番茄灰霉菌拮抗放线菌及抑菌活性物质研究
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摘要
放线菌是农业微生物的重要类群,其代谢产物是农用抗生素的重要来源。本文从土壤样品中筛选得到对番茄灰霉病菌(Botrytis cinerea)具有高抑菌活性的拮抗放线S-091菌株,并对其进行了种类鉴定、抑菌活性评价、发酵条件、抑菌机制以及抑菌活性物质的分离、纯化及结构解析等研究。
从采自不同地区的33份土壤样品中分离得到了112个放线菌菌株,并从中筛选出1株对番茄灰霉菌具有强烈抑菌活性的S-091菌株,该菌株还对多种植物病原真菌均表现出强烈的拮抗活性。采用传统放线菌分类方法和分子生物学相结合的鉴定方法对S-091菌株进行了种类的鉴定。结果表明S-091菌株的菌丝表面光滑,孢子丝成螺旋卷曲形态,孢子为椭圆形,表面长有丰富的疣刺,结合菌株的形态、培养特征及生理生化指标及16srDNA遗传分析,鉴定S-091菌株为链霉菌属不吸水链霉菌武夷山变种(Streptomyces. ahygrocopicus var. wuyishan)。
采用单因子测定和正交试验方法对S-091菌株的发酵条件进行了筛选优化。结果表明：可溶性淀粉显著影响活性物质产量,花生饼粉次之,微量元素中KC1显著影响活性物质产量。最优培养基配方：以花生饼粉1%、可溶性淀粉4.5%、氯化钾0.4%、酵母粉0.2%及硝酸铵0.1%为最佳配比。最适宜发酵温度为32℃、摇床转速为140r·min-1,pH值7.0,空气量以250mL三角瓶装液100-125mL为最适。
采用显微观察方法研究了S-091菌株发酵液活性成分的粗提物对灰霉菌的抑菌机制。用S-091菌株发酵液粗提物分别处理灰霉菌菌丝和分生孢子3hr和0.5hr后,菌丝停止生长,孢子停止萌发,菌丝体的细胞和分生孢子的内溶物产生浓缩凝集现象,并被处理的幼嫩菌丝体细胞在生长点处出现畸形膨大,但未见细胞壁和细胞膜破损、内溶物外溢,电导率测定表明支持无电解质外渗结论。
对S-091菌株发酵液活性成分理化性质研究结果表明发酵液活性成分对可见光、紫外光及热等环境因子具有很好的稳定性,对pH5以下的酸性条件下活性不稳定。有机溶剂萃取试验表明发酵液活性成分不溶于乙酸乙酯等有机溶剂；草酸沉淀法、丙酮沉淀法处理发酵液后沉淀部分无活性。通过pH纸层析、捷克八溶剂系统和Betina溶剂系统纸层析及纸电泳试验初步确定活性成分为两种或多种组分的混合物,并且活性成分极性中等或偏大,其中一部分为弱碱性,一部分强碱性。
用大孔吸附树脂柱、硅胶柱层析等分离方法对发酵液的活性物质粗提物进行去杂分离后,再采用制备型高效液相色谱仪对样品纯化,获得了主要活性成分的纯化合物,种白色无定形粉末状物质。将获得的纯化合物进行质谱与核磁谱的测定及结构解析,确定该活性化合物分子量为695,分子式为C35H53NO13,化学名称为：(1S,3R,5R,7E,13E,15E,17E,19ES,21S,23R,24S,25R)-11-甲基-12-乙基-1,3,5,25-四羟基-21-((2R,3S,4S,5S,6R)-四氢-6-甲基-3,5-二羟基-4-氨基-2H-吡喃-2-烷氧基)-10,27-二嗯-9-氧代-二环[21.3.1]二十七-7,13,15,17,19-五烯-24-羧酸,确定该化合物四烯类大环内酯类活性物质。通过文献检索发现该化合物为Tetramycin A,Tetramycin A在农用抗生素研究方面未见文献报道。
Actinomycete is one kind of very important microbiotic populations for agriculture production. Its secondary metabolites are the major sources of agricultural antibiotics. In this study, an antagonistic actinomycete strain S-091for tomato grey mold pathogen Botrytis cinerea was isolated from soil samples and its biological classification, antimicrobial activity evaluation, antimicrobial mechanisms, active component(s) isolation and purification and molecular structure characterization have been conducted.
Only S-091among112actinomycete strains isolated from33soil samples collected from different regions across China was found to be with strong suppression activity against Botrytis cinerea. Besides, it shows significantly antagonistic activities against several plant fungal pathogens. S-091was identified with classic actinomycete classification principles combined molecular identification methods. Its mycelium is smooth with coiled spore-bearing mycelium and the spores are oval and finely echinulate on the surface. Based on morphological characteristics, cultivation, physiological and biochemical properties and16s rDNA sequence analysis, S-091was identified as Streptomyces ahygrocopicus var. wuyishan.
With single-factorial evaluation and orthogonal experimental design, the fermentation conditions for S-091were selected and optimized. Based on the results, we found that starch content affects most significantly the S-091antibiotic productivity than peanut cake. Among the tested microelements, KC1is a key factor for S-091antibiotic productivity. The optimum medium was composed of1%peanut cake,4.5%starch soluble,0.4%KC1,0.2%yeast extract and0.1%NH4NO3. The optimum fermentation conditions are temperature at32℃,140r·min-1and medium pH at7.0. It is suitable for250mL triangular flask to contain100-125mL of fermentation medium.
The suppression mechanisms of S-091against Botrytis cinerea was conducted through microscopic investigation. When mycellium and spores of Botrytis cinerea were treated with the crude extract of S-091broths, the cell contents of the fungal pathogen were concentrated, and the mycellium ceased growth and the spores could not germinate. The cells at the growth point of mycellium were unmormally swelled after treatment. There were no any evidences that cell wall and cell membrane of Botrytis cinerea were broken and its cell contents outflowed ater treatment with the crude extract of S-091broths. Additionally, conductivity detection experiment showed that there is no electrolyte leakage of the pathogen cells treated with the crude extract.
The physicochemical property of antibiotics from fermented broth of the strain S-091were studied. The results showed that the antibiotics from fermented broth was stability to visible light, ultraviolet light and heat, but lost antibiotic activity in acidic conditions of pH value less than5. The test result of organic solvent extraction to fermented broth showed that the active ingredients of fermentation liquid were insoluble in ethyl acetate and other organic solvents. The deposit when it was under pH value5was not antagonistic activity to pathogen, and the deposit from fermented broth when it was added5times acetone was not antagonistic activity to pathogen either. The active ingredients of fermentation liquid were tested by pH paper chromatography, Czech eight solvent system and Betina solvent system paper chromatography and paper electrophoresis, the results showed that active ingredients were composed by two or more components from fermented broth, and the molecular polarity of active components was nutral or large, one part of which was alkaline, the other part was strongly alkaline.
Using polymer adsorbents, silica gel column chromatograph and combined with other methods, the antibiotic crude extract of S-091broths was isolated and then purified with HPLC to produce the pure compound of the active component. It is an amorphic white powdery material. Through mass spectrometry and nuclear magnetic resonance spectrum analysis, the active component is confirmed as one kind of tetraene macrolide antibiotics with molecular weight of695and its chemical structure formula C35H53NO13, and it was named (1S,3R,5R,7E,13E,15E,17E,19E,21S,23R,24S,25R)-21-((2R,3S,4S,5S,6R)-4-amino-tetrahydr o-3,5-dihydroxy-6-methyl-2H-pyran-2-yloxy)-12-ethyl-1,3,5,25-tetrahydroxy-11-methyl-9-ox o-10,27-dioxa-bicyclo[21.3.1]heptacosa-7,13,15,17,19-pentaene-24-carboxylic acid. It was found to be tetramycin A through literature retrieval, which has not been reported to be used as an agricultural antibiotic.
从采自不同地区的33份土壤样品中分离得到了112个放线菌菌株,并从中筛选出1株对番茄灰霉菌具有强烈抑菌活性的S-091菌株,该菌株还对多种植物病原真菌均表现出强烈的拮抗活性。采用传统放线菌分类方法和分子生物学相结合的鉴定方法对S-091菌株进行了种类的鉴定。结果表明S-091菌株的菌丝表面光滑,孢子丝成螺旋卷曲形态,孢子为椭圆形,表面长有丰富的疣刺,结合菌株的形态、培养特征及生理生化指标及16srDNA遗传分析,鉴定S-091菌株为链霉菌属不吸水链霉菌武夷山变种(Streptomyces. ahygrocopicus var. wuyishan)。
采用单因子测定和正交试验方法对S-091菌株的发酵条件进行了筛选优化。结果表明：可溶性淀粉显著影响活性物质产量,花生饼粉次之,微量元素中KC1显著影响活性物质产量。最优培养基配方：以花生饼粉1%、可溶性淀粉4.5%、氯化钾0.4%、酵母粉0.2%及硝酸铵0.1%为最佳配比。最适宜发酵温度为32℃、摇床转速为140r·min-1,pH值7.0,空气量以250mL三角瓶装液100-125mL为最适。
采用显微观察方法研究了S-091菌株发酵液活性成分的粗提物对灰霉菌的抑菌机制。用S-091菌株发酵液粗提物分别处理灰霉菌菌丝和分生孢子3hr和0.5hr后,菌丝停止生长,孢子停止萌发,菌丝体的细胞和分生孢子的内溶物产生浓缩凝集现象,并被处理的幼嫩菌丝体细胞在生长点处出现畸形膨大,但未见细胞壁和细胞膜破损、内溶物外溢,电导率测定表明支持无电解质外渗结论。
对S-091菌株发酵液活性成分理化性质研究结果表明发酵液活性成分对可见光、紫外光及热等环境因子具有很好的稳定性,对pH5以下的酸性条件下活性不稳定。有机溶剂萃取试验表明发酵液活性成分不溶于乙酸乙酯等有机溶剂；草酸沉淀法、丙酮沉淀法处理发酵液后沉淀部分无活性。通过pH纸层析、捷克八溶剂系统和Betina溶剂系统纸层析及纸电泳试验初步确定活性成分为两种或多种组分的混合物,并且活性成分极性中等或偏大,其中一部分为弱碱性,一部分强碱性。
用大孔吸附树脂柱、硅胶柱层析等分离方法对发酵液的活性物质粗提物进行去杂分离后,再采用制备型高效液相色谱仪对样品纯化,获得了主要活性成分的纯化合物,种白色无定形粉末状物质。将获得的纯化合物进行质谱与核磁谱的测定及结构解析,确定该活性化合物分子量为695,分子式为C35H53NO13,化学名称为：(1S,3R,5R,7E,13E,15E,17E,19ES,21S,23R,24S,25R)-11-甲基-12-乙基-1,3,5,25-四羟基-21-((2R,3S,4S,5S,6R)-四氢-6-甲基-3,5-二羟基-4-氨基-2H-吡喃-2-烷氧基)-10,27-二嗯-9-氧代-二环[21.3.1]二十七-7,13,15,17,19-五烯-24-羧酸,确定该化合物四烯类大环内酯类活性物质。通过文献检索发现该化合物为Tetramycin A,Tetramycin A在农用抗生素研究方面未见文献报道。
Actinomycete is one kind of very important microbiotic populations for agriculture production. Its secondary metabolites are the major sources of agricultural antibiotics. In this study, an antagonistic actinomycete strain S-091for tomato grey mold pathogen Botrytis cinerea was isolated from soil samples and its biological classification, antimicrobial activity evaluation, antimicrobial mechanisms, active component(s) isolation and purification and molecular structure characterization have been conducted.
Only S-091among112actinomycete strains isolated from33soil samples collected from different regions across China was found to be with strong suppression activity against Botrytis cinerea. Besides, it shows significantly antagonistic activities against several plant fungal pathogens. S-091was identified with classic actinomycete classification principles combined molecular identification methods. Its mycelium is smooth with coiled spore-bearing mycelium and the spores are oval and finely echinulate on the surface. Based on morphological characteristics, cultivation, physiological and biochemical properties and16s rDNA sequence analysis, S-091was identified as Streptomyces ahygrocopicus var. wuyishan.
With single-factorial evaluation and orthogonal experimental design, the fermentation conditions for S-091were selected and optimized. Based on the results, we found that starch content affects most significantly the S-091antibiotic productivity than peanut cake. Among the tested microelements, KC1is a key factor for S-091antibiotic productivity. The optimum medium was composed of1%peanut cake,4.5%starch soluble,0.4%KC1,0.2%yeast extract and0.1%NH4NO3. The optimum fermentation conditions are temperature at32℃,140r·min-1and medium pH at7.0. It is suitable for250mL triangular flask to contain100-125mL of fermentation medium.
The suppression mechanisms of S-091against Botrytis cinerea was conducted through microscopic investigation. When mycellium and spores of Botrytis cinerea were treated with the crude extract of S-091broths, the cell contents of the fungal pathogen were concentrated, and the mycellium ceased growth and the spores could not germinate. The cells at the growth point of mycellium were unmormally swelled after treatment. There were no any evidences that cell wall and cell membrane of Botrytis cinerea were broken and its cell contents outflowed ater treatment with the crude extract of S-091broths. Additionally, conductivity detection experiment showed that there is no electrolyte leakage of the pathogen cells treated with the crude extract.
The physicochemical property of antibiotics from fermented broth of the strain S-091were studied. The results showed that the antibiotics from fermented broth was stability to visible light, ultraviolet light and heat, but lost antibiotic activity in acidic conditions of pH value less than5. The test result of organic solvent extraction to fermented broth showed that the active ingredients of fermentation liquid were insoluble in ethyl acetate and other organic solvents. The deposit when it was under pH value5was not antagonistic activity to pathogen, and the deposit from fermented broth when it was added5times acetone was not antagonistic activity to pathogen either. The active ingredients of fermentation liquid were tested by pH paper chromatography, Czech eight solvent system and Betina solvent system paper chromatography and paper electrophoresis, the results showed that active ingredients were composed by two or more components from fermented broth, and the molecular polarity of active components was nutral or large, one part of which was alkaline, the other part was strongly alkaline.
Using polymer adsorbents, silica gel column chromatograph and combined with other methods, the antibiotic crude extract of S-091broths was isolated and then purified with HPLC to produce the pure compound of the active component. It is an amorphic white powdery material. Through mass spectrometry and nuclear magnetic resonance spectrum analysis, the active component is confirmed as one kind of tetraene macrolide antibiotics with molecular weight of695and its chemical structure formula C35H53NO13, and it was named (1S,3R,5R,7E,13E,15E,17E,19E,21S,23R,24S,25R)-21-((2R,3S,4S,5S,6R)-4-amino-tetrahydr o-3,5-dihydroxy-6-methyl-2H-pyran-2-yloxy)-12-ethyl-1,3,5,25-tetrahydroxy-11-methyl-9-ox o-10,27-dioxa-bicyclo[21.3.1]heptacosa-7,13,15,17,19-pentaene-24-carboxylic acid. It was found to be tetramycin A through literature retrieval, which has not been reported to be used as an agricultural antibiotic.
引文
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