George R. Price
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George Robert Price (October 6, 1922 – January 6, 1975) was an American
population geneticist 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 popu ...
. Price is often noted for his formulation of the
Price equation In the theory of evolution and natural selection, the Price equation (also known as Price's equation or Price's theorem) describes how a trait or allele changes in frequency over time. The equation uses a covariance between a trait and fitness, ...
in 1967. Originally a
physical chemist Physical chemistry is the study of macroscopic and microscopic phenomena in chemical systems in terms of the principles, practices, and concepts of physics such as motion, energy, force, time, thermodynamics, quantum chemistry, statistical me ...
and later a
science journalist Science journalism conveys reporting about science to the public. The field typically involves interactions between scientists, journalists, and the public. Origins Modern science journalism dates back to '' Digdarshan'' (means showing the d ...
, he moved to
London London is the capital and largest city of England and the United Kingdom, with a population of just under 9 million. It stands on the River Thames in south-east England at the head of a estuary down to the North Sea, and has been a majo ...
in 1967, where he worked in
theoretical biology Mathematical and theoretical biology, or biomathematics, is a branch of biology which employs theoretical analysis, mathematical models and abstractions of the living organisms to investigate the principles that govern the structure, development a ...
at the
Galton Laboratory The Galton Laboratory was a laboratory for research into eugenics and then into human genetics based at University College London in London, England. It was originally established in 1904, and became part of UCL's biology department in 1996. The a ...
, making three important contributions: first, rederiving
W.D. Hamilton William Donald Hamilton (1 August 1936 – 7 March 2000) was a British evolutionary biologist, recognised as one of the most significant evolutionary theorists of the 20th century. Hamilton became known for his theoretical work expounding a ...
's work on
kin selection Kin selection is the evolutionary strategy that favours the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin altruism can look like Altruism in animals, altruistic behavio ...
with the new Price equation that vindicated
group selection Group selection is a proposed mechanism of evolution in which natural selection acts at the level of the group, instead of at the level of the individual or gene. Early authors such as V. C. Wynne-Edwards and Konrad Lorenz argued that the behav ...
; second, introducing (with
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 und ...
) the concept of the
evolutionarily stable strategy An evolutionarily stable strategy (ESS) is a strategy (or set of strategies) that is ''impermeable'' when adopted by a population in adaptation to a specific environment, that is to say it cannot be displaced by an alternative strategy (or set of ...
(ESS), a central concept in
game theory Game theory is the study of mathematical models of strategic interactions among rational agents. Myerson, Roger B. (1991). ''Game Theory: Analysis of Conflict,'' Harvard University Press, p.&nbs1 Chapter-preview links, ppvii–xi It has appli ...
; and third, formalizing Fisher's fundamental theorem of natural selection. Price converted to Christianity and gave all his possessions to the poor. Price grew depressed in the setting of thyroid disease and non-compliance with his medications, and committed
suicide Suicide is the act of intentionally causing one's own death. Mental disorders (including depression, bipolar disorder, schizophrenia, personality disorders, anxiety disorders), physical disorders (such as chronic fatigue syndrome), and s ...
.


Early life

Price was born in 1922 in the U.S. state of New York. His father, an electrician, died when Price was four. His mother was a former theater actress, and the family struggled through the
Great Depression The Great Depression (19291939) was an economic shock that impacted most countries across the world. It was a period of economic depression that became evident after a major fall in stock prices in the United States. The economic contagio ...
. Price attended
Birch Wathen School The Birch Wathen Lenox School is a college preparatory K-12 school on the Upper East Side of Manhattan in New York City. Birch Wathen Lenox comprises approximately 500 students from all around New York City. The Birch Wathen Lenox School is one o ...
followed by
Stuyvesant High School Stuyvesant High School (pronounced ), commonly referred to among its students as Stuy (pronounced ), is a State school, public university-preparatory school, college-preparatory, Specialized high schools in New York City, specialized high school ...
in
New York New York most commonly refers to: * New York City, the most populous city in the United States, located in the state of New York * New York (state), a state in the northeastern United States New York may also refer to: Film and television * '' ...
. He graduated with a degree in
chemistry Chemistry is the science, scientific study of the properties and behavior of matter. It is a natural science that covers the Chemical element, elements that make up matter to the chemical compound, compounds made of atoms, molecules and ions ...
from University of Chicago in 1943 and received his doctorate in the subject from the same institution in 1946. In 1947 he married Julia Madigan, but their relationship was contentious as George was a strong atheist while his wife was a practicing Roman Catholic. They divorced in 1955, having had two daughters, Annamarie and Kathleen.


Career


Early career

Price was a member of the Manhattan Project as a chemist looking into the characteristics of
plutonium-235 Plutonium (94Pu) is an artificial element, except for trace quantities resulting from neutron capture by uranium, and thus a standard atomic weight cannot be given. Like all artificial elements, it has no stable isotopes. It was synthesized long ...
. Between 1946 and 1948, he was an instructor in chemistry at Harvard University and consultant to
Argonne National Laboratory Argonne National Laboratory is a science and engineering research United States Department of Energy National Labs, national laboratory operated by University of Chicago, UChicago Argonne LLC for the United States Department of Energy. The facil ...
. Later, he took a position at Bell Laboratories to work on the chemistry of transistors. He then worked as a research associate in medicine at the University of Minnesota, working on, among other things, fluorescence microscopy and liver perfusion. In 1955 and 1956, he published two papers in the journal '' Science'' criticizing the
pseudoscientific Pseudoscience consists of statements, beliefs, or practices that claim to be both scientific and factual but are incompatible with the scientific method. Pseudoscience is often characterized by contradictory, exaggerated or unfalsifiable claim ...
claims of extrasensory perception. Continuing with science journalism, Price tried to write a book titled ''No Easy Way'' about the United States'
Cold War The Cold War is a term commonly used to refer to a period of geopolitical tension between the United States and the Soviet Union and their respective allies, the Western Bloc and the Eastern Bloc. The term '' cold war'' is used because the ...
with the Soviet Union and the People's Republic of China, but complained that "the world kept changing faster than I could write about it", and so the book was never finished. From 1961 to 1967, Price was employed by IBM as a consultant on graphic data processing. In 1966, he was treated for thyroid cancer, but the operation to remove the tumour left his shoulder partially paralysed and left him reliant on thyroxine medication. With the money from his medical insurance, he moved to the United Kingdom to start a new life in November 1967.


In Britain

W.D. Hamilton William Donald Hamilton (1 August 1936 – 7 March 2000) was a British evolutionary biologist, recognised as one of the most significant evolutionary theorists of the 20th century. Hamilton became known for his theoretical work expounding a ...
failed to recall when Price first contacted him, but says Price had read Hamilton's 1964 papers on
kin selection Kin selection is the evolutionary strategy that favours the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin altruism can look like Altruism in animals, altruistic behavio ...
, and with no training in population genetics or
statistics Statistics (from German language, German: ''wikt:Statistik#German, Statistik'', "description of a State (polity), state, a country") is the discipline that concerns the collection, organization, analysis, interpretation, and presentation of ...
devised the Price equation, a covariance equation that generated the change in allele frequency of a population. Although the first part of the equation had previously been derived by Alan Robertson and
C. C. Li Ching Chun Li (; October 27, 1912 – October 20, 2003) was a Chinese-American population geneticist and human geneticist. He was known for his research and the book ''An Introduction to Population Genetics''. Biography Ching Chun Li was bor ...
, its second component allowed it to be applied to all levels of selection, meiotic drive, traditional natural selection with an extension into
inclusive fitness In evolutionary biology, inclusive fitness is one of two metrics of evolutionary success as defined by W. D. Hamilton in 1964: * Personal fitness is the number of offspring that an individual begets (regardless of who rescues/rears/supports them ...
and
group selection Group selection is a proposed mechanism of evolution in which natural selection acts at the level of the group, instead of at the level of the individual or gene. Early authors such as V. C. Wynne-Edwards and Konrad Lorenz argued that the behav ...
.


Conversion to Christianity

On 6 June 1970, Oren Harman (2010) ''The Price of Altruism: George Price and the Search for the Origins of Kindness'', New York :
W.W. Norton W. W. Norton & Company is an American publishing company based in New York City. Established in 1923, it has been owned wholly by its employees since the early 1960s. The company is known for its Norton Anthologies (particularly ''The Norton Ant ...
,
Price had a religious experience and became an ardent scholar of the New Testament. He believed that there had been too many coincidences in his life. In particular, he wrote a lengthy essay titled ''The Twelve Days of Easter'', arguing that the calendar of events surrounding Jesus of
Nazareth Nazareth ( ; ar, النَّاصِرَة, ''an-Nāṣira''; he, נָצְרַת, ''Nāṣəraṯ''; arc, ܢܨܪܬ, ''Naṣrath'') is the largest city in the Northern District of Israel. Nazareth is known as "the Arab capital of Israel". In ...
's death in Easter Week was actually slightly longer. Later he turned away from
Biblical scholarship Biblical criticism is the use of critical analysis to understand and explain the Bible. During the eighteenth century, when it began as ''historical-biblical criticism,'' it was based on two distinguishing characteristics: (1) the concern to ...
and instead dedicated his life to community work, helping the needy of North London.


Other work in evolutionary theory

Price developed a new interpretation of Fisher's fundamental theorem of natural selection, the Price equation, which has now been accepted as the best interpretation of a formerly enigmatic result. He wrote what is still widely held to be the best mathematical, biological and evolutionary representation of
altruism Altruism is the principle and moral practice of concern for the welfare and/or happiness of other human beings or animals, resulting in a quality of life both material and spiritual. It is a traditional virtue in many cultures and a core as ...
. He also pioneered the application of
game theory Game theory is the study of mathematical models of strategic interactions among rational agents. Myerson, Roger B. (1991). ''Game Theory: Analysis of Conflict,'' Harvard University Press, p.&nbs1 Chapter-preview links, ppvii–xi It has appli ...
to evolutionary biology, in a co-authored 1973 paper with John Maynard Smith. Furthermore, Price reasoned that in the same way as an organism may sacrifice itself and further its genes (
altruism Altruism is the principle and moral practice of concern for the welfare and/or happiness of other human beings or animals, resulting in a quality of life both material and spiritual. It is a traditional virtue in many cultures and a core as ...
) an organism may sacrifice itself to eliminate others of the same species if it enabled closely related organisms to better propagate their related genes. This negative altruism was described in a paper published by W. D. Hamilton and is termed
Hamiltonian spite Within the field of Sociobiology, social evolution, Hamiltonian spite is a term for spite (game theory), spiteful behaviours occurring among conspecifics that have a cost for the actor and a negative impact upon the recipient. Theories on altruism ...
. Price's 'mathematical' theory of altruism reasons that organisms are more likely to show altruism toward each other as they become more genetically similar to each other. Thus, in a species that requires two parents to reproduce, an organism is most likely to show altruistic behavior to a biological parent, full sibling, or direct offspring. The reason for this is that each of these relatives' genetic makeup contains (on average in the case of siblings) 50% of the genes that are found in the original organism. So if the original organism dies as a result of an altruistic act it can still manage to propagate its full genetic heritage as long as two or more of these close relatives are saved. Consequently, an organism is less likely to show altruistic behavior to a biological grandparent, grandchild, aunt/uncle, niece/nephew or half-sibling (each carry one-fourth of the genes found in the original organism); and even less likely to show altruism to a first cousin (carrying one-eighth of the genes found in the original organism). The theory then asserts that the further genetically removed two organisms are from each other, the less likely they are to show altruism to each other.


Helping the homeless

Price grew increasingly depressed by the implications of his equation. As part of an attempt to prove his theory right or wrong, he began showing an ever-increasing amount (in both quality and quantity) of random kindness to complete strangers. In this way, he dedicated the latter part of his life to helping the
homeless Homelessness or houselessness – also known as a state of being unhoused or unsheltered – is the condition of lacking stable, safe, and adequate housing. People can be categorized as homeless if they are: * living on the streets, also kn ...
, often inviting homeless people to live in his house. Sometimes, when the people in his house became a distraction, he slept in his office at the
Galton Laboratory The Galton Laboratory was a laboratory for research into eugenics and then into human genetics based at University College London in London, England. It was originally established in 1904, and became part of UCL's biology department in 1996. The a ...
. He also gave up everything to help alcoholics; yet as he helped them steal his belongings, he increasingly fell into depression. He was eventually evicted from his rented house owing to a construction project in the area, making him unhappy because he could no longer provide housing for the homeless. He moved to various squats in the North London area, and became depressed over Christmas, 1974.


Death

Possibly due to the long-term complications of his thyroid treatment, Price committed suicide on January 6, 1975, by cutting his
carotid artery Carotid artery may refer to: * Common carotid artery, often "carotids" or "carotid", an artery on each side of the neck which divides into the external carotid artery and internal carotid artery * External carotid artery, an artery on each side of t ...
with a pair of nail scissors. His body was identified by his close colleague, W.D. Hamilton. A memorial service was held for Price in Euston. The only persons present from academia were Hamilton and Maynard Smith, the other few mourners being those who had come to know him through his community work. He is buried in St Pancras Cemetery.


Recognition

Price's contributions were largely overlooked for 20 years; he had worked in
theoretical biology Mathematical and theoretical biology, or biomathematics, is a branch of biology which employs theoretical analysis, mathematical models and abstractions of the living organisms to investigate the principles that govern the structure, development a ...
for only a short time and was not very thorough in publishing papers. This has changed in recent years. An article by James Schwartz published in 2000 (also available at http://linguafranca.mirror.theinfo.org/0007/altruist.html, https://bio.kuleuven.be/ento/pdfs/schwartz2000.pdf and https://pages.uoregon.edu/myagkov/Death%20of%20an%20Altruist.html) was the beginning of the historical redress. In 2010, Oren Harman's
LA Times Book Prize Since 1980, the ''Los Angeles Times'' has awarded a set of annual book prizes. The Prizes currently have nine categories: Los Angeles Times Book Prize for Biography, biography, Los Angeles Times Book Prize for Current Interest, current interest, ...
winning biography, ''The Price of Altruism: George Price and the Search for the Origins of Kindness'' was published, and has received major attention, finally bringing Price and his story to the general public. A stage play about Price, ''The Altruists'' by Craig Baxter, won the fourth STAGE International Script Competition. On 29 March 2016, Farnworth's play ''Calculating Kindness'' opened for a sold-out three-week run at the Camden People's Theatre.


In media

*The story of Price's equation, its application to altruism, and the profound effects this supposedly had on its discoverer's life were featured in an episode of '' Dark Matters: Twisted But True'', in a segment entitled "Killed by Kindness". *Similar coverage, both of Price's contributions to studies of altruism and his efforts to disprove the supposed selfishness of altruism, was discussed in
The Good Show
episode of the NPR series '' Radiolab'' in December 2010. *The life and work of Price and his work with Bill Hamilton is the central theme of the third and final documentary of the series ''
All Watched Over by Machines of Loving Grace "All Watched Over by Machines of Loving Grace" is a poem by Richard Brautigan first published in his 1967 collection of the same name, his fifth book of poetry. It presents an enthusiastic description of a technological utopia in which machine ...
'' by Adam Curtis and includes an interview with his daughter Kathleen. *The life story of George R. Price was dramatized in 2010 by Craig Baxter, whose pla
"The Altruists"
won th

International Playwriting Competition in that year; and again in 2016 by Lydia Adetunji as
play
titled "Calculating Kindness". *Northern Irish singer songwriter
Joshua Burnside Joshua Burnside (born 11 July 1989) is a Northern Irish folk singer-songwriter based in Belfast. His music incorporates elements of Irish folk, and Scottish folk rock, Americana, world music, sound collage and electronica. His debut album ''E ...
wrote about Price and his fate in his song ''George Price''


Legacy

The Papers of George R Price are housed at the British Library. The papers can be accessed through the British Library catalogue.The Papers of George R Price
archives and manuscripts catalogue, the British Library. Retrieved 15 May 2020


Bibliography

* * * * * * * * * (written circa 1971) * Harman, Oren. (2010). ''The Price of Altruism: George Price and the Search for the Origins of Kindness'', Bodley Head.


References


Further reading

* Harman, Oren (2010). The Price of Altruism: George Price and the Search for the Origins of Kindness (New York, W.W. Norton

*Frank, S.A. (1995). George Price's contributions to Evolutionary Genetics. ''Journal of Theoretical Biology'' 175: 373-38
abstract

full text, pdf 412 KB
(both from http://www.stevefrank.org) * Frank, S.A. (1997). The Price Equation, Fisher's fundamental theorem, kin selection, and causal analysis. '' Evolution (journal), Evolution'' 51:1712–172
full text, pdf 551 KB
* Frank, S.A. (2002). Price, George. In: M. Pagel (ed) ''
Encyclopedia of Evolution The ''Encyclopedia of Evolution'' is a print encyclopedia of evolutionary biology edited by Mark Pagel and published in 2002 by Oxford University Press. It consists of 370 original articles written by leading experts including Richard Dawkins, S ...
'' pp930–
pdf file
* * * * * {{DEFAULTSORT:Price, George R. 1922 births 1975 suicides 20th-century American mathematicians Academics of University College London American physical chemists Critics of parapsychology Converts to Protestantism from atheism or agnosticism Game theorists Squatters Harvard University faculty Population geneticists Suicides by sharp instrument in England American Christian writers Suicides in London University of Chicago alumni Christian radicals Burials at St Pancras and Islington Cemetery Stuyvesant High School alumni Mathematicians from New York (state) Birch Wathen Lenox School alumni 1975 deaths