As a starting point for a phylogenetic study of self-incompatibility (SI) in crucifers and to elucidate the genetic basis of transitions between outcrossing and self-fertilizing mating systems in this family, we investigated the SI system of Arabidopsis lyrata . A. lyrata is an outcrossing close relative of the self-fertile A. thaliana and is thought to have diverged from A. thaliana ف 5 million years ago and from Brassica spp 15 to 20 million years ago. Analysis of two S (sterility) locus haplotypes demonstrates that the A. lyrata S locus contains tightly linked orthologs of the S locus receptor kinase ( SRK ) gene and the S locus cysteine-rich protein ( SCR ) gene, which are the determinants of SI specificity in stigma and pollen, respectively, but lacks an S locus glycoprotein gene. As described previously in Brassica , the S haplotypes of A. lyrata differ by the rearranged order of their genes and by their variable physical sizes. Comparative mapping of the A. lyrata and Brassica S loci indicates that the S locus of crucifers is a dynamic locus that has undergone several duplication events since the Arabidopsis-Brassica split and was translocated as a unit between two distant chromosomal locations during diversification of the two taxa. Furthermore, comparative analysis of the S locus region of A. lyrata and its homeolog in self-fertile A. thaliana identified orthologs of the SRK and SCR genes and demonstrated that self-compatibility in this species is associated with inactivation of SI specificity genes.
INTRODUCTIONSelf-incompatibility (SI) is the major outcrossing mechanism in the family Brassicaceae (de Nettancourt, 1977). Species in this family have been grouped into 19 tribes on the basis of morphological criteria (Schultz, 1936), and SI has been described in all tribes analyzed to date. When Bateman (1955) surveyed 182 species distributed in 11 tribes, he found that approximately half of these species included selfincompatible accessions. In a survey of 59 taxa in the subtribe Brassicineae of the tribe Brassiceae (which includes Brassica and Raphanus ), 50 taxa were self-incompatible (Takahata and Hinata, 1980). In all cases analyzed, SI has been shown to be controlled sporophytically by a single S (sterility) locus, with multiple alleles or variants and complex dominance relationships between alleles (Bateman, 1954(Bateman, , 1955Thompson and Taylor, 1966): in self-incompatible plants, pollen will not develop on a stigma that expresses the same S alleles as the pollen parent.Molecular analysis of the Brassica S locus region has shown that this mendelian locus is a gene complex consisting of distinct stigma-expressed and anther-expressed genes that determine SI specificity in stigma and pollen, respectively (reviewed in Nasrallah, 2000). The SRK (for S locus receptor kinase) gene (Stein et al., 1991) encodes a plasma membrane-spanning receptor serine/threonine kinase specific to the stigma epidermis (Stein et al., 1996) and is the determinant of SI specificity in the stigma (Takasaki et al., 2000). ...