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Alan Hodgkin

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Sir Alan Lloyd HodgkinOMKBEFRS[1](5 February 1914 – 20 December 1998) was an Englishphysiologistandbiophysicistwho shared the 1963Nobel Prize in Physiology or MedicinewithAndrew HuxleyandJohn Eccles.

Sir
Alan Lloyd Hodgkin
OMKBEFRS
Born(1914-02-05)5 February 1914
Banbury,Oxfordshire, England
Died20 December 1998(1998-12-20)(aged 84)
Cambridge,England
NationalityEnglish
CitizenshipBritish
Alma materUniversity of Cambridge
Known forHodgkin cycle
Hodgkin–Huxley model
Hodgkin–Huxley sodium channels
Goldman–Hodgkin–Katz flux equation
Goldman–Hodgkin–Katz voltage equation
SpouseMarion Rous
ChildrenSarah, Deborah,Jonathan Hodgkin,and Rachel
RelativesHodgkin family
Awards
Scientific career
FieldsPhysiology
Biophysics

Early life and education

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See also:Hodgkin family

Hodgkin was born inBanbury,Oxfordshire, on 5 February 1914. He was the oldest of three sons ofQuakersGeorge Hodgkin and Mary Wilson Hodgkin. His father was the son ofThomas Hodgkinand had read for theNatural Science TriposatCambridgewhere he had befriended electrophysiologistKeith 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 theSociety of Friends,George and Mary opposed theMilitary 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.[3]In 1916, George Hodgkin travelled to Armenia as part of an investigation of distress. Moved by the misery and suffering ofArmenian refugeeshe attempted to go back there in 1918 on a route through the Persian Gulf (as the northern route was closed because of theOctober Revolutionin Russia). He died of dysentery inBaghdadon 24 June 1918, just a few weeks after his youngest son, Keith, had been born.[4]

From an early life on, Hodgkin and his brothers were encouraged to explore the country around their home, which instilled in Alan an interest innatural history,particularlyornithology.At the age of 15, he helpedWilfred Backhouse Alexanderwith surveys ofheronriesand later, atGresham's School,he overlapped and spent a lot of time withDavid Lack.[5][6]In 1930, he was the winner of a bronze medal in the Public Schools Essay Competition organised by theRoyal Society for the Protection of Birds.[7]

School and university

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Alan started his education atThe Downs Schoolwhere his contemporaries included future scientistsFrederick Sanger,Alec Bangham,"neither outstandingly brilliant at school" according to Hodgkin,[8]as well as future artistsLawrence GowingandKenneth Rowntree.After the Downs School, he went on toGresham's Schoolwhere he overlapped with future composerBenjamin Brittenas well asMaury Meiklejohn.[9]He ended up receiving a scholarship atTrinity College, Cambridgein botany, zoology and chemistry.[10]

Between school and college, he spent May 1932 at theFreshwater Biological StationatWray Castlebased on a recommendation of his future Director of Studies atTrinity,Carl Pantin.[11]After Wray Castle, he spent two months with a German family inFrankfurtas "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 ofA. L. Smith,whose daughter Dorothy was also married to Alan's uncleRobert Howard Hodgkin.[14]

In the autumn of 1932, Hodgkin started as a freshman scholar at Trinity College where his friends includedclassicistsJohn RavenandMichael Grant,fellow-scientistsRichard SyngeandJohn H. Humphrey,[15]as well asPollyandDavid Hill,the children ofNobel laureateArchibald Hill.[5][16]He took physiology with chemistry and zoology for the first two years, including lectures byNobel laureateE.D. Adrian.[17]For Part II of thetriposhe decided to focus on physiology instead of zoology. Nevertheless, he participated in a zoological expedition to theAtlas Mountainsin Morocco led byJohn Pringlein 1934.[18]He finished Part II of the tripos in July 1935 and stayed atTrinityas 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 knewJames Klugmann[23]andJohn Cornford,[24]but he emphasised in his autobiography that none of his friends "made any serious effort to convert me [toCommunism], either then or later. "[25]From 1935 to 1937, Hodgkin was a member of theCambridge Apostles.[26]

Pre-war research

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Trinity College at theUniversity of Cambridgewhere Hodgkin was a student from 1932 to 1935, worked as a research fellow from 1936 on and finally served asMasterfrom 1978 to 1984

Hodgkin started conducting experiments on how electrical activity is transmitted in thesciatic nerveof frogs in July 1934.[27]He found that anerve impulsearriving at a cold or compression block, can decrease the electrical threshold beyond the block, suggesting that the impulse produces a spread of anelectrotonic potentialin the nerve beyond the block.[28]In 1936, Hodgkin was invited byHerbert Gasser,then director of theRockefeller Institutein New York City, to work in his laboratory during 1937–38. There he metRafael Lorente de Nó[29]andKenneth Stewart Colewith whom he ended up publishing a paper.[30]During that year he also spent time at theWoods Hole Marine Biological Laboratorywhere he was introduced to thesquid giant axon,[31]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 visitedJoseph ErlangeratWashington University in St. Louiswho 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.[32]Working with single nerve fibres fromshore crabsandsquids,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 withAndrew Huxleywho had enteredTrinityas a freshman in 1935, three years after Hodgkin.[34]With a £300 equipment grant from theRockefeller 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 thePlymouth Marine Laboratoryin July 1939.[35]There, he and Huxley managed to insert a finecannulainto the giant axon of squids and recordaction potentialsfrom inside the nerve fibre. They sent a short note of their success toNaturejust before the outbreak ofWorld War II.[36]

Wartime activities

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Despite hisQuakerupbringing, Hodgkin was eager to join the war effort as contact with the Nazis during his stay in Germany in 1932 had removed all hispacifistbeliefs. His first post was at theRoyal Aircraft Establishmentwhere he worked on issues in aviation medicine, such as oxygen supply for pilots at high altitudes and thedecompression sicknesscaused by nitrogen bubbles coming out of the blood.[37]In February 1940 he transferred to theTelecommunications Research Establishment(TRE) where he worked on the development of centimetricradar,including the design of thevillage InnAGLT airborne gun-laying system. He was a member ofE.G. Bowen'sgroup inSt Athanin South Wales and lived in a local guest house together with John Pringle andRobert Hanbury Brown.The group moved toSwanagein May 1940 where Pringle replaced Bowen as leader of the group.[38]In March 1941, Hodgkin flew on the test flight of aBristol Blenheimfitted with the first airborne centimetric radar system. In February and March 1944, he visited theMIT Radiation Laboratoryto 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 autobiographyChance and Design: Reminiscences of Science in Peace and War.[40]

1945–1963: Action potential theory and Nobel Prize

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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 (Cm). Voltage-gated and leak ion channels are represented by nonlinear (gn) and linear (gL) 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 (Ip).

As theAllied Forces'invasion of Franceand their continuedadvance towards Germanyin 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 withW. A. H. Rushtonand 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]

After being released from military service in August 1945 uponAdrian'srequest, Hodgkin was able to restart his experiments in collaboration withBernard Katzand his pre-war collaboratorAndrew Huxley.They spent the summers of 1947, 1948, and 1949 at thePlymouth Marine Laboratorywhere they continued to measure resting and action potentials from inside the giant axon of the squid.[42]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.[43] The data they had obtained in 1949 resulted in a series of five papers published inThe Journal of Physiologythat described what became later known as theHodgkin–Huxley modelof theaction potentialand eventually earned Hodgkin and Huxley theNobel Prize in Physiology or Medicine.[44][45][46][47][48]Building on work byKenneth S. Cole[49]they used a technique ofelectrophysiology,known as thevoltage clampto 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 theaction potentialis caused by changes in the selective permeability of the membrane for different ions, specificallysodium,potassium,andchloride.Amodelthat relies on a set ofdifferential equationsand 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 theHodgkin cycle.[52]

In addition, Hodgkin and Huxley's findings led them to hypothesize the existence ofion channelson cell membranes, which were confirmed only decades later. Confirmation of ion channels came with the development of thepatch clampleading to a Nobel prize in 1991 forErwin NeherandBert Sakmann,and in 2003 forRoderick 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 withRichard Keyneshe 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 scientistJens Christian Skoudiscovered the enzymeNa+/K+-ATPasethat usesATPto export three sodium ions in exchange for two potassium ions that are imported,[55]for which he received theNobel Prize in Chemistryin 1997.[56]

Hodgkin was nominated for the Nobel Prize in Physiology or Medicine in 1953 byLord Adrian.[57]In October 1961, he was told by Swedish journalists that he,Huxley,andEccleshad 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 toGeorg von Békésy.It was only two years later that Hodgkin,Huxley,andEccleswere 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 theNobel Banqueton 10 December 1963, Hodgkin gave the traditional speech on behalf of the three neurophysiologists, thanking thekingand theNobel Committee for Physiology or Medicinefor 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

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From 1951 to 1969, Hodgkin was the Foulerton Professor of theRoyal Societyat 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 aspresident of the Royal Society.Together withDenis Baylorand Peter Detwiler he published a series of papers on turtlephotoreceptors.[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 theOrder of Meritin 1973.[69] From 1978 to 1985 he was the 34thMaster of Trinity College, Cambridge.[70]

He served on the Royal Society Council from 1958 to 1960 and on theMedical Research Councilfrom 1959 to 1963. He was foreign secretary ofthe Physiological Societyfrom 1961 to 1967. He also held additional administrative posts such as Chancellor,University of Leicester,from 1971 to 1984

Awards and honours

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A portrait of Hodgkin byMichael Noakeshangs in Trinity College's collection.[76]

Publications

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  • The Conduction of the Nervous Impulse(1964)
  • Chance and Design: Reminiscences of Science in Peace and War(1992)

Personal life

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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 atSwarthmore 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 (seeWartime 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 atMacmillan Publishing Companyand a successful writer of children's literature, includingYoung Winter's TalesandDead Indeed.Jonathan Hodgkinbecame a molecular biologist at Cambridge University. Deborah Hodgkin is also a successful psychologist.

Thomas Hodgkin(1798–1866), who first describedHodgkin's lymphoma,was Alan Hodgkin's great-uncle.[85]

Death

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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 thespinal cordfrom one of theintervertebral discsin his neck, which left him unable to walk without support, and with progressive disablement.[1]Hodgkin died in 1998 inCambridge.[86]

See also

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References

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  1. ^abcdHuxley, Andrew (2000)."Sir Alan Lloyd Hodgkin, O.M., K.B.E. 5 February 1914 – 20 December 1998: Elected F.R.S. 1948".Biographical Memoirs of Fellows of the Royal Society.46:219–241.doi:10.1098/rsbm.1999.0081.
  2. ^Hodgkin 1992,p. 5
  3. ^Hodgkin 1992,pp. 7–9
  4. ^Hodgkin 1992,p. 10
  5. ^abHodgkin 1992,pp. 11–12
  6. ^Hodgkin, Alan (1983). "Beginning: Some Reminiscences of my early life (1914–1947)".Annual Review of Physiology.45:1–16.doi:10.1146/annurev.ph.45.030183.000245.PMID6342510.
  7. ^Protection of Birds Measures Urged By Royal SocietyinThe Times,Saturday, 29 March 1930; pg. 14; Issue 45474; col C
  8. ^Hodgkin 1992,p. 25
  9. ^Hodgkin 1992,pp. 27–29
  10. ^Hodgkin 1992,p. 30
  11. ^Hodgkin 1992,p. 31
  12. ^Hodgkin 1992,p. 34
  13. ^"SMITH, Arthur Lionel Forster"(PDF).Retrieved3 February2023.
  14. ^Hodgkin 1992,p. 40
  15. ^Hodgkin 1992,p. 49
  16. ^Hodgkin, Alan (1983). "Beginning: Some Reminiscences of my early life (1914–1947)".Annual Review of Physiology.45:1–16.doi:10.1146/annurev.ph.45.030183.000245.PMID6342510.
  17. ^Hodgkin 1992,p. 50
  18. ^Hodgkin 1992,p. 55
  19. ^Hodgkin 1992,p. 66
  20. ^Hodgkin 1992,p. 38
  21. ^Hodgkin 1992,p. 48
  22. ^Hodgkin 1992,p. 53
  23. ^Hodgkin 1992,p. 79
  24. ^Hodgkin 1992,pp. 83–87
  25. ^Hodgkin 1992,p. 80
  26. ^Hodgkin 1992,p. 87
  27. ^Hodgkin 1992,p. 63
  28. ^Hodgkin, Alan (1937)."Evidence for Electrical Transmission in Nerve. Part I".The Journal of Physiology.90(2): 183–210.doi:10.1113/jphysiol.1937.sp003507.PMC1395060.PMID16994885.
  29. ^Hodgkin 1992,p. 92
  30. ^Cole, Kenneth; Hodgkin, Alan (1939)."Membrane and Protoplasm Resistance in the Squid Giant Axon".The Journal of General Physiology.22(5): 671–687.doi:10.1085/jgp.22.5.671.PMC2142005.PMID19873126.
  31. ^Hodgkin, Alan (1983). "Beginning: Some Reminiscences of my early life (1914–1947)".Annual Review of Physiology.45:1–16.doi:10.1146/annurev.ph.45.030183.000245.PMID6342510.
  32. ^Hodgkin 1992,p. 113
  33. ^Hodgkin, Alan (1939)."The Relation between Conduction Velocity and the Electrical Resistance outside a Nerve Fibre".The Journal of Physiology.94(4): 560–570.doi:10.1113/jphysiol.1939.sp003702.PMC1393884.PMID16995066.
  34. ^Huxley, Andrew(2004)."Andrew Huxley".In Squire, Larry R. (ed.).The history of neuroscience in autobiography.Washington DC: Society for Neuroscience. pp. 282–318.ISBN0-12-660246-8.
  35. ^Hodgkin, Alan (1976)."Chance and Design in Electrophysiology: An Informal Account of Certain Experiments on Nerve Carried out between 1934 and 1952".Journal of Physiology.263(1): 1–21.doi:10.1113/jphysiol.1976.sp011620.PMC1307686.PMID796420.
  36. ^Hodgkin, A. L.; Huxley, A. F. (1939). "Action Potentials Recorded from Inside a Nerve Fibre".Nature.144(3651): 710–711.Bibcode:1939Natur.144..710H.doi:10.1038/144710a0.S2CID4104520.
  37. ^Hodgkin 1992,p. 140
  38. ^Hodgkin 1992,pp. 141–156
  39. ^Hodgkin 1992,pp. 233–239
  40. ^Hodgkin 1992,pp. ix–xi
  41. ^Hodgkin, A. L.; Rushton, W.A. H. (1946)."The electrical constants of a crustacean nerve fibre".Proceedings of the Royal Society of London. Series B.133(873): 444–479.Bibcode:1946RSPSB.133..444H.doi:10.1098/rspb.1946.0024.PMID20281590.
  42. ^Hodgkin 1992,pp. 271–290
  43. ^Hodgkin, A. L.; Katz, B. (1949)."The Effect of Sodium Ions on the Electrical Activity of the Giant Axon of the Squid".Journal of Physiology.108(1): 37–77.doi:10.1113/jphysiol.1949.sp004310.PMC1392331.PMID18128147.
  44. ^Hodgkin, A. L.; Huxley, A. F.; Katz, B. (1952)."Measurement of Current-Voltage Relations in the Membrane of the Gian Axon of Loligo".The Journal of Physiology.116(4): 424–448.doi:10.1113/jphysiol.1952.sp004716.PMC1392219.PMID14946712.
  45. ^Hodgkin, A. L.; Huxley, A. F. (1952)."Currents carried by sodium and potassium ions through the membrane of the giant axon of Loligo".The Journal of Physiology.116(4): 449–472.doi:10.1113/jphysiol.1952.sp004717.PMC1392213.PMID14946713.
  46. ^Hodgkin, A. L.; Huxley, A. F. (1952)."The components of membrane conductance in the giant axon of Loligo".The Journal of Physiology.116(4): 473–496.doi:10.1113/jphysiol.1952.sp004718.PMC1392209.PMID14946714.
  47. ^Hodgkin, A. L.; Huxley, A. F. (1952)."The dual effect of membrane potential on sodium conductance in the giant axon of loligo".The Journal of Physiology.116(4): 497–506.doi:10.1113/jphysiol.1952.sp004719.PMC1392212.PMID14946715.
  48. ^abHodgkin, A. L.; Huxley, A. F. (1952)."A Quantitative Description of Membrane Current and its Application to Conduction and Excitation in Nerve".The Journal of Physiology.117(4): 500–544.doi:10.1113/jphysiol.1952.sp004764.PMC1392413.PMID12991237.
  49. ^Cole, Kenneth S. (1949). "Dynamic electrical characteristics of the squid axon membrane".Archives des sciences physiologique.3(2): 253–258.
  50. ^Hodgkin 1992,pp. 299–303
  51. ^Hodgkin, Alan (1951). "The ionic basis of electrical activity in nerve and muscle".Biological Reviews.26(4): 339–409.doi:10.1111/j.1469-185X.1951.tb01204.x.S2CID86282580.
  52. ^Noble, D. (2010)."Biophysics and systems biology".Philosophical Transactions of the Royal Society A: Mathematical, Physical and Engineering Sciences.368(1914): 1125–1139.Bibcode:2010RSPTA.368.1125N.doi:10.1098/rsta.2009.0245.PMC3263808.PMID20123750.
  53. ^Schwiening, C. J. (2012)."A brief historical perspective: Hodgkin and Huxley".The Journal of Physiology.590(11): 2571–2575.doi:10.1113/jphysiol.2012.230458.PMC3424716.PMID22787170.
  54. ^Hodgkin, A. L.; Keynes, R. D. (1955)."Active transport of cations in giant axons from Sepia and Loligo".The Journal of Physiology.128(1): 28–60.doi:10.1113/jphysiol.1955.sp005290.PMC1365754.PMID14368574.
  55. ^Skou, J. C. (1957). "The influence of some cations on an adenosine triphosphatase from peripheral nerves".Biochimica et Biophysica Acta.23(2): 394–401.doi:10.1016/0006-3002(57)90343-8.PMID13412736.S2CID32516710.
  56. ^"The Nobel Prize in Chemistry 1997".Nobelprize.org.Nobel Media AB.Retrieved12 July2020.
  57. ^"Nomination Archive".nobelprize.org.April 2020.Retrieved25 June2020.
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  59. ^"Alan Hodgkin Facts".nobelprize.org.Retrieved25 June2020.
  60. ^"Alan Hodgkin Banquet speech".nobelprize.org.Retrieved25 June2020.
  61. ^"The Nobel Prize in Physiology or Medicine 1966".Nobelprize.org.Nobel Media AB.Retrieved25 June2020.
  62. ^Baylor, Denis A.; Hodgkin, A. L. (1973)."Detection and resolution of visual stimuli by turtle photoreceptors".The Journal of Physiology.234(1): 163–198.doi:10.1113/jphysiol.1973.sp010340.PMC1350657.PMID4766219.
  63. ^Baylor, Denis A.; Hodgkin, A. L. (1974)."Changes in time scale and sensitivity in turtle photoreceptors".The Journal of Physiology.242(3): 729–758.doi:10.1113/jphysiol.1974.sp010732.PMC1330660.PMID4449053.
  64. ^Baylor, Denis A.; Hodgkin, A. L. (1974)."The electrical response of turtle cones to flashes and steps of light".The Journal of Physiology.242(3): 685–727.doi:10.1113/jphysiol.1974.sp010731.PMC1330659.PMID4449052.
  65. ^Baylor, Denis A.; Hodgkin, A. L.; Lamb, T. D. (1974)."Reconstruction of the electrical responses of turtle cones to flashes and steps of light".The Journal of Physiology.242(3): 759–791.doi:10.1113/jphysiol.1974.sp010733.PMC1330661.PMID4449054.
  66. ^Detwiler, P. B.; Hodgkin, A. L.; McNaughton, P. A. (1978). "A surprising property of electrical spread in the network of rods in the turtle's retina".Nature.274(5671). Springer Science and Business Media LLC: 562–565.Bibcode:1978Natur.274..562D.doi:10.1038/274562a0.ISSN0028-0836.PMID672987.S2CID4257477.
  67. ^Detwiler, Peter B.; Hodgkin, A. L. (1979)."Electrical coupling between cones in the turtle retina".The Journal of Physiology.291:75–100.doi:10.1113/jphysiol.1979.sp012801.PMC1280889.PMID225478.
  68. ^Detwiler, Peter B.; Hodgkin, P. A.; McNaughton, A. L. (1980)."Temporal and spatial characteristics of the voltage response of rods in the retina of the snapping turtle".The Journal of Physiology.300:213–250.doi:10.1113/jphysiol.1980.sp013159.PMC1279352.PMID7381784.
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  76. ^"Trinity College, University of Cambridge".BBC Your Paintings. Archived fromthe originalon 11 May 2014.
  77. ^abHodgkin 1992,p. 100
  78. ^"The Nobel Prize in Physiology or Medicine 1966".Nobelprize.org.Nobel Media AB.Retrieved23 November2019.
  79. ^Hodgkin 1992,p. 112
  80. ^Hodgkin 1992,pp. 235–239
  81. ^Hodgkin 1992,p. 254
  82. ^Hodgkin 1992,p. 310
  83. ^Hodgkin 1992,p. 319
  84. ^Hodgkin 1992,p. 352
  85. ^Saxon, Wolfgang (22 December 1998)."Sir Alan Lloyd Hodgkin, 84, A Nobelist in Nerve Research".The New York Times.
  86. ^Lamb, Trevor (1999)."Obituary: Alan Hodgkin (1914–98)".Nature.397(6715): 112.Bibcode:1999Natur.397..112L.doi:10.1038/16362.PMID9923671.

Bibliography

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  • Hodgkin, Alan (1992).Chance & Design - Reminiscences of Science in Peace and War.Cambridge University Press.ISBN978-0-521-45603-6.
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Academic offices
Preceded by 33rdMaster of Trinity College, Cambridge
1978–1984 		Succeeded by
Preceded by 2ndChancellorof theUniversity of Leicester
1971–1984 		Succeeded by
Professional and academic associations
Preceded by 53rdPresident of the Royal Society
1970–1975 		Succeeded by
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