George R. Price

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 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.

Having attended the prestigious 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, having worked on the Manhattan Project.

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. 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 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, 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 Alan Robertson and 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 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. He also pioneered the application of game theory 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) 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-forth 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 farther 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

Unable to accept the selfish reasoning for kindness found in his own mathematical theory of altruism Price began trying to prove the theory wrong by personally 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 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.

Death

Price committed suicide on January 6, 1975, using a pair of nail scissors to cut his own carotid artery. His body was identified by his close colleague Bill Hamilton. 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. He is buried in an unmarked grave in St. Pancras' Cemetery.

Recognition

Price's contributions were 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. An article by James Schwartz published in 2000 was the beginning of the historical redress. More recently, Oren Harman's biography, The Price of Altruism: George Price and the Search for the Origins of Kindness (Norton, 2010), has received major attention, finally bringing George Price and his story to the general public.

Bibliography

• Price, G.R. (1955). "Science and the supernatural". Science. 122 (3165): 359–367. doi:10.1126/science.122.3165.359. JSTOR 1750914.
• Price, G.R. (1956). "Where is the definitive experiment?". Science. 123 (3184): 17–18. doi:10.1126/science.123.3184.17. JSTOR 1750107.
• Price, G.R. (1970). "Selection and covariance" (PDF). Nature. 227 (5257): 520–521. doi:10.1038/227520a0. PMID 5428476.
• Price, G.R. (1971). "Extension of the Hardy—Weinberg law to assortative mating". Annals of Human Genetics. 34: 455–458. doi:10.1111/j.1469-1809.1971.tb00259.x.
• Price, G.R. (1972a). "Extension of covariance selection mathematics". Annals of Human Genetics. 35: 485–490. doi:10.1111/j.1469-1809.1957.tb01874.x. PMID 5073694.
• Price, G.R.; Smith, C.A.B. (1972b). "Fisher's Malthusian parameter and reproductive value". Annals of Human Genetics. 36: 1–7. doi:10.1111/j.1469-1809.1972.tb00577.x. PMID 4656569.
• Price, G.R. (1972c). "Fisher's fundamental theorem made clear". Annals of Human Genetics. 36: 129–140. doi:10.1111/j.1469-1809.1972.tb00764.x. PMID 4656569.
• 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.
• Price, G.R. (1995). The nature of selection. Journal of Theoretical Biology 175:389-396 (written circa 1971)
• Harman, Oren. (2010). The Price of Altruism: George Price and the Search for the Origins of Kindness, Bodley Head. ISBN 978-1-84792-062-1

Further reading

• 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. (1997). The Price Equation, Fisher's fundamental theorem, kin selection, and causal analysis. Evolution 51:1712-1729 full text, pdf 551 KB
• Frank, S.A. (2002). Price, George. In: M. Pagel (ed) Encyclopedia of Evolution pp930–1 pdf file
• Van Veelen, M. (2005). "On the use of the Price equation". Journal of Theoretical Biology. 237 (4): 412–426. doi:10.1016/j.jtbi.2005.04.026. PMID 15953618.
• Selflessness of strangers, The Economist, May 20, 2010