垦殖与自然恢复黑土微生物群落结构及生态功能的季节变化
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摘要
东北黑土有机碳含量较高，是土壤碳库的重要组成部分。土壤有机碳变化与土地利用方式、气候条件及土壤微生物群落结构关系密切。黑土区气候四季分明，土壤温湿度及地表覆盖物变化明显，这些都预示着土壤微生物群落结构及功能可能存在季节性差异。本论文以中国科学院海伦农业生态试验站为研究平台，选取垦殖3种不同的施肥处理（无肥NoF、化肥CF和化肥配施有机肥CFM）和自然恢复黑土（NR）为研究对象，于不同季节采集土壤样品，通过测定可溶性有机碳、微生物量碳研究土壤活性有机碳库季节性结构组成；采用PCR-DGGE和克隆测序相结合的方法研究土壤微生物群落结构季节性变化规律；通过Biolog法比较微生物对底物碳源利用上的差异，同时测定土壤呼吸和土壤酶活性，解析土壤微生物生态功能的季节性差异，主要研究结果如下：
土壤微生物量碳的最高值都出现在积雪融化期，NR处理土壤微生物量碳5个采样时期平均值较NoF、CF和CFM处理分别提高102%、89%和52%；土壤可溶性有机碳含量总体表现为夏季比冬季高；同一处理在不同季节土壤呼吸差异显著，4种处理在不同温度条件下的土壤呼吸具有相似的变化趋势，均表现为NR处理高于垦殖处理。NR处理的过氧化氢酶、转化酶和磷酸酶活性在5个采样时期均具有较高活性，而NR处理的脲酶活性只在作物生长季节较高。单施化肥或化肥配施有机肥可以提高转化酶、磷酸酶和脲酶活性，而单施化肥有抑制过氧化氢酶活性的作用。除垦殖的3种施肥处理土壤转化酶活性较稳定、季节性变化不显著外，所有处理4种土壤酶活性均表现出明显的季节性变化。5个采样时期的土壤微生物量碳与土壤酶活性之间的正相关关系体现不明显。
采用PCR-DGGE和克隆测序相结合的方法研究土壤细菌和真菌群落结构的季节性变化规律，结果表明，垦殖处理与自然恢复处理相比，前者改变了土壤微生物群落结构；土壤细菌群落结构在植物非生长季节3个采样时期相似度高而在植物生长季节相似度较低；真菌对植被、土壤理化性状和垦殖的敏感性要高于细菌，真菌在作物生长季节（2007年8月28日）和作物收获期（2006年10月15日）的土壤样品群落相似度较高。DGGE条带的测序结果显示，不可培养微生物比例较高，变形杆菌门（Proteobacteria（α-, β-and γ-））和放线菌门（Actinobacteria）是细菌的主要成员；担子菌门（Ascomycota）和子囊菌门（Basidiomycota）是真菌的主要成员。
黑土区氨氧化细菌分布在目前已知的cluster1、cluster2、cluster3、cluster4、cluster7、cluster9和cluster10当中，其中Nitrosospira cluster3是黑土区农田当中的优势氨氧化细菌。土地利用方式改变了氨氧化细菌的群落结构，同时垦殖与自然恢复黑土当中的氨氧化细菌也存在季节性变化，表现为有作物生长季节之间以及非作物生长季节之间群落相似性较高。另外，本研究还发现一类目前尚不能分类到任何已知氨氧化细菌cluster当中的、黑土区特有的氨氧化细菌类群。
对4℃、15℃和28℃培养温度下4个处理黑土微生物代谢功能研究表明，在同一采样时期不同培养温度下，随着培养温度的升高每个处理的微生物代谢活性提高，不同采样季节垦殖与自然恢复处理在3个培养温度下微生物对碳源的代谢能力总体均呈现NR＞CFM＞CF＞NoF变化趋势。微生物在积雪融化期2009年3月27日对碳源的利用能力较强。28℃培养温度下，NoF处理较其它3种处理微生物代谢活性有较大不同，夏季作物生长季节CF、CFM和NR3个处理细菌群落对碳源的利用能力差异最大。
The black soils in Northeast China are commonly fertile and productive withhigh organic carbon content, which is closely related with soil management practice,climate and soil microorganisms. The climate in Northeast China is characterized bythe extreme annual variation in temperature and precipitation with hot and wet insummer and cold and dry in winter. Consequently, seasonal fluctuations of soiltemperature, soil water content and plant cover are very large for black soils, whichlead to the prediction that soil microbial communities show distinct seasonalvariations in this region. A long term fertilization experimental station which has threefertilizer treatments: no fertilizer (NoF), chemical fertilizers (CF) and chemicalfertilizers plus manure (CFM), and natural restoration (NR) was established in1985.Soil samples (0-20cm depth) were collected on different seasons. Soil dissolvedorganic carbon (DOC) and microbial biomass carbon (MBC) were measured toexplore the seasonal shifts of the active soil organic carbon pool. PCR, DGGE, cloneand sequencing were applied to compare the seasonal variations of microbialcommunities. Microbial metabolic activities (Biolog), soil respiration and soil enzymewere also measured to elucidate how microbial functions changed among differentseasons and treatments. The results were as follows:
The value of MBC was highest in spring and the mean value of NR of fivesampling times was higher102%、89%and52%than NoF、CF and CFM, respectively.Soil organic carbon content was higher in samples collected in summer comparedwith those collected in winter. Soil respiration rate from a same treatment but differentseasons was significantly different, and NR was the highest compared with other threecultivated treatments regardless the incubation temperature. NR has the highestactivities of soil catalase, invertase and phosphatase in five sampling times, while thehighest urease activities of NR only appeared in plant growing season. Theapplication of fertilization increased the activities of soil invertase, phosphatase andurease, but the activities of catalase were decreased when only applied chemicalfertilizer. The activities of soil enzymes had obvious seasonal shifts except theactivities of soil invertase in NoF, CF and CFM. The positive relationship betweenMBC and soil enzymes was not obvious in five sampling times.
The dominant members of the soil microbial communities were primarilyregulated by land use and also changed seasonally associated with plant growth inblack soils in the region. There was no distinct difference in dominant members ofboth bacterial and fungal communities among different fertilizer treatment plots incultivated black soils. The change of dominant members of fungal communities alongwith land use and season was more obvious than that of bacterial communities,suggesting greater sensitivity of fungi to vegetation, soil disturbance of tillage and soilphysico-chemical properties. Sequence analysis of the DGGE bands revealed thatbacteria belonging to Proteobacteria and Actinobacteria, and fungi of theBasidiomycota and Ascomycota inhabit predominantly in the black soil.
Ammonia oxidizing bacteria (AOB) in black soils were distributed in cluster1,cluster2, cluster3, cluster4, cluster7, cluster9and cluster10by phylogeneticanalysis, while Nitrosospira cluster3was dominant in cultivated black soils andNitrosospira cluster1was dominant in NR. The PCoA analysis showed that AOBcommunities of plant growing seasons and no plant growing seasons had highsimilarities, respectively. We found a new cluster (defined as cluster NR in this study)which was not affiliated with any known AOB cluster.
Biolog analysis revealed that the microbial metabolic activities (expressed asAWCD，average well color development) of samples from a same sampling time butunder different incubation temperature followed as the order of：28℃＞15℃＞4℃.Samples collected in spring had higher AWCD. For each incubation temperature,AWCD of samples from different sampling time followed as the order of：NR＞CFM＞CF＞NoF. Principal component analysis (28℃,72h) showed that the metabolicactivities of NoF had low similarities with other treatments in four sampling time andmetabolic activities of CF、CFM and NR in plant growing season were obviousdifferent.
土壤微生物量碳的最高值都出现在积雪融化期，NR处理土壤微生物量碳5个采样时期平均值较NoF、CF和CFM处理分别提高102%、89%和52%；土壤可溶性有机碳含量总体表现为夏季比冬季高；同一处理在不同季节土壤呼吸差异显著，4种处理在不同温度条件下的土壤呼吸具有相似的变化趋势，均表现为NR处理高于垦殖处理。NR处理的过氧化氢酶、转化酶和磷酸酶活性在5个采样时期均具有较高活性，而NR处理的脲酶活性只在作物生长季节较高。单施化肥或化肥配施有机肥可以提高转化酶、磷酸酶和脲酶活性，而单施化肥有抑制过氧化氢酶活性的作用。除垦殖的3种施肥处理土壤转化酶活性较稳定、季节性变化不显著外，所有处理4种土壤酶活性均表现出明显的季节性变化。5个采样时期的土壤微生物量碳与土壤酶活性之间的正相关关系体现不明显。
采用PCR-DGGE和克隆测序相结合的方法研究土壤细菌和真菌群落结构的季节性变化规律，结果表明，垦殖处理与自然恢复处理相比，前者改变了土壤微生物群落结构；土壤细菌群落结构在植物非生长季节3个采样时期相似度高而在植物生长季节相似度较低；真菌对植被、土壤理化性状和垦殖的敏感性要高于细菌，真菌在作物生长季节（2007年8月28日）和作物收获期（2006年10月15日）的土壤样品群落相似度较高。DGGE条带的测序结果显示，不可培养微生物比例较高，变形杆菌门（Proteobacteria（α-, β-and γ-））和放线菌门（Actinobacteria）是细菌的主要成员；担子菌门（Ascomycota）和子囊菌门（Basidiomycota）是真菌的主要成员。
黑土区氨氧化细菌分布在目前已知的cluster1、cluster2、cluster3、cluster4、cluster7、cluster9和cluster10当中，其中Nitrosospira cluster3是黑土区农田当中的优势氨氧化细菌。土地利用方式改变了氨氧化细菌的群落结构，同时垦殖与自然恢复黑土当中的氨氧化细菌也存在季节性变化，表现为有作物生长季节之间以及非作物生长季节之间群落相似性较高。另外，本研究还发现一类目前尚不能分类到任何已知氨氧化细菌cluster当中的、黑土区特有的氨氧化细菌类群。
对4℃、15℃和28℃培养温度下4个处理黑土微生物代谢功能研究表明，在同一采样时期不同培养温度下，随着培养温度的升高每个处理的微生物代谢活性提高，不同采样季节垦殖与自然恢复处理在3个培养温度下微生物对碳源的代谢能力总体均呈现NR＞CFM＞CF＞NoF变化趋势。微生物在积雪融化期2009年3月27日对碳源的利用能力较强。28℃培养温度下，NoF处理较其它3种处理微生物代谢活性有较大不同，夏季作物生长季节CF、CFM和NR3个处理细菌群落对碳源的利用能力差异最大。
The black soils in Northeast China are commonly fertile and productive withhigh organic carbon content, which is closely related with soil management practice,climate and soil microorganisms. The climate in Northeast China is characterized bythe extreme annual variation in temperature and precipitation with hot and wet insummer and cold and dry in winter. Consequently, seasonal fluctuations of soiltemperature, soil water content and plant cover are very large for black soils, whichlead to the prediction that soil microbial communities show distinct seasonalvariations in this region. A long term fertilization experimental station which has threefertilizer treatments: no fertilizer (NoF), chemical fertilizers (CF) and chemicalfertilizers plus manure (CFM), and natural restoration (NR) was established in1985.Soil samples (0-20cm depth) were collected on different seasons. Soil dissolvedorganic carbon (DOC) and microbial biomass carbon (MBC) were measured toexplore the seasonal shifts of the active soil organic carbon pool. PCR, DGGE, cloneand sequencing were applied to compare the seasonal variations of microbialcommunities. Microbial metabolic activities (Biolog), soil respiration and soil enzymewere also measured to elucidate how microbial functions changed among differentseasons and treatments. The results were as follows:
The value of MBC was highest in spring and the mean value of NR of fivesampling times was higher102%、89%and52%than NoF、CF and CFM, respectively.Soil organic carbon content was higher in samples collected in summer comparedwith those collected in winter. Soil respiration rate from a same treatment but differentseasons was significantly different, and NR was the highest compared with other threecultivated treatments regardless the incubation temperature. NR has the highestactivities of soil catalase, invertase and phosphatase in five sampling times, while thehighest urease activities of NR only appeared in plant growing season. Theapplication of fertilization increased the activities of soil invertase, phosphatase andurease, but the activities of catalase were decreased when only applied chemicalfertilizer. The activities of soil enzymes had obvious seasonal shifts except theactivities of soil invertase in NoF, CF and CFM. The positive relationship betweenMBC and soil enzymes was not obvious in five sampling times.
The dominant members of the soil microbial communities were primarilyregulated by land use and also changed seasonally associated with plant growth inblack soils in the region. There was no distinct difference in dominant members ofboth bacterial and fungal communities among different fertilizer treatment plots incultivated black soils. The change of dominant members of fungal communities alongwith land use and season was more obvious than that of bacterial communities,suggesting greater sensitivity of fungi to vegetation, soil disturbance of tillage and soilphysico-chemical properties. Sequence analysis of the DGGE bands revealed thatbacteria belonging to Proteobacteria and Actinobacteria, and fungi of theBasidiomycota and Ascomycota inhabit predominantly in the black soil.
Ammonia oxidizing bacteria (AOB) in black soils were distributed in cluster1,cluster2, cluster3, cluster4, cluster7, cluster9and cluster10by phylogeneticanalysis, while Nitrosospira cluster3was dominant in cultivated black soils andNitrosospira cluster1was dominant in NR. The PCoA analysis showed that AOBcommunities of plant growing seasons and no plant growing seasons had highsimilarities, respectively. We found a new cluster (defined as cluster NR in this study)which was not affiliated with any known AOB cluster.
Biolog analysis revealed that the microbial metabolic activities (expressed asAWCD，average well color development) of samples from a same sampling time butunder different incubation temperature followed as the order of：28℃＞15℃＞4℃.Samples collected in spring had higher AWCD. For each incubation temperature,AWCD of samples from different sampling time followed as the order of：NR＞CFM＞CF＞NoF. Principal component analysis (28℃,72h) showed that the metabolicactivities of NoF had low similarities with other treatments in four sampling time andmetabolic activities of CF、CFM and NR in plant growing season were obviousdifferent.
引文
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