Department of Molecular and Cellular Biology, Roswell Park Cancer Institute, Buffalo, NY 14263, USA.
Proc Natl Acad Sci U S A. 2012 Sep 11;109(37):E2457-65. doi: 10.1073/pnas.1206274109. Epub 2012 Aug 15.
The circadian clock controls many physiological parameters including immune response to infectious agents, which is mediated by activation of the transcription factor NF-κB. It is widely accepted that circadian regulation is based on periodic changes in gene expression that are triggered by transcriptional activity of the CLOCK/BMAL1 complex. Through the use of a mouse model system we show that daily variations in the intensity of the NF-κB response to a variety of immunomodulators are mediated by core circadian protein CLOCK, which can up-regulate NF-κB-mediated transcription in the absence of BMAL1; moreover, BMAL1 counteracts the CLOCK-dependent increase in the activation of NF-κB-responsive genes. Consistent with its regulatory function, CLOCK is found in protein complexes with the p65 subunit of NF-κB, and its overexpression correlates with an increase in specific phosphorylated and acetylated transcriptionally active forms of p65. In addition, activation of NF-κB in response to immunostimuli in mouse embryonic fibroblasts and primary hepatocytes isolated from Clock-deficient mice is significantly reduced compared with WT cells, whereas Clock-Δ19 mutation, which reduces the transactivation capacity of CLOCK on E-box-containing circadian promoters, has no effect on the ability of CLOCK to up-regulate NF-κB-responsive promoters. These findings establish a molecular link between two essential determinants of the circadian and immune mechanisms, the transcription factors CLOCK and NF-κB, respectively.
生物钟控制着许多生理参数,包括对感染因子的免疫反应,而这种反应是由转录因子 NF-κB 的激活介导的。人们普遍认为,生物钟的调节是基于基因表达的周期性变化,而这种变化是由 CLOCK/BMAL1 复合物的转录活性触发的。通过使用小鼠模型系统,我们表明,对各种免疫调节剂的 NF-κB 反应强度的日变化是由核心生物钟蛋白 CLOCK 介导的,CLOCK 可以在没有 BMAL1 的情况下上调 NF-κB 介导的转录;此外,BMAL1 抵消了 CLOCK 依赖性的 NF-κB 反应基因激活的增加。与它的调节功能一致,CLOCK 与 NF-κB 的 p65 亚基存在于蛋白质复合物中,其过表达与特定磷酸化和乙酰化的转录活性形式的 p65 增加相关。此外,与 WT 细胞相比,Clock 缺陷型小鼠来源的胚胎成纤维细胞和原代肝细胞中免疫刺激物对 NF-κB 的激活显著降低,而 Clock-Δ19 突变降低了 CLOCK 对 E 盒包含的生物钟启动子的转录激活能力,对 CLOCK 上调 NF-κB 反应启动子的能力没有影响。这些发现确立了生物钟和免疫机制的两个基本决定因素,即转录因子 CLOCK 和 NF-κB 之间的分子联系。