Motoo Kimura
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(November 13, 1924 – November 13, 1994) was a
Japanese Japanese may refer to: * Something from or related to Japan, an island country in East Asia * Japanese language, spoken mainly in Japan * Japanese people, the ethnic group that identifies with Japan through ancestry or culture ** Japanese diaspor ...
biologist A biologist is a scientist who conducts research in biology. Biologists are interested in studying life on Earth, whether it is an individual Cell (biology), cell, a multicellular organism, or a Community (ecology), community of Biological inter ...
best known for introducing the
neutral theory of molecular evolution The neutral theory of molecular evolution holds that most evolutionary changes occur at the molecular level, and most of the variation within and between species are due to random genetic drift of mutant alleles that are selectively neutral. The ...
in 1968. He became one of the most influential theoretical
population geneticists 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 ...
. He is remembered in genetics for his innovative use of diffusion equations to calculate the probability of fixation of beneficial, deleterious, or neutral
allele An allele (, ; ; modern formation from Greek ἄλλος ''állos'', "other") is a variation of the same sequence of nucleotides at the same place on a long DNA molecule, as described in leading textbooks on genetics and evolution. ::"The chro ...
s. Combining theoretical population genetics with
molecular evolution Molecular evolution is the process of change in the sequence composition of cellular molecules such as DNA, RNA, and proteins across generations. The field of molecular evolution uses principles of evolutionary biology and population genetics ...
data, he also developed the neutral theory of molecular evolution in which
genetic drift Genetic drift, also known as allelic drift or the Wright effect, is the change in the frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene variants to disappear completely and there ...
is the main force changing
allele 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 ...
. James F. Crow, himself a renowned population geneticist, considered Kimura to be one of the two greatest evolutionary geneticists, along with
Gustave Malécot Gustave Malécot (28 December 1911 – November 1998) was a French mathematician whose work on heredity had a strong influence on population genetics. Biography Malécot grew up in L'Horme, a small village near St. Étienne in the Loire ...
, after the great trio of the
modern synthesis Modern synthesis or modern evolutionary synthesis refers to several perspectives on evolutionary biology, namely: * Modern synthesis (20th century), the term coined by Julian Huxley in 1942 to denote the synthesis between Mendelian genetics and s ...
,
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 ...
,
J. B. S. Haldane John Burdon Sanderson Haldane (; 5 November 18921 December 1964), nicknamed "Jack" or "JBS", was a British-Indian scientist who worked in physiology, genetics, evolutionary biology, and mathematics. With innovative use of statistics in biolog ...
, and
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 ...
.


Life and work

Kimura was born in Okazaki,
Aichi Prefecture is a prefecture of Japan located in the Chūbu region of Honshū. Aichi Prefecture has a population of 7,552,873 () and a geographic area of with a population density of . Aichi Prefecture borders Mie Prefecture to the west, Gifu Prefectur ...
. 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 is the largest city in the Chūbu region, the fourth-most populous city and third most populous urban area in Japan, with a population of 2.3million in 2020. Located on the Pacific coast in central Honshu, it is the capital and the most pop ...
, Kimura focused on plant morphology and cytology; he worked in the laboratory of M. Kumazawa studying the
chromosome structure Eukaryotic chromosome structure refers to the levels of packaging from raw DNA molecules to the chromosomal structures seen during metaphase in mitosis or meiosis. Chromosomes contain long strands of DNA containing genetic information. Compared to ...
of
lilies ''Lilium'' () is a genus of herbaceous flowering plants growing from bulbs, all with large prominent flowers. They are the true lilies. Lilies are a group of flowering plants which are important in culture and literature in much of the world. M ...
. With Kumazawa, he also discovered how to connect his interests in botany and mathematics:
biometry Biostatistics (also known as biometry) are the development and application of statistical methods to a wide range of topics in biology. It encompasses the design of biological experiments, the collection and analysis of data from those experimen ...
. Due to
World War II World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the vast majority of the world's countries—including all of the great powers—forming two opposin ...
, Kimura left high school early to enter
Kyoto Imperial University , mottoeng = Freedom of academic culture , established = , type = Public (National) , endowment = ¥ 316 billion (2.4 billion USD) , faculty = 3,480 (Teaching Staff) , administrative_staff = 3,978 (Total Staff) , students = ...
in 1944. On the advice of the prominent geneticist
Hitoshi Kihara was a Japanese geneticist known for his work on the genetics of wheat. Biography Hitoshi Kihara was born on 21 October 1893 in Tokyo, Japan. He graduated from the Faculty of Agriculture, Hokkaido Imperial University and served as a professor a ...
, 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 The National Institute of Genetics ("Japanese Institute of Genetics") is a Japanese institution founded in 1949. It hosts the DNA Data Bank of Japan The DNA Data Bank of Japan (DDBJ) is a biological database that collects DNA sequences. It i ...
in
Mishima, Shizuoka Mishima City Hall is a city located in eastern Shizuoka Prefecture, Japan. , the city had an estimated population of 109,803 in 49,323 households, and a population density of 1800 persons per km². The total area of the city is . Geography ...
. 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 migration than
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 ...
's earlier "island model". After meeting visiting American geneticist Duncan McDonald (part of the
Atomic Bomb Casualty Commission The Atomic Bomb Casualty Commission (ABCC) ( Japanese:原爆傷害調査委員会, ''Genbakushōgaichōsaiinkai'') was a commission established in 1946 in accordance with a presidential directive from Harry S. Truman to the National Academy of ...
), Kimura arranged to enter graduate school at
Iowa State College Iowa State University of Science and Technology (Iowa State University, Iowa State, or ISU) is a public land-grant research university in Ames, Iowa. Founded in 1858 as the Iowa Agricultural College and Model Farm, Iowa State became one of the n ...
in the 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 to join a strong intellectual community of like-minded geneticists, including Newton Morton and most significantly,
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 ...
. Near the end of his graduate study, Kimura gave a paper at the 1955
Cold Spring Harbor Symposium Cold Spring Harbor Laboratory (CSHL) is a private, non-profit institution with research programs focusing on cancer, neuroscience, plant biology, genomics, and quantitative biology. It is one of 68 institutions supported by the Cancer Centers ...
; 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 John Burdon Sanderson Haldane (; 5 November 18921 December 1964), nicknamed "Jack" or "JBS", was a British-Indian scientist who worked in physiology, genetics, evolutionary biology, and mathematics. With innovative use of statistics in biolog ...
. 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 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 ...
's
fundamental theorem of natural selection Fisher's fundamental theorem of natural selection is an idea about genetic variance in population genetics developed by the statistician and evolutionary biologist Ronald Fisher. The proper way of applying the abstract mathematics of the theorem t ...
. He also built on the work of Wright with the Fokker–Planck equation by introducing the
Kolmogorov backward equation In probability theory, Kolmogorov equations, including Kolmogorov forward equations and Kolmogorov backward equations, characterize continuous-time Markov processes. In particular, they describe how the probability that a continuous-time Markov pr ...
to population genetics, allowing the calculation of the probability of an allele to become
fixed Fixed may refer to: * ''Fixed'' (EP), EP by Nine Inch Nails * ''Fixed'', an upcoming 2D adult animated film directed by Genndy Tartakovsky * Fixed (typeface), a collection of monospace bitmap fonts that is distributed with the X Window System * ...
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 Takeo may refer to: *Takéo Province, a province of Cambodia **Doun Kaev (town), formerly known as Takéo, the capital of Takéo province *Ta Keo, an Angkorian temple in Cambodia *Takeo, Saga, a city in Saga Prefecture, Japan *Takeo (given name), a ...
. He introduced the " infinite alleles", " infinite sites", and " stepwise" models of mutation, all of which would be used widely as the field of
molecular evolution Molecular evolution is the process of change in the sequence composition of cellular molecules such as DNA, RNA, and proteins across generations. The field of molecular evolution uses principles of evolutionary biology and population genetics ...
grew alongside the number of available
peptide Peptides (, ) are short chains of amino acids linked by peptide bonds. Long chains of amino acids are called proteins. Chains of fewer than twenty amino acids are called oligopeptides, and include dipeptides, tripeptides, and tetrapeptides. A ...
and
genetic sequence A nucleic acid sequence is a succession of bases signified by a series of a set of five different letters that indicate the order of nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. By convention, sequences are us ...
s. The stepwise mutation model is a "ladder model" that can be applied to electrophoresis studies where homologous proteins differ by whole units of charge. An early statement of his approach was published in 1960, in his ''An Introduction to Population Genetics''. He also contributed an important review article on the ongoing controversy over
genetic load Genetic load is the difference between the fitness of an average genotype in a population and the fitness of some reference genotype, which may be either the best present in a population, or may be the theoretically optimal genotype. The average i ...
in 1961. 1968 marked a turning point in Kimura's career. In that year he introduced the
neutral theory of molecular evolution The neutral theory of molecular evolution holds that most evolutionary changes occur at the molecular level, and most of the variation within and between species are due to random genetic drift of mutant alleles that are selectively neutral. The ...
, the idea that, at the molecular level, the large majority of genetic change is neutral with respect to
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 heritable traits characteristic of a population over generations. Charle ...
—making
genetic drift Genetic drift, also known as allelic drift or the Wright effect, is the change in the frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene variants to disappear completely and there ...
a primary factor in
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
. The field of
molecular biology Molecular biology is the branch of biology that seeks to understand the molecular basis of biological activity in and between cells, including biomolecular synthesis, modification, mechanisms, and interactions. The study of chemical and physi ...
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. Kimura spent the rest of his life developing and defending the neutral theory. As James Crow put it, "much of Kimura's early work turned out to be
pre-adapted Exaptation and the related term Co-option (biology), co-option describe a shift in the function of a trait during evolution. For example, a trait can evolve because it served one particular function, but subsequently it may come to serve another. E ...
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. Kimura produced a monograph on the neutral theory in 1983, ''
The Neutral Theory of Molecular Evolution ''The Neutral Theory of Molecular Evolution'' is an influential monograph written in 1983 by Japanese evolutionary biologist Motoo Kimura. While the neutral theory of molecular evolution existed since his article in 1968, Kimura felt the need to ...
'', and also worked to promote the theory through popular writings such as ''My Views on Evolution'', a book that became a best-seller in Japan. 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 The Darwin Medal is one of the medals awarded by the Royal Society for "distinction in evolution, biological diversity and developmental, population and organismal biology". In 1885, International Darwin Memorial Fund was transferred to the ...
from 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 ...
, and the following year he was made a
Foreign Member of the Royal Society Fellowship of the Royal Society (FRS, ForMemRS and HonFRS) is an award granted by the judges of the Royal Society of London to individuals who have made a "substantial contribution to the improvement of natural knowledge, including mathematic ...
. Kimura suffered from progressive weakening caused by amyotrophic lateral sclerosis later in life. After an accidental fall at his home in Shizuoka, Japan, Kimura struck his head and died on November 13, 1994, of a cerebral hemorrhage. He was married to Hiroko Kimura. They had one child, a son, Akio, and a granddaughter, Hanako. Brenner's Encyclopedia of Genetics, 2nd Edition

/ref>


Honors

* 1959 – Genetics Society of Japan Prize * 1965 –
Weldon Memorial Prize The Weldon Memorial Prize, also known as the Weldon Memorial Prize and Medal, is given yearly by the University of Oxford. The prize is to be awarded without regard to nationality or membership of any University to the person who, in the judgeme ...
, Oxford * 1968 – Japan Academy Prize * 1973 – Foreign member of the
National Academy of Sciences of the USA The National Academy of Sciences (NAS) is a United States nonprofit, 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 Natio ...
* 1976 – Person of Cultural Merit * 1976 –
Order of Culture The is a Japanese order, established on February 11, 1937. The order has one class only, and may be awarded to men and women for contributions to Japan's art, literature, science, technology, or anything related to culture in general; recipient ...
* 1982 – Member of the
Japan Academy The Japan Academy ( Japanese: 日本学士院, ''Nihon Gakushiin'') is an honorary organisation and science academy founded in 1879 to bring together leading Japanese scholars with distinguished records of scientific achievements. The Academy is ...
* 1986 – Chevalier de l'Ordre Nationale de Merite * 1986 –
Asahi Prize The , established in 1929, is an award presented by the Japanese newspaper ''Asahi Shimbun'' and Asahi Shimbun Foundation to honor individuals and groups that have made outstanding accomplishments in the fields of arts and academics and have greatl ...
* 1987 – John J. Carty Award of the
National Academy of Sciences The National Academy of Sciences (NAS) is a United States nonprofit, 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 Nati ...
in evolutionary biology * 1988 –
International Prize for Biology The is an annual award for "outstanding contribution to the advancement of research in fundamental biology." The Prize, although it is not always awarded to a biologist, is one of the most prestigious honours a natural scientist can receive. Ther ...
* 1992 –
Darwin Medal The Darwin Medal is one of the medals awarded by the Royal Society for "distinction in evolution, biological diversity and developmental, population and organismal biology". In 1885, International Darwin Memorial Fund was transferred to the ...
Royal Society
archived record
/ref> * 1993 – Foreign member of
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 ...


See also

*
History of biology The history of biology traces the study of the living world from ancient to modern times. Although the concept of ''biology'' as a single coherent field arose in the 19th century, the biological sciences emerged from traditions of medicine a ...
*
History of evolutionary thought Evolutionary thought, the recognition that species change over time and the perceived understanding of how such processes work, has roots in antiquity—in the ideas of the ancient Greeks, Romans, Chinese, Church Fathers as well as in medie ...
*
History of molecular biology The history of molecular biology begins in the 1930s with the convergence of various, previously distinct biological and physical disciplines: biochemistry, genetics, microbiology, virology and physics. With the hope of understanding life at its m ...
*
Molecular evolution Molecular evolution is the process of change in the sequence composition of cellular molecules such as DNA, RNA, and proteins across generations. The field of molecular evolution uses principles of evolutionary biology and population genetics ...


References

{{DEFAULTSORT:Kimura, Motoo 1924 births 1994 deaths Kyoto University alumni University of Wisconsin–Madison alumni People from Okazaki, Aichi Population geneticists Evolutionary biologists Theoretical biologists Japanese scientists Japanese biologists Japanese molecular biologists Japanese geneticists Foreign Members of the Royal Society Foreign associates of the National Academy of Sciences Recipients of the Order of Culture People with motor neuron disease 20th-century biologists Neutral theory