[1] Univ Tokyo, Grad Sch Sci, Dept Biol Sci, Bunkyo Ku, Tokyo 1130033, Japan
[2] Kyoto Univ, Grad Sch Biostudies, Sakyo Ku, Kyoto 6068501, Japan
[3] Nara Inst Sci & Technol, Plant Global Educ Project, Grad Sch Biol Sci, Nara 6300192, Japan
[4] CSIC, Ctr Nacl Biotecnol, Dept Genet Mol Plantas, E-28049 Madrid, Spain