Microgravity and root gravitropism

Abstract

Space experiments permit to understand better some phases of the gravitropic reaction which occurs when the orientation of the root changes in the gravitational field. In gravisensing cells (statocytes in the root cap), the nucleus is attached to the cell periphery, close to the plasma membrane, by actin filaments. The location of the amyloplasts (statoliths) depends also greatly on these elements of the cytoskeleton. A short period in microgravity (5 min.) modifies the location of the nucleus and of the amyloplasts in the statocytes. The tensions exerted by these very dense organelles on the actin network disappear and this network undergoes a relaxation. The kinetics of gravitropic curvature is also better understood. In fact, gravitropic reaction is regulated by a mechanism depending on gravity. In roots grown in space, then stimulated for 1 h on a 1 g centrifuge, and replaced in microgravity, the regulation limiting the curvature does not occur. It is hypothesized that the sedimentation of the amyloplasts on the endoplasmic reticulum placed at the basal pole of the statocytes could be responsible for this regulation. The contacts between these two organelles should have also a role in root growth. This hypothesis will be tested in our next space experiment (July 94). The experiments in near weightlessness also permit to determine the presentation time which is the duration of stimulation necessary to evoke a slight but significant curvature. Presentation time is 27 s. This short period allows a slight movement of the amyloplasts only (around 0.45 micrometer). The sequence of events leading to the curvature of the root is now well established: the first signal is the separation of the endoplasmic reticulum and the amyloplasts, when the root is subjected to a change in orientation. It is followed by the pressure of these organelles on the actin network which transmits this mechanical effect to the plasma membrane. The transduction of the effect occurs then by the activation of the ions channels (Ca++) and the carrier of a growth inhibitor (auxin), both located in the plasma membrane. This growth inhibitor provokes an asymmetrical growth in the distal part of the meristem and the proximal part of the cell elongation zone. At last, when the root tip reaches the direction of gravity, the amyoloplasts sediment on the endoplasmic reticulum and induce a signal of termination of the curavature.