自然生态恢复过程中铜尾矿废弃地微生物学性质及细菌群落结构研究
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摘要
铜尾矿废弃地是一种人工原生裸地,铜尾矿废弃地上生态系统的形成和发育表现为典型的原生演替。铜陵铜尾矿废弃地大部分都是在通过自然方式恢复,弃置后的尾矿裸地表面首先形成以藻类、苔藓为优势的隐花植物结皮,而后草本植物定居并逐渐形成自然植物群落。本研究以弃置时间分别为19 a和30 a的铜陵杨山冲尾矿废弃地和铜官山尾矿废弃地为对象,探讨自然生态恢复过程中,随着植物群落的演替和基质理化性质的变化,铜尾矿废弃地中微生物量、微生物活性及土壤酶活性等微生物学性质的演变,并利用聚合酶链式反应-变性梯度凝胶电泳(PCR-DGGE)技术对尾矿废弃地细菌(包括自由固氮微生物)的群落结构进行研究。
随着铜尾矿废弃地生态系统原生演替的进行,废弃地中有机质、含水量、总氮及速效磷含量增加,pH降低；尾矿废弃地表层微生物量碳、氮及脲酶和碱性磷酸酶活性随生态系统的发展而提高,而微生物代谢熵有所降低；铜官山老尾矿废弃地较杨山冲废弃地表现出高的养分含量和微生物量,表明尾矿废弃地的理化性质、微生物量和土壤酶活性受到生态系统发育时间的重要影响。
铜尾矿库废弃地不同恢复阶段细菌16S rDNA V3可变区的DGGE图谱分析表明,不同植物群落演替阶段尾矿中细菌多样性无明显的变化趋势；但同一植物群落下的尾矿废弃地,A-层较C-层表现出高的细菌多样性,这与尾矿中有机质、氮和磷等养分含量的变化相一致。细菌16S rDNA V3-V5可变区的DGGE分析及基因测序发现,铜尾矿废弃地中细菌的主要类群为变形菌门(Proteobacteria)(包括α-变形菌纲(alpha-proteobacteria),β-变形菌纲(beta-proteobacteria)和γ-变形菌纲(gamma-proteobacteria)),其次为厚壁菌门(Firmicutes) (包括芽孢杆菌纲(Bacilli)和梭菌纲(Clostridia)),此外还包括硝化螺旋菌门(Nitrospirae)、放线菌门(Actinobacteria)、异常球菌嗜热菌门(Deinococcus-Thermus)和拟杆菌门(Bacteroidetes)等。数据分析表明,铜尾矿废弃地中细菌的多样性同时受到养分的积累状况和植被类型影响。
自由固氮微生物为铜尾矿废弃地的自然生态恢复提供了重要的氮源，nifH基因(编码固氮铁蛋白)的DGGE指纹图谱分析表明,随着植物群落从裸地到维管植物群落的演替,杨山冲尾矿废弃地固氮微生物多样性未表现出明显的变化趋势；相对于弃置时间较短的杨山冲尾矿废弃地,铜官山老尾矿废弃地表现出高的固氮微生物多样性。测序结果表明,尾矿废弃地中自由固氮微生物主要类群包括变形菌门和蓝藻门。基于条带分布和铜尾矿废弃地理化性质的典范对应分析显示,铜尾矿废弃地中有机质含量、含水量、pH对自由固氮微生物多样性影响显著(p<0.05)。植物群落通过改善基质pH、提高有机质和水分含量影响尾矿废弃地中自由固氮微生物多样性和群落结构。
铜尾矿废弃地上生长的优势植物白茅(Imperata cylindrica var. major)、狗牙根(Cynodon dactylon)、中华结缕草(Zoysia sinica)、芒(Miscanthus sinensis)、五节芒(Miscanthus floridulus)和木贼(Hippochaete ramosissimum)的根际和非根际nifH基因多样性分析表明：在杨山冲铜尾矿废弃地,优势植物根际尾矿固氮微生物多样性低于非根际尾矿；相对于杨山冲尾矿废弃地,铜官山老尾矿废弃地白茅根际尾矿具有高的固氮微生物多样性；随植物定居时间的增加,杨山冲尾矿废弃地白茅和木贼根际尾矿中固氮微生物多样性提高,而中华结缕草表现出相反的趋势。系统发育分析显示,铜尾矿废弃地根际自由际固氮微生物主要类群包括α-变形菌,β-变形菌和蓝藻。大多数nifH基因序列与已知可培养的微生物及蓝藻表现出较低的相似性,说明铜尾矿废弃地的固氮微生物可能代表着新的固氮微生物类群。尾矿废弃地优势植物类型、植物群落定居时间、植物生活型及尾矿的理化性质皆影响优势植物根际自由固氮微生物多样性。
The wastes of copper mine tailings is a pedogenetic substrate. Formation and development of ecosystem on the wastelands of copper mine tailings is a typical primary succession. Most wastelands of copper mine tailings in Tongling, China, are restored in natural way. The tailings wastelands firstly come into being cryptogamic crust after the disposal, which is dominated by algae and moss, then herbaceous vascular plants settle down and gradually form a natural plant community. The changes in physico-chemical properties, microbial characteristics, enzyme activity in wastelands during primary succession were investigated in two wastelands of copper mine tailings near Tongling, and the structure of bacterial community, free-living nitrogen-fixing bacteria included, was examined using Polymerase Chain Reaction-Denatured Gradient Gel Electrophoresis (PCR-DGGE) approach. The Yangshanchong wasteland and Tongguanshan wasteland studied were discarded for 19 a and 30 a, respectively.
Soil organic matter, total nitrogen, available phosphorus and water holding increased with primary succession, while the pH showed opposite trend. The microbial biomass C and N, and activities of urease and alkaline phosphatase in surface layer increased, while metabolic quotient decreased with development of ecosystem on the wastelands of copper mine tailings.
As the development of wastelands, no significant trend of bacterial diversity was found in different successional stages of the same horizon. However, A-horizon displayed higher bacterial diversity than C-horizon, which was consistent with the change of soil organic matter and total nitrogen in content. Phylogenetic analysis of 16S rDNA V3-V5 variable region gene sequences retrieved from DGGE gels indicated that there were Proteobacteria, alpha-proteobacteria, beta-proteobacteria and garma-proteobacteria included, Firmicute, Bacilli and Clostridia included, Nitrospirae, Actinobacteria, Deinococcus-Thermus and Bacteroidetes detected in wastelands of copper mine tailings. Both the accumulation of nutrients and the type of vegetation influenced the changes of bacterial diversity in naturally reclaimed wastelands of copper mine tailings.
Biological nitrogen fixing is an important source of nitrogen input in the natural ecological restoration of mine wastelands. The analysis of diversity of nifH gene in tailings samples with different plant communities from wastelands using PCR-DGGE demonstrated that the diversity of nifH gene in Yangshanchong wasteland did not display a consistent successional tendency with development of plant communities. The nitrogen-fixing microorganism community in the upper layer of tailings in Tongguanshan wasteland showed higher Shannon-Wiener diversity index than that in Yangshanchong wasteland. Phylogenetic analysis of nifH gene sequences retrieved from the DGGE gels indicated that there were mainly two taxa of free-living nitrogen-fixing microorganisms, Proteobacteria and Cyanobacteria living in the wastelands investigated. Canonical correspondence analysis based on the relationship between band patterns of DGGE profile and physico-chemical properties of tailings samples showed that the diversity of nifH gene in different tailings samples was mainly affected by loss of ignition, water content and pH of wastelands (p< 0.05). The dominant plant species and development period of plant communities by ameliorating pH, increasing organic matter and water content affected the diversity and structure of the free-living nitrogen-fixing microorganisms in wastelands of copper mine tailings.
The study on free-living nitrogen-fixing microorganism communities in rhizosphere and non-rhizosphere of pioneer plants growing on wastelands of copper mine tailings, including Imperata cylindrica var. major, Cynodon dactylon, Zoysia sinica, Miscanthus sinensis, Miscanthus floridulus and Hippochaete ramosissimum, indicated that, free-living nitrogen-fixing microorganisms in the bulk soil in Yangshanchong wasteland showed higher diversity indices than that in the rhizosphere of dominant plant; and the nitrogen-fixing microorganism community in the rhizosphere of Imperata cylindrica var. major in Tongguanshan wasteland showed higher Shannon-Wiener diversity index than that in Yangshanchong wasteland. The diversity of free-living nitrogen-fixing microorganism communities in rhizosphere of Imperata cylindrica var. major and Hippochaete ramosissimum increased with the plant ages while Zoysia sinica showed the opposite trend. Phylogenetic analysis indicated that nifH gene sequences retrieved from DGGE gels clustered in the Proteobacteria and Cyanobacteria. Most of nifH gene fragments sequenced were not closely related to any known cultivated nitrogen-fixing bacteria and cyanobacteria, suggesting that the nifH gene sequences in wastelands of copper mine tailings investigated are unique and may represent novel sequences of nitrogen-fixing community. Our results indicated that diversity of nifH gene sequences in rhizosphere and non-rhizosphere tailings samples was affected by development period of plant community, plant species, plant life forms and physico-chemical properties of wastelands.
随着铜尾矿废弃地生态系统原生演替的进行,废弃地中有机质、含水量、总氮及速效磷含量增加,pH降低；尾矿废弃地表层微生物量碳、氮及脲酶和碱性磷酸酶活性随生态系统的发展而提高,而微生物代谢熵有所降低；铜官山老尾矿废弃地较杨山冲废弃地表现出高的养分含量和微生物量,表明尾矿废弃地的理化性质、微生物量和土壤酶活性受到生态系统发育时间的重要影响。
铜尾矿库废弃地不同恢复阶段细菌16S rDNA V3可变区的DGGE图谱分析表明,不同植物群落演替阶段尾矿中细菌多样性无明显的变化趋势；但同一植物群落下的尾矿废弃地,A-层较C-层表现出高的细菌多样性,这与尾矿中有机质、氮和磷等养分含量的变化相一致。细菌16S rDNA V3-V5可变区的DGGE分析及基因测序发现,铜尾矿废弃地中细菌的主要类群为变形菌门(Proteobacteria)(包括α-变形菌纲(alpha-proteobacteria),β-变形菌纲(beta-proteobacteria)和γ-变形菌纲(gamma-proteobacteria)),其次为厚壁菌门(Firmicutes) (包括芽孢杆菌纲(Bacilli)和梭菌纲(Clostridia)),此外还包括硝化螺旋菌门(Nitrospirae)、放线菌门(Actinobacteria)、异常球菌嗜热菌门(Deinococcus-Thermus)和拟杆菌门(Bacteroidetes)等。数据分析表明,铜尾矿废弃地中细菌的多样性同时受到养分的积累状况和植被类型影响。
自由固氮微生物为铜尾矿废弃地的自然生态恢复提供了重要的氮源，nifH基因(编码固氮铁蛋白)的DGGE指纹图谱分析表明,随着植物群落从裸地到维管植物群落的演替,杨山冲尾矿废弃地固氮微生物多样性未表现出明显的变化趋势；相对于弃置时间较短的杨山冲尾矿废弃地,铜官山老尾矿废弃地表现出高的固氮微生物多样性。测序结果表明,尾矿废弃地中自由固氮微生物主要类群包括变形菌门和蓝藻门。基于条带分布和铜尾矿废弃地理化性质的典范对应分析显示,铜尾矿废弃地中有机质含量、含水量、pH对自由固氮微生物多样性影响显著(p<0.05)。植物群落通过改善基质pH、提高有机质和水分含量影响尾矿废弃地中自由固氮微生物多样性和群落结构。
铜尾矿废弃地上生长的优势植物白茅(Imperata cylindrica var. major)、狗牙根(Cynodon dactylon)、中华结缕草(Zoysia sinica)、芒(Miscanthus sinensis)、五节芒(Miscanthus floridulus)和木贼(Hippochaete ramosissimum)的根际和非根际nifH基因多样性分析表明：在杨山冲铜尾矿废弃地,优势植物根际尾矿固氮微生物多样性低于非根际尾矿；相对于杨山冲尾矿废弃地,铜官山老尾矿废弃地白茅根际尾矿具有高的固氮微生物多样性；随植物定居时间的增加,杨山冲尾矿废弃地白茅和木贼根际尾矿中固氮微生物多样性提高,而中华结缕草表现出相反的趋势。系统发育分析显示,铜尾矿废弃地根际自由际固氮微生物主要类群包括α-变形菌,β-变形菌和蓝藻。大多数nifH基因序列与已知可培养的微生物及蓝藻表现出较低的相似性,说明铜尾矿废弃地的固氮微生物可能代表着新的固氮微生物类群。尾矿废弃地优势植物类型、植物群落定居时间、植物生活型及尾矿的理化性质皆影响优势植物根际自由固氮微生物多样性。
The wastes of copper mine tailings is a pedogenetic substrate. Formation and development of ecosystem on the wastelands of copper mine tailings is a typical primary succession. Most wastelands of copper mine tailings in Tongling, China, are restored in natural way. The tailings wastelands firstly come into being cryptogamic crust after the disposal, which is dominated by algae and moss, then herbaceous vascular plants settle down and gradually form a natural plant community. The changes in physico-chemical properties, microbial characteristics, enzyme activity in wastelands during primary succession were investigated in two wastelands of copper mine tailings near Tongling, and the structure of bacterial community, free-living nitrogen-fixing bacteria included, was examined using Polymerase Chain Reaction-Denatured Gradient Gel Electrophoresis (PCR-DGGE) approach. The Yangshanchong wasteland and Tongguanshan wasteland studied were discarded for 19 a and 30 a, respectively.
Soil organic matter, total nitrogen, available phosphorus and water holding increased with primary succession, while the pH showed opposite trend. The microbial biomass C and N, and activities of urease and alkaline phosphatase in surface layer increased, while metabolic quotient decreased with development of ecosystem on the wastelands of copper mine tailings.
As the development of wastelands, no significant trend of bacterial diversity was found in different successional stages of the same horizon. However, A-horizon displayed higher bacterial diversity than C-horizon, which was consistent with the change of soil organic matter and total nitrogen in content. Phylogenetic analysis of 16S rDNA V3-V5 variable region gene sequences retrieved from DGGE gels indicated that there were Proteobacteria, alpha-proteobacteria, beta-proteobacteria and garma-proteobacteria included, Firmicute, Bacilli and Clostridia included, Nitrospirae, Actinobacteria, Deinococcus-Thermus and Bacteroidetes detected in wastelands of copper mine tailings. Both the accumulation of nutrients and the type of vegetation influenced the changes of bacterial diversity in naturally reclaimed wastelands of copper mine tailings.
Biological nitrogen fixing is an important source of nitrogen input in the natural ecological restoration of mine wastelands. The analysis of diversity of nifH gene in tailings samples with different plant communities from wastelands using PCR-DGGE demonstrated that the diversity of nifH gene in Yangshanchong wasteland did not display a consistent successional tendency with development of plant communities. The nitrogen-fixing microorganism community in the upper layer of tailings in Tongguanshan wasteland showed higher Shannon-Wiener diversity index than that in Yangshanchong wasteland. Phylogenetic analysis of nifH gene sequences retrieved from the DGGE gels indicated that there were mainly two taxa of free-living nitrogen-fixing microorganisms, Proteobacteria and Cyanobacteria living in the wastelands investigated. Canonical correspondence analysis based on the relationship between band patterns of DGGE profile and physico-chemical properties of tailings samples showed that the diversity of nifH gene in different tailings samples was mainly affected by loss of ignition, water content and pH of wastelands (p< 0.05). The dominant plant species and development period of plant communities by ameliorating pH, increasing organic matter and water content affected the diversity and structure of the free-living nitrogen-fixing microorganisms in wastelands of copper mine tailings.
The study on free-living nitrogen-fixing microorganism communities in rhizosphere and non-rhizosphere of pioneer plants growing on wastelands of copper mine tailings, including Imperata cylindrica var. major, Cynodon dactylon, Zoysia sinica, Miscanthus sinensis, Miscanthus floridulus and Hippochaete ramosissimum, indicated that, free-living nitrogen-fixing microorganisms in the bulk soil in Yangshanchong wasteland showed higher diversity indices than that in the rhizosphere of dominant plant; and the nitrogen-fixing microorganism community in the rhizosphere of Imperata cylindrica var. major in Tongguanshan wasteland showed higher Shannon-Wiener diversity index than that in Yangshanchong wasteland. The diversity of free-living nitrogen-fixing microorganism communities in rhizosphere of Imperata cylindrica var. major and Hippochaete ramosissimum increased with the plant ages while Zoysia sinica showed the opposite trend. Phylogenetic analysis indicated that nifH gene sequences retrieved from DGGE gels clustered in the Proteobacteria and Cyanobacteria. Most of nifH gene fragments sequenced were not closely related to any known cultivated nitrogen-fixing bacteria and cyanobacteria, suggesting that the nifH gene sequences in wastelands of copper mine tailings investigated are unique and may represent novel sequences of nitrogen-fixing community. Our results indicated that diversity of nifH gene sequences in rhizosphere and non-rhizosphere tailings samples was affected by development period of plant community, plant species, plant life forms and physico-chemical properties of wastelands.
引文
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