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Plastids: dynamic components of plant cell development

The gravitropic bending of maize roots, as a response to reorientation of the root within a gravitational field, was examined for sensitivity to exogenous applications of the cytoskeletal inhibitor, cytochalasin D. Agar blocks were impregnated with this inhibitor, and were applied either to the root cap or to the zone of root cell elongation. Root growth was normal with either treatment, if the roots were not repositioned with respect to the gravitational vector. When untreated roots were placed in a horizontal position with respect to gravity, a 40 degree bending response was observed within one hour. This bending also occurred when cytochalasin D was applied at high concentrations to the zone of root cell elongation. However, when cytochalasin D above 40 micrograms/ml was applied to the root cap, roots lost the ability of directional reorientation within the gravitational field, causing a random bending.

NASA Discipline Cell Biology↗

Effects of clinorotation and microgravity on sweet clover columella cells treated with cytochalasin D

The cytoskeleton of columella cells is believed to be involved in maintaining the developmental polarity of cells observed as a reproducible positioning of cellular organelles. It is also implicated in the transduction of gravitropic signals. Roots of sweet clover (Melilotus alba L.) seedlings were treated with a microfilament disrupter, cytochalasin D, on a slowly rotating horizontal clinostat (2 rpm). Electron micrographs of treated columella cells revealed several ultrastructural effects including repositioning of the nucleus and the amyloplasts and the formation of endoplasmic reticulum (ER) whorls. However, experiments performed during fast clinorotation (55 rpm) showed an accumulation (but no whorling) of a disorganized ER network at the proximal and distal pole and a random distribution of the amyloplasts. Therefore, formation of whorls depends upon the speed of clinorotation, and the overall impact of cytochalasin D suggests the necessity of microfilaments in organelle positioning. Interestingly, a similar drug treatment performed in microgravity aboard the US Space Shuttle Endeavour (STS-54, January 1993) caused a displacement of ER membranes and amyloplasts away from the distal plasma membrane. In the present study, we discuss the role of microfilaments in maintaining columella cell polarity and the utility of clinostats to simulate microgravity.

manned↗

The effect of the external medium on the gravitropic curvature of rice (Oryza sativa, Poaceae) roots

The roots of rice seedlings, growing in artificial pond water, exhibit robust gravitropic curvature when placed perpendicular to the vector of gravity. To determine whether the statolith theory (in which intracellular sedimenting particles are responsible for gravity sensing) or the gravitational pressure theory (in which the entire protoplast acts as the gravity sensor) best accounts for gravity sensing in rice roots, we changed the physical properties of the external medium with impermeant solutes and examined the effect on gravitropism. As the density of the external medium is increased, the rate of gravitropic curvature decreases. The decrease in the rate of gravicurvature cannot be attributed to an inhibition of growth, since rice roots grown in 100 Osm/m3 (0.248 MPa) solutions of different densities all support the same root growth rate but inhibit gravicurvature increasingly with increasing density. By contrast, the sedimentation rate of amyloplasts in the columella cells is unaffected by the external density. These results are consistent with the gravitational pressure theory of gravity sensing, but cannot be explained by the statolith theory.

Non-NASA Center↗

Tip-growing cells of the moss Ceratodon purpureus Are gravitropic in high-density media

Gravity sensing in plants and algae is hypothesized to rely upon either the mass of the entire cell or that of sedimenting organelles (statoliths). Protonemata of the moss Ceratodon purpureus show upward gravitropism and contain amyloplasts that sediment. If moss sensing were whole-cell based, then media denser than the cell should prevent gravitropism or reverse its direction. Cells that were inverted or reoriented to the horizontal displayed distinct negative gravitropism in solutions of iodixanol with densities of 1.052 to 1.320 as well as in bovine serum albumin solutions with densities of 1.037 to 1.184 g cm(-3). Studies using tagged molecules of different sizes and calculations of diffusion times suggest that both types of media penetrate through the apical cell wall. Estimates of the density of the apical cell range from 1.004 to 1.085. Because protonemata grow upward when the cells have a density that is lower than the surrounding medium, gravitropic sensing probably utilizes an intracellular mass in moss protonemata. These data provide additional support for the idea that sedimenting amyloplasts function as statoliths in gravitropism.

NASA Discipline Plant Biology↗