Explaining the persistence of populations is an important quest in ecology, and is a modern manifestation of the balance of nature metaphor. Increasingly, however, ecologists see populations (and ecological systems generally) as not being in equilibrium or balance. The portrayal of ecological systems as ''non-equilibrium'' is seen as a strong alternative to deterministic or equilibrium ecology, but this approach fails to provide much theoretical or practical guidance, and warrants formalisation at a more fundamental level. This is available in adaptation theory, which allows population persistence to be explained as an epiphenomenon stemming from the maintenance, survival, movement and reproduction of individual organisms. These processes take place within a physicochemical and biotic environment that persists through structured annual cycles, but which is also spatiotemporally dynamic and subject to stochastic variation. The focus is thus shifted from the overproduction of offspring and the consequent density dependent population pressure thought to follow, to the adaptations and ecological circumstances that support those relatively few individuals that do survive.