安徽铜陵铜尾矿原生演替过程中的土壤固氮菌研究
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摘要
生物可利用氮的缺乏是原生演替早期生物生长的主要限制因子之一。固氮微生物通过生物固氮作用为这种氮素匾乏的生态系统提供稳定的N来源,提高土壤基质的营养水平,从而促进后期演替物种成功定居,固氮菌在尾矿原生演替进程中扮演重要角色。
本文以安徽铜陵已弃置20年铜尾矿的原生演替过程为模式,选择裸地(BT)、藻类结皮(AC)、苔藻混合结皮(AMC)、苔藓结皮(MC)和高等植物覆盖区域(VEG)作为尾矿原生演替的系列样地;选择邻近山上土壤结构好且物种丰富的草本植被区作为演替后期的对照点(CK)。通过末端限制性片段长度多态性(TRFLP)、克隆文库和荧光定量PCR(qPCR)等方法,结合土壤理化特征相关分析,阐明铜尾矿原生演替过程中固氮微生物的群落结构及其丰度变化规律。主要研究结果如下:
(1)固氮菌多样性在尾矿演替过程中总体上呈现先升后降的趋势,其中以裸地最低,生物结皮系列最高。这些固氮微生物主要分布在蓝藻门和变形细菌门(主要是α-Proteobacteria),前者主要存在于结皮系列中,而后者在演替各个阶段都有分布。生物结皮是尾矿原生演替过程中的一个重要阶段,而固氮蓝藻是生物结皮系列主要的N素来源。
(2)铜尾矿中固氮菌数量丰富。在所研究的样地中,每克土含有的nifH基因拷贝数在5.06×10~5(BT)和3.47×10~8(CK)之间,总体来说,随着铜尾矿原生演替的发展,固氮微生物数量呈现逐渐增加的趋势。nifH基因拷贝数和土壤重金属含量负相关(p < 0.05),而与碳氮水平呈显著正相关(p < 0.01),这可能与演替后期生物结皮和高等植物在改善土壤基质和减少环境干扰方面的贡献有关。
(3)铜尾矿原生演替过程(BT-BSCs-VEG)是一个土壤基质发展的过程,固氮微生物在演替早期发挥重要作用,主要表现为缓解土壤氮素缺乏的环境胁迫,为后期生物结皮和高等植物的定居提供必需的氮素来源。
本研究通过比较铜尾矿原生演替过程中固氮微生物的种类组成及其丰度变化,揭示固氮菌在促进原生演替进程中的作用,对完善尾矿废弃地生态恢复理论和指导尾矿生态修复实践具有重要意义。
Bio-available nitrogen in primary successional series is one of the key limitations for ecosystem development. N-fixing microorganisms often serve as early and abundant colonizers in these N-deficient terrestrial ecosystems and are tightly linked to the accumulation of soil N. Consequently, diazotrophic communities are a primary driving force for the improvement of substrate nutritional status and other physiochemical characteristics which facilitate recolonization of plants and other macroorganisms during later successional stages.
Based on detailed ecological surveys and the landscape pattern, we selected the bare tailings (BT), algae crust (AC), moss/algae crust (AMC), moss crust (MC) and vegetated area (VEG) as succession series of copper mine tailings at Tongling, Anhui Province, P.R. China. An undisturbed off-site control area (CK) with good soil structure and high herbaceous diversity was chosen for comparison purpose.
The abundance and diversity of N-fixing community along the succession series of mine tailings were determined using multiple molecular approaches including terminal restriction fragment length polymorphism (TRFLP), clone library analysis and real time quantitative PCR (qPCR) of the nifH gene. The main results were summarized as follows:
1) In general, the diazotrophic diversity was lowest in BT, and there was an increase in the BSCs followed by a decrease in site VEG. Nitrogen-fixing microorganisms detected are mainly affiliated with the Proteobacteria and Cyanobacteria, with the former present in all series while the latter exclusively associated with the BSCs. It is likely that nitrogen-fixng Cyanobacteria are the dominant N suppliers in the BSCs, a crucial stage during the primary succession.
2) Real-time PCR revealed that diazotrophs were abundant in the copper mine tailings. The nifH gene number ranged from 5.06×10~5(BT)to 3.47×10~8(CK)copies per gram of soil (dried weight).Variations in the nifH gene abundance showed a significant positive correlation with changes in soil C and N concentrations (p < 0.01) and a negative correlation with heavy metal (Cu and Zn) concentrations (p < 0.05). Overall, the number of nitrogen fixing bacteria increased with the progressing succession. This could be attributed to the new environmental niches and especially nutrient resources opened by the BSCs and plants in the later successional stages.
3) Primary succession was a process with soil development. In newly unvegetated and nitrogen-deficient tailings, diazotrophs played an important role in accumulating soil N, which was indispensible for further recolonization of biological soil crusts (BSCs) and plants.
This represents the first report using cultivation-independent molecular approaches to elucidate the diazotrophic diversity and community structure evolution associated with a successional series of mine tailings. Our findings could be of importance in developing ecological theory and restoration practice in mine tailings.
本文以安徽铜陵已弃置20年铜尾矿的原生演替过程为模式,选择裸地(BT)、藻类结皮(AC)、苔藻混合结皮(AMC)、苔藓结皮(MC)和高等植物覆盖区域(VEG)作为尾矿原生演替的系列样地;选择邻近山上土壤结构好且物种丰富的草本植被区作为演替后期的对照点(CK)。通过末端限制性片段长度多态性(TRFLP)、克隆文库和荧光定量PCR(qPCR)等方法,结合土壤理化特征相关分析,阐明铜尾矿原生演替过程中固氮微生物的群落结构及其丰度变化规律。主要研究结果如下:
(1)固氮菌多样性在尾矿演替过程中总体上呈现先升后降的趋势,其中以裸地最低,生物结皮系列最高。这些固氮微生物主要分布在蓝藻门和变形细菌门(主要是α-Proteobacteria),前者主要存在于结皮系列中,而后者在演替各个阶段都有分布。生物结皮是尾矿原生演替过程中的一个重要阶段,而固氮蓝藻是生物结皮系列主要的N素来源。
(2)铜尾矿中固氮菌数量丰富。在所研究的样地中,每克土含有的nifH基因拷贝数在5.06×10~5(BT)和3.47×10~8(CK)之间,总体来说,随着铜尾矿原生演替的发展,固氮微生物数量呈现逐渐增加的趋势。nifH基因拷贝数和土壤重金属含量负相关(p < 0.05),而与碳氮水平呈显著正相关(p < 0.01),这可能与演替后期生物结皮和高等植物在改善土壤基质和减少环境干扰方面的贡献有关。
(3)铜尾矿原生演替过程(BT-BSCs-VEG)是一个土壤基质发展的过程,固氮微生物在演替早期发挥重要作用,主要表现为缓解土壤氮素缺乏的环境胁迫,为后期生物结皮和高等植物的定居提供必需的氮素来源。
本研究通过比较铜尾矿原生演替过程中固氮微生物的种类组成及其丰度变化,揭示固氮菌在促进原生演替进程中的作用,对完善尾矿废弃地生态恢复理论和指导尾矿生态修复实践具有重要意义。
Bio-available nitrogen in primary successional series is one of the key limitations for ecosystem development. N-fixing microorganisms often serve as early and abundant colonizers in these N-deficient terrestrial ecosystems and are tightly linked to the accumulation of soil N. Consequently, diazotrophic communities are a primary driving force for the improvement of substrate nutritional status and other physiochemical characteristics which facilitate recolonization of plants and other macroorganisms during later successional stages.
Based on detailed ecological surveys and the landscape pattern, we selected the bare tailings (BT), algae crust (AC), moss/algae crust (AMC), moss crust (MC) and vegetated area (VEG) as succession series of copper mine tailings at Tongling, Anhui Province, P.R. China. An undisturbed off-site control area (CK) with good soil structure and high herbaceous diversity was chosen for comparison purpose.
The abundance and diversity of N-fixing community along the succession series of mine tailings were determined using multiple molecular approaches including terminal restriction fragment length polymorphism (TRFLP), clone library analysis and real time quantitative PCR (qPCR) of the nifH gene. The main results were summarized as follows:
1) In general, the diazotrophic diversity was lowest in BT, and there was an increase in the BSCs followed by a decrease in site VEG. Nitrogen-fixing microorganisms detected are mainly affiliated with the Proteobacteria and Cyanobacteria, with the former present in all series while the latter exclusively associated with the BSCs. It is likely that nitrogen-fixng Cyanobacteria are the dominant N suppliers in the BSCs, a crucial stage during the primary succession.
2) Real-time PCR revealed that diazotrophs were abundant in the copper mine tailings. The nifH gene number ranged from 5.06×10~5(BT)to 3.47×10~8(CK)copies per gram of soil (dried weight).Variations in the nifH gene abundance showed a significant positive correlation with changes in soil C and N concentrations (p < 0.01) and a negative correlation with heavy metal (Cu and Zn) concentrations (p < 0.05). Overall, the number of nitrogen fixing bacteria increased with the progressing succession. This could be attributed to the new environmental niches and especially nutrient resources opened by the BSCs and plants in the later successional stages.
3) Primary succession was a process with soil development. In newly unvegetated and nitrogen-deficient tailings, diazotrophs played an important role in accumulating soil N, which was indispensible for further recolonization of biological soil crusts (BSCs) and plants.
This represents the first report using cultivation-independent molecular approaches to elucidate the diazotrophic diversity and community structure evolution associated with a successional series of mine tailings. Our findings could be of importance in developing ecological theory and restoration practice in mine tailings.
引文
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黄铭洪(2003)环境污染与生态恢复。北京:科学出版社。
束文圣,杨开颜,张志权,杨兵.蓝崇钰(2001)湖北铜绿山古铜矿冶炼渣植被与优势植物的重金属含量研究。应用与环境生物学报7:7-12。
束文圣,叶志鸿,张志权,黄铭洪,蓝崇钰(2003)华南铅锌尾矿生态恢复的理论与实践。生态学报23:1629-1639。
许宝健(2009)危险的隐患沉寂的金山。中国环境报第6版(2009年3月6日)。
祝玉学(1998)关于尾矿库工程中几个问题的讨论。金属矿山10:7-10。