The modern synthesis was the early 20th-century synthesis of Charles Darwin's theory of evolution and Gregor Mendel's ideas on heredity into a joint mathematical framework. Julian Huxley coined the term in his 1942 book, Evolution: The Modern Synthesis. The synthesis combined the ideas of natural selection, Mendelian genetics, and population genetics. It also related the broad-scale macroevolution seen by palaeontologists to the small-scale microevolution of local populations.
The synthesis was defined differently by its founders, with Ernst Mayr in 1959, G. Ledyard Stebbins in 1966, and Theodosius Dobzhansky in 1974 offering differing basic postulates, though they all include natural selection, working on heritable variation supplied by mutation. Other major figures in the synthesis included E. B. Ford, Bernhard Rensch, Ivan Schmalhausen, and George Gaylord Simpson. An early event in the modern synthesis was R. A. Fisher's 1918 paper on mathematical population genetics, though William Bateson, and separately Udny Yule, had already started to show how Mendelian genetics could work in evolution in 1902.
Different syntheses followed, 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 and Gerd B. Müller's proposed extended evolutionary synthesis of 2007. In the view of 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.
Developments leading up to the synthesis
Darwin's evolution by natural selection, 1859
Charles Darwin's 1859 book, On the Origin of Species, convinced most biologists that evolution had occurred, but not that natural selection was its primary mechanism. In the 19th and early 20th centuries, variations of Lamarckism (inheritance of acquired characteristics), orthogenesis (progressive evolution), saltationism (evolution by jumps) and mutationism (evolution driven by mutations) were discussed as alternatives. Darwin himself had sympathy for Lamarckism, but Alfred Russel Wallace advocated natural selection and totally rejected Lamarckism. In 1880, Samuel Butler labelled Wallace's view neo-Darwinism.
The eclipse of Darwinism, 1880s onwards
From the 1880s onwards, biologists grew skeptical of Darwinian evolution. This eclipse of Darwinism (in Julian Huxley's words) grew out of the weaknesses in Darwin's account, with respect to his view of inheritance. Darwin believed in blending inheritance, which implied that any new variation, even if beneficial, would be weakened by 50% at each generation, as the engineer Fleeming Jenkin noted in 1868. This in turn meant that small variations would not survive long enough to be selected. Blending would therefore directly oppose natural selection. In addition, Darwin and others considered Lamarckian inheritance of acquired characteristics entirely possible, and Darwin's 1868 theory of pangenesis, with contributions to the next generation (gemmules) flowing from all parts of the body, actually implied Lamarckism as well as blending.
Weismann's germ plasm, 1892
August Weismann's idea, set out in his 1892 book Das Keimplasma: eine Theorie der Vererbung ("The Germ Plasm: a Theory of Inheritance"), was that the hereditary material, which he called the germ plasm, and the rest of the body (the soma) had a one-way relationship: the germ-plasm formed the body, but the body did not influence the germ-plasm, except indirectly in its participation in a population subject to natural selection. If correct, this made Darwin's pangenesis wrong, and Lamarckian inheritance impossible. His experiment on mice, cutting off their tails and showing that their offspring had normal tails, demonstrated that inheritance was 'hard'. He argued strongly and dogmatically for Darwinism and against Lamarckism, polarising opinions among other scientists. This increased anti-Darwinian feeling, contributing to its eclipse.
Disputed beginnings
Genetics, mutationism and biometrics, 1900–1918
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. 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. This reinforced a major division of thought, already present in the 1890s, between gradualists who followed Darwin, and saltationists such as Bateson.: Mutations (as 'sports') and polymorphisms were well known long before the Mendelian recovery.
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; and the biometric school, led by Karl Pearson and Walter Weldon. The biometricians 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.
A traditional view is that the biometricians and the Mendelians rejected natural selection and argued for their separate theories for 20 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. 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 heterozygous combinations might, on seriation, give so near an approach to a continuous curve, that the purity of the elements would be unsuspected". Also in 1902, the statistician Udny Yule showed mathematically that given multiple factors, Mendel's theory enabled continuous variation. Yule criticised Bateson's approach as confrontational, but failed to prevent the Mendelians and the biometricians from falling out.
Castle's hooded rats, 1911
Starting in 1906, William Castle carried out a long study of the effect of selection on coat colour in rats. The piebald or hooded pattern was recessive to the grey wild type. He crossed hooded rats with both wild and "Irish" types, and then back-crossed the offspring with pure hooded rats. The dark stripe on the back was bigger. He then tried selecting different groups for bigger or smaller stripes for 5 generations and found that it was possible to change the characteristics considerably beyond the initial range of variation. This effectively refuted de Vries's claim that continuous variation was caused by the environment and could not be inherited. By 1911, Castle noted that the results could be explained by Darwinian selection on a heritable variation of a sufficient number of Mendelian genes.
Morgan's fruit flies, 1912
Thomas Hunt Morgan began his career in genetics as a saltationist and started out trying to demonstrate that mutations could produce new species in fruit flies. However, the experimental work at his lab with the fruit fly, Drosophila melanogaster showed that rather than creating new species in a single step, mutations increased the supply of genetic variation in the population. By 1912, after years of work on the genetics of fruit flies, Morgan showed that these insects had many small Mendelian factors (discovered as mutant flies) on which Darwinian evolution could work as if the variation was fully continuous. The way was open for geneticists to conclude that Mendelism supported Darwinism.
An obstruction: Woodger's positivism, 1929
The theoretical biologist and philosopher of biology Joseph Henry Woodger led the introduction of positivism into biology with his 1929 book Biological Principles. He saw a mature science as being characterised by a framework of hypotheses that could be verified by facts established by experiments. He criticised the traditional natural history style of biology, including the study of evolution, as immature science, since it relied on narrative. Woodger set out to play the role of Robert Boyle's 1661 Sceptical Chymist, intending to convert the subject of biology into a formal, unified science, and ultimately, following the Vienna Circle of logical positivists like Otto Neurath and Rudolf Carnap, to reduce biology to physics and chemistry. His efforts stimulated the biologist J. B. S. Haldane to push for the axiomatisation of biology, and by influencing thinkers such as Huxley, helped to bring about the modern synthesis. The positivist climate made natural history unfashionable, and in America, research and university-level teaching on evolution declined almost to nothing by the late 1930s. The Harvard physiologist William John Crozier told his students that evolution was not even a science: "You can't experiment with two million years!" The tide of opinion turned with the adoption of mathematical modelling and controlled experimentation in population genetics, combining genetics, ecology and evolution in a framework acceptable to positivism.
Elements of the synthesis Fisher and Haldane's mathematical population genetics, 1918–1930
In 1918, R. A. Fisher wrote "The Correlation between Relatives on the Supposition of Mendelian Inheritance," "Paper read by J. Arthur Thomson on July 8, 1918 to the Royal Society of Edinburgh." which showed how continuous variation could come from a number of discrete genetic loci. In this and other papers, culminating in his 1930 book The Genetical Theory of Natural Selection, Fisher showed how Mendelian genetics was consistent with the idea of evolution by natural selection.
In the 1920s, a series of papers by J. B. S. Haldane analyzed real-world examples of natural selection, such as the evolution of industrial melanism in peppered moths. and showed that natural selection could work even faster than Fisher had assumed. Both of these scholars, and others, such as Dobzhansky and Wright, wanted to raise biology to the standards of the physical sciences by basing it on mathematical modeling and empirical testing. Natural selection, once considered unverifiable, was becoming predictable, measurable, and testable.
De Beer's embryology, 1930