The Starch Statolith Hypothesis and the Optimum Angle of Geotropic Stimulation

Abstract

AbstractWhen roots of cress seedlings (Lepidium sativum L.) are stimulated for 10 min at an angle of 135° (i.e. the root tips are pointing obliquely upward), the resulting geotropic curvatures become larger than after 10 min stimulation at 45°. This well‐known behavior has been explained by the better conditions for statoliths, initially located in the floor end of the statocytes, to slide along the cell walls when root tips are pointing upward at 135° than when pointing downward at 45°. Accepting this explanation, one would predict the optimum angle of stimulation to be near 45° when roots had first been kept inverted long enough for their statoliths to accumulate in the opposite end of each functional statocyte. This prediction has been verified in experiments with cress seedlings which were first kept inverted for 16 min, then stimulated for 10 min at given angles, and subsequently rotated parallel to the horizontal axis of the klinostat at 2 rpm. Under these conditions, roots stimulated at 45° curve faster during a 20 to 30 min period on the klinostat than roots stimulated at 135°, but thereafter they stop curving. Roots stimulated at 135°, on the other hand, although initially curving slower than those at 45°, continue curving for at least a whole hour, and attain larger curvatures than the others after 40 min. The optimum shifts from near 45° to near 135° during the course of the klinostat rotation. The behavior of normal and pre‐invertcd roots is interpreted as the result of at least two effects: (1) a stimulation due to the movement of amyloplasts, which is enhanced if these are allowed to slide along the cell walls, and (2) a modification of the development of the resulting curvatures by tonic effects, which are inhibitory between stimulation angles 0° and 90°, and absent or enhancing between 90° and 180°.