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Table 1

HR prevalence among apparently susceptible bacteria strains and observed correlation with overexpression level EPs.

Bacteria N Country/isolation sourceAntibiotic/heteroresistant rate (%*)Efflux pumpRef
P. aeruginosa 131China/clinical isolatesIMP/35.1Increased expression of the mexE and mexY compared to their respective native ones.[50]
106China/hospitalized patientsIMP/18.87Higher mexAB expression level compared to IMP-susceptible strains.[15]
4Greece/clinical isolatesIMP and MEM/-Fourfold higher MIC compared to native cells for IMP and MEM[51]
Increased transcription levels of the mexB and mexY genes.
451China/sterile body fluidsCarbapenems/84.9Identical PFGE profiles HR isolates with native cells[48]
HR isolates grew in higher MICs.
All HR isolates are negative for MBL production.
Upregulation in all HR, including mexB, mexC, mexE, and mexX genes.
Reduced expression of the oprD gene in HR isolates
Higher biofilm formation HR isolates
A. baumannii 260South Korea/clinical isolatesTGC/52HR isolates have eightfold-higher TGC MICs than the original isolates.[7]
In HR isolates adeABC efflux pumps are upregulated as a result of ISAba1 insertion into adeS genes.
3Portugal/MDR clinical isolatesColistin/-HR isolates have upregulation of adeB, adeJ, adeG, craA, amvA, abeS, and abeM Eps genes.[14]
576China/clinical isolatesColistin/1.84Colistin HR had weaker biofilm formation capacity than the ATCC19606 strain.[52]
Colistin HR has upregulation of adeI and adeB genes compared to ATCC19606 strain.
K. pneumoniae 74China/carbapenem-resistantTGC/66.2HR isolates have upregulation of acrB in the TGC-resistant subpopulations.[22]
90PMB/100
56Both TGC and PMB/48.4
393China/clinical isolatesEravacycline/5.08Overexpression of oqxAB and macAB efflux pumps and the transcriptional regulator ramA.[34]
Overexpress acrA, acrB, and tolC genes.
Mutations in the acrR and ramR in some HR isolates.
oqxAB and macB mutations in some isolates.
H. influenzae 59Switzerland/clinical isolatesIMP/77.9Mutation in the ftsI gene (encodes PBP3).[53]
Mutation in the OmpP2.
Partial deletion of acrR.
Enterobacter spp.—Laos and Nigeria/-Colistin/-Inactivation of tolC in the acrAB-tolC pump change HR to susceptible strains.[54]
In HR strains, overexpression of acrAB, tolC, and soxSR genes was observed.
In addition, PAβN significantly decreases the MIC of colistin on the HR strain.

Note. N: number, Ref: reference, IMP: imipenem, MEM: meropenem, TGC: tigecycline, and PMB: polymyxin B.