[No authors listed]
In eukaryotic cells a molecular chaperone network associates with translating ribosomes, assisting the maturation of emerging nascent polypeptides. Hsp70 is perhaps the major eukaryotic ribosome-associated chaperone and the first reported to bind cotranslationally to nascent chains. However, little is known about the underlying principles and function of this interaction. Here, we use a sensitive and global approach to define the cotranslational substrate specificity of the yeast Hsp70 SSB. We find that SSB binds to a subset of nascent polypeptides whose intrinsic properties and slow translation rates hinder efficient cotranslational folding. The SSB-ribosome cycle and substrate recognition is modulated by its ribosome-bound cochaperone, RAC. Deletion of SSB leads to widespread aggregation of newly synthesized polypeptides. Thus, cotranslationally acting Hsp70 meets the challenge of folding the eukaryotic proteome by stabilizing its longer, more slowly translated, and aggregation-prone nascent polypeptides.
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VAC17, MIC10, PRD1, KAR4, PBN1, APA1, POF1, MGR1, GLK1, SRO9, GFD2, GRX1, LSB5, RNQ1, HBN1, DCC1, GBP2, ILV6, STP22, CDC10, YCP4, CIT2, RVS161, PGK1, RHB1, RIM1, SYP1, RPS14A, RRP43, YCR043C, YCR051W, THR4, CTR86, PWP2, YIH1, TAH1, HCM1, RSA4, SOL2, FUB1, PAT1, AHC2, TRX3, TUP1, CSM1, YCR090C, MSH3, YCR102C, SNZ3, SWP82, RGD2, SEC53, OTU1, ACT1, YPT1, TUB2, MOB2, RPL22B, HAC1, CAF16, FRS2, GAT1, LPD1, BLM10, SEC4, VTC2, SPB4, RPN11, YFR006W, YFH7, IOC3, ATG18, LSB3, HIS2, CDC14, RPL2A, DUG1, BNA6, RMD8, YMR31, PRE4, RET2, RPN12, HXK1, IRC7, GRC3, RIX7, YLL032C, FRA1, ISA1, HSP104, SSA2, HIF1, DPS1, LMO1, ORC3, RTT109, MHT1, YLL058W, YLL053C, COF1, RPL8B, SDH2, DNM1, NOC3, SSK1, NSE1, MEU1, IRC25, SDO1, AAT2, ADE16, RPL15A, RAD5, YLR042C, TRX1, RPS0B, YLR049C, SHM2, REX2, FRS1, RPL22A, BMT6, XYL2, LAM6, RFU1, RPL10, ARP6, GIS3, IOC2, HRT3, ICT1, ERG27, CDC45, REX3, AHP1, CCW12, HOG1, MSL5, YLR118C, YLR126C, DCN1, DIP2, CKI1, PDC5, TIS11, RKM5, RRN5, DPH6, SPE4, SMD3, YLR149C, STM1, ACS2, RNH203, YLR157C-A, MAS1, RPS31, UPS2, DPH5, YLR179C, SAM1, TOS4, EMG1, SKG3, MMR1, UPS1, NMT1, PWP1, PBA1, COQ9, MSS51, SEC13, PNP1, CLB4, CDC123, CPR6, CCC1, CDC42, BNA5, VPS34, GPN3, MAP1, YEF3, GSY2, HSP60, LCB5, YPT6, SEC22, DCS1, MCM5, DBP9, NNT1, YLR287C, COQ11, SEC72, GSP1, ATP14, HRI1, MRPL15, SFH1, RPL38, NMA1, SGD1, RPP0, RPL26A, YLR345W, KAP95, DIC1, ORM2, NIT3, YLR352W, TAL1, ILV5, ATG33, RSC2, ADE13, STE11, RPS22B, ARC18, STP3, CSR1, IKI3, REH1, STE23, AFG2, DUS3, SFP1, DUS4, RPL31B, UTP21, VIP1, YLR419W, URA4, MAG2, CRN1, IMD3, CNA1, TSR2, DIF1, RPS1A, YLR446W, VMA6, RPL6B, SST2, YLR460C, RBG1, POP5, SNC1, CCR4, TPD3, CYS3, FUN14, ERP2, BDH2, BDH1, ECM1, AIM2, AIM1, YAL044W-A, GCV3, PTA1, SSA1, EFB1, TFC3, NUP60, ERP1, SWD1, RFA1, SEN34, ADE1, KIN3, CDC15, WHI4, RRI1, GDH2, YDL211C, NHP2, HEM3, ACK1, TRM8, GGC1, ARF1, RPL35A, UFD2, RBS1, PPH22, VMA1, LYS20, INH1, PHO13, PTP1, SSB1, HO, PCL9, SFA1, NRP1, CDC36, CDC9, DHH1, STE7, CLB3, RPN5, COP1, YDL144C, CCT4, CRD1, BPL1, RPO21, ARF2, RPL35B, RDI1, PPH21, LYS21, RPP1B, YDL129W, CDC48, YDL124W, YFH1, CYK3, NUP84, RRP42, KIN28, PHO2, QRI1, POL3, DUN1, GET3, RPN6, SRP14, RAM1, ASM4, YDL086W, SUB2, RPS16B, RPL13A, MDH3, RXT3, RPL31A, IDP1, UBC9, TSR1, PSA1, LHP1, STP4, SIT4, SIR2, NAT1, PUS9, SLM3, PRP9, ARP2, MPS1, RTK1, GPD1, RPN4, CDC7, NOP1, APC11, RPT2, PTC1, ATP16, RMD1, NTH1, YRB1, MAF1, TRP1, RPL4B, DAS2, FAL1, SES1, RPS11A, VPS54, PST2, ARO3, EHD3, KRS1, RSM10, HEM13, HEM12, TPI1, DET1, UBC5, AIM7, RPS13, PAA1, PPH3, VPS41, PDC2, RRP1, YDR089W, RLI1, UBC13, MSH6, GRX3, BMH2, ARX1, STE5, TRS85, FOB1, ALT2, MRX14, TMA64, TRM1, KIN1, ARO1, MTC5, SAC6, YDR132C, YDR134C, RGP1, PEX7, EKI1, KGD2, CPR1, RPA14, HOM2, ACL4, NBP2, TRM82, CDC37, STB3, SUP35, HMO1, RSM24, UBC1, SDH4, ATC1, UPS3, CCT6, SLY1, RVB1, NUP42, REF2, COQ4, EBS1, YDR210C-C, GCD6, TCP1, AHA1, YDR222W, HTB1, ADK1, PCF11, MRPL7, SEC26, PRP28, TRS23, VHS1, PAM1, SWM1, CIA1, GLO2, PMP3, RRP45, PHM6, SSD1, ATP5, PRO1, CFT1, GIC2, ASP1, UTP4, SKP1, UBX5, YDR341C, SVF1, TRR1, TRP4, BCP1, TFC6, CDC40, YDR365W-A, YPR1, XRS2, DXO1, VPS74, FRQ1, EFT2, RVS167, SAC7, UBA2, RPT3, UTP5, HPT1, URH1, RPB7, ADE8, SYF1, RPL12B, RAD30, RPN9, BNA7, TIF35, NPL3, PPM1, APT2, RPS17B, UTP6, RPS18A, YHP1, GUK1, MRPL28, STP1, RMT2, TLG1, RPL27B, TRS31, YDR476C, SNF1, DIG2, RIB3, SLD5, PKH1, MZM1, SEC20, SAM2, SPG3, SMT3, SDH7, GRX2, EMI2, GRH1, URC2, HLR1, QCR7, APA2, CAB1, HSP31, PAD1, FDC1, YDR541C, SRO77, PKC1, RTG3, BNA4, ROX3, SCS22, MAP2, RPL23A, ILS1, RPS8A, AST1, PRS4, PRX1, SKT5, SHP1, PTC3, YBL055C, TOD6, SEC17, COR1, PRE7, URA7, MRPL16, APL3, YBL036C, POL12, RIB1, HEK2, PET9, RPL19B, NCL1, MCM2, FUS3, RRN6, ALK2, HIR1, SLA1, LDB7, HTB2, NTH2, RER2, DSF2, HHF1, IPP1, YBR016W, KAP104, SCO2, OLA1, YPK3, RPL4A, HMT1, PDX3, SCO1, ATP3, ZTA1, RPS11B, RFS1, YBR053C, PRP6, YBR056W, UBP14, AKL1, ORC2, TRM7, YBR062C, TIP1, YBR071W, HSP26, RDH54, SLM4, SEC18, UBC4, TEC1, RPL19A, AAC3, RFC5, POL30, PBY1, FES1, EXO84, SIF2, VID24, YSA1, LYS2, TEF2, GRS1, MRPL36, PTC4, TPS1, VMA2, OPY1, HSL7, CKS1, BMT2, MAK5, SUP45, ADH5, MRPS9, ARA1, APD1, SLI15, AMN1, CDC28, ARL1, UBS1, TYR1, POP7, SSE2, NPL4, UMP1, SWD3, EHT1, RPS6B, RPS9B, RPL21A, MSI1, PGI1, BEM1, MCM7, LDH1, AME1, NGR1, MET8, PYC2, PDB1, PCS60, TDP1, YBR225W, ARC40, ABD1, YBR238C, ERT1, YBR242W, GPX2, ISW1, RRT2, ENP1, HIS7, ARO4, MRPS5, DUT1, TRS20, YBR255C-A, RIB5, TAE1, SHM1, MIC12, YPT10, TSC10, MRPL37, SDH8, EFM2, HSM3, UBX7, CHK1, DUG2, APM3, CTP1, GUS1, YGL242C, ADE5,7, EMC4, OST5, MTC3, ADH4, HXK2, RTG2, RAI1, SDT1, COG1, NIF3, FRA2, KIP3, SKI8, NCS6, YPT32, MIG2, SIP2, SPT16, CHC1, ARO8, EMP24, DSD1, COX13, CDC55, COX4, YGL185C, GTS1, ATG1, MPT5, XRN1, NUP49, ROK1, SUA5, YGL159W, ARI1, LYS5, ARO2, RPL9A, RRT6, HUL5, SEC27, MRM2, RPL1B, CEG1, RSM23, CWC23, MET13, RPS2, PRP43, CDC20, SNF4, TAF6, NSA1, MLC1, ARC1, YGL101W, SEH1, LSG1, SRM1, VPS45, NUP145, SCY1, YGL082W, MPC1, DBP3, RPL7A, MNP1, NPY1, PYC1, SDS23, TYW3, RPT6, ALG13, RNA15, HEM2, YGL039W, PNC1, AGA2, RPL24A, RPL30, TRP5, ALK1, CKB1, ATE1, KAP122, PUF4, SCL1, LEU1, RPN14, CDH1, ERG26, SWC4, ECT1, NMA2, YGR017W, UGA1, RPS25A, ERV1, RPL26B, KSS1, BUD9, RME1, TFC4, YGR054W, ADE6, SPT4, ART5, SMD1, TWF1, TOM20, MRP13, RPL11B, PIL1, UTP22, DBF2, VAS1, RRP46, ASK10, PCP1, CLB1, CLB6, YGR111W, YGR117C, RPS23A, PPT1, ASN2, UTP8, PHB1, PEX4, PRE9, RPL24B, RSR1, CYS4, PTI1, MTR3, GTR2, ERG1, PBP1, RNR4, TDH3, YPP1, YCH1, ADE3, SER2, TRX2, YGR210C, ZPR1, RPS0A, RSM27, MRPL9, PET54, PHO81, YHB1, MIC26, ENO1, APL6, BUD32, MES1, YGR266W, FOL2, CAB4, YGR283C, ZUO1, NUC1, ECM25, YJL218W, REE1, RPS14B, RPS22A, GON7, YJL181W, RPL17B, RFA3, ERG20, HSP150, CIS3, FAR1, VPS35, DAS1, IDS2, SFH5, TIM17, YAK1, TIF2, GLG2, RPS21B, MRS3, URA2, PBS2, NIT2, ALB1, RPE1, CCT7, GZF3, CHS6, MRPL49, DPB11, ARG3, TRL1, SCP160, PRY1, UTP18, YJL068C, MPM1, DLS1, MRPL8, NUP82, BNA3, YJL055W, PEP8, TDH1, MTR4, RTT101, SNX4, TAD2, BET4, RNR2, APS3, CCT3, VTC4, NOP9, CCT8, CTK2, PRE3, MPP10, APL1, POL31, SUI2, TDH2, MET3, ESS1, TES1, MDE1, BNA1, YJR026W, CPR7, RAV1, PET191, SSC1, TAH11, CYC1, RAD7, YJR056C, NTA1, RPA12, CCT5, ARP3, RFC2, HAM1, LIA1, MOG1, CDC11, MIR1, YJR084W, YJR085C, EMC2, YJR096W, YUH1, AIM25, URA8, SOD1, ADO1, CPA2, YJR111C, RSM7, JHD2, ATP2, RPS5, RSF2, XPT1, TTI2, HOM6, YJR142W, MGM101, RPS4A, BAT2, ADD66, PTK1, YKT6, SDS22, ACP1, DPH2, CNB1, HYM1, SPE1, PRS1, SRY1, URA1, OXP1, YRA2, DOA1, SAC1, TRP3, UBA1, RPL17A, MRPL38, PIR1, GPM1, MCR1, AVT3, RPT1, RPC25, MRP8, SDH3, MRPL31, MYO3, PMU1, PGM1, YPK1, RRN3, VPH2, PRR1, YKL106C-A, AAT1, GFA1, APE1, UTP11, MTC2, CWP2, CWP1, YJU3, CYT2, MDH1, TEF4, VMA5, SMY1, DHR2, MUD2, STB6, NUP100, YNK1, MNR2, MSN4, FBA1, MPE1, NUP120, TMA19, ANR2, PRI2, PHD1, SPC42, NFU1, UGP1, YKL033W-A, IXR1, MAE1, PAN3, URA6, MAK11, SPT23, RAM2, SWD2, HCS1, ATP7, ARC19, MRT4, CAP1, RPL14A, MRP17, MET14, VPS1, PAP1, OSH6, MRPL13, RSC4, YKR011C, YPT52, HEL1, GCN3, BCH2, SPC34, KAE1, UTH1, SHB17, YKR045C, PET10, NAP1, FMP46, MRS4, TRM2, RPS21A, TIF1, UTP30, TFA2, PAM17, CCP1, BET3, YKR070W, DRE2, AIM29, ECM4, MSA2, RPF2, NUP133, HBS1, MRPL20, PXL1, BAS1, SIR1, MED7, HRT1, ALR1, RPL25, MDH2, TRM13, TRM11, GRE2, PEX11, PSF3, RIB4, RRP40, CDC33, RPS19A, RPL18A, PAP2, MDY2, TPT1, SDD3, WRS1, COQ3, ADH1, ATG34, REX4, AVO1, BRX1, THP1, SDH5, NBA1, RIB2, MET22, APM4, PRS5, GPD2, ARG1, YOL057W, PSH1, SPE2, GAL11, GSH2, PEX15, RPS15, PRE6, MIM1, TSR4, DIS3, HTZ1, RCL1, COQ10, RRP6, DNL4, SGT2, TIR2, ROD1, HSP10, SFM1, AHC1, HST3, BUB3, STI1, HIR2, WHI2, RAT1, TMC1, SGT1, CKA2, RPL3, CYT1, MSA1, GYP1, CDC21, VPS21, RKI1, RPS7A, NUP1, RAS1, OST2, INP53, YOR111W, CEX1, AZF1, YOR114W, TRS33, RPO31, RPT5, PFY1, CAT5, IAH1, ADE2, ORT1, VPS17, EFT1, IDH2, RUP1, LSC1, THI80, PNO1, RPB2, ISN1, NFI1, PUP1, SME1, DDP1, GET4, SEY1, RPS28A, GLN4, LCB4, HEM15, SER1, GSP2, MSB1, MCA1, MRM1, HIS3, DED1, RET1, NPT1, STE4, SAS5, RFC1, MCT1, ODC2, RPB8, WTM2, WTM1, MKK1, MGE1, KIN4, RPL33B, DFR1, YOR238W, MET7, PUS7, ESA1, ENV9, CLP1, TUM1, NAT5, HNT3, RPN8, GPN2, RBL2, PAC1, YTM1, RIM20, CAF20, HEM4, FSH3, PLP2, YOR283W, RDL1, RDL2, RPS10A, RRS1, TIM18, MBF1, CPA1, RPL20B, FAA1, LDB19, PRO2, FRT1, VMA4, ALA1, RPA190, PYK2, CIN1, TFB6, VTS1, PDE2, PRT1, PRE10, SCP1, RPS12, MRS6, NDD1, NUD1, GDH1, ATF1, FIT2, FIT3, SUC2, COA1, IMP2', RRD1, RPL40A, PAN6, SSL2, CCT2, TMA108, RPL16A, FKH1, AYR1, RHO3, HIS5, POR2, HPM1, SEC24, YIL108W, PFK26, MOB1, SHQ1, DPH1, BMT5, LYS12, RSM25, UTP25, CAB2, AIR1, THS1, SEC28, SER33, MAM33, RPS24B, RNR3, FIS1, EFM4, ARC15, SNP1, GPP1, RPL34B, MMF1, PCL7, MET30, GVP36, NOT3, CST6, CKA1, CAP2, BCY1, IRR1, RPB3, HIS6, RPL2B, DOT5, FAA3, YIA6, BET1, CFD1, PAN1, EGH1, PRI1, STS1, SQT1, YVH1, DCG1, LYS1, YIR035C, IRC24, HYR1, GTT1, PML39, URA5, SEC65, NUP188, CAC2, VPS9, YML096W, GIM5, PRE8, TUB1, YML082W, DUS1, YML079W, RPL6A, YML131W, ERG13, TUB3, CTK3, BUL2, COQ5, DAK1, POB3, TEM1, RPS1B, SML1, CMP2, IMD4, RSE1, YML040W, CGI121, AMD1, RAD52, TSA1, YOX1, RPS18B, YML6, RPS17A, APT1, UNG1, TRM9, RAD33, ERG6, YAP1, TRM12, GLO1, YPT7, CDC5, MIX17, CLU1, SOK2, FMS1, YMR027W, ARP9, MIH1, MSN2, CCS1, ARA2, NUP116, CSM3, ERB1, YMR051C, BUB2, AAC1, RNA14, UBX4, AVO2, ABF2, IRC21, SDD2, SEC14, NAM7, ADH3, NPL6, AIP1, UTP15, YMR099C, YMR102C, ILV2, MYO5, HFD1, MED11, FOL3, ASC1, ADE17, RPL15B, EPO1, ECM16, YMR130W, RRB1, CIN4, RIM11, RPL13B, RPS16A, YMR144W, TIF34, YIM1, NUP53, RIM13, AIM36, MRPS8, YMR160W, ALD2, YMR178W, SSO2, ADD37, HSC82, GCV2, SPG5, RPL36A, YMR196W, CLN1, TOM40, INP1, PFK2, ERG12, YMR210W, EFR3, SKY1, GUA1, ERG8, FSH2, YMR226C, RRP5, RPS10B, PEP5, TRI1, RNA1, TAF9, BCH1, RPL20A, FAA4, GAD1, ROY1, TPS3, YMR265C, PRM15, NGL2, ABZ2, HAS1, TDA1, YMR295C, ADE4, UBP15, SCW10, YMR310C, ELP6, PRE5, YMR315W, ADH6, YMR321C, SNZ2, RPD3, PFS2, PHA2, ATP11, EMW1, RFA2, ZIM17, STB1, MCK1, YPT11, RPS19B, RPL18B, TOS6, CLA4, PUS4, RFC3, PCL1, CAF40, MRPL10, HCH1, CAF120, MET2, PDR17, YIF1, POL2, ORC5, FOL1, TEX1, NRD1, RPA49, YNL247W, VPS75, SUI1, SLA2, ZWF1, NAR1, LAP3, YNL234W, CSL4, URE2, SSU72, POP1, ADE12, MGS1, SSB2, RIO2, YNL200C, WHI3, YNL193W, DUG3, SRP1, NPR1, IPI3, YNL181W, RPS3, MRPL22, FMP41, YNL165W, RIA1, RPL42A, CBK1, CUZ1, GIM3, INN1, RPC31, MFA2, AAH1, SRV2, FPR1, YNL134C, KRE33, NRK1, NMA111, NCS2, DMA2, DBP2, CYB5, YNL108C, YAF9, INP52, POL1, RPS7B, RHO2, END3, PMS1, NIS1, IMP4, MLF3, RNH201, RPL16B, FKH2, RPL9B, YDJ1, GCD10, OCA2, POR1, YNL054W-A, MSG5, COX5A, SFB2, LAP2, YIP3, IDH1, YNL035C, CRZ1, SAM50, FAP1, RCM1, ARK1, YNL010W, SIS1, LST8, DOM34, RPC34, ATG3, NRM1, URK1, SMM1, ACC1, RCF2, ATP23, SNF12, SEC12, BUD17, YNR029C, PPG1, ABZ1, SOL1, ARC35, MRPS12, RSM19, DBP6, COQ2, MVD1, TRM112, LYS9, POP2, NOG2, AIF1, CLN2, VIK1, YAH1, RPL36B, YPL247C, YPL245W, SRP68, CIN2, HSP82, RVB2, SAM4, ACM1, DIM1, KEL3, FUM1, APM1, NEW1, YPL225W, RPL1A, SAR1, CBP3, THI6, PUS1, SRP72, RKM1, PGC1, TPK2, AFT2, RPL7B, OXR1, MRN1, RTC6, RTT10, TCO89, PPQ1, CBC2, CUP9, MRPL40, DAP1, MEX67, BEM4, CDC60, TGS1, KIP2, RRD2, PRP46, YPL150W, KES1, RPL33A, MKK2, UME1, ODC1, COX11, RPL5, TAF14, TBF1, HHO1, NAN1, KAP120, TFB2, VPS30, DBP1, MRP51, IDI1, YPL113C, CAR1, GDE1, SSE1, ELP4, ATG21, INA17, NOG1, GLR1, RPS6A, ELP3, BRO1, RPS9A, RPL21B, ATP4, MUK1, BTS1, YPL068C, RGL1, VPS28, ALD6, GRX5, CAM1, SSN3, MRX11, YPL039W, SVL3, PHO85, TRM44, ERG10, MET12, VTC3, SWI1, HST2, RET3, LSP1, ULA1, SNF8, ICL2, SUT2, RPA135, YPR011C, YPR015C, TIF6, MCM4, APL4, NTO1, SRO7, HTS1, ARP7, GLN1, VMA13, TIP41, TIF5, TAH18, TFB4, YMC1, ARO7, YPR063C, ISA2, SPE3, TKL1, TEF1, DIB1, SUA7, YPR098C, RPL11A, PRE2, ISR1, RPN7, RPC40, DBF20, PIS1, MRI1, CLB2, AXL1, YPR127W, ANT1, SCD6, NAT3, CTF4, RRP9, NOC4, ASN1, YPR147C, GPH1, SGV1, ORC4, TIF3, RHO1, YPR172W, DPB2, BET2, AOS1, SEC23, DPM1, MLC2, QCR2, ARR1, EFM1, RPL8A, GOS1, RPS20, PRS3, STE20, HSE1, RPL14B, OSH7, SOD2, RPL27A, VPS29, ARD1, YSC84, ARG4, DED81, THR1, YHI9, SLT2, RRM3, PIH1, DOG2, DOG1, INM1, AAP1, FSH1, MED6, PAN5, SSZ1, DYS1, RRP4, TRM5, ERG7, QNS1, PTC7, KSP1, SAM35, NAM8, GAR1, SFB3, GEP4, SBE22, GRE3, TRR2, CDC12, GGA2, CTM1, UBA4, YHR112C, APE4, BZZ1, DMA1, SET1, LSM12, CIA2, FUR1, YHR131C, ARO9, RPL42B, DSE2, DCD1, MRPL6, YAP1801, MPC2, SOL3, THP2, DBP8, NMD3, ATG7, OYE2, GND1, IKI1, ERG9, CTF8, EGD2, AIM18, AIM46, RPN10, PPX1, RPS4B, SCH9, SKN7, BAT1, IMD2, DLD3, HPA3, PRB1, PCM1, HAT2, POL5, RPL12A, VMA8, RML2, GLY1, UTR4, RAD23, HYP2, MCM3, BUD16, RIP1, EDC3, GCN4, IRC22, PMI40, FMP52, NUG1, TMA20, NTF2, YER010C, TIR1, PRE1, PRP22, ISC1, RPN3, PRO3, GCD11, GAL83, MIG3, YPT31, FIR1, ARB1, MXR1, SAH1, TPA1, HOM3, HIS1, RPL34A, GPP2, VHR2, MOT2, RNR1, VTC1, RPS24A, PTP3, SBH1, DOT6, PTC2, TRP2, MET6, AIM11, PUP3, RAD51, SHC1, PRS2, RPS8B, NUP157, GLE2, BOI2, RPL23B, SCS2, GLO3, DSE1, RSP5, VFA1, YER134C, GDI1, YER137C-A, DDI1, LSM5, SPT15, YER152C, OXA1, BEM2, YER156C, YER158C, GCG1, PAB1, BCK2, CCA1, RPH1, RAD3, GRX4, BMH1, PDA1, FMP10, FAU1, YER186C
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