HERP Binds TBK1 To Activate Innate Immunity and Repress Virus Replication in Response to Endoplasmic Reticulum Stress.
Identifieur interne : 000305 ( PubMed/Corpus ); précédent : 000304; suivant : 000306HERP Binds TBK1 To Activate Innate Immunity and Repress Virus Replication in Response to Endoplasmic Reticulum Stress.
Auteurs : Maolin Ge ; Zhen Luo ; Zhi Qiao ; Yao Zhou ; Xin Cheng ; Qibin Geng ; Yanyan Cai ; Pin Wan ; Ying Xiong ; Fang Liu ; Kailang Wu ; Yingle Liu ; Jianguo WuSource :
- Journal of immunology (Baltimore, Md. : 1950) [ 1550-6606 ] ; 2017.
English descriptors
- KwdEn :
- Animals, Endoplasmic Reticulum Stress (genetics), Endoplasmic Reticulum Stress (immunology), Female, Humans, Immunity, Innate, Interferon Regulatory Factor-3 (genetics), Interferon Regulatory Factor-3 (immunology), Interferons (genetics), Interferons (immunology), Male, Membrane Proteins (genetics), Membrane Proteins (immunology), Mice, Mice, Inbred BALB C, Protein-Serine-Threonine Kinases (genetics), Protein-Serine-Threonine Kinases (immunology), RNA Virus Infections (genetics), RNA Virus Infections (immunology), RNA Virus Infections (pathology), RNA Viruses (physiology), Virus Replication (immunology).
- MESH :
- chemical , genetics : Interferon Regulatory Factor-3, Interferons, Membrane Proteins, Protein-Serine-Threonine Kinases.
- genetics : Endoplasmic Reticulum Stress, RNA Virus Infections.
- immunology : Endoplasmic Reticulum Stress, Interferon Regulatory Factor-3, Interferons, Membrane Proteins, Protein-Serine-Threonine Kinases, RNA Virus Infections, Virus Replication.
- pathology : RNA Virus Infections.
- physiology : RNA Viruses.
- Animals, Female, Humans, Immunity, Innate, Male, Mice, Mice, Inbred BALB C.
Abstract
Host innate immunity is crucial for cellular responses against viral infection sensed by distinct pattern recognition receptors and endoplasmic reticulum (ER) stress. Enterovirus 71 (EV71) is a causative agent of hand, foot, and mouth disease and neurological diseases. However, the exact mechanism underlying the link between ER stress induced by EV71 infection and host innate immunity is largely unknown. In this study, we demonstrated that EV71 infection induces the homocysteine-induced ER protein (HERP), a modulator of the ER stress response which is dependent on the participation of MAVS. Virus-induced HERP subsequently stimulates host innate immunity to repress viral replication by promoting type-I IFNs (IFN-α and IFN-β) and type-III IFN (IFN-λ1) expression. Through interacting with TANK-binding kinase 1, HERP amplifies the MAVS signaling and facilitates the phosphorylation and nuclear translocation of IFN regulatory factor 3 and NF-κB to enhance the expression of IFNs, which leads to a broad inhibition of the replication of RNA viruses, including EV71, Sendai virus, influenza A virus, and vesicular stomatitis virus. Therefore, we demonstrated that HERP plays an important role in the regulation of host innate immunity in response to ER stress during the infection of RNA viruses. These findings provide new insights into the mechanism underlying the replication of RNA viruses and the production of IFNs, and also demonstrate a new role of HERP in the regulation of host innate immunity in response to viral infection.
DOI: 10.4049/jimmunol.1700376
PubMed: 28954889
Links to Exploration step
pubmed:28954889Le document en format XML
<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">HERP Binds TBK1 To Activate Innate Immunity and Repress Virus Replication in Response to Endoplasmic Reticulum Stress.</title><author><name sortKey="Ge, Maolin" sort="Ge, Maolin" uniqKey="Ge M" first="Maolin" last="Ge">Maolin Ge</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Luo, Zhen" sort="Luo, Zhen" uniqKey="Luo Z" first="Zhen" last="Luo">Zhen Luo</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Qiao, Zhi" sort="Qiao, Zhi" uniqKey="Qiao Z" first="Zhi" last="Qiao">Zhi Qiao</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Zhou, Yao" sort="Zhou, Yao" uniqKey="Zhou Y" first="Yao" last="Zhou">Yao Zhou</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Cheng, Xin" sort="Cheng, Xin" uniqKey="Cheng X" first="Xin" last="Cheng">Xin Cheng</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Geng, Qibin" sort="Geng, Qibin" uniqKey="Geng Q" first="Qibin" last="Geng">Qibin Geng</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Cai, Yanyan" sort="Cai, Yanyan" uniqKey="Cai Y" first="Yanyan" last="Cai">Yanyan Cai</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Wan, Pin" sort="Wan, Pin" uniqKey="Wan P" first="Pin" last="Wan">Pin Wan</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; 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and jwu@whu.edu.cn mvlwu@whu.edu.cn wukailang@whu.edu.cn.</nlm:affiliation></affiliation></author><author><name sortKey="Wu, Jianguo" sort="Wu, Jianguo" uniqKey="Wu J" first="Jianguo" last="Wu">Jianguo Wu</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and jwu@whu.edu.cn mvlwu@whu.edu.cn wukailang@whu.edu.cn.</nlm:affiliation></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PubMed</idno><date when="2017">2017</date><idno type="RBID">pubmed:28954889</idno><idno type="pmid">28954889</idno><idno type="doi">10.4049/jimmunol.1700376</idno><idno type="wicri:Area/PubMed/Corpus">000305</idno><idno type="wicri:explorRef" wicri:stream="PubMed" wicri:step="Corpus" wicri:corpus="PubMed">000305</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en">HERP Binds TBK1 To Activate Innate Immunity and Repress Virus Replication in Response to Endoplasmic Reticulum Stress.</title><author><name sortKey="Ge, Maolin" sort="Ge, Maolin" uniqKey="Ge M" first="Maolin" last="Ge">Maolin Ge</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Luo, Zhen" sort="Luo, Zhen" uniqKey="Luo Z" first="Zhen" last="Luo">Zhen Luo</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Qiao, Zhi" sort="Qiao, Zhi" uniqKey="Qiao Z" first="Zhi" last="Qiao">Zhi Qiao</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Zhou, Yao" sort="Zhou, Yao" uniqKey="Zhou Y" first="Yao" last="Zhou">Yao Zhou</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Cheng, Xin" sort="Cheng, Xin" uniqKey="Cheng X" first="Xin" last="Cheng">Xin Cheng</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Geng, Qibin" sort="Geng, Qibin" uniqKey="Geng Q" first="Qibin" last="Geng">Qibin Geng</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Cai, Yanyan" sort="Cai, Yanyan" uniqKey="Cai Y" first="Yanyan" last="Cai">Yanyan Cai</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Wan, Pin" sort="Wan, Pin" uniqKey="Wan P" first="Pin" last="Wan">Pin Wan</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Xiong, Ying" sort="Xiong, Ying" uniqKey="Xiong Y" first="Ying" last="Xiong">Ying Xiong</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Liu, Fang" sort="Liu, Fang" uniqKey="Liu F" first="Fang" last="Liu">Fang Liu</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</nlm:affiliation></affiliation></author><author><name sortKey="Wu, Kailang" sort="Wu, Kailang" uniqKey="Wu K" first="Kailang" last="Wu">Kailang Wu</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and jwu@whu.edu.cn mvlwu@whu.edu.cn wukailang@whu.edu.cn.</nlm:affiliation></affiliation></author><author><name sortKey="Liu, Yingle" sort="Liu, Yingle" uniqKey="Liu Y" first="Yingle" last="Liu">Yingle Liu</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and jwu@whu.edu.cn mvlwu@whu.edu.cn wukailang@whu.edu.cn.</nlm:affiliation></affiliation></author><author><name sortKey="Wu, Jianguo" sort="Wu, Jianguo" uniqKey="Wu J" first="Jianguo" last="Wu">Jianguo Wu</name><affiliation><nlm:affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and jwu@whu.edu.cn mvlwu@whu.edu.cn wukailang@whu.edu.cn.</nlm:affiliation></affiliation></author></analytic><series><title level="j">Journal of immunology (Baltimore, Md. : 1950)</title><idno type="eISSN">1550-6606</idno><imprint><date when="2017" type="published">2017</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass><keywords scheme="KwdEn" xml:lang="en"><term>Animals</term><term>Endoplasmic Reticulum Stress (genetics)</term><term>Endoplasmic Reticulum Stress (immunology)</term><term>Female</term><term>Humans</term><term>Immunity, Innate</term><term>Interferon Regulatory Factor-3 (genetics)</term><term>Interferon Regulatory Factor-3 (immunology)</term><term>Interferons (genetics)</term><term>Interferons (immunology)</term><term>Male</term><term>Membrane Proteins (genetics)</term><term>Membrane Proteins (immunology)</term><term>Mice</term><term>Mice, Inbred BALB C</term><term>Protein-Serine-Threonine Kinases (genetics)</term><term>Protein-Serine-Threonine Kinases (immunology)</term><term>RNA Virus Infections (genetics)</term><term>RNA Virus Infections (immunology)</term><term>RNA Virus Infections (pathology)</term><term>RNA Viruses (physiology)</term><term>Virus Replication (immunology)</term></keywords><keywords scheme="MESH" type="chemical" qualifier="genetics" xml:lang="en"><term>Interferon Regulatory Factor-3</term><term>Interferons</term><term>Membrane Proteins</term><term>Protein-Serine-Threonine Kinases</term></keywords><keywords scheme="MESH" qualifier="genetics" xml:lang="en"><term>Endoplasmic Reticulum Stress</term><term>RNA Virus Infections</term></keywords><keywords scheme="MESH" qualifier="immunology" xml:lang="en"><term>Endoplasmic Reticulum Stress</term><term>Interferon Regulatory Factor-3</term><term>Interferons</term><term>Membrane Proteins</term><term>Protein-Serine-Threonine Kinases</term><term>RNA Virus Infections</term><term>Virus Replication</term></keywords><keywords scheme="MESH" qualifier="pathology" xml:lang="en"><term>RNA Virus Infections</term></keywords><keywords scheme="MESH" qualifier="physiology" xml:lang="en"><term>RNA Viruses</term></keywords><keywords scheme="MESH" xml:lang="en"><term>Animals</term><term>Female</term><term>Humans</term><term>Immunity, Innate</term><term>Male</term><term>Mice</term><term>Mice, Inbred BALB C</term></keywords></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en">Host innate immunity is crucial for cellular responses against viral infection sensed by distinct pattern recognition receptors and endoplasmic reticulum (ER) stress. Enterovirus 71 (EV71) is a causative agent of hand, foot, and mouth disease and neurological diseases. However, the exact mechanism underlying the link between ER stress induced by EV71 infection and host innate immunity is largely unknown. In this study, we demonstrated that EV71 infection induces the homocysteine-induced ER protein (HERP), a modulator of the ER stress response which is dependent on the participation of MAVS. Virus-induced HERP subsequently stimulates host innate immunity to repress viral replication by promoting type-I IFNs (IFN-α and IFN-β) and type-III IFN (IFN-λ1) expression. Through interacting with TANK-binding kinase 1, HERP amplifies the MAVS signaling and facilitates the phosphorylation and nuclear translocation of IFN regulatory factor 3 and NF-κB to enhance the expression of IFNs, which leads to a broad inhibition of the replication of RNA viruses, including EV71, Sendai virus, influenza A virus, and vesicular stomatitis virus. Therefore, we demonstrated that HERP plays an important role in the regulation of host innate immunity in response to ER stress during the infection of RNA viruses. These findings provide new insights into the mechanism underlying the replication of RNA viruses and the production of IFNs, and also demonstrate a new role of HERP in the regulation of host innate immunity in response to viral infection.</div></front></TEI><pubmed><MedlineCitation Status="MEDLINE" Owner="NLM"><PMID Version="1">28954889</PMID><DateCompleted><Year>2017</Year><Month>10</Month><Day>26</Day></DateCompleted><DateRevised><Year>2018</Year><Month>01</Month><Day>29</Day></DateRevised><Article PubModel="Print-Electronic"><Journal><ISSN IssnType="Electronic">1550-6606</ISSN><JournalIssue CitedMedium="Internet"><Volume>199</Volume><Issue>9</Issue><PubDate><Year>2017</Year><Month>11</Month><Day>01</Day></PubDate></JournalIssue><Title>Journal of immunology (Baltimore, Md. : 1950)</Title><ISOAbbreviation>J. Immunol.</ISOAbbreviation></Journal><ArticleTitle>HERP Binds TBK1 To Activate Innate Immunity and Repress Virus Replication in Response to Endoplasmic Reticulum Stress.</ArticleTitle><Pagination><MedlinePgn>3280-3292</MedlinePgn></Pagination><ELocationID EIdType="doi" ValidYN="Y">10.4049/jimmunol.1700376</ELocationID><Abstract><AbstractText>Host innate immunity is crucial for cellular responses against viral infection sensed by distinct pattern recognition receptors and endoplasmic reticulum (ER) stress. Enterovirus 71 (EV71) is a causative agent of hand, foot, and mouth disease and neurological diseases. However, the exact mechanism underlying the link between ER stress induced by EV71 infection and host innate immunity is largely unknown. In this study, we demonstrated that EV71 infection induces the homocysteine-induced ER protein (HERP), a modulator of the ER stress response which is dependent on the participation of MAVS. Virus-induced HERP subsequently stimulates host innate immunity to repress viral replication by promoting type-I IFNs (IFN-α and IFN-β) and type-III IFN (IFN-λ1) expression. Through interacting with TANK-binding kinase 1, HERP amplifies the MAVS signaling and facilitates the phosphorylation and nuclear translocation of IFN regulatory factor 3 and NF-κB to enhance the expression of IFNs, which leads to a broad inhibition of the replication of RNA viruses, including EV71, Sendai virus, influenza A virus, and vesicular stomatitis virus. Therefore, we demonstrated that HERP plays an important role in the regulation of host innate immunity in response to ER stress during the infection of RNA viruses. These findings provide new insights into the mechanism underlying the replication of RNA viruses and the production of IFNs, and also demonstrate a new role of HERP in the regulation of host innate immunity in response to viral infection.</AbstractText><CopyrightInformation>Copyright © 2017 by The American Association of Immunologists, Inc.</CopyrightInformation></Abstract><AuthorList CompleteYN="Y"><Author ValidYN="Y"><LastName>Ge</LastName><ForeName>Maolin</ForeName><Initials>M</Initials><Identifier Source="ORCID">0000-0002-8803-1463</Identifier><AffiliationInfo><Affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Luo</LastName><ForeName>Zhen</ForeName><Initials>Z</Initials><Identifier Source="ORCID">0000-0002-1142-2845</Identifier><AffiliationInfo><Affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</Affiliation></AffiliationInfo><AffiliationInfo><Affiliation>Institute of Medical Microbiology, Jinan University, Guangzhou 510632, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Qiao</LastName><ForeName>Zhi</ForeName><Initials>Z</Initials><AffiliationInfo><Affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Zhou</LastName><ForeName>Yao</ForeName><Initials>Y</Initials><Identifier Source="ORCID">0000-0003-3994-3068</Identifier><AffiliationInfo><Affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Cheng</LastName><ForeName>Xin</ForeName><Initials>X</Initials><Identifier Source="ORCID">0000-0003-3331-9993</Identifier><AffiliationInfo><Affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Geng</LastName><ForeName>Qibin</ForeName><Initials>Q</Initials><Identifier Source="ORCID">0000-0003-4872-2400</Identifier><AffiliationInfo><Affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Cai</LastName><ForeName>Yanyan</ForeName><Initials>Y</Initials><AffiliationInfo><Affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Wan</LastName><ForeName>Pin</ForeName><Initials>P</Initials><Identifier Source="ORCID">0000-0003-0527-8952</Identifier><AffiliationInfo><Affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Xiong</LastName><ForeName>Ying</ForeName><Initials>Y</Initials><AffiliationInfo><Affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Liu</LastName><ForeName>Fang</ForeName><Initials>F</Initials><Identifier Source="ORCID">0000-0003-0972-9968</Identifier><AffiliationInfo><Affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Wu</LastName><ForeName>Kailang</ForeName><Initials>K</Initials><Identifier Source="ORCID">0000-0002-0926-9300</Identifier><AffiliationInfo><Affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and jwu@whu.edu.cn mvlwu@whu.edu.cn wukailang@whu.edu.cn.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Liu</LastName><ForeName>Yingle</ForeName><Initials>Y</Initials><AffiliationInfo><Affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and jwu@whu.edu.cn mvlwu@whu.edu.cn wukailang@whu.edu.cn.</Affiliation></AffiliationInfo><AffiliationInfo><Affiliation>Institute of Medical Microbiology, Jinan University, Guangzhou 510632, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Wu</LastName><ForeName>Jianguo</ForeName><Initials>J</Initials><AffiliationInfo><Affiliation>State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan 430072, China; and jwu@whu.edu.cn mvlwu@whu.edu.cn wukailang@whu.edu.cn.</Affiliation></AffiliationInfo><AffiliationInfo><Affiliation>Institute of Medical Microbiology, Jinan University, Guangzhou 510632, China.</Affiliation></AffiliationInfo></Author></AuthorList><Language>eng</Language><PublicationTypeList><PublicationType UI="D016428">Journal Article</PublicationType><PublicationType UI="D013485">Research Support, Non-U.S. Gov't</PublicationType></PublicationTypeList><ArticleDate DateType="Electronic"><Year>2017</Year><Month>09</Month><Day>27</Day></ArticleDate></Article><MedlineJournalInfo><Country>United States</Country><MedlineTA>J Immunol</MedlineTA><NlmUniqueID>2985117R</NlmUniqueID><ISSNLinking>0022-1767</ISSNLinking></MedlineJournalInfo><ChemicalList><Chemical><RegistryNumber>0</RegistryNumber><NameOfSubstance UI="C485932">HERPUD1 protein, human</NameOfSubstance></Chemical><Chemical><RegistryNumber>0</RegistryNumber><NameOfSubstance UI="C415829">Herpud1 protein, mouse</NameOfSubstance></Chemical><Chemical><RegistryNumber>0</RegistryNumber><NameOfSubstance UI="C494232">IRF3 protein, human</NameOfSubstance></Chemical><Chemical><RegistryNumber>0</RegistryNumber><NameOfSubstance 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