PHYSIOLOGY OF THE CLADOCERA Mud-living Ilyocryptus colored red by hemoglobin. Photographed by A.A. Kotov. PHYSIOLOGY OF THE CLADOCERA NIKOLAI N. SMIRNOV D.SC. Institute of Ecology, Moscow, Russia WITH ADDITIONAL CONTRIBUTIONS AMSTERDAM • BOSTON • HEIDELBERG • LONDON NEW YORK • OXFORD • PARIS • SAN DIEGO SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO Academic Press is an imprint of Elsevier Academic Press is an imprint of Elsevier 32 Jamestown Road, London NW1 7BY, UK 225 Wyman Street, Waltham, MA 02451, USA 525 B Street, Suite 1800, San Diego, CA 92101-4495, USA Copyright r 2014 Elsevier Inc. 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Because of rapid advances in the medical sciences, in particular, independent verification of diagnoses and drug dosages should be made British Library Cataloguing-in-Publication Data A catalogue record for this book is available from the British Library Library of Congress Cataloging-in-Publication Data A catalog record for this book is available from the Library of Congress ISBN : 978-0-12-396953-8 For information on all Academic Press publications visit our website at elsevierdirect.com Typeset by MPS Limited, Chennai, India www.adi-mps.com Printed and bound in United States of America 1415161710987654321 Preface Over 700 species of Cladocera are known, the role of iron are also discussed. Studies on and representatives of this group are often the impact of xenobiotics on respiration are dominant in the freshwater fauna and some- reviewed and the blood flows and separating times occur in enormous quantities. They live membranes are described. Heart rate and its in both small and large water bodies from arctic controlling factors are also reviewed. Occasional to tropical latitudes, in open water, on the bot- heart arrest and adhesion of blood cells are tom, in mud, among inshore vegetation, in acid reported. Studies on phagocytosis are pointed pools on bogs, in small accumulations of water out. The data related to excretion are reviewed in epiphytic plants, in narrow aquatic spaces and the distribution of xenobiotics in the body, between moist sand grains. A few species even their bioaccumulation and the transformation left the water and live in moist moss-like of xenobiotics are described. Studies on routes growth on tree trunks in tropical cloud forests. of elimination of xenobiotics and on detoxifica- Some species are specialized for life in saline tion are indicated. lakes and in the sea. Ion exchange is realized via the neck organ The data on organic and inorganic constitu- and limb epipodites. The principal types of ents of cladocerans are reviewed. They may osmotic regulation are: hyperosmotic regulation accumulate physiologically important sub- (in freshwater species), hypoosmotic regulation stances, those of no such importance, or toxi- (in marine Penilia), and amphiosmotic cants. The section on nutrition of Cladocera regulation. The effect of xenobiotics on osmotic comprises a brief review of their anatomical regulation is discussed. Numerous enzymes and environmental backgrounds, with special have been determined in Cladocera and studies consideration of food quality (algae, algal on their role are reviewed. The impact of xeno- lipids, bacteria, and organic debris). Feeding biotics on metabolic links is described, as well and digestion are discussed for both littoral as detoxification in the cladoceran body. and pelagic species and the digestion of pro- Cladocera live for a week up to several tein, carbohydrate, and lipids is reviewed. months, depending on the species and the Cladocera modify only a small amount of the environmental conditions. Body length incre- lipids consumed with algae, and then transfer ments occur between molts. Mechanical dam- them to fish and thus further, to man. The fate age is repaired by regeneration. Effects of of ingested chlorophyll, starvation and the environmental factors and xenobiotics on mor- impact of xenobiotics on feeding and digestion phology and cytology are described and data are also commented on. on the impact of xenobiotics on molting and life Oxygen consumption and the production duration are reviewed. The state of investiga- of CO2 are considered in littoral and pelagic tions on senescence and mortality is also com- Cladocera. Littoral and bottom-dwelling spe- mented on. cies often live under conditions of hypoxia Cladocera are remarkable by their partheno- and anoxia. The dynamics of hemoglobin and genetic reproduction, which is sometimes ix x PREFACE interrupted by bisexual (gamogenetic) reproduc- xenobiotics on behavior are also reviewed. tion. Physiological aspects of parthenogenetic Ecological aspects discussed comprise consid- and gamogenetic reproduction are discussed. eration of physiological background and the Differences of movement and trajectories of litto- limits of physiological factors, synergism and ral and pelagic species are indicated and data antagonism of factors, pathway of lipids from on the physiology of muscles, immobilization, algae via Cladocera to fish, conditioning of the fatigue, stress, and impact of xenobiotics are also environment by Cladocera, as well as the use reviewed. There is information on neurosecretion, of Cladocera in water quality testing. vision, ability to discern polarized and colored Within certain limits, Cladocera can support light, photoperiod, chemoreception, mechanore- the body’s homeostasis in a dynamic environ- ception, ability for orientation in space, endoge- ment, including the homeostatic level of their nous rhythms, impact of xenobiotics. chemical constituents and of osmotic pressure. Cladocera possess complicated behavior Along with special discussions, introductory patterns, differing in particular species. These remarks are made whenever it seemed to be include migration, swarming, akinesis and necessary to make the matter useful both for escape behavior. The data on the impact of specialists and for non-specialists. Contributors Stuart K. J. R. Auld School of Natural Sciences, University of Notre Dame, Notre Dame, IN, University of Stirling, Stirling, Scotland, UK USA Margaret J. Beaton Biology Department, Mount Joseph R. Shaw School of Public and Allison University, Sackville, NB, USA Environmental Affairs, Indiana University, Do¨rthe Becker Environmental Genomics Group, Bloomington, IN, USA School of Biosciences, University of Birmingham, Nikolai N. Smirnov Institute of Ecology, Leninski Birmingham, UK Prospect 33, Moscow, Russia John K. Colbourne Environmental Genomics Group, Elizabeth Turner School of Public and School of Biosciences, University of Birmingham, Environmental Affairs, Indiana University, Birmingham, UK Bloomington, IN, USA Carli M. Peters Biology Department, Mount Allison Kay Van Damme Environmental Genomics Group, University, Sackville, NB, USA School of Biosciences, University of Birmingham, Michael E. Pfrender Department of Biological Birmingham, UK Sciences, Environmental Change Initiative, xi Acknowledgments The present volume is an attempt at making Many librarians, mostly personally unknown a summary of work of many experts through- to the author, retrieved numerous publications out the world in various fields using special from different countries and time periods. Their methods. A substantial contribution to physiol- care and labor are appreciated, including those ogy of the Cladocera is made recently by of the Biological Department Library of Russian toxicologists. Academy of Sciences, including Ms N.I. Special thanks are due to Dr M.J. Burgis (UK) Gotovskaya and Ms E.V. Morozova. who generously used her time and experience to The present study is supported in part by improve the manuscript. grants on Cladocera projects from the Russian The present review is motivated by the Foundation for Basic Rssearch (12-04-00207, etc.) author’s observations on living, mostly littoral, and from the Program “Living Nature”. cladocerans. Some recent observations are made My wife L.A. Smirnova, Ph.D. (cited here as at the Hydrobiological Station “Glubokoe Lake” L.A. Luferova) is tolerant (mostly) towards using (Russia). The author is grateful to his imme- a big part of my time for such ventures as this. diate colleagues from the ‘Cladocera team’ Formulation of ideas included in this book for help and discussions: O.S. Boikova, and its composition was much favored by N.M. Korovchinsky, A.A. Kotov, E.I. Bekker, creative environment at the Institute of Ecology A. Yu. Sinev, as well as to Dr Yu.B. Manteifel and Evolution of the Russian Academy of and Dr E.P. Zinkevich for advice. Sciences, and by personal attention of academi- Dr A.A. Kotov critically read the draft man- cian D.S. Pavlov, Director, and of academician uscript, suggested numerous useful additions, Yu.Yu. Dgebuadze,