The nexus of long noncoding RNAs, splicing factors, alternative splicing and their modulations

References

• Djebali S, Davis CA, Merkel A, et al. Landscape of transcription in human cells. Nature. 2012;489(7414):101–108. doi: 10.1038/nature11233
   PubMed Web of Science ®Google Scholar
• Ulitsky I, Bartel DP. lincRNAs: genomics, evolution, and mechanisms. Cell. 2013;154(1):26. doi: 10.1016/j.cell.2013.06.020
   PubMed Web of Science ®Google Scholar
• Yang L, Lin C, Jin C, et al. lncRNA-dependent mechanisms of androgen-receptor-regulated gene activation programs. Nature. 2013;500(7464):598–602. doi: 10.1038/nature12451
   PubMed Web of Science ®Google Scholar
• Gutschner T, Hämmerle M, Diederichs S. MALAT1 — a paradigm for long noncoding RNA function in cancer. J Mol Med (Berl). 2013;91(7):791–801. doi: 10.1007/s00109-013-1028-y
   PubMed Web of Science ®Google Scholar
• Malakar P, Chartarifsky L, Hija A, et al. Insulin receptor alternative splicing is regulated by insulin signaling and modulates beta cell survival. Sci Rep. 2016;6(1):6. doi: 10.1038/srep31222
   PubMedGoogle Scholar
• Karni R, Hippo Y, Lowe SW, et al. The splicing-factor oncoprotein SF2/ASF activates mTORC1. Proc Natl Acad Sci U S A. 2008;105(40):15323–15327. doi: 10.1073/pnas.0801376105
   PubMed Web of Science ®Google Scholar
• Jbara A, Lin KT, Stossel C, et al. RBFOX2 modulates a metastatic signature of alternative splicing in pancreatic cancer. Nature. 2023;617(7959):147–153. doi: 10.1038/s41586-023-05820-3
   PubMed Web of Science ®Google Scholar
• Ben-Hur V, Denichenko P, Siegfried Z, et al. S6K1 alternative splicing modulates its oncogenic activity and regulates mTORC1. Cell Rep. 2013;3(1):103–115. doi: 10.1016/j.celrep.2012.11.020
   PubMed Web of Science ®Google Scholar
• Sever R, Brugge JS. Signal Transduction in cancer. Cold Spring Harb Perspect Med. 2015;5(4):a006098–a006098. doi: 10.1101/cshperspect.a006098
   PubMed Web of Science ®Google Scholar
• Venables JP. Aberrant and alternative splicing in cancer. Cancer Res. 2004;64(21):7647–7654. doi: 10.1158/0008-5472.CAN-04-1910
   PubMed Web of Science ®Google Scholar
• David CJ, Manley JL. Alternative pre-mRNA splicing regulation in cancer: pathways and programs unhinged. Genes Dev. 2010;24(21):2343. doi: 10.1101/gad.1973010
   PubMed Web of Science ®Google Scholar
• Srebrow A, Kornblihtt AR. The connection between splicing and cancer. J Cell Sci. 2006;119(13):2635–2641. doi: 10.1242/jcs.03053
   PubMed Web of Science ®Google Scholar
• Kim E, Goren A, Ast G. Insights into the connection between cancer and alternative splicing. Trends Genet. 2008;24(1):7–10. doi: 10.1016/j.tig.2007.10.001
   PubMed Web of Science ®Google Scholar
• Black DL. Mechanisms of alternative pre-messenger RNA splicing. Annu Rev Biochem. 2003;72(1):291–336. doi: 10.1146/annurev.biochem.72.121801.161720
   PubMedGoogle Scholar
• Gonçalves V, Pereira JFS, Jordan P. Signaling pathways driving aberrant splicing in cancer cells. Genes (Basel). 2018;9(1):9. doi: 10.3390/genes9010009
   Web of Science ®Google Scholar
• Shilo A, Siegfried Z, Karni R. The role of splicing factors in deregulation of alternative splicing during oncogenesis and tumor progression. Mol Cell Oncol. 2014;2(1):e970955. doi: 10.4161/23723548.2014.970955
   PubMedGoogle Scholar
• Busch A, Hertel KJ. Evolution of SR protein and hnRNP splicing regulatory factors. Wiley Interdiscip Rev RNA. 2012;3(1):1–12. doi: 10.1002/wrna.100
   PubMed Web of Science ®Google Scholar
• Chen M, Manley JL. Mechanisms of alternative splicing regulation: insights from molecular and genomics approaches. Nat Rev Mol Cell Biol. 2009;10(11):741. doi: 10.1038/nrm2777
   PubMed Web of Science ®Google Scholar
• Karni R, De Stanchina E, Lowe SW, et al. The gene encoding the splicing factor SF2/ASF is a proto-oncogene. Nat Struct Mol Biol. 2007;14(3):185–193. doi: 10.1038/nsmb1209
   PubMed Web of Science ®Google Scholar
• Dreyfuss G, Matunis MJ, Piñol-Roma S, et al. hnRNP proteins and the biogenesis of mRNA. Annu Rev Biochem. 1993;62(1):289–321. doi: 10.1146/annurev.bi.62.070193.001445
   PubMedGoogle Scholar
• Dreyfuss G, Kim VN, Kataoka N. Messenger-RNA-binding proteins and the messages they carry. Nat Rev Mol Cell Biol. 2002;3(3):195–205. doi: 10.1038/nrm760
   PubMed Web of Science ®Google Scholar
• Krecic AM, Swanson MS. hnRNP complexes: composition, structure, and function. Curr Opin Cell Biol. 1999;11(3):363–371. doi: 10.1016/S0955-0674(99)80051-9
   PubMed Web of Science ®Google Scholar
• Carpenter B, MacKay C, Alnabulsi A, et al. The roles of heterogeneous nuclear ribonucleoproteins in tumour development and progression. Biochim Biophys Acta. 2006;1765(2):85–100. doi: 10.1016/j.bbcan.2005.10.002
   PubMed Web of Science ®Google Scholar
• Golan-Gerstl R, Cohen M, Shilo A, et al. Splicing factor hnRNP A2/B1 regulates tumor suppressor gene splicing and is an oncogenic driver in glioblastoma. Cancer Res. 2011;71(13):4464–4472. doi: 10.1158/0008-5472.CAN-10-4410
   PubMed Web of Science ®Google Scholar
• Tauler J, Zudaire E, Liu H, et al. hnRNP A2/B1 modulates epithelial-mesenchymal transition in lung cancer cell lines. Cancer Res. 2010;70(18):7137–7147. doi: 10.1158/0008-5472.CAN-10-0860
   PubMed Web of Science ®Google Scholar
• Tang J, Chen Z, Wang Q, et al. Promotes colon cancer progression via the MAPK pathway. Front Genet. 2021;12:843. doi: 10.3389/fgene.2021.666451. hnRNPA2B1.
   Web of Science ®Google Scholar
• Romero-Barrios N, Legascue MF, Benhamed M, et al. Splicing regulation by long noncoding RNAs. Nucleic Acids Res. 2018;46(5):2169. doi: 10.1093/nar/gky095
   PubMed Web of Science ®Google Scholar
• Ji Q, Zhang L, Liu X, et al. Long non-coding RNA MALAT1 promotes tumour growth and metastasis in colorectal cancer through binding to SFPQ and releasing oncogene PTBP2 from SFPQ/PTBP2 complex. Br J Cancer. 2014;111(4):736–748. doi: 10.1038/bjc.2014.383
   PubMed Web of Science ®Google Scholar
• Li L, Feng T, Lian Y, et al. Role of human noncoding RNAs in the control of tumorigenesis. Proc Natl Acad Sci U S A. 2009;106(31):12956–12961. doi: 10.1073/pnas.0906005106
   PubMed Web of Science ®Google Scholar
• Wang G, Cui Y, Zhang G, et al. Regulation of proto-oncogene transcription, cell proliferation, and tumorigenesis in mice by PSF protein and a VL30 noncoding RNA. Proc Natl Acad Sci U S A. 2009;106(39):16794–16798. doi: 10.1073/pnas.0909022106
   PubMed Web of Science ®Google Scholar
• Romero-Barrios N, Legascue MF, Benhamed M, et al. Splicing regulation by long noncoding RNAs. Nucleic Acids Res. 2018;46(5):2169–2184. doi: 10.1093/nar/gky095
   PubMed Web of Science ®Google Scholar
• West JA, Davis CP, Sunwoo H, et al. The Long noncoding RNAs NEAT1 and MALAT1 bind active chromatin sites. Mol Cell. 2014;55(5):791–802. doi: 10.1016/j.molcel.2014.07.012
   PubMed Web of Science ®Google Scholar
• Statello L, Guo CJ, Chen LL, et al. Gene regulation by long non-coding RNAs and its biological functions. Nat Rev Mol Cell Biol. 2020;22(2):96–118. doi: 10.1038/s41580-020-00315-9
   PubMed Web of Science ®Google Scholar
• Pan Q, Shai O, Lee LJ, et al. Deep surveying of alternative splicing complexity in the human transcriptome by high-throughput sequencing. Nat Genet. 2008;40(12):1413–1415. doi: 10.1038/ng.259
   PubMed Web of Science ®Google Scholar
• WANG Y, LIU J, HUANG B, et al. Mechanism of alternative splicing and its regulation. Biomed Rep. 2015;3(2):152. doi: 10.3892/br.2014.407
   PubMed Web of Science ®Google Scholar
• Fu XD, Ares M. Context-dependent control of alternative splicing by RNA-binding proteins. Nat Rev Genet. 2014;15(10):689. doi: 10.1038/nrg3778
   PubMed Web of Science ®Google Scholar
• Bonnal SC, López-Oreja I, Valcárcel J. Roles and mechanisms of alternative splicing in cancer — implications for care. Nat Rev Clin Oncol. 2020;17(8):457–474. doi: 10.1038/s41571-020-0350-x
   PubMed Web of Science ®Google Scholar
• Suzuki H, Kumar SA, Shuai S, et al. Recurrent noncoding U1 snRNA mutations drive cryptic splicing in SHH medulloblastoma. Nature. 2019;574(7780):707–711. doi: 10.1038/s41586-019-1650-0
   PubMed Web of Science ®Google Scholar
• Obeng EA, Chappell RJ, Seiler M, et al. Physiologic expression of Sf3b1K700E causes impaired erythropoiesis, aberrant splicing, and sensitivity to therapeutic spliceosome modulation. Cancer Cell. 2016;30(3):404–417. doi: 10.1016/j.ccell.2016.08.006
   PubMed Web of Science ®Google Scholar
• Shirai CL, Ley JN, White BS, et al. Mutant U2AF1 expression alters hematopoiesis and pre-mRNA splicing in vivo. Cancer Cell. 2015;27(5):631–643. doi: 10.1016/j.ccell.2015.04.008
   PubMed Web of Science ®Google Scholar
• Yoshida K, Sanada M, Shiraishi Y, et al. Frequent pathway mutations of splicing machinery in myelodysplasia. Nature. 2011;478(7367):64–69. doi: 10.1038/nature10496
   PubMed Web of Science ®Google Scholar
• Darman RB, Seiler M, Agrawal AA, et al. Cancer-associated SF3B1 hotspot mutations induce cryptic 3′ splice site selection through use of a different branch point. Cell Rep. 2015;13(5):1033–1045. doi: 10.1016/j.celrep.2015.09.053
   PubMed Web of Science ®Google Scholar
• DeBoever C, Ghia EM, Shepard PJ, et al. Transcriptome sequencing reveals potential mechanism of cryptic 3’ splice site selection in SF3B1-mutated cancers. PLoS Comput Biol. 2015;11(3):e1004105. doi: 10.1371/journal.pcbi.1004105
   PubMed Web of Science ®Google Scholar
• Shiozawa Y, Malcovati L, Gallì A, et al. Aberrant splicing and defective mRNA production induced by somatic spliceosome mutations in myelodysplasia. Nat Commun. 2018;9(1):1–16. doi: 10.1038/s41467-018-06063-x
   PubMedGoogle Scholar
• Zhang Y, Qian J, Gu C, et al. Alternative splicing and cancer: a systematic review. Signal Transduction and Targeted Therapy 2021; 6:1–14.
   PubMed Web of Science ®Google Scholar
• Elliott K, Sakamuro D, Basu A, et al. Bin1 functionally interacts with Myc and inhibits cell proliferation via multiple mechanisms. Oncogene. 1999;18(24):3564–3573. doi: 10.1038/sj.onc.1202670
   PubMed Web of Science ®Google Scholar
• Ge K, Duhadaway J, Du W, et al. Mechanism for elimination of a tumor suppressor: aberrant splicing of a brain-specific exon causes loss of function of Bin1 in melanoma. Proc Natl Acad Sci U S A. 1999;96(17):9689–9694. doi: 10.1073/pnas.96.17.9689
   PubMed Web of Science ®Google Scholar
• Shimoni-Sebag A, Lebenthal-Loinger I, Zender L, et al. RRM1 domain of the splicing oncoprotein SRSF1 is required for MEK1-MAPK-ERK activation and cellular transformation. Carcinogenesis. 2013;34(11):2498–2504. doi: 10.1093/carcin/bgt247
   PubMed Web of Science ®Google Scholar
• Das S, Anczuków O, Akerman M, et al. Oncogenic Splicing Factor SRSF1 Is a Critical Transcriptional Target of MYC. Cell Rep. 2012;1(2):110–117. doi: 10.1016/j.celrep.2011.12.001
   PubMed Web of Science ®Google Scholar
• Choi S, Lee HS, Cho N, et al. RBFOX2-regulated TEAD1 alternative splicing plays a pivotal role in hippo-YAP signaling. Nucleic Acids Res. 2022;50(15):8658–8673. doi: 10.1093/nar/gkac509
   PubMed Web of Science ®Google Scholar
• Étienne HM, Vogel G, Zabarauskas A, et al. The Sam68 STAR RNA-Binding Protein Regulates mTOR Alternative Splicing during Adipogenesis. Mol Cell. 2012;46(2):187–199. doi: 10.1016/j.molcel.2012.02.007
   PubMed Web of Science ®Google Scholar
• Marani M, Tenev T, Hancock D, et al. Identification of novel isoforms of the BH3 domain protein bim which directly activate bax to trigger apoptosis. Mol Cell Biol. 2002;22(11):3577. doi: 10.1128/MCB.22.11.3577-3589.2002
   PubMed Web of Science ®Google Scholar
• Urbanski LM, Leclair N, Anczuków O. Alternative-splicing defects in cancer: splicing regulators and their downstream targets, guiding the way to novel cancer therapeutics. Wiley Interdiscip Rev RNA. 2018;9(4). doi: 10.1002/wrna.1476
   PubMed Web of Science ®Google Scholar
• Escobar-Hoyos L, Knorr K, Abdel-Wahab O. Aberrant RNA splicing in cancer. Annu Rev Cancer Biol. 2019;3(1):167–185. doi: 10.1146/annurev-cancerbio-030617-050407
   PubMedGoogle Scholar
• Yoshida K, Sanada M, Shiraishi Y, et al. Frequent pathway mutations of splicing machinery in myelodysplasia. Nature. 2011;478(7367):64–69. doi: 10.1038/nature10496
   PubMed Web of Science ®Google Scholar
• Papaemmanuil E, Gerstung M, Malcovati L, et al. Consortium on behalf of the CMD working group of the ICG, et al. Clinical and biological implications of driver mutations in myelodysplastic syndromes. Blood. 2013;122(22):3616–27; quiz 3699. doi: 10.1182/blood-2013-08-518886
   PubMed Web of Science ®Google Scholar
• Graubert TA, Shen D, Ding L, et al. RECURRENT MUTATIONS IN THE U2AF1 SPLICING FACTOR IN MYELODYSPLASTIC SYNDROMES. Nat Genet. 2012;44(1):53. doi: 10.1038/ng.1031
   Web of Science ®Google Scholar
• Madan V, Kanojia D, Li J, et al. Aberrant splicing of U12-type introns is the hallmark of ZRSR2 mutant myelodysplastic syndrome. Nat Commun. 2015;6(1):1–14. doi: 10.1038/ncomms7042
   Google Scholar
• Kim E, Ilagan JO, Liang Y, et al. SRSF2 mutations contribute to myelodysplasia by mutant-specific effects on exon recognition. Cancer Cell. 2015;27(5):617–630. doi: 10.1016/j.ccell.2015.04.006
   PubMed Web of Science ®Google Scholar
• Lasho TL, Finke CM, Hanson CA, et al. SF3B1 mutations in primary myelofibrosis: clinical, histopathology and genetic correlates among 155 patients. Leukemia. 2012;26(5):1135–1137. doi: 10.1038/leu.2011.320
   PubMed Web of Science ®Google Scholar
• Quesada V, Conde L, Villamor N, et al. Exome sequencing identifies recurrent mutations of the splicing factor SF3B1 gene in chronic lymphocytic leukemia. Nature Genet. 2012;44(1):47–52. doi: 10.1038/ng.1032
   Web of Science ®Google Scholar
• Wang L, Lawrence MS, Wan Y, et al. SF3B1 and other novel cancer genes in chronic lymphocytic leukemia. N Engl J Med. 2011;365(26):2497–2506. doi: 10.1056/NEJMoa1109016
   PubMed Web of Science ®Google Scholar
• Alsafadi S, Houy A, Battistella A, et al. Cancer-associated SF3B1 mutations affect alternative splicing by promoting alternative branchpoint usage. Nat Commun. 2016;7(1):1–12. doi: 10.1038/ncomms10615
   Google Scholar
• Saez B, Walter MJ, Graubert TA. Splicing factor gene mutations in hematologic malignancies. Blood. 2017;129(10):1260. doi: 10.1182/blood-2016-10-692400
   PubMed Web of Science ®Google Scholar
• Cohen-Eliav M, Golan-Gerstl R, Siegfried Z, et al. The splicing factor SRSF6 is amplified and is an oncoprotein in lung and colon cancers. J Pathol. 2013;229(4):630–639. doi: 10.1002/path.4129
   PubMed Web of Science ®Google Scholar
• Gonçalves V, Jordan P. Posttranscriptional regulation of splicing factor SRSF1 and its role in cancer cell Biology. Biomed Res Int. 2015;2015:1–10. doi: 10.1155/2015/287048
   Web of Science ®Google Scholar
• David CJ, Chen M, Assanah M, et al. HnRNP proteins controlled by c-myc deregulate pyruvate kinase mRNA splicing in cancer. Nature. 2010;463(7279):364–368. doi: 10.1038/nature08697
   PubMed Web of Science ®Google Scholar
• Bradley T, Cook ME, Blanchette M. SR proteins control a complex network of RNA-processing events. RNA. 2015;21(1):75–92. doi: 10.1261/rna.043893.113
   PubMed Web of Science ®Google Scholar
• Martinez-Contreras R, Cloutier P, Shkreta L, et al. hnRNP proteins and splicing control. Adv Exp Med Biol. 2007;623:123–147.
   PubMed Web of Science ®Google Scholar
• Graveley BR, Maniatis T. Arginine/serine-rich domains of SR proteins can function as activators of pre-mRNA splicing. Mol Cell. 1998;1(5):765–771. doi: 10.1016/S1097-2765(00)80076-3
   PubMed Web of Science ®Google Scholar
• Anczuków O, Akerman M, Cléry A, et al. SRSF1-regulated alternative splicing in breast cancer. Mol Cell. 2015;60(1):105. doi: 10.1016/j.molcel.2015.09.005
   PubMed Web of Science ®Google Scholar
• Jiang L, Huang J, Higgs BW, et al. Genomic landscape survey identifies SRSF1 as a key oncodriver in small cell lung cancer. PLoS Genet. 2016;12(4):12. doi: 10.1371/journal.pgen.1005895
   Web of Science ®Google Scholar
• Wan L, Yu W, Shen E, et al. SRSF6-regulated alternative splicing that promotes tumour progression offers a therapy target for colorectal cancer. Gut. 2019;68(1):118–129. doi: 10.1136/gutjnl-2017-314983
   PubMed Web of Science ®Google Scholar
• Mattick JS, Amaral PP, Carninci P, et al. Long non-coding RNAs: definitions, functions, challenges and recommendations. Nat Rev Mol Cell Biol. 2023;17:1–17.
   Google Scholar
• Wu H, Yin QF, Luo Z, et al. Unusual processing generates SPA LncRNAs that sequester multiple RNA binding proteins. Mol Cell. 2016;64(3):534–548. doi: 10.1016/j.molcel.2016.10.007
   PubMed Web of Science ®Google Scholar
• Wilusz JE, Freier SM, Spector DL. 3′ end processing of a Long nuclear-Retained noncoding RNA yields a tRNA-like cytoplasmic RNA. Cell. 2008;135(5):919–932. doi: 10.1016/j.cell.2008.10.012
   PubMed Web of Science ®Google Scholar
• Cheetham SW, Faulkner GJ, Dinger ME. Overcoming challenges and dogmas to understand the functions of pseudogenes. Nat Rev Genet. 2019;21(3):191–201. doi: 10.1038/s41576-019-0196-1
   PubMed Web of Science ®Google Scholar
• Patop IL, Wüst S, Kadener S. Past, present, and future of circRNAs. EMBO J. 2019;38(16):e100836. cited 2023. doi: 10.15252/embj.2018100836
   PubMed Web of Science ®Google Scholar
• Mercer TR, Wilhelm D, Dinger ME, et al. Expression of distinct RNAs from 3′ untranslated regions. Nucleic Acids Res. 2011;39(6):2393–2403. doi: 10.1093/nar/gkq1158
   PubMed Web of Science ®Google Scholar
• Seal RL, Chen L-L, Griffiths-Jones S, et al. A guide to naming human non-coding RNA genes. EMBO J. 2020;39(6):e103777. doi: 10.15252/embj.2019103777
   PubMed Web of Science ®Google Scholar
• Uszczynska-Ratajczak B, Lagarde J, Frankish A, et al. Towards a complete map of the human long non-coding RNA transcriptome. Nat Rev Genet. 2018;19(9):535–548. doi: 10.1038/s41576-018-0017-y
   PubMed Web of Science ®Google Scholar
• Mattick JS, Rinn JL. Discovery and annotation of long noncoding RNAs. Nat Struct Mol Biol. 2015;22(1):5–7. doi: 10.1038/nsmb.2942
   PubMed Web of Science ®Google Scholar
• Frankish A, Diekhans M, Jungreis I, et al. GENCODE 2021. Nucleic Acids Res. 2021;49(D1):D916–23. doi: 10.1093/nar/gkaa1087
   PubMed Web of Science ®Google Scholar
• Mattick JS, Amaral PP, Carninci P, et al. Long non-coding RNAs: definitions, functions, challenges and recommendations. Nat Rev Mol Cell Biol. 2023;17:1–17.
   Google Scholar
• Wu P, Mo Y, Peng M, et al. Emerging role of tumor-related functional peptides encoded by lncRNA and circRNA. Mol Cancer. 2020;19(1):1–14. doi: 10.1186/s12943-020-1147-3
   PubMed Web of Science ®Google Scholar
• Williamson L, Saponaro M, Boeing S, et al. UV irradiation induces a non-coding RNA that functionally opposes the protein encoded by the same gene. Cell. 2017;168(5):843–855.e13. doi: 10.1016/j.cell.2017.01.019
   PubMed Web of Science ®Google Scholar
• Hube F, Guo J, Chooniedass-Kothari S, et al. Alternative splicing of the first intron of the steroid receptor RNA activator SRA. Participates In The Generation Of Coding And Noncoding RNA Isoforms In Breast Cancer Cell Lines. 2006;25(7):418–428. doi: 10.1089/dna.2006.25.418
   Google Scholar
• Gonzàlez-Porta M, Frankish A, Rung J, et al. Transcriptome analysis of human tissues and cell lines reveals one dominant transcript per gene. Genome Biol. 2013;14(7):1–11. doi: 10.1186/gb-2013-14-7-r70
   Web of Science ®Google Scholar
• Kung JTY, Colognori D, Lee JT. Long Noncoding RNAs: Past, Present, and Future. Genetics. 2013;193(3):651. doi: 10.1534/genetics.112.146704
   PubMed Web of Science ®Google Scholar
• Statello L, Guo CJ, Chen LL, et al. Gene regulation by long non-coding RNAs and its biological functions. Nat Rev Mol Cell Biol. 2020;22(2):96–118. doi: 10.1038/s41580-020-00315-9
   PubMed Web of Science ®Google Scholar
• Malakar P, Shilo A, Mogilevsky A, et al. Long noncoding RNA MALAT1 promotes hepatocellular carcinoma development by SRSF1 upregulation and mTOR activation. Cancer Res. 2017;77(5):1155–1167. doi: 10.1158/0008-5472.CAN-16-1508
   PubMed Web of Science ®Google Scholar
• Huang GW, Zhang YL, Di LL, et al. Natural antisense transcript TPM1-AS regulates the alternative splicing of tropomyosin I through an interaction with RNA-binding motif protein 4. Int J Biochem Cell Biol. 2017;90:59–67. doi: 10.1016/j.biocel.2017.07.017
   PubMed Web of Science ®Google Scholar
• Duan Y, Jia Y, Wang J, et al. Long noncoding RNA DGCR5 involves in tumorigenesis of esophageal squamous cell carcinoma via SRSF1-mediated alternative splicing of Mcl-1. Cell Death Dis. 2021;12:587. doi: 10.1038/s41419-021-03858-7
   PubMed Web of Science ®Google Scholar
• Tripathi V, Ellis JD, Shen Z, et al. The nuclear-Retained noncoding RNA MALAT1 regulates alternative splicing by modulating SR splicing factor phosphorylation. Mol Cell. 2010;39(6):925. doi: 10.1016/j.molcel.2010.08.011
   PubMed Web of Science ®Google Scholar
• Pruszko M, Milano E, Forcato M, et al. The mutant p53‐ID4 complex controls VEGFA isoforms by recruiting lncRNA MALAT1. EMBO Rep. 2017;18(8):1331. doi: 10.15252/embr.201643370
   PubMed Web of Science ®Google Scholar
• Ji Q, Zhang L, Liu X, et al. Long non-coding RNA MALAT1 promotes tumour growth and metastasis in colorectal cancer through binding to SFPQ and releasing oncogene PTBP2 from SFPQ/PTBP2 complex. Br J Cancer. 2014;111(4):736–748. doi: 10.1038/bjc.2014.383
   PubMed Web of Science ®Google Scholar
• Wu H, Zheng J, Deng J, et al. LincRNA-uc002yug.2 involves in alternative splicing of RUNX1 and serves as a predictor for esophageal cancer and prognosis. Oncogene. 2015;34(36):4723–4734. doi: 10.1038/onc.2014.400
   PubMed Web of Science ®Google Scholar
• Meng LD, Shi GD, Ge WL, et al. Linc01232 promotes the metastasis of pancreatic cancer by suppressing the ubiquitin-mediated degradation of HNRNPA2B1 and activating the A-Raf-induced MAPK/ERK signaling pathway. Cancer Lett. 2020;494:107–120. doi: 10.1016/j.canlet.2020.08.001
   PubMed Web of Science ®Google Scholar
• Tripathi V, Song DY, Zong X, et al. SRSF1 regulates the assembly of pre-mRNA processing factors in nuclear speckles. Mol Biol Cell. 2012;23(18):3694–3706. doi: 10.1091/mbc.e12-03-0206
   PubMed Web of Science ®Google Scholar
• Liu Y, Zhang YM, Ma FB, et al. Long noncoding RNA HOXA11-AS promotes gastric cancer cell proliferation and invasion via SRSF1 and functions as a biomarker in gastric cancer. World J Gastroenterol. 2019;25(22):2763. doi: 10.3748/wjg.v25.i22.2763
   PubMed Web of Science ®Google Scholar
• Xu L, Wang Z, Yin C, et al. Long noncoding RNA LINC02580 suppresses the invasion–metastasis cascade in hepatocellular carcinoma by targeting SRSF1. Biochem Biophys Res Commun. 2020;533(4):685–691. doi: 10.1016/j.bbrc.2020.10.061
   PubMed Web of Science ®Google Scholar
• Dong M, Dong Z, Zhu X, et al. Long non-coding RNA MIR205HG regulates KRT17 and tumor processes in cervical cancer via interaction with SRSF1. Exp Mol Pathol 2019; 111.
   Web of Science ®Google Scholar
• Li H, Guo S, Zhang M, et al. Long non-coding RNA AGAP2-AS1 accelerates cell proliferation, migration, invasion and the EMT process in colorectal cancer via regulating the miR-4,668-3p/SRSF1 axis. J Gene Med 2020; 22.
   Web of Science ®Google Scholar
• Zhou X, Li X, Yu L, et al. The RNA-binding protein SRSF1 is a key cell cycle regulator via stabilizing NEAT1 in glioma. Int J Biochem Cell Biol. 2019;113:75–86. doi: 10.1016/j.biocel.2019.06.003
   PubMed Web of Science ®Google Scholar
• Wu Z-H, Liu C-C, Zhou Y-Q, et al. OnclncRNA-626 promotes malignancy of gastric cancer via inactivated the p53 pathway through interacting with SRSF1. Am J Cancer Res. 2019;9:2249.
   PubMed Web of Science ®Google Scholar
• SUN Q, HAO Q, LIN YC, et al. Antagonism between splicing and microprocessor complex dictates the serum-induced processing of lnc-MIRHG for efficient cell cycle reentry. RNA. 2020;26(11):1603–1620. doi: 10.1261/rna.075309.120
   PubMed Web of Science ®Google Scholar
• Wu J, Wang N, Yang Y, et al. Correction: LINC01152 upregulates MAML2 expression to modulate the progression of glioblastoma multiforme via notch signaling pathway. Cell Death Dis. 2021;12(10):12. doi: 10.1038/s41419-021-04106-8
   PubMed Web of Science ®Google Scholar
• Spiniello M, Knoener RA, Steinbrink MI, et al. HyPR-MS for multiplexed Discovery of MALAT1, NEAT1, and NORAD lncRNA protein interactomes. J Proteome Res. 2018;17(9):3022–3038. doi: 10.1021/acs.jproteome.8b00189
   PubMed Web of Science ®Google Scholar
• West JA, Davis CP, Sunwoo H, et al. The Long noncoding RNAs NEAT1 and MALAT1 bind active chromatin sites. Mol Cell. 2014;55(5):791–802. doi: 10.1016/j.molcel.2014.07.012
   PubMed Web of Science ®Google Scholar
• Mao Q, Li L, Zhang C, et al. Long non coding RNA NRON inhibited breast cancer development through regulating miR-302b/SRSF2 axis. Am J Transl Res. 2020;12:4683–4692.
   PubMed Web of Science ®Google Scholar
• Chen Y, Shen T, Ding X, et al. lncRNA MRUL suppressed non-small cell lung cancer cells proliferation and invasion by targeting miR-17-5p/SRSF2 axis. Biomed Res Int;2020. doi: 10.1155/2020/9567846
   Google Scholar
• Yu B, Qu L, Wu T, et al. A novel LncRNA, AC091729.7 promotes sinonasal squamous cell carcinomas proliferation and invasion through binding SRSF2. Front Oncol. 2020;9: doi: 10.3389/fonc.2019.01575
   Web of Science ®Google Scholar
• El Bassit G, Patel RS, Carter G, et al. MALAT1 in human adipose stem cells modulates survival and alternative splicing of PKCδII in HT22 cells. Endocrinology. 2017;158:183–195. doi: 10.1210/en.2016-1819
   PubMed Web of Science ®Google Scholar
• Wang H, Liu M, Fang L, et al. The cisplatin-induced lncRNA PANDAR dictates the chemoresistance of ovarian cancer via regulating SFRS2-mediated p53 phosphorylation. Cell Death Dis. 2018;9(11). doi: 10.1038/s41419-018-1148-y
   Google Scholar
• Gao H, Sun Y, Chen J, et al. Long non-coding RNA AFAP1-AS1 promotes cell growth and inhibits apoptosis by binding to specific proteins in germinal center B-cell-like diffuse large B-cell lymphoma. Am J Transl Res. 2020;12:8225–8246.
   PubMed Web of Science ®Google Scholar
• Wang Y, Zhang W, Liu W, et al. Long noncoding RNA VESTAR regulates lymphangiogenesis and Lymph Node metastasis of esophageal squamous cell carcinoma by enhancing VEGFC mRNA stability. Cancer Res. 2021;81(12):3187–3199. doi: 10.1158/0008-5472.CAN-20-1713
   PubMed Web of Science ®Google Scholar
• Kong J, Sun W, Li C, et al. Long non-coding RNA LINC01133 inhibits epithelial–mesenchymal transition and metastasis in colorectal cancer by interacting with SRSF6. Cancer Lett. 2016;380(2):476–484. doi: 10.1016/j.canlet.2016.07.015
   PubMed Web of Science ®Google Scholar
• Zhang F, Wang H, Yu J, et al. LncRNA CRNDE attenuates chemoresistance in gastric cancer via SRSF6-regulated alternative splicing of PICALM. Mol Cancer. 2021;20(1):20. doi: 10.1186/s12943-020-01299-y
   PubMed Web of Science ®Google Scholar
• Si Z, Yu L, Jing H, et al. Oncogenic lncRNA ZNF561-AS1 is essential for colorectal cancer proliferation and survival through regulation of miR-26a-3p/miR-128-5p-SRSF6 axis. J Exp Clin Cancer Res. 2021;40(1):40. doi: 10.1186/s13046-021-01882-1
   PubMed Web of Science ®Google Scholar
• Song J, Su ZZ, Shen QM. Long non-coding RNA MALAT1 regulates proliferation, apoptosis, migration and invasion via miR-374b-5p/SRSF7 axis in non-small cell lung cancer. Eur Rev Med Pharmacol Sci. 2020;24(4):1853–1862. doi: 10.26355/eurrev_202002_20363
   PubMed Web of Science ®Google Scholar
• He R, Wu S, Gao R, et al. Identification of a Long noncoding RNA TRAF3IP2-AS1 as key regulator of IL-17 signaling through the SRSF10–IRF1–Act1 axis in autoimmune diseases. J Immunol. 2021;206(10):2353–2365. doi: 10.4049/jimmunol.2001223
   PubMed Web of Science ®Google Scholar
• Zhou B, Wang Y, Jiang J, et al. The long noncoding RNA colon cancer-associated transcript-1/miR-490 axis regulates gastric cancer cell migration by targeting hnRNPA1. IUBMB Life. 2016;68(3):201–210. doi: 10.1002/iub.1474
   PubMed Web of Science ®Google Scholar
• Lan Z, Yao X, Sun K, et al. The interaction between lncRNA SNHG6 and hnRNPA1 contributes to the growth of colorectal cancer by enhancing aEROBIC GLYCOLYsis through the regulation of alternative splicing of PKM. Front Oncol. 2020;10:10. doi: 10.3389/fonc.2020.00363
   PubMed Web of Science ®Google Scholar
• Zhang Y, Huang W, Yuan Y, et al. Long non-coding RNA H19 promotes colorectal cancer metastasis via binding to hnRNPA2B1. J Exp Clin Cancer Res. 2020;39(1):39. doi: 10.1186/s13046-020-01619-6
   Web of Science ®Google Scholar
• Yu PF, Kang AR, Jing LJ, et al. Long non-coding RNA CACNA1G-AS1 promotes cell migration, invasion and epithelial-mesenchymal transition by HNRNPA2B1 in non-small cell lung cancer. Eur Rev Med Pharmacol Sci. 2018;22(4):993–1002. doi: 10.26355/eurrev_201802_14381
   PubMed Web of Science ®Google Scholar
• Wang H, Liang L, Dong Q, et al. Long noncoding RNA miR503HG, a prognostic indicator, inhibits tumor metastasis by regulating the HNRNPA2B1/NF-κB pathway in hepatocellular carcinoma. Theranostics. 2018;8(10):2814–2829. doi: 10.7150/thno.23012
   PubMed Web of Science ®Google Scholar
• Chen T, Gu C, Xue C, et al. LncRNA-uc002mbe.2 interacting with hnRNPA2B1 mediates AKT deactivation and p21 up-regulation induced by Trichostatin in liver cancer cells. Front Pharmacol. 2017;8: doi: 10.3389/fphar.2017.00669
   Web of Science ®Google Scholar
• Singh R, Gupta SC, Peng WX, et al. Regulation of alternative splicing of Bcl-x by BC200 contributes to breast cancer pathogenesis. Cell Death Dis. 2016;7(6):e2262. doi: 10.1038/cddis.2016.168
   PubMedGoogle Scholar
• Zhang Y, Chen W, Pan T, et al. LBX2-AS1 is activated by ZEB1 and promotes the development of esophageal squamous cell carcinoma by interacting with HNRNPC to enhance the stability of ZEB1 and ZEB2 mRNAs. Biochem Biophys Res Commun. 2019;511(3):566–572. doi: 10.1016/j.bbrc.2019.02.079
   PubMed Web of Science ®Google Scholar
• McHugh CA, Chen CK, Chow A, et al. The Xist lncRNA interacts directly with SHARP to silence transcription through HDAC3. Nature. 2015;521(7551):232–236. doi: 10.1038/nature14443
   PubMed Web of Science ®Google Scholar
• Xiang Z, Lv Q, Zhang Y, et al. Long non-coding RNA DDX11-AS1 promotes the proliferation and migration of glioma cells by combining with HNRNPC. Mol Ther Nucleic Acids. 2022;28:601–612. doi: 10.1016/j.omtn.2022.04.016
   PubMedGoogle Scholar
• Li J, He M, Xu W, et al. RETRACTED ARTICLE: knockdown of FOXO3a induces epithelial-mesenchymal transition and promotes metastasis of pancreatic ductal adenocarcinoma by activation of the β-catenin/TCF4 pathway through SPRY2. J Exp Clin Cancer Res. 2019;38(1):38. doi: 10.1186/s13046-019-1046-x
   PubMedGoogle Scholar
• Zheng F, Chen J, Zhang X, et al. The HIF-1α antisense long non-coding RNA drives a positive feedback loop of HIF-1α mediated transactivation and glycolysis. Nat Commun. 2021;12(1):12. doi: 10.1038/s41467-021-21535-3
   PubMed Web of Science ®Google Scholar
• Liu N, Zhou KI, Parisien M, et al. N6-methyladenosine alters RNA structure to regulate binding of a low-complexity protein. Nucleic Acids Res. 2017;45(10):6051–6063. doi: 10.1093/nar/gkx141
   PubMed Web of Science ®Google Scholar
• Li M, Li L, Zhang X, et al. LncRNA RP11-670E13.6, interacted with hnRNPH, delays cellular senescence by sponging microRNA-663a in UVB damaged dermal fibroblasts. Aging. 2019;11(16):5992. doi: 10.18632/aging.102159
   PubMedGoogle Scholar
• Li D, Wang X, Mei H, et al. Long noncoding RNA pancEts-1 promotes neuroblastoma progression through hnRNPK-Mediated β-catenin stabilization. Cancer Res. 2018;78(5):1169–1183. doi: 10.1158/0008-5472.CAN-17-2295
   PubMed Web of Science ®Google Scholar
• Toki N, Takahashi H, Sharma H, et al. SINEUP long non-coding RNA acts via PTBP1 and HNRNPK to promote translational initiation assemblies. Nucleic Acids Res. 2020;48(20):11626–11644. doi: 10.1093/nar/gkaa814
   PubMed Web of Science ®Google Scholar
• Zhang Z, Zhou C, Chang Y, et al. Long non-coding RNA CASC11 interacts with hnRNP-K and activates the WNT/β-catenin pathway to promote growth and metastasis in colorectal cancer. Cancer Lett. 2016;376(1):62–73. doi: 10.1016/j.canlet.2016.03.022
   PubMed Web of Science ®Google Scholar
• Van Alphen RJ, Wiemer EAC, Burger H, et al. The spliceosome as target for anticancer treatment. Br J Cancer. 2009;100:2. doi: 10.1038/sj.bjc.6604801. 2008; 100:228–232.
   Web of Science ®Google Scholar
• Lee CC, Chang WH, Chang YS, et al. Alternative splicing in human cancer cells is modulated by the amiloride derivative 3,5‐diamino‐6‐chloro‐N‐(N‐(2,6‐dichlorobenzoyl)carbamimidoyl)pyrazine‐2‐carboxide. Mol Oncol. 2019;13(8):1744. doi: 10.1002/1878-0261.12524
   PubMed Web of Science ®Google Scholar
• Chang JG, Yang DM, Chang WH, et al. Small Molecule Amiloride Modulates Oncogenic RNA Alternative Splicing to Devitalize Human Cancer Cells. PLoS One. 2011;6(6):e18643. doi: 10.1371/journal.pone.0018643
   PubMed Web of Science ®Google Scholar
• Kaida D, Motoyoshi H, Tashiro E, et al. Spliceostatin a targets SF3b and inhibits both splicing and nuclear retention of pre-mRNA. Nat Chem Biol. 2007;3(9):576–583. 3. doi: 10.1038/nchembio.2007.18
   PubMed Web of Science ®Google Scholar
• Nakajima H, Hori Y, Terano H, et al. New antitumor substances, FR901463, FR901464 and FR901465. II. Activities against experimental tumors in mice and mechanism of action. J Antibiot (Tokyo). 1996;49(12):1204–1211. doi: 10.7164/antibiotics.49.1204
   PubMed Web of Science ®Google Scholar
• Convertini P, Shen M, Potter PM, et al. Sudemycin E influences alternative splicing and changes chromatin modifications. Nucleic Acids Res. 2014;42(8):4947. doi: 10.1093/nar/gku151
   PubMed Web of Science ®Google Scholar
• Lee SCW, Abdel-Wahab O. Therapeutic targeting of splicing in cancer. Nat Med. [cited 2023 Feb 17] 2016;22:976. Available from:/pmc/articles/PMC5644489/ doi:10.1038/nm.4165.
   PubMed Web of Science ®Google Scholar
• Xiao SH, Manley JL. Phosphorylation of the ASF/SF2 RS domain affects both protein-protein and protein-RNA interactions and is necessary for splicing. Genes Dev. 1997;11(3):334–344. doi: 10.1101/gad.11.3.334
   PubMed Web of Science ®Google Scholar
• Cao W, Jamison SF, Garcia-Blanco MA. Both phosphorylation and dephosphorylation of ASF/SF2 are required for pre-mRNA splicing in vitro. RNA. 1997;3(12):1456.
   PubMed Web of Science ®Google Scholar
• Gui JF, Tronchere H, Chandler SD, et al. Purification and characterization of a kinase specific for the serine- and arginine-rich pre-mRNA splicing factors. Proc Natl Acad Sci U S A. 1994;91(23):10824–10828. doi: 10.1073/pnas.91.23.10824
   PubMed Web of Science ®Google Scholar
• Kuroyanagi N, Onogi H, Wakabayashi T, et al. Novel SR-protein-specific kinase, SRPK2, disassembles nuclear speckles. Biochem Biophys Res Commun. 1998;242(2):357–364. doi: 10.1006/bbrc.1997.7913
   PubMed Web of Science ®Google Scholar
• Rossi F, Labourier E, Forné T, et al. Specific phosphorylation of SR proteins by mammalian DNA topoisomerase I. Nature. 1996;381(6577):80–82. 1996 381:6577. doi: 10.1038/381080a0
   PubMed Web of Science ®Google Scholar
• Colwill K, Pawson T, Andrews B, et al. The Clk/Sty protein kinase phosphorylates SR splicing factors and regulates their intranuclear distribution. EMBO J. 1996;15(2):265. doi: 10.1002/j.1460-2075.1996.tb00357.x
   PubMed Web of Science ®Google Scholar
• Naro C, Sette C. Phosphorylation-mediated regulation of alternative splicing in cancer. Int J Cell Biol. 2013;2013:1–15. doi: 10.1155/2013/151839
   Google Scholar
• Pilch B, Allemand E, Facompré M, et al. Specific inhibition of serine- and arginine-rich splicing factors phosphorylation, spliceosome assembly, and splicing by the antitumor drug NB-506. Cancer Res. 2001;61(18):6876–6884.
   PubMed Web of Science ®Google Scholar
• Soret J, Bakkour N, Maire S, et al. Selective modification of alternative splicing by indole derivatives that target serine-arginine-rich protein splicing factors. Proc Natl Acad Sci U S A. 2005;102(24):8764–8769. doi: 10.1073/pnas.0409829102
   PubMed Web of Science ®Google Scholar
• Muraki M, Ohkawara B, Hosoya T, et al. Manipulation of alternative splicing by a newly developed inhibitor of clks. J Biol Chem. 2004;279(23):24246–24254. doi: 10.1074/jbc.M314298200
   PubMed Web of Science ®Google Scholar
• Fukuhara T, Hosoya T, Shimizu S, et al. Utilization of host SR protein kinases and RNA-splicing machinery during viral replication. Proc Natl Acad Sci U S A. 2006;103(30):11329–11333. doi: 10.1073/pnas.0604616103
   PubMed Web of Science ®Google Scholar
• Bennett CF, Swayze EE. RNA targeting therapeutics: molecular mechanisms of antisense oligonucleotides as a therapeutic platform. Annu Rev Pharmacol Toxicol. 2010;50(1):259–293. doi: 10.1146/annurev.pharmtox.010909.105654
   PubMedGoogle Scholar
• Dominski Z, Kole R. Restoration of correct splicing in thalassemic pre-mRNA by antisense oligonucleotides. Proc Natl Acad Sci U S A. 1993;90(18):8673–8677. doi: 10.1073/pnas.90.18.8673
   PubMed Web of Science ®Google Scholar
• Arechavala-Gomeza V, Anthony K, Morgan J, et al. Antisense oligonucleotide-mediated exon skipping for Duchenne muscular dystrophy: progress and challenges. Curr Gene Ther. 2012;12(3):152–160. doi: 10.2174/156652312800840621
   PubMed Web of Science ®Google Scholar
• Rigo F, Hua Y, Krainer AR, et al. Antisense-based therapy for the treatment of spinal muscular atrophy. J Cell Bio. 2012;199(1):21–25. doi: 10.1083/jcb.201207087
   PubMed Web of Science ®Google Scholar
• Kim YJ, Krainer AR. Antisense oligonucleotide therapeutics for cystic fibrosis: recent developments and perspectives. Mol Cells. 2023;46(1):10–20. doi: 10.14348/molcells.2023.2172
   PubMed Web of Science ®Google Scholar
• Roberts TC, Langer R, Wood MJA. Advances in oligonucleotide drug delivery. Nat Rev Drug Discov. 2020;19(10):673–694. 2020 19:10. doi: 10.1038/s41573-020-0075-7
   PubMed Web of Science ®Google Scholar
• Lee JM, Nobumori C, Tu Y, et al. Modulation of LMNA splicing as a strategy to treat prelamin a diseases. J Clin Invest. 2016;126(4):1592–1602. doi: 10.1172/JCI85908
   PubMed Web of Science ®Google Scholar
• Gonzalo S, Kreienkamp R, Askjaer P. Hutchinson-Gilford progeria syndrome: a premature aging disease caused by LMNA gene mutations. Ageing Res Rev. 2017;33:18. doi: 10.1016/j.arr.2016.06.007
   PubMed Web of Science ®Google Scholar
• Scaffidi P, Misteli T. Reversal of the cellular phenotype in the premature aging disease Hutchinson-Gilford progeria syndrome. Nat Med. 2005;11(4):440–445. 2005 11:4. doi: 10.1038/nm1204
   PubMed Web of Science ®Google Scholar
• Osorio FG, Navarro CL, Cadiñanos J, et al. Splicing-directed therapy in a new mouse model of human accelerated aging. Sci Transl Med. 2011;3(106):3. doi: 10.1126/scitranslmed.3002847
   Web of Science ®Google Scholar
• Aartsma-Rus A, Van Ommen GJB. Antisense-mediated exon skipping: a versatile tool with therapeutic and research applications. RNA. 2007;13(10):1609. doi: 10.1261/rna.653607
   PubMed Web of Science ®Google Scholar
• Arechavala-Gomeza V, Khoo B, Aartsma-Rus A. Splicing modulation therapy in the treatment of genetic diseases. Appl Clin Genet. 2014;7:245–252. doi: 10.2147/TACG.S71506
   PubMedGoogle Scholar
• McManus MT, Sharp PA. Gene silencing in mammals by small interfering RNAs. Nat Rev Genet. 2002;3(10):737–747. doi: 10.1038/nrg908
   PubMed Web of Science ®Google Scholar
• Grünweller A, Wyszko E, Bieber B, et al. Comparison of different antisense strategies in mammalian cells using locked nucleic acids, 2’-O-methyl RNA, phosphorothioates and small interfering RNA. Nucleic Acids Res. 2003;31(12):3185–3193. doi: 10.1093/nar/gkg409
   PubMed Web of Science ®Google Scholar
• McCaffrey AP, Meuse L, Karimi M, et al. A potent and specific morpholino antisense inhibitor of hepatitis C translation in mice. Hepatology. 2003;38(2):503–508. doi: 10.1053/jhep.2003.50330
   PubMed Web of Science ®Google Scholar
• Du M, Jillette N, Zhu JJ, et al. CRISPR artificial splicing factors. Nat Commun. 2020;11(1):1–11. doi: 10.1038/s41467-020-16806-4
   PubMed Web of Science ®Google Scholar
• Havens MA, Hastings ML. Splice-switching antisense oligonucleotides as therapeutic drugs. Nucleic Acids Res. 2016;44(14):6549. 2023. doi: 10.1093/nar/gkw533
   PubMed Web of Science ®Google Scholar
• Denichenko P, Mogilevsky M, Cléry A, et al. Specific inhibition of splicing factor activity by decoy RNA oligonucleotides. Nat Commun. 2019;10(1):10. doi: 10.1038/s41467-019-09523-0
   PubMedGoogle Scholar
• Roberts TC, Langer R, Wood MJA. Advances in oligonucleotide drug delivery. Nat Rev Drug Discov. 2020;19(10):673–694. doi: 10.1038/s41573-020-0075-7
   PubMed Web of Science ®Google Scholar
• Bennett Frank C, Krainer AR, Cleveland DW. Antisense Oligonucleotide Therapies for Neurodegenerative Diseases. Annu Rev Neurosci. 2019;42(1):385. doi: 10.1146/annurev-neuro-070918-050501
   PubMedGoogle Scholar
• Gapinske M, Luu A, Winter J, et al. CRISPR-SKIP: programmable gene splicing with single base editors. Genome Biol. 2018;19(1):1–11. doi: 10.1186/s13059-018-1482-5
   PubMedGoogle Scholar
• Malakar P, Stein I, Saragovi A, et al. Long noncoding RNA MALAT1 regulates cancer glucose metabolism by enhancing mTOR-Mediated translation of TCF7L2. Cancer Res. 2019;79(10):2480–2493. doi: 10.1158/0008-5472.CAN-18-1432
   PubMed Web of Science ®Google Scholar
• Huarte M, Guttman M, Feldser D, et al. A large intergenic noncoding RNA induced by p53 mediates global gene repression in the p53 response. Cell. 2010;142(3):409–419. doi: 10.1016/j.cell.2010.06.040
   PubMed Web of Science ®Google Scholar
• Khalil AM, Guttman M, Huarte M, et al. Many human large intergenic noncoding RNAs associate with chromatin-modifying complexes and affect gene expression. Proc Natl Acad Sci U S A. 2009;106(28):11667–11672. doi: 10.1073/pnas.0904715106
   PubMed Web of Science ®Google Scholar
• Guttman M, Donaghey J, Carey BW, et al. lincRNAs act in the circuitry controlling pluripotency and differentiation. Nature. 2011;477(7364):295–300. 2011 477:7364. doi: 10.1038/nature10398
   PubMed Web of Science ®Google Scholar
• Liu SJ, Lim DA. Modulating the expression of long non-coding RNAs for functional studies. EMBO Rep. 2018;19(12). doi: 10.15252/embr.201846955
   Web of Science ®Google Scholar
• Doench JG, Petersen CP, Sharp PA. siRNAs can function as miRNAs. Genes Dev. 2003;17(4):438. doi: 10.1101/gad.1064703
   PubMed Web of Science ®Google Scholar
• Zeng Y, Yi R, Cullen BR. MicroRNAs and small interfering RNAs can inhibit mRNA expression by similar mechanisms. Proc Natl Acad Sci U S A. 2003;100(17):9779–9784. doi: 10.1073/pnas.1630797100
   PubMed Web of Science ®Google Scholar
• Bartel DP. MicroRNAs: Genomics, Biogenesis, Mechanism, and Function. Cell. 2004;116(2):281–297. doi: 10.1016/S0092-8674(04)00045-5
   PubMed Web of Science ®Google Scholar
• Zeng Y, Cullen BR. RNA interference in human cells is restricted to the cytoplasm. RNA. 2002;8(7):855. doi: 10.1017/S1355838202020071
   PubMed Web of Science ®Google Scholar
• Lennox KA, Behlke MA. Cellular localization of long non-coding RNAs affects silencing by RNAi more than by antisense oligonucleotides. Nucleic Acids Res. 2016;44(2):863–877. doi: 10.1093/nar/gkv1206
   PubMed Web of Science ®Google Scholar
• Gagnon KT, Li L, Chu Y, et al. RNAi factors are present and active in human cell nuclei. Cell Rep. 2014;6(1):211–221. doi: 10.1016/j.celrep.2013.12.013
   PubMed Web of Science ®Google Scholar
• McKay RA, Miraglia LJ, Cummins LL, et al. Characterization of a potent and specific class of antisense oligonucleotide inhibitor of human protein kinase C-α expression. J Biol Chem. 1999;274(3):1715–1722. doi: 10.1074/jbc.274.3.1715
   PubMed Web of Science ®Google Scholar
• Kurreck J, Wyszko E, Gillen C, et al. Design of antisense oligonucleotides stabilized by locked nucleic acids. Nucleic Acids Res. 2002;30(9):1911–1918. doi: 10.1093/nar/30.9.1911
   PubMed Web of Science ®Google Scholar
• Laxton C, Brady K, Moschos S, et al. Selection, optimization, and pharmacokinetic properties of a novel, potent antiviral locked nucleic acid-based antisense oligomer targeting hepatitis C virus internal ribosome entry site. Antimicrob Agents Chemother. 2011;55(7):3105–3114. doi: 10.1128/AAC.00222-11
   PubMed Web of Science ®Google Scholar
• Geary RS, Norris D, Yu R, et al. Pharmacokinetics, biodistribution and cell uptake of antisense oligonucleotides. Adv Drug Deliv Rev. 2015;87:46–51. doi: 10.1016/j.addr.2015.01.008
   PubMed Web of Science ®Google Scholar
• Shen X, Corey DR. Chemistry, mechanism and clinical status of antisense oligonucleotides and duplex RNAs. Nucleic Acids Res. 2018;46(4):1584–1600. doi: 10.1093/nar/gkx1239
   PubMed Web of Science ®Google Scholar
• Crooke ST. Molecular Mechanisms of Antisense Oligonucleotides. Nucleic Acid Ther. 2017;27(2):70–77. doi: 10.1089/nat.2016.0656
   PubMed Web of Science ®Google Scholar
• Arun G, Diermeier SD, Spector DL. Therapeutic Targeting of Long Non-Coding RNAs in Cancer. Trends Mol Med. 2018;24(3):257. doi: 10.1016/j.molmed.2018.01.001
   PubMed Web of Science ®Google Scholar
• Arun G, Diermeier S, Akerman M, et al. Differentiation of mammary tumors and reduction in metastasis upon Malat1 lncRNA loss. Genes Dev. 2016;30(1):34–51. doi: 10.1101/gad.270959.115
   PubMed Web of Science ®Google Scholar
• Shmakov S, Abudayyeh OO, Makarova KS, et al. Discovery and functional characterization of diverse class 2 CRISPR-Cas systems. Mol Cell. 2015;60(3):385–397. doi: 10.1016/j.molcel.2015.10.008
   PubMed Web of Science ®Google Scholar
• Zhang Z, Chen J, Zhu Z, et al. CRISPR-Cas13-mediated knockdown of lncRNA-GACAT3 inhibited cell proliferation and motility, and induced apoptosis by increasing p21, Bax, and E-Cadherin expression in bladder cancer. Front Mol Biosci. 2020;7:627774. doi: 10.3389/fmolb.2020.627774
   PubMed Web of Science ®Google Scholar
• Xu D, Cai Y, Tang L, et al. A CRISPR/Cas13-based approach demonstrates biological relevance of vlinc class of long non-coding RNAs in anticancer drug response. Sci Rep. 2020;10(1):1–13. 2020 10:1. doi: 10.1038/s41598-020-58104-5
   PubMed Web of Science ®Google Scholar
• Maxwell IH, Harrison GS, Wood WM, et al. A DNA cassette containing a trimerized SV40 polyadenylation signal which efficiently blocks spurious plasmid-initiated transcription. Biotechniques. 1989;7:276–280.
   PubMed Web of Science ®Google Scholar
• Beaulieu YB, Kleinman CL, Landry-Voyer AM, et al. Polyadenylation-dependent control of Long noncoding RNA expression by the Poly(A)-binding protein nuclear 1. PLoS Genet. 2012;8(11):1003078. doi: 10.1371/journal.pgen.1003078
   PubMed Web of Science ®Google Scholar
• Gilbert LA, Horlbeck MA, Adamson B, et al. Genome-scale CRISPR-Mediated control of gene repression and activation. Cell. 2014;159(3):647–661. doi: 10.1016/j.cell.2014.09.029
   PubMed Web of Science ®Google Scholar
• Gilbert LA, Larson MH, Morsut L, et al. XCRISPR-mediated modular RNA-guided regulation of transcription in eukaryotes. Cell. 2013;154(2):442. doi: 10.1016/j.cell.2013.06.044
   PubMed Web of Science ®Google Scholar
• Yang F, Song HX, Xian YS, et al. Repression of the long noncoding RNA-LET by histone deacetylase 3 contributes to hypoxia-mediated metastasis. Mol Cell. 2013;49(6):1083–1096. doi: 10.1016/j.molcel.2013.01.010
   PubMed Web of Science ®Google Scholar
• Grote P, Wittler L, Hendrix D, et al. The tissue-specific lncRNA fendrr is an essential regulator of heart and body wall development in the mouse. Dev Cell. 2013;24(2):206–214. doi: 10.1016/j.devcel.2012.12.012
   PubMed Web of Science ®Google Scholar
• Ulitsky I, Shkumatava A, Jan CH, et al. Conserved function of lincRNAs in vertebrate embryonic development despite rapid sequence evolution. Cell. 2011;147(7):1537–1550. doi: 10.1016/j.cell.2011.11.055
   PubMed Web of Science ®Google Scholar
• Xiang JF, Yin QF, Chen T, et al. Human colorectal cancer-specific CCAT1-L lncRNA regulates long-range chromatin interactions at the MYC locus. Cell Res. 2014;24(5):513–531. doi: 10.1038/cr.2014.35
   PubMed Web of Science ®Google Scholar
• Konermann S, Brigham MD, Trevino AE, et al. Genome-scale transcriptional activation by an engineered CRISPR-Cas9 complex. Nature. 2014;517:7536. doi: 10.1038/nature14136
   Web of Science ®Google Scholar
• Shechner DM, Hacisuleyman E, Younger ST, et al. Multiplexable, locus-specific targeting of long RNAs with CRISPR-Display. Nat Methods. 2015;12(7):664–670. doi: 10.1038/nmeth.3433
   PubMed Web of Science ®Google Scholar
• Xu X, Yang D, Ding JH, et al. ASF/SF2-regulated CaMKIIδ alternative splicing temporally reprograms excitation-contraction coupling in cardiac muscle. Cell. 2005;120(1):59–72. doi: 10.1016/j.cell.2004.11.036
   PubMed Web of Science ®Google Scholar
• Wang HY, Xu X, Ding JH, et al. SC35 plays a role in T cell development and alternative splicing of CD45. Mol Cell. 2001;7(2):331–342. doi: 10.1016/S1097-2765(01)00181-2
   PubMed Web of Science ®Google Scholar
• Jumaa H, Wei G, Nielsen PJ. Blastocyst formation is blocked in mouse embryos lacking the splicing factor SRp20. Curr Biol. 1999;9(16):899–902. doi: 10.1016/S0960-9822(99)80394-7
   PubMed Web of Science ®Google Scholar
• Fernandes JCR, Acuña SM, Aoki JI, et al. Long non-coding RNAs in the regulation of gene expression: physiology and disease. Noncoding RNA. 2019;5(1):17. doi: 10.3390/ncrna5010017
   PubMed Web of Science ®Google Scholar