不同作物与施肥对黑土氨氧化微生物的影响
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摘要
土壤微生物是农田生态系统的重要组成，在土壤养分转化、有机质分解、腐殖质形成、土壤肥力保持、营养元素的转化和循环中发挥了重要作用，而发挥作用的微生物属性是丰度和种群结构。影响丰度和种群结构的外界条件是水、热、气和营养物质，而水、热、气和营养物质又决定于农田管理方式如种植作物、肥料施用等直接或间接地影响土壤微生物群落结构及其功能的演化。本研究基于长期定位试验平台，利用新一代454高通量测序技术，以16S rRNA基因为分子标靶，原位条件下研究农田管理措施对土壤微生物群落结构的影响规律；进一步结合稳定性同位素示踪土壤微生物核酸DNA技术研究长期施肥条件下，土壤硝化微生物群落结构的主要变化规律及主要作用者。主要研究结果如下：
东北黑土区主要的种植制度下（大豆、小麦和玉米）田间原位研究结果表明：大豆对根际微生物总量影响最大，小麦次之，玉米影响最小。根际微生物的种群组成中，细菌占绝对优势。三种作物对土壤氨氧化细菌和古菌丰度影响不显著。施用氮肥导致土壤氨氧化细菌的丰度显著增加，而氨氧化古菌的丰度明显降低。施用磷肥对于氨氧化古菌和氨氧化细菌丰度的影响不显著。
温室盆栽实验研究发现：小麦、肥料、小麦肥料耦合作用三种因素对土壤微生物数量、酶活性和氨氧化微生物丰度的影响不同。小麦种植增加了微生物数量、脲酶和磷酸酶活性，增幅分别为39.0%、302%和38.5%；不同肥料处理对三者的影响分别为24.6%~47.0%、17.2%~32.7%和21.2%~25.9%；小麦施肥耦合作用的影响分别为3.30%~31.4%、59.0%~168%和15.2%~26.7%。小麦降低了土壤氨氧化细菌的丰度达44.1%，增加了氨氧化古菌的丰度达29.5%；氮肥显著增加了氨氧化细菌的丰度264%~313%，而磷肥对于氨氧化细菌增加仅为4.94%，氮磷肥均显著降低了氨氧化古菌的丰度达23.8%~48.5%；小麦和肥料耦合显著降低了氨氧化细菌的丰度，增加了氨氧化古菌的丰度。三种因素中小麦和肥料耦合对于氨氧化细菌和古菌的影响最大，最终表现为小麦-施肥较施肥显著增加了氨氧化细菌丰度，而降低了氨氧化细菌丰度。
针对长期定位试验的土壤样品，新一代高通量测序分析微生物16S rRNAgene，在整体水平上对长期不同施肥土壤中微生物群落变化进行研究。结果表明：长期施肥后，土壤微生物以不施肥土壤中微生物数量4.39%为分界点，长期施肥土壤中微生物总量>4.39%的土壤微生物生长被抑制；相反，促进了<4.39%的土壤微生物的生长。并且长期施肥引起了土壤微生物多样性的降低，同时，施用氮肥土壤微生物多样性较磷肥土壤微生物多样性低。
进一步采用稳定性同位素示踪（DNA-SIP）和新一代高通量测序技术，研究了长期施肥对东北黑土中氨氧细菌和古菌功能的影响。长期施肥引起土壤铵态氮浓度增加达8倍，并且土壤原位硝化活性（~(15)N示踪）显著增强，与此同时，氨氧化细菌丰度显著增加，而氨氧化古菌变化不显著。进一步利用稳定性同位素示踪CK和NPK土壤中的氨氧化微生物核酸DNA发现，氨氧化细菌在长期不施肥CK和施肥NPK土壤中均具有氨氧化功能，而氨氧化古菌仅在不施肥CK土壤中具有活性。这充分说明东北黑土长期施肥导致了土壤中氨氧化古菌功能的退化。
综上所述，在东北黑土中施用氮肥增加了土壤氨氧化细菌的丰度，降低了氨氧化古菌的丰度，并且氮肥的施用降低了土壤微生物多样性。黑土中长期施肥导致了氨氧化古菌功能的退化，氨氧化细菌功能的强化。
Soil microorganisms are an important part of soil, and they are the main power insoil nutrients transformation, soil organic matter resolving and the forming of humus.Besides, crops, fertilization and the interaction of crop-fertilization may influence thesoil microbial communities and the function in farm ecosystem. In this study, wetaking16S rRNA gene as molecular target and adopted the next generation454pyrosequencing to investigate the effect of farm management on the microbialabuncance and community. At the same time we adopted the454pyrosequencing andDNA-stable isotope probing technology (DNA-SIP) investigate the contribution ofammonia oxidizing bacteria and archaea under long-term fertilization. The mainconclusions were drowm as follows:
Based on Hailun Agricultural Ecology Station of Chinese Academy of Sciences,the microorganism (bacteria, fungi and actinomyces) and abundance ofammonia-oxidizing bacteria (AOB) and ammonia-oxidizing archaea (AOA) affectedby different crops (such as soybean, maize and wheat) were investigated. The resultsshowed that among the three different kinds of crops, the amount of bacteria wasmuch larger than that of fungus and actinomycetes in soils. The soil microorganismamount was significantly affected by different crops roots（P<0.05）and was in thefollowing order: soybean>wheat>maize. The abundance of AOB and AOA was notsignificantly influenced by different crops. The fertilization of nitrogen significantlyincreased the abundance of AOB in maize and wheat, and has no significant effect onsoybean, while the nitrogen fertilization decreased the abundance of AOA. Thephosphorus fertilization did not change the abundance of AOB and AOA abundance.
A pot experiment was carried out in greenhouse to study the effect of fertilization and root system of wheat on the soil microbial amount, the abundance of AOB andAOA, urease and phosphatase activity. The microbial amount, urease activity andphosphatase activity of treatment CK were significantly increased by39.0%,302%and38.5%by wheat. The microbial amount, urease activity and phosphatase activitywere significantly increased by24.6%~47.0%,17.2%~32.7%and21.2%~25.9%withfertilizer application but without wheat planting. Fertilization treatments with wheatplanting increased the soil microbial amount, urease activity and phosphatase activityby3.3%~31.4%,59.0%~168%and15.2%~26.7%, respectively. With wheat planting,the abundance of AOB was significantly decreased by44.1%, while the AOAabundance was increased by29.5%. N fertilization significantly increased the AOBabundance by264%~313%, while P fertilization only increased by the AOAabundance by4.94%. Fertilization significantly decreased the AOA abundance by23.8%~48.5%. Fertilization treatments with wheat planting significantly decreased theabundance of AOB and increased the abundance of AOA.
Based on21-year fertilization experiment, we found that after21-yearfertilization the microorganisms which proportion>4.39%in no fertilization soilwere suppressed, while the microorganisms which proportion <4.39%were promoted.And the long-term fertilization decreased the microorganism diversity, the Ninfluenced the microbial community diversity much more than the P fertilization.
Further more, the functional importance of AOB and AOA was studied asaffected by21-year fertilization.~(15)N-isotope tracing demonstrated that ammoniaoxidation activity was significantly stimulated under near-in situ condition infertilized NPK soils when compared to the control soil of CK treatment that receivedno fertilization. Pyrosequencing results clearly showed that AOB abundance wassignificantly increased while the AOA was not by the21-year fertilization.Furthermore,~(13)CO_2-based DNA stable isotope probing (SIP) suggested thefunctional importance of bacterial ammonia oxidation in both CK and NPK soils. However, strong proliferation of archaeal ammonia oxidizers was demonstrated incontrol but not fertilized soil.
Therefore, N fertilization significantly increased the AOB abundance anddecreased the AOA abundance, and decreased the microbial diversity. And long-termfertilization of urea leads to the functional extinction of AOA.
东北黑土区主要的种植制度下（大豆、小麦和玉米）田间原位研究结果表明：大豆对根际微生物总量影响最大，小麦次之，玉米影响最小。根际微生物的种群组成中，细菌占绝对优势。三种作物对土壤氨氧化细菌和古菌丰度影响不显著。施用氮肥导致土壤氨氧化细菌的丰度显著增加，而氨氧化古菌的丰度明显降低。施用磷肥对于氨氧化古菌和氨氧化细菌丰度的影响不显著。
温室盆栽实验研究发现：小麦、肥料、小麦肥料耦合作用三种因素对土壤微生物数量、酶活性和氨氧化微生物丰度的影响不同。小麦种植增加了微生物数量、脲酶和磷酸酶活性，增幅分别为39.0%、302%和38.5%；不同肥料处理对三者的影响分别为24.6%~47.0%、17.2%~32.7%和21.2%~25.9%；小麦施肥耦合作用的影响分别为3.30%~31.4%、59.0%~168%和15.2%~26.7%。小麦降低了土壤氨氧化细菌的丰度达44.1%，增加了氨氧化古菌的丰度达29.5%；氮肥显著增加了氨氧化细菌的丰度264%~313%，而磷肥对于氨氧化细菌增加仅为4.94%，氮磷肥均显著降低了氨氧化古菌的丰度达23.8%~48.5%；小麦和肥料耦合显著降低了氨氧化细菌的丰度，增加了氨氧化古菌的丰度。三种因素中小麦和肥料耦合对于氨氧化细菌和古菌的影响最大，最终表现为小麦-施肥较施肥显著增加了氨氧化细菌丰度，而降低了氨氧化细菌丰度。
针对长期定位试验的土壤样品，新一代高通量测序分析微生物16S rRNAgene，在整体水平上对长期不同施肥土壤中微生物群落变化进行研究。结果表明：长期施肥后，土壤微生物以不施肥土壤中微生物数量4.39%为分界点，长期施肥土壤中微生物总量>4.39%的土壤微生物生长被抑制；相反，促进了<4.39%的土壤微生物的生长。并且长期施肥引起了土壤微生物多样性的降低，同时，施用氮肥土壤微生物多样性较磷肥土壤微生物多样性低。
进一步采用稳定性同位素示踪（DNA-SIP）和新一代高通量测序技术，研究了长期施肥对东北黑土中氨氧细菌和古菌功能的影响。长期施肥引起土壤铵态氮浓度增加达8倍，并且土壤原位硝化活性（~(15)N示踪）显著增强，与此同时，氨氧化细菌丰度显著增加，而氨氧化古菌变化不显著。进一步利用稳定性同位素示踪CK和NPK土壤中的氨氧化微生物核酸DNA发现，氨氧化细菌在长期不施肥CK和施肥NPK土壤中均具有氨氧化功能，而氨氧化古菌仅在不施肥CK土壤中具有活性。这充分说明东北黑土长期施肥导致了土壤中氨氧化古菌功能的退化。
综上所述，在东北黑土中施用氮肥增加了土壤氨氧化细菌的丰度，降低了氨氧化古菌的丰度，并且氮肥的施用降低了土壤微生物多样性。黑土中长期施肥导致了氨氧化古菌功能的退化，氨氧化细菌功能的强化。
Soil microorganisms are an important part of soil, and they are the main power insoil nutrients transformation, soil organic matter resolving and the forming of humus.Besides, crops, fertilization and the interaction of crop-fertilization may influence thesoil microbial communities and the function in farm ecosystem. In this study, wetaking16S rRNA gene as molecular target and adopted the next generation454pyrosequencing to investigate the effect of farm management on the microbialabuncance and community. At the same time we adopted the454pyrosequencing andDNA-stable isotope probing technology (DNA-SIP) investigate the contribution ofammonia oxidizing bacteria and archaea under long-term fertilization. The mainconclusions were drowm as follows:
Based on Hailun Agricultural Ecology Station of Chinese Academy of Sciences,the microorganism (bacteria, fungi and actinomyces) and abundance ofammonia-oxidizing bacteria (AOB) and ammonia-oxidizing archaea (AOA) affectedby different crops (such as soybean, maize and wheat) were investigated. The resultsshowed that among the three different kinds of crops, the amount of bacteria wasmuch larger than that of fungus and actinomycetes in soils. The soil microorganismamount was significantly affected by different crops roots（P<0.05）and was in thefollowing order: soybean>wheat>maize. The abundance of AOB and AOA was notsignificantly influenced by different crops. The fertilization of nitrogen significantlyincreased the abundance of AOB in maize and wheat, and has no significant effect onsoybean, while the nitrogen fertilization decreased the abundance of AOA. Thephosphorus fertilization did not change the abundance of AOB and AOA abundance.
A pot experiment was carried out in greenhouse to study the effect of fertilization and root system of wheat on the soil microbial amount, the abundance of AOB andAOA, urease and phosphatase activity. The microbial amount, urease activity andphosphatase activity of treatment CK were significantly increased by39.0%,302%and38.5%by wheat. The microbial amount, urease activity and phosphatase activitywere significantly increased by24.6%~47.0%,17.2%~32.7%and21.2%~25.9%withfertilizer application but without wheat planting. Fertilization treatments with wheatplanting increased the soil microbial amount, urease activity and phosphatase activityby3.3%~31.4%,59.0%~168%and15.2%~26.7%, respectively. With wheat planting,the abundance of AOB was significantly decreased by44.1%, while the AOAabundance was increased by29.5%. N fertilization significantly increased the AOBabundance by264%~313%, while P fertilization only increased by the AOAabundance by4.94%. Fertilization significantly decreased the AOA abundance by23.8%~48.5%. Fertilization treatments with wheat planting significantly decreased theabundance of AOB and increased the abundance of AOA.
Based on21-year fertilization experiment, we found that after21-yearfertilization the microorganisms which proportion>4.39%in no fertilization soilwere suppressed, while the microorganisms which proportion <4.39%were promoted.And the long-term fertilization decreased the microorganism diversity, the Ninfluenced the microbial community diversity much more than the P fertilization.
Further more, the functional importance of AOB and AOA was studied asaffected by21-year fertilization.~(15)N-isotope tracing demonstrated that ammoniaoxidation activity was significantly stimulated under near-in situ condition infertilized NPK soils when compared to the control soil of CK treatment that receivedno fertilization. Pyrosequencing results clearly showed that AOB abundance wassignificantly increased while the AOA was not by the21-year fertilization.Furthermore,~(13)CO_2-based DNA stable isotope probing (SIP) suggested thefunctional importance of bacterial ammonia oxidation in both CK and NPK soils. However, strong proliferation of archaeal ammonia oxidizers was demonstrated incontrol but not fertilized soil.
Therefore, N fertilization significantly increased the AOB abundance anddecreased the AOA abundance, and decreased the microbial diversity. And long-termfertilization of urea leads to the functional extinction of AOA.
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