Induced Resistance for Plant Defence: A Sustainable Approach to Crop Protection

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  • Induced Resistance forPlant Defence

    A Sustainable Approach to Crop Protection

    Edited by

    Dale WaltersCrop and Soil Systems Research Group,

    Scottish Agricultural College, Edinburgh, UK

    Adrian NewtonScottish Crop Research Institute, Invergowrie, Dundee, UK

    Gary LyonScottish Crop Research Institute, Invergowrie, Dundee, UK

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  • Induced Resistance for Plant Defence

  • This Page Intentionally Left Blank

  • Induced Resistance forPlant Defence

    A Sustainable Approach to Crop Protection

    Edited by

    Dale WaltersCrop and Soil Systems Research Group,

    Scottish Agricultural College, Edinburgh, UK

    Adrian NewtonScottish Crop Research Institute, Invergowrie, Dundee, UK

    Gary LyonScottish Crop Research Institute, Invergowrie, Dundee, UK

  • 2007 Blackwell PublishingBlackwell Publishing editorial offices:

    Blackwell Publishing Ltd, 9600 Garsington Road, Oxford OX4 2DQ, UKTel: 44 (0)1865 776868Blackwell Publishing Professional, 2121 State Avenue, Ames, Iowa 50014-8300, USATel: 1 515 292 0140Blackwell Publishing Asia Pty Ltd, 550 Swanston Street, Carlton, Victoria 3053, AustraliaTel: 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 DataInduced 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. PlantsDisease and pest resistanceGenetic aspects. 2. PlantsDisease and pest resistanceMolecular aspects. I. Walters, Dale. II. Newton, Adrian C. III. Lyon, Gary D.

    SB750.I4745 2007632.9dc222006026449

    A catalogue record for this title is available from the British Library

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    Printed and bound in Singaporeby COS Printers Pte Ltd

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  • Contents

    List of contributors ixPreface xi

    Chapter 1 Introduction: definitions and some history 1Ray Hammerschmidt1.1 Induced resistance: an established phenomenon 11.2 Terminology and types of induced resistance 11.3 A little history 31.4 Its all about interactions 61.5 Acknowledgements 71.6 References 7

    Chapter 2 Agents that can elicit induced resistance 9Gary Lyon2.1 Introduction 92.2 Compounds inducing resistance 102.3 Conclusions 212.4 Acknowledgements 232.5 References 23

    Chapter 3 Genomics in induced resistance 31Kemal Kazan and Peer M. Schenk3.1 Introduction 313.2 Transcriptome analyses for discovery of genes

    involved in induced resistance 323.3 Proteome analyses and induced resistance 403.4 Metabolome analysis and induced resistance 413.5 Forward genetic approaches for discovery of genes

    involved in induced resistance 433.6 Reverse genetic approaches 453.7 Manipulation of master switches for activation of

    induced resistance 503.8 Suitable promoters for defence gene expression 513.9 Conclusions: a systems biological approach to induced

    plant defence? 543.10 Acknowledgements 563.11 References 56

    Chapter 4 Signalling cascades involved in induced resistance 65Corn M.J. Pieterse and L.C. Van Loon4.1 Introduction 654.2 SA, JA and ET: important signals in primary defence 66


  • 4.3 SA, JA and ET: important signals in induceddisease resistance 68

    4.4 Crosstalk between signalling pathways 784.5 Outlook 804.6 Acknowledgements 814.7 References 81

    Chapter 5 Types and mechanisms of rapidly induced plant resistanceto herbivorous arthropods 89Michael J. Stout5.1 Introduction: induced resistance in context 895.2 Comparison of the threats posed by pathogens

    and herbivores 905.3 Types of induced resistance 925.4 Establishing the causal basis of induced resistance 995.5 Arthropods as dynamic participants in

    plantarthropod interactions 1025.6 Conclusions 1035.7 References 104

    Chapter 6 Mechanisms of defence to pathogens: biochemistry andphysiology 109Christophe Garcion, Olivier Lamotte and Jean-Pierre Mtraux6.1 Introduction 1096.2 Structural barriers 1096.3 Phytoalexins 1126.4 The hypersensitive response (HR) 1176.5 Antifungal proteins 1216.6 Conclusions 1236.7 References 123

    Chapter 7 Induced resistance in natural ecosystems and pathogenpopulation biology: exploiting interactions 133Adrian Newton and Jrn Pons-Khnemann7.1 Introduction 1337.2 Environmental variability 1337.3 Ecology of the plant environment 1347.4 Environmental parameters 1367.5 Plant and pathogen population genetics 1367.6 Consequences of resistance induction 1387.7 Conclusions 1397.8 Acknowledgements 1407.9 References 140

    Chapter 8 Microbial induction of resistance to pathogens 143Dale Walters and Tim Daniell8.1 Introduction 1438.2 Resistance induced by plant growth promoting rhizobacteria 143

    vi Contents

  • 8.3 Induction of resistance by biological control agents 1488.4 Resistance induced by composts 1498.5 Disease control provided by an endophytic fungus 1498.6 Mycorrhizal symbiosis and induced resistance 1508.7 Acknowledgements 1528.8 References 152

    Chapter 9 Trade-offs associated with induced resistance 157Martin Heil9.1 Introduction 1579.2 Artificial resistance inducers 1599.3 Costs of SAR 1639.4 Conclusions 1699.5 Acknowledgements 1709.6 References 170

    Chapter 10 Topical application of inducers for disease control 179Philippe Reignault and Dale Walters10.1 Introduction 17910.2 Biotic inducers 17910.3 Abiotic inducers 18410.4 Conclusions 19410.5 Acknowledgements 19410.6 References 194

    Chapter 11 Integration of induced resistance in crop production 201Tony Reglinski, Elizabeth Dann and Brian Deverall11.1 Introduction 20111.2 Induced resistance for disease control 20211.3 Variable efficacy of induced resistance 20611.4 Compatibility of activators with other control methods 20911.5 Integration of plant activators in crop management 21611.6 Knowledge gaps 22111.7 Conclusions 22211.8 References 223

    Chapter 12 Exploitation of induced resistance: a commercial perspective 229Andy Leadbeater and Theo Staub12.1 Introduction 22912.2 Science and serendipitous discovery of resistance-

    inducing compounds 23012.3 Discovery of INAs and BTHs 23112.4 Identification of BION and other SAR activators 23112.5 The role of basic studies in the discovery of BION and

    other SAR/ISR products 23212.6 Identification of harpin 23312.7 The commercial development of an induced

    resistance product 234

    Contents vii

  • 12.8 Innovation in registration? 23612.9 Commercial experiences with induced resistance products 23712.10 Conclusions 24012.11 References 241

    Chapter 13 Induced resistance in crop protection: the future, drivers and barriers 243Gary Lyon, Adrian Newton and Dale Walters13.1 Introduction 24313.2 Strategies to increase efficacy and durability in the field 24313.3 What research is required to make induced resistance

    work in practice? 24413.4 Can we breed plants with enhanced responsiveness

    to inducers? 24613.5 The potential for GM plants containing SAR-related genes 24613.6 Political, economic and legislation issues 24713.7 Conclusion 24713.8 Acknowledgements 24813.9 References 248

    Index 251

    viii Contents

  • List of contributors

    Dr Tim Daniell Scottish Crop Research Institute, Invergowrie, Dundee DD2 5DA, UKE-mail:

    Dr Elizabeth Dann Department of Primary Industries and Fisheries, 80 Meiers Road, Queensland 4068, AustraliaE-mail:

    Professor Brian Deverall Faculty of Agriculture, Food and Natural Resources,University of Sydney, NSW 2006, AustraliaE-mail:

    Dr Christophe Garcion Departemente de Biologie, Universit de Fribourg, 1700Fribourg, SwitzerlandE-mail:

    Professor Ray Department of Plant Pathology, 107 CIPS Building,Hammerschmidt Michigan State University, East Lansing, MI

    48824-1311, USAE-mail:

    Professor Martin Heil Department of General Botany Plant Ecology,University of Duisburg-Essen, Universitatsstr. 5, D-45117Essen, GermanyE-mail:

    Dr Kemal Kazan Commonwealth Scientific and Industrial ResearchOrganisation, Plant Industry, Queensland BiosciencePrecinct, St Lucia, Queensland 4069, AustraliaE-mail:

    Dr Olivier Lamotte Departemente de Biologie, Universit de Fribourg, 1700Fribourg,

    Dr Andy Leadbeater Syngenta Crop Protection AGE-mail:

    Dr Gary Lyon Scottish Crop Research Institute, Invergowrie, Dundee DD2 5DA, UKE-mail:

    Professor Jean-Pierre Departemente de Biologie, Universit de Fribourg, 1700 Mtraux Fribourg, Switzerland



  • Dr Adrian Newton Scottish Crop Research Institute, Invergowrie, Dundee DD2 5DA, UKE-mail:

    Professor Corn Phytopathology, Institute of Environmental Biology, M.J. Pieterse Utrecht University, PO Box 80084, 3508 TB Utrecht,

    The NetherlandsE-mail:

    Dr Jrn Pons-Khnemann Biometry and Population Genetics, Giessen University,Heinrich-Buff-Ring 26-32, 35392 Giessen, GermanyE-mail:

    Dr Tony Reglinski Bioprotection Group, HortResearch, Ruakura ResearchCentre, East Street, Private Bag 3123, Hamilton, New ZealandE-mail:

    Dr Philippe Reignault Mycologie/Phytopathologie/Environnement, Universitdu Littoral Cte dOpale, 17, avenue Louis Blriot BP699, F-62228 Calais Cedex, FranceE-mail:

    Dr Peer Schenk School of Integrative Biology, University of Queensland,St Lucia, Queensland 4072, AustraliaE-mail:

    Dr Theo Staub Syngenta Crop Protection AG

    Professor Mike Stout Department of Entomology, Louisiana State UniversityAgricultural Center, 402 Life Sciences Building,Louisiana State University, Baton Rouge, LA 70803,USAE-mail:

    Professor L.C. Phytopathology, Institute of Environmental Biology, Van Loon Utrecht University, PO Box 80084, 3508 TB Utrecht,

    The NetherlandsE-mail:

    Professor Dale Crop & Soil Systems Research Group, Walters Scottish Agricultural College, Kings Buildings,

    West Mains Road, Edinburgh EH9 3JG, UKE-mail:

    x List of contributors

  • 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. Wehave learned a great deal about plant diseases and how to control them in the interveningmillennia, 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 theresistance 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 ofPhytopathology 43, 83116) point out that more than 800 million people worldwide donot 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 thatthere will be an additional 1.5 billion people to feed by 2020, requiring farmers to produce39% more grain. Since it is estimated that some 12% of global crop production is lost toplant disease annually, it is clear that the need for efficient, reliable and affordable diseasecontrol measures has never been greater. Equally important from the modern perspectiveis 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 foundationbeing laid by Joe Kuc and his colleagues. There was a gradual awakening of interest, andinduced resistance has attracted increasing attention in the last 15 years or so. This interestis not surprising, since induced resistance offers the prospect of broad spectrum, long last-ing and, hopefully, environmentally benign disease control. However, this prospect will notbe realized unless we are able to translate our ever increasing understanding of the cellularbasis of induced resistance to the practical, field situation. This requires integration ofmolecular biology and biochemistry, with crop science and ecology. In this book, our aimis to provide plant pathologists, crop protectionists, agronomists and others with an updateof the broad and complex topic that is induced resistance and to highlight the efforts beingmade to provide the understanding necessary to allow induced resistance to be used inpractice. The various chapters in the book cover the cellular aspects of induced resistance,including signalling and defence mechanisms, the trade-offs associated with the expressionof induced resistance, work on integrating induced resistance into crop protection practiceand induced resistance from a commercial perspective. Our hope is that this book willexcite the interest of plant and crop scientists and encourage the collaboration betweenmolecular biologists, plant pathologists and ecologists that will be necessary to realize thegreat potential offered by induced resistance.

    Dale Walters, Adrian Newton and Gary Lyon


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  • Chapter 1

    Introduction: definitions and some history

    Ray HammerschmidtDepartment of Plant Pathology, Michigan State University, East Lansing, USA

    1.1 Induced resistance: an established phenomenon

    It is very well established that certain types of infection or other treatments can inducedisease resistance (e.g. Kuc, 1982; Hammerschmidt & Kuc, 1995; Sticher et al., 1997;Vallad & Goodman, 2004; da Rocha & Hammerschmidt, 2005). The induced plant is ableto resist attack by virulent pathogens and other p...


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