A universal role for inositol 1,4,5-trisphosphate-mediated signaling in plant gravitropism

Inositol 1,4,5- trisphosphate (InsP(3)) has been implicated in the early signaling events of plants linking gravity sensing to the initiation of the gravitropic response. However, at present, the contribution of the phosphoinositide signaling pathway in plant gravitropism is not well understood. To delineate the role of InsP(3) in plant gravitropism, we generated Arabidopsis (Arabidopsis thaliana) plants constitutively expressing the human type I inositol polyphosphate 5- phosphatase (InsP 5-ptase), an enzyme that specifically hydrolyzes InsP(3). The transgenic plants show no significant differences in growth and life cycle compared to wild-type plants, although basal InsP(3) levels are reduced by greater than 90% compared to wild-type plants. With gravistimulation, InsP(3) levels in inflorescence stems of transgenic plants show no detectable change, whereas in wild-type plant inflorescences, InsP(3) levels increase approximately 3-fold within the first 5 to 15 min of gravistimulation, preceding visible bending. Furthermore, gravitropic bending of the roots, hypocotyls, and inflorescence stems of the InsP 5-ptase transgenic plants is reduced by approximately 30% compared with the wild type. Additionally, the cold memory response of the transgenic plants is attenuated, indicating that InsP(3) contributes to gravisignaling in the cold. The transgenic roots were shown to have altered calcium sensitivity in controlling gravitropic response, a reduction in basipetal indole-3-acetic acid transport, and a delay in the asymmetric auxin-induced beta-glucuronidase expression with gravistimulation as compared to the controls. The compromised gravitropic response in all the major axes of growth in the transgenic Arabidopsis plants reveals a universal role for InsP(3) in the gravity signal transduction cascade of plants.