Genetic analysis of evolutionary processes

Abstract

Epigean and cave populations of A. fasciatus (Characidae, Pisces) differ in a series of morphological physiological, and ethological features. The interfertility of these populations made possible a genetic analysis of organs characteristic of interspecific divergence. The study of the regressive organs “eye” and “melanophore system” on the one hand and that of the constructively improved “gustatory equipment" and "feeding behaviour” on the other yielded identical principles of genetic manifestation: (1) All features have a polygenic basis with an at least di- to hexahybrid inheritance. (2) All polygenes have the same amount of expressivity. (3) After recombination of a minimum number of genes, discontinuous distributions (threshold effects) develop. (4) All features are independently inherited. (5) The genes responsible for a feature are unspecific. In the case of the eye this means that no “lens-” or “retinagenes” are analyzed; due to developmentally physiological interdependence within complex structures, only so-called “eye-genes” have as yet been described. Because of the developmentally physiological interdependence of complex organs, the process of reduction proceeds as a diminution in size, that of constructive evolution as enlargement. In both cases different allometric correlations of the single structures can be found. The convergent reduction of eyes in cave animals is caused by the loss of stabilizing selection which normally keeps the eye in its appropriate adapted form. It is not directional selection pressure, like f. ex. energy economy, but mutation pressure that causes eye reduction. By this, random mutations, which are mostly of deleterious character, are accumulated. The principles of regressive evolution are not restricted to the development of cave species. The absence of stabilizing selection regularly occurs during transitional evolutionary phases. These are f. ex. initial stages of speciation which may be observed when biotopes with little or no interspecific competition are colonized by an invader. Genotypic and phenotypic variability now arise and equilibria become punctuated, because stabilizing selection for a specific ecological niche which has once been acquired by the invading species is no longer acting. Examples include the evolution of species flocks in geologically young lakes or oceanic islands. Rapidly increasing variability now secondarily provides the material for directional selection which radiates such species into vacant niches. Genetic threshold effects as described above may accelerate this process. Variability will finally become lower again under the influence of inter- and intraspecific competition. A new equilibrium is attained.