Richard Dawkins on Visible Changes and Adaptationism

 
On another thread [Visible Mutations and Evolution by Natural Selection] we are discussing a common adaptationist claim that once a mutation has a visible phenotype it is almost certainly subject to selection. Some people have questioned whether there is anyone who actually believes in such a thing. Here's Richard Dawkins writing in The Extended Phenotype (1982).

The biochemical controversy over neutralism is concerned with the interesting and important question of whether all gene substitutions have phenotypic effects. The adaptationism controversy is quite different. It is concerned with whether, given that we are dealing with a phenotypic effect big enough to see and ask questions about, we should assume that it is the product of natural selection. The biochemist's 'neutral mutations' are more than neutral. As far as those of us who look at gross morphology, physiology and behaviour are concerned, they are not mutations at all. It was in this spirit that Maynard Smith (1976b) wrote: "I interpret 'rate of evolution' as a rate of adaptive change. In this sense, the substitution of a neutral allele would not constitute evolution ..." If a whole-organism biologist sees a genetically determined difference among phenotypes, he already knows he cannot be dealing with neutrality in the sense of the modern controversy among biochemical geneticists.

In 2007 Dawkins would probably admit to some neutral examples of "genetically determined differences among phenotypes" but his position hasn't changed very much from 1982. For example, in The Ancestor's Tale (2005) he writes,

Contrary to my rather ludicrous reputation as an "ultra-Darwinist" (a slander I would protest more vigorously if the name sounded less of a compliment than it does), I do not think that the majority of evolutionary change at the molecular level is favoured by natural selection. On the contrary, I have always had a lot of time for the so-called neutral theory associated with the great Japanese geneticist Motoo Kimura, or its extension, the "nearly neutral" theory of his collaborator Tomoko Ohta. The real world has no interest in human tastes, of course, but as it happens I positively want such theories to be true. This is because they give us a separate, independent chronicle of evolution, unlinked to the visible features of the creatures around us., and they hold out the hope that some kind of molecular clock might really work.

Just in case the point is misunderstood, I must emphasize that the neutral theory does not in any way denigrate the importance of selection in nature. Natural selection is all-powerful with respect to those visible changes that affect survival and reproduction. Natural selection is the only explanation we know for the functional beauty and apparently "designed" complexity of living things. But if there are any changes that have no visible effect—changes that pass right under natural selection's radar—they can accumulate in the gene pool with impunity and may supply just what we need for an evolutionary clock.

Pluralists believe that all kinds of alleles are neutral or nearly neutral and are fixed in a population by random genetic drift. This includes alleles that produce a visible phenotype. Pluralists do not believe that there is a major distinction between the mechanisms of evolution at the molecular level and the mechanisms at the morphological level.

Contrary to Dawkins, I believe that Neutral Theory has reduced the importance of selection in nature. Prior to 1968 it was common to attribute almost all changes to natural selection and it was common to advocate that the presence of variation in a population was due to balancing selection. Today, one has to consider the evidence for adaptation; you can no longer just assume that it is the only game in town.

Of course it's true that natural selection is the only mechanism that affects allele frequencies once you can demonstrate that a visible change affects survival and reproduction. But Dawkins goes farther than that. He strongly implies that all visible phenotypes are subject to selection and neutral alleles are confined to the molecular level.