Studies on the emission of nitrous oxide from terrestrial plants

doi:10.17521/cjpe.2024.0024

Abstract

Abstract:

As a greenhouse gas with a strong warming effect, nitrous oxide (N₂O) is also one of the main substances which destroy the atmospheric ozone layer, with a strong feedback effect on climate change. Plants are another important source of N₂O emissions in terrestrial ecosystems besides soil, and related studies have received extensive attention in recent years. Based on the existing research results, the related research methods, mechanisms and their influencing factors on N₂O emissions from terrestrial plants are summarized in this study. Existing studies have mainly determined the changes in N₂O emission from plants by in situ and in vitro methods. The potential mechanisms for N₂O emission from terrestrial plants are as follows: 1) the plant produces N₂O during nitrogen metabolism; 2) microbial activities in or on the surface of the plant produce N₂O; and 3) the plant acts as a soil N₂O channel during the gas exchange process. The N₂O emissions from plant are related to internal and external factors. The internal factors include plant species, different organs of the same plant and different developmental periods, etc.; external factors include light, temperature, water, nutrients and microorganisms. However, there is still a lack of in-depth analysis on the specific mechanisms. Therefore, combining metagenomic technology to determine plant microbial genomes under different soil environmental conditions, and analyzing their microbial community structure were applied to elucidate the microbial mechanisms underlying the flux differences of N₂O emissions from plants in different habitats. The metatranscriptome technology was used to determine total RNA in plant tissues and analyze the expression abundance changes of N₂O emission related genes under different environmental conditions. And isotope labeling technology was further used to track the nitrogen metabolism process of plants and reveal the mechanism of N₂O flux changes in plants. These results are of great significance for supplementing and improving N₂O prediction models such as “Ecosys” for global climate change, which provide theoretical reference for the formulation of greenhouse gas emission reduction measures.

Key words: greenhouse gases, plants N₂O flux, emission mechanisms, influencing factors, research method

JIANG Xiao-Yu, YU Xin-Miao, LIAO Qin, ZHANG Jin-Wei, WU Xue-Feng, WANG Xu, PAN Jun-Tong, WANG Jun-Feng, MU Chun-Sheng, SHI Yu-Jie. Studies on the emission of nitrous oxide from terrestrial plants[J]. Chin J Plant Ecol, 2025, 49(4): 513-525.

Add to citation manager EndNote|Ris|BibTeX

URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2024.0024

https://www.plant-ecology.com/EN/Y2025/V49/I4/513

Figures/Tables 2

References 124

[1]	Alegria AE, Sanchez S, Quintana I (2004). Quinone-enhanced ascorbate reduction of nitric oxide: role of quinone redox potential. Free Radical Research, 38, 1107-1112. PMID
[2]	Amundson RG, Davidson EA (1990). Carbon dioxide and nitrogenous gases in the soil atmosphere. Journal of Geochemical Exploration, 38, 13-41.
[3]	Ascenzi P, Marino M, Polticelli F, Santucci R, Coletta M (2014). Cardiolipin modulates allosterically the Nitrite reductase activity of horse heart cytochrome c. Journal of Biological Inorganic Chemistry, 19, 1195-1201. DOI PMID
[4]	Aslam M, Huffaker RC, Rains DW, Rao KP (1979). Influence of light and ambient carbon dioxide concentration on nitrate assimilation by intact barley seedlings. Plant Physiology, 63, 1205-1209. DOI PMID
[5]	Baggs EM (2011). Soil microbial sources of nitrous oxide: recent advances in knowledge, emerging challenges and future direction. Current Opinion in Environmental Sustainability, 3, 321-327.
[6]	Balogh E, Kalapos B, Ahres M, Boldizsár Á, Gierczik K, Gulyás Z, Gyugos M, Szalai G, Novák A, Kocsy G (2022). Far-red light coordinates the diurnal changes in the transcripts related to nitrate reduction, glutathione metabolism and antioxidant enzymes in barley. International Journal of Molecular Sciences, 23, 7479. DOI: 10.3390/ijms23137479.
[7]	Benamar A, Rolletschek H, Borisjuk L, Avelange-Macherel MH, Curien G, Mostefai HA, Andriantsitohaina R, Macherel D (2008). Nitrite-nitric oxide control of mitochondrial respiration at the frontier of Anoxia. Biochimica et Biophysica Acta-Bioenergetics, 1777, 1268-1275.
[8]	Blomberg MRA, Ädelroth P (2018). Mechanisms for enzymatic reduction of nitric oxide to nitrous oxide—A comparison between nitric oxide reductase and cytochrome c oxidase. Biochimica et Biophysica Acta-Bioenergetics, 1859, 1223-1234.
[9]	Bowatte S, Newton PCD, Brock S, Theobald P, Luo D (2015). Bacteria on leaves: a previously unrecognised source of N₂O in grazed pastures. The ISME Journal, 9, 265-267.
[10]	Bowatte S, Newton PCD, Theobald P, Brock S, Hunt C, Lieffering M, Sevier S, Gebbie S, Luo D (2014). Emissions of nitrous oxide from the leaves of grasses. Plant Soil, 374, 275-283.
[11]	Brudvig GW, Stevens TH, Chan SI (1980). Reactions of Nitric oxide with cytochrome c oxidase. Biochemistry, 19, 5275-5285. PMID
[12]	Butterbach-Bahl K, Baggs EM, Dannenmann M, Kiese R, Zechmeister-Boltenstern S (2013). Nitrous oxide emissions from soils: How well do we understand the processes and their controls? Philosophical Transactions of the Royal Society B: Biological Sciences, 368, 20130122. DOI: 10.1098/rstb.2013.0122.
[13]	Casella S, Leporini C, Nuti M (1984). Nitrous oxide production by nitrogen-fixing, fast-growing rhizobia. Microbial Ecology, 10, 107-114. DOI PMID
[14]	Castello PR, David PS, McClure T, Crook Z, Poyton RO (2006). Mitochondrial cytochrome oxidase produces nitric oxide under hypoxic conditions: implications for oxygen sensing and hypoxic signaling in eukaryotes. Cell Metabolism, 3, 277-287. DOI PMID
[15]	Chamizo-Ampudia A, Sanz-Luque E, Llamas A, Galvan A, Fernandez E (2017). Nitrate reductase regulates plant nitric oxide homeostasis. Trends in Plant Science, 22, 163-174. DOI PMID
[16]	Chang C, Janzen HH, Nakonechny EM, Cho CM (1998). Nitrous oxide emission through plants. Soil Science Society of America Journal, 62, 35-38.
[17]	Chen GX (1990). Research on N₂O emissions from plants. Journal of Applied Ecology, 4, 295-298.
	[陈冠雄 (1990). 植物释放N₂O的研究. 应用生态学报, 4, 295-298.]
[18]	Chen X, Boeckx P, Shen S, van Cleemput O (1999). Emission of N₂O from rye grass (Lolium perenne L.). Biology and Fertility of Soils, 28, 393-396.
[19]	Chen X, Shen SM, Zhang L, Wu J, Wang XQ (1995). Preliminary research on the effect of nitrogen and phosphorus supply on N₂O emission by crops. Journal of Applied Ecology, 6, 104-105.
	[陈欣, 沈善敏, 张璐, 吴杰, 王学强 (1995). N、P供给对作物排放N₂O的影响研究初报. 应用生态学报, 6, 104-105.]
[20]	Cho CM, Sakdinan L, Chang C (2010). Denitrification intensity and capacity of three irrigated alberta soils. Soil Science Society of America Journal, 43, 945-950.
[21]	Sparacino-Watkins CE, Tejero J, Sun B, Gauthier MC, Thomas J, Ragireddy V, Merchant BA, Wang J, Azarov I, Basu P, Gladwin MT (2014). Nitrite reductase and nitric-oxide synthase activity of the mitochondrial molybdopterin enzymes mARC₁ and mARC₂. The Journal of Biological Chemistry, 289, 10345-10358.
[22]	Cuhel J, Simek M, Laughlin RJ, Bru D, Chèneby D, Watson CJ, Philippot L (2010). Insights into the effect of soil pH on N₍₂₎O and N₍₂₎ emissions and denitrifier community size and activity. Applied and Environmental Microbiology, 76, 1870-1878. DOI PMID
[23]	Dai Z, Yu M, Chen H, Zhao H, Huang Y, Su W, Xia F, Chang SX, Brookes PC, Dahlgren RA, Xu JM (2020). Elevated temperature shifts soil N cycling from microbial immobilization to enhanced mineralization, nitrification and denitrification across global terrestrial ecosystems. Global Change Biology, 26, 5267-5276.
[24]	Davidson EA, Keller M, Erickson HE, Verchot LV, Veldkamp E (2000). Testing a conceptual model of soil emissions of nitrous and nitric oxides: using two functions based on soil nitrogen availability and soil water content, the hole-in-the-pipe model characterizes a large fraction of the observed variation of nitric oxide and nitrous oxide emissions from soils. Bioscience, 50, 667-680.
[25]	Ding L (2009). Preliminary Study of Processes of Nitrous Oxide Emission from Soil & Plant in the Northeast Farmland. Master degree dissertation, Jilin Agricultural University, Changchun.
	[丁雷 (2009). 东北农田土壤和植物N₂O排放过程初探. 硕士学位论文, 吉林农业大学, 长春.]
[26]	Ding SL (2015). Heavy Metal Stress on the Influence of Robiniapseudoaeaeia-rhizobia Symbiosis System Rootexudate and Rhizosphere Soil Microbial Diversity. Master degree dissertation, North West Agriculture and Forestry University, Yangling, Shaanxi.
	[丁淑兰 (2015). 重金属胁迫对刺槐—根瘤菌共生体系的根系分泌物及土壤微生物多样性的影响. 硕士学位论文, 西北农林科技大学, 陕西杨凌.]
[27]	Dowdell RJ, Burford RCJR, Cannell RQ (1979). Oxygen concentrations in a clay soil after ploughing or direct drilling. Journal of Soil Science, 30, 239-245.
[28]	Dowdell EB (2006). Alcohol use, smoking, and feeling unsafe: health risk behaviors of two urban seventh grade classes. Issues in Comprehensive Pediatric Nursing, 30, 239-245.
[29]	Du R, Peng YZ, Cao SB, Wang SY, Niu M (2016). Characteristic of nitrous oxide production in partial denitrification process with high nitrite accumulation. Bioresource Technology, 203, 341-347. DOI PMID
[30]	Frolking SE, Mosier AR, Ojima DS, Li C, Parton WJ, Potter CS, Priesack E, Stenger R, Haberbosch C, Dörsch P, Flessa H, Smith KA (1998). Comparison of N₂O emissions from soils at three temperate agricultural sites: simulations of year-round measurements by four models. Nutrient Cycling in Agroecosystems, 52, 77-105.
[31]	Gautam H, Sehar Z, Rehman MT, Hussain A, AlAjmi MF, Khan NA (2021). Nitric oxide enhances photosynthetic nitrogen and sulfur-use efficiency and activity of ascorbate-glutathione cycle to reduce high temperature stress-induced oxidative stress in rice (Oryza sativa L.) plants. Biomolecules, 11, 305. DOI: 10.3390/biom11020305.
[32]	Giweta M, Dyck M, Malhi SS (2017). Effects of long-term fertilization history and current N and S fertilizer applications on nitrous oxide production from S-deficient soils in a laboratory incubation. Canadian Journal of Soil Science, 97, 0105. DOI: 10.1139/cjss-2016-0105.
[33]	Goshima N, Mukai T, Suemori M, Takahashi M, Caboche M, Morikawa H (1999). Emission of nitrous oxide (N₂O) from transgenic tobacco expressing antisense NiR mRNA. The Plant Journal, 19, 75-80. PMID
[34]	Gu JF, Yang JC (2022). Nitrogen (N) transformation in paddy rice field: its effect on N uptake and relation to improved N management. Crop and Environment, 1, 7-14.
[35]	Guo S, Zhou Y, Li Y, Gao Y, Shen Q (2008). Effects of different nitrogen forms and osmotic stress on water use efficiency of rice (Oryza sativa). Annals of Applied Biology, 153, 127-134.
[36]	Gupta KJ, Igamberdiev AU (2011). The anoxic plant mitochondrion as a nitrite: no reductase. Mitochondrion, 11, 537-543. DOI PMID
[37]	Gupta KJ, Stoimenova M, Kaiser WM (2005). In higher plants, only root mitochondria, but not leaf mitochondria reduce nitrite to NO, in vitro and in situ. Journal of Experimental Botany, 56, 2601-2609.
[38]	Hakata M, Takahashi M, Zumft W, Sakamoto A, Morikawa H (2003). Conversion of the nitrate nitrogen and nitrogen dioxide to nitrous oxides in plants. Acta Biotechnologica, 23, 249-257.
[39]	Harper JE (1981). Evolution of nitrogen oxide(s) during in vivo nitrate reductase assay of soybean leaves. Plant Physiology, 68, 1488-1493. DOI PMID
[40]	He M, Xu QY, Xia Y, Yang LM, Fan YX, Yang YS (2023). Plant phosphorus acquisition mechanisms and their response to global climate changes. Chinese Journal of Plant Ecology, 47, 291-305. DOI
	[何敏, 许秋月, 夏允, 杨柳明, 范跃新, 杨玉盛 (2023). 植物磷获取机制及其对全球变化的响应. 植物生态学报, 47, 291-305.] DOI
[41]	He ZH (2018). Effects of Potassium Fertilizer Application Rate and Base-topdressing Ratio on Flue-cured Tobacco Growth. Master degree dissertation, Yan’an University, Yan’an, Shaanxi.
	[贺治慧 (2018). 钾肥施用量和基追比对烤烟生长的影响. 硕士学位论文, 延安大学, 陕西延安.]
[42]	Hoff T, Truong HN, Caboche M (1994). The use of mutants and transgenic plants to study nitrate assimilation. Plant, Cell & Environment, 17, 489-506.
[43]	Houghton JT, Meira Filho LG, Bruce J, Hoesung Lee, Callander BA, Haites E, Harris N, Maskell K (1995). Climate Change 1994: Radiative Forcing of Climate Change and an Evaluation of the IPCC 1992 IS92 Emission Scenarios. Cambridge University Press, Cambridge, UK.
[44]	Hu HW, Chen DL, He JZ (2015). Microbial regulation of terrestrial nitrous oxide formation: understanding the biological pathways for prediction of emission rates. FEMS Microbiology Reviews, 39, 729-749.
[45]	Hu L, Wu Y, Dawuda M, Liao W, Lv J, Li Y, Yu J, Xie J, Feng Z, Zhang G, Calderón-Urrea A (2021). Appropriate ammonium/nitrate mitigates low light stress in Brassica pekinensis by regulating the nitrogen metabolism and expression levels of key proteins. Journal of Plant Growth Regulation, 40, 574-593.
[46]	Huang GH, Chen GX, Xu H, Wu J, Wang YJ, Yu KW (1992). Research on N₂O release from sterile soybean plants. Acta Botanica Sinica, 34, 835-839.
	[黄国宏, 陈冠雄, 徐惠, 吴杰, 王玉杰, 于克伟 (1992). 无菌大豆植株释放N₂O研究. 植物学报, 34, 835-839.]
[47]	Huang R (2019). Effects of Organic Substitution on Greenhouse Gas Emission and Nitrogen Transformation in Vegetable Garden Soil. PhD dissertation, Southwest University, Chongqing.
	[黄容 (2019). 有机替代对菜园土壤温室气体排放和氮转化的影响. 博士学位论文, 西南大学, 重庆.]
[48]	Hussain S, Khaliq A, Noor MA, Tanveer M, Hussain HA, Hussain S, Shah T, Mehmood T (2019). Metal toxicity and nitrogen metabolism in plants: an overview//Datta R, Meena RS, Pathan SI, Ceccherini MT. Carbon and Nitrogen Cycling in Soil. Springer, Singapore. 221-248.
[49]	Igamberdiev AU, Hill RD (2009). Plant mitochondrial function during anaerobiosis. Annals of Botany, 103, 259-268. DOI PMID
[50]	IPCC Intergovernmental Panel on Climate Change (2013). The Physical Science Basis. Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change. Cambridge University Press, Cambridge, UK.
[51]	IPCC Intergovernmental Panel on Climate Change (2019). Refinement to the 2006 IPCC Guidelines for National Greenhouse Gas Inventories. Cambridge University Press, Cambridge, UK.
[52]	IPCC Intergovernmental Panel on Climate Change (2021). Climate Change 2021—The Physical Science Basis: Working Group I Contribution to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change. Cambridge University Press, Cambridge, UK.
[53]	Itakura M, Uchida Y, Akiyama H, Hoshino YT, Shimomura Y, Morimoto S, Tago K, Wang Y, Hayakawa C, Uetake Y, Sánchez C, Eda SM, Hayatsu M, Minamisawa K (2013). Mitigation of nitrous oxide emissions from soils by Bradyrhizobium japonicum inoculation. Nature Climate Change, 3, 208-212.
[54]	Jiang SY (2018). The Effect of Low Nitrogen Nutrition on Root Growth and Nitrogen Absorption and Utilization of Wheat Seedlings and Its Physiological Mechanisms. PhD dissertation, Nanjing Agricultural University, Nanjing.
	[姜苏育 (2018). 低氮营养对小麦幼苗根系生长与氮素吸收利用的影响及其生理机制. 博士学位论文, 南京农业大学, 南京.]
[55]	Kaur H, Kaur H, Kaur H, Srivastava S (2023). The beneficial roles of trace and ultratrace elements in plants. Plant Growth Regulation, 100, 219-236.
[56]	Kawamura Y, Takahashi M, Arimura G, Isayama T, Irifune K, Goshima N, Morikawa H (1996). Determination of levels of NO₃^-, NO₂^- and NO₄⁺ ions in leaves of various plants by capillary electrophoresis. Plant and Cell Physiology, 37, 878-880.
[57]	Kim GW, Kim PJ, Khan MI, Lee SJ (2021). Effect of rice planting on nitrous oxide (N₂O) emission under different levels of nitrogen fertilization. Agronomy, 11, 217. DOI: 10.3390/agronomy11020217.
[58]	Klepper LA (1987). Nitric oxide emissions from soybean leaves during in vivo nitrate reductase assays. Plant Physiology, 85, 96-99. DOI PMID
[59]	Köberl M, Dita M, Martinuz A, Staver C, Berg G (2015). Agroforestry leads to shifts within the gammaproteobacterial microbiome of banana plants cultivated in Central America. Frontiers in Microbiology, 6, 91. DOI: 10.3389/fmicb.2015.00091. PMID
[60]	Kohl L, Koskinen M, Polvinen T, Tenhovirta S, Rissanen K, Patama M, Zanetti A, Pihlatie M (2021). An automated system for trace gas flux measurements from plant foliage and other plant compartments. Atmospheric Measurement Techniques, 14, 4445-4460.
[61]	Lenhart K, Behrendt T, Greiner S, Steinkamp J, Well R, Giesemann A, Keppler F (2019). Nitrous oxide effluxes from plants as a potentially important source to the atmosphere. New Phytologist, 221, 1398-1408. DOI PMID
[62]	Levy-Booth DJ, Prescott CE, Grayston SJ (2014). Microbial functional genes involved in nitrogen fixation, nitrification and denitrification in forest ecosystems. Soil Biology & Biochemistry, 75, 11-25.
[63]	Li N, Chen GX (1993). Effects of N-deficiency supply on root growth in wheat seedlings and its physiological. Journal of Applied Ecology, 4, 295-298.
	[李楠, 陈冠雄 (1993). 控释肥和覆草旱种对晚稻田CH₄和N₂O排放的影响. 应用生态学报, 4, 295-298.]
[64]	Li YY, Chen GX, Xu H, Zhang Y, Zhang XD (2003). The contribution of maize and soybean to N₂O emission from the soil-plant system during seedling stage. Environmental Science, 24(6), 38-42.
	[李玥莹, 陈冠雄, 徐慧, 张颖, 张旭东 (2003). 苗期玉米、大豆在土壤-植物系统N₂O排放中的贡献. 环境科学, 24(6), 38-42.]
[65]	Liu H, Zheng X, Li Y, Yu J, Ding H, Sveen TR, Zhang Y (2022). Soil moisture determines nitrous oxide emission and uptake. Science of the Total Environment, 822, 153566. DOI: 10.1016/j.scitotenv.2022.153566.
[66]	Lundberg JO, Weitzberg E, Gladwin MT (2008). The nitrate-nitrite-nitric oxide pathway in physiology and therapeutics. Nature Reviews Drug Discovery, 7, 156-167. DOI PMID
[67]	Malvi UR (2011). Interaction of micronutrients with major nutrients with special reference to potassium. Karnataka Journal of Agricultural Sciences, 24, 106-109.
[68]	Mosier AR, Mohanty SK, Bhadrachalam A, Chakravorti SP (1990). Evolution of dinitrogen and nitrous oxide from the soil to the atmosphere through rice plants. Biology and Fertility of Soils, 9, 61-67.
[69]	Mu XH, Chen YL (2021). The physiological response of photosynthesis to nitrogen deficiency. Plant Physiology and Biochemistry, 158, 76-82. DOI PMID
[70]	Okubo T, Ikeda S, Sasaki K, Ohshima K, Hattori M, Sato T, Minamisawa K (2014). Phylogeny and functions of bacterial communities associated with field-grown rice shoots. Microbes and Environments, 29, 329-332. PMID
[71]	Pacheco PJ, Bedmar EJ, Mesa S, Tortosa G, Delgado MJ (2023). Ensifer meliloti denitrification is involved in infection effectiveness and N₂O emissions from alfalfa root nodules. Plant and Soil, 486, 519-534.
[72]	Pang YX (2017). Research on the Mechanism of N₂O Emission from Land Plant Leaves. Master degree dissertation, Fujian Agriculture and Forestry University, Fuzhou.
	[庞亚星 (2017). 陆地植物叶片排放N₂O的机理研究. 硕士学位论文, 福建农林大学, 福州.]
[73]	Pate JS (1973). Uptake, assimilation and transport of nitrogen compounds by plants. Soil Biology & Biochemistry, 5, 109-119.
[74]	Paul EA, Clark FE (1989). Soil Microbiology and Biochemistry. Academic Press, Pittsburgh, USA. 147-163.
[75]	Poh LS, Jiang X, Zhang ZB, Liu Y, Ng WJ, Zhou Y (2015). N₂O accumulation from denitrification under different temperatures. Applied Microbiology and Biotechnology, 99, 9215-9226.
[76]	Qin SP, Pang YX, Hu HX, Liu T, Yuan D, Clough T, Wrage-Mönnig N, Luo JF, Zhou SG, Ma L, Hu CS, Oenema O (2024). Foliar N₂O emissions constitute a significant source to atmosphere. Global Change Biology, 30, e17181. DOI: 10.1111/gcb.17181.
[77]	Ravishankara AR, Daniel JS, Portmann RW (2009). Nitrous oxide (N₂O): the dominant ozone-depleting substance emitted in the 21st century. Science, 326, 123-125. DOI PMID
[78]	Rice CW, Rogers KL (1993). Denitrification in subsurface environments: potential source for atmospheric nitrous oxide//Harper LA, Mosier AR, Duxbury JM, Rolston chair DE. Agricultural Ecosystem Effects on Trace Gases and Global Climate Change. American Society of Agronomy, Crop Science Society of America, and Soil Science Society of America, Madison, USA. 121-132.
[79]	Rockel P, Strube F, Rockel A, Wildt J, Kaiser WM (2002). Regulation of nitric oxide (NO) production by plant nitrate reductase in vivo and in vitro. Journal of Experimental Botany, 53, 103-110. PMID
[80]	Rolston DE, Duxbury JM, Harper LA, Mosier AR, Rogers KL (1993). Denitrification in subsurface environments: potential source for atmospheric nitrous oxide//Harper LA, Mosier AR, Duxbury JM, Rolston DE. Agricultural Ecosystem Effects on Trace Gases and Global Climate Change. American Society of Agronomy, Crop Science Society of America, and Soil Science Society of America, Madison, USA.
[81]	Rückauf U, Augustin J, Russow R, Merbach W (2004). Nitrate removal from drained and reflooded fen soils affected by soil N transformation processes and plant uptake. Soil Biology & Biochemistry, 36, 77-90.
[82]	Sanchez-Cruz P, Alegría AE (2009). Quinone-enhanced reduction of nitric oxide by xanthine/xanthine oxidase. Chemical Research in Toxicology, 22, 818-823. DOI PMID
[83]	Saraste M, Castresana J (1994). Cytochrome oxidase evolved by tinkering with denitrification enzymes. FEBS Letters, 341, 1-4. PMID
[84]	Saunois M, Jackson RB, Bousquet P, Poulter B, Canadell JG (2016). The growing role of methane in anthropogenic climate change. Environmental Research Letters, 11, 120207. DOI: 10.1088/1748-9326/11/12/120207.
[85]	Shi YF, Ke XS, Yang XX, Liu YH, Hou X (2022). Plants response to light stress. Journal of Genetics and Genomics, 49, 735-747. DOI PMID
[86]	Signore A, Bell L, Santamaria P, Wagstaff C, van Labeke MC (2020). Red light is effective in reducing nitrate concentration in rocket by increasing nitrate reductase activity, and contributes to increased total glucosinolates content. Frontiers in Plant Science, 11, 604. DOI: 10.3389/fpls.2020.00604. PMID
[87]	Smart DR, Bloom AJ (2001). Wheat leaves emit nitrous oxide during nitrate assimilation. Proceedings of the National Academy of Sciences of the United States of America, 98, 7875-7878. PMID
[88]	Song T, Zhang XL, Li J, Wu XY, Feng HX, Dong WY (2021). A review of research progress of heterotrophic nitrification and aerobic denitrification microorganisms (HNADMs). Science of the Total Environment, 801, 149319. DOI: 10.1016/j.scitotenv.2021.149319.
[89]	Stange CF, Spott O, Arriaga H, Menéndez S, Estavillo JM, Merino P (2013). Use of the inverse abundance approach to identify the sources of NO and N₂O release from Spanish forest soils under oxic and hypoxic conditions. Soil Biology & Biochemistry, 57, 451-458.
[90]	Stoimenova M, Igamberdiev AU, Gupta KJ, Hill RD (2007). Nitrite-driven anaerobic ATP synthesis in barley and rice root mitochondria. Planta, 226, 465-474. DOI PMID
[91]	Sun H (2018). Study on the Emission of N₂O from Plants Under the Co-denitrification of Endophytes and Plants. PhD dissertation, University of Chinese Academy of Sciences, Beijing.
	[孙浩 (2018). 内生菌-植物共反硝化作用下植物源N₂O的排放研究. 博士学位论文, 中国科学院大学, 北京.]
[92]	Sun H, Li YT, Xu H (2018). Is endophyte-plant co-denitrification a source of nitrous oxides emission? —An experimental investigation with soybean. Agronomy, 8, 108. DOI: 10.3390/agronomy8070108.
[93]	Sun HY, Xin QW, Lin XS, Luo HL, Lin H, Yan SJ, Liu WL, Lan SR (2019). Effects of plant species and diversity on methane emissions and functional gene abundances in constructed wetlands. Acta Ecologica Sinica, 39, 8565-8574.
	[孙红英, 辛全伟, 林兴生, 罗海凌, 林辉, 严少娟, 刘文莉, 兰思仁 (2019). 人工湿地植物种类及多样性对甲烷释放及功能基因丰度的影响. 生态学报, 39, 8565-8574.]
[94]	Sun YJ, Wu H, Wang YN (2011). The influence factors on N₂O emissions from nirification and denitrification reaction. Ecology and Environmental Sciences, 20, 384-388.
	[孙英杰, 吴昊, 王亚楠 (2011). 硝化反硝化过程中N₂O释放影响因素. 生态环境学报, 20, 384-388.] DOI
[95]	Syakila A, Kroeze C (2011). The global nitrous oxide budget revisited. Greenhouse Gas Measurement and Management, 1, 17-26.
[96]	Tang W, Guo HP, Baskin CC, Xiong WD, Yang C, Li ZY, Song H, Wang TR, Yin JN, Wu XL, Miao FH, Zhong SZ, Tao QB, Zhao YR, Sun J (2022). Effect of light intensity on morphology, photosynthesis and carbon metabolism of alfalfa (Medicago sativa) seedlings. Plants, 11, 1688. DOI: 10.3390/plants11131688.
[97]	Thörn M, Sörensson F (1996). Variation of nitrous oxide formation in the denitrification basin in a wastewater treatment plant with nitrogen removal. Water Research, 30, 1543-1547.
[98]	Tian H, Yang J, Lu C, Xu R, Canadell JG, Jackson RB, Arneth A, Chang J, Chen G, Ciais P, Gerber S, Ito A, Huang Y, Joos F, Lienert S, et al. (2018). The global N₂O model intercomparison project. Bulletin of the American Meteorological Society, 99, 1231-1251.
[99]	Timilsina A, Dong W, Luo J, Lindsey S, Wang Y, Hu C (2020). Nitrogen isotopic signatures and fluxes of N₂O in response to land-use change on naturally occurring saline-alkaline soil. Scientific Reports, 10, 21253. DOI: 10.1039/s41598-020-78149-w. PMID
[100]	Timilsina A, Oenema O, Luo J, Wang Y, Dong W, Pandey B, Bizimana F, Zhang Q, Zhang C, Yadav RKP, Li X, Liu X, Liu B, Hu C (2022). Plants are a natural source of nitrous oxide even in field conditions as explained by ¹⁵N site preference. Science of the Total Environment, 805, 150262. DOI: 10.1016/j.scitotenv.2021.150262.
[101]	Tischner R, Planchet E, Kaiser WM (2004). Mitochondrial electron transport as a source for nitric oxide in the unicellular green alga Chlorella sorokiniana. FEBS Letters, 576, 151-155. DOI PMID
[102]	Turner NC (1986). Crop water deficits: a decade of progress. Advances in Agronomy, 39, 1-51.
[103]	Ussiri D, Lal R (2012). Global sources of nitrous oxide//Ussiri D, Lal R. Soil Emission of Nitrous Oxide and Its Mitigation. Springer, Dordrecht, The Netherlands. 131-175.
[104]	Wang SJ, Duan SL, George TS, Feng G, Zhang L (2024). Adding plant metabolites improve plant phosphorus uptake by altering the rhizosphere bacterial community structure. Plant and Soil, 497, 503-522.
[105]	Wang WF, Zhai YY, Cao LX, Tan HM, Zhang RD (2017). Improvement of rice seedling growth and nitrogen use efficiency by seed inoculation with endophytic denitrifiers. Environmental Science and Pollution Research International, 24, 14477-14483. DOI PMID
[106]	Wang S, Duan S, George TS, Zhang L (2023). Adding plant metabolites improve plant phosphorus uptake by altering the rhizosphere bacterial community structure. Plant and Soil, 497, 503-522.
[107]	Wen Y, Corre MD, Rachow C, Chen L, Veldkamp E (2017). Nitrous oxide emissions from stems of alder, beech and spruce in a temperate forest. Plant and Soil, 420, 423-434.
[108]	Wrage N, Velthof GL, van Beusichem ML, Oenema O (2001). Role of nitrifier denitrification in the production of nitrous oxide. Soil Biology & Biochemistry, 33, 1723-1732.
[109]	Yan X, Shi S, Du L, Xing G (2000). Pathways of N₂O emission from rice paddy soil. Soil Biology & Biochemistry, 32, 437-440.
[110]	Yang SH, Chen GX, Lin JH, Wu J, Ma YQ (1995). N₂O emission from woody plants and its relation to their physiological activities. Chinese Journal of Applied Ecology, 6, 337-340.
	[杨思河, 陈冠雄, 林继惠, 吴杰, 马越强 (1995). 几种木本植物的N₂O释放与某些生理活动的关系. 应用生态学报, 6, 337-340.]
[111]	Yang Y, Chen GX, Li YY, Cui ZH, Zhang LJ (2005). N₂O emission from soybean in relation to light density and photosynthesis. Journal of Shenyang Agricultural University, 36(3), 282-285.
	[杨宇, 陈冠雄, 李玥莹, 崔震海, 张立军 (2005). 大豆N₂O释放与光照度和光合作用的关系. 沈阳农业大学学报, 36(3), 282-285.]
[112]	Yao H, Campbell CD, Chapman SJ, Freitag TE, Nicol GW, Singh BK (2013). Multi-factorial drivers of ammonia oxidizer communities: evidence from a national soil survey. Environmental Microbiology, 15, 2545-2556. DOI PMID
[113]	Yu KW, Huang B, Chen GX, Wu J (1997). Field measurement of N₂O flux from soybean plant and effect of light on it. Chinese Journal of Applied Ecology, 8, 171-176.
	[于克伟, 黄斌, 陈冠雄, 吴杰 (1997). 田间大豆植株N₂O通量的测定及光照的影响. 应用生态学报, 8, 171-176.]
[114]	Yu KW, Wang ZP, Chen GX (1997). Nitrous oxide and methane transport through rice plants. Biology and Fertility of Soils, 24, 341-343.
[115]	Zhai YJ, Li YH, Chen YH (2013). Major progress of global and China regional climate change projection. Journal of Arid Meteorology, 31, 803-813.
	[翟颖佳, 李耀辉, 陈玉华 (2013). 全球及中国区域气候变化预估研究主要进展简述. 干旱气象, 31, 803-813.] DOI
[116]	Zhang C, Ju X, Zhang J, Rees RM, Müller C (2023). Soil pH and long-term fertilization affect gross N transformation and N₂O production pathways in Chinese and UK croplands. Biology and Fertility of Soils, 59, 527-539.
[117]	Zhang J, Müller C, Cai Z (2015). Heterotrophic nitrification of organic N and its contribution to nitrous oxide emissions in soils. Soil Biology & Biochemistry, 84, 199-209.
[118]	Zhang LJ, Yang Y, Ruan YY, Fan JJ, Chen GX (2006). A new model of light regulation for N₂O emission from plants. Journal of Shenyang Agricultural University, 37(2), 131-136.
	[张立军, 杨宇, 阮燕晔, 樊金娟, 陈冠雄 (2006). 一种新的植物N₂O释放光调节模式. 沈阳农业大学学报, 37(2), 131-136.]
[119]	Zhang M, Huang SB, Xiao XN (2012). Effect of C/N ratio and pH on nitrous oxide production of themophic aerobic denitrifier. Chinese Journal of Environmental Engineering, 61, 275-279.
	[张苗, 黄少斌, 肖先念 (2012). C/N和pH值对高温好氧反硝化菌产N₂O的影响研究. 环境工程学报, 6, 275-279.]
[120]	Zhao D, Liu XM (2018). Changes in organic nitrogen components during the decomposition process of sheep manure and cow manure in typical grasslands of Inner Mongolia. Journal of Inner Mongolia University (Natural Science Edition), 49(2), 174-181.
	[赵东, 刘新民 (2018). 内蒙古典型草原羊粪和牛粪分解过程中有机氮组分的变化. 内蒙古大学学报(自然科学版), 49(2), 174-181.]
[121]	Zhao X, Sampath V, Caughey WS (1995). Cytochrome c oxidase catalysis of the reduction of nitric oxide to nitrous oxide. Biochemical and Biophysical Research Communications, 212, 1054-1060. PMID
[122]	Zhu CF, Luo HD, Luo LC, Wang KY, Liao Y, Zhang S, Huang SS, Guo XM, Zhang L (2022). Nitrogen and biochar addition affected plant traits and nitrous oxide emission from Cinnamomum camphora. Frontiers in Plant Science, 13, 905537. DOI: 10.3389/fpls.2022.905537.
[123]	Zhu TX, Gao K, Zhang YL (2011). 钾肥对小黑麦拔节期生物量及品质的影响. Effect of potassium fertilizer on biomass and quality at jointing stage of Triticale. Crops, (6), 57-59.
	[朱铁霞, 高凯, 张永亮 (2011). 钾肥对小黑麦拔节期生物量及品质的影响. 作物杂志, (6), 57-59.]
[124]	Zou JW, Huang Y, Sun WJ, Zheng XH (2005). Contribution of plants to N₂O emissions in soil-winter wheat ecosystem: pot and field experiments. Plant and Soil, 269, 205-211.