The role of fungal NADPH oxidase complexes in differentiation and virulence

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Reactive oxygen species (ROS) generated by NADPH oxidases (Nox) play an important role in pathogen defense reactions and a broad variety of differentiation processes in animals and plants. There is growing evidence that also in fungi these membrane complexes are involved in all major developmental processes: they have impact on polar growth, sexual differentiation and virulence. Still there is considerable lack of knowledge on complex composition, recruitment and localization, and on the molecular mechanisms underlying their impact on so many important cellular processes. Based on the broad analysis in the first project period of the Nox complexes and ROS signaling components in general in two fungi with completely different lifestyle and obviously significantly different Nox functions - the biotrophic ergot fungus Claviceps purpurea and the necrotrophic grey mold fungus Botrytis cinerea-, we will focus in this second period on a detailed functional characterization of the Nox complexes in the model system B.cinerea. We will investigate the specific composition/recruitment of the Nox complexes, the localization, linked to specific developmental stages/metabolic activities, focusing on a possible function in the ER. We will study the impact of Nox on the cytoskeleton, in the context of differentiation processes (e.g.penetration structures) and analyse regulatory circuits which affect Nox activity in mammalian systems, e.g. lipid signaling. In addition, we will try to identify alternative ROS sources, based on existing ROS overproducing mutants.