The Genetical Theory of Natural Selection
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''The Genetical Theory of Natural Selection'' is a book by
Ronald Fisher Sir Ronald Aylmer Fisher (17 February 1890 – 29 July 1962) was a British polymath who was active as a mathematician, statistician, biologist, geneticist, and academic. For his work in statistics, he has been described as "a genius who a ...
which combines
Mendelian genetics Mendelian inheritance (also known as Mendelism) is a type of biology, biological Heredity, inheritance following the principles originally proposed by Gregor Mendel in 1865 and 1866, re-discovered in 1900 by Hugo de Vries and Carl Correns, an ...
with
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended fr ...
's theory of
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Charle ...
, with Fisher being the first to argue that "Mendelism therefore validates Darwinism" and stating with regard to
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, mi ...
s that "The vast majority of large mutations are deleterious; small mutations are both far more frequent and more likely to be useful", thus refuting
orthogenesis Orthogenesis, also known as orthogenetic evolution, progressive evolution, evolutionary progress, or progressionism, is an obsolete biological hypothesis that organisms have an innate tendency to evolve in a definite direction towards some go ...
. First published in 1930 by
The Clarendon Press Oxford University Press (OUP) is the university press of the University of Oxford. It is the largest university press in the world, and its printing history dates back to the 1480s. Having been officially granted the legal right to print books ...
, it is one of the most important books of the
modern synthesis Modern synthesis or modern evolutionary synthesis refers to several perspectives on evolutionary biology, namely: * Modern synthesis (20th century), the term coined by Julian Huxley in 1942 to denote the synthesis between Mendelian genetics and s ...
, and helped define
population genetics Population genetics is a subfield of genetics that deals with genetic differences within and between populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as adaptation, speciation, and pop ...
. It is commonly cited in biology books, outlining many concepts that are still considered important such as
Fisherian runaway Fisherian runaway or runaway selection is a sexual selection mechanism proposed by the mathematical biologist Ronald Fisher in the early 20th century, to account for the evolution of ostentatious male ornamentation by persistent, directional fem ...
,
Fisher's principle Fisher's principle is an evolutionary model that explains why the sex ratio of most species that produce offspring through sexual reproduction is approximately 1:1 between males and females. A. W. F. Edwards has remarked that it is "probably the mo ...
, reproductive value,
Fisher's fundamental theorem of natural selection Fisher's fundamental theorem of natural selection is an idea about genetic variance in population genetics developed by the statistician and evolutionary biologist Ronald Fisher. The proper way of applying the abstract mathematics of the theorem ...
, Fisher's geometric model, the
sexy son hypothesis The sexy son hypothesis in evolutionary biology and sexual selection, proposed by Patrick J. Weatherhead and Raleigh J. Robertson of Queen's University in Kingston, Ontario in 1979, states that a female's ideal mate choice among potential mate ...
,
mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. Often, mimicry f ...
and the
evolution of dominance In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and ...
. It was dictated to his wife in the evenings as he worked at
Rothamsted Research Rothamsted Research, previously known as the Rothamsted Experimental Station and then the Institute of Arable Crops Research, is one of the oldest agricultural research institutions in the world, having been founded in 1843. It is located at Harp ...
in the day.


Contents

In the preface, Fisher considers some general points, including that there must be an understanding of natural selection distinct from that of
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
, and that the then-recent advances in the field of
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar wor ...
(see
history of genetics The history of genetics dates from the classical era with contributions by Pythagoras, Hippocrates, Aristotle, Epicurus, and others. Modern genetics began with the work of the Augustinian friar Gregor Johann Mendel. His work on pea plants, publis ...
) now allowed this. In the first chapter, Fisher considers the nature of inheritance, rejecting
blending inheritance Blending may refer to: * The process of mixing in process engineering * Mixing paints to achieve a greater range of colors * Blending (alcohol production), a technique to produce alcoholic beverages by mixing different brews * Blending (linguist ...
, 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 Parental investment, in evolutionary biology and evolutionary psychology, is any parental expenditure (e.g. time, energy, resources) that benefits offspring.Clutton-Brock, T.H. 1991. ''The Evolution of Parental Care''. Princeton, NJ: Princeton ...
, Fisher's geometric model, concerning how spontaneous
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, mi ...
s affect
biological fitness Fitness (often denoted w or ω in population genetics models) is the quantitative representation of individual reproductive success. It is also equal to the average contribution to the gene pool of the next generation, made by the same individua ...
, Fisher's principle which explains why the
sex ratio The sex ratio (or gender ratio) is usually defined as the ratio of males to females in a population. As explained by Fisher's principle, for evolutionary reasons this is typically about 1:1 in species which reproduce sexually. Many species devia ...
between
male Male (symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or ovum, in the process of fertilization. A male organism cannot reproduce sexually without access to ...
s and
female Female (Venus symbol, symbol: ♀) is the sex of an organism that produces the large non-motile ovum, ova (egg cells), the type of gamete (sex cell) that fuses with the Sperm, male gamete during sexual reproduction. A female has larger gamet ...
s is almost always 1:1, reproductive value, examining the
demography Demography () is the statistics, statistical study of populations, especially human beings. Demographic analysis examines and measures the dimensions and Population dynamics, dynamics of populations; it can cover whole societies or groups ...
of having girl children. Using his knowledge of
statistics Statistics (from German language, German: ''wikt:Statistik#German, Statistik'', "description of a State (polity), state, a country") is the discipline that concerns the collection, organization, analysis, interpretation, and presentation of ...
, the Fisherian runaway, which explores how
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of t ...
can lead to a
positive feedback Positive feedback (exacerbating feedback, self-reinforcing feedback) is a process that occurs in a feedback loop which exacerbates the effects of a small disturbance. That is, the effects of a perturbation on a system include an increase in the ...
runaway loop, producing features such as the peacock's
plumage Plumage ( "feather") is a layer of feathers that covers a bird and the pattern, colour, and arrangement of those feathers. The pattern and colours of plumage differ between species and subspecies and may vary with age classes. Within species, ...
. He also wrote about the evolution of dominance, which explores
genetic dominance In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and t ...
.


Eugenics

The last five chapters (8-12) include Fisher's concern about
dysgenics Dysgenics (also known as cacogenics) is the decrease in prevalence of traits deemed to be either socially desirable or well adapted to their environment due to selective pressure disfavoring the reproduction of those traits. The adjective "dysgeni ...
and proposals for
eugenics Eugenics ( ; ) is a fringe set of beliefs and practices that aim to improve the genetic quality of a human population. Historically, eugenicists have attempted to alter human gene pools by excluding people and groups judged to be inferior or ...
. Fisher attributed the fall of
civilization A civilization (or civilisation) is any complex society characterized by the development of a state, social stratification, urbanization, and symbolic systems of communication beyond natural spoken language (namely, a writing system). Ci ...
s 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.


Editions

A second, slightly revised edition was republished in 1958. In 1999, a third
variorum A variorum, short for ''(editio) cum notis variorum'', is a work that collates all known variants of a text. It is a work of textual criticism, whereby all variations and emendations are set side by side so that a reader can track how textual deci ...
edition (), 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 The University of Adelaide (informally Adelaide University) is a public research university located in Adelaide, South Australia. Established in 1874, it is the third-oldest university in Australia. The university's main campus is located on N ...
.


Dedication

The book is dedicated to Major
Leonard Darwin Leonard Darwin (15 January 1850 – 26 March 1943) was an English politician, economist and eugenicist. He was a son of the naturalist Charles Darwin, and also a mentor to Ronald Fisher, a statistician and evolutionary biologist. Biography L ...
, 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."


Reviews

The book was reviewed by
Charles Galton Darwin Sir Charles Galton Darwin (19 December 1887 – 31 December 1962) was an English physicist who served as director of the National Physical Laboratory (NPL) during the Second World War. He was a son of the mathematician George Howard Darwin an ...
, who sent Fisher his copy of the book, with notes in the margin, starting a correspondence which lasted several years. The book also had a major influence on
W. D. Hamilton William Donald Hamilton (1 August 1936 – 7 March 2000) was a British evolutionary biologist, recognised as one of the most significant evolutionary theorists of the 20th century. Hamilton became known for his theoretical work expounding a ...
's theories on the genetic basis of
kin selection Kin selection is the evolutionary strategy that favours the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin altruism can look like altruistic behaviour whose evolution i ...
. John Henry Bennett gave an account of the writing and reception of the book.
Sewall Wright Sewall Green Wright FRS(For) Honorary FRSE (December 21, 1889March 3, 1988) was an American geneticist known for his influential work on evolutionary theory and also for his work on path analysis. He was a founder of population genetics alongsi ...
, 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."
J. B. S. Haldane John Burdon Sanderson Haldane (; 5 November 18921 December 1964), nicknamed "Jack" or "JBS", was a British-Indian scientist who worked in physiology, genetics, evolutionary biology, and mathematics. With innovative use of statistics in biolog ...
described it as "brilliant."
Reginald Punnett Reginald Crundall Punnett FRS (; 20 June 1875 – 3 January 1967) was a British geneticist who co-founded, with William Bateson, the ''Journal of Genetics'' in 1910. Punnett is probably best remembered today as the creator of the Punnett ...
was negative, however. The book was largely overlooked for 40 years, and in particular Fisher's
fundamental theorem of natural selection Fisher's fundamental theorem of natural selection is an idea about genetic variance in population genetics developed by the statistician and evolutionary biologist Ronald Fisher. The proper way of applying the abstract mathematics of the theorem t ...
was misunderstood. The work had a great effect on
W. D. Hamilton William Donald Hamilton (1 August 1936 – 7 March 2000) was a British evolutionary biologist, recognised as one of the most significant evolutionary theorists of the 20th century. Hamilton became known for his theoretical work expounding a ...
, who discovered it as an undergraduate at the
University of Cambridge , mottoeng = Literal: From here, light and sacred draughts. Non literal: From this place, we gain enlightenment and precious knowledge. , established = , other_name = The Chancellor, Masters and Schola ...
and noted in these excerpts from the rear cover of the 1999 variorum edition: The publication of the variorum edition in 1999 led to renewed interest in the work and reviews by Laurence Cook,
Brian Charlesworth Brian Charlesworth (born 29 April 1945) is a British evolutionary biologist at the University of Edinburgh, and editor of ''Biology Letters''. Since 1997, he has been Royal Society Research Professor at the Institute of Evolutionary Biology (IE ...
, James F. Crow, and
A. W. F. Edwards Anthony William Fairbank Edwards, FRS (born 1935) is a British statistician, geneticist and evolutionary biologist. He is the son of the surgeon Harold C. Edwards, and brother of medical geneticist John H. Edwards. He has sometimes been called ...
.


References


Bibliography

* * * * "This book is based on a series of lectures delivered in January 1931 at the Prifysgol Cymru, Aberystwyth, and entitled 'A re-examination of Darwinism'."


External links

*
The Genetical Theory Of Natural Selection
{{DEFAULTSORT:Genetical Theory of Natural Selection, The 1930 non-fiction books Books about evolution Eugenics books Genetics in the United Kingdom History of genetics 1930 in biology Modern synthesis (20th century)