南京龙山钾矿物表生细菌的生物多样性及其与含钾矿物相互作用的研究
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摘要
微生物广泛分布于各种地质环境中,几乎参与了从地球表面到地下深部岩石圈中所有的地质地球化学过程,直接或间接地影响了地球化学环境。本论文通过研究不同风化程度钾矿物表生细菌的生物学特性及其对硅酸盐矿物的分解效应,为阐明从矿物到土壤过程中微生物群落变化及其与矿物风化之间的关系,以及对于深入理解矿物-微生物相互作用机制等提供理论依据,丰富微生物资源库、基因库并为微生物肥料的研制提供充足菌源。
通过构建16SrDNA克隆文库对南京龙山不同风化程度钾矿物表生细菌的群落多样性进行研究,结果表明该矿区细菌群落主要可归属于14大细菌类群,其中Proteobacteria(变形菌门)及Acidobacteria(酸杆菌门)在整个克隆文库中占主要优势。Gemmatimonadetes(芽单胞菌门)是仅在土壤中发现的细菌类群。低风化度矿物样品表生细菌种群以Acidobacteria, Actinobacteria(放线菌门)及α-Proteobacteria为优势种群；高风化度矿物样品中细菌种群以α-Proteobacteria, Acidobacteria, Bacteroidetes(拟杆菌门),Actinobacteria为优势种群；而土壤样品中细菌种群以Acidobacteria及β-Proteobacteria为优势种群。菌群β-,δ-Proteobacteria, Firmicutes(厚壁菌门),Planctomycetes (浮霉菌门),Chloroflexi(绿弯菌门),Nitrospira(硝化螺旋菌门),Ktedonobacteria(纤线杆菌纲)及Verrucomicrobia(疣微菌门)随着矿物风化程度的增高呈递增的趋势。
通过对供试矿物样品中有效性元素含量等基本理化性质与表征细菌种群多样性的16S rDNA克隆文库的香农-威纳指数,辛普森指数,均匀度及丰富度指数进行PCA及CCA分析,发现矿物风化程度与矿物表生细菌存在着相互影响,相互制约的关系,矿物风化程度的高低与其中细菌多样性之间有着明显的相关性。菌群α-Proteobacteria, Bacteroidetes, Gemmatimonadetes与样品中的K,Ca,Mg含量相关,而β-Proteobacteria, Planctomycetes, Ktedonobacteria, Firmicutes及Acidobacteria与Si,Al,Fe含量及样品pH相关。菌群Proteobacteria, Bacteroidetes及Acidobacteria在钾矿物风化及矿质元素转化过程中起了重要的作用。
通过平板分离法,从不同风化程度钾矿物表面及土壤中共分离筛选出矿物表生细菌63株。限制性酶切多态性分析(ARDRA)和16S rDNA序列系统发育分析表明供试细菌具有较丰富的物种多样性。分离自土壤样品的供试菌株归属于菌群α-,β-,γ-Proteobacteria, Firmicutes, Actinobacteria, Bacteroidetes。高风化度钾矿物样品中包含4个菌群,分别为Firmicutes, Actinobacteria, a-Proteobacteria, Bacteroidetes;低风化度钾矿物样品中有2个菌群,分别为Firmicutes和Actinobacteria。Bacillus成为该矿区可培养细菌的优势菌群,52.4%的供试菌株包括在其中,而Arthrobacter和Paenibacillus,分别包含了19%和11.1%的供试菌株。
通过摇瓶试验筛选出具有明显矿物分解能力的细菌菌株21株。经16S rDNA序列系统发育分析,结果表明矿物分解细菌主要归属于Bacillus(芽孢杆菌属),Paenibacillus(类芽孢杆菌属),Arthrobacter(节杆菌属),Flexibacter(屈挠杆菌属),Rhizobium(根瘤菌属),Ralstonia(雷尔氏菌属)和Pantoea(泛菌属)菌属。本研究首次发现Rhizobium, Ralstonia, Pantoea和Flexibacter菌属的菌株对含钾硅酸盐矿物也有较强的分解能力。高效矿物分解细菌菌株M65鉴定为Rhizobium sp.。
在短期摇瓶试验及长期静置试验中,菌株Q12及M65对黑云母的分解效果最好,对钾长石的分解效果略低。两株矿物分解细菌不但能够促进原生硅酸盐矿物的风化而且还具有一定的成矿效应。
在矿物存在的条件下,菌株M65和Q12的菌体形态发生了一定的改变。供试菌株对硅酸盐矿物中铁,钙和铝等金属元素的溶出效果显著,且随着时间的变化对矿物有反复溶解和沉淀的作用。菌株M65具备两种群体感应系统,即luxR/I型信息系统与luxS/AI-2型信息系统,且在矿物存在的条件下以luxR/I型信息系统为主要的信号传递系统,其生物膜的产生与其AHLs的产量有关。菌株Q12只具备luxS/AI-2型的信息系统。细菌信号分子的产量与细胞群体数量及生长状态有关,在细菌生长的对数期内达到最高值,进入稳定期后急剧下降。
Microbes inhabit diverse environments at and near the Earth's surface. Their potential to cause geochemical change is immense. These microorganisms directly or indirectly affect the Earth's geochemical environments. Research on mineral surface's bacterial biological characteristics and their effects on the silicate mineral weathering will further understand microbial resources and diversity, clarify microbial community changes from minerals to soils and the correlation with the mineral weathering in the natural conditions. This paper also isolated mineral-solubilizing bacteria and investigated mineral-microbes interaction which would provide the theoretical and experimental bases for further understanding of mineral-microbes interaction mechamisms.
Bacterial community and diversity of different weathered feldspar and soil samples were studied by16S rDNA clone libraries. The results showed that bacterial communities from the three series samples were affiliated with fourteen major groups. Proteobacteria and Acidobacteria were the most abundant groups in the clone libraries. Gemmatimonadetes were only found in the soil samples. Bacterial community composition may be related to the degree of mineral weathering. Clones from less-weathered feldspar mineral belonged to three major groups:Acidobacteria, Actinobacteria and a-Proteobacteria; Clones from more-weathered feldspar minerals belonged to four major groups:a-Proteobacteria, Acidobacteria, Bacteroidetes and Actinobacteria; Clones from the soils belonged to two major groups:Acidobacteria and (3-Proteobacteria. An increasing abundance of β,δ-Proteobacteria, Firmicutes, Planctomycetes, Chloroflexi, Nitrospira, Ktedonobacteria and Verrucomicrobia were found along the weathering gradient from the weathered feldspar minerals to the soils.
PCA and CCA analyses of the available element contents and the bacterial diversity index showed that the degree of mineral weathering was clearly correlation with bacterial community diversity. a-Proteobacteria, Bacteroidetes, Gemmatimonadetes were correlated with K, Ca, and Mg contents; while P-Proteobacteria, Planctomycetes, Ktedonobacteria, Firmicutes and Acidobacteria were correlated with Si, Al, Fe, and pH. Proteobacteria, Bacteroidetes and Acidobacteria played an important role in the process of mineral weathering and the elements mobilization.
Sixty-three strains were isolated from different weathered feldspar and soil samples by conventional plate culture technique. ARDRA and16S rDNA sequences phylogenetic analyses indicated that the bacteria showed abundant diversity. Bacteria isolated from the soils belonged to six groups:a-, β-and y-Proteobacteria, Firmicutes, Actinobacteria, and Bacteroidetes; bacteria isolated from more-weathered feldspar minerals belonged to four groups:Firmicutes, Actinobacteria, a-Proteobacteria, and Bacteroidetes; bacteria isolated from less-weathered feldspar belonged to two groups:Firmicutes and Actinobacteria. Bacteria belonging to Bacillus, Arthrobacter and Paenibacillus accounted for52.4%,19%, and11.1%, respectively.
Of the sixty-three isolates, twenty-one strains have the ability of dissolving feldspar mineral compared to the control.16S rDNA sequences phylogenetic analysis indicated that mineral-solubilizing bacteria belonged to Bacillus. Paenibacillus, Arthrobacter, Flexibacter, Rhizobium, Ralstonia and Pantoea. Rhizobium, Ralstonia, Pantoea and Flexibacter were the first reported bacteria which could promote the mineral solubilization. Strain M65identified as Rhizobium sp. was very effective in enhancing feldspar dissolution.
A short period flask-based and long period unflask-based experiments showed that the biotite was easier to be weathered by strains Q12and M65than feldspar. The presence of the bacteria not only promoted mineral solubilizition, but also induced mineral mineralization.
The morphogeneses of strains Q12and M65were changed in the presence of feldspar and biotite minerals. The release of Fe, Ca and Al was obviously increased in the presence of stains Q12and M65. Mineral solubilization and precipitation were existed alternatively. Strain M65had two quorum signalling:luxI/luxR quorum senxing system and luxS/AI-2quorum sensing system. luxI/luxR quorum sensing system as primary autoinducers-producing allowed strain M65to control the gene expression of bioflim formation in response to changes in cell numbers. Strain Q12only had luxS/AI-2quorum sensing system. The production of autoinducers was related with cell population desity and growth state. It reached the highest production of autoinducers during the bacterial logarithm phase, and decreased significantly in the stable phase of the strains Q12and M65.
通过构建16SrDNA克隆文库对南京龙山不同风化程度钾矿物表生细菌的群落多样性进行研究,结果表明该矿区细菌群落主要可归属于14大细菌类群,其中Proteobacteria(变形菌门)及Acidobacteria(酸杆菌门)在整个克隆文库中占主要优势。Gemmatimonadetes(芽单胞菌门)是仅在土壤中发现的细菌类群。低风化度矿物样品表生细菌种群以Acidobacteria, Actinobacteria(放线菌门)及α-Proteobacteria为优势种群；高风化度矿物样品中细菌种群以α-Proteobacteria, Acidobacteria, Bacteroidetes(拟杆菌门),Actinobacteria为优势种群；而土壤样品中细菌种群以Acidobacteria及β-Proteobacteria为优势种群。菌群β-,δ-Proteobacteria, Firmicutes(厚壁菌门),Planctomycetes (浮霉菌门),Chloroflexi(绿弯菌门),Nitrospira(硝化螺旋菌门),Ktedonobacteria(纤线杆菌纲)及Verrucomicrobia(疣微菌门)随着矿物风化程度的增高呈递增的趋势。
通过对供试矿物样品中有效性元素含量等基本理化性质与表征细菌种群多样性的16S rDNA克隆文库的香农-威纳指数,辛普森指数,均匀度及丰富度指数进行PCA及CCA分析,发现矿物风化程度与矿物表生细菌存在着相互影响,相互制约的关系,矿物风化程度的高低与其中细菌多样性之间有着明显的相关性。菌群α-Proteobacteria, Bacteroidetes, Gemmatimonadetes与样品中的K,Ca,Mg含量相关,而β-Proteobacteria, Planctomycetes, Ktedonobacteria, Firmicutes及Acidobacteria与Si,Al,Fe含量及样品pH相关。菌群Proteobacteria, Bacteroidetes及Acidobacteria在钾矿物风化及矿质元素转化过程中起了重要的作用。
通过平板分离法,从不同风化程度钾矿物表面及土壤中共分离筛选出矿物表生细菌63株。限制性酶切多态性分析(ARDRA)和16S rDNA序列系统发育分析表明供试细菌具有较丰富的物种多样性。分离自土壤样品的供试菌株归属于菌群α-,β-,γ-Proteobacteria, Firmicutes, Actinobacteria, Bacteroidetes。高风化度钾矿物样品中包含4个菌群,分别为Firmicutes, Actinobacteria, a-Proteobacteria, Bacteroidetes;低风化度钾矿物样品中有2个菌群,分别为Firmicutes和Actinobacteria。Bacillus成为该矿区可培养细菌的优势菌群,52.4%的供试菌株包括在其中,而Arthrobacter和Paenibacillus,分别包含了19%和11.1%的供试菌株。
通过摇瓶试验筛选出具有明显矿物分解能力的细菌菌株21株。经16S rDNA序列系统发育分析,结果表明矿物分解细菌主要归属于Bacillus(芽孢杆菌属),Paenibacillus(类芽孢杆菌属),Arthrobacter(节杆菌属),Flexibacter(屈挠杆菌属),Rhizobium(根瘤菌属),Ralstonia(雷尔氏菌属)和Pantoea(泛菌属)菌属。本研究首次发现Rhizobium, Ralstonia, Pantoea和Flexibacter菌属的菌株对含钾硅酸盐矿物也有较强的分解能力。高效矿物分解细菌菌株M65鉴定为Rhizobium sp.。
在短期摇瓶试验及长期静置试验中,菌株Q12及M65对黑云母的分解效果最好,对钾长石的分解效果略低。两株矿物分解细菌不但能够促进原生硅酸盐矿物的风化而且还具有一定的成矿效应。
在矿物存在的条件下,菌株M65和Q12的菌体形态发生了一定的改变。供试菌株对硅酸盐矿物中铁,钙和铝等金属元素的溶出效果显著,且随着时间的变化对矿物有反复溶解和沉淀的作用。菌株M65具备两种群体感应系统,即luxR/I型信息系统与luxS/AI-2型信息系统,且在矿物存在的条件下以luxR/I型信息系统为主要的信号传递系统,其生物膜的产生与其AHLs的产量有关。菌株Q12只具备luxS/AI-2型的信息系统。细菌信号分子的产量与细胞群体数量及生长状态有关,在细菌生长的对数期内达到最高值,进入稳定期后急剧下降。
Microbes inhabit diverse environments at and near the Earth's surface. Their potential to cause geochemical change is immense. These microorganisms directly or indirectly affect the Earth's geochemical environments. Research on mineral surface's bacterial biological characteristics and their effects on the silicate mineral weathering will further understand microbial resources and diversity, clarify microbial community changes from minerals to soils and the correlation with the mineral weathering in the natural conditions. This paper also isolated mineral-solubilizing bacteria and investigated mineral-microbes interaction which would provide the theoretical and experimental bases for further understanding of mineral-microbes interaction mechamisms.
Bacterial community and diversity of different weathered feldspar and soil samples were studied by16S rDNA clone libraries. The results showed that bacterial communities from the three series samples were affiliated with fourteen major groups. Proteobacteria and Acidobacteria were the most abundant groups in the clone libraries. Gemmatimonadetes were only found in the soil samples. Bacterial community composition may be related to the degree of mineral weathering. Clones from less-weathered feldspar mineral belonged to three major groups:Acidobacteria, Actinobacteria and a-Proteobacteria; Clones from more-weathered feldspar minerals belonged to four major groups:a-Proteobacteria, Acidobacteria, Bacteroidetes and Actinobacteria; Clones from the soils belonged to two major groups:Acidobacteria and (3-Proteobacteria. An increasing abundance of β,δ-Proteobacteria, Firmicutes, Planctomycetes, Chloroflexi, Nitrospira, Ktedonobacteria and Verrucomicrobia were found along the weathering gradient from the weathered feldspar minerals to the soils.
PCA and CCA analyses of the available element contents and the bacterial diversity index showed that the degree of mineral weathering was clearly correlation with bacterial community diversity. a-Proteobacteria, Bacteroidetes, Gemmatimonadetes were correlated with K, Ca, and Mg contents; while P-Proteobacteria, Planctomycetes, Ktedonobacteria, Firmicutes and Acidobacteria were correlated with Si, Al, Fe, and pH. Proteobacteria, Bacteroidetes and Acidobacteria played an important role in the process of mineral weathering and the elements mobilization.
Sixty-three strains were isolated from different weathered feldspar and soil samples by conventional plate culture technique. ARDRA and16S rDNA sequences phylogenetic analyses indicated that the bacteria showed abundant diversity. Bacteria isolated from the soils belonged to six groups:a-, β-and y-Proteobacteria, Firmicutes, Actinobacteria, and Bacteroidetes; bacteria isolated from more-weathered feldspar minerals belonged to four groups:Firmicutes, Actinobacteria, a-Proteobacteria, and Bacteroidetes; bacteria isolated from less-weathered feldspar belonged to two groups:Firmicutes and Actinobacteria. Bacteria belonging to Bacillus, Arthrobacter and Paenibacillus accounted for52.4%,19%, and11.1%, respectively.
Of the sixty-three isolates, twenty-one strains have the ability of dissolving feldspar mineral compared to the control.16S rDNA sequences phylogenetic analysis indicated that mineral-solubilizing bacteria belonged to Bacillus. Paenibacillus, Arthrobacter, Flexibacter, Rhizobium, Ralstonia and Pantoea. Rhizobium, Ralstonia, Pantoea and Flexibacter were the first reported bacteria which could promote the mineral solubilization. Strain M65identified as Rhizobium sp. was very effective in enhancing feldspar dissolution.
A short period flask-based and long period unflask-based experiments showed that the biotite was easier to be weathered by strains Q12and M65than feldspar. The presence of the bacteria not only promoted mineral solubilizition, but also induced mineral mineralization.
The morphogeneses of strains Q12and M65were changed in the presence of feldspar and biotite minerals. The release of Fe, Ca and Al was obviously increased in the presence of stains Q12and M65. Mineral solubilization and precipitation were existed alternatively. Strain M65had two quorum signalling:luxI/luxR quorum senxing system and luxS/AI-2quorum sensing system. luxI/luxR quorum sensing system as primary autoinducers-producing allowed strain M65to control the gene expression of bioflim formation in response to changes in cell numbers. Strain Q12only had luxS/AI-2quorum sensing system. The production of autoinducers was related with cell population desity and growth state. It reached the highest production of autoinducers during the bacterial logarithm phase, and decreased significantly in the stable phase of the strains Q12and M65.
引文
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