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Wong YK, Wang J, Lim TK, Lin Q, Yap CT, Shen HM. O-GlcNAcylation promotes fatty acid synthase activity under nutritional stress as a pro-survival mechanism in cancer cells. Proteomics 2022 35083852
Abstract:
Protein O-GlcNAcylation is a specific form of protein glycosylation that targets a wide range of proteins with important functions. O-GlcNAcylation is known to be deregulated in cancer and has been linked to multiple aspects of cancer pathology. Despite its ubiquity and importance, the current understanding of the role of O-GlcNAcylation in the stress response remains limited. In this study, we performed a quantitative chemical proteomics-based open study of the O-GlcNAcome in HeLa cells, and identified 163 differentially-glycosylated proteins under starvation, involving multiple metabolic pathways. Among them, fatty acid metabolism was found to be targeted and subsequent analysis confirmed that fatty acid synthase (FASN) is O-GlcNAcylated. O-GlcNAcylation led to enhanced de novo fatty acid synthesis activity, and fatty acids contributed to the cytoprotective effects of O-GlcNAcylation under starvation. Moreover, dual inhibition of O-GlcNAcylation and FASN displayed a strong synergistic effect in vitro in inducing cell death in cancer cells. Together, the results from this study provide novel insights into the role of O-GlcNAcylation in the nutritional stress response and suggest the potential of combining inhibition of O-GlcNAcylation and fatty acid synthesis in cancer therapy. This article is protected by copyright. All rights reserved.
O-GlcNAc proteins:
RUXGL, ADAS, DX39A, MYO1C, IPO5, PESC, NOP56, DDX3X, SCD, MGST3, HNRDL, XPO1, SURF4, OGT1, PPM1G, MOT4, DHX15, CYB5B, SERA, HNRPR, BUB3, ACTN4, MYO1B, GANP, HNRPQ, NDUS7, MPU1, H2AY, FLNB, SC22B, SF3B1, U520, UTP20, NU155, ATP5H, RL1D1, MTA2, RTN3, VAPB, IPO7, ACSL3, BAG2, TOM40, LDHA, DHE3, AATM, PGK1, ASSY, LMNA, TFR1, ALDOA, K2C1, G3P, HSPB1, RPN1, AT1A1, ADT2, PCCA, RLA1, RLA0, LA, K1C18, K2C8, ATPB, ENOA, NPM, TPM3, LDHB, PDIA1, ANXA2, TBB5, TRY1, PROF1, SYEP, HS90A, HNRPC, DAF, 4F2, HS90B, ODPA, RU17, VIME, RS17, K2C7, GNAI3, RSSA, LEG1, ROA1, PARP1, PRS56, HS71B, ODP2, THIO, MGST1, CH60, BIP, HSP7C, GTR1, TOP2A, PYC, PABP1, PCNA, ADT3, IMDH2, KCRU, XRCC6, XRCC5, EF2, K1C10, K2C5, PDIA4, PLST, ETFA, MIF, KPYM, ENPL, HNRPL, PLAK, EZRI, NDKA, RS2, DESP, H13, NCPR, AT2A2, DDX5, TCPA, PTN1, ARF4, RL7, RL17, NUCL, GSTM3, FLNA, FBRL, PUR6, UBA1, ROA2, QCR2, SFPQ, PPIB, RS3, SAHH, COF1, MCM3, RS12, ATPA, U2AF2, RL13, S10A4, PTBP1, SYVC, EF1G, STOM, RL10, APEX1, PYR1, CALX, TKT, ERP29, PRDX6, PRDX5, PRDX3, RL12, PDIA3, CPSM, HNRH1, STIP1, L1CAM, PRDX2, P5CR1, DUT, MCM7, GLYM, HSP74, PHB1, RL22, MYH9, SOAT1, DEK, K22E, RL4, LONM, NUP62, GRP75, IF4A3, RL3, RL13A, ARL1, STAT3, MDHM, RFC3, ECHA, SYIC, LAP2A, LPPRC, MATR3, MSH2, GPDM, VDAC2, KI67, BAG6, RL27A, RL5, RS9, STT3A, CAPZB, SYQ, RL29, AT5G3, TCPE, RL34, FAS, TCPG, EFTU, ACADV, TMEDA, NU153, RBP2, CPT1A, SERPH, RL14, TCPQ, TCPD, FXR1, RAB5C, RAB7A, HCFC1, ROA3, 6PGD, HNRPM, IMA1, HNRPF, MSH6, TXTP, ACLY, COPA, MOT1, SYRC, KAD2, P5CS, XPO2, TERA, NP1L1, DSRAD, ATPK, TMM33, TPIS, MYL6, IF4A1, RS20, S10AA, RAP1B, RL15, RL37A, HNRPK, RS8, RS16, 1433E, RS14, RS23, RS11, RUXE, RL7A, RS4X, RS6, H4, RAB1A, RAN, RL23, RS25, RS26, RL10A, RL11, RL8, PPIA, RS27A, RSMN, RACK1, ACTG, UBC9, TBA1B, TBB4B, GTF2I, TCPB, PRKDC, RL24, ARF5, RL19, SRSF3, MPCP, CLH1, HNRPU, SPTB2, EXOSX, RL18A, RL6, IF4G1, K1C17, PRDX1, RL18, C1QBP, KHDR1, DHX9, NCBP1, AHNK, NU160, SF3A3, ILF3, ACACA, PRDX4, CBX3, TIF1B, SPTN1, HNRPD, SAFB2, TTL12, CAPR1, ITPR1, RRP1B, GANAB, LBR, GOGB1, IMB1, NUMA1, SUZ12, U5S1, RRS1, PDIA6, PLEC, TEBP, NONO, PCBP1, PCBP2, DHC24, SF3B3, SF3A1, TRAM1, ELAV1, AAAT, RBBP7, H31T, PDS5A, TSR1, IF2GL, RRP12, NU188, HP1B3, EF1A3, PPR18, PRP8, C1TM, DHX30, CAND1, MISP, SPB1, PELP1, RDH10, CCAR2, TXND5, STT3B, BRX1, PO210, GEMI5, RT27, HS105, GCN1, NU205, AKAP1, AN32B, RBP56, DDX17, FUBP2, TNPO1, UBP7, UTP4, LRC59, PGAM5, FUBP3, MBOA7, MCCA, WRIP1, UHRF1, POP1, HCD2, ROAA, TM9S2, TCPH, ANM1, H2B1L, RNZ2, MEP50, MBB1A, ESYT1, H2AJ, GNL3, HDHD5, GTPB4, API5, RPF2, SFXN1, RDH14, ABCB6, DDX21, MDN1, DCA13, ATD3A, DDX18, MIC19, TEX10, TECR, MYOF, THYN1, HACD3, RRBP1, ABC3B, RLP24, ACINU, OGDHL, COR1C, PRP19, SSRG, TRI33, EIF3L, RUVB1, VDAC3, PDIP2, NOP58, SF3B6, RTCB, RL36, LAS1L, SRPRB, COPG1, MTCH2, CEPT1, ZNT1
Species: Homo sapiens
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Xie X, Wu Q, Zhang K, Liu Y, Zhang N, Chen Q, Wang L, Li W, Zhang J, Liu Y. O-GlcNAc modification regulates MTA1 transcriptional activity during breast cancer cell genotoxic adaptation. Biochimica et biophysica acta. General subjects 2021 1865(8) 34019948
Abstract:
Chromatin modifier metastasis-associated protein 1 (MTA1), closely associated with tumor angiogenesis in breast cancer, plays an important role in gene expression and cancer cell behavior. Recently, an association between O-GlcNAc transferase (OGT) and MTA1 was identified by mass spectroscopy. However, the potential relationship between MTA1 and O-GlcNAc modification has not yet explored.
O-GlcNAc proteins:
MED19, SBNO1, PGP, DPRX, SMHD1, RGPD3, ZSWM8, P121C, VGLL3, MCRI1, CK098, KIF2A, DX39A, PDLI1, BACH, MYO1C, MYO1F, AIP, SMAP, AP3B1, PSD11, PSD12, PSMD9, SPT5H, TAF4, DFFA, CLIC1, EIF3F, MATN2, LORF2, IPO5, SAP18, IF2B3, DNM1L, RTCA, AGRIN, PLOD2, HMGN4, IMA4, BCL9, PESC, NOP56, DDX3X, PODXL, CCN1, IMA3, NFIB, MA2B1, PDXK, ARI1A, TXND9, G3PT, CTRO, COPE, AP3D1, PDCD5, NHRF1, TPP1, TCRG1, TNPO2, PSA7, GIPC1, HAT1, HGS, MYPT1, HNRDL, XPO1, ZN609, SPTN2, SC16A, PLXB2, SR140, SET1A, LSM1, NPC1, ARC1B, ARPC2, ARPC3, TIF1A, PGRC2, PFD6, NKRF, LAMA5, ZN185, CASC3, OGT1, PMM2, HMGB3, PPM1G, FANCA, EIF3D, EIF3H, HDAC3, IPO8, STX7, NUP42, CLOCK, KDM6A, DHX15, MCES, FLRT2, PRP4, SERA, DC1L2, PSMD3, RFOX2, PAPS1, ZW10, SNUT1, SPIT2, TGFI1, M3K7, MCA3, HNRPR, PRPF3, TPD54, COCH, IF4G3, PPIH, ARK72, E41L2, TGON2, S26A4, FOXO3, DENR, XPOT, PLRG1, RGS10, ZN207, GET3, BUB3, ACTN4, BUD23, HTSF1, AP1G1, SYNC, KRT86, CPSF5, LANC1, AKAP8, CALU, RAD21, DHX16, MED14, SMCA5, JIP4, ZC3H1, TRAK2, AQR, GANP, KDM1A, GSDME, EVI5, SNX3, OGA, HNRPQ, CCNT1, PLOD3, AP180, PLIN3, MAFK, IMA7, UGDH, CTND1, SNX2, USO1, CCD22, PQBP1, DKC1, IF2P, EDF1, DNJA2, BRD4, PFD1, NBN, MCE1, WDR1, CPNE3, BRE1B, ZC11A, CLU, T22D2, PP6R2, CNOT3, CREST, ANR17, ZPR1, DUS11, PDCD6, TBCA, VATG1, H2AY, FLNB, NCOR1, SPAG7, SC22B, DEAF1, PR40A, VP26A, SAP30, MED24, KTNA1, PSIP1, SRS10, BAF, SF3B1, CSDE1, PRKRA, MED6, NPM3, LYPA1, U520, NU155, WDHD1, CRTAP, EIF3G, STAM2, CCNK, PRAF3, PIAS1, PIAS2, SPF27, DNJC8, SPF30, MPDZ, GLRX3, KRT36, RL1D1, CIAO1, SRP72, DDAH1, RECQ4, MTA2, MYO1D, TOX4, SC24D, PHF14, UBXN7, SUN1, PLPHP, ERLN2, PRP6, PCF11, NFAT5, HAUS5, AP2A2, SC31A, HEXI1, SEM3D, UBR5, AGFG2, SCAF4, ZFPL1, EPN2, ZRAB2, LC7L3, KAT7, MPZL1, FKBP9, CELF2, PAPS2, SMC2, LYPA2, IPO7, AGM1, CD2B2, BAG4, AHSA1, PSMG1, SC24A, SC24B, AGRL2, ACSL3, PCNT, CNOT4, CPSF4, EYA4, K2C75, SNP29, OXSR1, HS74L, PTBP3, AP2A1, STAU1, CAVN2, BAG2, M4K4, BPNT1, MBD3, CLPT1, ACL6A, CYB5, ADH1B, ADH1G, LDHA, DHE3, NB5R3, GSHR, SODC, PNPH, HPRT, AATM, EGFR, PGK1, KAD1, LALBA, FA10, TRYP, ASSY, A2MG, LDLR, HLAH, HBA, CO1A1, K1C14, K2C6A, LMNA, CASA1, CASA2, CASB, CASK, FINC, LACB, ALBU, ALBU, TFR1, GLCM, ALDOA, CYTB, ANXA1, GCR, PRIO, KITH, K2C6B, K2C1, G3P, CPNS1, HSPB1, RPN1, RPN2, GNAI2, AT1B1, ALDOB, A4, ARGI1, S10A8, HMGN1, ISG15, IF2A, HMGN2, ICAM1, RLA1, RLA2, JUN, LA, AP2A, ITB1, K1C18, K2C8, MYL1, GELS, PTMA, CDK1, ATPB, S10A6, IF4E, ENOA, PYGL, G6PI, DPOLB, NPM, TPM3, ITAV, LIPL, HEXA, ACBP, FABPL, LDHB, PDIA1, H10, CATD, ANXA2, CAN1, TBB5, TRY1, SAP, HEXB, PROF1, APT, K2C4, SYEP, CATB, HS90A, HNRPC, LAMB1, TPM2, FUMH, TSP1, SP1, ANXA6, DAF, MDR1, 4F2, HS90B, ASNS, ADH4, HEM3, RU2B, MET, RU17, ITA5, NFIC, VIME, RS17, K2C7, K1C13, ANXA5, K1C16, RSSA, CD63, SNRPA, GSTP1, RU1C, LEG1, HMGB1, CLCA, ROA1, RU2A, PARP1, UCHL1, LKHA4, ALDOC, CC175, ATX1L, ZN865, CT45A, HS71B, CALM3, RO60, H14, PTPRF, THIO, PPGB, KAP0, HEM4, ESTD, CH60, BIP, LAMC1, PPAL, HSP7C, GTR1, PYGB, LAMP1, TOP1, TOP2A, G6PD, C1TC, MPRI, ADHX, SRF, PABP1, PCNA, HARS1, IMDH2, TPR, KCRB, ANXA3, ACTN1, XRCC6, XRCC5, LAMP2, RINI, MYH6, ICAM2, AT1A3, EF2, K1C10, K2C5, PDIA4, TCTP, PLST, GYS1, T2FB, CD59, MIF, GLU2B, FPPS, TRI27, AK1A1, KPYM, ENPL, CCNB1, PO2F1, HNRPL, SYDC, ETS1, PLAK, ALDR, AMPN, PVR, ERF3A, EZRI, ATF2, FOSL1, FOSL2, MCP, NQO1, GNS, ZEP1, RS2, TFE2, DESP, RFA2, CD44, ITB4, CBR1, CREB1, BGAL, H15, H13, H12, AT2A2, STMN1, YBOX3, HSP76, HMGA1, JUNB, UBF1, JUND, ATF7, CAN2, PTN2, PYRG1, DDX5, PFKAL, LEG3, PRS6A, TCPA, PTN1, ARF4, RL7, VINC, SON, NELFE, RL17, PGAM1, RCC1, ATF1, XRCC1, CADH2, ML12A, NUCL, HXK1, RO52, SPEE, SL9A1, CSK22, RXRA, NFKB1, IF2B, ANXA7, TBP, BTF3, PSB1, MPRD, LMNB1, ICAL, CD11B, VATC1, CSRP1, FLNA, 5NTD, TAF1, CD9, TGM2, PIMT, FBRL, PUR2, PUR6, UBA1, NDKB, ROA2, RFX1, CBL, EST1, FBLN1, TCEA1, ITA6, SFPQ, PPIB, SYWC, RS3, NFYA, SAHH, COF1, IF4B, KTHY, CCND1, COT2, EF1B, PPAC, RPB1, CDK2, MCM3, NFYB, THTM, LAMA1, RS12, BRD2, TYY1, DNJB1, ATPA, PSA1, PSA3, PSA4, ITA3, MOES, DDX6, CTNA2, DNMT1, PAX6, U2AF2, RL13, HMGB2, PTBP1, SYTC, SYVC, EF1G, 1433T, ARNT, RL10, RFA1, APEX1, PYR1, CALR, MAP4, CALX, PSA5, DPOD1, MK01, RXRB, GRN, PTPRM, T2EA, CCN2, EPHA2, 3MG, KDM5A, TKT, RBMS1, PML, EF1D, ERP29, PRDX6, BLVRB, PRDX5, RL12, KCY, PEBP1, PDIA3, 2AAA, 2AAB, NMT1, HMOX2, PURA2, UFO, PUR8, CLIP1, SORCN, ILEU, RPB2, SDHA, METK2, CPSM, TIA1, ZEP2, DNJA1, AKT1, PUR9, HNRH3, HNRH1, 1433B, 1433S, STIP1, S10AB, L1CAM, PRDX2, OTX1, CDD, ELF1, RL9, KINH, CSTF2, DUT, TTK, MCM4, MCM5, MCM7, HSP74, GPC1, PROF2, CTNA1, CTNB1, PCGF2, SPB6, RADI, RFA3, RFC4, RFC2, RFC1, RL22, T2FA, SPRE, HXD13, CBS, K1C9, FBN2, GDE, MYH9, MYH10, COPB2, BASI, FUS, NU214, DEK, K22E, RL4, PP1G, GNL1, SRBP1, SRP14, NUP62, TAGL2, TALDO, RBMX, COIL, VKGC, GRP75, IF4A3, RS19, RL3, COIA1, OST48, AN32A, FEN1, CAPG, IL6RB, TXLNA, TCPZ, NNMT, PBX2, RL13A, STAT3, MDHC, MDHM, RFC5, IF2G, CETN2, BUD31, NAA10, CSK, GARS, SYIC, EIF1, KPCI, LAP2A, LAP2B, STAT1, MTREX, RS27, HELZ, RL35, CDN2A, PCP, ECE1, LIS1, MUC18, MATR3, MSH2, SSRA, RANG, NAMPT, PRS6B, NKX21, VDAC2, CBX5, UBP5, KI67, RAGP1, RECQ1, NOP2, CRK, CRKL, BAG6, NSF, NOTC1, MP2K3, RL27A, RL5, RL21, RL28, RS9, RS5, RS10, MAP1B, GNPI1, YAP1, UTRN, IQGA1, STT3A, CAPZB, IF1AX, SYQ, RL29, LIMS1, PPCE, RFX2, RFX5, SOX9, COPD, GSH0, PSMD8, PRC2A, TCPE, NEST, HSP13, KC1A, PIPNB, AGRE5, DPOD2, PAXI, RL34, NASP, FAS, CDK8, TCPG, EFTU, SRP09, PCY1A, SYAC, SYCC, SYSC, PRI2, CTCF, PSB2, MCM2, TSP3, YLPM1, TMEDA, RBM25, NUMB, HINT1, NU153, RBP2, GSK3A, GSK3B, TAF6, SPS1, GUAA, DNLI3, MRE11, CRIP1, GDIB, EMD, SERPH, F10A1, VASP, DYN2, CDK9, BCAM, PPT1, RL14, TCPQ, TCPD, ANX11, PAPOA, FXR1, FXR2, RAB5C, RAB7A, DUS3, SMCA4, GALK1, SSRD, HCFC1, UBP11, NR0B1, HDGF, ROA3, 6PGD, HNRPM, IMA1, IMA5, GDIR1, AGFG1, HNRPF, TF2AA, MSH6, ZN131, ZN143, RBM5, SPSY, EFNB2, CAZA1, CRIP2, NUP98, PPP5, ACLY, MAP11, COPB, COPA, CATC, AP2S1, PTTG, SMTN, IST1, SC24C, TCP4, ICLN, SYRC, ATX1, ATN1, SYYC, UBP14, HSP72, BCAT1, AT1B3, RD23B, PRRX1, HMCS2, SNAA, IF5, DRG2, XPO2, TERA, MFAP1, AF10, AF17, NP1L1, ADK, DSRAD, LAMB2, FOXG1, TPD52, SEC13, NH2L1, PSA, HNRH2, SUMO3, EIF3B, GFRA1, SYMC, IF6, CTBP2, TMM33, PCBP3, PCBP4, CORO7, NU107, MTPN, H2B1D, DEF3, ARPC4, SC61G, TPIS, EIF3E, SC61B, PP4C, MYL6, ACTB, IF4A1, RS20, PRPS1, PSA6, S10AA, CDC42, DEST, GMFB, RAB8A, SPCS3, RAB2A, RAB5B, RAB10, UBC12, UBE2N, ARP3, ARP2, ACTZ, CSN2, ABCE1, RS3A, RL26, PSME3, RL15, RL27, RL37A, RHOA, S61A1, LYSC, COPZ1, SUMO2, DCAF7, WDR5, NTF2, HNRPK, 1433G, RS7, PP1A, PP1B, PRS4, PRS8, RS8, RS15A, RS16, UB2G1, 1433E, RS14, RS23, RS18, RS29, RS13, RS11, RUXE, RUXF, RUXG, LSM3, LSM6, SMD1, SMD2, SMD3, TYB4, PRS10, RL7A, RB11A, ERF1, CNBP, RS4X, RL23A, RS6, H4, H2B1C, RAB1A, RAN, RL23, RAP1A, UB2D2, RS24, RS25, RS26, RS28, RS30, GBB1, RBX1, GBB2, RL30, RL31, RL10A, RL32, RL11, RL8, PPIA, FKB1A, RS27A, GRB2, TRA2B, RAC1, AP2B1, 1433Z, RSMN, SUMO1, DYL1, RL38, GBG5, RS21, IF5A1, RACK1, ACTG, SPT4H, UBC9, TYB10, PP2AA, YBOX1, CSK2B, TPM4, UB2L3, EF1A1, ACTS, TBA4A, TBB4B, CSK21, PA1B2, HBB, HBA, F193A, IGBP1, RBM6, IF4G2, RPP30, GTF2I, CNTP1, PHC1, TCPB, KRT85, RAE1L, PRKDC, MT1X, NUCB2, DCD, SSBP2, SARNP, RL24, CBX1, ARF1, ERH, RL19, SRSF3, FOXK1, DAB2, FBLN2, PGBM, RHG04, EFNB1, RBM10, RBM3, 2ABB, PIPNA, MPCP, VIGLN, TF2B, CDK5, CLH1, HSF1, FKBP3, DHSO, HNRPU, U2AF1, SPTB2, TIAR, SET, SRSF2, FOXK2, RUNX1, FABP5, CAP1, NC2B, TFAP4, EXOSX, OTUD4, PFKAP, EWS, TAGL, AK17A, MEF2A, KIF23, NRG1, DSG1, SP4, SP3, RL18A, MP2K1, FKBP4, PLOD1, NUCB1, RL6, TOP2B, DYST, CREM, KMT2A, LMNB2, TF65, UBXN1, GLGB, IF4G1, K1C17, NOTC2, TLE2, TLE4, LGUL, AK1C1, 1433F, CSTF1, PTN12, FAK1, SRS11, EF1A2, CALD1, PTN11, GFPT1, EXOS9, PSME1, SUH, GABPA, GABP1, RING1, PAX8, FMR1, PRDX1, RL18, CKAP4, ZO1, KHDR1, BAX, KLC1, LRP1, SRSF1, SRSF4, DHX9, GOGA2, LG3BP, CD47, PPID, SSRP1, NSUN2, RBBP4, EP300, AHNK, APOBR, HSP7E, MGAT2, GALT2, AP1B1, CPSF1, BST2, WASC5, NU160, SCAP, SRBP2, TBL3, ASPH, TROAP, TROP, SNF5, BPTF, CDC20, FSTL1, GRSF1, SFSWA, CHD3, SF3A3, TP53B, AIMP1, ILF2, ILF3, LMAN2, ELAV2, FOXC1, PP1R8, CSTF3, FLII, COASY, ACACA, CSN1, CAF1B, RED, MTAP, TADBP, ROA0, AIMP2, FADD, PRDX4, PAK2, CBX3, STK3, PSMD2, GPS2, SRSF9, SRSF5, SRSF6, TIF1B, G3BP1, CLN3, PTK7, PABP4, IFIT5, MTA1, EIF3I, 2A5G, CTBP1, UB2V1, DC1I2, ILK, MSLN, PPIG, TCOF, SF3B2, HAP28, ADAM9, FKBP5, SMAD4, PICAL, SQSTM, TBB3, ASAH1, PRP4B, PIN1, RIPK1, HDAC1, KCC2G, DCTN2, SNW1, STIM1, TRA2A, PWP1, CUL1, CUL2, CUL3, CUL4B, ASPP2, DYR1A, TPBG, FHL1, PM2PB, MOGS, CD166, NACA, THOC5, SPTN1, PKP1, DX39B, EXOS2, TBB2A, KLF5, BYST, IDI1, RUNX2, PEBB, NFYC, CDK13, CKAP5, CIRBP, COTL1, HNRPD, SCRB2, NID2, DAG1, VEZF1, DSG2, SAFB2, EIF3A, UBP2L, SCRIB, MLEC, TTL12, FHL2, DCTN1, DYHC1, EI2BA, IF4A2, SRC8, TRI25, FLNC, FA50A, MYPC2, GAMT, CAPR1, RBM39, KT33B, HLTF, PDIA5, DHX8, MCM6, ITPR3, MDC1, EPN4, SMC1A, RRP1B, NCOA6, GSE1, PSF1, UBP10, GANAB, 2A5D, LBR, GOGB1, MEF2D, CHD4, LASP1, NC2A, ZN638, IMB1, NOLC1, NUMA1, GAPD1, K2C72, NAA25, FL2D, SEPT2, SART3, U5S1, S39AE, SETB1, IF4H, BRD3, EEA1, PDIA6, PA1B3, PLEC, NOMO1, IPYR, TEBP, NECT1, NONO, PTPA, RBBP5, RCN1, K1H1, PCBP1, PCBP2, ELOC, SF3B3, DLGP5, RSU1, CNN3, SAFB1, SF3B4, SF3A2, RBMS2, SC23A, SC23B, SF3A1, SSXT, TAF5, TEAD4, TGIF1, NCOA2, NHRF2, TRAM1, SF01, TRIPB, PCH2, MED1, HMGN3, JHD2C, TRIP6, MARE1, ELAV1, ELF2, TAB1, AAAT, MLF2, TOM34, TBCE, UB2V2, STXB2, NEDD8, USF2, VAS1, VPS72, ZFHX3, ZYX, SEPT7, ADRM1, CCDC6, UAP1, AINX, PSMD5, CSRP2, DDB1, CDC37, DPYL2, RBBP7, TAF9, SRSF7, CPSF6, SMN, NRF1, FSCN1, IF16, GLO2, H2B2E, CGT, TRXR1, LY6K, HNRL2, Q1RMN8, PDS5A, QSER1, AAK1, WASC4, PRSR3, TSR1, QRIC1, SMU1, P3H1, MA7D1, CHD9, LSM12, GELS, A1AG, TBB8, GLYR1, FIL1L, AMOT, FA98B, EPC2, RHG29, FND3B, CRTC2, DHB12, GLE1, SNUT2, LACB2, P5CR3, PKHG4, H90B2, R39L5, ZN326, FILA2, BCORL, FIGN, K2026, AG10B, CC038, TGO1, PRC2B, RRP12, TOIP1, PCID2, SPXN3, SMAG2, BRM1L, TM201, NU188, HP1B3, CE170, ZN362, EYS, FKB15, DDX59, ZC3HD, LRIF1, UBR4, UBAP2, KPRP, XP32, RBM26, NT5D1, AHDC1, RPRD2, PR38B, BRE1A, ZN318, BROX, TASO2, ECM29, MBNL2, ZMYM4, K2C79, ARID2, USF3, C19L1, VIR, K1C40, HEAT6, BICRL, ATLA3, CARL2, CPIN1, TWF2, P4R3A, NIPBL, PDE12, LIN54, TET2, TM214, ARMC6, CAVN1, ERAP2, CDC73, TAF2, MED27, EDC4, PRP8, SCYL2, TTC27, GOLM2, NFRKB, MZT2A, CTR9, NCEH1, TRI65, MDEAS, BRAT1, ZC3HE, LARP1, BRWD3, SMRD3, FIP1, CRTC1, SAS6, CSPG4, CWC27, WDR82, ATL4, MCAF1, PACS1, MPRIP, GGYF2, CD109, CERS6, QSOX2, SRCAP, RIPR1, YJ005, UBN2, FGD6, PAXI1, RAPH1, IKIP, H32, LARP4, MED25, TBA1A, H2AV, HAKAI, ASXL2, CCD80, BT2A1, SPT6H, SND1, DDX46, BZW1, RSRC2, CYFP1, KDM3B, OLIG3, PHF5A, ZCCHV, KANL1, NUP54, POGZ, MYH14, NUFP2, CLAP1, PLGT3, HEAT3, HDGR2, EMSY, RAI1, I2BP2, RBBP6, SH3R1, HUWE1, YTHD3, KHDC4, TMED4, CENPV, T179B, Q862S4, KAISO, PATL1, MYPN, PEG10, NOP9, PABP2, MTA70, LDB1, PB1, LYRIC, TLK2, ZN598, KTN1, BCL9L, DTX2, LUZP1, THOC4, GP180, PACS2, CAND1, TSYL5, THOC6, MET16, NIPA, CARM1, PRSR1, MEX3D, DDX42, DAAF5, P66A, HORN, RB6I2, RIOX2, RBM45, PLD3, NUDC3, AHNK2, FOXP4, TEX2, Z3H7A, PHF6, CHERP, ANKH1, SUGP1, CCAR1, SLF2, PHC2, BAP18, RB12B, RAVR1, SPB1, IHO1, SRRM1, SMAP1, P20D2, KATL2, PHAR4, TRM2A, DCP1B, NALCN, MAML2, PELP1, ABI1, PORIM, CCAR2, NUP93, ZN687, CMTR1, LRC47, K2C78, ABD12, PALS1, STAG2, FNBP4, TAB3, DPP10, CPSF7, ARFG1, EMC1, PAF1, UBR7, ENAH, KRI1, PR38A, F184A, KDM1B, NHLC2, GT251, PGLT1, DHB11, TXND5, PAIRB, LEMD2, FA98A, RN214, LS14A, MISSL, TNR6A, PHC3, ABCF1, NGDN, SP20H, VP37A, KMT2C, ZMIZ2, ARI1B, NUP43, NUP37, NUP35, STXB6, KNL1, PLBL2, RLA0L, THOC2, WDR36, SMRC2, NPL4, PUM2, RDH11, STT3B, SPP2A, HM13, ALMS1, C99L2, MK15, BRX1, RBCC1, DOT1L, PO210, GEMI5, PARD3, IPO4, ZN384, K2C79, NOC3L, SCRB1, SMAP2, CCD12, TM263, NU133, PDC6I, GATD1, THEM6, CTF18, NUDC2, SCFD1, HNRLL, UBCP1, PCNP, LMO7, ATX2L, PHIP, PRP31, NELFB, PALLD, SREK1, PSPC1, P66B, DYH7, DNJC9, BBX, CTBL1, KAT6B, ELYS, ZCH14, DDX1, H1X, PSMF1, SMG7, RTF1, NICA, TM131, ELMO1, BAP1, PHF3, MAML1, NDRG1, HS105, CNOT9, ZN592, LAR4B, GCN1, PRP16, NU205, AN32B, NECT2, PRCC, TFG, TAF4B, HDAC2, STAM1, REQU, CBP, KAT6A, SYMPK, RBP56, GGH, DDX17, RAD50, CELF1, OSTF1, UFD1, RENT1, CSN5, GPKOW, SMRC1, SMRD2, FUBP2, TNPO1, NEP1, GLMN, USP9X, UBP7, CUL5, STMN2, LPP, MED12, H2A1C, BHMT1, SMCE1, MYDGF, RL36L, PSMG2, NCLN, UTP4, PP14A, MGN2, FERM2, FUBP1, LRC59, FKB10, PF21A, PPWD1, INT12, FWCH2, KLH29, CC124, AP2M1, REPS1, SNR40, MBD6, RMXL1, SEH1, TCAL4, THA11, GNA1, T3HPD, DAZP1, SAAL1, SGF29, RBM33, TMA16, ZN503, DYL2, CPNE2, THOC1, OTUB1, HMCES, MED8, DUS3L, DCNL1, SMRD1, DDX27, GWL, LIN37, ZCCHL, P121A, PDLI5, ERO1A, C2AIL, MED30, INT4, FUBP3, SPF45, ABHEB, PAWR, THOC3, KIRR1, WNK4, COG3, TMX3, CHAP1, LCOR, PDLI2, ANCHR, ZC3HA, TOPK, SCYL1, EHMT2, ZFR, EP400, PRRC1, FXL18, CCD43, NOL4L, DACH2, NSUN5, RPR1A, IPO9, RBM14, QKI, ALS2, PP1RA, S38A2, VPS35, PURB, PHF12, XPO6, NACC1, CIC, MED15, MCCA, NUDC1, ERBIN, WRIP1, RANB9, MYADM, SRPK1, STRBP, TM209, IWS1, SIN3A, RSF1, RBM15, RUFY1, MINT, PNKP, MMS19, UHRF1, NIBA2, HTF4, TBCB, CNN2, CDC5L, PSMD1, PFD5, RING2, PARK7, EYA3, VAT1, LGMN, NUP88, POP1, MNT, SCAFB, EIF3C, TTC1, DNJC7, KIF2C, RGPD5, ATX2, ROAA, NP1L4, SNAG, TS101, CPNE1, TCPH, EBP2, ARI3A, ANM1, H2A1J, PKP2, SEM3C, VRK1, RBM4B, DDX50,