Induced Resistance for Plant Defence 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 List of contributors Dr Tim Daniell Scottish Crop Research Institute, Invergowrie, Dundee DD2 5DA, UK E-mail: [email protected] Dr Elizabeth Dann Department of Primary Industries and Fisheries, 80 Meiers Road, Queensland 4068, Australia E-mail: [email protected] Professor Brian Deverall Faculty of Agriculture, Food and Natural Resources, University of Sydney, NSW 2006, Australia E-mail: [email protected] Dr Christophe Garcion Departemente de Biologie, Université de Fribourg, 1700 Fribourg, Switzerland E-mail: [email protected] Professor Ray Department of Plant Pathology, 107 CIPS Building, Hammerschmidt Michigan State University, East Lansing, MI 48824-1311, USA E-mail: [email protected] Professor Martin Heil Department of General Botany – Plant Ecology, University of Duisburg-Essen, Universitatsstr. 5, D-45117 Essen, Germany E-mail: [email protected] Dr Kemal Kazan Commonwealth Scientific and Industrial Research Organisation, Plant Industry, Queensland Bioscience Precinct, St Lucia, Queensland 4069, Australia E-mail: [email protected] Dr Olivier Lamotte Departemente de Biologie, Université de Fribourg, 1700 Fribourg, Switzerland [email protected] Dr Andy Leadbeater Syngenta Crop Protection AG E-mail: [email protected] Dr Gary Lyon Scottish Crop Research Institute, Invergowrie, Dundee DD2 5DA, UK E-mail: [email protected] Professor Jean-Pierre Departemente de Biologie, Université de Fribourg, 1700 Métraux Fribourg, Switzerland E-mail: [email protected] ix x List of contributors Dr Adrian Newton Scottish Crop Research Institute, Invergowrie, Dundee DD2 5DA, UK E-mail: [email protected] Professor Corné Phytopathology, Institute of Environmental Biology, M.J. Pieterse Utrecht University, PO Box 80084, 3508 TB Utrecht, The Netherlands E-mail: [email protected] Dr Jörn Pons-Kühnemann Biometry and Population Genetics, Giessen University, Heinrich-Buff-Ring 26-32, 35392 Giessen, Germany E-mail: [email protected] Dr Tony Reglinski Bioprotection Group, HortResearch, Ruakura Research Centre, East Street, Private Bag 3123, Hamilton, New Zealand E-mail: [email protected] Dr Philippe Reignault Mycologie/Phytopathologie/Environnement, Université du Littoral Côte d’Opale, 17, avenue Louis Blériot BP 699, F-62228 Calais Cedex, France E-mail: [email protected] Dr Peer Schenk School of Integrative Biology, University of Queensland, St Lucia, Queensland 4072, Australia E-mail: [email protected] Dr Theo Staub Syngenta Crop Protection AG Professor Mike Stout Department of Entomology, Louisiana State University Agricultural Center, 402 Life Sciences Building, Louisiana State University, Baton Rouge, LA 70803, USA E-mail: [email protected] Professor L.C. Phytopathology, Institute of Environmental Biology, Van Loon Utrecht University, PO Box 80084, 3508 TB Utrecht, The Netherlands E-mail: [email protected] Professor Dale Crop & Soil Systems Research Group, Walters Scottish Agricultural College, King’s Buildings, West Mains Road, Edinburgh EH9 3JG, UK E-mail: [email protected] Preface Plant diseases have been a problem for mankind since the very beginnings of agriculture. As we write this preface, some 12,000 years later, plant diseases are still a problem. We have learned a great deal about plant diseases and how to control them in the intervening millennia, but disease still takes its toll on our crops every year. The problem is the result, in large part, of the genetic adaptability of the pathogens responsible for causing plant dis- eases: they develop resistance to our crop protection chemicals and rapidly overcome the resistance bred into our new crop varieties. In the fight against plant disease, it is essen- tial therefore that we keep one (or preferably several) steps ahead of the pathogens. In their review of global food security, Strange & Scott (2005; Annual Review of Phytopathology43,83–116) point out that more than 800 million people worldwide do not have sufficient food, and some 1.3 billion people survive on less than $1 a day. Further, a survey by The Economist in 2000 (The Economist, March 25) estimated that there will be an additional 1.5 billion people to feed by 2020, requiring farmers to produce 39% more grain. Since it is estimated that some 12% of global crop production is lost to plant disease annually, it is clear that the need for efficient, reliable and affordable disease control measures has never been greater. Equally important from the modern perspective is the need to ensure that any new disease control measures maintain crop yield and qual- ity, without harming our fragile and long suffering environment. Although the first recorded observations of induced resistance date back to the 19th cen- tury, the phenomenon was largely ignored until the late 1950s and early 1960s. Even then, the concept of induced resistance was largely ignored, despite the very solid foundation being laid by Joe Kuc´ and his colleagues. There was a gradual awakening of interest, and induced resistance has attracted increasing attention in the last 15 years or so. This interest is not surprising, since induced resistance offers the prospect of broad spectrum, long last- ing and, hopefully, environmentally benign disease control. However, this prospect will not be realized unless we are able to translate our ever increasing understanding of the cellular basis of induced resistance to the practical, field situation. This requires integration of molecular biology and biochemistry, with crop science and ecology. In this book, our aim is to provide plant pathologists, crop protectionists, agronomists and others with an update of the broad and complex topic that is induced resistance and to highlight the efforts being made to provide the understanding necessary to allow induced resistance to be used in practice. The various chapters in the book cover the cellular aspects of induced resistance, including signalling and defence mechanisms, the trade-offs associated with the expression of induced resistance, work on integrating induced resistance into crop protection practice and induced resistance from a commercial perspective. Our hope is that this book will excite the interest of plant and crop scientists and encourage the collaboration between molecular biologists, plant pathologists and ecologists that will be necessary to realize the great potential offered by induced resistance. Dale Walters, Adrian Newton and Gary Lyon xi
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