DIAPAUSE IN AQUATIC INVERTEBRATES THEORY AND HUMAN USE MONOGRAPHIAE BIOLOGICAE VOLUME 84 Series Editor H.J. Dumont Aims and Scope The Monographiae Biologicae provide a forum for top-level, rounded- off monographs dealing with the biogeography of continents or major parts of continents, and the ecology of well individualized ecosystems such as islands, island groups, mountains or mountain chains. Aquatic ecosystems may include marine environments such as coastal ecosys- tems (mangroves, coral reefs) but also pelagic, abyssal and benthic ecosystems, and freshwater environments such as major river basins, lakes, and groups of lakes. In-depth, state-of-the-art taxonomic treat- ments of major groups of animals (including protists), plants and fungi are also eligible for publication, as well as studies on the comparative ecology of major biomes. Volumes in the series may include single- author monographs, but also multi-author, edited volumes. The titles published in this series are listed at the end of this volume. Diapause in Aquatic Invertebrates Theory and Human Use VICTOR R. ALEKSEEV Zoological Institute of the Russian Academy of Science St. Petersburg, Russia BART T. DE STASIO Department of Biology Lawrence University Appleton, WI, USA and JOHN J. GILBERT Department of Biological Sciences Dartmouth College Hanover, NH, USA A C.I.P. Catalogue record for this book is available from the Library of Congress ISBN-10 1-4020-5679-6 (HB) ISBN-13 978-1-4020-5679-6 (HB) ISBN-10 1-4020-5680-X (e-book) ISBN-13 978-1-4020-5680-2 (e-book) Published by Springer, P.O. Box 17, 3300 AA Dordrecht, The Netherlands. www.springer.com Printed on acid-free paper Cover illustration: Daphnia pulicaria, photo by Victor R. Alekseev All Rights Reserved ©2007 Springer No part of this work may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording or otherwise, without written permission from the Publisher, with the exception of any material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. We dedicate this book to Professor Alexander Danilevsky TABLE OF CONTENTS Preface xiii PART I: STRATEGIES AND MECHANISMS OF DIAPAUSE IN AQUATIC INVERTEBRATES 1. Introduction to Diapause 3 Victor R. Alekseev, Oscar Ravera, and Bart T. De Stasio 1.1 Diagnosis of diapause 3 1.2 Ecological causes of diapause in aquatic organisms 5 1.3 Terminology on dormancy 8 2. Timing of Diapause in Monogonont Rotifers: Mechanisms and Strategies 11 John J.Gilbert 2.1 Introduction 11 2.2 Female types and the fertilized resting egg 12 2.3 The timing of sex: environmental controls 16 2.3.1 Preface 16 2.3.2 Crowding 17 2.3.3 Dietary tocopherol 18 2.3.4 Photoperiod 19 2.3.5 General comments 20 2.4 The timing of sex: endogenous controls 21 2.4.1 Mixis delay 21 2.4.2 Mictic stem females 23 2.5 General mechanistic models for the control of mixis 23 2.6 Theoretical models for maximizing resting-egg production 24 2.7 Diapausing parthenogenetic eggs 25 2.7.1 Preface 25 2.7.2 The pseudosexual egg 25 2.7.3 The diapausing amictic egg of Synchaeta pectinata 26 Acknowledgments 27 vii viii TABLE OF CONTENTS 3. Diapause in Crustaceans: Peculiarities of Induction 29 Victor R. Alekseev 3.1 Introduction 29 3.2 Diapause in crustacean life cycles 29 3.2.1 Monocyclic species 29 3.2.2 Bicyclic and polycyclic species 30 3.2.3 Species with complicated life cycles 31 3.2.4 Species with life cycle without diapause 32 3.3 Presence of diapause among crustaceans 32 3.3.1 Embryonal diapause 34 3.3.2 Larval diapause 35 3.3.3 Adult diapause 38 3.4 Evolution of points of view on inducing factors 39 3.4.1 Embryonal diapause 39 3.4.2 Larval and adult diapause 40 3.5 Diapause as a photoperiodic response 41 3.5.1 Developmental stages in crustaceans responsible for perception of photoperiodic signal 44 3.6 Light as the source of information about the season 44 3.6.1 Peculiarities of crustaceans’ perception of photoperiodic signals 46 3.6.2 Role of photoperiod gradient in diapause induction 47 3.6.3 Geographical variability of photoperiodic reactions 48 3.7 Role of temperature and photoperiod in diapause induction 49 3.7.1 Embryonal diapause 50 3.7.2 Larval diapause 50 3.7.3 Adult diapause 51 3.8 Population density and manifestations of photoperiodic reactions 52 3.9 Food quality and diapause induction in the crustacea 55 3.10 Population polymorphism and inheritance of photoperiodic responses 56 3.10.1 Intrapopulation dimorphism for photoperiodic responses 57 3.10.2 Population polymorphism for photoperiodic responses 59 3.11 Heredity of photoperiodic responses 61 Acknowledgments 63 4. Reactivation of Diapausing Crustaceans 65 Victor R. Alekseev 4.1 Introduction 65 4.2 Patterns of reactivation processes for different types of diapause 65 4.2.1 Embryonal diapause 66 TABLE OF CONTENTS ix 4.2.2 Larval diapause 67 4.2.3 Adult diapause 69 4.3 Endogenous phase of diapause 70 4.4 Reactivation action of oxygen 75 4.5 Participation of carbon dioxide in reactivation 76 4.6 Hormonal basis of diapause 77 Acknowledgments 82 5. Diapause in Aquatic Insects, with Emphasis on Mosquitoes 83 Elena B. Vinogradova 5.1 Introduction 83 5.2 Mosquitoes (Culicidae) 85 5.2.1 Egg diapause 85 5.2.1.1 Diapause and quiescence 85 5.2.1.2 Hatching stimuli 86 5.2.1.3 Viability, drought, and cold hardiness 87 5.2.1.4 Photoperiodic and temperature induction of egg diapause 88 5.2.1.5 Diapause termination 94 5.2.2 Larval diapause 96 5.2.2.1 Syndrome of larval diapause 96 5.2.2.2 Photoperiodic and temperature induction and termination of larval diapause 97 5.2.3 Adult diapause 103 5.2.3.1 Syndrome of adult diapause 103 5.2.3.2 Photoperiod and temperature induction of adult diapause 105 5.2.3.3 Adult diapause termination 109 5.3 Other groups of aquatic insects 110 5.3.1 Chironomids (Chironomidae) 110 5.3.2 Biting midges (Ceratopogonidae) 111 5.3.3 Dragonflies (Odonata) 111 5.3.4 Heteroptera 112 5.3.5 Ephemeroptera 113 Acknowledgments 113 6. A Brief Perspective on Molecular Mechanisms of Diapause in Aquatic Invertebrates 115 Victor R. Alekseev 6.1 Introduction 115 6.2 Molecular mechanism of diapause in the nematode Caenorhabditis elegans 116 Acknowledgments 118 x TABLE OF CONTENTS PART II: THE ROLE OF DIAPAUSE IN SCIENCE AND HUMAN USES 7. Egg Bank Formation by Aquatic Invertebrates: A Bridge Across Disciplinary Boundaries 121 Bart T. De Stasio 7.1 Introduction 121 7.2 Dormancy processes 121 7.2.1 Dormancy initiation 121 7.2.2 Release from dormancy 123 7.2.2.1 Additional emergence data 123 7.2.3 Predation and infection of dormant stages 129 7.2.4 Deep burial of dormant stages 129 7.2.5 Senescence and egg viability 130 7.3 Egg bank size and dynamics 131 7.4 Creating an egg bank 131 7.5 Conclusions 133 Acknowledgments 133 8. Use of Cladoceran Resting Eggs to Trace Climate-driven and Anthropogenic Changes in Aquatic Ecosystems 135 Susanne L. Amsinck, Erik Jeppesen, and Dirk Verschuren 8.1 Introduction 135 8.2 Tracing acidification 137 8.3 Tracing eutrophication 141 8.4 Tracing fish introductions and biomanipulation 146 8.5 Tracing heavy metal pollution 149 8.6 Tracing climate change 150 8.7 Discussion and conclusion: limitations, concerns and future potentials 153 Acknowledgments 157 9. Reconstructing Microevolutionary Dynamics from Layered Egg Banks 159 Luc De Meester, Joachim Mergeay, Helen Michels, and Ellen Decaestecker 9.1 Introduction: dormant stages and the study of microevolution 159 9.2 A short survey of recent success stories 160 9.3 Pitfalls 162 9.4 Conclusions and future directions 164 Acknowledgments 166 10. Does Timing of Emergence within a Season Affect the Evolution of Post-diapause Traits? Post-diapause and Directly Developing Phenotypes of Daphnia 167 Kestutis Arbacˇauskas 10.1 Introduction 167 10.2 Daphnialife cycle 168 TABLE OF CONTENTS xi 10.3 Neonates: biochemical quality and body size 168 10.4 Physiology: respiration and starvation resistance 169 10.5 Life-history: growth, allocation, and relative fitness 170 10.6 Descendants of post-diapause and directly developing females 173 10.7 Conclusions 173 Acknowledgments 173 11. Diapause and its Consequences in the Daphnia galeata – cucullata – hyalinaSpecies Complex 177 Piet Spaak and Barbara Keller 11.1 Introduction 177 11.2 Hybridization in Daphnia 177 11.3 Genetic markers to identify parental and hybrid taxa within the D. galeata – cucullata – hyalinacomplex 178 11.4 Factors that determine sexual reproduction of parental Daphniaspecies 180 11.5 Are hybrids still produced? 181 11.5.1 Are hybrid diapausing eggs present in the sediment? 182 11.5.2 Do males and sexual females of hybridizing species temporally and spatially co-occur? 182 11.6 Taxon distribution of asexual and sexual daphnids as well as from their offspring 183 11.7 Can the sediment tell us something about past hybridization events? 184 11.8 Conclusions 185 Acknowledgments 185 12. Role of Diapause in Dispersal of Aquatic Invertebrates 187 Vadim E. Panov and Carla Caceres 12.1 Introduction 187 12.2 Mechanisms and vectors of dispersal of diapausing invertebrates 188 12.2.1 Natural vectors of dispersal 188 12.2.2 Human-mediated dispersal 189 12.3 Conclusions: generalized model of dispersal of aquatic invertebrates with prolonged diapause 193 Acknowledgments 195 13. The Role of within Trophic Level Chemical Interactions in Diapause Induction: Basic and Applied Aspects 197 Egor S. Zadereev 13.1 Introduction 197 13.2 The effect of chemical interactions on diapause induction at the individual level 198 13.3 The effect of chemical interactions on diapause induction at population and ecosystem levels 202
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