细胞核肌动蛋白参与基因转录调控的分子机制研究
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摘要
肌动蛋白是一类高度保守的蛋白质,存在于所有的真核细胞中,参与细胞分裂、运动、迁移、形态的维持、生长等许多重要生理活动。有研究指出,肌动蛋白也存在于细胞核内。但由于肌动蛋白是丰富的胞质蛋白,致使这一结论受到质疑。后来,依赖生物化学、免疫荧光、免疫电子显微镜和激光共聚焦显微镜等实验技术,证明肌动蛋白普遍存在于各类细胞的细胞核中。
虽然肌动蛋白在细胞质中的功能已经被研究的很透彻,但它在细胞核内的功能还不清楚。有实验指出,肌动蛋白在核运输方面发挥作用,可能参与RNA运输及mRNA加工。肌动蛋白还能与protein 4.1共定位,可能参与核组装过程。此外,肌动蛋白还能与染色质改构复合物和乙酰转移酶复合物相互作用,在染色质结构的调整方面也发挥重要作用。越来越多的证据表明肌动蛋白也能参与基因转录的调控。在早期的实验中,微注射肌动蛋白的抗体或结合蛋白能抑制灯刷染色体的转录。肌动蛋白还能与RNA聚合酶II共纯化,对RNA聚合酶II体外的有效转录起到作用。最近有实验指出,肌动蛋白能被募集到基因的启动子区,是转录前起始复合物的成分。但肌动蛋白在此复合物中的作用还有待于进一步研究。
本文通过实验证明,RNA解旋酶A是转录前起始复合物中能够与肌动蛋白相互作用的蛋白质。我们首先通过免疫共沉淀和免疫荧光标记技术证明RNA解旋酶A在体内可以与肌动蛋白相互作用。然后利用GST pull-down实验证实RNA解旋酶A主要是通过其RGG区域与肌动蛋白作用,并且这一负显性突变体能够抑制RNA聚合酶II募集到转录前起始复合物中。此外,过表达或干涉RNA解旋酶A都能够影响RNA聚合酶II与肌动蛋白之间的相互作用,并且也能影响肌动蛋白所参与的基因表达。综上所述,RNA解旋酶A对肌动蛋白参与的转录过程起到重要调节作用。
Actin is a highly conserved protein, which exists in almost all eukaryotic cells and has been proved to be essential for many cellular functions such as movement, morphology, growth, cytokinesis as well as other crucial events. Over the years actin has been reported to exist in the nucleus, but these results have always been disputed because they could have reflected the contamination due to the high concentration of actin in the cytoplasm. Recent years, based upon ultrstructural, histochemical, and immunofluorescence studies, the presence of actin in the nucleus has been confirmed in different cell types.
In contrast to its functions in cytoplasm, the knowledge concerning the role of actin in nucleus is still limited. It has been proposed that actin plays an important role in nuclear export, and actin might be involved in RNA transport and indirectly participate in mRNA processing. It was reported recently that nuclear actin co-localized with protein 4.1, linking actin to nuclear assembly processes. Furthermore, nuclear actin has been found to be associated with chromatin remodeling and histone acetyltransferase complexes, suggesting a role for actin in chromatin remodeling. A number of studies have suggested a role for actin in transcription. In early studies, it had been shown that injection of antibodies against actin or actin-binding proteins in amphibian oocyte nuclei could block the transcription of lampbrush chromosomes. Actin was also reported to co-purify with RNA polymerase II and to be required for efficient in vitro transcription by RNA polymerase II. Recently, it was reported that actin could be recruited to the promoter region of the gene, and is a component of pre-initiation complexes (PICs). However, as a part of PICs, the role of actin in this complex remains to be elucidated.
Here, we identified RHA (RNA helicase A) as an actin-interacting protein in PICs. Using immunoprecipitation and immunofluorescence techniques, we showed that RHA associated withβ-actin in the nucleus. GST pull-down assay using different deletion mutants revealed that RGG region of RHA was responsible for the interaction withβ-actin, and this dominant-negative mutant reduced the recruitment of Pol II (RNA polymerase II) into PICs. Moreover, overexpression or depletion of RHA could influence the interaction of Pol II withβ-actin andβ-actin-involved gene transcription regulation. These data suggest that RHA acts as a bridge factor linking nuclearβ-actin with Pol II.
虽然肌动蛋白在细胞质中的功能已经被研究的很透彻,但它在细胞核内的功能还不清楚。有实验指出,肌动蛋白在核运输方面发挥作用,可能参与RNA运输及mRNA加工。肌动蛋白还能与protein 4.1共定位,可能参与核组装过程。此外,肌动蛋白还能与染色质改构复合物和乙酰转移酶复合物相互作用,在染色质结构的调整方面也发挥重要作用。越来越多的证据表明肌动蛋白也能参与基因转录的调控。在早期的实验中,微注射肌动蛋白的抗体或结合蛋白能抑制灯刷染色体的转录。肌动蛋白还能与RNA聚合酶II共纯化,对RNA聚合酶II体外的有效转录起到作用。最近有实验指出,肌动蛋白能被募集到基因的启动子区,是转录前起始复合物的成分。但肌动蛋白在此复合物中的作用还有待于进一步研究。
本文通过实验证明,RNA解旋酶A是转录前起始复合物中能够与肌动蛋白相互作用的蛋白质。我们首先通过免疫共沉淀和免疫荧光标记技术证明RNA解旋酶A在体内可以与肌动蛋白相互作用。然后利用GST pull-down实验证实RNA解旋酶A主要是通过其RGG区域与肌动蛋白作用,并且这一负显性突变体能够抑制RNA聚合酶II募集到转录前起始复合物中。此外,过表达或干涉RNA解旋酶A都能够影响RNA聚合酶II与肌动蛋白之间的相互作用,并且也能影响肌动蛋白所参与的基因表达。综上所述,RNA解旋酶A对肌动蛋白参与的转录过程起到重要调节作用。
Actin is a highly conserved protein, which exists in almost all eukaryotic cells and has been proved to be essential for many cellular functions such as movement, morphology, growth, cytokinesis as well as other crucial events. Over the years actin has been reported to exist in the nucleus, but these results have always been disputed because they could have reflected the contamination due to the high concentration of actin in the cytoplasm. Recent years, based upon ultrstructural, histochemical, and immunofluorescence studies, the presence of actin in the nucleus has been confirmed in different cell types.
In contrast to its functions in cytoplasm, the knowledge concerning the role of actin in nucleus is still limited. It has been proposed that actin plays an important role in nuclear export, and actin might be involved in RNA transport and indirectly participate in mRNA processing. It was reported recently that nuclear actin co-localized with protein 4.1, linking actin to nuclear assembly processes. Furthermore, nuclear actin has been found to be associated with chromatin remodeling and histone acetyltransferase complexes, suggesting a role for actin in chromatin remodeling. A number of studies have suggested a role for actin in transcription. In early studies, it had been shown that injection of antibodies against actin or actin-binding proteins in amphibian oocyte nuclei could block the transcription of lampbrush chromosomes. Actin was also reported to co-purify with RNA polymerase II and to be required for efficient in vitro transcription by RNA polymerase II. Recently, it was reported that actin could be recruited to the promoter region of the gene, and is a component of pre-initiation complexes (PICs). However, as a part of PICs, the role of actin in this complex remains to be elucidated.
Here, we identified RHA (RNA helicase A) as an actin-interacting protein in PICs. Using immunoprecipitation and immunofluorescence techniques, we showed that RHA associated withβ-actin in the nucleus. GST pull-down assay using different deletion mutants revealed that RGG region of RHA was responsible for the interaction withβ-actin, and this dominant-negative mutant reduced the recruitment of Pol II (RNA polymerase II) into PICs. Moreover, overexpression or depletion of RHA could influence the interaction of Pol II withβ-actin andβ-actin-involved gene transcription regulation. These data suggest that RHA acts as a bridge factor linking nuclearβ-actin with Pol II.
引文
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