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19.5 mm 244 x 170 mm Discover the latest developments in herbicide and weed biology COBB In the newly revised Third Edition of Herbicides and Plant Physiology, distinguished researcher Professor Dr. Andrew H. Cobb delivers an insightful and comprehensive exam- H HERBICIDES ination of the interaction between herbicides and plant physiology. The book discusses E many of the advances in plant physiology, utilizing data from the Arabidopsis genome, and R gene editing techniques that have occurred in the last dozen years. AND PLANT B This latest edition includes a variety of new and recent references addressing the latest I C developments in plant research. In addition to a complete introduction to weed biology, the PHYSIOLOGY I book discusses the modern plant protection industry and the processes by which herbicides D are discovered and developed. E S Readers will find discussions of new targets for the future development of new herbicides, as well as the mechanisms by which modern herbicides interact with plants and achieve their A ANDREW H. COBB weed control objectives. N The book also offers: D THIRD EDITION • Thorough introductions to weed biology, the modern plant protection products P industry, and how herbicides are discovered and developed L • Comprehensive explorations of how herbicides gain entry into the plant and move A to their sites of action, as well as the basis of herbicide selectivity N • Practical discussions of how herbicides interact with the major physiological T processes in plants and accomplish weed control, including the inhibition of photosynthesis, pigment biosynthesis, and more P • Reviews recent developments following the use of genetically modified-herbicide H resistant crops Y S Perfect for plant biologists and agricultural scientists, this latest edition of Herbicides and I Plant Physiology is an indispensable resource for anyone seeking a comprehensive and O robust treatment of the latest advances in plant physiology and herbicide action. L O Andrew H. Cobb, Formerly Emeritus Professor of Plant Science, Dean of Academic Affairs G and Director of Research at Harper Adams University, Shropshire, UK. Y Cover Design: Wiley THIRD Cover Image: courtesy of Dr J P H Reade EDITION www.wiley.com ffirs.indd C2hapter No.: 13 Title Name: <TITLENAME> <WORKFLOW> ff0ir1s./i1n2d/d2022 17:55:33 Comp. by: <USER> Date: 12 Jan 2022 Time: 05:55:32 pm Stage: <STAGE> WorkFlow: Page Number: ii Herbicides and Plant Physiology ffirs.indd 1 Chapter No.: 13 Title Name: <TITLENAME> <WORKFLOW> 01ff/i1rs2.i/n2d0d22 17:55:33 Comp. by: <USER> Date: 12 Jan 2022 Time: 05:55:32 pm Stage: <STAGE> WorkFlow: Page Number: i ffirs.indd C2hapter No.: 13 Title Name: <TITLENAME> <WORKFLOW> ff0ir1s./i1n2d/d2022 17:55:33 Comp. by: <USER> Date: 12 Jan 2022 Time: 05:55:32 pm Stage: <STAGE> WorkFlow: Page Number: ii Herbicides and Plant Physiology Third Edition Professor Dr Andrew H. Cobb Formerly Emeritus Professor of Plant Science, Dean of Academic Affairs and Director of Research at Harper Adams University, Shropshire, UK ffirs.indd 3 01/12/2022 17:55:33 Chapter No.: 13 Title Name: <TITLENAME> WorkFlow:<WORKFLOW> ffirs.indd Comp. by: <USER> Date: 12 Jan 2022 Time: 05:55:32 pm Stage: <STAGE> Page Number: iii This edition first published 2022 © 2022 John Wiley & Sons Ltd Edition History (1e, 1992) Andrew Cobb, published by Chapman and Hall (2e, 2010) A.H. Cobb and J.P.H. Reade, published by Wiley- Blackwell 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 law. Advice on how to obtain permission to reuse material from this title is available at http://www.wiley.com/go/permissions. The right of Andrew H. Cobb to be identified as the author of this work has been asserted in accordance with law. Registered Offices John Wiley & Sons, Inc., 111 River Street, Hoboken, NJ 07030, USA John Wiley & Sons Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK Editorial Office The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK For details of our global editorial offices, customer services, and more information about Wiley products visit us at www.wiley.com. Wiley also publishes its books in a variety of electronic formats and by print- on- demand. Some content that appears in standard print versions of this book may not be available in other formats. Limit of Liability/Disclaimer of Warranty The contents of this work are intended to further general scientific research, understanding, and discussion only and are not intended and should not be relied upon as recommending or promoting scientific method, diagnosis, or treatment by physicians for any particular patient. 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Title: Herbicides and plant physiology / Professor Dr Andrew H. Cobb, formerly Emeritus Professor of Plant Science, Dean of Academic Affairs and Director of Research at Harper Adams University, Shropshire, UK. Description: Third edition. | Hoboken, NJ : Wiley-Blackwell, [2021] | Includes bibliographical references and index. Identifiers: LCCN 2021031562 (print) | LCCN 2021031563 (ebook) | ISBN 9781119157694 (paperback) | ISBN 9781119157717 (adobe pdf) | ISBN 9781119157700 (epub) Subjects: LCSH: Plants–Effect of herbicides on. | Plant physiology. | Herbicides–Physiological effect. | Weeds–Control. Classification: LCC SB951.4 .C63 2021 (print) | LCC SB951.4 (ebook) | DDC 632/.954–dc23 LC record available at https://lccn.loc.gov/2021031562 LC ebook record available at https://lccn.loc.gov/2021031563 Cover Design: Wiley Cover Image: courtesy of Dr J P H Reade Set in 10/12 pt TimesLTStd by Straive, Pondicherry, India 10 9 8 7 6 5 4 3 2 1 ffirs.indd 4 01/12/2022 17:55:33 Chapter No.: 13 Title Name: <TITLENAME> WorkFlow:<WORKFLOW> ffirs.indd Comp. by: <USER> Date: 12 Jan 2022 Time: 05:55:32 pm Stage: <STAGE> Page Number: iv Contents Preface x 1 An Introduction to Weed Biology 1 1.1 Introduction 1 1.2 Distribution 2 1.3 The importance of weeds 3 1.4 Problems caused by weeds 3 1.5 Biology of weeds 7 1.6 A few examples of problem weeds 17 1.7 Positive attributes of weeds 19 1.8 The ever- changing weed spectrum 21 1.9 Weed control 23 References 26 2 Herbicide Discovery and Development 28 2.1 Introduction 28 2.2 Markets 29 2.3 Prospects 33 2.4 Environmental impact and relative toxicology 35 2.5 Chemophobia 40 2.6 The search for novel active ingredients 42 2.7 The search for novel target sites 46 2.8 Mode of action studies 48 2.9 The role of natural chemistry 55 2.10 Recent developments 55 2.11 A lower limit for rates of herbicide application? 58 References 59 3 Herbicide Uptake and Movement 62 3.1 Introduction 62 3.2 The cuticle as a barrier to foliar uptake 63 3.3 Physicochemical aspects of foliar uptake 67 3.4 Herbicide formulation 69 v ftoc.indd 5 01/12/2022 18:12:48 vi Contents 3.5 Uptake by roots from soil 75 3.6 Herbicide translocation from roots to shoots 77 3.7 A case study: the formulation of acids 78 3.8 The formulation of glyphosate 81 3.9 Further developments 81 References 82 4 Herbicide Selectivity and Metabolism 84 4.1 Introduction 84 4.2 General principles 85 4.3 Herbicide safeners and synergists 96 References 104 5 Herbicides that Inhibit Photosynthesis 106 5.1 Introduction 106 5.2 Photosystems 106 5.3 Inhibition at Photosystem II 108 5.4 Photodamage and repair of Photosystem II 114 5.5 Structures and uses of Photosystem II inhibitors 118 5.6 Interference with electron flow at Photosystem I 120 5.7 RuBisCo activase 126 5.8 How treated plants die 127 5.9 Chlorophyll fluorescence 131 5.10 Inhibition of photosynthetic carbon reduction in C4 plants 135 References 136 6 Inhibitors of Pigment Biosynthesis 139 6.1 Introduction: structures and functions of photosynthetic pigments 139 6.2 Inhibition of chlorophyll biosynthesis 140 6.3 Inhibition of carotenoid biosynthesis 147 6.4 Inhibition of plastoquinone biosynthesis 153 6.5 How treated plants die 156 6.6 Selectivity and metabolism 157 6.7 Summary 158 References 160 7 Auxin- type Herbicides 162 7.1 Introduction 162 7.2 Structures and uses of auxin-t ype herbicides 163 7.3 Auxin, a natural plant growth regulator 168 7.4 Biosynthesis and metabolism of auxins 170 ftoc.indd 6 01/12/2022 18:12:48 Contents vii 7.5 Auxin receptors, gene expression and herbicides 172 7.6 Signal transduction 177 7.7 Auxin transport 180 7.8 Resistance to auxin-t ype herbicides 183 7.9 An ‘auxin overdose’ 184 7.10 How treated plants die 185 7.11 Selectivity and metabolism 187 References 191 8 Inhibitors of Lipid Biosynthesis 195 8.1 Introduction 195 8.2 Structures and uses of graminicides 196 8.3 Inhibition of lipid biosynthesis 196 8.4 Activity of graminicides in mixtures 207 8.5 How treated plants die 209 8.6 Plant oxylipins: lipids with key roles in plant defence and development 211 8.7 Selectivity 213 References 216 9 The Inhibition of Amino Acid Biosynthesis 219 9.1 Introduction 219 9.2 Overview of amino acid biosynthesis in plants 219 9.3 Inhibition of glutamine synthase 220 9.4 Inhibition of aromatic amino acid biosynthesis 224 9.5 Inhibition of branch-c hain amino acid biosynthesis 231 9.6 Inhibition of histidine biosynthesis 242 References 243 10 The Disruption of Cell Division 248 10.1 Introduction 248 10.2 The cell cycle 248 10.3 Control of the cell cycle 249 10.4 Microtubule structure and function 253 10.5 Herbicidal interference with microtubules 254 10.6 Selectivity 257 References 258 11 The Inhibition of Cellulose Biosynthesis 259 11.1 Introduction 259 11.2 Cellulose biosynthesis 259 11.3 Cellulose biosynthesis inhibitors 261 11.4 How treated plants die 265 ftoc.indd 7 01/12/2022 18:12:48 viii Contents 11.5 Selectivity 265 References 266 12 Plant Kinases, Phosphatases and Stress Signalling 267 12.1 Introduction 267 12.2 Plant kinases 268 12.3 Plant phosphatases 271 12.4 Cyclin- dependent kinases and plant stress 273 12.5 Post- translational modification of proteins 276 References 278 13 Herbicide Resistance 281 13.1 Introduction 281 13.2 Definition of herbicide resistance 281 13.3 How resistance occurs 282 13.4 A chronology of herbicide resistance 283 13.5 Mechanisms of herbicide resistance 285 13.6 Case study – black- grass (A. myosuroides Huds) 300 13.7 Strategies for the control of herbicide-r esistant weeds 303 13.8 The future development of herbicide resistance 304 References 306 14 Herbicide- tolerant Crops 313 14.1 Introduction 313 14.2 History of genetically modified, herbicide- tolerant crops 314 14.3 How genetically modified crops are produced 316 14.4 Genetically engineered herbicide tolerance to glyphosate 318 14.5 Genetically modified herbicide tolerance to glufosinate 321 14.6 Genetically modified herbicide tolerance to bromoxynil 322 14.7 Genetically modified herbicide tolerance to sulphonylureas 322 14.8 Genetically modified herbicide tolerance to 2,4- D 323 14.9 Genetically modified herbicide tolerance to fops and dims 323 14.10 Genetically modified herbicide tolerance to phytoene desaturase inhibitors 323 14.11 Herbicide tolerance owing to genetic engineering of enhanced metabolism 324 14.12 Herbicide tolerance through means other than genetic modification 324 14.13 Gene editing 326 14.14 Economic, environmental and human health benefits from the adoption of GM technology 327 14.15 Gene stacking 328 14.16 Will the rise of glyphosate be inevitably followed by a fall? 329 14.17 Why is there so much opposition to GM technology? 331 14.18 Future prospects 333 References 334 ftoc.indd 8 01/12/2022 18:12:48