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. 2017 Apr 25;8:681. doi: 10.3389/fmicb.2017.00681

Table 3.

Anti-mycobacterial drugs and mechanisms of drug resistance.

Agent Mode of action Target Proven utility for (MIC μg/ml) Gene Gene function Most prevalent mutation References
Isoniazid Inhibition of mycolic acid synthesis Mycolic acids M. tuberculosis (0.02–0.1 in 7H10) katG Catalase-peroxidase Ser-315-Thr OFFICIAL, 1990; National Committee for Clinical Laboratory Standards (NCCLS), 2000; Somoskovi et al., 2001; Ramaswamy et al., 2003; Zhang et al., 2005; Guo et al., 2006
inhA Enoyl ACP reductase C-15-T SNP
Ndh NADH dehydrogenase II Arg-13-Cys, Val-18-Ala
ahpC Alkyl hidroperoxidase C-39-T, G-9-A, SNPs
kasA b-ketoacyl-ACP synthase Gly-269-Ser
Rifampicin Inhibition of RNA polymerase RNA polymerase M. tuberculosis (1 in 7H10) M. kansasii (2) M. marinum (1) rpoB B-subunit of RNA polymerase Ser-450-Leu OFFICIAL, 1990; Heep et al., 2001; Somoskovi et al., 2001; Philley and Griffith, 2015
Pyrazinamide Inhibition of energy production and trans-translation Fatty acid synthase-I, ribosomal protein S1 M. tuberculosis (16–50 in LJ) pncA Pyrazinamidase Asp-12-Ala/Asn,Leu-85-Pro Somoskovi et al., 2001; Shi et al., 2011; Feuerriegel et al., 2013; Zhang et al., 2013, 2014
rpsA S1 ribosomal protein Deletion Ala438, Thr-5-Ala
panD Aspartate decarboxylase Ala-128-Ser, Val-138-Aal
Ethambutol Inhibition of arabinogalactan synthesis Arabinosyl transferases M. tuberculosis (5 in 7H10) M. kansasii (5) M. marinum (5) embCAB Arabinosyl transferases Met-306-Val/Ile/Leu OFFICIAL, 1990; Brown-Elliott et al., 2012; Palomino and Martin, 2014; Philley and Griffith, 2015
Streptomycin Inhibition of protein synthesis 30S ribosomal subunit M. tuberculosis (2–10 in 7H10) rpsL rrs gidB S12 ribosomal protein 16S rRNA 16S rRNAmethyltransferase Lis-43-Arg A-1401-G SNP Leu-16-Arg OFFICIAL, 1990; Finken et al., 1993; Da Silva and Palomino, 2011; Verma et al., 2014; Philley and Griffith, 2015
Amikacin/Kanamycin Inhibition of protein synthesis 30S ribosomal subunit M. tuberculosis (5 in 7H10) Rrs 16S rRNA A-1401-G SNP OFFICIAL, 1990; Zaunbrecher et al., 2009; Da Silva and Palomino, 2011; Georghiou et al., 2012; Kasperbauer and De Groote, 2015; Philley and Griffith, 2015
M. kansasii (32) Eis Aminoglycoside acetyltransferase G-37-T, G-10-A, G-14-T SNPs
M. marinum (32)
RGM (64)
Capreomycin/viomycin Inhibition of protein synthesis 30S and 50S ribosome subunits M. tuberculosis (10 in 7H10) rrs 16S rRNA A-1401-G SNP Johansen et al., 2006; Da Silva and Palomino, 2011; Georghiou et al., 2012
tylA rRNA methyltransferase G-223-T SNP
Eis Aminoglycoside acetyltransferase G-37-T, C-12-T SNPs
Ethionamide Inhibition of mycolic acid synthesis Mycolic acids M. tuberculosis (5 in 7H10) ethA Monooxygenase Leu-397-Arg, Leu-328-Met Morlock et al., 2003; Boonaiam et al., 2010; Brossier et al., 2011
inhA Enoyl-ACP reductase Ile-21-Thr/Val, Ser-94-Ala
Ndh NADH dehydrogenase Arg-13-Cys, Val-18-Ala
mshA Glycosyl transferase Val-171-Gly, Aal-187-Val
Fluoroquinolones Inhibition of DNA gyrase DNA gyrase M. tuberculosis (2 in 7H10) gyrA DNA gyrase subunit A Ala-90-Val, Asp-94-Gly/Tyr OFFICIAL, 1990; Cheng et al., 2004; Brown-Elliott et al., 2012; Kasperbauer and De Groote, 2015; Philley and Griffith, 2015
M. kansasii (2) gyrB DNA gyrase subunit B Asn-533-Thr
RGM (4)
P-aminosalicylicacid (PAS) Inhibition of folate synthesis Thymidylate synthase, Dihydrofolate synthase, Dihydrofolate reductases M. tuberculosis (2 in 7H10) thyA Thymidylate synthase Thr-202-Ala, Val-261-Gly Mathys et al., 2009; Da Silva and Palomino, 2011; Zheng et al., 2013; Zhao et al., 2014; Zhang X. et al., 2015
folC Dihydrofolate synthase Glu-153-Aal, Asn-73-Ser
ribD Dihydrofolate reductases G-11-A SNP
Cycloserine Inhibition of peptidoglycan synthesis Alanine racemase, D-Alanine-D-alanine ligase, D-serine/L- and D-alanine/glycine/D-cycloserine proton symporter, L-alanine dehydrogenase M. tuberculosis (5–10 in BACTEC) alr Alanine racemase G-10-T SNP Cáceres et al., 1997; Pelayo et al., 2009; Da Silva and Palomino, 2011; Chen et al., 2012; Gu et al., 2016
ddl D-Alanine-D-alanine ligase
cycA Ald D-serine/L- and D-alanine/glycine/D-cycloserine proton symporter Gly-122-Ser
L-alanine dehydrogenase
Macrolide- ketolide Inhibition of protein synthesis 50S ribosomal subunit M. avium (8–16 in BACTEC) rrl 23S ribosomal RNA A-2058-T, A-2059-C SNPs Meier et al., 1994; Da Silva and Palomino, 2011; Philley and Griffith, 2015
Clofazimine Interfering with redox NADH dehydrogenase M. tuberculosis (1) rv0678 Transcriptional regulator G193 deletion, C-466-T SNPs Hartkoorn et al., 2014; Zhang S. et al., 2015
Cycling, causing membrane destabilization and production of reactive oxygen species rv2535c rv1979c Peptidase Permease G-265-T SNP T-1052-C SNP
Oxazolidinones (Linezolid, Sutezolid [PNU-100480] and AZD5847 Inhibition of protein synthesis 50S ribosomal subunit M. tuberculosis (4–8) rrl rplC 23S ribosomal rRNA 50S ribosomal protein L3 G-2061-T, G-2576-T, G-2270-T T-460-C Hillemann et al., 2008; Williams et al., 2009; Balasubramanian et al., 2014
B-Lactams (In combination with belactamase inhibitor) Inhibition of peptidoglycan synthesis Transpeptidases M. tuberculosis (NR) RGM (128) blaC ponA Pbp beta-lactamase Penicillin-binding Proteins Penicillin-binding proteins – – – Fattorini et al., 1992; Hugonnet and Blanchard, 2007; Brown-Elliott et al., 2012; Wivagg et al., 2014
Tetracyclines and glycylcyclines Inhibition of protein synthesis 30S ribosomal subunit RGM (8–16) 16S rRNA gene 16S rRNA Brown-Elliott et al., 2012; Kasperbauer and De Groote, 2015
Trimethoprim and sulfonamides Inhibition of folate synthesis Dihydrofolate reductase, Dihydropteroate synthetase RGM (64) dfrA sulI and folP1 Dihydrofolate reductase Dihydropteroate synthetase Brown-Elliott et al., 2012; Kasperbauer and De Groote, 2015
Bedaquiline (TMC207) Inhibition of ATP synthesis ATP synthase M. tuberculosis (0.25) atpE ATP synthase Ala-63-Pro, Ile-66-Met Andries et al., 2005; Segala et al., 2012
Nitroimidazoles (PA-824 [Pretomanid] and OPC-67683 [Delamanid]) Inhibition of cell wall lipid and protein synthesis Dehydrogenase, Nitroreductase M. tuberculosis (0.5) Rv0407 Rv3547 Dehydrogenase Nitroreductase – – Stover et al., 2000; Matsumoto et al., 2006; Rivers and Mancera, 2008
SQ109 Inhibition of lipid synthesis Mycolic acids M. tuberculosis (0.5) mmpL3 Membrane transporter Protopopova et al., 2005; Tahlan et al., 2012
Phenothiazines (Thioridazine and Chlorpromazine) Inhibition of calcium transport, inhibition of type II NADH M. tuberculosis (1–32) Ordway et al., 2003; Martins et al., 2008
Benzothiazinones (BTZ043) Inhibition of cell wall arabinans synthesis Nitroreductase, M. tuberculosis (6) nfnB dprE1 Nitroreductase Makarov et al., 2009; Manina et al., 2010
Decaprenylphosphoryl epimerase Decaprenylphosphoryl epimerase
NAS-21 and NAS-91 Inhibition of mycolic acid synthesis FAS-II dehydratase M. tuberculosis (NR) rv0636 FAS-II dehydratase Gratraud et al., 2008

RGM, rapidly growing mycobacteria; SNP, single nucleotide polymorphism; NR, not reported.