242:
base pairing between the U6 snRNA in two highly conserved stem regions. It is suggested that this base-pairing interaction prevents the U6 snRNA from assembling with the U2 snRNA into the conformation required for catalytic activity. If the U4 snRNA is degraded and thereby removed from the spliceosome, splicing is effectively halted. The U4 and U6 snRNAs are demonstratively required for splicing in vitro.
251:
259:
172:
163:
40:
238:, involved with the U6 snRNA in the di-snRNP, as well as involved with both the U6 snRNA and the U5 snRNA in the tri-snRNP. The different formats have been proposed to coincide with different temporal events in the activity of the penta-snRNP, or as intermediates in the step-wise model of spliceosome assembly and activity.
278:
indicate that U4 snRNA secondary structure contains several conserved motifs, which serve structural as well as intermediary roles in establishing interactions with other splicing components. The putative U4/U6 snRNA base pairing secondary structure shown in Figure 2., is conserved across a diverse
287:
The U4 snRNA must be displaced from U6 snRNA in an ATP dependent process involving the protein Brr2 - before the spliceosome is made active. A cycle has been proposed including both Brr2 as well as the protein prp24 which selectively re-anneals U4 to the U6 snRNA. A ring of Sm proteins surround a
241:
The U4 snRNA (and its likely analog snR14 in Yeast) has been shown not to participate directly in the specific catalytic activities of the splicing reaction, and is proposed instead to act as a regulator of the U6 snRNA. The U4 snRNA inhibits spliceosome activity during assembly by complementary
992:
Mougin A, Gottschalk A, Fabrizio P, Lührmann R, Branlant C (April 2002). "Direct probing of RNA structure and RNA-protein interactions in purified HeLa cell's and yeast spliceosomal U4/U6.U5 tri-snRNP particles".
288:
conserved region of the U4 snRNA near the 3' end which are expected to promote favorable interactions between the different snRNPs as well as possibly protect the U4 snRNA from degradation by
279:
set of organisms suggesting the splicing machinery's ancient origins. It has been shown previously that a highly conserved Kinked-loop participates in specific protein interactions.
1030:"The trans-spliceosomal U4 RNA from the monogenetic trypanosomatid Leptomonas collosoma. Cloning and identification of a transcribed trna-like element that controls its expression"
643:
Siliciano PG, Brow DA, Roiha H, Guthrie C (August 1987). "An essential snRNA from S. cerevisiae has properties predicted for U4, including interaction with a U6-like snRNA".
773:
Madhani HD, Guthrie C (November 1992). "A novel base-pairing interaction between U2 and U6 snRNAs suggests a mechanism for the catalytic activation of the spliceosome".
292:
enzymes. Over 100 proteins have been identified that participate in spliceosomal pathway, several proteins of varying size are also known to interact with the U4 snRNP.
1159:
Blencowe BJ, Sproat BS, Ryder U, Barabino S, Lamond AI (November 1989). "Antisense probing of the human U4/U6 snRNP with biotinylated 2'-OMe RNA oligonucleotides".
1428:
Stark H, Dube P, Lührmann R, Kastner B (January 2001). "Arrangement of RNA and proteins in the spliceosomal U1 small nuclear ribonucleoprotein particle".
206:, and with each splicing round, it is displaced from the U6 snRNA (and the spliceosome) in an ATP-dependent manner, allowing U6 to re-fold and create the
816:
Berget SM, Robberson BL (August 1986). "U1, U2, and U4/U6 small nuclear ribonucleoproteins are required for in vitro splicing but not polyadenylation".
1205:
Raghunathan PL, Guthrie C (February 1998). "A spliceosomal recycling factor that reanneals U4 and U6 small nuclear ribonucleoprotein particles".
418:"Evidence for the existence of snRNAs U4 and U6 in a single ribonucleoprotein complex and for their association by intermolecular base pairing"
234:
The U4 snRNA has been shown to exist in a number of different formats including: bound to proteins as a small nuclear Ribo-Nuclear
Protein
175:
A 3D representation of a fragment of a U4 snRNA. The crystal structure of the spliceosomal 15.5KD protein is bound to a U4 snRNA fragment.
266:
The U4 snRNA secondary structure is suggested to alter depending on its interaction with the U6 snRNA. Several experiments involving
908:
Kambach C, Walke S, Nagai K (April 1999). "Structure and assembly of the spliceosomal small nuclear ribonucleoprotein particles".
516:"Biochemical and genetic analyses of the U5, U6, and U4/U6 x U5 small nuclear ribonucleoproteins from Saccharomyces cerevisiae"
691:"U4 small nuclear RNA dissociates from a yeast spliceosome and does not participate in the subsequent splicing reaction"
1647:
1291:"Evidence for a Prp24 binding site in U6 snRNA and in a putative intermediate in the annealing of U6 and U4 snRNAs"
1657:
275:
267:
49:
150:
1481:"Hierarchical, clustered protein interactions with U4/U6 snRNA: a biochemical role for U4/U6 proteins"
1340:"Specificity of Prp24 binding to RNA: a role for Prp24 in the dynamic interaction of U4 and U6 snRNAs"
1652:
1437:
1214:
467:"U5 small nuclear ribonucleoprotein: RNA structure analysis and ATP-dependent interaction with U4/U6"
1381:"Sm protein-Sm site RNA interactions within the inner ring of the spliceosomal snRNP core structure"
203:
45:
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369:"RNA unwinding in U4/U6 snRNPs requires ATP hydrolysis and the DEIH-box splicing factor Brr2"
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catalysis. A recycling process involving protein Brr2 releases U4 from U6, while protein
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184:
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1480:
1405:
1380:
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1290:
1250:"A stem/loop in U6 RNA defines a conformational switch required for pre-mRNA splicing"
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557:"Composition and functional characterization of the yeast spliceosomal penta-snRNP"
514:
Stevens SW, Barta I, Ge HY, Moore RE, Young MK, Lee TD, Abelson J (November 2001).
318:"Crystal structure of the spliceosomal 15.5kD protein bound to a U4 snRNA fragment"
211:
102:
861:"Pre-mRNA splicing in vitro requires intact U4/U6 small nuclear ribonucleoprotein"
555:
Stevens SW, Ryan DE, Ge HY, Moore RE, Young MK, Lee TD, Abelson J (January 2002).
1226:
943:
Comolli LR, Ulyanov NB, Soto AM, Marky LA, James TL, Gmeiner WH (October 2002).
207:
188:
250:
192:
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416:
Bringmann P, Appel B, Rinke J, Reuter R, Theissen H, Lührmann R (June 1984).
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1112:"Prp8p dissection reveals domain structure and protein interaction sites"
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Vidovic I, Nottrott S, Hartmuth K, Lührmann R, Ficner R (December 2000).
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17:
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Boon KL, Norman CM, Grainger RJ, Newman AJ, Beggs JD (February 2006).
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1071:"A simple principle to explain the evolution of pre-mRNA splicing"
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262:
Figure 2. Putative U4/U6 base pairing secondary structure.
1576:
Thomas, J; Lea K; Zucker-Aprison E; Blumenthal T (1990).
1379:
Urlaub H, Raker VA, Kostka S, Lührmann R (January 2001).
226:
of U4 in complex with a binding protein has been solved.
945:"NMR structure of the 3' stem-loop from human U4 snRNA"
738:
Guthrie C, Patterson B (1988). "Spliceosomal snRNAs".
1578:"The spliceosomal snRNAs of Caenorhabditis elegans"
198:
involved in the splicing of pre-messenger RNA (pre-
156:
144:
120:
108:
87:
82:
70:
62:
57:
32:
1479:Nottrott S, Urlaub H, Lührmann R (October 2002).
1028:Li L, Otake LR, Xu Y, Michaeli S (January 2000).
254:Figure 1. Naked U4 putative secondary structure.
311:
309:
307:
305:
1338:Ghetti A, Company M, Abelson J (April 1995).
1248:Fortner DM, Troy RG, Brow DA (January 1994).
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181:U4 small nuclear Ribo-Nucleic Acid (U4 snRNA)
8:
27:Non-coding RNA component of the spliceosome
38:
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1355:
1314:
1289:Jandrositz A, Guthrie C (February 1995).
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1069:Izquierdo JM, Valcárcel J (July 2006).
367:Raghunathan PL, Guthrie C (July 1998).
301:
187:component of the major or U2-dependent
29:
7:
859:Black DL, Steitz JA (August 1986).
752:10.1146/annurev.ge.22.120188.002131
1307:10.1002/j.1460-2075.1995.tb07060.x
465:Black DL, Pinto AL (August 1989).
434:10.1002/j.1460-2075.1984.tb01977.x
25:
689:Yean SL, Lin RJ (November 1991).
604:"Spliceosome assembly in yeast"
345:11858/00-001M-0000-0012-F75C-F
1:
922:10.1016/S0959-440X(99)80032-3
602:, Abelson J (November 1987).
574:10.1016/S1097-2765(02)00436-7
386:10.1016/S0960-9822(07)00345-4
335:10.1016/S1097-2765(00)00131-3
1629:Page for U4 spliceosomal RNA
1227:10.1126/science.279.5352.857
1173:10.1016/0092-8674(89)90036-6
878:10.1016/0092-8674(86)90345-4
830:10.1016/0092-8674(86)90344-2
787:10.1016/0092-8674(92)90556-R
657:10.1016/0092-8674(87)90031-6
274:, and chemical modification
202:). It forms a duplex with
1674:
218:re-anneals U4 and U6. The
37:
910:Curr. Opin. Struct. Biol
1397:10.1093/emboj/20.1.187
1047:10.1074/jbc.275.4.2259
1007:10.1006/jmbi.2002.5451
707:10.1128/MCB.11.11.5571
263:
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1594:10.1093/nar/18.9.2633
276:RNA structure probing
268:X-ray crystallography
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50:sequence conservation
1551:10.1093/nar/25.1.102
1497:10.1093/emboj/cdf544
621:10.1101/gad.1.9.1014
483:10.1128/MCB.9.8.3350
1535:"The uRNA database"
1442:2001Natur.409..539S
1267:10.1101/gad.8.2.221
1219:1998Sci...279..857R
1128:10.1261/rna.2281306
1088:10.1101/gad.1449106
46:secondary structure
33:U4 spliceosomal RNA
961:10.1093/nar/gkf560
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256:
177:
1648:Small nuclear RNA
1582:Nucleic Acids Res
1539:Nucleic Acids Res
1533:Zwieb, C (1997).
949:Nucleic Acids Res
220:crystal structure
196:molecular machine
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16:(Redirected from
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1653:Spliceosome
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208:active site
189:spliceosome
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1075:Genes Dev
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528:PMC
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