[1] Kyoto Univ, Res Div Microbial Sci, Sakyo Ku, Kyoto 6068502, Japan
[2] RIKEN BioResource Ctr, Japan Collect Microorganisms, Ibaraki 3050074, Japan
[3] Meiji Pharmaceut Univ, Dept Microbiol, Tokyo 2048588, Japan
[4] Kyoto Univ, Grad Sch Agr, Div Appl Life Sci, Sakyo Ku, Kyoto 6068502, Japan