker and Traylor,
1976); and (ii) the Prunus stem pitting disease,
which is present in North America and
Europe and is not associated with the vector
distribution (Németh, 1986). Even though the
virus is common on a wide range of orna-
mental and cultivated plants throughout the
world, both diseases are rarely associated
with severe losses in the peach industry.
However, substantial losses due to infection
with ToRSV still occur in orchards in Pennsyl-
vania, USA.
Symptoms
First described in 1936, the peach yellow bud
mosaic disease shows two phases. During the
first stage of infection, chlorotic spots and
rings or oak-leaf mottles are observed on
leaves (mosaic phase). This is followed by the
second phase, characterized by a severe
reduction of leaf growth on buds, giving a
rosette appearance (yellow bud phase) (Fig.
17.12/Plate 178). After a few years, infected
trees display a denuded appearance and fruit
production is reduced (Schlocker and Traylor,
1976). The most characteristic symptom for
the Prunus stem pitting disease is an abnor-
mal thickening of the bark, which shows a
spongy aspect associated with pits and
grooves in the wood (Barrat et al., 1968).
Causal agent and diagnosis
ToRSV is a member of the subgroup C of the
genus Nepovirus (Mayo and Robinson, 1996)
and has isometric particles about 28 nm in
Fig. 17.12. Severe reduction of leaf
growth on buds, giving a rosette
appearance, caused by Tomato ringspot
virus in ‘GF 305’ peach seedlings.
 Viruses and Viroids
diameter, sedimenting as three components.
It has a bipartite, single-stranded RNA genome
characterized by an unusually large RNA-2
(7273 nucleotides) similar in size to the RNA-1
(8114 nucleotides) (Stace-Smith, 1984). For
diagnosis purposes, the ToRSV can be detected
after mechanical transmission to the herba-
ceous hosts Chenopodium quinoa or Chenopo-
dium amaranticolor or, with greater sensitivity,
by indexing on ‘GF 305’ peach seedlings fol-
lowing graft inoculation by chip budding
(Desvignes, 1999). Owing to possible confu-
sion with symptoms of other nepoviruses in
particular, serological diagnosis by ELISA is,
however, recommended for definitive identi-
fication of ToRSV (Powell, 1984). In addition,
PCR-based assays have been developed and
are also available (Griesbach, 1995), including
a colorimetric-PCR assay that allows quantifi-
cation of the virus in samples from woody
hosts (Rowhani et al., 1998).
Epidemiology
On peach, ToRSV is naturally transmitted by
the North American dagger nematodes, Xiphi-
nema americanum Cobb, Xiphinema californicum
Lamberti and Bleve-Zacheo and Xiphinema
rivesi Dalmasso (Stace-Smith, 1984). In the field,
this mode of transmission is associated with
typical tree to neighbouring tree spreading.
Because of its wide host range, which includes
common weeds such as dandelion (Taraxacum
officinale) and sheep sorrel (Rumex acetosella),
the virus can persist on a site for many years
after removal of infected peach trees (Powell
et al., 1984). Propagation of infected plants and
grafting are also efficient transmission and dis-
semination means of the virus.
Control
The best way to control ToRSV is to prevent
its introduction and therefore the implemen-
tation of quarantine measures and the use of
planting material from a virus-free certifica-
tion programme are highly recommended. In
addition, long crop rotations with non-host
plants associated with a nematicide treatment
of soil before planting is strongly recommended.
449
In infected plots, diseased trees must be
pulled out. ToRSV is associated with union
necrosis in apples; thus replacing apple
orchards with peach is inadvisable as severe
and rapidly developing outbreaks of Prunus
stem pitting may result.
17.5 Strawberry Latent Ringspot Virus
Economic importance and geographic
distribution
Except for Canada, where this virus was
described once but was not shown to spread
naturally, Strawberry latent ringspot virus
(SLRSV) has been mainly described under
different names on peach trees in Europe. It
was first reported in 1962 in France as ‘court
noué du pêcher’, then in Germany as ‘shoot
dwarfing’, in Italy as ‘Rosetta a foglie salici-
formi’ and in Hungary (Fortusini et al., 1983;
Carles, 1984; Németh, 1986). In southern
European countries it is quite widespread in
the traditional production areas, where the
virus can be naturally maintained in weeds or
in wild-growing plants.
Symptoms
SLRSV is associated with mild symptoms on
peach. During spring, particularly with cold
weather, a delay in bud breaking, flowering
and leaf growth is observed. These early
symptoms are usually conspicuous in the
field but later in the season they can be much
less clear. Leaves become small, rolled and
chlorotic and the internodes are shortened
(Fig. 17.13/Plate 179), giving a stunted aspect
of the trees. When the weather becomes
warmer, new developing shoots are less
affected and the symptoms tend to disappear.
In mixed infections with PDV or PNRSV,
symptoms are much more severe due to syn-
ergistic effects between the viruses.
Causal agent and diagnosis
SLRSV is a tentative member of the genus Nep-
ovirus, displaying the typical 30 nm isometric
450 M. Cambra et al.
particle morphology. The particles sediment
as three components and encapsidate a bipar-
tite, single-stranded RNA genome (Kreiah
et al., 1994). Contrary to the majority of nepo-
viruses, which have only a single type of
capsid subunit, SLRSV particles contain two
different subunits (Mayo and Robinson,
1996). This virus is a good immunogen and is
easily identified in serological tests such as
ELISA. Being systemically distributed through-
out the trees, it can also be detected by bio-
logical indexing on ‘GF 305’ peach seedlings
(Desvignes, 1999). A PCR-based detection
assay has also been developed (Bertolini et al.,
2001), although it has not been tested exten-
sively on stone fruit tree samples.
Epidemiology
Many plant species are natural hosts for SLRSV.
Cherry (P. avium), European plum (P. domestica),
currants (Ribes spp.) and raspberries (Rubus
Fig. 17.13. Internode shortening
caused by Strawberry latent ringspot
virus in peach trees.
spp.) are susceptible, and natural latent infec-
tions occur in a number of other woody plants
such as black locust (Robinia pseudoacacia) and
elders (Sambuscus spp.), herbaceous plants
and weeds (Murant, 1974). Thus, SLRSV has a
potentially large number of reservoir hosts
for a new orchard infection. In peach, its seed
transmission has not been demonstrated. In
strawberry, the type strain of the SLRSV is
transmitted by the nematodes Xiphinema diver-
sicaudatum and Xiphinema coxi (Murant, 1974).
On peach its transmission by nematodes has
been assumed (Desvignes, 1999). The virus can
also easily be transmitted by grafting.
Control
Soil disinfections with nematicides and
use of virus-tested material from a national
certification programme are good options to
prevent introduction of the virus in a new
orchard. In a contaminated orchard, often due
 Viruses and Viroids
to the reduction/disappearance of the symp-
toms during the warmer months, growers
tend to retain infected trees until they are no
longer profitable. In areas where field transmis-
sion occurs, infected trees should be removed,
and weed or shrub proliferation controlled.
17.6 Peach Rosette Mosaic Virus
Economic importance and geographic
distribution
The disease was first observed in some peach
orchards in North America (Michigan, USA and
Ontario, Canada) and was also later associated
with a disease on ‘Concord’ grape (Dias and
Cation, 1976). It is not yet described in Europe,
where it is considered a quarantine pathogen.
With this limited distribution, the economic
importance of this disease in peach is limited.
Symptoms
The foliation of infected trees is delayed.
Leaves are small, curled and with chlorotic
mottling along the main vein (mosaic symp-
toms). Later in the season, shortening of inter-
nodes gives a stunted aspect to infected trees
and induces rosetting symptoms.
Causal agent and diagnosis
The causal agent has been identified as Peach
rosette mosaic virus (PRMV), a member of the
subgroup C of the genus Nepovirus. It has iso-
metric particles of 28 nm in diameter sedi-
menting as three components and possesses a
bipartite, single-stranded RNA genome. Par-
ticles contain one single polypeptide subunit
(Mayo and Robinson, 1996). PRMV can be
detected using serological assays or by index-
ing on P. persica cv. Elberta (Klos, 1976).
Epidemiology
PRMV is transmissible by grafting. It is
described as soil-borne and transmission by
451
X. americanum has been demonstrated. The
virus may also be transmitted by a Criconemoides
species (Ramsdell and Myers, 1978). It natu-
rally infects some weeds such as dandelion
(T. officinale), which may constitute reservoirs
for the virus in the environment, and it can be
experimentally transmitted to several other
herbaceous hosts (Chenopodium spp., Nicoti-
ana spp.) (Dias, 1975).
Control
As for other nepoviruses, it is recommended
to use certified virus-free peach trees, to
remove and destroy diseased trees and to
fumigate infected orchard sites against nema-
todes before replanting.
17.7 Peach Latent Mosaic
Economic importance and geographic
distribution
Peach latent mosaic (PLM) disease is econom-
ically significant because it affects fruit qual-
ity, reduces tree lifespan and causes peach
trees to be more susceptible to other biotic
and abiotic stresses. PLM disease, which is
induced by Peach latent mosaic viroid (PLMVd),
is present in most of the peach-growing areas
of the world: the Mediterranean basin, North
and South America, China, Japan and Austra-
lia, being particularly prevalent in those areas
where varieties of American or Japanese ori-
gin predominate.
Symptoms
PLM disease was initially described in France
as a consequence of sanitary controls on
imported peach material. It only affects peach
and peach hybrids (Desvignes, 1976, 1986).
The term ‘latent’ in the name of this disease
refers to the fact that most natural infections
occur without inducing leaf symptoms, with
the first pathological alterations being observed
at least 2 years after planting. Early field
452 M. Cambra et al.
symptoms consist of delayed flowering, folia-
tion and ripening, accompanied by altera-
tions of variable severity in different organs.
Flowers present pink streaks on petals, and
fruit show deformations and discolorations
with cracked sutures (Fig. 17.14/Plate 180)
and flattened stones. Bud necrosis and stem
pitting can also be observed. Leaf symptoms
appear sporadically as chlorotic blotches
(peach blotch), yellow-creamy mosaics (peach
yellow mosaic) (Fig. 17.15/Plate 181) and, in
the most severe cases, white patterns that
may cover most or all of the leaf area (peach
calico) (Fig. 17.16/Plate 182). The diseases
with these names reported previously in the
USA and Japan (Németh, 1986) are most likely
manifestations of distinct PLMVd strains/
isolates. Generally, 5 years after planting, the
trees present a reduced foliar density, a typi-
cal open growth habit and an increased sus-
ceptibility to biotic and abiotic stress, which
finally causes their premature death. In the
greenhouse, PLMVd isolates can be divided
into severe strains and latent strains, depend-
ing on whether or not they induce leaf symp-
toms on seedlings of the peach indicator ‘GF
305’. Pre-inoculation with latent strains incite
in ‘GF 305’ a cross-protection effect against
Fig. 17.14. Fruit symptoms (deformations and discolorations with cracked sutures) caused by Peach
latent mosaic viroid in peach.
challenge inoculations with severe strains
(see below). This effect is the basis of a bioas-
say for diagnosis (Desvignes, 1976, 1986).
Causal agent and diagnosis
The search for the causal agent of the PLM
disease, initially assumed to have a viral aeti-
ology, was unsuccessful for some time. Later,
a viroid RNA was isolated from plants exhib-
iting the typical PLM symptoms; this RNA,
which was absent in healthy controls, could
also be identified in asymptomatic ‘GF 305’
plants infected by a mild isolate of the aetiologi-
cal agent (as revealed by the cross-protection
assay) (Flores and Llácer, 1988). The involve-
ment of a viroid was reinforced when the
same RNA was found in naturally infected
plants of 20 different peach varieties, but not
in the plants resulting from sanitation by ther-
motherapy. A definitive causal relationship
between PLM disease and the viroid RNA was
established when, following inoculation of ‘GF
305’ with a purified preparation of the latter,
the plants developed the typical symptoms
and the same RNA could be recovered, which
 Viruses and Viroids
henceforth was termed Peach latent mosaic
viroid (PLMVd) (Flores et al., 1990).
Cloning and sequencing of PLMVd
revealed a RNA genome of 337 nucleotides.
Comparison of the motifs and structures pres-
ent in this sequence with other previously
described viroid sequences enable PLMVd to
be grouped in the genus Pelamoviroid within
the family Avsunviroidae (Flores et al., 2000).
Under greenhouse conditions, PLMVd
can be detected by a cross-protection bioassay
in ‘GF 305’. Briefly, ‘GF 305’ peach seedlings
are graft-inoculated with material from the
plants to be tested or from reference isolates.
Most PLMVd natural isolates do not incite leaf
symptoms in ‘GF 305’, whereas severe isolates
induce a wide range of symptoms (see above)
and are therefore detected in this first phase of
the assay. When challenge-inoculated with a
severe isolate, ‘GF 305’ plants pre-inoculated
453
Fig. 17.15. Leaf symptoms (yellow-
creamy mosaic) caused by Peach latent
mosaic viroid in peach.
with a latent PLMVd isolate do not display the
characteristic symptoms, allowing the identifi-
cation of latent PLMVd isolates in this second
phase of the assay. This bioassay was very
useful for the diagnosis and control of PLM in
France even when the viroid aetiology of the
disease was unknown (Desvignes, 1976, 1986).
PLMVd can now be detected more quickly
with different molecular approaches: (i) PAGE
(Flores and Llácer, 1988; Flores et al., 1990); (ii)
dot-blot hybridization with radioactive and
non-radioactive probes (Flores et al., 1990;
Ambrós et al., 1995; Loreti et al., 1995); and (iii)
reverse transcription and PCR using specific
primers (Shamloul et al., 1995). These tests
have shown the presence of PLMVd in Japa-
nese and Italian peach varieties displaying
peach yellow mosaic and peach calico, respec-
tively. Interestingly, in the latter case the
symptoms have been demonstrated to result
454 M. Cambra et al.
Fig. 17.16. White patterns (calico) covering most of the leaf area caused by Peach latent mosaic viroid
in leaves of nectarine cultivar ‘Red-Gin’.
from the presence of an insertion of 12–13
nucleotides (Malfitano et al., 2003).
Epidemiology
PLMVd is transmitted by grafting with
infected material and mechanically as a result
of agronomic practices, basically pruning
(Hadidi et al., 1997), but not by pollen, seed or
mites. The viroid might be also transmitted
by aphids (Myzus persicae), although with a
very low efficiency (Flores et al., 1992).
Control
The only effective control measure is the use
of viroid-free propagation material. PLMVd-
infected peach material can be cured by ther-
motherapy, followed by tip grafting and
subsequent propagation (Desvignes, 1986;
Barba et al., 1995). The use of disinfested prun-
ing tools is recommended.
17.8 Peach Dapple
Economic importance and geographic
distribution
Peach dapple (PD) disease, despite affecting
fruit quality, is of limited economic signifi-
cance because it has been reported only in
certain peach varieties grown in Japan (Sano,
2003). However, since its causal agent, Hop
stunt viroid (HSVd), has been found infecting
peach in other areas of the world (Flores and
Llácer, 1988), the disease may have a wider
geographic distribution but may have
remained unnoticed because symptoms are
expressed clearly only in some varieties.
Symptoms
Symptoms consist of chlorotic blotches on the
skin of the mature fruit, which can become
crinkled depending on varietal sensitivity.
Symptoms are not expressed in other organs.
 Viruses and Viroids
Causal agent and diagnosis
HSVd variants of 297 nucleotides, initially
reported from dapple peach (and plum) (Sano
et al., 1989), have been shown to be the causal
agent of PD disease (Terai et al., 1990). This
viroid, which has been found to naturally
infect a wide range of plants, can be detected
by bioassay in cucumber and other species of
the family Cucurbitaceae (symptoms include
leaf curling and stunting). PAGE can also be
applied for diagnostic purposes, although the
low titre of the viroid limits the reliability of
this technique. For this reason, dot-blot
hybridization or RT-PCR procedures are rec-
ommended.
Epidemiology
The viroid is transmitted by grafting with
infected material and, possibly, by mechanical
inoculation with contaminated pruning tools.
Control
Control is essentially the same as for PLMVd:
use of viroid-free propagation material and
decontamination of pruning instruments.
17.9 Peach Mosaic
Economic importance and geographic
distribution
The peach mosaic disease was first described
in Texas and Colorado, USA in the early 1930s
and later surveys indicated its presence in
other south-western states as well as in sev-
eral states of Mexico (Cochran and Pine, 1958;
Pine, 1976). Owing to its high transmissibility
and the severity of the symptoms, extensive
eradication efforts were carried out from 1930
to 1950 in the USA. This led to a very large
reduction in the prevalence of the disease so
that only a few cases were reported during
the 1980s. In the absence of such eradication
measures in Mexico, prevalence appears to
455
have remained very high in at least some
states (Larsen and Oldfield, 1995; Oldfield
et al., 1995). There is currently no evidence for
the presence of this disease outside North
America (Desvignes, 1999).
Symptoms
There has been quite a bit of confusion
between the peach mosaic disease caused by
the recently described Peach mosaic virus
(PcMV) and that caused by PLMVd (see
above). Since many of the early descriptions
of symptoms were made at a time when
PcMV and PLMVd were not known, the
actual identity of the causal agents at that
time is not clear.
Symptoms are reported to be variable,
depending on host cultivar, viral isolate, time
of the year and the existence of co-infections
with other agents. Flower breaking (discolor-
ation of petals) may be seen in some varieties,
as well as reduced petal size and petal defor-
mation. On leaves, typical symptoms are
chlorotic spots and streaks, together with
reduced leaf size, leaf deformation and crin-
kling of the leaf margin. Chlorotic areas may
become necrotic and abscise. Some growth
reduction may be observed, together with
rosetting or shoot proliferation. Fruit may
become deformed and unmarketable (Larsen
and Oldfield, 1995). Other hosts, such as
European and Japanese plums or apricot,
may also show symptoms of similar type.
Causal agent and diagnosis
A member of the genus Trichovirus, PcMV,
which is serologically related to Cherry mottle
leaf virus (CMLV), has been associated with
the peach mosaic disease (Gispert et al., 1998;
James and Howell, 1998). The two viruses
show similar genomic organizations.
PcMV can be identified by: (i) indexing
in susceptible peach varieties; (ii) serological
detection using cross-reacting polyclonal or
monoclonal antibodies reacting also to CMLV;
or (iii) use of the RT-PCR assay developed by
James and Upton (1999), which allows the
456 M. Cambra et al.
simultaneous detection and differentiation of
PcMV and CMLV.
Epidemiology
PcMV is transmitted by the peach bud mite
Eriophyes insidiosus (Keifer and Wilson, 1955;
Oldfield, 1970; Oldfield et al., 1995). The virus
has also been spread by this vector to other
commercially grown Prunus spp. such as
apricot and almond. Some American wild
Prunus spp. are usually symptomless hosts
but their potential as reservoirs for the virus
is not precisely known.
Control
The most effective control measures are: (i)
quarantine in unaffected regions; and (ii)
prompt removal and destruction of infected
trees coupled with the use of certified virus-
tested planting material. Such a strategy has
been used with success in the USA, greatly
reducing the incidence of the disease over a
long period of time.
Since the mite vectors have a strong asso-
ciation with the hosts (overwintering in buds),
control of the disease has been attempted
with some success through direct action
against the vectors with pre-bloom and petal-
fall miticide applications (Jones et al., 1970).
17.10 Diseases of Unknown Aetiology
and Experimental Infections in Peach
The use of biological indexing on woody dif-
ferential hosts is frequently a compulsory
step in a certification programme. Because
many graft-transmissible diseases could be
symptomless in the original host, this tech-
nique often uses pathogen-sensitive geno-
types of the same genus or family as the
indexing host. More than 100 virus-like dis-
eases have been reported to affect Prunus spp.
worldwide but for half of these diseases noth-
ing is known about the causal agent except
that it is graft-transmissible. Peach varieties
are frequently among the hosts used for
indexing for Prunus diseases, with the objec-
tive of detecting graft-transmissible patho-
gens, validating the absence of pathogens and
the elimination of such agents from propaga-
tive budwood.
These indexing techniques are commonly
performed on plants maintained in nurseries
or in greenhouses. In nurseries, the main
objective is to observe disease symptoms that
develop after several years in fruit and/or
bark. In greenhouses, the indexing is recom-
mended to detect diseases with symptoms
that develop rapidly, such as leaf symptoms
or growth-modifying symptoms. Such index-
ing assays on peach varieties have led to the
description of several diseases in peach that
have never been observed in the field. Among
these, some are caused by known viruses
while others are of unknown aetiology. Given
the susceptibility of peach and the potential
for transmission of viruses between Prunus
spp. including peach, the potential for the
natural occurrence of these diseases in peach
cannot be ruled out.
Apricot latent virus, Peach sooty ringspot
virus and Peach asteroid spot virus
Apricot latent virus, Peach sooty ringspot virus
and Peach asteroid spot virus are three related
agents belonging to the genus Foveavirus that
so far have only been reported in apricot. In
this host, they have been observed both in
symptomless plants and in plants displaying
scattered foliar symptoms on new growth
(Zemtchik et al., 1998; Gentit et al., 2001b). On
peach trees, they induce characteristic symp-
toms in the form of chlorotic spots on young
leaves that later develop into sooty green
rings on a yellow background as the leaves
mature and enter senescence (Figs 17.17 and
17.18/Plates 183 and 184).
Apple chlorotic leaf spot virus
Apple chlorotic leaf spot virus (ACLSV), which
is the type member of the genus Tricho-
virus, is common on many Prunus spp. hosts
 Viruses and Viroids
(Németh, 1986). In general, cultivars of plum,
cherry, apricot or peach are scarcely affected
by infection by ACLSV isolates (Hansen and
Gilmer, 1976; Desvignes, 1999). However,
some isolates of ACLSV, usually found on
cherry or on apricot, are particularly severe.
When experimentally transmitted to peach
they can induce stem pitting on bark, yellow-
ish line patterns or russet ring on leaves, as
well as chlorotic rings or mosaic on fruit (Fig.
17.19/Plate 185).
Nepoviruses
Nepoviruses generally have a wide host
range and can be detected in artificially inoc-
ulated annual or perennial weeds, woody
hosts, shrubs or trees. Artificial inoculation of
457
Fig. 17.17. Leaf symptoms (chlorotic
spots) caused by Peach sooty ringspot
virus in peach.
peach by several nepoviruses, for which
peach has not been reported to be a natural
host, can result in disease symptoms. This
includes reduced vigour with shortened
internodes (rosetting) associated with chloro-
tic mottling and curling of leaves (i.e. Arabis
mosaic virus, Myrobalan latent ringspot virus,
Stocky prune virus, Apricot latent ringspot virus);
xylem stem pitting (Myrobalan latent ringspot
virus); and union necrosis (Apricot latent ring-
spot virus) (Dunez and Dupont, 1976; Desvi-
gnes, 1999; Gentit et al., 2001a).
17.11 Concluding Remarks and
Perspectives
Many viruses and viroids of peach diseases
can be spread naturally in the orchard, with
458 M. Cambra et al.
different efficiencies, by vectors, through
pollen or through agricultural practices. In
addition, all viruses or virus-like agents are
propagated together with their host during
all vegetative multiplication practices. The
initial planting of certified virus-tested plant
material is therefore the first step in maintain-
ing a healthy and economically profitable
peach industry. Nurseries producing basic
propagation material need to be separated
from commercial orchards to prevent infec-
tions by viruliferous aphid species or pollen
flow from neighbouring diseased trees. Today,
in vitro propagation and subsequent appro-
priate cultivation under screenhouse or in
glasshouse facilities is, in many nurseries, a
routine practice that considerably reduces the
risks of infection.
Fig. 17.18. Sooty green rings on a
yellow background in senescent
leaves caused by Peach asteroid spot
virus in peach.
Prophylaxis to eliminate or at least
reduce the prevalence of graft-transmissible
agents such as viruses and viroids in the envi-
ronment is a second essential element of any
control strategy. Quarantine measures, eradi-
cation programmes, efforts to limit the spread
of the pathogens through the use of disinfes-
tation measures for pruning tools and, when
appropriate, the use of nematicides are
strongly recommended to complement the
use of certified virus-tested plant material.
Thus, if a viral disease is detected in an estab-
lished orchard, a prudent measure would be
the prompt removal and destruction of the
diseased trees and, if appropriate, treatment
of the site prior to any replanting.
In a globalized world, the emergence of
new graft- and vector-transmitted diseases of
 Viruses and Viroids 459

Fig. 17.19. Leaf symptoms (yellowish line patterns and russet rings) and fruit symptoms (chlorotic
rings and mosaic) caused by Apple chlorotic leaf spot virus in peach.

peach trees, but also the development of new
detection methods and tools, is expected. In
the near future, the detection of plant patho-
gens will probably be based on standardized
and validated diagnosis protocols making
use of more reliable and robust techniques
and reagents. These techniques may gradu-
ally replace indexing, contributing to a more
accurate control of viruses and viroids in
propagative plant materials. However, it
should be stressed that the development of
more powerful detection techniques is only
feasible once the agent to be detected has
been characterized. For diseases of unknown
aetiology, biological indexing will remain the
sole diagnostic option as long as the pathogen
responsible has not been identified and char-
acterized, which stresses the importance of
continued research in this area.
In parallel with direct and indirect con-
trol measures, the development of peach trees
resistant to different viruses and viroids needs
to be more actively explored. Both conven-
tional breeding and biotechnological-based
programmes are already being developed for
PPV resistance in some countries, but very
little work has been devoted to the develop-
ment of resistance towards other agents. The
screening of different available Prunus spp. or
related germplasms (sexually compatible
with peach or not) has great potential to iden-
tify resistance sources against viruses and
viroids. Genetic transformation and the sub-
sequent recovery of transgenic peach culti-
vars have so far stumbled on serious technical
difficulties. Even for other Prunus spp. in
which genetic transformation has been suc-
cessfully developed, such as apricot and
plum, the techniques still suffer from serious
limitations, such as the obligate use of embryo
tissues, with then ensuing loss of varietal
genetic make-up. However, should the prob-
lems currently encountered with peach genetic
transformation be lifted, the availability of
field-validated genetic constructions ensures
that the development of resistant transgenic
peach could be optimistically envisioned.
The generation of new transformation
vectors (some probably based on the viral
genomes), the development of new strategies
(expression of recombinant antibodies specific
for key biological targets and addressed to
460 M. Cambra et al.

the appropriate cell compartment), the search vivo antiviral or antimicrobial activities are
of genomic libraries for peach resistance likely, in time, to contribute to a better protec-
genes or for other defence factors, and the tion of peach trees or to the development of
identification of substances with putative in resistant or tolerant peach varieties.

References

Alayasa, N., Al Rwahnih, M., Myrta, A., Herranz, M.C., Minafra, A., Boscia, D. and Pallás, V. (2003) Identifica-
tion and characterization of an American plum line pattern virus isolate from Palestine. Journal of Plant
Pathology 85, 3–7.
Ambrós, S., Desvignes, J.C., Llácer, G. and Flores, R. (1995) Peach latent mosaic and pear blister canker vi-
roids: detection by molecular hybridization and relationships with specific maladies affecting peach and
pear trees. Acta Horticulturae 386, 515–521.
Aparicio, F., Sánchez-Pina, M.A., Sánchez-Navarro, J.A. and Pallás, V. (1999) Location of prunus necrotic
ringspot ilarvirus within pollen grains of infected nectarine trees: evidence from RT-PCR, dot-blot and in
situ hybridization. European Journal of Plant Pathology 105, 623–627.
Aparicio, F., Sánchez-Navarro, J.A., Herranz, M.C., Myrta, A., Minafra, A. and Pallás, V. (2003) Detection and
identification of seven stone fruit tree viruses by a single multiplex RT-PCR assay with an internal control
RNA. Presented at 19th International Symposium on Virus and Virus-like Diseases of Temperate Fruit
Crops, Valencia, Spain, 21–25 July; abstract book, p. 106.
Asai, W.K. and Uyemoto, J.K. (1991) Peach stunt disease affects on yield. Cling Peach Review 26, 26–27.
Atanasoff, D. (1932) Plum pox. A new virus disease. Yearbook University of Sofia, Bulgaria, Faculty of Agri-
culture 11, 49–69.
Atanasoff, D. (1935) Mosaic of stone fruits. Phytopathologische Zeitschrift 8, 259–284.
Barba, M., Cupidi, A., Loreti, S., Faggioli, F. and Martino, L. (1995) In vitro micrografting: a technique to
eliminate peach latent mosaic viroid from peach. Acta Horticulturae 386, 531–535.
Barrat, J.G., Mircetich, S.M. and Fogle, H.W. (1968) Stem pitting of peach. Plant Disease Reporter 52, 91–94.
Bassi, D. (2006) Breeding for resistance: breeding for resistance to Plum pox virus in Italy. Bulletin OEPP/EPPO
Bulletin 36, 327–329.
Bertolini, E., Olmos, A., Martínez, M.C., Gorris, M.T. and Cambra, M. (2001) Single-step multiplex RT-PCR for
simultaneous and colorimetric detection of six RNA viruses in olive trees. Journal of Virological Methods
96, 33–41.
Boscia, D., Zeramdini, H., Cambra, M., Potere, O., Gorris, M.T., Myrta, A., DiTerlizzi, B. and Savino, V. (1997)
Production and characterization of a monoclonal antibody specific to the M serotype of plum pox poty-
virus. European Journal of Plant Pathology 103, 477–480.
Bousalem, M., Candresse, T., Quiot-Douine, L. and Quiot, J.B. (1994) Comparison of three methods for assess-
ing plum pox virus variability: further evidence for the existence of major groups of isolates. Journal of
Phytopathology 142, 163–172.
Cambra, M., Asensio, M., Gorris, M.T., Pérez, E., Camarasa, E., García, J.A., Moya, J.J., López-Abella, D., Vela,
C. and Sanz, A. (1994) Detection of plum pox potyvirus using monoclonal antibodies to structural and
non-structural proteins. Bulletin OEPP/EPPO Bulletin 24, 569–577.
Cambra, M., Olmos, A., Gorris, M.T., Durán, N., Román, M.P., Camarasa, E. and Dasí, M.A. (1997) Sensitive
detection of plant pathogens by using immobilized targets in tissue imprinted membranes. In: Dehne,
H.W., Adam, G., Diekmann, M., Frahm, J., Mauler-Machnik, A. and van Halteren, P. (eds) Diagnosis and
Identification of Plant Pathogens. Kluwer Academic Publishers, London, pp. 95–99.
Cambra, M., Capote, N., Myrta, A. and Llácer, G. (2006) Plum pox virus and the estimated costs associated
with sharka disease. Bulletin OEPP/EPPO Bulletin 36, 202–204.
Candresse, T., Macquaire, G., Lanneau, M., Bousalem, M., Wetzel, T., Quiot-Douine, L., Quiot, J.B. and
Dunez, J. (1994) Detection of plum pox virus and analysis of its molecular variability using immunocap-
ture-PCR. EPPO Bulletin 24, 585–594.
Candresse, T., Macquaire, G., Lanneau, M., Bousalem, M., Quiot-Douine, L., Quiot, J.B. and Dunez, J. (1995)
Analysis of plum pox virus variability and development of strain-specific PCR assays. Acta Horticulturae
386, 357–369.
Candresse, T., Cambra, M., Dallot, S., Lanneau, M., Asensio, M., Gorris, M.T., Revers, F., Macquaire, G., Olmos, A.,
Boscia, D., Quiot, J.B. and Dunez, J. (1998a) Comparison of monoclonal antibodies and polymerase
 Viruses and Viroids 461

chain reaction assays for the typing of isolates belonging to the M and D serotypes of plum pox potyvirus.
Phytopathology 88, 198–204.
Candresse, T., Kofalvi, S.A., Lanneau, M. and Dunez, J. (1998b) A PCR-ELISA procedure for the simultaneous
detection and identification of prunus necrotic ringspot (PNRSV) and apple mosaic (ApMV) ilarviruses.
Acta Horticulturae 472, 219–224.
Capote, N., Gorris, M.T., Martínez, M.C., Asensio, M., Olmos, A. and Cambra, M. (2006) Interference
between D and M types of Plum pox virus assessed by specific monoclonal antibodies and quantitative
real-time RT-PCR. Phytopathology 96, 320–325.
Carles, L. (1984) Le court-noué du pêcher. Arboriculture Fruitière 364, 31–33.
Cochran, L.C. and Pine, T.S. (1958) Present status of information on host range and host reactions to peach
mosaic virus. Plant Disease Reporter 42, 1225–1228.
Cole, A., Mink, G.I. and Regev, S. (1982) Location of prunus necrotic ringspot virus on pollen grains from
infected almond and cherry trees. Phytopathology 72, 1542–1545.
Crosslin, J.M., Hammond, R.W. and Hammerschlag, F.A. (1992) Detection of prunus necrotic ring spot virus
in herbaceous hosts and Prunus hosts with a complementary RNA probe. Plant Disease 76, 1132–
1136.
Dal Zotto, A., Ortego, J.M., Raigón, J.M., Caloggero, S., Rossini, M. and Duchase, D.A. (2006) First report in
Argentina of plum pox virus causing sharka disease in Prunus. Plant Disease Note 90, 523.
Desvignes, J.C. (1976) The virus diseases detected in greenhouse and in field by the peach seedling GF 305
indicator. Acta Horticulturae 67, 315–323.
Desvignes, J.C. (1986) Peach latent mosaic and its relation to peach mosaic and peach yellow mosaic virus
diseases. Acta Horticulturae 193, 51–57.
Desvignes, J.C. (1999) Virus Diseases of Fruit Trees. CTIFL, Paris.
Dias, H.F. (1975) Peach rosette mosaic virus. In: Harrison, B.D. and Murant, A.F. (eds) CMI/AAB Descriptions
of Plant Viruses. Commonwealth Agricultural Bureaux Press, Slough, UK, No. 150.
Dias, H.F. and Cation, D. (1976) The characterization of a virus responsible for peach rosette mosaic and
grape decline in Michigan. Canadian Journal of Botany 54, 1228–1239.
Dicenta, F. and Audergon, J.M. (1998) Inheritance of resistance to plum pox potyvirus (PPV) in Stella apricot
seedlings. Plant Breeding 117, 579–581.
Digiaro, M., Di Terlizzi, B. and Savino, V. (1992) Ilarviruses in apricot and plum pollen. Acta Horticulturae
309, 94–98.
Dosba, F. (1992) Plum pox potyvirus. In: Smith, I.M., McNamara, D.G., Scott, P.R. and Harris, K.M. (eds)
Quarantine Pests for Europe. Data Sheets on Quarantine Pests for the European Communities and for the
European and Mediterranean Plant Protection Organization. CAB International, Wallingford, UK and
European and Mediterranean Plant Protection Organization, Paris, pp. 922–927.
Dosba, F., Lansac, M., Audergon, J.M. and Massonie, G. (1989) Tolerance to plum pox in apricot. Acta Horti-
culturae 235, 275–282.
Dosba, F., Maison, P., Denise, F., Audergon, J.M. and Massonie, G. (1991) Plum pox resistance in apricot. Acta
Horticulturae 293, 569–573.
Dunez, J. and Dupont, G. (1976) Myrobalan latent ringspot virus. In: Harrison, B.D. and Murant, A.F. (eds) CMI/
AAB Descriptions of Plant Viruses. Commonwealth Agricultural Bureaux Press, Slough, UK, No. 160.
Dunez, J. and Sutic, D. (1988) Plum pox virus. In: Smith, I.M., Dunez, J., Elliot, R.A., Phillips, D.H. and
Arches, S.A. (eds) European Handbook of Plant Diseases. Blackwell, London, pp. 44–46.
EPPO (2004) Diagnostic protocol for regulated pests. Plum pox potyvirus. Bulletin OEPP/EPPO Bulletin 34,
247–256.
Escalettes, V., Lansac, M., Eyquard, J.P. and Dosba, F. (1998) Genetic resistance to plum pox potyvirus in
peaches. Acta Horticulturae 465, 689–697.
Esteban, O., García, J.A., Gorris, M.T., Domínguez, E. and Cambra, M. (2003) Generation and characterisa-
tion of functional recombinant antibody fragments against RNA replicase NIb from plum pox virus.
Biochemical and Biophysical Research Communications 301, 167–175.
Flores, R. and Llácer, G. (1988) Isolation of a viroid-like RNA associated with peach latent mosaic disease.
Acta Horticulturae 235, 325–332.
Flores, R., Hernández, C., Desvignes, J.C. and Llácer, G. (1990) Some properties of the viroid inducing the
peach latent mosaic disease. Research in Virology 141, 109–118.
Flores, R., Hernández, C., Avinent, L., Hermoso, A., Llácer, G., Juárez, J., Arregui, J.M., Navarro, L. and Des-
vignes, J.C. (1992) Studies on the detection, transmission and distribution of peach latent mosaic viroid
in peach trees. Acta Horticulturae 309, 325–330.
462 M. Cambra et al.

Flores, R., Randles, J.W., Bar-Joseph, M. and Diener, T.O. (2000) Viroids. In: van Regenmortel, M.H.V., Fauquet,
C.M., Bishop, D.H.L., Carstens, E.B., Estes, M.K., Lemon, S.M., Maniloff, J., Mayo, M.A., McGeoch, D.J.,
Pringle, C.R. and Wickner, R.B. (eds) Virus Taxonomy, 7th Report of the International Committee on
Taxonomy of Viruses. Academic Press, San Diego, California, pp. 1009–1024.
Fortusini, A., Bianco, P.A. and Belli, G. (1983) Sintomatologia e danni da rosetta a foglie saliciformi in pe-
scheti del Piemonte. L’Informatore Agrario 30, 26837–26840.
Fulton, R.W. (1982) Ilar-like characteristics of American plum line pattern virus and its serological detection
in Prunus. Phytopathology 72, 1345–1348.
García, J.A. and Cambra, M. (2007) Plum pox virus and sharka disease. Plant Viruses 1, 69–79.
Gentit, P., Delbos, R.P., Candresse, T. and Dunez, J. (2001a) Characterization of a new nepovirus infecting apricot
in southeastern France: apricot latent ringspot virus. European Journal of Plant Pathology 107, 485–494.
Gentit, P., Foissac, X., Svanella-Dumas, L. and Candresse, T. (2001b) Variants of apricot latent foveavirus
(ALV), isolated from south European orchards associated with peach asteroid spot and peach sooty ring-
spot diseases. Acta Horticulturae 550, 213–219.
Gispert, C., Perring, T.M. and Creamer, R. (1998) Purification and characterization of peach mosaic virus.
Plant Disease 82, 905–908.
Glasa, M., Palkovics, L., Komínek, P., Labonne, G., Pittnerová, S., Kúdela, O., Candresse, T. and Šubr, Z.
(2004) Geographically and temporally distant natural recombinant isolates of plum pox virus (PPV) are
genetically very similar and form a unique PPV group. Journal of General Virology 85, 2671–2681.
Greber, R.S., Teakle, D.S. and Mink, G.I. (1992) Thrips-facilitated transmission of prune dwarf and prunus
necrotic ringspot viruses from cherry pollen to cucumber. Plant Disease 76, 1039–1041.
Griesbach, J.A. (1995) Detection of tomato ringspot virus by polymerase chain reaction. Plant Disease 79,
1054–1056.
Hadidi, A., Giunchedi, L., Shamloul, A.M., Poggi-Pollini, C. and Amer, M.A. (1997) Occurrence of peach latent
mosaic viroid in stone fruits and its transmission with contaminated blades. Plant Disease 81, 154–158.
Halk, E.L., Hsu, H.T. and Franke, J. (1984) Production of antibodies against three ilarviruses and alfalfa mo-
saic virus and their use in serotyping. Phytopathology 74, 367–372.
Hamilton, R.I., Nichols, C. and Valentina, B. (1984) Survey of prunus necrotic ringspot and other viruses con-
taminating the exine of pollen collected by bees. Canadian Journal of Plant Pathology 6, 196–199.
Hansen, A.J. and Gilmer, R.M. (1976) In: Virus Diseases and Non-Infectious Disorders of Stone Fruits in North
America. USDA Handbook No. 437. US Department of Agriculture–Agricultural Research Service,
Washington, DC, pp. 206–208.
Helguera, P.R., Taborda, R., Docampo, D.M. and Ducasse, D.A. (2001) Immunocapture reverse transcription-
polymerase chain reaction combined with nested PCR greatly increases the detection of prunus necrotic
ring spot virus in the peach. Journal of Virological Methods 95, 93–100.
Heuss-La Rosa, K., Hammond, R., Crosslin, J.M., Hazel, C. and Hammerschlag, F.A. (1995) Monitoring of
prunus necrotic ringspot virus infection by hybridization with a cRNA probe after in vitro micrografting.
Journal of the American Society for Horticultural Science 120, 928–931.
Hily, J.M., Scorza, R., Malinowski, T., Zawadzka, B. and Ravelonandro, M. (2004) Stability of gene silencing-
based resistance to plum pox virus in transgenic plum (Prunus domestica L.) under field conditions.
Transgenic Research 13, 427–436.
James, D. and Howell, W.E. (1998) Isolation and partial characterization of a filamentous virus associated with
peach mosaic disease. Plant Disease 82, 909–913.
James, D. and Upton, C. (1999) Single primer pair designs that facilitate simultaneous detection and differen-
tiation of peach mosaic virus and cherry mottle leaf virus. Journal of Virological Methods 83, 103–111.
James, D., Varga, A., Thompson, D. and Hayes, S. (2003) Detection of a new and unusual isolate of plum pox
potyvirus in plum (Prunus domestica). Plant Disease 87, 1119–1124.
Johansen, E., Edwards, M.C. and Hampton, R.O. (1994) Seed transmission of viruses: current perspectives.
Annual Review of Phytopathology 32, 363–386.
Jones, L.S., Wilson, N.S., Burr, W. and Barnes, M.M. (1970) Restriction of the peach mosaic virus spread through
control of the vector mite, Eriophyes insidiosus. Journal of Economic Entomology 63, 1551–1552.
Karayiannis, I., Mainou, A. and Tsaftaris, I. (1999) Apricot breeding in Greece for fruit quality and resistance
to plum pox virus. Acta Horticulturae 488, 111–117.
Keifer, H.H. and Wilson, N.S. (1955) A new species of eriophyd mite responsible for the vection of peach
mosaic virus. Bulletin of the Californian Department of Agriculture 44, 145–146.
Kelley, R.D. and Cameron, H.R. (1986) Location of prune dwarf and prunus necrotic ringspot viruses associ-
ated with sweet cherry pollen and seed. Phytopathology 76, 317–322.
 Viruses and Viroids 463

Kerlan, C. and Dunez, J. (1976) Some properties of plum pox virus and its nucleic acid and protein compo-
nents. Acta Horticulturae 67, 185–192.
Klos, E.J. (1976) Rosette mosaic. In: Virus Diseases and Non-Infectious Disorders of Stone Fruits in North
America. USDA Handbook No. 437. US Department of Agriculture–Agricultural Research Service,
Washington, DC, pp. 135–138.
Kölber, M., Nemeth, M., Krizbai, L., Szemes, M. and Kiss-Toth, E. (1998) Detectability of prunus necrotic
ringspot and plum pox virus by RT-PCR, ELISA and indexing on woody indicators. Acta Horticulturae
472, 243–247.
Korschineck, I., Himmler, G., Sagl, R., Steinkellner, H. and Kattinger, H.W.D. (1991) A PCR membrane spot
assay for the detection of plum pox virus RNA in bark of infected trees. Journal of Virological Methods
31, 139–146.
Kreiah, S., Strunk, G. and Cooper, J.I. (1994) Sequence analysis and location of capsid proteins within RNA2
of strawberry latent ringspot virus. Journal of General Virology 75, 2527–2532.
Kunze, L. and Krczal, H. (1971) Transmission of sharka virus by aphids. Annals de Phytopathologie, Hors
Série 255–260.
Labonne, G., Yvon, M., Quiot, J.B., Avinent, L. and Llácer, G. (1995) Aphids as potential vectors of plum pox
virus: comparison of methods of testing and epidemiological consequences. Acta Horticulturae 386,
207–218.
Laimer da Camara Machado, M., da Camara Machado, A. and Hanzer, V. (1992) Regeneration of transgenic plants
of Prunus armeniaca containing the coat protein gene of plum pox virus. Plant Cell Reports 11, 25–29.
Larsen, H.J. and Oldfield, G.N. (1995) Peach mosaic. In: Ogawa, J.M., Zehr, E.I., Bird, G.W., Ritchie, D.F., Uriu, K.
and Uyemoto, J.K. (eds) Compendium of Stone Fruit Diseases. APS Press, St. Paul, Minnesota, pp. 67–68.
Levy, L., Lee, I.M. and Hadidi, A. (1994) Simple and rapid preparation of infected plant tissue extracts for PCR
amplification of virus, viroid and MLO nucleic acids. Journal of Virological Methods 49, 295–304.
Levy, L., Damsteegt, V. and Welliver, R. (2000) First report of plum pox virus (sharka disease) in Prunus persi-
cae in the United States. Plant Disease 84, 202 (Disease note).
López-Moya, J.J. and García, J.A. (1999) Potyviruses (Potyviridae). In: Webster, R.G. and Granoff, A. (eds)
Encyclopaedia of Virology, 2nd edn, Vol. 3. Academic Press, London, pp. 1369–1375.
López-Moya, J.J., Cantó, T., Díaz-Ruíz, J.R. and López-Abella, D. (1995) Transmission by aphids of a naturally
non-transmissible plum pox virus isolate with the aid of potato virus Y helper component. Journal of
General Virology 76, 2293–2297.
Loreti, S., Faggioli, F. and Barba, M. (1995) A rapid extraction method to detect peach latent mosaic viroid by
molecular hybridization. Acta Horticulturae 386, 560–564.
Maiss, E., Timpe, U., Brisske, A., Jelkmann, W., Casper, R., Himmler, G., Mattanovich, D. and Katinger, H.W.D. (1989)
The complete nucleotide sequence of plum pox virus RNA. Journal of General Virology 70, 513–524.
Malfitano, M., di Serio, F., Covelli, L., Ragozzino, A., Hernández, C. and Flores, R. (2003) Peach latent mo-
saic viroid variants inducing peach calico contain a characteristic insertion that is responsible for this
symptomatology. Virology 313, 492–501.
Mayo, M.A. and Robinson, D.J. (1996) Nepoviruses: molecular biology and replication. In: Harrison, B.D. and
Murant, A.F. (eds) The Plant Viruses. Vol. 5. Polyhedral Virions and Bipartite RNA Genomes. Plenum
Press, New York, pp. 139–185.
Mink, G.I. (1992) Ilarvirus vectors. Advances in Disease Vector Research 9, 261–281.
Mink, G.I. and Aichele, M.D. (1984) Detection of prunus necrotic ringspot and prune dwarf viruses in Prunus
seed and seedlings by enzyme-linked immunosorbent assay. Plant Disease 68, 378–381.
Murant, A.F. (1974) Strawberry latent ringspot virus. In: Harrison, B.D. and Murant, A.F. (eds) CMI/AAB De-
scriptions of Plant Viruses. Commonwealth Agricultural Bureaux Press, Slough, UK, No. 126.
Myrta, A., Potere, O., Boscia, D., Candresse, T., Cambra, M. and Savino, V. (1998) Production of a monoclonal
antibody specific to the El Amar strain of plum pox virus. Acta Virologica 42, 248–250.
Myrta, A., Potere, O., Crescenzi, A., Nuzzaci, M. and Boscia, D. (2000) Production of two monoclonal antibod-
ies specific to cherry strain of plum pox virus (PPV-C). Journal of Plant Pathology 82(Suppl. 2), 95–103.
Myrta, A., Abbadi, H., Herranz, Al Rwahnih, M., Di Terlizzi, B., Minafra, A. and Pallás, V. (2002) First report
of American plum line pattern virus (PLPV) in Albania, Italy and Tunisia. Journal of Plant Pathology 84,
188 (Disease note).
Navrátil, M., Safarova, D., Karesova, R. and Petrzik, K. (2005) First incidence of plum pox virus on apricot
trees in China. Plant Disease Note 89, 338.
Nemchinov, L. and Hadidi, A. (1996) Characterization of the sour cherry strain of plum pox virus. Phytopa-
thology 86, 575–580.
464 M. Cambra et al.

Nemchinov, L., Hadidi, A., Maiss, E., Cambra, M., Candresse, T. and Damsteegt, V. (1996) Sour cherry strain
of plum pox potyvirus (PPV): molecular and serological evidence for a new subgroup of PPV strains.
Phytopathology 86, 1215–1221.
Németh, M. (1986) Virus, Mycoplasma and Rickettsia Diseases of Fruit Trees, 1st edn. Martinus Nijhoff
Publishers, Dordrecht, The Netherlands.
Németh, M. (1994) History and importance of plum pox in stone-fruit production. Bulletin OEPP/EPPO
Bulletin 24, 525–536.
Oldfield, G.N. (1970) Mite transmission of plant viruses. Annual Review of Phytopathology 69, 854–858.
Oldfield, G.N., Creamer, R., Gispert, C., Osorio, F., Rodríguez, R. and Perring, T.M. (1995) Incidence and
distribution of peach mosaic and its vector, Eriophyes insidiosus (Acari: Eriophyidae) in Mexico. Plant
Disease 79, 186–189.
Olmos, A., Dasí, M.A., Candresse, T. and Cambra, M. (1996) Print-capture-PCR: a simple and highly sensitive
method for the detection of plum pox virus (PPV) in plant tissues. Nucleic Acids Research 24, 2192–
2193.
Olmos, A., Cambra, M., Dasí, M.A., Candresse, T., Esteban, O., Gorris, M.T. and Asensio, M. (1997) Simulta-
neous detection and typing of plum pox potyvirus (PPV) isolates by heminested-PCR and PCR-ELISA.
Journal of Virological Methods 68, 127–137.
Olmos, A., Cambra, M., Esteban, O., Gorris, M.T. and Terrada, E. (1999) New device and method for capture,
reverse transcription and nested PCR in a single closed tube. Nucleic Acids Research 27, 1564–1565.
Olmos, A., Bertolini, E. and Cambra, M. (2002) Simultaneous and co-operational amplification (Co-PCR) for
detection of plant viruses. Journal of Virological Methods 106, 51–59.
Olmos, A., Esteban, O., Bertolini, E. and Cambra, M. (2003) Nested RT-PCR in a single closed tube. In:
Bartlett, J.M.S. and Stirling, D. (eds) Methods in Molecular Biology, 2nd edn, Vol. 226. PCR Protocols.
Humana Press, Totowa, New Jersey, pp. 153–161.
Olmos, A., Bertolini, E., Gil, M. and Cambra, M. (2005) Real-time assay for quantitative detection of non-
persistently transmitted plum pox virus RNA targets in single aphids. Journal of Virological Methods 128,
151–155.
Pallás, V., Sánchez-Navarro, J.A., Más, P., Cañizares, M.C., Aparicio, F. and Marcos, J.F. (1998) Molecular
diagnostic techniques and their potential role in stone fruit certification schemes. Options Méditerranée-
nnes 19, 191–208.
Parakh, D.R., Shamloul, A.M., Hadidi, A., Scott, S.W., Waterworth, H.E., Howell, H.E. and Mink, G.I. (1995)
Detection of prune dwarf ilarvirus from infected stone fruits using reverse transcription-polymerase chain
reaction. Acta Horticulturae 386, 421–430.
Pascal,T., Kervella, J., Pfeiffer, F., Sauge, M.H. and Esmenjaud, M. (1997) Evaluation of inter-specific progeny
of Prunus persica cv. Summergrand × Prunus davidiana for disease resistance and some agronomic fea-
tures. Acta Horticulturae 465, 185–192.
Pasquini, G. and Barba, M. (1994) Serological characterization of Italian isolates of plum pox potyvirus. Bul-
letin OEPP/EPPO Bulletin 24, 615–624.
Pine, T.S. (1976) Peach mosaic. In: Virus Diseases and Non-Infectious Disorders of Stone Fruits in North
America. USDA Handbook No. 437. US Department of Agriculture–Agricultural Research Service,
Washington, DC, pp. 61–70.
Powell, C.A. (1984) Comparison of enzyme-linked immunosorbent assay procedures for detection of tomato
ringspot virus in woody and herbaceous hosts. Plant Disease 68, 908–909.
Powell, C.A., Forer, L.B., Stouffer, R.F., Cummins, J.N., Gonsalves, D., Rosenberger, D.A., Hoffman, J. and
Lister, R.M. (1984) Orchard weeds as hosts of tomato ringspot and tobacco ringspot viruses. Plant Dis-
ease 68, 242–244.
Pusey, P.L. and Yadava, U.L. (1991) Influence of prunus necrotic ringspot virus on growth, productivity, and
longevity of peach trees. Plant Disease 75, 847–851.
Ramsdell, D.C. and Myers, R.L. (1978) Epidemiology of peach rosette mosaic virus in a Concord grape vine-
yard. Phytopathology 68, 447–450.
Ravelonandro, M., Scorza, R., Bachelier, J.C., Labonne, G., Levy, L., Damsteegt, V., Callahan, A. and Dunez, J. (1997)
Resistance of transgenic Prunus domestica to plum pox virus infection. Plant Disease 81, 1231–1235.
Ravelonandro, M., Scorza, R., Callahan, A., Levy, L., Jacquet, C., Monsion, M. and Damsteegt, V. (2000) The
use of transgenic fruit trees as a resistance strategy for virus epidemics: the plum pox (sharka) model.
Virus Research 71, 63–69.
Ravelonandro, M., Scorza, R., Minoiu, M., Zagrai, I. and Platon, I. (2002) Field tests of transgenic plums in
Romania. Sanatatea Plantelor special edition, 16–18.
 Viruses and Viroids 465

Revers, F., Gall, O.L., Candresse, T. and Maule, A.J. (1999) New advances in understanding the molecular
biology of plant–potyvirus interactions. Molecular Plant–Microbe Interactions 12, 367–376.
Riechmann, J.L., Laín, S. and García, J.A. (1992) Highlights and prospects of potyvirus molecular biology.
Journal of General Virology 73, 1–16.
Rowhani, A., Maningas, M.A., Lile, S.D., Daubert, S.D. and Golino, D.A. (1995) Development of a detection system
for viruses of woody plants based on PCR analysis of immobilized virions. Phytopathology 85, 347–352.
Rowhani, A., Biardi, L., Routh, G., Daubert, S.D. and Golino, D. (1998) Development of a sensitive colorimet-
ric-PCR assay for detection of viruses in woody plants. Plant Disease 82, 880–884.
Roy, A.S. and Smith, I.M. (1994) Plum pox situation in Europe. Bulletin OEPP/EPPO Bulletin 24, 515–523.
Saade, M., Aparicio, F., Sánchez-Navarro, J.A., Herranz, M.C., Myrta, A., Di-Terlizzi, B. and Pallás, V. (2000)
Simultaneous detection of the three ilarviruses affecting stone fruit trees by nonisotopic molecular hybrid-
ization and multiplex reverse-transcription polymerase chain reaction. Phytopathology 90, 1330–1336.
Sánchez-Navarro, J.A. and Pallás, V. (1997) Evolutionary relationships in the Ilarviruses: nucleotide sequence
of prunus necrotic ringspot virus RNA 3. Archives of Virology 142, 749–763.
Sánchez-Navarro, J.A., Aparicio, F., Rowhani, A. and Pallás, V. (1998) Comparative analysis of ELISA, nonra-
dioactive molecular hybridisation and PCR for the detection of prunus necrotic ringspot virus in herba-
ceous and Prunus hosts. Plant Pathology 47, 780–786.
Sano, T. (2003) Hop stunt viroid in plum and peach. In: Hadidi, A., Flores, R., Randles, J.W. and Semancik,
J.S. (eds) Viroids. CSIRO Publishing, Collingwood, Australia, pp. 165–167.
Sano, T., Hataya, T., Terai, Y. and Shikata, E. (1989) Hop stunt viroid strains from dapple fruit disease of plum
and peach in Japan. Journal of General Virology 70, 1311–1319.
Saunier, R. (1972) Incidence d’un virus du type ringspot sur le comportement de deux cultivars du pêcher. La
Pomologie Française 14(7), 175–185.
Schlocker, A. and Traylor, J.A. (1976) Yellow bud mosaic. In: Virus Diseases and Non-Infectious Disorders of
Stone Fruits in North America. USDA Handbook No. 437. US Department of Agriculture–Agricultural
Research Service, Washington, DC, pp. 156–165.
Schneider, W.L., Sherman, D.J., Stone, A.L., Damsteegt, V.D. and Frederick, R.D. (2004) Specific detection and
quantification of plum pox virus by real-time fluorescent reverse transcription-PCR. Journal of Virologi-
cal Methods 120, 97–105.
Scorza, R., Ravelonandro, M., Callahan, A.M., Cordts, J.M., Fuchs, M., Dunez, J. and Gonsalves, D. (1994)
Transgenic plums (Prunus domestica L.) express the plum pox virus coat protein gene. Plant Cell Reports
14, 18–22.
Scorza, R., Callahan, A., Levy, L., Damsteegt, V., Webb, K. and Ravelonandro, M. (2001) Post-transcriptional
gene silencing in plum pox virus resistant transgenic European plum containing the plum pox potyvirus
coat protein gene. Transgenic Research 10, 201–209.
Scott, S.W. and Zimmerman, M.T. (2001) American plum line pattern virus is a distinct ilarvirus. Acta Horti-
culturae 550, 221–227.
Scott, S.W., Barnett, O.W. and Burrows, P.M. (1989) Incidence of prunus necrotic ringspot virus in selected
peach orchards of South Carolina. Plant Disease 73, 913–916.
Scott, S.W., Bowman-Vance, V. and Bachman, E.J. (1992) The use of nucleic acid probes for the detection of
prunus necrotic ringspot virus and prune dwarf virus. Acta Horticulturae 309, 79–83.
Scott, S.W., Zimmerman, M.T., Yilmaz, S., Zehr, E.I. and Bachman, E. (2001) The interaction between prunus
necrotic ringspot virus and prune dwarf virus in peach stunt disease. Acta Horticulturae 550, 229–236.
Shamloul, A.M., Minafra, A., Hadidi, A., Giunchedi, L., Waterworth, H.E. and Allam, E.K. (1995) Peach latent
mosaic viroid: nucleotide sequence of an Italian isolate, sensitive detection using RT-PCR and geo-
graphic distribution. Acta Horticulturae 386, 522–530.
Shukla, D.D., Ward, C.W. and Brunt, A.A. (1994) Genome structure, variation and function. In: Shukla, D.D.,
Ward, C.W. and Brunt, A.A. (eds) The Potyviridae. CAB International, Wallingford, UK, pp. 74–112.
Smith, I.M., McNamara, D.G., Scott, P.R. and Harris, K.M. (eds) (1992) Quarantine Pests for Europe. Data
sheets on Quarantine Pests for the European Communities and for the European and Mediterranean
Plant Protection Organization. CAB International, Wallingford, UK and European and Mediterranean
Plant Protection Organization, Paris.
Smith, P.R. and Challen, D.K. (1977) Initial and subsequent yield reduction of peach trees affected by peach
rosette and decline disease. Australian Journal of Agricultural Research 28, 441–444.
Spiegel, S., Scott, S.W., Bowman-Vance, V., Tam, V., Galiakparov, N.N. and Rosner, A. (1996) Improved detec-
tion of prunus necrotic ringspot virus by the polymerase chain reaction. European Journal of Plant
Pathology 102, 681–685.
466 M. Cambra et al.

Spiegel, S., Kovalenko, E., Varga, A. and James, D. (2004) Detection and partial molecular characterization of
two plum pox virus isolates from plum and wild apricot in Southeast Kazakhstan. Plant Disease 88,
973–979.
Stace-Smith, R. (1984) Tomato ringspot virus. In: Harrison, B.D. and Murant, A.F. (eds) CMI/AAB Descriptions
of Plant Viruses. Commonwealth Agricultural Bureaux Press, Slough, UK, No. 290 (No. 18 revised).
Terai, Y., Sano, T. and Shikata, E. (1990) Back inoculation of plum dapple fruit disease and graft transmission
of peach dapple fruit disease. Annals of the Phytopathology Society of Japan 56, 428.
Thompson, D., McCann, M., McLeod, M., Iue, D., Green, M. and James, D. (2001) First report of plum pox
potyvirus in Canada. Plant Disease 85, 97 (Report).
Uyemoto, J.K., Luhn, C.F., Asai, W.K., Beede, R., Beutel, J.A. and Fenton, R. (1989) Incidence of ilarviruses in
young peach trees in California. Plant Disease 73, 217–220.
Uyemoto, J.K., Asai, W.K. and Luhn, F. (1992) Ilarviruses: evidence for rapid spread and effects on vegetative
growth and fruit yields of peach trees. Plant Disease 76, 71–74.
van Regenmortel, M.H.V., Fauquet, C.M., Bishop, D.H.L., Carstens, E.B., Estes, M.K., Lemon, S.M., McGeoch,
D.J., Maniloff, J., Mayo, M.A., Pringle, C.R. and Wickner, R.B. (eds) (2000) Virus Taxonomy. Classifica-
tion and Nomenclature of Viruses. Seventh ICTV Report. Academic Press, New York.
Varga, A. and James, D. (2005) Detection and differentiation of plum pox virus using real-time multiplex PCR
with SYBR green and melting curve analysis: a rapid method for strain typing. Journal of Virological
Methods 123, 213–220.
Varveri, C., Candresse, T., Cugusi, M., Ravelonandro, M. and Dunez, J. (1988) Use of a 32P labelled tran-
scribed RNA probe for dot hybridization detection of plum pox virus. Phytopathology 78, 1280–1283.
Vilanova, S., Romero, C., Abbott, A.G., Llácer, G. and Badenes, M.L. (2003) An apricot (Prunus armeniaca L.)
F2 progeny linkage map based on SSR and AFLP markers, mapping plum pox virus resistance and self-
incompatibility traits. Theoretical and Applied Genetics 107, 239–247.
Wetzel, T., Candresse, T., Ravelonandro, M., Delbos, R.P., Mazyad, H., Aboul-Ata, A.E. and Dunez, J. (1991a)
Nucleotide sequence of the 3′-terminal region of the RNA of the El Amar strain of plum pox potyvirus.
Journal of General Virology 72, 1741–1746.
Wetzel, T., Candresse, T., Ravelonandro, M. and Dunez, J. (1991b) A polymerase chain reaction assay adapted
to plum pox potyvirus detection. Journal of Virological Methods 33, 355–365.
Wetzel, T., Candresse, T., Macquaire, G., Ravelonandro, M. and Dunez, J. (1992) A highly sensitive immuno-
capture polymerase chain reaction method for plum pox potyvirus detection. Journal of Virological
Methods 39, 27–37.
Zemtchik, E.Z., Verderevskaya, T.D. and Kalashian Yu, A. (1998) Apricot latent ringspot virus: transmission,
purification and serology. Acta Horticulturae 472, 153–158.
 18 Insects and Mites

D.L. Horton,1 J. Fuest1 and P. Cravedi2

1Department of Entomology, University of Georgia, Athens, Georgia, USA
2Instituto di Entomologia e Patologia Vegetale, Università del Sacro Cuore,
Piacenza, Italy

18.1 Introduction 467

18.2 Direct Insect Pests 468
Oriental fruit moth, Grapholita molesta (Busck) (Lepidoptera: Tortricidae) 468
Codling moth, Cydia pomonella Linnaeus (Lepidoptera: Tortricidae) 472
Peach twig borer, Anarsia lineatella Zeller (Lepidoptera: Gelechiidae) 473
Tufted apple bud moth, Platynota idaeusalis (Walker) (Lepidoptera: Tortricidae) 475
Plum curculio, Conotrachelus nenuphar (Herbst) (Coleoptera: Curculionidae) 477
Scarab beetles (Coleoptera: Scarabaeidae) 479
Plant bugs (Hemiptera: Miridae), stink bugs (Hemiptera: Pentatomidae) and
leaf-footed bugs (Hemiptera: Coreidae) 480
Thrips (Thysanoptera) 484
Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) 486
18.3 Indirect Insect Pests 487
Peachtree borers, Synanthedon spp. (Lepidoptera: Sesiidae) 487
Armoured scale (Hemiptera: Diaspididae) 491
Aphids (Hemiptera: Aphididae) 494
18.4 Mite Pests 496
Two-spotted spider mite, Tetranychus urticae Koch (Acari: Tetranychidae) 497
European red mite, Panonychus ulmi (Koch) (Acari: Tetranychidae) 498

18.1 Introduction vary with production area and are normally

Insects and, to a more modest extent, mites
are important pests of peaches and nectarines
worldwide. Peaches in commercial culture
today are derived from selections of Prunus
spp. ecotypes native to south-eastern China
(Rieger, 2006). In most of the world’s produc-
tion areas, peaches are an introduced species.
Accordingly, peach arthropod pest complexes
an assortment of insect and mite species
native to the region and exotic species that
have adapted to peach. As is typical for orchard
crops, peaches have a complement of cosmo-
politan pests that are injurious across numer-
ous production areas.
China is the world’s leading peach pro-
ducer. Although southern China and the south-
eastern USA are climatically similar, many of

© CAB International 2008. The Peach: Botany, Production and Uses

(eds D.R. Layne and D. Bassi) 467
468 D.L. Horton et al.
the world’s significant peach production areas
are in arid to semi-arid production areas of
Italy, Spain, Greece, France, Iran, Chile and
the western USA (Rieger, 2006).
Orchard pest management is a pragmatic
discipline that strives to protect fruit while
mitigating tree-attacking pests to sustain
yield potential and enhance tree longevity.
Peaches are a relatively short-lived (Brittain
and Miller, 1978) but high-value crop. Growers
must be mindful of direct, fruit-attacking pests
and they must also diligently address com-
plexes of indirect or tree-attacking insects and
mites. Unless managed properly, tree-attacking
pests can reduce vigour, longevity and profit-
ability. The grower adage ‘take care of the trees,
the trees will take care of you’ is particularly
appropriate for peaches. Nectarine fruits are
particularly sensitive to damage by insects with
piercing–sucking mouthparts like aphids and
thrips. This chapter’s review of peach arthro-
pod pests and current management options is
an effort to address the current status of peach
entomology while highlighting the crop’s
important entomological research needs.
18.2 Direct Insect Pests
Fruit-attacking insect pests, species that feed
on and/or lay eggs in the fruit, may be
referred to as direct pests. Orchardists aim to
produce fruit of a quality that competes well
in the marketplace while protecting tree
health and longevity. Management decisions
must cater to market expectations. Fruit qual-
ity expectations are often on a sliding quality
scale, which shifts with the overall availabil-
ity of that day’s premium grade. Sound fruit
with minor cosmetic insect injury may be dis-
counted or fail to sell. The grower’s best
option is to produce peaches as free from
insect-related blemishes as possible, because
other fruit quality variables, such as cold
injury or low sugar from excessive rainfall at
harvest, are beyond his/her control. Accord-
ingly, peach growers often opt for insecticide-
intensive pest management programmes,
because optimizing the percentage of fruit
that make top grades is frequently rewarded
with better price, lower grading costs, and in
some cases, better control of indirect tree-
attacking pests. Fortunately, newer manage-
ment strategies (insect growth regulator (IGR)
insecticides and mating disruption) are less
disruptive to beneficial insects and allow
greater integration of parasitoids.
Oriental fruit moth, Grapholita molesta
(Busck) (Lepidoptera: Tortricidae)
Oriental fruit moth, Grapholita molesta (Busck)
(OFM), is a serious pest of peaches, nectarines
and quince worldwide. In California, prune,
cherry, almond and apricot are occasional
OFM hosts, and pear and plum suffer vegeta-
tive shoot strikes (Strand, 1999). OFM is
increasingly important as a pest of apples in
many regions (Reis et al., 1988).
OFM larvae feed on succulent terminal
growth during the initial spring and subse-
quent autumn vegetative growth flushes.
Stem feeding by OFM larvae produces with-
ered or dead shoots, which are referred to as
‘flagged’ (Fig. 18.1/Plate 186). Vegetative
shoot feeding typically destroys the shoot’s
distal or terminal bud, increasing lateral
branching as multiple lateral buds break to
vie for dominance. Larvae may continue to
burrow in the original shoot or enter other
shoots before they reach maturity (Hogmire,
1995). Injury from OFM terminal feeding is
more important in young trees, because rapid
growth of young trees is sought to quickly fill
each tree’s space, which optimizes early
yields. Conversely, OFM terminal feeding on
mature trees is of little consequence.
In young trees, the influence of OFM on
terminal growth varies according to tree
training style. OFM injury to terminals in
trees trained to decidedly flat, open-vase sys-
tems is seldom significant. Open-vase train-
ing systems emphasize heading cuts to
scaffold limbs (Lockwood and Myers, 2005).
These cuts systematically prune out terminal
growth to stimulate development of second-
ary and tertiary scaffold limbs. These pre-
scribed heading cuts remove the terminals of
most scaffolds, rendering OFM injury to
them irrelevant. Conversely, upright-vase or
V-tree systems require unbranched scaffold
limbs, because loss of terminal dominance in
 Insects and Mites
a scaffold limb slows growth, decreases limb
flexibility under fruit load and reduces early
yields (Marini and Rossi, 1985). Accordingly,
OFM feeding on terminals is intuitively detri-
mental in trees trained to upright systems.
The importance of OFM as a fruit-feed-
ing pest varies with region; however, it is a
key fruit-feeding pest in most major peach
production areas. Although it was thought to
have originated in Asia, OFM is now cosmo-
politan. In the USA, OFM is the key direct
pest of peach in the western states, the mid-
Atlantic region, where it produces four or five
generations per year, and the upper Midwest
(Hogmire, 1995; Horton et al., 2005). In the
USA’s largest production area, California’s
Central Valley, it completes up to six genera-
tions per year and is a dominant pest (Pickel
et al., 2006). OFM’s status is less defined in the
south-eastern USA, particularly in the warmer
production areas of South Carolina, Georgia
and Alabama. Cochran et al. (1955) observed
that OFM was an occasionally injurious, but
erratic, pest that seldom merited chemical
control on varieties ripening before ‘Hiley’
(early July in South Carolina’s Ridge produc-
tion area). The senior author’s on-farm obser-
vations in Georgia and South Carolina support
Cochran’s characterization of OFM’s modest
pest status in the south-eastern USA.
469
Fig. 18.1. Oriental fruit moth injury
to peach terminal growth. (Courtesy
of the University of Georgia, Bugwood
Network.)
In production areas with diverse fruit-
attacking pest complexes, such as the south-
eastern and mid-Atlantic regions of the USA,
where plum curculio rivals or exceeds OFM
in importance, standard management practice
relies on broad-spectrum insecticides which
simultaneously control OFM, plum curculio
and fruit-feeding hemipterans (plant bugs,
stink bugs, leaf-footed bugs). In California,
and in processing peach production scattered
across eastern North America, pheromone
mating disruption is more often the manage-
ment option of choice for OFM.
In Europe, OFM was first reported in
1920 in Italy. Since then it has distributed itself
into all production areas, becoming one of
their most dangerous pests.
OFM is a polyphagous pest which in
recent years has become increasingly injuri-
ous to pear and apple, creating a new and
complex situation owing to the proximity of
fruit orchards of mixed species and the pos-
sibility of seasonal migration of the popula-
tions (Sciarretta and Trematerra, 2006).
OFM adults are 6–7 mm long, herring-
bone grey moths with dark bands that may be
seen across the wings when at rest (Fig. 18.2/
Plate 187). Adult OFM have a wingspan of
approximately 12 mm (Strand, 1999). Larvae
are pinkish-white caterpillars with three pairs
470 D.L. Horton et al.

of thoracic legs and fleshy abdominal prolegs. OFM larvae (Fig. 18.3/Plate 188) from cod-
OFM larvae have brown heads and a black anal ling moth larvae, which are otherwise quite
comb on the last body segment, which is a key similar in appearance but lacking anal plates
diagnostic characteristic for differentiating (Hogmire, 1995).

Fig. 18.2. Oriental fruit moth adult. (Courtesy of the University of California IPM Program.)

Fig. 18.3. Anal plate of oriental fruit

moth larva. (Courtesy of the University
of California IPM Program.)
 Insects and Mites
OFM overwinter as mature larvae in
silken hibernacula on the tree, in dried fruits,
on leaves or twigs beneath trees, or other pro-
tected areas such as field bins. Pupation takes
place in late winter. Adult emergence is fre-
quently initiated a few days before peaches
begin to bloom. Egg laying begins a few days
after adult emergence. Females lay up to 200
white, flattish eggs, most often on twigs or
leaves near the distal ends of shoots (Howitt,
1993). Depending on fruit phenology, newly
hatched OFM larvae burrow into either the
tender, succulent tissue at the distal end of
shoots or ripening fruits.
Newly hatched, neonate OFM larvae are
known for quickly tunnelling beneath the
surface and into the fruit (Fig. 18.4/Plate 189).
OFM injury to harvested peaches typically
falls into two classes (Howitt, 1993). ‘Old
injury’ occurs when larvae move into green
fruit after abandoning twigs, leaves or tight
places where fruit touch. As fruit matures,
gum often extrudes from the entrance
wounds; this exudate darkens with time and
may be seen as a blackish blotch at harvest.
Larval entrances for many of these green fruit
infestations are through the sides of fruit,
Fig. 18.4. Oriental fruit moth larva tunnelling in peach. (Courtesy of the University of California IPM
Program.)
471
which makes them more visible than infesta-
tions to more mature fruit. ‘New injury’ pre-
dominates as the season progresses and fruit
ripens. Larval infestations in maturing fruit
are often difficult to discern. Many are associ-
ated with the stem cavity, leaving very mod-
est frass accumulation adjacent to the stem.
OFM larvae also enter the peach through the
stem, leaving no visible entrance wounds and
very little indication of infestation until
mature larvae exit the fruit (Hogmire, 1995).
As the season progresses into mid- and
late-season cultivars, OFM females will
increasingly oviposit directly on fruits, pre-
ferring fruit that are at least halfway through
their development (Myers et al., 2006). Once
inside the fruit, OFM larvae may wander
about or confine themselves to small areas.
Very often they do much of their injury near
the pit, where they leave considerable
amounts of sawdust-like frass. These accu-
mulations of frass differentiate OFM injury
from that of plum curculio, which leave
cleaner cavities with far less accumulation of
frass (Howitt, 1993). In summer OFM pupate
on the tree itself, while overwintering pupae
more often form hibernacula in cracks of tree
472 D.L. Horton et al.
bark, on refuse or weeds on the orchard floor,
or in degraded fruits (Hogmire, 1995).
Efficient OFM management requires
timely initiation of controls. Pest management
efforts for OFM begin with pheromone moni-
toring of adult male OFM. Capture patterns
indicate generational start-up dates, known
as biofixes, which trigger accumulation of
heat units in the OFM degree-day (°D) devel-
opmental model. These tactics accurately pre-
dict the timing of various life stages (Croft
et al., 1980).
Resistance of OFM to conventional insec-
ticides – organophosphates, carbamates and
pyrethroids – has been observed in at least
two North American production areas
(Ontario and New Jersey) (Kanga et al., 1999;
Shearer and Usmani, 2001). A 4-year study by
Kanga et al. (2003) suggests that resistance is
at least somewhat unstable, and that rotation
of insecticides may reduce and stabilize resis-
tance levels. Because the OFM neonate is the
primary surface-feeding life stage, the win-
dow for insecticidal control of larvae is quite
narrow. Effective insecticide-based OFM pro-
grammes rely on precise timing of applica-
tions to optimize control, while alternating
classes of chemistry to moderate selective
pressure for pest resistance.
Pheromone mating disruption is an
adaptation of traditional pheromone use that
saturates the orchard atmosphere with phero-
mone at levels that confuse, delay and impede
the ability of male moths to locate receptive
females. Mating disruption is non-toxic and
species-specific, which conveys obvious
advantages for integrated pest management
(IPM). However, the narrow spectrum of
activity for mating disruption requires the
use of additional controls to address other
insect pests, which adds to the cost of pest
control. Most often, pheromone mating dis-
ruption is used to control key pest species in
systems with relatively narrow pest com-
plexes or pesticide resistance issues. Though
utilization of pheromone mating disruption
varies considerably by pest species and
region, disruption of OFM typically performs
well. Mating disruption is commonly the
OFM management tool of choice in regions
where the technique is economically competi-
tive. For example, OFM mating disruption
has been widely used with good success on
stone fruit in California (Pickel et al., 2006),
Australia (Il’ichev et al., 2006) and Europe
(Rouzet et al., 1995; Cravedi and Molinari,
1996; Cravedi et al., 2001).
Codling moth, Cydia pomonella Linnaeus
(Lepidoptera: Tortricidae)
Codling moth, Cydia pomonella L. (CM), is
widely distributed as a pest of temperate-
zone fruit production areas worldwide,
including all of the USA, southern Canada,
Europe, south-eastern Australia, New Zealand,
the southernmost and northernmost limits of
Africa, the western and eastern portions of
Asia, as well as the South American countries
Argentina, Bolivia, Brazil, Chile, Colombia,
Peru and Uruguay.
CM is a serious, direct pest worldwide
on pome fruits, primarily apples and pears
(Epstein et al., 2006). The importance of CM as
a pest of peaches varies by region. In Califor-
nia, CM is an occasional, but serious, pest of
peach (Neven et al., 2006). CM infestations of
peaches in California are often associated
with pheromone mating disruption for CM in
adjacent pome fruits or walnuts (Strand,
1999). CM is also of import as a quarantine
pest restricting export of American-grown
pome fruit, walnuts and peaches (Neven
et al., 2006). In eastern North America, CM is
seldom seen as a peach pest; in Europe CM is
of little consequence as a peach pest.
CM overwinters as mature larvae in
debris on the orchard floor or under bark on
trees, and pupates there in early spring.
Adults are mottled grey moths with brown-
ish flecking and tented wings (Fig. 18.5/Plate
190). They emerge, mate and oviposit on fruit
and adjacent leaves, often preferring to
remain on the host species in which they
developed as larvae (Myers et al., 2006). Upon
hatching, first instar CM larvae tunnel into
available stages of fruit development, leaving
characteristic piles of frass at the entry site.
Three generations per year are common in
Central Valley locations in California, with a
possible fourth generation when very mild
winters are followed by temperate springs
(Strand, 1999).
 Insects and Mites
Fig. 18.5. Adult codling moth. (Courtesy of the University of California IPM Program.)
Pheromone monitoring and use of
degree-day developmental models are very
important tools for optimizing the timing of
IPM interventions. In Californian peaches,
large-scale monitoring efforts for CM are nor-
mally reserved for orchards with a history of
CM injury (Strand, 1999). These programmes
use the CM sex pheromone ((E,E)-8,10-dodec-
adien-1-ol) (Epstein et al., 2006) and degree-
day accumulations to optimize the timing of
IPM responses. In CM-prone peach orchards,
it is recommended that a single insecticide
application be applied at 250–300°D or 400–
500°D, depending on whether light or heavy
damage is expected (Strand, 1999).
Pheromone mating disruption is a very
reliable technique when used for managing
OFM in arid peach production areas with
relatively narrow complexes of direct pests.
However, mating disruption of CM is a more
challenging and problematic proposition. The
technique is seldom viewed as a reliable
stand-alone technology when employed
against CM, especially where CM abundance
is high (Knight, 2000).
CM control is primarily insecticide-based.
However, in apples, insecticide resistance and
473
greater inherent challenges with pheromone
mating disruption frequently require use of
multiple IPM tactics to ensure control and
slow the development of resistance. Multiple
insecticide classes, organophosphates, IGRs
and CM granulosis virus, as well as phero-
mone mating disruption, must sometimes
be used in concert to manage resistant CM
populations.
Peach twig borer, Anarsia lineatella (Zeller)
(Lepidoptera: Gelechiidae)
Peach twig borer, Anarsia lineatella (Zeller), is
a pest of peaches and other stone fruit, par-
ticularly almond (Hathaway et al., 1985) and
apricot. Peach twig borer is present in tem-
perate regions worldwide, occurring through-
out Europe, coastal North Africa, Israel,
Lebanon, Pakistan, Persia, Syria, Turkey, Iraq
and China. It is common throughout the USA
and southern British Columbia (Schlamp
et al., 2006). Peach twig borer is a frequently
encountered pest of almonds, peaches and
most other stone fruits in the western USA
474 D.L. Horton et al.
(Strand, 1999). Although widely distributed
across North America, peach twig borer is sel-
dom injurious to peaches in production areas
east of the Rocky Mountains (Horton et al.,
2005). In Europe, peach twig borer is widely
distributed, and its importance as a pest of
peach and apricot is thought to be on the rise.
In areas where peach twig borer is a
perennial peach pest, its management focuses
on prevention of early-season vegetative
injury, which minimizes subsequent risk of
direct injury to ripening fruit.
Peach twig borer adults are small, mot-
tled grey moths with distinct snout-like pro-
jections from the head (Fig. 18.6/Plate 191).
Larvae are initially white with black heads
and brown rings. Mature larvae are distinctly
ringed by brown body segments separated by
whitish intersegmental membranes, which
readily distinguish them from other caterpil-
lars feeding on stone fruit (Pickel et al., 2006)
(Fig. 18.7/Plate 192).
Peach twig borer overwinters as a first or
second instar larva within a silken chamber
or hibernaculum, which is usually tucked
into the crotches of 2- or 3-year-old wood,
pruning wounds, etc. Chimney-like mounds
of frass are often seen at the entrance of the
hibernaculum, although winter rains can
easily wash the frass away. Peach twig borer
larvae normally emerge from their hibernac-
ula as peaches come into bloom. Larvae feed
initially on opening vegetative and flower
buds. As terminal growth reaches sufficient
size, peach twig borer larvae tunnel into ter-
minals. A single larva may injure several
stems. As with vegetative feeding by OFM
larvae, the peach twig borer’s indirect injury
to terminal growth is most injurious in tree
training systems that emphasize upright-vase
or V-tree forms, which feature unbranched
scaffold limbs. Later in the season, as fruit
begin to colour, peach twig borer larvae feed
shallowly on the fruit, which results in off-
grade or cull fruit (Strand, 1999).
Peach twig borer is an important, but eas-
ily managed, pest in California. Preventive
controls are commonly used prior to fruit set.
Dormant or delayed dormant applications of
oil plus an organophosphate insecticide will
provide very reliable control of peach twig
borer. Oil alone does not control peach twig
borer. Alternatively, two bloom sprays of the
entomopathogenic bacterium Bacillus thuringi-
ensis also provide very reliable control of peach
twig borer. While B. thuringiensis treatments
lack efficacy against scale or mites, they pre-
serve natural enemies and eliminate the envi-
ronmental risks of dormant organophosphate
applications (Pickel et al., 2006).
In-season management options for peach
twig borer are based on observation of shoot
strikes and use of pheromone trapping to ini-
tiate accumulation of degree-day heat units.
Fig. 18.6. Adult of the peach twig
borer moth. (Courtesy of the Catholic
University, Piacenza, Italy.)
 Insects and Mites
Fig. 18.7. Mature peach twig borer larva. (Courtesy of the University of California IPM Program.)
An average of three to four shoot strikes per
tree is sufficient to warrant insecticide appli-
cation. Each peach twig borer generation
requires about 950–1080°D using a lower
threshold of 10°C and upper threshold of
31°C. Shoot strike monitoring should begin as
degree-day accumulations approach 400. As-
needed insecticides are applied at 400–500°D.
Twice-weekly monitoring for the biofixes of
each subsequent generation should begin as
degree-days reach 900. Treatments to protect
ripening fruit are more commonly required
late in the season (Pickel et al., 2006). Phero-
mone mating disruption of peach twig borer
has also worked effectively in California
(Pickel et al., 2006). Some European research-
ers have observed an inconsistent attractive-
ness of the peach twig borer pheromone,
which results in monitoring that is less reli-
able than for OFM. Cases of mating disrup-
tion failure have been reported for pheromone
dispensers that combine peach twig borer
and OFM pheromones, with damage pro-
duced mainly by peach twig borer. Research
is ongoing on the complex of natural enemies
affecting peach twig borer (Molinari et al.,
475
2005a), as well as on the development of fore-
casting models (Molinari et al., 2005b).
Tufted apple bud moth, Platynota idaeusalis
(Walker) (Lepidoptera: Tortricidae)
Tufted apple bud moth, Platynota idaeusalis
(Walker) (TABM), is a bivoltine pest of both
peach (Hogmire, 1995) and apple (Myers and
Hull, 2003) in the eastern USA (Fig. 18.8/Plate
193). On peach, larval feeding produces a suc-
cession of injury types as the maturing larvae
feed on fruit surfaces, gumming and later
feeding internally (Fig. 18.9/Plate 194). As
areas of damage develop, rolled leaves and
leaf shelters may be seen over protected areas
where larvae are actively feeding (Hogmire,
1995).
The second generation, as with many
other fruit-feeding lepidopteran pests, is the
most damaging, owing to the potential for
greater pest abundance and the presence of
maturing fruit. Mature fruit wounded by split
pits, cold injury or TABM infestation are
476 D.L. Horton et al.

Fig. 18.8. Tufted bud moth adult. (Courtesy of G. Krawczyk, University Park, Pennsylvania, USA.)

Fig. 18.9. Tufted bud moth larva on fruit. (Courtesy of G. Krawczyk, University Park, Pennsylvania,
USA.)

more susceptible to brown rot and other fun- Control measures for TABM have relied
gal diseases. Such fruits may serve as a source heavily on organophosphate pesticides, but
inoculum for fungal diseases in the orchard the development of resistance has forced
(Hogmire, 1995). growers to use pyrethroids and carbamates,
 Insects and Mites
to which the moths have also developed resis-
tance in Pennsylvania, USA. First tebufenoz-
ide and then methoxyfenozide, a growth
regulator that induces moulting, were regis-
tered in some measure as a response to TABM
resistance and concerns over the susceptibil-
ity of coccinellid predators to pyrethroids
(Biddinger et al., 2006). The application of
insecticides for TABM controls early larval
instars. The TABM degree-day model devel-
oped for use in apples has been used as a
decision-making aid in peaches as well.
Plum curculio, Conotrachelus nenuphar
(Herbst) (Coleoptera: Curculionidae)
Plum curculio, Conotrachelus nenuphar (Herbst)
(PC), is a key fruit-feeding pest of North
American peach (Quaintance and Jenne, 1912)
and apple (Racette et al., 1992) production
areas east of the Rocky Mountains. This wee-
vil is found across the eastern half of North
477
America, with small populations occurring as
far west as Montana and Utah. Particularly in
the south-eastern USA, where PC is multivol-
tine (multiple generations per year), it is the
key fruit-feeding pest of peaches (Horton
et al., 2005). PC is a native pest of indigenous
plum and crabapple species that has adapted
well to peach, nectarine and plum (Maier,
1990; Jenkins et al., 2006).
PC adults are brownish-grey weevils
with four pairs of bumps on the wing covers.
They are approximately 5 mm long (Quain-
tance and Jenne, 1912) (Fig. 18.10/Plate 195).
North of Virginia, most populations exhibit
an obligate diapause and thus are univoltine
(one generation per year). However, in the
south-east, a facultative diapause produces a
bivoltine cycle (two generations per year)
(Padula and Smith, 1971). Overwintering adults
emerge from orchard debris and ground cover
to feed on buds, flowers and developing fruits
(Hogmire, 1995). PC produce feeding injury
which is seen as round, puncture wounds and
Fig. 18.10. Plum curculio adult. (Courtesy
of the University of Georgia, Bugwood
Network.)
478 D.L. Horton et al.
D-shaped oviposition wounds (Fig. 18.11/
Plate 196). In young peaches, the abundant
fuzz makes attacks by overwintered adults
difficult to discern. Successful oviposition
produces yellowish-white, legless larvae that
tunnel through fruit and feed near the pit
(Hogmire, 1995). Larval infestations from
overwintered adults normally cause peach
fruitlets to abort, while field or second-gener-
ation larvae produce wormy, unmarketable
fruit (Quaintance and Jenne, 1912).
Throughout eastern North America, PC
utilizes a range of native fruits, primarily
rosaceous and ericaceous species (Quaintance
and Jenne, 1912; Mampe and Neunzig, 1967).
Maier (1990) found that 19 species of native
and exotic (primarily cultivated) rosaceous
hosts were acceptable PC hosts in Connecti-
cut. In that region of eastern North America,
PC is univoltine, favouring Crataegus spp. as
hosts, while readily feeding on apples. Fur-
ther south in Georgia, where PC is multivolt-
ine, the Chickasaw plum (Prunus angustifolia)
is a preferred early host, while peach has
become a very significant host for PC’s field
generation (Jenkins et al., 2006).
The abundance of native PC hosts in
adjacent woodland and roadside habitats
across much of eastern North America con-
founds management of PC. Especially in the
south-eastern USA, where PC from wild hosts
move to peach during their field generation
(Jenkins et al., 2006), the prolonged threat of
PC attack is a major concern which forces
employment of very conservative manage-
ment strategies.
Monitoring options for PC remain rudi-
mentary. Leskey and Wright’s (2004) work on
apples in the north-eastern USA showed that
traps baited with the aggregation pheromone
(grandisoic acid and benzaldehyde) are not
reliable enough to direct as-needed insecticide
Fig. 18.11. Plum curculio oviposition
wound. (Courtesy of J.A. Payne,
University of Georgia, Bugwood
Network.)
 Insects and Mites
applications for PC in commercial orchards.
Subsequent work suggests that more competi-
tive attractants will likely require complex,
multiple-component blends of tree volatiles
(Leskey et al., 2005).
Owing to the lack of commercially reli-
able sampling technologies to direct as-needed
insecticide application, PC control is insecti-
cide-based and preventive. In production areas
where PC is univoltine, early-season preven-
tive insecticide applications, applied in part
for OFM, plant bugs and scale crawlers, pro-
vide very reliable control of PC. In the south-
eastern USA, insecticide applications for PC
are initiated at petal fall. The absence of reli-
able sampling technologies forces most grow-
ers, certainly those who are wholesale shippers
of peaches, to spray preventively for PC
season-long, normally on a 14-day schedule
(Horton et al., 2007).
Prevailing market standards for control of
internal fruit-feeding insects are stringent, and
in wholesale shipping are market-driven. In
essence, fruit must be totally free of internal
fruit feeders such as OFM or PC. This rigorous
market standard has perpetuated highly reli-
able but risk-aversive and insecticide-intensive
control programmes. In the eastern USA, the
organophosphate insecticide phosmet and, to
a lesser degree, high-rate applications of pyre-
throids form the basis of control for PC and
other fruit-attacking pests. Alternative, reduced-
risk insecticides, the application of entomo-
pathogenic nematodes (Shapiro-Ilan et al., 2004),
the use of cellulose sheeting to limit overwin-
tering populations (Benoit et al., 2006) and the
application of finely ground kaolin particle
films (Lalancette et al., 2005) have all shown
promise, but are not in wide commercial utili-
zation for PC control.
Scarab beetles (Coleoptera: Scarabaeidae)
Scarab beetles are large, elongate to oval,
heavy-bodied, convex beetles typically bear-
ing five-segmented tarsi and eight- to 11-
segmented, lamellate antennae. Scarabs are
plant feeders (Triplehorn and Johnson, 2005b)
that are primarily direct fruit feeders in peach,
although Japanese beetles are also foliage
479
feeders. Scarab larvae are soil-dwelling, most
often feeding on organic matter or roots. They
are white, C-shaped grubs. Green June beetles
(Subfamily Cetoniinae), rose chafers (Mycro-
dactylus subspinosus (Fabricius)) (Subfamily
Melonlonthinae) and Japanese beetles (Sub-
family Rutelinae) are the common scarab
pests of peaches in eastern North America.
Japanese beetle, Popillia japonica Newman
Japanese beetle, Popillia japonica (Newman)
(JB), is known for its exceptionally broad host
range, which encompasses more than 275
plant species. On peach, JB feeds on foliage,
producing a characteristic lace-like defolia-
tion, and as fruit matures, JB feeds heavily on
fruit (Hogmire, 1995). JB is native to Japan,
where natural enemies typically keep its pop-
ulations in check. The life history of JB is typi-
cal of many white beetle grub species that
feed on the roots of grass. Eggs laid in the soil
in summer produce larvae (white grubs)
which may be diagnosed by their raster pat-
tern, i.e. a series of V-shaped bristles found at
the tip of the abdomen on the underside. The
larvae overwinter in the soil, pupate and
emerge as adults from mid-May to July (Shet-
lar and Johnson, 2005). The adults are charac-
terized by the presence of five white tufts of
hairs on each side of their copper-coloured
wing covers (Shetlar and Johnson, 2005) (Fig.
18.12/Plate 197).
JB is largely univoltine, and the presence
of large, synchronized, adult populations in
summer can pose significant problems for
growers. Adults are long-lived; they feed
voraciously throughout their 30–45-day
lifespan (Hogmire, 1995). Adult beetles are
attracted both to the aggregation pheromone
produced by feeding adults (Shetlar and
Johnson, 2005) and by olfactory cues and kai-
romonal signals emitted from damaged fruit
(Loughrin et al., 1995). Furthermore, the emis-
sion of sex pheromone by females may attract
more beetles to damaged fruit and increase
the localized concentrations of beetles seen in
orchards (Potter and Held, 2002). Orchards
can easily be monitored by visual observa-
tion, or trapping with a combination of the
sex pheromone (japonilure) and fruit volatiles
is also feasible (Potter and Held, 2002).
480 D.L. Horton et al.
Control of JB is based on as-needed use
of insecticides. Especially in orchards near
pastures, controls should be implemented
before significant defoliation occurs and cer-
tainly before JB begins to attack fruit. In North
America, attempts at classical biological con-
trol have been made by importation of natu-
ral enemies (Jackson and Klein, 2006). Despite
successful establishment of some parasitoid
species, suppression or control of JB is
sporadic.
JB is a quarantine pest in Europe. Although
a single occurrence of JB in the Portuguese
Azores Islands (Atlantic Ocean) has been
observed, no further observations of this
serious scarabeid pest have been reported in
Europe (Tremblay, 2000).
Green June beetle, Cotinis nitida (Linnaeus)
The green June beetle, Cotinis nitida (Lin-
naeus) (GJB), is native to much of the eastern
half of the USA. GJB is seasonally abundant
in peaches, especially in the south-eastern
USA. It can be a very important fruit-feeding
pest on late varieties (Flanders and Johnson,
2005). GJB is a large, metallic green- and
bronze-coloured scarab beetle (Fig. 18.13/
Plate 198) which emerges and often moves to
Fig. 18.12. Japanese beetle adult.
(Courtesy of the University of Georgia,
Bugwood Network.)
ripening peaches in late June to July (Flanders
and Johnson, 2005). Females attract males
with sex pheromone, mate and lay their eggs
in tightly packed masses in moist soils, pre-
ferring those rich with organic matter (Brand-
horst-Hubbard et al., 2001). Each female
continues a cycle of feeding, mating and egg
laying that lasts approximately 2 weeks
(Brandhorst-Hubbard et al., 2001). Regular
orchard monitoring, which may be aug-
mented by the use of fermenting fruit trays to
detect adults, readily identifies developing
GJB infestations. Initiation of fruit feeding
very often warrants application of insecti-
cides with short preharvest intervals (Flan-
ders and Johnson, 2005).
Plant bugs (Hemiptera: Miridae), stink bugs
(Hemiptera: Pentatomidae) and leaf-footed
bugs (Hemiptera: Coreidae)
Plant bugs (Hemiptera: Miridae), stink bugs
(Hemiptera: Pentatomidae) and leaf-footed
bugs (Hemiptera: Coreidae) are fruit-feeding
pests of peaches and nectarines. As a group,
they are polyphagous; most feed successively
through the growing season on flower buds,
 Insects and Mites
blooms, fruitlets and mature fruits of numer-
ous broadleaved hosts. Many of their pre-
ferred hosts are herbaceous. They have
stylet-like mouthparts that inject saliva which
breaks down host tissues, allowing the bugs
to suck in the liquefied cellular contents.
Plant bugs are small, soft-bodied true bugs.
The antennae and beak are four-segmented
(Triplehorn and Johnson, 2005a). The most
important plant bug pest of peach in eastern
North America is the tarnished plant bug,
Lygus lineolarius (Palisot de Beauvois) (TPB)
(Horton et al., 2005), which is a variable, mot-
tled brass-brown colour with a red thorax
(Sanderson and Peairs, 1921) (Fig. 18.14/Plate
199). In California, Lygus hesperus (Knight)
and Lygus elisus (Van Duzee) are the key plant
bug pests of stone fruit (Strand, 1999). L. hes-
perus adults are yellowish to reddish-brown,
while L. elisus adults are a pale green. In
Europe, damage has been reported for Lygus
rugulipennis (Poppius) (Tavella et al., 1996). In
peach, TPB are most abundant from bloom to
shuck off, although they may be injurious
through harvest. Rings (1958) thoroughly
characterized TPB injury. He showed that
TPB feeding typically aborted peach blos-
soms and small fruitlets from bloom to shuck
off. TPB-induced fruit deformation or scarring,
481
Fig. 18.13. Green June beetle adult.
(Courtesy of the University of Georgia,
Bugwood Network.)
often called ‘cat-facing’, typically occurred
from shuck off to 35 days after bloom. Bluish-
grey scarring and shallow pitting of the fruit
predominated from 35 days post-bloom to
harvest. TPB also produced gumming and
water soaking, both of which were more com-
mon later in the season.
Stink bugs, and to a lesser degree leaf-
footed bugs, feed on green and ripening
peaches. Stink bugs, often green or brown
(Fig. 18.15/Plate 200) in colour, are large,
round to oval, true bugs with five-segmented
antennae. As the name implies, many stink
bugs have an unpleasant odour. Leaf-footed
bugs are elongate, often brown, with the head
narrower and shorter than the first thoracic
segment, and expanded, leaf-like hind tibiae
(Triplehorn and Johnson, 2005a) (Fig. 18.16/
Plate 201). The brown stink bug (Euschistus
servus (Say)), the dusky stink bug (Euschistus
tristigmus (Say)), the southern green stink bug
(Nezara viridula (Linnaeus)), the green stink
bug (Acrosternum hilare (Say)) and Thyanta spp.
are peach pests in the eastern USA (Johnson
et al., 2005). In the arid peach production areas
of the western USA, Euschistus conspersus
(Uhler), A. hilare and Thyanta pallidovirens (Stål)
are significant fruit feeders (Strand, 1999).
The leaf-footed bug, Leptoglosus phyllopus
482 D.L. Horton et al.
(Linnaeus), is also a pest of peaches in the
south-eastern USA (Johnson et al., 2005).
Stink bug injury to peaches parallels the
injuries produced by plant bugs. Rings (1957)
showed that fruit phenology at the time of
feeding dramatically influenced the type of
injury produced. Early-season feeding, from
bloom to fruitlets of 13 mm diameter, pro-
duced fruit abortion, ‘cat-facing’ or fruit
Fig. 18.14. Tarnished plant bug adult.
(Courtesy of S. Bauer, USDA.)
Fig. 18.15. Brown stink bug adult.
(Courtesy of J. Greene, Clemson
University, South Carolina, USA.)
deformation (Fig. 18.17/Plate 202) and nor-
mally took place from bloom to 49 days post-
bloom. Scarring, characterized by brown,
corky scars, and some loss of fuzz in the
affected areas took place 42–56 days after
bloom. Gumming, the most common injury,
took place from 42 to 115 days after bloom.
Water-soaked injury also developed during
the same time interval. All common stink
 Insects and Mites 483

Fig. 18.16. Leaf-footed bug adult.

(Courtesy of D. Cappaert, University of
Georgia, Bugwood Network.)

Fig. 18.17. Cat-facing injury on peach. (Courtesy of University of Georgia, Bugwood Network.)

bug species produce similar injury to
peaches.
The hemipteran pests of peaches are highly
mobile and are difficult to monitor (Legrand
and Los, 2003). Close visual monitoring for
bugs and injury, emphasizing the vulnerable
orchard edges, should be integral in any
orchard monitoring plan (Johnson et al., 2005).
Bug populations often build up on herbaceous
hosts, particularly when moisture is abundant.
Movement to peaches, a deep-rooted perennial,
is most evident when herbaceous hosts dry
down and senesce. Trap-based monitoring
techniques for plant bugs and stink bugs are
available and, in some peach production areas,
trapping is utilized as an adjunct to observa-
tion. Pink and, to a lesser extent, white sticky
traps are used to monitor for TPB (Legrand
484 D.L. Horton et al.
and Los, 2003). Yellow pyramid traps baited with
the Euschistus spp. (Hemiptera: Pentatomidae)
aggregation pheromone have shown promise
for monitoring stink bugs in pecans (Cottrell,
2001) and apples (Hogmire and Leskey, 2006).
However, additional work defining the rela-
tionship between injury and capture remains
to be done before this system can be fully
incorporated as a commercial IPM decision-
making tool (Hogmire and Leskey, 2006).
Cultural control, in the form of selective
use of herbicides, is a basic tenet of managing
the plant bug and stink bug pests of peaches.
Orchard floor management does not elimi-
nate the need for insecticides, but it consis-
tently reduces pest pressure, thus improving
the performance of insecticide applications
and occasionally diminishing the need for
sprays (Horton et al., 2005). The prescribed
orchard floor management practices mini-
mize in-orchard populations of late-winter
and spring broadleaved annual weeds.
Removal of these preferred hosts renders the
orchard less attractive to heteropteran flower
and fruit feeders, consistently reducing injury
(Killian and Meyer, 1984; Atanassov et al.,
2002).
Thrips (Thysanoptera)
Thrips are occasional, but sometimes quite
injurious, pests of peach. Thrips are typically
more injurious to stone fruit grown in arid
production areas. However, even in produc-
tion areas with abundant rainfall, thrips can
be problematic, especially when dry weather
prevails just before and during bloom. Thrips
are typically more important as pests of nec-
tarines. Western flower thrips, Frankliniella
occidentalis (Pergande), is a well-documented
pest of nectarine and, to a less significant
extent, peaches in California (Weldon, 1921).
In the south-eastern USA thrips are less com-
monly pests of peach, but when abundant,
western flower thrips are injurious (Yonce
et al., 1990). Other western flower thrips,
Thrips meridionalis (Priesner) and Thrips major
(Uzel), are pests of nectarines and peach in
various Italian peach-producing regions
(Cravedi et al., 2001).
Adult thrips are slender, cigar-shaped
and quite small; western flower thrips, one of
the key pest species of peach, is about 1 mm
in length (Fig. 18.18/Plate 203). Adult thrips
have long, narrow, fringed wings which are
folded along the back when at rest. The
mouthparts of thrips scar plant tissues by
piercing the surface, releasing cellular fluids
which are then consumed. Oviposition injury
occurs when females insert their ovipositors
into tender flower parts.
Thrips produce two distinct types of
injury to stone fruit (Weldon, 1921). Russet-
ing, the more severe injury, is seen as a rough-
ened, corky brownish scar on the surface of
fruit (Fig. 18.19/Plate 204). Russeting is pro-
duced early in the season when flower thrips,
which overwinter on weed species, move to
peach as bloom is initiated. Adult thrips may
enter flower buds soon after the buds begin to
swell significantly. Adults feed inside buds
and in open blooms; females further injure
the nascent fruit by inserting their eggs into
succulent flower parts. Feeding by the resul-
tant nymphs adds their injury to that of the
adults. Silvering occurs as maturing fruit
begin to colour. Injury at this time produces
numerous small, clear blemishes where the
pigment has been evacuated with the cellular
contents. In hand, silvering is seen as a silver
sheen (Fig. 18.20/Plate 205). Light to moder-
ate silvering is sometimes ignored, especially
when demand for fruit is high.
Fresh market peaches should be moni-
tored by shaking or flailing flowers over a
cigar box or a stiff piece of paper. Unopened
blooms can be sampled by removing soon-to-
open buds and placing them into small, sealed
plastic bags. Adult and immature thrips will
leave the buds if the bags are allowed to warm
on a window sill, which drives them out of
hiding.
Formal treatment thresholds for flower
thrips do not exist in peaches. High thrips
numbers are required to damage peach. How-
ever, pest management practitioners must
consider the limited availability of insecti-
cides for thrips control and the difficulty of
obtaining spray coverage thorough enough
to reach thrips inside of soon-to-open flowers.
Cultural control in the form of orchard floor
management to minimize the abundance of
 Insects and Mites 485

Fig. 18.18. Western flower thrips on peach leaf. (Courtesy of J.A. Payne, University of Georgia,
Bugwood Network.)

Fig. 18.19. Western flower thrips russeting on fruit. (Courtesy of J.A. Payne, University of Georgia,
Bugwood Network.)

weed hosts in and adjacent to the orchard has tioned to avoid cutting lucerne as nearby
been shown to be of value (Cravedi et al., peaches mature to minimize movement from
2001). Similarly, California growers are cau- lucerne to peach (Strand, 1999).
486 D.L. Horton et al.
Mediterranean fruit fly, Ceratitis capitata
(Wiedemann) (Diptera: Tephritidae)
Mediterranean fruit fly, Ceratitis capitata (Wie-
demann), which is thought to have originated
in Africa, is a serious pest of numerous fruit
and vegetable species. It is well adapted to
diverse climates and has in excess of 350 host
fruit species (Liquido et al., 1991). Mediterra-
nean fruit fly is well established in Central
America, South Africa, Israel, much of the
Mediterranean and Australia (Peck and
McQuate, 2000). In general, it is found in most
tropical and subtropical areas of the world.
Numerous temporary populations have been
established in the continental USA but, to
date, eradication efforts have prevented the
establishment of permanent or adventive
populations. Mediterranean fruit fly was
introduced in the state of Hawaii, USA, in
1910, where it became established (Anon.,
2006).
The adult Mediterranean fruit fly is
slightly smaller than a house fly (Fig. 18.21,
Plate 206). It has dark blue eyes and a shiny
thorax of glistening black overlain with a
mosaic of yellowish white. The abdomen is
yellowish with two silvery cross bands. Its
wings, which are normally drooping, are
banded and blotched with yellow, brown and
black. As with other fruit fly species, the adult
diet is exclusively liquids. Females insert their
Fig. 18.20. Western flower thrips
silvering on fruit. (Courtesy of J.A.
Payne, University of Georgia,
Bugwood Network.)
eggs into the flesh of suitable fruits. Larvae
complete development inside fruit, at which
time the mature larvae exit fruit and move into
the soil to pupate. Under the most favourable
of conditions the life cycle may be completed
in as little as 17 days. In various parts of the
world Mediterranean fruit fly completes from
1 to 12 generations per year (Anon., 2006).
The economic impacts of fruit fly species
are twofold: (i) larval feeding directly injures
fruit, rendering it unmarketable; and (ii) fruit
fly infestations limit access to markets which
impose quarantine restrictions (Malavasi
et al., 1994). Management of Mediterranean
fruit fly in production areas with persistent
infestations varies according to the intensity
of infestation and the type of markets utilized
to sell fruit. In regions where export of pro-
duce to uninfested regions is not pursued, a
variety of traditional pest management
approaches are employed to mitigate losses.
Control relies primarily on broad-spectrum
insecticides applied in baits or as cover sprays
(Roessler, 1989). The sterile insect technique is
an alternative approach for suppressing or
eradicating fruit flies on an area-wide scale
(Hendrichs, 1996). Large numbers of males,
which have been irradiated to render them
sterile, are released to dramatically reduce the
probability of virgin females successfully
mating with viable males. Sterile male releases
are normally preceded by use of insecticidal
 Insects and Mites
Fig. 18.21. Mediterranean fruit fly mating. (Courtesy of Catholic University, Piacenza, Italy.)
baits and/or cover sprays to reduce the ambi-
ent population of flies. Use of insecticides to
ensure a relative paucity of viable males
increases the probability of females laying
infertile eggs.
Quarantine restrictions often prompt cre-
ation of area-wide management protocols
which prescribe aggressive, region-wide trap-
ping for adults. Trapping of even very low
numbers of fruit fly adults will trigger prompt
area-wide use of insecticidal baits, as-needed
foliar insecticide and, in some cases, release of
sterile male fruit flies.
18.3 Indirect Insect Pests
Indirect pests of peaches are the tree-attack-
ing pests, those which feed on the foliage,
bark, inner bark or the roots. They are indirect
in that their injury impacts the tree itself
before exacting costs from the fruit. Scale spe-
cies are key indirect pests of peaches wher-
ever they are grown. Borers of various sorts
are regionally important, as is the case with
the peach tree borer and lesser peach tree
borer in eastern North America. Aphids and
leafhoppers are frequently present in peaches,
487
although except for occasional stunting of
young trees, they are seldom truly injurious.
However, aphids are important vectors of
plant diseases, including Plum pox virus.
Management of indirect peach insect
pests should focus on tree health, productiv-
ity and orchard longevity. Profits are returned
to the grower from the time trees generate
sufficient income to pay for themselves until
the orchard’s eventual demise. Productivity
or longevity lost to borers or scale very
quickly eats into an orchard’s lifetime income
potential. Cultural and pest management
efforts should focus on maintaining tree health
and vigour to optimize economic returns.
Peachtree borers, Synanthedon spp.
(Lepidoptera: Sesiidae)
Peachtree borer, Synanthedon exitiosa (Say)
(PTB), and lesser peachtree borer, Synanthe-
don pictipes (Grote & Robinson) (LPTB), are
key tree-attacking pests of peach in eastern
North America. Native Prunus spp. are their
seminal hosts, but both PTB and LPTB have
adapted quite well to peaches. The larvae
(Fig. 18.22/Plate 207) of both species feed
488 D.L. Horton et al.
Fig. 18.22. Peachtree borer larva. (Courtesy of University of Georgia, Bugwood Network.)
beneath the bark on phloem tissue. As sub-
surface vascular feeders, they are initially dif-
ficult to diagnose and often do substantial
damage before infestations are evident. Both
species are quite injurious; if uncontrolled,
they debilitate fruiting wood, scaffold limbs
and entire trees, often leading to premature
orchard decline (Fig. 18.23/Plate 208).
PTB is generally univoltine; the white
larvae (Fig. 18.22/Plate 207) construct pupal
cases below ground, pupate and produce
adults that mate and oviposit on tree trunks
and exposed roots shortly after emergence.
After 10 days the neonates hatch, gain access
to cambial tissue through cracks or wounds
in the bark, and overwinter in various stages
of development. This sexually dimorphic diur-
nal PTB moth is also an effective wasp mimic;
its shining clear wings, metallic blue-black
body and, particularly in the males, superb
flight capabilities make it easily mistaken for
a wasp (Hogmire, 1995) (female – Fig. 18.24/
Plate 209; and male – Fig. 18.25/Plate 210).
LPTB is a similar species; however, it is
less sexually dimorphic: both males and females
are similar to PTB males, while lacking the
triangular shape of the terminal abdominal
segments and yellow scales between the
antennae. The LPTB lays its eggs on above-
ground structural wood. The larvae tunnel
through cambial tissues, producing a gum–
frass mixture that is highly diagnostic of
infestation (Fig. 18.26/Plate 211). Remnants
of previously eclosed pupae that extrude out
of the damaged, sticky bark are also diagnos-
tic. LPTB is bivoltine in the south-eastern USA
(Hogmire, 1995).
Control options for PTB and LPTB vary
with production area. Barrier insecticide
applications and pheromone mating disrup-
tion are the commercial standards for man-
agement of these pests. Both methods are
preventive in nature. Nematode-based con-
trol of PTB (Cossentine et al., 1990; Cottrell
and Shapiro-Ilan, 2006) and LPTB (Shapiro-
Ilan and Cottrell, 2006) has shown promise.
In eastern North America diverse fruit-
feeding pest complexes necessitate at least
some measure of repetitive, in-season cover
spray. Cover spray insecticide applications
are thought to diminish the levels of biologi-
cal control of PTB and LPTB which would
otherwise be provided by natural enemies.
However, history suggests that both barrier
insecticides and mating disruption may have
been augmented by cover spray suppression
 Insects and Mites 489

Fig. 18.23. Branches damaged by lesser peachtree borer. (Photo by J. Fuest, University of Georgia.)

Fig. 18.24. Adult female peachtree borer. (Photo by J. Fuest, University of Georgia.)
490
Fig. 18.26. Lesser peachtree borer wounding. (Photo by J. Fuest, University of Georgia.)
of the borer species. Organophosphate insec-
ticides have been the peach insecticides of
choice in eastern North America since the
mid-1950s. In the early 1990s, regulatory
mandate transitioned growers in the eastern
USA from encapsulated methyl parathion to
phosmet. Phosmet has performed quite well
as a fruit protectant. However, after a few
D.L. Horton et al.
Fig. 18.25. Adult male peachtree
borer. (Courtesy of University of
Georgia, Bugwood Network.)
years of widespread reliance on phosmet,
borer control in the south-eastern USA began
to deteriorate. LPTB, which is multivoltine,
and to a lesser degree PTB have asserted
themselves as aggressive, often debilitating
tree pests and led to premature decline and
removal of heavily infested orchards (Horton
et al., 2005).
 Insects and Mites
Chlorpyrifos, also an organophosphate,
is the most effective barrier insecticide treat-
ment for borers. As a protective barrier, it pro-
vides excellent control of PTB (Yonce, 1980),
and at least a measure of LPTB suppression.
Chlorpyrifos barrier sprays target neonate
and first instar larvae, with some measure of
ovicidal and adulticide activity. In much of
the eastern USA, chlorpyrifos barrier sprays
are the commercial standard. While targeted
primarily at PTB, barrier sprays at least sup-
press LPTB. Chlorpyrifos barrier insecticides
are applied either postharvest or a minimum
of 14 days preharvest. Application is by hand-
gun-directed coarse sprays to optimize cover-
age and, in the mid-Atlantic states, where
preharvest sprays are needed, to allow appli-
cators to scrupulously minimize residue
deposition on the fruit.
In eastern North America, pheromone
mating disruption of borer species in peaches
is in common use in the mid-Atlantic produc-
tion area, Ontario and Michigan. Acreages
using mating disruption in these areas are
now similar to those receiving the more con-
ventional barrier insecticide treatment with
chlorpyrifos. Rates required for efficacious
pheromone mating disruption of the borer
species in the south-eastern USA, where the
growing season is substantially longer than in
the above areas, are significantly higher and
hence more costly.
Armoured scale (Hemiptera: Diaspididae)
White peach scale, Pseudaulacaspis
pentagona (Targioni-Tozzetti)
White peach scale, Pseudaulacaspis pentagona
(Targioni-Tozzetti) (WPS), is an important
cosmopolitan pest of peaches. WPS has an
extremely wide host range, including nearly
all non-coniferous plants (Nalepa and Meyer,
1990). Although native to south-eastern Asia,
WPS has become a pest with a worldwide
distribution (Erkiliç and Uygun, 1997). Since
its introduction to the USA in the late 1800s,
WPS has been most notable as a peach pest in
the south-eastern states (Van Duyn and
Murphey, 1971; Yonce and Jacklin, 1974). The
range of WPS in the eastern USA extends west
491
to Texas and northwards to Maryland and
Tennessee (Hanks and Denno, 1993). This
insect also causes economic damage in Turkey,
where it is a dominant pest of peach (Erkiliç
and Uygun, 1997), Italy (Pedata et al., 1995),
Asia, Australia, Africa, the Caribbean and the
Pacific Islands (Hanks and Denno, 1993).
Female WPS are creamy white to red-
dish-orange, sac-like, sessile insects that are
covered in waxy, brownish coverings approx-
imately 1 mm in diameter (Hodges, 2005)
(Fig. 18.27/Plate 212). Female WPS overwin-
ter beneath their scale covers, becoming active
pre-bloom. Adult females emit sex phero-
mone in a rhythm that matches the daily egg
hatching cycles of males, which are, in turn,
controlled by light and temperature (McLaugh-
lin et al., 1990). The tiny, yellowish, winged
males are attracted to calling females and can
inseminate multiple females in each aggrega-
tion (McLaughlin et al., 1990). Crawlers, or
nymphs, disperse from beneath the females
to seek unoccupied feeding sites on the tree
bark (Hodges, 2005). In as little as a week
crawlers have begun to settle, at which time
they become sessile and lay down scale covers.
WPS infestations of peach result in loss
of tree vigour, with reduced fruit production
and ultimately death of fruiting wood, scaf-
folds and trees. In the absence of management
steps specifically for scale, heavy WPS infes-
tations can kill portions of a tree or even the
entire tree in as few as 2 or 3 years (Kuitert,
1967). WPS is a multivoltine pest, having
three generations in central Virginia (Bobb
et al., 1973) and four generations in north Flor-
ida, where it is a major pest of peaches (Kuit-
ert, 1967).
In Europe, WPS is present in that conti-
nent’s central and southern production areas,
where two or three generations per year can
develop (Kozar et al., 1997). In addition to
peach and cherry, WPS is an important pest of
kiwifruit (Paloukis and Navrozidis, 1997).
In the USA, WPS is most injurious in the
south-east’s warmer, coastal plain production
areas. Peaches are quite susceptible to injury
from WPS. Kuitert (1967) observed that
peaches were severely debilitated by WPS
infestations when visually similar infestations
did not appear to be detrimental to chinaberry,
privet, black walnut or mulberry. WPS is,
492 D.L. Horton et al.
however, widely noted as a pest of mulberry
(Hanks and Denno, 1993).
Control of WPS in the south-eastern USA
is reliably obtained by annual application of
two dormant-season horticultural oil sprays
(Bobb et al., 1973). Control of WPS via cover
sprays can occur if applications coincide with
crawler emergence. However, determination
of the optimal timing for spray applications is
difficult to obtain and, thus, good WPS con-
trol via cover sprays seldom occurs. Biologi-
cal control of WPS, primarily from parasitoids
such as Encarsia berlesei (Howard), can be quite
effective in the south-eastern USA; unfortu-
nately natural control is severely impeded by
the necessary in-season insecticide applica-
tions to control the region’s diverse complexes
of fruit-feeding pests (Nalepa and Meyer,
1990). Insecticides applied to control other
peach pests have been shown to exacerbate
WPS infestation levels by reducing parasitoid
abundance in Italy (Pedata et al., 1995), Tur-
key (Erkiliç and Uygun, 1997) and Florida
(Collins and Whitcomb, 1975).
San Jose scale, Quadraspidiotus
perniciosus (Comstock)
San Jose scale, Quadraspidiotus perniciosus (Com-
stock), is a cosmopolitan species which is a
Fig. 18.27. White peach scale: adult
females and crawlers. (Courtesy of
Catholic University, Piacenza, Italy.)
key pest of peach worldwide. San Jose scale
was inadvertently introduced to the USA via
San Jose, California, in 1873, which explains
how an insect thought to be native to north-
ern China, eastern Siberia and Korea (Rosen
and DeBach, 1978) came to have such a decid-
edly inappropriate common name (Flanders,
1960). San Jose scale is difficult to see and
readily spread on infested fruit, bud wood or
young trees, accounting for its global distri-
bution. San Jose scale has a broad host range,
attacking at least 34 host families and over
700 plant species. San Jose scale is often par-
ticularly problematic in stone fruit (Vasseur
and Schvester, 1957; Gentile and Summer,
1958). San Jose scale is a major pest of peach
which benefits from annual management
inputs in virtually all production areas (Rice
et al., 1979; Kyparissoudas, 1987a,b; Masoodi
et al., 1989).
San Jose scale is an armoured scale and,
except for the first nymphal stage and the
adult males, all life stages live under hard,
protective scale coverings. The scales of adult
female San Jose scale are round, about 1.6–2
mm across, grey and flattened, with a raised
projection or nipple in the centre (Fig. 18.28/
Plate 213). Beneath her scale covering, the
adult female is yellow, flattened, somewhat
pear-shaped and immobile. Immature male
 Insects and Mites
Fig. 18.28. San Jose scale: adult females and crawlers. (Courtesy of Catholic University, Piacenza, Italy.)
San Jose scales are similarly coloured, but
found beneath elongated scale coverings. The
adult male is a tiny, yellow, winged insect
which seeks out and mates with virgin females.
San Jose scale males can be differentiated from
other small, yellow, gnat-like insects by a dark
band across the back, their prominent beaded
antennae and small black eyes (Strand, 1999).
Fertilized females produce eggs which are
deposited under the mother’s protective scale.
Eggs very quickly hatch and the small, yel-
low, mite-like crawlers disperse to search for
a suitable site to feed and mature. Shortly
after settling, the crawler inserts its stylet-like
mouthparts into the host plant to feed, loses
its antennae, eyes and legs, and secretes a
greyish-white protective scale called a white
cap. Females remain immobile throughout
their lives, while the males regain their mobil-
ity as adults to seek out and mate with the
immobile females. After only a few days, the
white cap scale secretes a series of dark rings,
which indicates attainment of the immature
scale’s next stage, which is called a black cap.
After a subsequent moult, the immature scale
takes on its dark grey adult colour, and the
elongate males can be distinguished from the
round females (Strand, 1999).
493
San Jose scale is very prolific, each female
being able to produce up to 400 crawlers per
season (Quaintance, 1960). When abundant,
San Jose scale can kill fruiting wood, scaffold
limbs and trees within 3 years (Marlatt, 1953).
Although biocontrol can be impressive, natu-
ral enemies are seldom able to hold scale pop-
ulations in peach below damaging levels
(Rice et al., 1979). Heavily infested trees will
have scale on the fruit, producing inflamed
red lesions around the scale, which will still
have its characteristic nipple. Fruit infesta-
tions are an indicator of very high San Jose
scale populations, which should be expected
to reduce vigour and fruit size, and ultimately
kill limbs and trees.
Management of San Jose scale, and other
armoured scale species which attack peach,
requires annual inputs in order to prevent
injury and loss of productivity. Insecticidal
applications for San Jose scale target either
the sessile overwintering stages or the crawl-
ers (Downing and Logan, 1977). Dormant oil
applications should be the foundation of scale
control in peach. The concentration of oil
applied may be varied, with lower concentra-
tions being appropriate early in the dormant
season and again shortly before bud break.
494 D.L. Horton et al.
Higher concentrations are somewhat more
effective, but they should be reserved for the
cooler months when trees are more fully dor-
mant. As a general rule, two dormant oil
applications each winter will consistently
provide better control than a single applica-
tion. This is especially so in regions with heavy
scale pressure and mixed populations of San
Jose scale and WPS (Cochran et al., 1953). Oil
applications may be made 10–14 days apart
or several months apart.
IGR chemistries are valuable, though
expensive, corrective materials that are best
reserved for problematic blocks which expe-
rienced heavy scale build-up in the previous
season, but did not receive two dormant oil
applications. Growth regulators may be applied
pre-bloom as buds break or they may be
applied as the first crawlers appear. In like
fashion, where pest resistance is not a problem,
high-rate application of a broad-spectrum,
organophosphate insecticide cover spray,
applied as crawler movement is initiated, is
also very effective. As with growth regulator
applications, this management approach is
dependent on timing the applications to coin-
cide with the onset of crawlers. The cryptic
nature of scale and the high attention to detail
required to successfully time spray applica-
tion to crawler movement make this approach
challenging.
However, reliable techniques are avail-
able for monitoring San Jose scale life stages.
San Jose crawlers may be monitored by plac-
ing transparent, double-sided Scotch™ tape
around selected, scale-infested branches of a
consistent size (2 cm diameter) on several
trees per block (Badenes-Perez et al., 2002).
Tapes should be hung well before bud break.
Old tapes should be removed and new ones
put in place, with a 2-week schedule working
well. Tapes should be examined for the pres-
ence of crawlers using a dissecting micro-
scope. The San Jose scale phenology model of
Jorgensen et al. (1981) may be used with
meteorological data to predict scale life cycle
events. A biofix, consisting of the season’s
first capture of male San Jose scale in phero-
mone traps, is necessary to properly time
initiation of heat unit accumulation. Phero-
mone traps are the most practical means to
monitor adult male emergence.
Aphids (Hemiptera: Aphididae)
Aphids are soft-bodied, teardrop-shaped
insects which use their piercing and sucking
mouthparts to imbibe a liquid diet of plant
juices (Fig. 18.29/Plate 214). Most aphid spe-
cies have very high reproductive rates, and it
is common to see dense aggregations of these
slow-moving insects on succulent plant ter-
minals. In peach, the impact of direct aphid
injury, the result of aphid feeding, is typically
modest, and many peach growers regard aphids
as minor, or even inconsequential, pests.
Aphids of several species are known to
utilize peach as a host. In the USA, surveys by
Stoetzel and Miller (1998), Gildow et al. (2004)
and Wallis et al. (2005) overviewed the peach
aphids of North America, with particular
emphasis on their potential as viral vectors.
Blackman and Eastop (1985) included peach
aphids in their survey of crop-feeding aphids.
Naturally occurring biological control is
normally effective in mitigating aphid abun-
dance in peach. Even in productions systems
that must utilize regular application of broad-
spectrum insecticides to control key fruit or
tree pests, a robust complement of predacious
and parasitic insects is commonly successful
in bringing even the occasional high aphid
population down to low or moderate abun-
dances.
Aphids are, however, important vectors
of plant viruses. Numerous aphid species
feed on peach and many of them are vectors.
The virus vector potential of these aphid spe-
cies is far more significant to the health and
performance of peaches than direct feeding
injury. The vectors of Plum pox virus (PPV) in
peach in the eastern USA have been surveyed
(Stoetzel and Miller, 1998; Gildow et al., 2004).
However, the detailed biology and behaviour
of these aphids, which at least in the USA
were regarded as minor peach pests, is inad-
equately understood.
PPV, which causes plum pox, also known
as sharka disease, is perhaps the most serious
of the viruses of peach. Plum pox-infected
trees suffer dramatic reductions in fruit qual-
ity and yield (Kölber et al., 2001). PPV is
spread over long distances by human activi-
ties, primarily movement of asymptomatic
virus-infested bud wood or young trees and
 Insects and Mites
Fig. 18.29. Green peach aphid: parthenogenetic female and nymphs. (Courtesy of Catholic Univer-
sity, Piacenza, Italy.)
rootstocks. Movement within and among
adjacent orchards is by aphid transmission
(Shukla et al., 1994). PPV was first observed in
the Balkans, but it has since spread through-
out most key peach production areas in
Europe, the Middle East, China and South
America. For many years plum pox was suc-
cessfully excluded from North America.
However, in 1999 the disease was confirmed
in the north-eastern USA, in Adams County,
Pennsylvania. A second PPV infestation was
subsequently confirmed in south-eastern Can-
ada, on the Niagara Peninsula in 2000 (Levy
et al., 2000). This infestation has since spread
across the Saint Lawrence River to stone fruit
orchards in neighbouring New York State,
USA. Orchard destruction programmes have
been implemented in each of these areas in an
effort to eradicate the disease.
In eastern Europe Hyalopterous pruni
(Geoffroy) and Phorodon humuli (Schrank) are
felt to be key PPV vectors. In western Europe
and Pennsylvania, USA, the green peach
aphid, Myzus persicae (Sulzer), and the spirea
aphid, Aphis spiraecola (Patch), are considered
to be the more important vectors of PPV (Wallis
et al., 2005). Plum pox is not easy to detect,
495
and infected trees may be visually asymp-
tomatic. Aphid transmission of PPV is non-
persistent, meaning the virus can be acquired
by aphids which either feed or briefly probe
infested material to determine host suitabil-
ity. It is generally accepted that potyviruses
such as PPV are seldom carried long distances
because the vector’s infectivity is short-lived
(Wallis et al., 2005).
Green peach aphid is a cosmopolitan
species which is important as a pest of numer-
ous vegetable, ornamental and fruit crops
around the world (Blackman and Eastop,
1985). Green peach aphid is discussed here
because of its widespread distribution as both
a direct pest of stone fruit and its importance
as a virus vector of peach. In southern Europe,
green peach aphid is a major direct pest of
peaches, owing to its injury of succulent ter-
minal growth and leaves and feeding on the
fruit, which can reduce fruit quality (Pascal
et al., 2002). Green peach aphid is also regarded
as one of the key European vectors of PPV
(Pascal et al., 2002). Management of green
peach aphid in southern Europe is compli-
cated to some degree by the aphid’s resistance
to insecticides (Devonshire et al., 1998). In the
496 D.L. Horton et al.
USA peach aphids, including the green peach
aphid and spirea aphid, have not been regarded
as important pests. Control, when needed, has
been insecticidal. In warmer production areas
such as California (Strand, 1999) and the south-
eastern USA (Baker, 1994), green peach aphid
moves from one crop or weed host to the next.
In these regions, they show no dramatic pref-
erence for peach and there is no egg stage.
Both green peach aphid and spirea aphid are
more commonly seen on peach in these regions
during the early spring. In cooler production
areas, such as Michigan, green peach aphid
overwinters on peach as either eggs or mated
females. In the spring, as peach foliage emerges,
female green peach aphids give birth partho-
genetically to wingless offspring (Fig. 18.29/
Plate 214). By June winged adults develop and
disperse, leaving peach for summer hosts. They
return to peach in the autumn (Howitt, 1993).
18.4 Mite Pests
Two-spotted spider mite (TSM), European
red mite (ERM) and other foliage-feeding
mites are indirect pests of peaches. Relative to
other deciduous fruits, peaches are very toler-
ant of mite feeding. It is generally accepted
that mites are more problematic in drier pro-
duction areas and that peaches with adequate
moisture are more tolerant of mites. It is also
believed that mite outbreaks in many orchard
systems are at least partially an unintended
consequence of broad-spectrum insecticide
use for fruit- or tree-attacking pests (Croft
et al., 1987). Kovach and Gorsuch (1986)
asserted that pyrethroid insecticides should
have only a modest role on south-eastern
peaches because of their tendency to encour-
age TSM populations.
Multiple studies indicate that peach yield
and/or fruit quality were not reduced until
mite numbers were quite high. Kovach and
Gorsuch (1985) reported that more than 48
TSM/leaf were required to reduce the per-
centage of fruits that reached the more desir-
able, large size categories. In like fashion,
Bailey (1979) showed that TSM densities of
40–50 mites/leaf did not reduce peach yield
in South Australia. Working with ERM on
peaches, McClernan and Marini (1986) showed
no reduction of yield, fruit quality or tree
growth parameters at mite densities of less
than 100 per leaf. In Europe, IPM strategies in
northern Italy significantly reduced the dam-
age produced by tetranychid mites, whose
importance still remains high in more arid
climates. Problematic situations have been
reported in Greece.
Peach’s ability to tolerate mite infesta-
tions with very modest risk of yield loss must
be balanced with tree health issues. Defoli-
ation from heavy mite infestations exposes
scaffold limbs and ripening fruit to sunburn.
Miticides should be applied in advance of
defoliation to protect the scaffold limbs and
fruit from sun injury. Hogmire (1995) advo-
cated conservative provisional mite thresh-
olds for peaches, suggesting treatment for
>10 mites/leaf during mid-season and >20
mites/leaf as fruit approach maturity. These
values are approximately twice those used in
apples at similar points in crop development.
Hogmire (1995) also noted, as have others,
that it is often necessary to treat peaches for
mites preharvest to lessen the dermal irrita-
tion experienced by pickers.
Resistance management is a logical con-
sideration in formulating pest management
plans. While miticide use in peach is modest,
it is well to be aware that various TSM popu-
lations have shown themselves to be resistant
to chlorpyrifos, dicofol, cyhexatin, abamectin
and two ovicides, clofentezine and hexythi-
azox, which have a common mode of action
(Stumpf and Nauen, 2001). Overuse of the
‘best’ miticides, and use of least-cost cover
spray insecticides (pyrethroids) which may
adversely impact natural enemy populations
and enemies, can work in concert to increase
the selective pressures favouring resistance
(Croft et al., 1987). The most practical means
of managing miticide resistance is the alterna-
tion of miticides with differing modes of
action and the incorporation of horticultural
mineral oils into a management programme.
Resistance management plans should be
standard for mites and other resistance-prone
pests. Pest management advisors should help
growers develop plans by describing use pat-
terns to ensure rotation between unlike modes
of action. Such plans can easily be developed
164 165

166 167

169

168

Plate 164. Leaves on branch infected with European stone fruit yellows phytoplasma (left) are small in size, rolled
upward longitudinally, and have a pale yellow colour compared with leaves on a non-infected branch (right) (from
A. Ragozzino).
Plate 165. Peach tree infected with European stone fruit yellows phytoplasma with leaf midrib and lateral veins
enlarged and necrotic (from L. Giunchedi).
Plate 166. Two trees of the same age of cultivar ‘Babygold 7’. The tree in foreground is infected with the
European stone fruit yellows phytoplasma (exhibiting reduced vigour) compared with the non-infected tree in the
background (from A. Ragozzino).
Plate 167. Discoloration symptoms of Plum pox virus type M on peach cv. ‘Baby Gold 6’ flowers (courtesy of J.C.
Desvignes, La Force, France).
Plate 168. Colour break symptoms of Plum pox virus type M on petals of peach cv. ‘Gladys’ flowers (courtesy of
M.A. Cambra, Zaragoza, Spain).
Plate 169. Leaf symptoms of Plum pox virus on ‘GF 305’ peach seedlings used as indicator plant.
170 171

172 173

174 175

Plate 170. Leaf symptoms of Plum pox virus type M on peach cv. ‘Royal Gem’.
Plate 171. Fruit symptoms (under-pigmented yellow rings) caused by Plum pox virus in peach cv. ‘Springcrest’.
Plate 172. Chlorotic rings and line pattern caused by Plum pox virus in nectarine cv. ‘Arm King’ fruits.
Plate 173. Chlorotic spots and under-pigmented yellow rings caused by Plum pox virus in peach cv. ‘Catherine’
fruits (courtesy of M.A. Cambra, Zaragoza, Spain).
Plate 174. Leaf symptoms (chlorotic rings and deformation) caused by Prune necrotic ring spot virus in peach.
Plate 175. Necrotic spots caused by Prunus necrotic ring spot virus in peach leaves.
176 177

178 179

180

Plate 176. Dense canopy due to shortening of shoot internodes caused by Prune dwarf virus in peach trees.
Plate 177. Leaf symptoms (bright yellow mosaic and spots) caused by Apple mosaic virus in peach.
Plate 178. Severe reduction of leaf growth on buds giving a rosette appearance caused by Tomato ringspot virus
in ‘GF 305’ peach seedlings.
Plate 179. Internode shortening caused by Strawberry latent ringspot virus in peach trees.
Plate 180. Fruit symptoms (deformations and discolorations with cracked sutures) caused by Peach latent mosaic
viroid in peach.
181 182

183 184

186
185

Plate 181. Leaf symptoms (yellow-creamy mosaic) caused by Peach latent mosaic viroid in peach.
Plate 182. White patterns (calico) covering most of the leaf area caused by Peach latent mosaic viroid in
nectarine cv. ‘Red-Gin’ leaves.
Plate 183. Leaf symptoms (chlorotic spots) caused by Peach sooty ringspot virus in peach.
Plate 184. Sooty green rings on a yellow background in senescent leaves caused by Peach asteroid spot virus in peach.
Plate 185. Leaf symptoms (yellowish line patterns and russet rings) and fruit symptoms (chlorotic rings and
mosaic) caused by Apple chlorotic leaf spot virus in peach.
Plate 186. Oriental fruit moth injury to peach terminal (University of Georgia, Bugwood).
187 188

189
190

191 192

Plate 187. Oriental fruit moth adult (University of California, IPM).

Plate 188. Anal plate of oriental fruit moth larva (University of California, IPM).
Plate 189. Oriental fruit moth larva tunnelling in peach (University of California, IPM).
Plate 190. Adult codling moth (University of California, IPM).
Plate 191. Adult of the peach twig borer moth (Catholic University, Piacenza).
Plate 192. Mature peach twig borer larva (University of California, IPM).
193 194

195 196

198
197

Plate 193. Tufted bud moth adult (courtesy of G. Krawczyk, University Park, Pennsylvania, USA).
Plate 194. Tufted bud moth larva on fruit (courtesy of G. Krawczyk, University Park, Pennsylvania, USA).
Plate 195. Plum curculio adult (University of Georgia, Bugwood).
Plate 196. Plum curculio oviposition wound (courtesy of J.A. Payne, University of Georgia).
Plate 197. Japanese beetle adult (University of Georgia, Bugwood).
Plate 198. Green June beetle adult (University of Georgia, Bugwood).
199 200

201
202

203
204

Plate 199. Tarnished plant bug adult (courtesy of S. Bauer, USDA).

Plate 200. Brown stink bug adult (courtesy of J. Greene, Clemson University, South Carolina, USA).
Plate 201. Leaffooted bug adult (courtesy of D. Cappaert, University of Georgia, Bugwood).
Plate 202. Cat-facing injury on peach (University of Georgia, Bugwood).
Plate 203. Western flower thrips on peach leaf (courtesy of J.A. Payne, University of Georgia, Bugwood).
Plate 204. Western flower thrips russeting on fruit (courtesy of J.A. Payne, University of Georgia, Bugwood).
205 206

207 208

209 210

Plate 205. Western flower thrips silvering on fruit (courtesy of J.A. Payne, University of Georgia, Bugwood).
Plate 206. Mediterranean fruit fly mating (Catholic University, Piacenza).
Plate 207. Peachtree borer larva (University of Georgia, Bugwood).
Plate 208. Branches damaged by lesser peachtree borer (courtesy of J. Fuest, University of Georgia).
Plate 209. Adult female peachtree borer (courtesy of J. Fuest, University of Georgia).
Plate 210. Adult male peachtree borer (University of Georgia, Bugwood).
211 212

213 214

215 216

Plate 211. Lesser peachtree borer wounding (courtesy of J. Fuest, University of Georgia).
Plate 212. White peach scale. Adult females and crawlers (Catholic University, Piacenza).
Plate 213. San José scale. Adult females and crawlers (Catholic University, Piacenza).
Plate 214. Green peach aphid. Parthenogenetic female and nymphs (Catholic University, Piacenza).
Plate 215. Twospotted spider mite with egg (University of Georgia, Bugwood).
Plate 216. European red mite adult (INRA Montepellier).
217

218

Plate 217. Galls on ‘Okinawa’ (A) and ‘Nemaguard’ (B) peach roots caused by Meloidogyne floridensis (courtesy
of W.B. Sherman, Gainesville, Florida, USA).
Plate 218. Galls on peach (A and B) and Myrobalan plum (C) roots caused by root-knot nematode (RKN),
Meloidogyne sp. Severe early symptoms (A); root decay following an attack (B); two Myrobalan plum individuals
from a progeny segregating for the Ma RKN resistance gene (left = host; right = resistant) (C) (photos by A.P. Ny-
czepir (A) and D. Esmenjaud (B, C)).
219

220 221

Plate 219. Reproductive life cycle of Meloidogyne spp. (N = female nematode; * = giant cells) (courtesy of INRA,
Sophia-Antipolis, France).
Plate 220. Galls on Myrobalan plum root caused by the association of Meloidogyne spp. and Agrobacterium
tumefaciens (crown gall) (photo by D. Esmenjaud, INRA).
Plate 221. Influence of Mesocriconema xenoplax (Mx) on ‘Nemaguard’ peach feeder root growth after 6 months
((–)Mx, uninoculated; (+)Mx, inoculated; Pi = 14,000 nematodes/1500 cm3 soil, where Pi is initial nematode
population density) (photo by A.P. Nyczepir).
222 223

224 225

226

Plate 222. Dead peach tree with suckers at the crown during summer in the presence of Mesocriconema xeno-
plax and the peach tree short life disease complex (photo by A.P. Nyczepir).
Plate 223. Typical cambial tissue damage of the trunk above the soil line and healthy viable tissue below the soil
line in tree dying from peach tree short life disease (photo by A.P. Nyczepir).
Plate 224. Eight dead 3-year-old peach trees on ‘Nemaguard’ rootstock (foreground) and live trees on all
‘Guardian®’ rootstock (background, same row) in the presence of Mesocriconema xenoplax and the peach tree
short life disease complex (photo by A.P. Nyczepir).
Plate 225. Influence of Pratylenchus vulnus on ‘G × N No. 15’ almond–peach hybrid root growth after 24 months
(left, uninoculated; right, inoculated; Pi = 1000 nematodes/plant where Pi is initial nematode population density)
(courtesy of J. Pinochet, Agromillora Catalana SA, Barcelona, Spain).
Plate 226. Peaches picked at different maturity levels.
227 228

229

230

231
232

Plate 227. The influence of increased nitrogen fertilization (kg/ha) on red skin coloration of 'Fantasia' nectarine.
Plate 228. Water stress late in the summer causes fruit defects such as deep sutures and double-fruit formation.
Plate 229. Canopy position affects fruit size, red colour development and storage potential.
Plate 230. Leaf removal around the fruit improves red colour but may decrease fruit size.
Plate 231. Peach girdling (removal of a strip of scaffold bark) at the main scaffolds advances maturity and in-
creases fruit size.
Plate 232. Internal breakdown symptoms in peaches (top of image) include flesh mealiness, flesh browning and
loss of flavour.
233

235

234

237

236

Plate 233. Peach inking or staining as a consequence of abrasion combined with heavy metal contamination dur-
ing harvesting and hauling operations.
Plate 234. Bin of peaches being pre-cooled on conveyor-type hydro-cooler prior to packing.
Plate 235. Packaged fruit in unitized pallet loads are stacked to form a forced-air cooling tunnel.
Plate 236. Storage temperature influences incidence and severity of internal breakdown in susceptible cultivars.
Plate 237. Peaches for fresh market are hand picked. Harvesters work on ladders using picking bags or baskets.
238 239

240

241

Plate 238. Harvesters transfer peaches to field bins which are moved through the field on low trailers.
Plate 239. View of a shaded loading area to protect fruit from excess heating while awaiting transportation to the
packing house.
Plate 240. Dry bin dumping of fruit on to a commercial packing line.
Plate 241. Sorting peaches by skin colour and removing blemished fruit.
242

243

244

245

Plate 242. Packers sizing, sorting and packing fruit by hand into two-layer tray packs.
Plate 243. Fruit moving on to an electronic weight sizer.
Plate 244. Cull removal and disposal can be a major problem and expense in peach packing.
Plate 245. Peach fruit display at a retail store.
 Insects and Mites
by referring to the Insecticide Resistance
Action Committee’s eClassification site (http://
www.irac-online.org/eClassification/), which
facilitates this process by assigning numbers
to classes of toxins. Alternating among miti-
cides of differing numbers substantially reduces
selective pressure for resistance.
Two-spotted spider mite, Tetranychus
urticae Koch (Acari: Tetranychidae)
Two-spotted spider mite, Tetranychus urticae
Koch, is a cosmopolitan, indirect, foliage-feed-
ing pest of fruits, vegetables and nursery crops
worldwide, and is well known as a pest of
peach (Unwin, 1971) (Fig. 18.30/ Plate 215).
TSM is the most commonly injurious
phytophagous mite of peaches. In California,
TSM and Pacific spider mite, Tetranychus paci-
ficus (McGregor), are the key mite pests of
peaches and other stone fruits. Pacific spider
mite feeds on both the upper and lower sides
of leaves, while TSM feeds primarily on the
undersides. The biology and management of
these web-spinning mites are quite similar;
hence the discussion that follows will focus on
the more widely problematic TSM. In Europe,
Fig. 18.30. Two-spotted spider mite with egg. (Courtesy of University of Georgia, Bugwood
Network.)
497
TSM is more problematic as a peach pest in
warmer and drier areas like southern Italy
and Greece.
In most peach-producing areas, TSM
overwinters as adult females on the orchard
floor or at the base of trees. The overwinter-
ing females are reddish-orange and their
namesake spots are less visible. In North Car-
olina, TSM continues to reproduce on hosts
such as henbit during mild winters, though at
a reduced rate (Meagher and Meyer, 1990). As
peaches leaf out in spring, TSM moves first to
foliage in the lower part of trees. Eggs, which
are spherical and translucent, are laid primar-
ily on the underside of leaves. Eggs hatch into
six-legged larvae, which subsequently pass
through two eight-legged nymphal stages
before reaching the eight-legged adult stage.
TSM adults are small, about 0.4 mm long,
pale green, greenish-amber or yellowish, with
two, sometimes four, spots on their sides.
TSM development is rapid and temperature-
dependent, with generation times as short as
5 days being seen under optimal conditions.
Mite injury within an orchard is often clumped;
however, when densities are high, air-borne
dispersal of mites occurs with even gentle
breezes.
498 D.L. Horton et al.
TSM feeds by piercing the epidermis of
leaves, releasing sap which the mite ingests.
Injured mesophyll cells collapse and produce
a small chlorotic spot at each feeding site. A
delicate stippling develops and, as infesta-
tions progress, the leaves become off-colour,
often turning yellow or bronze (Meagher and
Meyer, 1990). Leaf discoloration, webbing and
defoliation can develop quickly. TSM develop-
ment is favoured by hot, dry conditions, which
lends more significant pest status to TSM in
California and other arid production areas.
European red mite, Panonychus ulmi
(Koch) (Acari: Tetranychidae)
European red mite, Panonychus ulmi (Koch), is
an occasional pest of peaches (Fig. 18.31/
Plate 216) that is generally considered to be
less important than TSM. However, ERM is
abundant on peaches, producing stippling
and bronzing of leaves and defoliation. Popu-
lation growth of ERM is favoured by humid,
shady conditions, which may partially explain
its greater importance as a mite pest of
peaches in the more humid production areas
of eastern North America and northern Italy
in Europe. McClernan and Marini (1986)
Fig. 18.31. European red mite adult. (Courtesy of INRA, Montepellier, France.)
observed that ERM was the most prevalent
phytophagous mite of peaches in New Jersey,
and that its numbers often increase dramati-
cally in mid-summer after orchards are treated
with carbamate insecticide for scarab beetle
infestations. Similar observations by Croft
et al. (1987) from work with apples under-
score the potential consequences of inadver-
tently shifting the predator/phytophagous mite
balance to favour the plant-feeding mites.
ERM overwinters in the egg stage on
host trees. Eggs are dark red in colour, slightly
flattened, with a hair-like spine that projects
up from the centre (Hogmire, 1995). The eggs
begin to hatch as peaches leaf out, and the
resultant nymphal mites move directly to
foliage. As is the case with TSM, ERM passes
through a six-legged larval stage, followed by
two eight-legged nymphal stages before
moulting to an eight-legged adult. ERM adults
are similar in size to TSM, but are a dark
brick-red colour, with long curved dorsal hairs
projecting from whitish spots on the mites’
back (Hogmire, 1995).
As is the case with TSM and silver mite, the
presence of stable, low numbers of ERM popu-
lations should be viewed positively, because
non-injurious, low-level infestations provide
a food source for predacious complexes that
 Insects and Mites 499

normally provide biological control of these
species in peaches (Strand, 1999).
Management of ERM in many peach
orchards is aided by one to two dormant oil
applications made for scale control. Infesta-
tions can normally be monitored by being
alert for foliage that is off-colour or stippled.
The work of McClernan and Marini (1986)
noted that outbreaks of ERM do occur on
peach, but that ERM is a minor pest that
should be dealt with as needed using curative
miticide treatments. Treatments, particularly
those made late in the season, are often made
for the comfort of harvesters who sometimes
suffer dermal irritation from disoriented
mites that crawl over their skin.

References

Anon. (2006) The Mediterranean Fruit Fly, APHIS Plant Protection and Quarantine, Factsheet. http://www.
aphis.usda.gov/publications/plant_health/content/printable_version/fs_phmedfly.pdf (accessed 2006).
Atanassov, A., Shearer, P.W., Hamilton, G. and Polk, D. (2002) Development and implementation of a
reduced risk peach arthropod management program in New Jersey. Journal of Economic Entomology
95, 803–812.
Badenes-Perez, F.R., Zalom, F.G. and Bentley, W.J. (2002) Effects of dormant insecticide treatments on the San
Jose scale (Homoptera: Diaspididae) and its parasitoids Encarsia perniciosi and Aphytis spp. (Hy-
menoptera: Aphelinidae). International Journal of Pest Management 48, 291–296.
Bailey, P. (1979) Effect of late season populations of twospotted mite on yield of peach trees. Journal of Eco-
nomic Entomology 72, 8–10.
Baker, J.R. (1994) Mites. In: Baker, J.R. (ed.) Insect and Related Pests of Flowers and Foliage Plants. AG-136.
North Carolina State University, Cooperative Extension Service, Raleigh, North Carolina, pp. 57–62.
Benoit, D.L., Vincent, C. and Chouinard, G. (2006) Management of weeds, apple sawfly (Hoplocampa
testudinea Klug) and plum curculio (Conotrachelus nenuphar Herbst) with cellulose sheeting. Crop
Protection 25, 331–337.
Biddinger, D., Hull, H., Huang, H., McPheron, B. and Loyer, M. (2006) Sublethal effects of chronic exposure
to tebufenozide on the development, survival, and reproduction of the tufted apple bud moth (Lepi-
doptera: Tortricidae). Journal of Economic Entomology 99, 834–842.
Blackman, R.L. and Eastop, V.F. (1985) Aphids on the World’s Crops. John Wiley & Sons, Chichester, UK.
Bobb, M.L., Weidhaus, J.A. and Ponton, L.F. (1973) White peach scale: life history and control studies. Journal
of Economic Entomology 66, 1290–1292.
Brandhorst-Hubbard, J.L., Flanders, K. and Appel, A.G. (2001) Oviposition site and food preference of the
green June beetle (Coleoptera: Scarabaeidae). Journal of Economic Entomology 94, 628–633.
Brittain, J.A. and Miller, R.W. (1978) Managing peach tree short life in the Southeast. Circular 585. Clemson
University Cooperative Extension Service, Clemson, South Carolina.
Cochran, J.H., Ferree, R.J., Foster, H.H., Nettles, W.C. and Petersen, D.H. (1953) Peach Pest Control in South
Carolina. Circular 360. Clemson Agricultural College, Cooperative Extension Service, Clemson, South
Carolina.
Cochran, J.H., Ferree, R.J., Foster, H.H., Nettles, W.C. and Petersen, D.H. (1955) Peach Pest Control in South
Carolina. Circular 360. Clemson University Cooperative Extension Service, Clemson, South Carolina.
Collins, F.A. and Whitcomb, W.H. (1975) Natural enemies of the white peach scale, Pseudaulacaspis pen-
tagona (Homoptera: Coccidae), in Florida. The Florida Entomologist 58, 15–21.
Cossentine, J.E., Banham, F.L. and Jensen, L.B. (1990) Efficacy of the nematode, Heterorhabditis heliothidis
(Rhabditida: Heterorhabditidae) against the peachtree borer, Synanthedon exitiosa (Lepidoptera: Sesiidae)
in peach trees. Journal of the Entomological Society of British Columbia 87, 82–84.
Cottrell, T.E. (2001) Improved trap capture of Euschistus servus and Euschistus tristigmus (Hemiptera: Pentato-
midae) in pecan orchards. The Florida Entomologist 84, 731–732.
Cottrell, T.E. and Shapiro-Ilan, D.I. (2006) Susceptibility of the peachtree borer, Synanthedon exitiosa, to Stein-
ernema carpocapsae and Steinernema riobrave in laboratory and field trials. Journal of Invertebrate Pa-
thology 92, 85–88.
Cravedi, P. and Molinari, F. (1996) Mating disruption method against Cydia molesta (Busck) in Italy. Acta
Horticulturae 374, 71–76.
500 D.L. Horton et al.

Cravedi, P., Guarino, F. and Tocci, A. (2001) Valuations about mating disruption method application in Cydia
molesta (Busck) control on nearly 400 hectares of peach tree in the Plain of Sibari (Calabria, South Italy).
Bulletin OILB/SROP 24(5), 79–84.
Croft, B., Weakley, C. and Rice, R. (1980) Validation of a PETE timing model for the oriental fruit moth in
Michigan and central California (Lepidoptera: Olethreutidae). Great Lakes Entomologist 13, 211–217.
Croft, B.A., Hoyt, S.C. and Westigard, P.H. (1987) Spider mite management on pome fruits, revisited: organo-
tin, and acaricide resistance management. Journal of Economic Entomology 80, 304–311.
Devonshire, A.L., Field L.N., Foster, S.T., Moores, G.D., Williamson, M.S. and Blackman, L. (1998) The evolu-
tion of insecticide resistance in the peach-potato aphid, Myzus persica. Transactions of the Royal Society
of London, B 353, 1677–1684.
Downing, R. and Logan, D. (1977) A new approach to San Jose scale control (Hemiptera: Diaspididae). Ca-
nadian Entomologist 109, 1249–1252.
Epstein, D.L., Stelinski, L.L., Reed, T.P., Miller, J.R. and Gut, L.J. (2006) Higher densities of distributed phero-
mone sources provide disruption of codling moth (Lepidoptera: Tortricidae) superior to that of lower
densities of clumped sources. Journal of Economic Entomology 99, 1327–1333.
Erkiliç, L.B. and Uygun, N. (1997) Development time and fecundity of the white peach scale, Pseudaulacaspis
pentagona, in Turkey. Phytoparasitica 25, 9–16.
Flanders, K.L. and Johnson, D. (2005) Green June beetle. In: Horton, D. and Johnson, D. (eds) Southeastern
Peach Growers Handbook. GES Handbook No. 1. University of Georgia College of Agricultural & Envi-
ronmental Sciences, Athens, Georgia, pp. 259–260.
Flanders, S.E. (1960) The status of San Jose scale parasitization (including biological notes). Journal of Eco-
nomic Entomology 53, 757–759.
Gentile, A.G. and Summer, F.M. (1958) The biology of the San Jose scale on peaches with special reference to
the behavior of males and juveniles. Hilgardia 27, 269–285.
Gildow, F., Damsteegt, V., Stone, A., Schneider, W., Lustre, D. and Levy, L. (2004) Plum pox in North America:
identification of aphid vector and a potential role for fruit in virus spread. Phytopathology 94, 868–
874.
Hanks, L.M. and Denno, R.F. (1993) The white peach scale, Pseudaulacaspis pentagona (Targioni-Tozzetti)
(Homoptera: Diaspididae) life history in Maryland, host plants and natural enemies. Proceedings of the
Entomological Society of Washington 95, 79–98.
Hathaway, D.O., Tamaki, G., Moffitt, H.R. and Burditt, A.K. (1985) Impact of removal of males with sex-
pheromone-baited traps on suppression of the peach-twig borer, Anarsia lineatella (Zeller). The Cana-
dian Entomologist 117, 643–645.
Hendrichs, J. (1996) Action programs against fruit flies of economic importance: session review. In: McPher-
on, B.A. and Steck, G.J. (eds) Fruit Fly Pests: A World Assessment of Their Biology and Management. St
Lucie Press, Boca Raton, Florida, pp. 513–519.
Hodges, G. (2005) Scale insects. In: Horton, D. and Johnson, D. (eds) Southeastern Peach Growers Hand-
book. GES Handbook No. 1. University of Georgia College of Agricultural & Environmental Sciences,
Athens, Georgia, pp. 230–235.
Hogmire, H.W. (ed.) (1995) Mid-Atlantic Orchard Monitoring Guide. Northeast Regional Agricultural Engi-
neering Service (75), Cooperative Extension, Ithaca, New York.
Hogmire, H.W. and Leskey, T.C. (2006) An improved trap for monitoring stink bugs (Heteroptera: Pentatomi-
dae) in apple and peach orchards. Journal of Entomological Science 41, 9–21.
Horton, D., Bellinger, B., Pettis, G., Brannen, P. and Mitchem, W. (2005) Pest management plan for eastern
peaches. USDA Office of Pest Management Policy, Pest Management Strategic Plan. http://www.ipmcenters.
org/pmsp/pdf/EastPeach.pdf (accessed 2005).
Horton, D., Brannen, P., Bellinger, B. and Ritchie, D. (2007) Southeastern Peach, Nectarine and Plum Pest
Management and Culture Guide. Cooperative Extension Bulletin No. 1171. University of Georgia, Ath-
ens, Georgia.
Howitt, A.H. (1993) Oriental fruit moth. In: Howitt, A.H. (ed.) Common Fruit Tree Pests. North Central Re-
gional Extension Publication No. 63. Michigan State University, East Lansing, Michigan, pp. 43–44,
202–207.
Il’ichev, A.L., Stelinski, L.L., Williams, D.G. and Gut, L.J. (2006) Sprayable microencapsulated sex pheromone
formulation for mating disruption of oriental fruit moth (Lepidoptera: Tortricidae) in Australian peach and
pear orchards. Journal of Economic Entomology 99, 2048–2054.
Jackson, T.A. and Klein, M.G. (2006) Scarabs as pests: a continuing program. Coleopterists Society Monograph
5, 102–119.
 Insects and Mites 501

Jenkins, D., Cottrell, T., Horton, D., Hodges, A. and Hodges, G. (2006) Hosts of plum curculio, Conotrachelus
nenuphar (Coleoptera: Curculionidae), in central Georgia. Environmental Entomology 35, 48–55.
Johnson, D., Cottrell, T. and Horton, D. (2005) Plant bugs and stink bugs. In: Horton, D. and Johnson, D. (eds)
Southeastern Peach Growers Handbook. GES Handbook No. 1. University of Georgia College of
Agricultural & Environmental Sciences, Athens, Georgia, pp. 236–239.
Jorgensen, C.D., Rice, R.E., Hoyt, S.C. and Westigard, P.H. (1981) Phenology of the San Jose scale (Ho-
moptera: Diaspididae). The Canadian Entomologist 113, 149–159.
Kanga, L.H.B., Pree, D.J., van Lier, J.L. and Walker, G.M. (1999) Monitoring for resistance to organophosphorus,
carbamate, and pyrethroid insecticides in the oriental fruit moth (Lepidoptera: Tortricidae). The Canadian
Entomologist 131, 441–450.
Kanga, L.H.B., Pree, D.J., van Lier, J.L. and Walker, G.M. (2003) Management of insecticide resistance in ori-
ental fruit moth (Grapholita molesta; Lepidoptera: Tortricidae) populations from Ontario. Pest Manage-
ment Science 59, 921–927.
Killian, J.C. and Meyer, J.R. (1984) Effect of orchard weed management on catfacing damage to peaches in
North Carolina. Journal of Economic Entomology 77, 1596–1600.
Knight, A.L. (2000) Monitoring codling moth (Lepidoptera: Tortricidae) with passive interception traps in sex
pheromone-treated apple orchards. Journal of Economic Entomology 93, 1744–1751.
Kölber, M., Nemeth, M., Chernets, A., Kalashian, Y., Dulic-Markovic, I., Glasa, M., Isac, M., Kriska, B.,
Malinowski, T., Zawadzka, B., Minoiu, N., Myrta, A., Navratil, M., Prichodko, Y., Slovakova, L. and Topchi-
iska, M. (2001) Current situation of plum pox disease on stone fruit species in middle and eastern Europe.
Acta Horticulturae 550, 73–78.
Kovach, J. and Gorsuch, C. (1985) Effect of Tetranychus urticae populations on peach production in South
Carolina. Journal of Agricultural Entomology 2, 46–51.
Kovach, J. and Gorsuch, C. (1986) Response of the twospotted spider mite, Tetranychus urticae Koch, to various
insecticides and fungicides used in South Carolina peach orchards. Journal of Agricultural Entomology 3,
175–178.
Kozar, F., Mazzoni, E. and Cravedi, P. (1997) Comparison of flight periods of male Pseudaulacaspis pentagona
in Hungary and northern Italy. Bulletin OILB/SROP 20, 43–49.
Kuitert, L.C. (1967) Observations on the biology, bionomics and control of white peach scale, Pseudaulacaspis
pentagona (Targ.). Proceedings of the Florida State Horticultural Society 80, 376–381.
Kyparissoudas, D.S. (1987a) The occurrence of Encarsia perniciosi in areas of northern Greece as assessed by
sex pheromone traps of its host Quadraspidiotus perniciosus. Entomologia Hellenica 5, 7–12.
Kyparissoudas, D.S. (1987b) Flight of San Jose scale Quadraspidiotus perniciosus males and time of crawler
appearance in orchards of northern Greece. Entomologica Hellenica 5, 75–80.
Lalancette, N., Belding, R.D., Shearer, P.W., Frecon, J.L. and Tietjen, W.H. (2005) Evaluation of hydrophobic
and hydrophilic kaolin particle films for peach crop, arthropod and disease management. Pest Manage-
ment Science 61, 25–39.
Legrand, A. and Los, L. (2003) Visual responses of Lygus lineolarius and Lygocoris spp. (Hemiptera: Miridae)
on peaches. The Florida Entomologist 86, 424–428.
Leskey, T.C. and Wright, S.E. (2004) Monitoring plum curculio, Conotrachelus nenuphar (Coleoptera: Curcu-
lionidae), populations in apple and peach orchards in the Mid-Atlantic. Journal of Economic Entomology
97, 79–88.
Leskey, T.C., Zhang, A. and Herzog, M. (2005) Nonfruiting host tree volatile blends: novel attractants for the
plum curculio (Coleoptera: Curculionidae). Environmental Entomology 34, 785–793.
Levy, L., Damsteegt, V. and Welliver, R. (2000) First report of plum pox virus (sharka disease) in Prunus persica
in the United States. Plant Disease 84, 202.
Liquido, N.J., Shinoda, L.A. and Cunningham, R.T. (1991) Host plants of the Mediterranean fruit fly (Diptera:
Tephritidae): an annotated world review. Miscellaneous Publications of the Entomological Society of
America 77, 1–52.
Lockwood, D. and Myers, S. (2005) Tree density, orchard design and training systems. In: Horton, D. and
Johnson, D. (eds) Southeastern Peach Growers Handbook. GES Handbook No. 1. University of Georgia
College of Agricultural & Environmental Sciences, Athens, Georgia, pp. 51–64.
Loughrin, J.H., Potter, D.A. and Hamilton-Kemp, T.R. (1995) Volatile compounds induced by herbivory act as
aggregation kairomones for the Japanese beetle (Popillia japonica Newman). Journal of Chemical Ecol-
ogy 21, 1457–1467.
McClernan, W.A. and Marini, R.P. (1986) European red mite on yield, fruit quality, and growth in peach trees.
HortScience 21, 244–246.
502 D.L. Horton et al.

McLaughlin, J.R., Heath, R.R. and Ashley, T.R. (1990) Periodicity of pheromone release from female white
peach scale. Physiological Entomology 15, 193–197.
Maier, C.T. (1990) Native and exotic rosaceous hosts of apple, plum, and quince curculio larvae (Coleoptera:
Curculionidae) in the northeastern United States. Journal of Economic Entomology 83, 1326–1332.
Malavasi, A., Rohwer, G.G. and Campbell, D.S. (1994) Fruit fly free areas: strategies to develop them. In:
Calkins, C.O., Klassen, W. and Liedo, P. (eds) Fruit Flies and Sterile Release Insect Technique. CRC Press,
Boca Raton, Florida, pp. 165–180.
Mampe, C.D. and Neunzig, H.H. (1967) The biology, parasitism and population sampling of the plum curculio
on blueberry in North Carolina. Journal of Economic Entomology 60, 807–812.
Marini, R.P. and Rossi, D. (1985) A partial economic analysis of three pruning treatments on mature peach
trees. HortScience 20, 242–243.
Marlatt, C.L. (1953) An Entomologist’s Quest; The Story of the San Jose Scale; The Diary of a Trip around the
World 1901–1902. Monumental Printing Co., Washington, DC.
Masoodi, M.A., Trali, A.R., Bhat, A.M., Tiku, R.K. and Neru, R.K. (1989) Establishment of Encarsia (+ Prospal-
tella) perniciosi, a specific parasite of San Jose scale, on apple in Kashmir. Biocontrol 34, 39–43.
Meagher, R.L. and Meyer, J.R. (1990) Influence of ground cover and herbicide treatments on Tetranychus ur-
ticae populations in peach orchards. Experimental and Applied Acarology 9, 149–158.
Molinari, F., Chiappini, E. and Sambado, P. (2005a) Preliminary investigation on the natural enemies of the
peach twig borer, Anarsia lineatella (Zeller) in northern Italy. Bulletin OILB/SROP 28, 135–138.
Molinari, F., Tiso, R., Butturini, A., Ceredi, G., Sambado, P. and Rossi, E. (2005b) A forecasting model for the
peach twig borer, Anarsia lineatella (Zeller). Bulletin OILB/SROP 28, 115–118.
Myers, C.T. and Hull, L.A. (2003) Insect growth regulator impact on fecundity and fertility of adult tufted apple
bud moth, Playnota idaeusalis Walker. Journal of Entomological Science 38, 420–430.
Myers, C.T., Hull, L.A. and Krawczyk, G. (2006) Comparative survival rates of oriental fruit moth (Lepidoptera:
Tortricidae) larvae on shoots and fruit of apple and peach. Journal of Economic Entomology 99, 1299–1309.
Nalepa, C.A. and Meyer, J.A. (1990) The seasonal history of the white peach scale (Homoptera: Diaspididae) and
its hymenopteran natural enemies in North Carolina. Journal of Entomological Science 25, 303–310.
Neven, L.G., Rehfield-Ray, L.M. and Obenland, D. (2006) Confirmation and efficacy tests against codling
moth and oriental fruit moth in peaches and nectarines using combination heat and controlled atmo-
sphere treatments. Journal of Economic Entomology 99, 1610–1619.
Padula, A.L. and Smith, E.H. (1971) Reproductive incompatibility between univoltine males and multivoltine
females of the plum curculio. Annals of the Entomological Society of America 64, 665–668.
Paloukis, S.S. and Navrozidis, E.I. (1997) Integrated control of Pseudaulacaspis pentagona (Targioni-Tozzetti)
(Homoptera, Diaspididae) on peach and kiwi trees in northern Greece. Bollettino del Laboratorio di
Entomologia Agraria ‘Filippo Silvestri’ 52, 111–116.
Pascal, T., Pfeiffer, F., Kervella, J., Lacroze, J.P. and Sauge, M.H. (2002) Inheritance of green peach aphid resis-
tance in the peach cultivar ‘Rubira.’ Plant Breeding 121, 459–461.
Peck, S.L. and McQuate, G.T. (2000) Field test of environmentally friendly malathion replacements to sup-
press Mediterranean fruit fly (Diptera: Tephritidae) populations. Journal of Economic Entomology 93,
280–289.
Pedata, P.A., Hunter, M.S., Godfray, H.C.J. and Viggiani, G. (1995) The population dynamics of the white
peach scale and its parasitoids in a mulberry orchard in Campania, Italy. Bulletin of Entomological Re-
search 85, 531–539.
Pickel, C., Bentley, W.J., Hasey, J.K. and Day, K.R. (2006) UC IPM Pest Management Guidelines: Peach. UC
ANR Publication 3454. University of California, Davis, California.
Potter, D.A. and Held, D.W. (2002) Biology and management of the Japanese beetle. Annual Review of Ento-
mology 47, 175–205.
Quaintance, A.L. (1960) The San Jose Scale and Its Control. USDA Farmers’ Bulletin No. 650. US Department
of Agriculture, Washington, DC.
Quaintance, A.L. and Jenne, E.L. (1912) The Plum Curculio. USDA Bureau of Entomology Bulletin No. 103.
US Department of Agriculture, Washington, DC.
Racette, G., Chouinard, G., Vincent, C. and Hill, S.B. (1992) Ecology and management of plum curculio,
Conotrachelus nenuphar [Coleoptera: Curculionidae], in apple orchards. Phytoprotection 73, 85–100.
Reis, W., Nora, I. and Melzer, R. (1988) Population dynamics of Grapholitha molesta, and its adaptation on
apple in south Brazil. Acta Horticulturae 232, 204–212.
Rice, R.E., Hoyt, S.C. and Westigard, P.H. (1979) Chemical control of male San Jose scale (Homoptera: Dias-
pididae) in apples, pears and peaches. The Canadian Entomologist 111, 827–831.
 Insects and Mites 503

Rieger, M. (2006) Peach. In: Rieger, M. (ed.) Introduction to Fruit Crops, text edn. Haworth Food and Agricul-
tural Products Press, New York, pp. 311–323.
Rings, R.W. (1957) Types and seasonal incidence of stink bug injury to peaches. Journal of Economic Entomol-
ogy 50, 599–604.
Rings, R.W. (1958) Types and seasonal incidence of plant bug injury to peaches. Journal of Economic Ento-
mology 51, 27–32.
Roessler, Y. (1989) Insecticidal bait and cover sprays. In: Robinson, A.S. and Hooper, G. (eds) Fruit Flies, Their
Biology, Natural Enemies and Control. Elsevier, Amsterdam, pp. 329–335.
Rosen, D. and DeBach, P. (1978) Homoptera: Diaspididae. In: Clausen, C.P. (ed.) Introduced Parasites and
Predators of Arthropod Pests and Weeds – A World Review. US Department of Agriculture, Washington,
DC, pp. 78–128.
Rouzet, J., Gendrier, J.P. and Audemard, H. (1995) Lutte par confusion sexuelle contre la tordeuse orientale
Cydia molesta en vergers de pecher dans le sud-est de la France. Bulletin OILB/SROP 18, 1–4.
Sanderson, E.D. and Peairs, L.M. (1921) Tarnished plant bug. In: Insect Pests of Farm, Garden and Orchard.
Wiley, New York, pp. 571–574.
Schlamp, K.K., Brown, K., Gries, R., Hart, M. and Gries, G. (2006) Diel periodicity of sexual communication
in Anarsia lineatella (Lepidoptera: Gelechiidae). The Canadian Entomologist 138, 384–389.
Sciarretta, A. and Trematerra, P. (2006) Geostatistical characterization of the spatial distribution of Grapholita mo-
lesta and Anarsia lineatella males in an agricultural landscape. Journal of Applied Entomology 130, 73–83.
Shapiro-Ilan, D.I. and Cottrell, T.E. (2006) Susceptibility of the lesser peachtree borer (Lepidoptera: Sesiidae) to
entomopathogenic nematodes under laboratory conditions. Environmental Entomology 35, 358–365.
Shapiro-Ilan, D.I., Mizell, R.F. III and Horton, D.L. (2004) Measuring field efficacy of Steinernema feltiae and
Steinernema riobrave for suppression of plum curculio, Conotrachelus nenuphar, larvae. Biological
Control 30, 496–503.
Shearer, P.W. and Usmani, K.A. (2001) Sex-related response to organophosphorus and carbamate insecticides
in adult oriental fruit moth, Grapholita molesta (Busck). Pest Management Science 57, 822–826.
Shetlar, D.J. and Johnson, D. (2005) Japanese beetle. In: Horton, D. and Johnson, D. (eds) Southeastern Peach
Growers Handbook. GES Handbook No. 1. University of Georgia College of Agricultural & Environmen-
tal Sciences, Athens, Georgia, pp. 257–258.
Shukla, D.D., Ward, C.W. and Brunt, A.A. (1994) Plum pox virus. In: Shukla, D.D., Ward, C.W. and Brunt,
A.A. (eds) The Potyviridae. CAB International, Wallingford, UK, pp. 382–385.
Stoetzel, M.B. and Miller, G.L. (1998) Aphids (Homoptera: Aphididae) colonizing peach in the United States
or with potential for introduction. The Florida Entomologist 81, 325–345.
Strand, L. (1999) Integrated Pest Management for Stone Fruits. University of California Statewide Integrated
Pest Management Program. University of California Agricultural and Natural Resources, Publication No.
3389. University of California, Oakland, California.
Stumpf, N. and Nauen, R. (2001) Cross-resistance, inheritance, and biochemistry of mitochondrial electron
transport inhibitor-acaricide resistance in Tetranychus urticae (Acari: Tetranychidae). Journal of Eco-
nomic Entomology 94, 1577–1583.
Tavella, L., Arzone, A., Alma, A. and Galliano, A. (1996) IPM application in peach orchards against Lygus
rugulipennis (Poppius). Bulletin OILB/SROP 19, 160–164.
Tremblay, E. (2000) Entomologia applicata, Vol. IV. Liguori Editore, Naples, Italy, pp. 74–75.
Triplehorn, C.A. and Johnson, N.F. (2005a) Hemiptera: Miridae, Coreoidea, Pentatomidae. In: Triplehorn, C.A.
and Johnson, N.F. (eds) Borror and Delong’s Study of Insects, 7th edn. Thomson Brooks/Cole, Belmont,
California, p. 294, 301, 302–303.
Triplehorn, C.A. and Johnson, N.F. (2005b) Scarabaeidae scarab beetles. In: Triplehorn, C.A. and Johnson, N.F.
(eds) Borror and Delong’s Study of Insects, 7th edn. Thomson Brooks/Cole, Belmont, California, pp.
412–415.
Unwin, B. (1971) Biology and control of the two-spotted spider mite, Tetranychus urticae (Koch). Journal of
the Australian Institute of Agricultural Science 17, 192–211.
Van Duyn, J. and Murphey, M. (1971) Life history and control of white peach scale, Pseudaulacaspis pen-
tagona (Homoptera: Coccoidea). The Florida Entomologist 54, 91–93.
Vasseur, R. and Schvester, D. (1957) Bilogie et ecoloie du pou de San Jose in France. Annales des Epiphyties,
Pathologie, Vegetaux, Zoologie, Agriculture, et Phytopharmacie 8, 5–66.
Wallis, C.M., Fleisher, S.J., Luster, D. and Gildow, F.E. (2005) Aphid (Hemiptera: Aphididae) species composi-
tion and potential aphid vectors of plum pox virus in Pennsylvania peach orchards. Journal of Economic
Entomology 98, 1441–1450.
504 D.L. Horton et al.

Weldon, G.P. (1921) Thrips injury to peaches in southern California. Journal of Economic Entomology 13,
424–428.
Yonce, C.E. (1980) Effectiveness of chlorpyrifos for control of Synanthedon pictipes and S. exitiosa in peach
orchard tests of young trees with emphasis on timing applications. Journal of Economic Entomology 73,
827–828.
Yonce, C.E. and Jacklin, S.W. (1974) Life history of the white peach scale in central Georgia. Journal of the
Georgia Entomological Society 9, 213–216.
Yonce, C.E., Beshear, R.J., Payne, J.A. and Horton, D.L. (1990) Thrips associated with unsprayed and sprayed
peaches in Georgia. Journal of Economic Entomology 83, 511–512.
 19 Nematodes

A.P. Nyczepir1 and D. Esmenjaud2

1USDA-ARS, Southeastern Fruit and Tree Nut Research Laboratory, Byron,
Georgia, USA
2INRA, UMR ‘Interactions Biotiques et Santé Végétale’ (IBSV), 400 Route des

Chappes, F-06560 Sophia-Antipolis Cedex, France

19.1 Introduction 506

19.2 Root-knot Nematodes 506
Species 506
Specific identification and polymorphism 507
Symptoms 507
Biology 508
Survival and dissemination 509
Economic importance 510
Environmental factors 510
Host range 510
Control 510
19.3 Ring Nematodes 515
Species 515
Symptoms 515
Biology 516
Survival and dissemination 517
Economic importance 517
Disease complexes 517
Host range 518
Control 518
19.4 Root-lesion Nematodes 521
Species 521
Symptoms 521
Biology 521
Survival and dissemination 522
Economic importance 522
Disease complexes 523
Host range 523
Control 523
19.5 Dagger Nematodes 524
Species 524
Symptoms 525
© CAB International 2008. The Peach: Botany, Production and Uses
(eds D.R. Layne and D. Bassi) 505
506 A.P. Nyczepir and D. Esmenjaud
Biology
Nepovirus diseases
Survival, dissemination and host range
Economic importance
Methods of diagnosis
Control
19.6 Other Nematodes
19.7 Outlook
19.1 Introduction
Nematodes are microscopic, true roundworms
that are non-segmented and have bilateral
symmetry and no appendages (Hirschmann,
1971). Most plant-parasitic nematodes are
generally thread-like in shape (at least during
one of their life stages) and range from 0.4 mm
(e.g. Meloidogyne, Mesocriconema and Praty-
lenchus spp.) to 5 mm in length (e.g. Xiphinema
spp.). Plant-parasitic nematodes are associ-
ated with many forms of plant life including
orchard crops such as peach (Prunus persica
(L.) Batsch). Due to their soil localization and
their non-specific above-ground symptoms,
the presence of nematodes and their economic
importance to the fruit industry are frequently
ignored and underestimated.
Nematodes, if not managed, can cause
some of the most important rootstock dis-
eases of peach, contributing to reduced yield
and vigour and even tree death. Successful
nematode management in peach begins with
site selection. A preferred site is one which is
suitable for peach culture and does not have a
history of stone fruit or nematode problems.
If nematode-free sites are not available, then
nematode problems need to be identified and
proper management practices implemented.
Recommended control practices include pre-
plant and post-plant nematicide application,
resistant rootstocks, cultural practices and bio-
logical control agents, when available. Addi-
tionally, proper sanitation is recommended to
prevent reinfestation of treated sites and
nematode-free and certified virus-free root-
stocks should be planted to circumvent any
future problems.
There are several nematode genera known
to be associated with causing severe losses
in peach orchards worldwide. This chapter
525
525
525
525
526
526
526
526
focuses on four major nematode pests of peach:
root-knot (Meloidogyne spp.), ring (Mesocriconema
spp.), root-lesion (Pratylenchus spp.) and dag-
ger (Xiphinema spp.) nematodes.
19.2 Root-knot Nematodes
Root-knot nematodes, Meloidogyne spp., are
considered to be the most damaging nema-
todes in the world and are distributed in the
temperate, tropical and equatorial areas (Lam-
berti, 1979; Sasser, 1979; Sasser and Freckman,
1987). Root-knot nematodes reduce fruit pro-
duction in several economically important
Prunus species, including peach.
Species
The three predominant root-knot nematode
species affecting Prunus species are Meloidog-
yne arenaria, Meloidogyne incognita and Meloid-
ogyne javanica, all of which feed on many hosts
(polyphagous) and reproduce asexually via
parthenogenesis. The northern root-knot
nematode, Meloidogyne hapla, develops poorly
on Prunus spp. (Esmenjaud et al., 1994). A
more recently described species, Meloidogyne
hispanica (Hirschmann, 1986; Esmenjaud et al.,
1994), appears to be restricted to the southern
part of the Iberian Peninsula, but is destructive
when present (C. Scotto La Massèse, France,
1992, personal communication). As with other
plant species, host suitability tests performed
on Prunus rootstocks in different countries
show high variations in aggressivity among
the populations (Pinochet et al., 1989, 1992;
Marull and Pinochet, 1991; Esmenjaud et al.,
1994; Fernández et al., 1994a).
 Nematodes
A less prominent Meloidogyne sp. was
reported on peach in Florida in the 1960s
(Sharpe and Perry, 1967; Sharpe et al., 1969) as
overcoming the resistance of ‘Nemaguard’
and ‘Okinawa’ rootstocks (Fig. 19.1/Plate
217). This nematode had initially been identi-
fied as M. incognita race 3 (Sherman et al.,
1981; Sherman and Lyrene, 1983) and pro-
duces smaller galls that do not coincide and
are more aligned on the rootlets than those of
the parthenogenetic species. Recently this iso-
late was described as a new species and
named the ‘peach root-knot nematode’, Melo-
idogyne floridensis Handoo (Handoo et al.,
2004). From phylogenetic analysis, M. floriden-
sis was intermediate between two distinct
and individualized groups: M. arenaria, M.
incognita and M. javanica, on the one hand and
M. hapla on the other. Although the distribu-
tion of this new species is unknown (Nyczepir
et al., 1998b), M. floridensis has been detected
in several areas in Florida (W. Sherman, Flor-
ida, 2004, personal communication).
Specific identification and polymorphism
Species identification among the partheno-
genetic root-knot nematodes cannot be differ-
entiated on the basis of below-ground root
Fig. 19.1. Galls on ‘Okinawa’ (A) and ‘Nemaguard’ (B) peach roots caused by Meloidogyne floridensis.
(Courtesy of W.B. Sherman, Gainesville, Florida, USA.)
507
galling symptoms. Therefore, integrated tools
were developed for nematode identification
such as morphological and morphometrical
characters (Golden, 1976; Eisenback et al.,
1981) in conjunction with electrophoresis (i.e.
esterases) (Janati et al., 1982; Esbenshade and
Triantaphyllou, 1985, 1990). The study of the
nucleus has shown high variations in the num-
ber of chromosomes due to polyploidy and
aneuploidy (Triantaphyllou, 1971, 1985), sug-
gesting that each species is composed of clones.
Moreover, a low genetic polymorphism was
observed among these species, which is related
to their parthenogenetic reproduction (Dalmasso
and Berge, 1975, 1978). Molecular data have con-
firmed the previous separation of species (Piotte
et al., 1992; Castagnone-Sereno et al., 1993; Pow-
ers and Harris, 1993; Petersen et al., 1997; Ran-
dig et al., 2001) and simplified molecular
diagnosis is now possible by PCR using specific
primers from ribosomal genes (Zijlstra, 1997;
Zijlstra et al., 2000) that allows direct identifica-
tion of M. arenaria, M. incognita and M. javanica
from a single juvenile nematode (i.e. J2).
Symptoms
The nematodes belonging to this genus
have a sedentary endoparasitic feeding habit.
508 A.P. Nyczepir and D. Esmenjaud
Attacks by root-knot nematodes on peach often
cause typical below-ground root galls (Fig. 19.2/
Plate 218) and the associated above-ground
stunted growth of young peach trees. Attacks
during early root development may lead to
extensive tree death across an orchard. Addi-
tional above-ground symptoms include a
reduction in tree vigour and early defoliation,
which ultimately results in yield reduction.
Symptom expression is enhanced in sandy
soils, especially when trees are exposed to
drought conditions. Extensive galling has some-
times been observed without associated reduc-
tions in tree vigour or stunted growth (Bertrand,
1985). In such cases, it is believed that the
orchard was established on a site with low ini-
tial soil population levels of Meloidogyne, thus
allowing the tree to develop a more extensive
root system.
Biology
Meloidogyne spp. are extremely polyphagous
and can parasitize hundreds of plant species
(a) (b)
(c)
from numerous botanical families (de Guiran
and Netscher, 1970; Sasser, 1979). M. arenaria,
M. incognita and M. javanica are mitotic and
reproduce exclusively by parthenogenesis
(Triantaphyllou, 1971, 1985), and have thus
been designated as the parthenogenetic Melo-
idogyne complex. The reproductive life cycle
is summarized in Fig. 19.3/Plate 219. Eggs
are deposited in gelatinous egg masses and
can survive in the soil or on plant residues.
The mobile second-stage juveniles (J2) hatch
directly from the egg and move to root tips,
where they penetrate and migrate intercellu-
larly within root tissues. The J2 initiate feed-
ing cells in the region of what will become the
vascular area, where they develop into swol-
len third- and fourth-stage juveniles. They
finally develop into sedentary adult females
which can each lay several hundred eggs in a
single egg mass, usually positioned exterior
to the root surface. Adult females feed with
their protruded stylet, which perforates the
cell wall but not the cell plasma membrane.
Feeding juveniles and adults induce the for-
mation of a few ‘giant cells’ (‘feeding sites’)
Fig. 19.2. Galls on peach (a and b)
and Myrobalan plum (c) roots caused
by root-knot nematode (RKN), Meloid-
ogyne sp. Severe early symptoms (a);
root decay following an attack (b);
two Myrobalan plum individuals from
a progeny segregating for the Ma
RKN resistance gene (left, host; right,
resistant) (c). (Photos by A.P. Nyczepir
(a) and D. Esmenjaud (b,c).)
 Nematodes
Fig. 19.3. Reproductive life cycle of Meloidogyne spp. (N = female nematode; *= giant cells).
(Courtesy of INRA, Sophia-Antipolis, France.)
around the nematode head along with the
multiplication of cells surrounding cortical
and vascular parenchyma. Nutrient uptake
by root-knot nematodes suppresses plant
development.
The parasitism of the plant by the root-
knot nematode described above is a compati-
ble interaction. In certain plant species,
Meloidogyne do not develop on all accessions
and an incompatible interaction results. In
Prunus, resistant rootstocks have been detected
but few studies have been conducted on the
characterization of the resistance mechanisms
involved. For instance, in Myrobalan plum, M.
arenaria J2 penetrated resistant and host acces-
sions (carrying or lacking, respectively, the
Ma gene for resistance to Meloidogyne spp.)
(see section on ‘Resistance’ below) and
showed an equivalent early penetration of
the nematodes into the roots during the 48 h
after their inoculation into the soil. The J2
numbers increased rapidly in the host acces-
sion, but decreased slowly in the resistant
accessions (Voisin et al., 1999). The results
substantiate the hypothesis that parasitized
cells of the resistant accessions express a
hypersensitive reaction to the juveniles. In
contrast, a more delayed resistance reaction
termed ‘walling off’ allows the J2 to penetrate
509
roots and form small galls, but prevents mat-
uration and reproduction. Such a phenome-
non was observed with M. javanica in the
resistant peach stocks of ‘Nemared’ and ‘Oki-
nawa’ (Malo, 1967; Marull et al., 1994) and
with M. incognita in ‘Guardian®’ (Nyczepir
et al., 1999).
Survival and dissemination
Root-knot nematodes survive as eggs grouped
in egg masses. At this stage, they may survive
for several years in the soil under Mediterra-
nean and temperate conditions. This persistence
during unfavourable conditions is attributed to
an egg diapause (de Guiran, 1979; de Guiran
and Villemain, 1980). These polyphagous
nematodes can also develop on numerous
annual crops including weed species associ-
ated with the orchard floor (de Guiran and
Netscher, 1970; Sasser, 1979). Nematodes are
readily disseminated by the use of nematode-
infected nursery stock. Seedlings or rooted
cuttings constitute the most effective long-
distance way of disseminating these pests.
Because of the increasing exchange of plant
material at national and international levels,
510 A.P. Nyczepir and D. Esmenjaud
populations originating from distant geo-
graphic origins are expected to be extensively
mixed. Nevertheless, such dissemination
should be limited by the increased use of cer-
tified nematode-free plant material.
Economic importance
Peach production is negatively influenced by
the presence of Meloidogyne spp. In South
Carolina, USA the use of either a pre-plant
fumigant or ‘S-37’ resistant rootstock increased
peach yields in Meloidogyne-infested soil com-
pared with the untreated check (Foster et al.,
1972). In Georgia, USA, pre-plant fumigation
with methyl bromide to control Meloidogyne
spp. increased yield in the third to fifth grow-
ing seasons of ‘Redhaven’ on ‘Lovell’ root-
stock by 2535 kg/ha (Sharpe et al., 1993). In
Spain as well as in North Africa, root-knot
nematodes are involved in replant disease
and tend to be a serious problem in warm,
well-drained sandy soils. The development of
specific replant problems caused by root-knot
nematodes in these regions has been attrib-
uted to the high degree of rootstock suscepti-
bility to Meloidogyne spp. (Calvet et al., 2000).
Environmental factors
Meloidogyne spp. are very damaging pests of
stone fruits especially when cultivated on
sandy soils (Stirling, 1975). Plant damage is
increased under drought conditions and high
soil temperature favours nematode parasit-
ism of the rootstock (Canals et al., 1992;
Fernández et al., 1994b). In water-saturated
soils, hatching of juveniles is inhibited (Baxter
and Blake, 1969).
Nematodes have been shown to facilitate
penetration of the bacterium Agrobacterium
tumefaciens into the root tissues and the subse-
quent development of symptoms of crown
gall in almond (Orion and Zutra, 1971), peach
(Dhanvantari et al., 1975) and plum (Rubio-
Cabetas et al., 2001) (Fig. 19.4/Plate 220).
M. incognita in combination with Fusarium
oxysporum has been shown to reduce growth
of peach seedlings more than either organism
alone (Wehunt and Weaver, 1972).
Host range
Root-knot nematodes parasitize a wide range of
plant species that are either crop plants (annu-
als or perennials) or weeds. Among Prunus,
rootstocks may express differing host suscepti-
bilities such as those summarized in Table 19.1.
Control
Control measures must be applied against
root-knot nematodes in both the plant and
soil. After their introduction into the orchard,
eradication of nematodes from soil is difficult.
Therefore, management strategies rely on the
combination of different methods to keep the
nematode density below economic damage
levels (Bernhard et al., 1985). Pre- and post-
plant chemical nematicides have been shown
to be effective tools in managing root-knot
nematodes in peach. However, because most
of the nematicides are being progressively
removed from the market, alternatives to
chemical control methods are encouraged
(Batchelor, 2002). For example, methyl bro-
mide has been associated with atmospheric
ozone depletion, which has resulted in a ban
(according to the 1992 Montreal Protocol) on
its importation and manufacture in the USA
and Western Europe since January 2005
(Clean Air Act, 1990). Additionally, the manu-
facturer of fenamiphos, the one remaining
recommended post-plant nematicide on
peach in the south-eastern USA, was volun-
tarily cancelled of all product registrations
effective from 31 May 2007 (Anon., 2003).
Nematicides
Root-knot nematodes can be controlled by pre-
plant soil fumigation plus post-plant applica-
tion of non-fumigant nematicides. Because
state/country recommendations for nematicide
use are not the same and change frequently,
current local sources of information need to be
consulted for updated recommendations.
Rotation
It is recommended to plant a non-host such as
cereal grains or leave the soil fallow prior to
 Nematodes
planting a peach orchard in order to suppress
nematode populations to undetectable levels
(McKenry, 1985). However, weeds also must
be controlled because many of them are good
hosts for root-knot nematodes, thus allowing
them to survive between successive orchards
(McKenry, 1985; Nyczepir and Halbrendt,
1993).
Resistance
The most economic and environmentally
sound method for managing Meloidogyne in
Prunus species is the use of resistant root-
stocks (Cook and Evans, 1987; Layne, 1987).
The search for peach rootstock resistance
began in the USA (Tufts, 1929; Day and Tufts,
1939; Weinberger et al., 1943) but has since
been investigated and documented in other
countries (Kochba and Spiegel-Roy, 1976;
Scotto La Massèse et al., 1984; Fernández et al.,
511
Fig. 19.4. Galls on Myrobalan plum
root caused by the association of
Meloidogyne spp. and Agrobacterium
tumefaciens (crown gall). (Photo by
D. Esmenjaud.)
1994a; Pinochet et al., 1999). A summary of
Prunus rootstock reactions to Meloidogyne
spp. is reported in Table 19.1.
The impact of root-knot nematodes on
peach has justified specific breeding efforts.
In the USA, breeding programmes have focused
mainly on the creation of peach or peach–
almond rootstocks using peach germplasm
(such as ‘Nemaguard’ and ‘Okinawa’) as the
resistance sources. For example, ‘Nemared’
(Ramming and Tanner, 1983) used ‘Nema-
guard’ as its source of resistance, whereas
‘Hansen 2168’ and ‘Hansen 536’ (Kester and
Asay, 1986) and ‘Flordaguard’ (Sherman et al.,
1991) both relied on ‘Okinawa’ and Prunus
davidiana. The source of root-knot nematode
resistance in ‘Guardian®’ comes from ‘Nema-
guard’ and ‘S-37’ (Okie et al., 1994a). In Europe,
an extensive evaluation of known or new
germplasm sources that include many new
selections of peach-based hybrids has been
512 A.P. Nyczepir and D. Esmenjaud

Table 19.1. Reaction of rootstocks used for peach to root-knot (Meloidogyne spp.), root-lesion
(Pratylenchus spp.) and ring (Mesocriconema xenoplax) nematodes. (From Westcott and Zehr, 1991;
Pinochet et al., 1992; Nyczepir and Halbrendt, 1993; Okie et al., 1994b; Alcaniz et al., 1996; Esmenjaud
et al., 1997; Nyczepir et al., 1999; Nyczepir and Beckman, 2000; Nyczepir and Pinochet, 2001;
D. Esmenjaud, France, 2007, personal communication.)

Nematodesa

Species/accession Origin Propagation MA MI MJ MF Pp Pv Mx

Peach
Prunus persica
‘Bokhara’ USA Seedling Rb R H
‘B-S6’ Italy Seedling MR?
‘Cadaman’ France Seedling R R R H
‘Elberta’ Australia Seedling H H H H
‘Flordaguard’ USA Seedling R H
‘GF 305’ France Seedling R/H R/H H H H H H
‘Guardian®’ USA Seedling R R/H H T H T
‘Harrow Blood’ Canada Seedling H H H H
‘Lovell’ USA Seedling H H H H H H
‘Missour’ Tunisia/ Seedling H
Morocco
‘Montclar’ France Seedling H H H
‘Nemaguard’ USA Seedling R R R/H H H H H
‘Nemared’ USA Seedling R R R/H H H
‘Okinawa’ USA Seedling R R R H H
‘Rancho Resistant’ USA Seedling R H H
‘Rubira’ France Seedling H H H
‘Rutgers Red Leaf’ USA Seedling H H/T H H
‘S-37’ USA Seedling R H H
‘Shalil’ USA Seedling R H H
‘Siberian C’ Canada Seedling H H
‘Yunnan’ USA Seedling R H H

Almond–peach hybrids
Prunus dulcis × P. persica
‘Adafuel’ Spain Cutting H H H H H
‘Alcaniz’ Spain Cutting H H H
‘Bergasa’ Spain Cutting H H
‘Cachirulo’ Spain Seedling R H
‘Felinem’ Spain Cutting R R R/H H
‘Fermoselle’ Spain Cutting H H H
‘Garnem’ Spain Cutting R R R/H H
‘GF 557’ France Cutting R R H H
‘GF 677’ France Cutting H H H H H
‘Hansen 2168’ USA Cutting R R R
‘Hansen 536’ USA Cutting R R R
‘Titan’ (hybrids) USA Seedling R R R H
P. persica × Prunus belsiana
‘Citation’ USA Cutting R H

Plums
Prunus cerasifera
‘Myrabi’ France Cutting H H H H
‘Myrobalan P-2175’ France Cutting R R R R

(Continued)
 Nematodes 513

Table 19.1. continued

Nematodesa

Species/accession Origin Propagation MA MI MJ MF Pp Pv Mx

‘Myrobalan P-2980’ France Cutting R R R R

‘Myrobalan 29C’ USA Cutting R R H
‘Myrobalan franc’ France Seedling H H H
‘Myrobalan 605’ Spain Cutting H H H
‘Myrobalan B’ UK Cutting MR
P. cerasifera × Prunus salicina
‘GF 31’ France Cutting R H
P. cerasifera × Prunus
munsoniana
‘GF 8-1’ France Seedling R R R
‘Marianna 2624’ USA Cutting R R R H
‘Redglow’ USA Cutting MR
Prunus domestica
‘Brompton’ UK Cutting R R R
‘Damas’ France Cutting R R R
‘GF 43’ France Cutting H H H
‘Torinel’ France Cutting R R R MR
P. domestica × Prunus spinosa
‘GF 1869’ France Cutting R
Prunus insititia
‘Montizo’ Spain Cutting R R H
‘PSM 101’ Spain In vitro R R H
‘Saint Julien 655-2’ France Cutting R R R H
aMA, Meloidogyne arenaria; MI, Meloidogyne incognita; MJ, Meloidogyne javanica; MF, Meloidogyne
floridensis; Pp, Pratylenchus penetrans; Pv, Pratylenchus vulnus; Mx, Mesocriconema xenoplax.
bR, resistant; MR, moderately resistant; H, host; T, tolerant.

performed to characterize resistance in parent
material according to their: (i) spectrum of
activity (Marull et al., 1994; Pinochet et al.,
1996a); (ii) tolerance to high population levels
(Esmenjaud et al., 1996a,b, 1997); (iii) heat sta-
bility (Canals et al., 1992; Fernández et al.,
1994b); and (iv) influence of age of the plant on
resistance expression (Fernández et al., 1995).
Resistance in tested peach and almond
cultivars is not complete and does not control
M. floridensis (Esmenjaud et al., 1997) (Table 19.2).
However, ‘Flordaguard’ has been reported to
be tolerant to this nematode, but its spectrum
of genetic resistance is currently unknown
(Sherman et al., 1991). In contrast, Myrobalan
plum (Prunus cerasifera from the subgenus
Prunophora, grouping plums and apricot)
comprises certain clones that also resist M.
floridensis and carry a complete spectrum and
high level of resistance to other Meloidogyne
spp. (Esmenjaud et al., 1996a, 1997). Some of
these clones impart favourable agronomic
features for peach scions such as broad graft
compatibility and high adaptation to water-
logged soils (Kester and Grasselly, 1987; Layne,
1987; Salesses et al., 1992). In Myrobalan plum,
resistance to M. arenaria, M. incognita, M.
javanica and M. floridensis is conferred by the
single major gene Ma, which behaves as com-
pletely dominant in the accessions ‘P-2175’
(allele Ma1, heterozygous) (Esmenjaud et al.,
1996b; Lecouls et al., 1997) and ‘P-2980’ (allele
Ma3, heterozygous) (Rubio-Cabetas et al.,
1998) (Table 19.2). This gene also controls Mel-
oidogyne mayaguensis (Rubio-Cabetas et al.,
1999), a species of tropical origin that over-
comes the resistance conferred by the Mi gene
from tomato (Rammah and Hirschmann, 1988;
Fargette et al., 1996), and M. hispanica (Stalin
et al., 1998).
514 A.P. Nyczepir and D. Esmenjaud

Table 19.2. Spectrum of resistance of main sources to root-knot nematodes used in Prunus rootstock
breeding.

Resistance status toa

Resistance gene
Accession MA MI MJ MF and genotype References

Amygdalus
Peach (Prunus persica)
‘Shalil’
‘GF 557’ RMia557 gene Esmenjaud et al.
(= almond × ‘Shalil’) controlling MA and MI (1994, 1997)
‘GF 557’ Rc R H H (RMia557/rMia557) Claverie et al. (2004)
‘Nemaguard’ RMiaNem gene
controlling MA and MI
‘Nemaguard’ R R R/Hd H (RMiaNem/RMiaNem) Esmenjaud et al. (1997)
‘Nemared’ R R R/H H (RMiaNem/RMiaNem) Claverie et al. (2004)
‘Rubira’b H H H H (rMiaNem/rMiaNem)
Prunophora
Myrobalan plum Ma gene controlling
(Prunus cerasifera) MA, MI, MJ and MF
‘P-2175’ R R R R (Ma1/ma) Esmenjaud et al. (1994;
‘P-2980’ R R R R (Ma3/ma) 1996b, 1997); Lecouls
‘P-2032’b H H H H (ma/ma) et al. (1997);
Rubio-Cabetas et al.
(1999)
aMA, Meloidogyne arenaria; MI, Meloidogyne incognita; MJ, Meloidogyne javanica; MF, Meloidogyne
floridensis.
bHost control accession.
cR, resistant; H, host.
dR/H, variable behaviour in function of M. javanica isolates.

In ‘Shalil’ and ‘Nemared’ peach, the genet-
ics of resistance is still not clear and at least
one major gene for resistance to both M. are-
naria and M. incognita, designated RMia, has
been reported and shown to be independent
from the Ma gene (Claverie et al., 2004). Yama-
moto and Hayashi (2002) have also reported
two tightly linked genes controlling either M.
incognita alone (Mia) or M. javanica (Mja) in
the Japanese accession ‘Juseitou’. The SCAR
(sequence characterized amplified regions)
markers designed by these authors for Mia
were closely linked to the RMia gene (Claverie
et al., 2004). Consequently, the RMia and Mia
genes from peach are presumably the same.
In contrast, the corresponding Mi (for M.
incognita) and Mij genes (for both M. incognita
and M. javanica) obtained by Lu et al. (1999,
2000) in ‘Nemared’ were only found distantly
linked to RMia (Claverie et al., 2004).
Based on the Ma and RMia genes, a breed-
ing programme for new Meloidogyne-resistant
rootstocks for stone fruits is being developed
in France in collaboration with Spain and
Italy. Interspecific rootstocks of the type
Myrobalan × peach, Myrobalan × almond or
Myrobalan × almond–peach have been cre-
ated in order to pyramid at least one Ma allele
and one resistance gene from peach. Their
selection is in progress for complementary
adaptative characteristics putatively inher-
ited from their parents (tolerance to waterlog-
ging, drought and chlorosis). Several SCAR
markers co-segregate with the Ma gene and
are available for marker-assisted selection for
all interspecific crosses carrying this gene
(Lecouls et al., 1999, 2004; Dirlewanger et al.,
2004). This development can enable highly
efficient procedures for evaluating woody
plants for Meloidogyne spp. resistance. The Ma
 Nematodes
gene has also been demonstrated to have a
protective effect against A. tumefaciens when
its expression follows root-knot nematode
attacks (Rubio-Cabetas et al., 2001). This pro-
tective effect of Ma and presumably of other
root-knot nematode resistance genes against
Meloidogyne-transmitted crown gall is an addi-
tional argument for their introgression into
Prunus rootstocks. Additional peach rootstock
programmes that evaluate germplasm for
resistance to root-knot nematodes using dif-
ferent selection criteria are currently under
way in other countries (Reighard, 2002).
19.3 Ring Nematodes
Species
Ring nematodes are widely distributed through-
out the world with certain species considered
to be economically important to the peach
industry. The two most important ring nema-
tode species associated with peach decline
diseases are Mesocriconema xenoplax (Raski)
Loof & de Grisse and Mesocriconema curvatum
(Raski) Loof & de Grisse (Nyczepir, 2001;
Nyczepir and Becker, 1998). Of less impor-
tance is Mesocriconema rusticum (Micoletzky)
Loof & De Grisse, which Jaffee et al. (1987b)
reported on peach in Pennsylvania, USA.
515
Symptoms
Ring nematodes are ectoparasites. ‘Nema-
guard’ peach parasitized by M. xenoplax was
characterized by a lack of feeder roots
(Fig. 19.5/Plate 221) (Nyczepir et al., 1987).
Extensive pits and lesions were observed on
cultured peach seedling roots parasitized by
M. curvatum (Hung and Jenkins, 1969).
In the south-eastern USA, peach trees
parasitized by M. xenoplax are predisposed to
cold injury and bacterial canker infection
(Pseudomonas syringae pv. syringae van Hall)
or a combination of both. This disease com-
plex is termed peach tree short life (PTSL)
(Brittain and Miller, 1978; Nyczepir, 1990).
Late winter/early spring temperature fluctu-
ations are associated with PTSL. In California,
M. xenoplax also predisposes peach trees to
bacterial canker infection, a disease complex
termed bacterial canker complex (BCC)
(McKenry, 1989) (see section on ‘Disease com-
plexes’ below).
Typical above-ground PTSL symptoms
commonly occur when trees are 3–6 years of
age, although younger or older trees have
been reported to be affected. Symptoms occur
suddenly in the spring with developing
leaves collapsing as if deprived of water, ulti-
mately resulting in premature death of the
tree. One or two branches may be stricken,
Fig. 19.5. Influence of Mesocriconema
xenoplax (Mx) on ‘Nemaguard’ peach
feeder root growth after 6 months ((–)
Mx, uninoculated; (+) Mx, inoculated,
Pi = 14,000 nematodes/1500 cm3 soil,
where Pi is initial nematode population
density). (Photo by A.P. Nyczepir.)
516 A.P. Nyczepir and D. Esmenjaud

but frequently the entire tree collapses (Brit-
tain and Miller, 1978). In severe cases, neither
flower nor leaf buds open at all.
Bacterial canker infection is generally
linked with delay of bloom or foliation of
individual limbs, which usually die during
the summer. If infection is severe, entire trees
may collapse with branches exhibiting alter-
nate zones of darkened and healthy tissue
with the presence of a sour sap odour (Fig. 19.6/
Plate 222). Since the soil buffers low tempera-
ture fluctuations, dead trunk tissue usually
does not extend below the soil line, thus leav-
ing the primary root system alive (Fig. 19.7/
Plate 223). Suckers are usually produced at
the tree crown during summer, resulting from
the live primary root system (Fig. 19.6/Plate
222). In BCC, limbs and entire trees infected
by P. syringae die in the spring.
With the presence of cold injury, trees
begin to leaf out until additional water is
required by the tree. It is at that time the leaves
collapse and the bark may crack and separate
from the scaffold limbs and tree trunk.
Shepard et al. (1999) reported that M.
xenoplax also was responsible for predisposing
peach trees (cv. ‘Suwanee’) to bacterial spot
(Xanthomonas arboricola pv. pruni) (see Chap-
ter 16). Trees growing in M. xenoplax-infested
soil exhibited more severe bacterial spot dam-
age than trees in soil where the nematode
populations had been suppressed.
Biology
M. xenoplax completes its life cycle in 25–34
days at 22–26°C, under laboratory conditions
(Seshadri, 1964). Nematode life stage devel-
opment requires 11–13 days for egg, 3–5 days
for J2, 4–7 days for J3, 5–6 days for J4, and 2–3
days for adult. Gravid females lay 8 to 15 eggs
over a 2–3-day period. Eggs are deposited in
close proximity to host roots or on the root
surface. Sixty-six per cent of M. xenoplax eggs
were observed to hatch between 13°C and
32°C, and aborted at temperatures above
32.5°C (Westcott and Burrows, 1991). Repro-
duction is presumed to be primarily by par-
thenogenesis, since males are rarely found.
Feeding is required for juveniles to moult and

Fig. 19.6. Dead peach tree with suckers at the crown during summer in the presence of
Mesocriconema xenoplax and the peach tree short life disease complex. (Photo by A.P. Nyczepir.)
 Nematodes 517

Fig. 19.7. Typical cambial tissue damage of the trunk above the soil line and healthy viable tissue
below the soil line in tree dying from peach tree short life disease. (Photo by A.P. Nyczepir.)

for oocyte maturation in adult females. Feed- to transplants. M. xenoplax generally prefers
ing activity of M. xenoplax occurs along peach woody perennials, but this nematode can also
roots and root tips (Thomas, 1959). Unlike survive on weeds found in peach orchards in
other ectoparasitic feeders, this ring nema- the south-eastern USA (Seshadri, 1964; Zehr
tode was reported to feed from a single root et al., 1990).
cortical cell for up to 8 days with no necrotic
tissue development at the feeding site (West-
cott and Hussey, 1992). Individual cortical cells Economic importance
were modified by M. xenoplax into discrete
‘food cells’ in order to sustain ingestion of food
Tree loss due to PTSL varies among years.
for long periods of time (Hussey et al., 1992).
Miller (1994) estimated losses during 1980–
M. xenoplax reproduces faster in sandy
1992 as over $6 million per year in South
soils than in loam or silty loam soils and pre-
Carolina alone. Managing M. xenoplax on a
fers soil temperatures ranging from 22°C to
known PTSL site, following pre-plant methyl
26°C (Seshadri, 1964; Stirling, 1975). Ring
bromide fumigation, in a test orchard dur-
nematode populations also were reported to
ing its third to fifth year of production
be higher in wetter soils (i.e. 15.5% moisture)
resulted in 20% greater yields in fumigated
than in soils that were drier (i.e. 11.6% and
compared with unfumigated soil (Sharpe
7.8% moisture) (Seshadri, 1964).
et al., 1993).

Survival and dissemination

Unlike root-knot nematode, dissemination of
ring nematodes occurs primarily via infested
soil transported on farm equipment and the
feet of animals, in water, and in soil clinging
Disease complexes
Since the late 1600s, the unsatisfactory perfor-
mance of trees grown on sites where peach
trees were previously removed, commonly
518 A.P. Nyczepir and D. Esmenjaud
referred to as replant sites, has plagued peach
growers (Brittain and Miller, 1978). In the
south-eastern USA a specific type of disease
complex problem exists which has been
termed PTSL (see above). It has been clearly
demonstrated through long-term research
that the ring nematode, M. xenoplax, is a key
biotic component of the PTSL disease com-
plex (Nyczepir et al., 1983). Equally important
was the discovery that other plant-parasitic
nematodes such as the root-knot nematode,
M. incognita, do not play a role in PTSL tree
death (Nyczepir et al., 1997). Furthermore, it
was established that a reduction in tree
growth was more severe with trees growing
in the presence of both of these nematode
species. M. incognita appears to be the more
dominant nematode species in this interac-
tion and is a stronger competitor than M.
xenoplax for food on ‘Lovell’ peach roots.
Severity of PTSL in orchards infested
with M. xenoplax increases and orchard
lifespan decreases with each successive peach
planting (Nyczepir and Okie, 1996). In a fol-
low-up study, Nyczepir et al. (2004a) were
able to create a PTSL site on land with no
known history of peach production, 3 years
after M. xenoplax introduction. It appears that
development of PTSL varies with exposure
of trees to the cumulative population of M.
xenoplax.
In South Africa, peach trees parasitized by
a concomitant infestation of M. xenoplax and
M. javanica defoliated 3 to 4 months before
natural leaf drop (Hugo and Meyer, 1995). In
California, M. xenoplax has been associated
with increasing the susceptibility of peach
trees to BCC (McKenry, 1989). An important
distinction between PTSL and BCC is that cold
injury is not typically associated with bacterial
canker tree death in California, even though
BCC is more likely to occur in cold spots within
an orchard. It is also important to note that the
California ‘replant problem’ as described by
McKenry (1999) is not the same as the PTSL or
BCC disease complexes described above. This
‘replant problem’, which is known to occur
worldwide, is associated with leaf yellowing
and tree stunting following the replanting of a
site within several years of removing the pre-
vious orchard (http://www.uckac.edu/nema-
tode/).
Host range
M. xenoplax prefers woody perennials to annu-
als, with some exceptions (Seshadri, 1964;
Zehr et al., 1986, 1990). Peach rootstock reac-
tion to M. xenoplax is summarized in Table
19.1. All tested materials are hosts to the nem-
atode, except ‘Guardian®’ peach, which is
tolerant.
Control
Nematicides
Ring nematodes can be controlled by pre-plant
soil fumigation and post-plant application of
non-fumigant nematicides. Because state/
country recommendations for nematicide use
are not the same and change frequently, cur-
rent local sources of information need to be
consulted for updated recommendations.
Under South Carolina, USA field condi-
tions, a nematicide treatment threshold of >50
M. xenoplax/100 cm3 soil is recommended in
peach orchards for prolonging tree life on
PTSL sites (E.I. Zehr, South Carolina, 2005,
personal communication). In Georgia, the
nematicide treatment threshold is ≥1 M.
xenoplax/100 cm3 soil (Nyczepir and Hal-
brendt, 1993; Davis et al., 1996). In North Car-
olina, it was estimated that PTSL tree death
was likely at average cumulative population
densities of 38–83 M. xenoplax/100 cm3 soil
(Ritchie, 1988). It is important to note that
extraction efficiency of M. xenoplax from
sandy soils can be improved if the dry soil is
first moistened and allowed to stand for up
to 7 days before extracting the nematode
(Lawrence and Zehr, 1978).
Cultural practices
In the early 1970s, a 10-Point Management
Program was developed and recommended
to peach growers in the south-eastern USA to
help reduce tree losses from PTSL (Brittain
and Miller, 1978; Ritchie and Zehr, 1995). It is
important to note that this programme is not
a cure for PTSL, but rather a management
scheme for the disease complex. One of the
major points to this programme is pre- and
 Nematodes
post-plant soil fumigation as a management
strategy for M. xenoplax. However, a contin-
ued reduction in the availability of chemical
nematicides as a result of apprehension asso-
ciated with potential environmental problems
is leaving growers with fewer nematode man-
agement options (e.g. methyl bromide and
fenamiphos – see control of root-knot nema-
todes above). Therefore, finding a cost-effec-
tive and environmentally safe alternative to
chemical control of ring nematodes is war-
ranted.
Alternatives to chemical control for nem-
atode management have been explored over
the past 14 years with some promising results.
Several ground covers appear promising as
either pre-plant or post-plant management
strategies for M. xenoplax. Nimblewill (Muhlen-
bergia schreberi J. Gmelin) planted around
peach trees suppressed populations of M.
xenoplax without being highly competitive for
nutrients and water (Meyer et al., 1992; Nyc-
zepir and Bertrand, 2000). Two disadvantages
in using nimblewill are that: (i) it supports
reproduction by M. javanica and M. arenaria,
and to a lesser extent M. incognita (McKenry,
1990; A.P. Nyczepir, 1995, personal observa-
tion); and (ii) since it is a cool-season grass,
nimblewill establishment in the south-eastern
USA peach orchards (i.e. Georgia and South
Carolina) has proved difficult (Olien et al.,
1994; A.P. Nyczepir, 2004, personal observa-
tion). In Georgia, chemically mowed bahiagrass
(Paspalum notatum Flugge) sod previously
grown in orchard row middles infested with
M. xenoplax suppressed the nematode popu-
lation density after 3 years (Nyczepir and
Bertrand, 2000). When peach trees were
planted into the killed-sod row middles they
generally grew better in the killed bahiagrass
sod, but tree survival was no different from
that in unfumigated weed ground cover soil
on a PTSL site. Additionally, wheat alone or
double-cropping wheat (Triticum aestivum L.
emend. Thell cv. ‘Stacy’) and sorghum (Sor-
ghum vulgare Pers. cv. ‘NK2660’) for 3 years
was as effective as pre-plant methyl bromide
fumigation in suppressing M. xenoplax popu-
lations and prolonging tree survival on a
PTSL site (Nyczepir and Bertrand, 2000;
Nyczepir, 2003a). In southern Brazil, selected
double-crop rotation schemes which include
519
alternating summer and winter non-host
plants have also shown promise in managing
M. xenoplax and Meloidogyne spp. in peach
orchards and nurseries (Carneiro et al., 1998).
Interplanting wheat (i.e. after orchard estab-
lishment) around newly planted or 4-year-old
well-established peach trees did not suppress
M. xenoplax populations after 3 years (Nyczepir
et al., 1998a). Besides ‘Stacy’ there are other
wheat varieties that suppress ring nematode
populations (Nyczepir, 2003b). Integration of
a 3-year pre-plant wheat rotation in conjunc-
tion with the improved ‘Guardian®’ rootstock
are the basis for developing a non-chemical
pre- and post-plant recommendation for M.
xenoplax management on PTSL sites in the
south-eastern USA.
Biofumigation is a non-chemical approach
of planting various crops which produce
breakdown products upon decomposition
that are toxic to nematodes, weeds and various
soil diseases. Some potential green manure
crops include rapeseed, barley, sudangrass
and velvetbean. When sorghum was used as
a green manure with and without a plastic
tarp, it suppressed M. xenoplax populations in
the early stages of a 5-year experiment. This
suppression, however, did not last as long as
pre-plant methyl bromide fumigation (i.e. 19
versus 24 months, respectively) (Nyczepir
and Rodriguez-Kabana, 2007). On the other
hand, double-cropping rapeseed (Brassica
napus L. ‘Humus’) as a green manure and sor-
ghum for 3 years was as effective as pre-plant
methyl bromide fumigation in suppressing
M. xenoplax populations and prolonging tree
survival on a PTSL site (Nyczepir, 2003a).
One matter that every grower must decide
upon when considering utilizing any pre-plant
cultural practice is whether or not that particu-
lar practice can be incorporated into their farm
operation. For the grower with much available
land for peach production this may not be an
issue. However, for the growers who have lim-
ited land suitable for peach production, this
may not be an economically feasible nematode
management approach.
Resistance
‘Lovell’ and ‘Nemaguard’ are two commonly
used peach rootstocks in the USA. Trees on
520 A.P. Nyczepir and D. Esmenjaud

‘Lovell’ generally outlive those on ‘Nema-
guard’ in the south-east, particularly where
M. xenoplax is prevalent; however, both are
subject to cold injury, bacterial canker infec-
tion and eventual PTSL tree death (Sharpe
et al., 1989). ‘Nemaguard’ has been shown to
be a better host to M. xenoplax than ‘Lovell’,
and this may explain why ‘Lovell’ trees sur-
vive longer on PTSL sites (Table 19.1) (Nyc-
zepir, 1990; Beckman et al., 1993). ‘Lovell’
rootstock was the recommended rootstock for
south-eastern USA stone fruit growers for
over 20 years (1973–1994). ‘Guardian®’, a new
commercially available rootstock that toler-
ates M. xenoplax parasitism, increases scion
longevity on PTSL sites compared with ‘Nem-
aguard’ or ‘Lovell’ rootstocks (Fig. 19.8/Plate
224) (Okie et al., 1994a,b). ‘Guardian®’ has
also been reported to be a poor host to some
Meloidogyne spp., but not all (Nyczepir et al.,
1999, 2006; Nyczepir and Beckman, 2000).
Although M. incognita J2 penetrated ‘Guard-
ian®’ roots and formed galls, the majority of
the nematodes failed to mature and repro-
duce (Nyczepir et al., 1999).
Molecular DNA studies are currently
under investigation to determine the genetic
basis of tolerance in ‘Guardian®’ to M. xeno-
plax and how this may relate to prolonged
tree life on PTSL sites (Blenda et al., 2002,
2006).
Biological
The endoparasitic fungus, Hirsutella rhossil-
iensis Minter & Brady, suppressed popula-
tions of M. xenoplax in five of nine controlled
experiments (Eayre et al., 1987). Furthermore,
amending the soil with KCl did not appear to
stimulate fungal parasitism or affect the level
of nematode suppression in most tests.
Kluepfel et al. (2002) reported that a
Pseudomonas sp. (BG33R) isolated from a PTSL
suppressive site in South Carolina suppressed
M. xenoplax reproduction. Under orchard
conditions, ring nematode populations on
newly established peach seedlings inoculated
with BG33R and planted into solarized soil
remained at or below the nematicide treat-
ment threshold for South Carolina for up to

Fig. 19.8. Eight dead 3-year-old peach trees on ‘Nemaguard’ rootstock (foreground) and live trees on
all ‘Guardian®’ rootstock (background, same row) in the presence of Mesocriconema xenoplax and the
peach tree short life disease complex. (Photo by A.P. Nyczepir.)
 Nematodes
2 years (Nyczepir et al., 1998c). Furthermore,
it was observed that significant and stable
qualitative changes in the microbial commu-
nity induced by soil solarization were corre-
lated with this reduction in M. xenoplax
population. In fact, solarization plus BG33R
applications and solarization alone were
shown to provide effective long-term nema-
tode suppression. In a second orchard trial
(D.A. Kluepfel, California, 2004, personal com-
munication), BG33R was delivered to trees
through a microsprinkler irrigation system
and similar M. xenoplax control was achieved.
Nyczepir et al. (2004b) also demonstrated
that multiple applications of the entomo-
pathogenic nematodes, Steinernema riobrave
Cabanillas, Poinar, & Raulston and Heter-
orhabditis bacteriophora Poinar, were not effec-
tive in suppressing populations of M. xenoplax
under controlled conditions for peach in rep-
licated tests. Results with other Mesocriconema
spp. have been inconsistent. Smitley et al.
(1992) reported that applications of H. bacte-
riophora did not reduce M. rusticum popula-
tions on turf, whereas S. riobrave applications
reduced recovery of a Mesocriconema sp. on
turf in Georgia (Grewal et al., 1997). One
explanation as to why there was a lack of
M. xenoplax suppression may be the result of
parasitic behaviour. The entomopathogenic
nematodes (i.e. Steinernema glaseri) are known
to be attracted to root tips, whereas M. xeno-
plax does not have a partiality towards feed-
ing at any specific root region other than the
root cortex tissue (Thomas, 1959; Bird and
Bird, 1986).
19.4 Root-lesion Nematodes
Species
At least nine root-lesion nematode species
have been reported on peach throughout the
world, they include: Pratylenchus penetrans
(Cobb) Chitwood & Oteifa, Pratylenchus vul-
nus Allen & Jensen, Pratylenchus pratensis (de
Man) Filipjev, Pratylenchus brachyurus (Godfrey)
Goodey, Pratylenchus zeae Graham, Pratylenchus
convallariae Seinhorst, Pratylenchus neglectus
(Rensch) Filipjev & Schuurmans Stekhoven,
521
Pratylenchus thornei Sher & Allen and Praty-
lenchus sefaensis Fortuner from Turkey (Nyczepir
and Becker, 1998; Kepenekci, 2001).
Symptoms
These nematodes are migratory endoparasites
that cause extensive root damage resulting from
their intra- and intercellular movement while
feeding on cortical cells. Above-ground symp-
toms can resemble nutrient deficiency and
include reduced shoot growth and general tree
vigour, and a reduction in fruit size. In Georgia
USA, Fliegel (1969) reported P. vulnus being
associated with reduced peach tree vigour. In
Canada, P. penetrans was affiliated with peach
tree decline (Mountain and Patrick, 1959).
Below-ground symptoms of Pratylenchus
spp. damage include a reduction in feeder
root number, root darkening and necrotic
lesions (McKenry, 1989) (Fig. 19.9/Plate 225).
Pitcher et al. (1960) related root tissue necrosis
following nematode injury to phenol content
or the ability of damaged plant cells to syn-
thesize phenols.
Biology
Unlike the adult stage of root-knot nematode,
Pratylenchus spp. have the capability of both
entering and exiting the root. Histological
studies of the peach–almond hybrid, ‘G × N
No. 1’, revealed that all life stages of P. vulnus
were observed in ‘large pockets’ and cavities
of living root cortical parenchyma cells (Mar-
ull and Pinochet, 1991). Both P. penetrans
(Corbett, 1973) and P. vulnus (Corbett, 1974)
reproduce sexually. Generally, adult females lay
eggs singly within individual living root cells.
However, some eggs can be found in necrotic
tissue resulting from nematode feeding and
movement. The first moult occurs in the egg
with the second-stage juvenile emerging at
hatch. Developing juveniles moult three more
times between feeding intervals before becom-
ing adults. Generally, the complete life cycle
for P. penetrans is temperature-dependent and
varies between 30 and 86 days at 30°C or
20/24°C, respectively.
522 A.P. Nyczepir and D. Esmenjaud
Fig. 19.9. Influence of Pratylenchus vulnus on ‘G × N No. 15’ almond–peach hybrid root growth after
24 months (left, uninoculated; right, inoculated, Pi = 1000 nematodes/plant, where Pi is initial
nematode population density). (Courtesy of J. Pinochet, Agromillora Catalana SA, Barcelona, Spain.)
Both P. penetrans and P. vulnus are more
damaging to plants growing in sandy loam
(coarse) than in finer-textured soils (Corbett,
1973, 1974). In Georgia, USA, P. vulnus soil
populations were highest from August to
December (Fliegel, 1969). Corbett (1973)
reported that P. penetrans populations on most
crops were highest in late summer and early
autumn and lowest in late spring and early
summer.
Survival and dissemination
P. vulnus is typically found in warmer cli-
mates, whereas P. penetrans is usually associ-
ated with cooler climates and higher
elevations (McKenry, 1989). Nematode-in-
fested nursery stock and transport of infested
soil on machinery account for most of Praty-
lenchus spp. movement to uninfested areas.
Economic importance
P. penetrans in Canada (Mountain and Patrick,
1959) and P. vulnus in the USA (McKenry,
1989), Spain (Pinochet et al., 2000) and France
(Scotto La Massèse, 1975) are economically
important pests of peach.
In California, P. vulnus damage to peach
rootstocks is estimated to cause a reduction of
about 16% in marketable fruit size and yield
(McKenry, 1989). In field microplots, P. vulnus
was associated with reduced peach tree
growth of ‘Guardian®’, ‘Lovell’ and ‘Nema-
guard’ rootstocks (Nyczepir and Pinochet,
2001). Furthermore, tree growth suppression
in the same study was greatest in the P. vulnus
(GA-peach isolate) infested plots than in the
P. vulnus (ID-apple isolate) plots. In Spain,
damage to ‘Nemared’ peach and ‘Garnem’
and ‘Monegro’ peach–almond hybrids was
evidenced at the end of the second growing
season in P. vulnus-infested microplots. All
tested rootstocks were good hosts for P. vul-
nus, which reached a high population density
in the roots (Pinochet et al., 1996b). In another
study also conducted in Spain, ‘Cadaman’
peach rootstock growth was suppressed in
the presence of P. vulnus (Spain-plum isolate)
(Hernandez-Dorrego et al., 1999). Pathogenic
diversity among P. vulnus isolates attacking
Prunus rootstocks appears to be high (Pinochet
et al., 2000). P. penetrans is also associated with
the peach replant problem in Canada (see fol-
lowing section).
 Nematodes
Disease complexes
P. penetrans is associated with the peach
replant problem in Canada (Mountain and
Patrick, 1959). The invading nematode causes
peach root necrosis in the absence of bacteria
and fungi. It is believed that P. penetrans initi-
ates root degeneration, thereby providing
sites for secondary infection by other soil
microbes.
P. vulnus is associated with peach replant
problems in Italy (Ricciardi et al., 1975) and
the USA (Fliegel, 1969). In the south-eastern
USA, P. vulnus was the primary root-lesion
nematode species most often associated with
extensive feeder root damage/loss and reduced
tree vigour. After 6 years in a field microplot
study, ‘Nemaguard’ peach trees growing in
soil infested solely with P. vulnus lived while
trees growing in soil infested solely with M.
xenoplax died from PTSL (A.P. Nyczepir, 2004,
personal observation).
Host range
The host range of P. vulnus and P. penetrans
includes over 80 and 350 plant species, respec-
tively; most hosts of P. vulnus are woody
perennials (Corbett, 1973, 1974; Pinochet et al.,
1992). Peach rootstocks (i.e. ‘Cadaman’,
‘Flordaguard’, ‘Guardian®’, ‘Lovell’, ‘Mont-
clar’, ‘Nemaguard’ and ‘Rutgers Red Leaf’)
tested under controlled conditions were all
hosts to P. vulnus and/or P. penetrans (Table
19.1) (Mountain and Patrick, 1959; Pinochet
et al., 2000; Nyczepir and Pinochet, 2001).
Control
Nematicides
Pre-plant soil fumigation and post-plant
application of recommended nematicides
provided adequate control of Pratylenchus
spp. in peach (Nyczepir, 1991). In Canada,
tree growth was improved and P. penetrans
populations suppressed for up to 4 years fol-
lowing soil fumigation with Vorlex (Olthof
et al., 1989). In Georgia, USA, the nematicide
treatment threshold is ≥1 P. vulnus/100 cm3
523
soil (Davis et al., 1996). Because state/country
recommendations for nematicide use are not
the same and change frequently, current local
sources of information need to be consulted
for updated recommendations.
Cultural practices
McFadden-Smith et al. (1993) noted that by-
products (i.e. allyl glucosinolate) of Brassica
green manure were toxic to P. penetrans. How-
ever, under field conditions, no root-lesion
nematode suppression was observed using
this green manure.
Resistance
Little progress has been made in identifying
sources of resistance to P. vulnus in commer-
cial peach rootstocks (McKenry, 1989; Pinochet,
1997; Pinochet et al., 2000; Nyczepir and
Pinochet, 2001). Resistance to root-lesion
nematodes has been difficult to detect and it
is also difficult to transmit from wild Prunus
or existing germplasm (e.g. obsolete plum
rootstocks of American origin) into commer-
cial rootstocks. Sources of resistance have
been identified in two wild Prunus species
(i.e. Prunus fremonti and Prunus tomentosa)
(Scotto La Massèse, 1975) and recently in
‘Redglow’ plum. Unfortunately, these sources
are non-viable from a breeding standpoint
since they do not graft or cross with existing
commercial rootstocks (Pinochet et al., 2000).
Findings in the last decade also indicate dif-
ferences in pathogenicity among P. vulnus iso-
lates, which further complicates the evaluation
of plant material (Pinochet et al., 1994, 1996a,
2000). Despite all these constraints, a few
commercial rootstocks have been found to be
moderately resistant to some P. vulnus iso-
lates. Since only partial sources of resistance
have been found to individual nematode iso-
lates, pooling genes from different sources
appears to be the only long-term effective
breeding strategy for now (Pinochet, 1997).
Resistance mechanisms to P. vulnus are
unknown. Tolerance to P. penetrans was iden-
tified in ‘Rutgers Red Leaf’, ‘Tzim Pee Tao’,
‘Bailey’, ‘BY520-8’, ‘Higama’ and ‘Guardian®’
peach rootstocks (Table 19.1) (Potter et al., 1984;
McFadden-Smith et al., 1998).
524 A.P. Nyczepir and D. Esmenjaud
19.5 Dagger Nematodes
Dagger nematodes (Xiphinema spp.) are
ectoparasites of Prunus that also transmit
nepoviruses (i.e. nematode-transmitted poly-
hedral viruses) which are lethal to stone fruits
(see section on ‘Nepovirus diseases’ below).
Species
There are more than 280 Xiphinema spp., but
only seven have been associated with stone
fruit disease. Two species, Xiphinema diversi-
caudatum and Xiphinema vuittenezi, originated
from Europe and five species including
Xiphinema americanum, Xiphinema brevicolle,
Xiphinema californicum, Xiphinema pachtaicum
and Xiphinema rivesi are from the X. america-
num group (Table 19.3). The species of the lat-
ter group were considered as a single species
until 1979 and are found mainly in North
America. Since that date, X. americanum
(today referred to as X. americanum sensu lato)
has been redefined by Lamberti and Bleve-
Zacheo (1979) into several distinct species
(mainly X. americanum sensu stricto, X. califor-
nicum, Xiphinema bricolensis and X. rivesi).
Nevertheless, the current status of those spe-
cies is not clear. Separation of the predomi-
nant species from North America can be
confirmed by molecular analysis using IST-
RFLP (internal transcribed spacer–restriction
fragment length polymorphism) markers
Table 19.3. Dagger nematode species and their associated nepoviruses in peach rootstocks.
Nematode
Xiphinema diversicaudatum
Xiphinema vuittenezi
Xiphinema pachtaicum
Xiphinema americanum sensu stricto
Xiphinema californicum
Xiphinema rivesi
Xiphinema bricolensis
(Vrain, 1993; Vrain et al., 1992) and those spe-
cies will be discussed in more detail within
this chapter. More recently, a complete listing
of the putative species from the X. americanum
group, their geographical occurrence and dis-
tribution was published (Lamberti et al., 2000,
2002). The authors have reclassified all spe-
cies into three categories: (i) the widespread
species (i.e. those previously mentioned);
(ii) the localized species; and (iii) the rare spe-
cies. For example, one of the rare species,
Xiphinema pacificum, was recently reported for
the first time in peach in Georgia, USA (Nyczepir
and Lamberti, 2001).
X. diversicaudatum is widely dispersed in
the temperate climates of Europe (Dalmasso,
1969, 1970; Scotto La Massèse, 1985) and is
occasionally found in North America, where
it was introduced on woody plants (Robbins
and Brown, 1991). X. vuittenezi is commonly
found in soils from stone fruit orchards in
Central and Eastern Europe (Taylor and
Brown, 1997), whereas X. pachtaicum, a Euro-
pean species of X. americanum sensu lato, is often
found on Prunus in France (C. Scotto La Massèse,
France, 1990, personal communication).
X. americanum sensu stricto is common in
North and South America but has also been
reported in South Africa (Lamberti and
Golden, 1984; Loots and Heyns, 1984; Luc
and Doucet, 1990; Robbins and Brown, 1991),
whereas X. rivesi is present in north-eastern
USA, eastern Canada, the former USSR, France,
Germany and Spain (Romanenko and Ste-
garescu, 1985; Luc and Doucet, 1990; Robbins
Nepovirus
Strawberry latent ringspot virus (SLRV)
Arabis mosaic virus (ArMV)
–
–
Peach rosette mosaic virus (PRMV)
Cherry rasp leaf virus (CRLV)
Tomato ringspot virus (ToRSV)
Tobacco ringspot virus (TRSV)
ToRSV
ToRSV
ToRSV
 Nematodes
and Brown, 1991). X. rivesi is suspected to
have been introduced to Europe (Taylor and
Brown, 1997).
X. californicum has frequently been
detected along the western seaboard of the
Americas, but it has also been reported in the
eastern USA (Lamberti et al., 1988; Robbins
and Brown, 1991). X. bricolensis is present in
British Columbia, Canada and also in Wash-
ington State and California, USA, where it is
more frequent in vineyards than in orchards
(Brown et al., 1994b).
Symptoms
Some species of Xiphinema induce gall forma-
tion at feeder root tips, which may appear
swollen and curled.
Biology
Little is known about the biology and life
cycle of most Xiphinema spp. X. diversicauda-
tum reproduces by amphimixis (i.e. true sex-
ual reproduction with fusion of sperm and
egg nuclei) and males are abundant. X. diver-
sicaudatum has a long life cycle (up to 3 years)
and showed no comparable annual cycle
(Flegg, 1968; Jaffee et al., 1987a). The life cycle
may take from several months to several
years depending on the environmental condi-
tions (temperature, soil moisture, soil type)
(D. Esmenjaud, unpublished results). Other
Xiphinema spp. reproduce by parthenogenesis
and the male stage is uncommon. Halbrendt
and Brown (1992) have shown that X. america-
num sensu stricto, X. californicum and X. rivesi
have only three juvenile stages and not four
as usually encountered with European spe-
cies such as X. diversicaudatum.
Nepovirus diseases
Most nepoviruses associated with Prunus
have been observed and described in North
America. A list of the nepoviruses and their
corresponding vectors is reported in Table 19.3
525
and detailed information can be found else-
where (Nyczepir and Halbrendt, 1993). The
most economically important nepovirus is
the Tomato ringspot virus (ToRSV) affecting
stone fruit in North and South America
(Brown et al., 1994a). The established vectors
of ToRSV are nematodes from the X. america-
num sensu lato group, including X. americanum
sensu stricto, X. bricolensis, X. californicum and
X. rivesi. This virus causes Prunus stem pit-
ting in peach (Mircetich and Fogle, 1976).
Other peach virus diseases include tobacco
ringspot caused by Tobacco ringspot virus
(TRSV) and peach rosette mosaic caused by
Peach rosette mosaic virus (PRMV), both vec-
tored by X. americanum sensu stricto (Klos,
1976), and strawberry latent ringspot caused
by Strawberry latent ringspot virus (SLRV), vec-
tored by X. diversicaudatum (Crossa-Reynaud
and Audergon, 1987). Strawberry latent ring-
spot is the most economically important nep-
ovirus disease in Europe.
Survival, dissemination and host range
All stone fruits and many other woody plant
species are hosts of Xiphinema spp. Nepovi-
ruses that are vectored by Xiphinema spp. are
naturally found in common broadleaved
weeds, which serve as natural reservoirs of
latent infections. Nematodes acquire the virus
by feeding on infected plants and subse-
quently transmit it when feeding on the peach
tree (Taylor and Robertson, 1970). Soil mois-
ture, temperature and texture affect nematode
survival and virus-vectoring efficiency
(Nyczepir and Becker, 1998).
Economic importance
The total economic impact of Xiphinema spp.
and nepoviruses to the European and US
peach industries has been difficult to estimate
(Nyczepir and Halbrendt, 1993). This is
because many factors impact orchard life and
productivity and it is difficult to attribute loss
to a single factor. Crop losses of 10% due to X.
americanum and P. penetrans damage have
been estimated by Klonsky and Bird (1981)
526 A.P. Nyczepir and D. Esmenjaud
for the Michigan tree fruit industry. Nema-
todes or weeds putatively harbouring ToRSV
as a primary causal agent for peach tree death
in Pennsylvania have been found in 90% of
the locations sampled (Greene et al., 1988).
Methods of diagnosis
Characteristic symptoms such as stem pit-
ting, constriction, brown line, rasp leaf and
rosette symptoms are the primary indicators
of a virus disease. Identification of ToRSV can
be confirmed by ELISA or dot-blot hybridiza-
tion (Powell et al., 1991). Subsequently, detec-
tion of the vector nematode can be attempted,
but sampling for Xiphinema vectors is difficult
on the established crop. The occurrence of X.
diversicaudatum and X. vuittenezi can be con-
firmed using multiplexed PCR primers
designed from ribosomal genes, which permit
a direct identification from a single nematode
(Wang et al., 2002), but molecular tools for the
identification of species from the X. america-
num group are lacking.
Control
Since X. diversicaudatum and SLRV share
many common hosts in the Rosaceae family,
the risk of virus transmission by the nema-
tode between different plant species is high.
Therefore, fields where rosaceous species
have been previously cultivated should be
avoided. In situations where soil transmis-
sion by dagger nematodes is clearly estab-
lished, pre-plant fumigation is an effective
means of managing virus diseases (Bernhard
et al., 1985). Applications of post-plant nem-
aticides and broadleaf herbicides to control
spread of nepoviruses are highly recom-
mended. Because state/country recommen-
dations for nematicide and herbicide use are
not the same and change frequently, current
local sources of information need to be con-
sulted for updated recommendations. Root-
stock resistance to ToRSV exists in Prunus
(Hoy and Mircetich, 1984), but is of limited
commercial interest for peach because of graft
union incompatibility problems.
19.6 Other Nematodes
Paratylenchus prunii Sharma, Sharma & Khan
suppressed peach growth under controlled
conditions and its reproduction rate was
inversely proportional to the initial popula-
tion density (Sharma and Sharma, 1987, 1988;
Nyczepir and Becker, 1998). A single species
of Longidorus, Longidorus diadecturus, appears
to be involved with transmitting PRMV, is
widespread in the central USA and extends
northwards to Ontario, Canada (Allen et al.,
1982; Halbrendt, 1993). Molecular identifica-
tion of certain Longidorus spp. found in Ger-
many and Western Europe has been developed
(Hubschen et al., 2004). Lists of nematodes
found in association with, but not necessar-
ily causing economic damage to, peach are
recorded elsewhere (Wehunt and Nyczepir,
1988).
19.7 Outlook
Utilization of nematicides for nematode
control in commercial nursery and orchard
systems is still heavily relied upon. In the
south-eastern USA alone, present manage-
ment practices include: (i) pre-plant fumiga-
tion (i.e. Telone II), which gets the trees off to a
good start and gives control for up to 2 years,
depending on the quality of the fumigation
and the nematode involved; and (ii) resistant
rootstocks (when available). However, with
the continued scrutiny and removal of the var-
ious nematicides by the regulatory agencies,
fewer and fewer chemical management options
are available to the grower. As a result, current
research efforts have shifted towards various
forms of alternative (non-chemical) nematode
control. Emphasis on non-chemical control is
partly due to the apprehension about the envi-
ronmental problems associated with soil fumi-
gation with methyl bromide and most recently
the voluntary removal in registration of the
post-plant nematicide, fenamiphos. As a result
of methyl bromide’s reputed role in ozone
depletion, the importation and manufacture of
methyl bromide in the USA have been banned
since January 2005 (Clean Air Act, 1990), with
certain exceptions (i.e. Critical Use Exemption,
 Nematodes 527

quarantine and pre-shipment exemptions, and
emergency exemptions). Therefore, finding an
alternative to chemical control of nematodes is
warranted.
Alternative non-chemical research areas
under investigation include the search for
biological control agent(s), biofumigation,
rotation/cover crops, rootstock resistance
and soil solarization, which could be used in
an integrated nematode management system.
We think that the key phrase here is integrated
pest management system (i.e. IPM). The term
IPM is not new, but it appears to be evident
that a combined nematode management sys-
tem approach for nematode control is what
needs to be developed and/or improved upon
for growers in the near future. Development of
such nematode management systems through
fundamental and applied research will pro-
vide the basis for nematode control that is
more effective, economical, and less hazard-
ous to man and the environment.

References

Alcaniz, A., Pinochet, J., Fernández, C., Esmenjaud, D. and Felipe, A. (1996) Evaluation of Prunus rootstocks
for root-lesion nematode resistance. HortScience 31, 1013–1016.
Allen, W.R., Van Schagen, J.G. and Everleigh, E.S. (1982) Transmission of peach rosette mosaic virus to peach,
grape, and cucumber by Longidorus diadecturus obtained from diseased orchards in Ontario. Canadian
Journal of Plant Pathology 4, 16–18.
Anon. (2003) Propanil and fenamiphos; use deletion and product cancellation order. Federal Register 68(237),
68901–68904.
Batchelor, T.A. (2002) International and European community controls on methyl bromide and the status of
methyl bromide use and alternatives in the European Community. In: Batchelor, T.A. and Bolivar, J.M.
(eds) Proceedings of the International Conference on Alternatives to Methyl Bromide ‘The Remaining
Challenges’. European Commission, Brussels, pp. 28–32.
Baxter, R.I. and Blake, C.D. (1969) Oxygen and the hatch of eggs and migration of larvae of Meloidogyne
javanica. Annals of Applied Biology 63, 191–203.
Beckman, T.G., Okie, W.R. and Nyczepir, A.P. (1993) Use of clonally replicated seedlings in field screening
for resistance to peach tree short life. Journal of the American Society for Horticultural Science 118,
115–118.
Bernhard, R., Bouquet, A. and Scotto La Massèse, C. (1985) Diversité des problems nématologiques en verg-
ers, solutions chimiques et génétiques. Création de variétés résistantes aux nématodes des cultures:
intérêt, possibilités et limites. Compte-Rendus de l’Académie d’Agriculture de France 71, 705–718.
Bertrand, P.F. (1985) Peach Nematodes and Their Control in Georgia. Cooperative Extension Service, Univer-
sity of Georgia, Athens, Georgia L347.
Bird, A.F. and Bird, J. (1986) Observations on the use of insect parasitic nematodes as a means of biological
control of root-knot nematodes. International Journal of Parasitology 16, 511–516.
Blenda, A.V., Reighard, G.L., Baird, W.V., Wang, Y. and Abbott, A.G. (2002) Application of molecular markers
in the development of peach rootstocks tolerant to ring nematode (Mesocriconema xenoplax). Acta Hor-
ticulturae 592, 229–231.
Blenda, A.V., Wechter, W.P., Reighard, G.L., Baird, W.V. and Abbott, A.G. (2006) Development and character-
ization of diagnostic AFLP markers in Prunus persica for its response to peach tree short life syndrome.
Journal of Horticultural Science & Biotechnology 81, 281–288.
Brittain, J.A. and Miller, R.W. (1978) Managing Peach Tree Short Life in the Southeast. Bulletin 585. Clemson
University Extension Service, Clemson, South Carolina, p. 19.
Brown, D.J.F., Halbrendt, J.M., Jones, A.T., Vrain, T.C. and Robbins, R.T. (1994a) Transmission of three North
American nepoviruses by populations of four distinct Xiphinema americanum-group species (Nematoda,
Dorylaimida). Phytopathology 84, 646–649.
Brown, D.J.F., Vrain, T.C., Jones, A.T., Robertson, W.M., Halbrendt, J.M. and Robbins, R.T. (1994b) Xiphinema
bricolensis – a natural vector of three serologically distinguishable strains of tomato ringspot virus. Journal
of Nematology 26, 94.
Calvet, C., Estaún, V., Camprubí, A. and Pinochet, J. (2000) Enfermedades de replantación en frutales. In:
Montesinos, E., Melgarejo, P., Cambra, M. and Pinochet, J. (eds) Enfermedades de Los Frutales de Pepita
y Hueso. Ediciones Mundi-Prensa, Madrid/Barcelona/México, pp. 107–109.
528 A.P. Nyczepir and D. Esmenjaud

Canals, J., Pinochet, J. and Felipe, A. (1992) Temperature and age of plants affect resistance in peach–almond
hybrids infected with Meloidogyne javanica. HortScience 27, 1211–1213.
Carneiro, R.M.D.G., Carvalho, F.L.C. and Kulczynski, S.M. (1998) Plant selection to control Mesocriconema
xenoplax and Meloidogyne spp. with crop rotation. Nematolgia-Brasileira 22, 41–48.
Castagnone-Sereno, P., Piotte, C., Uijthof, J., Abad, P., Wajnberg, E., Vanlerberghe-Masutti, F., Bongiovanni,
M. and Dalmasso, A. (1993) Genetic relationships between amphimictic and parthenogenetic nema-
todes of the genus Meloidogyne as inferred from repetitive DNA analysis. Heredity 70, 195–204.
Claverie, M., Bosselut, N., Lecouls, A.C., Voisin, R., Poizat, C., Dirlewanger, E., Kleinhentz, M., Lafargue, B.,
Laigret, F. and Esmenjaud, D. (2004) Location of independent root-knot nematode resistance genes in
the plum and peach species. Theoretical and Applied Genetics 108, 765–773.
Clean Air Act (1990) Title VI. Stratospheric Ozone Protection Publication L. Section 6001. US Congress,
Washington, DC, pp. 101–549.
Cook, R. and Evans, K. (1987) Resistance and tolerance. In: Brown, R.H. and Kerry, B.R. (eds) Principles and
Practice of Nematode Control in Crops. Academic Press, Sydney, Australia, pp. 179–231.
Corbett, D.C.M. (1973) Pratylenchus penetrans. CIH Descriptions of Plant Parasitic Nematodes, Set 2, No. 25.
Commonwealth Institute of Helminthology. William Cloves & Sons Ltd, St Albans, UK.
Corbett, D.C.M. (1974) Pratylenchus vulnus. CIH Descriptions of Plant Parasitic Nematodes, Set 3, No. 37.
Commonwealth Institute of Helminthology. William Cloves & Sons Ltd, St Albans, UK.
Crossa-Reynaud, P. and Audergon, J.M. (1987) Apricot rootstocks, In: Rom, R.C. and Carlson, R.F. (eds) Root-
stocks for Fruit Crops. Wiley, New York, pp. 321–360.
Dalmasso, A. (1969) Etude anatomique et taxonomique des genres Xiphinema, Longidorus et Paralongidorus (Nema-
toda: Longidoridae). Mémoires du Muséum National d’Histoire Naturelle. Série A. Zoologie 61, 33–82.
Dalmasso, A. (1970) Influence directe de quelques facteurs écologiques sur l’activité biologique et la distribution
des espèces françaises de la famille des Longidoridae (Dorylaimida). Annales de Zoologie et d’Ecologie
Animale 2, 163–200.
Dalmasso, A. and Berge, J.B. (1975) Variabilité génétique chez les Meloidogyne et tout particulièrement chez
M. hapla. Cahiers ORSTOM. Série Biologie 10, 233–238.
Dalmasso, A. and Berge, J.B. (1978) Molecular polymorphism and genetic relationship in some Meloidogyne
spp.: application to the taxonomy of Meloidogyne. Journal of Nematology 10, 323–332.
Davis, R.F., Bertrand, P., Gay, J.D., Baird, R.E., Padgett, G.B., Brown, E.A., Hendrix, F.F. and Balsdon, J.A.
(1996) Guide for Interpreting Nematode Assay Results. Bulletin 834. University of Georgia Cooperative
Extension Service, Athens, Georgia, p. 16.
Day, L.H. and Tufts, W.P. (1939) Further notes on nematode-resistant rootstocks for deciduous fruit trees. Pro-
ceedings of the American Society for Horticultural Science 37, 327–329.
de Guiran, G. (1979) A necessary diapause in root-knot nematodes. Observations on its distribution and in-
heritance in Meloidogyne incognita. Revue de Nematologie 2, 223–231.
de Guiran, G. and Netscher, G. (1970) Les nématodes du genre Meloidogyne parasites des cultures tropicales.
Cahiers de l’ORSTOM, Serie Biologie 11, 151–185.
de Guiran, G. and Villemain, M.A. (1980) Spécificité de la diapause embryonnaire des œufs de Meloidogyne
(Nematoda). Revue de Nematologie 3, 115–121.
Dhanvantari, B.N., Johnson, P.W. and Dirks, V.A. (1975) The role of nematodes in crown gall infection of
peach in Southern Ontario. Plant Disease Reporter 59, 109–112.
Dirlewanger, E., Kleinhentz, M., Xiloyannis, C., Dichio, B., Claverie, M., Bosselut, N., Howad, W., Voisin, R.,
Gomez-Aparisi, J., Rubio-Cabetas, M.J., Poessel, J.L., Di Vito, M., Arús, P., Laigret, F. and Esmenjaud, D.
(2004) Breeding for a new generation of Prunus rootstocks based on marker-assisted selection: a Euro-
pean initiative. Eucarpia Symposium on Fruit Breeding and Genetics, Angers, France 1–5 September
2003. Acta Horticulturae 663, 829–833.
Eayre, C.G., Jaffee, B.A. and Zehr, E.I. (1987) Suppression of Criconemella xenoplax by the nematophagous
fungus Hirsutella rhossiliensis. Plant Disease 71, 832–834.
Eisenback, J.D., Hirschmann, H., Sasser, J.N. and Triantaphyllou, A.C. (1981) A Guide to the Four Most Com-
mon Species of Root-knot Nematodes (Meloidogyne species), with a Pictorial Guide. Department of
Plant Pathology and Genetics, North Carolina State University and US Agency for International Develop-
ment, Raleigh, North Carolina.
Esbenshade, P.R. and Triantaphyllou, A.C. (1985) Use of enzyme phenotypes for identification of Meloidogyne
species. Journal of Nematology 17, 6–20.
Esbenshade, P.R. and Triantaphyllou, A.C. (1990) Isozyme phenotypes for the identification of Meloidogyne
species. Journal of Nematology 22, 5–15.
 Nematodes 529

Esmenjaud, D., Minot, J.C., Voisin, R., Pinochet, J. and Salesses, G. (1994) Inter- and intraspecific resistance
variability in Myrobalan plum, peach and peach–almond rootstocks using 22 root-knot nematode popu-
lations. Journal of the American Society for Horticultural Science 119, 94–100.
Esmenjaud, D., Minot, J.C. and Voisin, R. (1996a) Effect of durable inoculum pressure and high temperature
on root galling, nematode numbers and survival of Myrobalan plum genotypes (Prunus cerasifera Ehr.)
highly resistant to Meloidogyne spp. Fundamental and Applied Nematology 19, 85–90.
Esmenjaud, D., Minot, J.C., Voisin, R., Bonnet, A. and Salesses, G. (1996b) Inheritance of resistance to the
root-knot nematode Meloidogyne arenaria in Myrobalan plum. Theoretical and Applied Genetics 92,
873–879.
Esmenjaud, D., Minot, J C., Voisin, R., Pinochet, J., Simard, M.H. and Salesses, G. (1997) Differential response
to root-knot nematodes in Prunus species and correlative genetic implications. Journal of Nematology
29, 372–380.
Fargette, M., Phillips, M.S., Block, V.C., Waugh, R. and Trudgill, D.L. (1996) An RFLP study of relationships
between species, populations, and resistance breaking lines of tropical Meloidogyne. Fundamental and
Applied Nematology 19, 193–200.
Fernández, C., Pinochet, J., Esmenjaud, D., Salesses, G. and Felipe, A. (1994a) Resistance among new Prunus
rootstocks and selections to root-knot nematodes in Spain and France. HortScience 29, 1064–1067.
Fernández, C., Pinochet, J. and Felipe, A. (1994b) Influence of temperature on the expression of resistance in
six Prunus rootstocks infected with Meloidogyne incognita. Nematropica 23, 195–202.
Fernández, C., Pinochet, J., Esmenjaud, D., Gravato-Nobre, M.J. and Felipe, A. (1995) Age of plant material
influences resistance of some Prunus rootstocks to Meloidogyne incognita. HortScience 30, 582–585.
Flegg, J.J.M. (1968) Life-cycle studies of some Xiphinema and Longidorus species in southeastern England.
Nematologica 14, 197–210.
Fliegel, P. (1969) Population dynamics and pathogenicity of three species of Pratylenchus on peach. Phytopa-
thology 59, 120–124.
Foster, H.H., Gambrell, C.E. Jr, Rhodes, W.H. and Byrd, W.P. (1972) Effects of preplant nematicides and resis-
tant rootstocks on growth and fruit production of peach trees in Meloidogyne spp. infested soil of South
Carolina. Plant Disease Reporter 56, 169–173.
Golden, A.M. (1976) Meloidogyne taxonomy – current status, some problems, and needs. In: Proceedings of
the Research Planning Conference on Root-Knot Nematodes Meloidogyne spp. North Carolina State
University Graphics, Raleigh, North Carolina, pp. 13–17.
Greene, G.M., Burns, G., Crassweller, R.M., Dickert, M.F., Forer, L.B., Hickey, K.D., Hull, L.A., Jaffee, B.A.,
Powell, C.A., Smith, C.B. and Travis, J.W. (1988) Causes of peach tree death in Pennsylvania. In: Childers,
N.F. and Sherman, W.B. (eds) The Peach. Horticultural Publications, Gainesville, Florida, pp. 719–730.
Grewal, P.S., Martin, W.R., Miller, R.W. and Lewis, E.E. (1997) Suppression of plant-parasitic nematode popu-
lations in turfgrass by application of entomopathogenic nematodes. Biocontrol Science and Technology
7, 393–399.
Halbrendt, J.M. (1993) Virus-vector Longidoridae and their associated viruses in the Americas. Russian Journal
of Nematology 1, 65–68.
Halbrendt, J.M. and Brown, D.J.F. (1992) Morphometric evidence for three juvenile stages in some species of
Xiphinema americanum sensu lato. Journal of Nematology 24, 305–309.
Handoo, Z.A., Nyczepir, A.P., Esmenjaud, D., van der Beek, J.G., Castagnone-Sereno, P., Carta, L.K., Skantar,
A.M. and Higgins, J.A. (2004) Morphological, molecular and differential-host characterization of Melo-
idogyne floridensis n. sp. (Nematoda: Meloidogynidae), a root-knot nematode parasitizing peach in
Florida. Journal of Nematology 36, 20–35.
Hernandez-Dorrego, A., Pinochet, J. and Calvet, C. (1999) Growth response of peach and plum rootstocks
infected with Pratylenchus vulnus in microplots. Supplement to the Journal of Nematology 31(4S),
656–661.
Hirschmann, H. (1971) Comparative morphology and anatomy. In: Zuckerman, B.M. and Mai, W.F. (eds) Plant
Parasitic Nematodes. Vol. 1. Academic Press, New York, pp. 11–63.
Hirschmann, H. (1986) Meloidogyne hispanica n. sp. (Nematoda: Meloidogynidae), the ‘Seville root-knot
nematode’. Journal of Nematology 18, 520–532.
Hoy, J.W. and Mircetich, S.M. (1984) Prune brown line disease: susceptibility of prune rootstocks and tomato
virus detection. Phytopathology 74, 272–276.
Hubschen, J., Kling, L., Ipach, U., Zinkernagel, V., Brown, D.J.F. and Nielson, R. (2004) Development and valida-
tion of species-specific primers that provide a molecular diagnostic for virus-vector longidorid nematodes
and related species in German viticulture. European Journal of Plant Pathology 110, 883–891.
530 A.P. Nyczepir and D. Esmenjaud

Hugo, H.J. and Meyer, A.J. (1995) Severe nematode damage to peach trees in South Africa. Nematologica
41, 310.
Hung, C.L.P. and Jenkins, W.R. (1969) Criconemoides curvatum and the peach tree decline problem. Journal
of Nematology 1, 12.
Hussey, R.S., Mims, C.W. and Westcott, S.W. III. (1992) Ultrastructure of root cortical cells parasitized by the
ring nematode, Criconemella xenoplax. Protoplasma 167, 55–65.
Jaffee, B.A., Harrison, M.B., Shaffer, R.L. and Strang, M.B. (1987a) Seasonal population fluctuation of Xi-
phinema americanum and X. rivesi in New York and Pennsylvania orchards. Journal of Nematology 19,
369–378.
Jaffee, B.A., Nyczepir, A.P. and Golden, A.M. (1987b) Criconemella spp. in Pennsylvania peach orchards with
morphological observations of C. curvata and C. ornata. Journal of Nematology 19, 420–423.
Janati, A., Bergé, J.B., Triantaphyllou, A.C. and Dalmasso, A. (1982) Nouvelles données sur l’utilisation des
isoestérases pour l’identification des Meloidogyne. Revue de Nématologie 5, 147–154.
Kepenekci, I. (2001) Plant parasitic nematodes of Tylenchida (Nematoda) associated with stone fruits (apricot
and peaches) in southern Turkey. Pakistan Journal of Nematology 19, 49–61.
Kester, D.E. and Asay, R.N. (1986) ‘Hansen 2168’ and ‘Hansen 536’: two new Prunus rootstock clones. Hort-
Science 21, 331–332.
Kester, D.E. and Grasselly, C. (1987) Almond rootstocks. In: Rom, R.C. and Carlson, R.F. (eds) Rootstocks for
Fruit Crops. Wiley, New York, pp. 265–293.
Klonsky, K. and Bird, G.W. (1981) Economic Assessment of the Michigan Tart Industry in Relation to the Soil
Fumigant EDB. Agricultural Economics Report 401. Michigan State University, East Lansing, Michigan,
pp. 16–26.
Klos, E.J. (1976) Rosette mosaic. In: Virus Diseases and Non-Infectious Disorders of Stone Fruits in North
America. USDA Agriculture Handbook No. 437. US Department of Agriculture–Agricultural Research
Service, Washington, DC, pp. 137–138.
Kluepfel, D.A., Nyczepir, A.P., Lawrence, J.E., Wechter, P.W. and Leverentz, B. (2002) Biological control of the
phytoparasitic nematode Mesocriconema xenoplax on peach trees. Journal of Nematology 34, 120–123.
Kochba, J. and Spiegel-Roy, P. (1976) ‘Alnem 1’, ‘Alnem 88’, ‘Alnem 201’ almonds: nematode-resistant root-
stock seed source. HortScience 11, 270.
Lamberti, F. (1979) Economic importance of Meloidogyne spp. in subtropical and Mediterranean climates. In:
Lamberti, F. and Taylor, C.E. (eds) Root-Knot Nematodes (Meloidogyne Species): Systematics, Biology
and Control. Academic Press, New York, pp. 341–357.
Lamberti, F. and Bleve-Zacheo, T. (1979) Studies on Xiphinema americanum sensu lato with description of
fifteen new species (Nematoda, Longidoridae). Nematologia Mediterranea 7, 51–106.
Lamberti, F. and Golden, A.M. (1984) Redescription of Xiphinema americanum Cobb 1913 with comments
on its morphological variations. Journal of Nematology 16, 204–209.
Lamberti, F., Roca, F. and Agostinelli, A. (1988) On the identity of Xiphinema americanum in Chile with a key
to the Xiphinema species occurring in Chile. Nematologia Mediterranea 16, 67–68.
Lamberti, F., Molinari, S., Moens, M. and Brown, D.J.F. (2000) The Xiphinema americanum group. I. Putative
species, their geographical occurrence and distribution, and regional polytomous identification keys for
the group. Russian Journal of Nematology 8, 65–84.
Lamberti, F., Molinari, S., Moens, M. and Brown, D.J.F. (2002) The Xiphinema americanum group. II. Morpho-
metric relationships. Russian Journal of Nematology 10, 99–112.
Lawrence, E.G. and Zehr, E.I. (1978) Improvement of techniques for determining populations of Macropost-
honia xenoplax in dry soil. Phytopathology 68, 1102–1105.
Layne, R.E.C. (1987) Peach rootstocks. In: Rom, R.C. and Carlson, R.F. (eds) Rootstocks for Fruit Crops. Wiley,
New York, pp.185–216.
Lecouls, A.C., Salesses, G., Minot, J.C., Voisin, R., Bonnet, A. and Esmenjaud, D. (1997) Spectrum of the Ma
genes for resistance to Meloidogyne spp. in Myrobalan plum. Theoretical and Applied Genetics 95,
1325–1334.
Lecouls, A.C., Rubio-Cabetas, M.J., Minot, J.C., Voisin, R., Bonnet, A., Salesses, G., Dirlewanger, E. and Es-
menjaud, D. (1999) RAPD and SCAR markers linked to the Ma1 root-knot nematode resistance gene in
Myrobalan plum (Prunus cerasifera Ehr.). Theoretical and Applied Genetics 99, 328–335.
Lecouls, A.C., Bergougnoux, V., Rubio-Cabetas, M.J., Bosselut, N., Voisin, R., Poessel, J.L., Faurobert, M., Bon-
net, A., Salesses, G., Dirlewanger, E. and Esmenjaud, D. (2004) Marker-assisted selection for the wide-
spectrum resistance to root-knot nematodes conferred by the Ma gene from Myrobalan plum (Prunus
cerasifera) in interspecific Prunus material. Molecular Breeding 13, 113–124.
 Nematodes 531

Loots, G.C. and Heyns, J. (1984) A study of Xiphinema americanum sensu lato Heyns 1974 (Nematoda).
Phytophilactica 16, 313–319.
Lu, Z.X., Sossey-Alaoui, K., Reighard, G.L., Baird, W.V. and Abbott, A.G. (1999) Development and character-
ization of a codominant marker linked to root-knot nematode resistance, and its application to peach
rootstocks breeding. Theoretical and Applied Genetics 99, 115–123.
Lu, Z.X., Reighard, G.L., Nyczepir, A.P., Beckman, T.G. and Ramming, D.W. (2000) Inheritance of resistance
to root-knot nematodes in Prunus rootstocks. HortScience 35, 1344–1346.
Luc, M. and Doucet, M.E. (1990) La familia Longidoridae Thorne, 1935 (Nemata) en Argentina. 1. Distribu-
cion. Revista de Ciencias Agropecuarias 7, 19–25.
McFadden-Smith, W., Potter, J.W., Hawke, M. and Lazarovits, G. (1993) Use of Brassica green manures for
control of Verticillium dahliae and Pratylenchus penetrans. In: Proceedings of the Sixth International
Congress of Plant Pathology. National Research Council of Canada, Ottawa, p. 282.
McFadden-Smith, W., Miles, N.W. and Potter, J.W. (1998) Greenhouse evaluation of Prunus rootstocks for
resistance or tolerance to the root lesion nematode (Pratylenchus penetrans). Acta Horticulturae 465,
723–729.
McKenry, M.V. (1985) Nematodes. In: Flint, M.L. (ed.) Integrated Pest Management for Almonds. Publication
No. 3308. University of California Division of Natural Resources, Oakland, California, pp. 127–133.
McKenry, M.V. (1989) Nematodes. In: LaRue, J.H. and Johnson, R.S. (eds) Peaches, Plums, and Nectarines:
Growing and Handling for Fresh Market. Publication No. 3331. University of California Division of
Agriculture and Natural Resources, Oakland, California, pp. 139–147.
McKenry, M.V. (1990) Cover crops and nematode species. In: Proceedings of the International Conference on Ag-
riculture for the 21st Century. Pacific Culture Center & MOA Foundation, Honolulu, Hawaii, pp. 67–70.
McKenry, M.V. (1999) The Replant Problem and its Management. Catalina Publishing, Fresno, California.
Malo, S.E. (1967) Nature of resistance of ‘Okinawa’ and ‘Nemaguard’ peach to the root-knot nematode Melo-
idogyne javanica. Proceedings of the American Society for Horticultural Science 90, 39–46.
Marull, J. and Pinochet, J. (1991) Host suitability of Prunus rootstocks to four Meloidogyne species and Praty-
lenchus vulnus in Spain. Nematropica 21, 185–195.
Marull, J., Pinochet, J., Felipe, A. and Cenis, J.L. (1994) Resistance verification in Prunus selections to a mix-
ture of 13 Meloidogyne isolates and resistance mechanisms of a peach-almond hybrid to M. javanica.
Fundamental and Applied Nematology 17, 85–92.
Meyer, J.R., Zehr, E.I., Meager, R.L. Jr and Salvo, S.K. (1992) Survival and growth of peach trees and pest
populations in orchard plots managed with experimental ground covers. Agriculture, Ecosystems, and
Environment 41, 353–363.
Miller, R.W. (1994) Estimated peach tree losses 1980 to 1992 in South Carolina – causes and economic
impact. In: Nyczepir, A.P., Bertrand, P.F. and Beckman, T.G. (eds) Proceedings of the Sixth Stone Fruit
Decline Workshop, Fort Valley, Georgia. USDA-ARS, ARS-122. National Technical Information Service,
Springfield, Virginia, pp. 121–127.
Mircetich, S.M. and Fogle, H.W. (1976) Peach stem pitting. In: Virus Diseases and Non-Infectious Disorders of
Stone Fruits in North America. USDA Agriculture Handbook No. 437. US Department of Agriculture–
Agricultural Research Service, Washington, DC, pp. 77–87.
Mountain, W.B. and Patrick, Z.A. (1959) The peach replant problem in Ontario. VII. The pathogenicity of
Pratylenchus penetrans (Cobb, 1917) Filip. & Stek. 1941. Canadian Journal of Botany 37, 459–470.
Nyczepir, A.P. (1990) Influence of Criconemella xenoplax and pruning time on short life of peach trees. Journal
of Nematology 22, 97–100.
Nyczepir, A.P. (1991) Nematode management strategies in stone fruits in the United States. Journal of Nema-
tology 23, 334–341.
Nyczepir, A.P. (2001) Criconematids. In: Maloy, O.C. and Murray, T.D. (eds) Encyclopedia of Plant Pathology.
Wiley, New York, pp. 258–260.
Nyczepir, A.P. (2003a) Field evaluation of nonchemical alternatives for control of ring nematode on peach. In:
Proceedings of the Annual International Research Conference on Methyl Bromide Alternatives and Emis-
sions Reductions. Methyl Bromide Alternatives Outreach Press, Fresno, California, pp. 118/1–118/2.
Nyczepir, A.P. (2003b) Preplant crop rotation for nematode management – a replacement for ‘Stacy’ wheat.
In: Brannen, P.M. (ed.) Southeastern Regional Peach Newsletter No. 3. University of Georgia Cooperative
Extension Service, Athens, Georgia, 10–11.
Nyczepir, A.P. and Becker, O.J. (1998) Fruit and citrus trees. In: Barker, K.R., Pederson, G.A. and Windham,
G.L. (eds) Plant Nematode Interactions. American Society of Agronomy Monograph Series No. 36.
American Society of Agronomy, Madison, Wisconsin, pp. 637–684.
532 A.P. Nyczepir and D. Esmenjaud

Nyczepir, A.P. and Beckman, T.G. (2000) Host status of Guardian™ peach rootstock to Meloidogyne sp. and
M. javanica. HortScience 35, 772.
Nyczepir, A.P. and Bertrand, P.F. (2000) Preplanting bahia grass or wheat compared for controlling Mesocri-
conema xenoplax and short life in a young peach orchard. Plant Disease 84, 789–793.
Nyczepir, A.P. and Halbrendt, J.M. (1993) Nematode pests of deciduous fruit and nut trees. In: Evans, K.,
Trudgill, D.L. and Webster, J.M. (eds) Plant Parasitic Nematodes in Temperate Agriculture. CAB Interna-
tional, Wallingford, UK, pp. 381–425.
Nyczepir, A.P. and Lamberti, F. (2001) First record of Xiphinema pacificum from a peach orchard in Georgia.
Plant Disease 85, 1119.
Nyczepir, A.P. and Okie, W.R. (1996) Occurrence of peach tree short life on a field site with no history of
peach production. HortScience 31, 163.
Nyczepir, A.P. and Pinochet, J. (2001) Assessment of Guardian peach rootstock for resistance to two isolates
of Pratylenchus vulnus. Supplement to the Journal of Nematology 33(4S), 302–305.
Nyczepir, A.P. and Rodriguez-Kabana, R. (2007) Preplant biofumigation with sorghum or methyl bromide com-
pared for managing Criconemoides xenoplax in a young peach orchard. Plant Disease 91, 1607–1611.
Nyczepir, A.P., Zehr, E.I., Lewis, S.A. and Harshman, D.C. (1983) Short life of peach trees induced by Cricone-
mella xenoplax. Plant Disease 67, 507–508.
Nyczepir, A.P., Reilly, C.C. and Okie, W.R. (1987) Effect of initial population density of Criconemella xeno-
plax on reducing sugars, free amino acids, and survival of peach seedlings over time. Journal of Nema-
tology 19, 296–303.
Nyczepir, A.P., Wood, B.W. and Reighard, G.L. (1997) Impact of Meloidogyne incognita on the incidence of
peach tree short life in the presence of Criconemella xenoplax. Supplement to the Journal of Nematol-
ogy 29, 725–730.
Nyczepir, A.P., Bertrand, P.F., Parker, M.L., Meyer, J.R. and Zehr, E.I. (1998a) Interplanting wheat is not an ef-
fective postplant management tactic for Criconemella xenoplax in peach production. Plant Disease 82,
573–577.
Nyczepir, A.P., Esmenjaud, D. and Eisenback, J.D. (1998b) Pathogenicity of Meloidogyne sp. (FL-isolate) on
Prunus in the southeastern United States and France. Journal of Nematology 30, 509.
Nyczepir, A.P., Kluepfel, D.A., Lawrence, J. and Zehr, E.I. (1998c) Effect of preplant solarization on ring
nematode in a peach tree short life site. In: Proceedings of the Annual International Research Conference
on Methyl Bromide Alternatives and Emissions Reductions. Methyl Bromide Alternatives Outreach Press,
Fresno, California, pp. 4/1–4/2.
Nyczepir, A.P., Beckman, T.G. and Reighard, G.L. (1999) Reproduction and development of Meloidogyne
incognita and M. javanica on Guardian peach rootstock. Journal of Nematology 31, 334–340.
Nyczepir, A.P., Okie, W.R. and Beckman, T.G. (2004a) Creating a short life site for Prunus rootstock evaluation
on land with no innate Mesocriconema xenoplax population. HortScience 39, 124–126.
Nyczepir, A.P., Shapiro-Ilan, D.A., Lewis, E.E. and Handoo, Z.A. (2004b) Effect of entomopathogenic nematodes
on Mesocriconema xenoplax populations in peach and pecan. Journal of Nematology 36, 181–185.
Nyczepir, A.P., Beckman, T.G. and Reighard, G.L. (2006) Field evaluation of ‘Guardian®’ peach rootstock to
different root-knot nematode species. Acta Horticulturae 713, 303–309.
Okie, W.R., Beckman, T.G., Nyczepir, A.P., Reighard, G.L., Newell, W.C. Jr and Zehr, E.I. (1994a) BY520-9, a peach
rootstock for the southeastern United States that increases scion longevity. HortScience 29, 705–706.
Okie, W.R., Reighard, G.L., Beckman, T.G., Nyczepir, A.P., Reilly, C.C., Zehr, E.I., Newell, W.C. Jr and Cain,
D.W. (1994b) Field screening Prunus for longevity in the southeastern United States. HortScience 29,
673–677.
Olien, W.C., Ridley, J.D., Whitwell, T., Watson, W.A. and Newall, W.C. (1994) Methods for establishing nim-
blewill from seed as a ground cover for peach orchards. In: Nyczepir, A.P., Bertrand, P.F. and Beckman,
T.G. (eds) Proceedings of the Sixth Stone Fruit Decline Workshop, Fort Valley, Georgia. USDA-ARS, ARS-
122. National Technical Information Service, Springfield, Virginia, pp. 19–23.
Olthof, T.H.A., Johnson, P.W. and Potter, J.W. (1989) Establishment of a Redhaven peach orchard with Praty-
lenchus penetrans-infested and noninfested rootstocks in a fumigated and nonfumigated Niagara soil.
Canadian Journal of Plant Science 69, 285–295.
Orion, D. and Zutra, D. (1971) The effect of the root knot nematode on the penetration of crown gall bacteria
in almond roots. Israeli Journal of Agricultural Research 2, 27–29.
Petersen, D.J., Zijlstra, C., Wishert, J., Blok, V.C. and Vrain, T.C. (1997) Specific probes efficiently distinguish
root-knot nematodes species using signature sequences in ribosomal intergenic spacers. Fundamental
and Applied Nematology 20, 619–626.
 Nematodes 533

Pinochet, J. (1997) Breeding and selection for resistance to root-knot and lesion nematodes in Prunus root-
stocks adapted to Mediterranean conditions. Phytoparasitica 25, 271–274.
Pinochet, J., Verdejo, S. and Marull, J. (1989) Evaluacion de siete patrones de Prunus a tres especies de
Meloidogyne en Espana. Nematropica 19, 125–134.
Pinochet, J., Marull, J. and Felipe, A. (1992) Response of newly introduced peach, plum, and cherry rootstocks
to Meloidogyne javanica in Spain. Nematropica 22, 99–102.
Pinochet, J., Cenis, J.L., Fernández, C., Doucet, M. and Marull, J. (1994) Reproductive fitness and random
amplified polymorphic DNA variation among isolates of Pratylenchus vulnus. Journal of Nematology
26, 271–277.
Pinochet, J., Anglés, M., Dalmau, E., Fernández, C. and Felipe, A. (1996a) Prunus rootstock evaluation to root-
knot and lesion nematodes in Spain. Journal of Nematology 28(4S), 616–623.
Pinochet, J., Fernández, C., Alcañiz, E. and Felipe, A. (1996b) Damage by a lesion nematode, Pratylenchus
vulnus, to Prunus rootstocks. Plant Disease 80, 754–757.
Pinochet, J., Calvet, C., Hernández Dorrego, A., Bonet, A., Felipe, A. and Moreno, M. (1999) Resistance of
peach and plum rootstocks from Spain, France, and Italy to root-knot nematode Meloidogyne javanica.
HortScience 34, 1259–1262.
Pinochet, J., Fernández, C., Calvet, C., Hernandez-Dorrego, A. and Felipe, A. (2000) Selection against Praty-
lenchus vulnus populations attacking Prunus rootstocks. HortScience 35, 1333–1337.
Piotte, C., Castagnone-Sereno, P., Uijthof, J., Abad, P., Bongiovanni, M. and Dalmasso, A. (1992) Molecular
characterization of species and populations of Meloidogyne from various geographic origins with
repeated-DNA homologous probes. Fundamental and Applied Nematology 15, 271–276.
Pitcher, R.S., Patrick, Z.A. and Mountain, W.B. (1960) Studies on the host–parasite relations of Pratylenchus
penetrans (Cobb) to apple seedlings. I. Pathogenicity under sterile conditions. Nematologica 5, 309–314.
Potter, J.W., Dirks, V.A., Johnson, P.W., Olthof, T.H.A., Layne, R.E.C. and McDonnell, M.M. (1984) Response
of peach seedlings to infection by the root lesion nematode Pratylenchus penetrans under controlled
conditions. Journal of Nematology 16, 317–322.
Powell, C.A., Hadidi, A. and Halbrendt, J.M. (1991) Detection of tomato ringspot virus in nectarine trees using
ELISA and transcribed RNA probes. HortScience 26, 1290–1292.
Powers, T.O. and Harris, T.S. (1993) A polymerase chain reaction method for identification of five major
Meloidogyne species. Journal of Nematology 25, 1–6.
Rammah, A. and Hirschmann, H. (1988) Meloidogyne mayaguensis n. sp. (Meloidogynidae), a root-knot
nematode from Puerto Rico. Journal of Nematology 20, 58–69.
Ramming, D.W. and Tanner, O. (1983) Nemared peach rootstock. HortScience 18, 376.
Randig, O., Leroy, F., Bongiovanni, M. and Castagnone-Sereno, P. (2001) RAPD characterization of single fe-
males of the root-knot nematodes, Meloidogyne spp. European Journal of Plant Pathology 107, 639–643.
Reighard, G.L. (2002) Current directions of peach rootstock programs worldwide. Acta Horticulturae 592,
421–427.
Ricciardi, P., Amici, A. and Lalatta, F. (1975) Ulteriori indagini sul ruolo di Pratylenchus vulnus nel problema
del reimpianto del pesco. Rivista dell’Ortoflorofrutticoltura Italiana 59, 109–123.
Ritchie, D.F. (1988) Population dynamics of ring nematodes and peach tree short life in North Carolina. In:
Proceedings of the Third Stone Fruit Decline Workshop. National Technical Information Service, Spring-
field, Virginia, pp. 34–37.
Ritchie, D.F. and Zehr, E.I. (1995) Peach tree short life. In: Ogawa, J.M., Zehr, E.I., Bird, G.W., Ritchie, D.F.,
Uriu, K. and Uyemoto, J.K. (eds) Compendium of Stone Fruit Diseases. APS Press, St. Paul, Minnesota,
pp. 45–46.
Robbins, R.T. and Brown, D.J.F. (1991) Comments on the taxonomy, occurrence and distribution of Longidori-
dae (Nematoda) in North America. Nematologica 37, 395–418.
Romanenko, N.D. and Stegarescu, O.P. (1985) Taxonomic problem in nematodes of the Xiphinema america-
num group (Nematoda, Dorylaimoidea). Parazitologia 19, 434–442.
Rubio-Cabetas, M.J., Lecouls, A.C., Salesses, G., Bonnet, A., Minot, J.C., Voisin, R. and Esmenjaud, D. (1998)
Evidence of a new gene for high resistance to Meloidogyne spp. in Myrobalan plum (Prunus cerasifera).
Plant Breeding 117, 567–571.
Rubio-Cabetas, M.J., Minot, J.C., Voisin, R., Esmenjaud, D., Salesses, G. and Bonnet, G. (1999) Response of
the Ma genes from Myrobalan plum to Meloidogyne hapla and M. mayaguensis. HortScience 34, 1266–
1268.
Rubio-Cabetas, M.J., Minot, J.C., Voisin, R. and Esmenjaud, D. (2001) Interaction of root-knot nematodes
(RKN) and Agrobacterium tumefaciens in roots of Prunus cerasifera: evidence of the protective effect of
534 A.P. Nyczepir and D. Esmenjaud

the Ma RKN resistance against expression of crown gall symptoms. European Journal of Plant Pathology
107, 433–441.
Salesses, G., Grasselly, C., Renaud, R. and Claverie, J. (1992) Les porte-greffe des espèces fruitières à noyau
du genre Prunus. In: Gallais, A. and Bannerot, H. (eds) Amélioration des espèces végétales cultivées:
objectifs et critères de sélection. INRA Editions, Paris, pp. 605–650.
Sasser, J.N. (1979) Pathogenicity, host range and variability in Meloidogyne species. In: Lamberti, F. and
Taylor, C.E. (eds) Root-Knot Nematodes (Meloidogyne Species): Systematics, Biology and Control.
Academic Press, New York, pp. 257–268.
Sasser, J.N. and Freckman, D.W. (1987) A world perspective in nematology: the role of the society. In:
Veech, J.A. and Dickson, D.W. (eds) Vistas on Nematology. Society of Nematologists, Inc., Hyattsville,
Maryland, pp. 7–14.
Scotto La Massèse, C. (1975) Tests d’hôte de quelques porte-greffe et variétés fruitières à l’égard de Pratylenchus
vulnus Allen and Jensen. Compte-Rendus de l’Académie d’Agriculture de France 61, 1088–1095.
Scotto La Massèse, C. (1985) Atlas of plant parasitic nematodes of France. In: Alphey, T.J.W. (ed.) Distribution
of Longidoridae, Xiphinemidae and Trichodoridae. European Plant Parasitic Nematode Survey, Scottish
Crop Research Institute, Invergowrie, UK and INRA Station de Nematologie et de Genetique Molécu-
laire des Invertébrés, Antibes, France, pp. 1–43.
Scotto La Massèse, C., Grasselly, C., Minot, J.C. and Voisin, R. (1984) Différence de comportement de 23
clones et hybrides de Prunus à l’égard de quatre espèces de Meloidogyne. Revue de Nématologie 7,
265–270.
Seshadri, A.R. (1964) Investigations on the biology and life cycle of Criconemoides xenoplax Raski, 1952
(Nematoda: Criconematidae). Nematologica 10, 540–562.
Sharma, G.C. and Sharma, N.K. (1987) Effect of initial inoculum levels and multiplication potential of Paraty-
lenchus prunii on peach. Indian Journal of Nematology 17, 211–213.
Sharma, G.C. and Sharma, N.K. (1988) Effect of Paratylenchus prunii and Meloidogyne incognita on peach
seedlings. Nematologia Mediterranea 16, 117.
Sharpe, R.H. and Perry, V.G. (1967) Root-knot nematode populations on peaches in Florida. Proceedings of
the Florida State Horticultural Society 80, 342–344.
Sharpe, R.H., Hesse, C.O., Lownsbery, B.A., Perry, V.G. and Hansen, C.J. (1969) Breeding peaches for root-
knot nematode resistance. Journal of the American Society for Horticultural Science 94, 209–212.
Sharpe, R.R., Reilly, C.C., Nyczepir, A.P. and Okie, W.R. (1989) Establishment of peach in a replant site as
affected by soil fumigation, rootstock, and pruning date. Plant Disease 73, 412–415.
Sharpe, R.R., Pusey, P.L., Nyczepir, A.P. and Florkowski, W.J. (1993) Yield and economics of intervention with
peach tree short life disease. Journal of Production Agriculture 6, 241–244.
Shepard, D.P., Zehr, E.I. and Bridges, W.C. (1999) Increased susceptibility to bacterial spot of peach trees
growing in soil infested with Criconemella xenoplax. Plant Disease 83, 961–963.
Sherman, W.B. and Lyrene, P.M. (1983) Improvement of peach rootstock resistant to root-knot nematodes.
Proceedings of the Florida State Horticultural Society 96, 207–208.
Sherman, W.B., Lyrene, P.M. and Hansche, P.E. (1981) Breeding peach rootstocks resistant to root-knot nema-
tode. HortScience 16, 523–524.
Sherman, W.B., Lyrene, P.M. and Sharpe, R.H. (1991) Flordaguard peach rootstock. HortScience 26, 427–428.
Smitley, D.R., Warner, F.W. and Bird, G.W. (1992) Influence of irrigation and Heterorhabditis bacteriophora
on plant-parasitic nematodes in turf. Supplement to Journal of Nematology 24, 637–641.
Stalin, N., Salesses, G., Pinochet, J., Minot, J.C., Voisin, R. and Esmenjaud, D. (1998) Comparative host suit-
ability to Meloidogyne spp. and Pratylenchus vulnus in Myrobalan plum genotypes (Prunus cerasifera
Ehr.). Plant Pathology 47, 211–215.
Stirling, G.R. (1975) A survey of the plant-parasitic nematodes in Riverland peach orchards. Agricultural Re-
cord 2, 11–13.
Taylor, C.E. and Brown, D.J.F. (1997) Geographical distribution of Longidoridae. Nematode Vectors of Plant
Viruses. CAB International, Wallingford, UK, pp. 66–95.
Taylor, C.E. and Robertson, W.M. (1970) Sites of virus retention in the alimentary tract of the nematode vec-
tors, Xiphinema diversicaudatum (Micol.) and X. index (Thorne and Allen). Annals of Applied Biology 66,
375–380.
Thomas, H.A. (1959) On Criconemoides xenoplax Raski, with special reference to its biology under labora-
tory conditions. Proceedings of the Helminthological Society of Washington 26, 55–59.
Triantaphyllou, A.C. (1971) Genetics and cytology. In: Zuckerman, B.M., Mai, W.F. and Rohde, R.A. (eds)
Plant Parasitic Nematodes, Vol. II. Academic Press, New York, pp. 1–34.
 Nematodes 535

Triantaphyllou, A.C. (1985) Cytogenetics, cytotaxonomy and phylogeny of root-knot nematodes. In: Sasser,
J.N. and Carter, C.C. (eds) An Advanced Treatise on Meloidogyne, Vol. I. North Carolina State University
Graphics, Raleigh, North Carolina, pp. 113–126.
Tufts, W.P. (1929) Nematode resistance of certain peach seedlings. Proceedings of the American Society for
Horticultural Science 26, 98–110.
Voisin, R., Minot, J.C. and Esmenjaud, D. (1999) Penetration, development and emigration of juveniles of the
nematode Meloidogyne arenaria in Myrobalan plum (Prunus cerasifera) clones bearing the Ma resis-
tance genes. European Journal of Plant Pathology 105, 103–108.
Vrain, T.C. (1993) Restriction fragment length polymorphism separates species of the Xiphinema americanum
group. Journal of Nematology 25, 361–364.
Vrain, T.C., Wakarchuk, D.A., Levesque, A.C. and Hamilton, R.I. (1992) Intraspecific rDNA restriction frag-
ment polymorphism in the Xiphinema americanum group. Fundamental and Applied Nematology 15,
563–573.
Wang, X., Bosselut, N., Castagnone, C., Voisin, R., Abad, P. and Esmenjaud, D. (2002) Multiplex PCR identi-
fication of single individuals of the Longidorid nematodes, Xiphinema index, X. diversicaudatum, X.
vuittenezi and X. italiae using specific primers from ribosomal genes. Phytopathology 93, 160–166.
Wehunt, E.J. and Nyczepir, A.P. (1988) Nematodes on peaches in the US – possible controls. In: Childers, N.F.
and Sherman, W.B. (eds) The Peach. Horticultural Publications, Gainesville, Florida, pp. 739–750.
Wehunt, E.J. and Weaver, D.J. (1972) Effect of nematodes and Fusarium oxysporum on the growth of peach
seedlings in the greenhouse. Journal of Nematology 4, 236.
Weinberger, J.H., Marth, P.C. and Scott, D.H. (1943) Inheritance study of root-knot nematode resistance in
certain peach varieties. Proceedings of the American Society for Horticultural Science 42, 321–325.
Westcott, S.W. III and Burrows, P.M. (1991) Degree-day models for predicting egg hatch and population in-
crease of Criconemella xenoplax. Journal of Nematology 23, 386–392.
Westcott, S.W. III and Hussey, R.S. (1992) Feeding behavior of Criconemella xenoplax in monoxenic cultures.
Phytopathology 82, 936–940.
Westcott, S.W. III and Zehr, E.I. (1991) Evaluation of host suitability in Prunus for Criconemella xenoplax.
Journal of Nematology 23, 393–401.
Yamamoto, T. and Hayashi, T. (2002) New root-knot nematode resistance genes and their STS markers in
peach. Scientia Horticulturae 96, 81–90.
Zehr, E.I., Lewis, S.A. and Bonner, M.J. (1986) Some herbaceous hosts of the ring nematode (Criconemella
xenoplax). Plant Disease 70, 1066–1069.
Zehr, E.I., Aitken, J.B., Scott, J.M. and Meyer, J.R. (1990) Additional hosts for the ring nematode, Criconemella
xenoplax. Journal of Nematology 22, 86–89.
Zijlstra, C. (1997) A fast PCR assay to identify Meloidogyne hapla, M. chitwoodi, and M. fallax and to sensi-
tively differentiate them from each other and from M. incognita mixtures. Fundamental and Applied
Nematology 20, 505–511.
Zijlstra, C., Donker-Venne, D.T.H.M. and Fargette, M. (2000) Identification of Meloidogyne incognita, M. javanica
and M. arenaria using sequence characterised amplified region based PCR assays. Nematology 2, 847–853.
 20 Preharvest Factors Affecting
Peach Quality

C.H. Crisosto1 and G. Costa2

1Universityof California, Davis, Department of Plant Sciences; located at Kearney
Agricultural Center, Parlier, California, USA
2Department of Fruit Trees and Woody Plant Sciences, University of Bologna,

Bologna, Italy

20.1 Introduction 536

20.2 Quality Definition 537
20.3 Maturity and Quality 537
20.4 Genotype 539
20.5 Mineral Nutrition 540
Nitrogen 540
Calcium 541
Potassium 542
Iron 542
20.6 Irrigation 542
20.7 Canopy Manipulation 544

20.1 Introduction on fruit flavour and postharvest life potential

This chapter describes and discusses exclu-
sively the impact of preharvest factors on
peach fruit flavour and postharvest life (stor-
age and shelf-life). It does not include the role
of environmental factors and the use of plant
growth regulators on peach quality; these
topics are covered well in other chapters in
this book. The present chapter begins by
emphasizing quality definitions from the con-
sumer point of view and follows with a series
of short sections that update knowledge on
preharvest orchard factors. The relationship
between maturity and quality is covered, and
the role of genotype (cultivar and rootstock)
is described. Then the effect of mineral nutri-
tion on peach quality is discussed with
detailed attention on N and Ca as the most
studied nutrients. Updated information on
the effect of foliar nutrient application on fruit
quality including foliar Ca sprays is also
reported. Detailed practical information on
the effect of different irrigation regimes on
fruit quality is described next, followed by a
section on canopy management. The canopy
management section describes practical infor-
mation on leaf removal, girdling techniques,
and the use of reflective materials to improve
fruit size and enhance red skin colour. Exam-
ples that illustrate the influence of crop load

536 © CAB International 2008. The Peach: Botany, Production and Uses
(eds D.R. Layne and D. Bassi)
 Preharvest Factors Affecting Quality
and canopy position on fruit quality and post-
harvest storage potential are presented.
20.2 Quality Definition
Fruit ‘quality’ is a concept encompassing sen-
sory properties (appearance, texture, taste
and aroma), nutritive value, mechanical prop-
erties, safety and defects. Altogether, these
attributes give the fruit a degree of excellence
and an economic value (Abbott, 1999). Every-
one in the peach production and marketing
chain from the grower to the consumer looks
for fruit with no or few defects. However, in
each step of this chain, the term ‘quality’ takes
on different meanings and the economic rele-
vance of the various quality traits is largely
variable. For example, the grower is inter-
ested in high yield, in fruit with large size and
high disease resistance, and in the opportu-
nity to reduce the number of pickings. The
definition of ‘quality’ for packers, shippers,
distributors and wholesalers is mainly based
on flesh firmness, which is considered a good
indication to predict fruit potential storage
and market life. Peaches and nectarines ripen
and deteriorate quickly at ambient tempera-
ture and cold storage is required to slow
down these processes, especially for some
cultivars and/or long-distance market situa-
tions. However, firmness is an erroneous and
incomplete way to estimate peach posthar-
vest potential for domestic distribution. In
fact, in some production areas such as Cali-
fornia and Chile, the development of internal
breakdown symptoms such as lack of flavour,
flesh mealiness and flesh browning limits the
storage life and the postharvest quality of
tasty cultivars. For retailers, red colour, size
and firmness have historically represented
the main components of fruit quality, as they
need fruit that are attractive to the consumer,
resistant to handling and have a long shelf-
life. From the consumer’s point of view, in
general peach fruit quality has declined,
mainly because of premature harvesting,
chilling injury and lack of ripening prior to
consumption, resulting in consumer dissatis-
faction. In addition, quality is badly defined
and the only parameters being considered are
fruit size and skin colour. Other characters
537
such as flesh firmness, sugar content, acidity
and aroma, which are perceived by the con-
sumer as fruit quality, are completely disre-
garded by the grower and other individuals
along the chain. In fact, the grower, identifying
fruit quality almost exclusively with the fruit
size, does not consider that these are only the
first characters perceived by the consumer and
they orient him just in his very first choice. As
soon as he realizes that the fruit, even with good
size and attractive colour, is tasteless, with low
sugar content, poor aroma and rapidly perish-
able, he redirects his interest towards other
types of fruit. As a consequence, it is imperative
for the grower and other individuals in the
delivery chain to direct their attention to fruit
quality from the consumer’s perspective in
order to regain the confidence of the consumer.
In addition, there is now an increasing
appreciation that ‘quality’ of fruit also includes
nutritional properties (e.g. vitamins, minerals,
dietary fibre) and health benefits (e.g. antioxi-
dants); and these are becoming important fac-
tors in consumer preferences. Experimental,
epidemiological and clinical studies provide
evidence that diet has an important role in the
prevention of the chronic degenerative diseases
such as tumours, cardiovascular diseases and
atherosclerosis. It is supposed, in fact, that the
consumption of fresh fruit and vegetables exerts
a protective role against the development of
such pathologies (Doll, 1990; Ames et al., 1993;
Dragsted et al., 1993; Anderson et al., 2000).
Changes in quality definition that are
focusing more on consumer demand can
increase peach consumption if marketing pro-
motion and education programmes are well
executed. Because the consumer quality of
peaches cannot be improved after harvest, it
is important to understand the role of prehar-
vest factors in consumer acceptance and mar-
ket life (Kader, 1988; Crisosto et al., 1997).
20.3 Maturity and Quality
Peaches and nectarines are climacteric fruit char-
acterized by a sharp rise in ethylene biosynthe-
sis at the onset of ripening, which is associated
with changes in sensitivity to the hormone
itself and changes in colour, texture, aroma and
other biochemical features (Fig. 20.1/Plate 226).
538 C.H. Crisosto and G. Costa
Ethylene plays a key role in peach fruit rip-
ening by coordinating the expression of
ripening-related genes responsible for flesh
softening, colour development and sugar
accumulation, as well as other processes such
as abscission (Ruperti et al., 2002; Trainotti
et al., 2003, 2006).
The definition of the proper harvest time
is essential, as fruit maturity at harvest greatly
influences peach fruit market life potential
and quality. Recently, the most important
peach-producing countries in Europe have
lost considerable market share mainly due to
excessive early harvesting. A delayed harvest
could lead to a better fruit organoleptic qual-
ity but also to faster softening and a shorter
shelf-life. In fact, different from other species,
in peach fruit there is a close link between
‘on-tree physiological maturity’ and evolu-
tion of key traits responsible for peach quality
during the postharvest phase. On the other
hand, melting flesh peaches and nectarines
undergo a rapid softening after harvest,
which leads to dramatic losses in the market-
ing chain, as soft fruit are easily bruised dur-
ing handling and more susceptible to decay.
Therefore, they are often picked at an early
Fig. 20.1. Peaches picked at different maturity levels.
stage of ripening, and they never reach their
full flavour and aroma potential.
Modulation of pre- and postharvest
peach fruit ripening by the means of chemi-
cals that interfere with ethylene biosynthesis
and/or perception, such as aminoethoxyvi-
nylglycine and 1-methylcyclopropene, has
already been reported (Mathooko et al., 2001;
Bregoli et al., 2002, 2005; Ziosi et al., 2006). A bet-
ter understanding of the physiological basis of
the peach fruit ripening process should make
it possible to develop further strategies to reg-
ulate ripening. Such strategies need objective
parameters able to accurately describe fruit
ripeness stages and internal quality changes
occurring in pre- and postharvest conditions.
Until recently, few studies have been car-
ried out on this topic, and mainly by using
traditional fruit quality traits (flesh firmness,
soluble solids concentration (SSC) and titrat-
able acidity (TA)) which are assessed with
simple devices such as penetrometers, refrac-
tometers and titrators. Early studies carried out
in Europe and the USA have associated peach
fruit consumer acceptance with high SSC
(Mitchell et al., 1990; Parker et al., 1991; Rava-
glia et al., 1996; Anon., 1999). In California,
 Preharvest Factors Affecting Quality
a minimum of 10% SSC for yellow-fleshed
peaches and nectarines was proposed as a
quality standard (Kader, 1995). In France, a
minimum of 10% SSC for low-acidity (TA
<0.9%) and 11% SSC for high-acidity (TA
≥0.9%) peaches was proposed as part of their
quality standard (Hilaire, 2003). In Italy, a
minimum of 10% SSC for early-season, 11%
for mid-season and 12% for late-season culti-
vars was suggested for yellow-fleshed peaches
(Testoni, 1995; Ventura et al., 2000). In prelimi-
nary studies carried out in California by using
trained panels and ‘in-store’ consumer accep-
tance tests on ‘Ivory Princess’ (white flesh/low
TA), ‘Elegant Lady’, ‘O’Henry’ and ‘Spring
Bright’ (yellow flesh/high TA) peaches and
‘Honey Kist’ (yellow flesh/low TA) nectarine,
it was shown that acceptance correlated well
with ripe soluble solids concentration or the
ratio of ripe soluble solids concentration to
ripe titratable acidity; it was also shown that
the relationship was strictly dependent on
cultivar and/or maturity and that consumer
acceptance was not a linear relationship
(Crisosto and Crisosto, 2005).
The analyses of traditional fruit quality
traits are cheap and fast, but they do not con-
sider other fundamental aspects of quality,
such as antioxidant capacity, aroma volatile
emission, soluble sugars and organic acids
content. A more accurate definition of fruit
quality would require sophisticated analyses
(high-performance liquid chromatography,
gas chromatography or mass spectrometry)
that are not usually run because they should
be carried out only in well-equipped laborato-
ries with trained personnel. In any case, simple
or more complex destructive analyses can be
performed only on samples of a limited num-
ber of fruit, often not fully representative of
the entire lot (Costa et al., 2002, 2003b). In recent
years, extensive research has been focused on
the development of non-destructive techniques
for assessing internal fruit quality attributes.
These techniques offer a number of advan-
tages, including: the possibility to extend the
assessments on a large number of, or even on
all, the fruit in a lot; to repeat the analysis on
the same samples, monitoring their physio-
logical evolution; and to achieve real-time
information on several fruit quality parameters
at the same time (Abbott, 1999). Among the
539
non-destructive techniques, near infrared
spectroscopy can be used efficiently for deter-
mining traditional peach fruit quality traits
and concentrations of the main organic acids
and simple sugars. In addition, this technique
allowed definition of a new maturity index
strictly related to the fruit ethylene emission
and ripening stage. This index, called ‘absor-
bance difference’ (AD), can be effectively
used for determining harvest date and for
grouping harvested fruit into homogeneous
classes which show different ripening rates
during shelf-life (Costa et al., 2006).
As a final consideration, as new plant-
ings are based on new cultivars with different
organoleptic characteristics (low- and high-
acid, high SSC, highly aromatic, non-melting,
etc.) and since new markets and consumer
groups with different ethnic backgrounds are
being reached (Liverani et al., 2002; Crisosto,
2003), it is important to understand which
characters are determining consumer accep-
tance and segregate cultivars into different
organoleptic categories prior to proposing
any quality index (Crisosto, 2002, 2003).
As a long-term solution, it is expected
that breeding programmes will include qual-
ity characteristics in their screening process.
The creation of peach categories with their
own quality indices according to an organo-
leptic description may help marketing and
promotion.
20.4 Genotype
Genotype (cultivar and/or rootstock) has an
important role in flavour quality, nutrient
composition and postharvest life potential.
SSC and acidity are determined by several
factors such as cultivar (Crisosto et al., 1995,
1997; Frecon et al., 2002; Liverani et al., 2002;
Byrne, 2003) and rootstock (Reighard, 2002).
Reduction of physiological disorders and
even decay and insect losses can be achieved
by choosing the correct genotype for given
environmental conditions. Extensive harvest
quality and postharvest storage potential eval-
uations have been carried out since 1970 by
several researchers in all the main important
peach cultivation areas, such as the USA,
540 C.H. Crisosto and G. Costa
Italy, Spain, France, Chile and South Africa.
Brown rot and grey mould resistance have
not been successfully included in recently
released cultivars. These are the main dis-
eases, although other ones have been investi-
gated (Frecon et al., 2002; Reighard, 2002), but
current breeding programmes are constantly
creating new cultivars with improved pro-
duction and visual appearance attributes.
Unfortunately, an ideal cultivar(s) with all of
the current consumer quality attributes for
domestic and long-distance shipping has
(have) not been developed yet.
20.5 Mineral Nutrition
Nutritional status is an important factor of
quality and postharvest life potential. Defi-
ciencies, excesses or imbalances of various
nutrients may result in disorders that can limit
storage life. Fertilization rates vary widely
among growers, locations and cultivars, and
generally depend upon soil type, cropping
history and field testing results.
Nitrogen
This is the nutrient that has been studied the
most. N has the single greatest effect on peach
quality. Detailed and extensive research per-
formed since the early 1990s at the Kearney
Agricultural Center (Parlier, California, USA)
has evaluated the role of N in peach and nec-
tarine production and quality under Califor-
nia conditions (Daane et al., 1995). Based on
Table 20.1. Relationship between leaf nitrogen and per cent of fruit surface that is red, yield and fruit
size (mean for 3 years) on ‘Fantasia’ nectarine. (Adapted from Daane et al., 1995.)
N-fertilization
treatment (kg N/ha) Leaf N (%)
0 2.7a
112 3.0b
196 3.1c
280 3.5d
364 3.5d
a,b,cValues within columns with unlike superscript letters were significantly different by the Least
Significant Difference test (P < 0.05).
this work, in California, N should be kept
between 2.6 and 3.0% leaf N for best fruit
quality without reduction in production or
size (Table 20.1). Similarly, optimal fruit qual-
ity in nectarines in the Eastern Po Valley area
(Italy) was obtained in trials having 3.0% leaf
N concentration (Tagliavini et al., 1997; Scu-
dellari et al., 1999). Response of peach and nec-
tarine trees to N fertilization is dramatic; high
N levels stimulate vigorous vegetative growth,
causing shading out and death of lower fruit-
ing wood. Although high-N trees may look
healthy and lush, excess N does not increase
fruit size, production or SSC. Furthermore,
excessive N delays peach maturity because it
induces poor visual red colour development
(Fig. 20.2/Plate 227) and inhibits ground
colour change from green to yellow. As grow-
ers delay harvest waiting for fruit colour
changes from green to yellow and red colour
development, high-N fruit are picked soft
especially when measured on the softest posi-
tion on the fruit such as tips, which generally
ripen faster than the rest of the fruit in warm
production areas. These fruits then have fast
softening rates during postharvest handling
and are more susceptible to bruising and
decay development. N deficiency leads to
small fruit with poor flavour and unproduc-
tive trees. Fruit water loss from fruit with the
highest N rate (3.6% leaf N) was greater than
that from the lowest rate (2.6% leaf N). The
relationship between fruit N concentration
and fruit susceptibility to decay produced by
brown rot (Monilinia fructicola (Wint.) Honey)
has been studied extensively on stored fruit
(Daane et al., 1995). Wounded and brown rot-
inoculated fruit from ‘Fantasia’ and ‘Flavortop’
Fruit visual
redness (%) Yield (kg/tree) Fruit weight (g)
92a 132a 131a
80b 207b 166b
72c 193b 168b
69c 222b 169b
70c 197b 167b
 Preharvest Factors Affecting Quality
trees having more than 2.6% leaf N were more
susceptible to brown rot than fruit from trees
with 2.6% leaf N or less. Fruit anatomical
observations and cuticle density measure-
ments indicated differences in cuticle thick-
ness among ‘Fantasia’ fruit from the low,
middle and high N treatments. These changes
in cuticle and epidermis anatomy can par-
tially explain the differences in fruit suscepti-
bility to this disease and water loss.
Calcium
The nutrient Ca is involved in numerous bio-
chemical and morphological processes in
plants and has been implicated in many dis-
orders of considerable economic importance
to production and postharvest quality. While
Ca accumulation in apple, kiwifruit and grape
occurs predominantly in the first stages of
541
Fig. 20.2. Influence of increased
nitrogen fertilization (kg/ha) on red
skin coloration of ‘Fantasia’ nectarine.
fruit development, in peaches, owing to their
ability to maintain significant transpiration
rates, Ca continues to accumulate until har-
vest (Tagliavini et al., 2000). Foliar Ca sprays
have not been successful and are not used
commercially to maintain peach fruit quality.
Over the last decade, trials carried out in Cal-
ifornia using several commercial Ca foliar
sprays on peach and nectarine (applied every
14 days, starting 2 weeks after full bloom and
continuing until 1 week before harvest) showed
no effect on fruit quality of mid- or late-season
cultivars (Crisosto et al., 2000). These foliar
spray formulations and new formulations did
not affect fruit SSC, firmness, decay incidence,
fruit flesh Ca concentration or postharvest
disorders. Fruit flesh Ca concentration mea-
sured at harvest varied among cultivars from
200 to 300 µg/g dry weight basis. A lack of
decay control was also reported on ‘Jerseyland’
peaches, grown in Pennsylvania, treated with
ten weekly preharvest Ca sprays of CaCl2 at 0,
542 C.H. Crisosto and G. Costa
34, 67 or 101 kg/ha (Conwall, 1987). Even
fruit treated at a rate of 101 kg/ha, which had
70% more flesh Ca (490 versus 287 µg/g dry
weight basis) than untreated fruit, showed no
reduction in decay severity. Our recent
research suggests that any Ca spray formula-
tions and timing on peaches and nectarines
should be treated with caution because their
heavy metal content (Fe, Al, Cu, etc.) may
contribute to peach and nectarine skin discol-
oration (Crisosto et al., 1999). A moderate and
cultivar-dependent effect of Ca sprays on the
reduction of skin russeting development has
been reported for nectarines in Italy (Scudellari
et al., 1995).
Potassium
K is the major nutrient present in peaches
(about 2–2.5 kg/t fresh weight basis), where it
accumulates progressively as fruit approach
maturity (Tagliavini et al., 2000). Optimal K
nutrition usually leads to high photosynthetic
rates and reallocation of sugars and organic
acids that will enhance fruit quality.
Iron
Fe, as a micronutrient, is taken up by fruit
trees in relatively small amounts; however, its
deficiency not only affects fruit yields but also
peach fruit quality (Álvarez-Fernández et al.,
2003). In a study carried out in Spain, only
47% of fruits from Fe-deficient trees had opti-
mal fruit size compared with 95% from green
trees (Álvarez-Fernández et al., 2003). Peach
fruit colour could also be affected by Fe defi-
ciency: in a red-skin peach cultivar (‘Baby-
gold’) Fe deficiency caused decreases in the
mean ‘a’ colour coordinate and increases in
the mean ‘L’ and ‘b’ colour coordinates (Álvarez-
Fernández et al., 2003).
20.6 Irrigation
Despite the important role of water in fruit
growth and development, few specific studies
have been done on the influence of the amount
and the timing of water applications on peach
quality at harvest and postharvest perfor-
mance (Prashar et al., 1976). An early report
indicated that when trees were allowed to
grow without irrigation during the growing
season on a shallow soil under California
conditions, yield and fruit size were reduced,
SSC increased and fruit developed an abnor-
mal texture (Uriu et al., 1964). Reducing the
amount of applied water after harvest of early-
season peaches (postharvest stress) has shown
no negative effects on yield in California;
however, timing of the water deficit interval
is important. An increase in fruit defects such
as deep suture and double-fruit formation
has been reported for early-season ‘Regina’
peaches as a consequence of imposing a post-
harvest water stress (50% evapotranspiration;
ET) in mid- and late summer during the pre-
vious season (Fig. 20.3/Plate 228). These
defects reduced the final packout for the next
season’s crop (Johnson et al., 1992).
The regulated irrigation deficit (RID) tech-
nique has been evaluated for peach perfor-
mance in different production areas (Chalmers
et al., 1981; Ben Mechlia et al., 2002; Girona,
2002; Goldhamer et al., 2002). In general, this
technique imposes a moderate stress (30–50%
ET) to reduce vegetative growth and save water
use (4–30%) at a given physiological stage with-
out affecting yield. Researchers agree that the
water stress-tolerant phases in peach, which
has a double-sigmoid fruit development pat-
tern, have been identified as stage II, the lag
phase of fruit growth and the postharvest
period (Goldhamer et al., 2002). In some situa-
tions, besides saving water, the RID technique
also increased fruit size and SSC. Researchers
claim that consistency of the benefits of the
RID technology will depend on the under-
standing of local climatic conditions, soil
depth and composition, identification of the
fruit growth stages and fruit crop load (Berman
and DeJong, 1996; Girona, 2002). In California,
during three seasons, the influence of three dif-
ferent irrigation regimes applied 4 weeks before
harvest on ‘O’Henry’ peach quality and post-
harvest performance was evaluated: (i) normal
irrigation (100% evapotranspiration); (ii) over-
irrigation (150% ET); and (iii) RID irrigation
(50% ET) (Crisosto et al., 1994; Johnson and
 Preharvest Factors Affecting Quality
Fig. 20.3. Water stress late in the summer causes fruit defects such as deep sutures and double-fruit
formation.
Handley, 2000). Yield, flesh firmness, per cent
red surface, acidity and pH were not altered
at harvest by any of these irrigation regimes
in any season. Average fruit size, measured as
fruit weight, was lower but SSC was higher
for fruit from 50% ET than from the other
treatments. Ripe yellow-fleshed peaches and
nectarines with 10% SSC or higher with low
to moderate TA (<0.7%) are highly acceptable
to consumers. Although fruit from the 50%
ET treatment were smaller, they had higher
SSC and consumers would probably prefer
their eating quality over fruit from the other
two treatments. An economic study showed
that peaches with a higher SSC may have a
higher retail value (Parker et al., 1991). The
irrigation regimes (100%, 50% and 150% ET
applied 4 weeks before harvest) did not affect
‘O’Henry’ peach postharvest storage poten-
tial based on internal breakdown develop-
ment during 2, 4 and 6 weeks in cold storage
at 0°C or 5°C. Fruit from 50% ET had a lower
water loss rate than fruit from 150% ET or
100% ET. Fruit from 150% ET lost nearly 35%
543
more water than fruit from 50% ET or 100%
ET after 24 h. Light microscopy studies indi-
cated that fruits from 50% ET and 100% ET
had a continuous and much thicker cuticle
and a higher density of trichomes than fruits
from the 150% ET. These differences in exo-
dermis structure may explain the higher per-
centage of water loss from fruit from 150% ET
compared with the others (Crisosto et al.,
1994).
Recently, RID and partial root zone dry-
ing (PRD) were evaluated on white-fleshed
peach growing under California conditions
(Goldhamer et al., 2002). PRD involves induc-
ing partial stomatal closure by exposing some
part of the root zone to continual soil drying.
After 2 years of evaluations, yield and fruit
quality were affected equally by the PRD and
the RID treatments. Except for a few studies
which have comprehensively tested a broad
range of water management practices and
conditions and their impacts on postharvest
quality, it is often difficult to generalize about
the effects of water management from the
544 C.H. Crisosto and G. Costa

site-specific irrigation regimes that have been 14

reported. (a) O’Henry r2 = 0.72
13

12
20.7 Canopy Manipulation

SSC (%)
11
In most cultivars, fruitlet thinning increases
fruit size while also reducing total yield, thus 10
May Glo r2 = 0.67
a balance between yield and fruit size must be
9
achieved. Some cultivars must not be thinned
too much because their fruit will crack easily. 8
In some cases, fruit size, SSC and TA are mod- 300
ified without affecting fruit cracking. In other (b)
cultivars the fruit do not ripen properly when 250 O’Henry r2 = 0.82
trees are carrying too high a fruit load. In gen- Fruit weight (g)
eral, the number of fruit that can ripen on a 200
tree will depend on the cultivar and orchard
150
conditions. Thus detailed information about
cultivar response to crop load adjustment and 100
potential benefits should be developed for
each specific situation. Historically, maximum 50 May Glo r2 = 0.64
profit does not occur at maximum marketable
yield since larger fruit bring a higher market 0 100 200 300 400 500
price. Furthermore, new market trends for Crop load (1000s fruit/ha)
highly tasty fruit may force a review of this
topic. The crop load and fruit quality relation- Fig. 20.4. Relationship between (a) crop load
ship has been studied by researchers in vari- and soluble solids concentration (SSC) and
ous countries (Forlani et al., 2002; Giacalone (b) crop load and fruit weight for ‘O’Henry’
et al., 2002; Luchsinger et al., 2002; Costa et al., peach and ‘May Glo’ nectarine. (Adapted from
2003a). Leaving too many fruit on a tree Crisosto et al., 1997.)
reduces fruit size and SSC in the early-season
‘May Glo’ nectarine and late-season ‘O’Henry’
peach (Fig. 20.4). Crop load on ‘O’Henry’ peach a longer shelf-life (storage and market) than
trees affected the incidence of internal break- fruit grown under a low light environment
down. In general, the overall incidence of (inside canopy). During our work, we found
mealiness and flesh browning in fruit from that fruit that developed in the more shaded
the high crop load was low, intermediate in inner canopy positions have a greater inci-
fruit from the commercial crop load, and the dence of internal breakdown than fruit from
highest in fruit from the low crop load the high light, outer canopy positions (Fig. 20.5/
(Crisosto et al., 1997). Plate 229). Thus, fruit from the outer canopy
Fruit quality measured at harvest and have a longer potential market life, especially
during storage for several peach and nectar- for cultivars susceptible to internal break-
ine cultivars varied according to fruit canopy down. The use of more efficient training sys-
position in different production areas (Marini tems which allow more sunlight penetration
et al., 1991; Crisosto et al., 1997; Iannini et al., into the centre and lower canopy areas is rec-
2002). Large differences in SSC, acidity and ommended to reduce the number of shaded
fruit size were detected between fruit obtained fruit, thus extending postharvest life (Crisosto
from the outside and inside canopy positions et al., 1997).
of open-vase trained trees (Marini et al., 1991; Summer pruning and leaf removal
Crisosto et al., 1997). During the last decade, around the fruit increase fruit light exposure
we have observed that fruit grown under a and, when performed properly, can increase
high light environment (outside canopy) has fruit colour without affecting fruit size and
 Preharvest Factors Affecting Quality
SSC (Fig. 20.6/Plate 230). Excessive leaf pull-
ing or leaf removal executed too close to har-
vest can reduce both fruit size and SSC in
peaches and nectarines (Crisosto et al., 1997;
Day, 1997). Girdling (removal of bark) 4–6
weeks before harvest is performed to increase
peach and nectarine fruit size (Fig. 20.7/Plate
231) and to advance and synchronize matu-
rity (Day, 1997). Girdling increases fruit SSC
in some cases but also increases fruit acidity
and phenolics, so the flavour resulting from
the additional SSC may be masked. Girdling
can also cause the pits of peaches and nectar-
ines to split, especially if it is done too early
during pit hardening. Fruit with split pits
soften more quickly than intact fruits and are
more susceptible to decay.
Reports on the benefits of using different
reflective materials to improve peach red
545
Fig. 20.5. Canopy position affects
fruit size, red colour development and
storage potential.
colour and fruit size and speed up maturation
varied according to cultivar, orchard situation
and location (Layne et al., 2001; Fiori et al.,
2002). Under California’s long and hot grow-
ing season, canopy manipulations including
water sprout removal and leaf removal
around fruit become necessary to achieve the
benefit of red colour development in vigor-
ous orchards. Also, even when reflected light
was reaching fruit in the canopy, but temper-
atures remained high during that maturation
period, improvement in red colour develop-
ment was not observed.
In spite of the limited literature available
on the role of preharvest factors in consumer
quality, there is strong evidence that fruit fla-
vour quality, market life and physiological
disorders are related to preharvest factors. We
therefore encourage more detailed work on
546 C.H. Crisosto and G. Costa

Fig. 20.6. Leaf removal around the fruit improves red colour but may decrease fruit size.

Fig. 20.7. Peach girdling (removal of a strip of scaffold bark) at the main scaffolds advances maturity
and increases fruit size.
 Preharvest Factors Affecting Quality 547

this subject with an emphasis on consumer
satisfaction. In order to maximize ‘orchard
quality potential’, all of the preharvest factors
influencing quality must be investigated by
physiologists and understood by pomologists.
Detailed information on how these factors are
controlling peach consumer quality combined
with an effective marketing programme will
help to increase peach consumption (Crisosto,
2002).

References

Abbott, J.A. (1999) Quality measurement of fruits and vegetables. Postharvest Biology and Technology 15,
207–223.
Álvarez-Fernández, A., Paniagua, P., Abadía, J. and Abadía, A. (2003) Effects of Fe deficiency-chlorosis on
yield and fruit quality in peach (Prunus persica L. Batsch). Journal of Agricultural and Food Chemistry 51,
5738–5744.
Ames, B.N., Shigenaga, M.K. and Hagen, T.M. (1993) Oxidants, antioxidants and the degenerative diseases of
aging. Proceedings of the National Academy of Sciences USA 90, 7915–7922.
Anderson, J.W., Allgood, L.D., Lawrence, A., Altringer, L.A., Jerdack, G.R., Hengehold, D.A. and Morel, J.G. (2000)
Cholesterol-lowering effects of psyllium intake adjunctive to diet therapy in men and women with hyper-
cholesterolemia: meta-analysis of 8 controlled trials. American Journal of Clinical Nutrition 71, 472–479.
Anon. (1999) Peche-nectarine: eliminer la non quality N. INFOS-CTFL 151, 11.
Ben Mechlia, N., Ghrab, M., Zitouna, R., Ben Mimoun, M. and Masmoudi, M. (2002) Cumulative effect over
five years of deficit irrigation on peach yield and quality. Acta Horticulturae 592, 301–308.
Berman, M.E. and DeJong, T.M. (1996) Water stress and crop load effects on fruit fresh and dry weights in
peach (Prunus persica). Tree Physiology 16, 859–864.
Bregoli, A.M., Scaramagli, S., Costa, G., Sabatini, E., Ziosi, V., Biondi, S. and Torrigiani, P. (2002) Peach (Pru-
nus persica) fruit ripening: aminoethoxyvinylglycine (AVG) and exogenous polyamines affect ethylene
emission and flesh firmness. Physiologia Plantarum 114, 472–481.
Bregoli, A.M., Ziosi, V., Biondi, S., Rasori, A., Ciccioni, M., Costa, G. and Torrigiani, P. (2005) Postharvest
1-methylcyclopropene application in ripening control of ‘Stark Red Gold’ nectarines: temperature-
dependent effects on ethylene production and biosynthetic gene expression, fruit quality, and polyamine
levels. Postharvest Biology and Technology 37, 111–121.
Byrne, D. (2003) Breeding peaches and nectarines for mild-winter climate areas: state of the art and future
directions. In: Marra, F. and Sottile, F. (eds) Proceedings of the First Mediterranean Peach Symposium,
Agrigento, Italy, 10 September, pp. 102–109.
Chalmers, D.J., Mitchell, P. and van Heek, D. (1981) Control of peach growth and productivity by regulated
water supply, tree density and summer pruning. Journal of the American Society for Horticultural
Science 106, 307–312.
Conwall, W. (1987) Effects of preharvest and postharvest calcium treatments of peach on decay caused by
Monilinia fructicola. Plant Disease 71, 1084–1086.
Costa, G., Miserocchi, O. and Bregoli, A.M. (2002) NIRs evaluation of peach and nectarine fruit quality in
pre- and post-harvest conditions. Acta Horticulturae 592, 593–599.
Costa, G., Bregoli, A.M. and Vizzotto, G. (2003a) La regolazione della carica dei frutti nel pesco: analisi del
processo e possibili soluzioni. In: Atti IV Convegno Nazionale sulla Peschicoltura Meridionale, Campo-
bello di Licata, Agrigento, Italy, 11–12 September, pp. 52–61.
Costa, G., Noferini, M., Montefiori, M. and Brigati, S. (2003b) Non-destructive assessment methods of kiwi-
fruit quality. Acta Horticulturae 610, 179–190.
Costa, G., Noferini, M., Fiori, G. and Ziosi, V. (2006) Internal fruit quality: how to influence it, how to define
it. Acta Horticulturae 712, 339–345.
Crisosto, C.H. (2002) How do we increase peach consumption? Acta Horticulturae 592, 601–605.
Crisosto, C.H. (2003) Searching for consumer satisfaction: new trend in the California peach industry. In:
Marra, F. and Sottile, F. (eds) Proceedings of the First Mediterranean Peach Symposium, Agrigento, Italy,
10 September, pp. 113–118.
Crisosto, C.H. and Crisosto, G.M. (2005) Relationship between ripe soluble solids concentration (RSSC) and
consumer acceptance of high and low acid melting flesh peach and nectarine (Prunus persica (L.) Batsch)
cultivars. Postharvest Biology and Technology 38, 239–246.
548 C.H. Crisosto and G. Costa

Crisosto, C.H., Johnson, R.S., Luza, J.G. and Crisosto, G.M. (1994) Irrigation regimes affect fruit soluble solids
content and the rate of water loss of ‘O’Henry’ peaches. HortScience 29, 1169–1171.
Crisosto, C.H., Mitchell, F.G. and Johnson, R.S. (1995) Factors in fresh market stone fruit quality. Postharvest
News and Information 6, 17–21.
Crisosto, C.H., Johnson, R.S., Day, K.R. and DeJong, T. (1997) Orchard factors affecting postharvest stone fruit
quality. HortScience 32, 820–823.
Crisosto, C.H., Johnson, R.S., Day, K.R., Beede, B. and Andris, H. (1999) Contaminants and injury induce
inking on peaches and nectarines. California Agriculture 53, 19–23.
Crisosto, C.H., Day, K.R., Johnson, R.S. and Garner, D. (2000) Influence of in-season foliar calcium sprays on
fruit quality and surface discoloration incidence of peaches and nectarines. Journal of the American
Pomological Society 54, 118–122.
Daane, K.M., Johnson, R.S., Michailides, T.J., Crisosto, C.H., Dlott, J.W., Ramirez, H.T., Yokota, G.T. and
Morgan, D.P. (1995) Excess nitrogen raises nectarine susceptibility to disease and insects. California
Agriculture 49, 13–17.
Day, K.R. (1997) Production practices for quality peaches. Proceedings of Florida State Horticultural Association
77, 59–61.
Doll, R. (1990) An overview of the epidemiological evidence linking diet and cancer. Proceedings of the
Nutrition Society 49, 119–131.
Dragsted, L.O., Strube, M. and Larsen, J.C. (1993) Cancer-protective factors in fruit and vegetables: biochemical
and biological background. Pharmacology and Toxicology 72, 116–135.
Fiori, G., Bucchi, F., Corelli Grappadelli, L. and Costa, G. (2002) Effetto dteli riflettenti a terra sugli scambi
gassosi e la qualità delle produzioni in pesco. In: Atti VI Giornate Scientifiche SOI, Spoleto, Italy, 23–25
April, pp. 157–158.
Forlani, M., Basile, B., Cirillo, C. and Iannini, C. (2002) Effects of harvest date and fruit position along the tree
canopy on peach fruit quality. Acta Horticulturae 592, 459–466.
Frecon, J.L., Belding, R. and Lokaj, G. (2002) Evaluation of white-fleshed peach and nectarine varieties in
New Jersey. Acta Horticulturae 592, 467–478.
Giacalone, G., Peano, C. and Bounous, G. (2002) Correlation between thinning amount and fruit quality in
peaches and nectarines. Acta Horticulturae 592, 479–484.
Girona, J. (2002) Regulated deficit irrigation in peach. A global analysis. Acta Horticulturae 592, 335–342.
Goldhamer, D.A., Salinas, M., Crisosto, C., Day, K.R., Soler, M. and Moriana, A. (2002) Effects of regulated
deficit irrigation and partial root zone drying on late harvest peach tree performance. Acta Horticulturae
592, 343–350.
Hilaire, C. (2003) The peach industry in France: state of art, research and development. In: Marra, F. and Sottile,
F. (eds) Proceedings of the First Mediterranean Peach Symposium, Agrigento, Italy, 10 September, pp.
27–34.
Iannini, C., Cirillo, C., Basile, B. and Forlani, M. (2002) Estimation of peach yield efficiency and light intercep-
tion by the canopy in different training systems. Acta Horticulturae 592, 357–366.
Johnson, R.S. and Handley, D.F. (2000) Using water stress to control vegetative growth and productivity of
temperate fruit trees. HortScience 35, 1048–1050.
Johnson, R.S., Handley, D.F. and DeJong, T. (1992) Long-term response of early maturing peach trees to post-
harvest water deficit. Journal of the American Society for Horticultural Science 69, 1035–1041.
Kader, A.A. (1988) Influence of preharvest and postharvest environment on nutritional composition of fruits
and vegetables. In: Quebedeaux, B. and Bliss, F.A. (eds) Horticulture and Human Health Contributions
of Fruits and Vegetables. Prentice-Hall, Englewood Cliffs, New Jersey, pp. 18–22.
Kader, A.A. (1995) Fruit maturity, ripening, and quality relationships. Perishables Handling Newsletter 80, 2.
Layne, D.R., Jiang, Z.W. and Rushing, J.W. (2001) Tree fruit reflective film improves red skin coloration and
advances maturity in peach. HortTechnology 11, 234–242.
Liverani, A., Giovannini, D. and Brandi, F. (2002) Increasing fruit quality of peaches and nectarines: the main
goals of ISF-FO (Italy). Acta Horticulturae 592, 507–514.
Luchsinger, L., Ortin, P., Reginato, G. and Infante, R. (2002) Influence of canopy fruit position on the maturity
and quality of ‘Angelus’ peaches. Acta Horticulturae 592, 515–522.
Marini, R.P., Sowers, D. and Marini, M.C. (1991) Peach fruit quality is affected by shade during final swell of
fruit growth. Journal of the American Society for Horticultural Science 116, 383–389.
Mathooko, F.M., Tsunashima, Y., Owino, W.Z.O., Kubo, Y. and Inaba, A. (2001) Regulation of genes encoding
ethylene biosynthetic enzymes in peach (Prunus persica L.) fruit by carbon dioxide and 1-methylcyclo-
propene. Postharvest Biology and Technology 21, 265–281.
 Preharvest Factors Affecting Quality 549

Mitchell, F.G., Mayer, G., Biasi, W., Gulli, D. and Faubian, D. (1990) Selecting and handling high quality stone
fruit. California Tree Fruit Agreement 1989 Research Report. California Tree Fruit Agreement, Sacramento,
California.
Parker, D., Ziberman, D. and Moulton, K. (1991) How quality relates to price in California fresh peaches.
California Agriculture 45, 14–16.
Prashar, C.R.K., Pearl, R. and Hagan, R.M. (1976) Review on water and crop quality. Scientia Horticulturae 5,
193–205.
Ravaglia, G., Sansavini, S., Ventura, M. and Tabanelli, D. (1996) Indici di maturazione e miglioramento qual-
itativo delle pesche. Frutticoltura 3, 61–66.
Reighard, G.L. (2002) Current directions of peach rootstock programs worldwide. Acta Horticulturae 592,
421–428.
Ruperti, B., Cattivelli, L., Pagni, S. and Ramina, A. (2002) Ethylene-responsive genes are differentially regu-
lated during abscission, organ senescence and wounding in peach (Prunus persica). Journal of Experi-
mental Botany 53, 429–437.
Scudellari, D., Tagliavini, M. and Pelliconi, F. (1995) Aspetti teorici ed applicativi del calcio nelle colture
arboree. L’Informatore Agrario LI 15, 45–49.
Scudellari, D., Toselli, M., Marangoni, B. and Tagliavini, M. (1999) La diagnostica fogliare nelle piante arboree
da frutto a foglia caduca. Bollettino della Società Italiana di Scienza del Suolo 48, 829–842.
Tagliavini, M., Scudellari, D., Corelli Grappadelli, L. and Pelliconi, F. (1997) Valutazione di metodi rapidi per
stimare il livello azotato del pescheto. In: Atti XXII Convegno Peschicolo, Cesena, Italy, 5–7 October
1995, pp. 141–150.
Tagliavini, M., Zavalloni, C., Rombolà, A.D., Quartieri, M., Malaguti, D., Mazzanti, F., Millard, P. and
Marangoni, B. (2000) Mineral nutrient partitioning to fruits of deciduous trees. Acta Horticulturae 512,
131–140.
Testoni, A. (1995) Momento di racolta, qualita, condizionamento e confezionamento delle pesche. In: Pro-
ceedings of the Symposium ‘La peschicoltura Veronesa alle soglie del 2000’, Verona, Italy, 25 February,
pp. 327–354.
Trainotti, L., Zanin, D. and Casadoro, G. (2003) A cell-oriented genomic approach reveals a new and unex-
pected complexity of the softening in peaches. Journal of Experimental Botany 54, 1821–1832.
Trainotti, L., Bonghi, C., Ziliotto, F., Zanin, D., Rasori, A., Casadoro, G., Ramina, A. and Tonutti, P. (2006) The
use of microarray µPEACH1.0 to investigate transcriptome changes during transition from pre-climacteric
to climacteric phase in peach fruit. Plant Science 170, 606–613.
Uriu, K., Wereniels, P.G., Retan, A. and Fox, D. (1964) Cling peach irrigation. California Agriculture 18, 10–11.
Ventura, M., Sama, A., Minguzzi, A., Lanzón, S. and Sansavini, S. (2000) Ottimizzazione del carico di fruti
per migliorare la produzione e la qualita delle nectarine Supercrimson e Venus. In: Proceedings of XXIV
Convengo Peschiolo, Cesena, Italy, 24–25 February, pp. 173–176.
Ziosi, V., Bregoli, A.M., Borghi, C., Fossati, T., Biondi, S., Costa, G. and Torrigiani, P. (2006) Transcript levels of
ethylene perception and biosynthesis genes as altered by putrescine, spermidine and aminoethoxyvinyl-
glycine (AVG) during the course of ripening in peach fruit (Prunus persica L. Batsch). New Phytologist
172, 229–238.
 21 Ripening, Nutrition and Postharvest
Physiology

A. Ramina,1 P. Tonutti2 and W. McGlasson3

1Department of Environmental Agronomy and Crop Science, University of Padova,
Legnaro, Italy
2Sant’Anna School of Advanced Studies, Pisa, Italy
3Centre for Plant and Food Science, University of Western Sydney, Sydney,

New South Wales, Australia

21.1 Introduction 550

21.2 The Main Processes Defining the Ripening Syndrome 551
Respiration 551
Ethylene biosynthesis and perception 552
Flesh softening and melting 555
Organic acid and sugar metabolism 557
Colour and aroma development 561
21.3 Nutritional Value and Healthiness 562
21.4 Postharvest Physiology 564
Genetic differences among cultivars 564
Effect of low temperature 564
Effect of controlled atmosphere and modified atmosphere packaging 565
21.5 Future Perspectives and Conclusions 566

21.1 Introduction which have higher quality but shorter storage

Peach fruits ripen rapidly and have a short
postharvest life, usually limited to 3–4 weeks
depending on storage conditions. They are
climacteric fruit that ripen on the tree or after
harvest if picked mature. Crop maturity at
harvest strongly influences quality and shelf-
life. Determination of ideal harvest maturity
is critical to maximize yield, fruit size and
consumer acceptability. Generally, early-har-
vested fruits have good storability but they
are of lower quality than late-harvested fruits,
life. Immature peaches will not ripen to des-
sert quality, whereas those harvested too late
will be too soft to withstand commercial han-
dling. Early picking results in poor flavour
and a bitter acid taste, which is due to rela-
tively low sugar concentrations and a high
content of organic acids, polyphenolics and
aldehydes (Robertson et al., 1988; Horvat et al.,
1990). Typical peach flavour compounds such
as d-decalactone, g-decalactone, linalool and
benzaldehyde that contribute to fruit fra-
grance are low or absent in fruit harvested

550 © CAB International 2008. The Peach: Botany, Production and Uses
(eds D.R. Layne and D. Bassi)
 Ripening, Nutrition and Postharvest Physiology
immature (Horvat et al., 1990; Visai et al., 1993).
Change in ground colour is a useful maturity
index (Eccher Zerbini et al., 1991), because it
changes along with other important parame-
ters such as soluble solids, flesh firmness and
volatile compounds. Ground colour can be
estimated by comparison with colour charts
(Meredith et al., 1989) or it can be measured
with an electronic colorimeter according to the
CIE system (McGuire, 1992). Neri and Brigati
(1994) proposed several minimal instrumental
parameters that peach fruits should possess at
harvest in order to meet quality standards.
These parameters include firmness no more
than 45 N, positive ‘a’ values and soluble sol-
ids content not less than 12%. The relationship
between changes in ground colour and firm-
ness of fruit ripening on the tree is affected by
the light environment in which a fruit devel-
ops (Lewallen and Marini, 2003). Newer tech-
nologies that use time-resolved reflectance
spectroscopy offer the prospect of machine
grading of fruit maturity (Eccher Zerbini et al.,
2006; Tijskens et al., 2007). These technologies
are based on the relationship between de-
greening and softening of the flesh during rip-
ening. In practice, compromises among fruit
maturity, yield and possible higher prices for
early-picked fruit are often adopted. An immu-
nological assay of ripening-related proteins is
feasible for establishing the physiological age
of the fruit (Abdi et al., 2002). The changes in
the ripening-related proteins would be related
to changes in external appearance to assist
pickers to select mature fruit.
Although perishability is the limiting
factor in postharvest handling and marketing
of peaches, final quality and consumer accep-
tance and some postharvest responses are
also affected by preharvest factors, such as
growing conditions, cultural practices and
genetic potential. Various preharvest stresses
may have significant impact on fruit flavour,
weight, overall appearance and storage
potential (Crisosto et al., 1997).
Peach fruit ripening can be slowed by
application of postharvest technologies that
reduce metabolic activity. These technologies
include cooling the fruit and/or establishing
controlled and/or modified atmosphere condi-
tions (low concentration of oxygen and elevated
concentration of carbon dioxide). Application
551
of these technologies reduces respiration rate,
ethylene biosynthesis and action as well as
other biochemical processes. However, these
technologies may negatively affect specific
quality parameters and induce the develop-
ment of physiological disorders (Lill et al., 1989).
Thus, the knowledge of ripening physiology
and the postharvest behaviour of peach fruits
provides the basis for improving quality and
maintaining high eating quality along the mar-
keting chain. In the last decade many biochemi-
cal and molecular processes that operate during
peach fruit ripening have been elucidated and
ways to optimize postharvest practices to main-
tain shelf-life have been proposed.
21.2 The Main Processes Defining the
Ripening Syndrome
Respiration
Peach is classified as a climacteric fruit because
respiration increases during ripening. Com-
pared with other species the average respira-
tion rates of peaches are classified as moderate
(Wills et al., 2007). According to the growth
stages defined by Tonutti et al. (1991), respiration
rates are high (about 150 cm3 CO2/kg per h)
during stage I of fruit development, when
growth occurs predominantly by cell divi-
sion, decreasing through stage II (pit harden-
ing) and part of stage III (second exponential
growth), rising gradually at the end of stage
III and finally reaching the climacteric peak at
stage IV (ripening). However, climacteric
peaks are highly variable and range from 15
to 80 cm3 CO2/kg per h according to geno-
type, including yellow versus white flesh,
melting versus non-melting (Brady, 1993;
Ventura et al., 1998; Brovelli et al., 1999). In
some genotypes, the increase in respiration at
ripening is so moderate that the typical cli-
macteric behaviour is not readily apparent.
Marked differences have also been observed
in relation to the duration of the fruit devel-
opmental cycle, which dictates the harvest
time: the shorter the developmental cycle
(and the earlier the harvest date), the higher
and more pronounced the respiration rate at
the climacteric (Ventura et al., 1998).
552 A. Ramina et al.
Ethylene biosynthesis and perception
Ethylene plays a crucial role in the initiation
and continuation of ripening of climacteric
fruit. An increase of ethylene biosynthesis is
associated with several plant developmental
events. Internal ethylene concentrations are
less than 1 µl/l during fruit growth and devel-
opment up to the onset of ripening. This basal
level refers to ‘system 1’ of ethylene biosyn-
thesis. At stage IV ethylene evolution begins
to increase, reaching a peak usually associ-
ated with the last stage of ripening (Tonutti
et al., 1991). This constitutes the ‘system 2’ of
ethylene biosynthesis, characterized by the
autocatalytic action of the hormone (McMurchie
et al., 1972). As a consequence of this autocata-
lytic action, the application of exogenous ethyl-
ene or one of its analogues, such as propylene,
to mature fruit at 10 µl/l for 24–48 h at 20–25°C
is sufficient to enhance ripening, although less
mature fruits need continuous exposure to the
hormone for longer periods. Also in peach
fruit, the biosynthesis of ethylene requires the
action of two key enzymes, 1-aminocyclopro-
pane-1-carboxylate (ACC) synthase (ACS)
and ACC oxidase (ACO), responsible for the
formation of ACC and the oxidation of ACC
to ethylene, respectively, as demonstrated in
early works (Miller et al., 1988; Amoros et al.,
1989; Tonutti et al., 1991). At ripening, peach
whole fruit and mesocarp tissue show a tem-
poral difference in ethylene biosynthesis. A
gradient in biosynthesis between mesocarp
and epicarp has been detected. The epicarp is
more efficient in converting ACC to ethylene
(Tonutti et al., 1991). In several peach cultivars
(‘Springcrest’, ‘Redhaven’, ‘Fayette’, ‘Bailey’,
‘Suncrest’) ethylene evolution begins to increase
when fruits have already started to soften and
the highest biosynthetic rates occur in the lat-
est stage of ripening (Fig. 21.1a). Thus, peaches
should be listed among those fruit species in
which the ethylene climacteric coincides with
or follows eating ripeness. In peach this stage
corresponds to flesh firmness values of about
30–40 N. Huge variation exists among culti-
vars in terms of ethylene evolution rates at
ripening (Ventura et al., 1998). Low levels of
ethylene biosynthesis are not necessarily
related to slower rates of loss of firmness or to
the non-melting trait (Brovelli et al., 1999).
Stony hard (hd) peaches are characterized by
the absence of both ethylene production at
ripening and postharvest softening. How-
ever, melting and non-melting hd genotypes
have been distinguished by the degree of
softening following exogenous ethylene treat-
ments (Haji et al., 2003, 2005). In the ‘Fantasia’
nectarine cultivar and in some of its progenies
a correlation between ethylene evolution and
softening rate has been observed (Brecht and
Kader, 1984). Within these progenies an inter-
esting ripening mutation was found (see
Chapter 1, ‘Time of ripening’ section) called
‘slow ripening’. This mutant does not show a
characteristic pattern of ripening in terms of
ethylene evolution and respiration: in fact, it
remains firm and green and does not exhibit a
rise in carbon dioxide or ethylene production
for at least 4 weeks after harvest at 20°C. Eth-
ylene production remains at a low basal level
for 4 weeks and begins to increase before the
respiratory rise, reaching only 15–33% of that
detected in the wild type. When exposed to
exogenous ethylene during the pre-climacteric
stage the response of these fruits is similar to
that of non-climacterics. Short exposure to eth-
ylene provokes a respiratory rise but does not
affect the rate of ethylene production (Brecht
and Kader, 1984). The accumulation of high lev-
els of conjugated ACC in peaches is a curiosity.
Amoros et al. (1989) and Uthairatanakij et al.
(2005) reported conjugated ACC concentrations
ten times greater or more than those of ACC.
Three ACS isogenes (Pp-ACS1, Pp-ACS2
and Pp-ACS3) have been reported in peach
but only Pp-ACS1 is associated with ripening
(Mathooko et al., 2001; Tatsuki et al., 2006).
Ethylene production at ripening is strictly
related to increases in ACC content and ACS
activity and related transcripts. Analysis of
the accumulation pattern of specific mRNA
indicates that, in peach, Pp-ACS1 is develop-
mentally controlled. Southern blot analysis
revealed that, as in other species, ACS is
encoded by a multigene family. In tomato
these members are differently regulated and
the polymorphism of ACS has been claimed
to be involved in the transition from system 1
to system 2 of ethylene biosynthesis (Nakat-
suka et al., 1998; Barry et al., 2000). Preharvest
application of aminoethoxyvinylglycine (AVG)
has been shown to delay ripening and to
 Ripening, Nutrition and Postharvest Physiology 553

(a) 70 10

60 8

50
6
Firmness (N)

40
4

30
2

20
0
10
75 80 85 90
Days after full bloom

(b)

21,5 30,8 60,3 185,7

Pp-ACO1 expression

75 80 85 90
Days after full bloom

0,014
(c)
Pp-ETR1 Pp-ERS1
0,012

0,010
Transcripts (A.U.)

0,008

0,006

0,004

0,002

0,000
Pre-climacteric Climacteric
Ripening stage

Fig. 21.1. Firmness and ethylene evolution (a), 1-aminocyclopropane-1-carboxylate oxidase (ACO)
activity and Pp-ACO1 transcript accumulation (b) and Pp-ETR1 and Pp-ERS1 expression (c) during the
final stages of fruit development of cultivar ‘Springcrest’. (Parts a and b modified from Tonutti et al.,
1997; part c modified from Rasori et al., 2002.)
554 A. Ramina et al.
allow further increase in fruit size (Rath and
Prentice, 2004). AVG is a strong inhibitor of
ACS activity (Boller et al., 1979). Work with
nectarines suggested that AVG may act on
other enzyme systems in addition to ACS.
Softening was retarded in AVG-treated fruit
ripened at 20°C soon after harvest, whereas
AVG-treated fruit ripened after 1 week of
storage at 1.0–1.5°C softened at the same rate
as untreated fruit (McGlasson, et al., 2005).
Recently it has been shown that the reduced
level of ethylene production in stony-hard
peaches, causing a lack of softening, is due to
a suppression of Pp-ACS1 transcription at rip-
ening (Hayama et al., 2006; Tatsuki et al., 2006)
(for flesh texture phenotypes, see Chapter 1,
‘Flesh texture’ section). Interestingly, the same
gene appeared to be transcribed in wounded
immature, pre-climacteric and climacteric
fruits. This would indicate that one possible
mechanism of repression of Pp-ACS1 mRNA
is the interruption of the ripening-related
transcriptional activity by some insertion or
deletion in the 5′-flanking region of Pp-ACS1,
which contains a cis-regulatory domain of the
ripening-related sequences. Another possible
reason is the disruption of a transcriptional
factor specifically activated to induce Pp-
ACS1 mRNA at ripening (Tatsuki et al., 2006).
Callahan et al. (1992, 1993) and Lester et al.
(1994) were the first to isolate ACO cDNA
clones. They demonstrated that ACO tran-
scripts strongly accumulated during the late
stages of ripening, consistent with the patterns
of ethylene evolution and ACO activity (Fig.
21.1b) (Tonutti et al., 1991). In mature peach
fruit, the first detectable increase in ACO-
related transcripts takes place when ethylene
biosynthesis is still at a basal level and before
textural changes occur (Lester et al., 1996).
This pattern indicates that peach ACO gene
expression is under developmental control
and that it is ethylene-regulated. This has
been confirmed by Ruperti et al. (2001), who
isolated and characterized two members of
the peach ACO gene family, named Pp-ACO1
and Pp-ACO2. Pp-ACO1 is organized in four
exons interrupted by three introns whereas
Pp-ACO2 has only two of the three introns of
Pp-ACO1. Comparison of deduced amino
acid sequences of the two genes revealed
77.7% identity. Pp-ACO1 transcripts accumu-
late strongly in ripe mesocarp as well as in
abscising fruitlets and senescing leaves, and are
enhanced by ethylene. Pp-ACO2 mRNA is
detected in fruits only during early develop-
ment and is unaffected by ethylene. Functional
analysis (Rasori et al., 2003) showed that, within
the promoter region of the gene, ethylene-re-
sponsive elements are present in Pp-ACO1 but
not in Pp-ACO2. This may account for the dif-
ferent responsiveness to exogenous ethylene of
the two genes. In addition, two auxin-respon-
sive elements, probably responsible for the
auxin suppression of the ethylene induction of
Pp-ACO1 gene expression are present upstream
of the ethylene-responsive elements. Specific
regulatory elements affecting transcription in a
tissue-specific manner as well as a region con-
trolling the temporal expression of the gene
have been identified in the promoter of Pp-
ACO1 (Moon and Callahan, 2004).
There have been tremendous advances
in elucidation of the mechanisms of ethylene
perception and signal transduction in Arabi-
dopsis (Chang et al., 1993; Hua et al., 1995).
Recently, two peach genes, Pp-ETR1 and Pp-
ERS1, showing a similar organization to that
of the corresponding genes in Arabidopsis
have been isolated (Rasori et al., 2002). As
observed in other species (Bleeker, 1999),
these two genes belong to a multigene family.
Southern blot analysis carried out with the
specific probe for Pp-ERS1 suggests that, in
peach, at least one other gene related to Pp-
ERS1 exists. The presence of two genes encod-
ing ERS type protein has only been reported
in Arabidopsis (Hua et al., 1998). The deduced
proteins of the two genes contain a sensor
domain and a hystidine kinase domain, in
which residues thought to be important for
the normal function of ETR and ERS type pro-
tein as ethylene receptors are conserved.
These results indicate that Pp-ETR1 and Pp-
ERS1 could be putative ethylene receptors
with the ability to bind ethylene in peach.
Quantitative RT-PCR data showed that Pp-
ETR1 and Pp-ERS1 transcripts are differen-
tially expressed in immature and ripe fruit
(Fig. 21.1c). Pp-ETR1 appears to be constitu-
tive and ethylene-independent during fruit
development and ripening, whereas Pp-ERS1
transcripts increase during fruit ripening and
its expression appears to be upregulated by
 Ripening, Nutrition and Postharvest Physiology
ethylene (Rasori et al., 2002). The correlation
between ripening and expression of Pp-ERS1
has been confirmed by experiments with
1-methylcyclopropene (1-MCP), a competitor of
ethylene for the binding sites. The continuous
application of the chemical delayed fruit ripen-
ing, evaluated in terms of softening, decay and
ethylene evolution, and concurrently downreg-
ulated Pp-ERS1 while Pp-ETR1 transcription
was unaffected. 1-MCP action was rapidly abol-
ished after moving fruits to air, when a stimula-
tion of ethylene evolution and a concurrent
increase in Pp-ERS1 mRNAs were observed.
Recent investigations pointed out that the ethyl-
ene biosynthetic and signal transduction path-
ways are differently affected by 1-MCP in apple
and peach (Dal Cin et al., 2006). The chemical is
effective in delaying ripening in apples while
in peaches it has only a limited effect. This dif-
ferent behaviour has been attributed to differ-
ences in terms of ratio, expression patterns
and/or turnover of the ethylene receptors.
Flesh softening and melting
Most fruit soften during ripening and this is
the major quality attribute that often dictates
shelf-life. Fruit softening could arise from dif-
ferent processes: loss of turgor, degradation
of starch and breakdown of the fruit cell walls.
Loss of turgor is largely a non-physiological
process associated with postharvest dehydra-
tion of the fruit. Degradation of starch proba-
bly results in a pronounced textural change,
especially in fruits like banana, where starch
accounts for a high percentage of the fresh
weight. Changes in texture occurring during
the ripening of most fruit are thought to be
largely the result of cell wall degradation.
Carbohydrate polymers make up 90–95% of
the structural components of the wall, the
remaining 5–10% being largely hydroxyproline-
rich glycoproteins. The carbohydrate polymers
can be grouped together as cellulose, hemicel-
luloses and pectins. In melting peaches flesh
firmness decreases slowly at the beginning of
ripening (softening stage). During the later
stage of ripening the loss of firmness is rapid
(melting stage). Also in the peach, flesh soften-
ing has been related to the modification of
555
pectic polymers which undergo solubilization
and depolymerization, particularly during the
melting stage (Dawson et al., 1992). Besides
pectins, hemicelluloses also change their
apparent molecular mass during peach soften-
ing, resulting in decreased tissue cohesion
(Hedge and Maness, 1998). Cell wall enzymes
are responsible for these changes and a number
of cell wall hydrolases have been identified
in fruit tissue. The most important enzymes
include pectinmethylesterase (PME) (EC
3.1.1.15), polygalacturonase (PG) (EC 3.2.1.15),
b-(1,4)-glucanase (EG) (EC 3.2.1.4) and b-galac-
tosidase (b-GAL) (EC 3.2.1.23). These are either
constitutive throughout development and
ripening, or present in very low amounts and
exist as several isoforms within the fruit tis-
sue. In general, PME and b-GAL activities are
present during the development of the fruit.
Both PG and EG tend to be absent in mature
green fruit but their activities become mea-
surable only with the onset of ripening and
increase dramatically afterwards. Both of these
enzymes tend to be predominantly endo-act-
ing hydrolases, although exo-acting PG activ-
ity is also found during ripening in several
fruits. Considering the complexity of both wall
structure and enzyme profiles it is unlikely
that a single enzyme is responsible for textural
change. This process probably involves a com-
plex interaction of enzyme activity with physi-
cochemical changes in the cell wall.
In melting peaches, PG activity increases
during ripening (Pressey et al., 1971) and exo-
and endo-acting PG have been identified in
ripe peaches (Pressey and Avants, 1973). The
endo-acting enzyme (EC 3.2.1.15; endoPG) is
found in significant amounts only in melting
peaches; the exo-enzyme (EC 3.2.1.67; exoPG)
has comparable activity in the mesocarp of
both melting and non-melting fruit (Pressey
and Avants, 1978). Downs and Brady (1990)
found two forms of exoPG that increase dur-
ing ripening. EndoPG and exoPG activity
increase gradually as fruit soften, but the rate
of increase in activity accelerates when the
fruit are very soft (<20 N) (Downs et al., 1992;
Orr and Brady, 1993). These findings suggest
that initiation of softening is not associated
with exo- and endoPG activity. In the early-
ripening nectarine cultivar ‘Armking’, PME
activity appeared to be more closely related
556 A. Ramina et al.
with the decrease in flesh firmness than PG,
even though the increase in enzyme activity
measured throughout ripening was higher
for PG than for PME (Artes et al., 1996). The
approach to study PG at the molecular level
in peach started from the evidence that tomato
and peach endoPG are immunologically
related. Using a tomato PG cDNA, a 3.5 kb
fragment of peach DNA was isolated and 3.25
kb was subcloned and sequenced. The 3′ ends
of tomato and peach PG showed extensive
homology in the coding region but some dif-
ferences in part of the gene structures. The
presence of several bands in Southern blot
analyses suggests that endoPG in peach is
encoded by a multigene family with more
than three genes (Lester et al., 1994). Expres-
sion analyses carried out with the partial PG
cDNA confirmed that the accumulation of
specific mRNA in ‘Flavorcrest’ (melting) fruit
occurred in a pattern similar to that of
increases in endoPG activity (Lester et al.,
1994). Thus, no hybridization was detected in
fruits with flesh firmness values ranging from
120 to 40 N, but only when the firmness value
dropped below 5 N did a strong signal appear.
These data confirm the role played by endoPG
during the melting stage in peach fruit. A
cDNA, PRF5, was identified as a fruit-related
endoPG that may be involved with the tex-
ture differences (Lester et al., 1996). Callahan
et al. (2004) and Morgutti et al. (2006) found a
relationship between the non-melting trait
and a deletion in at least one of the PRF5-
related PG genes. Recent findings revealed the
existence of a single locus containing at least
one gene encoding an endoPG controlling
both freestone and melting flesh traits with at
least three effective alleles (Peace et al., 2005).
As far as EG (also known as cellulase) is
concerned, it hydrolyses the b-1,4-glucan link-
ages of plant cell wall polymers that contribute
to weakening of the wall structure during
processes (such as ripening) characterized by
cell separation phenomena. Hinton and Pressey
(1974) found during ripening that EG activity
increases before any significant change in
fruit firmness. Similar results have been
obtained by Bonghi et al. (1998), who found
that the increase of EG activity, occurring at
early softening and stimulated by propylene,
was due to a pronounced increase in the basic
isoform. These data confirm that there is a
relationship between ethylene and EG and
show that EG is mainly involved in the initial
phase of peach fruit softening, before the
action of PME and PG.
The presence of different EG isoforms
suggests that a multigene family is present in
peach as observed in other plant species. Three
cDNA clones (pCel10, pCel20 and pCel30) have
been isolated from peach leaf and fruit abscis-
sion zone RNA: the three clones showed a
high degree of divergence and different
expression patterns (Trainotti et al., 1997).
Only transcripts related to pCel10 have been
detected with a Northern blot analysis in leaf
and, to lesser extent, in flower abscission zones.
RNase protection assay (RPA) confirmed that
pCel20- and pCel30-related mRNAs scarcely
accumulate during abscission and ripening
and that pCel10-related transcripts are barely
detectable during fruit softening. The screen-
ing of a genomic library with the cDNA clone
pCel10 allowed the isolation of a peach EG
gene named PpEG1, which encodes a poly-
peptide of 497 amino acids with a predicted
molecular mass of 54.3 kDa that shares the
highest similarity (76.3%) to the avocado rip-
ening EG. PpEG1, a member of the small
peach EG family, is present in the peach
genome as a single-copy gene. Using a 753 bp
PCR product, identical to the corresponding
sequence of PpEG1, Bonghi et al. (1998) were
unable to detect related transcripts when
Northern blot analysis was performed on
poly(A) + RNA extracted from peach meso-
carp during early and late climacteric stages.
A lack of signal in these tissues was also
observed when RPA was performed, indicating
the low abundance of EG transcripts in peach
fruit tissues. EG transcripts were detected in
fruits at all stages of fruit development only
following RT-PCR. The highest accumulation
of EG-specific transcript corresponded to the
onset of the ethylene climacteric and the early
phase of flesh softening. Treating pre-climac-
teric peach fruits with propylene, besides
enhancing EG activity, strongly accelerated
specific transcript accumulation and the loss
of firmness within 24 h. In the same experi-
ments the use of a specific inhibitor of ethyl-
ene action (2,5-norbornadiene) reduced these
parameters. Differently from PpEG1, PpEG4,
 Ripening, Nutrition and Postharvest Physiology
another member of the peach EG family, shows
a decreasing expression pattern through rip-
ening (Trainotti et al., 2006b), being down-
regulated by ethylene (Begheldo et al., 2008).
Expansins are proteins that have been
shown to contribute to tomato fruit softening.
Recently, some expansins have been related
to a loss of fruit firmness in peach (Hayama
et al., 2003). Three expansins, named Pp-Exp1,
Pp-Exp2 and Pp-Exp3, have been isolated from
ripe peach fruit: all three expansins were
detected in the fruit and not in other tissue,
although each showed a different pattern of
expression during fruit development. Pp-Exp2
RNA was expressed constitutively throughout
fruit development but was more abundant
in stage III, during exponential growth and
maturation. Pp-Exp1 and Pp-Exp3 were up-
regulated at the onset of ripening but Pp-Exp1
was induced at an earlier stage. However, by
comparing cultivars with different rates of
softening, Pp-Exp3 shows a closer association
with softening.
A cell wall-oriented genomic approach
revealed a new and unexpected complexity in
the softening of peaches (Trainotti et al., 2003).
The genes analysed encode proteins involved
in the main metabolic aspects of a primary
cell wall (degradation, synthesis, structure)
(Fig. 21.2). In addition, some genes encoding
cell wall-related proteins with an unknown
function have been found. The gene expres-
sion profiles showed that softening begins
well before the ethylene climacteric rise and
continues thereafter. Genes whose expression
begins before the climacteric rise are mostly
downregulated by ethylene, while those that
are ripening-specific are mostly upregulated
by the hormone. A few other genes are appar-
ently insensitive to ethylene. According to the
expression analysis of the genes involved in
cell wall degradation, pectatelyase seems to
play an interesting role. Specific transcripts
for this enzyme are downregulated by ethyl-
ene but they increase markedly during the
early stage of softening and before the highest
accumulation of PG transcripts. Besides the
expected partial degradation, softening pro-
cesses include some repair of the cell walls per-
formed by enzymes involved in the synthesis
of cell wall polysaccharides, especially by pro-
teins with structural functions. The newly
557
synthesized polysaccharides and structural
proteins would thus help to maintain cell wall
integrity while not preventing softening (Trai-
notti et al., 2003).
Organic acid and sugar metabolism
Peach organoleptic quality depends on colour,
texture and flavour attributes that are mostly
determined during ripening (Brady, 1993). In
contrast, acidity, which is an essential compo-
nent of the fruit taste, is thought to be deter-
mined during the early stages of fruit
development. In peach, as in most fruits, the
two major organic anions are malate and cit-
rate (amounting to 140 and 120 mM H+,
respectively), which account for most of the
titratable acidity and, as a general trend in
high-acid cultivars, tend to decrease during
ripening. Major differences exist between
high- and low-acid (sub-acid) cultivars: for
example, in ripe fruit of ‘Fantasia’, which is
an acidic cultivar, juice titrable acidity is about
400 mmol l−1 H+, while in ‘Jalousia’ (low-acid)
it is around 40 mmol l−1 H+ (Moing et al., 1998)
(Fig. 21.3). Low acidity is a quantitative trait
regulated by a single dominant gene (Yoshida,
1970; Monet, 1979; see Chapter 1, ‘Flesh com-
pounds’ section). In acidic cultivars, malate
accumulates mainly in early fruit develop-
ment, slows down during stage II and
increases moderately in stage III, while citrate
increases mainly during late stage III. In low-
acid cultivars a reduced accumulation of
malate at early growth stage and citrate at
late stage III has been observed (Moing et al.,
2000). It was proposed that phosphoenolcar-
boxylase (PEPC) plays a major role in control-
ling the fruit organic acid metabolism (Moing
et al., 1999). PEPC has been regarded as the
key enzyme because it catalyses the b-carbox-
ylation of phosphoenolpyruvate to yield
oxaloacetate and inorganic phosphate. The
oxaloacetate is reduced by NAD-dependent
malate dehydrogenase to produce malate.
Oxaloacetate and malate can enter the tricar-
boxylic acid cycle to produce citrate and other
metabolites. PEPC mRNA and specific pro-
tein levels peaked at early development and
late stage III in ‘Fantasia’. In ‘Jalousia’, both
558 A. Ramina et al.

(a) (b)
3.5 2.0 Fuc-tra Ctg 190
PG Ctg 219
PAE Ctg 337 Gls-tra Ctg 285
3.0
1,3-EG Ctg 364 Cell Syn Ctg 907
1.5
2.5 Exp Ctg 404 Acyl-tra Ctg 6
PL Ctg 251 Glu-tran Ctg 254
2.0
PL Ctg 257 1.0 SuSy Ctg 393
1.5 PME Ctg 874
1,4-EG Ctg 124
1.0 0.5
β-Gal Ctg 167

0.5

0.0 0.0

ctg
(c) (d)
187
4.0 PRP Ctg 125 1.00 528
3.5 Leu-rich p Ctg 152 188

E6-like p Ctg 10 191

3.0 0.75 418
Extensin like Ctg 95
2.5 444
Leu-rich p Ctg 288
501
2.0 0.50 612
1.5 821
153
1.0 0.25
839
0.5 762

0.0 0.0
S1 S2 S3 S4 S1 S2 S3 S4

Fig. 21.2. Using a genomic approach, expression patterns of cDNA clones throughout fruit development
and ripening (stage I to stage IV, S1 to S4) were grouped according to three different functions as cell
wall weakening (a), cell wall synthesis (b), cell wall structure (c) and unknown (d). Among the previ-
ous clones assessed for sensitivity to ethylene at pre-climacteric and climacteric stages, 11 (ctg 219,
404, 167, 364, 874, 254, 10, 152, 187, 418, 839) are upregulated, and nine (ctg 257, 251, 124, 125, 95, 393,
6, 153, 188) are downregulated by ethylene. Five (ctg 190, 285, 907, 528, 821) are apparently insensitive
to ethylene. Values on the ordinate axis are expressed as the ratio of the subtracted volumes of the
indicated expressed sequence tags and ubiquitin. Arrows indicate ethylene climacteric. (Modified
from Trainotti et al., 2003.)

parameters were very low at early fruit devel-
opment and reached levels similar to ‘Fanta-
sia’ at late stage III. For both cultivars in vitro
PEPC activity was maximal at early fruit devel-
opment, decreased from 24 to 60 days after
bloom and then remained constant. The activ-
ity of fruit PEPC appears to be extremely sen-
sitive to malate and low pH. PEPC may
participate in the control of organic acid accu-
mulation through fruit development in high-
acid cultivars. However, mechanisms other than
organic acid synthesis may account for differ-
ences in acidity between acidic and low-acid
fruits. Six peach cDNAs encoding key proteins
in organic acid metabolism and solute accu-
mulation have been isolated and character-
ized (Etienne et al., 2002). The genes involved in
organic acid metabolism (mitochondrial citrate
synthase, cytosolic NAD-dependent malate
dehydrogenase and cytosolic NADP-depen-
dent isocitrate dehydrogenase) showed a
stronger expression in ripening fruit than
during the earlier phase of development,
although their expression patterns were not
necessarily correlated with changes in organic
acid content. Concerning genes involved in

5.0
4.5
Juice pH
4.0
3.5
3.0
0 20
500
Juice titratable acidity (mM H+)
400
300
200
100
0 standard deviation represented
0 20
storage, the tonoplast proton pump showed a
biphasic expression pattern more consistent
with the pattern of organic acid accumulation
and the tonoplast pyrophosphatases were more
highly expressed in the fruit of low-acid culti-
var during the second rapid growth phase.
Sweetness is the most important factor
affecting consumer acceptability of peaches.
Sucrose, glucose and fructose in proportion of
about 3:1:1 are the main sugars in peaches
(Genard et al., 2003). Peaches with high eating
quality are considered to have relatively large
amounts of fructose and low quantities of
glucose and sorbitol (Brooks et al., 1993)
because fructose is considered to be 3.0, 2.3 and
1.7 times sweeter than sorbitol, glucose and
sucrose, respectively (Kulp et al., 1991). The
sugars comprise more than 60% of the soluble
solids concentration (SSC) as measured with
Ripening, Nutrition and Postharvest Physiology 559
Jalousia
Fantasia
40 60 80 100 120 140
Time after bloom (days)
Jalousia
Fantasia
Fig. 21.3. pH and titratable acidity
of ‘Jalousia’ and ‘Fantasia’ peaches
during growth and development.
Values are means, with their
40 60 80 100 120 140 by vertical bars (n = 5 fruit). (From
Time after bloom (days) Moing et al., 1998.)
a refractometer. There is a general consensus
that an SSC reading of at least 12% is required
to ensure that peaches are acceptable to most
consumers. However, acidity can modify fla-
vour perception. Cultivars have been bred
with low, medium or high acidity (Moing
et al., 1998). In practice, SSC among fruit of
similar size harvested at similar maturity
from the same group of trees can vary by 8%.
This variation among a population of fruit
has serious implications for retailers wishing
to provide peaches with guaranteed sweet-
ness, especially in seasons when assimilation
rates are low. Genard et al. (2003) developed
models to describe the relationships between
sugar concentrations in fruit and assimilate
supply, metabolism and dilution. The SUGAR
model (Fig. 21.4; Genard and Souty, 1996) has
been used to relate the fluxes in carbon among
560 A. Ramina et al.

Sucrose k1(t)

λph CO2
k1(t)

Carbon supply Glucose Fructose

k4(t)

k2(t)

1–λph Other
compounds
Sorbitol
k3(t)

Fig. 21.4. Diagram of the SUGAR model. The arrows and boxes represent carbon fluxes and carbon
components, respectively. The two ellipses represent carbon supply and losses by respiration. The
proportion of sucrose in the phloem-sourced sugar pool (lph) and the relative rates of sugar
transformation k1(t), k2(t), k3(t) and k4(t) for each arrow are indicated. The model predicts the
partitioning of carbon into sucrose, sorbitol, glucose and fructose into the mesocarp of peach fruit.
The model can be used to simulate the effects of changing assimilate supply and fruit volume on
sugar concentrations. (From Genard et al., 2003.)

sugars, sorbitol, other compounds and respi-
ration. Glucose and fructose are accumulated
at a nearly constant rate throughout develop-
ment and ripening (Souty et al., 1998). The
glucose:fructose ratio decreases during matu-
ration: being nearly 1 during stage III, it falls
to 0.8 at ripening, showing that glucose is
preferentially used for respiration. Sucrose
accounts for the major part of the increase in
dry weight, and its concentration at ripening
represents more than 50% of the dry weight.
There is a good correlation between sucrose
accumulation and the increase in fruit dry
weight. Sucrose accumulates rapidly during the
last few days of maturation on the tree, reaching
a plateau at the ethylene climacteric. These data
suggest a negative role for ethylene in sucrose
accumulation since pre-climacteric applica-
tions of AVG and polyamines delay the ethyl-
ene climacteric and SSC increase at harvest
(Bregoli et al., 2002; Vizzotto et al., 2002).
Although sorbitol is generally low in
peach fruit, this sugar alcohol and sucrose
account for most of the carbon translocated in
the phloem from leaves to the fruit. After
unloading in the fruit, sorbitol is rapidly
metabolized to glucose and fructose by sorbi-
tol oxidase and sorbitol dehydrogenase,
respectively. It has been shown that the meso-
carp exhibits a capacity to actively accumu-
late sugars and that this tissue has different
sucrose synthesis and cleavage activities,
including sorbitol dehydrogenases, sucrose
synthases and invertases (Vizzotto et al., 1996).
A full-length cDNA for NAD-dependent sor-
bitol dehydrogenase (NAD-SDH) from peach
has been isolated and characterized (Yamada
et al., 2001). Peach NAD-SDH activity on a
fresh weight basis was very strong in imma-
ture fruit, declined temporarily and then
increased again with fruit maturation. Protein
level and enzyme activity paralleled tran-
script accumulation. The authors concluded
that NAD-SDH plays an important role in
sorbitol metabolism during peach fruit devel-
opment, and that the increase in NAD-SDH
activity as fruit matures is regulated by gene
transcription. More recently RT-PCR gene
expression analyses carried out with specific
primers have allowed grouping of genes
according to their differential regulation
(Nonis et al., 2003). Three genes encoding,
 Ripening, Nutrition and Postharvest Physiology
respectively, a putative sucrose synthase
(SuSy2), a hexose transporter and a vacuolar
invertase showed higher levels of transcrip-
tion at the early stages of fruit development
(stage I), when fruit growth was character-
ized by active cell division. Subsequently, the
transcripts of these genes gradually decreased
towards stage II, when endocarp (pit) hard-
ening and seed development primarily take
place. Genes for a hexose transporter and
SuSy2, together with a gene encoding a puta-
tive sucrose phosphate synthase, were clearly
upregulated during fruit ripening (stage IV),
a phase characterized by cell expansion. Three
genes, encoding respectively a cell wall inver-
tase, a cytoplasmic invertase and a sucrose
transporter, also appeared to be transcribed to
some extent during early fruit development.
In addition, there was a transient increase in
their mRNAs coincident with the onset of the
second phase of exponential fruit growth
(stage III) and of sucrose accumulation in
mesocarp cells. The transcription of a neutral
invertase (Pp-NT1) appeared to be regulated
in response to sugar signals only in the phase
of fruit expansion coincident with the onset of
sucrose accumulation (Nonis et al., 2007). A
gene encoding an additional sucrose synthase
(SuSy1) was expressed at low levels in fruits.
Expression data obtained for genes related to
sugar metabolism may point to a role for
these genes during peach fruit growth. In fact,
an association was found between the expres-
sion of some of these genes and the stages of
fruit development during which growth is
powered by cell division. A relationship was
also evident between the transient upregula-
tion of a different group of genes and the
developmental switch leading to sucrose
accumulation and to growth processes driven
by cell enlargement.
Colour and aroma development
Colour changes that are associated with rip-
ening strongly influence visual and eating
quality of peaches. Genotypical differences
markedly affect colour and intensity; for
example, non-melting peaches have signifi-
cantly higher total carotenes and xanthophylls
561
than melting-flesh peaches (Karakurt et al.,
2000). Colour changes are associated with
degradation of chlorophyll, which in turn, in
the yellow-fleshed cultivars, unmasks other
pigments such as carotenoids. The loss of
chlorophyll is accompanied by the biosynthe-
sis of anthocyanins in the epicarp and deep
mesocarp. Lessertois and Moneger (1978),
working on fruits of ‘Earlyglo’ (yellow flesh),
showed that fruit pigments of the mesocarp
are of the foliar type, i.e. chlorophylls, a- and
b-carotene, lutein, epoxylutein, violaxanthin
and neoxanthin. The transformation of the
ovary into the green adult fruit is accompa-
nied by a decline in the content of chloro-
phylls; total carotenoids decrease to levels as
low as 55% of the initial content and subse-
quently transitorily accumulate. In mature
fruits, 42% of the carotenoids are oxidized
and all carotenoids are of the b-configuration.
Later on, the carotenoids disappear but the
chlorophylls remain in traces. More recent
data from ‘Red Star’ (yellow flesh, late ripen-
ing) showed that carotenoid content increases
throughout fruit ripening and this occurs con-
currently with an accumulation of phytoene
synthase, z-carotene desaturase, lycopene
b-cyclase and b-carotene hydroxylase tran-
scripts, as reported, in ‘Fantasia’, by Trainotti
et al. (2006a). However, carotenoid content
decreases during the postharvest phase. Eth-
ylene biosynthesis and action might be
involved in the regulation of carotenoid bio-
synthesis during ripening, since anoxia or
1-MCP treatments applied at a pre-climacteric
stage reduce the level of pigments. 1-MCP
treatment performed in a confined environ-
ment on the tree at early ripening maintains
carotenoid content while it is totally ineffec-
tive if applied during the postharvest phase
(Mencarelli et al., 1998).
Aroma is determined by volatile compo-
nents produced by peaches in concentrations
that can be perceived by the human nose.
Aroma compounds arise from several differ-
ent substrates including fatty acids, amino
acids, phenolics and terpenoids. Aldehydes,
esters, ketones, terpenoids and sulfur-con-
taining compounds account for most of the
important flavour volatiles in fruits. Takeoka
et al. (1988) emphasized the important role of
lactones and other products of the peroxidation
562 A. Ramina et al.
of unsaturated fatty acids by lipoxygenase
(LOX) in nectarines. In unripe peaches most
of the compounds that impart green aroma
are aldehydes and C6 alcohols (hexanal, trans-
2-hexanal, hexanol, trans-2-hexanol), while in
ripe fruit the main components are lactones
(g-decalactone and d-decalactone). Significant
increases in benzaldehyde and linalool have
also been detected. A good marker for peach
aroma is cis-3-hexenylacetate (De Santis and
Mencarelli, 2001). At least some part of the
development of peach aroma is controlled by
ethylene. Application of 1-MCP results in
higher levels of C6 compounds (hexenals and
hexanols) and lower amounts of esters (Men-
carelli et al., 2003). Similarly, it has been shown
in banana fruit that initiation of volatile pro-
duction occurs at a later stage in ripening and
1-MCP stimulates production of aldehydes
and alcohols and suppresses ester production
(Golding et al., 1999).
21.3 Nutritional Value and Healthiness
Fruits are an important part of our daily diet,
given the protective role played by some fruit
components against serious human diseases.
Recent experimental evidence has shown that
fruit have higher ‘global’ antioxidant activity
than either isolated natural single compo-
nents or conventional synthetic antioxidants
used as food additives (Eberhardt et al., 2000).
From a dietary point of view, the most impor-
tant fruit constituents are carotenoids, pheno-
lics and fibre.
Carotenoids, besides being an important
component of fruit colour, are a good source
of vitamin A, which is essential for normal
growth, reproduction and resistance to infec-
tions in man. Carotenoids are converted to
vitamin A after ingestion. Yellow-fleshed
peaches are considered a good source of pro-
vitamin A carotenoids, mainly b-carotene and
b-cryptoxanthin (Gross, 1987). Carotenoids
are unstable when exposed to low pH, oxy-
gen and light (Klein, 1987). Wounding, by
triggering ethylene production and senes-
cence, provokes the oxidation of fatty acids
by LOX and may cause degradation of carote-
noids by co-oxidation (Thompson et al., 1987).
Antioxidant activity of phenolic com-
pounds is closely related to their chemical
structure (Robards et al., 1999), particularly
the hydroxylation level of the B-ring (Dziedzie
and Hudson, 1983) and the extent of glycosy-
lation. Glycosylated forms are normally char-
acterized by a lower antioxidant activity
compared with the aglycones. Flavonols,
which are glycosylated forms of quercetin
and kaempferol, are the most abundant phe-
nolics in peach and other stone fruits (Hen-
ning and Herrmann, 1980; Young et al., 1989,
Bengoechea et al., 1997). Myricetin has been
proposed as a phenolic marker for peaches
since this flavonol is not present in other fruit
species (Fernandez de Simon et al., 1990,
1992). Recent work by Costa and co-workers
(G. Costa, Italy, 2007, personal communica-
tion) showed that the chromatographic pro-
file of phenolics in peaches is characterized
by 17 main peaks, identified on the basis of
their retention times and ultraviolet absor-
bance, and by co-chromatography with com-
mercial standards. The phenolic profile appears
to be consistent among cultivars, although the
concentration of each compound may vary
within cultivars and tissue. Cyanidin and gly-
cosylated quercetin are present mainly in the
epicarp, while the cinnamic acids including
chlorogenic and neochlorogenic acids are the
most abundant and widely distributed in the
mesocarp and epicarp (1.27 and 0.5 mg/g dry
weight, respectively).
In fruits and vegetables complex carbo-
hydrates such as cellulose, hemicellulose and
pectins, which are not digested by humans,
comprise dietary fibre (Salunkhe et al., 1991;
Kader and Barret, 1996; Kader, 2002). It has
been shown that fibre consumption, mainly
the soluble forms, decreases plasma choles-
terol (Ebihara et al., 1979) and glycaemic
response (Sharaftedinov et al., 1999; Southon,
2000). Recent studies have shown that dietary
fibre concentrates from by-products of pro-
cessed fruit and vegetables have total fibre
content (35–58%) higher than cereal brans and
a better insoluble:soluble ratio (Holloway,
1983; Grigelmo and Martin-Belloso, 1999).
Grigelmo et al. (1999) reported that total dietary
fibre constitutes about 31–36% of dry matter
in peach concentrates, with insoluble fibre
being the major fraction (20–24%). The dietary
 Ripening, Nutrition and Postharvest Physiology
fibre content of peach baby foods is higher
than that found in other types of baby foods
(Torija Isasa et al., 1985). Melting and non-
melting flesh peach types differ in pectin con-
tent and solubility, and this influences fruit
softening (Pressey and Avants, 1978; Maness
et al., 1993; Karakurt et al., 2000). These differ-
ences may be of interest from a nutritional
point of view because of the well-recognized
role of fibre in regulating intestine function,
peristaltic movement and, as a consequence,
the hunger and/or satiety sensation. Chemo-
protective activity of peach fruit is under
evaluation in rats and man. Preliminary results
showed that a diet enriched with peach fruit
downregulates several isoforms of the cyto-
chrome P450-dependent monooxygenases
which are involved in the onset of degenerative
diseases. The haematic residual antioxidant
activity varies among cultivars. According to
these preliminary data, cultivars with high
dietary values can be identified (S. Ciapellano,
Italy, 2007, personal communication).
According to recent epidemiological
studies food allergies are increasing in many
Western countries. Allergies to fruits are an
important problem and research should be
aimed at selection of hypoallergenic fruits.
The major allergens in Rosaceae fruits are lip-
id-transfer proteins (LTPs). LTPs are abun-
dant and widespread in nature and form a
broad family of proteins whose role is not
clearly defined. Besides their biological func-
tions, LTPs have been recently identified as
the major allergen of peach (Pastorello et al.,
1999; Sanchez-Monge et al., 1999) and other
stone fruits (Pastorello et al., 2000, 2001). They
cause an allergy syndrome usually restricted
to the oral cavity (oral allergy syndrome),
although systemic reactions, including ana-
phylactic shock, have also been reported
(Ortolani et al., 1988). Recently, Asero et al.
(2000) showed that immunoglobulin E (IgE)
antibodies to stone and pome fruit LTPs react
to a broad range of extracts of different plant
species such as groundnut, walnut, pistachio,
broccoli, carrot, tomato, melon and kiwifruit,
indicating that LTPs represent a genuine pan-
allergen. The 9 kDa peach LTP has been puri-
fied and sequence analysis of the protein
(named Pru p 1) shows a high degree of
homology (average of 65%) with LTPs from
563
other plant species (Pastorello et al., 1999).
The protein extracted from peach fruit skin,
where most LTPs are found (Lleonart et al.,
1992), retains its IgE-binding capacity and
demonstrates a high stability following heat
treatment (Brenna et al., 2000) and pepsin
digestion (Asero et al., 2000).
Two LTP-related cDNAs have been iden-
tified. As observed in several plant species
(Kader, 1996), it has been shown that a multi-
gene family encoding LTPs is present in
peach. Southern blot analysis clearly demon-
strates the presence of at least four or five
genes. The two isolated peach LTP clones, Pp-
LTP1 and Pp-LTP2, share the common features
of plant LTPs (positions of cysteine residues,
lipid-binding motifs) but show only 49.6%
identity of deduced amino acid sequences
(Botton et al., 2002). Low similarity values and
phylogenetic analysis, which grouped Pp-
LTP1 and Pp-LTP2 in two distinct clusters,
show that these are not duplicated genes, as
observed for some members of the LTP family
in other species (Pelese-Siebenbourg et al.,
1994; Pyee and Kolattukudy, 1995; Arondel
et al., 2000), but they have evolved indepen-
dently from a common ancestor. Like Pp-
LTP1, Pp-LTP2 appears to encode a type-1
(9 kDa) LTP but it lacks the conserved proline
and tyrosine residues that characterize the
type 2 (7 kDa) LTPs (Monnet et al., 2001) and
differs significantly from the 7 kDa LTP
sequenced by Conti et al. (2001) in apricot.
The differential expression patterns of
the two isolated clones reinforce the hypoth-
esis of multiple roles for LTPs (Fig. 21.5). In
contrast to Pp-LTP1, Pp-LTP2 transcripts have
been strongly detected in non-pollinated ova-
ries and their level declined during a 4-week
period after pollination. The absence of Pp-
LTP2 transcripts in petals, sepals, stamens
and fruit tissues supports the hypothesis that
the role of this gene is highly specific and con-
fined to pistils, in which it may have different
functions (Botton et al., 2002).
In fruits only Pp-LTP1 is expressed and
its transcripts and related protein abundantly
accumulate in epicarp but not in mesocarp,
with different patterns according to the geno-
type (Botton et al., 2006). Since peach aller-
genic activity is largely concentrated in the
skin, it is likely that Pp-LTP1 encodes the
564 A. Ramina et al.

in Pp-LTP1 transcripts and related protein

AB

-28 B
B
B

A
DA
accumulation indicate that it is possible to

P- 7DA
P- 14D
P- 21D
select hypoallergenic cultivars (A. Botton, G.

-
-
-
-O
Ov
Ov
Ov
Ov
Ov
Pasini and P. Tonutti, Italy, 2007, personal
Np
P-
P-
(a) Ptl Spl Stm
communication).
Pp-LTP1

21.4 Postharvest Physiology

Pp-LTP2 Genetic differences among cultivars

In addition to differences in flesh colour, sugar

(b) SI SII SIII PC CR CP
Pp-LTP1
Pp-LTP2
Fig. 21.5. Northern blot analysis carried out with
Pp-LTP1 and Pp-LTP2 cDNA probes on total RNA
(10 µg) extracted from (a) petals (Ptl), sepals (Spl),
stamens (Stm), ovaries at different developmental
stages (Np-Ov, non-pollinated ovary; P-Ov, pol-
linated ovary; P-Ov-7DAB, P-Ov-14DAB,
P-Ov-21DAB and P-Ov-28DAB, pollinated ovary
7, 14, 21 and 28 days after full bloom, respectively)
and (b) epicarp collected at stage I (SI, first
exponential growth phase), stage II (SII,
endocarp lignification), stage III (SIII, second
exponential growth phase), pre-climacteric (PC),
climacteric rise (CR) and climacteric peak (CP).
(Modified from Botton et al., 2002.)
major allergen of peach, although it cannot be
ruled out that also other members of the peach
LTP family (or other proteins) may account for
allergenicity. These results confirm that peel-
ing is essential for the production of hypoal-
lergenic peach foods. The similar transcript
levels detected throughout ripening suggests
that the allergenicity due to LTP does not
change during the last developmental stage
(Botton et al., 2002). Moreover, it has been
shown that postharvest storage at low tem-
perature does not affect Pp-LTP1 transcript
accumulation. In addition, preliminary assays
showing a large variation within genotypes
levels and acidity, peaches display differences
in duration of fruit development, flesh texture,
fruit shape and strength of attachment of flesh
to the pit. Worldwide, most cultivars are mid- to
high-chill, which mature fruit in early summer
to autumn. These cultivars have a fruit devel-
opment period (FDP) in excess of 100 days.
More than 30 years ago low-chill cultivars that
flower after less than 250 chill units were
introduced on to the market. In countries
with a wide range of climatic conditions such
as Australia, the introduction of low-chill cul-
tivars has extended the peach season to about
6 months. The FDP for low-chill peaches can
be as short as 70 days, but even in medium-
chill cultivars very early-ripening peaches are
available. The slow second phase of fruit
growth is absent in these fruit and their growth
in size follows essentially a sigmoidal pattern.
Early-ripening cultivars may receive high
prices when they first arrive on the market
but frequently consumers are disappointed
because the fruit rarely accumulate more than
10% SSC. A disappointing experience early in
the season can deter consumers from buying
peaches until much later, even though SSC
has reached acceptable levels. Breeders are
continuously seeking ways to improve SSC
levels in early-maturing cultivars by selec-
tion, use of rootstocks that grow well when
soil temperatures are still low and enclosures
to create microclimates that improve peach
tree growth (Nissen et al., 2005a,b).
Effect of low temperature
Storage at low temperatures is the most prac-
tical way to slow metabolism and extend the
 Ripening, Nutrition and Postharvest Physiology
commercial life of fruits (see also Chapter 22).
Both respiration and ethylene biosynthesis
are decreased when peach fruits are exposed
to low temperatures, which also reduce the
activity of cell wall hydrolases, thus delaying
softening. Therefore, cold storage is used to
slow ripening and decay development even
though chilling injury may limit the storage
life of peaches and nectarines under low tem-
perature (Lurie and Crisosto, 2005). Consid-
ering ethylene physiology, no increase in ACC
accumulation has been reported in peaches or
nectarines stored at low temperatures either
before physiological disorders had begun or
after their appearance (Uthairatanakij et al.,
2005). It must be noticed that, in prolonged
cold storage, maintaining the ability of the
fruit to produce ethylene may be crucial to
prevent chilling-related physiological disor-
ders (Dong et al., 2001; Zhou et al., 2001).
These disorders include a loss of the ability to
soften normally when the fruit are returned to
normal temperatures, flesh browning, spread
of anthocyanin pigments through the flesh
(bleeding) and gel breakdown. The most com-
mon symptom is a loss of juiciness and the
development of a mealy texture. This disor-
der is called mealiness or woolliness. Meali-
ness develops in fruit stored at less than 8°C.
These disorders are not typical of chilling
injury in tropical fruit because best storage
life of 2–4 weeks depending on cultivar is
achieved near 0°C, but symptoms may also
develop following a week of storage at about
5°C (Anderson, 1979). Conditioning at about
20°C for up to 48 h before refrigerated storage
or delayed storage sometimes improves the
cold storage life of peaches (Zhou et al., 2001;
Crisosto et al., 2004). Mealy peaches have a
dry texture (cork-like), lack flavour, become
brown and less juice can be extracted from
affected fruit. Ben-Arie and Sonego (1980)
reported that pectin metabolism was abnor-
mal in chilled peaches, leading to the forma-
tion of a gel when the soluble pectins were
mixed with water. The viscosity of the soluble
pectin from chilled peaches was higher than
from peaches ripened normally. The binding
of water in pectin gels may explain the reduc-
tion of extractable juice in mealy fruit. Higher
PME activity during cold storage of peach
fruit may account for this increase in viscosity
565
of soluble pectin. PME produces de-esterified
pectins of high molecular mass that incorpo-
rate water in a gel. Normally endoPG breaks
down de-esterified pectins into small frag-
ments but low-temperature storage reduces
PG activity, resulting in the accumulation of
larger-sized pectin polymers and high juice
viscosity in mealy fruits (Zhou et al., 1999).
The incidence of mealiness depends on
genetic factors and maturity stage; Fernan-
dez-Trujillo et al. (1998) found that less mature
fruit were more susceptible to mealiness.
Generally, susceptibility to mealiness is less
in early-season cultivars than for those that
mature late in the season. According to this
pattern slow-ripening genotypes are supposed
to be more susceptible to mealiness (Brecht
et al., 1984; Kader and Chordas, 1984).
Effect of controlled atmosphere and modified
atmosphere packaging
Reduction in oxygen concentration and an
increase in carbon dioxide concentration may
retard ripening of peaches and, in some cases,
delay or prevent the appearance of chilling
symptoms (Lill et al., 1989). With the decreas-
ing availability of molecular oxygen a decline
in the general metabolism of the cells occurs.
However, large changes do not take place until
oxygen falls below about 5%. The critical fac-
tor is the actual concentration of oxygen within
the cells. Internal oxygen concentration is
controlled by resistance of the flesh to oxygen
diffusion and the rate of respiration. Changes
in these parameters lead to variations in the
optimum external oxygen concentration requi-
red for various products. Large, dense prod-
ucts generally have a higher external oxygen
requirement than smaller and/or less dense
products. Due to their sensitivity to chilling
injuries some products must be stored at
higher temperatures than others, thus increas-
ing their oxygen utilization rate.
When low oxygen (1–5% O2) and/or
enhanced carbon dioxide concentrations are
applied to peaches, softening and colour
development may be retarded and the rates
of respiration and ethylene production redu-
ced (Smilanick and Fouse, 1989). Oxygen at
566 A. Ramina et al.
0.25% reduces respiration by 45%, ethylene
production rates by more than 90% (Ke et al.,
1991) and delays the increase in PG activity
(Lurie and Pesis, 1992). Carbon dioxide con-
centrations are important during storage at
low temperature. An atmosphere containing
5% CO2 may delay the appearance of physio-
logical disorders caused by low temperature
(Lill et al., 1989). However, there are no spe-
cific recommendations for the composition of
the controlled atmosphere because relatively
few cultivars give a beneficial response. Wade
(1981) found that 20% CO2 (without reduc-
tion of O2) effectively decreased the incidence
of physiological disorders in ‘J.H. Hale’
peaches during 6 weeks’ storage at 0°C. Sub-
sequent research has shown that only a few
cultivars respond to high carbon dioxide lev-
els (Uthairatanakij, 2004).
The storage of peaches under controlled
atmosphere for longer periods may cause the
development of off-flavours and excessive
accumulation of acetaldehyde and ethanol
(Kader, 1986; Smilanick and Fouse, 1989). The
development of off-flavours is a result of pro-
longed storage in atmospheres particularly
rich in carbon dioxide. Considering the effects
of low oxygen or high carbon dioxide concen-
tration on ripening physiology, short treat-
ments (a few hours to a few days) may be
beneficial to prolong the storage life. Lurie
and Pesis (1992) showed that short exposure
of fruit to anaerobic conditions (24 h) retards
the softening of peach in successive 6-day
storage without changing other quality
parameters, such as soluble solids and acid-
ity. Gaseous shocks such as ultra-low oxygen
(ULO) and 30% CO2 applied for up to 24 and
48 h at 20°C have been shown to maintain
flesh firmness during the post-treatment
phase (in air) (Bonghi et al., 1999). This effect
is accompanied by a strong inhibition of eth-
ylene evolution, particularly in those fruits
harvested at an early stage of ripening when
endogenous ethylene production is low. The
effect of these treatments on the softening
process has been related to the temporary
inhibition of EG activity. Levels of O2 below
1% or CO2 levels above 20% induce acetalde-
hyde and ethanol accumulation in several fruit
species including peaches. Prolonged anaero-
biosis results in excessive accumulation of
acetaldehyde and ethanol in peach mesocarp
(Tonutti et al., 1998). Short-term treatments
with low oxygen, besides reducing the soft-
ening rate, induce a prompt accumulation of
acetaldehyde and ethanol. However, their
concentrations in fruit tissue decline gradu-
ally when the fruit are transferred to air. Acet-
aldehyde accumulates naturally during normal
ripening, although its concentration does not
exceed that in anaerobically stored fruit. The
decrease in acetaldehyde is a result of its
metabolism to other compounds (Ke et al.,
1995) or diffusion out of the fruit tissue. At
4°C much less acetaldehyde and ethanol
accumulate than at 20°C. Fruit stored at lower
temperature are apparently less sensitive to
anaerobic conditions compared with higher
temperature. Induction of alcohol dehydroge-
nase (ADH) is regarded as one of the reasons
for anaerobic fermentation and accumulation
of ethanol (Kennedy et al., 1992). Extractable
ADH activity depends on oxygen or carbon
dioxide concentration as well as on the dura-
tion of anaerobic treatment and temperature.
Bonghi et al. (1999) detected the presence of a
multigene ADH family in ‘Springcrest’ and
showed that anaerobic metabolism is rapidly
activated following exposure to ULO and
high CO2 concentrations.
21.5 Future Perspectives and
Conclusions
Challenges for peach physiologists include
finding ways to reduce the variability in SSC
among fruit within trees and to ensure accu-
mulation of levels that are acceptable to con-
sumers. This is particularly important in
low-chill cultivars that have a short FDP.
Advances in portable near infra-red technol-
ogy that permit measurements of sugar levels
in developing fruit should greatly assist
development of tree management strategies
that will reduce within-tree variability (Gold-
ing et al., 2006). Peach growers are in the
‘fashion business’ because customers are con-
stantly seeking products with different appear-
ance and flavours. Since the generation time
is only about 3 years for peaches, an enor-
mous number of new cultivars are released
 Ripening, Nutrition and Postharvest Physiology 567

each year. However, breeders must keep upper-
most the need to maximize the yield of mar-
ketable fruit with high eating quality that will
withstand the many transfer points along the
distribution chain from orchard to consumer.
A challenge for geneticists and breeders is to
improve the cool storage life of peaches.
Although the seasonality of peaches is one of
their attractions to consumers, there is increas-
ing interest in global trading arrangements.
The short storage life of peaches restricts the
time and distance the fruit can be shipped. An
increase in cool storage life of 50% would
enable peaches to be shipped by cheaper sea
freight to many markets and at the same time
achieve in-transit non-chemical quarantine
disinfestation treatment of some insect pests.
It is an academic challenge to understand
why peaches suffer physiological disorders
when stored below about 8°C yet maximum
storage life is achieved at near freezing tem-
peratures. Findings at the molecular and bio-
chemical levels of the observation that only
some cultivars respond beneficially to atmo-
spheres containing up to 20% CO2 could open
up new insights into ways to improve cool
storage life.
Recent advances have shed light on the
molecular basis of peach fruit ripening and
postharvest physiology. Examination of some
aspects of ethylene biosynthesis and percep-
tion has demonstrated homology with the
basic model defined in Arabidopsis and tomato.
Research focused on fruit softening showed
that cell wall metabolism during peach ripen-
ing is complex and involves the sequential
action of different cell wall hydrolases and
ethylene, and is developmentally regulated.
Genetic and molecular characterization of sugar
metabolism, pigmentation, aroma, dietary and
allergenic properties of peach fruit are pro-
ceeding in several laboratories. Each one of
these physiological aspects may represent a
target for practical manipulation that aims to
improve peach fruit quality. The continuing
development of genomic tools, including
expressed sequence tags (ESTs) and cDNA
microarrays, will rapidly advance knowledge
of fruit development and ripening in peach as
well as other species. In this context, a signifi-
cant contribution has been made by the
ESTree Consortium (http://linuxbox.itb.cnr.
it/estree_mar2004/, accessed August 2007),
which developed the first oligo-based micro-
array (µPEACH 1.0), corresponding to 4806
unigenes expressed in peach fruit (ESTree
Consortium, 2005). This array has been used to
investigate transcriptome changes during the
transition from the pre-climacteric to climac-
teric phases (Trainotti et al., 2006a). Large-
scale analysis showed that 267 and 109 genes
are up- and downregulated, respectively,
during this transition. These genes have been
classified according to the TAIR gene ontol-
ogy, allowing the identification of a new
member of the ETR family in peach and a
number of transcription factors belonging to
several families, putatively involved in the
regulation of peach ripening. The number of
peach fruit ESTs is expected to increase along
with the availability of genomics tools since
several research programmes around the
world are moving in this direction, and peach
is a candidate model for genetic studies
among Rosaceae species because of the small
size of its genome (see also Chapter 4).

References

Abdi, N., Holford, P. and McGlasson, B. (2002) Application of two-dimensional gel electrophoresis to detect
proteins associated with harvest maturity in stonefruit. Postharvest Biology and Technology 26, 1–13.
Amoros, A., Serrano, M., Riquelme, F. and Romojaro, F. (1989) Levels of ACC and physical and chemical
parameters in peach development. Journal of Horticultural Science 64, 673–677.
Anderson, R.E. (1979) The influence of storage temperature and warming during storage on peaches and nec-
tarine fruit quality. Journal of the American Society for Horticultural Science 104, 459–461.
Arondel, V., Vergnolle, C., Cantrel, C. and Kader, J.C. (2000) Lipid transfer proteins are encoded by a small
multigene family in Arabidopsis thaliana. Plant Science 157, 1–12.
Artes, F., Cano, A. and Fernandez-Trujillo, J.P. (1996) Pectolytic enzyme activity during intermittent warming
storage of peaches. Journal of Food Science 61, 311–313.
568 A. Ramina et al.

Asero, R., Mistrello, G., Roncarolo, D., de Vries, S.C., Gautier, M.F., Ciurana, C.L.F., Verbeek, E., Mohammadi,
T., Knul-Brettlova, V., Akkerdaas, J.H., Bulder, I., Aalberse, R.C. and van Ree, R. (2000) Lipid transfer
protein: a pan-allergen in plant-derived foods that is highly resistant to pepsin digestion. International
Archives of Allergy and Immunology 122, 20–32.
Barry, C.S., Llop-Tous, M.I. and Grierson, D. (2000) The regulation of 1-aminocyclopropane-1-carboxylic acid
synthase gene expression during the transition from system-1 to system-2 ethylene synthesis in tomato.
Plant Physiology 123, 979–986.
Begheldo, M., Manganaris, G.A., Bonghi, C. and Tonutti, P. (2008) Different post harvest conditions modulate
ripening and ethylene biosynthetic and signal transductive pathway in stony hard peaches. Postharvest
Biology and Technology 48, 84–91.
Ben-Arie, R. and Sonego, L. (1980) Pectolytic enzyme activity involved in woolly breakdown of stored peach-
es. Phytochemistry 19, 2553–2555.
Bengoechea, M.L., Sancho, A.I., Bartolome, B., Estrella, I., Gomez-Cordoves, C. and Hernandez, M. (1997)
Phenolic composition of industrially manufactured purees and concentrates from peach and apple fruits.
Journal of Agricultural and Food Chemistry 45, 4071–4075.
Bleeker, A.B. (1999) Ethylene perception and signaling: an evolutionary perspective. Trends in Plant Science
4, 269–274.
Boller, T., Herner, R.C. and Kende, H. (1979) Assay for and enzymic formation of an ethylene precursor,
1-aminocyclopropane-1-carboxylic acid. Planta 145, 293–303.
Bonghi, C., Ferrarese, L., Ruperti, B., Tonutti, P. and Ramina, A. (1998) Endo-b-1,4-glucanases are involved in
peach fruit growth and ripening and regulated by ethylene. Physiologia Plantarum 102, 346–352.
Bonghi, C., Ramina, A., Ruperti, B., Vidrih, R. and Tonutti, P. (1999) Peach fruit ripening and quality in relation
to picking time, and hypoxic and high CO2 short-term postharvest treatments. Postharvest Biology and
Technology 16, 213–222.
Botton, A., Begheldo, M., Rasori, A., Bonghi, C. and Tonutti, P. (2002) Differential expression of two lipid transfer
protein genes in reproductive organs of peach (Prunus persica L. Batsch). Plant Science 163, 993–1000.
Botton, A., Vegro, M., De Franceschi, F., Ramina, A., Gemignani, C., Marcer, G., Pasini, G. and Tonutti, P.
(2006) Different expression of Pp-LTP1 and accumulation of Pru p 3 in fruits of two Prunus persica L.
Batsch genotypes. Plant Science 171, 106–113.
Brady, C.J. (1993) Stone fruit. In: Seymour, G.B., Taylor, J.E. and Tucker, G.A. (eds) Biochemistry of Fruit Ripen-
ing. Chapman & Hall, London, pp. 379–404.
Brecht, J.K. and Kader, A.A. (1984) Ethylene production by fruit of some slow-ripening nectarine genotypes.
Journal of the American Society for Horticultural Science 109, 763–767.
Brecht, J.K., Kader, A.A. and Ramming, D.W. (1984) Description and postharvest physiology of some slow-
ripening nectarine genotypes. Journal of the American Society for Horticultural Science 109, 596–600.
Bregoli, A.M., Scaramagli, S., Costa, G., Sabatini, E., Ziosi, V., Biondi, S. and Torrigiani, P. (2002) Peach (Pru-
nus persica) fruit ripening: aminoethoxyvinylglicine (AVG) and exogenous polyamines affect ethylene
emission and flesh firmness. Physiologia Plantarum 114, 472–481.
Brenna, O., Pompei, C., Ortolani, C., Pravettoni, V., Farioli, L. and Pastorello, E.A. (2000) Technological pro-
cesses to decrease the allergenicity of peach juice and nectar. Journal of Agricultural and Food Chemistry
48, 493–497.
Brooks, S.J., Moore, J.N. and Murphy, J.B. (1993) Quantitative and qualitative changes in sugar content of
peach genotypes (Prunus persica L. Batsch). Journal of the American Society for Horticultural Science
118, 97–100.
Brovelli, E.A., Brecht, J.K., Sherman, W.B. and Sims, C.A. (1999) Nonmelting-flesh trait in peaches is not
related to low ethylene production rates. HortScience 34, 313–315.
Callahan, A.M., Morgens, P.H., Wright, P. and Nichols, K.E. Jr (1992) Comparison of Pch313 (pTOM13
homolog) RNA accumulation during fruit softening and wounding of two phenotypically different peach
cultivars. Plant Physiology 100, 482–488.
Callahan, A.M., Fishel, D. and Dunn, L.J. (1993) Relationship of ACC [1-aminocyclopropane-1-carboxylate]
oxidase RNA, ACC [1-aminocyclopropane-1-carboxylate] synthase RNA, and ethylene in peach fruit. In:
Pech, J.C., Latché, A. and Balagué, C. (eds) Cellular and Molecular Aspects of the Plant Hormone Ethyl-
ene. Kluwer Academic Publishers, Dordrecht, The Netherlands, pp. 31–32.
Callahan, A.M., Scorza, R., Bassett, C., Nickerson, M. and Abeles, F.B. (2004) Deletions in an endopolygalac-
turonase gene correlate with non-melting flesh texture in peach. Functional Plant Biology 3, 159–168.
Chang, C., Kwok, S.F., Bleeker, A.B. and Meyerowitz, E.M. (1993) Arabidopsis ethylene responsive gene ETR1:
similarity of product to two-component regulators. Science 262, 1807–1809.
 Ripening, Nutrition and Postharvest Physiology 569

Conti, A., Fortunato, D., Ortolani, C., Giuffrida, M.G., Pravettoni, V., Napoletano, L., Farioli, L., Perono Ga-
roffo, L., Trambaioli, C. and Pastorello, E.A. (2001) Determination of the primary structure of two lipid
transfer proteins from apricot (Prunus armeniaca). Journal of Chromatography B: Biomedical Sciences
and Applications 756, 123–129.
Crisosto, C.H., Johnson, R.S., DeJong, T. and Day, K.R. (1997) Orchard factors affecting postharvest stone fruit
quality. HortScience 32, 820–822.
Crisosto, C.H., Garner, D., Andris, H.L. and Day, K.R. (2004) Controlled delayed cooling extends peach
market life. HortTechnology 14, 99–104.
Dal Cin, V., Rizzini, F.M., Botton, A. and Tonutti, P. (2006) The ethylene biosynthetic and signal transduction
pathways are differently affected by 1-MCP in apple and peach fruit. Postharvest Biology and Technology
42, 125–133.
Dawson, D.M., Melton, L.D. and Watkins, C.D. (1992) Cell wall changes in nectarines (Prunus persica). Plant
Physiology 100, 1203–1210.
De Santis, D. and Mencarelli, F. (2001) Influenza del propylene e del 1-metilciclopropene sull’aroma delle
pesche. Rivisita di Frutticoltura e di Ortofloricoltura 6, 79–80.
Dong, L., Zhou, H.W., Sonego, L., Lers, A. and Lurie, S. (2001) Ethylene involvement in the cold storage dis-
order of Flavortop nectarine. Postharvest Biology and Technology 23, 105–115.
Downs, C.G. and Brady, C.J. (1990) Two forms of exopolygalacturonase increase as peach fruits ripen. Plant,
Cell & Environment 13, 523–530.
Downs, C.G., Brady, C.J. and Gooley, A. (1992) Exopolygalacturonase protein accumulates late in peach fruit
ripening. Physiologia Plantarum 85, 133–140.
Dziedzie, S. and Hudson, B. (1983) Polyhydroxy chalcones and flavanones as antioxidants for edible oils.
Food Chemistry 12, 205–212.
Eberhardt, M.V., Lee, C.Y. and Liu, R.H. (2000) Antioxidant activity of fresh apples. Nature 405, 903–904.
Ebihara, K., Kiriyama, S. and Manabe, M. (1979) Cholesterol-lowering activity of various natural pectins and
synthetic pectin-derivatives with different physico-chemical properties. Nutrition Reports International
20, 519–526.
Eccher Zerbini, P., Gorini, F., Spada, G. and Liverani, C. (1991) Il colore di fondo come indice di raccolta delle
pesche. Rivisita di Frutticoltura e di Ortofloricoltura 6, 27–33.
Eccher Zerbini, P., Vanoli, M., Grassi, M., Rizzolo, A., Fabiani, M., Cubeddu, R., Pifferi, A., Spinelli, L. and
Torricelli, A. (2006) A model for the softening of nectarines based on sorting fruit by time-resolved reflec-
tance spectroscopy. Postharvest Biology and Technology 39, 223–232.
ESTree Consortium (2005) Development of an oligo-based microarray (µPEACH 1.0) for genomics studies in
peach fruit. Acta Horticulturae 682, 263–268.
Etienne, C., Moing, A., Dirlewanger, E., Raymond, P., Monet, R. and Rothan, C. (2002) Isolation and character-
ization 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.
Fernandez de Simon, B., Perez-Ilzarbe, J., Hernandez, T., Gomez Cordovez, C. and Estrella, I. (1990) HPLC
study of efficiency of extraction of phenolic compounds. Chromatography 30, 35–37.
Fernandez de Simon, B., Perez-Ilzarbe, J., Hernandez, T., Gomez Cordovez, C. and Estrella, I. (1992) Impor-
tance of phenolic compounds for characterization of fruit. Journal of Agricultural and Food Chemistry
40, 1531–1535.
Fernandez-Trujillo, J.P., Cano, A. and Artes, F. (1998) Physiological changes in peaches related to chilling
injury and ripening. Postharvest Biology and Technology 13, 109–119.
Genard, M. and Souty, M. (1996) Modeling the peach sugar contents in relation to fruit growth. Journal of the
American Society for Horticultural Science 121, 1122–1131.
Genard, M., Lescourret, F., Gomez, L. and Habib, R. (2003) Changes in fruit sugar concentrations in response
to assimilate supply, metabolism and dilution: a modeling approach applied to peach fruit (Prunus per-
sica). Tree Physiology 23, 373–385.
Golding, J.B., Shearer, D., McGlasson, W.B. and Wyllie, S.G. (1999) Relationships between respiration, ethylene,
and aroma production in banana. Journal of Agricultural and Food Chemistry 47, 1646–1651.
Golding, J.B., Satyan, S., McGlasson, W.B., Liebenberg, C. and Walsh, K. (2006) Application of portable NIR
for measuring soluble solids in peaches. Acta Horticulturae 713, 461–464.
Grigelmo, M.N. and Martin-Belloso, O. (1999) Comparison of dietary fibre from by-products of processing
fruit and greens and from cereals. Lebensmittel-Wissenschaft & Technologie 32, 503–508.
Grigelmo, M., Gorinstein, N. and Martin-Belloso, O. (1999) Characterisation of peach dietary fibre concen-
trate as food ingredient. Food Chemistry 65, 175–181.
570 A. Ramina et al.

Gross, J. (1987) Pigments in Fruits. Academic Press, New York.

Haji, T., Yaegaki, H. and Yamaguchi, M. (2003) Softening of stony hard peach by ethylene and the induction
of endogenous ethylene by 1-aminocyclopropane-1-carboxylic acid (ACC). Journal of the Japanese
Society for Horticultural Science 72, 212–217.
Haji, T., Yaegaki, H. and Yamaguchi, M. (2005) Inheritance and expression of fruit texture melting, no-melting
and stony hard in peach. Scientia Horticulturae 105, 241–248.
Hayama, H., Ito, A., Moriguchi, T. and Kashimura, Y. (2003) Identification of a new expansin gene closely
associated with peach fruit softening. Postharvest Biology and Technology 29, 1–10.
Hayama, H., Shimada, T., Fujii, H., Ito, A. and Kashimura, Y. (2006) Ethylene regulation of fruit softening and
softening-related genes in peach. Journal of Experimental Botany 57, 4071–4077.
Hedge, S. and Maness, O. (1998) Changes in apparent molecular mass of pectin and hemicellulose extracts
during peach softening. Journal of the American Society for Horticultural Science 123, 445–456.
Henning, H. and Herrmann, K. (1980) Flavonol glycosides of apricots (Prunus armeniaca L.) and peaches
(Prunus persica Batsch). 13, Phenolics of fruits. Zeitschrift für Lebensmittel-Untersuchung und Forschung
171, 183–188.
Hinton, D.M. and Pressey, R. (1974) Cellulase activity in peaches during ripening. Journal of Food Science 39,
783–785.
Holloway, W.D. (1983) Composition of fruit, vegetable and cereal dietary fibre. Journal of the Science of Food
and Agriculture 34, 1236–1240.
Horvat, R.J., Chapman, G.V., Robertson, J.A., Meredith, F., Scorza, R., Callahan, A.M. and Morgens, P. (1990)
Comparison of the volatile compounds from several commercial peach cultivars. Journal of Agricultural
and Food Chemistry 38, 234–237.
Hua, J., Chang, C., Sun, Q. and Mayerowitz, E.M. (1995) Ethylene insensitivity conferred by Arabidopsis ERS
gene. Science 269, 1712–1714.
Hua, J., Sakai, H., Nourizadeh, S., Chen, C., Bleecker, A.B., Ecker, J.R. and Meyerowitz, E.M. (1998) EIN4 and
ERS2 are members of the putative ethylene receptor gene family in Arabidopsis. Plant Cell 10, 1321–
1332.
Kader, A.A. (1986) Biochemical and physiological basis for effects of controlled and modified atmosphere in
fruits and vegetables. Journal of Experimental Botany 40, 99–100.
Kader, A.A. (2002) Fruits in the global market. In: Knee, M. (ed.) Fruit Quality and Its Biological Basis. CRC
Press, Boca Raton, Florida, pp.1–16.
Kader, A.A. and Barret, D. (1996) Classification, composition of fruits, and postharvest maintenance of
quality. In: Somogy, L.P. (ed.) Processing Fruits: Science and Technology, Vol. 1. Technomic Publishing
Co., Lancaster, Pennsylvania, pp. 1–24.
Kader, A.A. and Chordas, K.R. (1984) Evaluating the browning potential of peaches. California Agriculture 38,
14–15.
Kader, J.C. (1996) Lipid transfer proteins in plants. Annual Reviews of Plant Physiology and Plant Molecular
Biology 47, 627–654.
Karakurt Y., Huber, D.J. and Sherman, W.B. (2000) Quality characteristics of melting and non-melting flesh
peach genotypes. Journal of the Science of Food and Agriculture 80, 1848–1853.
Ke, D., Rodriguez, L. and Kader, A.A. (1991) Physiological responses and quality attributes of peaches kept in
low-oxygen atmospheres. Scientia Horticulturae 47, 295–303.
Ke, D., Yahia, E., Hess, B., Zhou, L. and Kader, A.A. (1995) Regulation of fermentative metabolism in avocado
fruit under oxygen and carbon dioxide stresses. Journal of the American Society for Horticultural Science
120, 481–490.
Kennedy, R.A., Rumpho, M.E. and Fox, T.C. (1992) Anaerobic metabolism in plants. Plant Physiology 100, 1–6.
Klein, B.P. (1987) Nutritional consequences of minimal processing of fruits and vegetables. Journal of Food
Quality 10, 179–183.
Kulp, K., Lorenz, K. and Stone, M. (1991) Functionality of carbohydrate ingredients in bakery products. Jour-
nal of Experimental Botany 45, 136–142.
Lessertois, D. and Moneger, R. (1978) Evolution des pigments pendant la croissance et la maturation du fruit
Prunus persica. Phytochemistry 17, 411–415.
Lester, D.R., Speirs, J., Orr, G. and Brady, C.J. (1994) Peach (Prunus persica) endopolygalacturonase cDNA
isolation and mRNA analysis in melting and nonmelting peach cultivars. Plant Physiology 105, 225–
231.
Lester, D.R., Sherman, W.B. and Atwell, B.J. (1996) Endopolygalacturonase and the melting flesh (M) locus in
peach. Journal of the American Society for Horticultural Science 121, 231–235.
 Ripening, Nutrition and Postharvest Physiology 571

Lewallen, K.S. and Marini, R.P. (2003) Relationship between flesh firmness and ground colour in peach as
influenced by light and canopy position. Journal of the American Society for Horticultural Science 128,
163–170.
Lill, R.E., O’Donoghue, E.M. and King, G.A. (1989) Postharvest physiology of peaches and nectarines. Horti-
cultural Reviews 11, 413–452.
Lleonart, R., Cistero, A., Carriera, J., Batista, A. and Moscoso del Prado, J. (1992) Food allergy: identification
of the major IgE-binding component of peach (Prunus persica). Annals of Allergy 69, 128–130.
Lurie, S. and Crisosto, C. (2005) Chilling injury in peach and nectarine. Postharvest Biology and Technology
37, 195–208.
Lurie, S. and Pesis, E. (1992) Effect of acetaldehyde and anaerobiosis as postharvest treatments on the quality
of peaches and nectarines. Postharvest Biology and Technology 1, 317–326.
McGlasson, W.B., Rath, A.C. and Legendre, L. (2005) Preharvest application of aminovinylglycine (AVG)
modifies harvest maturity and cool storage life of ‘Arctic Snow’ nectarines. Postharvest Biology and
Technology 36, 93–102.
McGuire, R.G. (1992) Reporting of objective color measurements. HortScience 27, 1254–1255.
McMurchie, E.J., McGlasson, W.B. and Eaks, I.L. (1972) Treatment of fruit with propylene gives information
about the biogenesis of ethylene. Nature 237, 235–236.
Maness, N.O., Chrz, D., Hedge, S. and Goffreda, J.C. (1993) Cell wall changes in ripening peach fruit from
cultivars differing in softening rate. Acta Horticulturae 343, 200–203.
Mathooko, F.M., Tsunashima, Y., Owino, W.Z.O., Kubo, Y. and Inaba, A. (2001) Regulation of genes encoding
ethylene biosynthetic enzymes in peach (Prunus persica L.) fruit by carbon dioxide and 1-methylcyclo-
propene. Postharvest Biology and Technology 21, 265–281.
Mencarelli, F., Garosi, F. and Tonutti, P. (1998) Postharvest physiology of peaches and nectarines slices. Acta
Horticulturae 465, 462–470.
Mencarelli, F., Cecchi, F., De Santis, D., Aquilani, R., Giuliano, G. and Rotondi, R. (2003) Influence of ethylene
on volatiles and carotenoids biosynthesis of peach. In: Vendrell, M., Klee, H., Pech, J.C. and Romojaro, F.
(eds) Biology and Biotechnology of the Plant Hormone Ethylene III. NATO Science Series 348. IOS Press,
Amsterdam, pp. 222–226.
Meredith, F.I., Robertson, J.A. and Horvath, R.J. (1989) Changes in physical and chemical parameters associated
with quality and post harvest ripening of harvested peaches. Journal of Agricultural and Food Chemistry 37,
1210–1214.
Miller, A.N., Krizek, B.A. and Walsh, C.S. (1988) Whole-fruit ethylene evolution and ACC content of peach
pericarp and seeds during development. Journal of the American Society for Horticultural Science 113,
119–124.
Moing, A., Svanella, L., Rolin, D., Gaudillere, M., Gaudillere, J.P. and Monet, R. (1998) Compositional chang-
es during the fruit development of two peach cultivars differing in juice acidity. Journal of the American
Society for Horticultural Science 123, 770–775.
Moing, A., Svanella, L., Gaudillere, M., Gaudillere, J.P. and Monet, R. (1999) Organic acid concentration is
little controlled by phosphoenolpryruvate carboxylase activity in peach fruit. Australian Journal of Plant
Physiology 26, 579–585.
Moing, A., Rothan, C., Svanella, L., Just, D., Diakou, P., Raymond, P., Gaudillere, J.P. and Monet, R. (2000) Role
of phosphoenolpyruvate carboxylase in organic acid accumulation during peach fruit development. Phys-
iologia Plantarum 108, 1–10.
Monet, R. (1979) Transmission génétique du caractère ‘fruit doux’ chez le pêcher. Incidence sur la sélection
pour la qualité. In: Proceedings of the EUCARPIA ‘Tree Fruit Breeding’ Symposium. INRA, Angers,
France, pp. 273–276.
Monnet, F.P., Dieryck, W., Boutron, F., Jourdier, P. and Gautier, M.F. (2001) Purification, characterisation
and cDNA cloning of a type 2 (7 kDa) lipid transfer protein from Triticum durum. Plant Science 16,
747–755.
Moon, H.S. and Callahan, A.M. (2004) Developmental regulation of peach ACC oxidase promoter–GUS fusion
in transgenic tomato fruits. Journal of Experimental Botany 55, 1519–1528.
Morgutti, S., Negrini, N., Nocito, F.F., Ghiani, A., Bassi, D. and Cocucci, M. (2006) Changes in endopolygalac-
turonase levels and characterization of putative endo-PG gene during fruit softening in peach genotypes
with nomelting and melting flesh fruit phenotypes. New Phytologist 171, 315–328.
Nakatsuka, A., Murachi, S., Okunishi, H., Shiomi, S., Nakano, R., Kubo, Y. and Inaba, A. (1998) Differential ex-
pression and internal feedback regulation of 1-aminocyclopropane-1-carboxylate synthase, and ethylene
receptor genes in tomato fruit during development and ripening. Plant Physiology 118, 1295–1305.
572 A. Ramina et al.

Neri, F. and Brigati, S. (1994) Sensory and objective evaluation of peaches. In: De Jager, A., Jhonson, A. and
Hohn, E. (eds) COST 94: The postharvest treatment of fruit and vegetables. European Commission, Brussels,
pp. 107–115.
Nissen, R.J., George, A.P. and Topp, B.L. (2005a) Producing super sweet and firm peaches and nectarines.
Acta Horticulturae 694, 311–314.
Nissen, R.J., George, A.P., Waite, G., Lloyd, A. and Hamacek, E. (2005b) Innovative new production systems
for low-chill stonefruit in Australia and South-East Asia: a review. Acta Horticulturae 694, 247–251.
Nonis, A., Ruperti, B., Casatta, E. and Vizzotto, G. (2003) Expression of sugar metabolism-related genes during
peach fruit and leaf development. In: Proceedings of ‘Phloem 2003’ Symposium. University of Bayreuth,
Bayreuth, Germany, p. 214.
Nonis, A., Ruperti, B., Falchi, R., Casatta, E., Enferadi, S.T. and Vizzotto, G. (2007) Differential expression and
regulation of a natural invertase encoding gene from peach (Prunus persica): evidence for a role in fruit
development. Physiologia Plantarum 129, 436–446.
Orr, G. and Brady, C. (1993) Relationship of endopolygalacturonase activity to fruit softening in a freestone
peach. Postharvest Biology and Technology 3, 121–130.
Ortolani, C., Ispano, M., Pastorello, E.A., Bigi, A. and Ansaloni, R. (1988) The oral allergy syndrome. Annals
of Allergy 61, 47–52.
Pastorello, E.A., Farioli, L., Pravettoni, V., Ortolani, C., Ispano, M., Monza, M., Broglio, C., Scibola, E., Ansa-
loni, R., Incorvaia, C. and Conti, A. (1999) The major allergen of peach (Prunus persica) is a lipid transfer
protein. Journal of Allergy and Clinical Immunology 103, 520–526.
Pastorello, E.A., D’Ambrosio, F., Pravettoni, V., Farioli, L., Giuffrida, G., Monza, M., Ansaloni, R., Fortunato, D.,
Scibola, E., Rivolta, F., Incorvaia, C., Bengtsson, A., Conti, A. and Ortolani, C. (2000) Evidence for a lipid
transfer protein as the major allergen of apricot. Journal of Allergy and Clinical Immunology 105, 371–377.
Pastorello, E.A., Farioli, L., Pravettoni, V., Giuffrida, M.G., Ortolani, C., Fortunato, D., Trambaioli, C., Scibola, E.,
Calamari, A.M., Robino, A.M. and Conti, A. (2001) Characterization of the major allergen of plum as a
lipid transfer protein. Journal of Chromatography B: Biomedical Sciences and Applications 756, 95–103.
Peace, C.P., Crisosto, C.H. and Gradziel, T.M. (2005) Endopolygalacturonase: a candidate gene for freestone
and melting flesh in peach. Molecular Breeding 16, 21–31.
Pelese-Siebenbourg, J., Caellas, C., Kader, J.K., Delseny, M. and Puigdomenech, P. (1994) A pair of genes coding
for lipid-transfer protein in Sorghum vulgare. Gene 148, 305–308.
Pressey, R. and Avants, J.K. (1973) Two forms of polygalacturonase in tomatoes. Biochimica et Biophysica
Acta 309, 363–369.
Pressey, R. and Avants, J.K. (1978) Difference in polygalacturonase composition of clingstone and freestone
peaches. Journal of Food Science 43, 1415–1417.
Pressey, R., Hinton, D.M. and Avants, K. (1971) Polygalacturonase activity and solubilization of pectin in
peaches during ripening. Journal of Food Science 36, 1070–1073.
Pyee, J. and Kolattukudy, P.E. (1995) The gene for the major cuticular wax associated protein and three homolo-
gous genes from broccoli (Brassica oleracea) and their expression patterns. The Plant Journal 7, 49–59.
Rasori, A., Ruperti, B., Bonghi, C., Tonutti, P. and Ramina, A. (2002) Characterization of two putative ethylene
receptor genes expressed during peach fruit development and abscission. Journal of Experimental Bota-
ny 53, 2333–2339.
Rasori, A., Bertolasi, B., Furini, A., Bonghi, C., Tonutti, P. and Ramina, A. (2003) Functional analysis of peach
ACC oxidase promoters in transgenic tomato and in ripening peach fruit. Plant Science 165, 523–530.
Rath, A.C. and Prentice, A.J. (2004) Application of Retain plant growth regulator increases the yield and fresh
firmness of ‘Arctic Snow’ nectarines both at harvest in Australia and after export to Taiwan. Australian
Journal of Experimental Agriculture 44, 343–351.
Robards, K., Prenzler, P., Tucker, G., Swatsitang, P. and Glover, W. (1999) Phenolic compounds and their role
in oxidative processes in fruits. Food Chemistry 66, 401–436.
Robertson, J.A., Meredith, F.I. and Scorza, R. (1988) Characteristics of fruit from high and low quality peach
cultivars. HortScience 23, 1032–1034.
Ruperti, B., Bonghi, C., Rasori, A., Ramina, A. and Tonutti, P. (2001) Characterization and expression of two
members of the peach 1-aminocyclopropane-1-carboxylate oxidase gene family. Physiologia Plantarum
111, 336–344.
Salunkhe, D.K., Bolin, H.R. and Reddy, N.R. (1991) Storage, Processing and Nutritional Quality of Fruits and
Vegetables, 2nd edn, Vol. 1. CRC Press, Boca Raton, Florida.
Sanchez-Monge, R., Lombardero, M., Garcia-Selles, F.J., Barber, D. and Salcedo, G. (1999) Lipid-transfer pro-
teins are relevant allergens in fruit allergy. Journal of Allergy and Clinical Immunology 103, 514–519.
 Ripening, Nutrition and Postharvest Physiology 573

Sharaftedinov, K.K., Mescheryakova, V.A., Plotnikova, O.A., Nikolskaya, G.V. and Pavlova, Y.V. (1999) The
influence of drinks with fructose on glycemic parameters of patient with type II diabetes. Voprosy
Pitaniya 1, 42–45.
Smilanick, J.L. and Fouse, D.C. (1989) Quality of nectarines in insecticidal low O2 atmospheres at 5°C. Journal
of the American Society for Horticultural Science 114, 431–436.
Southon, S. (2000) Increased fruit and vegetable consumption within the EU. Potential health benefits. Food
Research International 33, 211–217.
Souty, M., Reich, M., Albagnac, G. and Génard, M. (1998) Quality of peach fruit in relation to carbon supply.
Acta Horticulturae 465, 481–490.
Takeoka, G.R., Flath, R.A., Gunter, M. and Jennings, W. (1988) Nectarine volatiles: vacuum steam distillation
versus headspace sampling. Journal of Agricultural and Food Chemistry 36, 553–560.
Tatsuki, M., Takashi, H. and Yamaguchi, M. (2006) The involvement of 1-aminocyclopropane-1-carboxylic acid
synthase isogene, Pp-ACS1, in peach fruit softening. Journal of Experimental Botany 57, 1281–1289.
Thompson, J.E., Legge, R.J. and Barber, R.F. (1987) The role of free radicals in senescence and wounding. New
Phytology 105, 317–324.
Tijskens, T.M.M., Eccher-Zerbini, P., Schouten, R.E., Vanoli, M., Jacob, S., Grassi, M., Cubbedu, R., Spinelli, L.
and Torricelli, A. (2007) Assessing harvest maturity in nectarines. Postharvest Biology and Technology
45, 204–213.
Tonutti, P., Casson, P. and Ramina, A. (1991) Ethylene biosynthesis during peach fruit development. Journal of
the American Society for Horticultural Science 116, 274–279.
Tonutti, P., Bonghi, C., Ruperti, B., Tornielli, G.B. and Ramina, A. (1997) Ethylene evolution and 1-aminocy-
clopropane-1-carboxylate oxidase gene expression during early development and ripening of peach
fruit. Journal of the American Society for Horticultural Science 122, 642–647.
Tonutti, P., Bonghi, C., Ramina, A. and Vidrih, R. (1998) Molecular and biochemical effects of anoxia,
hypoxia and CO2-enriched atmosphere on Springcrest peaches. Acta Horticulturae 465, 439–446.
Torija Isasa, M.E., Carballido, A. and Barragan Ruiz, M.R. (1985) Bromatological study of baby foods. 6. Contents
of crude and dietary fibre. Anales de Bromatologia 36, 137–149.
Trainotti, L., Spolaore, S., Ferrarese, L. and Casadoro, G. (1997) Characterization of ppEG1, a member of a mul-
tigene family which encodes endo-b,1,4-glucanase in peach. Plant Molecular Biology 34, 791–802.
Trainotti, L., Zanin, D. and Casadoro, G. (2003) A cell wall-oriented genomic approach reveals a new and
unexpected complexity of the softening in peaches. Journal of Experimental Botany 54, 1821–1832.
Trainotti, L., Bonghi, C., Ziliotto, F., Zanin, D., Rasori, A., Csadoro, G., Ramina, A. and Tonutti, P. (2006a) The
use of microarray µPEACH 1.0 to investigate transcriptome changes during transition from pre-climacteric
to climacteric phase in peach fruit. Plant Science 170, 606–613.
Trainotti, L., Pavanello, A. and Zanin, B. (2006b) PpEG4 is a peach endo-b-1,4-glucanase gene whose expression
in climacteric peaches does not follow a climacteric pattern. Journal of Experimental Botany 57, 589–598.
Uthairatanakij, A. (2004) Responses of nectarines to atmospheres containing high carbon dioxide concentra-
tions. PhD thesis, The University of Western Sydney, New South Wales, Australia.
Uthairatanakij, A., Penchaiya, P., McGlasson, B. and Holford, P. (2005) Changes in ACC and conjugated ACC
following CA storage of nectarines. Australian Journal of Experimental Agriculture 45, 1635–1641.
Ventura, M., Magnanimi, E. and Sansavini, S. (1998) Sistema automatico di misurazione dei gas nella mat-
urazione dei frutti. Rivista di Frutticoltura e di Ortofloricoltura 60, 63–67.
Visai, C., Vanoli, M. and Rizzolo, A. (1993) Caratteristiche aromatiche durante l’accrescimento e la maturazi-
one di frutti di pesco. In: Proceedings of ‘XXI Convegno peschicolo’. Camera di Commercio di Ravenna
e Forlì, Ravenna, Italy, pp. 295–304.
Vizzotto, G., Pinton, R., Varanini, Z. and Costa, G. (1996) Sucrose accumulation in developing peach fruit.
Physiologia Plantarum 96, 225–230.
Vizzotto, G., Casatta, E., Bomben, C., Bregoli, A.M., Sabatini, E. and Costa, G. (2002) Peach ripening as
affected by AVG. Acta Horticulturae 592, 561–566.
Wade, N.L. (1981) Effects of storage atmosphere and calcium on low-temperature injury of peach fruit.
Scientia Horticulturae 15, 145–154.
Wills, R., McGlasson, B., Graham, D. and Joyce, D. (2007) Physiology and biochemistry. In: Wills, R.,
McGlasson, B., Graham, D. and Joyce, D. (eds) Postharvest: An Introduction to the Physiology and
Handling of Fruit, Vegetables and Ornamentals, 5th edn. CABI, Wallingford, UK, p. 202.
Yamada, K., Niwa, N., Shiratake, K. and Yamaki, S. (2001) cDNA cloning of NAD-dependent sorbitol dehy-
drogenase from peach fruit and its expression during fruit development. Journal of Horticultural Science
and Biotechnology 76, 581–587.
574 A. Ramina et al.

Yoshida, M. (1970) Genetical studies on the fruit quality of peach varieties. 1. Acidity. Bulletin of the Fruit Tree
Research Station, Series A 9, 1–15.
Young, E.G., Ballard, R.E. and Coston, D.C. (1989) The identification and comparison of flavonoid com-
pounds in the fruit, skins and leaves of peach cultivars. Acta Horticulturae 254, 35–40.
Zhou, H.W., Sonego, L., Ben-Arie, R. and Lurie, S. (1999) Analysis of cell wall components in juice of Flavor-
top nectarines during normal ripening and woolliness development. Journal of the American Society for
Horticultural Science 124, 424–429.
Zhou, H.W., Dong, L., Ben Arie, R. and Lurie, S. (2001) The role of ethylene in the prevention of chilling
injury in nectarines. Journal of Plant Physiology 158, 55–61.
 22 Harvesting and Postharvest Handling of
Peaches for the Fresh Market

C.H. Crisosto1 and D. Valero2

1University of California, Davis, Department of Plant Sciences; located at Kearney
Agricultural Center, Parlier, California, USA
2University Miguel Hernández, Department of Food Technology, Alicante, Spain

22.1 Origin 576

22.2 Fruit Consumption and Antioxidant Capacity 576
Fruit composition 576
Ascorbic acid, carotenoids and phenolic composition 576
Antioxidant capacity 577
22.3 Deterioration Problems 577
Internal breakdown 577
Mechanical injury 579
Inking 580
22.4 Peach Maturity 580
Maturity and quality 580
Maturity definition 581
Maturity indices 581
Field application of maturity indices 581
22.5 Temperature Requirements and Management 582
Ideal storage conditions 582
Temperature management 582
Water loss control 584
New temperature management approach 584
22.6 Field Harvesting, Hauling and Packaging 585
Harvesting 585
Fruit hauling 586
Fruit packaging 587
Sorting and sizing operation 588
Shipping and transportation 590
22.7 Cull Utilization 590
Potential uses 590
Situation in California 591
Other uses 592
22.8 Fruit Handling at Retail Distribution 592
Fruit preparation for consumers 592

© CAB International 2008. The Peach: Botany, Production and Uses

(eds D.R. Layne and D. Bassi) 575
576 C.H. Crisosto and D. Valero
Fruit buyer handling
Peach handling at retail stores
22.1 Origin
Peach (Prunus persica (L.) Batsch) is native to
China; at one time it was called ‘Persian apple’.
Chinese literature dates its cultivation in China
to 1000 BC. Probably carried from China to
Persia (Iran), the peach quickly spread from
there to Europe. In the 16th century, peaches
were established in Mexico, probably by the
Spaniards. In the 18th century Spanish mis-
sionaries introduced the peach to California,
which turned out to be the most important
production area after China and Italy (LaRue,
1989). In recent years, an important develop-
ment of fruit with high soluble solids concen-
tration (SSC), high aromatic white flesh, and
low-acidity white and yellow cultivars has
occurred in all the production areas (Okie,
1998; Sansavini et al., 2000; Crisosto et al.,
2001a; Crisosto, 2002).
22.2 Fruit Composition and
Antioxidant Capacity
Fruit composition
Peaches are characteristically soft-fleshed and
highly perishable fruit, with a limited market
life potential. A peach fruit is approximately
87% water with 180 kJ (43 kcal) and contains
carbohydrates, organic acids, pigments, phe-
nolics, vitamins, volatiles, antioxidants and
trace amounts of proteins and lipids, which
make it very attractive to consumers (Kader
and Mitchell, 1989b; USDA, 2003). Immature
peach fruit contain very low or no starch grains
and these starch grains are rapidly converted
into soluble sugars as the fruit mature and
ripen. Consequently, there is no significant
increase in soluble sugars during storage and
ripening (Romani and Jennings, 1971). Solu-
ble sugars contribute approximately 7–18% of
total weight and fibre contributes approxi-
mately 0.3% of fresh weight (FW) of total fruit.
Sucrose, glucose and fructose represent about
75% of peach fruit soluble sugars. Malic acid
592
593
is the predominant organic acid in mature
peach fruit followed by citric acid. These
organic acids (0.4–1.2% FW) are important
because it has been reported that the ratio of
soluble solids to titratable acidity determines
consumer perception in most ripe peach culti-
vars. In most peach cultivars, we found that
acidity decreased about 30% during ripening.
Peach fruit has low protein content (0.5 to
0.8% FW) but these small-size proteins have
an important function as enzymes catalysing
the various chemical reactions responsible for
compositional changes. Despite lipids consti-
tuting only 0.1 to 0.2% FW, they are important
because they make up surface wax that con-
tributes to fruit cosmetic appearance and
cuticle that protects fruit against water loss
and pathogens. Lipids are also important
constituents of cell membranes, which influ-
ence physiological activities of fruits. Miner-
als in fruits include base-forming elements
(Ca, Mg, K, Na) and acid-forming elements
(P, Cl, S). Ca associated with cell wall struc-
ture is important in fruit softening and Ca in
the apoplast has been related to senescence.
Postharvest changes in mineral content in
fruits are small. Volatile compounds in very
low concentrations include esters, alcohols,
aldehydes, ketones and acids, and these are
responsible for the characteristic fruit aroma.
Lactones may be organoleptically important
in peach flavour but more detailed studies are
needed on this topic.
Ascorbic acid, carotenoids and phenolic
composition
Peach fruit has ascorbic acid (vitamin C),
carotenoids (provitamin A) and phenolic
compounds which are good sources of anti-
oxidants (Tomás-Barberán et al., 2001; Byrne,
2002). Since these compounds are located in
a high concentration in the fruit peel, which
constitutes only about 15% of total fruit FW,
most of the antioxidant potential is restricted
to the peel; thus, it is recommended to eat
 Harvesting and Postharvest Handling
peaches with the peel to ensure intake of most
of the antioxidant compounds. The total
ascorbic acid (vitamin C) content in a survey
of ten cultivars of California peach ranged
from 6 to 9 mg/100 g in white flesh and from
4 to 13 mg/100 g in yellow flesh (Gil et al.,
2002). Accordingly, similar concentrations of
ascorbic acid (5–6 mg/100 g) have been found
in European peach cultivars (Carbonaro et al.,
2002; Proteggente et al., 2002). Total carote-
noids concentration was in the range of
71–210 mg/100 g FW for yellow-fleshed and
7–20 mg/100 g FW for white-fleshed peach
cultivars (Gil et al., 2002). Thus, there were
about ten times more carotenoids in yellow-
fleshed than in white-fleshed peach cultivars.
The main carotenoid detected was b-carotene
(provitamin A), but also small quantities of
a-carotene and b-cryptoxanthin are present in
some peach cultivars.
The total phenolics concentration
expressed as mg/100 g FW varied from 28 to
111 for white-fleshed and from 21 to 61 for
yellow-fleshed California cultivars (Gil et al.,
2002). Other European cultivars had values of
38 mg/100 g (Proteggente et al., 2002), while
the Spanish cultivar ‘Caterina’ showed values
of 240 and 470 mg/100 g for pulp and peel,
respectively (Goristein, et al., 2002). Fruit phe-
nolics have a role in fruit visual appearance
(colour), taste (astringency) and health antioxi-
dant property (Tomás-Barberán et al., 2001).
The predominant hydrocinnamic acid is chlo-
rogenic acid. Catechin and epicatechin are the
main procyanidins identified and their con-
centrations are higher in white-fleshed than
in yellow-fleshed peaches. Cyanidin-3-glucoside
is the predominant anthocyanin, which, along
with other anthocyanins, is present mainly in
the skin. Concentrations of flavonols (includ-
ing quercetin and kaempferol) are higher in
yellow-fleshed than in white-fleshed peaches
(Gil et al., 2002).
Antioxidant capacity
The antioxidant capacity per peach fruit serv-
ing based on the intake of a fruit serving of
100 g (peel + flesh) varied widely according to
cultivar. In general, white-fleshed peaches
were slightly higher in total antioxidant
577
capacities than yellow-fleshed peaches. The
total antioxidants ranged from 13 to 107.3 mg
of ascorbic acid equivalents when evaluated
by the DPPH (2,2-diphenyl-1-picrylhydrazyl)
free radical method and from 19 to 119.6 mg
of ascorbic acid equivalents when evaluated
by the FRAP (ferric reducing ability plasma)
method (Tomás-Barberán et al., 2001). When
these values are compared with the amount
of ascorbic acid equivalents provided by 100
ml of red wine, 100 g of ‘Snow Skin’ (white
flesh) or ‘September Sun’ (yellow flesh) will
provide the same amount, while approxi-
mately 1000 g of ‘Summer Sweet’ (white flesh)
or ‘Flavorcrest’ (yellow flesh) would have to
be consumed to match the same amount of
antioxidant capacity in 100 ml of red wine. In
fact, the total antioxidant activity of peach is
similar to that reported for pear, apple and
tomato; and significantly lower than those
observed in strawberry, raspberry and red
plum (Proteggente et al., 2002).
22.3 Deterioration Problems
Commercial postharvest losses are mainly
due to decay and internal breakdown (IB) or
chilling injury (CI) (Ceponis et al., 1987; Mitch-
ell and Kader, 1989a). Postharvest loss of
stone fruits to decay-causing fungi is consid-
ered the greatest deterioration problem.
Worldwide, the most important pathogen of
fresh stone fruits is grey mould or Botrytis rot,
caused by the fungus Botrytis cinerea. In Cali-
fornia, an even greater cause of loss due to
decay is caused by the fungus Monilinia fruc-
ticola (brown rot). Details on the fungi life
cycle, epidemiology, orchard sanitation prac-
tices, fungicide applications and pre-/post-
harvest management to reduce these problems
are presented in Chapter 15 of this book.
Internal breakdown
This phenomenon (IB or CI) is genetically con-
trolled and triggered by storage temperature.
It manifests itself as dry, mealy, woolly or hard-
textured fruit (not juicy), flesh or pit cavity
browning, and flesh translucency usually
578 C.H. Crisosto and D. Valero
radiating through the flesh from the pit (Fig.
22.1/Plate 232). An intense red colour devel-
opment of the flesh (‘bleeding’) usually radi-
ating from the pit may be associated with this
problem in some peach cultivars. Recently
released cultivars rich in skin red pigment
showed flesh bleeding that is not affecting
fruit taste. The development of this symptom
has been associated with fruit maturity rather
than storage temperature. In all of the cases,
in susceptible cultivars flavour is lost before
visual CI symptoms are evident (Crisosto and
Labavitch, 2002). There is large variability in
IB susceptibility among peach cultivars (Mitch-
ell and Kader, 1989a; Crisosto et al., 1999c). In
general, most of the mid-season and late-season
peach cultivars are more susceptible to CI
than early-season cultivars (Mitchell and Kader,
1989a), although as new cultivars are being
released from a new genetic pool, the suscep-
tibility to CI is becoming random in the new
Fig. 22.1. Internal breakdown symptoms in peaches (top of image) include flesh mealiness, flesh
browning and loss of flavour.
cultivar population (Crisosto et al., 1999c;
Crisosto, 2002). It has been widely reported
that the expression of CI symptoms develops
faster and more intensely when susceptible
fruit are stored at temperatures between about
2.2°C and 7.6°C (‘killing zone’ temperature)
than when stored at 0°C or below but above
their freezing point (Harding and Haller,
1934; Smith, 1934; Mitchell and Kader, 1989a).
Therefore, market life is dramatically reduced
when fruit are exposed to the ‘killing zone’
temperature (Crisosto et al., 1999c). In addition,
the severity of CI depends on the ripening stage
at harvest since higher incidence was reported
for ‘Maycrest’ cultivar picked at more advanced
ripening stage (Valero et al., 1997), although
the opposite has been found in other cultivars
(Von Mollendorff et al., 1992).
Several treatments to delay and limit devel-
opment of this disorder have been tested, such
as controlled atmosphere (CA) environment,
 Harvesting and Postharvest Handling
calcium applications, warming cold storage
interruptions (Anderson, 1979; Nanos and
Mitchell, 1991; Garner et al., 2001), plant
growth regulators and controlled delayed
cooling. The major benefits of CA during stor-
age/shipment are retention of fruit firmness
and ground colour. CA conditions of 6%
O2+17% CO2 at 0°C have shown a limited
benefit for reduction of IB during shipments
for yellow-fleshed cultivars (Crisosto et al.,
1999a) and white-fleshed cultivars (Garner
et al., 2001). The CA efficacy is related to culti-
var (Mitchell and Kader, 1989a), preharvest
factors (Von Mollendorff, 1987; Crisosto et al.,
1997), temperature, fruit size (Crisosto et al.,
1999a), marketing period and shipping time
(Crisosto et al., 1999c). Another tool that
reduced CI in peach was modified atmosphere
packaging (MAP). Thus, ‘Paraguayo’ cultivar
(flat type) showed reductions in CI severity
using polypropylene standard film with
steady-state atmosphere of 12% CO2 and 4%
O2, or oriented polypropylene (23% CO2 and
2% O2) (Fernández-Trujillo et al., 1998). The
preconditioning treatment (Crisosto et al.,
2004) prior to storage/shipment has shown to
be effective in delaying IB symptoms and is
successfully being used commercially on Cal-
ifornian and Chilean fruit shipped to the USA
and England (Crisosto et al., 2004). The ‘Para-
guayo’ cultivar subjected to intermittent warm-
ing cycles of 1 day at 20°C every 6 days of
storage at 2°C was also effective in reducing
CI symptoms although scald and translucency
occurred (Fernández-Trujillo and Artés, 1998).
Mechanical injury
Peaches are susceptible to mechanical injuries
including impact, compression, abrasion (or
vibration), bruising, and wounds or cuts,
which can occur during harvest and transport
(Mitchell and Kader, 1989a). Impact bruising
is the result of dropping, bouncing or jarring.
Compression bruising occurs primarily when
bins are overfilled and stacked, and fruits are
‘crushed’ against each other. Abrasion bruis-
ing results from fruit rubbing against each
other or against container surfaces. Proper
fruit handling and transport will reduce these
579
types of injury and contribute to the produc-
tion of a high-quality final product. Careful
handling during harvesting, hauling and
packing operations to minimize such injuries
is important because the injuries result in
reduced appearance quality, accelerated
physiological activity, potentially more entry
points for and inoculation by fruit decay
organisms, and greater water loss. Incidence
of impact and compression bruising has
become a greater concern as a large part of the
peach industry is harvesting fruit at more
advanced maturity (softer) to maximize fruit
flavour quality. Our observations indicate
that most impact bruising damage occurs
during long hauling from orchard to packing-
house and during the packinghouse opera-
tion. Critical impact bruising thresholds (the
minimum fruit firmness measured at the
weakest point to tolerate impact abuse) have
been developed for many peach and nectar-
ine cultivars (Crisosto et al., 2001b). Physical
wounding or cuts on peaches can occur at
any time from harvest until consumption.
Good worker supervision assures adequate
protection against impact bruising during
picking, handling and transport of fruit.
Abrasion damage can occur at any time
during postharvest handling. Protection against
abrasion damage involves procedures to reduce
vibrations during transport and handling by
immobilizing the fruit. These procedures
include: installing air-suspension systems on
axles of field and highway trucks, plastic film
liners inside field bins, the use of plastic bins,
installing special bin top pads before trans-
port, avoiding abrasion on the packing line,
and using packing procedures that immobilize
the fruit within the shipping container before
they are transported to market. It is also help-
ful to grade farm roads to reduce roughness,
avoid rough roads during transport, and
establish strict speed limits for trucks operat-
ing between orchards and packinghouses.
Some research indicates that treatment of
peaches with plant growth regulators (i.e.
polyamines and gibberellic acid) before han-
dling and storage is also effective in reducing
the fruit susceptibility to mechanical damage
by increasing fruit firmness and thus inducing
resistance to compression forces (Martínez-
Romero et al., 2000). Additionally, preharvest
580 C.H. Crisosto and D. Valero

application of Ca+Mg+Ti spray led to firmer
fruits (Serrano et al., 2004), which would be
another way to increase the fruit resistance to
mechanical damage.
Inking
In situations when abrasion damage occurs
during harvesting on fruit that have heavy
metal contaminants on their skin (i.e. Fe, Cu
and/or Al), a dark discoloration referred to as
inking, staining or peach skin discoloration
occurs on the skin (Cheng and Crisosto, 1997).
These dark or brown spots or stripes on the
fruit surface are a cosmetic problem that is
limited to the skin but they lead to market
rejection and financial loss to the grower (Fig.
22.2/Plate 233). Light brown spots or stripes
are also produced on the surface of white-
fleshed peaches and nectarines as a conse-
quence of abrasion occurring mainly during
harvesting and hauling operations. These
symptoms appear generally 24–48 h after har-
vest. This problem is usually triggered during
the harvesting and hauling operations, but it
may also occur later during postharvest han-
dling (packaging). Heavy metal contaminants
on the surface of the fruit can occur as a conse-
quence of foliar nutrients and/or fungicides
sprayed within 15 or 7 days before harvest,
respectively. Gentle fruit handling, short-
distance hauling, avoiding any foliar nutrient
sprays within 15 days of harvest, and following
the suggested preharvest fungicide spray
interval guidelines are our recommendations to
reduce inking incidence (Crisosto et al., 1999b).
22.4 Peach Maturity
Maturity and quality
The maturity at which peaches are harvested
greatly influences their ultimate flavour, mar-
ket life and quality potential (Von Mollen-
dorff, 1987; Lill et al., 1989; Kader and Mitchell,
1989a; Crisosto et al., 1995). Peach maturity con-
trols the fruit’s flavour components, physio-
logical deterioration problems, susceptibility

Fig. 22.2. Peach inking or staining as a consequence of abrasion combined with heavy metal
contamination during harvesting and hauling operations.
 Harvesting and Postharvest Handling
to mechanical injuries, resistance to moisture
loss, susceptibility to invasion by rot organ-
isms, market life and ability to ripen (Shew-
felt et al., 1987; Crisosto, 1994). Peaches that
are harvested too soon (immature) may fail to
ripen properly or may ripen abnormally.
Immature fruit typically soften slowly and
irregularly, never reaching the desired melt-
ing texture of fully matured fruit. Green
ground colour of fruit picked immature may
never fully disappear. Because immature fruit
lack a fully developed surface cuticle, they
are more susceptible to water loss than prop-
erly matured fruit. Immature and low-maturity
fruit have lower SSC and higher acids than
harvested matured fruit, all of which contrib-
ute to inadequate flavour development and
low consumer acceptance. Over-mature fruit
have a shortened postharvest life, primarily
because of rapid softening and they are
already approaching a senescent stage at har-
vest. Such fruit have partially ripened, and
the resulting flesh softening renders them
highly susceptible to mechanical injury and
fungus invasion. By the time such fruit reach
the consumer they may have become over-
ripe (senescent), with poor eating quality
including off-flavours and irregular or mushy
texture.
Maturity definition
Optimum maturity must be defined for each
peach cultivar to assure maximum taste and
storage quality but in all cases it should assure
that the fruit has the ability to ripen satisfac-
torily (Kader and Mitchell, 1989a). Peach
quality was discussed in detail in Chapter 20
of this book. The ideal maturity varies accord-
ing to markets; for example, a more advanced
maturity is recommended for near-distance
markets than for long-distance markets.
Maturity indices
Several information sources from different pro-
duction areas have reported that flesh colour,
firmness and background colour changes are
well correlated to chemical and physical fruit
changes during maturation and ripening
581
(Crisosto, 1994). Based on this information,
maturity indices based on skin background
colour and firmness are being used to deter-
mine and supervise harvesting operations. In
California and other places, harvest date is
determined by skin background colour changes
from green to yellow in most cultivars. A
colour chip guide is used to determine matu-
rity of most cultivars except for white-fleshed
cultivars. Fruit skin background colour is a
useful, non-destructive method of estimating
fruit maturity, and is most easily employed
and understood by field workers during har-
vesting operations. Since the proper back-
ground colour for estimating optimum harvest
maturity varies by cultivar, experience with a
particular cultivar is helpful in making the
correct decision. In California, for new culti-
var releases where skin ground colour is
masked by full red colour development prior
to maturation, fruit firmness is being used to
determine how long fruit can be left on the
tree before harvest. In Europe, fruit firmness
on fresh market peach is not very reliable, so
maximum maturity index is recommended.
Maximum maturity is defined as the mini-
mum flesh firmness at which fruits can be
handled without bruising damage. Maximum
maturity varies among peach cultivars and
handling situations (Crisosto et al., 2001b).
Field application of maturity indices
In applying either one of these two maturity
indices (background colour and flesh firm-
ness) at the start of a block or cultivar, proper,
easily understood directions for estimating
maturity should be given to the workers. By
selecting a few fruit of varying maturity and
demonstrating what maturity level is accept-
able and unacceptable, many mistakes can be
avoided. It is recommended to leave these
samples with the crew leader as a reference
throughout the day. When a maturity index
based on fruit firmness is used, the instruc-
tions to the harvesters will also imply mini-
mum size and location of the fruit in the tree
canopy. The value of a good and expert crew
leader cannot be overemphasized. This person
should be considered essential and integral in
the harvesting process. He should be instructed
582 C.H. Crisosto and D. Valero
to continually monitor the fruit being picked
and the fruit remaining on the tree to deter-
mine if the correct balance is achieved.
Orchard managers should involve the crew
leader in all stages of the decision-making
process when determining optimum harvest
maturity. Doing so will give him greater
understanding and experience in the process.
More importantly, it will solidify in his mind
the importance of his role in harvesting fruit
at the proper maturity.
A number of factors can affect how
quickly fruits ripen. Trees tend to ripen from
top to bottom and periphery to interior. This is
probably related to the amount of sunlight
they receive. Consequently, fruit of a given
cultivar on weak trees tend to ripen earlier
than on strong trees, as do fruit on summer-
pruned trees. Fruit of a given cultivar on gir-
dled trees or trees in sandy areas ripen earlier
than fruit on non-girdled trees or in loamy
soils. These fruit also tend to ripen more uni-
formly within the tree from top to bottom. A
skilful manager will consider these factors, as
well as others, and judge when and how often
an orchard should be harvested, and how
much fruit can be removed in any one pick-
ing. Because of the complexity of these factors,
there is no substitute for experience in making
these decisions. Strategies that are effective for
one grower may be ineffective for another
because of different organizational and mar-
keting situations and tactics. An example of
differing strategies is demonstrated by grower
A, who prefers to harvest five to eight times
for each cultivar where each harvest is 2 to 3
days apart. This is in contrast to grower B,
who prefers to pick only two or three times
with a longer interval in between harvests.
Grower A may decide that he does not mind
spending the extra money on increased labour
because he is achieving a higher packout per-
centage (less cullage). Grower B may not mind
a reduced packout percentage (more cullage)
because he is saving money on labour.
In Spain, the indigenous cultivar ‘Calanda’
is much appreciated by European consumers
and it dominates the late fresh market
because of its special characteristics. Fruit is
individually wrapped in a paper bag, is free
of pesticides, and during the development on
the tree reaches a uniform cream or straw
colour (Ferrer et al., 2005). For marketing pur-
poses, only slight blush is accepted, but green
or orange-yellowish colours are refused. In
this cultivar, firmness and skin colour charts
have been proposed to estimate the optimal
harvest point.
22.5 Temperature Requirements and
Management
Ideal storage conditions
The ideal peach storage temperature is −1°C
to 0°C. The flesh freezing point varies depend-
ing on SSC. Storage-room relative humidity
should be maintained at 90–95% and air
velocity of approximately 0.0236 m3/s is sug-
gested during storage (Lill et al., 1989; Thomp-
son et al., 1998).
Temperature management
The application of the ideal cooling require-
ments will depend on the specific operation
and the way to apply these requirements
depends on the scheduling of the packing
operation (Mitchell, 1987). Fruit can be cooled
in field bins by hydro-cooling or pre-cooling
(Fig. 22.3/Plate 234). Hydro-cooling is nor-
mally done by a conveyor-type hydro-cooler.
Fruit in field bins can be cooled to intermedi-
ate temperatures (5–10°C) provided packing
will occur the next day. If packing is to be
delayed beyond the next day, then fruit
should be thoroughly cooled in the bins to
near 0°C. In IB-susceptible cultivars fast cool-
ing within 8 h and maintaining fruit tempera-
ture near 0°C are traditionally recommended
(Mitchell, 1987). Fruit in packed containers
should be cooled to near 0°C. Even fruit that
were thoroughly cooled in the bins will warm
substantially during packing and should be
thoroughly re-cooled after packing. Forced-
air cooling is normally indicated after pack-
ing (Fig. 22.4/Plate 235). A rare exception to
the need for cooling after packing would be a
system that handles completely cold fruit and
provides protection against warming during
packing.
 Harvesting and Postharvest Handling 583

Fig. 22.3. Bin of peaches being pre-cooled on a conveyor-type hydro-cooler prior to packing.

Fig. 22.4. Packaged fruit in unitized pallet loads are stacked to form a forced-air cooling tunnel.
584 C.H. Crisosto and D. Valero
Water loss control
Economic loss to the grower can result when
as little as 8% of the fruit fresh weight is lost
(Crisosto et al., 1994). The economic loss is
due to both decreased weight of the fruit and
the unsightly shrivelling that occurs. While
there is a large variability in susceptibility to
water loss among cultivars, all cultivars must
be protected to ensure the best postharvest
life. Fruit waxes or coatings that are com-
monly used as carriers for postharvest fungi-
cides can reduce the rate of water loss when
excess surface brushing has not occurred.
Mineral oil waxes can potentially control
water loss better than vegetable oil and edible
coatings, although the use of waxes or coat-
ings is regulated according to destination
point requirements. The main ways to limit
fruit water loss include short cooling delays,
efficient waxing with gentle brushing, fast
cooling followed by storage under constant
low temperature and high relative humidity.
New temperature management approach
A new technique to delay IB symptoms and pre-
ripen fruit has been successfully introduced
Fig. 22.5. Storage temperature influences incidence and severity of internal breakdown in suscep-
tible cultivars.
to the California and Chilean industries. This
technique, described above, consists of a ~48 h
controlled cooling delay (Crisosto et al., 2004).
A preconditioning protocol has been devel-
oped and promoted among packers/shippers
(Crisosto et al., 2004). In this delivery system,
preconditioned peaches should be arriving at
the distribution centre at ~2.3–3.6 kgf firm-
ness, measured at the weakest point on the
cheeks. This new fruit delivery system is one
more approach to limit IB and enhance the
fruit-eating experience for consumers. Due to
physical and chemical changes occurring in
the fruit during a well-controlled precondi-
tioning treatment, peaches undergo fruit soft-
ening to the ‘ready to buy’ stage (~2.7–3.6
kgf). Thus, fruit become tastier, more aromatic
and juicier, resulting in high consumer accep-
tance. Generally, all peach cultivars should be
kept out of the ‘killing zone’ temperature range
of 2.2–10°C (Fig. 22.5/Plate 236). The ideal
storage temperature is from 0°C to 1.7°C.
Keeping fruit at this temperature will slow
softening and reduce shrivelling, decay and
the incidence of IB or mealy fruit. The exact
temperature management will be part of a
broader fruit preparation for consumers that
takes into account the firmness on arrival of
fruit and the fruit turning schedule (time that
 Harvesting and Postharvest Handling
fruit remain on display tables). This needs to
be coordinated with the store-level demand
and it will depend on a particular company’s
anticipated sales/consumption schedule
(fruit turning schedule). Cheek firmness is a
good tool to determine ripening stage (trans-
fer point, ready to buy, ready to eat, etc.),
while firmness measured at the weakest posi-
tion (shoulder, tips or suture) is well related
to potential impact and transportation dam-
ages. Fruit firmness does not accurately cer-
tify the quality of the preconditioning
execution, however.
22.6 Field Harvesting, Hauling
and Packaging
The goals of fruit harvesting should be to pick
fruit at optimum maturity and transport fruit
to the packing facility with no deterioration
in fruit quality. To do this requires proper
coordination between human resources, fruit
585
maturity, environmental factors, technical
resources and equipment. An understanding
of these factors and their relationship is essen-
tial to making the proper management deci-
sions for a given orchard situation.
Harvesting
Peach fruit are hand picked using bags (Fig.
22.6/Plate 237), baskets or totes. Most com-
mercial peach fruit operations use picking
bags and bins in their harvest operations
(Corelli Grappadelli, 2001). Peaches are dumped
in bins (Fig. 22.7/Plate 238) that are on the top
of trailers between rows in the orchard. If
totes are being used, they are placed directly
inside the bins. Baskets are placed on top of
modified trailers. Fruit picked at advanced
maturity stages and white-fleshed peaches
are generally picked and placed into baskets
or totes. Because of the availability of new
cultivars that adapt well to harvesting more
mature (softer), the increase in popularity of
Fig. 22.6. Peaches for fresh market are
hand picked. Harvesters work on lad-
ders using picking bags or baskets.
586 C.H. Crisosto and D. Valero
high-quality, less firm fruit (more mature)
and use of more sophisticated packinghouse
equipment, a large proportion of stone fruits
are being picked at a more advanced maturity
stage than historically.
Regardless of maturity, a number of pre-
cautions should be taken with any harvest
operation (Mitchell and Kader, 1989b). Har-
vesters should be instructed to treat the fruit
as gently as possible at every stage of the har-
vest process. When emptying bags into the
transport bins, care should be taken to ensure
that the fruit are not dumped into the bin
from a high height. Again, this is where the
crew leader is helpful in reducing problems.
Picking bags and buckets should be kept
clean. There appears to be a relationship
between inking, surface abrasion and dirty
containers. Washing picking bags at regular
intervals may be helpful in reducing this
problem. After harvest, but while still in the
field, fruit should be protected from exposure
to direct sunlight and excess heating (Fig. 22.8/
Fig. 22.7. Harvesters transfer peaches
to field bins which are moved through
the field on low trailers.
Plate 239). Insulated bin covers are the most
beneficial shading technique. Some growers
use cloth coverings to protect the fruit. On
very hot days these should be supported
above the fruit because direct contact can allow
enough heat to pass through to cause fruit
scald. After harvest fruit should be hauled to
a cooling facility as quickly as possible. If
there is a delay in transportation, fruit should
be stored in a cool, shaded area. Temporary
structures near the harvest location are often
constructed from shade cloth material. Care
should be taken as the harvested fruit are
being loaded for transport to the packing
facility. Forklift drivers should be informed of
the importance of treating fruit gently when
loading and unloading bins of fruit.
Fruit hauling
Fruit are hauled for short distances by trailer,
but if the distance is longer than 10 km, bins
 Harvesting and Postharvest Handling
Fig. 22.8. View of a shaded loading area to protect fruit from excess heating while awaiting transpor-
tation to the packinghouse.
are loaded on trucks for transportation to
packinghouses. Peaches are transported from
orchard to packinghouse and cooler as soon
as possible after harvest. Fruit should be
shaded during any delay between harvest
and transport. Tractor drivers should be
instructed to drive slowly and smoothly. Severe
fruit damage can result from poor driving
practices, especially on turns and starts. There
appears to be a benefit to using ‘suspension-
type’ bin trailers instead of solid axle trailers.
These trailers tend to ride more smoothly.
Similar results can be obtained to a lesser
degree by lowering tyre air pressure. Both of
these procedures are probably more helpful
for road transport conditions than field trans-
port. Unloading of trailers should also be per-
formed as gently as possible. Care should be
taken to educate workers as to the importance
of this process. It is helpful if the unloading
area is smooth and spacious to eliminate
bumping and jarring. During hauling, drivers
should reduce and eliminate jarring and bounc-
ing. By choosing proper transportation routes
and avoiding rough, bumpy roads fruit injury
can be minimized. Position of fruit on the
trailer is also important. Within-bin vibration
587
levels are highest at the front of the trailer,
intermediate in the rear, and lowest in the
middle of the trailer. The addition of air-sus-
pension systems to trailers has been shown to
be of tremendous value in reducing this type
of fruit damage. Plastic bin liners and padded
bin covers have also been demonstrated to
reduce transport injury. Research has shown
that thick bubble padding is more beneficial
than thin, and that larger bubbles are pre-
ferred to small (Mitchell and Kader, 1989b).
Fruit packaging
At the packinghouse the fruit are dumped
(mostly using dry bin dumps, Fig. 22.9/Plate
240) and cleaned (Mitchell and Kader, 1989b).
Here debris is removed and fruit may be
washed with chlorinated water. Peaches are
normally wet-brushed to remove the trichomes
(fuzz), which are single-cell extensions of
epidermal cells and protect fruit from new
inoculations. Waxing and fungicide treatment
may follow, depending on country regulations
(http://www.fas.usda.gov/htp/MRL.asp).
588 C.H. Crisosto and D. Valero
Water-emulsifiable waxes are normally used,
and fungicides may be incorporated into
the wax.
Sorting and sizing operation
Sorting is carried out to eliminate fruit with
visual defects, cuts and wounded areas and
sometimes to divert fruit of high surface colour
to a high-maturity pack (Fig. 22.10/Plate 241).
Attention to details of sorting line efficiency is
especially important with peaches, where a
range of fruit colours, sizes and shapes can be
encountered. Sizing segregates fruit by either
weight or dimension. Sorting and sizing
equipment must be flexible to efficiently han-
dle large volumes of small fruit or smaller
volumes of larger fruit. In California, most
yellow-fleshed peaches are packed into two-
layer (trays) boxes (Fig. 22.11/Plate 242). In
the eastern USA, most are volume-fill packed.
Fig. 22.9. Dry bin dumping of fruit on
to a commercial packing line.
Electronic weight sizers are used to automati-
cally fill shipping containers (Fig. 22.12/Plate
243). Most of the white-fleshed peaches and
‘tree ripe’ peaches are packed into one-layer
(tray) boxes (flat). In some cases, peaches are
also packed in small-size plastic bags or clam-
shell plastic containers. In some operations,
mechanical place-packing units use hand-as-
sisted fillers where the operator can control
the belt speed to match the flow of fruit into
plastic trays. Limited volumes of high-matu-
rity peaches are ‘ranch’ or ‘field’ packed at the
point of production. In a typical ‘ranch’ or
‘field’ packed operation, fruit of high maturity
and quality are picked into buckets or totes
that are carried by trailer to the packing area.
These packers work directly from the buckets
to sort, grade, size, and pack fruit into plastic
trays. In these cases, the fruit are not washed,
brushed, waxed or fungicide-treated. In other
cases, fruit are picked into buckets or totes
but then dumped into a smooth-operating,
 Harvesting and Postharvest Handling 589

Fig. 22.10. Sorting peaches by skin colour and removing blemished fruit.

Fig. 22.11. Packers sizing, sorting and packing fruit by hand into two-layer tray packs.
590 C.H. Crisosto and D. Valero
low-volume packing line for washing, brush-
ing, waxing, sorting and packaging. Because
of less handling of the fruit, a higher maturity
can be used, and growers can benefit from
increased fruit size, red colour and greater
yield. High-quality fruit can also be produced
by managing orchard factors properly and
picking fruit that are firm. But in this latter
case, ripening at the retailer will be essential
to ensure good flavour quality for consumers.
Shipping and transportation
At the shipping point, fruit should be cooled
and held near or below 0°C according to their
freezing point. During transportation, if IB-
susceptible cultivars are exposed to 5°C their
postharvest life can be significantly reduced
(Mitchell and Kader, 1989a). Peach storage
and overseas shipments should be at or below
0°C. Maintaining these low pulp temperatures
requires knowledge of the freezing point of
Fig. 22.12. Fruit moving on to an
electronic weight sizer.
the fruit, the temperature fluctuations in the
storage system and equipment performance
(Thompson et al., 1998; Thompson, 2002).
Holding peaches at these low temperatures
minimizes both the losses associated with rot-
ting organisms, excessive softening and water
losses, and the deterioration resulting from IB
in susceptible cultivars, therefore optimizing
their postharvest life (Mitchell, 1987).
22.7 Cull Utilization
Potential uses
The main use of peach culls is for cattle feed
because culled peach is palatable and a good
source of energy (Fig. 22.13/Plate 244), but it
is low in protein and has other characteristics
that make it different from other feed sources
(Thompson, 2002). For example, peaches con-
tain ~85% water, 9% digestible dry matter, 5%
pits and 2% indigestible dry matter. The high
 Harvesting and Postharvest Handling
Fig. 22.13. Cull removal and disposal can be a major problem and expense in peach packing.
water content diminishes the real value as
feed because it makes culls expensive to
transport, requires large trough volumes, and
allows the feed to spoil quickly. If fed in large
proportions, culled fruit causes almost con-
tinuous urination and consequently the ani-
mals require a high amount of salt. The only
potential advantage to the high water content
is that animals in a remote, dry location will
not need extra water hauled to them. Low
protein levels in culled fruit limit the quantity
that can be fed where rapid weight gain is
important, such as in feed lots. For example,
only about 20% of the ration can be composed
of culls (Thompson, 2002).
The use of culls for fuel alcohol produc-
tion is limited mainly by the low sugar con-
tent; thus peach is not included in this group
(Thompson, 2002). The 8 to 12% sugar content
of most culled peaches results in an alcohol
yield of about 42 l/t (10 gal/t) of fruit, which
is too low compared with potatoes (83 to 104
l/t) or maize (375 l/t). This low yield makes it
uneconomical in addition to the waste man-
agement problem. Unfortunately, the limits to
the use of culls often result in large portions
of them being discarded. Improper disposal
can cause sanitary and pollution problems
(Thompson, 2002). Flies and odour problems
can be prevented by ensuring rapid drying.
591
Fly maggots hatch into adults within 7 to 10
days, and odour problems can develop before
flies appear. The culls should be crushed and
spread no more than one or two layers deep;
sometimes this is done on orchard roads or
fallow fields. Culls can be disked into the soil,
although this tends to cover the fruit with soil
and slows drying; also, insects or diseases that
may have caused the fruit to be culled in the
first place may infect a future crop. Disposal
sites should be as far away from neighbours as
possible. Flies can travel up to 8 km (5 miles)
from the place where they hatch. Culls should
not be dumped near streambeds. Fruit cull
piles can attract the dumping of many other
kinds of refuse. If culls are deposited away
from the point of production, use municipal
solid waste disposal sites if available. Some
culls can be turned into dried fruit for human
consumption. However, good-quality dried
fruit is made only from good-quality fresh
fruit. Only undersized or slightly overripe fruit
should be considered for drying.
Situation in California
In general, peach culls are going for frozen or
canned peaches or juice, dried for charity dona-
tion, or used for livestock feed. When fruit have
592 C.H. Crisosto and D. Valero
worms and decay they are utilized as green
waste for compost. The amount of culls varies
according to season, cultivar and other condi-
tions from ~10 to 30% of total production. The
decision on the cull utilization is made based
on returns. In general, when the reason for
disposal has been small sizes and mainly cos-
metic blemishes, fruit still have value for
human consumption and can be frozen, canned
or used for making juice or other value-added
products.
Other uses
A very limited peach fresh-cut business has
been developed because of the short market
life of this produce (Gorny et al., 1999). The
optimal ripeness for preparing fresh-cut
peach slices is when the flesh firmness reaches
1.4–2.7 kgf, and these slices can retain good
eating quality for 2–8 days (depending on
cultivar) while kept at 5°C and 90–95% rela-
tive humidity. Post-slice dips in ascorbic acid
and calcium lactate or use of MAP may
slightly prolong the shelf-life of peach slices.
Recently, mild heat pre-treatments (40°C for
70 min) before minimal processing and pack-
ing under passive MAP conditions were effec-
tive in inducing firmness (Steiner et al., 2006),
while preserving nutritional quality (organic
acids and vitamins).
22.8 Fruit Handling at Retail Distribution
Fruit preparation for consumers
Because peaches are a climacteric fruit they are
usually harvested when they reach a minimum
or higher maturity, but are not completely
ripe (‘ready to eat’). Initiation of the ripening
process must occur before consumption to
satisfy consumers. It has been demonstrated
that most consumers are satisfied after eating
ripe peaches. A ripe or ‘ready to eat’ peach is
defined when flesh firmness is approximately
0.9–1.4 kgf. Peaches with firmness below 2.7–
3.6 kgf (‘ready to buy’) are becoming attrac-
tive to consumers while still tolerating retail
handling. For this reason, this range is also
called the transfer point. Thus, a delivery sys-
tem should target store displays of peaches
with firmness below 2.7–3.6 kgf and ensure
that consumers are eating peaches that are
‘ready to eat’. Promotional programmes should
be established to educate consumers on rip-
ening issues.
As the market for fresh produce is grow-
ing steadily, the need for assuring quality is
increasing in European markets. In this sense,
the market splits into two classes of produce:
commodity (low price) and high-quality fruits,
which are in demand from the new export
markets. Accordingly, the combination of
colour measurements using two wavelengths
(450 and 680 nm) with non-destructive firm-
ness testing gave a good procedure for classi-
fying peaches for ripeness (Ruiz-Altisent
et al., 2006). It has been reported that the pres-
ence of g-decalactone, d-octalactone and g-oc-
talactone can be used to indicate the maturity
stage for harvesting peaches (Lavilla et al., 2002).
Two sensors based on solid-state detection of
gas concentration of g- and d-decalactone
(which increase significantly during the final
stages of ripeness) were able to grade peaches
by ripening stages. Moreover, the sensors
were capable of detecting skin breakage pro-
duced by mechanical or pathological causes
and showed a good correlation with firmness
measurements (Moltó et al., 1999).
Fruit buyer handling
If retailers are receiving mature peaches (4.5–
7.3 kgf), the ripening process can be initiated
at the distribution centres (receivers). Detailed
ripening protocols for retail handlers, ware-
house and produce managers have been
developed and well promoted (Crisosto and
Parker, 1997). In general, peach cultivars har-
vested commercially will ripen properly
without exogenous ethylene application.
Temperature conditions for peaches during
and after ripening should be adjusted accord-
ing to the desired sales/consumption sched-
ule. We encourage that further fruit ripening,
if necessary, be done at the distribution level.
The rate of fruit softening (pressure loss (kgf)
 Harvesting and Postharvest Handling
per day) varies among peach cultivars and
can be controlled by the storage temperature
used. Fruit stored at 2.2°C will soften slower
than fruit stored at 20°C. When the fruit reaches
the transfer firmness mentioned above, the
rate of softening slows. However, rate of soft-
ening also varies according to orchard and
season, so firmness measurements should be
taken to protect fruit integrity during the rip-
ening process. These fruit will reach their
‘ready to eat’ firmness of 0.9–1.8 kgf after 2–3
days at room temperature (15–20°C dry retail
display). Firmness is measured mid-cheek,
perpendicular to the fruit suture.
When kept at 2.2°C or below, peaches
should be shipped out of the distribution cen-
tre within 4–5 days (ideally within 2–3 days).
To the extent that the distribution centre does
not have rooms that can maintain tempera-
tures at this 2.2°C and below range, it might
make more sense to set up two shipments per
week from the shipper to ensure better tem-
perature control and extend the market life of
the product. In general, soft fruit are more
susceptible to bruising than hard fruit. To
reduce potential physical damage occurring
during transportation from the distribution
centres to retail stores and handling at the
retail stores, we suggest transferring fruit to
Fig. 22.14. Peach fruit display at a retail store.
593
the retail store before fruit reaches no lower
than 1.8–2.3 kgf measured on the weakest
position for tray-packed peaches and nectar-
ines. In general, the shoulder position is the
weakest point on mid- or late-season fruit. As
bruising incidence varies among cultivars,
and bruising potential is related to each spe-
cific operation, producers should fine-tune
their transfer points for their handling situa-
tion. These are general handling guidelines
but they need to be modified and assessed in
light of one’s particular company facilities,
logistics and customer requirements.
Peach handling at retail stores
Ideally, peaches should be transported at
0–1.7°C from the distribution centre and kept
at 0–1.7°C prior to transfer to dry/warm table
for display. In situations where fruit tempera-
ture cannot be maintained out of the ‘killing
zone’, it would be preferable to move fruit
fast. Firmness measurements need to be con-
sidered in the decision-making process
(Crisosto and Mitchell, 2002).
Peaches should ideally be arriving from
the distribution centre to the retail stores with
594 C.H. Crisosto and D. Valero

firmness in the range of 1.8–2.7 kgf (weakest
position) or 2.7–3.6 kgf (cheeks). This fruit is
at the ‘ready to buy’ or ‘transfer point’ stage
of ripening and within ~48–72 h at 20°C
should be ‘ready to eat’ in the 0.9–1.8 kgf
firmness range. This is the firmness range at
which most consumers claim the highest sat-
isfaction when eating peaches.
Produce managers need to be educated
about this new ‘ready to buy’ type of fruit (pre-
conditioned) to minimize mechanical damage
and expedite an effective rotation (first in,
first out). Peaches should be displayed on dry
tables and labelled well as ‘ready to buy/eat’,
and consumers should understand that this
fruit is riper than conventionally packed tree
fruit (Fig. 22.14/Plate 245). In order to protect
these fruit, the display should be no more
than two layers deep and in-box display
should be attempted. As tree fruit will con-
tinue to ripen on the display warm/dry table,
they should be checked often and the softest
fruit be placed at the front of the display. Fruit
that reach the ‘ready to eat’ ripeness of 0.9–1.4
kgf cheek firmness need to be sold quickly or
refrigerated to extend their shelf-life. It is
essential that consumers be instructed that
this type of fruit should be refrigerated if it is
not going to be consumed within 3 days of
purchase.

References

Anderson, R.E. (1979) The influence of storage temperatures and warming during storage on peach and nec-
tarine fruit quality. Journal of the American Society for Horticultural Science 104, 459–461.
Byrne, D.H. (2002) Peach breeding trends. Acta Horticulturae 592, 49–59.
Carbonaro, M., Mattera, M., Nicoli, S., Bergamo, P. and Cappelloni, M. (2002) Modulation of antioxidant
compounds in organic vs conventional fruit (peach, Prunus persica L., and pear, Pyrus communis L.).
Journal of Agricultural and Food Chemistry 50, 5458–5462.
Ceponis, M.J., Cappellini, R.A., Wells, J.M. and Lightner, G.W. (1987) Disorders in plum, peach and nectarine
shipments to the New York market, 1972–1985. Plant Disease 71, 947–952.
Cheng, G.W. and Crisosto, C.H. (1997) Iron–polyphenol complex formation and skin discoloration in peach-
es and nectarines. Journal of the American Society for Horticultural Science 122, 95–99.
Corelli Grappadelli, L. (2001) Peach handling and marketing in Italy. In: Proceedings of 60th National Peach
Council Convention, Hershey, Pennsylvania, 30 January–1 February, pp. 39–45.
Crisosto, C.H. (1994) Stone fruit maturity indices: a descriptive review. Postharvest News and Information 5,
65–68.
Crisosto, C.H. (2002) How do we increase peach consumption? Acta Horticulturae 592, 601–605.
Crisosto, C.H. and Labavitch, J.M. (2002) Developing a quantitative method to evaluate peach (Prunus
persica) flesh mealiness. Postharvest Biology and Technology 25, 151–158.
Crisosto, C.H. and Mitchell, F.G. (2002) Peach, nectarine and plum. In: Kader A.A. (ed.) Postharvest Technology
of Horticultural Crops. Special Publication No. 3311. University of California, Division of Agriculture
and Natural Resources, Oakland, California, pp. 345–351.
Crisosto, C.H. and Parker, D. (1997) Stone fruit ripening protocol for receivers. Slide set v98-c with cassette.
University of California, Division of Agriculture and Natural Resources, Oakland, California.
Crisosto, C.H., Johnson, R.S., Luza, J.G. and Crisosto, G.M. (1994) Irrigation regimes affect fruit soluble solids
concentration and rate of water loss of ‘O’Henry’ peaches. HortScience 29, 1169–1171.
Crisosto, C.H., Mitchell, F.G. and Johnson, S. (1995) Factors in fresh market stone fruit quality. Postharvest
News and Information 6, 17–21.
Crisosto, C.H., Johnson, R.S., DeJong, T. and Day, K.R. (1997) Orchard factors affecting postharvest stone fruit
quality. HortScience 32, 820–823.
Crisosto, C.H., Garner, D., Cid, L. and Day, K.R. (1999a) Peach size affects storage, market life. California
Agriculture 53, 33–36.
Crisosto, C.H., Johnson, R.S., Day, K.R., Beede, B. and Andris, H. (1999b) Contaminants and injury induce
inking on peaches and nectarines. California Agriculture 53, 19–23.
Crisosto, C.H., Mitchell, F.G. and Ju, Z. (1999c) Susceptibility to chilling injury of peach, nectarine, and plum
cultivars grown in California. HortScience 34, 1116–1118.
Crisosto, C.H., Day, K.R., Crisosto, G.M. and Garner, D. (2001a) Quality attributes of white flesh peaches and
nectarines grown under California conditions. Journal of the American Pomological Society 55, 45–51.
 Harvesting and Postharvest Handling 595

Crisosto, C.H., Slaughter, D., Garner, D. and Boyd, J. (2001b) Stone fruit critical bruising thresholds. Journal
of the American Pomological Society 55, 76–81.
Crisosto, C.H., Garner, D.T., Andris, H.L. and Day, K.R. (2004) Controlled delayed cooling extends peach
market life. HortTechnology 14, 99–104.
Fernández-Trujillo, J.P. and Artés, F. (1998) Chilling injuries in peaches during conventional and intermittent
warming storage. International Journal of Refrigeration 21, 265–272.
Fernández-Trujillo, J.P., Martínez, J.A. and Artés, F. (1998) Modified atmosphere packaging affects the
incidence of cold storage disorders and keeps ‘flat’ peach quality. Food Research International 31,
571–579.
Ferrer, A., Remón, S., Negueruela, A.I. and Oria, R. (2005) Changes during the ripening of the very late season
Spanish peach cultivar Calanda. Feasibility of using CIELAB coordinates as maturity indices. Scientia
Horticulturae 105, 435–446.
Garner, D., Crisosto, C.H. and Otieza, E. (2001) Controlled atmosphere storage and aminoethoxyvinyl-
glycine postharvest dip delay post cold storage softening of ‘Snow King’ peach. HortTechnology 11,
598–602.
Gil, M.I., Tomás-Barberán, F.A., Hess-Pierce, B. and Kader, A.A. (2002) Antioxidant capacities, phenolics
compounds, carotenoids, and vitamin C content of nectarine, peach, and plum cultivars from California.
Journal of Agricultural and Food Chemistry 50, 4976–4982.
Goristein, S., Martín-Belooso, O., Lojek, A., Ciz, M., Soliva-Fortuny, R., Park, Y.S., Caspi, A., Libman, I. and
Trakhtenberg, S. (2002) Comparative content of some phytochemicals in Spanish apples, peaches and
pears. Journal of the Science of Food and Agriculture 82, 1166–1170.
Gorny, J.R., Hess-Pierce, B. and Kader, A.A. (1999) Quality changes in fresh-cut peach and nectarine slices as
affected by cultivar, storage atmosphere and chemical treatments. Journal of Food Science 64, 429–
432.
Harding, P.L. and Haller, M.H. (1934) Peach storage with special reference to breakdown. Proceedings of the
American Society for Horticultural Science 32, 160–163.
Kader, A.A. and Mitchell, F.G. (1989a) Maturity and quality. In: LaRue, J.H. and Johnson, R.S. (eds) Peaches,
Plums, and Nectarines: Growing and Handling for Fresh Market. Publication No. 3331. University of
California, Division of Agriculture and Natural Resources, Oakland, California, pp. 191–196.
Kader, A.A. and Mitchell, F.G. (1989b) Postharvest physiology. In: LaRue, J.H. and Johnson, R.S. (eds) Peaches,
Plums, and Nectarines: Growing and Handling for Fresh Market. Publication No. 3331. University of
California, Division of Agriculture and Natural Resources, Oakland, California, pp. 158–164.
LaRue, J. (1989) Introduction. In: LaRue, J.H. and Johnson, R.S. (eds) Peaches, Plums, and Nectarines: Growing
and Handling for Fresh Market. Publication No. 3331. University of California, Division of Agriculture
and Natural Resources, Oakland, California, pp. 1–2.
Lavilla, T., Recasens, I., López, M.L. and Puy, J. (2002) Multivariate analysis of maturity stages, including quality
and aroma, in ‘Royal Glory’ peaches and ‘Big Top’ nectarines. Journal of the Science of Food and
Agriculture 82, 1842–1849.
Lill, R.E., O’Donoghue, E.M. and King, G.A. (1989) Postharvest physiology of peaches and nectarines. Horti-
cultural Reviews 11, 413–452.
Martínez-Romero, D., Valero, D., Serrano, M., Burló, F., Carbonell, A., Burgos, L. and Riquelme, F. (2000)
Exogenous polyamines and gibberellic acid effects on peach (Prunus persica L) storability improvement.
Journal of Food Science 65, 288–294.
Mitchell, F.G. (1987) Influence of cooling and temperature maintenance on the quality of California grown
stone fruit. International Journal of Refrigeration 10, 77–81.
Mitchell, F.G. and Kader, A.A. (1989a) Factors affecting deterioration rate. In: LaRue, J.H. and Johnson, R.S.
(eds) Peaches, Plums, and Nectarines: Growing and Handling for Fresh Market. Publication No. 3331.
University of California, Division of Agriculture and Natural Resources, Oakland, California, pp. 165–
178.
Mitchell, F.G. and Kader, A.A. (1989b) Field handling and packing. In: LaRue, J.H. and Johnson, R.S. (eds)
Peaches, Plums, and Nectarines: Growing and Handling for Fresh Market. Publication No. 3331.
University of California, Division of Agriculture and Natural Resources, Oakland, California, pp. 197–
208.
Moltó, E., Selfa, E., Ferriz, J., Conesa, E. and Gutierrez, A. (1999) An aroma sensor for assessing peach quality.
Journal of Agricultural Engineering Research 72, 311–316.
Nanos, G.D. and Mitchell, F.G. (1991) High-temperature conditioning to delay internal breakdown develop-
ment in peaches and nectarines. HortScience 26, 882–885.
596 C.H. Crisosto and D. Valero

Okie, W.R. (1998) Handbook of Peach and Nectarine Varieties: Performance in the Southeastern United
States and Index of Names. USDA Agriculture Handbook No. 714. US Department of Agriculture,
Washington, DC.
Proteggente, A.R., Pannala, A.S., Paganga, G., Van Buren, L., Wagner, E., Wiseman, S., Van de Put, F. and
Dacombe, C. (2002) The antioxidant activity of regularly consumed fruit and vegetables reflects their
phenolic and vitamin C composition. Free Radical Research 36, 217–233.
Romani, R.J. and Jennings, W.G. (1971) Stone fruits. In: Hulme, A.C. (ed.) The Biochemistry of Fruits and Their
Products, Vol. 2. Academic Press, New York, pp. 411–436.
Ruiz-Altisent, M., Lleó, L. and Riquelme, F. (2006) Instrumental quality assessment of peaches: fusion of
optical and mechanical parameters. Journal of Food Engineering 74, 490–499.
Sansavini, S., Corelli Grappadelli, L., Costa, G., Lugli, S., Marangoni, B., Tagliavini, M., Ventura, M., Abeti, D.,
Feralli, S., Marani, G., Mascanzoni, G., Molducci, S., Proni, R., Sama, A., Spada, G., Vitali, S., Turroni, P.,
Minguzzi, A. and Randi, M. (2000) Ricostituzione degli impianti e nuovi indirizzi produttivi della peschi-
coltura romagnola. In: Atti XXIII Convegno Peschicolo, Ravenna, 12–13 September 1997, pp. 62–74.
Serrano, M., Martínez-Romero, D., Castillo, S., Guillén, F. and Valero, D. (2004) Effects of pre-harvest sprays
containing calcium, magnesium and titanium on the quality of peaches and nectarines at harvest and
during post-harvest storage. Journal of the Science of Food and Agriculture 84, 1270–1276.
Shewfelt, R.L., Meyers, S.C., Prussia, S.E. and Jordan, J.L. (1987) Quality of fresh-market peaches within the
postharvest handling system. Journal of Food Science 52, 361–364.
Smith, W.H. (1934) Cold storage of Elberta peaches. Ice and Cold Storage 37, 54–57.
Steiner, A., Abreu, M., Correia, L., Beirão-da-Costa, S., Leitão, E., Beirão-da-Costa, M.L., Empis, J. and Moldao-
Martins, M. (2006) Metabolic response to combined mild heat pre-treatments and modified atmosphere
packaging on fresh-cut peach. European Food Research Technology 222, 217–222.
Thompson, J.F. (2002) Cull utilization. In: Kader, A.A. (ed.) Postharvest Technology of Horticultural Crops.
Special Publication No. 3311. University of California, Division of Agriculture and Natural Resources,
Oakland, California, pp. 41–43.
Thompson, J.F., Mitchell, F.G., Rumsey, T.R., Kasmire, R.F. and Crisosto, C.H. (eds) (1998) Commercial Cooling
of Fruits, Vegetables, and Flowers. Publication No. 21567. University of California, Division of Agriculture
and Natural Resources, Oakland, California.
Tomás-Barberán, F.A., Gil, M.I., Cremin, P., Waterhouse, A.L., Hess-Pierce, B. and Kader, A.A. (2001) HPLC-
DAD-ESIMS analysis of phenolic compounds in nectarines, peaches, and plums. Journal of Agricultural
and Food Chemistry 49, 4748–4760.
USDA (2003) Composition of Foods: Fruits and Fruit Juices – Raw, Processed, Prepared. USDA Agriculture
Handbook No. 8–9. US Department of Agriculture, Washington, DC (www.nal.usda.gov/fnic/foodcomp).
Valero, D., Serrano, M., Martínez-Madrid, M.C. and Riquelme, F. (1997) Polyamines, ethylene, and physio-
chemical changes in low-temperature-stored peach (Prunus persica L. cv. Maycrest). Journal of Agricul-
tural and Food Chemistry 45, 3406–3410.
Von Mollendorff, L.J. (1987) Woolliness in peaches and nectarines: a review. 1. Maturity and external factors.
Horticultural Science/Tuinbouwetenskap 5, 1–3.
Von Mollendorff, L.J., Jacobs, G. and De Villiers, O.T. (1992) Postharvest factors involved in the development
of chilling injuries in peaches and nectarines. Journal South African Horticultural Science 2, 58–68.
 Index

10-Point Management Programme 395, 518 chlorogenic acid 577

see also Peach tree short life (PTSL) cyanidin-3-glucoside 577
epicatechin 577
flavonols 577
Acari see Mites Aphids 55, 88, 334, 341, 436, 468, 487, 494–496
Adaptation 69 green peach aphid, Myzus persicae 55,
AFLP see Amplified fragment length 495–496
polymorphism Peach latent mosaic viroid (PLMVd) vector
Aggregation pheromone 478, 484 454
Agrobacterium spp. Plum pox virus (PPV) vector 441–442, 494
Agrobacterium radiobacter 420, 423 resistance 11, 63, 70, 74–75, 162, 164, 204
Agrobacterium tumefaciens 99–100, 194, Apical meristem culture 447
203–204, 206, 208, 409, 418, 510–511 see also Propagation
see also Crown gall Apoplastic phloem loading 248–249
Allergenicity 563–564 Appearance see Fruit
see also Nutritional value Apple chlorotic leaf spot virus (ACLSV) 456–457,
Alternate host 357 459
Amplified fragment length polymorphism (AFLP) Apricot latent virus 456
86, 93, 97 Arabidopsis 85, 92, 96
Anarsia lineatella see Peach twig borer A. thaliana 554, 567
Aneuploid 68 Arabis mosaic virus (ArMV) 457, 524
Anthocyanin 10, 18–19, 30, 63, 176, 183, 186, 188, Armillaria root rot (syn. oak root rot, shoe string
341, 561, 565, 577 root rot) 355, 386, 388–392, 394
anthocyaninless 10, 18, 22, 63, 67, 71, 165 A. mellea 355, 389, 390, 403
deficiency 10, 63, 67 A. mellea (oak root rot) 198, 205, 207, 210,
Anthracnose 212, 214
Colletotrichum acutatum and Colletotrichum A. ostoyae 355, 389
gloeosporiodes 401–403 A. tabescens 198, 355, 389
Gloeosporium laeticolor Berk 55 A. tabescens (oak root rot) 198, 205
Antioxidants 537, 539, 562–563 rhizomorph 355, 389–392, 403
ascorbic acid 576–577, 592 Armoured scale (Hemiptera: Diaspididae)
capacity 577 San Jose scale (SJS), Quadraspidiotus
carotenoids 576–577 perniciosus 492–494
phenolics 576–577 white peach scale (WPS), Pseudaulacaspis
catechin 577 pentagona 491–492, 494

597
598 Index

ArMV see Arabis mosaic virus Biological control 480, 492, 494
Aroma (fragrance) 17, 19–21, 537–539, 550, 561, Black knot (Apiosorina morbosa) 401–402
576, 584 Blush (overcolour) see Colour
benzaldehyde 550, 562 Boron 314–316
δ-decalactone 550, 562 deficiency 314–316
γ-decalactone 550, 562 toxicity 315
linalool 550, 562 Botryosphaeria dothidea see Fungal gummosis
Aromia bungii (Trunk borer) 55 Botryosphaeria fruit rot (B. dothidea, B. obtusa and B.
Ascorbic acid 576–577, 592 rhodinia) 401–402
see also Antioxidants Botrytis cinerea (grey mould) 356, 396, 398, 401, 577
ATS 294 Brachytic dwarf 165–166, 199
‘Axis central’ see Training system Breeding (classical) 68–81
Breeding goals
cold-hardiness 142–144, 151–152, 154,
Bacillus thuringiensis 474 165–167
Bacteria and bacterial diseases 407–409, 411–423 low-chill breeding
characteristics of bacteria 407–408 Australia 113
bacterial streaming 419 Brazil 117–118
environment 416 current 110–121
epiphytes 416 heritability 122–123, 125, 132
Bacterial artificial chromosome (BAC) 92–96 history 108–110
BAC insert (contig) 93–96, 97 inbreeding 110
Bacterial canker complex (BCC) 409, 411–416, Mexico 118–119
515–516, 518 nectarine 145, 147–151, 153–163, 165,
bacterial canker (Pseudomonas syringae pv. 167–169
syringae) 197, 205, 209–211, 214, South Africa 119
515–516, 518, 520 Taiwan 119
Bacterial decline syndrome 411 Thailand 119
symptoms 412–415 tree architecture 146, 151, 153–154,
canker 413 165–166
cold injury 411, 414, 415 USA 119–121
dead bud 412 low-chill breeding management 129–133
discoloured streaks in wood 413, 414 hybrid seed production 129–131
sucker growth 412 seedling populations 132–133
Bacterial leaf spot (Xanthomonas pruni) 55, 409, 416 testing advanced selections 133
infection conditions 417 low-chill breeding objectives 121–129
management 417–418 blind nodes 123, 124
chemical 418 bud drop 123
copper 418 disease resistance 127–129
host resistance 417 flower bud density 123
oxytetracycline 418 fruit development period 123–125
pathogen overwintering 417 fruit firmness 126–127
resistance 143, 145–146, 153 fruit shape 125–126
symptoms 416–422 low chilling requirement 121–122
leaf chlorosis and lesions 416–417 time of flowering 122–123
lesions on mature fruit 416, 420 Breeding programmes 139–174
lesions on young fruit 416, 419 Anderson 147–149
newly formed leaf lesions 416, 418 Arkansas 153
spring cankers 416–417, 421–422 Bradford 159
summer cankers 417, 421 Bulgaria 161
Xanthomonas arboricola pv. pruni 409 Burchell 159–160
BCC see Bacterial canker complex California 146–150, 154–160
Beer 51 China 166, 168
BG33R 520–521 conventional 44, 45
Biofix 472, 475, 494 early history 140–141
Biofumigation 519, 527 England 141, 173
Biolistics (particle bombardment) 99–100 flat shape 151, 154, 158, 162, 167–168
 Index 599

France 161–163, 166 Carbaryl 296

Georgia 141, 144–145, 150, 154 Carbon
Greece 163 assimilation 245
Hungary 163 partitioning 284–285
Ilinois 142–143 Carboxylation efficiency 245
Iowa 142–143 Carotenes 52, 561
Italy 146, 161, 163–166 Carotenoids 17–18, 182–183, 562, 576–577
Japan 168–169 β-carotene 562
Korea 169 β-cryptoxanthin 562
Louisiana 146, 153–154 Vitamin A 562
Maryland 141, 144 Cat-facing injury 334–335, 341, 481–483
Massachusetts 143 Catechin 577
Merrill 147 Catharathus roseus 411
Michigan 144, 151–153 Cells 290, 292
New Hampshire 143–144 Cellulose 562
New Jersey 143, 151 Central leader 55, 70, 255, 274–276
New York 142 density 274
New Zealand 169 photographs 275–276
North Carolina 145–146, 154 pruning 70, 274
Ontario 143, 153 training 274
Poland 165 see also Training system
Romania 166–167 Certification programmes 403, 442–443, 447,
Serbia 167 449–450, 458
South Africa 169 see also Viruses, indexing
Spain 167 Character inheritance 61–66
Sun World 160–161 Chemicals 293–294, 296
Texas 141, 144, 160 fertilizers 51
Ukraine 167 plant growth regulation 48
USDA 144–145, 150–151, 154–158 Cherry rasp leaf virus (CRLV) 524
Virginia 143 Chilling injury see Deterioration problems; Internal
West Virginia 144, 151 breakdown (IB)
Zaiger 158–159 Chilling requirement 17, 26–27, 30, 48, 55, 107,
Brown rot (Monilinia fructicola, M. fructigena, 117, 119, 121–122, 131–132, 144, 159–160,
M. laxa) 55, 118, 127–128, 338, 353–354, 164–165, 212, 223, 231, 252, 279
356–360, 396, 401 China Fruit Plant Monograph – Peach Flora 45
blossom blight 357–359, 361, 396 Chinese Spring Festival holiday 57
green fruit rot 354, 357–358, 361, 396 Chip budding 449
fruit rot 356–359, 396–397 see also Propagation, clonal
mummified fruit (mummy) 357–358, 400 Chipmunk, northwestern (Eutamias amoenus)
resistance 143, 163–164, 166–167 340
twig blight 357–358 Chlorogenic acid 562, 577
Budding 237, 239 Chlorophyll 561
see also Propagation Chromosomal walk 97
Bulked segregant analysis 88, 97 Chromosome 85
Citrate 557
Codling moth Cydia pomonella 472–473
Calcareous soil 195, 206–207, 209–212, 214 distinction from oriental fruit moth 470
Calcium 310–311 Coefficient of regression 66
deficiency 310–311 Cold hardiness 69, 92, 141–143, 151, 164–166, 168,
fruit quality 311 196, 201–202, 204–205, 214, 270, 319, 378,
Candida inconspicua (sour pit) 401–402 394
Canker (Valsa leucostoma (Pers.) Fr.) 55, 70, 151, Coleoptera 477–480
205, 295, 354, 375–378, 394 Colour, flesh 17–19, 21, 27, 30
see also Leucostoma canker see also Fruit, flesh
Canopy assimilation 247, 255 Colour, skin 12, 17–19, 537–538, 540–546, 551, 557,
Canopy shading 246 561–562
Carbamate insecticides 472, 476, 477, 498 anthocyanins 561
600 Index

Colour, skin continued named cultivars (partial list)

blush (overcolour) 29, 110, 118–119, 140, ‘Ailihong’ 54
168–169, 183, 186, 551, 582 ‘Armking’ 45, 278–280, 555
carotenes 561 ‘Babygold’ 44–45, 542
chlorophyll 561 ‘Bai Tao’ 42
CIE electronic colorimeter 551 ‘Baifeng-2’ 45
ground (undercolour) 551 ‘Baihua’ 44, 46
xanthophylls 561 ‘Baimangpantao’ 45
Comparative test for cultivar release 80 ‘Baixianglu’ 45
Conotrachelus nenuphar see Plum curculio ‘Baiyu Tao’ 42
Constriction canker (Phomopsis amygdali) 355, ‘Beinong Zaoyan’ 45
380–383 ‘Beinong-2’ 45
confused with brown rot twig blight 380 ‘Bositao’ 46
fusicoccin 381 ‘Chan Tao’ 42
Consumer acceptance 537–539, 543 ‘Chengxiang’ 45
Controlled atmosphere storage 551, 565 ‘Chenpupantao’ 45
high carbon dioxide (CO2) 566 ‘Chinese Cling’ 2, 69, 141–142, 168
acetaldehyde, ethanol 566 ‘chrysanthemum’ peach 55
low oxygen 565 ‘Chunhua’ 45
Cooling 551, 564–565 ‘Chunle’ 45
see also Temperature ‘Cullinan’ 45
Copper 318 ‘Datuanmilu’ 45
Cotyledon 98–99 ‘Dayu Tao’ 42
Covering materials and reflective films 48, 341, ‘Dong Tao’ 42
545 ‘Dunhuadongtao’ 46
Crop load 246 ‘Early red-2’ 46
interactions with stress 297–298 ‘Elberta’ 2, 46, 69, 108–109, 124, 141–144,
modifying 291–297 146, 154, 161, 166, 200, 223, 290,
quantifying 290–291 304, 451, 512
Cross-breeding 68 ‘Erse Tao’ 42
Crown gall ‘Fang Tao’ 42
Agrobacterium radiobacter 420, 423 ‘Feicheng Tao’ 44
Agrobacterium tumefaciens 99–100, 194, ‘Feichengtao’ 46
203–204, 206, 208, 409, 418, 510–511, ‘Fenghuapantao’ 45
515 ‘Fenzhou Tao’ 42
galls on peach seedlings 422 ‘Fertilia Morettini’ 45
infection 420, 423 ‘Fillips’ 46
T-DNA 420 ‘Flavorlate’ 45
management 423 ‘Guang Tao’ 42
Crown rot see Phytophthora root and crown rots ‘Guoyan red’ 42
Cull utilization 590–592 ‘Hakuho’ 44
other uses 592 ‘Hangshengpantao’ 45
potential uses 590–591 ‘Hanli Tao’ 42
cattle feed 590–591 ‘Hanlum’ 44
drying 591 ‘Hehuan Erse Tao’ 42
fuel alcohol 591 ‘Hongliguang’ 46
situation in California 591–592 ‘Hongrang Tao’ 42
Cultivar replacement time 53 ‘Huaguang’ 45
Cultivars ‘Huangroupantao’ 46
Chinese traditional cultivars and landraces ‘Huayumi’ 44
52 ‘Huiyulu’ 45
development 52 ‘Japan-89’ 45
early 45, 52 ‘J.H. Hale’ 14, 64, 69, 109, 141–142, 144,
flat peach and ornamental peach 45 146, 161, 164, 166, 179, 566
high-performance 45 ‘Jiaqingpantao’ 46
hypoallergenic 564 ‘Jin Cheng Tao’ 40
mid-season 45, 52 ‘Jin Tao’ 42
 Index 601

‘Jincheng’ 46 ‘Zaohongzhu’ 45
‘Jingmi’ 46 ‘Zaohuangpantao’ 45
‘Jingyu’ 46 ‘Zaokuimitao’ 45
‘Jinxiu’ 46 ‘Zaolupantao’ 45
‘Kunlun Tao’ 42 ‘Zaoshoumi’ 45
‘Kurakato’ 45 ‘Zaoxialu’ 45
‘Li Tao’ 42 ‘Zaoxiangyu’ 45
‘Lianhuang’ 45 ‘Zhaohui’ 44, 45
‘Lihepantao’ 46 ‘Zhaoxiang’ 45
‘Long 1-2-4’ 46 ‘Zhonghuashoutao’ 46
‘longevity’ peach 55, 56 ‘Zhongyoupantao’ 45
‘Longhuashuimi’ 46 ‘Zi Wen Tao’ 40
‘Luosi white’ 42 ‘Ziye DaTao’ 42
‘Mi Tao’ 42 ornamental peach 41, 45, 55
‘Myoujou’ 45 Cyanidin 562, 577
‘Nai Tao’ 40
‘NJC88’ 45
‘Okubo’ 44 Dagger nematodes (Xiphinema spp.) 196–198, 334,
‘Pan Tao’ 42 449–451
‘Pang Tao’ 42 biology 525
‘Putian Tao’ 42 control 526
‘Qi Di Tao’ 40 diagnosis 526
‘Qiangyefei’ 42 economic importance 525–526
‘Qianye Tao’ 42 geographical distribution 524–525
‘Qin Tao’ 40 host range 525
‘Qingzhoubaipimitao’ 46 identification 526
‘Redhaven’ 5, 45, 75, 81, 100, 142, species 524–525
144–145, 151–152, 163, 166, 198, survival 525
278, 510, 552 see also Nematodes; Xiphinema
‘Renmian Tao’ 42 Dandelion (Taraxicum officinale) 334, 341
‘Ruiguang-2’ 45 alternate host for Tomato ringspot virus 198,
‘Ruiguang-3’ 45 334, 449
‘Sahuahongpantao’ 45 alternate host for ring nematode 341
‘Shanghaishuimi’ 44 Decay see Deterioration problems
‘Shenzhou’ 46 Deer mouse (Peromyscus maniculatus) 340
‘Shenzhouhongmi’ 46 Degree day (°D) development models 472–473,
‘Shenzhoushuim’ 44 474, 475, 477
‘Shiyue Tao’ 42 Deterioration problems 577–580
‘Shouxing Tao-1’ 45 Botrytis cinerea (grey mould) 356, 396, 398,
‘Shuang Tao’ 40 401, 577
‘Shuguang’ 45 chilling injury (CI) see internal breakdown
‘Sunagawase’ 45 (below)
‘Wanshuomi’ 46 decay 538–542, 545, 577, 579, 584, 592
‘Wuyuexuanbiangang’ 45 defects 537, 542, 543
‘Xianghui-1’ 45 inking 580, 586
‘Xiao Tao’ 42 foliar nutrients 580
‘Xinbaihua’ 46 fungicides 580, 584, 588
‘Xizhuang-1’ 46 heavy metal contaminants 580
‘Yanguang’ 45 internal breakdown (IB) 537–538, 543–544,
‘Yanzhi Tao’ 42 565, 577–579, 582, 584, 590
‘Yexiandongtao’ 46 bleeding 578
‘Yin Tao’ 42 controlled atmosphere (CA) 578–579
‘Yinhualu’ 45 flesh translucency 577
‘You Tao’ 42 modified atmosphere packaging (MAP)
‘Yuhualu’ 44, 45 579, 592
‘Yulupantao’ 45 severity 578–579, 584
‘Zao Tao’ 42 susceptibility 578
602 Index

Deterioration problems continued analogue (propylene) 552, 556

internal breakdown (IB) continued autocatalytic action 552
taste 578 biosynthesis 551–552, 554
texture 577 1-aminocyclopropane-1-carboxylate
mechanical injury 41, 579–580 (ACC) 552, 565
bruising, abrasion 579 1-aminocyclopropane-1-carboxylate
bruising, compression 579 oxidase (ACO) 552–553
bruising, impact 579 1-aminocyclopropane-1-carboxylate
bruising, vibration 579 synthase (ACS) 552, 554
critical impact bruising thresholds 579 ACO isogenes (Pp-ACO1, Pp-ACO2) 554
plant growth regulators 579 ACS isogenes (Pp-ACS1, Pp-ACS2, Pp-
Monilinia fructicola see Brown rot ACS3) 552, 554
peach skin discoloration see inking (above) perception 551–552, 554
staining see inking (above) European red mite see Mites
Dichocrocis punctiferalis (Pyralid moth) 55 European stone fruit yellows (ESFY) 409–410,
Diplodina fruit rot (D. persicae) 401–402 428–431
Diptera 486–487 apple proliferation (AP) group 430
Disease resistance 127–129, 442 apricot chlorotic leaf roll (ACLR) 429
bacterial spot 143, 145–146, 153 group 16Sr X 430
brown rot 143, 163–164, 166–167 management 430
leaf curl 161–162, 164, 166–167 plant hosts 430
powdery mildew 161–162, 164, 166–167 plum leptonecrosis (PLN) 429
DNOC 294 psylla 430
Dodder (Cuscuta spp.) 409, 427 symptoms 429, 430
Domestication 38, 40, 43, 57 vector hosts 430
Dormancy 27, 30, 43, 49, 75, 78, 92, 97, 107, Evapotranspiration (ET) 228, 319–321, 542
121–122, 128, 223–224, 249, 251, 411, 416 Evergreen 97
Dormant oil 474, 493–494, 499 Evergrowing 97
Double petals 52 Expansins (Pp-Exp1, Pp-Exp2, Pp-Exp3) 557
Drought 2, 48, 97–98, 167, 194, 199, 206–209, Expressed sequence tag (EST) 93–96, 567
212–214, 245, 248, 295, 297, 298, 340, 508,
510, 514
tolerance 43 Feeding site 509, 517
see also Water stress Fibre 562
Dwarfism 4–5, 45, 48, 52–55, 62, 66, 70, 90, 100, Field harvesting, hauling and packaging 585–590
108, 140, 146, 151, 153, 161, 165–167, 199 fruit hauling 579, 580, 585–586
fruit packaging 580, 587–588, 590
harvesting 579–582, 585–586, 592
Emasculation 74, 76–77 shipping and transportation 590
Embryo 98–100 sorting and sizing operation 588–590
Embryo culture 25, 78–79, 98, 131, 155 ranch or field packed 588
definition 236 tray-packed 588–589, 593
methodology 236, 238 volume-fill 588
Endocarp 2, 15, 16, 19, 22, 24, 25, 30 Flavonols 577
Entomopathogenic nematode 521 Flower bud 3, 291–293
see also Heterorhabditis; Steinernema density 123, 270, 292
Epicatechin 577 differentiation 270
Eradication 442–443, 458 Flower
Ermine (Mustela erminea) 340 colours 10, 42, 52, 89
ESTs see Expressed sequence tags diversity 14
Ethephon 293, 296 double colours 42
Ethylene 22–24, 27, 30, 65, 127, 185, 293, 296, double flower 64
537–539, 551–558, 560–562, 565–567, 592 purple 42
1-methylcyclopropene (1-MCP) 555, 561–562 red 42
action inhibitor (2, 5-norbornadiene) 556 type 13, 64
aminoethoxyvinylglycine (AVG) 552, 554, white 42
560 Flowering, time of 122–123, 207, 408–409, 426
 Index 603

Food cell 517 cat-facing 481, 483

Fresh market, harvest and postharvest handling gumming 482
576–594 russetting 484
Frosty mildew (Leucotelium pruni-persicae) 401–402 silvering 484, 485
Fructose 247–251, 559–560 water soaking 482
Fruit 11 internal breakdown see Deterioration
colour 42 problems
see also Colour maturity see Ripening
composition 576 photosynthesis 246
aroma 576, 584 preharvest factors affecting peach quality
immature 576 536–547
mature 576 quality, definition 537
minerals 576 quality, factors affecting
organic acids 576, 592 canopy manipulation 544–547
titratable acidity (TA) 537–539, 543–545, canopy position 544, 545
576 cracking 544, 542
development 123–125, 551 crop load 544
fruit development period 564, 566 fruitlet thinning 544
stage I (cell division) 551, 561, 564 see also Thinning
stage II (pit hardening) 551, 557, 561, girdling 545–546
564 leaf removal 544–546
stage III (cell elongation/final swell) reflective materials 48, 341, 545
551, 557–558, 560–561, 564 summer pruning 544
stage IV (ripening) 551–552, 561 quality, genotype 539–540
flesh quality, irrigation 542–544
acids 17, 19, 21 calcium 541–542
colour 17–19, 21, 27, 30 iron 542
firmness (puncture pressure) 537–538, mineral nutrition 540–542
543, 551–553, 556, 581, 592 nitrogen 540–541
firmness, at retail stores 590, 592–594 partial root zone drying (PRD) 543
flavour 19 potassium 542
melting type 18, 21–24, 30, 42–46, 51, 53, regulated irrigation deficit (RID)
65, 67, 108, 110, 119–121, 127, 140, 542–543
151, 153–154, 176, 185–186, 538, sensory properties of quality
551–552, 555–556, 561, 563, 581 appearance 537, 540
non-melting type 2, 12, 18, 21–22, 24, 27, aroma 537–539
30, 41, 53, 65, 67, 71, 108, 110, flavour 537–540, 545
118–121, 126–127, 140, 153, 155, taste 537
163–167, 169, 176–177, 185–186, texture 537, 542
189, 539, 551–552, 555–556, 561, respiration 246, 252, 255–256
563 ripening
phenolics 19, 21 delaying 48
ready to buy 592 maturity definition 581
ready to eat 593 stage 578, 585, 592
slow-ripening mutant 552 stimulation 48
softening and melting 555 ripening, maturity indices 581
stony-hard (hd) 552, 554 background colour 581
see also Stony-hard field application 581–582
sugars 19–21, 27, 30 flesh colour 581
texture 16, 21–25, 27, 30 see also Fruit, flesh
growth 254–255 flesh firmness 581, 592
handling harvest date 581
fruit buyer handling 592–593 maximum maturity 581
fruit preparation for consumers 592 skin colour 581–582, 589
at retail distribution 592–594 ripening, maturity and quality 537–539,
hauling 579–580, 585–586 580–581
injury 468–486 ethylene 537–539
604
Fruit continued
ripening, maturity and quality continued
flavour 576, 578–581, 590
harvest 537–545
immature fruit 581
market life 537–538, 544–545, 580
mature 581, 592
non-destructive techniques 539
over-mature 581
quality potential 580
ripening 537–540, 544
storage potential 539, 543, 545
shape 15–16, 18, 25, 30, 42
size 15–16, 25, 29, 537, 540, 542–546
skin 2, 10, 12, 16, 17, 18, 19, 21, 25, 27, 30
storage
cold storage 51
controlled-atmosphere storage 51
low-pressure storage 51
low temperature storage 564–565
see also Postharvest; Temperature
Fungal gummosis (Botryosphaeria dothidea) 55, 355,
378–381
Fungicide resistance 360
Fusetto see Central-leader
Fusicladosporium carpophilum see Scab
Genetic diversity, assessment 52
Genetic map 95
Genetic resistance see Resistance
Genetic transformation see Transformation
Genome 85, 88, 91–92, 94, 96–97, 100, 215, 419, 446,
449–451, 453, 459, 556, 567
Genome Database for Rosaceae (GDR) 93, 97, 100,
256
Genotyping, high-throughput 55
Geotrichum candidum (sour rot) 356, 398, 400–401
Germplasm 44–45, 52, 68, 91, 99, 108, 110, 117–118,
120, 122–123, 131, 133–134, 143, 153,
163–168, 170, 181, 189, 213, 443, 459, 511,
515, 523
Giant cell 508, 509
Gibberellins 292–293
Gilbertella decay (G. persicaria) 356, 396, 398–399
Girdling 41, 48, 545–546
Glucose 247–251, 559–560
Graft
compatibility 194–199, 202–205, 207–213
incompatibility 194, 199, 206–207, 211
Grafting techniques 43
see also Propagation, clonal
Grapholita molesta 468–472, 475
see also Oriental fruit moth
Green June beetle (GJB), Cotinis nitida 480–481
Green peach aphid, Myzus persicae 204, 454
Greenhouse cultivation 48, 55
Index
management 49
microclimate 48
semi-ground 49
spacing 49
see also Protected cultivation
Grey mould (Botrytis cinerea) 356, 396, 398, 401, 577
Ground cover 339, 519
beneficial effects 334
fertilization 341–342
organic mulch 342
insect habitat 339–340
predacious mite, Neoseiulus fallacies 339
twospotted spider mite, Tetranychus
urticae 339
irrigation 341
killed sod 333
small mammals 340
types 340–341
chewing fescue (Festuca rubra) 340
common bermudagrass (Cynodon
dactylon) 333
fodder radish (Raphanus sativus v.
olieferus) 344
hard fescue (Festuca longifolia) 340
Kentucky bluegrass (Poa pratensis) 340,
346
perennial ryegrass (Lolium perenne) 340
Phacelia (Phacelia tenacetifolia) 344
subterranean clover (Trifolium
subterraneum) 340
tall fescue (Festuca arundinacea) 333
turnip (Brassica rapa v. rapa) 344
vetch (Vicia spp.) 340
white clover (Trifolium repens) 335, 342,
345
winter rye (Secale cereale) 344
Growth habit see Tree
Gummosis see Fungal gummosis
Haploid 66, 68–69, 74, 93, 96
Hardiness see Cold hardiness
Hardwood cuttings 203–204, 206–211, 213
see also Propagation, clonal
Harvesting 576–594
Heat requirements 26
Helicobasidium mompa (violet root rot) 401–402
Hemicellulose 562
Hemiptera 480–484, 491–496
Herbicides 296, 335–337, 346
for organic production 337, 346
pre-emergent types 335–336
post-emergent types 335–336
residue carryover 337
resistance development 335
Heritability 3–4, 21, 27, 66, 68, 72, 122–123, 125,
132, 162, 179, 181, 187
 Index 605

Heterorhabditis bacteriophora 521 Irrigation 318–321

Higher-density orchard 53 Isozyme 86
High-throughput genotyping 55
History, in China 38–40
Eastern and Western Han Dynasty Jacket rot (Botrytis cinerea, Sclerotinia sclerotiorum)
ErYa 40 354, 360–362, 396
HanShu 40 Japanese beetle (JB), Popillia japonica 479–480
from Wei–Jin Dynasty to Sui–Tang Dynasty Juice 44, 51, 175–176, 187–188, 557, 559, 565,
and Five Dynasties period 591–592
Eastern Jin period 40
GuangZhi 40
KuaiYuanTianBaoYiShi 41 KAC-V see Training system; Y-shaped tree
Northern Wei period 40 Kaempferol 562
QiMinYaoShu 40, 41 Kaolin particle films 479
Tang Dynasty 41
TangShu 41
TaoFu 41 LAI 268
Wei–Jin to Sui–Tang 41 Late season cultivars 46, 52
Pre-Qin Dynasty 38, 40 Layering 41, 206, 208, 210, 237
GuanZi 40 Leaf 6
HanFeiZhi 40 age 245
LüShiChunQiu 39 area index 247
ShiJing 38, 39 mass area 267
YeZhongJi 40 nitrogen content 267
Hormones 252 specific weight 247
HuangHe, Yellow River 40 water potential 245
Hydrolases Leaf curl (Taphrina deformans (Berk) Tul.) 2, 9, 55,
β-galactosidase (β-GAL) 555 70, 88–90, 161–162, 164, 166–167, 358,
β-(1, 4)-glucanase (syn. cellulase) (EG) 368–370
555–556, 566 resistance 161–162, 164, 166–167
pectinmethylesterase (PME) 555–556, 565 Leaf:fruit ratio 254, 290–291
polygalacturonase (PG) 555–556, 565 Leaf-footed bugs (Hemiptera: Coreidae) 481–483
Hypoallergenic cultivars 564 Leaf moth (Thosea sinensis) 55
Leaf rust (Tranzchelia discolour) 127–128
Lepidoptera 468–477, 487–491
Ilarviruses 443–447 Leptoglosus phyllopus 481–483
In vitro culture 98–99 Leucostoma (Cytospora or Valsa) canker (Leucostoma
see also Propagation, clonal cincta) 205, 214, 354, 375–378
Inbreeding 16, 73, 110 trunk painting 378
Inking see Deterioration problems see also Peach tree short life
Insecticides, organophosphate 472–474, 476, 479, Light
490–491, 494 distribution 268
Insects 55 interception 247, 269–270
growth regulator (IGR) 468, 473, 477, 494 penetration 268
scale see Armoured scale Linkage 61, 67–68
see also named insect species and groups Linkage groups 62–65, 86–87, 89–92
Integrated pest management (IPM) 472–473, 484, Linkage map 86
496 Lipid-transfer proteins (LTPs) 563–564
see also Ground cover Low acid 21, 23
Intercropping systems 49 see also Fruit, flesh; Organic acids
International Rosaceae Genome Consortium Low-chill 48, 196
(IRGC) 93 genetic resources 52
Intraspecific cross 72, 86 Low-chill cultivars 107, 108, 109, 110, 112, 113,
Iron 316–317 114–117, 118, 119, 120, 121, 122, 123, 124,
chlorosis 204, 207, 212–213 125, 127, 128, 130, 131, 133
deficiency symptoms and correction definition 107, 121
316–317 Low-density orchards see Orchard
606 Index

Lygus spp., plant bugs 334 economic importance 517

Lygus elisus 334, 481 host range 518
Lygus hesperus 334, 481 M. curvatum 515
Lygus lineolarius (tarnished plant bug) 341, M. rusticum 515, 521
481–482, 484 M. xenoplax (ring nematode) 196–197,
Lygus rugulipennis 481 207–208, 210–211, 341, 394, 512–513,
515–521, 523
survival 517
Magnesium 311 symptoms 515
Manganese 229, 317–318 see also Ring nematode
Marker-assisted selection 22, 55, 88, 162, 179, 212, Methyl bromide 222, 338, 392, 395, 510, 517, 519,
514 526
Mating distruption (pheromone) 129, 468–469, MIA see Y-shaped tree
472–473, 475, 488, 491 Microclimate see Greenhouse cultivation
Maturity see Fruit ripening Micrografting 236, 446
Meadow-orchard 278–279 see also Propagation, clonal
density 279 Micropropagation see Propagation, clonal
spacing 278 Mites
see also Orchard European red mite (ERM), Panonychus ulmi
Mealiness (chilling injury) see Deterioration 496–499
problems; Internal breakdown (IB) Pacific spider mite, Tetranychus pacificus 497
Mechanical injury see Deterioration problems Peach bud mite, Eriophyes insidiosus 456
Mediterranean fruit fly, Ceratitis capitata 486–487 Predacious mite, Neoseiulus fallacies 339
sterile male release 486 Silver mite 254, 498
Meloidogyne spp. 506–515 Two-spotted spider mite (TSM), Tetranychus
biological cycle 508–509 urticae 339, 496–499
biology 508–509 Model 247, 250–251, 254, 256–257
control 510–515 Modified atmosphere packaging (MAP) 565, 579,
dissemination 509 592
economic importance 510 Molecular marker 61, 86, 93, 96, 179
environmental factors 510 Mollicutes 407
host range 510 Monilinia spp. see Brown rot
identification 507 Monoploid see Haploid
M. arenaria 506, 512–514 Mouse-ear 43
M. floridensis 196–197, 203–204, 207–210, Mucor decay (Mucor piriformis) 356, 396, 398
212–215, 507, 511–514 Mutagenesis 72
M. hapla 196–197, 203–204, 207–210, 212–215, Mycelium 353, 355, 361, 364, 367–368, 371, 387,
506 391, 393, 396, 398
M. hispanica 506, 513 Mycoplasma-like organisms 239, 407
M. incognita 196–197, 203–204, 207–210, characteristics 408–409, 410
212–215, 506, 512–514 classification 409–410
M. javanica 196–197, 203–204, 207–210, detection 409
212–215, 506, 512–514 general introduction 407–410
M. mayaguensis 513 geographical distribution 410
polymorphism 507 hosts 410
survival 509 management 409
symptoms 507–508 myricetin 562
see also Root-knot nematode phytoplasma environment 407
Melting 18, 21, 22, 23, 24, 30, 42, 45, 51, 53 vectors 407, 410
see also Fruit, flesh dodder (Cuscuta spp.) 409, 427
Mendelian inheritance, traits 3–4, 7, 24, 29, 62, 71, leafhoppers 409
74, 80, 86, 93 psyllids 409
Mesocriconema spp. 506, 521 periwinkle 409, 411, 427
biology 516 Myrobalan plum (Prunus cerasifera) 194–198, 200,
control 518–517 202, 210, 212–213
disease complexes 515, 517–518 resistance gene 513–515
dissemination 517 resistance to Meloidogyne spp. 513–515
 Index 607

NAA 296 floor vegetation

Nematicide treatment threshold 518, 520, 523 cultural dwarfing 343
Nematodes 196, 479, 488, 506–527 establishment 342–344
dagger 524–525 influences 346–347
ectoparasitic 515, 524 soil temperature 346
endoparasitic 506, 521 management systems 342–346
entomopathogenic 521 permanent vegetation management by
identification mowing 345
Meloidogyne 507 vegetation-free tree rows in summer
Xiphinema 526 with control of cover crops 344
peach root-knot 196–197, 203–204, 207–210, vegetation-free tree rows with vegetated
212–215, 507, 511–514 drive alleys 342
plant parasitic 506–527 vegetation-free tree rows year-round
ring 196–197, 207–208, 210–211, 515–521, 345
523 year-round vegetation-free 342
root-knot 88, 196–197, 203–204, 207–210, Organic acids 550, 557
212–215, 506–515 citrate 557
root-lesion 521–523 high-acid fruit types 557, 559
resistance 201–202 low-acid fruit types (sub-acid) 21, 557, 559
see also Heterorhabditis; Steinernema malate 557
Neochlorogenic acid 562 phosphoenolcarboxylase (PEPC) 557–558
Nepoviruses 198, 448–451, 457, 524–526 Oriental fruit moth (Grapholita molesta) 468–472,
Nimblewill 335, 340, 519 475
Nitrogen 267, 305–308, 540–541 distinction from codling moth 470
deficiency 306, 308 Origin 1–2, 37–38, 576
excess 306 Gansu 37, 38, 45, 48
fertilization efficiency 307–308 Hemudu village 37
foliar applications 306–307 Neolithic 37
Non-melting see Fruit, flesh Taixi village of Gaochen city 37
Nutrient sampling 304–305 Tibet 37, 38
deviation from optimum percentage (DOP) Osmotic adjustment 248, 322–323
305
diagnostic and recommendation integrated
systems (DRIS) 305 Packaging, modified atmosphere (MAP) 565, 579,
flowers 304 592
leaves 304 Palmette 272–274
Nutritional value 562–563 density 273
allergenicity 563–564, 567 hedgerow 273
hypoallergenic cultivars 564 photograph 274
lipid-transfer proteins (LTPs) 563–564 pruning 273
peeling 564 training 273
antioxidant activity 562 yield 273
carotenoids 562 see also Training system
β-carotene 562 Particle bombardment (biolistics) 99–100
β-cryptoxanthin 562 Peach asteroid spot virus 456, 458
vitamin A 562 Peach bud mite (Eriophyes insidiosus) 456
fibre 562 Peach dapple (PD) 454–455
phenolics 562 Hop stunt viroid (HSVd) 454–455
cyanidin, chlorogenic acid, Peach latent mosaic disease (PLM) 451–454
neochlorogenic acid 562 Peach latent mosaic viroid (PLMVd) 451,
quercetin, kaempferol, myricetin 562 453–454
Peach leaf curl (Taphrina deformans) 2, 9, 55, 70,
88–90, 161–162, 164, 166–167, 354, 358,
Open centre see Training system; Vase 368–370
Orchard resistance 161–162, 164, 166–167
design 265, 267, 270 Peach mosaic virus (PcMV) 455–456
floor management 196, 332–351, 484–485 Cherry mottle leaf virus (CMLV) 455
608 Index

Peach rosette (PR) 410, 427–428 Pheromones 129, 468, 469, 472–473, 474, 475, 488,
group 16Sr III 428 491
management 428 monitoring 473, 475, 478–480, 494
peach yellows strain 428 see also Mating disruption
X-disease group 428 Phony (phoney) peach disease 408–409, 423
Peach rosette mosaic virus (PRMV) 451, 524–526 dwarfing symptoms 423
Peach sooty ringspot virus 456–457 hosts, reservoir and secondary spread 424
Peachtree borers (Synanthedon spp.) vectors 424
Lesser peachtree borer (LPTB), Synanthedon Phosphoenolpyruvate carboxylase 247
pictipes 487–491 Phosphorus 308
Peachtree borer (PTB), Synanthedon exitiosa deficiency 308
487–491 excess 308
Wasp mimic 488 Photoperiod 233, 247–248
Peach tree short life (PTSL) 197, 203–206, 210–211, Photosynthesis 17, 244–248, 254–255, 267, 281, 290,
355, 393–395, 515, 517–520, 523 296–297, 316–318, 321
‘Guardian’ rootstock 395 C3 267
Pseudomonas syringae 394 CER 267
ring nematode 394–395 whole-canopy 281
Peach tree short life complex 411 Photosynthetic assimilate 290, 295–298
bacterial fluorescence 415 photosynthetically active radiation (PAR)
bacterial toxins 415 245
management 416 Phymatotrichum root rot (Phymatotrichopsis
persicomycin 415 omnivora) 401–402
syringomycin 415 Physical map 91, 93–94, 97, 100
Pseudomonas syringae pv. persicae 409, 414 Phytophthora root and crown rots 198, 203, 205,
Pseudomonas syringae pv. syringae 204, 394, 207–210, 212, 319, 353, 355, 383–387, 403
409, 413–414, 515 chlamydospores 353, 355–356, 383, 385, 387
Peach twig borer (Anarsia lineatella) 473–475 oospores 353, 383, 385, 387
Peach X-disease 410, 424–426 P. cactorum 355, 383
group 16Sr III 426 P. cambivora 355, 383
leafhoppers 426 P. cinnamomi 355, 383
little peach 426 P. citricola 355, 383
management 426 P. citrophthora 355, 383
plant hosts 424 P. cryptogea 355, 383
plant reservoirs 426 P. drechsleri 355, 383
red suture 426 P. megasperma 355, 383
rosette 426 P. parasitica 356, 383
symptoms 424, 425 P. syringae 356, 383
Peach yellow leaf roll (PYLR) 410, 428 zoospore 353, 355–356, 383, 385, 387
apple proliferation (AP) group 428–429 Phytoplasmas and phytoplasma diseases 80, 407,
group 16Sr X 428 424–429
pear decline (PD) 428, 429 Pip fruit 44
psylla 429 Pit see Endocarp
Peach yellows 410, 426–427 Plant bugs (Hemiptera: Miridae) 480
group 16Sr III 427 tarnished plant bug (TPB), Lygus lineolarius
little peach 426 481–482, 483
management 427 other species, L. hesperus, L. elisus, L.
plant hosts 426 rugulipennis 481
plum leafhopper 427 Plantation distance 78
red suture 426, 427 see also Orchard
Pectin 21–22, 24, 186, 555, 562–563, 565 Planting density and yield 281–282
Pest management see Integrated pest management see also Orchard
Phenolics 19, 21, 127, 186–187, 225, 233, 237, 545, Platforms 272–274, 284
550, 561–562, 576–577 Platynota idaeusalis (Tufted apple bud moth)
Phenology 29 475–477
phenological classification 30 Plum, rootstock 512–514
phenological phases 27 see also Rootstocks
 Index 609

Plum curculio (PC) (Conotrachelus nenuphar) P. vulnus 513, 521–523

477–479 P. zeae 521
Plum pox virus (PPV/Sharka) 2, 80, 88, 96, 162, survival 522
437–443, 447, 487, 494–495 symptoms 521
Potyvirus 440, 442 see also Nematodes
PPV-D 441 PRMV see Peach rosette mosaic virus
PPV-M 441 Processing peach 117, 119, 175–192, 200, 205, 223,
PPV-EA 441 255–256, 469
PPV-C 441 blanching 186–187
PPV-W 441 breeding 178, 180, 185
PPV-Rec 441 canning 184–185, 187
Plum rust (Tranzschelia pruni-spinosae) 209 endopolygalacturonase 186
see also Rust flavour 176
Podosphaera spp., powdery mildew flesh texture 177
P. clandestina 354, 369 freezing 187
P. leucotricha 354, 369 fruit quality 182
P. pannosa 209, 254, 354, 368–369, 371–374 colour 182–183
Pollen and pollination 13–14, 44, 64, 66, 68, 74, 77, firmness 183, 185–186
122, 129, 131, 141, 310, 314, 409, 436, 447, flesh browning 187
454, 458 pit quality 184
Polymerase chain reaction (PCR) 24, 100, 389, 409, soluble solids 187
419, 424, 428, 441, 447, 449–450, 453, 507, germplasm 178–179, 189
526, 554, 556, 560 grading 182
cooperational-PCR (Co-PCR) 441 harvest 176–177
heminested-PCR 441 marker-assisted selection 179
nested-PCR 441 marketing 188
PCR-ELISA 447 maturity index 184
real-time-PCR (RT-PCR) 441, 447, 455, 554, nectarine 188
556, 560 orchard life 178
reverse transcription and PCR 453 pasteurization 188
squash capture-PCR 441 peeling 186
Polyphenolics 21, 550 phytonutrients 183, 188, 189
Polyploidization 72, 507 postharvest 182
Pomology 68–69, 163, 165 processed products
Postharvest handling 49–52, 98, 181–182, 398–401, beer 51
540, 551, 576–594 drinks 51
Potassium 229, 308–310, 542 fruit jelly 51
deficiency 309, 310 peach candies 51
Powdery mildew 209, 254, 354, 368–369, uniform ripening 178
371–374 varieties 178–179
resistance 161–162, 164, 166–167 ripening times 179
see also Podosphaera spp. yield 178
PPV see Plum pox virus (PPV/Sharka) case yield 188
Pratylenchus spp. (root-lesion nematodes) 196–197, heritability 179
202–205, 207–208, 212–214, 506, 512 Production
biology 521–522 in China 46–47
control 523 systems 53
disease complexes 523 see also Orchard
dissemination 522 top five producing countries 46
economic importance 522 Progeny 3–5, 17, 66, 72–73, 80, 86, 91, 97, 122, 145,
host range 523 162, 166, 180–182, 185, 189, 212, 508
P. brachyurus 521 Prokaryotes 407–434
P. convallariae 521 see also Bacteria; Phytoplasmas
P. neglectus 521 Propagation, clonal
P. penetrans 513, 521–523, 525 apical meristem culture 447
P. pratensis 521 budding 237, 239
P. sefaensis 521 chip budding 238, 239, 449
610
Propagation, clonal continued
hardwood cutting 203–204, 206–211, 213, 224
auxin treatment 225
bottom heat 226
collecting time 225
collapse 226
definition 224
indole-3-butyric acid 225
mother plants 224
rootstocks 226
transplanting 226
washing 226
graft take 238
herbaceous grafting 240
in vitro propagation (micropropagation) 55,
98–99, 206, 208–210, 215, 230–236,
458
application of mycorrhizal fungi 236
explants sterilization 231
material preparation 230
plantlets acclimatization 233
problems during in vitro culture 235
problems during plantlet acclimatization
235–236
shoot proliferation 232
shoot proliferation, culture medium
232, 234
shoot proliferation, cytokinins 232
shoot rooting 233
shoot rooting, auxins 233
June budding 239
micrografting 239, 446
see also Micrografting
semi-hardwood cutting 227
acclimatization 229
auxin leaf spraying 228
auxin treatment 228
basal cutting 229
grafting techniques 43
mist propagation 227
perlite 228
soluble auxins treatment 228
time for budding 229
softwood cuttings 195, 208
vegetative bud 239
Propagation, seed
nursery site 221
peach seedling propagation 223
Prunus sylvestris 223
seed dormancy 223
seed supply 223
soil type 221
Protected cultivation 48, 52, 58, 59, 279–280
see also Greenhouse cultivation
Pruning 41, 55, 70, 151, 177, 199, 251–253, 265–269,
271–274, 276, 278–283, 285, 293–294,
296–297, 306, 321–322, 343, 346, 354–355,
Index
365, 374–375, 378–379, 382, 395, 400, 416,
424, 454–455, 458, 474, 544
heading cuts 273
roots 48
summer 48, 49, 55, 266, 272–274, 279–280,
283, 424, 544
thinning cuts 273
‘tutta cima’ 265
Pruning systems 55
see also Training system
Prunus spp. 2, 194, 442, 456, 459
P. americana 195–196
P. amygdalus 100
P. angustifolia (Chickasaw plum) 195, 428, 478
P. armeniaca 86, 442
P. avium 100, 442
P. belsiana 512
P. besseyi 195, 208, 442
P. cerasifera (Myrobalan plum) 86, 194–198,
200, 202, 210, 212–213, 430, 442,
512–514
P. cerasus 86, 442
P. cistena 442
P. davidiana 2, 40, 85, 87, 143, 162, 194–195,
200, 202, 204, 207, 209–210, 212–213,
443, 511
P. domestica 86, 194–196, 200, 202, 206,
210–211, 430, 442, 513
P. dulcis 2, 85, 185, 194–195, 200, 202, 442, 512
P. emarginata 426
P. ferganensis 3, 85, 87, 89, 161, 194
P. fremonti 523
P. hortulana 195, 428
P. humilis Bunge 53
Chinese dwarf cherry 54
P. incana 196, 202
P. insititia 194–195, 200, 202, 206, 211, 213,
442, 513
P. japonica 195, 199, 215
P. kansuensis 3, 38, 45, 85, 194
‘GansuTao-1’ 45
P. mariana 442
P. mira 3, 38, 85, 194
P. munsoniana 428, 513
P. persica 1, 40, 97, 193–198, 200, 202, 204,
207–210, 212–213, 215, 256, 442, 451,
506, 512, 514, 576
P. polytricha Koehne 53
dense-pubescent cherry 53
P. pseudocerasus 100
P. pumila 195, 199, 202, 207
P. salicina 86, 194–195, 197, 200, 202, 208,
210–211, 410, 416, 442–443, 513
P. serrulata 410, 443
P. spinosa 194–195, 200, 206, 210–211, 430,
442, 513
P. subcordata 195
 Index 611

P. subhirtella autumno rosa 100 Root-knot nematodes (Meloidogyne spp.) 55, 63,
P. sylvestris 223 88, 196–197, 202–205, 207–210, 212–215,
P. tomentosa 195–196, 199, 202, 215, 256, 442, 334, 506–512, 514–515, 517–519, 521
523 resistance 45, 88–89
P. triloba 442 see also Meloidogyne spp.
P. virginiana 410, 424, 426 Root-lesion nematodes (Pratylenchus vulnus,
P. vulgaris 200 Pratylenchus penetrans) 196–197, 202–205,
Pseudomonas syringae pv. syringae 515–516 207–208, 212–214, 512, 521, 523
see also Bacterial canker; Peach tree short life see also Nematode
Pseudosclerotia 353, 396 Rootstock 2–3, 10–11, 48, 53–55, 88, 93, 99, 117, 120,
PTSL see Peach tree short life 128–131, 140, 145, 150, 158, 163, 166–167,
Pure line 73–74 193–215, 222–226, 229–230, 232–235,
Pyralid moth (Dichocrocis punctiferalis) 55 237–240, 246, 254, 256, 265, 267, 270,
Pyrethroid insecticides 472, 476, 477, 479, 496 272–273, 278, 283–284, 317, 355–356, 385,
392–397, 412, 416, 423–424, 430, 442, 444,
495, 506–507, 509–512, 514–515, 518–520,
QTL see Quantitative trait locus 522–524, 526–527, 536, 539, 564
Quantitative character 66 ‘Adafuel’ 195, 201, 209, 226, 512
Quantitative trait locus (QTL) 86, 88–90, 92–93 ‘Adarcias’ 195, 199, 201, 209
Quarantine 198, 230, 403, 408–409, 431, 442–443, ‘Adesoto’ 101 195–197, 199, 202, 206, 226
449, 451, 456, 458, 527, 567 ‘Alcaniz’ 512
Quarantine pests 472, 480, 486–487 ‘B-S6’ 512
Quercetin 233, 562 ‘Bailey’ 89, 99, 143, 197, 199–201, 203, 205,
214, 223, 523, 552
‘Barrier 1’ 195–197, 199, 202, 210
Raffinose 250 ‘Bergasa’ 512
Raised bed 333–334, 338, 355, 385, 392 ‘Bokhara’ 512
Randomly amplified polymorphic DNA (RAPD) ‘Brompton’ 225, 513
86 ‘Cachirulo’ 512
RAPD see Randomly amplified polymorphic DNA ‘Cadaman’ 195, 202, 209–210, 512, 522–523
Reflective films 48, 341, 545 ‘Castore’ 201, 208–209
Regeneration 98–100, 278, 333, 443 ‘Citation’ 214, 226, 512
Replant 518, 522–523 ‘Controller 5’ 199, 202, 208
see also Orchard ‘Damas’ 63, 194, 206, 210, 237, 513
Resistance to nematodes dwarfing 48, 194–195, 199–200, 208–209,
Meloidogyne spp. 63, 511–515 211–212, 214–215, 256, 267
Mesocriconema spp. 519 see also Dwarfism
Pratylenchus spp. 523 ‘Elberta’ 200, 512
Xiphinema spp. 526 ‘Felinem’ 195, 197, 201, 214, 512
Resistance gene 96, 162, 212, 460, 508, 513–515 ‘Fermoselle’ 512
gene analogue (RGA) 96 ‘Flordaguard’ 10, 99, 116, 128, 196–197, 214,
management 472–473, 477, 496–497 511–513, 523
Respiration, climacteric 550–554, 557–558, 560, 567 ‘Garnem’ 195, 197, 201, 214, 512, 522
Restriction fragment length polymorphism (RFLP) ‘GF 31’ 513
86, 88, 91–94, 524 ‘GF 43’ 206–207, 225, 234, 513
RFLP see Restriction fragment length ‘GF 305’ 197–198, 201, 203, 210, 438, 442,
polymorphism 448–450, 452–453, 512
Rhizopus rot (Rhizopus stolonifer) 396, 398–399, 401 ‘GF 557’ 208, 512, 514
Ribulose bisphosphate carboxylase (RUBISCO) ‘GF 655/2’ 206, 211, 225, 237
245 ‘GF 677’ 195, 198, 201–204, 207–210, 214,
Ring nematode 196–197, 202, 204–205, 207–208, 225–226, 233–234, 267, 278, 317, 430,
210–211, 341, 378, 394–395, 512–513, 512
515–521, 523 ‘GF 1869’ 194, 206, 210, 225, 513
see also Mesocriconema spp. ‘GF 8-1’ 513
RNAse protection assay (RPA) 556 ‘Guardian’ 197–198, 200–201, 203, 205, 214,
Root pruning 48 223, 395, 509, 511–512, 518–523
Root rots see Phytophthora root and crown rots ‘Halford’ 178–179, 200–201, 203, 205, 223
612
Rootstock continued
‘Hansen 2168’ 201, 209, 214, 226, 232,
511–512
‘Hansen 536’ 201, 209, 214, 226, 232, 511–512
‘Harrow Blood’ 196, 214, 223, 512
‘Ishtara’ 197–199, 202, 210–211
‘Julior’ 195–196, 199, 202, 211
‘Krymsk 1’ 196–199, 202, 214
‘Krymsk 2’ 196–197, 199, 202, 214
‘Krymsk 86’ 196–197, 201, 214
‘Lovell’ 87, 93, 96, 142, 146, 197–198, 200–203,
205, 207, 223, 510, 512, 518–520,
522–523
‘Marianna 2624’ 194, 513
‘Missour’ 200, 512
‘Montclar’ 198, 201, 203–204, 223, 430, 512,
523
‘Montizo’ 195–196, 202, 513
‘Mr.S. 2/5’ 195–196, 198–199, 201, 210,
233–234, 239
‘Mr.S. 2/8’ 195, 198, 210
‘Myrabi’ 512
‘Myran’ 197–199, 202, 210, 226
‘Myrobalan P-2175’ 512
‘Myrobalan P-2980’ 513
‘Myrobalan 29C’ 513
‘Myrobalan 605’ 513
‘Myrobalan B’ 513
‘Myrobalan franc’ 513
‘Nemaguard’ 99, 145, 150, 197, 200–205,
207, 210–211, 214, 223, 234, 254, 507,
511–512, 514–515, 519–520, 522–523
‘Nemared’ 87–94, 96–97, 99, 150, 197, 200,
203, 205, 214, 223, 509, 511–512, 514,
522
‘Nickels’ 201, 214
‘Okinawa’ 110, 122–123, 128, 131, 196–197,
214, 228–229, 507, 509, 511–512
‘P-2032’ 514
‘P-2175’ 512–514
‘P-2980’ 513–514
‘P.S.B2’ 197, 199, 201, 203–204, 223, 225–226,
232
‘PSM 101’ 513
‘Penta’ 196–197, 201, 207, 226
plum 512–514
‘Polluce’ 201, 208–209
‘Pumiselect’ 195, 199, 202, 207
‘Rancho Resistant’ 512
‘Redglow’ 513, 523
resistance to nematodes see Resistance
‘Rubira’ 11, 162, 197, 199, 201, 203–204, 223,
430, 512, 514
‘Rutgers Red Leaf’ 430, 512, 523
‘Saint Julien 655-2’ 206, 513
‘S-37’ 214, 510–512
‘Shalil’ 197, 214, 512, 514
Index
‘Siberian C’ 166, 196, 200–201, 205, 214, 223,
512
‘Sirio’ 195, 199, 201, 208
‘Tetra’ 196–197, 199, 201, 207, 226
‘Torinel’ 513
‘Titan’ 512
‘Viking’ 202, 208, 214
‘Yunnan’ 197, 210, 214, 512
Rose chafers, Mycrodactylus subspinosus 479
Rosellinia root rot (Rosellinia necatrix) 401–402
Rust (Tranzschelia discolor, Tranzschelia pruni-
spinosae) 209, 354, 365–368
Sanitary state 80, 221, 240, 431, 451, 591
Sanitation 129, 354–357, 365, 368, 374, 379, 382,
400, 402, 409, 443, 452, 506, 577
Scab (Fusicladosporium carpophilum) 354, 363–365,
383, 416
confusion of symptoms with bacterial leaf
spot (Xanthomonas arboricola)
364–365
Scarab beetles (Coleoptera: Scarabaeidae) 479–481,
498
green June beetle (GJB), Cotinis nitida 480–481
Japanese beetle (JB), Popillia japonica 479–480
rose chafers, Mycrodactylus subspinosus 479
Sclerotium stem rot (S. rolfsii) 401–402
Seed 2, 3, 15, 16, 25, 41, 66, 68–69, 75, 78, 94, 97,
99–100, 108, 122, 129–131, 133, 140–142,
144, 146, 150, 155, 163, 167, 195, 200,
204–205, 212, 222–224, 236, 254, 333,
335–338, 342, 344, 409, 423, 447, 450, 454,
561
Selection 14, 19, 22, 27, 40–41, 44–45, 54–55, 57, 66,
72, 74, 78, 80, 88, 92, 99, 113, 119–123, 126,
129, 131–134, 145, 141, 153, 162–164, 167,
175–176, 178–179, 181–182, 184, 194–195,
199, 205–207, 212–214, 254
Selection index 80
Self-compatible 86
Self-pollination 4–5, 16, 21, 25, 68, 72–75, 78, 163,
165, 182
Semi-ground greenhouses 49
Sequence tagged site (STS) 97
Sharka see Plum pox virus
Shot hole
Corineum beijerinckii 209–210
Wilsonomyces carpophilus 354, 361–364, 403
Silver leaf disease (Chondrostereum purpureum) 206,
354, 373–376
Silver mite 254, 498
Simple sequence repeat marker (SSR) 86, 88, 91
Single trait 61
Mendelian trait 62–65
see also Qualitative character
Sink demand 246
 Index 613

Sink strength 246 Tarnished plant bug (TPB), Lygus lineolarius 339,
Site selection 355–356, 395, 408–409, 506 341, 481–482, 483
Slender spindle see Central leader; Training Taphrina deformans see Leaf curl
system Temperature, affecting fruit 245, 252, 255, 357, 361,
Softwood cuttings 195, 208–209 363–364, 368, 371, 374, 381, 392, 396, 398,
see also Propagation, clonal 400, 564–565
Soil solarization 222, 333, 393, 521, 527 chilling injury see Deterioration problems;
Soluble solids concentration (SSC) 19–21, 198–199, Internal breakdown (IB)
209–210, 249, 538–545, 551, 559–560, 564, ideal storage conditions 582, 584
566, 576, 581–582 low temperature storage 564–565
Somatic embryogenesis 98, 240 new temperature management approach
Sorbitol 20, 187, 247–251, 255–256, 314–315, 322, 584–585
559–560 fresh-cut 592
Sour pit (Candida inconspicua/Torulopsis forced-air cooling 582–583
inconsipicua) 401–402 hydro-cooling 400, 582–583
Sour rot (Geotrichum candidum) 356, 398, 400–401 preconditioning 579, 584–585, 594
Sphaerotheca pannosa see Podosphera pannosa ready to buy 584, 585, 592, 594
Stachyose 250 ready to eat 585, 592–594
Starch 15, 237, 247–251, 290–291, 555, 576 water loss control 576, 579, 581, 584, 590
Steinernema glaseri 521 fruit waxes or coatings 584, 587–588,
Steinernema riobrave 521 590
Sterile male 486–487 Thermotherapy 447, 452, 454
see also Mediterranean fruit fly Thinning 15, 25, 123, 142, 177, 179, 182, 199,
Stink bugs (Hemiptera: Pentatomidae) 334–335, 252–253, 255, 273, 282, 285, 289–299, 357,
339, 341, 469, 480–484 368, 544
brown, Euschistus servus 481–482 by hand 289, 293–296
Euschistus tristigmus 481 Thosea sinensis (Leaf moth) 55
green, Acrosternum hilare 481 Thrips (Thysanoptera) 17, 341, 447, 468, 484–485
southern green, Nezara viridula 481 other species, T. meridionalis, T. major 484
Stomatal conductance 245–246, 321–322 Western flower thrips, Frankliniella occidentalis
Stone see Endocarp 484–486
Stony-hard 18, 22, 23, 24, 30, 31, 32, 65, 127, 140, Tobacco ringspot virus (TRSV) 524–525
146, 151, 165, 185, 552, 554 Tomato ringspot virus (ToRSV) 198, 334, 447–449,
see also Fruit, flesh 524–526
Stoolbed and layering 208, 237 Prunus stem pitting disease 198, 448–449, 525
see also Propagation, clonal ToRSV see Tomato ringspot virus
Stooling 207–208 Torulopsis inconspicua (sour pit) 401–402
see also Propagation, clonal Training system 264–284
Strawberry latent ringspot virus (SLRV) 449–451, ‘axis central’ see Central leader
524–526 central leader 55, 274–276
Suckering 202, 204–207, 210–211, 213, 394, 412, delayed vase 203, 269, 271, 273
414, 444, 516 free shape 273
Sucrose 20, 90, 187, 234–237, 247–251, 255–256, ‘forma libera’ 272
322, 416, 559–561, 576 Fs-Tatura 278
SUGAR model 250, 559–560 ‘fusetto’ see Central leader
Sulfur 311–312, 561 hedgerow see Palmette
Summer pruning 48–49, 55, 266, 272–274, 279, 280, KAC-V see Y-shaped tree
283, 424, 544, 582 meadow-orchard 278–279
Summer tipping 49 MIA see Y-shaped tree
Surfactant 294 open centre see Vase
Sweet kernels 52, 65, 67, 88, 90 palmette 269, 272–274
Sweetness (sugars) 88, 187, 255, 537, 559–560 PCR pruning 49
Synteny 88, 92, 96 slender spindle see Central leader
V-form 271, 468, 474
vase 55, 271–272, 468, 474
Taraxicum officinale see Dandelion ‘vaso californiano’ 271
Target leaf spot (Phyllosticta persicae) 401–402 ‘vaso de plataformas’ 271
614 Index

Training system continued herbicides 335–337, 346

‘vaso italiano’ 271
Y-shaped tree 55, 269, 276–278, 474
Y-trellis see Y-shaped tree
Transformation 55, 97–101, 240, 443, 459
transgenic 99
Tranzchelia discolor (leaf rust) 127–128
Tranzschelia pruni-spinosae (plum rust) 209, 354,
365–368
Tree wall systems, pillar 70
Triploid 68
TRSV see Tobacco ringspot virus
Trunk borer (Aromia bungii) 55
Tufted apple bud moth (TABM/Platynota
idaeusalis) 475–477
Two-spotted spider mite see Mites
mechanical cultivation 334, 337
johnsongrass, Sorghum halepense 333
mulching 337–339
Nicotiana spp. 451
pest habitat 334
poison ivy, Toxicodenron radicans 333
preplant control 333
sheep sorrel, Rumex acetosella 449
Virginia creeper, Parthenocissus quinquefolia
333
yellow nutsedge, Cyperus esculentus 333
Winter chilling see Chilling requirement
Wilthin 294
Woolliness (chilling injury) see Deterioration
problems; Internal breakdown (IB)

Unique expressed gene (unigene) 94, 96, 567 Xiphinema spp. 196–198, 506, 524–527
biology 525
control 526
Vase see Training system diagnosis 526
Verticillium wilt (Verticillium dahliae) 203–204, economic importance 525–526
208–209, 356, 392–394, 403 geographical distribution 524–525
microsclerotia 356, 392 host range 525
Violet root rot (Helicobasidium mompa) 401–402 identification 526
Viruses 80 species 524–525
indexing 200, 436, 442, 449–451, 455–456, survival 525
459 X. americanum group 196–198, 449,
see also Nepoviruses 524–526
Vole, montane (Microtus montanus) 340 sensu lato 524–525
sensu stricto 524–525
X. brevicolle 524
Water stress 228, 236, 245, 253, 297–298, 319, X. bricolensis 524–525
321–323, 542–543 X. californicum 449, 524–525
beneficial effects 321–322 X. coxi 450
managing 322–323 X. diversicaudatum 450, 524–526
negative effects 321 X. pachtaicum 524
regulated deficit irrigation (RDI) 322–323 X. pacificum 524
see also Drought X. rivesi 449, 524–525
Waterlogging 43, 194–196, 202–212, 319, 333, X. vuittenezi 524, 526
513–514 see also Nematodes
Weeds
alternate host for virus 334
Chenopodium spp. 449, 451 Yield 3, 25, 43, 46–47, 51, 53, 55, 74, 80, 118, 128,
chickweed, Stellaria spp. 334 164, 176–179, 181–182, 188–189, 193, 196,
host for stink bugs, plant bugs, cat- 198–200, 203–204, 208–211, 214, 254–256,
facing injury to fruit 334, 339, 269–274, 276, 278–285, 291–292, 295–297,
341 304–306, 308, 311, 313, 317, 321–322,
dandelion, Taraxicum officinale 334, 341, 333–335, 339, 341, 343–344, 346–347, 379,
451 384, 424, 444, 468–469, 494, 496, 506, 508,
alternate host for Tomato ringspot 510, 517, 522, 537, 540, 542–544, 550–551,
virus 334 567, 590
alternate host for ring nematode 341 average peach and nectarine 47
economic thresholds 335 planting density 281–282
flame weeding 337 training systems 280–283
 Index 615

Y-shaped tree 276–278 Y-trellis see Training system

density 277, 278
light interception 277
photograph 277 Zinc 312–314
protected cultivation 278
spacings 278
see also Training system