{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T18:45:26Z","timestamp":1789497926837,"version":"build-2803163510"},"reference-count":60,"publisher":"MDPI AG","issue":"7","license":[{"start":{"date-parts":[[2022,6,28]],"date-time":"2022-06-28T00:00:00Z","timestamp":1656374400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"the Natural Science Foundation of China","doi-asserted-by":"publisher","award":["81974318"],"award-info":[{"award-number":["81974318"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"the Natural Science Foundation of China","doi-asserted-by":"publisher","award":["2018A030310170"],"award-info":[{"award-number":["2018A030310170"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"the Natural Science Foundation of China","doi-asserted-by":"publisher","award":["2018A030313279"],"award-info":[{"award-number":["2018A030313279"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"the Natural Science Foundation of China","doi-asserted-by":"publisher","award":["2019B020209001"],"award-info":[{"award-number":["2019B020209001"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"the Natural Science Foundation of China","doi-asserted-by":"publisher","award":["2021A0505030007"],"award-info":[{"award-number":["2021A0505030007"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"the Natural Science Foundation of China","doi-asserted-by":"publisher","award":["B17018"],"award-info":[{"award-number":["B17018"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003453","name":"Natural Science Foundation of Guangdong Province","doi-asserted-by":"publisher","award":["81974318"],"award-info":[{"award-number":["81974318"]}],"id":[{"id":"10.13039\/501100003453","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003453","name":"Natural Science Foundation of Guangdong Province","doi-asserted-by":"publisher","award":["2018A030310170"],"award-info":[{"award-number":["2018A030310170"]}],"id":[{"id":"10.13039\/501100003453","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003453","name":"Natural Science Foundation of Guangdong Province","doi-asserted-by":"publisher","award":["2018A030313279"],"award-info":[{"award-number":["2018A030313279"]}],"id":[{"id":"10.13039\/501100003453","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003453","name":"Natural Science Foundation of Guangdong Province","doi-asserted-by":"publisher","award":["2019B020209001"],"award-info":[{"award-number":["2019B020209001"]}],"id":[{"id":"10.13039\/501100003453","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003453","name":"Natural Science Foundation of Guangdong Province","doi-asserted-by":"publisher","award":["2021A0505030007"],"award-info":[{"award-number":["2021A0505030007"]}],"id":[{"id":"10.13039\/501100003453","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003453","name":"Natural Science Foundation of Guangdong Province","doi-asserted-by":"publisher","award":["B17018"],"award-info":[{"award-number":["B17018"]}],"id":[{"id":"10.13039\/501100003453","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Guangdong Province Science and Technology Innovation Strategy Special Fund","award":["81974318"],"award-info":[{"award-number":["81974318"]}]},{"name":"Guangdong Province Science and Technology Innovation Strategy Special Fund","award":["2018A030310170"],"award-info":[{"award-number":["2018A030310170"]}]},{"name":"Guangdong Province Science and Technology Innovation Strategy Special Fund","award":["2018A030313279"],"award-info":[{"award-number":["2018A030313279"]}]},{"name":"Guangdong Province Science and Technology Innovation Strategy Special Fund","award":["2019B020209001"],"award-info":[{"award-number":["2019B020209001"]}]},{"name":"Guangdong Province Science and Technology Innovation Strategy Special Fund","award":["2021A0505030007"],"award-info":[{"award-number":["2021A0505030007"]}]},{"name":"Guangdong Province Science and Technology Innovation Strategy Special Fund","award":["B17018"],"award-info":[{"award-number":["B17018"]}]},{"name":"Guangdong International S&amp;T Cooperation Programme","award":["81974318"],"award-info":[{"award-number":["81974318"]}]},{"name":"Guangdong International S&amp;T Cooperation Programme","award":["2018A030310170"],"award-info":[{"award-number":["2018A030310170"]}]},{"name":"Guangdong International S&amp;T Cooperation Programme","award":["2018A030313279"],"award-info":[{"award-number":["2018A030313279"]}]},{"name":"Guangdong International S&amp;T Cooperation Programme","award":["2019B020209001"],"award-info":[{"award-number":["2019B020209001"]}]},{"name":"Guangdong International S&amp;T Cooperation Programme","award":["2021A0505030007"],"award-info":[{"award-number":["2021A0505030007"]}]},{"name":"Guangdong International S&amp;T Cooperation Programme","award":["B17018"],"award-info":[{"award-number":["B17018"]}]},{"name":"the 111 Project","award":["81974318"],"award-info":[{"award-number":["81974318"]}]},{"name":"the 111 Project","award":["2018A030310170"],"award-info":[{"award-number":["2018A030310170"]}]},{"name":"the 111 Project","award":["2018A030313279"],"award-info":[{"award-number":["2018A030313279"]}]},{"name":"the 111 Project","award":["2019B020209001"],"award-info":[{"award-number":["2019B020209001"]}]},{"name":"the 111 Project","award":["2021A0505030007"],"award-info":[{"award-number":["2021A0505030007"]}]},{"name":"the 111 Project","award":["B17018"],"award-info":[{"award-number":["B17018"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Antibiotics"],"abstract":"<jats:p>The spread of multidrug-resistant enterobacteria strains has posed a significant concern in public health, especially when the strain harbors metallo-beta-lactamase (MBL)-encoding and mobilized colistin resistance (mcr) genes as such genetic components potentially mediate multidrug resistance. Here we report an IncHI2\/2A plasmid carrying blaIMP-26 and mcr-9 in multidrug-resistant Serratia marcescens human isolates YL4. Antimicrobial susceptibility testing was performed by the broth microdilution method. According to the results, S. marcescens YL4 was resistant to several antimicrobials, including \u03b2-lactams, fluorquinolones, sulfanilamide, glycylcycline, and aminoglycosides, except for amikacin. To investigate the plasmid further, we conducted whole-genome sequencing and sequence analysis. As shown, S. marcescens YL4 possessed a circular chromosome with 5,171,477 bp length and two plasmids, pYL4.1 (321,744 bp) and pYL4.2 (46,771 bp). Importantly, sharing high similarity with plasmids pZHZJ1 and pIMP-26, pYL4.1 has an IncHI2\/2A backbone holding a variable region containing blaIMP-26, mcr-9, and two copies of blaTEM-1B. After comprehensively comparing relevant plasmids, we proposed an evolutionary pathway originating from ancestor pZHZJ1. Then, via an acquisition of the mcr-9 element and a few recombination events, this plasmid eventually evolved into pYL4.1 and pIMP-26 through two different pathways. In addition, the phage-like plasmid pYL4.2 also carried a blaTEM-1B gene. Remarkably, this study first identified a multidrug-resistant S. marcescens strain co-harboring blaIMP-26 and mcr-9 on a megaplasmid pYL4.1 and also included a proposed evolutionary pathway of epidemic megaplasmids carrying blaIMP-26.<\/jats:p>","DOI":"10.3390\/antibiotics11070869","type":"journal-article","created":{"date-parts":[[2022,6,28]],"date-time":"2022-06-28T23:59:22Z","timestamp":1656460762000},"page":"869","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":11,"title":["Occurrence of Serratia marcescens Carrying blaIMP-26 and mcr-9 in Southern China: New Insights in the Evolution of Megaplasmid IMP-26"],"prefix":"10.3390","volume":"11","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3128-2712","authenticated-orcid":false,"given":"Yuxia","family":"Zhong","sequence":"first","affiliation":[{"name":"Microbiome Medicine Center, Department of Laboratory Medicine, Zhujiang Hospital, Southern Medical University, Guangzhou 510282, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0354-9069","authenticated-orcid":false,"given":"Wanting","family":"Liu","sequence":"additional","affiliation":[{"name":"Microbiome Medicine Center, Department of Laboratory Medicine, Zhujiang Hospital, Southern Medical University, Guangzhou 510282, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peibo","family":"Yuan","sequence":"additional","affiliation":[{"name":"Microbiome Medicine Center, Department of Laboratory Medicine, Zhujiang Hospital, Southern Medical University, Guangzhou 510282, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ling","family":"Yang","sequence":"additional","affiliation":[{"name":"Department of Laboratory Medicine, The First Affiliated Hospital of Guangzhou Medical University, Guangzhou 510120, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2865-2441","authenticated-orcid":false,"given":"Zhenbo","family":"Xu","sequence":"additional","affiliation":[{"name":"School of Food Science and Engineering, Guangdong Province Key Laboratory for Green Processing of Natural Products and Product Safety, Engineering Research Center of Starch and Vegetable Protein Processing Ministry of Education, South China University of Technology, Guangzhou 510640, China"},{"name":"Department of Civil and Environmental Engineering, University of Maryland, College Park, MD 20742, USA"},{"name":"Research Institute for Food Nutrition and Human Health, Guangzhou 510640, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dingqiang","family":"Chen","sequence":"additional","affiliation":[{"name":"Microbiome Medicine Center, Department of Laboratory Medicine, Zhujiang Hospital, Southern Medical University, Guangzhou 510282, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2022,6,28]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"903","DOI":"10.1099\/00222615-46-11-903","article-title":"Serratia marcescens","volume":"46","author":"Hejazi","year":"1997","journal-title":"J. Med. Microbiol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"767","DOI":"10.1086\/339046","article-title":"Three consecutive outbreaks of Serratia marcescens in a neonatal intensive care unit","volume":"34","author":"Fleisch","year":"2002","journal-title":"Clin. Infect. Dis."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1016\/j.jhin.2008.06.028","article-title":"An outbreak of Serratia marcescens septicaemia in neonatal intensive care unit in Gaza City, Palestine","volume":"70","year":"2008","journal-title":"J. Hosp. Infect."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"4867134","DOI":"10.1155\/2018\/4867134","article-title":"An Outbreak of Serratia marcescens in a Moroccan Neonatal Intensive Care Unit","volume":"2018","author":"Daoudi","year":"2018","journal-title":"Adv. Med."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Cristina, M.L., Sartini, M., and Spagnolo, A.M. (2019). Serratia marcescens Infections in Neonatal Intensive Care Units (NICUs). Int. J. Environ. Res. Public Health, 16.","DOI":"10.3390\/ijerph16040610"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1016\/j.ajic.2018.08.026","article-title":"Serratia marcescens outbreak in a neonatology unit of a Spanish tertiary hospital: Risk factors and control measures","volume":"47","year":"2019","journal-title":"Am. J. Infect. Control."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1341","DOI":"10.1007\/s42770-021-00477-4","article-title":"Three challenging cases of infections by multidrug-resistant Serratia marcescens in patients admitted to intensive care units","volume":"52","author":"Rossato","year":"2021","journal-title":"Braz. J. Microbiol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1186\/s12941-021-00464-5","article-title":"Bloodstream Infections caused by Klebsiella pneumoniae and Serratia marcescens isolates co-harboring NDM-1 and KPC-2","volume":"20","author":"Bes","year":"2021","journal-title":"Ann. Clin. Microbiol. Antimicrob."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"ofab387","DOI":"10.1093\/ofid\/ofab387","article-title":"Meropenem Versus Piperacillin-Tazobactam for Definitive Treatment of Bloodstream Infections Caused by AmpC beta-Lactamase-Producing Enterobacter spp, Citrobacter freundii, Morganella morganii, Providencia spp, or Serratia marcescens: A Pilot Multicenter Randomized Controlled Trial (MERINO-2)","volume":"8","author":"Stewart","year":"2021","journal-title":"Open Forum. Infect. Dis."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"36","DOI":"10.7196\/SAMJ.2016.v106i4.10206","article-title":"Intracranial complications of Serratia marcescens infection in neonates","volume":"106","author":"Madide","year":"2016","journal-title":"S. Afr. Med. J."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1556\/AMicr.53.2006.1.6","article-title":"Virulence factors and resistance mechanisms of Serratia marcescens. A short review","volume":"53","author":"Rodrigues","year":"2006","journal-title":"Acta Microbiol. Immunol. Hung."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"2295","DOI":"10.1128\/JCM.03280-12","article-title":"Outbreak of a cluster with epidemic behavior due to Serratia marcescens after colistin administration in a hospital setting","volume":"51","author":"Merkier","year":"2013","journal-title":"J. Clin. Microbiol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"743312","DOI":"10.3389\/fmicb.2021.743312","article-title":"First Report of bla IMP-4 and bla SRT-2 Coproducing Serratia marcescens Clinical Isolate in China","volume":"12","author":"Huang","year":"2021","journal-title":"Front. Microbiol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"956","DOI":"10.3389\/fmicb.2020.00956","article-title":"Characterization of KPC-Producing Serratia marcescens in an Intensive Care Unit of a Brazilian Tertiary Hospital","volume":"11","author":"Ferreira","year":"2020","journal-title":"Front. Microbiol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"470","DOI":"10.1089\/mdr.2014.0240","article-title":"Outbreak of Serratia marcescens Coproducing ArmA and CTX-M-15 Mediated High Levels of Resistance to Aminoglycoside and Extended-Spectrum Beta-Lactamases, Algeria","volume":"21","author":"Batah","year":"2015","journal-title":"Microb. Drug Resist."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"440","DOI":"10.1128\/CMR.00001-07","article-title":"Carbapenemases: The versatile beta-lactamases","volume":"20","author":"Queenan","year":"2007","journal-title":"Clin. Microbiol. Rev."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1186\/s40560-020-0429-6","article-title":"Molecular diversity of extended-spectrum beta-lactamases and carbapenemases, and antimicrobial resistance","volume":"8","author":"Sawa","year":"2020","journal-title":"J. Intensive Care"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"2563","DOI":"10.1128\/JCM.01905-09","article-title":"Multilocus sequence types of carbapenem-resistant Pseudomonas aeruginosa in Singapore carrying metallo-beta-lactamase genes, including the novel bla(IMP-26) gene","volume":"48","author":"Koh","year":"2010","journal-title":"J. Clin. Microbiol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"6853","DOI":"10.1128\/AAC.01177-16","article-title":"Multidrug-Resistant Sequence Type 235 Pseudomonas aeruginosa Clinical Isolates Producing IMP-26 with Increased Carbapenem-Hydrolyzing Activities in Vietnam","volume":"60","author":"Tada","year":"2016","journal-title":"Antimicrob. Agents Chemother."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"270","DOI":"10.3343\/alm.2012.32.4.270","article-title":"Characterization of carbapenemase genes in Enterobacteriaceae species exhibiting decreased susceptibility to carbapenems in a university hospital in Chongqing, China","volume":"32","author":"Xia","year":"2012","journal-title":"Ann. Lab. Med."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1505","DOI":"10.3201\/eid2408.180151","article-title":"Novel Enterobacter Lineage as Leading Cause of Nosocomial Outbreak Involving Carbapenemase-Producing Strains","volume":"24","author":"Beyrouthy","year":"2018","journal-title":"Emerg. Infect. Dis."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"2820","DOI":"10.1093\/jac\/dkt269","article-title":"Dissemination of metallo-\u03b2-lactamase-producing Pseudomonas aeruginosa of sequence type 235 in Asian countries","volume":"68","author":"Kim","year":"2013","journal-title":"J. Antimicrob. Chemother."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1128\/AAC.01686-12","article-title":"Surveillance and molecular epidemiology of Klebsiella pneumoniae isolates that produce carbapenemases: First report of OXA-48-like enzymes in North America","volume":"57","author":"Lascols","year":"2013","journal-title":"Antimicrob. Agents Chemother."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"3034","DOI":"10.1093\/jac\/dky303","article-title":"Genomic characterization of IMP and VIM carbapenemase-encoding transferable plasmids of Enterobacteriaceae","volume":"73","author":"Matsumura","year":"2018","journal-title":"J. Antimicrob. Chemother."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"10212","DOI":"10.1038\/s41598-019-46777-6","article-title":"A Multidrug Resistance Plasmid pIMP26, Carrying blaIMP-26, fosA5, blaDHA-1, and qnrB4 in Enterobacter cloacae","volume":"9","author":"Wang","year":"2019","journal-title":"Sci. Rep."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"597","DOI":"10.2147\/IDR.S232514","article-title":"Carbapenem-Resistant Enterobacter hormaechei ST1103 with IMP-26 Carbapenemase and ESBL Gene bla (SHV-178)","volume":"13","author":"Gou","year":"2020","journal-title":"Infect. Drug Resist."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1610","DOI":"10.3389\/fmicb.2020.01610","article-title":"Molecular Characterization of an IncFII(k) Plasmid Co-harboring bla (IMP-26) and tet(A) Variant in a Clinical Klebsiella pneumoniae Isolate","volume":"11","author":"Yao","year":"2020","journal-title":"Front. Microbiol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"677720","DOI":"10.3389\/fmed.2021.677720","article-title":"Current Update on Intrinsic and Acquired Colistin Resistance Mechanisms in Bacteria","volume":"8","author":"Gogry","year":"2021","journal-title":"Front. Med."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1016\/S1473-3099(15)00424-7","article-title":"Emergence of plasmid-mediated colistin resistance mechanism MCR-1 in animals and human beings in China: A microbiological and molecular biological study","volume":"16","author":"Liu","year":"2016","journal-title":"Lancet Infect. Dis."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"3087","DOI":"10.1093\/jac\/dkaa205","article-title":"Epidemiology of mobile colistin resistance genes mcr-1 to mcr-9","volume":"75","author":"Ling","year":"2020","journal-title":"J. Antimicrob. Chemother."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"794","DOI":"10.1016\/j.tim.2018.02.006","article-title":"Towards Understanding MCR-like Colistin Resistance","volume":"26","author":"Sun","year":"2018","journal-title":"Trends Microbiol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"508","DOI":"10.1080\/22221751.2020.1732231","article-title":"Identification of novel mobile colistin resistance gene mcr-10","volume":"9","author":"Wang","year":"2020","journal-title":"Emerg. Microbes. Infect."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"e00853-19","DOI":"10.1128\/mBio.00853-19","article-title":"Identification of Novel Mobilized Colistin Resistance Gene mcr-9 in a Multidrug-Resistant, Colistin-Susceptible Salmonella enterica Serotype Typhimurium Isolate","volume":"10","author":"Carroll","year":"2019","journal-title":"mBio"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"8113","DOI":"10.1038\/s41598-020-65106-w","article-title":"Characterization of the global distribution and diversified plasmid reservoirs of the colistin resistance gene mcr-9","volume":"10","author":"Li","year":"2020","journal-title":"Sci. Rep."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"3144","DOI":"10.3389\/fmicb.2018.03144","article-title":"Insight Into the Diversity and Possible Role of Plasmids in the Adaptation of Psychrotolerant and Metalotolerant Arthrobacter spp. to Extreme Antarctic Environments","volume":"9","author":"Romaniuk","year":"2018","journal-title":"Front. Microbiol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"R1158","DOI":"10.1016\/j.cub.2020.07.003","article-title":"Plasmid evolution","volume":"30","author":"Wein","year":"2020","journal-title":"Curr. Biol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"2860","DOI":"10.1093\/molbev\/msw163","article-title":"Survival and Evolution of a Large Multidrug Resistance Plasmid in New Clinical Bacterial Hosts","volume":"33","author":"Porse","year":"2016","journal-title":"Mol. Biol. Evol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"e01801","DOI":"10.1128\/mBio.01801-14","article-title":"Movement of IS26-associated antibiotic resistance genes occurs via a translocatable unit that includes a single IS26 and preferentially inserts adjacent to another IS26","volume":"5","author":"Harmer","year":"2014","journal-title":"mBio"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"409","DOI":"10.1111\/mmi.13774","article-title":"Targeted conservative formation of cointegrates between two DNA molecules containing IS26 occurs via strand exchange at either IS end","volume":"106","author":"Harmer","year":"2017","journal-title":"Mol. Microbiol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1080\/2159256X.2014.998537","article-title":"Autonomous and non-autonomous Tn3-family transposons and their role in the evolution of mobile genetic elements","volume":"4","author":"Szuplewska","year":"2014","journal-title":"Mob. Genet. Elem."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1128\/microbiolspec.MDNA3-0060-2014","article-title":"The Tn3-family of Replicative Transposons","volume":"3","author":"Nicolas","year":"2015","journal-title":"Microbiol. Spectr."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1235","DOI":"10.1128\/AAC.00970-08","article-title":"Mobilization of qnrB2 and ISCR1 in plasmids","volume":"53","author":"Chen","year":"2009","journal-title":"Antimicrob. Agents Chemother."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"676113","DOI":"10.3389\/fmicb.2021.676113","article-title":"First Report of Coexistence of bla (SFO-1) and bla (NDM-1) \u03b2-Lactamase Genes as Well as Colistin Resistance Gene mcr-9 in a Transferrable Plasmid of a Clinical Isolate of Enterobacter hormaechei","volume":"12","author":"Ai","year":"2021","journal-title":"Front. Microbiol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"e0030221","DOI":"10.1128\/mSphere.00302-21","article-title":"First IncHI2 Plasmid Carrying mcr-9.1, bla(VIM-1), and Double Copies of bla(KPC-3) in a Multidrug-Resistant Escherichia coli Human Isolate","volume":"6","author":"Simoni","year":"2021","journal-title":"mSphere"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"756201","DOI":"10.3389\/fmicb.2021.756201","article-title":"Co-occurrence of bla (NDM-1) and mcr-9 in a Conjugative IncHI2\/HI2A Plasmid From a Bloodstream Infection-Causing Carbapenem-Resistant Klebsiella pneumoniae","volume":"12","author":"Liu","year":"2021","journal-title":"Front. Microbiol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"e00573-20","DOI":"10.1128\/AAC.00573-20","article-title":"The mcr-9 Gene of Salmonella and Escherichia coli Is Not Associated with Colistin Resistance in the United States","volume":"64","author":"Tyson","year":"2020","journal-title":"Antimicrob. Agents Chemother."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"e00629-19","DOI":"10.1128\/mSphere.00629-19","article-title":"First Report of bla (VIM-4)- and mcr-9-Coharboring Enterobacter Species Isolated from a Pediatric Patient","volume":"4","author":"Chavda","year":"2019","journal-title":"mSphere"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"e00033-21","DOI":"10.1128\/mSphere.00033-21","article-title":"The ABC-Type Efflux Pump MacAB Is Involved in Protection of Serratia marcescens against Aminoglycoside Antibiotics, Polymyxins, and Oxidative Stress","volume":"6","author":"Shirshikova","year":"2021","journal-title":"mSphere"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"302","DOI":"10.4103\/0255-0857.99491","article-title":"Gene expression analysis of the SdeAB multidrug efflux pump in antibiotic-resistant clinical isolates of Serratia marcescens","volume":"30","author":"Dalvi","year":"2012","journal-title":"Indian J. Med. Microbiol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"828","DOI":"10.3389\/fmicb.2018.00828","article-title":"The Genomic Basis of Intrinsic and Acquired Antibiotic Resistance in the Genus Serratia","volume":"9","author":"Vinuesa","year":"2018","journal-title":"Front. Microbiol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"6075","DOI":"10.1128\/IAI.73.9.6075-6084.2005","article-title":"Characterization of the dapA-nlpB genetic locus involved in regulation of swarming motility, cell envelope architecture, hemolysin production, and cell attachment ability in Serratia marcescens","volume":"73","author":"Soo","year":"2005","journal-title":"Infect. Immun."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"755","DOI":"10.1128\/CMR.00017-11","article-title":"Serratia infections: From military experiments to current practice","volume":"24","author":"Mahlen","year":"2011","journal-title":"Clin. Microbiol. Rev."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"5181","DOI":"10.1128\/AAC.00013-14","article-title":"Serratia marcescens arn, a PhoP-regulated locus necessary for polymyxin B resistance","volume":"58","author":"Lin","year":"2014","journal-title":"Antimicrob. Agents Chemother."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"e0052320","DOI":"10.1128\/JB.00523-20","article-title":"Serratia marcescens RamA Expression Is under PhoP-Dependent Control and Modulates Lipid A-Related Gene Transcription and Antibiotic Resistance Phenotypes","volume":"203","author":"Mariscotti","year":"2021","journal-title":"J. Bacteriol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"4915","DOI":"10.1038\/s41467-020-18668-2","article-title":"Piperacillin\/tazobactam resistance in a clinical isolate of Escherichia coli due to IS26-mediated amplification of blaTEM-1B","volume":"11","author":"Hubbard","year":"2020","journal-title":"Nat. Commun."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"3179","DOI":"10.1093\/jac\/dkz349","article-title":"Resistance to piperacillin\/tazobactam in Escherichia coli resulting from extensive IS26-associated gene amplification of blaTEM-1","volume":"74","author":"Hansen","year":"2019","journal-title":"J. Antimicrob. Chemother."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"608","DOI":"10.1038\/nrmicro1462","article-title":"Integrons: Agents of bacterial evolution","volume":"4","author":"Mazel","year":"2006","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"e24774","DOI":"10.4161\/mge.24774","article-title":"Costly Class-1 integrons and the domestication of the the functional integrase","volume":"3","author":"Harms","year":"2013","journal-title":"Mob. Genet. Elem."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1186\/s41182-020-00202-1","article-title":"Prevalence of class 1 integron in Escherichia coli isolated from animal sources in Iran: A systematic review and meta-analysis","volume":"48","author":"Halaji","year":"2020","journal-title":"Trop. Med. Health"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"413","DOI":"10.1089\/mdr.2021.0172","article-title":"Antimicrobial Resistance and Molecular Characterization of Gene Cassettes from Class 1 Integrons in Escherichia coli Strains","volume":"28","author":"Li","year":"2022","journal-title":"Microb. Drug Resist."}],"container-title":["Antibiotics"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2079-6382\/11\/7\/869\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T23:39:30Z","timestamp":1760139570000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2079-6382\/11\/7\/869"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,6,28]]},"references-count":60,"journal-issue":{"issue":"7","published-online":{"date-parts":[[2022,7]]}},"alternative-id":["antibiotics11070869"],"URL":"https:\/\/doi.org\/10.3390\/antibiotics11070869","relation":{},"ISSN":["2079-6382"],"issn-type":[{"value":"2079-6382","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,6,28]]}}}