裂殖酵母ING家族蛋白Png1p在DNA损伤应答过程中的功能研究
摘要
ING家族蛋白(INhibitor of Growth)作为一类肿瘤生长抑制因子,在几乎所有的真核细胞中都含有保守的同源蛋白,并广泛参与细胞凋亡,衰老,细胞周期调控,DNA损伤应答,细胞分化,肿瘤生长,激素信号传递,基因表达和染色质重构等多种过程。对ING家族蛋白参与DNA损伤修复的过程已经进行了许多的研究,但是由于ING家族蛋白参与DNA损伤修复的分子机制比较复杂,对于ING家族蛋白在DNA损伤信号传导通路上发挥的具体功能至今还未能完全有效解析。
裂殖酵母细胞含有两个ING家族蛋白：Png1p和Png2p。至今为止,在裂殖酵母细胞中还没有关于ING家族蛋白参与DNA损伤应答的报道。同源进化分析表明：Png1p和Png2p虽然同属ING家族,但两者位于系统发生树的不同分支上。进一步,我们通过同源重组的方法成功地敲除了两个基因的ORF,获得了突变体Δpng1和Δpng2。功能分析表明PNG1和PNG2在细胞生长和DNA损伤应答过程中执行了不同功能。
细胞生长和流式细胞术分析表明PNG1基因不仅是细胞正常生长所必需的,同时也是DNA损伤修复所必需的,PNG1的缺失使细胞阻滞在了S期。酵母双杂交,Pull-down实验,Co-IP实验,荧光共定位实验表明Png1p与组蛋白乙酰转移酶Mst1p密切相关,但两者可能没有直接的相互作用。TSA拯救实验和western blot检测表明Png1p参与了组蛋白H4的乙酰化修饰。PNG1基因的缺失使DNA损伤条件下裂殖酵母H4K5、K8、K12位的乙酰化水平下降,并且这种乙酰化程度的下调直接抑制了DNA损伤修复相关基因(如RAD22,ASF1,SPT3),染色质重构相关基因(如SNF5),细胞周期相关基因(如CDC20, CDC22, ENG1,AGN1)的表达。更进一步的研究发现,在PNG1缺失突变体中分别过量表达DNA损伤修复核心基因RAD22可以拯救PNG1缺失带来的DNA损伤敏感的表型。因此,PNG1很可能通过RAD22途径参与了DNA损伤应答过程。
DREB家族转录因子是一类参与调控植物体对干旱应答的重要的调节因子。这类转录因子可以特异性地结合到C-repeat/DRE顺式元件上(核心序列为A/GCCGAC),并以一种不依赖于ABA的方式,调控下游许多低温或干旱诱导基因的表达。在我们的研究中,我们在水稻中分离了3个新的水稻中的DREB家族转录因子：OsDREB1E、OsDREB1G和OsDREB2B。这三者与拟南芥中的DREB家族转录因子具有序列和结构上的高度同源性。酵母单杂交试验表明这三者可以特异性地结合到C-repeat/DRE顺式元件上。为了研究这三个基因在水稻中的功能,我们通过农杆菌介导的转基因方法将这三个基因在水稻中过量表达。对转基因水稻植株的分析表明,过表达OsDREB1G和OsDREB2B可以显著提高水稻对于干旱的耐受能力,但是过表达OsDREB1E只能稍微提高水稻对于干旱的耐受能力。这一实验结果表明,水稻中不同的DREB基因可能通过不同的方式参与干旱等压力应答的途径。
ING (INhibitor of Growth) tumor suppressor proteins are very important tumor inhibitors in species ranging from yeast to humans, and function in the process of cell apoptosis, senescence, cell cycle regulation, DNA damage response, cell proliferation, tumor growth, hormone signaling, gene transcription, chromatin remodeling and genome integrity. There are many studies about how ING family proteins function in the DNA damage response pathway. However, the molecular mechnism is so complicated that the blueprint of how ING family proteins function in the DNA damage signaling is still not very clear.
There are two homolog proteins in fission yeast, Png1p and Png2p. Until now, there is no report about how ING family proteins function in the DNA damage pathway in fission yeast. Phylogenic assay indicates although Png1p and Png2p belong to the same ING family, but they located at different branch. Here, we knock out PNG1 and PNG2 respectively. The functional analysis reveals that Png1p and Png2p have different functions in the process of cell growth and DNA damage reponse.
Cell growth and flow cytometry assay show PNG1 is not only required for normal cell growth, but also for DNA damage response.△png1 mutants get an intra-S-phase cell cycle arrest under DNA-damage induced conditions and are sensitive to DNA damage agents. Yeast-two-hybrid assay, Pull-down assay, Co-IP and fluorescent location assay indicate Png1p is collaborated with the fission yeast histone acetyltransferase Mst1p, but they might not have direct interaction。TSA rescue and western blot assay show Png1p is required for histone H4 acetylation at K5、K8、K12. The decrease of histone H4 acetylation at K5、K8、K12 leads to the down-regulation of many genes'expression, including DNA repair related genes (such as RAD22, ASF1, SPT3), chromatin remodeling related genes(such as SNF5), cell cycle related genes (such as CDC20, CDC22, ENG1, AGN1).
Further study shows over-expression of the DNA damage repair central gene RAD22 can recuse the DNA damage sensitivity in the△png1 cells, respectively. So, PNG1 might function in the DNA damage response pathway through RAD22.
The DREB transcription factors, which specifically interact with C-repeat/DRE (A/GCCGAC), play an important role in plant abiotic stress tolerance by controlling the expression of many cold or/and drought-inducible genes in an ABA-independent pathway. In our study, we have isolated three novel rice DREB genes, OsDREB1E, OsDREB1G, and OsDREB2B, which are homologous to Arabidopsis DREB genes. The yeast one-hybrid assay indicated that OsDREB1E, OsDREB1G, and OsDREB2B can specifically bind to the C-repeat/DRE element. To elucidate the function of respective OsDREB genes, we have stably introduced these to rice by agrobacterium-mediated transformation. Transgenic rice plants analysis revealed that over-expression of OsDREB1G and OsDREB2B in rice significantly improved their tolerance to water deficit stress, while over-expression of OsDREB1E could only slightly improved the tolerance to water deficit stress, suggesting that the OsDREBs might participate in the stress response pathway in different manners.
裂殖酵母细胞含有两个ING家族蛋白：Png1p和Png2p。至今为止,在裂殖酵母细胞中还没有关于ING家族蛋白参与DNA损伤应答的报道。同源进化分析表明：Png1p和Png2p虽然同属ING家族,但两者位于系统发生树的不同分支上。进一步,我们通过同源重组的方法成功地敲除了两个基因的ORF,获得了突变体Δpng1和Δpng2。功能分析表明PNG1和PNG2在细胞生长和DNA损伤应答过程中执行了不同功能。
细胞生长和流式细胞术分析表明PNG1基因不仅是细胞正常生长所必需的,同时也是DNA损伤修复所必需的,PNG1的缺失使细胞阻滞在了S期。酵母双杂交,Pull-down实验,Co-IP实验,荧光共定位实验表明Png1p与组蛋白乙酰转移酶Mst1p密切相关,但两者可能没有直接的相互作用。TSA拯救实验和western blot检测表明Png1p参与了组蛋白H4的乙酰化修饰。PNG1基因的缺失使DNA损伤条件下裂殖酵母H4K5、K8、K12位的乙酰化水平下降,并且这种乙酰化程度的下调直接抑制了DNA损伤修复相关基因(如RAD22,ASF1,SPT3),染色质重构相关基因(如SNF5),细胞周期相关基因(如CDC20, CDC22, ENG1,AGN1)的表达。更进一步的研究发现,在PNG1缺失突变体中分别过量表达DNA损伤修复核心基因RAD22可以拯救PNG1缺失带来的DNA损伤敏感的表型。因此,PNG1很可能通过RAD22途径参与了DNA损伤应答过程。
DREB家族转录因子是一类参与调控植物体对干旱应答的重要的调节因子。这类转录因子可以特异性地结合到C-repeat/DRE顺式元件上(核心序列为A/GCCGAC),并以一种不依赖于ABA的方式,调控下游许多低温或干旱诱导基因的表达。在我们的研究中,我们在水稻中分离了3个新的水稻中的DREB家族转录因子：OsDREB1E、OsDREB1G和OsDREB2B。这三者与拟南芥中的DREB家族转录因子具有序列和结构上的高度同源性。酵母单杂交试验表明这三者可以特异性地结合到C-repeat/DRE顺式元件上。为了研究这三个基因在水稻中的功能,我们通过农杆菌介导的转基因方法将这三个基因在水稻中过量表达。对转基因水稻植株的分析表明,过表达OsDREB1G和OsDREB2B可以显著提高水稻对于干旱的耐受能力,但是过表达OsDREB1E只能稍微提高水稻对于干旱的耐受能力。这一实验结果表明,水稻中不同的DREB基因可能通过不同的方式参与干旱等压力应答的途径。
ING (INhibitor of Growth) tumor suppressor proteins are very important tumor inhibitors in species ranging from yeast to humans, and function in the process of cell apoptosis, senescence, cell cycle regulation, DNA damage response, cell proliferation, tumor growth, hormone signaling, gene transcription, chromatin remodeling and genome integrity. There are many studies about how ING family proteins function in the DNA damage response pathway. However, the molecular mechnism is so complicated that the blueprint of how ING family proteins function in the DNA damage signaling is still not very clear.
There are two homolog proteins in fission yeast, Png1p and Png2p. Until now, there is no report about how ING family proteins function in the DNA damage pathway in fission yeast. Phylogenic assay indicates although Png1p and Png2p belong to the same ING family, but they located at different branch. Here, we knock out PNG1 and PNG2 respectively. The functional analysis reveals that Png1p and Png2p have different functions in the process of cell growth and DNA damage reponse.
Cell growth and flow cytometry assay show PNG1 is not only required for normal cell growth, but also for DNA damage response.△png1 mutants get an intra-S-phase cell cycle arrest under DNA-damage induced conditions and are sensitive to DNA damage agents. Yeast-two-hybrid assay, Pull-down assay, Co-IP and fluorescent location assay indicate Png1p is collaborated with the fission yeast histone acetyltransferase Mst1p, but they might not have direct interaction。TSA rescue and western blot assay show Png1p is required for histone H4 acetylation at K5、K8、K12. The decrease of histone H4 acetylation at K5、K8、K12 leads to the down-regulation of many genes'expression, including DNA repair related genes (such as RAD22, ASF1, SPT3), chromatin remodeling related genes(such as SNF5), cell cycle related genes (such as CDC20, CDC22, ENG1, AGN1).
Further study shows over-expression of the DNA damage repair central gene RAD22 can recuse the DNA damage sensitivity in the△png1 cells, respectively. So, PNG1 might function in the DNA damage response pathway through RAD22.
The DREB transcription factors, which specifically interact with C-repeat/DRE (A/GCCGAC), play an important role in plant abiotic stress tolerance by controlling the expression of many cold or/and drought-inducible genes in an ABA-independent pathway. In our study, we have isolated three novel rice DREB genes, OsDREB1E, OsDREB1G, and OsDREB2B, which are homologous to Arabidopsis DREB genes. The yeast one-hybrid assay indicated that OsDREB1E, OsDREB1G, and OsDREB2B can specifically bind to the C-repeat/DRE element. To elucidate the function of respective OsDREB genes, we have stably introduced these to rice by agrobacterium-mediated transformation. Transgenic rice plants analysis revealed that over-expression of OsDREB1G and OsDREB2B in rice significantly improved their tolerance to water deficit stress, while over-expression of OsDREB1E could only slightly improved the tolerance to water deficit stress, suggesting that the OsDREBs might participate in the stress response pathway in different manners.
引文
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