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. 2023 Jul 26;14(7):468. doi: 10.1038/s41419-023-05955-1

Table 2.

Overview of the candidate drugs selected for in vitro validation.

Drug Approved as cancer drug Metabolic targets (Recon 2.04) Main metabolic targets Non-metabolic main targets Indication Mode of action Use in cancer research
Fluvastatin No HMGCR, CYP3A4, SLC15A1, CYP2C8, SLCO1B1, CYP2C9, CYP2C19 HMGCR Hypercholesterolemia

HMG-CoA reductase

Inhibitor

Anti-proliferative effects in breast cancer (Garwood et al. [76]), could prevent the onset of renal cancer (Horiguchi et al. [12]), potential synergistic effects with gemcitabine (Bocci et al. [77]) and cisplatin (Taylor-Harding et al. [78])
Ellagic Acid No CYP2E1, CA6, SQLE, CA12, CA3, CA9, CYP1A1, CA1, CA4, CA5B, CA5A, CA7, CA2, CA14 CA6, SQLE, CA12, CA3, CA9, CA1, CA4, CA5B, CA5A, CA7, CA2, CA14 CSNK2A1, GSK3B, PRKACA, PRKCA, PRKCB, SYK Phytochemical abundant in fruits and vegetables Squalene epoxidase (SQLE) inhibitor) Apoptotic and anti-angiogenic effects in cancer cells (Losso et al. [60])
Icatibant No ANPEP BDKRB2

Orphan drug used for hereditary angioedema treatment

(Cicardi et al. [79])

Alanyl Aminopeptidase protein (ANPEP)

Inhibitor

Terbinafine No CYP19A1, SQLE, CYP11A1, CYP2D6 SQLE Antifungal agent

Possibly by targeting

SQLE

Demonstrated anticancer effects in vitro (Chien et al. [80]; Lee et al. [81])
Tioconazole No CYP2E1, CYP19A1, CYP51A1, CYP3A4, CYP2C8, CYP2C19 Antifungal agent Inhibits the ergosterol synthesis Desensitises cancer cells to chemotherapy (P.-F. Liu et al. [65]))
Lovastatin No HMGCR, CYP3A4, SLCO1A2, CYP2D6, CYP2C8, SLCO1B1, CYP2C9 HMGCR HDAC2, ITGAL, NR1I2 Hypercholesterolemia Mevalonate pathway and cholesterol synthesis inhibitor Anti-proliferative properties in cancers (Agarwal et al. [82]; Martirosyan et al. [83]),
Gemcitabine Yes TYMS, CMPK1, RRM1 TYMS, CMPK1, RRM1 Pyrimidine analogue Inhibits DNA replication (Noble and Goa, [84]) and has been approved for the treatment of several cancers
Cladribine Yes RRM1, RRM2, RRM2B RRM1, RRM2, RRM2B POLA1, POLE, POLE2, POLE3, POLE4 Purine analogue Ribonuclease reductase inhibitor Used in the treatment of hairy cell leukaemia (Bryson and Sorkin, [85])
Butenafine No SQLE SQLE Antifungal SQLE inhibitor Reduces cancer proliferation (Cirmena et al. [86])
Cerulenin No FASN FASN Antifungal FASN inhibitor Induces apoptosis in human breast cancer (Liu et al. [87]; Thupari et al. [88]) and in A375 melanoma cell line (Ho et al. [89]). Suppression of colon cancer metastasis in mice liver (Murata et al. [90])
Atovaquone No DHODH, CYP2C9 Ubiquinone analogue used for malaria DHODH inhibitor Inhibits oxidative phosphorylation in cancer (Fiorillo et al. [61]), (Ashton et al. [91])
Tamoxifen Yes CYP19A1 CYP1B1, EBP, CYP3A4, CYP2B6, CYP2D6, CYP2C8, ABCB11, CYP2C9 EBP ESR1, ESR2, GPER1, PRKCA, PRKCB, PRKCD, PRKCE, PRKCG, PRKCI, PRKCQ, PRKCZ Anti-oestrogen Developed to treat breast cancer (Buckley and Goa [92])

12 candidate drugs for repurposing in melanoma were selected for experimental validation. Half of the drugs are already FDA-approved as anticancer agents. Metabolic targets represent inhibited targets in the generic metabolic reconstruction Recon 2.04. The main metabolic targets in addition to non-metabolic targets were identified from the manually curated database Drug Repurposing Hub. Indication, mode of action, and use in cancer research were retrieved from Drug Bank and from literature.