小麦全蚀病菌的遗传组成和病害的防治技术研究
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摘要
近年来,小麦全蚀病和纹枯病在我国黄淮和长江下游麦区的发生呈上升趋势,现已成为该地区小麦上的主要病害。了解该地区小麦全蚀病菌组成、分布,对了解病害的发展和预测病害发生具有重要意义。为减少病害的危害,目前也急需对病害的控制技术进行筛选。
为此,首先从黄淮麦区的河南、山东、安徽和江苏4省采集和分离了104株小麦全蚀病病菌,采用GGT-RP:NS5和GGA-RP:NS5两对特异性引物对病菌基因组DNA进行PCR扩增,从而鉴定这些菌株的变种类型。结果表明,所分离菌株都为禾顶囊壳小麦变种。对3种小麦全蚀病菌基因型的分析方法进行了比较,发现ITS序列分析的方法是禾顶囊壳小麦变种下分型的可靠方法。对核糖体rDNA部分序列进行的系统发育学分析结果表明,根据rDNA ITS1区的序列,供试菌株可划分为两个基因型,与国外小麦全蚀病菌基因型的分类描述有所不同,我们暂将这两个基因型命名为Q1和Q2型。所分离菌株中,Q1型菌数量高于Q2型。河南菌株中,2010年和2011年两种基因型菌株的比例不同,这种变化与病害发展和防控措施的实行有无关系还需进行进一步的研究。
为了解该区域病菌的致病力,首先比较了小麦全蚀病菌致病力的3种接种测定方法和2种病情分级标准。比较结果表明,以蛭石为培养基质、麦种下2cm接种菌丝块的接种方法和0-6级的分级标准,可比较有效和准确的测定小麦全蚀病菌的致病力。对2010年和2011年分离自黄淮麦区的104株小麦全蚀病菌进行了致病力测定,结果表明供试菌株的致病力存在明显差异。不同地区菌群间致病力差异不显著,Q1型和Q2型菌株间的致病力没有显著差异。
利用公共的真菌基因组数据库资源,对小麦全蚀病病菌G. graminis及与其亲缘关系较近的M. oryzae和M. poae三种真菌基因组中SSRs的结构类型、分布、丰度及最长序列等进行了系统分析并做了比较。结果表明：这三种真菌基因组中的SSRs非常丰富,分别为13221、9977和11422个。编码基因的序列中二、四、五核苷酸重复基序的SSRs丰富度要低于全基因组中SSRs的丰富度,而三、六核苷酸重复基序的SSRs丰富度要高于全基因组中SSRs的丰富度。G. graminis基因组中二、三、四、五、六核苷酸重复基序的SSRs丰富度要远高于M. oryzae和M. poae基因组中的同类SSRs,说明G. graminis基因组可能具有较高的变异度。本研究对禾谷顶囊壳小麦变种基因组中SSRs的丰度、出现频率及最长基序的分析为快速、便捷地设计多态性丰富的SSRs引物提供了有益的信息。
在对已公布的禾顶囊壳小麦变种菌株R3-111a-1基因组中SSR组成和结构分析的基础上,选取不同核苷酸重复方式的较长重复基序,根据其两端的保守序列设计17条引物。利用2010年分离菌株对这些引物进行筛选,获得4对扩增多态性较好的引物G7、G9、G14、G17。这4个SSR标记共可以获得50个等位基因,每个基因座的等位基因数为2-7个,平均数为5个。应用以上的4个SSR标记对2010年从黄淮麦区分离的52个菌株进行了遗传分析。4个标记可把52个菌株分为两个大的簇。群体遗传分析的结果表明,江苏病菌群体的遗传多样性水平最高,其次为山东群体,河南和安徽群体的多样性水平最低。84%的变异来自于群体内,只有16%的变异来自于群体间,4个群体间的基因流(Nm)为2.604。以安徽和山东群体的遗传相似性最高。群体分析的结果说明,安徽和河南的小麦全蚀病菌可能分别来自于山东病区。
杀菌剂种子处理是目前小麦全蚀病防治的主要方法,为了筛选高效安全的药剂。首先测定了多种杀菌剂对小麦全蚀病菌的抑菌活性,并对杀菌剂在不同浓度下的防效以及对小麦的安全性进行了盆栽试验。室内毒力测定结果表明：甾醇脱甲基化抑制剂类杀菌剂(DMIs)对小麦全蚀病病菌菌丝生长有显著的抑制作用,其中氟喹唑、氟环唑、三唑醇、咪鲜胺、戊唑醇和苯醚甲环唑的平均EC50值分别为0.03,0.03,0.06,0.07,0.10和0.24mg/L。盆栽试验结果表明：硅噻菌胺和苯醚甲环唑在推荐浓度下对小麦全蚀病有很好的防治效果,并且对小麦出苗、苗高没有显著影响；咯菌腈的防效与苯醚甲环唑相似,但对小麦出苗有轻微的抑制作用；其它几种杀菌剂对小麦全蚀病也能起到很好的防治效果,但是对小麦的生长有一定的抑制作用。从防治效果、对小麦安全性以及抗药性治理方面综合考虑,将硅噻菌胺、苯醚甲环唑和咯菌腈等交替或混合使用是防治小麦全蚀病比较好的策略。
纹枯病是长江中下游小麦上的重要病害,稻秸秆还田是该地区广泛推广的农业栽培措施。为了解稻秸秆还田对小麦根围拮抗细菌和小麦纹枯病发生的影响,本研究通过多年的田间试验,利用基于培养的方法比较了稻秸秆还田2年和3年小区小麦根围和非根围的细菌和真菌群体。结果表明,稻秸秆还田可增加小麦根围和非根围的产荧光假单胞菌菌落形成单位的数量。稻秸秆还田土壤中的产荧光假单胞菌在总细菌中的比例高于非覆盖土壤。利用平板对对峙法测定了分离细菌菌株对禾谷丝核菌Rhizoctonia cerealis的拮抗作用,并根据16s rDNA的部分序列对菌株进行了鉴定。测定结果表明,与其他种的细菌相比,假单胞杆菌对禾谷丝核菌具有较强的拮抗作用,80%以上的小麦根围发荧光假单胞菌对禾谷丝核菌具有拮抗作用。稻秸秆还田小区的小麦纹枯病病情指数低于未还田小区。研究结果表明,稻麦轮作地区的稻秸秆还田可以增加土壤中的产荧光假单胞菌的群体数量,同时产荧光假单胞菌可以对小麦纹枯病的发生起控制作用。
The incidence of take-all caused by Gaeumannomyces graminis var. tritici (Ggt) and sharp eyespot caused by Rhizoctonia cerealis has been increasing in wheat growing areas of Huang-Huai and lower reaches of Yangtze river and becoming the important diseases of wheat recently. In order to predict and control the occurrence of wheat take-all, it is necessary to understand the genetic structure of the pathogen and the trend of population structure changes. It is also urgently needed to find the effective fungicides and other measures to prevent the disease.
In the present study,104isolates of G graminis collected from Henan, Shandong, Anhui, and Jiangsu province in Huang-Huai wheat growing area were identified by amplifying the genomic DNA with two specific primers GGT-RP:NS5and GGA-RP:NS5, respectively. The results showed that all the isolates were G graminis var. tritici (Ggt) genotype. ITS sequencing was the most reliable method to analyze Ggt genotype. All tested isolates were distributed into two main genetic groups, named as Q1and Q2, respectively. Both of which were different from any isolates out of China. The genetic frequency in Q1was higher than that of in Q2. Q1/Q2frequencies in Henan province were different during2years. What resulted in the diffenence of Q1/Q2frequency in Henan province needs further study.
To determine the virulence of Gaeumannomyces graminis isolates, three inoculation and two investigation methods were compared. The results suggested that the method with fungal-colonized agar plugs2cm below the wheat seeds combined with the0-6disease grade standard was better than the others, which presented appropriate disease severity and less standard error. The virulence of104G. graminis isolates collected from Henan, Jiangsu, Anhui, and Shandong province in2010and2011was then tested using the method mentioned above. The results showed that the virulence was different significantly among the isolates. There is no significant difference on virulence between populations collected from different provinces. The virulence between genotypes Q1and Q2has also no significant difference.
Base on the public genome database, we examined the SSRs in the completely sequenced G. graminis var. tritici genome. The occurrences, relative abundance, relative density, the commonest, and the longest SSRs were analyzed, and compared with other plant pathogenic fungal species:Magnaporthe oryzae and M. poae. The results revealed that the SSRs were abundant in these three genomes, total of13221,9977and11422respectively. In the gene sequences, the Di-, Tetra-and Pentanucleotide repeats were less abundant than in the total genome sequences, while the Tri-and Hexanucleotide repeats were more abundant than in the total genome sequences. In the G. graminis var. tritici genome sequence, Di-, Tri-, Tetra-, Penta-and Hexanucleotide repeats were more abundant than the same repeats in M. oryzae and M. poae genome sequences, indicated that the G. graminis var. tritici maybe is a species with more genetic diversity. The analysis in this study provided new information that could be useful for a variety of applications in population genetics of G. graminis var. tritici.
For analysis of genetic diversity and genetic structure of the take-all pathogen, based on the analysis of simple sequence repeats in genomes of Gaeumannomyces graminis var. tritici, we isolated longer SSR motifs of different repeat types and designed17primers for screening. Four SSRs markers with reproducible patterns and diversity between different Ggt isolates were developed. Their usefulness for gentic analysis was assessed on52isolates collected in2010from Huang-Huai wheat growing areas. Four markers displayed each from2to7alleles, with an average of5alleles per locus. Phylogenetic analysis based on these markers showed that55isolates could be clustered into2large clusters. Jiangsu population showed the highest level of genetic diversity, followed by Shandong, Henan and Anhui population.84%genetic variance was within populations. The gene flow (Nm) among populations was2.604.The genetic identities between Anhui and Shandong population was the highest.
In order to screening the effective and safe fungicides for the control of wheat take-all, the inhibitory activity of different fungicides in vitro against the pathogen Gaeumammomyces graminis causing wheat take-all was tested, and control efficacy and safety of fungicides at different concentrations were also investigated in pot experiments. In vitro, sterol demethylation inhibitors fungicides (DMIs) exhibited better inhibitory activity in hypha growth of G. graminis, and the EC50values of fluquinconazole, flusilazole, riadimenol, prochloraz, tebuconazole and difenoconazole were0.03,0.03,0.06,0.07, 0.10and0.24mg/L, respectively. The results of pot experiments indicated that silthiopham and difenoconazole exhibited good control efficacy in controlling wheat take-all, and did not affect the emergence and height of seedlings. The control efficiency of fludloxonll was similar to that of difenoconazole, but slightly inhibited the emergence of seedling. The others also provided good control efficacy, but significantly inhibited the growth of the wheat seedlings. Taken together with the aspects of safety to wheat and resistance management, silthiopham, difenoconazole and fludloxonll were considered suitable for the control of wheat take-all, and should be used alternatively.
Wheat sharp eyespot caused by Rhizoctonia cerealis is one of the main diseases in the lower reaches of the Yangtze River, where rice-wheat rotation has been adopted. To assess the effect of rice straw mulching on changes of antagonistic bacteria and the incidence of wheat sharp eyespot, a multi-year field study was performed to compare unmulched plots and plots mulched with rice straw for two or three years. Bacterial and fungal populations were evaluated in the cultures prepared from the wheat rhizosphere and bulk soil. Rice straw mulching increased the number of pseudomonas colony forming units (cfu) in wheat rhizosphere and bulk soil. The proportion of total bacteria that were fluorescent pseudomonads was higher in mulched than in unmulched soil. Bacterial isolates antagonistic to R. cerealis were identified using an inhibition zone test. A series of these isolates were typed by partial sequencing of the16S rRNA gene. Pseudomonads had higher antagonistic activity against R. cerealis than other species, and more than80%of rhizosphere fluorescent pseudomonads were antagonistic to R. cerealis. The disease indices in the mulched plots were lower than in the unmulched control. These suggest that rice straw mulching in a rice-wheat rotation increases the number of fluorescent pseudomonads. Additionally, these fluorescent pseudomonads may contribute to the control of wheat sharp eyespot.
为此,首先从黄淮麦区的河南、山东、安徽和江苏4省采集和分离了104株小麦全蚀病病菌,采用GGT-RP:NS5和GGA-RP:NS5两对特异性引物对病菌基因组DNA进行PCR扩增,从而鉴定这些菌株的变种类型。结果表明,所分离菌株都为禾顶囊壳小麦变种。对3种小麦全蚀病菌基因型的分析方法进行了比较,发现ITS序列分析的方法是禾顶囊壳小麦变种下分型的可靠方法。对核糖体rDNA部分序列进行的系统发育学分析结果表明,根据rDNA ITS1区的序列,供试菌株可划分为两个基因型,与国外小麦全蚀病菌基因型的分类描述有所不同,我们暂将这两个基因型命名为Q1和Q2型。所分离菌株中,Q1型菌数量高于Q2型。河南菌株中,2010年和2011年两种基因型菌株的比例不同,这种变化与病害发展和防控措施的实行有无关系还需进行进一步的研究。
为了解该区域病菌的致病力,首先比较了小麦全蚀病菌致病力的3种接种测定方法和2种病情分级标准。比较结果表明,以蛭石为培养基质、麦种下2cm接种菌丝块的接种方法和0-6级的分级标准,可比较有效和准确的测定小麦全蚀病菌的致病力。对2010年和2011年分离自黄淮麦区的104株小麦全蚀病菌进行了致病力测定,结果表明供试菌株的致病力存在明显差异。不同地区菌群间致病力差异不显著,Q1型和Q2型菌株间的致病力没有显著差异。
利用公共的真菌基因组数据库资源,对小麦全蚀病病菌G. graminis及与其亲缘关系较近的M. oryzae和M. poae三种真菌基因组中SSRs的结构类型、分布、丰度及最长序列等进行了系统分析并做了比较。结果表明：这三种真菌基因组中的SSRs非常丰富,分别为13221、9977和11422个。编码基因的序列中二、四、五核苷酸重复基序的SSRs丰富度要低于全基因组中SSRs的丰富度,而三、六核苷酸重复基序的SSRs丰富度要高于全基因组中SSRs的丰富度。G. graminis基因组中二、三、四、五、六核苷酸重复基序的SSRs丰富度要远高于M. oryzae和M. poae基因组中的同类SSRs,说明G. graminis基因组可能具有较高的变异度。本研究对禾谷顶囊壳小麦变种基因组中SSRs的丰度、出现频率及最长基序的分析为快速、便捷地设计多态性丰富的SSRs引物提供了有益的信息。
在对已公布的禾顶囊壳小麦变种菌株R3-111a-1基因组中SSR组成和结构分析的基础上,选取不同核苷酸重复方式的较长重复基序,根据其两端的保守序列设计17条引物。利用2010年分离菌株对这些引物进行筛选,获得4对扩增多态性较好的引物G7、G9、G14、G17。这4个SSR标记共可以获得50个等位基因,每个基因座的等位基因数为2-7个,平均数为5个。应用以上的4个SSR标记对2010年从黄淮麦区分离的52个菌株进行了遗传分析。4个标记可把52个菌株分为两个大的簇。群体遗传分析的结果表明,江苏病菌群体的遗传多样性水平最高,其次为山东群体,河南和安徽群体的多样性水平最低。84%的变异来自于群体内,只有16%的变异来自于群体间,4个群体间的基因流(Nm)为2.604。以安徽和山东群体的遗传相似性最高。群体分析的结果说明,安徽和河南的小麦全蚀病菌可能分别来自于山东病区。
杀菌剂种子处理是目前小麦全蚀病防治的主要方法,为了筛选高效安全的药剂。首先测定了多种杀菌剂对小麦全蚀病菌的抑菌活性,并对杀菌剂在不同浓度下的防效以及对小麦的安全性进行了盆栽试验。室内毒力测定结果表明：甾醇脱甲基化抑制剂类杀菌剂(DMIs)对小麦全蚀病病菌菌丝生长有显著的抑制作用,其中氟喹唑、氟环唑、三唑醇、咪鲜胺、戊唑醇和苯醚甲环唑的平均EC50值分别为0.03,0.03,0.06,0.07,0.10和0.24mg/L。盆栽试验结果表明：硅噻菌胺和苯醚甲环唑在推荐浓度下对小麦全蚀病有很好的防治效果,并且对小麦出苗、苗高没有显著影响；咯菌腈的防效与苯醚甲环唑相似,但对小麦出苗有轻微的抑制作用；其它几种杀菌剂对小麦全蚀病也能起到很好的防治效果,但是对小麦的生长有一定的抑制作用。从防治效果、对小麦安全性以及抗药性治理方面综合考虑,将硅噻菌胺、苯醚甲环唑和咯菌腈等交替或混合使用是防治小麦全蚀病比较好的策略。
纹枯病是长江中下游小麦上的重要病害,稻秸秆还田是该地区广泛推广的农业栽培措施。为了解稻秸秆还田对小麦根围拮抗细菌和小麦纹枯病发生的影响,本研究通过多年的田间试验,利用基于培养的方法比较了稻秸秆还田2年和3年小区小麦根围和非根围的细菌和真菌群体。结果表明,稻秸秆还田可增加小麦根围和非根围的产荧光假单胞菌菌落形成单位的数量。稻秸秆还田土壤中的产荧光假单胞菌在总细菌中的比例高于非覆盖土壤。利用平板对对峙法测定了分离细菌菌株对禾谷丝核菌Rhizoctonia cerealis的拮抗作用,并根据16s rDNA的部分序列对菌株进行了鉴定。测定结果表明,与其他种的细菌相比,假单胞杆菌对禾谷丝核菌具有较强的拮抗作用,80%以上的小麦根围发荧光假单胞菌对禾谷丝核菌具有拮抗作用。稻秸秆还田小区的小麦纹枯病病情指数低于未还田小区。研究结果表明,稻麦轮作地区的稻秸秆还田可以增加土壤中的产荧光假单胞菌的群体数量,同时产荧光假单胞菌可以对小麦纹枯病的发生起控制作用。
The incidence of take-all caused by Gaeumannomyces graminis var. tritici (Ggt) and sharp eyespot caused by Rhizoctonia cerealis has been increasing in wheat growing areas of Huang-Huai and lower reaches of Yangtze river and becoming the important diseases of wheat recently. In order to predict and control the occurrence of wheat take-all, it is necessary to understand the genetic structure of the pathogen and the trend of population structure changes. It is also urgently needed to find the effective fungicides and other measures to prevent the disease.
In the present study,104isolates of G graminis collected from Henan, Shandong, Anhui, and Jiangsu province in Huang-Huai wheat growing area were identified by amplifying the genomic DNA with two specific primers GGT-RP:NS5and GGA-RP:NS5, respectively. The results showed that all the isolates were G graminis var. tritici (Ggt) genotype. ITS sequencing was the most reliable method to analyze Ggt genotype. All tested isolates were distributed into two main genetic groups, named as Q1and Q2, respectively. Both of which were different from any isolates out of China. The genetic frequency in Q1was higher than that of in Q2. Q1/Q2frequencies in Henan province were different during2years. What resulted in the diffenence of Q1/Q2frequency in Henan province needs further study.
To determine the virulence of Gaeumannomyces graminis isolates, three inoculation and two investigation methods were compared. The results suggested that the method with fungal-colonized agar plugs2cm below the wheat seeds combined with the0-6disease grade standard was better than the others, which presented appropriate disease severity and less standard error. The virulence of104G. graminis isolates collected from Henan, Jiangsu, Anhui, and Shandong province in2010and2011was then tested using the method mentioned above. The results showed that the virulence was different significantly among the isolates. There is no significant difference on virulence between populations collected from different provinces. The virulence between genotypes Q1and Q2has also no significant difference.
Base on the public genome database, we examined the SSRs in the completely sequenced G. graminis var. tritici genome. The occurrences, relative abundance, relative density, the commonest, and the longest SSRs were analyzed, and compared with other plant pathogenic fungal species:Magnaporthe oryzae and M. poae. The results revealed that the SSRs were abundant in these three genomes, total of13221,9977and11422respectively. In the gene sequences, the Di-, Tetra-and Pentanucleotide repeats were less abundant than in the total genome sequences, while the Tri-and Hexanucleotide repeats were more abundant than in the total genome sequences. In the G. graminis var. tritici genome sequence, Di-, Tri-, Tetra-, Penta-and Hexanucleotide repeats were more abundant than the same repeats in M. oryzae and M. poae genome sequences, indicated that the G. graminis var. tritici maybe is a species with more genetic diversity. The analysis in this study provided new information that could be useful for a variety of applications in population genetics of G. graminis var. tritici.
For analysis of genetic diversity and genetic structure of the take-all pathogen, based on the analysis of simple sequence repeats in genomes of Gaeumannomyces graminis var. tritici, we isolated longer SSR motifs of different repeat types and designed17primers for screening. Four SSRs markers with reproducible patterns and diversity between different Ggt isolates were developed. Their usefulness for gentic analysis was assessed on52isolates collected in2010from Huang-Huai wheat growing areas. Four markers displayed each from2to7alleles, with an average of5alleles per locus. Phylogenetic analysis based on these markers showed that55isolates could be clustered into2large clusters. Jiangsu population showed the highest level of genetic diversity, followed by Shandong, Henan and Anhui population.84%genetic variance was within populations. The gene flow (Nm) among populations was2.604.The genetic identities between Anhui and Shandong population was the highest.
In order to screening the effective and safe fungicides for the control of wheat take-all, the inhibitory activity of different fungicides in vitro against the pathogen Gaeumammomyces graminis causing wheat take-all was tested, and control efficacy and safety of fungicides at different concentrations were also investigated in pot experiments. In vitro, sterol demethylation inhibitors fungicides (DMIs) exhibited better inhibitory activity in hypha growth of G. graminis, and the EC50values of fluquinconazole, flusilazole, riadimenol, prochloraz, tebuconazole and difenoconazole were0.03,0.03,0.06,0.07, 0.10and0.24mg/L, respectively. The results of pot experiments indicated that silthiopham and difenoconazole exhibited good control efficacy in controlling wheat take-all, and did not affect the emergence and height of seedlings. The control efficiency of fludloxonll was similar to that of difenoconazole, but slightly inhibited the emergence of seedling. The others also provided good control efficacy, but significantly inhibited the growth of the wheat seedlings. Taken together with the aspects of safety to wheat and resistance management, silthiopham, difenoconazole and fludloxonll were considered suitable for the control of wheat take-all, and should be used alternatively.
Wheat sharp eyespot caused by Rhizoctonia cerealis is one of the main diseases in the lower reaches of the Yangtze River, where rice-wheat rotation has been adopted. To assess the effect of rice straw mulching on changes of antagonistic bacteria and the incidence of wheat sharp eyespot, a multi-year field study was performed to compare unmulched plots and plots mulched with rice straw for two or three years. Bacterial and fungal populations were evaluated in the cultures prepared from the wheat rhizosphere and bulk soil. Rice straw mulching increased the number of pseudomonas colony forming units (cfu) in wheat rhizosphere and bulk soil. The proportion of total bacteria that were fluorescent pseudomonads was higher in mulched than in unmulched soil. Bacterial isolates antagonistic to R. cerealis were identified using an inhibition zone test. A series of these isolates were typed by partial sequencing of the16S rRNA gene. Pseudomonads had higher antagonistic activity against R. cerealis than other species, and more than80%of rhizosphere fluorescent pseudomonads were antagonistic to R. cerealis. The disease indices in the mulched plots were lower than in the unmulched control. These suggest that rice straw mulching in a rice-wheat rotation increases the number of fluorescent pseudomonads. Additionally, these fluorescent pseudomonads may contribute to the control of wheat sharp eyespot.
引文
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