Seed dormancy in grasses Seed dormancy in grasses G. M. Simpson Department of Crop Science and Plant Ecology, University of Saskatchewan, Saskatoon, Canada The right of the University of Cambridge to print and sell all manner of books was granted by Henry VIII in 1534. The University has printed and published continuously since 1584. CAMBRIDGE UNIVERSITY PRESS Cambridge New York Port Chester Melbourne Sydney CAMBRIDGE UNIVERSITY PRESS Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, Sao Paulo Cambridge University Press The Edinburgh Building, Cambridge CB2 8RU, UK Published in the United States of America by Cambridge University Press, New York www. c ambridge. org Information on this title: www.cambridge.org/9780521372886 © Cambridge University Press 1990 This publication is in copyright. Subject to statutory exception and to the provisions of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. First published 1990 This digitally printed version 2007 A catalogue record for this publication is available from the British Library Library of Congress Cataloguing in Publication data Simpson, G. M. Seed dormancy in grasses / G. M. Simpson. p. cm. Bibliography: p. Includes index. ISBN 0 521 37288 7 1. Grasses-Seeds-Dormancy. I. Title. QK495.G74S613 1990 584'.90446-dc20 89-17255 CIP ISBN 978-0-521-37288-6 hardback ISBN 978-0-521-03930-7 paperback Contents Preface page vii Acknowledgements ix Introduction 1 1 The occurrence of dormancy in the Gramineae 3 1.1 Wild grasses 6 1.2 Forage grasses 7 1.3 Cereal grasses 19 1.4 Weed grasses 30 1.5 Dormancy and persistence of individuals and populations 34 1.6 Xerophytes, mesophytes and hydrophytes 42 1.7 Terminology and definitions of dormancy 43 1.8 A conceptual approach for understanding seed dormancy 48 1.9 A venafatua, a prototype model of grass seed dormancy 59 2 Mutual influences of inflorescence and caryopsis parts on dormancy 60 2.1 Position of the caryopsis on the parent plant 60 2.2 Lemma, palea, glumes and awns 65 2.3 The grain coat - pericarp and testa 83 2.4 Embryo and scutellum 94 2.5 Interrelation between embryo and maternal storage tissue 106 3 Environmental influences on seed dormancy 114 3.1 Water - liquid and vapour phases 114 3.2 Effects of radiation - heat/cold, light, microwaves, magnetism, photoperiodism 119 3.3 Latitudinal influences - tropical and temperate zones 158 3.4 Gases - oxygen and anoxia, carbon dioxide, ethylene, volatile organics; pressure 161 3.5 Growth regulators - promoters, inhibitors, limiting and allelopathic factors 170 3.6 Soil and agricultural practice 187 Contents 4 Timing of dormancy 195 4.1 Semantic considerations 195 4.2 Timing in the induction, maintenance and release of dormancy 197 5 Modelling the induction, maintenance and termination of dormancy in grass seeds 207 5.1 Modelling theory for biological systems 207 5.2 A 'black box' perspective of seed behaviour 210 5.3 An energy flow model for grass seed dormancy 218 5.4 Similarity and diversity among dormancy states in grass seeds 224 5.5 Conclusion 228 Bibliography 232 Index 281 VI Preface A number of books have been published recently on the subject of germination physiology of seeds. They often have a chapter or two about seed dormancy, either to demonstrate the diversity of mechanisms among seed plants, or to try and simplify the complexity of dormancy mechanisms by establishing general models. A somewhat different approach is used here. Firstly, the subject is confined to seed dormancy in grasses. Secondly, experimental evidence is considered in depth for a single species, the wild oat (Avena fatua L.), probably the most widely studied species for understanding seed dormancy in the plant kingdom. The evidence for this member of the family Gramineae is compared with other examples among the Gramineae to reach some general conclusions about the nature of seed dormancy in grasses. There are several reasons for confining the book to grasses. The grass family is one of the largest (25 tribes and 600 genera) and most diverse in the plant kingdom. From a human nutrition perspective it is the most important family. Grasses are the principal plant life form covering more than 70% of the land surface of the globe and they are of critical importance to the stability of the fragile arid and semi-arid zones. While seed dormancy is of great adaptive significance for survival in nearly all plant species with seeds, it is also the main reason why grass species cause the most serious weed problems in cultivated crops around the globe. It is paradoxical that one of the great achievements of the Neolithic Age was selection against seed dormancy, so that planted crops would germinate quickly and uniformly. Today we are only beginning to understand the nature of dormancy in seeds and how we might use this knowledge to better advantage. Finally, cereal grass seeds in our present age are the main source of diet for the majority of the human race. It is therefore for reasons of great practical, economic and human survival interest that we should fully vn Preface understand the trait of seed dormancy because it is a primary means for survival of populations, indeed of species, in the plant kingdom. This book is dedicated to the memory of my colleague for 25 years, the late Prof. J. M. Naylor, who was a pioneer in the field of seed dormancy studies. I am indebted to my students and research colleagues who contributed to our present understanding of seed dormancy in A. fatua. The pathway to our understanding has not been along a straight line but something akin to a much compressed helix with much energy spent in going around the circles. Vlll A cknowledgemen ts Permission for reproduction was granted by Annals of Botany Company and the editors of Annals of Botany (Figs. 3.1,3.13); Association of Applied Biologists and editors of Annals of Applied Biology (Fig. 2.S(a)); Cambridge University Press, editors of BioEssays and Dr. A. J. Trewavas (Fig. 4.1); Crop Science Society of America and editors of Crop Science (Fig. 2.6); Editorial Office of Physiologia Plantarum (Figs. 3.6, 3.7, 3.12, 3.15); editors of New Phytologist (Fig. 3.8); editor of Phytochemistry (Table 2.3); editors of Proceedings of the Association of Official Seed Analysts (Figs. 2.8(b), 2.&(c)); editors of Seed Science and Technology (Fig. 3.11); John Wiley and Sons, New York Ltd., (Symbols of Prof. H. T. Odum in Figs. 5.7, 5.8); National Research Council of Canada and editor of Canadian Journal of Botany (Figs. 2.1, 2.2(a), 2.10, 3.2, 3.3, 3.14, 3.16); Professor I. N. Morrison (Fig. 2.1 (b)) and Professor M. V. S. Raju (Fig. 22(b)). I am particularly grateful to Mr. J. Diduck for help with the figures, to Dr. F. Turel for editorial asistance and to my wife Margarete for patience with me during preparation of the manuscript. IX