In such species, selection maintains lean and mostly functional melanogaster-like genomes. Although the neutral theory will survive Andolfatto's demonstration that there is pervasive selection in Drosophila , its original justification may not. Kimura 1 claimed that most substitutions must be neutral because positive selection driving all of them would incur too high a fitness cost. So Andolfatto's conclusion that one selection-driven substitution has occurred about every ten generations since D.
If flies can do it, so could others, invalidating Kimura's original argument. In Drosophila , the relatively junk-free regions between genes, which probably regulate gene expression, seem to be a major target of positive selection.
It is therefore fair to assume that the accumulation of beneficial mutations also had a major role in the evolution of functional segments in the intergenic regions in mammal-like genomes. However, nature is queerer than we may think. Since the human—chimpanzee divergence, functionally important intergenic segments have incorporated many deleterious mutations, rather than beneficial ones, perhaps because of the relatively recent decline of the effective population size in hominids We do not yet know how quickly beneficial mutations accumulate in functional segments of mammal-like genomes in the lineages where efficiency of selection did not decline.
The total number of functionally important nucleotides in the genome, T , is crucially important for estimating the genomic deleterious mutation rate U , a key parameter in evolutionary genetics Andolfatto's data and analysis suggest that, when U is estimated for D.
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