Bl-ology Guido di Prisco Bruno Maresca Antarctic =~Tobf Fish With 105 Figures and 30 Tables Springer-Verlag Berlin Heidelberg New York London Paris Tokyo Hong Kong Barcelona Budapest Professor Dr. Guido di Prisco Institute of Protein Biochemistry and Enzymology (lBPE) Italian National Research Council Via Marconi 10,1-80125 Naples Italy Professor Dr. Bruno Maresca International Institute of Genetics and Biophysics, CNR 1-80125 Naples Italy Professor Dr. Bruno Tota University of Calabria 1-87030 Arcavacata di Rende (CS) Italy ISBN-13:978-3-642-76219-2 e-ISBN-13:978-3-642-76217-8 DOl: 10.1007/978-3-642-76217-8 Library of Congress Cataloging-in-Publication Data Biology of antarctic fish/ Guido de Prisco, Bruno Maresca, Bruno Tota, editors. p. em. Includes bibliographical references and index. 1. Marine fishes -Antarctic Ocean -Congresses. I. Di Prisco, Guido, 1937- . II. Maresca, B. (Bruno), 1950- . III. Tola, Bruno. QL637.2.B56 1991 597.092'4-dc20 91-35000 CIP 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, broad casting, 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 September9, 1965, in its current version, and permission foruse must always be obtained from Springer Verlag. Violations are liable for prosecution under the German Copyright Law. © Springer-Verlag Berlin Heidelberg 1991 Softcover reprint of the hardcover 1st edition 1991 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 protec tive laws and regulations and therefore free for general use. Product liability: The publishers cannot guarantee the accuracy of any information about dosage and application contained in this book. In every individual case the user must check such information by consulting the relevant literature. Production Editor: Martin Langner Typesetting, International Typesetters, Inc. Makati, M. M., The Philippines 31/3145 -5 43 21 0 -Printed on acid-free paper Preface Antarctica originated as a fragment of the ancient supercontinent Gondwana, which drifted to its present polar position and was isolated climatically and biologically by the establishment of a circumpolar oceanic circulation. This gave rise to a well-defmed, roughly circular ocean front: the Antarctic Convergence, During the past 25 million years, fIsh have remained in isolation south of the Convergence, where the progressive cooling of the environment became a key factor in influencing their evolution. At present, the year-round temperature of the coastal waters is very near -1.87°C, the equilibrium temperature of the ice-seawater mixture. The unique characteristics produced in antarctic fIsh in the course of their evolution during the long isolation south of the Convergence encourage the investigation of this ecosystem. Populations of complex organisms such as fIsh contain a hierarchy oflevels ofo rganization: indi viduals, organ systems, cells, organelles, membranes, molecules (in order of decreasing complexity). Indeed, the temperature of the en vironment affects each of the levels ofo rganization offlsh life. Thus, the biology of fIsh of the Antarctic Ocean, where the extremely low tem perature is a feature of the utmost importance, can be best explored when multidisciplinary approaches and efforts are combined in a general framework. The need to produce a clear, up-to-date picture ofthe present state of knowledge in a large number off Ields oft he biology off Ish belonging to this unique environment, gave rise to a strong motivation to organize a meeting along these lines. The Second International Conference: Biology of Antarctic Fishes ("Ravello 2") took place in Ravello (Italy) at the end of May 1990. It was sponsored by the Italian National Research Council and by the National "Programme for Antarctic Research. ''Ravello 2" was a follow-up of the fIrst Conference ("Ravello I", 1986), which dealt with a much broader range of topics inherent to marine biology of antarctic organisms. The Conference lectures were focused on ecology, evolution, life history, physiology, biochemistry and molecular biology of antarctic fIsh, and are all contained in this volume. Weare possibly dealing with the most comprehensive and complete contribution so far available in the fIeld. VI Preface We express our gratitude to all the speakers, thanks to whom the Conference has been so stimulating and productive. We also wish to express our sincere hope to reconvene in "Ravello 3" in the near future. NapleslItaly, October 1990 Guido di Prisco Bruno Maresca Bruno Tota The speakers at the Second International Conference: Biology of Antarctic Fishes ("Ravello 2"). From left to right: front row, Eastman, Detrich, Targett, Duhamel, Radtke; second row, Hemmingsen, Ekau, Hubold, Wells, Sidell, North, EI-Sayed, Somera, Clarke, Johnston, DeVries, Macdonald, Kellermann, accompanying person; at back, White, Tota, di Prisco, Maresca. Contents Part 1 Ecology, Evolution and Life History Ecology of Notothenioid Fish in the Weddell Sea (With 8 Figures) ............................................ 3 G.Hubold Morphological Adaptations and Mode of Life in High Antarctic Fish (With 6 Figures) ...................... 23 WEkau The Biological and Demographic Peculiarities of the Icefish Champsocephalus gunnari L6nnberg, 1905 from the Kerguelen Plateau (With 8 Figures) ................ 40 G.Duhamel Is the Growth of Polar Fish Limited by Temperature? (With 6 Figures) ............................................ 54 A. Clarke and A. W North Review of the Early Life History of Antarctic Notothenioid Fish (With 1 Figure) 70 A. WNorth Age Determination in Antarctic Fish (With 4 Figures) ......... 87 MG. White Microstructural Analysis of Growth Patterns in the Early Life History of Antarctic Fishes (With 7 Figures) .. 101 R. L. Radtke and A. Kellermann The Fossil and Modern Fish Faunas of Antarctica: Evolution and Diversity (With 3 Figures) 116 J T.Eastman The Contribution of the BIOMASS Program to Antarctic Marine Ecosystem Research (With 3 Figures) ..... 131 S. Z. El-Sayed x Contents Part 2 Physiology, Biochemistry and Molecular Biology The Sensory Biology of Notothenioid Fish (With 9 Figures) ... 145 1.A.Macdonald and 1. C.Montgomery Viscosity of Body Fluids from Antarctic Notothenioid Fish (With 9 Figures) ............. 163 1.A.Macdonald and R.M. G. Wells Low Temperature Limits Burst Swimming Performance in Antarctic Fish (With 6 Figures) ............................ 179 LA. Johnston, T.P.Johnson, and 1. C.Battram Respiratory and Cardiovascular Adaptations in Hemoglobin-Free Fish: Resolved and Umesolved Problems (With 2 Figures) .......... 191 E. A. Hemmingsen Structural and Mechanical Characteristics of the Heart of the Icefish Chionodraco hamatus (LOnnberg) (With 10 Figures) ........................................... 204 B. Tota, C.Agnisola, M.Schioppa, R.Aciemo, P. Harrison, and G. Zummo Physiological Roles of High Lipid Content in Tissues of Antarctic Fish Species (With 5 Figures) .......... 220 B. D. Sidell Biochemical Mechanisms of Cold Adaptation and Stenothermality in Antarctic Fish (With 10 Figures) ....... 232 G.N.Somero Polymerization of Microtubule Proteins from Antarctic Fish (With 6 Figures) ......................... 248 H. W. Detrich III The Biochemistry of Oxygen Transport in Red-Blooded Antarctic Fish (With 2 Figures) ............... 263 G. di Prisco, R. D'Avino, C. Caruso, M. Tamburrini, L. Camardella, B. Rutigliano, V. Ca"atore, and M. Romano Subject Index ............................................... 283 Part I Ecology, Evolution and Life History Ecology of Notothenioid Fish in the Weddell Sea G. Hubold 1 Introduction Low water temperature, and the seasonality and scarcity of food are key factors in the life of polar ectotherms (Clarke 1983). Life in cold water requires special adaptations against freezing in teleost fish (DeVries 1971), and the need to accumu late energy reserves during summer may govern the seasonal activity pattern. In addition to these typical high-latitude features, environmental factors such as water mass distribution and current patterns also affect the life of fish in the Antarctic. Hydrography becomes a critical factor when pelagic stages are included in the life history (Harden Jones 1968; Cushing 1974; Sinclair 1988). For evolutionary adaptation, stability or predictability of the environment is important. Antarctic fish occur over a latitudinal range of more than 20° in the Atlantic sector. While temperature and production decrease towards the south, stability of temperature and predictability of hydrographic features increase in the permanent pack-ice zone surrounding the continent (Hempel 1990). Environmental conditions similar to those presently found may have prevailed over the past 20 million years, although several major advances and retreats of the continental ice sheet will have had effects on the composition of the fish fauna (DeWitt 1970; Hubold and Ekau 1987; Clarke and Crame 1989). High latitude species may therefore by confronted with very severe conditions, but may also have had the possibility to develop very specialized adaptations to match these conditions in the course of evolution. Fish species from high antarctic latitu.des (e.g., McMurdo Sound) were found to have more specialized biochemical adaptations than subant arctic species (De Vries and Eastman 1981). It is therefore of interest to investigate a wider spectrum of biological characteristics of high-latitude antarctic fish and relate these to the key factors of the particular environment, such as the southern Weddell Sea. Institute for Polar Ecology, University of Kiel, Olshausenstr. 40-60 D-2300 Kiel I(W), FRO
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