Skip Navigation


Brain Advance Access originally published online on May 29, 2007
Brain 2007 130(11):2758-2765; doi:10.1093/brain/awm098
This Article
Right arrow Abstract Freely available
Right arrow FREE Full Text (PDF) Freely available
Right arrow All Versions of this Article:
130/11/2758    most recent
awm098v2
awm098v1
Right arrow Alert me when this article is cited
Right arrow Alert me if a correction is posted
Services
Right arrow Email this article to a friend
Right arrow Similar articles in this journal
Right arrow Similar articles in ISI Web of Science
Right arrow Similar articles in PubMed
Right arrow Alert me to new issues of the journal
Right arrow Add to My Personal Archive
Right arrow Download to citation manager
Right arrowRequest Permissions
Right arrow Disclaimer
Google Scholar
Right arrow Articles by Feindel, W.
Right arrow Search for Related Content
PubMed
Right arrow PubMed Citation
Right arrow Articles by Feindel, W.
Social Bookmarking
 Add to CiteULike   Add to Connotea   Add to Del.icio.us  
What's this?

© The Author (2007). Published by Oxford University Press on behalf of the Guarantors of Brain. All rights reserved. For Permissions, please email: journals.permissions@oxfordjournals.org

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: william.feindel{at}mcgill.ca


    Summary
 Top
 Summary
 Introduction
 Sherrington as teacher
 Graduation from Oxford
 Graduate studies at Oxford
 Osler
 Penfield's tribute to...
 References
 
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.

Key Words: Sherrington; Penfield; Oxford; McGill; Brain

Received March 16, 2007. Accepted March 30, 2007.


    Introduction
 Top
 Summary
 Introduction
 Sherrington as teacher
 Graduation from Oxford
 Graduate studies at Oxford
 Osler
 Penfield's tribute to...
 References
 
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, 1977Go) (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, 1975Go). 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.


Figure 1
View larger version (100K):
[in this window]
[in a new window]
[Download PowerPoint slide]
 
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).

 

    Sherrington as teacher
 Top
 Summary
 Introduction
 Sherrington as teacher
 Graduation from Oxford
 Graduate studies at Oxford
 Osler
 Penfield's tribute to...
 References
 
On the 100th anniversary of Sherrington's birth, Penfield (1957Go) 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 ..."


Figure 2
View larger version (82K):
[in this window]
[in a new window]
[Download PowerPoint slide]
 
Fig. 2 Photograph of Charles Sherrington inscribed to his student Wilder Penfield (Photo by Elliott and Fry, from the Wilder Penfield Archive).

 
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.

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, 1919Go). 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.


Figure 3
View larger version (148K):
[in this window]
[in a new window]
[Download PowerPoint slide]
 
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).

 

Figure 4
View larger version (63K):
[in this window]
[in a new window]
[Download PowerPoint slide]
 
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, 1919Go).

 
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, 1919Go), involved a laboratory project on the treatment of severe and progressive haemorrhage by intravenous injections.


Figure 5
View larger version (20K):
[in this window]
[in a new window]
[Download PowerPoint slide]
 
Fig. 5 Text Fig. 11, sketch (A) and text (B) to detail the use of the scalpel in dissection (Sherrington, 1919Go).

 
As Sherrington modestly stated in the preface to his laboratory manual (Sherrington, 1919Go), ‘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 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, 1952Go).


    Graduation from Oxford
 Top
 Summary
 Introduction
 Sherrington as teacher
 Graduation from Oxford
 Graduate studies at Oxford
 Osler
 Penfield's tribute to...
 References
 
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, 1981Go). 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 resident of one of the great American surgeon–scientists, William Halsted; he then became Chairman of Surgery at Stanford University.


Figure 6
View larger version (127K):
[in this window]
[in a new window]
[Download PowerPoint slide]
 
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).

 

    Graduate studies at Oxford
 Top
 Summary
 Introduction
 Sherrington as teacher
 Graduation from Oxford
 Graduate studies at Oxford
 Osler
 Penfield's tribute to...
 References
 
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 post-war period Penfield (1957Go) recalled, about Sherrington,
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.

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, 2007Go).

After reading his former student's biography of Harvey Cushing (Fulton, 1946Go), 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, 1966Go).

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, 1979Go).

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, 1920Go) 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, 1922Go). 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.


Figure 7
View larger version (44K):
[in this window]
[in a new window]
[Download PowerPoint slide]
 
Fig. 7 Camera lucida drawings by Penfield (1920Go) 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).

 
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, 1977aGo; Eccles and Feindel, 1978Go).


    Osler
 Top
 Summary
 Introduction
 Sherrington as teacher
 Graduation from Oxford
 Graduate studies at Oxford
 Osler
 Penfield's tribute to...
 References
 
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). Penfield, captivated by Osler's charm and erudition, took him as his lifelong medical hero (Penfield, 1927Go, 1949Go; Feindel, 2003Go). 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, 1967Go). In the last year of his life, Osler initiated a gift from the Rockefeller Foundation of 5 million dollars for Canadian medical schools (Fedunkiw, 2005Go). 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 center for neurology, neurosurgery and what we now call the neurosciences (Fig. 9; Feindel, 2006Go). Sherrington's teachings were thus well vindicated (Elvidge and Penfield, 1935Go). 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.


Figure 8
View larger version (127K):
[in this window]
[in a new window]
[Download PowerPoint slide]
 
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).

 

Figure 9
View larger version (110K):
[in this window]
[in a new window]
[Download PowerPoint slide]
 
Fig. 9 McGill University's Montreal Neurological Institute opened by Penfield and his associates in 1934 (MNI Neuro-Archives).

 

    Penfield's tribute to Sherrington
 Top
 Summary
 Introduction
 Sherrington as teacher
 Graduation from Oxford
 Graduate studies at Oxford
 Osler
 Penfield's tribute to...
 References
 
Penfield dedicated the last study he published in his lifetime, The Mystery of the Mind (Penfield, 1975Go), to Sir Charles Sherrington, ‘physiologist and explorer of the nervous system’. And he observed (Penfield, 1952Go),
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 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).


Figure 10
View larger version (139K):
[in this window]
[in a new window]
[Download PowerPoint slide]
 
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).

 

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, 1977aGo, bGo). In his later writings (Penfield, 1958Go, 1975Go) Penfield often referred to Sherrington's views on the endlessly vexatious mind/brain problem, expressed in the Gifford Lectures (Sherrington, 1942Go) and later in his brief but still questioning Introductory to the broadcast series by others on The Physical Basis of Mind (Sherrington, 1950Go).

Wilder Penfield ended his centennial tribute to Charles Sherrington in Brain (Penfield, 1957Go) by quoting lines from a poem by Sherrington, written in his first spring term at Oxford, entitled ‘Oxford’ (Sherrington, 1925Go), 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.


    References
 Top
 Summary
 Introduction
 Sherrington as teacher
 Graduation from Oxford
 Graduate studies at Oxford
 Osler
 Penfield's tribute to...
 References
 
Bazett HC, Penfield W. A study of the Sherrington decerebrate animal in the chronic as well as the acute condition. Brain (1922) 45:185–264.[Free Full Text]

Cooke AN. William Harvey at Oxford. J Roy Coll Physins Lond (1975) 9:181–8.

Eccles J, Feindel W. Wilder Graves Penfield 1891-1976. Biog Memoirs Fellows Roy Soc (1978) 24:472–513. [includes CV and Bibliography].[CrossRef]

Eccles JC, Gibson WC. Sherrington, his life and thought. (1979) Berlin Heidelberg: Springer-Verlag.

Elvidge AR, Penfield W. Sir Charles Sherrington. Arch Neurol Psychiat (1935) 34:1299–309.

Fedunkiw MP. Rockefeller Foundation funding and medical education in Toronto, Montreal and Halifax. (2005) Montreal and Kingston: McGill-Queen's University Press.

Feindel W. Wilder Penfield (1891-1976): The man and his work. Neurosurgery (1977a) 1:93–100.[Medline]

Feindel W, ed. Wilder Penfield: his legacy to neurology. CMAJ (1977b) 116:3–16.

Feindel W. Osler and "the medico-chirurgical neurologists": Horsley, Cushing and Penfield. J Neurosurg (2003) 99:188–99.[Medline]

Feindel W. The Rockefellers to the rescue. CMAJ (2006) 175:1252–3.[Free Full Text]

Fulton JF. Harvey Cushing, a biography. (1946) Springfield, Illinois: Charles C. Thomas.

Gibson WC. A student recalls Sir Charles Sherrington, O.M. (1859-1952). Brain (2007) [this issue].

Granit R. Charles Scott Sherrington, an appraisal. (1966) London: Thomas Nelson and Sons Ltd.

Lewis J. Something hidden, a biography of Wilder Penfield. (1981) Toronto: Doubleday Canada Limited.

Penfield W. The treatment of severe and progressive hemorrhage by intravenous injection. Am J Physiol (1919) 48:121–32.[Free Full Text]

Penfield WG. Alterations of the Golgi apparatus in nerve cells. Brain (1920) 43:295–305.

Penfield W. A medical student's memories of the Regius Professor. (1927) 385–7. Bull Int Assc Med Mus No. 9 (Sir William Osler Memorial volume).

Penfield W. Neurology in Canada and the Osler Centennial. CMAJ (1949) 61:69–73.

Penfield W. Sir Charles Sherrington, O.M. G.B.E. F.R.S. Nature (1952) 169:698.

Penfield W. Charles Scott Sherrington, Poet and Philospher. Brain (1957) 80:402–10.[Free Full Text]

Penfield W. The excitable cortex in conscious man. (1958) (Sherrington Lecture) Liverpool: Liverpool University Press.

Penfield W. Sir Charles Sherrington, O.M. F.R.S. (1857-1952). Notes Records Roy Soc (1962) 17:162–8.

Penfield W. The difficult art of giving: the epic of Alan Gregg. (1967) Boston/Toronto: Little, Brown and Company.

Penfield W. The mystery of the mind. (1975) Princeton: Princeton University Press.

Penfield W. No man alone: a neurosurgeon's life. (1977) Boston/Toronto: Little, Brown and Company.

Sherrington CS. Mammalian physiology: a course of practical exercises. (1919) Oxford: Clarendon Press.

Sherrington CS. The assaying of Brabantius and other verse. (1925) Oxford: Oxford University Press.

Sherrington CS. Man on his nature. (1942) Cambridge: Cambridge University Press.

Sherrington CS. Introductory. In: The physical basis of mind.—Laslett P, ed. (1950) Oxford: Basil Blackwell. 1–4.


Add to CiteULike CiteULike   Add to Connotea Connotea   Add to Del.icio.us Del.icio.us    What's this?



This Article
Right arrow Abstract Freely available
Right arrow FREE Full Text (PDF) Freely available
Right arrow All Versions of this Article:
130/11/2758    most recent
awm098v2
awm098v1
Right arrow Alert me when this article is cited
Right arrow Alert me if a correction is posted
Services
Right arrow Email this article to a friend
Right arrow Similar articles in this journal
Right arrow Similar articles in ISI Web of Science
Right arrow Similar articles in PubMed
Right arrow Alert me to new issues of the journal
Right arrow Add to My Personal Archive
Right arrow Download to citation manager
Right arrowRequest Permissions
Right arrow Disclaimer
Google Scholar
Right arrow Articles by Feindel, W.
Right arrow Search for Related Content
PubMed
Right arrow PubMed Citation
Right arrow Articles by Feindel, W.
Social Bookmarking
 Add to CiteULike   Add to Connotea   Add to Del.icio.us  
What's this?