Onions and Other Vegetable Alliums J.L
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CROP PRODUCTION SCIENCE IN HORTICULTURE SERIES Series Editors: Jeff Atherton, Professor of Tropical Horticulture, University of the West Indies, Barbados, and Alun Rees, Horticultural Consultant and former Editor, Journal of Horticultural Science and Biotechnology. This series examines economically important horticultural crops selected from the major production systems in temperate, subtropical and tropical climatic areas. Systems represented range from open field and plantation sites to protected plastic and glass houses, growing rooms and laboratories. Emphasis is placed on the scientific principles underlying crop production practices rather than on providing empirical recipes for uncritical acceptance. Scientific understanding provides the key to both reasoned choice of practice and the solution of future problems. Students and staff at universities and colleges throughout the world involved in courses in horticulture, as well as in agriculture, plant science, food science and applied biology at degree, diploma or certificate level will welcome this series as a succinct and readable source of information. The books will also be invaluable to progressive growers, advisers and end-product users requiring an authoritative, but brief, scientific introduction to particular crops or systems. Keen gardeners wishing to understand the scientific basis of recommended practices will also find the series very useful. The authors are all internationally renowned experts with extensive experience of their subjects. Each volume follows a common format covering all aspects of production, from background physiology and breeding, to propagation and planting, through husbandry and crop protection, to harvesting, handling and storage. Selective references are included to direct the reader to further information on specific topics. Titles Available: 1. Ornamental Bulbs, Corms and Tubers A.R. Rees 2. Citrus F.S. Davies and L.G. Albrigo 3. Onions and Other Vegetable Alliums J.L. Brewster 4. Ornamental Bedding Plants A.M. Armitage 5. Bananas and Plantains J.C. Robinson 6. Cucurbits R.W. Robinson and D.S. Decker-Walters 7. Tropical Fruits H.Y. Nakasone and R.E. Paull 8. Coffee, Cocoa and Tea K.C. Willson 9. Lettuce, Endive and Chicory E.J. Ryder 10. Carrots and Related Vegetable Umbelliferae V.E. Rubatzky, C.F. Quiros and P.W. Simon 11. Strawberries J.F. Hancock 12. Peppers: Vegetable and Spice Capsicums P.W. Bosland and E.J. Votava 13. Tomatoes E. Heuvelink 14. Vegetable Brassicas and Related Crucifers G. Dixon 15. Onions and Other Vegetable Alliums, 2nd Edition J.L. Brewster This page intentionally left blank ONIONS AND OTHER VEGETABLE ALLIUMS, 2ND EDITION James L. Brewster Formerly at Horticulture Research International (now known as Warwick HRI), Wellesbourne, Warwick CV35 9EF, UK CABI is a trading name of CAB International CABI Head Office CABI North American Office Nosworthy Way 875 Massachusetts Avenue Wallingford 7th Floor Oxfordshire OX10 8DE Cambridge, MA 02139 UK USA Tel: +44 (0)1491 832111 Tel: +1 617 395 4056 Fax: +44 (0)1491 833508 Fax: +1 617 354 6875 E-mail: [email protected] E-mail: [email protected] Website: www.cabi.org © J.L. Brewster 2008. 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. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data Brewster, James L. Onions and other vegetable alliums / J.L. Brewster. -- 2nd ed. p. cm. -- (Crop production science in horticulture series ; 15) Includes bibliographical references and index. ISBN 978-1-84593-399-9 (alk. paper) 1. Onions. 2. Allium. I. Title. II. Series: Crop production science in horticulture ; 15. SB341.B74 2008 635Ј.25--dc22 2007045092 ISBN: 978 1 84593 399 9 Typeset by Columns Design Ltd, Reading, UK. Printed and bound in the UK by Biddles Ltd, King’s Lynn. CONTENTS PREFACE ix 1 THE CLASSIFICATION, ORIGINS, DISTRIBUTION AND ECONOMIC IMPORTANCE OF THE MAJOR VEGETABLE CROPS 1 Classification of Alliums 1 Distribution and Ecology 4 The Types, History and Evolution of the Edible Crops 5 Economic Importance 22 2 THE STRUCTURE OF EDIBLE ALLIUMS 27 Visible Growth Stages 27 Outline of the Overall Plant Structure 27 The Structure of the Vegetative Shoot Apex 32 Stem Structure 33 Bulb Structure 36 Foliage Leaf Structure 39 The Root System 40 Inflorescence and Flower Structure and Development 45 Seeds 47 3 THE GENETICS AND PLANT BREEDING OF ALLIUM CROPS 51 Introduction 51 The Chromosomes 52 Breeding Systems 54 Genetics 56 v vi Contents Genetic Linkage Maps 59 Genomics 64 Hybrid Vigour 65 Genetic Male Sterility and Hybrid Breeding 65 Techniques of Breeding and Cultivar Improvement 67 Pollination Control in Breeding 73 Breeding Objectives in Edible Alliums 73 Selection Methods and Environmental Influences 75 Genetic Transformation of Alliums 77 Breeding for Disease and Pest Resistance 79 Conclusions 83 4 PHYSIOLOGY OF CROP GROWTH, DEVELOPMENT AND YIELD 85 Fundamental Determinants of Yield 85 Agronomic Factors Influencing Bulb Onion Yields 90 Seed Viability, Germination and Emergence 93 Vegetative Growth 116 Onion Bulbing 123 Models for Onion Growth and Bulbing 135 Physiology and Environmental Control of Flowering 138 Interrelations of Growth, Flowering and Bulbing in Onion 149 Shallots 151 Wakegi Onion 152 Garlic 156 Leek 160 Japanese Bunching Onion 166 Rakkyo 168 Chives 168 Chinese Chives 169 Concluding Remarks 169 5 INTERACTIONS WITH OTHER ORGANISMS: WEEDS, PESTS, DISEASES AND SYMBIONTS 171 Introduction 171 Weeds and Weed Control 173 Arthropod Pests 186 Nematode Pests 198 Viral Diseases 201 Bacterial Diseases 209 Contents vii Fungal Leaf Diseases 216 Fungal Diseases Infecting from the Soil 231 Fungal Diseases of Stored Bulbs 240 Mycorrhizal Fungi 246 6 AGRONOMY AND CROP PRODUCTION 251 Onion Production 251 Seed Production 293 Garlic Production 299 Leek Production 300 Japanese Bunching Onion Production 306 Chive Production 307 Chinese Chives Production 307 7 CROP STORAGE AND DORMANCY 309 Bulb Crop Storage and Dormancy 309 Garlic Dormancy and Storage 341 Leek Storage 342 Green Onion Storage 344 8 BIOCHEMISTRY, HEALTH BENEFITS AND FOOD SCIENCE OF ALLIUMS 347 The Biochemistry of Flavour 347 Carbohydrate Biochemistry 356 Flavonoids 361 Medicinal Effects of Alliums 364 Processed Products from Alliums 370 REFERENCES 373 INDEX 413 The colour plate section can be found following p. 212 This page intentionally left blank PREFACE A number of people, including lecturers, researchers and seed company staff, have told me that they found the first edition of this book useful. This is the best reward for writing such a book. A lot of new allium science has been published since the first edition and progress has been rapid, particularly in the fields of genetics, plant breeding and plant pathology, where the techniques of molecular biology have had great impact. So, an attempt at an update was clearly worthwhile. Shortly before writing the first edition I had co-edited the multi-author Onions and Allied Crops with Haim Rabinowitch, so up to date reviews by experts in the relevant science were to hand and familiar to me. For this edition, I have had to resort more to the original research literature, although the reviews by contributors to Rabinowitch and Currah’s (2002) Allium Crop Science: Recent Advances were key sources for many topics. A book of this breadth cannot attempt to be an exhaustive review of the research literature and, inevitably, in choosing what research to mention, there is some bias to what I am familiar with, notably towards work from Wellesbourne, from the UK and what is published in English. I apologize to those who have done interesting work which is not mentioned but, nevertheless, I hope the book gives a reasonable overview of the current scientific understanding about the vegetable alliums. I hope this second edition will help to raise and maintain awareness of the huge and continuing achievement of allium scientists in discovering so much fascinating and useful information about these crops. These researchers are the real heroes and heroines behind this book. I hope I have not misrepresented their work by error, omission or misplaced emphasis. The aim of the book is to introduce the scientific principles that underlie production practices, rather than to give detailed instruction on how to produce allium crops. If the crop technologist has an understanding of the basic principles fundamental to his craft he/she should be better able to adapt his/her methods to changing circumstances and opportunities, and better be able to understand and cope with any abnormal or unusual problems that he/she encounters. Understanding of principles should also improve his/her ix x Preface judgement in deciding between possible alternative techniques. Furthermore, the details of production methods change as new products, legislation or market conditions arise, whereas the underlying principles remain true and relevant, although of course our knowledge of principles improves and increases as science progresses. It may seem over-ambitious to cover the wide range of science closely relevant to allium production, but one advantage of such an undertaking is that it reveals connections between different aspects of the underlying science and, sometimes, conflicts in the recommendations for practice coming from them. Furthermore, the crop grower has to embody the integral of all the underlying science in his/her production methods. The book is obviously aimed at students and professionals with a special interest in the allium vegetables, but I hope it might interest the occasional, more general, reader. On my first day as an undergraduate agricultural student at Wye College, UK, I remember the then Principal, Dunstan Skilbeck, remarking in his intro- ductory talk to new students, that the studies they were about to embark on were not only a source of practically applicable information but also offered a ‘window’ into many fields of knowledge; a possibility particularly well provided for by broadly based courses like agriculture or horticulture.