杨树烂皮病病原菌Cytosporaspp.的系统发育与遗传多样性研究
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摘要
本研究广泛收集中国杨树上的Cytospora spp.及其相关有性型（Valsa spp., Leucostomaspp., Valseutella spp., Valsella spp.）真菌，在依据子座和腔室排列为主要特征的形态学鉴定基础上，结合核糖体内转录间隔区间和Beta微管蛋白的部分序列同源性的分子数据进行分类鉴定和系统发育分析，并通过序列分析和RAPD分子标记对广布优势种C.chrysosperma种内遗传多样性研究，该篇研究有助于理解和认识Cytospora spp.及其有性型系统发育地位，尤其是界定该类真菌在中国杨树、柳树上的分类地位，为杨树产业中抗病树种、杂交种、无性系的选择提供重要依据，实现对该类病害的防治。
1本研究对我国黑龙江、吉林、辽宁、内蒙古、北京、四川、重庆6个省2个直辖市中的17个县(乡)杨柳树烂皮病进行了调查，对青杨派、白杨派、黑杨派3派内多个杨树树种、无性系和杂交种及分布于以上地区的柳树主要品种上的Cytospora spp.及其有性型进行了收集，共得到腐烂病标本276份，其中具典型症状的标本128份，经分离纯化，初步鉴定得到烂皮病菌Cytospora spp.菌株249株。
2对采集到的烂皮病菌进行形态学特征观察和依据分子数据进行系统发育分析。通过观察寄主材料和培养基上的子实体显微特征及菌落颜色、生长温度、子实体大小、数量、颜色等培养学特征，并依据Cytospora（Valsa）最新腔室排列方式术语，共发现4种无性型的分生孢子器类型，分别是rosette cytosporoid、labyrinthine cytosporiod、torselliod和Leucocytosporoid。将分离得到的所有来自杨树上的典型的、代表各个种形态学菌株的ITS-rDNA和β-tubulin序列与世界其他国家多个寄主上的Cytospora spp.（有性型Valsa）的参比序列进行比较，并采用MP和BI方法分别在PAUP*4.0b10和贝叶斯软件中构建系统发育树，发现供试菌株分别存在于5个已知组内和2个单系起源的独立分枝中。综合形态特征观察和分子检测结果，鉴定了我国杨树Cytospora spp.真菌的6个种，其中包括：新种1种Cytospora davidiana Y.L.Wang&X Y Zhang&Q. Lv, sp. nov.；新记录种2种C.atrocirrhata, C. kantschavelii；已知种3种C. chrysosperma，C. translucens和C. fugax及1种因菌株量少未能准确定名种Cytospora sp.1。其中杨树、柳树为C. translucens在中国的新寄主植物，C. chrysosperma为广布优势种类，菌株分离率为全部分离株的82%，在上述样品采集区均分离得到。
Cytospora davidiana sp. nov.形态学描述：产孢体子座散生，均匀分布，颜色由外而内为浅灰到深棕色，埋生在树皮内，常顶破表皮而出，形状多为卵圆形或长圆柱形，高0.2-1.2cm至1.5cm。产孢体盘深褐色，近平展，圆形至卵圆形，直径0.1-2.0mm。产孢体下部各腔具单一孔口，各孔口向中心聚集最后融合为1个中心孔口，孔口周围为灰色或灰白色内子座或无定型物质包围，常与产孢体盘顶部齐平，直径约20-50μm。产孢体类型为torselliod，多腔，腔壁无内陷，具单独腔壁，直径90-210×200-600μm。分生孢子梗无色透明，基部嵌入到胶状基质中，通常在基部或高度的1/2处分枝，产生4个小梗，大小为12-15×1.0-1.5μm；产孢细胞略呈圆筒状的分生孢子梗，顶端较尖，平周加厚，9-17×0.7-1μm。分生孢子无色透明，无隔，腊肠形，4.2-5.5×0.7-1.0μm。
3将鉴定为C. chrysosperma的标本进行分离培养，选择培养形态上有明显差异的41个代表性菌株作为种内遗传多样性分析材料，进行ITS-rDNA序列分析及RAPD分子标记。结果表明，所有供试菌株都能扩增出稳定一致的多态性条带。序列分析结果表明，各菌株ITS-rDNA序列在27个碱基位点存在差异，平均遗传距离为0.001，MP系统发育树不能将各菌株区分开。对RAPD标记的结果进行遗传多样性评估，得到多态性条带(N)79条，多态位点百分率(p)为91.86%。聚类分析结果显示相似系数在0.41水平上所有菌株被分为为4个聚类群类，该4个聚类群与依据培养学特征划分的4各组基本上是一一对应的，地理分化和寄主分化不明显。从DNA水平上证实C. chrysosperma种内存在明显的遗传分化并且具有非常宽阔的地理分布范围。RAPD分子标记技术所扩增的多态性条带表明该技术可以C. chrysosperma及Cytospora spp.及其相关有性型真菌的遗传多样性进行研究。
To confirm the phylogenetic placement and diversity of this kind fungus on Populus spp. inChina, extensive collection of Cytospora spp. and their teleomorph samples (Valsa spp.,Leucostoma spp., Valseutella spp., Valsella spp.) were gathered from Populus in China,identification and phylogeny of isolate were performed by morphological characteristicsincluding locule forms (Cytospora spp.) and stroma features (Valsa spp.) and DNA sequencehomology of the ITS-rDNA and β-tubulin partial sequences, furthermore, the genetic diversityof Cytospora chrysosperma was studyed by sequence analysis and RAPD marker. It is hopedthat to screen disease-resistant poplar varieties and create a new clue in control strategies forthis kind of pathogen in the long term.
1Samples of poplar stems and branches with canker symptoms were collected fromcultivated poplar factions including Leuce, Tacamahaca, Aigeiros, which from in the mainpoplar-producing regions of China, including Heilongjiang, Jilin, Liaoning, Inner Mongolia,Henan, Sichuan provinces as well as2municipalities of Beijing and Chongqing from2011to2012.276canker specimens were obtained, and128of these were of seriously infected byCytospora spp, strains were isolated from lesions of infected stems or branches. In total,249Cytospora spp. strains were isolated from Populus from the main poplar producing regions inChina.
2Isolates were identified based on morphology and DNA sequence homology of ITS-rDNAand β-tubulin. Microscopic structures of the fungi were examined on natural subtrates and inculture media by using the latest descriptive terms of locule forms. We found4conidiomatatypes: rosette cytosporoid, labyrinthine cytosporiod, Leucocytosporoid, torselliod. Isolatesreference were compared with the reference strains collected in other parts of the world,phylogenesis was performed in PAUP*4.0b10and Mrbayes using method of MP and BI.33representative genetically distinct sequences from249sequences of isolate residing within thepopulations of5known species and2unique lineages.1of them was described as new:Cytospora davidiana Y.L.Wang&X Y Zhang&Q. Lv, sp. nov. C. atrocirrhata and C.kantschavelii represented new reports from China, in addition,3of them corresponded with theprevious morphology-based taxa: C. chrysosperma, C. translucens, C. fugax, and remaining1phylogenetic species Cytospora sp.1was not nominated accurately due to the paucity of thematerials. Moreover, poplar and willow was found be new hosts of C. translucens in China, andC. chrysosperma was of a dominant species accounting for82%of the strains obtained in inthis study.
Description of Cytospora davidiana sp. nov.: Conidiomatal stromata immersed in bark,erumpent, ovoid to elongate cylindrical,0.2-1.2cm up to1.5cm diam. Discs dark brown,nearly flat, circular to lenticular,0.1-2.0mm diam,1-10ostiole(s). Ostiole, dark grey, oftenabove the disc surfaces,20-50μm, surrounded by pale brown entostromata of texura globose or amorphous material. Locules globose,90-210×200-600μm diam, simple undivided, not sharingcommon walls, brown walls of textura epidermoidea. Conidiophores, frequently branchedabove the base near the mid-height, with up to four verticillate phialides,12-15×1-1.5μminclusive of phialides, embedded in a continuous gelatinous matrix. Conidiogenous cellssubcylindrical phialides, tapering to the apices, with minute collarettes and periclinalthickenings,9-17×0.7-1μm. Conidia hyaline, eguttulate, allantoid, aseptate,4.2-5.5×0.7-1.0μm.
341representative isolates of C. chrysosperma confirmed by morphology and moleculardata were selected according to their obvious culture characters to study the genetic diversityusing ITS-rDNA sequence analysis and random amplified polymorphic deoxyribonucleic acid(RAPD) marker. Sequence analysis results showed that there were27different base in sequenceamplied from ITS-rDNA, the mean overal genetic distance was0.001, MP phylogenetic treecould not differentiate the strains which was clustered in one main clade. The RAPD resultdemonstrated all the random primers amplified reproducible polymorphic banding patterns. Theevaluation of genetic diversity got the79polymorphism bands, which polymorphic loci (p) wasof91.86%. Based on0.41Dice similarity coefficient, cluster analysis of the data divided theisolates into4groups which were almostly correspond to the groups subdivided by culturefeatures, showing a high genetic diversity among populations of C. chrysosperma. The resultalso demonstrated that there was no correlation between geographical origins and the resultinggroups of RAPD analysis. The amount of observed polymorphism showed by RAPD indicatedthat it was appreciate to study genetic diversity in C. chrysosperma, and could be used in otherspecies of Cytospora spp. and associated teleomorphs.
1本研究对我国黑龙江、吉林、辽宁、内蒙古、北京、四川、重庆6个省2个直辖市中的17个县(乡)杨柳树烂皮病进行了调查，对青杨派、白杨派、黑杨派3派内多个杨树树种、无性系和杂交种及分布于以上地区的柳树主要品种上的Cytospora spp.及其有性型进行了收集，共得到腐烂病标本276份，其中具典型症状的标本128份，经分离纯化，初步鉴定得到烂皮病菌Cytospora spp.菌株249株。
2对采集到的烂皮病菌进行形态学特征观察和依据分子数据进行系统发育分析。通过观察寄主材料和培养基上的子实体显微特征及菌落颜色、生长温度、子实体大小、数量、颜色等培养学特征，并依据Cytospora（Valsa）最新腔室排列方式术语，共发现4种无性型的分生孢子器类型，分别是rosette cytosporoid、labyrinthine cytosporiod、torselliod和Leucocytosporoid。将分离得到的所有来自杨树上的典型的、代表各个种形态学菌株的ITS-rDNA和β-tubulin序列与世界其他国家多个寄主上的Cytospora spp.（有性型Valsa）的参比序列进行比较，并采用MP和BI方法分别在PAUP*4.0b10和贝叶斯软件中构建系统发育树，发现供试菌株分别存在于5个已知组内和2个单系起源的独立分枝中。综合形态特征观察和分子检测结果，鉴定了我国杨树Cytospora spp.真菌的6个种，其中包括：新种1种Cytospora davidiana Y.L.Wang&X Y Zhang&Q. Lv, sp. nov.；新记录种2种C.atrocirrhata, C. kantschavelii；已知种3种C. chrysosperma，C. translucens和C. fugax及1种因菌株量少未能准确定名种Cytospora sp.1。其中杨树、柳树为C. translucens在中国的新寄主植物，C. chrysosperma为广布优势种类，菌株分离率为全部分离株的82%，在上述样品采集区均分离得到。
Cytospora davidiana sp. nov.形态学描述：产孢体子座散生，均匀分布，颜色由外而内为浅灰到深棕色，埋生在树皮内，常顶破表皮而出，形状多为卵圆形或长圆柱形，高0.2-1.2cm至1.5cm。产孢体盘深褐色，近平展，圆形至卵圆形，直径0.1-2.0mm。产孢体下部各腔具单一孔口，各孔口向中心聚集最后融合为1个中心孔口，孔口周围为灰色或灰白色内子座或无定型物质包围，常与产孢体盘顶部齐平，直径约20-50μm。产孢体类型为torselliod，多腔，腔壁无内陷，具单独腔壁，直径90-210×200-600μm。分生孢子梗无色透明，基部嵌入到胶状基质中，通常在基部或高度的1/2处分枝，产生4个小梗，大小为12-15×1.0-1.5μm；产孢细胞略呈圆筒状的分生孢子梗，顶端较尖，平周加厚，9-17×0.7-1μm。分生孢子无色透明，无隔，腊肠形，4.2-5.5×0.7-1.0μm。
3将鉴定为C. chrysosperma的标本进行分离培养，选择培养形态上有明显差异的41个代表性菌株作为种内遗传多样性分析材料，进行ITS-rDNA序列分析及RAPD分子标记。结果表明，所有供试菌株都能扩增出稳定一致的多态性条带。序列分析结果表明，各菌株ITS-rDNA序列在27个碱基位点存在差异，平均遗传距离为0.001，MP系统发育树不能将各菌株区分开。对RAPD标记的结果进行遗传多样性评估，得到多态性条带(N)79条，多态位点百分率(p)为91.86%。聚类分析结果显示相似系数在0.41水平上所有菌株被分为为4个聚类群类，该4个聚类群与依据培养学特征划分的4各组基本上是一一对应的，地理分化和寄主分化不明显。从DNA水平上证实C. chrysosperma种内存在明显的遗传分化并且具有非常宽阔的地理分布范围。RAPD分子标记技术所扩增的多态性条带表明该技术可以C. chrysosperma及Cytospora spp.及其相关有性型真菌的遗传多样性进行研究。
To confirm the phylogenetic placement and diversity of this kind fungus on Populus spp. inChina, extensive collection of Cytospora spp. and their teleomorph samples (Valsa spp.,Leucostoma spp., Valseutella spp., Valsella spp.) were gathered from Populus in China,identification and phylogeny of isolate were performed by morphological characteristicsincluding locule forms (Cytospora spp.) and stroma features (Valsa spp.) and DNA sequencehomology of the ITS-rDNA and β-tubulin partial sequences, furthermore, the genetic diversityof Cytospora chrysosperma was studyed by sequence analysis and RAPD marker. It is hopedthat to screen disease-resistant poplar varieties and create a new clue in control strategies forthis kind of pathogen in the long term.
1Samples of poplar stems and branches with canker symptoms were collected fromcultivated poplar factions including Leuce, Tacamahaca, Aigeiros, which from in the mainpoplar-producing regions of China, including Heilongjiang, Jilin, Liaoning, Inner Mongolia,Henan, Sichuan provinces as well as2municipalities of Beijing and Chongqing from2011to2012.276canker specimens were obtained, and128of these were of seriously infected byCytospora spp, strains were isolated from lesions of infected stems or branches. In total,249Cytospora spp. strains were isolated from Populus from the main poplar producing regions inChina.
2Isolates were identified based on morphology and DNA sequence homology of ITS-rDNAand β-tubulin. Microscopic structures of the fungi were examined on natural subtrates and inculture media by using the latest descriptive terms of locule forms. We found4conidiomatatypes: rosette cytosporoid, labyrinthine cytosporiod, Leucocytosporoid, torselliod. Isolatesreference were compared with the reference strains collected in other parts of the world,phylogenesis was performed in PAUP*4.0b10and Mrbayes using method of MP and BI.33representative genetically distinct sequences from249sequences of isolate residing within thepopulations of5known species and2unique lineages.1of them was described as new:Cytospora davidiana Y.L.Wang&X Y Zhang&Q. Lv, sp. nov. C. atrocirrhata and C.kantschavelii represented new reports from China, in addition,3of them corresponded with theprevious morphology-based taxa: C. chrysosperma, C. translucens, C. fugax, and remaining1phylogenetic species Cytospora sp.1was not nominated accurately due to the paucity of thematerials. Moreover, poplar and willow was found be new hosts of C. translucens in China, andC. chrysosperma was of a dominant species accounting for82%of the strains obtained in inthis study.
Description of Cytospora davidiana sp. nov.: Conidiomatal stromata immersed in bark,erumpent, ovoid to elongate cylindrical,0.2-1.2cm up to1.5cm diam. Discs dark brown,nearly flat, circular to lenticular,0.1-2.0mm diam,1-10ostiole(s). Ostiole, dark grey, oftenabove the disc surfaces,20-50μm, surrounded by pale brown entostromata of texura globose or amorphous material. Locules globose,90-210×200-600μm diam, simple undivided, not sharingcommon walls, brown walls of textura epidermoidea. Conidiophores, frequently branchedabove the base near the mid-height, with up to four verticillate phialides,12-15×1-1.5μminclusive of phialides, embedded in a continuous gelatinous matrix. Conidiogenous cellssubcylindrical phialides, tapering to the apices, with minute collarettes and periclinalthickenings,9-17×0.7-1μm. Conidia hyaline, eguttulate, allantoid, aseptate,4.2-5.5×0.7-1.0μm.
341representative isolates of C. chrysosperma confirmed by morphology and moleculardata were selected according to their obvious culture characters to study the genetic diversityusing ITS-rDNA sequence analysis and random amplified polymorphic deoxyribonucleic acid(RAPD) marker. Sequence analysis results showed that there were27different base in sequenceamplied from ITS-rDNA, the mean overal genetic distance was0.001, MP phylogenetic treecould not differentiate the strains which was clustered in one main clade. The RAPD resultdemonstrated all the random primers amplified reproducible polymorphic banding patterns. Theevaluation of genetic diversity got the79polymorphism bands, which polymorphic loci (p) wasof91.86%. Based on0.41Dice similarity coefficient, cluster analysis of the data divided theisolates into4groups which were almostly correspond to the groups subdivided by culturefeatures, showing a high genetic diversity among populations of C. chrysosperma. The resultalso demonstrated that there was no correlation between geographical origins and the resultinggroups of RAPD analysis. The amount of observed polymorphism showed by RAPD indicatedthat it was appreciate to study genetic diversity in C. chrysosperma, and could be used in otherspecies of Cytospora spp. and associated teleomorphs.
引文
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