Muller's Ratchet and the evolution of supernumerary chromosomes

Abstract

Supernumerary chromosomes arise from portions of the normal chromosome complement through nondisjunction, fragmentation, or other mechanisms. Once present in the genome, they are subject to virtually the same genetic conditions that affect the evolutionary degeneration of heteromorphic sex chromosomes. Y or W chromosomes occur only in the presence of X or Z chromosomes, respectively, just as supernumeraries never occur except in the presence of the complete regular karyotype containing their progenitor sequences. Thus, mechanisms that can account for the evolution of sex-chromosome heteromorphism can also be invoked to explain the degeneration process of supernumerary chromosomes after their origination. Incipient supernumeraries initially have genes identical with those on progenitor chromosomes. This frees them from the evolutionary constraint of carrying nonduplicated genetic information, just as in Y and W chromosomes during early stages of sex-chromosome differentiation. The degeneration of supernumerary chromosomes may thus proceed via the mechanism of Muller's Ratchet. This hypothesis predicts that supernumerary chromosomes should lose functional loci, lose sequence homology with the regular genome, and gain heterochromatin over time, resulting in multiple heteromorphic forms of degenerate supernumeraries within and between populations, as is commonly observed.Key words: supernumerary chromosomes, B chromosomes, evolution, origin, Muller's Ratchet.