CAS Key Laboratory of Pathogenic Microbiology and Immunology CASPMI,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China ,University of Chinese Academy of Sciences,Beijing,PR China ,China-Japan Joint Laboratory of Molecular Immunology and Microbiology,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China,CAS Key Laboratory of Pathogenic Microbiology and Immunology CASPMI,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China ,University of Chinese Academy of Sciences,Beijing,PR China ,China-Japan Joint Laboratory of Molecular Immunology and Microbiology,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China,CAS Key Laboratory of Pathogenic Microbiology and Immunology CASPMI,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China ,University of Chinese Academy of Sciences,Beijing,PR China ,China-Japan Joint Laboratory of Molecular Immunology and Microbiology,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China,CAS Key Laboratory of Pathogenic Microbiology and Immunology CASPMI,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China ,University of Chinese Academy of Sciences,Beijing,PR China ,China-Japan Joint Laboratory of Molecular Immunology and Microbiology,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China,CAS Key Laboratory of Pathogenic Microbiology and Immunology CASPMI,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China ,University of Chinese Academy of Sciences,Beijing,PR China ,China-Japan Joint Laboratory of Molecular Immunology and Microbiology,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China,CAS Key Laboratory of Pathogenic Microbiology and Immunology CASPMI,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China ,University of Chinese Academy of Sciences,Beijing,PR China ,China-Japan Joint Laboratory of Molecular Immunology and Microbiology,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China,CAS Key Laboratory of Pathogenic Microbiology and Immunology CASPMI,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China ,University of Chinese Academy of Sciences,Beijing,PR China ,China-Japan Joint Laboratory of Molecular Immunology and Microbiology,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China,CAS Key Laboratory of Pathogenic Microbiology and Immunology CASPMI,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China ,University of Chinese Academy of Sciences,Beijing,PR China ,China-Japan Joint Laboratory of Molecular Immunology and Microbiology,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China,CAS Key Laboratory of Pathogenic Microbiology and Immunology CASPMI,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China ,University of Chinese Academy of Sciences,Beijing,PR China ,China-Japan Joint Laboratory of Molecular Immunology and Microbiology,Institute of Microbiology,Chinese Academy of Sciences,Beijing,PR China

This work was supported by grants from National Basic Research Program of China (2013CB531502), National Science and Technology Major Project of China (2012ZX1004701-001-002) and the National Natural Science Foundation of China (31370889, 31170829)