George R. Price
Encyclopedia
George Robert Price 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
.
.
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 divorce
d in 1955, having had two daughters, Annamarie and Kathleen.
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 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.
, 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
.
. He believed that there had been too many coincidence
s 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.
, 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.
. 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.
. 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
.
'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. A stage play about Price 'The Altruists' by Craig Baxter won the 4th STAGE International Script Competition
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...
population geneticist
Population genetics
Population genetics is the study of allele frequency distribution and change under the influence of the four main evolutionary processes: natural selection, genetic drift, mutation and gene flow. It also takes into account the factors of recombination, population subdivision and population...
. Originally a physical chemist
Physical chemistry
Physical chemistry is the study of macroscopic, atomic, subatomic, and particulate phenomena in chemical systems in terms of physical laws and concepts...
and later a science journalist
Scientific journalism
Scientific journalism is the practice of including primary sources along with journalistic stories. The concept has been championed by Julian Assange of Wikileaks and is inspired by the philosophy of Karl Popper.-Technology:...
, he moved to London
London
London is the capital city of :England and the :United Kingdom, the largest metropolitan area in the United Kingdom, and the largest urban zone in the European Union by most measures. Located on the River Thames, London has been a major settlement for two millennia, its history going back to its...
in 1967, where he worked in theoretical biology at the Galton Laboratory
Galton Laboratory
The Galton Laboratory, was a laboratory for research into human genetics based at University College London in London, United Kingdom. It was originally established in 1904, and became part of UCL's biology department in 1996....
, making three important contributions: first, rederiving W.D. Hamilton's work on kin selection
Kin selection
Kin selection refers to apparent strategies in evolution that favor the reproductive success of an organism's relatives, even at a cost to the organism's own survival and reproduction. Charles Darwin was the first to discuss the concept of group/kin selection...
with a new Price equation
Price equation
The Price equation is a covariance equation which is a mathematical description of evolution and natural selection. The Price equation was derived by George R. Price, working in London to re-derive W.D. Hamilton's work on kin selection...
; second, introducing (with John Maynard Smith
John Maynard Smith
John Maynard Smith,His surname was Maynard Smith, not Smith, nor was it hyphenated. F.R.S. was a British theoretical evolutionary biologist and geneticist. Originally an aeronautical engineer during the Second World War, he took a second degree in genetics under the well-known biologist J.B.S....
) the concept of the evolutionarily stable strategy
Evolutionarily stable strategy
In game theory and behavioural ecology, an evolutionarily stable strategy , which is sometimes also called an evolutionary stable strategy, is a strategy which, if adopted by a population of players, cannot be invaded by any alternative strategy that is initially rare. An ESS is an equilibrium...
(ESS), a central concept in game theory
Game theory
Game theory is a mathematical method for analyzing calculated circumstances, such as in games, where a person’s success is based upon the choices of others...
; and third, formalising Fisher's fundamental theorem of natural selection
Fisher's fundamental theorem of natural selection
In population genetics, R. A. Fisher's fundamental theorem of natural selection was originally stated as:Or, in more modern terminology:- History :...
. After giving all his possessions to the poor, he committed suicide
Suicide
Suicide is the act of intentionally causing one's own death. Suicide is often committed out of despair or attributed to some underlying mental disorder, such as depression, bipolar disorder, schizophrenia, alcoholism, or drug abuse...
.
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 DepressionGreat Depression
The Great Depression was a severe worldwide economic depression in the decade preceding World War II. The timing of the Great Depression varied across nations, but in most countries it started in about 1929 and lasted until the late 1930s or early 1940s...
.
Having attended the prestigious Stuyvesant High School
Stuyvesant High School
Stuyvesant High School , commonly referred to as Stuy , is a New York City public high school that specializes in mathematics and science. The school opened in 1904 on Manhattan's East Side and moved to a new building in Battery Park City in 1992. Stuyvesant is noted for its strong academic...
in New York
New York
New York is a state in the Northeastern region of the United States. It is the nation's third most populous state. New York is bordered by New Jersey and Pennsylvania to the south, and by Connecticut, Massachusetts and Vermont to the east...
, Price graduated with a degree in chemistry
Chemistry
Chemistry is the science of matter, especially its chemical reactions, but also its composition, structure and properties. Chemistry is concerned with atoms and their interactions with other atoms, and particularly with the properties of chemical bonds....
from University of Chicago
University of Chicago
The University of Chicago is a private research university in Chicago, Illinois, USA. It was founded by the American Baptist Education Society with a donation from oil magnate and philanthropist John D. Rockefeller and incorporated in 1890...
in 1943 and received his doctorate
Doctorate
A doctorate is an academic degree or professional degree that in most countries refers to a class of degrees which qualify the holder to teach in a specific field, A doctorate is an academic degree or professional degree that in most countries refers to a class of degrees which qualify the holder...
in the subject from the same institution in 1946, having worked on the Manhattan Project
Manhattan Project
The Manhattan Project was a research and development program, led by the United States with participation from the United Kingdom and Canada, that produced the first atomic bomb during World War II. From 1942 to 1946, the project was under the direction of Major General Leslie Groves of the US Army...
.
In 1947 he married Julia Madigan, but their relationship was contentious because George was a strong atheist
Strong atheism
Positive atheism is a term popularly used to describe the form of atheism that maintains that "There is no god" is a true statement. Negative atheism refers to any other type of non-theism, wherein a person does not believe in the existence of any deity, but does not explicitly claim that the...
whilst his wife was a practicing Roman Catholic. They divorce
Divorce
Divorce is the final termination of a marital union, canceling the legal duties and responsibilities of marriage and dissolving the bonds of matrimony between the parties...
d in 1955, having had two daughters, Annamarie and Kathleen.
Early career
Between 1946 and 1948, Price was an instructor in chemistry at Harvard UniversityHarvard University
Harvard University is a private Ivy League university located in Cambridge, Massachusetts, United States, established in 1636 by the Massachusetts legislature. Harvard is the oldest institution of higher learning in the United States and the first corporation chartered in the country...
and consultant to Argonne National Laboratory
Argonne National Laboratory
Argonne National Laboratory is the first science and engineering research national laboratory in the United States, receiving this designation on July 1, 1946. It is the largest national laboratory by size and scope in the Midwest...
. Later, he worked as a research associate in medicine
Medicine
Medicine is the science and art of healing. It encompasses a variety of health care practices evolved to maintain and restore health by the prevention and treatment of illness....
at the University of Minnesota
University of Minnesota
The University of Minnesota, Twin Cities is a public research university located in Minneapolis and St. Paul, Minnesota, United States. It is the oldest and largest part of the University of Minnesota system and has the fourth-largest main campus student body in the United States, with 52,557...
, working on, amongst other things, fluorescence microscopy and liver perfusion. In 1955 and 1956, he published two papers in the journal Science
Science (journal)
Science is the academic journal of the American Association for the Advancement of Science and is one of the world's top scientific journals....
criticising the apparently pseudoscientific
Pseudoscience
Pseudoscience is a claim, belief, or practice which is presented as scientific, but which does not adhere to a valid scientific method, lacks supporting evidence or plausibility, cannot be reliably tested, or otherwise lacks scientific status...
claims of extra-sensory perception
Extra-sensory perception
Extrasensory perception involves reception of information not gained through the recognized physical senses but sensed with the mind. The term was coined by Frederic Myers, and adopted by Duke University psychologist J. B. Rhine to denote psychic abilities such as telepathy, clairaudience, and...
.
Continuing with science journalism, he tried to write a book entitled No Easy Way about the United States
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...
' Cold War
Cold War
The Cold War was the continuing state from roughly 1946 to 1991 of political conflict, military tension, proxy wars, and economic competition between the Communist World—primarily the Soviet Union and its satellite states and allies—and the powers of the Western world, primarily the United States...
with the Soviet Union
Soviet Union
The Soviet Union , officially the Union of Soviet Socialist Republics , was a constitutionally socialist state that existed in Eurasia between 1922 and 1991....
and the People's Republic of China
People's Republic of China
China , officially the People's Republic of China , is the most populous country in the world, with over 1.3 billion citizens. Located in East Asia, the country covers approximately 9.6 million square kilometres...
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
IBM
International Business Machines Corporation or IBM is an American multinational technology and consulting corporation headquartered in Armonk, New York, United States. IBM manufactures and sells computer hardware and software, and it offers infrastructure, hosting and consulting services in areas...
as a consultant on graphic data processing. In 1966 he was treated for thyroid cancer
Thyroid cancer
Thyroid neoplasm is a neoplasm or tumor of the thyroid. It can be a benign tumor such as thyroid adenoma, or it can be a malignant neoplasm , such as papillary, follicular, medullary or anaplastic thyroid cancer. Most patients are 25 to 65 years of age when first diagnosed; women are more affected...
, but the operation to remove the tumour left his shoulder partially paralysed and left him reliant on thyroxine
Thyroxine
Thyroxine, or 3,5,3',5'-tetraiodothyronine , a form of thyroid hormones, is the major hormone secreted by the follicular cells of the thyroid gland.-Synthesis and regulation:...
medication. With the money from his medical insurance, he moved to the United Kingdom
United Kingdom
The United Kingdom of Great Britain and Northern IrelandIn the United Kingdom and Dependencies, other languages have been officially recognised as legitimate autochthonous languages under the European Charter for Regional or Minority Languages...
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 selectionKin selection
Kin selection refers to apparent strategies in evolution that favor the reproductive success of an organism's relatives, even at a cost to the organism's own survival and reproduction. Charles Darwin was the first to discuss the concept of group/kin selection...
, and with no training in population genetics
Population genetics
Population genetics is the study of allele frequency distribution and change under the influence of the four main evolutionary processes: natural selection, genetic drift, mutation and gene flow. It also takes into account the factors of recombination, population subdivision and population...
or statistics
Statistics
Statistics is the study of the collection, organization, analysis, and interpretation of data. It deals with all aspects of this, including the planning of data collection in terms of the design of surveys and experiments....
devised the Price equation
Price equation
The Price equation is a covariance equation which is a mathematical description of evolution and natural selection. The Price equation was derived by George R. Price, working in London to re-derive W.D. Hamilton's work on kin selection...
, a covariance
Covariance
In probability theory and statistics, covariance is a measure of how much two variables change together. Variance is a special case of the covariance when the two variables are identical.- Definition :...
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
Alan Robertson
Alan Robertson FRS was an English population geneticist. Originally a chemist, he was recruited after the Second World War to work on animal genetics on behalf of the British government, and continued in this sphere until his retirement in 1985...
and C. C. Li, its second component allowed it to be applied to all levels of selection, meiotic drive, traditional natural selection
Natural selection
Natural selection is the nonrandom process by which biologic traits become either more or less common in a population as a function of differential reproduction of their bearers. It is a key mechanism of evolution....
with an extension into inclusive fitness
Inclusive fitness
In evolutionary biology and evolutionary psychology, the inclusive fitness of an organism is the sum of its classical fitness and the number of equivalents of its own offspring it can add to the population by supporting others...
and group selection
Group selection
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 alleles' effect on the fitness of individuals within that group....
.
Conversion
On 6 June 1970, Price had a religious experience and became an ardent scholar of the New TestamentNew Testament
The New Testament is the second major division of the Christian biblical canon, the first such division being the much longer Old Testament....
. He believed that there had been too many coincidence
Coincidence
A coincidence is an event notable for its occurring in conjunction with other conditions, e.g. another event. As such, a coincidence occurs when something uncanny, accidental and unexpected happens under conditions named, but not under a defined relationship...
s in his life. In particular, he wrote a lengthy essay entitled The Twelve Days of Easter, arguing that the calendar of events surrounding Jesus
Jesus
Jesus of Nazareth , commonly referred to as Jesus Christ or simply as Jesus or Christ, is the central figure of Christianity...
of Nazareth
Nazareth
Nazareth is the largest city in the North District of Israel. Known as "the Arab capital of Israel," the population is made up predominantly of Palestinian Arab citizens of Israel...
's death in Easter Week
Easter
Easter is the central feast in the Christian liturgical year. According to the Canonical gospels, Jesus rose from the dead on the third day after his crucifixion. His resurrection is celebrated on Easter Day or Easter Sunday...
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 selectionFisher's fundamental theorem of natural selection
In population genetics, R. A. Fisher's fundamental theorem of natural selection was originally stated as:Or, in more modern terminology:- History :...
, the Price equation
Price equation
The Price equation is a covariance equation which is a mathematical description of evolution and natural selection. The Price equation was derived by George R. Price, working in London to re-derive W.D. Hamilton's work on kin selection...
, 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
Altruism
Altruism is a concern for the welfare of others. It is a traditional virtue in many cultures, and a core aspect of various religious traditions, though the concept of 'others' toward whom concern should be directed can vary among cultures and religions. Altruism is the opposite of...
. 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
Altruism
Altruism is a concern for the welfare of others. It is a traditional virtue in many cultures, and a core aspect of various religious traditions, though the concept of 'others' toward whom concern should be directed can vary among cultures and religions. Altruism is the opposite of...
) 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
Hamiltonian spite
Within the field of social evolution, Hamiltonian spite is a term for those behaviours that occur between conspecifics, and have a cost for the actor and a negative impact for the recipient. Why do animals and humans exhibit spiteful behaviours?-Theories on altruism and spitefulness:W. D...
.
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 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 LaboratoryGalton Laboratory
The Galton Laboratory, was a laboratory for research into human genetics based at University College London in London, United Kingdom. It was originally established in 1904, and became part of UCL's biology department in 1996....
. 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
Squatting
Squatting consists of occupying an abandoned or unoccupied space or building, usually residential, that the squatter does not own, rent or otherwise have permission to use....
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 arteryCarotid artery
Carotid artery can refer to:* Common carotid artery* External carotid artery* Internal 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
St. Pancras and Islington Cemetery
The St. Pancras and Islington Cemetery in East Finchley, North London while situated in the London Borough of Barnet is actually two cemeteries, owned by two other London Boroughs, Camden and Islington...
.
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 HarmanOren Harman
Oren Harman is a writer and a professor of the history of science at Bar Ilan University in Israel. He is best known as the author of several critically acclaimed scholarly works as well as works published by commercial publishers popularizing scientific ideas for a mass audience.- Biography :Oren...
'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. A stage play about Price 'The Altruists' by Craig Baxter won the 4th STAGE International Script Competition
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. EvolutionEvolution (journal)Evolution, the International Journal of Organic Evolution, is a leading monthly scientific journal that publishes significant new results of empirical or theoretical investigations concerning facts, processes, mechanics, or concepts of evolutionary phenomena and events. Evolution is published by...
51:1712-1729 full text, pdf 551 KB - Frank, S.A. (2002). Price, George. In: M. Pagel (ed) Encyclopedia of EvolutionEncyclopedia of EvolutionThe Encyclopedia of Evolution is a print encyclopedia of evolutionary biology, edited by Mark Pagel. A similar book, the Cambridge Encyclopedia of Evolution is edited by Steve Jones.-Bibliography:...
pp930–1 pdf file - Selflessness of strangers, The Economist, May 20, 2010