微生物有机肥对烟草土传黑胫病的防控效果及机理研究
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摘要
烟草土传黑胫病是烟草连作障碍的主要病害之一,该病害的防控方法已成为烟草生产的研究热点,以生物防控为主导的综合防控是防控该病害的有效方法。本研究以从云南和安徽黑胫病重病烟田中采集的健康烟株根际土壤中分离出的显著拮抗烟草寄生疫霉(Phytophthora parasitica var. nicotianae)的多粘类芽孢杆菌(Paenibacillus polymyxa)菌株C5(CGMCC No.3303)及其微生物有机肥为材料,采用盆栽试验和室内分析及室内培养等方法,研究了微生物有机肥对烟草土传黑胫病的生防效果及其机理,获得了如下主要结果：
1、利用菌株C5制备的微生物有机肥能有效防控烟草土传黑胫病的发生,苗期防控率达80%；菌株C5及其微生物有机肥的施用能改变烟草根际土壤的微生物区系,与连作土壤(CK2)相比,施用微生物有机肥处理(BOF)的土壤中真菌数量减少,细菌和放线菌数量有显著增加。其中,细菌和放线菌分别为CK2的2.4倍左右和4.4倍左右。与CK2相比,BOF的鲜重、干重和株高分别增加约1.8倍、1.7倍和1.4倍；BOF处理的烟草鲜重和株高也显著高于健康土处理(CK1)和施用有机肥处理(OF)处理,增加量达1.2-1.4倍。说明菌株C5对烟草也具有明显的促生作用,能提高烟草产量。
2、菌株C5发酵液的粗提物对病原菌有拮抗作用,其发酵粗提物中的抗真菌活性物质在低温和中性条件下稳定,对蛋白酶稳定。利用高效液相色谱,获得了1个对烟草寄生疫霉具有较强抑制作用的色谱峰。通过串联质谱仪测定其分子量分别为883.5和897.5Da；947.5和961.5Da。这两组物质都属于多肽类物质Fusaricidins。
3、烟草根系能分泌绿原酸、咖啡酸、丁香酸、阿魏酸、水杨酸和肉桂酸6种酚酸类物质。其中绿原酸、咖啡酸和丁香酸是在菌株C5或者烟草寄生疫霉的刺激下产生的,3种酚酸在低浓度时促进烟草寄生疫霉生长。烟草分泌的有机酸类物质主要是草酸和苹果酸,在浓度为40μM时,促进菌株C5的生长,菌株C5对有机酸有最大的趋化性。
4、标记菌株C5-GFP在根际的定殖行为结果表明,菌株C5-GFP在烟草根系的定殖部位主要分布在根尖和伸长区。菌株C5-GFP能够在烟草根际和根系大量定殖,接种6d以后大量菌体在根表和根际定殖,菌体数量达到106cfug-1root和106cfu g-1root.
综上所述,烟草施用含有菌株C5的微生物有机肥后,其中多粘类芽孢杆菌菌株C5通过在根际大量定殖、产生拮抗物质、刺激根系产生有利生长的物质等方式保护烟草免受病原真菌的侵害,是其防控烟草土传黑胫病的主要机制。
Phytophthora parasitica var. nicotianae, a soilborne pathogen, causes tobacco black shank. A lot of methods have been used to prevent the disease, among which biocontrol has been proved to be most promising. 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, an antagonistic strain of Paenibacillus polymyxa C5effective against Ph. parasitica var. nicotianae, was isolated from the rhizosphere soil of a healthy tobacco plant grown in the field strongly attacked by tobacco black shank. A novel bio-organic fertilizer (BOF) was developed by inocalating P. polymyxa C5and doing the second solid fermentation. Pot experiments were carried out to evaluate the biocontrol effect of this BIO against tobacco black shank, and the related mechanisms were also investigated. The main results obtained are listed as follows:
1. Pot experiments revealed that the application of bio-organic fertilizer containing strain C5(BOF) could decrease the incidence of tobacco black shank by80%compared with control (CK2) in which neither C5strain nor organic fertilizer were used in continuous cropping soil (diseased soil). The application of BOF could influence the microbial community of rhizosphere soil. The population of bacteria and actinomyces was increased by2.4times and4.4times compared with control, respectively.
The fresh weight, dried weight and height of tobacco in BOF were increased by about1.8times,1.7times and1.4times, respectively, compared with CK2. The fresh weight and plant height of tobacco in BOF were increased by about1.4times and1.3times compared with those of CK1(healthy soil). The fresh weight, dried weight and plant height of tobacco in BIO were increased by about1.4times,1.4times and1.2times compared to those of OF. These results indicate that the bio-organic fertilizer could not only control the tobacco black shank, but also promote tobacco plant growth.
2. The extracts of P. polymyxa C5had a significantly antifungal activity against the pathogen. Compounds with antifungal activity were detected in the extracts, which were stable at neutral pH and low temperatures, and resistant to proteinase. One peak of the crude compounds with antifungal activities was detected by high-performance liquid chromatography (HPLC). Liquid chromatography-electrospray ionization-mass spectrometry (LC-ESI-MS) analysis showed that the mass spectra of the peaks were characterized by two series of homologous ion peaks, one with molecular weights of883.5and897.5Da and the other with molecular weights of947.5and961.5Da. The two series of compounds were identified as Fusaricidins.
3. Six kinds of phenolic acids, including chlorogenic acid, caffeic acid, syringic acid, ferulic acid, salicylic acid and cinnamic acid were detected in the root exudates of tobacco by HPLC. However, chlorogenic acid, caffeic acid and syringic acid were excreted when the tobacco roots were excited by C5or the pathogen. The growth of Phytophthora parasitica var. nicotianae was promoted by low concentration of chlorogenic acid, caffeic acid and syringic acid. The main organic acids of root exudates from tobacco plants were oxalic acid and malic acid. The growth of stain C5was promoted in the presence of40μM oxalic acid and malic acid of40μM. Furthermore, C5showed the highest chemotaxis reaction response to these organic acids.
4. Colonization assay indicated that the green fluorescent P. polymyxa C5cells mainly colonized the root tips and elongation zone of the root surfaces, as well as root hairs. High densities of the introduced GFP-tagged C5strain could be recovered on the root surface and in the rhizosphere soil reaching approximately106cfu g-1root and106cfu g-1soil six days after the inoculation.
In conclusion, our studies showed that the bio-organic fertilizer, appropriate combinations of organic fertilizer with the antagonistic strain P. polymyxa C5used in the present study, could effectively suppress the tobacco black shank. To reduce the losses of tobacco production caused by Phytophthora parasitica var. nicotianae, the mechanism of the biocontrol of tobacco black shank should be further studied.
1、利用菌株C5制备的微生物有机肥能有效防控烟草土传黑胫病的发生,苗期防控率达80%；菌株C5及其微生物有机肥的施用能改变烟草根际土壤的微生物区系,与连作土壤(CK2)相比,施用微生物有机肥处理(BOF)的土壤中真菌数量减少,细菌和放线菌数量有显著增加。其中,细菌和放线菌分别为CK2的2.4倍左右和4.4倍左右。与CK2相比,BOF的鲜重、干重和株高分别增加约1.8倍、1.7倍和1.4倍；BOF处理的烟草鲜重和株高也显著高于健康土处理(CK1)和施用有机肥处理(OF)处理,增加量达1.2-1.4倍。说明菌株C5对烟草也具有明显的促生作用,能提高烟草产量。
2、菌株C5发酵液的粗提物对病原菌有拮抗作用,其发酵粗提物中的抗真菌活性物质在低温和中性条件下稳定,对蛋白酶稳定。利用高效液相色谱,获得了1个对烟草寄生疫霉具有较强抑制作用的色谱峰。通过串联质谱仪测定其分子量分别为883.5和897.5Da；947.5和961.5Da。这两组物质都属于多肽类物质Fusaricidins。
3、烟草根系能分泌绿原酸、咖啡酸、丁香酸、阿魏酸、水杨酸和肉桂酸6种酚酸类物质。其中绿原酸、咖啡酸和丁香酸是在菌株C5或者烟草寄生疫霉的刺激下产生的,3种酚酸在低浓度时促进烟草寄生疫霉生长。烟草分泌的有机酸类物质主要是草酸和苹果酸,在浓度为40μM时,促进菌株C5的生长,菌株C5对有机酸有最大的趋化性。
4、标记菌株C5-GFP在根际的定殖行为结果表明,菌株C5-GFP在烟草根系的定殖部位主要分布在根尖和伸长区。菌株C5-GFP能够在烟草根际和根系大量定殖,接种6d以后大量菌体在根表和根际定殖,菌体数量达到106cfug-1root和106cfu g-1root.
综上所述,烟草施用含有菌株C5的微生物有机肥后,其中多粘类芽孢杆菌菌株C5通过在根际大量定殖、产生拮抗物质、刺激根系产生有利生长的物质等方式保护烟草免受病原真菌的侵害,是其防控烟草土传黑胫病的主要机制。
Phytophthora parasitica var. nicotianae, a soilborne pathogen, causes tobacco black shank. A lot of methods have been used to prevent the disease, among which biocontrol has been proved to be most promising. 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, an antagonistic strain of Paenibacillus polymyxa C5effective against Ph. parasitica var. nicotianae, was isolated from the rhizosphere soil of a healthy tobacco plant grown in the field strongly attacked by tobacco black shank. A novel bio-organic fertilizer (BOF) was developed by inocalating P. polymyxa C5and doing the second solid fermentation. Pot experiments were carried out to evaluate the biocontrol effect of this BIO against tobacco black shank, and the related mechanisms were also investigated. The main results obtained are listed as follows:
1. Pot experiments revealed that the application of bio-organic fertilizer containing strain C5(BOF) could decrease the incidence of tobacco black shank by80%compared with control (CK2) in which neither C5strain nor organic fertilizer were used in continuous cropping soil (diseased soil). The application of BOF could influence the microbial community of rhizosphere soil. The population of bacteria and actinomyces was increased by2.4times and4.4times compared with control, respectively.
The fresh weight, dried weight and height of tobacco in BOF were increased by about1.8times,1.7times and1.4times, respectively, compared with CK2. The fresh weight and plant height of tobacco in BOF were increased by about1.4times and1.3times compared with those of CK1(healthy soil). The fresh weight, dried weight and plant height of tobacco in BIO were increased by about1.4times,1.4times and1.2times compared to those of OF. These results indicate that the bio-organic fertilizer could not only control the tobacco black shank, but also promote tobacco plant growth.
2. The extracts of P. polymyxa C5had a significantly antifungal activity against the pathogen. Compounds with antifungal activity were detected in the extracts, which were stable at neutral pH and low temperatures, and resistant to proteinase. One peak of the crude compounds with antifungal activities was detected by high-performance liquid chromatography (HPLC). Liquid chromatography-electrospray ionization-mass spectrometry (LC-ESI-MS) analysis showed that the mass spectra of the peaks were characterized by two series of homologous ion peaks, one with molecular weights of883.5and897.5Da and the other with molecular weights of947.5and961.5Da. The two series of compounds were identified as Fusaricidins.
3. Six kinds of phenolic acids, including chlorogenic acid, caffeic acid, syringic acid, ferulic acid, salicylic acid and cinnamic acid were detected in the root exudates of tobacco by HPLC. However, chlorogenic acid, caffeic acid and syringic acid were excreted when the tobacco roots were excited by C5or the pathogen. The growth of Phytophthora parasitica var. nicotianae was promoted by low concentration of chlorogenic acid, caffeic acid and syringic acid. The main organic acids of root exudates from tobacco plants were oxalic acid and malic acid. The growth of stain C5was promoted in the presence of40μM oxalic acid and malic acid of40μM. Furthermore, C5showed the highest chemotaxis reaction response to these organic acids.
4. Colonization assay indicated that the green fluorescent P. polymyxa C5cells mainly colonized the root tips and elongation zone of the root surfaces, as well as root hairs. High densities of the introduced GFP-tagged C5strain could be recovered on the root surface and in the rhizosphere soil reaching approximately106cfu g-1root and106cfu g-1soil six days after the inoculation.
In conclusion, our studies showed that the bio-organic fertilizer, appropriate combinations of organic fertilizer with the antagonistic strain P. polymyxa C5used in the present study, could effectively suppress the tobacco black shank. To reduce the losses of tobacco production caused by Phytophthora parasitica var. nicotianae, the mechanism of the biocontrol of tobacco black shank should be further studied.
引文
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