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PER FREDRIK THORKELSSON SCHOLANDER November 29, ~ 905-fune 13, ~ 980 BY KNUT SCHMIDT-NIELSEN THE MOST IMPRESSIVE ASPECT of Scholanders scien- tific life is his versatility as a biologist and his ability to make significant contributions in a broad range of fields. He was first of all a physiologist, but his work always signified a fresh approach to broader biological problems ant! prin- ciples. He had an ability to ask the right questions, to conceive of simple experiments and (resign the necessary equipment, to utilize novel approaches, and to present simple and logical answers to important questions. As a person he was warm and enthusiastic, generous ant! kind, and utterly uncon- cernect with the hassles of form and bureaucracy. Scholancler was born in Sweden and grew up in a family of talent and culture. His father was an engineer; his mother, Norwegian born, was an accomplished professional pianist; his grandfather was a prominent architect, as well as a writer anc! musician, and a professor at the Royal Academy of Arts. Scholander has related that, as a small boy, he crawlecl un- erneath his mother's Steinway granct piano when she prac- ticed, smothered! by the waves of emotion in the music. He himself became an accomplished violinist, which I first ctis- coverec! when late one night ~ found him playing in the lab- oratory to a record that omitted the first violin. 387
388 BIOGRAPHICAL MEMOIRS Because of the divorce of his parents, Scholancler moved to Norway, where in 1924 he matriculated at the Faculty of Medicine of the University of OsIo (then Christiania). At that time mectical students started with Latin, philosophy, chem- istry, and physics, whereupon followocl years of preclinical work in anatomy, physiology, and so on. Although Scholan- der was terribly borer! by much of this, he continued his med- ical studies and completed his medical education in 1932, within the normal time span. Scholancler has described how his textbooks were always full of marginal notes about expla- nations he ctict not believe, but his grades clic! not reflect his brilliance. There was a rumor among the university stu- dents probably untrue that he completed his final medi- cal examinations with the unique distinction of having ob- tained the lowest gracles ever given a passing student. Scholancler's years as a medical student became significant in a very cti~erent ant! unexpected] way. Walking home from the University one day, bored with tedious coursework, he picked some lichens off the trees along the street. At home he found a flora for lower plants ant] soon tract the lichens identified. He continucct collecting, and when his flora proved inadequate, he sought further help from the profes- sor of botany, Bernt Lynge. Lynge recognizes! the unusual talents of the young student, who rapidly became an out- standing lichen specialist. Lynge himself, because of severe arthritis, was unable to go to GreenIanc! and offered Scho- lander the opportunity to take his place as a botanist on sev- eral arctic expeditions. As a result, Scholancler spent three summers (1930-1932) in Greenland and Spitzbergen. His contributions to lichenology, particularly his revision of the family Umbilicariaceae ~ ~ 934b), placer! him among the worIct's foremost lichenologists at that time. The young physician who was never to practice medi- cine-tract become a botanist. He was approached by the pro
PER FREDRIK THORKELSSON SCHOLANDER 389 fessor of systematic botany, lens Holmboe, who suggester! that he should present himself for a doctorate in botany, and in 1934 he was awarded the Dr. Philos. by the University of Oslo. It might be expected that his dissertation wouIcT deal with lichens, but this would be uncharacteristic for Scholan- der he was awarcled the doctorate basect on a monograph of the vascular plants of Spitzbergen (1934a). During the arctic expeditions, Scholancler had been in trigucct by the many seals, polar bears, and diving birds he saw. He clearly saw that many important questions needect answers, such as "How do diving seals get enough oxygen?" and "Why clon't they get divers' disease as humans do after descending to similar depths?" He obtained working space in the basement of the University Physiological Institute, and with the aid of the professor of anatomy, the geneticist Otto Mohr, he was awarded a small university research fellowship that enabler! him to pursue his interests in physiology. He clevelopect new methods for the continuous recording of the respiratory metabolism of diving animals, and one of his pub- lications (1937b) caught the attention of the distinguished Danish physiologist August Krogh, who immediately under- stoocT Scholancler's genius for design of experimental appa- ratus. At that time ~ was a stuclent in (Copenhagen, and when Scholancler came to-give a lecture on his studies of diving animals, ~ sat there completely spelIbouncl by his brilliant presentation and the simple and logical answers he provided to questions that long tract puzzled physiologists who contem- platecl the mysteries of diving physiology. The most characteristic differences between a diving seal ant! a human are strikingly simple. First, when a seal begins a ctive, it exhales and clives with a minimal volume of air in the lungs. It may seem counterproductive to dive with little air in the lungs, but it greatly recluces the amount of nitrogen taken up by the blooct and the tissues during the dive. This
390 BIOGRAPHICAL MEMOIRS seems to be the key factor in how seals and whales avoid divers' disease, or the bencts, which is so tragically well known from humans who ascend after dives to great depths. The oxygen-carrying capacity of the bloocI, however, is much greater in the seal than in humans. Its blood volume is rela- tively larger, anct both blooct and muscles contain much larger amounts of hemoglobin, ant! thus hold more oxygen, than in mammals in general. A seal's most characteristic re- sponse to an experimental dive is to slow the heart down to a few beats per minute; the blood is diverted to the most vital organs, notably the central nervous system and the eyes. The muscles, which are able to function anaerobically through the formation of lactic acid, receive no blooct ant! thus acquire an oxygen debt that is repaid when oxygen is again available at the termination of the dive. These important results were . . . . . . . . . summarized in a monograph (1940a) that remains the foun dation for what we unclerstand today of the physiology of diving animals. At about the same time, Laurence Irving at Swarthmore College had also clone distinguished work on diving physi- ology, anct August Krogh arranger! for a Rockefeller fellow- ship for Scholancler to continue his studies of diving physi- ology at Swarthmore. Scholancler, however, was still in Norway when the Second WorIct War broke out, and the Rockefeller Foundation cancellecT all fellowship awards. When Krogh was informer! about this, he sent an urgent telegram to Scholancler that he should immediately leave for the USA, and Scholancler obtained space on the last ship that left Norway for the USA. Once lanclecI, he of course received the cancellecI fellowship, and the studies of diving animals were continued in collaboration with Irving. However, as the Uniter] States became involved in the Sec- oncl World War, Laurence Irving went to the Air Force, and Scholancler followed him there. Both Irving and Scholancler
PER FREDRIK THORKELSSON SCHOLANDER 391 made substantial scientific and practical contributions during their military service. Scholander became involved in the test- ing of military equipment and survival gear. He developecl simple and reliable field methods; for example, he stucTiecl the conditions under which field stoves causer! carbon mon- oxicle poisoning in tents and snow houses, he tried sleeping bags under blizzard conditions, and tested coverer! life rafts cluring storms in the Aleutian Islands. be_ · ~ · · ~ 1 . During this time he also arranged a quick and unauthor- ~zec! rescue mission to an airplane that tract crashed at the tip of the Alaskan peninsula. One survivor had been seen, but rescue teams had failed to reach the site over land and Scho- lancler's camp commander hac! vetoed parachute jumps. Going in by plane would be insubordination, but Scholancler found a young pilot who, risking his career, volunteered. An- other medical cloctor and a priestjoinecT them, ant! the three, who had never iumpec! before, came clown successfully, al .. . .. 1 , tnougn warn some difficulty. They found that two men had perished, but three survivors were taken care of by the M.D. while Scholancler took on the cooking, greatly helped by the plane's cargo of sacramental wine for Passover, or "Hang- over" as Scholander called it. Eventually the survivors and the rescuers were taken out by a bush plane from Anchorage. ~ have later heard that Scholander, as a result of the suc- cessfu! Aleutian rescue, was to be courtmartialed for insub- orclination. As the case slowly moved through the military channels, it became known to Detlev Bronk, who was an- gered by the bureaucratic stupidity of punishing one of the most elective scientists in the Air Force. By some unknown intervention, the case was aborted at the highest level in Washington. After the years of military work, Irving and Scholancler, with support from the Navy, established a research laboratory at Point Barrow, Alaska. Among the most important results
392 BIOGRAPHICAL MEMOIRS were measurements of the metabolic heat production ant! the insulation neecled for warmblooded animals, mammals and birds, to keep warm in the arctic climate. It turner! out that larger animals, such as foxes and eskimo clogs, could sleep at temperatures as low as - 30 or - 40°C without any increase in heat production. Smaller animals, however, started to shiver at temperatures well above freezing, and they neecled increases! heat production to stay warm and maintain a nor- mal body core temperature of 37°C. The information was in agreement with the simple physical laws of heat exchange, anti the studies clearly showed that below a certain temper- ature, the lower critical temperature, the metabolic rate in- creased linearly with the decrease in temperature. This mode} for the responses of warmbloocled animals to low tem- peratures has remained! the mocle} for virtually all later stud- ies of this nature (1950c,d,e). The arctic work incluclec] studies of coldblooded animals and of plants, ant! also the tolerance of various organisms to freezing. Scholancler devisect a simple and elegant method for determining the amount of body water in a mosquito larva that was actually frozen to ice inside the intact animal. He showed that these, when they were thawed out, were still alive and unharmed after more than 80 percent of their body water hac! been frozen to ice (1953e). Scholander's many ingenious methods for the analysis of minute samples of gas became very helpful in an unexpected area. He noticed that gas bubbles tendect to clevelop under ice, and also that a chunk of ice chopper! from a glacier gave off bubbles of gas when put into a drink to coo} it. Scholander determinect that ice is virtually impermeable to gases; he froze thin sheets of ice, 0. ! mm thick, and floated them on cold mercury. He introduced small volumes of gases under the ice and attempter] to measure changes in their composi- tion. He was never able to detect any penetration, and con
PER FREDRIK THORKELSSON SCHOLANDER 393 cludect that ice at -10°C is at least 70,000 to 80,000 times more impermeable to gases than a layer of water. This made him wonder whether the air trapped creep in the Greenland glaciers contains a permanent record of the composition of the atmosphere in earlier periods (1956b,e; 195Sc,g; 1960c; 196Ic; 1962g). After the two years of work at Barrow, Scholander left the Arctic Research Laboratory. A. Bairc! Hastings, professor of biological chemistry at Harvard University, was a person who fully unclerstood Scholancler's genius, and he offered him a position as research fellow. During two years in Hastings' lab- oratory, Scholancler clevelope(1 a series of elegant micrometh- o(ls for blood and gas analysis, and started applying these to solving some of the intriguing problems of how gases are secreted into the swimblacicler of fish. Depending on the depth at which a fish is found, the gases in the swimblacicler are uncler high pressure. Gases that are dissolvecI in the sur- rounding seawater are at partial pressures close to those in the atmosphere, and they are secreted into the swimblacIder against pressures that in deep sea fish may amount to several hundred atmospheres. The micromethocts Scholander hacI clevelopect were ideal for these studies. He now demonstratect that not only oxygen, but also nitrogen, is secreted against tremendous concentration gradients. That this is possible for oxygen, a relatively reactive gas, was perhaps not surprising, but how couIct nitrogen, a presumably totally inert gas, also be secreted actively against such concentration gradients? The answer to both puzzles lay in the unusual vascular supply to the swimblacIder and in physical principles that, once they were understood, were simple to explain. The blood supply to the swimbladder forms a counter-current system that is an essential part of the secretion mechanism (1954a,b). The swimbladcler studies were continued cluring the three years of Scholancler's association with the Woods Hole
394 BIOGRAPHICAL MEMOIRS Oceanographic Institution as a physiologist ( 1952-19551. The result was a series of papers that now form the foun- dation for our unclerstancling of swimblaclcler function 955g, ~ 958f, and others). In 1955 Scholander was callecI to the University of OsIo as professor and director of a new Institute of Zoophysi- ology. His three years in OsIo were very active. He continued studies of the adaptation of humans to cold and answered questions of how primitive humans can keep warm in cold climates. Eskimoes, Lapps, and Australian aborigines were comparer! with Norwegian students. The latter slept out- doors at temperatures around 0°C, naked in a single-blanket sleeping bag. For an unacclimatized! person this is intensely uncomfortable and it is impossible to sleep, but after five or six days of acclimatization the students had no trouble. The most notable change in their physiology was a substantial in- crease in metabolic heat production. (The motivation of the students to go through with these uncomfortable experi- ments seemed to be related to their being granted permission to hunt reindeer.) When the Australian aborigines were studiecI, it was found that they could sleep through a coin night with a nor- mal metabolic rate. The difference was that the aborigines permitted the temperatures of the legs to ctrop to around 10°C. In other words, they had adapted by permitting their peripheral temperatures to drop instead of increasing their metabolic rate. This, of course, is a more economical ap- proach; keeping the legs warm requires a substantial increase in metabolic heat production. After three highly productive years in Norway, Scholan- cler was brought to the Scripps Institution of Oceanography through the efforts of its director, the oceanographer Roger Revelle, anc! he remained there until the enct of his life. With tremendous enthusiasm he took up one problem after an- other. Among these were some important problems of plant
PER FRED RI K TH O RKELS SO N S C H O LAN DER 395 physiology. The question of how water can be brought to the top of the tallest trees had always intrigued him, and studies that he hac! initiated while he was at Woods Hole and in OsIo were continuccI ~ ~ 957a, ~ 958b, ~ 96 ~ d). The fact that suction, in combination with the cohesion of water, can explain the ascent of the sap lect to later studies of osmotic phenomena in other plants, and eventually to the clevelopment of the solvent tension theory of osmosis. Important in this connec- tion was a series of studies of mangroves and their salt bal- ance and the question of how they manage with their roots in seawater. These studies started on the Cape York Peninsula in northern Australia ~ ~ 962h) and continucct for several years ~ ~ 965a,c; ~ 966a; ~ 96Sb). The solvent tension theory of osmosis was founded on Scholander's experience in plant physiology. A series of ele- gant experiments, in part carried out in collaboration with H. T. Hammel, also of Scripps, provided easily compre- hendect support for Scholancler's theories ~197 Ic,e). The theories were severely criticized by several physical chemists, who based their arguments on classical thermodynamic ar- guments. Scholander's ingenious experiments remained] un- challenged; only their interpretation could be disputed. Although Scholancler's view is simple and intuitive, it is un- doubtedly more convenient to adhere to the traditional way of thinking about osmotic phenomena. I can only regret that I do not have the competence to evaluate the controversy and give proper perspective to the significance of Scholancler's contributions in this field. A most important development during .~cholander's . ~. . . . many years at the Scripps Institution was the design and builcling of the research vessel Alpha Helix. During his many field expeditions, Scholancler tract understood the impor- tance of bringing adequate equipment into the field, where the important physiological problems are evident. He con- ceived of a laboratory ship that should be able to take a group
396 BIOGRAPHICAL MEMOIRS of a dozen or so scientists to any area of the worIcI. The ship should be equipped with modern laboratories anti an excel- lent machine shop. This was based on Scholander's own ex- perience; he was a skilled machinist and this hacT helped him during his many successes in designing new methods ant! equipment. He saw the absolute necessity of always having at his disposal a good machine shop that could solve problems as they arise uncler field conditions. Support for building the ship was obtained from the Na- tional Science Foundation, and in 1966 the ship started on its first cruise, the BilIabong Expedition to tile Great Barrier Reef. A National Advisory Board, initially chaired by Baird Hastings, evaluated applications for use of the ship. Over the fourteen years the ship remained in service as a floating phys- iological laboratory, expeditions went to Australia, the South Seas, the Amazon, the Antarctic, the Galapagos, the Bering Sea, and other sites. Several hundred scientists from all over the work! have participated actively in these expeditions, and the records of Scripps Institution show that the work on the Alpha Helix has resulted! in a total of 547 publications in rec- ognized scientific journals an impressive record for the rel- atively modest funds invested in the Alpha Helix. Not only through the Alpha Helix, but throughout his life, Scholander became a seminal figure for physiologists who were concerned with the problems that animals and plants encounter in nature. He was an immensely enthusiastic per- son, a true naturalist who perceived interesting scientific problems wherever he moved. He lee! a restless ant! highly productive life. A large number of scientists, now active anti recognized arounc] the world, have been associated with Scholancler and have been influenced by his stimulating and dynamic personality. In 1951 he married the daughter of Laurence Irving, Susan, who remained his devoted compan- ion the remainder of his life.
PER FREDRI K THORKELS S ON SC H O LAN DER 397 Throughout his productive life, Scholancler retained the ability to open up new fields of investigation. He was not only a brilliant scientist, but also an enthusiastic and charming person, engaging, warm, and generous. His way of managing the increasing constraints imposed by the bureaucracy of uni- versity life and government support of science was merely to ignore them. As an example of his charming and untradi- tional personality, ~ wish to relate his study of wave-ricling dolphins. Many of us have seen dolphins or porpoises that playfully rifle along just in front of the bow wave of a ship, motionless and effortlessly remaining in place. This, of course, arouser! Scholancler's curiosity. It belongs to the story that the free rifle of the dolphins had been the subject of theoretical analysis by fluid clynamicists, but Scholander thought that the theory, to use his own words, "missort the boat." When on board a Norwegian sealing vessel, he rigged up a simple device with a streamlinect vane attached to a spring to measure the forces in the bow wave-anti he found that these might indeed explain how a dolphin gets the neecled forwarc! thrust to remain in the wave ~ ~ 959a,ct). Characteristically, Scholander concludecI his paper with these words: "This, ~ believe, is the way dolphins rifle the bow wave, and if it is not, they shouIcI try." Scholander cliect in his home in La Volta, California, on tune 13, 1980. He was seventy-four years oict. Ten days ear- lier he had broken his hip in a fall in his laboratory, but after a brief hospital stay he was home again and enthusiastically planning for the summer's activities, which included a trip to Europe. A sucIden collapse ended his long and productive life. T H E M A T E R ~ A E ~ H A V E U S E D to prepare this biography has been assembled with the help of Susan Irving Scholander and the pre- sent director of the Physiological Research Laboratory at Scripps
398 BlOGRAPHICAE MEMOIRS Institution of Oceanography, Dr. Fred N. White. Archival material has been made available to me by Elizabeth N. Shor, also of Scripps Institution of Oceanography. My own familiarity with P. F. Scho- lander's life comes from my admiration for his scientific contribu- tions and from many years' close friendship. Other good friends have helped me by reading my manuscript.
PER FREDRIK THORKELS SON SC HOLANDER 399 HONORS AND DISTINCTIONS EDUCATION M.D., University of Oslo, 1932 Dr. Philos. (Botany), University of Oslo, 1934 PROFESSIONAL APPOINTMENTS 1932-1934 Instructor of Anatomy, University of Oslo 1932-1939 Research Fellow in Physiology, University of Oslo 1939-1941 Rockefeller Fellow 1939-1943 Research Associate, Department of Zoology, Swarth- more College 1943-1946 U.S. Air Force. Commissioned as Captain, 1943; Ma- jor, 1946 1946-1949 Research Biologist, Swarthmore College 1949-1951 Research Fellow, Department of Biological Chemistry Harvard Medical School 1952-1955 Physiologist, Woods Hole Oceanographic Institution 1955-1958 Professor of Zoophysiology, University of Oslo, and Director of Institute of Zoophysiology 1958 - 1973 Professor of Physiology, Scripps Institution of Ocean- ography, University of California, San Diego 1963-1970 Director, Physiological Research Laboratory, Univer- sity of California, San Diego 1973 - 1980 Emeritus Professor, Scripps Institution of Oceanog- raphy, University of California, San Diego HONORS AND AWARDS Rockefeller Fellow, 1939-1941 Soldiers' Medal for Valor (for Aleutian rescue), 1945 Legion of Merit (for pilot ejection seat), 1946 Norwegian Academy of Sciences, 1955 American Academy of Arts & Sciences, 1959 National Academy of Sciences, 1961 American Philosophical Society, 1962 Cosmos Club, 1964 John Simon Guggenheim Fellow, 1969-1970 Doctor of Science, University of Alaska, 1973 Royal Swedish Academy of Sciences, 1974 Doctor of Science, Uppsala University, 1977 Fridtjof Nansen Prize, Oslo, 1979
400 BIOGRAPHICAL MEMOIRS BIBLIOGRAPHY This list is based on one compiled by Susan Irving Scholander in 1964 and is updated with information received from Scripps Institution of Oceanography. It includes items published in books and journals, but not reports to government agencies and grant-giving foundations. 1932 With B. Lynge. Lichens from N.E. Greenland. Skr. Svalb. Ishavet, no. 41. 120 pp. 1933 a. Notes on Peltigera erumpens (Tayl.) Vain. s. 1. Nyt Mag. Naturv., 73:21-54. b. With l. Devoid. Flowering plants and ferns of Southeast Green- land. Skr. Svalb. Ishavet, no. 56. 220 pp. 1934 a. Vascular plants from northern Svalbard. Skr. Svalb. Ishavet, no. 62. 155 pp. b. On the apothecia in the lichen family Umbilicariaceae. Nyt Mag. Naturv., 75:1-41. 1937 a. With Eilif Dahl and B. Lynge. Lichens from Southeast Green- land. Skr. Svalb. Ishavet, no. 70. 90 pp. b. New graphic methods for the recording of the respiratory gas- eous exchange. Skr. Nor. Vidensk. Akad. Oslo, no. 3. 73 pp. 1938 a. A modified manometric blood gas apparatus. Scand. Arch. Physiol., 78: 145 -48. b. New method for the determination of the blood volume in an- imals. Scand. Arch. Physiol., 78:189-96. With W. Bjerknes. Method for continual airbreathing in closed- circuit apparatus. Scand. Arch. Physiol., 79: 164-68. c. 1940 a. Experimental investigations on the respiratory function in div- ing mammals and birds. Hvalrad. Skr., no. 22. 131 pp.
PER FREDRIK THORKELSSON SCHOLANDER 401 b. On the respiratory adjustment to prolonged diving in the seal. Am. I. Physiol., 129:456-57. With L. Irving and S. W. Grinnell. Respiratory metabolism of the porpoise. Science, 91:455. d. With L. Irving and S. W. Grinnell. Experimental studies on the physiology of diving mammals. Science, 92:483. 1941 a. With L. Irving and S. W. Grinnell. The respiration of the por- poise, Tursiops truncates. ]. Cell. Comp. Physiol., 1 7: 145-68. b. With L. Irving. Experimental investigations on the respiration and diving of the Florida manatee. I. Cell. Comp. Physiol., 17: 169-91. c. With L. Irving and S. W. Grinnell. The depression of metabo- lism during diving. Am. i. Med. Sci., 202:915 -16. d. With L. Irving and S. W. Grinnell. Significance of the heart rate to the diving ability of seals. J. Cell. Comp. Physiol., 18:283- 97. 1942 a. Method for the determination of the gas content of tissue. I. Biol. Chem., 142:427-30. b. With L. Irving and S. W. Grinnell. Aerobic and anaerobic changes in seal muscles during diving. I. Biol. Chem., 142:431- 40. With L. Irving and S. W. Grinnell. The regulation of arterial blood pressure in the seal during diving. Am. J. Physiol., 135:557-66. ct. With L. Irving and S. W. Grinnell. On the temperature and metabolism of the seal during diving. J. Cell. Comp. Physiol., 19:67-78. With S. W. Grinnell and L. Irving. Experiments on the relation between blood flow and heart rate in the diving seal. I. Cell. Comp. Physiol., 19:341-50. f. Microburette. Science, 95: 177-78. g. Analyzer for one ml of respiratory gas. Rev. Sci. Instrum., 13:27-31. h. Volumetric microrespirometers. Rev. Sci. Instrum., 13:32-33. i. A micro-gas-analyzer. Rev. Sci. Instrum., 13:264-66. c.
402 BIOGRAPHICAL MEMOIRS With G. A. Edwards. Volumetric microrespirometer for aquatic organisms. Rev. Sci. Instrum., 13:292-95. k. Analyzer for quick estimation of respiratory gases. I. Biol. Chem., 146: 159 - 62. Microgasometric determination of nitrogen in blood and saliva. Rev. Sci. Instrum., 13: 362- 64. m. With G. A. Edwards. Nitrogen clearance from the blood and saliva by oxygen breathing. Am. J. Physiol., 137:715 -16. n. With L. Irving and G. A. Edwards. Experiments on carbon monoxide poisoning in tents and snow houses. i. Ind. Hyg., 24:213-17. With F. I. W. Roughton. A simple microgasometric method of estimating carbon monoxide in blood. I. Ind. five.. 24:218-21. O. ~ , ~ . p. With L. Irving and S. W. Grinnell. Experimental studies of the respiration of sloths. i. Cell. Comp. Physiol., 20:189-210. 1943 a. With L. Irving and S. W. Grinnell. Respiration of the armadillo with possible implications as to its burrowing. I. Cell. Comp. Physiol., 21 :53-63. With L. Irving and G. A. Edwards. Factors producing carbon monoxide from camp stoves. i. Ind. Hyg., 25: 132-36. With G. A. Edwards and L. Irving. Improved micrometer bur- ette. I. Biol. Chem., 148:495 -500. d. With F. J. W. Roughton. Micro gasometric estimation of the blood gases. I. Oxygen. }. Biol. Chem., 148:541-50. e. With F. i. W. Roughton. Micro gasometric estimation of the blood gases. II. Carbon monoxide. i. Biol. Chem., 148:551-63. f. With G. A. Edwards and F. J. W. Roughton. Micro gasometric estimation of the blood gases. III. Nitrogen. I. Biol. Chem., 148:565-71. . g. With F. I. W. Roughton. Micro gasometric estimation of the blood gases. IV. Carbon dioxide. i. Biol. Chem., 148:573-80. h. With N. Haugaard and L. Irving. A volumetric respirometer for aquatic animals. Rev. Sci. Instrum., 14:48 - 51. 1947 a. With L. Irving and O. Hebel. Apparatus for complete record- ing of respiratory exchange in man. Fed. Proc. Fed. Am. Soc. Exp.Biol.,6:134 - 35.
PER FREDRI K TH ORKELS SON S C H O LAN DER 403 b. Accurate analysis of respiratory gases in 0.5 cubic centimeter samples. Fed. Proc. Fed. Am. Soc. Exp. Biol., 6:197-98. Analyser for accurate estimation of respiratory gases in one- half cubic centimer samples. i. Biol. Chem., 167:235 -50. d. Simple syringe burette. Science, 105:581. With S. C. Flemister and L. Irving. Microgasometric estimation of the blood gases. V. Combined carbon dioxide and oxygen. }. Biol. Chem., 169: 173 -81. f. With H. }. Evans. Microanalysis of fractions of a cubic milli- meter of gas. I. Biol. Chem., 169:551- 60. g. With L. Irving. Micro blood gas analysis in fractions of a cubic millimeter of blood. i. Biol. Chem., 169:561-69. c. 1949 Volumetric respirometer for aquatic animals. Rev. Sci. Instrum., 20:885-87. 1950 a. Volumetric plastic micro respirometer. Rev. Sci. Instrum.q 21 :378-80. b. With H. Niemeyer and C. L. Claff. Simple calibrator for War- burg respirometers. Science, 112 :437-38. c. With V. Walters, R. Hock, and L. Irving. Body insulation of some arctic tropical mammals and birds. Biol. Bull. Woods Hole, Mass., 99:225-36. d. With R. Hock, V. Walters, and L. Irving. Heat regulation in some arctic and tropical mammals and birds. Biol. Bull. Woods Hole, Mass., 99:237-58. e. With R. Hock, V. Walters, and L. Irving. Adaptation to cold in arctic and tropical mammals and birds in relation to body tem- perature, insulation, and basal metabolic rate. Biol. Bull. Woods Hole, Mass., 99: 259-71. 1951 a. Nitrogen tension in the swimbladder of marine fishes in rela- tion to the depth. Fed. Proc. Fed. Am. Soc. Exp. Biol., 10:121. b. With C. L. Claff, C. T. Teng, and V. Walters. Nitrogen tension in the swimbladder of marine fishes in relation to the depth. Biol. Bull. Woods Hole, Mass., 101:178-93. c. With H. Erikson and L. Irving. Apparatus for complete volu
404 BIOGRAPHICAL MEMOIRS metric recording of the respiratory gaseous exchange in man. Scand. i. Clin. Lab. Invest., 3:228-33. 1952 c' a. With C. L. Claff, }. R. Andrews, and D. F. Wallach. Microvolu- metric respirometry. I. Gen. Physiol., 35:375-95. b. With C. L. Claff and S. L. Sveinsson. Oxygen consumption dur- ing the cleavage of single cells. Fed. Proc. Fed. Am. Soc. Exp. Biol.,11:141. With C. L. Claff and S. L. Sveinsson. Respiratory studies of single cells. I. Methods. Biol. Bull. Woods Hole, Mass., 102: 157- 77. d. With C. L. Claff and S. L. Sveinsson. Respiratory studies of single cells. II. Observations on the oxygen consumption in single protozoans. Biol. Bull. Woods Hole, Mass., 102:178-84. e. With C. L. Claff, S. L. Sveinsson, and Susan I. Scholander. Res- piratory studies of single cells. III. Oxygen consumption during cell division. Biol. Bull. Woods Hole, Mass., 102: 185-99. With i. H. Kinoshita and I. P. Bunker. The use of the volumetric respirometer in the determination of plasma carbon dioxide. I. Lab. Clin. Med., 40:156-60. g. With }. Wyman, fir., G. A. Edwards, and L. Irving. On the sta- bility of gas bubbles in sea water. J. Mar. Res., 11 :47-62. h. With W. Flagg, V. Walters, and L. Irving. Respiration in some arctic and tropical lichens in relation to temperature. Am. I. Bot., 39: 707-13. i. With R. I. Hock, H. Erikson, W. Flagg, and L. Irving. Compo- sition of the ground-level atmosphere at Point Barrow, Alaska. i. Met., 9:441-42. 1953 a. With W. Flagg, V. Walters, and L. Irving. Climatic adaptation in arctic and tropical poikilotherms. Physiol. Zool., 26:67-92. b. With L. van Dam. Composition of the swimbladder gas in deep sea fishes. Biol. Bull. Woods Hole, Mass., 104:75-86. c. With L. van Dam. Concentration of hemoglobin in the blood of deep sea fishes. }. Cell. Comp. Physiol., 41:522-24. d. Studies on the physiology of frozen plants and animals in the Arctic. Abstracts Communications, 19th Int. Physiol. Congr., pp. 741-42.
e. PER FREDRIK THORKELSSON SCHOLANDER 405 With W. Flagg, R. J. Hock, and L. Irving. Studies on the phys- iology of frozen plants and animals in the Arctic. }. Cell. Comp. Physiol., 42, suppl. 1. 56 pp. 1954 a. With L. van Dam. Secretion of gases against high pressures in the swimbladder of deep sea fishes. I. Oxygen dissociation in blood. Biol. Bull. Woods Hole, Mass., 107:247-59. b. Secretion of gases against high pressures in the swimbladder of deep sea fishes. II. The rete mirabile. Biol. Bull. Woods Hole, Mass., 107:260-77. 1955 a. With L. van Dam and Susan I. Scholander. Gas exchange in the roots of mangroves. Am. }. Bot., 42 :92-98. b. Evolution of climatic adaptation in homeotherms. Evolution, 9:15-26. With W. E. Love and i. Id. Kanwisher. The rise of sap in tall grapevines. Plant Physiol., 30:93-104. d. With L. van Dam, C. L. Clam, and J. W. Kanwisher. Micro ga- sometric determination of dissolved oxygen and nitrogen. Biol. Bull. Woods Hole, Mass., 109:328-34. e. With W. E. Schevill. Counter-current vascular heat exchange in the fins of whales. J. Appl. Physiol., 8:279-82. Hydrostatic pressure in coconuts. Plant Physiol., 30:560-61. g. With L. van Dam and T. Enns. Secretion of inert gases and oxygen by the swimbladder of fishes. Biol. Bull. Woods Hole, Mass., 109:338-39. h. With Alan C. Burton and Otto G. Edholm. Man in a cold en- vironment. Physiological and pathological ejects of exposure to low temperatures. Scand. }. Clin. Lab. Invest., 7:349. c. 1956 a. With L. van Dam and T. Enns. Nitrogen secretion in the swim- bladder of whitefish. Science, 123 :59-60. b. With }. W. Kanwisher and D. C. Nutt. Gases in icebergs. Science, 123: 104-5. c. With H. Erikson, }. Krog, and K. Lange Andersen. The critical temperature in naked man. Acta Physiol. Scand., 37:35-39. d. Climatic rules. Evolution, 10:339-40.
406 BIOGRAPHICAL MEMOIRS With L. K. Coachman and E. Hemmingsen. Gas enclosures in a temperate glacier. Tellus, 8:415 -23. f. With L. van Dam and T. Enns. The source of oxygen secreted into the swimbladder of cod. }. Cell. Comp. Physiol., 48:517- 22. g. Observations on the gas gland in living fish. J. Cell. Comp. Phys- iol., 48:523-28. h. With L. van Dam. Micro gasometric determination of oxygen in fish blood. }. Cell. Comp. Physiol., 48:529-32. 1957 a. With Berthe Ruud and H. Leivestad. The rise of sap in a trop- ical liana. Plant Physiol., 32: 1-6. b. With L. van Dam. The concentration of hemoglobin in some cold water arctic fishes. }. Cell. Comp. Physiol., 49: 1-4. c. With L. van Dam, i. W. Kanwisher, H. T. Hammel, and M. S. Gordon. Supercooling and osmoregulation in arctic fish. J. Cell. Comp. Physiol., 49:5 -24. d. With K. Lange Andersen, }. Krog, F. Vogt Lorentzen, and }. Steen. Critical temperature in Lapps. J. Appl. Physiol., 10:231- 34. e. With H. T. Hammel, K. Lange Andersen, and Y. L0yning. Met- abolic acclimation to cold in man. Fed. Proc. Fed. Am. Soc. Exp. Biol., 16:114-15. f. "The wonderful net." Sci. Am., 196~4) :96 -107. g. With i. Krog. Countercurrent heat exchange and vascular bundles in sloths. }. Appl. Physiol., 10:405-11. h. With H. Leivestad and H. Andersen. Physiological response to air exposure in codfish. Science, 126:505. i. Oxygen dissociation curves in fish blood. Acta Physiol. Scand., 41 :340-44. J k . With H. T. Andersen and H. Leivestad. "Air diving" in fishes. Acta Physiol. Scand., 42, suppl. 145:6-7. With H. T. Hammel, K. Lange Andersen, and Y. L0yning. Met- abolic acclimation to cold in man. Acta Physiol. Scand., 42, suppl. 145:63-64. With }. Krog. Counter current vascular heat exchange, with special reference to the arteriovenous bundles in sloths. Acta Physiol. Scand., 42, suppl. 145: 89-90.
PER FREDRIK THORKELS SON S C H O LAN DER 407 m. With G. Sundnes. Gas secretion in fishes lacking "rete mirabile." Acta Physiol. Scand., 42, suppl. 145:125-26. 1958 a. With H. T. Hammel, K. Lange Andersen, and Y. L0yning. Met- abolic acclimation to cold in man. i. Appl. Physiol., 12: 1-8. b. The rise of sap in lianas. In: The Physiology of Forest Trees, ed. Kenneth V. Thimann, pp. 3-17. New York: Ronald Press. c. With L. K. Coachman, E. Hemmingsen, T. Enns, and H. de Vries. Gases in glaciers. Science, 127:1288-89. d. With H. Jensen. Bag spirometer. Scand. i. Clin. Lab. Invest., 10:225-26. e. With H. T. Hammel, I. S. Hart, D. H. LeMessurier, and I. Steen. Cold adaptation in Australian aborigines. }. Appl. Physiol., 13:211-18. With G. Sundnes and T. Enns. Gas secretion in fishes lacking rete mirabile. i. Exp. Biol., 35:671-76. g. With L. K. Coachman and T. Enns. Gas loss from a temperate glacier. Tellus, 10:493 -95. h. Counter current exchange. A principle in biology. Hvalrad. Skr., no. 44. 24 pp. . . . . i. Studies on man exposed to cold. Fed. Proc. Fed. Am. Soc. Exp. Biol., 17: 1054 - 57. With H. Leivestad and G. Sundnes. Cycling in the oxygen con sumption of cleaving eggs. Exp. Cell. Res., 15:505-11. k. With O. Iversen. New design of volumetric respirometer. Scand. i. Clin. Lab. Invest., 10:429-31. 1959 a. Wave-riding dolphins: How do they do it? Science, 129:1085- 87. b. Experimental studies on asphyxia in animals. In: Oxygen Supply to the Human Foetus, ed. lames Walker and A. C. Turnbull, pp. 267-74. Oxford: Blackwell Scientific Publications. c. Supercooling and freezing in poikilotherms. In: Simposios Con- ferencias, 21st Congr. Int. Ciencias Fisiol., Buenos Aires, pp. 77- 81. d. Wave-riding dolphins. Science, 130: 1658.
408 b. BIOGRAPHICAL MEMOIRS 1960 Oxygen transport through hemoglobin solutions. Science, 131:585-90. Oxygen diffusion. Science, 132:368. With D. C. Nutt. Bubble pressure in Greenland icebergs. I. Gla- ciol., 3:671-78. d. With E. Hemmingsen. Specific transport of oxygen through hemoglobin solutions. Science, 132: 1379-81. Man in cold environment. Discussion. Fed. Proc. Fed. Am. Soc. Exp. Biol., 19, suppl. 5: 8-10, 11-12. 1961 a. With H. T. Hammel, D. H. LeMessurier, E. Hemmingsen, and W. Garey. Circulatory adjustment in pearl divers. Fed. Proc. Fed. Am. Soc. Exp. Biol., 20:103. b. With D. C. Nutt and L. K. Coachman. Dissolved nitrogen in West Greenland waters. i. Mar. Res., 10:6-11. With E. A. Hemmingsen, L. K. Coachman, and D. C. Nutt. Composition of gas bubbles in Greenland icebergs. }. Glaciol., ~, 3:813-22. d. With E. Hemmingsen and W. Garey. Cohesive lift of sap in the rattan vine. Science, 134: 1835 -38. 1962 a. With M. S. Gordon and B. H. Amdur. Freezing resistance in some northern fishes. Biol. Bull. Woods Hole, Mass., 122:52- 62. With H. T. Hammel, D. H. LeMessurier, E. Hemmingsen, and W. Garey. Circulatory adjustment in pearl divers. J. Appl. Phys- iol., 17: 184-90. c. With C. R. Olsen and D. D. Fanestil. Some effects of breath holding and apneic underwater diving on cardiac rhythm in man. J. Appl. Physiol., 17:461-66. d. With Edda Bradstreet. Microdetermination of lactic acid in blood and tissues. }. Lab. Clin. Med., 60:164-66. e. With Edda Bradstreet and W. F. Garey. Lactic acid response in the grunion. Comp. Biochem. Physiol., 6:201-3. With R. W. Elsner and E. Hemmingsen. The work of maintain- ing flotation in sea water. Physiologist, 5: 136.
PER FREDRIK THORKELSSON SCHOLANDER 409 g. With W. Dansgaard, D. C. Nutt, H. de Vries, L. K. Coachman, and E. Hemmingsen. Radio-carbon age and oxygen-18 content of Greenland icebergs. Medd. Gronl., 165(1). 26 pp. h. With H. T. Hammel, E. Hemmingsen, and W. Garey. Salt bal 1. ance in mangroves. Plant Physiol., 37:722-29. With C. R. Olsen and D. D. Fanestil. Some effects of apneic underwater diving on blood gases, lactate, and pressure in man. J. Appl. Physiol., 17 :938-42. i. Physiological adaption to diving in animals and man. Harvey Lect., 57:93 -110. 1963 a. With R. W. Elsner and W. F. Garey. Selective ischemia in diving man. Am. Heart }., 65:571-72. b. The master switch of life. Sci. Am., 209~6~:92-106. 1964 Animals in aquatic environments: Diving mammals and birds. In: Handbook of Physiology. Sect. 4: adaptation to the environment, ed. D. B. Dill, pp. 729-39. Bethesda, Md.: American Physio- logical Society. b. With T. Enns. Oxygen transport by hemoglobin collision. Fed. Proc. Fed. Am. Soc. Exp. Biol., 23:468. c. With H. T. Hammel, E. A. Hemmingsen, and Edda D. Brad- street. Hydrostatic pressure and osmotic potential in leaves of mangroves and some other plants. Proc. Natl. Acad. Sci., USA, 52:119-25. d. With R. W. Elsner, A. B. Craig, E. G. Dimond, L. Irving, M. Pilson, K. Johansen, and Edda Bradstreet. A venous blood ox- ygen reservoir in the diving elephant seal. Physiologist, 7:124. From the frozen forest to tropical mangroves. In: Program Ab- stracts, Pt. 2, Proc. Alaska Sci. Conf., p. 38. Washington, D.C.: American Association for the Advancement of Science. 1965 a. With H. T. Hammel, Edda D. Bradstreet, and E. A. Hemming sen. Sap pressure in vascular plants. Science, 148:339 - 46. b. With T. Enns and E. D. Bradstreet. Effect of hydrostatic pres sure on gases dissolved in water. t. Phys. Chem., 69:389-91.
410 BIOGRAPHICAL MEMOIRS c. With H. T. Hammel, E. D. Bradstreet, and E. A. Hemmingsen. Sap pressure in plants. Science, 149:920 - 21. d. Tension gradients accompanying accelerated oxygen transport in a membrane. Science, 149:876-77. e. Reverse osmosis and sap pressure in vascular plants. Science, 150:384. f. With R. Elsner. Circulatory adaptations to diving in animals and Nan. In: Physiology of Breath-Hold Diving and the Ama of Japan, pp. 281-94. Washington, D.C.: National Academy of Sciences Publ. 131. 1966 a. With E. D. Bradstreet, H. T. Hammel, and E. A. Hemmingsen. Sap concentrations in halophytes and some other plants. Plant Physiol., 41:529-32. b. The role of solvent pressure in osmotic systems. Proc. Natl. Acad. Sci. USA, 55:1407-14. 1967 a. Osmotic mechanism and negative pressure. Science, 156:67- 69. b. Negative pressure in plants and osmotic mechanism. Science, 156:541. c. Osmotic pressure. Science, 158: 1212. d. With T. Enns and E. Douglas. Role of the swimbladder rete of fish in secretion of inert gas and oxygen. Adv. Biol. Med. Phys., 2:231-44. 1968 a. With R. W. Elsner. A comparative view of cardiovascular de- fense against acute asphyxia. Proc. 2nd Int. Symp. Emergency Resuscitation, Oslo. Acta Anaesthesiol. Scand., suppl. 29:15- 33. b. How mangroves desalinate seawater. Physiol. Plant,21:251-61. c. With A. R. Hargens and S. L. Miller. Negative pressure in the interstitial fluid of animals. Science, 161:321-28. d. With R. S. Bandurski and E. Bradstreet. Metabolic changes in the mud-skipper during asphyxia or exercise. Comp. Biochem. Physiol., 24:271-74.
PER FREDRIK THORKELSSON SCHOLANDER 411 e. With M. de Oliveira Perez. Sap tension in flooded trees and bushes -of the Amazon. Plant Physiol., 43: 1870-73. 1969 a. With L. Irving, E. A. Hemmingsen, and E. Bradstreet. Ultra- violet absorption in the cornea of arctic and alpine animals. In: The Biologic Elects of Ultraviolet Radiation, ed. F. Urbach, pp. 469-71. London: Pergamon Press. b. With S. B. Stromme and i. E. Maggert. Interstitial fluid pres- sure in terrestrial and semiterrestrial animals. I. Appl. Physiol., 27: 123-26. c. With A. R. Hargens. Stretch mounting for osmotic membranes. Microvasc. Res., 1 :417-19. 1971 a. State of water in osmotic processes. Microvasc. Res., 3:215-32. b. Imbibition and osmosis in plants. In: Topics in the Study oiLife: The Bio Source Book, ed. Amy Kramer, pp. 138-47. New York: Harper & Row. With M. Perez. Experiments on osmosis with magnetic fluid. Proc. Natl. Acad. Sci. USA, 68:1093-94. d. With i. E. Maggert. Supercooling and ice propagation in blood from arctic fishes. Cryobiology, 8:371-74. With M. Perez. Effect of gravity on osmotic equilibria. Proc. Natl. Acad. Sci. USA, 68:1569-71. c. 1972 a. With M. Perez. Molecular buoyancy and osmotic equilibrium. Proc. Natl. Acad. Sci. USA, 69:301-2. b. Tensile water. Am. Sci., 60:584-90. 1973 With H. T. Hammel. Thermal motion and forced migration of colloidal particles generate hydrostatic pressure in solvent. Proc. Natl. Acad. Sci. USA, 70:124-28. 1975 Water states and water gates in osmotic processes, and the inop- erative concept of molfraction of water. l. Exp. Zool., 194:241- 48.
412 BIOGRAPHICAL MEMOIRS 1976 With H. T. Hammel. Osmosis and Tensile Solvent. Berlin: Springer Verlag. 1978 c. a. Rhapsody in science. Annul Rev. Physiol., 40:1-17. b. Water under tension, its fundamental role in capillarity, osmosis and colligative properties. In: Frontiers Or Human Knowledge, Skrifter rorande Uppsala universitet, C:38, Acta Univ. Ups. Nova Acta Regiae Soc. Sci. Ups. Ser. VC, pp. 297-308. With A. R. Hargens and W. L. Orris. Positive tissue fluid pres- sure in the feet of antarctic birds. Microvasc. Res., 15:239-44.