[1] Univ Calif San Francisco, Howard Hughes Med Inst, Dept Physiol, San Francisco, CA 94143 USA
[2] Univ Calif San Francisco, Howard Hughes Med Inst, Dept Biochem, San Francisco, CA 94143 USA
[3] Kyoto Univ, Grad Sch Biostudies, Dept Cell & Dev Biol, Kyoto 6068502, Japan
[4] Kyoto Univ, Sch Sci, Dept Biophys, Kyoto 6068502, Japan