COMPARATIVE PHYSIOLOGY OF TEMPERATURE REGULATION PART 1 : v:~ j: ^^'^•-"'v Editors JOHN P. HANNON m ELEANOR VIERECK o_D D D m a ARCTIC AEROMEDICAL LABORATORY FORT WAINWRIGHT ALASKA 1962 ACKNOWLEDGEMENTS Grateful acknowledgement is made to the following for permission to publish copyrighted figures: American Physiological Society, George Banta Publishing Co., St. Martin's Press, the Missouri Agricultural Ex- periment Station, and Reinhold Publisher's. PROCEEDINGS SYMPOSIA ON ARCTIC BIOLOGY AND MEDICINE II. COMPARATIVE PHYSIOLOGY OF TEMPERATURE REGULATION Symposium held July 17, 18, 19, 1961 at the Arctic Aeromedical Laboratory Fort Wainwright, Alaska Edited by John P. Hannon, Head, Department of Physiology Arctic Aeromedical Laboratory Eleanor G. Viereck, Research Physiologist Arctic Aeromedical Laboratory Symposium held under the auspices of Geophysical Institute University of Alaska College, Alaska 1962 TABLE OF CONTENTS Part I. The Evolution of Homeothermy. 1. Introductory Remarks, John P. Hannon ^^^ 2. Some Temperature Adaptations of Poikilothermic Vertebrates, C. Ladd Prosser 1 3. The Evolution of Avian Temperature Regulation William R. Dawson 45 4. The Evolution of Mammalian Temperature Regulation, Kjell Johansen 73 Part II. The Maintenance of Homeothermy. 5. Heterothermy in the Cold Adaptations of Warm Blooded Animals, Laurence Irving 133 6. Maximal Metabolism and Organ Thermogenesis in Mammals, Ladislav Jansky 175 7. The Physiology of Mammalian Cold Acclimation, J. Sanford Hart 203 8. Temperature Responses and Adaptations in Domestic Mammals, Max Kleiber 243 9. Temperature Responses and Adaptations in Birds, George C. West 291 10. Racial Variations in Human Responses to Temperature, Frederick A. Milan 325 11. Temperature Regulation in Animals Native to Tropical and High Altitude Environments, Peter R. Morrison 381 12. Temperature Regulation in Animals Native to the Desert Environment, Jack W. Hudson 413 ii BIOGRAPHICAL NOTES ON AUTHORS G. Ladd Prosser obtained his B.A. at the University of Rochester in 19 29, his Ph.D. at the Johns Hopkins University Zoology Department in 1932, and now holds the title of Professor of Physiology, University of Illinois. He has performed outstanding work in the fieldsofphysiology of nervous systems of invertebrates, comparative pharmacology of hearts, conduction in non-striated muscles, and physiological adaotation: (1) mechanisms and (2) role in evolution. The text. Comparative Animal Physiology, edited by Prosser, et al., needs no introduction. William R. Dawson received his Ph.D. from Stanford, Depart- ment of Zoology, in 1953, and is now assistant professor in the Department of Zoology at the University of Michigan. He has worked on temperature regulation and water balance of birds and reptiles, reptile metabolism, and avian paleontology. Kjell Johansen has the degree Gand. Real, from Oslo, Norway. He has worked with many kinds of vertebrates and higher inverte- brates. He is concerned with physiological phylogeny, and compara- tive cardiovascular physiology. He is now atthe University of Oslo, Institute for Experimental Medical Research. Laurence Irving received the degree of A.B. at Bowdoin in 1916, the A.M. from Harvard in 1917, and the Ph.D. from Stanford in 19 25. He has been awarded honorary degrees also: an honorary M.D. from Oslo in 1956 and an honorary D.Sc. from Bowdoin in 1959. His long and productive list of research accomplishments include work in diverse fields such as adaptations to cold in mam- mals and birds, migrations of Alaskan birds, and respiratory exchange in diving mammals. He has elucidated the principle of peripheral heterothermy. 82663 Ladislav Jansky received the P.H. degree in 1958 from Charles University, Prague, £tepartment of Animal Physiology. He now holds the title of Assistant Professor, Department of Comparative Phys- iology, Charles University. His research interests are: metabolism in the cold, cold adaptation, maximal metabolism, organ thermo- genesis, and hibernation. J. Sanford Hart received the B.A. 1939, the M.A. 1943, the Ph.D. 1949 all from Toronto, and was elected a F.R.S.C. in 1956. His present title is Head, Environmental Physiology section, Division of Applied Biology, NRC, Ottawa, Canada. His general interest encompasses theenvironmentalphysiology of homeotherms, acclimatization in mammals and birds native to cold climates metabolic mechanisms of acclimation to cold in rats, shivering mechanisms and organ thermogenesis, and cardiovascular res- ponses to exercise and cold exposure. Max Kleiber obtained the D.Sci. in 1924 and the Privat Dozent in 1928 from the Federal Institute of Technology, Zurich, Switzer- land. He is now Professor Emeritus at the University of California and has been awarded an honorary LL.D. from California in 1961. His numerous contributions to physiology have been in the fields of energy metabolism and feed utilization, environmental tempera- ture, energy metabolism and food utilization (in chicks and cows), measurement of replacement equivalents for food in animal produc- tion, energy metabolism and nutritive deficiencies, intermediary metabolism especially in milk formation using radiocarbon (C ) as a metabolic tracer, and phosphorus and calcium metabolism using P and Ca as tracers. His recent book. Fire of Life, is destined to become a classic. George C. West received his B.A. degree from Middlebury College in 1953, and his M.S. and Ph.D. in Zoology under Dr. S. C. Kendeigh at the University of Illinois in 1956 and 1958 respectively. He spent a year doing post-doctoral research at Illinois and a year as a post-doctoral fellow of the National Research Council of Canada with Dr. J. S. Hart. He is now Assistant Professor of IV Zoology at the University of Rhode Island and on the summer staff of the Canadian National Research Council. His major research interests and publications are in the fields of ecology and physio- logy of avian energetics and migration. Frederick A. Milan obtained his B.A. in 19 52 from the Univer- sity of Alaska and his M.A. from the University of Wisconsin, 1959. He is currently studying for the Ph.D. degree at the Univer- sity of Wisconsin, Department of Anthropology. His investigations of human response to cold have taken him to the Australian interior, the arctic coast of Alaska, the Tierra del Fuego, Lappland, the Antarctic, the Canadian Archipelago, and mountain summits in Alaska. Peter Reed Morrison has a B.S. from Swarthmore College in 1940 and a Ph.D. from Harvard University Biology Department in 1947. Presently he is Professor of Physiology and Zoology, University of Wisconsin. His work includes the fieMs of tempera- ture regulation and energy metabolism in wild mammals, altitudinal adaptations in wild mammals, coagulation and the polymerization of fibrinogen, hibernation, physiological descriptions of intact animals, and individual tissue specializations. Jack W. Hudson obtained his M.A, from Occidental College, Biology Department, 1950 and his Ph.D. from University of California, Los Angeles, Zoology Department, 1960. He is now a lecturer in that department. The list of his research interests and is.: Aestivation and hibernation in mammals, water metabolism kidney function in desert mammals, uric acid excretion in birds, relation of heart rate, body size and thermal neutrality in birds. LIST OF DISCUSSANTS Adams, Thomas Hannon, John P. Department of Physiology Physiology Department University of Washington Arctic Aeromedical Laboratory Fort Wainwright, Alaska Durrer, John P. Physiology Department Kessel, Brina Arctic Aeromedical Laboratory Department of Biology Fort Wainwright, Alaska University of Alaska College, Alaska Eagan, Charles J. Physiology Department Miller, Keith Arctic Aeromedical Laboratory Arctic Public Health Research Fort Wainwright, Alaska Laboratory P. O. Box 960 Anchorage, Alaska Evonuk, Eugene Physiology Department Arctic Aeromedical Laboratory Opsahl, James F. Fort Wainwright, Alaska Department of Zoology Winona State College Winona, Minnesota Folk, G. Edgar, Jr. Department of Physiology University of Iowa Pitelka, Frank Iowa City, Iowa Museum of Vertebrate Zoology Berkeley 4, California VI INTRODUCTORY REMARKS John P. Harmon On behalf of the Commander and Staff of the Arctic Aeromedical Laboratory, it is indeed a pleasure to welcome such distinguished guests to this second Symposium on Arctic Biology and Medicine. We hope that your trip was an enjoyable one, despite the long distances many of you were forced to travel. It is our desire that you remem- ber this visit to our Laboratory and to the State of Alaska as a pleasant one as well as a scientifically profitable experience, and if there is anything that I or the other Staff members might do to assure this end, please do not hesitate to call on us. Before we begin the formal portions of our program, I would like to say a few words regarding our reasons for selecting the Comparative Phys- iology of Vertebrate Temperature Regulation as a symposium topic. As you are all well aware, the ability to adapt to an adverse or unusual environment is one of the more fundamental characteristics of all living things. In fact, we might go so far as to say that the ability to adapt to such environments is an essential prerequisite to the successful perpetuationof any population of plants or animals. Thus, when a species is unable to adapt to an adverse environment it becomes extinct. To those ofus wholivein arctic or subarctic areas, the adapta- tions of plants and animals to adverse environmental temperatures