About the Authors

Hidehiko Suzuki

Affiliations Laboratory of Vaccine Materials, National Institute of Biomedical Innovation, Health and Nutrition, Osaka 567–0085, Japan, Laboratory of Bio-Functional Molecular Chemistry, Graduate School of Pharmaceutical Sciences, Osaka University, Suita, Osaka 565–0871, Japan

Akihiro Watari

Affiliation Laboratory of Bio-Functional Molecular Chemistry, Graduate School of Pharmaceutical Sciences, Osaka University, Suita, Osaka 565–0871, Japan

Eri Hashimoto

Affiliations Laboratory of Vaccine Materials, National Institute of Biomedical Innovation, Health and Nutrition, Osaka 567–0085, Japan, Division of Mucosal Immunology, Department of Microbiology and Immunology, The Institute of Medical Sciences, The University of Tokyo, Tokyo 108–8639, Japan

Miki Yonemitsu

Affiliation Laboratory of Vaccine Materials, National Institute of Biomedical Innovation, Health and Nutrition, Osaka 567–0085, Japan

Hiroshi Kiyono

Affiliations Division of Mucosal Immunology, Department of Microbiology and Immunology, The Institute of Medical Sciences, The University of Tokyo, Tokyo 108–8639, Japan, International Research and Development Center for Mucosal Vaccines, The Institute of Medical Science, The University of Tokyo, Tokyo 108–8639, Japan, Core Research for Evolutional Science and Technology (CREST), Japan Science and Technology Agency, Tokyo, Japan

Kiyohito Yagi

Affiliation Laboratory of Bio-Functional Molecular Chemistry, Graduate School of Pharmaceutical Sciences, Osaka University, Suita, Osaka 565–0871, Japan

Masuo Kondoh

kunisawa@nibiohn.go.jp (JK); masuo@phs.osaka-u.ac.jp (MK)

Affiliation Laboratory of Bio-Functional Molecular Chemistry, Graduate School of Pharmaceutical Sciences, Osaka University, Suita, Osaka 565–0871, Japan

Jun Kunisawa

kunisawa@nibiohn.go.jp (JK); masuo@phs.osaka-u.ac.jp (MK)

Affiliations Laboratory of Vaccine Materials, National Institute of Biomedical Innovation, Health and Nutrition, Osaka 567–0085, Japan, Division of Mucosal Immunology, Department of Microbiology and Immunology, The Institute of Medical Sciences, The University of Tokyo, Tokyo 108–8639, Japan, International Research and Development Center for Mucosal Vaccines, The Institute of Medical Science, The University of Tokyo, Tokyo 108–8639, Japan, Core Research for Evolutional Science and Technology (CREST), Japan Science and Technology Agency, Tokyo, Japan, Department of Microbiology and Infectious Diseases, Kobe University Graduate School of Medicine, Kobe 650–0017, Japan, Graduate School of Medicine, Graduate School of Pharmaceutical Sciences, and Graduate School of Dentistry, Osaka University, Osaka 565–0871, Japan

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: HS HK KY MK JK. Performed the experiments: HS AW EH MY. Analyzed the data: HS AW MK JK. Wrote the paper: HS MK JK.