ENVIRONMENTAL SCIENCES An Interdisciplinary Monograph Series EDITORS DOUGLAS H. K. LEE E. WENDELL HEWSON DANIEL OKUN National Institute of Department of Department of Environmental Environmental Health Sciences Atmospheric Science Sciences and Engineering Research Triangle Park Oregon State University University of North Carolina North Carolina Corvaliis, Oregon Chapel Hill, North Carolina ARTHUR C. STERN, editor, AIR POLLUTION, Second Edition. In three volumes, 1968 L. FISHBEIN, W. G. FLAMM, and H. L. FALK, CHEMICAL MUTAGENS: Environ­ mental Effects on Biological Systems, 1970 DOUGLAS H. K. LEE and DAVID MINARD, editors, PHYSIOLOGY, ENVIRON­ MENT, AND MAN, 1970 KARL D. KRYTER, THE EFFECTS OF NOISE ON MAN, 1970 R. E. MUNN, BIOMETEOROLOGICAL METHODS, 1970 M. M. KEY, L. E. KERR, and M. BUNDY, PULMONARY REACTIONS TO COAL DUST: "A Review of U. S. Experience," 1971 DOUGLAS H. K. LEE, editor, METALLIC CONTAMINANTS AND HUMAN HEALTH, 1972 DOUGLAS H. K. LEE, editor, ENVIRONMENTAL FACTORS IN RESPIRA­ TORY DISEASE, 1972 H. ELDON SUTTON and MAUREEN I. HARRIS, editors, MUTAGENIC EFFECTS OF ENVIRONMENTAL CONTAMINANTS, 1972 RAY T. OGLESBY, CLARENCE A. CARLSON, and JAMES A. MCCANN, editors, RIVER ECOLOGY AND MAN, 1972 LESTER V. CRALLEY, LEWIS T. CRALLEY, GEORGE D. CLAYTON, and JOHN A. JURGIEL, editors, INDUSTRIAL ENVIRONMENTAL HEALTH: The Worker and the Community, 1972 MOHAMED Κ. YOUSEF, STEVEN M. HORVATH, and ROBERT W. BULLARD, PHYSIO­ LOGICAL ADAPTATIONS: Desert and Mountain, 1972 In preparation DOUGLAS H. K. LEE and PAUL KOTIN, editors, MULTIPLE FACTORS IN THE CAUSATION OF ENVIRONMENTALLY INDUCED DISEASE MERRIL EISENBUD, ENVIRONMENTAL RADIOACTIVITY, Second Edition PHYSIOLOGICAL ADAPTATIONS DESERT AND MOUNTAIN Edited by MOHAMED Κ. YOUSEF STEVEN M. HORVATH DEPARTMENT OF BIOLOGICAL SCIENCES INSTITUTE OF ENVIRONMENTAL STRESS UNIVERSITY OF NEVADA, LAS VEGAS UNIVERSITY OF CALIFORNIA, SANTA BARBARA ROBERT W. BULLARD DEPARTMENT OF ANATOMY AND PHYSIOLOGY INDIANA UNIVERSITY, BLOOMINGTON 1972 Academic Press New York and London COPYRIGHT © 1972, BY ACADEMIC PRESS, INC. ALL RIGHTS RESERVED NO PART OF THIS BOOK MAY BE REPRODUCED IN ANY FORM, BY PHOTOSTAT, MICROFILM, RETRIEVAL SYSTEM, OR ANY OTHER MEANS, WITHOUT WRITTEN PERMISSION FROM THE PUBLISHERS. ACADEMIC PRESS, INC. Ill Fifth Avenue, New York, New York 10003 United Kingdom Edition published by ACADEMIC PRESS, INC. (LONDON) LTD. 24/28 Oval Road, London NW1 LIBRARY OF CONGRESS CATALOG CARD NUMBER: 70-187240 PRINTED IN THE UNITED STATES OF AMERICA List of Contributors Numbers in parentheses indicate the pages on which the authors' contributions may be found. E. F. Adolph, Department of Physiology, University of Rochester, Rochester, New York(l) B. Balke, Department of Physical Education, University of Wisconsin, Madison, Wisconsin (195) H. S. Bel ding, Department of Occupational Health, Graduate School of Public Health, University of Pittsburgh, Pittsburgh, Pennsylvania (9) W. G. Bradley, Department of Biological Sciences, University of Nevada, Las Vegas, Nevada (127) Robert W. Bullard,* Department of Anatomy and Physiology, Indiana University, Bloomington, Indiana L. D. Carlson, Department of Human Physiology, School of Medicine, University of California, Davis, California (65) C. F. Consolazio, U.S. Army Medical Research and Nutrition Laboratory, Fitz- simons General Hospital, Denver, Colorado (227) F. N. Craig, Medical Research Laboratory, Edgewood Arsenal, Maryland (53) W. H. Forbes, Harvard School of Public Health, Boston, Massachusetts (191) A. Pharo Gagge, John B. Pierce Foundation Laboratory, New Haven, Connecti­ cut (23) S. M. Horvath, Institute of Environmental Stress, University of California, Santa Barbara, California (183) H. L. Johnson, U. S. Army Research and Nutrition Laboratory, Fitzsimons Gen­ eral Hospital, Denver, Colorado (227) R. E. Johnson, University of Illinois, Urbana, Illinois (99) H.J. Krzywick, U. S. Medical Research and Nutrition Laboratory, Fitzsimons General Hospital, Denver, Colorado (227) D. H. K. Lee, National Institutes of Environmental Health Science, Department of Health, Education, and Welfare, Research Triangle Park, North Carolina (109) •Deceased. ix χ List of Contributors U. C. Luft, Department ofPhysiology, Lovelace Foundation, Albuquerque, New Mexico (143) R. A. McFarland, Harvard School of Public Health, Boston, Massachusetts (157) S. Robinson, Department of Anatomy and Physiology, Indiana University, Bloomington, Indiana (77) Mohamed Κ. Yousef, Department of Biological Sciences, University of Nevada, Las Vegas, Nevada (127) Preface and Dedication This book is published as a tribute to Dr. David Bruce Dill, Research Professor and Director of the Laboratory of Environmental Patho-Physiology, Desert Re­ search Institute, University of Nevada System, Boulder City, Nevada in cele­ bration of his eightieth birthday. Many of his students or associates during the last five decades of his life — at the Fatigue Laboratory, Harvard University, Army Chemical Center, Indiana University, and University of Nevada — gathered for a symposium held at the University of Nevada (UNLV) on April 19-20, 1971 to pay him tribute. All were very pleased not only to see him enjoying life in good health but to note his highly active research endeavors carried out six days a week in his new laboratory at the Desert Research Institute, University of Nevada System. It is our fortune to be reminded that Dr. Dill, an outstanding scholar, has a brilliant record of achievements in the fields of environmental physiology, exer­ cise physiology, and comparative physiology. His contributions have appeared in many scientific journals where one can find over 250 manuscripts. In collabora­ tion with Α. V. Bock, Dill wrote the third edition of the "Physiology of Mus­ cular Exercise," published in 1931. This book includes 46 diagrams and 418 references and is considered one of the fundamental foundations of the field of exercise physiology. It was followed by a famous contribution on the physi­ ological responses of living organisms to stress entitled "Life, Heat, and Alti­ tude" published by Harvard University Press in 1938. Recently, with the help of E. F. Adolph and C. G. Wilber, Dill edited the "Handbook of Physiology," Sec­ tion 4, entitled "Adaptation to the Environment." This book represents the first authoritative reference on the subject and includes 65 chapters written by 66 eminent scientists from many different countries. The impact of Bruce Dill on the state of physiology is recognized by many colleagues. He has been in the forefront not only as an investigator and teacher but also as an organizer and leader of several expeditions to places such as the Andes, the Canal Zone, and the Colorado deserts. In recognition of his efforts, UNLV and the Desert Research Institute (DRI), University of Nevada System, organized this symposium as a special honor for him. In order to do this, some of his former associates were invited to review the current concepts of physi­ ological adaptations to desert and mountain areas of research to which Dill devoted a great deal of effort and research experience. We wish to acknowledge the help and support of Dr. Donald H. Baepler, Vice xi xii Preface and Dedication President for Academic Affairs, UNLV, and of Drs, John M. Ward, President of DRI, Robert B. Smith, Dean, College of Sciences and Mathematics, UNLV, and W. Glen Bradley, Professor of Biology, UNLV, in the organization of the sym­ posium. Dr. Baepler's encouragement and support throughout the preparation of this book are appreciated. The financial support given by the University of Nevada's Board of Regents made this symposium possible. Mohamed K. Yousef Steven M. Horvath Robert W- Bullard (1929-1971) Robert Winslow Bullard, distinguished physiologist and associate editor of this volume, died on Mt. McKinley June 24, 1971. He was making physiological observations on the members of the National Outdoor Leadership School of Lander, Wyoming. A group of four, including Bob, had climbed to 9300 feet and, while roped and wearing snowshoes, tramped out an air-drop area. Bob then took off his snowshoes, unfastened the rope, and stepped back on snow that covered a crevasse. A snowbridge collapsed and he fell 130 feet. Postmortem investigation indicated that death was virtually instantaneous. Born in Waltham, Massachusetts, June 13, 1929, Bob exemplified many tradi­ tional characteristics of the New England Yankees. His keen sense of humor, integrity of character, and brilliance in teaching and research were evident to those of us who came to know him in his years at Indiana. After his graduation from Springfield College in Massachusetts in 1951, he received his M.A. degree at the University of Massachusetts in 1954, and his Ph.D. with Edward Adolph at Rochester in 1956. The rest of his career was spent at Indiana University, first at Bloomington and then four years at the Medical School in Indianapolis. In 1962 he returned to Bloomington where from 1964 to 1969 he headed the Depart­ ment of Anatomy and Physiology. His sabbatical year of 1969—1970 was spent with Hardy in New Haven. During that year he was a Special Research Fellow of the National Institutes of Health. In 1970 he returned to his professorship at Bloomington. He became Dill's colleague in 1962 when he agreed to write a chapter on Annelids and Molluscs for the volume on "Adaptation to the Environment" that Adolph and Wilber helped Dill edit. That chapter demonstrated his intellectual capacity. In the same year Dill and he were at the Barcroft Laboratory, White Mt. Research Station. While working on different projects they shared the enjoy­ ment of dinner table discussions. When the National Science Foundation underwrote the establishment of the Laboratory of Environmental Patho-Physiology in 1966 Bob agreed to be a consulting physiologist. In 1968, he and Marlene and their three daughters, Kristen, Carolyn, and Alicia, spent a month in Boulder City, staying at the Hitchin Post Motel. Marlene, a former airline hostess, and the girls proved as adaptable as Bob to the desert environment. Bob's paper on "Responses of the burro to desert heat stress" with Dill and Yousef as joint authors was published in the Journal of Applied Physiology in August, 1970. Yousef presented another xiii xiv Robert W. Bullard paper on the two burros (Mabel and Maud) at the Federation meeting in April, 1971. Bob, as chairman, introduced him and suggested an alternate title "Further adventures of Mable and Maud." Bob and Marlene hoped to return with their daughters to Boulder City for the month of August, 1971. Plans for his summer studies were discussed when he came in April to take part in the symposium. His notable contribution to the symposium and his part in the discussion are matters of record. He and Horvath agreed to help Yousef edit the proceedings. For such a man as Bullard to be cut down at age 42 is a disaster for his family, for his friends, for Indiana University, and for physiology. But we can be thank­ ful that during his few years he left deep and lasting imprints on the family circle, on his many students, and on his colleagues in the fields of environmental physiology, temperature regulation, respiration, and sports medicine. D. B. Dill Sid Robinson Some General Concepts of Physiological Adaptations E. F. ADOLPH Introduction The study of physiological adaptations has recurred throughout my research career. In 1920 I spent a summer at a high altitude laboratory in the Alps and saw that mountain climbing became easy after I had lived on Monte Rosa for a couple of weeks. Some years later (1927) I was interested in the possible transition of aquatic animals from freshwater to seawater, and measured the adjustments of frogs to immersion in saline waters (1). In the next decade (1937), I came to Nevada with Bruce Dill and observed that we did the same walk day after day with less and less rise of body temperature and pulse rate (2). What did these three studies have in common? In each case, exposure to a particular environment modified the body in specific ways. But the specific ways had a uniform pattern in that the modifications increased for a few days and persisted for some days after exposure ceased. Evidently the results of adaptation have long time scales and often enable the body to respond to subsequent exposure in advantageous ways. At the symposium held when Dill "retired" from Edgewood, Maryland (1961) our theme was physiological adaptation. Upon that and other occasions, I, for one, tried to formulate concepts of physiological adaptations. Now, in 1971, new light can be thrown upon adaptations. As I see the general problem of physiological adaptations, we need three concepts. First, a precise notion of what these adaptations are; second, a