polygenic adaptation

Stetter MG, Thornton K & Ross-Ibarra J 2018 Genetic architecture and selective sweeps after polygenic adaptation to distant trait optima. PLoS Genet 14:e1007794.

  • during the stationary phase before the shift and after reaching the new optimum we followed a Gaussian fitness function appropriate for a trait under stabilizing selection
  • during the optimum shift, however, such a model would be problematic, as only a few individuals in the upper tail of the fitness distribution would have extremely high relative fitness, inducing a strong population bottleneck
  • instead, we applied a model of truncation selection, first calculating fitness under the Gaussian fitness function but then assigning a fitness of 1 to the top half of the population and 0 to the bottom half
  • such a model is reasonable for sudden shifts in trait optima that do not lead to the extinction of a population, but where higher trait values are unambiguously advantageous and the maximum population size is limited
  • in natural populations these factors can be observed when sudden changes in the environment favor a specific phenotype for invasive species
  • we modeled 20 QTL resembling 50kb regions, each with a 4 kb "genic" region centered in a 46 kb "intergenic" region
  • fitness
  • w = exp[− (zzopt)2 / (2VS)] ... (1)
  • this standard model for traits under stabilizing selection is well suited for populations at equilibrium
  • under strong directional selection, however, this model greatly amplifies fitness differences among individuals in the tails of the phenotypic distribution
  • during the adaptive phase of the simulation, we calculated individual fitness following Eq 1, but then apply truncation selection by assigning a fitness of 1 to the top 50% of the distribution of w and 0 for the remaining 50%
  • this model allowed for truncation selection on z, while the population was distant from the new optimum, but allows for selection against phenotypes that surpass the new optimum during the final stages of adaptation
  • we stopped truncation selection once the population mean reached the new optimum
  • initial genetic variance
  • the genetic variance at equilibrium can be approximated by the house of cards (HoC) model
  • E[VG] = 4μVS ... (2)
  • and the stochastic HoC approximation
  • E[VG]SHC = 4μVS / (1 + VS / (m2)) ... (3)
  • background
  • computational limitations do not allow simulation of an entire eukaryotic genomes
  • we added a heritable background (GB) to our simulations to account for the adaptive potential of the rest of the genome
  • sweeps
  • we defined as a sweep any mutation that fixed faster than 99% of neutral alleles