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Xu S, Yin K, Xu X, Fu L, Wu R. O-GlcNAcylation reduces proteome solubility and regulates the formation of biomolecular condensates in human cells. Nature communications 2025 16(1) 40307207
Abstract:
O-GlcNAcylation plays critical roles in the regulation of protein functions and cellular activities, including protein interactions with other macromolecules. While the formation of biomolecular condensates (or biocondensates) regulated by O-GlcNAcylation in a few individual proteins has been reported, systematic investigation of O-GlcNAcylation on the regulation of biocondensate formation remains to be explored. Here we systematically study the roles of O-GlcNAcylation in regulating protein solubility and its impacts on RNA-protein condensates using mass spectrometry-based chemoproteomics. Unexpectedly, we observe a system-wide decrease in the solubility of proteins modified by O-GlcNAcylation, with glycoproteins involved in focal adhesion and actin binding exhibiting the most significant decrease. Furthermore, O-GlcNAcylation sites located in disordered regions and with fewer acidic and aromatic residues nearby are related to a greater drop in protein solubility. Additionally, we discover that a specific group of O-GlcNAcylation events promotes the dissociation of RNA-protein condensates under heat stress, while some enhance the formation of RNA-protein condensates during the recovery phase. Using site mutagenesis, inhibition of O-GlcNAc transferase, and fluorescence microscopy, we validate that O-GlcNAcylation regulates the formation of biocondensates for YTHDF3 and NUFIP2. This work advances our understanding of the functions of protein O-GlcNAcylation and its roles in the formation of biomolecular condensates.
O-GlcNAc proteins:
MEX3A, CNOT1, RGPD3, P121C, MYO1G, H7C0N4, PDLI1, CBX4, TAF4, RFXAP, PODXL, ARI1A, UTY, ABLM1, DVL1, KMT2D, HGS, MYPT1, XPO1, SET1A, PUR4, FYB1, PPM1G, EIF3H, IPO8, NUP42, KDM6A, ZZEF1, TET3, SI1L1, CP110, SYNJ1, IF4G3, TGON2, WIPF1, HTSF1, MYPT2, SMCA5, GANP, MA1A2, DIAP1, PLIN3, MAFK, KIN17, CUTA, N4BP1, CLAP2, BRE1B, PP6R2, ANR17, ZN217, FLNB, NCOR1, SPAG7, TACC1, GGYF1, ERLN1, WBP4, U520, WDHD1, RNH2A, STAM2, CBPD, STK10, TOX4, ST65G, TOX, PCF11, ENDD1, UBR5, AGFG2, KIF4A, MPZL1, IPO7, BAG4, SC24B, HERC2, ZMYM6, BAG3, CLPT1, ABL1, ADA, LDLR, HLAB, TFR1, CATA, GCR, KITH, HLAA, RPN2, ITB2, RLA2, RLA0, ITB1, RB, TPM3, SAP, LAMB1, SP1, PTPRC, ITA5, VIME, SNRPA, ROA1, ATX1L, DERPC, MYBA, CH60, PABP1, TPR, SKI, XRCC6, LAMP2, ITA4, HCLS1, KPYM, PO2F1, RAC2, IGLL1, RS2, TFE2, LEUK, CREB1, GCFC2, ATF7, SON, ATF1, TFE3, NFKB1, AL5AP, PUR2, ROA2, CBL, IF4B, AT2B4, APC, MGAT1, ARNT, DCK, PYR1, CALR, MAP4, CALX, NDUS1, PML, PDIA3, HMOX2, AMRP, CLIP1, ZEP2, HNRH1, ELF1, ICAM3, TTK, CAH8, BMI1, PTN7, RFC1, ADDA, NU214, MP2K2, HPCL1, VATA, CUX1, TXLNA, MLH1, PEX19, S19A1, LAP2A, LAP2B, STAT1, EPS15, MATR3, KI67, CRKL, UTRN, CAPZB, RFX5, PRC2A, NASP, FAS, CDK8, YLPM1, NU153, RBP2, TAF6, EMD, TCPQ, PAPOA, PLCD1, HCFC1, AFF3, ROA3, AGFG1, RBM5, ACLY, SMTN, RD23A, AF10, DSRAD, CADH4, SEC13, IF6, STX17, CORO7, NU107, SELB, TPIS, RAB2A, DCAF7, TIM10, CNBP, TBA1B, NOMO3, F193A, IGBP1, IF4G2, GTF2I, PHC1, NUCB2, TMF1, RT09, SARNP, FOXK1, FBLN2, FAM3A, RBM10, CDK6, HNRPU, SPTB2, FOXK2, RUNX1, CAP1, MEF2A, SP2, KMT2A, IF4G1, TLE1, TLE2, TLE3, TLE4, REL, GFPT1, GABPA, RING1, VAC14, EP300, AHNK, BRME1, MGAT2, GALT2, TROAP, TROP, BPTF, SFSWA, DPYD, TP53B, IRAG2, NFAC3, ROA0, AIMP2, GPS2, PABP4, S39A6, PICAL, HDAC1, STIM1, MTMR2, CUL4B, ASPP2, PEBB, NFYC, LRP8, VEZF1, BOP1, SEPT6, UBP2L, MLEC, ZMYM3, SRC8, LAGE3, PUM1, EPN4, RRP1B, NCOA6, GSE1, MEF2D, NFIX, ZN638, NUMA1, CND2, CND1, RB3GP, NECT1, PCBP1, PCBP2, ELOC, KS6A1, SC23B, JHD2C, ELF2, TAB1, NCOA1, ZYX, ADRM1, DPYL2, TAF9, DREB, DGUOK, QSER1, RHG31, AAK1, QRIC1, LRRF1, GNPTA, C2CD3, EPC2, PDCD4, BOREA, PAN3, BRD10, TGO1, PRC2B, PCID2, P4R3B, DCA10, SNP47, MANEA, ZN362, LRIF1, CAF17, CAMP1, UBAP2, CPTP, RC3H1, AHDC1, VPS53, RPRD2, ZN318, TASO2, ZMYM4, PAPD7, RN123, MAP1S, ARID2, NIPBL, LIN54, ANR54, PPR18, EDC4, SCYL2, GOLM2, NFRKB, RSBNL, MDEAS, BRAT1, ZC3HE, LARP1, FIP1, CRTC3, SHSA2, PI16, MCAF1, BCOR, GGYF2, NFXL1, QSOX2, YJ005, UBN2, MTSS2, NOL8, SSH2, ASXL2, SPT6H, LRC8D, EIF3M, MEPCE, KDM3B, TRM1L, ZCCHV, EFL1, KANL1, POGZ, ZFY16, NUFP2, MAVS, EMSY, I2BP2, HUWE1, YTHD3, KAISO, LDB1, LYRIC, RETR3, TSYL5, DZIP3, CACL1, P66A, C2CD5, HIPK1, ERI1, DEN1C, VRK3, ARI3B, MGAP, Z3H7A, CDAN1, ANKH1, SUGP1, GLT12, K2013, PHAR4, DCP1B, XRN1, RPTOR, NUP93, ZN687, FNBP4, NOPC1, ZBTB2, EMC1, CB069, EIF1A, KRI1, AFG2A, NHLC2, AVL9, XXLT1, GOLM1, SUMF2, TXND5, KCT2, CHSTE, SLAI1, TNR6A, PHC3, SP20H, KMT2C, BD1L1, ARI1B, NUP35, KNL1, MSPD2, NEDD1, GABP2, PIGX, STT3B, ALMS1, NEK9, NEK7, PO210, GEMI5, PIGO, MZB1, RAB2B, TM263, NUDT8, ZFN2B, HNRLL, PCNP, LMO7, ATX2L, P66B, GBF1, SMG7, RTF1, TOPB1, TSC1, PRCC, RREB1, CBP, DDX17, SEM4D, GSLG1, RENT1, GPKOW, FUBP2, ARHG2, UBP7, LPP, IGSF8, MRTFA, ACD, RAB37, PF21A, DOC10, DCPS, FAF2, TMCO6, ESS2, RMI2, TONSL, PDLI5, ZN414, VCIP1, PDLI2, ANCHR, ZFR, EP400, PRRC1, RBM14, QKI, PURB, SPAG5, CIC, MED15, ERGI2, ERBIN, LYSM1, MYADM, SEBP2, RUFY1, MINT, HTF4, CNN2, AGAP2, CSN8, CGRE1, DOK1, CTIP, SMAD5, S29A1, ARI3A, ANM1, DPH2, PELO, ERP44, ESYT1, CNPY3, WAC, MMTA2, HNRL1, ALG8, TMUB1, YJU2, PAPOG, SSBP3, JAM3, OSB11, NAA15, EIF2A, MK67I, YTHD1, KPCD2, UCK2, FTO, AMRA1, CEP44, BC11B, YIPF3, BRD8, XYLT2, I2BPL, MEGF9, EPC1, SLK, ADNP, ZN106, TM245, TMX1, CENPH, WNK1, ZHX3, DOCK5, PKHG2, KANL2, PKHA5, CLSPN, MYG1, RC3H2, LY9, TAF9B, S38AA, TM165, NCOA5, TANC2, TNR6C, CHD8, RESF1, APMAP, ARFG3, UBN1, GP108, INCE, ANLN, GEPH, BMP2K, RBM12, STAU2, INT10, GPT2L, CARF, PP4R2, TAB2, RM39, EURL, CDK12, HPBP1, ITSN2, ARHGC, CHMP5, FOXJ2, SLAI2, CT2NL, TBC14, RBM27, KANL3, FRM4A, ANFY1, GMPR2, ZN219, SE1L1, B4GT7, STK39, SEC63, SUN2, SRP68, SEPT9, UBQL2, S30BP, PFD2, NRBP, EI2BD, BTNL2, CDC23, GGA2, GMEB2, DP13A, MA1B1, ACINU, NUP50, MRTFB, ZBT21, YETS2, HECD1, TPX2, MGT4A, SYNRG, MINP1, SCAF8, SET1B, TBD2B, DICER, SRRM2, SMC3, SCML2, ZN148, CLC11, POLH, PLAP, TPPC8, WBP11, GIT1, ZN281, SIT1, HBS1L, SALL2, TRRAP, ARIP4, HECD4, CRBG1, HYOU1, ATG4B, KLF12, PRC2C, RBM7, CD2AP, SRPRB, GMEB1, TACC3, STK24, NCOA3, M3K4, ZHX2, NS1BP
Species: Homo sapiens
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Zhang W, Liu T, Dong H, Bai H, Tian F, Shi Z, Chen M, Wang J, Qin W, Qian X. Synthesis of a Highly Azide-Reactive and Thermosensitive Biofunctional Reagent for Efficient Enrichment and Large-Scale Identification of O-GlcNAc Proteins by Mass Spectrometry. Analytical chemistry 2017 89(11) 28510447
Abstract:
O-linked β-N-acetylglucosamine (O-GlcNAc) is a ubiquitous post-translational modification of proteins in eukaryotic cells. Despite their low abundance, O-GlcNAc-modified proteins play many important roles in regulating gene expression, signal transduction, and cell cycle. Aberrant O-GlcNAc proteins are correlated with many major human diseases, such as Alzheimer's disease, diabetes, and cancer. Because of the extremely low stoichiometry of O-GlcNAc proteins, enrichment is required before mass spectrometry analysis for large-scale identification and in-depth understanding of their cellular function. In this work, we designed and synthesized a novel thermosensitive immobilized triarylphosphine reagent as a convenient tool for efficient enrichment of azide-labeled O-GlcNAc proteins from complex biological samples. Immobilization of triarylphosphine on highly water-soluble thermosensitive polymer largely increases its solubility and reactivity in aqueous solution. As a result, facilitated coupling is achieved between triarylphosphine and azide-labeled O-GlcNAc proteins via Staudinger ligation, due to the increased triarylphosphine concentration, reduced interfacial mass transfer resistance, and steric hindrance in homogeneous reaction. Furthermore, solubility of the polymer from complete dissolution to full precipitation can be easily controlled by simply adjusting the environmental temperature. Therefore, facile sample recovery can be achieved by increasing the temperature to precipitate the polymer-O-GlcNAc protein conjugates from solution. This novel immobilized triarylphosphine reagent enables efficient enrichment and sensitive detection of more than 1700 potential O-GlcNAc proteins from HeLa cell using mass spectrometry, demonstrating its potential as a general strategy for low-abundance target enrichment.
O-GlcNAc proteins:
A0A024R5Q9, A0A0B4J1V8, A0A0B4J203, GAL3B, RBM47, UBA6, ESYT2, SHOT1, NDUF8, VWA8, GTPBA, A5Y5A3, CNOT1, PGP, TBAL3, SMHD1, LR14B, NCF1B, A8K005, B4DLC0, B4DLN1, B4DQP1, B4E259, EIFCL, B7Z2B5, B7Z532, B7Z5W1, DSA2D, D7RF68, E9PL57, E9PLN8, I1VE18, L0R5A1, S35U4, ADAS, KIF2A, PDLI1, AIP, SMAP, NDUS8, PSD12, PSMD9, SPT5H, DFFA, HIP1, EIF3F, IPO5, SAP18, IF2B3, DNM1L, NDUA4, PSDE, BIN1, NOP56, DDX3X, IMA3, TRI38, SCD, DHSD, TXND9, ACPM, UBFD1, AP3D1, DVL2, CHD1, TOR1A, PTGES, ACOT8, NHRF1, TPP1, RPC1, PSA7, MGST3, HAT1, QCR8, HGS, S27A2, HNRDL, BTAF1, SR140, PUR4, NPC1, DEGS1, SCAM2, ARPC2, ARPC3, BET1, RPAC1, PGRC2, NDUA1, RER1, OGT1, PMM2, HMGB3, EIF3H, ARPC5, DHX15, CYB5B, PRP4, SERA, NDUS4, PAPS1, ZW10, SNUT1, RIPK2, PRPF3, PPIH, E41L2, XPOT, DNPH1, PSCA, TSN6, ZN207, NDUB5, BUB1, BUD23, GATC, HTSF1, SGTA, SYNC, PM34, CPSF5, SCO2, ORC5, TIM8A, DHX16, SMCA5, MAST3, OPA1, TBCD4, OGA, HNRPQ, DIAP1, MOT2, IMA7, CTND1, DPM1, Z354A, RT14, PQBP1, TI17B, DKC1, IF2P, EDF1, MCE1, ABCB7, WDR1, BRE1B, ZC11A, CLU, ANR17, COQ9, NDUS7, NIPS2, VPS4B, CISY, PR40A, NDK6, VP26A, MPPB, PSIP1, ECI2, BAF, SF3B1, CSDE1, PRKRA, NPM3, ERAL1, TIPRL, KS6A4, NU155, XRP2, WDHD1, CRTAP, S2512, RNH2A, EIF3G, AP1G2, IDHC, CCNK, SPF27, FLOT1, GLRX3, RL1D1, CLPX, CIAO1, SRP72, LTN1, UFL1, UBXN7, PRP6, PCF11, GLSK, SC31A, RPP14, YIF1A, AGFG2, NDUB6, UBE4B, ELP1, NDUB8, LC7L3, KIF4A, MPZL1, 6PGL, PAPS2, TRI16, ARI2, PSMG1, MPC2, ASML, RPP29, HS74L, AP2A1, TTC4, TM50A, BAG2, EMAL2, BPNT1, NDUBA, ACL6A, COX1, COX2, PNPH, AATM, EGFR, ADA, ATP6, RASN, LDLR, HLAB, LMNA, ALBU, NU1M, NU3M, NU4M, KITH, K2C1, G3P, HLAA, TYSY, RPN2, HMGN1, ISG15, LA, K1C18, DCUP, CDK1, NPM, HEXA, HYEP, H10, SAP, SYEP, HNRPC, YES, SP1, DAF, PFKAM, HEM3, CY1, RU17, ITA5, RS17, RSSA, RM03, SRP19, RU1C, HMOX1, RU2A, PARP1, LKHA4, ALDOC, F86B2, GSTT2, ST1A3, HS71B, RPAC2, ECE2, EFMT4, EFCE2, MYBA, HLAC, ARAF, H14, ODP2, MGST1, ESTD, GTR3, UMPS, ODB2, ACADM, TOP1, TOP2A, G6PD, PYC, C1TC, ADHX, PRPS2, PABP1, ADT1, TPR, KCRB, RINI, K2C5, ACPH, GYS1, ENOB, CD59, MTDC, MIF, CX7A2, CCNB1, CX6B1, PO2F1, SYDC, PLAK, QCR7, ERF3A, EZRI, UCHL3, MCP, PHKG2, RS2, RFA2, CBR1, BGAL, EPCAM, NCPR, AT2A2, UBF1, NDUB7, KAPCA, CAN2, PYRG1, DDX5, LEG3, TCPA, PTN1, RL7, NELFE, PGAM1, RCC1, XRCC1, RPAB1, NDUV2, SPEE, IF2B, BTF3, H2A1D, COX5A, LMNB1, IMDH1, FLNA, ACOHC, ODBB, MUTA, OSBP1, PIMT, FBRL, PUR2, HEMH, TCEA1, AT2B4, CPT2, KTHY, RIR1, EF1B, AT5F1, THIL, CDK2, APC, MCM3, THTM, RS12, TYY1, ITA3, DDX6, DNMT1, U2AF2, HMGB2, SYVC, EF1G, AOFB, RL10, RFA1, APEX1, MAP4, CALX, PSA5, NDUS1, RXRB, AMPL, T2EA, T2EB, IMPA1, 3MG, CO4A5, SPB3, ATPD, PDIA3, 2AAB, CDC27, NMT1, PURA2, AMRP, PUR8, CLIP1, GSTT1, CTR1, AL1B1, RPB2, METK2, RIR2, HNRH3, HNRH1, STIP1, L1CAM, P5CR1, TF2H1, ACSL1, CSTF2, MCM4, MCM5, MCM7, RFC4, RFC2, RL22, T2FA, GDE, SOAT1, BASI, NU214, DEK, K22E, PRS7, HEM6, RL4, LONM, SRP14, NUP62, TALDO, RBMX, COIL, RS19, FEN1, CAP2, TCPZ, NNMT, PSB10, RL13A, PEX19, MDHM, RFC5, RFC3, BUD31, GARS, SYIC, EIF1, LAP2A, LAP2B, MTREX, AK1C3, LPPRC, THIM, MATR3, GPDM, SSRA, SSRB, RANG, PRS6B, EFTS, VDAC2, UBP5, RECQ1, ATRX, IF2M, MP2K3, BRCC3, RL27A, RS10, GNPI1, CAPZB, IF1AX, SYQ, RL29, MAOX, PTSS1, IDHP, PIPNB, DPOD2, TCPG, RM19, INPP, SRP09, SYCC, PSB2, MCM2, ACADV, TMEDA, RBM25, NU153, RBP2, NDUV1, GSK3B, 5NTC, SPS1, GUAA, DNLI3, GDIB, CPT1A, F10A1, DYN2, MAP2, CDK7, CDK9, LRBA, RL14, PAPOA, FXR1, FXR2, RAB7A, RB27A, RT29, SMCA4, SSRD, TPMT, SSDH, DHB4, PSMD7, STA5B, UBP11, DYLT3, CCNH, ROA3, HNRPM, IMA1, NCBP2, MP2K6, KIF11, HXK2, BIEA, PLK1, NUBP1, MAP11, SUCA, MVD1, PGTB2, CATC, CCHL, IST1, TCP4, PMS2, SYYC, UBP14, RD23A, P5CS, IF5, PSMD4, BI1, MFAP1, AFAD, NP1L1, CASP9, ADK, DSRAD, SEC13, HNRH2, SCOT1, EIF3B, ATP5E, IF6, CF298, TMM33, CORO7, SELB, MTPN, YBEY, ARPC4, CD81, SC61B, ROMO1, MYL6, ACTB, IF4A1, PRPS1, PSA6, CDC42, DEST, GMFB, RAB8A, SPCS3, SRP54, RAB5B, UBC12, UBE2K, RAB14, TM258, CSN2, RAP1B, RS3A, VA0D1, NAA20, S61A1, B2MG, COPZ1, UFM1, AP1S1, HNRPK, TIM10, PRS4, RS15A, RS23, RUXE, SMD3, PRS10, ERF1, RS6, RAN, RL23, RS15, RS24, RS25, RS26, RBX1, RL39, RL31, RL10A, RL32, PPIA, TRA2B, AP2B1, DYL1, DYLT1, RL38, SKP1, ACTG, UBC9, CSK2B, UB2L3, EF1A1, CSK21, SERB, IF4G2, RPP38, GTF2I, RAE1L, GSTO1, PRKDC, IN35, RT25, RT35, RT05, RT11, RT15, RT06, RT09, LACTB, RL36A, ARF5, ERH, H33, FOXK1, PRR5, RBM10, TFAM, PIPNA, TF2B, CDK16, NFKB2, FKBP3, HNRPU, U2AF1, SET, GALK2, FABP5, CAP1, HMCS1, IFM3, RL18A, NUCB1, RL6, M2OM, LMNB2, TAP1, TAP2, CEBPZ, TF65, UBXN1, IF4G1, ATP7A, CSTF1, FAK1, SRS11, EF1A2, PUR1, PSME1, GABPA, FMR1, RL18, C1QBP, KHDR1, KLC1, SRSF1, QOR, LG3BP, PPID, SSRP1, CAMP2, NSUN2, RBBP4, NCBP1, HSP7E, BST2, MPPA, WASC5, TBL3, SNF5, STX4, GRSF1, NFIA, SF3A3, TP53B, ILF2, ILF3, PTPRJ, TRAP1, TRAF2, MYO1E, PP1R8, BNIP2, CSTF3, STRN3, RM28, ACACA, CSN1, CAF1B, TADBP, ROA0, AIMP2, PAK2, CBX3, STK3, STX5, SRSF5, SRSF6, TIF1B, PABP4, EIF3I, 2A5G, UB2V1, RM49, DC1I2, ILK, SNTB2, PPIG, TCOF, SF3B2, HAP28, ROCK1, PICAL, SQSTM, PRP4K, HDAC1, DCTN2, SNW1, STIM1, PWP1, MTMR2, CUL2, CUL4A, FCL, RAB31, RAB32, NOG2, TBB2A, BYST, PEBB, HNRPD, IL18, VEZF1, TRI29, TTL12, DPOA2, ICT1, DCTN1, ELOA1, TRI25, FASTK, PDE3A, RBM39, HLTF, MCM6, ITPR1, TRIPC, PUM1, EPN4, SMC1A, RRP1B, PSF1, UBP10, GANAB, LBR, CHD4, NOLC1, NUMA1, PSME4, GAPD1, CND2, SPCS2, EMC2, FL2D, PSMD6, ABRX2, SART3, CND1, EXOS7, U5S1, SYLM, WDR43, ACOX1, EBP, PCM1, NOMO1, PON2, TEBP, NONO, RAB35, RNPS1, PCBP2, ELOC, UBE3C, DHC24, SF3B3, RSU1, SF3B4, SF3A1, SKI2, MARE2, NHRF2, TSN, SF01, CIP4, TRIPB, PCH2, ELAV1, SMAD2, TBCE, VAMP3, ZYX, SEPT7, CCDC6, UAP1, PSMD5, MK14, RM23, DPYL2, SYPL1, RBBP7, IF16, KYNU, CAH9, NDUA9, PCKGM, HCDH, UGPA, MIC60, HNRL2, INF2, PDS5A, TSR1, ALG11, QRIC1, SMU1, LRRF1, RPA43, LSM12, ACSF3, PPCEL, TB10B, MIX23, RHG29, PDCD4, Q53FH6, DHB12, UBP39, LACB2, EIPR1, H90B4, TM41B, HERC4, PP6R3, PREP, PSMG4, RRP12, TOIP1, SYAM, PCID2, S27A3, SAMD9, CYTSB, TDIF2, TTC38, EXOS6, NU188, CE170, SNPC4, CHCH9, DDX59, GPTC4, HECD3, SKT, RM02, UBAP2, LCE1C, ACBD5, EEIG2, RBM26, DCAF8, ATPF1, DDI2, NT5D1, LYRM7, EMC10, RIF1, OBSCN, CE350, AXA81, PR38B, BRE1A, ARHGG, LYPL1, ECM29, MIC13, MAP1S, UTP25, TENS3, INT3, C19L1, CPIN1, SMYD5, PTRD1, CNDH2, MRM1, P4R3A, ACD10, DE10B, SCMC1, TM214, HIBCH, TTI2, DHX57, CDC73, RM14, L2GL2, MET2B, PRP8, TTC27, SKI3, RICTR, LCLT1, WDR82, HACD2, GGYF2, KAT3, UBR3, LARP4, SPT6H, SND1, COX15, DDX46, TM10C, 5MP2, EIF3M, MEPCE, CYFP1, ENOF1, FAKD5, MOB1B, KDM3B, HS2ST, K2C74, TRM1L, ZCCHV, NUP54, MAVS, DHX29, COMD6, HDDC2, LIMS2, I2BP2, PATL1, SETD3, PABP2, MTA70, ZN598, AACS, LUZP1, GP180, CAND1, CMTD1, PRSR2, CARM1, COMD2, DDX42, SPAS2, CONA1, NDUAB, DAAF5, STX12, DLP1, P66A, HORN, RB6I2, FABD, CEP97, CTL2, PHF6, UBR1, CHERP, ANKH1, DJC10, NELFD, MIRO1, RB12B, RAVR1, SPB1, SRRM1, CA174, MICU2, TRM2A, PELP1, ABI1, CCAR2, NDUF2, NUP93, LRC47, GPD1L, TOM5, OGFD1, TTC9C, JAGN1, CPSF7, PAF1, ZN433, F241A, ENAH, CL029, NHLC2, SUMF2, UBAC2, RDH13, UXS1, FA98A, NNRE, LS14A, PAPD5, CNNM3, MCU, FBX22, FAD1, NUP35, Q8NFH9, ABHDB, KNL1, OR1M1, ATLA2, MSPD2, DDX55, NCOA7, THOC2, WDR36, G45IP, CX038, FBX30, S35B2, PUM2, KLDC4, UBA3, KISHA, TBC15, RM30, PNPT1, SSH3, GEMI5, IPO4, ABHD5, ZC3HF, TF3C2, TM263, PTPM1, NU133, CHAC2, THEM6, LEO1, CPLX3, NUDC2, TFB1M, SCFD1, HNRLL, CD003, SEN15, SPRY4, TRUB1, LMO7, ARAP3, PRP31, PALLD, CTBL1, DDB2, DDX1, DHB8, PSMF1, RTF1, RT27, AP3S1, LAR4B, GCN1, PRP16, NU205, TFG, RBP56, DDX17, RAD50, CELF1, OSTF1, ARHG1, UFD1, RENT1, SMRC1, FUBP2, ARHG2, UBP13, USP9X, SCAM4, OSBP2, F162A, RM24, SYAP1, EXOC4, FERM2, PBIP1, RPE, EXOS8, OTUL, CHCH1, PPWD1, COA7, TAM41, F136A, P5CR2, DCPS, FAF2, FXRD1, AP2M1, GCP3, F241B, NMD3, RMD1, MSI2H, SNR40, RM38, RRFM, SIR1, YIPF6, KBP, GNA1, RT24, MOCOS, DAZP1, SAAL1, SIM12, DNJA3, TMA16, PTCD3, OTUB1, TRM61, HMCES, DUS3L, OTUD5, VMP1, DCNL1, SMRD1, TOE1, DDX27, MTNB, ZC21A, PGAM5, FUBP3, SPF45, SUCB2, HPDL, ATPMK, BT3L4, CLP1L, SCYL1, SNX27, YTDC1, ZN830, NSUN5, RBM14, UHRF2, PUS7, TRNT1, TEFM, EFGM, NUDC1, PRPK, PIGS, WRIP1, RANB9, NEB2, IWS1, SIN3A, OSBL9, INT14, ZSC10, RUFY1, MMS19, UHRF1, NIBA2, YMEL1, MYCBP, TBCB, PSB7, CNN2, CDC5L, PSMD1, PFD5, MPH6, TSNAX, TTC1, CSN8, KIF2C, MGLL, HCD2, NP1L4, ACON, TCPH, ANM1, VRK1, CND3, ARP5L, DDX50, RNZ2, GRWD1, MACD1, CMS1, MEP50, RT63, TBA1C, MBB1A, FERM1, CG050, PDD2L, MIC25, UQCC2, UTP23, LXN, NTPCR, TACO1, GCP2, SEN34, ALG1, PSMG3, CSN4, DIDO1, RBM42, MCMBP, AASD1, DCNL5, AN32E, P4K2A, MMTA2, TBB6, PAXX, HNRL1, MTNA, WDR18, THUM3, NTM1A, ASHWN, TM109, GNL3, NUP85, ELOV1, IFT27, SYTM, THIC, FUND2, SF3B5, SFXN3, CARD6, OSB10, B2L13, SRRT, HDHD5, HDAC8, RTF2, EIF2A, PDIP3, RM13, RM04, RM01, MK67I, YTHD1, RT26, NEUL, ABCA2, RM37, UACA, WDR11, CRNL1, TBL1R, UBL5, API5, FTO, UBE2O, XPO4, S39A8, WDR12, YIPF3, DERL2, SKI8, SLIRP, EGLN1, UBA5, MKRN2, T126A, PAIP1, CLPB, LSG1, NAT10, ILKAP, XRN2, NSRP1, CSTFT, 5NT3A, GBRL1, DDX47, RM18, SNAB, ATG5, GAN, DPH5, DHX36, IPYR2, SPNS1, RM46, PININ, GPT, GHITM, NELFA, OSGL1, HEAT1, TFB2M, YTDC2, RPAP3, WDR26, CH033, UBE2Z, ANKZ1, ZWILC, MED20, ARMT1, L2HDH, ELP3, RPA2, KT3K, SYCM, PPCS, NJMU, XPO5, GRPE1, RHOF, POPD2, EMAL4, GLOD4, MCCB, MOV10, SYF1, TM9S3, WDR6, TRXR2, SYSM, RT30, M1IP1, UBE2T, NOP10, OSGEP, MED4, RM40, RTN4, RPR1B, RRAGD, PFD4, EXOS3, ANLN, XPP1, BIRC6, ACSA, DBLOH, DDX21, SAR1A, DPOE4, SIAS, SHLB2, EI2BG, MBNL1, PYRG2, DPOE3, CHRC1, COXM2, ABC3C, PNO1, RM17, HELLS, SYFB, SYIM, ATG3, SMC4, OLA1, CUTC, ECHD1, RBM12, MDN1, DECR2, ABHDA, LIN7C, TXLNG, SPS2L, ABCF3, DD19A, UQCC1, EXD2, DJC11, FANCI, ATD3A, ARL8B, ANM7, FAIM1, RT18A, ARMC1, RPC5, SDA1, RBM28, RBM22, DMAC2, WDR70, PK1IP, RM22, CZIB, BABA1, NDUBB, RM16, NHP2, OCAD1, ADPRS, LYAR, MIC19, THUM1, TRM1, PP4R2, DPP3, BCLF1, FKB11, TMOD3, ELOV5, GTSE1, TECR, ERAP1, ARP10, CISD1, F120A, IF2B1, COQ3, HPBP1, MAT2B, CNOT2, OGFR, CNIH4, RM15, THYN1, KCMF1, TM14C, HM20B, RCC2, RELCH, BCCIP, STK26, FPRP, IBTK, RRBP1, SYLC, SUCB1, DPM3, ATX10, TFP11, SAE1, COMD3, MTRR, ASH2L, DHCR7, EIF3K, SAE2, WNT16, CSN7A, QCR9, ZO2, LRWD1, ABCF2, SYWM, SEC63, SRP68, CHRD1, BAP29, S30BP, PFD2, GPN3, PUF60, NRBP, ENOPH, NDUAC, EI2BD, VATH, XPO7, VPS51, S2513, DCTN4, FBX2, CCNL1, DBR1, FEM1B, DJC12, MRT4, CPSF3, NUDT5, RCOR1, TF3C4, PACN3, ACINU, AGO2, PSME2, PALD, COR1C, AKP8L, ALK, DD19B, PRP19, UBQL1, SNX12, CHIP, PACN2, MAGD2, SNX6, SSRG, FAF1, PPIE, MINP1, SCAF8, TRI33, PHF8, LIMC1, UBP24, SMC3, SCML2, PIN4, PSF2, EIF3L, PLAP, RUVB1, BCS1, SYFA, CNPY2, PKHA6, RRP44, RT07, PDIP2, WBP11, NOP58, SGT1, ACOT9, AAR2, MEMO1, TMX2, LC7L2, DHRS7, RT02, SBDS, TMED5, EXOS1, SF3B6, RM11, RRP15, NOP16, TPRKB, PPIL1, CIA2B, RT16, RT23, PTH2, RTCB, FBX7, NOC2L, RL36, SAMH1, DCAF1, UBP15, TBL2, TELO2, LAS1L, AFG32, ARI1, SAM50, PRC2C, PPME1, AP4S1, YTHD2, CTDP1, SP16H, UTP18, TNPO3, GMPPB, RBM8A, SNX8, FHOD1, SERC, NCOR2, ORNT1, ALG5, RT18B, CLIC4, NFS1, SPCS1, MD1L1, STK24, 5MP1, COMDA, DC1L1, TX264, S4A7, NDUB9, MORC2, S23IP
Species: Homo sapiens
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