338:
how U4 is displaced from U6 snRNA, although RNA has been implicated in spliceosome assembly, and may function to unwind U4/U6 and promote the formation of a U2/U6 snRNA interaction. The interactions of U4/U6 stem loops I and II dissociate and the freed stem loop II region of U6 folds on itself to form an intramolecular stem loop and U4 is no longer required in further spliceosome assembly. The freed stem loop I region of U6 base pairs with U2 snRNA forming the U2/U6 helix I. However, the helix I structure is mutually exclusive with the 3' half of an internal 5' stem loop region of U2 snRNA.
70:
221:
197:(also known as heterogeneous nuclear RNA, hn-RNA) at each exon:intron junction. The pre-mRNA introns contains specific sequence elements that are recognized and utilized during spliceosome assembly. These include the 5' end splice site, the branch point sequence, the polypyrimidine tract, and the 3' end splice site. The spliceosome catalyzes the removal of introns, and the ligation of the flanking exons.
322:
step of splicing. The U4/U5/U6 tri-snRNP (see Figure 1) is recruited to the assembling spliceosome to form complex B, and following several rearrangements, complex C is activated for catalysis. It is unclear how the tri-snRNP is recruited to complex A, but this process may be mediated through protein-protein interactions and/or base pairing interactions between U2 snRNA and U6 snRNA.
314:(U2 snRNP auxiliary factor) and possibly U1 snRNP. In an ATP-dependent reaction, U2 snRNP becomes tightly associated with the branch point sequence (BPS) to form complex A. A duplex formed between U2 snRNP and the pre-mRNA branch region bulges out the branch adenosine specifying it as the nucleophile for the first transesterification.
334:
5' splice site. Binding of U2 snRNP to the branch point sequence (BPS) is one example of an RNA-RNA interaction displacing a protein-RNA interaction. Upon recruitment of U2 snRNP, the branch binding protein SF1 in the commitment complex is displaced since the binding site of U2 snRNA and SF1 are mutually exclusive events.
376:, together with U5, are subunits of the minor spliceosome that splices a rare class of pre-mRNA introns, denoted U12-type. The minor spliceosome is located in the nucleus like its major counterpart, though there are exceptions in some specialised cells including anucleate platelets and the dendroplasm (
328:
Upon recruitment of the tri-snRNP, several RNA-RNA rearrangements precede the first catalytic step and further rearrangements occur in the catalytically active spliceosome. Several of the RNA-RNA interactions are mutually exclusive; however, it is not known what triggers these interactions, nor the
300:
is a biochemical reaction, and like all biochemical reactions, its rate depends on the concentration of enzymes and substrates. In this case, the enzymes are the spliceosomes, and the substrates are the pre-mRNAs. By varying the concentration of spliceosomes and pre-mRNAs based on their proximity to
337:
Within the U2 snRNA, there are other mutually exclusive rearrangements that occur between competing conformations. For example, in the active form, stem loop IIa is favored; in the inactive form a mutually exclusive interaction between the loop and a downstream sequence predominates. It is unclear
333:
from the 5' splice site and formation of a U6 snRNA interaction. It is known that U1 snRNP is only weakly associated with fully formed spliceosomes, and U1 snRNP is inhibitory to the formation of a U6-5' splice site interaction on a model of substrate oligonucleotide containing a short 5' exon and
321:
residue in U2 snRNA, nearly opposite of the branch site, results in an altered conformation of the RNA-RNA duplex upon the U2 snRNP binding. Specifically, the altered structure of the duplex induced by the pseudouridine places the 2' OH of the bulged adenosine in a favorable position for the first
309:
The model for formation of the spliceosome active site involves an ordered, stepwise assembly of discrete snRNP particles on the pre-mRNA substrate. The first recognition of pre-mRNAs involves U1 snRNP binding to the 5' end splice site of the pre-mRNA and other non-snRNP associated factors to form
295:
known as nuclear speckles. It was originally postulated that nuclear speckles are either sites of mRNA splicing or storage sites of mRNA splicing factors. It is now understood that nuclear speckles help concentrate splicing factors near genes that are physically located close to them. Genes located
236:
Cryo-EM has been applied extensively by Shi et al. to elucidate the near-/atomic structure of spliceosome in both yeast and humans. The molecular framework of spliceosome at near-atomic-resolution demonstrates Spp42 component of U5 snRNP forms a central scaffold and anchors the catalytic center in
43:) and numerous proteins. Small nuclear RNA (snRNA) molecules bind to specific proteins to form a small nuclear ribonucleoprotein complex (snRNP, pronounced "snurps"), which in turn combines with other snRNPs to form a large ribonucleoprotein complex called a spliceosome. The spliceosome removes
232:) tri-snRNPs. Below left is a schematic illustration of the interaction of tri-snRNP proteins with the U4/U6 snRNA duplex. Below right is a cartoon model of the yeast tri-snRNP with shaded areas corresponding to U5 (gray), U4/U6 (orange) and the linker region (yellow).
237:
yeast. The atomic structure of the human spliceosome illustrates the step II component Slu7 adopts an extended structure, poised for selection of the 3'-splice site. All five metals (assigned as Mg2+) in the yeast complex are preserved in the human complex.
310:
the commitment complex, or early (E) complex in mammals. The commitment complex is an ATP-independent complex that commits the pre-mRNA to the splicing pathway. U2 snRNP is recruited to the branch region through interactions with the E complex component
216:
Many proteins exhibit a zinc-binding motif, which underscores the importance of zinc in the splicing mechanism. The first molecular-resolution reconstruction of U4/U6.U5 triple small nuclear ribonucleoprotein (tri-snRNP) complex was reported in 2016.
100:
that is not involved in protein expression. The split gene structure was found when adenoviral mRNAs were hybridized to endonuclease cleavage fragments of single stranded viral DNA. It was observed that the mRNAs of the mRNA-DNA hybrids contained
109:
tails of non-hydrogen bonded regions. When larger fragments of viral DNAs were used, forked structures of looped out DNA were observed when hybridized to the viral mRNAs. It was realised that the looped out regions, the
200:
Introns typically have a GU nucleotide sequence at the 5' end splice site, and an AG at the 3' end splice site. The 3' splice site can be further defined by a variable length of polypyrimidines, called the
743:
Häcker I, Sander B, Golas MM, Wolf E, Karagöz E, Kastner B, et al. (November 2008). "Localization of Prp8, Brr2, Snu114 and U4/U6 proteins in the yeast tri-snRNP by electron microscopy".
1466:
Konforti BB, Koziolkiewicz MJ, Konarska MM (December 1993). "Disruption of base pairing between the 5' splice site and the 5' end of U1 snRNA is required for spliceosome assembly".
510:
Chow LT, Roberts JM, Lewis JB, Broker TR (August 1977). "A map of cytoplasmic RNA transcripts from lytic adenovirus type 2, determined by electron microscopy of RNA:DNA hybrids".
325:
The U5 snRNP interacts with sequences at the 5' and 3' splice sites via the invariant loop of U5 snRNA and U5 protein components interact with the 3' splice site region.
96:
labs revealed that genes of higher organisms are "split" or present in several distinct segments along the DNA molecule. The coding regions of the gene are separated by
1819:
206:
142:(snRNA) and a range of associated protein factors. When these small RNAs are combined with the protein factors, they make RNA-protein complexes called
114:, are excised from the precursor mRNAs in a process Sharp named "splicing". The split gene structure was subsequently found to be common to most
1863:
1812:
1069:
Seraphin B, Rosbash M (October 1989). "Identification of functional U1 snRNA-pre-mRNA complexes committed to spliceosome assembly and splicing".
1415:
Moore MJ, Sharp PA (September 1993). "Evidence for two active sites in the spliceosome provided by stereochemistry of pre-mRNA splicing".
1805:
330:
296:
farther from speckles can still be transcribed and spliced, but their splicing is less efficient compared to those closer to speckles.
66:
However, sometimes the RNA within the intron acts as a ribozyme, splicing itself without the use of a spliceosome or protein enzymes.
2026:
2021:
1245:
Burge CB, Tuschl T, Sharp PA (1999). "Splicing precursors to mRNAs by the spliceosomes". In
Gesteland RF, Cech TR, Atkins JF (eds.).
1202:
Newby MI, Greenbaum NL (December 2002). "Sculpting of the spliceosomal branch site recognition motif by a conserved pseudouridine".
2031:
2006:
2104:
786:
Yan C, Hang J, Wan R, Huang M, Wong CC, Shi Y (September 2015). "Structure of a yeast spliceosome at 3.6-angstrom resolution".
1728:
1254:
127:
1345:
604:"Stalling of spliceosome assembly at distinct stages by small-molecule inhibitors of protein acetylation and deacetylation"
205:(PPT), which serves the dual function of recruiting factors to the 3' splice site and possibly recruiting factors to the
1873:
1828:
555:"Semiquantitative proteomic analysis of the human spliceosome via a novel two-dimensional gel electrophoresis method"
653:"Molecular architecture of zinc chelating small molecules that inhibit spliceosome assembly at an early stage"
1316:"snRNA interactions at 5' and 3' splice sites monitored by photoactivated crosslinking in yeast spliceosomes"
373:
369:
59:. An analogy is a film editor, who selectively cuts out irrelevant or incorrect material (equivalent to the
1711:
Butcher SE (2011). "Chapter 8. The
Spliceosome and Its Metal Ions". In Sigel A, Sigel H, Sigel RK (eds.).
1868:
2142:
1746:
Nilsen TW (December 2003). "The spliceosome: the most complex macromolecular machine in the cell?".
2137:
1791:
48:
2074:
1895:
2147:
1599:
Denis MM, Tolley ND, Bunting M, Schwertz H, Jiang H, Lindemann S, et al. (August 2005).
880:"Drosophila Dscam is an axon guidance receptor exhibiting extraordinary molecular diversity"
553:
Agafonov DE, Deckert J, Wolf E, Odenwälder P, Bessonov S, Will CL, et al. (July 2011).
63:) from the initial film and sends the cleaned-up version to the director for the final cut.
2127:
1958:
1661:
1648:
Glanzer J, Miyashiro KY, Sul JY, Barrett L, Belt B, Haydon P, et al. (November 2005).
1424:
795:
464:
246:
202:
1601:"Escaping the nuclear confines: signal-dependent pre-mRNA splicing in anucleate platelets"
8:
2132:
1951:
1163:"Branch nucleophile selection in pre-mRNA splicing: evidence for the bulged duplex model"
225:
119:
89:
83:
1787:
1665:
1428:
997:
972:
799:
468:
329:
order of these rearrangements. The first rearrangement is probably the displacement of
1684:
1649:
1625:
1600:
1576:
1549:
1548:
Pessa HK, Will CL, Meng X, Schneider C, Watkins NJ, Perälä N, et al. (June 2008).
1530:
1448:
1392:
1368:"Evidence that U5 snRNP recognizes the 3' splice site for catalytic step II in mammals"
1367:
1332:
1227:
1094:
945:
920:
819:
768:
725:
677:
652:
628:
603:
579:
554:
535:
425:
400:
194:
166:
roteins, pronounced "snurps"). The snRNAs that make up the major spliceosome are named
123:
93:
52:
1291:
1274:
1138:
1113:
1046:
1021:
896:
879:
487:
452:
1946:
1763:
1734:
1724:
1689:
1630:
1581:
1522:
1483:
1479:
1440:
1397:
1337:
1296:
1250:
1219:
1184:
1143:
1086:
1082:
1051:
1002:
950:
901:
860:
811:
760:
729:
717:
682:
633:
584:
527:
523:
492:
430:
357:
347:
256:
139:
24:
1797:
1534:
1231:
1098:
823:
772:
539:
1755:
1716:
1679:
1669:
1620:
1612:
1571:
1561:
1514:
1475:
1452:
1432:
1387:
1379:
1327:
1286:
1211:
1174:
1133:
1125:
1078:
1041:
1033:
992:
984:
940:
932:
891:
878:
Schmucker D, Clemens JC, Shu H, Worby CA, Xiao J, Muda M, et al. (June 2000).
850:
803:
752:
709:
672:
664:
623:
615:
574:
566:
519:
482:
472:
420:
412:
32:
1905:
1888:
1883:
1858:
700:
Cate JH (March 2016). "STRUCTURE. A Big Bang in spliceosome structural biology".
1720:
1315:
936:
416:
2069:
1968:
1853:
1848:
1654:
Proceedings of the
National Academy of Sciences of the United States of America
1616:
1554:
Proceedings of the
National Academy of Sciences of the United States of America
1502:
1383:
988:
855:
838:
457:
Proceedings of the
National Academy of Sciences of the United States of America
301:
nuclear speckles, cells could potentially regulate the efficiency of splicing.
292:
260:
97:
69:
2121:
1715:. Metal Ions in Life Sciences. Vol. 9. RSC Publishing. pp. 235–51.
973:"Genome organization around nuclear speckles drives mRNA splicing efficiency"
602:
Kuhn AN, van Santen MA, Schwienhorst A, Urlaub H, LĂĽhrmann R (January 2009).
318:
1674:
1566:
1275:"Mechanical devices of the spliceosome: motors, clocks, springs, and things"
807:
713:
651:
Patil V, Canzoneri JC, Samatov TR, LĂĽhrmann R, Oyelere AK (September 2012).
263:
have been used to account for the relatively small number of protein coding
1915:
1836:
1782:
1767:
1738:
1693:
1634:
1585:
1526:
1223:
1037:
1006:
954:
905:
864:
815:
764:
721:
686:
668:
637:
588:
477:
434:
297:
268:
56:
28:
1505:(December 2003). "Splicing double: insights from the second spliceosome".
1487:
1444:
1401:
1341:
1300:
1188:
1147:
1090:
1055:
2094:
1900:
1550:"Minor spliceosome components are predominantly localized in the nucleus"
1129:
570:
531:
496:
1783:
3D macromolecular structures of
Spliceosomes from the EM Data Bank(EMDB)
1179:
1162:
619:
279:, has been speculated to be alternatively spliced into 38,000 different
1759:
272:
756:
291:
Pre-mRNA splicing factors were originally found to be concentrated in
1436:
365:
361:
353:
210:
115:
16:
Molecular machine that removes intron RNA from the primary transcript
1518:
283:, assuming all of its exons can splice independently of each other.
1215:
377:
190:, and participate in several RNA-RNA and RNA-protein interactions.
183:
179:
175:
171:
167:
220:
2061:
2041:
2036:
187:
60:
601:
2016:
2011:
1991:
1927:
1878:
1365:
1114:"Early commitment of yeast pre-mRNA to the spliceosome pathway"
453:"Spliced segments at the 5' terminus of adenovirus 2 late mRNA"
111:
44:
1366:
Chiara MD, Palandjian L, Feld Kramer R, Reed R (August 1997).
2084:
2079:
2001:
1996:
1986:
1981:
1976:
1936:
1494:
1465:
650:
552:
311:
276:
143:
106:
102:
40:
36:
2099:
2089:
1931:
837:
Zhang X, Yan C, Hang J, Finci LI, Lei J, Shi Y (May 2017).
280:
264:
252:
1647:
1598:
394:
392:
1022:"The spliceosome assembly pathway in mammalian extracts"
1019:
877:
271:, currently estimated at around 20,000. One particular
130:
for the discovery of introns and the splicing process.
1547:
1111:
509:
389:
251:
Alternative splicing (the re-combination of different
1827:
1313:
1020:
Jamison SF, Crow A, Garcia-Blanco MA (October 1992).
742:
1650:"RNA splicing capability of live neuronal dendrites"
1713:
Structural and catalytic roles of metal ions in RNA
1314:Newman AJ, Teigelkamp S, Beggs JD (November 1995).
1112:Legrain P, Seraphin B, Rosbash M (September 1988).
1249:. Cold Spring Harbor Lab. Press. pp. 525–60.
971:Bhat P, Chow A, Emert B, et al. (May 2024).
836:
2119:
1244:
450:
1160:
1068:
785:
193:The assembly of the spliceosome occurs on each
1268:
1266:
1201:
839:"An Atomic Structure of the Human Spliceosome"
1813:
1272:
970:
918:
871:
1592:
966:
964:
451:Berget SM, Moore C, Sharp PA (August 1977).
398:
55:. This process is generally referred to as
1500:
1459:
1359:
1263:
1161:Query CC, Moore MJ, Sharp PA (March 1994).
1105:
736:
1820:
1806:
1414:
925:Cold Spring Harbor Perspectives in Biology
405:Cold Spring Harbor Perspectives in Biology
1790:at the U.S. National Library of Medicine
1683:
1673:
1624:
1575:
1565:
1391:
1331:
1290:
1178:
1137:
1045:
996:
961:
944:
895:
854:
745:Nature Structural & Molecular Biology
676:
627:
578:
486:
476:
424:
356:have a second spliceosome, the so-called
213:required for the first step of splicing.
27:(RNP) complex found primarily within the
1307:
446:
444:
219:
68:
1710:
286:
240:
209:(BPS). The BPS contains the conserved
2120:
1745:
1507:Nature Reviews. Molecular Cell Biology
1273:Staley JP, Guthrie C (February 1998).
1801:
441:
186:, so-called because they are rich in
138:Each spliceosome is composed of five
128:Nobel Prize in Physiology or Medicine
699:
401:"Spliceosome structure and function"
341:
228:fields of negatively stained yeast (
35:. The spliceosome is assembled from
503:
360:. A group of less abundant snRNAs,
13:
1703:
919:Spector DL, Lamond AI (Feb 2011).
14:
2159:
1829:Post-transcriptional modification
1776:
399:Will CL, LĂĽhrmann R (July 2011).
1641:
1541:
1408:
1348:from the original on 2005-02-23
1238:
1195:
1154:
1062:
1013:
912:
830:
1118:Molecular and Cellular Biology
1026:Molecular and Cellular Biology
779:
693:
644:
595:
559:Molecular and Cellular Biology
546:
380:cytoplasm) of neuronal cells.
133:
1:
1292:10.1016/S0092-8674(00)80925-3
897:10.1016/S0092-8674(00)80878-8
383:
1480:10.1016/0092-8674(93)90531-T
1083:10.1016/0092-8674(89)90296-1
524:10.1016/0092-8674(77)90294-X
7:
1721:10.1039/9781849732512-00235
937:10.1101/cshperspect.a000646
417:10.1101/cshperspect.a003707
304:
73:Spliceosomal splicing cycle
10:
2164:
1617:10.1016/j.cell.2005.06.015
989:10.1038/s41586-024-07429-6
856:10.1016/j.cell.2017.04.033
345:
244:
81:
77:
2060:
1967:
1923:
1914:
1844:
1835:
1204:Nature Structural Biology
1792:Medical Subject Headings
1384:10.1093/emboj/16.15.4746
230:Saccharomyces cerevisiae
1896:Poly(A)-binding protein
1675:10.1073/pnas.0503783102
1567:10.1073/pnas.0803646105
1167:Genes & Development
808:10.1126/science.aac7629
714:10.1126/science.aaf4465
255:) is a major source of
1038:10.1128/MCB.12.10.4279
669:10.1261/rna.034819.112
478:10.1073/pnas.74.8.3171
233:
126:were awarded the 1993
74:
223:
207:branch point sequence
88:In 1977, work by the
72:
1959:Alternative splicing
1130:10.1128/MCB.8.9.3755
571:10.1128/mcb.05266-11
287:Location of splicing
247:Alternative splicing
241:Alternative splicing
224:Figure 1. Above are
203:polypyrimidine tract
51:pre-mRNA, a type of
1666:2005PNAS..10216859G
1429:1993Natur.365..364M
1180:10.1101/gad.8.5.587
800:2015Sci...349.1182Y
620:10.1261/rna.1332609
469:1977PNAS...74.3171B
226:electron microscopy
84:Splicing (genetics)
2070:5′ cap methylation
1760:10.1002/bies.10394
921:"Nuclear speckles"
849:(5): 918–929.e14.
317:The presence of a
234:
140:small nuclear RNAs
124:Richard J. Roberts
75:
53:primary transcript
37:small nuclear RNAs
2143:Protein complexes
2115:
2114:
2056:
2055:
2052:
2051:
1969:pre-mRNA factors
1730:978-1-84973-094-5
1256:978-0-87969-380-0
794:(6253): 1182–91.
757:10.1038/nsmb.1506
358:minor spliceosome
348:Minor spliceosome
342:Minor spliceosome
257:genetic diversity
25:ribonucleoprotein
2155:
1921:
1920:
1854:5′ cap formation
1842:
1841:
1822:
1815:
1808:
1799:
1798:
1771:
1742:
1698:
1697:
1687:
1677:
1660:(46): 16859–64.
1645:
1639:
1638:
1628:
1596:
1590:
1589:
1579:
1569:
1545:
1539:
1538:
1498:
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1463:
1457:
1456:
1437:10.1038/365364a0
1412:
1406:
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1395:
1372:The EMBO Journal
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1141:
1109:
1103:
1102:
1066:
1060:
1059:
1049:
1017:
1011:
1010:
1000:
983:(5): 1165–1173.
968:
959:
958:
948:
916:
910:
909:
899:
875:
869:
868:
858:
834:
828:
827:
783:
777:
776:
740:
734:
733:
708:(6280): 1390–2.
697:
691:
690:
680:
648:
642:
641:
631:
599:
593:
592:
582:
550:
544:
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507:
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33:eukaryotic cells
2163:
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2158:
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2138:Gene expression
2118:
2117:
2116:
2111:
2048:
1963:
1910:
1906:Polyuridylation
1859:Polyadenylation
1831:
1826:
1779:
1774:
1731:
1706:
1704:Further reading
1701:
1646:
1642:
1597:
1593:
1560:(25): 8655–60.
1546:
1542:
1519:10.1038/nrm1259
1499:
1495:
1464:
1460:
1423:(6444): 364–8.
1413:
1409:
1378:(15): 4746–59.
1364:
1360:
1351:
1349:
1312:
1308:
1271:
1264:
1257:
1243:
1239:
1200:
1196:
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1067:
1063:
1032:(10): 4279–87.
1018:
1014:
969:
962:
917:
913:
876:
872:
835:
831:
784:
780:
751:(11): 1206–12.
741:
737:
698:
694:
649:
645:
600:
596:
565:(13): 2667–82.
551:
547:
508:
504:
449:
442:
397:
390:
386:
350:
344:
307:
289:
261:Splice variants
259:in eukaryotes.
249:
243:
136:
86:
80:
17:
12:
11:
5:
2161:
2151:
2150:
2145:
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2112:
2110:
2109:
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2102:
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2075:mRNA decapping
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2019:
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1849:Precursor mRNA
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1777:External links
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1513:(12): 960–70.
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1237:
1216:10.1038/nsb873
1210:(12): 958–65.
1194:
1153:
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1104:
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911:
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829:
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663:(9): 1605–11.
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245:Main article:
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98:non-coding DNA
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15:
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1329:
1326:(9): 968–80.
1325:
1321:
1317:
1310:
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1298:
1293:
1288:
1285:(3): 315–26.
1284:
1280:
1276:
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1258:
1252:
1248:
1247:The RNA World
1241:
1233:
1229:
1225:
1221:
1217:
1213:
1209:
1205:
1198:
1190:
1186:
1181:
1176:
1173:(5): 587–97.
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1131:
1127:
1123:
1119:
1115:
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1077:(2): 349–58.
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890:(6): 671–84.
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844:
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621:
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614:(1): 153–75.
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581:
576:
572:
568:
564:
560:
556:
549:
541:
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518:(4): 819–36.
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470:
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463:(8): 3171–5.
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319:pseudouridine
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120:Phillip Sharp
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58:
54:
50:
46:
42:
38:
34:
30:
26:
22:
2148:RNA splicing
1941:
1930: /
1916:RNA splicing
1788:Spliceosomes
1751:
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1712:
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1643:
1608:
1604:
1594:
1557:
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1543:
1510:
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1350:. Retrieved
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408:
404:
351:
336:
327:
324:
316:
308:
298:RNA splicing
290:
269:human genome
250:
235:
229:
215:
199:
192:
163:
159:
155:
151:
147:
137:
87:
65:
20:
18:
2128:Spliceosome
1942:Spliceosome
1901:RNA editing
134:Composition
49:transcribed
23:is a large
21:spliceosome
2133:Organelles
2122:Categories
1501:Patel AA,
1352:2008-03-07
923:. Review.
384:References
354:eukaryotes
346:See also:
273:Drosophila
116:eukaryotic
82:See also:
2062:Cytosolic
1748:BioEssays
1503:Steitz JA
730:206648185
211:adenosine
1768:14635248
1739:22010274
1694:16275927
1635:16096058
1586:18559850
1535:21816910
1527:14685174
1346:Archived
1232:39628664
1224:12426583
1099:18553973
1007:38720076
998:11164319
955:20926517
906:10892653
865:28502770
824:22194712
816:26292707
773:22982227
765:18953335
722:27013712
687:22832025
638:19029308
589:21536652
540:37967144
435:21441581
378:dendrite
331:U1 snRNP
305:Assembly
195:pre-mRNA
118:genes.
57:splicing
2042:PRPF40B
2037:PRPF40A
2027:PRPF38B
2022:PRPF38A
1837:Nuclear
1685:1277967
1662:Bibcode
1626:4401993
1577:2438382
1488:8252623
1453:4361512
1445:8397340
1425:Bibcode
1402:9303319
1393:1170101
1342:8548661
1333:1369345
1301:9476892
1189:7926752
1148:3065622
1091:2529976
1056:1383687
946:3039535
796:Bibcode
788:Science
702:Science
678:3425776
629:2612777
580:3133382
465:Bibcode
426:3119917
267:in the
188:uridine
154:uclear
112:introns
94:Roberts
78:History
61:introns
47:from a
45:introns
29:nucleus
2032:PRPF39
2017:PRPF31
2012:PRPF19
2007:PRPF18
1992:PRPF4B
1928:Intron
1794:(MeSH)
1766:
1737:
1727:
1692:
1682:
1633:
1623:
1584:
1574:
1533:
1525:
1486:
1451:
1443:
1417:Nature
1400:
1390:
1340:
1330:
1299:
1253:
1230:
1222:
1187:
1146:
1139:365433
1136:
1097:
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1047:360351
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1005:
995:
977:Nature
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943:
904:
863:
822:
814:
771:
763:
728:
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685:
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538:
532:890740
530:
497:269380
495:
488:431482
485:
433:
423:
374:U6atac
372:, and
370:U4atac
275:gene,
182:, and
144:snRNPs
2085:DCP1B
2080:DCP1A
2002:PRPF8
1997:PRPF6
1987:PRPF4
1982:PRPF3
1977:PLRG1
1947:minor
1937:snRNP
1531:S2CID
1449:S2CID
1228:S2CID
1095:S2CID
820:S2CID
769:S2CID
726:S2CID
536:S2CID
352:Some
281:mRNAs
277:Dscam
265:genes
253:exons
162:ucleo
150:mall
90:Sharp
41:snRNA
2105:EDC4
2100:EDC3
2095:DCPS
2090:DCP2
1932:Exon
1889:CFII
1879:PAB2
1869:CstF
1864:CPSF
1764:PMID
1735:PMID
1725:ISBN
1690:PMID
1631:PMID
1605:Cell
1582:PMID
1523:PMID
1484:PMID
1468:Cell
1441:PMID
1398:PMID
1338:PMID
1297:PMID
1279:Cell
1251:ISBN
1220:PMID
1185:PMID
1144:PMID
1087:PMID
1071:Cell
1052:PMID
1003:PMID
951:PMID
902:PMID
884:Cell
861:PMID
843:Cell
812:PMID
761:PMID
718:PMID
683:PMID
634:PMID
585:PMID
528:PMID
512:Cell
493:PMID
431:PMID
312:U2AF
122:and
105:and
92:and
1884:CFI
1874:PAP
1756:doi
1717:doi
1680:PMC
1670:doi
1658:102
1621:PMC
1613:doi
1609:122
1572:PMC
1562:doi
1558:105
1515:doi
1476:doi
1433:doi
1421:365
1388:PMC
1380:doi
1328:PMC
1320:RNA
1287:doi
1212:doi
1175:doi
1134:PMC
1126:doi
1079:doi
1042:PMC
1034:doi
993:PMC
985:doi
981:629
941:PMC
933:doi
892:doi
888:101
851:doi
847:169
804:doi
792:349
753:doi
710:doi
706:351
673:PMC
665:doi
657:RNA
624:PMC
616:doi
608:RNA
575:PMC
567:doi
520:doi
483:PMC
473:doi
421:PMC
413:doi
366:U12
362:U11
158:ibo
31:of
2124::
1952:U1
1762:.
1752:25
1750:.
1733:.
1723:.
1688:.
1678:.
1668:.
1656:.
1652:.
1629:.
1619:.
1607:.
1603:.
1580:.
1570:.
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1552:.
1529:.
1521:.
1509:.
1482:.
1472:75
1470:.
1447:.
1439:.
1431:.
1419:.
1396:.
1386:.
1376:16
1374:.
1370:.
1344:.
1336:.
1322:.
1318:.
1295:.
1283:92
1281:.
1277:.
1265:^
1226:.
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1169:.
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1142:.
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1073:.
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1040:.
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1028:.
1024:.
1001:.
991:.
979:.
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963:^
949:.
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900:.
886:.
882:.
859:.
845:.
841:.
818:.
810:.
802:.
790:.
767:.
759:.
749:15
747:.
724:.
716:.
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681:.
671:.
661:18
659:.
655:.
632:.
622:.
612:15
610:.
606:.
583:.
573:.
563:31
561:.
557:.
534:.
526:.
516:11
514:.
491:.
481:.
471:.
461:74
459:.
455:.
443:^
429:.
419:.
407:.
403:.
391:^
368:,
364:,
184:U6
180:U5
178:,
176:U4
174:,
172:U2
170:,
168:U1
107:3'
103:5'
19:A
1821:e
1814:t
1807:v
1770:.
1758::
1741:.
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415::
409:3
164:p
160:n
156:r
152:n
148:s
146:(
39:(
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