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,
The Selfish Gene to behave selflessly with each other. A lineage is expected to evolve to maximise its inclusive fitness—the number of copies of its genes passed on globally (rather than by a particular individual). As a result, populations will tend towards an evolutionarily stable strategy. The book also coins the term "meme" for a unit of human cultural evolution analogous to the gene, suggesting that such "selfish" replication may also model human culture, in a different sense. Memetics has become the subject of many studies since the publication of the book. In raising awareness of Hamilton's ideas, as well as making its