George R. Price

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George R. Price
File:George Robert Price.jpg
Born October 6, 1922
United States
Died January 6, 1975(1975-01-06) (aged 52)
London, United Kingdom
Fields Population genetics, physical chemistry
Alma mater University of Chicago
Spouse Julia Madigan (m. 1947 div. 1955)

George Robert Price (October 6, 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 converting to Christianity and giving all his possessions to the poor, he committed suicide.

Early life

Price was born in 1922. His father, an electrician, died when Price was four. His mother was a former opera singer, and the family struggled through the Great Depression.[1]

Having attended Stuyvesant High 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.[1]

In 1947 he married Julia Madigan, but their relationship was contentious because George was a strong atheist whilst his wife was a practicing Roman Catholic.[1] 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 took a position at Bell Laboratories to work on the chemistry of transistors.[2] 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 criticising the pseudoscientific claims of extrasensory perception.[3][4]

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 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.

To Britain

W.D. Hamilton failed to recall when Price first contacted him, but says Price had read Hamilton's 1964 papers on kin selection,[5][6] and with no training in population genetics or statistics devised the Price equation,[7] a covariance equation that generated the change in allele frequency of a population.[8] Although the first part of the equation had previously been derived by Alan Robertson[9] and C. C. Li,[10] 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 to Christianity

On 6 June 1970,[2] 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.[7] He wrote what is still widely held to be the best mathematical, biological and evolutionary representation of altruism. He also pioneered the application of game theory to evolutionary biology, in a co-authored 1973 paper with John Maynard Smith.[11] Furthermore, Price reasoned that in the same way as an organism may sacrifice itself and further its genes (altruism) 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.

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. As such, 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 make up 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 contain one-fourth of the genes found in the original organism); and even less likely to show altruism to a first cousin (contains one-eighth of the genes found in the original organism). The theory then holds that the further genetically removed two organisms are from each other the less likely they are to show altruism to each other. If true then altruistic (kind) behavior is not truly selfless and is instead an adaptation that organisms have in order to promote their own genetic heritage.

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 Price began showing an ever increasing amount (in both quality and quantity) of random kindness to complete strangers. As such Price dedicated the latter 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, yet as he helped them some stole his belongings, and he increasingly fell into depression.[citation needed]

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.[citation needed]


Unable to prove his theory right or wrong, Price committed suicide on January 6, 1975. His body was identified by his close colleague W.D. Hamilton.[12]

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 an unmarked grave in St Pancras' Cemetery.[13] [14]


Price's contributions were largely overlooked for twenty years; he had worked in theoretical biology 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 was the beginning of the historical redress. More recently, Oren Harman's LA Times Book Prize winning biography, The Price of Altruism: George Price and the Search for the Origins of Kindness, has received major attention, finally bringing George 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.[15]

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 Radio Lab.
  • 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" by Adam Curtis and includes an interview with his daughter Kathleen.



  1. 1.0 1.1 1.2 Dugatkin, Lee Alan (2011). The Altruism Equation. Princeton University Press. p. 108.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  2. 2.0 2.1 Oren Harman (2010) The Price of Altruism: George Price and the Search for the Origins of Kindness, New York : W.W. Norton, ISBN 978-0-393-06778-1
  3. Price, G.R. (1955). "Science and the supernatural". Science. 122 (3165): 359–367. Bibcode:1955Sci...122..359P. doi:10.1126/science.122.3165.359. JSTOR 1750914.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  4. Price, G.R. (1956). "Where is the definitive experiment?". Science. 123 (3184): 17–18. Bibcode:1956Sci...123...17P. doi:10.1126/science.123.3184.17. JSTOR 1750107.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  5. Hamilton, W.D. (1964). "The evolution of social behaviour I". Journal of Theoretical Biology. 7 (1): 1–16. doi:10.1016/0022-5193(64)90038-4. PMID 5875341.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  6. Hamilton, W.D. (1964). "The evolution of social behaviour II". Journal of Theoretical Biology. 7 (1): 17–52. doi:10.1016/0022-5193(64)90039-6. PMID 5875340.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  7. 7.0 7.1 Price, G.R. (1972). "Fisher's fundamental theorem made clear". Annals of Human Genetics. 36 (2): 129–140. doi:10.1111/j.1469-1809.1972.tb00764.x. PMID 4656569.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  8. Hamilton, W.D. (1996). Narrow Roads of Gene Land, Volume 1. Oxford University Press, Oxford. ISBN 0-7167-4530-5.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  9. Robertson, Alan (1966). "A Mathematical Model of the Culling Process in Dairy Cattle". Animal Production. 8: 95–108. doi:10.1017/S0003356100037752.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  10. Li, C.C. (April 1967). "Fundamental Theorem of Natural Selection". Nature. 214 (5087): 505–506. Bibcode:1967Natur.214..505L. doi:10.1038/214505a0.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  11. Maynard Smith, J.; Price, G.R. (1973). "The logic of animal conflict". Nature. 246 (5427): 15–18. Bibcode:1973Natur.246...15S. doi:10.1038/246015a0.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  12. Brown, Andrew (2000). The Darwin Wars: The Scientific Battle for the Soul of Man. London: Touchstone Books. p. 1. ISBN 0-684-85145-8.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  13. Harman, O. (2010). The Price of Altruism: George Price and the Search for the Origins of Kindness. W. W. Norton & Company.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  14. George Robert Price at Find a Grave
  15. THE ALTRUISTS, by Craig Baxter

Further reading