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Progress in Plant Breeding—1 Edited by G.E. RuSSell, MA, PhD, ScD, DipAgricSci, FIBiol, FRES Emeritus Professor of Agricultural Biology, University of Newcastle upon Tyne Butterworths London Boston Durban Singapore Sydney Toronto Wellington All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, including photocopying and recording, without the written permission of the copyright holder, application for which should be addressed to the Publishers. Such written permission must also be obtained before any part of this publication is stored in a retrieval system of any nature. This book is sold subject to the Standard Conditions of Sale of Net Books and may not be re-sold in the UK below the net price given by the Publishers in their current price list. First published 1985 © Butterworth & Co. (Publishers) Ltd, 1985 British Library Cataloguing in Publication Data Progress in plant breeding.—1 1. Plant-breeding 631.5'3 SB123 ISBN 0-407-00780-6 Library of Congress Cataloging in Publication Data Main entry under title: Progress in plant breeding. Bibliography: v. 1, p. Includes index. 1. Plant-breeding—Collected works. 2. Food crops— Breeding—Collected works. I. Russell, G.E. (Gordon E.) SB123.P744 1984 633'.083 84-19961 ISBN 0-407-00780-6 (v. 1) Typeset by Scribe Design, Gillingham, Kent Printed in Great Britain at the University Press, Cambridge Preface Progress in the breeding of improved crop varieties has been particularly rapid during the past 15-20 years. Considerable advances, in terms of increased yielding ability, improved quality and agronomic characteristics, and better resistance to pests and diseases, have been made by the plant breeder and associated scientists and this has made a major contribution to crop productivity. Nevertheless, the need to develop even better varieties is greater than ever because of the ever-increasing world population. This series of volumes will review and critically assess progress in the major crops of the world, not only in terms of variety production but also across all the many associated disciplines. Plant breeding is such a complex, interdisciplinary subject involving, for example, genetics, plant physiology, plant pathology, applied zoology, biometrics and biochemistry, that it has become increasingly difficult for a specialist in one crop or discipline to keep abreast with developments in other fields. I hope that Progress in Plant Breeding will facilitate and improve communication between specialists in different fields. This volume contains 11 major review articles on a wide range of important plant-breeding topics. Several of the chapters review breeding progress in specific crops, for example in grain protein crops (Chapter 3), cassava (Chapter 4) and oil palm (Chapter 6); sometimes in certain geographical areas, for example in sugar beet in the USA (Chapter 2) and potatoes at OP (Chapter 5). Other chapters deal with certain breeding objectives in major crops, for example dwarfing genes in wheat (Chapter 1); drought response in small-grain cereals (Chapter 7); bird resistance in maize and sorghum (Chapter 8) and disease resistance in rice (Chapter 10); and genetic conservation in vegetable crops (Chapter 11). Certain themes run throughout the book: for example, the international nature of modern plant breeding is stressed repeatedly, reflecting the important role of the International Institutes in plant breeding. Another recurring subject is the importance of selecting for resistance to pests and diseases in most plant-breeding programmes; this topic is dealt with in detail particularly in Chapters 8 and 10, which illustrate quite different types of problems. The importance of plant physiology and of fundamental genetical studies is stressed in several chapters but particularly in Chapters 1 and 7 respectively. These recurring themes reinforce my view, held over a very long period, that experience with one crop or problem can sometimes be relevant, often to an unexpected extent, to an apparently dissimilar situation in a different crop. I hope therefore that readers who are interested mainly in a v vi Preface particular problem or a certain crop will not ignore chapters which do not appear immediately to be relevant to their present interests. If they do, they may save a little reading time but they may miss much that might be of interest and value to them in the future. Gordon E. Russell Contributors D. ASTLEY, National Vegetable Research Station, Wellesbourne, Warwick CV35 9EF, England R.W. BULLARD, US Fish and Wildlife Service, Denver Wildlife Research Center, Building 16, Federal Center, Denver, CO 80225, USA P. CRISP, National Vegetable Research Station, Wellesbourne, Warwick CV35 9EF, England M.D. GALE, Plant Breeding Institute, Maris Lane, Trumpington, Cambridge CB2 2LQ, England J.J. HARD ON, Directorate of Agricultural Research, Ministry of Agriculture and Fisheries, The Netherlands R.J. HECKER, US Department of Agriculture, Agricultural Research Service, Fort Collins, CO 80523, USA R.H. HELMERICK, Great Western Sugar Company, Longmont, CO 80501, USA C.H. HERSHEY, Centro Internationale de Agricultura Tropical, Apartado Aereo 6713, Cali, Colombia D.L. JENNINGS, Scottish Crop Research Institute, Invergowrie, Dundee DD2 5DA, Scotland GURDEV S. KHUSH, International Rice Research Institute (IRRI), Los Banos, Laguna, Philippines H.A. MENDOZA, International Potato Center (CIP), P.O. Box 5969, Lima, Peru N. RAJANAIDU, Palm Oil Research Institute of Malaysia, Malaysia V. RAO, Palm Oil Research Institute of Malaysia, Malaysia R.L. SAWYER, International Potato Center (CIP), P.O. Box 5969, Lima, Peru J.B. SMITHSON, Pulses Improvement Program, International Crops Research Institute for the Semi-Arid Tropics (ICRISAT), ICRISAT Center, Patancheru PO, Andhra Pradesh - 502 324, India R.E. SOJKA, Coastal Plains Soil and Water Conservation Research Center, US Department of Agriculture, Agricultural Research Service, Florence, SC 29502, USA vii viii Contributors S.S. VIRMANI, International Rice Research Institute (IRRI), Los Banos, Laguna, Philippines WATKIN WILLIAMS, Department of Agricultural Botany, Plant Science Laboratories, University of Reading, Whiteknights, Reading RG6 2AS, England J.O. YORK, Department of Agronomy, University of Arkansas, Fayetteville, Arkansas 72701, USA S. YOUSSEFIAN, Plant Breeding Institute, Maris Lane, Trumpington, Cambridge CB2 2LQ, England Chapter 1 Dwarfing genes in wheat M.D. Gale S. Youssefian Plant Breeding Institute, Mar is Lane, Trumpington, Cambridge CB2 2LQ, England Introduction Over the past two decades global wheat production has risen at an annual rate of 3.4% (Byerlee and Hesse de Polanco, 1983). Yield increases of this order have been achieved in both developing and developed nations, and are associated with the utilization of improved or more intensive husbandry methods. However, the more dramatic increases in production have, in most cases, become possible only with the concurrent widespread adoption of new semi-dwarf varieties. These varieties have stiff straw and are not prone to stem collapse or 'lodging' before harvest. Severe lodging can make harvesting impossible with modern agricultural equipment and even moderate lodging can result in a reduced final yield of grain with inferior processing quality. For example, the more humid environment of a lodged crop can often lead to increased preharvest sprouting, which can render the crop unsuitable for commercial bread or noodle production. Although several options are available for improving lodging resistance, the genetic strategy chosen by wheat breeders has generally been to reduce plant height. In the West, two important series of crosses have resulted in the world-wide use of probably four major dwarfing genes. These genes are now employed in over half the world wheat crop and their exploitation has been extremely rapid. For example, in Britain the first semi-dwarf variety, Fundin, was released in 1974 and by 1982 more than three-quarters of the acreage was sown to varieties carrying a Norin 10 dwarfing gene. The national average yields over the past 30 years are shown in Figure 1.1. As elsewhere, the large increases since the adoption of semi-dwarf varieties have been attributable both to genetic improvement and to more intensive crop husbandry, particularly increased fertilizer use which is permissible with these genotypes. The breeding and exploitation of semi-dwarf wheat varieties has proceeded with very little appreciation of the formal genetics of the genes controlling plant height or their effects on other plant characters besides straw length. However, over the past ten years a considerable amount of information regarding these genes has become available and the realization that only a few identifiable genes are responsible for the short stature of many of the world's wheat varieties has raised new genetic and agronomic questions. For example, which of the available dwarfing genes are intrinsically superior, which are better adapted to particular 1 2 Dwarfing genes in wheat 6.5 —1100 o^ Oto CDin CD "O CO Qj 2"S O ca 1950 1960 1970 1980 Year of harvest Figure 1.1 Wheat yields in England and Wales, the contribution of new varieties to production and the introduction of semi-dwarfs. The national spring and winter wheat yields and the effect of variety up to 1978 are five year moving averages from Silvey (1981). Yield figures from 1978 to 1983 were obtained from MAFF statistics; the most recent values are four- and three-year averages. The figures for area grown to semi-dwarf varieties were obtained from breeders' royalty returns agronomic environments and how will specific gene combinations perform? Furthermore, now that the acclaim following the 'Green Revolution1 is subsiding, the effects of the genes on yield stability are being questioned. For example, field surveys in developed and developing nations indicate that, ever since varieties based on Norin 10 have been grown on large areas, fluctuations in annual wheat production correlated with weather patterns have increased. This review attempts to summarize the agricultural impact of the various dwarfing genes in wheat; to outline the present understanding of their genetics; to summarize the known physiological mechanisms involved; and to describe, particularly in terms of yield characters, their known pleiotropic effects. The information reviewed is heavily weighted towards the Norin 10 genes. Their major impact on world agriculture has prompted intensive investigation and so they Brief history of dwarfing genes in wheat breeding 3 comprise the most completely understood genetic dwarfing system in wheat. This bias may not be out of place, however. Today's breeders must be aware of the relative merits and demerits of current germplasm derived from Norin 10 when considering the strategies to be adopted in producing the even more high-yielding and better 'quality' varieties for use in the latter part of this century. Brief history of dwarfing genes in wheat breeding The development of short-strawed varieties of bread wheat is not a recently formulated breeding objective. Breeders in Japan in the 19th century were probably the first to consciously employ genetic sources of short straw (Nonaka, 1984). A measure of their success is provided by the fact that most present-day short varieties of wheat owe their semi-dwarf characteristics to two Japanese genotypes. The first of these was the local variety Akakomugi, seed of which was obtained by Nazareno Strampelli at Rieti in Italy in 1911. His aims were to introduce early maturity and straw strength into Italian varieties. The varieties produced from his crosses with Akakomugi, for example Ardito and Villa Glori, incorporated both target traits (Strampelli, 1933) and have since formed the backbone of Italian hexaploid wheat breeding. The formal genetics of dwarfism from Akakomugi is only now becoming clear (see page 11). Nevertheless the genes involved have been transferred to other European programmes. The Italian wheats have played an important part in the development of the short, 'daylength-insensitive' (early maturing) genotypes produced by the Rockefeller International Wheat Improvement Project in Mexico which later, in 1963, became the International Maize and Wheat Improvement Center (CIMMYT). Mentana, which was one of Strampelli's taller varieties, was used in many breeding programmes and is cited by Norman Borlaug (1968) as one of the three key varieties in the Mexican programme in the 1940s. Ardito, one of the shortest of Strampelli's varieties, also has been grown outside Italy and was employed in the parentage of the important Russian variety, Bezostaya (FAO, 1971). The second Japanese source of dwarfism was Daruma. The initial crosses of Shirodaruma (white) and Akadaruma (red) with American varieties were made early this century in Japan (Gotoh, 1977). The most important and famous derivative of Shirodaruma is Norin 10, bred as Tohuko No. 34 in 1932 and renamed in 1934 (Matsumoto, 1968; Inazuka, 1971). Norin 10 was never an important variety in Japan; however, it was included in grain samples received by Orville Vogel at Washington State University in 1946. At that time, with the increasing use of artificial nitrogen fertilizers, Vogel was looking for sources of short straw specifically for use in the Pacific Northwest region of the USA. He made the first crosses in 1948 and, after initial difficulties with disease susceptibility and sterility, produced the line Norin 10-Brevor 14. This genotype was to become the main source of the two Norin 10 dwarfing genes (see pages 9-10) for both the USA and Mexican breeding programmes (Reitz and Salmon, 1968). Other sources of these genes, now known as Rhtl and Rht2 (Mclntosh, 1979), include Norin 16 and the Korean varieties Suweon 92 and Seu Seun 27, all of which originated from crosses with Daruma (Dalrymple, 1980). The spread of these genes through the world wheat crop has been meteoric by plant breeding standards. The first 'Norin 10 semi-dwarf variety, Gaines, was bred 4 Dwarfing genes in wheat by Vogel and released in 1961 (Vogel, 1964). This was followed by Nugaines in 1965 (Vogel and Peterson, 1974) and by 1979 a further 93 varieties derived from Norin 10 had been bred in North America (Dalrymple, 1980). In Mexico, the use of Norin 10-Brevor 14 in crosses with tall 'daylength insensitive' wheats produced semi-dwarf varieties of immense international significance. Borlaug was keenly aware of the need for short straw in his programme (Borlaug, 1968). He made his first successful crosses in 1955 and the Mexican government wheat programme released grain of the varieties Pitic 62 and Penjamo 62 to farmers in 1962. Since then all CIMMYT bread wheat lines have carried one or both of the Norin 10 dwarfing genes (Gale et aL, 1981). The international success of Borlaug's varieties owes much to the CIMMYT breeding methods. Alternate generations with selection under the different climatic and daylength regimes at sites near Mexico City and Cindad Obregon produces genotypes which are already adapted to agricultural conditions in N. Africa, India and Pakistan (Rao, 1974). Initially the genotypes developed by CIMMYT were used as varieties in these countries. Selections from the II 8156 (Penjamo 62/Gabo 55) such as Mexipak, Kalyansona, Super X and Siete Cerros 66 were particularly important in this respect. However, more recently, because of particular disease-resistance requirements, the most prominent varieties in these developing nations have been bred locally, but almost always from crosses involving CIMMYT germplasm. Their economic impact has been immense. For example India, where CIMMYT wheats were first grown in 1962, has, in spite of increased consumption by a growing population, moved from being an importer to being a wheat-exporting nation. In the more temperate wheat-producing countries, Mexican varieties have generally not been adequately adapted for direct use. However, the Rhtl and Rhtl genes have been almost universally introduced. Crosses made in Chile between French wheats and two of Vogel's lines, Vg 9144 and Vg 8058, were introduced into Britain through the Plant Breeding Institute (PBI) programme in 1964 (Lupton, 1975). Since 1974 all PBI winter wheats have carried Rhtl. In Australia, the first generation of semi-dwarf varieties were derived from the Mexican Rhtl genotype WW15 (Pugsley, 1974; Syme and Pugsley, 1975). In the USSR the importance of crosses made with the variety Red River 68 (A.A. Sosinov, personal communication), a sister line of Tobari 66 produced by CIMMYT, suggests that Rht2 may be prominent there. Dwarfing genes from Akakomugi and Norin 10 have also been transferred to tetraploid durum (macaroni) wheats. In Italy semi-dwarf durums were obtained by crossing the tall variety Cappelli with the short Akakomugi-derived hexaploids, Forlani and Acciaio (Vallega and Zitelli, 1973). Transfer of Norin 10 dwarfism to durums was first reported by Lebsock (1963) and appears to have been independently achieved on several subsequent occasions. The first semi-dwarf variety bred by CIMMYT was Oviachic 65 in 1965 (CIMMYT, 1977). Since that time varieties such as Jori 69, Cocorit 71, and Mexicali 75 (all Rhtl genotypes) have been employed internationally in much the same manner as the Mexican bread wheats. In the USA Rhtl has been transferred to varieties such as Modoc in California (Puri, Qualset and Vogt, 1978), Cando and Calvin (Quick, Miller and Donnelly, 1976; Quick, Donnelly and Miller, 1979) in North Dakota. In Italy, Mexican crosses were used to breed the Giorgi and Gerardo groups of semi-dwarfs (Vallega and Zitelli, 1973). Finally, in addition to the genetic sources described above, there have been many attempts to induce useful dwarfing genes by mutation breeding. Very few of the

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