weak selection
Lion S, Jansen VAA & Day T 2011 Evolution in structured populations: beyond the kin versus group debate. TREE 26:193-201.
- biological assumptions
- the parallel between the MS and IF methodologies is sometimes obfuscated because many practitioners of the IF methodology in fact make two additional, interrelated simplifying assumptions as a routine part of the IF partitioning
- in such cases, the IF partitioning not only yields conceptual insight, but also provides a means of making analytical progress
- (i) fitness is linearly related to genotype
- (ii) selection is weak
- (i.e. mutations have small phenotypic effects)
- the selection coefficient is often still too difficult to calculate analytically because the expected value of a neighbour's genotype will depend on the action of the mutant allele
- if we further assume that selection is weak, however, then these expected genotypes can be calculated under the assumption that the mutant allele is neutral
- if mutations have small effects, as typically assumed in IF methodology, fitness will be approximately linear in the genotype
- hence, these two assumptions are distinct, but interrelated
- much of IF theory typically assume mutations of small phenotypic effects
- it has sometimes been argued that IF cannot deal with strong selection while the MS methodology can
- however, partitioning of the selection coefficient between within-group and between-group components does not make its analytical calculation any simpler under strong selection
- in other words, the MS methodology does not provide a superior characterization of genetic structure under strong selection
- in fact most MS models also typically resort to a weak selection approximation or assume linear fitness to make analytical progress
- the relevant question is not what approach is inherently better, but how to derive analytical approximations to characterize genetic structure under strong selection in an empirically meaningful way
- is one interpretation better than the other?
- neutral measures of genetic associations are central components of the IF methodology
- for neutral alleles, the expected value of a neighbour's genotype corresponds exactly to the genealogical definition of relatedness and can be calculated for any type of population structure using standard population genetics techniques
- under weak selection, the genealogical relatedness among individuals provides a sufficient description of the population's genetic structure to make evolutionary predictions
- in the light of a recent article [9] that gives a misleading representation of current IF theory, we stress that extensions of the theory we have presented exist for fluctuating demography [32–34], non-linear genotype-phenotype maps [35] and non-pairwise interactions [36]
- mutation and strong selection
- when mutations are rare and have a small phenotypic effect (weak selection), genetic and demographic structuring tend to be decoupled
- the evolution of reduced parasite virulence in structured populations can be understood as a form of altruism among genetically related parasites, and thereby falls under the scope of social evolution theory
- these results typically rely on the assumption that mutation rates are low and that selection is weak