Modern synthesis (20th century) a gene-centred view of evolution following Hamilton's concepts, disputing the idea of evolutionary progress, and attacking the then widespread theory of group selection. Williams argued that natural selection worked by changing the frequency of alleles, and could not work at the level of groups. Gene-centred evolution was popularised by Richard Dawkins in his 1976 book "The Selfish Gene" and developed in his more technical writings. In 1975, E. O. Wilson published his controversial book "", the subtitle alluding to the modern synthesis as he attempted to bring the study of animal society into the evolutionary fold. This appeared radically new,
Alternatives to evolution by natural selection through a population when they were beneficial, helped by natural selection. This represented the germ of the modern synthesis, and the beginning of the end for mutationism as an evolutionary force. Contemporary biologists accept that mutation and selection both play roles in evolution; the mainstream view is that while mutation supplies material for selection in the form of variation, all non-random outcomes are caused by natural selection. Masatoshi Nei argues instead that the production of more efficient genotypes by mutation is fundamental for evolution, and that evolution is often mutation-limited. The endosymbiotic theory implies rare but major events of saltational