Gene Symbol: YRA1
Description: RNA-binding protein YRA1
Alias: SHE11, RNA-binding protein YRA1
Species: Saccharomyces cerevisiae S288c
Products:     YRA1

Top Publications

  1. Strasser K, Masuda S, Mason P, Pfannstiel J, Oppizzi M, Rodriguez Navarro S, et al. TREX is a conserved complex coupling transcription with messenger RNA export. Nature. 2002;417:304-8 pubmed
    The essential yeast proteins Yra1 and Sub2 are messenger RNA export factors that have conserved counterparts in metazoans, designated Aly and UAP56, respectively...
  2. Mackellar A, Greenleaf A. Cotranscriptional association of mRNA export factor Yra1 with C-terminal domain of RNA polymerase II. J Biol Chem. 2011;286:36385-95 pubmed publisher
    ..In Saccharomyces cerevisiae, the mRNA export protein Yra1 (ALY/RNA export factor in metazoa) cotranscriptionally associates with mRNA and delivers it to the nuclear pore ..
  3. Faza M, Kemmler S, Jimeno S, González Aguilera C, Aguilera A, Hurt E, et al. Sem1 is a functional component of the nuclear pore complex-associated messenger RNA export machinery. J Cell Biol. 2009;184:833-46 pubmed publisher
    ..Thus, Sem1 is a versatile protein that might stabilize multiple protein complexes involved in diverse pathways. ..
  4. González Aguilera C, Tous C, Gómez González B, Huertas P, Luna R, Aguilera A. The THP1-SAC3-SUS1-CDC31 complex works in transcription elongation-mRNA export preventing RNA-mediated genome instability. Mol Biol Cell. 2008;19:4310-8 pubmed publisher
  5. Gaillard H, Wellinger R, Aguilera A. A new connection of mRNP biogenesis and export with transcription-coupled repair. Nucleic Acids Res. 2007;35:3893-906 pubmed
    ..In this study we show by molecular DNA repair analysis, that Sub2-Yra1 and Thp1-Sac3, two main mRNA export complexes, are required for efficient TCR in yeast...
  6. Zenklusen D, Vinciguerra P, Strahm Y, Stutz F. The yeast hnRNP-Like proteins Yra1p and Yra2p participate in mRNA export through interaction with Mex67p. Mol Cell Biol. 2001;21:4219-32 pubmed
    ..describe a second nonessential Saccharomyces cerevisiae family member, called Yra2p, which is able to complement a YRA1 deletion when overexpressed...
  7. Iglesias N, Tutucci E, Gwizdek C, Vinciguerra P, Von Dach E, Corbett A, et al. Ubiquitin-mediated mRNP dynamics and surveillance prior to budding yeast mRNA export. Genes Dev. 2010;24:1927-38 pubmed publisher
    The evolutionarily conserved mRNA export receptor Mex67/NXF1 associates with mRNAs through its adaptor, Yra1/REF, allowing mRNA ribonucleoprotein (mRNP) exit through nuclear pores...
  8. Strasser K, Hurt E. Yra1p, a conserved nuclear RNA-binding protein, interacts directly with Mex67p and is required for mRNA export. EMBO J. 2000;19:410-20 pubmed
    ..Mutants of YRA1 are impaired in nuclear poly(A)+ RNA export at restrictive growth conditions...
  9. Stutz F, Bachi A, Doerks T, Braun I, Seraphin B, Wilm M, et al. REF, an evolutionary conserved family of hnRNP-like proteins, interacts with TAP/Mex67p and participates in mRNA nuclear export. RNA. 2000;6:638-50 pubmed
    ..Conditional expression in yeast cells implicates Yra1 p in the export of cellular mRNAs...

More Information


  1. Strasser K, Hurt E. Splicing factor Sub2p is required for nuclear mRNA export through its interaction with Yra1p. Nature. 2001;413:648-52 pubmed
    ..Here, we report a genetic interaction between YRA1 and SUB2, the gene for a DEAD box helicase involved in splicing...
  2. Fischer T, Strasser K, Rácz A, Rodriguez Navarro S, Oppizzi M, Ihrig P, et al. The mRNA export machinery requires the novel Sac3p-Thp1p complex to dock at the nucleoplasmic entrance of the nuclear pores. EMBO J. 2002;21:5843-52 pubmed
    ..Taken together, our data suggest that the novel Sac3p-Thp1p complex functions by docking the mRNP to specific nucleoporins at the nuclear entrance of the NPC. ..
  3. Zenklusen D, Vinciguerra P, Wyss J, Stutz F. Stable mRNP formation and export require cotranscriptional recruitment of the mRNA export factors Yra1p and Sub2p by Hpr1p. Mol Cell Biol. 2002;22:8241-53 pubmed
    ..We also identified functional interactions between Yra1p and the exosome components Rrp45p and Rrp6p. We show that yra1, sub2, and Deltahpr1 mutants all present defects in mRNA accumulation and that deletion of RRP6 in yra1 mutants ..
  4. Johnson S, Kim H, Erickson B, Bentley D. The export factor Yra1 modulates mRNA 3' end processing. Nat Struct Mol Biol. 2011;18:1164-71 pubmed publisher
    The Saccharomyces cerevisiae mRNA export adaptor Yra1 binds the Pcf11 subunit of cleavage-polyadenylation factor CF1A that links export to 3' end formation...
  5. Vinciguerra P, Iglesias N, Camblong J, Zenklusen D, Stutz F. Perinuclear Mlp proteins downregulate gene expression in response to a defect in mRNA export. EMBO J. 2005;24:813-23 pubmed
    ..The mRNA export adaptor Yra1p/REF contributes to nascent mRNP assembly and recruitment of the export receptor Mex67p. yra1 mutants exhibit mRNA export defects and a decrease in LacZ reporter and certain endogenous transcripts...
  6. Dong S, Li C, Zenklusen D, Singer R, Jacobson A, He F. YRA1 autoregulation requires nuclear export and cytoplasmic Edc3p-mediated degradation of its pre-mRNA. Mol Cell. 2007;25:559-73 pubmed
    ..We show that YRA1 exon 1 sequences in cis and Yra1p in trans inhibit YRA1 pre-mRNA splicing and commit the pre-mRNA to nuclear export...
  7. Johnson S, Cubberley G, Bentley D. Cotranscriptional recruitment of the mRNA export factor Yra1 by direct interaction with the 3' end processing factor Pcf11. Mol Cell. 2009;33:215-26 pubmed publisher
    We investigated recruitment of the yeast mRNA export factor Yra1 to the transcription elongation complex (TEC). Previously, the Sub2 helicase subunit of TREX was proposed to recruit Yra1...
  8. Swaminathan S, Kile A, MacDonald E, Koepp D. Yra1 is required for S phase entry and affects Dia2 binding to replication origins. Mol Cell Biol. 2007;27:4674-84 pubmed
    ..Using an affinity approach, we identified Yra1, a transcription-coupled mRNA export protein, as a Dia2 interaction partner...
  9. Ren Y, Schmiege P, Blobel G. Structural and biochemical analyses of the DEAD-box ATPase Sub2 in association with THO or Yra1. elife. 2017;6: pubmed publisher
    ..In yeast, one of these reactions involves loading of the mRNA-binding protein Yra1 by the DEAD-box ATPase Sub2 as assisted by the hetero-pentameric THO complex...
  10. Gomar Alba M, del Olmo M. Hot1 factor recruits co-activator Sub1 and elongation complex Spt4/5 to osmostress genes. Biochem J. 2016;473:3065-79 pubmed publisher
    ..Instead, other data presented herein indicate a key function of the Hot1 transcription factor in the recruitment of these proteins to the promoter or the 5'-coding region of the genes under its control. ..
  11. Salvi J, Chan J, Szafranski K, Liu T, Wu J, Olsen J, et al. Roles for Pbp1 and caloric restriction in genome and lifespan maintenance via suppression of RNA-DNA hybrids. Dev Cell. 2014;30:177-91 pubmed publisher
    ..Thus, we find roles for Pbp1 in genome maintenance and reveal that caloric restriction counteracts Pbp1 deficiencies by engaging RNaseH and Pif1...
  12. Del Rosario B, Pemberton L. Nap1 links transcription elongation, chromatin assembly, and messenger RNP complex biogenesis. Mol Cell Biol. 2008;28:2113-24 pubmed publisher
    ..One factor investigated is an essential mRNA export and TREX complex component, Yra1. Nap1 interacts directly with Yra1 and genetically with other TREX complex components and the mRNA export factor ..
  13. Blair L, Liu Z, Labitigan R, Wu L, Zheng D, Xia Z, et al. KDM5 lysine demethylases are involved in maintenance of 3'UTR length. Sci Adv. 2016;2:e1501662 pubmed publisher
    ..In contrast, both KDM5A and KDM5B are involved in the lengthening of DICER1. Our findings suggest both a novel role for this family of demethylases and a novel targetable mechanism for 3'UTR processing. ..
  14. Ma W, Paudel B, Xing Z, Sabath I, Rueda D, Tran E. Recruitment, Duplex Unwinding and Protein-Mediated Inhibition of the Dead-Box RNA Helicase Dbp2 at Actively Transcribed Chromatin. J Mol Biol. 2016;428:1091-1106 pubmed publisher
    ..cerevisiae is required to promote efficient assembly of the co-transcriptionally associated mRNA-binding proteins Yra1, Nab2, and Mex67 onto poly(A)(+)RNA...
  15. Ma W, Cloutier S, Tran E. The DEAD-box protein Dbp2 functions with the RNA-binding protein Yra1 to promote mRNP assembly. J Mol Biol. 2013;425:3824-38 pubmed publisher
    ..We now provide evidence that Dbp2 interacts genetically and physically with the mRNA export factor Yra1. In addition, we find that Dbp2 is required for in vivo assembly of mRNA-binding proteins Yra1, Nab2, and Mex67 ..
  16. Felberbaum R, Wilson N, Cheng D, Peng J, Hochstrasser M. Desumoylation of the endoplasmic reticulum membrane VAP family protein Scs2 by Ulp1 and SUMO regulation of the inositol synthesis pathway. Mol Cell Biol. 2012;32:64-75 pubmed publisher
    ..These results provide the first evidence of cross-regulation between the SUMO and inositol pathways, including the sumoylation of an ER membrane protein central to phospholipid synthesis and phosphoinositide signaling...
  17. Palancade B, Zuccolo M, Loeillet S, Nicolas A, Doye V. Pml39, a novel protein of the nuclear periphery required for nuclear retention of improper messenger ribonucleoparticles. Mol Biol Cell. 2005;16:5258-68 pubmed
    ..Together, these data indicate that Pml39 is an upstream effector of the Mlps, involved in the retention of improper mRNPs in the nucleus before their export. ..
  18. Kerr S, Azzouz N, Fuchs S, Collart M, Strahl B, Corbett A, et al. The Ccr4-Not complex interacts with the mRNA export machinery. PLoS ONE. 2011;6:e18302 pubmed publisher
    ..These results shed further insight into the biological functions of Ccr4-Not and suggest that this complex is involved in all aspects of mRNA biogenesis, from the regulation of transcription to mRNA export and turnover. ..
  19. NEAL S, Mak R, Bennett E, HAMPTON R. A Cdc48 "Retrochaperone" Function Is Required for the Solubility of Retrotranslocated, Integral Membrane Endoplasmic Reticulum-associated Degradation (ERAD-M) Substrates. J Biol Chem. 2017;292:3112-3128 pubmed publisher
    ..Thus, in addition to the long known ATP-dependent extraction of ERAD substrates during retrotranslocation, the Cdc48 complex is generally and critically needed for the solubility of retrotranslocated ERAD-M intermediates. ..
  20. Ellisdon A, Dimitrova L, Hurt E, Stewart M. Structural basis for the assembly and nucleic acid binding of the TREX-2 transcription-export complex. Nat Struct Mol Biol. 2012;19:328-36 pubmed publisher
  21. Gavaldá S, Santos Pereira J, García Rubio M, Luna R, Aguilera A. Excess of Yra1 RNA-Binding Factor Causes Transcription-Dependent Genome Instability, Replication Impairment and Telomere Shortening. PLoS Genet. 2016;12:e1005966 pubmed publisher
    b>Yra1 is an essential nuclear factor of the evolutionarily conserved family of hnRNP-like export factors that when overexpressed impairs mRNA export and cell growth...
  22. Preker P, Guthrie C. Autoregulation of the mRNA export factor Yra1p requires inefficient splicing of its pre-mRNA. RNA. 2006;12:994-1006 pubmed
    Yra1p is an essential RNA-binding protein that couples transcription to export. The YRA1 gene is one of only approximately 5% of genes that undergo splicing in budding yeast, and its intron is unusual in several respects, including its ..
  23. Wang Q, He J, Lynn B, Rymond B. Interactions of the yeast SF3b splicing factor. Mol Cell Biol. 2005;25:10745-54 pubmed
    ..These data establish the concordance of yeast and mammalian SF3b complexes, implicate accessory splicing factors in U2 snRNP function, and support SF3b contribution from early pre-mRNP recognition to late steps in splicing. ..
  24. Nedea E, Nalbant D, Xia D, Theoharis N, Suter B, Richardson C, et al. The Glc7 phosphatase subunit of the cleavage and polyadenylation factor is essential for transcription termination on snoRNA genes. Mol Cell. 2008;29:577-87 pubmed publisher
    ..Swd2 is also a subunit of the Set1c histone H3K4 methyltransferase complex and is required for its stability and optimal methyltransferase activity. ..
  25. Rollenhagen C, Hodge C, Cole C. Following temperature stress, export of heat shock mRNA occurs efficiently in cells with mutations in genes normally important for mRNA export. Eukaryot Cell. 2007;6:505-13 pubmed
    ..Heat shock mRNA was exported efficiently in temperature-sensitive npl3, yra1, and npl3 yra1 mutant strains. Nevertheless, Yra1p was recruited to heat shock mRNA, as were Nab2p and Npl3p...
  26. Zander G, Hackmann A, Bender L, Becker D, Lingner T, Salinas G, et al. mRNA quality control is bypassed for immediate export of stress-responsive transcripts. Nature. 2016;: pubmed publisher
    ..Thus, at the cost of error-free mRNA formation, heat-shock mRNAs are exported and translated without delay, allowing cells to survive extreme situations. ..
  27. Pascual García P, Govind C, Queralt E, Cuenca Bono B, Llopis A, Chávez S, et al. Sus1 is recruited to coding regions and functions during transcription elongation in association with SAGA and TREX2. Genes Dev. 2008;22:2811-22 pubmed publisher
    ..In addition, Sus1 copurifies with the essential mRNA export factors Yra1 and Mex67, which bind to the mRNA cotranscriptionally...
  28. Röther S, Clausing E, Kieser A, Strasser K. Swt1, a novel yeast protein, functions in transcription. J Biol Chem. 2006;281:36518-25 pubmed
    ..Importantly, deletion of Swt1 leads to decreased transcription. Taken together, these data suggest that Swt1 functions in gene expression in conjunction with the TREX complex. ..
  29. Brockmann C, Soucek S, Kuhlmann S, Mills Lujan K, Kelly S, Yang J, et al. Structural basis for polyadenosine-RNA binding by Nab2 Zn fingers and its function in mRNA nuclear export. Structure. 2012;20:1007-18 pubmed publisher
    ..is more closely linked to hyperadenylation and suppression of mutant alleles of the nuclear RNA export adaptor, Yra1, than to affinity for polyadenosine-RNA...
  30. Vitaliano Prunier A, Babour A, Hérissant L, Apponi L, Margaritis T, Holstege F, et al. H2B ubiquitylation controls the formation of export-competent mRNP. Mol Cell. 2012;45:132-9 pubmed publisher
    ..the recruitment of the different Mex67 adaptors to mRNPs, the effect of Swd2 ubiquitylation is restricted to Yra1 and Nab2, which, in turn, controls poly(A) tail length...
  31. Meinel D, Burkert Kautzsch C, Kieser A, O Duibhir E, Siebert M, Mayer A, et al. Recruitment of TREX to the transcription machinery by its direct binding to the phospho-CTD of RNA polymerase II. PLoS Genet. 2013;9:e1003914 pubmed publisher
    ..Genome-wide analyses show that the occupancy of THO and the TREX components Sub2 and Yra1 increases from the 5' to the 3' end of the gene in accordance with the CTD S2 phosphorylation pattern...
  32. Erce M, Abeygunawardena D, Low J, Hart Smith G, Wilkins M. Interactions affected by arginine methylation in the yeast protein-protein interaction network. Mol Cell Proteomics. 2013;12:3184-98 pubmed publisher
    ..Interactions between the yeast hub protein Npl3 and yeast proteins Air2, Ded1, Gbp2, Snp1, and Yra1 were first validated in the absence of methylation...
  33. Estruch F, Hodge C, Gómez Navarro N, Peiró Chova L, Heath C, Cole C. Insights into mRNP biogenesis provided by new genetic interactions among export and transcription factors. BMC Genet. 2012;13:80 pubmed publisher
  34. Dong S, Jacobson A, He F. Degradation of YRA1 Pre-mRNA in the cytoplasm requires translational repression, multiple modular intronic elements, Edc3p, and Mex67p. PLoS Biol. 2010;8:e1000360 pubmed publisher
    ..When shunted to the cytoplasm by autoregulated nuclear export, the intron-containing yeast YRA1 pre-mRNA evades NMD and is targeted by a cytoplasmic decay pathway mediated by the decapping activator Edc3p...
  35. Wang Q, Rymond B. Rds3p is required for stable U2 snRNP recruitment to the splicing apparatus. Mol Cell Biol. 2003;23:7339-49 pubmed
    ..Together these data establish Rds3p as an essential component of the U2 snRNP SF3b complex and suggest a new link between the nuclear processes of pre-mRNA splicing and RNA export. ..
  36. Fasken M, Stewart M, Corbett A. Functional significance of the interaction between the mRNA-binding protein, Nab2, and the nuclear pore-associated protein, Mlp1, in mRNA export. J Biol Chem. 2008;283:27130-43 pubmed publisher
    ..RNA and that this nab2 F73D mutant genetically interacts with alleles of two essential mRNA export genes, MEX67 and YRA1. These data provide in vivo evidence for a model of mRNA export in which Nab2 is important for targeting mRNAs to ..