BIOTECHNOLOGY IN AGRICULTURE SERIES General Editor: Gabrielle J. Persley, International Service for National Agri- cultural Research (The Hague, Netherlands), and Project Manager, World Bank/lSNAR/ACIAR/AIDAB Biotechnology Study. For a number of years, biotechnology has held out the prospect for major advances in agricultural production, but only recently have the results of this new revolution started to reach application in the field. The potential for further rapid developments is however immense. The aim of this new book series is to review advances and current knowledge in key areas of biotechnology as applied to crop and animal production, forestry and food science. Some titles will focus on individual crop species or groups of species, others on specific goals such as plant protection or animal health, with yet others addressing particular methodo- logies such as tissue culture, transtormation or immunoassay. In some cases, relevant molecular and cell biology and genetics will also be covered. Issues of relevance to both industrialized and developing countries will be addressed, and social, economic and legal implications will also be considered. Most titles will be written for research workers in the biological sciences and agriculture, but some will also be useful as text- books for senior-level students in these disciplines. Editorial Advisory Board: P.J. Brumby, formerly of the World Bank, Washington DC, USA. E.P. Cunningham, FAO, Rome, Italy. P. Day, Rutgers University, New Jersey, USA. J.H. Dodds, International Potato Center (CIP), Peru. J.J. Doyle, International Laboratory tor Research on Animal Diseases, Nairobi, Kenya. S.L. Krugman, United States Department of Agriculture, Forest Service. W,J. Peacock, CSIRO, Division of Plant Industry, Australia. Titles Available: 1 : Beyond Mendel’s Garden: Biotechnology in the Service of World Agriculture G.J. Pers/ey 2: Agricultural Biotechnology: Opportunities for International Development Edited by G.J. Persley 3: The Molecular and Cellular Biology of the Potato Edited by M.E. Vayda and W.D. Park 4: Advanced Methods in Plant Breeding and Biotechnology Edited by D. R. Murray 5: Barley: Genetics, Biochemistry, Molecular Biology and Biotechnology Edited by P. R. Shewry 6: Kice Biotechnology Edited by G.S. Khush and G.H. Toenniessen 7: Plant Genetic Manipulation for Crop Protection Edited by A. Gatehouse, V. Hilder, dnd D. Boulter Titles in Preparation Biotechnology of Perennial Fruit Crops Edited by F. Hammerschlag and R. Litz Non-Food Uses of Agricultural Raw Materials: Economics, Biotechnology and Politics C. Spelman Rice Biotechnology Edited by G S Khush International Rice Research Institute The Philippines and G H Toenniessen The Rockefeller Foundation, New York USA C AB International in association with the International Rice Research Institute To Robert F. and Sunny Chandler ~ ~ ~~ ~ C . A. B International Tel: Wallingford (0491) 32111 Wallingford Telex: 847964 (COMAGG G) Oxon OX10 8DE Telecom Gofd/Dialcom: 84: CAU00l UK Fax: (0491) 33508 © C • A • B International 1 991. 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. Published in Association with: international Rice Research Institute P.O. Box 933 1099 Manila Philippines A catalogue record for this hook is available from the British library ISBN 0 85198 712 5 (CABI) ISSN 0960-202X (CABI) ISBN 971 22 0013 2 (IRRI) Typeset by Leaper & Gard Ltd, Brisrol Printed and bound in the UK by The Alden Press Ltd, Oxford Contents Foreword vii Preface ix 1. The World Rice Economy: Challenges Ahead 1 Cristina C David 2. Research Priorities for Rice Biotechnology 19 Robert W Herdt 3. Genetic Diversity of Wild and Cultivated Rice 55 H I Oka 4. Rice Karyotype, Marker Genes, and Linkage Croups 83 Gurdev S Khush and Toshiro Kinoshita 5. Development and Use of Restriction Fragment Length Polymorphism in Rice Breeding and Genetics 109 S R McCouch and S D Tanksley 6. Rice Tissue Culture and its Application 135 P T Lynch, R P Finch, M R Davey and E C Cocking 7. Transformation and Kegeneration of Kice Protoplasts 157 Thomas K Hodges, Jianying Peng, Leszek A Lyznik, and David S Koetje v vi Contents 8. Assessment of Rice Genetic Transformation Techniques 175 Jun Cao, Wanggen Zhang, David McElroy, and Ray Wu 9. The Identification and Characterization of Rice Nuclear Genes 199 Thomas W Okita 10. Gene Expression in Rice 225 Virginia Walbot and Daniel Gallie 11. Potentially Useful Genes for Rice Genetic Engineering 253 Gary H Toenniessen 12. Molecular Probes for Disease Diagnosis and Monitoring 281 Jan E Leach and Frank F White 13. Trospects for the Future 309 Gary H Toenniessen and Gurdev S Khush Index 315 Foreword Populations in rice growing countries were increasing at a faster rate than food production in the 1950s and 1960s. Because of these trends, several authorities predicted large-scale food shortages in Asia resulting in famines and social upheavals, by the 1970s. Concerned at the impending crisis, the Rockefeller Foundation, the Ford Foundation, and the Government of the Philippines estab- lished the International Rice Research Institute (IRRI) in 1960 to harness the power of science for solving the problem of stagnant rice yields in developing countries. A series of high yielding rice varieties and other technology developed at IRRI and national agricultural research systems started the green revolution in rice and led to major increases in rice production. Most of the rice growing countries in Asia, where 92% of the world's rice is grown, became self-sufficient in food production. Worldwide, rice production doubled in a 25-year period, from 257 million tons in 1965 to 520 million tons in 1990. But the spectre of food shortages looms once again. The annual rate of increase in rice production has slowed down and there are no new areas that can be brought into rice production. World population which stands at 5.3 billion now is likely to exceed 6 billion by the turn of the century and 8 billion by 2020. The population of rice eaters is increasing at an exceptionally fast rate; the number of rice consumers will probably double by the year 2020. The Inter- national Food Policy Research Institute predicts that demand for rice will exceed production by the end of this century. We must therefore accelerate the rate of growth in rice production. New rice varieties with higher yield potential are essential to achieve this goal. In IRRI’s long-term research strategy, we have conceptualized rice varieties of the future for five rice-growing ecologies. Such varieties must have higher yield potential than present high-yielding-varieties, excellent grain quality, resistance to biotic and abiotic stresses, and should use inputs more efficiently. Recent developments in cellular and molecular genetics give us tools that hold great vii VIII Foreword promise in expediting such genetic improvement. Little has been known until recently, however, about the cellular and molecular genetics of rice. Fortunately, the Rockefeller Foundation again took the initiative and organized an Inter- national Program on Rice Biotechnology in 1985. The Foundation is funding research on rice biotechnology in carefully selected laboratories in industrial countries, and helping build capacity to use biotechnology in rice improvement in developing countries. Much useful information has been generated during the last 5 years; rice is now considered a ‘model’ cereal crop for biotechnology research. The Rockefeller Foundation and IRRI have collaborated in producing this book, which reviews the latest breakthroughs in rice biotechnology. I hope this volume will prove useful to rice breeders, geneticists and biotechnologists alike, and will help pave way for another scientific breakthrough toward a higher yield plateau for rice. K L Lampe Director General International Rice Research Institute Los Baños, The Philippines Preface Rice is the most important food crop of the developing world and rice genetic improvement through breeding has been an effective mechanism for delivering the benefits of science and technology to hundreds of millions of resource-poor people. A research system with regional, national, and international components is now in place for producing improved rice varieties and making them available to farmers that need them. Biotechnology can significantly strengthen rice breeding programs and help produce new varieties with higher yield potential and greater yield stability. These should improve the efficiency of rice production and allow an expansion of the rice-growing area, thereby preventing future food shortages from occurring as demand for rice increases because of population growth and economic development. A strong breeding program is a prerequisite to the application of the new biotechnologies for rice genetic improvement. Biotechnology will enable rice breeders to achieve results more quickly and efficiently and will help them to attain breeding goals not feasible using conventional techniques. Biotechnology will complement, not replace, breeding. In less than a decade research on rice biotechnology has moved from a position of neglect to a position where some now consider rice a model plant for cereal research. A useful knowledge base on rice molecular genetics and cellular biology has been established and the tools necessary for applying this knowledge to rice genetic improvement are being refined and becoming available. Tissue culture techniques such as anther culture, embryo rescue, and use of somaclonal variants have contributed to the release of new rice varieties. These technologies are of proven benefit to rice breeding and through research are becoming applic- able to a broader range of rice cultivars and breeding objectives. Most molecular genetic techniques are still being developed, but progress has been more rapid with rice than any other cereal. An RFLP map with 350 markers ix I x Preface has been prepared and several genes of economic importance have already been tagged with markers.