George C. Williams
Adaptation and Natural Selection
Princeton, NJ: Princeton University Press, 1996/1966.

Many of the contributions of evolutionary thought in the past century can be put in one of two opposed groups. One group emphasizes natural selection as the primary or exclusive creative force. The other minimizes the role of selection in relation to other proposed factors. R. A. Fisher (1930, 1954) showed that many of the proposed alternatives could be discounted with the acceptance of Mendelian genetics and a logical investigation of its relation to selection. Even without Mendelian genetics, Weismann (1904) effectively championed natural selection against some of its rivals of the nineteenth century. His only serious errors are traceable to his ignorance of the Mendelian gene.

The contest was decisively won by natural selection, in my opinion, when by 1932 the classic works of Fisher, Haldane, and Wright had been published. Yet even though this theory may now reign supreme, its realm still supports some opposition, perhaps more than is generally realized. Many recent discussions seem on the surface to conform to the modern Darwinian tradition, but on careful analysis they are found to imply something rather different. I believe that modern opposition, both overt and cryptic, to natural selection, still derives from the same sources that led to the now discredited theories of the nineteenth century. The opposition arises, as Darwin himself observed, not from what reason dictates but from the limits of what the imagination can accept. It is difficult for many people to imagine that an individual's role in evolution is entirely contained in its contribution to vital statistics. It is difficult to imagine that an acceptable moral order could arise from vital statistics, and difficult to dispense with belief in a moral order in living nature. It is difficult to imagine that the blind play of the genes could produce man. Major difficulties also arise from the current absence of rigorous criteria for deciding whether a given character is adaptive, and, if so, to precisely what is it an adaptation. As I will argue at some length, adaptation is often recognized in purely fortuitous effects, and natural selection is invoked to resolve problems that do not exist. If natural selection is shown to be inadequate for the production of a given adaptation, it is a matter of basic importance to decide whether the adaptation is real.

(From the introduction, pp. 3-4)

The essence of the genetical theory of natural selection is a statistical bias in the relative rates of survival of alternatives (genes, individuals, etc.). The effectiveness of such bias in producing adaptation is contingent on the maintenance of certain quantitative relationships among the operative factors. One necessary condition is that the selected entity must have a high degree of permanence and a low rate of endogenous change, relative to the degree of bias (differences in selection coefficients). Permanence implies reproduction with a potential geometric increase.

(From Chapter 1, "Natural selection, adaptation, and progress", pp. 21-22.)


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