Evolution of Mhc class IIB genes in Darwin's finches and their closest relatives:: birth of a new gene

The 15 extant species of Darwin's finches on the Galapagos and Cocos Islands are the products of an unfinished adaptive radiation from a founder flock of birds related to the South American species Tiaris obscura. Molecular characterization of their major histocompatibility complex (Mhc) class 11 B genes has revealed the existence of several related groups of sequences (presumably encoded in distinct loci) from which one (group 5) stands out because of its low divergence over extended time periods. Analysis of group 5 exon 2 and intron 2 sequences has revealed that the encoding locus apparently arose 2-3 million years ago in the Tiaris group of South and Central American Thraupini. The locus shows no evidence of inactivation, but displays a very low degree of polymorphism, both in terms of number of alleles and genetic distances between alleles. Some of the polymorphism. however. appears to be trans-specific. All the observed intergenic differences can be explained by point mutations and most of the exon 2 changes represent nonsynonymous substitutions, although the rate of nonsynonymous and synonymous substitutions appears to be the same. The origin of the new locus is explained by the birth-and-death model of Mhc evolution with two important extensions. First, the ancestor of the group 5 genes may have arisen without new gene duplication and second, the birth of the new group may have been brought about by a switch from balancing to directional selection. The ancestor of the group 5 genes may have been a classical class 11 B allele (one of many) which directional selection fixed in the ancestral population and drove into the category of nonclassical genes.