拮抗植物寄生线虫的细菌菌株筛选及其杀线虫相关基因的鉴定
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摘要
植物寄生线虫是世界范围内严重危害农业生产的重要病原生物之一,每年给全世界主要农作物生产造成的损失高达1250亿美元。植物寄生线虫地理分布和寄主范围非常广泛,并且多数生存在土壤中,因此给防治带来很大的困难。目前,常采用化学防治、改进栽培措施和使用抗性品种等来防治植物寄生线虫,但是都有其局限性。因此,植物寄生线虫的生物防治引起了人们越来越多的重视。芽孢杆菌是一类植物根围促生细菌,能够防治多种植物寄生线虫、真菌和细菌等病害。由于其生长快、易于培养且对线虫有较好的杀灭作用等特性,已成为生物防治线虫新的研究热点。在研究中发现大肠杆菌具有杀线虫的能力。大肠杆菌是分子生物学操作中常用的一类菌株,关于其具有杀线作用的研究还未见报道。本研究以芽孢杆菌和大肠杆菌为材料,对拮抗植物寄生线虫的两种细菌菌株进行了筛选,并对其杀线活性物质的性质、杀线相关基因进行了研究,还对这两种细菌的温室生防实验效果进行了评估。主要研究结果如下：
1、选取2个枯草芽孢杆菌(Bacillus subtilis)菌株OKB105、69和2个解淀粉芽孢杆菌(Bacillus amyloliquefaciens)菌株FZB42、B3,经LANDY培养基发酵,离心后取培养滤液处理线虫,离体条件下测定4个菌株的培养滤液对植物寄生线虫水稻干尖线虫(Aphelenchoides besseyi)、马铃薯腐烂线虫(Ditylenchus detructor)、松材线虫(Bursaphelenchus xylophilus)和爪哇根结线虫(Meloidogyne javanica)的杀线率。结果显示：处理12h后观察,对水稻干尖线虫防效最佳的是B3菌株,杀线率达10.6%;对马铃薯腐烂线虫防效最佳的是OKB105菌株,杀线率达27.6%；对松材线虫防效最佳的是69菌株,杀线率达35.6%；对爪哇根结线虫防效最佳的是OKB105菌株,杀线率为100%。选择OKB105菌株和爪哇根结线虫作为进一步研究的对象,结果显示OKB105菌株培养滤液原液稀释至30%,处理60h杀线率仍为100%;OKB105菌株培养滤液原液还能抑制爪哇根结线虫的卵块孵化,抑制率为98.3%。温室生防实验显示：OKB105菌株处理的番茄苗与对照植株比较,单株鲜根重有显著性差异,单株卵块数则没有显著性差异。
2、枯草芽孢杆菌(B.subtilis) OKB105菌株对爪哇根结线虫(M. jvanica)具有较高的杀线活性。本试验对OKB105菌株分泌的杀线活性物质的基本性质进行了研究。结果表明：该菌株产生的杀线活性物质是可随水分一同蒸发的非蛋白质类物质；分子量小于1kDa；该物质在4℃、-20℃下保存40d仍能保持杀线活性；对热不敏感；碱性条件下具有较高的杀线活性；极性强,正丁醇、氯仿、乙酸乙酯等有机溶剂不能将其活性物质从培养滤液中提取。
3、对构建的枯草芽孢杆菌OKB105菌株突变体文库中的2000个突变体进行筛选,结果表明：1个突变体M1在72h时的杀线率为0%。反向PCR扩增结果表明M1菌株中的purL基因被破坏。purL基因编码甲酰甘氨脒核苷酸合成酶II(Phosphoribosylformylglycinamidine synthase II)在嘌呤生物合成途径中催化甲酰甘氨酰胺核苷酸合成甲酰甘氨脒核苷酸。构建两种质粒pMA5-purL和pUC18-purL对M1突变体进行活性回复,结果证明purL基因在OKB105菌株的杀线活性中起着非常重要的作用。这也是首次报道purL基因影响枯草芽孢杆菌的杀线活性。M1菌株不仅能通过功能互补来回复杀线活性,而且在含有5-氨基咪唑-4-甲酰胺核苷酸(5-aminoimidazole-4-carboxamide riboside,AICAR)或腺嘌呤(Adenine)和硫胺(Thiamine)的LANDY培养基中发酵后,其杀线活性也能回复。结果显示purL基因是通过调控嘌呤生物合成中的一些代谢中间产物来影响OKB105菌株的杀线活性。此外,purL基因还影响M1菌株的培养性状。
4、从实验中选出六个大肠杆菌(Escherichia coli)菌株用LANDY培养基发酵,离心后取培养滤液处理线虫,离体条件下测定菌株的培养滤液原液对爪哇根结线虫(M.javanica)的杀线率。结果显示：处理12h后观察,6株大肠杆菌的杀线率都为100%,并且都能使线虫虫体内部结构发生改变,但是线虫虫体内部结构出现变化的时间不一致。以BL21菌株作为进一步研究的对象。结果显示BL21菌株培养滤液原液稀释至20%,处理36h杀线率仍为100%。对其分泌的杀线活性物质的基本性质进行了初步研究。结果表明：该菌株产生的杀线活性物质可随水分一同蒸发；4℃、-25℃下保存90d仍能保持杀线活性；对热不敏感；在一定的酸性范围内具有较高的杀线活性。BL21菌株培养滤液还能抑制爪哇根结线虫的卵块孵化,抑制率为98.9%。温室生防实验显示：处理组番茄苗与对照组番茄苗相比,单株鲜根重和单株卵块数都有显著性差异。
5、为了找到BL21菌株的杀线虫相关基因,利用Tn5转座子构建了大肠杆菌BL21菌株的突变体文库。对该文库的5000个突变体进行筛选,结果表明：2个突变体M12、M13在72h时的杀线率为0%,14个突变体能延缓线虫虫体的变化。研究结果表明这16个突变体都是通过改变酸性环境来影响其杀线活性。采用TAIL-PCR技术对这16个突变体Tn5转座子插入的杀线相关基因进行了鉴定。确定的7个突变体M2、M4、 M5、M7、M9、M10、M12的Tn5转座子插入的基因分别是cysH、serA、metC、pyrD、 ilvC、purA、carB,其余的9个突变体没有获得结果。
Plant-parasitic nematodes, known as one of the important plant pathogens, caused huge damage to many economically crops. The annual loss in agriculture caused by plant-parasitic nematodes amounted to＄125billion worldwide. It is difficult to control plant-parasitic nematodes because of their wide geographic distribution and host range, and most of them living in the soil. Currently, several methods, such as chemical control, cultivation measures and resistant cultivar, have been used to control plant-parasitic nematodes. But those control methods have led to limited success. Biocontrol of plant-parasitic nematodes has therefore given rise to vast attention. Bacillus spp, one kind of plant growth-promoting rhizobacteria(PGPR), can control nematode, fungal, bacterial, and other disease. Bacillus spp. controlling the nematodes has become a new hotspot for its fast growth, easily culturing, having a good effect on nematodes. Escherichia spp, one type of strains used in molecular biology in the lab, is not been reported that they are able to act against nematodes. The present work includes:screening of two types of bacteria antagonistic against plant-parasitic nematodes; characterizing the property of nematicidal activity substances respectively produced by OKB105and BL21; identifying the nematicidal-related genes; assessing control efficacy in the greenhouse experiment. The results obtained as below:
1. Bacillus strains OKB105,69, FZB42and B3were selected to ferment in the LANDY culture medium. The results had showed that the LANDY ferment filtrates of Bacillus strains OKB105,69, and B3all had the nematicidal activity toward A. besseyi, D. detructor, B. xylophilus and M. javanica, and the strain FZB42had the antagonistic activity toward other three nematodes except A. besseyi. The highest mortality at12hours, which were obtained from four optimum treatments A. besseyis/B3, D. detructor/OKB105, B. xylophilu169and M. javanica/OKB105,were10.6%,27.6%,35.6%and100%, respectively. The M. javanica/OKB105treatment was used for the further study. Low dilution culture filtrates of OKB105also showed a strong effect on M. javanica. The mortality rates of the nematodes tested after incubation for60h in30%culture filtrates were100%. The culture filtrates of OKB105could also inhibit the eggs of M. javanica hatching and the inhibition rate was98.3%. The greenhouse experiments showed that the fresh root weight/per plant had significant difference at the0.01levels, but the number of egg masses/per plant had no significant difference at the0.05levels, compared with the control.
2. Bacillus subtilis strain OKB105had high nematicidal activity to M. jvanica. In order to isolate and purify the nematicidal components, the elementary characteristics of the nematicidal active substances had been studied in our experiment. The results demonstrated that the nematicidal active ingredients from the culture filtrates of OKB105were nonprotein, and had thermophilic, psychrotolerant and strong polar characteristics, and could be evaporated with water but not extracted by the organic solvent, such as ethyl acetate, chloroform, normal butanol. The molecular weight of the nematicidal ingredients was less than1000Da. pH could affect its nematicidal activity. It had strong nematicidal activity at the alkalinity condition.
3. To identify the nematicidal-related genes,2000mutants of OKB105were screened. Mutant M1had eliminated the nematicidal activity compared with the original strain OKB105at72h. The inverse PCR result had showed that the purL gene of M1mutant was disrupted by transposon TnYLB-1. To restore the function of the purL gene, two plasmids pMA5-purL and pUC18-purL were constructed and transformed into M1mutant. The two complementation of the M1mutant showed the same activity with the original strain OKB105. Those results indicated that the purL gene of OKB105played a key role in the nematicidal activity. This was the first report that the purL gene affected the nematicidal activity of Bacillus subtilis strain OKB105. The purL gene encodes the5-phosphoribos-ylformylglycinamidine synthase Ⅱ, which involves in the purine biosythesis pathway. Interestingly, when M1mutant growed in LANDY medium supplemented with ademine and thiamin or5-aminoimidazole-4-carboxamide riboside, the culture filtrates were also restored the same nematicidal activity. We concluded that the purL gene regulated the biosythesis of the intermediates in purine synthesis pathway to affect the nematicidal activity of Bacillus subtilis strian OKB105. And the purL gene also affected the culture characteristics of the M1mutant.
4. The LANDY ferment filtrates of Escherichia strains TOP10, JM109, DH5α, BL21, Rosseta and K88all showed the nematicidal effects, killing100%M. javanica, and made the body of nematodes change. But the time of the body variation was different. The strain BL21was used for the further study. Low dilution culture filtrates of BL21also showed a strong effect on M. javanica. The mortality rates of the nematodes tested after incubation for36hours in20%culture filtrates were100%. The elementary characteristics of the nematicidal substance of BL21had been studied in our lab. The results demonstrated that the active ingredients from the culture filtrates were thermophilic, psychrotolerant and could be evaporated with water. pH could affect its nematicidal activity. It had strong nematicidal activity at a certain acid condition. The culture filtrates of BL21could also inhibit the eggs of M. javanica hatching and the inhibition rate was98.9%. The greenhouse experiments showed that fresh root weight/per plant at the0.01levels and the number of egg masses/per plant at the0.05levels was significant difference, compared with the control.
5. In order to identify the nematicidal-related genes, a mutant library of Escherichia coli BL21(DE3) was constructed by using Tn5transposon. The nematicidal proterties of5000mutants were evalated and2mutants M12, M13were unable to kill the root-knot nematodes M. javanica in72hours,14mutant could delay the body variation of the nematodes. The results showed that the nematicidal activity of16mutants was affected by changing their acidic environments. Hermal asymmetric interlaced PCR (TAIL-PCR) was adopted to verify16mutants screened from the Tn5inserted library of the strain BL21based on nematicidal assay. The results revealed that the inserted genes of7mutans M2, M4, M5, M7, M9, M10, M12were confirmed, which were cysH、serA、metC、pyrD、 ilvC、purA、 carB genes, and the other9mutants couldnot get the results.
1、选取2个枯草芽孢杆菌(Bacillus subtilis)菌株OKB105、69和2个解淀粉芽孢杆菌(Bacillus amyloliquefaciens)菌株FZB42、B3,经LANDY培养基发酵,离心后取培养滤液处理线虫,离体条件下测定4个菌株的培养滤液对植物寄生线虫水稻干尖线虫(Aphelenchoides besseyi)、马铃薯腐烂线虫(Ditylenchus detructor)、松材线虫(Bursaphelenchus xylophilus)和爪哇根结线虫(Meloidogyne javanica)的杀线率。结果显示：处理12h后观察,对水稻干尖线虫防效最佳的是B3菌株,杀线率达10.6%;对马铃薯腐烂线虫防效最佳的是OKB105菌株,杀线率达27.6%；对松材线虫防效最佳的是69菌株,杀线率达35.6%；对爪哇根结线虫防效最佳的是OKB105菌株,杀线率为100%。选择OKB105菌株和爪哇根结线虫作为进一步研究的对象,结果显示OKB105菌株培养滤液原液稀释至30%,处理60h杀线率仍为100%;OKB105菌株培养滤液原液还能抑制爪哇根结线虫的卵块孵化,抑制率为98.3%。温室生防实验显示：OKB105菌株处理的番茄苗与对照植株比较,单株鲜根重有显著性差异,单株卵块数则没有显著性差异。
2、枯草芽孢杆菌(B.subtilis) OKB105菌株对爪哇根结线虫(M. jvanica)具有较高的杀线活性。本试验对OKB105菌株分泌的杀线活性物质的基本性质进行了研究。结果表明：该菌株产生的杀线活性物质是可随水分一同蒸发的非蛋白质类物质；分子量小于1kDa；该物质在4℃、-20℃下保存40d仍能保持杀线活性；对热不敏感；碱性条件下具有较高的杀线活性；极性强,正丁醇、氯仿、乙酸乙酯等有机溶剂不能将其活性物质从培养滤液中提取。
3、对构建的枯草芽孢杆菌OKB105菌株突变体文库中的2000个突变体进行筛选,结果表明：1个突变体M1在72h时的杀线率为0%。反向PCR扩增结果表明M1菌株中的purL基因被破坏。purL基因编码甲酰甘氨脒核苷酸合成酶II(Phosphoribosylformylglycinamidine synthase II)在嘌呤生物合成途径中催化甲酰甘氨酰胺核苷酸合成甲酰甘氨脒核苷酸。构建两种质粒pMA5-purL和pUC18-purL对M1突变体进行活性回复,结果证明purL基因在OKB105菌株的杀线活性中起着非常重要的作用。这也是首次报道purL基因影响枯草芽孢杆菌的杀线活性。M1菌株不仅能通过功能互补来回复杀线活性,而且在含有5-氨基咪唑-4-甲酰胺核苷酸(5-aminoimidazole-4-carboxamide riboside,AICAR)或腺嘌呤(Adenine)和硫胺(Thiamine)的LANDY培养基中发酵后,其杀线活性也能回复。结果显示purL基因是通过调控嘌呤生物合成中的一些代谢中间产物来影响OKB105菌株的杀线活性。此外,purL基因还影响M1菌株的培养性状。
4、从实验中选出六个大肠杆菌(Escherichia coli)菌株用LANDY培养基发酵,离心后取培养滤液处理线虫,离体条件下测定菌株的培养滤液原液对爪哇根结线虫(M.javanica)的杀线率。结果显示：处理12h后观察,6株大肠杆菌的杀线率都为100%,并且都能使线虫虫体内部结构发生改变,但是线虫虫体内部结构出现变化的时间不一致。以BL21菌株作为进一步研究的对象。结果显示BL21菌株培养滤液原液稀释至20%,处理36h杀线率仍为100%。对其分泌的杀线活性物质的基本性质进行了初步研究。结果表明：该菌株产生的杀线活性物质可随水分一同蒸发；4℃、-25℃下保存90d仍能保持杀线活性；对热不敏感；在一定的酸性范围内具有较高的杀线活性。BL21菌株培养滤液还能抑制爪哇根结线虫的卵块孵化,抑制率为98.9%。温室生防实验显示：处理组番茄苗与对照组番茄苗相比,单株鲜根重和单株卵块数都有显著性差异。
5、为了找到BL21菌株的杀线虫相关基因,利用Tn5转座子构建了大肠杆菌BL21菌株的突变体文库。对该文库的5000个突变体进行筛选,结果表明：2个突变体M12、M13在72h时的杀线率为0%,14个突变体能延缓线虫虫体的变化。研究结果表明这16个突变体都是通过改变酸性环境来影响其杀线活性。采用TAIL-PCR技术对这16个突变体Tn5转座子插入的杀线相关基因进行了鉴定。确定的7个突变体M2、M4、 M5、M7、M9、M10、M12的Tn5转座子插入的基因分别是cysH、serA、metC、pyrD、 ilvC、purA、carB,其余的9个突变体没有获得结果。
Plant-parasitic nematodes, known as one of the important plant pathogens, caused huge damage to many economically crops. The annual loss in agriculture caused by plant-parasitic nematodes amounted to＄125billion worldwide. It is difficult to control plant-parasitic nematodes because of their wide geographic distribution and host range, and most of them living in the soil. Currently, several methods, such as chemical control, cultivation measures and resistant cultivar, have been used to control plant-parasitic nematodes. But those control methods have led to limited success. Biocontrol of plant-parasitic nematodes has therefore given rise to vast attention. Bacillus spp, one kind of plant growth-promoting rhizobacteria(PGPR), can control nematode, fungal, bacterial, and other disease. Bacillus spp. controlling the nematodes has become a new hotspot for its fast growth, easily culturing, having a good effect on nematodes. Escherichia spp, one type of strains used in molecular biology in the lab, is not been reported that they are able to act against nematodes. The present work includes:screening of two types of bacteria antagonistic against plant-parasitic nematodes; characterizing the property of nematicidal activity substances respectively produced by OKB105and BL21; identifying the nematicidal-related genes; assessing control efficacy in the greenhouse experiment. The results obtained as below:
1. Bacillus strains OKB105,69, FZB42and B3were selected to ferment in the LANDY culture medium. The results had showed that the LANDY ferment filtrates of Bacillus strains OKB105,69, and B3all had the nematicidal activity toward A. besseyi, D. detructor, B. xylophilus and M. javanica, and the strain FZB42had the antagonistic activity toward other three nematodes except A. besseyi. The highest mortality at12hours, which were obtained from four optimum treatments A. besseyis/B3, D. detructor/OKB105, B. xylophilu169and M. javanica/OKB105,were10.6%,27.6%,35.6%and100%, respectively. The M. javanica/OKB105treatment was used for the further study. Low dilution culture filtrates of OKB105also showed a strong effect on M. javanica. The mortality rates of the nematodes tested after incubation for60h in30%culture filtrates were100%. The culture filtrates of OKB105could also inhibit the eggs of M. javanica hatching and the inhibition rate was98.3%. The greenhouse experiments showed that the fresh root weight/per plant had significant difference at the0.01levels, but the number of egg masses/per plant had no significant difference at the0.05levels, compared with the control.
2. Bacillus subtilis strain OKB105had high nematicidal activity to M. jvanica. In order to isolate and purify the nematicidal components, the elementary characteristics of the nematicidal active substances had been studied in our experiment. The results demonstrated that the nematicidal active ingredients from the culture filtrates of OKB105were nonprotein, and had thermophilic, psychrotolerant and strong polar characteristics, and could be evaporated with water but not extracted by the organic solvent, such as ethyl acetate, chloroform, normal butanol. The molecular weight of the nematicidal ingredients was less than1000Da. pH could affect its nematicidal activity. It had strong nematicidal activity at the alkalinity condition.
3. To identify the nematicidal-related genes,2000mutants of OKB105were screened. Mutant M1had eliminated the nematicidal activity compared with the original strain OKB105at72h. The inverse PCR result had showed that the purL gene of M1mutant was disrupted by transposon TnYLB-1. To restore the function of the purL gene, two plasmids pMA5-purL and pUC18-purL were constructed and transformed into M1mutant. The two complementation of the M1mutant showed the same activity with the original strain OKB105. Those results indicated that the purL gene of OKB105played a key role in the nematicidal activity. This was the first report that the purL gene affected the nematicidal activity of Bacillus subtilis strain OKB105. The purL gene encodes the5-phosphoribos-ylformylglycinamidine synthase Ⅱ, which involves in the purine biosythesis pathway. Interestingly, when M1mutant growed in LANDY medium supplemented with ademine and thiamin or5-aminoimidazole-4-carboxamide riboside, the culture filtrates were also restored the same nematicidal activity. We concluded that the purL gene regulated the biosythesis of the intermediates in purine synthesis pathway to affect the nematicidal activity of Bacillus subtilis strian OKB105. And the purL gene also affected the culture characteristics of the M1mutant.
4. The LANDY ferment filtrates of Escherichia strains TOP10, JM109, DH5α, BL21, Rosseta and K88all showed the nematicidal effects, killing100%M. javanica, and made the body of nematodes change. But the time of the body variation was different. The strain BL21was used for the further study. Low dilution culture filtrates of BL21also showed a strong effect on M. javanica. The mortality rates of the nematodes tested after incubation for36hours in20%culture filtrates were100%. The elementary characteristics of the nematicidal substance of BL21had been studied in our lab. The results demonstrated that the active ingredients from the culture filtrates were thermophilic, psychrotolerant and could be evaporated with water. pH could affect its nematicidal activity. It had strong nematicidal activity at a certain acid condition. The culture filtrates of BL21could also inhibit the eggs of M. javanica hatching and the inhibition rate was98.9%. The greenhouse experiments showed that fresh root weight/per plant at the0.01levels and the number of egg masses/per plant at the0.05levels was significant difference, compared with the control.
5. In order to identify the nematicidal-related genes, a mutant library of Escherichia coli BL21(DE3) was constructed by using Tn5transposon. The nematicidal proterties of5000mutants were evalated and2mutants M12, M13were unable to kill the root-knot nematodes M. javanica in72hours,14mutant could delay the body variation of the nematodes. The results showed that the nematicidal activity of16mutants was affected by changing their acidic environments. Hermal asymmetric interlaced PCR (TAIL-PCR) was adopted to verify16mutants screened from the Tn5inserted library of the strain BL21based on nematicidal assay. The results revealed that the inserted genes of7mutans M2, M4, M5, M7, M9, M10, M12were confirmed, which were cysH、serA、metC、pyrD、 ilvC、purA、 carB genes, and the other9mutants couldnot get the results.
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