Mushrooms, such as Coprinus cinereus, possess large families of pheromones and G-protein-coupled receptors that are sequestered at the B mating-type locus and whose function is to confer vast numbers of different mating types. This ability results from complex patterns of cognate and noncognate pheromone/ receptor pairings, which potentially offer a unique insight into the molecular interaction between receptor and ligand. In this study we have identified many more members of these families by molecular analysis of strains collected worldwide. There are three groups of genes at each B locus. We have identified two alleles of group 1, five alleles of group 2, and seven alleles of group 3, encoding in total 14 different receptors and 29 different pheromones. The specificity of many newly identified alleles was determined by transformation analysis. One striking finding was that receptors fall into groups based on sequence homology but these do not correspond to the groups defined by position, indicating that complex evolutionary processes gave rise to the B loci. While additional allelic versions may occur in nature, the number of B specificities possible by combination of the alleles that we describe is 70, close to previous estimates based on population analysis.
T HE role of pheromone signaling in fungal matingand receptors were first identified as mating-type deterhas largely been elucidated from studies of the budminants and map to what has been designated the B ding yeast Saccharomyces cerevisiae. This fungus has just mating-type locus. A second mating-type locus, A, entwo mating types and pheromones are secreted to act codes the subunits of a transcription factor belonging as chemoattractants for identifying compatible mating to the homeodomain family (Kü es et al. 1994a) that is partners. Binding of a pheromone to an appropriate necessary, together with pheromone signaling, to proreceptor on the cell surface triggers an intracellular mote the initial stages of sexual development. Compati-G-protein-linked MAP kinase cascade that results in ble A genes encode versions of the proteins that can changes in growth direction to permit cell fusion, which heterodimerize following cell fusion, an interaction that is then followed immediately by nuclear fusion (reis analogous to that between the MATa1 and MAT␣2 viewed by Kurjan 1993). Once cells are diploid, pheromating-type proteins in mated cells of S. cerevisiae (remone signaling ceases.viewed by Johnson 1995). Pheromone signaling also plays an essential role in In contrast to S. cerevisiae, compatible cell fusion in mating in basidiomycete fungi (reviewed by Casselton C. cinereus is not followed immediately by nuclear fusion, and Olesnicky 1998), but in mushroom species such as but by an extended vegetative phase in which the nuclei Coprinus cinereus, unlike S. cerevisiae, there is no evidence from each mate remain paired in each cell, a phase that pheromones have a role in mate attraction (Olesknown as the dikaryophase or dikaryon. The B matingnicky et ...