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George R. Price
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Born

October 6, 1922

Died

January 6, 1975 (aged 52)

London

Fields

Population genetics, physical chemistry

Alma mater

University of Chicago

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.

Contents
[hide]

1 Early life

2 Early career

3 To Britain

4 Conversion

5 Other work in evolutionary theory

6 Helping the homeless

7 Death

8 Recognition

9 Bibliography

10 References

11 Further reading

Early life[edit]
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 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.[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[edit]
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, among 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.
[2][3]
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[edit]
W.D. Hamilton failed to recall when Price first contacted him, but says Price had read
Hamilton's 1964 papers on kin selection,[4][5] and with no training in population
genetics or statistics devised the Price equation,[6] a covariance equation that generated

the change in allele frequency of a population.[7] Although the first part of the equation
had previously been derived by Alan Robertson[8] and C. C. Li,[9] 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[edit]
On 6 June 1970,[10] 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[edit]

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.[6] 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.
Prices '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[edit]

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 they
stole his belongings causing him to fall 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]

Death[edit]
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 W.D. Hamilton.[12] 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. 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]
In his suicide note he had mentioned the poem 'The Hound of Heaven' by Francis
Thompson. No one has figured out why he had mentioned it.

Recognition[edit]
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 (Norton, 2010), 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 4th STAGE International Script Competition [14]

Bibliography[edit]