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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">The structural and accessory proteins M, ORF 4a, ORF 4b, and ORF 5 of Middle East respiratory syndrome coronavirus (MERS-CoV) are potent interferon antagonists</title><author><name sortKey="Yang, Yang" sort="Yang, Yang" uniqKey="Yang Y" first="Yang" last="Yang">Yang Yang</name><affiliation><nlm:aff id="Aff1"><institution-wrap><institution-id institution-id-type="GRID">grid.198530.6</institution-id><institution-id institution-id-type="ISNI">0000000088032373</institution-id><institution>Key Laboratory of Medical Virology, Ministry of Health, National Institute for Viral Disease Control and Prevention,</institution><institution>Chinese Center for Disease Control and Prevention,</institution></institution-wrap>Beijing, 102206 China</nlm:aff></affiliation></author><author><name sortKey="Zhang, Ling" sort="Zhang, Ling" uniqKey="Zhang L" first="Ling" last="Zhang">Ling Zhang</name><affiliation><nlm:aff id="Aff1"><institution-wrap><institution-id institution-id-type="GRID">grid.198530.6</institution-id><institution-id institution-id-type="ISNI">0000000088032373</institution-id><institution>Key Laboratory of Medical Virology, Ministry of Health, National Institute for Viral Disease Control and Prevention,</institution><institution>Chinese Center for Disease Control and Prevention,</institution></institution-wrap>Beijing, 102206 China</nlm:aff></affiliation></author><author><name sortKey="Geng, Heyuan" sort="Geng, Heyuan" uniqKey="Geng H" first="Heyuan" last="Geng">Heyuan Geng</name><affiliation><nlm:aff id="Aff1"><institution-wrap><institution-id institution-id-type="GRID">grid.198530.6</institution-id><institution-id institution-id-type="ISNI">0000000088032373</institution-id><institution>Key Laboratory of Medical Virology, Ministry of Health, National Institute for Viral Disease Control and Prevention,</institution><institution>Chinese Center for Disease Control and Prevention,</institution></institution-wrap>Beijing, 102206 China</nlm:aff></affiliation></author><author><name sortKey="Deng, Yao" sort="Deng, Yao" uniqKey="Deng Y" first="Yao" last="Deng">Yao Deng</name><affiliation><nlm:aff id="Aff1"><institution-wrap><institution-id institution-id-type="GRID">grid.198530.6</institution-id><institution-id institution-id-type="ISNI">0000000088032373</institution-id><institution>Key Laboratory of Medical Virology, Ministry of Health, National Institute for Viral Disease Control and Prevention,</institution><institution>Chinese Center for Disease Control and Prevention,</institution></institution-wrap>Beijing, 102206 China</nlm:aff></affiliation></author><author><name sortKey="Huang, Baoying" sort="Huang, Baoying" uniqKey="Huang B" first="Baoying" last="Huang">Baoying Huang</name><affiliation><nlm:aff id="Aff1"><institution-wrap><institution-id institution-id-type="GRID">grid.198530.6</institution-id><institution-id institution-id-type="ISNI">0000000088032373</institution-id><institution>Key Laboratory of Medical Virology, Ministry of Health, National Institute for Viral Disease Control and Prevention,</institution><institution>Chinese Center for Disease Control and Prevention,</institution></institution-wrap>Beijing, 102206 China</nlm:aff></affiliation></author><author><name sortKey="Guo, Yin" sort="Guo, Yin" uniqKey="Guo Y" first="Yin" last="Guo">Yin Guo</name><affiliation><nlm:aff id="Aff2"><institution-wrap><institution-id institution-id-type="GRID">grid.413106.1</institution-id><institution-id institution-id-type="ISNI">0000000098896335</institution-id><institution>Institute of Materia Medica (IMM),</institution><institution>Chinese Academy of Medical Sciences and Peking Union Medical College,</institution></institution-wrap>Beijing, 100050 China</nlm:aff></affiliation></author><author><name sortKey="Zhao, Zhengdong" sort="Zhao, Zhengdong" uniqKey="Zhao Z" first="Zhengdong" last="Zhao">Zhengdong Zhao</name><affiliation><nlm:aff id="Aff3"><institution-wrap><institution-id institution-id-type="GRID">grid.410318.f</institution-id><institution-id institution-id-type="ISNI">0000000406323409</institution-id><institution>Key Laboratory of Pathogen System Biology, Ministry of Health, Institute of Pathogen Biology,</institution><institution>Chinese Academy of Medical Sciences,</institution></institution-wrap>Beijing, 100176 China</nlm:aff></affiliation></author><author><name sortKey="Tan, Wenjie" sort="Tan, Wenjie" uniqKey="Tan W" first="Wenjie" last="Tan">Wenjie Tan</name><affiliation><nlm:aff id="Aff1"><institution-wrap><institution-id institution-id-type="GRID">grid.198530.6</institution-id><institution-id institution-id-type="ISNI">0000000088032373</institution-id><institution>Key Laboratory of Medical Virology, Ministry of Health, National Institute for Viral Disease Control and Prevention,</institution><institution>Chinese Center for Disease Control and Prevention,</institution></institution-wrap>Beijing, 102206 China</nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">24318862</idno><idno type="pmc">4875403</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4875403</idno><idno type="RBID">PMC:4875403</idno><idno type="doi">10.1007/s13238-013-3096-8</idno><date when="2013">2013</date><idno type="wicri:Area/Pmc/Corpus">000421</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000421</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">The structural and accessory proteins M, ORF 4a, ORF 4b, and ORF 5 of Middle East respiratory syndrome coronavirus (MERS-CoV) are potent interferon antagonists</title><author><name sortKey="Yang, Yang" sort="Yang, Yang" uniqKey="Yang Y" first="Yang" last="Yang">Yang Yang</name><affiliation><nlm:aff id="Aff1"><institution-wrap><institution-id institution-id-type="GRID">grid.198530.6</institution-id><institution-id institution-id-type="ISNI">0000000088032373</institution-id><institution>Key Laboratory of Medical Virology, Ministry of Health, National Institute for Viral Disease Control and Prevention,</institution><institution>Chinese Center for Disease Control and Prevention,</institution></institution-wrap>Beijing, 102206 China</nlm:aff></affiliation></author><author><name sortKey="Zhang, Ling" sort="Zhang, Ling" uniqKey="Zhang L" first="Ling" last="Zhang">Ling Zhang</name><affiliation><nlm:aff id="Aff1"><institution-wrap><institution-id institution-id-type="GRID">grid.198530.6</institution-id><institution-id institution-id-type="ISNI">0000000088032373</institution-id><institution>Key Laboratory of Medical Virology, Ministry of Health, National Institute for Viral Disease Control and Prevention,</institution><institution>Chinese Center for Disease Control and Prevention,</institution></institution-wrap>Beijing, 102206 China</nlm:aff></affiliation></author><author><name sortKey="Geng, Heyuan" sort="Geng, Heyuan" uniqKey="Geng H" first="Heyuan" last="Geng">Heyuan Geng</name><affiliation><nlm:aff id="Aff1"><institution-wrap><institution-id institution-id-type="GRID">grid.198530.6</institution-id><institution-id institution-id-type="ISNI">0000000088032373</institution-id><institution>Key Laboratory of Medical Virology, Ministry of Health, National Institute for Viral Disease Control and Prevention,</institution><institution>Chinese Center for Disease Control and Prevention,</institution></institution-wrap>Beijing, 102206 China</nlm:aff></affiliation></author><author><name sortKey="Deng, Yao" sort="Deng, Yao" uniqKey="Deng Y" first="Yao" last="Deng">Yao Deng</name><affiliation><nlm:aff id="Aff1"><institution-wrap><institution-id institution-id-type="GRID">grid.198530.6</institution-id><institution-id institution-id-type="ISNI">0000000088032373</institution-id><institution>Key Laboratory of Medical Virology, Ministry of Health, National Institute for Viral Disease Control and Prevention,</institution><institution>Chinese Center for Disease Control and Prevention,</institution></institution-wrap>Beijing, 102206 China</nlm:aff></affiliation></author><author><name sortKey="Huang, Baoying" sort="Huang, Baoying" uniqKey="Huang B" first="Baoying" last="Huang">Baoying Huang</name><affiliation><nlm:aff id="Aff1"><institution-wrap><institution-id institution-id-type="GRID">grid.198530.6</institution-id><institution-id institution-id-type="ISNI">0000000088032373</institution-id><institution>Key Laboratory of Medical Virology, Ministry of Health, National Institute for Viral Disease Control and Prevention,</institution><institution>Chinese Center for Disease Control and Prevention,</institution></institution-wrap>Beijing, 102206 China</nlm:aff></affiliation></author><author><name sortKey="Guo, Yin" sort="Guo, Yin" uniqKey="Guo Y" first="Yin" last="Guo">Yin Guo</name><affiliation><nlm:aff id="Aff2"><institution-wrap><institution-id institution-id-type="GRID">grid.413106.1</institution-id><institution-id institution-id-type="ISNI">0000000098896335</institution-id><institution>Institute of Materia Medica (IMM),</institution><institution>Chinese Academy of Medical Sciences and Peking Union Medical College,</institution></institution-wrap>Beijing, 100050 China</nlm:aff></affiliation></author><author><name sortKey="Zhao, Zhengdong" sort="Zhao, Zhengdong" uniqKey="Zhao Z" first="Zhengdong" last="Zhao">Zhengdong Zhao</name><affiliation><nlm:aff id="Aff3"><institution-wrap><institution-id institution-id-type="GRID">grid.410318.f</institution-id><institution-id institution-id-type="ISNI">0000000406323409</institution-id><institution>Key Laboratory of Pathogen System Biology, Ministry of Health, Institute of Pathogen Biology,</institution><institution>Chinese Academy of Medical Sciences,</institution></institution-wrap>Beijing, 100176 China</nlm:aff></affiliation></author><author><name sortKey="Tan, Wenjie" sort="Tan, Wenjie" uniqKey="Tan W" first="Wenjie" last="Tan">Wenjie Tan</name><affiliation><nlm:aff id="Aff1"><institution-wrap><institution-id institution-id-type="GRID">grid.198530.6</institution-id><institution-id institution-id-type="ISNI">0000000088032373</institution-id><institution>Key Laboratory of Medical Virology, Ministry of Health, National Institute for Viral Disease Control and Prevention,</institution><institution>Chinese Center for Disease Control and Prevention,</institution></institution-wrap>Beijing, 102206 China</nlm:aff></affiliation></author></analytic><series><title level="j">Protein & Cell</title><idno type="ISSN">1674-800X</idno><idno type="eISSN">1674-8018</idno><imprint><date when="2013">2013</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>The newly emerged Middle East respiratory syndrome coronavirus (MERS-CoV) is a highly pathogenic respiratory virus with pathogenic mechanisms that may be driven by innate immune pathways. The goal of this study is to characterize the expression of the structural (S, E, M, N) and accessory (ORF 3, ORF 4a, ORF 4b, ORF 5) proteins of MERS-CoV and to determine whether any of these proteins acts as an interferon antagonist. Individual structural and accessory protein-coding plasmids with an N-terminal HA tag were constructed and transiently transfected into cells, and their native expression and subcellular localization were assessed using Wes tern blotting and indirect immunofluorescence. While ORF 4b demonstrated majorly nuclear localization, all of the other proteins demonstrated cytoplasmic localization. In addition, for the first time, our experiments revealed that the M, ORF 4a, ORF 4b, and ORF 5 proteins are potent interferon antagonists. Further examination revealed that the ORF 4a protein of MERS-CoV has the most potential to counteract the antiviral effects of IFN via the inhibition of both the interferon production (IFN-β promoter activity, IRF-3/7 and NF-κB activation) and ISRE promoter element signaling pathways. Together, our results provide new insights into the function and pathogenic role of the structural and accessory proteins of MERS-CoV.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Basler, Cf" uniqKey="Basler C">CF Basler</name></author><author><name sortKey="Wang, X" uniqKey="Wang X">X Wang</name></author><author><name sortKey="Muhlberger, E" uniqKey="Muhlberger E">E Mühlberger</name></author><author><name sortKey="Volchkov, V" uniqKey="Volchkov V">V Volchkov</name></author><author><name sortKey="Paragas, J" uniqKey="Paragas J">J Paragas</name></author><author><name sortKey="Klenk, Hd" uniqKey="Klenk H">HD Klenk</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Breban, R" uniqKey="Breban R">R Breban</name></author><author><name sortKey="Riou, J" uniqKey="Riou J">J Riou</name></author><author><name sortKey="Fontanet, A" uniqKey="Fontanet A">A 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<front><journal-meta><journal-id journal-id-type="nlm-ta">Protein Cell</journal-id><journal-id journal-id-type="iso-abbrev">Protein Cell</journal-id><journal-title-group><journal-title>Protein & Cell</journal-title></journal-title-group><issn pub-type="ppub">1674-800X</issn><issn pub-type="epub">1674-8018</issn><publisher><publisher-name>Higher Education Press</publisher-name><publisher-loc>Beijing</publisher-loc></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">24318862</article-id><article-id pub-id-type="pmc">4875403</article-id><article-id pub-id-type="publisher-id">3096</article-id><article-id pub-id-type="doi">10.1007/s13238-013-3096-8</article-id><article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group></article-categories><title-group><article-title>The structural and accessory proteins M, ORF 4a, ORF 4b, and ORF 5 of Middle East respiratory syndrome coronavirus (MERS-CoV) are potent interferon antagonists</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Yang</surname><given-names>Yang</given-names></name><xref ref-type="aff" rid="Aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Zhang</surname><given-names>Ling</given-names></name><xref ref-type="aff" rid="Aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Geng</surname><given-names>Heyuan</given-names></name><xref ref-type="aff" rid="Aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Deng</surname><given-names>Yao</given-names></name><xref ref-type="aff" rid="Aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Huang</surname><given-names>Baoying</given-names></name><xref ref-type="aff" rid="Aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Guo</surname><given-names>Yin</given-names></name><xref ref-type="aff" rid="Aff2">2</xref></contrib><contrib contrib-type="author" corresp="yes"><name><surname>Zhao</surname><given-names>Zhengdong</given-names></name><address><email>timjszzd@163.com</email></address><xref ref-type="aff" rid="Aff3">3</xref></contrib><contrib contrib-type="author" corresp="yes"><name><surname>Tan</surname><given-names>Wenjie</given-names></name><address><email>tanwj28@yahoo.cn</email></address><xref ref-type="aff" rid="Aff1">1</xref></contrib><aff id="Aff1"><label>1</label><institution-wrap><institution-id institution-id-type="GRID">grid.198530.6</institution-id><institution-id institution-id-type="ISNI">0000000088032373</institution-id><institution>Key Laboratory of Medical Virology, Ministry of Health, National Institute for Viral Disease Control and Prevention,</institution><institution>Chinese Center for Disease Control and Prevention,</institution></institution-wrap>Beijing, 102206 China</aff><aff id="Aff2"><label>2</label><institution-wrap><institution-id institution-id-type="GRID">grid.413106.1</institution-id><institution-id institution-id-type="ISNI">0000000098896335</institution-id><institution>Institute of Materia Medica (IMM),</institution><institution>Chinese Academy of Medical Sciences and Peking Union Medical College,</institution></institution-wrap>Beijing, 100050 China</aff><aff id="Aff3"><label>3</label><institution-wrap><institution-id institution-id-type="GRID">grid.410318.f</institution-id><institution-id institution-id-type="ISNI">0000000406323409</institution-id><institution>Key Laboratory of Pathogen System Biology, Ministry of Health, Institute of Pathogen Biology,</institution><institution>Chinese Academy of Medical Sciences,</institution></institution-wrap>Beijing, 100176 China</aff></contrib-group><pub-date pub-type="epub"><day>8</day><month>12</month><year>2013</year></pub-date><pub-date pub-type="ppub"><month>12</month><year>2013</year></pub-date><volume>4</volume><issue>12</issue><fpage>951</fpage><lpage>961</lpage><history><date date-type="received"><day>29</day><month>9</month><year>2013</year></date><date date-type="accepted"><day>1</day><month>11</month><year>2013</year></date></history><permissions><copyright-statement>© Higher Education Press and Springer-Verlag Berlin Heidelberg 2013</copyright-statement><license><license-p>This article is made available via the PMC Open Access Subset for unrestricted research re-use and secondary analysis in any form or by any means with acknowledgement of the original source. These permissions are granted for the duration of the World Health Organization (WHO) declaration of COVID-19 as a global pandemic.</license-p></license></permissions><abstract id="Abs1"><p>The newly emerged Middle East respiratory syndrome coronavirus (MERS-CoV) is a highly pathogenic respiratory virus with pathogenic mechanisms that may be driven by innate immune pathways. The goal of this study is to characterize the expression of the structural (S, E, M, N) and accessory (ORF 3, ORF 4a, ORF 4b, ORF 5) proteins of MERS-CoV and to determine whether any of these proteins acts as an interferon antagonist. Individual structural and accessory protein-coding plasmids with an N-terminal HA tag were constructed and transiently transfected into cells, and their native expression and subcellular localization were assessed using Wes tern blotting and indirect immunofluorescence. While ORF 4b demonstrated majorly nuclear localization, all of the other proteins demonstrated cytoplasmic localization. In addition, for the first time, our experiments revealed that the M, ORF 4a, ORF 4b, and ORF 5 proteins are potent interferon antagonists. Further examination revealed that the ORF 4a protein of MERS-CoV has the most potential to counteract the antiviral effects of IFN via the inhibition of both the interferon production (IFN-β promoter activity, IRF-3/7 and NF-κB activation) and ISRE promoter element signaling pathways. Together, our results provide new insights into the function and pathogenic role of the structural and accessory proteins of MERS-CoV.</p></abstract><kwd-group xml:lang="en"><title>Keywords</title><kwd>MERS-CoV</kwd><kwd>structural proteins</kwd><kwd>accessory proteins</kwd><kwd>interferon antagonists</kwd></kwd-group><custom-meta-group><custom-meta><meta-name>issue-copyright-statement</meta-name><meta-value>© HEP and Springer 2013</meta-value></custom-meta></custom-meta-group></article-meta></front><back><fn-group><fn><p>These authors contributed equally to the work.</p></fn></fn-group><ref-list id="Bib1"><title>References</title><ref id="CR1"><element-citation publication-type="journal"><person-group 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