We survey the distribution of haplotypes at the self-incompatibility (SI) locus of Arabidopsis lyrata (Brassicaceae) at 12 locations spread over the species' natural distribution in Iceland. Previous investigations of the system have identified 34 functionally different S haplotypes maintained by frequency-dependent selection and arranged them into four classes of dominance in their phenotypic expression. On the basis of this model of dominance and the island model of population subdivision, we compare the distribution of S haplotypes with that expected from population genetic theory. We observe 18 different S haplotypes, recessive haplotypes being more common than dominant ones, and dominant ones being shared by fewer populations. As expected, differentiation, although significant, is very low at the S locus even over distances of up to 300 km. The frequency of the most recessive haplotype is slightly larger than expected for a panmictic population, but consistent with a subdivided population with the observed differentiation. Frequencies in nature reflect effects of segregation distortion previously observed in controlled crosses. The dynamics of the S-locus variation are, however, well represented by a 12-island model and our simplified model of dominance interactions.
MOST homomorphic self-incompatibility (SI) systems are determined by a single multi-allelic locus (the S locus), composed of at least two genes. In the sporophytic self-incompatibility (SSI) system of Brassicaceae, the S-receptor kinase (SRK) is responsible for the determination of female specificity, and the S-cysteinerich (SCR) protein is responsible for the determination of male specificity (Schopfer et al. 1999;Takasaki et al. 2000). In a panmictic population with gametophytic SI the dynamics were modeled quite accurately by Sewall Wright (1939). Gametophytic self-incompatibility systems are simple, in that incompatibility is determined by matching of a haploid pollen with a diploid stigma expressing both its haplotypes codominantly. In a panmictic population all haplotypes are expected to have a frequency drawn from the same unimodal distribution. Sporophytic self-incompatibility systems are found within several families, in particular Brassicaceae (Bateman 1954). They are more complex to model because pollen phenotype derives from the genotype of the pollen parent, and dominance may occur in both stigma and pollen (Bateman 1952). In simple sex-symmetric dominance models, where selection on the male and female sides is equally strong, phenotypes are expected to have the same mean frequency (although with different distributions) leading to a higher expected frequency of relatively recessive haplotypes because they display their phenotype less often (Bateman 1952;Cope 1962;Sampson 1967;Imrie et al. 1972;Charlesworth 1988;Schierup et al. 1997;Charlesworth et al. 2000). In cases of more complicated dominance relations among the haplotypes, the homogeneity of phenotype frequencies fails (Schierup et al. 2006). Recessive haplotypes, how...