13
WHEAT BREEDING Its scientific basis
WHEAT BREEDING Its scientific basis
SERIES EDITOR
E. H. Roberts Professor of Crop Production, Department of Agriculture and Horticulture, University of Reading
WHEAT BREEDING Its scientific hasis
Edited by
F. G. H. Lupton
SPRINGER-SCIENCE+BUSINESS MEDIA, B.V.
First published in 1987 by Chapman and Hali Ltd
© 1987 Springer Science+Business Media Dordrecht Originally published by Chapman & Hall Ltd in 1987
Softcover reprint of the hardcover 1st edition 1987
Ali rights reserved. No part of this book may be reprinted, or reproduced or utilized in any form
or by any electronic, mechanical or other means, now known or hereafter invented, including
photocopying and recording, or in any information storage and retrieval system, without permission in
writing from the publisher.
British Library Cataloguing in Publication Data
Wheat breeding. 1. Wheat-Breeding 1. Lupton, F. G. H. 633.1'13 SB19l.W5
Library of Congress Cataloging in Publication Data
Wheat breeding.
Includes bibliographies and index. 1. Wheat-Breeding. 2. Wheat. I. Lupton,
F. G. H. (Francis Geoffrey Hugh), 1921-SB19l.W5W494 1987 633.1'13 87-6599
ISBN 978-94-010-7908-2 ISBN 978-94-009-3131-2 (eBook) DOI 10.1007/978-94-009-3131-2
Contents
Preface xi
Contributors xii
Foreword by Sir Ralph Riley xiii
1 Systematics and evolution 1
T. E. Miller 1.1 Introduction 1
1.2 Classification of the Triticeae 1
1.3 The goat grasses, Aegilops L. 4
1.4 The ryes, Secale L. 14
1.5 The wheats, Triticum L. 15
References 28
2 The history of wheat cultivation 31
G. D. H. Bell 2.1 The earliest evidence: archaeology and palaeoethnobotany:
neolithicagriculture 31
2.2 The spread from the Fertile Crescent: Europe 34
2. 3 Production in historic times 37
2.4 The20thcentury 45
References 47
3 History of wheat breeding 51
F. G. H. Lupton 3.1 Wheat improvement in the 19th century 51
3.2 The development of modern plant breeding 52
3. 3 Wheat breeding in France and Belgium 53
3.4 Wheat breeding in Germany 56
3.5 Wheat breeding in the Netherlands 58
3.6 Swedish wheat breeding 59
3. 7 Hard red spring-wheat breeding in North America 61
VI
3.8 Wheat breeding in Australia 3. 9 Wheat breeding in Britain 3.10 Wheat breeding in Italy 3.11 Wheat breeding in Poland and the USSR 3.12 History of semi-dwarf wheat development
References
Contents
63 64 66 66 68 70
4 Aneuploidy in wheat and its uses in genetic analysis 71 C. N. Law, J. W. Snape and A. J. Worland 4.1 Introduction 71 4.2 The aneuploids of Chinese Spring 72 4.3 Monosomics and their use 74 4.4 The development of inter-varietal chromosome substitutions 88 4.5 Chromosome assays 92 4.6 Analysis of whole chromosomes 95 4. 7 Chromosome substitutions and prediction 100 4.8 Location of geneticfactors 101 4.9 The use of wheat aneuploids in the future 106
References 107
5 Conventional methods of genetic analysis in wheat 109 J. W.Snape 5.1 Introduction 109 5.2 Analysis of major gene variation 111 5.3 Genetic analysis of quantitative characters 113 5.4 Examining relationships between different characters 122 5. 5 Genetic analysis and wheat breeding 125
References 127
6 Wheat genetics 129 A. J. Worland, M. D. Gale and C. N. Law 6.1 Introduction 129 6.2 Morphological characters 130 6.3 Pigment characters 132 6.4 Adaptive characters 133 6.5 Agronomic and grain characters 136 6.6 Crossability 141 6. 7 Disease resistance 142 6.8 Geneticdisorders 145 6.9 Cytoplasmic inheritance 149 6.10 Biochemical and molecular markers 153 6.11 Chromosome maps and gene locations 158 6.12 Wheat genetics in the future 166
References 166
Contents Vll
7 The introduction of alien genetic variation into wheat 173 M.D. Gale and T. E. Miller 7.1 Introduction 173 7.2 Crossability within the Triticeae 174 7.3 Synthetic amphiploids 176 7.4 Wheat-alien chromosome addition lines 178 7.5 Wheat-alien chromosome substitution lines 184 7.6 The expression and transfer of alien genes to wheat 188 7.7 Conclusions and prospects 204
References 206
8 Chromosome structure and organization 211 R. B. Flavell, M. D. Bennett, A. G. Seal and J. Hutchinson 8.1 Introduction 211 8.2 The nuclear genome 211 8.3 The chloroplast genome 251 8.4 The mitochondrial genome 259 8.5 Concluding remarks 261
References 262
9 Triticale breeding 269 R. S. Gregory 9.1 Introduction 269 9.2 Breeding methods 271 9.3 Yield 273 9.4 Other physiological characters 276 9.5 Grain shrivelling 278 9.6 Disease resistance 278 9.7 Quality and utilization 281 9.8 Present production and future prospects 283
References 284
10 Development and structure of the wheat plant 287 E. J. M. Kirby and M. Appleyard 10.1 Introduction 287 10.2 Seedling development and emergence 287 10.3 Shoot apex development 291 10.4 Leaf development 300 10.5 Tillers 303 10.6 Stem growth 306 10.7 Ear development 307 10.8 Conclusion 310
References 310
Vlll Contents
11 Water relations 313 P. Innes and S. A. Quarrie 11.1 Introduction 313 11.2 Categories and critical periods of water shortage 314 11.3 Breeding for drought resistance 315 11.4 Effects of morphological and physiological characters on
yield and water economy 316 11.5 Effects of metabolic characters on water relations 325 11.6 Conclusions 333
References 333
12 Yield assessment 339 B. Westcott 12.1 Introduction 339 12.2 Designs for wheat variety trials 339 12.3 Factorial trials 348 12.4 Practical considerations 349 12.5 Genotype-environment interaction in wheat 352 12.6 The agricultural significance of wheat yield assessment 362
References 366
13 Breeding for disease resistance 369 R. Johnson and F. G. H. Lupton 13.1 Introduction 369 13.2 Pathogen variation 370 13.3 Sources of resistance 381 13.4 Durability of resistance 382 13.5 Disease escape 383 13.6 Practical aspects of breeding 387 13.7 Resistance to rust diseases 390 13.8 Resistance to other biotrophic pathogens 403 13.9 Resistance to necrotrophic pathogens 410 13.10 Conclusions 418
References 418
14 Breeding for resistance to insects 425 H.J. B. Lowe 14.1 Introduction 425 14.2 Terminology 425 14.3 Examples of breeding for resistance to insect pests of wheat 428 14.4 Otherpestsofwheat 439 14.5 Specificity and resistance 443 14.6 Practical aspects of selection for pest resistance 446
Contents ix
14.7 Resistance and pest control 448 14.8 Conclusion 449
References 449
15 Grain quality 455 J. A. Blackman and P. I. Payne 15.1 Introduction 455 15.2 The quality characters of wheat 455 15.3 Food products 459 15.4 Breeding for the quality character 465
References 483
16 Production of new varieties: an integrated research approacit to plant breeding 487 J. Bingham and F. G. H. Lupton 16.1 Introduction 487 16.2 Determination of breeding objectives 488 16.3 Breeding pure line varieties 494 16.4 Exploitation of male sterility 514 16.5 Selection for grain yield 519 16.6 Case histories of Plant Breeding Institute varieties 523 16.7 Impact of plant breeders' rights 528 16.8 Acquisition and maintenance of collections 530 16.9 Discussion and conclusions 532
References 536
17 Future prospects 539 P.R. Day and F. G. H. Lupton 17.1 Introduction 539 17.2 Improvement in physiological characters 539 17.3 Resistance to disease 542 17.4 Breeding techniques 543 17.5 Grain quality 545 17.6 Exploitation of alien genetic variation 546 17.7 Utilization of crop residues 547 17.8 The impact of biotechnology 548 17.9 Conclusion 551
References 551
Index 553
Preface
To be most effective, the work of the plant breeder requires the synthesis of knowledge in a wide range of subjects in the field of genetics and agricultural botany. During the 1970s and early 1980s, the Plant Breeding Institute at Cambridge provided a unique opportunity for collaborative work between plant breeders and those concerned with ancillary subjects likely to be of interest to them. The breeders had ready communication with specialists in plant physiology, plant pathology, entomology, cytogenetics and molecular biology, with whom they could discuss their work and from whom they could ask for help and guidance in dealing with specific problems, while departments concerned with chemistry and statistics were available to help in the planning and analysis of their experiments. At the same time the specialists had access to the most advanced breeding material for their studies, and could ensure that these studies were directed to problems of real importance to the breeders in the short or longer term.
It now seems likely that many of the specialist departments of which the Institute was formed may, for political or economic reasons, be dispersed or disbanded. This book offers a survey of the many aspects of wheat research carried out at the Plant Breeding Institute, with chapters written by specialists in each of the principal fields of work studied there. Although much of the work considered has not been comprehensively reviewed before, the book is primarily intended to give an overall view of wheat breeding research for students or specialists in other fields. Each subject is considered on an international basis, but discussion, particularly in the chapters dealing with the more practical aspects of breeding, is intentionally biased towards problems arising in Western Europe, as it is felt that these will be of most interest to the general reader, who can look to books published in other countries for reviews of their problems. No attempt has been made to give detailed accounts of aspects of wheat research not considered at the Institute; in such cases the reader is referred, where possible, to reviews in the literature.
Much of this book has been written during times of great uncertainty at the Plant Breeding Institute, and I would like to express my appreciation to all my colleagues for the friendly co-operation and support which I have received during its preparation. I hope that it will serve as a useful memorial to their years of co-operative work.
F. G. H. L.
Contributors
All contributors are present or former members of the staff of
The Plant Breeding Institute, Maris Lane, Trumpington, Cambridge CB2 2LQ
and may be contacted through this Institute or through the Editor,
Dr F. G. H. Lupton, I The Knapp, Haslingfield, Cambridge CB3 7JH.
Foreword
by Sir Ralph Riley, MA, DSc, PhD, FRS
Wheat is a major object of study both because of its primacy among all crops in feeding mankind and because of the intellectual challenge that it poses in a range of biological disciplines as well as in archaeology, and in social and economic history. In 1984, the last year for which world data are available, 512 million tonnes of wheat were produced compared with 464 million tonnes of rice. Not surprisingly, since wheat is well adapted to cultivation in temperate regions it is pre-eminently a crop of the developed world. In 1984, 60% of world wheat was produced in developed countries whereas only 5% of the rice came from the developed world.
A concentration of research on wheat was therefore in the self-interest of the developed world with its access to prolific resources and scientific manpower and its scientific traditions that led to the first agricultural revolution. The consequences of this concentration have been to enormously improve the efficiency of our capacity to produce wheat and have made wheat scientifically one of the best understood plant species and certainly the best understood polyploid, whether flowering plant, animal or lower organism. This understanding not only helps Man better to appreciate his place in the world but will provide that knowledge from which a second green revolution can be made when next the food: population equation gets out of balance. Then more of the world crop will have to be produced in the poorer countries. For the first green revolution, which saved Asia from hunger in the 1960s, was not built on new knowledge but on the adaptation of principles derived from earlier research. We must be sure that on the next occasion a similar store of knowledge is available especially about such crops as wheat.
A fine overview of the state of scientific work on wheat is provided by Wheat breeding. Its scient~fic basis. Under the guidance of Dr F. G. H. Lupton this book has brought together work from the informal 'Cambridge School of Wheat Scientists'. This school, which was never designated but was nevertheless an effective interactive grouping of scientists, came together at the Plant Breeding Institute over the thirty year period from the mid-1950s to the mid-80s. I am proud to have been a member of the school, for it has been unique both in the quality and breadth of its science and in its impact on practical
xiv Foreword
agriculture. Its membership has included practical wheat breeders and agronomists, pathologists, entomologists, geneticists and cytogeneticists, molecular and cell biologists, physiologists and protein chemists, and nowhere else in the world has such a broad-based assemblage of scientists been devoted to the study of a single crop. Over the years about 1000 scientists from most parts of the world have contributed to its work. The spread of scientific competence allowed for cross-feeding between disciplines and in particular demonstrated the ways in which applied and fundamental science can assist and stimulate each other. This ensured that those close to practice used new knowledge quickly and that those creating new knowledge were aware of practical problems.
This book may have to stand as a memorial to the 'Cambridge School of Wheat Scientists' for so long supported in an enlightened way by the British Agricultural Research Council (now the Agricultural and Food Research Council). This is because the United Kingdom Government has decided to divide the Institute, and so the School, into two-selling the applied activities to private enterprise while retaining in the public sector research that is not yet applied. In this casual way one of the great successes for all time in crop research could be destroyed. Whether either of the separated parts can long survive is unclear; certainly the strength of mutual support will be lost or will have to be rebuilt. As I write this, decisions on the implementation of the policy of splitting the 'school' are imminent. I hope that readers of this book will have evidence that scientifically aware authorities have recognized the strength of the 'Cambridge Wheat School' and the need for its continued coherence. I hope that they will not be reading a requiem for a school which was broken up because scientific enlightenment did not prevail.
Ralph Riley
Stapleford, Cambridge January, 1987
