Hypergravity can reduce but not enhance the gravitropic response of Chara globularis protonemata

The relationship between the position of the statoliths and the direction and rate of tip growth in negatively gravitropic protonemata of Chara globularis was studied with a centrifuge video microscope. Cells placed perpendicularly to the acceleration vector (stimulation angle 90 degrees) showed a gradual reduction of the gravitropic curvature with increasing accelerations from 1 g to 8 g despite complete sedimentation of all statoliths on the centrifugal cell flank. It is argued that the increased weight of the statoliths in hypergravity impairs their acropetal transport which is induced when the cell axis deviates from the normal upright orientation. When the statoliths were centrifuged deep into the apical dome at 6 g and a stimulation angle of 170 degrees the gravitropic curvature after 1 h was identical to that determined for the same cells at 1 g and the same stimulation angle. This indicates that gravitropism in Chara protonemata is either independent of the pressure exerted by the statoliths on an underlying structure or is already saturated at 1 g. When the statoliths were moved along the apical cell wall at 8 g and the stimulation angle was gradually increased from 170 degrees to 220 degrees the gravitropic curvature reverted sharply when the cluster of statoliths passed over the cell pole. This experiment supports the hypothesis that in Chara protonemata asymmetrically distributed statoliths inside the apical dome displace the Spitzenkorper and thus the centre of growth, resulting in gravitropic bending. In contrast to the positively gravitropic Chara rhizoids, no modifications either in the transport of statoliths during basipetal acceleration (6 g, stimulation angle 0 degree, 5 h) or in the subsequent gravitropic response could be detected in the protonemata. The different effects of centrifugation on the positioning of statoliths in Chara protonemata and rhizoids indicate subtle differences in the function of the cytoskeleton in both types of cells.