Alan_Hodgkin

Alan Hodgkin
Sir Alan Lloyd Hodgkin OM KBE FRS[1] (5 February

Sir
1914 – 20 December 1998) was an English
physiologist and biophysicist who shared the 1963 Alan Lloyd Hodgkin
OM KBE FRS
Nobel Prize in Physiology or Medicine with Andrew
Huxley and John Eccles.
Early life and education

Hodgkin was born in Banbury, Oxfordshire, on 5
February 1914. He was the oldest of three sons of
Quakers George Hodgkin and Mary Wilson Hodgkin.
His father was the son of Thomas Hodgkin and had
read for the Natural Science Tripos at Cambridge
where he had befriended electrophysiologist Keith
Lucas.[2]
Because of poor eyesight, he was unable to study

medicine and eventually ended up working for a bank
in Banbury. As members of the Society of Friends, Born 5 February 1914
George and Mary opposed the Military Service Act of Banbury, Oxfordshire, England
1916, which introduced conscription, and had to Died 20 December 1998 (aged 84)
endure a great deal of abuse from their local Cambridge, England
community, including an attempt to throw George in Nationality English
one of the town canals.[3] In 1916, George Hodgkin
Citizenship British
travelled to Armenia as part of an investigation of
distress. Moved by the misery and suffering of Alma mater University of Cambridge
Armenian refugees he attempted to go back there in Known for Hodgkin cycle
1918 on a route through the Persian Gulf (as the Hodgkin–Huxley model
northern route was closed because of the October Hodgkin–Huxley sodium channels
Revolution in Russia). He died of dysentery in Goldman–Hodgkin–Katz flux
Baghdad on 24 June 1918, just a few weeks after his equation
youngest son, Keith, had been born.[4] Goldman–Hodgkin–Katz voltage
equation
From an early life on, Hodgkin and his brothers were
Spouse Marion Rous
encouraged to explore the country around their home,
Children Sarah, Deborah, Jonathan
which instilled in Alan an interest in natural history,
Hodgkin, and Rachel
particularly ornithology. At the age of 15, he helped
Wilfred Backhouse Alexander with surveys of Relatives Hodgkin family
heronries and later, at Gresham's School, he Awards Royal Medal (1958)
overlapped and spent a lot of time with David
Lack.[5][6] In 1930, he was the winner of a bronze Physiological Society Annual
medal in the Public Schools Essay Competition Review Prize Lecture (1976)
organised by the Royal Society for the Protection of Nobel Prize in Physiology or
Birds.[7] Medicine (1963)
Copley Medal (1965)
School and university Foreign Associate of the National
Academy of Sciences (1964)
Alan started his education at The Downs School where
his contemporaries included future scientists Frederick Scientific career
Sanger, Alec Bangham, "neither outstandingly brilliant Fields Physiology
at school" according to Hodgkin,[8] as well as future Biophysics
artists Lawrence Gowing and Kenneth Rowntree. After
the Downs School, he went on to Gresham's School where he overlapped with future composer Benjamin
Britten as well as Maury Meiklejohn.[9] He ended up receiving a scholarship at Trinity College,
Cambridge in botany, zoology and chemistry.[10]
Between school and college, he spent May 1932 at the Freshwater Biological Station at Wray Castle
based on a recommendation of his future Director of Studies at Trinity, Carl Pantin.[11] After Wray Castle,
he spent two months with a German family in Frankfurt as "in those days it was thought highly desirable
that anyone intending to read science should have a reasonable knowledge of German."[12] After his
return to England in early August 1932, his mother Mary was remarried to Lionel Smith (1880–1972),[13]
the eldest son of A. L. Smith, whose daughter Dorothy was also married to Alan's uncle Robert Howard
Hodgkin.[14]
In the autumn of 1932, Hodgkin started as a freshman scholar at Trinity College where his friends
included classicists John Raven and Michael Grant, fellow-scientists Richard Synge and John H.
Humphrey,[15] as well as Polly and David Hill, the children of Nobel laureate Archibald Hill.[5][16] He
took physiology with chemistry and zoology for the first two years, including lectures by Nobel laureate
E.D. Adrian.[17] For Part II of the tripos he decided to focus on physiology instead of zoology.
Nevertheless, he participated in a zoological expedition to the Atlas Mountains in Morocco led by John
Pringle in 1934.[18] He finished Part II of the tripos in July 1935 and stayed at Trinity as a research
fellow.[19]
During his studies, Hodgkin, who described himself as "having been brought up as a supporter of the
British Labour Party"[20] was friends with communists[21] and actively participated in the distribution of
anti-war pamphlets.[22] At Cambridge, he knew James Klugmann[23] and John Cornford,[24] but he
emphasised in his autobiography that none of his friends "made any serious effort to convert me [to
Communism], either then or later."[25] From 1935 to 1937, Hodgkin was a member of the Cambridge
Apostles.[26]
Pre-war research
Hodgkin started conducting experiments on how electrical activity is transmitted in the sciatic nerve of
frogs in July 1934.[27] He found that a nerve impulse arriving at a cold or compression block, can
decrease the electrical threshold beyond the block, suggesting that the impulse produces a spread of an
electrotonic potential in the nerve beyond the block.[28] In 1936, Hodgkin was invited by Herbert Gasser,
then director of the Rockefeller Institute in New York
City, to work in his laboratory during 1937–38. There
he met Rafael Lorente de Nó[29] and Kenneth Stewart
Cole with whom he ended up publishing a paper.[30]
During that year he also spent time at the Woods Hole
Marine Biological Laboratory where he was
introduced to the squid giant axon,[31] which ended
up being the model system with which he conducted
most of the research that eventually led to his Nobel
Trinity College at the University of Cambridge
Prize. In the spring of 1938, he visited Joseph where Hodgkin was a student from 1932 to 1935,
Erlanger at Washington University in St. Louis who worked as a research fellow from 1936 on and
told him he would take Hodgkin's local circuit theory finally served as Master from 1978 to 1984
of nerve impulse propagation seriously if he could
show that altering the resistance of the fluid outside a
nerve fibre made a difference to the velocity of nerve impulse conduction.[32] Working with single nerve
fibres from shore crabs and squids, he showed that the conduction rate was much faster in seawater than
in oil, providing strong evidence for the local circuit theory.[33]
After his return to Cambridge he started collaborating with Andrew Huxley who had entered Trinity as a
freshman in 1935, three years after Hodgkin.[34] With a £300 equipment grant from the Rockefeller
Foundation, Hodgkin managed to set up a similar physiology setup to the one he had worked with at the
Rockefeller Institute. He moved all his equipment to the Plymouth Marine Laboratory in July 1939.[35]
There, he and Huxley managed to insert a fine cannula into the giant axon of squids and record action
potentials from inside the nerve fibre. They sent a short note of their success to Nature just before the
outbreak of World War II.[36]
Wartime activities
Despite his Quaker upbringing, Hodgkin was eager to join the war effort as contact with the Nazis during
his stay in Germany in 1932 had removed all his pacifist beliefs. His first post was at the Royal Aircraft
Establishment where he worked on issues in aviation medicine, such as oxygen supply for pilots at high
altitudes and the decompression sickness caused by nitrogen bubbles coming out of the blood.[37] In
February 1940 he transferred to the Telecommunications Research Establishment (TRE) where he
worked on the development of centimetric radar, including the design of the village Inn AGLT airborne
gun-laying system. He was a member of E.G. Bowen's group in St Athan in South Wales and lived in a
local guest house together with John Pringle and Robert Hanbury Brown. The group moved to
Swanage in May 1940 where Pringle replaced Bowen as leader of the group.[38] In March 1941, Hodgkin
flew on the test flight of a Bristol Blenheim fitted with the first airborne centimetric radar system. In
February and March 1944, he visited the MIT Radiation Laboratory to help foster the interchange of
information on developments in radar between Britain and America.[39]
Providing a readable account of the little-known piece of military history that he was a part of during
World War II was a main motivation for Hodgkin to write his autobiography Chance and Design:
Reminiscences of Science in Peace and War.[40]
1945–1963: Action potential theory and Nobel Prize

As the Allied Forces' invasion of France and their
continued advance towards Germany in autumn 1944
suggested an end of the war in the foreseeable future,
Hodgkin started to plan his return to a career in
research at Cambridge. He renewed his collaboration
with W. A. H. Rushton and they published an article
on how to calculate a nerve fibre's membrane
resistance, membrane capacity, its axoplasm's
resistance, and the resistance of the external fluid in
which the fibre is placed, from experimental
observations.[41]
Basic components of Hodgkin–Huxley-type
After being released from military service in August models. Hodgkin–Huxley type models represent
the biophysical characteristics of cell membranes.
1945 upon Adrian's request, Hodgkin was able to
The lipid bilayer is represented as a capacitance
restart his experiments in collaboration with Bernard (Cm). Voltage-gated and leak ion channels are
Katz and his pre-war collaborator Andrew Huxley. represented by nonlinear (gn) and linear (gL)
They spent the summers of 1947, 1948, and 1949 at conductances, respectively. The electrochemical
the Plymouth Marine Laboratory where they gradients driving the flow of ions are represented
continued to measure resting and action potentials by batteries (E), and ion pumps and exchangers
from inside the giant axon of the squid.[42] Together are represented by current sources (Ip).
with Katz, he provided evidence that the permeability

of the neuronal cell membrane for sodium increased during an action potential, thus allowing sodium ions
to diffuse inward.[43] The data they had obtained in 1949 resulted in a series of five papers published in
The Journal of Physiology that described what became later known as the Hodgkin–Huxley model of the
action potential and eventually earned Hodgkin and Huxley the Nobel Prize in Physiology or
Medicine.[44][45][46][47][48] Building on work by Kenneth S. Cole[49] they used a technique of
electrophysiology, known as the voltage clamp to measure ionic currents through the membranes of squid
axons while holding the membrane voltage at a set level. They proposed that the characteristic shape of
the action potential is caused by changes in the selective permeability of the membrane for different ions,
specifically sodium, potassium, and chloride. A model that relies on a set of differential equations and
describes each component of an excitable cell as an electrical element was in good agreement with their
empirical measurements.[48][50]
The cell membrane depolarisation sequence where a small depolarization leads to an increase in sodium
permeability, which leads to influx of sodium ions, which in turn depolarizes the membrane even
more[51] is now known as the Hodgkin cycle.[52]
In addition, Hodgkin and Huxley's findings led them to hypothesize the existence of ion channels on cell
membranes, which were confirmed only decades later. Confirmation of ion channels came with the
development of the patch clamp leading to a Nobel prize in 1991 for Erwin Neher and Bert Sakmann, and
in 2003 for Roderick MacKinnon.[53]
After establishing ion movements across a selectively

permeable cell membrane as the mechanism of the
action potential, Hodgkin turned his attention to how
the ionic interchange that occurs during the action
potential could be reversed afterwards. Together with
Richard Keynes he demonstrated that in addition to
the changes in permeability that lead to an action
potential, there is a secretory mechanism that ejects
sodium and absorbs potassium against the
electrochemical gradients.[54] A few years later, the
Danish scientist Jens Christian Skou discovered the
enzyme Na+/K+-ATPase that uses ATP to export three sodium ions in exchange for two potassium ions
that are imported,[55] for which he received the Nobel Prize in Chemistry in 1997.[56]
Hodgkin was nominated for the Nobel Prize in Physiology or Medicine in 1953 by Lord Adrian.[57] In
October 1961, he was told by Swedish journalists that he, Huxley, and Eccles had been awarded the
Nobel Prize. This turned out to be a false alarm, however, when shortly thereafter it was announced that
the 1961 Prize was awarded to Georg von Békésy. It was only two years later that Hodgkin, Huxley, and
Eccles were finally awarded the Prize "for their discoveries concerning the ionic mechanisms involved in
excitation and inhibition in the peripheral and central portions of the nerve cell membrane".[58][59] During
the Nobel Banquet on 10 December 1963, Hodgkin gave the traditional speech on behalf of the three
neurophysiologists, thanking the king and the Nobel Committee for Physiology or Medicine for the
award.[60] Incidentally, Hodgkin and his wife attended the Nobel Prize ceremony a second time, three
years later, when Hodgkin's father-in-law, Francis Peyton Rous, was awarded the 1966 Nobel Prize in
Physiology or Medicine.[61]
Later career and administrative positions

From 1951 to 1969, Hodgkin was the Foulerton Professor of the Royal Society at Cambridge. In 1970 he
became the John Humphrey Plummer Professor of Biophysics at Cambridge. Around this time he also
ended his experiments on nerve at the Plymouth Marine Laboratory and switched his focus to visual
research which he could do in Cambridge with the help of others while serving as president of the Royal
Society. Together with Denis Baylor and Peter Detwiler he published a series of papers on turtle
photoreceptors.[62][63][64][65][66][67][68]
From 1970 to 1975 Hodgkin served as the 53rd president of the Royal Society (PRS). During his tenure
as PRS, he was knighted in 1972 and admitted into the Order of Merit in 1973.[69] From 1978 to 1985 he
was the 34th Master of Trinity College, Cambridge.[70]
He served on the Royal Society Council from 1958 to 1960 and on the Medical Research Council from
1959 to 1963. He was foreign secretary of the Physiological Society from 1961 to 1967. He also held
additional administrative posts such as Chancellor, University of Leicester, from 1971 to 1984
Awards and honours

1988 – W.H. Helmerich III Award of the Retina Research Foundation[71]
1983 – Lord Crook Medal of the Worshipful Company of Spectacle Makers[72]
1982 – F.O. Schmitt Medal and Award 1983
1977 – Hon. DSc, University of Oxford
1975 – Hon. Fellow, Indian Academy of Sciences
1974 – Foreign Associate, National Academy of Sciences of the USA[73]
1973 – Order of Merit (O.M.)
1973 – Foreign Member, Royal Swedish Academy of Sciences (Medical Sciences, VIII
Class)
1972 – Knight Commander of the Order of the British Empire (K.B.E.)
1972 – Hon. Fellow, Indian National Science Academy
1970 – President of the Royal Society (PRS)[1]
1968 – Member, Pontifical Academy of Sciences
1968 – Foreign Member, American Philosophical Society
1966 – President of the Marine Biological Association of the United Kingdom
1965 – Copley Medal of the Royal Society
1964 – Foreign Member, Royal Danish Academy of Sciences and Letters
1964 – Member, German National Academy of Sciences Leopoldina[74]
1963 – Nobel Prize for Physiology or Medicine together with Andrew Fielding Huxley and
John Carew Eccles (for their research on synapses)[75]
1962 – Foreign Hon. Member, American Academy of Arts and Sciences
1958 – Royal Medal of The Royal Society
1958 – Hon. MD, University of Louvain
1956 – Hon. MD, University of Berne
1955 – Baly Medal of the Royal College of Physicians
1948 – Fellow of the Royal Society (FRS)[1]
A portrait of Hodgkin by Michael Noakes hangs in Trinity College's collection.[76]
Publications
The Conduction of the Nervous Impulse (1964)
Chance and Design: Reminiscences of Science in Peace and War (1992)
Personal life
During his stay at the Rockefeller Institute in 1937, Hodgkin got to know the American pathologist
Francis Peyton Rous[77] who was later awarded the 1966 Nobel Prize in Physiology or Medicine.[78]
When Rous invited him for dinner to his home, Hodgkin got to know Rous' daughter, Marni, who was
then a student at Swarthmore College.[77] He proposed to her before going back to England in 1938, but
she rejected him.[79] When Hodgkin briefly returned to the US in 1944 (see Wartime activities), they
reunited and got married on 31 March.[80]
Their first daughter, Sarah, was born in April 1945, shortly before the Hodgkins moved back to
Cambridge.[81] They had three more children: Deborah Hodgkin (born 2 May 1947),[82] Jonathan
Hodgkin (born 24 August 1949),[83] and Rachel Hodgkin (born June 1951).[84] Marni became a
Children's Book Editor at Macmillan Publishing Company and a successful writer of children's literature,
including Young Winter's Tales and Dead Indeed. Jonathan Hodgkin became a molecular biologist at
Cambridge University. Deborah Hodgkin is also a successful psychologist.
Thomas Hodgkin (1798–1866), who first described Hodgkin's lymphoma, was Alan Hodgkin's great-
uncle.[85]
Death
Hodgkin suffered from a series of medical problems that began soon after his retirement as Master of
Trinity. In 1989 he had surgery to relieve pressure on the spinal cord from one of the intervertebral discs
in his neck, which left him unable to walk without support, and with progressive disablement.[1] Hodgkin
died in 1998 in Cambridge.[86]
See also
List of presidents of the Royal Society
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Bibliography
Hodgkin, Alan (1992). Chance & Design - Reminiscences of Science in Peace and War.
Cambridge University Press. ISBN 978-0-521-45603-6.
External links
The Master of Trinity (http://www.trin.cam.ac.uk/index.php?pageid=172) at Trinity College,
Cambridge
Alan Hodgkin (https://www.nobelprize.org/laureate/376) on Nobelprize.org including the
Nobel Lecture on 11 December 1963 The Ionic Basis of Nervous Conduction
Portraits of Alan Hodgkin (https://www.npg.org.uk/collections/search/person.php?LinkID=mp
05411) at the National Portrait Gallery, London
BBC obituary (http://news.bbc.co.uk/1/hi/sci/tech/239286.stm)
Action Potential Paper (http://www.sfn.org/-/media/SfN/Documents/ClassicPapers/ActionPot
entials/hodgkin5.ashx?la=en&hash=B7491DA5CF6B4F269B13D2AAAD37F77BFBDCE5D
0)
Imperial War Museum Interview (http://www.iwm.org.uk/collections/item/object/80012991)
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Alan Hodgkin

Sir Alan Lloyd Hodgkin OM KBE FRS[1] (5 February

Early life and education

Because of poor eyesight, he was unable to study

1945–1963: Action potential theory and Nobel Prize

with Katz, he provided evidence that the permeability

After establishing ion movements across a selectively

Later career and administrative positions

Awards and honours

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