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The Genetical Theory of Natural Selection

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Book by Ronald Aylmer Fisher
The Genetical Theory of Natural Selection
First edition title page
AuthorRonald Fisher
LanguageEnglish
SubjectEvolutionary biology
PublisherThe Clarendon Press
Publication date
1930
Publication placeUnited Kingdom
OCLC18500548
575.423
LC ClassQH366 .F5
Preceded byStatistical Methods for Research Workers 
Followed byThe Design of Experiments 

The Genetical Theory of Natural Selection is a book byRonald Fisher which combinesMendelian genetics withCharles Darwin's theory ofnatural selection,[1] with Fisher being the first to argue that "Mendelism therefore validates Darwinism"[2] and stating with regard tomutations that "The vast majority of large mutations are deleterious; small mutations are both far more frequent and more likely to be useful", thus refutingorthogenesis.[3] First published in 1930 byThe Clarendon Press, it is one of the most important books of themodern synthesis,[4] and helped definepopulation genetics. It had been described byJ. F. Crow as the "deepest book on evolution since Darwin".[5]

It is commonly cited in biology books, outlining many concepts that are still considered important such asFisherian runaway,Fisher's principle,reproductive value,Fisher's fundamental theorem of natural selection,Fisher's geometric model, thesexy son hypothesis,mimicry and theevolution of dominance. It was dictated to his wife in the evenings as he worked atRothamsted Research in the day.[6]

History

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In the early 20th century, biologists were discussing on how to reconcile Mendelian genetics with Darwinian natural Section.[7] In England, at this time Ronald Fisher was working at the Rothamsted Experimental,[8] and he developed statistical ways to analyze inheritance and variation in experiments within agriculture. When he applied these methods to evolutionary problems, he showed how Mendelian inheritance could produce the continuous variation observed by Darwin mathematically. This unified selection and hereditary, with the guidance of natural selection, population and mutation. Fisher's book,The Genetical Theory of Selection (1930) emphasized the importance of sexual production and genetic recombination in evolution.[9]

After the publication of his book, it was recognized by the theoretical biologists, and considered important in population genetics. Researchers like,J.B.S. Haldane andSewall Wright expanded on Fisher's framework to describegene frequencies,genetic drift and mutation. The book also influenced later architects of the modern synthesis, likeTheodosius Dobzhansky,Ernst Mayr, andJulian Huxley. The book also contained chapters on human fertility and eugenics which was the reflection of what was socially accepted in the early 20th century in Britain. Scholars in recent years have re-evaluated these sections, and they noted that fisher's eugenic views may be outdated both scientifically and ethically. But his findings understood with the context of what time period it was written and his scientific contributions helped get to the current findings and is very important.[10]

Ronald Fisher

Contents

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The peacock plumage is a classic example of the hypothesizedFisherian runaway.

In the preface, Fisher considers some general points, including that there must be an understanding of natural selection distinct from that ofevolution, and that the then-recent advances in the field ofgenetics (seehistory of genetics) now allowed this. In the first chapter, Fisher considers the nature of inheritance, rejectingblending inheritance, because it would eliminate genetic variance, in favour of particulate inheritance. The second chapter introduces Fisher's fundamental theorem of natural selection. The third considers the evolution of dominance, which Fisher believed was strongly influenced by modifiers. Other chapters discussparental investment, Fisher's geometric model, concerning how spontaneousmutations affectbiological fitness, Fisher's principle which explains why thesex ratio betweenmales andfemales is almost always 1:1, reproductive value, examining thedemography of having girl children. Using his knowledge ofstatistics, the Fisherian runaway, which explores howsexual selection can lead to apositive feedback runaway loop, producing features such as thepeacock'splumage. He also wrote about the evolution of dominance, which exploresgenetic dominance.

Eugenics

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The last five chapters (8-12) include Fisher's concern aboutdysgenics and proposals foreugenics. Fisher attributed the fall ofcivilizations 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 fewer 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.[11][12][13]

Editions

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A second, slightly revised edition was republished in 1958. In 1999, a thirdvariorum edition (ISBN 0-19-850440-3), with the original 1930 text, annotated with the 1958 alterations, notes and alterations accidentally omitted from the second edition was published, edited by professor John Henry Bennett of theUniversity of Adelaide.

Biography

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The Rothamsted Experimental Station, Harpenden, England, where Ronald A. Fisher worked during the 1920s and early 1930s while developing the ideas later published inThe Genetical Theory of Natural Selection

In East Finchley, London, Ronald Aylmer Fisher was born on February 17, 1890. He was the youngest of eight children, including a twin, however, his twin brother passed away at infancy.[14] Tragically, his mother passed away fromperitonitis when he was fourteen, leaving his family in difficult circumstances during his early years.[14] Fisher received his education atHarrow School, where he excelled in mathematics and science.[14] Afterward, he obtained a scholarship to attendGonville and Caius College,Cambridge where he graduated in 1912.[14] He concentrated on mathematics while attending Cambridge, but eventually discovered that biology and statistics sparked his interest.[14]

Following Cambridge, he concentrated on a number of research and academic roles. Fisher established the groundwork for modern statistical theory while employed as astatistician at Rothamsted Experimental Station in Hertfordshire from 1919 to 1933.[14] Additionally, establishing the experiments that subsequently influenced agricultural biology and genetics.[14] In 1933, he accepted the Galton Professorship of Eugenics atUniversity College London, and ten years later was named theArthur Balfour Professor of Genetics atCambridge University.[14]

Friendships and connections with influential biologists were among Fisher's discoveries. Fisher later acknowledged Darwin's influence by dedicatingThe Genetical Theory of Natural Selection to MajorLeonard Darwin, Charles Darwin's son.[14] According to biologistE. B. Ford, Fisher had a significant impact on biological methodology and theory.[15] Ford wrote, “Fisher’s combination of mathematics, statistics, and genetics gave biology a quantitative backbone that was previously lacking” .[15] According to Ford, Fisher's work was occasionally misinterpreted by his peers, but his depth and range of thought won him lasting respect.[15] Following his retirement in 1957, Fisher moved toAdelaide, Australia, where he worked and published articles until his passing on July 29, 1962.[14]

This article is part ofa series on
Eugenics
Historical trajectory

Controversy

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Scholars have long discussed Fisher's bookThe Genetical Theory of Natural Selection because of its comprehensive treatment of eugenics and social policy, despite the fact that it is praised for its groundbreaking findings and unique theories, making it an influential book. According to historian A. Aylward, the book's last five chapters are a reflection on eugenics and the decline of civilizations.[16] The first portion of the book concentrates on the mathematical foundations ofmodern evolutionary theory, which are then applied tohuman civilization in the following chapters.[16]

Fisher argued that disparities infertility between different social classes were the cause of civilization's downfall.[16] Government incentives to promote reproduction among those with higher socioeconomic positions were part of Fisher's proposals.[16] Fisher's evolutionary theory is associated with the eugenic ideology of the early twentieth century, which is now considered discriminatory andpseudoscientific.[16] These texts containracial andclassist assumptions that are at odds with modern ethics and genetics.[16] According to Aylward, proponents have rejected these arguments as careless but ultimately meaningless ideas that have no bearing on the book's scientific validity.[16]

Additionally, Aylward challenges traditional readings of the book's composition.[16] Looking at the amended unpublished typescript from 1919, there are observations that numerous pages of the eugenics material are replicated exactly from this earlier draft.[16] According to Aylward, the book is backwards since Fisher published the eugenics writings before he developed his genetic theory.[16] This suggested that Fisher's political and social theories were conceptual foundations that influenced his research rather than being afterthoughts.[16] This controversy has sparked discussion over how to understand Fisher's legacy, whether as a pioneer in science alone or as a person whose social philosophy and genetics were deeply intertwined.[16]

Legacy

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The significant influence that Ronald A. Fisher had on genetics and heredity did not go unnoticed.[17] He was a cornerstone that advanced the work of Charles Darwin by linking it to Mendelian genetics.[17] Fisher's work also contributed to the development of many genetical studies, as his influence was great among scientists, making him a key figure in modernevolutionary biology.[17]

Reviews

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The book was reviewed byCharles Galton Darwin, who sent Fisher his copy of the book, with notes in the margin, starting a correspondence which lasted several years.[18] The book also had a major influence onW. D. Hamilton's theories on the genetic basis ofkin selection.

John Henry Bennett gave an account of the writing and reception of the book.[19]

Sewall Wright, who had many disagreements with Fisher, reviewed the book and wrote that it was "certain to take rank as one of the major contributions to the theory of evolution."[20]J. B. S. Haldane described it as "brilliant."[21]Reginald Punnett was negative, however.[22]

The book was largely overlooked for 40 years, and in particular Fisher'sfundamental theorem of natural selection was misunderstood. The work had a great effect onW. D. Hamilton, who discovered it as an undergraduate at theUniversity of Cambridge[23] and noted in these excerpts from the rear cover of the 1999 variorum edition:

This is a book which, as a student, I weighed as of equal importance to the entire rest of my undergraduate CambridgeBA course and, through the time I spent on it, I think it notched down my degree. Most chapters took me weeks, some months;...And little modified even bymolecular genetics, Fisher's logic and ideas still underpin most of the ever broadening paths by whichDarwinism continues its invasion of human thought.

For a book that I rate only second in importance in evolution theory to Darwin's "Origin" (this as joined with its supplement "of Man"), and also rate as undoubtedly one of the greatest books of the twentieth century the appearance of a variorum edition is a major event....

Unlike in 1958, natural selection has become part of the syllabus of our intellectual life and the topic is certainly included in every decent course in biology. By the time of my ultimate graduation, will I have understood all that is true in this book and will I get a First? I doubt it. In some ways some of us have overtaken Fisher; in many, however, this brilliant, daring man is still far in front.

The publication of the variorum edition in 1999 led to renewed interest in the work and reviews byLaurence Cook,[24]Brian Charlesworth,[25]James F. Crow,[26] andA. W. F. Edwards.[1]

References

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  1. ^abEdwards, A. W. F. (April 2000)."The Genetical Theory of Natural Selection".Genetics.154 (4). Bethesda, MD:Genetics Society of America:1419–1426.doi:10.1093/genetics/154.4.1419.ISSN 0016-6731.PMC 1461012.PMID 10747041.
  2. ^The Structure of Evolutionary Theory (2002) byStephen Jay Gould, Chapter 7, section "Synthesis as Restriction" Gould quotes Fisher “The whole group of theories which ascribe to hypothetical physiological mechanisms, controlling the occurrence of mutations, a power of directing the course of evolution, must be set aside, once the blending theory of inheritance is abandoned. The sole surviving theory is that of Natural Selection”The Genetical Theory of Natural Selection (1930, p. 20)
  3. ^The Structure of Evolutionary Theory (2002) byStephen Jay Gould, Chapter 7, section "Synthesis as Restriction" Gould further quotes Fisher “For mutations to dominate the trend of evolution it is thus necessary to postulate mutation rates immensely greater than those which are known to occur, and of an order of magnitude which, in general, would be incompatible with particulate inheritance”The Genetical Theory of Natural Selection (1930, p. 20)
  4. ^Grafen & Ridley 2006, p. 69
  5. ^Ford, E. B. (2005)."R. A. Fisher: An Appreciation".Genetics.171 (2):415–417.doi:10.1093/genetics/171.2.415.ISSN 0016-6731.PMC 1456759.PMID 16249595.
  6. ^"Sir Ronald Aylmer Fisher".History of Statistics & Probability. Minneapolis, MN:University of Minnesota. Retrieved2015-11-25.
  7. ^"Charles Darwin",Wikipedia, 2025-10-19, retrieved2025-10-29
  8. ^"Rothamsted Research",Wikipedia, 2025-10-11, retrieved2025-10-29
  9. ^Asher, Robert (August 27, 2025)."Ronald Fisher and group selection".History and Philosophy of the Life Sciences.47 (3) 42.doi:10.1007/s40656-025-00691-5.PMC 12391204.PMID 40864366.
  10. ^Alyward, Alex (October 11, 2021)."R.A. Fisher, eugenics, and the campaign for family allowances in interwar Britain".The British Journal for the History of Science.54 (4):485–505.doi:10.1017/S0007087421000674.
  11. ^"Series 12. Sir Ronald Aylmer Fisher (1890–1962) Statistician and geneticist. Papers 1911–2005. Papers on Eugenics. 1911–1920, 1936". University of Adelaide. Retrieved7 September 2017.
  12. ^Norton, Bernard (27 April 1978)."A 'fashionable fallacy' defended".New Scientist.Fisher worked as he didbecause he was an ardent eugenist. (original italics) ... Careful study of Fisher's writings, moreover, enables one to establish strong connections between the problems that Fisher facedqua eugenist and the work in genetics outlined above.
  13. ^Andrade da Cruz, Rodrigo (1980)."Ronald Fisher and eugenics: Statistics, evolution and genetics in the quest for permanent civilization".Circumscribere: International Journal for the History of Science.19. Pontifical Catholic University of São Paulo, Brazil (PhD Thesis): 53.doi:10.23925/1980-7651.2017v19;p153.
  14. ^abcdefghijYates, Frank; Mather, Kenneth (January 1997)."Ronald Aylmer Fisher, 1890-1962".Biographical Memoirs of Fellows of the Royal Society.9:91–129.doi:10.1098/rsbm.1963.0006.
  15. ^abcFienberg, Stephen E.; Hinkley, David V. (2012-12-06).R.A. Fisher: An Appreciation. Springer Science & Business Media.ISBN 978-1-4612-6079-0.
  16. ^abcdefghijklAylward, Alex (2025-08-20)."A backwards book? Eugenics and the evolution of R. A. Fisher's The genetical theory of natural selection".Notes and Records: The Royal Society Journal of the History of Science.0 20240060.doi:10.1098/rsnr.2024.0060.
  17. ^abcPiegorsch, Walter W. (1990)."Fisher's Contributions to Genetics and Heredity, with Special Emphasis on the Gregor Mendel Controversy".Biometrics.46 (4):915–924.doi:10.2307/2532437.ISSN 0006-341X.JSTOR 2532437.
  18. ^Fisher 1999, Appendix 2
  19. ^Bennett 1983,Introduction
  20. ^Wright, Sewall (August 1930)."The Genetical Theory of Natural Selection: A Review".Journal of Heredity.21 (8). Oxford, UK:Oxford University Press for theAmerican Genetic Association:349–356.doi:10.1093/oxfordjournals.jhered.a103361.ISSN 0022-1503. Retrieved2015-11-25.
  21. ^Haldane 1932
  22. ^Punnett, Reginald (October 18, 1930). "The Genetical Theory of Natural Selection".Nature (Book review).126 (3181). London:Nature Publishing Group:595–597.Bibcode:1930Natur.126..595P.doi:10.1038/126595a0.ISSN 0028-0836.S2CID 4120195.
  23. ^Grafen, Alan (2004)."William Donald Hamilton"(PDF).Biographical Memoirs of Fellows of the Royal Society.50. London:Royal Society:109–132.doi:10.1098/rsbm.2004.0009.ISSN 0080-4606.S2CID 56905497. Retrieved2015-11-25.
  24. ^Cook, Laurence (March 2000)."The Genetical Theory of Natural Selection — A Complete Variorum Edition".Heredity (Book review).84 (3). London: Nature Publishing Group:390–391.Bibcode:2000Hered..84..390C.doi:10.1046/j.1365-2540.2000.0713b.x.ISSN 0018-067X.PMC 2762834.
  25. ^Charlesworth, Brian (2000)."The Genetical Theory of Natural Selection. A Complete Variorum Edition. By R. A. Fisher (edited with foreword and notes by J. H. Bennett). Oxford University Press. 1999. ISBN 0-19-850440-3. xxi+318 pages. Price £25.00".Genetical Research (Book review).75 (3). Cambridge, UK:Cambridge University Press:369–373.doi:10.1017/s0016672300219228.ISSN 0016-6723.
  26. ^Crow, James F. (May 1, 2000). "Second only to Darwin: The Genetical Theory of Natural Selection. A Complete Variorum Edition by R.A. Fisher".Trends in Ecology & Evolution (Book review).15 (5). Cambridge, MA:Cell Press:213–214.Bibcode:2000TEcoE..15..213C.doi:10.1016/S0169-5347(00)01842-5.ISSN 0169-5347.

Bibliography

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