doi:10.1093/brain/awm098
Brain (2007), 130, 2758 ^2765
OCC ASIONAL PAPER
The physiologist and the neurosurgeon: the enduring
influence of Charles Sherrington on the career of
Wilder Penfield
William Feindel
Correspondence to: William Feindel, Director Emeritus, Curator, Wilder Penfield Archive, Montreal Neurological Institute,
McGill University, 3801 University St, Montreal, Quebec, Canada H3A 2B4
E-mail: [email protected]
Wilder Penfield, a Rhodes scholar from Princeton University, New Jersey, was a student in the first course on
mammalian physiology given in 1915 at Oxford University by Charles Sherrington, newly arrived from Liverpool
where, as Holt Professor of Physiology for 20 years, he had become a leading authority on the physiology of the
nervous system. The practical ‘exercises’ as well as graduate research on the Golgi apparatus and the decerebrate preparation, carried out by Penfield in Sherrington’s laboratory, gave him the groundwork to develop his
career as a physiological surgeon, who made fundamental observations on functional localization in the human
brain during the surgical treatment of patients afflicted with epilepsy.
Keywords: Sherrington; Penfield; Oxford; McGill; Brain
Received March 16, 2007. Accepted March 30, 2007. Advance Access publication May 29, 2007
Introduction
Sherrington as teacher
In January 1915, Wilder Penfield, a Rhodes scholar elected
from Princeton, became an undergraduate at Merton
College, Oxford University and enrolled in the course of
mammalian physiology directed by Professor Charles
Sherrington.
At Merton, Penfield settled into rooms in staircase
number 6 of the medieval section of the College called Mob
Quad (Fig. 1), claimed to be the oldest college quadrangle
in the Kingdom; for Penfield, in the winter of 1915,
it seemed certainly the coldest (Penfield, 1977) (As part of
the celebration of Merton’s 700th anniversary, a medical
symposium was held in the inclement month of April,
1964; Wilder Penfield and I attended and were kindly
assigned rooms by the Bursar in Mob Quad—they were still
cold.). His rooms were next to one where William Harvey,
who was Warden of the College in 1645, did some of
his work on the development of the chicken egg
which appeared in his de Generatione Animalium of 1651
(Cooke, 1975). Across the quadrangle, rooms were occupied
by Penfield’s Princeton classmate and fellow Rhodes
scholar, Wilburt Davison, also a student in Sherrington’s
class, and another Mertonian, a Harvard graduate, Tom
(TS) Eliot, who was reading philosophy but would move
off to London after that academic year.
On the 100th anniversary of Sherrington’s birth, Penfield
(1957) wrote in Brain:
It is clear that his physiological and philosophical teachings
will live on through the coming century but for most people
Sherrington, the man, is no more than a legend. Even those
who have met him may well remember only that he was a shy
man with a small body and a preoccupied manner; a man
oblivious of his own great stature and reputation.
For me he is much more than a legend. He was my teacher and
he is still my scientific hero. When I went to Oxford as an
undergraduate I met him for the first time. He was fifty-seven
years old then, a quick-moving man of medium height with
a small moustache and nose-glasses (Fig. 2), who hurried along
the corridors in a white coat, who moved from table to table
while we worked on our ‘‘preparations’’. How well I remember
him, appearing short-sighted, over my embarrassed shoulder.
His face was smooth and almost expressionless, but I watched
for the faint smile and the twinkle of humour and understanding in his eyes. ‘‘Hm, Penfield, you may be right’’ he
would say, ‘‘but, I should have thought . . .’’
Then he would pick up the delicately pointed forceps and
change the tissues of the preparation so as to set the youthful
experimenter back on ‘‘the right track.’’ Sherrington seemed
quite sincere in the expectation that each student would teach
him something, sometime.
ß The Author (2007). Published by Oxford University Press on behalf of the Guarantors of Brain. All rights reserved. For Permissions, please email: [email protected]
Sherrington and Penfield
Brain (2007), 130, 2758 ^2765
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Fig. 1 View of the North corner of the 13th century Mob Quad at
Merton Penfield’s rooms were in staircase six (the entrance on
the right under the lantern). In 1645, when he was Warden of
Merton, William Harvey carried out laboratory studies on
regeneration in a room over the left arched doorway
(William Feindel Photo Archive, circa 1958).
Fig. 2 Photograph of Charles Sherrington inscribed to his student
Wilder Penfield (Photo by Elliott and Fry, from the Wilder Penfield
Archive).
Penfield’s impressions of Sherrington as a teacher were
based on his participation in the laboratory course of
mammalian physiology, a highly organized series of 25
‘exercises’, as Sherrington termed them, that his pupils
carried out proceeding from simple to complex.
During the ‘exercises’, animals were prepared and
examined on a table with a heated chamber, with the
physiological changes such as nerve and muscle reflex
action or blood pressure recorded on a ‘smoked drum’ or
kymograph (Fig. 3). A sample of such an experimental
result, showing the effect of pituitary extract on the carotid
artery pressure, was obtained by Penfield working with his
partner Emile Holman, another Rhodes scholar in
Sherrington’s class (Fig. 4). This result along with those
from the work of other pupils figured in a laboratory
manual titled, Mammalian Physiology: A Course of Practical
Exercises (Sherrington, 1919). The detailed instructions and
the drawings for the surgical procedures cover some
sophisticated techniques, such as dissecting out the
splanchnic and sympathetic nerves or laminectomy to
expose the spinal cord.
In these laboratory instructions, Sherrington left little
room to chance. This is exemplified by his meticulous
directions for use of the scalpel (Fig. 5A and B). These
exercises proved of great value to Penfield since he learned
how to handle living tissues gently, to use fine surgical
dissection instruments and to maintain the vital function of
the experimental animals. His first application of this
experience, while still a medical student at Johns Hopkins
(Penfield, 1919), involved a laboratory project on the
treatment of severe and progressive haemorrhage by
intravenous injections.
As Sherrington modestly stated in the preface to his
laboratory manual (Sherrington, 1919), ‘By some of those
who have taken the course I have been told that they traced
to it a measure of facility in technique which proved of
Fig. 3 Wilder Penfield in Sherrington’s laboratory for mammalian physiology (1916). He wears a white collar and tie with his lab coat,
to take his examination in the Honours School of Physiology (Wilder Penfield Archive).
Fig. 4 Text-Fig. 24 from Sherrington’s Mammalian Physiology showing on the kymograph the effect of pituitary extract on carotid blood
pressure, an ‘Exercise’ carried out by the Rhodes scholars, Emile Holman and Wilder Penfield (Sherrington, 1919).
A
B
Fig. 5 Text Fig. 11, sketch (A) and text (B) to detail the use of the scalpel in dissection (Sherrington, 1919).
Sherrington and Penfield
Brain (2007), 130, 2758 ^2765
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Fig. 6 Graduating class of 1916 from the Honours School of Physiology, Oxford. Penfield sits in the middle row, third from the left and once
removed from Holman on his left. Davison, the third American Rhodes scholar is on the right in the front row. Sherrington, on the left in
the top row, is next to Professor Gowland Hopkins, biochemist and external examiner from Cambridge, later to receive a Nobel prize
(Wilder Penfield Archive).
service to them when entering upon independent investigation later on in one field or another of experimental
medicine.’
In a tribute to Sherrington, many years later, Penfield
noted.
It was not the example of Horsley or Cushing that led me into
surgery of the nervous system. It was the inspiration of
Sherrington. He was, so it seemed to me from the first, a
surgical physiologist, and I hoped then to become a
physiological surgeon (Penfield, 1952).
resident of one of the great American surgeon–scientists,
William Halsted; he then became Chairman of Surgery at
Stanford University.
Graduate studies at Oxford
After a surgical internship at the Peter Bent Brigham
Hospital where Harvey Cushing was Chief of Surgery,
Penfield returned to Oxford in 1919 for the third and final
year of his scholarship. Looking back on that happy postwar period Penfield (1957) recalled, about Sherrington,
Graduation from Oxford
Penfield completed the 3-year course in the Honours
School of Physiology in 2 years by taking the anatomy
course at Edinburgh during the summer vacation, a
shortcut arranged for him by Sir William Osler
(Lewis, 1981). The photograph of Sherrington’s first
graduating class of 1916 shows Sherrington, Penfield and
his two Rhodes scholar friends Davison and Holman, along
with others. The external examiner was Professor Gowland
Hopkins from Cambridge, who was later awarded a
Nobel prize for his biochemical work on vitamins
(Fig. 6). The three Rhodes scholars, with the help of
Osler’s letters of reference, were admitted to finish
their medical course at Johns Hopkins Hospital. Wilburt
Davison eventually became the founding Dean of the
new medical school at Duke University. Emile Holman
stayed on at Johns Hopkins, where he was the last
In later years, when I returned to Oxford for graduate
study, and during recurrent visits with him through the
long period of his retirement, I came to appreciate the brilliant
mind and the broad culture of this modest seeker after
the truth.
Harvey Cushing was an observer of Sherrington’s work in
Liverpool for a time, though never a regular worker in his
laboratory. He reported the Professor to be a man in a hurry
who wrote too much and had a poor memory for many things
including his glasses.
This is a curiously superficial misconception on the part of the
great American neurosurgeon!
I would say from the long acquaintance with him that
Sherrington’s memory excelled that of any man I have ever
known for accuracy of detail, whether employed in the telling
of thrilling tales of his adventurous youth, or in the recollection
of scientific detail.
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W. Feindel
B
A
Fig. 7 Camera lucida drawings by Penfield (1920) of the normal Golgi apparatus (A) in a nerve cell of the anterior horn of the spinal cord,
demonstrated by Cajal’s silver method. An anterior horn cell (B) shows retispersion of the Golgi apparatus 7 days after section of the
sciatic nerve (Wilder Penfield Archive).
However, despite Cushing’s rather brash critique of
Sherrington, they maintained a happy relationship for the
rest of their lives, affably exchanging letters and gifts of
books. And, as William Gibson relates, Sherrington, in
post-retirement from Oxford, made a special trip there
in July 1938 to witness Cushing receive a Honourary D.Sc.
in the Sheldonian Theatre (Gibson, 2007).
After reading his former student’s biography of Harvey
Cushing (Fulton, 1946), Sherrington commented to Ragnar
Granit, one of his students, ‘You know, the life of a great
surgeon is not really exciting, except, of course, from the
patient’s point of view’. Sherrington regarded as exceptions
Lord Lister and Penfield, ‘both also scientists and
physiologists.’ (Granit, 1966).
During Penfield’s second stage at Oxford, Sherrington
was an infrequent visitor to the laboratory, being occupied
with work on health commissions, as President of the
Royal Society (1920–1925) and, in 1922, President of
the British Association for the Advancement of Science.
He spent much of his time in London dealing with these
non-academic matters (Eccles and Gibson, 1979).
For his graduate work, Penfield carried out two projects
in the Oxford laboratory. First was a neat study of the Golgi
apparatus of the nerve cells located in the anterior horn of
the spinal cord. Following section of the sciatic nerve he
noted changes in the Golgi apparatus, characterized by its
eccentric displacement and retispersion in the nerve
cell body (Fig. 7A and B). He published his results in
Brain (Penfield, 1920) and received the Oxford degree of
B.Sc. for this research. It was Sherrington who had
originally suggested to Penfield that he might try Cajal’s
techniques for demonstrating components of the
nerve cells.
A more extensive project, a detailed study of spinal
reflexes in the chronic as well as the acute decerebrate
animal, was carried out with Cuthbert Bazett and
also reported at length in Brain (Bazett and Penfield,
1922). The decerebrate preparation had been used
by Sherrington since 1898, during his Liverpool period;
but most of his studies had been in the acute stage
after decerebration. It was a tour de force for Bazett
and Penfield to keep their animals surviving for as long as
3 weeks.
These two research projects in the Sherrington laboratory
epitomized Penfield’s later career; he became an expert
neurocytologist and neuropathologist on the one hand,
and on the other, a neurosurgeon well-versed in the
fundamentals of the nervous system (Feindel, 1977a; Eccles
and Feindel, 1978).
Osler
Penfield’s post-war session at Oxford was marred by the
illness and death of Sir William Osler in December 1919.
As mentioned, Osler had befriended Penfield, helping him
take the summer course in anatomy at Edinburgh (Fig. 8).
Sherrington and Penfield
Brain (2007), 130, 2758 ^2765
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Fig. 8 Sir William Osler in his library at Oxford (circa 1912);
over the mantle is the triptych of Linacre, Harvey and
Sydenham, now in the Osler Library at McGill (Osler Library
Photo Archive).
Penfield, captivated by Osler’s charm and erudition, took
him as his lifelong medical hero (Penfield, 1927, 1949;
Feindel, 2003). Osler, as Regius Professor of Medicine at
Oxford, had chaired the selection committee in 1913 that
recommended Sherrington to be Waynflete Professor of
Physiology. Osler’s great textbook of medicine published in
1892, when he was at Johns Hopkins Hospital, played
a critical role in directing the Rockefeller fortune into
medical research (Penfield, 1967). In the last year of his life,
Osler initiated a gift from the Rockefeller Foundation
of 5 million dollars for Canadian medical schools
(Fedunkiw, 2005). At McGill, this program began with
a new biology building, opened in May, 1922 by
Charles Sherrington, then President of the Royal Society
and Harvey Cushing, who was Moseley Professor of Surgery
at Harvard. By an historical coincidence they gave addresses
on the McGill campus, where their student, Wilder
Penfield, was to arrive 6 years later. The culminating
feature of the McGill/Rockefeller program was the foundation in 1934 of McGill’s Montreal Neurological Institute
where Penfield and his associates later established a world
Fig. 9 McGill University’s Montreal Neurological
Institute opened by Penfield and his associates in 1934
(MNI Neuro-Archives).
center for neurology, neurosurgery and what we now call
the neurosciences (Fig. 9; Feindel, 2006). Sherrington’s
teachings were thus well vindicated (Elvidge and Penfield,
1935). His name appears with those of other luminaries on
the frieze at the entrance to the MNI in what Penfield
called the Neurological Hall of Fame.
Penfield’s tribute to Sherrington
Penfield
lifetime,
Charles
nervous
dedicated the last study he published in his
The Mystery of the Mind (Penfield, 1975), to Sir
Sherrington, ‘physiologist and explorer of the
system’. And he observed (Penfield, 1952),
As years passed his influence did not grow less but stronger.
Indeed, it often happens now that, during a routine operation,
when the human brain of a conscious patient lies exposed and
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W. Feindel
Fig. 10 Penfield and his surgical team operating on an epilepsy patient who is awake under local anaesthesia, in order to identify the
epileptic focus by brain mapping. Herbert Jasper, in the glazed gallery, records the electrocorticogram (Wilder Penfield Archive, 1954).
happy chance makes possible some observation that should
throw light on physiological mechanism, I often seem to feel
him looking over my shoulder (Fig. 10).
Under the tutelage of Sherrington, Penfield had
formed the groundwork for his career as a physiological
surgeon of the human brain, with the ever-present aim to
reduce or cure his patients of their epileptic seizures
(Feindel, 1977a, b). In his later writings (Penfield, 1958,
1975) Penfield often referred to Sherrington’s views on the
endlessly vexatious mind/brain problem, expressed in the
Gifford Lectures (Sherrington, 1942) and later in his brief
but still questioning Introductory to the broadcast series by
others on The Physical Basis of Mind (Sherrington, 1950).
Wilder Penfield ended his centennial tribute to Charles
Sherrington in Brain (Penfield, 1957) by quoting lines from
a poem by Sherrington, written in his first spring term at
Oxford, entitled ‘Oxford’ (Sherrington, 1925), lines that will
touch anyone who has lived and studied in that magical
towered city,
The night is fallen and still thou speakst to me,
what though with one voice sole, with accents many,
tongued turret and tongued stream, tracked pasture fenny
And cloister spirit trod, and centuried tree.
Acknowledgements
The author thanks Marcus Arts and Helmut Bernhard of
the Department of Neurophotography, MNI, for preparing
the Figures and Ann Watson for formatting the text.
Permission to use the illustrations has been kindly granted
by the Osler Library of the History of Medicine (Fig. 8) and
by Oxford University Press (Figs 4 and 5 and the lines
from Sherrington’s poem ‘Oxford’). The author appreciates
support from the Wilder Penfield Archive Fund provided
by the Class of Medicine (1945) of McGill University and
the Thomas Willis Fund of the MNI.
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