嗜热毛壳菌超氧化物歧化酶基因的克隆、表达及转基因烟草耐盐性研究
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摘要
超氧化物歧化酶(Superoxide Dismutase,SOD)能够催化超氧阴离子(O2.~-)发生歧化反应,在生物体防御氧化损伤的过程中,发挥着重要作用。根据酶分子中所含金属辅基的不同,超氧化物歧化酶主要可分为Cu,Zn-SOD,Mn-SOD,Fe-SOD等类型。
超氧化物歧化酶已在食品、医药、化妆品和农业等领域中得到了广泛的应用,具有重要商业价值。但市场应用的超氧化物歧化酶主要来源于常温生物,因此酶的半衰期短,热稳定性差,储存期短,导致产品成本高,而制约超氧化物歧化酶在农业、工业中的应用。由此可见,热稳定性超氧化物歧化酶的研究和开发具有重要意义。
嗜热毛壳菌(Chaetomium thermophilum CT2)是广泛分布的,生长上限温度较高的嗜热真菌,从这种真菌中已分离了多种嗜热酶,具有极大的研究和应用价值。本研究通过RT-PCR和Tail-PCR从嗜热毛壳菌中分离到了一个新的Mn-SOD酶基因,序列分析表明该基因全长cDNA序列为684bp,编码227个氨基酸,具有信号肽。将该基因在毕赤酵母中表达,经甲醇诱导表达6d后酶活最高可达1020U/ml,蛋白量达0.93mg/ml。经硫酸铵沉淀、DEAE-Sepharose Fast Flow阴离子交换等步骤纯化了该重组蛋白,SDS-PAGE显示该重组蛋白大小约为22.7kDa,与推测的蛋白分子量相似。重组酶的最适反应温度和最适pH值分别为60℃和6.0,该酶具有较高的热稳定性,70℃处理60min后,残余酶活为71%。
本研究还从嗜热毛壳菌中克隆了一种Cu,Zn-SOD基因并将其在毕赤酵母中进行了高效表达。经甲醇诱导表达6d后酶活最高可达1660U/ml,蛋白量达1.54mg/ml。经硫酸铵沉淀、DEAE-Sepharose Fast Flow阴离子交换等步骤纯化了该重组蛋白,SDS-PAGE显示该重组蛋白大小约为17 kDa,该重组酶的最适反应温度和最适pH值分别为60℃和6.5。70℃处理60min后,残余酶活为65%,在80和90℃下的半衰期分别为22和7min。
比较上述构建的重组酵母GS115(pPIC9K/mnsod)和GS115(pPIC9K/czsod)与非重组酵母(pPIC9K)在不同浓度NaCl、百草枯、甲萘醌和H_2O_2胁迫下的生长状况,验证嗜热毛壳菌mnsod及czsod基因对不同非生物胁迫的抵抗能力。结果表明嗜热毛壳菌mnsod及czsod基因对包括盐、百草枯、甲萘醌、过氧化物等多种胁迫具有抵抗能力。
盐胁迫是自然界中主要的非生物胁迫之一,是影响植物生长发育的主要因素,盐胁迫的适应机制在许多模式生物中广泛研究。利用基因工程技术提高植物耐盐性是解决这一问题的最有效途径之一。研究抗逆相关基因的抗逆功能是基因工程抗性育种的前提。当植物遭受包括高盐、干旱、低温和高温等环境胁迫时可产生大量活性氧。活性氧进而破坏细胞内的大分子,从而使DNA裂解,造成膜的损伤或影响蛋白质的合成及稳定性等。因此,对植物来说有效的活性氧清除系统非常重要。
植物的超氧化物歧化酶活性与其在盐胁迫下的防御能力密切相关。关于逆境胁迫下SOD活性增高的生理关系已有报道,表明植物体内的SOD活性增高可以增强其对逆境胁迫的抵抗能力。到目前为止已报道多种转SOD基因植物对盐胁迫的抗性。
为了验证盐胁迫下嗜热毛壳菌mnsod及czsod的基因功能,将mnsod及czsod基因分别构建到35s启动子驱动的植物表达载体pROKⅡ中,通过农杆菌介导法转化烟草。经PCR、southern杂交和RT-PCR检测表明嗜热毛壳菌的mnsod及czsod基因分别整合到烟草基因组中。
通过分析高盐胁迫下非转基因和转基因烟草的种子萌发率表明,转mnsod基因和转czsod基因的烟草种子比野生型烟草种子的萌发率高,且两种转基因烟草的根生长受到的抑制都比野生型小。说明转嗜热毛壳菌sod基因提高了烟草在种子萌发及小苗生长阶段的耐盐性。
盐分胁迫会使植物膜系统受到伤害,导致植株的叶片MDA含量升高,膜脂过氧化作用增强。本实验中盐胁迫处理使转基因植株与野生型植物叶片的丙二醛含量都有所上升,但两种转基因烟草植株的MDA含量显著低于未转基因植株。说明转基因烟草膜脂过氧化程度低于野生型烟草,膜伤害程度较轻。该结果表明在盐胁迫下转基因植株与野生型植株相比具有明显提高的抗盐性。
在盐胁迫条件下,转mnsod基因和转czsod基因烟草株系的SOD活性与野生型一样都呈先升高后降低的趋势,但两种转基因烟草SOD活性都比非转基因烟草高的多。研究了两种转基因烟草植株的抗病害能力,结果表明两种转基因植株对烟草炭疽病和赤星病有一定的抵抗能力,且转mnsod基因比转czsod基因烟草的抗病能力更强。
The superoxide dismutases (SOD, EC1.15.1.1) are metallo-enzymes that catalyze the dismutation of superoxide(O2.~-) to hydrogen peroxide(H_2O_2) and molecular oxygen(O2). They have play a critical role in the defense against oxidative stress. There are three general classes of SODs in organisms, which differ in their metal cofactors: copper zinc-containing SOD (Cu,Zn-SOD), manganese-containing SOD (Mn-SOD)and iron-containing SOD (Fe-SOD).
In recent years, the important role of SOD in food, pharmaceutical, cosmeceutical industries and agriculture has been demonstrated. The SOD used in the market mainly from room temperature organisms. However, SOD is easily degraded at high temperatures and it can not be stored for long periods. Due to high production costs, poor thermal stability, low activity and yield, the application of SOD is restricted in agriculture and industrial applications. Therefore, there has been increasing interest in research and development of thermostable SODs from thermophilic organisms.
Chaetomium.thermophilum is a widely distributed soil-inhibiting fungi of considerable interest producers of thermostable enzymes, including glucoamylase, cellulase, endocellulase, xylanase, and laccase. The fungi has great value in research and application.
In this study, a Mn-SOD gene was isolated firstly from C.thermophilum CT2 by RT-PCR and Tail-PCR. Sequence analysis showed that the full cDNA of the gene was 684bp, encoding 227 amino acids. The gene was highly expressed in Paster.pastoris and the recombinant protein was purified by ammonium sulfate precipitation, DEAE-Sepharose Fast Flow anion exchange. SDS-PAGE showed that the recombinant protein was about 23kDa, similar to the deduced molecular weight. The optimal temperature and pH were 60℃and 6.0, separately. The enzyme was relatively thermostable, after incubated at 70℃for 60 min still remained 71% activity.
In this study, a Cu,Zn-SOD gene from C.thermophilum was also cloned and highly expressed in Paster.pastoris, Six days after induction, this strain had the highest SOD activity of 1,660 U/ml culture, and its expression level was 1.54 mg/ml. The recombinant protein was purified by ammonium sulfate precipitation and DEAE-Sepharose Fast Flow anion exchange. SDS-PAGE showed that the recombinant protein was about 17kDa. The recombinant Cu,Zn-SOD exhibited optimum activity at pH 6.5 and 60℃. The enzyme retained 65% of the maximum activity at 70℃for 60min and the half-life was 22min and 7min at 80 and 90℃, respectively.
The recombinant yeast GS115 (harboring pPIC9K/mnsod or pPIC9K/czsod) and control GS115 (harboring empty pPIC9K) were treated with different concentration of salt and oxidation stresses such as paraquat, mandione and H_2O_2. The results revealed that the recombinant yeast cells have a higher stress resistance than control cells. It also proved that the mnsod and czsod genes from C.thermophilum have the ability of salt and oxidation resistance.
Salt stress is the main factor that seriously affect on plant growth and development. Therefore, there is interest in breeding transgenic plants that are tolerant of saline soil. Salt-stress adaptation mechanisms are being intensively studied in model organisms. The genetic engineering is an effective way to improve plant salt stress resistance. One of the basis for plant genetic engineering is study on the function of stress- related gene. Exposure of plants to environmental stresses, including salinity, drought, cold and high temperatures, can lead to the generation of reactive oxygen species (ROS). ROS can attack cellular macromolecules, generate lesions in DNA, cause membrane damage, and affect protein synthesis and stability, therefore it is important for plants to have effective ROS scavenging mechanisms.
Superoxide dismutase activity is closely associated with defense ability of plants under the condition of salt stress. Physiological correlations between elevated SOD activity and stress tolerance have been reported, suggesting that the upregulation of SOD levels may enhance the stress-defense potential of plants. Various transgenic plants that express increased amounts of SODs have been generated for enhanced tolerance to environmental stress.
To investigate the function of mnsod and czsod from C.thermophilum under salt stress in plant,the full-length mnsod and czsod cDNAs were subcloned into the expression vector pROKⅡdownstream of the 35S-CaMV promoter, respectively. The constructs were first introduced into tobacco via Agrobacterium tumefaciens LBA4404 by the freezing transformation method and verified by PCR, Southern blot and RT-PCR. It was indicated that the mnsod and czsod gene had been recombined into tobacco genome, respectively, and transgenic tobacco plants were obtained.
On the medium containing the different concentration of NaCl, expression of mnsod or czsod gene in tobacco can showed the higher percentage of seed germination. Both of the transgenic lines exhibited increased root growth compare to WT plants.These results indicated that transgenic tobacco plants could enhance salt tolerance than the wild plants at the seed germination and seedlings growth stage.
Salt stress resulted in membrane injury and caused MDA content and the extent of the membrance lipid perioxidation rising in plants. In this study, salt stress increased MDA content in all Plants. However,the values of transgenic plants measured were much lower than those of wild-type, which indicated that the extent of the membrance lipid perioxidation in transgenic tobacco was lighter than that in wild type tobacco. These results indicated that the transgenic plants showed enhanced salt tolerance compared with wild-type under salt stress conditions.
Under salt stress, the activity of SOD increased first and then decreased in both wild types and transgenic tobacco plants. Furthermore, the transgenic tobacco plants always sustained higher SOD activity than wild-type plants.
The study showed that the SOD-transformed plants exhibited increased resistance to leaf infection with the fungi Colletotrichum nicotianae and Alternaria alternate.
The enhanced tolerance of transgenic lines to salt stress suggests that the mnsod and czsod genes from C.thermophilum are benefit for genetic engineering to improve plant tolerance to salt stress.
超氧化物歧化酶已在食品、医药、化妆品和农业等领域中得到了广泛的应用,具有重要商业价值。但市场应用的超氧化物歧化酶主要来源于常温生物,因此酶的半衰期短,热稳定性差,储存期短,导致产品成本高,而制约超氧化物歧化酶在农业、工业中的应用。由此可见,热稳定性超氧化物歧化酶的研究和开发具有重要意义。
嗜热毛壳菌(Chaetomium thermophilum CT2)是广泛分布的,生长上限温度较高的嗜热真菌,从这种真菌中已分离了多种嗜热酶,具有极大的研究和应用价值。本研究通过RT-PCR和Tail-PCR从嗜热毛壳菌中分离到了一个新的Mn-SOD酶基因,序列分析表明该基因全长cDNA序列为684bp,编码227个氨基酸,具有信号肽。将该基因在毕赤酵母中表达,经甲醇诱导表达6d后酶活最高可达1020U/ml,蛋白量达0.93mg/ml。经硫酸铵沉淀、DEAE-Sepharose Fast Flow阴离子交换等步骤纯化了该重组蛋白,SDS-PAGE显示该重组蛋白大小约为22.7kDa,与推测的蛋白分子量相似。重组酶的最适反应温度和最适pH值分别为60℃和6.0,该酶具有较高的热稳定性,70℃处理60min后,残余酶活为71%。
本研究还从嗜热毛壳菌中克隆了一种Cu,Zn-SOD基因并将其在毕赤酵母中进行了高效表达。经甲醇诱导表达6d后酶活最高可达1660U/ml,蛋白量达1.54mg/ml。经硫酸铵沉淀、DEAE-Sepharose Fast Flow阴离子交换等步骤纯化了该重组蛋白,SDS-PAGE显示该重组蛋白大小约为17 kDa,该重组酶的最适反应温度和最适pH值分别为60℃和6.5。70℃处理60min后,残余酶活为65%,在80和90℃下的半衰期分别为22和7min。
比较上述构建的重组酵母GS115(pPIC9K/mnsod)和GS115(pPIC9K/czsod)与非重组酵母(pPIC9K)在不同浓度NaCl、百草枯、甲萘醌和H_2O_2胁迫下的生长状况,验证嗜热毛壳菌mnsod及czsod基因对不同非生物胁迫的抵抗能力。结果表明嗜热毛壳菌mnsod及czsod基因对包括盐、百草枯、甲萘醌、过氧化物等多种胁迫具有抵抗能力。
盐胁迫是自然界中主要的非生物胁迫之一,是影响植物生长发育的主要因素,盐胁迫的适应机制在许多模式生物中广泛研究。利用基因工程技术提高植物耐盐性是解决这一问题的最有效途径之一。研究抗逆相关基因的抗逆功能是基因工程抗性育种的前提。当植物遭受包括高盐、干旱、低温和高温等环境胁迫时可产生大量活性氧。活性氧进而破坏细胞内的大分子,从而使DNA裂解,造成膜的损伤或影响蛋白质的合成及稳定性等。因此,对植物来说有效的活性氧清除系统非常重要。
植物的超氧化物歧化酶活性与其在盐胁迫下的防御能力密切相关。关于逆境胁迫下SOD活性增高的生理关系已有报道,表明植物体内的SOD活性增高可以增强其对逆境胁迫的抵抗能力。到目前为止已报道多种转SOD基因植物对盐胁迫的抗性。
为了验证盐胁迫下嗜热毛壳菌mnsod及czsod的基因功能,将mnsod及czsod基因分别构建到35s启动子驱动的植物表达载体pROKⅡ中,通过农杆菌介导法转化烟草。经PCR、southern杂交和RT-PCR检测表明嗜热毛壳菌的mnsod及czsod基因分别整合到烟草基因组中。
通过分析高盐胁迫下非转基因和转基因烟草的种子萌发率表明,转mnsod基因和转czsod基因的烟草种子比野生型烟草种子的萌发率高,且两种转基因烟草的根生长受到的抑制都比野生型小。说明转嗜热毛壳菌sod基因提高了烟草在种子萌发及小苗生长阶段的耐盐性。
盐分胁迫会使植物膜系统受到伤害,导致植株的叶片MDA含量升高,膜脂过氧化作用增强。本实验中盐胁迫处理使转基因植株与野生型植物叶片的丙二醛含量都有所上升,但两种转基因烟草植株的MDA含量显著低于未转基因植株。说明转基因烟草膜脂过氧化程度低于野生型烟草,膜伤害程度较轻。该结果表明在盐胁迫下转基因植株与野生型植株相比具有明显提高的抗盐性。
在盐胁迫条件下,转mnsod基因和转czsod基因烟草株系的SOD活性与野生型一样都呈先升高后降低的趋势,但两种转基因烟草SOD活性都比非转基因烟草高的多。研究了两种转基因烟草植株的抗病害能力,结果表明两种转基因植株对烟草炭疽病和赤星病有一定的抵抗能力,且转mnsod基因比转czsod基因烟草的抗病能力更强。
The superoxide dismutases (SOD, EC1.15.1.1) are metallo-enzymes that catalyze the dismutation of superoxide(O2.~-) to hydrogen peroxide(H_2O_2) and molecular oxygen(O2). They have play a critical role in the defense against oxidative stress. There are three general classes of SODs in organisms, which differ in their metal cofactors: copper zinc-containing SOD (Cu,Zn-SOD), manganese-containing SOD (Mn-SOD)and iron-containing SOD (Fe-SOD).
In recent years, the important role of SOD in food, pharmaceutical, cosmeceutical industries and agriculture has been demonstrated. The SOD used in the market mainly from room temperature organisms. However, SOD is easily degraded at high temperatures and it can not be stored for long periods. Due to high production costs, poor thermal stability, low activity and yield, the application of SOD is restricted in agriculture and industrial applications. Therefore, there has been increasing interest in research and development of thermostable SODs from thermophilic organisms.
Chaetomium.thermophilum is a widely distributed soil-inhibiting fungi of considerable interest producers of thermostable enzymes, including glucoamylase, cellulase, endocellulase, xylanase, and laccase. The fungi has great value in research and application.
In this study, a Mn-SOD gene was isolated firstly from C.thermophilum CT2 by RT-PCR and Tail-PCR. Sequence analysis showed that the full cDNA of the gene was 684bp, encoding 227 amino acids. The gene was highly expressed in Paster.pastoris and the recombinant protein was purified by ammonium sulfate precipitation, DEAE-Sepharose Fast Flow anion exchange. SDS-PAGE showed that the recombinant protein was about 23kDa, similar to the deduced molecular weight. The optimal temperature and pH were 60℃and 6.0, separately. The enzyme was relatively thermostable, after incubated at 70℃for 60 min still remained 71% activity.
In this study, a Cu,Zn-SOD gene from C.thermophilum was also cloned and highly expressed in Paster.pastoris, Six days after induction, this strain had the highest SOD activity of 1,660 U/ml culture, and its expression level was 1.54 mg/ml. The recombinant protein was purified by ammonium sulfate precipitation and DEAE-Sepharose Fast Flow anion exchange. SDS-PAGE showed that the recombinant protein was about 17kDa. The recombinant Cu,Zn-SOD exhibited optimum activity at pH 6.5 and 60℃. The enzyme retained 65% of the maximum activity at 70℃for 60min and the half-life was 22min and 7min at 80 and 90℃, respectively.
The recombinant yeast GS115 (harboring pPIC9K/mnsod or pPIC9K/czsod) and control GS115 (harboring empty pPIC9K) were treated with different concentration of salt and oxidation stresses such as paraquat, mandione and H_2O_2. The results revealed that the recombinant yeast cells have a higher stress resistance than control cells. It also proved that the mnsod and czsod genes from C.thermophilum have the ability of salt and oxidation resistance.
Salt stress is the main factor that seriously affect on plant growth and development. Therefore, there is interest in breeding transgenic plants that are tolerant of saline soil. Salt-stress adaptation mechanisms are being intensively studied in model organisms. The genetic engineering is an effective way to improve plant salt stress resistance. One of the basis for plant genetic engineering is study on the function of stress- related gene. Exposure of plants to environmental stresses, including salinity, drought, cold and high temperatures, can lead to the generation of reactive oxygen species (ROS). ROS can attack cellular macromolecules, generate lesions in DNA, cause membrane damage, and affect protein synthesis and stability, therefore it is important for plants to have effective ROS scavenging mechanisms.
Superoxide dismutase activity is closely associated with defense ability of plants under the condition of salt stress. Physiological correlations between elevated SOD activity and stress tolerance have been reported, suggesting that the upregulation of SOD levels may enhance the stress-defense potential of plants. Various transgenic plants that express increased amounts of SODs have been generated for enhanced tolerance to environmental stress.
To investigate the function of mnsod and czsod from C.thermophilum under salt stress in plant,the full-length mnsod and czsod cDNAs were subcloned into the expression vector pROKⅡdownstream of the 35S-CaMV promoter, respectively. The constructs were first introduced into tobacco via Agrobacterium tumefaciens LBA4404 by the freezing transformation method and verified by PCR, Southern blot and RT-PCR. It was indicated that the mnsod and czsod gene had been recombined into tobacco genome, respectively, and transgenic tobacco plants were obtained.
On the medium containing the different concentration of NaCl, expression of mnsod or czsod gene in tobacco can showed the higher percentage of seed germination. Both of the transgenic lines exhibited increased root growth compare to WT plants.These results indicated that transgenic tobacco plants could enhance salt tolerance than the wild plants at the seed germination and seedlings growth stage.
Salt stress resulted in membrane injury and caused MDA content and the extent of the membrance lipid perioxidation rising in plants. In this study, salt stress increased MDA content in all Plants. However,the values of transgenic plants measured were much lower than those of wild-type, which indicated that the extent of the membrance lipid perioxidation in transgenic tobacco was lighter than that in wild type tobacco. These results indicated that the transgenic plants showed enhanced salt tolerance compared with wild-type under salt stress conditions.
Under salt stress, the activity of SOD increased first and then decreased in both wild types and transgenic tobacco plants. Furthermore, the transgenic tobacco plants always sustained higher SOD activity than wild-type plants.
The study showed that the SOD-transformed plants exhibited increased resistance to leaf infection with the fungi Colletotrichum nicotianae and Alternaria alternate.
The enhanced tolerance of transgenic lines to salt stress suggests that the mnsod and czsod genes from C.thermophilum are benefit for genetic engineering to improve plant tolerance to salt stress.
引文
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