昆虫病原线虫共生菌的鉴定、培养及其抑菌活性研究
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摘要
昆虫病原线虫共生菌代谢产物在医药卫生和农业领域具有抗细菌、抗真菌、杀虫、杀线虫、抗病毒、抗癌等多种生物活性,是一类新型的具有较大开发潜力和广泛应用前景的生物资源。前期研究发现,从陕西杨凌、太白山筛选的3株昆虫病原线虫中分离的共生菌YL001、YL002和TB菌株代谢物具有较强的抑菌活性,为进一步确定其分类及系统进化地位,并为该生物资源的后续研究及开发应用奠定基础,本论文对其分类地位、培养及抑菌活性进行了研究,得出以下主要研究结果与结论:
1. PCR扩增、克隆并测定了YL001、YL002和TB菌株16S rRNA全序列,将此序列与GenBank中该类菌部分已知种的16S rRNA序列进行同源性比较及聚类分析。在构建的系统发育树中,3株菌与嗜线虫致病杆菌(Xenorhabdus nematophila)种内菌株形成一个类群,序列同源性均大于99%,且位于不同分支。结合前期生理生化等特征的鉴定结果,认为YL001、YL002和TB菌株属于嗜线虫致病杆菌种内的不同菌株。
2. 3株菌对供试的26种病原真菌均有不同程度拮抗作用,不同菌株对不同病菌的拮抗作用不同,但均对辣椒疫霉病菌有很强的拮抗作用;3株菌胞外代谢物对病原菌菌丝生长的抑制作用有较大差异,但其10倍稀释液对辣椒疫霉、番茄灰霉、油菜菌核和小麦根腐病菌的抑制率都在90%以上;对金黄色葡萄球菌抑制作用最强,抑菌圈直径达30 mm;活体组织试验表明3菌株对番茄灰霉病防效均在65%以上,不同菌株治疗和保护效果存在差异,其中YL002菌株治疗效果最好(77.31%),TB菌株保护效果最好(75.30%);对番茄灰霉病、辣椒疫霉病和黄瓜霜霉病的防效均在60%以上,其中YL001菌株对黄瓜霜霉病治疗效果最好(78.21%),YL002菌株对番茄灰霉病保护效果最好(77.73%),TB菌株对辣椒疫霉病保护效果最好(76.13%),TB菌株对小麦白粉病防效在60%以上,优于其他菌株,仅TB菌株对玉米大斑病有一定的保护效果。
3株菌胞外代谢物甲醇提取物对供试菌菌丝生长均有较强抑制作用,其中YL001菌株对辣椒疫霉病菌、YL002菌株对番茄灰霉病菌、TB菌株对油菜菌核病菌菌丝生长的抑制作用最强,EC50分别为:35.11、42.16和18.95 mg/L;对供试菌孢子萌发亦有较强抑制作用,其中对玉米大斑病菌抑制作用最强,EC50分别为28.03、36.77和26.40 mg/L;1000 mg/L时,对番茄灰霉病的防效均在70%以上(活体组织法),对番茄灰霉病和辣椒疫霉病的防效均在65%以上,不同菌株治疗和保护效果存在差异(盆栽试验)。
3株菌胞内代谢物乙醇提取物抑菌活性测定结果表明:1000 mg/L时,对辣椒疫霉病菌菌丝生长的抑制率在80%以上;不同菌株乙醇提取物对番茄灰霉病的防效不同,其中YL002菌株的治疗和保护效果最好,分别为79.16和61.66%(活体组织法);盆栽试验表明YL002菌株对番茄灰霉病、辣椒疫霉病和黄瓜霜霉病的治疗效果最好,分别为82.19%、70.75%和81.12%,TB菌株对其保护效果最好,分别为74.40%、65.87%和78.92%。
3.培养特性研究结果表明YL001、YL002和TB菌株的生长符合细菌生长的一般规律,发酵液中主要营养成分还原糖和氨基氮含量的变化趋势相似:在培养前期还原糖和氨基氮含量均迅速下降,后期保持相对稳定;其适宜发酵周期分别为54、66和60h。
4.综合运用Plackett–Burman设计和响应面等方法对TB菌株发酵工艺进行了整体优化。TB菌株最佳发酵培养基组成为(g/L):蛋白胨25.60、葡萄糖5.00、NaCl 5.00、K2HPO4 2.50;最佳发酵条件为:发酵时间54.07 h、初始pH 7.59、接种量9.95%、种龄(OD600: 2.00),装液量(100/250 mL)、转速150 rpm、温度25°C。最佳工业发酵培养基组成为(g/L):麸皮10.00、棉饼粉24.20、豆饼粉6.41、NaCl 5.00、K2HPO4 5.00;最佳发酵条件为:发酵时间72 h、种龄(OD600: 2.00)、装液量(50/250 mL)、初始pH 7.96、接种量10.02%、转速150 rpm。优化后TB菌株的抑菌活性较优化前分别提高了73.52%和82.58%。
5.不同培养方式对TB菌株生长和抗菌活性的研究结果表明70L发酵罐培养最有利于TB菌株的生长,DCW达16.70 g/L,较摇瓶和5L发酵罐分别增加25.85%和41.41%;摇瓶培养最有利于抑菌物质的产生,抑菌活性达358.3 U/mL,较5L和70L发酵罐分别增加10.83%和14.36%。
6.以番茄灰霉病菌和枯草芽孢杆菌为靶标菌,采用活性跟踪法,对TB菌株代谢物抑菌活性成分进行了分离。从胞内代谢物中分离出1种具有抑菌活性的化合物,结构已初步确定,为双(1-羟基-2-甲氧基乙酯)邻苯二甲酸酯;从胞外代谢物中分离出5种具有抑菌活性的化合物,其结构正在鉴定之中。活性测定结果表明:0.5 mg/mL时,4#和5#对番茄灰霉病菌菌丝生长的抑制作用较强,分别为69.1%和72.8%;4#、5#和6#对枯草芽孢杆菌的抑制作用较强,抑菌圈直径分别为10.9、11.6和10.2 mm。
The metabolites of symbiotic bacteria associated with entomopathogenic nematodes own much bioactivity such as antifungal, antibacterial, insecticidal, nematicidal, antivirus and anticancer in the field of medicine, public health and agriculture, which make it become a kind of novel bioresource with high potential for development and broad prospects for application. Previous study found that strains YL001, YL002 and TB obtained from the entomopathogenic nematodes screened from Yangling, Shaanxi Province and Taibai Moutain showed obvious antimicrobial activity. In order to determine the taxonomy and systemic evolutional status of the strains, and lay basis for further study, development and application of the bioresource, we studied their taxonomic status, culture and antimicrobial activity of the metabolites. The main results and conclusions of the thesis are as follows:
1. The 16S rRNA complete sequences of strains YL001, YL002 and TB were amplified by PCR, cloned and measured. The homology comparison and clustering analysis of the 16S rRNA sequences of the strains were conducted by comparison with the known strains in GenBank. In constructed phylogenetic tree, strains YL001, YL002 and TB formed a monophyletic clade with strains of X. nematophilus with sequence homology higher than 99%, and located at different branch. Based on previous research, we can conclude that strains YL001, YL002 and TB belong to the different strain of X. nematophila.
2. All the three strains exhibited antagonism effect on tested 26 plant pathogenic fungi, and different strains showed different antagonism effect, but all showed high effect on P. capsici. The extracellular metabolite of different strains exhibited a greater difference in inhibitory effect on pathogen mycelium, but all exhibited high inhibitory rate (>90%) on mycelium growth of P. capsici, B. cinerea, S. sclerotiorum and B. sorokiniana at ten times diluted liquid. The strains all showed strongest inhibitory effect on S. aureus. Tissue experiment showed that the efficacy of the strains on B. cinerea all higher than 65%, and different strains showed different therapeutic and protective effect on B. cinerea, among which, YL002 showed best therapeutic effect (77.31%) and TB showed best protective effect (75.30%). Potted plant experiment showed that the efficacy of the strains on B. cinerea, P. capsici and P. cubensis all higher than 60%, among which, YL001 showed best therapeutic effect (78.21%) on P. cubensis, YL002 showed best protective effect (77.73%) on B. cinerea and TB showed best protective effect (76.13%) on P. capsici; The efficacy of strain TB to B. graminis was higher than 60%, which was better that the other strains; only strain TB showed some protective effect on E. turctcum.
The methanol extracts of the three strains all showed strong effect on mycelium growth of tested pathogens, among which, YL001 showed strongest effect on P. capsici, YL002 showed strongest effect on B. cinerea and TB showed strongest effect on S. sclerotiorum, with EC50 of 35.11, 42.16 and 18.95 mg/L, respectively. The methanol extracts all showed strongest effect on spore germination of tested pathogens, and all exhibited highest inhibitory effect on E. turctcum with EC50 of 28.03, 36.77and 26.40 mg/L. At 1000 mg/L, the efficacy of methanol extracts to B. cirerea (tissue experiment) were all higher than 70% and to B. cirerea and P. capsici were all higher than 65% (potted plant experiment), different strains showed different therapeutic and protective effect on B. cirerea and P. capsici.
At 1000 mg/L, the inhibitory rates of ethanol extracts of the three strains on mycelium growth of P. capsici were all higher than 80%. Tissue experiment showed that ethanol extracts of different strains showed different efficacy to B. cirerea, among which, strain YL002 showed strongest therapeutic effect (79.16%) and protective effect (61.66%). Potted plant experiment showed that strain YL002 showed best therapeutic effect on B. cirerea, P. capsici and P. cubensis with 82.19, 70.75 and 81.12%, respectively, and strain TB showed best protective effect with 74.40, 65.87 and 78.92%, respectively.
3. Study on the culture characteristics of strains YL001, YL002 and TB showed that the strains growth was in accordance with the general discipline of bacteria growth; The changing trends of glucose and amido nitrogen of the strains were similar, which decreased sharply at first and then kept stable; The optimum fermentation period was 54, 66 and 66h, respectively.
4. Statistical methods such as Plackett–Burman design and response surface methodology were employed synthetically to optimize the fermentation medium and conditions together so as to improve antibiotic activity of X. nematophila TB. The optimum composition of the fermentation medium (g/L): peptone 25.60, glucose 5.00, NaCl 5.00, K_2HPO_4 2.50, and the optimum fermentation condition: fermentation time 54.07 h,initial pH 7.59, inoculation volume 9.95%, inoculation age (OD_(600): 2.00), medium volume (100/250 mL),rotary speed 150 rpm,temperature 25°C. The optimum composition of the industrial fermentation medium (g/L): bran 10.00, cotton cake powder 24.20, bean cake powder 6.41, NaCl 5.00, K_2HPO_4 5.00, and the optimum fermentation condition: fermentation time 72 h, inoculation age (OD_(600): 2.00), medium volume (50/250 mL), initial pH 7.96, inoculation volume 10.02%, rotary speed 150 rpm. After optimization, the antibiotic activity of strain TB improved by 73.52% and 82.58%, respectively.
5. Experiments of different culture mode on cell growth and antibiotic activity of strain TB showed that 70L fermenter was optimum for the cell growth with DCW of 16.70 g/L, which improved by 25.85 and 41.41% than that of 250 mL flask and 5L fermenter, respectively. Flask was optimum for the antibiotic production with antibiotic activity of 358.3 U/mL, which improved by 10.83 and 14.36% than that of 5L and 70L fermenter.
6. The antimicrobial components of the extra and intra cellular metabolite of strain TB was separated based on activity tracking using B. cirerea and B. subtilis as the target pathogens. One compound with antimicrobial activity was separated from the intracellular metabolite, and its structure was determined initially, which was bis (1-hydroxy-2-methoxye thy) phthalate. Five compounds were separated from extracellular metabolite, and their structures are under identification. Activity experiments indicated that 4#and 5# showed stronger inhibitory effect on mycelium growth of B. cirerea, which was 69.1 and 72.8%, respectively; 4#, 5# and 6#showed stronger inhibitory effect on B. subtilis with the inhibition zone of 10.9, 11.6 and 10.2 mm, respectively.
1. PCR扩增、克隆并测定了YL001、YL002和TB菌株16S rRNA全序列,将此序列与GenBank中该类菌部分已知种的16S rRNA序列进行同源性比较及聚类分析。在构建的系统发育树中,3株菌与嗜线虫致病杆菌(Xenorhabdus nematophila)种内菌株形成一个类群,序列同源性均大于99%,且位于不同分支。结合前期生理生化等特征的鉴定结果,认为YL001、YL002和TB菌株属于嗜线虫致病杆菌种内的不同菌株。
2. 3株菌对供试的26种病原真菌均有不同程度拮抗作用,不同菌株对不同病菌的拮抗作用不同,但均对辣椒疫霉病菌有很强的拮抗作用;3株菌胞外代谢物对病原菌菌丝生长的抑制作用有较大差异,但其10倍稀释液对辣椒疫霉、番茄灰霉、油菜菌核和小麦根腐病菌的抑制率都在90%以上;对金黄色葡萄球菌抑制作用最强,抑菌圈直径达30 mm;活体组织试验表明3菌株对番茄灰霉病防效均在65%以上,不同菌株治疗和保护效果存在差异,其中YL002菌株治疗效果最好(77.31%),TB菌株保护效果最好(75.30%);对番茄灰霉病、辣椒疫霉病和黄瓜霜霉病的防效均在60%以上,其中YL001菌株对黄瓜霜霉病治疗效果最好(78.21%),YL002菌株对番茄灰霉病保护效果最好(77.73%),TB菌株对辣椒疫霉病保护效果最好(76.13%),TB菌株对小麦白粉病防效在60%以上,优于其他菌株,仅TB菌株对玉米大斑病有一定的保护效果。
3株菌胞外代谢物甲醇提取物对供试菌菌丝生长均有较强抑制作用,其中YL001菌株对辣椒疫霉病菌、YL002菌株对番茄灰霉病菌、TB菌株对油菜菌核病菌菌丝生长的抑制作用最强,EC50分别为:35.11、42.16和18.95 mg/L;对供试菌孢子萌发亦有较强抑制作用,其中对玉米大斑病菌抑制作用最强,EC50分别为28.03、36.77和26.40 mg/L;1000 mg/L时,对番茄灰霉病的防效均在70%以上(活体组织法),对番茄灰霉病和辣椒疫霉病的防效均在65%以上,不同菌株治疗和保护效果存在差异(盆栽试验)。
3株菌胞内代谢物乙醇提取物抑菌活性测定结果表明:1000 mg/L时,对辣椒疫霉病菌菌丝生长的抑制率在80%以上;不同菌株乙醇提取物对番茄灰霉病的防效不同,其中YL002菌株的治疗和保护效果最好,分别为79.16和61.66%(活体组织法);盆栽试验表明YL002菌株对番茄灰霉病、辣椒疫霉病和黄瓜霜霉病的治疗效果最好,分别为82.19%、70.75%和81.12%,TB菌株对其保护效果最好,分别为74.40%、65.87%和78.92%。
3.培养特性研究结果表明YL001、YL002和TB菌株的生长符合细菌生长的一般规律,发酵液中主要营养成分还原糖和氨基氮含量的变化趋势相似:在培养前期还原糖和氨基氮含量均迅速下降,后期保持相对稳定;其适宜发酵周期分别为54、66和60h。
4.综合运用Plackett–Burman设计和响应面等方法对TB菌株发酵工艺进行了整体优化。TB菌株最佳发酵培养基组成为(g/L):蛋白胨25.60、葡萄糖5.00、NaCl 5.00、K2HPO4 2.50;最佳发酵条件为:发酵时间54.07 h、初始pH 7.59、接种量9.95%、种龄(OD600: 2.00),装液量(100/250 mL)、转速150 rpm、温度25°C。最佳工业发酵培养基组成为(g/L):麸皮10.00、棉饼粉24.20、豆饼粉6.41、NaCl 5.00、K2HPO4 5.00;最佳发酵条件为:发酵时间72 h、种龄(OD600: 2.00)、装液量(50/250 mL)、初始pH 7.96、接种量10.02%、转速150 rpm。优化后TB菌株的抑菌活性较优化前分别提高了73.52%和82.58%。
5.不同培养方式对TB菌株生长和抗菌活性的研究结果表明70L发酵罐培养最有利于TB菌株的生长,DCW达16.70 g/L,较摇瓶和5L发酵罐分别增加25.85%和41.41%;摇瓶培养最有利于抑菌物质的产生,抑菌活性达358.3 U/mL,较5L和70L发酵罐分别增加10.83%和14.36%。
6.以番茄灰霉病菌和枯草芽孢杆菌为靶标菌,采用活性跟踪法,对TB菌株代谢物抑菌活性成分进行了分离。从胞内代谢物中分离出1种具有抑菌活性的化合物,结构已初步确定,为双(1-羟基-2-甲氧基乙酯)邻苯二甲酸酯;从胞外代谢物中分离出5种具有抑菌活性的化合物,其结构正在鉴定之中。活性测定结果表明:0.5 mg/mL时,4#和5#对番茄灰霉病菌菌丝生长的抑制作用较强,分别为69.1%和72.8%;4#、5#和6#对枯草芽孢杆菌的抑制作用较强,抑菌圈直径分别为10.9、11.6和10.2 mm。
The metabolites of symbiotic bacteria associated with entomopathogenic nematodes own much bioactivity such as antifungal, antibacterial, insecticidal, nematicidal, antivirus and anticancer in the field of medicine, public health and agriculture, which make it become a kind of novel bioresource with high potential for development and broad prospects for application. Previous study found that strains YL001, YL002 and TB obtained from the entomopathogenic nematodes screened from Yangling, Shaanxi Province and Taibai Moutain showed obvious antimicrobial activity. In order to determine the taxonomy and systemic evolutional status of the strains, and lay basis for further study, development and application of the bioresource, we studied their taxonomic status, culture and antimicrobial activity of the metabolites. The main results and conclusions of the thesis are as follows:
1. The 16S rRNA complete sequences of strains YL001, YL002 and TB were amplified by PCR, cloned and measured. The homology comparison and clustering analysis of the 16S rRNA sequences of the strains were conducted by comparison with the known strains in GenBank. In constructed phylogenetic tree, strains YL001, YL002 and TB formed a monophyletic clade with strains of X. nematophilus with sequence homology higher than 99%, and located at different branch. Based on previous research, we can conclude that strains YL001, YL002 and TB belong to the different strain of X. nematophila.
2. All the three strains exhibited antagonism effect on tested 26 plant pathogenic fungi, and different strains showed different antagonism effect, but all showed high effect on P. capsici. The extracellular metabolite of different strains exhibited a greater difference in inhibitory effect on pathogen mycelium, but all exhibited high inhibitory rate (>90%) on mycelium growth of P. capsici, B. cinerea, S. sclerotiorum and B. sorokiniana at ten times diluted liquid. The strains all showed strongest inhibitory effect on S. aureus. Tissue experiment showed that the efficacy of the strains on B. cinerea all higher than 65%, and different strains showed different therapeutic and protective effect on B. cinerea, among which, YL002 showed best therapeutic effect (77.31%) and TB showed best protective effect (75.30%). Potted plant experiment showed that the efficacy of the strains on B. cinerea, P. capsici and P. cubensis all higher than 60%, among which, YL001 showed best therapeutic effect (78.21%) on P. cubensis, YL002 showed best protective effect (77.73%) on B. cinerea and TB showed best protective effect (76.13%) on P. capsici; The efficacy of strain TB to B. graminis was higher than 60%, which was better that the other strains; only strain TB showed some protective effect on E. turctcum.
The methanol extracts of the three strains all showed strong effect on mycelium growth of tested pathogens, among which, YL001 showed strongest effect on P. capsici, YL002 showed strongest effect on B. cinerea and TB showed strongest effect on S. sclerotiorum, with EC50 of 35.11, 42.16 and 18.95 mg/L, respectively. The methanol extracts all showed strongest effect on spore germination of tested pathogens, and all exhibited highest inhibitory effect on E. turctcum with EC50 of 28.03, 36.77and 26.40 mg/L. At 1000 mg/L, the efficacy of methanol extracts to B. cirerea (tissue experiment) were all higher than 70% and to B. cirerea and P. capsici were all higher than 65% (potted plant experiment), different strains showed different therapeutic and protective effect on B. cirerea and P. capsici.
At 1000 mg/L, the inhibitory rates of ethanol extracts of the three strains on mycelium growth of P. capsici were all higher than 80%. Tissue experiment showed that ethanol extracts of different strains showed different efficacy to B. cirerea, among which, strain YL002 showed strongest therapeutic effect (79.16%) and protective effect (61.66%). Potted plant experiment showed that strain YL002 showed best therapeutic effect on B. cirerea, P. capsici and P. cubensis with 82.19, 70.75 and 81.12%, respectively, and strain TB showed best protective effect with 74.40, 65.87 and 78.92%, respectively.
3. Study on the culture characteristics of strains YL001, YL002 and TB showed that the strains growth was in accordance with the general discipline of bacteria growth; The changing trends of glucose and amido nitrogen of the strains were similar, which decreased sharply at first and then kept stable; The optimum fermentation period was 54, 66 and 66h, respectively.
4. Statistical methods such as Plackett–Burman design and response surface methodology were employed synthetically to optimize the fermentation medium and conditions together so as to improve antibiotic activity of X. nematophila TB. The optimum composition of the fermentation medium (g/L): peptone 25.60, glucose 5.00, NaCl 5.00, K_2HPO_4 2.50, and the optimum fermentation condition: fermentation time 54.07 h,initial pH 7.59, inoculation volume 9.95%, inoculation age (OD_(600): 2.00), medium volume (100/250 mL),rotary speed 150 rpm,temperature 25°C. The optimum composition of the industrial fermentation medium (g/L): bran 10.00, cotton cake powder 24.20, bean cake powder 6.41, NaCl 5.00, K_2HPO_4 5.00, and the optimum fermentation condition: fermentation time 72 h, inoculation age (OD_(600): 2.00), medium volume (50/250 mL), initial pH 7.96, inoculation volume 10.02%, rotary speed 150 rpm. After optimization, the antibiotic activity of strain TB improved by 73.52% and 82.58%, respectively.
5. Experiments of different culture mode on cell growth and antibiotic activity of strain TB showed that 70L fermenter was optimum for the cell growth with DCW of 16.70 g/L, which improved by 25.85 and 41.41% than that of 250 mL flask and 5L fermenter, respectively. Flask was optimum for the antibiotic production with antibiotic activity of 358.3 U/mL, which improved by 10.83 and 14.36% than that of 5L and 70L fermenter.
6. The antimicrobial components of the extra and intra cellular metabolite of strain TB was separated based on activity tracking using B. cirerea and B. subtilis as the target pathogens. One compound with antimicrobial activity was separated from the intracellular metabolite, and its structure was determined initially, which was bis (1-hydroxy-2-methoxye thy) phthalate. Five compounds were separated from extracellular metabolite, and their structures are under identification. Activity experiments indicated that 4#and 5# showed stronger inhibitory effect on mycelium growth of B. cirerea, which was 69.1 and 72.8%, respectively; 4#, 5# and 6#showed stronger inhibitory effect on B. subtilis with the inhibition zone of 10.9, 11.6 and 10.2 mm, respectively.
引文
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