植物细菌性青枯病颉颃菌筛选策略研究及生防机理初探
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摘要
由青枯劳尔氏菌(Ralstonia solanacearum Yabuuchi et al.)引起的作物细菌性萎蔫病是一种毁灭性的土传病害,是中国南方许多省份农业生产中的一大威胁。目前生产上主要是采用抗病品种和轮作套种等农业措施和化学药物来防治青枯病,但由于受到各种条件和原因的限制,防治效果仍很不理想。这使得青枯病的生物防治受到了国内外的高度重视。已有研究报道,生物防治可以有效地降低青枯病的病害程度并减少其造成的产量损失。尽管在中国已有数种防治青枯的生防活菌制剂投入商业化生产,但生防活菌制剂并没有被广大农户接受。导致这种现象出现的最根本原因就是生防制剂的防效不稳定。本论文就青枯病生防菌筛选策略展开研究,旨在建立一个合理、快速的评价体系,通过一系列温室和大田试验对筛选出的生防菌进行有效性和稳定性验证,并借助分子生物学和遗传学手段对生防菌的生防机理进行初步探讨。
1.番茄青枯颉颃菌Acinetobacter sp.Xa6和Enterobacter sp.Xy3的筛选与评价
根据平板上对青枯及其它土传真菌的颉颃试验和温室防效试验结果,我们从14个候选菌株中挑选出两个—Xa6(Acinetobacter sp.,不动杆菌属)和Xy3(Enterobacter sp肠杆菌属)作为番茄青枯病的颉颃细菌。为了寻求到合适的颉颃菌接种方法,我们比较了温室条件下蘸根和土壤浇灌处理后颉颃菌的根围定殖力、防效和促生长作用。结果显示,土壤浇灌处理后颉颃菌的根围定殖力较好、防效和促生长率较高,并且该方法在田间大范围应用时易于操作。我们还发现,无论在温室试验还是大田试验中,菌株Xy3都表现出较高的防效,而菌株Xa6则对番茄表现出较好的促生长作用。在江苏淮安和福建龙岩的大田试验中,生防菌处理后75天时,Xy3的防效都在65%左右,Xa6引起的增产效果则为40.7%和32.4%。本研究也是第一次报道将不动杆菌属细菌作为番茄青枯病的颉颃菌。
2.一种基于根围定殖能力和对代表不同遗传多样性的青枯菌株的颉颃作用的生防菌株筛选策略研究
本实验室前期研究中将采集自中国15个省份14中寄主的319个青枯劳尔氏菌菌株进行了遗传多样性分析。选择了10个代表不同遗传多样性群体的青枯病菌开展以下研究。本研究中测定了8个颉颃细菌对上述青枯菌的平板拮抗活性和温室防治效果。结果显示,颉颃细菌对不同青枯菌株表现出的室内(包括平板和温室)拮抗活性有显著差异。我们从8个候选菌株中选择了沙雷氏菌属细菌菌株XY21,因为其在平板拮抗性测试中对10个参试青枯菌均表现出较好的抑制作用,且在番茄植株根围显示出良好的定殖能力。温室条件下,菌株XY21对由具有遗传多样性的不同青枯菌引起的病害的防效19%至70%不等。我们发现,温室防治效果与平板拮抗活性之间相关性良好(相关系数达到0.746)。在淮安(辣椒地)和福州(番茄地)两地的大田试验中,菌株XY21也有效地控制了青枯病的发生,明显增加了番茄和辣椒的产量。本研究首次报道根据菌株对具有遗传多样性的不同青枯菌的拮抗活性和根围定殖力来筛选植物青枯病的生防菌。
3.颉颃菌株XY21防治植物青枯病的机制初探
在本章中,首先对颉颃菌XY21进行RFP标记,并确定标记后颉颃菌的拮抗性(主要是针对青枯菌的拮抗性)和产酶活性等与生防相关的特性均未发生变化。通过平板计数与激光共聚焦显微镜观察相结合的方法检测其定殖情况。第一次温室试验结果显示,菌株XY21-RFP(?)(?)种处理后定殖于番茄主根根表；灌根处理后,菌株XY21-RFP在植株根表和根围土壤颗粒中均有分布；而且菌株XY21-RFP的荧光表达不稳定；青枯菌株Rs-GFP荧光表达稳定,在萎蔫植株根内会堵塞维管束组织。第二次温室试验,在只接种颉颃菌的处理组中,即使在颉颃菌处理后27天(播种后41天),菌株XY21-rif和XY21-RFP在番茄根部的定殖量仍能达到107CFU/g RFW。变性梯度凝胶电泳(DGGE)用来分析颉颃菌接种后对番茄根围细菌种群的影响,其结果也显示菌株XY21-rif和XY21-RFP在所有取样时间都可以被检测到,这进一步验证了菌株XY21的优良定殖力。此外,对番茄根围细菌种群的DGGE分析结果显示,菌株XY21-rif和XY21-RFP只在后两次取样时(播种后24天和41天)对细菌种群产生了一些影响；在前两次取样时,颉颃菌处理组中除了属于颉颃菌自身的条带外,与不接菌的对照组没有差异。另外,颉颃菌XY21-rif和XY21-RFP的引入对芽孢杆菌属的种群在所有取样时间点都没有影响,对假单胞菌属的影响也只在取样后期。
4.总结
本研究的主要发现与创新：第一、建立了一种基于根围定殖能力和对表现遗传多样性的不同青枯菌株的颉颃作用的生防菌筛选策略；第二、筛选得到了三株对作物青枯病有良好防效的颉颃细菌Xa6、Xy3和XY21,其中不动杆菌属细菌Xa6是首次报道用于青枯病的生物防治；第三、对沙雷氏菌属细菌XY21的生防机理进行了初步探讨,发现其与青枯菌存在位点竞争,并且其引入番茄植株根围后对原本的细菌种群影响不大。
本研究有两个主要缺点：第一、没有研究颉颃菌XY21野生型对番茄根围细菌种群的影响；第二、对菌株XY21的RFP标记不太成功,荧光表达不稳定。
Bacterial wilt of tomato, Lycopersicon esculentum Miller, caused by Ralstonia solanacearum (Yabuuchi et al.), is a destructive soil-borne disease and a serious threat for agricultural production in many provinces of Southern China. So far, Ralstonia wilt control approaches, including field sanitation, crop rotation and application of resistant varieties, have proven limited success. Applying chemical pesticides is generally considered as the most effective and fastest strategy for plant diseases management, however, no effective chemical product has yet been available for Ralstonia wilt. Therefore, more and more efforts need to be devoted to biological control with living microbes. Biological control is one of the most promising approaches to reduce the disease severity and yield loss caused by the diseases. Although several products have been commercialized, living microbial BCAs have not been widely accepted by farmers, since their biocontrol efficiency is not quite stable. This study was focused on development of a novel screen strategy of antagonist against R.solanacearum, evaluation of antagonists identified by the strategy and investigation of mechanism of the antagonistic strain XY21.
1. Evaluation of the strains of Acinetobacter and Enterobacter as potential biocontrol agents against ralstonia wilt of tomato
Based on antagonistic activity against R. solanacearum and three other soil-borne fungal pathogens as well as biocontrol efficiency in greenhouse, we select from a total of14candidates two bacterial strains Xa6(Acinetobacter sp.) and Xy3(Enterobacter sp.) as potential biocontrol agents. To seek a suitable antagonist inoculation method, root-dipping and drenching treatments were compared for bacterial rhizocompetence, biocontrol efficiencies and plant growth promotion potential under greenhouse conditions. Drenching treatment resulted in higher rhizocompetence, biocontrol efficiency and plant growth promotion, and this method was easier to operate in field in large scale. Field trails were conducted for further evaluation of these two antagonists. In both greenhouse and field experiments, strain Xy3had a better control effect against bacterial wilt than Xa6did, while Xa6caused more yield increase. Biocontrol efficiencies of Xy3were both about65%, and the yield increases caused by Xa6were32.4%and40.7%at the75th day after treatment in two field experiments. This is the first report of an Acinetobacter sp. strain used as a BCA agaisnt R. solanacearum of tomato.
2. Rhizocompetence and antagonistic activity towards a range of genetically diverse Ralstonia solanacearum strains-a novel approach for selecting biocontrol agents
Eight antagonistic bacterial strains were tested for their inhibition towards a range of genetically diverse R. solanacearum strains from China and for their efficiency to control Ralstonia wilt of tomato under greenhouse condition. The results indicated that the same antagonists showed markedly different in-vitro antagonistic activity towards different Ralstonia strains, and biocontrol efficiency. From the antagonists tested, Serratia sp. strain XY21was selected because of its good in-vitro inhibition of all ten R. solanacearum strains and its good rhizocompetence on tomato plants. Biocontrol efficacy of XY21towards seven R. solanacearum strains ranged from19%to70%. A good correlation of in-vitro antagonistic activity and the actual biocontrol efficiency towards different R. solanacearum strains was found. Efficient biocontrol and yield increases were observed in field experiments with pepper and tomato. This is the first report to select potential biocontrol strains of R. solanacearum based on their rhizocompetence and their ability to antagonize a range of genetically diverse R. solanacearum isolates.
3. Mechinisms of antagonist XY21against Ralstonia solanacearum strains
In this study, antagonistic strain XY21was labeled with RFP firstly. The antagonistic activity towards genetically diverse Ralstonia solanacearum strains and enzyme activity of strain XY21-RFP were tested and compared to that of wild type strain XY21. No difference was found between wild type strain XY21and mutants XY21-RFP and XY21-rif. The results of first greenhouse experiment showed that strain XY21-RFP colonized surface of primary root after seed inoculation, and after root inoculation, the strain XY21-RFP colonized root surface and soil particles. The expression of florescence of antagonistic strain was unstably, but the expression of florescence of pathogen strain Rs-GFP was stable and the vessel of the wilt plant was full of the pathogen bacteria. The results of second greenhouse experiment displayed that the CFU counts of Serratia sp. XY21-rif and XY21-RFP in the rhizosphere of tomato plants determined by selective plating at all sampling time were around107g-1RFW (root fresh weight). DGGE analysis of16S rRNA gene fragments amplified from total community DNA was used to detect shifts in the relative abundance of ribotypes in the tomato rhizosphere as a consequence of inoculation with antagonists. The comparison of the bacterial DGGE fingerprints revealed that at10and17days after seed inoculation, the patterns of all four replicates of both treatments were highly similar except that strong bands corresponding to the electrophoretic mobility16S rRNA gene products of XY21were detected in the profiles of XY21-mutants-inoculated plants. At24and41days after seed inoculation, several differentiating bands were detected. Some of these bands were only observed in the control or the inoculated plants. Remarkably,27days after the last drenching a dominant band resembling XY21mutants was still detected in three of four replicates suggesting that the inoculant established as dominant rhizosphere population. Highly similar Bacillus profiles were found for both the inoculated and control plants indicating that the inoculation had no effect on the Bacillus community composition, while the inoculation with XY21mutants clearly affected the abundance of some Pseudomonas populations.
1.番茄青枯颉颃菌Acinetobacter sp.Xa6和Enterobacter sp.Xy3的筛选与评价
根据平板上对青枯及其它土传真菌的颉颃试验和温室防效试验结果,我们从14个候选菌株中挑选出两个—Xa6(Acinetobacter sp.,不动杆菌属)和Xy3(Enterobacter sp肠杆菌属)作为番茄青枯病的颉颃细菌。为了寻求到合适的颉颃菌接种方法,我们比较了温室条件下蘸根和土壤浇灌处理后颉颃菌的根围定殖力、防效和促生长作用。结果显示,土壤浇灌处理后颉颃菌的根围定殖力较好、防效和促生长率较高,并且该方法在田间大范围应用时易于操作。我们还发现,无论在温室试验还是大田试验中,菌株Xy3都表现出较高的防效,而菌株Xa6则对番茄表现出较好的促生长作用。在江苏淮安和福建龙岩的大田试验中,生防菌处理后75天时,Xy3的防效都在65%左右,Xa6引起的增产效果则为40.7%和32.4%。本研究也是第一次报道将不动杆菌属细菌作为番茄青枯病的颉颃菌。
2.一种基于根围定殖能力和对代表不同遗传多样性的青枯菌株的颉颃作用的生防菌株筛选策略研究
本实验室前期研究中将采集自中国15个省份14中寄主的319个青枯劳尔氏菌菌株进行了遗传多样性分析。选择了10个代表不同遗传多样性群体的青枯病菌开展以下研究。本研究中测定了8个颉颃细菌对上述青枯菌的平板拮抗活性和温室防治效果。结果显示,颉颃细菌对不同青枯菌株表现出的室内(包括平板和温室)拮抗活性有显著差异。我们从8个候选菌株中选择了沙雷氏菌属细菌菌株XY21,因为其在平板拮抗性测试中对10个参试青枯菌均表现出较好的抑制作用,且在番茄植株根围显示出良好的定殖能力。温室条件下,菌株XY21对由具有遗传多样性的不同青枯菌引起的病害的防效19%至70%不等。我们发现,温室防治效果与平板拮抗活性之间相关性良好(相关系数达到0.746)。在淮安(辣椒地)和福州(番茄地)两地的大田试验中,菌株XY21也有效地控制了青枯病的发生,明显增加了番茄和辣椒的产量。本研究首次报道根据菌株对具有遗传多样性的不同青枯菌的拮抗活性和根围定殖力来筛选植物青枯病的生防菌。
3.颉颃菌株XY21防治植物青枯病的机制初探
在本章中,首先对颉颃菌XY21进行RFP标记,并确定标记后颉颃菌的拮抗性(主要是针对青枯菌的拮抗性)和产酶活性等与生防相关的特性均未发生变化。通过平板计数与激光共聚焦显微镜观察相结合的方法检测其定殖情况。第一次温室试验结果显示,菌株XY21-RFP(?)(?)种处理后定殖于番茄主根根表；灌根处理后,菌株XY21-RFP在植株根表和根围土壤颗粒中均有分布；而且菌株XY21-RFP的荧光表达不稳定；青枯菌株Rs-GFP荧光表达稳定,在萎蔫植株根内会堵塞维管束组织。第二次温室试验,在只接种颉颃菌的处理组中,即使在颉颃菌处理后27天(播种后41天),菌株XY21-rif和XY21-RFP在番茄根部的定殖量仍能达到107CFU/g RFW。变性梯度凝胶电泳(DGGE)用来分析颉颃菌接种后对番茄根围细菌种群的影响,其结果也显示菌株XY21-rif和XY21-RFP在所有取样时间都可以被检测到,这进一步验证了菌株XY21的优良定殖力。此外,对番茄根围细菌种群的DGGE分析结果显示,菌株XY21-rif和XY21-RFP只在后两次取样时(播种后24天和41天)对细菌种群产生了一些影响；在前两次取样时,颉颃菌处理组中除了属于颉颃菌自身的条带外,与不接菌的对照组没有差异。另外,颉颃菌XY21-rif和XY21-RFP的引入对芽孢杆菌属的种群在所有取样时间点都没有影响,对假单胞菌属的影响也只在取样后期。
4.总结
本研究的主要发现与创新：第一、建立了一种基于根围定殖能力和对表现遗传多样性的不同青枯菌株的颉颃作用的生防菌筛选策略；第二、筛选得到了三株对作物青枯病有良好防效的颉颃细菌Xa6、Xy3和XY21,其中不动杆菌属细菌Xa6是首次报道用于青枯病的生物防治；第三、对沙雷氏菌属细菌XY21的生防机理进行了初步探讨,发现其与青枯菌存在位点竞争,并且其引入番茄植株根围后对原本的细菌种群影响不大。
本研究有两个主要缺点：第一、没有研究颉颃菌XY21野生型对番茄根围细菌种群的影响；第二、对菌株XY21的RFP标记不太成功,荧光表达不稳定。
Bacterial wilt of tomato, Lycopersicon esculentum Miller, caused by Ralstonia solanacearum (Yabuuchi et al.), is a destructive soil-borne disease and a serious threat for agricultural production in many provinces of Southern China. So far, Ralstonia wilt control approaches, including field sanitation, crop rotation and application of resistant varieties, have proven limited success. Applying chemical pesticides is generally considered as the most effective and fastest strategy for plant diseases management, however, no effective chemical product has yet been available for Ralstonia wilt. Therefore, more and more efforts need to be devoted to biological control with living microbes. Biological control is one of the most promising approaches to reduce the disease severity and yield loss caused by the diseases. Although several products have been commercialized, living microbial BCAs have not been widely accepted by farmers, since their biocontrol efficiency is not quite stable. This study was focused on development of a novel screen strategy of antagonist against R.solanacearum, evaluation of antagonists identified by the strategy and investigation of mechanism of the antagonistic strain XY21.
1. Evaluation of the strains of Acinetobacter and Enterobacter as potential biocontrol agents against ralstonia wilt of tomato
Based on antagonistic activity against R. solanacearum and three other soil-borne fungal pathogens as well as biocontrol efficiency in greenhouse, we select from a total of14candidates two bacterial strains Xa6(Acinetobacter sp.) and Xy3(Enterobacter sp.) as potential biocontrol agents. To seek a suitable antagonist inoculation method, root-dipping and drenching treatments were compared for bacterial rhizocompetence, biocontrol efficiencies and plant growth promotion potential under greenhouse conditions. Drenching treatment resulted in higher rhizocompetence, biocontrol efficiency and plant growth promotion, and this method was easier to operate in field in large scale. Field trails were conducted for further evaluation of these two antagonists. In both greenhouse and field experiments, strain Xy3had a better control effect against bacterial wilt than Xa6did, while Xa6caused more yield increase. Biocontrol efficiencies of Xy3were both about65%, and the yield increases caused by Xa6were32.4%and40.7%at the75th day after treatment in two field experiments. This is the first report of an Acinetobacter sp. strain used as a BCA agaisnt R. solanacearum of tomato.
2. Rhizocompetence and antagonistic activity towards a range of genetically diverse Ralstonia solanacearum strains-a novel approach for selecting biocontrol agents
Eight antagonistic bacterial strains were tested for their inhibition towards a range of genetically diverse R. solanacearum strains from China and for their efficiency to control Ralstonia wilt of tomato under greenhouse condition. The results indicated that the same antagonists showed markedly different in-vitro antagonistic activity towards different Ralstonia strains, and biocontrol efficiency. From the antagonists tested, Serratia sp. strain XY21was selected because of its good in-vitro inhibition of all ten R. solanacearum strains and its good rhizocompetence on tomato plants. Biocontrol efficacy of XY21towards seven R. solanacearum strains ranged from19%to70%. A good correlation of in-vitro antagonistic activity and the actual biocontrol efficiency towards different R. solanacearum strains was found. Efficient biocontrol and yield increases were observed in field experiments with pepper and tomato. This is the first report to select potential biocontrol strains of R. solanacearum based on their rhizocompetence and their ability to antagonize a range of genetically diverse R. solanacearum isolates.
3. Mechinisms of antagonist XY21against Ralstonia solanacearum strains
In this study, antagonistic strain XY21was labeled with RFP firstly. The antagonistic activity towards genetically diverse Ralstonia solanacearum strains and enzyme activity of strain XY21-RFP were tested and compared to that of wild type strain XY21. No difference was found between wild type strain XY21and mutants XY21-RFP and XY21-rif. The results of first greenhouse experiment showed that strain XY21-RFP colonized surface of primary root after seed inoculation, and after root inoculation, the strain XY21-RFP colonized root surface and soil particles. The expression of florescence of antagonistic strain was unstably, but the expression of florescence of pathogen strain Rs-GFP was stable and the vessel of the wilt plant was full of the pathogen bacteria. The results of second greenhouse experiment displayed that the CFU counts of Serratia sp. XY21-rif and XY21-RFP in the rhizosphere of tomato plants determined by selective plating at all sampling time were around107g-1RFW (root fresh weight). DGGE analysis of16S rRNA gene fragments amplified from total community DNA was used to detect shifts in the relative abundance of ribotypes in the tomato rhizosphere as a consequence of inoculation with antagonists. The comparison of the bacterial DGGE fingerprints revealed that at10and17days after seed inoculation, the patterns of all four replicates of both treatments were highly similar except that strong bands corresponding to the electrophoretic mobility16S rRNA gene products of XY21were detected in the profiles of XY21-mutants-inoculated plants. At24and41days after seed inoculation, several differentiating bands were detected. Some of these bands were only observed in the control or the inoculated plants. Remarkably,27days after the last drenching a dominant band resembling XY21mutants was still detected in three of four replicates suggesting that the inoculant established as dominant rhizosphere population. Highly similar Bacillus profiles were found for both the inoculated and control plants indicating that the inoculation had no effect on the Bacillus community composition, while the inoculation with XY21mutants clearly affected the abundance of some Pseudomonas populations.
引文
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