Bioenergetics of Aquatic Animals Bioenergetics of Aquatic Animals ALBERT LUCAS Translator: Dr Jennifer J.Watson Translation Editor: Dr I.G.Priede Department of Zoology University of Aberdeen Scotland, UK UK Taylor & Francis Ltd, 1 Gunpowder Square, London EC4A 3DE USA Taylor & Francis Inc., 1900 Frost Road, Suite 101, Bristol, PA 19007 This edition published in the Taylor & Francis e-Library, 2002. This English edition is translated from the original French edition Bioénergétique des Animaux Aquatiques, © Masson, S.A., Paris 1992. © English edition, Taylor & Francis 1996 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, electrostatic, magnetic tape, mechanical, photocopying, recording or otherwise, without the prior permission of the copyright owner. British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library. ISBN 0-7484-0564-X (cased) (formerly ISBN 013 225905 2) (Print Edition) Library of Congress Cataloguing in Publication data are available Cover design by Youngs Design in Production ISBN 0-203-48350-2 Master e-book ISBN ISBN 0-203-79174-6 (Glassbook Format) Dedication Albert Lucas died suddenly at his home in Brest shortly before the completion of the English version of this book. He had been not only an excellent teacher, but also a versatile and competent naturalist. An unselfish and wise man, he worked with several generations of students who are now involved with fundamental and applied fields of marine biology all over the world. This book will remain an enduring legacy to his students and all his friends. Yves Le Gal March, 1996 Deputy Director, Marine Biology Laboratory, College de France, Concarneau Contents T r a n s la to r s ’ N o te pag e ix Preface xi Introduction 1 1 Physical Concepts of Bioenergetics 3 1.1 The basis of thermodynamics 3 1.2 First law of thermodynamics 5 1.3 Second law of thermodynamics 7 1.4 Present understanding of thermodynamics 10 1.5 Application of thermodynamics to living organisms 12 1.6 Physical measurements used in bioenergetics 14 1.7 Comments and clarification 19 2 Cellular Bioenergetics 21 2.1 The means of cellular work 21 2.2 The conditions of cellular work 27 2.3 Types of cellular work 29 2.4 The evaluation of cellular work 34 3 Bioenergetics of Organisms: Concepts 40 3.1 Analysis of metabolism 40 3.2 Energetic equilibrium equations 47 3.3 Some misconceptions to avoid 51 3.4 Time frame of energy budgets 53 3.5 Indices and efficiencies 56 3.6 Modelling of energy budgets 57 4 Bioenergetics of Organisms: Methods 65 4.1 The processing of organic matter 65 4.2 Calorimetry 67 vii Contents 4.3 Experimental methods 70 4.4 Ecological methods 76 4.5 Differences between the two measures 78 4.6 Mixed methods 79 5 Bioenergetics of Organisms: Analysis of Results 82 5.1 Macrobioenergetics 82 5.2 Microbioenergetics 94 5.3 Correlations between metabolism and biomass 98 5.4 Reproductive effort 103 5.5 Factors influencing energy budgets 104 5.6 Summary of the energetics of organisms 108 6 Population Bioenergetics 110 6.1 Population characteristics 110 6.2 Demographic kinetics 117 6.3 Demographic production 120 6.4 Other elements of the energy budget 125 6.5 Comments on ecosystem energetics 130 7 The Scope of Bioenergetics Applications 133 7.1 Applied sciences in the aquatic environment 133 7.2 Production in molluscan aquaculture 137 7.3 Bioenergetics in fish farming 140 7.4 Bioenergetics and fundamental sciences 145 Conclusion 147 Appendix: Symbols 149 References 153 Index 165 viii Translators’ Note This work has been translated in a way that is intended to retain, as far as possible, the accuracy, meaning and scientific content of all descriptions and explanations. Thus, where appropriate, slight adaptations of the original text have been made, in order to ensure that the English version reads as fluently as possible. J.J.Watson I.G.Priede ix