George R. Price (1922 - January 6, 1975) was a 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: Firstly, rederiving W.D. Hamilton's work on kin selection with a new Price equation; secondly introducing (with John Maynard Smith) the concept of the evolutionarily stable strategy (ESS), a central concept in game theory; and thirdly, formalising Fisher's fundamental theorem of natural selection. A troubled man, Price converted from atheism to Christianity, and after giving all his possessions to the poor, committed suicide. 1922 was a common year starting on Sunday (see link for calendar). ... January 6 is the 6th day of the year in the Gregorian Calendar. ... 1975 was a common year starting on Wednesday (the link is to a full 1975 calendar). ... Population genetics is the study of the distribution of and change in allele frequencies under the influence of the five evolutionary forces: natural selection, genetic drift, mutation, migration and nonrandom mating. ... Physical Chemistry is the combined science of physics, chemistry, thermodynamics, and quantum mechanics which functions to provide molecular-level interpretations of observed macroscopic phenomena. ... The clock tower of the Palace of Westminster, which contains Big Ben London is the capital city of the United Kingdom and of England. ... 1967 was a common year starting on Sunday of the Gregorian calendar. ... Theoretical biology is an interdisciplinary field of academic study and research that involves the use of quantitative tools in biology. ... The Galton Laboratory, based at University College London, conducts research into human genetics. ... This article is about the British biologist Bill Hamilton. ... Kin selection was first suggested by Darwin as an explanation of the sterile castes of social insects and has later been mathematically defined by W. D. Hamilton as a mechanism for the evolution of apparently altruistic acts. ... The Price equation (also known as Prices equation) is a covariance equation which is a mathematical description of evolution and natural selection. ... John Maynard Smith Professor John Maynard Smith, F.R.S. (6 January 1920 – 19 April 2004) was a British evolutionary biologist and geneticist. ... The evolutionarily stable strategy (or ESS; also evolutionary stable strategy) is a central concept in game theory introduced by John Maynard Smith and George R. Price in 1973 (a full account is given by Maynard Smiths 1982 book Evolution and the Theory of Games). ... Game theory is a branch of applied mathematics that uses models to study interactions with formalised incentive structures (games). Unlike decision theory, which also studies formalised incentive structures, game theory encompasses decisions that are made in an environment where various players interact strategically. ... In population genetics, Ronald Fishers fundamental theorem of natural selection was originally stated as: The rate of increase in fitness of any organism at any time is equal to its genetic variance in fitness at that time. ... Atheism, in its broadest sense, is characterized by an absence of belief in the existence of gods, thus contrasting with theism. ... Christianity is a monotheistic religion based on the life and teachings of Jesus of Nazareth as presented in the New Testament writings of his early followers. ... The Death of Socrates by Jacques-Louis David, 1787 Suicide (from Latin sui caedere, to kill oneself) is the act of willfully ending ones own life; it is sometimes a noun for one who has committed or attempted the act. ...

Biography

James Swartz (2000) has written an excellent 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. This article needs to be cleaned up to conform to a higher standard of quality. ...

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. Public education is schooling provided by the government, and paid for by taxes, in countries other than England where public schools are privately funded. ... State nickname: Empire State Other U.S. States Capital Albany Largest city New York City Governor George Pataki (R) Official languages None (English is de facto) Area 141,205 km² (27th)  - Land 122,409 km²  - Water 18,795 km² (13. ... 1943 is a common year starting on Friday. ... Chemistry (in Greek: χημεία) is the science of matter that deals with the composition, structure, and properties of substances and with the transformations that they undergo. ... The University of Chicago is a private co-educational university located in Chicago, Illinois. ... Control panels and operators for calutrons at the Y-12 Plant in Oak Ridge, Tennessee. ...

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. The Roman Catholic Church, most often spoken of simply as the Catholic Church, is the largest Christian church, with over one billion members. ... Divorce or dissolution of marriage is the ending of a marriage, which can be contrasted with an annulment which is a declaration that a marriage is void, though the effects of marriage may be recognized in such unions, such as spousal support, child custody and distribution of property. ... 1955 is a common year starting on Saturday of the Gregorian calendar. ...

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. 1946 was a common year starting on Tuesday. ... 1948 is a leap year starting on Thursday (link will take you to calendar). ... Harvard University is a private university in Cambridge, Massachusetts, USA, and a member of the Ivy League. ... Argonne National Laboratory is one of the United States governments oldest and largest science and engineering research national laboratories and is the largest in the Midwest. ... 1955 is a common year starting on Saturday of the Gregorian calendar. ... Medicine on the Web NLM (National Library of Medicine, contains resources for patients and healthcare professionals) Virtual Hospital (digital health sciences library by the University of Iowa) Online Medical Dictionary Collection of links to free medical resources Categories: Medicine | Health ... University of Minnesota, Twin Cities This article is about the oldest and largest campus of the University of Minnesota. ... Microscopy is any technique for producing visible images of structures or details too small to otherwise be seen by the human eye. ... 1956 was a leap year starting on Sunday of the Gregorian calendar. ... Science is the journal of the American Association for the Advancement of Science. ... Phrenology is seen today as a classic example of pseudoscience. ... Extra-sensory perception, or ESP, is the name given to any ability to acquire information by means other than the five canonical senses (taste, sight, touch, smell, and hearing), or any other sense well known to science (balance, proprioception, etc). ...

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. For the generic term for a high-tension rivalry between countries, see cold war (war). ...

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. 1961 was a common year starting on Sunday (link will take you to calendar). ... 1967 was a common year starting on Sunday of the Gregorian calendar. ... International Business Machines Corporation (IBM, or colloquially, Big Blue) NYSE: IBM (incorporated June 15, 1911, in operation since 1888) is headquartered in Armonk, NY, USA. The company manufactures and sells computer hardware, software, and services. ... Thyroid cancer is cancer of the thyroid gland. ... The thyroid hormones, thyroxine (T4) and triiodothyronine (T3), are tyrosine-based hormones produced by the thyroid gland. ... Health Insurance is a type of insurance whereby the insurer pays the medical costs of the insured if the insured becomes sick due to covered causes, or due to accidents. ... For other uses, see November (disambiguation). ...

To Britain

W.D. Hamilton (1996) failed to recall when Price first contacted him, but says he 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. This article is about the British biologist Bill Hamilton. ... Kin selection was first suggested by Darwin as an explanation of the sterile castes of social insects and has later been mathematically defined by W. D. Hamilton as a mechanism for the evolution of apparently altruistic acts. ... Population genetics is the study of the distribution of and change in allele frequencies under the influence of the five evolutionary forces: natural selection, genetic drift, mutation, migration and nonrandom mating. ... Statistics is a type of data analysis which practice includes the planning, summarizing, and interpreting of observations of a system possibly followed by predicting or forecasting of future events based on a mathematical model of the system being observed. ... The Price equation (also known as Prices equation) is a covariance equation which is a mathematical description of evolution and natural selection. ... In probability theory and statistics, the covariance between two real-valued random variables X and Y, with expected values and is defined as: where E is the expected value. ... Ching Chun Li Ching Chun Li (李景均; pinyin: Lǐ Jǐngjūn; 27 October 1912 — 20 October 2003) was a Chinese American population geneticist and human geneticist. ... Natural selection is a process by which biological populations are altered over time, as a result of the propagation of heritable traits that affect the capacity of individual organisms to survive and reproduce. ... Inclusive fitness encompasses conventional Darwinian fitness with the addition of behaviors that contribute to an organism’s individual fitness through altruism. ... In evolutionary biology, group selection refers to the idea that alleles can become fixed or spread in a population because of the benefits they bestow on groups, regardless of the fitness of individuals within that group. ...

Conversion

On the 6th 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. The New Testament, sometimes called the Greek Testament or Greek Scriptures is the name given to the part of the Christian Bible that was written after the birth of Jesus. ... This article is about the figure known by both Jesus of Nazareth and Jesus Christ. For other usages, see Jesus (disambiguation). ... Easter is the most important religious holiday of the Christian liturgical year, observed in March, April, or May to celebrate the resurrection of Jesus from the dead after his death by crucifixion (see Good Friday), which Christians believe happened at about this time of year around AD 30-33. ...

Fisher's fundamental theorem

He thus clarified Fisher's fundamental theorem of natural selection, which had caused some controversy and misunderstanding. He became to believe that this equation had been a gift from God, a miracle after a religious experience. In population genetics, Ronald Fishers fundamental theorem of natural selection was originally stated as: The rate of increase in fitness of any organism at any time is equal to its genetic variance in fitness at that time. ... The term God is capitalized in the English language as a proper noun when used to refer to a specific monotheistic concept of a Supreme Being in accordance with Christian, Jewish (sometimes as G-d - cf. ... According to many religions, a miracle, derived from the old Latin word miraculum meaning something wonderful, is a striking interposition of divine intervention by God in the universe by which the operations of the ordinary course of Nature are overruled, suspended, or modified. ...

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. The Galton Laboratory, based at University College London, conducts research into human genetics. ...

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. The word squat has different meanings: The act of occupying a building without permission, as practiced by a squatter. ...

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 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 Prices is mentioned in Swartz's biography.

References

Works by Price

• Price, G.R. (1955). Science and the supernatural. Science 122:359-367. JSTOR
• Price, G.R. (1956). Where is the definitive experiement? Science 123:17-18. JSTOR
• Price, G.R. (1970). Selection and covariance. Nature 227:520-521.
• Price, G.R. (1971) Extension of the Hardy--Weinberg law to assortative mating .Annals of Human Genetics 23:344-347
• Price, G.R. and C.A.B. Smith (1972) Fisher's Malthusian parameter and reproductive value.Annals of Human Genetics 3:1-7
• Price, G.R. (1972a). Fisher's fundamental theorem made clear. Annals of Human Genetics 36:129-140.
• Price, G.R. (1972b). Extension of covariance selection mathematics. Annals of Human Genetics 35:485-490.
• Maynard Smith, J. and G.R. Price. (1973). The logic of animal conflict. Nature 246:16-18.
• Price, G.R. (1995). The nature of selection Journal of Theoretical Biology 175:389-396 (written circa 1971)

Science is the journal of the American Association for the Advancement of Science. ... Science is the journal of the American Association for the Advancement of Science. ... Nature is one of the oldest and most reputable general-purpose scientific journals, first published on November 4, 1869. ... Annals of Human Genetics, formerly Annals of Eugenics is a scientific journal. ... Cedric Austen Bardell Smith (February 5, 1917, — January 16, 2002) was a British statistician Categories: Scientist stubs | 1917 births | 2002 deaths ... Annals of Human Genetics, formerly Annals of Eugenics is a scientific journal. ... Annals of Human Genetics, formerly Annals of Eugenics is a scientific journal. ... Annals of Human Genetics, formerly Annals of Eugenics is a scientific journal. ... John Maynard Smith Professor John Maynard Smith, F.R.S. (6 January 1920 – 19 April 2004) was a British evolutionary biologist and geneticist. ... Nature is one of the oldest and most reputable general-purpose scientific journals, first published on November 4, 1869. ... The Journal of Theoretical Biology is a scientific journal dealing with all mathematical and computational aspects of biology. ...

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 0716745305
• Swartz, 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 pdf file