The Hamilton-Williams theory of evolution focuses on genes as the “beneficiaries” of natural selection. It is alleged that genes, and genes alone, fulfil the necessary condition of having high enough copying fidelity. But natural selection is a substrate-neutral template, that is, it could in principle apply to any entity that fulfilled Darwin's necessary conditions of heredity, variation, population growth and limited resources. For all that I have said so far, there is nothing special about DNA that makes it alone the only thing that could be the locus of natural selection. Moreover, the move from Darwin's own organism-centred view to the gene-centred view of the 1970s was generally perceived to preserve a fundamental core of Darwin's theory as the same substrate-neutral template was still being employed. “Pure” gene selectionists are only claiming that as a matter of empirical fact, DNA is the only thing with the required high copying fidelity, not that it is the only thing that in principle could have it. Dawkins himself says that life on any other planet would have to have come about by natural selection, even though it might have a very different chemical basis from life on earth (Dawkins 1983).

Not everyone accepts the gene selectionists' claim about the uniqueness of genes. There are dissenting views over what can be said to be inherited, what it is that natural selection acts on, and what it is that is benefited when natural selection produces traits.