Tsukuda Toyoko, Fleming Alastair B, Nickoloff Jac A, Osley Mary Ann
Department of Molecular Genetics and Microbiology, University of New Mexico School of Medicine, 915 Camino de Salud Albuquerque, New Mexico 87131, USA.
Nature. 2005 Nov 17;438(7066):379-83. doi: 10.1038/nature04148.
The repair of DNA double-strand breaks (DSBs) is crucial for maintaining genome stability. Eukaryotic cells repair DSBs by both non-homologous end joining and homologous recombination. How chromatin structure is altered in response to DSBs and how such alterations influence DSB repair processes are important issues. In vertebrates, phosphorylation of the histone variant H2A.X occurs rapidly after DSB formation, spreads over megabase chromatin domains, and is required for stable accumulation of repair proteins at damage foci. In Saccharomyces cerevisiae, phosphorylation of the two principal H2A species is also signalled by DSB formation, which spreads approximately 40 kb in either direction from the DSB. Here we show that near a DSB phosphorylation of H2A is followed by loss of histones H2B and H3 and increased sensitivity of chromatin to digestion by micrococcal nuclease; however, phosphorylation of H2A and nucleosome loss occur independently. The DNA damage sensor MRX is required for histone loss, which also depends on INO80, a nucleosome remodelling complex. The repair protein Rad51 (ref. 6) shows delayed recruitment to DSBs in the absence of histone loss, suggesting that MRX-dependent nucleosome remodelling regulates the accessibility of factors directly involved in DNA repair by homologous recombination. Thus, MRX may regulate two pathways of chromatin changes: nucleosome displacement for efficient recruitment of homologous recombination proteins; and phosphorylation of H2A, which modulates checkpoint responses to DNA damage.
DNA双链断裂(DSB)的修复对于维持基因组稳定性至关重要。真核细胞通过非同源末端连接和同源重组两种方式修复DSB。染色质结构如何响应DSB而改变,以及这种改变如何影响DSB修复过程是重要的问题。在脊椎动物中,组蛋白变体H2A.X的磷酸化在DSB形成后迅速发生,扩散到兆碱基染色质结构域,并且是修复蛋白在损伤位点稳定积累所必需的。在酿酒酵母中,两种主要H2A物种的磷酸化也由DSB形成发出信号,其从DSB向任一方向扩散约40 kb。在这里,我们表明在DSB附近,H2A磷酸化之后是组蛋白H2B和H3的丢失以及染色质对微球菌核酸酶消化的敏感性增加;然而,H2A磷酸化和核小体丢失是独立发生的。DNA损伤传感器MRX是组蛋白丢失所必需的,这也依赖于INO80,一种核小体重塑复合物。在没有组蛋白丢失的情况下,修复蛋白Rad51(参考文献6)显示出对DSB的延迟募集,这表明MRX依赖性核小体重塑通过同源重组调节直接参与DNA修复的因子的可及性。因此,MRX可能调节染色质变化的两条途径:核小体置换以有效募集同源重组蛋白;以及H2A磷酸化,其调节对DNA损伤的检查点反应。