生物有机肥防控土传番茄青枯病的效果及其机制研究
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摘要
番茄青枯病是由青枯菌Ralstonia solanacearum引起的一种毁灭性土传细菌病害,严重制约着番茄产业的发展和经济效益的提高。生物防治因对环境、生态和人类健康安全而受到了国内外的关注,并在防治土传病害中发挥着越来越重要的作用。利用有益微生物防治青枯病已有大量的报道,但总体还处在生防菌资源收集阶段,缺乏稳定有效的生防产品。本研究从健康番茄植株根际分离获得解淀粉芽孢杆菌Bacillus amyloliquefaciens QL-18和皮氏劳尔氏菌Ralstonia pickettii QL-A6,分别研制出抗番茄青枯病的生物有机肥和液体菌剂。通过温室盆栽和田间试验研究了生物有机肥和液体菌剂单独或复合施用控制番茄青枯病的效果,并探讨了两者的作用机理。主要结果如下：
1、调查了南京麒麟镇后村蔬菜种植区番茄青枯病的现状,结果表明：春季发病率一般低于秋季,前茬为寄主作物的田块发病率高于前茬为非寄主作物的田块。灌溉水源中没有检测到致病菌,但在灌溉沟渠中检测到致病菌。利用选择性培养基M-SMSA从番茄植株根际分离获得流动型(疑似致病力青枯菌)和非流动型菌株(疑似无致病力青枯菌),根据形态特征、生理生化、特异性基因片段扩增、16S rRNA基因序列、致病性鉴定等手段,最终确定流动型菌株为强致病力R. solanacearum (其中一株命名为QL-Rs1115);非流动型菌株为R. pickettii (其中一株命名为QL-A6)和R. mannitolilytica。
2、从健康番茄植株根际分离到37株对青枯菌QL-Rs1115有抑制能力的细菌,其中B. amyloliquefaciens QL-18对青枯菌的平板抑制作用最强,在健康土壤(水稻土)和带病土壤(番茄连作土)中均显著抑制了青枯病发病率,减少了青枯菌在番茄植株根际和茎基部的数量。生防菌QL-18具有合成Fengycin、Subtilisin和Iturin等抗生素的能力,在PDA平板上能抑制Rhizoctonia solani, Phytophthora capsici, P. nicotianae, Fusarium oxysporum f. sp. niveum, F. oxysporum f. sp. cucumuerimu, F. oxysporum f. sp. melonis和Verticillium dahliae Kleb等植物土传病原真菌。GFP标记研究发现生防菌QL-18在番茄植株根表具有较好的定殖能力。
3、以菜粕堆肥(RCC)、猪粪堆肥(PMC)、菜粕和猪粪堆肥1：1混合物(R&P)、鸡粪堆肥(CMC)、牛粪堆肥(CDC)和中药渣堆肥(HRC)和2种泥炭(高位泥炭MP和低位泥炭FP)为原料,筛选适合生防菌QL-18的营养载体。结果表明,RCC不但能为生防菌QL-18提供大量的可溶性C、N养分,使其在载体中更好的繁殖和存活；而且能抑制青枯菌QL-Rs1115的生长,是菌株QL-18的最佳营养载体。在FP中生防菌QL-18也能保持很高的数量,但与其它堆肥载体相比,FP延长了青枯菌QL-Rs1115的存活时间,因此不是最适的生防菌营养载体。
4、将生防菌QL-18与菜粕堆肥二次固体发酵制成生物有机肥BOF。盆栽试验中,在育苗穴盘和移栽盆钵中均施用BOF的处理,番茄植株根际青枯菌的数量显著减少,有效降低了青枯病发病率。田间条件下BOF的控病效果因季节不同而异,春季施用BOF防效比秋季好,且育苗和移栽时均施用BOF的处理防效高于单施处理。番茄植株青枯病发病率与土壤中初始青枯菌的含量呈显著正相关,土壤初始病原菌数量越高,BOF的防病效果越差。
5、盆栽试验研究发现R. pickettii QL-A6在番茄根际和茎部定殖数量达到107cfu/gdw和108cfu/g fps,具备和青枯菌竞争的能力。灌根法施用R. pickettii QL-A6,能显著降低根际病原菌数量,有效防控苗期番茄青枯病的发生；增加R. pickettii QL-A6的施用量,能显著提高其生防效率。茎部注射法施用R. pickettii QL-A6的防控效果显著高于灌根法,R. pickettii QL-A6在地上部各组织大量定殖,抑制了病原菌对番茄的侵染。田间试验结果表明,茎部注射法施用R. pickettii QL-A6能有效降低番茄青枯病的发病率。
6、温室和田间试验结果表明,育苗施用BOF和茎部注射施用R. pickettii QL-A6两种方法配施能稳定防控番茄青枯病(70%-86%)。田间单独施用BOF或R. pickettiiQL-A6的防效均受季节影响：BOF处理的春季防效好(54.7%),秋季防效差(30.5%)；R. pickettii QL-A6处理春季防效差(31.6%),秋季防效好(80.0%)。育苗施用BOF后,生防菌QL-18能在番茄根际大量定殖,降低根际青枯菌的数量；茎部注射施用后,R. pickettii QL-A6在地上部快速大量定殖,减缓了青枯菌在地上部的侵染速度。
Bacterial wilt of tomato (BWT), a destructive soilborne disease caused by Ralstonia solanacearum, is a threat to tomato production and economical benefit. No effective measures have been developed yet. Environmentally friendly pesticide alternatives such as biolgical control have been suggested as integrated control strategies for BWT. Many beneficial microorganisms (BMs) have been screened for suppression of this disease; however, bio-products based on these BMs that could stably and efficiently control the disease are scare. In present study, BMs such as Bacillus amyloliquefaciens QL-18and Ralstonia pickettii QL-A6were isolated from the rhizosphere soil of healthy tomato plants. A bio-organic fertilizer (BOF) fortified with B. amyloliquefaciens QL-18and a liquid bio-product based on R. pickettii QL-A6were developed for suppression of BWT. A serial of experiments were conducted to test their control efficacies of BWT under greenhouse and field conditions. The possible control mechanisms were also discussed.
The main results obtained were listed as follows:
1. Field surveys on the epidemiolgy of BWT were undertaken in major tomato growing districts of the Qilin town, China. Disease incidences varied along with the crop seasons (spring and autumn) and previous crops (host and non-host plants). The pathogen was frequently detected from plant, soil and irrigation canals, not from irrigation water reservoir. Two phenotypically different colonies, fluidal (virulent R. solanacearum-Wke) and afluidal (avirulent R. solanacearum-liko) types were obtained from the rhizosphere soil using a modified semi-selective medium, South Africa (M-SMSA). Based on the results of the morpholgical characteristics, physiolgical and biochemical tests, amplication of R. solanacearum-specific genes,16S rRNA gene sequence and pathogenicity tests, the fluidal types were conformed as virulent R. solanacearum strains; and the afluidal types belong to R. pickettii and R. mannitolilytica.
2. Thirty-seven out of503rhizobacteria isolated from the rhizosphere soil of healthy tomato plants showed inhibitory effects against R. solanacearum strain QL-Rs1115. B. amyloliquefaciens QL-18performed best in suppression of R. solanacearum in vitro and efficiently reduced the disease incidences and populations of R. solanacearum in the rhizosphere soil and crown of tomato plants growing in both of paddy soil and soil continuously cropped with tomato plants. B. amyloliquefaciens QL-18has the capacities of producing antibiotics, such as fengycin, subtilisin and iturin, and inhibiting the growth of the phytopathogens including R. solanacearum, R. solani, P. capsici, P. nicotianae, F. oxysporum f. sp. Niveum, F. oxysporum f. sp. Cucumuerimu, F. oxysporum f. sp. melonis and V. dahliae Kleb. B. amyloliquefaciens QL-18successfully colonized the surface of tomato roots; this was conformed by labeling B. amyloliquefaciens QL-18with gfp and detecting the fluorescent cells under a fluorescence microscope.
3. Six composts, i.e., rapeseed cake compost (RCC), pig manure compost (PMC), mixture (1:1w/w) of RCC and PMC (R&P), chicken manure compost (CMC), cow dung compost (CDC), herb residue compost (HRC), and two peats, i.e., mosspeat (MP) and fenpeat (FP) were used as carriers of B. amyloliquefaciens QL-18. The results showed that RCC was the best carrier for B. amyloliquefaciens QL-18, because (1) B. amyloliquefaciens QL-18survived best in RCC, which could supply B. amyloliquefaciens QL-18with enough dissolvable organic carbon and nitrogen for multiplication;(2) RCC inhibited the growth of R. solanacearum. B. amyloliquefaciens QL-18could also survive well in the carrier FP, but the pathogen R. solanacearum survived in FP with large population (>105cfu/g dc) for more than60days, while R. solanacearum in RCC could not be detected15days after conservation. Thus, FP is unsuitable to be a carrier of a BM antagonistic against R. solanacearm.
4. The effect of BOF, a bio-product of RCC fortified with B. amyloliquefaciens QL-18was tested on suppression of BWT under both of the greenhouse and field condition. The best biocontrol efficiency was obtained by application of BOF into substrate during the nursery phase of tomato seedling followed by a second application to pot soil when seedlings were transplanted in greenhouse. However, the suppressiveness in field depended on the crop seasons. Disease incidence in the spring crop season was significantly reduced, while a low suppressive effect was observed in the autumn crop season. Disease incidences were significantly and positively correlated with the initial (before transplanting) R. solanacearum population in the soil.
5. Due to the good performances in colonization of the rhizosphere soil and stem of tomato, R. pickettii QL-A6was selected and developed as a liquid bio-product for suppression of R. solanacearum by use of the soil drench (SD) and stem injection (SI) methods in greenhouse. By the SI method, disease incidence was reduced by71.2%on the average with an inoculation dosage only about105cfu of R. pickettii QL-A6per plant. By the SD method, disease incidence was reduced by52.9%on the average but needed inoculation dosage was as high as about109cfu of R. pickettii QL-A6per plant. Thus, the SI method was chosen further test in field. The field disease incidence in R. pickettii QL-A6treated plots was8.8%at harvest time, while that in the sterilized water treated plots was as high as33.1%. It is concluded that direct injection of R. pickettii QL-A6in stem of tomato could be an alternative to chemical pesticides for biocontrol of R. solanacearum.
6. The effects of application of BOF and injection of R. pickettii QL-A6alone or in combination on suppression of BWT were investigated under greenhouse and field conditions. The results showed that combined treatment achieved better control effects (70-86%). The control efficacies of application of BOF and R. pickettii QL-A6alone altered along with the crop seasons in field. The control efficacies of BOF treatment was high (54.7%) in spring crop season, while was low (30.5%) in fall crop season. The control efficacies of stem injection of R. pickettii QL-A6was high (80.0%) in fall crop season, while was low (31.6%) in spring crop season. As BOF was amended with the seedling substrate, B. amyloliquefaciens QL-18colonized in tomato rhizosphere with large population (>106cfu/g dw) and reduced the population of R. solanacearum in the rhizosphere soil. The delay of multiplication of R. solanacearum in the aboveground parts of plants may due to the systemically colonization of both shoots and roots after R. pickettii QL-A6cells were directly injected into the plant stem, in which R. pickettii QL-A6may form a series of biofilms or other barriers and compete against the pathogen for spaces and nutrients.
1、调查了南京麒麟镇后村蔬菜种植区番茄青枯病的现状,结果表明：春季发病率一般低于秋季,前茬为寄主作物的田块发病率高于前茬为非寄主作物的田块。灌溉水源中没有检测到致病菌,但在灌溉沟渠中检测到致病菌。利用选择性培养基M-SMSA从番茄植株根际分离获得流动型(疑似致病力青枯菌)和非流动型菌株(疑似无致病力青枯菌),根据形态特征、生理生化、特异性基因片段扩增、16S rRNA基因序列、致病性鉴定等手段,最终确定流动型菌株为强致病力R. solanacearum (其中一株命名为QL-Rs1115);非流动型菌株为R. pickettii (其中一株命名为QL-A6)和R. mannitolilytica。
2、从健康番茄植株根际分离到37株对青枯菌QL-Rs1115有抑制能力的细菌,其中B. amyloliquefaciens QL-18对青枯菌的平板抑制作用最强,在健康土壤(水稻土)和带病土壤(番茄连作土)中均显著抑制了青枯病发病率,减少了青枯菌在番茄植株根际和茎基部的数量。生防菌QL-18具有合成Fengycin、Subtilisin和Iturin等抗生素的能力,在PDA平板上能抑制Rhizoctonia solani, Phytophthora capsici, P. nicotianae, Fusarium oxysporum f. sp. niveum, F. oxysporum f. sp. cucumuerimu, F. oxysporum f. sp. melonis和Verticillium dahliae Kleb等植物土传病原真菌。GFP标记研究发现生防菌QL-18在番茄植株根表具有较好的定殖能力。
3、以菜粕堆肥(RCC)、猪粪堆肥(PMC)、菜粕和猪粪堆肥1：1混合物(R&P)、鸡粪堆肥(CMC)、牛粪堆肥(CDC)和中药渣堆肥(HRC)和2种泥炭(高位泥炭MP和低位泥炭FP)为原料,筛选适合生防菌QL-18的营养载体。结果表明,RCC不但能为生防菌QL-18提供大量的可溶性C、N养分,使其在载体中更好的繁殖和存活；而且能抑制青枯菌QL-Rs1115的生长,是菌株QL-18的最佳营养载体。在FP中生防菌QL-18也能保持很高的数量,但与其它堆肥载体相比,FP延长了青枯菌QL-Rs1115的存活时间,因此不是最适的生防菌营养载体。
4、将生防菌QL-18与菜粕堆肥二次固体发酵制成生物有机肥BOF。盆栽试验中,在育苗穴盘和移栽盆钵中均施用BOF的处理,番茄植株根际青枯菌的数量显著减少,有效降低了青枯病发病率。田间条件下BOF的控病效果因季节不同而异,春季施用BOF防效比秋季好,且育苗和移栽时均施用BOF的处理防效高于单施处理。番茄植株青枯病发病率与土壤中初始青枯菌的含量呈显著正相关,土壤初始病原菌数量越高,BOF的防病效果越差。
5、盆栽试验研究发现R. pickettii QL-A6在番茄根际和茎部定殖数量达到107cfu/gdw和108cfu/g fps,具备和青枯菌竞争的能力。灌根法施用R. pickettii QL-A6,能显著降低根际病原菌数量,有效防控苗期番茄青枯病的发生；增加R. pickettii QL-A6的施用量,能显著提高其生防效率。茎部注射法施用R. pickettii QL-A6的防控效果显著高于灌根法,R. pickettii QL-A6在地上部各组织大量定殖,抑制了病原菌对番茄的侵染。田间试验结果表明,茎部注射法施用R. pickettii QL-A6能有效降低番茄青枯病的发病率。
6、温室和田间试验结果表明,育苗施用BOF和茎部注射施用R. pickettii QL-A6两种方法配施能稳定防控番茄青枯病(70%-86%)。田间单独施用BOF或R. pickettiiQL-A6的防效均受季节影响：BOF处理的春季防效好(54.7%),秋季防效差(30.5%)；R. pickettii QL-A6处理春季防效差(31.6%),秋季防效好(80.0%)。育苗施用BOF后,生防菌QL-18能在番茄根际大量定殖,降低根际青枯菌的数量；茎部注射施用后,R. pickettii QL-A6在地上部快速大量定殖,减缓了青枯菌在地上部的侵染速度。
Bacterial wilt of tomato (BWT), a destructive soilborne disease caused by Ralstonia solanacearum, is a threat to tomato production and economical benefit. No effective measures have been developed yet. Environmentally friendly pesticide alternatives such as biolgical control have been suggested as integrated control strategies for BWT. Many beneficial microorganisms (BMs) have been screened for suppression of this disease; however, bio-products based on these BMs that could stably and efficiently control the disease are scare. In present study, BMs such as Bacillus amyloliquefaciens QL-18and Ralstonia pickettii QL-A6were isolated from the rhizosphere soil of healthy tomato plants. A bio-organic fertilizer (BOF) fortified with B. amyloliquefaciens QL-18and a liquid bio-product based on R. pickettii QL-A6were developed for suppression of BWT. A serial of experiments were conducted to test their control efficacies of BWT under greenhouse and field conditions. The possible control mechanisms were also discussed.
The main results obtained were listed as follows:
1. Field surveys on the epidemiolgy of BWT were undertaken in major tomato growing districts of the Qilin town, China. Disease incidences varied along with the crop seasons (spring and autumn) and previous crops (host and non-host plants). The pathogen was frequently detected from plant, soil and irrigation canals, not from irrigation water reservoir. Two phenotypically different colonies, fluidal (virulent R. solanacearum-Wke) and afluidal (avirulent R. solanacearum-liko) types were obtained from the rhizosphere soil using a modified semi-selective medium, South Africa (M-SMSA). Based on the results of the morpholgical characteristics, physiolgical and biochemical tests, amplication of R. solanacearum-specific genes,16S rRNA gene sequence and pathogenicity tests, the fluidal types were conformed as virulent R. solanacearum strains; and the afluidal types belong to R. pickettii and R. mannitolilytica.
2. Thirty-seven out of503rhizobacteria isolated from the rhizosphere soil of healthy tomato plants showed inhibitory effects against R. solanacearum strain QL-Rs1115. B. amyloliquefaciens QL-18performed best in suppression of R. solanacearum in vitro and efficiently reduced the disease incidences and populations of R. solanacearum in the rhizosphere soil and crown of tomato plants growing in both of paddy soil and soil continuously cropped with tomato plants. B. amyloliquefaciens QL-18has the capacities of producing antibiotics, such as fengycin, subtilisin and iturin, and inhibiting the growth of the phytopathogens including R. solanacearum, R. solani, P. capsici, P. nicotianae, F. oxysporum f. sp. Niveum, F. oxysporum f. sp. Cucumuerimu, F. oxysporum f. sp. melonis and V. dahliae Kleb. B. amyloliquefaciens QL-18successfully colonized the surface of tomato roots; this was conformed by labeling B. amyloliquefaciens QL-18with gfp and detecting the fluorescent cells under a fluorescence microscope.
3. Six composts, i.e., rapeseed cake compost (RCC), pig manure compost (PMC), mixture (1:1w/w) of RCC and PMC (R&P), chicken manure compost (CMC), cow dung compost (CDC), herb residue compost (HRC), and two peats, i.e., mosspeat (MP) and fenpeat (FP) were used as carriers of B. amyloliquefaciens QL-18. The results showed that RCC was the best carrier for B. amyloliquefaciens QL-18, because (1) B. amyloliquefaciens QL-18survived best in RCC, which could supply B. amyloliquefaciens QL-18with enough dissolvable organic carbon and nitrogen for multiplication;(2) RCC inhibited the growth of R. solanacearum. B. amyloliquefaciens QL-18could also survive well in the carrier FP, but the pathogen R. solanacearum survived in FP with large population (>105cfu/g dc) for more than60days, while R. solanacearum in RCC could not be detected15days after conservation. Thus, FP is unsuitable to be a carrier of a BM antagonistic against R. solanacearm.
4. The effect of BOF, a bio-product of RCC fortified with B. amyloliquefaciens QL-18was tested on suppression of BWT under both of the greenhouse and field condition. The best biocontrol efficiency was obtained by application of BOF into substrate during the nursery phase of tomato seedling followed by a second application to pot soil when seedlings were transplanted in greenhouse. However, the suppressiveness in field depended on the crop seasons. Disease incidence in the spring crop season was significantly reduced, while a low suppressive effect was observed in the autumn crop season. Disease incidences were significantly and positively correlated with the initial (before transplanting) R. solanacearum population in the soil.
5. Due to the good performances in colonization of the rhizosphere soil and stem of tomato, R. pickettii QL-A6was selected and developed as a liquid bio-product for suppression of R. solanacearum by use of the soil drench (SD) and stem injection (SI) methods in greenhouse. By the SI method, disease incidence was reduced by71.2%on the average with an inoculation dosage only about105cfu of R. pickettii QL-A6per plant. By the SD method, disease incidence was reduced by52.9%on the average but needed inoculation dosage was as high as about109cfu of R. pickettii QL-A6per plant. Thus, the SI method was chosen further test in field. The field disease incidence in R. pickettii QL-A6treated plots was8.8%at harvest time, while that in the sterilized water treated plots was as high as33.1%. It is concluded that direct injection of R. pickettii QL-A6in stem of tomato could be an alternative to chemical pesticides for biocontrol of R. solanacearum.
6. The effects of application of BOF and injection of R. pickettii QL-A6alone or in combination on suppression of BWT were investigated under greenhouse and field conditions. The results showed that combined treatment achieved better control effects (70-86%). The control efficacies of application of BOF and R. pickettii QL-A6alone altered along with the crop seasons in field. The control efficacies of BOF treatment was high (54.7%) in spring crop season, while was low (30.5%) in fall crop season. The control efficacies of stem injection of R. pickettii QL-A6was high (80.0%) in fall crop season, while was low (31.6%) in spring crop season. As BOF was amended with the seedling substrate, B. amyloliquefaciens QL-18colonized in tomato rhizosphere with large population (>106cfu/g dw) and reduced the population of R. solanacearum in the rhizosphere soil. The delay of multiplication of R. solanacearum in the aboveground parts of plants may due to the systemically colonization of both shoots and roots after R. pickettii QL-A6cells were directly injected into the plant stem, in which R. pickettii QL-A6may form a series of biofilms or other barriers and compete against the pathogen for spaces and nutrients.
引文
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