Desiccation prels 19/3/02 1:42 pm Page i Desiccation and Survival in Plants Drying Without Dying Desiccation prels 19/3/02 1:42 pm Page ii Desiccation prels 19/3/02 1:42 pm Page iii Desiccation and Survival in Plants Drying Without Dying Edited by M. Black King’s College University of London UK and H.W. Pritchard Royal Botanic Gardens, Kew Wakehurst Place UK CABI Publishing Desiccation prels 4/4/02 2:16 pm Page iv CABI Publishing is a division of CAB International CABI Publishing CABI Publishing CAB International 10 E 40th Street Wallingford Suite 3203 Oxon OX10 8DE New York, NY 10016 UK USA Tel: +44 (0)1491 832111 Tel: +1 212 481 7018 Fax: +44 (0)1491 833508 Fax: +1 212 686 7993 Email: [email protected] Email: [email protected] Web site: www.cabi-publishing.org © CAB International 2002. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechanically, by photocopying, recording or otherwise, without prior permission of the copyright owners. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data Desiccation and survival in plants : drying without dying / edited by M. Black and H.W. Pritchard. p. cm. Includes bibliographical references (p. ). ISBN 0-85199-534-9 (alk. paper) 1. Plants--Drying. 2. Plant-water relationships. 3. Plants--Adaptation. I. Black, Michael. II. Pritchard, H. W. QK870 .D57 2002 581.4--dc21 2001043835 ISBN 0 85199 534 9 Typeset in Melior by Columns Design Ltd, Reading Printed and bound in the UK by Biddles Ltd, Guildford and King’s Lynn Desiccation prels 19/3/02 1:42 pm Page v Contents Contributors vii Preface ix PART I. INTRODUCTION 1 1 Drying Without Dying 3 Peter Alpert and Melvin J. Oliver PART II. METHODOLOGY 45 2 Methods for the Study of Water Relations Under Desiccation Stress 47 Wendell Q. Sun 3 Experimental Aspects of Drying and Recovery 93 Norman W. Pammenter, Patricia Berjak, James Wesley-Smith and Clare Vander Willigen 4 Biochemical and Biophysical Methods for Quantifying Desiccation Phenomena in Seeds and Vegetative Tissues 111 Olivier Leprince and Elena A. Golovina PART III. BIOLOGY OF DEHYDRATION 147 5 Desiccation Sensitivity in Orthodox and Recalcitrant Seeds in Relation to Development 149 Allison R. Kermode and Bill E. Finch-Savage 6 Pollen and Spores: Desiccation Tolerance in Pollen and the Spores of Lower Plants and Fungi 185 Folkert A. Hoekstra 7 Vegetative Tissues: Bryophytes, Vascular Resurrection Plants and Vegetative Propagules 207 Michael C.F. Proctor and Valerie C. Pence 8 Systematic and Evolutionary Aspects of Desiccation Tolerance in Seeds 239 John B. Dickie and Hugh W. Pritchard v Desiccation prels 19/3/02 1:42 pm Page vi vi Contents PART IV. MECHANISMS OF DAMAGE AND TOLERANCE 261 9 Desiccation Stress and Damage 263 Christina Walters, Jill M. Farrant, Norman W. Pammenter and Patricia Berjak 10 Biochemistry and Biophysics of Tolerance Systems 293 Julia Buitink, Folkert A. Hoekstra and Olivier Leprince 11 Molecular Genetics of Desiccation and Tolerant Systems 319 Jonathan R. Phillips, Melvin J. Oliver and Dorothea Bartels 12 Rehydration of Dried Systems: Membranes and the Nuclear Genome 343 Daphne J. Osborne, Ivan Boubriak and Olivier Leprince PART V. RETROSPECT AND PROSPECT 365 13 Damage and Tolerance in Retrospect and Prospect 367 Michael Black, Ralph L. Obendorf and Hugh W. Pritchard Glossary 373 Taxonomic Index 383 Subject Index 401 Desiccation prels 19/3/02 1:42 pm Page vii Contributors Peter Alpert, Biology Department, University of Massachusetts, Amherst, Massachusetts 01003-5810, USA. [email protected] Dorothea Bartels, Institute of Botany, University of Bonn, Kirschallee 1, D-53115 Bonn, Germany. [email protected] Patricia Berjak, School of Life and Environmental Sciences, University of Natal, Durban 4041, South Africa. [email protected] Michael Black, Division of Life Sciences, King’s College, Franklin Wilkins Building, 150 Stamford Street, London SE1 6NN, UK. [email protected] Ivan Boubriak, The Oxford Research Unit, Open University, Foxcombe Hall, Boars Hill OX1 5HR, UK. [email protected] Julia Buitink, UMR Physiologie Moléculaire des Semences, Institut National d’Horticulture, 16 Bd Lavoisier, F49045 Angers, France. [email protected] John B. Dickie, Seed Conservation Department, Royal Botanic Gardens Kew, Wakehurst Place, Ardingly, West Sussex RH17 6TN, UK. [email protected] Jill M. Farrant, Department of Molecular and Cellular Biology, University of Cape Town, 7700, South Africa. [email protected] Bill E. Finch-Savage, Horticulture Research International, Wellesbourne, Warwick CV35 9EF, UK. bill.fi[email protected] Elena A. Golovina, Laboratory of Plant Physiology, Department of Plant Sciences, University of Wageningen, Arboretumlaan 4, 6703 BD Wageningen, The Netherlands and Timiryazev Institute of Plant Physiology, Botanicheskaya 35, Moscow 127276, Russia. [email protected] Folkert A. Hoekstra, Laboratory of Plant Physiology, Department of Plant Sciences, University of Wageningen, Arboretumlaan 4, 6703 BD Wageningen, The Netherlands. Folkert. [email protected] Allison R. Kermode, Department of Biological Sciences, Simon Fraser University, Burnaby, BC, V5A 1S6, Canada. [email protected] Olivier Leprince, UMR Physiologie Moléculaire des Semences, Institut National d’Horticulture, 16 Bd Lavoisier, F49045 Angers, France. [email protected] Ralph L. Obendorf, Seed Biology, Department of Crop and Soil Sciences, Cornell University, Ithaca, New York, USA. [email protected] Melvin J. Oliver, USDA-ARS Plant Stress and Germplasm Development Unit, 3810 4th Street, Lubbock, Texas 79415, USA. [email protected] vii Desiccation prels 19/3/02 1:42 pm Page viii viii Contributors Daphne J. Osborne, The Oxford Research Unit, Open University, Foxcombe Hall, Boars Hill OX1 5HR, UK. [email protected] Norman W. Pammenter, School of Life and Environmental Sciences, University of Natal, Durban 4041, South Africa. [email protected] Valerie C. Pence, CREW, Cincinnati Zoo and Botanical Garden, 3400 Vine Street, Cincinnati, OH 45220, USA. [email protected] Jonathan R. Phillips, Max-Planck-Institute for Plant Breeding Research, Carl-von-Linné- Weg 10, D-550829 Köln, Germany. [email protected] Hugh W. Pritchard, Seed Conservation Department, Royal Botanic Gardens Kew, Wakehurst Place, Ardingly, West Sussex RH17 6TN, UK. [email protected] Michael C.F. Proctor, School of Biological Sciences, University of Exeter, Washington Singer Laboratories, Perry Road, Exeter EX4 4QG, UK. [email protected] Wendell Q. Sun, Department of Biological Sciences, National University of Singapore, Kent Ridge Crescent, Singapore 119260. [email protected] Clare Vander Willigen, Department of Botany, University of Capetown, Private Bag, Rondebosch 7701, South Africa. [email protected] Christina Walters, USDA-ARS National Seed Storage Laboratory, 1111 South Mason Street, Fort Collins, CO 80521, USA. [email protected] James Wesley-Smith, School of Life and Environmental Sciences, University of Natal, Durban 4041, South Africa. [email protected] Desiccation prels 19/3/02 1:42 pm Page ix Preface Plant survival of desiccation as sporophytic and gametophytic tissues was last reviewed in detail in two books published in 1980. The first, by J. Levitt, on Responses of Plants to Environmental Stresses, Volume II, Water, Radiation, Salt and Other Stresses is a classic. The topic of plant water stress consumes about 200 pages and is set mainly at the introductory level but is still of sufficient detail to stimulate post-graduate researchers. Interestingly, there is no mention in Levitt’s book of desiccation sensitivity in seeds. However, this latter topic was specifically covered in another book by H.F. Chin and E.H. Roberts (Recalcitrant Seeds). At that time recalcitrant seed behaviour was something of a novelty and the book deals mainly with descriptions of germination, a listing of species with such seeds and an indication of how best to store the material in the short term. Aspects of plant desiccation have been considered in other publications dealing with the biology and biophysics of dehydration and in contributions to general works on seeds but a comprehensive treatment of desiccation and plant survival is not yet available. Since 1980 there has been a revolution in plant science as new methods of cell and molecular biology and biophysics have been applied to environmental stress, particularly in relation to desiccation tolerance. The basic level of under- standing of how plant cells cope with extreme water stress has increased tremendously and considerable effort has been made in the last 10 years to develop diagnostic markers for desiccation tolerance. At the physiological level, studies have often focused on seed material and on the responses of resurrection plants. At a more mechanistic level, model membrane systems have been used extensively, and exploration of the molecular genetics of desiccation tolerance has begun on developmental mutants, especially of seeds of crop species. These progressive but fundamental changes in approach to investigating the basis of survival of plant tissues under desiccation since the 1980s have meant that our perceptions of this subject have altered significantly. It seems particular appropriate now to take stock of these recent developments, to assess critically the importance of the experimental systems available for investigation and to ix Desiccation prels 19/3/02 1:42 pm Page x x Preface consider possible foci for future research work. This book sets out to address these issues. The Introduction surveys the topic of desiccation, and the remain- der of the book is divided into four parts, dealing with: (i) the technical back- ground to desiccation tolerance studies; (ii) the frequency and levels of dehydration stress tolerance in biological systems; (iii) mechanisms of damage and tolerance; and (iv) a brief retrospect and prospect.