{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T02:01:41Z","timestamp":1790733701424,"version":"4.1.0"},"reference-count":26,"publisher":"Elsevier BV","issue":"7","license":[{"start":{"date-parts":[[2021,4,1]],"date-time":"2021-04-01T00:00:00Z","timestamp":1617235200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2021,4,1]],"date-time":"2021-04-01T00:00:00Z","timestamp":1617235200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2022,4,1]],"date-time":"2022-04-01T00:00:00Z","timestamp":1648771200000},"content-version":"vor","delay-in-days":365,"URL":"http:\/\/www.elsevier.com\/open-access\/userlicense\/1.0\/"},{"start":{"date-parts":[[2021,4,1]],"date-time":"2021-04-01T00:00:00Z","timestamp":1617235200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-017"},{"start":{"date-parts":[[2021,4,1]],"date-time":"2021-04-01T00:00:00Z","timestamp":1617235200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2021,4,1]],"date-time":"2021-04-01T00:00:00Z","timestamp":1617235200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-012"},{"start":{"date-parts":[[2021,4,1]],"date-time":"2021-04-01T00:00:00Z","timestamp":1617235200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2021,4,1]],"date-time":"2021-04-01T00:00:00Z","timestamp":1617235200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-004"}],"funder":[{"DOI":"10.13039\/100012394","name":"Robert J. Kleberg, Jr. and Helen C. Kleberg Foundation","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100012394","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000183","name":"Army Research Office","doi-asserted-by":"publisher","award":["W911NF-15-1-0120"],"award-info":[{"award-number":["W911NF-15-1-0120"]}],"id":[{"id":"10.13039\/100000183","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000928","name":"Welch Foundation","doi-asserted-by":"publisher","award":["F-1604"],"award-info":[{"award-number":["F-1604"]}],"id":[{"id":"10.13039\/100000928","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000928","name":"Welch Foundation","doi-asserted-by":"publisher","award":["F-1938"],"award-info":[{"award-number":["F-1938"]}],"id":[{"id":"10.13039\/100000928","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100004917","name":"Cancer Prevention and Research Institute of Texas","doi-asserted-by":"publisher","award":["RR160088"],"award-info":[{"award-number":["RR160088"]}],"id":[{"id":"10.13039\/100004917","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100004917","name":"Cancer Prevention and Research Institute of Texas","doi-asserted-by":"publisher","award":["W911NF-19-1-0021"],"award-info":[{"award-number":["W911NF-19-1-0021"]}],"id":[{"id":"10.13039\/100004917","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01AI110577"],"award-info":[{"award-number":["R01AI110577"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100005564","name":"Gilead Sciences","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100005564","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100008562","name":"University of Texas at Austin","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100008562","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000060","name":"National Institute of Allergy and Infectious Diseases","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000060","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000057","name":"National Institute of General Medical Sciences","doi-asserted-by":"publisher","award":["R01114223"],"award-info":[{"award-number":["R01114223"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000057","name":"National Institute of General Medical Sciences","doi-asserted-by":"publisher","award":["R35GM138348"],"award-info":[{"award-number":["R35GM138348"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["cell.com","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Molecular Cell"],"published-print":{"date-parts":[[2021,4]]},"DOI":"10.1016\/j.molcel.2021.01.035","type":"journal-article","created":{"date-parts":[[2021,1,28]],"date-time":"2021-01-28T16:19:51Z","timestamp":1611850791000},"page":"1548-1552.e4","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":132,"title":["Remdesivir is a delayed translocation inhibitor of SARS-CoV-2 replication"],"prefix":"10.1016","volume":"81","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0456-0753","authenticated-orcid":false,"given":"Jack P.K.","family":"Bravo","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tyler L.","family":"Dangerfield","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6198-1194","authenticated-orcid":false,"given":"David W.","family":"Taylor","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6575-2823","authenticated-orcid":false,"given":"Kenneth A.","family":"Johnson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.molcel.2021.01.035_bib1","doi-asserted-by":"crossref","first-page":"531","DOI":"10.1107\/S2059798318006551","article-title":"Real-space refinement in PHENIX for cryo-EM and crystallography","volume":"74","author":"Afonine","year":"2018","journal-title":"Acta Crystallogr. D Struct. Biol."},{"key":"10.1016\/j.molcel.2021.01.035_bib2","doi-asserted-by":"crossref","first-page":"e00221-e18","DOI":"10.1128\/mBio.00221-18","article-title":"Coronavirus Susceptibility to the Antiviral Remdesivir (GS-5734) Is Mediated by the Viral Polymerase and the Proofreading Exoribonuclease","volume":"9","author":"Agostini","year":"2018","journal-title":"MBio"},{"key":"10.1016\/j.molcel.2021.01.035_bib3","doi-asserted-by":"crossref","first-page":"1813","DOI":"10.1056\/NEJMoa2007764","article-title":"Remdesivir for the Treatment of Covid-19 - Final Report","volume":"383","author":"Beigel","year":"2020","journal-title":"N.\u00a0Engl. J. Med."},{"key":"10.1016\/j.molcel.2021.01.035_bib4","doi-asserted-by":"crossref","first-page":"1560","DOI":"10.1016\/j.cell.2020.07.033","article-title":"Structural Basis for Helicase-Polymerase Coupling in the SARS-CoV-2 Replication-Transcription Complex","volume":"182","author":"Chen","year":"2020","journal-title":"Cell"},{"key":"10.1016\/j.molcel.2021.01.035_bib26","series-title":"COSMIC\u00b2: A Science Gateway for Cryo-Electron Microscopy Structure Determination","author":"Cianfrocco","year":"2017"},{"key":"10.1016\/j.molcel.2021.01.035_bib5","doi-asserted-by":"crossref","first-page":"101849","DOI":"10.1016\/j.isci.2020.101849","article-title":"Remdesivir Is Effective in Combating COVID-19 because It Is a Better Substrate than ATP for the Viral RNA-Dependent RNA Polymerase","volume":"23","author":"Dangerfield","year":"2020","journal-title":"iScience"},{"key":"10.1016\/j.molcel.2021.01.035_bib6","doi-asserted-by":"crossref","DOI":"10.1016\/j.xpro.2021.100357","article-title":"Expression and purification of tag-free SARS-CoV-2 RNA-dependent RNA polymerase in Escherichia coli","author":"Dangerfield","year":"2021","journal-title":"STAR Protoc."},{"key":"10.1016\/j.molcel.2021.01.035_bib7","doi-asserted-by":"crossref","first-page":"538","DOI":"10.1021\/bi00216a031","article-title":"Kinetic partitioning between the exonuclease and polymerase sites in DNA error correction","volume":"30","author":"Donlin","year":"1991","journal-title":"Biochemistry"},{"key":"10.1016\/j.molcel.2021.01.035_bib8","doi-asserted-by":"crossref","first-page":"2126","DOI":"10.1107\/S0907444904019158","article-title":"Coot: model-building tools for molecular graphics","volume":"60","author":"Emsley","year":"2004","journal-title":"Acta Crystallogr. D Biol. Crystallogr."},{"key":"10.1016\/j.molcel.2021.01.035_bib9","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1002\/pro.3235","article-title":"UCSF ChimeraX: meeting modern challenges in visualization and analysis","volume":"27","author":"Goddard","year":"2018","journal-title":"Protein Sci."},{"key":"10.1016\/j.molcel.2021.01.035_bib10","doi-asserted-by":"crossref","first-page":"6785","DOI":"10.1074\/jbc.RA120.013679","article-title":"Remdesivir is a direct-acting antiviral that inhibits RNA-dependent RNA polymerase from severe acute respiratory syndrome coronavirus 2 with high potency","volume":"295","author":"Gordon","year":"2020","journal-title":"J.\u00a0Biol. Chem."},{"key":"10.1016\/j.molcel.2021.01.035_bib11","doi-asserted-by":"crossref","first-page":"154","DOI":"10.1038\/s41586-020-2368-8","article-title":"Structure of replicating SARS-CoV-2 polymerase","volume":"584","author":"Hillen","year":"2020","journal-title":"Nature"},{"key":"10.1016\/j.molcel.2021.01.035_bib12","doi-asserted-by":"crossref","first-page":"38097","DOI":"10.1074\/jbc.M106046200","article-title":"Exonuclease proofreading by human mitochondrial DNA polymerase","volume":"276","author":"Johnson","year":"2001","journal-title":"J.\u00a0Biol. Chem."},{"key":"10.1016\/j.molcel.2021.01.035_bib13","doi-asserted-by":"crossref","first-page":"279","DOI":"10.1038\/s41467-020-20542-0","article-title":"Mechanism of SARS-CoV-2 polymerase inhibition by remdesivir","volume":"12","author":"Kokic","year":"2020","journal-title":"Nat. Commun."},{"key":"10.1016\/j.molcel.2021.01.035_bib14","doi-asserted-by":"crossref","first-page":"36","DOI":"10.1016\/j.jsb.2005.07.007","article-title":"Automated electron microscope tomography using robust prediction of specimen movements","volume":"152","author":"Mastronarde","year":"2005","journal-title":"J.\u00a0Struct. Biol."},{"key":"10.1016\/j.molcel.2021.01.035_bib15","doi-asserted-by":"crossref","first-page":"1074","DOI":"10.1107\/S0907444909029436","article-title":"electronic Ligand Builder and Optimization Workbench (eLBOW): a tool for ligand coordinate and restraint generation","volume":"65","author":"Moriarty","year":"2009","journal-title":"Acta Crystallogr. D Biol. Crystallogr."},{"key":"10.1016\/j.molcel.2021.01.035_bib16","doi-asserted-by":"crossref","first-page":"290","DOI":"10.1038\/nmeth.4169","article-title":"cryoSPARC: algorithms for rapid unsupervised cryo-EM structure determination","volume":"14","author":"Punjani","year":"2017","journal-title":"Nat. Methods"},{"key":"10.1016\/j.molcel.2021.01.035_bib25","article-title":"DeepEMhancer: a deep learning solution for cryo-EM volume post-processing","author":"Sanchez-Garcia","year":"2020","journal-title":"bioRxiv"},{"key":"10.1016\/j.molcel.2021.01.035_bib17","article-title":"Favipiravir strikes the SARS-CoV-2 at its Achilles heel, the RNA polymerase","author":"Shannon","year":"2020","journal-title":"bioRxiv"},{"key":"10.1016\/j.molcel.2021.01.035_bib18","doi-asserted-by":"crossref","first-page":"16156","DOI":"10.1074\/jbc.AC120.015720","article-title":"Template-dependent inhibition of coronavirus RNA-dependent RNA polymerase by remdesivir reveals a second mechanism of action","volume":"295","author":"Tchesnokov","year":"2020","journal-title":"J.\u00a0Biol. Chem."},{"key":"10.1016\/j.molcel.2021.01.035_bib24","doi-asserted-by":"crossref","first-page":"1146","DOI":"10.1038\/s41592-019-0580-y","article-title":"Real-time cryo-electron microscopy data preprocessing with Warp","volume":"16","author":"Tegunov","year":"2019","journal-title":"Nat Methods"},{"key":"10.1016\/j.molcel.2021.01.035_bib19","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1038\/s41422-020-0282-0","article-title":"Remdesivir and chloroquine effectively inhibit the recently emerged novel coronavirus (2019-nCoV) in\u00a0vitro","volume":"30","author":"Wang","year":"2020","journal-title":"Cell Res."},{"key":"10.1016\/j.molcel.2021.01.035_bib20","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1016\/j.cell.2020.05.034","article-title":"Structural Basis for RNA Replication by the SARS-CoV-2 Polymerase","volume":"182","author":"Wang","year":"2020","journal-title":"Cell"},{"key":"10.1016\/j.molcel.2021.01.035_bib21","doi-asserted-by":"crossref","first-page":"5874","DOI":"10.1038\/s41467-020-19770-1","article-title":"Architecture of a SARS-CoV-2\u00a0mini replication and transcription complex","volume":"11","author":"Yan","year":"2020","journal-title":"Nat. Commun."},{"key":"10.1016\/j.molcel.2021.01.035_bib22","doi-asserted-by":"crossref","first-page":"184","DOI":"10.1016\/j.cell.2020.11.016","article-title":"Cryo-EM Structure of an Extended SARS-CoV-2 Replication and Transcription Complex Reveals an Intermediate State in Cap Synthesis","volume":"184","author":"Yan","year":"2021","journal-title":"Cell"},{"key":"10.1016\/j.molcel.2021.01.035_bib23","doi-asserted-by":"crossref","first-page":"1499","DOI":"10.1126\/science.abc1560","article-title":"Structural basis for inhibition of the RNA-dependent RNA polymerase from SARS-CoV-2 by remdesivir","volume":"368","author":"Yin","year":"2020","journal-title":"Science"}],"container-title":["Molecular Cell"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S1097276521000551?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S1097276521000551?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2025,10,22]],"date-time":"2025-10-22T00:37:07Z","timestamp":1761093427000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S1097276521000551"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,4]]},"references-count":26,"journal-issue":{"issue":"7","published-print":{"date-parts":[[2021,4]]}},"alternative-id":["S1097276521000551"],"URL":"https:\/\/doi.org\/10.1016\/j.molcel.2021.01.035","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2020.12.14.422718","asserted-by":"object"}]},"ISSN":["1097-2765"],"issn-type":[{"value":"1097-2765","type":"print"}],"subject":[],"published":{"date-parts":[[2021,4]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Remdesivir is a delayed translocation inhibitor of SARS-CoV-2 replication","name":"articletitle","label":"Article Title"},{"value":"Molecular Cell","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.molcel.2021.01.035","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2021 Elsevier Inc.","name":"copyright","label":"Copyright"}]}}