细菌B-FS01抗菌物质的鉴定以及对串珠镰刀菌生长和伏马菌素B1产生的抑制效应
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摘要
串珠镰刀菌(Fusarium verticillioides或Fusarium moniliforme Sheldon)为非专性、非宿主特异性的病原菌，主要污染玉米、高粱、小麦、棉花、西红柿、花生、香蕉、大豆、青椒及某些饲料。串珠镰刀菌的水溶性代谢产物伏马菌素(Fumonisin FB)，可以引起动物各种疾病，如马脑白质软化症、猪肺水肿或大鼠肝癌，还有证据表明伏马菌素与人类食管癌的高发率有关。
本实验室从被油菜菌核病菌(Sclerotinia sclerotiorum)感染的油菜茎杆内分离得到一株内生拮抗菌，通过形态特征、培养性状、生理生化特性试验以及16S rDNA序列分析鉴定其为枯草芽孢杆菌(Bacillus subtilis)，暂定代号为B-FS01。研究发现B-FS01对串珠镰刀菌有强烈的拮抗作用，而且能够分泌抗菌物质到发酵液中。对其抗菌物质粗提液生化性质分析表明，该抗菌物质的抑菌性质对热稳定，耐碱不耐酸，对蛋白酶K和胃蛋白酶均不敏感。
通过硫酸铵沉淀、DE52离子交换层析、Sephadex G100分子筛层析、HPLC等手段，分离得到了该抗菌物质。通过HPLC-ESI MS分析发现，该类抗菌物质包含一簇分子量非常相近的化合物([M+H]m／z=1，434.0；1，435.9；1，450.6；1，464.0；1，479.0；1，492.9；1，506.0；1，519.9)，与Fengycins家族成员相似。进一步对各个分子进行的二级质谱分析证实这些化合物属于Fengycins家族，包括Fengycins A、Fengycins B和一类新发现的fengcyin。此后，我们对纯化手段进行了改进，引进中压制备系统，采用盐酸酸沉定与硅胶柱层析相结合，简化了Fengycins纯化步骤，可以进行半制备规模的Fengycins分离纯化工作。Fengycins家族成员构成非常接近，在一般研究和应用中，只需要对Fengycins家族的整体分析即可。液质联用仪过于昂贵，普通实验室难以具备，而SDS-PAGE不太适合小分子脂肽的分析。本研究发现在高于临界胶团浓度以上时，Fengycins能够像核酸分子一样在紫外光下被染料染色，基于这一性质，建立了通过琼脂糖电泳对Fengycins进行半定量的快速检测方法。
通过各种生测手段，测定了Fengycins对串珠镰刀菌抑制活性。结果显示，Fengycins在PDA固体培养基上对串珠镰刀菌菌丝径向生长的IC_(50)为20μg／mL，对孢子生长的MIC为0.78μg／杯；在PD液体培养基中对串珠镰刀菌菌丝生长量的IC_(50)为48μg／mL。此外Fengycins能够延滞串珠镰刀菌孢子萌发，而且能在活体玉米种子上表现出对串珠镰刀菌生长的抑制作用。
普通显微观察结果显示，Fengycins处理能使部分串珠镰刀菌菌丝顶端破裂。进一步通过PI染色与荧光显微观察发现，Fengycins处理会导致串珠镰刀菌菌丝膜的损伤。琼脂糖电泳结果表明，Fengycins能够与细胞膜上卵磷脂结合。Fengycins是两性抗菌肽，容易形成聚合体。所以我们认为Fengycins与膜上卵磷脂结合，并在那里生成聚合物造成膜穿孔，是一个导致膜破坏的主要原因。而向培养基里加入卵磷脂能缓解Fengycins的抑菌作用，在一定程度上支持了这一推论。最近研究表明，磷脂酶与真菌毒性密切相关。目前已证实磷脂酶是很多真菌的致病因子，而且通过以磷脂酶B为靶标筛选抗真菌药物的实验发现磷脂酶的抑制剂能够抑制病原菌的生长。我们发现Fengycins能够抑制串珠镰刀菌分泌的磷脂酶A_2的活性。该性质很可能也在Fengycins的抑菌活性中起到了一定的作用。
Fengycins除了能够抑制串珠镰刀菌的生长以外，还能够抑制其伏马菌素B1的产生。通过HPLC对串珠镰刀菌产生的伏马菌素分析，发现50μg／mLFengycins处理的串珠镰刀菌菌丝的产毒能力明显下降，单位重菌丝的产毒量仅为对照的28％。串珠镰刀菌体内控制伏马菌素产生的基因在基因组内呈簇状分布，其中FUM1和FUM8是负责伏马菌素产生的关键基因。RT-PCR实验结果表明：Fengycins处理对串珠镰刀菌体FUM1的转录水平没有明显影响，但是能够下调FUM8的转录水平。
Fusarium moniliforme Sheldon (Fusarium verticiIIioides) is a facultative and non-host necrotrophic pathogen, which broadly contaminates maize, sorghum, wheat, cotton, tomato, peanut, bananas, soybean, pimiento and feed. Fumonisins are the water-soluble metabolites produced by E moniliforme. They are acutely toxic to livestocks and cause diseses like leukoencephalomalacia in equines, pulmonary edema and hydrothorax in swine, and liver cancer in rats. In humans, the occurrence of fumonisin is correlated with higher incidence of esophageal cancer.
In this study, an antagonistic bacterial strain B-FS01 was isolated from the rape stem infected by SIerotinia Sclerotiorum. Following analysis of its cultural, morphological, biochemical and physiological characters and partial sequencing of 16S rDNA, strain B-FS01 was identified as Bacillus subtilis. B-FS01 showed strong antagonism against F. moniIiforme and released antifungal substances to culture. The characterization of crude extractions containing antifungal substances indicated that the antifungal activity was thermostable, tolerant to basic pH, untolerant to acid pH, unsensible to proteinase K and pepsin.
By approaches of ammonium sulphate precipitation, DE52 ion-exchange chromatography, Sephadex G100 gel filtration and HPLC, antifungal substances were able to be isolated and purified from the culture of B-FS01. The HPLC/ESI mass spectrum of antifungal substances revealed a cluster containing several molecules that was observed at [M+H] m/z =1,434.0; 1,435.9; 1,450.6; 1,464.0; 1,479.0; 1,492.9; 1,506.0 and 1,519.9. On the base of further CID spectrums of respective molecules, these compounds were identified as fengycin homologues containing Fengycins A, Fengycins B and a new fengycin type. In addition, by HC1 precipitation and silica gel column chromatography in the middle pressure preparative chromatography system, the purification of Fengycins from B-FS01 culture was greatly improved. The new approches are well suited for the semi-scale preparation of fengyicns.
HPLC/MS is competent for analysis, but it is too expensive to be accepted by most labs. Also, SDS-PAGE is not a good approach for analysis of lipopeptide with low molecular weight. Fortunately, it was found that the Fengycins at the concentration upper critical micelle concentration (CMC) could be dyed as well as nucleic acid in the ultraviolet radiation. With this finding, we established a method for the rapid semi -quantitative detection of Fengycins by agar-gel electrophoresis.
The bioassay of Fengycins against the growth of E moniliforme was performed. The results showed that IC_(50) of AAC inhibiting the mycelial radial growth was 20μg/mL; MIC of AAC by agar diffusion inhibiting the spore growth was 0.78/μg per cup and IC_(50) of AAC inhibiting the mycelial growth-weight was 48μg/mL. Additionaly, Fengycins could delay the germination of E moniliforme spores, and treatment with Fengycins significantly modified infection of maize seeds by ATCC 38932.
Microscopic observation on morphology of F. moniliforme mycelia revealed that tips of partial mycelia were lysed, and intensive PI staining of Fengycins-treated F. moniliforme mycelia was observed under fluorescence microscope, indicating the toxin-mediated damage of cytoplasmic membrane. We also performed agar gel electrophoresis and found that Fengycins could bind the phosphatidylcholine. Fengycins are amphiphilic compounds and can aggregate at a concentration. So it can be hypothesized that Fengycins interacted with the memberane by binding the phosphatidylcholine, then formed Fengycins/phospholipid aggregates, and eventually caused the formation of ion-conducting pores as well as the damage of F. moniliforme mycelia membrane. Our hypothesis could be supported by the anatagonism of phosphatidylcholine aganist the antifungal activity of Fengycins in agar media. Recently studies demonstrate that phopholipase is closely relevant to the virulence of fungi, and is the virulence factor for many fungi. In the screening of antifungal agents using phopholipase as a target, inhibitors of this enzyme also could arrest the growth of many pathogens. It was noted that Fengycins could depress activity of PLA_2 secreted by F. moniliforme, which might contribute to the antifungal activity against F.moniliforme.
In addition to the inhibitory activity against the growth of F. moniliforme, Fengycins was also shown to inhibit production of fumonisin B1 (FB1) . It was found that the mycotoxin-producd ability of F. moniliforme treated with 50/μg/mL Fengycins was significantly reduced, and the quantity of FB1 produced by unit mass mycelia decreased to 28% of the control. Genes involved in the fumonisins biosynthesis were identified and showed to be clustered in the genome of F. moniIiforme. Expression patern by reverse transcription-PCR (RT-PCR) showed that treatment of Fengycins did not affect transcriptional level of FUM1, but the level of FUM8 was down-regulated
本实验室从被油菜菌核病菌(Sclerotinia sclerotiorum)感染的油菜茎杆内分离得到一株内生拮抗菌，通过形态特征、培养性状、生理生化特性试验以及16S rDNA序列分析鉴定其为枯草芽孢杆菌(Bacillus subtilis)，暂定代号为B-FS01。研究发现B-FS01对串珠镰刀菌有强烈的拮抗作用，而且能够分泌抗菌物质到发酵液中。对其抗菌物质粗提液生化性质分析表明，该抗菌物质的抑菌性质对热稳定，耐碱不耐酸，对蛋白酶K和胃蛋白酶均不敏感。
通过硫酸铵沉淀、DE52离子交换层析、Sephadex G100分子筛层析、HPLC等手段，分离得到了该抗菌物质。通过HPLC-ESI MS分析发现，该类抗菌物质包含一簇分子量非常相近的化合物([M+H]m／z=1，434.0；1，435.9；1，450.6；1，464.0；1，479.0；1，492.9；1，506.0；1，519.9)，与Fengycins家族成员相似。进一步对各个分子进行的二级质谱分析证实这些化合物属于Fengycins家族，包括Fengycins A、Fengycins B和一类新发现的fengcyin。此后，我们对纯化手段进行了改进，引进中压制备系统，采用盐酸酸沉定与硅胶柱层析相结合，简化了Fengycins纯化步骤，可以进行半制备规模的Fengycins分离纯化工作。Fengycins家族成员构成非常接近，在一般研究和应用中，只需要对Fengycins家族的整体分析即可。液质联用仪过于昂贵，普通实验室难以具备，而SDS-PAGE不太适合小分子脂肽的分析。本研究发现在高于临界胶团浓度以上时，Fengycins能够像核酸分子一样在紫外光下被染料染色，基于这一性质，建立了通过琼脂糖电泳对Fengycins进行半定量的快速检测方法。
通过各种生测手段，测定了Fengycins对串珠镰刀菌抑制活性。结果显示，Fengycins在PDA固体培养基上对串珠镰刀菌菌丝径向生长的IC_(50)为20μg／mL，对孢子生长的MIC为0.78μg／杯；在PD液体培养基中对串珠镰刀菌菌丝生长量的IC_(50)为48μg／mL。此外Fengycins能够延滞串珠镰刀菌孢子萌发，而且能在活体玉米种子上表现出对串珠镰刀菌生长的抑制作用。
普通显微观察结果显示，Fengycins处理能使部分串珠镰刀菌菌丝顶端破裂。进一步通过PI染色与荧光显微观察发现，Fengycins处理会导致串珠镰刀菌菌丝膜的损伤。琼脂糖电泳结果表明，Fengycins能够与细胞膜上卵磷脂结合。Fengycins是两性抗菌肽，容易形成聚合体。所以我们认为Fengycins与膜上卵磷脂结合，并在那里生成聚合物造成膜穿孔，是一个导致膜破坏的主要原因。而向培养基里加入卵磷脂能缓解Fengycins的抑菌作用，在一定程度上支持了这一推论。最近研究表明，磷脂酶与真菌毒性密切相关。目前已证实磷脂酶是很多真菌的致病因子，而且通过以磷脂酶B为靶标筛选抗真菌药物的实验发现磷脂酶的抑制剂能够抑制病原菌的生长。我们发现Fengycins能够抑制串珠镰刀菌分泌的磷脂酶A_2的活性。该性质很可能也在Fengycins的抑菌活性中起到了一定的作用。
Fengycins除了能够抑制串珠镰刀菌的生长以外，还能够抑制其伏马菌素B1的产生。通过HPLC对串珠镰刀菌产生的伏马菌素分析，发现50μg／mLFengycins处理的串珠镰刀菌菌丝的产毒能力明显下降，单位重菌丝的产毒量仅为对照的28％。串珠镰刀菌体内控制伏马菌素产生的基因在基因组内呈簇状分布，其中FUM1和FUM8是负责伏马菌素产生的关键基因。RT-PCR实验结果表明：Fengycins处理对串珠镰刀菌体FUM1的转录水平没有明显影响，但是能够下调FUM8的转录水平。
Fusarium moniliforme Sheldon (Fusarium verticiIIioides) is a facultative and non-host necrotrophic pathogen, which broadly contaminates maize, sorghum, wheat, cotton, tomato, peanut, bananas, soybean, pimiento and feed. Fumonisins are the water-soluble metabolites produced by E moniliforme. They are acutely toxic to livestocks and cause diseses like leukoencephalomalacia in equines, pulmonary edema and hydrothorax in swine, and liver cancer in rats. In humans, the occurrence of fumonisin is correlated with higher incidence of esophageal cancer.
In this study, an antagonistic bacterial strain B-FS01 was isolated from the rape stem infected by SIerotinia Sclerotiorum. Following analysis of its cultural, morphological, biochemical and physiological characters and partial sequencing of 16S rDNA, strain B-FS01 was identified as Bacillus subtilis. B-FS01 showed strong antagonism against F. moniIiforme and released antifungal substances to culture. The characterization of crude extractions containing antifungal substances indicated that the antifungal activity was thermostable, tolerant to basic pH, untolerant to acid pH, unsensible to proteinase K and pepsin.
By approaches of ammonium sulphate precipitation, DE52 ion-exchange chromatography, Sephadex G100 gel filtration and HPLC, antifungal substances were able to be isolated and purified from the culture of B-FS01. The HPLC/ESI mass spectrum of antifungal substances revealed a cluster containing several molecules that was observed at [M+H] m/z =1,434.0; 1,435.9; 1,450.6; 1,464.0; 1,479.0; 1,492.9; 1,506.0 and 1,519.9. On the base of further CID spectrums of respective molecules, these compounds were identified as fengycin homologues containing Fengycins A, Fengycins B and a new fengycin type. In addition, by HC1 precipitation and silica gel column chromatography in the middle pressure preparative chromatography system, the purification of Fengycins from B-FS01 culture was greatly improved. The new approches are well suited for the semi-scale preparation of fengyicns.
HPLC/MS is competent for analysis, but it is too expensive to be accepted by most labs. Also, SDS-PAGE is not a good approach for analysis of lipopeptide with low molecular weight. Fortunately, it was found that the Fengycins at the concentration upper critical micelle concentration (CMC) could be dyed as well as nucleic acid in the ultraviolet radiation. With this finding, we established a method for the rapid semi -quantitative detection of Fengycins by agar-gel electrophoresis.
The bioassay of Fengycins against the growth of E moniliforme was performed. The results showed that IC_(50) of AAC inhibiting the mycelial radial growth was 20μg/mL; MIC of AAC by agar diffusion inhibiting the spore growth was 0.78/μg per cup and IC_(50) of AAC inhibiting the mycelial growth-weight was 48μg/mL. Additionaly, Fengycins could delay the germination of E moniliforme spores, and treatment with Fengycins significantly modified infection of maize seeds by ATCC 38932.
Microscopic observation on morphology of F. moniliforme mycelia revealed that tips of partial mycelia were lysed, and intensive PI staining of Fengycins-treated F. moniliforme mycelia was observed under fluorescence microscope, indicating the toxin-mediated damage of cytoplasmic membrane. We also performed agar gel electrophoresis and found that Fengycins could bind the phosphatidylcholine. Fengycins are amphiphilic compounds and can aggregate at a concentration. So it can be hypothesized that Fengycins interacted with the memberane by binding the phosphatidylcholine, then formed Fengycins/phospholipid aggregates, and eventually caused the formation of ion-conducting pores as well as the damage of F. moniliforme mycelia membrane. Our hypothesis could be supported by the anatagonism of phosphatidylcholine aganist the antifungal activity of Fengycins in agar media. Recently studies demonstrate that phopholipase is closely relevant to the virulence of fungi, and is the virulence factor for many fungi. In the screening of antifungal agents using phopholipase as a target, inhibitors of this enzyme also could arrest the growth of many pathogens. It was noted that Fengycins could depress activity of PLA_2 secreted by F. moniliforme, which might contribute to the antifungal activity against F.moniliforme.
In addition to the inhibitory activity against the growth of F. moniliforme, Fengycins was also shown to inhibit production of fumonisin B1 (FB1) . It was found that the mycotoxin-producd ability of F. moniliforme treated with 50/μg/mL Fengycins was significantly reduced, and the quantity of FB1 produced by unit mass mycelia decreased to 28% of the control. Genes involved in the fumonisins biosynthesis were identified and showed to be clustered in the genome of F. moniIiforme. Expression patern by reverse transcription-PCR (RT-PCR) showed that treatment of Fengycins did not affect transcriptional level of FUM1, but the level of FUM8 was down-regulated
引文
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