{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T20:11:19Z","timestamp":1786738279437,"version":"build-2736575974"},"reference-count":108,"publisher":"American Chemical Society (ACS)","issue":"11","license":[{"start":{"date-parts":[[2021,3,10]],"date-time":"2021-03-10T00:00:00Z","timestamp":1615334400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2021,3,23]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Severe acute respiratory coronavirus 2 (SARS-CoV-2) is a newly identified virus that has resulted in over 2.5 million deaths globally and over 116 million cases globally in March, 2021. Small-molecule inhibitors that reverse disease severity have proven difficult to discover. One of the key approaches that has been widely applied in an effort to speed up the translation of drugs is drug repurposing. A few drugs have shown in vitro activity against Ebola viruses and demonstrated activity against SARS-CoV-2 in vivo. Most notably, the RNA polymerase targeting remdesivir demonstrated activity in vitro and efficacy in the early stage of the disease in humans. Testing other small-molecule drugs that are active against Ebola viruses (EBOVs) would appear a reasonable strategy to evaluate their potential for SARS-CoV-2. We have previously repurposed pyronaridine, tilorone, and quinacrine (from malaria, influenza, and antiprotozoal uses, respectively) as inhibitors of Ebola and Marburg viruses in vitro in HeLa cells and mouse-adapted EBOV in mice in vivo. We have now tested these three drugs in various cell lines (VeroE6, Vero76, Caco-2, Calu-3, A549-ACE2, HUH-7, and monocytes) infected with SARS-CoV-2 as well as other viruses (including MHV and HCoV 229E). The compilation of these results indicated considerable variability in antiviral activity observed across cell lines. We found that tilorone and pyronaridine inhibited the virus replication in A549-ACE2 cells with IC50 values of 180 nM and IC50 198 nM, respectively. We used microscale thermophoresis to test the binding of these molecules to the spike protein, and tilorone and pyronaridine bind to the spike receptor binding domain protein with Kd values of 339 and 647 nM, respectively. Human Cmax for pyronaridine and quinacrine is greater than the IC50 observed in A549-ACE2 cells. We also provide novel insights into the mechanism of these compounds which is likely lysosomotropic.<\/jats:p>","DOI":"10.1021\/acsomega.0c05996","type":"journal-article","created":{"date-parts":[[2021,3,10]],"date-time":"2021-03-10T05:13:12Z","timestamp":1615353192000},"page":"7454-7468","source":"Crossref","is-referenced-by-count":69,"title":["Repurposing the Ebola and Marburg Virus Inhibitors\nTilorone, Quinacrine, and Pyronaridine:\n                    <i>In Vitro<\/i>\n                    Activity\nagainst SARS-CoV\u20112 and\nPotential Mechanisms"],"prefix":"10.1021","volume":"6","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1456-8882","authenticated-orcid":true,"given":"Ana C.","family":"Puhl","sequence":"first","affiliation":[{"name":"Collaborations Pharmaceuticals, Inc. , 840 Main Campus Drive, Lab 3510 , , ,","place":["Raleigh, North Carolina, United States, 27606"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ethan J.","family":"Fritch","sequence":"additional","affiliation":[{"name":"University of North Carolina School of Medicine , , , ,","place":["Chapel Hill, North Carolina, United States, 27599"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thomas R.","family":"Lane","sequence":"additional","affiliation":[{"name":"Collaborations Pharmaceuticals, Inc. , 840 Main Campus Drive, Lab 3510 , , ,","place":["Raleigh, North Carolina, United States, 27606"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Longping V.","family":"Tse","sequence":"additional","affiliation":[{"name":"University of North Carolina School of Medicine , , , ,","place":["Chapel Hill, North Carolina, United States, 27599"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Boyd L.","family":"Yount","sequence":"additional","affiliation":[{"name":"University of North Carolina School of Medicine , , , ,","place":["Chapel Hill, North Carolina, United States, 27599"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Carolina Q.","family":"Sacramento","sequence":"additional","affiliation":[{"name":"Fundac\u0327a\u0303o Oswaldo Cruz (Fiocruz) , , ,","place":["Rio de Janeiro, RJ, Brazil, 21040-900"]},{"name":"Fiocruz , , ,","place":["Rio de\nJaneiro, Brazil, 21040-900"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Natalia","family":"Fintelman-Rodrigues","sequence":"additional","affiliation":[{"name":"Fundac\u0327a\u0303o Oswaldo Cruz (Fiocruz) , , ,","place":["Rio de Janeiro, RJ, Brazil, 21040-900"]},{"name":"Fiocruz , , ,","place":["Rio de\nJaneiro, Brazil, 21040-900"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tatyana Almeida","family":"Tavella","sequence":"additional","affiliation":[{"name":"University of Campinas-UNICAMP , , Campinas, ,","place":["Sa\u0303o Paulo, Brazil, 13083-970"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9969-7300","authenticated-orcid":true,"given":"Fabio Trindade","family":"Maranha\u0303o Costa","sequence":"additional","affiliation":[{"name":"University of Campinas-UNICAMP , , Campinas, ,","place":["Sa\u0303o Paulo, Brazil, 13083-970"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stuart","family":"Weston","sequence":"additional","affiliation":[{"name":"University of Maryland School of Medicine , , , ,","place":["Baltimore, Maryland, United States, 21201"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7410-9741","authenticated-orcid":true,"given":"James","family":"Logue","sequence":"additional","affiliation":[{"name":"University of Maryland School of Medicine , , , ,","place":["Baltimore, Maryland, United States, 21201"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Matthew","family":"Frieman","sequence":"additional","affiliation":[{"name":"University of Maryland School of Medicine , , , ,","place":["Baltimore, Maryland, United States, 21201"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lakshmanane","family":"Premkumar","sequence":"additional","affiliation":[{"name":"University of North Carolina School of Medicine , , , ,","place":["Chapel Hill, North Carolina, United States, 27599"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kenneth H.","family":"Pearce","sequence":"additional","affiliation":[{"name":"University of North Carolina , , , ,","place":["Chapel Hill, North Carolina, United States, 27599"]},{"name":"UNC Lineberger Comprehensive Cancer Center , , ,","place":["Chapel Hill, North Carolina, United States, 27599"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Brett L.","family":"Hurst","sequence":"additional","affiliation":[{"name":"Utah State University , , , ,","place":["Logan, Utah, United States, 84322"]},{"name":"Utah State University , , , ,","place":["Logan, Utah, United States, 84322"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Carolina Horta","family":"Andrade","sequence":"additional","affiliation":[{"name":"University of Campinas-UNICAMP , , Campinas, ,","place":["Sa\u0303o Paulo, Brazil, 13083-970"]},{"name":"Universidade Federal de Goia\u0301s , , ,","place":["Goia\u0302nia,\nGO, Brazil, 74605-170"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"James A.","family":"Levi","sequence":"additional","affiliation":[{"name":"North Carolina State University , , , ,","place":["Raleigh, North Carolina, United States, 27695"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nicole J.","family":"Johnson","sequence":"additional","affiliation":[{"name":"North Carolina State University , , , ,","place":["Raleigh, North Carolina, United States, 27695"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Samantha C.","family":"Kisthardt","sequence":"additional","affiliation":[{"name":"North Carolina State University , , , ,","place":["Raleigh, North Carolina, United States, 27695"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Frank","family":"Scholle","sequence":"additional","affiliation":[{"name":"North Carolina State University , , , ,","place":["Raleigh, North Carolina, United States, 27695"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thiago Moreno L.","family":"Souza","sequence":"additional","affiliation":[{"name":"Fundac\u0327a\u0303o Oswaldo Cruz (Fiocruz) , , ,","place":["Rio de Janeiro, RJ, Brazil, 21040-900"]},{"name":"Fiocruz , , ,","place":["Rio de\nJaneiro, Brazil, 21040-900"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nathaniel John","family":"Moorman","sequence":"additional","affiliation":[{"name":"University of North Carolina School of Medicine , , , ,","place":["Chapel Hill, North Carolina, United States, 27599"]},{"name":"University of North Carolina , , , ,","place":["Chapel Hill, North Carolina, United States, 27599"]},{"name":"University of North Carolina at Chapel Hill , , , ,","place":["Chapel\nHill, North Carolina, United States, 27599"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ralph S.","family":"Baric","sequence":"additional","affiliation":[{"name":"University of North Carolina School of Medicine , , , ,","place":["Chapel Hill, North Carolina, United States, 27599"]},{"name":"University of North Carolina School of Medicine , , , ,","place":["Chapel Hill, North Carolina, United States, 27599"]},{"name":"University of North Carolina at Chapel Hill , , , ,","place":["Chapel\nHill, North Carolina, United States, 27599"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peter B.","family":"Madrid","sequence":"additional","affiliation":[{"name":"SRI International , 333 Ravenswood Avenue , , ,","place":["Menlo Park, California, United States, 94025"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5691-5790","authenticated-orcid":true,"given":"Sean","family":"Ekins","sequence":"additional","affiliation":[{"name":"Collaborations Pharmaceuticals, Inc. , 840 Main Campus Drive, Lab 3510 , , ,","place":["Raleigh, North Carolina, United States, 27606"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2021,3,10]]},"reference":[{"key":"2026081415080944600_cit1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-020-2202-3","article-title":"Author Correction: A new coronavirus associated with\nhuman respiratory disease in 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Microbiol."},{"key":"2026081415080944600_cit3","doi-asserted-by":"publisher","first-page":"491","DOI":"10.1002\/jmv.25709","article-title":"Overlapping and discrete aspects of the pathology and pathogenesis\nof the emerging human pathogenic coronaviruses SARS-CoV, MERS-CoV,\nand 2019-nCoV","volume":"92","author":"Liu","year":"2020","journal-title":"J. Med. Virol."},{"key":"2026081415080944600_cit4","unstructured":"WHO Naming the coronavirus disease (COVID-2019) and the virus that\ncauses it. https:\/\/www.who.int\/emergencies\/diseases\/novel-coronavirus-2019\/technical-guidance\/naming-the-coronavirus-disease-(covid-2019)-and-the-virus-that-causes-it (accessed Dec 10, 2020)."},{"key":"2026081415080944600_cit5","doi-asserted-by":"publisher","first-page":"3306","DOI":"10.1007\/s00330-020-06731-x","article-title":"Initial CT findings and temporal\nchanges in patients with the novel coronavirus pneumonia (2019-nCoV):\na study of 63 patients in Wuhan, China","volume":"30","author":"Pan","year":"2020","journal-title":"Eur.\nRadiol."},{"key":"2026081415080944600_cit6","doi-asserted-by":"publisher","first-page":"2512","DOI":"10.1001\/jama.2020.839","article-title":"Olfactory Dysfunction\nin COVID-19: Diagnosis and Management","volume":"323","author":"Whitcroft","year":"2020","journal-title":"JAMA"},{"key":"2026081415080944600_cit7","doi-asserted-by":"publisher","first-page":"681","DOI":"10.1038\/s41591-020-0868-6","article-title":"SARS-CoV-2\nentry factors are highly expressed in nasal epithelial cells together\nwith innate immune genes","volume":"26","author":"Sungnak","year":"2020","journal-title":"Nat Med."},{"key":"2026081415080944600_cit8","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.abc5801","article-title":"Non-neuronal expression of SARS-CoV-2 entry genes in\nthe olfactory system suggests mechanisms underlying COVID-19-associated\nanosmia","volume":"6","author":"Brann","year":"2020","journal-title":"Sci. Adv."},{"key":"2026081415080944600_cit9","doi-asserted-by":"crossref","unstructured":"World Health Organization\n          . WHO Coronavirus Disease (COVID-19)\nDashboard. https:\/\/covid19.who.int (accessed Dec 10, 2020).","DOI":"10.46945\/bpj.10.1.03.01"},{"key":"2026081415080944600_cit10","doi-asserted-by":"publisher","first-page":"2293","DOI":"10.1056\/nejmoa1910993","article-title":"A Randomized,\nControlled Trial of Ebola Virus Disease Therapeutics","volume":"381","author":"Mulangu","year":"2019","journal-title":"N. Engl. J. Med."},{"key":"2026081415080944600_cit11","doi-asserted-by":"publisher","first-page":"6771","DOI":"10.1073\/pnas.1922083117","article-title":"Prophylactic\nand therapeutic remdesivir (GS-5734) treatment in the rhesus macaque\nmodel of MERS-CoV infection","volume":"117","author":"de\nWit","year":"2020","journal-title":"Proc. Natl. Acad.\nSci. U.S.A."},{"key":"2026081415080944600_cit12","doi-asserted-by":"publisher","DOI":"10.1126\/scitranslmed.aal3653","article-title":"Broad-spectrum\nantiviral GS-5734 inhibits both epidemic and zoonotic coronaviruses","volume":"9","author":"Sheahan","year":"2017","journal-title":"Sci. Transl. Med."},{"key":"2026081415080944600_cit13","doi-asserted-by":"publisher","first-page":"107940","DOI":"10.1016\/j.celrep.2020.107940","article-title":"Remdesivir Inhibits SARS-CoV-2 in Human Lung Cells\nand Chimeric SARS-CoV Expressing the SARS-CoV-2 RNA Polymerase in\nMice","volume":"32","author":"Pruijssers","year":"2020","journal-title":"Cell Rep."},{"key":"2026081415080944600_cit14","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1038\/s41422-020-0282-0","article-title":"Remdesivir and chloroquine\neffectively inhibit the recently emerged novel coronavirus (2019-nCoV)\nin vitro","volume":"30","author":"Wang","year":"2020","journal-title":"Cell Res."},{"key":"2026081415080944600_cit15","doi-asserted-by":"publisher","first-page":"1569","DOI":"10.1016\/s0140-6736(20)31022-9","article-title":"Remdesivir\nin adults with severe COVID-19: a randomised, double-blind, placebo-controlled,\nmulticentre trial","volume":"395","author":"Wang","year":"2020","journal-title":"Lancet"},{"key":"2026081415080944600_cit16","doi-asserted-by":"publisher","first-page":"1355","DOI":"10.1007\/s40265-020-01378-w","article-title":"Remdesivir:\nFirst Approval","volume":"80","author":"Lamb","year":"2020","journal-title":"Drugs"},{"key":"2026081415080944600_cit17","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1038\/s41421-020-0156-0","article-title":"Hydroxychloroquine,\na less toxic derivative of chloroquine, is effective in inhibiting\nSARS-CoV-2 infection in vitro","volume":"6","author":"Liu","year":"2020","journal-title":"Cell Discovery"},{"key":"2026081415080944600_cit18","doi-asserted-by":"publisher","first-page":"517","DOI":"10.1056\/nejmoa2016638","article-title":"A Randomized Trial\nof Hydroxychloroquine as Postexposure Prophylaxis for Covid-19","volume":"383","author":"Boulware","year":"2020","journal-title":"N. Engl. J. Med."},{"key":"2026081415080944600_cit19","doi-asserted-by":"publisher","DOI":"10.2196\/21758","article-title":"Efficacy of\nHydroxychloroquine and\nTocilizumab in Patients With COVID-19: Single-Center Retrospective\nChart Review","volume":"22","author":"Roomi","year":"2020","journal-title":"J. Med. Internet Res."},{"key":"2026081415080944600_cit20","doi-asserted-by":"publisher","first-page":"829","DOI":"10.1126\/science.abb9332","article-title":"Rapid repurposing\nof drugs for COVID-19","volume":"368","author":"Guy","year":"2020","journal-title":"Science"},{"key":"2026081415080944600_cit21","doi-asserted-by":"publisher","first-page":"661","DOI":"10.1016\/j.drudis.2018.01.018","article-title":"A bibliometric\nreview\nof drug repurposing","volume":"23","author":"Baker","year":"2018","journal-title":"Drug Discovery Today"},{"key":"2026081415080944600_cit22","doi-asserted-by":"crossref","DOI":"10.1101\/2020.05.27.117184","article-title":"Morphological Cell Profiling\nof SARS-CoV-2 Infection Identifies Drug Repurposing Candidates for\nCOVID-19","author":"Mirabelli","year":"2020"},{"key":"2026081415080944600_cit23","doi-asserted-by":"publisher","first-page":"104960","DOI":"10.1016\/j.phrs.2020.104960","article-title":"Discovery of the\nFDA-approved drugs\nbexarotene, cetilistat, diiodohydroxyquinoline, and abiraterone as\npotential COVID-19 treatments with a robust two-tier screening system","volume":"159","author":"Yuan","year":"2020","journal-title":"Pharmacol. 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Agents Chemother."},{"key":"2026081415080944600_cit27","doi-asserted-by":"publisher","first-page":"289","DOI":"10.1038\/s41586-020-2223-y","article-title":"Structure of Mpro from\nSARS-CoV-2 and discovery of its inhibitors","volume":"582","author":"Jin","year":"2020","journal-title":"Nature"},{"key":"2026081415080944600_cit28","doi-asserted-by":"crossref","DOI":"10.1101\/2020.06.16.154765","article-title":"Machine Learning Models Identify Inhibitors\nof SARS-CoV-2","author":"Gawriljuk","year":"2020"},{"key":"2026081415080944600_cit29","doi-asserted-by":"publisher","DOI":"10.1128\/aac.01142-19","article-title":"Repurposing\nQuinacrine against Ebola Virus InfectionIn Vivo","volume":"63","author":"Lane","year":"2019","journal-title":"Antimicrob. Agents Chemother."},{"key":"2026081415080944600_cit30","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pntd.0007890","article-title":"Repurposing\nThe Antimalarial Pyronaridine Tetraphosphate To Protect Against Ebola\nVirus Infection","volume":"13","author":"Lane","year":"2019","journal-title":"PLoS Neglected Trop. Dis."},{"key":"2026081415080944600_cit31","doi-asserted-by":"publisher","DOI":"10.1128\/aac.01711-17","article-title":"Efficacy\nof Tilorone Dihydrochloride\nagainst Ebola Virus Infection","volume":"62","author":"Ekins","year":"2018","journal-title":"Antimicrob. Agents\nChemother."},{"key":"2026081415080944600_cit32","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1007\/s11095-020-02799-8","article-title":"Tilorone:\na Broad-Spectrum Antiviral\nInvented in the USA and Commercialized in Russia and beyond","volume":"37","author":"Ekins","year":"2020","journal-title":"Pharm. Res."},{"key":"2026081415080944600_cit33","doi-asserted-by":"publisher","first-page":"1653","DOI":"10.1021\/acsmedchemlett.0c00298","article-title":"Toward the Target:\nTilorone, Quinacrine, and Pyronaridine Bind to\nEbola Virus Glycoprotein","volume":"11","author":"Lane","year":"2020","journal-title":"ACS Med. Chem. Lett."},{"key":"2026081415080944600_cit34","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1007\/978-1-4939-0597-3_2","article-title":"Pharmacology\nof Chloroquine and Hydroxychloroquine","volume-title":"Hydroxychloroquine\nand chloroquine retinopathy","author":"Browning","year":"2014"},{"key":"2026081415080944600_cit35","doi-asserted-by":"publisher","first-page":"11","DOI":"10.2174\/1874104501105010011","article-title":"Acridine\nand acridinones: old and new structures with antimalarial activity","volume":"5","author":"Valde\u0301s","year":"2011","journal-title":"Open Med. Chem. J."},{"key":"2026081415080944600_cit36","doi-asserted-by":"publisher","first-page":"859","DOI":"10.1515\/bchm2.1984.365.2.859","article-title":"Tilorone acts\nas a lysosomotropic agent in fibroblasts","volume":"365","author":"Gupta","year":"1984","journal-title":"Hoppe\nSeylers Z. Physiol. Chem."},{"key":"2026081415080944600_cit37","doi-asserted-by":"publisher","DOI":"10.1002\/bip.23398","article-title":"Pyronaridine:\nAn update of its pharmacological activities and mechanisms of action","author":"Bailly","year":"2020","journal-title":"Biopolymers"},{"key":"2026081415080944600_cit38","doi-asserted-by":"publisher","DOI":"10.1128\/aac.01142-19","article-title":"Repurposing\nQuinacrine against Ebola Virus Infection","volume":"63","author":"Lane","year":"2019","journal-title":"Antimicrob.\nAgents Chemother."},{"key":"2026081415080944600_cit39","doi-asserted-by":"publisher","first-page":"104863","DOI":"10.1016\/j.antiviral.2020.104863","article-title":"Pyronaridine tetraphosphate\nefficacy against Ebola virus infection in guinea pig","volume":"181","author":"Lane","year":"2020","journal-title":"Antiviral Res."},{"key":"2026081415080944600_cit40","doi-asserted-by":"publisher","DOI":"10.1128\/aac.00440-20","article-title":"Tilorone, a Broad-Spectrum\nAntiviral for Emerging Viruses","volume":"64","author":"Ekins","year":"2020","journal-title":"Antimicrob.\nAgents Chemother."},{"key":"2026081415080944600_cit41","article-title":"The in vitro antiviral activity\nof the anti-hepatitis C virus (HCV) drugs daclatasvir and sofosbuvir\nagainst SARS-CoV-2","author":"Sacramento","year":"2020"},{"key":"2026081415080944600_cit42","doi-asserted-by":"publisher","first-page":"3039","DOI":"10.1128\/jvi.01792-08","article-title":"Severe\nacute respiratory syndrome (SARS) coronavirus-induced lung epithelial\ncytokines exacerbate SARS pathogenesis by modulating intrinsic functions\nof monocyte-derived macrophages and dendritic cells","volume":"83","author":"Yoshikawa","year":"2009","journal-title":"J. Virol."},{"key":"2026081415080944600_cit43","doi-asserted-by":"publisher","first-page":"11065","DOI":"10.1128\/jvi.76.21.11065-11078.2002","article-title":"Systematic assembly\nof a full-length infectious cDNA of mouse hepatitis virus strain A59","volume":"76","author":"Yount","year":"2002","journal-title":"J. Virol."},{"key":"2026081415080944600_cit44","doi-asserted-by":"publisher","first-page":"9864","DOI":"10.1073\/pnas.1604472113","article-title":"Link of a ubiquitous\nhuman coronavirus to dromedary camels","volume":"113","author":"Corman","year":"2016","journal-title":"Proc.\nNatl. Acad. Sci. 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