新归圃药用植物白头翁叶斑病病原学及其致病机理研究
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摘要
白头翁(Pulsatilla spp.)为毛莨科多年生草本药用植物,具有较高的药用价值及观赏价值,随着对其药理研究的不断深入,人们对白头翁需求量不断增加,致使野生资源受到了严重破坏,因此白头翁的归圃栽培越来越受到重视,在野生种质资源归圃的过程中,发现白头翁病害呈逐年加重的趋势,其中白头翁叶斑病(windflower leaf spot)发生普遍且危害严重,迄今国内外尚未见该病的系统研究报道。因此,本文针对白头翁叶斑病菌(Ascochyta anemones Kab.et Bub.)的种类鉴定、遗传多样性、致病机理、侵染特性及该病的流行规律进行了系统研究,主要研究结果如下：
1.明确了白头翁叶斑病菌的分类地位。2010-2011年从采自沈阳农业大学百草园、抚顺市清源县英额门镇、抚顺市清原县夏家堡镇、抚顺市抚顺县马圈子乡白头翁人工栽培基地的白头翁叶斑病典型病叶进行分离,经柯赫氏法则证病,通过形态学及ITS序列分析,确定白头翁叶斑病的病原菌为银莲花壳二孢(Ascochyta anemones Kab.et Bub.)。
2.首次在自然寄主上发现了白头翁叶斑病菌(Ascochyta anemones Kab.et Bub.)的有性态,并确认其有性态为亚隔孢壳属(Didymella)新种,定名为Didymella anemonessp·nOv.。
3.首次从DNA分子水平上研究白头翁叶病斑菌遗传多样性。应用RAPD及ISSR两种分子标记手段,以采自辽宁省不同生态区的白头翁叶病斑菌菌株为研究对象,筛选出11条RAPD引物和8条ISSR引物,分别扩增出108条和95条谱带。采用NTSYS软件进行病原菌的聚类分析,结果表明：RAPD标记的30株供试菌株遗传相似系数在0.56～0.98之间,当遗传相似系数为0.622时,供试菌株被划分为4个类群；ISSR标记的30个供试菌株相似系数在0.58～1之间,当遗传相似系数为0.635时,供试菌株被划分为4个类群。RAPD及ISSR分析结果均表明辽宁省白头翁叶斑病菌具有丰富的遗传多样性,聚类组群与菌株的地理来源有一定的相关性,不同寄主来源的菌株之间存在明显的遗传差异。
4.首次明确了白头翁叶斑病菌的致病机理。白头翁叶斑病菌能够产生一系列细胞壁降解酶,但其产生条件存在明显差异。静置培养利于各种细胞壁降解酶的产生；PG和PMG在培养6d时活性达到高峰,PGTE、PMTE、Cx和p-葡萄糖苷酶在培养12d时活性最高；PG和PMG产生的最适温度为25℃, PGTE、 PMTE、Cx和p-葡萄糖苷酶产生的最适温度为30℃；PG、PMG的活性在pH为5时最高。利用酸碱沉淀法提取了白头翁叶斑病菌黑色素,采用红外光谱研究了黑色素的性质。通过对黑色素的基本性质及其在致病性中的作用进行研究,初步证明白头翁叶斑病菌黑色素为DHN黑色素。采用RACE技术,获得白头翁叶斑病菌(Ascochyta anemones)黑色素合成过程中的聚酮合成酶AaPKS基因的cDNA序列,其全长为6278bp,(GenBank登录号为KC190500),最大开放阅读框(ORF)长5853bp,编码1950个氨基酸。预测该蛋白是由35.38%的a-螺旋(alpha helix)、14.21%的延伸链(extended strand)、4.15%的p-转角(beta turn)和46.26%的无规则卷曲(random coil)组成。序列比对和同源性分析表明,AaPKS基因与鹰嘴豆褐斑病菌(Ascochyta rabiei) PKS基因的氨基酸同源性最高,达95%。
5.系统研究了白头翁叶斑病菌的侵染特性。研究结果表明：接种4h,病原菌即可萌发；接种12-24h,芽管不断伸长并在末端形成附着胞,附着胞大多在细胞间隙产生并直接侵入寄主角质层;一旦病原菌成功定殖,寄主组织便迅速瓦解；病原菌接种3-6d后,即可显症。寄主范围测定结果表明,白头翁叶斑病菌对草莓、苹果、番茄和辣椒均有致病性,其中对草莓致病性最强,发病率最高为80%。
6.初步构建了白头翁叶斑病时间流行动态模型。2010-2011年对白头翁叶斑病病害发生情况进行了系统调查,应用SPSS软件进行回归分析,确认Logistic模型为最佳模型,并推导得出沈阳地区白头翁叶斑病指数增长期为5月末到6月中旬；逻辑斯蒂期为6月中旬到8月中旬；衰退期为8月中旬到白头翁生育后期。
7.初步进行了白头翁叶斑病药剂防治研究。选用7种药剂对白头翁叶斑病菌进行室内药剂筛选,结果表明：25%嘧菌酯SC和40%氟硅唑EC对白头翁叶斑病菌的抑菌效果最好,EC5o值分别为3.1741mg/L和9.3088mg/Lo田间药剂试验结果表明：50%腐霉利可湿性粉剂对白头翁叶斑病的防效最高,达80.0%；40%嘧霉胺悬浮剂次之,防效可达78.0%;25%嘧菌酯悬浮剂对该病也有较好的防治效果。
Pulsatilla spp. is one of the most important Chinese traditional medical plant in Liaoning Province with upper economic and medical value., With the adjustment of Agricultural Structure in Liaoning province, the wild windflower of artificial cultivation caused by more attention by people. We found that disease of windflower became more and more serious, windflower leaf spot is one of the most serious diseases. The disease caused by Ascochyta anemones Kab.et Bub. and the system research has not been reported at home and abroad. For this reason, pathogen identification, the genetic differentiation, pathogenic mechanism, infection characteristics and epidemic regulation of Ascochyta anemones were systematically studied. The results were as follows:
1. Identification of pathology of windflower leaf spot. The samples which were collected from planting areas of windflower in Shenyang Agriculture University herbs garden, Yingemen Town of Qingyuan County, Xiajiapu Town of Qingyuan County from2010to2011were isolated. Based on Koch's Rule and ITS sequence analysis, it is determined that the windflower leaf spot is caused by the fungi of Ascochyta anemones.
2. The teleomorph of Ascochyta anemones has been discovered for the first time on overwintering windflower. Large numbers of documents indicated that the teleomorph is an undescribed species of Didymella.
3. Genetic diversity of Ascochyta anemones of windflower leaf spot was analyzed from DNA molecular level for the first time, which showed rich polymorphic loci. RAPD and ISSR were used to explore genetic diversity of Ascochyta anemones from different localities in Liaoning,11RAPD primers and8ISSR primers were screened obtained108and95polymorphic DNA bands respectively. The result of NTS YS cluster analysis showed that the genetic similarity coefficient of RAPD among the30Ascochyta isolates was from0.56-0.98and these strains were divided into4genetic groups when the similarity coefficient was0.622. The result also showed that the genetic similarity coefficient of ISSR among these isolates was from0.58-1and these strains were divided into4genetic groups when the similarity coefficient was0.635. The analysis results of RAPD and ISSR suggested that the pathogens of windflower leaf spot in Liaoning province had rich genetic diversity. There existed definite correations between the clustering groups and their geographic distribution, and an obvious genetic difference among the isolates from different host sources.
4. The study defined the pathogenic mechanism of Ascochyta anemones preliminary. A list of C WDEs as PG, PMG, PGTE, PMTE, Cx and β-glucosidase were found in A anemones, but the producing enzyme conditions of them were different significantly. Stationary culture made more for the production of cell-wall degradation compared to the shake culture. The activities of PG and PMG reached a peak when cultured6d, while PGTE, PMTE, Cx and β-glucosidase reached a peak when cultured12d.èPG and PMG showed maximum activities at25℃. PGTE, PMTE,(3-glucosidase and Cx showed maximum activities at30℃. PG and PMG showed maximum activities at culture pH5. Acidity alkali sedimentation method was used to extract the melanin from A. anemones. The basic characteristic of melanin and its effect of pathogenicity were studied, suggesting that the melnin was a DHN melanin. A novel polyketide synthase gene, named AaPKS (GenBank accession number KC190500), was cloned from A.anemones using RACE method. The full length cDNA of AaPKS consists of6278bp, with a5853bp ORF encoding1950amino acid residues. The predicted secondary structure composition for the protein contained about35.38%alpha helix,14.21%extended strand,4.15%beta turn and46.26%random coil. Phylogenetic analysis indicated that AaPKS of A.anemones shares95%identity to Ascochyta rabiei.
5. Study on the infection characteristics of Ascochyta anemones of windflower leaf spot and the primary source of infection of windflower were fist define. Histological studies of the artificially inoculated leaf tissues with Ascochyta anemones conidia were observed by light and electron microscopy. Conidial germination and germ tube foramtion began after4hours of inoculation. Each conidium produced one to germ tubes to peneatrate the host surface. The fungus gain entry of the host by direct penetration of the leaf cuticle, following the formation of appressorium after12-24h of inoculation. Most appressoria were formed in the grooves between adjacent epidermal cells. Once the fungus was fully established, it destroyed internal tissues, resulting in diseased lesions on the leaves after3-6days of inoculation. The test result of the host range of Ascochyta anemones demonstrated that the pathogen could infect strawberry, apple, tomatoes and peppers. In addition, strawberry had the hightest disease rate80%.
6. The temporal dynamic models of windflower leaf spot were first established. According to systematic investigation during2010-2011, making regression analysis via SPSS software, logistic model was comfirmed as the best model. The model deduced that exponential phase of windflower leaf spot was from late May to mid June, the logistic phase was from mid June to mid August, the decline phase was mid August to the end of windflower grow stage in Shenyang.
7. Studies on primary management of windflower leaf spot. The fungistatic action of seven fungicides to A.anemones was determined. Fungicide tests proved that azoxystrobin25%SC and flusilazole40%EC exhibited the best inhibitory effect on the colony growth which its EC50values reached3.1741mg/L and9.3088mg/L respectively. Field trials showed that the disease could be effectively controlled by using the procymidone50%WP which control effect reached80%, pyrimethanil40%SC and azoxystrobin25%SC also have better control effect.
1.明确了白头翁叶斑病菌的分类地位。2010-2011年从采自沈阳农业大学百草园、抚顺市清源县英额门镇、抚顺市清原县夏家堡镇、抚顺市抚顺县马圈子乡白头翁人工栽培基地的白头翁叶斑病典型病叶进行分离,经柯赫氏法则证病,通过形态学及ITS序列分析,确定白头翁叶斑病的病原菌为银莲花壳二孢(Ascochyta anemones Kab.et Bub.)。
2.首次在自然寄主上发现了白头翁叶斑病菌(Ascochyta anemones Kab.et Bub.)的有性态,并确认其有性态为亚隔孢壳属(Didymella)新种,定名为Didymella anemonessp·nOv.。
3.首次从DNA分子水平上研究白头翁叶病斑菌遗传多样性。应用RAPD及ISSR两种分子标记手段,以采自辽宁省不同生态区的白头翁叶病斑菌菌株为研究对象,筛选出11条RAPD引物和8条ISSR引物,分别扩增出108条和95条谱带。采用NTSYS软件进行病原菌的聚类分析,结果表明：RAPD标记的30株供试菌株遗传相似系数在0.56～0.98之间,当遗传相似系数为0.622时,供试菌株被划分为4个类群；ISSR标记的30个供试菌株相似系数在0.58～1之间,当遗传相似系数为0.635时,供试菌株被划分为4个类群。RAPD及ISSR分析结果均表明辽宁省白头翁叶斑病菌具有丰富的遗传多样性,聚类组群与菌株的地理来源有一定的相关性,不同寄主来源的菌株之间存在明显的遗传差异。
4.首次明确了白头翁叶斑病菌的致病机理。白头翁叶斑病菌能够产生一系列细胞壁降解酶,但其产生条件存在明显差异。静置培养利于各种细胞壁降解酶的产生；PG和PMG在培养6d时活性达到高峰,PGTE、PMTE、Cx和p-葡萄糖苷酶在培养12d时活性最高；PG和PMG产生的最适温度为25℃, PGTE、 PMTE、Cx和p-葡萄糖苷酶产生的最适温度为30℃；PG、PMG的活性在pH为5时最高。利用酸碱沉淀法提取了白头翁叶斑病菌黑色素,采用红外光谱研究了黑色素的性质。通过对黑色素的基本性质及其在致病性中的作用进行研究,初步证明白头翁叶斑病菌黑色素为DHN黑色素。采用RACE技术,获得白头翁叶斑病菌(Ascochyta anemones)黑色素合成过程中的聚酮合成酶AaPKS基因的cDNA序列,其全长为6278bp,(GenBank登录号为KC190500),最大开放阅读框(ORF)长5853bp,编码1950个氨基酸。预测该蛋白是由35.38%的a-螺旋(alpha helix)、14.21%的延伸链(extended strand)、4.15%的p-转角(beta turn)和46.26%的无规则卷曲(random coil)组成。序列比对和同源性分析表明,AaPKS基因与鹰嘴豆褐斑病菌(Ascochyta rabiei) PKS基因的氨基酸同源性最高,达95%。
5.系统研究了白头翁叶斑病菌的侵染特性。研究结果表明：接种4h,病原菌即可萌发；接种12-24h,芽管不断伸长并在末端形成附着胞,附着胞大多在细胞间隙产生并直接侵入寄主角质层;一旦病原菌成功定殖,寄主组织便迅速瓦解；病原菌接种3-6d后,即可显症。寄主范围测定结果表明,白头翁叶斑病菌对草莓、苹果、番茄和辣椒均有致病性,其中对草莓致病性最强,发病率最高为80%。
6.初步构建了白头翁叶斑病时间流行动态模型。2010-2011年对白头翁叶斑病病害发生情况进行了系统调查,应用SPSS软件进行回归分析,确认Logistic模型为最佳模型,并推导得出沈阳地区白头翁叶斑病指数增长期为5月末到6月中旬；逻辑斯蒂期为6月中旬到8月中旬；衰退期为8月中旬到白头翁生育后期。
7.初步进行了白头翁叶斑病药剂防治研究。选用7种药剂对白头翁叶斑病菌进行室内药剂筛选,结果表明：25%嘧菌酯SC和40%氟硅唑EC对白头翁叶斑病菌的抑菌效果最好,EC5o值分别为3.1741mg/L和9.3088mg/Lo田间药剂试验结果表明：50%腐霉利可湿性粉剂对白头翁叶斑病的防效最高,达80.0%；40%嘧霉胺悬浮剂次之,防效可达78.0%;25%嘧菌酯悬浮剂对该病也有较好的防治效果。
Pulsatilla spp. is one of the most important Chinese traditional medical plant in Liaoning Province with upper economic and medical value., With the adjustment of Agricultural Structure in Liaoning province, the wild windflower of artificial cultivation caused by more attention by people. We found that disease of windflower became more and more serious, windflower leaf spot is one of the most serious diseases. The disease caused by Ascochyta anemones Kab.et Bub. and the system research has not been reported at home and abroad. For this reason, pathogen identification, the genetic differentiation, pathogenic mechanism, infection characteristics and epidemic regulation of Ascochyta anemones were systematically studied. The results were as follows:
1. Identification of pathology of windflower leaf spot. The samples which were collected from planting areas of windflower in Shenyang Agriculture University herbs garden, Yingemen Town of Qingyuan County, Xiajiapu Town of Qingyuan County from2010to2011were isolated. Based on Koch's Rule and ITS sequence analysis, it is determined that the windflower leaf spot is caused by the fungi of Ascochyta anemones.
2. The teleomorph of Ascochyta anemones has been discovered for the first time on overwintering windflower. Large numbers of documents indicated that the teleomorph is an undescribed species of Didymella.
3. Genetic diversity of Ascochyta anemones of windflower leaf spot was analyzed from DNA molecular level for the first time, which showed rich polymorphic loci. RAPD and ISSR were used to explore genetic diversity of Ascochyta anemones from different localities in Liaoning,11RAPD primers and8ISSR primers were screened obtained108and95polymorphic DNA bands respectively. The result of NTS YS cluster analysis showed that the genetic similarity coefficient of RAPD among the30Ascochyta isolates was from0.56-0.98and these strains were divided into4genetic groups when the similarity coefficient was0.622. The result also showed that the genetic similarity coefficient of ISSR among these isolates was from0.58-1and these strains were divided into4genetic groups when the similarity coefficient was0.635. The analysis results of RAPD and ISSR suggested that the pathogens of windflower leaf spot in Liaoning province had rich genetic diversity. There existed definite correations between the clustering groups and their geographic distribution, and an obvious genetic difference among the isolates from different host sources.
4. The study defined the pathogenic mechanism of Ascochyta anemones preliminary. A list of C WDEs as PG, PMG, PGTE, PMTE, Cx and β-glucosidase were found in A anemones, but the producing enzyme conditions of them were different significantly. Stationary culture made more for the production of cell-wall degradation compared to the shake culture. The activities of PG and PMG reached a peak when cultured6d, while PGTE, PMTE, Cx and β-glucosidase reached a peak when cultured12d.èPG and PMG showed maximum activities at25℃. PGTE, PMTE,(3-glucosidase and Cx showed maximum activities at30℃. PG and PMG showed maximum activities at culture pH5. Acidity alkali sedimentation method was used to extract the melanin from A. anemones. The basic characteristic of melanin and its effect of pathogenicity were studied, suggesting that the melnin was a DHN melanin. A novel polyketide synthase gene, named AaPKS (GenBank accession number KC190500), was cloned from A.anemones using RACE method. The full length cDNA of AaPKS consists of6278bp, with a5853bp ORF encoding1950amino acid residues. The predicted secondary structure composition for the protein contained about35.38%alpha helix,14.21%extended strand,4.15%beta turn and46.26%random coil. Phylogenetic analysis indicated that AaPKS of A.anemones shares95%identity to Ascochyta rabiei.
5. Study on the infection characteristics of Ascochyta anemones of windflower leaf spot and the primary source of infection of windflower were fist define. Histological studies of the artificially inoculated leaf tissues with Ascochyta anemones conidia were observed by light and electron microscopy. Conidial germination and germ tube foramtion began after4hours of inoculation. Each conidium produced one to germ tubes to peneatrate the host surface. The fungus gain entry of the host by direct penetration of the leaf cuticle, following the formation of appressorium after12-24h of inoculation. Most appressoria were formed in the grooves between adjacent epidermal cells. Once the fungus was fully established, it destroyed internal tissues, resulting in diseased lesions on the leaves after3-6days of inoculation. The test result of the host range of Ascochyta anemones demonstrated that the pathogen could infect strawberry, apple, tomatoes and peppers. In addition, strawberry had the hightest disease rate80%.
6. The temporal dynamic models of windflower leaf spot were first established. According to systematic investigation during2010-2011, making regression analysis via SPSS software, logistic model was comfirmed as the best model. The model deduced that exponential phase of windflower leaf spot was from late May to mid June, the logistic phase was from mid June to mid August, the decline phase was mid August to the end of windflower grow stage in Shenyang.
7. Studies on primary management of windflower leaf spot. The fungistatic action of seven fungicides to A.anemones was determined. Fungicide tests proved that azoxystrobin25%SC and flusilazole40%EC exhibited the best inhibitory effect on the colony growth which its EC50values reached3.1741mg/L and9.3088mg/L respectively. Field trials showed that the disease could be effectively controlled by using the procymidone50%WP which control effect reached80%, pyrimethanil40%SC and azoxystrobin25%SC also have better control effect.
引文
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