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{{Short description|British physiologist and biophysicist}}
{{EngvarB|date=July 2020}}
{{Use dmy dates|date=December 2019}}
{{Infobox scientist
| honorific_prefix  = Sir
| name              = Alan Lloyd Hodgkin
| honorific_suffix  = {{postnominals|country=GBR|OM|KBE|FRS|size=100%}}
| image             = Alan Lloyd Hodgkin nobel.jpg
| image_size        = 
| caption           = 
| birth_name        = 
| birth_date        = {{birth date|df=yes|1914|02|05}}
| birth_place       = [[Banbury]], Oxfordshire, England
| death_date        = {{death date and age|df=yes|1998|12|20|1914|02|05}}
| death_place       = [[Cambridge]], England
| known_for         = [[Hodgkin cycle]]<br />[[Hodgkin–Huxley model]]<br />[[Sodium channel|Hodgkin–Huxley sodium channels]]<br />[[Goldman–Hodgkin–Katz flux equation]]<br />[[Goldman–Hodgkin–Katz voltage equation]]
| alma_mater        = [[University of Cambridge]]
| spouse            = [[Marni Hodgkin|Marion Rous]]
| children          = Sarah, Deborah, [[Jonathan Hodgkin]], and Rachel
| prizes            = {{Plainlist|
* [[Royal Medal]] (1958)
* [[Physiological Society Annual Review Prize Lecture]] (1976)
* [[Nobel Prize in Physiology or Medicine]] (1963)
* [[Copley Medal]] (1965)
}}
| field             = [[Physiology]]<br />[[Biophysics]]
| relatives         = [[Hodgkin family]]
}}
''' Sir Alan Lloyd Hodgkin''' (5 February 1914 – 20 December 1998) was a British [[physiology|physiologist]] and [[biophysics|biophysicist]] who shared the 1963 [[Nobel Prize in Physiology or Medicine]] with [[Andrew Huxley]] and [[John Eccles (neurophysiologist)|John Eccles]].

==Early life and education==
{{See also|Hodgkin family}}
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 (historian)|Thomas Hodgkin]] and had read for the [[Natural Sciences (Cambridge)|Natural Science Tripos]] at [[University of Cambridge|Cambridge]] where he had befriended electrophysiologist [[Keith Lucas (scientist)|Keith Lucas]].<ref>{{harvnb|Hodgkin|1992|p=5}}</ref>

Because of poor eyesight, he was unable to study medicine and eventually ended up working for a bank in Banbury. As members of the [[Quakers|Society of Friends]], George and Mary opposed the [[Military Service Act 1916|Military Service Act of 1916]], which introduced conscription, and had to endure a great deal of abuse from their local community, including an attempt to throw George in one of the town canals.<ref>{{harvnb|Hodgkin|1992|pp=7–9}}</ref> In 1916, George Hodgkin travelled to Armenia as part of an investigation of distress. Moved by the misery and suffering of [[Armenian genocide|Armenian refugees]] he attempted to go back there in 1918 on a route through the Persian Gulf (as the northern route was closed because of the [[October Revolution]] in Russia). He died of dysentery in [[Baghdad]] on 24 June 1918, just a few weeks after his youngest son, Keith, had been born.<ref>{{harvnb|Hodgkin|1992|p=10}}</ref>

From an early life on, Hodgkin and his brothers were encouraged to explore the country around their home, which instilled in Alan an interest in [[Natural History|natural history]], particularly [[ornithology]]. At the age of 15, he helped [[Wilfred Backhouse Alexander]] with surveys of [[Heronry|heronries]] and later, at [[Gresham's School]], he overlapped and spent a lot of time with [[David Lack]].<ref name="Hodgkin 1992 11–12">{{harvnb|Hodgkin|1992|pp=11–12}}</ref><ref>{{cite journal|last=Hodgkin|first=Alan|title=Beginning: Some Reminiscences of my early life (1914–1947)|journal=Annual Review of Physiology|year=1983|volume=45|pages=1–16|doi=10.1146/annurev.ph.45.030183.000245|pmid=6342510}}</ref> In 1930, he was the winner of a bronze medal in the Public Schools Essay Competition organised by the [[Royal Society for the Protection of Birds]].<ref>''Protection of Birds Measures Urged By Royal Society'' in ''The Times'', Saturday, 29 March 1930; pg. 14; Issue 45474; col C</ref>

===School and university===
Alan started his education at [[The Downs Malvern|The Downs School]] where his contemporaries included future scientists [[Frederick Sanger]], [[Alec Bangham]], "neither outstandingly brilliant at school" according to Hodgkin,<ref>{{harvnb|Hodgkin|1992|p=25}}</ref> as well as future 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 [[M. F. M. Meiklejohn|Maury Meiklejohn]].<ref>{{harvnb|Hodgkin|1992|pp=27–29}}</ref> He ended up receiving a scholarship at [[Trinity College, Cambridge]] in botany, zoology and chemistry.<ref>{{harvnb|Hodgkin|1992|p=30}}</ref>

Between school and college, he spent May 1932 at the [[Freshwater Biological Association|Freshwater Biological Station]] at [[Wray Castle]] based on a recommendation of his future Director of Studies at [[Trinity College, Cambridge|Trinity]], [[Carl Pantin]].<ref>{{harvnb|Hodgkin|1992|p=31}}</ref> 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."<ref>{{harvnb|Hodgkin|1992|p=34}}</ref> After his return to England in early August 1932, his mother Mary was remarried to Lionel Smith (1880–1972),<ref>{{cite web|url=https://www.sant.ox.ac.uk/sites/default/files/gb165-0266-alf-smith-collection.pdf|title=SMITH, Arthur Lionel Forster|access-date=2023-02-03}}</ref> the eldest son of [[Arthur Smith (historian)|A. L. Smith]], whose daughter Dorothy was also married to Alan's uncle [[Robert Howard Hodgkin]].<ref>{{harvnb|Hodgkin|1992|p=40}}</ref>

In the autumn of 1932, Hodgkin started as a freshman scholar at Trinity College where his friends included [[Classics|classicists]] [[John Raven]] and [[Michael Grant (classicist)|Michael Grant]], fellow-scientists [[Richard Laurence Millington Synge|Richard Synge]] and [[John H. Humphrey]],<ref>{{harvnb|Hodgkin|1992|p=49}}</ref> as well as [[Polly Hill (economist)|Polly]] and [[David Keynes Hill|David Hill]], the children of [[List of Nobel laureates in Physiology or Medicine|Nobel laureate]] [[A.V. Hill|Archibald Hill]].<ref name="Hodgkin 1992 11–12"/><ref>{{cite journal|last=Hodgkin|first=Alan|title=Beginning: Some Reminiscences of my early life (1914–1947)|journal=Annual Review of Physiology|year=1983|volume=45|pages=1–16|doi=10.1146/annurev.ph.45.030183.000245|pmid=6342510}}</ref> He took physiology with chemistry and zoology for the first two years, including lectures by [[List of Nobel laureates in Physiology or Medicine|Nobel laureate]] [[Edgar Adrian|E.D. Adrian]].<ref>{{harvnb|Hodgkin|1992|p=50}}</ref> For Part II of the [[Natural Sciences (Cambridge)|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 William Sutton Pringle|John Pringle]] in 1934.<ref>{{harvnb|Hodgkin|1992|p=55}}</ref> He finished Part II of the tripos in July 1935 and stayed at [[Trinity College, Cambridge|Trinity]] as a research fellow.<ref>{{harvnb|Hodgkin|1992|p=66}}</ref>

During his studies, Hodgkin, who described himself as "having been brought up as a supporter of the British Labour Party"<ref>{{harvnb|Hodgkin|1992|p=38}}</ref> was friends with communists<ref>{{harvnb|Hodgkin|1992|p=48}}</ref> and actively participated in the distribution of anti-war pamphlets.<ref>{{harvnb|Hodgkin|1992|p=53}}</ref> At Cambridge, he knew [[James Klugmann]]<ref>{{harvnb|Hodgkin|1992|p=79}}</ref> and [[John Cornford]],<ref>{{harvnb|Hodgkin|1992|pp=83–87}}</ref> but he emphasised in his autobiography that none of his friends "made any serious effort to convert me [to [[Communism]]], either then or later."<ref>{{harvnb|Hodgkin|1992|p=80}}</ref> From 1935 to 1937, Hodgkin was a member of the [[Cambridge Apostles]].<ref>{{harvnb|Hodgkin|1992|p=87}}</ref>

== Pre-war research==
[[File: Trinity_College_-_Great_Court_02.jpg|thumb|right|upright=1.35|Trinity College at the [[University of Cambridge]] where Hodgkin was a student from 1932 to 1935, worked as a research fellow from 1936 on and finally served as [[Master (college)|Master]] from 1978 to 1984]]
Hodgkin started conducting experiments on how electrical activity is transmitted in the [[sciatic nerve]] of frogs in July 1934.<ref>{{harvnb|Hodgkin|1992|p=63}}</ref> He found that a [[Action potential|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 [[Graded potential|electrotonic potential]] in the nerve beyond the block.<ref>{{cite journal|last=Hodgkin|first=Alan|title=Evidence for Electrical Transmission in Nerve. Part I|journal=The Journal of Physiology|year=1937|volume=90|issue=2|pages=183–210|doi=10.1113/jphysiol.1937.sp003507|pmid=16994885|pmc=1395060}}</ref> In 1936, Hodgkin was invited by [[Herbert Spencer Gasser|Herbert Gasser]], then director of the [[Rockefeller University|Rockefeller Institute]] in New York City, to work in his laboratory during 1937–38. There he met [[Rafael Lorente de Nó]]<ref>{{harvnb|Hodgkin|1992|p=92}}</ref> and [[Kenneth Stewart Cole]] with whom he ended up publishing a paper.<ref>{{cite journal|last1=Cole|first1=Kenneth|last2=Hodgkin|first2=Alan|title=Membrane and Protoplasm Resistance in the Squid Giant Axon|journal=The Journal of General Physiology|year=1939|volume=22|issue=5|pages=671–687|doi=10.1085/jgp.22.5.671|pmid=19873126|pmc=2142005}}</ref> During that year he also spent time at the [[Marine Biological Laboratory|Woods Hole Marine Biological Laboratory]] where he was introduced to the [[squid giant axon]],<ref>{{cite journal|last=Hodgkin|first=Alan|title=Beginning: Some Reminiscences of my early life (1914–1947)|journal=Annual Review of Physiology|year=1983|volume=45|pages=1–16|doi=10.1146/annurev.ph.45.030183.000245|pmid=6342510}}</ref> which ended up being the model system with which he conducted most of the research that eventually led to his Nobel Prize. 
In the spring of 1938, he visited [[Joseph Erlanger]] at [[Washington University in St. Louis]] who told him he would take Hodgkin's local circuit theory 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.<ref>{{harvnb|Hodgkin|1992|p=113}}</ref> Working with single nerve fibres from [[Carcinus maenas|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.<ref>{{cite journal|last=Hodgkin|first=Alan|title=The Relation between Conduction Velocity and the Electrical Resistance outside a Nerve Fibre|journal=The Journal of Physiology|year=1939|volume=94|pages=560–570|issue=4|doi=10.1113/jphysiol.1939.sp003702|pmid=16995066|pmc=1393884}}</ref>

After his return to Cambridge he started collaborating with [[Andrew Huxley]] who had entered [[Trinity College, Cambridge|Trinity]] as a freshman in 1935, three years after Hodgkin.<ref>{{Cite book|publisher = Society for Neuroscience|isbn = 0-12-660246-8|pages = 282–318|editor-first = Larry R.|editor-last = Squire|last = Huxley|first = Andrew| author-link = Andrew Huxley|title = The history of neuroscience in autobiography|chapter = Andrew Huxley|location = Washington DC|year = 2004|chapter-url = http://www.sfn.org/-/media/SfN/Documents/TheHistoryofNeuroscience/Volume-4/c8.ashx?la=en&hash=2B26C8338409076E6889C89B47EAFB0F6178167B}}</ref> 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.<ref>{{cite journal|last=Hodgkin|first=Alan|title=Chance and Design in Electrophysiology: An Informal Account of Certain Experiments on Nerve Carried out between 1934 and 1952|journal=Journal of Physiology|year=1976|volume=263|issue=1|pages=1–21|doi=10.1113/jphysiol.1976.sp011620|pmid=796420|pmc=1307686}}</ref> 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 (journal)|'' Nature'']] just before the outbreak of [[World War II]].<ref>{{cite journal|doi=10.1038/144710a0|title=Action Potentials Recorded from Inside a Nerve Fibre|year=1939|last1=Hodgkin|first1=A. L.|last2=Huxley|first2=A. F.|journal=Nature|volume=144|issue=3651|pages=710–711|bibcode = 1939Natur.144..710H |s2cid=4104520}}</ref>

==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 [[Pacifism|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.<ref>{{harvnb|Hodgkin|1992|p=140}}</ref> 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 [[Automatic Gun-Laying Turret|village Inn]] AGLT airborne gun-laying system. He was a member of [[Edward George Bowen|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.<ref>{{harvnb|Hodgkin|1992|pp=141–156}}</ref> 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.<ref>{{harvnb|Hodgkin|1992|pp=233–239}}</ref>

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''.<ref>{{harvnb|Hodgkin|1992|pp=ix–xi}}</ref>

==1945–1963: Action potential theory and Nobel Prize==
[[File:Hodgkin-Huxley.svg|thumb|right|upright=1.35|Basic components of Hodgkin–Huxley-type models. Hodgkin–Huxley type models represent the biophysical characteristics of cell membranes. The lipid bilayer is represented as a capacitance (''C''<SUB>m</SUB>). Voltage-gated and leak ion channels are represented by nonlinear (''g''<SUB>n</SUB>) and linear (''g''<SUB>L</SUB>) conductances, respectively. The electrochemical gradients driving the flow of ions are represented by batteries (E), and ion pumps and exchangers are represented by current sources (''I''<SUB>p</SUB>).]]

As the [[Allies of World War II|Allied Forces']] [[Invasion of Normandy|invasion of France]] and their continued [[Allied advance from Paris to the Rhine|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.<ref>{{cite journal|doi=10.1098/rspb.1946.0024|pmid=20281590|bibcode=1946RSPSB.133..444H|title=The electrical constants of a crustacean nerve fibre|year=1946|last1=Hodgkin|first1=A. L.|last2=Rushton|first2=W.A. H.|journal=Proceedings of the Royal Society of London. Series B|volume=133|issue=873|pages=444–479|doi-access=free}}</ref>

After being released from military service in August 1945 upon [[Edgar Adrian|Adrian's]] request, Hodgkin was able to restart his experiments in collaboration with [[Bernard Katz]] and his pre-war collaborator [[Andrew Huxley]]. They spent the summers of 1947, 1948, and 1949 at the [[Plymouth Marine Laboratory]] where they continued to measure resting and action potentials from inside the giant axon of the squid.<ref>{{harvnb|Hodgkin|1992|pp=271–290}}</ref> Together 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.<ref>{{cite journal|doi=10.1113/jphysiol.1949.sp004310|pmid=18128147|title=The Effect of Sodium Ions on the Electrical Activity of the Giant Axon of the Squid|year=1949|last1=Hodgkin|first1=A. L.|last2=Katz|first2=B.|journal=Journal of Physiology|volume=108|issue=1|pages=37–77|pmc=1392331}}</ref> 
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]].<ref>{{cite journal|doi=10.1113/jphysiol.1952.sp004716|pmid=14946712|title=Measurement of Current-Voltage Relations in the Membrane of the Gian Axon of Loligo|year=1952|last1=Hodgkin|first1=A. L.|last2=Huxley|first2=A. F.|last3=Katz|first3=B.|journal=The Journal of Physiology|volume=116|issue=4|pages=424–448|pmc=1392219}}</ref><ref>{{cite journal|doi=10.1113/jphysiol.1952.sp004717|pmid=14946713|title=Currents carried by sodium and potassium ions through the membrane of the giant axon of Loligo|year=1952|last1=Hodgkin|first1=A. L.|last2=Huxley|first2=A. F.|journal=The Journal of Physiology|volume=116|issue=4|pages=449–472|pmc=1392213}}</ref><ref>{{cite journal|doi=10.1113/jphysiol.1952.sp004718|pmid=14946714|title=The components of membrane conductance in the giant axon of Loligo|year=1952|last1=Hodgkin|first1=A. L.|last2=Huxley|first2=A. F.|journal=The Journal of Physiology|volume=116|issue=4|pages=473–496|pmc=1392209}}</ref><ref>{{cite journal|doi=10.1113/jphysiol.1952.sp004719|pmid=14946715|title=The dual effect of membrane potential on sodium conductance in the giant axon of loligo|year=1952|last1=Hodgkin|first1=A. L.|last2=Huxley|first2=A. F.|journal=The Journal of Physiology|volume=116|issue=4|pages=497–506|pmc=1392212}}</ref><ref name=HodgkinHuxley1952>{{cite journal|doi=10.1113/jphysiol.1952.sp004764|pmid=12991237|title=A Quantitative Description of Membrane Current and its Application to Conduction and Excitation in Nerve|year=1952|last1=Hodgkin|first1=A. L.|last2=Huxley|first2=A. F.|journal=The Journal of Physiology|volume=117|issue=4|pages=500–544|pmc=1392413}}</ref> Building on work by [[Kenneth Stewart Cole|Kenneth S. Cole]]<ref>{{cite journal|title=Dynamic electrical characteristics of the squid axon membrane.|year=1949|last1=Cole|first1=Kenneth S.|journal=Archives des sciences physiologique|volume=3|issue=2|pages=253–258}}</ref> 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 [[Hodgkin–Huxley model|model]] that relies on a set of [[differential equation]]s and describes each component of an excitable cell as an electrical element was in good agreement with their empirical measurements.<ref name=HodgkinHuxley1952 /><ref>{{harvnb|Hodgkin|1992|pp=299–303}}</ref>

[[File:Hodgkin-Cycle.jpg|thumb|right|upright=1.35]]
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<ref>{{cite journal|last=Hodgkin|first=Alan|title=The ionic basis of electrical activity in nerve and muscle|journal=Biological Reviews|year=1951|volume=26|issue=4|pages=339–409|doi=10.1111/j.1469-185X.1951.tb01204.x|s2cid=86282580}}</ref> is now known as the [[Hodgkin cycle]].<ref>{{cite journal|last=Noble|first=D.|title=Biophysics and systems biology|journal=Philosophical Transactions of the Royal Society A: Mathematical, Physical and Engineering Sciences|year=2010|volume=368|issue=1914|pages=1125–1139|doi=10.1098/rsta.2009.0245|pmid=20123750| pmc=3263808 |bibcode = 2010RSPTA.368.1125N }}</ref>

In addition, Hodgkin and Huxley's findings led them to hypothesize the existence of [[ion channel]]s 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]].<ref>{{cite journal|last=Schwiening|first=C. J.|title=A brief historical perspective: Hodgkin and Huxley|journal=The Journal of Physiology|year=2012|volume=590|issue=11|pages=2571–2575|doi=10.1113/jphysiol.2012.230458|pmid=22787170|pmc=3424716}}</ref>

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.<ref>{{cite journal|doi=10.1113/jphysiol.1955.sp005290|title=Active transport of cations in giant axons from Sepia and Loligo|year=1955|last1=Hodgkin|first1=A. L.|last2=Keynes|first2=R. D.|journal=The Journal of Physiology|volume=128|issue=1|pages=28–60|pmid=14368574|pmc=1365754}}</ref> A few years later, the Danish scientist [[Jens Christian Skou]] discovered the enzyme [[Na+/K+-ATPase]] that uses [[adenosine triphosphate|ATP]] to export three sodium ions in exchange for two potassium ions that are imported,<ref>{{cite journal|doi=10.1016/0006-3002(57)90343-8|title=The influence of some cations on an adenosine triphosphatase from peripheral nerves|year=1957|last1=Skou|first1=J. C.|journal=Biochimica et Biophysica Acta|volume=23|issue=2|pages=394–401|pmid=13412736|s2cid=32516710 }}</ref> for which he received the [[Nobel Prize in Chemistry]] in 1997.<ref>{{cite web|title=The Nobel Prize in Chemistry 1997|url=https://www.nobelprize.org/prizes/chemistry/1997/summary/|website=Nobelprize.org|publisher=Nobel Media AB|access-date=12 July 2020}}</ref>

Hodgkin was nominated for the Nobel Prize in Physiology or Medicine in 1953 by [[Edgar Adrian|Lord Adrian]].<ref>{{cite web|url=https://www.nobelprize.org/nomination/archive/show.php?id=13161|title=Nomination Archive|website=nobelprize.org|date=April 2020|access-date=25 June 2020}}</ref> In October 1961, he was told by Swedish journalists that he, [[Andrew Huxley|Huxley]], and [[John Eccles (neurophysiologist)|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, [[Andrew Huxley|Huxley]], and [[John Eccles (neurophysiologist)|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".<ref>{{harvnb|Hodgkin|1992|pp=360–369}}</ref><ref>{{cite web|url=https://www.nobelprize.org/prizes/medicine/1963/hodgkin/facts/|title=Alan Hodgkin Facts|website=nobelprize.org|access-date=25 June 2020}}</ref> During the [[Nobel Banquet]] on 10 December 1963, Hodgkin gave the traditional speech on behalf of the three neurophysiologists, thanking the [[Gustaf VI Adolf|king]] and the [[Nobel Committee for Physiology or Medicine]] for the award.<ref>{{cite web|url=https://www.nobelprize.org/prizes/medicine/1963/hodgkin/speech/|title=Alan Hodgkin Banquet speech|website=nobelprize.org|access-date=25 June 2020}}</ref> 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.<ref>{{cite web|title=The Nobel Prize in Physiology or Medicine 1966|url=https://www.nobelprize.org/prizes/medicine/1966/rous/facts/|website=Nobelprize.org|publisher=Nobel Media AB|access-date=25 June 2020}}</ref>

==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 [[List of presidents of the Royal Society|president of the Royal Society]]. Together with [[Denis Baylor]] and Peter Detwiler he published a series of papers on turtle [[Photoreceptor cell|photoreceptors]].<ref>{{cite journal|doi=10.1113/jphysiol.1973.sp010340|title=Detection and resolution of visual stimuli by turtle photoreceptors|year=1973|last1=Baylor|first1=Denis A.|last2=Hodgkin|first2=A. L.|journal=The Journal of Physiology|volume=234|issue=1|pages=163–198|pmid=4766219|pmc=1350657}}</ref><ref>{{cite journal|doi=10.1113/jphysiol.1974.sp010732|title=Changes in time scale and sensitivity in turtle photoreceptors|year=1974|last1=Baylor|first1=Denis A.|last2=Hodgkin|first2=A. L.|journal=The Journal of Physiology|volume=242|issue=3|pages=729–758|pmid=4449053|pmc=1330660}}</ref><ref>{{cite journal|doi=10.1113/jphysiol.1974.sp010731|title=The electrical response of turtle cones to flashes and steps of light|year=1974|last1=Baylor|first1=Denis A.|last2=Hodgkin|first2=A. L.|journal=The Journal of Physiology|volume=242|issue=3|pages=685–727|pmid=4449052|pmc=1330659}}</ref><ref>{{cite journal|doi=10.1113/jphysiol.1974.sp010733|title=Reconstruction of the electrical responses of turtle cones to flashes and steps of light|year=1974|last1=Baylor|first1=Denis A.|last2=Hodgkin|first2=A. L.|last3=Lamb|first3=T. D.|journal=The Journal of Physiology|volume=242|issue=3|pages=759–791|pmid=4449054|pmc=1330661}}</ref><ref>{{cite journal | last1=Detwiler | first1=P. B. | last2=Hodgkin | first2=A. L. | last3=McNaughton | first3=P. A. | title=A surprising property of electrical spread in the network of rods in the turtle's retina | journal=Nature | publisher=Springer Science and Business Media LLC | volume=274 | issue=5671 | year=1978 | issn=0028-0836 | doi=10.1038/274562a0 | pages=562–565| pmid=672987 | bibcode=1978Natur.274..562D | s2cid=4257477 }}</ref><ref>{{cite journal|doi=10.1113/jphysiol.1979.sp012801|title=Electrical coupling between cones in the turtle retina.|year=1979|last1=Detwiler|first1=Peter B.|last2=Hodgkin|first2=A. L.|journal=The Journal of Physiology|volume=291|pages=75–100|pmid=225478|pmc=1280889}}</ref><ref>{{cite journal|doi=10.1113/jphysiol.1980.sp013159|title=Temporal and spatial characteristics of the voltage response of rods in the retina of the snapping turtle|year=1980|last1=Detwiler|first1=Peter B.|last2=Hodgkin|first3=A. L.|last3=McNaughton|first2=P. A.|journal=The Journal of Physiology|volume=300|pages=213–250|pmid=7381784|pmc=1279352}}</ref>

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.<ref>{{cite web|url=https://www.britannica.com/biography/Alan-Hodgkin|title=Sir Alan Hodgkin|website=britannica.com|access-date=25 June 2020}}</ref> 
From 1978 to 1985 he was the 34th [[List of Masters of Trinity College, Cambridge|Master of Trinity College, Cambridge]].<ref>{{Cite web |title=Sir Alan Hodgkin {{!}} Nobel Prize, Action Potentials & Neurophysiology {{!}} Britannica |url=https://www.britannica.com/biography/Alan-Hodgkin |access-date=2024-09-07 |website=www.britannica.com |language=en}}</ref>

He served on the Royal Society Council from 1958 to 1960 and on the [[Medical Research Council (United Kingdom)|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<ref>{{cite web|access-date=25 June 2020|title=History of Major Award Recipients|url=https://retinaresearchfnd.org/research-programs/awardhistory/|website=retinaresearchfnd.org}}</ref> 
* 1983 – Lord Crook Medal of the [[Worshipful Company of Spectacle Makers]]<ref>{{cite web|url=https://www.spectaclemakers.com/supporting-research#:~:text=The%20Company's%20Gold%20medal%20is,Company%20from%201963%20to%201965.|title=Support for Research|website=spectaclemakers.com|access-date=25 June 2020}}</ref>   
* 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]]<ref name=nasmember>{{cite web|url=http://www.nasonline.org/member-directory/deceased-members/46132.html|title=Alan Hodgkin|website=Nasonline.org|access-date=28 April 2019}}</ref>
* 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)<ref name=rsbm>{{cite journal|last=Huxley|first=Andrew|title=Sir Alan Lloyd Hodgkin, O.M., K.B.E. 5 February 1914 – 20 December 1998: Elected F.R.S. 1948|journal=Biographical Memoirs of Fellows of the Royal Society|year=2000|volume=46|pages=219–241|doi=10.1098/rsbm.1999.0081|doi-access=free}}</ref>
* 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]]<ref>{{cite web|title=Sir Alan Lloyd Hodgkin|url=https://www.leopoldina.org/en/members/list-of-members/list-of-members/member/Member/show/sir-alan-lloyd-hodgkin/|website=leopoldina.org|publisher=German National Academy of Sciences Leopoldina|access-date=25 June 2020}}</ref>
* 1963 – Nobel Prize for Physiology or Medicine together with Andrew Fielding Huxley and John Carew Eccles (for their research on [[synapse]]s)<ref>{{cite web|title=The Nobel Prize in Physiology or Medicine 1963|url=https://www.nobelprize.org/nobel_prizes/medicine/laureates/1963/|website=Nobelprize.org|publisher=Nobel Media AB|access-date=16 June 2014}}</ref>
* 1962 – Foreign Hon. Member, [[American Academy of Arts and Sciences]]
* 1958 – [[Royal Medal]] of [[The Royal Society]]
* 1958 – Hon. MD, [[Université catholique de Louvain|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)<ref name=rsbm/>

A portrait of Hodgkin by [[Michael Noakes]] hangs in Trinity College's collection.<ref>{{cite web|title=Trinity College, University of Cambridge|url=https://www.bbc.co.uk/arts/yourpaintings/paintings/search/located_at/trinity-college-cambridge-5846_locations|archive-url=https://archive.today/20140511164255/http://www.bbc.co.uk/arts/yourpaintings/paintings/search/located_at/trinity-college-cambridge-5846_locations|url-status=dead|archive-date=11 May 2014|publisher=BBC Your Paintings}}</ref>

===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<ref name="Hodgkin 1992 100">{{harvnb|Hodgkin|1992|p=100}}</ref> who was later awarded the 1966 Nobel Prize in Physiology or Medicine.<ref>{{cite web|title=The Nobel Prize in Physiology or Medicine 1966|url=https://www.nobelprize.org/prizes/medicine/1966/rous/facts/|website=Nobelprize.org|publisher=Nobel Media AB|access-date=23 November 2019}}</ref> When Rous invited him for dinner to his home, Hodgkin got to know Rous' daughter, [[Marni Hodgkin|Marni]], who was then a student at [[Swarthmore College]].<ref name="Hodgkin 1992 100"/> He proposed to her before going back to England in 1938, but she rejected him.<ref>{{harvnb|Hodgkin|1992|p=112}}</ref> When Hodgkin briefly returned to the US in 1944 (see [[#Wartime activities|Wartime activities]]), they reunited and got married on 31 March.<ref>{{harvnb|Hodgkin|1992|pp=235–239}}</ref>

Their first daughter, Sarah, was born in April 1945, shortly before the Hodgkins moved back to Cambridge.<ref>{{harvnb|Hodgkin|1992|p=254}}</ref> They had three more children: Deborah Hodgkin (born 2 May 1947),<ref>{{harvnb|Hodgkin|1992|p=310}}</ref> Jonathan Hodgkin (born 24 August 1949),<ref>{{harvnb|Hodgkin|1992|p=319}}</ref> and Rachel Hodgkin (born June 1951).<ref>{{harvnb|Hodgkin|1992|p=352}}</ref>
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.<ref>{{cite news|last1=Saxon|first1=Wolfgang|title=Sir Alan Lloyd Hodgkin, 84, A Nobelist in Nerve Research|url=https://www.nytimes.com/1998/12/22/world/sir-alan-lloyd-hodgkin-84-a-nobelist-in-nerve-research.html|work=The New York Times|date=22 December 1998}}</ref>

===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.<ref name=rsbm/> Hodgkin died in 1998 in [[Cambridge]].<ref>{{cite journal|last=Lamb|first=Trevor|title=Obituary: Alan Hodgkin (1914–98)|journal=Nature|year=1999|volume=397|issue=6715|pages=112|pmid=9923671|doi=10.1038/16362|bibcode = 1999Natur.397..112L |doi-access=free}}</ref>

==See also==
* [[List of presidents of the Royal Society]]
==References==
{{Reflist}}

== Bibliography ==
{{refbegin}}
* {{cite book|last=Hodgkin|first=Alan|year=1992|title=Chance & Design - Reminiscences of Science in Peace and War|publisher=Cambridge University Press|isbn=978-0-521-45603-6}}
{{refend}}

==External links==
*[http://www.trin.cam.ac.uk/index.php?pageid=172 The Master of Trinity] at [[Trinity College, Cambridge]]
*{{Nobelprize}} including the Nobel Lecture on 11 December 1963 ''The Ionic Basis of Nervous Conduction''
*{{NPG name}}
*[https://news.bbc.co.uk/2/hi/science/nature/239286.stm BBC obituary]
*[http://www.sfn.org/-/media/SfN/Documents/ClassicPapers/ActionPotentials/hodgkin5.ashx?la=en&hash=B7491DA5CF6B4F269B13D2AAAD37F77BFBDCE5D0 Action Potential Paper]
*[http://www.iwm.org.uk/collections/item/object/80012991 Imperial War Museum Interview]

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{{s-bef|before=[[Rab Butler|The Lord Butler of Saffron Walden]]}}
{{s-ttl|title=[[List of masters of Trinity College, Cambridge|Master of Trinity College, Cambridge]]|order=33rd|years=1978–1984}}
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{{s-bef|before=[[Edgar Adrian, 1st Baron Adrian|The Lord Adrian]]}}
{{s-ttl|title=[[Chancellor (education)|Chancellor]] of the [[University of Leicester]]|order=2nd|years=1971–1984}}
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{{s-bef|before=[[Patrick Blackett]]}}
{{s-ttl|title=[[Royal Society|President of the Royal Society]]|order=53rd|years=1970–1975}}
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{{Copley Medallists 1951-2000}}
{{Nobel Prize in Physiology or Medicine Laureates 1951-1975}}
{{1963 Nobel Prize winners}}
{{Royal Society presidents 1900s}}
{{Masters of Trinity College, Cambridge}}

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[[Category:1914 births]]
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[[Category:Fellows of the Indian National Science Academy]]
[[Category:Fellows of the Royal Society]]
[[Category:Foreign associates of the National Academy of Sciences]]
[[Category:Hodgkin family|Alan]]
[[Category:Knights Commander of the Order of the British Empire]]
[[Category:Masters of Trinity College, Cambridge]]
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[[Category:Nobel laureates in Physiology or Medicine]]
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[[Category:Presidents of the Royal Society]]
[[Category:Radar pioneers]]
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[[Category:Systems biologists]]
[[Category:English expatriates in Switzerland]]
[[Category:20th-century Quakers]]
[[Category:John Humphrey Plummer Professors]]
[[Category:Members of the Royal Swedish Academy of Sciences]]
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