生物有机肥对土传马铃薯青枯病的防控技术及机理研究
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摘要
马铃薯青枯病是由茄科劳尔氏菌(Ralstonia solanacearum)引起的一种毁灭性的细菌性土传病害,是世界范围内马铃薯上的第二大病害,其重要性仅次于晚疫病。目前马铃薯青枯病的防控方法已成为马铃薯生产的研究热点,以生物防控为主导的综合防控是防控该病害的有效方法。本研究从黑龙江、南京、山东和海南等地采集健康植株根际土壤,从中分离出对青枯菌有拮抗作用的根际细菌,并通过进一步室内试验和温室试验筛选出两株能显著抑制马铃薯青枯菌的根际细菌解淀粉芽孢杆菌LH23和枯草芽孢杆菌LH36。另外通过一系列的室内分析、温室盆栽和田间试验,研究了拮抗菌LH23和LH36及其制成的生物有机肥(BI023和BI036)对土传马铃薯青枯病的生防效果及其机理,主要结果如下：
1、从番茄、茄子、马铃薯、烟草和香蕉等作物根际土壤中通过喷雾法初筛获得对马铃薯青枯菌有抑菌作用的140多株根际菌株,分离纯化后多次筛选得到两株生长速度快、抗生素抗性强、产芽孢、抗菌谱广泛的高效拮抗菌株LH23和LH36
2、根据16S rRNA基因序列分析和生理生化特性,菌株LH23被鉴定为解淀粉芽孢杆菌(Bacillus amyloliquefaciens),菌株LH36被鉴定为枯草芽孢杆菌(Bacillus subtilis)。同时,研究了拮抗菌LH23和LH36生长的最适发酵条件以及不同生长条件对生防菌LH23和LH36产的拮抗物质抑菌活性的影响。进一步的实验室结果表明：生防菌LH23和LH36具有产吲哚乙酸(IAA)、铁载体、蛋白酶、β-1,3-葡聚糖酶、蛋白类拮抗物质和脂肽类拮抗物质的能力,同时还具有解磷能力。生防菌LH23和LH36的无菌发酵液加入青枯菌培养基,测得的胞外多糖含量减少。
3、灌根法接种与蘸根法接种相比,灌根法接种拮抗菌能更好的在土壤中定殖,促进马铃薯植株生长。温室和田间试验结果表明,生物有机肥(BI023和BI036)能显著促进马铃薯生长和提高马铃薯块茎产量。温室盆栽试验结果表明,生物有机肥(BI023和BI036)处理,马铃薯植株株高、干重、鲜重、根冠比、根总长、根总表面积、根总体积和根平均直径显著高于CK和OF处理(OF即氨基酸肥料和猪粪堆肥1：1混匀制成的普通有机肥)；生物有机肥(BI023和BI036)促进马铃薯植株根际土壤细菌和放线菌数量增加,真菌数量减少。田间试验结果表明,生物有机肥BI023和B1036处理与OF处理相比,产量分别增加17.7%和10.8%,其差异达显著水平。
4、不同土壤验证防控效果试验结果显示：生物有机肥能有效防控马铃薯青枯病的发生,促进马铃薯植株生长,提高马铃薯产量。连作带病土温室盆栽试验中,CK发病率为39.7%,BI023和BI036处理对马铃薯青枯病的防病率皆为100%；健康土添加青枯菌温室盆栽试验中,cK发病率为57.7%,BI023和BI036处理对马铃薯青枯病的防病率分别为84.6%和80.8%。在两块青枯病感染程度不同的两块马铃薯田块(田块1感染程度较轻,青枯病发病率为8.3%；田块2感染程度较重,青枯病发病率为18.3%),进行马铃薯青枯病生防试验,结果表明：在田块1中,BI023和BI036处理的防病率分别为92.0%和84.0%,与OF处理相比产量分别提高42.3%和28.8%；在田块2中,BI023和BI036处理的防病率分别为65.6%和57.7%,与OF处理相比产量分别提高56.8%和45.1%。生物有机肥能有效防控马铃薯青枯病,并显著提高马铃薯产量。
5、施用生物有机肥可以改善马铃薯植株根际土壤微生物区系。不同土壤盆栽试验结果表明,对不同时期马铃薯植株根际土壤微生物计数结果显示,与CK相比,施用生物有机肥(BI023和BI036)显著促进马铃薯植株根际土壤细菌、放线菌的数量增加,真菌的数量减少。同时,与单一施用生防菌菌悬液处理(LH23和LH36)相比,生物有机肥(BI023和BI036)处理马铃薯植株根际土壤拮抗菌数量显著增加,马铃薯植株根际土壤青枯菌数量显著降低,防病率显著提高。田间试验中的根际土壤微生物区系变化与盆栽结果一致。同时生物有机肥显著提高马铃薯植株叶片过氧化物酶(POD)、过氧化氢酶(CAT)、超氧化物歧化酶(SOD)和多酚氧化酶(PPO)活性,降低丙二醛(MDA)含量,减轻发病情况,提高马铃薯植物的系统抗性。
综上所述,拮抗菌株与有机肥结合制成的生物有机肥(BI023和BI036)能有效防控马铃薯青枯病的发生,并促进马铃薯植株的生长、提高产量和改善品质。施用生物有机肥能促进拮抗菌在土壤和根际中的定殖和生存,有效降低青枯菌的数量,同时改善马铃薯植株根际微生物区系,提高植株系统抗性。本文的研究结果有助于研发马铃薯土传病害防控技术,为马铃薯青枯病的有效防控提供理论基础。
Potato bacterial wilt(Ralstonia solanacearum) is a soil-borne disease that seriously suppressed the potato plant(Solanum tuberosum) growth worldwide and causes serious economic losses. A lot of methods have been used to prevent the disease, among which biocontrol has been proved to be the most promising one. Biological control is one of the important measures for overcoming soil-borne diseases, because of its effectiveness and environment-friendly use. Screening antagonists against pathogens is the vital work for the biocontrol of plant diseases.
In this study, two bacterial strains LH23(Bacillus amyloliquefaciens) and LH36(Bacillus subtilis) with antagonistic activity against R. solanacearum, were selected from the rhizosphere soil of healthy potato plant grown in the field strongly diseased by bacterial wilt. Pot and field experiments were conducted to evaluate the LH23(Bacillus amyloliquefaciens) and LH36(Bacillus subtilis) strains and their derived bio-organic fertilizers (BIO23and BIO36) as potential biocontrol agents against potato bacterial wilt and the related mechanisms were also investigated. The main results obtained are listed as follows:
(1) In this study, more than140bacterial strains (bacterial antagonists against R.solanacearum) were isolated from rhizosphere soil of potato, tomato, eggplant and banana. Cell-free filtrate of six antagonistic strains had much stronger antagonistic activity against four R. solanacearum strains with inhibitory zones over20mm. Based on antagonistic activity against four R. solanacearum strains and ten soil-borne fungal pathogens as well as biocontrol efficacy in the greenhouse, two bacterial strains LH23and LH36were selected out of many candidates as potential biocontrol agents.
(2) Based on the results of morphologic characteristics, physiological and biochemical properties and phylogenetic analysis of16S rRNA, the strains LH23and LH36, were identified as Bacillus amyloliquefaciens and Bacillus subtilis, respectively. Measurement from laboratory tests showed that the LH23and LH36were able to produce growth-promoting substances indole acetic acid (IAA), siderophore, prolease, amylase, glucanase, antagonism of extracellular protein and dissolve P in medium. And the LH23and LH36could decrease the content of extracellular Polysaeeharides produced by R. solanacearum, suggesting that LH23and LH36were effective antagonists against the pathogen of potato bacterial wilt.
(3) In order to find a suitable antagonist inoculation method, we compared the methods of root-dipping with soil-drenching in the aspects including rhizocompetence, and effect of promoting plant growth under greenhouse conditions. The drenching treatment resulted in a higher rhizocompetence and plant-yield increase, and this method was also easier to operate in the field on a large scale. Pot and field experiments were performed to investigate the promoting effects of LH23(Bacillus amyloliquefaciens) and LH36(Bacillus subtilis) strains and their derived bio-organic fertilizers (BIO23and BIO36) in potato growth. In greenhouse experiment, BIO23and BIO36significantly promoted potato plant growth, especially the root growth. The root fresh weight of BIO23and BIO36treatments were1.8and2.1times higher than those of control, respectively. The root lengths, root SurfArea, root AvgDiam and RootVolume of the plants treated with BIO23and BIO36were significantly higher than those of control. The potato yield increments on85days after the application of BIO23and BIO36were17.7%and10.8%, respectively as compared with OF (organic fertilizer) treatment in the field experiments.
(4) Application of BIO23and BIO36decreased the incidence of bacterial wilt disease and increased potato yields. In one greenhouse experiment, the biocontrol efficiency of BIO23and BIO36applied were100%and100%, which were the mostly successful treatment while the application of the strains LH23and LH36alone resulted in a biocontrol efficiency of85.1%and81.1%, respectively. In another greenhouse experiment, the biocontrol efficiency of BIO23was84.6%, which was the most successful treatment and BIO36was the second in terms of the biocontrol efficiency (80.8%); the increased percentages of potato yields when compared with the control (chemical fertilizer) were63.5%(BIO23),64.7%(BIO36)34.8%(LH23),33.6%(LH36) and20.7%(OF). In field experiments, the biocontrol efficacies of BIO23and BIO36treatments were92.0%and84.0%, and the yield increases of BIO23and BIO36treatments were42.3%and28.8%, respectively, when compared with the OF treatment.
(5) Bio-organic fertilizers (BIO23and BIO36) could alter the microbial community structure and improved plant health. The counts of bacteria and actinomyces in the rhizosphere soils were significantly increased in BIO23and BIO36treatments, whereas the fungi in the soil in the both treatments were decreased; the counts of antagonists, bacteria and actinobacteria in the rhizosphere soil were significantly increased in BIO23and BIO36treatments, whereas the counts of R. solanacearum and fungi in the soil in the both treatments were decreased in the greenhouse experiments. In addition, the changes in the microbial populations in field experiments were the same as those observed in the greenhouse experiment. In the pot experiment, Bio-organic fertilizers (BIO23and BIO36) significantly increased the activities of plant defence enzymes such as POD, CAT, SOD and PPO, in plant leaves, and decreased MDA content, and consequently relieved the symptoms of potato.
In conclusion, our studies showed that the bio-organic fertilizer, an appropriate combination of organic fertilizer (OF) with the antagonists LH23or LH36used in the present study, could effectively suppress the potato wilt, promote the growth of potato plants and thus the quality of potato tubers, which mainly depends on producing many antibiotics and growth-promoting substances and the alternation of soil microbial community, after the successful colonization of the strains in soils. Results in this study would be useful in the developping new methods to better control potato bacterial wilt disease. Further researches should be done in the molecular microbilogical aspects of the functions performed by bio-organic fertilizers in remediation of continuously cropping obstacles and in the promotion of plant growth and health exerted by functional microbes added.
1、从番茄、茄子、马铃薯、烟草和香蕉等作物根际土壤中通过喷雾法初筛获得对马铃薯青枯菌有抑菌作用的140多株根际菌株,分离纯化后多次筛选得到两株生长速度快、抗生素抗性强、产芽孢、抗菌谱广泛的高效拮抗菌株LH23和LH36
2、根据16S rRNA基因序列分析和生理生化特性,菌株LH23被鉴定为解淀粉芽孢杆菌(Bacillus amyloliquefaciens),菌株LH36被鉴定为枯草芽孢杆菌(Bacillus subtilis)。同时,研究了拮抗菌LH23和LH36生长的最适发酵条件以及不同生长条件对生防菌LH23和LH36产的拮抗物质抑菌活性的影响。进一步的实验室结果表明：生防菌LH23和LH36具有产吲哚乙酸(IAA)、铁载体、蛋白酶、β-1,3-葡聚糖酶、蛋白类拮抗物质和脂肽类拮抗物质的能力,同时还具有解磷能力。生防菌LH23和LH36的无菌发酵液加入青枯菌培养基,测得的胞外多糖含量减少。
3、灌根法接种与蘸根法接种相比,灌根法接种拮抗菌能更好的在土壤中定殖,促进马铃薯植株生长。温室和田间试验结果表明,生物有机肥(BI023和BI036)能显著促进马铃薯生长和提高马铃薯块茎产量。温室盆栽试验结果表明,生物有机肥(BI023和BI036)处理,马铃薯植株株高、干重、鲜重、根冠比、根总长、根总表面积、根总体积和根平均直径显著高于CK和OF处理(OF即氨基酸肥料和猪粪堆肥1：1混匀制成的普通有机肥)；生物有机肥(BI023和BI036)促进马铃薯植株根际土壤细菌和放线菌数量增加,真菌数量减少。田间试验结果表明,生物有机肥BI023和B1036处理与OF处理相比,产量分别增加17.7%和10.8%,其差异达显著水平。
4、不同土壤验证防控效果试验结果显示：生物有机肥能有效防控马铃薯青枯病的发生,促进马铃薯植株生长,提高马铃薯产量。连作带病土温室盆栽试验中,CK发病率为39.7%,BI023和BI036处理对马铃薯青枯病的防病率皆为100%；健康土添加青枯菌温室盆栽试验中,cK发病率为57.7%,BI023和BI036处理对马铃薯青枯病的防病率分别为84.6%和80.8%。在两块青枯病感染程度不同的两块马铃薯田块(田块1感染程度较轻,青枯病发病率为8.3%；田块2感染程度较重,青枯病发病率为18.3%),进行马铃薯青枯病生防试验,结果表明：在田块1中,BI023和BI036处理的防病率分别为92.0%和84.0%,与OF处理相比产量分别提高42.3%和28.8%；在田块2中,BI023和BI036处理的防病率分别为65.6%和57.7%,与OF处理相比产量分别提高56.8%和45.1%。生物有机肥能有效防控马铃薯青枯病,并显著提高马铃薯产量。
5、施用生物有机肥可以改善马铃薯植株根际土壤微生物区系。不同土壤盆栽试验结果表明,对不同时期马铃薯植株根际土壤微生物计数结果显示,与CK相比,施用生物有机肥(BI023和BI036)显著促进马铃薯植株根际土壤细菌、放线菌的数量增加,真菌的数量减少。同时,与单一施用生防菌菌悬液处理(LH23和LH36)相比,生物有机肥(BI023和BI036)处理马铃薯植株根际土壤拮抗菌数量显著增加,马铃薯植株根际土壤青枯菌数量显著降低,防病率显著提高。田间试验中的根际土壤微生物区系变化与盆栽结果一致。同时生物有机肥显著提高马铃薯植株叶片过氧化物酶(POD)、过氧化氢酶(CAT)、超氧化物歧化酶(SOD)和多酚氧化酶(PPO)活性,降低丙二醛(MDA)含量,减轻发病情况,提高马铃薯植物的系统抗性。
综上所述,拮抗菌株与有机肥结合制成的生物有机肥(BI023和BI036)能有效防控马铃薯青枯病的发生,并促进马铃薯植株的生长、提高产量和改善品质。施用生物有机肥能促进拮抗菌在土壤和根际中的定殖和生存,有效降低青枯菌的数量,同时改善马铃薯植株根际微生物区系,提高植株系统抗性。本文的研究结果有助于研发马铃薯土传病害防控技术,为马铃薯青枯病的有效防控提供理论基础。
Potato bacterial wilt(Ralstonia solanacearum) is a soil-borne disease that seriously suppressed the potato plant(Solanum tuberosum) growth worldwide and causes serious economic losses. A lot of methods have been used to prevent the disease, among which biocontrol has been proved to be the most promising one. Biological control is one of the important measures for overcoming soil-borne diseases, because of its effectiveness and environment-friendly use. Screening antagonists against pathogens is the vital work for the biocontrol of plant diseases.
In this study, two bacterial strains LH23(Bacillus amyloliquefaciens) and LH36(Bacillus subtilis) with antagonistic activity against R. solanacearum, were selected from the rhizosphere soil of healthy potato plant grown in the field strongly diseased by bacterial wilt. Pot and field experiments were conducted to evaluate the LH23(Bacillus amyloliquefaciens) and LH36(Bacillus subtilis) strains and their derived bio-organic fertilizers (BIO23and BIO36) as potential biocontrol agents against potato bacterial wilt and the related mechanisms were also investigated. The main results obtained are listed as follows:
(1) In this study, more than140bacterial strains (bacterial antagonists against R.solanacearum) were isolated from rhizosphere soil of potato, tomato, eggplant and banana. Cell-free filtrate of six antagonistic strains had much stronger antagonistic activity against four R. solanacearum strains with inhibitory zones over20mm. Based on antagonistic activity against four R. solanacearum strains and ten soil-borne fungal pathogens as well as biocontrol efficacy in the greenhouse, two bacterial strains LH23and LH36were selected out of many candidates as potential biocontrol agents.
(2) Based on the results of morphologic characteristics, physiological and biochemical properties and phylogenetic analysis of16S rRNA, the strains LH23and LH36, were identified as Bacillus amyloliquefaciens and Bacillus subtilis, respectively. Measurement from laboratory tests showed that the LH23and LH36were able to produce growth-promoting substances indole acetic acid (IAA), siderophore, prolease, amylase, glucanase, antagonism of extracellular protein and dissolve P in medium. And the LH23and LH36could decrease the content of extracellular Polysaeeharides produced by R. solanacearum, suggesting that LH23and LH36were effective antagonists against the pathogen of potato bacterial wilt.
(3) In order to find a suitable antagonist inoculation method, we compared the methods of root-dipping with soil-drenching in the aspects including rhizocompetence, and effect of promoting plant growth under greenhouse conditions. The drenching treatment resulted in a higher rhizocompetence and plant-yield increase, and this method was also easier to operate in the field on a large scale. Pot and field experiments were performed to investigate the promoting effects of LH23(Bacillus amyloliquefaciens) and LH36(Bacillus subtilis) strains and their derived bio-organic fertilizers (BIO23and BIO36) in potato growth. In greenhouse experiment, BIO23and BIO36significantly promoted potato plant growth, especially the root growth. The root fresh weight of BIO23and BIO36treatments were1.8and2.1times higher than those of control, respectively. The root lengths, root SurfArea, root AvgDiam and RootVolume of the plants treated with BIO23and BIO36were significantly higher than those of control. The potato yield increments on85days after the application of BIO23and BIO36were17.7%and10.8%, respectively as compared with OF (organic fertilizer) treatment in the field experiments.
(4) Application of BIO23and BIO36decreased the incidence of bacterial wilt disease and increased potato yields. In one greenhouse experiment, the biocontrol efficiency of BIO23and BIO36applied were100%and100%, which were the mostly successful treatment while the application of the strains LH23and LH36alone resulted in a biocontrol efficiency of85.1%and81.1%, respectively. In another greenhouse experiment, the biocontrol efficiency of BIO23was84.6%, which was the most successful treatment and BIO36was the second in terms of the biocontrol efficiency (80.8%); the increased percentages of potato yields when compared with the control (chemical fertilizer) were63.5%(BIO23),64.7%(BIO36)34.8%(LH23),33.6%(LH36) and20.7%(OF). In field experiments, the biocontrol efficacies of BIO23and BIO36treatments were92.0%and84.0%, and the yield increases of BIO23and BIO36treatments were42.3%and28.8%, respectively, when compared with the OF treatment.
(5) Bio-organic fertilizers (BIO23and BIO36) could alter the microbial community structure and improved plant health. The counts of bacteria and actinomyces in the rhizosphere soils were significantly increased in BIO23and BIO36treatments, whereas the fungi in the soil in the both treatments were decreased; the counts of antagonists, bacteria and actinobacteria in the rhizosphere soil were significantly increased in BIO23and BIO36treatments, whereas the counts of R. solanacearum and fungi in the soil in the both treatments were decreased in the greenhouse experiments. In addition, the changes in the microbial populations in field experiments were the same as those observed in the greenhouse experiment. In the pot experiment, Bio-organic fertilizers (BIO23and BIO36) significantly increased the activities of plant defence enzymes such as POD, CAT, SOD and PPO, in plant leaves, and decreased MDA content, and consequently relieved the symptoms of potato.
In conclusion, our studies showed that the bio-organic fertilizer, an appropriate combination of organic fertilizer (OF) with the antagonists LH23or LH36used in the present study, could effectively suppress the potato wilt, promote the growth of potato plants and thus the quality of potato tubers, which mainly depends on producing many antibiotics and growth-promoting substances and the alternation of soil microbial community, after the successful colonization of the strains in soils. Results in this study would be useful in the developping new methods to better control potato bacterial wilt disease. Further researches should be done in the molecular microbilogical aspects of the functions performed by bio-organic fertilizers in remediation of continuously cropping obstacles and in the promotion of plant growth and health exerted by functional microbes added.
引文
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