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Hung YW, Ouyang C, Ping X, Qi Y, Wang YC, Kung HJ, Ann DK. Extracellular arginine availability modulates eIF2α O-GlcNAcylation and heme oxygenase 1 translation for cellular homeostasis. Journal of biomedical science 2023 30(1) 37217939
Abstract:
Nutrient limitations often lead to metabolic stress during cancer initiation and progression. To combat this stress, the enzyme heme oxygenase 1 (HMOX1, commonly known as HO-1) is thought to play a key role as an antioxidant. However, there is a discrepancy between the level of HO-1 mRNA and its protein, particularly in cells under stress. O-linked β-N-acetylglucosamine (O-GlcNAc) modification of proteins (O-GlcNAcylation) is a recently discovered cellular signaling mechanism that rivals phosphorylation in many proteins, including eukaryote translation initiation factors (eIFs). The mechanism by which eIF2α O-GlcNAcylation regulates translation of HO-1 during extracellular arginine shortage (ArgS) remains unclear.
O-GlcNAc proteins:
TVBY1, TX13D, CC195, SPEM3, TVB42, FR1L5, HFM1, CCD66, SBNO1, Z804B, LRRD1, XIRP2, POTEF, PSD1, BDP1, NKX26, SWAHB, F181B, SMCO2, MGT4D, CAN8, EPOP, SMHD1, AKD1B, T200C, AQP73, AXA2L, NTM2B, ZAR1L, AN36A, NEU1B, GG6L6, B2R853, KDM4E, HNRC2, ERVV1, HNRC3, TEX22, TM232, MEIOS, CK098, SO1B7, TEX46, KIF2A, BACH, MYO1C, AP3B1, IKBE, HIP1, TULP1, WASL, PDE2A, SAP18, DNM1L, IFRD1, ZN593, FAAH1, PESC, SDCB1, MPP10, NOP56, DDX3X, CCN1, PLPP3, NCKP5, AGRB1, AP3D1, IDLC, CHD1, CHD2, PDCD5, ANM5, H17B6, FPGT, TCRG1, KIF3C, GEMI2, ML12B, CLGN, MYPT1, S27A2, XPO1, AT2A1, PLXB2, SR140, PER2, GRRE1, MCF2L, NACAD, ATS3, ANR28, VILL, LAMA5, MATN3, ATX7, INP4B, TRI18, GPR37, RPA34, MAGB3, PER1, KDM6A, DHX15, TTI1, PRP4, SERA, ADA12, IRAK2, ZW10, SNUT1, SPIT2, SRGP3, VIP2, PRPF3, GRID2, TGON2, LAT, SYT7, PLRG1, BUB3, PLPP2, PI15, ACTN4, SCO2, IDH3B, CALU, AGRB3, MED14, FGF17, PRKN, JIP4, KIF5C, ZC3H1, KATIP, OPA1, KIF1B, NMD3B, PEPL, PLXC1, ADCY9, HNRPQ, TPST1, RNBP6, CCNT1, FAIM3, ANM3, HCN1, RT14, HNRC1, H2B1K, PQBP1, PRAF2, DKC1, IF2P, TRI13, DNJA2, BRD4, NBN, IGSF3, DEN4B, ADA23, MFA3L, CLAP2, OBSL1, BRE1B, RFIP3, KDM4A, DJC13, ANR17, ZPR1, NIPS2, HMMR, ZN217, SC22B, PR40A, DPOLQ, ZN254, PSIP1, ERLN1, ECI2, KHDR3, TADA3, SF3B1, MED6, FOXH1, SPB7, U520, UTP20, TCEA3, SPI2, KLF7, MMP23, RASF9, CCNK, PRAF3, DYSF, SPF27, ATP5H, TRIO, RL1D1, NEBL, OCTN2, PDE9A, SRP72, B3GA3, UBP1, SLIT2, TOM70, MICA2, NFASC, FARP2, ERLN2, PRP6, GLSK, RNF37, WDR47, AP2A2, CE152, HEXI1, UBE4B, CSPG5, LETM1, LC7L3, KIF4A, GOSR1, KAT7, MRP6, TNKS1, KCNK5, VAPB, IPO7, M3K6, UTS2, CD2B2, FMNL1, SGPL1, G6PE, SC24A, ACSL3, DUS4L, ZBT11, CPSF4, K2C75, HERC2, SNP29, AP2A1, WIZ, STAU1, BAG2, ONEC2, TOP3B, APCL, PX11B, PAK4, LDHA, NB5R3, F13A, MOS, PGK1, TRYP, OVAL, IGHG1, IGHG2, IGHG3, IGHG4, HLAH, HBE, LMNA, ALBU, VTDB, TRFE, FRIH, MIS, VTNC, TDT, ALDOA, ANXA1, APOB, OAT, K2C1, SEMG1, TBB4A, G3P, NTRK1, P53, HSPB1, RPN1, RPN2, B3A2, AT1A1, CP17A, ITB2, HMGN1, CFAI, PCCA, IF2A, HMGN2, RLA1, RLA2, RLA0, AP2A, INSR, AGAL, RB, CDK1, ATPB, ENOA, NPM, TPM3, H2B1J, LDHB, H10, CATD, ANXA2, CO8A, TBB5, PROF1, HS90A, HNRPC, TPM2, GLI1, ACM2, CEL2A, HS90B, SRPRA, ODPA, CO4A2, MET, RU17, VIME, RS17, K2C7, ANXA5, K1C16, RSSA, GSTP1, RU1C, VILI, HMGB1, TPM1, WNT2, ROA1, C1S, PARP1, H2BFM, ATX1L, MYZAP, ZCC18, POTEI, POTEJ, SP9, UBB, UBC, MPC1L, ST1A4, HS71B, C1C1L, IGG1, D130B, M14OS, TBA3D, PR20G, TZAP, ACRO, H14, ODP2, THIO, PPGB, CH60, THA, BIP, HSP7C, ODPB, PYGB, PYGM, BCR, SPTB1, TOP1, PYC, DMD, C1TC, PABP1, K2C3, CO6A2, SKI, FETUA, ACTN1, MYH7, XRCC6, XRCC5, EF2, K1C10, K1C13, K2C5, TNNT1, COBA2, SUIS, KPYM, ENPL, CCNB1, HNRPL, MYOD1, EZRI, AREG, NQO1, RS2, CD44, H15, H13, H12, YBOX3, ZNF18, ZN182, ZNF43, HSP76, HMGA1, ZNF8, KPCA, CXA1, UBF1, DESM, NEUM, CR1, IBP3, TCPA, ITB5, RL7, SON, RL17, SDC1, K2C4, NUCL, E2AK2, CSK22, NCF2, FST, IF2B, ANXA7, CD33, BTF3, TNR1B, RAB3B, CAN3, ICAL, CO5A1, PHOS, CD11B, CSRP1, FLNA, PERL, TENX, PUR6, UBA1, H1T, ROA2, SAV, FBLN1, TCEA1, ITA6, SFPQ, PPIB, RS3, PTPRE, H2B1O, COF1, IF4B, CPT2, CCND1, RAE1, KPCL, THIL, RPB1, LAMA1, DNJB1, ATPA, DDX6, CTNA2, DNMT1, U2AF2, RL13, S10A4, HMGB2, PTBP1, HGFL, STOM, PON1, 1433T, RL10, CALR, MAP4, CALX, PSA5, ABCD3, LYOX, HXD10, PTPRM, PTN4, T2EB, EPHA2, EPHA8, TKT, LMOD1, EF1D, RL12, PDIA3, 2AAA, MPIP3, BKRB2, NKTR, KPYR, SDHA, HXA10, HXA9, GBRA5, DNJA1, AKT1, HNRH1, 1433B, S10AB, P5CR1, IFNA8, DSG3, RL9, KINH, CP2CI, H2B1B, EVI2B, HS71L, HSP74, PHB1, MERL, RADI, RL22, T2FA, SCN1A, K1C9, FBN1, FBN2, IRS1, MYH9, MYH10, COPB2, SOAT1, FUS, DEK, CYLC1, MYH11, AHR, K22E, ACHA7, RL4, LONM, ACV1B, SRP14, ZEB1, TAGL2, BRCA1, VATA, GRP75, RS19, RL3, OST48, FEN1, TXLNA, TCPZ, ARL1, UBP8, RFC3, ECHA, IF2G, PERI, BUD31, NAA10, KDM5C, GARS, SYIC, NBL1, LAP2A, LAP2B, EPS15, RS27, LPPRC, RL35, PF2R, MUC18, MATR3, GPDM, RANG, NAMPT, EFTS, KI67, RAGP1, ATRX, IF2M, BAG6, RL27A, RL5, RL21, RS9, RS5, RS10, MAP1B, IQGA1, IF1AX, OMP, RL29, ATPO, TCPE, K2C6C, KC1A, MASP1, HSP77, TCPG, EFTU, SRP09, SYAC, SYCC, SYSC, YLPM1, TMEDA, RBM25, TSC2, GUAA, ENTP1, MEOX1, ST1A1, ST1A2, GATM, SERPH, KNTC1, CDK9, R144A, RL14, TCPQ, TCPD, FXR1, RAB7A, ZNF83, SSRD, AL3A2, DHB4, PSMD7, CCL11, ZNF41, ROA3, MYOM1, HNRPM, IMA1, IMA5, MP2K6, HNRPF, MSH6, HMGA2, CRIP2, COPB, COPA, CRBB1, SMTN, MOT1, IST1, SC24C, TCP4, SYRC, ATX3, TERF1, SYYC, HSP72, ATNG, EPHA4, GYS2, DRG2, XPO2, MFAP1, PAR2, BRPF1, NP1L1, ANM2, PSA, EIF3B, SYMC, HDAC4, IF6, RP1, NRG3, H2BFS, UBS3A, WDR4, SNX16, SELB, SESN2, H2B1D, NAL11, DJB13, TPIS, MYPR, MYL6, ACTB, IF4A1, RS20, S10AA, CDC42, DEST, SRP54, RAB2A, RAB14, ZC3H6, ACTZ, ARF3, ABCE1, RS3A, RL26, RL15, RL27, RL37A, DCAF7, HNRPK, RRAS2, RS7, RS8, RS15A, RS16, 1433E, RS14, RS18, RS29, RS13, RS11, SMD1, SMD2, ARF6, RL7A, RS4X, ACTA, RL23A, RS6, VISL1, H4, H2B1C, RAB1A, RAN, RL23, RS15, RS24, RS25, RS26, RS30, RL39, RL31, RL10A, RL32, RL11, RL8, PPIA, RS27A, RL40, TRA2B, AP2B1, 1433Z, 2ABA, RL38, RACK1, ACTG, ACTH, YBOX1, CSK2B, TPM4, ACTC, EF1A1, ACTS, TBA1B, TBA4A, TBB4B, CSK21, HBG1, HBG2, NOP14, RPP30, PHC1, S35A1, PRKDC, M3K9, SSBP2, RT22, RT11, RT09, ATL3, RL24, RL19, SRSF3, MUC5A, TFAM, CLH1, NFKB2, HNRPU, MYPC1, MDM2, U2AF1, AMPD2, DIAC, CAP1, K22O, CAC1D, EXOSX, PFKAP, EWS, CO4A3, AK17A, CENPE, MMSA, BDH, DSG1, H11, GHRHR, M3K10, PLOD1, RL6, PO3F1, CAV1, KMT2A, TAP2, ZN117, IF4G1, SRS11, EF1A2, CALD1, AT5G2, PTN11, GABP1, ZN33A, FMR1, PRDX1, RL18, C1QBP, CKAP4, PDE4B, KHDR1, SLC31, SOS2, SRSF1, ACTN3, TLE5, SRSF4, DHX9, GOGA2, DSC1, SSRP1, CAMP2, NSUN2, AHNK, DIK1C, LR74A, LMOD3, CF210, GALT2, AP1B1, HMGX3, NU160, TBL3, TF3C1, ASPH, TROP, BPTF, CNTN1, SFSWA, CHD3, SF3A3, NMDE1, TP53B, ILF3, TIAM1, AKAP6, FLII, TRDN, ACACA, ZBT17, REST, TADBP, ROA0, CBX3, STK3, CBLB, DNJC3, M3K1, SRSF6, I15RA, G3BP1, ATM, AP3B2, PPIG, TCOF, SF3B2, GOGA4, ROCK1, TOP3A, SQSTM, MED21, MTX1, TBB3, PRP4K, ATR, KCC2D, TDG, SNW1, DGKZ, CUL1, S12A1, MOGS, NACA, THOC5, NOG2, PTK6, TBB2A, CAC1C, CKAP5, CO4A6, LRP8, PDE1C, BOP1, SEPT6, MORC3, EFR3A, UBP2L, SIM2, DYHC1, ELOA1, TRI25, MYPC2, FRG1, CAPR1, RBM39, SCN5A, PDIA5, HS902, ZN266, ZN235, SMC1A, RRP1B, NCOA6, GANAB, 2A5D, MVP, LTBP1, GOGB1, GRM7, CHD4, LASP1, RB39A, IMB1, PSME4, CUL7, NAA25, RTP5, MO4L2, SEPT2, SUZ12, U5S1, BRD3, EEA1, PDIA6, PCOC1, PDK2, PLEC, PCM1, NONO, RASA2, RNPS1, PCBP1, PCBP2, SF3B3, RYR3, CNN3, SF3B4, RBMS2, SF3A1, SIX1, MED22, TRAM1, SF01, TRIPB, OASL, MED1, TRIP6, TWST1, PI5PA, CEBPE, AAAT, TOM34, NCOA1, GRK1, SCN9A, ZYX, SEPT7, UAP1, GBRA6, CSRP2, CDC37, IL17, DPYL2, M3K11, SRSF7, CPSF6, AT2B3, H2A2C, H2B2E, HCDH, FA98C, TIGD6, NKPD1, KNOP1, ESRG, PDS5A, MOONR, BLTP1, KIF7, TSR1, C19L2, CO3, PRTG, GAB4, ZN750, P3H1, RA1L2, MA7D1, CHD9, TKFC, PTHB1, RABL6, TBB8, CRFM7, GLYR1, C2CD3, LARP7, TB10B, A20A4, GRAP1, MIDN, PKDCC, CN177, CEP55, KLH22, PLA1A, SGO2, ACTBL, BRDT, H90B4, H90B3, H90B2, HS905, ODFP2, ESCO1, SH319, AN36C, PPM1J, FGD3, RRP12, CF157, ZN484, GNAS1, GASP1, EOGT, BRM1L, H2B2F, PPM1L, TM201, CA167, TT39A, HP1B3, CE170, FREM2, CE85L, SZT2, AXDN1, EYS, ZC3HD, GPTC4, RBM20, UBR4, GNTK, ER6L2, HENMT, TTC22, FRY, B2L15, KAD9, M3K21, DDI2, MROH9, EFCB6, A20A1, CROCC, CL060, OBSCN, FA78B, EF1A3, PR38B, BRE1A, SMBT2, SPO16, TASO2, MALR1, ECM29, SHE, NAA35, TM268, RES18, K2C79, KFA, WSCD1, F91A1, CP135, AR6P4, MROH7, UTP25, RHG17, Z518A, SCAR3, KDM4D, IQEC1, XAF1, OVOS1, CAPR2, IQCE, ZN770, F166A, NIPBL, LIN54, TET2, TEN4, OGFD2, TM214, CAVN1, CDC73, ARMX5, MED27, ANM9, PRP8, PLBL1, NFRKB, ALKB5, TRI40, TBA3E, PONL1, TTC33, MYO7B, RICTR, BRWD3, POTEE, POTEA, CC171, S4A10, FIP1, PRS55, LRSM1, SBSN, CLM9, LRIG3, RSPO2, TMC5, RFIP1, CD109, HSDL2, B3GN6, CCNI2, M3K15, LN28B, NFXL1, DYH12, RN207, ITPI1, CRACD, YC006, PLPL7, FGD6, PHLP2, ANKS3, YS043, NEK10, KRA55, KRA52, VP13C, UBP34, RAPH1, UBP51, UBP45, UBP50, IKIP, MD13L, TBA1A, B4GN4, ATS13, HECW1, BT2A1, SND1, DDX46, TM10C, RSRC2, DET1, H2A3, FAKD5, NLRC3, SSX6, MOT9, OVCH1, PLPR4, PCLI1, NRK, MON2, K1C28, K1C27, K1C25, CSMD3, NUFP2, SZRD1, NPHP3, TT21B, GDF7, KI21A, CFA91, ZN438, HDGR2, SDK1, SPRE1, SRGP1, RBBP6, NECA2, SH3R1, SEN54, K2C1B, CNTRL, CTU1, RM55, VP13B, AGRE4, GLRX5, ZN605, DAAM2, K1328, PATL1, DDX53, TRPT1, LYRIC, AKA28, KTN1, GTD2A, ABCAD, EST1A, PHLB1, LUZP1, THOC4, IFTAP, LRRT4, ZCH18, ARHGP, MON1A, THOC6, AMGO1, AMGO3, AROS, CHSS1, DDX42, NIBA3, CEP57, DAAF5, LYPD6, STX12, DZIP1, HORN, KLOTB, WFDC8, GLT13, ZDH13, CRLF3, CRERF, ARMC8, K0825, ANKY2, AHNK2, DYH10, P3H2, SPNS2, VSTM4, WDFY1, PHF6, SULF2, CHERP, ANKH1, PHC2, OD3L1, F217A, DDIAS, RB12B, SP8, DDX60, RAVR1, SPB1, DRC7, EFC13, CKP2L, SRRM1, CCD13, ELOA2, CC105, ZN100, SMYD4, MICU2, CFA46, RN168, CHSS2, RTKN2, DOCK3, AGRF3, XRN1, ASPM, DSEL, RBIS, SPICE, TAGAP, NDUF2, VPS52, LRC47, K2C78, MYO1H, KLH34, H2B3B, ANR35, GAPT, UB2J2, AN36B, LAT4, EH1L1, PALS1, FNBP4, GUF1, RIBC1, NIN, ZN276, ZN561, SRBD1, NKAP, MACOI, PAR16, AADAT, IQCF1, EFNMT, KLDC1, PLD5, ANR31, CDYL2, AMER3, NEPR1, YJ012, MUT7, BANP, SR1IP, TOPZ1, CO053, PR38A, UBP38, UN13C, FANCB, GOLM1, GT251, DHB11, TM145, SERB1, DYHC2, CC020, GIMA8, TNR6A, ZN740, DIK2A, ZN738, ABCF1, DUS18, ZN548, EXPH5, KMT2C, ABHDB, MYRIP, KNL1, RDM1, O52E5, DDX55, MAFA, ZFP28, THOC2, KCNG3, TTC8, PI42C, TBC15, MIEAP, CIP2A, ZN507, CERK1, BICD2, CHD6, HELQ, CHD5, KCNG4, BRX1, CLM1, OTU7A, ATS16, DYH5, DQX1, TP4AP, WHAMM, UBS3B, AT8B4, SHRM3, WIPF2, F222B, ZC3HF, PPIL4, CCD12, HDAC7, PDC6I, CATIN, CO040, SFT2A, SNR27, HNRLL, UBCP1, CQ050, CTL1, CKAP2, SPIR2, ATX2L, ABCAA, SREK1, CRGN, SYNE2, MUC16, P66B, DNJC9, TITIN, DDX1, H1X, TM131, IP6K1, AP3S1, HS105, SEPT8, PRP16, AN32B, TFG, AP2C, STAM1, DPF3, REQU, RBP56, EVPL, DDX17, KIFA3, SEM4D, XPF, SMRD2, RAB1C, FUBP2, KCNB2, USP9X, UBP7, CUL5, VPP1, MED12, TATD2, H2B1H, FBXW7, YIPF5, UBE2F, RL36L, GTPB3, H2B1A, CCD47, FERM2, PLPL2, LRC59, PBIP1, TTC28, BORC7, SPSB1, ANKR9, CC127, DOC10, P5CR2, ROP1L, FAF2, ZN689, AP2M1, NTNG2, KCD12, PGBD4, MTBP, RMXL1, HOOK2, RBM33, DTBP1, TMA16, ALG14, CPNE2, KIF12, SIPA1, THOC1, RM48, DDX27, P121A, TONSL, PDLI5, PGAM5, COX14, AJUBA, A26L1, PAWR, VPS50, JPH4, NEUL4, HSH2D, ZN514, AGRG7, RPGR1, ZFR, NAR5, ALPK3, IFT74, PPR36, LRGUK, DNHD1, KLH15, ZN512, RIPR3, RN145, CFA44, ZDH15, PWP2A, DOCK7, ZN830, BROMI, SCLT1, CNT3B, ANR27, GP101, IPO9, LRC3B, PGRP2, INP4A, ADCYA, GBP4, ZN317, NED4L, SLIK1, NEK1, PP1RA, XPO6, H2A1A, MAGI1, SNX18, S26A8, MCCA, CTGE2, HMCN1, S22AC, RANB9, CCNL2, SMC6, ZN347, STRBP, SIN3A, RBM15, PNKP, SETB2, NIBA2, SCN2A, NELL2, PSB7, SEC62, CDC5L, STAC, AGAP2, GA2L1, DNJC2, PHB2, KIF2C, HCD2, CPNE1, EBP2, H2B1N, H2B1M, H2B1L, MYOC, SEM3C, GDF15, AKAP9, MICU1, TRIR, SLX1, MEP50, TBA1C, SELS, MBB1A, SYT3, NRIP2, FYCO1, GPTC1, RIOK1, LLPH, WDR83, S35A5, PIMRE, SOX7, WAC, MTA3, MCMBP, MED10, LRRC1, TBB6, HNRL1, DDX23, TBB2B, TMED9, RIOK2, ELOV1, RFIP5, ASPN, AP1M1, NUSAP, TDRD1, EMIL2, AN30B, SCAPE, RTF2, DUS16, MK67I, HOIL1, RT26, KRA41, SETD2, TBD2A, ACTBM, CNTP3, PC11X, PC11Y, PUS3, UACA, REN3B, CRNL1, EDEM3, UBXN6, API5, ZFHX2, CFA74, CE295, ASXL3, ZN407, LRC27, WDR12, IMUP, SENP6, CHST5, WN10A, EGLN1, PEG3, CHST6, NACA2, PAIP1, LSG1, KLC2, ZN541, INT2, DDX47, F234A, NUCKS, S6A17, ABCG5, EHD4, P2Y12, VPS11, TARA, SLK, TNKS2, RBP17, RM46, PININ, SNX25, MOD5, MTRFR, HRH4, NDST4, PTN23, SEM6B, MFSD1, PIGU, PLK3, FA83D, TRABD, EHD1, MROH8, PDZD7, TFB2M, DHX35, ZN576, BICC1, Z385D, CCD86, PRR36, RANB3, AT133, NRDE2, METL8, ZN430, CAAP1, NOL11, ZFAN3, F204A, VRTN, MARH7, SFXN1, SPE39, VP37B, UBP42, KANL2, LN28A, EAF6, ZN556, LIN7B, RENT2, XPO5, CP343, MKNK2, UBP29, PLRKT, SPC25, GPR35, SPON1, LST2, TANC2, EPG5, K1586, ZBT26, FBSL, ZN334, SPCS, MYO10, APMAP, ZCC17, VTA1, SYSM, SIX2, DCP1A, MED4, GP108, LUC7L, SCUB2, RM40, CASS4, RTN4, IFT46, RPR1B, ANLN, BIRC6, DDX21, SAR1A, ASH1L, ARHG4, ARHG3, MXRA5, SPTN5, SNRK, STRN4, TRYG1, SOSSC, F1142, LTBP3, KRT84, DIAP3, PDS5B, SMC4, MA2C1, RN146, MDN1, LYRM2, SPS2L, ABCF3, MRGBP, SEP11, MED17, FANCI, DDX18, KLH11, RPC5, SDA1, CS073, ARGL1, PK1IP, HPIP, CB042, SDHF2, NHP2, LYAR, MED29, CWC25, THUM1, SCN3A, TBA8, DYH9, HXC10, FKB11, CELR1, CDK12, FMN2, RTEL1, IL36B, SACS, E2AK3, PA2GE, MYOF, BICRA, NOP53, HACD3, KCMF1, RAI14, VAPA, SLAF8, SIRPG, HCN3, DYH2, RCC2, TEN3, RRBP1, CPSF2, SYLC, ANKY1, HOP2, IKBL1, COPG2, MYOTI, P2RX2, SPAST, TRMB, FA8A1, DBF4A, MO4L1, MALT1, DAXX, MARCO, ZBT47, CGBP1, GCP4, KDM5B, SYWM, HMGX4, NOL12, ARMX3, SWP70, SUN2, AFF4, MKRN1, NPCL1, CHRD1, SEPT9, PCS1N, NB5R1, S30BP, PUF60, SCNBA, DNAI1, BAZ2B, PHF11, VAX2, RABX5, TRM6, ZN391, APC7, STML2, VPS28, CCNL1, FBX9, MRT4, PITC1, GPTC8, PARP4, MYO15, TF3C4, ADA28, NUP50, ZN236, ZN346, HCN2, ZN214, NRX1A, ZN608, MRTFB, ATD2B, NEB1, SHRM4, TMCC3, MYO5B, TPX2, NP1L2, MAFF, NOTC3, PRP19, G3BP2, RHG26, CHIP, MAGD2, SSRG, ING4, DIM1, TTF2, L10K, LIMC1, AGAP1, TNR6B, ZC3H4, THS7A, RIMS2, SRRM2, CTND2, BAIP2, SCN8A, PPR3C, LAMP3, RTRAF, RUVB2, LZTS1, TN13B, SYFA, SIGL7, DRG1, NCKP1, TLR6, ATF5, DIP2C, DLGP4, FYV1, KDM2A, TPPC8, KLH20, S27A6, YBOX2, TR150, WBP11, NOP58, NOSIP, LC7L2, SF3B6, ORN, NOP16, TIM16, RT23, RTCB, HCN4, ZN330, RL36, CCDC9, HBS1L, DMXL1, ZAN, KDM3A, MFHA1, PLXD1, HECD4, WDR7, IRS2, HYOU1, OARD1, SP16H, PCDGH, HEY1, SRPRB, O51B2, CORIN, DMRT2, MRCKB, NOX1, RBM8A, ZN706, SNX9, SCNAA, HCFC2, PUS1, MYH4, SPIN1, COPG1, CFA20, EMIL1, PTC2, DC1L1, STYL1, OAS3, TLL2, ZNT1, MYO16, GAK19
Species: Homo sapiens, Gallus gallus, Streptomyces avidinii, Bos taurus, Sus scrofa
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Woo CM, Lund PJ, Huang AC, Davis MM, Bertozzi CR, Pitteri SJ. Mapping and Quantification of Over 2000 O-linked Glycopeptides in Activated Human T Cells with Isotope-Targeted Glycoproteomics (Isotag). Molecular & cellular proteomics : MCP 2018 17(4) 29351928
Abstract:
Post-translational modifications (PTMs) on proteins often function to regulate signaling cascades, with the activation of T cells during an adaptive immune response being a classic example. Mounting evidence indicates that the modification of proteins by O-linked N-acetylglucosamine (O-GlcNAc), the only mammalian glycan found on nuclear and cytoplasmic proteins, helps regulate T cell activation. Yet, a mechanistic understanding of how O-GlcNAc functions in T cell activation remains elusive, partly because of the difficulties in mapping and quantifying O-GlcNAc sites. Thus, to advance insight into the role of O-GlcNAc in T cell activation, we performed glycosite mapping studies via direct glycopeptide measurement on resting and activated primary human T cells with a technique termed Isotope Targeted Glycoproteomics. This approach led to the identification of 2219 intact O-linked glycopeptides across 1045 glycoproteins. A significant proportion (>45%) of the identified O-GlcNAc sites lie near or coincide with a known phosphorylation site, supporting the potential for PTM crosstalk. Consistent with other studies, we find that O-GlcNAc sites in T cells lack a strict consensus sequence. To validate our results, we employed gel shift assays based on conjugating mass tags to O-GlcNAc groups. Notably, we observed that the transcription factors c-JUN and JUNB show higher levels of O-GlcNAc glycosylation and higher levels of expression in activated T cells. Overall, our findings provide a quantitative characterization of O-GlcNAc glycoproteins and their corresponding modification sites in primary human T cells, which will facilitate mechanistic studies into the function of O-GlcNAc in T cell activation.
O-GlcNAc proteins:
UBA6, ESYT2, HACL2, DEND3, SBNO1, XIRP2, CNOT1, PINLY, MT21E, SWAHB, P121B, TCAF2, MET15, F177B, P121C, GNAT3, MYO1G, SPT5H, TAF4, PK3CD, DNM1L, P3C2A, BT3A1, PSDE, BIN1, PITM1, DDX3X, RNT2, ARI1A, NCKP5, TRAD1, RHG33, ABLM1, KMT2D, IFIT3, HGS, MYPT1, S27A2, GAK, SC16A, SET1A, KDM6B, ARHGB, FYB1, ATX7, SHIP2, EIF3D, EIF3H, TOX3, NUP42, MEFV, DHX15, ZZEF1, PHF1, ZW10, PRPF3, TPD54, EMC8, SYNJ1, IF4G3, E41L2, WIPF1, LAT, OX1R, PLRG1, ZN207, ST1B1, LANC1, AKAP8, PLIN1, ZN292, AQR, GANP, HBP1, LY75, OGA, DIAP1, MAFK, HCN1, CCD22, BRD4, PP1RB, ABCB7, KI21B, LRP4, N4BP1, CPNE3, OBSL1, BRE1B, CAND2, T22D2, PP6R2, ANR17, H2AY, FLNB, NCOR1, PR40A, LRCH4, MPPB, PSIP1, NDUS3, KS6A5, MYCB2, U520, CCNK, CBPD, CYTF, LTN1, TOX4, PHF14, SUN1, PCF11, FRYL, TRI37, SC31A, CE152, AGFG2, SCAF4, SPN1, RTN3, APOL3, ATE1, CELF2, 6PGL, IPO7, CD2B2, ABCA1, SC24A, SC24B, PCNT, CNOT4, HERC2, HS74L, RIGI, M4K4, AIFM1, TXD12, LDHA, COX1, A1AT, FOS, LDLR, LMNA, ALBU, CYTB, GCR, HG2A, K2C1, G3P, HLAA, CPNS1, RPN1, RPN2, GNAI2, AT1A1, RLA2, JUN, ATPB, CD2, NPM, ANXA2, SYEP, TSP1, SP1, ANXA6, MDR1, HS90B, INHBA, ODPA, PTPRC, RU2B, HCK, VIME, GNAI3, ADA2A, HMGB1, ROA1, LKHA4, DERPC, F231L, GLI2, GRAB, RO60, RARB, HSP7C, EGR2, ODPB, LAMP1, SRF, FA5, IMDH2, TPR, SKI, ACTN1, K1C10, CEAM1, PLSL, GLU2B, HCLS1, PO2F1, RAC2, ATF2, FOSL2, PGCA, LEUK, CREB1, GDC, PECA1, MGMT, ZNF25, JUNB, UBF1, JUND, ATF7, PTN2, DDX5, EGR1, PTPRA, SON, RCC1, ATF1, ML12A, PLCG1, NUCL, NFKB1, LMNB1, CAN3, HNF1A, FLNA, TNAP3, PIMT, UBA1, ROA2, RFX1, CBL, QCR2, MAOM, SP100, NFYA, IF4B, AT2B4, RPB1, BRD2, ATPA, DDX6, PTBP1, ARNT, RFA1, APEX1, PYR1, CALR, MAP4, ERCC5, PTN6, SPB3, PDIA3, 2AAA, HLAF, HMOX2, CLIP1, RPB2, COR1A, ZEP2, HNRH3, HNRH1, STIP1, ELF1, KINH, LSP1, PHB1, PTN7, RFC4, MYH9, MYH10, COPB2, ACTN2, SOAT1, ADDA, FUS, NU214, ATP7B, MYH11, GLRX1, PPM1A, K22E, MP2K2, NUP62, GRP75, IF4A3, COIA1, STAT3, MDHM, ECHA, IF2G, PERI, ELK3, LAP2A, LAP2B, STAT1, RHG25, DPP6, HD, MATR3, GPDM, ZAP70, TNR4, VDAC2, MP2K4, NOP2, NOTC1, UTRN, IQGA1, STT3A, NPBW1, COPD, AGRE5, NASP, FAS, EFTU, CENPF, MA2A2, YLPM1, CLK1, NU153, RBP2, TAF6, GUAA, IDH3A, EMD, LRBA, AT1A2, MECP2, HCFC1, CCR3, KS6A3, LUM, ROA3, GDIR2, AGFG1, STAT2, TF2AA, CAZA1, NUP98, FOSB, SUCA, COPA, ITA8, SC24C, ATX1, UBP14, RD23B, EPHB3, AF17, CASP6, DSRAD, PSA, TPIS, SC61B, ACTB, ARF3, HNRPK, RS16, ACTA, GBB1, PPIA, RS27A, AP2B1, 1433Z, IF5A1, RACK1, ACTG, ACTS, TBA1B, TBA4A, PHC1, PRKDC, BTG2, SSBP2, ATL3, TXN4A, FOXK1, RHG04, NFKB2, SPTB2, FOXK2, RUNX1, AMPD2, CAP1, FLI1, OTUD4, PFKAP, SATB1, EWS, MEF2A, SP2, RHAG, SP4, SP3, RL18A, NUCB1, DYST, CREM, KMT2A, TF65, IF4G1, TLE3, TLE4, REL, UBE3A, GABPA, GABP1, CD69, ZO1, TLE5, DHX9, GOGA3, SLFN5, S38AB, RBBP4, NCBP1, AHNK, MN1, FOXO1, TBL3, TF3C1, AKP13, BPTF, NFIA, CHD3, TP53B, ANK3, PP1R8, AKAP6, ROA0, PAK2, TBX2, M3K1, ATM, DC1I2, IKZF1, TCOF, ROCK1, NFAC2, SMAD4, PICAL, PRP4K, SNW1, IQGA2, MTMR1, MTMR3, CUL4A, CUL4B, RUNX3, NFYC, KGP1, CDK13, IL16, CKAP5, CO4A6, VEZF1, MORC3, UBP2L, SCRIB, GIT2, DYHC1, ELOA1, FLNC, CAPR1, CASL, SCN5A, SEM3A, ITPR2, PLSI, LAGE3, PUM1, EPN4, RRP1B, NCOA6, LBR, STAT4, MEF2D, LASP1, NUMA1, GAPD1, SPCS2, SUZ12, ACAP1, R3HD1, SYK, ARHG6, ACAP2, BRD3, PLEC, L2GL1, EPHA7, SF3B3, RYR3, TAF5, MARE2, TSN, SF01, MED1, JHD2C, T22D1, ELF2, NAB2, TAB1, SPEG, USF2, ZFHX3, ZYX, SEPT7, ADRM1, PKN1, DDB1, TAF9, OBF1, NRF1, PTPRO, ZN827, EX3L4, HNRL2, AAK1, CCD57, QRIC1, PRTG, CEA16, TM249, FR1L6, LRRF1, EMAL3, UAP1L, GON4L, LARP7, EPC2, CRTC2, PAR10, TYW2, RHG15, H90B3, BCORL, ZN831, TGO1, DOC11, PRC2B, TOIP1, CEP78, CD158, TDIF2, KMCP1, ZN362, FKB15, ZEP3, ODAD2, MPP7, LRIF1, UBR4, UBAP2, GNTK, RBM26, CE350, RPRD2, AGAP9, MYOME, TASO2, RN213, GL8D1, PDPK2, BICRL, OTU7B, RGPA1, TWF2, SDE2, NIPBL, LIN54, ZN544, PPR18, ZCHC8, CDC73, ARMX5, SCYL2, NFRKB, LMOD2, LEG1H, TMM81, PDXD1, RSBNL, MDEAS, ZC3HE, LARP1, SCND3, POTEE, ZN322, ANR11, SPIT4, AFTIN, FIP1, CRTC3, MCAF1, PACS1, BCOR, DJC14, DG2L6, LR74B, OTOG, RHG36, YJ005, RHG27, TMTC3, UN13D, HAKAI, NOL8, HECW1, SPT6H, SND1, KDM3B, S26A9, DYM, PRS41, APTX, ZCCHV, SETX, NUP54, GLUCM, POGZ, MYH14, NUFP2, MAVS, HDGR2, EMSY, I2BP2, AB12B, DHB13, CMKMT, SRGP1, RBBP6, RHG30, NRARP, TCPR1, HUWE1, YTHD3, CENPV, ATL2, YRDC, GPAT4, ZFHX4, ABCAD, BCL9L, KIF27, LRRT3, IQGA3, VS10L, CEP57, FRAS1, CACL1, P66A, I2BP1, CRLF3, CRERF, DYH10, GID4, ARI3B, WDR75, MGAP, ANKH1, SUGP1, SUGP2, CCAR1, BAP18, PLPL6, CMIP, TIGD4, YAF2, IHO1, SRRM1, FANCM, CC116, A16A1, DCP1B, PELP1, WDFY3, ABCA7, LGI4, NUP93, LRC47, ABD12, FNBP4, GALT4, RN175, CARME, AF1L2, TAB3, CPSF7, EFNMT, MAGB6, LRTM2, KRI1, TTC29, POC5, LR75A, S43A3, SUMF2, NETO2, NF2IP, LS14A, MISSL, CA131, TNR6A, PHC3, SRFB1, SP20H, VP37A, PCAT1, DOCK8, SYNE1, ARI1B, ENASE, TET1, MYRIP, OR6K3, CFA61, THOC2, WDR36, GABP2, MARH1, ALMS1, PREX1, PKHO2, DYH3, DSCL1, DTX3L, NETO1, NEK7, MICA1, ATS18, RN128, SNX29, SMCR8, ZN384, HASP, SMAP2, SCFD1, LMO7, ATX2L, PHIP, RUFY2, CSKI1, MADD, AGRV1, SYNE2, MUC16, P66B, AUTS2, BBX, TITIN, CTTB2, GBF1, SMG7, SNX19, PHF3, HS105, ZN592, HMHA1, TFG, TAF4B, CBP, KAT6A, SYMPK, SHIP1, DDX17, TANK, RAD50, CELF1, SMRD2, RAB8B, FUBP2, DVL3, LPP, TATD2, AT2A3, MRTFA, PLPL2, SH3K1, PF21A, DOC10, INT12, ACSF2, GCP3, SLAF6, RSPRY, MTEF3, SIR1, THA11, GLT14, CERS2, SYMM, PDLI5, FUBP3, PP16A, COG3, VCIP1, CHAP1, PDLI2, ANCHR, UBP47, Z512B, ZFR, EP400, CNO6L, CA074, PRRC1, ZN512, CNT3B, LRRC7, ARAP1, AGRA2, INP4A, RBM14, NED4L, LENG8, TRNT1, MCCA, PCX1, CCNL2, SIN3A, SEBP2, MINT, HTF4, CDC5L, EYA3, LGMN, MNT, SCAFB, TTC1, OSMR, ATX2, METH, ACON, CPNE1, TBA1C, MBB1A, GPTC1, ERP44, ESYT1, CCM2, FUZZY, DIDO1, MCMBP, CABL2, NDC1, PAXX, HNRL1, NUP58, RIOK2, THIC, RBM4, NADAP, SSBP3, NAA15, AP1M1, M10L1, YTHD1, BACH2, PANK2, PC11Y, ASPC1, UCK2, TRI31, UNK, FTO, AMRA1, CE295, DRC3, SP130, BRD8, CSTFT, ZCPW1, LMA2L, CK054, SLIK2, CSRN2, I2BPL, VPS16, EPC1, ADNP, IPYR2, FOXP1, PTN23, WNK1, AMPB, E41L1, GSX1, ELOV6, CH033, VISTA, SFR19, GORS2, LN28A, MLXIP, GBB4, PKHA1, RISC, TAF9B, MRM3, ZBT20, NCOA5, TANC2, TNR6C, CHD8, AT131, VTA1, SYSM, UBN1, DCP1A, KI13B, PRD10, XPP1, PDLI7, DDX21, MBNL1, SIR7, TULP4, ABCBA, LATS2, UBQL4, THSD1, CENPM, PDS5B, RBM12, MED9, SLTM, MIC19, NSMA3, THUM1, CARF, SNTG2, MTMR4, TE2IP, TAB2, CDK12, GGA3, ITSN2, BICRA, CNOT2, TMOD2, THYN1, PDP1, VAPA, TEN3, CHD7, DYH1, SYLC, KLH42, KANL3, RERE, HDC, TRPC4, MALT1, ADDG, TCF20, NDRG3, SUN2, NDOR1, UBQL2, S30BP, RPGFL, AGRE2, NRBP, BAZ2A, HOOK1, S2513, TASOR, AKA11, GMEB2, PARP4, C8AP2, IKZF2, ACINU, CNO11, AT7L1, K1210, YETS2, HECD1, NOTC3, PRP19, UBQL1, FAF1, PPIE, DIM1, MACF1, SCAF8, SET1B, JIP3, ZC3H4, SRRM2, CLCA2, SMC3, ZN148, MTMR6, ENTP4, SAC2, MAST1, FYV1, WDR37, TR150, ZN281, FBX7, 3BP1, OR2W1, NOC2L, SAMH1, DMXL1, ARIP4, MTCL1, RIPR2, PKHM1, RPGF2, CRBG1, PRC2C, YTHD2, SP16H, ANGL3, PCDG3, SNX13, NUBP2, NCOR2, COPG1, GMEB1, DC1L1, ROBO1, NCOA3, M3K4, PCLO, CAN7, SCC4, ZHX2, S23IP
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, HCFC1, 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, TLE3, 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, SNUT2, 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, CC168, 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, HBS1L, 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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Shen B, Zhang W, Shi Z, Tian F, Deng Y, Sun C, Wang G, Qin W, Qian X. A novel strategy for global mapping of O-GlcNAc proteins and peptides using selective enzymatic deglycosylation, HILIC enrichment and mass spectrometry identification. Talanta 2017 169 28411811
Abstract:
O-GlcNAcylation is a kind of dynamic O-linked glycosylation of nucleocytoplasmic and mitochondrial proteins. It serves as a major nutrient sensor to regulate numerous biological processes including transcriptional regulation, cell metabolism, cellular signaling, and protein degradation. Dysregulation of cellular O-GlcNAcylated levels contributes to the etiologies of many diseases such as diabetes, neurodegenerative disease and cancer. However, deeper insight into the biological mechanism of O-GlcNAcylation is hampered by its extremely low stoichiometry and the lack of efficient enrichment approaches for large-scale identification by mass spectrometry. Herein, we developed a novel strategy for the global identification of O-GlcNAc proteins and peptides using selective enzymatic deglycosylation, HILIC enrichment and mass spectrometry analysis. Standard O-GlcNAc peptides can be efficiently enriched even in the presence of 500-fold more abundant non-O-GlcNAc peptides and identified by mass spectrometry with a low nanogram detection sensitivity. This strategy successfully achieved the first large-scale enrichment and characterization of O-GlcNAc proteins and peptides in human urine. A total of 474 O-GlcNAc peptides corresponding to 457 O-GlcNAc proteins were identified by mass spectrometry analysis, which is at least three times more than that obtained by commonly used enrichment methods. A large number of unreported O-GlcNAc proteins related to cell cycle, biological regulation, metabolic and developmental process were found in our data. The above results demonstrated that this novel strategy is highly efficient in the global enrichment and identification of O-GlcNAc peptides. These data provide new insights into the biological function of O-GlcNAcylation in human urine, which is correlated with the physiological states and pathological changes of human body and therefore indicate the potential of this strategy for biomarker discovery from human urine.
O-GlcNAc proteins:
TX13C, ESYT2, BICL2, ODAM, SRCRL, SYTC2, Z804B, O2A25, XIRP2, NKX26, SMCO2, MFS2B, O51F1, NCF1B, ABTB3, SRRM4, D11L8, YV023, LEUTX, MEIOS, RB27B, CACB4, SOCS6, CHD1, NDC80, KIF3C, MYPT1, IPP2C, MUSK, MRP3, PA2GX, ARPC5, P2RX6, ZW10, ZN749, KCAB3, ACTN4, VEGFD, BNI3L, ZN292, PI51C, MABP1, CCG3, NOBOX, REV3L, TBL1X, NBN, KI21B, PX11A, UBP2, CAN15, ATRN, SPF30, MYO1D, SUN1, ENDD1, M4K4, CFAB, LV151, K1C14, K2C1, HSPB1, CHLE, SAP, SRPRA, GNAI3, IL6RA, VILI, AT8OS, GLI3, RNAS2, LYAG, SPTB1, PSG2, LAMP2, CSPG2, SC5A1, F261, DPEP1, EPB42, ITB6, LMNB1, NEBU, RYR1, TENX, SP100, ANX13, ADH6, GNA11, NMT1, CPSM, GBRA5, AKT2, I5P2, MYH11, HMGCL, PGM1, CDN1A, MLH1, SATT, DCC, MAGAA, MMP13, ABCG1, MP2K3, UTRN, IDHP, PSB3, RBP2, MRE11, ETV1, IRAK1, ARSD, NEK4, SPSY, COPA, SMTN, ATN1, PMS1, PMS2, ATNG, PRRX1, AF17, NXPH1, NAL12, IF4A1, STX1B, KCNJ2, HPCA, PKD1, REEP5, CAC1D, MMSA, SEMG2, ACY1, P, ZNF91, LG3BP, PDE4C, SCAP, PO4F2, NMDE1, OVGP1, ANK3, NFAC3, AKAP6, CENPR, SF3B2, TRA2A, CUL4B, ALKB1, CO9A2, FA53B, WRN, SLBP, GOGB1, ZN169, ODFP1, FRPD4, MELT, SART3, IF4H, KIF14, PLCL1, PLEC, PSG5, DLGP5, MARE2, ENOX2, CNGA2, AINX, HCDH, CSPP1, QSER1, ZN423, SPKAP, PRSR3, ATOSA, K2C71, GON4L, GNPTA, LEGL, PAR14, PCDP1, PLCH1, AARD, RHG29, SPIN4, FA76B, CX066, BRM1L, ODR4, SZT2, SYRM, EEIG2, CE162, CL060, MYOME, ARHGG, CA140, CARL1, ZMYM4, EST4A, F219B, WDR25, F90A2, LAR1B, ATG9B, ARID2, FTM, USF3, KCD16, ZNF57, K1C39, Z518A, BLT3A, CV042, RTL6, CAPR2, ADAM5, ZNT6, CA094, VIP1, AGRD1, CC171, KLH24, ABRX1, PAMR1, TM14E, ITIH6, RFIP1, IGS10, WDR87, SHSA6, FGD6, RGMC, NEK10, UBP31, NOL8, MARK2, BEND5, PCAT2, EPMIP, DPH6, OLIG3, TRPM8, CC186, MYCPP, TMC7, Z804A, TAF8, RALYL, RBM23, CC190, PKHL1, GA2L3, TCAM2, STX12, KI18B, RB6I2, ARMC8, K0825, STH, RP1L1, TPH2, F217A, PLPL6, EFC13, CJ067, PSYR, TBC21, DOCK3, AGRF4, SYCE1, CADM3, CP4Z2, CLASR, ZN786, CLIP4, CBPA6, NKAP, TM156, CPSF7, EFNMT, IGS22, LMBL4, ZFP62, PHC3, DDHD1, EXPH5, NAV1, BD1L1, BPIB6, TET1, MYRIP, FBH1, O10K1, ST3L4, CHD6, DMXL2, MIPT3, ES8L3, DOT1L, NAT14, CKAP2, ARAP3, CSKI1, ATRIP, MUC16, ELYS, TITIN, LZIC, PHF3, TBCD5, K0232, PRCC, TFG, SFRP3, COR2A, ARHG2, TATD2, LENG9, FAM3D, ALKB8, CHM4C, LRC58, REPS1, PGBD4, P3IP1, CDCA5, HMCES, DMAC1, IGS21, PAWR, ITCH, M4A14, CLMN, S41A2, HSH2D, TM87B, ZN514, P12L2, C295L, CQ10A, ROBO3, WWAS2, DB118, KI20B, PHF12, SPAG5, OR2M4, HMCN1, RANB9, CCNL2, IWS1, K1C12, NPY6R, S1PR3, LYST, CDC6, EBP2, NIPS1, DDX50, FA83C, PIMRE, NDC1, MTNA, UT14A, NUP85, TDRD1, MSTRO, SETD2, OSBL8, UACA, TSG10, TB182, EGLN1, CRLD2, CSTFT, NKX24, CT191, ESF1, PIEZ2, RANB3, CSR2B, UCK1, ZN556, MLXIP, TNR6C, HMX1, EQTN, PRDM9, TLR8, HELLS, TOPRS, KIF15, RAD18, HOME2, BRWD1, TEN2, CCM2L, FGOP2, MCUB, MIC19, KCTD5, CARF, FAT2, DTL, SACS, KLHL8, CE126, WDR35, NRX2A, DNM3B, COPG2, GCP4, PARP2, TCF20, ASIC3, RABX5, STAG3, NGAP, FBX5, MKLN1, ZBT21, PKHH1, SOX13, LIMC1, EXOC7, CBPC1, TUTLB, XCT, SHOC2, RUVB2, PDE10, PRLD1, FBW1A, DLGP4, WDR37, ZBTB1, NSG2, LSM2, DOP2, C170B, SAM50, PCDBB, PCDA3, TF3C3, CCG2, BIG1, S4A4, PCLO
Species: Homo sapiens
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