I turn now to one of the central objections to the gene's-eye view: that it or ignores, or obscures, the causal structure of selection processes. Construing evolution as the substitution of fitter genes for their alleles is all very well, the objection goes, but it tells us nothing about why the substitutions have occurred. Thus gene's-eye theorists are accused of 'confusing bookkeeping with causality'. Versions of this objection have beenmadeby Wright (1930), Mayr (1963), Sober and Lewontin (1982), Sober (1984), Brandon (1990), Gould (2001, 2002), and others.
Proponents of this argument usually allow that the outcome of selection can be described in genic terms. Thus for example, Gould (2001) accepts that gene frequencies provide the best way of 'keeping the books' of evolution, since selection at all levels does eventuate in gene frequency change, he thinks. So the gene's-eye perspective is always available, he argues, but rarely useful, for it omits causal information.
There are two separate issues here. First, is it true that the outcome of any selection process can be described in genic terms? Is Gould right to concede this point to the gene's-eye theorists? Secondly, in cases where the gene's-eye perspective is available, what determines whether it is useful? I tackle these questions in turn.
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