[1] Kyoto Univ, Grad Sch Biostudies, Dept Cell & Dev Biol, Kyoto 6068502, Japan
[2] Kyoto Univ, Fac Sci, Dept Biophys, Kyoto 6068502, Japan
[3] Japan Sci & Technol Corp, PRESTO, Osaka 5650082, Japan
[4] Stanford Univ, Dept Mol Cellular Physiol, Sch Med, Stanford, CA 94305 USA