{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,15]],"date-time":"2026-07-15T23:56:48Z","timestamp":1784159808027,"version":"3.55.0"},"reference-count":41,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,8,11]],"date-time":"2025-08-11T00:00:00Z","timestamp":1754870400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2025,8,11]],"date-time":"2025-08-11T00:00:00Z","timestamp":1754870400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["BMC Microbiol"],"DOI":"10.1186\/s12866-025-04235-6","type":"journal-article","created":{"date-parts":[[2025,8,11]],"date-time":"2025-08-11T03:04:30Z","timestamp":1754881470000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":5,"title":["ST11 carbapenem-resistant Klebsiella pneumoniae integrates virulence plasmid fragments into the chromosome via insertion sequence"],"prefix":"10.1186","volume":"25","author":[{"given":"Xu","family":"Liu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lijuan","family":"Xu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Huiyue","family":"Dong","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shangshang","family":"Qin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yan","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hong","family":"Yao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,8,11]]},"reference":[{"key":"4235_CR1","doi-asserted-by":"publisher","first-page":"785","DOI":"10.1038\/s41467-025-56095-3","volume":"16","author":"CL Holmes","year":"2025","unstructured":"Holmes CL, Dailey KG, Hullahalli K, Wilcox AE, Mason S, Moricz BS, et al. Patterns of Klebsiella pneumoniae bacteremic dissemination from the lung. Nat Commun. 2025;16:785.","journal-title":"Nat Commun"},{"key":"4235_CR2","doi-asserted-by":"publisher","first-page":"2658","DOI":"10.1016\/j.scib.2023.09.040","volume":"68","author":"D Pu","year":"2023","unstructured":"Pu D, Zhao J, Chang K, Zhuo X, Cao B. Superbugs with hypervirulence and carbapenem resistance in Klebsiella pneumoniae: the rise of such emerging nosocomial pathogens in China. Sci Bull (Beijing). 2023;68:2658\u201370.","journal-title":"Sci Bull (Beijing)"},{"key":"4235_CR3","doi-asserted-by":"publisher","first-page":"65","DOI":"10.1016\/j.tim.2020.04.012","volume":"29","author":"X Yang","year":"2021","unstructured":"Yang X, Dong N, Chan EW, Zhang R, Chen S. Carbapenem resistance-encoding and virulence-encoding conjugative plasmids in Klebsiella pneumoniae. Trends Microbiol. 2021;29:65\u201383.","journal-title":"Trends Microbiol"},{"key":"4235_CR4","doi-asserted-by":"publisher","unstructured":"Russo TA, Marr CM. Hypervirulent Klebsiella pneumoniae. Clin Microbiol Rev. 2019. https:\/\/doi.org\/10.1128\/CMR.00001-19.","DOI":"10.1128\/CMR.00001-19"},{"key":"4235_CR5","doi-asserted-by":"publisher","DOI":"10.1016\/j.micres.2024.127839","author":"TY Lei","year":"2024","unstructured":"Lei TY, Liao BB, Yang LR, Wang Y, Chen XB. Hypervirulent and carbapenem-resistant Klebsiella pneumoniae: a global public health threat. Microbiol Res. 2024. https:\/\/doi.org\/10.1016\/j.micres.2024.127839.","journal-title":"Microbiol Res"},{"key":"4235_CR6","doi-asserted-by":"publisher","first-page":"101124","DOI":"10.1016\/j.drup.2024.101124","volume":"77","author":"C Liu","year":"2024","unstructured":"Liu C, Guo J, Fan S, Guo W, Qi H, Baker S, et al. An increased prevalence of carbapenem-resistant hypervirulent Klebsiella pneumoniae associated with the COVID-19 pandemic. Drug Resist Updat. 2024;77:101124.","journal-title":"Drug Resist Updat"},{"key":"4235_CR7","doi-asserted-by":"publisher","first-page":"603","DOI":"10.1128\/iai.54.3.603-608.1986","volume":"54","author":"X Nassif","year":"1986","unstructured":"Nassif X, Sansonetti PJ. Correlation of the virulence of Klebsiella pneumoniae K1 and K2 with the presence of a plasmid encoding aerobactin. Infect Immun. 1986;54:603\u20138.","journal-title":"Infect Immun"},{"issue":"10","key":"4235_CR8","doi-asserted-by":"publisher","first-page":"e621","DOI":"10.1016\/S1473-3099(24)00610-8","volume":"24","author":"M Das","year":"2024","unstructured":"Das M. Global update on hypervirulent Klebsiella pneumoniae. Lancet Infect Dis. 2024;24:e621.https:\/\/doi.org\/10.1016\/S1473-3099(24)00610-8","journal-title":"Lancet Infect Dis"},{"key":"4235_CR9","doi-asserted-by":"publisher","first-page":"825","DOI":"10.1080\/14737159.2019.1649145","volume":"19","author":"YC Lai","year":"2019","unstructured":"Lai YC, Lu MC, Hsueh PR. Hypervirulence and carbapenem resistance: two distinct evolutionary directions that led high-risk Klebsiella pneumoniae clones to epidemic success. Expert Rev Mol Diagn. 2019;19:825\u201337.","journal-title":"Expert Rev Mol Diagn"},{"key":"4235_CR10","doi-asserted-by":"publisher","first-page":"101137","DOI":"10.1016\/j.drup.2024.101137","volume":"77","author":"XM Jia","year":"2024","unstructured":"Jia XM, Zhu Y, Jia PY, Li CD, Chu XB, Sun TS, et al. The key role of iroBCDN-lacking pLVPK-like plasmid in the evolution of the most prevalent hypervirulent carbapenem-resistant ST11-KL64 Klebsiella pneumoniae in China. Drug Resist Update. 2024;77:101137.","journal-title":"Drug Resist Update."},{"key":"4235_CR11","doi-asserted-by":"publisher","DOI":"10.1128\/mSystems.00140-20","author":"X Yang","year":"2020","unstructured":"Yang X, Ye L, Chan EW, Zhang R, Chen S. Tracking recombination events that occur in conjugative virulence plasmid p15WZ-82_vir during the transmission process. mSystems. 2020. https:\/\/doi.org\/10.1128\/mSystems.00140-20.","journal-title":"mSystems"},{"key":"4235_CR12","doi-asserted-by":"publisher","first-page":"4187","DOI":"10.1038\/s41467-024-48206-3","volume":"15","author":"Q Nguyen","year":"2024","unstructured":"Nguyen Q, Nguyen YTP, Ha TT, Tran DTN, Voong PV, Chau V, et al. Genomic insights unveil the plasmid transfer mechanism and epidemiology of hypervirulent in Vietnam. Nat Commun. 2024;15:4187.","journal-title":"Nat Commun."},{"key":"4235_CR13","doi-asserted-by":"publisher","first-page":"4275","DOI":"10.1128\/AAC.02813-13","volume":"58","author":"S Qin","year":"2014","unstructured":"Qin S, Fu Y, Zhang Q, Qi H, Wen JG, Xu H, et al. High incidence and endemic spread of NDM-1-positive Enterobacteriaceae in Henan province, China. Antimicrob Agents Chemother. 2014;58:4275\u201382.","journal-title":"Antimicrob Agents Chemother"},{"key":"4235_CR14","doi-asserted-by":"publisher","first-page":"e109","DOI":"10.1093\/cid\/cis737","volume":"55","author":"S Borgia","year":"2012","unstructured":"Borgia S, Lastovetska O, Richardson D, Eshaghi A, Xiong J, Chung C, et al. Outbreak of carbapenem-resistant enterobacteriaceae containing blaNDM-1, ontario, Canada. Clin Infect Dis. 2012;55:e109\u201317.","journal-title":"Clin Infect Dis"},{"key":"4235_CR15","doi-asserted-by":"publisher","first-page":"455","DOI":"10.1089\/cmb.2012.0021","volume":"19","author":"A Bankevich","year":"2012","unstructured":"Bankevich A, Nurk S, Antipov D, Gurevich AA, Dvorkin M, Kulikov AS, et al. SPAdes: a new genome assembly algorithm and its applications to single-cell sequencing. J Comput Biol. 2012;19:455\u201377.","journal-title":"J Comput Biol"},{"key":"4235_CR16","doi-asserted-by":"publisher","first-page":"e1005595","DOI":"10.1371\/journal.pcbi.1005595","volume":"13","author":"RR Wick","year":"2017","unstructured":"Wick RR, Judd LM, Gorrie CL, Holt KE. Unicycler: resolving bacterial genome assemblies from short and long sequencing reads. PLoS Comput Biol. 2017;13:e1005595.","journal-title":"PLoS Comput Biol"},{"key":"4235_CR17","doi-asserted-by":"publisher","first-page":"3491","DOI":"10.1093\/jac\/dkaa345","volume":"75","author":"V Bortolaia","year":"2020","unstructured":"Bortolaia V, Kaas RS, Ruppe E, Roberts MC, Schwarz S, Cattoir V, et al. Resfinder 4.0 for predictions of phenotypes from genotypes. J Antimicrob Chemother. 2020;75:3491\u2013500.","journal-title":"J Antimicrob Chemother"},{"key":"4235_CR18","doi-asserted-by":"publisher","first-page":"3895","DOI":"10.1128\/AAC.02412-14","volume":"58","author":"A Carattoli","year":"2014","unstructured":"Carattoli A, Zankari E, Garcia-Fernandez A, Larsen MV, Lund O, Villa L, et al. In Silico detection and typing of plasmids using plasmidfinder and plasmid multilocus sequence typing. Antimicrob Agents Ch. 2014;58:3895\u2013903.","journal-title":"Antimicrob Agents Ch"},{"key":"4235_CR19","doi-asserted-by":"publisher","first-page":"77","DOI":"10.1186\/s13073-018-0587-5","volume":"10","author":"MMC Lam","year":"2018","unstructured":"Lam MMC, Wyres KL, Judd LM, Wick RR, Jenney A, Brisse S, et al. Tracking key virulence loci encoding aerobactin and salmochelin siderophore synthesis in Klebsiella pneumoniaeGenome Med 2018;10:77","journal-title":"Genome Med"},{"key":"4235_CR20","doi-asserted-by":"publisher","first-page":"3691","DOI":"10.1093\/bioinformatics\/btv421","volume":"31","author":"AJ Page","year":"2015","unstructured":"Page AJ, Cummins CA, Hunt M, Wong VK, Reuter S, Holden MT, et al. Roary: rapid large-scale prokaryote pan genome analysis. Bioinformatics. 2015;31:3691\u20133.","journal-title":"Bioinformatics"},{"key":"4235_CR21","doi-asserted-by":"publisher","first-page":"1641","DOI":"10.1093\/molbev\/msp077","volume":"26","author":"MN Price","year":"2009","unstructured":"Price MN, Dehal PS, Arkin AP. FastTree: computing large minimum evolution trees with profiles instead of a distance matrix. Mol Biol Evol. 2009;26:1641\u201350.","journal-title":"Mol Biol Evol"},{"key":"4235_CR22","doi-asserted-by":"publisher","first-page":"402","DOI":"10.1186\/1471-2164-12-402","volume":"12","author":"NF Alikhan","year":"2011","unstructured":"Alikhan NF, Petty NK, Ben Zakour NL, Beatson SA. BLAST ring image generator (BRIG): simple prokaryote genome comparisons. BMC Genomics. 2011;12:402.","journal-title":"BMC Genomics"},{"key":"4235_CR23","doi-asserted-by":"publisher","first-page":"1009","DOI":"10.1093\/bioinformatics\/btr039","volume":"27","author":"MJ Sullivan","year":"2011","unstructured":"Sullivan MJ, Petty NK, Beatson SA. Easyfig: a genome comparison visualizer. Bioinformatics. 2011;27:1009\u201310.","journal-title":"Bioinformatics"},{"key":"4235_CR24","doi-asserted-by":"publisher","first-page":"101083","DOI":"10.1016\/j.drup.2024.101083","volume":"74","author":"Q Wang","year":"2024","unstructured":"Wang Q, Wang R, Wang S, Zhang A, Duan Q, Sun S, et al. Expansion and transmission dynamics of high risk carbapenem-resistant Klebsiella pneumoniae subclones in china: an epidemiological, spatial, genomic analysis. Drug Resist Updat. 2024;74:101083.","journal-title":"Drug Resist Updat"},{"key":"4235_CR25","doi-asserted-by":"publisher","unstructured":"Russo TA, MacDonald U, Hassan S, Camanzo E, LeBreton F, Corey B, et al. An assessment of siderophore production, mucoviscosity, and mouse infection models for defining the virulence spectrum of hypervirulent Klebsiella pneumoniae. mSphere. 2021. https:\/\/doi.org\/10.1128\/mSphere.00045-21.","DOI":"10.1128\/mSphere.00045-21"},{"key":"4235_CR26","doi-asserted-by":"publisher","first-page":"2696","DOI":"10.1093\/jac\/dkaa226","volume":"75","author":"SS Qin","year":"2020","unstructured":"Qin SS, Zhang CL, Schwarz S, Li LY, Dong HY, Yao H, et al. Identification of a novel conjugative mcr-8.2-bearing plasmid in an almost pan-resistant hypermucoviscous Klebsiella pneumoniae ST11 isolate with enhanced virulence. J Antimicrob Chemoth. 2020;75:2696\u20139.","journal-title":"J Antimicrob Chemoth"},{"key":"4235_CR27","doi-asserted-by":"publisher","first-page":"210","DOI":"10.1038\/s41579-024-01105-2","volume":"23","author":"CL Holmes","year":"2025","unstructured":"Holmes CL, Albin OR, Mobley HLT, Bachman MA. Bloodstream infections: mechanisms of pathogenesis and opportunities for intervention. Nat Rev Microbiol. 2025;23:210\u201324.","journal-title":"Nat Rev Microbiol"},{"key":"4235_CR28","doi-asserted-by":"publisher","DOI":"10.1128\/JCM.01206-19","author":"S Harada","year":"2019","unstructured":"Harada S, Aoki K, Yamamoto S, Ishii Y, Sekiya N, Kurai H, et al. Clinical and molecular characteristics of Klebsiella pneumoniae isolates causing bloodstream infections in Japan: occurrence of hypervirulent infections in health care. J Clin Microbiol. 2019. https:\/\/doi.org\/10.1128\/JCM.01206-19.","journal-title":"J Clin Microbiol"},{"key":"4235_CR29","doi-asserted-by":"publisher","DOI":"10.1016\/j.watres.2023.120263","volume":"242","author":"H Zou","year":"2023","unstructured":"Zou H, Zhou Z, Berglund B, Zheng B, Meng M, Zhao L, et al. Persistent transmission of carbapenem-resistant, hypervirulent Klebsiella pneumoniae between a hospital and urban aquatic environments. Water Res. 2023;242: 120263.","journal-title":"Water Res"},{"key":"4235_CR30","doi-asserted-by":"publisher","first-page":"2349768","DOI":"10.1080\/21505594.2024.2349768","volume":"15","author":"T Chen","year":"2024","unstructured":"Chen T, Wang Y, Chi X, Xiong L, Lu P, Wang X, et al. Genetic, virulence, and antimicrobial resistance characteristics associated with distinct morphotypes in ST11 carbapenem-resistant Klebsiella pneumoniae. Virulence. 2024;15: 2349768.","journal-title":"Virulence"},{"issue":"4","key":"4235_CR31","doi-asserted-by":"publisher","first-page":"106747","DOI":"10.1016\/j.ijantimicag.2023.106747","volume":"61","author":"D Pu","year":"2023","unstructured":"Pu D, Zhao J, Lu B, Zhang Y, Wu Y, Li Z, et al. Within-host resistance evolution of a fatal ST11 hypervirulent carbapenem-resistant Klebsiella pneumoniae. Int J Antimicrob Agents. 2023;61(4): 106747.","journal-title":"Int J Antimicrob Agents"},{"key":"4235_CR32","doi-asserted-by":"publisher","first-page":"12758","DOI":"10.1021\/acs.chemrev.6b00003","volume":"116","author":"AB Hickman","year":"2016","unstructured":"Hickman AB, Dyda F. DNA transposition at work. Chem Rev. 2016;116:12758\u201384.","journal-title":"Chem Rev"},{"key":"4235_CR33","doi-asserted-by":"publisher","first-page":"1936","DOI":"10.1080\/22221751.2022.2103454","volume":"11","author":"D Tian","year":"2022","unstructured":"Tian D, Liu X, Chen W, Zhou Y, Hu D, Wang W, et al. Prevalence of hypervirulent and carbapenem-resistant Klebsiella pneumoniae under divergent evolutionary patterns. Emerg Microbes Infect. 2022;11:1936\u201349.","journal-title":"Emerg Microbes Infect"},{"key":"4235_CR34","doi-asserted-by":"publisher","first-page":"2658","DOI":"10.1016\/j.scib.2023.09.040","volume":"68","author":"DN Pu","year":"2023","unstructured":"Pu DN, Zhao JK, Chang K, Zhuo XX, Cao B. Superbugs with hypervirulence and carbapenem resistance in the rise of such emerging nosocomial pathogens in China. Sci Bull. 2023;68:2658\u201370.","journal-title":"Sci Bull"},{"key":"4235_CR35","doi-asserted-by":"publisher","first-page":"107245","DOI":"10.1016\/j.ijantimicag.2024.107245","volume":"64","author":"S Wang","year":"2024","unstructured":"Wang S, Ma S, Sun S, Wang Q, Ding Q, Jin L, et al. Global evolutionary dynamics of virulence genes in ST11-KL47 carbapenem-resistant Klebsiella pneumoniae Int J Antimicrob Agents. 2024;64: 107245.","journal-title":"Int J Antimicrob Agents"},{"key":"4235_CR36","doi-asserted-by":"publisher","first-page":"100918","DOI":"10.1016\/j.drup.2023.100918","volume":"67","author":"Y Zhou","year":"2023","unstructured":"Zhou Y, Wu C, Wang B, Xu Y, Zhao H, Guo Y, et al. Characterization difference of typical KL1, KL2 and ST11-KL64 hypervirulent and carbapenem-resistant Klebsiella pneumoniae. Drug Resist Updat. 2023;67:100918.","journal-title":"Drug Resist Updat"},{"key":"4235_CR37","doi-asserted-by":"publisher","first-page":"e01179","DOI":"10.1128\/mSphere.01179-20","volume":"5","author":"X Yang","year":"2020","unstructured":"Yang X, Ye L, Li Y, Chan EW, Zhang R, Chen S. Identification of a chromosomal integrated DNA fragment containing the rmpA2 and IucABCDiutA virulence genes in Klebsiella pneumoniae. mSphere. 2020;5:e01179.","journal-title":"mSphere."},{"key":"4235_CR38","doi-asserted-by":"publisher","first-page":"2039","DOI":"10.1038\/s41564-019-0566-7","volume":"4","author":"X Yang","year":"2019","unstructured":"Yang X, Wai-Chi Chan E, Zhang R, Chen S. A conjugative plasmid that augments virulence in Klebsiella pneumoniae. Nat Microbiol. 2019;4:2039\u201343.","journal-title":"Nat Microbiol"},{"key":"4235_CR39","doi-asserted-by":"publisher","first-page":"104428","DOI":"10.1016\/j.isci.2022.104428","volume":"25","author":"X Yang","year":"2022","unstructured":"Yang X, Xie M, Xu Q, Ye L, Yang C, Dong N, et al. Transmission of pLVPK-like virulence plasmid in Klebsiella pneumoniae mediated by an Incl1 conjugative helper plasmid. iScience. 2022;25:104428.","journal-title":"iScience"},{"key":"4235_CR40","doi-asserted-by":"publisher","DOI":"10.1128\/mmbr.00119-22","volume":"88","author":"CJ Harmer","year":"2024","unstructured":"Harmer CJ, Hall RM. IS26 and the IS26 family: versatile resistance gene movers and genome reorganizers. Microbiol Mol Biol Rev. 2024;88: e0011922.","journal-title":"Microbiol Mol Biol Rev"},{"key":"4235_CR41","doi-asserted-by":"publisher","first-page":"wrad032","DOI":"10.1093\/ismejo\/wrad032","volume":"18","author":"X Wang","year":"2024","unstructured":"Wang X, Zhang H, Yu S, Li D, Gillings MR, Ren H, et al. Inter-plasmid transfer of antibiotic resistance genes accelerates antibiotic resistance in bacterial pathogens. ISME J. 2024;18:wrad032.","journal-title":"ISME J."}],"container-title":["BMC Microbiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12866-025-04235-6.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s12866-025-04235-6\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12866-025-04235-6.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,9,8]],"date-time":"2025-09-08T16:11:20Z","timestamp":1757347880000},"score":1,"resource":{"primary":{"URL":"https:\/\/bmcmicrobiol.biomedcentral.com\/articles\/10.1186\/s12866-025-04235-6"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,8,11]]},"references-count":41,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["4235"],"URL":"https:\/\/doi.org\/10.1186\/s12866-025-04235-6","relation":{},"ISSN":["1471-2180"],"issn-type":[{"value":"1471-2180","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,8,11]]},"assertion":[{"value":"22 April 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"16 July 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"11 August 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"All mouse experiments and the patient\u2019s clinical record data were approved by the Ethics Committee of Zhengzhou University (ZDYWYJY2019018) and was conducted in compliance with the principles of the Declaration of Helsinki. As this study was retrospective and did not involve clinical interven-tion in patients, the Ethics Committee of Zhengzhou University waived the requirement for informed consent.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"Not Applicable.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare no competing interests.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"493"}}