Small Nuclear Ribonucleoprotein (RNP) U2 Contains Numerous Additional Proteins and Has a Bipartite RNP Structure Under Splicing Conditions

REFERENCES

• Abelson, J. Personal communication.
   Google Scholar
• Abovich, N., P. Legrain, and Μ. Rosbach. 1990. The yeast PRP6 gene encodes a U4/U6 small nuclear ribonucleoprotein particle (snRNP) protein, and the PRP, gene encodes a protein required for U2 snRNP binding. Mol. Cell. Biol. 10:6417–6425.
   PubMedGoogle Scholar
• Anderson, G. J., Μ. Bach, R. Lührmann, and J. D. Beggs. 1989. Conservation between yeast and man of a protein associated with U5 small nuclear ribonucleoprotein. Nature (London) 342:819–821.
   PubMedGoogle Scholar
• Ares, M., Jr., and A. H. Igel. 1990. Lethal and temperaturesensitive mutations and their suppressors identify an essential structural element in U2 small nuclear RNA. Genes Dev. 4:2132–2145.
   PubMed Web of Science ®Google Scholar
• Bach, M., P. Bringmann, and R. Liihrmann. 1990. Purification of small nuclear ribonucleoprotein particles with antibodies against modified nucleotides of small nuclear RNAs. Methods EnzymoL 181:232–257.
   PubMedGoogle Scholar
• Bach, M., G. Winkelmann, and R. Lührmann. 1989. 20S small nuclear ribonucleoprotein U5 shows a surprisingly complex protein composition. Proc. Natl. Acad. Sci. USA 86:6038–6042.
   PubMedGoogle Scholar
• Barabino, S. Μ. L., B. J. Blencowe, U. Ryder, B. S. Sproat, and A. I. Lamond. 1990. Targeted snRNP depletion reveals an additional role for mammalian U1 snRNP in Spliceosome assembly. Cell 63:293–302.
   PubMed Web of Science ®Google Scholar
• Behrens, S.-E., F. Gallison, P. Legrain, and R. Lührmann. Unpublished data.
   Google Scholar
• Behrens, S.-E., and R. Lührmann. 1991. Immunoaffinity purification of a [U4/U6.U5]tri-snRNP from human cells. Genes Dev. 5:1439–1452.
   PubMed Web of Science ®Google Scholar
• Bentley, R. C., and J. D. Keene. 1991. Recognition of U1 and U2 small nuclear RNAs can be altered by a 5-amino-acid segment in the U2 small nuclear ribonucleoprotein particle (snRNP) B″ protein and through interactions with U2 snRNP-A′ protein. MoL Cell. Biol. 11:1829–1839.
   PubMed Web of Science ®Google Scholar
• Black, D. L., B. Chabot, and J. A. Steitz. 1985. U2 as well as U1 small ribonucleoproteins are involved in premessenger RNA splicing. Cell 42:737–750.
   PubMed Web of Science ®Google Scholar
• Black, D. L., and A. L. Pinto. 1989. U5 small nuclear ribonucleoprotein: RNA structure analysis and ATP-dependent interaction with U4/U6. MoL Cell. Biol. 9:3350–3359.
   PubMedGoogle Scholar
• Bochnig, P., R. Reuter, P. Bringmann, and R. Liihrmann. 1987. A monoclonal antibody against 2,2,7-trimethylguanosine that reacts with intact U snRNPs as well as with 7-methylguanosine- capped RNAs. Eur. J. Biochem. 168:461–467.
   PubMedGoogle Scholar
• Bringmann, P., and R. Lührmann. 1986. Purification of the individual snRNPs Ul, U2, U5 and U4/U6 from HeLa cells and characterization of their protein constituents. EMBO J. 5:3509–3516.
   PubMed Web of Science ®Google Scholar
• Bringmann, P., and R. Liihrmann. 1987. Antibodies specific for N6-methyladenosine react with intact snRNPs U2 and U4/U6. FEBS Lett. 213:309–315.
   PubMed Web of Science ®Google Scholar
• Bringmann, P., J. Rinke, B. Appel, R. Reuter, and R. Liihr- mann. 1983. Purification of snRNPs Ul, U2, U4, U5 and U6 with 2,2,7-trimethylguanosine-specific antibody and definition of their constituent proteins reacting with anti-Sm and anti- (Ul)RNP antisera. EMBO J. 2:1129–1135.
   PubMed Web of Science ®Google Scholar
• Buvoli, M., G. Biamonti, P. Tsoulfor, Μ. T. Bassi, A. Ghetti, S. Riva, and C. Morandi. 1988. cDNA cloning of human hnRNP protein A1 reveals the existence of multiple mRNA isoforms. Nucleic Acids Res. 16:3751–3770.
   PubMed Web of Science ®Google Scholar
• Datta, B., and A. Μ. Weiner. 1991. Genetic evidence for base pairing between U2 and U6 snRNA in mammalian mRNA splicing. Nature (London) 352:821–824.
   PubMed Web of Science ®Google Scholar
• Dignam, J. D., R. Μ. Lebovitz, and R. G∙ Roeder. 1983. Accurate transcription initiation by RNA polymerase II in a soluble extract from isolated mammalian nuclei. Nucleic Acids Res. 11:1475–1489.
   PubMed Web of Science ®Google Scholar
• Ge, H., and J. L. Manley. 1990. A protein factor, ASF, controls cell-specific alternative splicing of SV40 early pre-mRNA in vitro. Cell 62:25–34.
   PubMed Web of Science ®Google Scholar
• Hamm, J., N. A. Dathan, and I. W. Mattaj. 1989. Functional analysis of mutant Xenopus U2 snRNAs. Cell 59:159–169.
   PubMed Web of Science ®Google Scholar
• Hausner, T.-P., L. Μ. Giglio, and A. Μ. Weiner. 1990. Evidence for base-pairing between mammalian U2 and U6 small nuclear ribonucleoprotein particles. Genes Dev. 4:2146–2156.
   PubMedGoogle Scholar
• Kastner, B., Μ. Bach, and R. Liihrmann. 1990. Electron microscopy of small nuclear ribonucleoprotein (snRNP) particles U2 and U5: evidence for a common structure-determining principle in the major U snRNP family. Proc. Natl. Acad. Sci. USA 87:1710–1714.
   PubMed Web of Science ®Google Scholar
• Kastner, B., and R. Liihrmann. 1989. Electron microscopy of U1 small nuclear ribonucleoprotein particles: shape of the particle and position of the 5′ RNA terminus. EMBO J. 8:277–286.
   PubMed Web of Science ®Google Scholar
• Konarska, Μ. Μ., and P. A. Sharp. 1987. Interactions between small nuclear ribonucleoprotein particles in formation of spliceosomes. Cell 49:763–774.
   PubMed Web of Science ®Google Scholar
• Krainer, A. R., G. C. Conway, and D. Kozak. 1990. Purification and characterization of pre-mRNA splicing factor SF2 from HeLa cells. Genes Dev. 4:1158–1171.
   PubMed Web of Science ®Google Scholar
• Krainer, A. R., and T. Maniatis. 1985. Multiple factors including the small nuclear ribonucleoproteins U1 and U2 are necessary for pre-mRNA splicing in vitro. Cell 42:725–736.
   PubMed Web of Science ®Google Scholar
• Krämer, A. 1988. Presplicing complex formation requires two proteins and U2 snRNP. Genes Dev. 2:1155–1167.
   PubMedGoogle Scholar
• Lehmeier, T., K. Foulaki, and R. Lührmann. 1990. Evidence for three distinct D proteins, which react differentially with anti-Sm autoantibodies, in the cores of the major snRNPs U1, U2, U4/U6 and U5. Nucleic Acids Res. 18:6475–6484.
   PubMedGoogle Scholar
• Lührmann, R., B. Kastner, and Μ. Bach. 1990. Structure of Spliceosomal snRNPs and their role in pre-mRNA splicing. Biochim. Biophys. Acta 1087:265–292.
   PubMed Web of Science ®Google Scholar
• Mattaj, I. W., and E. Μ. De Robertis. 1985. Nuclear segregation of U2 snRNA requires binding of specific snRNP proteins. Cell 40:111–118.
   PubMed Web of Science ®Google Scholar
• Mattaj, I. W., W. J. Habets, and W. J. van Venrooij. 1986. Monospecific antibodies reveal details of U2 snRNP structure and interaction between U1 and U2 snRNPs. EMBO J. 5:997–1002.
   PubMed Web of Science ®Google Scholar
• Moazed, D., S. Stern, and H. F. Noller. 1986. Rapid chemical probing of conformation in 16S ribosomal RNA and 30S ribosomal subunits using primer extension. J. Mol. Biol. 187:399–418.
   PubMed Web of Science ®Google Scholar
• Parker, R., P. G. Siliciano, and C. Guthrie. 1987. Recognition of the TACTAAC box during mRNA splicing in yeast involves base pairing to the U2-like snRNP. Cell 49:229–239.
   PubMed Web of Science ®Google Scholar
• Reed, R., and T. Maniatis. 1988. The role of the mammalian branchpoint sequence in pre-mRNA splicing. Genes Dev. 2:1268–1276.
   PubMedGoogle Scholar
• Reuter, R., and R. Liihrmann. 1986. Immunization of mice with purified U1 small nuclear ribonucleoprotein (RNP) induces a pattern of antibody specificities characteristic of the anti-Sm and anti-RNP autoimmune response of patients with lupus erythematosus, as measured by monoclonal antibodies. Proc. Natl. Acad. Sci. USA 83:8689–8693.
   PubMed Web of Science ®Google Scholar
• Ruby, S. W., and J. Abelson. 1988. An early hierarchic role of U1 small nuclear ribonucleoprotein in Spliceosome assembly. Science 242:1028–1035.
   PubMed Web of Science ®Google Scholar
• Ruskin, B., and Μ. R. Green. 1985. Specific and stable intronfactor interactions are established early during in vitro pre- mRNA splicing. Cell 43:131–142.
   PubMed Web of Science ®Google Scholar
• Ruskin, B., J. Μ. Greene, and Μ. R. Green. 1985. Cryptic branchpoint activation allows accurate in vitro splicing of human beta-globin intron mutants. Cell 41:833–844.
   PubMedGoogle Scholar
• Ruskin, B., P. D. Zamore, and Μ. R. Green. 1988. A factor, U2AF, is required for U2 snRNP binding and splicing complex assembly. Cell 52:207–219.
   PubMed Web of Science ®Google Scholar
• Scherly, D., N. A. Dathan, W. Boelens, W. J. van Venrooij, and I. W. Mattaj. 1990. The U2B″ RNP motif as a site of proteinprotein interaction. EMBO J. 9:3675–3681.
   PubMedGoogle Scholar
• Séraphin, B., L. Kretzner, and Μ. Rosbash. 1988. A U1 snRNA: pre-mRNA base pairing interaction is required early in yeast Spliceosome assembly but does not uniquely define the 5′ cleavage site. EMBO J. 7:2533–2538.
   PubMed Web of Science ®Google Scholar
• Séraphin, B., and Μ. Rosbash. 1989. Identification of functional U1 snRNA-Pre-mRNA complexes committed to Spliceosome assembly and splicing. Cell 59:349–358.
   PubMed Web of Science ®Google Scholar
• Temsamani, J., Μ. Rhoadhouse, and T. Pederson. 1991. The U2 small nuclear ribonucleoprotein particle associates with nuclear factors in a pre-mRNA independent reaction. J. Biol. Chem. 266:20356–20362.
   PubMedGoogle Scholar
• Utans, U., S.-E. Behrens, R. Liihrmann, R. Kole, and A. Kramer. 1992. A splicing factor which is inactivated during in vivo heat shock is functionally equivalent to the [U4/ U6.U5]triple-snRNP-specific proteins. Genes Dev. 5:1439–1452.
   Google Scholar
• Wu, J., and J. L. Manley. 1989. Mammalian pre-mRNA branch site selection by U2 snRNP involves base pairing. Genes Dev. 3:1553–1561.
   PubMed Web of Science ®Google Scholar
• Wu, J., and J. L. Manley. 1991. Base pairing between U2 and U6 snRNAs is necessary for splicing of a mammalian pre-mRNA. Nature (London) 352:818–821.
   PubMedGoogle Scholar
• Zamore, P. D., and Μ. R. Green. 1991. Biochemical characterization of U2 snRNP auxiliary factor: an essential pre-mRNA splicing factor with a novel intranuclear distribution. EMBO J. 10:207–214.
   PubMed Web of Science ®Google Scholar
• Zhuang, Y., and A. Μ. Weiner. 1989. A compensatory base change in human U2 snRNA can suppress a branch site mutation. Genes Dev. 3:1545–1552.
   PubMed Web of Science ®Google Scholar