土传烟草青枯病的生物防控及其机理研究
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摘要
烟草青枯病由茄科劳尔氏菌(Ralstonia solanacearum)引起,在我国长江流域及西南烟区普遍发生,危害严重,造成烟叶产量和品质降低甚至绝收。青枯病已经成为我国烟草生产上的主要限制因子之一。本研究主要从寄主-病原-环境三者生态互作角度出发,针对烟草青枯病的发生机理,运用生物防控和生物修复的手段防控青枯病的发生,并从拮抗菌在根系形成的生物保护膜和根系分泌物对病原菌与拮抗菌生长的促进与抑制作用两方面着重阐述了青枯病生物防控机理。
从烟草青枯病发生严重的田间根际土壤及病株中采用茄科劳尔氏菌选择性培养基分离、筛选到青枯病病原菌119株。通过菌落形态、生理生化和16S rRNA基因序列分析,并结合病原菌回接实验,鉴定其中一株为茄科劳尔氏菌烟草专化型致病型(Ralstonia solanacearum E.F.Smith f. sp. nicotianae, Rs)。通过分别改变土壤病原菌数量及温湿度条件研究青枯病的发病条件,结果表明当土壤中Rs的浓度达到107cfug-1soil以上土壤温度达到30℃,湿度维持在75%以上时烟株最易发病。
从健康烟株根际土壤分离、筛选得到两株高效的拮抗菌L-9和L-25,经16S rRNA基因序列和生理生化分析,分别将两株拮抗菌鉴定为Brevibacillus brevis (短短芽孢杆菌)和Streptomyces rochei(娄彻氏链霉菌)。平板拮抗实验结果表明,这两株拮抗菌不仅对茄科劳尔氏菌有较强的拮抗作用,平板拮抗圈分别达到11和151mm,而且对番茄青枯病、茄子青枯病、辣椒青枯病及烟草黑胫病等多种病害病原菌也有较好的平板拮抗作用。
利用BIOLOG表型芯片鉴定系统对拮抗菌的特征性碳、氮源进行分析,比较了两种拮抗菌的平均颜色变化率(AWCD)和三种固体有机废弃物菜粕有机肥、小麦麸、燕麦麸主要成分的利用情况。研究结果表明,L-9和L-25在菜粕有机肥中的9种主要氨基酸的AWCD值都分别大于在小麦麸和燕麦麸主要氨基酸的AWCD值。综合其他的因素,确定菜粕有机肥与牛粪1：1(w:w)的混合物为两个拮抗菌二次发酵的最佳有机载体。
通过拮抗菌发酵液和拮抗青枯病生物有机肥(BOF)的盆栽和大田试验,研究了两株拮抗菌的拮抗效果。结果表明,L-9生物有机肥与L-25生物有机肥1：1(w:w)处理在盆栽试验的各处理中防控效果最好,其防控率高达100%,大田试验的防治效果分别达到了95.4%(安徽试验大田)和30.1%(贵州试验大田)。BOF处理防控效果显著好于拮抗菌发酵液处理；L-9生物有机肥与L-25生物有机肥1：1配比施入土壤,要比单菌生物有机肥处理防控效果更好,发病率和病情指数最低,烟株生物量最大,根际土壤中病原菌数量最少,浓度控制在不易发病的106cfug-1soil以下。
BOF处理根际的细菌和放线菌数量也显著高于病土处理,而真菌和病原菌的数量却显著低于病土处理；BOF处理的烟草的各种抗性酶活性如POD、CAT、PPO、SOD等与病土处理有显著差异,土壤各种酶活性与病土处理也有显著差异；扫描电镜结果显示,发病烟株的维管束导管中有大量粘性物质堵塞于其中,而且维管束严重变形,而健康烟株和施用BOF后未发病烟株的维管束长势正常,未有严重变形；利用梯度变性凝胶电脉(DGGE)和BIOLOG-Ecoplate分析根际土壤的微生物变化,结果表明,施用BOF后微生物群落结构发生了改变,逐渐由“真菌型”向“细菌型”过渡；BOF处理的Shannon指数、Simpson指数和Mclntosh指数均高于病土处理；研究结果同时表明,施用BOF不但可以抵制病害的发生,对烟叶产量也有较好的促进作用,盆栽试验与田间试验BOF处理与病土对照相比分别增产3.43和3.27倍。
拮抗菌的根际定殖试验结果表明,烟草种子用拮抗菌发酵液处理后,拮抗菌会沿根系生长并附着在根和根毛上；而接种病原菌后,病原菌无法侵染根系,被阻挡在拮抗菌形成的生物保护壳(膜)外。荧光标记的病原菌跟踪结果表明,拮抗菌优先定殖于烟草根表利于抑制病原菌的定殖。通过高效液相色谱法(HPLC),分离并鉴定了烟草根系分泌物中的酚酸类物质。研究结果发现,烟草根系分泌物中检测到0.25μgg-1root DW根干重的苯甲酸和1.15μgg-1root DW的苯丙酸。外源添加苯甲酸和苯丙酸后,两种物质都是低浓度促进病原菌与拮抗菌的生长,高浓度抑制生长,而且两种酚酸类物质对病原菌促生浓度(苯甲酸为300mg L-1,苯丙酸为600mg L-1)大于对两株拮抗菌(L-9和L-25)的促生浓度,说明烟草土传病原菌更能有效地利用根系分泌物。
综上所述,高效拮抗烟草青枯病生防菌Brevibacillus brevis和Streptomyces rochei,在最佳发酵底物-菜粕堆肥与牛粪有机肥(1：1)二次发酵,其混合生物有机肥防控效果最佳且拮抗菌优于青枯菌在烟株根际定殖。BOF通过改善土壤微生物群落结构、增强烟株自身抗性,从而生物防控青枯病、增加烟叶产量。烟草根系分泌物对青枯菌的促进作用大于对拮抗菌的促进作用,这是烟草产生连作障碍的主要原因之一。
Tobacco bacterial wilt, which was caused by Ralstonia solanacearum, spread widely and caused very severe damages in Changjiang River and Southwest tobacco field in China. The wilt disease can decrease tobacco product and quality or even cause total destruction. In a word, tobacco bacterial wilt has been one of the mort crucial limiting factors in Chinese tobacco production. From the viewpoint of host-pathogen-environment ecological interaction triangle, this study investigated the biological control of tobacco bacterial wilt, bioremediation of soils, the bio-film protection from antagonists to R. solanacearum and the effet on the growth of pathogen and antagonists of tobacco root exudates on the basis of pathogenesis of tobacco bacterial wilt.
Rhizosphere soil or infested tobacco plants were collected from a severely wilt-diseased field. One hundred and nineteen colonies similar to R. solanacearum was screened on a semi-selective medium (SMSA) from roots and stem spots of infested plants and rhizosphere soils. A strain of R. solanacearum with strong virulence was identified by its colony and microscopic morphology,16S rRNA sequence homologies and its physo-biochemical characteristics. The isolated strain was identified as Ralstonia solanacearum E.F.Smith f. sp. Nicotianae (Rs) by re-inoculating pathogen to healthy tobacco plant and other four kinds of solanaceous plants. The results of pathogenic factors for R. solanacearum in soil showed that tobacco plants were more susceptible to wilt disease when the population of R. solanacearum reached to10cfu g-1soil, the environmental temperature was30℃and the humidity kept75%above.
In lab condition, two antagonists, L-9and L-25, were isolated and screened from uninfected tobacco plants and rhizosphere soils. Through16S rRNA sequence homologies and physo-biochemical analysis, one strain was Brevibacillus brevis, the other was Streptomyces rochei. These two strains not only had antagonictic effect on R. solanacearum with antagonism zone11mm and15mm, but also suppressed other kinds of soil borne disease on plates in out lab, such as tomato bacterial wilt, eggplant bacterial wilt, pepper bacterial wilt and tobacco black shank.
The carbon and nitrogen sources used by antagonists were analysed by BIOLOG phenotype microarray microplates. In microplates, the average well color development (A WCD) of amino acids, which were mainly contained in rapeseed meal, wheat barn and oat bran, was the highest in rapeseed meal among these three solid wastes. These results indicated that the rapeseed meal was the best substance for reproducetion of antagonists. Considering the comprehensive factors for tobacco plants, an organic fertilizer consisted of rapeseed meal and dairy manure at the proportion of1:1(w:w) was the best compost for the secondary solid fermentation of antagonists.
Biological experiment was conducted using antagonistis cell suspention or bioorganic fertilizer (BOF) to proof bio-control efficacy of antagonists. The results showed that the mixed BOF treatment had the best biocontrol effect in all antagonists applied treatments in pot experiment. The control efficacy of mixed BOF treatment was up to100%in pot experiment and that was95.4%in Anhui field experiemt and30.1%in Guizhou field experiment. The effect of BOF applied treatments was better than antagonists-cell-suspension applied treatments. Besides, the result of treatment where the two BOF second solid fermentated were applied to the soil together at the proportion of1:1, was better than those of sigle BOF applied treatments. The disease incidence and index were the lowest in mixed BOF treatment among all treatments. Moreover, in mixed BOF treatment, tobacco biomass was the largest while R. solancearum population was the least and kept106cfu g-1soil.
The counts of bacteria and actinomycetes were significantly higher while populations of fungi and R. solanacearum were much lower in BOF applied treatment than that of control treatment (infected soil). Furthermore, defence enzymes in leaves and roots, such as POD, CAT, PPO, SOD and so on, and the enzymes in soil were significantly different between the BOF applied treatments and the control treatment (infected soil). Observations of vascular bundle of healthy tobacco plant, wilt plant in control treatment (infected soil) and uninfected plant in antagonists applied treatment were made by scanning electron microscope (SEM). The SEM results showed that there was viscous material deforming and blocking vessel of vascular bundle in wilt tobacco while the vascular bundle of healthy tobacco in control and BOF treatments grew well with a normal shape. Denaturing Gradient Gel Electrophoresis (DGGE) and BIOLOG-Ecoplate were used to analyze the microbial changees in rhizosphere soil in pot experiment. Soil was transformed from fungal type to bacterial type, which indicated that continuous cropping soil was gradually recovered to a new microbial system, sustainable soil ecology and good productive functions. The Shannon index, Simpson index and McIntosh index were higher in BOF treatment than that of control treatment (infected soil). The application of BOF could not only suppress pathogen, but also have a promotion to tobacco yield. Biomass was increased3.43times and3.27times in pot and field experiment, respectivily.
Rhizosphere colonization of antagonists experiment showed that, antagonistic strain adhered on root and root hairs along with the root growth due to the exudates of roots after cell suspensions of antagonists were inoculated to tobacco seeds before its germination. After R. solanacearum was inoculated, antagonists could colonize in rhizosphere firstly, forming a protecting biofilm to suppress R. solanacearum and prevent pathogens from infecting roots. Plasmid pJZ383with green fluorescent protein (gfp) was successfully transformed to R. solanacearum. Antagonists could colonize on the root surface prior to pathogen, thus suppressing R. solanacearum in the rhizosphere. Phenolic acids in tobacco root exudates (TRE) were isolated and identified by high performance liquid chromatography (HPLC). Benzoic acid with the concentration of0.25μg g-1dry roots and3-phenylpropanoic acid with the concentration of1.15μg g-1dry roots were identified in TRE. Exogenous phenolic acids were used to investigate the effect of some phenolic acids, which existed in TRE, on the growth of pathogen and antagonists. It revealed that lower concentration of benzoic acid and3-phenylpropanoic acid promoted the growth of pathogen and antagonists, while higher concentration of those phenolic acids inhibited pathogen and antagonists. Besides, the promotion concentration of these phenolic acids for R. solanacearum (benzoic acid:300mg L-1and3-phenylpropanoic acid:600mg L-1) were higher than that for antagonists. This suggested that R. solanacearum made a better use of tobacco root exudates.
Above all, two antagonistic microbes, Brevibacillus brevis and Streptomyces rochei were screened. They were applied to aan orgnic fertilizer consisted of rapeseed meal and dairy manure at the proportion of1:1(w:w) for the secondary solid fermentation to make a BOF. The mixed BOF had the best control efficacy and the antagonists could colonize on root surface prior to pathogen. BOF could biocontrol tobacco bacterial wilt and increase yield through improve soil microbial diversities and strengthened tobacco self-resistance. Tobacco root exudates had a larger role in growth promoting of R. solanacearum than antagonists. This was one of the most important reseaons for tobacco obstacle of continuous mono-cropping.
从烟草青枯病发生严重的田间根际土壤及病株中采用茄科劳尔氏菌选择性培养基分离、筛选到青枯病病原菌119株。通过菌落形态、生理生化和16S rRNA基因序列分析,并结合病原菌回接实验,鉴定其中一株为茄科劳尔氏菌烟草专化型致病型(Ralstonia solanacearum E.F.Smith f. sp. nicotianae, Rs)。通过分别改变土壤病原菌数量及温湿度条件研究青枯病的发病条件,结果表明当土壤中Rs的浓度达到107cfug-1soil以上土壤温度达到30℃,湿度维持在75%以上时烟株最易发病。
从健康烟株根际土壤分离、筛选得到两株高效的拮抗菌L-9和L-25,经16S rRNA基因序列和生理生化分析,分别将两株拮抗菌鉴定为Brevibacillus brevis (短短芽孢杆菌)和Streptomyces rochei(娄彻氏链霉菌)。平板拮抗实验结果表明,这两株拮抗菌不仅对茄科劳尔氏菌有较强的拮抗作用,平板拮抗圈分别达到11和151mm,而且对番茄青枯病、茄子青枯病、辣椒青枯病及烟草黑胫病等多种病害病原菌也有较好的平板拮抗作用。
利用BIOLOG表型芯片鉴定系统对拮抗菌的特征性碳、氮源进行分析,比较了两种拮抗菌的平均颜色变化率(AWCD)和三种固体有机废弃物菜粕有机肥、小麦麸、燕麦麸主要成分的利用情况。研究结果表明,L-9和L-25在菜粕有机肥中的9种主要氨基酸的AWCD值都分别大于在小麦麸和燕麦麸主要氨基酸的AWCD值。综合其他的因素,确定菜粕有机肥与牛粪1：1(w:w)的混合物为两个拮抗菌二次发酵的最佳有机载体。
通过拮抗菌发酵液和拮抗青枯病生物有机肥(BOF)的盆栽和大田试验,研究了两株拮抗菌的拮抗效果。结果表明,L-9生物有机肥与L-25生物有机肥1：1(w:w)处理在盆栽试验的各处理中防控效果最好,其防控率高达100%,大田试验的防治效果分别达到了95.4%(安徽试验大田)和30.1%(贵州试验大田)。BOF处理防控效果显著好于拮抗菌发酵液处理；L-9生物有机肥与L-25生物有机肥1：1配比施入土壤,要比单菌生物有机肥处理防控效果更好,发病率和病情指数最低,烟株生物量最大,根际土壤中病原菌数量最少,浓度控制在不易发病的106cfug-1soil以下。
BOF处理根际的细菌和放线菌数量也显著高于病土处理,而真菌和病原菌的数量却显著低于病土处理；BOF处理的烟草的各种抗性酶活性如POD、CAT、PPO、SOD等与病土处理有显著差异,土壤各种酶活性与病土处理也有显著差异；扫描电镜结果显示,发病烟株的维管束导管中有大量粘性物质堵塞于其中,而且维管束严重变形,而健康烟株和施用BOF后未发病烟株的维管束长势正常,未有严重变形；利用梯度变性凝胶电脉(DGGE)和BIOLOG-Ecoplate分析根际土壤的微生物变化,结果表明,施用BOF后微生物群落结构发生了改变,逐渐由“真菌型”向“细菌型”过渡；BOF处理的Shannon指数、Simpson指数和Mclntosh指数均高于病土处理；研究结果同时表明,施用BOF不但可以抵制病害的发生,对烟叶产量也有较好的促进作用,盆栽试验与田间试验BOF处理与病土对照相比分别增产3.43和3.27倍。
拮抗菌的根际定殖试验结果表明,烟草种子用拮抗菌发酵液处理后,拮抗菌会沿根系生长并附着在根和根毛上；而接种病原菌后,病原菌无法侵染根系,被阻挡在拮抗菌形成的生物保护壳(膜)外。荧光标记的病原菌跟踪结果表明,拮抗菌优先定殖于烟草根表利于抑制病原菌的定殖。通过高效液相色谱法(HPLC),分离并鉴定了烟草根系分泌物中的酚酸类物质。研究结果发现,烟草根系分泌物中检测到0.25μgg-1root DW根干重的苯甲酸和1.15μgg-1root DW的苯丙酸。外源添加苯甲酸和苯丙酸后,两种物质都是低浓度促进病原菌与拮抗菌的生长,高浓度抑制生长,而且两种酚酸类物质对病原菌促生浓度(苯甲酸为300mg L-1,苯丙酸为600mg L-1)大于对两株拮抗菌(L-9和L-25)的促生浓度,说明烟草土传病原菌更能有效地利用根系分泌物。
综上所述,高效拮抗烟草青枯病生防菌Brevibacillus brevis和Streptomyces rochei,在最佳发酵底物-菜粕堆肥与牛粪有机肥(1：1)二次发酵,其混合生物有机肥防控效果最佳且拮抗菌优于青枯菌在烟株根际定殖。BOF通过改善土壤微生物群落结构、增强烟株自身抗性,从而生物防控青枯病、增加烟叶产量。烟草根系分泌物对青枯菌的促进作用大于对拮抗菌的促进作用,这是烟草产生连作障碍的主要原因之一。
Tobacco bacterial wilt, which was caused by Ralstonia solanacearum, spread widely and caused very severe damages in Changjiang River and Southwest tobacco field in China. The wilt disease can decrease tobacco product and quality or even cause total destruction. In a word, tobacco bacterial wilt has been one of the mort crucial limiting factors in Chinese tobacco production. From the viewpoint of host-pathogen-environment ecological interaction triangle, this study investigated the biological control of tobacco bacterial wilt, bioremediation of soils, the bio-film protection from antagonists to R. solanacearum and the effet on the growth of pathogen and antagonists of tobacco root exudates on the basis of pathogenesis of tobacco bacterial wilt.
Rhizosphere soil or infested tobacco plants were collected from a severely wilt-diseased field. One hundred and nineteen colonies similar to R. solanacearum was screened on a semi-selective medium (SMSA) from roots and stem spots of infested plants and rhizosphere soils. A strain of R. solanacearum with strong virulence was identified by its colony and microscopic morphology,16S rRNA sequence homologies and its physo-biochemical characteristics. The isolated strain was identified as Ralstonia solanacearum E.F.Smith f. sp. Nicotianae (Rs) by re-inoculating pathogen to healthy tobacco plant and other four kinds of solanaceous plants. The results of pathogenic factors for R. solanacearum in soil showed that tobacco plants were more susceptible to wilt disease when the population of R. solanacearum reached to10cfu g-1soil, the environmental temperature was30℃and the humidity kept75%above.
In lab condition, two antagonists, L-9and L-25, were isolated and screened from uninfected tobacco plants and rhizosphere soils. Through16S rRNA sequence homologies and physo-biochemical analysis, one strain was Brevibacillus brevis, the other was Streptomyces rochei. These two strains not only had antagonictic effect on R. solanacearum with antagonism zone11mm and15mm, but also suppressed other kinds of soil borne disease on plates in out lab, such as tomato bacterial wilt, eggplant bacterial wilt, pepper bacterial wilt and tobacco black shank.
The carbon and nitrogen sources used by antagonists were analysed by BIOLOG phenotype microarray microplates. In microplates, the average well color development (A WCD) of amino acids, which were mainly contained in rapeseed meal, wheat barn and oat bran, was the highest in rapeseed meal among these three solid wastes. These results indicated that the rapeseed meal was the best substance for reproducetion of antagonists. Considering the comprehensive factors for tobacco plants, an organic fertilizer consisted of rapeseed meal and dairy manure at the proportion of1:1(w:w) was the best compost for the secondary solid fermentation of antagonists.
Biological experiment was conducted using antagonistis cell suspention or bioorganic fertilizer (BOF) to proof bio-control efficacy of antagonists. The results showed that the mixed BOF treatment had the best biocontrol effect in all antagonists applied treatments in pot experiment. The control efficacy of mixed BOF treatment was up to100%in pot experiment and that was95.4%in Anhui field experiemt and30.1%in Guizhou field experiment. The effect of BOF applied treatments was better than antagonists-cell-suspension applied treatments. Besides, the result of treatment where the two BOF second solid fermentated were applied to the soil together at the proportion of1:1, was better than those of sigle BOF applied treatments. The disease incidence and index were the lowest in mixed BOF treatment among all treatments. Moreover, in mixed BOF treatment, tobacco biomass was the largest while R. solancearum population was the least and kept106cfu g-1soil.
The counts of bacteria and actinomycetes were significantly higher while populations of fungi and R. solanacearum were much lower in BOF applied treatment than that of control treatment (infected soil). Furthermore, defence enzymes in leaves and roots, such as POD, CAT, PPO, SOD and so on, and the enzymes in soil were significantly different between the BOF applied treatments and the control treatment (infected soil). Observations of vascular bundle of healthy tobacco plant, wilt plant in control treatment (infected soil) and uninfected plant in antagonists applied treatment were made by scanning electron microscope (SEM). The SEM results showed that there was viscous material deforming and blocking vessel of vascular bundle in wilt tobacco while the vascular bundle of healthy tobacco in control and BOF treatments grew well with a normal shape. Denaturing Gradient Gel Electrophoresis (DGGE) and BIOLOG-Ecoplate were used to analyze the microbial changees in rhizosphere soil in pot experiment. Soil was transformed from fungal type to bacterial type, which indicated that continuous cropping soil was gradually recovered to a new microbial system, sustainable soil ecology and good productive functions. The Shannon index, Simpson index and McIntosh index were higher in BOF treatment than that of control treatment (infected soil). The application of BOF could not only suppress pathogen, but also have a promotion to tobacco yield. Biomass was increased3.43times and3.27times in pot and field experiment, respectivily.
Rhizosphere colonization of antagonists experiment showed that, antagonistic strain adhered on root and root hairs along with the root growth due to the exudates of roots after cell suspensions of antagonists were inoculated to tobacco seeds before its germination. After R. solanacearum was inoculated, antagonists could colonize in rhizosphere firstly, forming a protecting biofilm to suppress R. solanacearum and prevent pathogens from infecting roots. Plasmid pJZ383with green fluorescent protein (gfp) was successfully transformed to R. solanacearum. Antagonists could colonize on the root surface prior to pathogen, thus suppressing R. solanacearum in the rhizosphere. Phenolic acids in tobacco root exudates (TRE) were isolated and identified by high performance liquid chromatography (HPLC). Benzoic acid with the concentration of0.25μg g-1dry roots and3-phenylpropanoic acid with the concentration of1.15μg g-1dry roots were identified in TRE. Exogenous phenolic acids were used to investigate the effect of some phenolic acids, which existed in TRE, on the growth of pathogen and antagonists. It revealed that lower concentration of benzoic acid and3-phenylpropanoic acid promoted the growth of pathogen and antagonists, while higher concentration of those phenolic acids inhibited pathogen and antagonists. Besides, the promotion concentration of these phenolic acids for R. solanacearum (benzoic acid:300mg L-1and3-phenylpropanoic acid:600mg L-1) were higher than that for antagonists. This suggested that R. solanacearum made a better use of tobacco root exudates.
Above all, two antagonistic microbes, Brevibacillus brevis and Streptomyces rochei were screened. They were applied to aan orgnic fertilizer consisted of rapeseed meal and dairy manure at the proportion of1:1(w:w) for the secondary solid fermentation to make a BOF. The mixed BOF had the best control efficacy and the antagonists could colonize on root surface prior to pathogen. BOF could biocontrol tobacco bacterial wilt and increase yield through improve soil microbial diversities and strengthened tobacco self-resistance. Tobacco root exudates had a larger role in growth promoting of R. solanacearum than antagonists. This was one of the most important reseaons for tobacco obstacle of continuous mono-cropping.
引文
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