The white rot fungus Schizophyllum commune is used for the analysis of mating and sexual development in homobasidiomycete fungi. In this study, we isolated the gene gap1 encoding a GTPase-activating protein for Ras. Disruption of gap1 should therefore lead to strains accumulating Ras in its activated, GTP-bound state and to constitutive Ras signaling. Haploid ⌬gap1 monokaryons of different mating types did not show alterations in mating behavior in the four different mating interactions possible in fungi expressing a tetrapolar mating type system. Instead, the growth rate in ⌬gap1 monokaryons was reduced by ca. 25% and ca. 50% in homozygous ⌬gap1/⌬gap1 dikaryons. Monokaryons, as well as homozygous dikaryons, carrying the disrupted gap1 alleles exhibited a disorientated growth pattern. Dikaryons showed a strong phenotype during clamp formation since hook cells failed to fuse with the peg beside them. Instead, the dikaryotic character of the hyphae was rescued by fusion of the hooks with nearby developing branches. ⌬gap1/⌬gap1 dikaryons formed increased numbers of fruitbody primordia, whereas the amount of fruitbodies was not raised. Mature fruitbodies formed no or abnormal gills. No production of spores could be observed. The results suggest Ras involvement in growth, clamp formation, and fruitbody development.The homobasidiomycetous white-rot fungus Schizophyllum commune has been used as a model system for the investigation of mating and sexual development for decades since it can be grown from spore to spore through its entire life cycle within 14 days on defined media, and it shows easily distinguished phenotypes for a tetrapolar mating behavior (32, 53). The tetrapolar mating system consists of two sets of mating type genes. The A mating type genes encode homeodomain transcription factors that are assumed to directly regulate gene expression (39,67). A multiallelic pheromone/receptor system is encoded by the B genes. Homology to the yeast pheromone system has led to the expectation that mating is in part controlled by a mitogen-activated protein kinase signal transduction cascade that is activated after stimulation of the G protein-coupled pheromone receptor (20,76,82).A fully compatible mating between two strains of S. commune occurs when both differ in their A and B gene specificities (A B ). The specificity of a locus is defined by a lack of activation of downstream developmental processes after crossing two strains with identical specificities in their mating type genes (20). Several steps of subsequent development can be distinguished. After cell fusion, septal breakdown and fast nuclear migration allow reciprocal nuclear exchange between the two mates. Migrant and resident nuclei pair, and dikaryotic hyphal tips are established. Subsequent conjugate nuclear division is accompanied by formation of clamp connections. Clamp connections are short, backwardly directed branches that fuse with the subapical cell and provide a bypass for one of the nuclei produced during synchronous division of the dikaryon, en...