{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T01:39:25Z","timestamp":1791337165996,"version":"4.3.1"},"reference-count":77,"publisher":"Oxford University Press (OUP)","issue":"4","license":[{"start":{"date-parts":[[2024,1,31]],"date-time":"2024-01-31T00:00:00Z","timestamp":1706659200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100008398","name":"VILLUM FONDEN","doi-asserted-by":"publisher","award":["00028321"],"award-info":[{"award-number":["00028321"]}],"id":[{"id":"10.13039\/100008398","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100008398","name":"VILLUM FONDEN","doi-asserted-by":"publisher","award":["00050284"],"award-info":[{"award-number":["00050284"]}],"id":[{"id":"10.13039\/100008398","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2024,2,28]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Many bacteria form biofilms to protect themselves from predators or stressful environmental conditions. In the biofilm, bacteria are embedded in a protective extracellular matrix composed of polysaccharides, proteins and extracellular DNA (eDNA). eDNA most often is released from lysed bacteria or host mammalian cells, and it is the only matrix component most biofilms appear to have in common. However, little is known about the form DNA takes in the extracellular space, and how different non-canonical DNA structures such as Z-DNA or G-quadruplexes might contribute to its function in the biofilm. The aim of this study was to determine if non-canonical DNA structures form in eDNA-rich staphylococcal biofilms, and if these structures protect the biofilm from degradation by nucleases. We grew Staphylococcus epidermidis biofilms in laboratory media supplemented with hemin and NaCl to stabilize secondary DNA structures and visualized their location by immunolabelling and fluorescence microscopy. We furthermore visualized the macroscopic biofilm structure by optical coherence tomography. We developed assays to quantify degradation of Z-DNA and G-quadruplex DNA oligos by different nucleases, and subsequently investigated how these enzymes affected eDNA in the biofilms.<\/jats:p>\n                  <jats:p>Z-DNA and G-quadruplex DNA were abundant in the biofilm matrix, and were often present in a web-like structures. In vitro, the structures did not form in the absence of NaCl or mechanical shaking during biofilm growth, or in bacterial strains deficient in eDNA or exopolysaccharide production. We thus infer that eDNA and polysaccharides interact, leading to non-canonical DNA structures under mechanical stress when stabilized by salt. We also confirmed that G-quadruplex DNA and Z-DNA was present in biofilms from infected implants in a murine implant-associated osteomyelitis model. Mammalian DNase I lacked activity against Z-DNA and G-quadruplex DNA, while Micrococcal nuclease could degrade G-quadruplex DNA and S1 Aspergillus nuclease could degrade Z-DNA. Micrococcal nuclease, which originates from Staphylococcus aureus, may thus be key for dispersal of biofilm in staphylococci. In addition to its structural role, we show for the first time that the eDNA in biofilms forms a DNAzyme with peroxidase-like activity in the presence of hemin. While peroxidases are part of host defenses against pathogens, we now show that biofilms can possess intrinsic peroxidase activity in the extracellular matrix.<\/jats:p>","DOI":"10.1093\/nar\/gkae034","type":"journal-article","created":{"date-parts":[[2024,1,10]],"date-time":"2024-01-10T09:45:43Z","timestamp":1704879943000},"page":"1575-1590","source":"Crossref","is-referenced-by-count":55,"title":["Extracellular G-quadruplexes and Z-DNA protect biofilms from DNase I, and G-quadruplexes form a DNAzyme with peroxidase activity"],"prefix":"10.1093","volume":"52","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4946-0337","authenticated-orcid":false,"given":"Gabriel Antonio\u00a0Salvador","family":"Minero","sequence":"first","affiliation":[{"name":"Interdisciplinary Nanoscience Center (iNANO), Aarhus University , Gustav Wieds Vej 14 , 8000\u00a0 Aarhus , Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andreas","family":"M\u00f8llebjerg","sequence":"additional","affiliation":[{"name":"Interdisciplinary Nanoscience Center (iNANO), Aarhus University , Gustav Wieds Vej 14 , 8000\u00a0 Aarhus , Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Celine","family":"Thiesen","sequence":"additional","affiliation":[{"name":"Interdisciplinary Nanoscience Center (iNANO), Aarhus University , Gustav Wieds Vej 14 , 8000\u00a0 Aarhus , Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mikkel\u00a0Illemann","family":"Johansen","sequence":"additional","affiliation":[{"name":"Department Infectious Diseases, Aarhus University Hospital , Palle Juul-Jensens bvld 99 , 8200\u00a0 Aarhus N , Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nis\u00a0Pedersen","family":"J\u00f8rgensen","sequence":"additional","affiliation":[{"name":"Department Infectious Diseases, Aarhus University Hospital , Palle Juul-Jensens bvld 99 , 8200\u00a0 Aarhus N , Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1360-8870","authenticated-orcid":false,"given":"Victoria","family":"Birkedal","sequence":"additional","affiliation":[{"name":"Interdisciplinary Nanoscience Center (iNANO), Aarhus University , Gustav Wieds Vej 14 , 8000\u00a0 Aarhus , Denmark"},{"name":"Department of Chemistry, Aarhus University , Langelandsgade 140 , 8000\u00a0 Aarhus , Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daniel Erik","family":"Otzen","sequence":"additional","affiliation":[{"name":"Interdisciplinary Nanoscience Center (iNANO), Aarhus University , Gustav Wieds Vej 14 , 8000\u00a0 Aarhus , Denmark"},{"name":"Department of Molecular Biology and Genetics, Aarhus University , Universitetsbyen 81 , 8000\u00a0 Aarhus , Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6485-5134","authenticated-orcid":false,"given":"Rikke\u00a0Louise","family":"Meyer","sequence":"additional","affiliation":[{"name":"Interdisciplinary Nanoscience Center (iNANO), Aarhus University , Gustav Wieds Vej 14 , 8000\u00a0 Aarhus , Denmark"},{"name":"Department of Biology, Aarhus University , Ny Munkegade 114 , 8000\u00a0 Aarhus , Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2024,1,31]]},"reference":[{"key":"2024022807094785300_B1","doi-asserted-by":"crossref","first-page":"1198","DOI":"10.1056\/NEJMoa1306801","article-title":"Multistate point- prevalence survey of health care- associated infections","volume":"370","author":"Magill","year":"2014","journal-title":"N. Engl. J. Med."},{"key":"2024022807094785300_B2","doi-asserted-by":"crossref","first-page":"1187","DOI":"10.1086\/605694","article-title":"Management of the catheter in documented catheter-related coagulase-negative staphylococcal bacteremia: remove or retain?","volume":"49","author":"Raad","year":"2009","journal-title":"Clin. Infect. Dis."},{"key":"2024022807094785300_B3","doi-asserted-by":"crossref","first-page":"447","DOI":"10.7326\/0003-4819-98-4-447","article-title":"Staphylococcus epidermidis causing prosthetic valve endocarditis: microbiologic and clinical observations as guides to therapy","volume":"98","author":"Karchmer","year":"1983","journal-title":"Ann. Intern. Med."},{"key":"2024022807094785300_B4","doi-asserted-by":"crossref","first-page":"498","DOI":"10.1016\/j.jacep.2016.01.019","article-title":"Microbiology of cardiac implantable electronic device infections","volume":"2","author":"Hussein","year":"2016","journal-title":"JACC Clin. Electrophysiol."},{"key":"2024022807094785300_B5","doi-asserted-by":"crossref","first-page":"ofac048","DOI":"10.1093\/ofid\/ofac048","article-title":"Predictors of treatment success after periprosthetic joint infection: 24-month follow up from a multicenter prospective observational cohort study of 653 patients","volume":"9","author":"Davis","year":"2022","journal-title":"Open Forum Infect. Dis."},{"key":"2024022807094785300_B6","doi-asserted-by":"crossref","first-page":"40","DOI":"10.3390\/antibiotics12010040","article-title":"The clinical outcome of early periprosthetic joint infections caused by Staphylococcus epidermidis and managed by surgical debridement in an era of increasing resistance","volume":"12","author":"Shabana","year":"2023","journal-title":"Antibiotics"},{"key":"2024022807094785300_B7","first-page":"14","article-title":"Structural basis of Staphylococcus epidermidis biofilm formation: mechanisms and molecular interactions","volume":"5","author":"Buttner","year":"2015","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"2024022807094785300_B8","doi-asserted-by":"crossref","first-page":"1711","DOI":"10.1016\/j.biomaterials.2006.11.046","article-title":"Polysaccharide intercellular adhesin or protein factors in biofilm accumulation of Staphylococcus epidermidis and Staphylococcus aureus isolated from prosthetic hip and knee joint infections","volume":"28","author":"Rohde","year":"2007","journal-title":"Biomaterials"},{"key":"2024022807094785300_B9","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1186\/s12866-017-0930-9","article-title":"Methicillin resistance and virulence genes in invasive and nasal Staphylococcus epidermidis isolates from neonates","volume":"17","author":"Salgueiro","year":"2017","journal-title":"BMC Microbiol."},{"key":"2024022807094785300_B10","doi-asserted-by":"crossref","first-page":"1579","DOI":"10.1099\/jmm.0.055210-0","article-title":"Staphylococcus epidermidis with the icaA(-)\/icaD(-)\/IS256(-) genotype and protein or protein\/extracellular-DNA biofilm is frequent in ocular infections","volume":"62","author":"Juarez-Verdayes","year":"2013","journal-title":"J. Med. Microbiol."},{"key":"2024022807094785300_B11","doi-asserted-by":"crossref","first-page":"2083","DOI":"10.1099\/mic.0.2007\/006031-0","article-title":"Role of autolysin-mediated DNA release in biofilm formation of Staphylococcus epidermidis","volume":"153","author":"Qin","year":"2007","journal-title":"Microbiology (Reading)"},{"key":"2024022807094785300_B12","doi-asserted-by":"crossref","first-page":"840","DOI":"10.5301\/ijao.5000057","article-title":"Biofilm extracellular-DNA in 55 Staphylococcus epidermidis clinical isolates from implant infections","volume":"34","author":"Ravaioli","year":"2011","journal-title":"Int. J. Artif. Organs"},{"key":"2024022807094785300_B13","doi-asserted-by":"crossref","first-page":"341","DOI":"10.3109\/1040841X.2013.841639","article-title":"The role of extracellular DNA in the establishment, maintenance and perpetuation of bacterial biofilms","volume":"41","author":"Okshevsky","year":"2015","journal-title":"Crit. Rev. Microbiol."},{"key":"2024022807094785300_B14","doi-asserted-by":"crossref","first-page":"1487","DOI":"10.1126\/science.295.5559.1487","article-title":"Extracellular DNA required for bacterial biofilm formation","volume":"295","author":"Whitchurch","year":"2002","journal-title":"Science"},{"key":"2024022807094785300_B15","doi-asserted-by":"crossref","first-page":"150","DOI":"10.1016\/j.jconrel.2015.04.028","article-title":"Disassembling bacterial extracellular matrix with DNase-coated nanoparticles to enhance antibiotic delivery in biofilm infections","volume":"209","author":"Baelo","year":"2015","journal-title":"J Control Release"},{"key":"2024022807094785300_B16","doi-asserted-by":"crossref","first-page":"1428","DOI":"10.3389\/fmicb.2015.01428","article-title":"DNase-sensitive and -resistant modes of biofilm formation by Listeria monocytogenes","volume":"6","author":"Zetzmann","year":"2015","journal-title":"Front. Microbiol."},{"key":"2024022807094785300_B17","doi-asserted-by":"crossref","first-page":"5740","DOI":"10.1016\/j.cell.2021.10.010","article-title":"Z-form extracellular DNA is a structural component of the bacterial biofilm matrix","volume":"184","author":"Buzzo","year":"2021","journal-title":"Cell"},{"key":"2024022807094785300_B18","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1038\/s41522-021-00197-5","article-title":"The biofilm matrix scaffold of Pseudomonas aeruginosa contains G-quadruplex extracellular DNA structures","volume":"7","author":"Seviour","year":"2021","journal-title":"npj Biofilms Microbiomes"},{"key":"2024022807094785300_B19","doi-asserted-by":"crossref","first-page":"7772","DOI":"10.1038\/s41467-023-43533-3","article-title":"RNA is a key component of extracellular DNA networks in Pseudomonas aeruginosa biofilms","volume":"14","author":"Mugunthan","year":"2023","journal-title":"Nat. Commun"},{"key":"2024022807094785300_B20","doi-asserted-by":"crossref","first-page":"466","DOI":"10.3390\/microorganisms10020466","article-title":"Bacterial biofilms utilize an underlying extracellular DNA matrix structure that can be targeted for biofilm resolution","volume":"10","author":"Goodman","year":"2022","journal-title":"Microorganisms"},{"key":"2024022807094785300_B21","doi-asserted-by":"crossref","first-page":"9340","DOI":"10.1074\/jbc.271.16.9340","article-title":"Analysis of left-handed Z-DNA formation in short d(CG)n sequences in Escherichia coli and Halobacterium halobium plasmids. Stabilization by increasing repeat length and DNA supercoiling but not salinity","volume":"271","author":"Kim","year":"1996","journal-title":"J. Biol. Chem."},{"key":"2024022807094785300_B22","doi-asserted-by":"crossref","first-page":"358","DOI":"10.1126\/science.2678475","article-title":"Stabilization of Z DNA in vivo by localized supercoiling","volume":"246","author":"Rahmouni","year":"1989","journal-title":"Science"},{"key":"2024022807094785300_B23","doi-asserted-by":"crossref","first-page":"11388","DOI":"10.1021\/bi00112a005","article-title":"Structure of the pure-spermine form of Z-DNA (magnesium free) at 1-A resolution","volume":"30","author":"Egli","year":"1991","journal-title":"Biochemistry"},{"key":"2024022807094785300_B24","doi-asserted-by":"crossref","first-page":"4006","DOI":"10.1002\/anie.200501149","article-title":"A cationic zinc porphyrin as a chiroptical probe for Z-DNA","volume":"44","author":"Balaz","year":"2005","journal-title":"Angew. Chem. Int. Ed Engl."},{"key":"2024022807094785300_B25","doi-asserted-by":"crossref","first-page":"9943","DOI":"10.1038\/srep09943","article-title":"Formation of poly(d(A-T)2) specific Z-DNA by a cationic porphyrin","volume":"5","author":"Jang","year":"2015","journal-title":"Sci. Rep."},{"key":"2024022807094785300_B26","doi-asserted-by":"crossref","first-page":"e0198418","DOI":"10.1371\/journal.pone.0198418","article-title":"C2G4)n repeat expansion sequences from the C9orf72 gene form an unusual DNA higher-order structure in the pH range of 5-6","volume":"13","author":"Lat","year":"2018","journal-title":"PLoS One"},{"key":"2024022807094785300_B27","doi-asserted-by":"crossref","first-page":"e35558","DOI":"10.1371\/journal.pone.0035558","article-title":"The role of methylation in the intrinsic dynamics of B- and Z-DNA","volume":"7","author":"Temiz","year":"2012","journal-title":"PLoS One"},{"key":"2024022807094785300_B28","doi-asserted-by":"crossref","first-page":"2198","DOI":"10.1039\/C8OB00030A","article-title":"C8-Guanine modifications: effect on Z-DNA formation and its role in cancer","volume":"16","author":"Vongsutilers","year":"2018","journal-title":"Org. Biomol. Chem."},{"key":"2024022807094785300_B29","doi-asserted-by":"crossref","first-page":"e0057720","DOI":"10.1128\/JB.00577-20","article-title":"G-quadruplex structures in bacteria: biological relevance and potential as an antimicrobial target","volume":"203","author":"Yadav","year":"2021","journal-title":"J. Bacteriol."},{"key":"2024022807094785300_B30","doi-asserted-by":"crossref","first-page":"16405","DOI":"10.1021\/ja045154j","article-title":"Loop-length-dependent folding of G-quadruplexes","volume":"126","author":"Hazel","year":"2004","journal-title":"J. Am. Chem. Soc."},{"key":"2024022807094785300_B31","doi-asserted-by":"crossref","first-page":"4254","DOI":"10.1039\/D0AN00791A","article-title":"Characterization of intermolecular G-quadruplex formation over intramolecular G-triplex for DNA containing three G-tracts","volume":"145","author":"Zhang","year":"2020","journal-title":"Analyst"},{"key":"2024022807094785300_B32","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1007\/978-1-61779-142-0_7","article-title":"G-quartet, G-quadruplex, and G-wire regulated by chemical stimuli","volume":"749","author":"Miyoshi","year":"2011","journal-title":"Methods Mol. Biol."},{"key":"2024022807094785300_B33","doi-asserted-by":"crossref","first-page":"1959","DOI":"10.1038\/s41467-018-04016-y","article-title":"High-resolution AFM structure of DNA G-wires in aqueous solution","volume":"9","author":"Bose","year":"2018","journal-title":"Nat. Commun."},{"key":"2024022807094785300_B34","doi-asserted-by":"crossref","first-page":"38","DOI":"10.3389\/fchem.2016.00038","article-title":"Metal cations in G-quadruplex folding and stability","volume":"4","author":"Bhattacharyya","year":"2016","journal-title":"Front Chem"},{"key":"2024022807094785300_B35","doi-asserted-by":"crossref","first-page":"1883","DOI":"10.1021\/acs.jctc.0c01176","article-title":"Insights into G-quadruplex-hemin dynamics using atomistic simulations: implications for reactivity and folding","volume":"17","author":"Stadlbauer","year":"2021","journal-title":"J. Chem. Theory Comput."},{"key":"2024022807094785300_B36","doi-asserted-by":"crossref","first-page":"2390","DOI":"10.1021\/acs.biochem.2c00478","article-title":"8-Oxoguanine forms quartets with a large Central cavity","volume":"61","author":"Aleksic","year":"2022","journal-title":"Biochemistry"},{"key":"2024022807094785300_B37","doi-asserted-by":"crossref","first-page":"8038","DOI":"10.1093\/nar\/gky735","article-title":"i-motif DNA: structural features and significance to cell biology","volume":"46","author":"Abou\u00a0Assi","year":"2018","journal-title":"Nucleic Acids Res."},{"key":"2024022807094785300_B38","doi-asserted-by":"crossref","first-page":"6292","DOI":"10.1073\/pnas.85.17.6292","article-title":"Intramolecular DNA triplexes in supercoiled plasmids","volume":"85","author":"Hanvey","year":"1988","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"2024022807094785300_B39","doi-asserted-by":"crossref","first-page":"4455","DOI":"10.1021\/bi00232a012","article-title":"Polyamines favor DNA triplex formation at neutral pH","volume":"30","author":"Hampel","year":"1991","journal-title":"Biochemistry"},{"key":"2024022807094785300_B40","doi-asserted-by":"crossref","first-page":"52321","DOI":"10.1039\/C5RA90057C","article-title":"Electronic pH switching of DNA triplex reactions (vol 5, pg 27313, 2015)","volume":"5","author":"Minero","year":"2015","journal-title":"RSC Adv."},{"key":"2024022807094785300_B41","first-page":"10126","article-title":"Intrastrand triplex DNA repeats in bacteria: a source of genomic instability","volume":"43","author":"Holder","year":"2015","journal-title":"Nucleic Acids Res."},{"key":"2024022807094785300_B42","doi-asserted-by":"crossref","first-page":"e00450-17","DOI":"10.1128\/genomeA.00450-17","article-title":"Complete genome sequence of Staphylococcus epidermidis 1457","volume":"5","author":"Galac","year":"2017","journal-title":"Genome Announc."},{"key":"2024022807094785300_B43","doi-asserted-by":"crossref","first-page":"e01254-20","DOI":"10.1128\/MRA.01254-20","article-title":"Genome sequence of Staphylococcus epidermidis AUH4567, a clinical isolate from an infected Central venous catheter","volume":"10","author":"Meyer","year":"2021","journal-title":"Microbiol. Resour. Announc."},{"key":"2024022807094785300_B44","doi-asserted-by":"crossref","first-page":"e103688","DOI":"10.1371\/journal.pone.0103688","article-title":"A modified chronic infection model for testing treatment of Staphylococcus aureus biofilms on implants","volume":"9","author":"J\u00f8rgensen","year":"2014","journal-title":"PLoS One"},{"key":"2024022807094785300_B45","doi-asserted-by":"crossref","first-page":"200","DOI":"10.1111\/j.1462-2920.2005.00880.x","article-title":"daime, a novel image analysis program for microbial ecology and biofilm research","volume":"8","author":"Daims","year":"2006","journal-title":"Environ. Microbiol."},{"key":"2024022807094785300_B46","doi-asserted-by":"crossref","first-page":"116485","DOI":"10.1016\/j.desal.2023.116485","article-title":"Novel high-throughput screening platform identifies enzymes to tackle biofouling on reverse osmosis membranes","volume":"554","author":"M\u00f8llebjerg","year":"2023","journal-title":"Desalination"},{"key":"2024022807094785300_B47","doi-asserted-by":"crossref","first-page":"e2113723119","DOI":"10.1073\/pnas.2113723119","article-title":"The structural role of bacterial eDNA in the formation of biofilm streamers","volume":"119","author":"Secchi","year":"2022","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"2024022807094785300_B48","doi-asserted-by":"crossref","first-page":"13137","DOI":"10.1021\/jacs.2c02466","article-title":"DNA bending force facilitates Z-DNA formation under physiological salt conditions","volume":"144","author":"Yi","year":"2022","journal-title":"J. Am. Chem. Soc."},{"key":"2024022807094785300_B49","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1007\/s00775-022-01973-0","article-title":"DNA G-quadruplex-stabilizing metal complexes as anticancer drugs","volume":"28","author":"Zegers","year":"2023","journal-title":"J. Biol. Inorg. Chem."},{"key":"2024022807094785300_B50","doi-asserted-by":"crossref","first-page":"1681","DOI":"10.1016\/j.chembiol.2019.10.003","article-title":"G-quadruplexes sequester free heme in living cells","volume":"26","author":"Gray","year":"2019","journal-title":"Cell Chem Biol"},{"key":"2024022807094785300_B51","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1111\/j.1574-6968.2002.tb11432.x","article-title":"Regulation of icaR gene expression in Staphylococcus epidermidis","volume":"216","author":"Conlon","year":"2002","journal-title":"FEMS Microbiol. Lett."},{"key":"2024022807094785300_B52","doi-asserted-by":"crossref","first-page":"150","DOI":"10.3389\/fmicb.2011.00150","article-title":"Pseudomonas aeruginosa genomic structure and diversity","volume":"2","author":"Klockgether","year":"2011","journal-title":"Front. Microbiol."},{"key":"2024022807094785300_B53","doi-asserted-by":"crossref","first-page":"jmm001287","DOI":"10.1099\/jmm.0.001287","article-title":"Host factors abolish the need for polysaccharides and extracellular matrix-binding protein in Staphylococcus epidermidis biofilm formation","volume":"70","author":"Skovdal","year":"2021","journal-title":"J. Med. Microbiol."},{"key":"2024022807094785300_B54","doi-asserted-by":"crossref","first-page":"11353","DOI":"10.1073\/pnas.1503058112","article-title":"Pel is a cationic exopolysaccharide that cross-links extracellular DNA in the Pseudomonas aeruginosa biofilm matrix","volume":"112","author":"Jennings","year":"2015","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"2024022807094785300_B55","doi-asserted-by":"crossref","first-page":"5254","DOI":"10.1093\/nar\/gkaa281","article-title":"High specificity and tight spatial restriction of self-biotinylation by DNA and RNA G-quadruplexes complexed in vitro and in vivo with Heme","volume":"48","author":"Lat","year":"2020","journal-title":"Nucleic Acids Res."},{"key":"2024022807094785300_B56","doi-asserted-by":"crossref","first-page":"7817","DOI":"10.1021\/bi9006786","article-title":"General peroxidase activity of G-quadruplex-Hemin complexes and its application in ligand screening","volume":"48","author":"Cheng","year":"2009","journal-title":"Biochemistry"},{"key":"2024022807094785300_B57","doi-asserted-by":"crossref","first-page":"1311","DOI":"10.1002\/jor.22907","article-title":"Quantifying the natural history of biofilm formation in vivo during the establishment of chronic implant-associated Staphylococcus aureus osteomyelitis in mice to identify critical pathogen and host factors","volume":"33","author":"Nishitani","year":"2015","journal-title":"J. Orthop. Res."},{"key":"2024022807094785300_B58","first-page":"85","volume-title":"Toxicoepigenetics","author":"Davis","year":"2019"},{"key":"2024022807094785300_B59","doi-asserted-by":"crossref","first-page":"3481","DOI":"10.1093\/nar\/6.11.3481","article-title":"Rapid RNA sequencing: nucleases from Staphylococcus aureus and Neurospora crassa discriminate between uridine and cytidine","volume":"6","author":"Krupp","year":"1979","journal-title":"Nucleic Acids Res."},{"key":"2024022807094785300_B60","doi-asserted-by":"crossref","first-page":"2431","DOI":"10.1021\/ac303440d","article-title":"G-quadruplex-based fluorescent assay of S1 nuclease activity and K+","volume":"85","author":"Zhou","year":"2013","journal-title":"Anal. Chem."},{"key":"2024022807094785300_B61","doi-asserted-by":"crossref","first-page":"1343","DOI":"10.1016\/j.bbagen.2016.03.027","article-title":"Heme interacts with histidine- and tyrosine-based protein motifs and inhibits enzymatic activity of chloramphenicol acetyltransferase from Escherichia coli","volume":"1860","author":"Brewitz","year":"2016","journal-title":"Biochim. Biophys. Acta"},{"key":"2024022807094785300_B62","doi-asserted-by":"crossref","first-page":"3278","DOI":"10.1111\/febs.14192","article-title":"Characterization of plasma labile heme in hemolytic conditions","volume":"284","author":"Gouveia","year":"2017","journal-title":"FEBS J."},{"key":"2024022807094785300_B63","doi-asserted-by":"crossref","first-page":"e111168","DOI":"10.1371\/journal.pone.0111168","article-title":"The Porphyromonas gingivalis ferric uptake regulator orthologue binds hemin and regulates hemin-responsive biofilm development","volume":"9","author":"Butler","year":"2014","journal-title":"PLoS One"},{"key":"2024022807094785300_B64","doi-asserted-by":"crossref","first-page":"470","DOI":"10.1128\/AEM.02073-07","article-title":"Differential roles of poly-N-acetylglucosamine surface polysaccharide and extracellular DNA in Staphylococcus aureus and Staphylococcus epidermidis biofilms","volume":"74","author":"Izano","year":"2008","journal-title":"Appl. Environ. Microb."},{"key":"2024022807094785300_B65","doi-asserted-by":"crossref","first-page":"1083","DOI":"10.1111\/j.1365-2958.1996.tb02548.x","article-title":"Molecular basis of intercellular adhesion in the biofilm-forming Staphylococcus epidermidis","volume":"20","author":"Heilmann","year":"1996","journal-title":"Mol. Microbiol."},{"key":"2024022807094785300_B66","doi-asserted-by":"crossref","first-page":"884065","DOI":"10.3389\/fcimb.2022.884065","article-title":"Phenol-soluble modulins from Staphylococcus aureus biofilms form complexes with DNA to drive autoimmunity","volume":"12","author":"Grando","year":"2022","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"2024022807094785300_B67","doi-asserted-by":"crossref","first-page":"4985","DOI":"10.1073\/pnas.0911528107","article-title":"Minute negative superhelicity is sufficient to induce the B-Z transition in the presence of low tension","volume":"107","author":"Lee","year":"2010","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"2024022807094785300_B68","doi-asserted-by":"crossref","first-page":"12069","DOI":"10.1093\/nar\/gkx856","article-title":"The effects of DNA supercoiling on G-quadruplex formation","volume":"45","author":"Sekibo","year":"2017","journal-title":"Nucleic Acids Res."},{"key":"2024022807094785300_B69","doi-asserted-by":"crossref","first-page":"860","DOI":"10.1093\/nar\/gkt957","article-title":"Detection of G-quadruplex DNA in mammalian cells","volume":"42","author":"Henderson","year":"2014","journal-title":"Nucleic Acids Res."},{"key":"2024022807094785300_B70","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1038\/nchem.1805","article-title":"Visualization and selective chemical targeting of RNA G-quadruplex structures in the cytoplasm of human cells","volume":"6","author":"Biffi","year":"2014","journal-title":"Nat. Chem."},{"key":"2024022807094785300_B71","doi-asserted-by":"crossref","first-page":"3546","DOI":"10.1073\/pnas.78.6.3546","article-title":"Antibodies specific for left-handed Z-DNA","volume":"78","author":"Lafer","year":"1981","journal-title":"Proc. Natl. Acad. Sci.-Biol."},{"key":"2024022807094785300_B72","doi-asserted-by":"crossref","first-page":"7758","DOI":"10.1021\/jacs.8b13855","article-title":"Topologically constrained formation of stable Z-DNA from normal sequence under physiological conditions","volume":"141","author":"Zhang","year":"2019","journal-title":"J. Am. Chem. Soc."},{"key":"2024022807094785300_B73","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1111\/j.1574-6976.2003.tb00626.x","article-title":"Single-strand-specific nucleases","volume":"26","author":"Desai","year":"2003","journal-title":"FEMS Microbiol. Rev."},{"key":"2024022807094785300_B74","doi-asserted-by":"crossref","first-page":"104116","DOI":"10.1039\/C5RA20773H","article-title":"Simple and convenient G-quadruplex-based fluorescence assay of micrococcal nuclease activity","volume":"5","author":"He","year":"2015","journal-title":"RSC Adv."},{"key":"2024022807094785300_B75","doi-asserted-by":"crossref","first-page":"114281","DOI":"10.1016\/j.bcp.2020.114281","article-title":"Antimicrobials offered from nature: peroxidase-catalyzed systems and their mimics","volume":"182","author":"Tonoyan","year":"2020","journal-title":"Biochem. Pharmacol."},{"key":"2024022807094785300_B76","doi-asserted-by":"crossref","first-page":"1803","DOI":"10.1093\/nar\/gkab007","article-title":"A heme*DNAzyme activated by hydrogen peroxide catalytically oxidizes thioethers by direct oxygen atom transfer rather than by a Compound I-like intermediate","volume":"49","author":"Shumayrikh","year":"2021","journal-title":"Nucleic Acids Res."},{"key":"2024022807094785300_B77","doi-asserted-by":"crossref","first-page":"919","DOI":"10.1016\/j.cell.2020.07.006","article-title":"Extracellular DNA promotes efficient extracellular electron transfer by Pyocyanin in biofilms","volume":"182","author":"Saunders","year":"2020","journal-title":"Cell"}],"container-title":["Nucleic Acids Research"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/nar\/article-pdf\/52\/4\/1575\/56763241\/gkae034.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/nar\/article-pdf\/52\/4\/1575\/56763241\/gkae034.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,2,28]],"date-time":"2024-02-28T02:17:13Z","timestamp":1709086633000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/nar\/article\/52\/4\/1575\/7595402"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,1,31]]},"references-count":77,"journal-issue":{"issue":"4","published-online":{"date-parts":[[2024,1,31]]},"published-print":{"date-parts":[[2024,2,28]]}},"URL":"https:\/\/doi.org\/10.1093\/nar\/gkae034","relation":{},"ISSN":["0305-1048","1362-4962"],"issn-type":[{"value":"0305-1048","type":"print"},{"value":"1362-4962","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2024,2,28]]},"published":{"date-parts":[[2024,1,31]]}}}