About the Authors

Hiromi Miura

Affiliation Department of Molecular Life Science, Division of Basic Medical Science and Molecular Medicine, School of Medicine, Tokai University, 143 Shimokasuya, Isehara, Kanagawa 259–1193, Japan

Hidetoshi Inoko

Affiliation Department of Molecular Life Science, Division of Basic Medical Science and Molecular Medicine, School of Medicine, Tokai University, 143 Shimokasuya, Isehara, Kanagawa 259–1193, Japan

Masafumi Tanaka

Affiliation Department of Molecular Life Science, Division of Basic Medical Science and Molecular Medicine, School of Medicine, Tokai University, 143 Shimokasuya, Isehara, Kanagawa 259–1193, Japan

Hirofumi Nakaoka

Affiliation Division of Human Genetics, Department of Integrated Genetics, National Institute of Genetics, Yata 1111, Mishima, Shizuoka 411–8540, Japan

Minoru Kimura

Affiliation Department of Molecular Life Science, Division of Basic Medical Science and Molecular Medicine, School of Medicine, Tokai University, 143 Shimokasuya, Isehara, Kanagawa 259–1193, Japan

Channabasavaiah B. Gurumurthy

Affiliation Mouse Genome Engineering Core Facility, Department of Genetics, Cell Biology and Anatomy, University of Nebraska Medical Center, Omaha, Nebraska, 68198, United States of America

Masahiro Sato

Affiliation Section of Gene Expression Regulation, Frontier Science Research Center, Kagoshima University, 1-21-20 Korimoto, Kagoshima, Kagoshima 890–0065, Japan

Masato Ohtsuka

masato@is.icc.u-tokai.ac.jp

Affiliations Department of Molecular Life Science, Division of Basic Medical Science and Molecular Medicine, School of Medicine, Tokai University, 143 Shimokasuya, Isehara, Kanagawa 259–1193, Japan, The Institute of Medical Sciences, Tokai University, 143 Shimokasuya, Isehara, Kanagawa 259–1193, Japan

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: MO HM. Performed the experiments: HM. Analyzed the data: HM HI MT HN MK CBG MO. Wrote the paper: HM MT HN CBG MS MO.