P088386-FM 8/23/05 6:11 PM Page i Reproductive Allocation in Plants P088386-FM 8/23/05 6:11 PM Page ii This is a volume in the PHYSIOLOGICAL ECOLOGY Series Edited by Harold A. Mooney Stanford University, Stanford, California A complete list of books in this series appears at the end of the volume. P088386-FM 8/23/05 6:11 PM Page iii Reproductive Allocation in Plants Edited by Edward G. Reekie Biology Department Acadia University Wolfville, NS, Canada Fakhri A. Bazzaz Department of Organismic and Evolutionary Biology Harvard University Cambridge, MA, USA AMSTERDAM • BOSTON • HEIDELBERG • LONDON NEW YORK • OXFORD • PARIS • SAN DIEGO SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO Academic Press is an imprint of Elsevier P088386-FM 8/23/05 6:11 PM Page iv Cover photo credit: Eric Kershaw (Kalmia inflorescence) Elsevier Academic Press 30 Corporate Drive, Suite 400, Burlington, MA 01803, USA 525 B Street, Suite 1900, San Diego, California 92101-4495, USA 84 Theobald's Road, London WC1X 8RR, UK This book is printed on acid-free paper. Copyright © 2005, Elsevier Inc. All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording, or any information storage and retrieval system, without permission in writing from the publisher. Permissions may be sought directly from Elsevier’s Science & Technology Rights Department in Oxford, UK: phone: (+44) 1865 843830, fax: (+44) 1865 853333, E-mail: [email protected]. You may also complete your request on-line via the Elsevier homepage (http://elsevier.com), by selecting “Customer Support” and then “Obtaining Permissions.” Library of Congress Cataloging-in-Publication Data Application submitted British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library ISBN 13: 978-0-12-088386-8 ISBN 10: 0-12-088386-4 For all information on all Elsevier Academic Press publications visit our Web site at www.books.elsevier.com Printed in the United States of America 05 06 07 08 09 10 9 8 7 6 5 4 3 2 1 P088386-FM 8/23/05 6:11 PM Page v Contents Contributors ix 1. The Resource Economy of Plant Reproduction 1 P. Staffan Karlsson, Marcos Méndez I. Introduction 1 II. Historical Prelude 1 III. The Principle of Allocation 3 IV. Reproductive Effort 3 A. Definitions 3 V. Problems in Determining Reproductive Allocation 5 A. The Currency 6 B. Definition of Reproductive versus Nonreproductive Plant Parts 7 C. When Should Reproductive Allocation be Measured? 8 VI. Dynamic Resource Allocation 8 VII. Empirical Patterns in Reproductive Allocation 9 A. RA and Life History 10 B. RA in Relation to Succession, Competition, and Disturbance 13 C. RA in Relation to Environmental Stress 18 D. Genetic Variation in RA 19 E. What does the Evidence Say? 20 VIII. Costs of Reproduction 20 A. Methodological Issues 21 B. Quantitative Links between Reproductive Allocation and Costs 22 IX. Conclusions 29 Acknowledgments 30 References 30 Appendix 40 2. Meristem Allocation as a Means of Assessing Reproductive Allocation 50 Kari Lehtilä, Annika Sundås Larsson I. Introduction 50 II. Developmental and Physiological Background of Meristem Allocation 51 III. Meristem Structure and Generation of Plant Architecture 52 IV. Axillary Bud Formation and Subsequent Development of the Bud 54 V. Genetics and Physiology of the Floral Transition 55 VI. Meristem Types 56 VII. Meristem Models 57 v P088386-FM 8/23/05 6:11 PM Page vi vi Contents VIII. The Assumptions of the Models 61 IX. The Impact of Meristem Allocation on Reproductive Allocation 63 X. Plasticity of Meristem Allocation 64 XI. Major Genes of Meristem Allocation 66 XII. Resource Levels and Meristem Limitation 67 XIII. The Function of Dormant Buds 68 XIV. Meristem Allocation as a Surrogate in Estimation of Resource Allocation 69 XV. Conclusions 70 Acknowledgments 71 References 71 3. It Never Rains but then it Pours:The Diverse Effects of Water on Flower Integrity and Function 75 Candace Galen I. Introduction 75 II. The Functional Ecology of Water in the Life of a Flower 76 A. Water Use by Flowers 76 B. The Water Cost of Flowers 78 C. Water as a Regulator of Flower Microclimate 79 D. Water as a Conduit for Environmental Sources of Flower Damage 79 III. Water Relations and the Evolution of Floral Traits 80 A. Floral Traits as Resource Sinks: The Resource Cost Hypothesis 80 B. Floral Traits and Water in the Flower Microclimate: Parental Environmental Effects 82 C. Plastic Responses of Floral Traits to Water Availability: Impact on Plant–Pollinator Interactions 86 IV. Conclusions 88 Acknowledgments 89 References 90 4. The Allometry of Reproductive Allocation 94 Gregory P. Cheplick I. Introduction 94 II. Definition and Analysis of RA in Relation to Allometry 96 III. Allometry Theory and RA 98 IV. Relation of RA to Relative Fitness 102 V. Allometry of Modules 105 VI. Allometry of RA and Plant Life History 109 VII. Determinants of Allometry 119 VIII. Conclusions 120 References 121 P088386-FM 8/23/05 6:11 PM Page vii Contents vii 5. Sex-specific Physiology and its Implications for the Cost of Reproduction 126 Andrea L.Case, Tia-Lynn Ashman I. Introduction 126 II. Sexual Polymorphisms 128 III. Costs of Reproduction 128 A. Male Costs 129 B. Female Costs 130 C. Common Flowering Costs 130 D. Demographic Costs 131 IV. Avenues for Mitigating the Cost of Reproduction 133 A. Photosynthetic Reproductive Organs 133 B. Increased Vegetative Photosynthesis 135 C. Increased Resource Uptake and Resource Use Efficiency 135 D. Reabsorption 136 V.Predictions for Sex-specific Physiology Based on Differential Reproductive Costs 136 A. Predictions for Females and Males 137 B. Predictions for Hermaphrodites in Monomorphic Sexual Systems: Cosexuality, Monoecy, and Diphasy 138 C. Predictions for Hermaphrodites in Dimorphic Sexual Systems: Gynodioecy and Subdioecy 139 VI. Potential Causes of Sex-specific Physiology 140 A. Physiological Differences Reflect Plastic Responses to Contrasting Reproductive Allocation between Sexes 140 B. Selection Modifies Physiological Traits after the Separation of the Sexes to Meet Differential Reproductive Costs 141 C. Physiology Changes as a Correlated Response to Selection on Other Traits (e.g., via Pleiotropy or Linkage) 141 VII. Available Data on Sex-specific Physiology 142 VIII.Recommendations for Future Study 148 Acknowledgments 148 References 148 Appendix 154 6. Time of Flowering,Costs of Reproduction, and Reproductive Output in Annuals 155 Tadaki Hirose, Toshihiko Kinugasa, Yukinori Shitaka I. Introduction 155 II. Modeling of Reproductive Output 156 III. Timing of Reproduction 158 A. Effect of Nutrient Availability 158 B. Effect of Germination Dates 160 C. Effect of Change in Flowering Time 162 P088386-FM 8/23/05 6:11 PM Page viii viii Contents IV. Costs of Reproduction 168 A. Reproductive Effort and the Relative Somatic Cost 170 B. Nitrogen Use Efficiency 172 V. Reproductive Nitrogen 176 VI. Conclusions 182 References 182 7. The Shape of the Trade-off Function between Reproduction and Growth 185 Edward G. Reekie, German Avila-Sakar I. Introduction 185 II. Methods of Describing the Trade-off Function 187 III. The Shape of the Trade-off Function in Plantago 191 IV. Impact of Reproduction on Resource Uptake 192 V. Differences in the Resource Requirements of Vegetative versus Reproductive Tissue 195 VI. Effect of Nitrogen versus Light Limitation 201 VII. Effect of Growth Pattern 202 VIII. Conclusions 207 Acknowledgments 208 References 208 8. On Size,Fecundity,and Fitness in Competing Plants 211 Lonnie W. Aarssen I. Introduction 211 II. Defining the Components of Competitive Ability for Between-species Plant Competition: Lessons from Within-species Competition 215 III. Predicting Fecundity under Competition 216 IV. Relationships among Plant Traits Affecting Fecundity under Competition: Alternative Ways to Compete Intensely While Avoiding Competitive Exclusion 220 V. Preliminary Empirical Tests 227 VI. Predicting Winners from Rank Orders in Plant Competition: Lessons from Sports Competition 231 VII. Conclusions 237 Acknowledgments 238 References 238 Index 241 Previous Volumes in Series 245 P088386-FM 8/23/05 6:11 PM Page ix Contributors Number in parentheses after each name indicates the chapter number for the author’s contribution Lonnie W. Aarssen (8) Candace Galen (3) Department of Biology Division of Biological Sciences Queen’s University 105 Tucker Hall Kingston, ON University of Missouri, Columbia K7L 3N6, Canada MO 65211, USA Tia-Lynn Ashman (5) Tadaki Hirose (6) Department of Biological Tokyo University of Agriculture Sciences Department of International University of Pittsburgh Agricultural Development Pittsburgh, PA 15217, USA Sakuragaoka 1-1-1, Setagaya-ku Tokyo 156-8502, Japan German Avila-Sakar (7) Department of Biology P. Staffan Karlsson (1) Mount Saint Vincent University Department of Plant Ecology Bedford, NS Evolutionary Biology Centre B3M 2J6, Canada Uppsala University, Villavägen 14 SE-752 36 Uppsala, Sweden Andrea L. Case (5) Department of Biological Toshihiko Kinugasa (6) Sciences Graduate School of Life Sciences Duke University Tohoku University Box No. 90338 Sendai 980-8578 Durham, NC 27708, USA Japan Gregory P. Cheplick (4) Annika S. Larsson (2) Department of Biology Department of Physiological Botany College of Staten Island Uppsala University City University of New York Villav. 6, S-752 36 Uppsala Staten Island, NY 10314, USA Sweden ix