George R. Price

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. A troubled man, Price converted to Christianity from atheism, and after giving all his possessions to the poor, committed suicide.

Biography
James Schwartz (2000) has written a biography of Price.

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, in 1943 Price graduated with a degree in chemistry from University of Chicago 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. Between 1955 and he was 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 journalist papers in the journal Science critisicing the apparently pseudoscientific claims of extra-sensory perception.

Continuing with science journalism, he tried to write a book entitled No Easy Way about what the United States should "fight" the 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 he 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 been 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.

Conversion
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 social work, helping the needy of North London.

Fisher's fundamental theorem
He thus clarified Fisher's fundamental theorem of natural selection, which had caused some controversy and misunderstanding. He believed that this equation had been a gift from God, a miracle after a religious experience.

Helping the homeless
He then dedicated his life to helping the homeless, and invited many 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. At one point a violent man whose partner he had helped escape caused trouble at the office, however, he was able to smooth this over with the lab.

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, where he committed suicide with a pair of nail scissors by slashing his throat at Christmas 1974. Friends said he committed suicide because of despondency over his inability to continue helping the homeless.

Conclusion
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 he had come to know him through his social 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 as is mentioned in Schwartz's biography.

Works by Price

 * Price, G.R. (1955). Science and the supernatural. Science 122:359-367. JSTOR
 * Price, G.R. (1956). Where is the definitive experiment? Science 123:17-18. JSTOR
 * Price, G.R. (1970). Selection and covariance. Nature 227:520-521.PDF
 * Price, G.R. (1971). Extension of the Hardy--Weinberg law to assortative mating. Annals of Human Genetics 34:455-458
 * Price, G.R. (1972). Extension of covariance selection mathematics. Annals of Human Genetics 35:485-490.
 * Price, G.R. and C.A.B. Smith (1972a) Fisher's Malthusian parameter and reproductive value. Annals of Human Genetics 36:1-7
 * Price, G.R. (1972b). Fisher's fundamental theorem made clear. Annals of Human Genetics 36:129-140.
 * Maynard Smith, J. and G.R. Price. (1973). The logic of animal conflict. Nature 246:15-18.
 * Price, G.R. (1995). The nature of selection. Journal of Theoretical Biology 175:389-396 (written circa 1971)

Other references

 * Frank, S.A. (1995) George Price's contributions to Evolutionary Genetics. Journal of Theoretical Biology 175:373-388 abstract - full text, pdf 412 KB (both from http://www.stevefrank.org)
 * Frank, S. A. The Price Equation, Fisher's fundamental theorem, kin selection, and causal analysis. Evolution 51:1712-1729 download pdf file
 * Frank, S.A. (2002) Price, George. In: M. Pagel (ed) Encyclopedia of Evolution pp930-1 pdf file
 * Hamilton, W.D. (1964). The evolution of social behaviour I and II. Journal of Theoretical Biology 7: 1-16 and 17-52. pubmed I and II
 * Hamilton, W.D. (1996). Narrow Roads of Gene Land vol 1. esp ch5 and ch9. Oxford University Press, Oxford. ISBN 0-7167-4530-5
 * Schwartz, J. (2000) Death of an Altruist: Was the man who found the selfless gene too good for this world?. Lingua Franca 10.5: 51-61 (192 KB pdf file)

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