THE A-MATING-TYPE ALLELES OF USTILAGO-MAYDIS ARE IDIOMORPHS

Two unlinked incompatibility loci, a and b, control mating, pathogenicity, and sexual development in Ustilago maydis, a fungal pathogen of corn. Fusion of nonpathogenic haploid cells occurs when alleles of the a locus differ; fusion products that differ at the b locus will be pathogenic and complete sexual development. The two alleles of the a locus have been cloned. The a2 allele was isolated by exploiting the close linkage of the a locus to the genetic marker pan1. Several cosmids from a clonal a2 mating-type library allowed, via DNA-mediated transformation, an a1 mating type, pan1-1 auxotrophic recipient to grow prototrophically on medium lacking pantothenic acid. Cosmids containing functional a2 mating-type regions were identified by their ability to confer a2 mating behavior to the a1 recipient strain. The a1 allele was isolated from a plasmid library using DNA from the a2 region as a hybridization probe. The isolated clones contain a1 or a2 mating-type specificity as demonstrated by a plate mating assay, by pathogenicity tests on corn, and by gene replacement experiments. The a1 and a2 mating-type clones contain nonhomologous DNA segments that are flanked by similar nucleotide (nt) sequences. Restriction fragments containing a mating-type function are embedded within the nonhomologous DNA segments. U. maydis contains a single copy of either the a1 or a2 mating-type sequence within each haploid genome. This structural organization demonstrates that the a mating-type alleles of U. maydis are idiomorphs sensu Metzenberg and Glass (1990), nt sequences mapping to the same genetic locus that share no obvious evolutionary relationship.