Concerns regarding the reliability of slow-and fast-rotating uni-axial clinostats in simulating weightlessness have induced the construction of devices considered to simulate weightlessness more adequately. A new three-dimensional (3-D) clinostat equipped with two rotation axes placed at right angles has been constructed. In the clinostat, the rotation achieved with two motors is computer-controlled and monitored with encoders attached to the motors. By rotating plants three-dimensionally at random rates on the clinostat, their dynamic stimulation by gravity in every direction can be eliminated. Some of the vegetative growth phases of plants dependent on the gravity vector, such as morphogenesis, are shown to be influenced by rotation on the 3-D clinostat. The validity of 3-D clinostatting has been evaluated by comparing structural parameters of cress roots and Chara rhizoids obtained under real microgravity with those obtained after 3-D clinostatting. The parameters analyzed up to now (organization of the root cap, integrity and polarity of statocytes, dislocation of statoliths, amount of starch and ER) demonstrate that the 3-D clinostat is a valuable device for simulating weightlessness.

During five rocket flights (TEXUS 18, 19, 21, 23 and 25), experiments were performed to investigate the behaviour of statoliths in rhizoids of the green alga Charo globularia Thuill. and in statocytes of cress (Lepidium sativum L.) roots, when the gravitational field changed to approx. l0(-4) g (i.e. microgravity) during the parabolic flight (lasting for 301-390 s) of the rockets. The position of statoliths was only slightly influenced by the conditions during launch, e.g. vibration, acceleration and rotation of the rocket. Within approx. 6 min of microgravity conditions the shape of the statolith complex in the rhizoids changed from a transversely oriented lens into a longitudinally oriented spindle. The center of the statolith complex moved approx. 14 micrometers and 3.6 micrometers in rhizoids and root statocytes, respectively, in the opposite direction to the originally acting gravity vector. The kinetics of statolith displacement in rhizoids demonstrate that the velocity was nearly constant under microgravity whereas it decreased remarkably after inversion of rhizoids on Earth. It can be concluded that on Earth the position of statoliths in both rhizoids and root statocytes depends on the balance of two forces, i.e. the gravitational force and the counteracting force mediated by microfilaments.

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Horizontally positioned Chara rhizoids continue growth without gravitropic bending when the statoliths are removed from the apex by basipetal centrifugation. The transport of statoliths in centrifuged rhizoids is bidirectional: 50-60 % ofthe statoliths are retransported on a straight course to the apex at velocities from 1 to 14 pm -min-', increasing towards the rhizoid tip. The centrifuged statoliths which are located closest to the nucleus are basipetally transported and caught up in the cytoplasmic streaming of the cell. Those statoliths which are located near the apical side of the nucleus are transported either apically or basally. A de-novo-formation of statoliths was not observed. After retransport to the apex some statoliths transiently sediment, a process which can induce a local inhibition of cell wall growth. The rhizoid bends again gravitropically only if a few statoliths finally sediment in the apex; the more statoliths that sediment in the apex the shorter the radius of bending becomes. The transport of statoliths is mediated by actin filaments which form a network ofthin filaments in the apical and subapical zone of the rhizoid, and thicker parallel bundles in the basal zone where cytoplasmic streaming occurs. Both subpopulations of actin filaments overlap in the nucleus zone.