大麦白粉菌遗传多样性分析、抗病基因鉴定及致病型鉴别寄主体系建立
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摘要
大麦白粉病是由布氏白粉菌大麦专化型Blumeria graminis f. sp. hordei(简称Bgh)活体寄生大麦叶片引起的病害,多发生在温带潮湿、半潮湿地区,一般可造成20%以上的产量损失,严重时损失达30%,目前是中国冬大麦区重要的病害之一。近年来,由于作物种植密度提高,氮肥使用量增大,以及种植品种单一化,从而使该病害日益严重。大麦抗白粉病育种是控制该病害最有效的手段,不但能减少产量损失,还能减少化学农药的使用、降低环境污染,Bgh群体遗传结构研究和抗病基因鉴定是合理布局和利用抗白粉病基因的先决条件。本研究利用'Pallas'近等基因系和其它5个大麦品种作为鉴别寄主,鉴定分析了我国大麦白粉菌种群毒性与致病型构成、分布和动态；运用AFLP (amplified fragment length polymorphism, AFLP)分子标记技术,对来源于不同地区、不同致病型菌株基因组DNA进行多态性分析,进一步揭示我国大麦白粉菌种群遗传变异程度和演化规律。通过抗谱分析的方法,鉴定了我国大麦主产区大麦骨干亲本和育成品种的抗病性和抗病基因；选择112个具有广泛代表性的Bgh菌株接种备选大麦品种(系),筛选我国大麦白粉病致病型鉴别品种。
2006年,从我国7个冬大麦主产区不同栽培品种上采集575个Bgh菌株,利用26个鉴别寄主进行毒性鉴定。所有的菌株共划分为58个不同的致病型,81%的菌株包含于其中13个致病型中,最优势的致病型为0047(18.3%),其次分别为0045(11.8%)和(7.8%)；7个不同的Bgh群体中大多数毒性基因频率相似。因此,中国冬大麦区可视为统一的大麦白粉病害流行区,这些地区所种植的大麦品种可能具有相似的遗传背景。KOIND软件分析显示,Bgh不同群体内和群体间遗传多样性指数显著不同,不同地区之间的遗传距离大小和地理距离远近无相关性。由此表明,中国冬大麦区Bgh群体遗传多样性主要源于本地区病原菌的遗传分化和长距离病原菌传播的基因流动。
分子标记技术可对病原菌基因组的多个变异位点进行遗传分析,已成为研究病原菌群体遗传多样性的重要手段,本研究利用AFLP分子标记,对来源于7个不同地区、12个不同致病型的79个Bgh菌株基因组DNA进行多态性检测,筛选的7对引物组合共扩增出337个多态性位点。UPGMA (unweighted pair group method average,UPGMA)法聚类分析,供试菌株可划分为7个不同的遗传宗谱,遗传宗谱与致病型不存在一一对应关系,但具有一定的相关性,与菌株的地理来源密切关联。通过POPGENE软件分析可见,我国冬大麦区Bgh群体具有丰富的遗传多样性,不同地区Bgh群体遗传多样性与该地区大麦播种面积、主栽品种及白粉病易于发生和越夏的气候条件有关；不同地区间Bgh群体遗传一致度较大,遗传距离大小与地理距离远近呈正相关,Bgh群体基因流动性较大。分子标记与毒性鉴定方法分别对Bghg群体遗传多样性检测的部分结果相互矛盾,但运用分子标记技术得到的研究结果能更为合理地揭示Bgh不同地理群体遗传分化的特点。
对2005和2006年Bgh群体毒性频率进行监测,通过抗谱分析的方法鉴定了328份大麦品种(系)的白粉病抗性和抗病基因。结果显示：大麦白粉菌群体对抗病基因Mla3、Mla9、Mlp1、Mlal、Mla(A12)、Mla6Mlal4、Mla7Mla (No3)、Mla7Ml (Lg2)、Mla9Mk、Mla13MlaRu3、Mg (Cp)和mlo5的毒性频率为0,对Mla12MlaEm2、Mla7Mlk、Mlat Mla8、Ma10MaDu2和.Mlkl的毒性频率很低,分别为0.1%、0.4%、0.9%、2.8%和4.2%；这些抗病基因可应用于大麦抗病育种中。所鉴定的328份材料绝大多数感病,抗病材料仅37份,能明确推导出抗白粉病基因的品种(品系)很少,这些品种(品系)可能含有的抗白粉病基因为M(Bw)、Ma8、Mg、 Mlra Mla8、Ma9Mlk、Mla1Ma (A12)或mlo5。
来源于不同地区、不同致病型Bgh菌株112个,接种筛选的112个来源于不同地区的大麦主栽品种(系)和骨干亲本。根据寄主毒性频率的梯度间隔、反应型的稳定性和明确性及寄主不同的遗传背景,选择鉴别品种(系),建立一套适合我国大麦白粉菌致病型鉴别寄主体系。结果显示,供试菌株对寄主的毒性频率在0～60%范围内呈连续的变化,毒性频率在60%-70%范围内的品种无,1个品种的毒性频率在70%-80%范围内,1个品种的毒性频率在80%-90%范围内,1个品种的毒性频率为100%。大多数寄主对菌株的反应症状明确、稳定,中间类型较少。筛选出9个品种作为我国大麦白粉菌致病型分类的鉴别寄主,分别是：单二、肚里黄、云大麦2号、鄂大麦6号、苏啤4号、北青8号、莆大麦7号、浙农大7号和浙皮3号,以萧山立夏黄为感病对照。这套鉴别可将112菌株划分为50个致病型,且致病型优势性明显,具有很强的可行性。
Barley powdery mildew caused by Blumeria graminis. f.sp. hordei (Bgh), which is a windborne and obligate biotrophic fungal pathogen, is one of the most destructive foliar diseases of barley. It is particularly prevalent in cooler and more humid regions of temperate climates, and it is one of more serious barley diseases in the winter barley regions of China. Barley powdery mildew can reduce grain yield by20%to30%. In recent years, barley powdery mildew in China is getting increasingly serious because of densely seeding, using larger amount of nitrogenous fertilizer and planting the same barley cultivars. Breeding for genetic resistance is an economically and environmentally sustainable way to control powdery mildew. Growing resistant cultivars against powdery mildew not only diminishes crop losses and fungicide spraying costs, but also reduces environmental pollution. Knowledge of the genetic structure and evolution of Bgh populations and identification of resistance genes to powdery mildew is prerequisite for an efficient use of available resistance genes. In this study, the frequency, distribution and dynamics of virulence and pathotypes in Bgh populations in the winter barley regions of China were investigated on'Pallas near-isogenic lines' and other five barley varieties as differential hosts. The genetic diversity of79Bgh isolates, selected from different pathotypes in respective region, was detected by AFLP to demonstrate further genetic diversity level in Bgh populations of China. Resistance and resistance genes to powdery mildew in some leading barley cultivars and backbone parents from major barley growing areas in China were identified by analysis of resistance reaction spectra;112widely representative Bgh isolates were selected to inoculate some core barley varieties (lines) for screening differential barley hosts for determining Bgh pathotypes in China。
575isolates were collected from seven populations of Bgh on different cultivated barley in seven geographically distant locations in2006. Their virulence was determined by inoculation onto26differential host lines. All of the isolates belonged to58pathotypes and13of which included81%of these isolates. The most abundant pathotype was pathotype 0047(18.3%), the second most abundant was pathotype0045(11.8%) and the third most abundant was pathotype0057(7.8%). Most of virulent genes investigated in this study showed similar frequencies in the seven different areas. These indicate that the seven locations may be in a uniform epidemiological region and barley cultivars in these areas may have similar genetic background. Diversities within these populations and distances between these populations measured by KOIND package were different. Correlations were not found between the genetic distances and the geographical distances between different locations. This suggested that long distance spread and local epidemics existed in the major winter barley growing regions in China.
Molecular markers, which can detect high level of allelic polymorphism, have become a important tool for studying genetic diversity in pathogen populations. In this study, genetic diversity of79Bgh isolates collected from7different locations in the winter barley regions of China, which belong to12different pathotypes, was assessed using AFLP (amplified fragment polymorphism, AFLP) markers. Cluster analysis of UPGMA (unweighted pair group method average, UPGMA) based on a total of337polymorphic AFLP bands detected among the79isolates showed seven different genetic lineages. There was some correlation between pathotypes and AFLP groups, and a close correlation between the genetic lineages and origins of the genotypes. The diversity parameter obtained with POPGENE program showed the Bgh population in the winter barley regions of China possessed relatively high levels of genetic diversity, and the level of genetic diversity detected in some Bgh population was different. These results were related with some factors causing evolution of Bgh such as the size of barley area planted every year, the environmental conditions which impaired life cycle of Bgh on barley host. The higher genetic similarity between different Bgh populations was found, and the positive correlations between the geographical distances and the genetic distances amongst these locations were found. These results indicated that the main allelic variation existed within Bgh populations, and there is an extensive gene flow and migration of pathogen among the different regions. Some results of genetic variation of Bgh populations obtained with AFLP analysis were not in agreement with some of results detected by virulence assessments, but the causes for evolution in natural Bgh populations concluded from AFLP analysis was more reasonable. 'Pallas' isogenic lines possessing known Bgh resistant genes were used in virulence analysis for the population consisting of729isolates collected from the winter barley region of China in2005and2006, and resistance and resistance genes to powdery mildew in328barley varieties (lines) were identified. All isolates were avirulent to Mlal Mla (A12), Mla3, Mla6Mlal4, Mla7Mla (No3), Mla7Ml (Lg2), Mla9Mlk, Mla9, Mlal3MlaRu3, Mlpl, Mlg (Cp) and mlo5, and the frequencies of isolates overcoming the genes Mlal2MlaEm2(0.1%), Mla7Mlk (0.4%), Mlat Mla8(0.9%), Mla10MlaDu2(2.8%) and Mlkl (4.2%) were very low. These results indicated that these resistance genes can be using in barley breeding program. Majority of328varieties (lines) tested were susceptible to powdery mildew, but37of these were resistant to all dominant pathotypes. Resistance genes can be postulated in less of the lines investigated, and the resistance genes in these lines were Ml (Bw), Mla8, Mlg, Mlra Mla8, Mla9Mlk, Mlal Mla (A12) or mlo5.
112representative isolates of Bgh selected from different locations and different pathotypes, were inoculated on112barley varieties including leading barley cultivars and backbone parents in China. A set of differential hosts which is suitable to test pathotypes of Bgh in China was established by investigating on infection types and virulence frequency of112isolates on112barley varieties. The variation of virulence frequency from0to60%was continuous, and virulence frequency within60%to70%was not found, one was within70%to80%, one was within80%to90%, and one was100%. Infection symptom of most inoculated hosts was obvious and stable. Nine barley varieties were chosen differential hosts to identify pathotypes of Bgh in China, and they were Daner, Dulihuang, Yun2, E6, Supi4, Beiqing8, Pu7, Zhenongda7, and Zhepi3respectively, and more, Xiaoshanglixiahuang was used as a susceptible control variety. These barley varieties were used to test the112isolates,50pathotypes, of which a few were predominent pathotypes, were found. These results showed that these differential barley varieties were suitable to identify pathotypes of Bgh population in China.
2006年,从我国7个冬大麦主产区不同栽培品种上采集575个Bgh菌株,利用26个鉴别寄主进行毒性鉴定。所有的菌株共划分为58个不同的致病型,81%的菌株包含于其中13个致病型中,最优势的致病型为0047(18.3%),其次分别为0045(11.8%)和(7.8%)；7个不同的Bgh群体中大多数毒性基因频率相似。因此,中国冬大麦区可视为统一的大麦白粉病害流行区,这些地区所种植的大麦品种可能具有相似的遗传背景。KOIND软件分析显示,Bgh不同群体内和群体间遗传多样性指数显著不同,不同地区之间的遗传距离大小和地理距离远近无相关性。由此表明,中国冬大麦区Bgh群体遗传多样性主要源于本地区病原菌的遗传分化和长距离病原菌传播的基因流动。
分子标记技术可对病原菌基因组的多个变异位点进行遗传分析,已成为研究病原菌群体遗传多样性的重要手段,本研究利用AFLP分子标记,对来源于7个不同地区、12个不同致病型的79个Bgh菌株基因组DNA进行多态性检测,筛选的7对引物组合共扩增出337个多态性位点。UPGMA (unweighted pair group method average,UPGMA)法聚类分析,供试菌株可划分为7个不同的遗传宗谱,遗传宗谱与致病型不存在一一对应关系,但具有一定的相关性,与菌株的地理来源密切关联。通过POPGENE软件分析可见,我国冬大麦区Bgh群体具有丰富的遗传多样性,不同地区Bgh群体遗传多样性与该地区大麦播种面积、主栽品种及白粉病易于发生和越夏的气候条件有关；不同地区间Bgh群体遗传一致度较大,遗传距离大小与地理距离远近呈正相关,Bgh群体基因流动性较大。分子标记与毒性鉴定方法分别对Bghg群体遗传多样性检测的部分结果相互矛盾,但运用分子标记技术得到的研究结果能更为合理地揭示Bgh不同地理群体遗传分化的特点。
对2005和2006年Bgh群体毒性频率进行监测,通过抗谱分析的方法鉴定了328份大麦品种(系)的白粉病抗性和抗病基因。结果显示：大麦白粉菌群体对抗病基因Mla3、Mla9、Mlp1、Mlal、Mla(A12)、Mla6Mlal4、Mla7Mla (No3)、Mla7Ml (Lg2)、Mla9Mk、Mla13MlaRu3、Mg (Cp)和mlo5的毒性频率为0,对Mla12MlaEm2、Mla7Mlk、Mlat Mla8、Ma10MaDu2和.Mlkl的毒性频率很低,分别为0.1%、0.4%、0.9%、2.8%和4.2%；这些抗病基因可应用于大麦抗病育种中。所鉴定的328份材料绝大多数感病,抗病材料仅37份,能明确推导出抗白粉病基因的品种(品系)很少,这些品种(品系)可能含有的抗白粉病基因为M(Bw)、Ma8、Mg、 Mlra Mla8、Ma9Mlk、Mla1Ma (A12)或mlo5。
来源于不同地区、不同致病型Bgh菌株112个,接种筛选的112个来源于不同地区的大麦主栽品种(系)和骨干亲本。根据寄主毒性频率的梯度间隔、反应型的稳定性和明确性及寄主不同的遗传背景,选择鉴别品种(系),建立一套适合我国大麦白粉菌致病型鉴别寄主体系。结果显示,供试菌株对寄主的毒性频率在0～60%范围内呈连续的变化,毒性频率在60%-70%范围内的品种无,1个品种的毒性频率在70%-80%范围内,1个品种的毒性频率在80%-90%范围内,1个品种的毒性频率为100%。大多数寄主对菌株的反应症状明确、稳定,中间类型较少。筛选出9个品种作为我国大麦白粉菌致病型分类的鉴别寄主,分别是：单二、肚里黄、云大麦2号、鄂大麦6号、苏啤4号、北青8号、莆大麦7号、浙农大7号和浙皮3号,以萧山立夏黄为感病对照。这套鉴别可将112菌株划分为50个致病型,且致病型优势性明显,具有很强的可行性。
Barley powdery mildew caused by Blumeria graminis. f.sp. hordei (Bgh), which is a windborne and obligate biotrophic fungal pathogen, is one of the most destructive foliar diseases of barley. It is particularly prevalent in cooler and more humid regions of temperate climates, and it is one of more serious barley diseases in the winter barley regions of China. Barley powdery mildew can reduce grain yield by20%to30%. In recent years, barley powdery mildew in China is getting increasingly serious because of densely seeding, using larger amount of nitrogenous fertilizer and planting the same barley cultivars. Breeding for genetic resistance is an economically and environmentally sustainable way to control powdery mildew. Growing resistant cultivars against powdery mildew not only diminishes crop losses and fungicide spraying costs, but also reduces environmental pollution. Knowledge of the genetic structure and evolution of Bgh populations and identification of resistance genes to powdery mildew is prerequisite for an efficient use of available resistance genes. In this study, the frequency, distribution and dynamics of virulence and pathotypes in Bgh populations in the winter barley regions of China were investigated on'Pallas near-isogenic lines' and other five barley varieties as differential hosts. The genetic diversity of79Bgh isolates, selected from different pathotypes in respective region, was detected by AFLP to demonstrate further genetic diversity level in Bgh populations of China. Resistance and resistance genes to powdery mildew in some leading barley cultivars and backbone parents from major barley growing areas in China were identified by analysis of resistance reaction spectra;112widely representative Bgh isolates were selected to inoculate some core barley varieties (lines) for screening differential barley hosts for determining Bgh pathotypes in China。
575isolates were collected from seven populations of Bgh on different cultivated barley in seven geographically distant locations in2006. Their virulence was determined by inoculation onto26differential host lines. All of the isolates belonged to58pathotypes and13of which included81%of these isolates. The most abundant pathotype was pathotype 0047(18.3%), the second most abundant was pathotype0045(11.8%) and the third most abundant was pathotype0057(7.8%). Most of virulent genes investigated in this study showed similar frequencies in the seven different areas. These indicate that the seven locations may be in a uniform epidemiological region and barley cultivars in these areas may have similar genetic background. Diversities within these populations and distances between these populations measured by KOIND package were different. Correlations were not found between the genetic distances and the geographical distances between different locations. This suggested that long distance spread and local epidemics existed in the major winter barley growing regions in China.
Molecular markers, which can detect high level of allelic polymorphism, have become a important tool for studying genetic diversity in pathogen populations. In this study, genetic diversity of79Bgh isolates collected from7different locations in the winter barley regions of China, which belong to12different pathotypes, was assessed using AFLP (amplified fragment polymorphism, AFLP) markers. Cluster analysis of UPGMA (unweighted pair group method average, UPGMA) based on a total of337polymorphic AFLP bands detected among the79isolates showed seven different genetic lineages. There was some correlation between pathotypes and AFLP groups, and a close correlation between the genetic lineages and origins of the genotypes. The diversity parameter obtained with POPGENE program showed the Bgh population in the winter barley regions of China possessed relatively high levels of genetic diversity, and the level of genetic diversity detected in some Bgh population was different. These results were related with some factors causing evolution of Bgh such as the size of barley area planted every year, the environmental conditions which impaired life cycle of Bgh on barley host. The higher genetic similarity between different Bgh populations was found, and the positive correlations between the geographical distances and the genetic distances amongst these locations were found. These results indicated that the main allelic variation existed within Bgh populations, and there is an extensive gene flow and migration of pathogen among the different regions. Some results of genetic variation of Bgh populations obtained with AFLP analysis were not in agreement with some of results detected by virulence assessments, but the causes for evolution in natural Bgh populations concluded from AFLP analysis was more reasonable. 'Pallas' isogenic lines possessing known Bgh resistant genes were used in virulence analysis for the population consisting of729isolates collected from the winter barley region of China in2005and2006, and resistance and resistance genes to powdery mildew in328barley varieties (lines) were identified. All isolates were avirulent to Mlal Mla (A12), Mla3, Mla6Mlal4, Mla7Mla (No3), Mla7Ml (Lg2), Mla9Mlk, Mla9, Mlal3MlaRu3, Mlpl, Mlg (Cp) and mlo5, and the frequencies of isolates overcoming the genes Mlal2MlaEm2(0.1%), Mla7Mlk (0.4%), Mlat Mla8(0.9%), Mla10MlaDu2(2.8%) and Mlkl (4.2%) were very low. These results indicated that these resistance genes can be using in barley breeding program. Majority of328varieties (lines) tested were susceptible to powdery mildew, but37of these were resistant to all dominant pathotypes. Resistance genes can be postulated in less of the lines investigated, and the resistance genes in these lines were Ml (Bw), Mla8, Mlg, Mlra Mla8, Mla9Mlk, Mlal Mla (A12) or mlo5.
112representative isolates of Bgh selected from different locations and different pathotypes, were inoculated on112barley varieties including leading barley cultivars and backbone parents in China. A set of differential hosts which is suitable to test pathotypes of Bgh in China was established by investigating on infection types and virulence frequency of112isolates on112barley varieties. The variation of virulence frequency from0to60%was continuous, and virulence frequency within60%to70%was not found, one was within70%to80%, one was within80%to90%, and one was100%. Infection symptom of most inoculated hosts was obvious and stable. Nine barley varieties were chosen differential hosts to identify pathotypes of Bgh in China, and they were Daner, Dulihuang, Yun2, E6, Supi4, Beiqing8, Pu7, Zhenongda7, and Zhepi3respectively, and more, Xiaoshanglixiahuang was used as a susceptible control variety. These barley varieties were used to test the112isolates,50pathotypes, of which a few were predominent pathotypes, were found. These results showed that these differential barley varieties were suitable to identify pathotypes of Bgh population in China.
引文
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