Abstract

BackgroundAdaptive protein evolution is common in several Drosophila species investigated. Some studies point to very weak selection operating on amino-acid mutations, with average selection intensities on the order of Nes ~ 5 in D. melanogaster and D. simulans. Species with lower effective population sizes should undergo less adaptation since they generate fewer mutations and selection is ineffective on a greater proportion of beneficial mutations.ResultsHere I study patterns of polymorphism and divergence at 91 X-linked loci in D. miranda, a species with a roughly 5-fold smaller effective population size than D. melanogaster. Surprisingly, I find a similar fraction of amino-acid mutations being driven to fixation by positive selection in D. miranda and D. melanogaster. Genes with higher rates of amino-acid evolution show lower levels of neutral diversity, a pattern predicted by recurrent adaptive protein evolution. I fit a hitchhiking model to patterns of polymorphism in D. miranda and D. melanogaster and estimate an order of magnitude higher selection coefficients for beneficial mutations in D. miranda.ConclusionThis analysis suggests that effective population size may not be a major determinant in rates of protein adaptation. Instead, adaptation may not be mutation-limited, or the distribution of fitness effects for beneficial mutations might differ vastly between different species or populations. Alternative explanation such as biases in estimating the fraction of beneficial mutations or slightly deleterious mutation models are also discussed.

Highlights

  • Adaptive protein evolution is common in several Drosophila species investigated

  • Many studies aimed at detecting adaptive evolution have applied the McDonald-Kreitman (MK) test [7] or modifications of it, which contrasts the number of polymorphisms within a species to the number of substitutions between species at two classes of sites, a putatively neutral and a putatively selected class

  • High rates of protein adaptation in D. miranda If the rate of beneficial mutations and their selective effects are constant across species, larger populations are expected to show higher rates of adaptation, both because they generate more mutations and selection is effective on a greater proportion of mutations [18,46]

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Summary

Introduction

Adaptive protein evolution is common in several Drosophila species investigated. Many studies aimed at detecting adaptive evolution have applied the McDonald-Kreitman (MK) test [7] or modifications of it, which contrasts the number of polymorphisms within a species to the number of substitutions between species at two classes of sites, a putatively neutral and a putatively selected class. Several members in the Drosophila melanogaster species group show high rates of adaptive amino-acid evolution. Using the MK test and its extensions, about half (and up to 95%) of all amino-acid mutations fixed between species are inferred to be driven by positive selection [1,2,3,4]. Some uncertainty in estimates of , the fraction of aminoacid substitutions driven to fixation by adaptive evolution (page number not for citation purposes)

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