In the phytopathogenic fungus Ustilago hordei, one locus with two alternate alleles, MAT.1 and MAT-2, controls mating and the establishment of the infectious dikaryon (bipolar mating). In contrast, for U. maydis, these functions are associated with two different gene complexes, called a and b (tetrapolar mating); the a complex has two alternate specificities, and the b gene complex is multiallelic. We have found homologs for the b gene complex in U. bordei and have cloned one from each mating type using sequences f" one bEast allele of U. maydis as a probe. Sequence analysis revealed two divergent open reading frames in each b complex, which we called bW (bWest) and bE (bEast) in analogy with the b gene complex of U. maydis. The predicted bW and bE gene products from the two different mating types showed .v75% identity when homologous polypeptides were compared. All of the characterized bW and bE gene products have variable amino-terminal regions, consenred carboxy-terminal regions, and similar homeodomain motifs. Sequence comparisons with the bW1 and bE1 genes of U. maydis showed conservation in organization and structure. Ransformation of the U. bordei b gene complex into a U. bordei strain of opposite mating type showed that the b genes from the two mating types are functional alleles. The U. hordei b genes, when introduced into 0. maydis, rendered the haploid transforniants weakly pathogenic on maize. These results indicate that structurally and functionally consenred b genes are present in U, bordei.