|Sir Ronald Fisher|
17 February 1890|
East Finchley, London, England
|Died||29 July 1962
Adelaide, South Australia
|Residence||England and Australia|
|Fields||Statistics, Genetics, and Evolutionary biology|
|Institutions||Rothamsted Research, University College London, Cambridge University, University of Adelaide, Commonwealth Scientific and Industrial Research Organisation|
|Alma mater||University of Cambridge|
|Academic advisors||James Hopwood Jeans and F. J. M. Stratton|
|Doctoral students||C. R. Rao, D. J. Finney, and Walter Bodmer |
Sir Ronald Aylmer Fisher FRS (17 February 1890 – 29 July 1962), known as R.A. Fisher, was an English statistician, evolutionary biologist, mathematician, geneticist, and eugenicist. He began working at Rothamsted Research in 1919, where he developed the analysis of variance (ANOVA) to analyse its immense data from crop experiments since the 1840s, and established his reputation there in the following years as a biostatistician.
Fisher is known as one of the three principal founders of population genetics, creating a mathematical and statistical basis for biology and uniting natural selection with Mendelian genetics, and as one of the chief architects of the modern evolutionary synthesis. He outlined Fisher's principle as well as the Fisherian runaway and sexy son hypothesis theories of sexual selection, and made important contributions to statistics, including the maximum likelihood, fiducial inference, and the derivation of various sampling distributions.
Anders Hald called him "a genius who almost single-handedly created the foundations for modern statistical science", while Richard Dawkins named him "the greatest biologist since Darwin. Not only was he the most original and constructive of the architects of the neo-Darwinian synthesis. Fisher also was the father of modern statistics and experimental design. He therefore could be said to have provided researchers in biology and medicine with their most important research tools, as well as with the modern version of biology's central theorem." and Geoffrey Miller said of him "To biologists, he was an architect of the 'modern synthesis' that used mathematical models to integrate Mendelian genetics with Darwin's selection theories. To psychologists, Fisher was the inventor of various statistical tests that are still supposed to be used whenever possible in psychology journals. To farmers, Fisher was the founder of experimental agricultural research, saving millions from starvation through rational crop breeding programs."
Fisher gained a scholarship to study Mathematics at the University of Cambridge in 1909, gained a First in Astronomy in 1912. In 1915 he published a paper The evolution of sexual preference on sexual selection and mate choice. He published The Correlation Between Relatives on the Supposition of Mendelian Inheritance in 1918, in which he introduced the term variance and proposed its formal analysis. He put forward a genetics conceptual model showing that continuous variation amongst phenotypic traits measured by the biostatisticians could be produced by the combined action of many discrete genes and thus be the result of Mendelian inheritance. This was the first step towards the establishment of population genetics, which demonstrated that natural selection could change allele frequencies in a population, resulting in reconciling its discontinuous nature with gradual evolution. Joan Box, Fisher's biographer and daughter says that Fisher had resolved this problem in 1911.
In 1919 he was offered a position at the Galton Laboratory in University College London led by Karl Pearson, but instead accepted a temporary job at Rothamsted Research in Harpenden to investigate the possibility of analysing the vast amount of data accumulated since 1842 from the "Classical Field Experiments" where he analysed the data recorded over many years and published Studies in Crop Variation in 1921. In 1928 Joseph Oscar Irwin began a three-year stint at Rothamsted and became one of the first people to master Fisher's innovations.
His first application of the analysis of variance was published in 1921.
Fisher's article On a distribution yielding the error functions of several well known statistics (1924) presented Pearson's chi-squared test and William Gosset's Student's t-distribution in the same framework as the Gaussian distribution and where he developed Fisher's z-distribution a new statistical method, commonly used decades later as the F distribution. He pioneered the principles of the design of experiments and the statistics of small samples and the analysis of real data.
In 1925 he published Statistical Methods for Research Workers, one of the 20th century's most influential books on statistical methods. Fisher's method is a technique for data fusion or "meta-analysis" (analysis of analyses)
The Genetical Theory of Natural Selection was first published in 1930 by Clarendon Press. A core work of the neo-Darwinian modern evolutionary synthesis, it helped define population genetics, which Fisher founded alongside Sewall Wright and J. B. S. Haldane, and revived the idea of sexual selection, neglected since Darwin's death. Commonly cited in biology books, it outlines many important concepts, such as:
- Parental investment, is any parental expenditure (time, energy etc.) that benefits one offspring at a cost to parents' ability to invest in other components of fitness,
- Fisherian runaway, explaining how the desire for a phenotypic trait in one sex combined with the trait in the other sex (for example a peacock's tail) creates a runaway development of the trait.
- Fisher's principle, which explains why the sex ratio is mostly 1:1 in nature. *Reproductive value which implies that sexually reproductive value measures the contribution of an individual of a given age to the future growth of the population.
- Fisher's fundamental theorem of natural selection, which states that "the rate of increase in fitness of any organism at any time is equal to its genetic variance in fitness at that time."
- Fisher's geometric model, an evolutionary model of the effect sizes on fitness of spontaneous mutations proposed by Fisher to explain the distribution of effects of mutations that could contribute to adaptative evolution.
- Sexy son hypothesis, which explains why females choose promiscuous, unreliable fathers for their children in the hope of having similar sexy sons who give them lots of grandchildren.
- Mimicry, a similarity of one species to another that protects one or both
- Dominance, a relationship between alleles of one gene, in which the effect on phenotype of one allele masks the contribution of a second allele at the same locus.
- Heterozygote advantage which was later found to play a frequent role in genetic polymorphism.
- Demonstrating that the probability of a mutation increasing the fitness of an organism decreases proportionately with the magnitude of the mutation and that larger populations carry more variation so that they have a greater chance of survival.
In 1935, he published by The Design of Experiments, which was "also fundamental, [and promoted] statistical technique and application... The mathematical justification of the methods was not stressed and proofs were often barely sketched or omitted altogether .... [This] led H.B. Mann to fill the gaps with a rigorous mathematical treatment in his treatise". In this book Fisher also outlined the Lady tasting tea, now a famous design of experiments statistical randomized experiment which uses Fisher's exact test and is the original exposition of Fisher's notion of a null hypothesis.
In his 1937 paper The wave of advance of advantageous genes he proposed Fisher's equation in the context of population dynamics to describe the spatial spread of an advantageous allele and explored its travelling wave solutions.
In 1938 the Fisher–Yates shuffle was described in 1938 by Fisher and Frank Yates in their book Statistical tables for biological, agricultural and medical research. Their description of the algorithm used pencil and paper; a table of random numbers provided the randomness.
He was the first to use diffusion equations to attempt to calculate the distribution of allele frequencies and the estimation of genetic linkage by maximum likelihood methods among populations. In 1950 he published "Gene Frequencies in a Cline Determined by Selection and Diffusion" on the wave of advance of advantageous genes and on clines of gene frequency, being notable as the first application of a computer, the EDSAC, to biology. He developed computational algorithms for analyzing data from his balanced experimental designs, with various editions and translations, becoming a standard reference work for scientists in many disciplines. In ecological genetics he and E. B. Ford showed how the force of natural selection was much stronger than had been assumed, with many ecogenetic situations (such as polymorphism) being maintained by the force of selection.
Fisher publicly spoke out against the 1950 study showing that smoking tobacco causes lung cancer, arguing that correlation does not imply causation. To quote his biographers Yates and Mather, "It has been suggested that the fact that Fisher was employed as consultant by the tobacco firms in this controversy casts doubt on the value of his arguments. This is to misjudge the man. He was not above accepting financial reward for his labours, but the reason for his interest was undoubtedly his dislike and mistrust of puritanical tendencies of all kinds; and perhaps also the personal solace he had always found in tobacco."
He is also known for the following theories:
- Linear discriminant analysis
- Fisher information
- Fisher–Kolmogorov equation
- Fisher–Tippett distribution
- von Mises–Fisher distribution
- Inverse probability
- Fisher–Bingham distribution
- Fisher's permutation test
- Fisher's inequality
In 1910 he joined the Eugenics Society at Cambridge, whose members included John Maynard Keynes, R. C. Punnett, and Horace Darwin. He saw eugenics as addressing a pressing social and scientific issues that encompassed both genetics and statistics. During World War I Fisher started writing book reviews for the Eugenic Review and volunteered to undertake all such reviews for the journal, being hired for a part-time position. The last third of The Genetical Theory of Natural Selection focussed on eugenics, attributing the fall of civilizations to the fertility of their upper classes being diminished, and used British 1911 census data to show an inverse relationship between fertility and social class, partly due, he claimed, to the lower financial costs and hence increasing social status of families with less children. He proposed the abolition of extra allowances to large families, with the allowances proportional to the earnings of the father. He served in several official committees to promote Eugenics. In 1934, he resigned from the Eugenics Society over a dispute about increasing the power of scientists within the movement.
The Race Question
He opposed UNESCO's The Race Question, believing that evidence and everyday experience showed that human groups differ profoundly "in their innate capacity for intellectual and emotional development" and concluded that the "practical international problem is that of learning to share the resources of this planet amicably with persons of materially different nature", and that "this problem is being obscured by entirely well-intentioned efforts to minimize the real differences that exist". The revised statement titled "The Race Concept: Results of an Inquiry" (1951) was accompanied by Fisher's dissenting commentary.
Fisher was born to Kate and George, a successful auctioneer and fine arts dealer, in East Finchley in London, England, one of twins but his elder twin was still-born and grew up the youngest with three sisters and one brother. From 1896 until 1904 they lived at Inverforth House in London, where English Heritage installed a blue plaque in 2002 to mark his residency. Due to poor eyesight, he learnt mathematics without paper and pen, which developed his ability to visualize problems in geometrical terms, but not in writing proper derivations of mathematical solutions, especially proofs. He amazed his peers with his ability to conjecture mathematical solutions without justifying his conclusions by showing intermediate steps.
His mother died from acute peritonitis when he was 14, and his father then lost his business 18 months later. He won the Neeld Medal (a competitive essay in mathematics) at Harrow School age 16 and developed a strong interest in biology, and especially evolutionary biology. In 1909, he won a scholarship to the Gonville and Caius College at the University of Cambridge. On graduating in 1912, his tutor told him that despite his aptitude for scientific work and mathematical potential his disinclination to show calculations or to prove propositions rendered him unsuited for a career in applied mathematics, which required greater thoroughness, giving him a "lukewarm" recommendation, stating that if Fisher "had stuck to the ropes he would have made a first-class mathematician, but he would not."
Unable to join the British Army because of poor eyesight, Fisher worked for six years as a statistician for the City of London. He took up teaching physics and mathematics at a sequence of public schools, at the Thames Nautical Training College and Bradfield College, where, with his new bride, 17 year old Eileen Guinness, with whom he had two sons and six daughters, and her sister, they became subsistence farmers, with a large garden and animals, surviving on very little and living through the last two years of the war without using food coupons. In 1919 he began working at Rothamsted Research.
Leonard Darwin was a great supporter of Fisher. When Fisher was elected to the Royal Society in 1929, Darwin congratulated him. Fisher replied on 25 February 1929, "I knew you would be glad, and your pleasure is as good to me almost as though my own father were still living." Darwin always treated Fisher with tact and generosity. Some years before, after a disagreement, Fisher had resigned from the Royal Statistical Society. Darwin regretted the development and engineered Fisher's re-entry by making him the gift of a life-time subscription to the society. (Letters of 25 and 27 June.) Fisher's 1930 book The Genetical Theory of Natural Selection is dedicated to Darwin. After Darwin's death in 1943 at the age of 93, Fisher wrote to Darwin's niece, Margaret Keynes, "My very dear friend Leonard Darwin... was surely the kindest and wisest man I ever knew". Leonard was the last surviving as well as the longest lived of Charles Darwin's 10 children.
In 1933 he became Professor of Eugenics at University College London (UCL), but in 1939 UCL dissolved the eugenics department, ordering all of the animals to be destroyed. Fisher was dispatched back to Rothamsted with a much reduced staff. In 1943, he took the Balfour Chair of Genetics where the Italian researcher Luigi Luca Cavalli-Sforza was recruited in 1948, establishing a one-man unit of bacterial genetics. Fisher continued his work on mouse chromosome mapping; breeding the mice in laboratories in his own house. In 1947, Fisher cofounded the journal Heredity with Cyril Darlington and in 1949 he published The Theory of Inbreeding.
His fame grew and he began to travel and lecure widely. In 1931, he spent six weeks at the Statistical Laboratory at Iowa State College where he gave three lectures per week, and met many American statisticians, including George W. Snedecor before returning again in 1936. In 1937, he visited the Indian Statistical Institute in Calcutta, and its one part-time employee, P. C. Mahalanobis, often returnings to encouraging its development, being the guest of honour at its 25th anniversary in 1957 when it had 2000 employees.
His marriage disintegrated during World War II and his oldest son George, an aviator, was killed in combat. His daughter and one of his biographers, Joan, married the noted statistician George E. P. Box. Fisher was made a Knight Bachelor by Queen Elizabeth II in 1952 and awarded the Linnean Society of London Darwin–Wallace Medal in 1958. In 1957, a retired Fisher emigrated to Australia where he spent time as a senior research fellow at the Australian Commonwealth Scientific and Industrial Research Organisation (CSIRO) in Adelaide, where he died in 1962, with his remains interred within St Peter's Cathedral.
According to Yates and Mather, "His large family, in particular, reared in conditions of great financial stringency, was a personal expression of his genetic and evolutionary convictions." Fisher was noted for being loyal, and was seen as a patriot, a member of the Church of England, politically conservative, as well as a scientific rationalist. He developed a reputation for carelessness in his dress and was the archetype of the absent-minded professor. H. Allen Orr describes him in the Boston Review as a "deeply devout Anglican who, between founding modern statistics and population genetics, penned articles for church magazines". In a 1955 broadcast on Science and Christianity, he said:
|“||The custom of making abstract dogmatic assertions is not, certainly, derived from the teaching of Jesus, but has been a widespread weakness among religious teachers in subsequent centuries. I do not think that the word for the Christian virtue of faith should be prostituted to mean the credulous acceptance of all such piously intended assertions. Much self-deception in the young believer is needed to convince himself that he knows that of which in reality he knows himself to be ignorant. That surely is hypocrisy, against which we have been most conspicuously warned.||”|
- "Natural selection is a mechanism for generating an exceedingly high degree of improbability."
- Ronald Fisher at the Mathematics Genealogy Project
- Yates, F.; Mather, K. (1963). "Ronald Aylmer Fisher 1890-1962". Biographical Memoirs of Fellows of the Royal Society 9: 91–129. doi:10.1098/rsbm.1963.0006.
- Hald, Anders (1998). A History of Mathematical Statistics. New York: Wiley. ISBN 0-471-17912-4.
- Dawkins, R. (2010). WHO IS THE GREATEST BIOLOGIST SINCE DARWIN? WHY? Edge "Who is the greatest biologist since Darwin? That's far less obvious, and no doubt many good candidates will be put forward. My own nominee would be Ronald Fisher. Not only was he the most original and constructive of the architects of the neo-Darwinian synthesis. Fisher also was the father of modern statistics and experimental design. He therefore could be said to have provided researchers in biology and medicine with their most important research tools, as well as with the modern version of biology's central theorem."
- Miller, Geoffrey (2000). The mating mind: how sexual choice shaped the evolution of human nature, London, Heineman, ISBN 0-434-00741-2 (also Doubleday, ISBN 0-385-49516-1) p.54
- Sir Ronald Aylmer Fisher Published by University of Minnesota
- Fisher, R. A. (1915). "The evolution of sexual preference". Eugenic Review 7 (3): 184–192. PMC 2987134. PMID 21259607.
- The Correlation Between Relatives on the Supposition of Mendelian Inheritance. Ronald A. Fisher. Philosophical Transactions of the Royal Society of Edinburgh. 1918. (volume 52, pages 399–433)
- Box, R. A. Fisher, pp 50–61
- R A Fisher: the life of a scientist Preface
- Parametric statistical theory Pfanzagl, Johann, with the assistance of R. Hamböker 1994 publisher, Walter de Gruyter, Berlin, DE isbn=3-11-013863-8 pages=207–208
- On the "Probable Error" of a Coefficient of Correlation Deduced from a Small Sample. Ronald A. Fisher. Metron, 1: 3-32 (1921)
- Conniffe, Denis 1991. R.A. Fisher and the development of statistics—a view in his centenary year. Journal of the Statistical and Social Inquiry Society of Ireland. 26 (3): pp. 55–108.
- Fisher, R.A. (1925). Statistical Methods for Research Workers. Oliver and Boyd (Edinburgh). ISBN 0-05-002170-2.
- Fisher, R.A.; Fisher, R. A (1948). "Questions and answers #14". The American Statistician 2 (5): 30–31. doi:10.2307/2681650. JSTOR 2681650.
- Grafen, Alan; Ridley, Mark (2006). Richard Dawkins: How A Scientist Changed the Way We Think. New York, New York: Oxford University Press. p. 69. ISBN 0-19-929116-0.
- Sexual Selection and Summary of Population Genetics Accessed from uscs.edu 2-08-2015
- Clutton-Brock, T.H. 1991. The Evolution of Parental Care. Princeton, NJ: Princeton U. Press. pg. 9
- Trivers, R.L. (1972). Parental investment and sexual selection. In B. Campbell (Ed.), Sexual selection and the descent of man, 1871-1971 (pp. 136–179). Chicago, IL: Aldine. ISBN 0-435-62157-2.
- A theory of Fisher's reproductive value Published by PubMed.gov
- The Relation Between Reproductive Value and Genetic Contribution Published by the Genetics journal
- Fisher, R.A. (1930) The Genetical Theory of Natural Selection, Clarendon Press, Oxford
- Fisher, Ronald (1930). The Genetical Theory of Natural Selection. Oxford, UK: Oxford University Press.
- Orr, Allen (2005). "The genetic theory of adaptation: a brief history". Nature Reviews Genetics 6 (2): 119–127. doi:10.1038/nrg1523. PMID 15716908.
- "dominance". Oxford Dictionaries Online. Oxford University Press. Retrieved 14 May 2014.
- "express". Oxford Dictionaries Online. Oxford University Press. Retrieved 14 May 2014.
- Fisher R. 1930. The Genetical Theory of Natural Selection.
- Mann, H.B. (1949). Analysis and design of experiments: Analysis of variance and analysis of variance designs. New York, N. Y.: Dover. MR 32177.
- Fisher 1971, Chapter II. The Principles of Experimentation, Illustrated by a Psycho-physical Experiment, Section 8. The Null Hypothesis.
- OED quote: 1935 R. A. Fisher, The Design of Experiments ii. 19, "We may speak of this hypothesis as the 'null hypothesis', and it should be noted that the null hypothesis is never proved or established, but is possibly disproved, in the course of experimentation."
- Fisher, R. A. (1935) "The fiducial argument in statistical inference", Annals of Eugenics, 8, 391–398.
- R. A. Fisher's Fiducial Argument and Bayes' Theorem by Teddy Seidenfeld
- R. A. Fisher (1936). "The use of multiple measurements in taxonomic problems" (PDF). Annals of Eugenics 7 (2): 179–188. doi:10.1111/j.1469-1809.1936.tb02137.x.
- R. A. Fisher. The wave of advance of advantageous genes, Ann. Eugenics 7:353–369, 1937.
- Fisher, Ronald A.; Yates, Frank (1948) . Statistical tables for biological, agricultural and medical research (3rd ed.). London: Oliver & Boyd. pp. 26–27. OCLC 14222135. Note: the 6th edition, ISBN 0-02-844720-4, is available on the web, but gives a different shuffling algorithm by C. R. Rao.
- Agresti, Alan; David B. Hichcock (2005). "Bayesian Inference for Categorical Data Analysis" (PDF). Statistical Methods & Applications 14 (14): 298. doi:10.1007/s10260-005-0121-y.
- R. A. Fisher, and Balmukand, B. 1928. The estimation of linkage from the offspring of selfed heterozygotes. Journal of Genetics 20:79-92.
- Fisher, R. A. (1950) "Gene Frequencies in a Cline Determined by Selection and Diffusion", Biometrics, 6 (4), 353–361 JSTOR 3001780
- Box, R. A. Fisher, pp 93–166
- Marston, Jean (8 March 2008). "Smoking gun (letter)". New Scientist (2646): 21.
- http://unesdoc.unesco.org/images/0007/000733/073351eo.pdf "The Race Concept: Results of an Inquiry", p. 27. UNESCO 1952
- Fisher biography
- Box, R. A. Fisher, pp 8–16
- Sir John Russell. Letter to The Times of London.
- Box, R. A. Fisher, pp 35–50
- William G. Hill, Trudy F.C. Mackay (1 August 2004). "D. S. Falconer and Introduction to Quantitative Genetics". Genetics 167 (4): 1529–36. PMC 1471025. PMID 15342495.
- Box, R. A. Fisher, p 337
- Box, R. A. Fisher, p 396
- Box, Joan Fisher (1978) R. A. Fisher: The Life of a Scientist Preface, ISBN 0-471-09300-9
- Gould on God: Can religion and science be happily reconciled?
- The Genetical Theory of Natural Selection " It was one of his favorite aphorisms, first reported by Julian Huxley in 1936 and often repeated in Huxley's work (e.g., 1942, 1954) until it finally passed into the language unattributed through the writings of C. H. Waddington, Gavin de Beer, Ernst Mayr, and Richard Dawkins.
- Tommi Jaakkola and David Haussler (1998), Exploiting Generative Models in Discriminative Classifiers. In Advances in Neural Information Processing Systems 11, pages 487–493. MIT Press. ISBN 978-0-262-11245-1 PS, Citeseer
- Aldrich, John (1997). "R.A. Fisher and the making of maximum likelihood 1912–1922". Statistical Science 12 (3): 162–176. doi:10.1214/ss/1030037906.
- Box, Joan Fisher (1978) R. A. Fisher: The Life of a Scientist, New York: Wiley, ISBN 0-471-09300-9.
- William H. Kruskal: "The significance of Fisher: A review of R. A. Fisher. The Life of a Scientist, by Joan Fisher Box," Journal of the American Statistical Association, 75 (1980), 1019–1030.
- David Howie, "Interpreting Probability: Controversies and Developments in the Early Twentieth Century" (Cambridge University Press, 2002)
- Salsburg, David (2002) The Lady Tasting Tea: How Statistics Revolutionized Science in the Twentieth Century, ISBN 0-8050-7134-2
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- O'Connor, John J.; Robertson, Edmund F., "Ronald Fisher", MacTutor History of Mathematics archive, University of St Andrews.
- A Guide to R. A. Fisher by John Aldrich
- University of Adelaide Library for bibliography, biography, 2 volumes of correspondence and many articles
- Classics in the History of Psychology for the first edition of Statistical Methods for Research Workers
- A collection of Fisher quotations compiled by A. W. F. Edwards
Austin Bradford Hill
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