Charles Sherrington

Charles Scott Sherrington.
1857-1952
Author(s): E. G. T. Liddell
Source: Obituary Notices of Fellows of the Royal Society, Vol. 8, No. 21 (Nov., 1952), pp.
241-270
Published by: Royal Society
Stable URL: https://www.jstor.org/stable/768811
Accessed: 02-03-2019 18:22 UTC
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CHARLES SCOTT SHERRINGTON
1857-1952
CHARLES SCOTT SHERRINGTON was born on 27 November 1857 in Islington,

where his mother was on a visit, and died at Eastbourne on 4 March 1952. For
fifty-six years he was a considerable and influential contributor to knowledge of
nervous reflex action and muscle management. His father was James Norton
Sherrington of Caister, Great Yarmouth, who died when his children were
small, there being two younger sons, both destined for the Law. After her first
husband's death the widow married Dr Caleb Rose of Ipswich. His stepfather's
influence was to extend all through Charles Sherrington's life, for he was a good
classical scholar and an archaeologist of note. The home at Edgehill House was
a rendezvous for artists, the rooms being crammed with the works of David
Cox, Cotman and others of the Norwich School. Many of them now hang in
the Norwich Gallery. In later life, Charles was deeply appreciative of Vermeer.
His interest in medicine doubtless came from his stepfather, and his mother
made the home a true home in every aspect. As a schoolboy and in early life,
Sherrington was an athlete of no mean order, playing soccer for his school,
Ipswich Grammar School, to which he went in 1871. He played soccer for
Ipswich Town, and in due time rugger for St Thomas's Hospital and for
Caius, and also rowed for his college. His vigorous game in the Caius scrum was
long remembered by his contemporaries and he was made to admit, in after
years, that he ought to have played half-back if only his eyesight had been
better. At St Thomas's, 'We used to play football, rugby it was, on the
Archbishop's ground near the hospital, but it was very wet and rather
dangerous'. Later on and until his marriage he was a pioneer of winter sports
at Grindelwald.
At Ipswich School, Thomas Ashe was a great inspiration to Sherrington and
gave him a love of classics and the urge to travel. The headmaster at the time
was H. A. Holden, editor of Aristophanes. Sherrington passed his preliminary
examination in General Education at the Royal College of Surgeons of England
in June 1875. This was the preliminary examination for the Fellowship and
gave exemption from the similar examination for the Membership. In later life,
he was to say of the early days of his education: 'Science was socially not quite
the thing'. In September 1876 he entered as a 'perpetual pupil' at St Thomas's
Hospital, London, passing his Primary Examination for the Membership of
the Royal College of Surgeons in April 1878, and the Primary for the Fellowship
twelve months later. 'When I was a young man, the library of the Royal College
Q 241
6< -
242 Obituary Notices
of Surgeons was the only place in London wher
librarian didn't come forward and ask what he could
time at Edinburgh about which there is but little i
went to Cambridge as a non-collegiate student in
following year migrated to Gonville and Caius. By
were on the mend after a disastrous bank failure som
The move to Cambridge was to be a momentous c
when much was afoot in England in the field of ph
medicine. The German universities were spreading
Trinity, for instance, in 1870 had appointed Michael
of talent, as Praelector in Physiology.
In 1881 a medical congress took place in London
influence Sherrington and bring him to the beginning
research. It was the Seventh International Medical
was in the Chair of the Physiology Section and quo
Bell: 'Each muscle is connected with the brain by
fibres'. Goltz of Strasbourg had turned by then fr
frogs to the same sort of work in dogs. He demonst
had made cortical excisions and which seemed to him to
He maintained that localized function in the cortex
younger than Goltz by nine years, proceeded to de
one with hemiplegia after a cerebral lesion of seven
was the occasion which evoked from Charcot th
'C'est un malade'. The other monkey had had ex
superior temporo-sphenoidal convolution on both s
It had no paralysis but was deaf even to the exp
Ferrier argued strongly for localization of functio
'I reject his conclusions'. Gerald Yeo, F.R.C.S., Pr
King's College, and previously Assistant Surgeon, w
for Ferrier, said he thought that Goltz's dog did sho
mentioned rather pointedly the need for 'aseptic' sur
In this highly charged atmosphere, escape from dea
then by chloroforming the dog and the hemiplegic
committee composed of Gowers, Klein, Schafer and
material. Sherrington is not recorded as a member
Congress. The right hemisphere of the dog was taken
who had been a Fellow of Trinity since 1877, and w
papers, and was giving the Advanced Lectures for Fo
tology. And to the histological examination of that h
subsequently brought in as a junior colleague after
papers on it. They reported together in 1884. It was
Thus it was that young Sherrington began neurolog
at the time many questions, and few answers, but G
that experimental method might help to provide answ
was definitely on the move. The Physiological Socie
Charles Scott Sherrington 243
vigorous youngster whose history was later well set forth by Sharpey
with first-hand knowledge. The Neurological Society of London was h
lively meetings which were recorded in the early volumes of Brain. Fe
writing thus: ' . . . anatomy, experimental physiology and clinical me
ought mutually to support and shed light on each other . . .' This que
cortical localization was enduringly to the fore, both motor and sensory
wrote, 'The problem which attracts the attention of all classes of neu
whether clinical, experimental or anatomical is the identification of
with certain spots in the brain'. He wrote, too, of the 'so-called' motor
the 'sensori-motor area', while Bastian for the same conception introd
word 'kinaesthetic'. The nature of the recently discovered knee-jerk w
debated. Was it a true reflex with its short latency? Or was it just a ph
concerned with the alertness of live muscle? Did its disappearance in
depend on lesion of the posterior spinal column or on lesions of the
roots? Then there was the problem of muscle sensibility which evok
interest and produced writings from men as far apart intellectually a
the German physiologist and psychologist, and G. H. Lewes, author,
and part-time physiologist. Was there really sensibility in muscle or
indirectly a part of cutaneous sensibility? Hence the young Sherring
thoughts turning towards neurology readily became a junior part
J. N. Langley in examining nervous material supplied by Goltz.
During these years research was one of many occupations. In 1883 Pr
(later, Sir George) Humphry made Sherrington a Demonstrator in the
Department. Writing later of this appointment, Humphry used word
mutatis mutandis, ring true of Sherrington at any age: 'being much i
by his ability and assiduity. He did his work well and the students we
attached to him. His genial friendly qualities made him a general favo
a most agreeable colleague.' His name reappears in the Pupil's Entry B
St Thomas's for the winter session of 1883-1884, when he demonstrated
histology there. The M.R.C.S. was obtained on 4 August 1884, and a First
Class in the Natural Science Tripos Part II with the mark of distinction in 1885.
His first own paper as sole author appeared in 1885, again on the topic of Goltz's
dogs. In it occurs a striking neologism of the kind which was to become so
characteristic of Sherrington's writing in later years. lie suggested the term
'cord area' of the cerebral cortex rather than 'motor-area'-which, had it been
accepted into general use, would have given a richer and surer conception of
brain and cord relationship. In the winter of 1884-1885 Sherrington went to
work with Goltz in Strasbourg. In later years Sherrington said of him, 'He
taught one that in all things only the best is good enough'. And Goltz wrote of
Sherrington ten years after that first visit: 'habe ich in ihm einen ebenso gliick-
lichen Beobachter als iiberaus geschickten Experimentator schAtzen gelernt'.
It was in Strasbourg, then at a time of high political feeling, that Sherrington
saw and never forgot the arrogance of German militarism but experienced
nevertheless some lingering spirit of academic freedom among German scholars
who had been translated there after the Treaty of Frankfurt in 1872. In 1885
he obtained the degree of M.B., Cambridge, an
following year. In the summer of 1885, Sherrington
Brown were deputed by the Association for Rese
junction with the Royal Society and the Universi
Spain to investigate an outbreak of cholera. In 18
Venetia and Puglia, again investigating cholera and
painting and Italian incunabula. The morbid materi
was examined in Berlin under the eye of Virchow
turning to politics as an opponent of Bismarck. His
taken to the Reichstag to hear Bismarck speak. Whe
been in progress for some time, a door above a sm
wall opened and Bismarck appeared (like a cuckoo
berate the assembly with a voluble oration, withdrawin
as he had emerged. Later on Sherrington was sent b
for a six weeks' course in technique but he stayed wi
research until the following year, and was almost p
in the Indian Medical Service, but thought that his
though in fact it was not. While in Berlin, Sherrin
lectures and to Zuntz for experimental physiology.
pupilage were ended in 1887 by Sherrington's appoin
as Lecturer in Systematic Physiology at St Thomas'
as being elected a Fellow of Gonville and Caius Colle
Upon young Sherrington's development there wer
considerable influence. The first was W. H. Gaskell,
laboratory. Of him, in later years, Sherrington wrot
OXFORD,
23. xi. 1918.
DEAR HEAD,-SO busied have I been that I could not get to your n
about Gaskell. He was always an inspiration to me and to any work
able to try. Such inspiration is often subtle and part of its success sp
I imagine, from its subtlety. One does not like to be driven, but inspi
is not driving. My own work began by chance at the wrong en
cortex-pyramidal degenerations, etc. It was certainly through Gaskel
I very soon felt that. One could not talk with him long without re
that the cord offered a better point of attack physiologically. Also, th
cord was originally and still must essentially be a chain of 'ganglia',
would say, thinking of invertebrates. The spinal afferent root should
the limits of, and at least part of the make-up of the metamerism-functi
as well as morphological. But when one got on with that, came the stumb
block, i.e. that in the vertebrate the segments, as judged by the
afferents, had overflowed into one another. But at the same tim
showed the visceral segmentation and how it had persisted in purer f
I remember how pleased he and I were about your observation
results. IHe had expected it would be something like that. He had al
insisted to me that the bulb-segmentation was more primitive th
spinal. But the bulb lent itself less to laboratory experiment.
Later, too, when his own chief interest had passed to the or
vertebrates, he was still always a bulwark to me about inhibi
voluntary muscle. To both him and me it always seemed that the
vol. muscle was impossible without inhibition. But we both ex
efferent inhibitory nerves to them. You remember his vivid inte
Bierdermann and the Astacus claw. I used to search for similar th
mammalian muscle-nerve. I used to put down-he agreeing-each
to a swamping of the inhibitory fibres by excitatory mixed in th
efferent nerve. Gradually, in view of one's reflex experiments, i
upon me that for the vol. muscles, the inhibitories play not on th
direct but on the spinal motor cells driving them, and in that sens
afferent or central. When I told him how things were pointing he
of his own work, though up to the neck in it then, and went into m
and all its pros and cons, not stinting time for it. Finally, he said,
course, here again the somatic part of the segment is less primitive
muscle's inhibition is peripheral, but skeletal muscle's inhibit
become central! That gift of sympathetic attention, and unselfish s
off from his own problem to a pupil's, was characteristic of his n
And his transcendant sincerity lent such force to his criticism or a
He personified truth. In a hundred ways I owe him help and inspir
I wish that, when this war is over, some more adequate account
and his influence may appear; no proper appreciation of him h
given yet.
With kindest remembrances to you both,
Yours ever,
C. S. SHERRINGTON.
The second person to influence Sherrington's thought was the Spanish

histologist Ram6n y Cajal. The two had met in Spain during Sherrington's
visit in the summer of 1885. As Sherrington's obituary notice of Cajal bears
witness (Obituary Notices of Fellows of the Royal Society, 1932-1935, 1, 425),
it is to Cajal and not to Waldeyer that credit is due for establishing the separate-
ness of identity and existence of nerve cells and for putting forward the 'Ley
del la polarizacion dinamica de la celula nerviosa' to explain interneuronal
transmission (1889). Cajal's views, however, in the middle eighties, were still
to come. Gaskell's influence, near at hand, is seen in Sherrington's papers on
outlying cells of the spinal cord. He attempted to find in mammalia the existence
of ganglion cells at the periphery of the antero-lateral column of the spinal
cord, as Gaskell had found in the alligator. They exist there, and also in the
posterior column. Sherrington's last scientific paper (1940) returned to the
problem of antero-lateral border cells.
In the St Thomas's Hospital Report of 1891 (21, 145) is a very short paper on
the knee-jerk which shows how Sherrington's mind had turned to the action of
spinal reflexes. His question was the question of th
was a true reflex or a direct muscular reaction d
Sherrington pointed out that even if the knee-jerk
because its speed was so high, it nevertheless depend
integrity of the reflex arc. Sensory fibres from the m
of the nerves from antagonistic muscles enhances th
of the posterior spinal roots into which they run. O
in the knee extensor is abolished by stimulation of
knee flexors. Here then, in miniature, is an early pictu
of reflex inhibition. Several papers on this same to
inhibition was thus clarified as a definitely rapid and
in complete inactivity of the antagonistic muscle.
knee-jerk could be brought about not only by the artif
of a nerve of a knee flexor but by more natural pro
knee flexor or gently compressing it between the f
degree of tension in one muscle of an antagonistic c
degree of "tonus" in its opponent, not only mechan
through afferent and efferent channels and the spinal
and difference between this inhibition and that fo
claw of Astacus was yet to seek.
Very soon in the problem of the knee-jerk, because
parts of a muscle are supplied by what nerve roots
question of nerve-supply. It had been supposed tha
had a definite functional expression in one muscle.
muscle contraction might result from activity of one s
ton showed that excitation of a spinal root determin
supplies only a partial contraction. Several roots tog
muscle, and a single motor root supplies parts of sev
may actually be antagonistic. There is no connexion
lay-out of spinal roots and a purposive muscular ac
myotome has lost its wholeness and become div
distributed among neighbouring muscles. When th
root stimulation its contraction is scattered in sep
purposeful function.
In the period 1892-1894 several long systematic p
on this important question of the efferent nerve su
the general outlook was entirely altered and the m
A weighty practical point was the discovery of two
'prefixed' and 'postfixed' plexuses in which anatom
muscles lie either further headward in the one class or further hindward in the
other class. There are two maxima, but apparently no normal or mean. More-
over, 'the commingling of the motor fibres to various muscles is great even at
their very exit from the cord'. All these results were obtained for efferent roots
by their careful electrical stimulation in the spinal canal and observation of
muscles' contractions-anatomical facts elicited by experimental methods.
The exploration of sensory root distribution came later, in the pe
1897, and charted segmented skin fields as never before. These resu
obtained from animals in which it is not easy to design experiments for e
sensation, but the basic idea of overlapping segmented skin areas w
seed itself in clinical fields. 'Each bunch of many sensory nerve tru
limb consists of fibres which enter the cord by two or three distinc
roots.' 'So also a great interlapping is evident in the distribution of
fibres from the skin.' Each root, however, possesses a single undivide
cutaneous distribution, not a field composed of separated patches, a
previously supposed. At the end of one of these papers Sherring
'I recognize that the problem here attempted has been one imperfectl
to inquiry upon animals. It awaits further solution in the opportunitie
by human disease where the attention of an instructed human subje
enlisted and interrogated verbally.' In later life, Sherrington was kno
that this early work on spinal roots and the innervation of muscles
and boring but he had to do it because anatomical knowledge of the
so inaccurate and useless.
At this time further anatomical studies on the nerve supply of skeletal muscles
brought to light the very important discovery that about one-third of the fibres
in a muscular nerve are afferent, coming especially from Kiihne muscle-spindle
but also from Golgi tendon organs. Only the remaining two-thirds of fibres i
the nerve are motor fibres. This work threw entirely new light on the nerv
supply of the skeletal muscles.
Thus by 1895, when Sherrington left London for the Holt Chair o
Physiology at Liverpool, he had turned his main attention upon two problems.
The first was seen at once to be of prime importance and was the anatomy o
the pathways to and from the spinal cord. The second was the opening study o
muscle management which was in Sherrington's hands to continue so fruitfull
for so long, the functional behaviour of the knee-jerk which depended upon th
reaction of the spinal nerves and the spinal cord. But even in these early paper
on the knee-jerk, the reflex play of one muscle upon another was brought int
view. Sherrington thus followed, as he pointed out himself, in the steps of
Charles and John Bell (The anatomy and physiology of the human body, London
1826). They wrote, 'The nerves have been considered so generally as instru-
ments for stimulating the muscles without thought of their acting in the opposite
capacity, that some additional illustration may be necessary here. Through the
nerves is established the connection between the muscles . . . but also that
relation between classes of muscles by which the one relaxes and the other
contracts. I [sic] appended a weight to a tendon of an extensor muscle, which
gently stretched it and drew out the muscle, and I [sic] found that the contraction
of the opponent flexor was attended with the descent of the weight, which
indicated the relaxation of the extensor.' As the Bells had found, Sherrington
found too that reciprocal contractions of eye-muscles provided an analogous
field of study. Direct experiments with the eye-muscles themselves, because of
difficulties of access, were, however, less rewarding than observations of eyeball
movement on stimulation of parts of the brain surf
activity by such stimulation was clearly establish
innervation' dates from this time, which was January
In 1897, Foster's Textbook appeared in its 7th editio
assistant author. In that edition the new word 'synap
time and was later credited to Foster. It became a classical
and is described in these words: 'So far as our presen
led to think that the tip of a twig of the arborescence is
merely in contact with the dendrite or cell-body on w
Between 1885 and 1895 he had several times visit
Strasbourg, whence had emanated papers on the reac
his Croonian Lecture, Sherrington followed up Goltz's
dogs but described shock in monkeys and cats and var
that the temporary paralyses in shock are Hemmung
Ausfallserscheinungen. The headward end of the ner
shock although sharing in the lowered blood pressure.
in the monkey than in other laboratory types and mor
that the greater effect in the monkey must be related
complexity in the monkey, so that spinal section lea
defect of nerve activity. For each individual creatur
thus separated into two parts, the part that is 'me' and
'I think it was Lotze who said that doubtless to the tro
halves the trodden "me" shall surely appear the g
transection the splendid motor machinery of the ver
whole and at one stroke severed from all the univers
the "material me".'
Segmentation of the cord was a continuing source of labour and an important
'rule' was defined in the course of the Lecture. 'In response to excitation, even
approximately minimal of a single afferent root, or even of a single filament of
a single efferent root, the spinal discharge of centrifugal impulses evoked tends
to occur via more than one efferent root, i.e. is pluri-segmental.' 'I look upon
the solidarity of the limb as structurally expressed by the pluri-segmental
character of the motor nuclei.' Thus was expressed a further addition to modern
conception of spinal structure and function.
After settling in at Liverpool in 1895 Sherrington took up with renewed
vigour the problem of the reciprocal innervation of antagonistic muscles.
There was no shortage of questions-nor ways of attack. His papers at this time
were especially numerous and versatile. The retina as well as the extrinsic eye
muscles were laid under contribution. Reflex inhibition was shown to be an
important part in reciprocal innervation. By this process excitation is not just
stopped or made absent. It can be graded, and is inversely proportional to the
excitation of the opposing group of muscles. Without reflex grading of con-
traction, muscle management would not be a coordinated process. 'Desistence
from action may be as truly active as is the taking of action.' Work continued on
inhibition and excitation and reached a point in 1913 when Sherrington could
say that 'the process of E and I may be viewed as polar opposites . . .
is able to neutralize the other'. An inference from such corresponden
well be that the two processes consist essentially of one reaction which
ible in direction. So in about twenty-five years had developed one of
important parts of Sherrington's contributions to knowledge of the sp
arising from the directing suggestion of Gaskell to study the spinal
from the pregnant conception of Cajal that separated neurones each co
cated impulses to the next neurone across the synaptic space. Nerve p
the spinal cord and out of it had been mapped accurately. Earlier wor
Charles Bell and Marshall Hall, and later workers like Goltz and Pflui
laid bare foundations, but the latter two, in Sherrington's time, and
experiments, were to be shown to be seriously and patently wrong.
During all the years that reciprocal action of muscles was holding his atte
Sherrington had other lines of work. The connexion between the brain
spinal cord by the pyramidal tract was still full of interest and was
From the beginning Sherrington had followed Ferrier and incline
higher animals rather than cats and dogs, because of the nearer-to-hum
of brain in anthropoids. This interest culminated in the experiments
colleague A. S. F. Griinbaum, published in 1901-1903 and 1917. Not on
the brain of the ape a higher type, it was richer in nerve cells and r
to electrical stimulation more readily and more surely than the lowe
The cortical map of 'motor points' became well known and was widel
duced for many years, but all too often without the authors' hints of d
warnings against too dogmatic acceptance of a map of points, too perm
rigid. The method necessarily employed, namely electrical stimulation
tissue which has been anaesthetized, was indeed a method of contradi
that the answers forthcoming were not to be assayed at their barest f
There was so much that might be wrong with normal function impa
put, as it were, on a knife edge between somnolence and spurred activ
too, Sherrington did not forget that the area might more safely have b
a 'cord area' and that the nerve path from this 'cord area' of the brain
to the cord itself was but an internuncial path between the 'will-
(whatever that may be) and the muscle which was made to move.
While these experiments were in progress at Liverpool, a young Am
surgeon paid a visit. This was Harvey Cushing (later For.Mem.R
prolonged his stay to eight months. It was the beginning of a lifelong frie
As a recent biographer of Cushing has written of that time: 'Sherring
surprise to Cushing. He found him younger than he expected, almost
wearing, when he had not lost them, a pair of gold spectacles. He
Sherrington operated well for a physiologist, but too often, wrote and
too much, went at things too rapidly.' That picture, however, pa
biographer as well as his subject. (Harvey Cushing. E. H. Thomson. Sc
New York, 1950.)
These papers on cortical stimulation by Griinbaum and Sherrington
almost to a standstill the controversy about the post-central gyrus c
motor nerve cells. Horsley said that they 'supers
Beevor. From the results of Sherrington and Griinb
main the post-central gyrus was not a motor area.
in after years was not dogmatic on this subject. It was
had not found it to be so outspokenly motivating as
had found conditions of rebound activity in which i
In 1904, among the usual generous flow of various
tion of the 'final common path' which was soon
neurophysiology. 'The final or efferent, is, so to say
impulses arising at any of many sources.' On the co
'the receptive neurone forms a private path exclus
source only'. 'The final terminal path may, to distin
common paths, be called the final common path.' Th
later into the concept of the 'motor unit', i.e. one spina
as all in one team, by branchings of its axone, a larg
which act together in all respects.
One of the most colourful chapters of Sherringto
century was the scratch-reflex-a perfected fulf
conception (1850) of the spinal cord as a centre of
mediate nerve cells of the spinal cord had long bee
of some of them was shown by Sherrington in 190
paper on the scratch reflex in the spinal dog. By car
aspects of reflex muscle control were found and desc
of observation and description, Sherrington seldom
than in this paper. 'Electric tickling', as he was to cal
artificial flea on a tiny point of skin, brought into
muscles beating rhythmically five times a second, a
in steady postural action. The main characteristic of
neuronal synapses could all be depicted inspiringly
scratch reflex: summation of stimuli, facilitation
refractory phase, after-discharge, inhibitability, the
post-inhibitory rebound.
At the turn of the century Sherrington had been b
buting to the large textbook of physiology of his fr
part was the authorship of chapters on the spinal co
cerebellum, corpora quadrigemina, thalamus, cutaneo
They were full of recondite information and are stil
In them, and repeated verbatim in the Silliman Lectu
strong protest against the hopelessly inaccurate Laws
reflexes, which had been argued by their author aft
reports of clinical cases. In the later years of his lif
had to spend an undue amount of time in refuti
obtained doctrinal importance and were as erron
accepted. So recently as 1891 they had had a measur
and Horsley in their Croonian ILecture to the Roya
After the work on the scratch reflex (1906) came several full exposit
discussions of the role of inhibition in reciprocal activity, and its impo
grading the strength of muscular contraction, and the avoidance of w
energy. Spinal inhibition is certainly central, and acts apparently at o
the origin of the final common path to the muscle. The characters of
forms of inhibition in the heart, intestine, blood vessels, and in lower
were described and compared. Once again was viewed inhibition of
muscle by stimulation of the cortex cerebri, which was, after all, th
ganglion of the nervous system. And once again the similarities betwee
tion and excitation are discussed. 'Two delicately gradable antithetic p
or states.'
One of Sherrington's longest studies was concerned with the tonus of muscle.
'Tonus! What is tonus? Who can say what tonus is?' is a remark attributed to
Mosso and expresses the opinion of the time. Sherrington's study was based on
his phenomenon of 'decerebrate rigidity', and began in the following manner.
After Ferrier's pioneer stimulation of the brain surface in the seventies, a
number of workers from time to time had extended his observations. In this
country, both Horsley and Sherrington were almost on the same trail, and
almost too close to one another for mutual comfort and satisfaction. Besides
stimulating the brain surface with electric currents, they also stimulated the cut
surface of the brain stem. In such preparations, with the brain stem cut across,
Sherrington noticed a peculiar 'cataleptoid' state.
In 1897 he read a paper to the Royal Society on this phenomenon in monkeys.
He had found that a limb, after a quasi-harmful ('nociceptive') stimulus, assumed
a position of withdrawal which was steadily maintained for many minutes and
was unlike any response seen after stimulation of the cortex cerebri, or in the
spinal state. He named for the first time this state of 'decerebrate rigidity' in
antigravity muscles. Horsley (with Loewenthal) described the same phenomenon
in the following month under the name of 'acerebral tonus'. Sherrington's
account was, however, clearer and more readable. Horsley and Loewenthal
gave mainly their protocols. From time to time in the next thirty years Sherring-
ton was often to advance knowledge in this branch of physiology. His deductions,
founded on circumstantial evidence, established a firm knowledge of muscle
tonus which even now by modern methods is being only amplified without
much alteration.
Very early in his research it became clear that this increased tonus of anti-
gravity muscles depends on the integrity of afferent pathways from the limbs.
The 'adequate' stimulus is stretch of the muscle itself. Proprioceptive endings
in muscle are 'in large measure' responsible for the reflex posture. This modifying
clause was needed because contemporary opinion had not yet realized that the
continued low-grade activity of muscle which characterizes tonus could have
several sources. The cerebrum, the cerebellum and the labyr nth were already
recognized as having an influence on tone. What was really new was Sherring-
ton's conception of this plastic tonus of an extensor having an autogenous
origin, coming from the muscle itself. The muscle proprioceptors and their
arcs were an entity which had to be treated as one sy
came fuller examination of the resistance exerted b
is made to pull it out longer. There is a resistance w
and the muscle then lengthens. After much experim
comparing this decline in tension with reflex inhib
stimulation, Sherrington came to the conclusion tha
as he called it, was due to inhibition arising from wi
must be receptor organs in muscle which produced
tendon tension was high. In the 'lengthening react
thought about, the muscles gave way and released i
when inhibited by electrical stimulation of an inhibi
was often contraction of the corresponding mu
('crossed extensor response') which seemed good evi
tion. Autogenetic inhibition, he thus felt certain, ex
just as autogenetic excitation. Tonic muscle, observe
preparation, had also a shortening reaction, which in
of the lengthening reaction. When the leg is lifted u
so as to allow the knee extensor muscle to shorten,
knee just to 'stay put'. The now shortened knee exte
length. By reason of the lengthening and the shor
reflex contraction is in this sense plastic; hence the
than one paper, the tonus of skeletal muscle was co
smooth muscle in the gut or bladder and with the
forms of animal life.
As early as 1905 another aspect of decerebrate tonus had been noticed. When
the hind leg of a decerebrated cat, which is not too rigid, is allowed to flex at
the knee in response to gravity so as to extend quickly the knee extensor muscle
'the anti-crus . . . is seen to be suddenly checked by exciting a contraction of
the extensor of the knee. This contraction is different from a knee-jerk, for it
only slowly passes off.' And in 1907: 'The impression given is that of a contrac-
tion reflexly evoked in the extensors of the knee.' The stretch reflex was thus,
in fact, observed and described but not yet identified and named.
Years later, in 1924, at the end of the long research on 'tonus', came a descrip-
tion of the reflex development in muscle of considerable tension which was
called 'the stretch reflex'. The effect of initial tension on the reflex response of
muscle was being observed, when one day a small accidental increase in muscle
length elicited a large response of active reflex tension. That was the stretch
reflex which rounded off Sherrington's long pursuit of tonus. The earlier
manifestation of 'decerebrate rigidity' Sherrington often called reflex standing.
The newly discovered stretch reflex was the continuing muscular response to
continuing tension applied to tendons, and through them, to proprioceptive
sense organs in the muscle. The knee-jerk was the transient muscular response
to transient tension and was thus a fractional manifestation of the stretch reflex.
Once more, too, there was a background in the stretch reflex for showing the
still unresolved problem of which the solution was then just out of reach:
autogenetic inhibition and excitation. 'Like the heart, it (musc
opposed nerves, one of augmentive function, one of inhibitory
Sherrington pointed out that another problem was here concerne
still waiting for solution-how the muscle receptors connect a
mental region of the brain. 'In their sensual aspect the muscul
serve as means whence mental experience can work towards attainm
finer delicacy and precision for the muscular acts of the body.'
Sherrington's experimental methods had never been elaborate; sim
surgery and modest electrical apparatus, but used with meticulous
eye ready for a meaningful result, and a mind which pondered. But
he brought into customary use the isometric myograph and so, w
recording, he had an instrument for the accurate measurement of
active tension of innervated muscle was measurable and compar
with the much lower tension of the paralyzed, denervated, or refle
muscle. It could be used to measure the rate at which spinal ne
brought into activity in different kinds of reflex response and the
they could be put out of activity by inhibitory stimuli. The excitat
and the inhibitory process I could be balanced and measured
another. This use of the isometric lever fructified thought on the
neurone of the final common path, and on the number of muscle f
each motoneurone controlled as one team of tension producers.
the conception of the 'motor unit', a spinal motoneurone realized
muscle team of one or two hundred muscle fibres which are p
measurable tension. These teams or 'packets' of fibres may ru
hundred in any one muscle. The idea of the motor unit completed S
contribution to the physiological anatomy of the function of spinal
forty years before. Further experimental support for the idea wa
sought.
Experiments interpreted on the basis of motor units excitated or inhibited
revealed much information about the convergence of nerve impulses on a group
of spinal motor units with a single function-'spinal centre' as they were vaguely
called before, but now 'motoneurone pool'. Into that pool poured streams of
excitation or of inhibition from various axones of distant origin. Any one of
these streams would debouch with full force on particular motor units, all at
the same time, and play in smaller streams on a number of other functionally
less accessible units, but perhaps not with sufficient force to stir them into
action. This less accessible kind of motor unit was said to be in the fringe of
excitation. That weaker stream by itself could seldom stir fringe units into
action. The fringe units had to be played on as well by another stream over-
lapping from another source before they would discharge. This fringe effect in
a motoneurone pool is implicit in the convergence of streams of impulses
ingoing along different paths. Convergence of pathways is part of the anatomy
of the central nervous system, but the overlapping of streams and fringe effects
depend on physiological conditions. All contribute to integration. If I (inhibition)
is pitted against E (excitation) early in a reflex contraction, it may have little or
no evident effect so long as E is being poured into th
but it may have a clear diminishing effect on after
flow of E has ceased, even when applied a considera
earlier. I, therefore, can in some way or somehow,
circumstances I does have an effect on E, the effec
smoothness of muscular response which comes from
motor units, as though the less strongly excited uni
more strongly excited were not. That E (or 'cent
could thus have a variable grip on neurones, dep
terminal points to be excited, and I ('central inhibi
implied that c.e.s. or c.i.s. might often be sublimin
subliminal volley of c.e.s. following after an earlier
now producing evident activity would show that the ef
persisted, though latently. It was found that the pe
gradually diminishing but not disappearing for a
oligo-synaptic reflex like the flexor reflex but lastin
in a poly-synaptic reflex like the crossed extensor re
The kinship between inhibition and fatigue, touch
was shown still more clearly by the isometric met
effect when tissue is fatigued. In brief, fatigue fa
always said it did. Once again there were there good
intimate nature of excitation and inhibition. 'In
acceleration of the spontaneous dissipation of an ex
Such acceleration might occur conceivably in sev
of the medium, or by increasing the local tempe
a catalyst, or by increasing the local conditions
neutralization of the excitation-ions or by change
the concentration.'
Thus Sherrington had built up by many points of circumstantial evidence a
great weight of support for the theory of dynamic polarization of nerve cells
promulgated by Cajal in 1889. It appears most probable, and is still at least a
fruitful working hypothesis. Cajal's end-knobs, 'boutons terminaux' (or plain
unadorned thread-like terminations) are the point of ending of the axone from
other nerve cells which lie in proximity to but do not touch the dendrites or cell
body of the receiving nerve-cells. That is the nerve synapse, the place of inter-
cellular conduction, the nexus between cells which is a 'surface of separation'.
When a sufficiency of end-knobs are active in time and space, so as to produce
a change at the synapse which is primarily (or solely) ionic, the receiving nerve
cell is excited and discharges, and so the excitation passes on to another nerve
cell or to an effector organ.
From many methods of approach Sherrington and his school had produced
the circumstantial evidence for this belief. Reflex action of short latency, the
long recruitment of motoneurones through multiple synapses, after-discharge
and well-timed inhibition all worked together to ensure speed, smoothness and
adequate endurance of muscular acts.
His Hughlings Jackson Lecture (1931) and Chapter VII of Reflex ac
the spinal cord, 1932, were two of the last comprehensive reviews fr
on muscle management at spinal levels. All his life, his mind was to
happy with research to feel the urge for writing many books. The in
action of the nervous system (1906) had been a necessary publicat
Silliman Lectures at Yale, which to accord with the terms of the Fo
were 'the basis of a volume to form part of a series'. It was reprinte
1914, 1916, 1920 and 1947, and did much to enhance and widen his r
in neurophysiology.
Sherrington did not allow his early acquaintance with pathology to l
his start on the problems of brain and cord. His appointment to St Th
1887 brought him in touch with pathology and some questions su
formation of scar-tissue, leucocytes, the specific gravity of the blood
in the secretions, the infectivity of oysters. The additional appointme
as Professor Superintendent of the Brown Institute of Preventive M
gave greater opportunity for research in animals larger than the
laboratory size. So it came about that he was the first in England to
use diphtheria antitoxin-and that on a moribund nephew-with
success. His own words describing this dramatic case are to be found
and Records of the Royal Society of London, 1948, 5, 156, and there
account in Nature, 1948, 161, 266-267.
After 1895, when he accepted the Holt Chair of Physiology at L
Sherrington's active work in pathology, but not his interest, came t
except for a dreary revival on behalf of the War Office Committee o
in 1916-1917. But he continued to write on hygiene, fatigue of school
and their hours of sleep.
In 1889, the Physiological Society appointed Sherrington to be
Secretary in succession to Gerald F. Yeo, who had been Secretary
foundation in 1876. This post with its steady load of administrat
Sherrington continued to hold until 1905. His temperament and i
were well suited for the additional post of secretaryship of the Inte
Physiological Congress which occupied him for six triennial congres
1892 to 1907. These were happy events where an example in friendly
formal procedure had been given at the first meeting in 1889 by Mich
who 'seated himself in a front bench or chair and rather ostentatious
his pipe'. Sherrington's friendly unassuming manner and the distinc
freshness of his demonstrations-difficult enough in strange laborat
foreign land-were long remembered by his contemporaries. Ewald sa
at that time: 'Seine Arbeiten durch ihre Sauberkeit imponieren'. At
1892, he demonstrated in a monkey the cortical control of the anus a
(and read a paper on varieties of leucocytes); at Berne in 1895, r
innervation in a monkey; at Cambridge in 1898, inhibition of tonus in
muscle by excitation of its antagonist ('ebenso eleganter Versuch' as
noted). At Turia in 1901, he could, for obvious reasons, describe only
show, his experiments on cerebral localization in a gorilla and in chi
In 1904, the Congress in Brussels saw Sherrington
reflex in the spinal dog.
The Heidelberg Congress in 1907 was his last meet
Secretary of the English language-a title which ind
for the English and American physiologists. His ow
years had been truly international.
In physical build Sherrington was rather short
Thanks to his wiry constitution he was able in mid
experimental researches, oblivious of physical discom
bent on the experiment. Working as he did often w
he had long worn for myopia, his close peering
emphasis to the intensity of his concentration. Exce
seventy-five he arranged to have at least one long e
sometimes more, and, as well, spent many hours on
time was he a secluded worker. He had a welcome f
the laboratory. During the First World War, as C
Fatigue Board, he used to speculate with some curio
even existence of industrial fatigue. At the beginn
have experienced industrial fatigue himself, since h
in a shell factory at Birmingham. The day shift was
shift nine hours. He was then fifty-seven. Bench wo
then or ever. Later during the war he helped the L
scientific inquiries, and wrote part of the book Alco
organism.
Sherrington had inevitably lost the facilities of the Brown Institute when he
moved to Liverpool in 1895, but from October 1898 he was to have his share
of the new Thompson-Yates laboratories for physiology and pathology which
were very much to his liking and convenience, with ideas on equipment which
some laboratories, even now, are still without. It was there that his thoughts
began to turn to teaching the students some of the simpler classical experiments
in mammalian physiology as a partial substitute for many involved and meaning-
less procedures in frog nerve-muscle classes. After some years of trial and the
use of typewritten notes (of which there was only one carefully guarded copy,
so that each student had to find spare time to write out from it his practical work
in his own note-book) Sherrington's class book on Mammalian physiology: A
course of practical exercises was published in 1919, at the first possible moment
after his coming to Oxford and the end of the war. The book injected entirely
new possibilities into teaching physiology especially in countries like ours where
experimentation is circumscribed by law. The illustrations for it were taken
from students' results in class, so that a foreign reviewer might well complain
that the 'authorities' quoted were all English! He taught regularly in the class,
plunging into its rapidly moving work with a lively interest and zest which
opened windows in the minds of its members. His scorn of carelessness was
equally impressive.
Invitation to Oxford and the Waynflete Chair of Physiology had come in
1913. As early as 1895, Sherrington had applied for this post. Now
invited and he accepted. The beauties of the place had long fascinated
the young life.
'Sworn Priest of Beauty, these thy shrines among,

That kneelest with old folks and that dancest with young.'
If the laboratory and its equipment were disappointing there were c

tions. So when the 1914-1918 war came to an end, and in spite of the
attaching to the Presidency of the Royal Society (1920-1925), She
entered into a busy and fruitful autumn of scientific endeavour, in wh
of his earlier work came to full harvest. That happy time was to last
when he retired from Oxford. He was then nearly seventy-nine.
In 1925, The assaying of Brabantius and other verse, by C. S. Sherr
evoked the hope in the breast of one reviewer that 'Miss Sherrington
continue to publish. These poems were a collection of previously
war-time poems, in which some readers found resemblances to Thoma
verse, and they showed a man whose emotions were strained by the so
war, 'Hate's voice, and, through its felon, War', which if others had se
as he had seen, and for so long, into the minds of militarists, might
come to pass. He never forgot the militarism at Strasbourg in the ei
the time of his writing these verses, he was stirred by the thought of you
giving their lives unquestioningly to pay for the mental blindness of
Years before he had written, 'Thirty-seven is too young to die'-and n
half that age were dying in thousands:
'the young folk, splashed with death in the trenched loam.'
The Rede Lecture at Cambridge in 1933 showed how the bio

philosopher and poet combined in Sherrington. His theme was 'The-b
its mechanism'. 'What right have we to conjoin mental experience with
logical? No scientific right; only the right of what Keats with that su
Shakespearian gift of his dubbed "busy common sense".' In Sherringt
poet was never deep down. In any prose he cared much for 'word-musi
will remember how a few weeks after the publication of Bridges' The t
of beauty (1930), Sherrington, then at the age of seventy-three, was re
heart long passages from it during the preparation of laboratory expe
For Goethe the poet he felt a sympathetic friendship, but not for G
scientist. 'Poet though he was, he was yet life-long an ardent student
sciences of Nature. And . . . Nature was usually Nature with a c
'Goethe could never describe a character-only a type. He was nev
same class as Shakespeare. Still, I have often enjoyed singing his balla
the first part of Faust is as good as anything that Shakespeare wrote.' Of G
scientific writings, Sherrington said: 'To appraise them is not a congen
The philosopher had long dwelt in him, culminating in a study
Fernel, the sixteenth-century French physician who lived at a time w
R
Renaissance was beginning to reach France and who
of his time, struggled to the light as Sherrington
himself in his time struggling against the enemies
long on Fernel's thoughts and sayings and could ca
to memory, the conversations that Fernel might hav
on the deepest problems of Life. Fernel became his
in spirit but not in body. So it was that the Gifford
1937-1938 were formed around Fernel and his times
Man on his nature (1940) and (1951, revised), Sherr
problem. 'The principle of life, it may turn out, is n
convention. But what of the Mind? Mind knows itself and knows the world:
chemistry and physics, explaining so much, cannot undertake to explain Mind
itself. It can intensify knowledge of Nature but it cannot be shown that Mind
has hitherto directed the operations of Nature. In that sense Mind and Natu
are different.' In talking over the problems of dualism, he said once not lon
before he died: 'Some people think that Thinking is all electric currents
Well!' He could not. And he had written in his lectures: 'Man's analysis of hi
sensible world seems to have outstripped his analysis of his own mind.'
He was an antiquarian too. In those early days in Venice he had learned how
to identify the origins of Italian incunabula and had built up a small collection
of antiquarian books about which he was, however, usually very reticent. On
one occasion his enthusiasm bubbled over and he told friends of a newly
discovered reference to syphilis by Widman of Wildbad as early as 1457, we
before the later accepted date of 1495 for the Naples epidemic.
In 1932 Sherrington was awarded the Nobel Prize in Physiology (jointly with
E. D. Adrian). He had been created Knight Grand Cross of the Most Excellent
Order of the British Empire in 1922 and the Order of Merit was conferred
1924. He was elected Fellow of the Royal Society in 1893 and gave the Croonian
Lecture in 1897 and the Ferrier Lecture in 1929. He was awarded the Roy
Medal in 1905 and the Copley Medal in 1927. He held honorary Degree o
Doctorate in twenty-two universities: Oxford, Paris, Manchester, Strasbourg
Louvain, Uppsala, Lyons, Budapest, Athens, London, Toronto, Harvard,
Dublin, Edinburgh, Montreal, Liverpool, Brussels, Sheffield, Berne, Birming
ham, Glasgow and the University of Wales, and was a corresponding membe
of many learned societies.
In 1892 Sherrington married Ethel Mary, younger daughter of John Ely
Wright, of Preston Manor, Suffolk. She died in 1933. C. E. R. Sherrington is
their son. She was a loyal and lively helpmate. In the Oxford period on Sunday
afternoons, there might be as many as two score of visitors, dropping in to te
at 9 Chadlington Road, where all were certain of a gay and easy welcome from
their host and hostess, especially warm if they came from overseas. By many
those visitors Sherrington will be best remembered as brimming joyously wit
vivid memories of places and people and full of shrewd forgiving reflexions o
the ways of mankind. By closer friends he will be remembered for an un
surpassing warmth of affection, a genial generosity of advice and time. He
practised straightforward self-effacing friendship. 'Altruism has to
positive charity is wanted', and that meant in his own private life as well a
affairs of the world. Beneath that genial exterior there was, howeve
balanced highly critical mind. There had to be. He was highly critica
life of his own work, viewing it from every angle, and making it true
and he was just as critical of the work or thoughts of others, but in th
of maturity he sealed his lips against criticism of those others. He c
broadcast many a pithy judgment, but he felt that to speak thus wo
bruised and not cured. So with only a few was this special wisdom sh
knew well the value of his own immense contributions to neurophysio
his humility in social intercourse was a solvent, as he meant it to be,
silence or shy reverence from a junior. He loved young men and youn
men and wanted to know their thoughts and problems and not just
pedestal above them. Up to the last hour of his life (he died instantly o
heart failure) his mental faculties were crystal clear, but his bodily h
been tediously frail for some years before his end. He found tha
isn't pleasant. One can't do things for oneself.'
It has been said of Charles Sherrington that he achieved for the
system what William Harvey achieved for the circulation. That is broa
He rooted out confused ideas and put in their place clear new discove
to make a chart of function to guide physiologists as well as clinicia
owe him much.
E. G. T. LIDDELL
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almost entirely, is his compilation.
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1899. The teaching of physiology and histology. Brit. Med. J. 1, 878.
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1899. On the relation between structure and function as examined in the arm. (Inaugural
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1899. (With H. E. HERING.) Inhibition of the contraction of voluntary muscles by
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1899. Inhibition of the tonus of a voluntary muscle by excitation of its antagonist.
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1900. Experiments on the value of vascular and visceral factors for the genesis of emotion.
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1900. The spinal cord. Text-book of Physiology. Edited by E. A. Schifer, Edinburgh
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1900. The parts of the brain below cerebral cortex, viz. medulla oblongata, pons,
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1900. Cutaneous sensations. Text-book of Physiology. Edited by E. A. Schifer, Edin-
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1900. The muscular sense. Text-book of Physiology. Edited by E. A. Schifer, Edinburgh
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1900. Nature of tendon reflexes-a discussion of E. Jendrassik's paper before 13th Int.
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1901. The spinal roots and dissociative anaesthesia in the monkey. J. Physiol. 27, 360.
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1901. (With A. S. F. GRV)NBAUM.) An address on localization in the 'motor' ce
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1902. (With A. FROEHLICH.) Path of impulses for inhibition under decerebrate rigidity.
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1902. Observations on 'flicker' in binocular vision. Proc. Roy. Soc. 71, 71.
1902. (With A. S. F. GRitNBAUM.) A discussion on the motor cortex as exemplified in
the anthropoid apes. Brit. Med. J. 2, 784.
1902. (With E. W. WALLIS & Others.) Committee report on the conditions of health
essential to carrying on the work of instruction in schools. Brit. Assoc. Rep.
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1902. (With J. BARR & Others.) Committee reports of special chloroform committee of
the British Medical Association. Brit. Med. J. 1902, 2, 116; 1903, 2, cxli;
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1902. Fatigue. A lecture to the Froebel Society, Owens College, Manchester. Abstract
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1902. Address to the Conference on the Hygiene of Social Life. J. R. Sanit. Inst. 23,
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1902. C. S. Roy, 1854-1897. Year Book of the Royal Society, p. 231.
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1902. Remarks at discussion on pathology of nerve degeneration. Brit. Med. J. 2, 928.
1902. (With E. E. LASLETT.) Note upon descending intrinsic spinal tracts in the
mammalian cord. Proc. Roy. Soc. 71, 115.
1903. (With R. BOYCE & R. Ross.) The history of the discovery of trypanosomes in man.
Lancet, 1, 509. Preliminary letter, Lancet, 1902, 2, 1426; Brit. Med. J. 2, 1680.
1903. (With E. E. LASLETT.) Observations on some spinal reflexes and the interconnection
of spinal segments. J. Physiol. 29, 58.
1903. (With E. E. LASLETT.) Remarks on the dorsal spino-cerebellar tract. J. Physiol.
29, 188.
1903. Physiology and nervous diseases. An address delivered to 'Doctorate Graduates'
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1903. An address on science and medicine in the modern university (delivered at the
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1903. (With A. S. F. GRtiNBAUM.) Observations on the physiology of the cerebral cortex
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1903. Qualitative difference of spinal reflex corresponding with qualitative difference of
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1903. Address on medical science. Canad. J. Med. Surg. 14, 321; Dom. Med. Mon. 21,
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1903. (With S. C. M. SOWTON.) On the dosage of the mammalian heart by chloroform.
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1904. On binocular flicker and the correlation of activity of 'corresponding' retinal
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1904. On certain spinal reflexes in the dog. Proc. Physiol. Soc., J. Physiol. 13, xvii.
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31, 234.
1904. (With S. C. M. SOWTON.) On the dosage of the isolated mammalian heart by
chloroform. Appendix I to the Third Report of Special Chloroform Committee.
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1904. (With S. J. HICKSON & Others.) Committee report on madreporaria of the
Bermuda Islands. Brit. Assoc. Rep. 1904, p. 299; 1905, p. 186; 1906, p. 325.
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1905. On the relative effects of chloroform upon the heart and upon other muscular
organs. Brit. Med. J. 2, 181. (Appendix I of Fourth Report of Special Chloro-
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1905. Physiology: its scope and method. From Oxford Lectures on Methods of Science,
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1905. The importance of longer hours of sleep at public schools. Brit. Med. J. 2, 1469
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1905. Obituary. Sir John Burdon-Sanderson. Brit. Med. J. 2, 1491.
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1906. On the proprio-ceptive system, especially in its reflex aspect. Brain, 29, 467.
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1906. Observations on the scratch-reflex in the spinal dog. J. Physiol. 34, 1.
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dioxide on cardiac and other muscle. Appendix III of Fifth Report of Special
Chloroform Committee, see 1902, 1905. Brit. Med. J. 2, 85.
1906. (With H. E. ROAF.) Experiments in examination of the locked-jaw induced by
tetanus-toxin. J. Physiol. 34, 315; abstract, Lancet, 2, 810; Brit. Med. J. 2, 9.
1906. (With H. E. ROAF.) The mechanism of 'locked-jaw' produced by tetanus-toxin.
Brit. Med. J. 2, 1805.
1907. Appreciation of Sir Michael Foster. Brit. Med. J. 1, 351.
1907. The Association and medical research. (Commenting on Mott's Review of the
Integrative action of the nervous system.) Brit. Med. J. 1, 657.
1907. On reciprocal innervation of antagonistic muscles. Tenth note. Proc. Roy. Soc. B,
79, 337.
1907. Nerve as a master of muscle. Not. Proc. Roy. Instn, 18, 609; Sci. Amer. Suppl.
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1907. Strychnine and reflex inhibition of skeletal muscle. J. Physiol. 36, 185.
1907. On reciprocal innervation of antagonistic muscles. Eleventh note. Further
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1907. Spinal reflexes. Brit. Assoc. Rep. p. 667.
1907. Address on 'Inhibition' to the Royal Medical Society of Edinburgh. (Reported
in The Scotsman, 26 October.)
1908. A discussion on the scientific education of the medical student. (Meeting of the
British Medical Association.) Brit. Med. J. 2, 380; Lancet, 2, 480.
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reflexes. Proc. Roy. Soc. B, 80, 552; reprinted Folia neuro-biol. 1909, 2, 578;
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1908. On reciprocal innervation of antagonistic muscles. Thirteenth note. On the
antagonism between reflex inhibition and reflex excitation. Proc. Roy. Soc. B,
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1908. (With S. M. COPEMAN.) Committee report on body metabolism in
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1909. On plastic tonus and proprioceptive reflexes. Quart. 5. Exp. Physiol
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1909. Discussion on the deep afferents; their function and distribution (meeting of the
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1910. Obituary. W. Page May. Brit. Med. J. 1, 298.
1910. Flexion-reflex of the limb, crossed extension-reflex, and reflex stepping and
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1910. Remarks on the reflex mechanism of the step. Brain, 33, 1.
1910. (With F. M. TOZER.) Receptors and afferents of the third, fourth and sixth cranial
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1910. (With H. E. ROAF.) Further remarks on the mammalian spinal preparation.
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1910. Notes on the scratch reflex of the cat. Quart. J. Exp. Physiol. 3, 213.
1910. Brain, physiology of. In Encyclopaedia Britannica, London and New York,
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1910. Note on certain reflex actions connected with the mouth. Brit. Dent. J. 31, 785.
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fatigue. Brit. Assoc. Rep. 1910, p. 292; 1911, p. 174.
1911. (With S. C. M. SOWTON.) On reflex rebound. Proc. Physiol. Soc. (printed but
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1911. (With S. C. M. SOWTON.) Reversal of the reflex effect of an afferent nerve by
altering the character of the electrical stimulus applied. Proc. Roy. Soc. B, 83,
435; reprinted Z. allg. Physiol. 12, 485.
1911. (With F. W. MOTT & E. SCHUSTER.) Motor localization in the brain of the gibbon,
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1911. (With T. GRAHAM BROWN.) Notes on the pilomotor system. Quart. J. Exp.
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1911. Muscle and nerve. In Encyclopaedia Britannica, 11th ed. 19, 44.
1911. Spinal cord, physiology of. In Encyclopaedia Britannica, 11th ed. 25, 672.
1911. Sympathetic system. In Encyclopaedia Britannica, 11th ed. 26, 287.
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1911. The role of reflex inhibition. Sci. Progr. Twent. Cent. no. 20, p. 584; translated in
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1911. (With S. C. M. SOWTON.) Chloroform and reversal of reflex effect. J. Physiol.
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1911. Sir Rupert Boyce, 1863-1911. Proc. Roy. Soc. B, 84, 1.
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1911. (With A. G. W. OWEN.) Observations on strychnine reversal. J. Physiol. 43,
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1912. Note on present problems of nervous function. In Melanges biologiques, 1912,
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1912. (With T. GRAHAM BROWN.) On the instability of a cortical point. Proc. Roy. Soc.
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1912. Bewegung und Leben. Address to the students at Utrecht, May 1912.
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1912. Report of Departmental Committee on Sight Tests, Board of Trade.
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University Press. (Six chapters on physiology 'briefly explaining the principles
which underlie the precepts and practice described in the other chapters of the
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1913. Reciprocal innervation and symmetrical muscles. Proc. Roy. Soc. B, 86, 219.
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1913. Reflex inhibition as a factor in the co-ordination of movements and postures.
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1913. The sight tests of the Board of Trade. Lancet, 1, 1691.
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1913. An address on the provincial school of medicine and the provincial university.
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1913. Rhythmic reflex produced by antagonizing reflex excitation by reflex inhibition.
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1914. Report on reciprocal innervation. Trans. Int. Congr. Med. Sect. II, Physiol., p. 85.
1914. (With A. FORBES.) Acoustic reflexes in the decerebrate cat. Amer. J. Physiol. 35,
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1915. (With S. C. M. SOWTON.) Observations on reflex responses to single break-
shocks. J. Physiol. 49, 331.
1915. (With F. R. MILLER.) Some observations on the bucco-pharyngeal stage of reflex
deglutition in the cat. Quart. J. Exp. Physiol. 9, 147.
1915. Postural activity of muscle and nerve. Brain, 38, 191.
1915. Simple apparatus for obtaining a decerebrate preparation of the cat. Proc. Physiol.
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1915. (With S. KENT.) Committee report on the structure and function of the mammalian
heart. Brit. Assoc. Rep. 1915, p. 226; 1916, p. 304; 1917, p. 122.
1916. A simple apparatus for illustrating the Listing-Donders law. Proc. Physiol. Soc.,
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1917. (With A. S. F. LEYTON.) Observations on the excitable cortex of the chimpanzee,
orang-utan and gorilla. Quart. J. Exp. Physiol. 11, 135.
1917. Reflexes elicitable in the cat from pinna, vibrissae and jaws. J. Physiol. 51, 404.
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1917. Recent physiology and the war. Not. Proc. Roy. Instn, 22, 1; Science, 46, 502.
1918. Stimulation of the motor cortex in a monkey subject to epileptiform seizures.
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1918. (With N. B. DREYER.) Brevity, frequency of rhythm and amount of reflex nervous
discharge, as indicated by reflex contraction. Proc. Roy. Soc. B, 90, 270.
1918. Observations on the sensual r6le of the proprioceptive nerve-supply of the
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1920. Sir William Osler. Obituary. Brit. Med. J. i, 65.
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1920. Gateways of sense. Huxley lecture, Birmingham University, 26 No
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1921. (With K. SASSA.) On the myogram of the flexor-reflex evoked by a
shock. Proc. Roy. Soc. B, 92, 108.
1921. Break-shock reflexes and 'supramaximal' contraction-response of mammalian
nerve-muscle to single shock stimuli. Proc. Roy. Soc. B, 92, 245.
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1921. Sur la production d'influx nerveux dans l'arc nerveux reflexe. Arch. Int. Physiol.
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1922. Note on the after-discharge of reflex centres. In Libro en honor de Santiago
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1922. Some points regarding present-day views of reflex action. Address to Royal
Society of Edinburgh, 20 March. Nature, Lond. 109, 463.
1922. Some aspects of animal mechanism. Presidential address, British Association for
the Advancement of Science, Hull. Brit. Assoc. Rep. p. 1; Brit. Med. J. 2, 485;
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1922. Inaugural address, delivered at the opening of the Biological Building, McGill
University, Montreal, 5 October. Printed privately, Murray Printing Co. Ltd,
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1922. Anniversary address, delivered before the Royal Society, 30 November. Pro
Roy. Soc. A, 102, 373 (1923); Proc. Roy. Soc. B, 94, i; extract bearing title:
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1923. The position of psychology. Address to National Institute of Industrial Psycho
20 March. Abstract, Nature, Lond. 111, 439. (Apparently never printed in
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1923. (With E. G. T. LIDDELL.) Stimulus rhythm in reflex contraction. Proc. Roy. Soc.
B, 95, 142. Appendix on separation key.
1923. (With E. G. T. LIDDELL.) A comparison between certain features of the spinal
flexor reflex and of the decerebrate extensor reflex respectively. Proc. Roy.
Soc. B, 95, 299.
1923. (With E. G. T. LIDDELL.) Recruitment type of reflexes. Proc. Roy. Soc. B,
95, 407.
1923. Anniversary address, delivered before the Royal Society, 30 November. Proc.
Roy. Soc. A, 105, 1 (1924); Proc. Roy. Soc. B, 95, 485; Nature, Lond. 112, 845;
Brit. Med. J. 2, 1113.
1924. (With E. G. T. LIDDELL.) Reflexes in response to stretch (myotatic reflexes).
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1924. Problems of muscular receptivity. Nature, Lond. 113, 732, 892, 929.
1924. Anniversary address, delivered before the Royal Society, 1 December. Proc. Roy.
Soc. A, 107, 1 (1925); Proc. Roy. Soc. B, 97, 254; Nature, Lond. 113, 840.
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Proc. Roy. Soc. B, 97, 488.
1925. Remarks on some aspects of reflex inhibition. Proc. Roy. Soc. B, 97, 519.
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1925. An address on avenues in medicine. Delivered at the opening of the winter session
of the London (Royal Free Hospital) School of Medicine. Mag. Lond. (Roy.
Free Hosp.) School Med. for Women, 20, 133; Lancet, 2, 741. Abstract under
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1925. Address at the unveiling of the Wheatstone Memoria
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1925. J. N. Langley. Obituary. Brit. Med. J. 2, 925.
1925. Anniversary address, delivered before the Royal Society, 30 November. Proc.
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1926. (With R. S. CREED.) Observations on concurrent contraction of flexor muscles in
the flexion reflex. Proc. Roy. Soc. B, 100, 258. Appendix on double mirror
myograph.
1926. (With S. COOPER & D. E. DENNY-BROWN.) Reflex fractionation of a muscle.
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1927. (With S. COOPER & D. E. DENNY-BROWN.) Interaction between ipsilateral spinal
reflexes acting on the flexor muscles of the hind-limb. Proc. Roy. Soc. B, 101,
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1927. Whither?-A footnote. Nature, Lond. 119, 205.
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1927. Ernest Henry Starling. Obituary. Brit. Med. J. 2, 905.
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1927. Keith Lucas (1879-1916). Dictionary of National Biography, 1912-1921. [Supple-
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1927. (With H. E. ROAF & Others.) Committee report on colour vision, with particular
reference to classification of colour blindness. Brit. Assoc. Rep. p. 307.
1928. Foreword to L. J. J. Muskens, Epilepsy, comparative pathogenes, symptoms,
treatment. London: Bailliere, Tindall & Cox.
1928. Sir Dawson Williams. Obituary. Brit. Med. J. 1, 418.
1928. Eulogy of Harvey. The Harvey Tercentenary Celebrations, Royal College of
Physicians, 14 May. Brit. Med. J. 1, 866; Lancet, 1, 1034.
1928. Some physiological data toward functional analysis of a simple reflex centre.
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1928. Introduction to F. Mason, Creation by Evolution. A consensus of present-day
knowledge as set forth by leading authorities in non-technical language that all
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1928. A mammalian myograph. Proc. Physiol. Soc., J. Physiol. 66, iii.
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1929. Some functional problems attaching to convergence. Ferrier lecture. Proc. Roy.
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1929. (With J. C. ECCLES.) Improved bearing for the torsion myograph. Proc. Ph
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1929. The sympathetic system. Encyclopaedia Britannica, London and New York,
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Charles Scott Sherrington.

CHARLES SCOTT SHERRINGTON was born on 27 November 1857 in Islington,

The second person to influence Sherrington's thought was the Spanish

'Sworn Priest of Beauty, these thy shrines among,

If the laboratory and its equipment were disappointing there were c

'the young folk, splashed with death in the trenched loam.'

The Rede Lecture at Cambridge in 1933 showed how the bio

Grateful acknowledgement is due to Professor J. F. Fulton for the Bib

1884. (With J. N. LANGLEY.) On sections of the right half of the med

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