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George R. Price

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George R. Price (1922 – January 6, 1975) was an American population geneticist. Originally a physical chemist and later a science journalist, he moved to London in 1967, where he worked in theoretical biology at the Galton Laboratory, making three important contributions: first, rederiving W.D. Hamilton's work on kin selection with a new Price equation; second, introducing (with John Maynard Smith) the concept of the evolutionarily stable strategy (ESS), a central concept in game theory; and third, formalising Fisher's fundamental theorem of natural selection. After giving all his possessions to the poor, he committed suicide.

Early life

Price was born in 1922, the younger brother of Edison. His father, an electrician, died when he was four. His mother was a former opera singer, and the family struggled through the Great Depression.

Having attended public school in New York, Price graduated with a degree in chemistry from University of Chicago in 1943 and received his doctorate in the subject from the same institution in 1946, having worked on the Manhattan Project.

In 1947 he married Julia Madigan, but their relationship was contentious because George was a militant atheist whilst his wife was a practicing Roman Catholic. They divorced in 1955, having had two daughters, Annamarie and Kathleen.

Early career

Between 1946 and 1948, Price was an instructor in chemistry at Harvard University and consultant to Argonne National Laboratory. Later, he worked as a research associate in medicine at the University of Minnesota, working on, amongst other things, fluorescence microscopy and liver perfusion. In 1955 and 1956, he published two papers in the journal Science criticising the apparently pseudoscientific claims of extra-sensory perception.

Continuing with science journalism, he tried to write a book entitled No Easy Way about the United States' Cold War 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 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.

To Britain

W.D. Hamilton (1996) failed to recall when Price first contacted him, but says Price had read Hamilton's 1964 papers on kin selection, and with no training in population genetics or statistics 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 C. C. Li , its second component allowed it to be applied to all levels of selection, meiotic drive, traditional natural selection with an extension into inclusive fitness and group selection.


On 6 June 1970, 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 entitled The Twelve Days of Easter, arguing that the calendar of events surrounding Jesus of Nazareth's death in Easter Week was actually slightly longer. Later he turned away from Biblical scholarship 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 w\,\Delta z = \operatorname{cov}\left(w_i,z_i\right). He also pioneered the application of game theory to evolutionary biology, in a co-authored 1973 paper with John Maynard Smith.

Helping the homeless

Price dedicated the later part of his life to helping the homeless, 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. He also gave up everything to help alcoholics, as he helped them they stole his belongings causing him to fall into depression.

He was eventually thrown out of his rented house due to a construction project in the area, which made 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. He later committed suicide in the New Year on the 6th January 1975 using a pair of nail scissors to slash his throat. His body was identified by his close colleague Bill Hamilton.[1] Friends said he committed suicide because of despondency over his inability to continue helping the homeless.


A memorial service was held for Price in Euston (not in a church). 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. Price's contributions were then largely overlooked for twenty years; he had worked only in theoretical biology for a short time and was not very thorough in publishing papers. This has changed in recent years, according to a biography of Price written by James Schwartz and published in 2000.



  1. Brown, Andrew (2000). The Darwin Wars: The Scientific Battle for the Soul of Man. London: Touchstone. p. 1. ISBN 0-684-85145-8. 

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