Published online by Cambridge University Press: 14 April 2009
Using models which describe the change, by natural selection, of the actual numbers of genes rather than their relative frequencies, it is demonstrated that the equation familiar to geneticists, i.e. dp/dt = sp(1 − p), is appropriate under a wide range of circumstances. It was pointed out that, for realistic treatment of the evolutionary process through which gene substitutions are repeated, the models must have the property such that the total population number remains constant or nearly constant throughout the process, and is not appreciably influenced by the genes being substituted.
The load or cost for a gene substitution was studied assuming a haploid population and the effects on the load of such factors as epistatic gene interaction in fitness, finite population number and slow change of environment were investigated. The load may become very large under a strong ‘reinforcing’ type epistasis between advantageous genes. In a finite population, the load for one gene substitution may be inflated by about unity if the product of the effective population number (Ne) and the selection coefficient (s) is large but Nesp0 is much smaller than unity, where p0 is the initial gene frequency. On the other hand, slow change of environment may decrease the load somewhat. It was concluded that despite these and other complicating factors, Haldane's original formula, –logep0, for a haploid population (−2 logep0 for the case of a diploid without dominance) is still useful for assessing the approximate amount of selective elimination that accompanies the process of gene substitution in evolution.