耐热真菌多样性及分子系统学研究
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摘要
耐热真菌(thermotolerant fungi)是一类最高生长温度在50℃左右(通常为40℃或40℃以上),最低生长温度在20℃以下的特殊真菌类群。耐热真菌在高温条件下表现出独特的生存适应能力,其特殊的机制及特殊的产物,将使某些新的生物技术手段成为可能,是奠定高效率、低成本生物技术、新工艺的基础。我国耐热真菌研究比较薄弱,未进行系统研究,已报道耐热真菌23种,与国际上已经报道的89种耐热真菌的数量有较大差距。我国存在耐热真菌分布和生存的各种气候条件和生境条件,其多样性应该丰富。通过对我国不同气候区域和生境的耐热真菌种类及分布进行研究,以明确我国这一类群的真菌资源,将为我国高温真菌的理论和应用研究以及分类专著性研究奠定基础。
2009-2011年在山东、山西、河北、陕西、河南、云南、江西、海南、新疆、内蒙古、甘肃、广东、吉林、湖南等地采集堆肥、培养料、草堆、腐殖质、植物残体、木屑堆、土壤、动物巢穴等标本800余份,生态类型包括森林、草原、农田、沙漠、荒地、湿地、高山草甸等,采用稀释平板法、土壤平板法等分离培养,结合形态特征观察与rDNA-ITS序列分析,分离鉴定耐热真菌25属51种,包括新种2个,中国新记录种11个,还有15个首次报道为耐热真菌。所研究的属、种均作详细形态描述和地点、生境引证。研究过的标本及活菌种保存在山东农业大学植物病理学标本室(HSAUP)。
一、系统研究了我国不同生态类型耐热真菌的种类及多样性
2个新种：井冈带孢霉Taeniolella jinggangensis Y.Zhang&D.C.Li,耐热盾盘菌Scutellinia subthermophila Y.Zhang&D.C.Li;11个中国新记录种：瘤突毛壳Chaetomium strumarium,耐热梭孢壳Thielavia subthermophila,榛色钩囊菌Hamigera avellanea,草莓状炭团菌Hypoxylon fragiforme,芬尼新萨托菌Neosartorya fennelliae,加纳木霉Trichoderma ghanense,焦特普尔毛壳Chaetomium jodhpurense,黄色毛壳Chaetomium luteum,土星孢木霉Trichoderma saturnisporum,沙栖梭孢壳Thielavia arenaria,棕色毛壳Chaetomium atrobrunnneum;38个其他耐热菌为国内已知种,其中15个首次报道为耐热真菌,包括：刺孢小克银汉霉Cunninghamella echinulata,卵形孢球托霉Gongronella butleri,卷枝毛霉Mucor circinelloides,金色毛壳Chaetomium aureum,四脊裸胞壳Emericella quadrilineata,爪哇正青霉Eupenicillium javanicum,粪生粪壳菌Sordaria fimicola,焦曲霉Aspergillus ustus,新月弯孢Curvularia lunata,嘴突凸脐孢Exserohilum rostratum,棕黑腐殖霉Humicola fuscoatra,尖端赛多孢子菌Scedosporium apiospermum,绳状青霉Penicillium funiculosum,疣孢青霉Penicillium verruculosum,灰拟鬼伞菌Coprinopsis cinerea。调查研究的结果增进了对我国耐热真菌种类的了解,丰富了我国生物多样性的内容,并为我国耐热真菌的理论研究和应用研究以及分类专著性研究奠定了基础。
二、研究总结耐热真菌种级分类标准
目前形态学特征仍然是耐热真菌分类的主要依据。对于同属内的耐热真菌种类,尽量在相同的条件下培养和形态比较,尽量减少外部环境变化或不稳定对形态的影响。耐热性(最高温度、最适温度、最低温度)是同属内耐热与非耐热真菌种级鉴别的重要依据。测试耐热真菌温度生长范围间隔为5℃。
对于半知菌等无性态耐热真菌,分生孢子的特征(包括形状、大小、颜色、隔膜数和表面纹饰等)是种级分类的主要依据；其次,分生孢子梗的形态特征、产孢细胞的延伸方式及产孢方式对属级水平上的鉴别有重要意义。对于接合菌、子囊菌等有性态耐热真菌：主要以有性结构的形态特点作为种级分类的依据,如子囊果大小,形状、颜色,有无附属丝,附属丝形状,子囊大小,形状,子囊孢子大小,形状,颜色、有无萌发孔及萌发孔个数等。当子囊菌培养条件下多产生无性态,而有性态只有在特殊条件下才能产生且出现较晚,无性态命名优先采用,保证在应用研究中耐热真菌名称的唯一性和正确性。培养特征(菌落特征、颜色以及生长特性)及产孢表型可作为分类鉴定的必要、有益参考。
三、耐热真菌分子系统学研究
本研究在形态分类的基础上,对所分离鉴定的耐热真菌,进行5.8S rDNA-ITS序列和部分Mn-SOD中间片段序列测定。测得的序列提交美国生物信息中心(NCBI)公共数据库GenBank,并获得相应的注册号。
5.8S rDNA-ITS序列能有效区分大多数不同属、种的耐热真菌,序列属内种间相似性明显大于属间相似性；基于5.8S rDNA-ITS序列构建的系统发育树从分子水平将无性态与有性态联系起来,可以帮助判断耐热真菌疑难属、种之间以及有性态与无性态之间,在进化关系上的亲疏远近,较好的显示了耐热真菌的系统进化关系,有力地支持了形态学的观点。ITS基因可同时对于属内种间关系甚至于种内分化做出较好的解析,适合耐热真菌系统发育研究。耐热真菌分子系统学的分析结果很大程度上支持形态学分类结果,但不同的形态表型性状在分子分类标准上是不等价的。Mn-SOD氨基酸序列显示出高度保守性,其保守序列有：NHHQTYV、GGGH、QGSGW、WEHAYY。基于Mn-SOD中间片段氨基酸序列构建系统发育树,对多数耐热种类表现与形态特征一致的结果,但部分种类与形态分类特征表现不一致。亲缘关系较远,但显示同源性高的菌种间有可能发生了基因水平转移。
研究表明,针对耐热真菌形态分类还是最主要的分类方法,分子性状作为分类的一条重要依据。分子数据以形态分类为基础,是形态分类的补充,而且大多数情况下两种分类方法是可以取得统一的,因此在形态学的基础上,结合分子生物学的鉴定方法对耐热真菌资源进行分类会更科学、更合理。
Thermotolerant fungi are those that have a maximum growth temperature of near50℃(usually40℃and above) and a minimum growth temperature under20℃, showing the strong ability for adaptation and subsistence in high temperature. Thermotolerant fungi may perform bioconversion processes and produce industrially important metabolites. With more studies on the high temperature organisms, thermotolerant fungi have become a hot area for international research. A total of89strictly thermotolerant fungi are so far documented, while only23were recorded previously in China. Due to a variety of climatic conditions and habitat conditions, thermotolerant fungi diversity should be rich in China. Studies on thermotolerant fungi species and distribution in different climatic zones and habitats lay a foundation for the theory development and applied research of thermotolerant fungi.
More than800samples were collected from Shandong,Shanxi, Hebei,Shanxi, Henan, Yunnan, Jiangxi, Hainan, Xinjiang, Neimenggu, Gansu5Guangdong, Jilin, Hunan Etc. including forest, grassland, farmland, desert, alpine meadow, wetland, wasteland from2009to2011. From these samples, based on morphological characteristics combined with rDNA-ITS sequence analysis,51species in25genera were identified. Among them,2are new species,11species are new records to China, other15first reported as thermotolerant fungi. All genera and species are fully described and illustrated morphologically. Distribution sites and habitat are also cited. Dried and living cultures of all the fungi studied have been deposited in the Herbarium of Shandong Agricultural University:Plant Pathology (HSAUP).
1.Species diversity of thermotolerant fungi
New species:Taeniolella jinggangensis Y.Zhang&D.C.Li, Scutellinia subthermophila Y.Zhang&D.C.Li; New records to China:Chaetomium strumarium, Thielavia subthermophila, Hamigera avellanea, Hypoxylon fragiforme, Neosartorya fennelliae, Trichoderma ghanense, Chaetomium jodhpurense, Chaetomium luteum, Trichoderma saturnisporum, Thielavia arenaria, Chaetomium atrobrunnneum; The other38species have been known to China, include15first reported as thermotolerant fungi:Cunninghamella echinulata, Gongronella butleri, Mucor circinelloides, Chaetomium aureum, Emericella quadrilineata, Eupenicillium javanicum, Sordaria fimicola, Aspergillus ustus, Curvularia lunata, Exserohilum rostratum, Humicola fuscoatra,Scedosporium apiospermum, Penicillium funiculosum, Penicillium verruculosum, Coprinopsis cinerea. This study was aimed at investigating the species diversity and distribution of thermotolerant fungi in China to enrich the biological diversity of China and lay a useful foundation for the theory development and applied research of thermotolerant fungi.
2Taxonomic criteria at species level
Morphological features are still the main basis for classification of thermotolerant fungi. Aiming to reducing the influence of environment, all species in one genus should be cultured at the same or being close to the same conditions. heat-resistant (maximum temperature, the optimum temperature, minimum temperature) is an important basis for identification of heat-resistant species and common species. Testing temperature range is5℃interval.
For anamorphic thermotolerant fungi, the morphological characters of conidia, especially the shape, size, color, septum and ornamentation etc. are important characters in identification species. The characters of conidiophores and sporulation are also very useful in identification species in some genera. For the zygomycetes and ascomycetes:the morphological characteristics of the sexual state (teleomorph) should be used as the basis for species identification, such as ascocarp size, shape, color, appendages, ascus size, shape, ascospore size, shape, color, with or without germ pore and the number of germ pores. When ascomycetous taxa produce the anamorph regularly and the teleomorph only under specific cultural conditions, the name of the anamorph could be preferentially selected to introduce uniformity and correctness in name citations of thermotolerant fungi. Cultural characteristics (colony characteristics, color and growth characteristics) and sporulation phenotype can be used.as.the.necessary.and.useful.reference.
3Molecular phylogeny of thermotolerant fungi
Based on the morphological taxonomy, Sequencing and molecular systematics were performed on the5.8S rDNA-ITS and Mn-SOD intermediate fragment of thermotolerant fungi. The measured sequence data have been submitted to GenBank (NCBI), obtained GenBank accession numbers.
5.8S rDNA-ITS sequences can be used to differentiate the thermotolerant fungi belonging to various genera and species effectively. The sequences of5.8S rDNA-ITS are more similar among different species in the same genus than among different genera. Phylogenetic tree based on5.8S rDNA-ITS sequences from the molecular level links asexual state and the sexual state, and can help determine the thermotolerant fungi problematic genera or species. Phylogenetic tree can be used to reveal the phylogenetic relationships of thermotolerant fungi and support the viewpoint of the morphology. ITS genes can make a better resolution to the differentiation between the species and even within species, suitable for thermotolerant fungus phylogenetic studies. The results of molecular systematics supported the result of morphological classification in great extent, but the different phenotypic characteristics were not recognized equally by molecular analysis. Mn-SOD amino acid sequence is highly conserved. Conserved sequences include:NHHQTYV, GGGH, QGSGW and WEHAY. The phylogenetic tree constructed on the Mn-SOD intermediate fragment can reflect the phylogenetic relationship corresponded to morphology to the most thermotolerant fungi, but to some types the phylogeny was inconsistent with morphology. Further study should be performed since it is the initial step that using the amino acid sequence of Mn-SOD for phylogenetic analysis. It may be due to the horizontal gene transfer that the distantly related strains show high homology.
Studies have shown that morphological classification is still the most important to thermotolerant fungi. Molecular data are the complement of the morphological classification. In most cases, the two classification methods can achieve a unified. On the basis of morphology, combined with the molecular methods would be more scientific and more reasonable on the thermotolerant fungi classification.
2009-2011年在山东、山西、河北、陕西、河南、云南、江西、海南、新疆、内蒙古、甘肃、广东、吉林、湖南等地采集堆肥、培养料、草堆、腐殖质、植物残体、木屑堆、土壤、动物巢穴等标本800余份,生态类型包括森林、草原、农田、沙漠、荒地、湿地、高山草甸等,采用稀释平板法、土壤平板法等分离培养,结合形态特征观察与rDNA-ITS序列分析,分离鉴定耐热真菌25属51种,包括新种2个,中国新记录种11个,还有15个首次报道为耐热真菌。所研究的属、种均作详细形态描述和地点、生境引证。研究过的标本及活菌种保存在山东农业大学植物病理学标本室(HSAUP)。
一、系统研究了我国不同生态类型耐热真菌的种类及多样性
2个新种：井冈带孢霉Taeniolella jinggangensis Y.Zhang&D.C.Li,耐热盾盘菌Scutellinia subthermophila Y.Zhang&D.C.Li;11个中国新记录种：瘤突毛壳Chaetomium strumarium,耐热梭孢壳Thielavia subthermophila,榛色钩囊菌Hamigera avellanea,草莓状炭团菌Hypoxylon fragiforme,芬尼新萨托菌Neosartorya fennelliae,加纳木霉Trichoderma ghanense,焦特普尔毛壳Chaetomium jodhpurense,黄色毛壳Chaetomium luteum,土星孢木霉Trichoderma saturnisporum,沙栖梭孢壳Thielavia arenaria,棕色毛壳Chaetomium atrobrunnneum;38个其他耐热菌为国内已知种,其中15个首次报道为耐热真菌,包括：刺孢小克银汉霉Cunninghamella echinulata,卵形孢球托霉Gongronella butleri,卷枝毛霉Mucor circinelloides,金色毛壳Chaetomium aureum,四脊裸胞壳Emericella quadrilineata,爪哇正青霉Eupenicillium javanicum,粪生粪壳菌Sordaria fimicola,焦曲霉Aspergillus ustus,新月弯孢Curvularia lunata,嘴突凸脐孢Exserohilum rostratum,棕黑腐殖霉Humicola fuscoatra,尖端赛多孢子菌Scedosporium apiospermum,绳状青霉Penicillium funiculosum,疣孢青霉Penicillium verruculosum,灰拟鬼伞菌Coprinopsis cinerea。调查研究的结果增进了对我国耐热真菌种类的了解,丰富了我国生物多样性的内容,并为我国耐热真菌的理论研究和应用研究以及分类专著性研究奠定了基础。
二、研究总结耐热真菌种级分类标准
目前形态学特征仍然是耐热真菌分类的主要依据。对于同属内的耐热真菌种类,尽量在相同的条件下培养和形态比较,尽量减少外部环境变化或不稳定对形态的影响。耐热性(最高温度、最适温度、最低温度)是同属内耐热与非耐热真菌种级鉴别的重要依据。测试耐热真菌温度生长范围间隔为5℃。
对于半知菌等无性态耐热真菌,分生孢子的特征(包括形状、大小、颜色、隔膜数和表面纹饰等)是种级分类的主要依据；其次,分生孢子梗的形态特征、产孢细胞的延伸方式及产孢方式对属级水平上的鉴别有重要意义。对于接合菌、子囊菌等有性态耐热真菌：主要以有性结构的形态特点作为种级分类的依据,如子囊果大小,形状、颜色,有无附属丝,附属丝形状,子囊大小,形状,子囊孢子大小,形状,颜色、有无萌发孔及萌发孔个数等。当子囊菌培养条件下多产生无性态,而有性态只有在特殊条件下才能产生且出现较晚,无性态命名优先采用,保证在应用研究中耐热真菌名称的唯一性和正确性。培养特征(菌落特征、颜色以及生长特性)及产孢表型可作为分类鉴定的必要、有益参考。
三、耐热真菌分子系统学研究
本研究在形态分类的基础上,对所分离鉴定的耐热真菌,进行5.8S rDNA-ITS序列和部分Mn-SOD中间片段序列测定。测得的序列提交美国生物信息中心(NCBI)公共数据库GenBank,并获得相应的注册号。
5.8S rDNA-ITS序列能有效区分大多数不同属、种的耐热真菌,序列属内种间相似性明显大于属间相似性；基于5.8S rDNA-ITS序列构建的系统发育树从分子水平将无性态与有性态联系起来,可以帮助判断耐热真菌疑难属、种之间以及有性态与无性态之间,在进化关系上的亲疏远近,较好的显示了耐热真菌的系统进化关系,有力地支持了形态学的观点。ITS基因可同时对于属内种间关系甚至于种内分化做出较好的解析,适合耐热真菌系统发育研究。耐热真菌分子系统学的分析结果很大程度上支持形态学分类结果,但不同的形态表型性状在分子分类标准上是不等价的。Mn-SOD氨基酸序列显示出高度保守性,其保守序列有：NHHQTYV、GGGH、QGSGW、WEHAYY。基于Mn-SOD中间片段氨基酸序列构建系统发育树,对多数耐热种类表现与形态特征一致的结果,但部分种类与形态分类特征表现不一致。亲缘关系较远,但显示同源性高的菌种间有可能发生了基因水平转移。
研究表明,针对耐热真菌形态分类还是最主要的分类方法,分子性状作为分类的一条重要依据。分子数据以形态分类为基础,是形态分类的补充,而且大多数情况下两种分类方法是可以取得统一的,因此在形态学的基础上,结合分子生物学的鉴定方法对耐热真菌资源进行分类会更科学、更合理。
Thermotolerant fungi are those that have a maximum growth temperature of near50℃(usually40℃and above) and a minimum growth temperature under20℃, showing the strong ability for adaptation and subsistence in high temperature. Thermotolerant fungi may perform bioconversion processes and produce industrially important metabolites. With more studies on the high temperature organisms, thermotolerant fungi have become a hot area for international research. A total of89strictly thermotolerant fungi are so far documented, while only23were recorded previously in China. Due to a variety of climatic conditions and habitat conditions, thermotolerant fungi diversity should be rich in China. Studies on thermotolerant fungi species and distribution in different climatic zones and habitats lay a foundation for the theory development and applied research of thermotolerant fungi.
More than800samples were collected from Shandong,Shanxi, Hebei,Shanxi, Henan, Yunnan, Jiangxi, Hainan, Xinjiang, Neimenggu, Gansu5Guangdong, Jilin, Hunan Etc. including forest, grassland, farmland, desert, alpine meadow, wetland, wasteland from2009to2011. From these samples, based on morphological characteristics combined with rDNA-ITS sequence analysis,51species in25genera were identified. Among them,2are new species,11species are new records to China, other15first reported as thermotolerant fungi. All genera and species are fully described and illustrated morphologically. Distribution sites and habitat are also cited. Dried and living cultures of all the fungi studied have been deposited in the Herbarium of Shandong Agricultural University:Plant Pathology (HSAUP).
1.Species diversity of thermotolerant fungi
New species:Taeniolella jinggangensis Y.Zhang&D.C.Li, Scutellinia subthermophila Y.Zhang&D.C.Li; New records to China:Chaetomium strumarium, Thielavia subthermophila, Hamigera avellanea, Hypoxylon fragiforme, Neosartorya fennelliae, Trichoderma ghanense, Chaetomium jodhpurense, Chaetomium luteum, Trichoderma saturnisporum, Thielavia arenaria, Chaetomium atrobrunnneum; The other38species have been known to China, include15first reported as thermotolerant fungi:Cunninghamella echinulata, Gongronella butleri, Mucor circinelloides, Chaetomium aureum, Emericella quadrilineata, Eupenicillium javanicum, Sordaria fimicola, Aspergillus ustus, Curvularia lunata, Exserohilum rostratum, Humicola fuscoatra,Scedosporium apiospermum, Penicillium funiculosum, Penicillium verruculosum, Coprinopsis cinerea. This study was aimed at investigating the species diversity and distribution of thermotolerant fungi in China to enrich the biological diversity of China and lay a useful foundation for the theory development and applied research of thermotolerant fungi.
2Taxonomic criteria at species level
Morphological features are still the main basis for classification of thermotolerant fungi. Aiming to reducing the influence of environment, all species in one genus should be cultured at the same or being close to the same conditions. heat-resistant (maximum temperature, the optimum temperature, minimum temperature) is an important basis for identification of heat-resistant species and common species. Testing temperature range is5℃interval.
For anamorphic thermotolerant fungi, the morphological characters of conidia, especially the shape, size, color, septum and ornamentation etc. are important characters in identification species. The characters of conidiophores and sporulation are also very useful in identification species in some genera. For the zygomycetes and ascomycetes:the morphological characteristics of the sexual state (teleomorph) should be used as the basis for species identification, such as ascocarp size, shape, color, appendages, ascus size, shape, ascospore size, shape, color, with or without germ pore and the number of germ pores. When ascomycetous taxa produce the anamorph regularly and the teleomorph only under specific cultural conditions, the name of the anamorph could be preferentially selected to introduce uniformity and correctness in name citations of thermotolerant fungi. Cultural characteristics (colony characteristics, color and growth characteristics) and sporulation phenotype can be used.as.the.necessary.and.useful.reference.
3Molecular phylogeny of thermotolerant fungi
Based on the morphological taxonomy, Sequencing and molecular systematics were performed on the5.8S rDNA-ITS and Mn-SOD intermediate fragment of thermotolerant fungi. The measured sequence data have been submitted to GenBank (NCBI), obtained GenBank accession numbers.
5.8S rDNA-ITS sequences can be used to differentiate the thermotolerant fungi belonging to various genera and species effectively. The sequences of5.8S rDNA-ITS are more similar among different species in the same genus than among different genera. Phylogenetic tree based on5.8S rDNA-ITS sequences from the molecular level links asexual state and the sexual state, and can help determine the thermotolerant fungi problematic genera or species. Phylogenetic tree can be used to reveal the phylogenetic relationships of thermotolerant fungi and support the viewpoint of the morphology. ITS genes can make a better resolution to the differentiation between the species and even within species, suitable for thermotolerant fungus phylogenetic studies. The results of molecular systematics supported the result of morphological classification in great extent, but the different phenotypic characteristics were not recognized equally by molecular analysis. Mn-SOD amino acid sequence is highly conserved. Conserved sequences include:NHHQTYV, GGGH, QGSGW and WEHAY. The phylogenetic tree constructed on the Mn-SOD intermediate fragment can reflect the phylogenetic relationship corresponded to morphology to the most thermotolerant fungi, but to some types the phylogeny was inconsistent with morphology. Further study should be performed since it is the initial step that using the amino acid sequence of Mn-SOD for phylogenetic analysis. It may be due to the horizontal gene transfer that the distantly related strains show high homology.
Studies have shown that morphological classification is still the most important to thermotolerant fungi. Molecular data are the complement of the morphological classification. In most cases, the two classification methods can achieve a unified. On the basis of morphology, combined with the molecular methods would be more scientific and more reasonable on the thermotolerant fungi classification.
引文
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