Table Of ContentZoophysiology 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
