789:. Alternative splicing can occur in many ways. Exons can be extended or skipped, or introns can be retained. It is estimated that 95% of transcripts from multiexon genes undergo alternative splicing, some instances of which occur in a tissue-specific manner and/or under specific cellular conditions. Development of high throughput mRNA sequencing technology can help quantify the expression levels of alternatively spliced isoforms. Differential expression levels across tissues and cell lineages allowed computational approaches to be developed to predict the functions of these isoforms. Given this complexity, alternative splicing of pre-mRNA transcripts is regulated by a system of trans-acting proteins (activators and repressors) that bind to cis-acting sites or "elements" (enhancers and silencers) on the pre-mRNA transcript itself. These proteins and their respective binding elements promote or reduce the usage of a particular splice site. The binding specificity comes from the sequence and structure of the cis-elements, e.g. in HIV-1 there are many donor and acceptor splice sites. Among the various splice sites, ssA7, which is 3' acceptor site, folds into three stem loop structures, i.e. Intronic splicing silencer (ISS), Exonic splicing enhancer (ESE), and Exonic splicing silencer (ESSE3). Solution structure of Intronic splicing silencer and its interaction to host protein hnRNPA1 give insight into specific recognition. However, adding to the complexity of alternative splicing, it is noted that the effects of regulatory factors are many times position-dependent. For example, a splicing factor that serves as a splicing activator when bound to an intronic enhancer element may serve as a repressor when bound to its splicing element in the context of an exon, and vice versa. In addition to the position-dependent effects of enhancer and silencer elements, the location of the branchpoint (i.e., distance upstream of the nearest 3' acceptor site) also affects splicing. The secondary structure of the pre-mRNA transcript also plays a role in regulating splicing, such as by bringing together splicing elements or by masking a sequence that would otherwise serve as a binding element for a splicing factor.
745:
121:
202:) is: G-G--G-U-R-A-G-U (donor site) ... intron sequence ... Y-U-R-A-C (branch sequence 20-50 nucleotides upstream of acceptor site) ... Y-rich-N-C-A-G--G (acceptor site). However, it is noted that the specific sequence of intronic splicing elements and the number of nucleotides between the branchpoint and the nearest 3' acceptor site affect splice site selection. Also, point mutations in the underlying DNA or errors during transcription can activate a
222:
954:
The existence of backsplicing was first suggested in 2012. This backsplicing explains the genesis of circular RNAs resulting from the exact junction between the 3' boundary of an exon with the 5' boundary of an exon located upstream. In these exonic circular RNAs, the junction is a classic 3'-5'link.
801:
are regions with high concentrations of splicing factors. These speckles were once thought to be mere storage centers for splicing factors. However, it is now understood that nuclear speckles help concentrate splicing factors near genes that are physically located close to them. Genes located farther
185:
Within introns, a donor site (5' end of the intron), a branch site (near the 3' end of the intron) and an acceptor site (3' end of the intron) are required for splicing. The splice donor site includes an almost invariant sequence GU at the 5' end of the intron, within a larger, less highly conserved
916:
Allelic differences in mRNA splicing are likely to be a common and important source of phenotypic diversity at the molecular level, in addition to their contribution to genetic disease susceptibility. Indeed, genome-wide studies in humans have identified a range of genes that are subject to
958:
The exclusion of intronic sequences during splicing can also leave traces, in the form of circular RNAs. In some cases, the intronic lariat is not destroyed and the circular part remains as a lariat-derived circRNA.In these lariat-derived circular RNAs, the junction is a 2'-5'link.
638:, cleaves pre-tRNA at two sites in the acceptor loop to form a 5'-half tRNA, terminating at a 2',3'-cyclic phosphodiester group, and a 3'-half tRNA, terminating at a 5'-hydroxyl group, along with a discarded intron. Yeast tRNA kinase then phosphorylates the 5'-hydroxyl group using
554:. Group I and II introns perform splicing similar to the spliceosome without requiring any protein. This similarity suggests that Group I and II introns may be evolutionarily related to the spliceosome. Self-splicing may also be very ancient, and may have existed in an
802:
from speckles can still be transcribed and spliced, but their splicing is less efficient compared to those closer to speckles. Cells can vary their genomic positions of genes relative to nuclear speckles as a mechanism to modulate the expression of genes via splicing.
266:). Assembly and activity of the spliceosome occurs during transcription of the pre-mRNA. The RNA components of snRNPs interact with the intron and are involved in catalysis. Two types of spliceosomes have been identified (major and minor) which contain different
166:. The term intron refers to both the DNA sequence within a gene and the corresponding sequence in the unprocessed RNA transcript. As part of the RNA processing pathway, introns are removed by RNA splicing either shortly after or concurrent with
477:
transcripts. However, in some cases, especially in mRNAs with very long introns, splicing happens in steps, with part of an intron removed and then the remaining intron is spliced out in a following step. This has been found first in the
771:. Second, the 3'OH of the released 5' exon then performs a nucleophilic attack at the first nucleotide following the last nucleotide of the intron at the 3' splice site, thus joining the exons and releasing the intron lariat.
397:
U4 is released, U6/U2 catalyzes transesterification, making the 5'-end of the intron ligate to the A on intron and form a lariat, U5 binds exon at 3' splice site, and the 5' site is cleaved, resulting in the formation of the
681:
Because spliceosomal introns are not conserved in all species, there is debate concerning when spliceosomal splicing evolved. Two models have been proposed: the intron late and intron early models (see
412:
U2/U5/U6 remain bound to the lariat, and the 3' site is cleaved and exons are ligated using ATP hydrolysis. The spliced RNA is released, the lariat is released and degraded, and the snRNPs are recycled.
606:(also tRNA-like) splicing is another rare form of splicing that usually occurs in tRNA. The splicing reaction involves a different biochemistry than the spliceosomal and self-splicing pathways.
678:, on the other hand, splice rarely and mostly non-coding RNAs. Another important difference between these two groups of organisms is that prokaryotes completely lack the spliceosomal pathway.
797:
The location of pre-mRNA splicing is throughout the nucleus, and once mature mRNA is generated, it is transported to the cytoplasm for translation. In both plant and animal cells,
444:
is very similar to the major spliceosome, but instead it splices out rare introns with different splice site sequences. While the minor and major spliceosomes contain the same U5
186:
region. The splice acceptor site at the 3' end of the intron terminates the intron with an almost invariant AG sequence. Upstream (5'-ward) from the AG there is a region high in
898:
Mutation of a splice site reducing specificity. May result in variation in the splice location, causing insertion or deletion of amino acids, or most likely, a disruption of the
3424:
2314:
Pan Q, Shai O, Lee LJ, Frey BJ, Blencowe BJ (December 2008). "Deep surveying of alternative splicing complexity in the human transcriptome by high-throughput sequencing".
920:
In plants, variation for flooding stress tolerance correlated with stress-induced alternative splicing of transcripts associated with gluconeogenesis and other processes.
872:
The use of antisense oligonucleotides to modulate splicing has shown great promise as a therapeutic strategy for a variety of genetic diseases caused by splicing defects.
3759:
3295:
Salzman J, Gawad C, Wang PL, et al. Circular RNAs are the predominant transcript isoform from hundreds of human genes in diverse cell types. PLoS One 2012;7(2):e30733.
1245:
934:
In addition to RNA, proteins can undergo splicing. Although the biomolecular mechanisms are different, the principle is the same: parts of the protein, called
875:
Recent studies have shown that RNA splicing can be regulated by a variety of epigenetic modifications, including DNA methylation and histone modifications.
785:
In many cases, the splicing process can create a range of unique proteins by varying the exon composition of the same mRNA. This phenomenon is then called
2467:"Solution Structure of the HIV-1 Intron Splicing Silencer and Its Interactions with the UP1 Domain of Heterogeneous Nuclear Ribonucleoprotein (hnRNP) A1"
1142:"Deep sequencing of subcellular RNA fractions shows splicing to be predominantly co-transcriptional in the human genome but inefficient for lncRNAs"
759:
Spliceosomal and self-splicing transesterification reactions occur via two sequential transesterification reactions. First, the 2'OH of a specific
3803:
3752:
170:. Introns are found in the genes of most organisms and many viruses. They can be located in a wide range of genes, including those that generate
646:
group to the 5' end of the 3'-half and joins the two halves together. NAD-dependent 2'-phosphotransferase then removes the 2'-phosphate group.
3104:"Abnormally spliced beta-globin mRNAs: a single point mutation generates transcripts sensitive and insensitive to nonsense-mediated mRNA decay"
642:. Yeast tRNA cyclic phosphodiesterase cleaves the cyclic phosphodiester group to form a 2'-phosphorylated 3' end. Yeast tRNA ligase adds an
194:. Further upstream from the polypyrimidine tract is the branchpoint, which includes an adenine nucleotide involved in lariat formation. The
3202:"Transcriptomes of Eight Arabidopsis thaliana Accessions Reveal Core Conserved, Genotype- and Organ-Specific Responses to Flooding Stress"
3745:
2017:"The yeast tRNA splicing endonuclease: a tetrameric enzyme with two active site subunits homologous to the archaeal tRNA endonucleases"
3966:
3961:
3372:
3304:
Jeck WR, Sorrentino JA, Wang K, et al. Circular RNAs are abundant, conserved, and associated with ALU repeats. RNA 2013;19(2):141-57.
3971:
3946:
1496:
756:
reactions that occur between RNA nucleotides. tRNA splicing, however, is an exception and does not occur by transesterification.
1695:"Increased noncanonical splicing of autoantigen transcripts provides the structural basis for expression of untolerized epitopes"
905:
Displacement of a splice site, leading to inclusion or exclusion of more RNA than expected, resulting in longer or shorter exons.
302:
429:, which accounts for more than 99% of splicing. By contrast, when the intronic flanking sequences do not follow the GU-AG rule,
4044:
829:, localization, expression and activity. Furthermore, DNA damage often disrupts splicing by interfering with its coupling to
448:, the minor spliceosome has different but functionally analogous snRNPs for U1, U2, U4, and U6, which are respectively called
3540:
2629:"Next-generation SELEX identifies sequence and structural determinants of splicing factor binding in human pre-mRNA sequence"
869:
to snRNP binding sites, to the branchpoint nucleotide that closes the lariat, or to splice-regulatory element binding sites.
133:
Several methods of RNA splicing occur in nature; the type of splicing depends on the structure of the spliced intron and the
2095:
1233:
1318:
942:
instead of exons, are fused together. Protein splicing has been observed in a wide range of organisms, including bacteria,
594:
The 3'OH of the 5' exon triggers the second transesterification at the 3' splice site, thereby joining the exons together.
313:
are required for the assembly of the spliceosome. The spliceosome forms different complexes during the splicing process:
17:
3322:
Talhouarne GJ and Gall JG. Lariat intronic RNAs in the cytoplasm of
Xenopus tropicalis oocytes. RNA 2014;20(9):1476-87.
1611:"Kinetic role for mammalian SF1/BBP in spliceosome assembly and function after polypyrimidine tract recognition by U2AF"
3813:
3404:
114:
3768:
3512:
2167:"Circularly permuted tRNA genes: their expression and implications for their physiological relevance and development"
993:
576:
3'OH of the 5' exon becomes a nucleophile and the second transesterification results in the joining of the two exons.
2521:"Using positional distribution to identify splicing elements and predict pre-mRNA processing defects in human genes"
2118:"Identification of a human endonuclease complex reveals a link between tRNA splicing and pre-mRNA 3' end formation"
822:
3666:
3610:
3251:"Novel biochemistry: post-translational protein splicing and other lessons from the school of antigen processing"
2928:
Morcos PA (June 2007). "Achieving targeted and quantifiable alteration of mRNA splicing with
Morpholino oligos".
2832:"Inhibition of zebrafish fgf8 pre-mRNA splicing with morpholino oligos: a quantifiable method for gene knockdown"
826:
573:(or one located in the intron) or a nucleotide cofactor (GMP, GDP, GTP) attacks phosphate at the 5' splice site.
105:
that can catalyze their own excision from their parent RNA molecule. The process of transcription, splicing and
3605:
2359:"Systematically differentiating functions for alternatively spliced isoforms through integrating RNA-seq data"
833:. DNA damage also has an impact on the splicing and alternative splicing of genes intimately associated with
3693:
3624:
3365:
536:, performing the functions of the spliceosome by RNA alone. There are three kinds of self-splicing introns,
461:
457:
3429:
3313:
Zhang Y, Zhang XO, Chen T, et al. Circular intronic long noncoding RNAs. Molecular cell 2013;51(6):792-806.
1968:"Homologous SV40 RNA trans-splicing: a new mechanism for diversification of viral sequences and phenotypes"
1650:
Cheng Z, Menees TM (December 2011). "RNA splicing and debranching viewed through analysis of RNA lariats".
910:
3808:
2965:"Correction of aberrant FGFR1 alternative RNA splicing through targeting of intronic regulatory elements"
1189:
Roy SW, Gilbert W (March 2006). "The evolution of spliceosomal introns: patterns, puzzles and progress".
891:
Mutation of a splice site resulting in loss of function of that site. Results in exposure of a premature
278:
splices introns containing GU at the 5' splice site and AG at the 3' splice site. It is composed of the
1517:
Matlin AJ, Clark F, Smith CW (May 2005). "Understanding alternative splicing: towards a cellular code".
3644:
662:
of life, however, the extent and types of splicing can be very different between the major divisions.
453:
2727:"LEDGF/p75 interacts with mRNA splicing factors and targets HIV-1 integration to highly spliced genes"
1140:
Tilgner H, Knowles DG, Johnson R, Davis CA, Chakrabortty S, Djebali S, et al. (September 2012).
752:
Spliceosomal splicing and self-splicing involve a two-step biochemical process. Both steps involve
369:
The U5/U4/U6 snRNP trimer binds, and the U5 snRNP binds exons at the 5' site, with U6 binding to U2;
89:. For many eukaryotic introns, splicing occurs in a series of reactions which are catalyzed by the
4072:
3688:
3476:
3389:
3358:
3344:
1322:
614:
538:
167:
82:
683:
4014:
3835:
3649:
3466:
3451:
830:
643:
490:
78:
46:
38:
3335:
383:
The U1 snRNP is released, U5 shifts from exon to intron, and the U6 binds at the 5' splice site;
3654:
3579:
3471:
639:
588:
The 2'OH of a specific adenosine in the intron attacks the 5' splice site, thereby forming the
1742:
Patel AA, Steitz JA (December 2003). "Splicing double: insights from the second spliceosome".
4067:
3569:
3554:
3434:
1287:
978:
909:
Although many splicing errors are safeguarded by a cellular quality control mechanism termed
199:
106:
86:
3200:
van Veen H, Vashisht D, Akman M, Girke T, Mustroph A, Reinen E, et al. (October 2016).
4077:
3898:
3676:
3574:
3492:
2532:
2370:
2270:
1912:
1855:
1798:
1411:
1094:
1083:"Sequence of a mouse germ-line gene for a variable region of an immunoglobulin light chain"
1037:
866:
786:
780:
449:
207:
191:
98:
50:
8:
3891:
3502:
1003:
764:
753:
581:
562:
295:
291:
287:
283:
279:
3340:
2879:
Sazani P, Kang SH, Maier MA, Wei C, Dillman J, Summerton J, et al. (October 2001).
2702:
2677:
2627:
Reid DC, Chang BL, Gunderson SI, Alpert L, Thompson WA, Fairbrother WG (December 2009).
2536:
2374:
2274:
1916:
1859:
1802:
1415:
1098:
1041:
355:
The U2 snRNP displaces SF1 and binds to the branch point sequence and ATP is hydrolyzed;
85:
that contain introns, splicing is usually needed to create an mRNA molecule that can be
3637:
3520:
3278:
3226:
3201:
3177:
3152:
3133:
3079:
3054:
3030:
3005:
2861:
2807:
2780:
2753:
2726:
2653:
2628:
2604:
2579:
2555:
2520:
2493:
2466:
2442:
2417:
2393:
2358:
2339:
2291:
2258:
2193:
2166:
2147:
2046:
1992:
1967:
1943:
1930:
1900:
1876:
1843:
1819:
1786:
1785:
Sibley CR, Emmett W, Blazquez L, Faro A, Haberman N, Briese M, et al. (May 2015).
1767:
1719:
1694:
1675:
1586:
1561:
1542:
1483:
1458:
1434:
1399:
1375:
1350:
1299:
1214:
1166:
1141:
1063:
837:. For instance, DNA damages modulate the alternative splicing of the DNA repair genes
195:
2257:
Fica SM, Tuttle N, Novak T, Li NS, Lu J, Koodathingal P, et al. (November 2013).
2134:
2117:
2082:
2033:
2016:
1117:
1082:
3886:
3270:
3231:
3182:
3125:
3084:
3035:
2986:
2945:
2910:
2905:
2880:
2853:
2812:
2758:
2725:
Singh PK, Plumb MR, Ferris AL, Iben JR, Wu X, Fadel HJ, et al. (November 2015).
2707:
2658:
2609:
2560:
2498:
2447:
2398:
2331:
2296:
2239:
2198:
2139:
2087:
2038:
1997:
1948:
1881:
1824:
1759:
1724:
1667:
1632:
1591:
1534:
1488:
1439:
1380:
1291:
1206:
1171:
1122:
1055:
884:
655:
521:
or between an endogenous and an exogenous (such as from viruses) or artificial RNAs.
440:
34:
3737:
3721:
3282:
3137:
3102:
Danckwardt S, Neu-Yilik G, Thermann R, Frede U, Hentze MW, Kulozik AE (March 2002).
2881:"Nuclear antisense effects of neutral, anionic and cationic oligonucleotide analogs"
2865:
2418:"The emerging era of genomic data integration for analyzing splice isoform function"
2151:
2050:
1771:
1546:
1303:
1218:
3262:
3221:
3213:
3172:
3164:
3115:
3074:
3066:
3025:
3017:
2976:
2937:
2900:
2892:
2843:
2802:
2792:
2748:
2738:
2697:
2689:
2648:
2640:
2599:
2591:
2550:
2540:
2488:
2478:
2437:
2429:
2388:
2378:
2343:
2323:
2286:
2278:
2229:
2188:
2178:
2129:
2077:
2028:
1987:
1979:
1938:
1920:
1871:
1863:
1842:
Duff MO, Olson S, Wei X, Garrett SC, Osman A, Bolisetty M, et al. (May 2015).
1814:
1806:
1751:
1714:
1706:
1659:
1622:
1581:
1573:
1526:
1478:
1470:
1429:
1419:
1370:
1362:
1283:
1198:
1161:
1153:
1112:
1102:
1067:
1045:
929:
857:
Splicing events can be experimentally altered by binding steric-blocking antisense
798:
659:
550:
1679:
1400:"Genome-wide association between branch point properties and alternative splicing"
3845:
3828:
3823:
3798:
3698:
3535:
3381:
2383:
2065:
1901:"Cis- and trans-splicing of mRNAs mediated by tRNA sequences in eukaryotic cells"
1424:
1332:
988:
858:
811:
544:
262:, a large RNA-protein complex composed of five small nuclear ribonucleoproteins (
110:
2015:
Trotta CR, Miao F, Arn EA, Stevens SW, Ho CK, Rauhut R, Abelson JN (June 1997).
1844:"Genome-wide identification of zero nucleotide recursive splicing in Drosophila"
883:
It has been suggested that one third of all disease-causing mutations impact on
4009:
3908:
3793:
3788:
3525:
3497:
2941:
2693:
2595:
2525:
Proceedings of the
National Academy of Sciences of the United States of America
1905:
Proceedings of the
National Academy of Sciences of the United States of America
1710:
1328:
1087:
Proceedings of the
National Academy of Sciences of the United States of America
983:
671:
514:
506:
211:
3266:
2433:
1693:
Ng B, Yang F, Huston DP, Yan Y, Yang Y, Xiong Z, et al. (December 2004).
1663:
1474:
763:
nucleotide within the intron, defined during spliceosome assembly, performs a
4061:
3726:
3413:
3120:
3103:
2896:
2678:"Genome organization around nuclear speckles drives mRNA splicing efficiency"
2234:
2217:
2183:
913:(NMD), a number of splicing-related diseases also exist, as suggested above.
899:
480:
474:
175:
70:
54:
2545:
2483:
1925:
1271:
3776:
3600:
3399:
3274:
3235:
3186:
3129:
3088:
3055:"Context-dependent control of alternative splicing by RNA-binding proteins"
3039:
3006:"Context-dependent control of alternative splicing by RNA-binding proteins"
2990:
2949:
2914:
2857:
2816:
2762:
2743:
2711:
2662:
2613:
2564:
2502:
2451:
2402:
2335:
2300:
2202:
2143:
2001:
1952:
1885:
1828:
1763:
1728:
1671:
1636:
1627:
1610:
1595:
1538:
1492:
1443:
1398:
Corvelo A, Hallegger M, Smith CW, Eyras E (November 2010). Meyer IM (ed.).
1384:
1351:"Large-scale mapping of branchpoints in human pre-mRNA transcripts in vivo"
1295:
1210:
1175:
1107:
805:
619:
214:, which might otherwise affect only a single amino acid, can manifest as a
179:
74:
2243:
2091:
2042:
1349:
Taggart AJ, DeSimone AM, Shih JS, Filloux ME, Fairbrother WG (June 2012).
1157:
301:
and is active in the nucleus. In addition, a number of proteins including
120:
4034:
3881:
3840:
3703:
3632:
2981:
2964:
1126:
1059:
998:
767:
on the first nucleotide of the intron at the 5' splice site, forming the
323:
The U1 snRNP binds to the GU sequence at the 5' splice site of an intron;
259:
206:
in part of the transcript that usually is not spliced. This results in a
90:
3217:
2797:
2644:
2519:
Lim KH, Ferraris L, Filloux ME, Raphael BJ, Fairbrother WG (July 2011).
2357:
Eksi R, Li HD, Menon R, Wen Y, Omenn GS, Kretzler M, Guan Y (Nov 2013).
2282:
1934:
1867:
1810:
744:
3661:
892:
862:
834:
675:
570:
473:
In most cases, splicing removes introns as single units from precursor
326:
310:
187:
3168:
2848:
2831:
1983:
1366:
1050:
1025:
663:
555:
221:
3250:
3070:
3021:
1755:
1577:
1530:
1202:
3681:
3671:
3595:
2327:
1081:
Tonegawa S, Maxam AM, Tizard R, Bernard O, Gilbert W (March 1978).
973:
533:
518:
513:, and joins two exons that are not within the same RNA transcript.
230:
171:
162:, that is, a segment of DNA that is located between two exons of a
134:
102:
3350:
2116:
Paushkin SV, Patel M, Furia BS, Peltz SW, Trotta CR (April 2004).
1898:
4001:
3981:
3976:
3417:
2465:
Jain N, Morgan CE, Rife BD, Salemi M, Tolbert BS (January 2016).
943:
565:
characterize the mechanism in which group I introns are spliced:
3101:
3956:
3951:
3931:
3867:
3818:
939:
935:
635:
631:
623:
510:
58:
1459:"The role of U2AF35 and U2AF65 in enhancer-dependent splicing"
4024:
4019:
3941:
3936:
3926:
3921:
3916:
3876:
3439:
938:
instead of introns, are removed. The remaining parts, called
845:
839:
627:
610:
445:
338:
332:
306:
298:
267:
263:
215:
94:
4039:
4029:
3871:
968:
792:
667:
603:
163:
66:
42:
3199:
1899:
Di Segni G, Gastaldi S, Tocchini-Valentini GP (May 2008).
1397:
1348:
1139:
1080:
748:
Diagram illustrating the two-step biochemistry of splicing
3409:
2518:
580:
The mechanism in which group II introns are spliced (two
2626:
1784:
852:
806:
Role of splicing/alternative splicing in HIV-integration
2580:"Role of RNA structure in regulating pre-mRNA splicing"
2115:
2063:
498:
genes, but cases in humans have been reported as well.
2066:"Structure and function of the yeast tRNA ligase gene"
1456:
1272:"Mechanisms of alternative pre-messenger RNA splicing"
3767:
2724:
2464:
2014:
2930:
1841:
949:
2878:
2829:
2064:Westaway SK, Phizicky EM, Abelson J (March 1988).
817:
210:with a missing section of an exon. In this way, a
2676:Bhat P, Chow A, Emert B, et al. (May 2024).
2256:
433:is said to occur (see "minor spliceosome" below).
93:, a complex of small nuclear ribonucleoproteins (
4059:
3336:Virtual Cell Animation Collection: mRNA Splicing
2416:Li HD, Menon R, Omenn GS, Guan Y (August 2014).
2415:
1516:
1457:Graveley BR, Hertel KJ, Maniatis T (June 2001).
878:
2774:
2772:
2215:
1692:
2356:
2313:
1699:The Journal of Allergy and Clinical Immunology
1234:"RNA Splicing: Introns, Exons and Spliceosome"
584:reaction like group I introns) is as follows:
509:is a form of splicing that removes introns or
3753:
3366:
2830:Draper BW, Morcos PA, Kimmel CB (July 2001).
2778:
2675:
1787:"Recursive splicing in long vertebrate genes"
1608:
895:, loss of an exon, or inclusion of an intron.
2962:
2769:
2718:
2577:
2109:
2057:
1450:
2209:
1741:
1649:
825:affects splicing factors by altering their
814:, as HIV-1 targets highly spliced genes.
517:can occur between two different endogenous
3760:
3746:
3521:Precursor mRNA (pre-mRNA / hnRNA)
3373:
3359:
3248:
3150:
2008:
1559:
1553:
1188:
335:binds at the 3' splice site of the intron;
329:binds to the intron branch point sequence;
3343:at the U.S. National Library of Medicine
3225:
3176:
3119:
3078:
3029:
2980:
2963:Bruno IG, Jin W, Cote GJ (October 2004).
2904:
2847:
2806:
2796:
2781:"The RNA Splicing Response to DNA Damage"
2752:
2742:
2701:
2652:
2603:
2554:
2544:
2492:
2482:
2441:
2392:
2382:
2290:
2259:"RNA catalyses nuclear pre-mRNA splicing"
2233:
2192:
2182:
2133:
2081:
2032:
1991:
1965:
1942:
1924:
1875:
1818:
1718:
1643:
1626:
1585:
1482:
1433:
1423:
1374:
1355:Nature Structural & Molecular Biology
1344:
1342:
1165:
1116:
1106:
1049:
253:
2250:
2158:
1602:
1288:10.1146/annurev.biochem.72.121801.161720
793:Role of nuclear speckles in RNA splicing
743:
739:
220:
119:
2216:Abelson J, Trotta CR, Li H (May 1998).
1265:
1263:
1023:
810:The process of splicing is linked with
774:
409:Complex C* (post-spliceosomal complex)
303:U2 small nuclear RNA auxiliary factor 1
140:
14:
4060:
3052:
3003:
2927:
2514:
2512:
1744:Nature Reviews. Molecular Cell Biology
1566:Nature Reviews. Molecular Cell Biology
1562:"A day in the life of the spliceosome"
1519:Nature Reviews. Molecular Cell Biology
1510:
1339:
1231:
366:Complex B (pre-catalytic spliceosome)
3741:
3541:Histone acetylation and deacetylation
3354:
1609:Guth S, Valcárcel J (December 2000).
1269:
853:Experimental manipulation of splicing
468:
3606:Ribosome-nascent chain complex (RNC)
2779:Shkreta L, Chabot B (October 2015).
2350:
2164:
1260:
532:occurs for rare introns that form a
218:or truncation in the final protein.
128:
3380:
3151:Ward AJ, Cooper TA (January 2010).
2509:
2471:The Journal of Biological Chemistry
2222:The Journal of Biological Chemistry
2070:The Journal of Biological Chemistry
1615:The Journal of Biological Chemistry
1560:Matera AG, Wang Z (February 2014).
923:
77:either during or immediately after
24:
1310:
1248:from the original on 15 March 2011
394:Complex C (catalytic spliceosome)
341:binds to the polypyrimidine tract;
115:central dogma of molecular biology
25:
4089:
3769:Post-transcriptional modification
3329:
2578:Warf MB, Berglund JA (May 2024).
1966:Eul J, Patzel V (November 2013).
994:Post-transcriptional modification
501:
950:Splicing and genesis of circRNAs
598:
524:
421:This type of splicing is termed
137:required for splicing to occur.
61:(non-coding regions of RNA) and
57:). It works by removing all the
3611:Post-translational modification
3316:
3307:
3298:
3289:
3249:Hanada K, Yang JC (June 2005).
3242:
3193:
3144:
3095:
3046:
2997:
2956:
2921:
2872:
2823:
2669:
2620:
2571:
2458:
2409:
2307:
2098:from the original on 2018-11-18
1959:
1892:
1835:
1778:
1735:
1686:
1652:Molecular Genetics and Genomics
1499:from the original on 2018-11-20
827:post-translational modification
818:Splicing response to DNA damage
3153:"The pathobiology of splicing"
3053:Fu XD, Ares M (October 2014).
3004:Fu XD, Ares M (October 2014).
2584:Trends in Biochemical Sciences
1391:
1225:
1182:
1133:
1074:
1017:
13:
1:
3255:Journal of Molecular Medicine
2135:10.1016/S0092-8674(04)00342-3
2083:10.1016/S0021-9258(18)69050-7
2034:10.1016/S0092-8674(00)80270-6
1324:2012 Journal Citation Reports
1319:Molecular Biology of the Cell
1276:Annual Review of Biochemistry
1010:
879:Splicing errors and variation
258:Splicing is catalyzed by the
2384:10.1371/journal.pcbi.1003314
1425:10.1371/journal.pcbi.1001016
946:, plants, yeast and humans.
911:nonsense-mediated mRNA decay
732:
729:
721:
718:
710:
707:
649:
622:heterotetramer, composed of
352:Complex A (pre-spliceosome)
27:Process in molecular biology
7:
1024:Gilbert W (February 1978).
962:
666:splice many protein-coding
654:Splicing occurs in all the
10:
4094:
2942:10.1016/j.bbrc.2007.04.172
2694:10.1038/s41586-024-07429-6
2596:10.1016/j.tibs.2009.10.004
2363:PLOS Computational Biology
1711:10.1016/j.jaci.2004.09.006
1404:PLOS Computational Biology
927:
917:allele-specific splicing.
778:
154:is derived from the terms
145:
4000:
3907:
3863:
3854:
3784:
3775:
3714:
3623:
3588:
3562:
3553:
3511:
3485:
3459:
3450:
3388:
3267:10.1007/s00109-005-0652-6
2434:10.1016/j.tig.2014.05.005
1664:10.1007/s00438-011-0635-y
1475:10.1017/s1355838201010317
887:. Common errors include:
488:) gene of the fruit fly,
73:, splicing occurs in the
3672:sequestration (P-bodies)
3345:Medical Subject Headings
3157:The Journal of Pathology
3121:10.1182/blood.V99.5.1811
3059:Nature Reviews. Genetics
3010:Nature Reviews. Genetics
2969:Human Molecular Genetics
2235:10.1074/jbc.273.21.12685
2184:10.3389/fgene.2014.00063
1191:Nature Reviews. Genetics
618:, a yeast tRNA splicing
615:Saccharomyces cerevisiae
558:present before protein.
241:- Poly pyrimidine Tract
198:for an intron (in IUPAC
3836:Poly(A)-binding protein
3650:Gene regulatory network
2731:Genes & Development
2546:10.1073/pnas.1101135108
2484:10.1074/jbc.M115.674564
2165:Soma A (1 April 2014).
1926:10.1073/pnas.0800420105
644:adenosine monophosphate
569:3'OH of a free guanine
491:Drosophila melanogaster
124:Process of RNA splicing
87:translated into protein
39:precursor messenger RNA
3655:cis-regulatory element
2897:10.1093/nar/29.19.3965
2885:Nucleic Acids Research
2744:10.1101/gad.267609.115
1628:10.1074/jbc.M001483200
1270:Black DL (June 2003).
1108:10.1073/pnas.75.3.1485
1026:"Why genes in pieces?"
1006:, a splicing regulator
749:
640:adenosine triphosphate
254:Formation and activity
250:
125:
69:(coding regions). For
49:is transformed into a
2171:Frontiers in Genetics
1158:10.1101/gr.134445.111
979:Exon junction complex
867:Peptide nucleic acids
747:
740:Biochemical mechanism
431:noncanonical splicing
224:
200:nucleic acid notation
123:
99:self-splicing introns
71:nuclear-encoded genes
3899:Alternative splicing
3677:alternative splicing
3667:Post-transcriptional
3493:Transcription factor
1331:(Science ed.).
787:alternative splicing
781:Alternative splicing
775:Alternative splicing
563:transesterifications
226:Intron Exon Boundary
208:mature messenger RNA
192:polypyrimidine tract
141:Spliceosomal complex
51:mature messenger RNA
3601:Transfer RNA (tRNA)
3218:10.1104/pp.16.00472
2798:10.3390/biom5042935
2645:10.1261/rna.1821809
2537:2011PNAS..10811093H
2531:(27): 11093–11098.
2375:2013PLSCB...9E3314E
2283:10.1038/nature12734
2275:2013Natur.503..229F
2228:(21): 12685–12688.
1917:2008PNAS..105.6864D
1868:10.1038/nature14475
1860:2015Natur.521..376D
1811:10.1038/nature14466
1803:2015Natur.521..371S
1621:(48): 38059–38066.
1416:2010PLSCB...6E1016C
1099:1978PNAS...75.1485T
1042:1978Natur.271..501G
1004:SWAP protein domain
769:lariat intermediate
765:nucleophilic attack
754:transesterification
691:
690:Splicing diversity
582:transesterification
204:cryptic splice site
37:where a newly-made
18:Splicing (genetics)
4010:5′ cap methylation
3715:Influential people
3694:Post-translational
3513:Post-transcription
2982:10.1093/hmg/ddh272
2422:Trends in Genetics
750:
689:
494:, and a few other
469:Recursive splicing
423:canonical splicing
251:
196:consensus sequence
126:
4055:
4054:
3996:
3995:
3992:
3991:
3909:pre-mRNA factors
3735:
3734:
3619:
3618:
3549:
3548:
3425:Special transfers
3169:10.1002/path.2649
2975:(20): 2409–2420.
2891:(19): 3965–3974.
2849:10.1002/gene.1053
2737:(21): 2287–2297.
2639:(12): 2385–2397.
2590:(8014): 169–178.
2322:(12): 1413–1415.
2269:(7475): 229–234.
1984:10.4161/rna.26707
1978:(11): 1689–1699.
1911:(19): 6864–6869.
1854:(7552): 376–379.
1797:(7552): 371–375.
1367:10.1038/nsmb.2327
1232:Clancy S (2008).
737:
736:
441:minor spliceosome
327:Splicing factor 1
276:major spliceosome
237:- 3' Splice site
156:intragenic region
129:Splicing pathways
35:molecular biology
16:(Redirected from
4085:
3861:
3860:
3794:5′ cap formation
3782:
3781:
3762:
3755:
3748:
3739:
3738:
3560:
3559:
3457:
3456:
3375:
3368:
3361:
3352:
3351:
3323:
3320:
3314:
3311:
3305:
3302:
3296:
3293:
3287:
3286:
3246:
3240:
3239:
3229:
3206:Plant Physiology
3197:
3191:
3190:
3180:
3148:
3142:
3141:
3123:
3114:(5): 1811–1816.
3099:
3093:
3092:
3082:
3050:
3044:
3043:
3033:
3001:
2995:
2994:
2984:
2960:
2954:
2953:
2925:
2919:
2918:
2908:
2876:
2870:
2869:
2851:
2827:
2821:
2820:
2810:
2800:
2791:(4): 2935–2977.
2776:
2767:
2766:
2756:
2746:
2722:
2716:
2715:
2705:
2688:(5): 1165–1173.
2673:
2667:
2666:
2656:
2624:
2618:
2617:
2607:
2575:
2569:
2568:
2558:
2548:
2516:
2507:
2506:
2496:
2486:
2477:(5): 2331–2344.
2462:
2456:
2455:
2445:
2413:
2407:
2406:
2396:
2386:
2369:(11): e1003314.
2354:
2348:
2347:
2311:
2305:
2304:
2294:
2254:
2248:
2247:
2237:
2213:
2207:
2206:
2196:
2186:
2162:
2156:
2155:
2137:
2113:
2107:
2106:
2104:
2103:
2085:
2076:(7): 3171–3176.
2061:
2055:
2054:
2036:
2012:
2006:
2005:
1995:
1963:
1957:
1956:
1946:
1928:
1896:
1890:
1889:
1879:
1839:
1833:
1832:
1822:
1782:
1776:
1775:
1739:
1733:
1732:
1722:
1705:(6): 1463–1470.
1690:
1684:
1683:
1658:(5–6): 395–410.
1647:
1641:
1640:
1630:
1606:
1600:
1599:
1589:
1557:
1551:
1550:
1514:
1508:
1507:
1505:
1504:
1486:
1454:
1448:
1447:
1437:
1427:
1410:(11): e1001016.
1395:
1389:
1388:
1378:
1346:
1337:
1336:
1314:
1308:
1307:
1267:
1258:
1257:
1255:
1253:
1238:Nature Education
1229:
1223:
1222:
1186:
1180:
1179:
1169:
1152:(9): 1616–1625.
1137:
1131:
1130:
1120:
1110:
1093:(3): 1485–1489.
1078:
1072:
1071:
1053:
1051:10.1038/271501a0
1021:
930:Protein splicing
924:Protein splicing
799:nuclear speckles
692:
688:
684:intron evolution
249:- 5' splice site
83:eukaryotic genes
33:is a process in
21:
4093:
4092:
4088:
4087:
4086:
4084:
4083:
4082:
4073:Gene expression
4058:
4057:
4056:
4051:
3988:
3903:
3850:
3846:Polyuridylation
3799:Polyadenylation
3771:
3766:
3736:
3731:
3710:
3645:Transcriptional
3615:
3584:
3545:
3536:Polyadenylation
3507:
3481:
3446:
3440:Protein→Protein
3391:
3384:
3382:Gene expression
3379:
3332:
3327:
3326:
3321:
3317:
3312:
3308:
3303:
3299:
3294:
3290:
3247:
3243:
3198:
3194:
3149:
3145:
3100:
3096:
3071:10.1038/nrg3778
3065:(10): 689–701.
3051:
3047:
3022:10.1038/nrg3778
3016:(10): 689–701.
3002:
2998:
2961:
2957:
2926:
2922:
2877:
2873:
2828:
2824:
2777:
2770:
2723:
2719:
2674:
2670:
2625:
2621:
2576:
2572:
2517:
2510:
2463:
2459:
2414:
2410:
2355:
2351:
2316:Nature Genetics
2312:
2308:
2255:
2251:
2218:"tRNA splicing"
2214:
2210:
2163:
2159:
2114:
2110:
2101:
2099:
2062:
2058:
2013:
2009:
1964:
1960:
1897:
1893:
1840:
1836:
1783:
1779:
1756:10.1038/nrm1259
1750:(12): 960–970.
1740:
1736:
1691:
1687:
1648:
1644:
1607:
1603:
1578:10.1038/nrm3742
1558:
1554:
1531:10.1038/nrm1645
1515:
1511:
1502:
1500:
1455:
1451:
1396:
1392:
1347:
1340:
1333:Thomson Reuters
1316:
1315:
1311:
1268:
1261:
1251:
1249:
1230:
1226:
1203:10.1038/nrg1807
1187:
1183:
1146:Genome Research
1138:
1134:
1079:
1075:
1022:
1018:
1013:
989:Polyadenylation
965:
952:
932:
926:
881:
855:
820:
812:HIV integration
808:
795:
783:
777:
742:
672:non-coding RNAs
652:
601:
527:
504:
471:
256:
148:
143:
131:
111:gene expression
97:). There exist
28:
23:
22:
15:
12:
11:
5:
4091:
4081:
4080:
4075:
4070:
4053:
4052:
4050:
4049:
4048:
4047:
4042:
4037:
4032:
4027:
4022:
4015:mRNA decapping
4012:
4006:
4004:
3998:
3997:
3994:
3993:
3990:
3989:
3987:
3986:
3985:
3984:
3979:
3974:
3969:
3964:
3959:
3954:
3949:
3944:
3939:
3934:
3929:
3924:
3913:
3911:
3905:
3904:
3902:
3901:
3896:
3895:
3894:
3889:
3879:
3874:
3864:
3858:
3852:
3851:
3849:
3848:
3843:
3838:
3833:
3832:
3831:
3826:
3821:
3816:
3811:
3806:
3796:
3791:
3789:Precursor mRNA
3785:
3779:
3773:
3772:
3765:
3764:
3757:
3750:
3742:
3733:
3732:
3730:
3729:
3724:
3722:François Jacob
3718:
3716:
3712:
3711:
3709:
3708:
3707:
3706:
3701:
3691:
3686:
3685:
3684:
3679:
3674:
3664:
3659:
3658:
3657:
3652:
3642:
3641:
3640:
3629:
3627:
3621:
3620:
3617:
3616:
3614:
3613:
3608:
3603:
3598:
3592:
3590:
3586:
3585:
3583:
3582:
3577:
3572:
3566:
3564:
3557:
3551:
3550:
3547:
3546:
3544:
3543:
3538:
3533:
3528:
3523:
3517:
3515:
3509:
3508:
3506:
3505:
3500:
3498:RNA polymerase
3495:
3489:
3487:
3483:
3482:
3480:
3479:
3474:
3469:
3463:
3461:
3454:
3448:
3447:
3445:
3444:
3443:
3442:
3437:
3432:
3422:
3421:
3420:
3402:
3396:
3394:
3386:
3385:
3378:
3377:
3370:
3363:
3355:
3349:
3348:
3338:
3331:
3330:External links
3328:
3325:
3324:
3315:
3306:
3297:
3288:
3261:(6): 420–428.
3241:
3212:(2): 668–689.
3192:
3163:(2): 152–163.
3143:
3094:
3045:
2996:
2955:
2936:(2): 521–527.
2920:
2871:
2842:(3): 154–156.
2822:
2768:
2717:
2668:
2619:
2570:
2508:
2457:
2428:(8): 340–347.
2408:
2349:
2328:10.1038/ng.259
2306:
2249:
2208:
2157:
2128:(3): 311–321.
2108:
2056:
2027:(6): 849–858.
2007:
1958:
1891:
1834:
1777:
1734:
1685:
1642:
1601:
1572:(2): 108–121.
1552:
1525:(5): 386–398.
1509:
1469:(6): 806–818.
1449:
1390:
1361:(7): 719–721.
1338:
1329:Web of Science
1309:
1282:(1): 291–336.
1259:
1224:
1197:(3): 211–221.
1181:
1132:
1073:
1015:
1014:
1012:
1009:
1008:
1007:
1001:
996:
991:
986:
981:
976:
971:
964:
961:
951:
948:
928:Main article:
925:
922:
907:
906:
903:
896:
880:
877:
854:
851:
819:
816:
807:
804:
794:
791:
779:Main article:
776:
773:
741:
738:
735:
734:
731:
728:
724:
723:
720:
717:
713:
712:
709:
706:
702:
701:
698:
695:
668:messenger RNAs
651:
648:
600:
597:
596:
595:
592:
578:
577:
574:
526:
523:
515:Trans-splicing
507:Trans-splicing
503:
502:Trans-splicing
500:
470:
467:
466:
465:
435:
434:
427:lariat pathway
425:or termed the
418:
417:
416:
415:
414:
413:
404:
403:
402:
401:
400:
399:
389:
388:
387:
386:
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384:
375:
374:
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371:
370:
361:
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359:
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357:
356:
347:
346:
345:
344:
343:
342:
336:
330:
324:
315:
314:
255:
252:
245:- Branch site
212:point mutation
190:(C and U), or
147:
144:
142:
139:
130:
127:
65:back together
26:
9:
6:
4:
3:
2:
4090:
4079:
4076:
4074:
4071:
4069:
4066:
4065:
4063:
4046:
4043:
4041:
4038:
4036:
4033:
4031:
4028:
4026:
4023:
4021:
4018:
4017:
4016:
4013:
4011:
4008:
4007:
4005:
4003:
3999:
3983:
3980:
3978:
3975:
3973:
3970:
3968:
3965:
3963:
3960:
3958:
3955:
3953:
3950:
3948:
3945:
3943:
3940:
3938:
3935:
3933:
3930:
3928:
3925:
3923:
3920:
3919:
3918:
3915:
3914:
3912:
3910:
3906:
3900:
3897:
3893:
3890:
3888:
3885:
3884:
3883:
3880:
3878:
3875:
3873:
3869:
3866:
3865:
3862:
3859:
3857:
3853:
3847:
3844:
3842:
3839:
3837:
3834:
3830:
3827:
3825:
3822:
3820:
3817:
3815:
3812:
3810:
3807:
3805:
3802:
3801:
3800:
3797:
3795:
3792:
3790:
3787:
3786:
3783:
3780:
3778:
3774:
3770:
3763:
3758:
3756:
3751:
3749:
3744:
3743:
3740:
3728:
3727:Jacques Monod
3725:
3723:
3720:
3719:
3717:
3713:
3705:
3702:
3700:
3697:
3696:
3695:
3692:
3690:
3689:Translational
3687:
3683:
3680:
3678:
3675:
3673:
3670:
3669:
3668:
3665:
3663:
3660:
3656:
3653:
3651:
3648:
3647:
3646:
3643:
3639:
3636:
3635:
3634:
3631:
3630:
3628:
3626:
3622:
3612:
3609:
3607:
3604:
3602:
3599:
3597:
3594:
3593:
3591:
3587:
3581:
3578:
3576:
3573:
3571:
3568:
3567:
3565:
3561:
3558:
3556:
3552:
3542:
3539:
3537:
3534:
3532:
3529:
3527:
3524:
3522:
3519:
3518:
3516:
3514:
3510:
3504:
3501:
3499:
3496:
3494:
3491:
3490:
3488:
3484:
3478:
3475:
3473:
3470:
3468:
3465:
3464:
3462:
3458:
3455:
3453:
3452:Transcription
3449:
3441:
3438:
3436:
3433:
3431:
3428:
3427:
3426:
3423:
3419:
3415:
3411:
3408:
3407:
3406:
3405:Central dogma
3403:
3401:
3398:
3397:
3395:
3393:
3387:
3383:
3376:
3371:
3369:
3364:
3362:
3357:
3356:
3353:
3346:
3342:
3339:
3337:
3334:
3333:
3319:
3310:
3301:
3292:
3284:
3280:
3276:
3272:
3268:
3264:
3260:
3256:
3252:
3245:
3237:
3233:
3228:
3223:
3219:
3215:
3211:
3207:
3203:
3196:
3188:
3184:
3179:
3174:
3170:
3166:
3162:
3158:
3154:
3147:
3139:
3135:
3131:
3127:
3122:
3117:
3113:
3109:
3105:
3098:
3090:
3086:
3081:
3076:
3072:
3068:
3064:
3060:
3056:
3049:
3041:
3037:
3032:
3027:
3023:
3019:
3015:
3011:
3007:
3000:
2992:
2988:
2983:
2978:
2974:
2970:
2966:
2959:
2951:
2947:
2943:
2939:
2935:
2931:
2924:
2916:
2912:
2907:
2902:
2898:
2894:
2890:
2886:
2882:
2875:
2867:
2863:
2859:
2855:
2850:
2845:
2841:
2837:
2833:
2826:
2818:
2814:
2809:
2804:
2799:
2794:
2790:
2786:
2782:
2775:
2773:
2764:
2760:
2755:
2750:
2745:
2740:
2736:
2732:
2728:
2721:
2713:
2709:
2704:
2699:
2695:
2691:
2687:
2683:
2679:
2672:
2664:
2660:
2655:
2650:
2646:
2642:
2638:
2634:
2630:
2623:
2615:
2611:
2606:
2601:
2597:
2593:
2589:
2585:
2581:
2574:
2566:
2562:
2557:
2552:
2547:
2542:
2538:
2534:
2530:
2526:
2522:
2515:
2513:
2504:
2500:
2495:
2490:
2485:
2480:
2476:
2472:
2468:
2461:
2453:
2449:
2444:
2439:
2435:
2431:
2427:
2423:
2419:
2412:
2404:
2400:
2395:
2390:
2385:
2380:
2376:
2372:
2368:
2364:
2360:
2353:
2345:
2341:
2337:
2333:
2329:
2325:
2321:
2317:
2310:
2302:
2298:
2293:
2288:
2284:
2280:
2276:
2272:
2268:
2264:
2260:
2253:
2245:
2241:
2236:
2231:
2227:
2223:
2219:
2212:
2204:
2200:
2195:
2190:
2185:
2180:
2176:
2172:
2168:
2161:
2153:
2149:
2145:
2141:
2136:
2131:
2127:
2123:
2119:
2112:
2097:
2093:
2089:
2084:
2079:
2075:
2071:
2067:
2060:
2052:
2048:
2044:
2040:
2035:
2030:
2026:
2022:
2018:
2011:
2003:
1999:
1994:
1989:
1985:
1981:
1977:
1973:
1969:
1962:
1954:
1950:
1945:
1940:
1936:
1932:
1927:
1922:
1918:
1914:
1910:
1906:
1902:
1895:
1887:
1883:
1878:
1873:
1869:
1865:
1861:
1857:
1853:
1849:
1845:
1838:
1830:
1826:
1821:
1816:
1812:
1808:
1804:
1800:
1796:
1792:
1788:
1781:
1773:
1769:
1765:
1761:
1757:
1753:
1749:
1745:
1738:
1730:
1726:
1721:
1716:
1712:
1708:
1704:
1700:
1696:
1689:
1681:
1677:
1673:
1669:
1665:
1661:
1657:
1653:
1646:
1638:
1634:
1629:
1624:
1620:
1616:
1612:
1605:
1597:
1593:
1588:
1583:
1579:
1575:
1571:
1567:
1563:
1556:
1548:
1544:
1540:
1536:
1532:
1528:
1524:
1520:
1513:
1498:
1494:
1490:
1485:
1480:
1476:
1472:
1468:
1464:
1460:
1453:
1445:
1441:
1436:
1431:
1426:
1421:
1417:
1413:
1409:
1405:
1401:
1394:
1386:
1382:
1377:
1372:
1368:
1364:
1360:
1356:
1352:
1345:
1343:
1334:
1330:
1326:
1325:
1320:
1313:
1305:
1301:
1297:
1293:
1289:
1285:
1281:
1277:
1273:
1266:
1264:
1247:
1243:
1239:
1235:
1228:
1220:
1216:
1212:
1208:
1204:
1200:
1196:
1192:
1185:
1177:
1173:
1168:
1163:
1159:
1155:
1151:
1147:
1143:
1136:
1128:
1124:
1119:
1114:
1109:
1104:
1100:
1096:
1092:
1088:
1084:
1077:
1069:
1065:
1061:
1057:
1052:
1047:
1043:
1039:
1036:(5645): 501.
1035:
1031:
1027:
1020:
1016:
1005:
1002:
1000:
997:
995:
992:
990:
987:
985:
982:
980:
977:
975:
972:
970:
967:
966:
960:
956:
947:
945:
941:
937:
931:
921:
918:
914:
912:
904:
901:
900:reading frame
897:
894:
890:
889:
888:
886:
876:
873:
870:
868:
864:
860:
850:
848:
847:
842:
841:
836:
832:
831:transcription
828:
824:
815:
813:
803:
800:
790:
788:
782:
772:
770:
766:
762:
757:
755:
746:
726:
725:
716:Self-splicing
715:
714:
704:
703:
699:
696:
694:
693:
687:
685:
679:
677:
673:
669:
665:
661:
657:
647:
645:
641:
637:
633:
629:
625:
621:
617:
616:
612:
607:
605:
599:tRNA splicing
593:
591:
587:
586:
585:
583:
575:
572:
568:
567:
566:
564:
559:
557:
553:
552:
547:
546:
541:
540:
535:
531:
530:Self-splicing
525:Self-splicing
522:
520:
516:
512:
508:
499:
497:
493:
492:
487:
483:
482:
481:Ultrabithorax
476:
463:
459:
455:
451:
447:
443:
442:
437:
436:
432:
428:
424:
420:
419:
411:
410:
408:
407:
406:
405:
396:
395:
393:
392:
391:
390:
382:
381:
379:
378:
377:
376:
368:
367:
365:
364:
363:
362:
354:
353:
351:
350:
349:
348:
340:
337:
334:
331:
328:
325:
322:
321:
319:
318:
317:
316:
312:
309:(U2AF65) and
308:
304:
300:
297:
293:
289:
285:
281:
277:
273:
272:
271:
269:
265:
261:
248:
244:
240:
236:
233:
232:
227:
223:
219:
217:
213:
209:
205:
201:
197:
193:
189:
183:
181:
177:
176:ribosomal RNA
173:
169:
168:transcription
165:
161:
157:
153:
138:
136:
122:
118:
116:
112:
108:
104:
100:
96:
92:
88:
84:
80:
79:transcription
76:
72:
68:
64:
60:
56:
52:
48:
44:
40:
36:
32:
19:
4068:RNA splicing
3870: /
3856:RNA splicing
3855:
3704:irreversible
3589:Key elements
3530:
3486:Key elements
3400:Genetic code
3390:Introduction
3341:RNA+Splicing
3318:
3309:
3300:
3291:
3258:
3254:
3244:
3209:
3205:
3195:
3160:
3156:
3146:
3111:
3107:
3097:
3062:
3058:
3048:
3013:
3009:
2999:
2972:
2968:
2958:
2933:
2929:
2923:
2888:
2884:
2874:
2839:
2835:
2825:
2788:
2785:Biomolecules
2784:
2734:
2730:
2720:
2685:
2681:
2671:
2636:
2632:
2622:
2587:
2583:
2573:
2528:
2524:
2474:
2470:
2460:
2425:
2421:
2411:
2366:
2362:
2352:
2319:
2315:
2309:
2266:
2262:
2252:
2225:
2221:
2211:
2174:
2170:
2160:
2125:
2121:
2111:
2100:. Retrieved
2073:
2069:
2059:
2024:
2020:
2010:
1975:
1971:
1961:
1908:
1904:
1894:
1851:
1847:
1837:
1794:
1790:
1780:
1747:
1743:
1737:
1702:
1698:
1688:
1655:
1651:
1645:
1618:
1614:
1604:
1569:
1565:
1555:
1522:
1518:
1512:
1501:. Retrieved
1466:
1462:
1452:
1407:
1403:
1393:
1358:
1354:
1323:
1312:
1279:
1275:
1250:. Retrieved
1241:
1237:
1227:
1194:
1190:
1184:
1149:
1145:
1135:
1090:
1086:
1076:
1033:
1029:
1019:
984:mRNA capping
957:
953:
933:
919:
915:
908:
882:
874:
871:
856:
844:
838:
821:
809:
796:
784:
768:
760:
758:
751:
705:Spliceosomal
700:Prokaryotes
680:
653:
620:endonuclease
613:
608:
602:
589:
579:
560:
549:
543:
537:
529:
528:
505:
495:
489:
485:
479:
472:
439:
430:
426:
422:
275:
257:
246:
242:
238:
234:
228:
225:
203:
184:
180:transfer RNA
178:(rRNA), and
160:intracistron
159:
155:
151:
149:
132:
81:. For those
62:
31:RNA splicing
30:
29:
4078:Spliceosome
3882:Spliceosome
3841:RNA editing
3555:Translation
3392:to genetics
1972:RNA Biology
999:RNA editing
863:Morpholinos
761:branchpoint
676:Prokaryotes
380:Complex B*
260:spliceosome
188:pyrimidines
107:translation
101:, that is,
91:spliceosome
4062:Categories
3699:reversible
3662:lac operon
3638:imprinting
3633:Epigenetic
3625:Regulation
3580:Eukaryotic
3526:5' capping
3477:Eukaryotic
2102:2014-12-17
1503:2014-12-17
1011:References
893:stop codon
861:, such as
835:DNA repair
823:DNA damage
697:Eukaryotes
664:Eukaryotes
571:nucleoside
496:Drosophila
320:Complex E
305:(U2AF35),
109:is called
47:transcript
4002:Cytosolic
3570:Bacterial
3467:Bacterial
1244:(1): 31.
670:and some
650:Evolution
556:RNA world
551:Group III
519:pre-mRNAs
150:The word
135:catalysts
103:ribozymes
3682:microRNA
3596:Ribosome
3575:Archaeal
3531:Splicing
3503:Promoter
3472:Archaeal
3416: →
3412: →
3283:37698110
3275:15759099
3236:27208254
3187:19918805
3138:17128174
3130:11861299
3089:25112293
3040:25112293
2991:15333583
2950:17493584
2915:11574678
2866:32270393
2858:11477696
2817:26529031
2763:26545813
2712:38720076
2703:11164319
2663:19861426
2614:19959365
2565:21685335
2503:26607354
2452:24951248
2403:24244129
2336:18978789
2301:24196718
2203:24744771
2152:16049289
2144:15109492
2096:Archived
2051:16055381
2002:24178438
1953:18458335
1935:25461891
1886:25970244
1829:25970246
1772:21816910
1764:14685174
1729:15577853
1672:22065066
1637:10954700
1596:24452469
1547:14883495
1539:15956978
1497:Archived
1493:11421359
1444:21124863
1385:22705790
1304:23576288
1296:12626338
1252:31 March
1246:Archived
1219:33672491
1211:16485020
1176:22955974
974:DBASS3/5
963:See also
885:splicing
656:kingdoms
545:Group II
534:ribozyme
231:pre-mRNA
216:deletion
182:(tRNA).
172:proteins
63:splicing
3982:PRPF40B
3977:PRPF40A
3967:PRPF38B
3962:PRPF38A
3777:Nuclear
3435:RNA→DNA
3430:RNA→RNA
3418:Protein
3227:5047075
3178:2855871
3080:4440546
3031:4440546
2836:Genesis
2808:4693264
2754:4647561
2654:2779669
2605:2834840
2556:3131313
2533:Bibcode
2494:4732216
2443:4112133
2394:3820534
2371:Bibcode
2344:9228930
2292:4666680
2271:Bibcode
2244:9582290
2194:3978253
2092:3277966
2043:9200603
1993:3907479
1944:2383978
1913:Bibcode
1877:4529404
1856:Bibcode
1820:4471124
1799:Bibcode
1720:3902068
1587:4060434
1484:1370132
1435:2991248
1412:Bibcode
1376:3465671
1335:. 2013.
1167:3431479
1095:Bibcode
1068:4216649
1038:Bibcode
944:archaea
940:exteins
936:inteins
660:domains
609:In the
539:Group I
511:outrons
398:lariat;
146:Introns
75:nucleus
59:introns
3972:PRPF39
3957:PRPF31
3952:PRPF19
3947:PRPF18
3932:PRPF4B
3868:Intron
3347:(MeSH)
3281:
3273:
3234:
3224:
3185:
3175:
3136:
3128:
3087:
3077:
3038:
3028:
2989:
2948:
2913:
2903:
2864:
2856:
2815:
2805:
2761:
2751:
2710:
2700:
2682:Nature
2661:
2651:
2612:
2602:
2563:
2553:
2501:
2491:
2450:
2440:
2401:
2391:
2342:
2334:
2299:
2289:
2263:Nature
2242:
2201:
2191:
2177:: 63.
2150:
2142:
2090:
2049:
2041:
2000:
1990:
1951:
1941:
1933:
1884:
1874:
1848:Nature
1827:
1817:
1791:Nature
1770:
1762:
1727:
1717:
1680:846297
1678:
1670:
1635:
1594:
1584:
1545:
1537:
1491:
1481:
1442:
1432:
1383:
1373:
1302:
1294:
1217:
1209:
1174:
1164:
1127:418414
1125:
1118:411497
1115:
1066:
1060:622185
1058:
1030:Nature
859:oligos
636:TSEN15
634:, and
632:TSEN34
624:TSEN54
590:lariat
462:U6atac
460:, and
458:U4atac
299:snRNPs
294:, and
268:snRNPs
264:snRNPs
158:, and
152:intron
113:, the
95:snRNPs
4025:DCP1B
4020:DCP1A
3942:PRPF8
3937:PRPF6
3927:PRPF4
3922:PRPF3
3917:PLRG1
3887:minor
3877:snRNP
3563:Types
3460:Types
3279:S2CID
3134:S2CID
3108:Blood
2906:60237
2862:S2CID
2340:S2CID
2148:S2CID
2047:S2CID
1931:JSTOR
1768:S2CID
1676:S2CID
1543:S2CID
1300:S2CID
1215:S2CID
1064:S2CID
846:Ercc1
840:Brca1
628:TSEN2
611:yeast
446:snRNP
339:U2AF2
333:U2AF1
307:U2AF2
67:exons
41:(pre-
4045:EDC4
4040:EDC3
4035:DCPS
4030:DCP2
3872:Exon
3829:CFII
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