Article Figures & Data
Tables
- TABLE 1.
Overview of plasmid families associated with AmpC, ESBLs, 16S rRNA methylases, Qnr, and MBLs in Enterobacteriaceae
Enzymea Replicon(s)b Speciesc No. of plasmids Country(ies)e Source(s) Reference(s) or GenBank accession no. CMY-2 A/C Escherichia coli, S. enterica [Agona, Anatum, Bredeney, Heidelberg, Newport, Typhimurium] 155 Canada, France, Honduras, Iraq, Ireland, UK, USA Humans, cattle, pigs, poultry 1, 14, 32, 48, 73, 113 I1 E. coli, S. enterica [4,5,12:I:−, Ajiobo, Heidelberg, Thompson, Typhimurium] 30 Canada, France, Gambia, Italy, UK, USA Humans, cattle, horses, dogs, pigs, poultry 1, 14, 32, 38, 48 FIA-FIB E. coli 1 UK Humans 48 NT E. coli, S. enterica [Heidelberg] 6 Canada, UK Cattle, poultry, pigs 1, 73 CMY-4 A/C S. enterica [Senftenberg] 1 UK Humans 48 CMY-7 I1 E. coli 11 Pakistan, UK Humans 48 CMY-8 (CTX-M-3) HI2 Klebsiella pneumoniae 1 Taiwan Humans 19 CMY-21 I1 E. coli 1 UK Humans 48 CMY-31 ColE S. enterica [Newport] 1 USA Humans 118 CMY-36 ColE K. pneumoniae 1 Greece Humans 118 CTX-M-1 N E. coli, Klebsiella pneumoniae 29 Denmark, France, Spain Humans, pigs 30, 69, 72, 76 I1 E. coli 16 France, Italy Humans, poultry, dogs 38, 42, 69 FII K. pneumoniae 2 Spain Humans 30 L/M E. coli 2 Spain Humans 76 NT E. coli 1 France Poultry 42 CTX-M-2 A/C E. coli 14 Bolivia, France, Peru, UK Humans 48, 69, 81 HI2 E. coli 5 France, Belgium Humans, poultry 37, 69 P E. coli 2 France, Ireland Humans 48, 69 I1 E. coli 1 France Humans 69 FVII E. coli 1 Bolivia, Peru Humans 81 NT E. coli 3 Bolivia, Peru Humans 81 CTX-M-3 (ArmA) L/M E. coli, Enterobacter cloacae, Citrobacter freundii, K. pneumoniae, Klebsiella oxytoca, Proteus mirabilis, Salmonella enterica, Shigella flexneri, Serratia marcescens 73 Bulgaria, Croatia, France, Korea, Poland, Russia Humans 4, 5, 35, 43, 54, 66, 69, 100, 104 N E. coli, S. enterica [Virchow] 5 Australia, Spain, UK Humans 30, 48, 119 A/C E. coli 2 Spain Humans 29, 76 I1 S. enterica [Anatum, Potsdam] 2 Taiwan Humans 67 FII E. coli 3 Australia, Croatia Humans 66, 119 NT E. coli 3 Croatia, France Humans 43, 66, 69 CTX-M-9 HI2 E. coli, E. cloacae, K. pneumoniae, S. enterica [Virchow] 68 France, Honduras, Pakistan, Spain, UK Humans, poultry 30, 36, 37, 48, 77 P E. coli, K. pneumoniae 10 Spain Humans 77 FII, FIB E. coli 12 Australia, France, Spain Humans 30, 36, 69, 77, 119 I1 E. coli, K. pneumoniae 8 Spain Humans 30, 77 Y E. coli 2 Spain Humans 77 B/O E. coli 1 Spain Humans 77 K E. coli 1 Spain Humans 30 NT E. coli 2 France, Spain Humans 69, 77 CTX-M-10 K E. coli 1 Spain Humans 30 NT E. coli 1 Spain Humans 30 CTX-M-14 K E. coli 28 Australia, France, Spain, UK Cattle, humans 30, 48, 64, 69, 74, 119 FII, FIB E. coli, K. pneumoniae 13 Australia, France Humans 69, 118 I1 E. coli 8 Australia, Bolivia, France, Peru, Spain, UK Humans 4, 48, 74, 81, 119 HI2 E. coli 2 Spain Humans 74 B E. coli 2 Australia Humans 119 A/C E. coli 1 France Humans 69 NT E. coli, S. enterica [Stanley] 9 Australia, France, Spain, Thailand, UK Humans, cattle 4, 30, 48, 64, 69, 119 CTX-M-15 [TEM-1, AAC(6′)-IB-CR] FII, FIA, FIB E. coli, K. pneumoniae, S. sonnei, S. enterica [Enteritidis] 152 Australia, Bolivia, Canada, Central African Republic, Croatia, Czech Republic, France, Kuwait, India, Italy, Portugal, Spain, Switzerland, Tunisia, Turkey, Peru, UK Humans 8, 13, 25, 30, 44, 48, 50, 55, 61, 66, 69, 76, 81, 90, 119 I1 E. coli, S. enterica [Anatum, Ohio, Infantis, Typhimurium] 14 Australia, France, UK Humans 48, 69, 119 A/C E. coli 1 France Humans 69 L/M E. coli 1 France Humans 69 N E. coli 1 France Humans 69 NT E. coli, K. pneumoniae 22 Australia, Bolivia, Canada, France, Peru, Turkey Humans 8, 44, 48, 69, 81, 119 CTX-M-17 ColE K. pneumoniae 1 Vietnam Humans 10 CTX-M-24 FII E. coli 2 Australia Humans 119 I1 E. coli 2 Bolivia, Peru Humans 81 CTX-M-27 FII E. coli 1 Australia Humans 119 CTX-M-32 N E. coli 8 Spain Humans 30, 76 CTX-M-40 N E. coli 1 UK Humans 48 CTX-M-42 L/M E. coli 2 Russia Humans 104 CTX-M-53 Q S. enterica [Westhampton] 1 France DQ268764 CTX-M-56 A/C E. coli 2 Bolivia, Peru Humans 81 CTX-M-62 NT K. pneumoniae 1 Australia Humans 119 DHA-1 FII, FIA S. enterica [Senftenberg] 2 UK Humans 48 SHV-2 A/C E. coli 1 France Humans 69 FII, FIB E. coli 2 France Humans 69 NT E. coli 3 France Humans 69 SHV-4 NT E. coli 2 France Humans 69 SHV-5 L/M K. oxytoca, K. pneumoniae, E. coli, E. cloacae 118 Albania, USA Humans 98, 110 FII, FIB E. coli, K. pneumoniae 2 Spain, Poland Humans 30, 117 A/C E. coli 1 France Humans 69 NT E. coli 1 France Humans 69 SHV-12 I1 E. coli, K. pneumoniae 12 Italy, Spain Humans, poultry 14, 30, 38, 69 K E. coli, K. pneumoniae 5 Spain Humans 30 FII, FIB E. coli, K. pneumoniae Enterobacter aerogenes, S. marcescens 9 Australia, France, Italy, Spain Humans 14, 30, 69, 119 A/C E. coli, K. oxytoca 3 Australia, France, Italy Humans 14, 69, 119 HI2 E. coli 1 Spain Humans 30 FIIKd K. pneumoniae 1 USA CP000649 NT E. coli 4 France, Spain Humans 30, 69 TEM-1 FII, FIA, FIB E. coli 31 France Humans 69 I1 E. coli, S. enterica 7 Czech Republic, France Humans 51, 69 HI1 S. enterica 2 Czech Republic Humans 51 K E. coli 1 France Humans 69 ColE K. pneumoniae, S. enterica [Typhimurium] 2 Italy, USA Humans, rabbits 83, 101 NT E. coli 10 France Humans 69 Inhibitor-resistant TEM FII, FIA, FIB E. coli 35 France Humans 69 I1 E. coli 3 France Humans 69 NT E. coli 4 France Humans 69 TEM-3 A/C E. coli 2 France Humans 69 L/M E. coli 1 France Humans 69 NT E. coli 1 France Humans 69 TEM-10 L/M E. coli 1 France Humans 69 TEM-21 A/C E. coli 7 France Humans 69 NT E. coli 1 France Humans 69 TEM-24 A/C E. coli 27 France Humans 69 Y E. coli 1 France Humans 69 NT E. coli 3 France Humans 69 TEM-52 I1 E. coli, S. enterica [Agona, Derby, Infantis, Paratyphi B, Typhimurium] 20 France, Belgium Humans, poultry 22, 69 NT E. coli 3 France Humans 69 VEB-1 (QnrA1) A/C E. coli, E. cloacae, C. freundii Providencia stuartii, P. mirabilis 13 Algeria, Canada, France, Thailand, Turkey Humans 96 GES-5 Q E. cloacae 1 France Humans EU266532 KPC-2 N K. pneumoniae 1 USA Humans FJ223607 ColE K. pneumoniae 1 Colombia Humans EU176012 IMP-4 (QnrB2, QnrB-4, ArmA) L/M E. coli, K. pneumoniae, K. oxytoca, C. freundii, E. cloacae, Citrobacter amalonaticus, S. marcescens, Morganella morganii 22 Australia Humans 33 A/C E. coli, K. pneumoniae, S. marcescens, Citrobacter koseri, E. cloacae, K. oxytoca 15 Australia Humans 33 HI2 Citrobacter youngae 1 Australia Humans 33 NT K. pneumoniae 1 Australia Humans 33 IMP-8 (QnrB2, SHV-12) HI2 E. cloacae 2 Taiwan Humans 20 IMP-13 A/C, P S. enterica [Anatum, Typhimurium] 2 Colombia Chicken, cheese 79 VIM-1 N E. coli, K. pneumoniae 17 Greece Humans 14, 99 HI2 E. cloacae 4 Spain Humans 108 I1 E. coli, K. pneumoniae, 15 Spain Humans 108 W Serratia liquefaciens, K. oxytoca 2 Greece Humans 71 NT K. pneumoniae 1 Greece Humans 99 VIM-4 (CMY-4) A/C E. cloacae, K. pneumoniae 2 Italy Humans 14 ArmA L/M C. amalonaticus, C. freundii, E. coli, K. pneumoniae, M. morganii, S. marcescens 38 France, Korea Humans 35, 53 FII, FIIAsd E. coli, K. pneumoniae, E. cloacae, S. marcescens 28 Korea Humans 4, 53 A/C E. coli, K. pneumoniae, C. freundii 9 Korea Humans 53 HI2 E. cloacae 4 Korea Humans 53 N E. coli 1 Spain Pigs 45 NT E. coli, K. pneumoniae, Enterobacter spp. 79 Belgium, Korea Humans 5, 53 RmtB (CTX-M-14) A/C E. coli, K. pneumoniae, C. freundii 53 Korea Humans 53, 54 FII E. coli, K. pneumoniae 9 Belgium, France, Korea Humans 4, 5, 53 I1 E. coli 1 Korea Humans 53 MphA I1 S. sonnei 1 France Humans 7 QepA (QnrS1, RmtB, LAP-1) FII-FIA-FIB E. aerogenes, E. coli 3 France, Korea Humans 82, 86 QepA-2 FII-FIA E. coli 1 France Humans 16 QnrA1 [SHV-12, CTX-M-3, CTX-M-9, AAC(6′)-IB-CR] L/M E. cloacae, S. marcescens 6 Korea Humans 59 HI2 K. pneumoniae, E. cloacae 2 Australia, France Humans 96 FII E. aerogenes 1 France Humans 96 I1 K. pneumoniae 1 France Humans 96 QnrA3 N K. pneumoniae, K. ascorbata 2 France Humans 60 QnrB1 [AAC(6′)-IB-CR] L/M C. freundii, S. marcescens 2 Korea Humans 59 QnrB2 [CTX-M-3, AAC(6′)-IB-CR] FII, FIA E. coli 1 Portugal Dog 97 L/M E. cloacae 1 Korea Humans 59 N S. enterica [Bredeney] 1 The Netherlands Poultry 39 QnrB4 [ArmA, CTX-M-14, DHA-1, SHV-12, AAC(6′)-IB-CR] FIIAs,d FIA E. coli, K. pneumoniae, E. cloacae 34 Korea Humans 107 L/M C. freundii 1 Korea Humans 59 QnrB6 (ArmA, DHA-1) FIIAsd K. pneumoniae 1 Korea Humans 107 QnrB19 (SHV-12) N S. enterica [Typhimurium] 1 The Netherlands Poultry 39 L/M S. enterica [Typhimurium] 1 Italy Humans FJ790886 QnrS1 [LAP-2, AAC(6′)-IB-CR] ColE S. enterica [Typhimurium, Virginia, Corvallis, Anatum] 29 The Netherlands, Taiwan, UK Humans 39, 49, 56, 116 N S. enterica [Virchow, Kentucky, Saintpaul] 5 The Netherlands, UK Humans 39, 49 L/M E. cloacae, K. pneumoniae 2 Korea Humans 59, 107 R K. pneumoniae, S. enterica [Montevideo] 2 Taiwan, The Netherlands Humans 21, 39 HI2 S. enterica [Stanley] 1 The Netherlands Humans 39 NT S. enterica [Stanley, Virginia, Virchow] 6 Turkey, UK Humans, poultry 2, 49 QnrS2 U Aeromonas punctata, Aeromonas allosaccharophila, Aeromonas media 3 France, Switzerland Environment 15, 89 Q Not identified NDf Germany Environment 6 ↵ a Plasmids carrying more than one resistance gene are listed only one time with the most relevant resistant mechanism encoded, and additional resistance colocalizing on the same plasmid is reported in parentheses.
↵ b NT, plasmids that were not typeable.
↵ c Salmonella enterica serovars are listed in brackets.
↵ d FIIAs and FIIK indicates replicons homologous to those of the Salmonella virulence plasmids (13) and the pKPN4 plasmid of K. pneumoniae (GenBank accession no. CP000649), respectively.
↵ e UK, United Kingdom, USA, United States.
↵ f ND, not determined.
- TABLE 2.
Major plasmid families and associated resistance genes in drug-resistant Enterobacteriaceae isolated worldwide from human and animal sources
Replicon No. of plasmids Resistance genes Species HFEC (%)a AFEC (%)a F 331 aac(6′)-Ib-cr, blaCMY-2, blaCTX-M-1-2-3-9-14-15-24-27, blaDHA-1, blaSHV-2-5-12, blaTEM-1, armA, rmtB, qepA, qepA2, qnrA1, qnrB2, qnrB4, qnrB6, qnrB19, qnrS1 E. aerogenes, E. cloacae, E. coli, K. pneumoniae, S. enterica, S. marcescens, S. sonnei 53.5 67.0 A/C 317 bla CMY-2-4, blaCTX-M-2-3-14-15-56, blaSHV-2-5-12, blaTEM-3-21-24, blaIMP-4-8-13, blaVIM-4, blaVEB-1, armA, rmtB, qnrA1 C. freundii, C. koseri, E. cloacae, E. coli, K. oxytoca, K. pneumoniae, P. mirabilis, P. stuartii, S. enterica, S. marcescens 1.0 0.0 L/M 270 aac(6′)-Ib-cr, blaCTX-M-1-3-15-42, blaTEM-3-10, blaSHV-5, blaIMP-4-8, armA, qnrA1, qnrB1, qnrB2, qnrB4, qnrS1 C. amalonaticus, C. freundii, E. aerogenes, E. cloacae, E. coli, K. oxytoca, K. pneumoniae, M. morganii, P. mirabilis, S. enterica, S. flexneri, S. marcescens 0.0 0.0 I1 146 bla CMY-2-7-21, blaCTX-M-1-2-3-9-14-15-24, blaSHV-12, blaTEM-1-3-52, blaVIM-1, armA, rmtB, mphA, qnrA1 E. coli, K. pneumoniae, S. enterica, S. sonnei 6.9 17.4 HI2 90 bla CTX-M-2-3-9-14, blaSHV-12, blaIMP-4, blaVIM-1, armA, qnrA1, qnrS1 C. youngae, E. cloacae, E. coli, K. pneumoniae, S. enterica 0.0 3.3 N 70 bla KPC-2, blaCTX-M-1-3-15-32-40, blaVIM-1, qnrA3, qnrB2, qnrB19, qnrS1, armA E. coli, K. ascorbata, K. pneumoniae, S. enterica 0.0 10.9 ↵ a Occurrence of replicon types among 101 E. coli strains isolated from the feces of healthy, antibiotic-free humans (HFEC) and among 92 avian fecal E. coli strains (AFEC), detected by PBRT (52).