辣椒疫病生防ASD菌株筛选及对土壤微生态调控机理研究
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摘要
辣椒疫病(Phytophthora capsici)是保护地辣椒生产上造成产量损失极大的土传病害。由于连作、大水漫灌和化学药剂频繁使用,常导致保护地土壤中该病菌菌源积累增大、农药污染加重,微生态环境易遭到破坏。目前生产上缺少生防菌剂产品,缺乏有效的生物防治措施。本文以辽宁省辣椒疫病为研究对象,开展了辣椒疫霉病菌遗传多样性、生防菌和微生态调控研究,取得如下结果：
1.通过形态学和rDNA-ITS同源性分析表明,来源于辽宁省不同地区辣椒疫病病菌形态学相似,菌丝多为直角或锐角分枝,孢子囊梗无色、丝状,孢子囊单孢、卵圆形或梨形,乳突单生；病菌rDNA-ITS序列805-877bp。聚类分析表明,辽宁省不同地区的辣椒疫病菌表现出较高的同源性,无区域性差异。
2.通过SRAP技术对病菌遗传多样性分析表明,不同地区辣椒疫病病菌的群体遗传相似性很高,具出丰富的多态性,但区域性差异不明显；致病力测定表明,辽宁省辣椒疫病菌存在11种致病力类型,主要以中等致病力为主,致病力增强风险较大。辣椒疫病菌致病力与遗传相似性无明显相关性。
3.采用平板稀释法分离辣椒生产田56份土样,对获得的微生物进行对峙抑菌试验,得到对辣椒疫病菌有明显的拮抗作用的ASD菌株,其抑菌带达11.7mm。盆栽试验防治效果可达75.12%,优于80%代森锰锌可湿性粉剂300mg/L、50%烯酰吗啉水分散粒剂20mg/L。
4.形态学及分子鉴定结果表明,ASD菌株在PDA培养基上菌落丝绒状,菌丝初为白色至土黄色,偶有透明液渗出。分生孢子顶囊为辐射形,分生孢子球形,有小梗及链状排列的分生孢子；测定ASD菌株rDNA-ITS序列为621bp,与黄柄曲霉序列相似度达97%,确认ASD菌株为黄柄曲霉菌(Aspergillus flavipes)。
5.控病机理研究表明,生防菌ASD菌株可产生抑菌物质,可致病菌菌丝畸形,抑制病菌生长。加入疫病菌丝可诱导CMC酶活性、木聚糖酶活性显著提高,且随着底物浓度的增加酶活性升高,抑制疫病菌生长的主要因素是诱导产生的CMC酶、木聚糖酶,而与B-1,3葡聚糖酶和蛋白酶关系不大。
6.发酵条件优化结果表明,诱导ASD菌产生抑菌物质由高到低的碳源顺序为单糖、双糖和多糖,最佳碳源为葡萄糖；有机氮明显优于无机氮,最佳有机氮和无机氮分别为甘氨酸、硝酸铵；KCl、MgSO4·7H2O、FeSO4等盐类在低浓度下有利于抑菌物质产生结合正交试验确定生防菌ASD菌株产生抑菌物质的环境最佳发酵配方及条件是2%葡萄糖、0.1%甘氨酸、0.003%FeSO4、0.01%KCl、0.01%MgSO4·7H20,pH6,180rpm,30℃下,装液量30mL/100mL,培养5d。
7.微生态区系分析表明,采集不同地块不同年限病株与健株的根际土壤细菌、放线菌及真菌数量变化规律不明显。同一地块健株根际土壤细菌与放线菌数量比病株多,而真菌数量比病株少；病株土壤较健株土壤的细菌/真菌、放线菌/真菌、细菌/放线菌值高。
8.经对分离出的土壤优势微生物形态学观察、生理生化试验及16srDNA序列分析,鉴定出芽孢杆菌7种,占全部细菌种类43.75%；土壤放线菌17株。B33、B40、B22和B44菌株只鉴定到属,有待与进一步鉴定。
9.ASD菌株对土壤微生态区系影响研究表明,施用生防菌后,健株与病株根际土壤中细菌和放线菌数量均有所减少,真菌数量增加。随着生防菌ASD菌株定殖时间增加,健株根际土壤细菌数量逐渐增加,病株根际土壤细菌数量基本不变,而健株和病株土壤放线菌数量变化不大,健株和病株土壤真菌数量逐渐下降。
10.对土壤酶影响研究表明,施用生防菌ASD菌株后,健株与病株根际土壤脲酶活性、酸性磷酸酶活性、蔗糖酶活性及脱氢酶活性均有不同程度的提高,只有过氧化氢酶活性提高不显著；随着生防菌ASD菌病定殖时间增加,不同酶活性变化趋势基本一致,即增加后下降,但均高于对照；施用ASD菌株后影响各种土壤酶活性大小的顺序依次为脲酶、脱氢酶、蔗糖酶、酸性磷酸酶和过氧化氢酶,可以改良土壤环境。
Phytophthora capsici Leonian is one of soil borne diseases, which is prevalently occur and seriously reduce pepper yield in conservation field. As a result of continuously cropping, flood irrigation and chemical abuse, the accumulation of pathogen source is increasing, the problem of pesticide pollution is aggravating and the micro ecology environment is liable to be broken. At present, the production of biocontrol agent is scarce and the effective biocontrol technology is absent. In the present study, the researches on genetic polymorphism, biocontrol strain and micro ecology biocontrol are carried out for pepper Phytophthora blight. The results are as follows:
1. The analysis of morphology and rDNA-ITS homology for Phytophthora capsici showed:the various strains collected from different regions of Liaoning province are similar on the configuration. Most of mycelial branches are right or acute angle. The sporangiophore is colorless and filamentous. The sporangium is single, ovoid or pyriform, and the mastoid is single. The sequences of rDNA-ITS change from805bp to877bp. Clustering analysis show that those strains collected from different regions are obvious homology, and there are not regional characteristics.
2. Using Sequence Related Amplified Polymorphism (SRAP) technology, the genetic polymorphism of Phytophthora capsici Leonian collected from different regions of Liaoning province is evaluated. The results show that the genetic similarity of Phytophthora capsici Leonian is very high and the strains have abundant polymorphism, while the regional diversity is not significant. The pathogenicity determination show that these strains are divided11pathogenic types, which of the intermediate type is primary. There are not significant correlations between the genetic similarity and pathogenicity.
3. Fifty-six samples collected from different fields are separated by plate dilution method. The ASD strain is screened and has significant resistant to Phytophthora capsici Leonian by bacteriostatic test. The width of inhibition zone is11.7mm. The control efficiency of ASD strain reaches to75.12%against Phytophthora capsici in pot experiment, which is better than80%WP Mancozeb300mg/L and50%alkene acyl complexes of water dispersible granules20mg/L
4. The morphology and molecular determinations show that the ASD strain is velvet on PDA medium. The early myceliums vary from white to khaki, occasionally exude transparent liquid. The conidium is spherical, having small stems and arraying as catenulate type, which of top sporangium is radiate. The rDNA-ITS sequence of ASD strain is621bp, which of97%is similar to Aspergillus flavipes. According to the above result, the ASD strain is determined as Aspergillus flavipes.
5. The study of resistance mechanism shows that the biological control ASD strain can produce antibacterial substances, which can cause mycelia abnormality and inhibit the pathogen growth. The activities of CMC and xylem enzymes are significant improved by the biological control ASD. With the increase of substrate concentration, the enzyme activities are improved. The main factors inhibiting the growth of Phytophthora are CMC and xylanase enzymes, while there is little relation with β-1,3glucanase and protease.
6. The results of optimizing fermentation condition show that the order of carbon source induced ASD strain to produce antibacterial substances is monosaccharide, disaccharides and polysaccharides. The glucose is the most optimize carbon source. The organic nitrogen is significantly superior to the inorganic nitrogen. The best organic and inorganic nitrogen are glycine and ammonium nitrate, respectively. KCl, MgSO4·7H2O, FeSO4are benefit to produce the antibacterial substances when the concentrations are low. Combining with orthogonal test, the optimal condition is2%Glucose,0.1%Glycine,0.003%FeSO4,0.01%KCl, MgSO4·7H2O, pH6,180rpm,30℃, the Volume30mL/100mL, culturing for5days.
7. The analysis of micro ecological system shows that the varieties of numbers of bacteria, actinomycetes and fungus are not significant, which collected from capsicum rooting zone soil of the healthy and infected plants in different regions and various years,. The numbers of bacteria and actinomycetes in capsicum rooting zone soil of healthy plant are more than those in rooting zone soil of infected plants, while the number of fungus is lower. The scales of bacteria/actinomycetes, fungus/actinomycetes and bacteria/fungus in rooting zone soil of infected plants are higher than those in rooting zone soil of healthy plants.
8. By the morphological observation, physiological and biochemical test and16s rDNA sequence analysis, seven types of dominent bacillus are identified, which is43.75%of total bacteria. There are17soil funguses. The strains of B33、B40、B22and B44are only determined to genus level. The further study should be carried out in the future.
9. Research on the effect of micro ecological system in soil, the results show that the biocontrol strain ASD could significantly reduce the quantity of the bacteria and actinomycetes, and increase the number of the fungi in capsicum rooting zone soil of the healthy and infected plants. With the colonization time increase of the biocontrol strain ASD, the number of the bacteria is gradually increased in capsicum rooting zone soil of healthy plants, while it is not changed in the capsicum rooting zone soil of infected plants. The number of the actinomycetes is not changed in capsicum rooting zone soil of the healthy and infected plants, while the number of the fungi is gradually reduced.
10. Research on the effect of soil enzyme, the activities of urease, acid phosphatase, sucrose enzyme and dehydrogenase are significantly increased by the biocontrol strain ASD in capsicum rooting zone soil of the healthy and infected plants, while the activity of catalase is not significant increased. With the colonization time increase of the biocontrol strain ASD, the activities of various enzymes are higher than the control and their variation trends are very similar, which firstly increased and decreased afterward. The responses of soil enzyme activities affected by the biocontrol strain ASD from high to low are as follows:urease, dehydrogenase, sucrose enzyme, acid phosphatase and catalase. The results showed that the biocontrol strain ASD could improve the micro ecological systems in soil.
1.通过形态学和rDNA-ITS同源性分析表明,来源于辽宁省不同地区辣椒疫病病菌形态学相似,菌丝多为直角或锐角分枝,孢子囊梗无色、丝状,孢子囊单孢、卵圆形或梨形,乳突单生；病菌rDNA-ITS序列805-877bp。聚类分析表明,辽宁省不同地区的辣椒疫病菌表现出较高的同源性,无区域性差异。
2.通过SRAP技术对病菌遗传多样性分析表明,不同地区辣椒疫病病菌的群体遗传相似性很高,具出丰富的多态性,但区域性差异不明显；致病力测定表明,辽宁省辣椒疫病菌存在11种致病力类型,主要以中等致病力为主,致病力增强风险较大。辣椒疫病菌致病力与遗传相似性无明显相关性。
3.采用平板稀释法分离辣椒生产田56份土样,对获得的微生物进行对峙抑菌试验,得到对辣椒疫病菌有明显的拮抗作用的ASD菌株,其抑菌带达11.7mm。盆栽试验防治效果可达75.12%,优于80%代森锰锌可湿性粉剂300mg/L、50%烯酰吗啉水分散粒剂20mg/L。
4.形态学及分子鉴定结果表明,ASD菌株在PDA培养基上菌落丝绒状,菌丝初为白色至土黄色,偶有透明液渗出。分生孢子顶囊为辐射形,分生孢子球形,有小梗及链状排列的分生孢子；测定ASD菌株rDNA-ITS序列为621bp,与黄柄曲霉序列相似度达97%,确认ASD菌株为黄柄曲霉菌(Aspergillus flavipes)。
5.控病机理研究表明,生防菌ASD菌株可产生抑菌物质,可致病菌菌丝畸形,抑制病菌生长。加入疫病菌丝可诱导CMC酶活性、木聚糖酶活性显著提高,且随着底物浓度的增加酶活性升高,抑制疫病菌生长的主要因素是诱导产生的CMC酶、木聚糖酶,而与B-1,3葡聚糖酶和蛋白酶关系不大。
6.发酵条件优化结果表明,诱导ASD菌产生抑菌物质由高到低的碳源顺序为单糖、双糖和多糖,最佳碳源为葡萄糖；有机氮明显优于无机氮,最佳有机氮和无机氮分别为甘氨酸、硝酸铵；KCl、MgSO4·7H2O、FeSO4等盐类在低浓度下有利于抑菌物质产生结合正交试验确定生防菌ASD菌株产生抑菌物质的环境最佳发酵配方及条件是2%葡萄糖、0.1%甘氨酸、0.003%FeSO4、0.01%KCl、0.01%MgSO4·7H20,pH6,180rpm,30℃下,装液量30mL/100mL,培养5d。
7.微生态区系分析表明,采集不同地块不同年限病株与健株的根际土壤细菌、放线菌及真菌数量变化规律不明显。同一地块健株根际土壤细菌与放线菌数量比病株多,而真菌数量比病株少；病株土壤较健株土壤的细菌/真菌、放线菌/真菌、细菌/放线菌值高。
8.经对分离出的土壤优势微生物形态学观察、生理生化试验及16srDNA序列分析,鉴定出芽孢杆菌7种,占全部细菌种类43.75%；土壤放线菌17株。B33、B40、B22和B44菌株只鉴定到属,有待与进一步鉴定。
9.ASD菌株对土壤微生态区系影响研究表明,施用生防菌后,健株与病株根际土壤中细菌和放线菌数量均有所减少,真菌数量增加。随着生防菌ASD菌株定殖时间增加,健株根际土壤细菌数量逐渐增加,病株根际土壤细菌数量基本不变,而健株和病株土壤放线菌数量变化不大,健株和病株土壤真菌数量逐渐下降。
10.对土壤酶影响研究表明,施用生防菌ASD菌株后,健株与病株根际土壤脲酶活性、酸性磷酸酶活性、蔗糖酶活性及脱氢酶活性均有不同程度的提高,只有过氧化氢酶活性提高不显著；随着生防菌ASD菌病定殖时间增加,不同酶活性变化趋势基本一致,即增加后下降,但均高于对照；施用ASD菌株后影响各种土壤酶活性大小的顺序依次为脲酶、脱氢酶、蔗糖酶、酸性磷酸酶和过氧化氢酶,可以改良土壤环境。
Phytophthora capsici Leonian is one of soil borne diseases, which is prevalently occur and seriously reduce pepper yield in conservation field. As a result of continuously cropping, flood irrigation and chemical abuse, the accumulation of pathogen source is increasing, the problem of pesticide pollution is aggravating and the micro ecology environment is liable to be broken. At present, the production of biocontrol agent is scarce and the effective biocontrol technology is absent. In the present study, the researches on genetic polymorphism, biocontrol strain and micro ecology biocontrol are carried out for pepper Phytophthora blight. The results are as follows:
1. The analysis of morphology and rDNA-ITS homology for Phytophthora capsici showed:the various strains collected from different regions of Liaoning province are similar on the configuration. Most of mycelial branches are right or acute angle. The sporangiophore is colorless and filamentous. The sporangium is single, ovoid or pyriform, and the mastoid is single. The sequences of rDNA-ITS change from805bp to877bp. Clustering analysis show that those strains collected from different regions are obvious homology, and there are not regional characteristics.
2. Using Sequence Related Amplified Polymorphism (SRAP) technology, the genetic polymorphism of Phytophthora capsici Leonian collected from different regions of Liaoning province is evaluated. The results show that the genetic similarity of Phytophthora capsici Leonian is very high and the strains have abundant polymorphism, while the regional diversity is not significant. The pathogenicity determination show that these strains are divided11pathogenic types, which of the intermediate type is primary. There are not significant correlations between the genetic similarity and pathogenicity.
3. Fifty-six samples collected from different fields are separated by plate dilution method. The ASD strain is screened and has significant resistant to Phytophthora capsici Leonian by bacteriostatic test. The width of inhibition zone is11.7mm. The control efficiency of ASD strain reaches to75.12%against Phytophthora capsici in pot experiment, which is better than80%WP Mancozeb300mg/L and50%alkene acyl complexes of water dispersible granules20mg/L
4. The morphology and molecular determinations show that the ASD strain is velvet on PDA medium. The early myceliums vary from white to khaki, occasionally exude transparent liquid. The conidium is spherical, having small stems and arraying as catenulate type, which of top sporangium is radiate. The rDNA-ITS sequence of ASD strain is621bp, which of97%is similar to Aspergillus flavipes. According to the above result, the ASD strain is determined as Aspergillus flavipes.
5. The study of resistance mechanism shows that the biological control ASD strain can produce antibacterial substances, which can cause mycelia abnormality and inhibit the pathogen growth. The activities of CMC and xylem enzymes are significant improved by the biological control ASD. With the increase of substrate concentration, the enzyme activities are improved. The main factors inhibiting the growth of Phytophthora are CMC and xylanase enzymes, while there is little relation with β-1,3glucanase and protease.
6. The results of optimizing fermentation condition show that the order of carbon source induced ASD strain to produce antibacterial substances is monosaccharide, disaccharides and polysaccharides. The glucose is the most optimize carbon source. The organic nitrogen is significantly superior to the inorganic nitrogen. The best organic and inorganic nitrogen are glycine and ammonium nitrate, respectively. KCl, MgSO4·7H2O, FeSO4are benefit to produce the antibacterial substances when the concentrations are low. Combining with orthogonal test, the optimal condition is2%Glucose,0.1%Glycine,0.003%FeSO4,0.01%KCl, MgSO4·7H2O, pH6,180rpm,30℃, the Volume30mL/100mL, culturing for5days.
7. The analysis of micro ecological system shows that the varieties of numbers of bacteria, actinomycetes and fungus are not significant, which collected from capsicum rooting zone soil of the healthy and infected plants in different regions and various years,. The numbers of bacteria and actinomycetes in capsicum rooting zone soil of healthy plant are more than those in rooting zone soil of infected plants, while the number of fungus is lower. The scales of bacteria/actinomycetes, fungus/actinomycetes and bacteria/fungus in rooting zone soil of infected plants are higher than those in rooting zone soil of healthy plants.
8. By the morphological observation, physiological and biochemical test and16s rDNA sequence analysis, seven types of dominent bacillus are identified, which is43.75%of total bacteria. There are17soil funguses. The strains of B33、B40、B22and B44are only determined to genus level. The further study should be carried out in the future.
9. Research on the effect of micro ecological system in soil, the results show that the biocontrol strain ASD could significantly reduce the quantity of the bacteria and actinomycetes, and increase the number of the fungi in capsicum rooting zone soil of the healthy and infected plants. With the colonization time increase of the biocontrol strain ASD, the number of the bacteria is gradually increased in capsicum rooting zone soil of healthy plants, while it is not changed in the capsicum rooting zone soil of infected plants. The number of the actinomycetes is not changed in capsicum rooting zone soil of the healthy and infected plants, while the number of the fungi is gradually reduced.
10. Research on the effect of soil enzyme, the activities of urease, acid phosphatase, sucrose enzyme and dehydrogenase are significantly increased by the biocontrol strain ASD in capsicum rooting zone soil of the healthy and infected plants, while the activity of catalase is not significant increased. With the colonization time increase of the biocontrol strain ASD, the activities of various enzymes are higher than the control and their variation trends are very similar, which firstly increased and decreased afterward. The responses of soil enzyme activities affected by the biocontrol strain ASD from high to low are as follows:urease, dehydrogenase, sucrose enzyme, acid phosphatase and catalase. The results showed that the biocontrol strain ASD could improve the micro ecological systems in soil.
引文
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