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Wang J, Dou B, Zheng L, Cao W, Zeng X, Wen Y, Ma J, Li X. Synthesis of Na2S2O4 mediated cleavable affinity tag for labeling of O-GlcNAc modified proteins via azide-alkyne cycloaddition. Bioorganic & medicinal chemistry letters 2021 48 34229054
Abstract:
A facile and convergent procedure for the synthesis of azobenzene-based probe was reported, which could selectively release interested proteins conducted with sodium dithionite. Besides, the cleavage efficiency is closely associated with the structural features, in which an ortho-hydroxyl substituent is necessary for reactivity. In addition, the azobenzene tag applied in the Ac4GlcNAz-labled proteins demonstrated high efficiency and selectivity in comparison with Biotin-PEG4-Alkyne, which provides a useful platform for enrichment of any desired bioorthogonal proteomics.
O-GlcNAc proteins:
PGP, EIFCL, KIF2A, PDLI1, BACH, DFFA, CLIC1, EIF3F, IF2B3, RTCA, PSDE, PPP6, RPC1, PSA7, HNRDL, SC16A, RPAC1, NKRF, EIF3H, PAPS1, SNUT1, ARK72, MYO1B, IDH3B, SAHH2, PLIN3, IMA7, UGDH, CTND1, SNX2, BRD4, WDR1, TBCA, FLNB, PR40A, MPPB, NDUS3, ECI2, CSDE1, U520, WDHD1, EIF3G, PSD10, IDHC, GLRX3, RL1D1, CIAO1, PLPHP, ERLN2, GLSK, SC31A, UBR5, ELP1, VAPB, 6PGL, AGM1, AHSA1, PSMG1, SGPL1, AP2A1, STAU1, TTC4, BPNT1, MBD3, TOM40, ACL6A, GSHR, PNPH, CYTB, KITH, P53, TPM3, PROF1, FUMH, ODPA, CY1, SRP19, DLDH, RU2A, UCHL1, ALDOC, THIO, KAP0, ESTD, ODPB, PYGB, ACADM, G6PD, ADHX, CDK4, HARS1, PEPD, P4HA1, ETFA, MIF, AK1A1, CCNB1, GLNA, DESP, FER, UBF1, PRS6A, RL35A, NELFE, RCC1, E2AK2, SPEE, ANXA7, RAB6A, PSB1, IMDH1, GSTM3, VATB2, FLNA, ACOC, SDHB, PIMT, FBRL, NDKB, ADRO, TCEA1, TBG1, MAOM, IF4B, THTM, RS12, BRD2, DNJB1, PSA1, PSA2, PSA4, STOM, PYR1, PSB4, PSB6, NDUS1, DPOD1, AMPL, ERP29, PRDX3, ECHM, PEBP1, PDIA3, HMOX2, PURA2, PUR8, AL1B1, RPB2, GDIA, TIA1, QCR1, HNRH3, STIP1, PRDX2, P5CR1, DUT, PROF2, SPB6, RADI, T2FA, MYH9, MYH10, FUS, PRS7, MP2K2, HEM6, GNL1, ODO2, SRP14, TALDO, ETFB, VATA, IF4A3, TXLNA, BUD31, CSK, THIM, LIS1, NAMPT, PRS6B, RECQ1, NOP2, CRKL, NSF, CAPZB, COPD, IDHP, AL9A1, RL34, FAS, SYCC, PSB3, IDH3A, SERPH, ANX11, FXR1, FXR2, SMCA4, GALK1, ROA3, HNRPM, IMA5, GDIR1, HNRPF, KIF11, THOP1, CAZA1, BIEA, MAP11, SUCA, SC24C, DRG2, ECHB, DSRAD, HNRH2, IF6, CORO7, ARPC4, CD81, SC61B, MYL6, PSA6, CDC42, SRP54, UB2D3, UBC12, ARP3, RL37A, COPZ1, NTF2, 1433G, PP1A, PP1B, SMD2, PRS10, ERF1, CNBP, H4, RAP1A, RS30, GBB1, GBB2, TRA2B, 2ABA, DYL1, RL38, PP2AA, TBA1B, GSTO1, DCD, RT05, RT09, RL36A, H33, VIGLN, FKBP3, DHSO, EXOSX, ODO1, MMSA, TF65, LGUL, 1433F, CSTF1, SRS11, EF1A2, PTN11, PUR1, GFPT1, C1QBP, BAX, SRSF4, RBBP4, ASPH, GRSF1, AIMP1, ILF3, CSN1, RED, MTAP, TADBP, ROA0, STX5, SRSF9, SRSF5, IFIT5, EIF3I, DC1I2, PICAL, ULA1, SNW1, FHL1, BOP1, UBP2L, DYHC1, EI2BA, TRI25, FLNC, GNA13, CAPR1, KPRA, UBP10, CHD4, NUMA1, GAPD1, EMC2, SEPT2, IF4H, IPYR, CNN3, SC23B, SF01, TRIP6, MARE1, ELAV1, TOM34, VAMP3, ADRM1, PKN2, CSRP2, DPYL2, RBBP7, H2B2E, PCKGM, TRXR1, TIM50, FA98B, ZN326, PREP, RRP12, SYAM, EXOS6, CAF17, UBR4, NT5D1, PDE12, JMJD6, CDC73, EDC4, PRP8, RL22L, SYDM, GGYF2, HSDL2, TM10C, ZCCHV, DHX29, DCXR, HUWE1, ACOT1, KTN1, CARM1, STX12, HORN, SPB1, SRRM1, SUV3, TXND5, SCPDL, FA98A, PCAT1, FAD1, UBA3, NEK9, BRX1, ZC3HF, SCFD1, HNRLL, ATX2L, PSPC1, P66B, DNJC9, DDX1, H1X, PSMF1, RT27, LAR4B, ARC1A, RENT1, FUBP1, P5CR2, TRM61, ZCCHL, PGAM5, FUBP3, SPF45, THOC3, ZFR, SNX27, RBM14, PRPK, TBCB, CDC5L, PARK7, HCD2, ROAA, EBP2, VRK1, NIPS1, MEP50, TBA1C, ERP44, NTPCR, DDX23, MTNA, NTM1A, TM109, SYTM, THIC, RBM4, HDHD5, ITPA, EIF2A, PDIP3, MK67I, GTPB4, REN3B, API5, UBE2O, WDR12, SLIRP, NAA50, ILKAP, SLK, PININ, YTDC2, RPF2, QTRT2, ARMT1, CSN7B, ELP3, KT3K, MRM3, GLOD4, MCCB, CWC22, WDR6, VTA1, EXOS4, INO1, LUC7L, TIGAR, XPP1, SIAS, PHP14, HELLS, ECHD1, RBM12, DD19A, SEP11, TBC13, ATD3A, DDX18, PNPO, RBM28, LYAR, DPP3, BCLF1, F120A, HPBP1, MAT2B, RRBP1, GMPR2, GRHPR, TES, CHRD1, SEPT9, EI2BD, DBNL, DDX41, APC7, STML2, MRT4, ACINU, NUP50, PSME2, MYO6, CHIP, CSN3, SRRM2, CD11A, SMC3, RTRAF, PIN4, PLAP, NUDC, COF2, AP3M1, TR150, NOP58, SGT1, SYYM, SBDS, EXOS1, SF3B6, RRP15, RT23, STRAP, CHTOP, SAMH1, TLN1, HYOU1, ATG4B, TBL2, PRC2C, PPME1, YTHD2, SNX9, SERC, CLIC4, DC1L1, S23IP
Species: Homo sapiens
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Ramirez DH, Aonbangkhen C, Wu HY, Naftaly JA, Tang S, O'Meara TR, Woo CM. Engineering a Proximity-Directed O-GlcNAc Transferase for Selective Protein O-GlcNAcylation in Cells. ACS chemical biology 2020 15(4) 32119511
Abstract:
O-Linked β-N-acetylglucosamine (O-GlcNAc) is a monosaccharide that plays an essential role in cellular signaling throughout the nucleocytoplasmic proteome of eukaryotic cells. Strategies for selectively increasing O-GlcNAc levels on a target protein in cells would accelerate studies of this essential modification. Here, we report a generalizable strategy for introducing O-GlcNAc into selected target proteins in cells using a nanobody as a proximity-directing agent fused to O-GlcNAc transferase (OGT). Fusion of a nanobody that recognizes GFP (nGFP) or a nanobody that recognizes the four-amino acid sequence EPEA (nEPEA) to OGT yielded nanobody-OGT constructs that selectively delivered O-GlcNAc to a series of tagged target proteins (e.g., JunB, cJun, and Nup62). Truncation of the tetratricopeptide repeat domain as in OGT(4) increased selectivity for the target protein through the nanobody by reducing global elevation of O-GlcNAc levels in the cell. Quantitative chemical proteomics confirmed the increase in O-GlcNAc to the target protein by nanobody-OGT(4). Glycoproteomics revealed that nanobody-OGT(4) or full-length OGT produced a similar glycosite profile on the target protein JunB and Nup62. Finally, we demonstrate the ability to selectively target endogenous α-synuclein for O-GlcNAcylation in HEK293T cells. These first proximity-directed OGT constructs provide a flexible strategy for targeting additional proteins and a template for further engineering of OGT and the O-GlcNAc proteome in the future. The use of a nanobody to redirect OGT substrate selection for glycosylation of desired proteins in cells may further constitute a generalizable strategy for controlling a broader array of post-translational modifications in cells.
O-GlcNAc proteins:
SBNO1, CNOT1, P121C, DX39A, GTPB1, AP3B1, PGRC1, TAF4, EIF3F, IPO5, IF2B3, NOP56, DDX3X, ARI1A, IRS4, ANM5, TCRG1, PSA7, HGS, MYPT1, HNRDL, XPO1, SET1A, PUR4, NPC1, TIF1A, NKRF, OGT1, PPM1G, EIF3D, EIF3H, DHX15, SERA, HNRPR, IF4G3, E41L2, ZN207, BUB3, ACTN4, HTSF1, AP1G1, SYNC, AKAP8, CALU, SMCA5, JIP4, OGA, HNRPQ, DIAP1, TSN3, SNX2, DKC1, CLAP2, CPNE3, PHF2, ANR17, H2AY, FLNB, NCOR1, CISY, PR40A, SF3B1, CSDE1, U520, EIF3G, PRAF3, SRP72, MTA2, TOX4, SC24D, SC31A, SCAF4, ZRAB2, LC7L3, VAPB, IPO7, SC24B, ACSL3, AP2A1, AIFM1, LDHA, COX2, HPRT, AATM, PGK1, LMNA, TFR1, ALDOA, OAT, G3P, RPN1, RPN2, AT1A1, ADT2, PCCA, IF2A, RLA0, ITB1, ATPB, ENOA, PYGL, G6PI, NPM, LDHB, PDIA1, H10, TBB5, HEXB, PROF1, SYEP, HS90A, HNRPC, 4F2, HS90B, ASNS, ODPA, RU17, RSSA, SNRPA, GSTP1, HMGB1, DLDH, ROA1, PARP1, LKHA4, HS71B, H14, ODP2, THIO, CH60, BIP, HSP7C, EPB41, ODPB, LAMP1, ACADM, TOP1, TOP2A, PYC, C1TC, MPRI, PRPS2, PABP1, PCNA, HARS1, IMDH2, TPR, KCRB, XRCC6, XRCC5, EF2, PDIA4, PLST, GLU2B, KPYM, ENPL, PO2F1, HNRPL, SYDC, PLAK, ALDR, EZRI, GNS, RS2, CREB1, H12, AT2A2, JUNB, PYRG1, DDX5, PRS6A, TCPA, RL35A, RL7, VINC, SON, RCC1, NUCL, HXK1, E2AK2, SPEE, IF2B, ANXA7, LMNB1, FLNA, VDAC1, FBRL, PUR2, PUR6, UBA1, NDKB, ROA2, RFX1, TCEA1, SFPQ, PPIB, RS3, NFYA, SAHH, COF1, IF4B, EF1B, MCM3, BRD2, ATPA, PSA1, PSA3, PSA4, PAX6, U2AF2, RL13, PTBP1, SYTC, SYVC, EF1G, RFA1, APEX1, PYR1, CALR, MAP4, CALX, PSB5, TKT, PRDX6, PRDX5, PRDX3, RL12, PEBP1, PDIA3, 2AAA, CDC27, AMRP, SDHA, QCR1, PUR9, HNRH1, STIP1, PRDX2, RL9, CSTF2, MCM4, MCM5, MCM7, GLYM, HSP74, PHB, MYH9, COPB2, ADDA, BASI, FUS, NU214, DEK, MP2K2, ATPG, RL4, SRP14, NUP62, RBMX, GRP75, IF4A3, RS19, RL3, TXLNA, TCPZ, MDHC, MDHM, ECHA, IF2G, GARS, SYIC, LAP2A, LAP2B, MTREX, RS27, LPPRC, MATR3, RANG, VDAC2, UBP5, KI67, RAGP1, NOP2, CRKL, BAG6, RL27A, RL5, RL21, RL28, RS9, STT3A, COPD, PRC2A, TCPE, AL9A1, RL34, NASP, FAS, TCPG, EFTU, SYAC, SYSC, PSB3, MCM2, YLPM1, TMEDA, RBM25, NU153, RBP2, GSK3A, TAF6, GUAA, MRE11, GDIB, EMD, F10A1, LRBA, RL14, TCPQ, TCPD, ANX11, PAPOA, RAB7A, SMCA4, HCFC1, DHB4, ROA3, 6PGD, HNRPM, IMA1, AGFG1, HNRPF, MSH6, RBM5, NUP98, ACLY, COPB, COPA, MOT1, SC24C, SYRC, SYYC, AT1B3, RD23B, P5CS, IF5, XPO2, TERA, AFAD, DSRAD, PSA, SYMC, CTBP2, NU107, TPIS, ACTB, IF4A1, PSA6, ARF3, ABCE1, RAP1B, RS3A, RL26, RL15, S61A1, HNRPK, RS7, RS8, 1433E, RS14, RS23, RS11, SMD1, RL7A, RS4X, H4, RAN, RL23, GBB2, RL10A, RL11, RL8, PPIA, RL40, TRA2B, AP2B1, IF5A1, RACK1, YBOX1, EF1A1, TBB4B, CSK21, IF4G2, GTF2I, TCPB, PRKDC, RL24, RL19, SRSF3, FOXK1, RBM10, MPCP, CLH1, HNRPU, SPTB2, FOXK2, CAP1, LAT1, EXOSX, EWS, RL18A, FKBP4, RL6, TOP2B, KMT2A, LMNB2, TF65, IF4G1, TLE3, SRS11, PUR1, SUH, GABPA, PRDX1, RL18, C1QBP, KHDR1, SRSF1, DHX9, CD47, SSRP1, RBBP4, AHNK, AP1B1, NU160, BPTF, TP53B, AIMP1, ILF2, ILF3, LMAN2, TRAP1, PP1R8, ACACA, ROA0, PRDX4, CBX3, PSMD2, SRSF6, TIF1B, PTK7, PABP4, EIF3I, TCOF, SF3B2, TMED1, PICAL, RIPK1, HDAC1, CUL4B, CD166, IDI1, NFYC, CKAP5, HNRPD, SCRB2, DSG2, EIF3A, UBP2L, SCRIB, TTL12, DCTN1, DYHC1, SRC8, CAPR1, RBM39, MCM6, MDC1, EPN4, SMC1A, RRP1B, UBP10, GANAB, LBR, ZN638, IMB1, NUMA1, SEPT2, SART3, U5S1, SYK, BRD3, PDIA6, IPYR, TEBP, NONO, PWP2, RCN1, PCBP1, PCBP2, SF3B3, SAFB1, SF3A1, NCOA2, SF01, MARE1, NSDHL, TAB1, AAAT, VAS1, ZYX, CCDC6, PKN2, DDB1, CDC37, SRSF7, CPSF6, NRF1, H31T, QSER1, QRIC1, P3H1, TB10B, AMOT, DHB12, PRC2B, H2B2F, HP1B3, CE170, ZC3HD, RBM26, RIF1, RPRD2, ZN318, ECM29, ZMYM4, MAP1S, LIN54, EDC4, PRP8, SCYL2, NFRKB, ZC3HE, LARP1, FIP1, MCAF1, GGYF2, SPT6H, SND1, DHX30, KDM3B, ZCCHV, NUP54, POGZ, NUFP2, MAVS, I2BP2, RBBP6, HUWE1, YTHD3, CENPV, LYRIC, ZN598, GP180, CAND1, CARM1, DDX42, P66A, ARI3B, MGAP, PHF6, CHERP, ANKH1, SUGP1, CCAR1, SPB1, PHAR4, SPART, CCAR2, NUP93, S11IP, FNBP4, CPSF7, ARFG1, ENAH, AFG2H, TXND5, LS14A, Z280C, TNR6A, SMRC2, TBC15, PNPT1, HM13, PO210, GEMI5, ZN384, SMAP2, NU133, PDC6I, PCNP, CKAP2, ATX2L, P66B, ELYS, DDX1, GBF1, NICA, UBXN4, HS105, LAR4B, NU205, AKAP1, TFG, CBP, DDX17, CELF1, RENT1, SMRC1, FUBP2, TNPO1, UBP7, NCLN, FUBP1, FKB10, KBP, PDLI5, FUBP3, CHAP1, Z512B, ZFR, PRRC1, DOCK7, RBM14, VPS35, CIC, EFGM, SIN3A, MINT, CDC5L, PSMD1, EYA3, ATX2, HCD2, ACON, TS101, TCPH, ANM1, SH3G1, COR1B, DIDO1, HNRL1, DDX23, TMED9, NUP58, RBM4, NAA15, B2L13, YTHD1, UNK, ILKAP, SP130, BRD8, I2BPL, SLK, S6A15, PININ, NELFA, PTN23, WNK1, AMPB, GORS2, CYBP, TAF9B, GLOD4, CBX8, NCOA5, CHD8, APMAP, DCP1A, RTN4, ANLN, GEPH, PDLI7, DDX21, SYFB, SYIM, SMC4, RBM12, DDX18, CARF, UGGG1, CDK12, TECR, IF2B1, HPBP1, ITSN2, CNOT2, HACD3, RCC2, SYLC, SUCB1, UBQL2, PCYOX, S30BP, PUF60, NRBP, DACH1, BAZ2A, BAZ1B, CDC23, TASOR, ACINU, CDV3, MRTFB, YETS2, HECD1, PKCB1, DD19B, PRP19, MAGD2, FAF1, TRI33, SRRM2, PA2G4, RUVB2, RUVB1, VDAC3, E41L3, TR150, NOP58, SHLB1, LC7L2, TMED7, STRAP, RTCB, HBS1L, TLN1, HYOU1, PRC2C, SP16H, COPG1, DC1L1, S23IP
Species: Homo sapiens
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Zhu Y, Willems LI, Salas D, Cecioni S, Wu WB, Foster LJ, Vocadlo DJ. Tandem Bioorthogonal Labeling Uncovers Endogenous Cotranslationally O-GlcNAc Modified Nascent Proteins. Journal of the American Chemical Society 2020 142(37) 32870666
Abstract:
Hundreds of nuclear, cytoplasmic, and mitochondrial proteins within multicellular eukaryotes have hydroxyl groups of specific serine and threonine residues modified by the monosaccharide N-acetylglucosamine (GlcNAc). This modification, known as O-GlcNAc, has emerged as a central regulator of both cell physiology and human health. A key emerging function of O-GlcNAc appears to be to regulate cellular protein homeostasis. We previously showed, using overexpressed model proteins, that O-GlcNAc modification can occur cotranslationally and that this process prevents premature degradation of such nascent polypeptide chains. Here, we use tandem metabolic engineering strategies to label endogenously occurring nascent polypeptide chains within cells using O-propargyl-puromycin (OPP) and target the specific subset of nascent chains that are cotranslationally glycosylated with O-GlcNAc by metabolic saccharide engineering using tetra-O-acetyl-2-N-azidoacetyl-2-deoxy-d-galactopyranose (Ac4GalNAz). Using various combinations of sequential chemoselective ligation strategies, we go on to tag these analytes with a series of labels, allowing us to define conditions that enable their robust labeling. Two-step enrichment of these glycosylated nascent chains, combined with shotgun proteomics, allows us to identify a set of endogenous cotranslationally O-GlcNAc modified proteins. Using alternative targeted methods, we examine three of these identified proteins and further validate their cotranslational O-GlcNAcylation. These findings detail strategies to enable isolation and identification of extremely low abundance endogenous analytes present within complex protein mixtures. Moreover, this work opens the way to studies directed at understanding the roles of O-GlcNAc and other cotranslational protein modifications and should stimulate an improved understanding of the role of O-GlcNAc in cytoplasmic protein quality control and proteostasis.
Species: Homo sapiens
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Phoomak C, Park D, Silsirivanit A, Sawanyawisuth K, Vaeteewoottacharn K, Detarya M, Wongkham C, Lebrilla CB, Wongkham S. O-GlcNAc-induced nuclear translocation of hnRNP-K is associated with progression and metastasis of cholangiocarcinoma. Molecular oncology 2019 13(2) 30444036
Abstract:
O-GlcNAcylation is a key post-translational modification that modifies the functions of proteins. Associations between O-GlcNAcylation, shorter survival of cholangiocarcinoma (CCA) patients, and increased migration/invasion of CCA cell lines have been reported. However, the specific O-GlcNAcylated proteins (OGPs) that participate in promotion of CCA progression are poorly understood. OGPs were isolated from human CCA cell lines, KKU-213 and KKU-214, using a click chemistry-based enzymatic labeling system, identified using LC-MS/MS, and searched against an OGP database. From the proteomic analysis, a total of 21 OGPs related to cancer progression were identified, of which 12 have not been previously reported. Among these, hnRNP-K, a multifaceted RNA- and DNA-binding protein known as a pre-mRNA-binding protein, was one of the most abundantly expressed, suggesting its involvement in CCA progression. O-GlcNAcylation of hnRNP-K was further verified by anti-OGP/anti-hnRNP-K immunoprecipitations and sWGA pull-down assays. The perpetuation of CCA by hnRNP-K was evaluated using siRNA, which revealed modulation of cyclin D1, XIAP, EMT markers, and MMP2 and MMP7 expression. In native CCA cells, hnRNP-K was primarily localized in the nucleus; however, when O-GlcNAcylation was suppressed, hnRNP-K was retained in the cytoplasm. These data signify an association between nuclear accumulation of hnRNP-K and the migratory capabilities of CCA cells. In human CCA tissues, expression of nuclear hnRNP-K was positively correlated with high O-GlcNAcylation levels, metastatic stage, and shorter survival of CCA patients. This study demonstrates the significance of O-GlcNAcylation on the nuclear translocation of hnRNP-K and its impact on the progression of CCA.
Species: Homo sapiens
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Hahne H, Sobotzki N, Nyberg T, Helm D, Borodkin VS, van Aalten DM, Agnew B, Kuster B. Proteome wide purification and identification of O-GlcNAc-modified proteins using click chemistry and mass spectrometry. Journal of proteome research 2013 12(2) 23301498
Abstract:
The post-translational modification of proteins with N-acetylglucosamine (O-GlcNAc) is involved in the regulation of a wide variety of cellular processes and associated with a number of chronic diseases. Despite its emerging biological significance, the systematic identification of O-GlcNAc proteins is still challenging. In the present study, we demonstrate a significantly improved O-GlcNAc protein enrichment procedure, which exploits metabolic labeling of cells by azide-modified GlcNAc and copper-mediated Click chemistry for purification of modified proteins on an alkyne-resin. On-resin proteolysis using trypsin followed by LC-MS/MS afforded the identification of around 1500 O-GlcNAc proteins from a single cell line. Subsequent elution of covalently resin bound O-GlcNAc peptides using selective β-elimination enabled the identification of 185 O-GlcNAc modification sites on 80 proteins. To demonstrate the practical utility of the developed approach, we studied the global effects of the O-GlcNAcase inhibitor GlcNAcstatin G on the level of O-GlcNAc modification of cellular proteins. About 200 proteins including several key players involved in the hexosamine signaling pathway showed significantly increased O-GlcNAcylation levels in response to the drug, which further strengthens the link of O-GlcNAc protein modification to cellular nutrient sensing and response.
O-GlcNAc proteins:
UBA6, SHOT1, MEX3A, TPC13, CNOT1, PGP, CE051, SMHD1, CC85C, I4E1B, ZBBX, B2R8K8, B2RB32, B3KNS4, B3KUR4, B4DF70, B4DIH0, B4DIR9, B4DTN6, B8XCX8, DX39A, BACH, GTPB1, SMAP, AP3B1, PSD12, PSMD9, ATOX1, PGRC1, SPT5H, TAF4, DFFA, CLIC1, EIF3F, WASL, IPO5, EF2K, PLOD2, MEIS1, PSDE, IMA4, BCL9, SDCB1, NOP56, IMA3, ARI1A, UBFD1, CCS, DVL2, KMT2D, ANM5, TNPO2, GEMI2, HAT1, MYPT1, GAK, XPO1, ZN609, SC16A, SR140, PUR4, NKRF, ZN185, CASC3, OGT1, PMM2, HMGB3, PPM1G, SHIP2, NVL, NUP42, R113A, KDM6A, DHX15, MCES, RRP8, SERA, DC1L2, ZW10, M3K7, RIPK2, WDR62, TXNL1, IF4G3, E41L2, FOXO3, DENR, XPOT, BUB3, ACTN4, HTSF1, SGTA, SYNC, LANC1, DHX16, ZNRD2, KPRB, AQR, GANP, HNRPQ, BUB1B, DIAP1, PLIN3, ANM3, CTND1, EIF1B, USO1, IF2P, NBN, SRGP2, N4BP1, ROCK2, CLAP2, CPNE3, BRE1B, ZC11A, CLU, T22D2, ANR17, GGCT, HMMR, VATG1, FLNB, NCOR1, SPAG7, PR40A, GGYF1, PSIP1, NDUS3, SRS10, KHDR3, SF3B1, CSDE1, PRKRA, KS6A5, NPM3, LYPA1, U520, TIPRL, OFD1, WDHD1, CD123, EIF3G, PSD10, SPF27, CIAO1, RMP, TOX4, SC24D, ST65G, UBXN7, PLPHP, NFAT5, SOGA1, UBP19, WDR47, SC31A, HEXI1, UBR5, SCAF4, UBE4B, ELP1, ZRAB2, KIF4A, VAPB, SNAPN, 6PGL, SMC2, ATG7, IPO7, AGM1, ZFYV9, SVIL, BAG4, AHSA1, PARN, PSMG1, SC24A, SC24B, AGRL2, TYDP2, PCNT, YETS4, STABP, ASML, EYA4, CEP43, HS74L, WIZ, STAU1, BAG2, BAG3, BPNT1, ECD, MBD3, BCL10, MOC2B, TOM40, CHK2, GSHR, PNPH, HPRT, ADA, CYTB, OAT, KITH, CPNS1, P53, HSPB1, TYSY, RPN1, RLA1, JUN, LA, NPM, NFL, NFM, ANXA2, SYEP, HS90A, HNRPC, SP1, 4F2, HS90B, ASNS, VIME, GSTP1, LEG1, HMGB1, UCHL1, LKHA4, H2A1, UBC, GLI2, ODP2, LAMP1, G6PD, ADHX, CDK4, SRF, PCNA, HARS1, ADT3, IMDH2, ACTN1, PEPD, XRCC5, LAMP2, RINI, EF2, PLST, ACPH, KAP2, CD99, GLU2B, AK1A1, KPYM, PO2F1, SYDC, ALDR, ERF3A, GNS, ZEP1, DESP, CREB1, GCFC2, UBF1, ATF7, CAN2, PYRG1, TCPA, SON, NELFE, ATF1, NFKB1, ICAL, IMDH1, GSTM3, FLNA, COMT, FBRL, PUR2, PUR6, UBA1, ENPP1, CBL, SP100, NFYA, SAHH, COF1, TENA, THTM, RS12, DNJB1, PSA4, DNMT1, RAE2, PTBP1, SYTC, 1433T, RFA1, APEX1, PYR1, MAP4, CALX, PSA5, NDUS1, TPP2, SHC1, TKT, PML, MARCS, PRDX6, PRDX5, PRDX3, KCY, 2AAA, 2AAB, CDC27, NMT1, SDHA, GDIA, METK2, DNJA1, AKT1, PUR9, HNRH1, S10AB, ARRB2, DCTD, TF2H1, KINH, CSTF2, DUT, TTK, PROF2, CAH8, RFC4, RFC2, HXD13, MYH9, MYH10, ADDA, BASI, NU214, DEK, MYH11, PPM1A, PRS7, ARL3, MP2K2, RL4, 8ODP, NUP62, ZEB1, TAGL2, COIL, VATA, GRP75, TXLNA, STAT3, PEX19, RFC5, RFC3, IF2G, NAA10, KDM5C, CSK, GARS, SYIC, LAP2A, LAP2B, STAT1, EPS15, CASP3, LPPRC, HD, MSH2, GPDM, RANG, VDAC2, CBX5, UBP5, MK09, KI67, RECQ1, ATRX, CRKL, RL21, MAP1B, IQGA1, GLYG, SYQ, ID4, TISD, PPCE, RFX5, GSH0, PRC2A, NEST, RPIA, NASP, FAS, ARRB1, CENPF, SRP09, PCY1A, SYAC, SYSC, PSB2, ACOT2, RBM25, NUMB, NU153, RBP2, GUAA, MRE11, SERPH, PDLI4, VASP, MAP2, LRBA, TCPD, FXR1, FXR2, RAB5C, DUS3, GALK1, HCFC1, IRAK1, UBP11, HDGF, ROA3, 6PGD, IMA1, GDS1, GDIR1, AGFG1, HNRPF, TF2AA, MSH6, KIF11, ZN143, HXK2, THOP1, NUP98, BIEA, PPP5, ARFP2, NUBP1, ACLY, COPB, CCHL, ICLN, SYRC, SYYC, AAPK2, BCAT1, RD23B, KAD2, P5CS, IF5, XPO2, TERA, AFAD, AF17, ADK, DSRAD, PKNX1, ALR, HNRH2, EIF3B, BID, RRP1, IF6, GEMI4, EPIPL, MTPN, TPIS, EIF3E, MYL6, ACTB, IF4A1, RS20, PRPS1, S10AA, DEST, CSN2, ABCE1, RS3A, PSME3, MGN, PFD3, HNRPK, PP1A, RS15A, RUXF, PPIA, RS27A, AP2B1, 1433Z, DYL1, SKP1, IF5A1, RACK1, UBC9, UB2L3, ACTS, CSK21, PA1B2, IGBP1, IF4G2, GTF2I, PI42B, TCPB, RAE1L, GSTO1, PRKDC, TMF1, RL19, SRSF3, MXRA7, FOXK1, XIAP, TFAM, VIGLN, PURA, BORG5, CLH1, NFKB2, U2AF1, FOXK2, AMPD2, NC2B, TFAP4, OTUD4, EWS, CDR2, TOP2B, AKA12, KMT2A, UBXN1, IF4G1, K1C17, LGUL, REL, 1433F, UBE3A, PTN12, SRS11, CALD1, PTN11, PUR1, GFPT1, PSME1, GABPA, RING1, FMR1, KHDR1, KLC1, SRSF1, DHX9, GOGA2, LG3BP, SSRP1, VAC14, NSUN2, RBBP4, NCBP1, AHNK, HSP7E, SCRN1, NU160, FOXO1, TBL3, TFCP2, GRSF1, SFSWA, TP53B, ILF3, ELAV2, TRAP1, FOXC1, TAF10, NFAC3, CSTF3, UBP4, CAF1A, RED, MTAP, LIPA1, FADD, PRDX4, CBX3, STK3, PSMD2, SBP1, SRSF6, TIF1B, PABP4, IFIT5, PHLP, UB2V1, DC1I2, PPIG, TCOF, GOGA4, FKBP5, GAB1, 4EBP1, 4EBP2, RIPK1, HDAC1, KCC2B, DCTN2, ULA1, SNW1, SNX1, PWP1, CUL2, DYR1A, AAMP, DX39B, BLMH, CKAP5, DAG1, BOP1, UBE4A, KEAP1, SAFB2, UBP2L, SCRIB, TTL12, DPOA2, WRN, DPYL3, DYHC1, NPAT, SRC8, TRI25, FLNC, FRG1, CAPR1, SLBP, MCM6, PUM1, MDC1, SMC1A, RRP1B, NCOA6, UBP10, 2A5D, MEF2D, CHD4, NFAC4, ZN638, IMB1, NUMA1, NAA25, CND2, FL2D, CND1, EXOS7, U5S1, R3HD1, RB3GP, SETB1, RRS1, WDR43, PLEC, PCM1, IPYR, TEBP, NONO, RBBP5, L2GL1, PCBP1, ELOB, SF3B3, CNN3, KS6A1, SAFB1, SF3B4, SC23A, SF3A1, SKIV2, TCEA2, NCOA2, TSN, TRBP2, SF01, PCH2, MED1, TRIP6, ELAV1, ELF2, TAB1, AAAT, SMAD2, TBCE, ZYX, ADRM1, UAP1, E2F4, AINX, DDB1, CDC37, RBBP7, SMN, DREB, NRF1, KNOP1, HNRL2, INF2, PDS5A, QSER1, CL16A, ERC6L, IAH1, QRIC1, CTU2, LRRF1, RA1L2, RPA43, MA7D1, RABL6, CE022, VP26B, LARP7, ARI4B, CBWD6, BCL7A, PDCD4, CRTC2, H1BP3, Q59GF8, ZN326, RIPL1, PP6R3, SH319, WDR44, PRC2B, SYAM, DOP1, SMAG2, TBCEL, TDIF2, EXOS6, CE170, CHCH9, FKB15, ZC3HD, GPTC4, LRIF1, RBM20, UBR4, BEND3, UBAP2, RBM26, TUT4, ATPF1, DDI2, NT5D1, LYRM7, RIF1, CE350, RPRD2, ANGE2, BRE1A, MYOME, ZN318, ECM29, ZMYM4, FHI2A, K2C79, LAR1B, MAP1S, AR6P4, KAZRN, ARID2, INT3, CDCA2, C19L1, PDPK2, HEAT6, WWC2, CARL2, CPIN1, SMYD5, OTU7B, ELP2, CNDH2, SDE2, LIN54, IPP2B, ANR54, JMJD6, PPR18, ZCHC8, PMF1, F117B, C2D1A, MET2B, ZN529, EDC4, CEP85, SCYL2, TTC27, WDR73, NFRKB, RSBNL, AAGAB, TTC33, FIGL1, MDEAS, BRAT1, ZC3HE, LARP1, BL1S3, C1TM, AFTIN, FIP1, CRTC1, SAS6, MCAF1, RBP10, BCOR, RFIP1, CA122, YJ005, UBN2, STXB4, LARP4, NOL8, SND1, DDX46, RUFY3, TM10C, EIF3M, CD37L, CSN6, KDM3B, ZCCHV, EFL1, NUP54, POGZ, ZFY16, MON2, MAVS, CLAP1, HAUS6, HEAT3, HDGR2, EMSY, RAI1, WAPL, R3HCL, RBBP6, CE024, SH3R1, HUWE1, CNTRL, UB2Q1, CK096, PRUN1, YRDC, ZN546, UBP48, LUZP1, ZCH18, CAND1, GLCI1, HOOK3, PARG, CCD25, CDR2L, NIT1, PRSR1, SPAS2, HOME1, P66A, C2CD5, I2BP1, KCC1D, CRLF3, PLD3, VRK3, NUDC3, ARI3B, CEP97, MA7D3, CC117, NXP20, MGAP, ANKH1, SUGP1, SUGP2, CCAR1, SKA3, PHC2, BAP18, RB12B, SPB1, SRRM1, DI3L2, ZZZ3, CA174, P20D2, PHAR4, TRM2A, DCP1B, PELP1, TTC5, CCAR2, NUP93, LRC47, MTURN, MILK1, STAG2, FNBP4, KIF2B, MIA40, OXR1, ZGPAT, TAB3, RM50, ZC3H8, KKCC1, OTU6B, CA052, EFNMT, UBR7, CE112, PTGR2, RM43, JMY, ZIC4, ASCC1, BANP, EIF1A, ANR52, AFG2H, NHLC2, ATAD1, PAIRB, NF2IP, RN169, LS14A, SLAI1, TNR6A, PHC3, SIMC1, ABCF1, DDHD1, KMT2C, BD1L1, NUP35, GRPE2, CX038, FBX30, PI42C, CIP2A, ZN507, C99L2, NEK9, BL1S5, GEMI5, PARD3, IPO4, PP4R1, ZN384, SETD7, ZC3HF, PPIL4, CCD12, NU133, PDC6I, CHAC2, ZFN2B, LEO1, HNRLL, UBCP1, TRUB1, CKAP2, ATX2L, PALLD, SREK1, GEMI6, SRS12, P66B, DNJC9, BBX, PPM1E, DGCR8, IRGQ, DDX1, RTF1, TOPB1, WASF1, TSC1, UBXN4, PHF3, MAML1, NDRG1, HS105, SEPT8, LAR4B, GCN1, NU205, ANS1A, PGTA, TFG, ARC1A, HDAC2, TANK, RAD50, CELF1, ARHG1, CSN5, GPKOW, SMRC1, FUBP2, TNPO1, ARHG2, GLMN, UBP7, LPP, TCAL3, TSR2, RIFK, RRF2M, MRTFA, GTPB3, FAKD4, CPNE4, SYAP1, R51A1, CR025, PF21A, RNF25, OTUL, UPP, PTER, ATG2B, EFHD2, CSTOS, DCPS, PP14B, ENTR1, COAC, FWCH2, KCD12, CMBL, ATG4C, RMXL1, SIR1, T3HPD, SGTB, TMA16, MMAB, CYFP2, NRBF2, CCD97, EDC3, THOC1, OTUB1, HMCES, DUS3L, NUD11, OTUD5, LTV1, A7L3B, GWL, MTNB, LIN37, ERO1A, MED30, NAF1, PGAM5, SCLY, FUBP3, NSL1, PAWR, MPP4, ZFP91, UBP47, TOPK, ZFR, EP400, SNX27, Q96LS1, RNPC3, AP4AT, ZN583, SCLT1, RPR1A, IPO9, RBM14, QKI, F193B, PUS7, KI20B, RAB3I, VPS35, PURB, SPAG5, NACC1, PASK, CIC, KKCC2, NUDC1, UBP28, WRIP1, RANB9, SRPK1, CK5P2, SIN3A, INT14, AEDO, MINT, MMS19, UHRF1, TBCB, CNN2, RING2, EYA3, LGMN, NUP88, SCAFB, EIF3C, TTC1, DNJC7, KIF2C, HCD2, SNAG, TS101, CPNE1, HAUS7, ANM1, H2B1L, BAG1, SH3G1, AKAP9, CND3, F118B, GRWD1, MACD1, MEP50, FYCO1, CPPED, CG050, LZTS2, PDD2L, GPTC1, LLPH, SLD5, PSF3, CENPK, NTPCR, TACO1, ESYT1, UBAC1, CCM2, PSMG3, WAC, RBM42, MCMBP, AN32E, TBCD, HGH1, MMTA2, DOHH, SPNDC, PAXX, MSTO1, TCAB1, CA050, WDR18, THUM3, NTM1A, ASHWN, PBDC1, UT14A, NUP58, DPCD, GNL3, NUP85, RPAP1, NADAP, OSB11, NAA15, ITPA, GNB1L, MK67I, YTHD1, STK33, SETD2, ASPC1, UCK2, API5, UNK, TB182, UBE2O, CEP44, PITH1, SENP6, EGLN1, PIP30, MFR1L, PAIP1, CLPB, KLC2, ILKAP, XRN2, BRD8, NSRP1, CSTFT, HDHD2, SIL1, APC1, I2BPL, ASCC2, RBGPR, PININ, CPVL, BOLA2, GCP60, PTN23, UN45A, DSN1, IFG15, WNK1, AMPB, SMRCD, HEAT1, RABE2, CCD86, NOL6, WDCP, YTDC2, RPAP3, WDR55, RANB3, DCTP1, PEAK1, WDR26, SFR19, PHAX, NOL11, MINY3, GORS2, QTRT2, ARMT1, CNO10, EHMT1, VP37B, RM44, CSN7B, NHEJ1, NJMU, PKHA5, XPO5, CYBP, RRAGC, PATZ1, EMAL4, GLOD4, GPBL1, NCOA5, TNR6C, ZDBF2, GOPC, SYSM, INO1, BRD7, DCP1A, MED4, RIC8A, TIGAR, COPRS, HINT3, RRAGD, PFD4, AVEN, EXOS5, EXOS3, ANLN, XPP1, BIRC6, DDX21, SHLB2, EI2BG, MBNL1, CHRC1, AAAS, YAE1, STRN4, PNO1, LANC2, KIF15, RAD18, SYIM, KLC4, ATG3, OLA1, ZF64B, RBM12, P