Plant Reproduction Annual Plant Reviews A series for researchers and postgraduates in the plant sciences. Each volume in this annual series will focus on a theme of topical importance and emphasis will be placed on rapid publication. Editorial Board: Professor Jeremy A. Roberts (Editor-in-Chief), Plant Science Division, School of Biological Sciences, University of Nottingham, Sutton Bonington Campus, Loughborough, Leics, LEI2 5RD, UK. Professor Hidemasa Imaseki, Obata-Minami 2419, Moriyama-ku, Nagoya 463, Japan. Dr Michael McManus, Department of Plant Biology and Biotechnology. Massey University, Private Bay, Palmerston North, New Zealand. Professor Sharman D. O'Neill, Section of Plant Biology, Division of Biological Science, University of California, Davis, CA 95616-8537, USA. Professor David G. Robinson, Zellenlehre, Universitat Heidelberg, Im Neuenheimer Feld 230, D-69120 Heidelberg. Germany. Titles in the series: 1. Arabidopsis Edited by M. Anderson and J. 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. Plant Reproduction Edited by SHARMAN D. O'NEILL Section of Plant Biology Division of Biological Science University of California, Davis, USA and JEREMY A. ROBERTS Plant Science Division School of Biosciences University of Nottingham, UK CRC Press First published 2002 Copyright © 2002 Sheffield Academic Press Published by Sheffield Academic Press Ltd Mansion House, 19 Kingfield Road Sheffield SI 1 9AS. UK ISBN 1-84127-226-4 ISSN 1460-1494 Published in the U.S.A. and Canada (only) by CRC Press LLC 2000 Corporate Blvd., N.W. Boca Raton, FL 33431, U.S.A. Orders from the U.S.A. and Canada (only) to CRC Press LLC U.S.A. and Canada only: ISBN 0-8493-9791-X ISSN 1097-7570 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 or otherwise, without the prior permission of the copyright owner. This book contains information obtained from authentic and highly regarded sources. Reprinted material is quoted with permission, and sources are indicated. Reasonable efforts have been made to publish reliable data and information, but the author and the publisher cannot assume responsibility for the validity of all materials or for the consequences of their use. Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation, without intent to infringe. Printed on acid-free paper in Great Britain by Antony Rowe Ltd, Chippenham, Wiltshire British Library Cataloguing-in-Publication Data: A catalogue record for this book is available from the British Library Library of Congress Cataloging-in-Publication Data: A catalog record for this book is available from the Library of Congress Preface Reproduction is the final stage in the life cycle of a plant. The developmental changes occurring during this event result in the production of the inflores- cence and floral meristem, floral organ primordia, ovules and pollen, embryos and seeds. Each of these reproductive tissues is produced sequentially along the floral axis, reflecting the activity of the reproductive meristem as cell division affects the penultimate outcome of the genetic programme to reproduce sexually. This volume provides an overview of our understanding of plant repro- duction in a sequence that recapitulates the process of reproduction itself. The initial chapters focus on the transition to flowering and synthesize state- of-the-art knowledge of this topic from physiological, molecular and genetic perspectives. The sequence of events leading to pollen and ovule develop- ment is then described and the mechanisms by which gamete fusion leads to formation of the embryo and endosperm are considered. Self-incompatibility mechanisms are evaluated and formation of embryos in the absence of fer- tilization is discussed. The final chapter details mechanisms that regulate floral organ senescence. The volume combines the interdisciplinary perspectives of cell biology, biochemistry, physiology, molecular biology, genetics and developmental biology. Each chapter is authored by a chosen expert in the field. The next frontier will be to understand the cellular basis of developmental changes. For example, we will need to understand the molecular events that occur during cell division and the signaling events that drive developmental change, resulting in the differentiation of ovules, pollen, gametes, embryos and seeds. These frontier areas will require the application of interdisciplinary research, using advanced genomic technologies and approaches, and will require par- ticipation and collaboration of the international community of plant scientists in investigating the functions of gene networks responsible for reproductive development. Finally, with advances in the field of evolution and phylogeny, plant scientists will begin to view reproductive development from an evo- lutionary perspective. This volume is a strong foundation for these future endeavors. Sharman D. O'Neill Jeremy A. Roberts This page intentionally left blank Contributors Professor Richard Amasino Department of Biochemistry, University of Wisconsin, 433 Babcock Drive, Madison, WI 53706-1544, US A Professor Georges Bernier Laboratory of Plant Physiology, Institute of Botany, University of Liege, B22 Sart Tilman, B-4000 Liege, Belgium Dr Roy C. Brown Department of Biology, University of Lou- isiana at Lafayette, Lafayette, LA 70504- 2451, US A Mr David Chevalier Institute of Plant Biology, University of Zurich, Zollikerstrasse 107, CH-8008 Zurich, Switzerland Mr Mark Doyle Department of Biochemistry, University of Wisconsin, 433 Babcock Drive, Madison, WI 53706-1544, USA Dr Anna M. Koltunow CSIRO Plant Industry, Horticulture Unit, PO Box 350, Glen Osmond, South Aus- tralia, 5064, Australia Dr Betty E. Lemmon Department of Biology, University of Lou- isiana at Lafayette, Lafayette, LA 70504- 2451, US A Dr Jinhong Li Centre for Plant Sciences, University of Leeds, Leeds LS2 9JT, UK Dr Ed Newbigin Plant Cell Biology Research Centre, School of Botany, University of Melbourne, Vic- toria 3010, Australia Ms Hong Nguyen Department of Biology, University of Lou- isiana at Lafayette, Lafayette, LA 70504- 2451, US A viii CONTRIBUTORS Mr Nicholas Paech CSIRO Plant Industry, Horticulture Unit, PO Box 350, Glen Osmond, South Aus- tralia, 5064, Australia Dr Luis Perez-Grau J.R. Simplot Company, Plant Sciences Division, 5369 W. Irving Street, Boise, Idaho 83706, USA Dr Claire Perilleux Laboratory of Plant Physiology, Institute of Botany, University of Liege. B22 Sart Tilman, B-4000 Liege, Belgium Dr Kay Schneitz Institute of Plant Biology, University of Zurich, Zollikerstrasse 107. CH-8008 Zurich, Switzerland Mr Si-Bum Sung Department of Biochemistry, University of Wisconsin, 433 Babcock Drive, Madison. WI 53706-1544, USA Mr Patrick Sieber Institute of Plant Biology, University of Zurich, Zollikerstrasse 107. CH-8008 Zurich, Switzerland Mr Matthew R. Tucker Department of Plant Science, Waite Cam- pus, University of Adelaide, P.M.B., 1 Glen Osmond, South Australia 5064. Australia. Professor David Twell Department of Biology, University of Leicester, University Road. Leicester LEI 7RH, UK Dr Adam Vivian-Smith CSIRO Plant Industry, Horticulture Unit. PO Box 350, Glen Osmond, South Aus- tralia, 5064. Australia Professor William R. Woodson Department of Horticulture and Landscape Architecture, Purdue University, West Lafayette, IN 47907-1165, USA Contents 1 The control of flowering: do genetical and physiological 1 approaches converge? CLAIRE PERILLEUX and GEORGES BERNIER 1.1 Introduction 1 1.2 Floral meristem identity genes 3 1.3 Floral repression genes and pathway 5 1.4 Floral promotion genes and pathways 6 1.4.1 Photoperiod promotion pathway 7 .,4.2 Autonomous promotion pathway 17 1.4.3 GA promotion pathway 18 1.5 Interactions between different genes and pathways: welcome to complexity 20 1.5.1 Are the pathways linear? 20 1.5.2 Do the pathways act in parallel? 21 1.5.3 Are the pathways independent? 21 1.5.4 Where do the pathways converge? 23 1.6 Conclusions 25 Acknowledgements 25 References 26 2 The genetic control of flowering time 33 MARK DOYLE, SI-BUM SUNG and RICHARD AMASINO 2.1 Introduction 33 2.2 FLOWERING LOCUS C and FR1GIDA confer a vernalization requirement in Ambidopsis 36 2.3 Genes in the autonomous pathway of floral promotion 38 2.4 FLC as a key regulator for the competence to flower 40 2.5 The molecular basis of vernalization 41 2.6 Possible parallels between Arabidopsis and other species 42 2.7 Photoreceptors and flowering time 43 2.8 Photoperiodism and circadian clocks 45 2.8.1 Input genes 46 2.8.2 Central oscillator components 48 2.9 Photoperiod pathway genes downstream of the clock 50 2.10 Clocks and flowering—summary 51 2.11 Genes downstream of the major flowering pathways 51 2.12 The transition to flowering—activation of floral meristem identity genes 54 2.13 Summary 54 References 55