耐辐射奇球菌类胡萝卜素合成途径相关基因的研究以及两个新种的多相分类鉴定
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摘要
耐辐射奇球菌(Deinococcus radiodurans R1)是一株红色的细菌,对电离辐射、紫外线、干燥以及许多化学诱变剂和强氧化剂有很强的抗性。大量的研究表明,耐辐射奇球菌的超强抗性是特殊的细胞物质结构、活性氧(reactive oxygen species, ROS)的有效清除以及DNA损伤的高效修复等方面共同作用的结果。自然界中许多类胡萝卜素是天然的抗氧化剂。耐辐射奇球菌能产生大量红色的类胡萝卜素物质deinoxanthin,这种类胡萝卜素在抗氧化方面的生物学功能尚未完全阐明。
本研究通过敲除crtB和crtl这两个类胡萝卜素合成途径的上游基因成功构建两个无色突变株,并将其与野生型进行比较。比较的结果显示无色突变株对电离辐射、紫外线以及H2O2的抗性较野生型有所降低,但对丝裂霉素C(MMC)的抗性无明显变化。利用电子自旋共振(electron spin resonance, ESR)和自旋捕捉技术,在UV-C照射耐辐射奇球菌悬浮液体系时观察到自由基信号,信号的强弱受体系中耐辐射奇球菌细胞内类胡萝卜素水平的影响。在耐辐射奇球菌crtB无色突变株的悬浮液体系中加入野生型的色素抽提物,能明显抑制自由基信号的出现。因此,细胞体内与体外两方面的实验证据均说明野生型合成的类胡萝卜素可以通过清除自由基而保护细胞。
进而对耐辐射奇球菌无色突变株的菌悬液体系中产生的自由基信号的类型进行分析。在crtB突变株的菌悬液体系中加入羟自由基特异性清除剂D-甘露醇后自由基信号几乎完全消失,说明UV-C照射crtB突变株的过程中生成的自由基主要是羟自由基。不过,在该体系加入过氧化氢酶也能使大部分的自由基信号消失,说明过氧化氢是羟自由基产生的关键条件。从野生株中提取的类胡萝卜素能使crtB突变株体系的自由基信号消失,且在野生型的体系中这种自由基信号不会出现,说明野生株合成的类胡萝卜素能在细胞体内和体外清除羟自由基。此外,从耐辐射奇球菌野生型中抽提的类胡萝卜素还能明显地清除核黄素/EDTA体系产生的超氧阴离子自由基。综合这些结果,得出以下结论：耐辐射奇球菌野生型中的类胡萝卜素可以通过清除羟自由基和超氧阴离子参与细胞的活性氧防御机制。
耐辐射奇球菌的主要类胡萝卜素组分deinoxanthin具有独特的结构,说明耐辐射奇球菌中催化其合成的酶和途径有独特之处,值得深入研究。本研究对耐辐射奇球菌类胡萝卜素合成途径的另外两个基因crtLm (DR0801,编码番茄红素环化酶)和crtO(DR0093,编码p-胡萝卜素酮化酶)进行敲除,得到两个不同颜色的突变株。然后对这两个突变株的类胡萝卜素组分作了初步检测,为进一步研究耐辐射奇球菌的类胡萝卜素合成途径奠定了基础。此外,本研究通过分子生物学手段成功地构建了crtB、crtl及crtO基因的重组表达质粒。其中crtO基因在大肠杆菌中得到高水平的表达,而crtB和crtI基因的表达在SDS-PAGE电泳图谱中无法检测到。
另一方面,本研究从新疆维吾尔自治区吐鲁番地区的戈壁荒漠区域采集的土壤样品中分离到184个菌株。形态特征、16S rRNA基因序列、部分生理生化特性以及全细胞脂肪酸组分的初步鉴定结果表明这184株细菌分属于5个细菌门的31个属中,显示出较丰富的物种多样性。
在分离菌株中有多个潜在的新分类单元存在。本研究通过形态观察,系统发育分析,生理生化特性检测,醌组分、全细胞脂肪酸组分和极性脂组分的分析,DNA G+C含量测定和DNA-DNA分子杂交等多相分类鉴定技术对分离菌株X14-1T、X19-1和ZLB-3T进行了详细的鉴定,并在国际权威的系统分类学期刊IJSEM上发表。其中菌株X14-1T与X19-1被确定为Pontibacter属的一个新种,命名为Pontibacter korlensis sp. nov., X14-1T为这个种的模式菌株；菌株ZLB-3T被确定为Hymenobacter属的一个新种,命名为Hymenobacter deserti sp. nov.。
Deinococcus radiodurans R1, a red-pigmented strain of the extremely radioresistant genus Deinococcus, showed high resistance to ionizing radiation, ultraviolet light, desiccation and many chemical agents. The high resistance of Deinococcus radiodurans R1 was attributed to the special cell structure, the scavenging ability of reactive oxygen species and efficient repair capability for DNA damage. Many carotenoids are the natural antioxidants. Deinococcus radiodurans R1 contains a major carotenoid namely deinoxanthin. Its possible antioxidant role in this strain has not been completely elucidated.
In this study, two colorless mutants were constructed by knockout of crtB and crtl genes, respectively. Comparative analysis of the two colorless mutants and the wild type showed that the two colorless mutants were more sensitive to ionizing radiation, UV, and hydrogen peroxide, but not to mitomycin-C (MMC). With electron spin resonance (ESR) and spin trapping techniques, we observed that free radical signals occurred in the suspensions of UV irradiated Deinococcus radiodurans cells and the intensity of signals was influenced by carotenoids levels within the cells. We further showed that the carotenoid extract from the wild type could obviously scavenge the free radical signals generated by the irradiated crtB mutant cells. These results suggest that carotenoids in D. radiodurans R1 function as free radical scavengers.
To define which type of free radicals was generated in the process of UV radiation of crtB mutant cells, D-mannitol, a commonly used hydroxyl radical scavenger, was added to the mixture before radiation. The presence of D-mannitol resulted in the evident decrease of the free radical signals, suggesting that the free radical signals generated in UV-irradiated crtB mutant cells were mainly hydroxyl radicals. Furthermore, when catalase was added to the mixture, the major signals of free radicals also disappeared. This result indicated that hydrogen peroxide production was a prerequisite to the generation of hydroxyl radicals in the process of UV radiation of crtB mutant cells. The results also demonstrated that carotenoid extract from the wild type could scavenge the free radical signals generated in UV-irradiated crtB mutant cells, and no obvious free radicals were generated in the UV-irradiated wild type. Thus, carotenoids of D. radiodurans R1 act as hydroxyl radical scavengers in vivo. The ESR assay in vitro indicated that carotenoids from D. radiodurans R1 were also capable of scavenging superoxide anion. Therefore, carotenoids of D. radiodurans R1 play an important role in the defense mechanism of this organism by scavenge free radicals including hydroxyl radicals and superoxide anion.
Deinoxanthin, the major pigment in D. radiodurans R1, has particular structure which has not been found in other organisms. Thus, the genes in the carotenoid biosynthetic pathway should be much different and further study is required. The genes crtLm (DR0801, encoding lycopeneβ-cyclase) and crtO (DR0093, encodingβ-carotene ketolase) were knocked out and two colored mutants were obtained. The carotenoid extracts of these mutants were also tested by HPLC. The expression vectors of crtB, crtl and crtO genes were also constructed. However, onlyβ-carotene ketolase could be expressed in E. coli. The expression of crtB and crtl genes could not be detected in the SDS-PAGE gel electrophoresis.
On the other hand,184 bacterial isolates were obtained from soil samples from Xinjiang Province, China. On the basis of morphological characteristics,16S rRNA gene sequences, partial physiological and biochemical features, and whole-cell fatty acid profiles, these isolates belonged to 31 genera within 5 phyla.
Several strains of these isolates showed big differences with any recognized bacterial species and probably belonged to new species or new genera. And then three strains were identified by the approaches of polyphasic taxonomy. On the basis of phenotypic characteristics, chemotaxonomic data, phylogenetic analysis, DNA G+C content and DNA-DNA hybridization, strains X14-1T and X19-1 are considered to represent a novel species of the genus Pontibacter, for which the name Pontibacter korlensis sp. nov. is proposed. The type strain is X14-1T. Strain ZLB-3T is considered to represent a novel species of the genus Hymenobacter, for which the name Hymenobacter deserti sp. nov. sp. nov. is proposed.
本研究通过敲除crtB和crtl这两个类胡萝卜素合成途径的上游基因成功构建两个无色突变株,并将其与野生型进行比较。比较的结果显示无色突变株对电离辐射、紫外线以及H2O2的抗性较野生型有所降低,但对丝裂霉素C(MMC)的抗性无明显变化。利用电子自旋共振(electron spin resonance, ESR)和自旋捕捉技术,在UV-C照射耐辐射奇球菌悬浮液体系时观察到自由基信号,信号的强弱受体系中耐辐射奇球菌细胞内类胡萝卜素水平的影响。在耐辐射奇球菌crtB无色突变株的悬浮液体系中加入野生型的色素抽提物,能明显抑制自由基信号的出现。因此,细胞体内与体外两方面的实验证据均说明野生型合成的类胡萝卜素可以通过清除自由基而保护细胞。
进而对耐辐射奇球菌无色突变株的菌悬液体系中产生的自由基信号的类型进行分析。在crtB突变株的菌悬液体系中加入羟自由基特异性清除剂D-甘露醇后自由基信号几乎完全消失,说明UV-C照射crtB突变株的过程中生成的自由基主要是羟自由基。不过,在该体系加入过氧化氢酶也能使大部分的自由基信号消失,说明过氧化氢是羟自由基产生的关键条件。从野生株中提取的类胡萝卜素能使crtB突变株体系的自由基信号消失,且在野生型的体系中这种自由基信号不会出现,说明野生株合成的类胡萝卜素能在细胞体内和体外清除羟自由基。此外,从耐辐射奇球菌野生型中抽提的类胡萝卜素还能明显地清除核黄素/EDTA体系产生的超氧阴离子自由基。综合这些结果,得出以下结论：耐辐射奇球菌野生型中的类胡萝卜素可以通过清除羟自由基和超氧阴离子参与细胞的活性氧防御机制。
耐辐射奇球菌的主要类胡萝卜素组分deinoxanthin具有独特的结构,说明耐辐射奇球菌中催化其合成的酶和途径有独特之处,值得深入研究。本研究对耐辐射奇球菌类胡萝卜素合成途径的另外两个基因crtLm (DR0801,编码番茄红素环化酶)和crtO(DR0093,编码p-胡萝卜素酮化酶)进行敲除,得到两个不同颜色的突变株。然后对这两个突变株的类胡萝卜素组分作了初步检测,为进一步研究耐辐射奇球菌的类胡萝卜素合成途径奠定了基础。此外,本研究通过分子生物学手段成功地构建了crtB、crtl及crtO基因的重组表达质粒。其中crtO基因在大肠杆菌中得到高水平的表达,而crtB和crtI基因的表达在SDS-PAGE电泳图谱中无法检测到。
另一方面,本研究从新疆维吾尔自治区吐鲁番地区的戈壁荒漠区域采集的土壤样品中分离到184个菌株。形态特征、16S rRNA基因序列、部分生理生化特性以及全细胞脂肪酸组分的初步鉴定结果表明这184株细菌分属于5个细菌门的31个属中,显示出较丰富的物种多样性。
在分离菌株中有多个潜在的新分类单元存在。本研究通过形态观察,系统发育分析,生理生化特性检测,醌组分、全细胞脂肪酸组分和极性脂组分的分析,DNA G+C含量测定和DNA-DNA分子杂交等多相分类鉴定技术对分离菌株X14-1T、X19-1和ZLB-3T进行了详细的鉴定,并在国际权威的系统分类学期刊IJSEM上发表。其中菌株X14-1T与X19-1被确定为Pontibacter属的一个新种,命名为Pontibacter korlensis sp. nov., X14-1T为这个种的模式菌株；菌株ZLB-3T被确定为Hymenobacter属的一个新种,命名为Hymenobacter deserti sp. nov.。
Deinococcus radiodurans R1, a red-pigmented strain of the extremely radioresistant genus Deinococcus, showed high resistance to ionizing radiation, ultraviolet light, desiccation and many chemical agents. The high resistance of Deinococcus radiodurans R1 was attributed to the special cell structure, the scavenging ability of reactive oxygen species and efficient repair capability for DNA damage. Many carotenoids are the natural antioxidants. Deinococcus radiodurans R1 contains a major carotenoid namely deinoxanthin. Its possible antioxidant role in this strain has not been completely elucidated.
In this study, two colorless mutants were constructed by knockout of crtB and crtl genes, respectively. Comparative analysis of the two colorless mutants and the wild type showed that the two colorless mutants were more sensitive to ionizing radiation, UV, and hydrogen peroxide, but not to mitomycin-C (MMC). With electron spin resonance (ESR) and spin trapping techniques, we observed that free radical signals occurred in the suspensions of UV irradiated Deinococcus radiodurans cells and the intensity of signals was influenced by carotenoids levels within the cells. We further showed that the carotenoid extract from the wild type could obviously scavenge the free radical signals generated by the irradiated crtB mutant cells. These results suggest that carotenoids in D. radiodurans R1 function as free radical scavengers.
To define which type of free radicals was generated in the process of UV radiation of crtB mutant cells, D-mannitol, a commonly used hydroxyl radical scavenger, was added to the mixture before radiation. The presence of D-mannitol resulted in the evident decrease of the free radical signals, suggesting that the free radical signals generated in UV-irradiated crtB mutant cells were mainly hydroxyl radicals. Furthermore, when catalase was added to the mixture, the major signals of free radicals also disappeared. This result indicated that hydrogen peroxide production was a prerequisite to the generation of hydroxyl radicals in the process of UV radiation of crtB mutant cells. The results also demonstrated that carotenoid extract from the wild type could scavenge the free radical signals generated in UV-irradiated crtB mutant cells, and no obvious free radicals were generated in the UV-irradiated wild type. Thus, carotenoids of D. radiodurans R1 act as hydroxyl radical scavengers in vivo. The ESR assay in vitro indicated that carotenoids from D. radiodurans R1 were also capable of scavenging superoxide anion. Therefore, carotenoids of D. radiodurans R1 play an important role in the defense mechanism of this organism by scavenge free radicals including hydroxyl radicals and superoxide anion.
Deinoxanthin, the major pigment in D. radiodurans R1, has particular structure which has not been found in other organisms. Thus, the genes in the carotenoid biosynthetic pathway should be much different and further study is required. The genes crtLm (DR0801, encoding lycopeneβ-cyclase) and crtO (DR0093, encodingβ-carotene ketolase) were knocked out and two colored mutants were obtained. The carotenoid extracts of these mutants were also tested by HPLC. The expression vectors of crtB, crtl and crtO genes were also constructed. However, onlyβ-carotene ketolase could be expressed in E. coli. The expression of crtB and crtl genes could not be detected in the SDS-PAGE gel electrophoresis.
On the other hand,184 bacterial isolates were obtained from soil samples from Xinjiang Province, China. On the basis of morphological characteristics,16S rRNA gene sequences, partial physiological and biochemical features, and whole-cell fatty acid profiles, these isolates belonged to 31 genera within 5 phyla.
Several strains of these isolates showed big differences with any recognized bacterial species and probably belonged to new species or new genera. And then three strains were identified by the approaches of polyphasic taxonomy. On the basis of phenotypic characteristics, chemotaxonomic data, phylogenetic analysis, DNA G+C content and DNA-DNA hybridization, strains X14-1T and X19-1 are considered to represent a novel species of the genus Pontibacter, for which the name Pontibacter korlensis sp. nov. is proposed. The type strain is X14-1T. Strain ZLB-3T is considered to represent a novel species of the genus Hymenobacter, for which the name Hymenobacter deserti sp. nov. sp. nov. is proposed.
引文
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