The Clp1 Protein Is Required for Clamp Formation and Pathogenic Development ofUstilago maydis

In the phytopathogenic fungus Ustilago maydis, pathogenic development is controlled by a heterodimer of the two homeodomain proteins bE and bW, encoded by the b-mating-type locus. We have identified a b-dependently induced gene, clampless1 (clp1), that is required for the proliferation of dikaryotic filaments in planta. We show that U. maydis hyphae develop structures functionally equivalent to clamp cells that participate in the distribution of nuclei during cell division. In clp1 mutant strains, dikaryotic filaments penetrate the plant cuticle, but development is stalled before the first mitotic division, and the clamp-like structures are not formed. Although clp1 is immediately activated upon b-induction on the transcriptional level, nuclear-localized Clp1 protein is first observed at the stage of plant penetration prior to the first cell division. Induced expression of clp1 strongly interferes with b-dependent gene regulation and blocks b-dependent filament formation and b-dependent cell cycle arrest. We speculate that the Clp1 protein inhibits the activity of the bE/bW heterodimer to facilitate the cell cycle progression during hyphal growth.