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WALLACE OSGOOD PENN Al ugust 27, 1893-September 20, 1971 BY HERMANN RAHN WALLACE OSGOOD FENN was born in Lanesboro (Berkshire County), Massachusetts and died in Rochester, New York in his seventy-ninth year, after a brief illness. He is survived by his widows, Clara Bryce (Comstock) Fenn; his children, William Wallace Fenn, Ruth (Fenn) Starman, Priscilla (Fenn) Roslansky, and David Bryce Fenn; and ten grandchildren. He led a most vigorous life and up to his very last days was working in the laboratory; during his last three years he shaped the new direc- tions of the International Union of Physiological Sciences as its President. To many of his colleagues he was the Dean of Physio- logical Sciences, the last Renaissance Man, whose basic contri- butions covered so many areas and who had a remarkable per- spective on the whole field of biology. His forefathers settled in New England in the seventeenth century. William Wallace Fenn, his father, was a Unitarian minister who had married Faith Huntington Fisher, also from New England. Later his father became the Bussey Professor of Theology at Harvard and Dean of the Divinity School. Thus Wallace Fenn's childhood was spent in Cambridge, where he attended the Cambridge Latin School and entered Harvard with the goal of preparing himself for the ministry. However, when he started cutting his father's lectures to attend plant physi- ologist W. J. V. Osterhout's classes in biology, the foundations 141

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142 BIOGRAPHICAL MEMOIRS were laid for a career in physiology that was to span more than half a century. Fenn graduated in 1914. His graduate work at Harvard with Osterhout was interrupted by World War I, during which he served in the Sanitary Corps of the U.S. Army and was com- missioned a Second Lieutenant. Upon discharge in 1919, he fin- ished his doctoral thesis in Curie, married Clara Bryce Comstock in September, and began his appointment as an Instructor in Applied Physiology in the Department of Industrial Hygiene at the Harvard Medical School under Cecil K. Drinker. Here began his classical studies of phagocytosis of solid particles by white blood corpuscles. In 1922 he accepted a Rockefeller Travel Fellowship and was the first American to work in A. V. Hill's laboratory in London, England. This was followed by a six-month stay in H. H. Dale's laboratory at the National Institute for Medical Research in London. Returning to this country in 1924, he accepted the Chair of Physiology at the newly formed Medical School at the University of Rochester, New York. This position he filled for thirty-five years. In 1961 he was named Distinguished Professor of Physiology, a post he occupied until his death in 1971. THE SCIENTIST Fenn's first paper was published in 1916 in the Proceedings of the National A cademy of Sciences. It was entitled "Salt Antagonism in Gelatin." His last paper, "Partial Pressure of Gases Dissolved at Great Depth," was published posthumously in Science in 1972. During the intervening half-century his 267 publications can be conveniently divided into four general areas: the physiology of muscle, electrolytes, respiration, and high pressure. In each area he laid foundations of new concepts, and when he was satisfied that he had made new basic contributions, moved on to explore new fields.

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WALLACE OSGOOD FENN CONTRIBUTIONS TO MUSCLE AND ELECTROLYTE PHYSIOLOGY 143 The work that brought Fenn his first recognition was his study on the heat production of muscle, which he started in A. V. Hill's laboratory in 1922-1923. Fenn wrote: "In particular it can now be shown that there is a fairly good quantitative relation between the heat production of muscles and the work which they perform, and that a muscle which does work liberates, ipso facto, an extra supply of energy which does not appear in an isometric contraction."3t It was A. V. Hill who referred to this as the Fenn Effect, and so it has been known ever since. Fenn's heat data showed first of all that if a muscle shortens, no matter how little and no matter how lightly loaded, it pro- duces more heat than during an isometric contraction over the same time period. He then showed that this extra heat produc- tion was proportional to the external work done by the muscle. It was clearly not determined by the load alone, nor by the change in length. This was the first evidence, and remains today the best evidence, that shortening is an active process and that muscle is riot simply a prestretched spring shortening passively. The Fenn Effect has emerged as the nearest thing to a law that muscle physiologists have. Following his pioneer work on muscle heat production, Fenn began to measure gas exchange by nerve and by muscle. To this end he had to invent a number of ingenious instruments to obtain the necessary specificity and precision. In 1927 he meas- ured for the first time the quantitative amount of oxygen required by a nerve to conduct an impulse. Similar studies on the metabolism of contracting muscles led him to consider the role of electrolytes, particularly potassium, in nerve and muscle Wallace Osgood Fenn, "A Quantitative Comparison between the Energy Liberated and the Work Performed by the Isolated Sartorius Muscle of the Frog," Journal of Physiology, 58(1924): 175.

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144 BIOGRAPHICAL MEMOIRS activity. At the time, although it was known that muscle fibers were rich in potassium, almost nothing was known of the mecha- nisms by which cells accumulated and maintained a high potas- sium content. The work ushered in the era of electrolyte physiology. Begin- ning in 1933 Fenn virtually created the field of potassium metab- olism. He made the first determinations of potassium, sodium, magnesium, and calcium in nerve. He developed a new method for determining internal pH of muscle and nerve and obtained values that remain acceptable today. He showed that intra- cellular potassium was mobile, not fixed, and that muscle potas- sium shifted in response to various environmental factors. Most importantly, he showed that during contraction potas- sium was lost from muscle in exchange for sodium, and that the process was reversed in recovery. For the first time he showed that sodium could penetrate muscle. These observations were clearly the necessary foundation for the Hodgkin-Huxley hypo- theses concerning initiation and propagation of nerve and muscle impulses and the magnitude and polarity of electrical potential differences across cell membranes. As early as 1936, at the Cold Spring Harbor Symposium, Fenn said, "The explana- tion of a loss of potassium from a muscle during activity is a matter of fundamental theoretical importance. In terms of the theory which I have been using as a guide, it is interpreted as an increase in the permeability of the muscle membrane of sufficient extent to permit sodium, but not chloride, to enter. Every mole- cule of sodium which enters then displaces one molecule of potassium." ~ Fenn showed that potassium escaped from muscle during contraction in situ and that a large part of this potassium ap- peared in the liver. He demonstrated that potassium uptake was #Wallace Osgood Fenn, "Electrolytes in Muscle," Cold Spring Harbor Symp. Quant. Viol., 4(1936):252-59.

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WALLACE OSGOOD FEN N 145 linked with carbohydrate metabolism, particularly with glycogen deposition, and developed the concept that potassium tends to follow the Cori cycle. He was always quick to seize new oppor- tunities. When radioactive potassium became available to him in 1939, he ingested a sample. Using himself as subject, he was thus the first not only to study the kinetics of potassium metabo- lism but also to demonstrate potassium incorporation into blood cells, previously thought to be impermeable. He showed that nearly all muscle potassium in the body is exchangeable, proving that high intracellular potassium content is not maintained by binding or sequestration of potassium, an idea which was con- sonant with his notion that potassium is maintained by an active energetic process. CONTRIBUTIONS TO RESPIRATION PHYSIOLOGY The entrance of Wallace 0. Fenn into the history of respira- tory physiology can be precisely dated. It was within days after the U.S. entry into World War II. At that time he was forty-eight years old and had established himself as the acknowledged leader in the physiology of muscle and electrolytes. He was to be recog- nized in 1943 by election to the National Academy of Sciences. Wallace Fenn was drawn into respiratory physiology by his desire to contribute to the war effort. This was to be largely a war in the air, and from a military point of view, supremacy in altitude tolerance meant supremacy of air power. The airplanes of that day did not yet have pressurized cabins, but the possi- bility occurred that the human lung might be pressurized by application of positive pressure breathing. The question was whether man's lungs could tolerate a sufficient amount of pres- sure to raise the partial pressure of oxygen to a significant degree, or would the lungs rupture, or would the circulation stop? What were the limiting factors? What were the hazards? What was known about respiratory physiology in general? This can best be answered by listing some terms which did not

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146 BIOGRAPHICAL MEMOIRS appear in the physiology textbooks of that era, but which are commonplace today. Such terms are: positive and intermittent pressure breathing, pressure-volume diagram, work of breath- ing, pulmonary compliance, airway resistance, alveolar gas equa- tion, O'-CO2 diagram, ventilation-perfusion ratio, just to name a few. Wallace Fenn had never worked in the field of human respiration. The equipment in his laboratory would be regarded as primitive by current standards. Among the more useful items were a few assorted spirometers, two or three Haldane machines, an equal number of Van Slykes, and several U-tube manometers. The most sophisticated instrument was a Millikan ear oximeter, which had been loaned to him by the Military. It carried a security classification of a fairly high level, and since no instruc- tion manual came with it, it took some time and a visit to Glen Millikan himself before anyone could figure out how to use it properly. In addition to this modest inventory of equipment, Fenn had three young instructors, all trained in biology departments. They knew all about such things as how fast the drosophila can beat its wings, how and why the rattlesnake changes color, and how to activate or inhibit enzymes found in grasshopper eggs, but none of them had ever blown a vital capacity; neither did they know the difference between complemental and supplemen- tal air. L. E. Chadwick, A. B. Otis, and H. Rahn, living with their wives on postdoctoral stipends which were only a fraction of what a graduate student receives today, were the most unlikely crew to have been assembled for the unknown job that lay ahead of them. Neither the equipment nor the staff was very impressive, and it seems doubtful that by present standards the project could have qualified for a National Institutes of Health grant. How- ever, the major asset, recognizable even then, was Wallace Fenn himself. He was not put off by lack of ready-made equipment; he

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WALLACE OSGOOD FEN N 147 was well endowed with Yankee ingenuity, and he loved to impro- vise. He could, with whatever components happened to be handy, construct apparatus that would perform in a reliable and effective fashion. Everyone associated with him has memories of him in the laboratory surrounded by what at first sight appeared to be an unrelated jumble of strange wires and rubber bands, tubing, pulleys, lenses, light sources, mirrors, and other assorted bits and pieces. A more careful examination suggested there might be some order in the arrangement, and further observa- tion would reveal that something of physiological interest was actually being measured and perhaps even graphically recorded. A relatively refined example was a device for the automatic re- cording of blood flow through the finger and its alteration by pressure breathing. The high-altitude chamber was perhaps the crowning master- piece of Fenn's ingenuity. He had received from the Committee on Medical Research of the Office of Scientific Research and Development a contract which provided the sum of $500 (five hundred!) for special research equipment. From this budget he bought a steel tank designed for the processing or transport of beer, commandeered the tree-spraying pump from the Univer- sity Grounds Department, reversed its valves, and connected Dumb to tank. The result was a chamber which could go to 1 ~ . . simulated altitudes at the rate of 5,000 feet per minute. As he later said, "It surely was the worst high altitude chamber in the country, but a rare atmos ?here is the same wherever you find it." Not only could he get the most out of primitive pieces of equipment, but he also seemed somehow able to evoke the best output from his staff. He did not tell people to do things. Rather, he pointed out things that needed doing and waited for some- #Wallace Osgood Penn, "Born Fifty Years Too Soon," Annual Review of Physiology, 24(1962):1.

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148 BIOGRAPHICAL MEMOIRS thing to happen. He worked hard himself and expected others to do likewise, but he recognized that there were individual differ- ences in effective work patterns and did not try to impose his own habits on others. Although he kept rather regular working hours himself, he apparently was not perturbed by those with more erratic habits. Getting something done rather than com- pulsive adherence to a fixed schedule was the important thing. In starting a new experiment he frequently took the lead by setting up apparatus himself rather than asking someone else to do it. Typically, he would insist on being the first subject in a new experimental procedure, and in experiments with pressure breathing and in the altitude chamber he extended himself on a number of occasions to the point of losing consciousness. He was a pioneer in every sense, and it was a blessing that his work ante- dated the Human Subjects Review Committee. Fenn's intuitive approach to and logical analysis of the pres- sure breathing problem led him to develop two powerful con- cepts and to express them in the form of graphic relationships: the pressure-volume diagram of the lung and thorax, and the O2-CO2 diagram of the composition of alveolar gas. Although the basic pressure-volume (P-V) diagram had been previously developed by F. Rohrer, Penn conceived it independ- ently, elaborated it further, and distilled into it some ten years of work and thought. Like all his work, it defined physiological boundaries, limiting values for muscle forces and the corre- sponding volumes of gas and blood. Within these limits were centered the normal operating range of pulmonary mechanics and the response of the system to positive and negative pressure breathing. It was not only a beautiful composition both artistic- ally and scientifically, but it was also the foundation and frame- work of respiratory mechanics that would be further embellished by students during succeeding decades. Fenn's second masterpiece, the O2-CO2 diagram, did for pulmonary gas exchange what the P-V diagram did for respira- tory mechanics. With it he could represent all parameters of the

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WALLACE OSGOOD FENN 149 alveolar gas and ventilation equations. He never claimed to have originated these equations, but he derived them independently, made sure they were correct, and put them in graphic form. As somebody put it, "That's when he made them sing." On the diagram he could show all possible compositions of alveolar gas and the arterial blood under any specified set of conditions. He could indicate normal ranges and limits of survival as well as the pathways followed during hyperventilation and asphyxia and during exposure to CO2, altitude, or hyperbaric pressures. It could be used to demonstrate ranges of normal and impaired performance. It was indeed a theme that could be sung with . . many varlatlons. Although the P-V and O2-CO2 diagrams represent great masterpieces of Fenn's scientific artistry, he created, inspired, or contributed to many other works. To give a few examples: devel- opment of the concept of an optimal breathing frequency, meas- urement of alveolar pressure, dynamic pressure-volume curves presented for the first time on a cathode ray oscilloscope, devel- opment of an infrared CO2 meter, and probably the first pub- lished continuous recording of CO2 changes during a single breath. Finally, one must mention two special contributions to respiration physiology, a lasting monument to his effort in this area: his book, ~4 Graphical Analysis of the Respiratory Gas Exchange, which went through many reprintings, and his editor- ship of Respiration in the Handbook of Physiology series, both of these published by The American Physiological Society. CONTRIBUTIONS TO THE PHYSIOLOGY OF SPACE AND OCEANS From the mid-1950s Fenn became greatly intrigued with two new frontiers that began to unfold man's explorations in space and the ocean depths. While his research continued in very basic experiments, their application was obviously directed to filling in the gaps of knowledge so that man could exist successfully in

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150 BIOGRAPHICAL MEMOIRS these new environments. He was in great demand as a consultant by space physiologists and tried to convince his more earth- bound colleagues of the great new opportunities in physiology that unfolded as man ventured into space. Every problem Fenn "considered basic, if the investigator put some basic thinking into it." In that sense he felt that physiology as a science had gained immeasurably and would continue to grow as man went forth in orbit and into the oceans thoughts that he expressed so well in his address "Physiology in Orbit."* Wherever man went he needed oxygen as the life-sustaining gas, yet when it exceeded normal pressures it became poisonous. Fenn spent many years with his associates (R. Gerschman and D. L. Gilbert) in trying to understand the toxic nature of oxy- gen. Probably his most important insight was the recognition and demonstration that oxygen poisoning and X-irradiation effects have the same common mechanism. He also turned his attention to the effects of high inert gas pressures upon the metabolism of unicellular organisms and the effects of hydraulic pressure on biological reactions. His last benchwork emphasized the importance of partial molar volume concepts as a tool for determining the volume that Of occupies within the hemoglobin structure." His last research concerned itself with the theoretical concepts of partial pressures of gases dissolved at great depths. It was a thermodynamic interpretation published posthumously in Science,+ where with his great mod- esty he asked for the help of physical chemists to develop this concept in greater detail, help which shortly appeared. Wallace Osgood Fenn, "Physiology in Orbit," The Physiologist, 3(1960):20-26. ~ Wallace Osgood Fenn, "Partial Molar Volumes of Oxygen and Carbon Monoxide in Blood," Respiratory Physiology, 13~1971~ :129~0. ~ Wallace Osgood Fenn, "Partial Pressure of Gases Dissolved at Great Depth," Science, 176(1972):1011-12. F. C. Andrews, "Gravitational Effects on Concentrations and Partial Pressures in Solutions: A Thermodynamic Analysis," Science' 178(1972):1199-1201.

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WALLACE OSGOOD FENN 163 With M. Goettsch. Electrolytes in nutritional muscular dystrophy in rabbits. J. Biol. Chem., 120:41-50. Loss of potassium from stimulated frog muscle. Proc. Soc. Exp. Biol. Med., 37:71-74. 1938 With D. M. Cobb, i. F. Manery, and W. B. Bloor. Electrolyte changes in cat muscle during stimulation. Am. J. Physiol., 121: 595-608. With J. H. Wills. Potassium changes in submaxillary glands during stimulation. Am. l. Physiol., 124:72-76. Factors affecting the loss of potassium from stimulated muscles. Am. J. Physiol., 124:213-29. With B. H. Carleton. Studies on respiration of muscle in the presence of carbon monoxide. l. Cell. Comp. Physiol., 11:91-98. The potassium and water contents of cat nerves as affected by stimu- lation. J. Neurophysiol., 1: 1-3. 1939 The fate of potassium liberated from muscles during activity. Am. I. Physiol., 217: 356-73. With W. S. Wilde, R. A. Boak, and R. H. Koenen~ann. The effect of blood flow on potassium liberation from muscle. Am. J. Physiol., 128: 139~6. The deposition of potassium and phosphate with glycogen in rat livers. l. Biol. Chem., 128:297-307. The distribution of excess potassium in cats. In: Professor Alvaro e Miguel Ozorio de Almeida, Livro de Homenagem, pp. 197-202, Rio de ~aneiro, Brazil. 1940 With R. H. Koenemann, B. V. Favata, and E. T. Sheridan. The role of the lactic acid in the movements of potassium. Am. i. Physiol., 131 :494-508. With L. F. Haege. The deposition of glycogen with water in the livers of cats. T. Biol. Chem., 136:87-101. With R. H. Koenemann and E. T. Sheridan. Potassium exchange of

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164 BIOGRAPHICAL MEMOIRS perfused frog muscle during asphyxia. J. Cell. Comp. Physiol., 16:255-64. The role of potassium in physiological processes. Physiol. Rev., 20: 377-415. 1941 With T. R. Noonan and L. F. Haege. The distribution of injected radioactive potassium in rats. Am. J. Physiol., 132:474-88. With T. R. Noonan and L. F. Haege. The effects of denervation and of stimulation on exchange of radio-active potassium in muscle. Am. J. Physiol., 132: 612-21. With L. J. Mullins, T. R. Noonan, and L. F. Haege. Permeability of erythrocytes to radioactive potassium. Am. J. Physiol., 135: 93-101. With T. R. Noonan, L. l. Mullins, and L. F. Haege. The exchange of radioactive potassium with body potassium. Am. J. Physiol., 135: 149-163. Muscle. Annul Rev. Physiol., 3:209-32. Preface. In: Biological Symposia, ed. l. Cattell, 3:vii-ix. Lancaster, Pa.: Cattell Press. Introduction to muscle physiology. In: Biological Symposia, ed. I. Cattell, 3: 1-8. Lancaster, Pa.: Cattell Press. With R. B. Dean, T. R. Noonan, and L. F. Haege. Permeability of erythrocytes to radioactive potassium. J. Gen. Physiol., 24:353-65. 1942 With L. F. Haege. The penetration of magnesium into frog muscle. J. Cell. Comp. Physiol., 19:37-46. With W. F. Bale and L. J. Mullins. The radioactivity of potassium from human sources. l. Gen. Physiol., 25:345-53. 1944 With L. F. Haege, E. Sheridan, and l. B. Flick. The penetration of ammonia into frog muscle. l. Gen. Physiol., 28: 53-77. 1945 Muscles. In: Physical Chemistry of Cells and Tissues, ed. R. Hober. Philadelphia: Blakiston.

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WALLACE OSGOOD FEN N 1946 16S With H. Rahn, A. B. Otis, and L. E. Chadwick. The pressure volume diagram of the thorax and lung. Am. l. Physiol., 146: 161-78. With A. B. Otis, H. Rahn, and M. A. Epstein. Performance as related to composition of alveolar air. Am. I. Physiol., 146:207-21. With A. B. Otis and H. Rahn. Venous pressure changes associated with positive intra-pulmonary pressures: their relationship to the distensibility of the lung. Am. J. Physiol., 146:307-17. With H. Rahn and A. B. Otis. A theoretical study of the composition of the alveolar air at altitude. Am. I. Physiol., 146:637-53. With H. Rahn, A. B. Otis, M. Hodge, M. A. Epstein, and S. W. Hunter. The effects of hypocapnia on performance. l. Aviat. Med., 17: 164-72. With H. Rahn, I. Mohney, and A. B. Otis. A method for the con- tinuous analysis of alveolar air. i. Aviat. Med., 17: 173-78. With A. B. Otis, H. Rahn, M. Brontman, and L. I. Mullins. Ballisto- cardiographic study of changes in cardiac output due to respira- tion. J. Clin. Invest., 25:413-21. 1947 With A. B. Otis, H. Rahn, L. E. Chadwick, and A. H. Hegnauer. Displacement of blood from the lungs by pressure breathing. Am. J Physiol., 151 :258-69. With L. E. Chadwick. Effect of pressure breathing on blood flow through the fingers. Am. J. Physiol., 151 :270-75. With A. L. Barach, E. B. Ferris, and C. F. Schmidt. The physiology of pressure breathing: a brief review of its present status. J. Aviat. Med., 18: 73-87. With R. J. Dern. The effect of varying pulmonary pressure on the arterial pressures in men and anesthetized cats. J. Clin. Invest., 26:460-67. 1948 With A. B. Otis and H. Rahn. Alveolar gas changes during breath holding. Am. .T- Physiol., 152:674-86. Physiology of exposures to abnormal concentrations of the respira- tory gases. Proc. Am. Philos. Soc., 92: 145-54.

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166 BIOGRAPHICAL MEMOIRS 1949 Physiology on horseback. Am. J. Physiol., 159:~51-55. With R. Galambos, A. B. Otis, and H. Rahn. Corneoretinal potential in anoxia and acapnia. J. Appl. Physiol., 1:710-16. With H. Rahn and A. B. Otis. Daily variations of vital capacity, residual air, an expiratory reserve, including a study of the resid- ual air method. J. Appl. Physiol., 1: 725-36. With H. Rahn, A. B. Otis, and L. E. Chadwick. Voluntary pressure breathing at high altitudes. l. Appl. Physiol., 1:752-72. With H. Rahn, A. B. Otis, and L. E. Chadwick. Physiological obser- vations on hyperventilation at altitude with intermittent pressure breathing by the pneumolator. i. Appl. Physiol., 1: 773-89. With R. T. Clark, Jr., and J. N. Stannard. Evidence for the conver- sion of carbon monoxide to carbon dioxide by the intact animal. Science, 109:615-16. Potassium. Sci. Amer., 181 (No. 2:~16-21. 1950 With R. T. Clark, in and l. N. Stannard. The burning of CO to CO2 by isolated tissues as shown by the use of radioactive carbon. Am. J. Physiol., 161:40~6. With H. Rahn and A. B. Otis. Respiratory system. Annul Rev. Physiol., 7: 179-204. With A. B. Otis, W. O. Fenn, and H. Rahn. Mechanics of breathing in man. J. Appl. Physiol., 2:592-607. With R. Gerschman. The loss of potassium from frog nerves in anoxia and other conditions. l. Gen. Physiol., 33: 195-203. Department of physiology and vital economics. In: The Quarter Century 1925-50, pp. 53-60. Rochester, N.Y.: The University of Rochester. 1951 Mechanics of respiration. American journal of Medicine, 10:77-90. With R. Gerschman, G. Fischer, J. Lacy, M. Bailly, and J. L. Wright. Experiments on the role of potassium in the blocking of neuro- muscular transmission by curare and other drugs. J. Gen. Physiol., 34:~. Medical aspects of military manpower selection. In: The Selection of

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WALLACE OSGOOD FENN 167 Military Manpower, ed. L. Carmichael and L. C. Mead, pp. 28- 37, National Academy of Sciences Publ. No. 209. Wash., D.C.: National Academy of Sciences. Medical aspects of military manpower selection. Sci. Mon., 73: 209-12. With A. B. Otis and H. Rahn. Studies in respiratory physiology. Air Force Technical Report No. 6528. U.S. Air Force, Wright Air Development Center, August. lg52 With A. B. DuBois, R. C. Fowler, and A. Softer. Alveolar CO2 measured by expiration into the rapid infra-red gas analyzer. J. Appl. Physiol., 4:526-34. With A. B. DuBois and A. G. Britt. Alveolar CO2 during the respira- tory cycle. J. Appl. Physiol., 4:535~8. With A. B. DuBois and A. G. Britt. COP dissociation curve of lung tissue. J. Appl. Physiol., 5: 13-16. Cost of medical education. Bull. Monroe County Med. Soc. Roches- ter Acad. Med., 10:49-52. The great goldrush in medical research. University Tennessee Rec- ord, 55:78-89. lg53 With F. H. Freeman. Changes in carbon dioxide stores of rats due to atmosphere low in oxygen or high in carbon dioxide. Am. J. Physiol., 174:422-30. Acute and sustained high energy output. Symposium on stress, Army Medical Services Graduate School, Walter Reed Army Medical Center, Wash., D.C., March. lg54 With A. B. Otis and M. Suskind. The accumulation of carbon di- oxide in apneic dogs during intermittent oxygen insulation. Am. l. Physiol. Med., 33:299-312. With R. Gerschman. Ascorbic acid content of adrenal glands of rat in oxygen poisoning. Am. J. Physiol., 176:6-8. With R. Gerschman, D. L. Gilbert, S. W. Nye, and P. W. Nadig. Role of adrenalectomy and adrenal-cortical hormones in oxygen poisoning. Am. J. Physiol., 178:346-50.

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168 BIOGRAPHICAL MEMOIRS With R. Gerschman, D. L. Gilbert, S. W. Nye, and P. Dwyer. La intoxacacion par el oxigeno y par los rayos X: un mecanismo en comun. Ciencia e Investigacion, 10:346-50. With P. Dejours. Composition of alveolar air during breath holding with and without prior inhalation of oxygen and carbon dioxide. J. Appl. Physiol., 7: 313-19. With R. Gerschman, S. W. Nye, D. O. Gilbert, and P. Dwyer. Studies on oxygen poisoning; protective effect of beta-mercaptosthyla- mine. Proc. Soc. Exp. Biol. Med., 85:75-77. With J. Goodman. Effects of 3-(o-toloxy)-propane-1, 2-diol (mephene- sin) and 3-~2-methyl-6-chlorophenoxy)-propane-1, 2-diol (P105) on a-excitability of muscle. Proc. Soc. Exp. Biol. Med., 85:500-503. With R. Gerschman, D. Gilbert, and S. W. Nye. Influence of X-irra- diation on oxygen poisoning in mice. Proc. Soc. Exp. Biol. Med., 86:27-29. With R. Gerschman, D. Gilbert, S. W. Nye, and P. Dwyer. Oxygen poisoning and X-irradiation: a mechanism in common. Science, 1 1 9: 623-26. The pressure volume diagram of the breathing mechanism. In: Handbook of Respiratory Physiology, ed. W. M. Boothby, U.S. Air Force School of Aviation Medicine, Randolph Field, Texas, September. 1955 Recent advances in basic muscle chemistry, physiology and pharma- cology. Am. J. Physical Med., 34: 8-10. With D. L. Gilbert and R. Gerschman. Effects of fasting and X-irra- diation on oxygen poisoning in mice. Am. I. Physiol., 181:272-74. With H. Rahn. A graphical analysis of the respiratory gas exchange: the O2-CO2 diagram. Wash., D.C.: The American Physiological Society. With J. E. Drorbaugh. A barometric method for measuring ventila- tion in newborn infants. Pediatrics, 16:81-87. With R. Gerschman, D. L. Gilbert, S. W. Nye, and W. E. Price. Effects of autonomic drugs and of adrenal glands on oxygen poisoning. Proc. Soc. Biol. Med., 88:617-21. Una rassegna sui lavori esequiti di recense net campo delta respira-

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WALLACE OSGOOD FENN 169 zione, applicati alla respirazione a pressione volontaria. Rivista Di Medicina Aeronautica, 18: 3-22. With H. Rahn. Studies in respiratory physiology. Second series: Chemistry, mechanics and circulation of the lung. WADC Techn. Rep., 55-357, U.S. Air Force, Wright Development Center, November. 1956 With T. Velasquez. The effect of adaptation to CO atmospheres on the rate of burning of CO by frog muscle. Acta Physiol. Lat. Am., 6:23-26. With T. Asano. Effects of carbon dioxide inhalation on potassium liberation from the liver. Am. l. Physiol., 185:567-76. With B. S. Olsen. Effect of CO2 on blood lactic acid in cats. Proc. Sac. Exp. Biol. Med., 91:477-79. 1957 The mechanics of standing on the toes. Am. I. Physical Med., 36: 153-56. Some elasticity problems in the human body. In: Tissue Elasticity, ed. l. W. Remington, pp. 98-101. Wash., D.C.: American Physio- logical Society. Changes in length of blood vessels on inflation. In: Tissue Elasticity, ed. i. W. Remington, pp. 154-167. Wash., D.C.: American Physio- logical Society. Scientific manpower in Russia. Bull. Am. Inst. Biol. Sc.: 7:~22. With D. L. Gilbert. Calcium equilibrium in muscle. l. Gen. Physiol., 40:393~08. With D. F. Sears. Narcosis and emulsion reversal by inert gases. J. Gen. Physiol., 40:515-20. With R. Gerschman, D. L. Gilbert, D. E. Terwilliger, and F. V. Cothran. Mutagenic effects of high oxygen tensions on Esche- richia coli. Proc. Natl. Acad. Sci. USA, 43:1027-32. Sodium and potassium contents of frog muscle after extraction in 50% glycerol. Proc. Soc. Exp. Biol. Med., 96:783-85. Ionic transfer in muscle and nerve. In: Metabolic aspects of Trans- port across Cell Membranes. ed. Q. R. Murphy, pp. 151-58. Madison: Univ. of Wisconsin Press.

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170 BIOGRAPHICAL MEMOIRS 1958 With T. A. Rogers and E. A. Ohr. Muscle electrolytes in acid and alkaline solutions. Am. l. Physiol., 194:373-78. The challenge of space biology. Bull. Am. Inst. Biol. Sci., 8: 15. Concepts and problems concerning ion transport. Fed. Proc., 17:578. Remarks on acceptance of Gold Medal Award of the University of Rochester Medical Alumni. Rochester Alumni News-Views, 1: 2-3. 1959 Introduction, Symposium on Life in Space. Fed. Proc., 18: 1241. With E. Agostoni and F. F. Thimm. Comparative features of the mechanics of breathing. J. Appl. Physiol., 14:679-83. With H. Falsetti. Effect of oxygen tension on sodium transport across isolated frog skin. Proc. Soc. Exp. Biol. Med., 101:721-22. 1960 With l. G. Henrotte and E. Cosmos. Calcium exchange in isolated turtle ventricle. Am. l. Physiol., 199: 779-82. Front seats for biologists past president's address. Bull. Am. Inst. Biol. Sci., 10:13-18. With J. H. Knowles and W. Newman. Determination of oxygenated, mixed venous blood CO2 tension by a breath-holding method. J. Appl. Physiol., 15:225-28. With E. Agostoni. Velocity of muscle shortening as a limiting factor in respiratory air flow. J. Appl. Physiol., 15:349-53. Physiology in orbit. The Physiologist, 3:20-26. The philosophy of research. Rochester Med. Alumni News, 2: 12-14. Mechanism of breathing. Sci. Am., 202: 138-148. 1961 With A. B. Craig, Jr. and Eleanore A. Ohr. Factors affecting the sodium and potassium contents of glycerinated frog muscle. Am. I. Physiol., 200:561-64. Carbon dioxide and intracellular homeostatis. Ann. N. Y. Acad. Sci., 92:547-58. Biological communications, theoretical and philosophic aspects. Biol. Abstr., 36:17-23.

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WALLACE OSGOOD FENN 171 American Institute of Biological Sciences. The Physiologist, 4:57-60. Creativity in medicine. Rochester Alumni Review, ian.-Feb., p. 6. Physiologist, pharmacologist, physician. A tribute to Dr. Wedd. Univ. of Rochester Med. Alumni News, 6-7. John Raymond Murlin. Yearb. Am. Philos. Soc., pp. 145-52. 1962 Born fifty years too soon. Annul Rev. Physiol., 24: 1-10. Physiological effects of high pressures of nitrogen and oxygen. Circu- lation, 16: 1134~3. Comments on a paper by Dr. Whalen. Fed. Proc., 21 :999-1000. A comparison of respiratory and skeletal muscles. In: Perspectives in Biology, ed. C. F. Cori, V. G. Foglia, L. F. Leloir, and S. Ochoa, pp. 293-300. Amsterdam, New York, & London: Elsevier. 1963 Introductory remarks. Ann. N. Y. Acad. Sci., 109:415-17. La regolazione della Accademia Medica Lombarda, 18: 1-12. Man's survival in space. Bull. Monroe County Med. Soc. Rochester Acad. Med., 21:52-55. L'ossigeno e i suoi compiti in fisiologia. Bullettino delle Societa Italiana di Biologia Sperimentale, 39: 1703-13. With A. B. Craig, in Effect of CO2 on respiration using a new method of administering CO2. J. Appl. Physiol., 1023-24. International Union of Physiological Sciences. The Physiologist, 6:44-46. History of the American Physiological Society: The Third Quarter Century' 1937-1962. Wash., D.C.: American Physiological Society. 1964 Report on meeting, Minimum Ecological Systems for Man. Bio- Science, 14:32-33. With H. Rahn, eds. Handbook of Physiology, Respiration, vols. 1 and 2. Wash., D.C.: American Physiological Society. Introduction to the mechanics of breathing. In: Handbook of Phys- iology, Respiration, ed. W. O. Fenn and H. Rahn, vol. 1, pp. 357-62. Wash., D.C.: American Physiological Society. Introduction. In: Oxygen in the ~nimal Organism, ed. F. Dickens and E. Neil. Oxford: Pergamon Press.

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172 BIOGRAPHICAL MEMOIRS 1965 Inert gas narcosis. Ann. N.Y. Acad. Sci., 117:760-67. Some physiological differences between air and low pressure oxygen atmospheres. Astronautica Acta, 11:133~10. Alexander Forbes (1882-1964~. Yearb. Am. Philos. Soc., pp. 140~5. 1966 With I. l. Thomas and R. C. Baxter. Interactions of oxygen at high pressure and radiation in drosophila. l. Gen. Physiol., 49:537~9. 1967 With M. Philpott, C. Meehan, and M. Henning. Recovery from oxygen poisoning in drosophila. Am. J. Physiol., 213:663-70. Gaseous exchange in breathing (pulmonary gas exchange). In: Ency- clopedia of Biochemistry, ed. R. J. Williams and E. M. Lansford, pp. 352-54. N. Y.: Reinhold. With M. Henning and M. Philpott. Oxygen poisoning in drosophila. I. Gen. Physiol., 50: 1693-1707. Inert gases. In: Physiology in the Space Environment, Respiration, vol. 2, pp. 102-12, National Academy of Sciences Publ. No. 1485B. Wash., D.C.: National Academy of Sciences. Interactions of oxygen and inert gases in drosophila. Resp. Physiol., 3:117-29. Possible role of hydrostatic pressure in diving. In: Proceedings of the Third Symposium on Underwater Physiology, ed. C. l. Lambert- sen, pp. 395~03. Baltimore: Williams and Wilkins. 1968 Perspectives in phonation. Ann. N. Y. Acad. Sci., 155:~8. With R. E. Marquis. Growth of Streptococcus faecalis under high hydrostatic pressure and high partial pressures of inert gases. J. Gen. Physiol., 52:810-24. Editor, History of the International Congresses of Physiology, 1889- 1968. Wash., D.C.: American Physiological Society. Introduction to the problems posed to the conference. In: Human Ecology in Space Flight. ed. D. H. Calloway, p. 11. N. Y.: New York Academy of Sciences.

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WALLACE OSGOOD FENN 173 1969 With R. E. Marquis. Dilatometric study of streptococcal growth and metabolism. Canadian Journal of Microbiology, 15: 993~0. Alexander Forbes. In: Biographical Memoirs, 40:113~1, Wash., D.C.: National Academy of Sciences. Oxygen poisoning and inert gas narcosis in paramecium caudatum. Physiol. Zoo., 42:129-137. With S. H. Rodgers and A. B. Craig, Jr. The oxygen consumption of rat tissues in the presence of nitrogen, helium or hydrogen. Resp. Physiol., 6: 168-77. With V. Boschen. Oxygen consumption of frog tissues under high hydrostatic pressure. Resp. Physiol., 7:335~0. The physiological effects of hydrostatic pressures. In: The Physiology of Diving and Compressed A ir Work, ed. P. B. Bennett and D. H. Elliott, pp. 36-57. Baltimore: Williams and Wilkins. 1970 The burning of CO in tissues. Ann. N. Y. Acad. Sci., 174:6~71. Life under high pressures. Proc. Am. Philos. Soc., 114: 191-97. A study of aquatic life from the laboratory of Paul Bert: a review of "La Vie Dans Les Eaux" by Paul Regnard, Paris 1891. Resp. Physiol., 9:95-107. 1971 With R. E. Marquis and W; P. Brown. Pressure sensitivity of strepto- coccal growth in relation to catabolism. l. Bacterial., 105:50~11. With V. P. Boschen. Hemolysis under high hydrostatic pressure. Proc. Soc. Exp. Biol. Med., 137:847-51. Partial molar volumes of oxygen and carbon monoxide Resp. Physiol., 13: 129~0. 1972 in blood. Partial pressure of gases dissolved at great depth. Science, 176: 1011-12. 1974 CO2 and the sea. In: CON and Metabolic Regulations, ed. K. E. Schaefer and G. Nahas, pp. 19-25. New York: Springer-Verlag.