Adaptation, in contrast, is viewed here as a genuine non-equilibrium phenomenon: the genomic response to time-dependent selection. Our approach extends the static concept of fitness landscapes to dynamic fitness seascapes. It shows that adaptation requires a surplus of beneficial substitutions over deleterious ones.
We discuss the evolution of yeast and Drosophila genomes, providing examples where adaptive evolution can and where it cannot be inferred, despite the presence of positive selection.