四川松针散斑壳菌的种群结构及其遗传分析
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摘要
松树是北半球最重要的森林树种,在中国的分布遍及全国,包括引进栽培种在内共有38种12变种,四川是全国松树种类最多的省区,有7种2变种,主要有云南松、马尾松、华山松、油松、高山松、湿地松、白皮松、思茅松等。四川林区是青藏高原东部林区的重要组成部分,其森林植被是维系长江流域生态平衡的主要天然屏障。松落针病也是世界常见病害之一,主要危害松树苗木、幼林和成林,轻者引起松树部分松针枯死、脱落,重者可引起70%以上松针枯死脱落,严重影响松树的生长,材积损失率可高达80%,甚至导致松树整株枯死。松落针病也是我国松树林区常见病害,分布广,危害重,长期以来受到学者的高度重视。引起松落针病的病原菌是散斑壳属真菌,该属为斑痣盘菌科中最大一个属,己报道有100多个种,可在多种针叶树和阔叶树上寄生。由于该属真菌分布地域广,寄主种类多,其系统发育及种的分类鉴定也备受人们关注,松树生散斑壳菌的研究也是国内外学者研究的重点。本文是历经多年对四川松树落针病进行调查分析,并着重对松落针病病原菌的种类及其遗传多样性、系统发育关系进行全面系统的研究,以期完善我国松生散斑壳属真菌的研究、为维护四川林区森林健康及松落针病的防控提供科学依据。
1.参照Minter提出的松生散斑壳菌形态分类依据,对采自四川近20个市县的样品进行分类鉴定,共鉴定出散斑壳属真菌7种,即针叶树散斑壳(Lophodermium conigenum)、松针散斑壳(LophodermY pinastri)、四川散斑壳(Lophodermium sJchuanense)、印度散斑壳(Lophodermium indianum)、南方散斑壳(Lophodermium australe)、二郎山散斑壳(Lophodermium erlangshanense)(待发表新种)、库茂恩散斑壳(Lophodermiumkumaunicum)。并根据主要形态分类特征提出了四川松生散斑壳菌种的分类检索表。
2.以随机采取400颗松针统计针叶上是否有子囊果的方法,对来自四川松树林区29个采样点的样品进行统计分析表明,四川松树林区均有不同程度的落针病发生,其发病率可高达50%以上;针叶树散斑壳、松针散斑壳2个种在云南松、马尾松、湿地松、高山松、华山松上均有出现,且常有混生现象,在云南松、湿地松和马尾松上落地针叶上均有印度散斑壳菌存在,而南方散斑壳菌只在湿地松针叶上出现,库曼散斑壳菌只在高山松上出现,但数量不多;四川散斑壳菌只在二郎山云南松活针叶上有发现,二郎山散斑壳菌只在二郎山和康定跑马山的华山松活针叶和脱落针叶上有发现,数量较少。
3.采用子囊果组织分离法对采自四川各地松针散斑壳菌材料进行分离培养,共获得菌株24个(编号L01至L-24),并对其培养菌落形态特征进行了描述。通过降低养分浓度、改变碳源、更换不同种类的培养基等方法对生长于PDA平板上的散斑壳菌株进行子囊孢子诱发实验,23℃条件下培养60d后均无有性态产生。
4.采用改良二次沉淀法和改良氯化苄法提取散斑壳菌培养物总DNA,用于RAPD分析和ITS测序,其结果证明改良后的二种DNA提取法均能满足分析测试要求,改进方法是:在DNA提取之前,将各菌株培养物的菌丝体放入-20℃冰箱冷冻保存12小时以上,然后将冷冻后的菌丝体置于研钵中研磨,再加入相应的提取液。
5.从160条RAPD引物中通过初筛、复选,共筛选出11个引物用于24个散斑壳菌株的RAPD分析,并对PCR反应条件进行了优化,其最佳反应条件是:25μlPCR反应体积,10×Taq酶缓冲液(2.5μl),1.5UTaq酶,20ng DNA模板,0.4 mmol/LdNTPs,4.0mmol/LMg~(2+),0.48μmol/L引物,10.2μlddH_2O。
6.利用11个引物对24个菌株进行RAPD分析,共扩增出217个位点,其中多态性位点214个,占总数的98.6%,表明四川散斑壳菌遗传多样性非常丰富。对扩增结果采用UPGMA法进行聚类分析得到24个菌株的相似性系数聚类图。聚类图显示,在相似性系数为0.28处,所有的供试菌株被分为Ⅰ、Ⅱ、Ⅲ、Ⅳ和Ⅴ5类。其中,第1类又可分为Ⅰ-1和Ⅰ-2两个亚类:Ⅰ-1类包含了形态学上具有较多共同特征的L.pinastri、L.conigenum两个种。第Ⅳ类和第Ⅴ类因相似率很低而与其它类群分开,同样也应证了分子生物学与形态学的相关性。将寄主同为马尾松的L.conigenum各菌株单独进行分析,聚类结果显示同一种内各菌株由于地域分隔而产生较大的遗传分化;对来源于泸定二郎山林场的L.pinastri和L.conigenum种内各菌株构建相似性系数矩阵,结果表明,寄主来源不同可能造成种内不同程度的遗传分化。
7.利用真菌通用引物ITS_1、ITS_4成功对18个四川松上散斑壳菌株进行了rDNA的ITS区碱基序列分析,其结果已被GenBank接受注册(AY422489,AY422490,AY515318,AY515319,EU696766,EU696767,EU696768,EU696769,EU696770,EU696771,EU696772,EU696773,EU696774,EU696775,EU696776,EU696777,EU696778,EU679661)。采用DNAstar软件对ITS区序列结果进行多序列比较分析,构建系统发育树。与RAPD分析结果相似,18个菌株可以分成5大组(类)。与Genbank中已注册的欧美国家的9株散斑壳菌ITS区序列结果进行比对分析,其同源率最高为92.9%,四川散斑壳与欧美地区散斑壳菌的同源率较低,仅为70%至80%。
综上所述,由于四川林区地理环境复杂、松树种类多,其松落针病病原散斑壳菌多达7种,其中2种为四川报道新种,种间及种内遗传多样性丰富。其形态分类结果与RAPD、rDNA的ITS区碱基序列分析结果有较大的一致性。如何利用现代分子标记技术等手段进行松落针病的病原判定、病害早期诊断和防控是需进一步深入研究的课题。
Pine tree is one of the most important forestry species in the Northern Hemisphere.In China,there are 38 species and 12 variaties of pine tree including introduced cultivars,and their distribution is all over the entire country.Sichuan has the highest pine tree species among all of other provinces,main species are Pinus.yunnanensis,P massoniana,P. armandii,P.tabulaeformis,P.densata,P.elliottii,P.bungeanae,and P.kesiya var. langbianensis,etc.Sichuan forestry region is an important part of The Tibetan Plateau forestry, and its vegetation is the essential natural barrier to balance the ecosystem of the Yangtze River watershed area.
The pine needle-shedding is a common disease throughout the world,which harms all seedlings,nurseries and mature forests.This disease can cause different levels of damage to pine trees from partially needles shed and fall to loosing up to 70%of the total needles.The loosing of the needles can severely damage the growth of the pine tree,decrease the timber volume to up to 80%,and even result in the death of the entire pine tree.Due to the wide-range affecting areas and the severe consequences,attentions have been paid largely on this disease.
The pathogenic fungi which cause the pine needle-shedding disease belong to the genus Lophodermiun within the fungi family Rhytismataceae.More than 100 varieties have been reported in this species on both conifers and broadleaves.Due to its wide ranges of distribution and host plants,researches have been focusing on the variety identification and its development systems.This research based on multiple years investigation on the pine needle-shedding disease in Sichuan,and emphasized on identification of the virulent varieties, genetic diversity,and their systematic development relationships.The results from this research can provide the scientific basis to prevent the pine needle-shedding disease and maintain the health of Sichuan forestry region.
1.Based on the morphological classifications of Lophodermium on pine by Minter (1981),7 Lophodermium fungi from samples collected more than 20 geographic locations in Sichuan are identified.They are Lophodermium conigenum,Lophodermi pinastri, Lophodermium sichuanense,Lophodermium indianum,Lophodermium austral, Lophodermium erlangshanense(new species going to register),and Lophodermium kumaunicum.Based on the morphological traits,a classification searching index of Lophodermium on pine in Sichuan was also developed:
2.By the identification of the existing ascocarp on pine needle from 400 randomly selected pine trees in 29 different forestry regions in Sichuan province,the pine tree needle-shedding disease were found from all of the locations surveyed in this research,and the infection levels could be as high as 50%.The pathogen Lophodermium conigenum and Lophodermi pinastri existed on five pine tree cultivars P.yunnanensis,P massoniana,P. elliottii,P.densata,and P.armandii,and frequently growing together.Pathogen Lophodermium indianum existed on the needles collected from the ground of pine tree P.yunnanensis,P.elliottii,and P massoniana.Pathogen Lophodermium australe was only found growing on the needles of pine tree P.elliottii,and pathogen Lophodermium kumaunicum only appeared on pine tree P.densata as scarce events.It was also discovered that pathogen Lophodermium sichuanense only appeared on living needles of P.yunnanensis in Mount Erlang.Pathogen Lophodermium erlangshanense were found on both living or fallen needles of P.armandii with small amount on Mount Erlang and Mount Paoma in Kangding.
3.Ascocarp from fungi Lophodermium collected from various geographic locations in Sichuan were separated and cultivated.Totally 24 strains(No.from L-01 to L-24) were obtained and their morphological traits were described based on each colony.Various cultivation conditions were analyzed including nutrient concentrations,carbon sources and types of culture media,and the germination and growing of Lophodermium were assessed on PDA plates.No sexual organs were found after 60 days cultivation at 23℃.
4.Total DNA was isolated from the cultivated Lophodermium by two methods adapted from twice deposition and benzyl chloride extraction for RAPD analysis and ITS sequencing. Both of the methods proved to be effective in isolating DNA for this purpose.The essential steps of successfully extraction DNA were before DNA extraction,freeze the mycelium in-20℃for 12h in refrigerator,then homogenize the frozen mycelium in mortar,and then add corresponding extracting solutions.
5.After cycles of screening,11 primer pairs were selected from initial 160 primer pairs for RAPD analysis of 24 Lophodermium strains.The PCR reaction conditions were also optimized and the best conditions were:25μl PCR reaction volume,10×Taq enzyme buffer (2.5μl),15UTaq enzyme,20ng DNA template,0.4 mmol/L dNTPs,4.0 mmol/L Mg~(2+), 0.48μmol/L primers,and 10.2μl ddH_2O.
6.The results from the RAPD analysis of 24 strains of Lophodermium by using 11 primer pairs found 217 sites were amplified.Among the 217 bands,214 sites were indentified as polymorphisms,which accounted for 98.6%of total amplified sites,and evidenced the abundant genetic diversity of Lophodermium in Sichuan.The clustering chart of the 24 strains of Lophodermium was developed based on the coefficients of comparability by using UPGMA.It demonstrated that when comparability coefficient is 0.28,all the testing strains could be divided intoⅠ,Ⅱ,Ⅲ,Ⅳ,andⅤtypes,and typeⅠcould be further divided into subtypeⅠ-1 andⅠ-2,as subtypeⅠ-1 contains L.pinastri and L.conigenum which shares multiple morphologic traits.TypeⅣandⅤseparated from others due to their low affinity, and on the other hand evidenced the relationship between biology and morphology.Analysis on the fungi strains of L.conigenum which only infect the same host P massonictna found a broad genetic diversity within the same genus,and it was mainly due to the geographic differences.On the other hand,genetic diversity within same genus could also be results of different host origins,which was proved by the matrix analysis of the comparability coefficient for fungi strains from both L.pinastri and L.conigenum collected from Mount Erlang in Luding.
7.Analysis of base sequencing in ITS region of rDNA of 18 strains of pine Lophodermium in Sichuan was conducted by using fungi universal primer ITS_3 and ITS_4,and the result has registered in GenBank(AY422489,AY422490,AY515318,AY515319, EU696766,EU696767,EU696768,EU696769,EU696770,EU696771,EU696772, EU696773,EU696774,EU696775,EU696776,EU696777,EU696778,EU696779). Multi-sequence analysis of ITS region was conducted by using DNAstar,and the systematic development tree was constructed.Similar to the RAPD results,18 strains could be divided into 5 groups.Compared with 9 Lophodermium's ITS region registered in GenBank from Mexico,USA,and Finland,the homologous rate reached up to 92.9%.The homologous rate between Lophodermium sichuanense and Lophodermium from European is low and only ranged from 70%to 80%.
Above all,due to the complex geological environment in Sichuan forest and the diverse species of pine,the lophodermium on pine needle contained 7 genuses,and 2 of them are new species reported from Sichuan.Genetic diversity was largely observed within or between species.The classifications based on results from morphological traits,RAPD,and base sequencing analyses in ITS region of rDNA are consistent.How to utilize the modern technology of molecular marker to identify the pathogen of pine needle cast,to diagnosis in early stage,and to provide scientific basis of disease prevention and control are the future emphases that need to be studied further.
1.参照Minter提出的松生散斑壳菌形态分类依据,对采自四川近20个市县的样品进行分类鉴定,共鉴定出散斑壳属真菌7种,即针叶树散斑壳(Lophodermium conigenum)、松针散斑壳(LophodermY pinastri)、四川散斑壳(Lophodermium sJchuanense)、印度散斑壳(Lophodermium indianum)、南方散斑壳(Lophodermium australe)、二郎山散斑壳(Lophodermium erlangshanense)(待发表新种)、库茂恩散斑壳(Lophodermiumkumaunicum)。并根据主要形态分类特征提出了四川松生散斑壳菌种的分类检索表。
2.以随机采取400颗松针统计针叶上是否有子囊果的方法,对来自四川松树林区29个采样点的样品进行统计分析表明,四川松树林区均有不同程度的落针病发生,其发病率可高达50%以上;针叶树散斑壳、松针散斑壳2个种在云南松、马尾松、湿地松、高山松、华山松上均有出现,且常有混生现象,在云南松、湿地松和马尾松上落地针叶上均有印度散斑壳菌存在,而南方散斑壳菌只在湿地松针叶上出现,库曼散斑壳菌只在高山松上出现,但数量不多;四川散斑壳菌只在二郎山云南松活针叶上有发现,二郎山散斑壳菌只在二郎山和康定跑马山的华山松活针叶和脱落针叶上有发现,数量较少。
3.采用子囊果组织分离法对采自四川各地松针散斑壳菌材料进行分离培养,共获得菌株24个(编号L01至L-24),并对其培养菌落形态特征进行了描述。通过降低养分浓度、改变碳源、更换不同种类的培养基等方法对生长于PDA平板上的散斑壳菌株进行子囊孢子诱发实验,23℃条件下培养60d后均无有性态产生。
4.采用改良二次沉淀法和改良氯化苄法提取散斑壳菌培养物总DNA,用于RAPD分析和ITS测序,其结果证明改良后的二种DNA提取法均能满足分析测试要求,改进方法是:在DNA提取之前,将各菌株培养物的菌丝体放入-20℃冰箱冷冻保存12小时以上,然后将冷冻后的菌丝体置于研钵中研磨,再加入相应的提取液。
5.从160条RAPD引物中通过初筛、复选,共筛选出11个引物用于24个散斑壳菌株的RAPD分析,并对PCR反应条件进行了优化,其最佳反应条件是:25μlPCR反应体积,10×Taq酶缓冲液(2.5μl),1.5UTaq酶,20ng DNA模板,0.4 mmol/LdNTPs,4.0mmol/LMg~(2+),0.48μmol/L引物,10.2μlddH_2O。
6.利用11个引物对24个菌株进行RAPD分析,共扩增出217个位点,其中多态性位点214个,占总数的98.6%,表明四川散斑壳菌遗传多样性非常丰富。对扩增结果采用UPGMA法进行聚类分析得到24个菌株的相似性系数聚类图。聚类图显示,在相似性系数为0.28处,所有的供试菌株被分为Ⅰ、Ⅱ、Ⅲ、Ⅳ和Ⅴ5类。其中,第1类又可分为Ⅰ-1和Ⅰ-2两个亚类:Ⅰ-1类包含了形态学上具有较多共同特征的L.pinastri、L.conigenum两个种。第Ⅳ类和第Ⅴ类因相似率很低而与其它类群分开,同样也应证了分子生物学与形态学的相关性。将寄主同为马尾松的L.conigenum各菌株单独进行分析,聚类结果显示同一种内各菌株由于地域分隔而产生较大的遗传分化;对来源于泸定二郎山林场的L.pinastri和L.conigenum种内各菌株构建相似性系数矩阵,结果表明,寄主来源不同可能造成种内不同程度的遗传分化。
7.利用真菌通用引物ITS_1、ITS_4成功对18个四川松上散斑壳菌株进行了rDNA的ITS区碱基序列分析,其结果已被GenBank接受注册(AY422489,AY422490,AY515318,AY515319,EU696766,EU696767,EU696768,EU696769,EU696770,EU696771,EU696772,EU696773,EU696774,EU696775,EU696776,EU696777,EU696778,EU679661)。采用DNAstar软件对ITS区序列结果进行多序列比较分析,构建系统发育树。与RAPD分析结果相似,18个菌株可以分成5大组(类)。与Genbank中已注册的欧美国家的9株散斑壳菌ITS区序列结果进行比对分析,其同源率最高为92.9%,四川散斑壳与欧美地区散斑壳菌的同源率较低,仅为70%至80%。
综上所述,由于四川林区地理环境复杂、松树种类多,其松落针病病原散斑壳菌多达7种,其中2种为四川报道新种,种间及种内遗传多样性丰富。其形态分类结果与RAPD、rDNA的ITS区碱基序列分析结果有较大的一致性。如何利用现代分子标记技术等手段进行松落针病的病原判定、病害早期诊断和防控是需进一步深入研究的课题。
Pine tree is one of the most important forestry species in the Northern Hemisphere.In China,there are 38 species and 12 variaties of pine tree including introduced cultivars,and their distribution is all over the entire country.Sichuan has the highest pine tree species among all of other provinces,main species are Pinus.yunnanensis,P massoniana,P. armandii,P.tabulaeformis,P.densata,P.elliottii,P.bungeanae,and P.kesiya var. langbianensis,etc.Sichuan forestry region is an important part of The Tibetan Plateau forestry, and its vegetation is the essential natural barrier to balance the ecosystem of the Yangtze River watershed area.
The pine needle-shedding is a common disease throughout the world,which harms all seedlings,nurseries and mature forests.This disease can cause different levels of damage to pine trees from partially needles shed and fall to loosing up to 70%of the total needles.The loosing of the needles can severely damage the growth of the pine tree,decrease the timber volume to up to 80%,and even result in the death of the entire pine tree.Due to the wide-range affecting areas and the severe consequences,attentions have been paid largely on this disease.
The pathogenic fungi which cause the pine needle-shedding disease belong to the genus Lophodermiun within the fungi family Rhytismataceae.More than 100 varieties have been reported in this species on both conifers and broadleaves.Due to its wide ranges of distribution and host plants,researches have been focusing on the variety identification and its development systems.This research based on multiple years investigation on the pine needle-shedding disease in Sichuan,and emphasized on identification of the virulent varieties, genetic diversity,and their systematic development relationships.The results from this research can provide the scientific basis to prevent the pine needle-shedding disease and maintain the health of Sichuan forestry region.
1.Based on the morphological classifications of Lophodermium on pine by Minter (1981),7 Lophodermium fungi from samples collected more than 20 geographic locations in Sichuan are identified.They are Lophodermium conigenum,Lophodermi pinastri, Lophodermium sichuanense,Lophodermium indianum,Lophodermium austral, Lophodermium erlangshanense(new species going to register),and Lophodermium kumaunicum.Based on the morphological traits,a classification searching index of Lophodermium on pine in Sichuan was also developed:
2.By the identification of the existing ascocarp on pine needle from 400 randomly selected pine trees in 29 different forestry regions in Sichuan province,the pine tree needle-shedding disease were found from all of the locations surveyed in this research,and the infection levels could be as high as 50%.The pathogen Lophodermium conigenum and Lophodermi pinastri existed on five pine tree cultivars P.yunnanensis,P massoniana,P. elliottii,P.densata,and P.armandii,and frequently growing together.Pathogen Lophodermium indianum existed on the needles collected from the ground of pine tree P.yunnanensis,P.elliottii,and P massoniana.Pathogen Lophodermium australe was only found growing on the needles of pine tree P.elliottii,and pathogen Lophodermium kumaunicum only appeared on pine tree P.densata as scarce events.It was also discovered that pathogen Lophodermium sichuanense only appeared on living needles of P.yunnanensis in Mount Erlang.Pathogen Lophodermium erlangshanense were found on both living or fallen needles of P.armandii with small amount on Mount Erlang and Mount Paoma in Kangding.
3.Ascocarp from fungi Lophodermium collected from various geographic locations in Sichuan were separated and cultivated.Totally 24 strains(No.from L-01 to L-24) were obtained and their morphological traits were described based on each colony.Various cultivation conditions were analyzed including nutrient concentrations,carbon sources and types of culture media,and the germination and growing of Lophodermium were assessed on PDA plates.No sexual organs were found after 60 days cultivation at 23℃.
4.Total DNA was isolated from the cultivated Lophodermium by two methods adapted from twice deposition and benzyl chloride extraction for RAPD analysis and ITS sequencing. Both of the methods proved to be effective in isolating DNA for this purpose.The essential steps of successfully extraction DNA were before DNA extraction,freeze the mycelium in-20℃for 12h in refrigerator,then homogenize the frozen mycelium in mortar,and then add corresponding extracting solutions.
5.After cycles of screening,11 primer pairs were selected from initial 160 primer pairs for RAPD analysis of 24 Lophodermium strains.The PCR reaction conditions were also optimized and the best conditions were:25μl PCR reaction volume,10×Taq enzyme buffer (2.5μl),15UTaq enzyme,20ng DNA template,0.4 mmol/L dNTPs,4.0 mmol/L Mg~(2+), 0.48μmol/L primers,and 10.2μl ddH_2O.
6.The results from the RAPD analysis of 24 strains of Lophodermium by using 11 primer pairs found 217 sites were amplified.Among the 217 bands,214 sites were indentified as polymorphisms,which accounted for 98.6%of total amplified sites,and evidenced the abundant genetic diversity of Lophodermium in Sichuan.The clustering chart of the 24 strains of Lophodermium was developed based on the coefficients of comparability by using UPGMA.It demonstrated that when comparability coefficient is 0.28,all the testing strains could be divided intoⅠ,Ⅱ,Ⅲ,Ⅳ,andⅤtypes,and typeⅠcould be further divided into subtypeⅠ-1 andⅠ-2,as subtypeⅠ-1 contains L.pinastri and L.conigenum which shares multiple morphologic traits.TypeⅣandⅤseparated from others due to their low affinity, and on the other hand evidenced the relationship between biology and morphology.Analysis on the fungi strains of L.conigenum which only infect the same host P massonictna found a broad genetic diversity within the same genus,and it was mainly due to the geographic differences.On the other hand,genetic diversity within same genus could also be results of different host origins,which was proved by the matrix analysis of the comparability coefficient for fungi strains from both L.pinastri and L.conigenum collected from Mount Erlang in Luding.
7.Analysis of base sequencing in ITS region of rDNA of 18 strains of pine Lophodermium in Sichuan was conducted by using fungi universal primer ITS_3 and ITS_4,and the result has registered in GenBank(AY422489,AY422490,AY515318,AY515319, EU696766,EU696767,EU696768,EU696769,EU696770,EU696771,EU696772, EU696773,EU696774,EU696775,EU696776,EU696777,EU696778,EU696779). Multi-sequence analysis of ITS region was conducted by using DNAstar,and the systematic development tree was constructed.Similar to the RAPD results,18 strains could be divided into 5 groups.Compared with 9 Lophodermium's ITS region registered in GenBank from Mexico,USA,and Finland,the homologous rate reached up to 92.9%.The homologous rate between Lophodermium sichuanense and Lophodermium from European is low and only ranged from 70%to 80%.
Above all,due to the complex geological environment in Sichuan forest and the diverse species of pine,the lophodermium on pine needle contained 7 genuses,and 2 of them are new species reported from Sichuan.Genetic diversity was largely observed within or between species.The classifications based on results from morphological traits,RAPD,and base sequencing analyses in ITS region of rDNA are consistent.How to utilize the modern technology of molecular marker to identify the pathogen of pine needle cast,to diagnosis in early stage,and to provide scientific basis of disease prevention and control are the future emphases that need to be studied further.
引文
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