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Induced resistance offers the prospect of broad spectrum, long-lasting and potentially environmentally-benign disease and pest control in plants. Induced Resistance for Plant Defense 2e provides a comprehensive account of the subject, encompassing the underlying science and methodology, as well as research on application of the phenomenon in practice.

The second edition of this important book includes updated coverage of cellular aspects of induced resistance, including signalling and defenses, costs and trade-offs associated with the expression of induced resistance, research aimed at integrating induced resistance into crop protection practice, and induced resistance from a commercial perspective. Current thinking on how beneficial microbes induce resistance in plants has been included in the second edition.

The 14 chapters in this book have been written by internationally-respected researchers and edited by three editors with considerable experience of working on induced resistance. Like its predecessor, the second edition of Induced Resistance for Plant Defense will be of great interest to plant pathologists, plant cell and molecular biologists, agricultural scientists, crop protection specialists, and personnel in the agrochemical industry. All libraries in universities and research establishments where biological, agricultural, horticultural and forest sciences are studied and taught should have copies of this book on their shelves.

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Table of Contents

Title Page

Copyright

Contributors

Preface to Second Edition

Preface to First Edition

Chapter 1: Introduction: Definitions and Some History

1.1 Induced Resistance: An Established Phenomenon

1.2 Terminology and Types of Induced Resistance

1.3 A Little History

1.4 It's All About Interactions

1.5 Acknowledgements

References

Chapter 2: Agents That Can Elicit Induced Resistance

2.1 Introduction

2.2 Compounds Inducing Resistance

2.3 Redox Regulation

2.4 Elicitor Combinations and Synergism

2.5 Assays

2.6 Conclusions

References

Chapter 3: Transcriptome Analysis of Induced Resistance

3.1 Introduction

3.2 The Impact of

Arabidopsis thaliana

on Induced Resistance

3.3 Techniques Used for Studying Gene Expression

3.4 How Sequencing Helps Crop Research

3.5 Conclusion

3.6 Acknowledgements

References

Chapter 4: Signalling Networks Involved in Induced Resistance

4.1 Introduction

4.2 The SA–JA Backbone of the Plant Immune Signalling Network

4.3 SA and JA: Important Signals in Systemically Induced Defence

4.4 ISR and Priming for Enhanced Defence

4.5 Hormonal Crosstalk During Induced Defence

4.6 Outlook

4.7 Acknowledgements

References

Chapter 5: Types and Mechanisms of Rapidly Induced Plant Resistance to Herbivorous Arthropods

5.1 Introduction: Induced Resistance in Context

5.2 Comparison of the Threats Posed by Pathogens and Herbivores

5.3 Types of Induced Resistance

5.4 Establishing the Causal Basis of Induced Resistance

5.5 Arthropods as Dynamic Participants in Plant–Arthropod Interactions

5.6 Summary and Conclusions

References

Chapter 6: Mechanisms of Defence to Pathogens: Biochemistry and Physiology

6.1 Introduction

6.2 Structural Barriers

6.3 Phytoalexins

6.4 The Hypersensitive Response (HR)

6.5 Antimicrobial Proteins or Defence-Related Proteins

6.6 Conclusions

References

Chapter 7: Induced Resistance in Natural Ecosystems and Pathogen Population Biology: Exploiting Interactions

7.1 Introduction

7.2 Environmental Variability

7.3 Ecology of the Plant Environment

7.4 Environmental Parameters

7.5 Plant and Pathogen Population Genetics

7.6 Consequences of Resistance Induction

7.7 Conclusions

7.8 Acknowledgements

References

Chapter 8: Microbial Induction of Resistance to Pathogens

8.1 Introduction

8.2 Resistance Induced by Plant Growth Promoting Rhizobacteria and Fungi

8.3 Induction of Resistance by Biological Control Agents

8.4 Resistance Induced by Composts

8.5 Disease Control Provided by Endophytes

8.6 Arbuscular Mycorrhizal Symbiosis and Induced Resistance

8.7 Acknowledgements

References

Chapter 9: Trade-offs Associated with Induced Resistance

9.1 Introduction

9.2 Resistance Inducers

9.3 Costs of Induced Resistance

9.4 Outlook

References

Chapter 10: Topical Application of Inducers for Disease Control

10.1 Introduction

10.2 Biotic Inducers

10.3 Abiotic Inducers

10.4 Conclusions

10.5 Acknowledgements

References

Chapter 11: How do Beneficial Microbes Induce Systemic Resistance?

11.1 Plant-Beneficial Microbes

11.2 The Plant Immune System as a Regulator of Plant–Biotic Interactions

11.3 How do Beneficial Microbes Cope with the Plant Immune System?

11.4 The ISR Paradox: Local Suppression of Immunity Leads to Systemic Resistance

11.5 Concluding Remarks and Future Directions

References

Chapter 12: Implementation of Induced Resistance for Crop Protection

12.1 Introduction

12.2 Induced Resistance for Disease Control

12.3 Induced Resistance for Postharvest Disease Control

12.4 Compatibility of Activators with Other Control Methods

12.5 Influence of Genotype, Environment and Management Practices on Induced Resistance

12.6 Integration of Plant Activators in Crop Management

12.7 Challenges and Future Directions

12.8 Conclusions

References

Chapter 13: Exploitation of Induced Resistance: A Commercial Perspective

13.1 Introduction

13.2 Science and Serendipitous Discovery of Resistance-Inducing Compounds

13.3 Discovery of INAs and BTHs

13.4 Identification of BION® and Other SAR Activators

13.5 The Role of Basic Studies in the Discovery of BION® and other SAR/ISR Products

13.6 Identification of Harpin

13.7 Extracts from

Reynoutria sachalinensis

13.8 The Commercial Development of an Induced Resistance Product

13.9 Legislative Framework

13.10 Commercial Experiences with Induced Resistance Products

13.11 Conclusions

References

Chapter 14: Induced Resistance in Crop Protection: The Future, Drivers and Barriers

14.1 Introduction

14.2 Strategies to Increase Efficacy and Durability in the Field

14.3 What Research is Required to Make Induced Resistance Work in Practice?

14.4 Can We Breed Plants with Enhanced Responsiveness to Inducers?

14.5 The Potential for GM Plants Containing SAR-related Genes

14.6 Political, Economic and Legislation Issues

14.7 Conclusion

14.8 Acknowledgements

References

Index

End User License Agreement

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Guide

Cover

Table of Contents

Preface to Second Edition

Begin Reading

List of Illustrations

Figure 6.1

Figure 7.1

Figure 9.1

Figure 9.2

Figure 11.1

Figure 12.1

Figure 12.2

Figure 13.1

List of Tables

Table 3.1

Table 9.1

Table 9.2

Table 10.1

Table 13.1

Induced Resistance for Plant Defense

A Sustainable Approach to Crop Protection

Second Edition

Edited by

Dale R. Walters

Crop and Soil Systems Research Group, SRUC, Edinburgh, UK

Adrian C. Newton

James Hutton Institute, Invergowrie, Dundee, UK

Gary D. Lyon

Invergowrie, Dundee, UK

This edition first published 2014 © 2014 by John Wiley & Sons, Ltd

First edition © 2007 Blackwell Publishing

Registered office: John Wiley & Sons, Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK

Editorial offices: 9600 Garsington Road, Oxford, OX4 2DQ, UK

The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK

111 River Street, Hoboken, NJ 07030-5774, USA

For details of our global editorial offices, for customer services and for information about how to apply for permission to reuse the copyright material in this book please see our website at www.wiley.com/wiley-blackwell.

The right of the author to be identified as the author of this work has been asserted in accordance with the UK 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.

Designations used by companies to distinguish their products are often claimed as trademarks. All brand names and product names used in this book are trade names, service marks, trademarks or registered trademarks of their respective owners. The publisher is not associated with any product or vendor mentioned in this book.

Limit of Liability/Disclaimer of Warranty: While the publisher and author(s) have used their best efforts in preparing this book, they make no representations or warranties with respect to the accuracy or completeness of the contents of this book and specifically disclaim any implied warranties of merchantability or fitness for a particular purpose. It is sold on the understanding that the publisher is not engaged in rendering professional services and neither the publisher nor the author shall be liable for damages arising herefrom. If professional advice or other expert assistance is required, the services of a competent professional should be sought.

Library of Congress Cataloging-in-Publication Data

Induced resistance for plant defense : a sustainable approach to crop protection / edited by Dale R. Walters, Adrian C. Newton, Gary D. Lyon.—Second edition.

pages cm

Includes bibliographical references and index.

ISBN 978-1-118-37183-1 (cloth)

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 (Gary D.)

SB750.I4745 2015

632′.3–dc23

2014015270

A catalogue record for this book is available from the British Library.

Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic books.

Cover image: Hans van Pelt and Corné Pieterse.

Contributors

Dr Emily Beardon

Department of Animal and Plant Sciences, University of Sheffield, Western Bank, Sheffield S10 2TN, UK

Email:

[email protected]

Dr Alison E. Bennett

James Hutton Institute, Invergowrie, Dundee DD2 5DA, UK

E-mail :

[email protected]

Dr Jean-Luc Cacas

Université de Bourgogne, UMR 1347 Agroécologie 1347, Pôle Mécanismes et gestion des interactions Plantes-Micro-organismes, ERL CNRS6300, 17 rue Sully, F-21000 Dijon, France

Dr Elizabeth Dann

Queensland Alliance for Agriculture and Food Innovation, University of Queensland, St Lucia, Queensland 4072, Australia

Email:

[email protected]

Professor Brian Deverall

Faculty of Agriculture and Environment, University of Sydney, Sydney, New South Wales 2006, Australia

Dr Christophe Garcion

INRA, Univ. Bordeaux, UMR 1332 BFP, F-33140 Villenave d'Ornon, France

Dr Patrice Halama

Laboratoire BioGAP, GIS PhyNoPi, Institut Supérieur d'Agriculture de Lille, 48 Boulevard Vauban, 59800 Lille, France

Professor Ray Hammerschmidt

Department of Plant, Soil and Microbial Science, Michigan State University, East Lansing, MI 48824, USA

E-mail:

[email protected]

Professor Martin Heil

Departamento de Ingeniería Genética, CINVESTAV – Irapuato, Km. 9.6 Libramiento Norte, Irapuato, Guanajuato, CP 36821, Mexico

E-mail:

[email protected]

Dr Kemal Kazan

Commonwealth Scientific and Industrial Research, Organisation Plant Industry, Queensland Bioscience Precinct, St. Lucia, Queensland 4067, Australia

E-mail:

[email protected]

Dr Brendan Kidd

Commonwealth Scientific and Industrial Research, Organisation Plant Industry, Queensland Bioscience Precinct, St. Lucia, Queensland, 4067, Australia, and School of Agriculture and Food Sciences, University of Queensland, St. Lucia, Queensland, 4072, Australia

E-mail:

[email protected]

Dr Olivier Lamotte

CNRS, UMR 1347 Agroécologie 1347, Pôle Mécanismes et gestion des interactions Plantes-Micro-organismes, ERL CNRS6300, 17 rue Sully, F-21000 Dijon, France

Mr Andy Leadbeater

Syngenta Crop Protection AG, Schwarzwaldallee 215, 4058 Basel, Switzerland

E-mail:

[email protected]

Dr Gary D. Lyon

12 Greystane Road, Invergowrie, Dundee DD2 5JQ, UK

E-mail:

[email protected]

Professor Jean-Pierre Métraux

Département de Biologie, Université de Fribourg, 1700 Fribourg, Switzerland

E-mail:

[email protected]

Professor Adrian C. Newton

James Hutton Institute, Invergowrie, Dundee DD2 5DA, UK

E-mail:

[email protected]

Professor Corné M.J. Pieterse

Plant-Microbe Interactions, Institute of Environmental Biology, Utrecht University, P.O. Box 800.56, 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 Béatrice Randoux

Unité de Chimie Environnementale et Interactions sur le Vivant (UCEIV), GIS

PhyNoPi, Université du Littoral Côte d'Opale (ULCO), Université du Lille-Nord de France, CS 80699, F-62228 Calais Cedex, France

Dr Tony Reglinski

The New Zealand Institute for Plant and Food Research Ruakura, Private Bag 3123, Waikato Mail Centre, Hamilton 3240, New Zealand

Email:

[email protected]

Professor Philippe Reignault

Unité de Chimie Environnementale et Interactions sur le Vivant (UCEIV), GIS PhyNoPi, Université du Littoral Côte d'Opale (ULCO), Université du Lille-Nord de France, CS 80699, F-62228 Calais Cedex, France

Email:

[email protected]

Dr Peer M. Schenk

School of Agriculture and Food Sciences, University of Queensland, St. Lucia, Queensland 4072, Australia

E-mail:

[email protected]

Professor Julie Scholes

Department of Animal and Plant Sciences, University of Sheffield, Western Bank, Sheffield S10 2TN, UK

Email:

[email protected]

Dr Ali Siah

Laboratoire BioGAP, GIS PhyNoPi, Institut Supérieur d'Agriculture de Lille, 48 Boulevard Vauban, 59800 Lille, France

Dr Theo Staub

Syngenta Crop Protection AG, Schwarzwaldallee 215, 4058 Basel, Switzerland

Professor Michael J. Stout

Department of Entomology, 404 Life Sciences Building, Louisiana State University, Baton Rouge, LA 70803-1710, USA

Email:

[email protected]

Dr Christine Tayeh

Unité de Chimie Environnementale et Interactions sur le Vivant (UCEIV), GIS PhyNoPi, Université Littoral Côte d'Opale (ULCO), Université Lille-Nord de France, CS 80699, F-62228 Calais Cedex, France

Dr Jurriaan Ton

Department of Animal and Plant Sciences, University of Sheffield, Western Bank, S10 2TN, Sheffield S10 2TN, UK

Email:

[email protected]

Dr Dieuwertje Van der Does

Plant-Microbe Interactions,

Institute of Environmental Biology,

Utrecht University, P.O. Box 800.56, 3508 TB Utrecht, The Netherlands

Dr Saskia C.M. Van Wees

Plant-Microbe Interactions, Institute of Environmental Biology, Utrecht University, P.O. Box 800.56, 3508 TB Utrecht, The Netherlands

Professor Dale R. Walters

Crop and Soil Systems Research Group, SRUC, King's Buildings,

West Mains Road, Edinburgh EH9 3JG, UK

E-mail:

[email protected]

Dr Christos Zamioudis

Plant-Microbe Interactions, Institute of Environmental Biology, Utrecht University, P.O. Box 800.56, 3508 TB Utrecht, The Netherlands

Preface to Second Edition

Since the first edition of this book was published in 2007, considerable advances have been made in our understanding of induced resistance and many of these are discussed in the following chapters. The development of increasingly sophisticated techniques has greatly increased our ability to dissect the complexities of plant–microbe interactions. Although our understanding of how best to use induced resistance in crop protection practice lags behind the more fundamental aspects of induced resistance research, progress is being made. Indeed, interest in induced resistance has probably never been so great, stimulated by changes in legislation, especially in Europe, and the withdrawal of many pesticides from use. Coupled with the increasingly held view that crop protection should exert minimal impact on the environment, induced resistance, based on enhancing the plant's own defences, seems set to move away from the side-lines of crop protection.

Those of us who are fortunate enough to work on induced resistance will be aware of the pioneering contributions made by Professor Joe Kuć, who died in 2012. His painstaking and innovative research, coupled with his great enthusiasm for the topic, was instrumental in laying a solid foundation for future work in the area.

Dale R. WaltersAdrian C. NewtonGary D. LyonDecember 2013

Preface to First Edition

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 diseases: 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 essential therefore that we keep one (or preferably several) steps ahead of the pathogens.

In their review of global food security, Strange and 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 quality, without harming our fragile and long suffering environment.

Although the first recorded observations of induced resistance date back to the 19th century, 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 Kuć 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 lasting 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 R. WaltersAdrian C. NewtonGary D. Lyon

Chapter 1Introduction: Definitions and Some History

Ray Hammerschmidt

Department of Plant, Soil and Microbial Science, Michigan State University, East Lansing, MI, USA

1.1 Induced Resistance: An Established Phenomenon

Certain types of pathogen infection, non-pathogen interaction or other treatments are known to induce localized and systemic disease resistance (e.g. Kuć, 1982; Hammerschmidt and Kuć, 1995; Sticher et al., 1997; Hammerschmidt, 2009; Vallad and Goodman, 1994). The induced plant is believed to resist attack by virulent pathogens and other pests because of an enhanced ability to rapidly express defences upon infection and, in some cases, an increase in defences that are expressed in response to the inducing treatment. Although well established and studied, it is important to consider why induced resistance occurs. How can a plant that is known to be susceptible to a pathogen or even multiple pathogens be physiologically or biochemically changed so that it can now resist those infections?

Two basic assumptions must be considered to explain the overall phenomenon of induced resistance. First of all, plants must have all the genes that are necessary to mount an effective defence. Secondly, the inducing treatment should be capable of activating some of the defences directly and, more importantly, that the inducing treatment primes or sensitizes that plant in such a way that allows rapid expression of a broad set of defences upon infection by a pathogen.

The first assumption is easy to support. It is a well known plant pathology concept that plants resist the vast majority of pathogens which exist in nature, and that this phenomenon (non-host resistance) is associated with the expression of defences (Heath, 2000) and is the basis for innate immunity in plants (van Loon, 2009). Most plants, however, are susceptible to some pathogens or specific isolates or races of those pathogens. This does not mean that the plant lacks the defence needed to fend off the pathogen, but rather that the plant does not have the means to rapidly detect the presence of the pathogen (e.g. a major gene for resistance) and induce the expression of genes needed for defence. The second assumption also has significant support: plants that are induced have enhanced capacity to rapidly express defences after a challenge infection (Conrath et al., 2002).

1.2 Terminology and Types of Induced Resistance

Plant resistance to pathogens and pests can be active and/or passive (Hammerschmidt, and Nicholson, 1999). Passive resistance depends on defences that are constitutively expressed in the plant, while active resistance relies on defences that are induced after infection or attack. Induced resistance is an active process that can describe resistance at two levels. Firstly, active defence to an incompatible race or isolate of a pathogen is a form of induced resistance that is characterized by highly localized expression of defences such as phytoalexins and the hypersensitive response (Hammerschmidt and Nicholson, 1999). Secondly, induced resistance can also describe plants that express resistance to a broad range of compatible pathogens after some initial inducing treatment (Kuć, 1982). It is this latter form of induced resistance that is the focal point of this book. The term induced resistance in itself only describes the general phenomenon and does not imply any specific type of defence expression or regulation.

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