Revealing the hierarchy of processes and time-scales that control the tropic response of shoots to gravi-stimulations

Gravity is a major abiotic cue for plant growth. However, little is known about the responses of plants to various patterns of gravi-stimulation, with apparent contradictions being observed between the dose-like responses recorded under transient stimuli in microgravity environments and the responses under steady-state inclinations recorded on earth. Of particular importance is how the gravitropic response of an organ is affected by the temporal dynamics of downstream processes in the signalling pathway, such as statolith motion in statocytes or the redistribution of auxin transporters. Here, we used a combination of experiments on the whole-plant scale and live-cell imaging techniques on wheat coleoptiles in centrifuge devices to investigate both the kinematics of shoot-bending induced by transient inclination, and the motion of the statoliths in response to cell inclination. Unlike previous observations in microgravity, the response of shoots to transient inclinations appears to be independent of the level of gravity, with a response time much longer than the duration of statolith sedimentation. This reveals the existence of a memory process in the gravitropic signalling pathway, independent of statolith dynamics. By combining this memory process with statolith motion, a mathematical model is built that unifies the different laws found in the literature and that predicts the early bending response of shoots to arbitrary gravi-stimulations.