[1] Hokkaido Univ, Fac Grad Sch Fisheries Sci, 3-1-1 Minato Cho, Hakodate, Hokkaido 0418611, Japan
[2] Hokkaido Univ, Arctic Res Ctr, Kita 21 Nishi 11 Kita Ku, Sapporo, Hokkaido 0010021, Japan
[3] Japan Agcy Marine Earth Sci & Technol, Inst Arctic Climate & Environm, 2-15 Natsushima Cho, Yokosuka, Kanagawa 2370061, Japan
[4] Univ Tokyo, Atmosphere & Ocean Res Inst, 5-1-5 Kashiwanoha, Kashiwa, Chiba 2778564, Japan
[5] Helmholtz Ctr Polar & Marine Res, Polar Biol Oceanog, Alfred Wegener Inst, Handelshafen 12, DE-27570 Bremerhaven, Germany
[6] Marine Ecol Res Inst, Demonstrat Lab, 4-7-17 Arahama, Kashiwazaki, Niigata 9450017, Japan