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Phosphatidylinositol 3- and 4-Phosphate Are Required for Normal Stomatal Movements

^a Division of Molecular Life Sciences, Pohang University of Science and Technology, Pohang 790-784, Korea
^b Center for Plant Intracellular Trafficking, Pohang University of Science and Technology, Pohang 790-784, Korea
^c Division of Biology, Cell and Developmental Biology Section, and Center for Molecular Genetics, University of California, San Diego, La Jolla, California 92093-0116

² To whom correspondence should be addressed. E-mail ylee{at}postech.ac.kr; fax 82-54-279-2199

Phosphatidylinositol (PI) metabolism plays a central role in signaling pathways in both animals and higher plants. Stomatal guard cells have been reported to contain PI 3-phosphate (PI3P) and PI 4-phosphate (PI4P), the products of PI 3-kinase (PI3K) and PI 4-kinase (PI4K) activities. In this study, we tested the roles of PI3P and PI4P in stomatal movements. Both wortmannin (WM) and LY294002 inhibited PI3K and PI4K activities in guard cells and promoted stomatal opening induced by white light or the circadian clock. WM and LY294002 also inhibited stomatal closing induced by abscisic acid (ABA). Furthermore, overexpression in guard cells of GFP:EBD (green fluorescent protein:endosome binding domain of human EEA1) or GFP:FAPP1PH (PI-four-P adaptor protein-1 pleckstrin homology domain), which bind to PI3P and PI4P, respectively, increased stomatal apertures under darkness and white light and partially inhibited stomatal closing induced by ABA. The reduction in ABA-induced stomatal closing with reduced levels of PI monophosphate seemed to be attributable, at least in part, to impaired Ca²⁺ signaling, because WM and LY294002 inhibited ABA-induced cytosolic Ca²⁺ increases in guard cells. These results suggest that PI3P and PI4P play an important role in the modulation of stomatal closing and that reductions in the levels of functional PI3P and PI4P enhance stomatal opening.

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