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Sherrington, Charles Scott

SHERRINGTON, CHARLES SCOTT

(b. London, England, 27 November 1857; d. Eastbourne, England, 4 March 1952)

neurophysiology.

Sherrington was the son of Anne Brookes and James Norton Sherrington. After his father’s death, in Sherrington’s early childhood, his mother married Dr. Caleb Rose, Jr., of Ipswich. The Rose home, a gathering place for artists and scholars, helped to shape Sherrington’s broad interests in science, philosophy, history, and poetry. After attending the Ipswich Grammar School from 1870 to 1875, sherrington, encouraged by his stepfather, began medical training at St. Thomas’s Hospital in London. In 1879 improved family finances enabled him to enter Caius College, Cambridge, where he studied physiology in Sir Michael Foster’s laboratory. He worked chiefly under John Newport Langley and Walter Gaskell, who imparted t him their dominant interest in how anatomical structure reflects, or is expressed in, physiological function. From 1884 to 1887 Sherrington completed his medical courses and did graduate study and research in Europe under Friedrich Goltz, Rudolph Virchow, and Robert koch, gaining a superb grounding in physiology, morphology, histology, and pathology. In 1887 he was appointed a lecturer in systematic physiology at St. Thomas’s, and from 1891 to 1895 he served as physician-superintendent of the Brown Institution, a London animal hospital. From 1895 to 1912 Sherrington held the Holt professorship of physiology at Liverpool and from 1913 to 1935 the Wayneflete chair of physiology at Oxford.

Sherrington married Ethel Wright of Suffolk, England, on 27 August 1891; their only child, Carr E. R. Sherrington, was born in 1897. He himself was a man of diverse interests. Outside of the laboratory his activities included sports (a feature event during his years in London was Sunday morning parachute jumping from the tower of St. Thomas’s), work in many scientific organizations, academic affairs at Liverpool and Oxford, and studies for the government on such problems as industrial fatigue. After his retirement from Oxford, his pursuits included lecturing and writing, trusteeship of the British Museum, and service as governor of the Ipswich School and Ipswich town adviser on museums and health services. In a career that spanned sixty-nine years, Sherrington is remembered mainly for his scientific contributions but he was also the teacher who prepared Mammalian Physiology. A Course of Practical Exercises (1919), the poet who wrote The Assaying of Brabantius (1925), and the philosopher and historian whose writings included Man on His Nature (1941) and The Endeavour of Jean Fernel (1946). His numerous honors included the presidency of the Royal Society of London (1920–1925), Knight Grand Cross of the British Empire (1922), Order of Merit (1924), and the Nobel Prize for physiology or medicine (1932); at his death he was an honorary fellow, member, or associate of more than forty academies and had received honorary degrees from twenty-two universities.

Sherrington’s classic investigations dealt primarily with reflex motor behavior in vertebrates, detailing the nature of muscle management at the spinal level. The data, terms, and concepts that he introduced have become such a fundamental part of the neuroscience that it is perhaps not surprising their authorship is often forgotten: such terms as proprioceptive, nociceptive, recruitment, fractionation, occulusion, myotatic, neuron pool, motoneuron, and synapse, and such concepts as the final common path, the motor unit, the neuron threshold, central excitatory and inhibityory states, proprioception, reciprocal innervation, and the integrative action of the nervous system.

Sherrington’s scientific work may be broadly divided into two phases: from the 1880’s to the publication of The Integrative Action of the Nervous System in 1906, and from 1906 to his receipt of the Nobel Prize in 1932. When he began his work, in the 1880’s the data and theories about the structure and function of the nervous system that had developed over the centuries were at best piecemeal. Controversy was rampant in almost every area and apart from some textbook presentations, few attempts had been made to correlate structural and functional data within a given field of study, much less to interrelate the various separate channels of work on the nervous system.

The study of reflex actions, for example, went on almost independently of work on such problems as the structure and interconnection of nerve cells, the differentiation of the sensory and motor functions of the spinal cord, and the determination of brain structure and function. A fairly extensive fund of techniques, data, and theories about reflexes was available, but the field of reflex physiology was greatly in need of reorganization. Techniques were generally imprecise; sounder anatomical knowledge was needed; above all, experimentally based concepts—with which to interpret the known facts of reflex action and evaluate their role in the animal economy—were singularly lacking.

Sherrington decided to concentrate on neuro-physiology rather than on pathology, his initial interest, when he returned to England in 1887 from Koch’s laboratory in Berlin. He credited Gaskell with directing his attention from his first neuro-physiological investigations into brain–spinal cord. Sherrington began by studying the little-under-stood phenomenon of the knee jerk, reporting the muscles and nerves upon which the jerk depends. In these studies, however, he found that he could not deal satisfactorily with functional problems in the face of a major gap in neuroanatomical knowledge—the distribution of the sensory and motor fibers of the spinal cord. For a decade of what seemed to him often “boring” and “pedestrain” research, Sherrington therefore surveyed the field of distribution of each spinal root, creating the anatomical foundation necessary for physiological work. His three major contributions to neuronatomy were mapping motor pathways, chiefly those in the lumbosacral plexus (1892), establishing the existence of sensory nerves in muscles (1894), and tracing the cutaneous distribution of the posterior spinal roots (1894, 1898).

Concomitant with his anatomical work, and often deriving from it, was a profusion of ideas and observations on the reflex functions of the spinal cord. The two major, intertwined lines of these researches were the analyses of antagonistic muscle action and of larger pieces of reflex “machinery” such as the extension, flexion, and scratch reflexes of the hind limb. Out of these studies emerged Sherrington’s conviction that the “main secret of nervous co-ordination. . .lies in the compounding of reflexes,” a compounding built up by the play of reflex arc about their “common paths.” Behind this play lie the key process of inhibitory and excitatory actions at the junctional regions between nerve cells—the synapses.

Like other investigators of the nervous system, Sherrington faced the task of devising techniques for reducing and controlling the complexity of the nervous system to the point where meaningful data could be obtained. His first steps were to concentrate upon the reflex functions of the cord rather than on the more complex field of the brain; to choose an appropriate experimental animal, the monkey; and to make parallel control and comparison experiments on lower forms to establish the necessary points of anatomical knowledge.

Sherrington’s basic method was to study simple motor acts which could be made to occur in isolation. correlating his exacting analyses of input-out-put relations of reflex responses with anatomical and histological data. He used two types of experimental preparations: the classic spinal animal and the decerebate animal. The effects of decerebration had been partially described by many earlier workers, such as Magendie, Bernard, and Flourens, but it was Sherrington who named decerebrate rigidity and, in a fundamental paper of 1898 and later publications, established it as a phenomenon in its own right and as a major tool for examining the reflex functions of the spinal cord, particularly the nature of inhibition.

The last decades of the nineteenth century saw the rise and fall of numerous theories about the nature of central inhibition, such as the controversial and influential center theory advanced by Johann Setchenov in 1863. It was against the background of these theories and the emergence of the neuron theory that Sherrington began to work out his ideas on the roles of inhibition and excitation in motor behavior and on the reflex nature of inhibition itself. The most important theme in Sherrington’s functional researches up to 1900, for both his understanding of the operation of spinal reflexes per se and his comprehension of the machanisms of nervous coordination, was his analysis of th reciprocal innervation of antagonistic muscles. It was the principle of reciprocal innervation, as Lord Adrian commented, “which opened the way to th further advance from the simple to the complex. It was the clue to the whole system of traffic control in the spinal cord and throughout the central pathways”1.

The results of Sherrington’s exhaustive study of reciprocal innervation, which stemmed from his observations on the knee jerk, are found chiefly in his fourteen classic “Notes” in the Proceedings of the Royal Society from 1893 to 1909. He first used the term “reciprocal innervation” in the title of the third “Note,” read before the Royal Society on 21 January 1897; the term, he explained, denoted the “particular form of correlation” in which one muscle of an antagonistic couple is relaxed as its mechanical opponent actively contracts. Four months later, as the Royal Society’s Croonian lecturer, he proposed his classic definition of reciprocal innervation as that form of coordination in which “in hibito-motor spinal reflexes occur quite habitually and concurrently with many of the excito-motor.”

Another critical event of 1897 was Sherrington’s introduction of the term and concept of synapse in Michael Foster’s Textbook of Physiology, “So far as our present knowledge goes we are led to think that the tip of a twig of the [axon’s] arborescence is not continuous with but merely in contact with the substance of th dendrite or cell body on which it impinges. Such a connection of one nerve-cell with another might be called a synapsis”.

Sherrington’s statement reflects the impact upon ideas of the structural and functional interrelations of the nervous system created by the neuron theory, introduced in 1889 by Ramón y Cajal. Prior to Ramón y Cajal’s researches the dominat neurohistological view was the reticular theory, which held that nerve impulses are transmitted throughout the body over a continuous network or reticulum of anastomosing nerve processes. RamÓn y Cajal’s preparations showed that definitely limited conduction paths exist in the gray matter and that nerve impulses are somehow transmitted by contact or contiguity, not by continuity. The significance of Sherrington’s choice of the neuron theory and his coining of synapse has been well stated by Ragnar Granit: “When Sherrington decided in favor of nerve-cell contacts he refashioned thinking in this field along lines that determined its future course for all time and also tied it to the newly born science of electrophysiology.. . .Only a contact theory could bridge the gap between reflex transmission and electrophysiology; such is the power of a fundamental concept like the synapse.”2

Between 1897 and 1900 Sherrington formulated a comprehensive picture of the motor functions of the spinal cord. His conception of these functions, of the rules that govern them, their mechanisms of control, and their role in the unitary functioningof the nervous system, were set forth in his Croonian and Marshall Hall lectures and in E. A. Schäfer’s Text Book of Physiology. By 1900 Sherrington had assembled the major ingredients of the integrative action concept. From a study of the seemingly simple anatomy and physiology of the knee jerk he had become engaged in a series of broader problems, such as the nature and mechanisms of antagonistic muscle action, the production and maintenance of decerebrate rigidity, and the nature and significance of spinal shock. From these and other researche he developed a number of basic functional principles; reciprocal innervation; interaction between higher and lower level centers of motor control; and muscular sense, inhibition, and facilitation as the three key mechanisms of muscle management at the spinal level. Recognizing the import of the neuron theory, he had perceived that many of the characteristic properties of reflex pathways might be explicable by the events at the synapse.

The next phase of his work was to determine how reflex arcs combine to form successively larger and more complex reflex patterns. Sherrington’s analysis of the scratch and other hind limb reflexes confirmed his earlier findings: the same functional principles obtain in both the simplest and most complex reflex actions. And, because of their very complexity, the hind limb reflexes further illuminated a wide range of phenomena underlying motor coordination, such as inhibition, facilitation, spinal induction, and the events at the synapse. In his definitive paper on the scratch reflex (“Observations on the Scratch Reflex in the Spinal Dog,” Journal of Physiology, 34 [1906], 1–50), the properties of reflexes as found in the isolated spinal cord were described more minutely and fully than ever before in the annals of neurophysiolosy.

In 1904 Sherrington enunciated the essentials of the integrative action concept to the Physiological Section of the British Association for the Advancement of Science in a presidential address entitled “The Correlation of Reflexes and the Principle of the Common Path.” It was his most important published conceptual statement before The Integrative Action of the Nervous System (1906). The main theme was the reflex chain of the synaptic system: the receptive neuron forms a private path into the brain or cord; within the “great central organ” many private paths converge at an internuncial neuron to form a public or common path which runs to the motor neuron; from the motor neuron, impulses travel over a final common path to converge upon the effecter organ. “The singleness of action from moment to moment,” assured by the principle of the common path, Sherrington declared, “is a keystone in the construction of the individual whose unity it is the specific office of the nervous system to perfect.”

The immediate fruitfulness of the common path principle can be seen in Sherrington’s papers of 1905 and 1906, in which he extended his analysis of the mechanisms controlling reflex actions. Working with the hind limb reflexes, he now focused on spinal induction, inhibition, and his fundamental concept of the proprioceptive system.

Sherrington journeyed to the United States to deliver ten lectures on “Integrative Action by the Nervous System” as the second Silliman Memorial Lecturer at Yale University in April 1904. His oral delivery, complex and difficult to follow even for those familiar with his work, left the majority of his steadily dwinding audience less than enthusiastic; but publication of the lectures in 1906 was recognized as an epochal event in the development of neurophysiology. The lasting value of The Integrative Action of the Nervous System for students of neurophysiology is reflected in the numerous reviews of its fifth reprinting, in 1947. F. M. R. Walshe, writing in the British Medical Journal, asserted:

I have called it “an imperishable work,” for it is one of those works, rare in science, the permanent value of which is unquestionable, and I believe that future generations of physiologists will so acclaim it. In physiology, it holds a position similar to that of Newton’s Principia in physics.. . .For it is more than an orderly record of precise observations: it is a product of sustained thought upon what is essentially—though only his genius revealed it as such—a single problem—namely, the mode of nervous actions.3

The Integrative Action consits, in essence, of a synthesis of Sherrington’s own researches and concepts and a chronicle of relevant work by other investigators. The structure and major concepts of the book may be divided into six parts: (1) Sherrington’s definition of his topic in the first seven and one-half pages of lecture I;(2) lectures I–III, treating of coordination in the simple reflex; (3) lectures IV–VI, concerned with coordination between reflexes—their interaction and compounding by simultaneous and successive combination; (4) lecture VII, reflexes as adapted reactions; (5) lectures VIII–IX, the brain’s role in integrative motor action; and (6) lectures X, sensual fusion. Sherrington’s written analysis of the integrative action of the nervous system followed basically the same pattern as his research work. He began by delineating the characteristics of the simple reflex, the smallest functional unit of integrative action as seen in the spinal animal, and then built toward the complex patterns of reflex muscle management, guided by the brain, in the intact animal.

For a person unfamiliar with Sherrington’s work, the first pages of lecture I provide a succinct statement of the meaning and scope of the concept of integrative action. In them he laid down three central propositions: (1) the nervous system is one, if not the only, major integrating agent in complex multicellular organisms; (2) the reflex is the unit reaction in nervous integration; (3) there are two grades of reflex coordination, that effected by the simple reflex and that effected by the simultaneous and successive combination of reflexes. Working from these premises he proceeded to demonstrate in meticulous detail the basic theme of the integrative action concept: “The nervous synthesis of an individual from what without it were a mere aggregation of commensal organs resolves itself into coordination by reflex action.”

Sherrington continued his active life as researcher, teacher, writer, and prominent member of the international scientific community in the years after 1906, moving from Liverpool to Oxford in 1913. When his research was curtailed by world War I, Sherrington devoted himself to government war work with his customary drive and efficiency. Few episodes better illustrate his character than his activities during the summer of 1915, when he disappeared from home on a bicycle, presumably for a holiday, leaving no address. His whereabouts were finally revealed when he needed to replace a lost collar stud—having decided to study industrial fatigue in situ, he was working incognito as an unskilled workman at the Vickers-Maxim shell factory in Birmingham.

At the end of the war Sherrington resumed his extensive research program, and until he was almost seventy-five performed at least one long experiment every week and spent many hours analyzing data. When he retired in 1935, the “Sherrington school” at Oxford had issued a series of influential papers on such topics as afterdischarge, summation, recruitment, postural contraction and the motor unit, to illuminate the finer details of the reflex activity of the spinal cord. Two of Sherrington’s last comprehensive reviews of muscle management at the spinal level, summarizing and synthesizing the work of the Oxford years, are his 1931 Hughlings Jackson lecture, “Quantitative Management of Contraction in Lowest Level Coordination,” and chapter seven of Reflex Activity of the Spinal Cord (1932). His coauthors for the later work exemplify the continuing prominence of the Sherrington school in modern neurophysiology: R. S. Creed, D. Denny-Brown, J. C. Eccles, and E. G. T. Liddell.

Sherrington’s tenure at Oxford may be characterized as a period of quantitation, testing and reining the concepts set forth in The Integrative Action. New techniques, in particular the development of isometric myography, made possible the accurate measurement of muscle tensions in various preparations. Sherrington could now measure and balance excitatory and inhibitory processes against each other, learning virtually everything about reflexes that was possible without the aid of the more sophisticated electronic methods that were developed as his career drew to a close.

Three fundamental publications of the Oxford period were Sherrington’s papers on the stretch reflex (1924), central excitatory and inhibitory states (1925), and the motor unit (1930). These papers presented both the culmination of issues raised by his earlier work and the basis for many subsequent major advances in unraveling the operactions of the nervous system. Sherrington and Liddell’s analysis of the stretch reflex, the basic reflex used in standing, grew out of Sherrington’s studies in the 1890’s on the response of the “muscular sense organs” to stretching and contraction. By 1905 he had observed and described the stretch reflex, although its naming and definition awaited his and Liddell’s later work. In their 1924 and 1925 papers, they reported the results of using in dividual isolated knee extensor muscles in a decerebrate preparation, with the free ends of the muscle attached to an isometric myograph. These researches, in turn, led to the definition of the nature of autogenetic excitation and inhibition and to our present understanding of muscle tonus, attitude, and posture.

Sherrington’s work on central excitatory and inhibitory states and the motor unit were the culmination, for him, of the research he had begun forty years earlier on the functional anatomy of sensory and motor pathways, and led to the studies of the finer details of synaptic conduction, which now fill volumes.

After 1906 one of the chief problems occupying Sherrington’s attention was inhibition. By 1925, in “Remarks on Some Aspects of Reflex Inhibition,” he was ready to state his concept of central excitatory and inhibitory states and, as he had first suggested in 1908, of excitation and inhibition interacting algebraically at the synapse. In the 1925 paper, Sherrington marshaled the evidence developed in over twenty-five years of experiments, reasoning back from the phenomena of muscle contraction, as seen principally in the crossed extensor and ipsilateral flexor reflexes, to the events at the synapse. Inhibition, he demonstrated, is a distinct phenomenon although it is almost identical in its properties to excitation and obeys the same laws. The further testing and development of his ideas on central excitation and inhibition occupied much of Sherrington’s time during his last decade or research. His concept of central excitatory and inhibitory states has been confirmed, expanded, and reformulated in terms of postsynaptic excitatory and inhibitory potentials by Sir John Eccles and others, using such techniques as intracellular recordings from motoneurons.

The concept of the motor unit, Sherrington’s last major contribution to neurophysiology, can be seen as a more sophisticated, experimentally based development of the principle of the common path. The motor unit, in simple terms, is a spinal motoneuron (or motor cell in the ventral horn of the spinal cord) which, by the branching of its axon, controls and coordinates the actions of more than 100 muscle fibers. Years of exacting study by Sherrington and his colleagues went into the paper on the motor unit which he published with Eccles in 1930: the writing of the paper itself occupied Sherrington for over two years. A glimpse of Sherrington the scientist, a seventy-two, working on his last major paper, is offered by John Fulton’s diary entry for 2 April 1930:4

It is remarkable to see how at this time of his life he is beginning to correlate all the various aspects of his own work; structure of fibers, reflexes, series of central excitations, etc., etc., into a beautifully synthesized body of knowledge. For him the days simply weren’t long enough, and there are all manner of things that need investigation. The rest of the world in his eyes is a little slow because it does not see all these things staring it in the face.

Sherrington received the Nobel Prize in 1932 specifically for his isolation and functional analysis of the motor unit. Somewhat ironically, the man who developed the single-unit concept never used the then newly available electrical recording techniques, but Sherrington can scarcely be held at fault for not making such studies when he was in his seventies. Sherrington shared the Nobel award with Edgar Douglas Adrian, Foulerton professor of physiology at Cambridge, for his analysis of the frequency discharge of single units. Both men had long sought to define the properties of the nerve cell as the functional unit of the Central nervous system—Adrian through investigating the afferent input from sense organs, Sherrington the nature of motor input—and Adrian’s work during the 1920’s probably had the most immediate influence on the direction and development of Sherrington’s ideas. To many, particularly those who had worked with him over the years, Sherrington’s award was long overdue—a circumstance perhaps partly explained by the Nobel Committee’s difficulties in citing a specific discovery on which to base the prize.

The work noted above is a small, but highly important, portion of the experiments and ideas generated by Sherrington and his pupils and colleagues during his twenty-three years at Oxford. It would be a difficult task indeed to select the most important single achievement from among the vast program of researches in which Sherrington was engaged from the 1880’s to his retirement in 1935: brain-cord connections and spinal degenerations; the distribution of motor and sensory roots; the proprioceptive system; the characteristics of synaptic reflex and conduction; reciprocal innervation and the nature of central inhibition; the reflex paterns of the spinal, decerebrate, and intact animal; and the nature of supraspinal control as seen in the functional organization of the motor cortex. Perhaps most significant are his inseparable analysis of reciprocal innervation and inhibition, his studies of muscle tonus (posture), and his conceptual definition of the nature of synaptic action in effecting the unitary or integrated behavior of nerve cells.

In more general terms, by examining the antecedents to, and tracing the course and content of, Sherrington’s researches, one sees how strikingly the new outweighed the old in his work: in instance after instance Sherrington himself “made the time ripe” for answering a given problem. He pioneered new techniques and apparatus, such as the method of successive degenerations, surgical procedures for mammalian decerebration, and the use of the myograph for reflex recordings, and established new methodological canons with his meticulously designed and executed experiments. Second, he marshaled extant facts and theories and added a host of new ones about each topic he studied, emphasizing particularly the correlation of structural and functional data.

The scope of his specific contribution clearly marks Sherrington as a major figure in the history of the neurosciences. The greater significance of his work, however, lies in his “synthetic attitude.” his perception of his interrelatedness of his varied researches. One of his goals was to explain the functional unity of motor behavior, primarily by interpreting central nervous system function in histological terms. From the content of his 1897 Croonian lecture on the mammalian spinal cord as an organ of reflex action, it appears that he was moving toward the concept of integrative action by that date.

Sherrington’s work is resolvable into a threefold study of reflex actions, using the nerve cell and its interconnection as his basic analytical unit: their gross and histological architecture, the spinal and higher level mechanism controlling them, and their functions in vertebrate motor behavior. Although his most fundamental were performed around the turn of the century, it was the work during the 1920’s at Oxford which has shaped many facets of neurophysiological research up to the present.

Sherrington’s name remains linked most closely, however, with his integrative action concept, although the idea of nervous integration did not originate with him. The fact of motor coordination and the participation of the nervous system in its operation had been recognized since antiquity, and prototupes of the integrative action concept may be found from the ancient idea of “sympathy” to Flourens’s studies of how specific brain regions affect an animal’s functional unity. The uniqueness and significance of Sherrington’s work, epitomized by the integrative action concept, lies in the fact that it provided the first comprehensive, experimentally documented explanation of how the nervous system, through the unit mechanism of reflex action, produces an integrated or coordinated motor organism. It was this watershed achievement, synthesizing the work of one era and opening a new one, that led Sherrington’s peers to designate him as “the main architect of the nervous systems,” “the author of the Principia of physiology,” and the “man who almost singlehandedly crystallized the special field of neurophysiology.”

Sherrington’s work was motivated, in large measure, by a desire to explain organized, purposeful behavior, and he looked upon reflexes as very simple items of such behavior. Thus, in the Silliman lectures and in subsequent writings he voiced distinct reservations about the ability of his analysis of reflex action to account for the functional solidarity of vertebrates, especially among primate forms. These reservations stemmed, in part, from the realization that his work left unanswered many questions about the nature of reflex motor control that would be resolved by others using more sophisticated techniques and types of experiments.

His other reservations about his work stemmed from his belief that reflex action was only a small part of the integration of higher vertebrates. Sherrington himself might have written the words of the sixteenth-century physician Jean Fernel, quoted in Man on His Nature: “. . .our task, now that we have dealt with the excellent structure of the body, cannot stop there, because man is a body and a mind together.” As the quotation suggests, this aspect of Sherrington’s self-appraisal is intimately related to his position on the question of the relation of mind and body. In this realm Sherrington was a dualist. For Sherrington man is the product of natural forces, yet he encompasses a territory which neurophysiology cannot reach—the realm of mind and thought. One senses, however, that Sherrington was not a dualist by philosophical choice but, rather, felt constrained to adopt the position because the sciences of his day offered no evidence or means of bridging the gap between mind and brain.

In the foreword to the 1947 reissue of The Integrative Action, Sherrington summarized his years of study and thought about the roles and relations of body and mind in animal integration, distinguishing three systems or levels of integrative action. At the first level, physicochemical processes weld the body’s organs into a “unified machine.” This welding is exemplified by the integrative action of the nervous system, of which the unit mechanism, as Sherrington had shown, is reflex action.

Reflex action vanished completely as a mechanism in Sherrington’s second system of integrative action, “the field of the psyche.” At this level, he held, “the physical creates from psychical data a percipient, thinking, and endeavouring individual.” For Sherrington “the physical is never anything but physical, or the psychical anything but psychical,” yet the two systems “are largely complemental and life brings them co-operatively together at innumerable points.” This mind-body liaison was, to Sherrington, the third and highest level of integrative action.“In all of those types of organisms in which the physical and psychical coexist, each of the two achieves its aim only by reason of a contact utile between them. And this liaison can rank as the final and supreme integration completing the individual.”

To Sherrington, called the “supreme philosopher of the nervous system,” the most baffling and challenging problem for both scientists and philosophers was how the mind-body liaison is effected. In 1947, he commented succinctly that the issue “remains where Aristotle left it 2,000 years ago.” While Sherrington could not explain a dualistic interaction, neither could he find any valid basis for reducing mind to a manifestation of physical energy. Thus, as he explained in the closing sentence of the foreword, dualism seemed to be as reasonable an assumpton as monism. “That our being should consist of two fundamental entities offers I suppose no greater inherent probability than that it should rest on one only.”

Near the end of his life Sherrington made one of his final and most positive statements about the levels of integrative action that produce the totality of an animal such as man. During a conversation with Sir Russell Brain, he said “the reflex was a very useful idea, but it has served its purpose. What the reflex does is so banal. You don’t think that what we are doing now is reflex, do you? No, no, no.”5

Although Sherrington’s dualistic philosophy was disturbing to many of his scientific colleagues, it did not diminish their estimate of his contributions to neurophysiology. Of the scores of tributes which marked his death on 4 March 1952, none more simply expressed the sentiments of those who knew him than the words of Henry Viets: “A great and good man has died....We stand on mighty shoulders.”6

NOTES

1. E. D. Adrian, “The Analysis of the Nervous System: Sherrington Memorial Lecture,” in Proceedings of the Royal Society of Medicine, 50 (1957), 993.

2. Ragnar Granit. Charles Scott Sherrington, p. 43.

3. F. M. R. Walshe, “A Foundation of Neurology. The Integrative Action of the Nervous System,” in British Medical Journal(1947), 2 . 823.

4. John F. Fulton Papers, Historical Library, Yale University School of Medicine.

5. Quoted in G. E. W. Wolstenholme and C. M. O’Connor, eds., Ciba Foundation Symposium on the Neurological Bases of Behaviour, in Commemoration of Sir Charles Scott Sherrington (Boston, 1948), p. 24.

6. Henry Viets, “Charles Scott Sherrington, 1857–1952,” in New England Journal of Medicine, 246 (1952), 981.

BIBLIOGRAPHY

I. Original Works. A complete Sherrington bibliography is in Fulton (1952); an extensive although not complete listing is in Swazey (1969). Both are cited below. For those wishing to explore the development of Sherrington’s scientific ideas and his nonscientific works, the following are suggested: The Central Nervous System, vol. III of Michael Foster, A Textbook of Physiology, 7th ed. (London, 1897); “The Mammalian Spinal Cord as an Organ of Reflex Action. Croonian Lecture,” Proceedings of the Royal Society, 61 (1897) 220–221, an abstract that was printed in full as sec. 4 of Sherrington’s “Experiments in Examination of the Peripheral Distribution of the Fibres of the Posterior Roots of Some Spinal Nerves (II),” in Philosophical Transactions of the Royal Society, 190B (1898), 45–186: “Decerebrate Rigidity, and Reflex Co-ordination of Movements,” in Journal of Physiology, 22 (1898), 319–332; “On the Spinal Animal (The Marshall Hall Lecture),” in Medico-Chirurgical Transactions, 82 (1899), 449–477; “The Spinal Cord,” “The Parts of the Brain Below the Cerebral Cortex,” “Cutaneous Sensations,” and “The Muscular Sense,” in E. A. Schäfer, ed., Text Book of Physiology. II (Edinburgh. 1900). 783–1025: “The Correlation of Reflexes and the Principle of the Common Path,” in Report of the British Association for the Advancement of Science, 74 (1904), 1–14; The Integrative Action of the Nervous System (New Haven, 1906); “Reflex Inhibition as a Factor in the Co-ordination of Movements and Postures,” in Quarterly Journal of Experimental Physiology, 6 (1913), 251–310; “Some Aspects of Animal Mechanism. Presidential Address, British Association for the Advancement of Science,” in Report of the British Association for the Advancement of Science (1922), 1–15; “Reflexes in Response to Stretch (Myotatic Reflexes),” in Proceedings of the Royal Society. 86B (1924), 212–242, written with E. G. T. Liddell: The Assaying of Brabantius and Other Verses (Oxford, 1925): “Remarks on Some Aspects of Reflex Inhibition,” in Proceedings of the Royal Society, 97B (1925), 519–545; “Numbers and Contraction-values of Individual Motor-units Examined in Some Muscles of the Limb,” ibid., 106B (1930), 326–357, written with J. C. Eccles; Reflex Activity of the Spinal Cord (Oxford, 1932), written with R. S. Creed et al.; Inhibition as a Co-ordinative Factor (Stockholm, 1932), the Nobel lecture delivered at Stockholm, 12 Dec. 1932: The Brain and Its Mechanism (Cambridge, 1933): Man on His Nature (Cambridge, 1941), the Gifford lectures, Edinburgh, 1937–1938; The Endeavour of Jean Fernel (Cambridge, 1946); and “Marginalia,” in E. A. Underwood, ed., Science, Medicine and History, II (Oxford, 1954), 545–553.

II. Secondary Literature. See Edgar D. Adrian. “The Analysis of the Nervous System: Sherrington Memorial Lecture,” in Proceedings of the Royal Society of Medicine, 50 (1957), 991–998; Mary Brazier, “The Historical Development of Neurophysiology,” in John Field, H. W. Magoun, and V. E. Hall, eds., Handbook of Physiology. Section I: Neurophysiology (Washington, D.C., 1959): Georges Canguilhem, La formation du concept de réflexe au XVIIe et XVIIIe siècles (Paris, 1955); Lord Cohen of Birkenhead, Sherrington: Physiologist, Philosopher, Poet (Liverpool, 1958), vol. IV iologist, Philosopher, Poet (Liverpool, 1958), vol. IV of the University of Liverpool Sherrington Lectures; Derek Denny-Brown, “The Sherrington School of Physiology,” in Journal of Neurophysiology, 20 (1957), 543–548; Franklin Fearing, Reflex Action: A Study in the History of Physiological Psychology (Baltimore, 1930: repr. New York, 1964): John F. Fulton, “Sir Charles Scott Sherrington, O. M.,” in Journal of Neurophysiology. 15 (1952), 167–190: and “Historical Reflections on the Backgrounds of Neurophysiology: Inhibition, Excitation, and Integration of Activity,” in Chandler M. Brooks and P. F. Cranefield, eds., The Historical Development of Physiological Thought (New York, 1959): Ragnar Granit, Charles Scott Sherrington: An Appraisal (London, 1966): E. G. T. Liddell, “Charles Scott Sherrington, 1857–1952.” in Obituary Notices of Fellows of the Royal Society of London, 8 (1952), 241–259; and The Discovery of Reflexes (Oxford, 1960); Wilder Penfield, “Sir Charles Sherrington, Poet and Philosopher,” in Brain, 80 (1957), 402–410; Carr E. R. Sherrington, Memories (privately printed, 1957), the Beaumont lecture, Yale University, 15 Nov. 1957: Judith P. Swazey, “Sherrington’s Concept of Integrative Action,” in Journal of the History of Biology, 1 (1968), 57–89; and Reflexes and Motor Integration: Sherrington’s Concept of Integrative Action (Cambridge, Mass., 1969).

A major source of information about Sherrington and his life and work is contained in the papers of the late Dr. John F. Fulton, who had a long and close personal and professional association with him. Fultion first studied under Sherrington in 1921, when he went to Oxford as a Rhodes Scholar. The Fulton papers are housed at the Historical Library, Yale University School of Medicine, New Haven, Connecticut.

Judith P. Swazey

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