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J. B. S. Haldane
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J. B. S. Haldane : ウィキペディア英語版
J. B. S. Haldane

| death_place = Bhubaneswar, India
| residence = United Kingdom
| nationality = British (until 1961)
Indian
| field =
| work_institution = University of Cambridge University College London
| alma_mater = University of Oxford
| academic_advisors = Frederick Gowland Hopkins
| doctoral_students = Helen Spurway
Krishna Dronamraju
John Maynard Smith
| known_for =Oparin-Haldane hypothesis
Malaria theory
Population genetics
Enzymology
Neo-Darwinism
| prizes =
| parents = John Scott Haldane
| spouse =
| module =
}}
J. B. S. Haldane, born John Burdon Sanderson Haldane, known as Jack, FRS (; 5 November 1892 – 1 December 1964) (but who used 'J. B. S.' in his printed works), was a British-Indian scientist, well known for his works in physiology, genetics and evolutionary biology, mathematics where he made innovative contributions to statistics and biostatistics. He was son of the equally famous John Scott Haldane. He was a socialist, Marxist, atheist, and humanist whose political dissent led him to leave England in 1956 and live in India, and became a naturalised Indian citizen in 1961.
His first paper in 1915 demonstrated genetic linkage in mammals while subsequent works helped to create population genetics, establishing a unification of Mendelian genetics and Darwinian evolution by natural selection and laying the groundwork for modern evolutionary synthesis. His article on Abiogenesis in 1929 introduced the "Primordial Soup Theory", and is the foundation of modern understanding of the chemical origin of life. Haldane established human gene maps for haemophilia and colour blindness on the X chromosome, and formulated Haldane's rule on sterility in the heterogametic sex of hybrids in species.〔(【引用サイトリンク】title=The Dominance Theory of Haldane's Rule )〕 He correctly proposed that sickle-cell disease gives some immunity to malaria. He was the first to suggest the central idea of ''in vitro fertilisation'', as well as concepts such as hydrogen economy, ''cis trans'', ''coupling'', ''repulsion'', ''darwin'' (as a unit of evolution) and ''cloning''. In 1957 he articulated Haldane's dilemma, a limit on the speed of beneficial evolution which subsequently proved incorrect. He willed his body for medical studies, as he wanted to remain useful even in death.
Arthur C. Clarke credited him as "perhaps the most brilliant science populariser of his generation". Nobel laureate Peter Medawar called Haldane "the cleverest man I ever knew".
==Academic career==
With his sister Naomi Mitchison he started investigating Mendelian genetics in 1908, used guinea pigs and mice, publishing ''Reduplication in mice'' in 1915,〔 demonstrating genetic linkage in mammals, as well as in chickens.
Following his father's footsteps, Haldane's first publication was on the mechanism of gaseous exchange by haemoglobin.〔 and subsequently worked on the properties of blood. He investigated several aspects of kidney functions and mechanism of excretion.
In 1925, with G. E. Briggs, Haldane derived a new interpretation of the enzyme kinetics law described by Victor Henri in 1903, different from the 1913 Michaelis–Menten equation. Leonor Michaelis and Maud Menten assumed that enzyme (catalyst) and substrate (reactant) are in fast equilibrium with their complex, which then dissociates to yield product and free enzyme. The Briggs–Haldane equation was of the same algebraic form, but their derivation is based on the quasi steady state approximation, that is the concentration of intermediate complex (or complexes) does not change. As a result, the microscopic meaning of the "Michaelis Constant" (''Km'') is different. Although commonly referring it as Michaelis–Menten kinetics, most of the current models actually use the Briggs–Haldane derivation.
He was one of the three major figures to develop the mathematical theory of population genetics along with Ronald Fisher and Sewall Wright, and thus playing an important role in the modern evolutionary synthesis, the concept more popularly known as "neo-Darwinism" (popularised by Richard Dawkins' 1976 work titled ''The Selfish Gene''), which re-established natural selection as the premier mechanism of evolution by explaining it in terms of the mathematical consequences of Mendelian inheritance. He wrote a series of ten papers called A Mathematical Theory of Natural and Artificial Selection, on the mathematical theory behind natural selection. It showed the direction and rates of changes of gene frequencies and pioneered the interaction of natural selection with mutation and animal migration. Haldane's book, ''The Causes of Evolution'' (1932), summarised these results, especially in its extensive appendix.
His contributions to statistical human genetics included the first methods using maximum likelihood for estimation of human linkage maps, and pioneering methods for estimating human mutation rates. He was the first to estimate rate of mutation in humans, at 2 × 10−5 mutations per gene per generation for the X-linked haemophilia gene, and to introduce the idea of a "cost of natural selection". At the John Innes Horticultural Institution he developed the complicated linkage theory for polyploids,〔 and extended the idea of gene/enzyme relationships with the biochemical and genetic study of plant pigments. In his essay ''On Being the Right Size'' he outlines Haldane's principle, which states that the size very often defines what bodily equipment an animal must have: "Insects, being so small, do not have oxygen-carrying bloodstreams. What little oxygen their cells require can be absorbed by simple diffusion of air through their bodies. But being larger means an animal must have complicated oxygen pumping and distributing systems to reach all the cells." The conceptual metaphor to animal body complexity has been of use in energy economics and secession ideas.
Haldane introduced the modern concept of abiogenesis in an eight-page article titled ''The origin of life'' (1929), describing the primitive ocean as a "vast chemical laboratory" containing a mixture of inorganic compounds – like a "hot dilute soup" in which organic compounds could have formed. Under the solar energy the oxygenless atmosphere containing carbon dioxide, ammonia and water vapour gave rise to a variety of organic compounds, "living or half-living things". The first molecules reacted with one another to produce more complex compounds, and ultimately the cellular components. At some point a kind of "oily film" was produced that enclosed self-reproducing molecules, thereby becoming the first cell. J.D. Bernal named the hypothesis biopoiesis or biopoesis, the process of living matter evolving from self-replicating but nonliving molecules. The idea was generally dismissed as "wild speculation". In 1924 Alexander Oparin had suggested similar idea in Russian, and in 1936 he introduced it to the English-speaking people. The experimental basis of the theory began in 1953 with the classic Miller–Urey experiment. Since then the primordial soup theory became a dominant theory in the chemistry of life, and often is attributed as Oparin-Haldane hypothesis.〔(【引用サイトリンク】url=http://www.chem.ox.ac.uk/vrchemistry/chapter26/page10.htm )
He proposed in 1949 that genetic disorders in humans living in malaria-endemic regions provide beneficial effects in making people immune to malaria itself, noting that red blood cell mutations such as sickle-cell anemia and various thalassemias were prevalent only in tropical regions where malaria was endemic and were favourable traits for natural selection that prevented individuals from receiving malarial infection. This concept of resistance to malaria was eventually confirmed by A.C. Allison in 1954.

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