F.R.S. 1953 11 January 1998: Elected − Henry Barcroft. 18 October

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Henry Barcroft. 18 October 1904 − 11 January 1998: Elected
F.R.S. 1953
A.D.M. Greenfield and I.C. Roddie
Biogr. Mems Fell. R. Soc. 2000 46, 1-17, published 1 November 2000
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HENRY BARCROFT
18 October 1904 — 11 January 1998
Biog. Mems Fell. R. Soc. Lond. 46, 1–17 (2000)
HENRY BARCROFT
18 October 1904 — 11 January 1998
Elected F.R.S. 1953
B A.D.M. G*  I.C. R, C.B.E.†
*25 Sutton Passeys Crescent, Wollaton Park, Nottingham NG8 1BX, UK
†Calle San Bernabé 3, Lomas Club-Pueblo, 29600 Marbella, Spain
Henry Barcroft (H.B.) was born on 18 October 1904 at 92 Chesterton Road, Cambridge. He
was the first son of Joseph Barcroft (J.B.), later Professor Sir Joseph Barcroft, C.B.E., F.R.S.,
and Mary Agnetta (Minnie) Ball (M.A.B.), whom he had married in 1903. He had one
brother, Lt.-Col. Robert Ball Barcroft, born in 1909, and no sisters. Both of his parents had
distinguished ancestors.
T B 
The Barcrofts were an old Lancashire family. William Barcroft, born in 1611, went to Ireland
as an officer in Cromwell’s army in 1658, joined the Society of Friends and settled in what
was then King’s County. H.B.’s grandfather (1837–1905), also called Henry Barcroft, lived in
a large house called ‘The Glen’ in Newry, County Down. He became managing director of the
linen firm of Richardson and Owden in Bessbrook, a small town about three miles from
Newry. The blockade of cotton exports from the South in the American Civil War had
increased the demand for linen and the industry was booming. Henry Barcroft was a practical
innovator and inventor. He increased company productivity by his invention of the
‘Bessbrook Loom’, which could weave linen damask much faster and more economically than
the Jacquard Loom then in vogue. Other inventions included an electric tramway system
whose carriages could run on both road and rail and was served by gates that opened
automatically as the tram approached and closed when it passed. He also devised a partly
submerged propeller system for steamships. This versatility led to his election to the Institute
of Civil Engineers in London. In 1890 he became High Sheriff of County Armagh and
subsequently Deputy Lieutenant for County Down.
3
© 2000 The Royal Society
4
Biographical Memoirs
Henry Barcroft and his wife Anna had five children, the second of whom was H.B.’s
father, Joseph. Joseph became a pupil at the Friends’ School, York, and the Leys School,
Cambridge. At the latter he was one of a group, including Henry (later Sir Henry) Dale
(F.R.S. 1914), who achieved distinction in diverse fields. While still at school, he obtained a
London University BSc degree. At King’s College, Cambridge, he was awarded a double First
in the Natural Sciences Tripos. He embarked at once on research under Michael Foster’s
leadership among an array of brilliant men including J.N. Langley, F.R.S., W.H. Gaskell,
F.R.S., H.W.R. Rivers, F.G. Hopkins and H.M. Fletcher. He analysed the effects of
autonomic nerve stimulation on the submaxillary gland by studying the gas content in venous
blood. The combination of circulatory and respiratory investigation was a prelude to his life’s
work. He remained at Cambridge for the rest of his life, with ever increasing distinction.
T B 
H.B.’s mother’s family were descendants of the Balls of Bampton in Devon. His maternal
grandfather was Sir Robert Stowell Ball, F.R.S. (1840–1913), Astronomer Royal of Ireland
(1876–1892) and later Lowndean Professor of Geometry at Cambridge (1892–1913), where he
was a Professorial Fellow of King’s College. A brother of Sir Robert’s, Professor Valentine
Ball, F.R.S., held the chair of Geology in Dublin University. Another brother, Sir Charles
Ball, Bt., was Regius Professor of Surgery in Dublin University. Sir Robert’s father, the
zoologist Dr Robert Ball (d. 1857), was Secretary of the Dublin Zoo and the Queen’s
University of Ireland. He was proposed for election to the Fellowship of The Royal Society in
1857 by T.H. Huxley and eleven others but died before the election.
J.B. had met the Ball family in 1893 through his friendship with M.A.B.’s brother, W.V.
Ball, when they were freshmen together. A decade later, J.B. and M.A.B. were married in
August 1903.
C  C
J.B. and M.A.B. set up home in 92 Chesterton Road, Cambridge. Soon after H.B.’s birth in
1904, the family moved from there to 13 Grange Road, a house with a beautiful garden. J.B.
and M.A.B. were excellent hosts and loved good company. During his childhood, H.B. met
many famous personalities in Physiology when they visited the Cambridge laboratories and
the family home.
M.A.B. kept a record, ‘Henry Barcroft. His Journal’, from the time of his birth until the
birth of his own first child. According to the record, he was an exceptionally loving child who
was deeply attached to his parents and his younger brother Robert. He included them in all
his prayers. From the age of five, he had a German governess to whom he became very
attached, and spoke German as fluently as English at the age of ten. This enabled him later to
lecture confidently in German.
On holidays at his grandparents’ home in Newry, H.B. enjoyed meeting his relatives and
their friends and renewing his acquaintance with the donkeys and the golf. J.B., being a golf
enthusiast, had built a golf course in the grounds of ‘The Glen’.
Henry Barcroft
5
S   K’ C C S
 M C
From 1912 to 1918 H.B. was a dayboy at King’s College Choir School, where he found the
headmaster, C.R. Jelf, a strict disciplinarian and a good teacher of Latin. In 1918 he became a
boarder at Marlborough College. His mother thought it suited him well, but H.B. found the
conditions austere, the food poor and the experience not enjoyable. No doubt the war and its
aftermath contributed to this. However, while at Marlborough, he won a Bethune Baker
Scholarship, passed the Cambridge 1st MB examination and gained the ‘top’ Exhibition to
King’s College, Cambridge, with botany, zoology and chemistry as his subjects. He ascribed
much of the credit for this to the biology master, Mr Ashley G. Lowndes, among whose pupils
were P.B. Medawar, R.J. Pumphrey and J.Z. Young; all three were later elected F.R.S. In
September 1922, Mr Lowndes took H.B. to study at the Marine Biological Station at
Plymouth.
Between leaving Marlborough in March 1923 and entering King’s College that autumn,
H.B. and his father collaborated on two investigations. The first, at Cambridge, showed that
in rats given low concentrations of carbon monoxide to breathe, the increase in carboxyhaemoglobin in the splenic pulp lagged behind that in the circulating blood by up to thirty
minutes. This led to a paper in J. Physiol. (1)*. The second and more difficult, performed at
Cambridge and Plymouth with the use of oxygen capacity and spectral analysis with the
Hartridge reversion spectroscope, compared the red pigment in the marine worm Arenicola
with haemoglobin in vertebrates. J.B., as chairman of the the Haemoglobin Committee of the
MRC, was interested in the comparative chemistry and physiology of haemoglobin. Their
joint work led to a paper in Proc. R. Soc. Lond. B (2). H.B. modestly regarded his contribution as mere ‘bottle washing’, but J.B. and H.B. worked independently and recorded their
observations separately. The two projects provided a superb research apprenticeship.
C U
At King’s College, H.B. studied anatomy, physiology, zoology and chemistry. He was awarded
a First in Mays in 1924 and a Vintner Exhibition. In 1925 he was awarded a First in the
Natural Sciences Tripos Part I and the Cambridge 2nd MB. During the summer of that year,
J.B. arranged for H.B. to assist Professor Yandell Henderson, who was visiting from Yale, in
setting up the ethyl iodide method to determine cardiac output in man. Soon after Henderson
returned to Yale, H.B., while still an undergraduate, wrote his first solo publication (3) on a
source of error in Henderson and Haggard’s method of estimating cardiac output. His closely
reasoned argument took account of variations in ratios of ventilation to perfusion in different
parts of the lung.
During Lent Term 1926 he was ill, so he began Physiology for the Part II Tripos only in
the summer. By that time J.B. had succeeded Langley as Professor in the Department. He
arranged for his role as H.B.’s supervisor to be given to E.D. (later Lord) Adrian, F.R.S., in
exchange for one of Adrian’s pupils because he thought it inappropriate for H.B. to have a
* Numbers in this form refer to the bibliography at the end of the text.
6
supervisor who was both his professor and his parent. In 1927, H.B. obtained a First in the
Part II Tripos.
Among his classmates were G.W. (later Sir George) Pickering (F.R.S. 1960) and Rachel
Eckhart, who took Part II in 1926, and Alison Dale (later Lady Todd) and Bryan (later Sir
Bryan) Matthews (F.R.S. 1940), who took it in 1927. Pickering was to become a lifelong
friend.
H.B. was introduced by F.J.W. Roughton (F.R.S. 1936) to the Cambridge University
Natural Sciences Club and was its president for a term in 1927. He had been president of the
Cambridge University Medical Society in the previous year. At about that time, Ernest
Starling, F.R.S., visited Cambridge to lecture to the Society on the heart–lung preparation
that he had developed in dogs to show the effect of filling pressure on stroke volume in the
heart. H.B. gave an amusing account of Starling’s talk in his Bayliss–Starling Memorial
Lecture to the Physiological Society in 1976 (36).
I was meetings secretary at the time and in charge of the arrangements for showing the film. The
showing was to take place in the Examination Hall behind the Cavendish Laboratory. In those days
films were inflammable and regulations had to be observed before they could be shown to societies—
so many yards between the projector and the nearest person in the audience, so many water buckets,
and so many sand-buckets, readily available in case of fire. The projector itself was enormous and
made such a noise that Starling’s commentary was difficult to hear. What I remember chiefly is the
film itself. The light in Starling’s room at University College where the film was to have been made had
been insufficient. Starling had been told that better lamps would be available in Paris at the Sorbonne,
but there too the lights had been found not to be strong enough. Starling was adamant—the film had
to be made. Taking his scalpel he cut through the tissues connecting the heart and lungs to the body
and with the help of those present, carried the preparation into the forecourt of the Sorbonne and tied
them to the railings between the courtyard and the pavement of the public thoroughfare. What I
remember chiefly about the film is the look of amazement on the faces of the pedestrian passers-by
when they saw the beating heart on the railings, only a yard or two away.
In 1927 H.B. embarked on postgraduate work in Cambridge with a Harold Fry
Studentship of King’s College, and later with a George Henry Lewes Studentship in 1928.
He had decided on a career in physiology and, for this purpose, sought a fellowship at
King’s College. In January 1928 he demonstrated a simple mechanical stromuhr he had
developed (4) to the Physiological Society. Whereas most contemporary methods tended to
average blood flow over prolonged periods, the stromuhr provided moment-to-moment
measurements. At the suggestion of G. Anrep (F.R.S. 1928), H.B. used the stromuhr during
his Part II work to study the effect of intravenous adrenaline on aortic blood flow (6). Anrep
had also described an increase in left ventricular contractility that followed aortic occlusion in
the dog. It became known as the Anrep effect, and H.B. investigated it further using his
stromuhr. He found that occluding the descending thoracic aorta caused a paradoxical
increase in cardiac output (4). He continued this work for a further two years at Cambridge
and later at University College London. He thought it possible that the effect might be due to
an undiscovered baroreceptor reflex. The work formed the subject of his unsuccessful
fellowship submissions to King’s College in 1927, 1928 and 1929 and for another dissertation
that gained him the Gedge Prize (a prize that his father had won jointly with Henry Dale
thirty years earlier).
Henry Barcroft
7
S M’ H M S
On J.B.’s advice, he decided not to stay longer in Cambridge without a fellowship. Instead he
arranged to complete his medical qualifications to increase his eligibility for academic posts
elsewhere. So he went to St Mary’s Hospital London in October 1929 with a Harmsworth
Scholarship as a clinical student and part-time demonstrator in the Department of
Physiology of the Medical School. Although he did not enjoy clinical work much and suffered
a good deal from sinus infections, he qualified MRCS, LRCP in 1932. While at St Mary’s he
published the results of earlier investigations with the stromuhr under a variety of
experimental conditions (4–6).
U C L
After qualifying in medicine in 1932, H.B. proceeded directly to a lectureship in the
Department of Physiology at University College London (UCL) without taking house
appointments. The Head of Department, Professor Lovatt Evans, F.R.S., had been impressed
with H.B. when he examined him in the Cambridge Tripos.
At UCL, H.B. gave about 150 lectures a year, ran practical classes for 100 students and
supervised research students. He also organized fortnightly colloquia for the UCL Research
Club. His salary of £350 per annum increased to £410 on completion of his Cambridge MB,
BChir degree.
His time at UCL had an important bearing on his future research interests in that it
brought him into contact with the work on human physiology in Sir Thomas Lewis’s
Department of Clinical Research at University College Hospital. He particularly admired the
work of his friend G.W. Pickering. He was also impressed with the work of others there such
as R.T. Grant, Harold (later Sir Harold) Himsworth, John (later Sir John) McMichael and
C.R. (later Sir Charles) Harington, who were all later elected F.R.S. Their influence was one
factor that led H.B. to work on human physiology in Belfast a few years later.
In the meantime he continued at UCL to study the paradoxical increase in cardiac output
after occlusion of the descending aorta (7, 8). It was difficult and painstaking work requiring
heroic surgery, beautiful but delicate preparations, thoughtful experimental approaches and
clear analysis. It owed something to the elegant surgical techniques developed by Ernest
Starling and Ivan de Burgh Daly (F.R.S. 1943). He found that the paradoxical increase was
not a nervous reflex because it persisted after destruction of the central nervous system. It
was not due to a release of adrenaline because it persisted when adrenal venous blood was
prevented from returning to the central circulation. It was not an intrinsic response of the
heart because it occurred even when an artificial pump that maintained central venous
pressure at zero replaced the heart. However, it was blocked when the inferior vena cava was
clamped. H.B. concluded that the paradoxical increase in output was a mechanical
consequence of the redistribution of blood in the circulation and was not due to an
undiscovered circulatory reflex. Coming soon after the discovery of the baroreceptor reflexes,
this was an important finding and earned him his Cambridge MD.
Another significant event occurred during his time at UCL. In 1933 he married Bridget
Mary (Biddy) Ramsey, who had just graduated in that year from the London School of
8
Medicine for Women. She was the elder daughter of Arthur Ramsey, President of Magdalene
College Cambridge and sister to a future Archbishop of Canterbury. The marriage took place
in the same church (Coton Church, Cambridgeshire) with the same minister (The Revd
Canon Adams) who had officiated at his parents’ marriage thirty years earlier. H.B. and
Biddy then began a long and happy life of mutual companionship together and had four
children. Biddy was medical officer to an infant welfare clinic in Camden and Islington
Health Authority for twenty-five years. Their time together ended sadly when she died in
April 1990.
T Q’ U  B
In 1935, H.B., at the age of thirty, was appointed to the Dunville Chair of Physiology in the
Queen’s University of Belfast. It was a difficult appointment. As the only permanent member
of academic staff, he was expected to give all the lectures, six days a week from 9 to 10 a.m.
One of us (I.C.R.), as a preclinical medical student in Belfast, attended some of these. We
were deeply impressed by H.B.’s enthusiasm and patent love of his subject. This came over
clearly in his lectures, in which he described his physiological adventures with the ring of
authenticity. His blow by blow account of the controversy between his father and J.S.
Haldane, F.R.S., on the question of oxygen secretion by the lungs made a deep and lasting
impression. It was a beautiful example of how scientific knowledge inches forward. Students
had no doubt that he was one of the most honest and modest of men. He differed from many
of his contemporaries in that it was possible, even as junior students, to understand virtually
everything he said and, at the same time, value his impish sense of humour.
With large numbers of students, frequent class exams, essay style questions and two parttime demonstrators, the marking of scripts was a constant and formidable task. The
department had little modern equipment or laboratory space, and animal facilities were
rudimentary; so were departmental grants. There was one departmental technician who had
been with the department for forty years but his skills were largely histological. (Before
coming to Belfast in 1893, Harry Miller had worked in the Department of Chemistry at UCL
with Sir William Ramsay, F.R.S., the discoverer of argon and other rare gases.)
At that time the Belfast Medical School was the fourth largest in the British Isles. It had a
reputation for sound clinical instruction, producing sensible and competent doctors, but was
not noted for research. Most members of the faculty had little time for it. However, it was the
specific hope and expectation of the university that H.B. would stimulate research in the
Faculty of Medicine. In his own words:
At this time at Queen’s and in many other Medical Schools, the clinical professors had no facilities for
doing research. They realised however that the day had dawned for clinical departments to do research
but they had not the time, expertise, or facilities for training newly qualified clinical staff in how to do
laboratory work. So the solution was to spot outstanding promising graduates and after housejobs to
send them for a couple of years to the Anatomy, Physiology or Biochemistry Departments for basic
training. They worked for an MD degree and presented a thesis. I at once became the delighted
employer of very responsible and up and coming clinicians who combined working for the MD degree
with taking classes in practical work and I had the delightful experience of having keen research
collaborators.
Henry Barcroft
9
R   B
Soon after arrival in Belfast, H.B. and W.M. Loughridge, one of the demonstrators, studied
the reliability of the thermostromuhr method for measuring blood flow in animals (9) and
found it subject to considerable errors. This perhaps was the final straw that made H.B. decide
to move to the field of research on which he concentrated in his later years and that led to his
election to the Fellowship, the human peripheral circulation. His doubts about the relevance
of observations on anaesthetized and probably shocked animals, his clinical contacts at UCL
and the expectations of Queen’s University had all led him in this direction. He expressed his
feelings thus:
While at University College I realised the limited value of my work on severely shocked dogs and I was
looking around for other topics. My attraction to human physiology sprang partly from my
admiration for my friend G.W. Pickering’s work in Lewis’s Department of Clinical Research. I felt a
powerful attraction to all that went on in Lewis’s department. My dissatisfaction with my study of the
circulation in animals was further increased by the finding that the Rein thermostromuhr was in fact
of very doubtful value.
Muscle blood flow during muscular contractions
H.B.’s first study in humans was with J.L.E. Millen (10). It was designed to study blood flow
through contracting human calf muscle. It was found that mechanical compression of blood
vessels by the contracting muscles could arrest local blood flow. In these heroic experiments,
each lasting five or six hours, the subjects (usually themselves) stood in bins filled with water a
little above or a little below core body temperature. Thermoelectric junctions inserted into the
calf muscles were used to register temperature changes. Sustained contraction at 20–30% of
maximal strength nearly, or completely, arrested calf blood flow. In contrast, rhythmic
contractions, as in walking, increased flow.
O.G. Edholm
When the medical course in Physiology at the University was extended to five terms in 1937,
the staffing of the Department was increased by the appointment of O.G. Edholm (O.E.) to a
lectureship. He held this post with distinction until leaving for the chair at the Royal
Veterinary College in London in 1944. H.B.’s collaboration with O.E. was a very happy one
and resulted in a highly productive phase in both their research careers. Of O.E., H.B. wrote:
I can not exaggerate too much the privilege it was to have him join the departmental research. I owe
him very much indeed. A great many of the ideas in our joint work were his. His sound judgement and
knowledge of men were of the greatest value in the prosecution of the research.
When O.E. left in 1944, he was succeeded by Q.H. Gibson (F.R.S. 1969) and then by J.T.
Shepherd.
V  
The thermocouple method for estimating blood flow had a slow response time and was not
able to give moment-to-moment estimates of flow. So H.B., now joined by O.E., turned to
10
venous occlusion plethysmography. The principle is simple. Stop the venous return and
measure with a plethysmograph the initial rise in volume due to continuing arterial inflow.
Venous occlusion is achieved by inflating a cuff around the limb above the plethysmograph to
a suitable pressure. Each measurement requires a volume record for only a few seconds, so
measurements can be made several times a minute. The procedure is comfortable and noninvasive and experiments can often be completed in under two hours. The method had
previously been used by others, notably T. Lewis and R.T. Grant in 1925. It permitted the
frequent estimations of flow that H.B. had enjoyed earlier with his stromuhr.
H.B. and his colleagues used the method to make elegant systematic studies of the nervous
and chemical control of the circulation in human limbs, matters of medical and surgical as
well as physiological importance. First they looked at the effect of the water temperature in
the plethysmograph on local blood flow and on the temperature deep within the forearm (11).
This was a necessary preliminary for subsequent studies.
Vasoconstrictor innervation of muscle
H.B. and his team showed that blood vessels in human skeletal muscle are innervated with
sympathetic vasoconstrictor nerves. Several groups, including G. Anrep, A. Blalock and
A. Samaan in 1934 had demonstrated such innervation in certain animal species, but the
situation in humans was less clear. Some groups had looked for it in humans by using indirect
methods, but without success. In contrast, vascular surgeons were performing surgical
sympathectomy as a standard procedure in patients with poor peripheral circulation and
related conditions. H.B. was aware of this because one of his part-time lecturers in physiology,
James Loughridge, was a surgeon doing such work at the Royal Victoria Hospital, the main
teaching hospital in Belfast. H.B. recognized that these patients could provide information on
the importance of sympathetic nerves in regulating the circulation in human limbs: with
unilateral sympathectomy, vascular responses in the normal limb could be compared with
those in the contralateral sympathectomized limb.
H.B., W.McK. Bonnar, O.E. and A.S. Effron (12), in carefully designed experiments,
measured blood flow in both forearms of normal subjects with the radial, ulnar and median
nerves blocked near the elbow on one side. Blood flow in the nerve-blocked arm was increased
relative to the control arm; with the plethysmograph temperature at 35 °C, blood flow was
approximately doubled on the nerve-blocked side. The increase was not in the skin because it
occurred after the skin circulation had been suppressed by adrenaline ionophoresis. No such
increase occurred when motor nerves were blocked in sympathectomized limbs. This indicated
that the increase in flow was due to the blocking of sympathetic vasoconstrictor nerves and
was not a mechanical effect of skeletal muscle paralysis. It was concluded that the release of
vasoconstrictor tone with deep nerve block was in vessels deep to skin, presumably in skeletal
muscle. It was the demonstration of the vasoconstrictor nerves to muscle that H.B. felt to be
his most important contribution and gave him most satisfaction in retrospect fifty years later.
With another team he showed subsequently that the vasoconstrictor fibres to muscle were
not involved in the nerve-mediated vasodilatation that occurred in limbs in response to body
heating. With general body warming, the total blood flow through the limb, as measured by
plethysmography, increased; however, that through the muscles, as measured by a heated
thermocouple method, did not (27). Thus the vasodilatation with body heating seemed to be
caused by the release of vasoconstrictor tone in skin vessels only.
Henry Barcroft
11
V   
During World War II, H.B. and O.E. made a study of the effects of haemorrhage in
volunteers (mostly conscientious objectors) in collaboration with John McMichael and E.P.
Sharpey-Schafer (13) at the British Postgraduate Medical School at Hammersmith. They
measured forearm blood flow, arterial pressure and cardiac output. During the studies, one of
the subjects fainted. Though the faint caused some local commotion, H.B. said that O.E.
showed admirable composure and, realizing the importance of the opportunity, continued to
make plethysmographic observations throughout the faint. Despite the marked decrease in
arterial pressure, forearm flow increased at a time when cardiac output had not diminished.
This indicated that the initial decrease in the arterial pressure was due to a decrease in
peripheral resistance rather than in cardiac output, leading to the conclusion that, because the
skin was pale, vasodilatation occurred in muscle blood vessels.
H.B. and O.E. continued these studies in Belfast (14, 15). To induce fainting more
regularly, venesection was combined with inflation of pneumatic cuffs on the thighs. Trapping
venous blood in the legs was a useful way of increasing apparent blood loss that could be
reversed quickly. The combination caused fainting in nearly all young healthy subjects and
each faint led to an increase in forearm blood flow. The increase in flow, despite the fall in
perfusion pressure, indicated vasodilatation in the forearm during fainting. The vasodilatation
was deep to skin because (i) the skin was very pale during a faint, (ii) suppression of the skin
circulation by adrenaline electrophoresis did not prevent it and (iii) blood flow in the hand
(which is mainly through the skin) decreased during the faint. It was mediated by sympathetic
nerves and not by circulating adrenaline because it was not seen in sympathectomized
forearms. It was postulated that the nerves were vasodilator, rather than vasoconstrictor,
because flow increased to a higher level in normal arms than in nerve-blocked arms where
vasoconstrictor tone was fully released. Similar increases in forearm blood flow were found
also in fainting caused by hypoxia (15).
Surgical sympathectomy
Before leaving the Queen’s University of Belfast in 1948, H.B. began to examine the timecourse of the physiological changes in limbs after surgical sympathectomy (16–18). In limbs
tested one to six months after operation, vasomotor and sudomotor reflexes were absent but,
one to two years later, they had returned. Thus some sympathetic function can be regained
after pre-ganglionic sympathectomy.
S T’ H M S
H.B. was appointed to the chair of Physiology at St Thomas’ Hospital Medical School in
1948. As in Belfast thirteen years earlier, he had to overcome many problems. London was
still recovering from World War II, during which the hospital had been bombed and its
activities dispersed. The Department of Physiology, like most departments of physiology after
the war, was in a run-down state and starved of young research workers. Laboratory space for
research was limited; and there was little tradition of clinical research in the hospital. H.B.
wrote:
When I went to St. Thomas’ in 1948, clinical research was almost nil. I had several youngsters for
12
training [but] few from the hospital since Sharpey-Schafer was soon to be appointed and was a
marvellous research clinician himself. So the source of clinical physiologists dried up and I felt the loss.
But of course it was quite time for the clinical departments to grow up researchwise.
Despite these problems, H.B. was still able to attract many capable young people to
maintain an active research programme. With these and more senior colleagues from overseas,
he continued the peripheral vascular studies started in Belfast. This included work on muscle
blood flow in rhythmic exercise (20), the mechanism of functional hyperaemia (30–35) and
the vascular responses to catecholamines (21, 24, 26, 28, 29). In those days before the
production of sharp disposable needles, leak-proof three-way taps, reliable infusion pumps
and good quality plastic tubing, investigative work on humans presented many technical
problems. H.B.’s ingenuity allowed him to overcome most of these but the equipment used
was quite primitive by modern standards.
At St Thomas’ Hospital Medical School, H.B. continued his studies on the time-course of
the vascular changes after surgical sympathectomy (19, 22, 23). With Dr Lynn at the Royal
London Postgraduate Medical School, he examined the changes in blood flow in the feet for
up to three months after sympathectomy. In feet with normal arteries, flow was much
increased after surgery, reaching a maximum on the second day. By the sixth day, however,
vascular tone had returned almost to pre-operative levels.
Much of H.B.’s early work appears in an excellent monograph (25) that he published with
Jeremy Swan in 1953, the first monograph to be published in the Physiological Society’s
monograph series. Most of its contents are as valid and relevant today as they were at the
time it was printed. It became a classic.
Many scientific honours came his way. He was elected a Fellow of The Royal Society in
1953 and an honorary member of the Physiological Society in 1974. The University of
Western Australia (1963), the University of Innsbruck (1969) and the Queen’s University of
Belfast (1975) awarded him honorary degrees. He was Arris and Gale Lecturer to the Royal
College of Surgeons (England) in 1945, Bertram Louis Abrahams Lecturer to the Royal
College of Physicians in 1960, Robert Campbell Memorial Orator to the Ulster Medical
Society in 1975 and Bayliss–Starling Memorial Lecturer to the Physiological Society in 1976.
He was Secretary of the Research Defence Society from 1972 to 1977 and later its VicePresident. He was a Wellcome Trustee from 1966 to 1974.
R
After H.B. retired from the chair at St Thomas’ in 1971 he continued to live in north London
and remained active with the Research Defence Society and the Wellcome Trust. In 1975 he
gave a Bayliss–Starling Memorial Lecture entitled ‘Lymph formation by secretion or
filtration?’ to the Physiological Society (36). In 1984, H.B. and his wife attended a party given
by his colleagues to celebrate his eightieth birthday. It was held before the dinner of the
Physiological Society at the Institute of Neurology in London. When his wife later became ill,
he looked after her devotedly during her long illness until she died in April 1990. His mind
remained sharp and alert and he enjoyed visits from his many friends to discuss recent
developments and reminisce on old times. In October 1994, another party was held at the
Royal Society of Medicine to celebrate his ninetieth birthday. There were short tributes by a
number of his friends, and Jeremy Swan, the co-author of their joint monograph, spoke for
Henry Barcroft
13
all in expressing his gratitude to H.B. for the influence he had exerted on so many lives. He
said:
[I remember the] encouragement which you always gave your younger colleagues. I particularly
remember on my second or third presentation (filled with fear) at the Physiological Society with A.V.
Hill, Henry Dale, J.H. Burn in the front row, I had seen you talking to Dale earlier. My paper was the
last of the morning session and after a pleasant discussion, the chairman adjourned the meeting. Dale
got up from his seat in the front row and walked over towards me and boomed a congratulatory note
of approval. God had spoken, and I was anointed for the rest of my life. I always wondered if you had
something to do with that. Whether or not, you always did it, both directly or indirectly, in your
genuine enthusiasm for our successes, and concern about our failures. And then there was the train
ride to an MRC meeting in Manchester. In a First Class carriage, with McMichael, Pickering, Sir
Geoffrey Jefferson, Franklin (I think), yourself, and little me—the poor boy from the West of Ireland.
And you asked my opinion on something or other. I was a part of it all. Dear Henry, you are the
person who started me on my career. Without you it never would have happened. I will always
remember you.
H.B., by this time limited in mobility and somewhat frail, enjoyed the gathering and made
a short speech of thanks at the end.
On 7 November 1997, at the last meeting of the Physiological Society to be held in the
Sherrington School of Physiology at the St Thomas’ Hospital Medical School site, a lecture
was given by one of us (I.C.R.) to honour and commemorate H.B.’s contribution to
physiology and the Sherrington School. Happily, he was able to attend the lecture,
accompanied by his son John. He sat in a wheelchair in the front row and enjoyed the
recapitulation of his life’s work and activities. At the Society dinner that followed, he was
welcomed with great warmth and affection. Sadly, about nine weeks later, he had a stroke at
home that took away his ability to speak. He died peacefully on 11 January 1998. A service of
thanksgiving was held for him on 21 January 1998 in the Free Church, Hampstead Garden
Suburb. At the service, members of his family, former colleagues and friends talked warmly of
the great contribution that this kind and courteous man had made to all their lives.
P 
With such a famous and admired father, H.B. had, to some extent, to live in his enormous
shadow. But it was a shadow that he never seemed to resent. Of the important influences on
his life and work he wrote, ‘First and foremost [was] my Father’s influence and immense
encouragement and monumental kindness.’ J.B. was a person of singular intelligence,
enthusiasm and intellectual integrity and H.B. was fortunate in having such a role model. As a
duo, they make an interesting comparison. They shared many traits. Some of the descriptions
of J.B. by Roughton in his Biographical Memoir for The Royal Society could have been
written for H.B. with equal force. Both retained a youthful freshness of mind and sense of
wonder to the end of their days. Both could attract younger colleagues and infect them with
their own enthusiasm. In new ventures, both would buckle to and, if need be, devise methods
that were simple to the verge of crudity, but usually worked. By showing how good research
could be performed in a friendly, rapid and informal way, both were a great encouragement to
beginners who doubted their own ability to discover anything new or worthwhile. They
attracted young people by their free and unpretentious friendship, sense of fun, constant flow
14
of ideas, simple and direct methods of work, enthusiasm tempered by patience, and
imperturbability when things went wrong. H.B. was particularly good at helping juniors see
the flaws in the thesis that they were proposing in a way that never crushed or ended hope, but
seemed to open up new avenues of opportunity.
One of us (A.D.M.G.) never had the opportunity to work with H.B., but followed him in
the chair at Queen’s, Belfast, and found him unfailingly kind and encouraging to himself and
all the members of an expanding department. All looked forward to seeing H.B. at meetings
of the Physiological Society and his occasional visits to Belfast as an external examiner. He
was always interested in, and anxious to discuss, their work and progress.
J.B. and H.B. differed in their interest and aptitude for committee work. Although J.B.’s
prime working interest was his research, he had vast experience in committee work, in his
college, university, learned societies, various government advisory boards and institutions at
both national and international level. On committees he was highly effective both as chairman
and as an ordinary member. H.B. was more single-minded in his attachment to research. He
showed little interest in expanding his administrative responsibilities or in becoming active in
academic politics. When he served as a Wellcome Trustee, it was said of him that he was
on the whole a retiring person who had never served on any of the major organizations such as the
MRC. His interests were very narrow but deep. When he was intrigued by a problem, he would be like
a leech, quite unable to pass on to the next step, say, on a tour of a laboratory. He did not live in a
world of policy speculations and development. He therefore played a valuable role in a narrow range
of the Trustees’ activities. There could not have been a more loyal trustee than Henry Barcroft and no
more charming and kindly man.
H.B.’s kindness was legendary. He was loath to speak ill of anyone and this was one of the
reasons that he was so greatly loved by so many people. His parents had inculcated this style
in him from an early age. An entry in his mother’s journal dated 1910 stated:
Yesterday Joe and I made a resolve that, as far as we could, we would never let the children hear us
speak unkindly of anyone unless with a view to checking what we considered to be an imprudent
friendship.
H.B. followed this precept throughout his life and was at pains to see the best, never the
worst, in the people that he knew. Sheila McMichael in an address given at H.B.’s
thanksgiving service in January 1998 said:
Character assassination was a form of pursuit totally foreign to Henry’s nature. He was able to find a
good word to say about almost anyone.
When he was knocked down by a car that was going too fast in a crowded area, H.B. kept
insisting that ‘It was entirely my fault!’
He was fortunate in being born into a stable and loving family and this good fortune was
to stay with him for the rest of his life. He and his wife were bound by strong bonds of
affection and mutual support. He felt her to be a major contributor to whatever success he
had had in life and of her he wrote, ‘I can hardly overestimate my gratitude to my wife for
unselfish encouragement and invaluable advice.’
To quote Sheila McMichael again:
Henry was devoted to his family—his children, his grandchildren,—and he always spoke of them with
the greatest affection. He was lucky, especially in this day and age, that this affection was returned and
they gave him their love and their support right to the end.
Henry Barcroft
15
R 
One of H.B.’s strengths was his ability to break down complex phenomena into component
parts that could be described lucidly and tested separately. Similarly, his diagrams were clear,
easy to understand and convincing. Students found them invaluable in coming to grips with
difficult concepts. Technically, he could usually devise and make simple if crude equipment to
perform his experiments, so maintaining the family tradition of inventiveness. Once his mind
latched on to a problem, he pursued it relentlessly until it was solved. His quest for the
mechanism of functional hyperaemia was almost obsessional. He had an excellent memory
and stored in his mind a diverse collection of snippets of scientific knowledge that could
surface when needed to aid interpretation of new phenomena. The other-worldliness of his
outward demeanour tended to obscure a tough inner resolve to make the best of his
opportunities in the real world. In making decisions, he could be quite ruthless and
unswerving in his determination to do what he considered to be the right thing.
One of the most telling testimonies to his influence on others was that many of those
privileged to train with him followed his example by continuing in active research in many
parts of the world. Through his example and influence on others, he has had a remarkable
effect on the way in which we think and on our understanding of the regulation of the human
peripheral circulation.
A
We thank the Wellcome Trust for help in obtaining copies of H.B.’s publications. We are also deeply indebted
to Dr Sheila Howarth (Lady McMichael) for reviewing the manuscript and making many valuable suggestions.
We also thank two of H.B.’s children, Dr John Barcroft and Mrs Sarah Falk, for their help and advice.
The frontispiece photograph was taken by the author in London in August 1993.
B
The following publications are those referred to directly in the text. A full bibliography
appears on the accompanying microfiche, numbered as in the second column. A photocopy is
available from The Royal Society’s Library at cost.
(1)
(2)
(3)
(4)
(5)
(6)
(7)
(8)
(1)
1923 (With J. Barcroft) Observations on the taking up of carbon monoxide by the
haemoglobin in the spleen. J. Physiol. 58, 138–144.
(2) 1924 (With J. Barcroft) The blood pigment of Arenicola. Proc. R. Soc. Lond. B 96, 28–42.
(3) 1927 A source of error in the measurement of the circulation rate by Henderson and
Haggard’s method. J. Physiol. 63, 162–174.
(5) 1931 Cardiac output and blood distribution. J. Physiol. 71, 280–291.
(6)
Properties of the peripheral vascular system and their relation to the systemic output. J.
Physiol. 72, 186–188.
(7) 1932 A study on the influence of adrenaline on the systemic blood flow. J. Physiol. 76, 339–
346.
(9) 1934 (With P. Formijne) The relation of the central nervous system to the increase in systemic
flow produced by occlusion of the thoracic aorta. J. Physiol. 82, 377–384.
(10) 1935 (With A. Samaan) The explanation of the increase in systemic flow caused by occluding
the descending thoracic aorta. J. Physiol. 85, 47–61.
16
(9)
(10)
(11)
(12)
(13)
(14)
(15)
(16)
(17)
(18)
(19)
(20)
(21)
(22)
(23)
(24)
(25)
(26)
(27)
(28)
(29)
(30)
(31)
(32)
(13) 1938 (With W.M. Loughridge) On the accuracy of the thermostromuhr method for measuring
blood flow. J. Physiol. 93, 382–400.
(14) 1939 (With J.L.E. Millen) The blood flow through muscle during sustained contraction. J.
Physiol. 97, 17–31.
(15) 1943 (With O.G. Edholm) The effect of temperature on blood flow and deep temperature of
the human forearm. J. Physiol. 120, 5–20.
(16)
(With W.McK. Bonnar, O.G. Edholm & A.S. Effron) On sympathetic vasoconstrictor
tone in human skeletal muscle. J. Physiol. 105, 21–31.
(17) 1944 (With O.G. Edholm, J. McMichael & E.P. Sharpey-Schafer) Posthaemorrhagic fainting—
study by cardiac output and forearm flow. Lancet i, 489–490.
(18) 1945 (With O.G. Edholm) On the vasodilatation in human skeletal muscle during
posthaemorrhagic fainting. J. Physiol. 104, 161–175.
(21) 1946 (With W.J. Allen, D.P. Anderson, O.G. Edholm & G.W. Manning) Circulatory changes
during fainting and coma caused by oxygen lack. J. Physiol. 104, 426–434.
(27) 1948 (With G.T.C. Hamilton) Results of sympathectomy of the upper limb with special
reference to Raynaud’s disease. Lancet i, 441–444.
(29)
(With G.T.C. Hamilton) Further observations on the results of sympathectomy of the
upper limb. Lancet ii, 770–771.
(31)
(With G.T.C. Hamilton) On the return of sudomotor and vasomotor reflexes to the
sympathectomised hand. J. Physiol. 108, 18P.
(33) 1949 (With A.J. Walker) Return of tone in blood vessels of the upper limb after
sympathectomy. Lancet i, 1035–1039.
(36)
(With A.C. Dornhorst) The blood flow through the human calf during rhythmic exercise.
J. Physiol. 109, 402–411.
(37)
(With H. Konzett) On the actions of noradrenaline, adrenaline and isopropylnoradrenaline on the arterial blood pressure, heart-rate and muscle blood flow in man. J.
Physiol. 110, 194–204.
(38) 1950 (With R.B. Lynn) Circulatory changes in the foot after lumbar sympathectomy. Lancet i,
1105–1108.
(39)
(With A.J. Walker) On the circulatory changes in the hand and foot after sympathectomy.
St Thomas’ Hosp. Rep. 6, 18.
(44) 1951 (With I. Starr) Comparison of the actions of adrenaline and noradrenaline on the
cardiac output in man. Clin. Sci. 10, 297–301.
(48) 1953 (With H.J.C. Swan) Sympathetic control of human blood vessels (Monographs of the
Physiological Society No. 1). London: Edward Arnold.
(53) 1954 (With P. Gaskell, J.T. Shepherd & R.F. Whelan) The effect of noradrenaline infusions on
the blood flow through the human forearm. J. Physiol. 123, 443–450.
(54) 1955 (With H. Hensel & A.H. Kitchin) Die Muskeldurchblutung des Menschen bei indirektor
Erwarmung und Abkuhlung. Pflügers Arch. 261, 199–210.
(57)
Action of epinephrine in man. Shock and circulatory homeostasis. In Transactions of the
Fourth Conference sponsored by the Josiah Macy Foundation (ed. H.D. Green), pp. 9–78.
New York: Madison Printing Co.
(61) 1956 (With A.F. Cobbold) The action of adrenaline on muscle blood flow and blood lactate in
man. J. Physiol. 132, 372–378.
(78) 1963 (With B. Greenwood & R.F. Whelan) Blood flow and venous oxygen saturation during
sustained contraction of the forearm muscles. J. Physiol. 168, 848–856.
(79)
(With B. Greenwood & D.L. Rutt) pH, standard bicarbonate and pCO2 of blood in the
deep veins of the forearm, before, during and after strong sustained contraction of the
forearm muscles. J. Physiol. 169, 34P– 35P.
(86) 1968 Exercise and hyperaemia. Circulation in skeletal muscle. In Proceedings of an
International Symposium held at Smolenice, Czechoslovakia (ed. J. Brod & O. Hudlicke),
p. 121. Oxford: Pergamon Press.
Henry Barcroft
(33)
(34)
(35)
(36)
17
(88) 1969 Correlation of blood supply with metabolism and function. In Proceedings of an
International Symposium held in Tbilisi (ed. G.I. Mzhedlishvili), pp. 48–57. Metsniereba,
Tbilisi: Georgian Academy of Sciences.
(90) 1971 (With T.H. Foley & R.R. McSwiney) Experiments on the liberation of phosphate from
the muscles of the human forearm during vigorous exercise and on the action of sodium
phosphate on forearm muscle blood vessels. J. Physiol. 213, 411–420.
(92) 1972 An enquiry into the nature of the mediator of the vasodilatation in skeletal muscle in
exercise and during circulatory arrest. Review Lecture. J. Physiol. 222, 99P–118P.
(99) 1976 Lymph formation by secretion or filtration? Bayliss–Starling Memorial Lecture. J.
Physiol. 260, 1P–20P.

F.R.S. 1953 11 January 1998: Elected − Henry Barcroft. 18 October

Transcription

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