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RALPH WALDO GERARD October 7, 900-February 17, 1974 BY SEYMOUR S. KETY DWRING THE MIDDLE DECADES of the twentieth century, study of the nervous system became a major component of biological research, growing from a strong base in mor- phology and physiology to involve all of the biological and behavioral sciences. It was not by chance that this clevelop- ment coincided with the time anal span of Ralph Gerard's scientific career, for he was one of a small number of intellec- tual leaders who brought it about. Born in Harvey, Illinois at the beginning of this century, Ralph Gerard was blessed with an uncommon intellectual enclowment, a heritage that has traditionally held scholarship and ethics in high regard, and a remarkable father who nur- tured his scientific curiosity. His father, Maurice Gerard, who hacI come to America from Central Europe after receiving a degree in engineering in Britain, was a self-employecl con- sultant to industry. He named his son after Emerson, whom he admired, and saw for him the career in pure science that he had been unable to pursue. From his father, Ralph Gerard also gained an appreciation of mathematics and of chess, showing particular aptitude for the latter, so that in his teens he beat the American champion and the world cham- pion at different times when they were playing simultaneous matches in Chicago. 179

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180 BIOGRAPHICAL MEMOIRS He completed the four-year course at Chicago's Hyde Park High School in two years, by passing examinations in subjects he already knew or hacT taught himself outside of class. He entered the University of Chicago at the age of fifteen, where he took at least one course in every one of the sciences offerec} and in most of the other disciplines besides. In this way, the natural genius and irrepressible interest that had been stimulated and reinforced by his father was broadened, uncloubtecTly contributing to his comprehension of many fields as a scientist and his unique ability to compound and integrate that knowledge with ever-widening scope. He was strongly influenced by Julius Stieglitz in chemistry, and by Anton CarIson and Ralph Lillie in physiology and neuro- physiology. He received his doctorate in physiology in 1921. Shortly thereafter, he married Margaret Wilson, who had just completect her doctorate in neuroanatomy, and together they finished their meclical training at the Rush Medical Col- lege in 1925. Margaret Wilson Gerard went on to train in pediatrics and psychiatry and to become an outstanding scholar and practitioner of child psychiatry until her death in 1954. Ralph Gerard took an internship at the L`os Angeles Gen- eral Hospital, at the end of which he was faced with what he felt was the major career decision of his life. He was offered a much-coveted residency in medicine at the same time that he was awardecI a National Research Council fellowship in neurophysiology and neurochemistry. He accepted the research fellowship. In an interesting revelation of his restive nature and lack of complacency, he recalled that decision at times with some misgiving, even after achieving worIdwicle acclaim as a neurophysiologist. The research fellowship launched him most propitiously

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RALPH WALDO GERARD 181 on his career in neuroscience. With A. V. Hill and Otto Meyerhoff, two giants in biophysics and biochemistry, re- spectively, he carrier! out pioneering research leading to the recognition that the concluction of the nerve impulse cle- pencled on biochemical processes along the nerve. When he returned to the University of Chicago in 1928, he was faced with another major decision. An offer by Carison of an ap- pointment in physiology was more than matched by Dallas Phemister, who was in the process of establishing the Depart- ment of Surgery in the new Medical School there. Again he chose physiology and remained in that department for twenty-five years. His laboratory there, which encompassed all of the promising neurobiological clisciplines, trained a large number of graduate students and postdoctoral fellows, several of whom were later to become leaclers in neuro- sclence. in 1952 Gerard was asked to develop and direct the re- search laboratories of the Neuropsychiatric institute of the University of TIlinois, and he spent the next two years orga- nizing a multidisciplinary research program that brought neurology and psychiatry more closely together, as well as the fundamental disciplines that sustain them both. After the death of Dr. Margaret Gerard, he accepted an invitation from Ralph Tyler to join the first group of dis- tinguished fellows of the Center for Advanced Stucly in the Behavioral Sciences, which had been established adjacent to the campus of Stanford University. During the preceding sixteen years, while he was engaged in some of his most significant contributions to neurophysiol- ogy, he fount! time to contemplate and address philosophical and social problems that lay beyond neuroscience. He pub- lished "The Role of Pure Science" in 193S, "Organism, Soci- ety and Science" in 1940, "A Biological Basis for Ethics" and

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182 BIOGRAPHICAL MEMOIRS "Higher Levels of Integration" in 1942, "Extrapolation from the Biological to the Social" in 1945 and, in 1946, "The Bio- logical Basis of Imagination." At the Center in Palo Alto, Ralph Gerard's interest in the behavioral and social sciences expanded further in what must have been exciting interactions with Anato} Rapoport, Clyde Kluckhohn, Franz Alexancler, Paul Lazarsfeld, and Alex Ba- velas, from which emerged "Biological and Cultural Evolu- tion" in 1956 and, in 1957, "Problems in the Institutionaliza- tion of Higher Education." In January 1955 he married Leona Bachrach Chalkley, whom he had known since high school when they were cap- tains of opposing debating teams. "Frosty," as he preferred to call her, in addition to being a skilled debater, is an accom- plishecl writer and poet. In the course of the fellowship year at the Center, another salutary event occurred when James Miller invited Gerard to join him and Anato! Rapoport in founding the Mental Health Research Institute at the Uni- versity of Michigan, and Ralph Gerard spent many hours discussing with Anato! Rapoport the possibility of creating a new and broader consortium of sciences dedicated to the stucly of behavior. Convinced of the sincerity and commit- ment of the Commonwealth and University of Michigan to this important concept, he accepted this opportunity and spent more hours with Miller, planning the philosophy and scope of the new research institute. Ralph and Frosty spent the next nine years at Ann Arbor, where he was professor of neurophysiology and director of the Institute's laboratories. During that time the Mental Health Research Institute grew from imaginatively conceived plans to one of the outstanding behavioral and psychiatric research centers in the nation, with a scope that embraces! fundamental neurochemical and physiological research, the behavioral sciences, and information processing. He clevotecT

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RALPH WALDO GERARD 183 the bulk of his research efforts during that time to organizing and leading a multifaceted stucly of schizophrenia, the most important of the mental illnesses in terms of loss of fulfi~- ment to the inctividual and family and cost to society. At the age of sixty-four, when some might have thought of retiring, Ralph Gerard instead accepted a new and broa(ler challenge, moving to the new Irvine campus of the University of California as professor of biological sciences and clean of the Graduate Division. This enabled him to revive the imaginative interest in teaching and eclucational philosophy that earlier hac3 made him one of the most stimu- lating contributors to the new concepts of undergraduate education introcluced by Robert Hutchins shortly after Ralph Gerard's appointment to the University of Chicago. At Irvine he continued his nurture of neuroscience, participating in the establishment of the important Department of Psycho- biology. Most of his energies and wisclom, however, were cievotecT to the wicler fields of education, science, society, and the social aspects of medicine. At the age of seventy he retired to throw himself filthy into civic affairs. At the same time his wife and his colleagues became concerned over some changes in his personality and some slowing of his intellectual functions. An intracerebral tumor was discoverecI, but even for this ominous situation, his remarkable brain found a salutary resolution. The tumor turned out to be a benign meningioma, which was remover! successfully and with complete recovery. Frosty has related a remarkable incident at that time that describes his indomita- ble spirit: "Service for a year on the Orange County Grand Jury meant so much to him that two days following the neu- rosurgical operation, and while he still remained in the hos- pital's intensive care unit, he insisted on dictating to me a letter to the Grand Jurors. He wrote them about his feelings when tests hac! revealed the existence of a massive tumor. If

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184 BIOGRAPHICAL MEMOIRS further tests had suggested that the mass was malignant, he would have refused to undergo surgery anct merely awaited the end. Instead, a benign tumor was inclicatect, he had taken his chances, and won. He wanted them to know that he was eager to rejoin them and to carry his share of the loacl. Two months later he was among them." For the next three-and- one-half years Ralph Gerard remained active; he ctied of coronary insufficiency in 1974. Ralph Gerard was elected to the National Academy of Sciences and to the American Academy of Arts and Sciences in 1955. He was the recipient of numerous honors and awards. Honorary degrees were bestowed upon him by the Universities of Maryland, Leiden, and St. Andrews, and by Brown University and McGill University. He was awarded the Medal of Charles University and the Orcler of the White Lion in Prague in 1946, the Stanley Dean Award in 1964, the Alumni Medal of the University of Chicago in 1967, and the Extraordinarius Awarcl of the University of California at Ir- vine, posthumously. He was a Distinguished Fellow of the American Psychiatric Association ant! an Honorary Fellow of the Royal Society of Edinburgh. A long list of honorary lec- tureships in this country and abroad attests to his interna- tional esteem and his brilliance as a speaker. He was a consul- tant to numerous research arms of the federal government, including the Office of Naval Research, the National Institute of Mental Health, and the National Science Foundation, and an advisor to numerous private founclations. Ralph Gerarc] was so extraordinary a man that he became a legend cluring his lifetime. His intellectual power was his outstanding characteristic, expressed early on by scholastic precocity, in micl-career by creative insights and the careful execution of crucial experiments, and at the end of his career in encyclopeclic erudition ant! wisdom. His knowlecige of the scientific literature, perceptiveness, and ability to synthesize

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RALPH WALDO GERARD 185 observations in a great variety of clisciplines, coupled with an almost poetic fluency in articulating and crystallizing issues, macle him highly regarcled as a teacher and as a summarizer of scientific conferences, in which he was undoubtedly one of the worId's leaders for several decades. He also possessed a remarkable sense of humor and a comparable fund of anec- dotes, which were always to the point. Among his physical characteristics, more striking than his portly figure and bawl pate, were his eyes, which have been described as "bright and restless the visible edge of a keen and probing mine! . . . eyes that showed by their sparkle not only the excitement of ctis- covery, but also a reflection of profound awe before the intri- cacy and complexity of the natural order."* These attributes will remain in the memories of his students and colleagues and all who knew him, but his most enduring legacy will be in his contributions to science, and particularly to neuro- science. During the twenty-five years that he devoted to lab- oratory research, he was responsible for a remarkable num- ber of pioneering insights and discoveries that opened up areas of neurobiological research that are far from exhausted today. Ralph Gerard aptly clescribed the motivation anti signifi- cance of his scientific career as a commitment to "the minute experiment and the large picture.": His contributions ful- fi~lect that commitment generously, for they demonstrate his remarkable ability to design and conduct rigorous research that crucially examines a specific hypothesis. They also epito- mize his vision, imagination, and courage to perceive the implications of the experimental results to the broact picture * The Reverend Edward P. Allen, remarks on the occasion of a memorial convo- cation, University of California at Irvine, 7 March 1974. ~ R. W. Gerard, "The Minute Experiment and the Large Picture," in The Neuro- sciences: Paths of Discovery, ed. F. G. Worded,. Swazey, and G. Adelman (Cambridge, Mass.: MIT Press, 1975), pp. 456-74.

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186 BIOGRAPHICAL MEMOIRS that would eventually emerge. He was both an architect of neuroscience ant! a stone carver. He attributed his enduring interest in the nervous system to a brief encounter with Anton CarIson, his professor of ohvsiolo~Y. while a student at the UniversitY of Chicago. 1 J (JJ ' J Gerard successfully defended his unwillingness, on logical grounds, to draw the accepted conclusion from a laboratory experiment that had for years been user! to demonstrate the nonfatigability of nerve. CarIson appreciated the wisdom in what a lesser man might have seen as brashness, took a con- tinuing interest in young Gerard, anal, several years later, recommender! him for the National Research Fellowship he was awarded in 1925. That fellowship permitted him to participate in A. V. Hill's classical demonstration of heat production by nerve and to make his first major discovery in the delayed heat production that follows a period of stimulation. Gerard de- scribec! those observations at Hill's suggestion at the Interna- tional Physiological Congress in Stockholm in 1926 and in his paper on "The Two Phases of Heat Production of Nerve" in 1927. He tract found that of the total quantity of heat attribut- able to a period of stimulation, only ~ ~ percent was releaser! during the stimulation, the much larger moiety being liber- ated over a period as long as ten minutes immediately follow- ing the stimulation. Although the heat generated in muscular contraction had been clemonstratect and measured for a long time, the much smaller amounts associated with nerve conduction had re- mained elusive. Hill, thirty-three years after this successful demonstration, recounted his many previous unsuccessful attempts and those of others going back to Helmho~z's first attempt in 1848, explaining its importance: Why did people go on trying to measure the heat production of nerve, in spite of repeated failure? Chiefly, ~ suppose, in order to settle the

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RALPH WALDO GERARD 187 question of whether the nerve impulse is the sort of physical wave in which the whole of the energy for transmission is impressed on the system at the start.... If it could be shown that heat really was produced all along the nerve during transmission, then the purely physical theory of conduction would be untenable. A distributed relay system would be required, with energy derived presumably from chemical change.* During the second year of his fellowship, Gerard moved to the laboratory of Otto Meyerhoff in Berlin in order to examine some of the chemical processes involves! in axonal transmission ant! the differences he surmised wouIc! exist cluring stimulation and recovery. With the use of specially prepared chambers of small size, he was able to measure the oxygen consumer! and the carbon dioxide released by a seg- ment of nerve at rest and cluring stimulation. He founc! that whereas the resting oxygen consumption of nerve and mus- cle were equal, the increase during stimulation in muscle was 8,000 times greater than that achieved in nerve. In addition, he measured the temperature coefficient of the oxygen me- tabolism in nerve arid its respiratory quotient at rest ant! cluring stimulation, and found evidence for the development of an oxygen debt in nerve cluring anoxic stimulation. The increased oxygen consumption of stimulated nerve was soon challenged as an artifact resulting from unphysio- logical stimulation rather than the physiological activity that resulted. F. O. Schmitt was able to counter that criticism by demonstrating that the oxygen consumption was correlatecI with the number of transmitted impulses rather than the amount or intensity of the stimulation. Then, in the summer of 1933, Gerard and H. K. Hartline established that physio- logical transmission alone accounted for the increased oxy- gen consumption: Hartline and I agreed to test this out on the Limulus optic nerve, ~ A. V. Hill, `'The Heat Production of Muscle and Nerve, 1848- 1914," Annual Review of Physiology, 2 1 ( 1959): 1 - 18.

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188 BIOGRAPHICAL MEMOIRS isolated along with the attached eye. The first attempt, using small War- burg vessels, was clearly far below the required sensitivity; but the problem was solved that same night by threading the optic nerve into a capillary through a Vaseline seal, the eye being outside and the far end being closed with a measuring drop. Two such capillaries in a large closed test tube in a thermostat were arranged so that light could be shined on the eye of either nerve, and each one thus constituted a control for the other. The movement of the index drop was followed with an ocular micrometer minute by minute. The oxygen consumption when "natural" nerve im- pulses were carried was established, and a valuable microrespirometer became available. Since our time commitments were such that we had less than a week to work together, experiments were continued day and night and neither of us was out of his clothes for the entire period.* With H. M. Serota he looker! for a similar coupling of metabolism to functional activity within the mammalian brain, where, unfortunately, the elegant technique he had used on the optic nerve was inapplicable. Using temperature, the only approach available to them, but which they could measure accurately, they inserted five thermocouples into particular structures by means of a stereotaxic instrument. They recognized that a change in temperature accompany- ing functional activity at a point within the brain could be the result either of altered metabolism or altered perfusion. They also reasoned that where the temperature of the blood and brain was the same, a sudden increase in temperature was likely to indicate the liberation of metabolic heat. Record- ing temperature changes and electrical activity, they were able to demonstrate an increase in both in the optic raclia- tions, the lateral geniculate, ant] the visual cortex upon illumination of the eye. It was not until forty years later that Louis Sokoloff succeeded in conclusively demonstrating the highly localized increased metabolism that accompanies func- tional activity in the visual system. * R. W. Gerard, "The Minute Experiment."

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RALPH WALDO GERARD 1934 201 With O. M. Solandt, D. Y. Solandt, and E. Ross. Methemoglobin and methylene blue as cyanide antagonists. Proc. Soc. Exp. Biol. Med., 31: 539-41. With H. K. Hartline. Respiration due to natural nerve impulses. A method for measuring. I. Cell. Comp. Physiol., 4:141-60. With B. B. Rubinstein. A note on the respiration of arbacia eggs. J. Gen. Physiol., 17:375-81. With A. V. Hill, W. O. Fenn, and H. S. Gasser. Physical and chem- ical changes in nerve during activity. The chemical activity of nerve. Science, 79 (Suppl.~:20-26. With B. B. Rubinstein. Fertilization and the temperature coef- ficients of oxygen consumption in eggs of arbacia punctulata Gen. Physiol., 17:677-85. The irritability of a non-medullated nerve. I. Physiol., 83:24-25. 1935 .~. With McK. Cattell. The inhibitory effect of high-frequency stimula- tion and the excitation state of nerve. I. Physiol., 83:407-15. With M. Shaffer and T. H. Chang. The influence of blood con- stituents on oxygen consumption in nerve. Am. J. Physiol., 111:697-709. With T. H. Chang and M. Shaffer. The influence of electrolytes on respiration in nerve. Am. I. Physiol., 111:681-96. 1936 With G. L. Engel. The phosphorus metabolism of invertebrate nerve. I. Biol. Chem., 112:379-92. With H. Seroia. Localized thermal changes in brain. Am. l. Physiol., 116:59. With N. O. Brookens and L. Ectors. Respiration of local brain regions. Technique and applications. Am. J. Physiol., 116:16-17. With H. W. Magoun. Influence of potassium and calcium on motor discharges. Proc. Soc. Exp. Biol. Med., 34:755-56. With M. S. Kharasch, R. R. Legault, and A. B. Wilder. Metal cata- lysts in biologic oxidations. I. The simple system. I. Biol. Chem., 113:537-55. With M. S. Kharasch, R. R. Legault, and A. B. Wilder. Metal cata-

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202 BIOGRAPHICAL MEMOIRS lysts in biologic oxidations. II. Tissue inhibitors. J. Biol. Chem., 113:557-69. With W. H. Marshall and L. I. Saul. Electrical activity of the cat's brain. Arch. Neurol. Psychiatry, 36:675-735. With F. Offner. A high speed crystal ink writer. Science, 84: 20~10. Metabolism and excitation. Cold Spring Harbor Symp. Quant. Biol., 4:194-201. Factors controlling brain potentials. Cold Spring Harbor Symp. Quant. Biol., 4:292-304. Factors influencing brain potentials. Trans. Am. Neurol. Assoc., 62:55-60. 1937 The metabolism of brain and nerve. Annul Rev. Biochem., 6:419-44. With M. B. Cohen. Oxidizing enzymes in brain extracts. Am. I. Physiol., 119: 34-47. With H. Blake. Brain potentials during sleep. Am. I. Physiol., 119:692-703. With J. Z. Young. Electrical activity of the central nervous system of the frog. Proc. R. Soc. London, 122B:343-52. With L. Schoen. The role of dicarboxylic acids in brain oxidations. Am. J. Physiol., 119:397 - 98. With N. Tupikova. Salt content of neural structures. Am. J. Phys- iol., 119:414-15. With R. A. Cohen. Hyperthyroidism and brain oxidations. I. Cell. Comp. Physiol., 10:223 -40. With H. M. Serota. Localized thermal changes in the cat's brain. I. Neurophysiol., 1: 115-24. Brain metabolism and circulation. Proceedings, Association for Re- search in Nervous Mental Disease, 18:316-45. 1938 With B. Libet. Automaticity of central neurones after nicotine block of synapses. Proc. Soc. Exp. Biol. Med., 38:886-88. With L. E. Ectors and N. L. Brookens. Autonomic and motor lo- calization in the hypothalamus. Arch. Neurol. Psychiatry, 39:789-98. The role of pure science. Science, 88:361-68.

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RALPH WALDO GERARD 203 Anoxia and neural metabolism. Arch. Neurol. Psychiatry, 40: 985-96. With N. Tupikova. Creatine in nerve, muscle and brain. I. Cell. Comp. Physiol., 12:325-60. With O. Sugar. Anoxia and brain potentials. I. Neurophysiol., 1 :558-72. 1939 With N. Tupikova. Nerve and muscle phosphates. I. Cell. Comp. Physiol., 13: 1 - 12. With H. Blake and N. Kleitman. Factors influencing brain poten- tials during sleep. J. Neurophysiol., 2:48-60. With B. Libet. Control of the potential rhythm of the isolated frog brain. J. Neurophysiol., 2:153-69. With H. H. Dubner. Factors controlling brain potentials in the cat. J. Neurophysiol., 2: 142-52. With E. Tokaji. Avitaminosis B. and pigeon brain potentials. Proc. Soc. Exp. Biol. Med., 41 :653 - 58. Mechanism of action of "shock therapies." Arch. Neurol. Psychia- try, 42:564-65. 1940 With O. Sugar. Spinal cord regeneration in the rat. J. Neurophys- iol., 3:1-19. With B. Libet. The control of normal and "convulsive" brain potentials. Am. I. Psychiatry, 96: 1125-51. Organism, society and science. Sci. Mon., 50:340-50, 403- 12, 530-35. 1941 With F. Panimon and M. K. Horwitt. Orthophenanthroline as accelerator of brain tissue oxygen consumption. I. Cell. Comp. Physiol., 17:17-29. With F. Panimon and M. K. Horwitt. Iron induced oxidations in brain and other tissues. I. Cell. Comp. Physiol., 17:1-16. With B. Libet. Steady potential fields and neurone activity. J. Neu- rophysiol., 4:438-55. The interaction of neurones. Ohio J. Sci., 41:160-72. Intercellular electric fields and brain function. Schweiz. Med. Wo- chenschr., 12:555 -59.

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204 BIOGRAPHICAL MEMOIRS With J. Tobias. An improved capillary microrespirometer. Proc. Soc. Exp. Biol. Med., 47:531-33. Science at the celebration. Univ. Chicago Mag., 34:12-14. 1942 With }. Pearce. The respiration of neurones. Am. }. Physiol., 136: 49-65. A biological basis for ethics. Philos. Sci., 9:92-120. Higher levels of integration. Science, 95: 309- 13. Electrophysiology. Annul Rev. Physiol., 4:329-58. 1944 With F. P. Simon and M. K. Horwitt. The inhibition of catalyzed oxidations by hemins. }. Biol. Chem., 154:421-25. 1945 With A. E. Emerson. Extrapolation from the biological to the social. Science, 101:582 - 85. 1946 The biological basis of imagination. Sci. Mon., 62:477-99. With J. M. Tobias, C. C. Lushbaugh, H. M. Patt, S. Postel, and M. N. Swift. The pathology and therapy with 2, 3-dimercapto- propanol (BAL) of experimental cadmium poisoning. ]. Pharma- col. Exp. Ther. (Suppl.), 87:102-18. With H. M. Patt, }. M. Tobias, M. N. Swift, and S. Postel. Hemody- namics in pulmonary irritant poisoning. Am. }. Physiol., 147: 329-39. With }. Graham. Membrane potentials and excitation of impaled single muscle fibers. I. Cell. Comp. Physiol., 28:99-117. Nerve metabolism and function. Ann. N.Y. Acad. Sci.,47:575-600. With S. Postel, J. M. Tobias, and H. M. Patt. The effect of exercise on mortality of animals poisoned with diphosgene. Proc. Soc. Exp. Biol. Med., 63:432-36. 1947 With F. P. Simon and A. M. Potts. Metabolism of isolated lung tissue: Normal and in phosgene poisoning. J. Biol. Chem., 167:303- 11.

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RALPH WALDO GERARD 205 With F. P. Simon and A. M. Potts. Action of cadmium and thiols on tissues and enzymes. Arch. Biochem., 12:283-91. With L. L. Boyarsky and I. M. Tobias. Nerve conduction after inactivation of choline esterase. Proc. Soc. Exp. Biol. Med., 64: 106-8. The scope of science. Sci. Mon., 64:496-512. With S. W. Kuffler. The small-nerve motor system to skeletal mus- cle. J. Neurophysiol., 10:383 - 94. Science and the public. Science, 106:23-25. Anesthetics and cell metabolism. Anesthesiology, 8:453-63. With R. D. Tschirgi. The carotid-mandibular reflex in acute respi- ratory failure. Am. J. Physiol., 105:358-64. 1949 With A. M. Potts and F. P. Simon. The mechanism of action of phosgene and diphosgene. Arch. Biochem., 24:329-37. Physiology and psychiatry. Am. J. Psychiatry, 106: 161 - 73. With V. B. Brooks and R. E. Ransmeier. Action of anticholines- terases, drugs and intermediates on respiration and electrical activity of the isolated frog brain. Am. I. Physiol., 157:299-316. With }. M. Tobias, S. Postel, H. M. Patt, C. C. Lushbaugh, and M. N. Swift. Localization of the site of action of a pulmonary irri- tant, diphosgene. Am. J. Physiol., 158:173-83. With G. Ling. The normal membrane potential of frog sartorius fibers. }. Cell. Comp. Physiol., 34:383-96. With G. Ling. The influence of stretch on the membrane potential of the striated muscle fiber. I. Cell. Comp. Physiol., 34:397- 405. With G. Ling. The membrane potential and metabolism of muscle fibers. I. Cell. Comp. Physiol., 34:413-38. 1950 With A. M. Potts, F. P. Simon, }. M. Tobias, S. Postel, M. N. Swift, and H. M. Patt. Distribution and fate of cadmium in the animal body. Arch. Ind. Hyg. Occup. Med., 2:175-88. A biologist's view of society. Common Cause, 3:630-38. With G. Ling. Effect of external potassium concentration on the membrane potential of single muscle fibers. Nature, 165: 113-14.

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206 BIOGRAPHICAL MEMOIRS Some aspects of neural growth, regeneration and function. In: Genetic Neurology, ed. P. Weiss, pp. 199-207. Chicago: Univ. of Chicago Press. With R. W. Doty. Nerve conduction without increased oxygen con- sumption: Action of azide and fluoracetate. Am. I. Physiol., 162:458-68. Some of the problems concerning digital notions in the central nervous system. In: Cybernetics, ed. H. von Foerster, pp. 1 1-57. New York: Josiah Macy fir. Foundation. 1951 The physiology of pain. Abnormal neurone states in causalgia and related phenomena. Anesthesiology, 12: 1 - 13. With A. L. Samuels, L. L. Boyarsky, B. Libet, and M. Brust. Distri- bution, exchange and migration of phosphate compounds in the nervous system. Am. I. Physiol., 164:1-15. With L. G. Abood. A hexokinase inhibitor in nerve. Proc. Soc. Exp. Biol. Med., 77:438-41. 1952 The organization of science. Annul Rev. Physiol., 14:1-12. With L. G. Abood, J. Banks, and R. D. Tschirgi. Substrate and enzyme distribution in cells and cell fractions of the nervous system. Am. I. Physiol., 168:728-38. With L. G. Abood. Oxidative esterification of phosphate by neural tissue. Am. I. Physiol., 168:739-41. With L. G. Abood and S. Ochs. Electrical stimulation of metabolism of homogenates and particulates. Am. l. Physiol., 171:134-39. With L. G. Abood and R. D. Tschirgi. Spatial and chemical.ex- change of phosphate in the resting and active nervous system. In: Phosphorus Metabolism, vol. 2, ed. W. D. McElroy and B. Glass, pp. 798-821. Baltimore: Johns Hopkins Press. 1953 Central excitation and inhibition. In: Cybernetics, ed. H. von Foerster, Mead, and Teuber, pp. 127-50. New York: Josiah Macy fir. Foundation. With H. P. Jenerick. Membrane potential and threshold of single muscle fibers. I. Cell. Comp. Physiol., 42:79-102.

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RALPH WALDO GERARD 207 With L. G. Abood. A new group of cytochrome oxidase inhibitors. }. Pharmacol. Exp. Ther., 108:261-73. 1954 With L. G. Abood. Enzyme distribution in isolated particulates of rat peripheral nerve. }. Cell. Comp. Physiol., 43:379-92. With G. Falk. Effect of micro-injected salts and ATP on the mem- brane potential and mechanical response of muscle. J. Cell. Comp. Physiol., 43:393 -403. With E. Streicher. Phosphate exchange in brain phospholipids in vivo and in vitro. Proc. Soc. Exp. Biol. Med., 85: 174-77. With R. E. Ransmeier. Effects of temperature, convulsion, and metabolic factors on rodent memory and EEG. Am. I. Physiol., 179:663-64. Experiments in microevolution. Science, 120:727-32. 1955 With A. F. Lash and G. Falk. A microelectrode method in the diagnosis of carcinoma of the cervix and endometrium. Am. I. Obstet. Gynecol., 70:354-58. With L. G. Abood. A phosphorylation defect in the brains of mice susceptible to audiogenic seizure. In: Biochemistry of the Develop- ing Nervous System, ed. H. Waelsch, pp. 467-72. New York: Academic Press. The academic lecture: The biological roots of psychiatry. Am. J. Psychiatry, 112:81 -90. - O 1 J I . With M. E. Kosman. The effect of adrenaline on a conditioned avoidance response. I. Comp. Physiol. Psychol., 48:506-8. With E. Sigg and S. Ochs. Effects of the medullary hormones on the somatic nervous system in the cat. Am. I. Physiol., 183:419-26. 1956 With K. Koketsu. Effect of sodium fluoride on nerve-muscle trans- mission. Am. J. Physiol., 186:278-82. With C. Kluckhohn and A. Rapoport. Biological and cultural evolu- tion. Behav. Sci., 1:6-34. 1957 The units and concepts of biology. Science, 125:429-33.

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