Fisher's fundamental theorem of natural selection
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Fisher's fundamental theorem of natural selection is an idea about
genetic variance Genetic variance is a concept outlined by the England, English biologist and statistics, statistician Ronald Fisher in his Fisher's fundamental theorem of natural selection, fundamental theorem of natural selection. In his 1930 book ''The Genetica ...
in
population genetics Population genetics is a subfield of genetics that deals with genetic differences within and between populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as adaptation, speciation, and pop ...
developed by the
statistician A statistician is a person who works with theoretical or applied statistics. The profession exists in both the private and public sectors. It is common to combine statistical knowledge with expertise in other subjects, and statisticians may wor ...
and
evolutionary biologist Evolutionary biology is the subfield of biology that studies the evolutionary processes (natural selection, common descent, speciation) that produced the diversity of life on Earth. It is also defined as the study of the history of life for ...
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 ...
. The proper way of applying the abstract mathematics of the theorem to actual biology has been a matter of some debate. It states: :"The rate of increase in fitness of any
organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and ...
at any time is equal to its genetic variance in fitness at that time." Or in more modern terminology: :"The rate of increase in the mean fitness of any organism, at any time, that is ascribable to natural selection acting through changes in
gene frequencies Allele frequency, or gene frequency, is the relative frequency of an allele (variant of a gene) at a particular locus in a population, expressed as a fraction or percentage. Specifically, it is the fraction of all chromosomes in the population that ...
, is exactly equal to its genetic variance in fitness at that time".


History

The theorem was first formulated in Fisher's 1930 book ''
The Genetical Theory of Natural Selection ''The Genetical Theory of Natural Selection'' is a book by Ronald Fisher which combines Mendelian genetics with Charles Darwin's theory of natural selection, with Fisher being the first to argue that "Mendelism therefore validates Darwinism" and ...
''. Fisher likened it to the law of entropy in
physics Physics is the natural science that studies matter, its fundamental constituents, its motion and behavior through space and time, and the related entities of energy and force. "Physical science is that department of knowledge which r ...
, stating that "It is not a little instructive that so similar a law should hold the supreme position among the biological sciences". The model of
quasi-linkage equilibrium Quasi-linkage equilibrium (QLE) is a mathematical approximation used in solving population genetics problems. Motoo Kimura introduced the notion to simplify a model of Fisher's fundamental theorem. QLE greatly simplifies population genetic equati ...
was introduced by
Motoo Kimura (November 13, 1924 – November 13, 1994) was a Japanese biologist best known for introducing the neutral theory of molecular evolution in 1968. He became one of the most influential theoretical population geneticists. He is remembered in geneti ...
in 1965 as an approximation in the case of weak selection and weak
epistasis Epistasis is a phenomenon in genetics in which the effect of a gene mutation is dependent on the presence or absence of mutations in one or more other genes, respectively termed modifier genes. In other words, the effect of the mutation is dep ...
. Largely as a result of Fisher's feud with the American geneticist
Sewall Wright Sewall Green Wright FRS(For) Honorary FRSE (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 alongsi ...
about adaptive landscapes, the theorem was widely misunderstood to mean that the average fitness of a population would always increase, even though models showed this not to be the case. In 1972,
George R. Price George Robert Price (October 6, 1922 – January 6, 1975) was an American population geneticist. Price is often noted for his formulation of the Price equation in 1967. Originally a physical chemist and later a science journalist, he moved ...
showed that Fisher's theorem was indeed correct (and that Fisher's proof was also correct, given a typo or two), but did not find it to be of great significance. The sophistication that Price pointed out, and that had made understanding difficult, is that the theorem gives a formula for part of the change in gene frequency, and not for all of it. This is a part that can be said to be due to natural selection. Due to confounding factors, tests of the fundamental theorem are quite rare though Bolnick in 2007 did test this effect in a natural population.


References


Further reading

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External links

* {{Population genetics Population genetics Biological theorems Evolutionary dynamics Modern synthesis (20th century)