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Biographical Memoirs HERBERT SPENCER GASSER July 5, 1888–May 11, 1963 BY MERRILL W. CHASE AND CARLTON C. HUNT HERBERT GASSER was a major scientific figure. An outstanding physiologist, he was a pioneer in the field of neurophysiology. In addition, as Director of the Rockefeller Institute for Medical Research from 1935 until 1953, he exercised an important national and international influence on science. This long-overdue memoir has been written almost thirty years after his death. We both knew him at the Rockefeller Institute and have had access to extensive archival material. Though our information about his early years is limited, Gasser's autobiography (1964), written characteristically with great reserve and in the third person, contains much of interest. ORIGINS AND EARLY YEARS IN WISCONSIN Herbert Gasser was born in 1888 in Platteville, a small town in southwestern Wisconsin. His father, Herman, was an immigrant from the Tyrol, who, after working as a pharmacist, studied medicine and became a practicing physician. His mother, Jane Elizabeth Griswold Gasser, came from a family of early Connecticut settlers. The given names of their first child, Herbert Spencer, stemmed from his father's perusal of books by Wallace, Darwin, and Spencer after

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Biographical Memoirs local newspapers attacked the concept of evolution. There were two younger siblings, a sister and a brother. Gasser recalled his youth in Platteville: “. . .excursions into the countryside, fishing in the summer and skating in winter. There were few distractions, no cinema or radio, and travel was by horse and buggy. Children had to find their own amusement.” Reading was of great importance to him. He also built furniture, taking pleasure in benchwork. Fine craftsmanship remained a lifelong interest. Young Gasser had a simple box Kodak camera that he supplemented with much “improvised equipment.” As he remembered: “This experience later turned out to be a good training for a physiologist.” In accordance with his father's wishes, Gasser studied at the state normal school in Platteville. A year was lost because of an unspecified but serious illness; he graduated in 1907. Gasser wished to continue studies in engineering, but his father preferred that he enter medical school. Gasser was not attracted to a country doctor's life. He wanted further education but felt medical training was too great a price to pay. At last, he and his father struck a compromise that he later considered a turning point in his life: “The university would be allowed if no specialization were to take place.” Gasser attended the University of Wisconsin at Madison, receiving a bachelor's degree in 1910 and, in the next year, a master's degree. At Wisconsin he took some medical school courses. He wrote about himself: “That is how it came about that his introduction to physiology was through a lecture course given by Prof. Joseph Erlanger before the latter's departure to Washington University in St. Louis. Erlanger was a beautifully clear lecturer, but the subject matter he presented differed so widely from what was anticipated that it amounted to a revelation. Gasser listened to the lectures with bewilderment and felt he was getting

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Biographical Memoirs only a feeble grasp of their content. He had no realization that Erlanger was aware of his attendance.” Gasser remained at the university as an assistant in biochemistry, meanwhile completing his preclinical subjects, and then was appointed an instructor in physiology. As a student at Wisconsin, Gasser was involved in several studies with Loevenhart, an imaginative chemist, on the effects of oxygen want and with Meek on the responses of the heart to exercise. At the medical school in Wisconsin, there were young faculty who were enthusiastic about research and Gasser came to realize that “medical science could be considered a discipline in its own right. In this light medicine appeared in a form which Gasser found acceptable.” Because Wisconsin had only a two-year school, Gasser had to complete his medical education at another school, and he wisely chose Johns Hopkins. It apparently required some persuasion to get his father's consent, and the costs were a considerable strain. JOHNS HOPKINS The Johns Hopkins medical curriculum provided for elective time; Gasser utilized this for study in physiology. Professor William Howell suggested that he work on mechanisms of blood clotting, studies that he did not complete until after leaving Baltimore. Howell confirmed Gasser's results. Gasser was pleased that this independent work had been successful, but it was not a field of long-term interest to him. When Gasser received his M.D. in 1915, Howell offered him a position in physiology, but Gasser did not feel free to accept it and returned to Wisconsin where Loevenhart appointed him as an instructor in pharmacology. As he wrote: “The salary was higher and the location was adapted to

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Biographical Memoirs helping in the support of a younger sister and brother at the University of Wisconsin.” ST. LOUIS (1916-31) A year later Gasser had an opportunity to return to physiology. Joseph Erlanger invited him to take a position in his new department at the recently reorganized Washington University School of Medicine in St. Louis. In the spring of 1917 the United States entered the First World War. Gasser worked with Erlanger on problems related to the war effort, investigating wound shock and publishing eight papers on this topic. In the summer of 1918 he joined Loevenhart at a chemical warfare station at American University in Washington, D.C., and worked on Lewisite. At the war's end he returned to St. Louis, in December 1918. At this time he had no clear research direction but had some interest in the nervous system. There were discussions in the department about the problem of recording nerve activity, and Gasser was aware of the need for a new and innovative technical approach. A classmate from Johns Hopkins, H. S. Newcomer, who had a background in physics, had constructed a three-stage amplifier using thermionic valves. He visited St. Louis, where he and Gasser tried using this amplifier in conjunction with a string galvanometer to record impulses in the phrenic nerve. While the amplification was sufficient to detect the nerve action potentials with the string galvanometer, the latter was too slow to record the fast potential changes with fidelity. A recording device with sufficient speed did exist, the cathode ray or Braun tube, in which a beam of electrons was deflected by potentials applied to pairs of vertical or horizontal plates. Bernstein had mentioned its possible use for the recording of rapid nerve potential changes in his

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Biographical Memoirs book on “Electrobiologie” in 1912. But there was a problem of sensitivity. As Gasser wrote: “In the old Braun tubes, cathode beams were composed of high velocity electrons wrenched from cold cathodes by high voltages. There was still a large gap between the sizes of the potentials needed to deflect them and those which could be produced through the augmentation of a nerve action potential with the aid of the new amplification. Then there occurred an event crucial in its significance.” This crucial event occurred in 1920, when both the Physical Society and the American Physiological Society happened to meet at the same time in Chicago. Professor Horatio Williams, of Columbia, told Gasser there would be a paper read at the Physical Society meeting that would interest him. J. B. Johnson of Western Electric Laboratories was describing a modification of the Braun tube with a heated cathode, allowing the tube to be operated at lower voltages and thus increasing its sensitivity. Newcomer's amplifier had enough gain for the modified Braun tube to record nerve potentials. It was following this development that Erlanger joined Gasser in the studies on nerve potentials. Their first attempt to acquire a cathode ray tube met with resistance from Western Electric, which refused to sell them one. Undaunted, they made a tube by coating a phosphor inside an Erlenmeyer flask. Western Electric subsequently relented and sold them a tube. To provide for triggering of sweeps of the cathode ray oscillograph and for stimuli to be delivered to the nerve, much ancillary apparatus had to be constructed. This was done largely with mechanical devices. Condenser discharges were applied to the plates deflecting the beam horizontally for the time axis, while vertical deflection was produced by the amplified nerve signal. Records were obtained by holding film on the tube face in the dark room and repeating

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Biographical Memoirs the sweeps until a sufficient exposure was obtained. To minimize vibration, the apparatus was mounted in a basement laboratory on a foundation separate from that of the building. Nonetheless, passing streetcars often disturbed the delicate recordings. The images were blurred, and careful correction had to be made for the nonlinearities in horizontal and vertical deflections. In spite of technical difficulties, in shortly over a year Erlanger and Gasser were ready to publish a first paper on the nerve potentials recorded with the oscillograph. Writing some forty years later, Gasser recalled: “The most difficult step in opening up a new field had been taken. Ever afterward, Gasser never had any doubt about the direction he would follow.” It was evident from these earliest studies that the action potential recorded from the frog sciatic nerve was compound in nature (i.e., it resulted from the summation of potential changes in many individual axons). But the characteristics of the action potentials in the individual axons were not known. Much remained to be done. In 1921 Gasser was made head of the Department of Pharmacology at Washington University, a position he held until 1931. The pharmacology position was accepted with some misgivings and with the stipulation that he would be free to follow his own line of research. Gasser's interest in pharmacology probably stemmed from the influence of Loevenhart. Later, Gasser described taking the chair at Washington University as his “final excursion into Pharmacology.” Already, Gasser had come to view himself primarily as a physiologist. Shortly after Gasser became professor of pharmacology, Abraham Flexner visited the Executive Faculty and, referring to his boyish appearance, asked: “What have you been doing making freshmen Professors?” Soon thereafter Flexner indicated that the General Education Board of the Rockefeller

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Biographical Memoirs Foundation would support an extended stay in Europe, for him to gain further experience in pharmacology and to better his knowledge of foreign languages. Gasser had already planned to attend the International Physiological Congress in Edinburgh; there he delivered a paper on the new electrophysiological technique and met with European physiologists. Gasser was soon elected to membership in the Physiological Society of Great Britain (1924). Exploitation of the new oscillographic technique had barely begun when Gasser began his sojourn abroad, which was extended to a second year. He first worked on muscle with A. V. Hill at University College, London. They published a long paper on the dynamics of muscular contraction; and, in another paper with Hartree, Gasser found that the mechanical and thermal responses of muscle were inseparable. He then went to Straub's laboratory in Munich and, following that, to Louis Lapique in Paris. Lapique was well-known for his work on nerve and muscle excitation and for his ideas about “chronaxie,” an indirect measure of the speed of voltage change of the tissue to an applied current pulse. When Gasser showed his records of compound action potentials to Lapique, “he suggested forth-with that the velocity differences might be associated with the sizes of the fibers.” His visit with Lapique was the only one during the European trip that had direct relevance to Gasser's long-range interests and led to a special publication. Gasser next went to Henry Dale's laboratory. They studied the development of increased sensitivity of muscle to nicotine and acetylcholine following denervation. Doing no further work on muscle, he later wrote an important review on the subject of muscle contracture (1930). Back in St. Louis, Gasser resumed studies on peripheral nerve with Erlanger. George Bishop had joined in the ef-

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Biographical Memoirs fort and brought to the group a considerable array of skills, both practical and theoretical. His knowledge of physics was extensive, and he was very helpful in designing the recording conditions so as to avoid artifacts. In 1924 a paper titled “The Compound Nature of the Action Current in Nerve as Disclosed by the Cathode Ray Oscillograph” was authored by Erlanger and Gasser “with the collaboration in some of the experiments of George H. Bishop.” This important paper was a milestone in the development of knowledge about the physiology of peripheral nerve. No less than eight further papers carried Bishop as a joint author. The excised sciatic nerve of the bullfrog was the principal experimental subject, mounted in a closed humidified chamber on an array of electrodes for stimulation and recording. One recording electrode was placed at the cut end of the nerve, the other some distance centrally, and the potential difference between the two was recorded. With such extracellular recording, impulses arriving at the electrode located on the intact nerve produced a negative deflection. To provide a quantitative understanding of the action potential recorded from the multiaxonal nerve at different distances from the site of stimulation, it was necessary to know the number and sizes of the axons in the nerve, the form of the action potential in the individual axon, the speed of conduction of the action potential in different axons and its relation to axonal diameter. It would later be found that after-potentials also had to be taken into account. Gasser was particularly interested in reconstructing the “compound” action potential by summing the predicted potential changes in all the individual axons of the bullfrog nerve, at a particular conduction distance, and comparing this with the recorded action potential. This approach proved

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Biographical Memoirs to be very useful and one that Gasser used for his entire research career. From cross sections of the nerve, the numbers of axons (nerve fibers) of various sizes were counted, the action potential of each size group was approximated by a triangular waveform of depolarization, the group of axons of a given size being assumed to have a constant conduction velocity that was proportional to their diameter. Only the myelinated axons of bullfrog peripheral nerve, the so-called A fibers, were studied in their early work. When recording near the cathode, where all the axons were stimulated at the same instant, the action potential had a simple form, the axon potentials in all the axons being nearly synchronous. But with increasing conduction distance the difference in the time of arrival of the action potentials in the fastest and slowest axons increased progressively. The successive elevations in the potential record were produced by axons conducting progressively more slowly; the deflections were called alpha, beta, gamma, and delta. The best fit between the actual compound action potential and the reconstructed plot was obtained when the action potential in the individual axon was given a duration of 1 millisecond, the amplitude of the externally recorded action potential in each axon was made proportional to axonal size, and conduction velocity was related directly to axonal diameter. Other important questions were pursued. For example, was conduction in the various fibers independent? As the strength of a stimulus was increased, first the largest and then the smaller axons were excited. If a weak shock that excited only the largest (A alpha) axons was given and then, while they were refractory, a second stronger shock was delivered, the latter evoked no alpha wave but only later deflections. This indicated that action potentials in the larger and smaller fibers were conducted independently. Subsequently, it was possible to record the action potential in an

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Biographical Memoirs individual axon and to measure directly its time course. Studies were also made on the absolute and relative refractory periods following impulse activity. The first observation of the C fiber action potential was made by Peter Heinbecker, a research fellow with George Bishop. Applying exceedingly strong shocks to the cervical sympathetic nerve of the turtle, he saw small, very late potential elevations that conducted very slowly. These were the responses of unmyelinated or C fibers. Erlanger and Gasser explored the details of the C fiber potentials, publishing a detailed paper in 1930. Decades later Gasser was to return to this problem. The threshold of C fibers to electrical stimulation was nearly 100 times that of A fibers. C fiber action potentials were longer in duration than those in A fibers; the proportionality between conduction velocity and diameter was also different, although the relationship appeared to be linear. Myelinated preganglionic sympathetic fibers were designated B fibers. Their properties differed from A fibers in duration of action potential, relation between fiber diameter and conduction velocity, and in after-potentials. The order in which nerve fibers were blocked by cocaine was found to be from smallest to largest. Conversely, pressure blocked the largest fibers first, the smaller fibers later. This was useful in relating function to the various fiber groups, a topic of lasting interest to both Gasser and to Bishop. Toward the end of his St. Louis tenure, Gasser, together with Erlanger, became interested in the potential changes that follow the brief nerve impulse, the after-potentials, and their effect on excitability. After the nerve impulse or “spike,” the potential only gradually returned to the baseline level; the potential during this period being negative to the baseline, it was called the negative after-potential. During the nega-

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Biographical Memoirs and on appropriate contributions of after-potentials to the recorded action potentials. With the higher resolution of the electron microscope, Gasser examined olfactory nerve fibers in mammals and in the pike. The unmyelinated fibers were found to be extremely small. The length of the olfactory nerve in the pike permitted recording from these nerve fibers. Although afferent, they showed action potentials that differed from those of mammalian dorsal root C axons, showing a simple form consonant with all of the axons having nearly the same diameter. Gasser also sought an answer to a problem in the recording of the compound action potential, which had long bothered him. The problem arose from the fact that the potential was recorded by two electrodes, one at the end of the nerve and the other some distance centrally. Even if the nerve end was crushed and treated with a local anesthetic, impulses in some fibers propagated into the stretch between the recording electrodes, creating an artifact. The solution was to integrate the response recorded by a pair of electrodes located quite close together on the nerve, a procedure that he called a tangent lead. The results showed that some of the previously described elevations were artifactual; in the compound action potential produced by myelinated fibers of mammalian cutaneous nerves there were only two peaks, the alpha and delta. This finding was consonant with the size distribution of myelinated axons in these nerves. The paper describing these studies was his last. Working alone or with one or two collaborators, Gasser was a very focused investigator, intensely concentrated on his subject. Most of his research was, in fact, within a circumscribed although important area. An approach he utilized throughout his career was reconstruction of the com-

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Biographical Memoirs pound action potential from the calculated sum of the activities of the unitary axons. He returned to this approach, first used in St. Louis, after his retirement as Director. As Lloyd later wrote of Gasser's style of selecting problems: “. . . Gasser espoused the principle that there are two times for working on a problem—before anyone has thought of it and after everyone else has left it. As a result, Gasser was always the innovator or the finalist.” GASSER IN PERSON As Director of the Rockefeller Institute, Gasser was a striking figure. Tall, elegant, and graced for many years with an extraordinarily youthful appearance, his formidable intellect made him impatient with trivial conversation. While he was clearly a fastidious man of great integrity, those privileged to know him realized that he was also a warm, engaging person who treated friends with much kindness, loyalty, and concern. He had a keen sense of humor, enjoyed puns, and could be very good company. Fine art, classical music, and good food were among his pleasures. He read widely and his knowledge on many topics was profound. His highpitched voice reflected a hormonal deficiency, but a strong personality and rigorous intellect made this unimportant. Not a facile speaker, Gasser wrote beautifully, with clarity and grace. An example of Gasser's uncompromising standards can be seen in his response to a letter from Erlanger in 1938, requesting information to be transmitted to the Nobel committee. Their colleague, Dr. Evarts Graham, had nominated them. Gasser wrote Erlanger: It must be well known in Stockholm that nominations for the Nobel Prize coming to hand with full information must be made with the cooperation of the nominee. One is thus forced into the position of appearing, at least in some measure, to nominate oneself. . . . I am greatly pleased that Evarts

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Biographical Memoirs should value our work highly enough to place it in nomination, and I am grateful to him for the proposal, but my considered opinion about the effect of our becoming a party to the proposal impels me into not consenting to do so. While such an attitude may seem quaint today, it reflected Gasser's deeply rooted antipathy to any kind of self-promotion. Gasser's laboratory days ended when he suffered a cerebral accident on April 17, 1961, at the age of seventy-three, eight years after resigning as Director. Thereafter he resided in the New York Hospital. He made a partial recovery from his stroke but died from a respiratory infection in the hospital on May 11, 1963. The fact that this memoir is written almost thirty years after his death permits some perspective on Gasser's view that independence of scientific inquiry was essential. By the time he retired in 1953, grant support had expanded considerably, and Gasser's reservations seemed old-fashioned to many. There was then an abundance of governmental money, and research of quality found support with proposals that were liberally and flexibly reviewed. Gasser had admitted that under a grant system “there is no gain-saying that accomplishments of the highest type can come out of it in spite of its shortcomings.” Now, with the stringency of funding, an applicant for federal grant support must not only predict the results to be obtained but must also demonstrate that the proposed experiments will yield the anticipated results. Thus, the investigator's independence is even more compromised than Gasser once feared. From the vantage point of the 1990s, his concerns now appear prescient. VALUED SOURCES ARE GASSER'S autobiography, published posthumously with an introduction by Joseph C. Hinsey in Experimental

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Biographical Memoirs Neurology, 10 (Suppl. 1, 1964):1-38 (our quotations from Gasser come from this), also David P. C. Lloyd's obituary of Dr. Gasser in volume 5 of the Dictionary of Scientific Biography, pp. 290, 291 (New York: Charles Scribner's Sons, 1972) and Lord Adrian's article in the Biographical Memoir of Fellows of the Royal Society, 10(1964):75-82. The history of the axonologists comes from F. O. Schmitt to MWC and is recounted in volume 2 of Advances in American Medicine: Essays at the Bicentennial in the article “The Neurosciences” by Robert J. Frank, Louise H. Marshall, and H. J. Magoun, pp. 552-616 (New York: Josiah Macy, Jr., Foundation, 1976). The reference to the status of women in science, as related to H. T. Graham, comes from M. W. Rossiter's Women Scientists in America (Baltimore: Johns Hopkins University Press, 1984). The homemade cathode ray tube and its trundling to the XIII International Congress of Physiology in Boston, 1929, is described in Hallowell Davis' memoir of Joseph Erlanger in Biographical Memoirs, vol. 41, pp. 111-39 (Washington, D.C.: National Academy of Sciences, 1976). It is to be noted that the 1936 book by Erlanger and Gasser was reprinted thirty-one years later by the trustees of the University of Pennsylvania, along with Gasser's bibliography and a foreword by David P. C. Lloyd. Gasser's picture was taken at age forty-seven, shortly after he became Director of the Rockefeller Institute for Medical Research.

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Biographical Memoirs HONORS AND DISTINCTIONS AWARDS AND MEMBERSHIPS 1924 Physiological Society (British), Ordinary Member 1934 National Academy of Sciences 1935-53 Director, The Rockefeller Institute for Medical Research 1936 Sociedad Argentina de Biologia, Corresponding Member 1937 American Philosophical Society 1942 Asociacion Medica Argentina, Honorary Member 1943 Asociacion Medica Argentina de Buenos Aires, Honorary Member Royal Society of Edinburgh, Honorary Fellow 1944-45 Nobel Prize in Physiology and Medicine (with J. Erlanger) 1946 Royal Society of London, Foreign Member Royal Swedish Academy of Sciences, Foreign Member 1947 Finnish Academy of Sciences, Foreign Member Académie Royale de Médicine de Belgique, Corresponding Member American Society of Electroencephalography, Honorary Member 1948 American Academy of Arts and Sciences Accademia della Scienze dell' Istituto di Bologna, Corresponding Member 1949 Physiological Society (British), Honorary Member 1953 Société Philomathique de Paris, Corresponding Member 1954 Kober Medal, Association of American Physicians 1958 Société de Biologie, Collège de France, Associate Member HONORARY DEGREES 1936 Sc.D., University of Pennsylvania 1940 Sc.D., University of Rochester LL.D., Washington University 1941 Sc.D., University of Wisconsin 1945 Sc.D., Columbia University 1947 LL.D., Johns Hopkins University

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Biographical Memoirs   Sc.D., Oxford University 1948 Sc.D., Harvard University 1949 Doctor honoris causa, Université Libre de Bruxelles Doctor of Medicine, Honorary, Université Catholique de Louvain 1953 Honorique de docteur, Université de Paris 1959 Sc.D., The Rockefeller University PROFESSIONAL ORGANIZATIONS American Physiological Society American Association for the Advancement of Science, Fellow Society for Experimental Biology and Medicine, President, 1937-39 Harvey Society, President 1940-42 Association for Research in Nervous and Mental Diseases American Society of Pharmacology and Experimental Therapeutics Association of American Physicians History of Science Society New York State Society for Medical Research American Neurological Association (Associate Member)

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Biographical Memoirs SELECTED BIBLIOGRAPHY 1914 With A. S. Loevenhart. The mechanism of stimulation of the medullary centers by decreased oxidation. J. Pharmacol. Exp. Ther. 5:239-73. With W. J. Meek. A study of the mechanism by which muscular exercise produces acceleration of the heart. Am. J. Physiol. 31:48-71. 1917 The significance of prothrombin and of free and combined thrombin in blood-serum. Am. J. Physiol. 42:378-94. 1918 With W. J. Meek. Blood volume. A method for its determination with data for dogs, cats and rabbits. Am. J. Physiol. 47:302-17. 1919 With J. Erlanger and R. Gesell. Studies in secondary traumatic shock . I. The circulation in shock after abdominal injuries. Am. J. Physiol. 49:90-116. With J. Erlanger. Studies in secondary traumatic shock. II. Shock due to mechanical limitation of blood flow. Am. J. Physiol. 49:151-73. With J. Erlanger. Studies in secondary traumatic shock. III. Circulatory failure due to adrenalin. Am. J. Physiol. 49:345-76. With J. Erlanger. Hypertonic gum acacia and glucose in the treatment of secondary traumatic shock. Ann. Surg. 69:389-421. With J. Erlanger. Studies in secondary traumatic shock. V. Restoration of the plasma volume and of the alkali reserve. Am. J. Physiol. 50:104-18. 1921 With H. S. Newcomer. Physiological action currents in the phrenic nerve. An application of the thermionic vacuum tube to nerve physiology. Am. J. Physiol. 57:1-26.

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Biographical Memoirs 1922 With J. Erlanger. A study of the action currents of nerve with the cathode ray oscillograph . Am. J. Physiol. 62:496-524. 1924 With J. Erlanger and G. H. Bishop. The compound nature of the action current in nerve as disclosed by the cathode ray oscillograph. Am. J. Physiol. 70: 624-66. With A. V. Hill. The dynamics of muscular contraction. Proc. R. Soc. Ser. B. 96:398-437. With W. Hartree. The inseparability of the mechanical and thermal responses in muscle . J. Physiol. (Lond.) 58:396-404. The methods of recording the electrical potential change in nerve with special reference to the use of the Braun tube oscillograph . Br. J. Radiol. 20:105-11. 1925 With L. Lapique and A. Desoille. Relation entre le degré d'héterogénéité des nerfs et la complexité de leur courant d'action. C. R. Seances Soc. Biol. 92:9-10. With J. Erlanger. The nature of conduction of an impulse in the relatively refractory period. Am. J. Physiol. 73:613-35. With J. Erlanger and G. H. Bishop. On the conduction of the action potential wave through the dorsal root ganglion. Proc. Soc. Exp. Biol. Med. 23:372. 1926 With J. Erlanger and G. H. Bishop. The refractory phase in relation to the action potential of nerve . (Proc. Am. Physiol. Soc., Dec. 1925) Am. J. Physiol. 76:203. With J. Erlanger and G. H. Bishop. Experimental analysis of the simple action potential wave in nerve by the cathode ray oscillograph. Am. J. Physiol. 78:537-73. With J. Erlanger and G. H. Bishop. The action potential waves transmitted between the sciatic nerve and the spinal roots. Am. J. Physiol. 78:574-91. With G. H. Bishop and J. Erlanger. Distortion of action potentials as recorded from the nerve surface . Am. J. Physiol. 78:592-609.

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Biographical Memoirs With H. H. Dale. The pharmacology of denervated mammalian muscle. I. The nature of the substances producing contracture. J. Pharmacol. Exp. Ther. 29:53-67. With H. H. Dale. The pharmacology of denervated mammalian muscle. II. Some phenomena of antagonism, and the formation of lactic acid in chemical contracture . J. Pharmacol. Exp. Ther. 28:287-315. 1927 With J. Erlanger. The rôle played by the sizes of the constituent fibers of a nerve trunk in determining the form of the action potential wave. Am. J. Physiol. 80:522-47. 1928-29 With J. C. Hinsey. The Sherrington Phenomenon. Am. J. Physiol. 87:368-80. 1929 With J. Erlanger. The role of fiber size in the establishment of a nerve block by pressure or cocaine. Am. J. Physiol. 88:581-91. Arthur S.Loevenhart. Science 70:317-21. 1930 With J. Erlanger. The action potential in fibers of slow conduction in spinal roots and somatic nerves. Am. J. Physiol. 92:43-82. With J. C. Hinsey. The component of the dorsal root mediating vasodilation and the Sherrington contracture. Am. J. Physiol. 92:679-89. Contractures of skeletal muscle. Physiol. Rev. 10:35-109. With J. Erlanger. The ending of the action potential and its relation to other events in nerve activity. Am. J. Physiol. 94:247-77. 1931 With H. T. Graham. Modification of nerve response by veratrine, protoveratrine and aconitine . J. Pharmacol. Exp. Ther. 43:163-85. Nerve activity as modified by temperature changes. Am. J. Physiol. 97:254-70. 1932 With H. T. Graham. The end of the spike-potential of nerve and its

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Biographical Memoirs relation to the beginning of the after-potential . Am. J. Physiol. 101:316-30. 1933 With H. T. Graham. Potentials produced in the spinal cord by stimulation of dorsal roots . Am. J. Physiol. 103:303-20. With F. O. Schmitt. The relation between the after-potential and oxidative processes in medullated nerve. Am. J. Physiol. 104:320-30. 1934 With J. Hughes. Some properties of the cord potentials evoked by a single afferent volley. Am. J. Physiol. 108:295-306. With J. Hughes. The response of the spinal cord to two afferent volleys. Am. J. Physiol. 108:307-21. 1935 Changes in nerve-potentials produced by rapidly repeated stimuli and their relation to the responsiveness of nerve to stimulation . Am. J. Physiol. 111:35-50. With D. Clark and J. Hughes. Afferent function in the group of nerve fibers of slowest conduction velocity. Am. J. Physiol. 114:69-76. 1936 With H. Grundfest. Action and excitability in mammalian A fibers. Am. J. Physiol. 117:113-33. 1936-37The control of excitation in the nervous system. Harvey Lect. 169-93. 1937 With J. Erlanger. Electrical signs of nervous activity. In Eldridge Reeves Johnson Foundation for Medical Physics Lectures. Philadelphia : University of Pennsylvania Press. 1938 Recruitment of nerve fibers. Am. J. Physiol. 121:193-202. With C. H. Richards and H. Grundfest. Properties of the nerve

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Biographical Memoirs fibers of slowest conduction in the frog . Am. J. Physiol. 123:299-306. With H. Grundfest. Properties of mammalian nerve fibers of slowest conduction. Am. J. Physiol. 123:307-18. 1939 With H. Grundfest. Axon diameters in relation to the spike dimensions and the conduction velocity in mammalian A fibers. Am. J. Physiol. 127:393-414. Axons as samples of nervous tissue. J. Neurophysiol. 2:361-69. 1943 Pain producing impulses in peripheral nerves. Res. Publ. Assoc. Res. Nerv. Ment. Dis. 23:44-62. 1945 Mammalian nerve fibers. Nobel Lecture, December 12 . In Les Prix Nobel en 1940-1944, pp. 128-41. Stockholm : Nobelstiftelsen. 1950 Unmedullated fibers originating in dorsal root ganglia. J. Gen. Physiol. 33:651-90. 1955 Properties of dorsal root unmedullated fibers on the two sides of the ganglion. J. Gen. Physiol. 38:709-28. 1956 Olfactory nerve fibers. J. Gen. Physiol. 39:473-96. 1958 The postspike positivity of unmedullated fibers of dorsal root origin . J. Gen. Physiol. 41:613-32. Comparison of the structure, as revealed with the electron microscope, and the physiology of the unmedullated fibers in the skin nerves and in the olfactory nerves. Exp. Cell Res. (Suppl.) 5:3-17. 1960 Effect of the method of leading on the recording of the nerve fiber spectrum. J. Gen. Physiol. 43:927-40.