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 biological inheritance following the principles originally proposed by Gregor Mendel in 1865 and 1866, re-discovered in 1900 by Hugo de Vries and Carl Correns, and later popularized ...
with
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English Natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
'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 Heredity, heritable traits characteristic of a population over generation ...
, 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, ...
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also known as orthogenetic evolution, progressive evolution, evolutionary progress, or progressionism, is an Superseded theories in science, obsolete biological hypothesis that organisms have an innate tendency to evolution, evolve ...
. First published in 1930 by The Clarendon Press, it is one of the most important books of the modern synthesis, and helped define
population genetics Population genetics is a subfield of genetics that deals with genetic differences within and among populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as Adaptation (biology), adaptation, s ...
. It had been described by J. F. Crow as the "deepest book on evolution since Darwin". 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 f ...
, Fisher's principle, reproductive value, Fisher's fundamental theorem of natural selection, Fisher's geometric model, the sexy son hypothesis,
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. In the simples ...
and the evolution of dominance. 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 experiment station, agricultural research institutions in the world, having been founde ...
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 the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
, 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 Augustinians, Augustinian ...
(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. Experiments on Plant Hybridis ...
) now allowed this. In the first chapter, Fisher considers the nature of inheritance, rejecting
blending inheritance Blending inheritance is an obsolete theory in biology from the 19th century. The theory is that the progeny inherits any characteristic as the average of the parents' values of that characteristic. As an example of this, a crossing of a red flo ...
, 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, ...
s affect biological fitness, Fisher's principle which explains why the
sex ratio A sex ratio is 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. However, many species deviate from an even sex ratio, ei ...
between
male Male (Planet symbols, symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or Egg cell, ovum, in the process of fertilisation. A male organism cannot sexual repro ...
s and
female An organism's sex is female ( symbol: ♀) if it produces the ovum (egg cell), the type of gamete (sex cell) that fuses with the male gamete (sperm cell) during sexual reproduction. A female has larger gametes than a male. Females and ...
s is almost always 1:1, reproductive value, examining the
demography Demography () is the statistical study of human populations: their size, composition (e.g., ethnic group, age), and how they change through the interplay of fertility (births), mortality (deaths), and migration. Demographic analysis examine ...
of having girl children. Using his knowledge of
statistics Statistics (from German language, German: ', "description of a State (polity), state, a country") is the discipline that concerns the collection, organization, analysis, interpretation, and presentation of data. In applying statistics to a s ...
, the Fisherian runaway, which explores how
sexual selection Sexual selection is a mechanism of evolution in which members of one 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 the opposite sex ...
can lead to a
positive feedback Positive feedback (exacerbating feedback, self-reinforcing feedback) is a process that occurs in a feedback loop where the outcome of a process reinforces the inciting process to build momentum. As such, these forces can exacerbate the effects ...
runaway loop, producing features such as the peacock's
plumage Plumage () 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, there can b ...
. He also wrote about the evolution of dominance, which explores genetic dominance.


Eugenics

The last five chapters (8-12) include Fisher's concern about dysgenics and proposals for
eugenics Eugenics is a set of largely discredited beliefs and practices that aim to improve the genetic quality of a human population. Historically, eugenicists have attempted to alter the frequency of various human phenotypes by inhibiting the fer ...
. Fisher attributed the fall of
civilization A civilization (also spelled civilisation in British English) is any complex society characterized by the development of state (polity), the state, social stratification, urban area, urbanization, and symbolic systems of communication beyon ...
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 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 is a public university, public research university based in Adelaide, South Australia. Established in 1874, it is the third-oldest university in Australia. Its main campus in the Adelaide city centre includes many Sa ...
.


Dedication

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."


Reviews

The book was reviewed by Charles Galton Darwin, 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 a process whereby natural selection favours a trait due to its positive effects on the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin selection can lead ...
. John Henry Bennett gave an account of the writing and reception of the book.
Sewall Wright Sewall Green Wright ForMemRS HonFRSE (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 alongside ...
, 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 described it as "brilliant." Reginald 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 statistics, statistician and evolutionary biology, evolutionary biologist Ronald Fisher. The proper way of applying the abst ...
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 The University of Cambridge is a Public university, public collegiate university, collegiate research university in Cambridge, England. Founded in 1209, the University of Cambridge is the List of oldest universities in continuous operation, wo ...
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, James F. Crow, and
A. W. F. Edwards Anthony William Fairbank Edwards, Fellow of the Royal Society, FRS One or more of the preceding sentences incorporates text from the royalsociety.org website where: (born 1935) is a British statistician, geneticist and evolutionary biologist. Ed ...
.


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) Ronald Fisher