我国亚热带土壤碳氮转化微生物及其功能研究
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摘要
我国亚热带地区面积辽阔,占国土面积的21%,是重要的农业生产区。该地区酸性土壤在世界上占有独特的地位,该地区土壤具有pH低、高度风化、氧化势高等特点。我国亚热带地区土壤碳、氮生物地球化学循环与环境变化、农业生产等密切相关,了解碳、氮转化微生物特性及其生态功能对把握该地区碳、氮地球生物化学循环规律、发展该地区可持续绿色农业生产具有指导意义。
本研究选取亚热带地区典型水稻种植区(湖北武汉)稻田土壤为研究对象,测定田间原位和温室盆栽条件下种植转Bt基因水稻(以下简称Bt水稻)对土壤主要温室气体(CH4、CO2和N2O)排放的影响,采用定量PCR (Quantitative PCR, qPCR)、PCR-DGGE等现代分子生态学手段测定种植Bt水稻对温室气体排放相关功能微生物的影响,并采用稳定同位素核酸标记技术来分析种植转Bt基因水稻对土壤功能“活跃的”产甲烷古菌群落的影响,以了解该地区C转化微生物群落特征及其功能。主要结果如下：
(1)田间原位和温室盆栽实验结果均显示种植Bt水稻显著的、持久的降低了土壤CH4、N2O和CO2排放通量,对产甲烷古菌、甲烷氧化细菌、产N2O功能微生物、N2O还原功能微生物群落也有显著影响。种植Bt水稻显著的、持久的降低了土壤CH4排放通量,对根际产甲烷古菌和甲烷氧化细菌群落丰度和组成(香农多样性指数H’)也有显著的负面影响。在S2和S5阶段,Bt水稻根际产N2O功能微生物丰度显著低于Ck水稻(p<0.05),在S3阶段Bt水稻根际产N2O功能微生物丰度显著高于Ck水稻(p<0.05),而在S1和S4阶段,两种水稻根际产N2O功能微生物丰度无显著差异(p>0.05)。在S2阶段,Bt水稻根际N2O还原功能微生物丰度显著低于Ck水稻(p<0.05),在S1和S4阶段Bt水稻根际N2O还原功能微生物丰度显著高于Ck水稻(p<0.05),而在S3和S5阶段,两种水稻根际N2O还原功能微生物丰度无显著差异(p>0.05)。在苗期(S1)和分蘖期(S2),Bt水稻根际总细菌丰度显著低于Ck水稻(p<0.05)；而在晒田期(S3)、灌浆期(S4)和成熟期(S5),这种显著差异消失(p>0.05)。在S3阶段,Bt水稻根际总古菌丰度显著高于Ck水稻(p<0.05)；在其它生长阶段,Bt水稻和Ck水稻根际总古菌丰度无显著差异(p>0.05)。相关分析结果显示土壤CH4排放通量与产甲烷古菌丰度成显著正相关(R2=0.564,p<0.05)；土壤N2O排放通量与产N2O功能微生物丰度成显著正相关(R2=0.564,p<0.05)。推测：与Ck水稻相比,Bt水稻根系分泌物含量较低,产甲烷微生物、产N2O功能微生物丰度较低,苗期和分蘖期植株生物量较低,土壤氧化还原电位较高等是导致Bt水稻土壤温室气体排放通量较低的主要原因。
(2)稳定同位素核酸标记结果显示种植Bt水稻显著降低了功能“活跃的”产甲烷古菌群落丰度,但没有显著影响功能“活跃的”产甲烷古菌群落组成。Bt水稻和Ck水稻分蘖期根际土壤中功能“活跃的”产甲烷古菌类群有Rice cluster-Ⅰ (RC-Ⅰ)、Methanosaetaceae、Methanosarcinaceae和Methanobacteriaceae,其中RC-I占所有功能“活跃的”产甲烷古菌类型的60%～66.7%,为最活跃的产甲烷古菌类群。
我们还选取了亚热带地区福建万木林自然保护区典型森林土壤(细柄阿丁枫,ALG;浙江桂,CIC)及由其转变的农业利用土壤(杉木,CUL；桔园,ORG)为研究对象,采用qPCR, PCR-DGGE等现代分子生态学手段测定氨氧化古菌(ammonia-oxidizing archaea, AOA)和氨氧化细菌(ammonia-oxidizing bacteria, AOB)群落丰度和组成,研究森林土壤向农田土壤转变对土壤氨氧化微生物群落及其氨氧化功能的影响,以表征该地区N转化微生物群落特征及其功能。主要结果如下：
(1)人为利用和施肥显著提高了我国亚热带地区土壤氨氧化势。该地区森林土壤转变为农田土壤显著增加了AOA和AOB的群落丰度,且显著减小了AOA和AOB丰度比值。DGGE指纹图谱显示森林土壤转变为农田土壤也明显改变了AOA和AOB群落组成,其中隶属于Group1.la-associated lineage的AOA类群(A7～A10)在施肥利用后没有检测到,而隶属于Cluster3a lineage的AOB类群(B6)在施肥利用后也没有检测到。
(2)添加尿素培养后,天然林(ALG和CIC)土壤AOA丰度显著增加,AOB丰度无显著变化；未施肥种植林(CUL)土壤AOA和AOB丰度均显著增加;而施肥土壤(ORG) AOA和AOB丰度均无显著变化。添加尿素培养明显改变了AOA群落结构；但没有显著改变AOB群落结构。可见,天然林(ALG和CIC)土壤中AOA对添加尿素培养有明显响应；种植林土壤(CUL)和施肥土壤(ORG)中AOA和AOB对添加尿素培养有明显响应。因此,我们推断天然林(ALG和CIC)土壤中AOA对自养氨氧化过程贡献大,种植林土壤(CUL)和施肥土壤(ORG)中AOA和AOB对自养氨氧化过程均有贡献。即森林土壤向农田土壤转变改变了AOA和AOB对硝化作用的贡献,但相对贡献率尚不能确定。
The subtropical region of China, cover about21%of the country's land surface, is the main area of agricultural production in our country. Soils in this region, characterized with low pH, highly weathered, high oxidation potential, play a unique role in the world. Soil carbon and nitrogen biogeochemical cycles in this region are closely related to environmental change, agricultural production, etc. Therefore, it will be instructive to understand microbial characteristics and their functions related to carbon-nitrogen transformations for grasping the laws of carbon and nitrogen biogeochemical cycles and developing sustainable green agricultural production.
Typical rice-growing area in subtropical region (Wuhan, Hubei province, China) was selected to be the study site. Static closed chamber technique was employed to analyze the effect of planting transgenic Bt rice on greenhouse gases (GHQ such as CH4, CO2and N2O) emission from paddy soil under both field and greenhouse conditions. Modern molecular ecology methods were employed to investigate the effect of planting transgenic Bt rice on functional microorganisms related to GHGs emission. DNA-based stable isotope probing (DNA-SIP) method was employed to investigate the effect of planting transgenic Bt rice on functionally "active" methanogenic archaea. The objective of this study was to understand the microbial characteristics of carbon transformation and the functions in subtropical region. The results are as follows:
(1) Under both field and greenhouse conditions, planting transgenic Bt rice significantly and persistently reduced the CH4, N2O and CO2emission. Planting transgenic Bt rice did some significant and negative effects on community abundances and composition (characterized by Shannon-Weuner diversity index) of rhizospheric methanogenic archaea and methanotrophic bacteria. The community abundances of total bacteria in Bt rice rhizosphere at SI and S2stages were significantly lower than Ck rice rhizosphere (p<0.05), but these significant differences disappeared at following stages. The community abundance of total archaea in Bt rice rhizosphere at S3stage was significantly lower than Ck rice rhizosphere (p<0.05), but these significant differences disappeared at following stages. Regression analysis shown that the CH4emission flux significantly increased with the increasing of methanogenic archaeal abundance (R2=0.839, p<0.001), and the N2O emission flux significantly increased with the increasing of N2O production microorganism abundance (R2=0.564, p<0.05). We inferred that the lower root exudates contents, plant biomass and functional microorganism abudance and the higher soil redox potential of Bt rice were the key reasons resulting in the lower greenhouse gases emission fluxes.
(2) Results of SIP-DNA indicated that planting transgenic Bt rice significantly reduced the community abundance of functionally "active" methanogenic archaea, but did not affect the community composition. The populations of functional "active" methanogenic archaea in Bt and Ck rice rhizosphere were Rice cluster-I (RC-I), Methanosaetaceae, Methanosarcinaceae and Methanobacteriacea. RC-I is represented by the most species, accounting for60%～66.7%of total functional "active" methanogenic archaeal populations.
Typical forest soils (Altingia gralilipes, ALG; Cinnamomum chekiangense, CIC) and changed agriculture soils (Cunninghamia lanceolata, CUL; Orange orchard, ORG) in subtropical region were selected to be another study site. Modern molecular ecology methods were employed to investigate the effect of forest conversion to agriculture on ammonia oxidizers and their contributions to autotrophic ammonia oxidation process. The objective of this study was to understand the microbial characteristics of nitrogen transformation and the functions in subtropical region. The results are as follows:
(1) People use and fertilization significantly increased soil potential ammonia oxidation in this region. Forest conversion to agriculture significantly increased ammonia-oxidizing archaea (AOA) and ammonia-oxidizing bacteria (AOB) community abundances, and changed AOA and AOB community structures. The relative contributions of AOA and AOB to autotrophic ammonia oxidation process were also changed after the land-use conversion. Forest conversion to agriculture significantly changed the community compositions of AOA and AOB. The AOA populations (A7-A10), belong to Group1.la-associated lineage, could not be detected after fertilizing. The AOB population (B6), belong to Cluster3a lineage, could not be detected after fertilizing.
(2) The Urea addition significantly increased the AOA abundance in natural forests (ALG and CIC), significantly promoted the AOA and AOB abundances in unfertilized plantation soil (CUL), but had no significant effect on the AOB abundance in natural forests and the AOA and AOB abundances in fertilized soil (ORG). The Urea addition significantly changed the community compositions of AOA, but had no significant effect on the community compositions of AOB. We inferred that AOA may dominate the autotrophic ammonia oxidation process in natural forests (ALG and CIC), whereas both AOA and AOB may conduct autotrophic ammonia oxidation in agriculture soils (CUL and ORG). We supported that forest conversion to agriculture significantly changed the relative contributions of AOA and AOB to autotrophic ammonia oxidation, but the relative contribution ratio in these soils still remained unknown.
本研究选取亚热带地区典型水稻种植区(湖北武汉)稻田土壤为研究对象,测定田间原位和温室盆栽条件下种植转Bt基因水稻(以下简称Bt水稻)对土壤主要温室气体(CH4、CO2和N2O)排放的影响,采用定量PCR (Quantitative PCR, qPCR)、PCR-DGGE等现代分子生态学手段测定种植Bt水稻对温室气体排放相关功能微生物的影响,并采用稳定同位素核酸标记技术来分析种植转Bt基因水稻对土壤功能“活跃的”产甲烷古菌群落的影响,以了解该地区C转化微生物群落特征及其功能。主要结果如下：
(1)田间原位和温室盆栽实验结果均显示种植Bt水稻显著的、持久的降低了土壤CH4、N2O和CO2排放通量,对产甲烷古菌、甲烷氧化细菌、产N2O功能微生物、N2O还原功能微生物群落也有显著影响。种植Bt水稻显著的、持久的降低了土壤CH4排放通量,对根际产甲烷古菌和甲烷氧化细菌群落丰度和组成(香农多样性指数H’)也有显著的负面影响。在S2和S5阶段,Bt水稻根际产N2O功能微生物丰度显著低于Ck水稻(p<0.05),在S3阶段Bt水稻根际产N2O功能微生物丰度显著高于Ck水稻(p<0.05),而在S1和S4阶段,两种水稻根际产N2O功能微生物丰度无显著差异(p>0.05)。在S2阶段,Bt水稻根际N2O还原功能微生物丰度显著低于Ck水稻(p<0.05),在S1和S4阶段Bt水稻根际N2O还原功能微生物丰度显著高于Ck水稻(p<0.05),而在S3和S5阶段,两种水稻根际N2O还原功能微生物丰度无显著差异(p>0.05)。在苗期(S1)和分蘖期(S2),Bt水稻根际总细菌丰度显著低于Ck水稻(p<0.05)；而在晒田期(S3)、灌浆期(S4)和成熟期(S5),这种显著差异消失(p>0.05)。在S3阶段,Bt水稻根际总古菌丰度显著高于Ck水稻(p<0.05)；在其它生长阶段,Bt水稻和Ck水稻根际总古菌丰度无显著差异(p>0.05)。相关分析结果显示土壤CH4排放通量与产甲烷古菌丰度成显著正相关(R2=0.564,p<0.05)；土壤N2O排放通量与产N2O功能微生物丰度成显著正相关(R2=0.564,p<0.05)。推测：与Ck水稻相比,Bt水稻根系分泌物含量较低,产甲烷微生物、产N2O功能微生物丰度较低,苗期和分蘖期植株生物量较低,土壤氧化还原电位较高等是导致Bt水稻土壤温室气体排放通量较低的主要原因。
(2)稳定同位素核酸标记结果显示种植Bt水稻显著降低了功能“活跃的”产甲烷古菌群落丰度,但没有显著影响功能“活跃的”产甲烷古菌群落组成。Bt水稻和Ck水稻分蘖期根际土壤中功能“活跃的”产甲烷古菌类群有Rice cluster-Ⅰ (RC-Ⅰ)、Methanosaetaceae、Methanosarcinaceae和Methanobacteriaceae,其中RC-I占所有功能“活跃的”产甲烷古菌类型的60%～66.7%,为最活跃的产甲烷古菌类群。
我们还选取了亚热带地区福建万木林自然保护区典型森林土壤(细柄阿丁枫,ALG;浙江桂,CIC)及由其转变的农业利用土壤(杉木,CUL；桔园,ORG)为研究对象,采用qPCR, PCR-DGGE等现代分子生态学手段测定氨氧化古菌(ammonia-oxidizing archaea, AOA)和氨氧化细菌(ammonia-oxidizing bacteria, AOB)群落丰度和组成,研究森林土壤向农田土壤转变对土壤氨氧化微生物群落及其氨氧化功能的影响,以表征该地区N转化微生物群落特征及其功能。主要结果如下：
(1)人为利用和施肥显著提高了我国亚热带地区土壤氨氧化势。该地区森林土壤转变为农田土壤显著增加了AOA和AOB的群落丰度,且显著减小了AOA和AOB丰度比值。DGGE指纹图谱显示森林土壤转变为农田土壤也明显改变了AOA和AOB群落组成,其中隶属于Group1.la-associated lineage的AOA类群(A7～A10)在施肥利用后没有检测到,而隶属于Cluster3a lineage的AOB类群(B6)在施肥利用后也没有检测到。
(2)添加尿素培养后,天然林(ALG和CIC)土壤AOA丰度显著增加,AOB丰度无显著变化；未施肥种植林(CUL)土壤AOA和AOB丰度均显著增加;而施肥土壤(ORG) AOA和AOB丰度均无显著变化。添加尿素培养明显改变了AOA群落结构；但没有显著改变AOB群落结构。可见,天然林(ALG和CIC)土壤中AOA对添加尿素培养有明显响应；种植林土壤(CUL)和施肥土壤(ORG)中AOA和AOB对添加尿素培养有明显响应。因此,我们推断天然林(ALG和CIC)土壤中AOA对自养氨氧化过程贡献大,种植林土壤(CUL)和施肥土壤(ORG)中AOA和AOB对自养氨氧化过程均有贡献。即森林土壤向农田土壤转变改变了AOA和AOB对硝化作用的贡献,但相对贡献率尚不能确定。
The subtropical region of China, cover about21%of the country's land surface, is the main area of agricultural production in our country. Soils in this region, characterized with low pH, highly weathered, high oxidation potential, play a unique role in the world. Soil carbon and nitrogen biogeochemical cycles in this region are closely related to environmental change, agricultural production, etc. Therefore, it will be instructive to understand microbial characteristics and their functions related to carbon-nitrogen transformations for grasping the laws of carbon and nitrogen biogeochemical cycles and developing sustainable green agricultural production.
Typical rice-growing area in subtropical region (Wuhan, Hubei province, China) was selected to be the study site. Static closed chamber technique was employed to analyze the effect of planting transgenic Bt rice on greenhouse gases (GHQ such as CH4, CO2and N2O) emission from paddy soil under both field and greenhouse conditions. Modern molecular ecology methods were employed to investigate the effect of planting transgenic Bt rice on functional microorganisms related to GHGs emission. DNA-based stable isotope probing (DNA-SIP) method was employed to investigate the effect of planting transgenic Bt rice on functionally "active" methanogenic archaea. The objective of this study was to understand the microbial characteristics of carbon transformation and the functions in subtropical region. The results are as follows:
(1) Under both field and greenhouse conditions, planting transgenic Bt rice significantly and persistently reduced the CH4, N2O and CO2emission. Planting transgenic Bt rice did some significant and negative effects on community abundances and composition (characterized by Shannon-Weuner diversity index) of rhizospheric methanogenic archaea and methanotrophic bacteria. The community abundances of total bacteria in Bt rice rhizosphere at SI and S2stages were significantly lower than Ck rice rhizosphere (p<0.05), but these significant differences disappeared at following stages. The community abundance of total archaea in Bt rice rhizosphere at S3stage was significantly lower than Ck rice rhizosphere (p<0.05), but these significant differences disappeared at following stages. Regression analysis shown that the CH4emission flux significantly increased with the increasing of methanogenic archaeal abundance (R2=0.839, p<0.001), and the N2O emission flux significantly increased with the increasing of N2O production microorganism abundance (R2=0.564, p<0.05). We inferred that the lower root exudates contents, plant biomass and functional microorganism abudance and the higher soil redox potential of Bt rice were the key reasons resulting in the lower greenhouse gases emission fluxes.
(2) Results of SIP-DNA indicated that planting transgenic Bt rice significantly reduced the community abundance of functionally "active" methanogenic archaea, but did not affect the community composition. The populations of functional "active" methanogenic archaea in Bt and Ck rice rhizosphere were Rice cluster-I (RC-I), Methanosaetaceae, Methanosarcinaceae and Methanobacteriacea. RC-I is represented by the most species, accounting for60%～66.7%of total functional "active" methanogenic archaeal populations.
Typical forest soils (Altingia gralilipes, ALG; Cinnamomum chekiangense, CIC) and changed agriculture soils (Cunninghamia lanceolata, CUL; Orange orchard, ORG) in subtropical region were selected to be another study site. Modern molecular ecology methods were employed to investigate the effect of forest conversion to agriculture on ammonia oxidizers and their contributions to autotrophic ammonia oxidation process. The objective of this study was to understand the microbial characteristics of nitrogen transformation and the functions in subtropical region. The results are as follows:
(1) People use and fertilization significantly increased soil potential ammonia oxidation in this region. Forest conversion to agriculture significantly increased ammonia-oxidizing archaea (AOA) and ammonia-oxidizing bacteria (AOB) community abundances, and changed AOA and AOB community structures. The relative contributions of AOA and AOB to autotrophic ammonia oxidation process were also changed after the land-use conversion. Forest conversion to agriculture significantly changed the community compositions of AOA and AOB. The AOA populations (A7-A10), belong to Group1.la-associated lineage, could not be detected after fertilizing. The AOB population (B6), belong to Cluster3a lineage, could not be detected after fertilizing.
(2) The Urea addition significantly increased the AOA abundance in natural forests (ALG and CIC), significantly promoted the AOA and AOB abundances in unfertilized plantation soil (CUL), but had no significant effect on the AOB abundance in natural forests and the AOA and AOB abundances in fertilized soil (ORG). The Urea addition significantly changed the community compositions of AOA, but had no significant effect on the community compositions of AOB. We inferred that AOA may dominate the autotrophic ammonia oxidation process in natural forests (ALG and CIC), whereas both AOA and AOB may conduct autotrophic ammonia oxidation in agriculture soils (CUL and ORG). We supported that forest conversion to agriculture significantly changed the relative contributions of AOA and AOB to autotrophic ammonia oxidation, but the relative contribution ratio in these soils still remained unknown.
引文
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