海洋中稻瘟病拮抗菌的分离、发酵优化及其活性成分分析
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摘要
稻瘟病是造成水稻减产最严重的病害之一,目前主要以化学防治为主。由于化学防治的局限性及对环境的破坏性,这种防治手段已造成越来越严重的社会问题。以利用微生物天然次生代谢产物为主的生物防控,不仅能有效防治作物病害,而且对环境不会造成污染,符合现代农业发展的需求。
海洋是一个巨大的生物资源库,但由于其特殊的生存环境,大部分海洋生物资源尚未被开发。随着生物技术的发展；越来越多的海洋微生物被开发、利用,它们的次生代谢产物为生物防控提供了新的策略。
本研究从海洋样品中分离、鉴定出对稻瘟菌有拮抗作用的细菌,通过发酵优化,提高了活性物质代谢产率；采用各种色谱法对活性物质进行了分离纯化,并测定活性物质的分子量,为后续开发、利用提供了实验依据。本研究的结果有以下几点：
1.拮抗菌株的分离、鉴定
从海样样品中分离到了13株对水稻稻瘟菌有明显抑制活性的菌株,通过16SrDNA单引物序比对,发现有ZJ387与Cobetia marina有最高的同源性(99.351%),其他12株菌株与高地芽孢杆菌(Bacillus altitudinis)、同温层芽孢杆菌(Bacillus stratosphericus)及嗜气芽孢杆菌(Bacillus aerophilus)有高度同源性,推测这几种菌株可能为同一种菌。我们选择抑菌活性更强的ZJ186作为目标菌株,用于后续研究。
通过形态学观察、生理生化指标测定以及16S rDNA系统进化树分析,确定ZJ186属于高地芽孢杆菌。
本文首次从海洋材料中分离到了高地芽孢杆菌,并首次报道了高地芽孢杆菌对稻瘟菌有拮抗作用。
2.拮抗菌株ZJ186的发酵优化
通过发酵培养基的筛选实验,选用Landy作为发酵基础培养基进行优化。经单因素优化试验,确定以最佳碳源糖蜜10g/L、20g/L、30g/L,最佳氮源菜籽粉2.5g/L5g/L、10g/L, NaCl0.75g/L、1.5g/L、3g/L, MgSO40.5g/L,1g/L、2g/L,发酵时间36h、48h、72h,温度28℃、32℃、37℃,初始pH值5、6、7,接种量1%、2%、4%,装液量10%、30%、50%等9种因素各三个水平采用L27313正交表进行优化。
经显著性检验,糖蜜(碳源)、NaCl、发酵时间、初始ph值及装液量的F值均达极显著水平,菜籽粉(氮源)、MgSO4和温度的F值达到显著水平,而接种量F值不显著。最优发酵组合为：A2B2C3D2E2F1G2I2。
发酵优化后的发酵液滤液相对抑菌率为67.54%，较优化之前的54.1%,优化效率提高了24.83%,较阳性对照三环唑抑菌活性(21.64%)提高了212.06%。
通过发酵优化实验,我们提高了稻瘟菌拮抗菌株ZJ186产生抑菌活性物质的能力,为该菌株的进一步开发奠定了研究基础。
3.抗菌活性物质的分离纯化及活性成分分析
通过抗菌图谱检测,高地芽孢杆菌ZJ186对测试的不同生理小种的稻瘟菌及水稻纹枯病原菌有拮抗作用,对测试的青霉素类、大环内脂类等抗生素敏感,对杆菌肽、复达欣、环丙氟呱酸三种抗生素有耐药性。
田间检测发现,菌株ZJ186的发酵液处理后的植株感病指数为12%,显著低于阴性对照,能较好的防治稻瘟病。通过孢子抑制实验进一步证明,拮抗菌ZJ186的发酵液可以抑制孢子萌芽达6h,而阳性对照三环唑处理后的孢子在2h就已经开始产生萌芽。发酵液处理后的孢子芽管长度极显著低于同时期阴性和阳性对照。结果表明,菌株ZJ186的发酵液能有效的防治稻瘟病。
菌株ZJ186属于分泌型,可通过发酵获得活性物质。该菌株具有较高的遗传稳定性,在连续传代10次后,发酵液活性保持不变。ZJ186在Landy发酵培养基中抑菌活性与菌体生长状况呈极显著正相关(p<0.01),其最佳发酵时间为72h至80h。
通过对活性物质的理化性质研究,发现活性物质具有耐高温、耐酸性,但其在强碱性条件下容易失活。粗提物最小抑菌浓度为100ug/ml。
根据活性物质极性较大的特点,采用MCI反相色谱、Sephadex LH-20及高效液相色谱(HPLC)进行分离纯化,经质谱检测,其分子量为270.2。经分子量数据库的检索比对的结果推测,活性物质可能为一种新的抗生素。
Rice blast caused by fungus Magnaporthe oryzae is one of the most serious rice diseases of the world. Chemical control being as the main traditional measure is widely used in rice production to reduce the incidence of rice blast. But due to the restrictions and destructive effects, chemical control caused more and more serious social problems. Biocontrol based on secondary metabolites of microorganisms can not only control disease pathogens, but also be safe to the environment. This special advantage makes it eligible to meet the demand of modern agriculture development.
The marine environment is the largest habitat on earth. However, mainly because of their extreme living environments compared with terrestrial bacteria, the most majority of marine microbes have not yet been cultured. As the development of biotechnology, more and more marine microorganisms were explored and their secondary metabolites provide a new strategy to plant fungi diseases control.
A strain was isolated in this study which showed strong inhibition activity against Magnaporthe oryzae. The product yield was significantly increased by fermentation optimize. The active compounds were isolated and purified by chromatography methods, and the molecular mass was also determinated by MS. All the above achievements make it possible for the strain and its natural products to commercial production. The results of this study were summarized in following sections:
1. The isolation and identification of antifungal strains.
13strains which showed strong antifungal activity against Magnaporthe oryzae were isolated from marine samples. Strain ZJ387was classified into Cobetia marina by partial sequence analysis, and other12strains showed great homology to Bacillus altitudinis, Bacillus stratosphericus and Bacillus aerophilus, and speculated to be the same bacterium. ZJ186was chose as the target strain and used in following studies.
According to the morphology, physiological and chemical tests and phylogenetic tree based on the16S rDNA, the strain ZJ186was characterized as Bacillus altitudinis.
It's the first report of Bacillus altitudinis isolated from marine samples and showing antifungal activity against Magnaporthe oryzae.
2. Optimization of culture conditions.
Landy medium was chose as basic culture broth for fermentation of strain ZJ186. Orthogonal optimize with L27313array was performed in9factors and three levels, such as the optimal carbon source molasses (10,20and30g/L), the optimal nitrogen source rapeseedmeal (2.5,5and lOg/L), NaCl (0.75,1.5and3g/L), MgSO4(0.5,1and2g/L), time (36,48and72h), temperature (28,32and37℃), initial pH (5,6and7), inoculum volume (1%,2%and4%) and liquid medium volume (10%,30%and50%).
The F value of carbon source molasses, nitrogen source rapeseedmeal, NaCl, time, initial pH and liquid medium volume, respectively, were extremely significant (p<0.01) compared with error value. The F value of nitrogen source rapeseedmeal, MgSO4and temperature were significant (p<0.05), while that of inoculum volume was not significant. The best optimal combination was A2B2C3D2E2F1G2I2.
The relative inhibition of mycelium growth of optimal culture broth was67.54%, increased24.83%compared with that of basic culture broth, and improved nearly2times than that of positive control tricyclazole (0.75%).
3. The isolation, purification and analysis of active compounds.
Strain ZJ186showed antifungal activity to all tested physiological races of Magnaporthe oryzae and Rhizoctonia solani from the result of antifungal spectrum, and exhibited sensitive to penicillin antibiotics and macrolide antibiotic, and resistant to bacitracin, fortune and ring c fluorine gung acid.
The assay of field test indicated that the index of disease infection significantly lower than negative control. This result was confirmed by the experiment that the filtrate of broth culture could suppress germination of pathogen spore for6h, which significantly longer than that of both negative and positive control. The above results suggested the filtrate of culture broth of strain ZJ186might control rice blast disease.
The active compounds could be secreted, and it could be gathered by fermentation methods.
Strain ZJ186was capable of genetic stability since the activity of filtrate remained almost unchanged after10generations culture. And the antifungal activity became significant correlated with the cell growth at20h after inoculation till to the whole96h cell growth periods in Landy culture (p<0.01). The best fermentation time for strain ZJ186was72h to80h.
The active compounds were capable of resistant properties of high-temperature and strong acid, but sensitive to alkalinity. The minimal inhibition concentration (MIC) of primary extract from MCI chromatography was100ug/ml.
The active compound was isolated and purified with the chromatograph methods, such as MCI reverse phase chromatograph, Sephadex LH-20and HPLC. The molecular mass was270.2by MS detection. The active compound was speculated to be a new antibiotic from the result of Blastn in molecular database.
In summarize, the product yield of strain ZJ186was increased by optimization experiments. All the above achievements provided detail data for the further research on the Bacillus altitudinis strain ZJ186.
海洋是一个巨大的生物资源库,但由于其特殊的生存环境,大部分海洋生物资源尚未被开发。随着生物技术的发展；越来越多的海洋微生物被开发、利用,它们的次生代谢产物为生物防控提供了新的策略。
本研究从海洋样品中分离、鉴定出对稻瘟菌有拮抗作用的细菌,通过发酵优化,提高了活性物质代谢产率；采用各种色谱法对活性物质进行了分离纯化,并测定活性物质的分子量,为后续开发、利用提供了实验依据。本研究的结果有以下几点：
1.拮抗菌株的分离、鉴定
从海样样品中分离到了13株对水稻稻瘟菌有明显抑制活性的菌株,通过16SrDNA单引物序比对,发现有ZJ387与Cobetia marina有最高的同源性(99.351%),其他12株菌株与高地芽孢杆菌(Bacillus altitudinis)、同温层芽孢杆菌(Bacillus stratosphericus)及嗜气芽孢杆菌(Bacillus aerophilus)有高度同源性,推测这几种菌株可能为同一种菌。我们选择抑菌活性更强的ZJ186作为目标菌株,用于后续研究。
通过形态学观察、生理生化指标测定以及16S rDNA系统进化树分析,确定ZJ186属于高地芽孢杆菌。
本文首次从海洋材料中分离到了高地芽孢杆菌,并首次报道了高地芽孢杆菌对稻瘟菌有拮抗作用。
2.拮抗菌株ZJ186的发酵优化
通过发酵培养基的筛选实验,选用Landy作为发酵基础培养基进行优化。经单因素优化试验,确定以最佳碳源糖蜜10g/L、20g/L、30g/L,最佳氮源菜籽粉2.5g/L5g/L、10g/L, NaCl0.75g/L、1.5g/L、3g/L, MgSO40.5g/L,1g/L、2g/L,发酵时间36h、48h、72h,温度28℃、32℃、37℃,初始pH值5、6、7,接种量1%、2%、4%,装液量10%、30%、50%等9种因素各三个水平采用L27313正交表进行优化。
经显著性检验,糖蜜(碳源)、NaCl、发酵时间、初始ph值及装液量的F值均达极显著水平,菜籽粉(氮源)、MgSO4和温度的F值达到显著水平,而接种量F值不显著。最优发酵组合为：A2B2C3D2E2F1G2I2。
发酵优化后的发酵液滤液相对抑菌率为67.54%，较优化之前的54.1%,优化效率提高了24.83%,较阳性对照三环唑抑菌活性(21.64%)提高了212.06%。
通过发酵优化实验,我们提高了稻瘟菌拮抗菌株ZJ186产生抑菌活性物质的能力,为该菌株的进一步开发奠定了研究基础。
3.抗菌活性物质的分离纯化及活性成分分析
通过抗菌图谱检测,高地芽孢杆菌ZJ186对测试的不同生理小种的稻瘟菌及水稻纹枯病原菌有拮抗作用,对测试的青霉素类、大环内脂类等抗生素敏感,对杆菌肽、复达欣、环丙氟呱酸三种抗生素有耐药性。
田间检测发现,菌株ZJ186的发酵液处理后的植株感病指数为12%,显著低于阴性对照,能较好的防治稻瘟病。通过孢子抑制实验进一步证明,拮抗菌ZJ186的发酵液可以抑制孢子萌芽达6h,而阳性对照三环唑处理后的孢子在2h就已经开始产生萌芽。发酵液处理后的孢子芽管长度极显著低于同时期阴性和阳性对照。结果表明,菌株ZJ186的发酵液能有效的防治稻瘟病。
菌株ZJ186属于分泌型,可通过发酵获得活性物质。该菌株具有较高的遗传稳定性,在连续传代10次后,发酵液活性保持不变。ZJ186在Landy发酵培养基中抑菌活性与菌体生长状况呈极显著正相关(p<0.01),其最佳发酵时间为72h至80h。
通过对活性物质的理化性质研究,发现活性物质具有耐高温、耐酸性,但其在强碱性条件下容易失活。粗提物最小抑菌浓度为100ug/ml。
根据活性物质极性较大的特点,采用MCI反相色谱、Sephadex LH-20及高效液相色谱(HPLC)进行分离纯化,经质谱检测,其分子量为270.2。经分子量数据库的检索比对的结果推测,活性物质可能为一种新的抗生素。
Rice blast caused by fungus Magnaporthe oryzae is one of the most serious rice diseases of the world. Chemical control being as the main traditional measure is widely used in rice production to reduce the incidence of rice blast. But due to the restrictions and destructive effects, chemical control caused more and more serious social problems. Biocontrol based on secondary metabolites of microorganisms can not only control disease pathogens, but also be safe to the environment. This special advantage makes it eligible to meet the demand of modern agriculture development.
The marine environment is the largest habitat on earth. However, mainly because of their extreme living environments compared with terrestrial bacteria, the most majority of marine microbes have not yet been cultured. As the development of biotechnology, more and more marine microorganisms were explored and their secondary metabolites provide a new strategy to plant fungi diseases control.
A strain was isolated in this study which showed strong inhibition activity against Magnaporthe oryzae. The product yield was significantly increased by fermentation optimize. The active compounds were isolated and purified by chromatography methods, and the molecular mass was also determinated by MS. All the above achievements make it possible for the strain and its natural products to commercial production. The results of this study were summarized in following sections:
1. The isolation and identification of antifungal strains.
13strains which showed strong antifungal activity against Magnaporthe oryzae were isolated from marine samples. Strain ZJ387was classified into Cobetia marina by partial sequence analysis, and other12strains showed great homology to Bacillus altitudinis, Bacillus stratosphericus and Bacillus aerophilus, and speculated to be the same bacterium. ZJ186was chose as the target strain and used in following studies.
According to the morphology, physiological and chemical tests and phylogenetic tree based on the16S rDNA, the strain ZJ186was characterized as Bacillus altitudinis.
It's the first report of Bacillus altitudinis isolated from marine samples and showing antifungal activity against Magnaporthe oryzae.
2. Optimization of culture conditions.
Landy medium was chose as basic culture broth for fermentation of strain ZJ186. Orthogonal optimize with L27313array was performed in9factors and three levels, such as the optimal carbon source molasses (10,20and30g/L), the optimal nitrogen source rapeseedmeal (2.5,5and lOg/L), NaCl (0.75,1.5and3g/L), MgSO4(0.5,1and2g/L), time (36,48and72h), temperature (28,32and37℃), initial pH (5,6and7), inoculum volume (1%,2%and4%) and liquid medium volume (10%,30%and50%).
The F value of carbon source molasses, nitrogen source rapeseedmeal, NaCl, time, initial pH and liquid medium volume, respectively, were extremely significant (p<0.01) compared with error value. The F value of nitrogen source rapeseedmeal, MgSO4and temperature were significant (p<0.05), while that of inoculum volume was not significant. The best optimal combination was A2B2C3D2E2F1G2I2.
The relative inhibition of mycelium growth of optimal culture broth was67.54%, increased24.83%compared with that of basic culture broth, and improved nearly2times than that of positive control tricyclazole (0.75%).
3. The isolation, purification and analysis of active compounds.
Strain ZJ186showed antifungal activity to all tested physiological races of Magnaporthe oryzae and Rhizoctonia solani from the result of antifungal spectrum, and exhibited sensitive to penicillin antibiotics and macrolide antibiotic, and resistant to bacitracin, fortune and ring c fluorine gung acid.
The assay of field test indicated that the index of disease infection significantly lower than negative control. This result was confirmed by the experiment that the filtrate of broth culture could suppress germination of pathogen spore for6h, which significantly longer than that of both negative and positive control. The above results suggested the filtrate of culture broth of strain ZJ186might control rice blast disease.
The active compounds could be secreted, and it could be gathered by fermentation methods.
Strain ZJ186was capable of genetic stability since the activity of filtrate remained almost unchanged after10generations culture. And the antifungal activity became significant correlated with the cell growth at20h after inoculation till to the whole96h cell growth periods in Landy culture (p<0.01). The best fermentation time for strain ZJ186was72h to80h.
The active compounds were capable of resistant properties of high-temperature and strong acid, but sensitive to alkalinity. The minimal inhibition concentration (MIC) of primary extract from MCI chromatography was100ug/ml.
The active compound was isolated and purified with the chromatograph methods, such as MCI reverse phase chromatograph, Sephadex LH-20and HPLC. The molecular mass was270.2by MS detection. The active compound was speculated to be a new antibiotic from the result of Blastn in molecular database.
In summarize, the product yield of strain ZJ186was increased by optimization experiments. All the above achievements provided detail data for the further research on the Bacillus altitudinis strain ZJ186.
引文
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