Zoophysiology Volume 32 Editors S.D. Bradshaw W. Burggren H.C. Heller S. Ishii H. Langer G. Neuweiler D.l. Randall Zoophysiology Volumes already published in the series: Volume 1: P.l. Bentley Volume 18: E. Gwinner Endocrines and Osmoregulation Circannual Rhythms (1986) (1971) Volume 19: l. C. Ruegg Volume 2: L. Irving Calcium and Muscle Activation Arctic Life of Birds and Mammals (1986) Including Man (1972) Volume 20: l.-R. Truchot Volume 3: A.E. Needham Comparative Aspects of Extra The Significance of Zoochromes cellular Acid-Base Balance (1987) (1974) Volume 21: A. Epple and l.E. Volume 4/5: A.c. Neville Brinn Biology of the Arthropod Cuticle The Comparative Physiology of (1975) the Pancreatic Islets (1987) Volume 6: Schmidt-Koenig Volume 22: WHo Dantzler Migration and Homing in Animals Comparative Physiology of the (1975) Vertebrate Kidney (1988) Volume 7: E. Curio Volume 23: G.L. Kooyman The Ethology of Predation (1976) Diverse Divers (1989) Volume 8: W Leuthold Volume 24: S. S. Guraya African Ungulates (1977) Ovariant Follicles in Reptiles and Volume 9: E.B. Edney Birds (1989) Water Balance in Land Arthropods Volume 25: G.D. Pollak and (1977) l.H. Casseday Volume 10: H.-v. Thiele The Neural Basis of Echolocation Carabid Beetles in in Bats (1989) Their Environments (1977) Volume 26: G.A. Manley Volume 11: M.H.A. Keenleyside Peripheral Hearing Mechanisms in Diversity and Adaptation in Reptiles and Birds (1989) Fish Behaviour (1979) Volume 27: V.M. Norberg Volume 12: E. Skadhauge Vertebrate Flight (1990) Osmoregulation in Birds (1981) Volume 28: M. Nikinmaa Volume 13: S. Nilsson Vertebrate Red Blood Cells (1990) Autonomic Nerve Function in the Vertebrates (1983) Volume 29: B. Kramer Electrocommunication in Teleost Volume 14: A.D. Hasler Fishes (1990) Olfactory Imprinting and Homing in Salmon (1983) Volume 30: W Peters Volume 15: T. Mann Peri trophic Membranes (1991) Spermatophores (1984) Volume 31: M.S. Kaulenas Volume 16: P. Bouverot Insert Accessory Reproductive Adaption of Altitude-Hypoxia Structures (1992) in Vertebrates (1985) Volume 32: A.L. Val and Volume 17: R.J.F. Smith V.M.F. de Almeida-Val The Control of Fish Migration Fishes of the Amazon and their (1985) Environment (1995) A.L. Val V.M.F. de Almeida-Val Fishes of the Amazon and Their Environment Physiological and Biochemical Aspect With 76 Figures " Springer Dr. A.L. VAL Dr. Y.M.F. DE ALMEIDA-VAL Instituto Nacional de Pesquisas da Amazonia Alameda Cosme Ferreira, 1756 69083-000 Manaus, AM, Brazil Cover drawings by Izeni Pires Farias ISBN-13: 978-3-642-79231-1 e-ISBN-13: 978-3-642-79229-8 DOl: 10.1007/978-3-642-79229-8 Library of Congress Cataloging· in-Publication Data. Val, Adalberto Luis. Fishes of the Amazon and their environment: physiological and biochemical aspects/A.L. Val, V.M.F. de Almeida-Val. p. cm. - (Zoophysiology; v. 32) Includes bibliographical references (p. ) and index. 1. Fishes - Ecophysiology - Amazon River Watershed. 2. Fishes - Amazon River Watershed - Composition. 3. Fishes - Amazon River Watershed - Respiration. I. Almeida-Val, Y.M.F. de (Vera M.F.),1956- II. Title. III. Series. QL639.1.V35 1995 597'.01'09811 - dc20 95-1001 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer Verlag. Violations are liable for prosecution under the German Copyright Law. © Springer-Verlag Berlin Heidelberg 1995 Softeover reprint of the hardcover I st edition 1995 The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Typesetting: Best-set Typesetter Ltd., Hong Kong SPIN: 10426266 3113130/SPS - 5 432 10-Printed on acid-free paper To Fernando and Pedro, for the joy we have shared Preface The Amazon is a giant piece of "amphibian" land which is the result of complex geological and evolutionary processes. The number of living beings in such a land is difficult to estimate. The interactions between these organ isms and the environment are fascinating but barely understood. These features lured us to the Amazon in 1981. However, soon after, we realized that the dimensions of these interactions were overwhelming. This book is designed to review aspects of the physiology and biochemistry of fishes of the Amazon. The description of the pulsative nature of the environment and the distinct features of the ichthyofauna of the Amazon were central to the main goal. Nevertheless, any complete view is limited by the magnitude of the intraspecific variability coupled with the complex fluctuations of the environment. Thus, we have placed an emphasis on respiratory physiology and biochemistry. The reference list was made as complete as possible, particularly regarding special publications not readily available. We hope that this book is useful for comparative physiologists, tropical biologists, and the people interested in interactions between organ isms and their environment. We are grateful to many people who contributed to the making of this book. Our initial ideas were influenced by Drs. Arno Schwantes, Maria Lufza Schwantes, Jose Tundisi, Anna Emflia Vazzoler, and Naercio Menezes. New insights, particularly with respect to the respiration as an integrated system, have been learned with Profs. Peter Hochachka, David Randall, and the late Grant Bartlett. Much of the information was derived from the reports of and the discussions with our students Elizabeth Araujo, Mercia Caraciolo, Oscar Costa, Izeni Farias, Sebastiao Freire, Elizabeth Gusmao, Nadia Limeira, Jaydione Marcon, Lenise Mesquita, PHnio Monteiro, Marcia Moura, Nazare Paula, Alberto Peixoto, Hernando Ramirez, Roberto Souza, and Cacilda Yano. Prof. David Randall suggested we write this book during a meeting in Gottingen. Later, he made available to us all his facilities in Vancouver, always providing the necessary support, advice, and encouragement. As editor responsible for the book he made many valuable comments on the manuscript. Finally, we are grateful to Patricia O'Byrne for her patient help in revising the manuscript, to Izeni Pires Farias and Jorge Soares Dacio for VII their remarkable line drawings, and to Nazare Paula da Silva for her dedica tion in developing the photographs. The National Research Council of Brazil (CNPq), the National Institute of Amazon Research (INPA), and the Natural Science and Engineering Research Council of Canada (NSERC) have financially supported this work. Manaus, Rainy season 1995 A.L. VAL Y.M.F. DE ALMEIDA-VAL VIII Contents Chapter 1. Introduction ....................................... 1 Chapter 2. Amazonian Waters. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2.1 Geological Aspects of the Basin ........................ . 5 2.2 Dimensions of the Region ............................. . 7 2.3 Climate ............................................ . 8 2.3.1 Climate in the Past and in the Future ................... . 8 2.3.2 Precipitation ........................................ . 9 2.3.3 Solar Radiation ...................................... . 10 2.3.4 Temperature ........................................ . 10 2.4 The Amazon Basin ................................... . 10 2.4.1 Division of the Amazon Basin ......................... . 12 2.4.2 Basin Connections ................................... . 13 2.4.3 Types of Water ...................................... . 13 2.4.4 Chemical and Physical Characteristics of Amazon Waters .. 14 2.4.5 The Main Rivers ..................................... . 16 2.4.6 Varzea and Igapo .................................... . 17 2.4.7 Lakes .............................................. . 18 2.4.8 Annual Flood Cycle .................................. . 20 2.4.9 Metabolism in Amazon Waters ........................ . 21 2.5 Recent Modifications Produced by the Activities of Man .. . 24 2.5.1 Fish Culture in the Amazon ........................... . 26 Chapter 3. The Amazon Ichthyofauna . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 3.1 Ichthyofauna Composition ............................ . 29 3.1.1 The Elasmobranchii .................................. . 30 3.1.2 The Dipneusti ....................................... . 30 3.1.3 The Teleostei ....................................... . 31 3.1.3.1 Osteoglossiformes and Clupeiformes ................... . 31 3.1.3.2 Ostariophysi ........................................ . 32 3.1.3.3 Paracanthopterygii and Acanthopterygii ................ . 35 3.2 Feeding Behaviour ................................... . 37 3.2.1 What Do the Amazonian Fish Eat? ..................... . 39 3.2.2 When and How Much Is Eaten? ....................... . 41 IX 3.3 Migratory Cycles .................................... . 43 3.4 Breeding Patterns .................................... . 47 3.4.1 Larval Development ................................. . 50 3.5 Communication ..................................... . 50 3.6 Evolutionary and Genetic Features ..................... . 51 3.6.1 Karyotypes of Fishes of the Amazon .................... . 51 3.6.2 Isozymes ........................................... . 55 3.6.2.1 Lactate Dehydrogenase ............................... . 55 3.6.2.2 Malate Dehydrogenase ............................... . 60 3.6.3 Allozymes .......................................... . 61 3.6.4 Other Proteins ...................................... . 66 Chapter 4. Gas Exchange. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 4.1 The Obligatory Air-Breathers ......................... . 71 4.1.1 The Lungfish Lepidosiren paradoxa .................... . 75 4.1.2 Arapaima gigas ...................................... . 80 4.1.3 Electrophorus electricus ............................... . 86 4.1.4 Controversial Points .................................. . 89 4.2 The Facultative Air-Breathers ......................... . 90 4.2.1 Gills as Air-Breathing Organs ......................... . 90 4.2.2 The Stomach and Intestine as Air-Breathing Organs ...... . 95 4.2.3 Swim Bladder as an Accessory Air-Breathing Organ ...... . 98 4.3 Water-Breathing ..................................... . 100 4.3.1 Avoiding Low Oxygen Conditions ...................... . 101 4.3.2 Skimming the Water's Surface ......................... . 102 4.3.3 Improving Oxygen Transfer ........................... . 104 4.4 The Blood .......................................... . 106 4.4.1 The Erythrocytes .................................... . 106 4.4.2 The Hemoglobins .................................... . 109 4.4.2.1 Blood Levels and Solubility of Hemoglobins ............. . 109 4.4.2.2 Adjustments of Hemoglobin Proportions ................ . 111 4.4.2.3 Why So Many Hemoglobin Fractions? .................. . 113 4.4.2.4 Methemoglobin Levels ............................... . 114 4.4.3 Oxygen Affinity of Hemoglobins ....................... . 114 4.4.3.1 Effect of Temperature on Hb-02 Affinity ............... . 117 4.4.3.2 The Effect of Phosphates on Hb-02 Affinity ............ . 118 4.4.4 Oxygen Affinity of Blood ............................. . 119 4.4.5 The Root Effect ..................................... . 120 4.4.6 Intraerythrocytic Phosphates .......................... . 124 4.4.6.1 Ontogenetic Changes in the Red Blood Cell Organic Phosphate Concentrations ..................... . 124 4.4.6.2 Effects of Migration on Intraerythrocytic NTP Levels ..... . 128 4.4.6.3 Environmental Factors Affecting the Red Blood Cell Organic Phosphate Concentrations ..................... . 128 x 4.4.6.4 Interactive Effects .................................... 134 4.5 Effects of Specific Environmental Conditions on Gas Transfer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136 Chapter 5. Respiratory Metabolism ............................. 137 5.1 Optimizing Aerobic Metabolism ....................... . 138 5.1.1 Air-Breathers Versus Water-Breathers .................. . 138 5.1.2 Oxygen Consumption ................................ . 139 5.1.3 Enzyme-Level Adjustments ........................... . 141 5.1.3.1 LDH Isozymes: Channelling Pyruvate .................. . 144 5.1.3.2 Aerobic Versus Anaerobic Pathways ................... . 148 5.2 Effects of Hypoxia ................................... . 152 5.2.1 Metabolites ......................................... . 154 5.2.2 Tissue Metabolic Regulation .......................... . 156 5.3 Temperature Effects ................................. . 158 5.3.1 Isozyme Thermostability .............................. . 159 5.3.2 Km Versus Temperature .............................. . 162 5.3.3 Temperature and pH: Combined Effects ................ . 162 5.3.3.1 QIO ................................................ . 168 5.3.4 Thermal Acclimation ................................. . 169 5.3.4.1 Electrophoretic Pattern ............................... . 169 5.3.4.2 Km ................................................ . 171 5.3.4.3 Regulating Q 10 Values ................................ . 174 5.4 Metabolic Arrest .................................... . 176 5.4.1 Lungfish Estivation .................................. . 176 5.5 Effects of Specific Environmental Conditions ............ . 178 5.5.1 Accumulating End Products ........................... . 179 5.5.1.1 Lactate Accumulation ................................ . 179 5.5.2.2 Ammonia Accumulation .............................. . 180 Chapter 6. Conclusion and Perspectives. . . . . . . . . . . . . . . . . . . . . . . . . . 182 6.1 Perspectives ......................................... . 183 References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187 Systematic Index ..... ;....................................... 213 Subject Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217 XI
Description: