The Genetical Theory of Natural Selection

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First Edition cover

The Genetical Theory of Natural Selection is a book by Ronald Fisher which combines Mendelian genetics with Charles Darwin's theory of natural selection.[1] First published in 1930 by Clarendon Press, it is one of the most important books of the modern evolutionary synthesis,[2] and helped define population genetics. It is commonly cited in biology books, outlining many concepts that are still considered important such as Fisherian runaway, Fisher's principle, Reproductive value, Fisher's fundamental theorem of natural selection, Fisher's geometric model, the sexy son hypothesis, mimicry and the evolution of dominance. It was dictated to his wife in the evenings as he worked at Rothamsted Research in the day.[3]


A second, slightly revised edition was republished in 1958. In 1999, a third variorum 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 the University of Adelaide.


Ronald Fisher

It contains the following chapters:

  1. The Nature of Inheritance
  2. The Fundamental Theorem of Natural Selection
  3. The Evolution of Dominance
  4. Variation as determined by Mutation and Selection
  5. Variation etc.
  6. Sexual Reproduction and Sexual Selection
  7. Mimicry
  8. Man and Society
  9. The Inheritance of Human Fertility
  10. Reproduction in Relation to Social Class
  11. Social Selection of Fertility
  12. Conditions of Permanent Civilization


The peacock plumage is a classic example of the hypothesized Fisherian runaway

In the preface, Fisher considers some general points, including that there must be an understanding of natural selection distinct from that of evolution, and that the then-recent advances in the field of genetics (see history of genetics) now allowed this. In the first chapter, Fisher considers the nature of inheritance, rejecting blending 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 discuss parental investment, Fisher's geometric model, concerning how spontaneous mutations affect biological fitness, Fisher's principle which explains why the sex ratio between males and females is almost always 1:1, Reproductive value, examining the demography of having girl children. Using his knowledge of statistics, the Fisherian runaway, which explores how sexual selection can lead to a positive feedback runaway loop, producing features such as the peacock's plumage. He also wrote about the evolution of dominance, which explores genetic dominance. The last five chapters (8-12) include Fisher's more idiosyncratic views on eugenics.


The book is dedicated to Major Leonard Darwin, Fisher's friend, correspondent and son of Charles Darwin, "In gratitude for the encouragement, given to the author, during the last fifteen years, by discussing many of the problems dealt with in this book".


Reviewed by Charles Galton Darwin, who sent Fisher his copy of the book, with notes in the margin, which initiated a correspondence lasting several years.[4] This book also had a major influence on W. D. Hamilton's theories on the genetic basis of kin selection.

John Henry Bennett gave an account of the writing and reception of the book.[5]

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".[6] J.B.S. Haldane described it as "brilliant".[7] Reginald Punnett was negative, however.[8]

The book was largely overlooked for 40 years, and in particular Fisher's fundamental theorem of natural selection was misunderstood. The work had a great effect on W.D. Hamilton, who discovered it as an undergraduate at the University of Cambridge[9] and noted on 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 Cambridge BA 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 by molecular genetics, Fisher's logic and ideas still underpin most of the ever broadening paths by which Darwinism continues its invasion of human thought.
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.
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...
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 by Laurence Cook ("This is perhaps the most important book on evolutionary genetics ever written"),[10] Brian Charlesworth,[11] James F. Crow[12] and A. W. F. Edwards.[1]

See also[edit]


  1. ^ a b Edwards, A. W. F. (April 2000). "The Genetical Theory of Natural Selection". Genetics (Bethesda, MD: Genetics Society of America) 154 (4): 1419–1426. ISSN 0016-6731. PMC 1461012. PMID 10747041. 
  2. ^ Grafen & Ridley 2006, p. 69
  3. ^ "Sir Ronald Aylmer Fisher". History of Statistics & Probability. Minneapolis, MN: University of Minnesota. Retrieved 2015-11-25. 
  4. ^ Fisher 1999, Appendix 2
  5. ^ Bennett 1983, Introduction
  6. ^ Wright, Sewall (August 1930). "The Genetical Theory of Natural Selection: A Review" (PDF). Journal of Heredity (Oxford, UK: Oxford University Press for the American Genetic Association) 21 (8): 349–356. ISSN 0022-1503. Retrieved 2015-11-25. 
  7. ^ Haldane 1932
  8. ^ Punnett, Reginald (October 18, 1930). "The Genetical Theory of Natural Selection". Nature (Book review) (London: Nature Publishing Group) 126 (3181): 595–597. doi:10.1038/126595a0. ISSN 0028-0836. 
  9. ^ Grafen, Alan (2004). "William Donald Hamilton" (PDF). Biographical Memoirs of Fellows of the Royal Society (London: Royal Society) 50: 109–132. ISSN 0080-4606. Retrieved 2015-11-25. 
  10. ^ Cook, Laurence (March 2000). "The Genetical Theory of Natural Selection — A Complete Variorum Edition". Heredity (Book review) (London: Nature Publishing Group) 84 (3): 390–391. doi:10.1046/j.1365-2540.2000.0713b.x. ISSN 0018-067X. 
  11. ^ 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) (Cambridge, UK: Cambridge University Press) 75 (3): 369–373. ISSN 0016-6723. Retrieved 2015-11-26. 
  12. ^ 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) (Cambridge, MA: Cell Press) 15 (5): 213–214. doi:10.1016/S0169-5347(00)01842-5. ISSN 0169-5347.