Species as Individuals in the Hierarchical Theory of Selection 631
All major proponents of gene selectionism have unintentionally illustrated the
theory's incoherence by trying to "cash out" their system, and failing at the crucial
point of assigning causal agency in natural selection. For, however these
proponents may talk about genes as primary agents or units of selection, they
cannot deny that nature's Darwinian action generally unfolds between discrete
organisms and their environments. These authors therefore acknowledge this basic
fact and then tend to lapse into verbal obfuscation on the gene's behalf. I have
already noted a prime example in Williams's claim, quoted previously in another
context, that organismic selection should be regarded not "as a level of selection in
addition to that of the gene, but as the primary mechanism of selection at the genie
level." But what does this statement mean? Williams recognizes organismic
selection as the "primary mechanism" by which genes pass differentially from one
generation to the next. But primary mechanisms are efficient causes in any
standard analysis of the logic of science. Williams (1992, p. 38) presents an
accurate epitome of selection in the following passage: he states that selection must
always operate on interactors (and he knows, as the previous quotation shows, that
organisms usually constitute the relevant interactors in cases that he wishes to
describe as genie selection); he also recognizes that information must pass to future
generations by faithful heredity, and he seems to acknowledge that such biased
passage defines the result, not the cause, of selection. Yet he fails to take the final
step of acknowledging that these statements debar gene selectionism as the
mechanism of evolution. "Natural selection must always act on physical entities
(interactors) that vary in aptitude for reproduction, either because they differ in the
machinery of reproduction or in that of survival and resource capture on which
reproduction depends. It is also necessary that there be what Darwin called 'the
strong principle of inheritance,' so that events in the material domain can influence
the codical record. Offspring must tend to resemble their own parents more than
those of other offspring. Whenever these conditions are found there will be natural
selection."
Over the years, Dawkins has developed a litany of similar admissions. Of
course organisms must be regarded as the foci of selection, but since biased gene
passage occurs as a result of this process, we may identify genes as agents of
selection. (But results are not causes, although foci of action surely are): "Just as
whole boats win or lose races, it is indeed individuals [organisms] who live or die,
and the immediate manifestation of natural selection is nearly always the
individual level. But the long-term consequences of non-random individual death
and reproductive success are manifested in the form of changing gene frequencies
in the gene pool" (1976, p. 48).
Dawkins then apologizes for framing his descriptions in terms of organisms
as causal actors, excusing himself for succumbing to temptations of convenience.
(But perhaps we find this mode "convenient" because we achieve the best
description of a causal reality thereby—while the genie mode remains tortuous and
uncomfortable because we sense the central error in such formulation): "In practice
it is usually convenient, as an approximation, to regard the individual body as an
agent 'trying' to increase the number of all its