李文娟(1996-), 女, 湖南石门县人, 硕士研究生, 主要从事土壤氮素循环过程及微生物机理研究。E-mail:
氧化亚氮(N2O)是主要温室气体之一,土壤是N2O的重要排放源,其排放主要受N2O产生和还原的功能微生物影响。土壤团聚体是由原生颗粒(砂、粉、黏粒)、胶结物质和孔隙组成的土壤基本结构单元。土壤不同粒径团聚体之间因基质和孔隙差异形成特殊独立的微生境被视为N2O的生物化学反应器。在不同的微生境中,N2O产生和还原的功能微生物分布不同,因而土壤不同粒径团聚体N2O排放可能存在差异。目前在不同生态系统土壤全土N2O排放特征的报道较多,而对于不同粒径土壤团聚体N2O排放相对贡献尚不清楚、功能微生物分布还未知、N2O产生和还原热区尚未明确。本文综述了近年来国内外关于土壤团聚体对N2O产生和排放机制的研究,总结了土壤团聚体性状特征对N2O产生和还原的影响,阐述了不同粒径土壤团聚体对N2O排放影响的微生物学机制,进一步明确了今后需加强土壤团聚体N2O产生和还原的热区、环境因子阈值范围的确定、系列功能基因(酶)整体性的研究,以期为N2O模拟排放模型优化提供参考,为土壤N2O减排提供理论依据。
Nitrous oxide (N2O), a potent greenhouse gas, is produced and reduced mainly under the mediation of functional microorganisms in soil. In terrestrial ecosystems, soil is an important source of N2O emission. Soil aggregates, a key structural component of the soil, consist of sand, silt, clay (primary particles), organic matter (binding agents) and pore spaces. According to the hierarchy theory, soil aggregates can be divided into four fractions by size, that is, large macroaggregates (>2 mm), small macroaggregates (2-0.25 mm), microaggregates (0.25-0.053 mm) and silt plus clay-sized particles (< 0.053 mm). Large macroaggregates are high in pore connectivity and oxygen diffusion rate, fast in turnover, and rich in organic matter, and microaggregates high in water retention capacity and stable carbon content, and capable of protecting microorganisms from being predated. Hence, soil aggregates different in size may offer heterogeneous microhabitats for fungi and bacteria. And each independent microhabitat could be regarded as a biogeochemical reactor producing greenhouse gas. Nitrifiers and denitrifiers, which carry functional genes
氧化亚氮(Nitrous oxide,N2O)是大气中重要的温室气体,其单位质量的增温潜势是二氧化碳(CO2)的265倍[
土壤团聚体由原生颗粒(砂、粉、黏粒)、胶结物质和孔隙组成[
土壤团聚体是矿物颗粒在植物根系和土壤有机质、菌丝、土壤氧化物等有机和无机胶结物质作用下结合形成的二次颗粒。根据分级团聚理论,将土壤团聚体分成大团聚体(Large macroaggregates, > 2 mm)、小团聚体(Small macroaggregates,2~ 0.25 mm)、微团聚体(Microaggregates,0.25~ 0.053 mm)和粉-黏颗粒(Silt-plus Clay-size Particles, < 0.053 mm)[
土壤团聚体结构特征和周转过程概念图
A conception map of structural characteristics and turnover processes of soil aggregates
不同粒径土壤团聚体N2O排放速率
N2O emission rate in different sizes of soil aggregates
筛分粒径 |
团聚体分级 |
培养时间 |
N2O排放量 |
排放速率 |
文献 |
< 2 mm | 小团聚体 | 9 d | 12 mg·kg-1·9 d-1 | 56 | [ |
4.5 mm | 大团聚体 | 40.7 mg·kg-1·9 d-1 | 188.4 | ||
< 1 mm | 小团聚体 | 96 h | 7.637 μg·g-1·h-1 | 7637 | [ |
2~4 mm | 大团聚体 | 5.607 μg·g-1·h-1 | 5607 | ||
2~8 mm | 大团聚体 | 21 d | 23.5 mg·kg-1 | 46.6 | [ |
0.25~0.5 mm | 小团聚体 | 96 h | 0.04 mg·kg-1 | 0.42 | [ |
2~4 mm | 大团聚体 | — | 7.0 mg·kg-1 | 72.9 | |
< 0.25 mm | 微团聚体 | — | 4.03 μg·kg-1·h-1 | 4.03 | [ |
2~0.25 mm | 小团聚体 | — | 0.18 μg·kg-1·h-1 | 0.18 | |
2~6 mm | 大团聚体 | — | 0.96 μg·kg-1·h-1 | 0.96 | |
< 0.053 mm | 粉-黏颗粒 | 28 d | 14.3 μg·kg-1 | 0.021 | [ |
0.25~0.053 mm | 微团聚体 | — | 6.2 μg·kg-1 | 0.009 | |
2~0.25 mm | 小团聚体 | — | 10.47 μg·kg-1 | 0.016 | |
> 2 mm | 大团聚体 | — | 29.83 μg·kg-1 | 0.044 | |
1~2 mm | 小团聚体 | — | 1.2 ng·g-1·h-1 | 1.2 | [ |
2~4 mm | 大团聚体 | — | 1.8 ng·g-1·h-1 | 1.8 |
大量研究发现大团聚体较微团聚体具有更高的N2O排放能力[
土壤团聚体N2O排放主要与N2O产生和还原有关。土壤N2O主要通过微生物的硝化作用(Nitrification)、反硝化作用(Denitrification)以及硝酸盐异化还原成铵(Dissimilatory nitrate reduction to ammonium,DNRA)、硝化-反硝化耦合作用(Nitrification-Coupled denitrification)、共反硝化作用(Co-denitrification)和非生物分解等过程产生[
土壤N2O产生过程及其微生物机制
Biotic processes of nitrous oxide(N2O)production in soil
微生物硝化作用主要是将NH3氧化为
反硝化作用是微生物将
土壤N2O的产生还存在其他微生物过程,如:硝酸盐异化还原成铵、硝化-反硝化耦合作用和共反硝化等[
土壤N2O可以通过多个微生物途径产生,但N2O还原目前已知的过程仅为
土壤团聚体形成与周转过程中,微生物可以通过代谢物黏结周围土壤颗粒,形成对其有利的生境[
Tisdall和Oades[
微生物群落和结构受土壤团聚体粒径的影响,细菌和真菌群落在不同粒径团聚体中存在明显结构差异。细菌和古菌更倾向存活于微团聚体,一方面能躲避土壤动物的捕食;另一方面作为原核生物的细菌和古菌较真核生物的真菌小,更易于在微小生境栖息[
土壤氮素循环功能微生物在不同粒径团聚体的分布情况如
环境变化影响土壤团聚体功能微生物群落重新分布,进而影响N2O排放[
氮循环功能基因可作为一项指标来衡量N2O排放。目前已有研究表明,土壤N2O产生潜力能用
不同粒径土壤团聚体中微生物群落分布不同,携带的功能基因也存在一定差异(
不同粒径团聚体N2O排放速率及功能基因分布
Distribution of functional genes and N2O emission rate in different sizes of soil aggregates
团聚体粒径 |
N2O排放速率N2O flux/(μg·kg-1·h-1)[ |
主要功能微生物 |
主要功能基因[ |
< 0.25 mm | 0.044~5607 | 硝化细菌 |
|
0.25~2 mm | 0.016~7637 | ||
> 2 mm | 0.009~4.03 | 氨氧化菌AOA |
但是Liu等[
土壤团聚体是微生物进行硝化/反硝化作用的生化反应器。不同粒径土壤团聚体中孔隙特征、通气条件、底物浓度以及水分含量等性状分异能改变微生物群落结构和活性,从而导致不同粒径团聚体中N2O排放差异较大。目前功能微生物对N2O排放的研究重心逐渐从全土尺度转移到团聚体尺度,聚焦于微生物群落结构和多样性以及功能微生物与团聚体的生物物理关联机制下N2O排放研究。
今后的研究主要加强以下几个方面:
1)土壤团聚体N2O产生和还原热区探究。土壤团聚体内部和周围孔隙是微生物存活的生境,有机质、氧气浓度、水分条件和底物成分等差异可能是导致微生物群落差异的主要驱动因素。土壤大团聚体通气条件好,可利用有机碳含量高,但是容易被外界环境因素扰动,而微团聚体内部可能是微生物相对稳定和隐蔽的栖息地。微团聚体具有长期稳定的碳库,能够持续为硝化/反硝化微生物提供生长环境,并且可能为微生物提供物理保护。近年来大量研究表明微团聚体内存在较高的微生物丰度,氮循环功能基因也在微团聚体中大量分布。但是目前对土壤团聚体中氮循环功能基因分布特征研究较少,尚不能明确功能微生物对土壤微团聚体粒径的偏好。今后还需要结合室内控制试验和环境因子梯度,明确N2O产生和还原的潜在热区,进一步探究微团聚体和大团聚体对N2O排放的贡献。
2)环境因子阈值范围确定。大团聚体和微团聚体均可能为硝化/反硝化微生物提供有利的生境,不同粒径土壤团聚体N2O排放潜势存在差异,可能是由底物浓度、通气条件以及水分含量等异质性引起。一旦外界环境压力超出阈值范围,则对参与硝化/反硝化过程的微生物生存产生胁迫,从而改变硝化/反硝化过程中N2O排放。通过研究确定环境因子的临界范围,可进一步明晰不同粒径土壤团聚体N2O产生和还原所占的比例,可初步判断N2O排放情况,有利于更好地分析硝化/反硝化对N2O的贡献率以及微生物驱动机制。
3)对系列功能基因(酶)开展整体性研究,而不仅是单个主要基因(酶)。硝化/反硝化过程能够在微生物体内发生,并排放N2O,是一系列功能基因和酶的运转,而非某一个功能基因和酶发挥作用。例如:目前用
Pachauri R K, Meyer L A, Barros V R, et al. Climate change 2014: Synthesis report[M]//Contribution of working groups I, II and III to the fifth assessment report of the intergovernmental panel on climate change. Geneva, Switzerland: Intergovernmental Panel on Climate Change(IPCC), 2014.
Ravishankara A R, Daniel J S, Portmann R W. Nitrous oxide(N2O): The dominant ozone-depleting substance emitted in the 21st century[J]. Science, 2009, 326(5949): 123-125.
Solomon S, Qin D, Manning M, et al. Climate change 2007: The physical science basis[M]//Contribution of working group I to the fourth assessment report of the intergovernmental panel on climate change. Summary for policymakers. Geneva: Intergovernmental Panel on Climate Change(IPCC), 2007.
Stein L Y. The long-term relationship between microbial metabolism and greenhouse gases[J]. Trends in Microbiology, 2020, 28(6): 500-511.
Bosilj P, Gould I, Duckett T, et al. Estimating soil aggregate size distribution from images using pattern spectra[J]. Biosystems Engineering, 2020, 198: 63-77.
Li F Q, Xue C, Qiu P F, et al. Soil aggregate size mediates the responses of microbial communities to crop rotation[J]. European Journal of Soil Biology, 2018, 88: 48-56.
Six J, Bossuyt H, Degryze S, et al. A history of research on the link between(micro)aggregates, soil biota, and soil organic matter dynamics[J]. Soil & Tillage Research, 2004, 79(1): 7-31.
Diba F, Shimizu M, Hatano R. Effects of soil aggregate size, moisture content and fertilizer management on nitrous oxide production in a volcanic ash soil[J]. Soil Science and Plant Nutrition, 2011, 57(5): 733-747.
Blaud A, van der Zaan B, Menon M, et al. The abundance of nitrogen cycle genes and potential greenhouse gas fluxes depends on land use type and little on soil aggregate size[J]. Applied Soil Ecology, 2018, 125: 1-11.
Zhou H C, Zhang W Z, Liu Y, et al. Relationships of N2O emission with abundance and composition of denitrifying microorganisms in soil aggregates[J]. Acta Pedologica Sinica, 2015, 52(5): 1144-1152.
周汉昌, 张文钊, 刘毅, 等. 土壤团聚体N2O释放与反硝化微生物丰度和组成的关系[J]. 土壤学报, 2015, 52(5): 1144-1152.
Cambardella C A, Elliott E T. Carbon and nitrogen distribution in aggregates from cultivated and native grassland soils[J]. Soil Science Society of America Journal, 1993, 57(4): 1071-1076.
Ananyeva K, Wang W, Smucker A J M, et al. Can intra-aggregate pore structures affect the aggregate's effectiveness in protecting carbon?[J]. Soil Biology & Biochemistry, 2013, 57: 868-875.
Rillig M C, Muller L A, Lehmann A. Soil aggregates as massively concurrent evolutionary incubators[J]. The ISME Journal, 2017, 11(9): 1943-1948.
Wang B, Brewer P E, Shugart H H, et al. Soil aggregates as biogeochemical reactors and implications for soil-atmosphere exchange of greenhouse gases—A concept[J]. Global Change Biology, 2019, 25(2): 373-385.
Khalil K, Renault P, Mary B. Effects of transient anaerobic conditions in the presence of acetylene on subsequent aerobic respiration and N2O emission by soil aggregates[J]. Soil Biology & Biochemistry, 2005, 37(7): 1333-1342.
Kimura S D, Melling L, Goh K J. Influence of soil aggregate size on greenhouse gas emission and uptake rate from tropical peat soil in forest and different oil palm development years[J]. Geoderma, 2012, 185/186: 1-5.
Drury C F, Yang X M, Reynolds W D, et al. Influence of crop rotation and aggregate size on carbon dioxide production and denitrification[J]. Soil & Tillage Research, 2004, 79(1): 87-100.
Muñoz C, Torres P, Alvear M, et al. Physical protection of C and greenhouse gas emissions provided by soil macroaggregates from a Chilean cultivated volcanic soil[J]. Acta Agriculturae Scandinavica, Section B — Soil & Plant Science, 2012, 62(8): 739-748.
Uchida Y, Clough T, Kelliher F, et al. Effects of aggregate size, soil compaction, and bovine urine on N2O emissions from a pasture soil[J]. Soil Biology & Biochemistry, 2008, 40(4): 924-931.
Robinson A, Di H J, Cameron K C, et al. Effect of soil aggregate size and dicyandiamide on N2O emissions and ammonia oxidizer abundance in a grazed pasture soil[J]. Soil Use and Management, 2014, 30(2): 231-240.
Sey B K, Manceur A M, Whalen J K, et al. Small-scale heterogeneity in carbon dioxide, nitrous oxide and methane production from aggregates of a cultivated sandy-loam soil[J]. Soil Biology & Biochemistry, 2008, 40(9): 2468-2473.
Bandyopadhyay K K, Lal R. Effect of land use management on greenhouse gas emissions from water stable aggregates[J]. Geoderma, 2014, 232/233/234: 363-372.
Mangalassery S, Sjögersten S, Sparkes D L, et al. The effect of soil aggregate size on pore structure and its consequence on emission of greenhouse gases[J]. Soil & Tillage Research, 2013, 132: 39-46.
Butterbach-Bahl K, Baggs E M, Dannenmann M, et al. Nitrous oxide emissions from soils: How well do we understand the processes and their controls?[J]. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 2013, 368(1621): 20130122.
Hallin S, Philippot L, Löffler F E, et al. Genomics and ecology of novel N2O-reducing microorganisms[J]. Trends in Microbiology, 2018, 26(1): 43-55.
Kuypers M M M, Marchant H K, Kartal B. The microbial nitrogen-cycling network[J]. Nature Reviews Microbiology, 2018, 16(5): 263-276.
Prosser J I, Hink L, Gubry-Rangin C, et al. Nitrous oxide production by ammonia oxidizers: Physiological diversity, niche differentiation and potential mitigation strategies[J]. Global Change Biology, 2020, 26(1): 103-118.
Cao W C, Song H, Wang Y J, et al. Key production processes and influencing factors of nitrous oxide emissions from agricultural soils[J]. Journal of Plant Nutrition and Fertilizers, 2019, 25(10): 1781-1798.
曹文超, 宋贺, 王娅静, 等. 农田土壤N2O排放的关键过程及影响因素[J]. 植物营养与肥料学报, 2019, 25(10): 1781-1798.
Liu N, Yan Z S, Fan Y J, et al. Effect of different nitrogen application levels on the content of soluble protein and key enzyme activities in nitrogen metabolism of sugar beet[J]. Chinese Agricultural Science Bulletin, 2015, 31(30): 149-154.
刘娜, 闫志山, 范有君, 等. 不同氮素水平对甜菜氮代谢酶和可溶性蛋白含量的影响[J]. 中国农学通报, 2015, 31(30): 149-154.
Daims H, Lebedeva E V, Pjevac P, et al. Complete nitrification by
Kits K D, Jung M Y, Vierheilig J, et al. Low yield and abiotic origin of N2O formed by the complete nitrifier
Opperman D J, Murgida D H, Dalosto S D, et al. A three-domain copper-nitrite reductase with a unique sensing loop[J]. IUCrJ, 2019, 6(2): 248-258.
Priemé A, Braker G, Tiedje J M. Diversity of nitrite reductase(
Zumft W G. Nitric oxide reductases of prokaryotes with emphasis on the respiratory, heme-copper oxidase type[J]. Journal of Inorganic Biochemistry, 2005, 99(1): 194-215.
Blomberg M R A, Ädelroth P. Mechanisms for enzymatic reduction of nitric oxide to nitrous oxide-A comparison between nitric oxide reductase and cytochrome c oxidase[J]. Biochimica et Biophysica Acta-Bioenergetics, 2018, 1859(11): 1223-1234.
Suharti, Strampraad M J F, Schröder I, et al. A novel copper a containing menaquinol NO reductase from
Putz M, Schleusner P, Rütting T, et al. Relative abundance of denitrifying and DNRA bacteria and their activity determine nitrogen retention or loss in agricultural soil[J]. Soil Biology & Biochemistry, 2018, 123: 97-104.
Simon J. Enzymology and bioenergetics of respiratory nitrite ammonification[J]. FEMS Microbiology Reviews, 2002, 26(3): 285-309.
He J Z, Zhang L M. Key processes and microbial mechanisms of soil nitrogen transformation[J]. Microbiology China, 2013, 40(1): 98-108.
贺纪正, 张丽梅. 土壤氮素转化的关键微生物过程及机制[J]. 微生物学通报, 2013, 40(1): 98-108
Rütting T, Boeckx P, Müller C, et al. Assessment of the importance of dissimilatory nitrate reduction to ammonium for the terrestrial nitrogen cycle[J]. Biogeosciences, 2011, 8(7): 1779-1791.
Tanimoto T, Hatano K I, Kim D H, et al. Co-denitrification by the denitrifying system of the fungus
Graf D R H, Jones C M, Hallin S. Intergenomic comparisons highlight modularity of the denitrification pathway and underpin the importance of community structure for N2O emissions[J]. PLoS One, 2014, 9(12): e114118.
Sanford R A, Wagner D D, Wu Q Z, et al. Unexpected nondenitrifier nitrous oxide reductase gene diversity and abundance in soils[J]. Proceedings of the National Academy of Sciences of the United States of America, 2012, 109(48): 19709-19714.
Baggs E M. Soil microbial sources of nitrous oxide: Recent advances in knowledge, emerging challenges and future direction[J]. Current Opinion in Environmental Sustainability, 2011, 3(5): 321-327.
Pauleta S R, Carepo M S P, Moura I. Source and reduction of nitrous oxide[J]. Coordination Chemistry Reviews, 2019, 387: 436-449.
Li F Q, Qiu P F, Shen B, et al. Soil aggregate size modifies the impacts of fertilization on microbial communities[J]. Geoderma, 2019, 343: 205-214.
Dini-Andreote F, Stegen J C, van Elsas J D, et al. Disentangling mechanisms that mediate the balance between stochastic and deterministic processes in microbial succession[J]. Proceedings of the National Academy of Sciences of the United States of America, 2015, 112(11): E1326-E1332.
Rittershaus E S C, Baek S H, Sassetti C M. The normalcy of dormancy: Common themes in microbial quiescence[J]. Cell Host & Microbe, 2013, 13(6): 643-651.
Rillig M C, Lehmann A, Aguilar-Trigueros C A, et al. Soil microbes and community coalescence[J]. Pedobiologia, 2016, 59(1/2): 37-40.
Tisdall J M, Oades J M. Organic matter and water-stable aggregates in soils[J]. Journal of Soil Science, 1982, 33(2): 141-163.
Blagodatsky S, Smith P. Soil physics meets soil biology: Towards better mechanistic prediction of greenhouse gas emissions from soil[J]. Soil Biology & Biochemistry, 2012, 47: 78-92.
Xiao J J, Xing D, Mao M M, et al. Mechanism of arbuscular mycorrhizal fungal affecting soil aggregates in rhizosphere of mulberry(
肖玖军, 邢丹, 毛明明, 等. AM真菌对桑树根围土壤团聚体的影响机制[J]. 土壤学报, 2020, 57(3): 773-782.
Lehmann A, Zheng W S, Rillig M C. Soil biota contributions to soil aggregation[J]. Nature Ecology & Evolution, 2017, 1(12): 1828-1835.
Li N, Han X Z, You M Y, et al. Research review on soil aggregates and microbes[J]. Ecology and Environment, 2013, 22(9): 1625-1632.
李娜, 韩晓增, 尤孟阳, 等. 土壤团聚体与微生物相互作用研究[J]. 生态环境学报, 2013, 22(9): 1625-1632.
Tian X L, Wang C B, Bao X G, et al. Crop diversity facilitates soil aggregation in relation to soil microbial community composition driven by intercropping[J]. Plant and Soil, 2019, 436(1/2): 173-192.
Okiobe S T, Augustin J, Mansour I, et al. Disentangling direct and indirect effects of mycorrhiza on nitrous oxide activity and denitrification[J]. Soil Biology & Biochemistry, 2019, 134: 142-151.
Ran Y G, Ma M H, Liu Y, et al. Physicochemical determinants in stabilizing soil aggregates along a hydrological stress gradient on reservoir riparian habitats: Implications to soil restoration[J]. Ecological Engineering, 2020, 143: 105664.
Six J, Paustian K, Elliott E T, et al. Soil structure and organic matter I. Distribution of aggregate-size classes and aggregate-associated carbon[J]. Soil Science Society of America Journal, 2000, 64(2): 681-689.
Wright D A, Killham K, Glover L A, et al. Role of pore size location in determining bacterial activity during predation by protozoa in soil[J]. Applied and Environmental Microbiology, 1995, 61(10): 3537-3543.
Mummey D, Holben W, Six J, et al. Spatial stratification of soil bacterial populations in aggregates of diverse soils[J]. Microbial Ecology, 2006, 51(3): 404-411.
Yang C, Liu N, Zhang Y J. Soil aggregates regulate the impact of soil bacterial and fungal communities on soil respiration[J]. Geoderma, 2019, 337: 444-452.
Frey S D. Aggregation-microbial aspects[M]//Hillel D. Encyclopedia of soils in the environment. New York: Academic Press, 2005: 22-28.
Liao H, Zhang Y C, Zuo Q Y, et al. Contrasting responses of bacterial and fungal communities to aggregate-size fractions and long-term fertilizations in soils of northeastern China[J]. Science of the Total Environment, 2018, 635: 784-792.
Mothapo N, Chen H H, Cubeta M A, et al. Phylogenetic, taxonomic and functional diversity of fungal denitrifiers and associated N2O production efficacy[J]. Soil Biology & Biochemistry, 2015, 83: 160-175.
Poll C, Thiede A, Wermbter N, et al. Micro-scale distribution of microorganisms and microbial enzyme activities in a soil with long-term organic amendment[J]. European Journal of Soil Science, 2003, 54(4): 715-724.
Bach E M, Williams R J, Hargreaves S K, et al. Greatest soil microbial diversity found in micro-habitats[J]. Soil Biology & Biochemistry, 2018, 118: 217-226.
Chotte J L, Schwartzmann A, Bally R, et al. Changes in bacterial communities and
Nahidan S, Nourbakhsh F, Henneberger R, et al. Aggregate size distribution of ammonia-oxidizing bacteria and archaea at different landscape positions[J]. Geomicrobiology Journal, 2017, 34(10): 895-902.
Jiang Y J, Jin C, Sun B. Soil aggregate stratification of nematodes and ammonia oxidizers affects nitrification in an acid soil[J]. Environmental Microbiology, 2014, 16(10): 3083-3094.
Li P P, Han Y L, He J Z, et al. Soil aggregate size and long-term fertilization effects on the function and community of ammonia oxidizers[J]. Geoderma, 2019, 338: 107-117.
Lensi R, Clays-Josserand A, Jocteur Monrozier L. Denitrifiers and denitrifying activity in size fractions of a mollisol under permanent pasture and continuous cultivation[J]. Soil Biology & Biochemistry, 1995, 27(1): 61-69.
Kong A Y Y, Hristova K, Scow K M, et al. Impacts of different N management regimes on nitrifier and denitrifier communities and N cycling in soil microenvironments[J]. Soil Biology & Biochemistry, 2010, 42(9): 1523-1533.
Zhou J, Ning D. Stochastic community assembly: Does it matter in microbial ecology?[J]. Microbiology and Molecular Biology Reviews, 2017, 81(4): e00002-00017.
Rabbi S M F, Daniel H, Lockwood P V, et al. Physical soil architectural traits are functionally linked to carbon decomposition and bacterial diversity[J]. Scientific Reports, 2016, 6: 33012.
Xiao S S, Zhang W, Ye Y Y, et al. Soil aggregate mediates the impacts of land uses on organic carbon, total nitrogen, and microbial activity in a Karst ecosystem[J]. Scientific Reports, 2017, 7: 41402.
Aamer M, Shaaban M, Hassan M U, et al. Biochar mitigates the N2O emissions from acidic soil by increasing the
Huang R, Wang Y Y, Liu J, et al. Variation in N2O emission and N2O related microbial functional genes in straw- and biochar-amended and non-amended soils[J]. Applied Soil Ecology, 2019, 137: 57-68.
Mathieu O, Lévêque J, Hénault C, et al. Emissions and spatial variability of N2O, N2 and nitrous oxide mole fraction at the field scale, revealed with 15N isotopic techniques[J]. Soil Biology & Biochemistry, 2006, 38(5): 941-951.
Weier K, Doran J W, Power J F, et al. Denitrification and the dinitrogen/nitrous oxide ratio as affected by soil water, available carbon, and nitrate[J]. Soil Science Society of America Journal, 1993, 57(1): 66-72.
Ying J Y, Li X X, Wang N N, et al. Contrasting effects of nitrogen forms and soil pH on ammonia oxidizing microorganisms and their responses to long-term nitrogen fertilization in a typical steppe ecosystem[J]. Soil Biology & Biochemistry, 2017, 107: 10-18.
Senbayram M, Budai A, Bol R, et al. Soil NO3- level and O2 availability are key factors in controlling N2O reduction to N2 following long-term liming of an acidic sandy soil[J]. Soil Biology & Biochemistry, 2019, 132: 165-173.
Trivedi P, Rochester I J, Trivedi C, et al. Soil aggregate size mediates the impacts of cropping regimes on soil carbon and microbial communities[J]. Soil Biology & Biochemistry, 2015, 91: 169-181.
Bateman E J, Baggs E. Contributions of nitrification and denitrification to N2O emissions from soils at different water-filled pore space[J]. Biology and Fertility of Soils, 2005, 41(6): 379-388.
Ruser R, Flessa H, Russow R, et al. Emission of N2O, N2 and CO2 from soil fertilized with nitrate: Effect of compaction, soil moisture and rewetting[J]. Soil Biology & Biochemistry, 2006, 38(2): 263-274.
Jia W L, Liang S, Zhang J, et al. Nitrous oxide emission in low-oxygen simultaneous nitrification and denitrification process: Sources and mechanisms[J]. Bioresource Technology, 2013, 136: 444-451.
Zhu X, Burger M, Doane T A, et al. Ammonia oxidation pathways and nitrifier denitrification are significant sources of N2O and NO under low oxygen availability[J]. Proceedings of the National Academy of Sciences of the United States of America, 2013, 110(16): 6328-6333.
An S S, Mentler A, Mayer H, et al. Soil aggregation, aggregate stability, organic carbon and nitrogen in different soil aggregate fractions under forest and shrub vegetation on the Loess Plateau, China[J]. Catena, 2010, 81(3): 226-233.
Zhu K, Ma M H, Ran Y G, et al. In mitigating CO2 emission in the reservoir riparian: The influences of land use and the dam-triggered flooding on soil respiration[J]. Soil & Tillage Research, 2020, 197: 104522.
Vilain G, Garnier J, Tallec G, et al. Effect of slope position and land use on nitrous oxide(N2O)emissions(Seine Basin, France)[J]. Agricultural and Forest Meteorology, 2010, 150(9): 1192-1202.
Shi H A, Li L J, You M Y, et al. Impact of soil temperature and moisture on soil N2O emission from mollisols under different land-use types[J]. Journal of Agro-Environment Science, 2013, 32(11): 2286-2292.
石洪艾, 李禄军, 尤孟阳, 等. 不同土地利用方式下土壤温度与土壤水分对黑土N2O排放的影响[J]. 农业环境科学学报, 2013, 32(11): 2286-2292.
Li M, Qing J, Hong Y, et al. Effects of nitrogen addition on ecological stoichiometric characteristics of carbon, nitrogen and phosphorus in
李明, 秦洁, 红雨, 等. 氮素添加对贝加尔针茅草原土壤团聚体碳、氮和磷生态化学计量学特征的影响[J]. 草业学报, 2019, 28(12): 29-40.
Wei W, Isobe K, Shiratori Y, et al. N2O emission from cropland field soil through fungal denitrification after surface applications of organic fertilizer[J]. Soil Biology & Biochemistry, 2014, 69: 157-167.
Zhong L, Du R, Ding K, et al. Effects of grazing on N2O production potential and abundance of nitrifying and denitrifying microbial communities in meadow-steppe grassland in Northern China[J]. Soil Biology & Biochemistry, 2014, 69: 1-10.
Wang L, Li K, Song YQ, et al. The N2O consumption ability in the surface paddy soil layer and its coupling relationship to N2O reducing microorganisms[J]. Acta Ecologica Sinica, 2019, 39(20): 7602-7610.
王玲, 李昆, 宋雅琦, 等. 浅表层水稻土N2O消耗能力及其与N2O还原微生物的耦合关系[J]. 生态学报, 2019, 39(20): 7602-7610.
Rasche F, Knapp D, Kaiser C, et al. Seasonality and resource availability control bacterial and archaeal communities in soils of a temperate beech forest[J]. The ISME Journal, 2011, 5(3): 389-402.
Levy-Booth D J, Prescott C E, Grayston S J. Microbial functional genes involved in nitrogen fixation, nitrification and denitrification in forest ecosystems[J]. Soil Biology & Biochemistry, 2014, 75: 11-25.
Li S S, Chen C, Duan P P, et al. Effects of biochar application on N2O emissions and abundance of nitrogen related functional genes in an acidic vegetable soil[J]. Journal of Plant Nutrition and Fertilizers, 2018, 24(2): 414-423.
李双双, 陈晨, 段鹏鹏, 等. 生物质炭对酸性菜地土壤N2O排放及相关功能基因丰度的影响[J]. 植物营养与肥料学报, 2018, 24(2): 414-423.
Xin X, Liu Q, Liu W, et al. Distribution of nitrifiers and nitrification associated with different sizes of aggregates along a 2000 year chronosequence of rice cultivation[J]. Catena, 2014, 119: 71-77.
Liu X, Chen C R, Wang W J, et al. Soil environmental factors rather than denitrification gene abundance control N2O fluxes in a wet sclerophyll forest with different burning frequency[J]. Soil Biology & Biochemistry, 2013, 57: 292-300.
Wallenstein M D, Myrold D D, Firestone M, et al. Environmental controls on denitrifying communities and denitrification rates: Insights from molecular methods[J]. Ecological Applications, 2006, 16(6): 2143-2152.