Induced Resistance for Plant Defence A Sustainable Approach to Crop Protection Edited by Dale Walters Crop and Soil Systems Research Group, Scottish Agricultural College, Edinburgh, UK Adrian Newton Scottish Crop Research Institute, Invergowrie, Dundee, UK Gary Lyon Scottish Crop Research Institute, Invergowrie, Dundee, UK This Page Intentionally Left Blank Induced Resistance for Plant Defence This Page Intentionally Left Blank Induced Resistance for Plant Defence A Sustainable Approach to Crop Protection Edited by Dale Walters Crop and Soil Systems Research Group, Scottish Agricultural College, Edinburgh, UK Adrian Newton Scottish Crop Research Institute, Invergowrie, Dundee, UK Gary Lyon Scottish Crop Research Institute, Invergowrie, Dundee, UK ©2007 Blackwell Publishing Blackwell Publishing editorial offices: Blackwell Publishing Ltd, 9600 Garsington Road, Oxford OX4 2DQ, UK Tel:(cid:1)44 (0)1865 776868 Blackwell Publishing Professional, 2121 State Avenue, Ames, Iowa 50014-8300, USA Tel:(cid:1)1 515 292 0140 Blackwell Publishing Asia Pty Ltd, 550 Swanston Street, Carlton, Victoria 3053, Australia Tel:(cid:1)61 (0)3 8359 1011 The right of the Author to be identified as the Author of this Work has been asserted in accordance with the Copyright, Designs and Patents Act 1988. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by the UK Copyright, Designs and Patents Act 1988, without the prior permission of the publisher. First published 2007 by Blackwell Publishing Ltd ISBN: 978-1-4051-3447-7 Library of Congress Cataloging-in-Publication Data Induced resistance for plant defence: a sustainable approach to crop protection / edited by Dale Walters, Adrian Newton, Gary D. Lyon. – 1st. ed. p. cm. Includes bibliographical references and index. ISBN: 978-1-4051-3447-7 (hardback : alk. paper) 1. Plants–Disease and pest resistance–Genetic aspects. 2. Plants–Disease and pest resistance–Molecular aspects. I. Walters, Dale. II. Newton, Adrian C. III. Lyon, Gary D. SB750.I4745 2007 632(cid:2).9–dc22 2006026449 A catalogue record for this title is available from the British Library Set in 10/12.5 pt Times by Charon Tec Ltd (A Macmillan Company), Chennai, India www.charontec.com Printed and bound in Singapore by COS Printers Pte Ltd The publisher’s policy is to use permanent paper from mills that operate a sustainable forestry policy, and which has been manufactured from pulp processed using acid-free and elementary chlorine-free practices. Furthermore, the publisher ensures that the text paper and cover board used have met acceptable environmental accreditation standards. For further information on Blackwell Publishing, visit our website: www.blackwellpublishing.com Contents List of contributors ix Preface xi Chapter 1 Introduction: definitions and some history 1 Ray Hammerschmidt 1.1 Induced resistance: an established phenomenon 1 1.2 Terminology and types of induced resistance 1 1.3 A little history 3 1.4 It’s all about interactions 6 1.5 Acknowledgements 7 1.6 References 7 Chapter 2 Agents that can elicit induced resistance 9 Gary Lyon 2.1 Introduction 9 2.2 Compounds inducing resistance 10 2.3 Conclusions 21 2.4 Acknowledgements 23 2.5 References 23 Chapter 3 Genomics in induced resistance 31 Kemal Kazan and Peer M. Schenk 3.1 Introduction 31 3.2 Transcriptome analyses for discovery of genes involved in induced resistance 32 3.3 Proteome analyses and induced resistance 40 3.4 Metabolome analysis and induced resistance 41 3.5 Forward genetic approaches for discovery of genes involved in induced resistance 43 3.6 Reverse genetic approaches 45 3.7 Manipulation of master switches for activation of induced resistance 50 3.8 Suitable promoters for defence gene expression 51 3.9 Conclusions: a systems biological approach to induced plant defence? 54 3.10 Acknowledgements 56 3.11 References 56 Chapter 4 Signalling cascades involved in induced resistance 65 Corné M.J. Pieterse and L.C. Van Loon 4.1 Introduction 65 4.2 SA, JA and ET: important signals in primary defence 66 v vi Contents 4.3 SA, JA and ET: important signals in induced disease resistance 68 4.4 Crosstalk between signalling pathways 78 4.5 Outlook 80 4.6 Acknowledgements 81 4.7 References 81 Chapter 5 Types and mechanisms of rapidly induced plant resistance to herbivorous arthropods 89 Michael J. Stout 5.1 Introduction: induced resistance in context 89 5.2 Comparison of the threats posed by pathogens and herbivores 90 5.3 Types of induced resistance 92 5.4 Establishing the causal basis of induced resistance 99 5.5 Arthropods as dynamic participants in plant–arthropod interactions 102 5.6 Conclusions 103 5.7 References 104 Chapter 6 Mechanisms of defence to pathogens: biochemistry and physiology 109 Christophe Garcion, Olivier Lamotte and Jean-Pierre Métraux 6.1 Introduction 109 6.2 Structural barriers 109 6.3 Phytoalexins 112 6.4 The hypersensitive response (HR) 117 6.5 Antifungal proteins 121 6.6 Conclusions 123 6.7 References 123 Chapter 7 Induced resistance in natural ecosystems and pathogen population biology: exploiting interactions 133 Adrian Newton and Jörn Pons-Kühnemann 7.1 Introduction 133 7.2 Environmental variability 133 7.3 Ecology of the plant environment 134 7.4 Environmental parameters 136 7.5 Plant and pathogen population genetics 136 7.6 Consequences of resistance induction 138 7.7 Conclusions 139 7.8 Acknowledgements 140 7.9 References 140 Chapter 8 Microbial induction of resistance to pathogens 143 Dale Walters and Tim Daniell 8.1 Introduction 143 8.2 Resistance induced by plant growth promoting rhizobacteria 143 Contents vii 8.3 Induction of resistance by biological control agents 148 8.4 Resistance induced by composts 149 8.5 Disease control provided by an endophytic fungus 149 8.6 Mycorrhizal symbiosis and induced resistance 150 8.7 Acknowledgements 152 8.8 References 152 Chapter 9 Trade-offs associated with induced resistance 157 Martin Heil 9.1 Introduction 157 9.2 Artificial resistance inducers 159 9.3 Costs of SAR 163 9.4 Conclusions 169 9.5 Acknowledgements 170 9.6 References 170 Chapter 10 Topical application of inducers for disease control 179 Philippe Reignault and Dale Walters 10.1 Introduction 179 10.2 Biotic inducers 179 10.3 Abiotic inducers 184 10.4 Conclusions 194 10.5 Acknowledgements 194 10.6 References 194 Chapter 11 Integration of induced resistance in crop production 201 Tony Reglinski, Elizabeth Dann and Brian Deverall 11.1 Introduction 201 11.2 Induced resistance for disease control 202 11.3 Variable efficacy of induced resistance 206 11.4 Compatibility of activators with other control methods 209 11.5 Integration of plant activators in crop management 216 11.6 Knowledge gaps 221 11.7 Conclusions 222 11.8 References 223 Chapter 12 Exploitation of induced resistance: a commercial perspective 229 Andy Leadbeater and Theo Staub 12.1 Introduction 229 12.2 Science and serendipitous discovery of resistance- inducing compounds 230 12.3 Discovery of INAs and BTHs 231 12.4 Identification of BION®and other SAR activators 231 12.5 The role of basic studies in the discovery of BION®and other SAR/ISR products 232 12.6 Identification of harpin 233 12.7 The commercial development of an induced resistance product 234 viii Contents 12.8 Innovation in registration? 236 12.9 Commercial experiences with induced resistance products 237 12.10 Conclusions 240 12.11 References 241 Chapter 13 Induced resistance in crop protection: the future, drivers and barriers 243 Gary Lyon, Adrian Newton and Dale Walters 13.1 Introduction 243 13.2 Strategies to increase efficacy and durability in the field 243 13.3 What research is required to make induced resistance work in practice? 244 13.4 Can we breed plants with enhanced responsiveness to inducers? 246 13.5 The potential for GM plants containing SAR-related genes 246 13.6 Political, economic and legislation issues 247 13.7 Conclusion 247 13.8 Acknowledgements 248 13.9 References 248 Index 251
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