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::General Information
ID U0017
Creation Date 01 September 2005
Updating Date 28 September 2009
Standard Name UNR binding site
UTRSite Pattern
(RRRRRAAGUAR|RRRRRRRRAACRRR)
::Taxonomy
UTR Region All
Taxon Range Metazoa
::Description
Description Upstream of N-ras (UNR) is a cytoplasmic protein containing five cold shock domains (CSD) (Jacquemin- Sablon et al., 1994). Proteins of the CSD family perform a variety of functions and are thought to behave as RNA chaperones, promoting a linear conformation of the mRNA. In humans, UNR has been implicated in the destabilization of c-fos mRNA (Chang et al., 2004) and the activation of translation driven by the IRESs of a number of transcripts, including c-myc (Evans et al., 2003), rhinovirus (Hunt et al., 1999), poliovirus (Boussadia et al., 2003), PITSLRE protein kinase (Tinton et al., 2005), and the pro-apoptotic factor Apaf-1 (Mitchell et al., 2003).

The CSD is a five-stranded β-barrel fold with conserved RNP1 and RNP2 motifs that mediates binding to single stranded nucleic acids. Accordingly, UNR binds to single stranded RNA (Jacquemin-Sablon et al., 1994). SELEX experiments indicate that UNR binds to motifs AAGUA or AACG/A downstream of a 5-8 nt purine stretch (see Table 1) with a Kd of 10-25 nM (Triquineaux et al., 1999). Single base changes Pu-Py have limited effect on UNR-RNA recognition, and 3' purines seem to be dispensable for efficient binding.

Two isoforms of UNR are thought to arise by alternative skipping of exon 5 (Boussadia et al., 1993). In addition, three alternative polyadenylation sites have been described in the UNR gene (Jeffers et al., 1990).

UNR has been described to interact with PABP (Chang et al., 2004) and unrip (Hunt et al., 1999).

Table 1. mRNAs that interact with UNR
mRNA
DB ID
Location
Reference
SELEX
n.a.
n.a.
Triquineaux et al., 1999
PITSLRE kinase
NM_033537
745-1125
Tinton et al., 2005
Apaf-1
NM_001160
345-518
Mitchell et al., 2003
c-fos
NM_005252
533-640
Chang et al., 2004
HRV-2
K02121
10-611
Hunt et al., 1999
c-myc
NM_002467
1-524
Mitchell et al., 2001

::Features
Feature Key UNR-bs
::Transcript(s) / Gene(s)
UTR(s)
Description Species Link Ref.
Apoptotic protease activating factor (Apaf-1) 5'UTR mRNA Homo sapiens UTRef: 5HSAR027754 [U1]
c-myc, 5'UTR mRNA. Homo sapiens UTRef: 5HSAR031001 [U2]
cell division cycle 2-like 2 (PITSLRE proteins), 5'UTR mRNA. Homo sapiens UTRef: BR044502 [U3]
Transcript(s)
Description Species Link Ref.
Apoptotic protease activating factor (Apaf-1) mRNA Homo sapiens RefSeq: NM_001160 [T1]
c-myc, mRNA Homo sapiens RefSeq: NM_002467 [T2]
cell division cycle 2-like 2 (PITSLRE proteins), mRNA. Homo sapiens RefSeq: NM_033537 [T3]
c-fos, mRNA Homo sapiens RefSeq: NM_005252 [T4]
Gene(s)
Description Species Link Ref.
PITSLRE serine/threonine-protein kinase CDC2L2 Homo sapiens ENSEMBL: ENSG00000008128
Myc proto-oncogene protein (c-myc) Homo sapiens ENSEMBL: ENSG00000136997
Apoptotic protease activating factor 1 (Apaf-1) Homo sapiens ENSEMBL: ENSG00000120868
proto-oncogene c-fos Homo sapiens ENSEMBL: ENSG00000170345
::Protein(s)
Binding Protein(s)
Description Species Link Ref.
Cold shock domain protein E1 (UNR protein) (N-ras upstream gene protein) Homo sapiens UniProt: O75534 [B1][B2][B3][B4][B5][B6][B7][B8]
Interactor(s) of Binding Protein(s)
Description Species Link Ref.
Polyadenylate-binding protein 1 Homo sapiens UniProt: P11940 [P1]
Serine-threonine kinase receptor-associated protein (UNR interacting protein) Homo sapiens UniProt: Q9Y3F4 [P2]
::References
ID [1]
Authors Boussadia O, Jacquemin-Sablon H, Dautry F.
Title Exon skipping in the expression of the gene immediately upstream of N-ras (unr/NRU).
Citation Biochim Biophys Acta. 1993 Feb 20;1172(1-2):64-72.
Year 1993
ID [2]
Authors Boussadia O, Niepmann M, Creancier L, Prats AC, Dautry F, Jacquemin-Sablon H.
Title Unr is required in vivo for efficient initiation of translation from the internal ribosome entry sites of both rhinovirus and poliovirus.
Citation J Virol. 2003 Mar;77(6):3353-9.
Year 2003
ID [3]
Authors Chang TC, Yamashita A, Chen CY, Yamashita Y, Zhu W, Durdan S, Kahvejian A, Sonenberg N, Shyu AB.
Title UNR, a new partner of poly(A)-binding protein, plays a key role in translationally coupled mRNA turnover mediated by the c-fos major coding-region determinant.
Citation Genes Dev. 2004 Aug 15;18(16):2010-23.
Year 2004
ID [4]
Authors Evans JR, Mitchell SA, Spriggs KA, Ostrowski J, Bomsztyk K, Ostarek D, Willis AE.
Title Members of the poly (rC) binding protein family stimulate the activity of the c-myc internal ribosome entry segment in vitro and in vivo.
Citation Oncogene. 2003 Sep 11;22(39):8012-20.
Year 2003
ID [5]
Authors Hunt SL, Hsuan JJ, Totty N, Jackson RJ.
Title Unr, a cellular cytoplasmic RNA-binding protein with five cold-shock domains, is required for internal initiation of translation of human rhinovirus RNA.
Citation Genes Dev. 1999 Feb 15;13(4):437-48.
Year 1999
ID [6]
Authors Jacquemin-Sablon H, Triqueneaux G, Deschamps S, le Maire M, Doniger J, Dautry F.
Title Nucleic acid binding and intracellular localization of unr, a protein with five cold shock domains.
Citation Nucleic Acids Res. 1994 Jul 11;22(13):2643-50.
Year 1994
ID [7]
Authors Jeffers M, Paciucci R, Pellicer A.
Title Characterization of unr; a gene closely linked to N-ras.
Citation Nucleic Acids Res. 1990 Aug 25;18(16):4891-9.
Year 1990
ID [8]
Authors Mitchell SA, Brown EC, Coldwell MJ, Jackson RJ, Willis AE.
Title Protein factor requirements of the Apaf-1 internal ribosome entry segment: roles of polypyrimidine tract binding protein and upstream of N-ras.
Citation Mol Cell Biol. 2001 May;21(10):3364-74.
Year 2001
ID [9]
Authors Mitchell SA, Spriggs KA, Coldwell MJ, Jackson RJ, Willis AE.
Title The Apaf-1 internal ribosome entry segment attains the correct structural conformation for function via interactions with PTB and unr.
Citation Mol Cell. 2003 Mar;11(3):757-71.
Year 2003
ID [10]
Authors Tinton SA, Schepens B, Bruynooghe Y, Beyaert R, Cornelis S.
Title Regulation of the cell-cycle-dependent internal ribosome entry site of the PITSLRE protein kinase: roles of Unr (upstream of N-ras) protein and phosphorylated translation initiation factor eIF-2alpha.
Citation Biochem J. 2005 Jan 1;385(Pt 1):155-63.
Year 2005
ID [11]
Authors Triqueneaux G, Velten M, Franzon P, Dautry F, Jacquemin-Sablon H.
Title RNA binding specificity of Unr, a protein with five cold shock domains.
Citation Nucleic Acids Res. 1999 Apr 15;27(8):1926-34.
Year 1999
ID [U1]
Authors Mitchell SA, Spriggs KA, Coldwell MJ, Jackson RJ, Willis AE.
Title The Apaf-1 internal ribosome entry segment attains the correct structural conformation for function via interactions with PTB and unr.
Citation Mol Cell. 2003 Mar;11(3):757-71.
Year 2003
ID [U2]
Authors Mitchell SA, Brown EC, Coldwell MJ, Jackson RJ, Willis AE.
Title Protein factor requirements of the Apaf-1 internal ribosome entry segment: roles of polypyrimidine tract binding protein and upstream of N-ras.
Citation Mol Cell Biol. 2001 May;21(10):3364-74.
Year 2001
ID [U3]
Authors Tinton SA, Schepens B, Bruynooghe Y, Beyaert R, Cornelis S.
Title Regulation of the cell-cycle-dependent internal ribosome entry site of the PITSLRE protein kinase: roles of Unr (upstream of N-ras) protein and phosphorylated translation initiation factor eIF-2alpha.
Citation Biochem J. 2005 Jan 1;385(Pt 1):155-63.
Year 2005
ID [T1]
Authors Mitchell SA, Spriggs KA, Coldwell MJ, Jackson RJ, Willis AE.
Title The Apaf-1 internal ribosome entry segment attains the correct structural conformation for function via interactions with PTB and unr.
Citation Mol Cell. 2003 Mar;11(3):757-71.
Year 2003
ID [T2]
Authors Mitchell SA, Brown EC, Coldwell MJ, Jackson RJ, Willis AE.
Title Protein factor requirements of the Apaf-1 internal ribosome entry segment: roles of polypyrimidine tract binding protein and upstream of N-ras.
Citation Mol Cell Biol. 2001 May;21(10):3364-74.
Year 2001
ID [T3]
Authors Tinton SA, Schepens B, Bruynooghe Y, Beyaert R, Cornelis S.
Title Regulation of the cell-cycle-dependent internal ribosome entry site of the PITSLRE protein kinase: roles of Unr (upstream of N-ras) protein and phosphorylated translation initiation factor eIF-2alpha.
Citation Biochem J. 2005 Jan 1;385(Pt 1):155-63.
Year 2005
ID [T4]
Authors Chang TC, Yamashita A, Chen CY, Yamashita Y, Zhu W, Durdan S, Kahvejian A, Sonenberg N, Shyu AB.
Title UNR, a new partner of poly(A)-binding protein, plays a key role in translationally coupled mRNA turnover mediated by the c-fos major coding-region determinant.
Citation Genes Dev. 2004 Aug 15;18(16):2010-23.
Year 2004
ID [B1]
Authors Chang TC, Yamashita A, Chen CY, Yamashita Y, Zhu W, Durdan S, Kahvejian A, Sonenberg N, Shyu AB.
Title UNR, a new partner of poly(A)-binding protein, plays a key role in translationally coupled mRNA turnover mediated by the c-fos major coding-region determinant.
Citation Genes Dev. 2004 Aug 15;18(16):2010-23.
Year 2004
ID [B2]
Authors Hunt SL, Hsuan JJ, Totty N, Jackson RJ.
Title Unr, a cellular cytoplasmic RNA-binding protein with five cold-shock domains, is required for internal initiation of translation of human rhinovirus RNA.
Citation Genes Dev. 1999 Feb 15;13(4):437-48.
Year 1999
ID [B3]
Authors Jacquemin-Sablon H, Triqueneaux G, Deschamps S, le Maire M, Doniger J, Dautry F.
Title Nucleic acid binding and intracellular localization of unr, a protein with five cold shock domains.
Citation Nucleic Acids Res. 1994 Jul 11;22(13):2643-50.
Year 1994
ID [B4]
Authors Jeffers M, Paciucci R, Pellicer A.
Title Characterization of unr; a gene closely linked to N-ras.
Citation Nucleic Acids Res. 1990 Aug 25;18(16):4891-9.
Year 1990
ID [B5]
Authors Mitchell SA, Brown EC, Coldwell MJ, Jackson RJ, Willis AE.
Title Protein factor requirements of the Apaf-1 internal ribosome entry segment: roles of polypyrimidine tract binding protein and upstream of N-ras.
Citation Mol Cell Biol. 2001 May;21(10):3364-74.
Year 2001
ID [B6]
Authors Mitchell SA, Spriggs KA, Coldwell MJ, Jackson RJ, Willis AE.
Title The Apaf-1 internal ribosome entry segment attains the correct structural conformation for function via interactions with PTB and unr.
Citation Mol Cell. 2003 Mar;11(3):757-71.
Year 2003
ID [B7]
Authors Tinton SA, Schepens B, Bruynooghe Y, Beyaert R, Cornelis S.
Title Regulation of the cell-cycle-dependent internal ribosome entry site of the PITSLRE protein kinase: roles of Unr (upstream of N-ras) protein and phosphorylated translation initiation factor eIF-2alpha.
Citation Biochem J. 2005 Jan 1;385(Pt 1):155-63.
Year 2005
ID [B8]
Authors Triqueneaux G, Velten M, Franzon P, Dautry F, Jacquemin-Sablon H.
Title RNA binding specificity of Unr, a protein with five cold shock domains.
Citation Nucleic Acids Res. 1999 Apr 15;27(8):1926-34.
Year 1999
ID [P1]
Authors Chang TC, Yamashita A, Chen CY, Yamashita Y, Zhu W, Durdan S, Kahvejian A, Sonenberg N, Shyu AB.
Title UNR, a new partner of poly(A)-binding protein, plays a key role in translationally coupled mRNA turnover mediated by the c-fos major coding-region determinant.
Citation Genes Dev. 2004 Aug 15;18(16):2010-23.
Year 2004
ID [P2]
Authors Jeffers M, Paciucci R, Pellicer A.
Title Characterization of unr; a gene closely linked to N-ras.
Citation Nucleic Acids Res. 1990 Aug 25;18(16):4891-9.
Year 1990