Unpacking Conditional Neutrality: Genomic Signatures of Selection on Conditionally Beneficial and Conditionally Deleterious Mutations*

AbstractIt is common to look for signatures of local adaptation in genomes by identifying loci with extreme levels of allele frequency divergence among populations. This approach to finding genes associated with local adaptation often assumes antagonistic pleiotropy, wherein alternative alleles are strongly favored in alternative environments. Conditional neutrality has been proposed as an alternative to antagonistic pleiotropy, but conditionally neutral polymorphisms are transient, and it is unclear how much outlier signal would be maintained under different forms of conditional neutrality. Here, we use individual-based simulations and a simple analytical heuristic to show that a pattern that mimics local adaptation at the phenotypic level, where each genotype has the highest fitness in its home environment, can be produced by the accumulation of mutations that are neutral in their home environment and deleterious in nonlocal environments. Because conditionally deleterious mutations likely arise at a rat...