温室条件下芽孢杆菌的生物活性及对番茄青枯病菌的抑制机制研究
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摘要
由青枯病菌引起的番茄青枯病(TBW)是全球范围内最严重的细菌性病害之一。由于其寄主范围广并能够在土壤中长时间生存,至今尚无一种防治方法能在病害流行区对其进行有效控制。生物防治与其他防治方法相结合被证明具有持久、环保的特点,并且能够保障农业生态系统的生产力和稳定性。
本研究从番茄和马铃薯的根际土壤中分离到约200个根际细菌,并测定它们对青枯病菌T-91的拮抗作用,在离体试验中,Am1, Am2, Am5, Am50, Am47, D11, D12, D16, D29和H810株菌株表现出了高潜力的拮抗活性。在温室条件下,进一步筛选它们对青枯病菌的生防效果。结果发现,使用Am1, D16, D29和H8菌株处理的番茄植株,青枯病的发病率减少至81.1%-89.0%。与未处理组比较显著(P>0.05)增加了番茄的株高(22.7%-43.7%)和干重(47.93%-91.55%)并能有效抑制土传真菌赤霉病菌,瓜果腐霉菌和立枯丝核菌。采用16S rRNA基因序列、生理和生化测试、脂肪酸甲酯分析对这4株拮抗细菌菌进行鉴定,结果表明Aml和D29为解淀粉芽孢杆菌(Bacillus amyloliquefaciens),D16和H8分别为枯草芽孢杆菌(B. subtilis)和甲基营养型芽孢杆菌(B. methylotrophicus)。
为评估枯草芽孢杆菌4812和甲基营养型芽孢杆菌H8单独使用或结合两种植物防御诱导剂乙酰水杨酸(ASA)和DL-β-氨基丁酸(BABA)使用时的生防效果,分别在离体和活体条件下测试了它们对青枯病菌的抑制作用。结果表明,在温室和离体条件下,所有处理均对青枯菌具有显著的抑制作用。在离体条件下,H8+ASA和4812+H8被认为是最有效的处理。将处理剂在番茄种子上施用并浸泡土壤时,H8对该病害的抑制作用最强,而4812+H8+ASA的抑制作用最弱。此外,使用4812+H8,H8+ASA和4812+H8+ASA处理的植物中检测到高活性的苯丙氨酸解氨酶。使用H8,4812+H8和H8+ASA处理的植物中检测到过氧化物酶和多酚氧化酶的表达量最高。实时定量PCR结果证实,所有处理组的植物茎组织中的病菌浓度均显著降低,其中H8+ASA是最有效的处理组合。
生物活性机制研究显示,解淀粉芽孢杆菌Am1和D29,枯草芽孢杆菌D16和甲基营养型芽孢杆菌H8这4株菌都能产生吲哚乙酸和铁载体,而不产生氰化氢。此外,D29,AM1和H8都具有溶解磷的能力。芽孢杆菌菌株既具有形成生物膜的能力,也具有溶解和破坏青枯病菌生物膜的能力。常规PCR分析结果显示,这4株菌都具有合成脂肽的基因(bmyB,fenD,ituC,srfA和bacA),这些基因分别表达合成Bacillomycin,Fengycin,Iturin,Surfactin和Bacilysin.在温室试验中,分别观察到处理组的番茄植株的IAA含量,鲜重与干重之间存在正相关(0.777和0.686)的关系。
Tomato Bacterial wilt (TBW) caused by Ralstonia solanacearum is the most serious and worldwide bacterial disease. Due to its wide host range and ability to survive freely in the soil for extended period, no single strategy can provide effective control against this disease in regions where the pathogen is endemic. Biological control of TBW combined with other control approaches has proven to be more durable, eco-friendly and maintain stability and productivity of agro-ecosystems.
In this study, about200rhizobacterial isolates were isolated from tomato and potato rhizosphere and examined for their antagonistic activities against R. solanacearum T-91, the causal agent of TBW. Among these isolates,10isolates including Am1, Am2, Am5, Am50, Am47, D11, D12, D16, D29and H8showed high potential of antagonistic activity in in vitro assay and further screened for their biocontrol efficacy against R. solanacearum under greenhouse conditions. In greenhouse,81.1-89.0%reduction of disease incidence of TBW was recorded in the tomato plants treated with the isolates Am1, D16, D29and H8. The application of these biocontrol agents significantly (p>0.05) increased plant height by22.7-43.7%and dry weight by47.93-91.55%compared with non-treated control. The four Bacillus isolates showed ability to inhibit growth of the three soil-borne fungi(Fusarium graminearum, Pythium aphanidermatum and Rhizoctonia solani). Identification of four selected isolates using16SrRNA gene sequence, the biochemical and physiological tests and fatty acid methyl esters analysis assigned strains Am1and D29as Bacillus amyloliquefaciens, D16and H8as B. subtilis and B. methylotrophicus, respectively.
Another study was carried out to assess the biocontrol efficacy of B. subtilis4812and B. methylotrophicus H8individually or in combination with two plant defense inducers viz. Acetyl salicylic acid (ASA) and DL-Beta-aminobutyric acid (BABA) against R. solanacearum in vitro and in vivo. The results showed that the pathogen was significantly inhibited by all applied treatments in the greenhouse and in vitro tests. Under in vitro test the treatments H8+ASA and4812+H8were found to be the most effective treatments.
Applied on tomato seeds and as soil drenching, the disease was most inhibited by H8whereas4812+H8+ASA was the least effective treatment. High activity of phenylalanine ammonia-lyase was observed in the plants treated with4812+H8, H8+ASA and4812+H8+ASA. The highest expression of peroxidase and polyphenoloxidase enzymes was found in the plants treated with H8,4812+H8and H8+ASA. The result of real time PCR confirmed that concentration of the pathogen in stem tissues was significantly reduced in all treated plants and H8+ASA was the most effective treatment.
The study of bioactivity mechanisms revealed that all four strains viz. Bacillus amyloliquefaciens D29, Am1, B. subtilis D16and B. methylotrophicus H8were able to produce Indole acetic acid and siderophores and can not produce Hydrogen cyanide (HCN). Moreover three of them D29, Am1and H8had capability to solubilize phosphor. Bacillus strains were able to form biofilm beside their ability to both membrane lysis and the destruction of Ralstonia biofilm. Results of conventional PCR analysis revealed that all the four strains are harboring the lipopeptides biosynthetic genes bmyB,fenD, ituC, srfAA and bacA which were responsible for biosynthesis of lipopeptides (Bacillomycin, Fengycin, Iturin, Surfactin and Bacilysin), respectively. Subsequent reverse transcription-polymerase chain reaction analysis revealed that ituC and srfAA biosynthesis genes in Bacillus strains Am1and D16gave the highest expression levels during in vitro interaction with R solanacearum which could suggest the potential antibacterial related mechanism is associated to their ability to secret the corresponding lipopeptide in surrounding niche. A positive correlation value (0.777and0.686) was observed between IAA amount, fresh weight and dry weight, respectively in treated tomato plants in a greenhouse trial.
本研究从番茄和马铃薯的根际土壤中分离到约200个根际细菌,并测定它们对青枯病菌T-91的拮抗作用,在离体试验中,Am1, Am2, Am5, Am50, Am47, D11, D12, D16, D29和H810株菌株表现出了高潜力的拮抗活性。在温室条件下,进一步筛选它们对青枯病菌的生防效果。结果发现,使用Am1, D16, D29和H8菌株处理的番茄植株,青枯病的发病率减少至81.1%-89.0%。与未处理组比较显著(P>0.05)增加了番茄的株高(22.7%-43.7%)和干重(47.93%-91.55%)并能有效抑制土传真菌赤霉病菌,瓜果腐霉菌和立枯丝核菌。采用16S rRNA基因序列、生理和生化测试、脂肪酸甲酯分析对这4株拮抗细菌菌进行鉴定,结果表明Aml和D29为解淀粉芽孢杆菌(Bacillus amyloliquefaciens),D16和H8分别为枯草芽孢杆菌(B. subtilis)和甲基营养型芽孢杆菌(B. methylotrophicus)。
为评估枯草芽孢杆菌4812和甲基营养型芽孢杆菌H8单独使用或结合两种植物防御诱导剂乙酰水杨酸(ASA)和DL-β-氨基丁酸(BABA)使用时的生防效果,分别在离体和活体条件下测试了它们对青枯病菌的抑制作用。结果表明,在温室和离体条件下,所有处理均对青枯菌具有显著的抑制作用。在离体条件下,H8+ASA和4812+H8被认为是最有效的处理。将处理剂在番茄种子上施用并浸泡土壤时,H8对该病害的抑制作用最强,而4812+H8+ASA的抑制作用最弱。此外,使用4812+H8,H8+ASA和4812+H8+ASA处理的植物中检测到高活性的苯丙氨酸解氨酶。使用H8,4812+H8和H8+ASA处理的植物中检测到过氧化物酶和多酚氧化酶的表达量最高。实时定量PCR结果证实,所有处理组的植物茎组织中的病菌浓度均显著降低,其中H8+ASA是最有效的处理组合。
生物活性机制研究显示,解淀粉芽孢杆菌Am1和D29,枯草芽孢杆菌D16和甲基营养型芽孢杆菌H8这4株菌都能产生吲哚乙酸和铁载体,而不产生氰化氢。此外,D29,AM1和H8都具有溶解磷的能力。芽孢杆菌菌株既具有形成生物膜的能力,也具有溶解和破坏青枯病菌生物膜的能力。常规PCR分析结果显示,这4株菌都具有合成脂肽的基因(bmyB,fenD,ituC,srfA和bacA),这些基因分别表达合成Bacillomycin,Fengycin,Iturin,Surfactin和Bacilysin.在温室试验中,分别观察到处理组的番茄植株的IAA含量,鲜重与干重之间存在正相关(0.777和0.686)的关系。
Tomato Bacterial wilt (TBW) caused by Ralstonia solanacearum is the most serious and worldwide bacterial disease. Due to its wide host range and ability to survive freely in the soil for extended period, no single strategy can provide effective control against this disease in regions where the pathogen is endemic. Biological control of TBW combined with other control approaches has proven to be more durable, eco-friendly and maintain stability and productivity of agro-ecosystems.
In this study, about200rhizobacterial isolates were isolated from tomato and potato rhizosphere and examined for their antagonistic activities against R. solanacearum T-91, the causal agent of TBW. Among these isolates,10isolates including Am1, Am2, Am5, Am50, Am47, D11, D12, D16, D29and H8showed high potential of antagonistic activity in in vitro assay and further screened for their biocontrol efficacy against R. solanacearum under greenhouse conditions. In greenhouse,81.1-89.0%reduction of disease incidence of TBW was recorded in the tomato plants treated with the isolates Am1, D16, D29and H8. The application of these biocontrol agents significantly (p>0.05) increased plant height by22.7-43.7%and dry weight by47.93-91.55%compared with non-treated control. The four Bacillus isolates showed ability to inhibit growth of the three soil-borne fungi(Fusarium graminearum, Pythium aphanidermatum and Rhizoctonia solani). Identification of four selected isolates using16SrRNA gene sequence, the biochemical and physiological tests and fatty acid methyl esters analysis assigned strains Am1and D29as Bacillus amyloliquefaciens, D16and H8as B. subtilis and B. methylotrophicus, respectively.
Another study was carried out to assess the biocontrol efficacy of B. subtilis4812and B. methylotrophicus H8individually or in combination with two plant defense inducers viz. Acetyl salicylic acid (ASA) and DL-Beta-aminobutyric acid (BABA) against R. solanacearum in vitro and in vivo. The results showed that the pathogen was significantly inhibited by all applied treatments in the greenhouse and in vitro tests. Under in vitro test the treatments H8+ASA and4812+H8were found to be the most effective treatments.
Applied on tomato seeds and as soil drenching, the disease was most inhibited by H8whereas4812+H8+ASA was the least effective treatment. High activity of phenylalanine ammonia-lyase was observed in the plants treated with4812+H8, H8+ASA and4812+H8+ASA. The highest expression of peroxidase and polyphenoloxidase enzymes was found in the plants treated with H8,4812+H8and H8+ASA. The result of real time PCR confirmed that concentration of the pathogen in stem tissues was significantly reduced in all treated plants and H8+ASA was the most effective treatment.
The study of bioactivity mechanisms revealed that all four strains viz. Bacillus amyloliquefaciens D29, Am1, B. subtilis D16and B. methylotrophicus H8were able to produce Indole acetic acid and siderophores and can not produce Hydrogen cyanide (HCN). Moreover three of them D29, Am1and H8had capability to solubilize phosphor. Bacillus strains were able to form biofilm beside their ability to both membrane lysis and the destruction of Ralstonia biofilm. Results of conventional PCR analysis revealed that all the four strains are harboring the lipopeptides biosynthetic genes bmyB,fenD, ituC, srfAA and bacA which were responsible for biosynthesis of lipopeptides (Bacillomycin, Fengycin, Iturin, Surfactin and Bacilysin), respectively. Subsequent reverse transcription-polymerase chain reaction analysis revealed that ituC and srfAA biosynthesis genes in Bacillus strains Am1and D16gave the highest expression levels during in vitro interaction with R solanacearum which could suggest the potential antibacterial related mechanism is associated to their ability to secret the corresponding lipopeptide in surrounding niche. A positive correlation value (0.777and0.686) was observed between IAA amount, fresh weight and dry weight, respectively in treated tomato plants in a greenhouse trial.
引文
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