水稻细菌性条斑病菌遗传多样性研究与生防菌剂研发
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摘要
水稻细菌性条斑病由Xanthomonas oryzae pv. oryzicola (Xooc)引起,是水稻生产中一种重要的检疫性病害,其发生具有流行性、暴发性和毁灭性等特点。该病是我国南方稻区生产的重要病害,当气候条件适宜时,在感病品种上能引起15%～25%产量损失,严重时达40%～60%,对水稻的高产稳产造成严重威胁。品种抗病性是寄主与病原物互作的结果,深入了解病原物群体遗传结构的变化,可作为抗性基因合理分布的依据。主要研究如下：
本文通过rep-PCR指纹技术,分析了来源于江苏、云南、江西、湖南和安徽等省份69个水稻细菌性条斑病菌(X. oryzae pv. oryzicola)群体遗传多样性。用2对特异性引物BOX和ERIC对菌株基因组DNA进行了PCR扩增,以彼此间的带位相似率达80%为界,BOX扩增的谱型被分为10簇,ERIC的谱型被分为8簇。BOX的第3簇包含27个菌株,占总数的39.1%,ERIC的第5簇包含24个菌株,占总数的34.8%,均为优势簇。群体遗传多样性值BOX为0.9784,ERIC为0.9833。说明水稻细菌性条斑病菌群体的遗传分化明显,遗传多样性较高。全部69株菌株接种于含有不同抗病基因近等基因系及高感品种金刚30等6个鉴别品种上,被划分为13个致病型,其中C6致病型占26.1%,为优势致病型。69个菌株的BOX、ERIC的簇群分类与地理位置相关,与致病型不相关。
其次对来源于江苏、云南、江西、湖南和安徽等省细菌性条斑病菌菌株,通过采用苗期注射接种和成株期针刺接种法,研究其在6个鉴别品种(金刚30、IRBB4、 IRBB5、IRBB14、IRBB21和IRBB24)上致病力分化及其对水稻幼苗和成株致病力差异,为品种布局和条斑病菌致病力快速鉴定提供理论依据。试验结果显示,71个条斑病菌菌株在苗期和成株期可分别区分为18个和13个致病型；苗期的优势致病型为C1,占供试菌株总数的17.0%；成株期的优势致病型为C10,占供试菌株总数的25.4%。研究还表明：大多数菌株与水稻品种之间表现为弱互作关系,只有一部分菌株表现为强互作关系。就单个菌株而言,在苗期和成株期的致病力分化是不一致的。整体而言,71个菌株在6个鉴别品种的苗期上可侵染241次,侵染率为56.6%；在成株期上有245次侵染,总侵染率为57.5%。病原菌在不同水稻品种的苗期和成株期上的侵染能力是基本相同的。
同时采集江苏省南京、泰州、扬州和宿迁等地区水稻田水稻病叶、健叶、根围土等样品90份,分离纯化得到1173个细菌分离物,对其进行水稻细菌性条斑病菌(X. oryzae pv. oryzicola)皿内抑制试验,共获得12株拮抗能力较强的细菌,其中4株拮抗细菌抑菌圈直径大于27mm。盆栽试验结果表明,12株拮抗能力较强的细菌中有4株对水稻细菌性条斑病防效大于50%。其中菌株Lx-11盆栽防效达62.5%,大田示范试验防效达60.2%,显著高于化学药剂20%叶枯唑的防效(51.2%和45.8%)。通过形态特征、理化特性和分子鉴定结果确定菌株Lx-11为解淀粉芽孢杆菌Bacillus amyloliquefaciens.
对前期筛选获得和在田间试验中防治水稻细菌性条斑病效果较好的解淀粉芽孢杆菌Lx-11,通过Plackett-Burman单因素筛选及Box-Behnken响应曲面法(response surface methodology, RSM)对影响生防菌株Lx-11生物发酵工艺进行了优化。试验结果表明,培养温度、通气量和接种量是影响发酵活菌数和抑菌活性的主要因子,由所得响应曲面方程预测出这3个主要因子分别为30.84℃、59.68mL/250mL和1.52%时,发酵活菌数和抑菌活性的最大值为3.75×109cm·mL-1和8.2mm。经摇瓶发酵试验和抑菌活性验证该理论预测值与实际值无显著差异。
Lx-11菌株分泌的脂肽类抗生素物质包括surfactin, bacillomycin D和fengycin三大类,通过构建surfactin突变体菌株发现surfactin在防治细菌性条斑病中起主要作用。生防菌处理水稻植株后能够诱导植株防卫反应相关基因PR-la、PR-1b、NPR1和PAL基因的表达,并在处理24小时后达到最大值,表明Lx-11菌株引发植株产生系统免疫反应。
Bacterial leaf streak (BLS) is a quarantine rice disease caused by Xanthomonas oryzae pv. oryzicola (Xooc), which has characteristics of destruction, epidemic and outbreak in southern China. Available reports suggest that yield losses due to this disease typically range from15%to25%depending on the rice variety and climatic conditions. Under conditions favorable for spread, BLS may affect entire fields and cause damage reductions in grain weight of up to40%-60%. Variety resistance results from interaction between host and pathogen. The fundation of resistance gene distribution is to uderstand the varity of population genetic of plant pathogen. The main results were summarized as follows:
Genetic diversity and pathotype variability of X. oryzae pv. oryzicola strains collected from Jiangsu, Yunnan, Jiangxi, Hunan and Anhui were analyzed by using rep-PCR fingerprinting techniques. Genomic DNA from sixty nine strains was amplified with two specific primers BOX and ERIC. Dendrograms were generated from the data by using UPGMA analysis. The strains tested were similar each other at a level of80%,10and8clusters were grouped with BOX and ERIC, respectively. The predominant groups were cluster3and5for each primer BOX and ERIC involving27(39.1%) and24(34.8%) strains, respectively. The genetic diversities of the population of tested strains were0.9784(for BOX) and0.9833(for ERIC), respectively. The results indicated that the genetic diversity of Xanthomonas oryzae pv. oryzicola was various. Sixty nine strains inoculated onto five differential cultivars including near-isogenic rice lines carrying single resistance gene, IRBB4, IRBB5, IRBB14, IRBB21and IRBB24, and a highly susceptible cultivar Jinggang30, may group thirteen pathotypes. The predominant pathotype was C6(26.1%). No relationships were observed between UPGM A groups (BOX and ERIC) of the69strains and pathotypes, but each group was partially associated with the regional origin of strains.
To provide a theoretical basis for rice cultivar distribution and rapid identification of virulence of X. oryzae pv. oryzicola, The virulence differentiation of71isolates of Xanthomonas oryzae pv. oryzicola collected from Hunan, Jiangxi, Yunnan, Jiangsu and Anhui provinces were identified with six rice cultivars (Jingang30, IRBB4, IRBB5, IRBB14, IRBB21, IRBB24) containing different resistance genes against bacterial blight by the inoculation methods of leaf-infiltration with needleless syringes at seedling stage and leaf-needling at adult stage. The results showed that the71strains were classified into18pathotypes at seedling stage and13pathotypes at adult stage. The predominant pathotype was C1(17.0%of the total strains) at seedling stage, and C10(25.4%of the total strains) at adult stage. Most isolates had weak interactions with the rice cultivars, and some the isolates had specific interactions. However, for a single strain, the virulence differentiation of X. oryzae pv. oryzicola was not the same in different growth stages. It also found that all the strains were infected241times (the infection rate was56.6%) and245times (the infection rate was57.5%) at seedling and adult stages, respectively. The virulences of pathogenic bacteria in different rice varieties at seedling and adult stages were basically the same.
In this paper,1173bacterial strains were isolated from the disease leaves infected by X. oryzae pv. oryzicola, the healthy leaves and the rhizosphere of rice plants sampled in Nanjing, Taizhou, Yangzhou and Suqian in Jiangsu province. Among the tested strains,12strains demonstrated high inhibitory activity agaimst X. oryzae pv. oryzicola. The Diameter of inhibition zones of4strains were more than27mm. Control efficiencies of strain Lx-11were62.5%and60.2%in pot and field trials, respectively, significantly higher than those of20%bismerthlazol treatment (51.2%and45.8%). Based on morphological, physiological and biochemical characteristics and16S rDNA sequence analysis, the strain Lx-11was identified to be Bacillus amyloliquefaciens.
The Plackett-Burman experiment and response surface methodology were adopted to optimize the liquid fermentation conditions of the biocontrol agent of Bacillus amyloliquefaciens Lx-11. The results indicated that the temperature, the filling volume of shaking flask and inoculum volume were the major factors that affect the yield of Lx-11. The fermentation conditions were30.84℃, inoculum volume1.52%and the filling volume59.68mL in250mL shaking flask for48h, and initial pH at7.0. Under the optimized fermentation conditions the production of spores was up to3.75×109cfu·mL-1and the width of antibacterial activity was8.2mm. Fermentation experiments with shake flasks verified that there was no statistical difference between the yield value and antibacterial activity forecasted by the model.
We found that Lx-11secreted three kinds of lipopeptides including surfactin, bacillomycin D and fengycin, which exhibted antibacterial activity against X. oryzae pv. oryzicola. The antibacterial activity could be associated with surfactin-lipopeptides, and practically abolished in surfacin-deficient mutants. In addition, the defense-related genes PR-1a, PR-1b, NPR1and PAL were concurrently expressed in the leaves of rice after treated with Lx-11. The results implied that Lx-11triggered a systemic immunization activity.
本文通过rep-PCR指纹技术,分析了来源于江苏、云南、江西、湖南和安徽等省份69个水稻细菌性条斑病菌(X. oryzae pv. oryzicola)群体遗传多样性。用2对特异性引物BOX和ERIC对菌株基因组DNA进行了PCR扩增,以彼此间的带位相似率达80%为界,BOX扩增的谱型被分为10簇,ERIC的谱型被分为8簇。BOX的第3簇包含27个菌株,占总数的39.1%,ERIC的第5簇包含24个菌株,占总数的34.8%,均为优势簇。群体遗传多样性值BOX为0.9784,ERIC为0.9833。说明水稻细菌性条斑病菌群体的遗传分化明显,遗传多样性较高。全部69株菌株接种于含有不同抗病基因近等基因系及高感品种金刚30等6个鉴别品种上,被划分为13个致病型,其中C6致病型占26.1%,为优势致病型。69个菌株的BOX、ERIC的簇群分类与地理位置相关,与致病型不相关。
其次对来源于江苏、云南、江西、湖南和安徽等省细菌性条斑病菌菌株,通过采用苗期注射接种和成株期针刺接种法,研究其在6个鉴别品种(金刚30、IRBB4、 IRBB5、IRBB14、IRBB21和IRBB24)上致病力分化及其对水稻幼苗和成株致病力差异,为品种布局和条斑病菌致病力快速鉴定提供理论依据。试验结果显示,71个条斑病菌菌株在苗期和成株期可分别区分为18个和13个致病型；苗期的优势致病型为C1,占供试菌株总数的17.0%；成株期的优势致病型为C10,占供试菌株总数的25.4%。研究还表明：大多数菌株与水稻品种之间表现为弱互作关系,只有一部分菌株表现为强互作关系。就单个菌株而言,在苗期和成株期的致病力分化是不一致的。整体而言,71个菌株在6个鉴别品种的苗期上可侵染241次,侵染率为56.6%；在成株期上有245次侵染,总侵染率为57.5%。病原菌在不同水稻品种的苗期和成株期上的侵染能力是基本相同的。
同时采集江苏省南京、泰州、扬州和宿迁等地区水稻田水稻病叶、健叶、根围土等样品90份,分离纯化得到1173个细菌分离物,对其进行水稻细菌性条斑病菌(X. oryzae pv. oryzicola)皿内抑制试验,共获得12株拮抗能力较强的细菌,其中4株拮抗细菌抑菌圈直径大于27mm。盆栽试验结果表明,12株拮抗能力较强的细菌中有4株对水稻细菌性条斑病防效大于50%。其中菌株Lx-11盆栽防效达62.5%,大田示范试验防效达60.2%,显著高于化学药剂20%叶枯唑的防效(51.2%和45.8%)。通过形态特征、理化特性和分子鉴定结果确定菌株Lx-11为解淀粉芽孢杆菌Bacillus amyloliquefaciens.
对前期筛选获得和在田间试验中防治水稻细菌性条斑病效果较好的解淀粉芽孢杆菌Lx-11,通过Plackett-Burman单因素筛选及Box-Behnken响应曲面法(response surface methodology, RSM)对影响生防菌株Lx-11生物发酵工艺进行了优化。试验结果表明,培养温度、通气量和接种量是影响发酵活菌数和抑菌活性的主要因子,由所得响应曲面方程预测出这3个主要因子分别为30.84℃、59.68mL/250mL和1.52%时,发酵活菌数和抑菌活性的最大值为3.75×109cm·mL-1和8.2mm。经摇瓶发酵试验和抑菌活性验证该理论预测值与实际值无显著差异。
Lx-11菌株分泌的脂肽类抗生素物质包括surfactin, bacillomycin D和fengycin三大类,通过构建surfactin突变体菌株发现surfactin在防治细菌性条斑病中起主要作用。生防菌处理水稻植株后能够诱导植株防卫反应相关基因PR-la、PR-1b、NPR1和PAL基因的表达,并在处理24小时后达到最大值,表明Lx-11菌株引发植株产生系统免疫反应。
Bacterial leaf streak (BLS) is a quarantine rice disease caused by Xanthomonas oryzae pv. oryzicola (Xooc), which has characteristics of destruction, epidemic and outbreak in southern China. Available reports suggest that yield losses due to this disease typically range from15%to25%depending on the rice variety and climatic conditions. Under conditions favorable for spread, BLS may affect entire fields and cause damage reductions in grain weight of up to40%-60%. Variety resistance results from interaction between host and pathogen. The fundation of resistance gene distribution is to uderstand the varity of population genetic of plant pathogen. The main results were summarized as follows:
Genetic diversity and pathotype variability of X. oryzae pv. oryzicola strains collected from Jiangsu, Yunnan, Jiangxi, Hunan and Anhui were analyzed by using rep-PCR fingerprinting techniques. Genomic DNA from sixty nine strains was amplified with two specific primers BOX and ERIC. Dendrograms were generated from the data by using UPGMA analysis. The strains tested were similar each other at a level of80%,10and8clusters were grouped with BOX and ERIC, respectively. The predominant groups were cluster3and5for each primer BOX and ERIC involving27(39.1%) and24(34.8%) strains, respectively. The genetic diversities of the population of tested strains were0.9784(for BOX) and0.9833(for ERIC), respectively. The results indicated that the genetic diversity of Xanthomonas oryzae pv. oryzicola was various. Sixty nine strains inoculated onto five differential cultivars including near-isogenic rice lines carrying single resistance gene, IRBB4, IRBB5, IRBB14, IRBB21and IRBB24, and a highly susceptible cultivar Jinggang30, may group thirteen pathotypes. The predominant pathotype was C6(26.1%). No relationships were observed between UPGM A groups (BOX and ERIC) of the69strains and pathotypes, but each group was partially associated with the regional origin of strains.
To provide a theoretical basis for rice cultivar distribution and rapid identification of virulence of X. oryzae pv. oryzicola, The virulence differentiation of71isolates of Xanthomonas oryzae pv. oryzicola collected from Hunan, Jiangxi, Yunnan, Jiangsu and Anhui provinces were identified with six rice cultivars (Jingang30, IRBB4, IRBB5, IRBB14, IRBB21, IRBB24) containing different resistance genes against bacterial blight by the inoculation methods of leaf-infiltration with needleless syringes at seedling stage and leaf-needling at adult stage. The results showed that the71strains were classified into18pathotypes at seedling stage and13pathotypes at adult stage. The predominant pathotype was C1(17.0%of the total strains) at seedling stage, and C10(25.4%of the total strains) at adult stage. Most isolates had weak interactions with the rice cultivars, and some the isolates had specific interactions. However, for a single strain, the virulence differentiation of X. oryzae pv. oryzicola was not the same in different growth stages. It also found that all the strains were infected241times (the infection rate was56.6%) and245times (the infection rate was57.5%) at seedling and adult stages, respectively. The virulences of pathogenic bacteria in different rice varieties at seedling and adult stages were basically the same.
In this paper,1173bacterial strains were isolated from the disease leaves infected by X. oryzae pv. oryzicola, the healthy leaves and the rhizosphere of rice plants sampled in Nanjing, Taizhou, Yangzhou and Suqian in Jiangsu province. Among the tested strains,12strains demonstrated high inhibitory activity agaimst X. oryzae pv. oryzicola. The Diameter of inhibition zones of4strains were more than27mm. Control efficiencies of strain Lx-11were62.5%and60.2%in pot and field trials, respectively, significantly higher than those of20%bismerthlazol treatment (51.2%and45.8%). Based on morphological, physiological and biochemical characteristics and16S rDNA sequence analysis, the strain Lx-11was identified to be Bacillus amyloliquefaciens.
The Plackett-Burman experiment and response surface methodology were adopted to optimize the liquid fermentation conditions of the biocontrol agent of Bacillus amyloliquefaciens Lx-11. The results indicated that the temperature, the filling volume of shaking flask and inoculum volume were the major factors that affect the yield of Lx-11. The fermentation conditions were30.84℃, inoculum volume1.52%and the filling volume59.68mL in250mL shaking flask for48h, and initial pH at7.0. Under the optimized fermentation conditions the production of spores was up to3.75×109cfu·mL-1and the width of antibacterial activity was8.2mm. Fermentation experiments with shake flasks verified that there was no statistical difference between the yield value and antibacterial activity forecasted by the model.
We found that Lx-11secreted three kinds of lipopeptides including surfactin, bacillomycin D and fengycin, which exhibted antibacterial activity against X. oryzae pv. oryzicola. The antibacterial activity could be associated with surfactin-lipopeptides, and practically abolished in surfacin-deficient mutants. In addition, the defense-related genes PR-1a, PR-1b, NPR1and PAL were concurrently expressed in the leaves of rice after treated with Lx-11. The results implied that Lx-11triggered a systemic immunization activity.
引文
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