{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T18:46:49Z","timestamp":1790794009593,"version":"4.1.0"},"reference-count":52,"publisher":"Elsevier BV","issue":"3","license":[{"start":{"date-parts":[[2022,3,1]],"date-time":"2022-03-01T00:00:00Z","timestamp":1646092800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2022,3,1]],"date-time":"2022-03-01T00:00:00Z","timestamp":1646092800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2022,2,8]],"date-time":"2022-02-08T00:00:00Z","timestamp":1644278400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["jbc.org","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Journal of Biological Chemistry"],"published-print":{"date-parts":[[2022,3]]},"DOI":"10.1016\/j.jbc.2022.101710","type":"journal-article","created":{"date-parts":[[2022,2,9]],"date-time":"2022-02-09T20:56:31Z","timestamp":1644440191000},"page":"101710","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":67,"title":["Integrin mediates cell entry of the SARS-CoV-2 virus independent of cellular receptor ACE2"],"prefix":"10.1016","volume":"298","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1883-9165","authenticated-orcid":false,"given":"Jiamnin","family":"Liu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fan","family":"Lu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6295-7068","authenticated-orcid":false,"given":"Yinghua","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Edward","family":"Plow","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jun","family":"Qin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.jbc.2022.101710_bib1","doi-asserted-by":"crossref","first-page":"270","DOI":"10.1038\/s41586-020-2012-7","article-title":"A\u00a0pneumonia outbreak associated with a new coronavirus of probable bat origin","volume":"579","author":"Zhou","year":"2020","journal-title":"Nature"},{"key":"10.1016\/j.jbc.2022.101710_bib2","article-title":"Cryo-EM structure of the 2019-nCoV spike in the prefusion conformation","author":"Wrapp","year":"2020","journal-title":"bioRxiv"},{"key":"10.1016\/j.jbc.2022.101710_bib3","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1038\/s41586-020-2180-5","article-title":"Structure of the SARS-CoV-2 spike receptor-binding domain bound to the ACE2 receptor","volume":"581","author":"Lan","year":"2020","journal-title":"Nature"},{"key":"10.1016\/j.jbc.2022.101710_bib4","doi-asserted-by":"crossref","first-page":"894","DOI":"10.1016\/j.cell.2020.03.045","article-title":"Structural and functional basis of SARS-CoV-2 entry by using human ACE2","volume":"181","author":"Wang","year":"2020","journal-title":"Cell"},{"key":"10.1016\/j.jbc.2022.101710_bib5","doi-asserted-by":"crossref","first-page":"E1","DOI":"10.1161\/01.RES.87.5.e1","article-title":"A\u00a0novel angiotensin-converting enzyme-related carboxypeptidase (ACE2) converts angiotensin I to angiotensin 1-9","volume":"87","author":"Donoghue","year":"2000","journal-title":"Circ. Res."},{"key":"10.1016\/j.jbc.2022.101710_bib6","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"},{"key":"10.1016\/j.jbc.2022.101710_bib7","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1016\/j.cell.2020.02.052","article-title":"SARS-CoV-2 cell entry depends on ACE2 and TMPRSS2 and is blocked by a clinically proven protease inhibitor","volume":"181","author":"Hoffmann","year":"2020","journal-title":"Cell"},{"key":"10.1016\/j.jbc.2022.101710_bib8","doi-asserted-by":"crossref","first-page":"404","DOI":"10.1038\/s41590-021-00901-9","article-title":"Lessons learned: New insights on the role of cytokines in COVID-19","volume":"22","author":"Buszko","year":"2021","journal-title":"Nat. Immunol."},{"key":"10.1016\/j.jbc.2022.101710_bib9","doi-asserted-by":"crossref","first-page":"1200","DOI":"10.1038\/s41591-020-0965-6","article-title":"Clinical and immunological assessment of asymptomatic SARS-CoV-2 infections","volume":"26","author":"Long","year":"2020","journal-title":"Nat. Med."},{"key":"10.1016\/j.jbc.2022.101710_bib10","article-title":"Cytokine storm. Reply","volume":"384","author":"Fajgenbaum","year":"2021","journal-title":"N.\u00a0Engl. J. Med."},{"key":"10.1016\/j.jbc.2022.101710_bib11","doi-asserted-by":"crossref","DOI":"10.1128\/JVI.01511-21","article-title":"SARS-CoV-2 causes lung infection without severe disease in human ACE2 knock-in mice","volume":"96","author":"Winkler","year":"2022","journal-title":"J.\u00a0Virol."},{"key":"10.1016\/j.jbc.2022.101710_bib12","doi-asserted-by":"crossref","first-page":"100847","DOI":"10.1016\/j.jbc.2021.100847","article-title":"Role of host factors in SARS-CoV-2 entry","volume":"297","author":"Evans","year":"2021","journal-title":"J.\u00a0Biol. Chem."},{"key":"10.1016\/j.jbc.2022.101710_bib13","doi-asserted-by":"crossref","first-page":"124","DOI":"10.1016\/j.chom.2020.05.020","article-title":"A\u00a0mouse model of SARS-CoV-2 infection and pathogenesis","volume":"28","author":"Sun","year":"2020","journal-title":"Cell Host Microbe"},{"key":"10.1016\/j.jbc.2022.101710_bib14","article-title":"Tropism of SARS-CoV-2 for developing human cortical astrocytes","author":"Andrews","year":"2021","journal-title":"bioRxiv"},{"key":"10.1016\/j.jbc.2022.101710_bib15","doi-asserted-by":"crossref","first-page":"1438","DOI":"10.3390\/microorganisms9071438","article-title":"SARS-CoV-2 infection remodels the phenotype and promotes angiogenesis of primary human lung endothelial cells","volume":"9","author":"Caccuri","year":"2021","journal-title":"Microorganisms"},{"key":"10.1016\/j.jbc.2022.101710_bib16","article-title":"SARS-CoV-2 infection and persistence throughout the human body and brain","author":"Chertow","year":"2022","journal-title":"Biol. Sci."},{"key":"10.1016\/j.jbc.2022.101710_bib17","doi-asserted-by":"crossref","first-page":"104759","DOI":"10.1016\/j.antiviral.2020.104759","article-title":"A\u00a0potential role for integrins in host cell entry by SARS-CoV-2","volume":"177","author":"Sigrist","year":"2020","journal-title":"Antiviral Res."},{"key":"10.1016\/j.jbc.2022.101710_bib18","doi-asserted-by":"crossref","first-page":"146","DOI":"10.3390\/v13020146","article-title":"Biological and clinical consequences of integrin binding via a rogue RGD motif in the SARS CoV-2 spike protein","volume":"13","author":"Makowski","year":"2021","journal-title":"Viruses"},{"key":"10.1016\/j.jbc.2022.101710_bib19","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.jacbts.2020.10.003","article-title":"The integrin binding peptide, ATN-161, as a novel therapy for SARS-CoV-2 infection","volume":"6","author":"Beddingfield","year":"2021","journal-title":"JACC Basic Transl. Sci."},{"key":"10.1016\/j.jbc.2022.101710_bib20","doi-asserted-by":"crossref","first-page":"673","DOI":"10.1016\/S0092-8674(02)00971-6","article-title":"Integrins: Bidirectional, allosteric signaling machines","volume":"110","author":"Hynes","year":"2002","journal-title":"Cell"},{"key":"10.1016\/j.jbc.2022.101710_bib21","doi-asserted-by":"crossref","first-page":"2590","DOI":"10.1016\/j.jmb.2018.06.024","article-title":"Virus-receptor interactions: The key to cellular invasion","volume":"430","author":"Maginnis","year":"2018","journal-title":"J.\u00a0Mol. Biol."},{"key":"10.1016\/j.jbc.2022.101710_bib22","doi-asserted-by":"crossref","first-page":"645","DOI":"10.3390\/v13040645","article-title":"The spike glycoprotein of SARS-CoV-2 binds to beta1 integrins expressed on the surface of lung epithelial cells","volume":"13","author":"Park","year":"2021","journal-title":"Viruses"},{"key":"10.1016\/j.jbc.2022.101710_bib23","doi-asserted-by":"crossref","first-page":"496","DOI":"10.1016\/j.jinf.2021.07.007","article-title":"Low incidence rate of diarrhoea in COVID-19 patients is due to integrin","volume":"83","author":"Gao","year":"2021","journal-title":"J.\u00a0Infect."},{"key":"10.1016\/j.jbc.2022.101710_bib24","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0253347","article-title":"SARS-CoV-2 uses major endothelial integrin alphavbeta3 to cause vascular dysregulation in-vitro during COVID-19","volume":"16","author":"Nader","year":"2021","journal-title":"PLoS One"},{"key":"10.1016\/j.jbc.2022.101710_bib25","doi-asserted-by":"crossref","first-page":"912","DOI":"10.3389\/fphar.2020.00912","article-title":"New strategy for COVID-19: An evolutionary role for RGD motif in SARS-CoV-2 and potential inhibitors for virus infection","volume":"11","author":"Yan","year":"2020","journal-title":"Front. Pharmacol."},{"key":"10.1016\/j.jbc.2022.101710_bib26","doi-asserted-by":"crossref","first-page":"20398","DOI":"10.1038\/s41598-021-99893-7","article-title":"Integrin activation is an essential component of SARS-CoV-2 infection","volume":"11","author":"Simons","year":"2021","journal-title":"Sci. Rep."},{"key":"10.1016\/j.jbc.2022.101710_bib27","doi-asserted-by":"crossref","first-page":"318","DOI":"10.1016\/j.jinf.2020.03.046","article-title":"A\u00a0potential inhibitory role for integrin in the receptor targeting of SARS-CoV-2","volume":"81","author":"Luan","year":"2020","journal-title":"J.\u00a0Infect."},{"key":"10.1016\/j.jbc.2022.101710_bib28","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/j.cytogfr.2020.05.003","article-title":"The cytokine storm in COVID-19: An overview of the involvement of the chemokine\/chemokine-receptor system","volume":"53","author":"Coperchini","year":"2020","journal-title":"Cytokine Growth Factor. Rev."},{"key":"10.1016\/j.jbc.2022.101710_bib29","doi-asserted-by":"crossref","first-page":"1619","DOI":"10.1038\/s41390-020-01177-9","article-title":"The role of integrins in inflammation and angiogenesis","volume":"89","author":"Mezu-Ndubuisi","year":"2021","journal-title":"Pediatr. Res."},{"key":"10.1016\/j.jbc.2022.101710_bib30","doi-asserted-by":"crossref","first-page":"26566","DOI":"10.1074\/jbc.M003495200","article-title":"Cytokine signaling through Stat3 activates integrins, promotes adhesion, and induces growth arrest in the myeloid cell line 32D","volume":"275","author":"Wooten","year":"2000","journal-title":"J.\u00a0Biol. Chem."},{"key":"10.1016\/j.jbc.2022.101710_bib31","doi-asserted-by":"crossref","first-page":"18848","DOI":"10.1074\/jbc.M116.773713","article-title":"Pleiotrophin, a multifunctional cytokine and growth factor, induces leukocyte responses through the integrin Mac-1","volume":"292","author":"Shen","year":"2017","journal-title":"J.\u00a0Biol. Chem."},{"key":"10.1016\/j.jbc.2022.101710_bib32","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1038\/s41586-020-2179-y","article-title":"Structural basis of receptor recognition by SARS-CoV-2","volume":"581","author":"Shang","year":"2020","journal-title":"Nature"},{"key":"10.1016\/j.jbc.2022.101710_bib33","doi-asserted-by":"crossref","first-page":"3263","DOI":"10.1242\/jcs.02436","article-title":"A\u00a0novel protein kinase C alpha-dependent signal to ERK1\/2 activated by alphaVbeta3 integrin in osteoclasts and in Chinese hamster ovary (CHO) cells","volume":"118","author":"Rucci","year":"2005","journal-title":"J.\u00a0Cell Sci."},{"key":"10.1016\/j.jbc.2022.101710_bib34","doi-asserted-by":"crossref","first-page":"735","DOI":"10.1038\/nbt.1932","article-title":"The genomic sequence of the Chinese hamster ovary (CHO)-K1 cell line","volume":"29","author":"Xu","year":"2011","journal-title":"Nat. Biotechnol."},{"key":"10.1016\/j.jbc.2022.101710_bib35","doi-asserted-by":"crossref","first-page":"7981","DOI":"10.1074\/jbc.273.14.7981","article-title":"Integrin activation by dithiothreitol or Mn2+ induces a ligand-occupied conformation and exposure of a novel NH2-terminal regulatory site on the beta1 integrin chain","volume":"273","author":"Ni","year":"1998","journal-title":"J.\u00a0Biol. Chem."},{"key":"10.1016\/j.jbc.2022.101710_bib36","doi-asserted-by":"crossref","DOI":"10.1126\/sciadv.abe9716","article-title":"Structural insights into integrin alpha5beta1 opening by fibronectin ligand","volume":"7","author":"Schumacher","year":"2021","journal-title":"Sci. Adv."},{"key":"10.1016\/j.jbc.2022.101710_bib37","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1038\/ni1566","article-title":"HIV-1 envelope protein binds to and signals through integrin alpha4beta7, the gut mucosal homing receptor for peripheral T cells","volume":"9","author":"Arthos","year":"2008","journal-title":"Nat. Immunol."},{"key":"10.1016\/j.jbc.2022.101710_bib38","doi-asserted-by":"crossref","first-page":"6967","DOI":"10.1128\/JVI.78.13.6967-6973.2004","article-title":"Integrin alpha v beta 6 is an RGD-dependent receptor for coxsackievirus A9","volume":"78","author":"Williams","year":"2004","journal-title":"J.\u00a0Virol."},{"key":"10.1016\/j.jbc.2022.101710_bib39","doi-asserted-by":"crossref","first-page":"106","DOI":"10.1016\/j.cell.2020.12.004","article-title":"Genetic screens identify host factors for SARS-CoV-2 and common cold coronaviruses","volume":"184","author":"Wang","year":"2021","journal-title":"Cell"},{"key":"10.1016\/j.jbc.2022.101710_bib40","doi-asserted-by":"crossref","first-page":"92","DOI":"10.1016\/j.cell.2020.10.030","article-title":"Identification of required host factors for SARS-CoV-2 infection in human cells","volume":"184","author":"Daniloski","year":"2021","journal-title":"Cell"},{"key":"10.1016\/j.jbc.2022.101710_bib41","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1016\/j.cell.2020.10.028","article-title":"Genome-wide CRISPR screens reveal host factors critical for SARS-CoV-2 infection","volume":"184","author":"Wei","year":"2021","journal-title":"Cell"},{"key":"10.1016\/j.jbc.2022.101710_bib42","doi-asserted-by":"crossref","first-page":"629","DOI":"10.15252\/embj.201695803","article-title":"Conformational equilibria and intrinsic affinities define integrin activation","volume":"36","author":"Li","year":"2017","journal-title":"EMBO J."},{"key":"10.1016\/j.jbc.2022.101710_bib43","doi-asserted-by":"crossref","first-page":"15408","DOI":"10.1038\/ncomms15408","article-title":"Rules of engagement between alphavbeta6 integrin and foot-and-mouth disease virus","volume":"8","author":"Kotecha","year":"2017","journal-title":"Nat. Commun."},{"key":"10.1016\/j.jbc.2022.101710_bib44","doi-asserted-by":"crossref","first-page":"39805","DOI":"10.1038\/srep39805","article-title":"A\u00a0comprehensive evaluation of the activity and selectivity profile of ligands for RGD-binding integrins","volume":"7","author":"Kapp","year":"2017","journal-title":"Sci. Rep."},{"key":"10.1016\/j.jbc.2022.101710_bib45","doi-asserted-by":"crossref","first-page":"1053","DOI":"10.1128\/JVI.79.2.1053-1061.2005","article-title":"Deletion of penton RGD motifs affects the efficiency of both the internalization and the endosome escape of viral particles containing adenovirus serotype 5 or 35 fiber knobs","volume":"79","author":"Shayakhmetov","year":"2005","journal-title":"J.\u00a0Virol."},{"key":"10.1016\/j.jbc.2022.101710_bib46","doi-asserted-by":"crossref","first-page":"124","DOI":"10.1016\/j.tibs.2020.09.001","article-title":"Integrins control vesicular trafficking; new tricks for old dogs","volume":"46","author":"Nolte","year":"2021","journal-title":"Trends Biochem. Sci."},{"key":"10.1016\/j.jbc.2022.101710_bib47","doi-asserted-by":"crossref","first-page":"610","DOI":"10.1111\/j.1600-0854.2012.01327.x","article-title":"Distinct recycling of active and inactive beta1 integrins","volume":"13","author":"Arjonen","year":"2012","journal-title":"Traffic"},{"key":"10.1016\/j.jbc.2022.101710_bib48","doi-asserted-by":"crossref","first-page":"290","DOI":"10.1038\/cr.2008.15","article-title":"SARS coronavirus entry into host cells through a novel clathrin- and caveolae-independent endocytic pathway","volume":"18","author":"Wang","year":"2008","journal-title":"Cell Res."},{"key":"10.1016\/j.jbc.2022.101710_bib49","doi-asserted-by":"crossref","first-page":"1519","DOI":"10.1021\/acsinfecdis.1c00070","article-title":"Small-molecule inhibitors of the coronavirus spike: ACE2 protein-protein interaction as blockers of viral attachment and entry for SARS-CoV-2","volume":"7","author":"Bojadzic","year":"2021","journal-title":"ACS Infect. Dis."},{"key":"10.1016\/j.jbc.2022.101710_bib50","doi-asserted-by":"crossref","first-page":"600372","DOI":"10.3389\/fphar.2020.600372","article-title":"Methylene blue inhibits the SARS-CoV-2 spike-ACE2 protein-protein interaction-a mechanism that can contribute to its antiviral activity against COVID-19","volume":"11","author":"Bojadzic","year":"2020","journal-title":"Front. Pharmacol."},{"key":"10.1016\/j.jbc.2022.101710_bib51","article-title":"Differential gene expression by RNA-Seq in Sigma-2 receptor\/TMEM97 knockout cells reveals its role in complement activation and SARS-CoV-2 viral uptake","author":"Riad","year":"2021","journal-title":"bioRxiv"},{"key":"10.1016\/j.jbc.2022.101710_bib52","doi-asserted-by":"crossref","first-page":"109364","DOI":"10.1016\/j.celrep.2021.109364","article-title":"Systematic analysis of SARS-CoV-2 infection of an ACE2-negative human airway cell","volume":"36","author":"Puray-Chavez","year":"2021","journal-title":"Cell Rep."}],"container-title":["Journal of Biological Chemistry"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0021925822001508?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0021925822001508?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2025,9,19]],"date-time":"2025-09-19T00:24:23Z","timestamp":1758241463000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0021925822001508"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,3]]},"references-count":52,"journal-issue":{"issue":"3","published-print":{"date-parts":[[2022,3]]}},"alternative-id":["S0021925822001508"],"URL":"https:\/\/doi.org\/10.1016\/j.jbc.2022.101710","relation":{},"ISSN":["0021-9258"],"issn-type":[{"value":"0021-9258","type":"print"}],"subject":[],"published":{"date-parts":[[2022,3]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Integrin mediates cell entry of the SARS-CoV-2 virus independent of cellular receptor ACE2","name":"articletitle","label":"Article Title"},{"value":"Journal of Biological Chemistry","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.jbc.2022.101710","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2022 The Authors. Published by Elsevier Inc on behalf of American Society for Biochemistry and Molecular Biology.","name":"copyright","label":"Copyright"}],"article-number":"101710"}}