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Sewall Green Wright ForMemRS HonFRSE (December 21, 1889March 3, 1988) was an American geneticist known for his influential work on evolutionary theory and also for his work on path analysis. He was a founder of
population genetics Population genetics is a subfield of genetics that deals with genetic differences within and among populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as Adaptation (biology), adaptation, s ...
alongside
Ronald Fisher Sir Ronald Aylmer Fisher (17 February 1890 – 29 July 1962) was a British polymath who was active as a mathematician, statistician, biologist, geneticist, and academic. For his work in statistics, he has been described as "a genius who a ...
and J. B. S. Haldane, which was a major step in the development of the modern synthesis combining
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinians, Augustinian ...
with
evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
. He discovered the inbreeding coefficient and methods of computing it in pedigree animals. He extended this work to
population Population is a set of humans or other organisms in a given region or area. Governments conduct a census to quantify the resident population size within a given jurisdiction. The term is also applied to non-human animals, microorganisms, and pl ...
s, computing the amount of
inbreeding Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely genetic distance, related genetically. By analogy, the term is used in human reproduction, but more commonly refers to the genet ...
between members of populations as a result of random
genetic drift Genetic drift, also known as random genetic drift, allelic drift or the Wright effect, is the change in the Allele frequency, frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene va ...
, and along with Fisher he pioneered methods for computing the distribution of gene frequencies among populations as a result of the interaction of
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
,
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, ...
, migration and genetic drift. Wright also made major contributions to
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
ian and biochemical genetics.


Biography

Sewall Wright was born in Melrose, Massachusetts, to Philip Green Wright and Elizabeth Quincy Sewall Wright. His parents were first cousins, an interesting fact in light of Wright's later research on
inbreeding Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely genetic distance, related genetically. By analogy, the term is used in human reproduction, but more commonly refers to the genet ...
. The family moved three years later after Philip accepted a teaching job at Lombard College, a Universalist college in Galesburg, Illinois. As a child, Wright helped his father and brother print and publish an early book of poems by his father's student Carl Sandburg. At the age of seven, in 1897, he wrote his first "book", entitled ''Wonders of Nature'', and he published his last paper in 1988: he can be claimed, therefore, to be the scientist with the longest career of science writing. Wright's astonishing maturity at the age of seven may be judged from the following excerpt quoted in the obituary:
Have you ever examined the gizzard of a fowl? The gizzard of a fowl is a deep red colar with blu at the top. First on the outside is a very thick muscle. Under this is a white and fleecy layer. Holding very tight to the other. I expect you know that chickens eat sand. The next two layers are rough and rumply. These layers hold the sand. They grind the food. One night when we had company we had chicken-pie. Our Aunt Polly cut open the gizzard, and in it we found a lot of grain, and some corn.
He was the oldest of three gifted brothers—the others being the aeronautical engineer Theodore Paul Wright and the
political scientist Political science is the scientific study of politics. It is a social science dealing with systems of governance and Power (social and political), power, and the analysis of political activities, political philosophy, political thought, polit ...
Quincy Wright. From an early age Wright had a love and talent for
mathematics Mathematics is a field of study that discovers and organizes methods, Mathematical theory, theories and theorems that are developed and Mathematical proof, proved for the needs of empirical sciences and mathematics itself. There are many ar ...
and
biology Biology is the scientific study of life and living organisms. It is a broad natural science that encompasses a wide range of fields and unifying principles that explain the structure, function, growth, History of life, origin, evolution, and ...
. Wright attended Galesburg High School and graduated in 1906. He then enrolled in Lombard College where his father taught, to study
mathematics Mathematics is a field of study that discovers and organizes methods, Mathematical theory, theories and theorems that are developed and Mathematical proof, proved for the needs of empirical sciences and mathematics itself. There are many ar ...
. He was influenced greatly by Professor Wilhelmine Key, one of the first women to receive a Ph.D. in
biology Biology is the scientific study of life and living organisms. It is a broad natural science that encompasses a wide range of fields and unifying principles that explain the structure, function, growth, History of life, origin, evolution, and ...
. Wright received his Ph.D. from
Harvard University Harvard University is a Private university, private Ivy League research university in Cambridge, Massachusetts, United States. Founded in 1636 and named for its first benefactor, the History of the Puritans in North America, Puritan clergyma ...
, where he worked at the Bussey Institute with the pioneering mammalian geneticist William Ernest Castle investigating the inheritance of coat colors in mammals. He worked for the U.S. Department of Agriculture until 1925, when he joined the Department of Zoology at the
University of Chicago The University of Chicago (UChicago, Chicago, or UChi) is a Private university, private research university in Chicago, Illinois, United States. Its main campus is in the Hyde Park, Chicago, Hyde Park neighborhood on Chicago's South Side, Chic ...
. He remained there until his retirement in 1955, when he moved to the
University of Wisconsin–Madison The University of Wisconsin–Madison (University of Wisconsin, Wisconsin, UW, UW–Madison, or simply Madison) is a public land-grant research university in Madison, Wisconsin, United States. It was founded in 1848 when Wisconsin achieved st ...
. He received many honors in his long career, including the
National Medal of Science The National Medal of Science is an honor bestowed by the President of the United States to individuals in science and engineering who have made important contributions to the advancement of knowledge in the fields of behavioral science, behavior ...
(1966), the Balzan Prize (1984), and the Darwin Medal of the
Royal Society The Royal Society, formally The Royal Society of London for Improving Natural Knowledge, is a learned society and the United Kingdom's national academy of sciences. The society fulfils a number of roles: promoting science and its benefits, re ...
(1980). He was a member of the
National Academy of Sciences The National Academy of Sciences (NAS) is a United States nonprofit, NGO, non-governmental organization. NAS is part of the National Academies of Sciences, Engineering, and Medicine, along with the National Academy of Engineering (NAE) and the ...
, the
American Philosophical Society The American Philosophical Society (APS) is an American scholarly organization and learned society founded in 1743 in Philadelphia that promotes knowledge in the humanities and natural sciences through research, professional meetings, publicat ...
, the
American Academy of Arts and Sciences The American Academy of Arts and Sciences (The Academy) is one of the oldest learned societies in the United States. It was founded in 1780 during the American Revolution by John Adams, John Hancock, James Bowdoin, Andrew Oliver, and other ...
, and a Foreign Member of the Royal Society. The
American Mathematical Society The American Mathematical Society (AMS) is an association of professional mathematicians dedicated to the interests of mathematical research and scholarship, and serves the national and international community through its publications, meetings, ...
selected him as the Josiah Willards Gibbs lecturer for 1941. For his work on genetics of evolutionary processes, Wright was awarded the Daniel Giraud Elliot Medal from the National Academy of Sciences in 1945. He died in
Madison, Wisconsin Madison is the List of capitals in the United States, capital city of the U.S. state of Wisconsin. It is the List of municipalities in Wisconsin by population, second-most populous city in the state, with a population of 269,840 at the 2020 Uni ...
, on March 3, 1988.


Family

Wright married Louise Lane Williams (1895–1975) in 1921. They had three children: Richard, Robert, and Elizabeth. Sewall Wright worshipped as a Unitarian.


Scientific achievements and credits


Population genetics

His papers on
inbreeding Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely genetic distance, related genetically. By analogy, the term is used in human reproduction, but more commonly refers to the genet ...
, mating systems, and
genetic drift Genetic drift, also known as random genetic drift, allelic drift or the Wright effect, is the change in the Allele frequency, frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene va ...
make him a principal founder of theoretical population genetics, along with R. A. Fisher and J. B. S. Haldane. Their theoretical work is the origin of the modern evolutionary synthesis or neo-Darwinian synthesis. Wright was the inventor/discoverer of the inbreeding coefficient and F-statistics, standard tools in population genetics. He was the chief developer of the mathematical theory of
genetic drift Genetic drift, also known as random genetic drift, allelic drift or the Wright effect, is the change in the Allele frequency, frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene va ...
, which is sometimes known as the Sewall Wright effect, cumulative stochastic changes in gene frequencies that arise from random births, deaths, and Mendelian segregations in reproduction. In this work he also introduced the concept of
effective population size The effective population size (''N'e'') is the size of an idealised population that would experience the same rate of genetic drift as the real population. Idealised populations are those following simple one- locus models that comply with ass ...
. Wright was convinced that the interaction of
genetic drift Genetic drift, also known as random genetic drift, allelic drift or the Wright effect, is the change in the Allele frequency, frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene va ...
and the other evolutionary forces was important in the process of adaptation. He described the relationship between genotype or phenotype and fitness as fitness surfaces or evolutionary landscapes. On these landscapes mean population fitness was the height, plotted against horizontal axes representing the allele frequencies or the average phenotypes of the population.
Natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
would lead to a population climbing the nearest peak, while
genetic drift Genetic drift, also known as random genetic drift, allelic drift or the Wright effect, is the change in the Allele frequency, frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene va ...
would cause random wandering. He did not accept Fisher's genetic theory of dominance, but instead considered it to arise from biochemical considerations. Although set aside for many years, his interpretation is at the basis of modern ideas of dominance.


Evolutionary theory

Wright's explanation for stasis was that organisms come to occupy adaptive peaks. In order to evolve to another, higher peak, the species would first have to pass through a valley of maladaptive intermediate stages. This could happen by
genetic drift Genetic drift, also known as random genetic drift, allelic drift or the Wright effect, is the change in the Allele frequency, frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene va ...
if the population is small enough. If a species was divided into small populations, some could find higher peaks. If there was some
gene flow In population genetics, gene flow (also known as migration and allele flow) is the transfer of genetic variation, genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent ...
between the populations, these adaptations could spread to the rest of the species. This was Wright's
shifting balance theory The shifting balance theory is a theory of evolution proposed in 1932 by Sewall Wright, suggesting that adaptive evolution may proceed most quickly when a population divides into subpopulations with restricted gene flow. The name of the theory ...
of evolution. There has been much skepticism among evolutionary biologists as to whether these rather delicate conditions hold often in natural populations. Wright had a long-standing and bitter debate about this with R. A. Fisher, who felt that most populations in nature were too large for these effects of genetic drift to be important.


Path analysis

Wright's statistical method of path analysis, which he invented in 1921 and which was one of the first methods using a graphical model, is still widely used in social science. He was a hugely influential reviewer of manuscripts, as one of the most frequent reviewers for ''Genetics''.


Plant and animal breeding

Wright strongly influenced Jay Lush, who was the most influential figure in introducing quantitative genetics into
animal Animals are multicellular, eukaryotic organisms in the Biology, biological Kingdom (biology), kingdom Animalia (). With few exceptions, animals heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, ...
and plant breeding. From 1915 to 1925 Wright was employed by the Animal Husbandry Division of the U.S. Bureau of Animal Husbandry. His main project was to investigate the inbreeding that had occurred in the artificial selection that resulted in the leading breeds of livestock used in American beef production. He also performed experiments with 80,000 guinea pigs in the study of physiological genetics. Furthermore he analyzed characters of some 40,000 guinea pigs in 23 strains of brother-sister matings against a random-bred stock. (Wright 1922a-c). The concentrated study of these two groups of mammals eventually led to the Shifting Balance Theory and the concept of "surfaces of selective value" in 1932. He did major work on the genetics of guinea pigs, and many of his students became influential in the development of mammalian genetics. He appreciated as early as 1917 that genes acted by controlling
enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
s. An anecdote about Wright, disclaimed by Wright himself, describes a lecture during which Wright tucked an unruly guinea pig under his armpit, where he usually held a chalkboard eraser: according to the anecdote, at the conclusion of the lecture, Wright absent-mindedly began to erase the blackboard using the guinea pig.


Statistics

The creation of the
statistical Statistics (from German language, German: ', "description of a State (polity), state, a country") is the discipline that concerns the collection, organization, analysis, interpretation, and presentation of data. In applying statistics to a s ...
coefficient of determination has been attributed to Sewall Wright and was first published in 1921. This metric is commonly employed to evaluate regression analyses in computational statistics and
machine learning Machine learning (ML) is a field of study in artificial intelligence concerned with the development and study of Computational statistics, statistical algorithms that can learn from data and generalise to unseen data, and thus perform Task ( ...
.


Wright and philosophy

Wright was one of the few geneticists of his time to venture into
philosophy Philosophy ('love of wisdom' in Ancient Greek) is a systematic study of general and fundamental questions concerning topics like existence, reason, knowledge, Value (ethics and social sciences), value, mind, and language. It is a rational an ...
. He found a union of concept in Charles Hartshorne, who became a lifelong friend and philosophical collaborator. Wright endorsed a form of
panpsychism In philosophy of mind, panpsychism () is the view that the mind or a mind-like aspect is a fundamental and ubiquitous feature of reality. It is also described as a theory that "the mind is a fundamental feature of the world which exists throug ...
. He believed that the birth of the consciousness was not due to a mysterious property of increasing complexity, but rather an inherent property, therefore implying these properties were in the most elementary particles.


Legacy

Wright and Fisher, along with J.B.S. Haldane, were the key figures in the modern synthesis that brought genetics and evolution together. Their work was essential to the contributions of Dobzhansky, Mayr, Simpson, Julian Huxley, and Stebbins. The modern synthesis was the most important development in evolutionary biology after Darwin. Wright also had a major effect on the development of mammalian genetics and biochemical genetics. Judea Pearl and Dana Mackenzie's '' The Book of Why'' (2018) describes the contribution of Wright's work on path analysis and delays in its acceptance by several technical disciplines (specifically statistics and formal causal analysis). OpenMx has as its icon a representation of Wright's piebald guinea pig.


Bibliography

* ** vol. 1, Genetic & Biometric Foundations. ** vol. 2, Theory of Gene Frequencies. ** vol. 3, Experimental Results and Evolutionary Deductions. ** vol. 4, Variability within and Among Natural Populations.


References


Further reading

* Ghiselin, Michael T. (1997
''Metaphysics and the Origin of Species''
NY: SUNY Press. * * * Wright 1934
"The Method of Path Coefficients"
''Annals of Mathematical Statistics'', 5: 161-215 * * Wright 1983
"Path Analysis in Genetic Epidemiology: A Critique"


External links



by Edric Lescouflair and James F. Crow
Sewall Wright Papers
at the
American Philosophical Society The American Philosophical Society (APS) is an American scholarly organization and learned society founded in 1743 in Philadelphia that promotes knowledge in the humanities and natural sciences through research, professional meetings, publicat ...
* * {{DEFAULTSORT:Wright, Sewall Green 1889 births 1988 deaths American statisticians American Unitarians American evolutionary biologists Fellows of the Econometric Society Foreign members of the Royal Society Bussey Institution alumni History of genetics Lombard College alumni Members of the United States National Academy of Sciences Modern synthesis (20th century) National Medal of Science laureates Panpsychism Population geneticists 20th-century American biologists Members of the American Philosophical Society Presidents of the American Society of Naturalists Theistic evolutionists