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The Genetical Theory of Natural Selection

2000/04/01 by A. W. F. Edwards · 2 citations
Arts and Humanities · Biochemistry, Genetics and Molecular Biology · Psychology · #Philosophy and History of Science #Evolution and Science Education #Evolution and Genetic Dynamics #Genius #Nothing #Neglect #Natural selection #Natural (archaeology) #Darwin (ADL) #Epistemology #Selection (genetic algorithm) #Biology #Work (physics) #Philosophy #Literature #Psychology #Computer science #Paleontology #Art

paper · pdf · doi:10.1093/genetics/154.4.1419

openalex publication_date 2000/04/01 · openalex created_date 2025/10/10 · openalex updated_date 2026/07/30

Abstract

So melancholy a neglect of Darwin's work suggests reflections upon the use of those rare and precious possessions of man—great books. It was, we believe, the custom of the late Professor Freeman to warn his students that mastery of one great book was worth any amount of knowledge of many lesser ones. The tendency of modern scientific teaching is to neglect the great books, to lay far too much stress upon relatively unimportant modern work, and to present masses of detail of doubtful truth and questionable weight in such a way as to obscure principles … . How many biological students of today have read The Origin? The majority know it only from extracts, a singularly ineffective means, for a work of genius does not easily lend itself to the scissors; its unity is too marked. Nothing can really take the place of a first-hand study of the work itself. R. A. Fisher and C. S. Stock (1915) [Fisher was 25 years old and his student friend Stock 27 in 1915. Professor E. A. Freeman was Regius Professor of Modern History at Oxford, 1884–1892.] ON May 14, 1929, R. A. Fisher wrote to Oxford University Press, “I should call the book something like The Genetical Theory of Natural Selection.” Within the year Fisher's (1930) book had been published, and in May 2000 we can celebrate its seventieth birthday. Last year Oxford republished in facsimile the 1930 edition as a Variorum Edition, edited with a foreword, notes, and appendices by Henry Bennett. Once again, the book, which J. B. S. Haldane (1932) described as “brilliant” and Sewall Wright (1930) as “certain to take rank as one of the major contributions to the theory of evolution,” is in print. In the present panegyric I assume some familiarity with The Genetical Theory. The indefatigable scholarship of Professor Bennett has left us with two detailed descriptions of the book, his Variorum Edition foreword and an earlier essay in his volume of Fisher's biological correspondence (Bennett 1971–74, 1983), but, like The Origin of Species, The Genetical Theory cannot be appreciated second hand. The book is not long; the main text of the 1930 edition has 265 pages of 13-point lines. (“Fairly large print is a real antidote to stiff reading” wrote Fisher to his Oxford editor—all quotations from Fisher's correspondence are from Bennett 1983.) Bennett opens his foreword with the following paragraph: The genetical theory of natural selection is celebrated as the first major work to provide a synthesis of Darwinian selection and Mendelian genetics. Its publication in 1930 marked a turning point in the development of evolutionary thought, contributing fundamentally to the renaissance of Darwinism following a long period of neglect. It is a work remarkable for what it reveals of Fisher's creative genius and his insight into many of the problems facing evolutionary biologists today. Bennett (1983) In this Perspectives series, Crow (1990a) summarized the main themes of The Genetical Theory (“arguably the deepest and most influential book on evolution since Darwin”), and in another celebration of the Fisher centenary (Crow 1990b), he gave this succinct account: [In addition to enunciating the “fundamental theorem of natural selection,” Fisher] developed a totally novel way (now standard) for determining the probability of survival of a mutant gene. He worked out the partial differential equation for gene-frequency change, using a trigonometric transformation that made the variance independent of the allele frequencies. He generalized Haldane's formula for the probability of survival of a mutant gene, making it possible to treat deleterious as well as favorable mutants. The formula, further improved by Malécot and Kimura, plays a crucial role in the analytical treatment of the neutral theory of molecular evolution. He worked out the stationary distribution of allele frequencies. He gave us the first quantitative theories of sexual selection, mimicry, polymorphism, evolution of recombination rates, and supergenes. He explained why the sex ratio is nearly 1:1 even in highly polygamous species—a problem that baffled Darwin and that provides one of the best illustrations that natural selection does R. A. Fisher, about 1932. not always do what is best for the species. In doing this, Fisher introduced the concept of parental expenditure, thereby seeding a cloudburst of ecological literature. Rarely have so many new and deep ideas been put into a single book. Crow (1990b, p. 270) I discuss the problem of the sex ratio below, as well as mention further themes in The Genetical Theory, such as kin selection. The author: Ronald Aylmer Fisher was born in London, England, in 1890, and died in Adelaide, Australia, in 1962. In 1928–1929, when he was writing The Genetical Theory, Fisher was head of the statistics department at Rothamsted Experimental Station, already taking over from Karl Pearson the mantle of Britain's leading statistician. His Statistical Methods for Research Workers, which was to revolutionize the application of statistics, had appeared in 1925 and was now in its second edition (Fisher 1925). By mid-century, when his collection of papers Contributions to Mathematical Statistics (Fisher 1950) was published, it was clear that Fisher was the leading world figure in statistics, and as the century turns it is now generally accepted by historians of the subject that his contributions place him alongside Laplace (1749–1827) and Gauss (1777–1855). Hald (1998), the foremost historian of mathematical statistics, writes, “Fisher was a genius who almost single-handedly created the foundations for modern statistical science … .” Like Laplace and Gauss, Fisher also made outstanding contributions to science outside the fields of probability and statistics. While Laplace and Gauss contributed to the physical sciences such as astronomy and geodesy (not to mention pure mathematics itself), Fisher used his mathematical gifts to till the fertile ground of biology. C. G. Darwin (1930)—Sir Charles Darwin, physicist, grandson of Charles Darwin—opened his review of The Genetical Theory plaintively with the words, “When a man reveals himself as a master of two very different subjects it is hard to find a critic of his work.” How did Fisher, the equal of Laplace and Gauss in statistics, become “the greatest of Darwin's successors” (Dawkins 1986) as well? Joan Box in her superb biography of her father (Box 1978) tells a story about Fisher's biology teacher at Harrow School, Arthur Vassall, and the influence he had on the boy. Vassall once told E. B. Ford, in answer to his invitation to name the 10 or 12 cleverest boys he had taught, that it would be difficult to do so, but in terms of sheer brilliance he could divide them into two groups, Fisher in one and all the rest in the other. Writing to Vassall in 1929, as The Genetical Theory approached completion, Fisher said, “The fact is that nearly all my statistical work is based on biological material and much of it has been undertaken merely to clear up difficulties in experimental technique.” Fisher's early interest in natural history is reflected in the books chosen for school prizes, leading up to, in his last year, the complete works of Charles Darwin in 13 volumes. “He went up to Cambridge in possession of volumes he was to read and re-read with loving care throughout his life” (Box 1978). There was some uncertainty as to whether Fisher should try for a Cambridge scholarship in mathematics or in biology, but in the end mathematics won. When he arrived at Cambridge with a scholarship in 1909, the University had just celebrated the Darwin centenary and the fiftieth anniversary of the publication of On the Origin of Species (Darwin 1859). In connection with the centenary, an anonymous benefactor had endowed a Professorship of Biology at Cambridge “to be devoted to that branch of Biology now entitled Genetics (Heredity and Variation),” and William Bateson, who had coined the very word “genetics” only 3 years previously, had been elected to it in 1908. Bateson's (1909) book Mendel's Principles of Heredity, containing an English translation of Mendel's paper, and Francis Darwin's edition of his father's unpublished essays of 1842 and 1844 (Darwin 1909) were being printed by Cambridge University Press at one end of King's Parade just as the young mathematician-biologist entered Gonville and Caius College at the other end. Fisher bought a copy of Bateson (Fisher 1952), later remarking, “The new school of geneticists using Mendel's laws of inheritance was full of activity and confidence, and the shops were full of books good and bad from which one could see how completely many writers of this movement believed that Darwin's position had been discredited” (Fisher 1947). Fisher the mathematician developed and retained the highest opinion of biologists' work. In the Preface to The Genetical Theory he remarks: The types of mind which result from training in mathematics and in biology certainly differ but the does not to in the It would certainly be a to that the of mathematical in on the it much to the of the and its of and in very of an Fisher p. mathematics was Fisher's subject and he himself at it in the hard school of the Cambridge a in a in he was to for a year and at the the theory of J. and statistical and theory It is hard to a complete biological and mathematical for the young man who was to revolutionize statistics and mathematical and statistical into evolutionary as had a theory of out of the of so Fisher, in the school of had a theory of evolution out of the survival of Fisher even to that he to Darwin the as did to Fisher the of we should to the G. when he a on the by Fisher's Cambridge of on also have today to to Fisher the great to Fisher the greatest of all I should also like to a word about Fisher the great figure some of not have He has many to that he was one of the who to some the of the Theory of which and which has to a role in biology. the on his contributions to It to present this two of theory as in great of 1932. what in a had the theory would have developed in much the way from Fisher's book The Genetical Theory of Natural in years at a mathematical in the devoted to the analytical of … to R. A. and I two to one it be the R. A. Fisher we were had R. A. Fisher the much R. A. Fisher the So in this a to his us for once the full of his In his second year, Fisher was in up the Cambridge University with the of of the University such as R. C. to be the first Professor of and Darwin of in his to the of in 1909, had cannot be too that a great of statistical work has to be the science of can large in this have been Fisher's for the work what cannot be as it has been by some historians of is that Fisher's for was the for his in statistical and evolutionary Bennett his was of influence in another for it him to as of the Darwin, the and man I (Fisher in by Bennett Darwin was to the young Fisher what J. S. Professor of at had been to the young Charles In Charles wrote on his from the I do long to see have been the friend to that man In his (Darwin Charles his “I have not a which my any other. was my with Professor When Charles his with a that Fisher could so well have of over his with and his as should do in the highest in over his by was a influence on Fisher in many and the he with Charles Darwin is of The of The Genetical Theory is in for the to the the last by many of the problems with in this to the publication of The Genetical Theory and the two were in much of it in we find Fisher, as early as to his great “I to up to a great work on the mathematics of In he “I are at work on evolution book … . How about new statistical I can on but do on with did with Fisher at Rothamsted the and The Genetical Theory to his in the The of It is difficult for as the century to the that Darwin's theory of evolution by natural selection years and for us to that itself was in some as to In The of in detail the years and the Bennett (1983) opens his with a of the in also and Fisher The old of Darwin's that could the that also to It had appeared as an to The Origin of Species when had that Darwin's theory was “the of this I do not Darwin, it is very a in the second edition of his in in which he can the of and and natural selection as a of the and present we can the of books a as a one of and the in an by a be in to the of the of to them in a p. this to Fisher out to that evolution by natural selection on Mendelian was a theory not only of the that had Darwin to his but also in that for evolutionary as By the of mathematics to Mendel's genetical Fisher was to an of the of natural selection. The I once Fisher selection is a for an of It was one of his first by in and in work it into the the of C. and it does not in The Genetical Theory, Fisher's of “the which has been that the of Natural on a of the in The is in the of an of a for it for its upon the of the word in its The from a by its in one be to upon a of the a of to the of the of his It is any very to the a of from the laws of and on the other the and of It is on the that the of Natural Fisher p. In Fisher contributed to as a edited by A. C. and E. B. Ford, an of the of the Theory of Natural (Fisher In the of Fisher with the main which have been the theory of natural selection, and the main difficulties which its have He out that all of have been in or by of or It is when Fisher wrote this in the When he it to in for as a he “I wrote it a long when the of my out a second edition of the The Genetical Theory was in my It the of the of in two Fisher on his … it was Darwin's not only to Biology but to the of natural to have to a by which a are in the of an it is which highly Fisher p. of the in that … the of an at one of his sex has a of about the that this were of his and to the probability that a of his in the should have left at one The such a as it would have appeared to his the of would for of the this has certainly Fisher p. from The Genetical Theory in the also provide a to Fisher's of the foundations of statistical for the of any statistical of is that it on the of the probability of an the has been cannot be but the development of Fisher's concept of is of of the to the and its see that the theory of evolution by natural selection has a the ratio of the probability of the natural as we on the of natural selection to its probability on the of pure is so The Genetical The the the Fisher's and all that The Genetical Theory is not an book. years its publication the custom is now for such a such a book were to be to a far on to and text the of and like the at a Fisher's book, by is all be from the very first one is not of this something (Crow Darwin did his best as he read in one I a good he wrote to Fisher in has is my to the of Ronald Fisher in this the of with such and writing is my to The to Charles Darwin's not and it is to of The Genetical Theory as a of to The Origin of Fisher was the Darwinian of his and it is not that it his of The Genetical Theory that is of its is its way with The Origin of Species, it be at on the that it was but an of the much work that Darwin had which I now be I cannot and for my and I to the some in my (Darwin 1859). the most of Fisher's is his (Fisher the on the of Mendelian It is that the is not with but what modern could such a when he to his in the from Fisher's of has of evolutionary biologists to assume that the most single in The Genetical Theory, on and the was to Fisher was not that the was in the first edition of The of (Darwin he was certainly of some for it in a by in The did not to Darwin any Fisher did to and one that the was at the the of in such of The Genetical Bennett (1983) has the of The Genetical Theory. in a position to Haldane and who wrote long it a In of (1930) the book was not as appreciated in the as it was in and this was to the Darwin Fisher that its influence would be to be … my is … that it be as a very to the I it be so really all that it … .” Fisher himself on from Oxford University Press in of was so long I from them that I had made up my mind that it was one of those books which and and would have influence on biological In the appreciated the of The Genetical Theory, but in his influential Genetics and the Origin of Species he the of by Wright and in was of Fisher's by to have to terms with its in and the Origin of Species or when writing on the history of biology. In The first edition he contributed a on the history of the evolutionary once The Genetical Theory a about “the evolutionary of … R. A. as a In his The of about Fisher's work that can be explained only by an study of The Genetical Theory, and his Charles Darwin and the of Modern nearly but of Fisher's at all of In the of E. B. and in that The Genetical Theory was not in the early wrote to Fisher in has been an to that the edition did not out long the but, a book is to be not by its but by its upon and book of the century has had a upon biology has this the ideas out from it a of of the but that of is what and I are to find like to be Fisher, he was Arthur Professor of Genetics at Cambridge from his in a very which had at the and the leading on evolutionary in has been a student of The Theory of edition with a of books for further which with The Origin of Species and books by and as well as R. C. C. and G. C. but not The Genetical Theory Haldane's The of The Genetical Theory has not been into any and outside the world only has it The main influential from the early of The Genetical Theory were and the which to be as the and “the theorem of natural The from been the for has a great of and and only now can we Fisher's my and for a the history of the theorem and that of see of The Genetical By the of Fisher's in The Genetical Theory had had only a Like Mendel's work, its now on the influence it should have the influence it did It lay for just as long as did Mendel's The publication of the second edition in made it and the of evolutionary biologists was to or in some its I the point by the of G. C. influential book and Natural with has been for the modern that natural selection is the of evolution. The Genetical Theory in the first of his but it is his of what him in the writing of his book to which I that I read that year were also of great to and have from some was and on sex ratio The other was in and and superb work went it was by who made the equation a concept for his treatment of sex … . from its “The of was a I had a who as as I that natural selection is a real scientific It that are of that and such as for the of the (Fisher that could not … . I have been by a of in that I had the and was far of my of the work of and should have made doubtful about any such I was really to it by the of other that some of what I was works on p. point in this of that the works of and and all to Fisher in The Genetical Theory. and from The Genetical Theory like Fisher, only the second edition of Darwin's The of had Fisher about natural selection and the sex ratio in and he had told to read The Genetical Theory, which I his into mathematics with the of and the of parental expenditure, which and had we now the was not to Fisher, but it was The Genetical Theory that made it influential “The evolution of appeared alongside Fisher's in as a In the of in his the that is by selection … .” I can do from has a long and R. A. Fisher, to the problem in his book in an obscure work. Fisher's on have been in the with evolution and from which many Fisher himself the to Darwin, a of Charles Darwin did the He his treatment of the subject was but he was his father had the problem in the first edition of his of and the in the second edition the was and the on later biologists the first Darwin's to have been even by his a Charles Darwin gave the for the crucial insight to … to works on had with a that Fisher had the of in The Genetical Theory, to the evolution of in The first in a Fisher gave to the Cambridge University in (Fisher It was later up by to the publication of Fisher's we now have a succinct of it a to A. G. does not which are the survival of for to a or lesser the of the So the parental are to natural selection, and in my book I do discuss how far we of the development of in … .” I this on The Genetical Theory with a from Fisher and Stock (1915) on the of a great book. I it with a from is a book as a I as of equal to the rest of my Cambridge the I on I it my … . a book that I only second in in evolution theory to Darwin's as with its and also as one of the greatest books of the century the of a edition is a major … . By the of my I have all that is in this book and I a I In some some of us have in this man is far in the of the Variorum Genetical In the years since Fisher told to read The Genetical Theory, I have it with I can I to all of of could have the scholarship of Henry Bennett and the of

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