Physiology of Cold Adaptation in Birds

Physiology of Cold Adaptation in Birds

NATO ASI Series Advanced Science Institutes Series A series presenting the results of activities sponsored by the NATO Science Committee, which aims at the dissemination of advanced scientific and technological knowledge, with a view to strengthening links between scientific communities. The series is published by an international board of publishers in conjunction with the NATO Scientific Affairs Division A B c D E F G H Life Sciences Physics Mathematical and Physical Sciences Behavioral and Social Sciences Applied Sciences Computer and Systems Sciences Ecological Sciences Cell Biology Plenum Publishing Corporation New York and London Kluwer Academic Publishers Dordrecht, Boston, and London Springer-Verlag Berlin, Heidelberg, New York, London, Paris, and Tokyo Recent Volumes in this Series Volume 169-Evolutionary Tinkering in Gene Expression edited by Marianne Grunberg-Manago, Brian F.C. Clark, and Hans G. Zachau Volume 170-ras Oncogenes edited by Demetrios Spandidos Volume 171-Dietaryw3 and w6 Fatty Acids: Biological Effects and Nutritional Essentiality edited by Claudio Galli and Artemis P. Simopoulos Volume 172-Recent Trends in Regeneration Research edited by V. Kiortsis, S. Koussoulakos, and H. Wallace Volume 173-Physiology of Cold Adaptation in Birds edited by Claus Bech and Randi Eidsmo Reinertsen Volume 174-Cell and Molecular Biology of Artemia Development edited by Alden H. Warner, Thomas H. MacRae, and Joseph C. Bagshaw Volume 175-Vascular Endothelium: Receptors and Transduction Mechanisms edited by John D. Catravas, C. Norman Gillis, and Una S. Ryan Series A: Life Sciences

Physiology of Cold Adaptation in Birds Edited by Claus Bech and Randi Eidsmo Reinertsen University of Trondheim Trondheim, Norway Springer Science+Business Media, LLC

Proceedings of a NATO Advanced Research Workshop on Physiology of Cold Adaptation in Birds, held June 6-10, 1988, in Loen, Norway Library of Congress Cataloging in Publication Data NATO Advanced Research Workshop on Physiology of Cold Adaptation in Birds (1988: Loen, Norway) Physiology of cold adaptation in birds 1 edited by Claus Bech and Randi Eidsmo Reinertsen. p. cm.-(nato ASI series. Series A, Life sciences; voi. 173) "Proceedings of a NATO Advanced Research Workshop on Physiology of Cold Adaptation in Birds, held June 6-10, 1988, in Loen, Norway"-T.p. verso. Bibliography: p. lncludes index. ISBN 978-1-4757-0033-6 ISBN 978-1-4757-0031-2 (ebook) DOI 10.1007/978-1-4757-0031-2 1. Birds-Physiology-Congresses. 2. Cold adaptation-congresses. 1. Bech, Claus, 1951-11. Reinertsen, Randi Eidsmo. III. North Atlantic Treaty Organization. Scientific Affairs Division. IV. Title. V. Series: NATO ASI series. Series A, Life sciences; v. 173. QL698.N38 1988 89-16102 598.2'542-dc20 CIP 1989 Springer Science+Business Media New York Originally published by Plenum Press, New York in 1989 Softcocer reprint of the hardcover 1 st edition 1989 Ali rights reserved No part of this book may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher

PREFACE The papers in this volume were presented at the NATO Advanced Workshop: Physiology of Cold Adaptation in Birds, held 10.-15. June, 1988, at Hotel Alexandra, Loen, Norway. The workshop was generously supported by the NATO Scientific Affairs Division, Brussel, Belgia (grant ARW 585/87). We are also grateful to the College of Science, University of Trondheim, Norway, for supporting the participation of students. Most recent symposia and meetings dealing with thermoregulation in vertebrates have emphasized on mammalian studies, and this book seems to be the first one ever published which solely deals with thermoregulation in birds. Birds and mammals maintain their internal body temperature constant and independent of fluctuating ambient temperatures. The present knowledge of mam- malian and avian thermoregulation is extensive, but major questions are still unanswered. Furthermore, birds differ from mammals in many fundamental physiological aspects. Thus, even though both mammals and birds regulate their body temperature within strictly defined limits, the mechanisms utilized by birds are often different from those used by mammals. Only by knowing the mechanisms underlying thermoregulation in both groups that regulate their body temperature, can we fully understand the basic mechanisms of thermoregulation in homeotherms. The main themes for the workshop were central nervous control mechanisms regulating body temperature, the nature of effector mechanisms, including shivering and non-shivering thermogenesis, and also factors initiating changes in the effector mechanisms. We greatly appreciate, through the NATO Scientific Affair's Division, to have been able to organize such a workshop, and it is our hope that the publication of this book will stimulate further research in the field of avian thermoregulation. We greatly acknowledge and appreciate the enthusiastic responses of Professor William R. Dawson and Professor Eckhart Simon in participating in the organizing committee of the workshop. Thanks are also given to all individual participants who made it an outstanding event. We are especially indepted to Professor John 0. Krog, who, through his speech, reminded us about the great impact of Norwegian scientists on the field of comparative physiology. v

Lastly, we would like to thank the publisher, Plenum Press. Thanks are especially given to Janie Curtis and Gregory Safford, who provided valuable advice and assistance in publication of this book. Claus Bech Randi Eidsmo Reinertsen

CONTENTS CENTRAL MECHANISMS OF THERMOREGULATION Nervous control of cold defence in birds........ 1 E. Simon The shivering response in Common Eider ducks....... 17 J.B. Mercer On the thermosensitivity of the spinal cord in pigeons..... R.E. Reinertsen and c. Bech 27 MECHANISMS OF HEAT PRODUCTION Shivering and nonshivering thermogenesis in birds: A mammalian view E. Connolly, J. Nedergaard and B. Cannon 37 Muscular nonshivering thermogenesis in cold-acclimated ducklings............. 49 H. Barre, C. Duchamp, J.-L. Rouanet, A. Dittmar and G. Delhomme Nonshivering thermogenesis in winter-acclimatized King Penguin chicks........... C. Duchamp, H. Barre, D. Delage, G. Berne, P. Brebion and J.-L. Rouanet Apparent non-muscular thermogenesis in cold-exposed phasianid birds E. Hohtola, R. Imppola and R. Hissa 59 69 Shivering in aerobic and anaerobic muscles in bantams (Gallus domesticus)........ 77 A. Aulie and 0. Teien METABOLIC ADAPTATIONS Metabolic acclimatization to cold and season in birds... W.R. Dawson and R.L. Marsh 83

Enzyme activities in muscles of seasonally acclimatized house finches c. Carey, R.L. Marsh, A. Bekoff, R.M. Johnston and A.M. Olin 95 Energy substrates and metabolic acclimatization in small birds........... 105 R.L. Marsh and W.R. Dawson Thermogenic capacity of greenfinches and siskins in winter and summer... 115 s. Saarela, B. Klapper and G. Heldmaier Heat increment of feeding in the Kestrel, Falco tinnunculus, and its natural seasonal variation....... 123 D. Masman, S. Daan and M. Dietz Thermoregulation and energetics of arctic seabirds............... 137 G.W. Gabrielsen and F. Mehlum Thermoregulatory adaptations to cold in winter-acclimatized Longtailed ducks (Clangula hyemalis)... 147 B.M. Jenssen and M. Ekker RESPIRATION AND CIRCULATION Respiration and gas exchange in birds J. Piiper and P. Scheid...... 15 3 Gas exchange during cold exposure in Pekin ducks (Anas platyrhynchos)... 163 C. Bech and H. Johannesen Respiratory responses of the mallard to external and internal cooling... 173 H. Johannesen and C. Bech The respiratory pattern and expiratory gas concentrations in torpid hummingbirds Colibri coruscans... 179 M. Berger and K. Johansen Energy metabolism and patterns of ventilation in euthermic and torpid hummingbirds...... 187 T.L. Bucher and M.A. Chappell Respiration by birds at high altitude and in flight................... 197 M.H. Bernstein viii

Body and brain temperatures in pigeons at simulated high altitudes... 207 M.H. Bernstein Circulatory adaptations to cold in birds... 211 u. Midtgard Control of cardiorespiration during shivering thermogenesis in the pigeon............. 223 W. Rautenberg PHYSIOLOGY OF HYPOMETABOLISM Sleep, hypometabolism, and torpor in birds........ 231 H.C. Heller Adaptive capacity of the pigeon's daily body temperature rhythm............. 247 R. Graf, H.C. Heller, S. Krishna, W. Rautenberg and B. Misse Thermal and feeding reactions of pigeons during food scarcity and cold......... 255 M.E. Rashotte, D. Henderson and D.L. Phillips Metabolism and body temperature during circadian sleep and torpor in the fed and fasting pigeon............ 265 N.H. Phillips and R.J. Berger Autonomic and behavioral temperature regulation as a part of the response complex to food scarcity in the pigeon............. 275 J. Ostheim and w. Rautenberg Body mass, food habits, and the use of torpor in birds................... 283 B.K. McNab BREEDING AND INCUBATION Energy saving during breeding and molt in birds... 293 J.-P. Robin, Y. Handrich, Y. Cherel andy. LeMaho Effect of clutch size on efficiency of heat transfer to cold eggs in incubating Bantam hens............... 305 0. T0ien ix

Emu winter incubation: Thermal, water, and energy relations.......... 315 W.A. Buttemer and T.J. Dawson Energy saving in incubating Eiders............ 325 G.W. Gabrielsen ADAPTATIONS TO COLD IN CHICKS Adaptations to cold in bird chicks... R.E. Ricklefs..... 329 Energy partitioning in Arctic tern chicks (Sterna paradisaea) and possible metabolic adaptations in high latitude chicks......... 339 M. Klaassen, C. Bech, D. Masman and G. Slagsvold Energetics of avian growth: the causal link with BMR and metabolic scope....... 349 R.H. Drent and M. Klaassen Strategies for homeothermy in Eider ducklings (Somateria mollissima)... 361 J.B. Steen, H. Grav, B. Borch-Iohnsen and G.W. Gabrielsen Body temperatures under natural conditions of two species of alcid chicks... 371 D. Vongraven, F.J. Aarvik and C. Bech Participants 3 7 5 Index...... 379 X