{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,22]],"date-time":"2026-10-22T14:24:06Z","timestamp":1792679046004,"version":"4.5.0"},"reference-count":44,"publisher":"Elsevier BV","issue":"2","license":[{"start":{"date-parts":[[2009,10,1]],"date-time":"2009-10-01T00:00:00Z","timestamp":1254355200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2009,10,1]],"date-time":"2009-10-01T00:00:00Z","timestamp":1254355200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2013,8,22]],"date-time":"2013-08-22T00:00:00Z","timestamp":1377129600000},"content-version":"vor","delay-in-days":1421,"URL":"http:\/\/www.elsevier.com\/open-access\/userlicense\/1.0\/"}],"content-domain":{"domain":["clinicalkey.com","clinicalkey.com.au","clinicalkey.es","clinicalkey.fr","clinicalkey.jp","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Virology"],"published-print":{"date-parts":[[2009,10]]},"DOI":"10.1016\/j.virol.2009.07.038","type":"journal-article","created":{"date-parts":[[2009,8,30]],"date-time":"2009-08-30T04:09:23Z","timestamp":1251605363000},"page":"265-271","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":62,"title":["SARS-coronavirus spike S2 domain flanked by cysteine residues C822 and C833 is important for activation of membrane fusion"],"prefix":"10.1016","volume":"393","author":[{"given":"Ikenna G.","family":"Madu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sandrine","family":"Belouzard","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gary R.","family":"Whittaker","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"issue":"5","key":"10.1016\/j.virol.2009.07.038_bib1","doi-asserted-by":"crossref","first-page":"633","DOI":"10.1084\/jem.20021756","article-title":"Infectious hepatitis C virus pseudo-particles containing functional E1\u2013E2 envelope protein complexes","volume":"197","author":"Bartosch","year":"2003","journal-title":"J. Exp. Med."},{"issue":"14","key":"10.1016\/j.virol.2009.07.038_bib2","doi-asserted-by":"crossref","first-page":"5871","DOI":"10.1073\/pnas.0809524106","article-title":"Activation of the SARS coronavirus spike protein via sequential proteolytic cleavage at two distinct sites","volume":"106","author":"Belouzard","year":"2009","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"issue":"3","key":"10.1016\/j.virol.2009.07.038_bib3","doi-asserted-by":"crossref","first-page":"554","DOI":"10.1016\/j.bbrc.2004.11.063","article-title":"Implication of proprotein convertases in the processing and spread of severe acute respiratory syndrome coronavirus","volume":"326","author":"Bergeron","year":"2005","journal-title":"Biochem. Biophys. Res. Commun."},{"issue":"4","key":"10.1016\/j.virol.2009.07.038_bib4","doi-asserted-by":"crossref","first-page":"1210","DOI":"10.1016\/j.bbrc.2004.10.156","article-title":"Prediction of quaternary assembly of SARS coronavirus peplomer","volume":"325","author":"Bernini","year":"2004","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"10.1016\/j.virol.2009.07.038_bib5","series-title":"Nidoviruses","first-page":"157","article-title":"Nidovirus entry into cells","author":"Bosch","year":"2008"},{"issue":"22","key":"10.1016\/j.virol.2009.07.038_bib6","doi-asserted-by":"crossref","first-page":"8455","DOI":"10.1073\/pnas.0400576101","article-title":"Severe acute respiratory syndrome coroavirus (SARS-CoV) infection inhibition using spike protein heptad repeat-derived peptides","volume":"101","author":"Bosch","year":"2004","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"issue":"2","key":"10.1016\/j.virol.2009.07.038_bib7","doi-asserted-by":"crossref","first-page":"1688","DOI":"10.1128\/jvi.71.2.1688-1691.1997","article-title":"Human carcinoembryonic antigen and biliary glycoprotein can serve as mouse hepatitis virus receptors","volume":"71","author":"Chen","year":"1997","journal-title":"J. Virol."},{"issue":"7","key":"10.1016\/j.virol.2009.07.038_bib8","doi-asserted-by":"crossref","first-page":"3180","DOI":"10.1128\/JVI.80.7.3180-3188.2006","article-title":"The avian coronavirus infectious bronchitis virus undergoes direct low-pH-dependent fusion activation during entry into host cells","volume":"80","author":"Chu","year":"2006","journal-title":"J. Virol."},{"issue":"20","key":"10.1016\/j.virol.2009.07.038_bib9","doi-asserted-by":"crossref","first-page":"9738","DOI":"10.1128\/JVI.74.20.9738-9741.2000","article-title":"Critical role for the cysteines flanking the internal fusion peptide of avian sarcoma\/leukosis virus envelope glycoprotein","volume":"74","author":"Delos","year":"2000","journal-title":"J. Virol."},{"issue":"6","key":"10.1016\/j.virol.2009.07.038_bib10","doi-asserted-by":"crossref","first-page":"3131","DOI":"10.1128\/JVI.02266-07","article-title":"Cysteines flanking the internal fusion peptide are required for the avian sarcoma\/leukosis virus glycoprotein to mediate the lipid mixing stage of fusion with high efficiency","volume":"82","author":"Delos","year":"2008","journal-title":"J. Virol."},{"issue":"20","key":"10.1016\/j.virol.2009.07.038_bib11","doi-asserted-by":"crossref","first-page":"1967","DOI":"10.1056\/NEJMoa030747","article-title":"Identification of a novel coronavirus in patients with severe acute respiratory syndrome","volume":"348","author":"Drosten","year":"2003","journal-title":"New Engl. J. Med."},{"issue":"1","key":"10.1016\/j.virol.2009.07.038_bib12","doi-asserted-by":"crossref","first-page":"174","DOI":"10.1016\/j.bbrc.2007.05.092","article-title":"Cleavage of spike protein of SARS coronavirus by protease factor Xa is associated with viral infectivity","volume":"359","author":"Du","year":"2007","journal-title":"Biochem. Biophys. Res. Commun."},{"issue":"2","key":"10.1016\/j.virol.2009.07.038_bib13","doi-asserted-by":"crossref","first-page":"358","DOI":"10.1016\/j.virol.2006.02.003","article-title":"Furin cleavage of the SARS coronavirus spike glycoprotein enhances cell\u2013cell fusion but does not affect virion entry","volume":"350","author":"Follis","year":"2006","journal-title":"Virology"},{"issue":"6937","key":"10.1016\/j.virol.2009.07.038_bib14","doi-asserted-by":"crossref","first-page":"240","DOI":"10.1038\/423240a","article-title":"Aetiology \u2014 Koch's postulates fulfilled for SARS virus","volume":"423","author":"Fouchier","year":"2003","journal-title":"Nature"},{"issue":"21","key":"10.1016\/j.virol.2009.07.038_bib15","doi-asserted-by":"crossref","first-page":"12029","DOI":"10.1128\/JVI.00315-07","article-title":"Specific asparagine-linked glycosylation sites are critical for DC-SIGN- and L-SIGN-mediated severe acute respiratory syndrome coronavirus entry","volume":"81","author":"Han","year":"2007","journal-title":"J. Virol."},{"key":"10.1016\/j.virol.2009.07.038_bib16","first-page":"43","article-title":"Human biliary glycoproteins function as receptors for interspecies transfer of Mouse Hepatitis virus","volume":"Vol. 440","author":"Hensley","year":"1998"},{"issue":"17","key":"10.1016\/j.virol.2009.07.038_bib17","doi-asserted-by":"crossref","first-page":"8639","DOI":"10.1128\/JVI.00560-06","article-title":"Highly conserved regions within the spike proteins of human coronaviruses 229E and NL63 determine recognition of their respective cellular receptors","volume":"80","author":"Hofmann","year":"2006","journal-title":"J. Virol."},{"issue":"20","key":"10.1016\/j.virol.2009.07.038_bib18","doi-asserted-by":"crossref","first-page":"1948","DOI":"10.1056\/NEJMp030078","article-title":"SARS-associated coronavirus","volume":"348","author":"Holmes","year":"2003","journal-title":"New Engl. J. Med."},{"issue":"6","key":"10.1016\/j.virol.2009.07.038_bib19","doi-asserted-by":"crossref","first-page":"3198","DOI":"10.1074\/jbc.M508381200","article-title":"SARS coronavirus, but not human coronavirus NL63, utilizes cathepsin L to infect ACE2-expressing cells","volume":"281","author":"Huang","year":"2006","journal-title":"J. Biol. Chem."},{"issue":"2","key":"10.1016\/j.virol.2009.07.038_bib20","doi-asserted-by":"crossref","first-page":"366","DOI":"10.2307\/1592976","article-title":"Spike glycoprotein cleavage recognition site analysis of infectious bronchitis virus","volume":"45","author":"Jackwood","year":"2001","journal-title":"Avian Dis."},{"issue":"2","key":"10.1016\/j.virol.2009.07.038_bib21","doi-asserted-by":"crossref","first-page":"712","DOI":"10.1128\/JVI.01933-08","article-title":"Protease-mediated entry via the endosome of human coronavirus 229E","volume":"83","author":"Kawase","year":"2009","journal-title":"J. Virol."},{"issue":"6686","key":"10.1016\/j.virol.2009.07.038_bib22","doi-asserted-by":"crossref","first-page":"648","DOI":"10.1038\/31405","article-title":"Structure of an HIV gp120 envelope glycoprotein in complex with the CD4 receptor and a neutralizing human antibody","volume":"393","author":"Kwong","year":"1998","journal-title":"Nature"},{"issue":"22","key":"10.1016\/j.virol.2009.07.038_bib23","doi-asserted-by":"crossref","first-page":"19727","DOI":"10.1074\/jbc.M201837200","article-title":"Quaternary structure of coronavirus spikes in complex with carcinoembryonic antigen-related cell adhesion molecule cellular receptors","volume":"277","author":"Lewicki","year":"2002","journal-title":"J. Biol. Chem."},{"issue":"6965","key":"10.1016\/j.virol.2009.07.038_bib24","doi-asserted-by":"crossref","first-page":"450","DOI":"10.1038\/nature02145","article-title":"Angiotensin-converting enzyme 2 is a functional receptor for the SARS coronavirus","volume":"426","author":"Li","year":"2003","journal-title":"Nature"},{"issue":"2","key":"10.1016\/j.virol.2009.07.038_bib25","doi-asserted-by":"crossref","first-page":"367","DOI":"10.1016\/j.virol.2007.04.035","article-title":"The S proteins of human coronavirus NL63 and severe acute respiratory syndrome coronavirus bind overlapping regions of ACE2","volume":"367","author":"Li","year":"2007","journal-title":"Virology"},{"issue":"15","key":"10.1016\/j.virol.2009.07.038_bib26","doi-asserted-by":"crossref","first-page":"7411","DOI":"10.1128\/JVI.00079-09","article-title":"Characterization of a highly conserved domain within the severe acute respiratory syndrome coronavirus spike protein S2 domain with characteristics of a viral fusion peptide","volume":"83","author":"Madu","year":"2009","journal-title":"J. Virol."},{"issue":"1","key":"10.1016\/j.virol.2009.07.038_bib27","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1089\/152308603321223621","article-title":"Redox control on the cell surface: implications for HIV-1 entry","volume":"5","author":"Matthias","year":"2003","journal-title":"Antioxid. Redox Signal."},{"issue":"8","key":"10.1016\/j.virol.2009.07.038_bib28","doi-asserted-by":"crossref","first-page":"727","DOI":"10.1038\/ni815","article-title":"Disulfide exchange in domain 2 of CD4 is required for entry of HIV-I","volume":"3","author":"Matthias","year":"2002","journal-title":"Nat. Immunol."},{"issue":"6","key":"10.1016\/j.virol.2009.07.038_bib29","doi-asserted-by":"crossref","first-page":"2584","DOI":"10.1128\/JVI.02158-08","article-title":"Role of conserved cysteines in the alphavirus E3 protein","volume":"83","author":"Parrott","year":"2009","journal-title":"J. Virol."},{"issue":"2","key":"10.1016\/j.virol.2009.07.038_bib30","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1016\/j.virol.2005.06.046","article-title":"Genetic analysis of the SARS-coronavirus spike glycoprotein functional domains involved in cell-surface expression and cell-to-cell fusion","volume":"341","author":"Petit","year":"2005","journal-title":"Virology"},{"issue":"2","key":"10.1016\/j.virol.2009.07.038_bib31","doi-asserted-by":"crossref","first-page":"264","DOI":"10.1016\/j.virol.2006.10.034","article-title":"Palmitoylation of the cysteine-rich endodomain of the SARS-coronavirus spike glycoprotein is important for spike-mediated cell fusion","volume":"360","author":"Petit","year":"2007","journal-title":"Virology"},{"issue":"2","key":"10.1016\/j.virol.2009.07.038_bib32","doi-asserted-by":"crossref","first-page":"683","DOI":"10.1128\/JVI.78.2.683-691.2004","article-title":"The spacing between cysteines two and three of the LDL-A module of Tva is important for subgroup A avian sarcoma and leukosis virus entry","volume":"78","author":"Rai","year":"2004","journal-title":"J. Virol."},{"issue":"6","key":"10.1016\/j.virol.2009.07.038_bib33","doi-asserted-by":"crossref","first-page":"361","DOI":"10.1080\/09687860400017784","article-title":"Class I and class II viral fusion protein structures reveal similar principles in membrane fusion (Review)","volume":"21","author":"Schibli","year":"2004","journal-title":"Mol. Membr. Biol."},{"issue":"8","key":"10.1016\/j.virol.2009.07.038_bib34","doi-asserted-by":"crossref","first-page":"5634","DOI":"10.1128\/jvi.70.8.5634-5637.1996","article-title":"Transmissible gastroenteritis coronavirus, but not the related porcine respiratory coronavirus, has a sialic acid (N-glycolylneuraminic acid) binding activity","volume":"70","author":"Schultze","year":"1996","journal-title":"J. Virol."},{"issue":"19","key":"10.1016\/j.virol.2009.07.038_bib35","doi-asserted-by":"crossref","first-page":"10328","DOI":"10.1128\/JVI.78.19.10328-10335.2004","article-title":"Synthesis and characterization of a native, oligomeric form of recombinant severe acute respiratory syndrome coronavirus spike glycoprotein","volume":"78","author":"Song","year":"2004","journal-title":"J. Virol."},{"issue":"1","key":"10.1016\/j.virol.2009.07.038_bib36","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1016\/j.bbrc.2003.08.122","article-title":"Molecular modelling of S1 and S2 subunits of SARS coronavirus spike glycoprotein","volume":"310","author":"Spiga","year":"2003","journal-title":"Biochem. Biophys. Res. Commun."},{"issue":"6","key":"10.1016\/j.virol.2009.07.038_bib37","doi-asserted-by":"crossref","first-page":"2682","DOI":"10.1128\/JVI.78.6.2682-2692.2004","article-title":"Requirements for CEACAMs and cholesterol during murine coronavirus cell entry","volume":"78","author":"Thorp","year":"2004","journal-title":"J. Virol."},{"issue":"3","key":"10.1016\/j.virol.2009.07.038_bib38","doi-asserted-by":"crossref","first-page":"1280","DOI":"10.1128\/JVI.80.3.1280-1289.2006","article-title":"Palmitoylations on murine coronavirus spike proteins are essential for virion assembly and infectivity","volume":"80","author":"Thorp","year":"2006","journal-title":"J. Virol."},{"issue":"23","key":"10.1016\/j.virol.2009.07.038_bib39","doi-asserted-by":"crossref","first-page":"11985","DOI":"10.1128\/JVI.01412-08","article-title":"Entry from the cell surface of severe acute respiratory syndrome coronavirus with cleaved S protein as revealed by pseudotype virus bearing cleaved S protein","volume":"82","author":"Watanabe","year":"2008","journal-title":"J. Virol."},{"issue":"20","key":"10.1016\/j.virol.2009.07.038_bib40","doi-asserted-by":"crossref","first-page":"9741","DOI":"10.1128\/JVI.75.20.9741-9752.2001","article-title":"Molecular determinants of species specificity in the coronavirus receptor aminopeptidase N (CD13): influence of N-linked glycosylation","volume":"75","author":"Wentworth","year":"2001","journal-title":"J. Virol."},{"issue":"4","key":"10.1016\/j.virol.2009.07.038_bib41","doi-asserted-by":"crossref","first-page":"1159","DOI":"10.1016\/j.bbrc.2003.11.054","article-title":"The SARS-CoV S glycoprotein: expression and functional characterization","volume":"312","author":"Xiao","year":"2003","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"10.1016\/j.virol.2009.07.038_bib42","first-page":"89","article-title":"Requirement of proteolytic cleavage of the murine coronavirus MHV-2 spike protein for fusion activity","volume":"Vol. 440","author":"Yamada","year":"1998"},{"issue":"18","key":"10.1016\/j.virol.2009.07.038_bib43","doi-asserted-by":"crossref","first-page":"9904","DOI":"10.1128\/JVI.78.18.9904-9917.2004","article-title":"Genetic analysis of determinants for spike glycoprotein assembly into murine coronavirus virions: distinct roles for charge-rich and cysteine-rich regions of the endodomain","volume":"78","author":"Ye","year":"2004","journal-title":"J. Virol."},{"issue":"13","key":"10.1016\/j.virol.2009.07.038_bib44","doi-asserted-by":"crossref","first-page":"6938","DOI":"10.1128\/JVI.78.13.6938-6945.2004","article-title":"Identification of an antigenic determinant on the S2 domain of the severe acute respiratory syndrome coronavirus spike glycoprotein capable of inducing neutralizing antibodies","volume":"78","author":"Zhang","year":"2004","journal-title":"J. Virol."}],"container-title":["Virology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0042682209004759?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0042682209004759?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2025,9,23]],"date-time":"2025-09-23T00:02:02Z","timestamp":1758585722000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0042682209004759"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2009,10]]},"references-count":44,"journal-issue":{"issue":"2","published-print":{"date-parts":[[2009,10]]}},"alternative-id":["S0042682209004759"],"URL":"https:\/\/doi.org\/10.1016\/j.virol.2009.07.038","relation":{},"ISSN":["0042-6822"],"issn-type":[{"value":"0042-6822","type":"print"}],"subject":[],"published":{"date-parts":[[2009,10]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"SARS-coronavirus spike S2 domain flanked by cysteine residues C822 and C833 is important for activation of membrane fusion","name":"articletitle","label":"Article Title"},{"value":"Virology","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.virol.2009.07.038","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"Copyright \u00a9 2009 Elsevier Inc. All rights reserved.","name":"copyright","label":"Copyright"}]}}