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Mechanisms of Intrinsic Antibiotic Resistance in Enterococci Alexander Kiruthiga1,2, Kesavaram Padmavathy1*

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Mechanisms of Intrinsic Antibiotic Resistance in Enterococci Alexander Kiruthiga1,2, Kesavaram Padmavathy1* Review Article Mechanisms of intrinsic antibiotic resistance in enterococci Alexander Kiruthiga1,2, Kesavaram Padmavathy1* ABSTRACT Enterococci are considered as serious nosocomial pathogens as they are likely to exhibit resistance effectively to all antibiotics meant for clinical use. The most predominant species encountered frequently among human infections includes Enterococcus faecalis and Enterococcus faecium. Antibiotic resistance in enterococci may be either intrinsic or acquired through mutation of the intrinsic genes or horizontal gene transfer of resistance determinants. This paper reviews the mechanisms of intrinsic resistance in enterococci. KEY WORDS: Enterococcus faecalis, Enterococcus faecium, Enterococcus, Intrinsic resistance INTRODUCTION species and is not attributed to horizontal gene transfer.[4] The genes encoding intrinsic resistance Among Enterococci, Enterococcus faecalis and may either be expressed constitutively (always Enterococcus faecium are the most often encountered expressed) or induced (expressed only upon antibiotic species in various human infections ranging from exposure).[5] Due to the limited choice of antibiotics uncomplicated urinary tract infection to serious against enterococci, monotherapy with a single class bacteremia. Enterococci are considered as serious of antimicrobial agents often results in poor treatment nosocomial pathogens due to their intrinsic resistance outcomes and is significantly associated with and their potential to acquire resistance to various intrinsic resistance exhibited by them. Enterococci antimicrobial agents.[1] Besides exhibiting natural are proven to be intrinsically resistant to β-lactams, intrinsic resistance to multiple antimicrobial classes aminoglycosides, and sulfonamides.[6] (beta-lactams, aminoglycosides, and glycopeptides), they possess a remarkable ability to acquire resistance Intrinsic resistance in enterococci is found to be mediated to last resort of antibiotics (quinupristin-dalfopristin, by different mechanisms of resistance (Table 1). (1) Low linezolid, daptomycin, and tigecycline) that are or altered cell wall permeability, (2) class of modifying used to treat drug resistant enterococci.[2] Hence, enzymes such as ribosome modifying methyl transferase the therapeutic management of serious enterococcal and aminoglycoside modifying enzymes, (3) production infections is a cause of concern.[3] Recent reviews of penicillin binding proteins (PBPs), and (4) ATP- related to enterococcal resistance have mainly focused binding cassette (ABC) efflux pumps.[7] on acquired resistance in enterococci. Hence, this review narrates the different mechanisms of intrinsic INTRINSIC RESISTANCE resistance exhibited by E. faecalis, E. faecium, and TOWARDS VARIOUS CLASSES other clinically relevant Enterococcal species. OF ANTIBIOTICS Intrinsic resistance is an inherent property present β-lactams universally in the genome of a particular bacterial Ampicillin/penicillin Access this article online Due to low affinity PBPs Peptidoglycan synthesis and bacterial viability Website: jprsolutions.info ISSN: 0975-7619 have been a major target in antimicrobial therapy 1Department of Microbiology, Research Laboratory for Oral and Systemic Health, Sree Balaji Dental College and Hospital, Bharath Institute of Higher Education and Research, Chennai, Tamil Nadu, India, 2Department of Microbiology, Priyadarshini Dental College and Hospital, Pandur, Tamil Nadu, India *Corresponding author: Kesavaram Padmavathy, Department of Microbiology, Sree Balaji Dental College and Hospital, Bharath Institute of Higher Education and Research, Velachery Main Road, Chennai - 600 100, Tamil Nadu, India. Phone: 91-9884164212. E-mail: [email protected] Received on: 09-01-2020; Revised on: 19-02-2020; Accepted on: 15-03-2020 896 Drug Invention Today | Vol 13 • Issue 6 • 2020 Alexander Kiruthiga and Kesavaram Padmavathy Drug Invention Today | Today Drug Invention Table 1: Intrinsic resistance mechanisms in enterococci S. No. Name of the antibiotic Mode of action Mechanism of intrinsic Confers intrinsic Species involved References resistance resistance to Intrinsic resistance mechanisms of clinically effective antibiotics against enterococci 1. β – lactams Interferes with the (1) Production of β-lactamases, β-lactam antibiotics E. faecalis [8] Vol 13 •Issue62020 Vol synthesis of peptidoglycan which destroy the β-lactam ring E. faecium Ampicillin by inhibiting penicillin- (2) Presence of altered PBPs, which Low-level resistance to binding proteins have lower affinity for β-lactams penicillin Penicillin PBP4 E. faecalis [1] PBP5 E. faecium [8] (3) Altered cell wall β- lactam agents E. faecium [8,9] Cephalosporins Inhibition of cell wall (1) PBP-5 cross-linkage of β- lactam agents E. faecium [10,11] synthesis peptidoglycan (2) Regulation of signal β-lactam agents E. faecalis [12] transduction pathway (3) Kinase activity of IreK Cephalosporins E. faecalis [13] (4) Presence of mur AA gene Enhances intrinsic E. faecalis [3] resistance to cephalosporins 2. Aminoglycosides Binds to the 16S rRNA of (1) Low cell wall permeability Low-level aminoglycoside E. faecalis [14,16] the 30S ribosomal subunit resistance and interferes with protein (2) Aminoglycoside modifying Amikacin, kanamycin, E. faecium [18] synthesis enzymes netilmicin, and tobramycin (3) Ribosome modifying enzymes Tobramycin and kanamycin E. faecium [19] 3. Glycopeptides Inhibits bacterial growth Presence of 9 Van phenotypes reported till date [22] by interfering with the vanC; vanC1, C2, C3, C4 codes Low-level resistance to E. gallinarum [29-32] synthesis of peptidoglycan for intrinsic resistance vancomycin but not to E. casseliflavus teicoplanin E. flavescens vanD; vanD1, D2, D3, D4, D5 Low to high for both E. faecium [23] vancomycin and teicoplanin vanN Low-level resistance to E. faecium [28] vancomycin 4. Fluoroquinolones Inhibition of bacterial Presence of qnr gene homolog Ofloxacin and ciprofloxacin E. faecalis [34] topoisomerase enzymes, (qnr E.faecalis) namely DNA gyrase and topoisomerase IV (Contd...) 897 Alexander Kiruthiga and Kesavaram Padmavathy 898 Table 1: (Continued) S. No. Name of the antibiotic Mode of action Mechanism of intrinsic Confers intrinsic Species involved References resistance resistance to Intrinsic resistance mechanisms of clinically effective antibiotics against enterococci 5. Macrolides/lincosamides/ Inhibition of protein ABC efflux pump Quinupristin E. faecalis [43] streptogramins synthesis by binding to (streptogramin B) the 50S subunit of the dalfopristin (streptogramin E. faecalis [7] ribosome A class) Clindamycin (1) msrC gene Clindamycin, streptogramin E. faecium [8, 44] A and B (2) lsa gene Intrinsic resistance E. faecalis [8, 45, 46] to lincosamides and E. avium streptogramins E. gallinarum E. casseliflavus Intrinsic resistance mechanisms of least clinically important antibiotics against enterococci 6. Trimethoprim- Inhibition of Absorption of folic acid without Intrinsic resistance to TMP- Enterococcal species [7] sulfamethoxazole tetrahydrofolate synthesis the de novo folate synthesis SMX in the presence of and dihydropteroate folic acid Drug Invention Today | Today Drug Invention synthetase 7. Fusidic acid Inhibition of protein Reduced uptake and enzymatic Moderate resistance E. faecalis [46,50] synthesis by hindering inactivation E. faecium both peptide translocation and ribosome disassembly E. faecalis: Enterococcus faecalis, E. faecium Enterococcus faecium, E. avium: Enterococcus avium, E. casseliflavus: Enterococcus casseliflavus, E. gallinarum: Enterococcus gallinarum, TMP-SMX: Trimethoprim/sulfamethoxazole Vol 13 • Issue62020 Vol Alexander Kiruthiga and Kesavaram Padmavathy for decades. Cell wall active β-lactam antibiotics the presence of mur AA and mur AB (homolog such as ampicillin and penicillin inhibit of mur A in E. coli) of which mur AA enhances its peptidoglycan synthesis. PBPs considered essential intrinsic resistance to cephalosporin.[3] for the peptidoglycan synthesis can be divided as Class A (bifunctional enzymes responsible for D, Aminoglycosides D-transpeptidase, and transglycosylase activity) Enterococci exhibit a low-level intrinsic resistance and Class B (contains only transpeptidase domain to aminoglycosides. Nevertheless, the degree of and depends on the transglycosylase activity of resistance varies among various aminoglycosides with other enzymes). Six putative PBP genes have been minimum inhibitory concentrations (MICs) ranging identified among E. faecalis and E. faecium, of which from as low as 4 µg/mL to as high as 256 µg/mL.[14,15] three genes (ponA, pbpF, pbpZ) belong to Class A Decreased uptake of aminoglycosides leads to low- and other three genes (pbp5, pbpA, pbpB) belong level aminoglycoside resistance among Enterococcal to Class B.[8] Intrinsic resistance of enterococci to species. Enterococci being a facultative anaerobe, β-lactams is associated with the expression of species- its metabolic pathway interferes with the proteins specific low-affinity PBPs generally, PBP5 (in E. involved in electron transport and hence limits the faecium) and PBP4 (in E. faecalis) that bind weakly to uptake of aminoglycosides and thereby contributes beta-lactam antibiotics.[1,8] to intrinsic low level aminoglycoside resistance.[14,16] Nevertheless, a combination of aminoglycosides with β-lactamase-mediated resistance a cell wall active agent significantly enhances the Another mechanism of ampicillin resistance uptake of aminoglycosides in enterococci.[17] E. faecium described
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