[1] Kyoto Univ, Ctr Ecol Res, Otsu, Shiga 5202113, Japan
[2] Kyoto Univ, Global COE Program Evolut & Biodivers, Grad Sch Sci, Kyoto 6068502, Japan
[3] Univ Turin, Dept Plant Biol, Plant Physiol Unit, I-10135 Turin, Italy
[4] Univ Turin, Innovat Ctr, I-10135 Turin, Italy