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Biographical Memoirs: Volume 53 (1982)

Chapter: Ralph Waldo Gerard

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Suggested Citation:"Ralph Waldo Gerard." National Academy of Sciences. 1982. Biographical Memoirs: Volume 53. Washington, DC: The National Academies Press. doi: 10.17226/576.
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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

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

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

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

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

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

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.

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

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.

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."

RALPH WALDO GERARD 189 In 1931 Gerarc! carried out an imaginative series of ex- periments with D. D. Cook on the phenomenon of axonal degeneration. They reasoned that a nerve degenerates beyond a cut either because that portion is no longer stimu- latecI, or because an important nutrient flow of chemical substances clown the fiber is stopped. They tested the first possibility by chronically stimulating a cut sciatic nerve with buried electrocles ant! observed that the nerve lost its func- tion even more rapidly when stimulated than when at rest. They concludecl: "Degeneration of a nerve process isolated from its cell bocly might be clue to lack of impulses conclucted by it or of necessary substances spreading along it. The evi- dence (here) considered favors the second possibility."* This was perhaps the first suggestion, supported by experimental evidence, for the important process of axonal flow, which Paul Weiss was able to demonstrate thirteen years later. In 1951, employing isotopically labelled phosphorous, Ralph Gerard and four collaborators made the first measurements of the flow of a chemical substance, phosphoprotein, clown nerve trunks, which occurred at a rate of 3 millimeters per day. In 1940, with Oscar Sugar, Gerard tackled the controver- sial subject of regeneration in the transected! mammalian spinal cord. Using immature animals, impeccable surgical techniques, and following a suggestion of Cajal by implanting pieces of peripheral nerve to serve as a scaffold on which the sprouts might climb, they providecl the first demonstration that functional as well as structural regeneration could take place. Because of the prevailing belief that regeneration was impossible in the mammalian spinal cord, the report receiver! scant attention. In summarizing a conference on the subject thirty years later, Gerard was able to take some satisfaction * R. W. Gerard and D. D. Cook, "The Effect of Stimulation on the Degeneration of a Severed Peripheral Nerve," American Journal of Physiology, 97 (1931):412-25.

190 BIOGRAPHICAL MEMOIRS from the new evidence presented, commenting that "Today the question is rather 'how,' not 'if."'* Expressing a note of optimism, he congratulated William Windie for having kept the spark alive. In 1933, 1934, and more completely in 1936, Gerard published, with Wade Marshall and Leon Saul, the results of research that opened a new chapter in neurophysiology ant! made possible the systematic mapping of the mammalian brain by Clinton Woolsey, and of the human brain by Wilder Penfield. Recognizing the power of recently developed tools—the oscilloscope, powerful amplifiers, and sterotaxic instruments for precise localization they proceeded to ex- plore the cat brain for spontaneous activity in its various regions. Using what they called "evoked potentials," they were able to trace the pathways by which particular sensory stimuli proceeded to the cortex and to follow their ramifica- tions and ripples into quite unexpected regions. With the cooperation of two neurosurgeons, it was possible to clemon- strate evoked cortical potentials at the operating table the first demonstration of what was to become a powerful clinical too} for the diagnosis and further unclerstanding of dis- turbed cerebral function. With Benjamin Libet in 1939 ant! ~ 94 I, Gerard publisher! the first experimental observations of steady potentials in the brain and their potential relationship to excitability patterns ant! the form of brain waves. They also demonstrated that spread of neuronal activity need not necessarily be mediated by the usual synaptic transmission. Caffeine-induced epilep- tiform waves were shown to travel across a complete transec- tion of the frog brain. It became apparent that extracellular fields of electric current flow could provide a significant mode of neuronal interaction and synchronization. Addi- * R. W. Gerard, "Summary of the Paraplegia Conference, Palm Beach, Florida, May 1-3, 1972," R. W. Gerard Collection, University of California at Irvine.

RALPH WALDO GERARD 191 tional studies on isolates! frog brain and fragments thereof showed that spontaneous rhythmicity in brain tissue could be a function of localized neuron groups anti their immediate ionic environments. As a result of these and his earlier obser- vations, Gerard developed the now generally accepted con- cept that the electroencephalogram represents the summa- tion of envelopes of slow potentials rather than neuronal spikes. Although these remarkable contributions were macle with the existing Acirian-Bronk concentric electrodes or with other macroelectrodes, Gerard was convinced that a true microelectrode could be developecI that might record the physiological activity of indiviclual neurons in the brain. When Judith Graham joined his laboratory as a graduate student a few years later, she began work on that goal by pulling fine glass capillaries. Gerard traced the idea behind this to what was probably his first research project, imagina- tively inspired and ingeniously executed while he was an undergracluate at Chicago. From George Bartelmez, the pro- fessor of histology, he learner! about myof~brils and the con- tinuing controversy over whether they were real or fixation artifacts. He suggested to the professor "that if a quartz needle was moved steadily across a living muscle fiber, the tip would move smoothly if the protoplasm was homogeneous but in a sort of cogwheel fashion if viscous fibrils were em- bedded in fluid sarcoplasm, and this could be followed by reflecting a beam of light from a mirror attached to the needle."* Bartelmez was enthusiastic and presented Gerard with the original micromanipulator that had been developec! in the department by Kite. "It was a museum piece but it still worked and T had a lot of fun learning micromanipulation and that protoplasm was a more complex thing than ~ * R. W. Gerard, "Informal Talk by an Old-timer" (lecture of 30 March 1972), R. W. Gerard Collection, University of California at Irvine.

192 BIOGRAPHICAL MEMOIRS thought." He did not solve that problem but the experience was invaluable later on to him and to neurophysiology. Judith Graham was able to straw capillaries down to a diameter of several micra, fill them with a conducting potas- sium chloride solution, insert them into inctiviclual muscle cells, and record the intracellular potential. In their publica- tion of the findings in 1946, Graham reported an average membrane potential of 62 mV but with a large range (41- 80 mV). They felt that the variation was due largely to the injury of insertion, since the finest electrocles gave smooth penetration uncler the microscope and also the highest read- ings. Gilbert Ling, who was also a graduate student in the labo- ratory at that time, began to work on the problem and after two years found that it was possible to make finer micro- pipettes, considerably less than one micron at their tip anct with a taper gentle enough that the tip would not break off. These could be inserted into a muscle cell without any inclica- tion of injury. Using these, Ling and Gerarc! were able to report in 1949 a membrane potential (78 + 5 mV) consist- ently at the high range that was previously obtained. Alan Hodgkin, attending a meeting of the American Physiological Society, was much impresser by these microelectrodes, came to observe their manufacture and application, and asker! per- mission to take one back to Cambridge. There he modified it by increasing the concentration of electrolyte and acIding a cathode follower, which made it capable of recording the rapid changes in potential that accompany action spikes in single cells. John Eccles applied the microelectrode to studies of activity of inclividual units within the spinal cord and brain ant] Andrew Huxley usect it in muscle cells. It would be difficult to exaggerate the important role that the capillary microelectrode has played in neurophysiology in the thirty years since its development. It made possible the

RALPH WALDO GERARD 193 neurophysiological research for which several Nobel Prizes have been awarded, and many of the most exciting advances that have occurrent in the past two decacles regarding neu- ronal activity in sleep, sensory processing, voluntary muscu- lar movements, and attention—to name just a few couict not have occurred at the time without it. Ralph Gerard indicated more than once that he did not consider the clevelopment of the microelectrocle to be his most important contribution. This may have been because of its technical rather than its conceptual nature. It was the quality of imaginative and prescient conceptualizations, such as those that unclerIay his research on memory, of which he was most proud. In his Gregory Lecture on "Physiology and Psychiatry" in 194S, Gerard proposed a concept of memory that was to become the basis of his experimental work in the f~elc! ant! that, today, remains the most plausible and heuristic model that we have: All is not over when an impulse flashes across a synapse and on to its destination. It leaves behind ripples in the state of the system. The fate of a later impulse can thus be at least a little influenced by the past history of the neurons involved.... Reverberant circuits, in principle, could last indefinitely, but in practice their duration is doubtful.... Perhaps there is a short-lasting active memory, depending on circuits, and a more enduring static one.* In a later series of ingenious experiments, it was possible to show that by interrupting or confounding the electrical activity of the brain by induced hibernation or electroshock, recently acquired memories would be erased while those that had been former! an hour or more before would persist. He surmised that in a brief critical period following a learning experience, the memory was transformed from a dynamic * R. W. Gerard, "Physiology and Psychiatry," American journal of Psychiatry, 106 (1949): 161-73.

194 BIOGRAPHICAL MEMOIRS representation in electrical activity through some chemical process to a more permanent molecular, physiological, or morphological state. In 1963, with T. J. Chamberlain and P. Halick, he tested this concept in an experiment that hacl all of the elegance and simplicity of CIaucle Bernard's (lemonstra- tion of the site of action of curare. They founct that the postural asymmetry procluced by a unilateral cerebelIar le- sion induced in less than one hour a persistent asymmetry in function at the level of the lower motor neuron. In further experiments with G. H. Rothschild, evidence was adduced that this process couch be facilitated by an agent that stimu- lated RNA synthesis and was Startled by drugs that inhibited that process. There have been thousands of experiments car- ried out since that time with more sophisticated physiological and biochemical techniques anct the general thrust of all of the research has been to support the idea that short-term memory is consoliciated in long-term memory by means of chemical processes, and that in that process RNA and protein synthesis may play essential roles. Gerard was also proud of his contributions to psychiatry in recognition of which he was made a distinguishect fellow of the American Psychiatric Association. His contributions here were not at the level of the minute experiment but of the large picture. Indeed, he played an important role in foster- ing the development in psychiatry of critical and judicious scientific approaches. in the Gregory Lecture, he com- mented on the continuing controversy regarding the genesis of psychoses: The constitutionalists and the organicists and the environmentalists and the mentalists too often are quarreling with each other as to which of them has the cause. Now, it is obviously useful to find out that schizo- phrenics have abnormal capillaries in their fingers, that they had abnormal experiences in their childhood, and that they have abnormal individuals as parents or sibs; but one does not exclude the others and no one of them

RALPH WALDO GERARD 195 can possibly be the whole story.... It is never sensible to ask the question "Does heredity or environment determine some characteristic?" It always takes both . . . and the meaningful question which is mostly not asked is a quantitative one.* In his Academic Lecture at a convocation of the American Psychiatric Association ~ ~ 955), he developed further the theme of etiologic and typologic diversity in mental illness: Many different etiologies may initiate the same train of pathogenic events; many different pathogenic sequences may produce a single pathol- ogy; and many different pathologies may still lead to a single symp- tom.... Can it be doubted that mental disease also presents symptoms and even syndromes which may subsume multiple nosologic entities and which are almost certainly based on multiple chains of abnormalities? ~ Gerard prepared that lecture just before moving to the Mental Health Research Institute at the University of Michi- gan. During the eight years he spent there, he organized and (Erected a large multidisciplinary program that was to ex- amine his concept of the heterogeneity of schizophrenia. On the basis of a small number of objective psychological, physio- logical, and biochemical measurements, it was found that seven typologies conic! be distinguishecl an cl characterized as well by clinical and behavioral observations. Ralph Gerard's recognition of the importance of both genetic and environmental components in mental illness and the heterogeneity of the classical syndromes is by this time well established ant! has become part of the mainstream of modern psychiatry, where it has led to a more open-minded and less doctrinaire examination of plausible hypotheses that are not mutually exclusive. it has been possible in the foregoing account to delineate *Ibid. tR. W. Gerard, "The Academic Lecture: The Biological Roots of Psychiatry," American fournal of Psychiatry, 1 12 ( 1955): 8 1-90.

196 BIOGRAPHICAL MEMOIRS some of the peaks of Ralph Gerarcl's scientific work. There were other peaks as well, and all rose from a high plateau of phenomenal productivity extending over fifty years and more than 500 publications. By far the major portion of his dedication was to neuroscience, which he insisted was a single cliscipline. In a deep sense, the appreciation ant! command of biophysics, biochemistry, physiology, psychology, anal psy- chiatry that his own career exemplified offered a clear valicla- tion of that thesis. He played an important role in establishing the Society for Neuroscience and at its foundling meeting in 1969 he was elected! honorary president by unanimous ac- cIaim. His large picture, however, was even larger than neuro- science. It embraced all of science and human imagination as well, which he once described as the culminating efflores- cence of the process of evolution up to the present time. He was a strong exponent of the implications and responsibility of science to society, but a courageous mentor as well of the reciprocal responsibility of society to scientific freedom and growth. In 1~952, in an important paper entities! "The Orga- nization of Science," he wrote: Tt heart: r~r`~titi^m that the im~r~Oc:~ in mr<rOni7~ti~n iC On in~Y^rOh]~ ^ ~ B_ ~ ~ A ~ ~ i_ ~ ~ ~ ~ ~ ~ A . . . ~ 6 1 ~ ~ . ~ %_ ~ ~ -_ A 1 4 ~ A 5 ~ ^—A 4. LO ~ J—— A ~7 ~—— A 4 4 ~~ trend in evolution; our problem is to fight the diseases and enhance the uses of interrelatedness. The great danger is authoritarianism and con- formity. This can blight at any level from the petty bookkeeping practices of too many governmental agencies to the national murder of the free pursuit of truth. The Lysenko story in Russia and the earlier distortions under Hitler deserve the most careful attention by scientists. Although these represent excesses under totalitarian police states, the anlages of similar attitudes are clearly present in our country.* Ten years earlier, he saw the relationship of science to society in a remarkable perspective which is even more perti- nent today: *Annual Review of Physiology, ~4 (~952):~-~2.

RALPH WALDO GERARD 197 These are the New Frontiers of mankind in the illimitable domain of the mind which science penetrates and scholarship consolidates. Change is often uncomfortable but it is exhilarating. Societies like animals must evolve or retrogress. Science, created by the social organism to sensitize itself to a fuller environment, is stirring and shaking the body politic with the birth pangs of the new. Men may suffer on the way probably a caterpillar does not metamorphose painlessly into a butterfly and the direction of travel is still obscure. But mankind is on the march somewhere, not vegetating into decadence. Science has brought and will bring men both weal and woe, mostly weal; and it is not destroying and will not destroy, rather it is enhancing those values of human society which we call civilization.* That understanding and appreciation of science and its salu- tary role in social evolution was Ralph Gerarcl's enduring credo. He cried on February 17, 1974, survived by his son James and his wife Leona Bachrach ("Frosty") Gerard. A dedicated, creative, and compassionate person in her own right, she had made the two clecacles they spent together a happy and mu- tually enriching experience. Many of his colleagues and former students have tried to put into words the unique qualities of Ralph Waldo Gerard. Perhaps Lord Adrian, who was his onetime mentor ant] long- time friend, stated it best: "There were few physiologists or philosophers with his unclerstanding both of experimental techniques anct of human aspirations."! ~ AM GREATLY INDEBTED to Leona Bachrach Gerard, who shared with me her personal reminiscences, biographical memorabilia, and letters; to Benjamin Libet and Richard Thompson for notes on Gerard's scientific contributions; to Roger B. Berry of the Irvine University Library for complete bibliographies; and to Roxanne- Louise Nilan, who compiled the Gerard Microfiche Collection, for access to unpublished documents. * R. W. Gerard, "Science at the Celebration," University of Chicago Magazine, 34 (1941): 12- 14. t Lord Adrian to Leona Gerard, 10 March 1974.

198 BIOGRAPHICAL MEMOIRS BIBLIOGRAPHY 1922 Chemical studies on intestinal intoxication. I. The presence and significance of histamine in an obstructed bowel. I. Biol. Chem., 52:111-24. The lethal agent in acute intestinal obstruction. I. Am. Med. Assoc. 79: 1581-84. 1924 With A. lit. Abel, R. W. Backus, and H. Bourquin. Tryptophane and thyroid function. Am. l. Physiol., 73:287-95. 1926 With T. Koppanyi. Studies on spinal cord regeneration in the rat. Am. J. Physiol., 76:211-12. With A. C. Downing and A. V. Hill. The heat production of nerve. Proc. R. Soc. London, 100B:223-51. 1927 The two phases of heat production of nerve. l. Physiol., 62:349-63. With A. V. Hill and Y. Zotterman. The effect of frequency of stimulation on the heat production of nerve. I. Physiol., 63: 130-43. Studies on nerve metabolism. I. The influence of oxygen lack on heat production and action current. I. Physiol., 63:289-98. Studies on nerve metabolism. II. Respiration in oxygen and nitro- gen. Am. I. Physiol., 82:381 -404. With O. Meyerhoff. Studies on nerve metabolism. III. Chemismus and intermediarprozess. Biochem. Z., 191:125-46. The activity of nerve. Science, 66:495-99. 1928 With A. Forbes. A note on action currents and "equilibration" in the cat's peroneal nerve. Am. J. Physiol., 86:178-85. With A. Forbes. "Fatigue" of the flexion reflex. Am. J. Physiol., 86: 186-205. 1929 With l. Wallen. Studies on nerve metabolism. V. Phosphates. Am. J. Physiol., 89: 108-20.

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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-

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.

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.

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.

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.

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.

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.

208 BIOGRAPHICAL MEMOIRS With D. G. Marquis, E. L. Kelly, J. G. Miller, and A. Rapoport. Experimental studies of behavioral effects of meprobamate on normal subjects. In: Meprobamate and Other Agents Used in Mental Disturbances, ed. Otto Whitelock. Ann. N.Y. Acad. Sci., 67: 701-10. Problems in the institutionalization of higher education; an analysis based on historical materials. Behav. Sci., 2: 134-46. 1958 With E. L. Kelly,J. G. Miller, D. G. Marquis, and L. Uhr. Continued meprobamate and proclorperazine administration. Arch. Neu- rol. Psychiatry, 80:247-52. Concepts of biology. Behav. Sci., 3:92-215. Anxiety and tension (Hermann M. Biggs Lecture). Bull. N.Y. Acad. Med., 34:429-44. 1959 With A. Rabe. The influence of drugs on memory fixation time. Am. Psychol., 14:423. 1960 Physiology from physicians the nervous system. In: Disease and the Advancement of Basic Science (Lowell Lectures), ed. H. K. Beecher, pp. 234-51. Cambridge, Mass.: Harvard Univ. Press. 1961 The program of the Mental Health Research Institute. Behav. Sci., 6:66-71. The architecture of knowledge and neural functions. In: Lectures in Experimental Psychiatry, ed. H. W. Brosin, pp. 147-63. Pitts- burgh: Univ. of Pittsburgh Press. Fixation of experience. In: Brain Mechanisms and Learning, ed. A. Delafresnaye, pp. 21-35. Oxford: Blackwell Scientific Publica- tions. To prevent another world war truth detection. J. Conflict Resolu- tion, 5:212-18. 1962 With B. Libet. An analysis of some correlates of steady potentials in mammalian cerebral cortex. Electroencephalogr. Clin. Neuro- physiol., 14:445-52.

RALPH WALDO GERARD 1963 209 Material basis of memory. I. Verb. Learn. Verb. Behav., 2:22-63. The nosology of schizophrenia: A cooperative study. Am. l. Psy- chiatry, 120:16-29. With T. I. Chamberlain and G. H. Rothschild. Drugs affecting RNA and learning. Proc. Natl. Acad. Sci. USA, 49:918-24. With T. i. Chamberlain and P. Halick. Fixation of experience in the rat spinal cord. J. Neurophysiol., 26:662 - 73. 1965 Intelligence, information and education. Science, 148:762-66. 1966 Building better brains. In: Enzymes in Mental Health, ed. G. {. Martin and B. Kisch, pp. 194-200. New York: J. B. Lippincott. 1967 Shaping the mind: Computers in education. In: Applied Science and Technological Progress, pp. 207-28. Washington, D.C.: U.S. Govt. Print. Off. for the National Academy of Sciences. 1968 With N. B. Mattson. Typology of schizophrenia based on multidis- ciplinary observational vectors. In: Recent Advances in Biological Psychiatry, vol. 10, pp. 507-34. New York: Plenum Press. The neurophysiology of purposive behavior. In: Purposive Systems, Proceedings of First Annual Symposium of American Society of Cyber- netics, ed. H. van Foerster, J. D. White, L. J. Peterson, and J. K. Russell, pp. 25-34. New York: Spartan Books. Build thee more stately mansions, O my soul. Recent Adv. Biol. Psychiatry, 10: 1 -5. 1969 Computer assisted learning: Introduction & general considera- tions. Proc. Natl. Acad. Sci. USA, 63:573 - 79. Hierarchy, entitation & levels. In: Hierarchical Structures, Proceedings of the Symposium, ed. L. Whyte, A. G. Wilson, and D. Wilson, pp. 215-28. New York: American Elsevier.

210 BIOGRAPHICAL MEMOIRS 1975 Is the age of heroes ended? In: The Life and Contributions of Walter Bradford Cannon, ed. C. McC. Brooks, K. Koisumi, and l. O. Pinkston, pp. 197-208. State Univ. of New York. The minute experiment and the large picture. In: The Neurosciences: Paths of Discovery, ed. F. G. Worden, i. Swazey, and G. Adelman, pp. 456-74. Cambridge, Mass.: MIT Press.

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Biographic Memoirs: Volume 53 contains the biographies of deceased members of the National Academy of Sciences and bibliographies of their published works. Each biographical essay was written by a member of the Academy familiar with the professional career of the deceased. For historical and bibliographical purposes, these volumes are worth returning to time and again.

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