虫草属及其无性型关系研究
摘要
针对虫草属的不同发育阶段被用于新种描述和多型现象普遍出现，导致虫草属分、无性型鉴定及其在开发应用上的混乱现象。本文采用了RAPD、rDNA ITS的RFLP、rDNA的ITS序列分析、诱发微循环产孢、扫描电镜观察、子囊孢子分离等方法研究了虫草及其他虫生真菌的100多个标本或菌株。取得的主要结果如下：
通过经典生物学和分子生物学综合研究所提供的有力证据，证明了尼泊尔虫草和多枝虫草是冬虫夏草的同物异名，冬虫夏草的无性型是中国被毛孢，其他所对应的无性型为，拟布里班克虫草--大孢绿僵菌，大蝉虫草--蝉白僵菌，布氏虫草--布氏白僵菌，古尼虫草--古尼拟青霉(=霍克斯虫草—霍克斯拟青霉)，茧虫草--茧草玛利亚霉，蛹虫草--蛹草拟青霉等，研究结果初步证明甘肃虫草和阔孢虫草也可能是冬虫夏草的同物异名。
用ITS的RFLP分析方法对一些有争议和分类地位不明确的无性型研究表明：双型孢绿僵菌和柱孢绿僵菌应归为一个种，代氏绿僵菌和贵州绿僵菌也应归为一个种。古尼拟青霉和霍克斯虫草的无性型—霍克斯拟青霉是一个种。绿僵菌属RFLP，虫草无性型RFLP和拟青霉属RFLP都能把同一属中的不同种清楚地相互区分开。
虫草属rDNA ITS碱基序列分析结果表明：以子囊壳埋生和子囊壳非埋生或半埋生将虫草分成两类。在埋生类群中，子囊壳斜埋生的种，即属于新虫草亚属的沫蝉虫草和双头虫草，以及虫草亚属的下垂虫草都被归到一个类群之中，垂直埋生的类型被归到另一类群中，在埋生类群中，除大团囊虫草和C。cantheraloides为埋生型外，其他均为非埋生型。同时也发现，不同产地的同种虫草，有的分子水平上无差异如蛹虫草，冬虫夏草，有的虽有差异但仍然被归在一组中如大蝉虫草，种与种间差异较大。为此，对虫草属的分类提出了如下新原则：在虫草属分类中，子囊壳埋生与否应为第一位的特征，这与传统的虫草分类原则不同。
同时。对西南地区10多个不同生态区或自然保护区的虫草及其他的虫生真菌进行了调查和分离研究，发现了3个虫草新种和4个半知菌新种，既茂兰虫草(Cordyceps maolanensis)，贵州虫草(C.guizhouensis)，拟布里班克虫草(C.brittlebanksoides)，蜻蜒拟青霉(Paecilomycesodonatae)，双梭隔梭孢(Septofusidium bifusisporum)，香棒弯颈霉(Tolypocladium barnesii)，蝉白僵菌(Beauviria soboliferα)。
The different developing stages of Cordyceps spp. have been used in description of new species and pleomorphism is very common in the genus Cordyceps so that there are some problems in classification of Cordyceps, identification of anamorphs, development and application of both Cordyceps and its anamorphs. Restriction fragment lenth polymorphism (RFLP) analysis n random amplified polymorphic DNA analysis [RAPD], the sequence analysis of internal transcribed spacers (ITS) of ribosomal DNA (rDNA), induction of microcyclic conidiation, SEM (scanning electron microscopy) ascosporal isolation and other methods were applied to study more than 100 specimens or isolates of Cordyceps, its anamorphs and other entomogenous fungi. The main results were following:
Based on some powerful evidence provided by using the classic and molecular biological methods it is determined that C. nepalensis and C. multiaxialis are synonym of Cordyceps sinensis, the anamorph of C. sinensis is Hirsutella sinensis and the relative relationship between anamorphs and teleomorphs of other Cordyceps spp.are Paecilomyces gunnii-- C.gunnii (Paecilomyces hawkesii~C. hawkesii), Beauveria bronginartii-C. bronginartii, Metarhizium anisopliae var. Majus-C. Brittlebanksoides, Beauveria sobolifera--C. Sobolifera, Mariannaea pruinosa--C. Pruinosa, Paecilomyces militaris--C. Militaris, Septofusidium bifusisporum-C. Bifusispora etc. C. gansuensis and C. crassospora are possibly also synonym of C. sinensis.
The results from RFLP showed Metarhizium bifusisporium and M. Cylindrica are same fungus, Metarhizium taii and M. Guizhouensis are the same, Paecilomyces gunnii and Paecilomyces hawkesii maybe are same . Based on the results of RFLP there are clear differences between species in both genera Metarhizium and Paecilomyces.
The phylogenetic relationship of Cordyceps based on the ITS sequence analysis of rDNA showed all of the species in this experiment have been divided into two groups by molecular systematic classification, group one including the species with perithecia wholly immersed, and group two including those with perithecia not or half immersed. It means the position of perithecia in the stroma is very important characteristic. In the
immersed group, 3 species with perithecia obliquely immersed are located in same subgroup, and these species with perithecia vertical immersed in the other subgroup.In the not immersed group, all of species have perithecia not immersed or half immersed but C. ophioglossoides which grows on Ellomyces and C. cantheraloides which has stroma as Agaricus have. In subgroup, there are no differences between specimens of same species from different places for instance C. militaris or a ittle differences for example C. Sobolifera.According to above analysis a new classified criterion proposed is that in classification of the genus Cordyceps whether the perithecia are immersed or not should be the first classified characteristic, which is quite different from the classic ones. At the same time based on the investigations, isolation and identification of Cordyceps spp. and other entomogenous fungi collected from more than 10 different ecological places or natural preserves in southwest China. Seven new species includin
g 3 species of Cordyceps and 4 species of mitosporic fungi are described. They are Cordyceps maolanensis sp. Nov., C. guizhouensis sp.nov, C. brittlebanksoides sp. nov., Paecilomyces odonatae sp. nov., Septofusidium bifusisporum sp. nov., Tolypocladium barnesii sp. nov.,and Beauviria sobolifera sp. nov..
通过经典生物学和分子生物学综合研究所提供的有力证据，证明了尼泊尔虫草和多枝虫草是冬虫夏草的同物异名，冬虫夏草的无性型是中国被毛孢，其他所对应的无性型为，拟布里班克虫草--大孢绿僵菌，大蝉虫草--蝉白僵菌，布氏虫草--布氏白僵菌，古尼虫草--古尼拟青霉(=霍克斯虫草—霍克斯拟青霉)，茧虫草--茧草玛利亚霉，蛹虫草--蛹草拟青霉等，研究结果初步证明甘肃虫草和阔孢虫草也可能是冬虫夏草的同物异名。
用ITS的RFLP分析方法对一些有争议和分类地位不明确的无性型研究表明：双型孢绿僵菌和柱孢绿僵菌应归为一个种，代氏绿僵菌和贵州绿僵菌也应归为一个种。古尼拟青霉和霍克斯虫草的无性型—霍克斯拟青霉是一个种。绿僵菌属RFLP，虫草无性型RFLP和拟青霉属RFLP都能把同一属中的不同种清楚地相互区分开。
虫草属rDNA ITS碱基序列分析结果表明：以子囊壳埋生和子囊壳非埋生或半埋生将虫草分成两类。在埋生类群中，子囊壳斜埋生的种，即属于新虫草亚属的沫蝉虫草和双头虫草，以及虫草亚属的下垂虫草都被归到一个类群之中，垂直埋生的类型被归到另一类群中，在埋生类群中，除大团囊虫草和C。cantheraloides为埋生型外，其他均为非埋生型。同时也发现，不同产地的同种虫草，有的分子水平上无差异如蛹虫草，冬虫夏草，有的虽有差异但仍然被归在一组中如大蝉虫草，种与种间差异较大。为此，对虫草属的分类提出了如下新原则：在虫草属分类中，子囊壳埋生与否应为第一位的特征，这与传统的虫草分类原则不同。
同时。对西南地区10多个不同生态区或自然保护区的虫草及其他的虫生真菌进行了调查和分离研究，发现了3个虫草新种和4个半知菌新种，既茂兰虫草(Cordyceps maolanensis)，贵州虫草(C.guizhouensis)，拟布里班克虫草(C.brittlebanksoides)，蜻蜒拟青霉(Paecilomycesodonatae)，双梭隔梭孢(Septofusidium bifusisporum)，香棒弯颈霉(Tolypocladium barnesii)，蝉白僵菌(Beauviria soboliferα)。
The different developing stages of Cordyceps spp. have been used in description of new species and pleomorphism is very common in the genus Cordyceps so that there are some problems in classification of Cordyceps, identification of anamorphs, development and application of both Cordyceps and its anamorphs. Restriction fragment lenth polymorphism (RFLP) analysis n random amplified polymorphic DNA analysis [RAPD], the sequence analysis of internal transcribed spacers (ITS) of ribosomal DNA (rDNA), induction of microcyclic conidiation, SEM (scanning electron microscopy) ascosporal isolation and other methods were applied to study more than 100 specimens or isolates of Cordyceps, its anamorphs and other entomogenous fungi. The main results were following:
Based on some powerful evidence provided by using the classic and molecular biological methods it is determined that C. nepalensis and C. multiaxialis are synonym of Cordyceps sinensis, the anamorph of C. sinensis is Hirsutella sinensis and the relative relationship between anamorphs and teleomorphs of other Cordyceps spp.are Paecilomyces gunnii-- C.gunnii (Paecilomyces hawkesii~C. hawkesii), Beauveria bronginartii-C. bronginartii, Metarhizium anisopliae var. Majus-C. Brittlebanksoides, Beauveria sobolifera--C. Sobolifera, Mariannaea pruinosa--C. Pruinosa, Paecilomyces militaris--C. Militaris, Septofusidium bifusisporum-C. Bifusispora etc. C. gansuensis and C. crassospora are possibly also synonym of C. sinensis.
The results from RFLP showed Metarhizium bifusisporium and M. Cylindrica are same fungus, Metarhizium taii and M. Guizhouensis are the same, Paecilomyces gunnii and Paecilomyces hawkesii maybe are same . Based on the results of RFLP there are clear differences between species in both genera Metarhizium and Paecilomyces.
The phylogenetic relationship of Cordyceps based on the ITS sequence analysis of rDNA showed all of the species in this experiment have been divided into two groups by molecular systematic classification, group one including the species with perithecia wholly immersed, and group two including those with perithecia not or half immersed. It means the position of perithecia in the stroma is very important characteristic. In the
immersed group, 3 species with perithecia obliquely immersed are located in same subgroup, and these species with perithecia vertical immersed in the other subgroup.In the not immersed group, all of species have perithecia not immersed or half immersed but C. ophioglossoides which grows on Ellomyces and C. cantheraloides which has stroma as Agaricus have. In subgroup, there are no differences between specimens of same species from different places for instance C. militaris or a ittle differences for example C. Sobolifera.According to above analysis a new classified criterion proposed is that in classification of the genus Cordyceps whether the perithecia are immersed or not should be the first classified characteristic, which is quite different from the classic ones. At the same time based on the investigations, isolation and identification of Cordyceps spp. and other entomogenous fungi collected from more than 10 different ecological places or natural preserves in southwest China. Seven new species includin
g 3 species of Cordyceps and 4 species of mitosporic fungi are described. They are Cordyceps maolanensis sp. Nov., C. guizhouensis sp.nov, C. brittlebanksoides sp. nov., Paecilomyces odonatae sp. nov., Septofusidium bifusisporum sp. nov., Tolypocladium barnesii sp. nov.,and Beauviria sobolifera sp. nov..
引文
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