Sir Alan Lloyd Hodgkin
(5 February 1914 – 20 December 1998) was an English
physiologist and
biophysicist
Biophysics is an interdisciplinary science that applies approaches and methods traditionally used in physics to study biological phenomena. Biophysics covers all scales of biological organization, from molecular to organismic and populations. ...
who shared the 1963
Nobel Prize in Physiology or Medicine
The Nobel Prize in Physiology or Medicine is awarded yearly by the Nobel Assembly at the Karolinska Institute for outstanding discoveries in physiology or medicine. The Nobel Prize is not a single prize, but five separate prizes that, accord ...
with
Andrew Huxley
Sir Andrew Fielding Huxley (22 November 191730 May 2012) was an English physiologist and biophysicist. He was born into the prominent Huxley family. After leaving Westminster School in central London, he went to Trinity College, Cambridge ...
and
John Eccles.
Early life and education
Hodgkin was born in
Banbury
Banbury is a historic market town on the River Cherwell in Oxfordshire, South East England. It had a population of 54,335 at the 2021 Census.
Banbury is a significant commercial and retail centre for the surrounding area of north Oxfordshire ...
, Oxfordshire, on 5 February 1914. He was the oldest of three sons of
Quakers
Quakers are people who belong to a historically Protestant Christian set of denominations known formally as the Religious Society of Friends. Members of these movements ("theFriends") are generally united by a belief in each human's abil ...
George Hodgkin and Mary Wilson Hodgkin. His father was the son of
Thomas Hodgkin
Thomas Hodgkin RMS (17 August 1798 – 5 April 1866) was a British physician, considered one of the most prominent pathologists of his time and a pioneer in preventive medicine. He is now best known for the first account of Hodgkin's disease, ...
and had read for the
Natural Science Tripos
The Natural Sciences Tripos (NST) is the framework within which most of the science at the University of Cambridge is taught. The tripos includes a wide range of Natural Sciences from physics, astronomy, and geoscience, to chemistry and biology, w ...
at
Cambridge
Cambridge ( ) is a university city and the county town in Cambridgeshire, England. It is located on the River Cam approximately north of London. As of the 2021 United Kingdom census, the population of Cambridge was 145,700. Cambridge bec ...
where he had befriended electrophysiologist
Keith Lucas. 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, George and Mary opposed the
Military Service Act of 1916 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. In 1916 George Hodgkin travelled to Armenia as part of an investigation of distress. Moved by the misery and suffering of
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
The October Revolution,. officially known as the Great October Socialist Revolution. in the Soviet Union, also known as the Bolshevik Revolution, was a revolution in Russia led by the Bolshevik Party of Vladimir Lenin that was a key moment ...
in Russia). He died of dysentery in
Baghdad
Baghdad (; ar, بَغْدَاد , ) is the capital of Iraq and the second-largest city in the Arab world after Cairo. It is located on the Tigris near the ruins of the ancient city of Babylon and the Sassanid Persian capital of Ctesiphon ...
on 24 June 1918, just a few weeks after his youngest son, Keith, had been born.
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, particularly
ornithology
Ornithology is a branch of zoology that concerns the "methodological study and consequent knowledge of birds with all that relates to them." Several aspects of ornithology differ from related disciplines, due partly to the high visibility and th ...
. At the age of 15, he helped
Wilfred Backhouse Alexander
Wilfrid Backhouse Alexander (4 February 1885 – 18 December 1965) was an England, English ornithologist and entomologist. He was a brother of Horace Alexander and Christopher James Alexander.
Alexander was born at Croydon in Surrey, England in ...
with surveys of
heronries
A heronry, sometimes called a heron rookery, is a breeding ground for herons.
Notable heronries
Although their breeding territories are often on more protected small islands in lakes or retention ponds, herons breed in heronries (or also calle ...
and later, at
Gresham's School
Gresham's School is a Public school (United Kingdom), public school (English Independent school (United Kingdom), independent Day school, day and boarding school) in Holt, Norfolk, Holt, Norfolk, England, one of the top thirty International Bac ...
, he overlapped and spent a lot of time with
David Lack
David Lambert Lack FRS (16 July 1910 – 12 March 1973) was a British evolutionary biologist who made contributions to ornithology, ecology, and ethology. His 1947 book, ''Darwin's Finches'', on the finches of the Galapagos Islands was a landm ...
.
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
The Royal Society for the Protection of Birds (RSPB) is a charitable organisation registered in England and Wales and in Scotland. It was founded in 1889. It works to promote conservation and protection of birds and the wider environment thr ...
.
Alan started his education at
The Downs School where his contemporaries included future scientists
Frederick Sanger
Frederick Sanger (; 13 August 1918 – 19 November 2013) was an English biochemist who received the Nobel Prize in Chemistry twice.
He won the 1958 Chemistry Prize for determining the amino acid sequence of insulin and numerous other p ...
,
Alec Bangham
Alec Douglas Bangham FRS (10 November 1921 Manchester – 9 March 2010 Great Shelford) was a British biophysicist who first studied blood clotting mechanisms but became well known for his research on liposomes and his invention of clinically u ...
, "neither outstandingly brilliant at school" according to Hodgkin, as well as future artists
Lawrence Gowing
Sir Lawrence Burnett Gowing (21 April 1918 – 5 February 1991) was an English artist, writer, curator and teacher. Initially recognised as a portrait and landscape painter, he quickly rose to prominence as an art educator, writer, and eventuall ...
and
Kenneth Rowntree
Kenneth Rowntree (14 March 1915 – 21 February 1997) was a British artist.
Career and life
Kenneth Rowntree was born in Scarborough, the son of Howard Doncaster Rowntree (1879-1974).
He was educated at Bootham School, York.
He studied at th ...
. After the Downs School, he went on to
Gresham's School
Gresham's School is a Public school (United Kingdom), public school (English Independent school (United Kingdom), independent Day school, day and boarding school) in Holt, Norfolk, Holt, Norfolk, England, one of the top thirty International Bac ...
where he overlapped with future composer
Benjamin Britten
Edward Benjamin Britten, Baron Britten (22 November 1913 – 4 December 1976, aged 63) was an English composer, conductor, and pianist. He was a central figure of 20th-century British music, with a range of works including opera, other ...
as well as
Maury Meiklejohn. He ended up receiving a scholarship at
Trinity College, Cambridge
Trinity College is a constituent college of the University of Cambridge. Founded in 1546 by Henry VIII, King Henry VIII, Trinity is one of the largest Cambridge colleges, with the largest financial endowment of any college at either Cambridge ...
in Botany, Zoology and Chemistry.
Between school and college, he spent May 1932 at the
Freshwater Biological Station at
Wray Castle
Wray Castle is a Victorian neo-gothic building at Claife in the English county of Cumbria.
The house and grounds have belonged to the National Trust since 1929, and house has opened to the public on a regular basis since 2013. The grounds, whi ...
based on a recommendation of his future Director of Studies at
Trinity
The Christian doctrine of the Trinity (, from 'threefold') is the central dogma concerning the nature of God in most Christian churches, which defines one God existing in three coequal, coeternal, consubstantial divine persons: God the F ...
,
Carl Pantin. After Wray Castle, he spent two months with a German family in
Frankfurt
Frankfurt, officially Frankfurt am Main (; Hessian: , "Frank ford on the Main"), is the most populous city in the German state of Hesse. Its 791,000 inhabitants as of 2022 make it the fifth-most populous city in Germany. Located on its na ...
as "in those days it was thought highly desirable that anyone intending to read science should have a reasonable knowledge of German."
After his return to England in early August 1932, his mother Mary was remarried to Lionel Smith (1880–1972), the eldest son of
A. L. Smith
Sir Archibald Levin Smith (26 August 1836 – 20 October 1901) was a British judge and a rowing (sport), rower who competed at Henley and in the Oxford and Cambridge Boat Race.
Biography
Smith was the son of Francis Smith, J.P. of Salt ...
, whose daughter Dorothy was also married to Alan's uncle
Robert Howard Hodgkin.
In autumn of 1932 Hodgkin started as a freshman scholar at
Trinity College, Cambridge
Trinity College is a constituent college of the University of Cambridge. Founded in 1546 by Henry VIII, King Henry VIII, Trinity is one of the largest Cambridge colleges, with the largest financial endowment of any college at either Cambridge ...
where his friends included classicists
John Raven
John Earle Raven (13 December 1914 – 5 March 1980) was an English classical scholar, notable for his work on presocratic philosophy, and amateur botanist.
Early life and education
John Raven was born on 13 December 1914 in Cambridge, the ...
and
Michael Grant, fellow-scientists
Richard Synge
Richard Laurence Millington Synge FRS FRSE FRIC FRSC MRIA (Liverpool, 28 October 1914 – Norwich, 18 August 1994) was a British biochemist, and shared the 1952 Nobel Prize in Chemistry for the invention of partition chromatography with Arch ...
and
John H. Humphrey
John Herbert Humphrey Order of the British Empire#Commander, CBE Fellow of the Royal Society, FRS Fellow of the Royal College of Physicians, FRCP (16 December 1915 – 25 December 1987) was a UK, British bacteriologist and immunologist.
Education ...
, as well as
Polly
Polly is a given name, most often feminine, which originated as a variant of Molly (name), Molly (a diminutive of Mary (name), Mary). Polly may also be a short form of names such as Polina (given name), Polina, Polona (given name), Polona, Paula (g ...
and
David Hill, the children of
Nobel Laureate Archibald Hill
Archibald Vivian Hill (26 September 1886 – 3 June 1977), known as A. V. Hill, was a British physiologist, one of the founders of the diverse disciplines of biophysics and operations research. He shared the 1922 Nobel Prize in Physiology or ...
.
He took Physiology with Chemistry and Zoology for the first two years, including lectures by
Nobel Laureate E.D. Adrian. For Part II of the
tripos
At the University of Cambridge, a Tripos (, plural 'Triposes') is any of the examinations that qualify an undergraduate for a bachelor's degree or the courses taken by a student to prepare for these. For example, an undergraduate studying mathe ...
he decided to focus on physiology instead of zoology. Nevertheless, he participated in a zoological expedition to the
Atlas Mountains
The Atlas Mountains are a mountain range in the Maghreb in North Africa. It separates the Sahara Desert from the Mediterranean Sea and the Atlantic Ocean; the name "Atlantic" is derived from the mountain range. It stretches around through Moroc ...
in Morocco led by
John Pringle John Pringle may refer to:
*John Pringle, Lord Haining (c. 1674–1754), Scottish landowner, judge and politician, shire commissioner for Selkirk 1702–07, MP for Selkirkshire 1708–29, Lord of Session
*Sir John Pringle, 1st Baronet (1707–1782) ...
in 1934. He finished Part II of the tripos in July 1935 and stayed at
Trinity
The Christian doctrine of the Trinity (, from 'threefold') is the central dogma concerning the nature of God in most Christian churches, which defines one God existing in three coequal, coeternal, consubstantial divine persons: God the F ...
as a research fellow.
During his studies, Hodgkin, who described himself as "having been brought up as a supporter of the British Labour Party" was friends with communists and actively participated in distribution of Anti-War pamphlets. At Cambridge, he knew
James Klugmann
Norman John Klugmann (27 February 1912 – 14 September 1977), generally known as James Klugmann, was a leading British Communist writer and WW2 Soviet Spy, who became the official historian of the Communist Party of Great Britain.
Background ...
and
John Cornford
Rupert John Cornford (27 December 1915 – 28 December 1936) was an English poet and communist. During the first year of the Spanish Civil War, he was a member of the POUM militia and later the International Brigades. He died while fighting a ...
, but he emphasised in his autobiography that none of his friends "made any serious effort to convert me
o Communism either then or later." From 1935 to 1937, Hodgkin was a member of the
Cambridge Apostles
The Cambridge Apostles (also known as ''Conversazione Society'') is an intellectual society at the University of Cambridge founded in 1820 by George Tomlinson, a Cambridge student who became the first Bishop of Gibraltar.W. C. Lubenow, ''The Ca ...
.
Pre-war research
![Trinity_College_-_Great_Court_02](https://upload.wikimedia.org/wikipedia/commons/9/94/Trinity_College_-_Great_Court_02.jpg)
Hodgkin started conducting experiments on how electrical activity is transmitted in the
sciatic nerve of frogs in July 1934. He found that a
nerve impulse
An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, ...
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 physiology, electrotonus refers to the passive spread of charge inside a neuron and between cardiac muscle cells or smooth muscle cells. ''Passive'' means that voltage-dependent changes in membrane conductance do not contribute. Neurons and oth ...
in the nerve beyond the block.
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 Rafael Lorente de Nó (April 8, 1902 – April 2, 1990) was a Spanish neuroscientist who advanced the scientific understanding of the nervous system with his seminal research.
He was a member of the National Academy of Sciences.
The National Acad ...
and
Kenneth Stewart Cole with whom he ended up publishing a paper. During that year he also spent time at the
Woods Hole Marine Biological Laboratory where he was introduced to the squid giant axon, which ended up being the model system with which he conducted most of the research that eventually led to his Nobel Prize.
In spring 1938 he visited
Joseph Erlanger
Joseph Erlanger (January 5, 1874 – December 5, 1965) was an American physiologist who is best known for his contributions to the field of neuroscience. Together with Herbert Spencer Gasser, he identified several varieties of nerve fiber and es ...
at
Washington University in St. Louis
Washington University in St. Louis (WashU or WUSTL) is a private research university with its main campus in St. Louis County, and Clayton, Missouri. Founded in 1853, the university is named after George Washington. Washington University is r ...
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. Working with single nerve fibres from
shore crabs and
squids
True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting ...
, he showed that the conduction rate was much faster in sea water than in oil, providing strong evidence for the local circuit theory.
After his return to Cambridge he started collaborating with
Andrew Huxley
Sir Andrew Fielding Huxley (22 November 191730 May 2012) was an English physiologist and biophysicist. He was born into the prominent Huxley family. After leaving Westminster School in central London, he went to Trinity College, Cambridge ...
who had entered
Trinity
The Christian doctrine of the Trinity (, from 'threefold') is the central dogma concerning the nature of God in most Christian churches, which defines one God existing in three coequal, coeternal, consubstantial divine persons: God the F ...
as a freshman in 1935, three years after Hodgkin. With a £300 equipment grant from the
Rockefeller Foundation
The Rockefeller Foundation is an American private foundation and philanthropic medical research and arts funding organization based at 420 Fifth Avenue, New York City. The second-oldest major philanthropic institution in America, after the Carneg ...
, 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
Plymouth Marine Laboratory (abbreviated as PML) in the city of Plymouth, England, is a marine research organization and registered charity. It is a partner of the UK Research & Innovation's Natural Environment Research Council (NERC). PML's chai ...
in July 1939. There, he and Huxley managed to insert a fine cannula into the giant axon of squids and record
action potentials
An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells ...
from inside the nerve fibre. They sent a short note of their success to
''Nature'' just before the outbreak of
World War II
World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the vast majority of the world's countries—including all of the great powers—forming two opposin ...
.
Wartime activities
Despite his
Quaker
Quakers are people who belong to a historically Protestant Christian set of Christian denomination, denominations known formally as the Religious Society of Friends. Members of these movements ("theFriends") are generally united by a belie ...
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
Pacifism is the opposition or resistance to war, militarism (including conscription and mandatory military service) or violence. Pacifists generally reject theories of Just War. The word ''pacifism'' was coined by the French peace campaign ...
beliefs. His first post was at the
Royal Aircraft Establishment
The Royal Aircraft Establishment (RAE) was a British research establishment, known by several different names during its history, that eventually came under the aegis of the Ministry of Defence (United Kingdom), UK Ministry of Defence (MoD), bef ...
where he worked on issues in Aviation Medicine, such as oxygen supply for pilots at high altitude and the
decompression sickness
Decompression sickness (abbreviated DCS; also called divers' disease, the bends, aerobullosis, and caisson disease) is a medical condition caused by dissolved gases emerging from solution as bubbles inside the body tissues during decompressio ...
caused by nitrogen bubbles coming out of the blood. In February 1940 he transferred to the
Telecommunications Research Establishment
The Telecommunications Research Establishment (TRE) was the main United Kingdom research and development organization for radio navigation, radar, infra-red detection for heat seeking missiles, and related work for the Royal Air Force (RAF) ...
(TRE) where he worked on the development of centimetric
radar
Radar is a detection system that uses radio waves to determine the distance (''ranging''), angle, and radial velocity of objects relative to the site. It can be used to detect aircraft, ships, spacecraft, guided missiles, motor vehicles, w ...
, including the design of the
Village Inn
Village Inn is a casual-dining restaurant chain in the United States. Its restaurants are known for their breakfast menu items. Also, they feature a variety of salads, sandwiches, burgers, melts, and dinner items. Their pies have won numerous a ...
AGLT airborne gun-laying system. He was a member of
E.G. Bowen's group in
St Athan
St Athan ( cy, Sain Tathan) is a village and community (Wales), community in the Vale of Glamorgan in South Wales. The village and its parish church are dedicated to Saint Tathan. The church dates to the 13th–14th century, though an earlier ch ...
in South Wales and lived in a local guest house together with
John Pringle John Pringle may refer to:
*John Pringle, Lord Haining (c. 1674–1754), Scottish landowner, judge and politician, shire commissioner for Selkirk 1702–07, MP for Selkirkshire 1708–29, Lord of Session
*Sir John Pringle, 1st Baronet (1707–1782) ...
and
Robert Hanbury Brown
Robert Hanbury Brown, AC FRS (31 August 1916 – 16 January 2002) was a British astronomer and physicist born in Aruvankadu, India. He made notable contributions to the development of radar and later conducted pioneering work in the field of ...
. The group moved to
Swanage
Swanage () is a coastal town and civil parish in the south east of Dorset, England. It is at the eastern end of the Isle of Purbeck and one of its two towns, approximately south of Poole and east of Dorchester. In the 2011 census the civil ...
in May 1940 where Pringle replaced Bowen as leader of the group. In March 1941, Hodgkin flew on the test flight of a
Bristol Blenheim
The Bristol Blenheim is a British light bomber aircraft designed and built by the Bristol Aeroplane Company (Bristol) which was used extensively in the first two years of the Second World War, with examples still being used as trainers until ...
fitted with the first airborne centimetric radar system. In February and March 1944, Hodgkin visited the
MIT Radiation Laboratory
The Radiation Laboratory, commonly called the Rad Lab, was a microwave and radar research laboratory located at the Massachusetts Institute of Technology (MIT) in Cambridge, Massachusetts. It was first created in October 1940 and operated until 31 ...
to help foster the interchange of information on developments in radar between Britain and America.
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''.
1945–1963: Action potential theory and Nobel Prize
![Hodgkin-Huxley](https://upload.wikimedia.org/wikipedia/commons/9/98/Hodgkin-Huxley.svg)
As the
Allied Forces' invasion of France
France has been invaded on numerous occasions, by foreign powers or rival French governments; there have also been unimplemented invasion plans.
* the 1746 War of the Austrian Succession, Austria-Italian forces supported by the British navy attemp ...
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
Cambridge ( ) is a university city and the county town in Cambridgeshire, England. It is located on the River Cam approximately north of London. As of the 2021 United Kingdom census, the population of Cambridge was 145,700. Cambridge bec ...
. 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.
After being released from military service in August 1945 upon
Adrian's request, Hodgkin was able to restart his experiments in collaboration with
Bernard Katz and his pre-war collaborator
Andrew Huxley
Sir Andrew Fielding Huxley (22 November 191730 May 2012) was an English physiologist and biophysicist. He was born into the prominent Huxley family. After leaving Westminster School in central London, he went to Trinity College, Cambridge ...
. They spent the summers of 1947, 1948, and 1949 at the
Plymouth Marine Laboratory
Plymouth Marine Laboratory (abbreviated as PML) in the city of Plymouth, England, is a marine research organization and registered charity. It is a partner of the UK Research & Innovation's Natural Environment Research Council (NERC). PML's chai ...
where they continued to measure resting and action potentials from inside the giant axon of the squid. 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.
The data they had obtained in 1949 resulted in a series of five papers published in ''
The Journal of Physiology
''The Journal of Physiology'' is a biweekly peer-reviewed scientific journal that was established in 1878 and is published by Wiley-Blackwell on behalf of The Physiological Society. It covers research on all aspects of physiology, with an emphasis ...
'' that described what became later known as the
Hodgkin–Huxley model
The Hodgkin–Huxley model, or conductance-based model, is a mathematical model that describes how action potentials in neurons are initiated and propagated. It is a set of nonlinear differential equations that approximates the electrical charact ...
of the
action potential
An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, ...
and eventually earned Hodgkin and Huxley the
Nobel Prize in Physiology or Medicine
The Nobel Prize in Physiology or Medicine is awarded yearly by the Nobel Assembly at the Karolinska Institute for outstanding discoveries in physiology or medicine. The Nobel Prize is not a single prize, but five separate prizes that, accord ...
.
Building on work by
Kenneth S. Cole
Kenneth Stewart Cole (July 10, 1900 – April 18, 1984) was an American biophysicist described by his peers as "a pioneer in the application of physical science to biology". Cole was awarded the National Medal of Science in 1967.
Biography
He wa ...
they used a technique of
, known as the ''
voltage clamp
The voltage clamp is an experimental method used by electrophysiologists to measure the ion currents through the membranes of excitable cells, such as neurons, while holding the membrane voltage at a set level. A basic voltage clamp will itera ...
'' 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
An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, ...
is caused by changes in the selective permeability of the membrane for different ions, specifically
sodium
Sodium is a chemical element with the symbol Na (from Latin ''natrium'') and atomic number 11. It is a soft, silvery-white, highly reactive metal. Sodium is an alkali metal, being in group 1 of the periodic table. Its only stable iso ...
,
potassium
Potassium is the chemical element with the symbol K (from Neo-Latin ''kalium'') and atomic number19. Potassium is a silvery-white metal that is soft enough to be cut with a knife with little force. Potassium metal reacts rapidly with atmosphe ...
, and
chloride
The chloride ion is the anion (negatively charged ion) Cl−. It is formed when the element chlorine (a halogen) gains an electron or when a compound such as hydrogen chloride is dissolved in water or other polar solvents. Chloride salts ...
. A
model
A model is an informative representation of an object, person or system. The term originally denoted the plans of a building in late 16th-century English, and derived via French and Italian ultimately from Latin ''modulus'', a measure.
Models c ...
that relies on a set of
differential equation
In mathematics, a differential equation is an equation that relates one or more unknown functions and their derivatives. In applications, the functions generally represent physical quantities, the derivatives represent their rates of change, an ...
s and describes each component of an excitable cell as an electrical element was in good agreement with their empirical measurements.
![Hodgkin-Cycle](https://upload.wikimedia.org/wikipedia/commons/d/d0/Hodgkin-Cycle.jpg)
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 is now known as the
Hodgkin cycle
In membrane biology, the Hodgkin cycle is a key component of membrane physiology that describes bioelectrical impulses, especially prevalent in neural and muscle tissues. It was identified by British physiologist and biophysicist Sir Alan Lloy ...
.
In addition, Hodgkin and Huxley's findings led them to hypothesize the existence of
ion channel
Ion channels are pore-forming membrane proteins that allow ions to pass through the channel pore. Their functions include establishing a resting membrane potential, shaping action potentials and other electrical signals by gating the flow of io ...
s on cell membranes, which were confirmed only decades later. Confirmation of ion channels came with the development of the
patch clamp
The patch clamp technique is a laboratory technique in electrophysiology used to study ionic currents in individual isolated living cells, tissue sections, or patches of cell membrane. The technique is especially useful in the study of excita ...
leading to a Nobel prize in 1991 for
Erwin Neher
Erwin Neher (; ; born 20 March 1944) is a German biophysicist, specializing in the field of cell physiology. For significant contribution in the field, in 1991 he was awarded, along with Bert Sakmann, the Nobel Prize in Physiology or Medicine for ...
and
Bert Sakmann
Bert Sakmann (; born 12 June 1942) is a German cell physiologist. He shared the Nobel Prize in Physiology or Medicine with Erwin Neher in 1991 for their work on "the function of single ion channels in cells," and the invention of the patch cl ...
, and in 2003 for
Roderick MacKinnon
Roderick MacKinnon (born February 19, 1956) is an American biophysicist, neuroscientist, and businessman. He is a professor of Molecular Neurobiology and Biophysics at Rockefeller University who won the Nobel Prize in Chemistry together with Peter ...
.
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
Richard Darwin Keynes, CBE, FRS ( ; 14 August 1919 – 12 June 2010) was a British physiologist. The great-grandson of Charles Darwin, Keynes edited his great-grandfather's accounts and illustrations of Darwin's famous voyage aboard into ''T ...
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. A few year later, the Danish scientist
Jens Christian Skou
Jens Christian Skou (; 8 October 1918 – 28 May 2018) was a Danish biochemist and Nobel laureate.
Early life
Skou was born in Lemvig, Denmark to a wealthy family. His father Magnus Martinus Skou was a timber and coal merchant. His mother Ane-Ma ...
discovered the enzyme
Na+/K+-ATPase
NA, N.A., Na, nA or n/a may refer to:
Chemistry and physics
* Sodium, symbol Na, a chemical element
* Avogadro constant (''N''A)
* Nucleophilic addition, a type of reaction in organic chemistry
* Numerical aperture, a number that characterizes a ...
that uses
ATP to export three sodium ions in exchange for two potassium ions that are imported, for which he received the
Nobel Prize in Chemistry
)
, image = Nobel Prize.png
, alt = A golden medallion with an embossed image of a bearded man facing left in profile. To the left of the man is the text "ALFR•" then "NOBEL", and on the right, the text (smaller) "NAT•" then "M ...
in 1997.
Hodgkin was nominated for the
Nobel Prize in Physiology or Medicine
The Nobel Prize in Physiology or Medicine is awarded yearly by the Nobel Assembly at the Karolinska Institute for outstanding discoveries in physiology or medicine. The Nobel Prize is not a single prize, but five separate prizes that, accord ...
in 1953 by
Lord Adrian. 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
Georg von Békésy ( hu, Békésy György, ; 3 June 1899 – 13 June 1972) was a Hungarian-American biophysicist.
By using strobe photography and silver flakes as a marker, he was able to observe that the basilar membrane moves like a surface w ...
. 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". During the
Nobel Banquet
The Nobel Banquet ( sv, Nobelfesten) is an annual banquet held on 10 December in the Blue Hall of Stockholm City Hall, after the Nobel Prize ceremony. At the banquet, for which a formal dress code exists, a multi-course dinner is served and ent ...
on 10 December 1963, Hodgkin gave the traditional speech on behalf of the three neurophysiologists, thanking the
king
King is the title given to a male monarch in a variety of contexts. The female equivalent is queen, which title is also given to the consort of a king.
*In the context of prehistory, antiquity and contemporary indigenous peoples, the tit ...
and the
for the award. 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
Francis Peyton Rous () (October 5, 1879 – February 16, 1970) was an American pathologist at the Rockefeller University known for his works in oncoviruses, blood transfusion and physiology of digestion. A medical graduate from the Johns Hopki ...
, was awarded the 1966
Nobel Prize in Physiology or Medicine
The Nobel Prize in Physiology or Medicine is awarded yearly by the Nobel Assembly at the Karolinska Institute for outstanding discoveries in physiology or medicine. The Nobel Prize is not a single prize, but five separate prizes that, accord ...
.
Later career and administrative positions
From 1951 to 1969, Hodgkin was the Foulerton Professor of the
Royal Society
The Royal Society, formally The Royal Society of London for Improving Natural Knowledge, is a learned society and the United Kingdom's national academy of sciences. The society fulfils a number of roles: promoting science and its benefits, re ...
at
Cambridge
Cambridge ( ) is a university city and the county town in Cambridgeshire, England. It is located on the River Cam approximately north of London. As of the 2021 United Kingdom census, the population of Cambridge was 145,700. Cambridge bec ...
. In 1970 he became the John Humphrey Plummer Professor of Biophysics at
Cambridge
Cambridge ( ) is a university city and the county town in Cambridgeshire, England. It is located on the River Cam approximately north of London. As of the 2021 United Kingdom census, the population of Cambridge was 145,700. Cambridge bec ...
. Around this time he also ended his experiments on nerve at the
Plymouth Marine Laboratory
Plymouth Marine Laboratory (abbreviated as PML) in the city of Plymouth, England, is a marine research organization and registered charity. It is a partner of the UK Research & Innovation's Natural Environment Research Council (NERC). PML's chai ...
and switched his focus to visual research which he could do in
Cambridge
Cambridge ( ) is a university city and the county town in Cambridgeshire, England. It is located on the River Cam approximately north of London. As of the 2021 United Kingdom census, the population of Cambridge was 145,700. Cambridge bec ...
with the help of others while serving as
president of the Royal Society
The president of the Royal Society (PRS) is the elected Head of the Royal Society of London who presides over meetings of the society's council.
After informal meetings at Gresham College, the Royal Society was officially founded on 28 November ...
. Together with
Denis Baylor
Denis Aristide Baylor (January 30, 1940 – March 16, 2022) was an American neurobiologist. He was professor emeritus of neurobiology at Stanford University. He is known for his research on nerve cells in the retina of the eye. He developed a w ...
and Peter Detwiler he published a series of papers on turtle
photoreceptors.
From 1970 to 1975 Hodgkin served as the 53rd
president of the Royal Society
The president of the Royal Society (PRS) is the elected Head of the Royal Society of London who presides over meetings of the society's council.
After informal meetings at Gresham College, the Royal Society was officially founded on 28 November ...
(PRS). During his tenure as PRS, he was knighted in 1972 and admitted into the
Order of Merit in 1973.
From 1978 to 1984 he was the 34th
Master of Trinity College, Cambridge
The following have served as Master of Trinity College, Cambridge:
{, class="wikitable"
, -
!Name
!Portrait
!colspan=2, Term of office
, -
, John Redman
,
, 1546
, 1551
, -
, William Bill
,
, 1551
, 1553
, -
, John Christopherson
,
, 1553
, ...
.
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
, mottoeng = So that they may have life
, established =
, type = public research university
, endowment = £20.0 million
, budget = £326 million
, chancellor = David Willetts
, vice_chancellor = Nishan Canagarajah
, head_lab ...
, from 1971 to 1984
Awards and honours
* 1988 – W.H. Helmerich III Award of the Retina Research Foundation
* 1983 – Lord Crook Medal of the Worshipful Company of Spectacle Makers
* 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
* 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
The Indian National Science Academy (INSA) is a national academy in New Delhi for Indian scientists in all branches of science and technology.
In August 2019, Dr. Chandrima Shaha was appointed as the president of Indian National Science Acade ...
* 1970 –
President of the Royal Society
The president of the Royal Society (PRS) is the elected Head of the Royal Society of London who presides over meetings of the society's council.
After informal meetings at Gresham College, the Royal Society was officially founded on 28 November ...
(PRS)
[
* 1968 – Member, Pontifical Academy of Sciences
* 1968 – Foreign Member, ]American Philosophical Society
The American Philosophical Society (APS), founded in 1743 in Philadelphia, is a scholarly organization that promotes knowledge in the sciences and humanities through research, professional meetings, publications, library resources, and communit ...
* 1966 – President of the Marine Biological Association of the United Kingdom
The Marine Biological Association of the United Kingdom (MBA) is a learned society with a scientific laboratory that undertakes research in marine biology. The organisation was founded in 1884 and has been based in Plymouth since the Citadel H ...
* 1965 – Copley Medal
The Copley Medal is an award given by the Royal Society, for "outstanding achievements in research in any branch of science". It alternates between the physical sciences or mathematics and the biological sciences. Given every year, the medal is t ...
of the Royal Society
The Royal Society, formally The Royal Society of London for Improving Natural Knowledge, is a learned society and the United Kingdom's national academy of sciences. The society fulfils a number of roles: promoting science and its benefits, re ...
* 1964 – Foreign Member, Royal Danish Academy of Sciences and Letters
{{Infobox organization
, name = The Royal Danish Academy of Sciences and Letters
, full_name =
, native_name = Det Kongelige Danske Videnskabernes Selskab
, native_name_lang =
, logo = Royal ...
* 1964 – Member, German National Academy of Sciences Leopoldina
The German National Academy of Sciences Leopoldina (german: Deutsche Akademie der Naturforscher Leopoldina – Nationale Akademie der Wissenschaften), short Leopoldina, is the national academy of Germany, and is located in Halle (Saale). Founded ...
* 1963 – Nobel Prize for Physiology or Medicine together with Andrew Fielding Huxley, and John Carew Eccles (for his research on synapse
In the nervous system, a synapse is a structure that permits a neuron (or nerve cell) to pass an electrical or chemical signal to another neuron or to the target effector cell.
Synapses are essential to the transmission of nervous impulses from ...
s)
* 1962 – Foreign Hon. Member, American Academy of Arts and Sciences
The American Academy of Arts and Sciences (abbreviation: AAA&S) is one of the oldest learned societies in the United States. It was founded in 1780 during the American Revolution by John Adams, John Hancock, James Bowdoin, Andrew Oliver, and ...
* 1958 – Royal Medal
The Royal Medal, also known as The Queen's Medal and The King's Medal (depending on the gender of the monarch at the time of the award), is a silver-gilt medal, of which three are awarded each year by the Royal Society, two for "the most important ...
of The Royal Society
The Royal Society, formally The Royal Society of London for Improving Natural Knowledge, is a learned society and the United Kingdom's national academy of sciences. The society fulfils a number of roles: promoting science and its benefits, r ...
* 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
The Royal Society, formally The Royal Society of London for Improving Natural Knowledge, is a learned society and the United Kingdom's national academy of sciences. The society fulfils a number of roles: promoting science and its benefits, r ...
(FRS)[
A portrait of Hodgkin by ]Michael Noakes
Michael Noakes (28 October 1933 – 30 May 2018) was an English artist and portrait painter. hangs in Trinity College's collection.
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
Francis Peyton Rous () (October 5, 1879 – February 16, 1970) was an American pathologist at the Rockefeller University known for his works in oncoviruses, blood transfusion and physiology of digestion. A medical graduate from the Johns Hopki ...
who was later awarded the 1966 Nobel Prize in Physiology or Medicine
The Nobel Prize in Physiology or Medicine is awarded yearly by the Nobel Assembly at the Karolinska Institute for outstanding discoveries in physiology or medicine. The Nobel Prize is not a single prize, but five separate prizes that, accord ...
. When Rous invited him for dinner to his home, Hodgkin got to know Rous' daughter, Marni
The name Marni originates from several languages, including Hebrew, meaning "rejoice", and Latin as a variant of " Marina", meaning "of the sea". It also has derivations from Gaelic and Swahili. "Marni" and "Marnie" are the two most common spell ...
, who was then a student at Swarthmore College
Swarthmore College ( , ) is a Private college, private Liberal arts colleges in the United States, liberal arts college in Swarthmore, Pennsylvania. Founded in 1864, with its first classes held in 1869, Swarthmore is one of the earliest coeduca ...
. He proposed to her before going back to England in 1938, but she rejected him. When Hodgkin briefly returned to the US in 1944 (see Wartime activities), they reunited and got married on 31 March. Their first daughter, Sarah, was born in April 1945, shortly before the Hodgkins moved back to Cambridge. They had three more children: Deborah Hodgkin (born 2 May 1947), Jonathan Hodgkin (born 24 August 1949), and Rachel Hodgkin (born June 1951).
Marni became 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
Thomas Hodgkin RMS (17 August 1798 – 5 April 1866) was a British physician, considered one of the most prominent pathologists of his time and a pioneer in preventive medicine. He is now best known for the first account of Hodgkin's disease, ...
(1798–1866), who first described Hodgkin's lymphoma
Hodgkin lymphoma (HL) is a type of lymphoma, in which cancer originates from a specific type of white blood cell called lymphocytes, where multinucleated Reed–Sternberg cells (RS cells) are present in the patient's lymph nodes. The condition wa ...
, was Alan Hodgkin's ancestor.
Death
Hodgkin suffered from a series of medical problems that began soon after his retirement as Master of Trinity. In 1989 he had a 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.[ Hodgkin died in 1998 in ]Cambridge
Cambridge ( ) is a university city and the county town in Cambridgeshire, England. It is located on the River Cam approximately north of London. As of the 2021 United Kingdom census, the population of Cambridge was 145,700. Cambridge bec ...
.
References
Bibliography
*
External links
The Master of Trinity
at Trinity College, Cambridge
Trinity College is a constituent college of the University of Cambridge. Founded in 1546 by Henry VIII, King Henry VIII, Trinity is one of the largest Cambridge colleges, with the largest financial endowment of any college at either Cambridge ...
* including the Nobel Lecture on 11 December 1963 ''The Ionic Basis of Nervous Conduction''
*
BBC obituary
Action Potential Paper
Imperial War Museum Interview
{{DEFAULTSORT:Hodgkin, Alan Lloyd
1914 births
1998 deaths
Alumni of Trinity College, Cambridge
British physiologists
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Fellows of the Indian National Science Academy
Fellows of the Royal Society
Foreign associates of the National Academy of Sciences
Alan
Alan may refer to:
People
*Alan (surname), an English and Turkish surname
* Alan (given name), an English given name
**List of people with given name Alan
''Following are people commonly referred to solely by "Alan" or by a homonymous name.''
*A ...
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