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Different syntheses followed, accompanying the gradual breakup of the early 20th century synthesis, including with social behaviour
in E. O. Wilson's sociobiology in 1975, evolutionary developmental biology's integration of embryology with genetics and evolution,
starting in 1977, and Massimo Pigliucci's proposed extended evolutionary synthesisof 2007. In the view of the evolutionary biologist
Eugene Koonin in 2009, the modern synthesis will be replaced by a 'post-modern' synthesis that will include revolutionary changes in
molecular biology, the study of prokaryotes and the resulting tree of life, and genomics.[2]
Contents
Developments leading up to the synthesis
Darwin's evolution by natural selection, 1859
The eclipse of Darwinism, 1880s onwards
Weismann's germ plasm, 1892
Disputed beginnings
Genetics, mutationism and biometrics, 1900–1918
Castle's hooded rats, 1911
Morgan's fruit flies, 1912
An obstruction: Woodger's positivism, 1929
Events in the synthesis
Fisher and Haldane's mathematical population genetics, 1918–1930
De Beer's embryology, 1930
Wright's adaptive landscape, 1932
Dobzhansky's evolutionary genetics, 1937
Ford's ecological genetics, 1940
Schmalhausen's stabilizing selection, 1941
Huxley's popularising synthesis, 1942
Mayr's allopatric speciation, 1942
Simpson's palaeontology, 1944
The Society for the Study of Evolution, 1946
Stebbins's botany, 1950
Definitions by the founders
After the synthesis
Hamilton's inclusive fitness, 1964
Williams's gene-centred evolution, 1966
Wilson's sociobiology, 1975
Lewis's homeotic genes, 1978
Later syntheses
Pigliucci's extended evolutionary synthesis, 2007
Koonin's 'post-modern' evolutionary synthesis, 2009
Towards a replacement synthesis
Historiography
See also
Notes
References
Sources
Further reading
While carrying out breeding experiments to clarify the mechanism of inheritance in 1900,
Hugo de Vries and Carl Correns independently rediscovered Gregor Mendel's work. News of
this reached William Bateson in England, who reported on the paper during a presentation to
the Royal Horticultural Society in May 1900.[17] In Mendelian inheritance, the contributions
of each parent retain their integrity rather than blending with the contribution of the other
parent. In the case of a cross between two true-breeding varieties such as Mendel's round and
wrinkled peas, the first-generation offspring are all alike, in this case all round. Allowing these
to cross, the original characteristics reappear (segregation): about 3/4 of their offspring are
round, 1/4 wrinkled. There is a discontinuity between the appearance of the offspring; de
Vries coined the term allele for a variant form of an inherited characteristic.[18] This
reinforced a major division of thought, already present in the 1890s, between gradualists who
[19]
followed Darwin, and saltationists such as Bateson.
William Bateson
championed Mendelism. The two schools were the Mendelians, such as Bateson and de Vries, who favoured
mutationism, evolution driven by mutation, based on genes whose alleles segregated
discretely like Mendel's peas;[20][21] and the biometric school, led by Karl Pearson and Walter
Weldon. They argued vigorously against mutationism, saying that empirical evidence indicated that variation was continuous in most
organisms, not discrete as Mendelism seemed to predict; they wrongly believed that Mendelism inevitably implied evolution in
discontinuous jumps.[22][23]
A traditional view is that the biometricians and the Mendelians rejected natural selection and argued for their separate theories for 20
[22][24]
years, the debate only resolved by the development of population genetics.
A more recent view is that Bateson, de Vries, Thomas Hunt Morgan and Reginald Punnett had
by 1918 formed a synthesis of Mendelism and mutationism. The understanding achieved by
these geneticists spanned the action of natural selection on alleles (alternative forms of a
gene), the Hardy-Weinberg equilibrium, the evolution of continuously-varying traits (like
height), and the probability that a new mutation will become fixed. In this view, the early
geneticists accepted natural selection but rejected Darwin's non-Mendelian ideas about
variation and heredity, and the synthesis began soon after 1900.[25][26] The traditional claim
that Mendelians rejected the idea of continuous variation is false; as early as 1902, Bateson
and Saunders wrote that "If there were even so few as, say, four or five pairs of possible
allelomorphs, the various homo- and hetero-zygous combinations might, on seriation, give so
near an approach to a continuous curve, that the purity of the elements would be
unsuspected".[27] Also in 1902, the statistician Udny Yule showed mathematically that given Karl Pearson led the
multiple factors, Mendel's theory enabled continuous variation. Yule criticised Bateson's biometric school.
confrontational approach,[28] but failed to prevent the Mendelians and the biometricians from
falling out.[29]
During the 1920s, a series of papers by J. B. S. Haldane applied mathematical analysis to real-
world examples of natural selection, such as the evolution of industrial melanism in peppered
moths.[38] Haldane established that natural selection could work even faster than Fisher had
assumed.[40] Both workers, and others such as Dobzhansky and Wright, explicitly intended to
bring biology up to the philosophical standard of the physical sciences, making it firmly based
Ronald Fisher brought
in mathematical modelling, its predictions confirmed by experiment. Natural selection, once
mathematics into population
considered hopelessly unverifiable speculation about history, was becoming predictable,
genetics.
measurable, and testable.[41]
However, the book was not what it seemed. In the view of the philosopher of science Michael Ruse, and in Huxley's own opinion,
Huxley was "a generalist, a synthesizer of ideas, rather than a specialist".[67] Ruse observes that Huxley wrote as if he were just
adding empirical evidence to the mathematical framework established by Fisher and the population geneticists, but that this was not
so. Huxley avoided mathematics, for instance not even mentioningFisher's fundamental theorem of natural selection.Instead, Huxley
used a mass of examples to demonstrate that natural selection is powerful, and that it works on Mendelian genes. The book was
successful in its goal of persuading readers of the reality of evolution, effectively illustrating island biogeography, speciation,
competition and so on. Huxley further showed that the appearance oforthogenetic trends - predictable directions for evolution - in the
fossil record were readily explained as allometric growth (since parts are interconnected). All the same, Huxley did not reject
orthogenesis out of hand, but maintained a belief in progress all his life, with Homo sapiens as the end point, and he had since 1912
been influenced by the vitalist philosopher Henri Bergson, though in public he maintained an atheistic position on evolution.[67]
Huxley's belief in progress within evolution and evolutionary humanism was shared in various forms by Dobzhansky, Mayr, Simpson
and Stebbins, all of them writing about "the future of Mankind". Both Huxley and Dobzhansky admired the palaeontologist priest
Pierre Teilhard de Chardin, Huxley writing the introduction to Teilhard's 1955 book on orthogenesis, The Phenomenon of Man. This
.[70]
vision required evolution to be seen as the central and guiding principle of biology
Later syntheses
In 1982, a historical note on a series of evolutionary biology books[e] could state
without qualification that evolution is the central organizing principle of biology.
Smocovitis commented on this that "What the architects of the synthesis had worked
to construct had by 1982 become a matter of fact", adding in a footnote that "the
centrality of evolution had thus been rendered tacit knowledge, part of the received
wisdom of the profession".[93]
By the late 20th century, however, the modern synthesis was showing its age, and
fresh syntheses to remedy its defects and fill in its gaps were proposed from dif
ferent
directions. These have included such diverse fields as the study of society,[85]
developmental biology,[47] epigenetics,[94] molecular biology, microbiology,
genomics,[2] symbiogenesis, and horizontal gene transfer.[95] The physiologist Denis
Noble argues that these additions render neo-Darwinism in the sense of the early
20th century's modern synthesis "at the least, incomplete as a theory of Evolutionary developmental biology
evolution",[95] and one that has been falsified by later biological research.
[95] has formed a synthesis of
evolutionary and developmental
Michael Rose and Todd Oakley note that evolutionary biology, formerly divided and biology, discovering deep homology
between the embryogenesis of such
"Balkanized", has been brought together by genomics. It has in their view discarded
different animals as insects and
at least five common assumptions from the modern synthesis, namely that the
vertebrates.
genome is always a well-organised set of genes; that each gene has a single function;
that species are well adapted biochemically to their ecological niches; that species
are the durable units of evolution, and all levels from organism to organ, cell and molecule within the species are characteristic of it;
and that the design of every organism and cell is efficient. They argue that the "new biology" integrates genomics, bioinformatics,
and evolutionary genetics into a general-purpose toolkit for a Postmodern
" Synthesis".[96]
Historiography
Looking back at the conflicting accounts of the modern synthesis, the historian Betty Smocovitis notes in her 1996 book Unifying
Biology: The Evolutionary Synthesis and Evolutionary Biology that both historians and philosophers of biology have attempted to
grasp its scientific meaning, but have found it "a moving target";[105] the only thing they agreed on was that it was a historical
event.[105] In her words "by the late 1980s the notoriety of the evolutionary synthesis was recognized . . . So notorious did 'the
synthesis' become, that few serious historically minded analysts would touch the subject, let alone know where to begin to sort
[106]
through the interpretive mess left behind by the numerous critics and commentators".
See also
Developmental systems theory
Gene-centered view of evolution
History of evolutionary thought
Neo-Darwinism
Objections to evolution
Notes
a. Also known variously as thenew synthesis, the modern evolutionary synthesis, the evolutionary synthesis,
and the neo-Darwinian synthesis. These alternative terms are ambiguous as they could possibly include later
syntheses, so this article usesJulian Huxley's 1942 "modern synthesis" throughout.
b. Peter Gauthier has however argued thatWeismann's experiment showed only that injury did not affect the germ
plasm. It did not test the effect of Lamarckian use and disuse.[13]
c. Fisher also analysed sexual selection in his book, but his work was largely ignored, and Darwin's case for such
selection misunderstood, so it formed no substantial part of the modern synthesis.[39]
d. Though C. H. Waddington had called for embryology to be added to the synthesis in his 1953 paper "Epigenetics
and Evolution".[49]
e. In a reissue of Dobzhansky'sGenetics and the Origin of Species.
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