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Our results imply that mode and speed of influenza evolution are governed not only by positive selection within, but also by background selection outside antigenic epitopes: immune adaptation and conservation of other viral functions interfere with each other.
Amino acid residues likely to have been influenced by positive selection within a protein can be identified by estimating ω with site-based maximum likelihood methods [ 35], even though this approach may be overly conservative [ 36].
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P -values represent the significance of positive selection inferred by the branch-site test for positive selection.
To test if the higher than mean dN/dS estimates for individual sex-associated genes might be caused by positive selection on individual sites within genes, we chose five genes for additional analysis among a larger set of species, ranging from seven to twelve taxa for the different genes (Additional file 2: Table S2).
The fraction α of insertions that were fixed by positive selection between species is very high within all compartments: It reaches approximately 60% in exons and approximately 67% in noncoding regions.
In some cases, only a few sites are affected by an episode of positive selection within a given branch.
Using a site-by-site approach, we found that the he gene has two sites under positive selection within codons in positions 134 and 340.
Studies with mammalian defensins suggested a rapid duplication followed by positive selection and diversification within each group [ 13- 18].
We present a phylogeny describing the evolutionary history of the FoxO gene family and show that the genes have evolved through duplications followed by purifying selection except for four sites in FoxO6 fixed by positive selection lie mostly within the non-conserved optimal PKB motif in the C-terminal part.
Positive selection of Adam1 in Mus is restricted to Adam1b (Table 1), a pattern mirrored by the paralogs in Glires, with no evidence of positive selection within Laurasiatheria (Table 2).
Furthermore, each of the four branch-site analyses detected three sites under positive selection within one of the epitopes presented by HLA alleles commonly observed in the Japanese population (with frequency >10%) [29].
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