{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T19:21:17Z","timestamp":1790882477813,"version":"4.1.0"},"reference-count":97,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2025,9,24]],"date-time":"2025-09-24T00:00:00Z","timestamp":1758672000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Microbiol."],"abstract":"<jats:p>Antimicrobial resistance (AMR) is a critical global challenge, yet the role of environmental dissemination of antibiotic-resistant bacteria remains underexplored, particularly in developing regions. This study investigated urban wastewater overflows from public streets as vectors for extended-spectrum-\u03b2-lactamase (ESBL)- and carbapenemase-producing <jats:italic>Enterobacterales<\/jats:italic> and <jats:italic>Aeromonas<\/jats:italic> in the Suqu\u00eda River (C\u00f3rdoba, Argentina). Sixty-two water samples were analyzed for coliform counts, antimicrobial susceptibility, and resistance genes. Horizontal gene transfer was assessed by conjugation. Sixty-five ESBL- and\/or carbapenemase-producing isolates were recovered, including six carbapenemase producers subjected to whole-genome sequencing (WGS). Urban wastewater exhibited coliform levels &amp;gt;10<jats:sup>8<\/jats:sup> MPN\/100\u202fmL, while river counts increased 2\u20135 logs at urban and downstream sites compared to upstream, where no resistant strains were detected. ESBL- and\/or carbapenemase-producers occurred in ~70% of wastewater and river samples, mainly <jats:italic>Escherichia coli<\/jats:italic> harboring <jats:italic>bla<jats:sub>CTX-M<\/jats:sub><\/jats:italic>. Carbapenemase producers carried <jats:italic>bla<jats:sub>KPC-2<\/jats:sub><\/jats:italic> or <jats:italic>bla<jats:sub>NDM-1<\/jats:sub><\/jats:italic> in <jats:italic>Enterobacter<\/jats:italic>, <jats:italic>Klebsiella<\/jats:italic>, <jats:italic>Citrobacter<\/jats:italic>, and <jats:italic>Aeromonas caviae<\/jats:italic>. WGS revealed extensive resistomes, virulence genes, and plasmid replicons, including IncU and IncA\/C2 linked to carbapenemases. Conjugation confirmed plasmid-mediated transfer of \u03b2-lactamase genes, and genetic context analysis identified clinically recognized transposons. Notably, <jats:italic>Enterobacter kobei<\/jats:italic> and <jats:italic>Aeromonas caviae<\/jats:italic> from the river carried <jats:italic>bla<jats:sub>KPC-2<\/jats:sub><\/jats:italic> on plasmidic contigs combining clinical and environmental elements, consistent with genetic exchange within aquatic ecosystems and transfer of clinically significant resistance determinants to species adapted for riverine survival. These findings identify urban wastewater overflows as AMR hotspots that facilitate the dissemination of multidrug-resistant bacteria and mobile resistance elements into urban and peri-urban aquatic environments, underscoring the need for integrated environmental AMR surveillance.<\/jats:p>","DOI":"10.3389\/fmicb.2025.1669531","type":"journal-article","created":{"date-parts":[[2025,9,24]],"date-time":"2025-09-24T05:41:46Z","timestamp":1758692506000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":4,"title":["Urban wastewater overflows as hotspots for dissemination of bacteria producing extended-spectrum \u03b2-lactamases and carbapenemases in the Suqu\u00eda River, Argentina"],"prefix":"10.3389","volume":"16","author":[{"given":"Susana Eugenia","family":"Ruiz","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fabrizzio Nicolas","family":"Morandini","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mar\u00eda Emilia","family":"Panzetta","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Flavio Gabriel","family":"Lipari","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mar\u00eda Gabriela","family":"Irraz\u00e1bal","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ricardo","family":"Toselli","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mart\u00edn","family":"Der Ohannesian","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cristian","family":"Amieva","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mar\u00eda Eugenia","family":"Valdes","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Federico Javier","family":"Giraudo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mar\u00eda del Rosario","family":"Roll\u00e1n","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Valeria","family":"Am\u00e9","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Claudia","family":"Sola","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"H\u00e9ctor Alex","family":"Saka","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2025,9,24]]},"reference":[{"key":"ref1","doi-asserted-by":"publisher","first-page":"64","DOI":"10.1016\/j.scitotenv.2019.02.183","article-title":"Antibiotic resistance in urban runoff","volume":"667","author":"Almakki","year":"2019","journal-title":"Sci. Total Environ."},{"key":"ref2","doi-asserted-by":"publisher","first-page":"e17","DOI":"10.1093\/nar\/gkab1115","article-title":"Deeplasmid: deep learning accurately separates plasmids from bacterial chromosomes","volume":"50","author":"Andreopoulos","year":"2022","journal-title":"Nucleic Acids Res."},{"key":"ref3","doi-asserted-by":"publisher","first-page":"629","DOI":"10.1016\/S0140-6736(21)02724-0","article-title":"Global burden of bacterial antimicrobial resistance in 2019: a systematic analysis","volume":"399","author":"Antimicrobial Resistance","year":"2022","journal-title":"Lancet"},{"key":"ref4","doi-asserted-by":"publisher","first-page":"716","DOI":"10.1128\/am.22.4.716-717.1971","article-title":"Detection of R factors in naturally occurring Aeromonas salmonicida strains","volume":"22","author":"Aoki","year":"1971","journal-title":"Appl. Microbiol."},{"key":"ref5","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1016\/j.jenvman.2019.05.073","article-title":"High clonal diversity of multidrug-resistant and extended spectrum beta-lactamase-producing Escherichia coli in a wastewater treatment plant","volume":"245","author":"Aristizabal-Hoyos","year":"2019","journal-title":"J. Environ. Manag."},{"key":"ref6","doi-asserted-by":"publisher","first-page":"105","DOI":"10.3390\/tropicalmed6020105","article-title":"Extended spectrum beta-lactamase Escherichia coli in river waters collected from two cities in Ghana, 2018-2020","volume":"6","author":"Banu","year":"2021","journal-title":"Trop. Med. Infect. Dis."},{"key":"ref7","doi-asserted-by":"publisher","first-page":"2145","DOI":"10.1093\/jac\/dkx146","article-title":"Global epidemiology of CTX-M beta-lactamases: temporal and geographical shifts in genotype","volume":"72","author":"Bevan","year":"2017","journal-title":"J. Antimicrob. Chemother."},{"key":"ref8","doi-asserted-by":"publisher","first-page":"e0036623","DOI":"10.1128\/msphere.00366-23","article-title":"Nationwide molecular epidemiology of carbapenemase-producing Citrobacter spp. in France in 2019 and 2020","volume":"8","author":"Biez","year":"2023","journal-title":"mSphere"},{"key":"ref9","doi-asserted-by":"publisher","first-page":"2114","DOI":"10.1093\/bioinformatics\/btu170","article-title":"Trimmomatic: a flexible trimmer for Illumina sequence data","volume":"30","author":"Bolger","year":"2014","journal-title":"Bioinformatics"},{"key":"ref10","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1016\/j.jgar.2013.01.005","article-title":"Carbapenemases: partners in crime","volume":"1","author":"Bush","year":"2013","journal-title":"J. Glob. Antimicrob. Resist."},{"key":"ref11","doi-asserted-by":"publisher","first-page":"e01076-18","DOI":"10.1128\/AAC.01076-18","article-title":"Past and present perspectives on beta-lactamases","volume":"62","author":"Bush","year":"2018","journal-title":"Antimicrob. Agents Chemother."},{"key":"ref12","doi-asserted-by":"publisher","first-page":"e00047-19","DOI":"10.1128\/CMR.00047-19","article-title":"Epidemiology of beta-lactamase-producing pathogens","volume":"33","author":"Bush","year":"2020","journal-title":"Clin. Microbiol. Rev."},{"key":"ref13","doi-asserted-by":"publisher","first-page":"421","DOI":"10.1186\/1471-2105-10-421","article-title":"BLAST+: architecture and applications","volume":"10","author":"Camacho","year":"2009","journal-title":"BMC Bioinformatics"},{"key":"ref14","doi-asserted-by":"publisher","first-page":"3895","DOI":"10.1128\/AAC.02412-14","article-title":"In silico detection and typing of plasmids using PlasmidFinder and plasmid multilocus sequence typing","volume":"58","author":"Carattoli","year":"2014","journal-title":"Antimicrob. Agents Chemother."},{"key":"ref15","doi-asserted-by":"publisher","first-page":"2265","DOI":"10.3390\/microorganisms11092265","article-title":"Impact of environmental sub-inhibitory concentrations of antibiotics, heavy metals, and biocides on the emergence of tolerance and effects on the mutant selection window in E. coli","volume":"11","author":"Chukwu","year":"2023","journal-title":"Microorganisms"},{"key":"ref16","volume-title":"Methods for Antimicrobial dilution and disk susceptibility testing of infrequently isolated or fastidious Bacteria","year":"2016"},{"key":"ref17","volume-title":"Clinical and laboratory standards institute (CLSI). Performance standards for Antimicrobial susceptibility testing","year":"2024"},{"key":"ref18","doi-asserted-by":"publisher","first-page":"12853","DOI":"10.3390\/ijerph191912853","article-title":"Impact of anthropogenic activities on the dissemination of ARGs in the environment-A review","volume":"19","author":"Czatzkowska","year":"2022","journal-title":"Int. J. Environ. Res. Public Health"},{"key":"ref19","doi-asserted-by":"publisher","first-page":"1919","DOI":"10.1038\/s41564-019-0492-8","article-title":"Epidemic of carbapenem-resistant Klebsiella pneumoniae in Europe is driven by nosocomial spread","volume":"4","author":"David","year":"2019","journal-title":"Nat. Microbiol."},{"key":"ref20","doi-asserted-by":"publisher","first-page":"457","DOI":"10.1038\/nature10388","article-title":"Antibiotic resistance is ancient","volume":"477","author":"D'Costa","year":"2011","journal-title":"Nature"},{"key":"ref21","doi-asserted-by":"publisher","first-page":"8084","DOI":"10.1038\/s41467-024-51864-y","article-title":"An expanded database and analytical toolkit for identifying bacterial virulence factors and their associations with chronic diseases","volume":"15","author":"Dong","year":"2024","journal-title":"Nat. Commun."},{"key":"ref22","doi-asserted-by":"publisher","first-page":"e00891-19","DOI":"10.1128\/mSphere.00891-19","article-title":"Novel IncR\/IncP6 hybrid plasmid pCRE3-KPC recovered from a clinical KPC-2-producing Citrobacter braakii isolate","volume":"5","author":"Dong","year":"2020","journal-title":"mSphere"},{"key":"ref23","doi-asserted-by":"publisher","first-page":"90","DOI":"10.1111\/j.1469-0691.2007.01846.x","article-title":"Phenotypic detection of extended-spectrum beta-lactamase production in Enterobacteriaceae: review and bench guide","volume":"14","author":"Drieux","year":"2008","journal-title":"Clin. Microbiol. Infect."},{"key":"ref24","doi-asserted-by":"publisher","first-page":"513","DOI":"10.3390\/antibiotics12030513","article-title":"Characterization of third generation cephalosporin- and Carbapenem-resistant Aeromonas isolates from municipal and hospital wastewater","volume":"12","author":"Drk","year":"2023","journal-title":"Antibiotics (Basel)"},{"key":"ref25","doi-asserted-by":"publisher","first-page":"2024","DOI":"10.3389\/fmicb.2017.02024","article-title":"Partitioning of antibiotic Resistance genes and fecal indicators varies intra and inter-storm during combined sewer overflows","volume":"8","author":"Eramo","year":"2017","journal-title":"Front. Microbiol."},{"key":"ref26","doi-asserted-by":"publisher","first-page":"e0140622","DOI":"10.1128\/aac.01406-22","article-title":"Epidemiology of Resistance determinants identified in Meropenem-nonsusceptible Enterobacterales collected as part of a global surveillance study, 2018 to 2019","volume":"67","author":"Estabrook","year":"2023","journal-title":"Antimicrob. Agents Chemother."},{"key":"ref27","article-title":"European committee on Antimicrobial susceptibility testing (EUCAST)","year":"2024"},{"key":"ref28","doi-asserted-by":"publisher","first-page":"479","DOI":"10.3390\/pathogens12030479","article-title":"Emergence of hyper-epidemic clones of Enterobacterales clinical isolates co-producing KPC and Metallo-Beta-lactamases during the COVID-19 pandemic","volume":"12","author":"Faccone","year":"2023","journal-title":"Pathogens"},{"key":"ref29","doi-asserted-by":"publisher","first-page":"12728","DOI":"10.1038\/s41598-021-91456-0","article-title":"AMRFinderPlus and the reference gene catalog facilitate examination of the genomic links among antimicrobial resistance, stress response, and virulence","volume":"11","author":"Feldgarden","year":"2021","journal-title":"Sci. Rep."},{"key":"ref30","doi-asserted-by":"publisher","first-page":"129","DOI":"10.3390\/microorganisms8010129","article-title":"An update on the genus Aeromonas: taxonomy, epidemiology, and pathogenicity","volume":"8","author":"Fernandez-Bravo","year":"2020","journal-title":"Microorganisms"},{"key":"ref31","doi-asserted-by":"publisher","first-page":"1107","DOI":"10.1126\/science.1220761","article-title":"The shared antibiotic resistome of soil bacteria and human pathogens","volume":"337","author":"Forsberg","year":"2012","journal-title":"Science"},{"key":"ref32","doi-asserted-by":"publisher","first-page":"4750","DOI":"10.1128\/JB.00189-09","article-title":"Comparative genomics of the IncA\/C multidrug resistance plasmid family","volume":"191","author":"Fricke","year":"2009","journal-title":"J. Bacteriol."},{"key":"ref33","doi-asserted-by":"publisher","first-page":"722536","DOI":"10.3389\/fcimb.2021.722536","article-title":"Characterization of emerging pathogens carrying Bla(KPC-2) gene in IncP-6 plasmids isolated from urban sewage in Argentina","volume":"11","author":"Ghiglione","year":"2021","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"ref34","doi-asserted-by":"publisher","first-page":"1520","DOI":"10.1111\/j.1469-0691.2011.03600.x","article-title":"Clonal dissemination of Klebsiella pneumoniae ST258 harbouring KPC-2 in Argentina","volume":"17","author":"Gomez","year":"2011","journal-title":"Clin. Microbiol. Infect."},{"key":"ref35","doi-asserted-by":"publisher","first-page":"1174336","DOI":"10.3389\/fmicb.2023.1174336","article-title":"Spatiotemporal dissemination of ESBL-producing Enterobacterales in municipal sewer systems: a prospective, longitudinal study in the city of Basel, Switzerland","volume":"14","author":"Gomez-Sanz","year":"2023","journal-title":"Front. Microbiol."},{"key":"ref36","doi-asserted-by":"publisher","first-page":"100144","DOI":"10.1016\/j.crmicr.2022.100144","article-title":"Chromosomal integration of Bla (CTX-M) genes in diverse Escherichia coli isolates recovered from river water in Japan","volume":"3","author":"Gomi","year":"2022","journal-title":"Curr. Res. Microb. Sci."},{"key":"ref37","doi-asserted-by":"publisher","first-page":"210","DOI":"10.1016\/j.plasmid.2005.11.003","article-title":"Evidence that IncG (IncP-6) and IncU plasmids form a single incompatibility group","volume":"55","author":"Haines","year":"2006","journal-title":"Plasmid"},{"key":"ref38","doi-asserted-by":"publisher","first-page":"186","DOI":"10.3390\/antibiotics9040186","article-title":"The current burden of Carbapenemases: review of significant properties and dissemination among gram-negative Bacteria","volume":"9","author":"Hammoudi Halat","year":"2020","journal-title":"Antibiotics (Basel)"},{"key":"ref39","doi-asserted-by":"publisher","first-page":"2000118","DOI":"10.2807\/1560-7917.ES.2021.26.21.2000118","article-title":"Outbreak of OXA-48-producing Enterobacterales in a haematological ward associated with an uncommon environmental reservoir, France, 2016 to 2019","volume":"26","author":"Jolivet","year":"2021","journal-title":"Euro Surveill."},{"key":"ref40","doi-asserted-by":"publisher","first-page":"1005","DOI":"10.1099\/mic.0.055459-0","article-title":"Ribosomal multilocus sequence typing: universal characterization of bacteria from domain to strain","volume":"158","author":"Jolley","year":"2012","journal-title":"Microbiology (Reading)"},{"key":"ref41","doi-asserted-by":"publisher","first-page":"124","DOI":"10.12688\/wellcomeopenres.14826.1","article-title":"Open-access bacterial population genomics: BIGSdb software, the PubMLST.org website and their applications","volume":"3","author":"Jolley","year":"2018","journal-title":"Wellcome Open Res."},{"key":"ref42","doi-asserted-by":"publisher","first-page":"106848","DOI":"10.1016\/j.ijantimicag.2023.106848","article-title":"One health and Global Health view of Antimicrobial susceptibility through the \"eye\" of Aeromonas: systematic review and Meta-analysis","volume":"62","author":"Jones","year":"2023","journal-title":"Int. J. Antimicrob. Agents"},{"key":"ref43","doi-asserted-by":"publisher","first-page":"1053","DOI":"10.2166\/wh.2024.041","article-title":"Carbapenem-resistant Enterobacterales in wastewater resources and healthy carriers: A survey in Iran","volume":"22","author":"Khavandi","year":"2024","journal-title":"J. Water Health"},{"key":"ref44","doi-asserted-by":"publisher","first-page":"121989","DOI":"10.1016\/j.watres.2024.121989","article-title":"National-scale antimicrobial resistance surveillance in wastewater: A comparative analysis of HT qPCR and metagenomic approaches","volume":"262","author":"Knight","year":"2024","journal-title":"Water Res."},{"key":"ref45","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1111\/nyas.14223","article-title":"Plasmid evolution in carbapenemase-producing Enterobacteriaceae: a review","volume":"1457","author":"Kopotsa","year":"2019","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"ref46","doi-asserted-by":"publisher","first-page":"173554","DOI":"10.1016\/j.scitotenv.2024.173554","article-title":"River waters in Greece: A reservoir for clinically relevant extended-spectrum-beta-lactamases-producing Escherichia coli","volume":"941","author":"Kotzamanidis","year":"2024","journal-title":"Sci. Total Environ."},{"key":"ref47","doi-asserted-by":"publisher","first-page":"132","DOI":"10.1016\/j.envint.2018.04.041","article-title":"Critical knowledge gaps and research needs related to the environmental dimensions of antibiotic resistance","volume":"117","author":"Larsson","year":"2018","journal-title":"Environ. Int."},{"key":"ref48","doi-asserted-by":"publisher","first-page":"257","DOI":"10.1038\/s41579-021-00649-x","article-title":"Antibiotic resistance in the environment","volume":"20","author":"Larsson","year":"2022","journal-title":"Nat. Rev. Microbiol."},{"key":"ref49","volume-title":"El municipio admite 30 desbordes cloacales promedio cada d\u00eda. [Online]","year":"2017"},{"key":"ref50","article-title":"Desborde cloacal en barrio Alberdi: se agrava la crisis ambiental","year":"2021"},{"key":"ref51","doi-asserted-by":"publisher","first-page":"1057","DOI":"10.1016\/S1473-3099(13)70318-9","article-title":"Antibiotic resistance-the need for global solutions","volume":"13","author":"Laxminarayan","year":"2013","journal-title":"Lancet Infect. Dis."},{"key":"ref52","doi-asserted-by":"publisher","first-page":"1993","DOI":"10.1038\/s41396-023-01506-w","article-title":"Differential effects of wastewater treatment plant effluents on the antibiotic resistomes of diverse river habitats","volume":"17","author":"Lee","year":"2023","journal-title":"ISME J."},{"key":"ref53","doi-asserted-by":"publisher","first-page":"6991","DOI":"10.1128\/AEM.00567-11","article-title":"Microarray-based analysis of IncA\/C plasmid-associated genes from multidrug-resistant Salmonella enterica","volume":"77","author":"Lindsey","year":"2011","journal-title":"Appl. Environ. Microbiol."},{"key":"ref54","volume-title":"Standard methods for the examination of water and wastewater","author":"Lipps","year":"2023"},{"key":"ref55","doi-asserted-by":"publisher","first-page":"D912","DOI":"10.1093\/nar\/gkab1107","article-title":"VFDB 2022: a general classification scheme for bacterial virulence factors","volume":"50","author":"Liu","year":"2022","journal-title":"Nucleic Acids Res."},{"key":"ref56","doi-asserted-by":"publisher","first-page":"1543","DOI":"10.1007\/s00253-015-7202-0","article-title":"Antibiotic resistance in urban aquatic environments: can it be controlled?","volume":"100","author":"Manaia","year":"2016","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref57","doi-asserted-by":"publisher","first-page":"e0221960","DOI":"10.1371\/journal.pone.0221960","article-title":"Isolation of five Enterobacteriaceae species harbouring blaNDM-1 and mcr-1 plasmids from a single paediatric patient","volume":"14","author":"Martino","year":"2019","journal-title":"PLoS One"},{"key":"ref58","doi-asserted-by":"publisher","first-page":"i142","DOI":"10.1093\/bioinformatics\/bty266","article-title":"Versatile genome assembly evaluation with QUAST-LG","volume":"34","author":"Mikheenko","year":"2018","journal-title":"Bioinformatics"},{"key":"ref59","doi-asserted-by":"publisher","first-page":"179241","DOI":"10.1016\/j.scitotenv.2025.179241","article-title":"Carbapenemase-producing Enterobacterales and their carbapenemase genes are stably recovered across the wastewater-watershed ecosystem nexus","volume":"975","author":"Mollenkopf","year":"2025","journal-title":"Sci. Total Environ."},{"key":"ref60","doi-asserted-by":"publisher","first-page":"123706","DOI":"10.1016\/j.watres.2025.123706","article-title":"Quantitative study of ESBL and carbapenemase producers in wastewater treatment plants in Seville, Spain: a culture-based detection analysis of raw and treated water","volume":"281","author":"Monge-Olivares","year":"2025","journal-title":"Water Res."},{"key":"ref61","doi-asserted-by":"publisher","first-page":"6538","DOI":"10.1038\/s41598-024-57245-1","article-title":"In-depth analysis of Klebsiella aerogenes resistome, virulome and plasmidome worldwide","volume":"14","author":"Morgado","year":"2024","journal-title":"Sci. Rep."},{"key":"ref62","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1016\/j.eimc.2017.08.007","article-title":"The endless increase of antibiotic resistance in Enterobacteriaceae and the activity of new compounds to face the challenge","volume":"35","author":"Morosini","year":"2017","journal-title":"Enferm. Infecc. Microbiol. Clin."},{"key":"ref63","doi-asserted-by":"publisher","first-page":"152","DOI":"10.1007\/s11274-022-03334-0","article-title":"Occurrence of antibiotics and bacterial resistance genes in wastewater: resistance mechanisms and antimicrobial resistance control approaches","volume":"38","author":"Mutuku","year":"2022","journal-title":"World J. Microbiol. Biotechnol."},{"key":"ref64","volume-title":"Tackling drug-resistant infections globally: Final report and recommendations. Review on Antimicrobial Resistance","author":"O\u2019Neill","year":"2016"},{"key":"ref65","doi-asserted-by":"publisher","first-page":"1043","DOI":"10.1101\/gr.186072.114","article-title":"CheckM: assessing the quality of microbial genomes recovered from isolates, single cells, and metagenomes","volume":"25","author":"Parks","year":"2015","journal-title":"Genome Res."},{"key":"ref66","doi-asserted-by":"publisher","first-page":"561","DOI":"10.1016\/S1473-3099(19)30210-5","article-title":"Carbapenem-resistant Enterobacteriaceae: global action required","volume":"19","author":"Perez","year":"2019","journal-title":"Lancet Infect. Dis."},{"key":"ref67","doi-asserted-by":"publisher","first-page":"e102","DOI":"10.1002\/cpbi.102","article-title":"Using SPAdes de novo assembler","volume":"70","author":"Prjibelski","year":"2020","journal-title":"Curr. Protoc. Bioinform."},{"key":"ref68","doi-asserted-by":"publisher","first-page":"293","DOI":"10.1128\/CVI.00270-07","article-title":"Preparation of bacterial DNA template by boiling and effect of immunoglobulin G as an inhibitor in real-time PCR for serum samples from patients with brucellosis","volume":"15","author":"Queipo-Ortuno","year":"2008","journal-title":"Clin. Vaccine Immunol."},{"key":"ref69","doi-asserted-by":"publisher","first-page":"408","DOI":"10.1186\/s12864-024-10139-y","article-title":"Exploring the resistome, virulome, and mobilome of multidrug-resistant Klebsiella pneumoniae isolates: deciphering the molecular basis of carbapenem resistance","volume":"25","author":"Rahmat Ullah","year":"2024","journal-title":"BMC Genomics"},{"key":"ref70","doi-asserted-by":"publisher","first-page":"231","DOI":"10.1016\/j.bjid.2020.03.002","article-title":"Mobile genetic elements associated with carbapenemase genes in south American Enterobacterales","volume":"24","author":"Reyes","year":"2020","journal-title":"Braz. J. Infect. Dis."},{"key":"ref71","doi-asserted-by":"publisher","first-page":"1677","DOI":"10.1093\/jac\/dkad150","article-title":"Citrobacter freundii resistant to novel beta-lactamase inhibitor combinations and cefiderocol, co-producing class A, B and D carbapenemases encoded by transferable plasmids","volume":"78","author":"Riccobono","year":"2023","journal-title":"J. Antimicrob. Chemother."},{"key":"ref72","doi-asserted-by":"publisher","first-page":"e000206","DOI":"10.1099\/mgen.0.000206","article-title":"MOB-suite: software tools for clustering, reconstruction and typing of plasmids from draft assemblies","volume":"4","author":"Robertson","year":"2018","journal-title":"Microb. Genom."},{"key":"ref73","doi-asserted-by":"publisher","first-page":"22","DOI":"10.1016\/j.nmni.2015.02.007","article-title":"The global threat of antimicrobial resistance: science for intervention","volume":"6","author":"Roca","year":"2015","journal-title":"New Microbes New Infect."},{"key":"ref74","doi-asserted-by":"publisher","first-page":"1121","DOI":"10.1093\/jac\/dkx488","article-title":"Plasmids carrying antimicrobial resistance genes in Enterobacteriaceae","volume":"73","author":"Rozwandowicz","year":"2018","journal-title":"J. Antimicrob. Chemother."},{"key":"ref75","doi-asserted-by":"publisher","first-page":"D20","DOI":"10.1093\/nar\/gkae979","article-title":"Database resources of the National Center for biotechnology information in 2025","volume":"53","author":"Sayers","year":"2025","journal-title":"Nucleic Acids Res."},{"key":"ref76","doi-asserted-by":"publisher","first-page":"000685","DOI":"10.1099\/mgen.0.000685","article-title":"Bakta: rapid and standardized annotation of bacterial genomes via alignment-free sequence identification","volume":"7","author":"Schwengers","year":"2021","journal-title":"Microb. Genom."},{"key":"ref77","doi-asserted-by":"publisher","first-page":"589","DOI":"10.1111\/1758-2229.12772","article-title":"Potential KPC-2 carbapenemase reservoir of environmental Aeromonas hydrophila and Aeromonas caviae isolates from the effluent of an urban wastewater treatment plant in Japan","volume":"11","author":"Sekizuka","year":"2019","journal-title":"Environ. Microbiol. Rep."},{"key":"ref78","article-title":"pyGenomeViz: A genome visualization python package for comparative genomics","author":"Shimoyama","year":"2024"},{"key":"ref79","doi-asserted-by":"publisher","first-page":"D32","DOI":"10.1093\/nar\/gkj014","article-title":"ISfinder: the reference Centre for bacterial insertion sequences","volume":"34","author":"Siguier","year":"2006","journal-title":"Nucleic Acids Res."},{"key":"ref80","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1128\/CMR.00030-10","article-title":"Challenges of antibacterial discovery","volume":"24","author":"Silver","year":"2011","journal-title":"Clin. Microbiol. Rev."},{"key":"ref81","doi-asserted-by":"publisher","first-page":"1728","DOI":"10.3389\/fmicb.2016.01728","article-title":"Review of Antimicrobial Resistance in the environment and its relevance to environmental regulators","volume":"7","author":"Singer","year":"2016","journal-title":"Front. Microbiol."},{"key":"ref82","doi-asserted-by":"publisher","first-page":"114244","DOI":"10.1016\/j.envpol.2020.114244","article-title":"Antibiotic resistant and extended-spectrum beta-lactamase producing faecal coliforms in wastewater treatment plant effluent","volume":"262","author":"Smyth","year":"2020","journal-title":"Environ. Pollut."},{"key":"ref83","doi-asserted-by":"publisher","first-page":"118958","DOI":"10.1016\/j.envpol.2022.118958","article-title":"Occurrence and distribution of Carbapenem-resistant Enterobacterales and carbapenemase genes along a highly polluted hydrographic basin","volume":"300","author":"Teixeira","year":"2022","journal-title":"Environ. Pollut."},{"key":"ref84","doi-asserted-by":"publisher","first-page":"e0310026","DOI":"10.1371\/journal.pone.0310026","article-title":"Carbapenemase-producing bacteria recovered from Nairobi River, Kenya surface water and from nearby anthropogenic and zoonotic sources","volume":"19","author":"Too","year":"2024","journal-title":"PLoS One"},{"key":"ref85","doi-asserted-by":"publisher","first-page":"116133","DOI":"10.1016\/j.envpol.2020.116133","article-title":"Distribution of antibiotics in water, sediments and biofilm in an urban river (Cordoba, Argentina, LA)","volume":"269","author":"Valdes","year":"2021","journal-title":"Environ. Pollut."},{"key":"ref86","doi-asserted-by":"publisher","first-page":"127170","DOI":"10.1016\/j.jhazmat.2021.127170","article-title":"Impacts of municipal wastewater treatment plant discharge on microbial community structure and function of the receiving river in northwest Tibetan plateau","volume":"423","author":"Wang","year":"2022","journal-title":"J. Hazard. Mater."},{"key":"ref87","doi-asserted-by":"publisher","first-page":"13829","DOI":"10.3390\/ijerph192113829","article-title":"Clinically relevant beta-lactam resistance genes in wastewater treatment plants","volume":"19","author":"Wasko","year":"2022","journal-title":"Int. J. Environ. Res. Public Health"},{"key":"ref88","doi-asserted-by":"publisher","first-page":"2817","DOI":"10.3389\/fmicb.2019.02817","article-title":"IS26-mediated transfer of Bla (NDM-1) as the Main route of Resistance transmission during a polyclonal, multispecies outbreak in a German hospital","volume":"10","author":"Weber","year":"2019","journal-title":"Front. Microbiol."},{"key":"ref89","doi-asserted-by":"publisher","first-page":"404","DOI":"10.1007\/s10393-019-01415-5","article-title":"Critical importance of a one health approach to antimicrobial resistance","volume":"16","author":"White","year":"2019","journal-title":"EcoHealth"},{"key":"ref90","volume-title":"WHO bacterial priority pathogens list, 2024: Bacterial pathogens of public health importance to guide research, development and strategies to prevent and control antimicrobial resistance. Licence: CC BY-NC-SA 3.0 IGO","year":"2024"},{"key":"ref91","doi-asserted-by":"publisher","first-page":"e00115-18","DOI":"10.1128\/CMR.00115-18","article-title":"NDM Metallo-beta-lactamases and their bacterial producers in health care settings","volume":"32","author":"Wu","year":"2019","journal-title":"Clin. Microbiol. Rev."},{"key":"ref92","doi-asserted-by":"publisher","first-page":"698478","DOI":"10.3389\/fmicb.2021.698478","article-title":"Complete genome sequences of two novel KPC-2-producing IncU multidrug-resistant plasmids from international high-risk clones of Escherichia coli in China","volume":"12","author":"Wu","year":"2021","journal-title":"Front. Microbiol."},{"key":"ref93","doi-asserted-by":"publisher","first-page":"705","DOI":"10.2166\/wh.2021.112","article-title":"Prevalence of extended-spectrum beta-lactamase (ESBL)-producing Enterobacteriaceae in wastewater: a systematic review and meta-analysis","volume":"19","author":"Zaatout","year":"2021","journal-title":"J. Water Health"},{"key":"ref94","doi-asserted-by":"publisher","first-page":"82","DOI":"10.1016\/j.jhazmat.2018.06.052","article-title":"Contributions of the microbial community and environmental variables to antibiotic resistance genes during co-composting with swine manure and cotton stalks","volume":"358","author":"Zhang","year":"2018","journal-title":"J. Hazard. Mater."},{"key":"ref95","doi-asserted-by":"publisher","first-page":"1183","DOI":"10.3390\/ijerph17041183","article-title":"The prevalence and characterization of extended-Spectrum beta-lactamase- and Carbapenemase-producing Bacteria from hospital sewage, treated effluents and receiving Rivers","volume":"17","author":"Zhang","year":"2020","journal-title":"Int. J. Environ. Res. Public Health"},{"key":"ref96","doi-asserted-by":"publisher","first-page":"975","DOI":"10.1089\/mdr.2018.0431","article-title":"Evaluating dissemination mechanisms of antibiotic-resistant Bacteria in rural environments in China by using CTX-M-producing Escherichia coli as an Indicator","volume":"25","author":"Zou","year":"2019","journal-title":"Microb. Drug Resist."},{"key":"ref97","doi-asserted-by":"publisher","first-page":"3021","DOI":"10.1128\/AEM.00054-13","article-title":"Characteristics of extended-spectrum beta-lactamase- and carbapenemase-producing Enterobacteriaceae isolates from rivers and lakes in Switzerland","volume":"79","author":"Zurfluh","year":"2013","journal-title":"Appl. Environ. Microbiol."}],"container-title":["Frontiers in Microbiology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmicb.2025.1669531\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,9,24]],"date-time":"2025-09-24T05:41:49Z","timestamp":1758692509000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmicb.2025.1669531\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,9,24]]},"references-count":97,"alternative-id":["10.3389\/fmicb.2025.1669531"],"URL":"https:\/\/doi.org\/10.3389\/fmicb.2025.1669531","relation":{},"ISSN":["1664-302X"],"issn-type":[{"value":"1664-302X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,9,24]]},"article-number":"1669531"}}