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Xu S, Yin K, Xu X, Fu L, Wu R. O-GlcNAcylation reduces proteome solubility and regulates the formation of biomolecular condensates in human cells. Nature communications 2025 16(1) 40307207
Abstract:
O-GlcNAcylation plays critical roles in the regulation of protein functions and cellular activities, including protein interactions with other macromolecules. While the formation of biomolecular condensates (or biocondensates) regulated by O-GlcNAcylation in a few individual proteins has been reported, systematic investigation of O-GlcNAcylation on the regulation of biocondensate formation remains to be explored. Here we systematically study the roles of O-GlcNAcylation in regulating protein solubility and its impacts on RNA-protein condensates using mass spectrometry-based chemoproteomics. Unexpectedly, we observe a system-wide decrease in the solubility of proteins modified by O-GlcNAcylation, with glycoproteins involved in focal adhesion and actin binding exhibiting the most significant decrease. Furthermore, O-GlcNAcylation sites located in disordered regions and with fewer acidic and aromatic residues nearby are related to a greater drop in protein solubility. Additionally, we discover that a specific group of O-GlcNAcylation events promotes the dissociation of RNA-protein condensates under heat stress, while some enhance the formation of RNA-protein condensates during the recovery phase. Using site mutagenesis, inhibition of O-GlcNAc transferase, and fluorescence microscopy, we validate that O-GlcNAcylation regulates the formation of biocondensates for YTHDF3 and NUFIP2. This work advances our understanding of the functions of protein O-GlcNAcylation and its roles in the formation of biomolecular condensates.
O-GlcNAc proteins:
MEX3A, CNOT1, RGPD3, P121C, MYO1G, H7C0N4, PDLI1, CBX4, TAF4, RFXAP, PODXL, ARI1A, UTY, ABLM1, DVL1, KMT2D, HGS, MYPT1, XPO1, SET1A, PUR4, FYB1, PPM1G, EIF3H, IPO8, NUP42, KDM6A, ZZEF1, TET3, SI1L1, CP110, SYNJ1, IF4G3, TGON2, WIPF1, HTSF1, MYPT2, SMCA5, GANP, MA1A2, DIAP1, PLIN3, MAFK, KIN17, CUTA, N4BP1, CLAP2, BRE1B, PP6R2, ANR17, ZN217, FLNB, NCOR1, SPAG7, TACC1, GGYF1, ERLN1, WBP4, U520, WDHD1, RNH2A, STAM2, CBPD, STK10, TOX4, ST65G, TOX, PCF11, ENDD1, UBR5, AGFG2, KIF4A, MPZL1, IPO7, BAG4, SC24B, HERC2, ZMYM6, BAG3, CLPT1, ABL1, ADA, LDLR, HLAB, TFR1, CATA, GCR, KITH, HLAA, RPN2, ITB2, RLA2, RLA0, ITB1, RB, TPM3, SAP, LAMB1, SP1, PTPRC, ITA5, VIME, SNRPA, ROA1, ATX1L, DERPC, MYBA, CH60, PABP1, TPR, SKI, XRCC6, LAMP2, ITA4, HCLS1, KPYM, PO2F1, RAC2, IGLL1, RS2, TFE2, LEUK, CREB1, GCFC2, ATF7, SON, ATF1, TFE3, NFKB1, AL5AP, PUR2, ROA2, CBL, IF4B, AT2B4, APC, MGAT1, ARNT, DCK, PYR1, CALR, MAP4, CALX, NDUS1, PML, PDIA3, HMOX2, AMRP, CLIP1, ZEP2, HNRH1, ELF1, ICAM3, TTK, CAH8, BMI1, PTN7, RFC1, ADDA, NU214, MP2K2, HPCL1, VATA, CUX1, TXLNA, MLH1, PEX19, S19A1, LAP2A, LAP2B, STAT1, EPS15, MATR3, KI67, CRKL, UTRN, CAPZB, RFX5, PRC2A, NASP, FAS, CDK8, YLPM1, NU153, RBP2, TAF6, EMD, TCPQ, PAPOA, PLCD1, HCFC1, AFF3, ROA3, AGFG1, RBM5, ACLY, SMTN, RD23A, AF10, DSRAD, CADH4, SEC13, IF6, STX17, CORO7, NU107, SELB, TPIS, RAB2A, DCAF7, TIM10, CNBP, TBA1B, NOMO3, F193A, IGBP1, IF4G2, GTF2I, PHC1, NUCB2, TMF1, RT09, SARNP, FOXK1, FBLN2, FAM3A, RBM10, CDK6, HNRPU, SPTB2, FOXK2, RUNX1, CAP1, MEF2A, SP2, KMT2A, IF4G1, TLE1, TLE2, TLE3, TLE4, REL, GFPT1, GABPA, RING1, VAC14, EP300, AHNK, BRME1, MGAT2, GALT2, TROAP, TROP, BPTF, SFSWA, DPYD, TP53B, IRAG2, NFAC3, ROA0, AIMP2, GPS2, PABP4, S39A6, PICAL, HDAC1, STIM1, MTMR2, CUL4B, ASPP2, PEBB, NFYC, LRP8, VEZF1, BOP1, SEPT6, UBP2L, MLEC, ZMYM3, SRC8, LAGE3, PUM1, EPN4, RRP1B, NCOA6, GSE1, MEF2D, NFIX, ZN638, NUMA1, CND2, CND1, RB3GP, NECT1, PCBP1, PCBP2, ELOC, KS6A1, SC23B, JHD2C, ELF2, TAB1, NCOA1, ZYX, ADRM1, DPYL2, TAF9, DREB, DGUOK, QSER1, RHG31, AAK1, QRIC1, LRRF1, GNPTA, C2CD3, EPC2, PDCD4, BOREA, PAN3, BRD10, TGO1, PRC2B, PCID2, P4R3B, DCA10, SNP47, MANEA, ZN362, LRIF1, CAF17, CAMP1, UBAP2, CPTP, RC3H1, AHDC1, VPS53, RPRD2, ZN318, TASO2, ZMYM4, PAPD7, RN123, MAP1S, ARID2, NIPBL, LIN54, ANR54, PPR18, EDC4, SCYL2, GOLM2, NFRKB, RSBNL, MDEAS, BRAT1, ZC3HE, LARP1, FIP1, CRTC3, SHSA2, PI16, MCAF1, BCOR, GGYF2, NFXL1, QSOX2, YJ005, UBN2, MTSS2, NOL8, SSH2, ASXL2, SPT6H, LRC8D, EIF3M, MEPCE, KDM3B, TRM1L, ZCCHV, EFL1, KANL1, POGZ, ZFY16, NUFP2, MAVS, EMSY, I2BP2, HUWE1, YTHD3, KAISO, LDB1, LYRIC, RETR3, TSYL5, DZIP3, CACL1, P66A, C2CD5, HIPK1, ERI1, DEN1C, VRK3, ARI3B, MGAP, Z3H7A, CDAN1, ANKH1, SUGP1, GLT12, K2013, PHAR4, DCP1B, XRN1, RPTOR, NUP93, ZN687, FNBP4, NOPC1, ZBTB2, EMC1, CB069, EIF1A, KRI1, AFG2A, NHLC2, AVL9, XXLT1, GOLM1, SUMF2, TXND5, KCT2, CHSTE, SLAI1, TNR6A, PHC3, SP20H, KMT2C, BD1L1, ARI1B, NUP35, KNL1, MSPD2, NEDD1, GABP2, PIGX, STT3B, ALMS1, NEK9, NEK7, PO210, GEMI5, PIGO, MZB1, RAB2B, TM263, NUDT8, ZFN2B, HNRLL, PCNP, LMO7, ATX2L, P66B, GBF1, SMG7, RTF1, TOPB1, TSC1, PRCC, RREB1, CBP, DDX17, SEM4D, GSLG1, RENT1, GPKOW, FUBP2, ARHG2, UBP7, LPP, IGSF8, MRTFA, ACD, RAB37, PF21A, DOC10, DCPS, FAF2, TMCO6, ESS2, RMI2, TONSL, PDLI5, ZN414, VCIP1, PDLI2, ANCHR, ZFR, EP400, PRRC1, RBM14, QKI, PURB, SPAG5, CIC, MED15, ERGI2, ERBIN, LYSM1, MYADM, SEBP2, RUFY1, MINT, HTF4, CNN2, AGAP2, CSN8, CGRE1, DOK1, CTIP, SMAD5, S29A1, ARI3A, ANM1, DPH2, PELO, ERP44, ESYT1, CNPY3, WAC, MMTA2, HNRL1, ALG8, TMUB1, YJU2, PAPOG, SSBP3, JAM3, OSB11, NAA15, EIF2A, MK67I, YTHD1, KPCD2, UCK2, FTO, AMRA1, CEP44, BC11B, YIPF3, BRD8, XYLT2, I2BPL, MEGF9, EPC1, SLK, ADNP, ZN106, TM245, TMX1, CENPH, WNK1, ZHX3, DOCK5, PKHG2, KANL2, PKHA5, CLSPN, MYG1, RC3H2, LY9, TAF9B, S38AA, TM165, NCOA5, TANC2, TNR6C, CHD8, RESF1, APMAP, ARFG3, UBN1, GP108, INCE, ANLN, GEPH, BMP2K, RBM12, STAU2, INT10, GPT2L, CARF, PP4R2, TAB2, RM39, EURL, CDK12, HPBP1, ITSN2, ARHGC, CHMP5, FOXJ2, SLAI2, CT2NL, TBC14, RBM27, KANL3, FRM4A, ANFY1, GMPR2, ZN219, SE1L1, B4GT7, STK39, SEC63, SUN2, SRP68, SEPT9, UBQL2, S30BP, PFD2, NRBP, EI2BD, CDC23, GGA2, GMEB2, DP13A, MA1B1, ACINU, NUP50, MRTFB, ZBT21, YETS2, HECD1, TPX2, MGT4A, SYNRG, MINP1, SCAF8, SET1B, TBD2B, DICER, SRRM2, SMC3, SCML2, ZN148, CLC11, POLH, PLAP, TPPC8, WBP11, GIT1, ZN281, SIT1, HBS1L, SALL2, TRRAP, ARIP4, HECD4, CRBG1, HYOU1, ATG4B, KLF12, PRC2C, RBM7, CD2AP, SRPRB, GMEB1, TACC3, STK24, NCOA3, M3K4, ZHX2, NS1BP
Species: Homo sapiens
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Luna S, Malard F, Pereckas M, Aoki M, Aoki K, Olivier-Van Stichelen S. Studying the O-GlcNAcome of human placentas using banked tissue samples. Glycobiology 2024 34(4) 38253038
Abstract:
O-GlcNAcylation is a dynamic modulator of signaling pathways, equal in magnitude to the widely studied phosphorylation. With the rapid development of tools for its detection at the single protein level, the O-GlcNAc modification rapidly emerged as a novel diagnostic and therapeutic target in human diseases. Yet, mapping the human O-GlcNAcome in various tissues is essential for generating relevant biomarkers. In this study, we used human banked tissue as a sample source to identify O-GlcNAcylated protein targets relevant to human diseases. Using human term placentas, we propose (1) a method to clean frozen banked tissue of blood proteins; (2) an optimized protocol for the enrichment of O-GlcNAcylated proteins using immunoaffinity purification; and (3) a bioinformatic workflow to identify the most promising O-GlcNAc targets. As a proof-of-concept, we used 45 mg of banked placental samples from two pregnancies to generate intracellular protein extracts depleted of blood protein. Then, antibody-based O-GlcNAc enrichment on denatured samples yielded over 2000 unique HexNAc PSMs and 900 unique sites using 300 μg of protein lysate. Due to efficient sample cleanup, we also captured 82 HexNAc proteins with high placental expression. Finally, we provide a bioinformatic tool (CytOVS) to sort the HexNAc proteins based on their cellular localization and extract the most promising O-GlcNAc targets to explore further. To conclude, we provide a simple 3-step workflow to generate a manageable list of O-GlcNAc proteins from human tissue and improve our understanding of O-GlcNAcylation's role in health and diseases.
O-GlcNAc proteins:
TRGV5, TVBL4, HV320, TM275, MRCOL, EXC1L, TVB74, TVBK3, FEZF1, DEND3, MTMRB, PI4P2, POTEF, ARB2P, U17L3, LIRA5, S22AV, O6C75, SHSA7, TCPQL, SIG16, O4C45, P12LL, D11L8, ARG33, CD15L, YQ037, ZN487, TDR15, ERVV1, NACAM, ADAS, TAF4, CLIC1, MATN2, LORF2, ACACB, ARI1A, NCKP5, IRS4, ADA10, ZN609, SC16A, MCF2L, ERC2, ARHGB, FANCG, TNC18, MRP5, SIA8E, DDX3Y, MEFV, TET3, TTI1, SI1L1, SERA, M3K13, SNUT1, MTSS1, ZN208, HXA3, HNRPR, IF4G3, FOXO3, KCNQ2, LAT, BUD23, MYO1B, NRDC, S28A2, KIF1C, ADCY3, JIP4, SI1L3, TRAK2, L37A3, OPA1, GANP, KDM1A, FZD6, PLIN3, F262, SH21A, MMP20, ROCK2, LRP5, GLRA3, ZN217, USH2A, MYCB2, NU155, P3C2G, NALF2, CABYR, CDKA2, TRIO, C1QRF, CBPD, OR7C1, RECQ4, RECQ5, BAIP3, ATG12, NFASC, SRBS2, PCF11, ABLM3, SC31A, KCNK2, UBR5, AGFG2, SPN1, UBE4B, SCEL, CAC1H, RTN3, EPN2, STBD1, OR6A2, KCNH1, SMC2, TACC2, ZBT7A, CLD9, AGRL2, GLP2R, LGI1, MBD3, ANGT, EGF, IL2RA, LV223, IGHG3, IGHG4, IGHM, CO1A1, CO2A1, CO3A1, CO4A1, FETA, HMDH, A1BG, TBB4A, CP1A1, RPN2, ALDH2, HEP2, CO5A2, INSR, GELS, TY3H, ACBP, LDHB, NFM, CATD, TBB5, CBG, PFKAM, INHBA, CO4A2, IL6RA, PKHD1, HXB5, DLDH, IFI6, DANCR, DU4L4, O13C7, CC192, EFCE2, TZAP, LYAG, TGFB3, HSP7C, PSG1, MPRI, PRPS2, FGF5, COBA1, CO6A3, RLBP1, KCRU, MYH1, HEM1, K1C13, CO6, COBA2, ENPL, ATF2, VAV, IGLL1, PGCA, ZN823, HSP76, JUND, PFKAL, EGR1, XRCC1, K2C4, CADH2, TSN8, LFA3, ITIH1, TYPH, PSB1, CO9A1, C4BPB, NEBU, PRPS3, FLNA, NF1, MUTA, TENX, IDS, PROZ, KS6B1, PTPRZ, MYL9, CO8A2, ITA3, 3BHS2, SYVC, PON1, VATL, CD82, MAP4, TPP2, EPHA8, CO4A5, MPIP1, V2R, SC6A1, CLIP1, HXD11, 3HIDH, PRDX2, DSG3, C2TA, ACSL1, CSTF2, MRP1, MCM7, BMP8B, HSP74, TRY3, PCGF2, ADML, ADA1A, GGT2, VATA, HSPA9, COFA1, PK3CB, PI4KA, CNDD3, THIM, MAGAC, ABCG1, KI67, NOP2, UTRN, PA24A, OR1D4, TNFL6, SC6A8, IREB2, PK3CG, RPIA, SYCC, RBP2, RGS3, GSK3B, MNX1, F10A1, LRBA, TCPD, SMCA2, GALK1, IRAK1, DHB4, NEK3, ROA3, MYOM1, IMA1, JAK3, ZN142, EFNA4, SMTN, SC24C, ATX3, SYYC, HSP72, RD23A, GYS2, LMIP, SYMC, DPOE2, HDAC4, PER3, BACE1, BCAR1, CORO7, KCNKA, SESN2, FOXL2, ATS20, C1QT8, SIA8F, RS6, VPK21, POK7, HBA, RNF4, IRX4, DEN2B, SSBP2, RT05, EFNB1, AT8B2, EM55, HXC5, PLCB2, PSG9, E2F1, CO4A3, OCRL, NAAA, CO7A1, DSG1, MUC3A, GUC2D, AKA12, DYST, CREM, ROM1, KIME, COAA1, COEA1, KDSR, ZO1, SOS2, PP2BA, DHX9, CACB2, GOGA3, CLM6, SPIR1, EP300, AHNK, KLH30, MN1, SIAT6, WASC5, KCMA1, RB40B, DLG1, NFAC3, ABR, STK4, CSN1, IFT88, KLF10, RFA4, PSMD2, MMRN1, M3K1, IFIT5, PDE3B, ADAM9, BIRC2, OLIG2, CUL4B, DYR1A, PTC1, THOC5, CAYP1, CDK13, CO4A6, NID2, VEZF1, DELE1, BTG3, DYHC1, NPAT, E2F2, FLNC, LRC32, NF2L1, FAT1, HABP2, SCN5A, ITIH4, TRIPC, NCOA6, MBTP1, PTCA, ICAM4, MYPC3, COJA1, NAA25, CND1, ACAP1, SETB1, RRS1, RYR3, KS6A1, SURF1, TAF7, OR7A5, PCH2, MED1, JHD2C, U2AFM, HERC1, CAC1E, B2LA1, MIA, QPCT, RTN1, PPR3A, SIA4B, PTPRN, SYT16, COOA1, DOK7, CSPP1, INF2, MICB, KI26B, BLTP1, MGAL, SC5A9, RPEL1, TMM44, SMTL2, COSA1, KLH38, FR1L6, S4A8, RGL3, WDR72, CCD14, C2CD3, HYDIN, SRCRM, GREB1, SEZ6, DHB12, SETMR, SDK2, DCAF6, ACTMP, ACBG2, ESCO1, FREM1, PWP3B, BRD10, MTX3, AKAP4, PREP, RRP12, FRPD3, DOP1, SNP47, MUC21, SAPC1, SNPC4, EYS, MRP7, RBM20, ZFY27, CAMP1, RM02, ADIPL, LITD1, KANK4, TTBK1, MTCL3, BIG3, STPG1, RAIN, RIF1, OBSCN, MRCKA, EF1A3, AGAP9, TCRGL, RSBN1, C1QL3, DEN4C, RRAGB, USPL1, EST4A, RN123, RN213, EGFLA, TENS2, SAXO2, ICE2, MAP1S, B3GNL, RPGP2, NR3L1, USF3, ZFY26, CR063, S31H1, WWC2, BLT3A, MRCKG, RGPA1, NADE, CPZIP, ALKB2, DHX57, MSD2, ACSM6, BGP11, ZN549, CCD62, S2534, TXD11, XKR2, GP179, TRAF7, CORO6, ARL9, ANR11, CRTC1, CSPG4, CP20A, EST3, PRS57, XKR5, PAMR1, ITIH6, FAT4, NHLC1, BCOR, IGS10, GGYF2, ATL5, EMAL6, NFXL1, ALKMO, NBEL2, DMP46, NBEL1, RFLA, CFA47, TPGS1, UBN2, MROH5, TPRG1, PHLP2, ACD11, UBP54, UBP43, UB2R2, MD13L, KNDC1, SND1, TEDC2, NDUF7, FA2H, VASH1, GRP2, CXXC5, ZCCHV, GLUCM, VP35L, MIER3, CA2D4, ZFY16, CSMD3, LFG1, PK1L2, GDF7, HEAT3, MDGA2, BRAP, MUC19, RBBP6, SH3R1, RAB44, TAF8, TARSH, VP13B, TMED4, UBP37, K1328, ALPK2, ACOT1, CN023, PABP2, MTA70, LYRIC, BCL9L, GGN, CASZ1, VASH2, MOGT3, NAL14, NLRC5, HEAT4, RLGPB, CIR1, MICU3, HOGA1, GA2L3, PRSR1, ZGRF1, THNS2, C2CD5, KR191, TMTC1, AHNK2, MGAP, ANKH1, LMTD2, ZC3H3, SP8, MILK2, REM2, ELMD2, CORA1, MAML2, DSEL, MIER1, INT1, SCML4, LRC43, CIART, CLIP4, CMYA5, PRR18, T151A, PAQR4, FBXL6, SRBD1, NKAP, TBC19, MEIOB, IGSF1, TEX26, ARFG2, ARI2O, SIGL1, DCBD1, AMER3, ABTB2, SYT2, TRIML, DAAF3, LMTD1, LMBL4, FANCB, NHLC2, DRAXI, SIDT2, DHB11, P5I13, CAMKV, MACA2, SMG8, RN214, HMCN2, CC168, RB15B, TNR6A, BHE23, DIK2A, CC112, ZSWM2, ZMIZ2, COMA1, MYRIP, NEK11, O4F15, O51S1, OR6T1, OR6K6, O10R2, OR1FC, LIRB1, KI18A, NPL4, MUC7, ALMS1, LKAM1, MO2R1, DSCL1, GTR14, GRP4, ACTT1, CHD5, DMXL2, DLG5, HAVR2, LMX1A, SH3R3, FBF1, FNIP1, ZN526, SETD7, COG1, FRS2, LMO7, TCP11, NELFB, PALLD, ATRIP, STON2, AGRV1, CNKR2, MUC16, ASB15, TITIN, S39A7, NR4A3, MAML1, HS105, AKAP1, RYR2, TENR, RREB1, CBP, PRG4, GLMN, DVL3, TNR25, SBP2L, NKD2, POP5, P33MX, SOSB2, RPE, CR3L1, MSI2H, NXPE2, SGTB, TV23C, DTBP1, MCRS1, MISP3, LRC46, CFA36, FUBP3, BOD1, HPDL, LRIG1, M4A14, DYH8, VPS39, SPTCS, MAML3, ZN333, LCOR, MY15B, S35E1, PHAR3, P12L2, ALPK3, IFT74, PRRC1, CCDC7, DNHD1, HARB1, CFA44, SPT33, TTC14, HUTU, LRRC7, QRFPR, GPR78, NOE3, PGRP2, ARHGH, GBP4, SIG12, PP1RA, DEPD7, PHF12, MAGI1, TAAR9, MED15, GCP5, HMCN1, CRML, WDR24, S22AC, TM209, CP2S1, NIBA2, STAC, AGAP2, PLIN2, PKP4, PI3R4, PMYT1, EPYC, LYST, COCA1, SMAD5, ZNT3, TEP1, TBL1Y, SLF1, SYT15, TIRR, PSF3, ZN649, CRDL1, ZSA5A, TBB2B, TRPM6, C1QT4, PINK1, COPA1, TEX15, AN30A, PECR, K1671, CDHR2, GSDMC, TRI55, PITM3, CPEB1, BARH1, IRX2, CFA74, TANC1, LNP, ASXL3, SRCN1, WDR33, ARP3C, BRD8, RBM38, I2BPL, TRPV6, S6A17, GBA3, CAD23, VPS11, SLK, HIPK2, ATS10, E41LB, UBQL3, KDM4C, SEM6C, RIBC2, PDZD7, AT133, MT21D, CCNP, GORS2, PANK3, SPE39, AGO3, RPA2, S28A3, LY9, TENS1, TM165, NCKX3, MUC5B, CAN10, SPON1, NCOA5, TANC2, PLXA4, TRI39, PDE11, WDR6, MLXPL, OSGEP, UBN1, MEPE, ANO2, BIRC6, ARHG4, KCNQ5, MXRA5, 4ET, SH2B1, SNRK, KCNH7, SIA7A, LRC4B, MDN1, FX4L1, ZBT40, DECR2, SPS2L, HAUS2, AKIP, OXSM, SIR5, ZDHC7, ING3, SNTG2, SO1C1, DACT1, CELR1, CELR3, ERAP1, PDXL2, DTL, PA2GE, MYOF, CNOT2, LRP1B, FOXJ2, AKA7G, YP008, SIRPG, PHRF1, PLCE1, ZSWM5, DYH2, JCAD, FPRP, CT2NL, CHD7, HTR5B, F135A, UNC79, RRBP1, UBP35, KANL3, NYNRI, ZN219, HNF6, MYO1A, DBF4A, KCNN3, AAKG2, RPGFL, MED13, VAX2, ENPP5, DP13A, BORG2, TLK1, PO2F3, MUC12, CDV3, TEN1, TRPC5, PADI1, DEN2A, F184B, INO80, MRTFB, ZBT21, HECD1, ZMYD8, CAPS1, COHA1, KMT2B, BARX2, PCDGK, PACN2, ATS8, BSN, LHX6, MACF1, PHF8, IQEC3, LIMC1, ANR26, UBP24, CBPC1, TUTLB, SRRM2, CNTN6, CHKB, RT33, ZN652, SL9A8, ANKR6, PKHA6, SIK3, YBOX2, COG6, ZBT32, LSM2, PHOCN, RTCB, GRIP1, WNK2, LMBL1, DMXL1, ARIP4, MTCL1, CRBG1, PRC2C, YTHD2, PCDB7, SPO11, CD2AP, ENTP2, CERT, MRCKB, NR2E3, HEBP2, NCOR2, COX11, AP1M2, NCOA3, M3K4
Species: Homo sapiens
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Hou C, Wu C, Wu Z, Cheng Y, Li W, Sun H, Ma J. Systematic Evaluation of Affinity Enrichment Methods for O-GlcNAc Proteomics. Journal of proteome research 2024 39302247
Abstract:
O-Linked β-N-acetylglucosamine (O-GlcNAc) modification (i.e., O-GlcNAcylation) on proteins plays critical roles in the regulation of diverse biological processes. However, protein O-GlcNAcylation analysis, especially at a large scale, has been a challenge. So far, a number of enrichment materials and methods have been developed for site-specific O-GlcNAc proteomics in different biological settings. Despite the presence of multiple methods, their performance for the O-GlcNAc proteomics is largely unclear. In this work, by using the lysates of PANC-1 cells (a pancreatic cancer cell line), we provided a head-to-head comparison of three affinity enrichment methods and materials (i.e., antibody, lectin AANL6, and an OGA mutant) for site-specific O-GlcNAc proteomics. The enriched peptides were analyzed by HCD product-dependent EThcD (i.e., HCD-pd-EThcD) mass spectrometry. The resulting data files were processed by three different data analysis packages (i.e., Sequest HT, Byonic, and FragPipe). Our data suggest that each method captures a subpopulation of the O-GlcNAc proteins. Besides the enrichment methods, we also observe complementarity between the different data analysis tools. Thus, combining different approaches holds promise for enhanced coverage of O-GlcNAc proteomics.
O-GlcNAc proteins:
NBAS, SBNO1, PRS37, CNOT1, MGT4D, P121C, P20L1, MCRI1, SPT5H, MANBA, AGRIN, BCL9, NOP56, ARI1A, ABLM1, KMT2D, SCAM3, MYPT1, HNRDL, XPO1, SYN3, SC16A, SYNJ2, SYNEM, TIF1A, LAMA5, MCES, DC1L2, CP110, IF4G3, E41L2, ZN207, BUB1, HTSF1, AKAP8, DHX16, MGRN1, TRAK2, KIF1B, PLIN3, MAFK, IF2P, DTNB, MITF, SRGP2, ATG13, ZC11A, PP6R2, ANR17, LRP5, NDUS2, NCOR1, U520, UTP20, WDHD1, FACE1, TOX4, SC24D, SUN1, SC31A, CE152, UBR5, AGFG2, APOL3, IPO7, SVIL, SC24A, SC24B, ZMYM6, GPR75, LATS1, LMNA, GCR, OAT, CPNS1, HSPB1, GNAI2, A4, K1C18, ATPB, SAP, PROF1, HS90A, HS90B, NFIC, VIME, SNRPA, ATX1L, DERPC, PTPRF, ESTD, ODPB, C1TC, SKIL, XRCC6, CCNB1, ATF2, NDKA, GNS, ZEP1, AT2A2, ATF7, ERCC2, EGR1, ATF1, GGT1, AT2B1, FLNA, 5NTD, PUR2, RFX1, SFPQ, RS3, STOM, ARNT, MAP4, KPYR, CLIP1, HXC9, ZEP2, QCR1, I5P2, ELF1, MRP1, TRY3, CAH8, T2FA, NU214, NUP62, STAT3, ECHA, HNF4A, LAP2A, LAP2B, CACP, GATA4, BAG6, MAP1B, STT3A, RFX5, PAXI, LMAN1, CDK8, CENPF, YLPM1, NU153, RBP2, GSK3A, TAF6, DNLI3, EMD, TCPD, PAPOA, HNRPM, AGFG1, STAT2, NUP98, ATX1, ATN1, RD23B, AF10, BRPF1, NP1L1, EIF3B, MAZ, HDAC4, IF6, H2B1D, ACTB, ABCE1, RRAS2, ERF1, RL32, ACTG, EF1A1, CSK21, F193A, SARNP, FOXK1, ZBED6, RHG04, CLH1, SPTB2, TIAR, SP2, DYST, ATP7A, NOTC2, TLE4, PTN12, FAK1, ZO1, ACK1, DHX9, EP300, AHNK, NEXN, CPSF1, WASC5, TROAP, BPTF, SF3A3, TP53B, PTN13, TRAP1, DLG1, ATM, PPIL2, TCOF, SMAD4, PICAL, MAMD1, ASPP2, RIN1, CDK13, VEZF1, UBP2L, SRC8, CAPR1, ITPR1, MDC1, RRP1B, UBP10, COG2, MEF2D, LASP1, IMB1, PLEC, NONO, SF3A1, JHD2C, ELF2, TAB1, NCOA1, ZYX, UAP1, DREB, PDS5A, QSER1, TXIP1, FHOD3, P3H1, EMAL3, ZN791, CHD9, TB10B, FND3B, CRTC2, BCORL, BRD10, CE170, ZN362, ZEP3, UBR4, UBAP2, AHDC1, RPRD2, ZN318, TUT7, RN213, TENS2, WAC2A, MAP1S, ARID2, TENS3, ANR40, BICRL, ATLA3, CPIN1, GRAM4, NIPBL, LIN54, RINT1, L2GL2, C2D1A, SCYL2, NFRKB, ZC3HE, LARP1, ZNT9, CRTC3, GLB1L, MCAF1, SRCAP, CFA65, UBN2, VP13C, RAPH1, H32, TMUB2, MARK2, DDX46, EIF3M, KDM3B, GRIN1, KANL1, POGZ, NUFP2, MAVS, DHX29, EMSY, RAI1, I2BP2, HUWE1, YTHD3, KAISO, MYPN, LYRIC, KTN1, PRSR1, DDX42, P66A, RHPN2, NUDC3, AHNK2, GID4, ARI3B, MA7D3, MGAP, LMTK2, ANKH1, SUGP1, MILK2, SRRM1, PHAR4, DCP1B, PELP1, TENS4, CCAR2, HROB, TXND5, RB15B, TNR6A, PHC3, NAV1, VP37A, SYNE1, FAD1, NUP35, TOIP2, KNL1, O52N2, TRY6, COP1, GOGA5, ALMS1, DLG5, MORC4, GEMI5, PIGO, PARD3, IPO4, TM263, PDC6I, LMO7, ATX2L, PHIP, P66B, BBX, SMG7, F168A, PRCC, RYR2, US6NL, SYMPK, RENT1, PF21A, EFHD2, KBP, F120B, RBM33, THOC1, P121A, PDLI5, VPS50, DNJC1, Z512B, ZFR, EP400, SENP8, PRRC1, DOCK7, RBM14, NEK1, PP1RA, PHF12, SPAG5, CIC, ERBIN, SIN3A, MINT, HTF4, VAT1, PLIN2, DNJC2, MNT, KIF2C, ATX2, ARI3A, GDF15, WAC, DIDO1, RFIP5, SRRT, SETD2, ABCA2, TRIM5, TANC1, ZN407, SP130, PUS7L, I2BPL, NELFA, CHM4B, WNK1, ZHX3, CDCP1, Z385D, BCAS3, PEAK1, ECT2, JUPI2, ZN408, PKHA1, ILRL2, RC3H2, TAF9B, NCOA5, TANC2, SYTL2, CHD8, UBN1, DCP1A, SEM4B, HPS4, PDLI7, 4ET, CHRC1, F1142, LTBP3, SYIM, IGSF5, RBM12, STAU2, TYDP1, SLTM, CARF, HXC10, TAB2, CDK12, ERAP1, MYOF, BICRA, CNOT2, SPN90, F135A, RBM27, KANL3, ZMYM2, LIMD1, TCF20, UBQL2, S30BP, PUF60, SIX4, TRM6, NOCT, TASOR, GMEB2, ACINU, CNO11, ASAP1, YETS2, ZMYD8, MYO6, SPAT2, PRP19, NSF1C, MACF1, TRAK1, SRRM2, SCML2, PDE10, AP3M1, ZN281, ZN711, TLN1, ARIP4, DCAF1, WAC2C, RPGF2, CRBG1, PRC2C, UTP18, SNX9, NCOR2, BIG1, NCOA3, PCLO, 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, PHB1, MYH9, COPB2, ADDA, BASI, FUS, NU214, DEK, MP2K2, ATPG, RL4, SRP14, NUP62, RBMX, HSPA9, 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, 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, S25A3, CLH1, HNRPU, SPTB2, FOXK2, CAP1, LAT1, EXOSX, EWS, RL18A, FKBP4, RL6, TOP2B, KMT2A, LMNB2, TF65, IF4G1, 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, AFG2A, 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, ZMYD8, DD19B, PRP19, MAGD2, FAF1, TRI33, SRRM2, PA2G4, RUVB2, RUVB1, VDAC3, E41L3, TR150, NOP58, SHLB1, LC7L2, TMED7, STRAP, RTCB, TLN1, HYOU1, PRC2C, SP16H, COPG1, DC1L1, S23IP
Species: Homo sapiens
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Zhang W, Liu T, Dong H, Bai H, Tian F, Shi Z, Chen M, Wang J, Qin W, Qian X. Synthesis of a Highly Azide-Reactive and Thermosensitive Biofunctional Reagent for Efficient Enrichment and Large-Scale Identification of O-GlcNAc Proteins by Mass Spectrometry. Analytical chemistry 2017 89(11) 28510447
Abstract:
O-linked β-N-acetylglucosamine (O-GlcNAc) is a ubiquitous post-translational modification of proteins in eukaryotic cells. Despite their low abundance, O-GlcNAc-modified proteins play many important roles in regulating gene expression, signal transduction, and cell cycle. Aberrant O-GlcNAc proteins are correlated with many major human diseases, such as Alzheimer's disease, diabetes, and cancer. Because of the extremely low stoichiometry of O-GlcNAc proteins, enrichment is required before mass spectrometry analysis for large-scale identification and in-depth understanding of their cellular function. In this work, we designed and synthesized a novel thermosensitive immobilized triarylphosphine reagent as a convenient tool for efficient enrichment of azide-labeled O-GlcNAc proteins from complex biological samples. Immobilization of triarylphosphine on highly water-soluble thermosensitive polymer largely increases its solubility and reactivity in aqueous solution. As a result, facilitated coupling is achieved between triarylphosphine and azide-labeled O-GlcNAc proteins via Staudinger ligation, due to the increased triarylphosphine concentration, reduced interfacial mass transfer resistance, and steric hindrance in homogeneous reaction. Furthermore, solubility of the polymer from complete dissolution to full precipitation can be easily controlled by simply adjusting the environmental temperature. Therefore, facile sample recovery can be achieved by increasing the temperature to precipitate the polymer-O-GlcNAc protein conjugates from solution. This novel immobilized triarylphosphine reagent enables efficient enrichment and sensitive detection of more than 1700 potential O-GlcNAc proteins from HeLa cell using mass spectrometry, demonstrating its potential as a general strategy for low-abundance target enrichment.
O-GlcNAc proteins:
A0A024R5Q9, A0A0B4J1V8, A0A0B4J203, GAL3B, RBM47, UBA6, ESYT2, SHOT1, NDUF8, VWA8, GTPBA, A5Y5A3, CNOT1, PGP, TBAL3, SMHD1, LR14B, NCF1B, A8K005, B4DLC0, B4DLN1, B4DQP1, B4E259, EIFCL, B7Z2B5, B7Z532, B7Z5W1, DSA2D, D7RF68, E9PL57, E9PLN8, I1VE18, L0R5A1, S35U4, ADAS, KIF2A, PDLI1, AIP, SMAP, NDUS8, PSD12, PSMD9, SPT5H, DFFA, HIP1, EIF3F, IPO5, SAP18, IF2B3, DNM1L, NDUA4, PSDE, BIN1, NOP56, DDX3X, IMA3, TRI38, SCD, DHSD, TXND9, ACPM, UBFD1, AP3D1, DVL2, CHD1, TOR1A, PTGES, ACOT8, NHRF1, TPP1, RPC1, PSA7, MGST3, HAT1, QCR8, HGS, S27A2, HNRDL, BTAF1, SR140, PUR4, NPC1, DEGS1, SCAM2, ARPC2, ARPC3, BET1, RPAC1, PGRC2, NDUA1, RER1, OGT1, PMM2, HMGB3, EIF3H, ARPC5, DHX15, CYB5B, PRP4, SERA, NDUS4, PAPS1, ZW10, SNUT1, RIPK2, PRPF3, PPIH, E41L2, XPOT, DNPH1, PSCA, TSN6, ZN207, NDUB5, BUB1, BUD23, GATC, HTSF1, SGTA, SYNC, PM34, CPSF5, SCO2, ORC5, TIM8A, DHX16, SMCA5, MAST3, OPA1, TBCD4, OGA, HNRPQ, DIAP1, MOT2, IMA7, CTND1, DPM1, Z354A, RT14, PQBP1, TI17B, DKC1, IF2P, EDF1, MCE1, ABCB7, WDR1, BRE1B, ZC11A, CLU, ANR17, COQ9, NDUS7, NIPS2, VPS4B, CISY, PR40A, NDK6, VP26A, MPPB, PSIP1, ECI2, BAF, SF3B1, CSDE1, PRKRA, NPM3, ERAL1, TIPRL, KS6A4, NU155, XRP2, WDHD1, CRTAP, S2512, RNH2A, EIF3G, AP1G2, IDHC, CCNK, SPF27, FLOT1, GLRX3, RL1D1, CLPX, CIAO1, SRP72, LTN1, UFL1, UBXN7, PRP6, PCF11, GLSK, SC31A, RPP14, YIF1A, AGFG2, NDUB6, UBE4B, ELP1, NDUB8, LC7L3, KIF4A, MPZL1, 6PGL, PAPS2, TRI16, ARI2, PSMG1, MPC2, ASML, RPP29, HS74L, AP2A1, TTC4, TM50A, BAG2, EMAL2, BPNT1, NDUBA, ACL6A, COX1, COX2, PNPH, AATM, EGFR, ADA, ATP6, RASN, LDLR, HLAB, LMNA, ALBU, NU1M, NU3M, NU4M, KITH, K2C1, G3P, HLAA, TYSY, RPN2, HMGN1, ISG15, LA, K1C18, DCUP, CDK1, NPM, HEXA, HYEP, H10, SAP, SYEP, HNRPC, YES, SP1, DAF, PFKAM, HEM3, CY1, RU17, ITA5, RS17, RSSA, RM03, SRP19, RU1C, HMOX1, RU2A, PARP1, LKHA4, ALDOC, F86B2, GSTT2, ST1A3, HS71B, RPAC2, ECE2, EFMT4, EFCE2, MYBA, HLAC, ARAF, H14, ODP2, MGST1, ESTD, GTR3, UMPS, ODB2, ACADM, TOP1, TOP2A, G6PD, PYC, C1TC, ADHX, PRPS2, PABP1, ADT1, TPR, KCRB, RINI, K2C5, ACPH, GYS1, ENOB, CD59, MTDC, MIF, CX7A2, CCNB1, CX6B1, PO2F1, SYDC, PLAK, QCR7, ERF3A, EZRI, UCHL3, MCP, PHKG2, RS2, RFA2, CBR1, BGAL, EPCAM, NCPR, AT2A2, UBF1, NDUB7, KAPCA, CAN2, PYRG1, DDX5, LEG3, TCPA, PTN1, RL7, NELFE, PGAM1, RCC1, XRCC1, RPAB1, NDUV2, SPEE, IF2B, BTF3, H2A1D, COX5A, LMNB1, IMDH1, FLNA, ACOHC, ODBB, MUTA, OSBP1, PIMT, FBRL, PUR2, HEMH, TCEA1, AT2B4, CPT2, KTHY, RIR1, EF1B, AT5F1, THIL, CDK2, APC, MCM3, THTM, RS12, TYY1, ITA3, DDX6, DNMT1, U2AF2, HMGB2, SYVC, EF1G, AOFB, RL10, RFA1, APEX1, MAP4, CALX, PSA5, NDUS1, RXRB, AMPL, T2EA, T2EB, IMPA1, 3MG, CO4A5, SPB3, ATPD, PDIA3, 2AAB, CDC27, NMT1, PURA2, AMRP, PUR8, CLIP1, GSTT1, CTR1, AL1B1, RPB2, METK2, RIR2, HNRH3, HNRH1, STIP1, L1CAM, P5CR1, TF2H1, ACSL1, CSTF2, MCM4, MCM5, MCM7, RFC4, RFC2, RL22, T2FA, GDE, SOAT1, BASI, NU214, DEK, K22E, PRS7, HEM6, RL4, LONM, SRP14, NUP62, TALDO, RBMX, COIL, RS19, FEN1, CAP2, TCPZ, NNMT, PSB10, RL13A, PEX19, MDHM, RFC5, RFC3, BUD31, GARS, SYIC, EIF1, LAP2A, LAP2B, MTREX, AK1C3, LPPRC, THIM, MATR3, GPDM, SSRA, SSRB, RANG, PRS6B, EFTS, VDAC2, UBP5, RECQ1, ATRX, IF2M, MP2K3, BRCC3, RL27A, RS10, GNPI1, CAPZB, IF1AX, SYQ, RL29, MAOX, PTSS1, IDHP, PIPNB, DPOD2, TCPG, RM19, INPP, SRP09, SYCC, PSB2, MCM2, ACADV, TMEDA, RBM25, NU153, RBP2, NDUV1, GSK3B, 5NTC, SPS1, GUAA, DNLI3, GDIB, CPT1A, F10A1, DYN2, MAP2, CDK7, CDK9, LRBA, RL14, PAPOA, FXR1, FXR2, RAB7A, RB27A, RT29, SMCA4, SSRD, TPMT, SSDH, DHB4, PSMD7, STA5B, UBP11, DYLT3, CCNH, ROA3, HNRPM, IMA1, NCBP2, MP2K6, KIF11, HXK2, BIEA, PLK1, NUBP1, MAP11, SUCA, MVD1, PGTB2, CATC, CCHL, IST1, TCP4, PMS2, SYYC, UBP14, RD23A, P5CS, IF5, PSMD4, BI1, MFAP1, AFAD, NP1L1, CASP9, ADK, DSRAD, SEC13, HNRH2, SCOT1, EIF3B, ATP5E, IF6, CF298, TMM33, CORO7, SELB, MTPN, YBEY, ARPC4, CD81, SC61B, ROMO1, MYL6, ACTB, IF4A1, PRPS1, PSA6, CDC42, DEST, GMFB, RAB8A, SPCS3, SRP54, RAB5B, UBC12, UBE2K, RAB14, TM258, CSN2, RAP1B, RS3A, VA0D1, NAA20, S61A1, B2MG, COPZ1, UFM1, AP1S1, HNRPK, TIM10, PRS4, RS15A, RS23, RUXE, SMD3, PRS10, ERF1, RS6, RAN, RL23, RS15, RS24, RS25, RS26, RBX1, RL39, RL31, RL10A, RL32, PPIA, TRA2B, AP2B1, DYL1, DYLT1, RL38, SKP1, ACTG, UBC9, CSK2B, UB2L3, EF1A1, CSK21, SERB, IF4G2, RPP38, GTF2I, RAE1L, GSTO1, PRKDC, IN35, RT25, RT35, RT05, RT11, RT15, RT06, RT09, LACTB, RL36A, ARF5, ERH, H33, FOXK1, PRR5, RBM10, TFAM, PIPNA, TF2B, CDK16, NFKB2, FKBP3, HNRPU, U2AF1, SET, GALK2, FABP5, CAP1, HMCS1, IFM3, RL18A, NUCB1, RL6, M2OM, LMNB2, TAP1, TAP2, CEBPZ, TF65, UBXN1, IF4G1, ATP7A, CSTF1, FAK1, SRS11, EF1A2, PUR1, PSME1, GABPA, FMR1, RL18, C1QBP, KHDR1, KLC1, SRSF1, QOR, LG3BP, PPID, SSRP1, CAMP2, NSUN2, RBBP4, NCBP1, HSP7E, BST2, MPPA, WASC5, TBL3, SNF5, STX4, GRSF1, NFIA, SF3A3, TP53B, ILF2, ILF3, PTPRJ, TRAP1, TRAF2, MYO1E, PP1R8, BNIP2, CSTF3, STRN3, RM28, ACACA, CSN1, CAF1B, TADBP, ROA0, AIMP2, PAK2, CBX3, STK3, STX5, SRSF5, SRSF6, TIF1B, PABP4, EIF3I, 2A5G, UB2V1, RM49, DC1I2, ILK, SNTB2, PPIG, TCOF, SF3B2, HAP28, ROCK1, PICAL, SQSTM, PRP4K, HDAC1, DCTN2, SNW1, STIM1, PWP1, MTMR2, CUL2, CUL4A, FCL, RAB31, RAB32, NOG2, TBB2A, BYST, PEBB, HNRPD, IL18, VEZF1, TRI29, TTL12, DPOA2, ICT1, DCTN1, ELOA1, TRI25, FASTK, PDE3A, RBM39, HLTF, MCM6, ITPR1, TRIPC, PUM1, EPN4, SMC1A, RRP1B, PSF1, UBP10, GANAB, LBR, CHD4, NOLC1, NUMA1, PSME4, GAPD1, CND2, SPCS2, EMC2, FL2D, PSMD6, ABRX2, SART3, CND1, EXOS7, U5S1, SYLM, WDR43, ACOX1, EBP, PCM1, NOMO1, PON2, TEBP, NONO, RAB35, RNPS1, PCBP2, ELOC, UBE3C, DHC24, SF3B3, RSU1, SF3B4, SF3A1, SKI2, MARE2, NHRF2, TSN, SF01, CIP4, TRIPB, PCH2, ELAV1, SMAD2, TBCE, VAMP3, ZYX, SEPT7, CCDC6, UAP1, PSMD5, MK14, RM23, DPYL2, SYPL1, RBBP7, IF16, KYNU, CAH9, NDUA9, PCKGM, HCDH, UGPA, MIC60, HNRL2, INF2, PDS5A, TSR1, ALG11, QRIC1, SMU1, LRRF1, RPA43, LSM12, ACSF3, PPCEL, TB10B, MIX23, RHG29, PDCD4, Q53FH6, DHB12, UBP39, LACB2, EIPR1, H90B4, TM41B, HERC4, PP6R3, PREP, PSMG4, RRP12, TOIP1, SYAM, PCID2, S27A3, SAMD9, CYTSB, TDIF2, TTC38, EXOS6, NU188, CE170, SNPC4, CHCH9, DDX59, GPTC4, HECD3, SKT, RM02, UBAP2, LCE1C, ACBD5, EEIG2, RBM26, DCAF8, ATPF1, DDI2, NT5D1, LYRM7, EMC10, RIF1, OBSCN, CE350, AXA81, PR38B, BRE1A, ARHGG, LYPL1, ECM29, MIC13, MAP1S, UTP25, TENS3, INT3, C19L1, CPIN1, SMYD5, PTRD1, CNDH2, MRM1, P4R3A, ACD10, DE10B, SCMC1, TM214, HIBCH, TTI2, DHX57, CDC73, RM14, L2GL2, MET2B, PRP8, TTC27, SKI3, RICTR, LCLT1, WDR82, HACD2, GGYF2, KAT3, UBR3, LARP4, SPT6H, SND1, COX15, DDX46, TM10C, 5MP2, EIF3M, MEPCE, CYFP1, ENOF1, FAKD5, MOB1B, KDM3B, HS2ST, K2C74, TRM1L, ZCCHV, NUP54, MAVS, DHX29, COMD6, HDDC2, LIMS2, I2BP2, PATL1, SETD3, PABP2, MTA70, ZN598, AACS, LUZP1, GP180, CAND1, CMTD1, PRSR2, CARM1, COMD2, DDX42, SPAS2, CONA1, NDUAB, DAAF5, STX12, DLP1, P66A, HORN, RB6I2, FABD, CEP97, CTL2, PHF6, UBR1, CHERP, ANKH1, DJC10, NELFD, MIRO1, RB12B, RAVR1, SPB1, SRRM1, CA174, MICU2, TRM2A, PELP1, ABI1, CCAR2, NDUF2, NUP93, LRC47, GPD1L, TOM5, OGFD1, TTC9C, JAGN1, CPSF7, PAF1, ZN433, F241A, ENAH, CL029, NHLC2, SUMF2, UBAC2, RDH13, UXS1, FA98A, NNRE, LS14A, PAPD5, CNNM3, MCU, FBX22, FAD1, NUP35, Q8NFH9, ABHDB, KNL1, OR1M1, ATLA2, MSPD2, DDX55, NCOA7, THOC2, WDR36, G45IP, CX038, FBX30, S35B2, PUM2, KLDC4, UBA3, KISHA, TBC15, RM30, PNPT1, SSH3, GEMI5, IPO4, ABHD5, ZC3HF, TF3C2, TM263, PTPM1, NU133, CHAC2, THEM6, LEO1, CPLX3, NUDC2, TFB1M, SCFD1, HNRLL, CD003, SEN15, SPRY4, TRUB1, LMO7, ARAP3, PRP31, PALLD, CTBL1, DDB2, DDX1, DHB8, PSMF1, RTF1, RT27, AP3S1, LAR4B, GCN1, PRP16, NU205, TFG, RBP56, DDX17, RAD50, CELF1, OSTF1, ARHG1, UFD1, RENT1, SMRC1, FUBP2, ARHG2, UBP13, USP9X, SCAM4, OSBP2, F162A, RM24, SYAP1, EXOC4, FERM2, PBIP1, RPE, EXOS8, OTUL, CHCH1, PPWD1, COA7, TAM41, F136A, P5CR2, DCPS, FAF2, FXRD1, AP2M1, GCP3, F241B, NMD3, RMD1, MSI2H, SNR40, RM38, RRFM, SIR1, YIPF6, KBP, GNA1, RT24, MOCOS, DAZP1, SAAL1, SIM12, DNJA3, TMA16, PTCD3, OTUB1, TRM61, HMCES, DUS3L, OTUD5, VMP1, DCNL1, SMRD1, TOE1, DDX27, MTNB, ZC21A, PGAM5, FUBP3, SPF45, SUCB2, HPDL, ATPMK, BT3L4, CLP1L, SCYL1, SNX27, YTDC1, ZN830, NSUN5, RBM14, UHRF2, PUS7, TRNT1, TEFM, EFGM, NUDC1, PRPK, PIGS, WRIP1, RANB9, NEB2, IWS1, SIN3A, OSBL9, INT14, ZSC10, RUFY1, MMS19, UHRF1, NIBA2, YMEL1, MYCBP, TBCB, PSB7, CNN2, CDC5L, PSMD1, PFD5, MPH6, TSNAX, TTC1, CSN8, KIF2C, MGLL, HCD2, NP1L4, ACON, TCPH, ANM1, VRK1, CND3, ARP5L, DDX50, RNZ2, GRWD1, MACD1, CMS1, MEP50, RT63, TBA1C, MBB1A, FERM1, CG050, PDD2L, MIC25, UQCC2, UTP23, LXN, NTPCR, TACO1, GCP2, SEN34, ALG1, PSMG3, CSN4, DIDO1, RBM42, MCMBP, AASD1, DCNL5, AN32E, P4K2A, MMTA2, TBB6, PAXX, HNRL1, MTNA, WDR18, THUM3, NTM1A, ASHWN, TM109, GNL3, NUP85, ELOV1, IFT27, SYTM, THIC, FUND2, SF3B5, SFXN3, CARD6, OSB10, B2L13, SRRT, HDHD5, HDAC8, RTF2, EIF2A, PDIP3, RM13, RM04, RM01, MK67I, YTHD1, RT26, NEUL, ABCA2, RM37, UACA, WDR11, CRNL1, TBL1R, UBL5, API5, FTO, UBE2O, XPO4, S39A8, WDR12, YIPF3, DERL2, SKI8, SLIRP, EGLN1, UBA5, MKRN2, T126A, PAIP1, CLPB, LSG1, NAT10, ILKAP, XRN2, NSRP1, CSTFT, 5NT3A, GBRL1, DDX47, RM18, SNAB, ATG5, GAN, DPH5, DHX36, IPYR2, SPNS1, RM46, PININ, GPT, GHITM, NELFA, OSGL1, HEAT1, TFB2M, YTDC2, RPAP3, WDR26, CH033, UBE2Z, ANKZ1, ZWILC, MED20, ARMT1, L2HDH, ELP3, RPA2, KT3K, SYCM, PPCS, NJMU, XPO5, GRPE1, RHOF, POPD2, EMAL4, GLOD4, MCCB, MOV10, SYF1, TM9S3, WDR6, TRXR2, SYSM, RT30, M1IP1, UBE2T, NOP10, OSGEP, MED4, RM40, RTN4, RPR1B, RRAGD, PFD4, EXOS3, ANLN, XPP1, BIRC6, ACSA, DBLOH, DDX21, SAR1A, DPOE4, SIAS, SHLB2, EI2BG, MBNL1, PYRG2, DPOE3, CHRC1, COXM2, ABC3C, PNO1, RM17, HELLS, SYFB, SYIM, ATG3, SMC4, OLA1, CUTC, ECHD1, RBM12, MDN1, DECR2, ABHDA, LIN7C, TXLNG, SPS2L, ABCF3, DD19A, UQCC1, EXD2, DJC11, FANCI, ATD3A, ARL8B, ANM7, FAIM1, RT18A, ARMC1, RPC5, SDA1, RBM28, RBM22, DMAC2, WDR70, PK1IP, RM22, CZIB, BABA1, NDUBB, RM16, NHP2, OCAD1, ADPRS, LYAR, MIC19, THUM1, TRM1, PP4R2, DPP3, BCLF1, FKB11, TMOD3, ELOV5, GTSE1, TECR, ERAP1, ARP10, CISD1, F120A, IF2B1, COQ3, HPBP1, MAT2B, CNOT2, OGFR, CNIH4, RM15, THYN1, KCMF1, TM14C, HM20B, RCC2, RELCH, BCCIP, STK26, FPRP, IBTK, RRBP1, SYLC, SUCB1, DPM3, ATX10, TFP11, SAE1, COMD3, MTRR, ASH2L, DHCR7, EIF3K, SAE2, WNT16, CSN7A, QCR9, ZO2, LRWD1, ABCF2, SYWM, SEC63, SRP68, CHRD1, BAP29, S30BP, PFD2, GPN3, PUF60, NRBP, ENOPH, NDUAC, EI2BD, VATH, XPO7, VPS51, S2513, DCTN4, FBX2, CCNL1, DBR1, FEM1B, DJC12, MRT4, CPSF3, NUDT5, RCOR1, TF3C4, PACN3, ACINU, AGO2, PSME2, PALD, COR1C, AKP8L, ALK, DD19B, PRP19, UBQL1, SNX12, CHIP, PACN2, MAGD2, SNX6, SSRG, FAF1, PPIE, MINP1, SCAF8, TRI33, PHF8, LIMC1, UBP24, SMC3, SCML2, PIN4, PSF2, EIF3L, PLAP, RUVB1, BCS1, SYFA, CNPY2, PKHA6, RRP44, RT07, PDIP2, WBP11, NOP58, SGT1, ACOT9, AAR2, MEMO1, TMX2, LC7L2, DHRS7, RT02, SBDS, TMED5, EXOS1, SF3B6, RM11, RRP15, NOP16, TPRKB, PPIL1, CIA2B, RT16, RT23, PTH2, RTCB, FBX7, NOC2L, RL36, SAMH1, DCAF1, UBP15, TBL2, TELO2, LAS1L, AFG32, ARI1, SAM50, PRC2C, PPME1, AP4S1, YTHD2, CTDP1, SP16H, UTP18, TNPO3, GMPPB, RBM8A, SNX8, FHOD1, SERC, NCOR2, ORNT1, ALG5, RT18B, CLIC4, NFS1, SPCS1, MD1L1, STK24, 5MP1, COMDA, DC1L1, TX264, S4A7, NDUB9, MORC2, S23IP
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, RIMC1, 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, 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, HSPA9, TXLNA, STAT3, PEX19, RFC5, RFC3, IF2G, NAA10, KDM5C, CSK, GARS, SYIC, LAP2A, LAP2B, 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, 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, SKI2, 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, ZNG1F, 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, AFG2A, NHLC2, ATAD1, SERB1, 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, PDRG1, STAU2, LIN7C, TXLNG, MRGBP, SEP11, TBC13, ASF1B, KTU, PNPO, NECP2, BSDC1, MED9, F118A, CZIB, BABA1, UCKL1, NHP2, OCAD1, LYAR, MED29, THUM1, TRM1, NDE1, CARF, PP4R2, AATF, TE2IP, HXC10, BCLF1, TMOD3, MS18A, GTSE1, UBAP1, GGA3, F120A, UBIP1, COQ3, HPBP1, ITSN2, CNOT2, OGFR, PALS2, CHMP5, NDUF4, RAI14, RPA12, PHRF1, VPS18, RCC2, RELCH, JCAD, SLAI2, BCCIP, STK26, CT2NL, CPSF2, SYLC, KANL3, ATX10, NCDN, SAE1, RBX2, PI4KB, ASH2L, HDAC6, EIF3K, RNF14, SAE2, PEF1, CSN7A, MALT1, DAXX, STK39, CGBP1, TES, SRGEF, LIMD1, NDRG3, LIMA1, SEPT9, UBQL2, DDX20, BI2L1, S30BP, GPN3, NRBP, VATH, IPO11, TR112, SIX4, DBNL, GGA2, GGA1, STML2, DBR1, AKA11, CPSF3, DP13A, RCOR1, TF3C4, ACINU, NUP50, ZHX1, BAG5, RAB21, ZN346, MCTS1, MRTFB, YETS2, HECD1, ZMYD8, CIZ1, COR1C, TPX2, DD19B, NFU1, PRP19, UBQL1, SNX12, SYNRG, VPS4A, CHIP, SNX6, PSD13, FAF1, CSN3, SCAF8, PP6R1, ANR26, UBP22, UBP24, DICER, HPS5, SRRM2, SCML2, EIF3L, PLAP, RUVB1, NUDC, VDAC3, ASF1A, DRG1, NCKP1, ICE1, R3HD2, CHSP1, NOP58, ZN281, SGT1, AAR2, NOSIP, SHLB1, LC7L2, SF3B6, ORN, RRP15, NOP16, UFC1, HDGR3, STRAP, RBGP1, ZN330, R3HC1, CCDC9, SAMH1, SALL2, ARIP4, KDM3A, RBM19, PLXD1, WAC2C, UBP15, IRS2, ATG4B, WIPI2, LAS1L, ARI1, PRC2C, PPME1, SP16H, CD2AP, SNX5, COPG1, TACC3, STK24, DC1L1, NEMO, NCOA3, M3K4, WASF2, SCC4, ZHX2, S23IP
Species: Homo sapiens
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