[1] NARO, Hokkaido Agr Res Ctr, Sapporo, Hokkaido 0628555, Japan
[2] NARO, Inst Livestock & Grassland Sci, Tsukuba, Ibaraki 3050901, Japan
[3] Natl Livestock Breeding Ctr, Fukushima 9618511, Japan
[4] Livestock Improvement Assoc Japan, Tokyo 1350041, Japan
[5] Obihiro Univ Agr & Vet Med, Obihiro, Hokkaido 0808555, Japan