Monographs on 14 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

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© 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 regula­tions 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 solana­ceous 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 regard­ing 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 2.2 2.3

Introduction Mitosis Meiosis ................................................ .

Chapter 3 Genetics of Lycopersicon J. W. DEVERNA and A. H. PATERSON

11 11 12

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 8.2 8.3 8.4 8.5 8.6

Introduction Meaning of Heterosis ................................... . The Causes of Heterosis ................................. . Breeding of F 1 Tomato Hybrids .......................... . Development of Hybrid Seed Production .................. . Summary .............................................. .

83 83 84 85 91 98

Contents

Chapter 9 Disease Resistance in Tomato A. N. LUKYANENKO

9.1 9.2 9.3 9.4 9.5

Introduction Fungal Diseases ........................................ . Bacterial Diseases ...................................... . Virus Diseases ......................................... . Conclusions ............................................ .

Chapter 10 Insect and Mite Resistance in Tomato R.R. FARRAR Jr. and G.G. KENNEDY

IX

99 99

110 115 119

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

Chapter 16 Some Aspects of Breeding Tomato for Greenhouse Conditions S.1. IGNATOVA

XI

16.1 Introduction ............................................ 207 16.2 Development of Cultivars and Hybrids ..................... 207

Chapter 17 Breeding Tomato for Mechanized Harvesting A.N. LUKYANENKO

17.1 17.2 17.3 17.4

Introduction Requirements of Tomato Cultivars for Mechanized Harvesting Breeding for Uniform Ripening of Fruits .................. . Breeding Tomatoes for Enhanced Fruit Firmness and Marketable Yield ....................................... .

17.5 Tomato Breeding for Pedicel Characteristics ................ . 17.6 Conclusions ............................................ .

Chapter 18 Cell, Tissue and Organ Culture in Lycopersicon H. KOBLITZ

213 213 215

221 226 229

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

XII Contents

20.4 Variants or Mutants from Cell and Callus Cultures .......... 267 20.5 Factors Influencing Variation .. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 268 20.6 Conclusions............................................. 269

Chapter 21 Isozymes in Lycopersicon N. V. KUDRYAKOVA

21.1 Introduction ............................................ 277 21.2 Variations in Lycopersicon species ......................... 277 21.3 Applications in Breeding Practices ......................... 279

Chapter 22 Genetic Transformation of Tomato and Prospects for Gene Transfer J. HILLE, P. ZABEL, and M. KOORNNEEF

22.1 Introduction ............................................ 283 22.2 The Ti-Plasmid as Natural Gene Vector .................... 284 22.3 Agrobacterium-Mediated Plant Cell Transformation .......... 285 22.4 Direct Gene Transfer ..................................... 286 22.5 Application of Genetic 'fransformation in Lycopersicon ...... 287 22.6 Concluding Remarks ..................................... 291

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 293

Subject Index. . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 353

Contributors

You will find the addresses at the beginning of the respective contribution

Berry, S. Z. 197 Chalukova, M. 179 DeVerna, J. W. 21 Farrar, R. R. Jr. 121 Fassuliotis, G. 143 Garanko, I. B. 51 Georgiev, H. 83 Hille, J. 283 Ignatova, S. I. 207 Kalloo, G. 1, 73, 153 Kaul, M. L. H. 39 Kennedy, G. G. 121

Kirillova, G. A. 11 Koblitz, H. 231, 247 Koornneef, M. 283 Kudryakova, N. V. 277 Lukyanenko, A. N. 99, 167, 213 Manuelyan, H. 179 Paterson, A. H. 21 Peirce, L. C. 59 Ramulu, Sree K. 259 Uddin, M. R. 197 Zabel, P. 283