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Li G, Meng F, Zhong X, Yu K, Zhang N, Zhang K, Huang H, Li W, Zhang J, Wang W, Ren Y, Liu Y. DNA probe pulldown screening uncovers O-GlcNAcylation modulation of transcription factor DNA interactions. Scientific reports 2025 15(1) 40596516
Abstract:
O-linked β-N-acetylglucosamine (O-GlcNAc), a critical post-translational modification predominantly found in the nucleus, plays a substantial role in regulating gene expression by modulating transcription factors (TFs) activity. However, quantitative analysis investigating the influence of O-GlcNAcylation on protein-DNA interactions at a proteome scale remains undone. Herein, a pulldown screening approach using a consensus TF response element (catTFRE) was employed to unravel the impact of fluctuating levels of O-GlcNAcylation on the DNA binding efficiency of endogenous TFs/co-factors. Utilizing quantitative proteomics, we identified a substantial enhancement in the binding capacity of 241 nuclear proteins (NPs) to DNA sequences due to elevated levels of O-GlcNAcylation, whereas a decrease in DNA binding was observed for 2 NPs concurrently. Intriguingly, the O-GlcNAcylation elevation significantly enhanced the binding of 146 TFs/co-factors to specific DNA sequences. We further established that the O-GlcNAcylation of several Forkhead family TFs, including FOXA1 and FOXC1, notably enhances their binding to specific DNA sequences in living cells. Our research presents an efficacious approach to assessing the impact of O-GlcNAcylation on the interactions between proteins and DNA. This significantly enhances our understanding of the role O-GlcNAcylation plays in the regulation of transcription.
O-GlcNAc proteins:
RBM47, ESYT2, HACL2, VWA8, CNOT1, PGP, SMHD1, U119B, MET15, MYO1G, ADAS, PDLI1, BACH, HAX1, SPT5H, TAF4, CLIC1, BMAL1, SAP18, DNM1L, PLOD2, PSDE, BIN1, NOP56, DDX3X, TRI38, NFIB, PPP6, CDKA1, ABLM1, ADA10, CHK1, PSA7, TX1B3, TIM23, MYPT1, S27A2, XPO1, SYN3, ZN609, SR140, SET1A, ZN185, ATX7, OGT1, EIF3D, EIF3H, TNC18, MOT4, P4HA2, NUP42, ARPC5, CLOCK, MAFG, KDM6A, DHX15, RRP8, PRP4, SERA, PHF1, DC1L2, PSMD3, SNUT1, MCA3, HNRPR, PRPF3, TPD54, EMC8, MGA, KLF4, TIM44, PLRG1, ZN207, BUB3, ACTN4, BUD23, ENSA, SPOP, LANC1, AKAP8, CALU, ORC5, ZNRD2, MED14, NPA1P, ZC3H1, AQR, GANP, KDM1A, ACSL4, VINEX, CCNT1, PLOD3, TSN3, PLIN3, MAFK, UGDH, PQBP1, TBL1X, NBN, MITF, CPNE3, ZC11A, ANR17, H2AY, FLNB, NCOR1, SPAG7, RM33, SC22B, PR40A, MPPB, KTNA1, PSIP1, ERLN1, NDUS3, SRS10, SF3B1, CSDE1, U520, NU155, CRTAP, S2512, EIF3G, SOAT2, CCNK, SPF27, SPF30, RL1D1, CLPX, SYUG, DFFB, SRP72, RECQ5, ATG12, TOX4, K0754, SRBS2, SUN1, ERLN2, PRP6, SC31A, UBR5, AGFG2, SCAF4, DUS14, SPN1, NDUB8, RTN3, EPN2, LC7L3, MBD4, VAPB, 6PGL, ZBT7A, IPO7, SLU7, CD2B2, PSMG1, SC24B, ZBT11, SNP29, OXSR1, HS74L, LSM8, AP2A1, WIZ, BAG2, AIFM1, BPNT1, SNAI1, DDAH2, TXD12, MBD3, TOM40, NAPSA, ACL6A, CCNE2, LDHA, NB5R3, PGK1, A1AT, FOS, RASH, LDLR, LMNA, ALBU, TFR1, TRFL, FRIH, NU3M, MIS, VTNC, CATA, GBA1, FUCO, ALDOA, CYTB, GCR, SODM, KITH, K2C1, G3P, HLAA, CPNS1, HSPB1, TYSY, RPN1, RPN2, GNAI2, AT1A1, AT1B1, A4, APOD, ALDH2, PERM, PPBT, IF2A, RLA2, RLA0, JUN, ITB1, K1C18, K2C8, GELS, ATPB, IF4E, ENOA, NPM, ITAV, GDN, ACBP, LDHB, CATD, ANXA2, TBB5, SAP, CATL1, SYEP, CATB, HNRPC, FUMH, TSP1, SP1, ANXA6, HS90B, SRPRA, ODPA, RU2B, RU17, ITA5, NFIC, VIME, K2C7, GNAI3, ANXA5, SNRPA, HMGB1, DLDH, ROA1, RU2A, LKHA4, CTF8, MNS60, RO60, MYBB, LYAG, HLAC, KAP0, CH60, CLUS, BIP, HSP7C, GTR1, ODPB, LAMP1, ACADM, TOP1, TOP2A, G6PD, MPRI, PRPS2, PABP1, PCNA, ADT3, FA5, IMDH2, TPR, KCRU, SKI, ACTN1, XRCC6, NR2C1, LAMP2, EF2, K1C10, K1C13, PDIA4, PLST, T2FB, MIF, FPPS, KPYM, ENPL, MYL6B, RSMB, PO2F1, PLAK, ALDR, RAC2, ERF3A, EZRI, ATF2, FOSL2, MCP, NDKA, GNS, RS2, DESP, MUC1, CREB1, MGMT, CD36, STMN1, YBOX3, ZFX, JUNB, UBF1, JUND, ATF7, PTN2, DDX5, ARF4, PTPRA, SON, NELFE, RCC1, ATF1, ATF6A, XRCC1, LFA3, NUCL, ELK1, TFE3, CSK22, NFKB1, TYPH, TBP, RAB6A, PSB1, LMNB1, HXA5, CSRP1, FLNA, VDAC1, SDHB, CD9, PIMT, FBRL, UBA1, NDKB, ROA2, RFX1, KAPCB, QCR2, ITA6, SFPQ, PPIB, MAOM, SP100, NFYA, SAHH, COF1, IF4B, GATA2, THIL, MCM3, RS12, BRD2, TYY1, ATPA, PSA1, PSA4, ITA3, DDX6, PTBP1, SYVC, KAD4, 1433T, VATL, ARNT, RFA1, APEX1, PYR1, CALR, MAP4, CALX, PSA5, PSB4, PEX2, NDUS1, TEAD1, GRN, T2EB, 3MG, TKT, SPB3, ECHM, KCY, PEBP1, PDIA3, 2AAA, CDC27, HMOX2, PURA2, AMRP, RPB2, SDHA, METK2, HXA11, HXC9, CPSM, 3HIDH, PUR9, HNRH3, HNRH1, 1433S, STIP1, L1CAM, PRDX2, P5CR1, ELF1, CGL, RL9, CSTF2, MCM4, MCM7, HSP74, TRY3, GPC1, CTNA1, BMI1, PCGF2, PHB1, RADI, RFC4, RFC1, MYH9, MYH10, COPB2, SOAT1, ADDB, BASI, FUS, NU214, DEK, GLRX1, K22E, MP2K2, VATE1, RL4, PGM1, SPB5, ODO2, SRP14, NUP62, TAGL2, SYUA, ETFB, RBMX, VATA, HSPA9, IF4A3, TXLNA, TCPZ, PBX2, MLH1, STAT3, MDHM, ECHA, IF2G, S19A1, ELK3, ACTY, LAP2A, LAP2B, LPPRC, THIM, ECE1, MUC18, MATR3, GPDM, SSRA, RANG, PRS6B, VDAC2, MP2K4, KI67, RAGP1, RECQ1, NOP2, CRKL, NEDD4, IQGA1, STT3A, CAPZB, RL29, UCRI, RFX5, COPD, PSMD8, PRC2A, HSP13, AGRE5, DPOD2, NR2C2, RPIA, LMAN1, FAS, CDK8, RM19, AL7A1, CENPF, SYAC, SYCC, SYSC, MCM2, ACADV, YLPM1, RBM25, CLK1, NU153, RBP2, GSK3A, GSK3B, TAF6, GUAA, DNLI3, MRE11, IDH3A, EMD, CPT1A, SERPH, F10A1, MXI1, MAP2, PPT1, TCPQ, TCPD, FXR1, FXR2, RAB5C, RAB7A, SMCA4, IDH3G, GALK1, MECP2, DHB4, SPHM, HDGF, ROA3, 6PGD, IMA1, GDS1, MP2K6, AGFG1, HNRPF, TF2AA, MSH6, CAZA1, CRIP2, NUP98, PLK1, ACLY, SUCA, COPA, PTTG, SC24C, TCP4, ICLN, HIRA, SYYC, UBP14, HSP72, AT1B3, RD23A, RD23B, KAD2, SNAA, IF5, RAD, TERA, DSRAD, TPD52, SEC13, PSA, HNRH2, EIF3B, SYMC, DPOE2, ATP5I, IF6, NU107, SESN2, FOXL2, EPIPL, TPIS, SC61B, ACTB, IF4A1, PRPS1, SRP54, RAB2A, ARP2, ACTZ, CSN2, ARF3, CH10, STXB1, B2MG, WDR5, HNRPK, RS16, SMD2, ERF1, RS4X, ACTA, RL23A, RS6, H4, GBB1, RL32, RL8, PPIA, RS27A, RL40, AP2B1, 1433Z, IF5A1, RACK1, ACTG, YBOX1, TPM4, EF1A1, ACTS, TBA1B, TBA4A, TBB4B, CSK21, HBB, HBA, CXAR, NOP14, RBM6, PITX1, IF4G2, GTF2I, GSTO1, PO6F2, PRKDC, BASP1, RT22, RT05, RT11, RT34, RT09, HMGN5, SARNP, LACTB, TXN4A, RL19, FOXK1, RBM10, EM55, TFAM, VIGLN, CDK5, CLH1, HNRPU, SPTB2, TIAR, SET, FOXK2, RUNX1, CAP1, TFAP4, EXOSX, XPC, EWS, MEF2A, SP2, ODO1, KIF23, DSG1, SP3, RL18A, FKBP4, PLOD1, TOP2B, M2OM, DYST, CREM, KMT2A, CENPC, LMNB2, UPAR, KIME, TAP1, IF4G1, K1C17, SSBP, SRS11, EF1A2, CALD1, KDSR, SUH, GABPA, GABP1, PRDX1, RL18, C1QBP, CKAP4, ZO1, KHDR1, SRSF4, PP2BA, DHX9, LG3BP, MFGM, CD47, SSRP1, SLFN5, RBBP4, NCBP1, EP300, AHNK, MGAT2, GALT2, BST2, MPPA, SIA4C, NU160, TBL3, TF3C1, ASPH, SNF5, BPTF, NFIA, CHD3, TP53B, IFRD2, AIMP1, ILF3, LMAN2, FOXC1, PP1R8, ECH1, PRDM2, CSN1, KLF10, NR1H3, TADBP, ROA0, PAK2, PSMD2, DDX10, GPS2, SRSF5, SRSF6, TIF1B, G3BP1, STF1, NMI, KCAB2, SKP2, PABP4, GRB10, MTA1, DC1I2, ORC2, SNTB2, PPIG, TCOF, ADAM9, TUSC3, SMAD4, PICAL, TBB3, ASAH1, HDAC1, SNW1, STIM1, CUL4A, CUL4B, FHL1, RUNX3, THOC5, SPTN1, NOG2, KLF5, RUNX2, PEBB, NFYC, CKAP5, SCRB2, DAG1, VEZF1, DSG2, KEAP1, MORC3, SAFB2, EIF3A, UBP2L, MLEC, ZMYM3, DCTN1, DYHC1, EI2BA, ELOA1, MAP7, SRC8, FLNC, CAPR1, NF2L1, HABP2, DHX8, ITPR3, PLSI, TRIPC, PUM1, MDC1, EPN4, SMC1A, RRP1B, RRP5, BMS1, GANAB, MBTP1, 2A5D, LBR, CBX2, KIF22, MEF2D, CHD4, LASP1, ARI5B, NFIX, ZN638, NOLC1, NUMA1, GAPD1, SPCS2, EMC2, MO4L2, ABRX2, CND1, SUZ12, U5S1, SYK, RRS1, IF4H, BRD3, WDR43, ACOX1, EBP, PLEC, PON2, NONO, PTPA, PWP2, RNPS1, RCN1, PCBP1, MTFR1, SF3B3, PUM3, DLGP5, KS6A1, SF3A2, SC23A, SF3A1, SURF1, MED22, TAF5, TAF7, MARE2, TSN, SF01, PCH2, MED1, JHD2C, TRIP6, MARE1, ELF2, NAB2, AAAT, USF2, VAS1, RB11B, EZH2, ZYX, SEPT7, ADRM1, TAF12, DDB1, SNPC1, CDC37, DPYL2, SYPL1, TAF9, SRSF7, DREB, NRF1, PCKGM, DDR2, LY6K, HNRL2, PDS5A, ZN800, P3H1, RA1L2, MA7D1, TBRG1, LARP7, ACSF3, TPRN, EPC2, DHB12, UBP39, SETMR, YIF1B, ZN326, NOM1, SH319, RRP12, TOIP1, TDIF2, NU188, HP1B3, ZN362, SYRM, ZC3HD, LRIF1, RM02, UBAP2, RBM26, RIF1, MRCKA, RPRD2, BRE1A, TASO2, KHDR2, AR6P4, ARID2, GL8D1, NR3L1, MSL1, ABLM2, GLYAT, TWF2, K0930, SN12L, SDE2, LIN54, ALKB2, PPR18, CAVN1, CDC73, PRP8, SCYL2, GOLM2, RHGBA, NFRKB, RSBNL, KCD18, NCEH1, MDEAS, ZC3HE, SAMD1, C1TM, FIP1, CSPG4, PLGT2, SBSN, CF089, LCN15, CREL2, LRIG3, BCOR, MPRIP, CD109, SRCAP, UBN2, IKIP, CENPU, LARP4, HAKAI, NOL8, ASXL2, SND1, DDX46, TM10C, EIF3M, MCM10, NDUF7, CXXC5, APTX, ZCCHV, KANL1, NUP54, POGZ, NUFP2, SZRD1, CLAP1, HEAT3, HDGR2, BRAP, EMSY, I2BP2, RBBP6, YTHD3, TAF8, TMED4, CENPV, PHLB2, MYPN, ACOT1, PABP2, LDB1, PB1, YRDC, LYRIC, KTN1, LUZP1, CAND1, THOC6, DDX42, SPAS2, CACL1, P66A, RIOX2, CRLF3, VRK3, NAV2, CCD50, MISP, CTL2, MGAP, SFPA2, SFTA1, PHF6, CHERP, SUGP1, SUGP2, CCAR1, BAP18, RM41, KLHL7, MILK2, DHX37, YAF2, SRRM1, ZZZ3, K2013, ELMD2, CORA1, PELP1, TTC5, MIER1, NDUF2, NUP93, ZN687, CMTR1, LRC47, INT1, ABD12, CARME, FBXL6, SRBD1, NKAP, RP25L, ATPF2, RM50, CPSF7, ARFG2, EFNMT, ARFG1, PAF1, ENAH, DAND5, KRI1, MLKL, XXLT1, GOLM1, SUMF2, SUMF1, PGLT1, DHB11, TXND5, SERB1, CHSTE, LS14A, FSAF1, RB15B, PHC3, DIK2A, POPD1, SRFB1, SP20H, VP37A, PCAT1, BD1L1, NUP35, KNL1, PLBL2, DDX55, THOC2, WDR36, KI18A, GABP2, NPL4, STT3B, SPP2B, BRX1, STK35, NEK7, GPX8, PO210, GEMI5, FBF1, ZN384, WIPF2, SETD7, SMAP2, TM263, NU133, PDC6I, CO040, NUDC2, TWST2, SCFD1, PCNP, SPRY4, TRUB1, LMO7, CTL1, CKAP2, ATX2L, PHIP, RT4I1, NELFB, PALLD, PYDC1, PSPC1, P66B, BBX, CTBL1, ELYS, S39A7, H1X, RT27, NR4A3, PHF3, NDRG1, HS105, ZN592, LAR4B, NU205, PRCC, TFG, TAF4B, RREB1, HDAC2, SC65, SYMPK, GGH, DDX17, RAD50, CELF1, GSLG1, GPKOW, SMRC1, SMRD2, RAB8B, FUBP2, TNPO1, ARHG2, GLMN, KAT5, UBP7, POP5, LSM10, LMBL2, NCLN, FUBP1, SOSB2, PBIP1, SH3K1, PF21A, PINX1, P5CR2, INT12, FWCH2, FAF2, RMD1, MSI2H, RRFM, MTEF3, THA11, SGF29, DTBP1, PTCD3, MMAB, MCRS1, CERS2, SMRD1, TOE1, DDX27, SYMM, P121A, PDLI5, ERO1A, INT4, FUBP3, SYNM, VSIG2, HPDL, PAWR, RBM41, RSRC1, THOC3, CHAP1, S35E1, CLP1L, Z512B, ZFR, EP400, CNO6L, PRRC1, ZN512, YTDC1, SPT33, ZN830, RPR1A, RBM14, NED4L, QKI, LENG8, TRNT1, PP1RA, DEPD7, ATAD5, TEFM, MED15, EFGM, PIGS, TM209, IWS1, SIN3A, RBM15, MINT, NIBA2, YMEL1, HTF4, CNN2, SEC62, CDC5L, PSMD1, RING2, PARK7, EYA3, LGMN, PLIN2, POP1, SCAFB, TEAD3, TTC1, PHB2, CSN8, HCD2, SMAD5, ROAA, S29A1, CPNE1, MYD88, PKP2, SH3G1, GDF15, ARP5L, ZFY21, CMS1, DPH2, TBA1C, BCL7B, MBB1A, COR1B, TIRR, FSP1, PELO, ERP44, TACO1, ESYT1, CCM2, CNPY3, DIDO1, NDC1, ZSA5A, PAXX, HNRL1, RPP25, RCAF1, TBB2B, TM109, NUP58, GNL3, RIOK2, TIM21, THIC, RBM4, MND1, NADAP, SSBP3, RFIP5, ORNT2, NAA15, SRRT, AP1M1, NUSAP, HDHD5, RTF2, PECR, PDIP3, CEP41, RM37, GTPB4, UACA, UCK2, TB182, LNP, CEP44, WDR33, CTDS1, MFF, RB33B, NAT10, SP130, BRD8, RGAP1, CSTFT, LMA2L, CK054, F234A, RBM38, TDIF1, ALX4, SIL1, APC1, TMX4, WDR13, TARA, EPC1, ADNP, IPYR2, RM46, PININ, TM245, FOXP1, CPVL, GHITM, NELFA, BORG4, UN45A, SMOC2, AMPB, ZHX3, EMRE, REEP4, TM231, SH24A, DOCK5, CH033, SDS3, ACAD9, CQ10B, GORS2, JUPI2, SFXN1, SPE39, UBP42, RPF1, RPA2, GBB4, BRF2, RISC, CYBP, TENS1, TAF9B, TM165, MRM3, NCOA5, TRI39, SDF2L, AT131, RM47, APMAP, ARFG3, VTA1, PLPL8, SYSM, OSGEP, DMAP1, UBN1, RTN4, KMT5A, INCE, PRD10, ANLN, PDLI7, DDX21, MBNL1, SIR7, AAAS, ABCBA, PNO1, MBIP1, RAD18, SLX9, BMP2K, PDS5B, ECHD1, MDN1, DECR2, STAU2, DDX28, T106B, SPS2L, TYDP1, SEP11, DJC11, HAUS2, RBM22, MED9, ARGL1, SLTM, RMND1, AKIP, OXSM, THG1, HPF1, NDUBB, OCAD1, MARH5, MIC19, SIR5, CWC25, TEX10, THUM1, ING3, CARF, NSE4A, AATF, DDX56, TE2IP, ABI2, HXC10, BCLF1, TMOD3, MTCH1, HPBP1, MAT2B, ITSN2, CHMP5, THYN1, VAPA, RCC2, SLAI2, BCCIP, FPRP, RRBP1, CPSF2, RBM27, KANL3, GRHPR, NXF1, DAXX, TRM7, VTI1B, ABCF2, SYWM, SEC63, NDRG3, SUN2, LIMA1, SEPT9, TRPS1, DPP2, BAP29, NB5R1, S30BP, MED13, KAD3, HACL1, S2513, DBNL, CDC23, GMEB2, RCOR1, ACINU, NUP50, ZHX1, CDV3, CNO11, INT6, MCTS1, ZBT21, YETS2, ZMYD8, TMCO1, EPDR1, MYO6, KMT2B, PRP19, G3BP2, PACN2, MAGD2, PPIE, DIM1, MINP1, MACF1, PHF8, LIMC1, XCT, SRRM2, PA2G4, SMC3, SCML2, ZN148, RUVB2, VDAC3, RT33, PLAK2, ICE1, DDX52, MAN1, TR150, WBP11, MED16, NOP58, ZN281, SYYM, COQ6, LSM2, PHOCN, ORN, RT23, HDGR3, STRAP, RTCB, NOC2L, CHTOP, VPP2, TLN1, TRRAP, CRBG1, HYOU1, LAS1L, SAM50, PRC2C, RBM7, INSL6, YTHD2, PAXB1, SP16H, UTP18, CD2AP, SRPRB, TSSC4, SNX5, NUBP2, HCFC2, LRRF2, NPTN, 5MP1, DC1L1, CABIN, COX11, SCC4, ZHX2, MORC2
Species: Homo sapiens
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Vang S, Helton ES, Guo Y, Burpee B, Rose E, Easter M, Bollenbecker S, Hirsch MJ, Matthews EL, Jones LI, Howze PH 4th, Rajasekaran V, Denson R, Cochran P, Attah IK, Olson H, Clair G, Melkani G, Krick S, Barnes JW. O-GlcNAc transferase regulates collagen deposition and fibrosis resolution in idiopathic pulmonary fibrosis. Frontiers in immunology 2024 15 38665916
Abstract:
Idiopathic pulmonary fibrosis (IPF) is a chronic pulmonary disease that is characterized by an excessive accumulation of extracellular matrix (ECM) proteins (e.g. collagens) in the parenchyma, which ultimately leads to respiratory failure and death. While current therapies exist to slow the progression, no therapies are available to resolve fibrosis.
O-GlcNAc proteins:
NUD4B, GAL3B, UBA6, HACL2, SPD2B, NBAS, SBNO1, VP37C, CNOT1, ZN316, DDTL, ZSWM8, P121C, GNAT3, EIFCL, C2D4D, IGLL5, NACAM, DNS2A, ADAS, PDLI1, BACH, MYO1C, AIP, GTPB1, AP3B1, PSD11, PSD12, PSMD9, ATOX1, CBX4, PGRC1, SPT5H, TAF4, DFFA, CLIC1, TR11B, EIF3F, ODPX, MATN2, QSOX1, WASL, IPO5, EMAL1, IF2B3, DNM1L, MANBA, PLOD2, MEIS1, PSDE, ZN593, IMA4, BCL9, PESC, NOP56, DDX3X, CCN1, IMA3, MA2B1, SCD, PLPP3, ARI1A, TRAD1, HSPB6, COPE, IF1AY, CCS, DVL2, TOR1A, P5I11, ANM5, NHRF1, TPP1, TCRG1, NRP1, PSA7, SCAM3, ML12B, HGS, MYPT1, GAK, HNRDL, XPO1, ZN609, SC16A, PLXB2, SR140, SET1A, SYNJ2, PUR4, NPC1, TBX3, ARC1B, ARPC2, ARPC3, TIF1A, NKRF, CASC3, RER1, SURF4, SPTC1, SPTC2, OGT1, PMM2, PPM1G, SHIP2, EIF3D, EIF3H, IPO8, STX7, MOT4, MRP4, P4HA2, NUP42, ARPC5, CLOCK, TPSN, KDM6A, DHX15, ASAP2, PRP4, SERA, DC1L2, PSMD3, RFOX2, PAPS1, SNUT1, DAPK3, TGFI1, CTIF, M3K7, MCA3, HNRPR, PRPF3, TXNL1, TPD54, EMC8, SYNJ1, IF4G3, ARK72, E41L2, B4GA1, FOXO3, RNF13, DENR, XPOT, PLRG1, ZN207, NDUA2, BUB3, ACTN4, AP1G1, SNG2, SGTA, SYNC, KRT86, MYO1B, CPSF5, LANC1, U3IP2, AKAP8, NRDC, CALU, EDIL3, SAHH2, EXTL3, MED14, SMCA5, TRAK2, GANP, FADS1, PEPL, NRP2, MA1A2, ACSL4, SNX3, OGA, HNRPQ, TPST1, NEMF, GREM1, PLOD3, SEP15, TSN3, PLIN3, MAFK, UGDH, CTND1, DPM1, USO1, RT14, H2B1K, PRAF2, DKC1, IF2P, DNJA2, BRD4, MITF, SRGP2, WDR1, ROCK2, CPNE3, PHF2, T22D2, PP6R2, CREST, ANR17, GGCT, NDUS2, ZPR1, DUS11, NIPS2, SEM7A, LX12B, TBCA, VATG1, H2AY, FLNB, NCOR1, NDUS6, CISY, SPAG7, SC22B, DEAF1, PR40A, MPPB, PSIP1, ERLN1, CLN5, CLD11, SF3B1, CSDE1, WBP4, PRKRA, LYPA1, ERAL1, U520, TIPRL, CRTAP, EIF3G, EIF3J, CBR3, FACE1, IDHC, ATRN, STAM2, PIAS1, PIAS2, DNJC8, SPF30, ATP5H, TRIO, CBPD, KT33A, KRT34, KRT36, RL1D1, WFS1, CLPX, STC2, PDE5A, SRP72, DDAH1, B3GA3, MTA2, AL1A2, TOM70, TOX4, SC24D, PHF14, CSCL1, TOX, SUN1, ERLN2, PRP6, NFAT5, GLSK, SC31A, CSTN1, YIF1A, AGFG2, SCAF4, ZFPL1, ELP1, NDUB8, LETM1, ZRAB2, LC7L3, VAPB, MPZL1, NDUAA, FKBP9, 6PGL, PAPS2, SMC2, IPO7, AGM1, SVIL, AHSA1, PSMG1, NSA2, G6PE, SC24A, SC24B, ETHE1, ACSL3, CNOT4, EYA4, K2C75, OXSR1, AP2A1, WIZ, STAU1, BAG2, BAG3, AIFM1, LATS1, BPNT1, FADS2, DDAH2, ABHGA, TXD12, FBLN4, RECK, NUDT3, NDUBA, CLPT1, TOM40, ACL6A, CYB5, LDHA, DHE3, NB5R3, GSHR, COX2, PNPH, HPRT, AATM, EGFR, PGK1, C1R, PLMN, UROK, TPA, ANT3, A2MG, CO3, CO5, TIMP1, CYTA, RASN, LDLR, TGFB1, IGHG1, IGHA1, HLAB, CO1A1, CO3A1, CRYAB, K1C14, K2C6A, LMNA, APOA1, APOC3, FIBA, FIBB, FINC, AMBP, FETUA, FETA, VTDB, TFR1, TRFE, TRFL, CATA, RAF1, GBA1, FUCO, ALDOA, ANXA1, APOB, GCR, PRIO, SODM, OAT, HRG, THY1, K2C6B, K2C1, G3P, HLAA, CPNS1, AMYP, HSPB1, RPN1, RPN2, GNAI2, AT1A1, A4, ARGI1, APOD, ALDH2, ITB3, S10A8, PAI1, ADT2, IF2A, RLA2, RLA0, JUN, LA, HEP2, ITB1, K1C18, K2C8, CO5A2, DCUP, LCK, GELS, PTMA, CDK1, ATPB, S10A9, S10A6, IF4E, ENOA, G6PI, NPM, ITAV, HEXA, H2B1J, GDN, HYEP, LDHB, PGK2, PDIA1, H10, CATD, ANXA2, CAN1, TBB5, TRY1, PGS2, SAP, HEXB, CATL1, PROF1, APT, SYEP, CATB, HS90A, HNRPC, LAMB1, TPM2, FUMH, TSP1, SP1, CO1A2, ANXA6, 4F2, PFKAM, HS90B, SRPRA, ASNS, ODPA, CY1, RU2B, RU17, ITA5, NFIC, VIME, A2AP, RS17, GNAI3, ANXA5, K1C16, STS, RSSA, CD63, FGF2, GSTP1, RU1C, LEG1, DHPR, HMGB1, SPRC, TPM1, CLCA, ANXA4, CN37, PGFRB, DLDH, ROA1, PARP1, UCHL1, LKHA4, ALDOC, CO4A, H2AZ, UCRIL, ATX1L, POTEI, ZN865, ZN888, INAM2, HS71A, GLI2, LYAG, HLAC, H14, ODP2, THIO, PPGB, MGST1, KAP0, ESTD, CH60, CLUS, BIP, LAMC1, PPAL, HSP7C, UMPS, ODPB, ODB2, PYGB, RALA, LAMP1, ACADM, FGFR1, TOP1, G6PD, PYC, C1TC, MPRI, ADHX, SRF, PABP1, PCNA, HARS1, CO6A1, CO6A2, CO6A3, ADT3, IMDH2, TPR, ACTN1, PEPD, XRCC6, XRCC5, COX41, IFM1, LAMP2, RINI, CSPG2, EF2, K1C13, K2C5, PDIA4, P4HA1, TCTP, HLAE, PLST, ACPH, ETFA, KAP2, ENOB, CD59, MIF, CD99, GLU2B, FPPS, NID1, AK1A1, KPYM, ENPL, RSMB, IDE, PO2F1, HNRPL, SYDC, PLAK, ALDR, AMPN, PVR, ERF3A, B4GT1, EZRI, ATF2, UCHL3, FOSL1, FOSL2, NDKA, GNS, ZEP1, ARSB, RS2, ITF2, TFE2, DESP, RFA2, CD44, CBR1, CREB1, PGFRA, BGAL, H15, H13, H12, NCPR, AT2A2, FAAA, STMN1, NAGAB, AATC, KPCA, JUNB, ITA2, ASM, UBF1, HXB6, ATF7, CAN2, DESM, CEBPB, PYRG1, EGLN, DDX5, LEG3, IBP3, TCPA, PTN1, IBP2, RL35A, ARF4, RL7, EGR1, VINC, SON, RL17, PGAM1, RCC1, ATF1, NUCL, HXK1, E2AK2, SPEE, CSK22, RXRA, ITIH2, ITIH1, NFKB1, FST, IF2B, ANXA7, CD22, RAB3B, RAB5A, PSB1, MPRD, COX5A, LMNB1, HXA5, IMDH1, CO5A1, FILA, GSTM3, VATB2, CSRP1, FLNA, ACOHC, 5NTD, VDAC1, PGS1, SDHB, COMT, TGM2, PIMT, FBRL, PUR2, PUR6, SCP2, UBA1, NDKB, SPR1B, SPR2E, ROA2, RFX1, CBL, IBP4, QCR2, TGM1, FBLN1, PGH1, ITA6, SFPQ, PPIB, MAOM, SYWC, RS3, SAHH, COF1, IF4B, AT2B4, DGKA, GATA2, CPT2, KTHY, RIR1, EF1B, IBP5, PPAC, THIL, TENA, MYL9, RPB1, PAR1, MCM3, NFYB, ZA2G, THTM, RS12, TYY1, DNJB1, DNJB2, ATPA, PSA1, PSA2, PSA3, PSA4, ITA3, MOES, DDX6, DNMT1, U2AF2, RL13, IVD, HMGB2, PTBP1, SYTC, SYVC, EF1G, FKBP2, STOM, 1433T, DPP4, ARNT, RL10, APEX1, PYR1, CALR, MAP4, CALX, PSB8, PSA5, PSB4, PSB6, PSB5, ABCD3, ITPI2, LYOX, ELK4, NDUS1, MK01, GRN, AMPL, TPP2, IMPA1, CCN2, EPHA2, EPHB2, TKT, SPB3, RBMS1, PML, EF1D, MARCS, GNA11, AL4A1, ERP29, PRDX6, BLVRB, PRDX5, PRDX3, ATPD, RL12, ECHM, PEBP1, PDIA3, 2AAA, CDC27, T4S1, NMT1, PURA2, UFO, AMRP, TSPO, PUR8, CLIP1, SORCN, CTR1, AL1B1, RPB2, LCN1, SDHA, GDIA, S10A7, METK2, RIR2, TIA1, ZEP2, DNJA1, AKT1, QCR1, 3HIDH, PUR9, HNRH3, HNRH1, CASPE, 1433B, STIP1, S10AB, PRDX2, P5CR1, GBP1, ELF1, RL9, KINH, DUT, MRP1, MCM4, MCM5, MCM7, CD68, GLYM, HSP74, PROF2, CTNA1, CTNB1, PHB1, RADI, RFA3, RFC2, RL22, CBS, K1C9, FBN1, FBN2, IRS1, MYH9, MYH10, COPB2, ADDA, BASI, TIMP3, FUS, NU214, DEK, SOX5, GLRX1, PRS7, MP2K2, ATPG, RL4, LONM, PGM1, PP1G, PEDF, ODO2, GPX4, SRP14, NUP62, TAGL2, TALDO, ETFB, RBMX, VKGC, VATA, HSPA9, IF4A3, RS19, RL3, COIA1, OST48, AN32A, FEN1, CAPG, IL6RB, TXLNA, TCPZ, NNMT, RL13A, STAT3, MDHC, MDHM, ECHA, IF2G, ETV6, WNT5A, BUD31, NAA10, CSK, GARS, SYIC, KPCI, LAP2A, LAP2B, MTREX, HUTH, CASP3, RS27, HELZ, LPPRC, THIM, RL35, PCP, ECE1, LIS1, MUC18, MATR3, SSRA, RANG, PRS6B, PPIC, VDAC2, CBX5, UBP5, RAGP1, RECQ1, NOP2, CRKL, BAG6, NSF, NOTC1, MP2K3, RL27A, RL5, RL21, RL28, RS9, RS5, RS10, MAP1B, YAP1, UTRN, IQGA1, STT3A, CAZA2, CAPZB, SYQ, RL29, LEG7, TISD, ATPO, LIMS1, PPCE, MAOX, RFX2, RFX5, SOX9, COPD, LSS, PP2BC, CD151, PRC2A, TCPE, NEST, HSP13, KC1D, IDHP, PIPNB, PAXI, NR2C2, AL9A1, RL34, LMAN1, NASP, FAS, CDK8, TCPG, EFTU, AL7A1, SRP09, PCY1A, SYAC, SYCC, SYSC, MA2A2, PSB2, MCM2, COMP, ACADV, YLPM1, TMEDA, RBM25, NUMB, HINT1, NU153, RBP2, GSK3A, TAF6, 5NTC, SEPP1, GUAA, IDH3A, MMP14, GDIB, EMD, CPT1A, SERPH, PDLI4, F10A1, VASP, DYN2, MAP2, TOB1, CDK9, BCAM, PPT1, RL14, TCPQ, TCPD, ANX11, PAPOA, RAB5C, RAB7A, RAB9A, DAP1, RT29, SMCA4, IDH3G, SSRD, BAP31, DHB4, PSMD7, SPHM, CLCN5, KS6A3, HDGF, LUM, CNN1, NDUA8, ROA3, 6PGD, HNRPM, IMA1, NCBP2, GDIR1, AGFG1, HNRPF, STAT2, TF2AA, MSH6, ZN143, RBM5, SPSY, THOP1, CAZA1, CRIP2, NUP98, BIEA, TXTP, PPP5, ACLY, FOSB, MAP11, SUCA, PGTB2, COPB, COPA, CATC, RPAB4, MOT1, SC24C, TCP4, SYRC, ATX1, ATN1, CA2D1, SYYC, UBP14, AAKG1, HSP72, BCAT1, AT1B3, RD23A, RD23B, ANAG, KAD2, P5CS, SNAA, IF5, PSMD4, DRG2, XPO2, TERA, ECHB, MANF, AF10, DSRAD, LAMB2, CAD11, CAD13, SEC13, NH2L1, PSA, HNRH2, SCOT1, EIF3B, BID, SYMC, ITA1, MAZ, ATP5E, ATP5I, HDAC4, IF6, CTBP2, BCAR1, TMM33, PCBP3, NU107, MTPN, ARPC4, TPIS, EIF3E, SC61B, PP4C, GBRL2, MYL6, ACTB, IF4A1, RS20, PRPS1, PSA6, S10AA, CDC42, DEST, SPCS3, SRP54, RAB2A, RAB10, UBC12, UBE2K, UBE2N, RAB14, ARP3, ARP2, ARF3, ABCE1, RAP1B, RS3A, RL26, RL15, MGN, RL27, VA0D1, RL37A, RHOA, CH10, S61A1, LYSC, STXB1, B2MG, DAD1, NPC2, COPZ1, SUMO2, UFM1, NTF2, HNRPK, 1433G, RS7, PP1A, PP1B, PRS4, PRS8, RS8, RS15A, RS16, 1433E, RS14, RS23, RS18, RS29, RS13, RS11, RUXE, RUXF, SMD1, SMD2, PRS10, RL7A, RB11A, ERF1, CNBP, YPEL5, RS4X, PP2AB, ACTA, RL23A, RS6, H4, RAB1A, RAN, RL23, RS15, RS25, RS26, RS28, RS30, GBB1, RBX1, GBB2, RL30, RL31, RL10A, RL32, RL11, RL8, PPIA, FKB1A, RS27A, GRB2, RAC1, AP2B1, VAMP2, GNAS2, 1433Z, SUMO1, RL38, SKP1, RS21, IF5A1, RACK1, ACTG, ACTH, PP2AA, YBOX1, SC11A, CSK2B, TPM4, UB2L3, EF1A1, TBA4A, TBB4B, CSK21, HBB, NOMO3, HBA, F193A, IGBP1, SHPS1, SERB, PITX1, IF4G2, GTF2I, CNTP1, PHC1, TCPB, RAE1L, GSTO1, PRKDC, MAP1A, NUCB2, DCD, SSBP2, SARNP, RL24, RL36A, SMAD3, ERH, RHOG, RL19, SRSF3, FOXK1, DAB2, FBLN2, PGBM, XIAP, RBM10, RBM3, CYC, S25A3, VIGLN, PURA, BORG5, CLH1, FKBP3, REEP5, DHSO, HNRPU, U2AF1, SPTB2, TIAR, SET, SRSF2, FOXK2, RUNX1, AMPD2, FABP5, CAP1, K22O, HMCS1, LAT1, EXOSX, OTUD4, PFKAP, EWS, PLCB3, ROR2, TAGL, MEF2A, ODO1, DSG1, SP3, RL18A, FKBP4, PLOD1, NUCB1, RL6, TOP2B, CREB5, AKA12, M2OM, DYST, CAV1, KMT2A, LMNB2, UPAR, TF65, GLGB, IF4G1, K1C17, NOTC2, TLE4, LGUL, AK1C1, 1433F, PLP2, CSTF1, UBE3A, FAK1, CALD1, COEA1, PUR1, GFPT1, SUH, MEF2C, GABPA, GABP1, PRDX1, RL18, CKAP4, ZO1, KHDR1, BAX, KLC1, ACK1, LRP1, SRSF1, RHG01, TLE5, TGM3, PP2BA, DHX9, QOR, GOGA3, GOGA2, LG3BP, DSC1, CD47, PPID, SSRP1, NSUN2, RBBP4, NCBP1, MGT5A, EP300, AHNK, IFFO1, INCA1, NEXN, MGAT2, GALT2, GALT1, AP1B1, CPSF1, MPPA, SCRN1, NU160, SCAP, SRBP2, TWF1, ASPH, FBLN3, TROP, BPTF, FSTL1, GRSF1, NFIA, SFSWA, SF3A3, SEPR, TP53B, AIMP1, ILF3, LMAN2, EPS8, TRAP1, TAF10, ECH1, STRN3, CDC16, FLII, CSN1, MTAP, AAPK1, EI2BE, TADBP, ROA0, AIMP2, PRDX4, PAK2, CBX3, PSMD2, TBX2, SRSF9, SRSF5, SRSF6, TIF1B, G3BP1, SG2A2, PTK7, PABP4, EIF3I, CTBP1, UB2V1, DC1I2, ILK, NNTM, TCOF, S39A6, SF3B2, ADAM9, TMED1, LSAMP, MYO9B, GAB1, SMAD4, PICAL, MAMD1, SQSTM, TBB3, ASAH1, PRP4K, PIN1, RIPK1, HDAC1, KCC2D, DCTN2, TRA2A, SNX1, CUL1, CUL4A, CUL4B, DYR1A, FCL, FHL1, FHL3, CD166, RUNX3, SPTN1, PKP1, DX39B, BLMH, TBB2A, IDI1, RUNX2, PEBB, NFYC, CDK13, CKAP5, COTL1, HNRPD, SCRB2, NID2, DAG1, VEZF1, EIF3A, UBP2L, SCRIB, GIT2, MLEC, TTL12, FHL2, DPYL3, DCTN1, DYHC1, SRC8, RCN2, TRI25, FLNC, FKBP8, FZD2, GAS6, CAPR1, RBM39, FAT1, SEM3A, MCM6, DSC3, PUM1, EPN4, SMC1A, RRP1B, NCOA6, GSE1, GANAB, 2A5D, LBR, MVP, LTBP1, LTBP2, MEF2D, CHD4, LASP1, ARI5B, PTGR1, NC2A, ZN638, IMB1, NOLC1, NUMA1, K2C72, CND2, PSMD6, SEPT2, SART3, CND1, U5S1, R3HD1, TRAM2, RB3GP, SYK, IF4H, BRD3, POSTN, OXA1L, EEA1, PDIA6, PA1B3, PCOC1, PLEC, IPYR, TEBP, NONO, RBBP5, RCN1, PCBP1, PCBP2, RHEB, DHC24, SF3B3, PUM3, DLGP5, RSU1, CNN3, SF3B4, SF3A2, RBMS2, SC23A, SF3A1, CDSN, SSXT, BGH3, NCOA2, TRAM1, SF01, MED1, JHD2C, TRIP6, MARE1, T22D1, ELAV1, ELF2, NAB2, MYLK, TAB1, AAAT, MLF2, ITSN1, TBCC, VAS1, ZFHX3, ZYX, ETFD, SEPT7, ADRM1, UAP1, IBP7, AINX, LAMA4, EXT1, PSMD5, CSRP2, DDB1, CDC37, DPYL2, KCC4, RBBP7, TAF9, ECM1, SRSF7, CPSF6, DREB, NRF1, FSCN1, IF16, DECR, GLO2, H2A2C, NDUA9, PCKGM, DDR2, HCDH, UGPA, TRXR1, MIC60, CPEB4, HNRL2, INF2, RFX7, QSER1, AAK1, PRSR3, QRIC1, LRRF1, P3H1, MA7D1, AL1L2, K2C71, TIM50, FUT11, TB10B, ANO6, FIL1L, SVEP1, AMOT, EPC2, FND3B, CRTC2, QORX, DHB12, CYBR1, IN80D, PAN3, YIF1B, F133B, ZN326, FILA2, MEST, HERC4, PIGG, BCORL, FA76B, TGO1, PREP, PRC2B, RRP12, TOIP1, PCID2, EOGT, TTC38, TM201, NU188, HP1B3, CE170, ZN362, FBP1L, FKB15, ZEP3, ZC3HD, LRIF1, UBR4, STXB5, RHG21, RM02, UBAP2, KPRP, KPLCE, RBM26, WLS, TUT4, H37, NT5D1, AHDC1, MRCKA, RPRD2, MYOME, ZN318, BROX, TASO2, PEAR1, MBNL2, ZMYM4, FHI2A, CD276, TSH3, TENS2, KANK2, AR6P4, OLM2A, HGNAT, ARID2, TENS3, CRBG3, RHG17, BICRL, AMERL, ATLA3, NXN, IQEC1, VASN, CPIN1, PTRD1, ELP2, LTOR1, TWF2, ADNP2, K2C80, LIN54, SCMC1, TM214, PPR18, CAVN1, CDC73, RM14, TATD1, EDC4, PRP8, SCYL2, GOLM2, NFRKB, RL22L, ALKB5, NCEH1, MDEAS, ZC3HE, OGFD3, LARP1, C1TM, FIP1, CRTC1, CSPG4, ARAID, LRSM1, TMCO3, LCLT1, CREL2, PAMR1, FAT4, MCAF1, CP2R1, PACS1, BCOR, MPRIP, HACD2, GGYF2, HSDL2, KAT3, CERS6, THSD4, MEX3B, ZN574, RN111, BAKOR, SRCAP, RIPR1, YJ005, TSH1, BGIN, UBN2, TMTC3, RAPH1, MOB2, IKIP, H32, LARP4, TBA1A, RS27L, HAKAI, ASXL2, CCD80, SPT6H, SND1, DDX46, 5MP2, DHX30, EIF3M, CYFP1, TAOK1, KDM3B, PHF5A, ERMP1, ZCCHV, NUP54, POGZ, MON2, K1C27, NUFP2, MAVS, CLAP1, DHX29, PLGT3, LIMS2, HDGR2, DCXR, EMSY, RAI1, I2BP2, RBBP6, SH3R1, RABEK, HUWE1, YTHD3, K2C1B, KHDC4, TMED4, COXM1, CENPV, GALT5, PHLB2, GLT10, MYPN, SYVN1, ACOT1, PABP2, LDB1, LYRIC, ZN598, KTN1, BCL9L, PHLB1, LUZP1, GP180, CAND1, HOOK3, TSYL5, CARM1, PRSR1, MEX3D, DDX42, P66A, HORN, RB6I2, RELL1, PLD3, FOXP4, P3H2, P3H3, CCD50, SPA12, CTL2, WDFY1, TEX2, NXP20, MGAP, Z3H7A, CHERP, ANKH1, SUGP1, CCAR1, DJC10, MSRB3, PLPL6, MILK2, SPB1, SMAP1, P20D2, PHAR4, ZNF34, ELMD2, DCP1B, XRN1, MAML2, ABI1, SELH, SPART, RPTOR, CCAR2, SIX5, NUP93, K2C78, ZCH24, LTBP4, ASCC3, SYNPO, FNBP4, AFAP1, CERS5, NKAP, CPSF7, ARFG2, ARFG1, LRC17, EMC1, ENAH, RM43, YP023, ANR52, GT251, SUMF2, SUMF1, PGLT1, PCYXL, TXND5, SCPDL, SERB1, LEMD2, NECP1, FA98A, CHSTE, APCDL, RN214, LS14A, TNR6A, PHC3, ABCF1, SP20H, ZNT7, NAV1, VP37A, FBX22, PCAT1, ZMIZ2, ARI1B, NUP35, NEUA, PLBL2, MFSD8, NEDD1, ZNT5, G45IP, SMRC2, NPL4, PUM2, KISHA, RDH11, STT3B, SPP2B, HM13, ALMS1, NEK9, PKHO2, DDX54, DLG5, NEK7, GPX8, GEMI5, IPO4, ZN384, SETD7, F222B, SMAP2, CHUR, CEMIP, PDC6I, FBLI1, PHLA1, NUDC2, SCFD1, POF1B, LMO7, ATX2L, SELM, PALLD, SREK1, PSPC1, P66B, AUTS2, ZCH14, IRGQ, DDB2, ASM3A, DDX1, S39A7, FAM3C, GBF1, SMG7, RTF1, NICA, TM9S4, TM131, RT27, F168A, PHF3, MAML1, HS105, LAR4B, GCN1, ZN516, NU205, PXDN, AN32B, NECT2, TFG, ARC1A, STAM1, REQU, SC65, CBP, GGH, NELL1, DDX17, TANK, CELF1, GSLG1, RENT1, CSN5, GATA6, SMRC1, FUBP2, TNPO1, ARHG2, UBP13, USP9X, UBP7, LPP, RBPMS, SMCE1, CAVN3, MYDGF, YIPF5, PIGT, RL36L, ZN622, WBP2, NCLN, MRTFA, ERGI1, RFT1, FERM2, FUBP1, TTC17, LRC59, PBIP1, FKB10, AKTS1, PF21A, EFHD2, GALM, INT12, CTHR1, KLH29, OPTN, AP2M1, RCN3, REPS1, KBP, DAZP1, RBM33, CK024, ZN503, I17RA, OTUB1, PGM2, CERS2, ZC21A, P121A, PDLI5, CREL1, ERO1A, PGAM5, FUBP3, SUCB2, GMPPA, KIRR1, WNK4, TMX3, CHAP1, ZFP91, PDLI2, ZC3HA, CLP1L, SCYL1, Z512B, CNDP2, ZFR, B3GT6, EP400, VATG3, PRRC1, ZN512, NOL4L, MBOA7, DOCK7, TOX2, SPB12, ARHGH, RBM14, QKI, LENG8, GSDMA, PP1RA, S38A2, VPS35, PHF12, SNX18, CIC, MED15, ERGI2, NUDC1, ERBIN, PIGS, WRIP1, RANB9, CLCC1, UBL7, MYADM, SIN3A, RUFY1, MINT, MMS19, UHRF1, NIBA2, HTF4, TBCB, NELL2, PSB7, CNN2, SEC62, PCY2, K1C12, CDC5L, PSMD1, SDF2, PARK7, EYA3, NEUR1, VAT1, LGMN, PLIN2, NUP88, MNT, SCAFB, DNJC7, PHB2, RGPD5, PITX2, ATX2, HCD2, COCA1, SMAD5, ROAA, LITAF, NP1L4, ACON, TM9S2, TS101, CPNE1, TCPH, EBP2, ANM1, SH3G1, CND3, DDX50, MEP50, VKOR1, TBA1C, SELS, MBB1A, COR1B, TXD17, VPS25, RM45, MXRA8, CAB45, ADPGK, PELO, ERP44, LXN, ESYT1, CNPY3, LIMD2, CSN4, WAC, DIDO1, DCTN5, AASD1, TMM43, TBCD, TBB6, HNRL1, HTAI2, DDX23, DHRS6, CA050, WDR18, THUM3, RN126, TBB2B, TM109, TMED9, NUP58, ELOV1, THIC, REPI1, RBM4, SFXN3, SSBP3, NAA15, KLF16, SRRT, TGT, HDHD5, EIF2A, CADM1, MK67I, YTHD1, WNK3, NEUL, RM37, GTPB4, UACA, RAB34, TBL1R, UBL5, EDEM3, API5, K1C23, UNK, FTO, TB182, AMRA1, TTYH3, ADSSP, PITH1, SKI8, SLIRP, WWTR1, TWSG1, NAA50, MKRN2, PAIP1, LSG1, NAT10, XRN2, SP130, BRD8, DDX47, MAGT1, LMA2L, SIL1, I2BPL, NUCKS, EHD4, VPS16, SH3L3, EPC1, SLK, S38A1, PDCL3, RBGPR, ADNP, DPH5, HIPK2, FOXP1, CK068, BOLA2, TMX1, NELFA, GCP60, PTN23, UN45A, OFUT1, WNK1, AMPB, GAPR1, ZHX3, EHD1, ZN768, SG196, CC134, PEAK1, AAMDC, WDR26, PKHG2, ZN703, ACAD9, GORS2, JUPI2, SYP2L, SFXN1, FKRP, ELP3, KT3K, FBRS, MLXIP, PKHA5, GBB4, XPO5, GRPE1, MYG1, RC3H2, TENS1, TAF9B, EMAL4, GLOD4, ZBT20, NCOA5, TANC2, MOV10, ZN532, ZN160, TNR6C, CHD8, SDF2L, CD248, SEC12, S10AE, AT131, RM47, CHM1A, TM9S3, APMAP, TRXR2, ARFG3, RAB18, VTA1, CD320, UBN1, DCP1A, TF7L2, RTN4, NIT2, NECT3, EXOS3, ANLN, XPP1, GEPH, PDLI7, DDX21, SAR1A, SIAS, SHLB2, MBNL1, 4ET, AAAS, STRN4, UBQL4, HEBP1, ABC3C, PHP14, F1142, TM7S3, KRT84, SYFB, SYIM, BMP2K, ATG3, SMC4, SAC1, OLA1, ZF64B, RBM12, ABHDA, SHFL, T106B, SPS2L, DD19A, TBC23, CC50A, SEP11, MED17, DJC11, INT7, ATD3A, ARL8B, DDX18, RT18A, NLE1, ZN358, RBM22, ARGL1, RM22, CZIB, GEMI8, F120C, OCAD1, IMPA3, MIC19, CKLF6, P4HTM, TOR4A, TRM1, CARF, DPP3, DDX56, BCLF1, TAB2, M3K20, UGGG2, UGGG1, CDK12, TECR, ERAP1, GGA3, F120A, GDE1, IF2B1, MAT2B, MYOF, ITSN2, BICRA, EHD2, CNOT2, CALL5, CRIM1, HACD3, KCMF1, RAI14, RM27, SENP1, SH3B4, PHRF1, RCC2, SLAI2, TEN3, CT2NL, RRBP1, SYLC, RBM27, KANL3, ANFY1, RERE, SUCB1, LRIT1, DPM3, SAE1, COPG2, MRC2, GBG12, DHCR7, DKK3, VPS29, EIF3K, GRHPR, CATZ, SAE2, NXF1, B4GT7, PEF1, FBLN5, CLIP2, ZO2, DNJB4, ADDG, ABCF2, TES, LIMD1, Z3H7B, TCF20, SUN2, LIMA1, SRP68, CHRD1, SEPT9, UBQL2, PCYOX, DDX20, ATS1, DPP2, CEIP2, NB5R1, S30BP, MED13, PUF60, NRBP, VATH, CBPA4, XPO7, BAZ2B, SIX4, GGT7, MSRA, NAGK, DBNL, DCTN4, CDC23, STML2, DBR1, AKA11, GMEB2, CPSF3, PARP4, RALY, ITA11, NUP50, ZHX1, ZN346, DNPEP, MCTS1, S39AA, ASAP1, MRTFB, HEG1, ZMIZ1, PRR12, YETS2, HECD1, ZMYD8, COR1C, NOTC3, SPAT2, PPT2, PRP19, UBQL1, SNX12, SYNRG, G3BP2, CHIP, PACN2, MAGD2, SNX6, PSD13, FAF1, PPIE, CSN3, MINP1, S12A4, MACF1, SCAF8, TRI33, TNR6B, SMAG1, CE164, FOXJ3, SHAN2, COR2B, SRRM2, PA2G4, ZN148, RTRAF, RUVB2, CLC11, EIF3L, PLAP, RUVB1, NUDC, VDAC3, COF2, ERGI3, SYFA, DRG1, NCKP1, PLAK2, ZN652, MA2B2, OFUT2, FND3A, E41L3, R3HD2, RRP44, LRCH1, GSTK1, DDX52, RT07, TMA7, MAN1, CHSP1, TR150, NOP58, ZN281, SGT1, ACOT9, NOSIP, MEMO1, SHLB1, LC7L2, SBDS, TMED7, ORN, UFC1, STRAP, RTCB, ZN330, RL36, SAMH1, TLN1, ARIP4, MTCL1, TCAF1, UBP15, LOXL2, CRBG1, HYOU1, TBL2, KLF12, ARI1, LSM4, RN114, PRC2C, RBM7, YTHD2, SP16H, SRPRB, MRCKB, ZN706, MAGD1, SNX5, HCFC2, FHOD1, SERC, NCOR2, NPTN, CBPQ, SOX21, RT18B, COPG1, FKBP7, GMEB1, CLIC4, EMIL1, MTCH2, 5MP1, DC1L1, EPN1, NEMO, IF2B2, ZNT1, NCOA3, NUMBL, PIAS3, ZHX2, S23IP
Species: Bos taurus, 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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Hou C, Deng J, Wu C, Zhang J, Byers S, Moremen KW, Pei H, Ma J. Ultradeep O-GlcNAc proteomics reveals widespread O-GlcNAcylation on tyrosine residues of proteins. Proceedings of the National Academy of Sciences of the United States of America 2024 121(47) 39531497
Abstract:
As a unique type of glycosylation, O-linked β-N-acetylglucosamine (O-GlcNAc) modification (O-GlcNAcylation) on Ser/Thr residues of proteins was discovered 40 y ago. O-GlcNAcylation is catalyzed by two enzymes: O-GlcNAc transferase (OGT) and O-GlcNAcase (OGA), which add and remove O-GlcNAc, respectively. O-GlcNAcylation is an essential glycosylation that regulates the functions of many proteins in virtually all cellular processes. However, deep and site-specific characterization of O-GlcNAcylated proteins remains a challenge. We developed an ultradeep O-GlcNAc proteomics workflow by integrating digestion with multiple proteases, two mass spectrometric approaches (i.e., electron-transfer/higher-energy collision dissociation [EThcD] and HCD product-dependent electron-transfer/higher-energy collision dissociation [HCD-pd-EThcD]), and two data analysis tools (i.e., MaxQuant and Proteome Discoverer). The performance of this strategy was benchmarked by the analysis of whole lysates from PANC-1 (a pancreatic cancer cell line). In total, 2,831 O-GlcNAc sites were unambiguously identified, representing the largest O-GlcNAc dataset of an individual study reported so far. Unexpectedly, in addition to confirming known sites and identifying many other sites of Ser/Thr modification, O-GlcNAcylation was found on 121 tyrosine (Tyr) residues of 93 proteins. In vitro enzymatic assays showed that OGT catalyzes the transfer of O-GlcNAc onto Tyr residues of peptides and OGA catalyzes its removal. Taken together, our work reveals widespread O-GlcNAcylation on Tyr residues of proteins and that Tyr O-GlcNAcylation is mediated by OGT and OGA. As another form of glycosylation, Tyr O-GlcNAcylation is likely to have important regulatory roles.
O-GlcNAc proteins:
NPIA9, RBM47, E2F8, SBNO1, CNOT1, P121B, HMX3, RHG32, P121C, NACAM, PDLI1, TAF4, DNM1L, PLOD2, CAC1A, ARI1A, CDKA1, NCAN, ABLM1, KMT2D, HGS, MYPT1, ZN609, SC16A, SET1A, TRNK1, OGT1, TNC18, P4HA2, NUP42, KDM6A, ASAP2, SI1L1, RFOX2, PRPF3, SYNJ1, IF4G3, MTG8R, E41L2, PLRG1, ZN207, BUB1, MYPT2, KPRB, ZN292, SI1L3, GANP, TBC12, OGA, KLH41, PLIN3, MAFK, ABCC9, TBL1X, MITF, SRGP2, SLIT3, T22D2, PP6R2, ANR17, LIPA4, ZN217, NCOR1, U520, RASF9, DYSF, TOX4, SC24D, FCSD2, PCF11, NFAT5, SC31A, AGFG2, SCAF4, ELP1, MRP6, SVIL, SC24A, SC24B, CNOT4, ZMYM6, BAG3, LATS1, LMNA, CATA, GCR, CPNS1, HSPB1, RPN1, RLA2, ITB1, K1C18, K2C8, NPM, CATD, GP1BA, SAP, CATB, SP1, PTPRC, NFIC, VIME, K1C19, SNRPA, ATX1L, LAMC1, ARY2, SRF, PABP1, TPR, SKIL, ENPL, PO2F1, B4GT1, ATF2, FOSL1, FOSL2, GNS, ZEP1, RS2, TFE2, CREB1, HXB6, ATF7, EGR1, SON, ATF1, NUCL, ICAL, OSBP1, RFX1, CBL, NFYA, IF4B, GATA2, ARNT, MAP4, GRN, CDC27, CLIP1, HXC9, ZEP2, ELF1, NU214, NUP62, CUX1, STAT3, BUD31, HNF4A, LAP2A, MTOR, PCP, MATR3, GATA4, UTRN, STT3A, RFX5, SOX9, PRC2A, NEST, HSP13, NASP, CDK8, YLPM1, NU153, RBP2, TAF6, EMD, PPT1, PAPOA, FXR2, KCNQ1, MYOM1, AGFG1, HNRPF, NUP98, SMTN, SC24C, ATX1, RD23B, AF17, DSRAD, MAZ, CTBP2, CSK21, F193A, SERB, IF4G2, GTF2I, PHC1, NGAL, KGD4, SARNP, FOXK1, DAB2, PGBM, RHG04, SPTB2, FOXK2, OTUD4, EWS, SP2, NRG1, SP4, SP3, PAX2, KMT2A, IF4G1, NOTC2, PTN12, GABPA, GABP1, CKAP4, ZO1, ACK1, TLE5, EP300, AHNK, ASPH, TROAP, BPTF, NFIA, TP53B, FOXC1, TAF10, PICAL, MAMD1, SQSTM, RIPK1, STIM1, CUL4B, NFYC, CDK13, CKAP5, VEZF1, UBP2L, GIT2, SRC8, CAPR1, PUM1, MDC1, EPN4, TTLL4, RRP1B, NCOA6, GSE1, MEF2D, ZN638, NUMA1, HECAM, R3HD1, DAZP2, KIF14, PDK3, EBP, NONO, LRC41, RBMS2, SSXT, NCOA2, SF01, MED1, JHD2C, T22D1, ELF2, TAB1, NCOA1, TBCC, ZFHX3, ZYX, CCDC6, TAF9, DREB, NRF1, GEN, RFX7, QSER1, AAK1, PRSR3, RGPA2, QRIC1, TB10B, TPRN, FIL1L, CRTC2, IF44L, YIF1B, NOM1, BCORL, DDX6L, BRD10, SH319, TGO1, PRC2B, CE170, ZN362, DCST2, ZEP3, ZC3HD, CSCL2, LRIF1, RHG21, UBAP2, RBM26, CF163, AHDC1, RPRD2, ZN318, TASO2, ZMYM4, PAPD7, TENS2, KANK2, ICE2, ARID2, TENS3, USF3, ANR40, BICRL, SDE2, NIPBL, LIN54, TET2, ZNT6, EDC4, SCYL2, TTC27, NFRKB, RSBNL, TT23L, NCEH1, MDEAS, ZC3HE, FIP1, CRTC3, SAS6, ARSK, MCAF1, BCOR, GGYF2, AEBP2, BNC2, NFXL1, CO039, SRCAP, UBN2, TM1L2, RAPH1, HAKAI, ASXL2, SPT6H, BOP, KDM3B, ZCCHV, NUP54, RGPD4, POGZ, ZFY16, NUFP2, MAVS, EMSY, RAI1, I2BP2, SDK1, SRGP1, RBBP6, SH3R1, YTHD3, CENPV, KAISO, MYPN, LDB1, LYRIC, ABCAC, BCL9L, CASZ1, TSYL5, CARM1, PRSR1, DDX42, CACL1, P66A, HIPK1, RB6I2, AHNK2, NAV2, GID4, ARI3B, TEX2, MGAP, Z3H7A, ANKH1, SUGP1, CCAR1, MILK2, GAR4, TBC21, PHAR4, XRN1, MAML2, SPART, NUP93, DOCK4, SYNPO, FNBP4, AF1L2, TAB3, NOPC1, IGSF1, ARFG1, ENAH, TM102, WDR31, MISSL, TNR6A, PHC3, SP20H, NAV1, VP37A, KMT2C, ARI1B, NUP35, KNL1, O52H1, PLBL2, NEDD1, SMRC2, TBC15, ZN507, ALMS1, DLG5, FNIP1, ZN384, WIPF2, PYRD1, D42E1, PCNP, LMO7, ATX2L, P66B, BBX, ELYS, ZCH14, PIEZ1, GBF1, RTF1, PHF3, MAML1, ZN592, LAR4B, SORL, TFG, TAF4B, RREB1, CBP, DDX17, GPKOW, M4K1, FUBP2, LPP, WNT2B, P33MX, ACD, FKB10, TTC28, PF21A, ZN669, ZN501, REPS1, RBM33, CERS2, GWL, P121A, CC120, PDLI5, SENP5, VCIP1, ZN462, CHAP1, LCOR, Z512B, ZFR, EP400, BMF, PRRC1, NOL4L, RBM14, QKI, PHF12, CIC, MED15, ERBIN, SIN3A, MINT, HTF4, EYA3, DNJC2, ELF4, RGPD5, ATX2, TS101, ARI3A, TEP1, DPH2, GPTC1, WAC, DIDO1, HNRL1, CP250, NUP58, RBM4, UBP26, YTHD1, BCDO2, B2L14, AMRA1, CE295, TANC1, ZC12C, CEP44, SP130, BRD8, I2BPL, RBNS5, EPC1, ADNP, HIPK2, ZN106, PTN23, WNK1, ZHX3, ZN644, HAUS4, SLAP2, YTDC2, PEAK1, PKHG2, ECT2, GORS2, MLXIP, PKHA5, CLSPN, CDHR5, RC3H2, TAF9B, NCOA5, TANC2, ZN532, TNR6C, CHD8, APMAP, UBE2T, DMAP1, UBN1, DCP1A, RTN4, PRD10, PDLI7, EI2BG, MBNL1, RNC, RBM12, INT7, CARF, ABI2, HXC10, TAB2, CDK12, GRHL1, ITSN2, BICRA, CNOT2, FLRT1, KCMF1, DAPLE, CHD7, F135A, RBM27, KANL3, RERE, LIMD1, TCF20, SUN2, UBQL2, TRPS1, S30BP, PUF60, NRBP, BAZ2B, SIX4, HOOK1, GGA2, TASOR, AKA11, GMEB2, PARP4, MYO15, MYH13, NUP50, ZHX1, CDV3, MRTFB, ZBT21, PLXB3, PRR12, YETS2, HECD1, ZMYD8, SPAT2, PRP19, SOX13, SCAF8, TRI33, TNR6B, SET1B, SATB2, SHAN2, HPS5, SRRM2, SCML2, R3HD2, ZN281, KLK5, ARIP4, MTCL1, RPGF2, CRBG1, HYOU1, PRC2C, RBM7, YTHD2, HCFC2, NCOR2, GMEB1, NCOA3, S23IP
Species: Homo sapiens
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Liu J, Hao Y, Wang C, Jin Y, Yang Y, Gu J, Chen X. An Optimized Isotopic Photocleavable Tagging Strategy for Site-Specific and Quantitative Profiling of Protein O-GlcNAcylation in Colorectal Cancer Metastasis. ACS chemical biology 2022 17(3) 35254053
Abstract:
O-linked-β-N-acetylglucosamine (O-GlcNAc) glycosylation is a ubiquitous protein post-translational modification of the emerging importance in metazoans. Of the thousands of O-GlcNAcylated proteins identified, many carry multiple modification sites with varied stoichiometry. To better match the scale of O-GlcNAc sites and their dynamic nature, we herein report an optimized strategy, termed isotopic photocleavable tagging for O-GlcNAc profiling (isoPTOP), which enables quantitative and site-specific profiling of O-GlcNAcylation with excellent specificity and sensitivity. In HeLa cells, ∼1500 O-GlcNAcylation sites were identified with the optimized procedures, which led to quantification of ∼1000 O-GlcNAcylation sites with isoPTOP. Furthermore, we apply isoPTOP to probe the O-GlcNAcylation dynamics in a pair of colorectal cancer (CRC) cell lines, SW480 and SW620 cells, which represent primary carcinoma and metastatic cells, representatively. The stoichiometric differences of 625 O-GlcNAcylation sites are quantified. Of these quantified sites, many occur on important regulators involved in tumor progression and metastasis. Our results provide a valuable database for understanding the functional role of O-GlcNAc in CRC. IsoPTOP should be applicable for investigating O-GlcNAcylation dynamics in various pathophysiological processes.
O-GlcNAc proteins:
A0A0B4J203, A0A0C4DFX4, RBM47, E2F8, WDR27, SBNO1, CNOT1, P121B, P121C, H0YAE9, H0YHG0, H7C469, K7ELQ4, M0QZ24, PDLI1, HAX1, TAF4, BCL9, CAC1A, DDX3X, NFIB, PPP6, MA2B1, ARI1A, SOCS7, ABLM1, KMT2D, GBRD, RGRF2, TX1B3, HGS, MYPT1, SYN3, ZN609, TRI66, PDZD2, MAST4, SC16A, SET1A, CASC3, FOXP2, MOT4, P4HA2, ARPC5, CLOCK, MAFG, PER1, KDM6A, TET3, SI1L1, TGFI1, M3K7, MCA3, PRPF3, TPD54, SYNJ1, IF4G3, E41L2, WIPF1, FOXO3, TGM5, RNF13, SPY2, PLRG1, ZN207, AKAP8, CALU, ORC5, MYPT2, GANP, OGA, CCNT1, BUB1B, PLOD3, PLIN3, MOT2, MAFK, PQBP1, BRD4, TBL1X, PP1RB, NBN, MITF, SRGP2, N4BP1, ROCK2, PP6R2, CNOT3, ANR17, FLNB, NCOR1, SF3B1, REM1, CREG1, CRTAP, SYUG, CYTF, TOX4, TOX, SUN1, PCF11, AGFG2, UBE4B, CAC1H, SVIL, SC24A, SC24B, CNOT4, EYA4, ZMYM6, BAG3, LATS1, DDAH2, TXD12, ONEC2, CLPT1, ABL1, CRYAB, LMNA, TFR1, CATA, GBA1, FUCO, ALDOA, GCR, G3P, CPNS1, HSPB1, RLA2, RLA0, ITB1, K1C18, NPM, CATL1, CATB, MCR, BGLR, ITA5, NFIC, VIME, SNRPA, FGR, ATX1L, DERPC, ZN865, GLI2, MYBB, CLUS, PPAL, MPRI, PABP1, TPR, BMP3, SKIL, ENPL, PO2F1, PLAK, ATF2, ZEP1, RS2, TFE2, F261, ITB4, ZNF23, ZNF25, JUNB, ATF7, TPH1, DDX5, EGR1, SON, NELFE, ATF1, ATF6A, CADH2, ICAL, CSRP1, FLNA, RFX1, CBL, SFPQ, COF1, IF4B, GATA2, APC, DDX6, ARNT, MAP4, LYOX, HXD9, MZF1, CLIP1, 5HT1F, HXA11, ZEP2, ELF1, CTNB1, FBN1, ADDA, BASI, NU214, VGFR2, SRP14, NUP62, SYUA, VATA, CUX1, TXLNA, STAT3, LAP2A, EPS15, HELZ, MATR3, SSRA, SSRB, KI67, ATRX, MAP1B, YAP1, UTRN, STT3A, SC6A8, RFX5, SOX2, PRC2A, HSP13, NR2C2, NASP, CDK8, DHE4, YLPM1, NU153, RBP2, TAF6, MRE11, EMD, MXI1, MAP2, TOB1, PPT1, TCPQ, PAPOA, GDS1, AGFG1, CRIP2, NUP98, SMTN, SC24C, HIRA, ATX1, ATN1, AFAD, AF10, AF17, DSRAD, SEC13, NU107, ZN445, CSN2, RL37, WDR5, TIM10, F193A, RBM6, PITX1, IF4G2, PHC1, ADA17, KGD4, RL19, FOXK1, DAB2, RHG04, RBM10, HNRPU, SPTB2, FOXK2, RUNX1, MEF2A, SP2, SP3, PLOD1, KMT2A, TF65, IF4G1, NOTC2, TLE4, PTN12, CALD1, MEF2C, P5F1B, GABPA, ZO1, ACK1, EP300, AHNK, FCHO2, HMGX3, SRBP2, FOXO1, ASPH, TROAP, BPTF, FSTL1, NFIA, DPYD, TP53B, FOXC1, ECH1, ROA0, DDX10, TBX2, GPS2, G3BP1, PABP4, ADAM9, PICAL, NAB1, SERC3, RIPK1, IQGA2, STIM1, CUL4B, ASPP2, CAC1S, RUNX2, NFYC, CDK13, TOB2, VEZF1, UBP2L, GIT2, SRC8, CAPR1, LAGE3, PUM1, MDC1, EPN4, TTLL4, RRP1B, NCOA6, GSE1, MEF2D, LASP1, MYPC3, ZN638, NUMA1, SART3, CND1, R3HD1, KIF14, WDR43, PLCL1, PLEC, NOMO1, NONO, RCN1, RYR3, KS6A1, RBMS2, TAF1C, SF01, MED1, JHD2C, TRIP6, T22D1, ELF2, TAB1, HERC1, NCOA1, VAS1, ZFHX3, ZYX, ADRM1, SYPL1, TAF9, DREB, DGKD, CGT, GEN, LY6K, RFX7, QSER1, AAK1, PRSR3, QRIC1, MA7D1, WDR72, TBRG1, TB10B, TPRN, FIL1L, SVEP1, AMOT, EPC2, CRTC2, PAN3, HS904, YIF1B, AG10A, IGS11, ZN628, BCORL, FIGN, BRD10, SH319, TGO1, PRC2B, TOIP1, CEP78, P4R3B, HP1B3, CE170, ZN362, FKB15, AKND1, ZEP3, LRIF1, SWT1, RHG21, UBAP2, RBM26, DEP1A, OGRL1, AHDC1, F222A, RPRD2, RN220, ZN318, TASO2, ZMYM4, PAPD7, TENS2, KANK2, ARID2, USF3, RHG17, CYTSA, ANR40, BICRL, JADE1, PKHA7, NIPBL, LIN54, TET2, RINT1, CRCDL, ZNT6, TTC41, RHGBA, NFRKB, RSBNL, KCD18, NCEH1, MDEAS, ZC3HE, LARP1, NHS, CRTC3, SAS6, MCAF1, BCOR, MPRIP, DNMBP, GGYF2, THADA, BNC2, NFXL1, NBEL2, CO039, SRCAP, CBAR2, UBN2, XIRP1, RAPH1, LARP4, HAKAI, ASXL2, SPT6H, KDM3B, ZCCHV, KANL1, RGPD4, POGZ, ZFY16, NUFP2, MAVS, CLAP1, EMSY, I2BP2, SRGP1, RBBP6, SH3R1, HUWE1, YTHD3, NPM2, ILDR1, KAISO, MYPN, LDB1, LYRIC, BCL9L, LUZP1, NRAP, RTTN, PRSR1, DDX42, CEP57, CD20B, CACL1, P66A, HIPK1, KCC1D, RN135, MY18B, AHNK2, FOXP4, NAV3, NAV2, MISP, ARI3B, IPRI, TEX2, MGAP, CC28A, Z3H7A, ANKH1, SUGP1, RPAP2, MILK2, SRRM1, ZZZ3, GAR4, PHAR4, RTKN2, DCP1B, XRN1, PELP1, CKLF8, TENS4, SPART, RPTOR, NUP93, ZN687, DOCK4, RHG24, RUSC2, SYNPO, FNBP4, D2HDH, RP25L, ATPF2, CPSF7, ARFG1, ENAH, SPOT1, SUMF1, KCNH5, SLAI1, TNR6A, PHC3, DRC6, CBPC3, NAV1, VP37A, KMT2C, ZMIZ2, BD1L1, ARI1B, FLCN, NUP35, TOIP2, TNIP2, KNL1, OR2L2, PUM2, CC110, TBC15, STT3B, ZN507, ALMS1, DLG5, KCNV2, BRX1, DOT1L, GEMI5, PARD3, ZN384, SMAP2, IASPP, TM263, ZFN2B, NUDC2, PCNP, TRUB1, LMO7, ATX2L, PALLD, P66B, BBX, ZCH14, GBF1, SMG7, RTF1, NICA, PHF3, MAML1, ZN592, LAR4B, TFG, TAF4B, RREB1, SC65, CBP, SYMPK, DDX17, GPKOW, FUBP2, UBP7, LPP, LSM10, NCLN, MRTFA, FUBP1, TTC17, PBIP1, TTC28, TOM6, PF21A, INT12, REPS1, ESS2, MBD6, ELP4, SGF29, RBM33, ZN503, P121A, TONSL, PDLI5, ERO1A, DOCK6, FUBP3, RSRC1, ZN594, VCIP1, ZN462, LCOR, PDLI2, CLP1L, Z512B, ZFR, EP400, MRFL, H6ST2, TIGD1, NOL4L, DOCK7, RPR1A, RBM14, ADCYA, QKI, LENG8, TRNT1, PP1RA, PHF12, CIC, MED15, ERBIN, HMCN1, LMF1, PIGS, WRIP1, SIN3A, MINT, HTF4, EYA3, POP1, TEAD3, TTC1, CSN8, ATX2, ARI3A, ANM1, PKP2, TEP1, DPH2, WAC, DIDO1, HNRL1, RBM4, SSBP4, PRR14, SSBP3, YTHD1, KPCD2, ZCHC2, TB182, AMRA1, CE295, TANC1, ZC12C, CEP44, STRAB, SP130, BRD8, RGAP1, SMG9, APC1, I2BPL, TMX4, KI13A, WDR13, EPC1, ADNP, ZN106, TM245, FOXP1, PABP3, WNK1, E41L1, ZHX3, BICC1, PEAK1, PPR3E, ZN703, PKHA5, CLSPN, BCDO1, RC3H2, ZFYV1, TAF9B, EMAL4, ZBT20, NCOA5, TANC2, ZN532, NCK5L, TNR6C, CHD8, FBSL, APMAP, DMAP1, UBN1, DCP1A, INCE, ANLN, GEPH, PDLI7, TULP4, HOME2, SLX9, DIAP3, BMP2K, RBM12, STAU2, DDX28, CWC25, CARF, ETAA1, ABI2, HXC10, BCLF1, TAB2, CELR3, CDK12, GRHL1, SACS, ITSN2, BICRA, CNOT2, TMEM9, CAC1I, CAMP3, DAPLE, RCC2, DIP2B, MBD5, CT2NL, F135A, KANL3, RERE, SE1L1, TRM7, YM012, KDM5B, LIMD1, TCF20, SUN2, LIMA1, SEPT9, UBQL2, TRPS1, S30BP, NRBP, BAZ2B, SIX4, HOOK1, CDC23, TASOR, GMEB2, TNIK, PARP4, NUP50, ZHX1, CDV3, MCTS1, KCNH3, LRFN2, MRTFB, ZBT21, PRR12, YETS2, HECD1, ZMYD8, NOTC3, SPAT2, SOX13, G3BP2, MAGD2, MINP1, MACF1, CP131, SCAF8, TRI33, PHF8, LIMC1, TNR6B, SRRM2, SCML2, ZN148, POLH, INVS, ICE1, R3HD2, MAN1, TR150, WBP11, ZN281, STA13, WNK2, ARIP4, MTCL1, DCAF1, RPGF2, IRS2, CRBG1, HYOU1, SAM50, PRC2C, YTHD2, NCOR2, GMEB1, DC1L1, EPN1, NCOA3, ZHX2, S23IP, U3KPZ7, V9GYH0
Species: Homo sapiens
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Chen Y, Tang F, Qin H, Yue X, Nie Y, Huang W, Ye M. Endo-M Mediated Chemoenzymatic Approach Enables Reversible Glycopeptide Labeling for O-GlcNAcylation Analysis. Angewandte Chemie (International ed. in English) 2022 61(23) 35289036
Abstract:
To selectively enrich O-linked β-N-acetylglucosamine (O-GlcNAc) peptides in their original form from complex samples, we report the first reversible chemoenzymatic labeling approach for proteomic analysis. In this strategy, the O-GlcNAc moieties are ligated with long N-glycans using an Endo-M mutant, which enables the enrichment of the labeled glycopeptides by hydrophilic interaction liquid chromatography (HILIC). The attached glycans on the enriched glycopeptides are removed by wild-type Endo-M/S to restore the O-GlcNAc moiety. Compared with classic chemoenzymatic labeling, this approach enables the tag-free identification, and eliminates the interference of bulky tags in glycopeptide detection. This approach presents a unique avenue for the proteome-wide analysis of protein O-GlcNAcylation to promote its mechanism research.
O-GlcNAc proteins:
TM271, HEAT9, SBNO1, P121B, GGT3, TAF4, AGRIN, MEIS1, RNT2, NFIB, MA2B1, CDKA1, ABLM1, ADA10, KMT2D, MYPT1, ZN609, SC16A, PLXB2, SET1A, VA0E1, TNC18, TLR3, TET3, PRPF3, IF4G3, ZN207, AKAP8, CALU, EXTL3, PCDH7, TRAK2, GANP, TSN3, MAFK, KERA, SRGP2, ANR17, NCOR1, U520, CRTAP, ATRN, CBPD, RMP, TOX4, SC24D, SRBS2, SUN1, ERLN2, PCF11, SCAF4, VAPB, SMC2, SC24B, CNOT4, EGFR, HPT, GLHA, LMNA, NU3M, VTNC, GCR, A1BG, K2C1, HSPB1, RPN1, AT1B1, K1C18, K2C8, ITAV, GDN, TRY1, SAP, MET, NFIC, VIME, K2C7, SNRPA, UCRIL, ATX1L, NT2NC, LYAG, LAMC1, PPAL, LAMP1, MPRI, TPR, SKIL, T2FB, GLU2B, NID1, PO2F1, MCP, ZEP1, CD36, ATF7, EGR1, SON, ATF1, ICAL, CO5A1, ASPG, SFPQ, ITA3, NKG2A, ARNT, TEAD1, 3MG, TKT, UFO, HXA11, HXC9, ZEP2, HNRH1, ELF1, CD68, RFC1, FBN1, NU214, RL4, SCNNA, SRP14, NUP62, TAGL2, CUX1, IL6RB, PBX2, STAT3, NBL1, LAP2A, LAP2B, LIFR, PCP, ECE1, MUC18, MATR3, SSRB, VDAC2, ATRX, NOTC1, UTRN, RFX5, HSP13, CCN3, AGRE5, NR2C2, CDK8, CENPF, YLPM1, RBM25, NU153, RBP2, TAF6, EMD, PPT1, SMCA4, AGFG1, NUP98, PTTG, ATX1, AT1B3, AF17, DSRAD, LAMB2, CAD13, ITA1, NU107, TGFB2, ACTG, CXAR, PITX1, CNTP1, ABCA4, PHC1, ADA17, KGD4, SARNP, FOXK1, DAB2, HNRPU, SPTB2, FOXK2, MEF2A, SP3, PLOD1, KMT2A, UPAR, IF4G1, NOTC2, SRS11, SUH, MFGM, AHNK, TMM62, GALT1, BST2, SIA4C, ASPH, BPTF, NFIA, FOXC1, ADAM9, NFAC2, MAMD1, NAB1, TBB3, PKD2, LAMB3, KLF5, NFYC, CDK13, SCRB2, LRP8, VEZF1, UBP2L, LAGE3, MDC1, RRP1B, MEF2D, ARI5B, NUMA1, PON2, RCN1, TAF1C, SF01, MED1, JHD2C, ELF2, TAB1, ZFHX3, ZYX, ADRM1, LAMA4, TAF9, FOXD1, LAMA3, RFX7, QSER1, SVEP1, SYTL3, EPC2, LUZP6, CRTC2, YIF1B, BCORL, BRD10, PRC2B, ZN362, LRIF1, UBAP2, RBM26, VP13D, RPRD2, RN220, ZN318, TASO2, ECM29, ARID2, SE1L3, BICRL, SCAR3, NIPBL, LIN54, FSTL4, TET2, ZNT6, GOLM2, NFRKB, S39A4, ZC3HE, FIP1, CRTC3, PLGT2, SUSD1, MCAF1, BCOR, IGS10, B3GLT, CD109, FRRS1, SRCAP, NBEL1, UBN2, RAPH1, HAKAI, ASXL2, SPT6H, KDM3B, NRK, NUP54, POGZ, MAVS, PK1L3, PLGT3, EMSY, RAI1, YTHD3, LDB1, LYRIC, OSTM1, PRSR1, DDX42, P66A, FOXP4, CTL2, TEX2, MGAP, ANKH1, SUGP1, NUP93, PLD6, FNBP4, ARFG1, PMGT2, GOLM1, PGLT1, TM87A, FSAF1, PHC3, SP20H, ARI1B, NUP35, OR6C4, VA0E2, PLBL2, PUM2, SPP2B, DLG5, GPX8, PO210, ZN384, LMO7, MUC16, P66B, BBX, SMG7, NICA, TM131, PHF3, TAF4B, GGH, GSLG1, FUBP2, LPP, NCLN, TTC17, FKB10, TOM6, PF21A, RBM33, I17RA, P121A, PDLI5, CREL1, FUBP3, TXD15, LOXL4, VCIP1, CHAP1, Z512B, ZFR, EP400, HUTU, RBM14, KI20B, PHF12, PANX1, CIC, MED15, ERBIN, JMJD8, MINT, TEAD3, SEM3C, WAC, DIDO1, HNRL1, TM109, NUP58, M4A8, YTHD1, ULBP3, AMRA1, TANC1, TWSG1, S22A4, SP130, APC1, I2BPL, WNT5B, VP33B, EPC1, ADNP, ZN106, FOXP1, PTN23, WNK1, SIA7D, ZHX3, PIEZ2, SG196, TM231, PEAK1, DOCK5, CF155, CNO10, MLXIP, SIAE, PKHA5, RC3H2, TAF9B, ZBT20, NCOA5, TANC2, ZN532, CELR2, APMAP, ABCB9, CHSTB, UBN1, NECT3, PDLI7, RBM12, OCAD1, CARF, ETAA1, HXC10, TAB2, CELR1, CDK12, CNOT2, CRIM1, TMEM9, RCC2, CHD7, RBM27, KANL3, MRC2, SUCO, TCF20, SUN2, UBQL2, LCAP, GGT7, TASOR, GMEB2, ZHX1, DSE, WWC3, MRTFB, ZBT21, PRR12, YETS2, NOTC3, KMT2B, PRP19, MINP1, SCAF8, ZC3H4, SRRM2, SCML2, UST, ICE1, ZN281, DAAM1, IRS2, PRC2C, NCOR2, NPTN, GMEB1, POMT1, S4A7, S23IP
Species: Homo sapiens
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Xie X, Wu Q, Zhang K, Liu Y, Zhang N, Chen Q, Wang L, Li W, Zhang J, Liu Y. O-GlcNAc modification regulates MTA1 transcriptional activity during breast cancer cell genotoxic adaptation. Biochimica et biophysica acta. General subjects 2021 1865(8) 34019948
Abstract:
Chromatin modifier metastasis-associated protein 1 (MTA1), closely associated with tumor angiogenesis in breast cancer, plays an important role in gene expression and cancer cell behavior. Recently, an association between O-GlcNAc transferase (OGT) and MTA1 was identified by mass spectroscopy. However, the potential relationship between MTA1 and O-GlcNAc modification has not yet explored.
O-GlcNAc proteins:
MED19, SBNO1, PGP, DPRX, SMHD1, RGPD3, ZSWM8, P121C, VGLL3, MCRI1, CK098, KIF2A, DX39A, PDLI1, BACH, MYO1C, MYO1F, AIP, SMAP, AP3B1, PSD11, PSD12, PSMD9, SPT5H, TAF4, DFFA, CLIC1, EIF3F, MATN2, LORF2, IPO5, SAP18, IF2B3, DNM1L, RTCA, AGRIN, PLOD2, HMGN4, IMA4, BCL9, PESC, NOP56, DDX3X, PODXL, CCN1, IMA3, NFIB, MA2B1, PDXK, ARI1A, TXND9, G3PT, CTRO, COPE, AP3D1, PDCD5, NHRF1, TPP1, TCRG1, TNPO2, PSA7, GIPC1, HAT1, HGS, MYPT1, HNRDL, XPO1, ZN609, SPTN2, SC16A, PLXB2, SR140, SET1A, LSM1, NPC1, ARC1B, ARPC2, ARPC3, TIF1A, PGRC2, PFD6, NKRF, LAMA5, ZN185, CASC3, OGT1, PMM2, HMGB3, PPM1G, FANCA, EIF3D, EIF3H, HDAC3, IPO8, STX7, NUP42, CLOCK, KDM6A, DHX15, MCES, FLRT2, PRP4, SERA, DC1L2, PSMD3, RFOX2, PAPS1, ZW10, SNUT1, SPIT2, TGFI1, M3K7, MCA3, HNRPR, PRPF3, TPD54, COCH, IF4G3, PPIH, ARK72, E41L2, TGON2, S26A4, FOXO3, DENR, XPOT, PLRG1, RGS10, ZN207, GET3, BUB3, ACTN4, BUD23, HTSF1, AP1G1, SYNC, KRT86, CPSF5, LANC1, AKAP8, CALU, RAD21, DHX16, MED14, SMCA5, JIP4, ZC3H1, TRAK2, AQR, GANP, KDM1A, GSDME, EVI5, SNX3, OGA, HNRPQ, CCNT1, PLOD3, AP180, PLIN3, MAFK, IMA7, UGDH, CTND1, SNX2, USO1, CCD22, PQBP1, DKC1, IF2P, EDF1, DNJA2, BRD4, PFD1, NBN, MCE1, WDR1, CPNE3, BRE1B, ZC11A, CLU, T22D2, PP6R2, CNOT3, CREST, ANR17, ZPR1, DUS11, PDCD6, TBCA, VATG1, H2AY, FLNB, NCOR1, SPAG7, SC22B, DEAF1, PR40A, VP26A, SAP30, MED24, KTNA1, PSIP1, SRS10, BAF, SF3B1, CSDE1, PRKRA, MED6, NPM3, LYPA1, U520, NU155, WDHD1, CRTAP, EIF3G, STAM2, CCNK, PRAF3, PIAS1, PIAS2, SPF27, DNJC8, SPF30, MPDZ, GLRX3, KRT36, RL1D1, CIAO1, SRP72, DDAH1, RECQ4, MTA2, MYO1D, TOX4, SC24D, PHF14, UBXN7, SUN1, PLPHP, ERLN2, PRP6, PCF11, NFAT5, HAUS5, AP2A2, SC31A, HEXI1, SEM3D, UBR5, AGFG2, SCAF4, ZFPL1, EPN2, ZRAB2, LC7L3, KAT7, MPZL1, FKBP9, CELF2, PAPS2, SMC2, LYPA2, IPO7, AGM1, CD2B2, BAG4, AHSA1, PSMG1, SC24A, SC24B, AGRL2, ACSL3, PCNT, CNOT4, CPSF4, EYA4, K2C75, SNP29, OXSR1, HS74L, PTBP3, AP2A1, STAU1, CAVN2, BAG2, M4K4, BPNT1, MBD3, CLPT1, ACL6A, CYB5, ADH1B, ADH1G, LDHA, DHE3, NB5R3, GSHR, SODC, PNPH, HPRT, AATM, EGFR, PGK1, KAD1, LALBA, FA10, TRYP, ASSY, A2MG, LDLR, HLAH, HBA, CO1A1, K1C14, K2C6A, LMNA, CASA1, CASA2, CASB, CASK, FINC, LACB, ALBU, ALBU, TFR1, GBA1, ALDOA, CYTB, ANXA1, GCR, PRIO, KITH, K2C6B, K2C1, G3P, CPNS1, HSPB1, RPN1, RPN2, GNAI2, AT1B1, ALDOB, A4, ARGI1, S10A8, HMGN1, ISG15, IF2A, HMGN2, ICAM1, RLA1, RLA2, JUN, LA, ITB1, K1C18, K2C8, MYL1, GELS, PTMA, CDK1, ATPB, S10A6, IF4E, ENOA, PYGL, G6PI, DPOLB, NPM, ITAV, LIPL, HEXA, ACBP, FABPL, LDHB, PDIA1, H10, CATD, ANXA2, CAN1, TBB5, TRY1, SAP, HEXB, PROF1, APT, K2C4, SYEP, CATB, HS90A, HNRPC, LAMB1, TPM2, FUMH, TSP1, SP1, ANXA6, DAF, MDR1, 4F2, HS90B, ASNS, ADH4, HEM3, RU2B, MET, RU17, ITA5, NFIC, VIME, RS17, K2C7, K1C13, ANXA5, K1C16, RSSA, CD63, SNRPA, GSTP1, RU1C, LEG1, HMGB1, CLCA, ROA1, RU2A, PARP1, UCHL1, LKHA4, ALDOC, CC175, ATX1L, ZN865, CT45A, HS71B, CALM3, RO60, H14, PTPRF, THIO, PPGB, KAP0, HEM4, ESTD, CH60, BIP, LAMC1, PPAL, HSP7C, GTR1, PYGB, LAMP1, TOP1, TOP2A, G6PD, C1TC, MPRI, ADHX, SRF, PABP1, PCNA, HARS1, IMDH2, TPR, KCRB, ANXA3, ACTN1, XRCC6, XRCC5, LAMP2, RINI, MYH6, ICAM2, AT1A3, EF2, K1C10, K2C5, PDIA4, TCTP, PLST, GYS1, T2FB, CD59, MIF, GLU2B, FPPS, TRI27, AK1A1, KPYM, ENPL, CCNB1, PO2F1, HNRPL, SYDC, ETS1, PLAK, ALDR, AMPN, PVR, ERF3A, EZRI, ATF2, FOSL1, FOSL2, MCP, NQO1, GNS, ZEP1, RS2, TFE2, DESP, RFA2, CD44, ITB4, CBR1, CREB1, BGAL, H15, H13, H12, AT2A2, STMN1, YBOX3, HSP76, HMGA1, JUNB, UBF1, JUND, ATF7, CAN2, PTN2, PYRG1, DDX5, PFKAL, LEG3, PRS6A, TCPA, PTN1, ARF4, RL7, VINC, SON, NELFE, RL17, PGAM1, RCC1, ATF1, XRCC1, CADH2, ML12A, NUCL, HXK1, RO52, SPEE, SL9A1, CSK22, RXRA, NFKB1, IF2B, ANXA7, TBP, BTF3, PSB1, MPRD, LMNB1, ICAL, CD11B, VATC1, CSRP1, FLNA, 5NTD, TAF1, CD9, TGM2, PIMT, FBRL, PUR2, PUR6, UBA1, NDKB, ROA2, RFX1, CBL, EST1, FBLN1, TCEA1, ITA6, SFPQ, PPIB, SYWC, RS3, NFYA, SAHH, COF1, IF4B, KTHY, CCND1, COT2, EF1B, PPAC, RPB1, CDK2, MCM3, NFYB, THTM, LAMA1, RS12, BRD2, TYY1, DNJB1, ATPA, PSA1, PSA3, PSA4, ITA3, MOES, DDX6, CTNA2, DNMT1, PAX6, U2AF2, RL13, HMGB2, PTBP1, SYTC, SYVC, EF1G, 1433T, ARNT, RL10, RFA1, APEX1, PYR1, CALR, MAP4, CALX, PSA5, DPOD1, MK01, RXRB, GRN, PTPRM, T2EA, CCN2, EPHA2, 3MG, KDM5A, TKT, RBMS1, PML, EF1D, ERP29, PRDX6, BLVRB, PRDX5, RL12, KCY, PEBP1, PDIA3, 2AAA, 2AAB, NMT1, HMOX2, PURA2, UFO, PUR8, CLIP1, SORCN, ILEU, RPB2, SDHA, METK2, CPSM, TIA1, ZEP2, DNJA1, AKT1, PUR9, HNRH3, HNRH1, 1433B, 1433S, STIP1, S10AB, L1CAM, PRDX2, OTX1, CDD, ELF1, RL9, KINH, CSTF2, DUT, TTK, MCM4, MCM5, MCM7, HSP74, GPC1, PROF2, CTNA1, CTNB1, PCGF2, SPB6, RADI, RFA3, RFC4, RFC2, RFC1, RL22, T2FA, SPRE, HXD13, CBS, K1C9, FBN2, GDE, MYH9, MYH10, COPB2, BASI, FUS, NU214, DEK, K22E, RL4, PP1G, GNL1, SRBP1, SRP14, NUP62, TAGL2, TALDO, RBMX, COIL, VKGC, HSPA9, IF4A3, RS19, RL3, COIA1, OST48, AN32A, FEN1, CAPG, IL6RB, TXLNA, TCPZ, NNMT, PBX2, RL13A, STAT3, MDHC, MDHM, RFC5, IF2G, CETN2, BUD31, NAA10, CSK, GARS, SYIC, EIF1, KPCI, LAP2A, LAP2B, MTREX, RS27, HELZ, RL35, CDN2A, PCP, ECE1, LIS1, MUC18, MATR3, MSH2, SSRA, RANG, NAMPT, PRS6B, NKX21, VDAC2, CBX5, UBP5, KI67, RAGP1, RECQ1, NOP2, CRK, CRKL, BAG6, NSF, NOTC1, MP2K3, RL27A, RL5, RL21, RL28, RS9, RS5, RS10, MAP1B, GNPI1, YAP1, UTRN, IQGA1, STT3A, CAPZB, IF1AX, SYQ, RL29, LIMS1, PPCE, RFX2, RFX5, SOX9, COPD, GSH0, PSMD8, PRC2A, TCPE, NEST, HSP13, KC1A, PIPNB, AGRE5, DPOD2, PAXI, RL34, NASP, FAS, CDK8, TCPG, EFTU, SRP09, PCY1A, SYAC, SYCC, SYSC, PRI2, CTCF, PSB2, MCM2, TSP3, YLPM1, TMEDA, RBM25, NUMB, HINT1, NU153, RBP2, GSK3A, GSK3B, TAF6, SPS1, GUAA, DNLI3, MRE11, CRIP1, GDIB, EMD, SERPH, F10A1, VASP, DYN2, CDK9, BCAM, PPT1, RL14, TCPQ, TCPD, ANX11, PAPOA, FXR1, FXR2, RAB5C, RAB7A, DUS3, SMCA4, GALK1, SSRD, UBP11, NR0B1, HDGF, ROA3, 6PGD, HNRPM, IMA1, IMA5, GDIR1, AGFG1, HNRPF, TF2AA, MSH6, ZN131, ZN143, RBM5, SPSY, EFNB2, CAZA1, CRIP2, NUP98, PPP5, ACLY, MAP11, COPB, COPA, CATC, AP2S1, PTTG, SMTN, IST1, SC24C, TCP4, ICLN, SYRC, ATX1, ATN1, SYYC, UBP14, HSP72, BCAT1, AT1B3, RD23B, PRRX1, HMCS2, SNAA, IF5, DRG2, XPO2, TERA, MFAP1, AF10, AF17, NP1L1, ADK, DSRAD, LAMB2, FOXG1, TPD52, SEC13, NH2L1, PSA, HNRH2, SUMO3, EIF3B, GFRA1, SYMC, IF6, CTBP2, TMM33, PCBP3, PCBP4, CORO7, NU107, MTPN, H2B1D, DEF3, ARPC4, SC61G, TPIS, EIF3E, SC61B, PP4C, MYL6, ACTB, IF4A1, RS20, PRPS1, PSA6, S10AA, CDC42, DEST, GMFB, RAB8A, SPCS3, RAB2A, RAB5B, RAB10, UBC12, UBE2N, ARP3, ARP2, ACTZ, CSN2, ABCE1, RS3A, RL26, PSME3, RL15, RL27, RL37A, RHOA, S61A1, LYSC, COPZ1, SUMO2, DCAF7, WDR5, NTF2, HNRPK, 1433G, RS7, PP1A, PP1B, PRS4, PRS8, RS8, RS15A, RS16, UB2G1, 1433E, RS14, RS23, RS18, RS29, RS13, RS11, RUXE, RUXF, RUXG, LSM3, LSM6, SMD1, SMD2, SMD3, TYB4, PRS10, RL7A, RB11A, ERF1, CNBP, RS4X, RL23A, RS6, H4, H2B1C, RAB1A, RAN, RL23, RAP1A, UB2D2, RS24, RS25, RS26, RS28, RS30, GBB1, RBX1, GBB2, RL30, RL31, RL10A, RL32, RL11, RL8, PPIA, FKB1A, RS27A, GRB2, TRA2B, RAC1, AP2B1, 1433Z, RSMN, SUMO1, DYL1, RL38, GBG5, RS21, IF5A1, RACK1, ACTG, SPT4H, UBC9, TYB10, PP2AA, YBOX1, CSK2B, TPM4, UB2L3, EF1A1, ACTS, TBA4A, TBB4B, CSK21, PA1B2, HBB, HBA, F193A, IGBP1, RBM6, IF4G2, RPP30, GTF2I, CNTP1, PHC1, TCPB, KRT85, RAE1L, PRKDC, MT1X, NUCB2, DCD, SSBP2, SARNP, RL24, CBX1, ARF1, ERH, RL19, SRSF3, FOXK1, DAB2, FBLN2, PGBM, RHG04, EFNB1, RBM10, RBM3, 2ABB, PIPNA, S25A3, VIGLN, TF2B, CDK5, CLH1, HSF1, FKBP3, DHSO, HNRPU, U2AF1, SPTB2, TIAR, SET, SRSF2, FOXK2, RUNX1, FABP5, CAP1, NC2B, TFAP4, EXOSX, OTUD4, PFKAP, EWS, TAGL, AK17A, MEF2A, KIF23, NRG1, DSG1, SP4, SP3, RL18A, MP2K1, FKBP4, PLOD1, NUCB1, RL6, TOP2B, DYST, CREM, KMT2A, LMNB2, TF65, UBXN1, GLGB, IF4G1, K1C17, NOTC2, TLE2, TLE4, LGUL, AK1C1, 1433F, CSTF1, PTN12, FAK1, SRS11, EF1A2, CALD1, PTN11, GFPT1, EXOS9, PSME1, SUH, GABPA, GABP1, RING1, PAX8, FMR1, PRDX1, RL18, CKAP4, ZO1, KHDR1, BAX, KLC1, LRP1, SRSF1, SRSF4, DHX9, GOGA2, LG3BP, CD47, PPID, SSRP1, NSUN2, RBBP4, EP300, AHNK, APOBR, HSP7E, MGAT2, GALT2, AP1B1, CPSF1, BST2, WASC5, NU160, SCAP, SRBP2, TBL3, ASPH, TROAP, TROP, SNF5, BPTF, CDC20, FSTL1, GRSF1, SFSWA, CHD3, SF3A3, TP53B, AIMP1, ILF2, ILF3, LMAN2, ELAV2, FOXC1, PP1R8, CSTF3, FLII, COASY, ACACA, CSN1, CAF1B, RED, MTAP, TADBP, ROA0, AIMP2, FADD, PRDX4, PAK2, CBX3, STK3, PSMD2, GPS2, SRSF9, SRSF5, SRSF6, TIF1B, G3BP1, CLN3, PTK7, PABP4, IFIT5, MTA1, EIF3I, 2A5G, CTBP1, UB2V1, DC1I2, ILK, MSLN, PPIG, TCOF, SF3B2, HAP28, ADAM9, FKBP5, SMAD4, PICAL, SQSTM, TBB3, ASAH1, PRP4K, PIN1, RIPK1, HDAC1, KCC2G, DCTN2, SNW1, STIM1, TRA2A, PWP1, CUL1, CUL2, CUL3, CUL4B, ASPP2, DYR1A, TPBG, FHL1, PM2PB, MOGS, CD166, NACA, THOC5, SPTN1, PKP1, DX39B, EXOS2, TBB2A, KLF5, BYST, IDI1, RUNX2, PEBB, NFYC, CDK13, CKAP5, CIRBP, COTL1, HNRPD, SCRB2, NID2, DAG1, VEZF1, DSG2, SAFB2, EIF3A, UBP2L, SCRIB, MLEC, TTL12, FHL2, DCTN1, DYHC1, EI2BA, IF4A2, SRC8, TRI25, FLNC, FA50A, MYPC2, GAMT, CAPR1, RBM39, KT33B, HLTF, PDIA5, DHX8, MCM6, ITPR3, MDC1, EPN4, SMC1A, RRP1B, NCOA6, GSE1, PSF1, UBP10, GANAB, 2A5D, LBR, GOGB1, MEF2D, CHD4, LASP1, NC2A, ZN638, IMB1, NOLC1, NUMA1, GAPD1, K2C72, NAA25, FL2D, SEPT2, SART3, U5S1, S39AE, SETB1, IF4H, BRD3, EEA1, PDIA6, PA1B3, PLEC, NOMO1, IPYR, TEBP, NECT1, NONO, PTPA, RBBP5, RCN1, K1H1, PCBP1, PCBP2, ELOC, SF3B3, DLGP5, RSU1, CNN3, SAFB1, SF3B4, SF3A2, RBMS2, SC23A, SC23B, SF3A1, SSXT, TAF5, TEAD4, TGIF1, NCOA2, NHRF2, TRAM1, SF01, TRIPB, PCH2, MED1, HMGN3, JHD2C, TRIP6, MARE1, ELAV1, ELF2, TAB1, AAAT, MLF2, TOM34, TBCE, UB2V2, STXB2, NEDD8, USF2, VAS1, VPS72, ZFHX3, ZYX, SEPT7, ADRM1, CCDC6, UAP1, AINX, PSMD5, CSRP2, DDB1, CDC37, DPYL2, RBBP7, TAF9, SRSF7, CPSF6, SMN, NRF1, FSCN1, IF16, GLO2, H2B2E, CGT, TRXR1, LY6K, HNRL2, Q1RMN8, PDS5A, QSER1, AAK1, WASC4, PRSR3, TSR1, QRIC1, SMU1, P3H1, MA7D1, CHD9, LSM12, GELS, A1AG, TBB8, GLYR1, FIL1L, AMOT, FA98B, EPC2, RHG29, FND3B, CRTC2, DHB12, GLE1, UBP39, LACB2, P5CR3, PKHG4, H90B2, R39L5, ZN326, FILA2, BCORL, FIGN, BRD10, AG10B, CC038, TGO1, PRC2B, RRP12, TOIP1, PCID2, SPXN3, SMAG2, BRM1L, TM201, NU188, HP1B3, CE170, ZN362, EYS, FKB15, DDX59, ZC3HD, LRIF1, UBR4, UBAP2, KPRP, KPLCE, RBM26, NT5D1, AHDC1, RPRD2, PR38B, BRE1A, ZN318, BROX, TASO2, ECM29, MBNL2, ZMYM4, K2C79, ARID2, USF3, C19L1, VIR, K1C40, HEAT6, BICRL, ATLA3, CARL2, CPIN1, TWF2, P4R3A, NIPBL, PDE12, LIN54, TET2, TM214, ARMC6, CAVN1, ERAP2, CDC73, TAF2, MED27, EDC4, PRP8, SCYL2, TTC27, GOLM2, NFRKB, MZT2A, CTR9, NCEH1, TRI65, MDEAS, BRAT1, ZC3HE, LARP1, BRWD3, SMRD3, FIP1, CRTC1, SAS6, CSPG4, CWC27, WDR82, ATL4, MCAF1, PACS1, MPRIP, GGYF2, CD109, CERS6, QSOX2, SRCAP, RIPR1, YJ005, UBN2, FGD6, PAXI1, RAPH1, IKIP, H32, LARP4, MED25, TBA1A, H2AV, HAKAI, ASXL2, CCD80, BT2A1, SPT6H, SND1, DDX46, 5MP2, RSRC2, CYFP1, KDM3B, OLIG3, PHF5A, ZCCHV, KANL1, NUP54, POGZ, MYH14, NUFP2, CLAP1, PLGT3, HEAT3, HDGR2, EMSY, RAI1, I2BP2, RBBP6, SH3R1, HUWE1, YTHD3, KHDC4, TMED4, CENPV, T179B, Q862S4, KAISO, PATL1, MYPN, PEG10, NOP9, PABP2, MTA70, LDB1, PB1, LYRIC, TLK2, ZN598, KTN1, BCL9L, DTX2, LUZP1, THOC4, GP180, PACS2, CAND1, TSYL5, THOC6, MET16, ZC3C1, CARM1, PRSR1, MEX3D, DDX42, DAAF5, P66A, HORN, RB6I2, RIOX2, RBM45, PLD3, NUDC3, AHNK2, FOXP4, TEX2, Z3H7A, PHF6, CHERP, ANKH1, SUGP1, CCAR1, SLF2, PHC2, BAP18, RB12B, RAVR1, SPB1, IHO1, SRRM1, SMAP1, P20D2, KATL2, PHAR4, TRM2A, DCP1B, NALCN, MAML2, PELP1, ABI1, PORIM, CCAR2, NUP93, ZN687, CMTR1, LRC47, K2C78, ABD12, PALS1, STAG2, FNBP4, TAB3, DPP10, CPSF7, ARFG1, EMC1, PAF1, UBR7, ENAH, KRI1, PR38A, F184A, KDM1B, NHLC2, GT251, PGLT1, DHB11, TXND5, SERB1, LEMD2, FA98A, RN214, LS14A, MISSL, TNR6A, PHC3, ABCF1, NGDN, SP20H, VP37A, KMT2C, ZMIZ2, ARI1B, NUP43, NUP37, NUP35, STXB6, KNL1, PLBL2, RLA0L, THOC2, WDR36, SMRC2, NPL4, PUM2, RDH11, STT3B, SPP2A, HM13, ALMS1, C99L2, MK15, BRX1, RBCC1, DOT1L, PO210, GEMI5, PARD3, IPO4, ZN384, K2C79, NOC3L, SCRB1, SMAP2, CCD12, TM263, NU133, PDC6I, GATD1, THEM6, CTF18, NUDC2, SCFD1, HNRLL, UBCP1, PCNP, LMO7, ATX2L, PHIP, PRP31, NELFB, PALLD, SREK1, PSPC1, P66B, DYH7, DNJC9, BBX, CTBL1, KAT6B, ELYS, ZCH14, AP2B, DDX1, H1X, PSMF1, SMG7, RTF1, NICA, TM131, ELMO1, BAP1, PHF3, MAML1, NDRG1, HS105, CNOT9, ZN592, LAR4B, GCN1, PRP16, NU205, AN32B, NECT2, PRCC, TFG, TAF4B, HDAC2, STAM1, REQU, CBP, KAT6A, SYMPK, RBP56, GGH, DDX17, RAD50, CELF1, OSTF1, UFD1, RENT1, CSN5, GPKOW, SMRC1, SMRD2, FUBP2, TNPO1, NEP1, GLMN, USP9X, UBP7, CUL5, STMN2, LPP, MED12, H2A1C, BHMT1, SMCE1, MYDGF, RL36L, PSMG2, NCLN, UTP4, PP14A, MGN2, FERM2, FUBP1, LRC59, FKB10, PF21A, PPWD1, INT12, FWCH2, KLH29, CC124, AP2M1, REPS1, SNR40, MBD6, RMXL1, SEH1, TCAL4, THA11, GNA1, T3HPD, DAZP1, SAAL1, SGF29, RBM33, TMA16, ZN503, DYL2, CPNE2, THOC1, OTUB1, HMCES, MED8, DUS3L, DCNL1, SMRD1, DDX27, GWL, LIN37, ZCCHL, P121A, PDLI5, ERO1A, C2AIL, MED30, INT4, FUBP3, SPF45, ABHEB, PAWR, THOC3, KIRR1, WNK4, COG3, TMX3, CHAP1, LCOR, PDLI2, ANCHR, ZC3HA, TOPK, SCYL1, EHMT2, ZFR, EP400, PRRC1, FXL18, CCD43, NOL4L, DACH2, NSUN5, RPR1A, IPO9, RBM14, QKI, ALS2, PP1RA, S38A2, VPS35, PURB, PHF12, XPO6, NACC1, CIC, MED15, MCCA, NUDC1, ERBIN, WRIP1, RANB9, MYADM, SRPK1, STRBP, TM209, IWS1, SIN3A, RSF1, RBM15, RUFY1, MINT, PNKP, MMS19, UHRF1, NIBA2, HTF4, TBCB, CNN2, CDC5L, PSMD1, PFD5, RING2, PARK7, EYA3, VAT1, LGMN, NUP88, POP1, MNT, SCAFB, EIF3C, TTC1, DNJC7, KIF2C, RGPD5, ATX2, ROAA, NP1L4, SNAG, TS101, CPNE1, TCPH, EBP2, ARI3A, ANM1, H2A1J, PKP2, SEM3C, VRK1, RBM4B, DDX50, RNZ2, MEP50, TBA1C, MBB1A, COR1B, TXD17, CPPED, PDD2L, PYGO2, LLPH, PSF3, ERP44, GCP2, ESYT1, SEN34, CSN4, WAC, DIDO1, RBM42, MCMBP, DCNL5, RAMAC, AN32E, NDC1, MMTA2, TBB6, HNRL1, PHF23, DDX23, MCRI2, WDR18, MTND, NTM1A, TBB2B, ASHWN, TM109, UT14A, TMED9, NUP58, SELO, DPCD, GNL3, KIFC1, NUP85, ELOV1, THIC, RBM4, GATA5, LS14B, RBM24, TRPM6, NAA15, SRRT, AP1M1, MAK16, ITPA, RTF2, EIF2A, PDIP3, TMPSD, MK67I, YTHD1, WNK3, GTPB4, UACA, CRNL1, TBL1R, PP12C, API5, DPY30, TRIM4, GRB1L, UNK, TB182, AMRA1, TANC1, TTYH3, WDR33, TINAL, NDEL1, PIP30, MKRN2, ILKAP, XRN2, TOLIP, SP130, RGAP1, CSTFT, RAB1B, LMA2L, ING2, I2BPL, TMX4, EPN3, MED28, SH3L3, PPIL3, ADNP, DHX36, PININ, PABP3, CPVL, BOLA2, TMX1, NELFA, PTN23, UN45A, CHM4B, OFUT1, WNK1, AMPB, ZHX3, EHD1, HEAT1, SUV92, CCD86, PRR36, BCAS3, RANB3, PEAK1, SMYD3, WDR26, SDS3, SFR19, KAT8, PHAX, UBE2Z, RMD5A, BRD9, GORS2, JUPI2, GHC1, MED20, CNO10, KANL2, FBRS, SP30L, MLXIP, XPO5, MYG1, CYBP, RC3H2, TENS1, SPC25, TAF9B, GLOD4, NCOA5, MOV10, TNR6C, CHD8, SYF1, SRA1, CHM1A, TM9S3, APMAP, ARFG3, HM20A, SSU72, VTA1, ENY2, CD320, DMAP1, UBN1, DCP1A, MED4, RIC8A, LUC7L, TIGAR, RTN4, RPR1B, EXOS3, PRD10, ANLN, PDLI7, DDX21, SAR1A, SIAS, SHLB2, MBNL1, AAAS, BAZ1A, ADPPT, OSTC, UBQL4, RAB6B, ABC3C, PHP14, HELLS, KRT84, KRT82, SYFB, PLPL3, BMP2K, ATG3, SMC4, OLA1, ZF64B, RBM12, MDN1, STAU2, T106B, SPS2L, CYRIB, DD19A, MRGBP, CC50A, SEP11, MED17, INT13, DDX18, NLE1, RBM22, WDR70, SLTM, LYAR, MED29, NSMA3, THUM1, TRM1, CARF, GAR1, FA53C, BCLF1, TAB2, FKB11, TMOD3, UGGG1, CDK12, GGA3, TLS1, IF2B1, HPBP1, MAT2B, MYOF, ITSN2, BICRA, EHD2, CNOT2, SPN90, LMCD1, CHMP5, COMD9, CWC15, THYN1, HACD3, KCMF1, RAI14, VAPA, H2AW, TMEM9, SEP10, CACO1, DAPLE, RCC2, MBD5, SLAI2, BCCIP, RRBP1, CPSF2, SYLC, MYEF2, RBM27, KANL3, ANFY1, RERE, ATX10, EP15R, SAE1, COPG2, MRC2, ASH2L, DKK3, TRMB, GULP1, VPS29, GRHPR, CATZ, SAE2, NXF1, PEF1, DNJB4, STK39, CNOT8, ABCF2, GTR6, Z3H7B, TCF20, HMGX4, SWP70, SUN2, RLP24, LIMA1, SRP68, CHRD1, SEPT9, UBQL2, CEIP2, S30BP, MED13, PFD2, PUF60, NRBP, TR112, XPO7, LCAP, MBD1, SIX4, CNOT7, PLXA1, MAGL2, TRM6, FOXD3, DBNL, DDX41, VPS28, CCNL1, TASOR, JUPI1, GMEB2, MRT4, CPSF3, GPTC8, NXT1, RCOR1, RALY, TF3C4, ACINU, AGO2, NUP50, ZHX1, CDV3, CNO11, ZN236, PSME2, MCTS1, ASAP1, MRTFB, ZMIZ1, MED23, PRR12, YETS2, HECD1, ZMYD8, COR1C, TPX2, AKP8L, MAFF, TMCO1, EPDR1, DD19B, PPT2, PRP19, SYNP2, UBQL1, VPS4A, G3BP2, CHIP, PACN2, MAGD2, SNX6, SSRG, FAF1, EPCR, CSN3, NSF1C, SCAF8, TRI33, TNR6B, ZC3H4, SMAG1, TRAK1, FOXJ3, SRRM2, PA2G4, SPG7, SMC3, SCML2, ZN148, RTRAF, RUVB2, CDYL, EIF3L, PLAP, RUVB1, NUDC, COF2, SYFA, DRG1, NCKP1, OFUT2, PKHA6, WDR37, E41L3, MYH15, R3HD2, KDM2A, RRP44, RCL1, DDX52, TMA7, YBOX2, MAN1, CHSP1, TR150, WBP11, MED16, NOP58, GIT1, ZN281, SGT1, AAR2, NOSIP, MEMO1, LSM2, SHLB1, LC7L2, SBDS, TMED7, SF3B6, ORN, RRP15, NOP16, PPIL1, MED31, KAD6, STRAP, RTCB, FBX7, NOC2L, RL36, CHTOP, SAMH1, PKP3, SALL2, TLN1, TRRAP, ARIP4, MTCL1, MYO5A, CRBG1, HYOU1, ATG4B, TBL2, KLF12, LSM4, RN114, PRC2C, PPME1, RBM7, YTHD2, PAXB1, SP16H, CD2AP, TNPO3, RBM8A, SNX9, NUBP2, HCFC2, PUS1, SERC, NCOR2, SPIN1, COPG1, GMEB1, CLIC4, CFA20, TACC3, MD1L1, 5MP1, EPN1, TEST, IF2B2, S4A7, ROBO1, NCOA3, SCC4, ZHX2, S23IP
Species: Mus musculus, Sus scrofa, Bos taurus, Homo sapiens
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Liu Y, Chen Q, Zhang N, Zhang K, Dou T, Cao Y, Liu Y, Li K, Hao X, Xie X, Li W, Ren Y, Zhang J. Proteomic profiling and genome-wide mapping of O-GlcNAc chromatin-associated proteins reveal an O-GlcNAc-regulated genotoxic stress response. Nature communications 2020 11(1) 33214551
Abstract:
O-GlcNAc modification plays critical roles in regulating the stress response program and cellular homeostasis. However, systematic and multi-omics studies on the O-GlcNAc regulated mechanism have been limited. Here, comprehensive data are obtained by a chemical reporter-based method to survey O-GlcNAc function in human breast cancer cells stimulated with the genotoxic agent adriamycin. We identify 875 genotoxic stress-induced O-GlcNAc chromatin-associated proteins (OCPs), including 88 O-GlcNAc chromatin-associated transcription factors and cofactors (OCTFs), subsequently map their genomic loci, and construct a comprehensive transcriptional reprogramming network. Notably, genotoxicity-induced O-GlcNAc enhances the genome-wide interactions of OCPs with chromatin. The dynamic binding switch of hundreds of OCPs from enhancers to promoters is identified as a crucial feature in the specific transcriptional activation of genes involved in the adaptation of cancer cells to genotoxic stress. The OCTF nuclear factor erythroid 2-related factor-1 (NRF1) is found to be a key response regulator in O-GlcNAc-modulated cellular homeostasis. These results provide a valuable clue suggesting that OCPs act as stress sensors by regulating the expression of various genes to protect cancer cells from genotoxic stress.
O-GlcNAc proteins:
RBM47, SBNO1, CNOT1, RGPD3, P121C, PDLI1, MYO1C, AIP, PSD11, PGRC1, TAF4, CLIC1, IPO5, IF2B3, AGRIN, PLOD2, HMGN4, IMA4, PESC, NOP56, DDX3X, PODXL, IMA3, NFIB, ARI1A, G3PT, PDCD5, TCRG1, PSA7, SCAM3, HGS, MYPT1, HNRDL, XPO1, ZN609, SC16A, SR140, SET1A, NPC1, TBX3, ARC1B, TIF1A, PGRC2, PFD6, NKRF, ZN185, OGT1, HMGB3, PPM1G, EIF3D, IPO8, RPA34, NUP42, DHX15, PRP4, SERA, PSMD3, RFOX2, PAPS1, MCA3, HNRPR, PRPF3, TPD54, IF4G3, KLF4, E41L2, DENR, XPOT, PRC1, ZN207, GET3, BUB3, ACTN4, BUD23, SYNC, KRT86, CPSF5, U3IP2, CALU, SAHH2, MED14, SMCA5, ZN862, GANP, KDM1A, ACSL4, SNX3, OGA, HNRPQ, PLOD3, MAFK, IMA7, UGDH, PQBP1, DKC1, IF2P, EDF1, DNJA2, BRD4, PFD1, WDR1, CPNE3, ZC11A, CLU, T22D2, PP6R2, CREST, ANR17, PDCD6, TBCA, H2AY, FLNB, NCOR1, SC22B, PR40A, PSIP1, SRS10, SF3B1, CSDE1, NPM3, U520, NU155, WDHD1, CRTAP, IDHC, CCNK, PIAS2, SPF27, DNJC8, RL1D1, SRP72, MTA2, TOM70, TOX4, SC24D, SUN1, NFAT5, AP2A2, SC31A, SEM3D, AGFG2, ZRAB2, LC7L3, LYPD3, FKBP9, SMC2, IPO7, AHSA1, PSMG1, SC24A, SC24B, CNOT4, OXSR1, HS74L, AP2A1, BAG3, CLPT1, ACL6A, LDHA, AATM, EGFR, PGK1, ASSY, LDLR, K1C14, LMNA, APOA2, FINC, ALBU, TFR1, PROC, ALDOA, CYTB, ANXA1, GCR, KITH, THY1, K2C1, G3P, HSPB1, RPN1, GNAI2, AT1A1, AT1B1, ADT2, IF2A, HMGN2, ICAM1, RLA2, JUN, LA, ITB1, K1C18, K2C8, CDK1, ATPB, S10A6, ENOA, PYGL, G6PI, NPM, ITAV, ACBP, LDHB, PDIA1, H10, CATD, ANXA2, TBB5, PROF1, SYEP, HS90A, HNRPC, TSP1, SP1, ANXA6, RHOC, DAF, MDR1, 4F2, HS90B, SRPRA, ASNS, CY1, RU17, ITA5, NFIC, VIME, RS17, K2C7, ANXA5, K1C16, RSSA, SNRPA, GSTP1, LEG1, HMGB1, TPM1, ROA1, RU2A, PARP1, PPBI, UCHL1, ALDOC, ATX1L, HS71B, CALM3, RO60, H14, PTPRF, THIO, ESTD, CH60, BIP, HSP7C, LAMP1, TOP1, TOP2A, PYC, C1TC, MPRI, ADHX, PABP1, PCNA, HARS1, IMDH2, TPR, KCRB, ACTN1, XRCC6, XRCC5, RINI, EF2, K1C10, K2C5, PDIA4, P4HA1, PLST, T2FB, CD59, MIF, GLU2B, CBPM, AK1A1, KPYM, ENPL, CCNB1, PO2F1, HNRPL, SYDC, PLAK, ALDR, AMPN, ERF3A, EZRI, FOSL1, FOSL2, MCP, NQO1, GNS, ZEP1, RS2, DESP, MUC1, CD44, CBR1, CREB1, H15, H13, H12, NCPR, AT2A2, CD36, STMN1, HSP76, HMGA1, JUNB, UBF1, JUND, ATF7, CEBPB, PYRG1, DDX5, PFKAL, LEG3, TCPA, PTN1, RL35A, RL7, VINC, SON, RL17, PGAM1, RCC1, ATF1, ML12A, NUCL, SPEE, RXRA, NFKB1, IF2B, ANXA7, BTF3, PSB1, MPRD, LMNB1, CSRP1, FLNA, 5NTD, VDAC1, CD9, TGM2, PIMT, FBRL, PUR2, PUR6, UBA1, NDKB, ROA2, RFX1, CBL, TCEA1, ITA6, SFPQ, PPIB, SYWC, RS3, NFYA, SAHH, COF1, IF4B, KTHY, EF1B, PPAC, CDK2, MCM3, RS12, BRD2, DNJB1, ATPA, PSA1, PSA3, PSA4, S100P, ITA3, MOES, DDX6, DNMT1, PAX6, U2AF2, RL13, S10A4, HMGB2, PTBP1, SYTC, SYVC, EF1G, STOM, 1433T, ARNT, RL10, RFA1, APEX1, PYR1, CALR, MAP4, CALX, TEAD1, GRN, EPHA2, 3MG, TKT, RBMS1, PML, EF1D, ERP29, PRDX6, RL12, KCY, PEBP1, PDIA3, 2AAA, NMT1, PURA2, UFO, SORCN, ILEU, RPB2, METK2, TIA1, ZEP2, DNJA1, PUR9, HNRH3, HNRH1, 1433B, 1433S, STIP1, S10AB, L1CAM, PRDX2, CDD, ELF1, RL9, CD70, KINH, CSTF2, MCM4, MCM5, MCM7, GLYM, HSP74, PROF2, PHB1, SPB6, RFC4, RL22, K1C9, MYH9, MYH10, COPB2, BASI, FUS, NU214, DEK, K22E, PRS7, ATPG, RL4, PP1G, GNL1, SRP14, NUP62, TAGL2, TALDO, RBMX, VKGC, HSPA9, IF4A3, RS19, RL3, OST48, FEN1, CAPG, TXLNA, TCPZ, RL13A, STAT3, MDHC, MDHM, IF2G, GARS, SYIC, LAP2A, LAP2B, MTREX, RS27, LPPRC, RL35, CDN2A, ECE1, LIS1, MUC18, MATR3, MSH2, SSRA, RANG, VDAC2, CBX5, UBP5, KI67, RAGP1, RECQ1, NOP2, BAG6, NOTC1, RL27A, RL5, RL21, RL28, RS9, RS5, RS10, IQGA1, CAPZB, IF1AX, RL29, SOX9, COPD, GSH0, PSMD8, PRC2A, TCPE, PTSS1, K2C6C, AGRE5, PAXI, RL34, LMAN1, NASP, FAS, CDK8, TCPG, EFTU, SYAC, SYSC, MCM2, ACADV, YLPM1, TMEDA, RBM25, HINT1, NU153, RBP2, TAF6, GUAA, CRIP1, GDIB, EMD, SERPH, F10A1, MAP2, RL14, TCPQ, TCPD, ANX11, PAPOA, FXR1, FXR2, RAB7A, SMCA4, SSRD, HDGF, ROA3, 6PGD, HNRPM, IMA1, GDIR1, AGFG1, HNRPF, MSH6, CAZA1, CRIP2, NUP98, ACLY, COPA, SC24C, TCP4, SYRC, ATX1, ATN1, SYYC, UBP14, AT1B3, RD23B, SNAA, IF5, PSMD4, XPO2, TERA, AF10, AF17, NP1L1, ADK, DSRAD, SEC13, NH2L1, PSA, EIF3B, SYMC, IF6, CTBP2, TMM33, NU107, EPIPL, TPIS, EIF3E, SC61B, MYL6, ACTB, IF4A1, RS20, PRPS1, PSA6, S10AA, CDC42, DEST, RAB10, UBC12, UBE2N, ARP3, ABCE1, RS3A, RL26, PSME3, RL15, RL27, RL37A, S61A1, PFD3, B2MG, DAD1, SUMO2, WDR5, NTF2, HNRPK, 1433G, RS7, PP1A, PP1B, RS8, RS15A, RS16, 1433E, RS14, RS23, RS18, RS29, RS13, RS11, RUXE, SMD1, SMD2, SMD3, PRS10, RL7A, ERF1, CNBP, RS4X, RL23A, RS6, H4, RAN, RL23, RAP1A, RS24, RS25, RS26, RS30, GBB2, RL30, RL31, RL10A, RL32, RL11, RL8, PPIA, FKB1A, RS27A, TRA2B, AP2B1, 1433Z, RSMN, SUMO1, DYL1, RL38, RS21, RACK1, UBC9, YBOX1, CSK2B, TPM4, EF1A1, ACTS, TBA1B, TBA4A, TBB4B, CSK21, PA1B2, HBB, HBA, PITX1, GTF2I, PHC1, TCPB, RAE1L, PRKDC, SARNP, RL24, ARF1, ERH, RL19, SRSF3, FOXK1, DAB2, EFNB1, RBM10, RBM3, CYC, S25A3, VIGLN, CLH1, FKBP3, HNRPU, U2AF1, SPTB2, TIAR, SRSF2, FOXK2, RUNX1, FABP5, LAT1, TFAP4, OTUD4, PFKAP, XPC, EWS, MEF2A, SP3, H11, RL18A, FKBP4, PLOD1, RL6, M2OM, DYST, KMT2A, LMNB2, TF65, UBXN1, GLGB, IF4G1, K1C17, REL, 1433F, PLP2, CSTF1, SRS11, EF1A2, SUH, GABPA, PAX8, FMR1, PRDX1, RL18, CKAP4, KHDR1, LRP1, SRSF1, DHX9, LG3BP, PPID, SSRP1, NSUN2, RBBP4, EP300, AHNK, HSP7E, GALT2, BST2, NU160, TBL3, ASPH, TROAP, BPTF, NFIA, SF3A3, AIMP1, ILF2, ILF3, LMAN2, TRAP1, FOXC1, MYO1E, CSTF3, ECH1, ACACA, CAF1B, RED, MTAP, TADBP, ROA0, PRDX4, CBX3, PSMD2, GPS2, SRSF9, SRSF5, SRSF6, TIF1B, G3BP1, PTK7, PABP4, EIF3I, TCOF, SF3B2, HAP28, FKBP5, SMAD4, PICAL, TBB3, PRP4K, PIN1, RIPK1, HDAC1, DCTN2, SNW1, TRA2A, CUL4B, DYR1A, TPBG, FHL1, MOGS, CD166, SPTN1, DX39B, TBB2A, KLF5, BYST, RUNX2, CDK13, CKAP5, CIRBP, HNRPD, SCRB2, DAG1, VEZF1, DSG2, EIF3A, UBP2L, SCRIB, TTL12, FHL2, DPYL3, DYHC1, IF4A2, SRC8, TRI25, FLNC, FA50A, CAPR1, RBM39, MCM6, ITPR1, PUM1, MDC1, EPN4, SMC1A, RRP1B, NCOA6, GSE1, UBP10, GANAB, LBR, MEF2D, CHD4, LASP1, ZN638, IMB1, NOLC1, NUMA1, SEPT2, SART3, CND1, ACAP1, U5S1, SYK, IF4H, PDIA6, PLEC, NOMO1, PON2, IPYR, TEBP, NONO, PWP2, RNPS1, PCBP1, PCBP2, SF3B3, KS6A1, SAFB1, SF3A2, RBMS2, SC23A, SC23B, SF3A1, SSXT, NCOA2, TRAM1, SF01, MED1, HMGN3, JHD2C, TRIP6, MARE1, ELAV1, ELF2, TAB1, AAAT, TOM34, UB2V2, NEDD8, ZYX, SEPT7, ADRM1, UAP1, PSMD5, DDB1, CDC37, DPYL2, RBBP7, TAF9, SRSF7, CPSF6, NRF1, FSCN1, IF16, KYNU, H2A2C, H2B2E, TRXR1, HNRL2, PDS5A, QSER1, TSR1, SMU1, P3H1, LSM12, CRTC2, GLE1, H90B2, ZN326, BCORL, TGO1, PRC2B, RRP12, TOIP1, PCID2, NU188, HP1B3, CE170, ZN362, ZC3HD, LRIF1, UBR4, UBAP2, KPRP, RBM26, AHDC1, CROCC, RPRD2, ECM29, MBNL2, ZMYM4, AR6P4, STEA4, ARID2, BICRL, CPIN1, LIN54, TM214, CAVN1, CDC73, EDC4, PRP8, SCYL2, GOLM2, NFRKB, NCEH1, MDEAS, ZC3HE, LARP1, FIP1, CRTC3, SAS6, CSPG4, WDR82, MCAF1, PACS1, SRCAP, RIPR1, UBN2, FBX50, IKIP, H32, LARP4, H2AV, RS27L, HAKAI, SPT6H, SND1, DDX46, 5MP2, CYFP1, KDM3B, PHF5A, ZCCHV, NUP54, POGZ, NUFP2, HEAT3, EMSY, RAI1, I2BP2, RBBP6, SH3R1, HUWE1, YTHD3, KHDC4, CENPV, KAISO, MYPN, PEG10, PABP2, KTN1, THOC4, GP180, CAND1, CARM1, PRSR1, DDX42, DAAF5, P66A, RB6I2, CHERP, ANKH1, CCAR1, RAVR1, SPB1, SMAP1, PHAR4, MAML2, PORIM, CCAR2, NUP93, LRC47, MT1M, FNBP4, CPSF7, PR38A, GT251, TXND5, SERB1, FA98A, TNR6A, PHC3, ABCF1, VP37A, NUP43, NUP37, NUP35, RLA0L, THOC2, WDR36, SMRC2, PUM2, SPP2A, ALMS1, C99L2, DDX54, DLG5, BRX1, DOT1L, PO210, GEMI5, ZN384, SCRB1, ZC3HF, NU133, PDC6I, NUDC2, GLMP, LMO7, ATX2L, PALLD, PSPC1, P66B, DNJC9, ELYS, DDX1, H1X, NICA, TM131, MAML1, HS105, CNOT9, ZN592, LAR4B, GCN1, NU205, TFG, TAF4B, STAM1, CBP, RBP56, DDX17, CELF1, OSTF1, RENT1, SMRC1, SMRD2, FUBP2, TNPO1, NEP1, UBP7, STMN2, LPP, MED12, H2A1C, SMCE1, RL36L, NCLN, FERM2, FUBP1, LRC59, FKB10, PF21A, INT12, FWCH2, AP2M1, REPS1, CMBL, SNR40, SEH1, DAZP1, RBM33, S10AG, OTUB1, HMCES, DDX27, P121A, PDLI5, FUBP3, WNK4, CHAP1, ZC3HA, CLP1L, CNDP2, ZFR, EP400, PRRC1, NOL4L, RBM14, QKI, PLIN4, S38A2, VPS35, CIC, MED15, MCCA, WRIP1, STRBP, TM209, SIN3A, MINT, UHRF1, HTF4, CDC5L, PFD5, RING2, EYA3, NUP88, POP1, MNT, SCAFB, EIF3C, DNJC7, PHB2, ATX2, ROAA, NP1L4, TS101, CPNE1, TCPH, EBP2, ANM1, H2B1M, RNZ2, TBA1C, MBB1A, TXD17, ERP44, ESYT1, WAC, DIDO1, AN32E, TMM43, TBB6, HNRL1, DDX23, TBB2B, TM109, TMED9, NUP58, GNL3, KIFC1, NUP85, RBM4, NAA15, SRRT, PDIP3, YTHD1, WNK3, GTPB4, UNK, SLIRP, IF5A2, NAT10, ILKAP, XRN2, SP130, RGAP1, DDX47, I2BPL, PININ, BOLA2, PTN23, WNK1, FA83D, ZHX3, STEEP, RPF2, ZN703, GORS2, JUPI2, MLXIP, CYBP, RC3H2, TENS1, EMAL4, NCOA5, TNR6C, CHD8, APMAP, ENY2, CD320, UBN1, DCP1A, LUC7L, RTN4, RPR1B, NIT2, ANLN, PDLI7, DDX21, SIAS, SHLB2, MBNL1, OSTC, RAB6B, PHP14, SYFB, RBM12, DD19A, LYAR, CARF, BCLF1, TAB2, TMOD3, CDK12, IF2B1, MAT2B, MYOF, ITSN2, BICRA, CNOT2, SEP10, RCC2, BCCIP, RRBP1, RBM27, KANL3, ATX10, MRC2, SAE2, NXF1, ABCF2, TES, TCF20, SUN2, LIMA1, CHRD1, SEPT9, UBQL2, S30BP, MED13, PFD2, PUF60, XPO7, SIX4, DDX41, CCNL1, JUPI1, MRT4, GPTC8, NUDT5, RALY, ACINU, AGO2, NUP50, ZHX1, CDV3, TRPC5, MRTFB, ZMIZ1, YETS2, HECD1, ZMYD8, COR1C, TPX2, AKP8L, PRP19, UBQL1, G3BP2, CHIP, PACN2, SSRG, EPCR, SCAF8, TRI33, SRRM2, PA2G4, CD11A, SMC3, RTRAF, RUVB2, EIF3L, RUVB1, NUDC, SYFA, DRG1, E41L3, RRP44, TR150, WBP11, NOP58, ZN281, SGT1, NOSIP, LSM2, LC7L2, SBDS, STRAP, RTCB, NOC2L, RL36, CHTOP, TLN1, ARIP4, HYOU1, PRC2C, PPME1, YTHD2, SP16H, TNPO3, SRPRB, RBM8A, ZN706, NCOR2, NPTN, COPG1, CLIC4, MD1L1, 5MP1, IF2B2, SCC4, ZHX2, S23IP
Species: Homo sapiens
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Hao Y, Fan X, Shi Y, Zhang C, Sun DE, Qin K, Qin W, Zhou W, Chen X. Next-generation unnatural monosaccharides reveal that ESRRB O-GlcNAcylation regulates pluripotency of mouse embryonic stem cells. Nature communications 2019 10(1) 31492838
Abstract:
Unnatural monosaccharides such as azidosugars that can be metabolically incorporated into cellular glycans are currently used as a major tool for glycan imaging and glycoproteomic profiling. As a common practice to enhance membrane permeability and cellular uptake, the unnatural sugars are per-O-acetylated, which, however, can induce a long-overlooked side reaction, non-enzymatic S-glycosylation. Herein, we develop 1,3-di-esterified N-azidoacetylgalactosamine (GalNAz) as next-generation chemical reporters for metabolic glycan labeling. Both 1,3-di-O-acetylated GalNAz (1,3-Ac2GalNAz) and 1,3-di-O-propionylated GalNAz (1,3-Pr2GalNAz) exhibit high efficiency for labeling protein O-GlcNAcylation with no artificial S-glycosylation. Applying 1,3-Pr2GalNAz in mouse embryonic stem cells (mESCs), we identify ESRRB, a critical transcription factor for pluripotency, as an O-GlcNAcylated protein. We show that ESRRB O-GlcNAcylation is important for mESC self-renewal and pluripotency. Mechanistically, ESRRB is O-GlcNAcylated by O-GlcNAc transferase at serine 25, which stabilizes ESRRB, promotes its transcription activity and facilitates its interactions with two master pluripotency regulators, OCT4 and NANOG.
O-GlcNAc proteins:
A0A087X1C1, A0A0B4J203, A0A0C4DFX4, SRCRL, SBNO1, CNOT1, IQCAL, RGPD3, LRIQ3, P121C, YU005, H0YHG0, H3BMQ9, H8Y6P7, I3L521, PDLI1, TAF4, P3C2A, DDX3X, NFIB, ARI1A, ABLM1, KMT2D, ZN197, RGPD8, MYPT1, ZN609, ZN646, SET1A, SYNEM, ZN185, NUP42, SI1L1, TGFI1, VIP2, M3K7, TPD54, SYNJ1, IF4G3, WIPF1, SMAD6, MYPT2, PLIN3, MAFK, N4BP1, ANR17, NCOR1, GGYF1, PRDM1, STAM2, TOX4, AGFG2, UNC5C, VENTX, SC24B, PCNT, ZBT11, CNOT4, BAG3, TXD12, TTLL1, APCL, NSD2, EGFR, IGF2, CRYAB, LMNA, ALDOA, GCR, HSPB1, PCCA, RLA2, JUN, HEP2, K1C18, ANXA2, ADRB2, NFIC, VIME, 5HT1A, SNRPA, ROA1, CO4B, ATX1L, FMAS1, GRL1A, DERPC, C1C1L, GLI2, TPR, GYS1, MYL6B, PO2F1, ATF2, ZEP1, RS2, ITB4, JUNB, JUND, ATF7, SON, ATF1, NEBU, CSRP1, NF1, ROA2, RFX1, CBL, COF1, IF4B, ARNT, MAP4, CALX, TEAD1, PDIA3, CDC27, CLIP1, ZEP2, ELF1, TTK, PCKGC, ADDA, NU214, MP2K2, NUP62, VKGC, VATA, CUX1, TXLNA, PBX2, HELZ, UTRN, RFX5, PAXI, NR2C2, NASP, CENPF, YLPM1, NU153, RBP2, TAF6, EMD, PAPOA, AGFG1, NUP98, ATX1, MYOM2, AF10, AF17, DSRAD, NU107, RL8, F193A, PITX1, PHC1, KGD4, SARNP, FOXK1, HTD2, DAB2, RHG04, VIGLN, HNRPU, SPTB2, SCN7A, FOXK2, EWS, MEF2A, SP2, NUCB1, IF4G1, NOTC2, TLE4, NMDZ1, GABPA, ZO1, ACK1, CACB2, EP300, AHNK, MGAT2, GALT2, FOXO1, SNF5, BPTF, NFIA, DPYD, TP53B, ZN155, FOXC1, AKAP6, ROA0, GPS2, G3BP1, KCAB2, PABP4, PLD1, PICAL, MAMD1, RIPK1, SNW1, MTMR1, CUL4B, ASPP2, NFYC, CDK13, TOB2, DAG1, VEZF1, DSG2, UBP2L, GIT2, SRC8, MDC1, EPN4, RRP1B, NCOA6, GSE1, MEF2D, ARI5B, NUMA1, PSME4, SART3, KIF14, BRD3, EBP, PLEC, RBMS2, TAF1C, SF01, MED1, JHD2C, MARE1, ELF2, TAB1, ZFHX3, ZYX, ADRM1, TAF9, RFX7, QSER1, QRIC1, ZN800, LR75B, EPC2, CRTC2, YIF1B, ZN326, BRD10, PRC2B, SYAM, CE170, NHSL1, ZN362, ZEP3, LRIF1, UBR4, SKT, RHG21, UBAP2, RBM26, RC3H1, VP13D, RPRD2, ZN318, TASO2, TTC23, ERR2, KAZRN, ARID2, RHG17, ANR40, BICRL, NIPBL, LIN54, NFRKB, ZN449, RSBNL, MDEAS, ZC3HE, CRTC3, SAS6, MCAF1, BCOR, MPRIP, GGYF2, BNC2, FGD5, CO039, SRCAP, YJ005, UBN2, RAPH1, UBP54, PREX2, HAKAI, ASXL2, SPT6H, KDM3B, ZCCHV, RGPD4, POGZ, SZRD1, MAVS, EMSY, RAI1, I2BP2, SRGP1, SH3R1, YTHD3, STRA8, MYPN, TLK2, ZFHX4, BCL9L, IQCH, SNX32, PRSR1, MEX3D, DDX42, CACL1, P66A, CC125, KCC1D, HID1, AHNK2, FOXP4, NAV2, MGAP, RP1L1, PHF6, ANKH1, SUGP1, CCAR1, RPAP2, MILK2, EFC13, PHAR4, XRN1, TBC3A, TENS4, SPART, NUP93, ZN687, SYNPO, FNBP4, TAB3, CPSF7, ARFG1, ENAH, XXLT1, CHSTE, TNR6A, PHC3, SP20H, NAV1, VP37A, KMT2C, ARI1B, NUP35, TDRD7, NEDD1, PUM2, ALMS1, DLG5, ZN384, WIPF2, FRS2, F222B, SMAP2, IASPP, ZFN2B, TWST2, PCNP, LMO7, ATX2L, STAB2, PALLD, CSKI2, SRS12, P66B, BBX, SMG7, RTF1, PHF3, MAML1, LAR4B, PRP16, PRCC, CBP, EVPL, DDX17, GPKOW, FUBP2, LPP, FUBP1, TTC28, PF21A, KLH29, RBM33, MMAB, EF2KT, GWL, P121A, PDLI5, INT4, FUBP3, PAWR, ANCHR, Z512B, ZFR, EP400, COG8, RBM14, QKI, LENG8, CIC, MED15, ERBIN, MINT, HTF4, TEAD3, RGPD5, CGRE1, ATX2, CTIP, SH3G1, DPH2, WAC, DIDO1, TBCD, HNRL1, HIRP3, CAR11, YTHD1, GTPB4, AMRA1, TANC1, CEP44, MFF, SP130, BRD8, RGAP1, I2BPL, RBNS5, ADNP, FOXP1, PTN23, CA198, WNK1, E41L1, ZHX3, ILRUN, PEAK1, PKHG2, ECT2, JUPI2, PKHA5, RC3H2, TAF9B, ZBT20, NCOA5, TANC2, ZN532, ARFG3, PLPL8, UBN1, PCD12, INCE, PDLI7, 4ET, DIAP3, PDS5B, RBM12, CCD87, CARF, TAB2, MS18A, CDK12, ITSN2, WHRN, DAPLE, SLAI2, BAHC1, BCCIP, C2D2A, RBM27, KANL3, ZN219, LIMD1, TCF20, UBQL2, S30BP, NRBP, CTNA3, BAZ2B, HERC5, SIX4, TASOR, GMEB2, PARP4, NUP50, ZHX1, MRTFB, PRR12, YETS2, HECD1, SPAT2, SCAF8, LIMC1, ZC3H4, SRRM2, SCML2, ZN148, ZFP30, WDR37, MYH15, R3HD2, ZN281, COQ6, DCAF1, RPGF2, CRBG1, PRC2C, RBM7, CD2AP, TSSC4, HCFC2, NCOR2, GMEB1, DC1L1, NCOA3, ZHX2, S23IP, U3KPZ7
Species: Mus musculus, Homo sapiens
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Levine ZG, Fan C, Melicher MS, Orman M, Benjamin T, Walker S. O-GlcNAc Transferase Recognizes Protein Substrates Using an Asparagine Ladder in the Tetratricopeptide Repeat (TPR) Superhelix. Journal of the American Chemical Society 2018 140(10) 29485866
Abstract:
The essential mammalian enzyme O-GlcNAc Transferase (OGT) is uniquely responsible for transferring N-acetylglucosamine to over a thousand nuclear and cytoplasmic proteins, yet there is no known consensus sequence and it remains unclear how OGT recognizes its substrates. To address this question, we developed a protein microarray assay that chemoenzymatically labels de novo sites of glycosylation with biotin, allowing us to simultaneously assess OGT activity across >6000 human proteins. With this assay we examined the contribution to substrate selection of a conserved asparagine ladder within the lumen of OGT's superhelical tetratricopeptide repeat (TPR) domain. When five asparagines were mutated, OGT retained significant activity against short peptides, but showed limited limited glycosylation of protein substrates on the microarray. O-GlcNAcylation of protein substrates in cell extracts was also greatly attenuated. We conclude that OGT recognizes the majority of its substrates by binding them to the asparagine ladder in the TPR lumen proximal to the catalytic domain.
O-GlcNAc proteins:
E2F8, PRXD1, TSPY2, CGB1, ZN839, CENPX, PDLI1, WWP2, FOXN3, IMA4, DPYL4, ABLM1, MEIS2, HGS, GAK, PLS1, MAGB4, ARPC5, DC1L2, FGF16, WIPF1, FRS3, RNF13, ZN207, ENSA, SMCA5, CE104, OGA, AP180, PQBP1, E2F6, ADAP1, DNJC8, FLOT1, TPPP, SCEL, CELF2, RPGF3, CLD14, KLF8, BAG3, PAK4, IGHG3, MBP, NFIC, SNRPA, CEL3A, RU1C, IGLC1, MTAP2, ERG, DMD, IRF2, ENPL, FOSL2, F261, SDC1, ATF1, ATF3, ELK1, TFEB, RXRA, COMT, USF1, FGFR3, NR4A1, THAS, MARK3, ARNT, PSB4, RBMS1, FCERG, MPIP1, CSTF2, CD68, BMI1, COPB2, SOX5, RS19, ARL4A, ETV5, IPP2, STAT6, ABL2, KSYK, TFPI2, RFX5, RXRG, CDK8, GSK3B, TAF6, SPS1, DNLI3, DYN2, GALK1, AGFG1, SC24C, DLX1, HDAC4, CTBP2, KLF3, TIRAP, PPIA, CSK21, 3BP2, SSBP2, CDKL1, BORG5, TFAP4, ROR2, CACB1, CREM, MECOM, TLE4, FAK1, COEA1, MTG8, GABPA, GABP1, DERM, TLE5, DEMA, ASPH, PP1R8, LCP2, DUS4, MTAP, EI2BE, G3BP1, GRB10, MTA1, PPIL2, PICAL, DYR1A, FHL3, NCOA4, CAYP1, CIRBP, DGCR6, VGLL4, CAPR1, GRB7, CASL, MEF2D, LASP1, SPCS2, SEPT2, PCM1, SIX1, SF01, TRIP6, RHOH, TAB1, ADRM1, CCDC6, BATF, NFE2, MAR1, NRF1, SMTL2, LETM2, OD3L2, AX2R, AMOT, AKIR2, PRR30, LARGN, RGPS1, PRC2B, TEX30, SNP47, PHYD1, KANK4, EEIG1, CPTP, AHDC1, KHDR2, EF1DL, OLM2B, ANKS6, RHG17, CNDH2, GRHL2, HMBX1, JHY, PP4R4, JMJD6, SCYL2, GOLM2, RSBNL, PSRC1, DRIC1, KCD18, FIP1, SAS6, CLM9, CCBE1, DUSTY, E400N, TM1L2, UBP45, MARK2, KCTD9, MCM10, CC186, POGZ, FLIP1, TAF8, RM55, S1A7A, SYVN1, PHLB1, CASZ1, PIM3, GA2L3, DDX42, CEP57, SPAS2, Z280B, PDPN, KCC1D, TF2LX, LOXH1, ARI3B, APLF, CC28A, RHG12, SKA3, PHC2, HYCC2, CMIP, SMAP1, RUSD2, ABI1, SYCE1, RPTOR, BATF2, S35F6, KLH36, ANS4B, MARHA, ZSCA1, UXS1, NECP1, BBOF1, LIPB2, MISSL, PHC3, TMIE, NUP35, OR4D6, JOS2, WDR48, GABP2, DPTOR, ALAT2, BRSK1, GRP4, HELQ, BL1S5, BCAS4, LMX1A, SSH3, SH3R2, FBF1, DLRB2, HDAC7, STYX, GTSF1, NEUR4, CSRN3, AP2B, SHC3, RORB, STAM1, CELF1, ARHG1, CSN5, RBPMS, ZDH16, PMEPA, UTP4, FERM2, SOSB2, R51A1, CR025, LENG1, BIRC7, PBIR1, CC126, PF21B, NRBF2, EDC3, PDLI5, ANCHR, ZC3HA, PKNX2, EGLN2, MARK4, UB2E2, PRRC1, NOL4L, AP4AT, SCLT1, TOX2, QKI, PNMA5, YPEL2, NUDC1, RB40C, UBL7, CK2N2, OSBL9, PNKP, PLIN2, CDCA3, HCD2, SNAG, ARI3A, DPYL5, CG025, SOSB1, TCF25, COE4, MIC25, UTP23, ALKB7, RBM42, LRRC1, TRMO, MCRI2, RSRP1, BBLN, ASHWN, SARG, NADAP, SSBP3, PLVAP, OSB11, B2L13, STK31, DGC6L, RM01, KLC2, SIK2, REN3A, NELFA, PPDPF, ILRUN, ES8L2, KI3X1, SP14L, B2L12, PKHA3, PKHA1, TAF9B, ARNT2, SYTL2, CIP1, CASS4, HINT3, PRDM5, PAR11, MBNL1, BRWD1, KLC4, TB22B, MBNL3, NFIP2, ZSC32, TMOD3, UGGG2, CWC15, KANL3, RBPJL, DKK4, LIMD1, DPOLL, AFF4, UBQL2, PUF60, BAZ2B, GGA2, ING1, BORG2, AGO2, ZHX1, MUCEN, ANR50, PCDGK, POLI, LIMC1, SMAG1, BAIP2, RPC10, PPIL1, C2CD2, MTL5, TBL2, YTHD2, T10B, SNX14, RT18B, GMEB1, STON1, PCLO, ZHX2, S23IP
Species: Homo sapiens
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Qin K, Zhu Y, Qin W, Gao J, Shao X, Wang YL, Zhou W, Wang C, Chen X. Quantitative Profiling of Protein O-GlcNAcylation Sites by an Isotope-Tagged Cleavable Linker. ACS chemical biology 2018 13(8) 30059200
Abstract:
Large-scale quantification of protein O-linked β- N-acetylglucosamine (O-GlcNAc) modification in a site-specific manner remains a key challenge in studying O-GlcNAc biology. Herein, we developed an isotope-tagged cleavable linker (isoTCL) strategy, which enabled isotopic labeling of O-GlcNAc through bioorthogonal conjugation of affinity tags. We demonstrated the application of the isoTCL in mapping and quantification of O-GlcNAcylation sites in HeLa cells. Furthermore, we investigated the O-GlcNAcylation sensitivity to the sugar donor by quantifying the levels of modification under different concentrations of the O-GlcNAc labeling probe in a site-specific manner. In addition, we applied isoTCL to compare the O-GlcNAcylation stoichiometry levels of more than 100 modification sites between placenta samples from male and female mice and confirmed site-specifically that female placenta has a higher O-GlcNAcylation than its male counterpart. The isoTCL platform provides a powerful tool for quantitative profiling of O-GlcNAc modification.
O-GlcNAc proteins:
A0A0A6YVU8, A0A1B0GSG7, RBM47, ZN335, A2A8N0, SBNO1, CNOT1, PHRF1, ZN462, TAGAP, D3YUK0, E9PUR0, E9PVW1, E9PWI7, PARP4, E9PZS2, E9Q2C0, NU153, E9Q616, BD1L1, E9Q732, ARHG5, E9Q7N9, E9Q842, E9Q9B4, E9Q9Q2, E9QA22, E9QAE1, F6Y6L6, F8VQ29, F8VQM5, J9JI28, PDLI1, SPT5H, TAF4, ARI1A, ABLM1, KMT2D, MYPT1, ZN609, SET1A, SYNEM, PUR4, TNC18, KDM6A, DPOD2, M3K7, TPD54, SYNJ1, ZN207, SRPK2, ACK1, SYUA, MYPT2, KIF1B, HBP1, OGA, VINEX, PLIN3, MAFK, BRD4, PDLI1, KDM6A, SRPK1, N4BP1, ANR17, NCOR1, CREG1, CRTAP, MYO1A, MTR1L, CREG1, TOX4, SUN1, M3K6, PSMG1, SC24B, CNOT4, ABL1, ABL1, EGFR, LAMC1, LMNA, GBA1, GCR, HSPB1, PPBT, RLA2, ITB1, K1C18, K2C8, SAP, CATL1, LAMB1, ENPL, BGLR, NFIC, VIME, SNRPA, ROA1, ATX1L, TGAP1, GLI2, HLAC, CATB, TAU, BIP, FINC, K2C8, TPR, MSH3, ENPL, PO2F1, ATF2, GNS, ZEP1, RS2, MUC1, JUNB, ATF7, CATD, SON, SERPH, NELFE, BIP, ROA2, CBL, IF4B, APC, ARNT, MAP4, TEAD1, RXRA, RXRB, RXRG, CLIP1, AIMP1, HXA11, ELF1, NU214, MP2K2, VATA, CUX1, PBX2, MLH1, STAT3, SSRB, KI67, STT3A, RFX5, LMNA, DPOD2, PAXI, CDK8, YLPM1, NU153, RBP2, TAF6, EMD, PPT1, FXR1, ICAL, AGFG1, NUP98, ATX1, ATN1, PTN5, AF17, DSRAD, AMRP, ACYP2, NU107, ACOT8, S26A1, TB182, YTHD1, ASXL1, PI5PA, RIN3, MRTFB, RL37, KCNA2, RALA, STIM1, PITX1, IF4G2, SRPK2, RENBP, COG7, WNK1, SERF2, RPTN, SPSY, DAB2, RBM10, HNRPU, SPTB2, FOXK2, EWS, MEF2A, SP2, CO7A1, S30BP, NUCB1, ENL, IF4G1, K1C17, TLE4, TOP1, SUH, CBG, ACK1, DEMA, AHNK, FOXO1, TROAP, BPTF, NFIA, ROA0, G3BP1, PABP4, ATM, PICAL, MAMD1, RIPK1, STIM1, MTMR1, CUL4B, ASPP2, KLF5, NFYC, CDK13, VEZF1, DSG2, TRI29, UBP2L, SRC8, PUM1, EPN4, RRP1B, NCOA6, DIP2A, MEF2D, NUMA1, R3HD1, KIF14, EBP, RCN1, KS6A1, RBMS2, TAF1C, NCOA2, SF01, JHD2C, MARE1, ELF2, TAB1, ZFHX3, ZYX, ADRM1, CCDC6, TAF9, STX1A, RFX7, QSER1, QRIC1, PRC2C, PBIP1, GSE1, TNR6A, CEAM5, Q3UKP4, COBL1, ARH40, SC31A, PEG3, SRBS2, Q3UU43, Q3UUE0, F91A1, ARBK2, Q497W2, Q4KL65, PHAR4, EPC2, CRTC2, BCORL, BRD10, TGO1, PRC2B, TOIP1, SPG17, SHRM1, ZN362, LRIF1, RHG21, UBAP2, RBM26, RPRD2, ZN318, NCOR1, LAMA5, HCFC1, P3C2A, SAP, AP180, MAFK, SPTB2, SH3G1, ZYX, TSH3, INADL, WAPL, KAZRN, SBNO1, ARID2, DYH17, SAM9L, CDK13, LAR4B, BICRL, RHG21, HELZ, TTLL5, PANX2, PKHG2, NIPBL, LIN54, F135A, RPRD2, IF4G1, SPIC, SCYL2, NFRKB, INT1, ZN182, UGGG1, MDEAS, ZC3HE, RICTR, FIP1, CRTC3, SAS6, MCAF1, BCOR, GGYF2, NU188, CO039, UBN2, HAKAI, ASXL2, SPT6H, DDX46, KDM3B, PRC2B, OOG2, ZIC5, NRK, POGZ, MAVS, CLAP1, EMSY, I2BP2, SRGP1, SH3R1, HUWE1, YTHD3, NU214, TMC5B, ZN598, TOPRS, SHAN2, Q80ZX0, ZNF18, Q810G1, BCL9L, LUZP1, PRSR1, DDX42, PALB2, P66A, GNS, LPP, TB10B, TGO1, Q8BIB6, ZN771, ZNT6, AAPK2, CNOT4, SP110, IFFO1, YTHD3, NCBP3, DEFI6, RBM14, CNOT2, CABS1, Q8C6L9, TCAL5, TAB1, SCYL2, ASPP2, PHC3, EPN2, PDLI5, I2BP1, RN135, AHNK2, NAV2, MISP, MGAP, ANKH1, PHAR4, XRN1, PELP1, Q8JZK6, Q8K0U8, AGFG1, TXD11, IL23R, ARHG6, SPART, SPICE, NUP93, CLASR, ZN786, SYNPO, FNBP4, ARFG1, ENAH, TNR6A, PHC3, SP20H, NAV1, VP37A, KMT2C, BD1L1, NUP35, STXB6, KNL1, TCAL3, MTSS1, SPART, NUP35, PUM2, STT3B, ALMS1, GEMI5, WIPF2, MAVS, UTP6, PI3R4, AMOT, P66B, STAG1, PCNP, LMO7, ATX2L, CSKI2, P66B, BBX, TITIN, HNMT, UBAP2, DCP1A, NRIF1, SMG7, RTF1, MAML1, ZN592, LAR4B, TAF4B, SHIP1, DDX17, RENT1, GPKOW, FUBP2, LPP, TTC28, PF21A, INT12, RCN3, CERS2, PDLI5, FUBP3, MY15B, ANCHR, CLP1L, Z512B, ZFR, EP400, NOL4L, RBM14, CIC, MED15, PIGS, DCR1C, SIN3A, MINT, EYA3, TEAD3, ATX2, RFC4, DHX58, ANX13, GORS2, TAB2, EPN4, ANR17, DPH2, WAC, DIDO1, YTHD1, AMRA1, TANC1, TXD12, F133B, RBM33, GPI8, Q9D2U0, ZC21B, FUND2, F162A, APMAP, Q9D809, FIP1, CNPY3, Q9DAV5, Q9DB24, ALG2, PLIN3, MYPT1, WWTR1, Q9EQC8, SALL1, RBP2, GILT, MFF, SP130, APC1, I2BPL, RBNS5, EPC1, ADNP, ZN106, TM245, CPVL, PTN23, WNK1, E41L1, ZHX3, ZN335, PKHG2, CCSE2, CQ10B, MLXIP, PKHA5, RC3H2, TAF9B, ZBT20, NCOA5, ZN532, APMAP, HYOU1, ADRM1, GIT2, BAG3, UBN1, PDLI7, DIAP3, RBM12, CARF, ETAA1, HXC10, TAB2, UGGG1, CDK12, ITSN2, CNOT2, TMEM9, DAPLE, NYAP2, KANL3, SON, LIMD1, KI21B, KI21A, PPIE, PCM1, GALK1, MRP5, SE1L1, LIMD1, TCF20, SUN2, AFF4, UBQL2, S30BP, NRBP, SIX4, TASOR, GMEB2, PARP4, NUP50, ZHX1, YETS2, HECD1, SCAF8, SRRM2, SCML2, S22AL, NCOR2, DEMA, POLH, R3HD2, ZN281, FBX7, RPGF2, IRS2, HYOU1, PRC2C, NCOR2, GMEB1, S23IP, SRPK3, SLFN1, VNN1, KLK4, SE1L1, RGS6, E41L1
Species: Mus musculus, Homo sapiens
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