苹果黑星病菌遗传多样性及其快速分子检测研究
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摘要
黑星病是苹果树上的主要病害之一,具有流行速度快、为害严重、难于防治等特点,广泛分布于世界各苹果产区,特别是对欧洲、美洲苹果产区,每年均造成严重的产量和经济损失。苹果黑星病在我国主要分布在辽宁、黑龙江、甘肃,以及陕西、新疆、山东的部分苹果产区,曾多次大流行,已给我国苹果产业造成了严重影响。为了阐明苹果黑星病菌的传播路线、遗传变异等问题,以及探究其快速检测方法,为防治该病害提供比较系统的理论依据,本文开展了苹果黑星病菌遗传多样性和分子快速检测研究,取得了以下主要结果：
1、采用SSR分子标记技术,利用筛选出的18对SSR引物对中国、英国、印度三个国家的90株苹果黑星病菌进行遗传多样性分析,构建了遗传聚类分析图。结果表明：在相似系数为0.76的水平,供试的90个菌株全部聚到一类。在相似系数为0.80的水平,90个菌株被划分为2个类群：全部的英国菌株(共18株)被分在了第Ⅰ类群；中国和印度菌株(共72株)被分在了第Ⅱ类群。而在相似系数为0.83时,第Ⅱ类群又可分为3个亚群：全部的印度菌株被分在了A亚群；全部的兴平富士(4株)、全部的红星(16株)、全部的秦冠(9株)和11%的兴平嘎啦(1株)上采集的菌株被分在B亚群；全部的旬邑嘎啦(14株)、全部的旬邑富士(9株)和89%的兴平嘎啦(8株)上采集的菌株被分在C亚群。根据结果分析得出,三个国家的苹果黑星病菌与地理来源存在一定的相关性：中国菌株与印度菌株亲缘关系较近,而与英国菌株亲缘关系较远；从中国内部菌株来看,病菌亲缘关系与地理来源相关性不大,但与寄主苹果品种有一定相关性。同时采用5对SSR引物对来自欧洲的20株分属于7个不同生理小种的苹果黑星病菌进行PCR扩增,共检测到43个等位变异,等位位点数目范围6～11个,平均为8.6个；构建遗传聚类分析图,病菌遗传相似系数变异范围为0.72～1.00,平均值为0.85；在遗传距离为0.81的水平上,将20个苹果黑星病菌株分为六大类。除第一、四大类外,其余四类每类内的菌株都分属于不同的生理小种,说明苹果黑星病菌基于SSR标记划分的类群与生理小种之间无明显相关性。
2、采用AFLP分子标记技术,利用筛选出的25对AFLP引物对来自我国陕西兴平和旬邑的46株苹果黑星病菌进行遗传多样性分析,构建了遗传聚类分析图。结果表明,在相似系数为0.68的水平,46个菌株被划分为2个类群：第Ⅰ大类由全部的旬邑富士旬邑嘎啦和兴平嘎啦采集菌株组成；第Ⅱ大类由全部的旬邑红星、旬邑秦冠、兴平红星、兴平秦冠采集菌株组成。表明陕西兴平和旬邑两地的苹果黑星病菌遗传多样性与地理来源无明显相关性,而与其寄主品种间存在一定的相关性。
3、应用通用引物ITS1和ITS4对苹果黑星病菌(V. inaequalis)、苹果斑点落叶病菌(Alternaria mali)、苹果褐斑病菌(Marssonina cororlar)和苹果白粉病菌(Podospharea leucotricha)等4种苹果树常见病害的病原菌共26个菌株进行PCR扩增和测序,得到26条ITS序列。对其进行比对筛选,获得1对片段大小为320 bp的特异性引物320A/320B。应用该特异性引物对26个苹果病原真菌供试菌株和接有苹果黑星病菌的苹果叶片组织进行PCR分子检测,结果表明：特异性引物320A/320B能够特异的检测出供试菌株中的苹果黑星病菌,同时也能从接种过苹果黑星病菌但未显症的苹果组织中检测到苹果黑星病菌的存在；对苹果黑星病菌基因组DNA检测的灵敏度为100fg/μL。
Apple scab is one of the main diseases of apples. It is characterized by fast epidemic, serious damage and difficalf to control. It is one of the most important apple diseases in Europe and America and often causes yield and economic losses. In China, apple scab distributes mainly in Liaoning province, Heilongjiang province, Gansu province, and same areas of Shaanxi province, Xinjiang province and Shandong province. It has caused serious effect on industry of apple because of large-scale epidemic. In this study, we performed research on genetic diversity and molecular rapid detection of Venturia inaequalis and obtained the results as follows:
1. Eighteen pairs of primers were selected for study on genetic diversity of 90 isolates of V. inaequalis from China, India and United kingdom by using simple sequence repeats (SSR) technique. Genetic clustering map was constructed based on these results. The results indicated that, all the 90 isolates were clustered together at the 0.76 genetic similarity level. At the 0.80 genetic similarity level, all the isolates (18) from United kingdom were clustered in groupⅠand isolates from China and India (72) were clustered into groupⅡ. At the 0.83 genetic similarity level, isolates from groupⅡwere divided into 3 subgroup:all the Indian isolates belonged to subgroup A, all the isolates from Xingping Fuji, Starking Delicious, Qinguan and 11% isolates from Xingping Gala had clustered into subgroup B and all the isolates from Xunyi Gala, Xunyi fuji and 89% isolates from Xingping Gala were clustered into subgroup C. We can conclude that the isolates of V. inaequalis from three countries were relative relationship based on the regional source. The isolates from China were relatively close to that from India and far from the UK. All the isolates from China were not relatively close based on the source but had a relative relationship on cultivars. Five pairs of primers were used in PCR amplification of 20 isolates of V. inaequalis, which belong to 7 physiological races, through SSR technique. Fourty three alleles were detected with an average of 8.6 alleles per SSR primer pair.A genetic clustering map was constructed and genetic similarity for isolates of V. inaequalis was from 0.72 to 1.0 with an average of 0.85. At the genetic distance of 0.81,20 isolates were divided into 6 groups. Except group 1 and group 4, isolates in the same group belonged to different races, which indicated that genetic diversity of 20 isolates of V. inaequalis had minor relevance with their races.
2. Using amplified fragment length polymorphism (AFLP) technique, we obtained 25 pairs of primers for study on genetic diversity of 46 isolates of V. inaequalis from Xingping and Xunyi in shaanxi province. Genetic clustering map was constructed based on these results. The results indicated that, all the 46 isolates were clustered into two groups at the 0.68 genetic similarity level. Group I contained all the isolates from Xunyi fuji, Xunyi Gala and Xingping Gala. Group II contained all the isolates from Xunyi Starking Delicious, Xunyi Qinguan, Xingping Starking Delicious and Xingping Qinguan. These results showed that the isolates of Venturia inaequalis from Xingping and Xunyi didn't have a relative relationship based on the regional source but had a relative relationship on the host cultivars.
3. Universal primers ITS1 and ITS4 were used for PCR amplification of 26 isolates from V. inaequalis, Alternaria mali, Marssonina cororlar and Podospharea leucotricha. Twenty-six ITS sequences were obtained. Based on sequence alignment result, one specific primer pair was obtained and the size of PCR product was 320 bp. The primer pair 320A/320B was specific to V. inaequalis and the pathogen could be detected in apple leaves without symptom with the primer pair. The sensitivity of this primer pair for DNA of V. inaequalis was 100 fp/μl.
1、采用SSR分子标记技术,利用筛选出的18对SSR引物对中国、英国、印度三个国家的90株苹果黑星病菌进行遗传多样性分析,构建了遗传聚类分析图。结果表明：在相似系数为0.76的水平,供试的90个菌株全部聚到一类。在相似系数为0.80的水平,90个菌株被划分为2个类群：全部的英国菌株(共18株)被分在了第Ⅰ类群；中国和印度菌株(共72株)被分在了第Ⅱ类群。而在相似系数为0.83时,第Ⅱ类群又可分为3个亚群：全部的印度菌株被分在了A亚群；全部的兴平富士(4株)、全部的红星(16株)、全部的秦冠(9株)和11%的兴平嘎啦(1株)上采集的菌株被分在B亚群；全部的旬邑嘎啦(14株)、全部的旬邑富士(9株)和89%的兴平嘎啦(8株)上采集的菌株被分在C亚群。根据结果分析得出,三个国家的苹果黑星病菌与地理来源存在一定的相关性：中国菌株与印度菌株亲缘关系较近,而与英国菌株亲缘关系较远；从中国内部菌株来看,病菌亲缘关系与地理来源相关性不大,但与寄主苹果品种有一定相关性。同时采用5对SSR引物对来自欧洲的20株分属于7个不同生理小种的苹果黑星病菌进行PCR扩增,共检测到43个等位变异,等位位点数目范围6～11个,平均为8.6个；构建遗传聚类分析图,病菌遗传相似系数变异范围为0.72～1.00,平均值为0.85；在遗传距离为0.81的水平上,将20个苹果黑星病菌株分为六大类。除第一、四大类外,其余四类每类内的菌株都分属于不同的生理小种,说明苹果黑星病菌基于SSR标记划分的类群与生理小种之间无明显相关性。
2、采用AFLP分子标记技术,利用筛选出的25对AFLP引物对来自我国陕西兴平和旬邑的46株苹果黑星病菌进行遗传多样性分析,构建了遗传聚类分析图。结果表明,在相似系数为0.68的水平,46个菌株被划分为2个类群：第Ⅰ大类由全部的旬邑富士旬邑嘎啦和兴平嘎啦采集菌株组成；第Ⅱ大类由全部的旬邑红星、旬邑秦冠、兴平红星、兴平秦冠采集菌株组成。表明陕西兴平和旬邑两地的苹果黑星病菌遗传多样性与地理来源无明显相关性,而与其寄主品种间存在一定的相关性。
3、应用通用引物ITS1和ITS4对苹果黑星病菌(V. inaequalis)、苹果斑点落叶病菌(Alternaria mali)、苹果褐斑病菌(Marssonina cororlar)和苹果白粉病菌(Podospharea leucotricha)等4种苹果树常见病害的病原菌共26个菌株进行PCR扩增和测序,得到26条ITS序列。对其进行比对筛选,获得1对片段大小为320 bp的特异性引物320A/320B。应用该特异性引物对26个苹果病原真菌供试菌株和接有苹果黑星病菌的苹果叶片组织进行PCR分子检测,结果表明：特异性引物320A/320B能够特异的检测出供试菌株中的苹果黑星病菌,同时也能从接种过苹果黑星病菌但未显症的苹果组织中检测到苹果黑星病菌的存在；对苹果黑星病菌基因组DNA检测的灵敏度为100fg/μL。
Apple scab is one of the main diseases of apples. It is characterized by fast epidemic, serious damage and difficalf to control. It is one of the most important apple diseases in Europe and America and often causes yield and economic losses. In China, apple scab distributes mainly in Liaoning province, Heilongjiang province, Gansu province, and same areas of Shaanxi province, Xinjiang province and Shandong province. It has caused serious effect on industry of apple because of large-scale epidemic. In this study, we performed research on genetic diversity and molecular rapid detection of Venturia inaequalis and obtained the results as follows:
1. Eighteen pairs of primers were selected for study on genetic diversity of 90 isolates of V. inaequalis from China, India and United kingdom by using simple sequence repeats (SSR) technique. Genetic clustering map was constructed based on these results. The results indicated that, all the 90 isolates were clustered together at the 0.76 genetic similarity level. At the 0.80 genetic similarity level, all the isolates (18) from United kingdom were clustered in groupⅠand isolates from China and India (72) were clustered into groupⅡ. At the 0.83 genetic similarity level, isolates from groupⅡwere divided into 3 subgroup:all the Indian isolates belonged to subgroup A, all the isolates from Xingping Fuji, Starking Delicious, Qinguan and 11% isolates from Xingping Gala had clustered into subgroup B and all the isolates from Xunyi Gala, Xunyi fuji and 89% isolates from Xingping Gala were clustered into subgroup C. We can conclude that the isolates of V. inaequalis from three countries were relative relationship based on the regional source. The isolates from China were relatively close to that from India and far from the UK. All the isolates from China were not relatively close based on the source but had a relative relationship on cultivars. Five pairs of primers were used in PCR amplification of 20 isolates of V. inaequalis, which belong to 7 physiological races, through SSR technique. Fourty three alleles were detected with an average of 8.6 alleles per SSR primer pair.A genetic clustering map was constructed and genetic similarity for isolates of V. inaequalis was from 0.72 to 1.0 with an average of 0.85. At the genetic distance of 0.81,20 isolates were divided into 6 groups. Except group 1 and group 4, isolates in the same group belonged to different races, which indicated that genetic diversity of 20 isolates of V. inaequalis had minor relevance with their races.
2. Using amplified fragment length polymorphism (AFLP) technique, we obtained 25 pairs of primers for study on genetic diversity of 46 isolates of V. inaequalis from Xingping and Xunyi in shaanxi province. Genetic clustering map was constructed based on these results. The results indicated that, all the 46 isolates were clustered into two groups at the 0.68 genetic similarity level. Group I contained all the isolates from Xunyi fuji, Xunyi Gala and Xingping Gala. Group II contained all the isolates from Xunyi Starking Delicious, Xunyi Qinguan, Xingping Starking Delicious and Xingping Qinguan. These results showed that the isolates of Venturia inaequalis from Xingping and Xunyi didn't have a relative relationship based on the regional source but had a relative relationship on the host cultivars.
3. Universal primers ITS1 and ITS4 were used for PCR amplification of 26 isolates from V. inaequalis, Alternaria mali, Marssonina cororlar and Podospharea leucotricha. Twenty-six ITS sequences were obtained. Based on sequence alignment result, one specific primer pair was obtained and the size of PCR product was 320 bp. The primer pair 320A/320B was specific to V. inaequalis and the pathogen could be detected in apple leaves without symptom with the primer pair. The sensitivity of this primer pair for DNA of V. inaequalis was 100 fp/μl.
引文
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