soft sweep
Cutter AD & Payseur BA 2013 Genomic signatures of selection at linked sites: unifying the disparity among species. Nat Rev Genet 14:262-274.
- mutation-limited adaptation
- when mutational input into a population is sufficiently low, the rate of adaptation will be limited by the input of new beneficial mutations
- this common theoretical assumption will be violated in real species with populations that are large, subdivided or subject to frequent changes in selective regime
- this notion of mutation-limited adaptation at the molecular level might not be the norm in nature
- in Drosophila simulans, one approach that can quantify only hard sweeps suggests that ~13% of replacement-site substitutions were fixed in such a manner versus ~90% of such sites inferred to have fixed through either hard or soft sweeps
- soft sweeps could represent upwards of 85% of positively selected substitutions
- formal analyses are desperately needed to disentangle the relative contributions of hard and soft sweeps to divergence
- the relative incidence of hard and soft sweeps is a major unsolved problem
- relaxing the classic hard sweep assumption that selected alleles are rapidly driven to fixation generates a new suite of predictions
- a challenge is to determine why the incidence of hard versus soft sweeps in adaptation might differ among organisms and whether it varies in predictable ways
- species-wide soft sweeps could dominate when populations are subdivided
- the few theoretical studies to date on polygenic selective sweep effects suggest weaker signatures of selection at linked sites
- when a beneficial mutation is linked to deleterious mutations, those mildly deleterious mutations can hitch-hike to fixation
- the fixed region will be susceptible to repeated bouts of positive selection as compensatory alleles arise and replace the detrimental alleles that had hitch-hiked with the original beneficial mutation
- the linkage to deleterious mutations slows the fixation process and increases the incidence of recombinant genetic backgrounds, obscuring the signatures of a selective sweep
- even without positive selection, purifying selection against multiple linked targets similarly reduces the efficacy of selection
- this mitigation of classic background selection effects, owing to Hill–Robertson interference, has largely been explored through simulation
- a complete mathematical association with recombination rate has not yet been established
- fitness-class coalescent modelling appears to be promising
- much effort has been devoted to distinguishing background selection from hitch-hiking effects of recurrent selective sweeps
- this is difficult and is perhaps a false dichotomy
- background selection should be taken into account as a part of a null model of molecular evolution
- deleterious mutations arise inexorably and in turn are subjected to purifying selection