四川几种竹叶部微生物区系研究
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摘要
四川地区栽培的竹林主要分布于西南部,种类多,用途广。2008年特大冰雪灾害的发生对竹产业的生产和发展造成重大的损失,各地区竹林均表现出不同程度的受害,严重影响竹林林分健康。竹叶作为竹林健康状况的重要指示器官,同时也是许多微生物生存繁衍的重要场所,更是病原微生物侵害竹子的重要入口。其表面与内部均生活着大量不同种类的微生物,并构成稳定复杂的微生态系统。
本实验首次以四川雅安、宜宾、泸州三地6种栽培范围较广的竹种为研究材料,主要包括雅安地区(撑x绿杂交竹(Bambusa pervariabilis×Dendrocalamopsis grandis)、慈竹(Neosinocalamus affinis))、宜宾地区(毛竹(Phyllostachys heterocycla)、硬头黄(Bambusa rigida))、泸州地区(梁山慈(Dendrocalamus farinosus)、西凤竹(Bambusa multiplex))。采用传统林木病理学和现代分子生物学相结合的研究方法,主要对雪灾后6竹种竹林(实施清林等恢复措施)叶部微生物区系中可培养微生物进行了数量和种群的季节动态分析。结果表明：
①竹叶部微生物区系主要由真菌和细菌两大类群组成,且细菌数量明显高于真菌。
三地区6竹种叶部微生物总数量(慈竹附生微生物除外)只与细菌数量表现出显著相关(P≤0.05)或极显著相关(P≤0.01),另外,慈竹附生真菌、附生放线菌及附生微生物总数间也表现出显著相关(P≤0.05),硬头黄内生真菌与内生细菌、内生微生物总数也表现出显著相关(P≤0.05)；同一竹种微生物总数量的季节变化趋势与细菌表现出相似的规律,与真菌、放线菌存在差异,而竹种间微生物数量的季节变化趋势存在较大差异；对同一地区两竹种微生物数量的比较分析表明,除泸州梁山慈叶部微生物及各类群数量均显著高于西凤竹外,雅安、宜宾竹林叶部微生物数量在季节、竹种和类群间差异较大,无明显规律；竹种和季节是造成叶部微生物数量差异的两个主要因素,双因素方差分析结果表明,两者对不同微生物类群数量的影响程度不同。
②三地区6竹种叶部微生物的种群组成及优势菌存在较大差异,各微生物种群的季节演替均有不同的变化规律。
雅安地区共分离得到真菌43属60种,细菌14属24种,放线菌1属1种,明显多于宜宾(真菌26属33种,细菌12属17种)、泸州(真菌23属32种,细菌11属17种),且雅安地区两竹种叶部微生物的共有物种数要多于其余两地。在雅安两竹种的叶部微生物区系中,附生微生物和内生微生物类群均有其共有物种；在宜宾,两竹种在附生真菌、附生细菌、内生真菌的组成上有共有物种,而内生细菌种群组成差异较大；在泸州,两竹种附生真菌、内生真菌的种群组成上有共有物种,而附生细菌和内生细菌种群组成差异较大。
③三地区6竹种叶部微生物物种丰富度指数的变化无一致规律。
雅安慈竹叶部微生物各类群间均表现出不同的季节变化趋势,撑×绿杂交竹附生微生物各类群间变化趋势相反,内生微生物变化趋势却完全一致。而不同竹种间,除慈竹附生真菌物种丰富度指数在四季均大于撑×绿杂交竹外,其余微生物类群无明显规律。
宜宾毛竹叶部微生物各类群间表现出相似的季节变化规律,而硬头黄各类群指数变化趋势差异较大。在同一微生物类群的不同竹种间,硬头黄竹附生真菌物种丰富度在四季均大于毛竹,而毛竹内生细菌物种丰富度在四季(除冬季)大于硬头黄竹,两竹种附生细菌和内生真菌无明显规律。
泸州两竹种叶部微生物各类群间物种丰富度指数的变化趋势均有较大差异,同一微生物类群在不同竹种间的差异也较大,无明显规律。
④在调查采样的过程中,竹林内仍有部分常见病虫害的发生,不同季节不同竹林内病虫害种类与发生程度不同,主要病害为常见的煤污,这与煤污菌在叶片上的广泛存在(如Cladosporium cladosporioides、Alternaria alternate等)密不可分,而主要虫害为食叶害虫(如竹织叶野螟(Algedonia coclesalis)等)和蚜虫。在微生物区系的组成及优势种群中存在潜在病原菌(并未引起植物表现病状),同时也存在一些有益微生物。
Cultivated bamboo forest, distributed mainly in southwest Sichuan Province, consists of a wide range of species with various usages. In early 2008, an unusual rainfall and a snowstorm destroyed a large population of the bamboo forest, and caused a significant loss in bamboo production in different areas. These adverse weather conditions affected health of the bamboo plants, including bamboo leaves. Not only reflect the growing condition of the entire bamboo plant, bamboo leaves are also important organs which provide an ideal habitat for many microorganisms and an entry for pathogen to colonize the entire system. These microorganisms, both on the surface and inside of the leaves, are abundant, belong to various species, and can form complex yet steady micro-ecosystem.
Six species of bamboo plants cultivated in Ya'an (B. pervariabilis×D. grandis, N. affinis), Yibin (P. heterocycla, B. rigida) and Luzhou (D. farinosus, B. multiplex) were studied in this research. After the recovery from the snowstorm, traditional forest pathology and molecular biology methods were used to investigate seasonal dynamics of cultivable microorganisms'quantity and compositions of foliar sample from the bamboo plants. The following results have been obtained:
①The foliar microflora of bamboo plants mainly consisted of fungi and bacteria, with bacteria as the dominating microorganism.
The total microorganism counts were significantly correlated (P<0.05) or highly significantly correlated (.P<0.01) with bacteria counts. The counts of phylloplane fungi, actinomycetes and microorganism of N. affinis leaves were significantly correlated (P<0.05) with each other; the endophyte counts of B. rigida leaves had significant correlation (P<0.05) with bacteria's and total microorganism's counts as well. Seasonal dynamics of microorganism's total counts were similar to the bacteria counts, but different from fungi and actinomycetes of the same bamboo species. Seasonal dynamics of total microorganism counts were different among different species in different locations, and species in the same locations except the two species in Luzhou area (D. farinosus, and B. multiplex). In this study, bamboo species and seasonal changes were the two major factors that influenced the total microorganism counts. Results of double-factor ANOVA also showed that the two factors had different influences on different microorganism populations.
②The microorganisms isolated from the six species at the three locations demonstrated various differences in population composition, dominant population, and their responses to seasonal changes.
In Ya'an area, samples isolated contained 43 genera and 60 species of fungi,14 genera and 24 species of bacteria, and 1 genus and 1 species of actinomycetes. The microorganism population was significantly larger than the microorganisms isolated from samples collected from the other two locations. Samples from Yibin contained 26 genera and 33 species of fungi,12 genera and 17 species of bacteria, and sample collected from Luzhou contained 23 genera and 32 species of fungi, and 11 genera and 17 species of bacteria. Similar trend was found with the commonly seen microorganism counts from different locations as well. In Ya'an, the phylloplane and endophytic microflora of two bamboo forests had common species. Similarly, it was also in samples collected in Yibin except the endophytic bacteria. However, samples collected in Luzhou only had common species in phylloplane and endophytic fungi, the bacteria populations were remarkably difference on compositions between phylloplane microflora and endophytic microflora.
③The variations of species richness indexes between different bamboo forests in the same area were inconsistent.
In samples collected from Ya'an, responses to seasonal changes of different microorganism populations varied. For example, phylloplane population of N. affinis leaves showed the opposite tendency, but similar tendency among endophytic populations in B. pervariabilis×D. grandis leaves. Species richness indexes of same microorganism population between different bamboo species did not show a clear trend, except phylloplane fungi of N. affinis leaves were more abundant than that of B. pervariabilis×D. grandis leaves in four seasons.
In Yibin, similar seasonal responses were found among different microorganism populations of P. heterocycla leaves, but not that of B. rigida leaves. Between the two different bamboo species, the species richness indexes of phylloplane fungi of B. rigida leaves were larger than that of P. heterocycla leaves in four seasons, but the endophytic bacteria index were lower in B. rigida than that of P. heterocycla leaves in four seasons except for winter.
In Luzhou, results showed different variation tendency between different bamboo species and microorganism populations.
④During the survey, commonly seen pests were also discovered at different seasons and among different bamboo species. The main fungal disease was sooty blotch, which was directly related to the widespread phylloplane sooty moulds (C. cladosporioides, Alternaria alternate, et al.). The main insect was defoliator (Algedonia coclesalis, et al.) and aphids (Aphidoidea). There were also some potential pathogens in the foliar microflora and dominant species, and some beneficial microorganisms.
本实验首次以四川雅安、宜宾、泸州三地6种栽培范围较广的竹种为研究材料,主要包括雅安地区(撑x绿杂交竹(Bambusa pervariabilis×Dendrocalamopsis grandis)、慈竹(Neosinocalamus affinis))、宜宾地区(毛竹(Phyllostachys heterocycla)、硬头黄(Bambusa rigida))、泸州地区(梁山慈(Dendrocalamus farinosus)、西凤竹(Bambusa multiplex))。采用传统林木病理学和现代分子生物学相结合的研究方法,主要对雪灾后6竹种竹林(实施清林等恢复措施)叶部微生物区系中可培养微生物进行了数量和种群的季节动态分析。结果表明：
①竹叶部微生物区系主要由真菌和细菌两大类群组成,且细菌数量明显高于真菌。
三地区6竹种叶部微生物总数量(慈竹附生微生物除外)只与细菌数量表现出显著相关(P≤0.05)或极显著相关(P≤0.01),另外,慈竹附生真菌、附生放线菌及附生微生物总数间也表现出显著相关(P≤0.05),硬头黄内生真菌与内生细菌、内生微生物总数也表现出显著相关(P≤0.05)；同一竹种微生物总数量的季节变化趋势与细菌表现出相似的规律,与真菌、放线菌存在差异,而竹种间微生物数量的季节变化趋势存在较大差异；对同一地区两竹种微生物数量的比较分析表明,除泸州梁山慈叶部微生物及各类群数量均显著高于西凤竹外,雅安、宜宾竹林叶部微生物数量在季节、竹种和类群间差异较大,无明显规律；竹种和季节是造成叶部微生物数量差异的两个主要因素,双因素方差分析结果表明,两者对不同微生物类群数量的影响程度不同。
②三地区6竹种叶部微生物的种群组成及优势菌存在较大差异,各微生物种群的季节演替均有不同的变化规律。
雅安地区共分离得到真菌43属60种,细菌14属24种,放线菌1属1种,明显多于宜宾(真菌26属33种,细菌12属17种)、泸州(真菌23属32种,细菌11属17种),且雅安地区两竹种叶部微生物的共有物种数要多于其余两地。在雅安两竹种的叶部微生物区系中,附生微生物和内生微生物类群均有其共有物种；在宜宾,两竹种在附生真菌、附生细菌、内生真菌的组成上有共有物种,而内生细菌种群组成差异较大；在泸州,两竹种附生真菌、内生真菌的种群组成上有共有物种,而附生细菌和内生细菌种群组成差异较大。
③三地区6竹种叶部微生物物种丰富度指数的变化无一致规律。
雅安慈竹叶部微生物各类群间均表现出不同的季节变化趋势,撑×绿杂交竹附生微生物各类群间变化趋势相反,内生微生物变化趋势却完全一致。而不同竹种间,除慈竹附生真菌物种丰富度指数在四季均大于撑×绿杂交竹外,其余微生物类群无明显规律。
宜宾毛竹叶部微生物各类群间表现出相似的季节变化规律,而硬头黄各类群指数变化趋势差异较大。在同一微生物类群的不同竹种间,硬头黄竹附生真菌物种丰富度在四季均大于毛竹,而毛竹内生细菌物种丰富度在四季(除冬季)大于硬头黄竹,两竹种附生细菌和内生真菌无明显规律。
泸州两竹种叶部微生物各类群间物种丰富度指数的变化趋势均有较大差异,同一微生物类群在不同竹种间的差异也较大,无明显规律。
④在调查采样的过程中,竹林内仍有部分常见病虫害的发生,不同季节不同竹林内病虫害种类与发生程度不同,主要病害为常见的煤污,这与煤污菌在叶片上的广泛存在(如Cladosporium cladosporioides、Alternaria alternate等)密不可分,而主要虫害为食叶害虫(如竹织叶野螟(Algedonia coclesalis)等)和蚜虫。在微生物区系的组成及优势种群中存在潜在病原菌(并未引起植物表现病状),同时也存在一些有益微生物。
Cultivated bamboo forest, distributed mainly in southwest Sichuan Province, consists of a wide range of species with various usages. In early 2008, an unusual rainfall and a snowstorm destroyed a large population of the bamboo forest, and caused a significant loss in bamboo production in different areas. These adverse weather conditions affected health of the bamboo plants, including bamboo leaves. Not only reflect the growing condition of the entire bamboo plant, bamboo leaves are also important organs which provide an ideal habitat for many microorganisms and an entry for pathogen to colonize the entire system. These microorganisms, both on the surface and inside of the leaves, are abundant, belong to various species, and can form complex yet steady micro-ecosystem.
Six species of bamboo plants cultivated in Ya'an (B. pervariabilis×D. grandis, N. affinis), Yibin (P. heterocycla, B. rigida) and Luzhou (D. farinosus, B. multiplex) were studied in this research. After the recovery from the snowstorm, traditional forest pathology and molecular biology methods were used to investigate seasonal dynamics of cultivable microorganisms'quantity and compositions of foliar sample from the bamboo plants. The following results have been obtained:
①The foliar microflora of bamboo plants mainly consisted of fungi and bacteria, with bacteria as the dominating microorganism.
The total microorganism counts were significantly correlated (P<0.05) or highly significantly correlated (.P<0.01) with bacteria counts. The counts of phylloplane fungi, actinomycetes and microorganism of N. affinis leaves were significantly correlated (P<0.05) with each other; the endophyte counts of B. rigida leaves had significant correlation (P<0.05) with bacteria's and total microorganism's counts as well. Seasonal dynamics of microorganism's total counts were similar to the bacteria counts, but different from fungi and actinomycetes of the same bamboo species. Seasonal dynamics of total microorganism counts were different among different species in different locations, and species in the same locations except the two species in Luzhou area (D. farinosus, and B. multiplex). In this study, bamboo species and seasonal changes were the two major factors that influenced the total microorganism counts. Results of double-factor ANOVA also showed that the two factors had different influences on different microorganism populations.
②The microorganisms isolated from the six species at the three locations demonstrated various differences in population composition, dominant population, and their responses to seasonal changes.
In Ya'an area, samples isolated contained 43 genera and 60 species of fungi,14 genera and 24 species of bacteria, and 1 genus and 1 species of actinomycetes. The microorganism population was significantly larger than the microorganisms isolated from samples collected from the other two locations. Samples from Yibin contained 26 genera and 33 species of fungi,12 genera and 17 species of bacteria, and sample collected from Luzhou contained 23 genera and 32 species of fungi, and 11 genera and 17 species of bacteria. Similar trend was found with the commonly seen microorganism counts from different locations as well. In Ya'an, the phylloplane and endophytic microflora of two bamboo forests had common species. Similarly, it was also in samples collected in Yibin except the endophytic bacteria. However, samples collected in Luzhou only had common species in phylloplane and endophytic fungi, the bacteria populations were remarkably difference on compositions between phylloplane microflora and endophytic microflora.
③The variations of species richness indexes between different bamboo forests in the same area were inconsistent.
In samples collected from Ya'an, responses to seasonal changes of different microorganism populations varied. For example, phylloplane population of N. affinis leaves showed the opposite tendency, but similar tendency among endophytic populations in B. pervariabilis×D. grandis leaves. Species richness indexes of same microorganism population between different bamboo species did not show a clear trend, except phylloplane fungi of N. affinis leaves were more abundant than that of B. pervariabilis×D. grandis leaves in four seasons.
In Yibin, similar seasonal responses were found among different microorganism populations of P. heterocycla leaves, but not that of B. rigida leaves. Between the two different bamboo species, the species richness indexes of phylloplane fungi of B. rigida leaves were larger than that of P. heterocycla leaves in four seasons, but the endophytic bacteria index were lower in B. rigida than that of P. heterocycla leaves in four seasons except for winter.
In Luzhou, results showed different variation tendency between different bamboo species and microorganism populations.
④During the survey, commonly seen pests were also discovered at different seasons and among different bamboo species. The main fungal disease was sooty blotch, which was directly related to the widespread phylloplane sooty moulds (C. cladosporioides, Alternaria alternate, et al.). The main insect was defoliator (Algedonia coclesalis, et al.) and aphids (Aphidoidea). There were also some potential pathogens in the foliar microflora and dominant species, and some beneficial microorganisms.
引文
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