Review Article

Mechanisms of Antimicrobial Resistance in ESKAPE Pathogens

Table 1

Categorization of bacterial β-lactamase enzymes by the Bush-Jacoby and Ambler systems.

ClassificationAntimicrobial agents EnzymesDescriptionESKAPE pathogens
Bush-JacobyAmbler

1CNarrow and extended-spectrum cephalosporins, including cephamycinsACT-1, FOX-1, MIR-1, CMYCephalosporinases not inactivated by clavulanic acidEnterobacter spp.
2aAPenicillinsPC1Penicillinases inactivated by clavulanic acid Enterobacteriaceae (e.g., K. pneumoniae, Enterobacter spp.) and nonfermenters (i.e., P. aeruginosa, A. baumannii)
2bAPenicillins, cephalothinTEM-1, TEM-2, TEM-13, SHV-1, SHV-11Broad-spectrum enzymes inactivated by clavulanic acid
2beAPenicillins, oxyimino-cephalosporins (cefotaxime, ceftazidime, ceftriaxone, cefepime), monobactamsTEM-3, TEM-10, TEM-26, SHV-2, SHV-3, Klebsiella oxytoca K1, CTX-M, PER, VEBExtended broad-spectrum enzymes inactivated by clavulanic acid
2brAPenicillins, resistant to clavulanic acid, tazobactam, sulbactamTEM-30, TEM-31, SHV-10, SHV-72Broad-spectrum enzymes with reduced binding to clavulanic acid (inhibitor-resistant TEMs)
2berAPenicillins, oxyimino-cephalosporins, monobactams, resistant to clavulanic acid, tazobactam, sulbactamTEM-50, TEM-158Extended-spectrum enzymes with relative resistance to clavulanic acid
2cAPenicillins, carbenicillinPSE-1, CARB-3Carbenicillin-hydrolyzing enzymes inactivated by clavulanic acid
2ceACarbenicillin, cefepimeRTG-4 (CARB-10)Extended-spectrum carbenicillinase
2dDCloxacillin, oxacillinOXA-1, OXA-2, OXA-10Cloxacillin-hydrolyzing enzymes with variable inactivation by clavulanic acid
2deDCloxacillin, oxacillin, oxyimino-cephalosporins, monobactamsOXA-11, OXA-15
2dfDCloxacillin, oxacillin, carbapenemsOXA-23, OXA-51, OXA-58
2eACephalosporinsCepACephalosporinases inactivated by clavulanic acid
2fAAll -lactams, including carbapenemsKPC, SME, GES, IMI-1Carbapenem-hydrolyzing non-metallo--lactamases
3BAll -lactams, including carbapenems, with exception of monobactamsIMP, VIM, INDMetallo--lactamases
4PenicillinsPenicillinases from Burkholderia cepaciaPenicillinases not inactivated by clavulanic acid

—: no classification in Ambler system.