Table Of Content14
Monographs on
Theoretical and Applied Genetics
Edited by
R. Frankel (Coordinating Editor), Bet-Dagan
M. Grossman, Urbana . H. F. Linskens, Nijmegen
P. Maliga, Piscataway . R. Riley, London
Monographs on Theoretical and Applied Genetics
Volume 1 Meiotic Configurations
A Source of Information for Estimating Genetic Parameters
By J. Sybenga (1975)
Volume 2 Pollination Mechanisms, Reproduction and Plant Breeding
By R. Frankel and E. Galun (1977)
Volume 3 Incompatibility in Angiosperms
By D. de Nettancourt (1977)
Volume 4 Gene Interactions in Development
By L. I. Korochkin (1981)
Volume 5 The Molecular Theory of Radiation Biology
By K.H. Chadwick and H.P. Leenhouts (1981)
Volume 6 Heterosis
Reappraisal of Theory and Practice
Editor: R. Frankel (1983)
Volume 7 Induced Mutations in Plant Breeding
By W. Gottschalk and G. Wolff (1983)
Volume 8 Protoplast Fusion
Genetic Engineering in Higher Plants
By Y. Y. Gleba and K. M. Sytnik (1984)
Volume 9 Petunia
Editor: K. C. Sink (1984)
Volume 10 Male Sterility in Higher Plants
By M. L. H. Kaul (1988)
Volume 11 'fiee Breeding: Principles and Strategies
By G. Namkoong, H. C. Kang, and J. S. Brouard (1988)
Volume 12 The Wheat Rusts - Breeding for Resistance
By D. R. Knott (1989)
Volume 13 Population Dynamics and the liibolism Model
Genetics and Demography
By R. F. Costantino and R. A. Desharnais (1991)
Volume 14 Genetic Improvement of Tomato
Editor: G. Kalloo (1991)
G. Kalloo (Ed.)
Genetic Improvement
of Tomato
With 26 Figures
Springer-Verlag
Berlin Heidelberg New York
London Paris Tokyo
Hong Kong Barcelona
Budapest
Prof. Dr. G. KALLOO
Department of Vegetable Crops
Haryana Agricultural University
Hisar 125 004
Haryana
India
ISBN-13:978-3-642-84277-1 e-ISBN-13:978-3-642-84275-7
001: 10.1007/978-3-642-84275-7
Library of Congress Cataloging-in-Publication Data. Genetic improvement of tomato / Kalloo, editor. p. cm.
- (Monographs on theoretical and applied genetics; 14). Includes bibliographical references and index.
ISBN-13:978-3-642-84277-1
1. 1bmatoes - Breeding. 2. Tomatoes - Genetics. 3. Tomatoes - Biotechnology. I. Kalloo, Dr. II. Series.
SB349.G42 1991 635'.64223-dc20 91-11760 CIP
This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned,
specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on
microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is
permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version,
and permission for use must always be obtained from Springer-Verlag. Violations are liable for prosecution
under the German Copyright Law.
© Springer-Verlag Berlin Heidelberg 1991
Softcover reprint of the hardcover 1st edition 1991
The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even
in the absence of a specific statement, that such names are exempt from the relevant protective laws and regula
tions and therefore free for general use.
Reproduction of the figures: Gustav Dreher GmbH, Stuttgart
'!Ypesetting: International 'JYpesetters Inc., Makati, Philippines
3113145-543210 - Printed on acid-free paper
Preface
The esculent Lycopersicon esculentum, long thought
to be poisonous, has become a major U.S. food
crop and source of vitamins and minerals, thanks
largely to genetic modification and new production
technology
Rick (1978)
Tomato (Lycopersicon esculentum Mill.) is one of the most important solana
ceous vegetable crops grown worldwide under outdoor and indoor conditions.
It has become an important commercial crop so far as the area, production,
industrial values and its contribution to human nutrition is concerned. During
the past few decades tremendous developments have contributed to the
knowledge and understanding of various areas of genetics, breeding and
biotechnology and voluminous literature has been generated. The purpose of
preparing this monograph is to give a comprehensive up-to-date treatment to
the various aspects of genetic improvement of tomato. The emphasis has been
placed on cytology, classical and molecular genetics, reproductive biology,
germplasm resources, hybrid seed production, use of wild taxa, selection/
breeding methods, breeding for abiotic and biotic stresses, processing and
quality breeding, improvement for mechanical harvesting, and biotechnology:
tissue culture, protoplast fusion, and genetic transformation. These topics are
presented in 22 different chapters. However, a few aspects have been discussed
in more than one chapter. For example, seed production is treated in chapters
1, 4 and 8; molecular biology/genetic engineering in chapters 3 and 22 and
heterosis in chapters 8 and 16. Breeding for resistance to diseases, insects,
nematodes and environmental stresses has been presented in chapters 3, 16, 18
and 22 besides in their respective chapters. However, detail information regard
ing treatment of various topics can be consulted with the help of subject index.
The chapters have been prepared by specialists from different countries with
the support of relevant and current literature.
It is hoped that the monograph will serve the need of students, teachers and
researchers working on the subject or related areas.
I am thankful to various authors for contributing their chapters and Prof.
Rafael Frankel for valuable suggestions to improve the text.
Hisar, India, March 1991 G. KALLOo
Contents
Chapter 1 Introduction
G. KALWO
1.1 Origin and History ..................................... .
1.2 Taxonomy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.3 Economic Importance .................................... 2
1.4 Area and Production .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.5 History of Improvement .................................. 3
1.6 An Ideal Research Material ............................... 5
1. 7 Seed Production ......................................... 8
Chapter 2 Cytology of Lycopersicon
G. A. KIRILWVA
2.1 Introduction 11
2.2 Mitosis 11
2.3 Meiosis ................................................ . 12
Chapter 3 Genetics of Lycopersicon
J. W. DEVERNA and A. H. PATERSON
3.1 Tomato as a Research Vehicle ............................. 21
3.2 Variability. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
3.3 Mutants and Genetic Stocks .............................. 27
3.4 Gene Linkage ........................................... 31
3.5 Genetics of Quantitative Traits ............................ 35
Chapter 4 Reproductive Biology in Tomato
M.L.H. KAUL
4.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
4.2 Floral Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
4.3 Incompatibility. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
4.4 Artificial Hybridization .................................. 42
4.5 Genetic Male Sterility .................................... 43
VIII Contents
Chapter 5 Germplasm Resources in Lycopersicon
I. B. GARANKO
5.1 Introduction... . ......................... ....... ... ... .. 51
5.2 Collection and Exploration ............................... 51
5.3 Evaluation and Utilization ................................ 54
5.4 Maintenance and Conservation ............................ 56
5.5 Descriptor. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
5.6 Extinction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Chapter 6 Selection Systems for Tomato Improvement
L.C. PEIRCE
6.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
6.2 Improving Pedigree-Based Selection Systems ................ 60
6.3 Population Improvement Systems .......................... 61
6.4 Selection for Combining Ability ........................... 64
6.5 Gametophytic Selection .................................. 65
6.6 Selecting Superior Parents ................................ 65
6.7 Factors Affecting Selection and Genetic Advance ............ 66
Chapter 7 Interspecific and Intergeneric Hybridization in Tomato
G. KALLOO
7.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
7.2 Crossability. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
7.3 Hybrids and Segregating Progenies ........................ 75
7.4 Sesquidiploids. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
7.5 Barriers. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
7.6 Overcoming the Barriers .................................. 79
7.7 Transfer of Economic Attributes from Wild Species
to Commercial Cultivars .................................. 80
7.8 Breeding Strategies and Limitations ........................ 81
Chapter 8 Heterosis in Tomato Breeding
H. GEORGIEV
8.1 Introduction 83
8.2 Meaning of Heterosis ................................... . 83
8.3 The Causes of Heterosis ................................. . 84
8.4 Breeding of F 1 Tomato Hybrids .......................... . 85
8.5 Development of Hybrid Seed Production .................. . 91
8.6 Summary .............................................. . 98
Contents IX
Chapter 9 Disease Resistance in Tomato
A. N. LUKYANENKO
9.1 Introduction 99
9.2 Fungal Diseases ........................................ . 99
9.3 Bacterial Diseases ...................................... . 110
9.4 Virus Diseases ......................................... . 115
9.5 Conclusions ............................................ . 119
Chapter 10 Insect and Mite Resistance in Tomato
R.R. FARRAR Jr. and G.G. KENNEDY
10.1 Introduction ............................................ 121
10.2 Tomato Species in Which Resistance to Insects
or Mites Is Known . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
10.3 Insects and Mites to Which Resistance Is Known
in Tomato Species ....................................... 132
10.4 Known Mechanisms of Resistance ......................... 134
10.5 Conclusions............................................. 142
Chapter 11 Root-Knot Resistance in Tomato
G. F ASSULIOTIS
11.1 Introduction ............................................ 143
11.2 Distribution............................................. 143
11.3 Pathology .............................................. 143
11.4 Biology and Life History ................................. 144
11.5 Nematode Control ....................................... 145
11.6 Breeding for Resistance ................................... 145
11.7 Screening for Root-Knot Resistance ........................ 146
11.8 Genetics of Resistance. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 146
11.9 Stability of Resistance .................................... 147
11.10 Mechanism of Resistance ................................. 149
11.11 Relationship of Root-Knot Nematodes
to Other Tomato Root Diseases ........................... 151
11.12 Summary............................................... 152
Chapter 12 Breeding for Environmental Stress Resistance in Tomato
G. KALLOO
12.1 Introduction ............................................ 153
12.2 Cold Resistance ......................................... 153
12.3 Heat Resistance ......................................... 156
12.4 Drought Resistance ...................................... 159
x Contents
12.5 Excessive Moisture Resistance ............................. 161
12.6 Salt Resistance .......................................... 161
12.7 Efficient Mineral Nutrition Uptake ........................ 163
12.8 Resistance to Herbicides .................................. 164
12.9 Ozone Resistance ........................................ 165
Chapter 13 Parthenocarpy in Tomato
A. N. LUKYANENKO
13.1 Introduction ............................................ 167
13.2 'JYpes of Parthenocarpy .................................. 167
13.3 Sources ................................................ 168
13.4 Expression.............................................. 169
13.5 Genetics................................................ 171
13.6 Physiology.............................................. 173
13.7 Quality of the Parthenocarpic Fruits ....................... 174
13.8 Methods of Screening for Parthenocarpy ................... 175
13.9 Use of Parthenocarpy in Tomato Breeding .................. 176
Chapter 14 Breeding for Carotenoid Pigments in Tomato
M. CHALUKOVA and H. MANUELYAN
14.1 Introduction ............................................ 179
14.2 Regulation of Carotenogenesis ............................ 180
14.3 Function of Carotenoids ................... . . . . . .. . . . .. . . . 185
14.4 Characteristics of Carotenogenesis in Interspecific and
Intercultivar Hybridization of Tomato ...................... 186
14.5 Express Methods for Assessing the Carotenoid Composition
of Tomato Fruits ........................................ 193
Chapter 15 Breeding Tomato for Quality and Processing Attributes.
S. Z. BERRY and M. R. UDDIN
15.1 Introduction ............................................ 197
15.2 Fruit Quality. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
15.3 Breeding Methods ....................................... 199
15.4 Testing and Evaluation of Breeding Lines. . . . . . . . . .. . . . . . . .. 201
15.5 General Breeding Objectives .............................. 201
15.6 Specific Breeding Objectives .............................. 202
15.7 Future Prospect ......................................... 205
Contents XI
Chapter 16 Some Aspects of Breeding Tomato
for Greenhouse Conditions
S.1. IGNATOVA
16.1 Introduction ............................................ 207
16.2 Development of Cultivars and Hybrids ..................... 207
Chapter 17 Breeding Tomato for Mechanized Harvesting
A.N. LUKYANENKO
17.1 Introduction 213
17.2 Requirements of Tomato Cultivars for Mechanized Harvesting 213
17.3 Breeding for Uniform Ripening of Fruits .................. . 215
17.4 Breeding Tomatoes for Enhanced Fruit Firmness and
Marketable Yield ....................................... . 221
17.5 Tomato Breeding for Pedicel Characteristics ................ . 226
17.6 Conclusions ............................................ . 229
Chapter 18 Cell, Tissue and Organ Culture in Lycopersicon
H. KOBLITZ
18.1 Introduction ............................................ 231
18.2 Callus and Cell Suspension Cultures ....................... 231
18.3 Root, Shoot and Plant Regeneration from Vegetative Material. 237
18.4 Culture of Reproductive Organs (Anthers, Flowers, Ovaries,
Embryos) ............................................... 241
Chapter 19 Protoplast Culture and Somatic Hybridization
in Lycopersicon
H. KOBLITZ
19.1 Introduction ............................................ 247
19.2 Isolation and Culture of Protoplasts ....................... 247
19.3 Regeneration of Shoots and Plantlets
from Isolated Protoplasts ................................. 250
19.4 Protoplast Fusion and Production of Somatic Hybrids ....... 253
Chapter 20 Genetic Variation in in-Vitro Cultures
and Regenerated Plants in Tomato and Its Implications
K. SREE RAMULU
20.1 Introduction ............................................ 259
20.2 In Vitro Differentiation and Nuclear Processes .............. 260
20.3 Phenotypic and Genetic Variation in Regenerated Plants ...... 261
