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Univ Grenoble 1, Lab Physiol Cellulaire & Vegetate, Inst Rech Sci & Technol Vivant, CNRS,CEA,INRA,CEA Grenoble,iRTSV LPCV,UMR 5168, F-38054 Grenoble, France Univ Grenoble 1, Lab Physiol Cellulaire & Vegetate, Inst Rech Sci & Technol Vivant, CNRS,CEA,INRA,CEA Grenoble,iRTSV LPCV,UMR 5168, F-38054 Grenoble, France