Motoo Kimura

Motoo Kimura (木村資生 Kimura Motoo), (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 geneticist. In genetics, he is famous for his innovative use of diffusion equations to calculate the probability of fixation and time to fixation of beneficial, deleterious, or neutral alleles. Combining theoretical population genetics with molecular evolution data, he also developed the neutral theory of molecular evolution in which genetic drift is the main force changing allele frequencies. James F. Crow, himself a renowned population geneticist, considers Kimura to be one of the two greatest evolutionary geneticists, along with Gustave Malécot, after the great trio of the modern synthesis (Haldane, Wright, Fisher).

Life and work
Kimura was born in Okazaki, Aichi Prefecture. From an early age he was very interested in botany, though he also excelled at mathematics (teaching himself geometry and other maths during a lengthy convalescence due to food poisoning). After entering a selective high school in Nagoya, Kimura focused on plant morphology and cytology; he worked in the laboratory of M. Kumazawa studying the chromosome structure of lilies. With Kumazawa, he also discovered how to connect his interests in botany and mathematics: biometry.

Due to World War II, Kimura left high school early to enter Kyoto Imperial University in 1944. On the advice of the prominent geneticist Hitoshi Kihara, Kimura entered the botany program rather than cytology because the former, in the Faculty of Science rather than Agriculture, allowed him to avoid military duty. He joined Kihara's laboratory after the war, where he studied the introduction of foreign chromosomes into plants and learned the foundations of population genetics. In 1949, Kimura joined the National Institute of Genetics in Mishima. In 1953 he published his first population genetics paper (which would eventually be very influential), describing a "stepping stone" model for population structure that could treat more complex patterns of immigration than Sewall Wright's earlier "island model". After meeting visiting American geneticist Duncan McDonald (part of Atomic Bomb Casualty Commission), Kimura arranged to enter graduate school at Iowa State College in summer 1953 to study with J. L. Lush.

Kimura soon found Iowa State College too restricting; he moved to the University of Wisconsin to work on stochastic models with James F. Crow and join a strong intellectual community of like-minded geneticists, including Newton Morton and most significantly, Sewall Wright. Near the end of his graduate study, Kimura gave a paper at the 1955 Cold Spring Harbor Symposium; though few were able to understand it (both because of mathematical complexity and Kimura's English pronunciation) it received strong praise from Wright and later J. B. S. Haldane. His accomplishments at Wisconsin included a general model for genetic drift, which could accommodate multiple alleles, selection, migration, and mutations, as well as some work based on R. A. Fisher's fundamental theorem of natural selection. He also built on the work of Wright with the Fokker-Planck equation by introducing the Kolmogorov backward equation to population genetics, allowing the calculation of the probability of a gene to become fixed in a population. He received his PhD in 1956, before returning to Japan (where he would remain for the rest of his life, at the National Institute of Genetics).

Kimura worked on a wide spectrum of theoretical population genetics problems, many of them in collaboration with Takeo Maruyama. He introduced the "infinite allele" and "infinite site" models for the study of genetic drift, both of which would be used widely as the field of molecular evolution grew alongside the number of available peptide and genetic sequences. He also created the "ladder model" that could be applied to electrophoresis studies where homologous proteins differ by whole units of charge. He also contributed an important review article on the ongoing controversy over genetic load in 1961.

1968 marked a turning point in Kimura's career. In that year he introduced the neutral theory of molecular evolution, the idea that, at the molecular level, the large majority of genetic change is neutral with respect to natural selection&mdash;making genetic drift a primary factor in evolution. The field of molecular biology was expanding rapidly, and there was growing tension between advocates of the expanding reductionist field and scientists in organismal biology, the traditional domain of evolution. The neutral theory was immediately controversial, receiving support from many molecular biologists and attracting opposition from many evolutionary biologists (though with plenty of divergence from this pattern on both sides, especially among those who studied the theory seriously).

Kimura would spend most of the rest of his life developing and defending the neutral theory. As James F. Crow put it, "much of Kimura's early work turned out to be pre-adapted for use in the quantitative study of neutral evolution." . As new experimental techniques and genetic knowledge became available, Kimura expanded the scope of the neutral theory and created mathematical methods for testing it against the available evidence. In 1973, his student Tomoko Ohta developed a more general version of the theory, the "nearly neutral theory", that could account for high volumes of slightly deleterious mutations. Kimura produced a monograph on the neutral theory in 1983, The Neutral Theory of Molecular Evolution, and also worked to promote the theory through popular writing. Though difficult to test against alternative selection-centered hypotheses, the neutral theory has become part of modern approaches to molecular evolution.

In 1992, Kimura received the Darwin Medal from the Royal Society, and the following year he was made a Foreign Member. He was married to Hiroko Kimura. They had one child, a son, Akio, and had a granddaughter, Hanako.