Evolutionary Aspects of the S-Related Genes of the Brassica Self-Incompatibility System: Synonymous and Nonsynonymous Base Substitutions

In the Brassicaceae, self-vs. nonself-recognition in self-incompatibility is controlled by sporophytic S-alleles. Haplotypes specifying both SRK (S-receptor kinase) and SLG (S-locus glycoprotein) are considered to play an important role in the recognition reactions. We compared the nucleotide sequences of SRK(9)(Bc) and SRK(6)(Bo). The number of nonsynonymous substitutions per site (P(n)) was lower, constrained, in the kinase than the receptor domain, while the numbers of synonymous substitutions (P(s)) in the two domains were largely comparable. Pairwise values for P(s) and P(n) were calculated among 17 operational taxonomic units, including eight SLGs, the receptor domains of two SRKs, four SRAs (S-related A) and three SRBs (S-related B), which have high homologies with each other. The values of P(s) and P(n) of SLG were mostly comparable to those of the receptor domain of SRK. Dendrograms constructed on the basis of P(n) and P(s) indicated that SRA differentiated first, followed by SRB. The differentiation of SLG alleles is one of prerequisite factors for the establishment of self-incompatibility, and the allelic differentiation has occurred more than tens of million years ago.