About the Authors

Asami Nishimori

Affiliation Department of Disease Control, Graduate School of Veterinary Medicine, Hokkaido University, Sapporo, Japan

Satoru Konnai

konnai@vetmed.hokudai.ac.jp

Affiliation Department of Disease Control, Graduate School of Veterinary Medicine, Hokkaido University, Sapporo, Japan

Tomohiro Okagawa

Affiliation Department of Disease Control, Graduate School of Veterinary Medicine, Hokkaido University, Sapporo, Japan

Naoya Maekawa

Affiliation Department of Disease Control, Graduate School of Veterinary Medicine, Hokkaido University, Sapporo, Japan

Ryoyo Ikebuchi

Affiliation Department of Disease Control, Graduate School of Veterinary Medicine, Hokkaido University, Sapporo, Japan

Shinya Goto

Affiliation Department of Disease Control, Graduate School of Veterinary Medicine, Hokkaido University, Sapporo, Japan

Yamato Sajiki

Affiliation Department of Disease Control, Graduate School of Veterinary Medicine, Hokkaido University, Sapporo, Japan

Yasuhiko Suzuki

Affiliations Division of Bioresources, Research Center for Zoonosis Control, Hokkaido University, Sapporo, Japan, Global Station for Zoonosis Control, Global Institution for Collaborative Research and Education (GI-CoRE), Hokkaido University, Sapporo, Japan

Junko Kohara

Affiliation Animal Research Center, Agricultural Research Department, Hokkaido Research Organization, Shintoku, Japan

Satoshi Ogasawara

Affiliation Tohoku University Graduate School of Medicine, Sendai, Japan

Yukinari Kato

Affiliation Tohoku University Graduate School of Medicine, Sendai, Japan

Shiro Murata

Affiliation Department of Disease Control, Graduate School of Veterinary Medicine, Hokkaido University, Sapporo, Japan

Kazuhiko Ohashi

Affiliation Department of Disease Control, Graduate School of Veterinary Medicine, Hokkaido University, Sapporo, Japan

Competing Interests

This research was supported by JSPS KAKENHI, grants from the Science and Technology Research Promotion Program for Agriculture, Forestry, Fisheries and Food Industry, Japan (grant 26058B to S.K.) and the NARO, Bio-oriented Technology Research Advancement Institution (the special scheme project on regional developing strategy: grant 16817557 to S.K.). This work was also supported in part by the Platform for Drug Discovery, Informatics, and Structural Life Science (PDIS) from Japan Agency for Medical Research and development, AMED (to Y.K.). The authors, Nishimori, Konnai, Okagawa, Maekawa, Suzuki, Murata and Ohashi, have a pending patent derived from this work (Tokugan 2016-159089). This does not alter our adherence to all PLOS ONE policies on sharing data and materials.

Author Contributions

Conceptualization: AN SK KO. Funding acquisition: SK YK. Investigation: AN TO NM SG Y. Sajiki SO RI. Project administration: SK SM KO. Resources: Y. Suzuki JK SO YK. Supervision: SK SM KO. Writing – original draft: AN SK. Writing – review & editing: AN SK YK SM KO.