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Table 1 Human ubiquitin specific proteases (USPs) and their reported functions in the cancer progression

From: The role of ubiquitin-specific peptidases in cancer progression

Gene symbol Cellular location Substrate Function and remarks in cancer Inhibitor References
USP1 N FANCD2
PCNA 		DNA repair; Oncogene Pimozideb, ML323, GW7647, C527, 6-Amino-pyrimidines, SJB2-043, SJB3-019A, PR619 [92, 110,111,112,113,114]
USP2 C, N Fatty acid synthase, cyclin D1, MDM2 and 4 Fas/p53, NF-κB, c-Myc; Oncogene NSC632839, AM146, RA-9, RA-14, 2-cyano-pyrimidines and -triazinesb, ML364, PR619 [18, 31, 44, 114,115,116,117,118,119,120,121,122]
USP3 N H2A, H2B DDR, Oncogene   [123,124,125]
USP4 C, N TRAF2, TRAF6 TGFβ, NFκB, Wnt, p53; Oncogene Vialinin A, PR619 [81, 114, 126,127,128]
USP5 L, V, Ca   p53, DDR, Oncogene G9, Vialinin A, WP1130, EOAI3402143, AM146, RA-9, RA-14, PR619 [49, 93, 106, 114, 118, 127, 129,130,131,132]
USP6 Golgi, C   NFκB activation; Oncogene or Suppressor   [133]
USP7 N, C, PML body HDM2, p53, H2B, TP53, MDM2 & 4, FOXO4, PTEN Oncogene P5091, Cpd14, P22077, HBX41108, HBX 19818, HBX 28258, NSC632839, WO2013030218, P0050429, W02013030218, PR619 [114, 117, 121, 134,135,136,137,138,139,140,141,142,143,144,145,146]
USP8 C, N NRDP1, RNF128, STAM2 Oncogene HBX90397, HBX41108, AM146, RA-9, RA-14, Ethyloxyimino-9H-indeno[1,2-b] pyrazine-2,3-dicarbonitrile, PR619 [95, 114, 118, 147,148,149,150]
USP9X C, E, L, V β-catenin, epsins, AF-6, SMAD2 TGFβ, Mcl-1, ERG, AGS-3, ITCH, Wnt, Notch; Oncogene or suppressor G9, WP1130, PR619 [106, 107, 114, 130,131,132, 151,152,153,154]
USP9Y C   Spematogenesis   [155]
USP10 C, N TP53, SNX3, CFTR c-Myc, p53; Oncogene or suppressor P22077, HBX19818, Spautin-1, PR619 [32, 56, 114, 156,157,158]
USP11 N, C BRCA2, NFκBIA DDR, NFκB; Oncogene Mitoxantroneb [70, 104, 159,160,161]
USP12   Androgen receptor Oncogene GW7647 [92, 162,163,164]
USP13 L, V, C, Na MCL1, BECN1, USP10   Spautin-1 [157, 165,166,167]
USP14 C, PM   Wnt; Oncogene VLX1570b, IU1, WP1130, b-AP15, AC17, Auranofinb, Tricyclic heterocyclics, Azepan-4-ones, PR619 [106, 114, 132, 168,169,170,171,172,173,174,175]
USP15 C, N RBX1, SMAD1, 2, 3 & 7 NFκB, Wnt; Oncogene PR619 [114, 176,177,178,179]
USP16 N H2A Chromosome condensation; Oncogene PR619 [114, 180,181,182,183]
USP17   SUDS3 Oncogene   [184,185,186]
USP18 C, N TAK1, TAB1, PTEN JAK-STAT, NFκB; Oncogene   [187, 188]
USP19 ER RNF123 ERAD PR619 [114, 189,190,191]
USP20 C, N, DIO2, ADRB2, TRAF6, Tax Thyroid hormone, hypoxia, NFκB; Oncogene PR619 [114, 192, 193]
USP21 C, N H2A, RIPK1, DDX58, GATA3, IL33 NFκB, NEDD8; Oncogene   [72, 194,195,196,197,198]
USP22 N H2A c-Myc; Oncogene PR619 [114, 199,200,201,202]
USP24 C TP53, DDB2, MCL1, Bax, p300, E2F4, securin, βTrCP Cell growth repressor;
Metastasis promoter;
Overexpression in M2 macrophages 		G9, PR619 [23, 57, 75, 106, 114, 130, 131, 203]
USP25 C, N DDX58 ERAD; Oncogene   [204,205,206]
USP26 N (testis) AR Spermatogenesis   [207,208,209]
USP27X   BCL2L11 tumor suppressor   [58, 210]
USP28 N   CLSPN, c-MYC;
Oncogene or suppressor 		PR619 [114, 211, 212]
USP29 Na   p53 pathway; Oncogene   [213, 214]
USP30 M MFN1, MFN2, DRP1, Parkin Hepatocarcinogenesis   [215,216,217]
USP31 N, C   Inhibition of NFκB   [218]
USP32 PM, Golgi   Oncogene   [219]
USP33 C, N, centrosome HIF1-α DIO2, ADRB2, CCP110, ARRB Tumor suppressor   [192, 220,221,222,223]
USP34 C, N, PM, Extracellular AXIN1, AXIN2, Activation of Wnt; Inhibition of EMT and cancer stemness   [102, 224]
USP35 Na ABIN-2, Aurora B Tumor suppressor through inactivating NFκB   [225, 226]
USP36 N c-Myc Oncogene   
USP37 N, c-Myc Increase in DNA damage repair; Oncogene   
USP38 C, N, GAa     
USP39 N   Oncogene   
USP40 C, N, PM     
USP41 Na     
USP42 Na TP53 p53; Oncogene   [53, 227, 228]
USP43 Na H2BK120 Tumor suppressor   
USP44 N CDC20, EZH2 Oncogene   [229]
USP45 C, N     
USP46 L, Va   Oncogene Pimozideb [113]
USP47 C POLB Oncogene P5091, Cpd14, P22077, PR619 [114, 136, 137, 139, 230]
USP48 C, N Gli1 Oncogene PR619 [114]
USP49 N H2B Tumor suppressor   [231]
USP50 Na   G2/M checkpoint   [232]
USP51 Na     
PAN2 C, N     
USP53 Golgi, Na     
USP54 Ma   Oncogene   
USPL1 N, Cajal body     
DUB3   H2AX G2/M checkpoint;
Cancer associated 		  [233, 234]
CYLD C, N, PM TRAF2/6, NEMO, TRPA1, Tak1, Lck, Bcl3, Dvl, DDX58, K63polyUb-RIPK1, K63polyUβ-IKBKG NFκB and JNK-STAT; Familial tumor suppressor   [68, 235,236,237,238,239]
  1. The roles of USPs in the cancer progression. apredicted; C: Cytoplasm; N: Nucleus; L: Lysosome; V: Vacuole; ER: Endoplasmic reticulum; M: Mitochondria; E: Endosome; bClinic trial on going (https://clinicaltrials.gov/ct2/home)
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