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Among these, the UL30 DNA polymerase, complexed with the UL42 processivity factor, orchestrates leading and lagging strand replication of the 152 kb viral genome. UL30 polymerase is a prime target for antiviral therapy, and resistance to current drugs can arise in immunocompromised individuals. Using electron cryo-microscopy (cryo-EM), we unveil the dynamic changes of the UL30\/UL42 complex with DNA in three distinct states. First, a pre-translocation state with an open fingers domain ready for nucleotide incorporation. Second, a halted elongation state where the fingers close, trapping dATP in the dNTP pocket. Third, a DNA-editing state involving significant conformational changes to allow DNA realignment for exonuclease activity. Additionally, the flexible UL30 C-terminal domain interacts with UL42, forming an extended positively charged surface binding to DNA, thereby enhancing processive synthesis. These findings highlight substantial structural shifts in the polymerase and its DNA interactions during replication, offering insights for future antiviral drug development.<\/jats:p>","DOI":"10.1093\/nar\/gkae374","type":"journal-article","created":{"date-parts":[[2024,5,29]],"date-time":"2024-05-29T01:11:46Z","timestamp":1716945106000},"page":"7292-7304","source":"Crossref","is-referenced-by-count":21,"title":["Dynamics of the Herpes simplex virus DNA polymerase holoenzyme during DNA synthesis and proof-reading revealed by Cryo-EM"],"prefix":"10.1093","volume":"52","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0166-1786","authenticated-orcid":false,"given":"Emil","family":"Gustavsson","sequence":"first","affiliation":[{"name":"Department of Cell and Molecular Biology, Karolinska Institutet , 171 77\u00a0 Stockholm , Sweden"},{"name":"Centre for Structural Systems Biology , Deutsches Elektronen-Synchrotron DESY, Notkestra\u00dfe 85, Building 15, 22607\u00a0 Hamburg , Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kay","family":"Gr\u00fcnewald","sequence":"additional","affiliation":[{"name":"Centre for Structural Systems Biology , Deutsches Elektronen-Synchrotron DESY, Notkestra\u00dfe 85, Building 15, 22607\u00a0 Hamburg , Germany"},{"name":"Leibniz-Institute of Virology , Martinistra\u00dfe 52, 20251\u00a0 Hamburg , Germany"},{"name":"Department of Chemistry, University of Hamburg , Martin-Luther-King-Platz 6, 20146\u00a0 Hamburg , Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Per","family":"Elias","sequence":"additional","affiliation":[{"name":"Institute of Biomedicine, Department of Medical Biochemistry and Cell Biology, Sahlgrenska Academy, University of Gothenburg , Box 440, 405 30\u00a0 Gothenburg , Sweden"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"B\u00a0Martin","family":"H\u00e4llberg","sequence":"additional","affiliation":[{"name":"Department of Cell and Molecular Biology, Karolinska Institutet , 171 77\u00a0 Stockholm , Sweden"},{"name":"Centre for Structural Systems Biology , Deutsches Elektronen-Synchrotron DESY, Notkestra\u00dfe 85, Building 15, 22607\u00a0 Hamburg , Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2024,5,29]]},"reference":[{"key":"2024070813465214400_B1","doi-asserted-by":"crossref","first-page":"1513","DOI":"10.1016\/S0140-6736(00)04638-9","article-title":"Herpes simplex virus infections","volume":"357","author":"Whitley","year":"2021","journal-title":"Lancet"},{"key":"2024070813465214400_B2","doi-asserted-by":"crossref","first-page":"1726","DOI":"10.12688\/f1000research.16157.1","article-title":"Clinical management of herpes simplex virus infections: past, present, and future","volume":"7","author":"Whitley","year":"2018","journal-title":"F1000Research"},{"key":"2024070813465214400_B3","doi-asserted-by":"crossref","first-page":"114","DOI":"10.1128\/CMR.16.1.114-128.2003","article-title":"Herpes simplex virus resistance to Acyclovir and Penciclovir after two decades of antiviral therapy","volume":"16","author":"Bacon","year":"2003","journal-title":"Clin. 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