How many processes are responsible for phenotypic evolution?

In addressing phenotypic evolution, this article re-considers natural selection, random drift, developmental constraints and internal selection in the new extended context of evolutionary developmental biology. The change of perspective from the ‘evolution of phenotypes’ towards an ‘evolution of ontogenies’ (evo-devo perspective) affects the reciprocal relationships among these different processes. Random drift and natural selection are sibling processes: two forms of post-productional sorting among alternative developmental trajectories, random the former, non-random the latter. Developmental constraint is a compound concept; it contains even some forms of natural (‘external’ and ‘internal’) selection. A narrower definition (‘reproductive constraints’) is proposed. Internal selection is not a selection caused by an internal agent. It is a form of environment-independent selection depending on the level of the organism’s internal developmental or functional coordination. Selection and constraints are the main deterministic processes in phenotypic evolution but they are not opposing forces. Indeed they are continuously interacting processes of evolutionary change, but with different roles that should not be confused.