抗肿瘤药物NB-506对富含丝氨酸和精氨酸的剪接因子磷酸化、剪接体组装及剪接的特异性抑制作用
Specific inhibition of serine- and arginine-rich splicing factors phosphorylation, spliceosome assembly, and splicing by the antitumor drug NB-506.
作者信息
Pilch B, Allemand E, Facompré M, Bailly C, Riou J F, Soret J, Tazi J
机构信息
Institut de Génétique Moléculaire, UMR 5535, Centre National de la Recherché Scientifique, IFR 24, Université de Montpellier II, 34293 Montpellier Cedex, France.
出版信息
Cancer Res. 2001 Sep 15;61(18):6876-84.
Specific phosphorylation of serine- and arginine-rich pre-mRNA splicing factors (SR proteins) is one of the key determinants regulating splicing events. Several kinases involved in SR protein phosphorylation have been identified and characterized, among which human DNA topoisomerase I is known to have DNA-relaxing activity. In this study, we have investigated the mechanism of splicing inhibition by a glycosylated indolocarbazole derivative (NB-506), a potent inhibitor of both kinase and relaxing activities of topoisomerase I. NB-506 completely inhibits the capacity of topoisomerase I to phosphorylate, in vitro, the human splicing factor 2/alternative splicing factor (SF2/ASF). This inhibition is specific, because NB-506 does not demonstrate activity against other kinases known to phosphorylate SF2/ASF such as SR protein kinase 1 and cdc2 kinase. Importantly, HeLa nuclear extracts competent in splicing but not splicing-deficient cytoplasmic S100 extracts treated with the drug fail to phosphorylate SF2/ASF and to support splicing of pre-mRNA substrates containing SF2/ASF-target sequences. Native gel analysis of splicing complexes revealed that the drug affects the formation of the spliceosome, a dynamic ribonucleoprotein structure where splicing takes place. In the presence of the drug, neither pre-spliceosome nor spliceosome is formed, demonstrating that splicing inhibition occurs at early steps of spliceosome assembly. Splicing inhibition can be relieved by adding phosphorylated SF2/ASF, showing that extracts treated with NB-506 lack a phosphorylating activity required for splicing. Moreover, NB-506 has a cytotoxic effect on murine P388 leukemia cells but not on P388CPT5 camptothecin-resistant cells that carry two point mutations in conserved regions of topoisomerase I gene (Gly361Val and Asp709Tyr). After drug treatment, P388 cells accumulated hypophosphorylated forms of SR proteins and polyadenylated RNA in the nucleus. In contrast, neither SR protein phosphorylation nor polyadenylated mRNA distribution was affected in P388 CPT5-treated cells. Consistently, NB506 treatment altered the mRNA levels and/or splicing pattern of several tested genes (Bcl-X, CD 44, SC35, and Sty) in P388 cells but not in P388 CPT5 cells. The study shows for the first time that indolocarbazole drugs targeting topoisomerase I can affect gene expression by modulating pre-mRNA splicing through inhibition of SR proteins phosphorylation.
富含丝氨酸和精氨酸的前体mRNA剪接因子(SR蛋白)的特异性磷酸化是调节剪接事件的关键决定因素之一。已鉴定并表征了几种参与SR蛋白磷酸化的激酶,其中人DNA拓扑异构酶I具有DNA松弛活性。在本研究中,我们研究了一种糖基化吲哚咔唑衍生物(NB-506)抑制剪接的机制,该衍生物是拓扑异构酶I激酶和松弛活性的有效抑制剂。NB-506在体外完全抑制拓扑异构酶I磷酸化人剪接因子2/可变剪接因子(SF2/ASF)的能力。这种抑制是特异性的,因为NB-506对其他已知可磷酸化SF2/ASF的激酶如SR蛋白激酶1和cdc2激酶没有活性。重要的是,用该药物处理的具有剪接能力的HeLa核提取物而非缺乏剪接能力的细胞质S100提取物不能磷酸化SF2/ASF,也不能支持含有SF2/ASF靶序列的前体mRNA底物的剪接。对剪接复合物的天然凝胶分析表明,该药物影响剪接体的形成,剪接体是发生剪接的动态核糖核蛋白结构。在药物存在下,既不形成前剪接体也不形成剪接体,这表明剪接抑制发生在剪接体组装的早期步骤。通过添加磷酸化的SF2/ASF可以解除剪接抑制,这表明用NB-506处理的提取物缺乏剪接所需的磷酸化活性。此外,NB-506对小鼠P388白血病细胞具有细胞毒性作用,但对在拓扑异构酶I基因保守区域携带两个点突变(Gly361Val和Asp709Tyr)的P388CPT5喜树碱抗性细胞没有作用。药物处理后,P388细胞在细胞核中积累了低磷酸化形式的SR蛋白和多聚腺苷酸化RNA。相比之下,P388 CPT5处理的细胞中SR蛋白磷酸化和多聚腺苷酸化mRNA分布均未受影响。一致地,NB506处理改变了P388细胞中几个测试基因(Bcl-X、CD 44、SC35和Sty)的mRNA水平和/或剪接模式,但在P388 CPT5细胞中没有改变。该研究首次表明,靶向拓扑异构酶I的吲哚咔唑药物可通过抑制SR蛋白磷酸化调节前体mRNA剪接来影响基因表达。