耐辐射球菌细胞内锰离子水平调控相关蛋白研究
摘要
细菌细胞内的金属离子动态平衡与细胞的生理生长具有密切关系,许多过渡价态金属离子,包括Mn(Ⅱ)、Zn(Ⅱ)、Ca(Ⅱ)等,不但可以作为蛋白的辅因子参与细胞的多种氧化还原反应,还可以作为信号因子参与基因组转录调控,影响细胞的毒性以及极端抗性等。因此细胞内金属离子的调控对于细菌维持正常生理活动至关重要,而关于细菌细胞内金属离子调控的研究也一直是微生物领域的热点。
作为极端微生物家族的重要成员,耐辐射球菌对于电离辐射、紫外线、化学诱变剂、干燥等具有极强抗性,而越来越多的研究表明,耐辐射球菌对于极端环境的适应能力得益于其超强的抗氧化能力,其细胞内积累的超高锰离子水平一直被认为是其最重要的抗氧化机制之一,因此耐辐射球菌是极好的研究细菌细胞锰离子代谢调控的模式生物,通过研究其锰离子调控机制,不但可以进一步了解耐辐射球菌的极端抗性机制,而且对于拓展细菌细胞金属离子调控机制也具有重要意义。本文主要通过细胞吸收及外排两个方面,研究了耐辐射球菌的细胞内锰离子动态调控机制,研究结果如下：
1.我们首次鉴定得到耐辐射球菌锰离子外排蛋白(Dr1236)。在构建了Dr1236的基因缺失突变株后,我们发现突变株对于锰离子的抗性下降,而且细胞内的锰离子水平显著升高,说明Dr1236在耐辐射球菌锰离子抗性及细胞内锰离子动态调控机制中,发挥了重要的作用。进一步的研究发现,突变株细胞内锰离子浓度升高以后,其电离辐射、紫外、过氧化氢抗性都明显增强,表明细胞内锰离子确实参与了细胞的极端抗性,而这种保护作用很可能是通过降低自由基对蛋白的攻击实现的。然而,虽然突变株细胞在积累了高水平锰离子之后,极端抗性有了不同程度的增加,但是细胞的生长周期却远远短于野生株,表明锰离子的过高积累对细胞的整体机能造成了损伤,这也从侧面反映出锰离子外排蛋白对于细胞的重要作用。
2.dr0865编码了耐辐射球菌唯一一个Fur同源蛋白,Fur蛋白在其它细菌中参与了锰离子吸收通道蛋白的转录调控,因此我们利用大肠杆菌表达系统体外表达了dr0865,并研究了其生化性质。生物信息学研究表明,Dr0865很可能包含了结合三个金属离子的结构,这种结构可能赋予了其独特的生物学功能。DNA结合活性实验表明,Dr0865可以与dr2283、dr2284、dr2524的启动子结合,而且结合活性与金属离子的浓度成正比,另一方面却不能与dr1709的启动子结合；β-半乳糖甘酶活性实验表明Dr0865与dr2283、dr2284、dr2523的启动子的结合可能促进了其转录,考虑到Dr0865的DNA结合活性依赖于金属离子的浓度,及Dr0865突变株的过氧化氢抗性增强,因此我们推测dr2283、dr2284、dr2523(?)(?)可能编码了一种铁离子外排通道蛋白,而其具体调控机制还需要进一步研究。
3.dr2539编码耐辐射球菌的DtxR/MntR家族蛋白,这类蛋白家族对于细胞锰离子或铁离子调控至关重要,通过在Dr2539 N-端融合表达助溶片段IF,我们成功得到Dr2539蛋白。DNA结合活性实验表明,Dr2539只能与dr1709的启动子结合,而且EMSA和p-半乳糖甘酶活性实验表明,蛋白的结合活性受到锰离子和铁离子的调控,表明Dr2539与此前所报道的MntR蛋白结构上可能存在很大差异,因此我们构建了Dr2539突变体,并将突变体补偿Dr2539突变株,包括C-D11M、C-H98Y、C-E101D、C-Tru,通过检测补偿株对于锰离子的抗性,发现C-D11M及C-Tru锰离子抗性相对于野生Dr2539补偿株明显下降,而C-E101D的抗性并没有明显变化；惊奇的是,H98Y补偿株对6-7Mm Mn(Ⅱ)极度敏感,这与此前所报道的DtxR激活模型一致,因此我们推测,dr2539所编码的MntR蛋白很可能利用一种类似于DtxR的激活模式,而且可能参与了细胞内锰离子与铁离子动态平衡的调控。
综上所述,在研究了耐辐射球菌细胞内锰离子外排和吸收系统后,我们的结果表明锰离子的确参与了耐辐射球菌的极端抗性机制,此外,金属离子调控蛋白Dr0865和Dr2539的生化活性表明,耐辐射球菌的金属离子调控机制与目前发现的细菌调控机制存在很大差异,而这种差异可能与其极端环境生存能力有关。我们的研究为下一步深入研究耐辐射球菌金属离子调控蛋白Dr0865和Dr2539结构奠定了基础。
Homeostasis of metal ions is closely related to the physiological processes of growth in bacterial cells. Some transition metal ions, such as Fe(Ⅱ), Mn(Ⅱ), Zn(Ⅱ), Ca(Ⅱ), could not only consider as the protein cofactors joining in the reaction of oxidizing reduction, but also set as the signal factors involving in the regulation of gene transcription and affecting the cytotoxicity and extreme resistance. Therefore, it is essential in regulating metal ions as to maintain normal physiology activity in bacterial cells. Moreover, the study of the regulation of metal ions in bacterial cells has always been a hotspot in the field of microbiology.
Deinococcus radiodurans, an important member of extreme micro-organisms family, has high resistances to ionizing radiation, ultraviolet (UV) radiation, chemical mutation and desiccation. However, more and more studies show that adaptability of D. radiodurans is benefit from its anti-oxidative ability. Furthermore, the high concentration of intracellular manganese ions is always considered to be one of the most important antioxidation mechanisms. Therefore, D. radiodurans is a great kind of model organisms to study in regulating manganese ions metabolism. By studying the regulatory metabolism of manganese ions, we could not only to delve more deeply into the extremely resistant mechanism of D. radiodurans, but it is also significant to widen the regulatory mechanisms of metal ions in bacterial cells. In this study, by researching the manganese acquisition and efflux mechanism, we investigated the regulation of intracellular manganese homeostasis of D. radiodurans. The results of the studies are listed as follows:
1. First of all, we identified and evaluated a manganese efflux protein (Drl236) in D. radiodurans. Dr1236 is the second manganese efflux protein that indentified from bacterial and is also first verified as the third type of efflux protein. By constructing the Dr1236 deletion mutant, we found the resistance of Dr1236 to manganese ions was decreased. whereas the level of intracellular manganese concentration is markedly increased. This indicated that Dr1236 play an critical role in the manganese resistance mechanism and in regulating intracellular manganese levels. Further, compared with the wild-type strain, the bacteria has extremely higher resistance to ionizing radiation, ultraviolet (UV) radiation and H2O2 after intracellular manganese content in mutant increased. The results indicated that the manganese ions are indispensable involved in the resistance of D. radiodurans cells, and then the protective effects may be realized by reducing the challenge of free radical to protein. Though the mutant cells accumulated higher intracellular concentration of manganese ions and extreme resistance was also increased differently, the cell lifespan of mutant cells is far shorter than the wild-type strain. The result showed that manganese ions at high concentrations are toxic to cells. Moreover, manganese efflux protein plays the key role in D. radiodurans.
2. Dr0865, which encodes the only Fur homologue, may be involved in the regulation of intracellular manganese levels in D. radiodurans. E. coli expression system was used to express the dr0865 and its biochemical characteristics were also investigated. Additionally, bioinformatics study showed that Dr0865 probably has three metal-binding sites and add up to a distinct advantage of biological function. DNA binding activity assay showed that Dr0865 could bind with the promoters of dr2283、dr2284、dr2523, the binding activities are positive proportional to the concentration of metal ions. On the other hand, Dr0865 could not bind with the promoter of dr 1709. (3-galactosidase activity assay showed that Dr0865 binds with promoters of dr2283、dr2284、dr2523 could promote gene transcription. Moreover, it should be considered that the DNA binding activity of Dr0865 is dependent to metal ions concentration and the mutant resistance of Dr0865 is enhanced to H2O2. We supposed that dr2283、dr2284、dr2523 probably encode a new kind of Fe2+efflux protein. Furthermore, the regulatory mechanism of Fe ions is still unclear and needs further study.
3. Dr2539 belongs to the MntR/DtxR family, and this family plays important role in intracellular Mn- or Fe- homeostasis regulation. A solubility-enhancing partner of translation initiation factor (IF) was fused to dr2539, and Dr2539 was successfully expressed in vitro. EMSA results indicated that Dr2539 can bind with the dr1709 promoter and the DNA binding activity of Dr2539 depended on the concentration of Mn and Fe. It is quite different with previous reported MntR because it is always Mn specifically binding protein. Bioinformatic analysis has shown that Dr2539 has DtxR characteristics, and further site-directed mutagenesis was carried out in order to explain why Dr2539 can response to both Mn and Fe. Different Dr2539 mutants were constructed and complemented to dr2539 null mutant. By examine the manganese resistance of complemented strains (C-D11M、C-H98Y、C-E101D、C-Tru), we found that the manganese resistance of C-D11M and C-Tru decreased compared with wild strain. Surprisingly, the resistance of C-H98Y decreased dramatically in 6-7mM Mn(II), while that recovered in>7mM. It is consistent with previous reported three step active model of DtxR, therefore, we suppose that the Dr2539 may be active in DtxR-like pattern, and it may also be involved in Mn/Fe homeostasis regulation.
In sum, we investigated the manganese efflux ans acquisition system, and the results indicate that the manganese involved in the extreme resistance of D. radiodurans. Moreover, the biochemical characteristics of metalloregulator Dr0865 and Dr2539 suggested that the intracellular metal concentration regulated system in D. radiodurans may be different with that of presented reported bacteria, and our studies gave a good first step for further Dr0865 and Dr2539 structure analysis.
作为极端微生物家族的重要成员,耐辐射球菌对于电离辐射、紫外线、化学诱变剂、干燥等具有极强抗性,而越来越多的研究表明,耐辐射球菌对于极端环境的适应能力得益于其超强的抗氧化能力,其细胞内积累的超高锰离子水平一直被认为是其最重要的抗氧化机制之一,因此耐辐射球菌是极好的研究细菌细胞锰离子代谢调控的模式生物,通过研究其锰离子调控机制,不但可以进一步了解耐辐射球菌的极端抗性机制,而且对于拓展细菌细胞金属离子调控机制也具有重要意义。本文主要通过细胞吸收及外排两个方面,研究了耐辐射球菌的细胞内锰离子动态调控机制,研究结果如下：
1.我们首次鉴定得到耐辐射球菌锰离子外排蛋白(Dr1236)。在构建了Dr1236的基因缺失突变株后,我们发现突变株对于锰离子的抗性下降,而且细胞内的锰离子水平显著升高,说明Dr1236在耐辐射球菌锰离子抗性及细胞内锰离子动态调控机制中,发挥了重要的作用。进一步的研究发现,突变株细胞内锰离子浓度升高以后,其电离辐射、紫外、过氧化氢抗性都明显增强,表明细胞内锰离子确实参与了细胞的极端抗性,而这种保护作用很可能是通过降低自由基对蛋白的攻击实现的。然而,虽然突变株细胞在积累了高水平锰离子之后,极端抗性有了不同程度的增加,但是细胞的生长周期却远远短于野生株,表明锰离子的过高积累对细胞的整体机能造成了损伤,这也从侧面反映出锰离子外排蛋白对于细胞的重要作用。
2.dr0865编码了耐辐射球菌唯一一个Fur同源蛋白,Fur蛋白在其它细菌中参与了锰离子吸收通道蛋白的转录调控,因此我们利用大肠杆菌表达系统体外表达了dr0865,并研究了其生化性质。生物信息学研究表明,Dr0865很可能包含了结合三个金属离子的结构,这种结构可能赋予了其独特的生物学功能。DNA结合活性实验表明,Dr0865可以与dr2283、dr2284、dr2524的启动子结合,而且结合活性与金属离子的浓度成正比,另一方面却不能与dr1709的启动子结合；β-半乳糖甘酶活性实验表明Dr0865与dr2283、dr2284、dr2523的启动子的结合可能促进了其转录,考虑到Dr0865的DNA结合活性依赖于金属离子的浓度,及Dr0865突变株的过氧化氢抗性增强,因此我们推测dr2283、dr2284、dr2523(?)(?)可能编码了一种铁离子外排通道蛋白,而其具体调控机制还需要进一步研究。
3.dr2539编码耐辐射球菌的DtxR/MntR家族蛋白,这类蛋白家族对于细胞锰离子或铁离子调控至关重要,通过在Dr2539 N-端融合表达助溶片段IF,我们成功得到Dr2539蛋白。DNA结合活性实验表明,Dr2539只能与dr1709的启动子结合,而且EMSA和p-半乳糖甘酶活性实验表明,蛋白的结合活性受到锰离子和铁离子的调控,表明Dr2539与此前所报道的MntR蛋白结构上可能存在很大差异,因此我们构建了Dr2539突变体,并将突变体补偿Dr2539突变株,包括C-D11M、C-H98Y、C-E101D、C-Tru,通过检测补偿株对于锰离子的抗性,发现C-D11M及C-Tru锰离子抗性相对于野生Dr2539补偿株明显下降,而C-E101D的抗性并没有明显变化；惊奇的是,H98Y补偿株对6-7Mm Mn(Ⅱ)极度敏感,这与此前所报道的DtxR激活模型一致,因此我们推测,dr2539所编码的MntR蛋白很可能利用一种类似于DtxR的激活模式,而且可能参与了细胞内锰离子与铁离子动态平衡的调控。
综上所述,在研究了耐辐射球菌细胞内锰离子外排和吸收系统后,我们的结果表明锰离子的确参与了耐辐射球菌的极端抗性机制,此外,金属离子调控蛋白Dr0865和Dr2539的生化活性表明,耐辐射球菌的金属离子调控机制与目前发现的细菌调控机制存在很大差异,而这种差异可能与其极端环境生存能力有关。我们的研究为下一步深入研究耐辐射球菌金属离子调控蛋白Dr0865和Dr2539结构奠定了基础。
Homeostasis of metal ions is closely related to the physiological processes of growth in bacterial cells. Some transition metal ions, such as Fe(Ⅱ), Mn(Ⅱ), Zn(Ⅱ), Ca(Ⅱ), could not only consider as the protein cofactors joining in the reaction of oxidizing reduction, but also set as the signal factors involving in the regulation of gene transcription and affecting the cytotoxicity and extreme resistance. Therefore, it is essential in regulating metal ions as to maintain normal physiology activity in bacterial cells. Moreover, the study of the regulation of metal ions in bacterial cells has always been a hotspot in the field of microbiology.
Deinococcus radiodurans, an important member of extreme micro-organisms family, has high resistances to ionizing radiation, ultraviolet (UV) radiation, chemical mutation and desiccation. However, more and more studies show that adaptability of D. radiodurans is benefit from its anti-oxidative ability. Furthermore, the high concentration of intracellular manganese ions is always considered to be one of the most important antioxidation mechanisms. Therefore, D. radiodurans is a great kind of model organisms to study in regulating manganese ions metabolism. By studying the regulatory metabolism of manganese ions, we could not only to delve more deeply into the extremely resistant mechanism of D. radiodurans, but it is also significant to widen the regulatory mechanisms of metal ions in bacterial cells. In this study, by researching the manganese acquisition and efflux mechanism, we investigated the regulation of intracellular manganese homeostasis of D. radiodurans. The results of the studies are listed as follows:
1. First of all, we identified and evaluated a manganese efflux protein (Drl236) in D. radiodurans. Dr1236 is the second manganese efflux protein that indentified from bacterial and is also first verified as the third type of efflux protein. By constructing the Dr1236 deletion mutant, we found the resistance of Dr1236 to manganese ions was decreased. whereas the level of intracellular manganese concentration is markedly increased. This indicated that Dr1236 play an critical role in the manganese resistance mechanism and in regulating intracellular manganese levels. Further, compared with the wild-type strain, the bacteria has extremely higher resistance to ionizing radiation, ultraviolet (UV) radiation and H2O2 after intracellular manganese content in mutant increased. The results indicated that the manganese ions are indispensable involved in the resistance of D. radiodurans cells, and then the protective effects may be realized by reducing the challenge of free radical to protein. Though the mutant cells accumulated higher intracellular concentration of manganese ions and extreme resistance was also increased differently, the cell lifespan of mutant cells is far shorter than the wild-type strain. The result showed that manganese ions at high concentrations are toxic to cells. Moreover, manganese efflux protein plays the key role in D. radiodurans.
2. Dr0865, which encodes the only Fur homologue, may be involved in the regulation of intracellular manganese levels in D. radiodurans. E. coli expression system was used to express the dr0865 and its biochemical characteristics were also investigated. Additionally, bioinformatics study showed that Dr0865 probably has three metal-binding sites and add up to a distinct advantage of biological function. DNA binding activity assay showed that Dr0865 could bind with the promoters of dr2283、dr2284、dr2523, the binding activities are positive proportional to the concentration of metal ions. On the other hand, Dr0865 could not bind with the promoter of dr 1709. (3-galactosidase activity assay showed that Dr0865 binds with promoters of dr2283、dr2284、dr2523 could promote gene transcription. Moreover, it should be considered that the DNA binding activity of Dr0865 is dependent to metal ions concentration and the mutant resistance of Dr0865 is enhanced to H2O2. We supposed that dr2283、dr2284、dr2523 probably encode a new kind of Fe2+efflux protein. Furthermore, the regulatory mechanism of Fe ions is still unclear and needs further study.
3. Dr2539 belongs to the MntR/DtxR family, and this family plays important role in intracellular Mn- or Fe- homeostasis regulation. A solubility-enhancing partner of translation initiation factor (IF) was fused to dr2539, and Dr2539 was successfully expressed in vitro. EMSA results indicated that Dr2539 can bind with the dr1709 promoter and the DNA binding activity of Dr2539 depended on the concentration of Mn and Fe. It is quite different with previous reported MntR because it is always Mn specifically binding protein. Bioinformatic analysis has shown that Dr2539 has DtxR characteristics, and further site-directed mutagenesis was carried out in order to explain why Dr2539 can response to both Mn and Fe. Different Dr2539 mutants were constructed and complemented to dr2539 null mutant. By examine the manganese resistance of complemented strains (C-D11M、C-H98Y、C-E101D、C-Tru), we found that the manganese resistance of C-D11M and C-Tru decreased compared with wild strain. Surprisingly, the resistance of C-H98Y decreased dramatically in 6-7mM Mn(II), while that recovered in>7mM. It is consistent with previous reported three step active model of DtxR, therefore, we suppose that the Dr2539 may be active in DtxR-like pattern, and it may also be involved in Mn/Fe homeostasis regulation.
In sum, we investigated the manganese efflux ans acquisition system, and the results indicate that the manganese involved in the extreme resistance of D. radiodurans. Moreover, the biochemical characteristics of metalloregulator Dr0865 and Dr2539 suggested that the intracellular metal concentration regulated system in D. radiodurans may be different with that of presented reported bacteria, and our studies gave a good first step for further Dr0865 and Dr2539 structure analysis.
引文
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