Obituaries

A scroll through these entries illustrates the tremendous achievements that physiologists have achieved over many generations. Their valuable contributions to the discipline benefit us all.

This page provides brief notices on the deaths of Society Members as soon as we hear about them. Full PDF obituaries are presented where available.

J M Adam

2013

The Society regrets to announce the death Colonel J M Adam OBE OStJ FRCP who was elected a Member in 1960.

Takis Anagnostopolous

( 1936 - 2000 )

Takis Anagnostopoulos was a renal physiologist, with special interests in ion transport in the nephron. After medical studies in Paris and training in the US, he created the first French laboratory of renal electrophysiology. He developed this during the 1980s within the INSERM Unit at the Necker Hospital for Sick Children. Doctor of Medicine and Docteur des Sciences, he taught in three Paris universities and was an early advocate for the modernisation of medical and biological curricula.

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Declan John Anderson

d.2016

The Society is saddened to hear of the death of Declan Anderson, Emeritus Professor of Oral Biology at Bristol. Declan Anderson was a dentist and physiologist; but foremost, a scientist. He died at the age of 95 on Easter Day, 27 March 2016. He was educated at Christ’s Hospital School and Guy’s Hospital. After graduating in dentistry in 1942, he went on to obtain a B.Sc. in Physiology in 1946, and a Ph. D. in 1955. During these latter studies, he held clinical posts and a lectureship in Physiology at Guy’s. A full obituary will be appear in Physiology News 103.

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Roger Anwyl

d.2019

The Society regrets to hear of the death of Roger Anwyl on 21 May 2019 after a long illness. A graduate of the University of Birmingham (BSc in 1969 and PhD in 1972) Roger worked with Peter Usherwood at the University of Glasgow and Toshio Narahashi in Chicago before taking up a lectureship in 1978 at Trinity College Dublin where he subsequently became Professor of Neurophysiology and then Fellow Emeritus on his retirement. A Member of The Society from 1979, his research centred on mechanisms of glutamatergic transmission, including the roles of metabotropic glutamate receptors, in hippocampal synaptic plasticity.

David Armstrong

The Society regrets to hear of the death of David Armstrong, emeritus Professor of Physiology at the University of Bristol, after a long illness following a stroke. David was internationally recognised for his work on how the brain controls goal-directed movement, using a combination of anatomical, physiological and behavioural methods. Completing a PhD in Canberra with John Eccles, he moved to Bristol where he became head of Physiology in 1990. A Society member from 1969, he was an editor of The Journal of Physiology between 1979 and 1986. After retirement, David took up a second career in the Orkneys of ornithological photography.

Graham Francis Baker

( 1947 - 2004 )

Graham Baker graduated in physiology in 1969, gained a PhD and was appointed lecturer at Bedford College in 1981. Throughout his career he worked predominantly on transport systems, most notably in human erythrocytes (with Wilfred Widdas) using self-built equipment. He discovered the asymmetric action of 4,6 O-ethylidene-D-glucose in inhibiting glucose transport, investigated the effect of ATP on phloretin activity (with Martha Kaloyianni) and hypothesized, with Richard Naftalin, a structural explanation for the apparent variable affinity of the glucose transporter. He was also interested in transport across the nasal mucosa, evolving a number of techniques to study the human airways in vitro, with implications for the understanding of air pollution and asthma.

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W. E. Balfour

( 1927 - 2007 )

Bill Balfour graduated with a BSc from King’s College London. After PhD studies with Catherine Hebb in Edinburgh, he spent his working life at The Physiological Laboratory, Cambridge, as an Agricultural Research Fellow and, subsequently, on the academic staff. Early collaborative work included studies on acetylcholine synthesis in brain, carrier proteins for thyroxine and triiodothyronine, the role of the carotid body in the regulation of erythropoiesis, adrenal gland secretion of progesterone, and absorption of colostrum in the newborn calf. Later, his main interest was in the regulation of blood volume. Bill was a dedicated teacher and an excellent and caring Director of Studies in King’s College.

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A. D. Bangham

( 1922 - 2010 )

Working at Babraham, Alec Bangham made research contributions in many areas including mechanisms of anaesthesia, lung surfactants, haemoglobin polymorphisms, water structure and the evolutionary origin of life. His most significant discovery – that of the liposome – emerged from electron microscopy observations of hydrated lipid films with Bob Horne. Proteoliposomes subsequently became, and remain, a major tool for functional analysis of membrane transport and have also been developed as drug delivery systems.

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Jean Banister

( 1917 - 2013 )

Jean, a former teaching Fellow of Somerville College, Oxford, died on 15 February, aged 95. She had been a Member of The Society since 1950. Her research largely concentrated on the pulmonary vascular system. Jean is also remembered as a tireless champion of women’s education.

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James Alexander Barclay

( 1909 - 1999 )

James (Jimmy) Alexander Barclay, renal physiologist, studied Medicine in Aberdeen, He worked in the Department of Physiology at Birmingham University his entire professional life becoming a Reader and, for a while, acting Head of Department. His research centred on mechanisms of acid secretion by the kidney and, later, on the isolation of sodium-binding polypeptides from erythrocytes and cardiac muscle.

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Christopher Bell

( 1941 - 2008 )

Chris Bell took zoology at Melbourne University where he was one of Geoffrey Burnstock’s first PhD students. As a National Heart Foundation Overseas Research Fellow, he worked in the UK with Marthe Vogt at Babraham and with John Vane at the Royal College of Surgeons in London, studying the catecholaminergic innervation of peripheral tissues. Awarded the Sandoz Prize in 1972, Chris returned to the Department of Physiology in Melbourne, gaining his DSc (1980) and building an international reputation in vascular physiology. In 1995, he was appointed to the Chair in Physiology in Trinity College Dublin. He created new courses there in exercise physiology and neuroscience. He was editor of several international journals and wrote a number of popular books on physiology, for schoolchildren and others.

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Alan Benson

d. 2016

Alan Benson joined the Society in 1961. He specialised in aviation physiology and was Head of the Behavioural Sciences Division at the RAF Institute of Aviation Medicine at Farnborough.

Robert W. Berliner

( 1915 - 2002 )

Bob Berliner was one of the founding fathers of modern renal physiology. Director of the National Institutes of Health (NIH) Laboratory of Kidney and Electrolyte Metabolism for 12 years, in 1969 he became the first NIH Deputy Director for Science and, in 1973, Dean of the Yale University School of Medicine. Among his numerous honours and awards were the endowment by Yale of the Robert W Berliner Chair of Cardiology and Diagnostic Radiology, and the creation of the Robert W Berliner Lectureship in Renal Physiology.

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Gordon Bisset

d. 2013

The Society is sorry to hear of the death of Gordon Bisset, who was elected a Member in 1964 and was an Editor of The Journal of Physiology from 1966 to 1967.

Michael William Blackburn Bradbury

( 1930 - 2013 ) 

Professor of Physiology Kings College London 1977-1995, Emeritus 1995-2013. Mike Bradbury died peacefully on the 9th of February 2013 in Blandford, Dorset aged 82. He became a Member of the Society in 1964 and served on the Editorial Board of The Journal of Physiology from 1981-1988. His many friends and colleagues will miss him greatly.

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Alison Brading

( 1939 - 2011 )

Alison’s services to The Society were extensive. Alison studied Zoology at The University of Bristol, before working in the Department of Pharmacology in Oxford. Throughout her career she was a familiar and respected personality and made substantial contributions to understanding smooth muscle function.

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Hubert Britton

d. 2017

The Society notes the death of Member Hubert Britton in October 2017 at the age of 92. With a medical degree and a PhD in chemistry, Hubert joined the Physiology Department at St Mary’s Hospital Medical School, where he remained until retirement. A Member from 1965, he worked on the metabolic role of the placenta and the foetal endocrine system, and developed a method for studying enzyme function by induced transport. A staunch supporter of integrative physiology, he arranged many demonstrations linked to the clinical course he taught.

John Ramsey Bronk

( 1929 - 2007 )

Ramsey Bronk was first Professor of Biochemistry at the University of York. He came as a Rhodes Scholar to Oxford in 1952, studying for his doctorate with Fisher and Parsons, and returned to Washington (1956) to work in the National Institutes of Health. He was subsequently appointed Associate Professor of Zoology at Columbia University, becoming full Professor shortly before coming to York (1966) where he remained until his retirement (1997). There, he was closely involved in establishing the successful biochemistry degree course and the cancer research laboratories in the Biology Department, funded by Yorkshire Cancer Research. He wrote two major successful textbooks, Chemical biology and Human metabolism. His research interests included mitochondria, and thyroid hormones, and he worked extensively on metabolic aspects of chemotherapeutic drug delivery by peptide transport across the small intestine. He was a founding member of, and regular contributor to, the European Intestinal Transport Group and Chairman of the European Editorial Committee of Physiological Reviews, as well as a Distributing Editor for The Journal of Physiology.

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Alan G. Brown

( 1939 - 2006 )

Alan Brown, a medical graduate from the University of Edinburgh, qualified with a BSc in Physiology (1961) and MBChB (1964) and joined the Department of Veterinary Physiology at the Royal (Dick) Veterinary College. Awarded his PhD in 1968, he progressed to a Readership (1976), then full Professorship (1984). He was Head of Department of Preclinical Veterinary Sciences for much of the 1990s. He contributed prominently to our understanding of the diversity of sensory mechanisms in the dorsal horn. His research awards included a Beit Memorial Research Fellowship and MRC Research Fellowships. A Fellow of the Royal Society of Edinburgh (1984) and of the Institute of Biology (1988), he served as a member of the MRC Neurosciences Grants Committee and held editorial positions on The Journal of Physiology, the Quarterly Journal of Experimental Physiology, the Journal of Neurophysiology, Neuroscience and Brain Research/Brain Research Reviews.

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Eberhard Buhl

( 1959 - 2003 )

By the time of his appointment in 1999 as Professor of Neurobiology at Leeds, Eberhard Buhl had achieved international renown as an experimental neuroscientist. He pioneered the study of structure in relation to function in the brain, and of in vitro brain rhythms. He attracted major international collaborators and research grants and held important academic and managerial roles in physiology at local and national levels.

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Liam Burke

d. 2018

The Society belatedly notes the death of Member Liam (William) Burke at the age of 96. Following a first degree in pharmacy, completed in 1945, he was one of the only two UK PhD students to be supervised by Bernard Katz in the Department of Biophysics at UCL. After publishing work with Bernard Ginsborg on slow muscle fibres, he moved to Australia in 1956 where he subsequently carried out research with Peter O. Bishop recording from the visual system. He became Head of the Department of Physiology in Sydney, retiring as Emeritus in 1987. Much admired for his scientific curiosity and great sense of humour, he last visited UCL in 2016.

Dick Burn-Murdoch

( 1949 - 2000 )

Dick Burn-Murdoch qualified as a dentist in 1974 from Guy’s Hospital and joined the Physiology Department there as a Junior Lecturer. His teaching and research (into tooth eruption mechanisms, leading to a PhD in 1979) continued to the end of his life. He took on a wide range of responsibilities in administration and teaching – the latter, particularly on cardiovascular and respiratory physiology.

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Roger Carpenter

d. 2017

Roger Carpenter, Professor of Oculomotor Physiology at Cambridge, died on 27 October 2017, aged 72. His ‘Movements of the Eyes’ was a classic text. His own research, and his influential theoretical model, revealed how saccadic movements of the eyes could be used to study human decision making. His textbook of Neurophysiology has been used by generations of students. In his Cambridge college – Gonville and Caius – he was a much admired teacher: his engaging supervisions were intellectual in the noblest sense and inspired many medical students to combine experimental research with a clinical career.

Full Offsite Obituary

Alan Chipperfield

d. 2011

Alan Chipperfield died in October 2011 at the age of 73. He was elected a Member in 1978. His interests and expertise were in the areas of membrane transport physiology and he was distinguished for his work on sodium ion fluxes both through the sodium pump and through co-transporters (he was a pioneer in the area of sodium–potassium–chloride co-transport).

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Rob Clarke

( 1956 - 2004 )

Rob Clarke made an important contribution to physiology and to the Physiological Society. A physiology graduate from University of Manchester (1978), he studied for his PhD with Jim Pascoe at University College London. He later joined the Department of Physiology and Environmental Sciences (now Animal Sciences) at the University of Nottingham as lecturer, becoming Senior Lecturer in 1995. Working predominantly on nociception, he published more than 50 papers. He presented over 60 communications at Physiological Society meetings, convened the Somatosensory Physiology Special Interest Group, served as a Committee member and was closely involved in many of the Society’s educational initiatives. These included the Society’s booklet for schools (‘Understanding Life’), a major benchmarking statement for universities starting physiology related degrees, and – with the British Pharmacological Society – vacation courses on in vivo techniques for undergraduates.

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Ronald H. Cook

( 1914 - 2009 )

Ron Cook started work in the Cambridge Physiological Laboratory in 1948 as a Craftsman. He went on to work as Alan Hodgkin’s Instrument Maker, then Technical Officer until his retirement in 1987. With his interest in aquaria, he was able immediately to introduce cuttlefish (Sepia) to Cambridge. They were used by Hodgkin and Richard Keynes for the ion flux measurements that backed up the then new ionic theory of the action potential.

PDF Obituary 1 Oral history

Keith Cooper

d. 2011

Keith Cooper was elected a Member in 1950 and was on the Editorial Board of The Journal of Physiology from 1968 to 1972. Keith was active in the thermoregulatory field, initially at Mill Hill and subsequently in Canada at the University of Calgary.

John Coote

d. 2017

The Society is much saddened to hear of the death of Honorary Member John Coote, Bowman Professor of Physiology and Head of Department at the University of Birmingham from 1984 until his retirement. John joined the Society in 1967. He served on Council and as Chair of the editorial board of Experimental Physiology. An outstanding mentor to many students and colleagues, he was a distinguished autonomic neuroscientist with a wide range of interests including being a keen mountaineer and an expert in high altitude physiology. In 2003 he was awarded the Carl Ludwig Distinguished Lectureship of the American Physiological Society.

John Cotes

d. 2018

John Cotes died on 15 April 2018, aged 94. Working at the RAF Institute of Aviation Medicine, Farnborough (1950-2) then the MRC Pneumoconiosis Research Unit (PRU), Llandough (1953) his research concerned the design of an oxygen delivery system for the 1953 British Everest expedition. Oxygen equipment had a daunting reputation following failures on previous expeditions. As sole author of his first paper, he described an improved design and production of the oxygen masks and open circuit oxygen breathing circuits fundamental to the first successful Everest Summit. A member of The Society since 1954, he was well-known for his work on patients with industrial and other lung diseases at the PRU. Later, at the University of Newcastle, he developed systems to evaluate pulmonary function at rest and exercise, particularly lung gas transfer. The 6th Edition of his well-known text book, Lung Function: Physiology, Measurement and Application in Medicine, appeared in 2006.

David R Curtis

d. 2017

David Curtis FRS FAA died in December 2017, aged 90. David is famous for his development and application of the technique of multi-barrelled microelectrophoresis for recording activity from identified central neurones in vivo. Thus he was able to eject a range of compounds into the locality of single neurones. He combined this with activation of synaptic inputs either electrically or by application of other compounds. David, with his chemist colleagues, Jeff Watkins and Graham Johnston, demonstrated the excitatory and inhibitory effects of amino acids on single neurones. They developed the pharmacology of glutamate, GABA and glycine receptors along with novel agonists and antagonists. This provided the core evidence for establishing the role of these amino acids as synaptic transmitters in the mammalian CNS. David became the Howard Florey Professor of Medical Research and Director at the John Curtin School of Medical Research, Australian National University. David was elected to Fellowship of the Australian Academy of Science in 1965, becoming its President in 1986-90, and to Fellowship of the Royal Society in 1974.

Alan Cuthbert

( 1932 - 2016 )

Professor Alan Cuthbert passed away on 27 August following a short illness. Alan has been part of The Society since 1973. He was Shield Professor of Pharmacology at the University of Cambrige and Master of Fitzwilliam College, Cambridge. His work expanded our knowledge of the physiology and pharmacology of sodium transport in secretory epithelia.

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Michael de Burgh Daly

( 1922 - 2002 )

Michel de Burgh Daly was Emeritus Professor of Physiology in the University of London; Distinguished Visitor, Joint Department of Physiology Royal Free Hospital School of Medicine and University College, London (from 1984 until his death); and former Professor of Physiology and Head of Department of Physiology St Bartholomew’s Hospital Medical College, London (1958-84). His experiments on carotid body chemoreceptors and baroreceptors, in and out of the laboratory, have helped us to understand the mechanisms that underlie cardiovascular and respiratory reflexes.

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George Darlow

d. 2015

The Physiological Society is saddened to hear of the death of Dr George Darlow, who passed away in June.

Nicholas J. Davey

( 1957 - 2005 )

Nick Davey graduated from Bedford College London in 1980 with a zoology BSc, studying neural control of breathing for his subsequent PhD at Imperial College. His contributions to the science of spinal cord injury developed from early post-doctoral work on gamma motoneurone activity. Subsequent clinical research at the National Spinal Injuries Centre, Stoke Mandeville Hospital, led to a position there as Honorary Research Physiologist. After moving to the Department of Physiology at Charing Cross and Westminster Medical School (1988), he was promoted to Senior Lecturer in the Division of Neuroscience and Psychological Medicine at Imperial College in 2002. He was widely published and his findings that repetitive magnetic stimuli can have a therapeutic effect in spinal cord injury generated Clinical Initiative funding from the International Spinal Cord Trust. He also sought to engage the public with science, working with partners like the BBC and the Science Museum.

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C T M Davies

d. 2017

C.T. Mervyn Davies died on Sunday, 30 April 2017, after a short final illness, aged 83. He was at the forefront of research into human exercise physiology and part of a global network of like-minded scientists from the mid-60’s onwards. He was appointed founding Chair in Sport and Exercise Sciences at the University of Birmingham in 1986 and transformed a respected Physical Education department into a School that would become the world-ranked Sport and Exercise Sciences School it now is.

Joe Davison

( 1942 - 2013 )

The Society is sorry to hear of the death of Joe Davison, who joined The Society in 1974. Joe made important contributions to our understanding of the neural control of the gastrointestinal (GI) tract and its accessory organs. He died on 13 April at the age of 71.

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Hugh Davson

( 1909 - 1996 )

Hugh Davson’s work on the cell membrane was a major contribution to an area that is now recognised as fundamental for much of modern physiology and pharmacology. He was also a pioneering investigator of the physiology of eye fluids and the cerebrospinal fluid and a prolific and elegant writer of textbooks and research monographs. After wartime work on visual physiology with the military, he was employed as a member of staff of the Medical Research Council from 1954. Based at UCL for much of his career, he held several visiting professorships after his 1973 retirement in Britain and the US, where he was also awarded numerous prizes and honours. He was elected an honorary member of the Physiology Society in 1985.

Full Offsite Obituary Oral history

Eric J Denton

( 1923 - 2007 )

Professor Sir Eric Denton was one of the most distinguished marine biologists of the past century. A graduate of St John’s College, Cambridge, Denton’s early career included posts at TRE (the radar research establishment at Malvern), at the biophysics research unit, University College London, and as lecturer in physiology at Aberdeen. In 1956 he began an illustrious career at the Marine Biological Laboratory on Plymouth Hoe. He was first a staff physiologist, then a Royal Society research professor seconded from Bristol until 1974, and director of the laboratory from 1974 to 1987 – attracting visitors such as Sir Alan Hodgkin, Sir Andrew Huxley and Professor Richard Keynes. He continued at the laboratory as a research fellow until 2005. The scope of his work was notable: from buoyancy, to eyes and vision, photophores, camouflage and intraspecific signalling. He received many honours, from the Royal Society’s Royal Medal to the International Prize for Biology from the Japanese Society for the Promotion of Science.

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Jack Diamond

d. 2014

The Society was saddened to hear of the death of Jack Diamond, a prominent physiologist who helped establish neuroscience at the Michael G. DeGroote School of Medicine of McMaster University, at the age of 86.

Christopher John Dickinson

(1927 - 2015)

Christopher John Dickinson, Emeritus Professor of Medicine, Barts and The London School of Medicine and Dentistry (2002-15); Chairman, Department of Medicine, St Bartholomew’s Hospital Medical College (1975-92) died on the 30 September 2015 at the age of 88. He was a long time supporter of The Physiology Society, a member since 1966. He was Senior Lecturer at UCH London then Professor of Medicine at St Barts (London). He was interested in the nervous control of essential hypertension and co-authored a popular textbook (Clinical Physiology, E. J. Moran Campbell, C. J. Dickinson, J. H. D. Slater, C. R. W. Edwards, E. K. Sikora 1984 etc). In the late 1960s he developed a suite of computer programs to teach aspects of clinical physiology including respiration (Macpuf), kidney function (Macpee), drug action (Macdope) and integrative aspects of the circulation (Macman). (Despite their name, these programmes were initially written for microcomputers and then rewritten for PCs).

Alison Douglas

( 1961 - 2012 )

Alison worked in the School of Biomedical Sciences in Edinburgh, where she played an important role in teaching and administration, including as Organiser of the Honours Programme in Physiology (2001-2004) and as Chair of the Honours Exam Board in Medical Biology (2007-2011). In 2011 she was elected Chair of the British Society for Neuroendocrinology. She was elected a Member in 1995.

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William F Dryden

( 1941 - 2012 )

Bill Dryden, a neuroscientist and true nenaissance man, has died aged 71. Bill grew up in Paisley, just west of Glasgow, Scotland and received a BSc in Pharmacy from the University of Glasgow and a PhD in Pharmacology from the University of Strathclyde. Following postdoctoral studies in Neuroscience at the University of Pennsylvania, Bill returned to Strathclyde as a lecturer. He was appointed to the faculty in the Department of Pharmacology at the University of Alberta in 1976 where he remained until his retirement in 2008. He was acting chair of the Department between 1991 and 1992. Bill was a founding member of the Division of (later Centre for) Neuroscience and served as its director between 1995 and 1999.

His insight, breadth of knowledge, collegiality and sense of humour will be fondly remembered. He was a true renaissance man and mentor for us all. Bill is survived by his wife Angela, three children, Colin, Gillian and Anna and one grandchild.

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Helen Duke

d. 2014

The Society is sorry to hear of the death of Helen Duke. She became a Member in 1950 and published in respiratory physiology while at the Royal Free Hospital School of Medicine and the Middlesex Hospital Medical School.

Richard Edwards

( 1925 - 2013 )

Professor Richard HT Edwards was a physiologist and clinician with an international reputation for skeletal muscle research. A medical graduate from Middlesex Hospital Medical School, his interest in muscle developed during work at the Hammersmith Hospital and Royal Postgraduate Medical School and, as a Wellcome Trust Swedish Research Fellow, at the Karolinska Institute in 1970. He published a series of landmark papers in The Journal of Physiology, Clinical Science and The Lancet on muscle during exercise. With Professors David Hill and Victor Dubowitz, he secured a major grant to establish a Neuromuscular Centre at Hammersmith, then taking several of his research team with him when he was appointed Professor of Human Metabolism at University College Hospital (UCH, 1976). The group published important ground-breaking studies in muscle energetics, muscle protein turnover, muscle damage and muscle disorders. Richard became Head of the Department of Medicine at UCH (1982) and subsequently at University of Liverpool (1984), introducing major clinical and educational innovations there and developing an interest in chronic fatigue syndrome. He was subsequently appointed Professor of Research and Development for Health and Social Care at the University of Wales College of Medicine and Head of Research and Development for the NHS in Wales (1996-9)

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Sir Robert Edwards

( 1939 - 2009 )

Sir Robert was one of the pioneers of in vitro fertilisation, which is celebrated in Sir Richard Gardner’s personal account of Bob as a mentor, collaborator and friend in Physiology News. Bob was awarded Honorary Membership of The Society after receiving the Nobel Prize in Physiology or Medicine in 2010.

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Annie B Elliott

( 1931 - 2013 )

The Society is sorry to have to record that Annie B Elliott FRPharmS, PhD, of Sheffield, who was elected a Member in 1968, has died at the age of 98. She had worked in the Department of Biology in Nanyang University, Singapore.

Gerald Elliott

d. 2013

Gerald Elliot, founding Professor of Biophysics at the Open University and a Society Member of many years standing, has died in Oxford, aged 82. He gave his first presentation to a Physiological Society meeting in 1959, and his last in Dublin at the 2009 Summer meeting.

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John Ernsting

( 1928 - 2009 )

Air Vice-Marshall Professor John Ernsting’s work on aviation physiology and medicine had a profound and lasting influence on the design of crew support apparatus worldwide. Having qualified in Physiology and Medicine at Guy’s Hospital Medical School (1952), he was commissioned into the Medical Branch of the RAF. He subsequently led teams studying high altitude physiology, becoming deputy director of research, and then Commandant (1988-1992), at the RAF Institute of Aviation Medicine (IAM) at Farnborough. His work influenced international agreements on cabin pressures in modern airliners. On his retirement from the RAF, he came back to King’s as a visiting Professor where he was closely involved in educational innovations relating to aviation medicine and space physiology. He was appointed OBE in 1959, CB in 1992, and was a Queen’s honorary surgeon from 1989 to 1993. He was a past-president of the International Academy of Aviation and Space Medicine, and received the Louis Bauer Award from the Aerospace Medical Association in 2002.

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Gertrude Falk

( 1925 - 2008 )

Gertrude Falk completed a PhD at Rochester on diuresis in the rat and worked as a postdoctoral researcher with Gerrard in Chicago, studying a wide range of muscle types. She came on a Guggenheim Fellowship to the UCL Biophysics Department in 1961 from the University of Washington. An early microelectrode physiologist, she and Paul Fatt first studied muscle before turning their techniques to electrical studies of rod outer segments. Their work on synapses and retinal function generated a theoretical paper in 1974. Other publications followed, including a clutch of Nature papers with Jonathan Ashmore. Further discoveries about retinal physiology came later from her research with Richard Shiells.

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Paul Fatt

d. 2014

The Society is saddened to hear of the death of Professor Paul Fatt, Emeritus Professor of Biophysics, UCL, who passed away peacefully on 28 September. He was elected as an Honorary Member in 1990.

Oral history

Marianne Fillenz

d. 2012

Marianne was a neuroscientist who worked in Oxford having trained in Dunedin, New Zealand where she was taught by JC Eccles and was influenced by Karl Popper. Her own work included studies both on transmitter release and on inhibitory synaptic mechanisms in the CNS. She was medical tutor at St Anne’s College and had a remarkable influence on generations of science and medical students.

Tom Forrester

d. 2016

Tom Forrester was a Professor in the Department of Pharmacological and Physiological Sciences at the University of St Louis for over 30 years. Previously, he worked at the University of Glasgow and at University College London. He joined the Society in 1969 and, around that time, gave a number of communications on the release of ATP and its extracellular actions.

Venetia Franglen

( 1941 - 2007 )

Venetia Franglen was a PhD student at University College London in the 1960s, moving later to King’s College, where she was Sub-Dean in the medical faculty for some years. Working with senior physiologists like SE Dicker and Richard Durbin, she studied ion and electrolyte transport in skin (frog, fetal sheep and pig), and ouabain binding in gastric mucosa. She left Kings to become an Open University biology tutor and Curriculum Development facilitator at UCL.

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John Spence Gillespie

( 1926 - 2009 ) 

John Spence Gillespie, a medical graduate of Glasgow University, was later founding Head of the Department of Pharmacology there (1968-88), Professor of Pharmacology (1968-92) and Vice-Principal (1983-91). He served as Chairman of MRC Grants Committees, on the Committees of both the British Pharmacological Society (BPS) and The Physiological Society and, as BPS representative, on the Editorial Board of Pharmacological Reviews and on the Editorial Board for Monographs of The Physiological Society. He was Honorary Secretary of The Physiological Society from 1966 to 1972 and was a Fellow of the Royal Society of Edinburgh. His principal research interest was neurotransmission in the peripheral autonomic nervous system.

PDF Obituary 1 Oral history

Bernard Ginsborg

d. 2018

The Society regrets to hear of the death of Bernard Ginsborg at the age of 93. Born in London and trained as a physicist, he took a second degree in Physiology on AV Hill’s recommendation. With Paul Fatt at UCL he discovered voltage-gated calcium currents which led on to a long-standing research interest in synaptic neurotransmission and secretion. Moving to Edinburgh, he became the Head of the Department of Pharmacology in 1980, retiring in 1985. A member from 1957, Ginsborg was great supporter of The Society for many years and an outstanding Editor of The Journal of Physiology. He was elected a Fellow of the Royal Society of Edinburgh in 1971.

Margaret Hay Gladden

( 1940 - 2010 )

Margaret Gladden was a medical graduate from London University (1965), gaining her MRCS, LRCP, and DCh shortly after. She took up an MRC junior research fellowship in Geoffrey Kidd’s Liverpool laboratory, was awarded her PhD (1971) and published her first papers on the relationship between structure and function in the innervation of skeletal muscle. She later moved to Glasgow, joining Ian Boyd’s laboratory in the then Institute of Physiology as a research fellow and successively as lecturer (1973), senior lecturer (1983) and reader (1991). She published over 70 papers, mostly in The Journal of Physiology and almost entirely on the subject of muscle spindles.

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Constancio Gonzalez

( 1949 - 2015 )

The Society is sorry to hear of the sudden death of Dr Constancio Gonzalez, who was a JP Editor. Constancio had been a Member of The Society since 2012.

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John Archibald Browne Gray

( 1918 - 2011 )

John Gray studied medicine at Clare College, Cambridge and UCL Medical School, graduating in 1942. After wartime physiological research with the Pacific Fleet, he worked at the MRC’s National Institute for Medical Research (NIMR) on the sensory transduction of touch and pressure, moving later to the UCL Departmnent of Physiology as Reader. His series of papers in The Journal of Physiology (1950-7) significantly advanced our understanding of the Pacinian corpuscle and mechanoreception. He became a professor (1959), Dean of the UCL Science Faculty (early 1960s) and then ran the MRC (1966-77).After serving as MRC Secretary, he returned to research, studying the physiology of the lateral line in fish with Eric Denton at the Marine Biological Association Labs, Plymouth.

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John H. Green

( 1919 - 2000 )

Professor John Green spent most of his academic life in the Department of Physiology, Middlesex Hospital Medical School. A student at Cambridge and the Middlesex Hospital, he conducted cardiovascular and respiratory research, gaining a PhD in 1954 and a Professorial title in 1968. Author of numerous successful books for medical undergraduates, he was closely involved in the creation of teaching videos and held the patent, among others, on a finger cuff for monitoring post-operative blood pressure.

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David Greenfield

( 1917 - 2005 )

David Greenfield was Foundation Dean of the Faculty of Medicine and Professor of Physiology at the University of Nottingham between 1966 and 1981. A graduate of St Mary’s Hospital Medical School (1940), he started his physiology career as junior lecturer there – studying limb circulation, foetal haemodynamics and aviation medicine. Aged 31, he was appointed to the Dunville Chair of Physiology at Queen’s University of Belfast. His research there addressed the neural control of peripheral circulation, and many of his junior colleagues went on to prestigious academic careers. In 1963-4, he worked at San Francisco Medical Centre, developing a technique for testing cardiovascular reflex function later used by NASA. In 1966 he was appointed Dean at the newly-founded medical school at Nottingham and re-elected three times. His advice on the establishment of medical schools was sought by Universities around the globe. He was a member of the MRC, the GMC and several editorial boards and was elected an honorary member of The Physiological Society in 1987.

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Abraham Guz

d. 2014

The Society is sorry to hear of the death of Abraham Guz; he was 84 and had been a Member since 1966. He was Emeritus Professor at the National Heart and Lung Institute, Imperial College London, and had been Professor of Medicine at the Charing Cross Hospital Medical School 1973–82 and Professor and Head of the Department of Medicine of the joint Charing Cross and Westminster Medical School, University of London until September 1982–94.

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James Halliwell

d. 2012

James Halliwell, Emeritus Reader in Physiology at UCL, died on 7 March after a short illness. He was very much a ‘hands-on’ electrophysiologist who made substantial contributions to the physiology of ion channels in the CNS.

In recent years he focused on the roles of dopamine receptors in the mesolimbic system, and on the modulation of glutamatergic transmission in the olfactory tubercle. He was elected a Member in 1985.

A. Murray Harper

( 1933 - 2005 )

Murray Harper was a physiologist of major international and historical significance with a lifelong interest in cerebral blood flow. A Glasgow medical graduate, he led a research team there at the Wellcome Surgical Institute to investigate brain blood flow and cardiopulmonary bypass, ultimately becoming Professor of Surgical Physiology. A champion of interdisciplinary research, basic and clinical, he founded the International Society of Cerebral Blood Flow and Metabolism and was the first editor of its journal.

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Geoffrey Wingfield Harris

(1913 - 1971)

Geoffrey Wingfield Harris (1913 – 1971) was an English physiologist and fellow of the Royal Society, considered by many to be the father of neuroendocrinology. He published the ‘Neural Control of the Pituitary Gland’ in 1955 which predicted the subsequently discovered hormone ‘releasing factors’ acting on the hypothalamus. He was a demonstrator in Anatomy and then a lecturer in Physiology at Cambridge before working as a neuroendocrinologist at the Maudsley Hospital in London. Harris went to Oxford University in 1962 as Professor of Anatomy and a Fellow of Hertford College. He was an effective and popular teacher of anatomy, raising its profile, and contributing to the development of the new Physiological Sciences Final Honour School, which brought together for the first time, the five preclinical departments of anatomy, biochemistry, pathology, pharmacology and physiology. His teaching extended to include endocrinology as a special subject. In the same year he was appointed Honorary Director of the Medical Research Council’s Neuroendocrinology Research Unit in Oxford. Here he continued his scientific research: attempting to isolate the luteinizing hormone releasing factor, and studying the effect of gonadal hormones on the sexual differentiation of the brain. He also continued to practise clinical medicine at the Littlemore Hospital, where he was Honorary Consultant, investigating gonadal and pituitary hormones in psychiatric patients. In 1986, The Physiological Society instituted a triennial lecture in memory of the late Professor G. W. Harris.

Geoffrey Wingfield Harris

(1913 - 1971)

Geoffrey Wingfield Harris was an English physiologist and fellow of the Royal Society, considered by many to be the father of neuroendocrinology. He published the ‘Neural Control of the Pituitary Gland’ in 1955 which predicted the subsequently discovered hormone ‘releasing factors’ acting on the hypothalamus. He was a demonstrator in Anatomy and then a lecturer in Physiology at Cambridge before working as a neuroendocrinologist at the Maudsley Hospital in London.

Harris went to Oxford University in 1962 as Professor of Anatomy and a Fellow of Hertford College. He was an effective and popular teacher of anatomy, raising its profile, and contributing to the development of the new Physiological Sciences Final Honour School, which brought together for the first time, the five preclinical departments of anatomy, biochemistry, pathology, pharmacology and physiology. His teaching extended to include endocrinology as a special subject. In the same year he was appointed Honorary Director of the Medical Research Council’s Neuroendocrinology Research Unit in Oxford. Here he continued his scientific research: attempting to isolate the luteinizing hormone releasing factor, and studying the effect of gonadal hormones on the sexual differentiation of the brain. He also continued to practise clinical medicine at the Littlemore Hospital, where he was Honorary Consultant, investigating gonadal and pituitary hormones in psychiatric patients. In 1986, The Physiological Society instituted a triennial lecture in memory of the late Professor G. W. Harris.

David Keynes Hill

( 1915 - 2002 )

David Hill, son of A V Hill, was a physiologist and biophysicist whose principle interest was muscle contraction. A graduate of Trinity College, Cambridge, he started research on muscle there and was quickly appointed a research fellow. During the Second World War, he worked on crush injuries and ballistics, returning afterwards to his muscle research, first at Cambridge, then (1948–49) as physiologist at the laboratory of the Marine Biological Association in Plymouth, and finally as biophysicist back in Hammersmith (vice-dean, 1970–76). He used innovative techniques to investigate intracellular phenomena and, from 1971, collaborated with RHT Edwards and subsequently with PA Merton.

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Harold Hillman

d. 2016

Harold Hillman joined the Society in 1968. He spent most of his career at the University of Surrey. He was known as a critic of modern methods of electron microscopy and for his writing on methods of resuscitation and humane execution techniques. Read more about his life and achievements in obituaries by the Guardian and Telegraph.

Sidney Montagu Hilton

( 1921 - 2011 )

Sidney Montague Hilton was Bowman Professor and Head of the Department of Physiology at the Medical School in Birmingham from 1965 to 1983. He was a distinguished physiologist who made leading contributions to our understanding of brain control of stress-induced cardiovascular changes. He studied medicine at Cambridge and Guy’s Hospital, London. Following National Service at the Institute of Aviation Medicine, he worked in Professor Adrian’s Cambridge laboratory and then under Professor Feldberg at the Medical Research Council (MRC), London. After initial studies on vasodilatory mechanisms in muscle and salivary glands, he turned to the animal defense response – establishing the then revolutionary notion that blood pressure was not controlled by a single brain centre under these circumstances. He developed this work further in Warsaw and England, with Andrzej Zbrożyna of the Nencki Institute of Experimental Biology of the Polish Academy of Sciences. On his appointment to Birmingham, he built a department with an international research reputation and helped to establish the Physiology Department in the new Medical School in Salisbury, Southern Rhodesia (Harare, Zimbabwe). He was elected an Honorary Member of The Physiology Society in 2004.

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Sir Alan Hodgkin

( 1914 - 1998 )

Alan Hodgkin was distinguished for his contributions to the understanding of nerve conduction, the excitation of muscle, and phototransduction in the retina. With his colleagues Bernard Katz and Andrew Huxley, he established the ionic basis of the action potential, revolutionising neuroscience and laying the foundations for subsequent work on ion channels. In 1963, at the age of 49, he won the Nobel Prize for Physiology and Medicine with Andrew Huxley and John Eccles. He was a Fellow of the Royal Society at the age of 40, and held its Foulerton Research Chair from 1952 to 1969. In 1970 he succeeded F J W Roughton to the John Humphrey Plummer Chair of Biophysics in the University of Cambridge. He was President of the Royal Society from 1970 to 1975, its Croonian Lecturer in 1957, and winner of its Royal Medal in 1958 and its Copley Medal in 1965. He was Master of Trinity College from 1978 to 1984, President of the Marine Biological Association from 1966 to 1976, and Chancellor of the University of Leicester from 1971 to 1984. He served on the Committee of The Physiological Society from 1949 to 1953 and again as Foreign Secretary from 1961 to 1967. He gave the Annual Review Lecture in the Society’s Centenary year (1976) and became an Honorary Member in 1979. He was made KBE in 1972, and a Member of the Order of Merit in 1973.

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Oliver Holmes

( 1933 - 2004 )

Oliver Holmes was a BSc student at University College in the 1950s and subsequently Bayliss- Starling Scholar, studying for his MSc with GL Brown and completing his medical degree and pre-registration posts. His research focus was experimental epilepsy, as MRC Fellow with GD Dawson at the Institute of Psychiatry, then Senior Lecturer at Leicester, Special Wellcome Research Fellow to the Royal Postgraduate Medical School with Professor Sir Gordon Robson and finally, at the Institute of Physiology in Glasgow (from 1975). To enhance this research, he studied computer programming and mathematics and collaborated widely, contributing significantly to basic epileptology. In later life, he turned to work on gastric secretion and served the Physiological Society in several roles. After retiring as Senior Lecturer in 1998, he remained Honorary Research Fellow in the Faculty of Biomedical and Life Sciences, University of Glasgow.

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Sir Gabriel Horn

d. 2012

Sir Gabriel Horn, a Member of The Society from 1963, has died aged 85. Gabriel was one of the outstanding neuroscientists of his generation. He was responsible for numerous advances in neuroscience encompassing diverse areas, but especially concerning neural mechanisms of learning and memory. He was a most gifted teacher, delivering lectures appreciated by generations of students. He will be remembered by many students, friends and colleagues for the generous warmth of his personality.

David Horrobin

( 1939 - 2003 )

David Horrobin, appointed Professor of Medical Physiology in Nairobi at the age of 31, had research interests in polyunsaturated fatty acids and schizophrenia. After retiring as Professor of Medicine in Montreal in 1979, he became a controversial pharmaceutical scientist/entrepreneur, founding Scotia Holdings, then Laxdale Ltd.

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David Hubel

( 1926 - 2013 )

David H. Hubel was the John Franklin Enders University Professor at Harvard Medical School. Having attended McGill University Medical School he qualified as an MD in 1951. His research pioneered the understanding of the cerebral cortex, and knowledge in the fields of colour vision, motion perception and stereoscopic vision. In 1981 he won the Nobel Prize for Physiology or Medicine, shared with Torsten Wiesel and Roger Sperry. Professor Hubel was President of the Society for Neuroscience, and an Honorary Member of both The Physiological Society and the American Neurological Association. He was also a member of the American Academy of Arts and Sciences.

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Olga Hudlicka

d. 2014

The Society is sorry to hear of the death on 3 May of Olga Hudlicka, Professor Emeritus at the University of Birmingham. She had been a member of The Society since 1972. She died at the age of 87 and had continued working in the School of Clinical and Experimental Medicine until quite recently. She worked on factors influencing blood flow and capillary growth in normal and ischaemic skeletal and cardiac muscle and gave the Annual Review Prize Lecture in 1990.

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Sir Andrew Huxley

d. 2012

Sir Andrew Huxley OM FRS, one of the greatest physiologists of his generation, has died aged 94. Andrew, together with Alan Hodgkin, determined the basis for nerve cell excitability, applying the technique of the voltage clamp to identify the ionic currents in the squid axon. This pioneering research was carried out at the Marine Biological Association in Plymouth and at the Physiological Laboratory in Cambridge. Published exactly 60 years ago, the work was a fundamental breakthrough and led to the subsequent understanding of voltage-gated ion channels, the understanding of propagating action potentials and provided the framework for studying and analysing ion channel kinetics. With Alan Hodgkin and John Eccles, Andrew shared the 1963 Nobel Prize for Physiology or Medicine for this work. In his subsequent work, carried out largely at UCL, he developed the cross-bridge theory of muscle contraction. Andrew was elected a Fellow of the Royal Society in 1955 and was knighted in 1974 and subsequently appointed to the Order of Merit in 1983. He served as President of the Royal Society from 1980 to 1985.

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Ainsley Iggo

d. 2012

Ainsley Iggo was elected a Member in 1956. He was a member of the Editorial Board of The Journal of Physiology from 1962 to 1969, and a member of the Editorial Board of The Quarterly Journal of Experimental Physiology from 1980 to 1983.

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Fred Imms

d. 2012

Fred Imms was a familiar figure at Society meetings for many years. He was an applied human physiologist who worked at Guys Hospital Medical School and then, following merger, at the GKT School of Medicine.

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Douglas L Ingram

( 1929 - 2006 )

Doug Ingram’s research career spanned four decades during which time he made major contributions to the fields of reproductive physiology, thermal and environmental physiology, and nutritional and developmental physiology. A zoology undergraduate in Birmingham under Peter Medawar, then a PhD student in anatomy with Solly Zuckerman, the major part of Doug’s research achievement occurred between 1961 and 1989 at the ARC Institute of Animal Physiology, Cambridge/BBSRC Babraham Institute, in the Departments of Applied Biology, Cell Biology, and Molecular and Cellular Physiology. He was awarded Individual Merit Promotion to Senior Principal Scientific Officer in 1975, a DSc by the University of Birmingham and elected a Fellow of the Institute of Biology in 1976. Along with numerous publications in learned journals, including The Journal of Physiology, he was joint author of the definitive book on Man and animals in hot environments (1975).

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Colin Ingram

d. 2013

The Society is sorry to hear of the death of Colin Ingram, who was a Member from 1994 to 2008. He was Professor of Psychobiology in the School of Neurology, Neurobiology & Psychiatry from 2000 and Director of the Institute of Neuroscience from 2004 at the University of Newcastle upon Tyne. He elucidated the neural mechanisms controlling the stress response and led research into neuroinformatics and translational neuroscience. He died on 15 December.

Werner Ulrich Jacobson

( 1906 - 2000 )

Werner Jacobson graduated in Heidelberg in 1930 and moved to England in 1933. Honor Fell, the great cell biologist, recruited him to work at the Strangeways Laboratory in Cambridge. Doctorates in philosophy and of science followed in 1940 and 1960, and two sojourns in Harvard Medical School. In 1980, he was appointed Halley Stewart Professor of Experimental Medicine, first in the University Department of Haematology and later in that of Paediatrics in the Clinical School at Addenbrooke’s Hospital. His research was wide-ranging and he was an early and insistent advocate for the clinical significance of folic acid deficiency.

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Emilie Leslie Jervis

( 1927 - 2002 )

Emiline Jervis worked in Sheffield University Department of Physiology under the direction of Professor David Smyth as research assistant, lecturer and senior lecturer, investigating intestinal transport mechanisms of hexoses and amino acids. She also administered the dental student physiology course for many years.

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David Jordan

( 1953 - 2007 )

David Jordan studied at the University of Birmingham receiving a BSc in Biological Sciences (1974) and a PhD (1977) for his studies on The termination and excitability of sinus nerve afferents. He became research fellow in the Physiology Department, then physiology lecturer at the Royal Free Hospital School of Medicine, with subsequent promotion to senior lecturer (1988), Reader (1993), Professor (1999) and Head of Department. He collaborated over several years with his PhD supervisor, Mike Spyer, and subsequently with Andrew Ramage, funded by the Wellcome Trust and British Heart Foundation, The papers that flowed from their partnership have significantly increased our understanding of the role of 5-hydroxytryptamine receptors in the central nervous control of cardiovascular function. Author of several review articles and book chapters, he had many editorial, educational and administrative roles. He served as an undergraduate and higher degree external examiner both within the UK and overseas.

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C T Kappagoda

( 1943 - 2015 )

Tissa Kappagoda was Professor of Medicine in the Division of Cardiovascular Medicine at the University of California, Davis. He qualified in Medicine at the University of Ceylon (1965), moving to England the following year. He held junior hospital appointments in the National Health Service before joining the Cardiovascular Unit in the Department of Physiology at Leeds University. His PhD thesis was on the function of atrial receptors, the subject of a series of papers in The Journal of Physiology and Experimental Physiology as well as in The Society’s publication, Atrial Receptors, (with RJ Linden). He developed a method for measuring oxygen uptake during exercise. Kappagoda left Leeds in 1978 and became Research Professor of Medicine and Director of Cardiac Rehabilitation at the University of Alberta in Canada, where his team worked on aspects of cardiovascular physiology, including pulmonary receptors in heart failure and exercise training in ischaemic heart disease. He was then appointed Professor of Medicine, Division of Cardiovascular Medicine, at the University of California, Davis and Director of the Preventive Cardiology Programs at the University of California Medical Center. Here, his research focused on the role of endothelial cells in regulating vascular tone and the effects of fatty acids and polyphenolic products present in plants in human and animal models. A caring physician, his contribution to cardiovascular science included over 200 peer-reviewed publications.

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Bernard Katz

(1911 –2003)

Sir Bernard Katz was a German-born Nobel Prize winning physiologist and an icon of post-war biophysics. Katz was born to a Jewish family in Leipzig. After studying medicine at the University of Leipzig, he fled to England in 1935 to escape Nazi persecution. He went on to work at University College London (UCL) under AV Hill, and on finishing his PhD in 1938, won the Carnegie Fellowship to study with John Eccles at the Kanematsu Memorial Institute in Sydney. After being naturalised in 1941, Katz served in the Royal Australian Air Force for the duration of World War II. In 1946, he returned to UCL and continued his research on the synapse.

Katz became a Henry Head Fellow of the Royal Society in 1952, the same year in which he became Head of the newly designated Biophysics Department at UCL. He was knighted in 1969, shortly before receiving the Nobel Prize for Physiology or Medicine in 1970 (with Julius Axelrod and Ulf von Euler), for his discovery that neurotransmitter release at synapses is ‘quantal’, meaning that the amount of neurotransmitter release is never less than a fixed minimum and, if more, this is an integer multiple of this amount. This functional evidence was later complemented by ultrastructural evidence of transmitter-containing, membrane-bound vesicles common to nerve endings and secretory cells. Katz became Emeritus Professor at UCL in 1978.

On 20 April 2003, Sir Bernard Katz died aged 92. In 1999 The Physiological Society established the Hodgkin-Huxley-Katz Prize Lecture. This prestigious biennial lecture celebrates the contributions to the physiological sciences of Alan Hodgkin, Andrew Huxley and Bernard Katz.

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Michael Keating

( 1940 - 1998 )

Michael Keating did pioneering work on the role of activity in determining the specificity of neural connections in the vertebrate brain. With R.M.Gaze and others, he studied visual and auditory functional adaptation, becoming head of the Division of Neurophysiology at the National Institute for Medical Research in Mill Hill in 1976.

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William Keatinge

( 1931 - 2008 )

William Richard Keatinge (Bill) studied medicine at Cambridge University and St Thomas’s Hospital in London. He was Director of Studies in Medicine at Pembroke College Cambridge (1958-60) and, after an interval as a Fullbright scholar, subsequently held an MRC post there (1961- 68). He joined the London Hospital Medical College (LHMC) as Reader in Physiology (1969) and was soon promoted to a Personal Chair (1970), head of department (1981) and Head of Physiology in the joint school (1990) after LHMC and St Bartholomew’s Hospital Medical College (Barts) merged with Queen Mary and Westfield College (QMW). A great international collaborator, especially in Russia, much of his research addressed temperature regulation and the control of blood vessels. During challenging times at his academic institution, he served in many senior administrative positions and was appointed Emeritus Professor on retirement in 1995.

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Gerald A. Kerkut

( 1927 - 2004 )

Gerald Kerkut was a natural sciences undergraduate, and then zoology PhD student, at Cambridge. His subsequent work on slug ganglia took him to Southampton as lecturer. There, he helped establish the Department of Physiology and Biochemistry (1959), was appointed the second Professor of Physiology and Biochemistry at Southampton (1966) and later served as Dean of Science, Chairman of the School of Biochemical and Physiological Sciences and Head of the Department of Neurophysiology. During three decades of research on invertebrate models he made several major advances in neurophysiology, including the development of an electrode for intracellular chloride measurement. Later, he worked on dorsal horn field potentials using isolated mammalian CNS preparations. Editor of more than fifty volumes, his publications ranged from a complete revision of a standard text on invertebrates to his own: ‘The Implications of Evolution’. He was founder (1960) and editor of Comparative Biochemistry and Physiology, editor of Progress in Neurobiology and, at one stage, had his own publishing company – Scientechnica. In his last years, his website attracted up to 60,000 hits/week with articles on politics, student concerns, money and education – as well as science.

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Roderick P Kernan

d. 2014

The Society is sorry to hear of the death on 10 January of Roderick P (Roddy) Kernan, who was a member since 1963. He was Emeritus Professor of Physiology at University College Dublin.

David McKie Kerslake

( 1923 - 2005 )

David Kerslake made major contributions to the field of human thermal physiology. Medically qualified in 1946, he conducted his PhD research during RAF National Service at the Institute of Aviation Medicine (IAM). Appointed Officer in Charge of the IAM Climatic Section (1950), he was soon involved in developing the Institute’s climatic research facility. He investigated the physiological effects of thermal stresses on service personnel, publishing many papers in The Journal of Physiology and, later, a Monograph of The Physiological Society, The Stress of Hot Environments (1972). He was awarded an OBE (1962) and the Vernon Prize of the National Institute of Industrial Psychology (1966). Eschewing further promotion, he remained IAM Deputy Chief Scientific Officer until his retirement in 1978.

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Richard Darwin Keynes

( 1919 - 2010 )

Richard Keynes was an eminent cell physiologist. His undergraduate studies at Cambridge were interrupted by work on sonar and radar during the Second World War, after which he became research fellow at Trinity College, teaching fellow at Peterhouse and University Lecturer in Physiology (1948-60). He then joined the Agricultural Research Council Institute of Animal Physiology at Babraham, becoming Director in 1965. With Peter Lewis, he applied novel radioactive tracer techniques to measure ionic movements across membranes – work that directly supported Hodgkin and Huxley’s subsequent Nobel prize discoveries. He investigated ion transport in secretory epithelia and, fascinated by South America, studied the Electroporus electric organ, whose electrophysiological mechanism he had clarified early in his career. He published books on Darwin’s zoological research and served on Galapagos conservation bodies. Secretary General (1972), and then President (1981–84), of the International Union for Pure and Applied Biophysics, he helped establish the ICSU/Unesco International Biosciences Networks, later becoming its Chairman (1982–1993). He received the Order of Scientific Merit (Brazil) and Honorary Membership of the Latin American Academy of Sciences amongst many academic honours, and was awarded the CBE in 1984.

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Hirosi Kuriyama

( 1928 - 2003 )

Hirosi Kuriyama made immense contributions to our knowledge of smooth muscle, his lifelong research interest. He graduated in medicine from Fukuoka in 1951. After starting his career in Japanese physiology departments, he worked with Csapo in New York, using microelectrodes to record electrical activity in uterine smooth muscle. He subsequently completed an Oxford DPhil under Edith Bulbring, publishing work on the electrophysiology of guinea pig taenia coli. In 1964, he returned to Kyushu University and was later appointed head of the Department of Pharmacology, Faculty of Medicine (1976). Between 1970 and 1990, his group produced the majority of electrophysiological and physiological studies on smooth muscle which were published at the time. Many of his co-workers and assistants became heads of department in Japanese universities. He retired from Kyushu in 1992, worked briefly with Chugai Pharmaceutical Company, then joined Seinan Jo Gakuin University for nine years, eventually becoming President there.

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Max Lab

d. 2016

Professor Max Lab, an influential Physiologist working in the UK, sadly passed away in January. He has been affiliated to Imperial College for most of his career and we would like to organise a one day symposium in his honor, and celebrate his scientific discoveries. Max was an extremely popular figure in the Physiology community.

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Joe Lamb

d. 2015

Joe, JF Lamb, was born in 1928, formerly Chandos Professor of Physiology at St Andrews University and later headed the Rowett Research Institute at the University of Aberdeen. He was an Honorary Member (2005), a member since 1963 and had been The Society Secretary, 1982-5. Joe served on the Journal of Physiology Editorial Board and as Physiology News Editor.

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Yves Laporte

d. 2012

Yves Laporte, one of the most eminent physiologists of the second half of the 20th century, has died in Paris, aged 91. Yves was a great admirer and supporter of The Physiological Society, after he was introduced to it by David Barker in the early 60s.

Paul Lauterbur

( 1929 - 2007 )

Paul Lauterbur published the first magnetic resonance image in a short letter to Nature in 1973 (242, 190-191). The invention of MRI won him many prizes and awards, culminating in the 2003 Nobel Prize for Physiology or Medicine jointly with the British physicist Peter Mansfield. Having gained his Bachelor degree from Case Institute of Technology, he was drafted into the US army where he used an NMR spectrometer for the first time and published several papers. After military service, he completed graduate school and became an Associate Professor at the State University of New York (SUNY), where his Stony Brook laboratory became a focus for others interested in the idea of imaging with NMR.

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Peter Raymond Lewis

( 1924 - 2007 )

Peter Lewis followed undergraduate chemistry at Oxford with a DPhil thesis on the kinetics of bacterial growth. He joined the Cambridge Physiological Laboratory (1948), working on sodium and potassium flux in squid nerves with Richard Keynes. He studied diurnal rhythms on MRC-funded research trips to Spitzbergen. Working with the histologist, Charles Shute, he then turned to the cholinesterases that became central to his career, publishing landmark papers in Brain (1967) on cholinesterase-containing fibres in the rat brain. He continued his cholinergic work when he returned to The Physiological Laboratory (1970), with additional wide-ranging research interests including spectral sensitivity curves, monoamines, placental esterases and weeping lubrication in mammalian joints. A medical society founded (2007) in his college, Corpus Christi, was named the Lewis Society of Medicine by unanimous decision.

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Malcolm Lidierth

(1961-2019)

The Society is saddened to hear of the death of Member Malcolm Lidierth at the age of 58. A neurophysiologist with a rare talent for in vivo recording, he completed a PhD with David Armstrong in Bristol before becoming a lecturer in 1990 at the Sherrington School of Physiology at UMDS, later part of King’s College London. As well as collaborating with Pat Wall on studies of the dorsal horn and Lissauer tract, Malcolm had a particular flair for developing physiological analysis software, which he made freely available to others and thus it was widely used. An innovative and conscientious teacher, he ran a highly popular 3rd year neuroscience module for many years.

Ronald James Linden

( 1920 - 2010 )

Ron Linden founded a dynasty of Leeds-trained cardiovascular physiologists who have spread far and wide. A graduate of Leeds Medical School, he was awarded MB CHB (1951) and PhD (1958). He was then appointed research fellow at the National Heart Institute in Washington DC, returning to Leeds for his DSc (1965). In 1966 he was awarded a Personal Chair and embarked on the creation of the Cardiovascular Unit at Leeds, within the Department of Physiology. In 1977 he was appointed head of a separate Department of Cardiovascular Studies, his chair being one of the first endowed by the British Heart Foundation. He served on several editorial boards and was a member or chairman of several committees giving advice to government departments on physiological topics. He served on the University Grants Committee, and committees of the British Heart Foundation and the Ministry of Defence. Following retirement in 1985, he continued research collaboration with Gianni Losano at the University of Turin, from whom he received an honorary degree in Medicine and Surgery in 1993.

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Olof Lippold

d. 2016

Olof Lippold (1923-2016) was born in London to parents of Swedish and German ancestry. He studied medicine at University College London during WW2, qualifying in 1946. He became a member of The Physiological Society in 1953. After initial clinical work, he joined the Physiology Department at UCL where worked until 1983, then moving to Royal Holloway as Head of Department. He researched in areas as varied as tetanus toxin action and muscle spindle excitation, the latter led to work on the effects of Direct Current on long-term brain function. He observed that the EEG alpha rhythm originated from ocular muscle activity rather than the underlying cortex. And he worked on Parkinsonism.

Olof was also a renowned teacher, revealed to a wider audience through successful textbooks such as ‘Human Physiology’ (a title taken over from Winton & Bayliss), ‘Respiratory Physiology’, ‘Neurophysiology of the Cerebral Cortex’ (with Lynn Bindmann) and ‘Physiology Illustrated’ (with his daughter, Barbara Cogdell). Amongst his many accomplishments, Olof was also a talented painter and pianist.

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Alex Livingston

d. 2014

The Society is sorry to hear of the death of Alex Livingston, of the School of Veterinary Medicine at the University of Saskatchewan, Canada. He had been a member of The Society since 1998.

Margaret Lowrie

d. 2017

Margaret Lowrie gave her first communication to The Physiological Society in 1980 and joined the Society in 1986. In the 1980s and 1990s she worked with Professor Gerta Vrbova at University College London on nerve injury and she subsequently moved to Imperial College.

Hans-Cristoph Lüttagu

d. 2017

We are saddened to report that Hans-Christoph Lüttgau passed away on 5th November at the age of 91. He was an eminent physiologist who has made lasting contributions to our understanding of nerve function and the intricate events between excitation and contraction in muscle. He had been the founding Professor of Cell Physology at Ruhr University, Bochum in Germany, an Editor of The Journal of Physiology, a member of The Society since 1965 and an Honorary Member since 1997.

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Beverley R Mackenna

d. 2012

Bev has died at the age of 83. He became a Society Member in 1964. Recruited to the Physiology Department at Glasgow University by Robert Campbell Gary, much of his career at Glasgow University was devoted to the professional courses in Medicine and in Dentistry. He revised later editions of the successful textbook ‘Illustrated Physiology’ with Robin Callander. A calm and reserved character, Bev’s quiet professionalism and personal charm will be sadly missed by his many students and colleagues.

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Laurence Malcolm

( 1913 - 2001 )

Laurence Malcolm was the son of New Zealand physiologist Professor John Malcolm. His early career was spent in Otago, London and New York. He joined the National Institute for Medical Research, Mill Hill, in 1953 to study evoked potentials in spinal cord and brain. In 1959 he was awarded the Regius Chair in Physiology at Marischal College, University of Aberdeen. He served as Dean of the Science Faculty there in the early 1970s, introducing several notable teaching innovations.

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Archie McIntyre

( 1913 - 2002 )

Archie McIntyre was one of the founders of modern neuroscience in Australia. A medical graduate from Sydney University, his early research addressed the physiology of eye movements, uterine muscle and aviation. After the war, as a Rockefeller Fellow, he studied tendon reflexes with Herbert Gasser in New York, continuing this work as Nuffield scholar in Cambridge. He then accepted Jack Eccles’s offer of a senior lectureship in the physiology department at the University of Otago, becoming Head of Department there in 1952. In 1962 he was appointed professor of physiology at the new Monash University, Melbourne, and subsequently held a number of key national positions in Australian scientific institutions. He is best known for his work in neuroscience, publishing three landmark papers in 1960 on cutaneous sensory receptors. He is widely credited with designing the microelectrode equipment used in the work that brought Eccles the Nobel Prize in 1970.

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Hans Meves

( 1925 - 2008 )

Hans Meves qualified in medicine from the University of Marburg (1951). He worked on the physiology of nerve fibres with Hans Lullies at Saar University and then in Kiel, with B Frankenhauser in Stockholm (1957), and with R Stampfli in Homburg (1959 and 1961-7). Between 1962 and 1965, Meves collaborated with Alan Hodgkin and others on the effects of low ionic strength on action potentials. He was appointed Professor of Physiology in Kiel (1967) but left three years later for the Marine Biological Association, eventually becoming Senior Principal Scientific Officer. His work mostly addressed the selectivity (structure) and activation/inactivation mechanisms (gating) of ion channels, with several papers appearing in The Journal of Physiology. In 1980, Meves returned to succeed R Stampfli in the Chair of Physiology at Homburg. In 1990, he retired but continued to experiment, learning the patch clamp technique to do so. He became a Member of The Physiological Society in 1965 and was a member of the Editorial Board of The Journal of Physiology from 1975 to 1984.

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Ricardo Miledi

d. 2017

The Society is much saddened to hear of the death of Ricardo Miledi at the age of 90. Educated in Mexico and a skilled experimenter, he and Bernard Katz laid the foundations of our understanding of the role of calcium in synaptic neurotransmitter release. As well as making many basic discoveries on the acetylcholine receptor channel (including the first description of ‘acetylcholine noise’ with Katz), he was one of the early developers of the use of oocytes for drug-receptor expression studies. Succeeding Katz as head of the Department of Biophysics at UCL, he moved to the University of California Irvine in 1986. Amongst other awards, Ricardo Miledi held the Royal Society’s Fullerton Chair whilst at UCL and received its Royal Medal in 1998.

Peter Mott

d. 2014

The Society is saddened to learn of the death of Peter Mott on 14 March; he was 84. Peter worked at Cambridge University Press and was responsible for overseeing the publication of many journals including The Journal of Physiology and the meetings abstracts. When The Society took over the ailing Quarterly Journal of Experimental Physiology (now Experimental Physiology), Peter contributed to discussions on the change to monthly publication with a new title and cover design.

Vernon Mountcastle

d. 2015

The Society is sorry to hear of the death of Dr Vernon Mountcastle, who was an Honorary Member of The Society, elected in 1986. He passed away in January.

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Rolf Niedergerke

d. 2011

Rolf Niedergerke (1921–2011) worked principally to elucidate the links between excitation and contraction in muscle, especially cardiac muscle.

His 1954 Nature paper with Andrew Huxley – the first by either on muscle – described in clear terms the sliding filament mechanism of muscle contraction, now acknowledged as the paradigm for all forms of cellular motility. He was both a consummate experimentalist and a scrupulous reporter of his science.

Rolf was elected a Member of The Society in 1959 and an Honorary Member in 1987.

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Syogoro Nishi

d. 2014

The Society is sorry to hear of the death of Syogoro Nishi on 13 February. He was an Honorary Member of the Society, elected in 1999.

Robert O Nneli

d. 2017

We are saddened to hear that Professor Robert O. Nneli, the Dean of the School of Basic Medical Sciences at the Federal University, Ndufu-Alike Ebonyi, died in a motor accident in April 2017 at the age of 69. He joined the university in 2012 after spending close to 20 years at Abia State University. He was educated at St. Patrick’s Secondary School Emene in Enugu and the University of Calabar. His physiological work covered a wide range of issues on cardiovascular and occular problems in the Nigerian population.

Susan Noble

d. 2015

Dr Susan Noble passed away in the John Radcliffe Hospital early in the morning of Sunday 4 October 2015, at the age of 75. She worked in the Department of Physiology, Anatomy and Genetics at The University of Oxford as a researcher and lecturer between 1969 and 1997. She published more than 50 papers while working in the Department, including the 1979 article in Nature with Drs Hilary Brown and Dario DiFrancesco that identified the ‘funny’ pacemaker current, and that led to the development of ivabradine (Servier). She was a Member since 1976.

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Stephen O’Neill

d. 2012

Stephen O’Neill, a cardiac physiologist and Member of The Physiological Society since 1990, had died aged just 49 after maintaining a positive attitude since being diagnosed with a brain tumour six years ago. Stephen continued working and publishing until a matter of weeks before passing away. He has left warm and lasting memories of an excellent physiologist and loyal friend.

Autur Paintal

( 1925 - 2004 )

Autar Paintal was a distinguished physiologist from India. With an MD from Lucknow University, he came to Edinburgh as a Rockefeller Fellow in 1951 and conducted PhD research there under David Whitteridge. He established that it was possible to record from nerve fibres, and dissect them under liquid paraffin – a major advance in experimental technique. His research also shed light on many aspects of vagal innervation and included the discovery, in 1955, of the J receptor in the lungs. He subsequently returned to a research position in Kanpur, at the Indian Army Medical Centre, and later became head of the Indian MRC.

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John Richard Pappenheimer

( 1915 - 2007 )

John Pappenheimer’s work in capillary permeability and molecular sieving are classics in physiological literature. He contributed valuable research to a wide range of disciplines within physiology: capillary permeability, respiratory physiology, blood-brain CSF transport, the neurochemical aspects of sleep, and most recently to the understanding of the absorption of sugars and amino acids in the intestine. Born in NYC, Pappenheimer attended the Lincoln and Loomis schools. He received a BS from Harvard College and a PhD from Clare College, Cambridge, England. In 1953 he was awarded the lifetime Career Investigator position from the American Heart Association for his work on capillary permeability and in respiratory physiology. He was appointed the George Higginson Professor at Harvard Medical School in 1969. He was a member of the American Physiological Society (President 1964-1965), the Academy of Arts and Sciences, the National Academy of Sciences and Honorary Member of The Physiological Society. He published numerous articles of original work over a span of 70 years. He collaborated with colleagues from around the world, including Frank Winton, Glen Millikan, Bjorn Folkow, Manfred Karnofsky, JM Krueger, J Madara and Charles Michel.

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James Edward Pascoe

( 1924 - 2011 )

Jim Pascoe was a Reader in Physiology at University College London. Having studied pharmacy at Plymouth College and the School of Pharmacy, he became a junior lecturer at the latter. He then studied Physiology, coming to UCL at the invitation of G.L.Brown, the head of department, and then joining the staff. Elected a Member of The Physiological Society (1954), he joined the Committee in 1979. He worked with Tim Biscoe, writing computer software to enable neurophysiological research, and was closely involved in parliamentary discussions about vivisection legislation in the 1980s.

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Sir William Drummond Macdonald Paton

(1917 –1993)

Sir William Drummond Macdonald Paton (1917 –1993), always known as Bill Paton, was an English physiologist, pharmacologist and Fellow of the Royal Society, considered by many to be one of the world’s greatest pharmacologists. He was responsible for discovering two new classes of drug that acted on nicotinic acetylcholine receptors. His theorised multiple types of nicotinic receptor (confirmed in the 1970s) formed the foundation of the development of Decamethonium, the first specific neuromuscular blocking drug and Hexamethonium, the first drug that specifically and safely lowered blood pressure. Paton was also charged with finding the solution to the problem of convulsions suffered by deep-sea divers if they went more than 200ft below sea-level, having discovered that the high pressure causing the convulsions could be reversed with anaesthetics. He was awarded a CBE in 1968 and knighted in 1979 for his work. Paton not only made countless discoveries but was also heavily involved in numerous public committees and had a special interest in the history of medicine. He made a substantial donation to The Society that founded the Paton Prize Fund for historical research on physiology and physiologists. Paton was Honorary Director of the Wellcome Institute for History of Medicine from 1983 to 1987. Sir William Drummold Macdonald Paton died on 17 October 1993. In 1994, The Physiological Society introduced the Paton Prize Lecture, this annual lecture commemorates Paton’s support and initiatives for promoting interest in the history of scientific experiments and ideas.

John Patterson

d. 2018

It is with sadness that The Society reports the death of John Patterson on 25 February 2018. John was an Honorary Senior Lecturer in Medical Education (formerly Associate Dean, Undergraduate Medical Studies) at Barts and the Royal London, Queen Mary University (1978-2009). John taught physiology for 30 years and was three times elected ‘best preclinical teacher’. In his role as associate Dean for Undergraduate Medical Studies, he oversaw a revision of assessment structures across the programmes and as Head of Assessment he had oversight of the design, delivery and analysis of all MBBS examinations. More recently John was appointed as psychometrician and then, additionally, as Vice-Chairman of the Examinations Board, at the Society of Apothecaries of London.

Sir Stanley Peart

(1922-2019)

The Society notes with regret the death on 14 March of Honorary Member Sir Stanley Peart FRS, FMedSci, FRCP at the age of 96. Trained in medicine at St Mary’s Hospital Medical School in London, where he later served as Professor of Medicine, Peart was the first to purify and determine the structure of the peptide hormone angiotensin and to isolate renin. A trustee  of the Wellcome Trust from 1975, he headed the clinical research panel which was instrumental in promoting the beginnings of the human genome project and neuroimaging research. He became a Society Member in 1953 and was knighted in 1985.

Christopher Peers

d. 2018

It is with sadness that The Society reports the death of Christopher (Chris) Peers on 20 May 2018. Having obtained a degree in physiology and a PhD in pharmacology from RFHMS, University of London, a period of postdoctoral research at Oxford was followed by his appointment as Lecturer in Pharmacology at the University of Leeds in 1994. From then on, an exceptionally high level of research quality and productivity led to rapid career progression and promotion to Professor of Cellular Physiology in 2002. Chris will be remembered as a remarkably enthusiastic and gifted researcher with a warmth of personality that ensured numerous enduring collaborations.

Mary Pickford

( 1902 - 2002 )

Born in India, Mary Pickford graduated in science from Bedford College, London, in 1924. While conducting pharmacological research part-time at UCL, with AJ Clarke and EB Verney, she trained in medicine and qualified in 1933. After junior clinical jobs, she took a post as physiology lecturer at Edinburgh University, where she studied oxytocin and anti-diuretic hormone in dogs. She gained a DSc in 1951 (Edinburgh) and later was given an Honorary DSc by the Heriot-Watt University. In 1954 she became a Fellow of the Royal Society of Edinburgh, in 1977 of the Royal College of Physicians of Edinburgh, and in 1966 of the Royal Society of London. In 1966, she was the first woman to hold a chair in the Edinburgh Medical Faculty. She published extensively and in 1972 was appointed Special Professor of Endocrinology at Nottingham University.

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Vernon Pickles

d. 2012

Vernon Rycroft Pickles died on 27 April 2012. He became a Member in 1950.

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Philip Poole-Wilson

( 1943 - 2009 )

Philip Poole-Wilson studied at Cambridge and then St Thomas’ Hospital Medical School, qualifying in 1967 and joining the academic department of medicine. In 1973 he was awarded a British Heart Foundation (BHF) Fellowship to the ‘Heart Lab’ at UCLA in California, where he studied the effects of acidosis and ischaemia on myocardial function and Ca2+ exchange. He was appointed Senior Lecturer at the Cardiothoracic Institute (1976) with honorary consultant physician status at the National Heart Hospital, allowing him to investigate K+ loss from hypoxia and ischaemic tissue in both laboratory and catheter lab. Appointed Reader (1980) and given a chair by London University two years after, in 1988 he became Simon Marks and BHF Professor of Cardiology at the National Heart and Lung Institute, Imperial College. He served as councillor (1988), secretary (1990) and president (1994-6) of the European Society of Cardiology (ESC) and president of the World Heart Federation (2003-5). On retirement (2008), he was made Emeritus Professor and Senior Research Investigator.

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Jeffrey T. Potts

( 1958 - 2009 )

Jeff Potts received his Bachelor of Science degree in Physical Education (1982) from the University of New Brunswick, his hometown, and a Masters of Arts degree in Exercise Science from Indiana State University in Terre Haute, Indiana. In 1986 he entered the PhD program in the Department of Kinesiology at the University of Maryland, but moved to join Peter Raven’s research program, obtaining a PhD (1993) in Biomedical Sciences (Physiology). Focussing on the carotid arterial baroreflex, he published work in the American Journal of Physiology and developed the ‘built curve’ technique of modelling the carotid baroreflex. With a postdoctoral fellowship, he then joined Artin Shoukas in the Department of Biomedical Engineering at Johns Hopkins University School of Medicine to study the role of arterial compliance and venous resistance on the carotid baroreflex control of the circulation. Subsequently, at UT Southwestern Medical Center (UTSWMC) in Dallas, he worked with with Jere H. Mitchell MD and with Julian Paton, publishing several important papers identifying the neurotransmitters within the brainstem medullary centres involved in baroreflex and exercise pressor reflex control (1995-2000).

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Mike Rennie

d. 2017

Mike Rennie held the Symers Professor of Physiology at the University of Dundee for many years and was subsequently at the University of Nottingham in Derby. He was a member of the Physiological Society from 1978 and served on the Editorial Board of Journal of Physiology and on Council; he delivered the G.L. Brown Prize lecture in 2004/5. He is best known for his work in human physiology where he pioneered the use of stable isotopes in metabolic studies.

Paul S. Richardson

d. 2011

Paul Richardson was at St George’s Medical School before he retired. Paul became a Member in 1973 and was a Journal of Physiology editor from 1974 until 1978.

J. Murdoch Ritchie

( 1925 - 2008 )

J Murdoch Ritchie was Higgins Professor Emeritus of Pharmacology at Yale University School of Medicine, and a major figure in neuropharmacology in the second half of the twentieth century. With a BSc in mathematics & physics from Aberdeen University(1944), he became a research physicist under A V Hill at the Telecommunications Research Establishment, subsequently following Hill to University College London (UCL) in 1946. Murdoch received a BSc in physiology (1949), while working on the dynamics of skeletal muscle contraction. He soon moved to the National Institute for Medical Research at Mill Hill, receiving a PhD and DSc in biophysics (1952, 1960). He began work on the control of blood pressure by unmyelinated nerves with Bill Douglas, eventually following him (1958) to Al Gilman’s new Department of Pharmacology at Albert Einstein College of Medicine, New York. Influential papers on nerve and on local anaesthetics followed. Appointed Professor of Pharmacology at Yale (1968), he directed the pharmacology course there for second year medical students for 30 years. He became Director of the Yale Division of Biological Sciences (1975–1978), championing the establishment of its first core facility providing technical support for scientific and biomedical computing. He was also co-Director of the Interdepartmental Neuroscience Program (1993–1999). He was elected a Fellow of the Royal Society in 1976 and a Fellow of the Institute of Physics (London) in 1997.

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Ian Campbell Roddie

( 1928 - 2011 )

Ian Roddie led research in three major areas related to the circulation – effects of mental stress, thermoregulation and circulation of lymph. The papers he published (largely in The Journal of Physiology) are still widely quoted some 50 years later. At Queen’s University, Belfast, his career began under David Greenfield with an intercalated BSc in Physiology (1950), an MD with gold medal (1957) and a DSc (1962). Aged 35, he succeeded Greenfield as Dunville Professor of Physiology at Queen’s and subsequently served as Dean of the Faculty of Medicine and Pro Vice Chancellor. A Sherrington lecturer, he was Chairman of the Committee of The Physiological Society from 1986 to 1988 and an Honorary Member from 1998. He was also a member of the General Medical Council, the General Dental Council and the Medical Research Council and examiner for many years for the Fellowship examinations of the Royal Colleges of Surgeons. On retirement, he served as Visiting Professor at the Chinese University of Hong Kong, then Medical Director and Head of Medical Education in Jeddah. Latterly his influence extended worldwide, particularly in developing countries, when he advised the World Bank, the Asian Development Bank and various governments and institutions in some 30 countries. His many contributions were recognized nationally when he became Commander of the Order of the British Empire.

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Martin Rosenberg

d. 2011

Martin was elected a Member in 1968 when he was in the Department of Physiology, Basic Medical Sciences, Queen Mary & Westfield College. For many years he was effectively the official photographer for The Society and was heavily involved in ensuring that the images in the Archives were indexed and properly stored. As a member of the History & Archives Committee, he took a very active part in the recording of Oral Histories. By doing considerable homework before every session, he was able to get the maximum value from each interviewee.

James Ryder

( 1974 - 2004 )

James Ryder graduated with a BSc in Sports and Exercise Science from South Bank University. His MSc (University College Chester) addressed the effects of hydroxy-methyl-butyrate supplementation on muscle damage following eccentric exercise and for his PhD, he studied markers of physical fitness in elite young soccer players. At the time of his death, he had recently taken up a lectureship in Lars McNaughton’s team in the Department of Sports Science, University of Hull.

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Bengt Saltin

d. 2014

The Society is sorry to hear of the death of Professor Bengt Saltin of the Copenhagen Muscle Research Centre, who passed away on 12 September. He was elected a Member in 2001 and an Honorary Member in 2006.

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Melville Schachter

( 1920 - 2000 )

Melville Schachter’s early work concerned the control of gastric acid secretion, work which he initiated in 1947 as an assistant professor in the Department of Physiology at Dalhousie University, Nova Scotia. In 1950 he moved to the National Institute of Medical Research at Mill Hill, London, where he worked both with Wilhelm Feldberg and with Bill Paton, on problems related to histamine, its release and effects of antagonists. He soon coined the name “kinins” for this generic group of peptides, returning to Canada in 1965 where he became the Professor and Head of the Department of Physiology at the University of Alberta in Edmonton, a position he retained until his official retirement in 1986. His lifetime’s work on kinins was recognised in 1995, at the International Conference on Kinins held in Denver, Colorado, when he was awarded the Frey-Werle Medal, the highest honour in the field.

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Alexander A. Selyanko

( 1952 - 2001 )

Alex Selyanko, a graduate of Kiev State University, conducted life-long research into synaptic transmission in mammalian sympathetic ganglia. Working in London, Kiev and Alberta in 1984, he made major contributions to the understanding of potassium channels and the M-current.

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Stephen Semple

(1927-2018)

The Society regrets to hear of the death of Member Stephen (‘Steve’) Semple. Professor of Medicine at the Middlesex Hospital Medical School from 1969-91, he was a highly respected  respiratory clinician and physiologist, working with many key figures in the field. One of his important contributions was developing the oscillation hypothesis for control of breathing in exercise. A Member of the Society since 1964  he served on the Committee from 1982-6.

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Brian Setchell

(1931-2018)

The Society notes with regret the death on 11 July 2018 of Member Brian Setchell. A vet by training, he graduated from Sydney before completing a PhD in Cambridge and, in 1982, becoming Professor of Animal Sciences at the Waite Agricultural Research Institute at the University of Adelaide. He retired in 1996. A Member of The Society since 1971, Setchell is recognised for his extensive work on many aspects of the physiology of the testis and epididymis.

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Craig Sharp

d. 2018

The Society notes the death of Member Craig Sharp in March at the age of 84. Educated as a veterinary pathologist in Glasgow, his subsequent work laid the foundation of modern sports and exercise physiology in the UK. A secondment to Kenya in 1963 resulted inter alia in his record ascent of Kilimanjaro, and the first scientific measurement of a cheetah’s running speed (64.3 mph). As the scientific head of the British Olympic Medical Centre in the 1980s, and Professor of Sports Science at West London Institute of Higher Education (later Brunel University) 1994-2005, he was responsible for guiding the training of many notable UK sports champions.

Edward Sharpey-Schafer

(1850 –1935)

Sir Edward Albert Sharpey-Schafer (1850 –1935) was an English physiologist and Fellow of the Royal Society. Born Edward Schäfer, he studied under the physiologist William Sharpey and became the first Sharpey Scholar in 1873 at University College London (UCL). In 1874 he was appointed Assistant Professor of Practical Physiology at UCL where he went on to become Jodrell Professor. He was elected a Fellow of the Royal Society in 1878 at the age of just 28. Schäfer was appointed Chair of Physiology at the University of Edinburgh in 1899 where he would stay until his retirement. He was one of the nineteen founder members of the Physiological Society in 1876 and he also founded and edited [the Quarterly Journal of] Experimental Physiology from 1908 until 1933. Schäfer was knighted in 1913. He is renowned for his invention of the prone-pressure method or Schäfer method of artificial respiration.

He was very active as a facilitator, mentor, coordinator, teacher and organiser through much of his career. He had started as a histologist and always emphasised the importance of structural knowledge. He was the co-discoverer (in 1894, with George Oliver) of adrenaline (as in the adrenal-derived, circulating hormone) and he coined the term ‘endocrine’ as the generic term for such secretions. He intuited (as did a few others, independently) that insulin must exist (i.e. a pancreatic hormone to account for diabetes mellitus) and coined the name (originally as ‘insuline’). (Banting and Best actually discovered what S-S and the others had predicted). Thus, he had a founding role in modern endocrinology. He also did important early work on the localisation of function (e.g. motor centres) to brain regions. After the death of his eldest son, John Sharpey Schafer, and in memory of his late professor William Sharpey, he changed his surname to Sharpey-Schafer in 1918.

Sir Edward Albert Sharpey-Schafer died on 29 March 1935 aged 84. Funded by bequests from Sir Edward Sharpey-Schafer (1850–1935) and his daughter Miss GM Sharpey-Schafer and in memory of Sir Edward and his grandson Professor EP Sharpey-Schafer, The Physiological Society established the Sharpey-Schafer Prize Lecture. This is a triennial lecture given alternately by an established physiologist (preferably but not necessarily from abroad) and a young physiologist chosen by The Society.

John T Shepherd

( 1919 - 2011 )

John Shepherd was a visionary Mayo Clinic physiologist. He made major contributions to understanding the regulation of the circulatory system, producing more than 300 scientific publications and four books. He received his MB, BCh, MChir and MD from Queen’s University, Belfast – joining the Department of Physiology there soon after. In 1953, he was awarded a one-year Fulbright Scholarship to the Mayo Clinic to engage in cardiovascular research, returning there in 1957. He became Director of Research there ( 1969-76), Director for Education of the Mayo Foundation, and Dean of the new Mayo Medical School (1977-83), a post which included responsibility for the Mayo Graduate School of Medicine and Mayo School of Health-Related Sciences. He then chaired the Mayo Board of Development (1983-8) and was actively involved in establishment of the Mayo Clinic campus in Jacksonville, Florida. He was president of the American Heart Association (1975-6) and a fellow of the Royal College of Physicians, and of the Royal College of Physicians of Ireland. In the course his career, he headed the American Heart Association, served as a NASA adviser and chaired the Committee on Space Medicine at the National Academy of Sciences (1965-74). He led US scientific exchanges with the Soviet Union during the Cold War, working on space physiology with scientific colleagues there.

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David Shirley

d. 2011

David Shirley was a Senior Research Fellow and Honorary Reader, in the Department of Physiology and Division of Medicine (Nephrology) at UCL. David was a leading UK renal physiologist and skilled micropuncturist. His work resulted in many significant publications concerning the membrane events underlying renal tubular fluid and electrolyte transport. David joined The Physiological Society in 1973.

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Luis Silva-Carvalho

( 1954 - 2008 )

Luis Silva-Carvalho completed his MD in Coimbra, Portugal (1977). Afterwards he joined his father at the Department of Physiology, University of Lisbon, conducting studies on arterial chemoreception for his PhD (1984). He continued these and other studies on autonomic function, eventually succeeding his father as Head of Department (1998). In the interim, he spent several years (1989-97) in Mike Spyer’s laboratory at the Royal Free Hospital School of Medicine, collaborating on studies involving cerebellar and hypothalamic control of cardiorespiratory function. Back in Lisbon, he established an internationally recognised programme of autonomic research. He was instrumental in the formation of the Institute of Molecular Medicine in the Faculty of Medicine in Lisbon, leading its Unit of Autonomic Neuroscience, and served as the Vice-Rector of the University of Lisbon (1995-8). He became a Foreign Member of The Physiological Society (1989), and later an Ordinary Member. He used the services of The Society, and the Research Defence Society, in countering growing anti-vivisection tendencies in Portugal.

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John Sirs

( 1926 - 2005 )

John Sirs was Professor of Biophysics at the University of London between 1979 and 1991. He read physics at Durham University, later moving to the Department of Colloid Science, Cambridge as research student supervised by FJW Roughton. After gaining his PhD, he became interested in the physiology of blood gas transport, research he continued after his appointment as Lecturer in Physics at St Mary’s Hospital Medical School (1960), under SG Rowlands. Over the next twenty years his research addressed the physical properties of erythrocytes, often in collaboration with clinicians. Having taught physics for some years, he later developed a statistics course for medical undergraduates, which he ran for two decades.

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Ernest Starling

(1866 –1927)

Ernest Henry Starling (April 1866 –1927) was an English physiologist and Fellow of the Royal Society. Starling joined Guy’s Hospital, London in 1882, earning his qualifying degrees of MB, in 1889 and MD in 1890. Starling worked part-time from 1889 to 1899 at Guy’s and also lectured at the London School of Medicine for Women. In this period, after research experience in various German laboratories, he established ‘Starling’s Principle’ which describes tissue fluid formation and the importance of osmotic forces across the capillary wall. He was next appointed Professor of Physiology at University College London, serving from 1899 to 1923. It was here he met Sir William Maddock Bayliss, with whom he first described gut peristalsis and then discovered the peptide hormone secretin. This first discovery of a hormone significantly broadened the fields of research in physiology. Starling was elected a Fellow of the Royal Society in 1899, but it was not until 1915 that he uncovered the findings known now as the ‘Frank-Starling law of the Heart’, which was further revised in 1919. Together with Otto Frank, Starling had independently discovered and characteristed that the stroke volume of the heart (‘cardiac output’) increases in response to an increase in the volume of blood filling the heart (‘end diastolic volume’), when all other factors were held constant and that the force of the muscular contraction of the heart is directly proportional to the extent to which the muscle is stretched. On 2 May 1927, Starling died at the age of 61 whilst aboard a ship in the Atlantic. He was buried in Kingston, Jamaica. The Physiological Society created the Bayliss Starling Prize Lecture as a joint memorial in 1960 and in 1979, the Bayliss and Starling Society was established, focusing on central and autonomic peptide functions.

Donald Hulme Steven

( 1933 - 2005 )

Donald Steven read zoology at King’s College, Cambridge, and was subsequently appointed demonstrator in Veterinary Anatomy there (1961). A talented artist, he investigated the vascular anatomy of the placenta in the sheep and mare, guided by Robert Comline in the Physiological Laboratory, and edited Comparative placentation: essays in structure and function (1975). He subsequently became Director of the Sub-department of Veterinary Anatomy (1984-9). As a Fellow of Churchill College, Cambridge, he was the Physiological Society’s first archivist when its archive was established there in the 1970s.

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Sir John Sulston

The Society regrets to hear of the death of Honorary Member Sir John Sulston CH who was awarded the Nobel Prize for Physiology or Medicine in 2002 for his work on cell lineages and on the first complete DNA sequence to be published, of the nematode worm Caenorhaditis elegans. A forceful advocate for genome information remaining in the public domain, he became the founding director of the Wellcome Sanger Centre at Hinxton. In 2002, he gave The Society’s Annual Review Prize Lecture on the ethical uses of science.

John Warburton Thompson

d. 2012

John Warburton Thompson has died suddenly aged 88. John was a Member of The Society since 1958 when at the Royal College of Surgeons of England. Although he retired from the NHS in 1990, John continued to work in the field of chronic pain. He will be greatly missed by his friends and colleagues.

Hilda Tracy

( 1927 - 2010 )

Hilda Tracy worked in Rod Gregory’s physiology laboratory at the University of Liverpool, before studying and graduating in medicine. She subsequently joined Gregory’s academic team (1958), staying in the department until his death in 1990. She performed seminal work on the isolation and characterization of the acid-secretory hormone gastrin, the first gastrointestinal hormone to be sequenced. Subsequent investigations with Gregory laid the foundation for gastrin radioimmuno-assay to become a reliable diagnostic test for Zollinger-Ellison tumours. She retired in 1993.

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Roger Y. Tsien

d. 2016

The energetic and inspiring Roger Tsien will be greatly missed by The Society, and the scientific world. He is most famous for creating a vital research tool from green fluorescent protein. This work earned him a Nobel Prize in Chemistry in 2008. The decades of research he conducted changed the course of visualisation in molecular biology. Tsien served as professor of pharmacology, chemistry, and biochemistry at UC San Diego’s medical school for 27 years, and his passing is sad and sudden news.

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Stanislav Tucek

( 1932 -2002 )

Stan Tuček had a worldwide reputation as a neurochemist and author of the 1978 monograph, Acetylcholine Synthesis in Neurons. In the mid 1960s he worked with Catherine Hebb at Babraham, and later with S-C Cheng in New York. His Institute of Physiology laboratory in Prague was extremely productive and he was involved in arranging and editing numerous important symposia on the cholinergic nervous system.

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Sir John Vane

( 1927 - 2004 )

John Vane was a towering figure in the physiological tradition of pharmacology. He was awarded the 1982 Nobel Prize for Physiology or Medicine, with B Samuelsson and S Bergström, for their work on aspirin. After graduating in chemistry from University of Birmingham (1946), he studied experimental pharmacology with JH Burn and G Dawes in Oxford and A Welch in Yale, subsequently joining the Institute of Basic Medical Sciences at in London. He was awarded a personal chair there in 1966. Having developed his signature ‘blood bathed organ cascade’, he was then able to establish major features of bradykinin and angiotensin physiology – discoveries that led eventually to the development of ACE inhibitors. He began to study aspirin in 1971 and, while managing his research group at the Wellcome Foundation (1973-86), discovered prostacyclin with S Moncada and other colleagues. Prostacyclin analogues followed. Wellcome also produced other successful new drugs under Vane’s management, including Zovirax, Tracrium and Lamictal. Vane left Wellcome for St Bartholomew’s Hospital Medical School to build a new research team there – later The William Harvey Research Institute – largely focussed on cardiovascular hormones and the cyclooxygenase-2 inhibitors. Knighted in 1984, John Vane was subsequently awarded over 50 other honorary degrees and fellowships.

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Patrick Wall

(1925 - 2001)

Honorary Member Patrick David “Pat” Wall was a leading British neuroscientist described as ‘the world’s leading expert on pain’ and best known for the gate control theory of pain.

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E. Geoffrey Walsh

( 1922 - 2003 )

Geoffrey Walsh studied medicine at Oxford, graduating in Animal Physiology in 1943. He spent two years as a Rockefeller student at Harvard University, gaining an MD, and returning to Oxford to graduate as MA, BSc and BM BCh in 1947. In 1951 he was appointed lecturer in the Department of Physiology at University of Edinburgh, under David Whitteridge. His research concerned human neuromuscular control and vestibular dysfunction – especially balance and tremor. He studied paraplegic patients, eventually being promoted to Reader, and his research interest in spastic children was recognised in his appointment as Honorary Neurophysiological Specialist at the Royal Hospital for Sick Children, Edinburgh. He was awarded Fellowship of the Royal Society of Edinburgh in 1959, and then of the Royal Colleges of Physicians of London (1967) and Edinburgh (1968). A musician in later life, his inaugural lecture as extra-mural Professor at the University of Central England in 1998 was entitled: ‘Movement control in normals, the disabled and musicians: muscles, medicine and Mozart’.

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Anne Warner

d. 2012

Professor Anne Warner FRS, who has died aged 71, applied electrophysiological methods to the study of the development of the embryo. She was responsible for initiating the Microelectrode Techniques workshop at the MBA, Plymouth and more recently directed CoMPLEX, the systems biology centre at UCL.

She was elected a Member in 1968 and served on the Committee (1975 to 1979) and on the Editorial Board of The Journal of Physiology (1980-1987).

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R A Webster

d. 2017

Roy Webster died peacefully on the 2nd April 2017. A Senior Lecturer, in the Department of Pharmacology at UCL, he was a gifted experimentalist. His early research focussed on confirming the identity and physiological roles of the newly discovered amino-acid neurotransmitters in the spinal cord. Later he went on to study drugs, notably the benzodiazepines, which interact with the receptors for these amino acid transmitters in specific regions of the central nervous system. He joined The Society in 1967.

Silvio Weidmann

( 1921 - 2005 )

Silvio Weidmann was a Swiss electrophysiologist. His elegant microelectrode work on cardiac Purkinje fibres in Cambridge led to several discoveries, including the phenomenon of all-or-nothing repolarization, a series of 1950s papers in The Journal of Physiology, and his classic 1956 book: Elektrophysiologie der Herzmuskelfaser .

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Brian Whipp

d. 2011

Brian was elected as a Member in 1984. He served on the Committee from 1993 to 1997, and was an Editor on the Board of Experimental Physiology from 1994 to 2000. Brian and his collaborators produced defining research in muscle energetics, pulmonary gas exchange and ventilatory control. His first love was sport and, among his many accomplishments, he was Welsh AAA champion for high hurdles and decathlon.

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Saffron Whitehead

(1948-2019)

The Society is greatly saddened to hear of the death on 19 February of Member Saffron Whitehead, an emerita Professor of St George’s, University of London. With a PhD from McMaster University in 1974, she made significant contributions to endocrinology, providing early evidence for how endocrine disrupting chemicals and hormones impact on fertility and cancer. A Society member from 1980, Saffron edited Physiology News (1994-98) and served on Council (1995-99). As well as authoring an acclaimed textbook on clinical endocrinology, she was interviewed and wrote widely for the media on healthcare topics under her married name, Saffron Davies.

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Reginal James Whitney

( 1914 - 2001 )

Reg Whitney conducted outstanding research in biomechanics and human physiology and was a distinguished inventor. With a zoology doctorate from the University of Birmingham, he conducted Army operational research from 1941, initially into the problems facing tank-operators. Subsequently he studied posture and motion during extreme human activities as senior scientific officer with the War Office. Working at the MRC Climate and Working Efficiency Unit (Oxford) from 1948, he developed the Whitney strain gauge plethysmograph for quantifying human peripheral blood flow and, later, a force analysis platform for investigating physiological aspects of whole body activities. In 1958, he moved to the MRC Institute for Medical Research at Hampstead. There he continued to analyse human posture and motion. Among other projects, he embarked on construction of a simulator to permit investigation by the Navy of the effects of ship motion on human performance. After 1974, he continued this work at the RAF Institute of Aviation Medicine, Farnborough, retiring from the MRC in 1980.

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Wilfred F. Widdas

( 1916 - 2008 )

Wilfred Widdas was a stalwart of The Physiological Society. A member for over 50 years, he was on the Editorial Board of The Journal of Physiology , and was its chairman (1970–72). A medical graduate from Newcastle (1937), he served as a doctor in the Royal Army Medical Corps during the war. He then trained for his PhD on placental glucose transport at St Mary’s Hospital with Prof A. Huggett, contributing to placental and fetal research until 1960. His most important papers concerned erythrocyte glucose transport and he collaborated over a long period with Graham Baker. A Reader in Physiology at Kings College London by 1954, he later became Foundation and only Professor of Physiology at Bedford College, member of the Senate of London University and Chairman of the Board of Studies in Physiology, London University.

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John Guy Widdicombe

( 1925 - 2011 )

John Widdicombe has been described as one of the giants of respiratory physiology over the last 50 years. He studied at Oxford, graduating in medicine in 1949 and taking his DPhil there as an MRC scholar at the Nuffield Institute for Medical Research (1953). After two years conscripted at Porton Down Microbiological Research Establishment, he was appointed lecturer, and then senior lecturer, at St Bartholomew’s Hospital, London. His research focused on lung mechanics and reflexes. A Visiting Scientist in the Cardiovascular Research Institute in San Francisco (1960-1), he worked on the nervous control of breathing and on the regulation of bronchomotor tone. He was then appointed lecturer in Physiology and Fellow of New College, Oxford (1961) remaining until he became Professor and Chairman of the new Physiology Department at St George’s (1972-92). He received many awards/degrees in his career including his FRCP in London in 1976, an honorary MD from Helsinki University in 2000, nine international medal awards, honorary membership of four international societies and a Life Time Achievement award. He was an Editor of The Journal of Physiology and British Journal of Pharmacology, President of the British Association for Lung Research (1994-8) and and Honorary Treasurer of The Physiological Society (1990-6).

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Robin Gow Willison

d. 2012

Robin Willison, a Member of the Society from 1965 and, from 1980 to his retirement in 1990, head of the Department of Clinical Neurophysiology at the National.

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Gerald Wiseman

Gerry Wiseman’s life’s work, at University College Hospital, addressed active transfer mechanisms in the intestinal absorption of nutrients. For this work, he devised an acclaimed rodent intestinal preparation. He retired as Reader in Physiology in 1989.

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Roger Woledge

d. 2015

The Society is sad to hear of the death of Professor Roger Woledge on 13 March following a riding accident. He had been a Member since 1968.

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Maureen Young

d. 2013

Maureen Young was born on 16 October 1915 in Southwold, England. An eminent fetal physiologist, she published well over 100 peer-reviewed original papers, as well as reviews and chapters in books, in a publishing career that spanned 63 years. For many of these, she ran a research unit in O&G at St Thomas’, becoming Professor of Perinatal Physiology before her retirement in 1982. She was one of the first women to secure a teaching position in a medical school and championed the professional recognition of women in an era when female academics were rare. She was an active advocate of the need for basic research to underpin advances in medicine and will be remembered for her many kindnesses to younger academics and clinicians.

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John Atherton Young

( 1936 - 2004 )

John Young was a towering figure in Australian physiology, a global expert on exocrine secretion and the undoubted authority on salivary secretion, as evidenced by nine ISI defined citation classics and nearly 5,000 citations in all. After studying medicine in Brisbane, John travelled to Sydney for doctoral training in renal physiology, followed by post-doctoral studies in Berlin, before returning to Australia in 1966 to a Senior Lectureship in Physiology at Sydney University. Subsequently he was promoted to Professor and Head of Department, before becoming Dean of Medicine in 1989 and then Pro-Vice- Chancellor for Health Sciences in 1996, a post he held until his retirement in April 2003.

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David L. Yudilevich

( 1930 - 2006 )

David L Yudilevich made seminal contributions in his early career to the field of capillary permeability, studying microvascular permeability in the isolated perfused gastric mucosa, heart and brain. Born in Santiago, he received his MD (1957) from the Universidad de Chile and conducted post-doctoral research for some years in the USA. He returned to Chile but left during the 1973 military coup to work with Laurence Smaje at University College London. The next year, he was appointed Professor and Head of the Department of Physiology at Queen Elizabeth College, University of London – a post he held until 1985. A strong advocate of research links between Chile and Britain, he published more than 100 articles in peer-reviewed international journals. In his retirement, he wrote a series of books on Darwin and Humboldt and was awarded Emeritus Professorship of Physiology at King’s College London, Professor Titular in the Faculty of Medicine in the Universidad de Chile and Honorary Membership of the Sociedad Chilena de Ciencas Fisiologicas.

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To notify us of the death of a Member please email membership@physoc.org.​

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