Force sensitivity of plant gravisensing.

Rotation at 4, 10, 50 and 100 rpm on a horizontal clinostat and in microgravity exerts limited effects on the morphogenesis of lettuce and cress root statocytes and statoliths if compared with the vertical control or 1 g spaceflight reference centrifuge. However, the average distance of statoliths from the distal wall increases. The pattern of plastid location of microgravity-grown and that of clino-rotated samples has been determined at 10, 50, and 100 rpm. Experiments on the centrifuge-clinostat and spaceflight centrifuge (acceleration forces of 0.005 to 1 g) revealed that the average statolith location depends on the amplitude of acropetally or basipetally directed mass acceleration. Decreasing the acropetally directed force from 1 g to 0.4 g dislocates statoliths towards the cell center possibly mediated by the elastic forces of the cytoskeleton. In statocytes formed on the clinostat or in microgravity, the majority of statoliths are located at the center of the cell. To force the statoliths from the center of the statocyte towards one of its poles, a threshold mass acceleration of 0.01 g is required. Statocytes with centrally-located statoliths are considerably more effective in transducing a gravistimulus than those with distally-located plastids. The latent time of the graviresponse is shorter and the response itself is enhanced in roots grown on the clinostat compared to vertically grown samples. The early phases of graviperception are independent of root growth conditions since presentation time and g-threshold are similar for roots grown stationary and those on a clinostat. We propose a sequence of events in gravitropic stimulation that considers not only the lateral displacement of statoliths, as predicted by the starch-statolith hypothesis, but also its longitudinal motion, together with differential gravisensitivity of mechanotransducing structures along the lower-most longitudinal cell wall.