Murase Kazunori 村瀬一典

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Kazunori Murase 村瀬一典

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Researcher Number	40710869
Other IDs
Affiliation (Current)	2025: 京都大学, 医学研究科, 助教
Affiliation (based on the past Project Information) *help	2019 – 2024: 京都大学, 医学研究科, 助教 2016 – 2019: 京都大学, 医学研究科, 研究員 2017 – 2018: 宮崎大学, 医学部, 助教 2015: 京都大学, 医学(系)研究科(研究院), 特定助教 2014: 宮崎大学, フロンティア科学実験総合センター, 研究員
Review Section/Research Field	Principal Investigator Basic Section 49050:Bacteriology-related / Bacteriology (including mycology) Except Principal Investigator Basic Section 49050:Bacteriology-related / Hygiene and public health / Morphological basic dentistry
Keywords	Principal Investigator 細胞外小胞 / 外膜小胞 / 病原性メカニズム / 病原機構 / 免疫応答 / 細胞傷害性 / A群レンサ球菌 / ワクチン / エンドサイトーシス / 宿主細胞の免疫応答 … More / 病原性細菌 / 国際情報交換（フランス） / グラム陰性細菌 / HlyF / 国際共同研究 … More Except Principal Investigator A群レンサ球菌 / ゼノファジー / ゲノム / 比較ゲノム / TBK1 / ガレクチン３ / GBP1 / 認識 / 糖鎖認識 / 自然免疫 / VPS4 / SLO / Rab41 / ESCRT / Ａ群レンサ球菌 / ユビキチン / 細菌感染 / 生態 / 水圏 / ビブリオ / フィールド / 海外 / コレラ菌 / 魚病 / CRISPR / ゲノム疫学 / 薬剤耐性 / 比較ゲノム解析 / コレラ / 微生物 / 細菌 / 病原性 / トランスクリプトーム / メチローム / ゲノム解析 / GWAS / エピゲノム / 溶連菌 / 多様化 / 細菌ゲノム / 多様性 / 進化 Less

多様な細菌感染に対するロバストネスを支えるゼノファジー機構の解明
- Principal Investigator
  
  野澤孝志
- Project Period (FY)
  2024 – 2026
- Research Category
  
  Grant-in-Aid for Scientific Research (B)
- Review Section
  
  Basic Section 49050:Bacteriology-related
- Research Institution
  Kyoto University
Elucidation of the mechanism of competition between different bacterial species via extracellular vesicles and its application to the development of new drug discovery technologiesPrincipal Investigator
- Principal Investigator
  
  村瀬一典
- Project Period (FY)
  2023 – 2025
- Research Category
  
  Grant-in-Aid for Scientific Research (C)
- Review Section
  
  Basic Section 49050:Bacteriology-related
- Research Institution
  Kyoto University
Elucidation of Xenophagy Regulation by Bacterial Glycan Recognition
- Principal Investigator
  
  中川一路
- Project Period (FY)
  2022 – 2024
- Research Category
  
  Grant-in-Aid for Scientific Research (B)
- Review Section
  
  Basic Section 49050:Bacteriology-related
- Research Institution
  Kyoto University
Involvement in the pathogenic mechanism of Streptococcus pyogenes-derived extracellular vesiclesPrincipal Investigator
- Principal Investigator
  
  Murase Kazunori
- Project Period (FY)
  2020 – 2022
- Research Category
  
  Grant-in-Aid for Scientific Research (C)
- Review Section
  
  Basic Section 49050:Bacteriology-related
- Research Institution
  Kyoto University
Analysis of membrane vesicle production in gram negative bacteriaPrincipal Investigator
- Principal Investigator
  
  Murase Kazunori
- Project Period (FY)
  2017 – 2019
- Research Category
  
  Grant-in-Aid for Scientific Research (C)
- Research Field
  
  Bacteriology (including mycology)
- Research Institution
  Kyoto University
  University of Miyazaki
Elucidation of genomic diversification in Group A Streptococci based on GWAS and epigenome
- Principal Investigator
  
  MARUYAMA FUMITO
- Project Period (FY)
  2016 – 2018
- Research Category
  
  Grant-in-Aid for Scientific Research (B)
- Research Field
  
  Morphological basic dentistry
- Research Institution
  Kyoto University
Validation of microbial safety in drinking water in the world by comparative genomics of Vibrio cholerae
- Principal Investigator
  
  MARUYAMA Fumito
- Project Period (FY)
  2016 – 2019
- Research Category
  
  Grant-in-Aid for Scientific Research (B)
- Research Field
  
  Hygiene and public health
- Research Institution
  Hiroshima University
  Kyoto University
Elucidation of the mechanism of outer membrane vesicles production in Gram-negative bacteriaPrincipal Investigator
- Principal Investigator
  
  Kazunori Murase
- Project Period (FY)
  2014 – 2016
- Research Category
  
  Grant-in-Aid for Young Scientists (B)
- Research Field
  
  Bacteriology (including mycology)
- Research Institution
  Kyoto University
  University of Miyazaki

All 2023 2022 2021 2020 2019 2016 2015 Other

All Journal Article Presentation

[Journal Article] Group A Streptococcus cation diffusion facilitator proteins contribute to immune evasion by regulating intracellular metal concentrations2023
- Author(s)
  Aikawa Chihiro、Shimizu Akihide、Nakakido Makoto、Murase Kazunori、Nozawa Takashi、Tsumoto Kouhei、Nakagawa Ichiro
- Journal Title
  
  Biochemical and Biophysical Research Communications
  
  Volume: 676 Pages: 141-148
- DOI
  10.1016/j.bbrc.2023.07.052
- Peer Reviewed / Open Access
- Data Source
  KAKENHI-PROJECT-23K18214, KAKENHI-PROJECT-22KK0096, KAKENHI-PROJECT-22K07053, KAKENHI-PROJECT-21K07023, KAKENHI-PLANNED-19H05766, KAKENHI-PUBLICLY-22H04809, KAKENHI-PROJECT-23K24130
[Journal Article] Rab41-mediated ESCRT machinery repairs membrane rupture by a bacterial toxin in xenophagy2023
- Author(s)
  Nozawa Takashi、Toh Hirotaka、Iibushi Junpei、Kogai Kohei、Minowa-Nozawa Atsuko、Satoh Junko、Ito Shinji、Murase Kazunori、Nakagawa Ichiro
- Journal Title
  
  Nature Communications
  
  Volume: 14 Issue: 1 Pages: 6230-6230
- DOI
  10.1038/s41467-023-42039-2
- Peer Reviewed / Open Access
- Data Source
  KAKENHI-PROJECT-23K18214, KAKENHI-PROJECT-23KJ2178, KAKENHI-PROJECT-22KK0096, KAKENHI-PROJECT-23K06528, KAKENHI-PROJECT-23K06544, KAKENHI-PROJECT-21K07023, KAKENHI-PUBLICLY-22H04809, KAKENHI-PROJECT-23K24130
[Journal Article] Clonal Expansion of Multidrug-Resistant <i>Streptococcus dysgalactiae</i> Subspecies <i>equisimilis</i> Causing Bacteremia, Japan, 2005?20212023
- Author(s)
  Shinohara Koh、Murase Kazunori、Tsuchido Yasuhiro、Noguchi Taro、Yukawa Satomi、Yamamoto Masaki、Matsumura Yasufumi、Nakagawa Ichiro、Nagao Miki
- Journal Title
  
  Emerging Infectious Diseases
  
  Volume: 29 Issue: 3 Pages: 528-539
- DOI
  10.3201/eid2903.221060
- Peer Reviewed / Open Access
- Data Source
  KAKENHI-PROJECT-20K07495, KAKENHI-PROJECT-23K18214, KAKENHI-PROJECT-22KK0096, KAKENHI-PUBLICLY-22H04809, KAKENHI-PROJECT-23K24130
[Journal Article] Genomic dissection of the Vibrio cholerae O-serogroup global reference strains: reassessing our view of diversity and plasticity between two chromosomes2022
- Author(s)
  Murase Kazunori、Arakawa Eiji、Izumiya Hidemasa、Iguchi Atsushi、Takemura Taichiro、Kikuchi Taisei、Nakagawa Ichiro、Thomson Nicholas R.、Ohnishi Makoto、Morita Masatomo
- Journal Title
  
  Microbial Genomics
  
  Volume: 8 Issue: 8 Pages: 860-860
- DOI
  10.1099/mgen.0.000860
- Peer Reviewed / Open Access / Int'l Joint Research
- Data Source
  KAKENHI-PROJECT-22KK0096, KAKENHI-PROJECT-20K07495, KAKENHI-PROJECT-21K19376, KAKENHI-PUBLICLY-22H04809, KAKENHI-PROJECT-23K24130
[Journal Article] Global population structure of the Serratia marcescens complex and identification of hospital-adapted lineages in the complex2022
- Author(s)
  Ono Tomoyuki、Taniguchi Itsuki、Nakamura Keiji、Nagano Debora Satie、Nishida Ruriko、Gotoh Yasuhiro、Ogura Yoshitoshi、Sato Mitsuhiko P.、Iguchi Atsushi、Murase Kazunori、Yoshimura Dai、Itoh Takehiko、Shima Ayaka、Dubois Damien、Oswald Eric、Shiose Akira、Gotoh Naomasa、Hayashi Tetsuya
- Journal Title
  
  Microbial Genomics
  
  Volume: 8 Issue: 3
- DOI
  10.1099/mgen.0.000793
- Peer Reviewed / Open Access / Int'l Joint Research
- Data Source
  KAKENHI-PROJECT-18K16175, KAKENHI-PROJECT-19H03472, KAKENHI-PROJECT-20K07495
[Journal Article] Cytolysin A (ClyA): A Bacterial Virulence Factor with Potential Applications in Nanopore Technology, Vaccine Development, and Tumor Therapy2022
- Author(s)
  Murase Kazunori
- Journal Title
  
  Toxins
  
  Volume: 14 Issue: 2 Pages: 78-78
- DOI
  10.3390/toxins14020078
- Peer Reviewed / Open Access
- Data Source
  KAKENHI-PROJECT-20K07495
[Journal Article] Biological Effect of Streptococcus pyogenes-Released Extracellular Vesicles on Human Monocytic Cells, Induction of Cytotoxicity, and Inflammatory Response2021
- Author(s)
  Murase Kazunori、Aikawa Chihiro、Nozawa Takashi、Nakatake Ayako、Sakamoto Kuniyo、Kikuchi Taisei、Nakagawa Ichiro
- Journal Title
  
  Frontiers in Cellular and Infection Microbiology
  
  Volume: 11 Pages: 711144-711144
- DOI
  10.3389/fcimb.2021.711144
- Peer Reviewed / Open Access
- Data Source
  KAKENHI-PROJECT-18KK0193, KAKENHI-PUBLICLY-20H05504, KAKENHI-PROJECT-19H03471, KAKENHI-PROJECT-20K07495, KAKENHI-PROJECT-21K07023
[Journal Article] TBC1D9 regulates TBK1 activation through Ca 2+ signaling in selective autophagy.2020
- Author(s)
  Nozawa T, Sano S, Minowa-Nozawa A, Toh H, Nakajima S, Murase K, Aikawa C, Nakagawa I
- Journal Title
  
  Nature Communications
  
  Volume: 11(1) Pages: 1-16
- NAID
  120006840172
- Peer Reviewed
- Data Source
  KAKENHI-PROJECT-17K08825
[Journal Article] Characterization of pig saliva as the major natural habitat of Streptococcus suis by analyzing oral, fecal, vaginal, and environmental microbiota2019
- Author(s)
  Murase K, Watanabe T, Arai S, Kim H, Tohya M, Ishida-Kuroki K, Vo TH, Nguyen TPB, Nakagawa I, Osawa R, Nguyen NH, Sekizaki T
- Journal Title
  
  PLoS One
  
  Volume: 14(4)
- NAID
  120006634676
- Peer Reviewed
- Data Source
  KAKENHI-PROJECT-17K08825
[Journal Article] Phylogenetic relationship of prophages is affected by CRISPR selection in Group A Streptococcus2019
- Author(s)
  Yamada S, Shibasaki M, Murase K, Watanabe T, Aikawa C, Nozawa T, Nakagawa I. Phylogenetic relationship of prophages is affected by CRISPR selection in Group A Streptococcus
- Journal Title
  
  BMC Microbiology
  
  Volume: 19(1) Pages: 24-24
- Peer Reviewed
- Data Source
  KAKENHI-PROJECT-17K08825
[Journal Article] Distinct interacting core taxa in co-occurrence networks enable discrimination of polymicrobial oral diseases with similar symptoms2016
- Author(s)
  Shiba T, Watanabe T, Kachi H, Koyanagi T, Maruyama N, Murase K, Takeuchi Y, Maruyama F, Izumi Y, Nakagawa I
- Journal Title
  
  Scientific reports
  
  Volume: 6 Issue: 1 Pages: 30997-30997
- DOI
  10.1038/srep30997
- NAID
  120005841407
- Peer Reviewed / Acknowledgement Compliant / Open Access
- Data Source
  KAKENHI-PROJECT-26463129, KAKENHI-PROJECT-26860293, KAKENHI-PROJECT-16K08015
[Journal Article] DNA-based culture-independent analysis detects the presence of Group A Streptococcus in throat samples from healthy adults in Japan2016
- Author(s)
  T. Kulkarni, F. Maruyama, C. Aikawa, T. Nozawa, K. Murase, I. Nakagawa.
- Journal Title
  
  BMC Microbiology
  
  Volume: 16(1) Issue: 1 Pages: 237-237
- DOI
  10.1186/s12866-016-0858-5
- NAID
  120006223635
- Peer Reviewed / Open Access
- Data Source
  KAKENHI-PROJECT-26860293
[Journal Article] Hlyf Produced By Extraintestinal Pathogenic Escherichia Coli Is A Virulence Factor That Regulates Outer Membrane Vesicle Biogenesis2015
- Author(s)
  Kazunori Murase, Patricia Martin, Gaëlle Porcheron, Sébastien Houle, Emmanuelle Helloin, Marie Pénary, Jean-Philippe Nougayrède, Charles M. Dozois, Tetsuya Hayashi, and Eric Oswald
- Journal Title
  
  Journal of Infectious Diseases
  
  Volume: 213(5) Issue: 5 Pages: 856-865
- DOI
  10.1093/infdis/jiv506
- Peer Reviewed / Acknowledgement Compliant / Open Access / Int'l Joint Research
- Data Source
  KAKENHI-PROJECT-26860293
[Presentation] Genome diversity of Streptococcus dysgalactiae and the evolutional process with host switching2023
- Author(s)
  村瀬一典，柘植亮佑，中川一路
- Organizer
  第96回日本細菌学会総会
- Data Source
  KAKENHI-PROJECT-23K24130
[Presentation] Rab GTPase ネットワークが制御する細胞内膜輸送経路と細菌感染2023
- Author(s)
  野澤孝志，村瀬一典，中川一路
- Organizer
  第96回日本細菌学会総会
- Invited
- Data Source
  KAKENHI-PROJECT-23K24130
[Presentation] A 群レンサ球菌の金属恒常性タンパク質 AdcAII は亜鉛の獲得と病原性に必要である2023
- Author(s)
  相川知宏，清水玲秀，村瀬一典，野澤孝志，中川一路
- Organizer
  第96回日本細菌学会総会
- Data Source
  KAKENHI-PROJECT-23K24130
[Presentation] 液-液相分離体ストレス顆粒を介した生体防御システム2023
- Author(s)
  野澤孝志、野澤敦子、村瀬一典、中川一路
- Organizer
  第34回日本生体防御学会総会
- Invited
- Data Source
  KAKENHI-PROJECT-23K24130
[Presentation] 腸菌由来の細胞外小胞が A 群レンサ球菌に与える生物学的影響2023
- Author(s)
  河岸優，村瀬一典，中川一路
- Organizer
  第96回日本細菌学会総会
- Data Source
  KAKENHI-PROJECT-23K24130
[Presentation] 大腸菌由来の細胞外小胞が A 群レンサ球菌に与える生物学的影響2023
- Author(s)
  河岸優，村瀬一典，中川一路
- Organizer
  第96回日本細菌学会総会
- Data Source
  KAKENHI-PROJECT-23K24130
[Presentation] Intraspecific genome diversity of Streptococcus dysgalactiae and the evolutional process with host switching2022
- Author(s)
  村瀬一典
- Organizer
  第95回細菌学会総会
- Data Source
  KAKENHI-PROJECT-20K07495
[Presentation] 大腸菌由来の細胞外小胞がA群レンサ球菌に与える生物学的影響2022
- Author(s)
  河岸優、村瀬一典、中川一路
- Organizer
  第75回日本細菌学会関西支部会学術総会
- Data Source
  KAKENHI-PROJECT-23K24130
[Presentation] 大腸菌が産生する細胞外小胞を介した A 群レンサ球菌の細胞分裂障害2022
- Author(s)
  河岸優、村瀬一典、中川一路
- Organizer
  第52回レンサ球菌シンポジウム
- Data Source
  KAKENHI-PROJECT-23K24130
[Presentation] MntEを介した菌体内の金属ホメオスタシスの維持はA群レンサ球菌の生育や病原性に寄与する2022
- Author(s)
  清水玲秀, 相川知宏, 村瀬一典, 野澤孝志, 中川一路
- Organizer
  第75回日本細菌学会関西支部会学術総会
- Data Source
  KAKENHI-PROJECT-23K24130
[Presentation] A群レンサ球菌が産生する細胞外小胞の病原性への関与2021
- Author(s)
  村瀬一典
- Organizer
  第94回日本細菌学会総会
- Data Source
  KAKENHI-PROJECT-20K07495
[Presentation] Proteome analysis of extracellular vesicles produced by Streptococcus pyogenes2020
- Author(s)
  村瀬一典, 相川知宏, 中武彩子, 菊地泰生, 野澤孝志, 中川一路
- Organizer
  第93回日本細菌学会総会（2020）
- Data Source
  KAKENHI-PROJECT-17K08825
[Presentation] COMPREHENSIVE PHYLOGENETIC ANALYSIS AND GENOME DYNAMICS OF O-SEROGROUP REFERENCE STRAINS IN VIBRIO CHOLEARE2019
- Author(s)
  Kazunori Murase, Masatomo Morita, Taichiro Takemura, Eiji Arakawa, Hidemasa Izumiya, Ichiro Nakagawa, Taisei Kikuchi, Makoto Ohnishi
- Organizer
  8th Congress of European Microbiologists (FEMS 2019)
- Int'l Joint Research
- Data Source
  KAKENHI-PROJECT-17K08825
[Presentation] New mechanisms of outer membrane vesicles in Gram-negative bacteria2016
- Author(s)
  Kazunori Murase
- Organizer
  The International Seminar on Applied Microbial Ecology
- Place of Presentation
  Temuco, Chile
- Year and Date
  2016-11-25
- Invited / Int'l Joint Research
- Data Source
  KAKENHI-PROJECT-26860293
[Presentation] HlyFによって誘導される外膜小胞の産生機構の解明2016
- Author(s)
  村瀬一典, Patricia Martin, Jean-Philippe Nougayrede, 林哲也, Eric Oswald
- Organizer
  第89回日本細菌学会総会
- Place of Presentation
  大阪国際交流センター（大阪）
- Year and Date
  2016-03-23
- Data Source
  KAKENHI-PROJECT-26860293
[Presentation] HlyFによるOuter membrane vesicles（外膜小胞）産生誘導機構の解明
- Author(s)
  村瀬一典, Patricia Martin, Jean-Philippe Nougayrede, 林哲也, Eric Oswald
- Organizer
  第88回日本細菌学会総会
- Place of Presentation
  長良川国際会議場（岐阜県岐阜市）
- Year and Date
  2015-03-26 – 2015-03-28
- Data Source
  KAKENHI-PROJECT-26860293
[Presentation] Role for hemolysin F in the virulence of Escherichia coli
- Author(s)
  Martin P, Murase K, Houle S, Helloin E, Nougayrede JP, Dozois C, Hayashi T, Oswald E
- Organizer
  ASM 2014 (114th General Meeting, American Society for Microbiology)
- Place of Presentation
  Boston Convention & Exhibition Center (Boston, Massachusetts)
- Year and Date
  2014-05-17 – 2014-05-20
- Data Source
  KAKENHI-PROJECT-26860293

1. 中川一路 (70294113)

# of Collaborated Projects: 3 results

# of Collaborated Products: 13 results
2. 野澤孝志 (10598858)

# of Collaborated Projects: 3 results

# of Collaborated Products: 4 results
3. MARUYAMA FUMITO (30423122)

# of Collaborated Projects: 2 results

# of Collaborated Products: 1 results
4. 相川知宏 (70725499)

# of Collaborated Projects: 2 results

# of Collaborated Products: 4 results
5. 白石宗 (70725168)

# of Collaborated Projects: 2 results

# of Collaborated Products: 0 results
6. 小椋義俊 (40363585)

# of Collaborated Projects: 1 results

# of Collaborated Products: 0 results
7. 竹村太地郎 (60572899)

# of Collaborated Projects: 1 results

# of Collaborated Products: 0 results
8. 野中里佐 (70363265)

# of Collaborated Projects: 1 results

# of Collaborated Products: 0 results
9. 植木尚子 (50622023)

# of Collaborated Projects: 1 results

# of Collaborated Products: 0 results
10. 菊地泰生 (20353659)

# of Collaborated Projects: 1 results

# of Collaborated Products: 0 results
11. 藤博貴 (50963237)

# of Collaborated Projects: 1 results

# of Collaborated Products: 0 results
12. KOYANAGI Tatsuro

# of Collaborated Projects: 0 results

# of Collaborated Products: 1 results
13. MARUYAMA Noriko

# of Collaborated Projects: 0 results

# of Collaborated Products: 1 results
14. SHIBA Takahiko

# of Collaborated Projects: 0 results

# of Collaborated Products: 1 results
15. 竹内康雄

# of Collaborated Projects: 0 results

# of Collaborated Products: 1 results
16. 小野友行

# of Collaborated Projects: 0 results

# of Collaborated Products: 1 results

Murase Kazunori 村瀬 一典

Kazunori Murase 村瀬 一典

Research Projects

Research Products

Co-Researchers

多様な細菌感染に対するロバストネスを支えるゼノファジー機構の解明

Principal Investigator

Project Period (FY)

Research Category

Review Section

Research Institution

Elucidation of the mechanism of competition between different bacterial species via extracellular vesicles and its application to the development of new drug discovery technologiesPrincipal Investigator

Principal Investigator

Project Period (FY)

Research Category

Review Section

Research Institution

Elucidation of Xenophagy Regulation by Bacterial Glycan Recognition

Principal Investigator

Project Period (FY)

Research Category

Review Section

Research Institution

Involvement in the pathogenic mechanism of Streptococcus pyogenes-derived extracellular vesiclesPrincipal Investigator

Principal Investigator

Project Period (FY)

Research Category

Review Section

Research Institution

Analysis of membrane vesicle production in gram negative bacteriaPrincipal Investigator

Principal Investigator

Project Period (FY)

Research Category

Research Field

Research Institution

Elucidation of genomic diversification in Group A Streptococci based on GWAS and epigenome

Principal Investigator

Project Period (FY)

Research Category

Research Field

Research Institution

Validation of microbial safety in drinking water in the world by comparative genomics of Vibrio cholerae

Principal Investigator

Project Period (FY)

Research Category

Research Field

Research Institution

Elucidation of the mechanism of outer membrane vesicles production in Gram-negative bacteriaPrincipal Investigator

Principal Investigator

Project Period (FY)

Research Category

Research Field

Research Institution

[Journal Article] Group A Streptococcus cation diffusion facilitator proteins contribute to immune evasion by regulating intracellular metal concentrations2023

Author(s)

Journal Title

DOI

Data Source

[Journal Article] Rab41-mediated ESCRT machinery repairs membrane rupture by a bacterial toxin in xenophagy2023

Author(s)

Journal Title

DOI

Data Source

[Journal Article] Clonal Expansion of Multidrug-Resistant <i>Streptococcus dysgalactiae</i> Subspecies <i>equisimilis</i> Causing Bacteremia, Japan, 2005?20212023

Author(s)

Journal Title

DOI

Data Source

[Journal Article] Genomic dissection of the Vibrio cholerae O-serogroup global reference strains: reassessing our view of diversity and plasticity between two chromosomes2022

Author(s)

Journal Title

DOI

Data Source

[Journal Article] Global population structure of the Serratia marcescens complex and identification of hospital-adapted lineages in the complex2022

Author(s)

Journal Title

DOI

Data Source

[Journal Article] Cytolysin A (ClyA): A Bacterial Virulence Factor with Potential Applications in Nanopore Technology, Vaccine Development, and Tumor Therapy2022

Author(s)

Murase Kazunori 村瀬一典

Kazunori Murase 村瀬一典

[Presentation] A 群レンサ球菌の金属恒常性タンパク質 AdcAII は亜鉛の獲得と病原性に必要である2023