{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,27]],"date-time":"2026-02-27T08:26:14Z","timestamp":1772180774140,"version":"3.50.1"},"reference-count":58,"publisher":"Oxford University Press (OUP)","issue":"5","license":[{"start":{"date-parts":[[2026,2,27]],"date-time":"2026-02-27T00:00:00Z","timestamp":1772150400000},"content-version":"vor","delay-in-days":3,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"}],"funder":[{"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\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R15 AI090573"],"award-info":[{"award-number":["R15 AI090573"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000097","name":"National Center for Research Resources","doi-asserted-by":"publisher","award":["P20 RR-016464"],"award-info":[{"award-number":["P20 RR-016464"]}],"id":[{"id":"10.13039\/100000097","id-type":"DOI","asserted-by":"publisher"}]},{"name":"U.S. Department of Education GAANN fellowship","award":["P200A210055"],"award-info":[{"award-number":["P200A210055"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,2,24]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>In bacteria, nucleoid-structuring proteins bind and constrain DNA, often leading to transcriptional silencing. In Shigella spp., the histone-like nucleoid-structuring protein H-NS silences many genes on the large virulence plasmid. Upon a shift to human body temperature, VirB, a DNA-binding protein and key transcriptional regulator of the Shigella virulence cascade, is produced. VirB counteracts H-NS-mediated transcriptional silencing and belongs to a fast-evolving clade of the ParB superfamily. Like other ParB proteins, VirB binds the ligand CTP. While CTP is essential for the anti-silencing activity of VirB, the role of CTP in the mechanism of VirB-dependent anti-silencing has yet to be determined. This work shows that VirB does not require CTP for specific engagement of its DNA recognition site, but CTP is necessary for the formation of large VirB\u2013DNA complexes, which likely form when VirB dimers adopt a sliding clamp conformation. Furthermore, CTP-binding mutants do not trigger a loss of negative supercoils from plasmid DNA, a VirB-dependent activity proposed to destabilize H-NS\u2013DNA complexes. Together, our findings provide novel insight into the relationship between VirB, CTP, and DNA and reveal the role of the VirB ligand, CTP, in regulating Shigella virulence.<\/jats:p>","DOI":"10.1093\/nar\/gkag115","type":"journal-article","created":{"date-parts":[[2026,1,31]],"date-time":"2026-01-31T12:39:13Z","timestamp":1769863153000},"source":"Crossref","is-referenced-by-count":0,"title":["The role of the VirB ligand CTP in the molecular mechanism of transcriptional anti-silencing in\n                    <i>Shigella flexneri<\/i>"],"prefix":"10.1093","volume":"54","author":[{"given":"Taylor M","family":"Gerson","sequence":"first","affiliation":[{"name":"School of Life Sciences, University of Nevada , Las Vegas, NV 89154-4004 ,","place":["United States"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Monika M A","family":"Karney","sequence":"additional","affiliation":[{"name":"School of Life Sciences, University of Nevada , Las Vegas, NV 89154-4004 ,","place":["United States"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0123-5568","authenticated-orcid":false,"given":"Helen J","family":"Wing","sequence":"additional","affiliation":[{"name":"School of Life Sciences, University of Nevada , Las Vegas, NV 89154-4004 ,","place":["United States"]}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"286","published-online":{"date-parts":[[2026,2,27]]},"reference":[{"key":"2026022702152293000_B1","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/S0065-2164(08)01002-2","article-title":"Nucleoid-associated proteins and bacterial physiology","volume":"67","author":"Dorman","year":"2009","journal-title":"Adv Appl Microbiol"},{"key":"2026022702152293000_B2","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1016\/0966-842X(93)90114-7","article-title":"DNA topology and bacterial virulence gene regulation","volume":"1","author":"Dorman","year":"1993","journal-title":"Trends Microbiol"},{"key":"2026022702152293000_B3","doi-asserted-by":"publisher","first-page":"e3001557","DOI":"10.1371\/journal.pbio.3001557","article-title":"Correction: dynamic landscape of protein occupancy across the Escherichia coli chromosome","volume":"20","author":"Freddolino","year":"2022","journal-title":"PLoS Biol"},{"key":"2026022702152293000_B4","doi-asserted-by":"publisher","first-page":"e3001306","DOI":"10.1371\/journal.pbio.3001306","article-title":"Dynamic landscape of protein occupancy across the Escherichia coli chromosome","volume":"19","author":"Freddolino","year":"2021","journal-title":"PLoS Biol"},{"key":"2026022702152293000_B5","doi-asserted-by":"publisher","first-page":"148","DOI":"10.1016\/S1369-5274(99)80026-5","article-title":"Interplay of global regulators and cell physiology in the general stress response of Escherichia coli","volume":"2","author":"Hengge-Aronis","year":"1999","journal-title":"Curr Opin Microbiol"},{"key":"2026022702152293000_B6","doi-asserted-by":"publisher","first-page":"965","DOI":"10.1046\/j.1365-2958.1996.436961.x","article-title":"Antagonistic involvement of FIS and H-NS proteins in the transcriptional control of hns expression","volume":"19","author":"Falconi","year":"1996","journal-title":"Mol Microbiol"},{"key":"2026022702152293000_B7","doi-asserted-by":"publisher","first-page":"439","DOI":"10.1046\/j.1365-2958.2001.02646.x","article-title":"Involvement of FIS in the H-NS-mediated regulation of virF gene of Shigella and enteroinvasive Escherichia coli","volume":"42","author":"Falconi","year":"2001","journal-title":"Mol Microbiol"},{"key":"2026022702152293000_B8","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1111\/j.1574-6968.1994.tb07082.x","article-title":"Transcription of the Salmonella typhimurium spv virulence locus is regulated negatively by the nucleoid-associated protein H-NS","volume":"121","author":"O\u2019Byrne","year":"1994","journal-title":"FEMS Microbiol Lett"},{"key":"2026022702152293000_B9","doi-asserted-by":"publisher","first-page":"15333","DOI":"10.1074\/jbc.M111429200","article-title":"Molecular dissection of VirB, a key regulator of the virulence cascade of Shigella flexneri","volume":"277","author":"Beloin","year":"2002","journal-title":"J Biol Chem"},{"key":"2026022702152293000_B10","doi-asserted-by":"publisher","first-page":"6347","DOI":"10.1128\/JB.182.22.6347-6357.2000","article-title":"Silencing and activation of ClyA cytotoxin expression in Escherichia coli","volume":"182","author":"Westermark","year":"2000","journal-title":"J Bacteriol"},{"key":"2026022702152293000_B11","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1111\/j.1365-2958.2007.05907.x","article-title":"The RovA regulons of Yersinia enterocolitica and Yersinia pestis are distinct: evidence that many RovA-regulated genes were acquired more recently than the core genome","volume":"66","author":"Cathelyn","year":"2007","journal-title":"Mol Microbiol"},{"key":"2026022702152293000_B12","doi-asserted-by":"publisher","first-page":"119","DOI":"10.1046\/j.1365-2958.2002.02721.x","article-title":"Regulation of gene expression in Vibrio cholerae by ToxT involves both antirepression and RNA polymerase stimulation","volume":"43","author":"Yu","year":"2002","journal-title":"Mol Microbiol"},{"key":"2026022702152293000_B13","doi-asserted-by":"publisher","first-page":"314","DOI":"10.1111\/j.1365-2958.2008.06171.x","article-title":"Bicarbonate-mediated transcriptional activation of divergent operons by the virulence regulatory protein, RegA, from Citrobacter rodentium","volume":"68","author":"Yang","year":"2008","journal-title":"Mol Microbiol"},{"key":"2026022702152293000_B14","doi-asserted-by":"publisher","first-page":"5101","DOI":"10.1074\/jbc.M411840200","article-title":"A cis-spreading nucleoprotein filament is responsible for the gene silencing activity found in the promoter relay mechanism","volume":"280","author":"Chen","year":"2005","journal-title":"J Biol Chem"},{"key":"2026022702152293000_B15","doi-asserted-by":"publisher","first-page":"15111","DOI":"10.1074\/jbc.M414544200","article-title":"LeuO protein delimits the transcriptionally active and repressive domains on the bacterial chromosome","volume":"280","author":"Chen","year":"2005","journal-title":"J Biol Chem"},{"key":"2026022702152293000_B16","doi-asserted-by":"publisher","first-page":"2533","DOI":"10.1099\/mic.0.2008\/020693-0","article-title":"Anti-silencing: overcoming H-NS-mediated repression of transcription in Gram-negative enteric bacteria","volume":"154","author":"Stoebel","year":"2008","journal-title":"Microbiology (Reading)"},{"key":"2026022702152293000_B17","doi-asserted-by":"publisher","first-page":"1331","DOI":"10.1136\/bmj.2.2549.1331","article-title":"The identity of British ulcerative colitis and tropical bacillary dysentery","volume":"2","author":"Hawkins","year":"1909","journal-title":"BMJ"},{"key":"2026022702152293000_B18","doi-asserted-by":"publisher","first-page":"1503","DOI":"10.1128\/jb.88.5.1503-1518.1964","article-title":"Epithelial cell penetration as an essential step in the pathogenesis of bacillary dysentery","volume":"88","author":"Labrec","year":"1964","journal-title":"J Bacteriol"},{"key":"2026022702152293000_B19","doi-asserted-by":"publisher","first-page":"1141","DOI":"10.1111\/j.1469-0691.2007.01829.x","article-title":"Increasing antimicrobial resistance\u2014an emerging problem in the treatment of shigellosis","volume":"13","author":"Niyogi","year":"2007","journal-title":"Clin Microbiol Infect"},{"key":"2026022702152293000_B20","first-page":"133","article-title":"Shigellosis","volume":"43","author":"Niyogi","year":"2005","journal-title":"J Microbiol"},{"key":"2026022702152293000_B21","doi-asserted-by":"publisher","first-page":"825","DOI":"10.1046\/j.1365-2958.2003.03347.x","article-title":"An extended role for the nucleoid structuring protein H-NS in the virulence gene regulatory cascade of Shigella flexneri","volume":"47","author":"Beloin","year":"2003","journal-title":"Mol Microbiol"},{"key":"2026022702152293000_B22","doi-asserted-by":"publisher","first-page":"951","DOI":"10.1099\/mic.0.27639-0","article-title":"Analysis of virulence plasmid gene expression defines three classes of effectors in the type III secretion system of Shigella flexneri","volume":"151","author":"Gall","year":"2005","journal-title":"Microbiology (Reading)"},{"key":"2026022702152293000_B23","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1007\/s004380050550","article-title":"Differential regulation of the plasmid-encoded genes in the Shigella flexneri virulence regulon","volume":"256","author":"Porter","year":"1997","journal-title":"Mol Gen Genet"},{"key":"2026022702152293000_B24","doi-asserted-by":"publisher","first-page":"2562","DOI":"10.1128\/JB.00212-13","article-title":"Characterization of the ospZ promoter in Shigella flexneri and its regulation by VirB and H-NS","volume":"195","author":"Basta","year":"2013","journal-title":"J Bacteriol"},{"key":"2026022702152293000_B25","doi-asserted-by":"publisher","first-page":"3403","DOI":"10.1128\/JB.01813-06","article-title":"H-NS antagonism in Shigella flexneri by VirB, a virulence gene transcription regulator that is closely related to plasmid partition factors","volume":"189","author":"Turner","year":"2007","journal-title":"J Bacteriol"},{"key":"2026022702152293000_B26","doi-asserted-by":"publisher","first-page":"699","DOI":"10.1128\/JB.186.3.699-705.2004","article-title":"Regulation of IcsP, the outer membrane protease of the Shigella actin tail assembly protein IcsA, by virulence plasmid regulators VirF and VirB","volume":"186","author":"Wing","year":"2004","journal-title":"J Bacteriol"},{"key":"2026022702152293000_B27","doi-asserted-by":"publisher","first-page":"6142","DOI":"10.1128\/jb.175.19.6142-6149.1993","article-title":"Transcriptional control of the invasion regulatory gene virB of Shigella flexneri: activation by virF and repression by H-NS","volume":"175","author":"Tobe","year":"1993","journal-title":"J Bacteriol"},{"key":"2026022702152293000_B28","doi-asserted-by":"publisher","first-page":"5158","DOI":"10.1128\/JB.185.17.5158-5165.2003","article-title":"Determination of the InvE binding site required for expression of IpaB of the Shigella sonnei virulence plasmid: involvement of a ParB boxA-like sequence","volume":"185","author":"Taniya","year":"2003","journal-title":"J Bacteriol"},{"key":"2026022702152293000_B29","doi-asserted-by":"publisher","first-page":"e0151923","DOI":"10.1128\/mbio.01519-23","article-title":"VirB, a key transcriptional regulator of Shigella virulence, requires a CTP ligand for its regulatory activities","volume":"14","author":"Gerson","year":"2023","journal-title":"mBio"},{"key":"2026022702152293000_B30","first-page":"193","article-title":"DNA-related conditions controlling the initiation of sporulation in Bacillus subtilis","volume":"40","author":"Ireton","year":"1994","journal-title":"Cell Mol Biol Res"},{"key":"2026022702152293000_B31","doi-asserted-by":"publisher","first-page":"741","DOI":"10.1046\/j.1365-2958.2001.02643.x","article-title":"The chromosome partitioning protein, ParB, is required for cytokinesis in Caulobacter crescentus","volume":"42","author":"Mohl","year":"2001","journal-title":"Mol Microbiol"},{"key":"2026022702152293000_B32","doi-asserted-by":"publisher","first-page":"102289","DOI":"10.1016\/j.mib.2023.102289","article-title":"CTP switches in ParABS-mediated bacterial chromosome segregation and beyond","volume":"73","author":"McLean","year":"2023","journal-title":"Curr Opin Microbiol"},{"key":"2026022702152293000_B33","doi-asserted-by":"publisher","first-page":"1129","DOI":"10.1126\/science.aay3965","article-title":"Self-organization of parS centromeres by the ParB CTP hydrolase","volume":"366","author":"Soh","year":"2019","journal-title":"Science"},{"key":"2026022702152293000_B34","doi-asserted-by":"publisher","first-page":"3623","DOI":"10.1016\/j.molcel.2021.06.025","article-title":"CTP regulates membrane-binding activity of the nucleoid occlusion protein Noc","volume":"81","author":"Jalal","year":"2021","journal-title":"Mol Cell"},{"key":"2026022702152293000_B35","doi-asserted-by":"publisher","first-page":"1512","DOI":"10.1016\/j.cell.2019.11.015","article-title":"ParB-type DNA segregation proteins are CTP-dependent molecular switches","volume":"179","author":"Osorio-Valeriano","year":"2019","journal-title":"Cell"},{"key":"2026022702152293000_B36","doi-asserted-by":"crossref","first-page":"e53515","DOI":"10.7554\/eLife.53515","article-title":"ParB spreading on DNA requires cytidine triphosphate in vitro","volume":"9","author":"Jalal","year":"2020","journal-title":"eLife"},{"key":"2026022702152293000_B37","doi-asserted-by":"publisher","first-page":"167401","DOI":"10.1016\/j.jmb.2021.167401","article-title":"CTP-controlled liquid\u2013liquid phase separation of ParB","volume":"434","author":"Babl","year":"2022","journal-title":"J Mol Biol"},{"key":"2026022702152293000_B38","doi-asserted-by":"crossref","first-page":"gkaf623","DOI":"10.1093\/nar\/gkaf623","article-title":"Significance of the CTP-binding motif for the interactions of S. coelicolor ParB with DNA, chromosome segregation, and sporogenic hyphal growth","volume":"53","author":"Szymczak","year":"2025","journal-title":"Nucleic Acids Res"},{"key":"2026022702152293000_B39","doi-asserted-by":"publisher","first-page":"4047","DOI":"10.1128\/JB.00313-09","article-title":"VirB alleviates H-NS repression of the icsP promoter in Shigella flexneri from sites more than one kilobase upstream of the transcription start site","volume":"191","author":"Castellanos","year":"2009","journal-title":"J Bacteriol"},{"key":"2026022702152293000_B40","doi-asserted-by":"publisher","first-page":"505","DOI":"10.1111\/mmi.13932","article-title":"Insights into transcriptional silencing and anti-silencing in Shigella flexneri: a detailed molecular analysis of the icsP virulence locus","volume":"108","author":"Weatherspoon-Griffin","year":"2018","journal-title":"Mol Microbiol"},{"key":"2026022702152293000_B41","doi-asserted-by":"publisher","first-page":"3679","DOI":"10.1093\/nar\/gkad088","article-title":"Localized modulation of DNA supercoiling, triggered by the Shigella anti-silencer VirB, is sufficient to relieve H-NS-mediated silencing","volume":"51","author":"Picker","year":"2023","journal-title":"Nucleic Acids Res"},{"key":"2026022702152293000_B42","doi-asserted-by":"publisher","first-page":"1204","DOI":"10.1038\/s42003-023-05590-8","article-title":"VirB, a transcriptional activator of virulence in Shigella flexneri, uses CTP as a cofactor","volume":"6","author":"Antar","year":"2023","journal-title":"Commun Biol"},{"key":"2026022702152293000_B43","doi-asserted-by":"publisher","first-page":"318","DOI":"10.1038\/s41467-023-44509-z","article-title":"The virulence regulator VirB from Shigella flexneri uses a CTP-dependent switch mechanism to activate gene expression","volume":"15","author":"Jakob","year":"2024","journal-title":"Nat Commun"},{"key":"2026022702152293000_B44","doi-asserted-by":"crossref","first-page":"421","DOI":"10.1007\/978-1-0716-3930-6_19","article-title":"Methods to quantitatively measure topological changes induced by DNA-binding proteins in vivo and in vitro","volume":"2819","author":"Karney","year":"2024","journal-title":"Methods Mol Biol"},{"key":"2026022702152293000_B45","doi-asserted-by":"publisher","first-page":"4121","DOI":"10.1128\/jb.177.14.4121-4130.1995","article-title":"Tight regulation, modulation, and high-level expression by vectors containing the arabinose PBAD promoter","volume":"177","author":"Guzman","year":"1995","journal-title":"J Bacteriol"},{"key":"2026022702152293000_B46","doi-asserted-by":"publisher","first-page":"10529","DOI":"10.1093\/nar\/gkt748","article-title":"Structural insights into VirB\u2013DNA complexes reveal mechanism of transcriptional activation of virulence genes","volume":"41","author":"Gao","year":"2013","journal-title":"Nucleic Acids Res"},{"key":"2026022702152293000_B47","doi-asserted-by":"publisher","first-page":"3271","DOI":"10.1128\/IAI.69.5.3271-3285.2001","article-title":"Complete DNA sequence and analysis of the large virulence plasmid of Shigella flexneri","volume":"69","author":"Venkatesan","year":"2001","journal-title":"Infect Immun"},{"key":"2026022702152293000_B48","doi-asserted-by":"publisher","first-page":"265","DOI":"10.1111\/mmi.15344","article-title":"Remote regulation by VirB, the transcriptional anti-silencer of Shigella virulence genes, provides mechanistic information","volume":"123","author":"Cris","year":"2025","journal-title":"Mol Microbiol"},{"key":"2026022702152293000_B49","doi-asserted-by":"publisher","first-page":"eabn3299","DOI":"10.1126\/sciadv.abn3299","article-title":"ParB proteins can bypass DNA-bound roadblocks via dimer-dimer recruitment","volume":"8","author":"Tisma","year":"2022","journal-title":"Sci Adv"},{"key":"2026022702152293000_B50","doi-asserted-by":"crossref","first-page":"e00627","DOI":"10.1128\/JB.00627-20","article-title":"VirB, a key transcriptional regulator of virulence plasmid genes in Shigella flexneri, forms DNA-binding site-dependent foci in the bacterial cytoplasm","volume":"203","author":"Socea","year":"2021","journal-title":"J Bacteriol"},{"key":"2026022702152293000_B51","doi-asserted-by":"publisher","first-page":"718","DOI":"10.1128\/JB.01020-07","article-title":"Characterization of nucleotide pools as a function of physiological state in Escherichia coli","volume":"190","author":"Buckstein","year":"2008","journal-title":"J Bacteriol"},{"key":"2026022702152293000_B52","doi-asserted-by":"publisher","first-page":"120161","DOI":"10.1016\/j.talanta.2019.120161","article-title":"Analysis of nucleotide pools in bacteria using HPLC-MS in HILIC mode","volume":"205","author":"Zbornikova","year":"2019","journal-title":"Talanta"},{"key":"2026022702152293000_B53","doi-asserted-by":"publisher","first-page":"e00072","DOI":"10.1128\/JB.00072-20","article-title":"The antiactivator of type III secretion, OspD1, is transcriptionally regulated by VirB and H-NS from remote sequences in Shigella flexneri","volume":"202","author":"McKenna","year":"2020","journal-title":"J Bacteriol"},{"key":"2026022702152293000_B54","doi-asserted-by":"publisher","first-page":"eabj2854","DOI":"10.1126\/sciadv.abj2854","article-title":"Relief of ParB autoinhibition by parS DNA catalysis and recycling of ParB by CTP hydrolysis promote bacterial centromere assembly","volume":"7","author":"Antar","year":"2021","journal-title":"Sci Adv"},{"key":"2026022702152293000_B55","doi-asserted-by":"publisher","first-page":"3992","DOI":"10.1016\/j.molcel.2021.09.004","article-title":"The CTPase activity of ParB determines the size and dynamics of prokaryotic DNA partition complexes","volume":"81","author":"Osorio-Valeriano","year":"2021","journal-title":"Mol Cell"},{"key":"2026022702152293000_B56","doi-asserted-by":"publisher","first-page":"e69676","DOI":"10.7554\/eLife.69676","article-title":"A CTP-dependent gating mechanism enables ParB spreading on DNA","volume":"10","author":"Jalal","year":"2021","journal-title":"eLife"},{"key":"2026022702152293000_B57","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1016\/S0014-5793(03)00524-6","article-title":"In vitro DNA-binding properties of VirB, the Shigella flexneri virulence regulatory protein","volume":"545","author":"McKenna","year":"2003","journal-title":"FEBS Lett"},{"key":"2026022702152293000_B58","doi-asserted-by":"publisher","first-page":"e2527592122","DOI":"10.1073\/pnas.2527592122","article-title":"Versatile NTP recognition and domain fusions expand the functional repertoire of the ParB-CTPase fold beyond chromosome segregation","volume":"122","author":"Kaljevic","year":"2025","journal-title":"Proc Natl Acad Sci USA"}],"container-title":["Nucleic Acids Research"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/nar\/article-pdf\/54\/5\/gkag115\/67146774\/gkag115.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/nar\/article-pdf\/54\/5\/gkag115\/67146774\/gkag115.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,27]],"date-time":"2026-02-27T07:15:32Z","timestamp":1772176532000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/nar\/article\/doi\/10.1093\/nar\/gkag115\/8500547"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,24]]},"references-count":58,"journal-issue":{"issue":"5","published-print":{"date-parts":[[2026,2,24]]}},"URL":"https:\/\/doi.org\/10.1093\/nar\/gkag115","relation":{},"ISSN":["0305-1048","1362-4962"],"issn-type":[{"value":"0305-1048","type":"print"},{"value":"1362-4962","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,3,24]]},"published":{"date-parts":[[2026,2,24]]},"article-number":"gkag115"}}