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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">NMR assignments of the macro domain from Middle East respiratory syndrome coronavirus (MERS-CoV)</title><author><name sortKey="Huang, Yi Ping" sort="Huang, Yi Ping" uniqKey="Huang Y" first="Yi-Ping" last="Huang">Yi-Ping Huang</name><affiliation><nlm:aff id="Aff1"><institution-wrap><institution-id institution-id-type="GRID">grid.28665.3f</institution-id><institution-id institution-id-type="ISNI">0000000122871366</institution-id><institution>Genomics Research Center,</institution><institution>Academia Sinica,</institution></institution-wrap>Nankang, Taipei, 11529 Taiwan</nlm:aff></affiliation></author><author><name sortKey="Cho, Chao Cheng" sort="Cho, Chao Cheng" uniqKey="Cho C" first="Chao-Cheng" last="Cho">Chao-Cheng Cho</name><affiliation><nlm:aff id="Aff2"><institution-wrap><institution-id institution-id-type="GRID">grid.19188.39</institution-id><institution-id institution-id-type="ISNI">0000000405460241</institution-id><institution>Genome and Systems Biology Degree Program,</institution><institution>National Taiwan University and Academia Sinica,</institution></institution-wrap>Taipei, 10617 Taiwan</nlm:aff></affiliation><affiliation><nlm:aff id="Aff3"><institution-wrap><institution-id institution-id-type="GRID">grid.19188.39</institution-id><institution-id institution-id-type="ISNI">0000000405460241</institution-id><institution>Department of Agricultural Chemistry,</institution><institution>National Taiwan University,</institution></institution-wrap>Taipei, 10617 Taiwan</nlm:aff></affiliation></author><author><name sortKey="Chang, Chi Fon" sort="Chang, Chi Fon" uniqKey="Chang C" first="Chi-Fon" last="Chang">Chi-Fon Chang</name><affiliation><nlm:aff id="Aff1"><institution-wrap><institution-id institution-id-type="GRID">grid.28665.3f</institution-id><institution-id institution-id-type="ISNI">0000000122871366</institution-id><institution>Genomics Research Center,</institution><institution>Academia Sinica,</institution></institution-wrap>Nankang, Taipei, 11529 Taiwan</nlm:aff></affiliation></author><author><name sortKey="Hsu, Chun Hua" sort="Hsu, Chun Hua" uniqKey="Hsu C" first="Chun-Hua" last="Hsu">Chun-Hua Hsu</name><affiliation><nlm:aff id="Aff2"><institution-wrap><institution-id institution-id-type="GRID">grid.19188.39</institution-id><institution-id institution-id-type="ISNI">0000000405460241</institution-id><institution>Genome and Systems Biology Degree Program,</institution><institution>National Taiwan University and Academia Sinica,</institution></institution-wrap>Taipei, 10617 Taiwan</nlm:aff></affiliation><affiliation><nlm:aff id="Aff3"><institution-wrap><institution-id institution-id-type="GRID">grid.19188.39</institution-id><institution-id institution-id-type="ISNI">0000000405460241</institution-id><institution>Department of Agricultural Chemistry,</institution><institution>National Taiwan University,</institution></institution-wrap>Taipei, 10617 Taiwan</nlm:aff></affiliation><affiliation><nlm:aff id="Aff4"><institution-wrap><institution-id institution-id-type="GRID">grid.19188.39</institution-id><institution-id institution-id-type="ISNI">0000000405460241</institution-id><institution>Center for Systems Biology,</institution><institution>National Taiwan University,</institution></institution-wrap>Taipei, 10617 Taiwan</nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">26993639</idno><idno type="pmc">7091117</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7091117</idno><idno type="RBID">PMC:7091117</idno><idno type="doi">10.1007/s12104-016-9676-9</idno><date when="2016">2016</date><idno type="wicri:Area/Pmc/Corpus">000341</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000341</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">NMR assignments of the macro domain from Middle East respiratory syndrome coronavirus (MERS-CoV)</title><author><name sortKey="Huang, Yi Ping" sort="Huang, Yi Ping" uniqKey="Huang Y" first="Yi-Ping" last="Huang">Yi-Ping Huang</name><affiliation><nlm:aff id="Aff1"><institution-wrap><institution-id institution-id-type="GRID">grid.28665.3f</institution-id><institution-id institution-id-type="ISNI">0000000122871366</institution-id><institution>Genomics Research Center,</institution><institution>Academia Sinica,</institution></institution-wrap>Nankang, Taipei, 11529 Taiwan</nlm:aff></affiliation></author><author><name sortKey="Cho, Chao Cheng" sort="Cho, Chao Cheng" uniqKey="Cho C" first="Chao-Cheng" last="Cho">Chao-Cheng Cho</name><affiliation><nlm:aff id="Aff2"><institution-wrap><institution-id institution-id-type="GRID">grid.19188.39</institution-id><institution-id institution-id-type="ISNI">0000000405460241</institution-id><institution>Genome and Systems Biology Degree Program,</institution><institution>National Taiwan University and Academia Sinica,</institution></institution-wrap>Taipei, 10617 Taiwan</nlm:aff></affiliation><affiliation><nlm:aff id="Aff3"><institution-wrap><institution-id institution-id-type="GRID">grid.19188.39</institution-id><institution-id institution-id-type="ISNI">0000000405460241</institution-id><institution>Department of Agricultural Chemistry,</institution><institution>National Taiwan University,</institution></institution-wrap>Taipei, 10617 Taiwan</nlm:aff></affiliation></author><author><name sortKey="Chang, Chi Fon" sort="Chang, Chi Fon" uniqKey="Chang C" first="Chi-Fon" last="Chang">Chi-Fon Chang</name><affiliation><nlm:aff id="Aff1"><institution-wrap><institution-id institution-id-type="GRID">grid.28665.3f</institution-id><institution-id institution-id-type="ISNI">0000000122871366</institution-id><institution>Genomics Research Center,</institution><institution>Academia Sinica,</institution></institution-wrap>Nankang, Taipei, 11529 Taiwan</nlm:aff></affiliation></author><author><name sortKey="Hsu, Chun Hua" sort="Hsu, Chun Hua" uniqKey="Hsu C" first="Chun-Hua" last="Hsu">Chun-Hua Hsu</name><affiliation><nlm:aff id="Aff2"><institution-wrap><institution-id institution-id-type="GRID">grid.19188.39</institution-id><institution-id institution-id-type="ISNI">0000000405460241</institution-id><institution>Genome and Systems Biology Degree Program,</institution><institution>National Taiwan University and Academia Sinica,</institution></institution-wrap>Taipei, 10617 Taiwan</nlm:aff></affiliation><affiliation><nlm:aff id="Aff3"><institution-wrap><institution-id institution-id-type="GRID">grid.19188.39</institution-id><institution-id institution-id-type="ISNI">0000000405460241</institution-id><institution>Department of Agricultural Chemistry,</institution><institution>National Taiwan University,</institution></institution-wrap>Taipei, 10617 Taiwan</nlm:aff></affiliation><affiliation><nlm:aff id="Aff4"><institution-wrap><institution-id institution-id-type="GRID">grid.19188.39</institution-id><institution-id institution-id-type="ISNI">0000000405460241</institution-id><institution>Center for Systems Biology,</institution><institution>National Taiwan University,</institution></institution-wrap>Taipei, 10617 Taiwan</nlm:aff></affiliation></author></analytic><series><title level="j">Biomolecular Nmr Assignments</title><idno type="ISSN">1874-2718</idno><idno type="eISSN">1874-270X</idno><imprint><date when="2016">2016</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p id="Par1">The newly emerging human pathogen, Middle East respiratory syndrome coronavirus (MERS-CoV), contains a macro domain in the highly conserved N-terminal region of non-structural protein 3. Intense research has shown that macro domains bind ADP-ribose and other derivatives, but it still remains intangible about their exact function. In this study we report the preliminary structural analysis through solution NMR spectroscopy of the MERS-CoV macro domain. The near complete NMR assignments of MERS-CoV macro domain provide the basis for subsequent structural and biochemical investigation in the context of protein function.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Chen, Yd" uniqKey="Chen Y">YD Chen</name></author><author><name sortKey="Wu, Sh" uniqKey="Wu S">SH Wu</name></author><author><name sortKey="Hsu, Ch" uniqKey="Hsu C">CH Hsu</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="De Groot, Rj" uniqKey="De Groot R">RJ de Groot</name></author><author><name sortKey="Baker, Sc" uniqKey="Baker S">SC Baker</name></author><author><name sortKey="Baric, Rs" uniqKey="Baric R">RS Baric</name></author><author><name sortKey="Brown, Cs" uniqKey="Brown C">CS Brown</name></author><author><name sortKey="Drosten, C" uniqKey="Drosten C">C Drosten</name></author><author><name sortKey="Enjuanes, L" uniqKey="Enjuanes L">L Enjuanes</name></author><author><name sortKey="Fouchier, Ra" uniqKey="Fouchier R">RA Fouchier</name></author><author><name sortKey="Galiano, M" uniqKey="Galiano M">M Galiano</name></author><author><name sortKey="Gorbalenya, Ae" uniqKey="Gorbalenya A">AE Gorbalenya</name></author><author><name sortKey="Memish, Za" uniqKey="Memish Z">ZA Memish</name></author><author><name sortKey="Perlman, S" uniqKey="Perlman S">S Perlman</name></author><author><name sortKey="Poon, Ll" uniqKey="Poon L">LL Poon</name></author><author><name sortKey="Snijder, Ej" uniqKey="Snijder E">EJ Snijder</name></author><author><name sortKey="Stephens, Gm" uniqKey="Stephens G">GM Stephens</name></author><author><name sortKey="Woo, Pc" uniqKey="Woo P">PC Woo</name></author><author><name sortKey="Zaki, Am" uniqKey="Zaki A">AM Zaki</name></author><author><name sortKey="Zambon, M" uniqKey="Zambon M">M Zambon</name></author><author><name sortKey="Ziebuhr, J" uniqKey="Ziebuhr J">J Ziebuhr</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Eriksson, Kk" uniqKey="Eriksson K">KK Eriksson</name></author><author><name sortKey="Cevantes Barragan, L" uniqKey="Cevantes Barragan L">L Cevantes-Barragan</name></author><author><name sortKey="Ludewig, B" uniqKey="Ludewig B">B Ludewig</name></author><author><name sortKey="Thiel, V" uniqKey="Thiel V">V Thiel</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Fehr, Ar" uniqKey="Fehr A">AR Fehr</name></author><author><name sortKey="Perlman, S" uniqKey="Perlman S">S Perlman</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Guntert, P" uniqKey="Guntert P">P Güntert</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Han, W" uniqKey="Han W">W Han</name></author><author><name sortKey="Li, X" uniqKey="Li X">X Li</name></author><author><name sortKey="Fu, X" uniqKey="Fu X">X Fu</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Han, Hj" uniqKey="Han H">HJ Han</name></author><author><name sortKey="Wen, Hl" uniqKey="Wen H">HL Wen</name></author><author><name sortKey="Zhou, Cm" uniqKey="Zhou C">CM Zhou</name></author><author><name sortKey="Chen, Ff" uniqKey="Chen F">FF Chen</name></author><author><name sortKey="Luo, Lm" uniqKey="Luo L">LM Luo</name></author><author><name sortKey="Liu, Jw" uniqKey="Liu J">JW Liu</name></author><author><name sortKey="Yu, Xj" uniqKey="Yu X">XJ Yu</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Hurst Hess, Kr" uniqKey="Hurst Hess K">KR Hurst-Hess</name></author><author><name sortKey="Kuo, L" uniqKey="Kuo L">L Kuo</name></author><author><name sortKey="Masters, Ps" uniqKey="Masters P">PS Masters</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Karras, Gi" uniqKey="Karras G">GI Karras</name></author><author><name sortKey="Kustatscher, G" uniqKey="Kustatscher G">G Kustatscher</name></author><author><name sortKey="Buhecha, Hr" uniqKey="Buhecha H">HR Buhecha</name></author><author><name sortKey="Allen, Md" uniqKey="Allen M">MD Allen</name></author><author><name sortKey="Pugieux, C" uniqKey="Pugieux C">C Pugieux</name></author><author><name sortKey="Sait, F" uniqKey="Sait F">F Sait</name></author><author><name sortKey="Bycroft, M" uniqKey="Bycroft M">M Bycroft</name></author><author><name sortKey="Ladurner, Ag" uniqKey="Ladurner A">AG Ladurner</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Markley, Jl" uniqKey="Markley J">JL Markley</name></author><author><name sortKey="Bax, A" uniqKey="Bax A">A Bax</name></author><author><name sortKey="Arata, Y" uniqKey="Arata Y">Y Arata</name></author><author><name sortKey="Hilbers, Cw" uniqKey="Hilbers C">CW Hilbers</name></author><author><name sortKey="Kaptein, R" uniqKey="Kaptein R">R Kaptein</name></author><author><name sortKey="Sykes, Bd" uniqKey="Sykes B">BD Sykes</name></author><author><name sortKey="Wright, Pe" uniqKey="Wright P">PE Wright</name></author><author><name sortKey="Wuthrich, K" uniqKey="Wuthrich K">K Wuthrich</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Neuvonen, M" uniqKey="Neuvonen M">M Neuvonen</name></author><author><name sortKey="Ahola, T" uniqKey="Ahola T">T Ahola</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Parvez, Mk" uniqKey="Parvez M">MK Parvez</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Putic, A" uniqKey="Putic A">A Putic</name></author><author><name sortKey="Filipowicz, W" uniqKey="Filipowicz W">W Filipowicz</name></author><author><name sortKey="Hall, J" uniqKey="Hall J">J Hall</name></author><author><name sortKey="Gorvalenva, Ae" uniqKey="Gorvalenva A">AE Gorvalenva</name></author><author><name sortKey="Ziebuhr, J" uniqKey="Ziebuhr J">J Ziebuhr</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Sattler, M" uniqKey="Sattler M">M 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<front><journal-meta><journal-id journal-id-type="nlm-ta">Biomol NMR Assign</journal-id><journal-id journal-id-type="iso-abbrev">Biomol NMR Assign</journal-id><journal-title-group><journal-title>Biomolecular Nmr Assignments</journal-title></journal-title-group><issn pub-type="ppub">1874-2718</issn><issn pub-type="epub">1874-270X</issn><publisher><publisher-name>Springer Netherlands</publisher-name><publisher-loc>Dordrecht</publisher-loc></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">26993639</article-id><article-id pub-id-type="pmc">7091117</article-id><article-id pub-id-type="publisher-id">9676</article-id><article-id pub-id-type="doi">10.1007/s12104-016-9676-9</article-id><article-categories><subj-group subj-group-type="heading"><subject>Article</subject></subj-group></article-categories><title-group><article-title>NMR assignments of the macro domain from Middle East respiratory syndrome coronavirus (MERS-CoV)</article-title></title-group><contrib-group><contrib contrib-type="author" equal-contrib="yes"><name><surname>Huang</surname><given-names>Yi-Ping</given-names></name><xref ref-type="aff" rid="Aff1">1</xref></contrib><contrib contrib-type="author" equal-contrib="yes"><name><surname>Cho</surname><given-names>Chao-Cheng</given-names></name><xref ref-type="aff" rid="Aff2">2</xref><xref ref-type="aff" rid="Aff3">3</xref></contrib><contrib contrib-type="author" corresp="yes"><name><surname>Chang</surname><given-names>Chi-Fon</given-names></name><address><phone>+886-2-2787-1236</phone><email>chifon@gate.sinica.edu.tw</email></address><xref ref-type="aff" rid="Aff1">1</xref></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">http://orcid.org/0000-0002-0008-7383</contrib-id><name><surname>Hsu</surname><given-names>Chun-Hua</given-names></name><address><phone>+886-2-3366-4468</phone><email>andyhsu@ntu.edu.tw</email></address><xref ref-type="aff" rid="Aff2">2</xref><xref ref-type="aff" rid="Aff3">3</xref><xref ref-type="aff" rid="Aff4">4</xref></contrib><aff id="Aff1"><label>1</label><institution-wrap><institution-id institution-id-type="GRID">grid.28665.3f</institution-id><institution-id institution-id-type="ISNI">0000000122871366</institution-id><institution>Genomics Research Center,</institution><institution>Academia Sinica,</institution></institution-wrap>Nankang, Taipei, 11529 Taiwan</aff><aff id="Aff2"><label>2</label><institution-wrap><institution-id institution-id-type="GRID">grid.19188.39</institution-id><institution-id institution-id-type="ISNI">0000000405460241</institution-id><institution>Genome and Systems Biology Degree Program,</institution><institution>National Taiwan University and Academia Sinica,</institution></institution-wrap>Taipei, 10617 Taiwan</aff><aff id="Aff3"><label>3</label><institution-wrap><institution-id institution-id-type="GRID">grid.19188.39</institution-id><institution-id institution-id-type="ISNI">0000000405460241</institution-id><institution>Department of Agricultural Chemistry,</institution><institution>National Taiwan University,</institution></institution-wrap>Taipei, 10617 Taiwan</aff><aff id="Aff4"><label>4</label><institution-wrap><institution-id institution-id-type="GRID">grid.19188.39</institution-id><institution-id institution-id-type="ISNI">0000000405460241</institution-id><institution>Center for Systems Biology,</institution><institution>National Taiwan University,</institution></institution-wrap>Taipei, 10617 Taiwan</aff></contrib-group><pub-date pub-type="epub"><day>18</day><month>3</month><year>2016</year></pub-date><pub-date pub-type="ppub"><year>2016</year></pub-date><volume>10</volume><issue>2</issue><fpage>245</fpage><lpage>248</lpage><history><date date-type="received"><day>28</day><month>12</month><year>2015</year></date><date date-type="accepted"><day>11</day><month>3</month><year>2016</year></date></history><permissions><copyright-statement>© Springer Science+Business Media Dordrecht 2016</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 id="Par1">The newly emerging human pathogen, Middle East respiratory syndrome coronavirus (MERS-CoV), contains a macro domain in the highly conserved N-terminal region of non-structural protein 3. Intense research has shown that macro domains bind ADP-ribose and other derivatives, but it still remains intangible about their exact function. In this study we report the preliminary structural analysis through solution NMR spectroscopy of the MERS-CoV macro domain. The near complete NMR assignments of MERS-CoV macro domain provide the basis for subsequent structural and biochemical investigation in the context of protein function.</p></abstract><kwd-group xml:lang="en"><title>Keywords</title><kwd>Viral macro domain</kwd><kwd>MERS-CoV</kwd><kwd>ADP-ribose binding module</kwd><kwd>NMR spectroscopy</kwd></kwd-group><funding-group><award-group><funding-source><institution-wrap><institution-id institution-id-type="FundRef">http://dx.doi.org/10.13039/501100004663</institution-id><institution>Ministry of Science and Technology, Taiwan</institution></institution-wrap></funding-source><award-id>103-2113-M-002-009-MY2</award-id><principal-award-recipient><name><surname>Hsu</surname><given-names>Chun-Hua</given-names></name></principal-award-recipient></award-group></funding-group><funding-group><award-group><funding-source><institution-wrap><institution-id institution-id-type="FundRef">http://dx.doi.org/10.13039/501100006477</institution-id><institution>National Taiwan University</institution></institution-wrap></funding-source><award-id>NTU-ERP-104R8600</award-id><award-id>NTU-ICRP-104R7560-5</award-id><principal-award-recipient><name><surname>Hsu</surname><given-names>Chun-Hua</given-names></name></principal-award-recipient></award-group></funding-group><custom-meta-group><custom-meta><meta-name>issue-copyright-statement</meta-name><meta-value>© Springer Science+Business Media Dordrecht 2016</meta-value></custom-meta></custom-meta-group></article-meta></front><body><sec id="Sec1"><title>Biological context</title><p id="Par2">Middle East Respiratory Syndrome Coronavirus (MERS-CoV) is a kind of RNA virus belongs to genus <italic>Betacoronavirus</italic> in <italic>Coronavaridae</italic> of enveloped, which is a positive-stranded RNA virus and becomes a novel zoonotic virus in recent years (de Groot et al. <xref ref-type="bibr" rid="CR2">2013</xref>). MERS-CoV transmits respiratory tract infections from original host, the bat, to human by intermediate host, the camel, although the detail viral life cycle is still unknown (Han et al. <xref ref-type="bibr" rid="CR9">2015</xref>). The first human case was reported in 2012 at Saudi Arabia about a man with severe acute pneumonia and secondary renal failure (Zaki et al. <xref ref-type="bibr" rid="CR19">2012</xref>). Through close contact to infected patients, the human-to-human transmissions of the MERS-CoV virus spread very quickly. Since the first report, MERS-CoV has caused over 400 people died among 1100 cases in 23 countries. There is no efficient specific drug to use for this newly human pathogen yet, thus drug development is the emergency to conquer for the researchers.</p><p id="Par3">The macro domain, a protein module specifically recognizing ADP ribose (Karras et al. <xref ref-type="bibr" rid="CR11">2005</xref>), could be found in many bacteria, archaea, plants and vertebrates, suggesting their evolutionary conservation (Han et al. <xref ref-type="bibr" rid="CR8">2011</xref>). Some macro domains possess phosphatase activity towards ADP-ribose-1″-phosphate, which is a by-product of tRNA splicing (Putic et al. <xref ref-type="bibr" rid="CR15">2005</xref>). Furthermore, macro domains are found in nonstructural proteins (nsps) of several positive-strand RNA viruses, including alphaviruses, rubella virus, hepatitis E virus (HEV), and coronaviruses [including the severe acute respiratory syndrome coronavirus (SARS-CoV) and Middle East Respiratory Syndrome Coronavirus (MERS-CoV)] (Neuvonen and Ahola <xref ref-type="bibr" rid="CR13">2009</xref>; Fehr and Perlman <xref ref-type="bibr" rid="CR5">2015</xref>; Hurst-Hess et al. <xref ref-type="bibr" rid="CR10">2015</xref>; Parvez <xref ref-type="bibr" rid="CR14">2015</xref>). The enzymatic activity of viral macro domain is dispensable for coronavirus replication (Putic et al. <xref ref-type="bibr" rid="CR15">2005</xref>), however it seems to play a role in the pathogenesis of mouse hepatitis virus infection (Eriksson et al. <xref ref-type="bibr" rid="CR4">2008</xref>). In addition, viral macro proteins may act via ADP-ribose binding to influence the cellular macro domains-regulated pathways either to promote virus replication or to inhibit host responses directed against the virus (Neuvonen and Ahola <xref ref-type="bibr" rid="CR13">2009</xref>). Since the biochemical, enzymatic and structural analysis of MERS-CoV macro domain is not available, here we present the expression, purification, and chemical shift assignments of MERS-CoV macro domain. The NMR chemical shift assignments serve as the basis for further structural characterization and ligand screening, which provides insight into the conformational properties of this domain in solution and contributes to understanding its function.</p></sec><sec id="Sec2"><title>Materials and methods</title><sec id="Sec3"><title>Cloning, overexpression and purification of MERS-CoV macro domain</title><p id="Par4">The DNA sequence containing MERS-CoV macro domain (aa. 1110–1274) was chemical synthesized and cloned into NdeI/XhoI site of pET-28a(+) vector system (Novagen). Bacteria of the <italic>E. coli</italic> strain BL21(DE3) transformed with the pET-28a (+)-macro domain plasmid were grown in 4 liters of LB medium at 37 °C until the absorbance at 600 nm reach 1.0. Cells were then harvested by centrifugation and re-suspended in one liter of M9 minimal medium supplemented with 1 g/L of <sup>15</sup>NH<sub>4</sub>Cl and 2 g/L of <sup>13</sup>C<sub>6</sub>-glucose (Cambridge Isotope Laboratories). Isopropyl-β-D-thiogalactoside (IPTG, 1 mM) was added to induce His-tagged protein for 20 h at 16 °C. The cell culture was harvested by centrifugation at 6000 rpm.</p><p id="Par5">For purification, cell pellets were re-suspended in 50 mL buffer containing 25 mM sodium phosphate and 100 mM NaCl at pH 7.0 and then disrupted by sonication for 20 min. The cell extract was clarified by centrifugation at 12,500 rpm for 30 min at 4 °C to remove debris. The supernatant was then applied to Ni–NTA column (GE, Healthcare) equilibrated with the same re-suspension buffer, and His-tagged protein was eluted with 200 mM imidazole. The purified His-tagged macro domain was then digested with thrombin for 6 h at 16 °C to remove the His-tag. Finally, macro domain protein (20 kDa, with extra Gly, Ser, His and Met at N-terminus) was purified using size-exclusion Superdex75 XK 16/60 column (GE, Healthcare). The purified protein was concentrated to 0.1–0.5 mM in 20 mM sodium phosphate (pH 6.5) and 100 mM or 150 mM NaCl for NMR structural studies.</p></sec><sec id="Sec4"><title>NMR spectroscopy</title><p id="Par6">All NMR experiments were carried out at 293 K on Bruker Avance 600 MHz NMR or 800 MHz spectrometers equipped with 5 mm triple resonance cryoprobe and Z-gradient. The data was acquired and processed using the software Topspin2.1 (Bruker, Germany) and further analyzed using Sparky, version 3.114 (T. D. Goddard and D. G. Kneller, SPARKY 3, University of California, San Francisco), following the procedures as described previously (Yang et al. <xref ref-type="bibr" rid="CR18">2010</xref>; Chen et al. <xref ref-type="bibr" rid="CR1">2014</xref>). <sup>1</sup>H chemical shifts were externally referenced to 0 ppm of 2,2-dimethyl-2-silapentane-5-sulfonate, whereas <sup>13</sup>C and <sup>15</sup>N chemical shifts were indirectly referenced according to IUPAC recommendations (Markley et al. <xref ref-type="bibr" rid="CR12">1998</xref>). Protein backbone resonance assignments were based on standard triple resonance experiments (Sattler et al. <xref ref-type="bibr" rid="CR16">1999</xref>): HNCACB, CBCA(CO)NH, HNCO and HN(CA)CO. Aliphatic side-chain assignments were primarily done by HCCH-TOCSY and HCCH-COSY with the help of HCC(CO)NH and HBHA(CO)NH experiments.</p></sec></sec><sec id="Sec5"><title>Assignments extension and data deposition</title><p id="Par7">The backbone resonance assignments were nearly complete. Figure <xref rid="Fig1" ref-type="fig">1</xref> illustrates the 2D (<sup>1</sup>H–<sup>15</sup>N) HSQC spectrum and assignments of the amide resonances. Except for the first four amino acids on N-terminal (Gly<sup>−2</sup>, Ser<sup>−1</sup>, His<sup>0</sup> and Met<sup>1</sup>) and six Proline residues (Pro<sup>1</sup>, Pro<sup>72</sup>, Pro<sup>96</sup>, Pro<sup>118</sup>, Pro<sup>123</sup>, and Pro<sup>134</sup>), amides of all other residues (158 out of 168) have been assigned under the experimental conditions (pH 6.5 at 293 K). Among these 158 residues, all other backbone resonances (<sup>1</sup>Hα, <sup>13</sup>Cα, <sup>13</sup>Cβ and <sup>13</sup>C) are 100 % completed. Completeness of <sup>1</sup>H resonances assignment, including side-chain, calculated by CYANA3.9 (Güntert <xref ref-type="bibr" rid="CR7">2004</xref>) is 86.7 %. Secondary structure elements of MERS-CoV macro domain were identified by calculating the chemical shift deviations of the Cα(ΔδCα) and Cβ(ΔδCβ) from the random coil values and was corroborated by analysis of the chemical shift data using the program TALOS+ (Shen et al. <xref ref-type="bibr" rid="CR17">2009</xref>). Positive and negative values of the difference between ΔδCα and ΔδCβ correspond to α-helix and β-sheet secondary structure, respectively and correlated well with TALOS+ index (Fig. <xref rid="Fig2" ref-type="fig">2</xref>). Six helices and seven β-strands could be deduced for MERS-CoV moacro domain protein based on the secondary chemical shift analysis, which results in residues 21–27, 47–54. 58–69, 104–115, 134–144 and 156–162 in α-helices and residues 5–10, 14–19, 32–25, 78–81, 89–94, 119–124 and 148–153 forming β-sheets. The resonance assignments have been deposited to the BioMagResBank (<ext-link ext-link-type="uri" xlink:href="http://www.bmrb.wisc.edu/">http://www.bmrb.wisc.edu/</ext-link>) under the accession number 26657.<fig id="Fig1"><label>Fig. 1</label><caption><p><sup>1</sup>H–<sup>15</sup>N HSQC spectrum of MERS-CoV macro domain recorded at 600 MHz cryogenic-probe with pH 6.5 at 293 K. Backbone amide <sup>1</sup>H and <sup>15</sup>N cross peaks are shown. The <italic>horizontal lines</italic> connect pairs of the side chain NH<sub>2</sub> protons from Asn and Gln residues</p></caption><graphic xlink:href="12104_2016_9676_Fig1_HTML" id="MO1"></graphic></fig><fig id="Fig2"><label>Fig. 2</label><caption><p>The secondary structure assessment of MERS-CoV macro domain. <italic>Upper</italic> The parameter ∆δCα − ∆δCβ shows the deviation of Cα and Cβ experimental values from the corresponding random coil values. Positive and negative values suggest α-helix and β-strand structure, respectively. <italic>Middle</italic> Chemical shift analysis resulting secondary structure elements of macro domain are represented. <italic>Lower</italic> TALOS+ index shows the prediction of secondary structure distribution based on backbone N, H, Cα, Hα, C and side chain Cβ chemical shift values. Negative and positive values suggest α-helix (in <italic>red</italic>) and β-strand (in <italic>blue</italic>) structure, respectively</p></caption><graphic xlink:href="12104_2016_9676_Fig2_HTML" id="MO2"></graphic></fig></p></sec></body><back><fn-group><fn><p>Yi-Ping Huang and Chao-Cheng Cho have contributed equally to this work.</p></fn></fn-group><ack><p>The NMR spectra were obtained at High-Field Nuclear Magnetic Resonance Center (HF-NMRC) and GRC NMR Core Facility in Academia Sinica, Taiwan. This work was supported by the Ministry of Science and Technology, Taiwan (103-2113-M-002-009-MY2), and National Taiwan University (NTU-ERP-104R8600 and NTU-ICRP-104R7560-5) to C.-H. 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