[1] Kitasato Univ, Sch Fisheries Sci, Sanri Ku, Ofunato, Iwate 0220101, Japan
[2] Univ Tokyo, Grad Sch Agr & Life Sci, Dept Aquat Biosci, Bunkyo Ku, Tokyo 1138657, Japan
[3] Natl Inst Basic Biol, Dept Dev Biol, Reprod Biol Lab, Okazaki 4448585, Japan
[4] Iwate Fisheries Technol Ctr, Kamaishi, Iwate 0260001, Japan