Reorientation of Seedlings in the Earth's Gravitational Field Induces Cytosolic Calcium Transients

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Abstract
The gravitational field controls plant growth, morphology, and development. However, the underlying transduction mechanisms are not well understood. Much indirect evidence has implicated the cytoplasmic free calcium concentration ([Ca2+]c) as an important factor, but direct evidence for changes in [Ca2+]c is currently lacking. We now have made measurements of [Ca2+]c in groups of young seedlings of Arabidopsis expressing aequorin in the cytoplasm and reconstituted in vivo with cp-coelenterazine, a synthetic high-affinity luminophore. Distinct [Ca2+]c signaling occurs in response to gravistimulation with kinetics very different from [Ca2+]c transients evoked by other mechanical stimuli (e.g. movement and wind). [Ca2+]cchanges produced in response to gravistimulation are transient but with a duration of many minutes and dependent on stimulus strength (i.e. the angle of displacement). The auxin transport blockers 2,3,5-tri-iodo benzoic acid and N-(1-naphthyl) phthalamic acid interfere with gravi-induced [Ca2+]cresponses and addition of methyl indole-3-acetic acid to whole seedlings induces long-lived [Ca2+]ctransients, suggesting that changes in auxin transport may interact with [Ca2+]c. Permanent nonaxial rotation of seedlings on a two-dimensional clinostat, however, produced a sustained elevation of the [Ca2+]c level. This probably reflects permanent displacement of gravity-sensing cellular components and/or disturbance of cytoskeletal tension. It is concluded that [Ca2+]c is part of the gravity transduction mechanism in young Arabidopsis seedlings.