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in 1942. Though the 'Modern Synthesis' is the basis of current evolutionary thinking, it refers to a historical event that took place in the 1930s and 1940s. Major figures in the development of the modern synthesis include R. A. Fisher, Theodosius Dobzhansky, J.B.S. Haldane, Sewall Wright, Julian Huxley, Ernst Mayr, Bernhard Rensch, George Gaylord Simpson, and G. Ledyard Stebbins. The modern synthesis solved difficulties and confusions caused by the specialisation and poor communication between biologists in the early years of the twentieth century. Discoveries of early geneticists were difficult to reconcile with gradual evolution and the mechanism of natural selection. The synthesis reconciled the two schools of thought, while providing evidence that studies of populations in the field were crucial to evolutionary theory. It drew together ideas from several branches of biology that had become separated, particularly genetics, cytology, systematics, botany, morphology, ecology and paleontology. Modern evolutionary synthesis is also referred to as the new synthesis, the '''modern synthesis''', and the '''evolutionary synthesis'''. The ideas of the modern synthesis have sometimes been called '''neo-Darwinism'''. DEVELOPMENTS LEADING UP TO THE SYNTHESIS See Also: history of evolutionary thought 1859-1899 ''The Origin of Species'' was successful in convincing most of the scientific community of the fact that evolution had occurred, but was much less successful in convincing naturalists that natural selection was its primary mechanism. In the 19th and early 20th centuries variations of Lamarckism, orthogenesis ("progressive" evolution), and saltationism (evolution by "jumps" or mutations) were discussed as alternatives. Bowler Evolution:The History of an Idea pp.236-256 Also, Darwin did not offer a precise explanation of how new species arise. As part of the disagreement about whether natural selection alone was sufficient to explain speciation, George John Romanes coined the term "neo-Darwinism" to refer to the version of evolution advocated by Alfred Russel Wallace and August Weismann with its heavy dependence on natural selection. Gould The Structure of Evolutionary theory p. 216 Weismann and Wallace rejected the Lamarckian idea of inheritance of acquired characteristics, something that Darwin had not ruled out. Larson p. 86 Weismann's idea was that the relationship between the hereditary material, which he called the . 1900-1915 Gregor Mendel 's work was re-discovered by Hugo De Vries and Carl Correns . It showed that the contributions of each parent retained its integrity rather than blending with the contribution of the other parent. However, the early Mendelians viewed hard inheritance as incompatible with natural selection and favored saltationism (large mutations or jumps) instead. Larson pp. 157-166 The Biometric School , led by Karl Pearson and Walter Weldon , argued vigorously against it, saying that empirical evidence indicated that variation was continuous in most organisms not discrete as Mendelalism predicted. The relevance of Mendelism to evolution was unclear and hotly debated. T. H. 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 Drosophila Melanogaster , which helped establish the link between Mendelian genetics and the chromosomal theory of inheritance, demonstrated that rather than creating new species in a single step, mutations increased the genetic variation in the population. Bowler pp. 271-272 The foundation of population genetics The first step towards the synthesis was the development of . Haldane established that natural selection could work in the real world at a faster rate than even Fisher had assumed.Bowler Evolution:The history of an Idea pp. 325-339 Sewall Wright focused on combinations of genes that interacted as complexes, and the effects of inbreeding on small relatively isolated populations, which could exhibit Genetic Drift . In a 1932 paper he introduced the concept of an Adaptive Landscape in which phenomena such as cross breeding and genetic drift in small populations could push them away from adaptive peaks, which would in turn allow Natural Selection to push them towards new adaptive peaks. Wright's model would appeal to field naturalists such as Theodosius Dobzhansky and Ernst Mayr who were becoming aware of the importance of geographical isolation in real world populations. The work of Fisher, Haldane and Wright founded the discipline of Population Genetics . This is the precursor of the modern synthesis, which is an even broader coalition of ideas. Gould The Structure of Evolutionary Theory pp. 503-518 THE MODERN SYNTHESIS Theodosius Dobzhansky, a Russian emigre who had been a postdoctoral worker in T.H. Morgan's fruit fly lab, was one of the first to apply genetics to natural populations. He worked mostly with '''' was a key step in bridging the gap between population geneticists and field naturalists. It presented the conclusions reached by Fisher, Haldane, and especially Wright in their highly mathematical papers in a form that was easily accessible to others. It also emphasized that real world populations had far more genetic variability than the early population geneticists had assumed in their models, and that genetically distinct sub-populations were important. Dobzhansky argued that natural selection worked to maintain genetic diversity as well as driving change. Dobzhansky had been influenced by his exposure in the 1920s to the work of a Russian geneticist named Sergei Chetverikov who had looked at the role of recessive genes in maintaining a reservoir of genetic variability in a population before his work was shutdown by the rise of Lysenkoism in the Soviet Union . Ernst Mayr 's key contribution to the synthesis was '' Systematics And The Origin Of Species '', published in 1942. Mayr emphasized the importance of Allopatric Speciation , where geographically isolated sub-populations diverge so far that Reproductive Isolation occurs. He was sceptical of the reality of Sympatric Speciation believing that geographical isolation was a prerequisite for building up intrinsic (reproductive) isolating mechanisms. George Simpson was responsible for showing that the modern synthesis was compatible with paleontology in his book '' Tempo And Mode In Evolution '' published in 1944. Simpson's work was crucial because so many paleontologists had disagreed, in some cases vigorously, with the idea that natural selection was the main mechanism of evolution. It showed that the trends of linear progression (in for example the Evolution Of The Horse ) that earlier paleontologists had used as support for Neo-Lamarckism and Orthogenesis did not hold up under careful examination. Instead the fossil record was consistent with the irregular, branching, and non-directional pattern predicted by the modern synthesis. The botanist G. Ledyard Stebbins was another major contributor to the synthesis. His major work, '' Variation And Evolution In Plants '', was published in 1950. It extended the synthesis to encompass botany including the important effects of hybridization and Polyploidy in some kinds of plants. . TENETS OF THE MODERN SYNTHESIS According to the modern synthesis as established in the 1930s and 1940s , genetic variation in populations arises by chance through Mutation (this is now known to be sometimes caused by mistakes in DNA Replication ) and Recombination (crossing over of homologous Chromosome s during Meiosis ). Evolution consists primarily of changes in the Frequencies Of Alleles between one generation and another as a result of Genetic Drift , Gene Flow , and Natural Selection . Speciation occurs gradually when populations are reproductively isolated, for example by geographic barriers. FURTHER ADVANCES The modern evolutionary synthesis continued to be developed and refined after the initial establishment in the 1930s and 1940s. The work of W. D. Hamilton , George C. Williams , John Maynard Smith and others led to the development of a Gene-centric View Of Evolution in the 1960s. The synthesis as it exists now has extended the scope of the Darwinian idea of natural selection to include subsequent scientific discoveries and concepts unknown to Darwin, such as DNA and Genetics , which allow rigorous, in many cases mathematical, analyses of phenomena such as Kin Selection , Altruism , and Speciation . A particular interpretation of neo-Darwinism most commonly associated with Richard Dawkins , author of '' The Selfish Gene '', asserts that the gene is the only true Unit Of Selection . Bowler p.361 Dawkins further extended the Darwinian idea to include non-biological systems exhibiting the same type of selective behavior of the 'fittest' such as Meme s in culture. SEE ALSO REFERENCES IMPORTANT PUBLICATIONS
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