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  • Plant Hormone Signaling

    Edited by

    PETER HEDDEN Rothamsted Research

    Harpenden, UK

    and

    STEPHEN G. THOMAS Rothamsted Research

    Harpenden, UK

  • This page intentionally left blank

  • Plant Hormone Signaling

  • Annual Plant Reviews A series for researchers and postgraduates in the plant sciences. Each volume in this series focuses on a theme of topical importance and emphasis is placed on rapid publication.

    Editorial Board:

    Prof. Jeremy A. Roberts (Editor-in-Chief), Plant Science Division, School of Biosciences, University of Nottingham, Sutton Bonington Campus, Loughborough, Leicestershire, LE12 5RD, UK; Dr David Evans, School of Biological and Molecular Sciences, Oxford Brookes University, Headington, Oxford, OX3 0BP; Prof. Hidemasa Imaseki, Obata-Minami 2419, Moriyama-ku, Nagoya 463, Japan; Dr Michael T. McManus, Institute of Molecular BioSciences, Massey University, Palmerston North, New Zealand; Dr Jocelyn K.C. Rose, Department of Plant Biology, Cornell University, Ithaca, New York 14853, USA.

    Titles in the series:

    1. Arabidopsis Edited by M. Anderson and J.A. Roberts 2. Biochemistry of Plant Secondary Metabolism Edited by M. Wink 3. Functions of Plant Secondary Metabolites and Their Exploitation in

    Biotechnology Edited by M. Wink 4. Molecular Plant Pathology Edited by M. Dickinson and J. Beynon 5. Vacuolar Compartments Edited by D.G. Robinson and J.C. Rogers 6. Plant Reproduction Edited by S.D. O’Neill and J.A. Roberts 7. Protein–Protein Interactions in Plant Biology Edited by M.T. McManus,

    W.A. Laing and A.C. Allan 8. The Plant Cell Wall Edited by J.K.C. Rose 9. The Golgi Apparatus and the Plant Secretory Pathway Edited by D.G. Robinson

    10. The Plant Cytoskeleton in Cell Differentiation and Development Edited by P.J. Hussey

    11. Plant–Pathogen Interactions Edited by N.J. Talbot 12. Polarity in Plants Edited by K. Lindsey 13. Plastids Edited by S.G. Moller 14. Plant Pigments and Their Manipulation Edited by K.M. Davies 15. Membrane Transport in Plants Edited by M.R. Blatt 16. Intercellular Communication in Plants Edited by A.J. Fleming 17. Plant Architecture and Its Manipulation Edited by C. Turnbull 18. Plasmodesmata Edited by K.J. Oparka 19. Plant Epigenetics Edited by P. Meyer 20. Flowering and Its Manipulation Edited by C. Ainsworth 21. Endogenous Plant Rhythms Edited by A. Hall and H. McWatters 22. Control of Primary Metabolism in Plants Edited by W.C. Plaxton and M.T. McManus 23. Biology of the Plant Cuticle Edited by M. Riederer 24. Plant Hormone Signaling Edited by P. Hedden and S.G. Thomas 25. Plant Cell Separation & Adhesion Edited by J.R. Roberts and Z. Gonzalez-Carranza 26. Senescence Processes in Plants Edited by S. Gan 27. Seed Development, Dormancy and Germination Edited by K.J. Bradford and

    H. Nonogaki

  • Plant Hormone Signaling

    Edited by

    PETER HEDDEN Rothamsted Research

    Harpenden, UK

    and

    STEPHEN G. THOMAS Rothamsted Research

    Harpenden, UK

  • © 2006 by Blackwell Publishing Ltd

    Editorial Offices: Blackwell Publishing Ltd, 9600 Garsington Road, Oxford OX4 2DQ, UK

    Tel: �44 (0)1865 776868 Blackwell Publishing Professional, 2121 State Avenue, Ames, Iowa 50014-8300, USA

    Tel: �1 515 292 0140 Blackwell Publishing Asia Pty Ltd, 550 Swanston Street, Carlton, Victoria 3053, Australia

    Tel: �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 2006 by Blackwell Publishing Ltd

    ISBN-13: 978-14051-3887-1 ISBN-10: 1-4051-3887-4

    Library of Congress Cataloging-in-Publication Data

    Plant hormone signaling / edited by Peter Hedden & Stephen Thomas. p. cm.

    Includes bibliographical references and index. ISBN-13: 978-1-4051-3887-1 (hardback : alk. paper) ISBN-10: 1-4051-3887-4 (hardback : alk. paper) 1. Plant hormones. 2. Plant cellular signal transduction.

    I. Hedden, Peter. II. Thomas, Stephen (Stephen Gregory), 1969– QK898.H67P64 2006 571.7�42—dc22

    2006004364

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

    Set in 10/12 pt, Times by Charon Tec Ltd, Chennai, India www.charontec.com Printed and bound in India by Replika Press Pvt. Ltd, Kundli

    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 web site: www.blackwellpublishing.com

  • Contents

    Contributors xiii Preface xv

    1 Abscisic acid synthesis, metabolism and signal transduction 1 ANNIE MARION-POLL and JEFFREY LEUNG

    1.1 Introduction 1 1.2 Biosynthesis and catabolism pathways 2

    1.2.1 Main early steps of ABA biosynthesis 2 1.2.2 Epoxy-carotenoid cleavage 4 1.2.3 The conversion of xanthoxin to ABA 6 1.2.4 ABA catabolism 6

    1.3 Regulation of ABA synthesis and metabolism 7 1.3.1 Developmental regulation 7

    1.3.1.1 Vegetative tissues 7 1.3.1.2 Reproductive organs 8

    1.3.2 Regulation in response to abiotic stresses 9 1.3.3 Regulation by endogenous signals and factors 10

    1.4 ABA signaling in seed maturation processes: proteolysis and combinatorial protein interactions 12

    1.5 Stress responses in vegetative tissues: the five major nexuses 15 1.5.1 ABA recognition sites and the search for the receptors 15 1.5.2 Transcriptional network as the readout 17 1.5.3 RNA metabolism 17 1.5.4 Protein phosphatases 2C 19 1.5.5 Sucrose non-fermenting-related kinases 19

    1.6 ABA signaling in guard cells: simple movements controlled by complex mechanisms 22

    1.7 ABA as antagonizing signal to light in stomatal movement 23 1.8 Concluding remarks 24

    Acknowledgements 25 References 26

    2 Auxin metabolism and signaling 37 JERRY D. COHEN and WILLIAM M. GRAY

    2.1 Introduction 37 2.2 Auxin metabolism 37

    2.2.1 Indole-3-acetic acid biosynthesis 37 2.2.1.1 The tryptophan-independent pathway 37

  • 2.2.1.2 IAA biosynthesis from tryptophan 40 2.2.2 IAA conjugates in plants 42

    2.2.2.1 IAA-peptide conjugates 42 2.2.2.2 Amino acid conjugates 43 2.2.2.3 Amide conjugate hydrolysis 43 2.2.2.4 Ester conjugates 44

    2.2.3 IAA degradation 46 2.3 Auxin signaling 46

    2.3.1 Auxin-responsive genes 46 2.3.2 Auxin response factors 47 2.3.3 Regulation of auxin response by the SCFTIR1 ubiquitin–ligase 51 2.3.4 Regulation of SCFTIR1 activity 53 2.3.5 Identification of an auxin receptor 55

    2.4 Conclusions and future perspectives 58 Acknowledgements 59 References 59

    3 Integration of brassinosteroid biosynthesis and signaling 67 MIKLOS SZEKERES and GERARD J. BISHOP

    3.1 Introduction 67 3.2 Metabolism 67

    3.2.1 Biosynthesis 67 3.2.1.1 DET2 69 3.2.1.2 SAX1 72 3.2.1.3 DWF4 72 3.2.1.4 CPD 73 3.2.1.5 ROT3 and CYP90D1 73 3.2.1.6 CYP85A1 and CYP85A2 74 3.2.1.7 Other biosynthetic functions 74

    3.2.2 Inactivation 75 3.2.2.1 BAS1 75 3.2.2.2 CHI2/SHK1/SOB7 76 3.2.2.3 UGT73C5 77 3.2.2.4 BNST3 and BNST4 77

    3.2.3 Functional aspects of BR metabolism 77 3.2.3.1 Regulation of biosynthetic genes 77 3.2.3.2 Regulation of BR-inactivating genes 78 3.2.3.3 Conservation of BR synthesis in higher plants 79

    3.3 Signal transduction 80 3.3.1 BRI1 and BAK1 80 3.3.2 BIN2 and BSU1 81 3.3.3 BZR1 and BZR2/BES1 82 3.3.4 BIM1 82 3.3.5 Signaling mechanism and other putative components 83

    3.4 Future prospectives 84 3.4.1 Metabolism 84 3.4.2 Signal transduction 86

    vi CONTENTS

  • 3.4.3 Crops 86 Acknowledgements 87 References 87

    4 Cytokinin metabolism and signal transduction 93 ALEXANDER HEYL, TOMÁS̆ WERNER and THOMAS SCHMÜLLING

    4.1 Introduction 93 4.2 Cytokinin metabolism 93

    4.2.1 Cytokinin biosynthesis 94 4.2.2 Cytokinin interconversion and conjugation 97 4.2.3 Cytokinin catabolism 98

    4.3 Cytokinin signal transduction 101 4.3.1 Cytokinin signal perception 101 4.3.2 Cytokinin signal transduction 105

    4.4 Conclusions 117 References 118

    5 Ethylene biosynthesis and signaling: a puzzle yet to be completed 125 FILIP VANDENBUSSCHE, WIM H. VRIEZEN and DOMINIQUE VAN DER STRAETEN

    5.1 Introduction 125 5.2 Ethylene biosynthesis 126

    5.2.1 ACC synthase 127 5.2.2 ACC oxidase 130

    5.3 Ethylene signal transduction 131 5.4 A complex network 137

    Acknowledgements 139 References 139

    6 Gibberellin metabolism and signal transduction 147 STEPHEN G. THOMAS and PETER HEDDEN

    6.1 Introduction 147 6.2 The gibberellin metabolic pathway 148

    6.2.1 Biosynthesis of bioactive GAs 148 6.2.2 GA deactivation 150

    6.3 Genes of GA biosynthesis and their regulation 151 6.3.1 Developmental regulation 151 6.3.2 Hormonal regulation 154 6.3.3 Environmental regulation 154

    6.4 The gibberellin signal transduction pathway 157 6.4.1 The gibberellin receptor 159 6.4.2 DELLA proteins act as repressors of GA signaling 159

    CONTENTS vii