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John Burdon Sanderson Haldane (; 5 November 18921 December 1964), nicknamed "Jack" or "JBS", was a British-Indian scientist who worked in
physiology Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemic ...
,
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar work ...
,
evolutionary biology Evolutionary biology is the subfield of biology that studies the evolutionary processes (natural selection, common descent, speciation) that produced the diversity of life on Earth. It is also defined as the study of the history of life ...
, and
mathematics Mathematics is an area of knowledge that includes the topics of numbers, formulas and related structures, shapes and the spaces in which they are contained, and quantities and their changes. These topics are represented in modern mathematics ...
. With innovative use of statistics in biology, he was one of the founders of
neo-Darwinism Neo-Darwinism is generally used to describe any integration of Charles Darwin's theory of evolution by natural selection with Gregor Mendel's theory of genetics. It mostly refers to evolutionary theory from either 1895 (for the combinations of Dar ...
. He served in the
Great War World War I (28 July 1914 11 November 1918), often abbreviated as WWI, was one of the deadliest global conflicts in history. Belligerents included much of Europe, the Russian Empire, the United States, and the Ottoman Empire, with fightin ...
, and obtained the rank of captain. Despite his lack of an academic degree in the field, he taught biology at the
University of Cambridge , mottoeng = Literal: From here, light and sacred draughts. Non literal: From this place, we gain enlightenment and precious knowledge. , established = , other_name = The Chancellor, Masters and Schola ...
, the
Royal Institution The Royal Institution of Great Britain (often the Royal Institution, Ri or RI) is an organisation for scientific education and research, based in the City of Westminster. It was founded in 1799 by the leading British scientists of the age, inc ...
, and
University College London , mottoeng = Let all come who by merit deserve the most reward , established = , type = Public research university , endowment = £143 million (2020) , budget = ...
. Renouncing his British citizenship, he became an Indian citizen in 1961 and worked at the
Indian Statistical Institute Indian Statistical Institute (ISI) is a higher education and research institute which is recognized as an Institute of National Importance by the 1959 act of the Indian parliament. It grew out of the Statistical Laboratory set up by Prasanta C ...
for the rest of his life. Haldane's article on
abiogenesis In biology, abiogenesis (from a- 'not' + Greek bios 'life' + genesis 'origin') or the origin of life is the natural process by which life has arisen from non-living matter, such as simple organic compounds. The prevailing scientific hypothes ...
in 1929 introduced the " primordial soup theory", which became the foundation for the concept of the chemical origin of life. He established human gene maps for
haemophilia Haemophilia, or hemophilia (), is a mostly inherited genetic disorder that impairs the body's ability to make blood clots, a process needed to stop bleeding. This results in people bleeding for a longer time after an injury, easy bruisin ...
and
colour blindness Color blindness or color vision deficiency (CVD) is the decreased ability to see color or differences in color. It can impair tasks such as selecting ripe fruit, choosing clothing, and reading traffic lights. Color blindness may make some aca ...
on the
X chromosome The X chromosome is one of the two sex-determining chromosomes (allosomes) in many organisms, including mammals (the other is the Y chromosome), and is found in both males and females. It is a part of the XY sex-determination system and XO sex ...
, and codified Haldane's rule on sterility in the
heterogametic sex Heterogametic sex (digametic sex) refers to the individuals of a species in which the sex chromosomes are not the same. For example, in humans, males with an X and a Y sex chromosome would be referred to as the heterogametic sex, and females ...
of hybrids in
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
. He correctly proposed that
sickle-cell disease Sickle cell disease (SCD) is a group of blood disorders typically inherited from a person's parents. The most common type is known as sickle cell anaemia. It results in an abnormality in the oxygen-carrying protein haemoglobin found in red ...
confers some immunity to
malaria Malaria is a mosquito-borne infectious disease that affects humans and other animals. Malaria causes symptoms that typically include fever, tiredness, vomiting, and headaches. In severe cases, it can cause jaundice, seizures, coma, or death. ...
. He was the first to suggest the central idea of
in vitro fertilisation In vitro fertilisation (IVF) is a process of fertilisation where an egg is combined with sperm in vitro ("in glass"). The process involves monitoring and stimulating an individual's ovulatory process, removing an ovum or ova (egg or eggs) ...
, as well as concepts such as
hydrogen economy The hydrogen economy is using hydrogen to decarbonize economic sectors which are hard to electrify, essentially, the "hard-to-abate" sectors such as cement, steel, long-haul transport etc. In order to phase out fossil fuels and limit climate ch ...
, cis and
trans-acting In the field of molecular biology, ''trans''-acting (''trans''-regulatory, ''trans''-regulation), in general, means "acting from a different molecule" (''i.e.'', intermolecular). It may be considered the opposite of ''cis''-acting (''cis''-regula ...
regulation,
coupling reaction A coupling reaction in organic chemistry is a general term for a variety of reactions where two fragments are joined together with the aid of a metal catalyst. In one important reaction type, a main group organometallic compound of the type R-M (R = ...
, molecular repulsion, the darwin (as a unit of evolution), and organismal cloning. In 1957, Haldane articulated
Haldane's dilemma Haldane's dilemma, also known as "the waiting time problem", is a limit on the speed of beneficial evolution, calculated by J. B. S. Haldane in 1957. Before the invention of DNA sequencing technologies, it was not known how much polymorphism ...
, a limit on the speed of beneficial
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
, an idea which is still debated today. He willed his body for medical studies, as he wanted to remain useful even in death. He is also remembered for his work in
human biology Human biology is an interdisciplinary area of academic study that examines humans through the influences and interplay of many diverse fields such as genetics, evolution, physiology, anatomy, epidemiology, anthropology, ecology, nutrition, populat ...
, having coined " clone", "
cloning Cloning is the process of producing individual organisms with identical or virtually identical DNA, either by natural or artificial means. In nature, some organisms produce clones through asexual reproduction. In the field of biotechnology, c ...
", and "
ectogenesis Ectogenesis (from the Greek ''ἐκτός'', "outside," and '' genesis'') is the growth of an organism in an artificial environment outside the body in which it would normally be found, such as the growth of an embryo or fetus outside the mother's ...
". With his sister,
Naomi Mitchison Naomi Mary Margaret Mitchison, Baroness Mitchison (; 1 November 1897 – 11 January 1999) was a List of Scottish novelists, Scottish novelist and poet. Often called a doyenne of Scottish literature, she wrote over 90 books of historical and sci ...
, Haldane was the first to demonstrate
genetic linkage Genetic linkage is the tendency of DNA sequences that are close together on a chromosome to be inherited together during the meiosis phase of sexual reproduction. Two genetic markers that are physically near to each other are unlikely to be sep ...
in
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s. Subsequent works established a unification of
Mendelian genetics Mendelian inheritance (also known as Mendelism) is a type of biological inheritance following the principles originally proposed by Gregor Mendel in 1865 and 1866, re-discovered in 1900 by Hugo de Vries and Carl Correns, and later populari ...
and
Darwinian evolution Darwinism is a theory of biological evolution developed by the English naturalist Charles Darwin (1809–1882) and others, stating that all species of organisms arise and develop through the natural selection of small, inherited variations that ...
by
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
whilst laying the groundwork for
modern synthesis Modern synthesis or modern evolutionary synthesis refers to several perspectives on evolutionary biology, namely: * Modern synthesis (20th century), the term coined by Julian Huxley in 1942 to denote the synthesis between Mendelian genetics and ...
, and helped to create
population genetics Population genetics is a subfield of genetics that deals with genetic differences within and between populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as adaptation, speciation, and po ...
. Haldane was a professed
socialist Socialism is a left-wing economic philosophy and movement encompassing a range of economic systems characterized by the dominance of social ownership of the means of production as opposed to private ownership. As a term, it describes the ...
,
Marxist Marxism is a left-wing to far-left method of socioeconomic analysis that uses a materialist interpretation of historical development, better known as historical materialism, to understand class relations and social conflict and a dialecti ...
,
atheist Atheism, in the broadest sense, is an absence of belief in the existence of deities. Less broadly, atheism is a rejection of the belief that any deities exist. In an even narrower sense, atheism is specifically the position that there no ...
, and
secular humanist Secular humanism is a philosophy, belief system or life stance that embraces human reason, secular ethics, and philosophical naturalism while specifically rejecting religious dogma, supernaturalism, and superstition as the basis of morality ...
whose political dissent led him to leave England in 1956 and live in India, becoming a naturalised Indian citizen in 1961.
Arthur C. Clarke Sir Arthur Charles Clarke (16 December 191719 March 2008) was an English science-fiction writer, science writer, futurist, inventor, undersea explorer, and television series host. He co-wrote the screenplay for the 1968 film '' 2001: A Spac ...
credited him as "perhaps the most brilliant science populariser of his generation". Brazilian-British biologist and
Nobel laureate The Nobel Prizes ( sv, Nobelpriset, no, Nobelprisen) are awarded annually by the Royal Swedish Academy of Sciences, the Swedish Academy, the Karolinska Institutet, and the Norwegian Nobel Committee to individuals and organizations who make o ...
Peter Medawar Sir Peter Brian Medawar (; 28 February 1915 – 2 October 1987) was a Brazilian-British biologist and writer, whose works on graft rejection and the discovery of acquired immune tolerance have been fundamental to the medical practice of tissu ...
called Haldane "the cleverest man I ever knew". According to
Theodosius Dobzhansky Theodosius Grigorievich Dobzhansky (russian: Феодо́сий Григо́рьевич Добржа́нский; uk, Теодо́сій Григо́рович Добржа́нський; January 25, 1900 – December 18, 1975) was a prominent ...
, "Haldane was always recognized as a singular case"; Ernst Mayr described him as a "polymath"; Michael J. D. White described him as "the most erudite biologist of his generation, and perhaps of the century";
James Watson James Dewey Watson (born April 6, 1928) is an American molecular biologist, geneticist, and zoologist. In 1953, he co-authored with Francis Crick the academic paper proposing the double helix structure of the DNA molecule. Watson, Crick a ...
described him as "England's most clever and eccentric biologist" and
Sahotra Sarkar Sahotra Sarkar (born 1962) is an Indian-American professor at the University of Texas at Austin, specializing in the philosophy of biology. Education and career Sarkar is originally from India where he lived in Darjeeling until 1975. He earned ...
described him as "probably the most prescient biologist of this 0thcentury." According to a Cambridge student, "he seemed to be the last man who might know all there was to be known."


Biography


Early life and education

Haldane was born in
Oxford Oxford () is a city in England. It is the county town and only city of Oxfordshire. In 2020, its population was estimated at 151,584. It is north-west of London, south-east of Birmingham and north-east of Bristol. The city is home to the ...
in 1892. His father was
John Scott Haldane John Scott Haldane (; 2 May 1860 – 14/15 March 1936) was a British physician and physiologist famous for intrepid self-experimentation which led to many important discoveries about the human body and the nature of gases. He also experime ...
, a
physiologist Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemica ...
, scientist, a
philosopher A philosopher is a person who practices or investigates philosophy. The term ''philosopher'' comes from the grc, φιλόσοφος, , translit=philosophos, meaning 'lover of wisdom'. The coining of the term has been attributed to the Greek th ...
and a Liberal who was the grandson of evangelist James Alexander Haldane. His mother Louisa Kathleen Trotter, was a
Conservative Conservatism is a cultural, social, and political philosophy that seeks to promote and to preserve traditional institutions, practices, and values. The central tenets of conservatism may vary in relation to the culture and civilization in ...
, and descended from Scottish ancestry. His only sibling,
Naomi Naomi or Naomie may refer to: People and biblical figures * Naomi (given name), a female given name and a list of people with the name * Naomi (biblical figure), Ruth's mother-in-law in the Old Testament Book of Ruth * Naomi (Romanian singer) (bor ...
, became a writer and married
Dick Mitchison, Baron Mitchison Gilbert Richard Mitchison, Baron Mitchison, (23 March 1894 – 14 February 1970) was a British Labour politician. Born in Staines, Mitchison was educated at Eton College and New College, Oxford, and became a barrister (called to the bar in 191 ...
(thereby becoming Naomi Mary Margaret Mitchison, Baroness Mitchison), who was his best friend at
Eton College Eton College () is a Public school (United Kingdom), public school in Eton, Berkshire, England. It was founded in 1440 by Henry VI of England, Henry VI under the name ''Kynge's College of Our Ladye of Eton besyde Windesore'',Nevill, p. 3 ff. i ...
. His uncle was
Viscount Haldane A viscount ( , for male) or viscountess (, for female) is a title used in certain European countries for a noble of varying status. In many countries a viscount, and its historical equivalents, was a non-hereditary, administrative or judicial ...
and his aunt the author Elizabeth Haldane. Descended from an aristocratic and
secular Secularity, also the secular or secularness (from Latin ''saeculum'', "worldly" or "of a generation"), is the state of being unrelated or neutral in regards to religion. Anything that does not have an explicit reference to religion, either negativ ...
family of the
Clan Haldane Clan Haldane is a Lowland Scottish clan.Way, George and Squire, Romily. ''Collins Scottish Clan & Family Encyclopedia''. (Foreword by The Rt Hon. The Earl of Elgin KT, Convenor, The Standing Council of Scottish Chiefs). Published in 1994. Pages ...
, he would later claim that his
Y chromosome The Y chromosome is one of two sex chromosomes (allosomes) in therian mammals, including humans, and many other animals. The other is the X chromosome. Y is normally the sex-determining chromosome in many species, since it is the presence or abs ...
could be traced back to
Robert the Bruce Robert I (11 July 1274 – 7 June 1329), popularly known as Robert the Bruce (Scottish Gaelic: ''Raibeart an Bruis''), was King of Scots from 1306 to his death in 1329. One of the most renowned warriors of his generation, Robert eventuall ...
. Haldane grew up at 11 Crick Road, North Oxford. He learnt to read at the age of three, and at four, after injuring his forehead he asked the doctor of the bleeding, "Is this oxyhaemoglobin or carboxyhaemoglobin?" As a youth he was raised as an Anglican. From age eight he worked with his father in their home laboratory where he experienced his first
self-experimentation Self-experimentation refers to the special case of single-subject research in which the experimenter conducts the experiment on themselves. Usually this means that a single person is the designer, operator, subject, analyst, and user or reporte ...
, the method he would later be famous for. He and his father became their own "human guinea pigs", such as in their investigation on the effects of poison gases. In 1899 his family moved to "Cherwell", a late Victorian house at the outskirts of Oxford with its own private laboratory. At age 8, in 1901, his father brought him to the Oxford University Junior Scientific Club to listen to a lecture on
Mendelian genetics Mendelian inheritance (also known as Mendelism) is a type of biological inheritance following the principles originally proposed by Gregor Mendel in 1865 and 1866, re-discovered in 1900 by Hugo de Vries and Carl Correns, and later populari ...
, which had been recently rediscovered. Although he found the lecture given by Arthur Dukinfield Darbishire, Demonstrator of Zoology at
Balliol College, Oxford Balliol College () is one of the constituent colleges of the University of Oxford in England. One of Oxford's oldest colleges, it was founded around 1263 by John I de Balliol, a landowner from Barnard Castle in County Durham, who provided the ...
, "interesting but difficult," it influenced him permanently such that genetics became the field in which he made his most important scientific contributions. His formal education began in 1897 at Oxford Preparatory School (now
Dragon School ("Reach for the Sun") , established = 1877 , closed = , type = Preparatory day and boarding school and Pre-Prep school , religion = Church of England , president = , head_label = Head , head = Emma Goldsm ...
), where he gained a First Scholarship in 1904 to Eton. In 1905 he joined Eton, where he experienced severe abuse from senior students for allegedly being arrogant. The indifference of authority left him with a lasting hatred for the English education system. However, the ordeal did not stop him from becoming Captain of the school. He participated for the first time in scientific research as a volunteer subject for his father in 1906. John was the first to study the effects of decompression (relief from high pressure) in humans. He investigated the physiological condition called "bends," such as when goats lift and bend their legs if discomforted, that is also experienced by deep-sea divers. In July 1906, on board '' HMS Spanker'' off the west coast of Scotland, Rothesay, young Haldane jumped into the Atlantic Ocean with the experimental diving suit. The study was published in a 101-paged article in ''
The Journal of Hygiene ''Epidemiology and Infection'' is a peer-reviewed medical journal that contains original reports and reviews on all aspects of infection in humans and animals. Some of these aspects include zoonoses, tropical infections, food hygiene, and vac ...
'' in 1908; where Haldane was described as "Jack Haldane (age 13)" for whom it "was the first time ehad ever dived in a diving dress." The research became a foundation for a scientific theory called
Haldane's decompression model Haldane's decompression model is a mathematical model for Decompression (diving), decompression to sea level atmospheric pressure of divers breathing compressed air at ambient pressure that was proposed in 1908 by the Scottish physiologist, J ...
. He studied mathematics and
classics Classics or classical studies is the study of classical antiquity. In the Western world, classics traditionally refers to the study of Classical Greek and Roman literature and their related original languages, Ancient Greek and Latin. Classics ...
at
New College, Oxford New College is one of the constituent colleges of the University of Oxford in the United Kingdom. Founded in 1379 by William of Wykeham in conjunction with Winchester College as its feeder school, New College is one of the oldest colleges at ...
and obtained first-class honours in mathematical
moderations Honour Moderations (or ''Mods'') are a set of examinations at the University of Oxford at the end of the first part of some degree courses (e.g., Greats or '' Literae Humaniores''). Honour Moderations candidates have a class awarded (hence the ' ...
in 1912. He became engrossed in genetics and presented a paper on gene linkage in
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
s in the summer of 1912. His first technical paper, a 30-page long article on haemoglobin function, was published that same year, as a co-author alongside his father. He presented the mathematical treatment of the study on 19 October in the ''Proceedings of the Physiological Society'' and was published in December 1913. Haldane did not want his education to be confined to a specific subject. He took up Greats and graduated with first-class honours in 1914. While he had full intention of studying physiology, his plan was, as he described later, "somewhat overshadowed by other events" (referring to World War I). His only formal education in biology was an incomplete course in vertebrate anatomy.


Career

To support the war effort, Haldane volunteered for and joined the
British Army The British Army is the principal land warfare force of the United Kingdom, a part of the British Armed Forces along with the Royal Navy and the Royal Air Force. , the British Army comprises 79,380 regular full-time personnel, 4,090 Gurkha ...
, and was commissioned a temporary second lieutenant in the 3rd Battalion of the
Black Watch The Black Watch, 3rd Battalion, Royal Regiment of Scotland (3 SCOTS) is an infantry battalion of the Royal Regiment of Scotland. The regiment was created as part of the Childers Reforms in 1881, when the 42nd (Royal Highland) Regime ...
(Royal Highland Regiment) on 15 August 1914. He was assigned as the trench mortar officer, to lead his team for hand-bombing the enemy trenches, the experience of which he remarked "enjoyable." In his article in 1932 he described how "he enjoyed the opportunity of killing people and regarded this as a respectable relic of primitive man." He was promoted to temporary lieutenant on 18 February 1915 and to temporary captain on 18 October. While serving in France, he was wounded by an artillery fire for which he was sent back to Scotland. There he served as instructor of grenades for the Black Watch recruits. In 1916, he joined the war in Mesopotamia (Iraq) where an enemy bomb severely wounded him. He was relieved from war fronts and was sent to India and stayed there for the rest of the war. He returned to England in 1919 and relinquished his commission on 1 April 1920, retaining his rank of captain. For his ferocity and aggressiveness in battles, his commander Douglas Haig described him as the "bravest and dirtiest officer in my Army." Between 1919 and 1922, he served as
Fellow A fellow is a concept whose exact meaning depends on context. In learned or professional societies, it refers to a privileged member who is specially elected in recognition of their work and achievements. Within the context of higher education ...
of New College, Oxford, where he taught and researched in physiology and genetics, despite his lack of formal education in the field. During his first year at Oxford, he published six papers dealing with physiology of respiration and genetics. He then moved to the
University of Cambridge , mottoeng = Literal: From here, light and sacred draughts. Non literal: From this place, we gain enlightenment and precious knowledge. , established = , other_name = The Chancellor, Masters and Schola ...
, where he accepted a newly created readership in
Biochemistry Biochemistry or biological chemistry is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology and ...
, in 1923 and taught until 1932. During his nine years at Cambridge, he worked on
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products ...
s and
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar work ...
, particularly the mathematical side of genetics. While working as a visiting professor at the
University of California The University of California (UC) is a public land-grant research university system in the U.S. state of California. The system is composed of the campuses at Berkeley, Davis, University of California, Irvine, Irvine, University of Califor ...
in 1932, he was elected
Fellow of the Royal Society Fellowship of the Royal Society (FRS, ForMemRS and HonFRS) is an award granted by the judges of the Royal Society of London to individuals who have made a "substantial contribution to the improvement of natural knowledge, including mathemati ...
. Haldane worked part-time at the John Innes Horticultural Institution (later named
John Innes Centre The John Innes Centre (JIC), located in Norwich, Norfolk, England, is an independent centre for research and training in plant and microbial science founded in 1910. It is a registered charity (No 223852) grant-aided by the Biotechnology and ...
) at
Merton Park Merton Park is a suburb in the London Borough of Merton. It is situated between Colliers Wood, Morden, South Wimbledon and Raynes Park. It is 11 miles (11.7 km) southwest of Charing Cross. The area is part of the historic parish of Merton i ...
in Surrey from 1927 to 1937. When Alfred Daniel Hall became the Director in 1926, one of his earliest tasks was to appoint as assistant director "a man of high quality in the study of genetics" who could become his successor. Recommended by
Julian Huxley Sir Julian Sorell Huxley (22 June 1887 – 14 February 1975) was an English evolutionary biologist, eugenicist, and internationalist. He was a proponent of natural selection, and a leading figure in the mid-twentieth century modern synthesis. ...
, the council appointed Haldane in March 1927, with the terms: "Mr Haldane to visit the Institution fortnightly for a day and a night during the Cambridge terms, to put in two months also at Easter and long vacations in two continuous blocks and to be free in the Christmas vacation." He was Officer in charge of Genetical Investigations. He became the
Fullerian Professor of Physiology The Fullerian Chairs at the Royal Institution in London, England, were established by John 'Mad Jack' Fuller. Fullerian Professors of Physiology & Comparative Anatomy * 1834–1837 Peter Mark Roget * 1837–1838 Robert Edmond Grant * 1841–1844 ...
at the
Royal Institution The Royal Institution of Great Britain (often the Royal Institution, Ri or RI) is an organisation for scientific education and research, based in the City of Westminster. It was founded in 1799 by the leading British scientists of the age, inc ...
from 1930 to 1932 and in 1933 he became full Professor of Genetics at
University College London , mottoeng = Let all come who by merit deserve the most reward , established = , type = Public research university , endowment = £143 million (2020) , budget = ...
, where he spent most of his
academic An academy (Attic Greek: Ἀκαδήμεια; Koine Greek Ἀκαδημία) is an institution of secondary or tertiary higher learning (and generally also research or honorary membership). The name traces back to Plato's school of philosophy, ...
career. As Hall did not retire as early as expected – retiring in 1939, Haldane had to resign from the John Innes in 1936 to become the first Weldon Professor of
Biometry Biostatistics (also known as biometry) are the development and application of statistical methods to a wide range of topics in biology. It encompasses the design of biological experiments, the collection and analysis of data from those experimen ...
at University College London. Haldane's service was recorded to have helped the John Innes as "the liveliest place for research in genetics in Britain." At the height 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 World War II by country, vast majority of the world's countries—including all of the great power ...
, he moved his team to the
Rothamsted Experimental Station Rothamsted Research, previously known as the Rothamsted Experimental Station and then the Institute of Arable Crops Research, is one of the oldest agricultural research institutions in the world, having been founded in 1843. It is located at Har ...
in Hertfordshire during 1941 to 1944 to escape bombings. Complying an invitation of
Reginald Punnett Reginald Crundall Punnett FRS (; 20 June 1875 – 3 January 1967) was a British geneticist who co-founded, with William Bateson, the ''Journal of Genetics'' in 1910. Punnett is probably best remembered today as the creator of the Punnett ...
, who founded the ''Journal of Genetics'' in 1910 with
William Bateson William Bateson (8 August 1861 – 8 February 1926) was an English biologist who was the first person to use the term genetics to describe the study of heredity, and the chief populariser of the ideas of Gregor Mendel following their rediscove ...
, he became the editor since 1933 until his death.


In India

In 1956, Haldane left
University College London , mottoeng = Let all come who by merit deserve the most reward , established = , type = Public research university , endowment = £143 million (2020) , budget = ...
, and joined the
Indian Statistical Institute Indian Statistical Institute (ISI) is a higher education and research institute which is recognized as an Institute of National Importance by the 1959 act of the Indian parliament. It grew out of the Statistical Laboratory set up by Prasanta C ...
(ISI) in Calcutta (later renamed
Kolkata Kolkata (, or , ; also known as Calcutta , the official name until 2001) is the capital of the Indian state of West Bengal, on the eastern bank of the Hooghly River west of the border with Bangladesh. It is the primary business, comme ...
), India, where he worked in the
biometry Biostatistics (also known as biometry) are the development and application of statistical methods to a wide range of topics in biology. It encompasses the design of biological experiments, the collection and analysis of data from those experimen ...
unit. Haldane gave many reasons for moving to India. Officially he stated that he left the UK because of the
Suez Crisis The Suez Crisis, or the Second Arab–Israeli war, also called the Tripartite Aggression ( ar, العدوان الثلاثي, Al-ʿUdwān aṯ-Ṯulāṯiyy) in the Arab world and the Sinai War in Israel,Also known as the Suez War or 1956 Wa ...
, writing: "Finally, I am going to India because I consider that recent acts of the British Government have been violations of
international law International law (also known as public international law and the law of nations) is the set of rules, norms, and standards generally recognized as binding between states. It establishes normative guidelines and a common conceptual framework for ...
." He believed that the warm
climate Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologi ...
would do him good, and that India shared his socialist dreams. In an article "A passage to India" which he wrote in ''The Rationalists Annual'' in 1958, he stated: "For one thing I prefer Indian food to American. Perhaps my main reason for going to India is that I consider that the opportunities for scientific research of the kind in which I am interested are better in India than in Britain, and that my teaching will be at least as useful there as here." The university had sacked his wife Helen for being
drunk and disorderly Public intoxication, also known as "drunk and disorderly" and "drunk in public", is a summary offense in some countries rated to public cases or displays of drunkenness. Public intoxication laws vary widely by jurisdiction, but usually require an ...
and refusing to pay a fine, triggering Haldane's resignation. He declared he would no longer wear socks, "Sixty years in socks is enough." and always dressed in Indian attire. Haldane was keenly interested in inexpensive research. Explaining in "A passage to India," he said, "Of course, if my work required electron microscopes,
cyclotrons A cyclotron is a type of particle accelerator invented by Ernest O. Lawrence in 1929–1930 at the University of California, Berkeley, and patented in 1932. Lawrence, Ernest O. ''Method and apparatus for the acceleration of ions'', filed: Janu ...
, and the like, I should not get them in India. But the sort of facilities which Darwin and Bateson used for their researches—such as gardens, gardeners, pigeon lofts, and pigeons—are more easily obtained in India than in England." He wrote to Julian Huxley about his observations on ''Vanellus malabaricus'', the
yellow-wattled lapwing The yellow-wattled lapwing (''Vanellus malabaricus'') is a lapwing that is endemic to the Indian Subcontinent. It is found mainly on the dry plains of peninsular India and has a sharp call and is capable of fast flight. Although they do not mig ...
. He advocated the use of ''Vigna sinensis'' (
cowpea The cowpea (''Vigna unguiculata'') is an annual herbaceous legume from the genus ''Vigna''. Its tolerance for sandy soil and low rainfall have made it an important crop in the semiarid regions across Africa and Asia. It requires very few inputs, ...
) as a model for studying
plant genetics Plant genetics is the study of genes, genetic variation, and heredity specifically in plants.Hartl D, Jones E (2005) It is generally considered a field of biology and botany, but intersects frequently with many other life sciences and is strongly ...
. He took an interest in the
pollination Pollination is the transfer of pollen from an Stamen, anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by Anemophily, wind. Pollinating agents can ...
of ''
Lantana camara ''Lantana camara'' (common lantana) is a species of flowering plant within the verbena family ( Verbenaceae), native to the American tropics. It is a very adaptable species, which can inhabit a wide variety of ecosystems; once it has been introdu ...
''. He lamented that Indian universities forced those who took up biology to drop mathematics. He took an interest in the study of
floral symmetry Floral symmetry describes whether, and how, a flower, in particular its perianth, can be divided into two or more identical or mirror-image parts. Uncommonly, flowers may have no axis of symmetry at all, typically because their parts are spirall ...
. In January 1961 he befriended Canadian lepidopterist Gary Botting, the 1960 U.S. Science Fair winner in zoology (who had first visited the Haldanes along with Susan Brown, 1960 U.S. National Science Fair winner in botany), inviting him to share the results of his experiments hybridising '' Antheraea'' silk moths. He, his wife Helen Spurway and student Krishna Dronamraju were present at the Oberoi Grand Hotel in Kolkata when Brown reminded the Haldanes that she and Botting had a previously scheduled event that would prevent them from accepting an invitation to a banquet proposed by the Haldanes in their honour and had regretfully declined the honour. After the two students had left the hotel, Haldane went on his much-publicized hunger strike to protest what he regarded as a "U.S. insult." When the director of the ISI,
P. C. Mahalanobis Prasanta Chandra Mahalanobis OBE, FNA, FASc, FRS (29 June 1893– 28 June 1972) was an Indian scientist and statistician. He is best remembered for the Mahalanobis distance, a statistical measure, and for being one of the members of the first ...
, confronted Haldane about both the hunger strike and the unbudgeted banquet, Haldane resigned from his post (in February 1961), and moved to a newly established biometry unit in Bhubaneswar, the capital of Orissa (
Odisha Odisha (English: , ), formerly Orissa ( the official name until 2011), is an Indian state located in Eastern India. It is the 8th largest state by area, and the 11th largest by population. The state has the third largest population of ...
). Haldane took Indian citizenship; he was interested in
Hinduism Hinduism () is an Indian religion or '' dharma'', a religious and universal order or way of life by which followers abide. As a religion, it is the world's third-largest, with over 1.2–1.35 billion followers, or 15–16% of the global p ...
and became a
vegetarian Vegetarianism is the practice of abstaining from the consumption of meat (red meat, poultry, seafood, insects, and the flesh of any other animal). It may also include abstaining from eating all by-products of animal slaughter. Vegetariani ...
. In 1961, Haldane described India as "the closest approximation to the Free World."
Jerzy Neyman Jerzy Neyman (April 16, 1894 – August 5, 1981; born Jerzy Spława-Neyman; ) was a Polish mathematician and statistician who spent the first part of his professional career at various institutions in Warsaw, Poland and then at University Colleg ...
objected that "India has its fair share of scoundrels and a tremendous amount of poor unthinking and disgustingly subservient individuals who are not attractive." Haldane retorted:
Perhaps one is freer to be a scoundrel in India than elsewhere. So one was in the U.S.A in the days of people like
Jay Gould Jason Gould (; May 27, 1836 – December 2, 1892) was an American railroad magnate and financial speculator who is generally identified as one of the robber barons of the Gilded Age. His sharp and often unscrupulous business practices made him ...
, when (in my opinion) there was more internal freedom in the U.S.A than there is today. The "disgusting subservience" of the others has its limits. The people of Calcutta riot, upset trams, and refuse to obey police regulations, in a manner which would have delighted Jefferson. I don't think their activities are very efficient, but that is not the question at issue.
When on 25 June 1962 he was described in print as a " Citizen of the World" by
Groff Conklin Edward Groff Conklin (September 6, 1904 – July 19, 1968) was an American science fiction anthologist. He edited 40 anthologies of science fiction, one of mystery stories (co-edited with physician Noah Fabricant), wrote books on home improvemen ...
, Haldane responded:
No doubt I am in some sense a citizen of the world. But I believe with Thomas Jefferson that one of the chief duties of a citizen is to be a nuisance to the government of his state. As there is no world state, I cannot do this. On the other hand, I can be, and am, a nuisance to the government of India, which has the merit of permitting a good deal of criticism, though it reacts to it rather slowly. I also happen to be proud of being a citizen of India, which is a lot more diverse than Europe, let alone the U.S.A, the U.S.S.R or China, and thus a better model for a possible world organisation. It may of course break up, but it is a wonderful experiment. So, I want to be labeled as a citizen of India.


Personal life

Haldane was married twice, first to Charlotte Franken and then to
Helen Spurway Helen Spurway (12 June 1915 – 15 February 1978) was a British biologist and the second wife of J. B. S. Haldane. She emigrated to India in 1957 along with him and conducted research in field biology with Krishna Dronamraju, Suresh Jayakar, and o ...
. In 1924, Haldane met Charlotte Franken, who was a journalist for the ''
Daily Express The ''Daily Express'' is a national daily United Kingdom middle-market newspaper printed in tabloid format. Published in London, it is the flagship of Express Newspapers, owned by publisher Reach plc. It was first published as a broadsheet ...
'' and married to Jack Burghes. Following the publication of Haldane's ''Daedalus, or Science and the Future,'' she interviewed Haldane and they began a relationship. In order to marry Haldane, Franken filed a divorce suit, which resulted in controversy as Haldane was involved as co-respondent in the legal proceeding. Additionally, as
Sahotra Sarkar Sahotra Sarkar (born 1962) is an Indian-American professor at the University of Texas at Austin, specializing in the philosophy of biology. Education and career Sarkar is originally from India where he lived in Darjeeling until 1975. He earned ...
reported: "For her to secure a divorce, Haldane overtly committed adultery with her." Haldane's conduct was described as "gross immorality," for which he was formally dismissed by Cambridge's Sex Viri (a six-member disciplinary committee) from the university in 1925. Cambridge professors, including G. K. Chesterton,
Bertrand Russell Bertrand Arthur William Russell, 3rd Earl Russell, (18 May 1872 – 2 February 1970) was a British mathematician, philosopher, logician, and public intellectual. He had a considerable influence on mathematics, logic, set theory, linguistics, a ...
, and W. L. George, raised their defence for Haldane insisting that the university should not make such judgements, based solely on a professor's private life. The ouster was revoked in 1926. Haldane and Charlotte Franken were married in 1926. Following their separation in 1942, they divorced in 1945. Later that year he married
Helen Spurway Helen Spurway (12 June 1915 – 15 February 1978) was a British biologist and the second wife of J. B. S. Haldane. She emigrated to India in 1957 along with him and conducted research in field biology with Krishna Dronamraju, Suresh Jayakar, and o ...
, his former PhD student. Haldane once boasted about himself, saying, "I can read 11 languages and make public speeches in three; but am unmusical. I am a fairly competent public speaker." He had no children, but he and his father were important influences to his sister Naomi's children, of whom Denis Mitchison, Murdoch Mitchison and
Avrion Mitchison (Nicholas) Avrion Mitchison (born 5 May 1928) is a British zoologist and immunologist. Biography Mitchison was born in 1928, the son of the Labour politician Dick Mitchison (Baron Mitchison of Carradale in the County of Argyll, who died 197 ...
became professors of biology at the
University of London The University of London (UoL; abbreviated as Lond or more rarely Londin in post-nominals) is a federal public research university located in London, England, United Kingdom. The university was established by royal charter in 1836 as a degr ...
,
Edinburgh University The University of Edinburgh ( sco, University o Edinburgh, gd, Oilthigh Dhùn Èideann; abbreviated as ''Edin.'' in Post-nominal letters, post-nominals) is a Public university, public research university based in Edinburgh, Scotland. Granted ...
, and University College London, respectively. Inspired by his father, Haldane often used self-experimentation and would expose himself to danger to obtain data. To test the effects of acidification of the blood he drank dilute
hydrochloric acid Hydrochloric acid, also known as muriatic acid, is an aqueous solution of hydrogen chloride. It is a colorless solution with a distinctive pungent smell. It is classified as a strong acid. It is a component of the gastric acid in the dige ...
, enclosed himself in an airtight room containing 7% carbon dioxide, and found that it 'gives one a rather violent headache'. One experiment to study elevated levels of
oxygen saturation Oxygen saturation (symbol SO2) is a relative measure of the concentration of oxygen that is dissolved or carried in a given medium as a proportion of the maximal concentration that can be dissolved in that medium at the given temperature. It ca ...
triggered a fit which resulted in him suffering crushed
vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristi ...
. In his
decompression chamber A diving chamber is a vessel for human occupation, which may have an entrance that can be sealed to hold an internal pressure significantly higher than ambient pressure, a pressurised gas system to control the internal pressure, and a supply o ...
experiments, he and his volunteers suffered perforated eardrums. But, as Haldane stated in ''What is Life,'' "the drum generally heals up; and if a hole remains in it, although one is somewhat deaf, one can blow tobacco smoke out of the ear in question, which is a social accomplishment." Haldane made himself unpopular among his colleagues from the start of his academic career. In Cambridge, he annoyed most of the senior faculty due to his uninhibited behaviour, particularly at dinner. His partisan,
Edgar Adrian Edgar Douglas Adrian, 1st Baron Adrian (30 November 1889 – 4 August 1977) was an English electrophysiologist and recipient of the 1932 Nobel Prize for Physiology, won jointly with Sir Charles Sherrington for work on the function of neurons ...
(the 1932 Nobel laureate), had almost convinced the university to offer an appointment as Fellow of Trinity College, but that was ruined by an incident when Haldane arrived at the dining table, carrying a gallon jar of urine from his laboratory.


Later life and death

In the autumn of 1963, Haldane visited the US for a series of scientific conferences. At the
University of Wisconsin A university () is an institution of higher (or tertiary) education and research which awards academic degrees in several academic disciplines. Universities typically offer both undergraduate and postgraduate programs. In the United Stat ...
,
Sewall Wright Sewall Green Wright FRS(For) Honorary FRSE (December 21, 1889March 3, 1988) was an American geneticist known for his influential work on evolutionary theory and also for his work on path analysis. He was a founder of population genetics alongsi ...
introduced him before his speech, noting many of Haldane's achievements, after which Haldane modestly remarked that the introduction would have been more accurate if all the references to "Haldane" were replaced with "Wright". In Florida, he met, for the first and only time, the Russian biochemist
Alexander Oparin Alexander Ivanovich Oparin (russian: Александр Иванович Опарин; – April 21, 1980) was a Soviet biochemist notable for his theories about the origin of life, and for his book ''The Origin of Life''. He also studied the bi ...
, who had developed the origin of life theory quite independent of his own in the 1920s. It was while there that he started feeling abdominal pains. Haldane went to London for a diagnosis. He was found to have
colorectal cancer Colorectal cancer (CRC), also known as bowel cancer, colon cancer, or rectal cancer, is the development of cancer from the colon or rectum (parts of the large intestine). Signs and symptoms may include blood in the stool, a change in bowel ...
, and had a surgery in February 1964. Around that time Philip Dally was making a BBC documentary about eminent living scientists, which included Sewall Wright and the double Nobel laureate
Linus Pauling Linus Carl Pauling (; February 28, 1901August 19, 1994) was an American chemist, biochemist, chemical engineer, peace activist, author, and educator. He published more than 1,200 papers and books, of which about 850 dealt with scientific topi ...
. Dally's team approached Haldane at the hospital for the documentary profile, but instead of a filmed interview, Haldane gave them a self-obituary, the opening lines of which run:
I am going to begin with a boast. I believe that I am one of the riginally as "I am the most"most influential people living today, although I haven't got a scrap of power. Let me explain. In 1932 I was the first person to estimate the rate of mutation of a human gene.
He also wrote a comic poem while in the hospital, mocking his own incurable disease. It was read by his friends, who appreciated the consistent irreverence with which Haldane had lived his life. The poem first appeared in print on 21 February 1964 issue of the ''
New Statesman The ''New Statesman'' is a British political and cultural magazine published in London. Founded as a weekly review of politics and literature on 12 April 1913, it was at first connected with Sidney and Beatrice Webb and other leading members ...
'', and runs: The poem ends: He willed that his body be used for medical research and teaching at the
Rangaraya Medical College Rangaraya Medical College is a Government Medical College in Andhra Pradesh. It was established in 1958 and is in Kakinada, Andhra Pradesh, India. It is affiliated to Dr. NTR University of Health Sciences. History Dr. Datla Satyanarayana Ra ...
,
Kakinada Kakinada ( formerly called Kakinandiwada, Coringa, and Cocanada; ) is the sixth largest city of the Indian state of Andhra Pradesh and serves as the district headquarters of the Kakinada District. It lies on the coast of the Bay of Bengal. J. ...
. His surgery in London was declared successful. But the symptoms reappeared after returning to India in June, and in August, the Indian doctors confirmed that his condition was terminal. Writing to
John Maynard Smith John Maynard Smith (6 January 1920 – 19 April 2004) was a British theoretical and mathematical evolutionary biologist and geneticist. Originally an aeronautical engineer during the Second World War, he took a second degree in genetics un ...
on 7 September, he said, "I am not appreciably upset by the propect of dying fairly soon. But I am very angry
t the English doctor who performed the operation T, or t, is the twentieth letter in the Latin alphabet, used in the modern English alphabet, the alphabets of other western European languages and others worldwide. Its name in English is ''tee'' (pronounced ), plural ''tees''. It is der ...
" He died on 1 December 1964 in
Bhubaneswar Bhubaneswar (; ) is the capital and largest city of the Indian state of Odisha. The region, especially the old town, was historically often depicted as ''Ekamra Kshetra'' (area (''kshetra'') adorned with mango trees (''ekamra'')). Bhubaneswar i ...
. On that day the BBC broadcast his self-obituary as "Professor J.B.S. Haldane, obituary."


Scientific contributions

Following his father's footsteps, Haldane's first publication was on the mechanism of gaseous exchange by haemoglobin 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 emphasi ...
'', and he subsequently worked on the chemical properties of blood as a pH buffer. He investigated several aspects of
kidney function Assessment of kidney function occurs in different ways, using the presence of symptoms and signs, as well as measurements using urine tests, blood tests, and medical imaging. Functions of a healthy kidney include maintaining a person's fluid ...
s and mechanism of excretion.


Genetic linkage

In 1904, Arthur Dukinfield Darbishire published a paper on an experiment attempting to test
Mendelian inheritance Mendelian inheritance (also known as Mendelism) is a type of biological inheritance following the principles originally proposed by Gregor Mendel in 1865 and 1866, re-discovered in 1900 by Hugo de Vries and Carl Correns, and later popularize ...
between Japanese waltzing and albino mice. When Haldane came across the paper, he noticed that Darbishire had overlooked the possibility of genetic linkage in the experiment. Having sought advice from
Reginald Punnett Reginald Crundall Punnett FRS (; 20 June 1875 – 3 January 1967) was a British geneticist who co-founded, with William Bateson, the ''Journal of Genetics'' in 1910. Punnett is probably best remembered today as the creator of the Punnett ...
, a professor of biology at the University of Cambridge, he was ready to write a paper but only after an independent experiment. With his sister
Naomi Naomi or Naomie may refer to: People and biblical figures * Naomi (given name), a female given name and a list of people with the name * Naomi (biblical figure), Ruth's mother-in-law in the Old Testament Book of Ruth * Naomi (Romanian singer) (bor ...
and a friend Alexander Dalzell Sprunt, one year his senior, he started the experiment in 1908 using
guinea pigs The guinea pig or domestic guinea pig (''Cavia porcellus''), also known as the cavy or domestic cavy (), is a species of rodent belonging to the genus '' Cavia'' in the family Caviidae. Breeders tend to use the word ''cavy'' to describe the a ...
and
mice A mouse ( : mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus'' ...
. By 1912, the report was ready. But the paper titled ''Reduplication in mice'' was published in the '' Journal of Genetics'' only in December 1915 that became the first demonstration of
genetic linkage Genetic linkage is the tendency of DNA sequences that are close together on a chromosome to be inherited together during the meiosis phase of sexual reproduction. Two genetic markers that are physically near to each other are unlikely to be sep ...
in
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s, showing that certain genetic traits tend to be inherited together (as was later discovered, because of their proximity on chromosomes). (Between 1912 and 1914, genetic linkage had been reported in the fruit fly '' Drosophilla,''
silk moth The domestic silk moth (''Bombyx mori''), is an insect from the moth family Bombycidae. It is the closest relative of ''Bombyx mandarina'', the wild silk moth. The silkworm is the larva or caterpillar of a silk moth. It is an economically im ...
, and plants.) As the paper was written during Haldane's service during World War I, James F. Crow called it "the most important science article ever written in a front-line trench." Haldane himself recalled that he was the "only officer to complete a scientific paper from a forward position of the Black Watch." As was Haldane, Sprunt had joined 4th Battalion
Bedfordshire Regiment The Bedfordshire and Hertfordshire Regiment was the final title of a line infantry regiment of the British Army that was originally formed in 1688. After centuries of service in many conflicts and wars, including both the First and Second World Wa ...
at the start of World War I, and was killed at the
Battle of Neuve Chapelle The Battle of Neuve Chapelle (10–13 March 1915) took place in the First World War in the Artois region of France. The attack was intended to cause a rupture in the German lines, which would then be exploited with a rush to the Aubers Ridge a ...
on 17 March 1915. It was upon this news that Haldane submitted the paper for publication, in which he remarked: "Owing to the war it has been necessary to publish prematurely, as unfortunately one of us (A. D. S.) has already been killed in France." He was also the first to demonstrate linkage in
chicken The chicken (''Gallus gallus domesticus'') is a domesticated junglefowl species, with attributes of wild species such as the grey and the Ceylon junglefowl that are originally from Southeastern Asia. Rooster or cock is a term for an adu ...
s in 1921, and (with Julia Bell) in humans in 1937.


Enzyme kinetics

In 1925, with G.E. Briggs, Haldane derived a new interpretation of the
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products ...
kinetic law of Victor Henri in 1903, better known as the 1913 Michaelis–Menten equation.
Leonor Michaelis Leonor Michaelis (16 January 1875 – 8 October 1949) was a German biochemist, physical chemist, and physician, known for his work with Maud Menten on enzyme kinetics in 1913, as well as for work on enzyme inhibition, pH and quinones. Ear ...
and Maud Menten assumed that enzyme (catalyst) and substrate (reactant) are in fast equilibrium with their complex, which then dissociates to yield product and free enzyme. By contrast, at almost the same time, Donald Van Slyke and G. E. Cullen treated the binding step as an irreversible reaction. The Briggs–Haldane equation was of the same algebraic form as both of the earlier equations, but their derivation is based on the quasi-
steady state In systems theory, a system or a process is in a steady state if the variables (called state variables) which define the behavior of the system or the process are unchanging in time. In continuous time, this means that for those properties ''p' ...
approximation, which is the concentration of intermediate complex (or complexes) does not change. As a result, the microscopic meaning of the "Michaelis Constant" (''Km'') is different. Although commonly referring to it as Michaelis–Menten kinetics, most of the current models typically use the Briggs–Haldane derivation.


Haldane's principle

In his essay '' On Being the Right Size'' he outlines Haldane's principle, which states that the size very often defines what bodily equipment an animal must have: "Insects, being so small, do not have oxygen-carrying bloodstreams. What little oxygen their cells require can be absorbed by simple diffusion of air through their bodies. But being larger means an animal must have complicated oxygen pumping and distributing systems to reach all the cells."


Haldane's sieve

In 1927 Haldane pointed out that because selection mainly acts on heterozygotes, newly arisen dominant mutations are much more likely to be fixed than recessive ones, a mechanism now called '' Haldane's sieve''. This leads to the expectation that adaptation from new mutations in large outcrossing populations should primarily proceed via fixing non-recessive beneficial mutations.


Origin of life

Haldane introduced the modern concept of
abiogenesis In biology, abiogenesis (from a- 'not' + Greek bios 'life' + genesis 'origin') or the origin of life is the natural process by which life has arisen from non-living matter, such as simple organic compounds. The prevailing scientific hypothes ...
in an eight-page article titled ''The origin of life,'' in ''The'' ''Rationalist Annual'' in 1929, describing the primitive ocean as a "vast chemical laboratory" containing a mixture of inorganic compounds – like a "hot dilute soup" in which organic compounds could have formed. Under the solar energy the anoxic atmosphere containing
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is t ...
,
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous ...
and
water vapour (99.9839 °C) , - , Boiling point , , - , specific gas constant , 461.5 J/( kg·K) , - , Heat of vaporization , 2.27 MJ/kg , - , Heat capacity , 1.864 kJ/(kg·K) Water vapor, water vapour or aqueous vapor is the gaseous pha ...
gave rise to a variety of organic compounds, "living or half-living things". The first molecules reacted with one another to produce more complex compounds, and ultimately the cellular components. At some point a kind of "oily film" was produced that enclosed
self-replicating Self-replication is any behavior of a dynamical system that yields construction of an identical or similar copy of itself. Biological cells, given suitable environments, reproduce by cell division. During cell division, DNA is replicated and ca ...
nucleic acids, thereby becoming the first cell.
J. D. Bernal John Desmond Bernal (; 10 May 1901 – 15 September 1971) was an Irish scientist who pioneered the use of X-ray crystallography in molecular biology. He published extensively on the history of science. In addition, Bernal wrote popular book ...
named the hypothesis ''biopoiesis'' or ''biopoesis'', the process of living matter spontaneously evolving from self-replicating but lifeless molecules. Haldane further hypothesised that viruses were the intermediate entities between the prebiotic soup and the first cells. He asserted that prebiotic life would have been "in the virus stage for many millions of years before a suitable assemblage of elementary units was brought together in the first cell." The idea was generally dismissed as "wild speculation".
Alexander Oparin Alexander Ivanovich Oparin (russian: Александр Иванович Опарин; – April 21, 1980) was a Soviet biochemist notable for his theories about the origin of life, and for his book ''The Origin of Life''. He also studied the bi ...
had suggested a similar idea in Russian in 1924 (published in English in 1936). The hypothesis gained some empirical support in 1953 with the classic
Miller–Urey experiment The Miller–Urey experiment (or Miller experiment) is a famous chemistry experiment that simulated the conditions thought at the time (1952) to be present in the atmosphere of the early, prebiotic Earth, in order to test the hypothesis of the ...
. Since then, the primordial soup theory (Oparin–Haldane hypothesis) has become the foundation in the study of abiogenesis. Although Oparin's theory became widely known only after the English version in 1936, Haldane accepted Oparin's originality and said, "I have very little doubt that Professor Oparin has the priority over me."


Malaria and sickle-cell anemia

Haldane was the first to realise the evolutionary link between genetic disorder and infection in humans. While estimating the rates of human mutation in different situations and diseases, he noted that mutations expressed in red blood cells, like
thalassemia Thalassemias are inherited blood disorders characterized by decreased hemoglobin production. Symptoms depend on the type and can vary from none to severe. Often there is mild to severe anemia (low red blood cells or hemoglobin). Anemia can resul ...
s, were prevalent only in tropical regions where deadly infection like malaria has been
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...
. He further observed that these were favourable traits (heterozygous inheritance of sickle cell trait) for natural selection which protected individuals from receiving malarial infection. He introduced his hypothesis at the Eighth International Congress of Genetics held in 1948 at Stockholm on a topic "The Rate of Mutation of Human Genes". He proposed that genetic disorders in humans living in
malaria Malaria is a mosquito-borne infectious disease that affects humans and other animals. Malaria causes symptoms that typically include fever, tiredness, vomiting, and headaches. In severe cases, it can cause jaundice, seizures, coma, or death. ...
-endemic regions provided a condition (
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology (biology), morphology or physical form and structure, its Developmental biology, developmental proc ...
) that makes them relatively immune to malarial infections. He formalised in a technical paper published in 1949 in which he made a prophetic statement: "The corpuscles of the anaemic heterozygotes are smaller than normal, and more resistant to hypotonic solutions. It is at least conceivable that they are also more resistant to attacks by the sporozoa which cause malaria." This became known as "Haldane's malaria hypothesis", or concisely, the "malaria hypothesis". This hypothesis was eventually confirmed by Anthony C. Allison in 1954 in the case of
sickle-cell anemia Sickle cell disease (SCD) is a group of blood disorders typically inherited from a person's parents. The most common type is known as sickle cell anaemia. It results in an abnormality in the oxygen-carrying protein haemoglobin found in red bl ...
.


Population genetics

Haldane was one of the three major figures to develop the mathematical theory of
population genetics Population genetics is a subfield of genetics that deals with genetic differences within and between populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as adaptation, speciation, and po ...
, along with
Ronald Fisher Sir Ronald Aylmer Fisher (17 February 1890 – 29 July 1962) was a British polymath who was active as a mathematician, statistician, biologist, geneticist, and academic. For his work in statistics, he has been described as "a genius who ...
and
Sewall Wright Sewall Green Wright FRS(For) Honorary FRSE (December 21, 1889March 3, 1988) was an American geneticist known for his influential work on evolutionary theory and also for his work on path analysis. He was a founder of population genetics alongsi ...
. He thus played an important role in the modern evolutionary synthesis of the early 20th century. He re-established
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
as the central mechanism of
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
by explaining it as a mathematical consequence of
Mendelian inheritance Mendelian inheritance (also known as Mendelism) is a type of biological inheritance following the principles originally proposed by Gregor Mendel in 1865 and 1866, re-discovered in 1900 by Hugo de Vries and Carl Correns, and later popularize ...
. He wrote a series of ten papers, A Mathematical Theory of Natural and Artificial Selection, deriving expressions for the direction and rate of change of gene frequencies, and also analyzing the interaction of natural selection with
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, m ...
and
migration Migration, migratory, or migrate may refer to: Human migration * Human migration, physical movement by humans from one region to another ** International migration, when peoples cross state boundaries and stay in the host state for some minimum le ...
. The series consists of ten papers published between 1924 and 1934 in journals such as ''
Biological Reviews The Cambridge Philosophical Society (CPS) is a scientific society at the University of Cambridge. It was founded in 1819. The name derives from the medieval use of the word philosophy to denote any research undertaken outside the fields of l ...
'' (part II), ''
Mathematical Proceedings of the Cambridge Philosophical Society ''Mathematical Proceedings of the Cambridge Philosophical Society'' is a mathematical journal published by Cambridge University Press for the Cambridge Philosophical Society. It aims to publish original research papers from a wide range of pure ...
'' (parts I and from III to IX) and ''
Genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar work ...
'' (part X). He gave a set of lectures based on this series at the
University of Wales , latin_name = , image = , caption = Coat of Arms , motto = cy, Goreu Awen Gwirionedd , mottoeng = The Best Inspiration is Truth , established = , , type = Confederal, non-member ...
in 1931, and were summarised in a book, '' The Causes of Evolution'' in 1932. His first paper on the series in 1924 specifically treats the rate of natural selection in
peppered moth evolution The evolution of the peppered moth is an evolutionary instance of directional colour change in the moth population as a consequence of air pollution during the Industrial Revolution. The frequency of dark-coloured moths increased at that time ...
. He predicted that environmental condition can favour the increase or decline of either the dominant (in this case the black or melanic forms) or the recessive (the grey or
wild type The wild type (WT) is the phenotype of the typical form of a species as it occurs in nature. Originally, the wild type was conceptualized as a product of the standard "normal" allele at a locus, in contrast to that produced by a non-standard, "m ...
) moths. For a sooty environment such as Manchester, where the phenomenon was discovered in 1848, he predicted that the dominant melanic moths will have fifty times more survival fitness than the typical grey ones. According to his estimate, assuming 1% dominant form in 1848 and about 99% in 1898, "48 generations are needed for the change
or the dominant to appear Or or OR may refer to: Arts and entertainment Film and television * "O.R.", a 1974 episode of M*A*S*H * Or (My Treasure), a 2004 movie from Israel (''Or'' means "light" in Hebrew) Music * ''Or'' (album), a 2002 album by Golden Boy with Mis ...
.. After only 13 generations the dominants would be in a majority." Such mathematical prediction was considered improbable for natural selection in nature. But it was subsequently proven by an elaborate experiment (named
Kettlewell's experiment Kettlewell's experiment was a biological experiment in the mid-1950s to study the evolutionary mechanism of industrial melanism in the peppered moth (''Biston betularia''). It was executed by Bernard Kettlewell, working as a research fellow in th ...
) performed by an Oxford zoologist
Bernard Kettlewell Henry Bernard Davis Kettlewell (24 February 1907 – 11 May 1979) was a British geneticist, lepidopterist and medical doctor, who performed research on the influence of industrial melanism on peppered moth (''Biston betularia'') coloration, sho ...
between 1953 and 1958, and further by a Cambridge geneticists
Michael Majerus Michael Eugene Nicolas Majerus (13 February 1954 – 27 January 2009) was a British geneticist and professor of evolution at the University of Cambridge. He was also a teaching fellow at Clare College, Cambridge. He was an enthusiast in Darwin' ...
in his experiments conducted between 2001 and 2007. His contributions to statistical human genetics included: the first methods using
maximum likelihood In statistics, maximum likelihood estimation (MLE) is a method of estimating the parameters of an assumed probability distribution, given some observed data. This is achieved by maximizing a likelihood function so that, under the assumed stat ...
for the estimation of human linkage maps; pioneering methods for estimating human mutation rates; the first estimates of
mutation rate In genetics, the mutation rate is the frequency of new mutations in a single gene or organism over time. Mutation rates are not constant and are not limited to a single type of mutation; there are many different types of mutations. Mutation rates ...
in humans (2 × 10−5 mutations per gene per generation for the X-linked
haemophilia Haemophilia, or hemophilia (), is a mostly inherited genetic disorder that impairs the body's ability to make blood clots, a process needed to stop bleeding. This results in people bleeding for a longer time after an injury, easy bruisin ...
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a b ...
); and the first notion that there is a "cost of natural selection". He was the first to estimate the rate of human mutation in his 1932 book ''The Causes of Evolution.'' At the
John Innes Horticultural Institution The John Innes Centre (JIC), located in Norwich, Norfolk, England, is an independent centre for research and training in plant and microbial science founded in 1910. It is a registered charity (No 223852) grant-aided by the Biotechnology and B ...
, he developed the complicated linkage theory for polyploids; and extended the idea of gene/enzyme relationships with the biochemical and genetic study of plant pigments.


Political views

Haldane became a
socialist Socialism is a left-wing economic philosophy and movement encompassing a range of economic systems characterized by the dominance of social ownership of the means of production as opposed to private ownership. As a term, it describes the ...
during the First World War, supported the
Second Spanish Republic The Spanish Republic (), commonly known as the Second Spanish Republic (), was the form of government in Spain from 1931 to 1939. The Republic was proclaimed on 14 April 1931, after the deposition of King Alfonso XIII, and was dissolved on 1 ...
during the
Spanish Civil War The Spanish Civil War ( es, Guerra Civil Española)) or The Revolution ( es, La Revolución, link=no) among Nationalists, the Fourth Carlist War ( es, Cuarta Guerra Carlista, link=no) among Carlism, Carlists, and The Rebellion ( es, La Rebeli ...
, and then became an open supporter of the
Communist Party of Great Britain The Communist Party of Great Britain (CPGB) was the largest communist organisation in Britain and was founded in 1920 through a merger of several smaller Marxist groups. Many miners joined the CPGB in the 1926 general strike. In 1930, the CPGB ...
in 1937. A pragmatic dialectical-materialist
Marxist Marxism is a left-wing to far-left method of socioeconomic analysis that uses a materialist interpretation of historical development, better known as historical materialism, to understand class relations and social conflict and a dialecti ...
, he wrote many articles for the ''
Daily Worker The ''Daily Worker'' was a newspaper published in New York City by the Communist Party USA, a formerly Comintern-affiliated organization. Publication began in 1924. While it generally reflected the prevailing views of the party, attempts were m ...
''. In '' On Being the Right Size'', he wrote that "while
nationalization Nationalization (nationalisation in British English) is the process of transforming privately-owned assets into public assets by bringing them under the public ownership of a national government or state. Nationalization usually refers to p ...
of certain industries is an obvious possibility in the largest of states, I find it no easier to picture a completely socialized British Empire or United States than an elephant turning
somersaults A somersault (also ''flip'', ''heli'', and in gymnastics ''salto'') is an acrobatic exercise in which a person's body rotates 360° around a horizontal axis with the feet passing over the head. A somersault can be performed forwards, backwards ...
or a hippopotamus jumping a hedge." In 1938, Haldane proclaimed enthusiastically: "I think that Marxism is true." He joined the Communist Party in 1942. He was pressed to speak out about the rise of
Lysenkoism Lysenkoism (russian: Лысенковщина, Lysenkovshchina, ; uk, лисенківщина, lysenkivščyna, ) was a political campaign led by Soviet biologist Trofim Lysenko against genetics and science-based agriculture in the mid-20th ce ...
and the persecution of geneticists in the Soviet Union as anti-Darwinist and the political suppression of genetics as incompatible with
dialectical materialism Dialectical materialism is a philosophy of science, history, and nature developed in Europe and based on the writings of Karl Marx and Friedrich Engels. Marxist dialectics, as a materialist philosophy, emphasizes the importance of real-world co ...
. He shifted his polemic focus to the United Kingdom, criticizing the dependence of scientific research on financial patronage. In 1941, he wrote about the Soviet trial of his friend and fellow geneticist
Nikolai Vavilov Nikolai Ivanovich Vavilov ( rus, Никола́й Ива́нович Вави́лов, p=nʲɪkɐˈlaj ɪˈvanəvʲɪtɕ vɐˈvʲiləf, a=Ru-Nikolay_Ivanovich_Vavilov.ogg; – 26 January 1943) was a Russian and Soviet agronomist, botanist ...
: By the end of the
Second World War 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 opposi ...
, Haldane had become an explicit critic of the regime. He left the party in 1950, shortly after considering standing for Parliament as a Communist Party candidate. He continued to admire
Joseph Stalin Joseph Vissarionovich Stalin (born Ioseb Besarionis dze Jughashvili; – 5 March 1953) was a Georgian revolutionary and Soviet Union, Soviet political leader who led the Soviet Union from 1924 until his death in 1953. He held power as Ge ...
, describing him in 1962 as "a very great man who did a very good job". Haldane has been accused by authors including Peter Wright and Chapman Pincher of having been a Soviet
GRU The Main Directorate of the General Staff of the Armed Forces of the Russian Federation, rus, Гла́вное управле́ние Генера́льного шта́ба Вооружённых сил Росси́йской Федера́ци ...
spy codenamed ''Intelligentsia''.


Social and scientific views


Human cloning

Haldane was the first to have thought of the genetic basis for
human cloning Human cloning is the creation of a genetically identical copy (or clone) of a human. The term is generally used to refer to artificial human cloning, which is the reproduction of human cells and tissue. It does not refer to the natural concept ...
, and the eventual artificial breeding of superior individuals. For this he introduced the terms "clone" and "cloning", modifying the earlier "clon" which had been used in agriculture since the early 20th century (from Greek ''klōn'', twig). He introduced the term in his speech on "Biological Possibilities for the Human Species of the Next Ten Thousand Years" at the '' Ciba Foundation Symposium on Man and his Future'' in 1963. He said:


Ectogenesis and in vitro fertilisation

His essay '' Daedalus; or, Science and the Future'' (1924) posited the concept of
in vitro fertilisation In vitro fertilisation (IVF) is a process of fertilisation where an egg is combined with sperm in vitro ("in glass"). The process involves monitoring and stimulating an individual's ovulatory process, removing an ovum or ova (egg or eggs) ...
, which he called
ectogenesis Ectogenesis (from the Greek ''ἐκτός'', "outside," and '' genesis'') is the growth of an organism in an artificial environment outside the body in which it would normally be found, such as the growth of an embryo or fetus outside the mother's ...
. He envisioned ectogenesis as a tool for creating better individuals (eugenics). Haldane's work was an influence on Huxley's ''
Brave New World ''Brave New World'' is a dystopian novel by English author Aldous Huxley, written in 1931 and published in 1932. Largely set in a futuristic World State, whose citizens are environmentally engineered into an intelligence-based social hiera ...
'' (1932) and was also admired by Gerald Heard. Various essays on science were collected and published in a volume titled ''Possible Worlds'' in 1927. His book, ''A.R.P. (Air Raid Precautions)'' (1938) combined his physiological research into the effects of stress upon the human body with his experience of air raids during the
Spanish Civil War The Spanish Civil War ( es, Guerra Civil Española)) or The Revolution ( es, La Revolución, link=no) among Nationalists, the Fourth Carlist War ( es, Cuarta Guerra Carlista, link=no) among Carlism, Carlists, and The Rebellion ( es, La Rebeli ...
to provide a scientific account of the likely effects of the air raids that Britain was to endure during the
Second World War 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 opposi ...
.


Criticism of C. S. Lewis

Along with
Olaf Stapledon William Olaf Stapledon (10 May 1886 – 6 September 1950) – known as Olaf Stapledon – was a British philosopher and author of science fiction.Andy Sawyer, " illiamOlaf Stapledon (1886-1950)", in Bould, Mark, et al, eds. ''Fifty Key Figures ...
,
Charles Kay Ogden Charles Kay Ogden (; 1 June 1889 – 20 March 1957) was an English linguist, philosopher, and writer. Described as a polymath but also an eccentric and outsider, he took part in many ventures related to literature, politics, the arts, and philo ...
, I. A. Richards, and H. G. Wells, Haldane was accused by
C. S. Lewis Clive Staples Lewis (29 November 1898 – 22 November 1963) was a British writer and Anglican lay theologian. He held academic positions in English literature at both Oxford University (Magdalen College, 1925–1954) and Cambridge Univer ...
of
scientism Scientism is the opinion that science and the scientific method are the best or only way to render truth about the world and reality. While the term was defined originally to mean "methods and attitudes typical of or attributed to natural scientis ...
. Haldane criticised Lewis and his Ransom Trilogy for the "complete mischaracterisation of science, and his disparagement of the human race". Haldane wrote a book for children titled ''My Friend Mr Leakey'' (1937), containing the stories "A Meal With a Magician", "A Day in the Life of a Magician", "Mr Leakey's Party", "Rats", "The Snake with the Golden Teeth", and "My Magic Collar Stud"; later editions featured illustrations by
Quentin Blake Sir Quentin Saxby Blake, (born 16 December 1932) is an English cartoonist, caricaturist, illustrator and children's writer. He has illustrated over 300 books, including 18 written by Roald Dahl, which are among his most popular works. For his ...
. He also wrote an essay criticising Lewis's arguments for the existence of God, entitled "More Anti-Lewisite", a reference to the
poison gas Many gases have toxic properties, which are often assessed using the LC50 (median lethal dose) measure. In the United States, many of these gases have been assigned an NFPA 704 health rating of 4 (may be fatal) or 3 (may cause serious or perma ...
and its
antidote An antidote is a substance that can counteract a form of poisoning. The term ultimately derives from the Greek term φάρμακον ἀντίδοτον ''(pharmakon) antidoton'', "(medicine) given as a remedy". Antidotes for anticoagulants are s ...
.


Hydrogen-generating windmills

In 1923, in a talk given in Cambridge titled "Science and the Future", Haldane, foreseeing the exhaustion of
coal Coal is a combustible black or brownish-black sedimentary rock, formed as rock strata called coal seams. Coal is mostly carbon with variable amounts of other elements, chiefly hydrogen, sulfur, oxygen, and nitrogen. Coal is formed when ...
for power generation in Britain, proposed a network of
hydrogen Hydrogen is the chemical element with the symbol H and atomic number 1. Hydrogen is the lightest element. At standard conditions hydrogen is a gas of diatomic molecules having the formula . It is colorless, odorless, tasteless, non-to ...
-generating
windmill A windmill is a structure that converts wind power into rotational energy using vanes called sails or blades, specifically to mill grain (gristmills), but the term is also extended to windpumps, wind turbines, and other applications, in some ...
s. This is the first proposal of the hydrogen-based renewable energy economy.


Scientists

In his ''An Autobiography in Brief'', published shortly before his death in India, Haldane named four close associates as showing promise to become illustrious scientists: T. A. Davis,
Dronamraju Krishna Rao Dronamraju Krishna Rao (14 January 1937 – 3 December 2020) was an Indian-born geneticist and president of the Foundation for Genetic Research in Houston, Texas. He was born in Pithapuram, in the state of Andhra Pradesh, India. One focus of hi ...
,
Suresh Jayakar Suresh Dinakar Jayakar (21 September 1937, Bombay – 21 January 1988) was an Indian biologist who pioneered in the use of quantitative approaches in genetics and biology. He studied mathematical statistics, physics and mathematics at the Unive ...
and S. K. Roy.


Awards and honours

Haldane was elected a Fellow of the Royal Society in 1932. The
French Government The Government of France ( French: ''Gouvernement français''), officially the Government of the French Republic (''Gouvernement de la République française'' ), exercises executive power in France. It is composed of the Prime Minister, who ...
conferred him its
National Order of the Legion of Honour The National Order of the Legion of Honour (french: Ordre national de la Légion d'honneur), formerly the Royal Order of the Legion of Honour ('), is the highest French order of merit, both military and civil. Established in 1802 by Napoleon B ...
in 1937. In 1952, he received the
Darwin Medal The Darwin Medal is one of the medals awarded by the Royal Society for "distinction in evolution, biological diversity and developmental, population and organismal biology". In 1885, International Darwin Memorial Fund was transferred to the ...
from 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 ...
. In 1956, he was awarded the Huxley Memorial Medal of the
Anthropological Institute of Great Britain The Royal Anthropological Institute of Great Britain and Ireland (RAI) is a long-established anthropological organisation, and Learned Society, with a global membership. Its remit includes all the component fields of anthropology, such as biolo ...
. He received the
Feltrinelli Prize The Feltrinelli Prize (from the Italian "Premio Feltrinelli", also known as "International Feltrinelli Prize" or "Antonio Feltrinelli Prize") is an award for achievement in the arts, music, literature, history, philosophy, medicine, and physical and ...
from
Accademia Nazionale dei Lincei The Accademia dei Lincei (; literally the "Academy of the Lynx-Eyed", but anglicised as the Lincean Academy) is one of the oldest and most prestigious European scientific institutions, located at the Palazzo Corsini on the Via della Lungara in Rom ...
in 1961. He also received an Honorary Doctorate of Science, an Honorary Fellowship at New College, Oxford, and the Kimber Award of the US National Academy of Sciences. He was awarded the
Linnean Society of London The Linnean Society of London is a learned society dedicated to the study and dissemination of information concerning natural history, evolution, and taxonomy. It possesses several important biological specimen, manuscript and literature coll ...
's prestigious
Darwin–Wallace Medal The Darwin–Wallace Medal is a medal awarded by the Linnean Society of London for "major advances in evolutionary biology". Historically, the medals have been awarded every 50 years, beginning in 1908. That year marked 50 years after the joint p ...
in 1958.


Legacy

The Haldane Lecture at the
John Innes Centre The John Innes Centre (JIC), located in Norwich, Norfolk, England, is an independent centre for research and training in plant and microbial science founded in 1910. It is a registered charity (No 223852) grant-aided by the Biotechnology and ...
, where Haldane worked from 1927 to 1937 is named in his honour. The JBS Haldane Lecture of
The Genetics Society The Genetics Society is a British learned society. It was founded by William Bateson and Edith Rebecca Saunders in 1919 and celebrated its centenary year in 2019. It is therefore one of the oldest learned societies devoted to genetics. Its membe ...
is also named in his honour. Haldane was parodied as an obsessive self-experimenter, described as "the biologist too absorbed in his experiments to notice his friends bedding his wife" by his friend
Aldous Huxley Aldous Leonard Huxley (26 July 1894 – 22 November 1963) was an English writer and philosopher. He wrote nearly 50 books, both novels and non-fiction works, as well as wide-ranging essays, narratives, and poems. Born into the prominent Huxle ...
in the novel ''
Antic Hay ''Antic Hay'' is a comic novel by Aldous Huxley, published in 1923. The story takes place in London, and depicts the aimless or self-absorbed cultural elite in the sad and turbulent times following the end of World War I. The book follows the ...
'' (1923).


Quotations

* He is famous for the (possibly
apocryphal Apocrypha are works, usually written, of unknown authorship or of doubtful origin. The word ''apocryphal'' (ἀπόκρυφος) was first applied to writings which were kept secret because they were the vehicles of esoteric knowledge considered ...
) response that he gave when some theologians asked him what could be inferred about the mind of the Creator from the works of His Creation: "An inordinate fondness for beetles." or sometimes, "....stars and beetles." * "My own suspicion is that the universe is not only queerer than we suppose, but queerer than we ''can'' suppose." Emphasis in the original. * "It seems to me immensely unlikely that mind is a mere by-product of matter. For if my mental processes are determined wholly by the motions of atoms in my brain I have no reason to suppose that my beliefs are true. They may be sound chemically, but that does not make them sound logically. And hence I have no reason for supposing my brain to be composed of atoms." * "
Teleology Teleology (from and )Partridge, Eric. 1977''Origins: A Short Etymological Dictionary of Modern English'' London: Routledge, p. 4187. or finalityDubray, Charles. 2020 912Teleology" In ''The Catholic Encyclopedia'' 14. New York: Robert Appleton ...
is like a mistress to a biologist: he cannot live without her but he's unwilling to be seen with her in public." * "I had
gastritis Gastritis is inflammation of the lining of the stomach. It may occur as a short episode or may be of a long duration. There may be no symptoms but, when symptoms are present, the most common is upper abdominal pain (see dyspepsia). Other possi ...
for about fifteen years until I read
Lenin Vladimir Ilyich Ulyanov. ( 1870 – 21 January 1924), better known as Vladimir Lenin,. was a Russian revolutionary, politician, and political theorist. He served as the first and founding head of government of Soviet Russia from 1917 to 1 ...
and other writers, who showed me what was wrong with our society and how to cure it. Since then I have needed no magnesia." * "I suppose the process of acceptance will pass through the usual four stages: (i) This is worthless nonsense; (ii) This is an interesting, but perverse, point of view; (iii) This is true, but quite unimportant; (iv) I always said so." * "Three hundred and ten species in all of India, representing two hundred and thirty-eight genera, sixty-two families, nineteen different orders. All of them on the
Ark Ark or ARK may refer to: Biblical narratives and religion Hebrew word ''teva'' * Noah's Ark, a massive vessel said to have been built to save the world's animals from a flood * Ark of bulrushes, the boat of the infant Moses Hebrew ''aron'' * ...
. And this is only India, and only the ''birds''." * "The stupidity of the '' mynah'' shows that in birds, as in men, linguistic and practical abilities are not very highly correlated. A student who can repeat a page of a text book may get first class honours, but may be incapable of doing research." * When asked whether he would lay down his life for his brother, Haldane, presaging Hamilton's rule, supposedly replied "two brothers or eight cousins".


Publications

* '' Daedalus; or, Science and the Future'' (1924), E.P. Dutton and Company, Inc., a paper read to the Heretics, Cambridge, on 4 February 1923 ** second edition (1928), London: Kegan Paul, Trench & Co. ** see also ''Haldane's'' Daedalus ''Revisited'' (1995), ed. with an introd. by Krishna R. Dronamraju, Foreword by Joshua Lederberg; with essays by
M. F. Perutz Max Ferdinand Perutz (19 May 1914 – 6 February 2002) was an Austrian-born British molecular biologist, who shared the 1962 Nobel Prize for Chemistry with John Kendrew, for their studies of the structures of haemoglobin and myoglobin. He went ...
,
Freeman Dyson Freeman John Dyson (15 December 1923 – 28 February 2020) was an English-American theoretical physicist and mathematician known for his works in quantum field theory, astrophysics, random matrices, mathematical formulation of quantum m ...
, Yaron Ezrahi, Ernst Mayr, Elof Axel Carlson, D. J. Weatherall, N. A. Mitchison and the editor. Oxford University Press. * A Mathematical Theory of Natural and Artificial Selection, a series of papers beginning in 1924 * (With G.E. Briggs)
''Callinicus: A Defence of Chemical Warfare''
(1925), E. P. Dutton
''Possible Worlds and Other Essays''
(1927), Chatto & Windus; 2001 reprint, Transaction Publishers: (includes " On Being the Right Size" and
On Being One's Own Rabbit
) *The Last Judgment, an essay sequel to ''Daedalus'' (1927)''.''
''Possible Worlds and other Essays''
(1927), London: Chatto and Windus. * ''On Being the Right Size'' (1929)
"The origin of life"
in the ''Rationalist Annual'' (1929) * ''Animal Biology'' (1929) Oxford: Clarendon
''The Sciences and Philosophy''
(1929) NY: Doubleday, Doran and Company. By John Scott Haldane, JBS Haldane's father. * ''Enzymes'' (1930), MIT Press 1965 edition with new preface by the author written just prior to his death: *
''The Inequality of Man, and Other Essays''
(1932)

London: Longmans, Green, 1932. * ''Science and Human Life'' (1933), Harper and Brothers, Ayer Co. reprint: * ''Science and the Supernatural: Correspondence with Arnold Lunn'' (1935), Sheed & Ward, Inc, * ''Fact and Faith'' (1934), Watts Thinker's Library * ''Human Biology and Politics'' (1934) * "A Contribution to the Theory of Price Fluctuations", ''The Review of Economic Studies'', 1:3, 186–195 (1934).
''My Friend Mr Leakey''
(1937), Jane Nissen Books reprint (2004): * "A Dialectical Account of Evolution" in ''Science & Society'' Volume I (1937) * * (with Julia Bell) * (with C.A.B. Smith) * ''Air Raid Precautions (A.R.P.)'' (1938), Victor Gollancz
''Heredity and Politics''
(1938), Allen and Unwin. * "Reply to A.P. Lerner's Is Professor Haldane's Account of Evolution Dialectical?" in ''Science & Society'' volume 2 (1938)
''The Marxist Philosophy and the Sciences''
(1939), Random House, Ayer Co. reprint: * Preface to Engels' ''Dialectics of Nature'' (1939) * ''Science and Everyday Life'' (1940), Macmillan, 1941 Penguin, Ayer Co. 1975 reprint: * "Lysenko and Genetics" in ''Science & Society'' volume 4 (1940) * "Why I am a Materialist" in ''Rationalist Annual'' (1940) * "The Laws of Nature" in ''Rationalist Annual'' (1940) * ''Science in Peace and War'' (1941), Lawrence & Wishart Ltd
''New Paths in Genetics''
(1941), George Allen & Unwin
''Heredity & Politics''
(1943), George Allen & Unwin * ''Why Professional Workers should be Communists'' (1945), London: Communist Party (of Great Britain) In this four page pamphlet, Haldane contends that Communism should appeal to professionals because Marxism is based on the scientific method and Communists hold scientists as important; Haldane subsequently disavowed this position. * ''Adventures of a Biologist'' (1947) * ''Science Advances'' (1947), Macmillan
''What is Life?''
(1947), Boni and Gaer, 1949 edition: Lindsay Drummond * ''Everything Has a History'' (1951), Allen & Unwin—Includes "Auld Hornie, F.R.S."; C.S. Lewis's "Reply to Professor Haldane" is available in "On Stories and Other Essays on Literature," ed. Walter Hooper (1982), . * "The Origins of Life", ''New Biology'', 16, 12–27 (1954). Suggests that an alternative biochemistry could be based on liquid ammonia. * ''The Biochemistry of Genetics'' (1954) * * * * "Cancer's a Funny Thing", in ''New Statesman'', 21 February 1964.


See also

* ''
Experiments in the Revival of Organisms ''Experiments in the Revival of Organisms'' (russian: О́пыты по оживле́нию органи́зма) is a 1940 motion picture, directed by David Yashin, that documents Soviet research into the resuscitation of clinically dead organ ...
'', a 1940 Soviet film featuring Haldane in the introduction. * List of independent discoveries (" primordial soup" theory of the evolution of life from carbon-based molecules, ) * Precambrian rabbit * Timeline of hydrogen technologies


References


Citations


Further reading

* Clark, Ronald (1968). ''JBS: The Life and Work of J.B.S. Haldane''. Coward-McCann. * * * Foreword by Naomi Mitchison. * * * *
Subramanian Subramaniam, Subrahmanyam, Subramanyam or Subramanian ( ta, சுப்பிரமணியம்; te, సుబ్రహ్మణ్యం) is a South Indian male given name. Due to the South Indian tradition of using patronymic surnames it ...
, Samanth (2019). '' A Dominant Character: The Radical Science and Restless Politics of J. B. S. Haldane.'' Simon and Schuster. . *


External links

* *
Possible worlds, and other essays
' at
Toronto Public Library Toronto Public Library (TPL) (french: Bibliothèque publique de Toronto) is a public library system in Toronto, Ontario. It is the largest public library system in Canada, and in 2008 had averaged a higher circulation per capita than any other pub ...

Facsimiles of Haldane's books and some of his scientific papers, with photographs, a detailed bibliography of his publications and other materials



A review (from a modern perspective) of ''The Causes of Evolution''


* ttps://web.archive.org/web/20070615001014/http://www.iisc.ernet.in/academy/resonance/Dec1998/pdf/Dec1998p32-35.pdf Haldane's contributions to science in India
Marxist Writers: J.B.S. Haldane
* Th

on the Marxist Writers page has a photograph of Haldane when he was younger.

* ttp://wellcomelibrary.org/using-the-library/subject-guides/genetics/makers-of-modern-genetics/digitised-archives/j-b-s-haldane/ Codebreakers: Makers of Modern Genetics: the J B S Haldane papers
Haldane: a cantankerous and charismatic pioneer
{{DEFAULTSORT:Haldane, J. B. S. 1892 births 1964 deaths 20th-century British writers 20th-century Indian biologists 20th-century Indian chemists 20th-century Indian non-fiction writers Academics of University College London Black Watch officers British Army personnel of World War I British evolutionary biologists Communist Party of Great Britain members Critics of Lamarckism Deaths from colorectal cancer English atheists English biochemists English communists English emigrants to India English geneticists English people of Scottish descent English physiologists English political writers Fellows of New College, Oxford Fellows of the Royal Society Fellows of Trinity College, Cambridge Foreign associates of the National Academy of Sciences Foreign Members of the USSR Academy of Sciences Fullerian Professors of Physiology J. B. S Honorary Members of the USSR Academy of Sciences Hunger strikers Indian atheists Indian biochemists Indian communists Indian geneticists Indian Marxist writers Indian physiologists Indian political writers Indian Statistical Institute faculty Indian technology writers Linnean Medallists Members of the German Academy of Sciences at Berlin Modern synthesis (20th century) People educated at Eton College People educated at The Dragon School People from Oxford People who lost British citizenship People with acquired Indian citizenship Population geneticists