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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, LRTM3, 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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Boulard M, Rucli S, Edwards JR, Bestor TH. Methylation-directed glycosylation of chromatin factors represses retrotransposon promoters. Proceedings of the National Academy of Sciences of the United States of America 2020 117(25) 32522876
Abstract:
The mechanisms by which methylated mammalian promoters are transcriptionally silenced even in the presence of all of the factors required for their expression have long been a major unresolved issue in the field of epigenetics. Repression requires the assembly of a methylation-dependent silencing complex that contains the TRIM28 protein (also known as KAP1 and TIF1β), a scaffolding protein without intrinsic repressive or DNA-binding properties. The identity of the key effector within this complex that represses transcription is unknown. We developed a methylation-sensitized interaction screen which revealed that TRIM28 was complexed with O-linked β-N-acetylglucosamine transferase (OGT) only in cells that had normal genomic methylation patterns. OGT is the only glycosyltransferase that modifies cytoplasmic and nuclear protein by transfer of N-acetylglucosamine (O-GlcNAc) to serine and threonine hydroxyls. Whole-genome analysis showed that O-glycosylated proteins and TRIM28 were specifically bound to promoters of active retrotransposons and to imprinting control regions, the two major regulatory sequences controlled by DNA methylation. Furthermore, genome-wide loss of DNA methylation caused a loss of O-GlcNAc from multiple transcriptional repressor proteins associated with TRIM28. A newly developed Cas9-based editing method for targeted removal of O-GlcNAc was directed against retrotransposon promoters. Local chromatin de-GlcNAcylation specifically reactivated the expression of the targeted retrotransposon family without loss of DNA methylation. These data revealed that O-linked glycosylation of chromatin factors is essential for the transcriptional repression of methylated retrotransposons.
O-GlcNAc proteins:
DX39A, MPP10, NOP56, TCRG1, SR140, SNUT1, PRPF3, SMCA5, KDM1A, PR40A, SF3B1, U520, RL1D1, H2A1B, HNRPC, VIME, PARP1, H2AZ, TOP1, TOP2A, H2AX, H15, UBF1, DDX5, RL7, NUCL, LMNB1, FBRL, RS3, DNMT1, U2AF2, RL13, SMCA1, RFC1, DEK, RL4, RBMX, IF4A3, MSH2, CBX5, NOP2, RS9, CTCF, RBM25, SMCA4, HNRPM, MSH6, NUP98, RS3A, RS15A, RS13, RL7A, RS4X, RL23A, RS6, H4, H2B1C, RL8, TRA2B, RACK1, H31, H33, HNRPU, H11, RL18, SRSF1, DHX9, SSRP1, TBL3, CHD3, ILF3, REST, TIF1B, TCOF, SF3B2, HDAC1, NOG2, RBM39, BMS1, CHD4, CUL7, U5S1, SF3B3, PUM3, QSER1, LIN41, SMU1, TDIF2, LITD1, RIF1, TBA1A, EMSY, WDR75, FBXW8, WDR36, DDX17, SNR40, DDX27, ZFR, NACC1, SIN3A, CDC5L, MBB1A, DDX23, UT14A, GTPB4, SENP3, SMRCD, MOV10, DDX21, HELLS, DDX18, DNM3B, ZMYM2, DNM3L, RALY, PRP19, SCML2, DDX52, TR150, NOP58, LAS1L, SP16H, DNM3A
Species: Homo sapiens
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Huo B, Zhang W, Zhao X, Dong H, Yu Y, Wang J, Qian X, Qin W. A triarylphosphine-trimethylpiperidine reagent for the one-step derivatization and enrichment of protein post-translational modifications and identification by mass spectrometry. Chemical communications (Cambridge, England) 2018 54(98) 30379171
Abstract:
We report a new reagent that is capable of both chemical derivatization and selective enrichment of azide-labeled PTM peptides for sensitive identification by mass spectrometry (MS). Facile sample recovery, enhanced ionization and fragmentation in MS of the enriched PTM peptides are achieved, which leads to the identification of 3293 O-GlcNAc peptides and the location of 1706 sites in HeLa cells and efficiently expands the current mapping scale.
O-GlcNAc proteins:
MEIKN, BLT3B, SHOT1, TTC24, PIPSL, HFM1, ZN320, NBAS, CQ102, VWA8, SBNO1, Z804B, CC14C, SH321, EFCB5, XIRP2, ODAD3, CNOT1, BDP1, ANR62, CC88B, AN33B, NKX26, PGP, F186A, ARRD5, FBP12, ANKUB, CI092, CE052, FA47D, SMHD1, LRIQ4, GLOD5, ZN233, SH2D7, MARHB, TTLL8, ABTB3, A30BL, ZSWM8, P121C, ERI2, PP2D1, CEA18, R212B, YV023, ZN727, VGLL3, YD021, RUXGL, IQAK1, MAGBH, FOXO6, FHAD1, SRRM5, ZN879, S14L6, CJ142, KRBX1, MCRI1, FONG, CE067, F227A, CRCC2, KIF2A, MYO1C, EYA2, AP3B1, SPT5H, TAF4, HIP1, CDC7, ST1C2, WASL, IPO5, EMAL1, P3C2A, EXOC5, PSDE, USP9Y, BCL9, CAC1A, PITM1, MPP10, TRI38, NDKM, ARI1A, NCKP5, FCHO1, SOCS3, TRAD1, RHG33, TYB4Y, UTY, ENC1, ZN197, APAF, TERT, CHK1, GNB5, TCRG1, UN13B, RASM, PPE1, IFIT3, IKKB, TIM23, HGS, AURKA, MYPT1, XPO1, IPP2C, ZN646, PDZD2, SPTN2, SET1A, TRNK1, PER2, SYNJ2, ZN536, IKKA, TBX3, ARPC2, TBXT, VILL, NKRF, ZN185, CASC3, CLIC2, ACOX3, NPHP1, OGT1, PPM1D, RA51B, SHIP2, EIF3D, IPO8, MSH4, MAGB4, ARPC5, PER1, ZSC9, ZZEF1, TET3, SI1L1, PRP4, SERA, IRAK2, PSMD3, TEX33, AP5Z1, MTSS1, PAX4, WDR62, PRPF3, SYNJ1, IF4G3, KCAB3, E41L2, FOXO3, TGM5, SYT7, PPIP1, CHM2A, PLRG1, LDB2, BUB1, HTSF1, STX6, ENSA, CPSF5, CREB3, AK1BA, KALRN, MYPT2, MED14, SOX15, KLK8, PRKN, SMCA5, ZC3H1, TYW4, LZTS3, GANP, KIF1B, PHLP1, ZN861, BRNP1, MPZL2, OGA, VINEX, HNRPQ, RNBP6, GMDS, REV3L, JAK2, MAFK, MUSC, CTND1, HPGDS, Z354A, RT14, PQBP1, BRD4, ABCB7, PLCH2, SRGP2, DEN4B, N4BP1, ROCK2, COBL, CPNE3, ZBED4, LIPA3, OBSL1, BRE1B, PHF2, ZC11A, CLU, DJC13, ANR17, SIN3B, LIPA2, LIPA4, ZN253, ZN217, FLNB, NCOR1, NDUS6, RM33, DPOLQ, CDC45, MPPB, USH2A, SRS10, GEMI, ECI2, TAF5L, ZN821, DAB1, MYCB2, U520, OFD1, UTP20, NU155, S2512, PALM, ZN189, CCNK, SERF1, PIAS1, DNJC8, RAD17, ANXA9, CLPX, SEGN, CDKL5, NEBL, S14L2, RECQ4, RECQ5, RMP, AL1A2, TOM70, TOX4, MICA2, K0754, FCSD2, SRBS2, SASH1, CSCL1, LRIG2, SUN1, PRP6, PCF11, KDM4B, CE152, SOX30, UBR5, REC8, DUS14, NDUB8, KLHL2, LETM1, MBD4, KAT7, KCNH1, SMC2, IPO7, SLU7, SVIL, NPT2B, LCA5L, FMNL1, PCNT, CNOT4, CPSF4, NFAC1, EYA4, BRD1, HERC2, HS74L, RIGI, ZMYM6, BAG3, DCMC, DDAH2, TGM3L, TBX18, TBX6, BCL10, APBA3, NSD2, GSHR, CRYAB, LMNA, SPTA1, ESR1, ANGI, TDT, CYTB, GCR, SODM, MYCN, G3P, P53, HSPB1, ARGI1, PCCB, RLA2, RLA0, RPC4, K1C18, FYN, NFL, NFM, PERT, CRGC, SYEP, ANXA6, RHOC, GLI1, PFKAM, CATG, INHBA, VIME, ANXA5, SNRPA, VILI, CN37, HMOX1, DLDH, HXB7, DPOLA, IFIT2, WEE2, FBSP1, BNIP5, AN34C, ACSM4, PRR29, F90AO, YT014, R10B1, GRL1A, F200B, DERPC, CTF8, ZN888, RGPD2, GIMD1, ZN728, CH088, AT8OS, MYBA, ARAF, ODP2, KAP0, HEM4, HSP7C, SPTB1, TOP2A, VDR, DMD, ADHX, NFH, TPR, ANXA3, SKIL, NR2C1, HXD8, ENOB, T2FB, MTDC, 3BHS1, GFAP, KPYM, HXB1, PO2F1, ERF3A, EZRI, PHKG2, DESP, ANK1, NCPR, CBPE, PLCG2, ZFX, ZNF14, ZN708, ZNF17, ZNF19, ZNF20, ZKSC1, ZNF28, ZNF30, ZSC20, ZNF7, KPCA, UBF1, ATF7, DDX5, LOX12, EGR1, ARY1, SON, ATF1, ATF6A, NUCL, HXK1, TNNI3, HME2, SL9A1, RAB3B, LMNB1, GRAH, NEBU, AOFA, ZNF10, TGM2, MUTA, PERL, ROA2, CBL, KAPCB, HEMH, XPA, CRCM, SAHH, IF4B, PAX7, GATA2, CPT2, RIR1, EF1B, CCNC, DNJB2, PSA1, MOES, DDX6, RAE2, PTBP1, SYVC, 1433T, ARNT, RFA1, MAP4, PSB9, TMOD1, ITPI2, HXD9, ERCC5, EPHB2, PTN6, SHC1, KDM5A, LMOD1, TYK2, PRDX6, CDC27, WEE1, MPIP1, MPIP2, NKTR, CLIP1, RPB2, KAP1, CPSM, ZEP2, CASPE, 1433B, STIP1, FOXN2, TF2H1, C2TA, LSP1, DUT, GCYA2, MCM7, HYES, MPI, CTNA1, NOS2, SPB6, MERL, RFC4, RFC1, ADML, RORA, CBS, GDE, ACTN2, ADDB, NU214, DEK, CYLC1, ATP7B, SOX5, SOX11, ZNF93, VATE1, 8ODP, LONM, PGM1, GNL1, PEDF, ODO2, SRP14, IF4A3, SMBP2, CUX1, TYRP2, MLH1, STAT3, UBP8, PEX19, ECHA, ETV6, STAT6, AF9, HELZ, LPPRC, HD, ECE1, MUC18, BTD, MAGB1, KSYK, UBP5, KI67, KPB2, KPB1, RECQ1, IF2M, BRCC3, RL28, MAP1B, NEDD4, UTRN, CAPZB, CDX1, RL29, UCRI, PPCE, RFX5, PI42A, RXRG, NRIP1, PRC2A, CCN3, AL9A1, NASP, RM19, CENPF, MCM2, YLPM1, VPS41, NUMB, CLK1, NU153, RBP2, RGS3, RGS7, TSC2, TAF6, DNLI3, ETV1, DYN2, TUB, TCPD, PAPOA, PLCD1, ZNF80, ZNF81, ZNF84, BRCA2, RENBP, MECP2, SSDH, AFF2, AFF3, MYOM1, HNRPM, NCBP2, MP2K6, GDIR2, AGFG1, ZN135, ZN142, ZN143, RBM5, SPSY, HXK2, DGKQ, THOP1, TXTP, PPP5, LIMK2, SMTN, DPOG1, ICLN, BLM, PDE1A, GALC, KAD2, IF5, NR1H2, AFAD, AF17, ADK, DSRAD, UCP3, DLX1, SYMC, ATP5I, HDAC4, PEX3, PER3, RP1, SIK1, PRD15, UBS3A, NAC3, NU107, SNX16, RGS8, FBXW4, SESN2, TIRAP, H2B1D, CU082, MCCD1, EIF3E, PTEN, IF4A1, SNP25, DEST, ARP2, ISL1, REC21, STXB1, 1433G, UBP46, TIM10, PRS4, PRS10, ERF1, GAK8, RS4X, RBX1, TRA2B, ACTG, F193A, NOP14, RBM6, GTF2I, DLG4, IRX6, GSTO1, RELN, PRKDC, ELF3, MAP1A, SIM1, SSBP2, TMF1, RT05, RT11, RT09, HMGN5, SKOR1, FOXK1, PRR5, DAB2, EFNB1, RBM10, RBM3, VIGLN, BORG5, FANCC, CLH1, HSF1, MYPC1, U2AF1, SPTB2, TIAR, SET, FOXK2, RELB, AMPD2, ANK2, TFAP4, XPC, OCRL, AK17A, MEF2B, SP2, CENPE, KIF23, MMSA, ID2, ZNF45, APBA1, HEN1, FKBP4, RL6, TOP2B, DYST, KMT2A, CENPC, ERCC6, TAP1, ZN117, ARI5A, IF4G1, TLE4, UBE3A, DYN1, SRS11, CALD1, RN5A, EMAL5, BTK, EXOS9, MTG8, GABPA, CDK18, ZO1, TRHY, PDE4B, SPAG1, KHDR1, DPOE1, SRSF4, GOGA3, PDE4C, DEMA, PDE4D, RBL2, ZN844, ACSM1, DMWD, EP300, AHNK, ZNFS1, FCHO2, PLCX2, DYH14, K2012, CGNL1, CF210, FP100, MESP2, NEXN, CPSF1, MPPA, SCRN1, TF3C1, HYAL1, BPTF, SFSWA, SF3A3, DPYD, TP53B, IFRD2, ILF3, EPS8, FOXC1, ANK3, GNDS, SEC20, NFX1, TIAM1, AF1Q, AKAP6, PRDM2, CAF1A, TRAF3, AIMP2, CBX3, DDX10, TBX2, ERCC8, GPS2, SBP1, NMI, PABP4, CD5R2, PPIL2, MB211, PM2P3, ORC1, ORC2, SNTB2, PPIG, FKBP5, MYO9B, GAB1, BIRC2, PICAL, SQSTM, BFSP2, PRP4K, ATR, SNW1, IQGA2, MTMR1, CUL4B, ASPP2, DYR1A, PTC1, RAB32, FHL1, FHL3, PM2PB, CAC1S, THOC5, NCOA4, SPTN1, KCNC3, CDK13, CKAP5, CYLC2, DAG1, SRTD2, MORC3, SAFB2, UBP2L, SCRIB, DOC2B, DOCK1, BTG3, ZMYM3, DYHC1, EI2BA, SRC8, FAK2, FLNC, FA50A, GRB14, HES1, NF2L1, FAT1, DHX8, MCM6, ITPR3, ZN266, ZN235, ZN460, PLSI, PUM1, KANK1, MELK, SMC1A, RRP1B, NCOA6, GSE1, PSF1, BMS1, LBR, STAT4, CBX2, KIF22, MEF2D, MTF1, MYPC3, NC2A, NFIX, ZN638, NOLC1, NR1D2, PSME4, CUL7, SLMAP, GAPD1, FRPD4, CND2, MO4L2, SUZ12, R3HD1, SNX17, S39AE, LRC14, DPOD3, IF4H, KIF14, EEA1, CD5R1, NCF4, PLCL1, PDK1, PLCB4, PLEC, PTPRR, PWP2, IRF4, ZMY11, L2GL1, UBE3C, PUM3, DLGP5, RBY1F, KS6A1, SAFB1, SYCP1, PP1R7, SP17, MED22, TAF1C, NCOA2, TRIPB, PCH2, MED1, JHD2C, ZN174, DLG2, CEBPE, TAB1, STXB2, ZFHX3, ZBTB6, ZYX, SEPT7, ADRM1, AINX, PSMD5, PKN2, TAF12, CDC37, B2LA1, TAF9, MAPK3, TBR1, CP1B1, DGKD, THTR, PCKGM, ZN827, LONF1, EX3L4, TPRXL, GEN, HNRL2, LONF2, SAMTR, PDS5A, MOONR, QSER1, WASC4, EX3L2, TECT1, ERC6L, RGPA2, D19L1, ZXDC, CCD57, PWP3A, ATG2A, FHOD3, IF2GL, KLH38, HFE, BTBDG, LRRF1, RFX4, LRRK1, F161A, INAVA, MA7D1, PKHG6, CHD9, TBC25, C144B, OD3L2, ZN718, AL1L2, P12L1, PTHB1, GON4L, GRDN, GVQW3, PAR14, CP100, ZN404, CL036, SPAS1, SV2C, YJ016, ZFP69, CE022, C2CD3, KMT5B, LARP7, HYDIN, KCD21, IQCC, PLCH1, FIL1L, ARI4B, GRAP1, Z658B, PAP1L, TIGD2, FNDC1, VWA3B, PAN2, CN177, Z585B, SBK1, UBP39, TYW2, FAKD1, CTSRT, SGO2, CC178, ODFP2, CAVN4, FSIP2, UROL1, ZN628, TRI61, ESCO1, ZN789, BCORL, ZMAT1, PP6R3, DCA17, FSCB, BRD10, ZC12B, NHSL2, ZN831, AKAP4, MTUS2, FA47A, MEIG1, PRC2B, RRP12, IQEC2, SPIN3, ZN506, DOP1, F209B, GASP1, DGKK, CYTSB, CD158, SPXN4, FBX47, TCHL1, TDIF2, ERIC3, LRRK2, SAPC1, CE170, PR15B, CJ055, SNPC4, CTSRE, NHSL1, F120S, C2D1B, TEX35, FA83B, SYRM, AXDN1, FKB15, AKND1, ZC3HD, MPP7, UBR4, ZN684, SWT1, SKT, CC183, RHG21, BEND3, UBAP2, PHF19, LITD1, F27D1, TSTD2, ER6L2, PPR26, ACBD5, UT14C, TUT4, DEP1A, CE162, FRY, ZBT37, MAGI3, KAD9, DDI2, MEI1, CC181, LIN9, ZN658, CROCC, CBPC2, MCAF2, RIF1, ZN468, SPXN1, OBSCN, NTM2D, CE350, RPRD2, SYDE2, KLD7A, BRE1A, MYOME, P210L, ZN318, A20A3, ZN691, PPRC1, SPO16, MYSM1, NTM1B, TAF3, SNX30, TASO2, KHDR2, ECM29, NAA35, MBNL2, OBI1, USPL1, F219B, RN123, FBX31, FBW10, ZKSC2, RN213, WDR25, CP135, Z385C, AR6P4, ARID2, TENS3, SPT2, FMN1, ANKS6, USF3, MSL1, S31H1, CRBG3, FRPD2, RHG17, DUS29, C19L1, VIR, CYTSA, ZN660, REP15, BLT3A, SP5, CARL2, K0930, JADE1, SNR48, CAPR2, SPOPL, CCSAP, PKHA7, ADNP2, IF5AL, GRHL2, ZN425, ACD10, IFIX, NIPBL, LIN54, R7BP, TET2, TEN4, Z280D, NAA16, RADX, SANBR, ZSC25, SCMC1, ACSM5, JHY, CRCDL, SWAP1, ZN774, ZN544, JMJD6, CAVN1, FXL22, M18BP, CDC73, T2H2L, XRRA1, TTC41, NEK5, NFRKB, LMOD2, RL22L, PKN3, ERIC5, ANR16, FXL19, ZN880, TM10B, KLH35, PSRC1, BORA, IYD1, KCD18, GASP3, FBX38, FOXR1, ARMD1, BRAT1, ZC3HE, LARP1, ATAD2, SPG17, RICTR, BRWD3, TSSK4, F111B, SAMD1, SARM1, KLH17, PADI6, C1TM, ANR12, ANR11, FIP1, SAS6, UD2A3, CWC27, LRIG3, MIC27, BI2L2, MCAF1, BCOR, MPRIP, ZY11A, NSMF, RBMS3, RPTN, DNMBP, GGYF2, CCNI2, ZN782, EMAL6, SYNE3, MASTR, CU136, MEX3B, ZNF66, ZN793, LN28B, CCD81, RN111, ZN704, BICL1, CC141, WDR87, ZN493, CO039, CCD73, SRCAP, ZN662, NBEL1, YD022, SHSA6, YO027, YJ005, TSH1, CBAR2, CG065, MSD1, K0408, CFA65, UBN2, BEND4, F205A, RHG27, FRMD7, FOXNB, RFX8, FGD6, ANKS3, YS043, VP13C, UBP34, UBP31, UBP53, UBP43, ZN365, CEFIP, CENPU, SSH2, CEP68, SPT6H, RUFY3, ERIP6, DHX30, ZN765, MCM10, DOK3, GDPD3, RFIP2, FAKD5, OTU6A, ZN627, KDM3B, STRAA, MICA3, OTOP3, SSX7, U17L1, H2BWT, ZCCHV, GVIN1, TICRR, EFL1, PBIR2, SETX, KANL1, NUP54, PRIC2, ZN572, RGPD4, POGZ, ZFY16, MYCPP, TMC6, NUFP2, MAVS, CLAP1, DHX29, LIMS2, TT21B, HEAT3, CFA91, ZN438, HDGR2, EMSY, AKNA, RAI1, ANKAR, I2BP2, AB12B, SNX20, SPRE1, RBBP6, ANS1B, UBE3D, PTAR1, TCPR1, RABEK, HUWE1, YTHD3, FLIP1, TAF8, KHDC4, VP13B, S1A7A, PHLB2, CROL1, KAISO, ZN605, UBP37, K1328, RN180, PATL1, MYPN, PEG10, ATG4D, TTC7B, ISCA2, PABP2, LDB1, PB1, RIP, MARH3, NFE4, RIMS1, CATAC, NT5D3, CASZ1, VASH2, LUZP1, MORC1, MYPOP, NRAP, IQGA3, ZCH18, PACS2, MON1A, TSYL5, ZSC30, PARG, LIPB1, LONP2, RASF3, NLRC5, CHD1L, SKAP1, AROS, CDR2L, DAA11, EZHIP, CQ082, CIR1, HTR5A, ZFAN4, SNX32, HOGA1, DRC11, ZN761, GOLI, TRI50, Z354C, CD20B, KMT5C, ZN573, PRGC2, P66A, RASL3, MD12L, I2BP1, RB6I2, RIOX2, MY18B, RBM45, SIAH1, RELL1, LACC1, HID1, ANKY2, PSTK, VRK3, CROL2, PKHH2, AHNK2, TIAM2, AN18A, FMNL3, SAM9L, NAV3, NAV2, CCD50, FABD, MISP, HERC6, WDR75, MFN1, CCD60, TEX14, MA7D3, NXP20, CCD83, DNAI3, MGAP, GCC2, RP1L1, BRSK2, Z3H7A, TPH2, UBR2, ANKH1, SUGP1, SUGP2, KIBRA, GOR, CCAR1, SLF2, RM41, GPAM1, KLHL7, ETKMT, DDIAS, RB12B, ZC3H3, SMC5, DHX37, KBTB2, SPB1, NIM1, C144C, CKP2L, IHO1, SUV3, ERF3B, FANCM, ZZZ3, EXOC8, CF206, P20D2, DAAF8, CFA46, SAXO1, SACDR, RTKN2, SAM14, KMT2E, DOCK3, ABI1, ASPM, F185A, CA087, IQCK, SPG16, SPART, SPICE, TTC5, TAGAP, RPTOR, EID3, AHI1, ZN567, ZN687, REXO1, S6OS1, CRBG2, THEM5, ANR35, CLASR, AN36B, RUSC2, CF223, RHG18, PARP8, EH1L1, METL4, CMYA5, VPS8, PALS1, SYNPO, FNBP4, RDHE2, PRS35, PGBD5, NIN, SPA6L, SPEM1, TT30B, LRC71, AF1L2, CERS5, SRBD1, TAB3, MACOI, MINY1, CCD33, TM148, ESX1, PACRL, TRIMM, TM217, ZN676, NANG2, ANR31, CNTD1, TEANC, ZN615, K1958, ENAH, CDYL2, CO056, ZN709, JMY, ZN454, ZN565, YV006, SR1IP, CDRT4, TAC2N, GAR2, FSIP1, ZN383, PNDC1, FA47B, MARHA, KBTB3, CI106, CAST3, ZBT38, CH058, SMYD1, UN13C, KY, CK072, ZN114, SERB1, HJURP, SEN2, GLBL3, MACA2, SMG8, RN214, Z280C, MROH1, PAPD5, LRTM3, ICA1L, EHBP1, CBPC5, RB15B, SMC1B, PHC3, BEND2, SIMC1, NSUN7, SRFB1, DDHD1, SP20H, CBPC3, ARMC2, PDZD8, ZN555, RTL9, DP13B, NAV1, WDR19, KMT2C, FBX22, SYNE1, BD1L1, FAD1, FLCN, PXT1, DBF4B, IPMK, ABHDB, DMBX1, FBH1, PJA1, KNL1, NEK11, KDM2B, ARK74, CLHC1, CFA61, NEDD1, RLA0L, ZFP28, THOC2, INADL, KI18A, ZNT5, SNIP1, WDR48, UHMK1, NEIL3, FA76A, ZN671, PRR7, PUM2, KLDC4, NANP, AT2L1, RBM46, PMFBP, CC110, ZN502, TBC15, ZN596, GAR3, FANK1, ZN507, PHOP1, ALMS1, DC2L1, CHD6, HUMMR, SP7, ACTT1, HELQ, DMXL2, DLG5, MIPT3, OTU7A, S26A7, ATS17, S3TC1, TBCK, PO210, CE192, GEMI5, FBF1, RPGF6, SMCR8, PARD3, PNISR, ZN483, DI3L1, ZN526, GIPC2, CP078, DUS19, THAP3, ZDHC1, TB22A, PDC6I, CO040, DCAF4, MITD1, TFB1M, TPC14, DC2I1, UBCP1, TRUB1, LMO7, CKAP2, CCNB3, ATX2L, PHIP, RT4I1, PALLD, CRGN, SYNE2, CNKR2, ASB10, ACO11, P66B, ASB17, BBX, CTBL1, ELYS, TITIN, TRIB2, SMG7, RTF1, IP6K1, PHYIP, DCNL4, P55G, NR4A3, PHF3, NDRG1, HS105, ZN592, PRP16, ANS1A, FAN, AKAP1, ZN75C, CENPI, PRCC, RREB1, HDAC2, DPF1, DPF3, REQU, CBP, SYMPK, HYAS2, KAT2A, RPGR, DDX17, KIFA3, NEO1, RAD50, RAB8B, ARHG2, IRF7, TF3B, DVL3, USP9X, CUL5, LPP, TP4A1, HGD, SMCE1, FBXW7, FBX32, TOP1M, TRI63, LMBL2, ZN622, 5NT3B, FBXW5, PSMG2, TEKT1, FAKD4, TET5B, C102A, RM24, SPSB4, ZUP1, CC74A, TTC28, R51A1, DACT3, SH3K1, PF21A, ELMO3, PINX1, CC049, S1PBP, ZN775, DOC10, ATG2B, ZBTB9, ROP1L, COAC, CHM4C, CC124, OPTN, S7A6O, GASP2, RMD1, OTUB2, ATG4C, DRC10, MTBP, RRFM, MTEF3, ELP4, HOOK2, MTMR8, ZN837, RL39L, MITOK, SGF29, RBM33, DTBP1, ZN503, ELMD3, THOC1, HMCES, IMP4, KLD7B, SCMH1, TOE1, MSD3, ZCCHL, TONSL, NTAQ1, PDLI5, PRR11, FMC1, C2AIL, INT4, SYNM, REXO5, GMPPA, GMCL1, ZN496, NGLY1, RBM41, TPD55, ESIP1, MAP6, ZN594, ZN469, ZN462, FEM1C, PGBD1, ZN521, TOPK, PRAC1, COX42, KCNS1, Z512B, ZFR, F167A, P12L2, CD036, NSD1, EP400, IFT74, SGCZ, CX058, CEP19, CH048, CL037, MRFL, Z354B, PRIPO, RMD2, CJ090, IFAS1, TEKT5, TEX55, FEAS2, CCDC7, CFA53, ZN512, CQ10A, NSE2, CE034, RIPR3, TEAN2, ZSWM3, YTDC1, ZNF48, F161B, F218A, TTC14, CK037, ZN578, PWP2A, DOCK7, ZN641, ZN582, ZFP3, LRRC7, BIRC8, INP4A, TSSK2, RBM14, ADCYA, IPP2L, ZN317, PNMA5, K1908, FMNL2, PUS7, F111A, PLIN4, SMG1, ALKB3, KI20B, RHG07, ATAD5, TEFM, RAB3I, GCNA, PRAM, MAGI1, RET7, PANX2, MED15, ERBIN, TRIB1, HMCN1, CRML, PRR25, LYSM1, CCNL2, SMC6, ZN607, ZN587, IWS1, SEBP2, RSF1, RUFY1, IMP2L, MINT, SETB2, ACOX2, AGAP2, EYA3, DNJC2, MPP9, M3K14, S10AD, SCAFB, ELF4, KLF6, PHB2, CHP1, KIF2C, LYST, ATX2, SMAD5, BARD1, NPAS1, EPAS1, TS101, CPNE1, SMO, EBP2, PKP2, AKAP9, ARP5L, SIN1, RNZ2, CMS1, SENP7, SLF1, COE4, ST4A1, ZSWM1, COR1B, LSMD1, SRSF8, PYGO2, FSP1, GPTC1, LLPH, SIKE1, TRI56, ZPBP1, BTBDA, TACO1, PIMRE, S2533, HAUS8, ALKB7, DIDO1, MTA3, DCNL5, RAMAC, FAP24, LRRC1, MED18, RCAF1, SCMC3, THUM3, CP250, KTNB1, PHF20, RUSC1, RIOK2, TIM21, KIFC1, HIRP3, TRAIP, FUND2, CYREN, CDCA7, MND1, CAR10, NADAP, GATA5, SYCP2, FANCJ, SRBS1, CARD6, ZN471, OSB10, RFIP5, SPZ1, CAR14, CAR11, T121B, TGT, TDRD1, TEX15, RNF17, STK31, UBP26, OSBL1, FACD2, AN30A, MAK16, SCAPE, DGC6L, RTF2, PECR, LRRC3, RNF26, SCOT2, RM01, ACE2, MSTRO, HYCCI, YTHD1, LRRC2, NEUL, PBX4, BCDO2, CEP41, SETD2, TBD2A, LDH6B, PANK2, NSD3, NUF2, PUS3, BARH1, OSBL7, UACA, REN3B, NIBA1, UBXN6, TRI15, TRIM4, F117A, SPEF2, LRCC1, ZFHX2, CFA74, AMRA1, CE295, ZY11B, TANC1, XPO4, GRIP2, ASXL3, CEP44, ZN407, SRCN1, CCSE1, WDR33, TGIF2, ROGDI, REXO4, NXF2, MFF, DRC3, CLPB, LSG1, NUAK2, NAT10, IQCN, ILKAP, SP130, UBP44, BRD8, RGAP1, PUS7L, ZCPW1, GZF1, BC11A, I2BPL, ISCU, RSPH9, VPS11, AL8A1, TARA, EPC1, CI037, PDCL3, ADNP, TNMD, RBP17, IPYR2, MIXL1, HIPK2, ZN106, E41LB, UBQL3, YI012, MOD5, GHITM, BOLA2, BORG4, UN45A, DSN1, CA198, RBSK, FN3K, WNK1, PLK3, E41L1, EMRE, PRD12, ZN644, VIAAT, THAP9, BICC1, CCD86, MYH16, CIAO3, ACSS3, BCAS3, ZN552, ASB8, SH24A, SMYD3, MB12B, ZN665, CCSE2, ZN430, NOL11, PAAT, MINY3, ECT2, F204A, CF208, ANKZ1, GHC1, PPIP2, MARH7, CNO10, SPE39, ANRA2, UNKL, RPF1, RPA2, SYCM, NMNA1, MLXIP, RENT2, BRF2, CLSPN, BCDO1, PRD16, PKHA1, AS3MT, TENS1, NMRL1, SPC25, TAF9B, Z286A, TPC, MRM3, GPBL1, CENPJ, ZBT20, HES4, NCOA5, TANC2, ZN532, SYTL2, MAGE1, K1586, ZSWM6, ZDBF2, CHD8, GPAT1, FBSL, ZN304, NDF4, HMX1, S40A1, GCM1, VTA1, EXOS4, UBN1, MKKS, MED4, SCRT2, CK016, PLCB1, A1CF, DDX4, DMRT3, INCE, PRDM9, BIN3, ENTP7, BIRC6, PDLI7, ACSA, DYRK4, ANM8, ASH1L, KLHL1, TSH2, SMYD2, SNRK, TBG2, STRN4, ZN277, LATS2, C42S1, PNO1, TOPRS, MBIP1, RAD18, HOME2, SALL1, ZN275, BRWD1, KLC4, AT8A2, DEPP1, CTSRZ, ZF64B, ECHD1, RBM12, ANKE1, MDN1, UFSP2, MITOS, OTULL, TYDP1, POTE1, DCA13, NUD15, RN19A, TYW1, EXOC1, UQCC1, MED17, TBC13, DJC11, INT7, FANCI, INT13, DZAN1, FAIM1, INT10, KTU, TBCC1, RPC2, RBM28, RBM22, BSDC1, WDR70, SLTM, SMOX, PIHD1, PARI, RMND1, BABA1, THG1, HPF1, IRAK4, TXN4B, NDUBB, MARH5, SYBU, MTMRA, TEX10, CNTLN, TRM1, CARF, NSE4A, DDX43, DPP3, ETAA1, ABI2, DYH9, HXC10, DACT1, FA53C, BCLF1, TAB2, CDK12, SMAL1, SACS, MAT2B, MYOF, ITSN2, ARHGC, CNOT2, VSX1, CWC15, RAI14, H2AW, RM27, VPS54, CAMP3, PHRF1, DAPLE, ZFAT, RELCH, DIP2B, JCAD, SLAI2, BAHC1, PAK5, CHD7, F135A, RRBP1, MAP10, CE126, CPSF2, KLHL9, SYLC, C2D2A, DISP3, TBC14, RBM27, KANL3, HECW2, STAR9, RERE, ANKY1, IMPCT, HNF6, HSFX1, RBX2, MYOTI, PRGC1, ARP21, GULP1, SAE2, POLK, DBF4A, NXF1, ZMYM2, REV1, ZNF70, S14L4, DAXX, ADDG, HYPDH, FXL17, CCD39, ABCF2, S23A2, AAKG2, KDM5B, SYWM, TCF20, SUN2, NDOR1, AFF4, SEPT9, UBQL2, IP6K2, NUFP1, SMPX, S30BP, MED13, PFD2, NRBP, FEZ2, EVL, CTNA3, XPO7, BAZ2B, BAZ2A, BAZ1B, HERC5, SIX4, ZMYM5, HACL1, STAG3, HOOK1, NGAP, ZN391, SALL4, S2513, FBXL7, ZN112, DBNL, ZN229, ZN180, HSPB8, KS6A6, DBR1, FBXL4, FBXL2, GMEB2, PO2F3, GPTC8, C8AP2, FBX4, FBXW2, IKZF3, TRHDE, HDAC9, ACINU, AGO2, NUP50, ZHX1, DSE, SARDH, CFA45, AGO1, ZN236, ZN214, MKLN1, NRX1A, PADI1, ZN777, INTU, ZN608, WWC3, F184B, INO80, KDIS, MRTFB, ZBT21, NEB1, AT7L1, K1210, PKHG1, SHRM4, PKHH1, YETS2, CNOT6, HECD1, ZMYD8, HSF4, PADI4, APC10, MYO6, SPAT2, KMT2B, CAN11, PHTF1, MAGD2, CSN3, TTF2, BSN, MACF1, SCAF8, TRI33, PHF8, LIMC1, TRI35, TNR6B, PLCL2, EXOC7, SMAG1, CE164, CBPC1, SATB2, UN13A, DICER, SRRM2, PA2G4, CD11A, CTND2, BAIP2, SCML2, RUVB2, CDYL, HIG1A, AKT3, PRLD1, PLAP, T53G5, ADIP, ICE1, EFR3B, PKHA6, MAST1, AMOL2, ZBTB1, KDM2A, FRM4B, FAN1, KLH20, CRYL1, AXIN2, GUAD, MAN1, TR150, MED16, ZBT32, MTO1, SYYM, ZBT12, CAB39, LC7L2, RBMX2, HDGR3, BOLA1, STA13, T22D4, RL36, C2CD2, ZN175, DMXL1, TRRAP, MTCL1, ZN451, WDR7, C170B, RIPR2, TLN2, DTNA, CRBG1, LAS1L, AFG32, PRC2C, PPME1, TRI17, CEP83, PAXB1, SP16H, MAGD1, KLF2, SNX5, LRRF2, FHOD1, IRAK3, NCOR2, GMEB1, BIG2, MD1L1, RM42, ATP23, SNCAP, CABIN, OAS3, NEMO, SQOR, NCOA3, NUMBL, Z780B, CCD61, SCIN, PCLO, SETBP, MORC2, NS1BP
Species: Homo sapiens
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Hahne H, Gholami AM, Kuster B. Discovery of O-GlcNAc-modified proteins in published large-scale proteome data. Molecular & cellular proteomics : MCP 2012 11(10) 22661428
Abstract:
The attachment of N-acetylglucosamine to serine or threonine residues (O-GlcNAc) is a post-translational modification on nuclear and cytoplasmic proteins with emerging roles in numerous cellular processes, such as signal transduction, transcription, and translation. It is further presumed that O-GlcNAc can exhibit a site-specific, dynamic and possibly functional interplay with phosphorylation. O-GlcNAc proteins are commonly identified by tandem mass spectrometry following some form of biochemical enrichment. In the present study, we assessed if, and to which extent, O-GlcNAc-modified proteins can be discovered from existing large-scale proteome data sets. To this end, we conceived a straightforward O-GlcNAc identification strategy based on our recently developed Oscore software that automatically analyzes tandem mass spectra for the presence and intensity of O-GlcNAc diagnostic fragment ions. Using the Oscore, we discovered hundreds of O-GlcNAc peptides not initially identified in these studies, and most of which have not been described before. Merely re-searching this data extended the number of known O-GlcNAc proteins by almost 100 suggesting that this modification exists even more widely than previously anticipated and the modification is often sufficiently abundant to be detected without enrichment. However, a comparison of O-GlcNAc and phospho-identifications from the very same data indicates that the O-GlcNAc modification is considerably less abundant than phosphorylation. The discovery of numerous doubly modified peptides (i.e. peptides with one or multiple O-GlcNAc or phosphate moieties), suggests that O-GlcNAc and phosphorylation are not necessarily mutually exclusive, but can occur simultaneously at adjacent sites.
O-GlcNAc proteins:
MA2B1, HGS, MYPT1, PLXB2, IF4G3, MAFK, ANR17, CREG1, SYUG, ERLN2, SC24B, AACT, LMNA, FIBG, TFR1, FUCO, K2C1, HSPB1, RLA2, ITB1, CATD, SAP, CATL1, CD3E, BGLR, LYAG, BIP, TPR, CD99, GLU2B, SON, AL5AP, MAP4, CALX, GRN, AMRP, BASI, NU214, STAT3, SOX3, LAP2A, LAP2B, SOX2, TMEDA, NU153, RBP2, SERPH, RD23B, FOXK1, SPTB2, NUCB1, AHNK, BPTF, PICAL, UBP2L, RCN1, CNN3, JHD2C, TAB1, ADRM1, QSER1, TGO1, PRC2B, TOIP1, UBAP2, LITD1, RPRD2, NFRKB, CREL2, IKIP, KDM3B, POGZ, NUFP2, YTHD3, ARI3B, ANKH1, GT251, TXND5, MCFD2, MZB1, NUDC2, ATX2L, SMG7, FKB10, PF21A, PDLI5, ZFR, EP400, PRRC1, RBM14, SIN3A, COCA1, S29A1, H2B1L, ERP44, CNPY3, YTHD1, LACRT, SP130, CPVL, WNK1, SDF2L, APMAP, DCP1A, PDLI7, RBM12, CARF, FKB11, CDK12, S30BP, SALL4, GMEB2, ZMYD8, PRP19, MINP1, SCAF8, ERGI3, SALL2, HYOU1, PRC2C
Species: Homo sapiens
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