Siebel C W, Fresco L D, Rio D C
Whitehead Institute for Biomedical Research, Cambridge, Massachusetts 02142.
Genes Dev. 1992 Aug;6(8):1386-401. doi: 10.1101/gad.6.8.1386.
Somatic inhibition restricts splicing of the Drosophila P-element third intron (IVS3) to the germ line. We have exploited this simple system to provide a model for a mechanism of alternative pre-mRNA splicing. Biochemical complementation experiments revealed that Drosophila somatic extracts inhibited U1 snRNP binding to the 5' splice site. Using sensitive RNase protection and modification-interference assays, we found that U1 snRNP bound to a pseudo-5' splice site in the 5' exon and that multiprotein complexes bound to an adjacent site. Binding of these factors appeared to mediate the inhibitory effect, because mutations in the pseudo-5' splice sites blocked binding and activated splicing in vitro. Likewise, wild-type, but not mutant, 5' exon RNA titrated inhibitory factors away from the pre-mRNA and activated splicing. Thus, we have defined the pseudo-5' splice sites as crucial components of the regulatory element, correlated the inhibitory activity with specific RNA binding factors from Drosophila somatic cells, and provided a mechanistic description of somatic inhibition. Because the inhibitory activity involves general splicing functions such as protein recognition of 5' splice site sequences and changes in the distribution of bound U1 snRNP, our data may also provide insights into how splice sites are selected.
体细胞抑制将果蝇P因子的第三个内含子(IVS3)的剪接受限在生殖系中。我们利用这个简单的系统为可变前体mRNA剪接机制提供了一个模型。生化互补实验表明,果蝇体细胞提取物抑制U1 snRNP与5'剪接位点的结合。使用灵敏的核糖核酸酶保护和修饰干扰分析,我们发现U1 snRNP与5'外显子中的一个假5'剪接位点结合,并且多蛋白复合物与相邻位点结合。这些因子的结合似乎介导了抑制作用,因为假5'剪接位点的突变会阻断结合并在体外激活剪接。同样,野生型而非突变型的5'外显子RNA将抑制因子从前体mRNA上滴定下来并激活剪接。因此,我们将假5'剪接位点定义为调控元件的关键组成部分,将抑制活性与来自果蝇体细胞的特定RNA结合因子相关联,并提供了体细胞抑制的机制描述。由于抑制活性涉及一般的剪接功能,如蛋白质对5'剪接位点序列的识别以及结合的U1 snRNP分布的变化,我们的数据也可能为剪接位点的选择方式提供见解。