Gravitropic bending of cress roots without contact between amyloplasts and complexes of endoplasmic reticulum.

The polar arrangement of cell organelles in Lepidium root statocytes is persistently converted to a physical stratification during lateral centrifugation (the centrifugal force acts perpendicular to the root long axis) or by apically directed centrifugation combined with cytochalasin-treatment. Lateral centrifugation (10 min, 60 min at 10g or 50g) causes displacement of amyloplasts to the centrifugal anticlinal cell wall and shifting of the endoplasmic reticulum (ER) complex to the centripetal distal cell edge. After 60 min of lateral centrifugation at 10g or 50g all roots show a clear gravitropic curvature. The average angle of curvature is about 40 degrees and corresponds to that of roots stimulated gravitropically in the horizontal position at 1 g in spite of the fact that the gravistimulus is 10- or 50-fold higher. Apically directed centrifugation combined with cytochalasin B (25 micrograms ml-1) or cytochalasin D (2.5 micrograms ml-1) incubation yields statocytes with the amyloplasts sedimented close to the centrifugal periclinal cell wall and ER cisternae accumulated at the proximal cell pole. Gravitropic stimulation for 30 min in the horizontal position at 1 g and additional 3 h rotation on a clinostat result in gravicurvature of cytochalasin B-treated centrifuged (1 h at 50 g) roots, but because of retarded root growth the angle of curvature is lower than in control roots. Cytochalasin D-treatment during centrifugation (20 min at 50 g) does not affect either root growth or gravicurvature during 3 h horizontal exposure to 1-g relative to untreated roots. As lateral centrifugation enables only short-term contact between the amyloplasts and the distal ER complex at the onset of centrifugation and apically directed centrifugation combined with cytochalasin-treatment even exclude any contact the integrity of the distal cell pole need not necessarily be a prerequisite for graviperception in Lepidium root statocytes.