Ronald Fisher

From Wikipedia, the free encyclopedia
  (Redirected from Malthusian parameter)
Jump to: navigation, search
Sir Ronald Fisher
R. A. Fischer.jpg
Born (1890-02-17)17 February 1890
East Finchley, London, England
Died 29 July 1962(1962-07-29) (aged 72)
Adelaide, South Australia
Residence England and Australia
Nationality British
Fields Statistics, Genetics, and Evolutionary biology
Alma mater University of Cambridge
Academic advisors Sir James Jeans and F. J. M. Stratton
Doctoral students C. R. Rao, D. J. Finney, and Walter Bodmer [1]
Known for
Influences Leonard Darwin
Notable awards
He was the father-in-law of George E. P. Box.
Not to be confused with Irving Fisher or Michael Fisher.

Sir Ronald Aylmer Fisher FRS[2] (17 February 1890 – 29 July 1962), known as R.A. Fisher, was an English statistician, evolutionary biologist, mathematician, geneticist, and eugenicist. Fisher is known as one of the chief architects of the modern evolutionary synthesis, where he outlined Fisher's principle as well as the Fisherian runaway theory of sexual selection, as one of the three principal founders of population genetics and for his important contributions to statistics, including the analysis of variance (ANOVA), 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",[3] while Richard Dawkins named him "the greatest biologist since Darwin"[4] while 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."[5]

Personal life[edit]

Fisher was born in East Finchley in London, England, to Katie, who died from acute peritonitis when he was 14, and George Fisher, a successful auctioneer and fine arts dealer who then lost his business, 18 months after this tragedy. He had three older sisters and an older brother,[6] and 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 his poor eyesight, he was tutored in mathematics without the aid of paper and pen, which developed his ability to visualize problems in geometrical terms, without contributing to his interest 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. 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. Approaching Fisher's graduation in 1912, his tutor told his student that despite his enormous aptitude for scientific work and his mathematical potential his disinclination to show calculations or to prove propositions rendered him unsuited for a career in applied mathematics, which required greater thoroughness, gaving 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."[7]

Fisher was now eager to join the British Army but failed the medical examinations because of poor eyesight. Over the next six years, he worked as a statistician for the City of London. He took up teaching physics and mathematics at a sequence of public schools, including Bradfield College in Berkshire, as well as aboard H.M. Training Ship Worcester as war work.

He married Eileen Guinness in 1917 when she was only 17 and, with her sister's help, set up a subsistence farming operation on the Bradfield estate, whith a large garden and animals, learning to make do on very little and living through the rest of the war without using their food coupons.[8] 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.

After the end of World War I, Fisher went looking for a new job, calling himself "an egregious failure in two professions", that is, as a commercial statistician and as a teacher.[9] In 1919 he was offered a position at the Galton Laboratory led by Karl Pearson, the founder of mathematical statistics in Great Britain but because he saw the developing rivalry with Pearson as a professional obstacle he instead accepted a temporary job as a statistician at Rothamsted Experimental Station in Harpenden, Hertfordshire, England where he started a major study of the extensive collections of data recorded over many years. This resulted in a series of reports under the general title Studies in Crop Variation.

Stained glass window in the dining hall of Caius College, in Cambridge, commemorating Ronald Fisher and representing a Latin square.

Fisher received the recognition of his peers in 1929 when he was inducted into the Royal Society. His fame grew and he began to travel more and lecture to wider circles. In 1931, he spent six weeks at the Statistical Laboratory at Iowa State College in Ames, Iowa where he gave three lectures per week, and met many of the active American statisticians, including George W. Snedecor before returning for another visit in 1936. In 1933 he left Rothamsted to become a Professor of Eugenics at the University College London (UCL). In 1937, he visited the Indian Statistical Institute in Calcutta, which at the time consisted of one part-time employee, P. C. Mahalanobis. He visited there often in later years, encouraging its development and was the guest of honour at its 25th anniversary in 1957 when it had 2000 employees.[10]

In 1939 UCL tried to dissolve the eugenics department, ordering all of the animals to be destroyed. Fisher fought back, but then he was dispatched back to Rothamsted with a much reduced staff and resources and was then unable to find any suitable war work, and though he kept very busy with various small projects, he became discouraged, his marriage disintegrated and his oldest son George, an aviator,[11] was killed in combat. In 1943, he was offered the Balfour Chair of Genetics at the Cambridge. During the war, this department was almost destroyed, but the University promised him that he would be charged with rebuilding it after the war but the promises were largely unfulfilled, and the department grew slowly, though the Italian researcher Luigi Luca Cavalli-Sforza was recruited in 1948 and established a one-man unit of bacterial genetics. Fisher continued his work on mouse chromosome mapping—breeding the mice in laboratories in his own house—[12] and other projects. These culminated in the publication in 1949 of The Theory of Inbreeding. In 1947, Fisher cofounded the journal Heredity: An International Journal of Genetics with Cyril Darlington.

Fisher eventually received many awards for his work, and was dubbed a Knight Bachelor by Queen Elizabeth II in 1952 and awarded the Linnean Society of London's prestigious Darwin–Wallace Medal in 1958.

Memorial plaque Ronald Aylmer Fisher 1890-1962
Memorial plaque over remains of Ronald Aylmer Fisher, lectern-side aisle of St Peter's Cathedral, Adelaide

In 1957, a retired Fisher emigrated to Australia where he spent time as a senior research fellow at the Australian CSIRO in Adelaide, where he died in 1962, with his remains interred within St Peter's Cathedral.[13]

Fisher was noted for being loyal to a fault to his friends once he had formed a favourable opinion of anyone. A similar sense of loyalty bound him to his culture, being a patriot, a member of the Church of England, politically conservative, as well as a scientific rationalist. Once famous 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".[14] In a 1955 broadcast on Science and Christianity,[2] he said:

Academic achievements[edit]


Fisher was a major figure in 20th century statistics. He was a prominent opponent of Bayesian statistics, and was even the first to use the term "Bayesian".[15] His article On a distribution yielding the error functions of several well known statistics (1924) presented Pearson's chi-squared test and William Gosset's t in the same framework as the Gaussian distribution, and his own parameter in the analysis of variance Fisher's z-distribution, which were more commonly used decades later in the form of the F distribution.

He pioneered the principles of the design of experiments and elaborated his studies of analysis of variance. He furthered his studies of the statistics of small samples and began his systematic approach of the analysis of real data as the springboard for the development of new statistical methods. He developed computational algorithms for analyzing data from his balanced experimental designs. In 1925, this work resulted in the publication of his first book, Statistical Methods for Research Workers,[16] which had many editions and translations, becoming a standard reference work for scientists in many disciplines. In 1935, he published by The Design of Experiments, which was also widely used.

Fisher named and promoted the method of maximum likelihood estimation and originated the concepts of sufficiency, ancillary statistics, Fisher's linear discriminator and Fisher information.


While still an undergraduate, Fisher learned of the newly rediscovered theory of Mendelian genetics, reading a series of articles by Karl Pearson ("Mathematical Contributions to the Theory of Evolution"). He saw biometry and its growing corpus of statistical methods as a potential way to reconcile the discontinuous nature of Mendelian inheritance with continuous variation and gradual evolution, and he published several articles on biometry in the following years. This included the ground-breaking paper "The Correlation Between Relatives on the Supposition of Mendelian Inheritance", written in 1916, published in 1918 and laying the foundation for what came to be known as biometrical genetics, as well as introducing the methodology of the analysis of variance (being the first usage of the term "variance" in statistics, which was a considerable advance over the correlation methods used until then, and showing that the inheritance of traits measurable by real values (i.e., continuous or dimensional traits) is consistent with Mendelian principles.[17] This forms the basis of the genetics of complex trait inheritance and mitigated debates between biometricians and Mendelians, as well as the compatibility of particulate inheritance with natural selection.

His work on the theory of population genetics also made him one of the three great figures of that field, together with Sewall Wright and J. B. S. Haldane, and as such was one of the founders of the neo-Darwinian modern evolutionary synthesis. In addition to founding modern quantitative genetics with his 1918 paper, he was the first to use diffusion equations to attempt to calculate the distribution of gene frequencies among populations. He pioneered the estimation of genetic linkage and gene frequencies by maximum likelihood methods, and wrote early papers on the wave of advance of advantageous genes and on clines of gene frequency. His 1950 paper[18] on gene frequency clines is notable as the first application of a computer, the EDSAC, to biology.[citation needed]

His ground-breaking book The Genetical Theory of Natural Selection was started in 1928 and published in 1930. He developed ideas on sexual selection, including Fisher's principle and the Fisherian runaway, mimicry and the evolution of dominance. He famously showed that the probability of a mutation increasing the fitness of an organism decreases proportionately with the magnitude of the mutation. He also proved that larger populations carry more variation so that they have a larger chance of survival. It was in this book that he set forth the foundations of what was to become known as population genetics. The book was reviewed, among others, by physicist Charles Galton Darwin, a grandson of Charles Darwin's, and following publication of his review, C. G. Darwin sent Fisher his copy of the book, with notes in the margin. The marginal notes became the food for a correspondence running at least three years.[19] Fisher's book also had a major influence on the evolutionary biologist W. D. Hamilton and the development of his later theories on the genetic basis for the existence of kin selection.

Fisher had a long and successful collaboration with E. B. Ford in the field of ecological genetics. The outcome of this work was the general recognition that the force of natural selection was often much stronger than had been appreciated before, and that many ecogenetic situations (such as polymorphism) were not selectively neutral, but were maintained by the force of selection. Fisher was the original author of the idea of heterozygote advantage, which was later found to play a frequent role in genetic polymorphism.[20] The discovery of indisputable cases of natural selection in nature was one of the main strands in the modern evolutionary synthesis.


Fisher as a steward at the First International Eugenics Conference in 1912

Until World War II Fisher was an ardent promoter of eugenics, which also stimulated and guided much of his work in the genetics of humans. In 1910 at Cambridge he joined the Eugenics Society, whose members included John Maynard Keynes, R. C. Punnett, and Horace Darwin (the son of Charles Darwin), seeing the eugenics movement as a pressing social as well as scientific issue that encompassed both genetics and statistics. This group was active, and it held monthly meetings, often featuring addresses by leaders of mainstream eugenics organizations, such as the Eugenics Education Society of London, founded by Charles Darwin's half-cousin, Francis Galton in 1909.[21] The last third of his book The Genetical Theory concerned the applications of these ideas to humans, and presented the data available at that time. He presented a theory that attributed the decline and fall of civilizations to its arrival at a state where the fertility of the upper classes is forced down. Using the census data of 1911 for Britain, he showed that there was an inverse relationship between fertility and social class. This was partly due, he believed, to the rise in social status of families who were not capable of producing many children but who rose because of the financial advantage of having a small number of children. Therefore he proposed the abolition of the economic advantage of small families by instituting subsidies (he called them allowances) to families with larger numbers of children, with the allowances proportional to the earnings of the father. He himself had two sons and six daughters. 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."[2]

Between 1929 and 1934 the Eugenics Society also campaigned hard for a law permitting sterilization on eugenic grounds. They believed that it should be entirely voluntary, and a right, not a punishment. They published a draft of a proposed bill, and it was submitted to Parliament. Although it was defeated by a 2:1 ratio, this was viewed as progress, and the campaign continued. Fisher played a major role in this movement, and served in several official committees to promote it.[citation needed] In 1934, Fisher moved to increase the power of scientists within the Eugenics Society, but was ultimately thwarted by members with an environmentalist point of view[clarification needed (not obvious what "environmentalist" means here)], and he, along with many other scientists, resigned.[citation needed]

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.[22]

An inveterate pipe-smoker, Fisher was opposed to the conclusion that smoking causes lung cancer by Richard Doll and Austin Bradford Hill in their investigation into the health effects of tobacco smoking. He compared the correlations in their papers to a correlation between the import of apples and the rise of divorce in order to show that correlation does not imply causation.[23] To quote his biographers Yates and Mather,[2] "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."


Statistical Methods for Research Workers[edit]

Statistical Methods for Research Workers (ISBN 0-05-002170-2) published in 1925 book, is one of the 20th century's most influential books on statistical methods. According to Conniffe,[24]

Ronald A. Fisher was "interested in application and in the popularization of statistical methods... his early book Statistical Methods for Research Workers, published in 1925, went through many editions and influenced the practical use of statistics in many fields of


The Genetical Theory of Natural Selection[edit]

The Genetical Theory of Natural Selection was started in 1928 and published in 1930, with a second edition in 1950 and a third in 1997 where developed ideas on sexual selection, mimicry and the evolution of dominance. He showed that the probability of a mutation increasing the fitness of an organism decreases proportionately with the magnitude of the mutation. He also proved that larger populations carry more variation and have a larger chance of survival. It was here that he wrote the foundations of what was to become population genetics.

Design of Experiments[edit]

Main article: Design of Experiments

Fisher's Design of Experiments, published in 1935, 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".[24][25]


A selection from Fisher's 395 articles[edit]

These are available on the University of Adelaide website:

Books by Fisher[edit]

Full publication details are available on the University of Adelaide website:

Biographies of Fisher[edit]

Secondary literature[edit]

  • Edwards, A.W.F., 2005, "Statistical methods for research workers" in Grattan-Guinness, I., ed., Landmark Writings in Western Mathematics. Elsevier: 856–70.

See also[edit]


  1. ^ Ronald Fisher at the Mathematics Genealogy Project
  2. ^ a b c d e 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. K.K. edit
  3. ^ Hald, Anders (1998). A History of Mathematical Statistics. New York: Wiley. ISBN 0-471-17912-4. 
  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."
  5. ^ 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
  6. ^ Box, R. A. Fisher, pp 8–16
  7. ^ Sir John Russell. Letter to The Times of London.
  8. ^ Box, R. A. Fisher, pp 35–50
  9. ^ Box, R. A. Fisher, pp 35–36
  10. ^ Box, R. A. Fisher, p 337
  11. ^ Box, R. A. Fisher, p 396
  12. ^ 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. 
  13. ^
  14. ^ Gould on God: Can religion and science be happily reconciled?
  15. ^ 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. 
  16. ^ Box, R. A. Fisher, pp 93–166
  17. ^ Box, R. A. Fisher, pp 50–61
  18. ^ Fisher, R. A. (1950) "Gene Frequencies in a Cline Determined by Selection and Diffusion", Biometrics, 6 (4), 353–361 JSTOR 3001780
  19. ^ Fisher, R. A., 1999. The Genetical Theory of Natural Selection. Complete Variorum Edition. Oxford University Press. Appendix 2.
  20. ^ Fisher R. 1930. The Genetical Theory of Natural Selection.
  21. ^ Box, R. A. Fisher, pp 17–34
  22. ^ "The Race Concept: Results of an Inquiry", p. 27. UNESCO 1952
  23. ^ Marston, Jean (8 March 2008). "Smoking gun (letter)". New Scientist (2646): 21. 
  24. ^ a b c 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.
  25. ^ Mann, H.B. (1949). Analysis and design of experiments: Analysis of variance and analysis of variance designs. New York, N. Y.: Dover. MR 32177. 


External links[edit]

Academic offices
Preceded by
Austin Bradford Hill
Presidents of the Royal Statistical Society
Succeeded by
William Piercy