About the Authors

Takeshi Sato

Contributed equally to this work with: Takeshi Sato, Yoshimi Aiyama, Mayuko Ishii-Inagaki

Affiliations Department of Veterinary Anatomy, The University of Tokyo, Tokyo, Japan, Drug Safety Research Labs, Astellas Pharma Inc., Osaka, Japan

Yoshimi Aiyama

Contributed equally to this work with: Takeshi Sato, Yoshimi Aiyama, Mayuko Ishii-Inagaki

Affiliation Department of Veterinary Anatomy, The University of Tokyo, Tokyo, Japan

Mayuko Ishii-Inagaki

Contributed equally to this work with: Takeshi Sato, Yoshimi Aiyama, Mayuko Ishii-Inagaki

Affiliation Department of Veterinary Anatomy, The University of Tokyo, Tokyo, Japan

Kenshiro Hara

Affiliation Division of Germ Cell Biology, National Institute for Basic Biology and Department of Basic Biology, School of Life Science, Graduate University for Advanced Studies (SOKENDAI), Okazaki, Japan

Naoki Tsunekawa

Affiliation Department of Veterinary Anatomy, The University of Tokyo, Tokyo, Japan

Kyoko Harikae

Affiliation Department of Veterinary Anatomy, The University of Tokyo, Tokyo, Japan

Mami Uemura-Kamata

Affiliations Department of Veterinary Anatomy, The University of Tokyo, Tokyo, Japan, Center for Experimental Animal, Tokyo Medical and Dental University, Tokyo, Japan

Mai Shinomura

Affiliation Department of Veterinary Anatomy, The University of Tokyo, Tokyo, Japan

Xiao Bo Zhu

Affiliation Department of Veterinary Anatomy, The University of Tokyo, Tokyo, Japan

Seishi Maeda

Affiliation Division of Cell Biology, Department of Anatomy, Hyogo College of Medicine, Nishinomiya, Japan

Sachi Kuwahara-Otani

Affiliation Division of Cell Biology, Department of Anatomy, Hyogo College of Medicine, Nishinomiya, Japan

Akihiko Kudo

Affiliation Department of Anatomy, Kyorin University School of Medicine, Mitaka, Japan

Hayato Kawakami

Affiliation Department of Anatomy, Kyorin University School of Medicine, Mitaka, Japan

Masami Kanai-Azuma

Affiliation Center for Experimental Animal, Tokyo Medical and Dental University, Tokyo, Japan

Michio Fujiwara

Affiliation Drug Safety Research Labs, Astellas Pharma Inc., Osaka, Japan

Yoichi Miyamae

Affiliation Drug Safety Research Labs, Astellas Pharma Inc., Osaka, Japan

Shosei Yoshida

Affiliation Division of Germ Cell Biology, National Institute for Basic Biology and Department of Basic Biology, School of Life Science, Graduate University for Advanced Studies (SOKENDAI), Okazaki, Japan

Makoto Seki

Affiliation Division of Cell Biology, Department of Anatomy, Hyogo College of Medicine, Nishinomiya, Japan

Masamichi Kurohmaru

Affiliation Department of Veterinary Anatomy, The University of Tokyo, Tokyo, Japan

Yoshiakira Kanai

aykanai@mail.ecc.u-tokyo.ac.jp

Affiliation Department of Veterinary Anatomy, The University of Tokyo, Tokyo, Japan

Competing Interests

TS, MF, and YM are employed by Astellas Pharma Inc. However, this does not alter the authors' adherence to all the PLoS ONE policies on sharing data and materials.

Author Contributions

Conceived and designed the experiments: TS YA MI-I YK. Performed the experiments: TS YA MI-I K. Hara NT K. Harikae MU-K M. Shinomura XBZ SM SK-O AK H. Kawakami MK-A YM. Analyzed the data: TS YA MI-I K. Hara NT K. Harikae MU-K M. Shinomura XBZ M. Seki SM SK-O AK H. Kawakami MK-A YM. Contributed reagents/materials/analysis tools: K. Hara SY. Wrote the paper: SY MK YK. Permitted and indirectly supported Dr. Sato's (TS) work: MF YM.