Chromosomal speciation: a reply

The “genic” and the “non-genic” (chromosomal) hypotheses for the predominant mechanism by which species diverge into two have long been in contention. In 1998 Coyne and Orr attacked certain formulations of the chromosomal hypothesis on the grounds that they required macromutations (structural changes in chromosomes). In 1999 I replied that numerous independent micromutations (single DNA base changes) should suffice (GC% hypothesis). Kliman et al., with the support of Coyne and Charlesworth, have presented various counterarguments, to which the present paper responds with evidence that GC% differences are primary to genic differences and would operate by changing the structure of stem–loops extruded from duplex DNAs. Chromosomes attempting to align by means of complementary loop–loop interactions would fail if GC% differences exceeded a critical threshold. This would disrupt meiosis (hybrid sterility) and the parents of organisms with failed meiosis would be reproductively isolated from each other. If they could find new mates with which they were GC-compatible, then new species could emerge. The model leads to predictions consistent with several lines of evidence. The GC% version of the chromosomal hypothesis has a sound basis and deserves at least as much attention as its genic rival.