The Evolution of Cooperation insects, where the workers – which are sterile, and therefore incapable of passing on their genes – benefit the queen, who is essentially passing on copies of "their" genes. This is further elaborated in the "selfish gene" theory of Richard Dawkins, that the unit of evolution is not the individual organism, but the gene. (As stated by Wilson: "the organism is only DNA's way of making more DNA.") However, kinship selection works only where the individuals involved are closely related; it fails to explain the presence of altruism and cooperation between unrelated individuals, particularly across species. In a 1971 paper
The Selfish Gene and Natural Selection" (1966), which argued that altruism is not based upon group benefit "per se", but is a result of selection that occurs "at the level of the gene mediated by the phenotype" and any selection at the group level occurred only under rare circumstances. This approach was developed further during the 1960s by W. D. Hamilton and others who opposed group selection and selection aimed directly at benefit to the individual organism: Despite the principle of 'survival of the fittest' the ultimate criterion which determines whether [a gene] "G" will spread is not whether the behavior is to