1Center for Experimental Medicine, 2Department of Microbiology and Immunity, 3 Frontier Research Initiative, The Institute of Medical Science, The University of Tokyo, 4-6-1 Shirokanedai, Minato-ku, Tokyo, 108-8639, Japan. 4Core Research for Evolutional Science and Technology (CREST), Japan Science and Technology Agency, Saitama 332-0012, Japan. 5Present address: Animal Research Center, Tokyo Medical University, 6-1-1 Shinjyuku, Shinjyuku-ku, Tokyo 160-8402, Japan.

1. Department of Biological Information, Graduate School of Bioscience and Biotechnology, Tokyo Institute of Technology, B34 4259 Nagatsuta-cho, Midori-ku, Yokohama-shi, Kanagawa 226-8501, Japan 2. Department of Molecular Biophysics and Biochemistry, 3. Department of Chemistry, Yale University, New Haven, Connecticut 06520-8114, USA 4. Department of Basic Medical Sciences, Institute of Medical Science, The University of Tokyo, 4-6-1 Shirokanedai, Minato-ku, Tokyo 108-8639, Japan 5. These authors contributed equally to this work.

1. Department of Biological Information, Graduate School of Bioscience and Biotechnology, Tokyo Institute of Technology, 4259 Nagatsuta-cho, Midori-ku, Yokohama-shi, Kanagawa 226-8501, Japan 2. Institute for Virus Research, Kyoto University, Kyoto 606-8507, Japan 3. Division of Structural Biology, Department of Basic Medical Science, The Institute of Medical Science, The University of Tokyo, 4-6-1 Shirokanedai, Minato-ku, Tokyo 108-8639, Japan 4. Advanced Science Institute, 5. Biomolecular Characterization Team and CREST/JST, RIKEN, 2-1 Hirosawa, Wako, Saitama 351-0198, Japan 6. Department of Biochemistry and Molecular Genetics, University of Alabama at Birmingham, Schools of Medicine and Dentistry, 402B Kaul Genetics Building, 720 20th Street South, Birmingham, Alabama 35294, USA 7. These authors contributed equally to this work. 8. Present address: Structural Biology Laboratory, Life Science Division, Synchrotron Radiation Research Organization, The University of Tokyo, 211 General Research Building, Institute of Molecular and Cellular Biosciences, 1-1-1 Yayoi, Bunkyo-ku, Tokyo 113-0032, Japan.

Division of Cellular and Molecular biology, Institute of Medical Science, University of Tokyo, 4-6-1 Shirokane-dai, Minato-ku, Tokyo 108-8639, Japan.

1. Institute for Molecular Virology, 2. Howard Hughes Medical Institute, 3. Department of Pathobiological Sciences, and, 4. Departments of Statistics and of Biostatistics and Medical Informatics, University of Wisconsin-Madison, Madison, Wisconsin 53706, USA 5. Faculty of Veterinary Medicine, and, 6. Collaborating Research Center-Emerging & Reemerging Infectious Diseases, Tropical Disease Centre, Airlangga University, Surabaya 60115, Indonesia 7. Division of Virology, Department of Microbiology and Immunology, and, 8. International Research Center for Infectious Diseases, Institute of Medical Science, University of Tokyo, Tokyo 108-8639, Japan 9. Division of Zoonosis, Department of Microbiology and Infectious Diseases, Graduate School of Medicine, Kobe University, Kobe 650-0017, Japan 10. These authors contributed equally to this work. 11. Present address: First Department of Forensic Science, National Research Institute of Police Science, 6-3-1 Kashiwanoha, Kashiwa 277-0882, Japan.

1 Division of Oncology, Department of Cancer Biology, The University of Tokyo, Minato-ku, Tokyo, 108-8639, Japan
2 Division of Cell Biology, Center for Experimental Medicine, The University of Tokyo, Minato-ku, Tokyo, 108-8639, Japan
3 Laboratory of Proteomics, Institute of Medical Science, The University of Tokyo, Minato-ku, Tokyo, 108-8639, Japan
4 Present address: Laboratory for Pluripotent Cell Studies, RIKEN Center for Developmental Biology, Kobe, Hyogo, 650-0047, Japan.
5 Present address: National Eye Institute, NIH, Bethesda, MD, 20892, USA.
6 Present address: Animal Research Center, Tokyo Medical University, Shinjuku-ku, Tokyo, 160-8402, Japan.

1. Center for Experimental Medicine, The Institute of Medical Science, The University of Tokyo, 4-6-1 Shirokanedai, Minato-ku, Tokyo 108-8639, Japan.
2. Genodive Pharma, 411 Ikehata, Isehara-shi, Kanagawa 259-1144, Japan.
3. Department of Pathology, Ehime University Graduate School of Medicine, Shitsukawa, Toon, Ehime 791-0295, Japan.
4. Present addresses: Genodive Pharma Inc., 411 Ikehata, Isehara-shi, Kanagawa 259-1144, Japan (H.K.); Animal Research Center, Tokyo Medical University, 6-1-1, Shinjuku, Shinjuku-ku, Tokyo 160-8402, Japan (K.S.).

Division of Cell Biology, Center for Experimental Medicine, and Fine Morphology Laboratory, Department of Basic Medical Science, Institute of Medical Science, University of Tokyo, 4-6-1, Shirokanedai, Minato-ku, Tokyo 108-8639, Japan.

1: Institute of Medical Science, University of Tokyo, 4-6-1 Shirokanedai, Minato-ku, Tokyo, 108-8639, Japan
2: Atopy (Allergy) Research Center, Juntendo University School of Medicine, 2-1-1 Hongo, Bunkyo-ku, Tokyo, 113-8421, Japan
3: Finsen Laboratory, Rigshospitalet, Strandboulevarden 49, 2100 Copenhagen, Denmark
4: Department of Anatomy, University of California, San Francisco, 513 Parnassus Avenue, San Francisco, CA 94143-0452, USA

1: Division of Infectious Genetics, The Institute of Medical Science, The University of Tokyo
2: Division of Immunology, The Institute of Medical Science, The University of Tokyo
4: Department of Immunobiology and Pharmacological Genetics, Graduate School of Medicine and Pharmaceutical Sciences for Research, University of Toyama
3: These authors contributed equally to this study.