Plant Toxicology, Fourth Edition

Plant

Toxicology

Fourth Edition

DK1206_half-series-title 8/27/04 2:14 PM Page A

BOOKS IN SOILS, PLANTS, AND THE ENVIRONMENT

Editorial Board

Agricultural Engineering Robert M. Peart, University of Florida,

Gainesville

Animal Science Harold Hafs, Rutgers University, New Brunswick,

New Jersey

Crops Mohammad Pessarakli, University of Arizona, Tucson

Irrigation and Hydrology Donald R. Nielsen, University of California, Davis

Microbiology Jan Dirk van Elsas, Research Institute for Plant

Protection, Wageningen, The Netherlands

Plants L. David Kuykendall, U.S. Department of Agriculture,

Beltsville, Maryland

Kenneth B. Marcum, Texas A&M University,

El Paso, Texas

Soils Jean-Marc Bollag, Pennsylvania State University,

University Park, Pennsylvania

Tsuyoshi Miyazaki, University of Tokyo, Japan

Soil Biochemistry, Volume 1, edited by A. D. McLaren and G. H. Peterson

Soil Biochemistry, Volume 2, edited by A. D. McLaren and J. Skujins

Soil Biochemistry, Volume 3, edited by E. A. Paul and A. D. McLaren

Soil Biochemistry, Volume 4, edited by E. A. Paul and A. D. McLaren

Soil Biochemistry, Volume 5, edited by E. A. Paul and J. N. Ladd

Soil Biochemistry, Volume 6, edited by Jean-Marc Bollag and G. Stotzky

Soil Biochemistry, Volume 7, edited by G. Stotzky and Jean-Marc Bollag

Soil Biochemistry, Volume 8, edited by Jean-Marc Bollag and G. Stotzky

Soil Biochemistry, Volume 9, edited by G. Stotzky and Jean-Marc Bollag

DK1206_half-series-title 8/27/04 2:14 PM Page B

Organic Chemicals in the Soil Environment, Volumes 1 and 2,

edited by C. A. I. Goring and J. W. Hamaker

Humic Substances in the Environment, M. Schnitzer and S. U. Khan

Microbial Life in the Soil: An Introduction, T. Hattori

Principles of Soil Chemistry, Kim H. Tan

Soil Analysis: Instrumental Techniques and Related Procedures,

edited by Keith A. Smith

Soil Reclamation Processes: Microbiological Analyses and Applications,

edited by Robert L. Tate III and Donald A. Klein

Symbiotic Nitrogen Fixation Technology, edited by Gerald H. Elkan

Soil-–Water Interactions: Mechanisms and Applications, Shingo Iwata

and Toshio Tabuchi with Benno P. Warkentin

Soil Analysis: Modern Instrumental Techniques, Second Edition,

edited by Keith A. Smith

Soil Analysis: Physical Methods, edited by Keith A. Smith and Chris E. Mullins

Growth and Mineral Nutrition of Field Crops, N. K. Fageria, V. C. Baligar,

and Charles Allan Jones

Semiarid Lands and Deserts: Soil Resource and Reclamation,

edited by J. Skujins

Plant Roots: The Hidden Half, edited by Yoav Waisel, Amram Eshel,

and Uzi Kafkafi

Plant Biochemical Regulators, edited by Harold W. Gausman

Maximizing Crop Yields, N. K. Fageria

Transgenic Plants: Fundamentals and Applications, edited by Andrew Hiatt

Soil Microbial Ecology: Applications in Agricultural and Environmental

Management, edited by F. Blaine Metting, Jr.

Principles of Soil Chemistry: Second Edition, Kim H. Tan

Water Flow in Soils, edited by Tsuyoshi Miyazaki

Handbook of Plant and Crop Stress, edited by Mohammad Pessarakli

Genetic Improvement of Field Crops, edited by Gustavo A. Slafer

Agricultural Field Experiments: Design and Analysis, Roger G. Petersen

Environmental Soil Science, Kim H. Tan

Mechanisms of Plant Growth and Improved Productivity: Modern Approaches,

edited by Amarjit S. Basra

Selenium in the Environment, edited by W. T. Frankenberger, Jr.

and Sally Benson

Plant–Environment Interactions, edited by Robert E. Wilkinson

Handbook of Plant and Crop Physiology, edited by Mohammad Pessarakli

Handbook of Phytoalexin Metabolism and Action, edited by M. Daniel

and R. P. Purkayastha

DK1206_half-series-title 8/27/04 2:14 PM Page C

Soil–Water Interactions: Mechanisms and Applications, Second Edition,

Revised and Expanded, Shingo Iwata, Toshio Tabuchi,

and Benno P. Warkentin

Stored-Grain Ecosystems, edited by Digvir S. Jayas, Noel D. G. White,

and William E. Muir

Agrochemicals from Natural Products, edited by C. R. A. Godfrey

Seed Development and Germination, edited by Jaime Kigel and Gad Galili

Nitrogen Fertilization in the Environment, edited by Peter Edward Bacon

Phytohormones in Soils: Microbial Production and Function,

William T. Frankenberger, Jr., and Muhammad Arshad

Handbook of Weed Management Systems, edited by Albert E. Smith

Soil Sampling, Preparation, and Analysis, Kim H. Tan

Soil Erosion, Conservation, and Rehabilitation, edited by Menachem Agassi

Plant Roots: The Hidden Half, Second Edition, Revised and Expanded,

edited by Yoav Waisel, Amram Eshel, and Uzi Kafkafi

Photoassimilate Distribution in Plants and Crops: Source–Sink Relationships,

edited by Eli Zamski and Arthur A. Schaffer

Mass Spectrometry of Soils, edited by Thomas W. Boutton

and Shinichi Yamasaki

Handbook of Photosynthesis, edited by Mohammad Pessarakli

Chemical and Isotopic Groundwater Hydrology: The Applied Approach,

Second Edition, Revised and Expanded, Emanuel Mazor

Fauna in Soil Ecosystems: Recycling Processes, Nutrient Fluxes,

and Agricultural Production, edited by Gero Benckiser

Soil and Plant Analysis in Sustainable Agriculture and Environment,

edited by Teresa Hood and J. Benton Jones, Jr.

Seeds Handbook: Biology, Production, Processing, and Storage: B. B. Desai,

P. M. Kotecha, and D. K. Salunkhe

Modern Soil Microbiology, edited by J. D. van Elsas, J. T. Trevors,

and E. M. H. Wellington

Growth and Mineral Nutrition of Field Crops: Second Edition, N. K. Fageria,

V. C. Baligar, and Charles Allan Jones

Fungal Pathogenesis in Plants and Crops: Molecular Biology and Host Defense

Mechanisms, P. Vidhyasekaran

Plant Pathogen Detection and Disease Diagnosis, P. Narayanasamy

Agricultural Systems Modeling and Simulation, edited by Robert M. Peart

and R. Bruce Curry

Agricultural Biotechnology, edited by Arie Altman

Plant–Microbe Interactions and Biological Control, edited by Greg J. Boland

and L. David Kuykendall

Handbook of Soil Conditioners: Substances That Enhance the Physical

Properties of Soil, edited by Arthur Wallace and Richard E. Terry

DK1206_half-series-title 8/27/04 2:14 PM Page D

Environmental Chemistry of Selenium, edited by William T. Frankenberger, Jr.,

and Richard A. Engberg

Principles of Soil Chemistry: Third Edition, Revised and Expanded, Kim H. Tan

Sulfur in the Environment, edited by Douglas G. Maynard

Soil–Machine Interactions: A Finite Element Perspective, edited by Jie Shen

and Radhey Lal Kushwaha

Mycotoxins in Agriculture and Food Safety, edited by Kaushal K. Sinha

and Deepak Bhatnagar

Plant Amino Acids: Biochemistry and Biotechnology, edited by Bijay K. Singh

Handbook of Functional Plant Ecology, edited by Francisco I. Pugnaire

and Fernando Valladares

Handbook of Plant and Crop Stress: Second Edition, Revised and Expanded,

edited by Mohammad Pessarakli

Plant Responses to Environmental Stresses: From Phytohormones to Genome

Reorganization, edited by H. R. Lerner

Handbook of Pest Management, edited by John R. Ruberson

Environmental Soil Science: Second Edition, Revised and Expanded,

Kim H. Tan

Microbial Endophytes, edited by Charles W. Bacon and James F. White, Jr.

Plant–Environment Interactions: Second Edition, edited by Robert E. Wilkinson

Microbial Pest Control, Sushil K. Khetan

Soil and Environmental Analysis: Physical Methods, Second Edition, Revised

and Expanded, edited by Keith A. Smith and Chris E. Mullins

The Rhizosphere: Biochemistry and Organic Substances at the Soil–Plant

Interface, Roberto Pinton, Zeno Varanini, and Paolo Nannipieri

Woody Plants and Woody Plant Management: Ecology, Safety,

and Environmental Impact, Rodney W. Bovey

Metals in the Environment, M. N. V. Prasad

Plant Pathogen Detection and Disease Diagnosis: Second Edition, Revised

and Expanded, P. Narayanasamy

Handbook of Plant and Crop Physiology: Second Edition, Revised

and Expanded, edited by Mohammad Pessarakli

Environmental Chemistry of Arsenic, edited by William T. Frankenberger, Jr.

Enzymes in the Environment: Activity, Ecology, and Applications,

edited by Richard G. Burns and Richard P. Dick

Plant Roots: The Hidden Half, Third Edition, Revised and Expanded,

edited by Yoav Waisel, Amram Eshel, and Uzi Kafkafi

Handbook of Plant Growth: pH as the Master Variable,

edited by Zdenko Rengel

Biological Control of Major Crop Plant Diseases,

edited by Samuel S. Gnanamanickam

Pesticides in Agriculture and the Environment, edited by Willis B. Wheeler

DK1206_half-series-title 8/27/04 2:14 PM Page E

Mathematical Models of Crop Growth and Yield, Allen R. Overman

and Richard Scholtz

Plant Biotechnology and Transgenic Plants,

edited by Kirsi-Marja Oksman Caldentey and Wolfgang Barz

Handbook of Postharvest Technology: Cereals, Fruits, Vegetables, Tea,

and Spices, edited by Amalendu Chakraverty, Arun S. Mujumdar,

G. S. Vijaya Raghavan, and Hosahalli S. Ramaswamy

Handbook of Soil Acidity, edited by Zdenko Rengel

Additional Volumes in Preparation

Humic Matter: Issues and Controversies in Soil and Environmental Science,

Kim H. Tan

Molecular Host Resistance to Pests, S. Sadasivam and B. Thayumanavan

DK1206_half-series-title 8/27/04 2:14 PM Page F

Marcel Dekker New York

Plant

Toxicology

Fourth Edition

Edited by

Bertold Hock

Professor of Cell Biology

and Dean of the Center of Life and Food Sciences

Technische Universität München

Freising, Germany

Erich F. Elstner

Professor and Head of the Institute of Phytopathology

Technische Universität München

Freising, Germany

DK1206_half-series-title 8/27/04 2:14 PM Page i

Although great care has been taken to provide accurate and current information,

neither the author(s) nor the publisher, nor anyone else associated with this

publication, shall be liable for any loss, damage, or liability directly or indirectly

caused or alleged to be caused by this book. The material contained herein is not

intended to provide specific advice or recommendations for any specific situation.

Trademark notice: Product or corporate names may be trademarks or registered

trademarks and are used only for identification and explanation without intent to

infringe.

Library of Congress Cataloging-in-Publication Data

A catalog record for this book is available from the Library of Congress.

ISBN: 0-8247-5323-2

This book is printed on acid-free paper.

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Marcel Dekker, 270 Madison Avenue, New York, NY 10016, U.S.A.

tel: 212-696-9000; fax: 212-685-4540

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tel: 800-228-1160; fax: 845-796-1772

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Copyright
2005 by Marcel Dekker. All Rights Reserved.

Neither this book nor any part may be reproduced or transmitted in any form or by

any means, electronic or mechanical, including photocopying, microfilming, and

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Current printing (last digit):

10 9 8 7 6 5 4 3 2 1

PRINTED IN THE UNITED STATES OF AMERICA

Preface

Plant toxicology is dealing with poisons causing harmful effects in plants.

Not only humans and animals, but also plants are affected by a multitude of

toxins. Therefore plant toxicology is concerned with damages, which are

caused by toxic agents, either accidentally or deliberately. Considering the

growing number of environmental compounds interfering with plant

metabolism and development, and keeping in mind the role of plants as

primary producers of food, it is surprising that the term toxicology has been

confined almost exclusively to humans and animals. If recent damages such

as forest diebacks are taken into account, it is clear that plant toxicology

represents an important branch in biological sciences.

Understanding of toxic processes in plants requires a detailed

knowledge of molecular events when toxic compounds as well as elicitors

during host–pathogen interactions bind to their molecular targets. However,

toxicology involves the entire series of phases that are relevant for the toxic

process, i.e. exposure to toxic material, uptake, distribution, metabolism

and finally secretion. These topics are central to this book.

Another focus is the recognition and possible prevention of damage,

caused by environmental pollutants. Quantification of damage is therefore

crucial. But plant toxicology also deals with negative effects, which are

intended. Agriculture and horticulture provide many examples, such as the use

of herbicides. Questions concerning the uptake, metabolism and detoxification

have to be solved before suitable and justifiable applications can be considered.

Although exogenous compounds, which normally do not occur in the

metabolism of plants (xenobiotics) are central to this book, it should be

iii

noted that also endogenous compounds of the organism can become

harmful when certain thresholds are exceeded. Such effects may result from

over-fertilization and are therefore taken into account. Even physical factors

such as ionizing radiation or detrimental effects of biogenic origin such as

infestation with parasites have to be considered if damage arises similar to

the impact of xenobiotics.

The boundaries of plant toxicology are relatively wide. They are

primarily determined by practical aspects. This book should also help to

classify observed damage and, if possible, identify. The limitation on

eukaryotic plants as potential target groups will meet practical interests.

Methods of plant toxicology originate primarily from chemistry and

biochemistry. Chemical analysis provides mainly the methods, biochemical

techniques contribute to the elucidation of action mechanisms and

metabolism of toxic compounds. Progress in toxicology largely depends

on the development of new methods and techniques.

It is not sufficient to see plants as isolated organisms. The

consideration of the ecological context is an important requisite for the

evaluation and abolishment of toxic influences. Basics of biological

knowledge are essential and provided where needed.

The editors would like to thank Marcel Dekker, Inc., particularly

Theresa Stockton, who edited and guided the book throughout production.

Indeed we wish to thank the entire staff for their understanding, encourage￾ment, and practical help.

Bertold Hock

Erich F. Elstner

iv Preface

Contents

Preface iii

Contributors vii

1. Characteristics of Plant Life: Hazards from Pollutants 1

Bertold Hock and Nicola M. Wolf

2. Plant Stress: Avoidance, Adaptation, Defense 87

Harald Schempp, Susanne Hippeli, and Erich F. Elstner

3. Uptake and Transport of Xenobiotics 131

Markus Riederer

4. Air Pollution: Trace Gases as Inducers of Plant Damage 151

Harald Schempp, Susanne Hippeli, Erich F. Elstner,

and Christian Langebartels

5. Limitation of Salt Stress to Plant Growth 191

Yuncai Hu and Urs Schmidhalter

6. Mineral Element Toxicities: Aluminum and Manganese 225

Walter J. Horst, Angelika Staß, and

Marion M. Fecht-Christoffers

v

7. Herbicides 247

Carl Fedtke and Stephen O. Duke

8. Molecular Basis of Toxic Effects: Inhibition of

Cellular Pathways and Structural Components 331

K. Kramer and Bertold Hock

9. Metabolism and Elimination of Toxicants 469

K. K. Hatzios

10. Host–Pathogen Relations: Diseases Caused by Viruses,

Subviral Organisms, and Phytoplasmas 519

Bala´zs Barna and Lo´ra´nt Kira´ly

11. Interactions Between Host Plants and Fungal

and Bacterial Pathogens 555

Ingrid Heiser, Jo¨rg Durner, and Christian Langebartels

12. Allelopathy 597

Astrid Lux-Endrich and Bertold Hock

Index 621

vi Contents

Contributors

Bala´zs Barna Plant Protection Institute, Hungarian Academy of Sciences,

Budapest, Hungary

Jo¨rg Durner Institute of Biochemical Plant Pathology, National Research

Center for Environment and Health, Neuherberg, Germany

Stephen O. Duke U.S. Department of Agriculture, University, Mississippi,

U.S.A.

Erich F. Elstner Technische Universita¨t Mu¨nchen, Freising, Germany

Marion M. Fecht-Christoffers Universita¨t Hannover, Hannover, Germany

Carl Fedtke Ko¨ln, Germany

K. K. Hatzios Virginia Polytechnic Institute and State University,

Blacksburg, Virginia, U.S.A.

Ingrid Heiser Technische Universita¨t Mu¨nchen, Freising, Germany

Susanne Hippeli Technische Universita¨t Mu¨nchen, Freising, Germany

Bertold Hock Technische Universita¨t Mu¨nchen, Freising, Germany

vii

Walter J. Horst Universita¨t Hannover, Hannover, Germany

Yuncai Hu Technische Universita¨t Mu¨nchen, Freising, Germany

K. Kramer Technische Universita¨t Mu¨nchen, Freising, Germany

Lo´ra´nt Kira´ly Plant Protection Institute, Hungarian Academy of Sciences,

Budapest, Hungary

Christian Langebartels Institute of Biochemical Plant Pathology, National

Research Center for Environment and Health, Neuherberg, Germany

Astrid Lux-Endrich Technische Universita¨t Mu¨nchen, Freising, Germany

Markus Riederer Universita¨t Wu¨rzburg, Wu¨rzburg, Germany

Harald Schempp Technische Universita¨t Mu¨nchen, Freising, Germany

Urs Schmidhalter Technische Universita¨t Mu¨nchen, Freising, Germany

Angelika Staß Universita¨t Hannover, Hannover, Germany

Nicola M. Wolf Technische Universita¨t Mu¨nchen, Freising, Germany

viii Contributors