The sporangiophore of Phycomyces blakesleeanus: a tool to investigate fungal gravireception and graviresponses.

The giant sporangiophore of the single-celled fungus, Phycomyces blakesleeanus, utilises light, gravity and gases (water and ethylene) as environmental cues for spatial orientation. Even though gravitropism is ubiquitous in fungi (Naturwissenschaftliche Rundschau, 1996, 49, 174), the underlying mechanisms of gravireception are far less understood than those operating in plants. The amenability of Phycomyces to classical genetics and the availability of its genome sequence makes it essential to fill this knowledge gap and serve as a paradigm for fungal gravireception. The physiological phenomena describing the gravitropism of plants, foremost adherence to the so-called sine law, hold even for Phycomyces. Additional phenomena pertaining to gravireception, specifically adherence to the novel exponential law and non-adherence to the classical resultant law of gravitropism, were for the first time investigated for Phycomyces. Sporangiophores possess a novel type of gravisusceptor, i.e. lipid globules that act by buoyancy rather than sedimentation and that are associated with a network of actin cables (Plant Biology, 2013). Gravitropic bending is associated with ion currents generated by directed Ca(2+) and H(+) transport in the growing zone (Annals of the New York Academy of Sciences, 2005, 1048, 487; Planta, 2012, 236, 1817). A set of behavioural mutants with specific defects in gravi- and/or photoreception allowed dissection of the respective transduction chains. The complex phenotypes of these mutants led to abandoning the concept of simple linear transduction chains in favour of interacting networks with molecular modules of physically interacting proteins.