Department of Molecular Biology, Princeton University, Princeton, New Jersey 08544.
Mol Cell Proteomics. 2014 Jan;13(1):73-83. doi: 10.1074/mcp.M113.031377. Epub 2013 Oct 10.
It has been shown that SIRT7 regulates rDNA transcription and that reduced SIRT7 levels inhibit tumor growth. This anti-tumor effect could be due to reduced Pol I activity and perturbed ribosome biogenesis. In this study, using pulse labeling with RNA and amino acid analogs, we found that SIRT7 knockdown efficiently suppressed both RNA and protein synthesis. Surprisingly, SIRT7 knockdown preferentially inhibited protein synthesis over rDNA transcription, whereas the levels of both were reduced to similar extents following Pol I knockdown. Using an affinity purification mass spectrometry approach and functional analyses of the resulting SIRT7 interactome, we identified and validated SIRT7 interactions with proteins involved in ribosomal biogenesis. Indeed, SIRT7 co-fractionated with monoribosomes within a sucrose gradient. Using reciprocal isolations, we determined that SIRT7 interacts specifically with mTOR and GTF3C1, a component of the Pol III transcription factor TFIIIC2 complex. Further studies found that SIRT7 knockdown triggered an increase in the levels of LC3B-II, an autophagosome marker, suggesting a link between SIRT7 and the mTOR pathway. Additionally, we provide several lines of evidence that SIRT7 plays a role in modulating Pol III function. Immunoaffinity purification of SIRT7-GFP from a nuclear fraction demonstrated specific SIRT7 interaction with five out of six components of the TFIIIC2 complex, but not with the TFIIIA or TFIIIB complex, the former of which is required for Pol III-dependent transcription of tRNA genes. ChIP assays showed SIRT7 localization to the Pol III targeting genes, and SIRT7 knockdown triggered a reduction in tRNA levels. Taken together, these data suggest that SIRT7 may regulate Pol III transcription through mTOR and the TFIIIC2 complex. We propose that SIRT7 is involved in multiple pathways involved in ribosome biogenesis, and we hypothesize that its down-regulation may contribute to an antitumor effect, partly through the inhibition of protein synthesis.
已经表明,SIRT7 调节 rDNA 转录,并且 SIRT7 水平降低会抑制肿瘤生长。这种抗肿瘤作用可能是由于 Pol I 活性降低和核糖体生物发生受到干扰。在这项研究中,我们使用 RNA 和氨基酸类似物脉冲标记,发现 SIRT7 敲低有效地抑制了 RNA 和蛋白质的合成。令人惊讶的是,SIRT7 敲低优先抑制蛋白质合成而不是 rDNA 转录,而在 Pol I 敲低后,两者的水平都降低到相似的程度。使用亲和纯化质谱方法和对所得 SIRT7 相互作用组的功能分析,我们鉴定并验证了 SIRT7 与参与核糖体生物发生的蛋白质的相互作用。事实上,SIRT7 与蔗糖梯度中的单核糖体共分馏。通过相互分离,我们确定 SIRT7 与 mTOR 和 GTF3C1(Pol III 转录因子 TFIIIC2 复合物的一个组成部分)特异性相互作用。进一步的研究发现,SIRT7 敲低会导致 LC3B-II 水平升高,这是自噬体的标志物,表明 SIRT7 与 mTOR 途径之间存在联系。此外,我们提供了几条证据表明 SIRT7 在调节 Pol III 功能方面发挥作用。从核部分的 SIRT7-GFP 免疫亲和纯化证明了 SIRT7 与 TFIIIC2 复合物的六个成分中的五个特异性相互作用,但与 TFIIIA 或 TFIIIB 复合物没有相互作用,前者是 Pol III 依赖的 tRNA 基因转录所必需的。ChIP 测定显示 SIRT7 定位于 Pol III 靶向基因,并且 SIRT7 敲低会导致 tRNA 水平降低。总之,这些数据表明 SIRT7 可能通过 mTOR 和 TFIIIC2 复合物调节 Pol III 转录。我们提出 SIRT7 参与核糖体生物发生的多个途径,并且我们假设其下调可能部分通过抑制蛋白质合成来发挥抗肿瘤作用。