Obituary: Professor David Whitteridge

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David Whitteridge, physiologist: born London 22 June 1912; University Demonstrator in Physiology, Oxford University 1944-50; Professor of Physiology, Edinburgh University 1950-68; FRSE 1951; FRS 1953; Waynflete Professor of Physiology, Oxford University 1968-79 (Emeritus); married 1938 Gweneth Hutchings (died 1993; three daughters); died Oxford 15 June 1994.

DAVID WHITTERIDGE was one of those rare and favoured individuals who remain at the cutting edge of their profession for their entire lifetime.

He read medicine as a scholar at Magdalen College, Oxford, in 1931, but soon developed a passion for physiology, the study of the functions of the body, under the tutelage of MH MacKeith. The Waynflete Professor of Physiology, Sir Charles Sherrington, appointed 'DW' to a departmental demonstratorship in 1934 and he started working for a BSc degree (now called an MSc) supervised by J. C. 'Jack' Eccles. Early in 1935 he moved into Sherrington's own laboratory where he assisted Sherrington in his last demonstration of the operation of the motor cortex in primates.

After clinical training in London he returned to Oxford in 1938 and restarted the electrophysiological techniques that had been brought to a halt after Sherrington's retirement and Eccles's return to Australia. His 1989 monograph, in which he reviewed 100 years of international congresses of physiology, showed how extensive was his knowledge of general physiology and the scientists who worked in the field.

Whitteridge's long-standing interest in the stretch receptors in the heart and lungs began with a collaboration with Edith Bulbring in 1939. They were able to demonstrate new sensory nerve fibres from atrial and ventricular receptors of the heart. In the 1950s he was delighted to be corrected by one collaborator, Professor Autar Singh Paintal, who showed that the 'B-type' nerve fibres, which Whitteridge had supposed to come from the lungs, actually arose from the atria of the heart. After the Bhopal disaster in 1984, when due to human error poisonous gas escaped from a chemical plant, the two men resumed their collaboration in Delhi to study the mechanism of phosgene gas poisoning.

Whitteridge considered his most important work during the war to be that he did with Ludwig Guttman, Director of the Spinal Unit at Stoke Mandeville Hospital. The management of paraplegics was then in its infancy and the two men studied the physiological disturbances in paraplegics using apparatus developed and built by Whitteridge himself. They described for the first time the failure of control systems for blood pressure and temperature in paraplegics with lesions above thoracic vertebra 6, a condition now known as autonomic dysreflexia. For his research he was awarded the Radcliffe Prize and the degree of doctor of medicine in 1945. He also served as Secretary of the Physiological Society from 1946 to 1950 and was its Foreign Secretary from 1980 to 1986.

Whitteridge always sought the challenge of new fields and in 1948 began to study with Peter Daniel and Sybil Cooper the proprioceptive control - the sense of muscle and joint position - in eye movements. While investigating the origin of proprioceptive fibres Whitteridge discovered that the retinal image of the visual world was mapped topographically in the midbrain. This discovery ran counter to the prevailing ideas of a diffuse brain organisation. Following up this discovery was interrupted only by his move to the Chair of Physiology in Edinburgh in 1950 and his election to the Royal Societies of London and of Edinburgh, all before his 40th birthday.

Whitteridge's prime task in Edinburgh was to revive a dormant department and he achieved this with considerable distinction. In the laboratory he insisted on correct experimental technique and developed a reputation amongst his collaborators for having both a high IQ and a low pH. But his innate grace tempered his critical comments to clumsy experimentalists and his enthusiasm for research meant that even the most arduous experiments were a joy. Invariably they became extempore seminars that were always enlightening, seasoned with impish wit, and frequently strayed some distance from the business in hand. He became the mentor for the waifs and stray students that found their way to his door, and the many scientists who streamed in from all over the world for postgraduate training in his laboratory. A quintessential 'DW' scientific paper had a polyglot authorship and encapsulated a mass of experimental data in one or two simple diagrams.

Whitteridge's research in Edinburgh has now become standard lore in textbooks, it included investigation of the connections between the two halves of the brain and the neuromuscular control of breathing. His work with Peter Daniel revolutionised our understanding of how the visual field was represented in the primary visual cortex of the primate. Their map, based on extensive experimental data they collected over five summers, revealed that more cortical surface was devoted to the representation of the fovea, which is the area of highest acuity in the retina. Advised by the statistician David Kendall, they introduced the term 'cortical magnification' to express this concept mathematically, and suggested that there was a precise relationship between cortical magnification and visual acuity. More than 30 years later theirs remains the standard model.

From 1944 until Whitteridge returned to Oxford in 1968 as Waynflete Professor of Physiology, his right-hand man in the laboratory was W. T. S. 'Jock' Austin, who built apparatus and made sure that his 'Prof's' spectacular demonstrations to the students worked. After returning to Oxford, Whitteridge began a valuable collaboration with Peter Clarke and Ian Donaldson on the processing of binocular depth information in the visual cortex. VS Ramachandran came across from Cambridge and stimulated them to settle the debate about whether cortical depth cells were present at birth and therefore did not require visual experience. It was the kind of challenge that 'DW' loved. His final job before retiring was to convert the ocean of talk on reform of the Final Honour School of Physiology into a viable, beneficial scheme of enlarging the number of departments taking part.

After becoming an Emeritus Professor and Honorary Fellow of Magdalen College in 1979, Whitteridge had in mind to set up his microscope at home and to tend his collection of rhododendrons, but a lifetime of enthusiasm for physiological research could not be so easily contained. Professor Larry Weiskrantz supplied him with space in the Department of Experimental Psychology in Oxford, and Whitteridge persuaded the Medical Research Council to fund a pilot project in which he would begin to link structure and function of the visual cortex. Within five years, the team he formed was successful in charting the form, function, and intracortical projections of nerve cells in the visual cortex. Their discoveries were a critical factor in persuading the Medical Research Council to create a new unit to study the neural networks of the brain. Whitteridge's last paper was published in 1993 from the MRC's Anatomical Neuropharmacology Unit in Oxford.

Against the present age of extreme specialisation, Whitteridge's research endeavours may seem positively agoraphilic, but he loved the challenge of pioneering paths through thickets of ignorance, being content to move on afterwards and let others widen his trail. He was modest about his own contribution: for him success as a scientist was to have placed one anonymous brick in the edifice of knowledge. His endeavours over 60 years have left us with a finely crafted wall.

(Photograph omitted)