Who was John Fulton?
A pioneer in cognitive neuroscience
Gordon Shepherd and Cynthia Tsay
Cognitive neuroscience refers to studies of the brain that give insight into the neural basis of higher mental faculties involved in such behavior as perception, emotion, memory and decision making. It had its origins in the early years of neuroscience around 1900, when it became clear that the nervous system is built of cells called neurons, which form networks of connections that mediate behavior. The first steps were taken by anatomists and physiologists who began on the one hand to stimulate sites in the brain to reproduce specific types of behavior, and on the other to remove sites to demonstrate loss of specific functions. One of those leading the way was John Farquhar Fulton.
By 1940 Fulton was renowned in the world of medicine and brain studies, as illustrated by the following anecdote. A young medical doctor, Hsiang-Tsung Chang, was trapped in Nanking, China, during the Second World War. With battles raging all around he found one dayin the hospital library a copy of “Physiology of the Nervous System” by John Fulton. Inspired, hewrote a letter addressed to “John Fulton, Yale University, United States” expressing his desire to work with him. A month later a telegram arrived: “Admission accepted with support”. In a hair-raising circuitous journey by air and boat he managed to make it to Yale and presented himself to Fulton at his laboratory in the Sterling Hall of Medicine in 1943.
Who was Fulton, known world-wide, now largely forgotten? What did he do as a founder of cognitive neuroscience, and what has become of his legacy?
John Farquhar Fulton Jr. was born on November 1, 1899, in St Paul, Minnesota, the son of prominent physician. From early on he had an inquiring mind and a friendly outgoing personality. Graduating from high school he applied to Harvard for college, but was turned down as needing improvement in his academic record; undeterred, he gained this after a year at the University of Minnesota and admittance to Harvard as a sophomore. He immediately plunged into laboratory work in his biology classes and especially in two summer projects at a biological station in Bermuda, which produced multiple published studies (eventually a total of 8). These won him not just one but two of the highest undergraduate prizes at Harvard, and a Rhodes scholarship to Oxford (Fig. 1). This seemingly effortless success was in fact obtained by fanatical diligence in the laboratory, which was to characterize his academic career.
Beginning at Oxford in 1921, he entered the circles around Professor Charles Sherrington, widely regarded as the leading neurophysiologist of his generation, and around the widow of Sir William Osler, who had been the most famous physician of that era. Both Osler and Sherrington were avid antique book collectors and science historians. Harvey Cushing, who was creating the new discipline of neurosurgery, also an avid book collector and historian, was also a member of their circles on frequent visits from Harvard (Fig. 2A). They welcomed the young Fulton into their confidence, and in the case of Sherrington, as a student in his laboratory. And what a laboratory! Over the next few years John Eccles, Derek Denny-Brown, H.G.T. Liddell, R.S. Creed, and now Fulton, working with Sherrington, established the modern basis of the reflex physiology of the spinal cord. This work led to Sherrington’s Nobel Prize in 1932 together with Edgar Adrian of Cambridge.
Fulton’s work toward a DPhil as part of this activity was so extensive that it resulted in a thick tome published by Oxford University Press in 1925 as Muscular Contraction and the Spinal Cord. Reflecting his newfound passion for science history, the experimental results were accompanied by an exhaustive review of all earlier studies of movement physiology dating from the Greeks. Unfortunately, it also included a significant error in interpreting what was called “the angle”, a perturbation in the recording of the reflex, which turned out to be due to a flaw in the recording. It was the kind of misinterpretation of an instrumental artifact that plagues all experimental work.
During this time Fulton married Lucia Wheatstone, heiress to a timber fortune in Maine (Fig. 2B). This gave him the wherewithal to indulge his love of ancient books in science, and he began acquiring a collection that matched those of his mentors.
Cushing, notoriously tough on his staff, took a liking to Fulton and arranged for him to receive an accelerated medical degree at Harvard (some called Fulton the son that Cushing never had). During 1925-27, Fulton pursued medical studies, then spent a year as Cushing’s assistant in neurosurgery (Fig. 3). This not only gave him invaluable experience in Cushing’s tactics, strategy and philosophy in founding the new field of neurosurgery, but also a personal bond in his interactions with leading scientists of the day as well as his newfound passion for classic books and history. Fortune continued to smile on the young investigator: in the course of his work Fulton noted a case of increased blood flow to the visual cortex during visual experience, which is now realized as a step toward modern brain imaging. He also collaborated with Jaime Pi-Suner on several papers comparing the responses of muscle spindles and tendon organs to contraction and stretch, which became a part of the standard textbook description of muscle reflexes.
In 1928, Milton Winternitz, dean of the Yale School of Medicine, offered Fulton at the age of 28 a professorship of physiology at Yale as part of revitalizing the medical school following the Flexner Report. After much soul searching, Fulton instead leapt at the chance to return to Oxford as a lecturer in physiology and tutor at his beloved Magdalen College. He couldn’t pass up the chance to continue the exciting physiological studies, under the dreaming spires, surrounded by ancient books and congenial mentors. However, Winternitz, known as a “steam engine in pants”, didn’t give up; later in 1928 he increased his offer to Chair of Physiology and a Sterling professorship. With Cushing and Sherrington’s blessing, Fulton accepted, arriving in 1930 at the tender age of 30.
Fulton’s aim was to launch a Laboratory of Physiology in emulation of Sherrington’s at Oxford, and create a new field of primate brain research In emulation of Cushing, applying the principles of human neurosurgical skill and postoperative care he had learned under Cushing. As a result, during the decade of the 30s he established the first primate research laboratory in the United States, one of the leading laboratories in the world, especially opening the primate brain to physiological investigation. In parallel he brought in a senior neurophysiologist, Johannes Dusser de Barenne, as Sterling Professor of Physiology, who carried out pioneering studies of the organization of the cerebral cortex.
Among the many advances by Fulton and his collaborators, the study with Carlyle Jacobson in 1935 of the behavior of a chimpanzee named Becky attracted the most interest (Fig. 4). The dramatic effects of the removal of the prefrontal cortex in reducing agitated behavior stimulated a neurologist, Egaz Moniz, to apply this approach to ameliorating human mental disorders, which led to widespread use of prefrontal lobotomy on schizophrenic and depressed patients, especially in the United States. As a promising method for treatment of these disorders, prefrontal lobotomy gained a Nobel Prize for Moniz in 1949, recognition that many believed should have been shared with Jacobson and Fulton. Fulton, initially encouraged by the obvious docility of the previously agitated patients, became increasingly concerned with the abuses that occurred in rapidly producing the lobotomies. With the introduction of chlorpromazine as a major tranquilizer in 1954, control of mental disorders passed to the new generation of neuroleptic drugs.
Fulton’s experiments with Jacobson placed him among the pioneers in the scientific basis for physiological investigation of the human brain in relation to behavior, and among the founders of modern cognitive neuroscience. The next step in this direction was Brenda Milner’s study of the effects of bilateral removal of the hippocampus on loss of memory in patient HM beginning in the 1950s. As cognitive neuroscience began to form as a new discipline in the 1980s, Fulton’s legacy was taken up especially by Patricia Goldman-Rakic in the new Department of Neurobiology at Yale with experiments using single neuron recordings to identify mechanisms of learning and memory in primates (Fig. 5).
Along with the experiments, Fulton indulged his passion for writing about the history of physiology. Selected Readings in the History of Physiology in 1930 was followed in 1932 by The Sign of Babinski with A. D. Keller, based on his training with Cushing, and in the same year his historical magnum opus the Bibliography of Robert Boyle, the founder of modern chemistry, responsible among other things for Boyle’s Law, that the volume of a gas varies inversely with the pressure.
Meanwhile the department was humming with many projects. Dusser de Barenne carried out a series of studies of cortical physiology through the 30s until his early death in 1940. His main assistant was Warren McCulloch, with whom he developed novel methods such as activation by local strychnine application and local temperature probes. Margaret Kennard came to the laboratory in 1932 and performed experiments until 1943 on the effects of cortical and striatal ablations on motor functions.
Fulton loved the life of the laboratory and could be found regularly carrying out and analyzing primate brain surgeries with Jacobson, Kennard, and others, while also taking a keen interest in many of the other projects. He especially hated weekends and holidays, unless he could get in to the lab and do an experiment. The success of the laboratory, however, also meant that meetings, invited talks, and administrative duties increasingly drew him away. Even here he made important contributions. He had long thought of writing a textbook of Physiology of the Nervous System, which finally came out in 1938. This became an instant classic worldwide, and was the book that inspired Chang to write to Fulton. For this book Fulton persuaded Rafael Lorente de No, a former student of the founder of neuroanatomy Ramon y Cajal, to write a chapter on the neuronal organization of the cerebral cortex. This became a classic, and was a stimulus to the eventual concept of a cortical column and the breakthroughs in understanding the cerebral cortex as revealed by Vernon Mountcastle, David Hubel, Torsten Wiesel, and others in the 1950s and 60s.
With the increasing numbers of articles in neurophysiology there was a bottleneck in getting published, so Fulton founded the Journal of Neurophysiology, also in 1938. This was facilitated by his personal relations with the publisher, Charles C. Thomas, and by his personal financial support. The Chief Editor was Ralph Gerard, and the Editorial Board contained other prominent neurophysiologists of the day. The Journal of Neurophysiology quickly established itself beside the British Journal of Physiology as the prime site for publication of leading articles in the basic physiology of the nervous system
Meanwhile, as Cushing neared early retirement at Harvard, Fulton encouraged him to move to Yale, which he did in 1933. Together with Arnold Klebs, another medical historical enthusiast, Fulton and Cushing pledged to donate their book collections as the basis for building a new “Medical Historical Library” at Yale, which finally took place in 1940, just after Cushing’s death. This beautiful addition to the Yale Medical Library was an architectural amalgamation of ancient libraries in Europe. It immediately took its place as one of the great medical historical libraries in the world.
When in 1941 the U.S. entered the Second World War, Fulton began contributing to the war effort by developing a unit of Aeronautical Research to test the effects of altitude and gravity on brain function experienced by aircraft pilots. The unit continued through the war years (1942-45), ending in 1948. This activity was accompanied by visits to similar laboratories around the country
His best-known activity during the war was through his long-time friendship with Howard Florey, who around 1940 was developing penicillin into a therapeutic drug after its earlier discovery by Alexander Fleming. A female patient in New Haven, extremely ill with pneumonia, was brought to Fulton’s attention. He was able to arrange for a small amount of the new drug, which turned out to be just enough to bring about a dramatic recovery. It was one of the first demonstrations of the efficacy of antibiotics in treating human disease.
After the war Fulton turned to a long delayed project. Cushing had written the classic biography of the revered physician William Osler, and it had become obvious that Fulton should do a biography of the revered neurosurgeon Cushing. With help from his assistants Madeleine Stanton and Elizabeth Thompson, he wrote Harvey Cushing: A Biography, published by Oxford University Press in 1947, which won a Pulitzer Prize.
Physiological brain studies continued in the 1940s, with Fulton being invited to give numerous talks on the work on brain and behavior, with special interests in revealing relations between frontal lobe cortical physiology and autonomic functions. In 1943 H.T. Chang arrived from China. He opened a new field of research by obtaining the first microelectrode recordings from cells in the cerebral cortex, with evidence of the active physiological properties of dendrites and dendritic spines. In 1955 he returned to China to become the country’s leader of neurophysiology and brain research. Patrick Wall from England joined the laboratory from 1947-1950, working on problems in primates, and later became a leader in studies of mechanisms of pain. Paul MacLean came in July 1949 to take part in training both the chimpanzees and humans with lobotomies. He worked on limbic areas of the brain, which led to his concept of the “triune brain”, containing representations of a crocodile, horse, and monkey brain, a highly popular way of describing the new research on the brain. Moving to the NIH in 1957, MacLean was a leader in cellular investigation of emotions and other limbic functions.
Margaret Lennox was in the laboratory during the 1940s, focusing on the effects of brain ablations on the electrical activity of the brain as measured by the electroencephograph. A young neurosurgeon, Karl Pribram, came to the laboratory from 1947-1951, applying his surgical and clinical expertise to the chimpanzee studies. GMS studied with Pribram during a summer at MIT in 1956, when he was beginning to form his ideas of the holographic nature of brain function.
Fulton enjoyed good health early in his life, but on one of his many trips during the war he contracted a serious case of coccioidomycosis, requiring an extended recovery. After the war, a life style of consuming excessive alcohol began to take its toll. Ironically, among his letters home as a teenager was a vow never to come under the influence of alcohol. However, in his student days at Oxford, with other classmates, he turned to imbibing port to keep warm in his cold college rooms. Drinking continued with the intense social life that accompanied many academic careers in the 20s, 30s, and 40s. Along the way he became a diabetic. He had several hospitalizations in the late 40s and into the 50s to try to bring the alcoholism under control. It was painfully obvious to all his close associates and friends, and was the main factor in the reluctance of those who knew him best to take on the task of writing a biography.
In his diary for January 8, 1951 Fulton records: “Since Dr. Cushing’s death [ed. in 1939], I have been agitating for the establishment of a Chair of the History of Medicine at Yale, as it had been one of the things that Dr. Cushing had hoped might come about in connection with building the Historical Library; indeed, in his will he had asked whether it might be possible to transfer his Sterling Professorship of Neurology to one of medical history.” Fulton added that he never considered himself as a candidate.
In 1950, Whitney Griswold became the new President of Yale, and just before Christmas, the diary records that Griswold “summoned me to Woodbridge Hall to say that he had at his disposal a new Sterling Professorship and that he proposed to assign it to the Medical School, if I would accept it as Sterling Professor of the History of Medicine” (Fig. 6). After a period of “soul searching” Fulton agreed. Today, there is a program in the History of Science and Medicine, a semi-autonomous unit within the university Department of History. There is also a Section in the History of Medicine within the medical school.
Did Fulton’s campaign for a chair in medical history prove to be his undoing, used by his dean to move Fulton out and the dean into his Sterling professorship and department chair? It is possible. On the other hand, the diaries during this period begin to record problems with administrative red tape and interactions with colleagues. Fulton by nature had little patience with anything that stood in the way of academic work and especially that took time from doing experiments in the laboratory. This impatience was heightened by his enthusiastic good humor, and the confident impression that goes with financial independence. Couple that with the increasing evidence of alcoholism and there was an obvious opportunity to save embarrassment for both Fulton and Yale in a way that saved face for everybody.
Thus began the final chapter of Fulton at Yale, with the increasing impression of him more as a historian of science than as an experimental physiologist. The experiments on chimpanzees by several students and younger colleagues in physiology continued. Lectures in distinguished series continued. Honors flowed. In addition to the honorary OBE from Britain for services during the war were 35 honorary degrees and other awards from around the world, culminating in an honorary doctorate from Oxford in 1957 (significantly a D.Litt., in recognition of his services to medical history, not his many contributions to science). Fortunately he managed to make it to Oxford to receive it despite the ravages of alcohol, which can be seen clearly in the photograph of him and Lucia taken on the occasion. Although he published nothing in original science after 1954, his enthusiasm for understanding the brain and behavior continued to be recorded in the diaries up to the last volume (46) in early 1960. He died on March 29, 1960, at the age of 60.
Fulton is one of Yale’s most decorated faculty. Extensive materials assembled at the Sterling Memorial Library at Yale relate to his scientific achievements, historical writings, colleagues and students, and life as an administrator; the table of contents alone runs to 58 single-space pages, making him one of the most documented scientists anywhere. In addition are family letters during his early years, and a massive 46 volume diary kept for over 30 years of his professional life, housed in the Yale Medical Historical Library. This has been digitized and is available for scholarly study. Transcriptions of over 40 selected topics from the diaries by CT have been the basis for the present account. A full bibliography is available.
Fulton, J.F., A Bibliography of the Honourable Robert Boyle, 2nd edition (Oxford: Clarendon Press, 1961) (originally published in Proceedings of the Oxford Bibliographical Society in 1932)
Fulton’s special collecting interest was physiological works from the 16th to 18th century. In addition to his major texts in physiology, he authored or coauthored biographies of Harvey Cushing, Benjamin Silliman, and Michael Servetus, and bibliographies of Fracastoro’s poem Syphilis, Luigi Galvani and his nephew Aldini, Richard Lower and John Mayow, Joseph Priestley, and Robert Boyle.
Training: (1922-1930) Harvard B.A.; Rhodes Scholarship; Oxford DPhil; Harvard Medical School M.D., assistant to Harvey Cushing; instructor at Oxford University; Sterling Professor of Physiology; Sterling Professor of the History of Science and Medicine.
1928 Tendon organ with Pi-Suner
1930 Yale Sterling Professor age 30
1935 Sub-human primate frontal lobe behavior: Jacobson
1935 Jacobson frontal lobes and behavior: origins of cognitive neuroscience
1930s Dusser de Barenne cortical physiology
1938 Lorente de No cortical anatomy to columns
1940s Warren McCulloch brain and computer
1940s Paul MacLean and triune brain
1940-50s H-T Chang dendrites and China neurophysiology
1940s Aeronautics laboratory
1943 Introduction of penicillin
1940-50s Studies of frontal lobe behavior
Publishing and Libraries
1932 Boyle bibliography (2nd ed. 1961)
1938 Founding of the Journal of Neurophysiology
1938 Physiology of the Nervous System
1940 Founding of the Yale Medical Historical Library
1942 Founding of the Journal of Neurosurgery
1947 Cushing biography (Pulitzer Prize)
1951 Created the Section in the History of Medicine
1926 The “angle” artefact negated much of his thesis work
1950s Scandal of human lobotomies obscured his basic research establishing cognitive neuroscience as a new discipline
1940-50s Increasing alcoholism limited his research contributions
1947 Fulton’s Life of Harvey Cushing contains no mention of one of Cushing’s closest friends, John Fulton himself
1951 Removed to history of science and medicine chair, without promised funds for expansion
1927-1960 The diaries are a valuable source of information on what happened, but less on immediate insights into how it happened and reflections on what went right and wrong.
 It also enabled him to buy up the print run of Muscular Contraction to limit the propagation of the error of his ways.
 After moving to Chicago in 1941 McCulloch teamed with Walter Pitts to write the classic paper in 1943 on neurons as logic operators; their discussions with John von Neumann were central to the latter’s development of the “von Neumann architecture” in 1945 for modern digital computers.