George R. Price

George Robert Price (October 16, 1922 – January 6, 1975) was an American population geneticist.

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.

Price attended Birch Wathen School followed by Stuyvesant High School in New York.

He 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.

Between 1946 and 1948, he 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.

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 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.

Price was a member of the Manhattan Project as a chemist looking into the characteristics of plutonium-235.

In 1955 and 1956, he published two papers in the journal Science criticizing the pseudoscientific claims of extrasensory perception.

Continuing with science journalism, Price tried to write a book titled 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.

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.

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.

Price is often noted for his formulation of the Price equation in 1967.

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 the new Price equation that vindicated group selection; second, introducing (with John Maynard Smith) the concept of the evolutionarily stable strategy (ESS), a central concept in game theory; 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.

With the money from his medical insurance, he moved to the United Kingdom to start a new life in November 1967.

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 titled 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.

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.

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.

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.