First, the idea of 'coadapted genotypes'. A gene has the particular effect that it does only because there is an existing structure upon which to work. A gene can't affect the wiring up of a brain unless there is a brain being wired up in the first place. There won't be a brain being wired up in the first place, unless there is a complete developing embryo. And there won't be a complete developing embryo unless there is a whole program of chemical and cellular events, under the influence of lots and lots of other genes, and lots and lots of other, non-genetic, causal influences. The particular effects that genes have are not intrinsic properties of those genes. They are properties of embryological processes, existing processes whose details may be changed by genes, acting in particular places and at particular times during embryonic development. We saw this message demonstrated, in elementary form, by the development of the computer biomorphs.
In a sense, the whole process of embryonic development can be looked upon as a cooperative venture, jointly run by thousands of genes together. Embryos are put together by all the working genes in the developing organism, in collaboration with one another. Now comes the key to understanding how such collaborations come about. In natural selection, genes are always selected for their capacity to flourish in the environment in which they find themselves. We often think of this environment as the outside world, the world of predators and climate. But from each gene's point of view, perhaps the most important part of its environment is all the other genes that it encounters. And where does a gene 'encounter' other genes? Mostly in the cells of the successive individual bodies in which it finds itself. Each gene is selected for its capacity to cooperate successfully with the population of other genes that it is likely to meet in bodies.