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

USP1

N

FANCD2

PCNA

DNA repair; Oncogene

Pimozide^b, 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 -triazines^b, 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, C^a

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

Mitoxantrone^b

[70, 104, 159,160,161]

USP12

Androgen receptor

Oncogene

GW7647

[92, 162,163,164]

USP13

L, V, C, N^a

MCL1, BECN1, USP10

Spautin-1

[157, 165,166,167]

USP14

C, PM

Wnt; Oncogene

VLX1570^b, IU1, WP1130, b-AP15, AC17, Auranofin^b, 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

N^a

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

N^a

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, GA^a

USP39

N

Oncogene

USP40

C, N, PM

USP41

N^a

USP42

N^a

TP53

p53; Oncogene

[53, 227, 228]

USP43

N^a

H2BK120

Tumor suppressor

USP44

N

CDC20, EZH2

Oncogene

[229]

USP45

C, N

USP46

L, V^a

Oncogene

Pimozide^b

[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

N^a

G2/M checkpoint

[232]

USP51

N^a

PAN2

C, N

USP53

Golgi, N^a

USP54

M^a

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]