The close relationship between gravitropism and calcium ions has been around for over 10 years. There is evidence that wall calcium ion movement can regulate growth expansion and that calcium ions in free cells can mediate signaling in statocytes.
However, changes in wall calcium ion or intracellular calcium concentrations are unlikely to occur independently of each other. This creates ambiguity in the interpretation of experimental information and experiment.
Also, there is no certainty that the observed redistribution of calcium ions in walls is sufficient to cause bending, and there is currently no direct evidence from images or measurements were changes in [{Ca^{2+}}Ca
2+
] initiate or transform gravitational signals.
Relating gravity and gravitropism evidence is important, but impractical nonetheless. The free intracellular calcium ion involvement is tested best with a novel transgenic aequorin method targeting columella cells to address this critical issue.
New information is provided that links the control or orientation of growing cells, such as pollen tubes, to the manipulation of cytoplasmic calcium ions. This information can be applied directly to the fit method, a gravity modeling system that has been studied over the years.
Role of Calcium in Root Gravity Perception:
Calcium ions may have a crucial function in linking asymmetric growth with root cap gravity perception that occurs in the elongated root areas. The calcium chelator application to the root canopy without affecting growth suppresses gravitational curvature to one side of the root cap asymmetric application of calcium distorts the gravity stimulation and calcium source induces a polar movement of calcium ions applied to the base through the root cover.
The effect of calcium may be associated with the migration of auxin in the roots, since 1) inhibitors of auxin transport interfere with the polar transport of calcium and gravitational curvature under the influence of gravity, and 2) the introduction of asymmetric calcium increases the transport of auxin across elongation zone.
Indirect data indicate that the calcium-regulating regulatory protein calmodulin may be involved in the action of calcium or transport in the mechanisms of the root gravitropism.
Extracellular Free-Calcium Gradients Function in Gravitropic Signaling in Roots of Maize:
Since the applied radioactive calcium is polarized by gravity and the asymmetrically applied calcium causes root deformation, an extracellular calcium gradient in the root canopy has been proposed as a mediator for gravity transfer in the root. For these gradients to be a signal, one needs to detect the difference in calcium reactivity along the gradient.
The very high cation-binding pH-dependent capacity of the wall means that there can be only a small ratio between the calcium activity and calcium. In research, direct measurements of calcium activity show similar results when imaging curvature of this activity gradient, which begins to develop with a delay. If the physiological signal is extracellular calcium, a low apoplast diffusion constant means that extracellular calcium has a more localized effect on cell function than diffusion in the growth zone.
The proposed physiological event refers only to the response of state cells to gravitational stimulation, but not to growth. The root apoplastic initial calcium activity was higher compared to the nutrient medium, which indicates that calcium seeped out of these young seedlings and was not absorbed by the nutrient medium. There are no barriers, and the difference in activity directly determines the driving force behind the spread.
Since the elimination of this gradient by mobilizing calcium ions removes curvature, a gradient of calcium activity towards the root apex is required for gravity. Calcium and Dinitro-BAPTA bind with each other and the complex diffuses very strongly, but the binding constant is such that the calcium activity is constant throughout the apoplast and only slightly lower than that found elsewhere.
This may be because it is attached to the wall. The diffusion of these ions slows down due to a large number of binding sites. The mechanism that determines the slope of the apoplastic activity of calcium is unknown. One possibility relates to the difference in the pH of the non-plastic materials on both sides. Since protons can displace calcium bound in the wall and increase calcium activity, increased transport of protons from the base of the root to the top through the cell can lead to a calcium gradient.
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