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Li G, Meng F, Zhong X, Yu K, Zhang N, Zhang K, Huang H, Li W, Zhang J, Wang W, Ren Y, Liu Y. DNA probe pulldown screening uncovers O-GlcNAcylation modulation of transcription factor DNA interactions. Scientific reports 2025 15(1) 40596516
Abstract:
O-linked β-N-acetylglucosamine (O-GlcNAc), a critical post-translational modification predominantly found in the nucleus, plays a substantial role in regulating gene expression by modulating transcription factors (TFs) activity. However, quantitative analysis investigating the influence of O-GlcNAcylation on protein-DNA interactions at a proteome scale remains undone. Herein, a pulldown screening approach using a consensus TF response element (catTFRE) was employed to unravel the impact of fluctuating levels of O-GlcNAcylation on the DNA binding efficiency of endogenous TFs/co-factors. Utilizing quantitative proteomics, we identified a substantial enhancement in the binding capacity of 241 nuclear proteins (NPs) to DNA sequences due to elevated levels of O-GlcNAcylation, whereas a decrease in DNA binding was observed for 2 NPs concurrently. Intriguingly, the O-GlcNAcylation elevation significantly enhanced the binding of 146 TFs/co-factors to specific DNA sequences. We further established that the O-GlcNAcylation of several Forkhead family TFs, including FOXA1 and FOXC1, notably enhances their binding to specific DNA sequences in living cells. Our research presents an efficacious approach to assessing the impact of O-GlcNAcylation on the interactions between proteins and DNA. This significantly enhances our understanding of the role O-GlcNAcylation plays in the regulation of transcription.
O-GlcNAc proteins:
RBM47, ESYT2, HACL2, VWA8, CNOT1, PGP, SMHD1, U119B, MET15, MYO1G, ADAS, PDLI1, BACH, HAX1, SPT5H, TAF4, CLIC1, BMAL1, SAP18, DNM1L, PLOD2, PSDE, BIN1, NOP56, DDX3X, TRI38, NFIB, PPP6, CDKA1, ABLM1, ADA10, CHK1, PSA7, TX1B3, TIM23, MYPT1, S27A2, XPO1, SYN3, ZN609, SR140, SET1A, ZN185, ATX7, OGT1, EIF3D, EIF3H, TNC18, MOT4, P4HA2, NUP42, ARPC5, CLOCK, MAFG, KDM6A, DHX15, RRP8, PRP4, SERA, PHF1, DC1L2, PSMD3, SNUT1, MCA3, HNRPR, PRPF3, TPD54, EMC8, MGA, KLF4, TIM44, PLRG1, ZN207, BUB3, ACTN4, BUD23, ENSA, SPOP, LANC1, AKAP8, CALU, ORC5, ZNRD2, MED14, NPA1P, ZC3H1, AQR, GANP, KDM1A, ACSL4, VINEX, CCNT1, PLOD3, TSN3, PLIN3, MAFK, UGDH, PQBP1, TBL1X, NBN, MITF, CPNE3, ZC11A, ANR17, H2AY, FLNB, NCOR1, SPAG7, RM33, SC22B, PR40A, MPPB, KTNA1, PSIP1, ERLN1, NDUS3, SRS10, SF3B1, CSDE1, U520, NU155, CRTAP, S2512, EIF3G, SOAT2, CCNK, SPF27, SPF30, RL1D1, CLPX, SYUG, DFFB, SRP72, RECQ5, ATG12, TOX4, K0754, SRBS2, SUN1, ERLN2, PRP6, SC31A, UBR5, AGFG2, SCAF4, DUS14, SPN1, NDUB8, RTN3, EPN2, LC7L3, MBD4, VAPB, 6PGL, ZBT7A, IPO7, SLU7, CD2B2, PSMG1, SC24B, ZBT11, SNP29, OXSR1, HS74L, LSM8, AP2A1, WIZ, BAG2, AIFM1, BPNT1, SNAI1, DDAH2, TXD12, MBD3, TOM40, NAPSA, ACL6A, CCNE2, LDHA, NB5R3, PGK1, A1AT, FOS, RASH, LDLR, LMNA, ALBU, TFR1, TRFL, FRIH, NU3M, MIS, VTNC, CATA, GBA1, FUCO, ALDOA, CYTB, GCR, SODM, KITH, K2C1, G3P, HLAA, CPNS1, HSPB1, TYSY, RPN1, RPN2, GNAI2, AT1A1, AT1B1, A4, APOD, ALDH2, PERM, PPBT, IF2A, RLA2, RLA0, JUN, ITB1, K1C18, K2C8, GELS, ATPB, IF4E, ENOA, NPM, ITAV, GDN, ACBP, LDHB, CATD, ANXA2, TBB5, SAP, CATL1, SYEP, CATB, HNRPC, FUMH, TSP1, SP1, ANXA6, HS90B, SRPRA, ODPA, RU2B, RU17, ITA5, NFIC, VIME, K2C7, GNAI3, ANXA5, SNRPA, HMGB1, DLDH, ROA1, RU2A, LKHA4, CTF8, MNS60, RO60, MYBB, LYAG, HLAC, KAP0, CH60, CLUS, BIP, HSP7C, GTR1, ODPB, LAMP1, ACADM, TOP1, TOP2A, G6PD, MPRI, PRPS2, PABP1, PCNA, ADT3, FA5, IMDH2, TPR, KCRU, SKI, ACTN1, XRCC6, NR2C1, LAMP2, EF2, K1C10, K1C13, PDIA4, PLST, T2FB, MIF, FPPS, KPYM, ENPL, MYL6B, RSMB, PO2F1, PLAK, ALDR, RAC2, ERF3A, EZRI, ATF2, FOSL2, MCP, NDKA, GNS, RS2, DESP, MUC1, CREB1, MGMT, CD36, STMN1, YBOX3, ZFX, JUNB, UBF1, JUND, ATF7, PTN2, DDX5, ARF4, PTPRA, SON, NELFE, RCC1, ATF1, ATF6A, XRCC1, LFA3, NUCL, ELK1, TFE3, CSK22, NFKB1, TYPH, TBP, RAB6A, PSB1, LMNB1, HXA5, CSRP1, FLNA, VDAC1, SDHB, CD9, PIMT, FBRL, UBA1, NDKB, ROA2, RFX1, KAPCB, QCR2, ITA6, SFPQ, PPIB, MAOM, SP100, NFYA, SAHH, COF1, IF4B, GATA2, THIL, MCM3, RS12, BRD2, TYY1, ATPA, PSA1, PSA4, ITA3, DDX6, PTBP1, SYVC, KAD4, 1433T, VATL, ARNT, RFA1, APEX1, PYR1, CALR, MAP4, CALX, PSA5, PSB4, PEX2, NDUS1, TEAD1, GRN, T2EB, 3MG, TKT, SPB3, ECHM, KCY, PEBP1, PDIA3, 2AAA, CDC27, HMOX2, PURA2, AMRP, RPB2, SDHA, METK2, HXA11, HXC9, CPSM, 3HIDH, PUR9, HNRH3, HNRH1, 1433S, STIP1, L1CAM, PRDX2, P5CR1, ELF1, CGL, RL9, CSTF2, MCM4, MCM7, HSP74, TRY3, GPC1, CTNA1, BMI1, PCGF2, PHB1, RADI, RFC4, RFC1, MYH9, MYH10, COPB2, SOAT1, ADDB, BASI, FUS, NU214, DEK, GLRX1, K22E, MP2K2, VATE1, RL4, PGM1, SPB5, ODO2, SRP14, NUP62, TAGL2, SYUA, ETFB, RBMX, VATA, HSPA9, IF4A3, TXLNA, TCPZ, PBX2, MLH1, STAT3, MDHM, ECHA, IF2G, S19A1, ELK3, ACTY, LAP2A, LAP2B, LPPRC, THIM, ECE1, MUC18, MATR3, GPDM, SSRA, RANG, PRS6B, VDAC2, MP2K4, KI67, RAGP1, RECQ1, NOP2, CRKL, NEDD4, IQGA1, STT3A, CAPZB, RL29, UCRI, RFX5, COPD, PSMD8, PRC2A, HSP13, AGRE5, DPOD2, NR2C2, RPIA, LMAN1, FAS, CDK8, RM19, AL7A1, CENPF, SYAC, SYCC, SYSC, MCM2, ACADV, YLPM1, RBM25, CLK1, NU153, RBP2, GSK3A, GSK3B, TAF6, GUAA, DNLI3, MRE11, IDH3A, EMD, CPT1A, SERPH, F10A1, MXI1, MAP2, PPT1, TCPQ, TCPD, FXR1, FXR2, RAB5C, RAB7A, SMCA4, IDH3G, GALK1, MECP2, DHB4, SPHM, HDGF, ROA3, 6PGD, IMA1, GDS1, MP2K6, AGFG1, HNRPF, TF2AA, MSH6, CAZA1, CRIP2, NUP98, PLK1, ACLY, SUCA, COPA, PTTG, SC24C, TCP4, ICLN, HIRA, SYYC, UBP14, HSP72, AT1B3, RD23A, RD23B, KAD2, SNAA, IF5, RAD, TERA, DSRAD, TPD52, SEC13, PSA, HNRH2, EIF3B, SYMC, DPOE2, ATP5I, IF6, NU107, SESN2, FOXL2, EPIPL, TPIS, SC61B, ACTB, IF4A1, PRPS1, SRP54, RAB2A, ARP2, ACTZ, CSN2, ARF3, CH10, STXB1, B2MG, WDR5, HNRPK, RS16, SMD2, ERF1, RS4X, ACTA, RL23A, RS6, H4, GBB1, RL32, RL8, PPIA, RS27A, RL40, AP2B1, 1433Z, IF5A1, RACK1, ACTG, YBOX1, TPM4, EF1A1, ACTS, TBA1B, TBA4A, TBB4B, CSK21, HBB, HBA, CXAR, NOP14, RBM6, PITX1, IF4G2, GTF2I, GSTO1, PO6F2, PRKDC, BASP1, RT22, RT05, RT11, RT34, RT09, HMGN5, SARNP, LACTB, TXN4A, RL19, FOXK1, RBM10, EM55, TFAM, VIGLN, CDK5, CLH1, HNRPU, SPTB2, TIAR, SET, FOXK2, RUNX1, CAP1, TFAP4, EXOSX, XPC, EWS, MEF2A, SP2, ODO1, KIF23, DSG1, SP3, RL18A, FKBP4, PLOD1, TOP2B, M2OM, DYST, CREM, KMT2A, CENPC, LMNB2, UPAR, KIME, TAP1, IF4G1, K1C17, SSBP, SRS11, EF1A2, CALD1, KDSR, SUH, GABPA, GABP1, PRDX1, RL18, C1QBP, CKAP4, ZO1, KHDR1, SRSF4, PP2BA, DHX9, LG3BP, MFGM, CD47, SSRP1, SLFN5, RBBP4, NCBP1, EP300, AHNK, MGAT2, GALT2, BST2, MPPA, SIA4C, NU160, TBL3, TF3C1, ASPH, SNF5, BPTF, NFIA, CHD3, TP53B, IFRD2, AIMP1, ILF3, LMAN2, FOXC1, PP1R8, ECH1, PRDM2, CSN1, KLF10, NR1H3, TADBP, ROA0, PAK2, PSMD2, DDX10, GPS2, SRSF5, SRSF6, TIF1B, G3BP1, STF1, NMI, KCAB2, SKP2, PABP4, GRB10, MTA1, DC1I2, ORC2, SNTB2, PPIG, TCOF, ADAM9, TUSC3, SMAD4, PICAL, TBB3, ASAH1, HDAC1, SNW1, STIM1, CUL4A, CUL4B, FHL1, RUNX3, THOC5, SPTN1, NOG2, KLF5, RUNX2, PEBB, NFYC, CKAP5, SCRB2, DAG1, VEZF1, DSG2, KEAP1, MORC3, SAFB2, EIF3A, UBP2L, MLEC, ZMYM3, DCTN1, DYHC1, EI2BA, ELOA1, MAP7, SRC8, FLNC, CAPR1, NF2L1, HABP2, DHX8, ITPR3, PLSI, TRIPC, PUM1, MDC1, EPN4, SMC1A, RRP1B, RRP5, BMS1, GANAB, MBTP1, 2A5D, LBR, CBX2, KIF22, MEF2D, CHD4, LASP1, ARI5B, NFIX, ZN638, NOLC1, NUMA1, GAPD1, SPCS2, EMC2, MO4L2, ABRX2, CND1, SUZ12, U5S1, SYK, RRS1, IF4H, BRD3, WDR43, ACOX1, EBP, PLEC, PON2, NONO, PTPA, PWP2, RNPS1, RCN1, PCBP1, MTFR1, SF3B3, PUM3, DLGP5, KS6A1, SF3A2, SC23A, SF3A1, SURF1, MED22, TAF5, TAF7, MARE2, TSN, SF01, PCH2, MED1, JHD2C, TRIP6, MARE1, ELF2, NAB2, AAAT, USF2, VAS1, RB11B, EZH2, ZYX, SEPT7, ADRM1, TAF12, DDB1, SNPC1, CDC37, DPYL2, SYPL1, TAF9, SRSF7, DREB, NRF1, PCKGM, DDR2, LY6K, HNRL2, PDS5A, ZN800, P3H1, RA1L2, MA7D1, TBRG1, LARP7, ACSF3, TPRN, EPC2, DHB12, UBP39, SETMR, YIF1B, ZN326, NOM1, SH319, RRP12, TOIP1, TDIF2, NU188, HP1B3, ZN362, SYRM, ZC3HD, LRIF1, RM02, UBAP2, RBM26, RIF1, MRCKA, RPRD2, BRE1A, TASO2, KHDR2, AR6P4, ARID2, GL8D1, NR3L1, MSL1, ABLM2, GLYAT, TWF2, K0930, SN12L, SDE2, LIN54, ALKB2, PPR18, CAVN1, CDC73, PRP8, SCYL2, GOLM2, RHGBA, NFRKB, RSBNL, KCD18, NCEH1, MDEAS, ZC3HE, SAMD1, C1TM, FIP1, CSPG4, PLGT2, SBSN, CF089, LCN15, CREL2, LRIG3, BCOR, MPRIP, CD109, SRCAP, UBN2, IKIP, CENPU, LARP4, HAKAI, NOL8, ASXL2, SND1, DDX46, TM10C, EIF3M, MCM10, NDUF7, CXXC5, APTX, ZCCHV, KANL1, NUP54, POGZ, NUFP2, SZRD1, CLAP1, HEAT3, HDGR2, BRAP, EMSY, I2BP2, RBBP6, YTHD3, TAF8, TMED4, CENPV, PHLB2, MYPN, ACOT1, PABP2, LDB1, PB1, YRDC, LYRIC, KTN1, LUZP1, CAND1, THOC6, DDX42, SPAS2, CACL1, P66A, RIOX2, CRLF3, VRK3, NAV2, CCD50, MISP, CTL2, MGAP, SFPA2, SFTA1, PHF6, CHERP, SUGP1, SUGP2, CCAR1, BAP18, RM41, KLHL7, MILK2, DHX37, YAF2, SRRM1, ZZZ3, K2013, ELMD2, CORA1, PELP1, TTC5, MIER1, NDUF2, NUP93, ZN687, CMTR1, LRC47, INT1, ABD12, CARME, FBXL6, SRBD1, NKAP, RP25L, ATPF2, RM50, CPSF7, ARFG2, EFNMT, ARFG1, PAF1, ENAH, DAND5, KRI1, MLKL, XXLT1, GOLM1, SUMF2, SUMF1, PGLT1, DHB11, TXND5, SERB1, CHSTE, LS14A, FSAF1, RB15B, PHC3, DIK2A, POPD1, SRFB1, SP20H, VP37A, PCAT1, BD1L1, NUP35, KNL1, PLBL2, DDX55, THOC2, WDR36, KI18A, GABP2, NPL4, STT3B, SPP2B, BRX1, STK35, NEK7, GPX8, PO210, GEMI5, FBF1, ZN384, WIPF2, SETD7, SMAP2, TM263, NU133, PDC6I, CO040, NUDC2, TWST2, SCFD1, PCNP, SPRY4, TRUB1, LMO7, CTL1, CKAP2, ATX2L, PHIP, RT4I1, NELFB, PALLD, PYDC1, PSPC1, P66B, BBX, CTBL1, ELYS, S39A7, H1X, RT27, NR4A3, PHF3, NDRG1, HS105, ZN592, LAR4B, NU205, PRCC, TFG, TAF4B, RREB1, HDAC2, SC65, SYMPK, GGH, DDX17, RAD50, CELF1, GSLG1, GPKOW, SMRC1, SMRD2, RAB8B, FUBP2, TNPO1, ARHG2, GLMN, KAT5, UBP7, POP5, LSM10, LMBL2, NCLN, FUBP1, SOSB2, PBIP1, SH3K1, PF21A, PINX1, P5CR2, INT12, FWCH2, FAF2, RMD1, MSI2H, RRFM, MTEF3, THA11, SGF29, DTBP1, PTCD3, MMAB, MCRS1, CERS2, SMRD1, TOE1, DDX27, SYMM, P121A, PDLI5, ERO1A, INT4, FUBP3, SYNM, VSIG2, HPDL, PAWR, RBM41, RSRC1, THOC3, CHAP1, S35E1, CLP1L, Z512B, ZFR, EP400, CNO6L, PRRC1, ZN512, YTDC1, SPT33, ZN830, RPR1A, RBM14, NED4L, QKI, LENG8, TRNT1, PP1RA, DEPD7, ATAD5, TEFM, MED15, EFGM, PIGS, TM209, IWS1, SIN3A, RBM15, MINT, NIBA2, YMEL1, HTF4, CNN2, SEC62, CDC5L, PSMD1, RING2, PARK7, EYA3, LGMN, PLIN2, POP1, SCAFB, TEAD3, TTC1, PHB2, CSN8, HCD2, SMAD5, ROAA, S29A1, CPNE1, MYD88, PKP2, SH3G1, GDF15, ARP5L, ZFY21, CMS1, DPH2, TBA1C, BCL7B, MBB1A, COR1B, TIRR, FSP1, PELO, ERP44, TACO1, ESYT1, CCM2, CNPY3, DIDO1, NDC1, ZSA5A, PAXX, HNRL1, RPP25, RCAF1, TBB2B, TM109, NUP58, GNL3, RIOK2, TIM21, THIC, RBM4, MND1, NADAP, SSBP3, RFIP5, ORNT2, NAA15, SRRT, AP1M1, NUSAP, HDHD5, RTF2, PECR, PDIP3, CEP41, RM37, GTPB4, UACA, UCK2, TB182, LNP, CEP44, WDR33, CTDS1, MFF, RB33B, NAT10, SP130, BRD8, RGAP1, CSTFT, LMA2L, CK054, F234A, RBM38, TDIF1, ALX4, SIL1, APC1, TMX4, WDR13, TARA, EPC1, ADNP, IPYR2, RM46, PININ, TM245, FOXP1, CPVL, GHITM, NELFA, BORG4, UN45A, SMOC2, AMPB, ZHX3, EMRE, REEP4, TM231, SH24A, DOCK5, CH033, SDS3, ACAD9, CQ10B, GORS2, JUPI2, SFXN1, SPE39, UBP42, RPF1, RPA2, GBB4, BRF2, RISC, CYBP, TENS1, TAF9B, TM165, MRM3, NCOA5, TRI39, SDF2L, AT131, RM47, APMAP, ARFG3, VTA1, PLPL8, SYSM, OSGEP, DMAP1, UBN1, RTN4, KMT5A, INCE, PRD10, ANLN, PDLI7, DDX21, MBNL1, SIR7, AAAS, ABCBA, PNO1, MBIP1, RAD18, SLX9, BMP2K, PDS5B, ECHD1, MDN1, DECR2, STAU2, DDX28, T106B, SPS2L, TYDP1, SEP11, DJC11, HAUS2, RBM22, MED9, ARGL1, SLTM, RMND1, AKIP, OXSM, THG1, HPF1, NDUBB, OCAD1, MARH5, MIC19, SIR5, CWC25, TEX10, THUM1, ING3, CARF, NSE4A, AATF, DDX56, TE2IP, ABI2, HXC10, BCLF1, TMOD3, MTCH1, HPBP1, MAT2B, ITSN2, CHMP5, THYN1, VAPA, RCC2, SLAI2, BCCIP, FPRP, RRBP1, CPSF2, RBM27, KANL3, GRHPR, NXF1, DAXX, TRM7, VTI1B, ABCF2, SYWM, SEC63, NDRG3, SUN2, LIMA1, SEPT9, TRPS1, DPP2, BAP29, NB5R1, S30BP, MED13, KAD3, HACL1, S2513, DBNL, CDC23, GMEB2, RCOR1, ACINU, NUP50, ZHX1, CDV3, CNO11, INT6, MCTS1, ZBT21, YETS2, ZMYD8, TMCO1, EPDR1, MYO6, KMT2B, PRP19, G3BP2, PACN2, MAGD2, PPIE, DIM1, MINP1, MACF1, PHF8, LIMC1, XCT, SRRM2, PA2G4, SMC3, SCML2, ZN148, RUVB2, VDAC3, RT33, PLAK2, ICE1, DDX52, MAN1, TR150, WBP11, MED16, NOP58, ZN281, SYYM, COQ6, LSM2, PHOCN, ORN, RT23, HDGR3, STRAP, RTCB, NOC2L, CHTOP, VPP2, TLN1, TRRAP, CRBG1, HYOU1, LAS1L, SAM50, PRC2C, RBM7, INSL6, YTHD2, PAXB1, SP16H, UTP18, CD2AP, SRPRB, TSSC4, SNX5, NUBP2, HCFC2, LRRF2, NPTN, 5MP1, DC1L1, CABIN, COX11, SCC4, ZHX2, MORC2
Species: Homo sapiens
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Xiong Y, Lu Z, Shao Y, Meng P, Wang G, Zhou X, Yao J, Bao H, Lu H. Rapid and large-scale glycopeptide enrichment strategy based on chemical ligation. National science review 2024 11(11) 39534244
Abstract:
Protein glycosylation, the most universal post-translational modification, is thought to play a crucial role in regulating multiple essential cellular processes. However, the low abundance of glycoproteins and the heterogeneity of glycans complicate their comprehensive analysis. Here, we develop a rapid and large-scale glycopeptide enrichment strategy via bioorthogonal ligation and trypsin cleavage. The enrichment process is performed in one tube to minimize sample loss and time costs. This method combines convenience and practicality, identifying over 900 O-GlcNAc sites from a 500 μg sample. Surprisingly, it allows simultaneous identification of N-glycosites, O-GlcNAc sites, O-GalNAc sites and N-glycans via a two-step enzymatic release strategy. Combined with quantitative analysis, it reveals the distinct O-GlcNAcylation patterns in different compartments during oxidative stress. In summary, our study offers a convenient and robust tool for glycoproteome and glycome profiling, facilitating in-depth analysis to elucidate the biological functions of glycosylation.
O-GlcNAc proteins:
MEIOC, SBNO1, CNOT1, HMX3, RGPD3, P121C, TAF4, ARI1A, CDKA1, ABLM1, KMT2D, RGPD8, MYPT1, SC16A, SET1A, TIF1A, NUP42, SYNJ1, ZN207, AKAP8, ZC3H1, GANP, DIAP1, MAFK, BRD4, MITF, T22D2, ANR17, NCOR1, CCNK, PIAS1, TOX4, ST65G, PCF11, NFAT5, SC31A, SVIL, SC24A, SC24B, CNOT4, EGFR, LMNA, HSPB1, RPN1, RPN2, RLA2, RLA0, AP2A, K1C18, K2C8, NPM, VIME, K2C7, SNRPA, ATX1L, TPR, GLU2B, PO2F1, ZEP1, JUNB, JUND, ATF7, DDX5, SON, ATF1, CSRP1, SFPQ, BRD2, ARNT, MAP4, CALX, PSB4, TEAD1, HXA11, HXC9, ELF1, NU214, NUP62, CUX1, MATR3, SSRA, SSRB, UTRN, STT3A, NEST, LMAN1, CDK8, YLPM1, NU153, RBP2, TAF6, EMD, DYN2, SMCA4, AGFG1, HNRPF, NUP98, ATN1, RD23B, AF10, AF17, FOXA1, CTBP2, RS16, NOMO3, PHC1, RAE1L, SARNP, FOXK1, HNRPU, SPTB2, FOXK2, TFAP4, KMT2A, IF4G1, GABPA, ZO1, EP300, BPTF, NFIA, FOXC1, ROA0, G3BP1, PICAL, MOGS, RUNX3, KLF5, CDK13, VEZF1, UBP2L, ELOA1, CAPR1, PUM1, RRP1B, NCOA6, GSE1, MEF2D, LASP1, ARI5B, ZN638, NUMA1, DAZP2, KIF14, EBP, PLEC, PCM1, PUM3, RBMS2, TEAD4, SF01, MED1, JHD2C, ELAV1, ELF2, TAB1, ZFHX3, ZYX, CCDC6, TAF9, IF16, RFX7, QSER1, CHD9, CRTC2, BCORL, BRD10, NOMO2, TGO1, PRC2B, ZN362, LRIF1, RHG21, UBAP2, AHDC1, RPRD2, ZN318, ZMYM4, ARID2, BICRL, NIPBL, LIN54, Z280D, MFSD5, SCYL2, NFRKB, RSBNL, GLD2, MDEAS, ZC3HE, CRTC3, SAS6, MCAF1, BCOR, GGYF2, CO039, SRCAP, UBN2, TMTC3, RAPH1, UBP54, SPT6H, KDM3B, POGZ, NUFP2, EMSY, RAI1, I2BP2, RBBP6, SH3R1, YTHD3, BCL9L, TIKI1, PRSR1, DDX42, P66A, RB6I2, AHNK2, GID4, TEX2, MGAP, ANKH1, SUGP1, SPB1, ZN687, SYNPO, FNBP4, MACOI, ARFG1, S43A3, LS14A, TNR6A, PHC3, KMT2C, KNL1, CHD6, PO210, PARD3, LMO7, P66B, BBX, ELYS, ZCH14, SMG7, ZN592, PRCC, US6NL, RREB1, CBP, HXB13, GPKOW, FUBP2, LPP, S2546, TOM6, PF21A, INT12, SNR40, RBM33, P121A, PDLI5, FUBP3, CHAP1, ZC3HA, Z512B, ZFR, EP400, NOL4L, RBM14, QKI, LENG8, CIC, STRBP, SIN3A, MINT, HTF4, MNT, SCAFB, TEAD3, ATX2, ARI3A, RBM4B, CNPY3, WAC, DIDO1, RBM4, YTHD1, AMRA1, SP130, BRD8, I2BPL, ADNP, FOXP1, E41L1, PEAK1, ECT2, FBRS, PKHA5, RC3H2, TAF9B, ZBT20, NCOA5, TANC2, CHD8, UBN1, PDLI7, 4ET, FANCI, NSMA3, CARF, CDK12, CNOT2, CHD7, RBM27, KANL3, RERE, TCF20, DPP2, GT2D1, PUF60, SIX4, CDC23, TASOR, GMEB2, ZHX1, ZBT21, PRR12, YETS2, PRP19, SCAF8, TRI33, SRRM2, SCML2, ZN148, HPSE, ZN281, FBX7, ARIP4, KDM3A, PLXD1, CRBG1, HYOU1, PRC2C, YTHD2, SRPRB, NCOR2, S23IP
Species: Homo sapiens
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Liu J, Hao Y, Wang C, Jin Y, Yang Y, Gu J, Chen X. An Optimized Isotopic Photocleavable Tagging Strategy for Site-Specific and Quantitative Profiling of Protein O-GlcNAcylation in Colorectal Cancer Metastasis. ACS chemical biology 2022 17(3) 35254053
Abstract:
O-linked-β-N-acetylglucosamine (O-GlcNAc) glycosylation is a ubiquitous protein post-translational modification of the emerging importance in metazoans. Of the thousands of O-GlcNAcylated proteins identified, many carry multiple modification sites with varied stoichiometry. To better match the scale of O-GlcNAc sites and their dynamic nature, we herein report an optimized strategy, termed isotopic photocleavable tagging for O-GlcNAc profiling (isoPTOP), which enables quantitative and site-specific profiling of O-GlcNAcylation with excellent specificity and sensitivity. In HeLa cells, ∼1500 O-GlcNAcylation sites were identified with the optimized procedures, which led to quantification of ∼1000 O-GlcNAcylation sites with isoPTOP. Furthermore, we apply isoPTOP to probe the O-GlcNAcylation dynamics in a pair of colorectal cancer (CRC) cell lines, SW480 and SW620 cells, which represent primary carcinoma and metastatic cells, representatively. The stoichiometric differences of 625 O-GlcNAcylation sites are quantified. Of these quantified sites, many occur on important regulators involved in tumor progression and metastasis. Our results provide a valuable database for understanding the functional role of O-GlcNAc in CRC. IsoPTOP should be applicable for investigating O-GlcNAcylation dynamics in various pathophysiological processes.
O-GlcNAc proteins:
A0A0B4J203, A0A0C4DFX4, RBM47, E2F8, WDR27, SBNO1, CNOT1, P121B, P121C, H0YAE9, H0YHG0, H7C469, K7ELQ4, M0QZ24, PDLI1, HAX1, TAF4, BCL9, CAC1A, DDX3X, NFIB, PPP6, MA2B1, ARI1A, SOCS7, ABLM1, KMT2D, GBRD, RGRF2, TX1B3, HGS, MYPT1, SYN3, ZN609, TRI66, PDZD2, MAST4, SC16A, SET1A, CASC3, FOXP2, MOT4, P4HA2, ARPC5, CLOCK, MAFG, PER1, KDM6A, TET3, SI1L1, TGFI1, M3K7, MCA3, PRPF3, TPD54, SYNJ1, IF4G3, E41L2, WIPF1, FOXO3, TGM5, RNF13, SPY2, PLRG1, ZN207, AKAP8, CALU, ORC5, MYPT2, GANP, OGA, CCNT1, BUB1B, PLOD3, PLIN3, MOT2, MAFK, PQBP1, BRD4, TBL1X, PP1RB, NBN, MITF, SRGP2, N4BP1, ROCK2, PP6R2, CNOT3, ANR17, FLNB, NCOR1, SF3B1, REM1, CREG1, CRTAP, SYUG, CYTF, TOX4, TOX, SUN1, PCF11, AGFG2, UBE4B, CAC1H, SVIL, SC24A, SC24B, CNOT4, EYA4, ZMYM6, BAG3, LATS1, DDAH2, TXD12, ONEC2, CLPT1, ABL1, CRYAB, LMNA, TFR1, CATA, GBA1, FUCO, ALDOA, GCR, G3P, CPNS1, HSPB1, RLA2, RLA0, ITB1, K1C18, NPM, CATL1, CATB, MCR, BGLR, ITA5, NFIC, VIME, SNRPA, FGR, ATX1L, DERPC, ZN865, GLI2, MYBB, CLUS, PPAL, MPRI, PABP1, TPR, BMP3, SKIL, ENPL, PO2F1, PLAK, ATF2, ZEP1, RS2, TFE2, F261, ITB4, ZNF23, ZNF25, JUNB, ATF7, TPH1, DDX5, EGR1, SON, NELFE, ATF1, ATF6A, CADH2, ICAL, CSRP1, FLNA, RFX1, CBL, SFPQ, COF1, IF4B, GATA2, APC, DDX6, ARNT, MAP4, LYOX, HXD9, MZF1, CLIP1, 5HT1F, HXA11, ZEP2, ELF1, CTNB1, FBN1, ADDA, BASI, NU214, VGFR2, SRP14, NUP62, SYUA, VATA, CUX1, TXLNA, STAT3, LAP2A, EPS15, HELZ, MATR3, SSRA, SSRB, KI67, ATRX, MAP1B, YAP1, UTRN, STT3A, SC6A8, RFX5, SOX2, PRC2A, HSP13, NR2C2, NASP, CDK8, DHE4, YLPM1, NU153, RBP2, TAF6, MRE11, EMD, MXI1, MAP2, TOB1, PPT1, TCPQ, PAPOA, GDS1, AGFG1, CRIP2, NUP98, SMTN, SC24C, HIRA, ATX1, ATN1, AFAD, AF10, AF17, DSRAD, SEC13, NU107, ZN445, CSN2, RL37, WDR5, TIM10, F193A, RBM6, PITX1, IF4G2, PHC1, ADA17, KGD4, RL19, FOXK1, DAB2, RHG04, RBM10, HNRPU, SPTB2, FOXK2, RUNX1, MEF2A, SP2, SP3, PLOD1, KMT2A, TF65, IF4G1, NOTC2, TLE4, PTN12, CALD1, MEF2C, P5F1B, GABPA, ZO1, ACK1, EP300, AHNK, FCHO2, HMGX3, SRBP2, FOXO1, ASPH, TROAP, BPTF, FSTL1, NFIA, DPYD, TP53B, FOXC1, ECH1, ROA0, DDX10, TBX2, GPS2, G3BP1, PABP4, ADAM9, PICAL, NAB1, SERC3, RIPK1, IQGA2, STIM1, CUL4B, ASPP2, CAC1S, RUNX2, NFYC, CDK13, TOB2, VEZF1, UBP2L, GIT2, SRC8, CAPR1, LAGE3, PUM1, MDC1, EPN4, TTLL4, RRP1B, NCOA6, GSE1, MEF2D, LASP1, MYPC3, ZN638, NUMA1, SART3, CND1, R3HD1, KIF14, WDR43, PLCL1, PLEC, NOMO1, NONO, RCN1, RYR3, KS6A1, RBMS2, TAF1C, SF01, MED1, JHD2C, TRIP6, T22D1, ELF2, TAB1, HERC1, NCOA1, VAS1, ZFHX3, ZYX, ADRM1, SYPL1, TAF9, DREB, DGKD, CGT, GEN, LY6K, RFX7, QSER1, AAK1, PRSR3, QRIC1, MA7D1, WDR72, TBRG1, TB10B, TPRN, FIL1L, SVEP1, AMOT, EPC2, CRTC2, PAN3, HS904, YIF1B, AG10A, IGS11, ZN628, BCORL, FIGN, BRD10, SH319, TGO1, PRC2B, TOIP1, CEP78, P4R3B, HP1B3, CE170, ZN362, FKB15, AKND1, ZEP3, LRIF1, SWT1, RHG21, UBAP2, RBM26, DEP1A, OGRL1, AHDC1, F222A, RPRD2, RN220, ZN318, TASO2, ZMYM4, PAPD7, TENS2, KANK2, ARID2, USF3, RHG17, CYTSA, ANR40, BICRL, JADE1, PKHA7, NIPBL, LIN54, TET2, RINT1, CRCDL, ZNT6, TTC41, RHGBA, NFRKB, RSBNL, KCD18, NCEH1, MDEAS, ZC3HE, LARP1, NHS, CRTC3, SAS6, MCAF1, BCOR, MPRIP, DNMBP, GGYF2, THADA, BNC2, NFXL1, NBEL2, CO039, SRCAP, CBAR2, UBN2, XIRP1, RAPH1, LARP4, HAKAI, ASXL2, SPT6H, KDM3B, ZCCHV, KANL1, RGPD4, POGZ, ZFY16, NUFP2, MAVS, CLAP1, EMSY, I2BP2, SRGP1, RBBP6, SH3R1, HUWE1, YTHD3, NPM2, ILDR1, KAISO, MYPN, LDB1, LYRIC, BCL9L, LUZP1, NRAP, RTTN, PRSR1, DDX42, CEP57, CD20B, CACL1, P66A, HIPK1, KCC1D, RN135, MY18B, AHNK2, FOXP4, NAV3, NAV2, MISP, ARI3B, IPRI, TEX2, MGAP, CC28A, Z3H7A, ANKH1, SUGP1, RPAP2, MILK2, SRRM1, ZZZ3, GAR4, PHAR4, RTKN2, DCP1B, XRN1, PELP1, CKLF8, TENS4, SPART, RPTOR, NUP93, ZN687, DOCK4, RHG24, RUSC2, SYNPO, FNBP4, D2HDH, RP25L, ATPF2, CPSF7, ARFG1, ENAH, SPOT1, SUMF1, KCNH5, SLAI1, TNR6A, PHC3, DRC6, CBPC3, NAV1, VP37A, KMT2C, ZMIZ2, BD1L1, ARI1B, FLCN, NUP35, TOIP2, TNIP2, KNL1, OR2L2, PUM2, CC110, TBC15, STT3B, ZN507, ALMS1, DLG5, KCNV2, BRX1, DOT1L, GEMI5, PARD3, ZN384, SMAP2, IASPP, TM263, ZFN2B, NUDC2, PCNP, TRUB1, LMO7, ATX2L, PALLD, P66B, BBX, ZCH14, GBF1, SMG7, RTF1, NICA, PHF3, MAML1, ZN592, LAR4B, TFG, TAF4B, RREB1, SC65, CBP, SYMPK, DDX17, GPKOW, FUBP2, UBP7, LPP, LSM10, NCLN, MRTFA, FUBP1, TTC17, PBIP1, TTC28, TOM6, PF21A, INT12, REPS1, ESS2, MBD6, ELP4, SGF29, RBM33, ZN503, P121A, TONSL, PDLI5, ERO1A, DOCK6, FUBP3, RSRC1, ZN594, VCIP1, ZN462, LCOR, PDLI2, CLP1L, Z512B, ZFR, EP400, MRFL, H6ST2, TIGD1, NOL4L, DOCK7, RPR1A, RBM14, ADCYA, QKI, LENG8, TRNT1, PP1RA, PHF12, CIC, MED15, ERBIN, HMCN1, LMF1, PIGS, WRIP1, SIN3A, MINT, HTF4, EYA3, POP1, TEAD3, TTC1, CSN8, ATX2, ARI3A, ANM1, PKP2, TEP1, DPH2, WAC, DIDO1, HNRL1, RBM4, SSBP4, PRR14, SSBP3, YTHD1, KPCD2, ZCHC2, TB182, AMRA1, CE295, TANC1, ZC12C, CEP44, STRAB, SP130, BRD8, RGAP1, SMG9, APC1, I2BPL, TMX4, KI13A, WDR13, EPC1, ADNP, ZN106, TM245, FOXP1, PABP3, WNK1, E41L1, ZHX3, BICC1, PEAK1, PPR3E, ZN703, PKHA5, CLSPN, BCDO1, RC3H2, ZFYV1, TAF9B, EMAL4, ZBT20, NCOA5, TANC2, ZN532, NCK5L, TNR6C, CHD8, FBSL, APMAP, DMAP1, UBN1, DCP1A, INCE, ANLN, GEPH, PDLI7, TULP4, HOME2, SLX9, DIAP3, BMP2K, RBM12, STAU2, DDX28, CWC25, CARF, ETAA1, ABI2, HXC10, BCLF1, TAB2, CELR3, CDK12, GRHL1, SACS, ITSN2, BICRA, CNOT2, TMEM9, CAC1I, CAMP3, DAPLE, RCC2, DIP2B, MBD5, CT2NL, F135A, KANL3, RERE, SE1L1, TRM7, YM012, KDM5B, LIMD1, TCF20, SUN2, LIMA1, SEPT9, UBQL2, TRPS1, S30BP, NRBP, BAZ2B, SIX4, HOOK1, CDC23, TASOR, GMEB2, TNIK, PARP4, NUP50, ZHX1, CDV3, MCTS1, KCNH3, LRFN2, MRTFB, ZBT21, PRR12, YETS2, HECD1, ZMYD8, NOTC3, SPAT2, SOX13, G3BP2, MAGD2, MINP1, MACF1, CP131, SCAF8, TRI33, PHF8, LIMC1, TNR6B, SRRM2, SCML2, ZN148, POLH, INVS, ICE1, R3HD2, MAN1, TR150, WBP11, ZN281, STA13, WNK2, ARIP4, MTCL1, DCAF1, RPGF2, IRS2, CRBG1, HYOU1, SAM50, PRC2C, YTHD2, NCOR2, GMEB1, DC1L1, EPN1, NCOA3, ZHX2, S23IP, U3KPZ7, V9GYH0
Species: Homo sapiens
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Chen Y, Tang F, Qin H, Yue X, Nie Y, Huang W, Ye M. Endo-M Mediated Chemoenzymatic Approach Enables Reversible Glycopeptide Labeling for O-GlcNAcylation Analysis. Angewandte Chemie (International ed. in English) 2022 61(23) 35289036
Abstract:
To selectively enrich O-linked β-N-acetylglucosamine (O-GlcNAc) peptides in their original form from complex samples, we report the first reversible chemoenzymatic labeling approach for proteomic analysis. In this strategy, the O-GlcNAc moieties are ligated with long N-glycans using an Endo-M mutant, which enables the enrichment of the labeled glycopeptides by hydrophilic interaction liquid chromatography (HILIC). The attached glycans on the enriched glycopeptides are removed by wild-type Endo-M/S to restore the O-GlcNAc moiety. Compared with classic chemoenzymatic labeling, this approach enables the tag-free identification, and eliminates the interference of bulky tags in glycopeptide detection. This approach presents a unique avenue for the proteome-wide analysis of protein O-GlcNAcylation to promote its mechanism research.
O-GlcNAc proteins:
TM271, HEAT9, SBNO1, P121B, GGT3, TAF4, AGRIN, MEIS1, RNT2, NFIB, MA2B1, CDKA1, ABLM1, ADA10, KMT2D, MYPT1, ZN609, SC16A, PLXB2, SET1A, VA0E1, TNC18, TLR3, TET3, PRPF3, IF4G3, ZN207, AKAP8, CALU, EXTL3, PCDH7, TRAK2, GANP, TSN3, MAFK, KERA, SRGP2, ANR17, NCOR1, U520, CRTAP, ATRN, CBPD, RMP, TOX4, SC24D, SRBS2, SUN1, ERLN2, PCF11, SCAF4, VAPB, SMC2, SC24B, CNOT4, EGFR, HPT, GLHA, LMNA, NU3M, VTNC, GCR, A1BG, K2C1, HSPB1, RPN1, AT1B1, K1C18, K2C8, ITAV, GDN, TRY1, SAP, MET, NFIC, VIME, K2C7, SNRPA, UCRIL, ATX1L, NT2NC, LYAG, LAMC1, PPAL, LAMP1, MPRI, TPR, SKIL, T2FB, GLU2B, NID1, PO2F1, MCP, ZEP1, CD36, ATF7, EGR1, SON, ATF1, ICAL, CO5A1, ASPG, SFPQ, ITA3, NKG2A, ARNT, TEAD1, 3MG, TKT, UFO, HXA11, HXC9, ZEP2, HNRH1, ELF1, CD68, RFC1, FBN1, NU214, RL4, SCNNA, SRP14, NUP62, TAGL2, CUX1, IL6RB, PBX2, STAT3, NBL1, LAP2A, LAP2B, LIFR, PCP, ECE1, MUC18, MATR3, SSRB, VDAC2, ATRX, NOTC1, UTRN, RFX5, HSP13, CCN3, AGRE5, NR2C2, CDK8, CENPF, YLPM1, RBM25, NU153, RBP2, TAF6, EMD, PPT1, SMCA4, AGFG1, NUP98, PTTG, ATX1, AT1B3, AF17, DSRAD, LAMB2, CAD13, ITA1, NU107, TGFB2, ACTG, CXAR, PITX1, CNTP1, ABCA4, PHC1, ADA17, KGD4, SARNP, FOXK1, DAB2, HNRPU, SPTB2, FOXK2, MEF2A, SP3, PLOD1, KMT2A, UPAR, IF4G1, NOTC2, SRS11, SUH, MFGM, AHNK, TMM62, GALT1, BST2, SIA4C, ASPH, BPTF, NFIA, FOXC1, ADAM9, NFAC2, MAMD1, NAB1, TBB3, PKD2, LAMB3, KLF5, NFYC, CDK13, SCRB2, LRP8, VEZF1, UBP2L, LAGE3, MDC1, RRP1B, MEF2D, ARI5B, NUMA1, PON2, RCN1, TAF1C, SF01, MED1, JHD2C, ELF2, TAB1, ZFHX3, ZYX, ADRM1, LAMA4, TAF9, FOXD1, LAMA3, RFX7, QSER1, SVEP1, SYTL3, EPC2, LUZP6, CRTC2, YIF1B, BCORL, BRD10, PRC2B, ZN362, LRIF1, UBAP2, RBM26, VP13D, RPRD2, RN220, ZN318, TASO2, ECM29, ARID2, SE1L3, BICRL, SCAR3, NIPBL, LIN54, FSTL4, TET2, ZNT6, GOLM2, NFRKB, S39A4, ZC3HE, FIP1, CRTC3, PLGT2, SUSD1, MCAF1, BCOR, IGS10, B3GLT, CD109, FRRS1, SRCAP, NBEL1, UBN2, RAPH1, HAKAI, ASXL2, SPT6H, KDM3B, NRK, NUP54, POGZ, MAVS, PK1L3, PLGT3, EMSY, RAI1, YTHD3, LDB1, LYRIC, OSTM1, PRSR1, DDX42, P66A, FOXP4, CTL2, TEX2, MGAP, ANKH1, SUGP1, NUP93, PLD6, FNBP4, ARFG1, PMGT2, GOLM1, PGLT1, TM87A, FSAF1, PHC3, SP20H, ARI1B, NUP35, OR6C4, VA0E2, PLBL2, PUM2, SPP2B, DLG5, GPX8, PO210, ZN384, LMO7, MUC16, P66B, BBX, SMG7, NICA, TM131, PHF3, TAF4B, GGH, GSLG1, FUBP2, LPP, NCLN, TTC17, FKB10, TOM6, PF21A, RBM33, I17RA, P121A, PDLI5, CREL1, FUBP3, TXD15, LOXL4, VCIP1, CHAP1, Z512B, ZFR, EP400, HUTU, RBM14, KI20B, PHF12, PANX1, CIC, MED15, ERBIN, JMJD8, MINT, TEAD3, SEM3C, WAC, DIDO1, HNRL1, TM109, NUP58, M4A8, YTHD1, ULBP3, AMRA1, TANC1, TWSG1, S22A4, SP130, APC1, I2BPL, WNT5B, VP33B, EPC1, ADNP, ZN106, FOXP1, PTN23, WNK1, SIA7D, ZHX3, PIEZ2, SG196, TM231, PEAK1, DOCK5, CF155, CNO10, MLXIP, SIAE, PKHA5, RC3H2, TAF9B, ZBT20, NCOA5, TANC2, ZN532, CELR2, APMAP, ABCB9, CHSTB, UBN1, NECT3, PDLI7, RBM12, OCAD1, CARF, ETAA1, HXC10, TAB2, CELR1, CDK12, CNOT2, CRIM1, TMEM9, RCC2, CHD7, RBM27, KANL3, MRC2, SUCO, TCF20, SUN2, UBQL2, LCAP, GGT7, TASOR, GMEB2, ZHX1, DSE, WWC3, MRTFB, ZBT21, PRR12, YETS2, NOTC3, KMT2B, PRP19, MINP1, SCAF8, ZC3H4, SRRM2, SCML2, UST, ICE1, ZN281, DAAM1, IRS2, PRC2C, NCOR2, NPTN, GMEB1, POMT1, S4A7, S23IP
Species: Homo sapiens
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Qin K, Zhu Y, Qin W, Gao J, Shao X, Wang YL, Zhou W, Wang C, Chen X. Quantitative Profiling of Protein O-GlcNAcylation Sites by an Isotope-Tagged Cleavable Linker. ACS chemical biology 2018 13(8) 30059200
Abstract:
Large-scale quantification of protein O-linked β- N-acetylglucosamine (O-GlcNAc) modification in a site-specific manner remains a key challenge in studying O-GlcNAc biology. Herein, we developed an isotope-tagged cleavable linker (isoTCL) strategy, which enabled isotopic labeling of O-GlcNAc through bioorthogonal conjugation of affinity tags. We demonstrated the application of the isoTCL in mapping and quantification of O-GlcNAcylation sites in HeLa cells. Furthermore, we investigated the O-GlcNAcylation sensitivity to the sugar donor by quantifying the levels of modification under different concentrations of the O-GlcNAc labeling probe in a site-specific manner. In addition, we applied isoTCL to compare the O-GlcNAcylation stoichiometry levels of more than 100 modification sites between placenta samples from male and female mice and confirmed site-specifically that female placenta has a higher O-GlcNAcylation than its male counterpart. The isoTCL platform provides a powerful tool for quantitative profiling of O-GlcNAc modification.
O-GlcNAc proteins:
A0A0A6YVU8, A0A1B0GSG7, RBM47, ZN335, A2A8N0, SBNO1, CNOT1, PHRF1, ZN462, TAGAP, D3YUK0, E9PUR0, E9PVW1, E9PWI7, PARP4, E9PZS2, E9Q2C0, NU153, E9Q616, BD1L1, E9Q732, ARHG5, E9Q7N9, E9Q842, E9Q9B4, E9Q9Q2, E9QA22, E9QAE1, F6Y6L6, F8VQ29, F8VQM5, J9JI28, PDLI1, SPT5H, TAF4, ARI1A, ABLM1, KMT2D, MYPT1, ZN609, SET1A, SYNEM, PUR4, TNC18, KDM6A, DPOD2, M3K7, TPD54, SYNJ1, ZN207, SRPK2, ACK1, SYUA, MYPT2, KIF1B, HBP1, OGA, VINEX, PLIN3, MAFK, BRD4, PDLI1, KDM6A, SRPK1, N4BP1, ANR17, NCOR1, CREG1, CRTAP, MYO1A, MTR1L, CREG1, TOX4, SUN1, M3K6, PSMG1, SC24B, CNOT4, ABL1, ABL1, EGFR, LAMC1, LMNA, GBA1, GCR, HSPB1, PPBT, RLA2, ITB1, K1C18, K2C8, SAP, CATL1, LAMB1, ENPL, BGLR, NFIC, VIME, SNRPA, ROA1, ATX1L, TGAP1, GLI2, HLAC, CATB, TAU, BIP, FINC, K2C8, TPR, MSH3, ENPL, PO2F1, ATF2, GNS, ZEP1, RS2, MUC1, JUNB, ATF7, CATD, SON, SERPH, NELFE, BIP, ROA2, CBL, IF4B, APC, ARNT, MAP4, TEAD1, RXRA, RXRB, RXRG, CLIP1, AIMP1, HXA11, ELF1, NU214, MP2K2, VATA, CUX1, PBX2, MLH1, STAT3, SSRB, KI67, STT3A, RFX5, LMNA, DPOD2, PAXI, CDK8, YLPM1, NU153, RBP2, TAF6, EMD, PPT1, FXR1, ICAL, AGFG1, NUP98, ATX1, ATN1, PTN5, AF17, DSRAD, AMRP, ACYP2, NU107, ACOT8, S26A1, TB182, YTHD1, ASXL1, PI5PA, RIN3, MRTFB, RL37, KCNA2, RALA, STIM1, PITX1, IF4G2, SRPK2, RENBP, COG7, WNK1, SERF2, RPTN, SPSY, DAB2, RBM10, HNRPU, SPTB2, FOXK2, EWS, MEF2A, SP2, CO7A1, S30BP, NUCB1, ENL, IF4G1, K1C17, TLE4, TOP1, SUH, CBG, ACK1, DEMA, AHNK, FOXO1, TROAP, BPTF, NFIA, ROA0, G3BP1, PABP4, ATM, PICAL, MAMD1, RIPK1, STIM1, MTMR1, CUL4B, ASPP2, KLF5, NFYC, CDK13, VEZF1, DSG2, TRI29, UBP2L, SRC8, PUM1, EPN4, RRP1B, NCOA6, DIP2A, MEF2D, NUMA1, R3HD1, KIF14, EBP, RCN1, KS6A1, RBMS2, TAF1C, NCOA2, SF01, JHD2C, MARE1, ELF2, TAB1, ZFHX3, ZYX, ADRM1, CCDC6, TAF9, STX1A, RFX7, QSER1, QRIC1, PRC2C, PBIP1, GSE1, TNR6A, CEAM5, Q3UKP4, COBL1, ARH40, SC31A, PEG3, SRBS2, Q3UU43, Q3UUE0, F91A1, ARBK2, Q497W2, Q4KL65, PHAR4, EPC2, CRTC2, BCORL, BRD10, TGO1, PRC2B, TOIP1, SPG17, SHRM1, ZN362, LRIF1, RHG21, UBAP2, RBM26, RPRD2, ZN318, NCOR1, LAMA5, HCFC1, P3C2A, SAP, AP180, MAFK, SPTB2, SH3G1, ZYX, TSH3, INADL, WAPL, KAZRN, SBNO1, ARID2, DYH17, SAM9L, CDK13, LAR4B, BICRL, RHG21, HELZ, TTLL5, PANX2, PKHG2, NIPBL, LIN54, F135A, RPRD2, IF4G1, SPIC, SCYL2, NFRKB, INT1, ZN182, UGGG1, MDEAS, ZC3HE, RICTR, FIP1, CRTC3, SAS6, MCAF1, BCOR, GGYF2, NU188, CO039, UBN2, HAKAI, ASXL2, SPT6H, DDX46, KDM3B, PRC2B, OOG2, ZIC5, NRK, POGZ, MAVS, CLAP1, EMSY, I2BP2, SRGP1, SH3R1, HUWE1, YTHD3, NU214, TMC5B, ZN598, TOPRS, SHAN2, Q80ZX0, ZNF18, Q810G1, BCL9L, LUZP1, PRSR1, DDX42, PALB2, P66A, GNS, LPP, TB10B, TGO1, Q8BIB6, ZN771, ZNT6, AAPK2, CNOT4, SP110, IFFO1, YTHD3, NCBP3, DEFI6, RBM14, CNOT2, CABS1, Q8C6L9, TCAL5, TAB1, SCYL2, ASPP2, PHC3, EPN2, PDLI5, I2BP1, RN135, AHNK2, NAV2, MISP, MGAP, ANKH1, PHAR4, XRN1, PELP1, Q8JZK6, Q8K0U8, AGFG1, TXD11, IL23R, ARHG6, SPART, SPICE, NUP93, CLASR, ZN786, SYNPO, FNBP4, ARFG1, ENAH, TNR6A, PHC3, SP20H, NAV1, VP37A, KMT2C, BD1L1, NUP35, STXB6, KNL1, TCAL3, MTSS1, SPART, NUP35, PUM2, STT3B, ALMS1, GEMI5, WIPF2, MAVS, UTP6, PI3R4, AMOT, P66B, STAG1, PCNP, LMO7, ATX2L, CSKI2, P66B, BBX, TITIN, HNMT, UBAP2, DCP1A, NRIF1, SMG7, RTF1, MAML1, ZN592, LAR4B, TAF4B, SHIP1, DDX17, RENT1, GPKOW, FUBP2, LPP, TTC28, PF21A, INT12, RCN3, CERS2, PDLI5, FUBP3, MY15B, ANCHR, CLP1L, Z512B, ZFR, EP400, NOL4L, RBM14, CIC, MED15, PIGS, DCR1C, SIN3A, MINT, EYA3, TEAD3, ATX2, RFC4, DHX58, ANX13, GORS2, TAB2, EPN4, ANR17, DPH2, WAC, DIDO1, YTHD1, AMRA1, TANC1, TXD12, F133B, RBM33, GPI8, Q9D2U0, ZC21B, FUND2, F162A, APMAP, Q9D809, FIP1, CNPY3, Q9DAV5, Q9DB24, ALG2, PLIN3, MYPT1, WWTR1, Q9EQC8, SALL1, RBP2, GILT, MFF, SP130, APC1, I2BPL, RBNS5, EPC1, ADNP, ZN106, TM245, CPVL, PTN23, WNK1, E41L1, ZHX3, ZN335, PKHG2, CCSE2, CQ10B, MLXIP, PKHA5, RC3H2, TAF9B, ZBT20, NCOA5, ZN532, APMAP, HYOU1, ADRM1, GIT2, BAG3, UBN1, PDLI7, DIAP3, RBM12, CARF, ETAA1, HXC10, TAB2, UGGG1, CDK12, ITSN2, CNOT2, TMEM9, DAPLE, NYAP2, KANL3, SON, LIMD1, KI21B, KI21A, PPIE, PCM1, GALK1, MRP5, SE1L1, LIMD1, TCF20, SUN2, AFF4, UBQL2, S30BP, NRBP, SIX4, TASOR, GMEB2, PARP4, NUP50, ZHX1, YETS2, HECD1, SCAF8, SRRM2, SCML2, S22AL, NCOR2, DEMA, POLH, R3HD2, ZN281, FBX7, RPGF2, IRS2, HYOU1, PRC2C, NCOR2, GMEB1, S23IP, SRPK3, SLFN1, VNN1, KLK4, SE1L1, RGS6, E41L1
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