b.22 June 1912 d.15 June 1994
BA Oxon(1934) BSc(1936) BM BCh(1937) DM(1946) FRSE(1951) FRS(1953) FRCP(1966) Hon DSc Edin(1993)
David Whitteridge was Waynflete professor of physiology at the University of Oxford. He was born in South Norwood, London. His father, Walter Randall Whitteridge, was a woollen manufacturer's agent. His mother, Jeanne Hortense (née Carouge), was French. David was educated at Whitgift School, Croydon, and Magdalen College, Oxford; his clinical studies being undertaken at Kings College Hospital, London. After qualifying he spent a period as RMO at Finchley Memorial Hospital before returning to Oxford in 1938 as departmental demonstrator in physiology, a post he held concurrently with a Beit memorial research fellowship in 1939, and a Schorstein fellowship for medical research, 1940. At Oxford he started to investigate the sensory information that was conveyed from the heart and the lungs in the many afferent branches of the vagal nerve. At an early stage he began to collaborate with Edith Bulbring, who had recently arrived in the Oxford pharmacology department from Nazi Germany, via the School of Pharmacy. Together they applied the single fibre recording techniques which had been pioneered by Lord Adrian [Munk’s Roll, Vol.VII, p.3] for following the activity of mechanoreceptors of skin and muscle. The approach proved to be immensely successful and, after the Second World War, Whitteridge and his pupils were able to describe a range of phenomena, much of which was completely new. Whitteridge was delighted when his first research student in Edinburgh, A S Paintal, extended the experimental analysis of cardiac afferent activity and completely revised earlier conclusions. From 1944 to 1950 he was a demonstrator in physiology and a fellow by special election at Magdalen College. He moved to Edinburgh in 1950 as professor of physiology, transforming the department there into one of the most active in the British Isles. Largely for his experimental investigation of visceral afferent activity,Whitteridge was elected to the Royal Society in 1953.
David Whitteridge's quickness of mind and tongue were obvious after the briefest conversation. He had an outstanding career as an undergraduate. In his second year at Magdalen he won the Theodore Williams scholarship for part one of the BM examinations (anatomy and physiology) and at the end of his third year he gained first class honours in the final honour school of natural sciences (physiology). He then interrupted his medical studies to spend a year working on the electrophysiology of neuromuscular transmission under the supervision of J C Eccles. For this work he was awarded a BSc, a degree which the University later recognized as an MSc. He had a rather different experience of research while a clinical student at King’s. He acted as one of the heroic subjects of R A McCance [Munk’s Roll, Vol.IX, p.327] in the latter’s investigation of the effects of disturbances of fluid and electrolyte balance in normal subjects. In spite of temporally losing a third of his extracellular fluid volume from excess sweating during the course of this study, he formed a high opinion of McCance whom he regarded as one of the few physicians who were prepared to investigate physiological disturbances in initially healthy subjects, so as to treat patients in a more rational fashion.
Whitteridge was later to carry out two important investigations on human subjects. The first was started towards the end of the Second World War. Having spent the early years of the war working on blast injury, Whitteridge met Ludwig Guttmann, later Sir Ludwig [Munk’s Roll, Vol.VII, p.233] who was looking after patients with spinal injuries at Stoke Mandeville Hospital. At that time paraplegic patients were managed with little hope of rehabilitation, most of them dying from infections within a few weeks or months of their spinal injury. Guttman recognized the need to prevent pressure sores and also realized that further improvements in management would require a better understanding of derangements of autonomic function. Using apparatus largely constructed by Whitteridge, they demonstrated clear cut alterations in the regulation of arterial blood pressure and body temperature in patients with lesions above T6. Most important, they investigated the effect of bladder distension and showed how the altered responses of skin blood flow above the level of the lesion could be used by the patient as a signal that the bladder needed to be emptied. By making use of this information patients became continent at an early stage and risks of urinary tract infection were greatly reduced. For this work Whitteridge received his doctorate in 1945 and the Radcliffe prize in 1946. The second important investigation, on human subjects, was carried out approximately ten years after Whitteridge had moved to Edinburgh: using electromyography he and his colleagues demonstrated the involvement of the intercostal muscles in resting and stimulated levels of breathing. These studies provided real observations to replace dogma which was no more than speculation based on the idealized arrangement of the fibres of the internal and external intercostal muscles.
Before he moved to Edinburgh he had become interested in the proprioceptive control of the external eye muscles. With Sibyl Cooper and Peter Daniel he discovered that the retinal image is mapped in neurones in the mid-brain. This led him into the field of visual neurophysiology which was to remain his principle research interest for the rest of his life. Like Sherrington [Munk’s Roll, Vol.IV, p.523] Whitteridge believed that our understanding of the mechanisms of the brain required detailed knowledge of the fine structure of neural connections. This was an unfashionable view around 1950 and, in collaboration with Peter Daniel, Whitteridge set out to provide the basic correlates between structure and function in the visual pathways. Although Daniel was based in London the work was not halted by Whitteridge’s move to Edinburgh. Over a period of five summers they mapped the visual field on the cerebral cortex of the monkey, recognizing the columnar arrangement of the cortical neurones and its potential functional importance. This work laid the foundation for many of the spectacular advances in our understanding of the visual system which followed in the 1960s and 1970s.
In 1968 Whitteridge returned to Oxford as Waynflete professor of physiology and for several years pursued the problem of how the visual system processes binocular information concerning the depth of field. A series of solid contributions resulted from this work, but in 1977 he began to survey the form, function and intracortical projections of the nerve cells within the visual cortex in collaboration with a post-doctoral fellow, Kevin Martin. At the time when Whitteridge retired from the Waynflete chair in 1979 the work had started to attract interest and at the invitation of Larry Weiskrantz, he and Martin - with the support of the Medical Research Council - moved their laboratory to the department of experimental psychology. Over the next few years the work flourished and Whitteridge was delighted both by the recognition given to Martin and in feeling that he himself "... could still do something of interest to others".
Whitteridge's qualities as an academic leader became obvious when he was in his early thirties. During the years 1944 to 1950 several young men, of great potential but little research experience, were appointed to the Oxford laboratory of physiology. They were committed to teaching on a scale which would make today's lecturers blanch. Whitteridge saw their difficulties, helping them with their publications and providing both criticism and support. These same young men helped to make the Oxford laboratory the principal centre for experimental physiology in the UK when Whitteridge became Waynflete professor some 20 years later. The problems were somewhat different when he moved to Edinburgh. Although his predecessor had been a distinguished research worker, few of those remaining on the academic staff of the department were actively engaged in research. Whitteridge stirred his colleagues by setting a personal example both as a research worker and a teacher - enlivening instruction with demonstrations, often on human subjects, with new practical experiments and the introduction of an intercalated BSc in physiology for medical students. He waged war on those of his staff who confined their teaching to the content of textbooks and pointed out not only that didactic physiology was merely "..something easily purveyed and conveniently examined..." but also that many of the favoured textbooks of the time made statements which were demonstrably wrong. This vigorous approach enabled him to recruit intelligent and enthusiastic young people to his staff and to attract academic visitors and research students from overseas, many of whom came from the Indian subcontinent. Both young members of staff and visitors benefited from Whitteridge's interest in physiology and his analytical mind. Even those who were working on subjects distinct from Whitteridge’s own direct experience at the bench found that their ideas were clarified, and sometimes modified, by an informal conversation with him.
To those who did not know him, David Whitteridge appeared a rather severe figure. He had a reputation for uncompromising criticism of what he regarded as intellecutal slackness. Some of his more pungent remarks were reserved for the self-important and particularly for those who embroidered the truth. He was described by G L Brown as "...a man of high integrity and low pH". This epithet was almost immediately misquoted as "... a man of high IQ and low pH" and the misquoted form was the label for Whitteridge for the rest of his life. While lacking pseudo-bonhomie and trivial small talk, he showed warmth, wit and charm in informal conversation as well as a formidable array of interests and knowledge. Unlike many distinguished men, Whitteridge was modest about his own achievements and shared with younger men a contempt for the pompous. During his period as foreign secretary of the Physiological Society in the 1980s he was immensely popular with the younger members of the Society’s committee who, in view of his reputation, were surprised by his kindness and approachability and who admired his depth of understanding of modern physiology and his outspokenness on ethical issues.
He served on many national and international committees. India and Sri Lanka often sought his advice through his distinguished pupils. He was a member of the committee of the All India Institute of Medical Sciences in 1964 and an external examiner in Malaya, Burma and Ceylon. Apart from the usual run of grants committees, Whitteridge gave much of his time to the running of the Physiological Society, both at the beginning and end of his career. He was a member of the Society’s committee from 1944 to 1947 and from 1951 to 1954; secretary of the Society from 1947 to 1951 and foreign secretary from 1980 to 1986. Throughout his working life and after his official retirement he continued to be a loyal supporter of the Society’s scientific meetings and at eighty he could ask penetrating questions which would unsettle over-confident speakers.
Whitteridge had a deep interest in the history of physiology and valued his own contacts with Sherrington, Adrian and the generation of pre-war physiologists. As a research student in 1934 he had assisted Sherrington in the last demonstration the latter gave to students. His knowledge of the methods used by physiologists of the 19th and early 20th centuries was profound and both in Edinburgh and Oxford he discovered and restored equipment dating from this period, displaying it in an attractive and useful way.
He married Gweneth Hutchings [Munk’s Roll, Vol.IX, p.581] in 1938. She predeceased him. She was the distinguished Harveian scholar and David encouraged her in this field, appreciating that, when it came to understanding Harvey’s writings, her training as a scholar of medieval French and Latin was more appropriate than a knowledge of the classical Latin of the Roman Empire. He took great pride in the recogniton that Gweneth eventually achieved, particularly in her election as an honorary fellow of the College. They had three children.
C C Michel
[Brit.med.J., 1994,309,122; The Times, 25 June 1994;The Independent, 23 June 1994; The Guardian, 1 July 1994; The Daily Telegraph, 14 July 1994]
(Volume X, page 514)
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