The gene's eye view, the Gouldian knot, Fisherian swords and the causes of selection.

The biological units-of-selection debate has centred on questions of which units experience selection and adaptation. Here, I use a causal framework and the Price equation to develop the gene's eye perspective. Genes are causally special in being both replicators and interactors. Gene effects are tied together in a complex Gouldian knot of interactions, but Fisher deployed three swords to try to cut the knot. The first, Fisher's average excess, is non-causal, so not fully satisfactory in that respect. The Price equation highlights Fisher's other two swords, choosing to model only selection, and only the part that is transmissible across generations. The models developed here show that many causes of organismal fitness do not cause Gouldian complications. Only two kinds of elements must be added to the focal gene for a causal explanation of its selective change: co-replicators that are associated with the focal gene and co-interactors that interact non-additively with the focal gene. Identical equations for co-replication and co-interaction describe interactions between gene copies at a single locus or at separate loci, and also for genes situated within the same individual or in different individuals. These results resolve some of the objections to the gene's eye view. This article is part of the theme issue 'Fifty years of the Price equation'.