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施加生物炭缓解土壤氮流失机理的研究进展

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史广宇, 吴贝贝, 胡嘉源, 施维林. 施加生物炭缓解土壤氮流失机理的研究进展[J]. 环境化学. doi: 10.7524/j.issn.0254-6108.2023072502
引用本文: 史广宇, 吴贝贝, 胡嘉源, 施维林. 施加生物炭缓解土壤氮流失机理的研究进展[J]. 环境化学. doi: 10.7524/j.issn.0254-6108.2023072502
shu
SHI Guangyu, WU Beibei, HU Jiayuan, SHI Weilin. Advances in nitrogen loss reduction mechanism study by biochar application[J]. Environmental Chemistry. doi: 10.7524/j.issn.0254-6108.2023072502
Citation: SHI Guangyu, WU Beibei, HU Jiayuan, SHI Weilin. Advances in nitrogen loss reduction mechanism study by biochar application[J]. Environmental Chemistry. doi: 10.7524/j.issn.0254-6108.2023072502
shu

施加生物炭缓解土壤氮流失机理的研究进展

  • 1.

    苏州科技大学环境科学与工程学院，苏州，215009

  • 2.

    苏州科技大学天平学院，苏州，215009

    • 通讯作者: Email：shigy@usts.edu.cn; 
  • 基金项目:
    国家自然科学基金青年基金（41807142），苏州市科技局科技计划项目（SNG2020053）和苏州市科技局科技计划项目（SNG2022072）资助.

Advances in nitrogen loss reduction mechanism study by biochar application

  • 1.

    School of Environmental Science and Engineering, Suzhou University of Science and Technology, Suzhou, 215009, China

  • 2.

    Tianping College of Suzhou University of Science and Technology, Suzhou, 215009, China

    • Corresponding author: SHI Guangyu, shigy@usts.edu.cn ; 
  • Fund Project: the the National Natural Science Foundation of China (41807142), Suzhou Technology Office Program (SNG2020053) and the Suzhou Technology Office Program (SNG2022072).

  • 摘要: 施加生物炭不仅可以抑制农田土壤氮素流失，还可以实现农业废弃物资源化利用. 因此，本文分析了现有研究结果，重点剖析生物炭的NH4+-N和NO3−-N吸附容量的差异和施加生物炭对土壤理化性质、微生物群落结构和植物氮吸收能力的影响，综述生物炭抑制土壤氮素流失的机理，并对今后深入研究施加生物炭缓解土壤氮素流失的有关方向进行展望. 施加生物炭缓解土壤氮素流失的机理主要为：缓解NH4+-N和NO3−-N的流失、抑制N2O的逸散、提高植物吸收氮素的能力. 生物炭不仅拥有较强的NH4+-N吸附能力，还具有提高土壤碳含量、pH值和土壤含水量等能力，从而降低因地表径流和水分入渗导致的NH4+-N和NO3−-N流失量. 施加生物炭抑制土壤中N2O的逸散的主要原因是生物炭可以促进土壤中氨氧化古菌、氨氧化细菌、完全氨氧化菌等微生物的富集、提高nxrA、napAB和nrfA等基因的丰度和亚硝酸氧化还原酶活性，以此降低反硝化反应速率. 此外，施加生物炭有助于降低根系生长阻力，促进植物根系生长发育，并提高植物吸收氮素的能力，进一步降低土壤溶液中NO3−-N的浓度. 未来关于施加生物炭缓解农田土壤氮流失领域的研究建议从以下几个方向展开：探究生物炭热解方式、理化性质和施加方法等因素对农田土壤氮流失的影响；关注生物炭促进土壤中异化硝酸盐还原成铵反应和完全氨氧化反应对减少N2O逸散量的贡献；重视生物炭材料中金属离子对土壤中铁氨氧化反应、厌氧氨氧化反应等氮循环途径的影响.
    Abstract: Biochar application can not only reduce the nitrogen loss in farmland, but also realize the resource utilization of agricultural waste. Thus, in this paper, experimental results of existing studies were analyzed, as well as the differences in the adsorption capacity of biochar NH4+-N and NO3−-N and the effects on soil physicochemical properties, microbial community structure, and plant nitrogen absorption capacity , the mechanisms of reducing nitrogen loss from soil by biochar application was clarified, and future research priorities were prospected. The mechanisms of reducing nitrogen loss from soil by biochar application mainly include: reducing the loss of NH4+-N and NO3−-N, mitigating the N2O emission, and improving the ability of plants to absorb nitrogen. Biochar not only has strong NH4+-N adsorption capacity, but also has the ability to increase soil carbon content, pH value and soil water content, so as to reduce the loss of NH4+-N and NO3−-N caused by surface runoff and water infiltration. Biochar application can mitigate N2O emission in the soil is because biochar could enrich the microorganisms such as ammonia oxidizing archaea, ammonia oxidizing bacteria, and cmplete ammonia oxidizing bacteria in the soil, increase the abundance of genes such as nxrA, napAB, and nrfA, and enhance nitrite oxidoreductase activity, thereby reducing the rate of denitrification. Additionally, biochar application could reduce root growth resistance, promote plant root growth, and improve the ability of plants to absorb nitrogen, thereby further reducing the concentration of NO3−-N in the soil. Future research is suggested to be carried out in the following aspects: exploring the effects of biochar pyrolysis methods, physicochemical properties and application methods on nitrogen loss from soil; focusing on the contribution of biochar application to the reduction of N2O emission by improving the efficiency of dissimilatory nitrate reduction to ammonium reaction and anammox reaction; emphasizing the impact of metal ions in biochar on nitrogen cycle pathways such as feammox reaction and anammox reaction in soil.
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  • 图 1  生物炭NH4+-N和NO3−-N吸附容量箱形图

    Figure 1.  Box plot of NH4+-N and NO3−-N adsorption capacity of biochar

    下载: 全尺寸图片 幻灯片

    图 2  生物炭理化性质与NH4+-N吸附容量皮尔逊相关性分析

    Figure 2.  Pearson correlation analysis of physicochemical properties of biochar and NH4+-N adsorption capacity of biochar

    下载: 全尺寸图片 幻灯片

    图 3  土壤中NH4+-N、NO3−-N、NO2−-N、N2O和NH3参与的主要反应过程

    Figure 3.  Main reaction processes involved in NH4+-N, NO3−-N, NO2−-N, N2O and NH3 in soil

    下载: 全尺寸图片 幻灯片

    图 4  施加生物炭对植物株高、生物量、作物产量的影响

    Figure 4.  Effects of biochar returning on plant height, biomass and crop yield

    下载: 全尺寸图片 幻灯片

    表 1  施加生物炭对土壤理化性质的影响

    Table 1.  Effects of biochar returning on soil physicochemical properties

    材料
    Materials    		 pH 含水量
    Water content    		 容重
    Bulk density    		 SOC 总氮
    Total nitrogen    		 碱解氮
    Akaline nitrogen decomposition    		 参考文献
    References
    牛粪生物炭
    Cow dung biochar    		 42.5% — — 148.6% 72.0% — [54]
    棉花秸秆生物炭
    Cotton straw biochar    		 — 54.2% —7.6% 6.4% 2.5% — [55]
    玉米秸秆生物炭
    Corn stover biochar    		 15.0% — — 45.9% — 19.0% [56]
    5.3% — — 145.0% — 12.2% [57]
    6.9% 17.4% —9.87% — 6.73% — [58]
    1.2% 13.9% —14.3% 54.1% 26.2% 23.5% [59]
    小麦秸秆生物炭
    Wheat straw biochar    		 7.9% — — 344.8% 155.1% — [60]
    5.3% 36.7% — — — 50.0% [61]
    水稻秸秆生物炭
    Rice straw biochar    		 5.2% — — 29.3% 4.5% — [62]
    苹果枝生物炭
    Apple branch biochar    		 7.8% 24.6% −12.9% — — — [63]
    11.2% — — 207.4% 67.5% — [64]
    稻壳生物炭
    Rice husk biochar    		 5.4% 47.9% −30.4% — 29.8% — [65]
    秸秆生物炭
    Straw biochar    		 5.2% −8.0% −5.6% — 112.8% — [65]
    — 42.2% −19.6% 179.6% 30.5% — [66]
    2.4% 44.7% −32.5% 297.2% — — [67]
    　　表中所有数值均为与CK组对比换算后的各指标变化的百分比，“负数”表示该指标的数值因生物炭添加而降低.
    　　All values in the table are converted percentage changes in each indicator compared to the CK group. “—” indicates that the value of this indicator decreases due to the biochar application.
    材料
    Materials    		 pH 含水量
    Water content    		 容重
    Bulk density    		 SOC 总氮
    Total nitrogen    		 碱解氮
    Akaline nitrogen decomposition    		 参考文献
    References
    牛粪生物炭
    Cow dung biochar    		 42.5% — — 148.6% 72.0% — [54]
    棉花秸秆生物炭
    Cotton straw biochar    		 — 54.2% —7.6% 6.4% 2.5% — [55]
    玉米秸秆生物炭
    Corn stover biochar    		 15.0% — — 45.9% — 19.0% [56]
    5.3% — — 145.0% — 12.2% [57]
    6.9% 17.4% —9.87% — 6.73% — [58]
    1.2% 13.9% —14.3% 54.1% 26.2% 23.5% [59]
    小麦秸秆生物炭
    Wheat straw biochar    		 7.9% — — 344.8% 155.1% — [60]
    5.3% 36.7% — — — 50.0% [61]
    水稻秸秆生物炭
    Rice straw biochar    		 5.2% — — 29.3% 4.5% — [62]
    苹果枝生物炭
    Apple branch biochar    		 7.8% 24.6% −12.9% — — — [63]
    11.2% — — 207.4% 67.5% — [64]
    稻壳生物炭
    Rice husk biochar    		 5.4% 47.9% −30.4% — 29.8% — [65]
    秸秆生物炭
    Straw biochar    		 5.2% −8.0% −5.6% — 112.8% — [65]
    — 42.2% −19.6% 179.6% 30.5% — [66]
    2.4% 44.7% −32.5% 297.2% — — [67]
    　　表中所有数值均为与CK组对比换算后的各指标变化的百分比，“负数”表示该指标的数值因生物炭添加而降低.
    　　All values in the table are converted percentage changes in each indicator compared to the CK group. “—” indicates that the value of this indicator decreases due to the biochar application.
    下载: 导出CSV
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  • 收稿日期:  2023-07-25
  • 录用日期:  2023-09-27

施加生物炭缓解土壤氮流失机理的研究进展

    通讯作者: Email：shigy@usts.edu.cn; 
  • 1. 苏州科技大学环境科学与工程学院，苏州，215009
  • 2. 苏州科技大学天平学院，苏州，215009
  • 收稿日期:  2023-07-25
  • 录用日期:  2023-09-27
  • 网络出版日期:  2024-03-29
基金项目:
国家自然科学基金青年基金（41807142），苏州市科技局科技计划项目（SNG2020053）和苏州市科技局科技计划项目（SNG2022072）资助.

摘要: 施加生物炭不仅可以抑制农田土壤氮素流失，还可以实现农业废弃物资源化利用. 因此，本文分析了现有研究结果，重点剖析生物炭的NH4+-N和NO3−-N吸附容量的差异和施加生物炭对土壤理化性质、微生物群落结构和植物氮吸收能力的影响，综述生物炭抑制土壤氮素流失的机理，并对今后深入研究施加生物炭缓解土壤氮素流失的有关方向进行展望. 施加生物炭缓解土壤氮素流失的机理主要为：缓解NH4+-N和NO3−-N的流失、抑制N2O的逸散、提高植物吸收氮素的能力. 生物炭不仅拥有较强的NH4+-N吸附能力，还具有提高土壤碳含量、pH值和土壤含水量等能力，从而降低因地表径流和水分入渗导致的NH4+-N和NO3−-N流失量. 施加生物炭抑制土壤中N2O的逸散的主要原因是生物炭可以促进土壤中氨氧化古菌、氨氧化细菌、完全氨氧化菌等微生物的富集、提高nxrA、napAB和nrfA等基因的丰度和亚硝酸氧化还原酶活性，以此降低反硝化反应速率. 此外，施加生物炭有助于降低根系生长阻力，促进植物根系生长发育，并提高植物吸收氮素的能力，进一步降低土壤溶液中NO3−-N的浓度. 未来关于施加生物炭缓解农田土壤氮流失领域的研究建议从以下几个方向展开：探究生物炭热解方式、理化性质和施加方法等因素对农田土壤氮流失的影响；关注生物炭促进土壤中异化硝酸盐还原成铵反应和完全氨氧化反应对减少N2O逸散量的贡献；重视生物炭材料中金属离子对土壤中铁氨氧化反应、厌氧氨氧化反应等氮循环途径的影响.

English Abstract

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  • 大量施用氮肥满足了我国日益增长的粮食需求，但也造成极大的环境风险. 研究表明，我国平均施氮量和氮利用率分别为全球平均水平的412%和60%[1- 2]，约50%氮肥通过径流、淋溶和挥发等途径流失[3]. 氮肥进入土壤后，聚合物形式的有机氮先降解为可溶性有机氮，再矿化为极易流失的可溶性无机氮（主要为铵态氮（NH4+-N）和硝态氮（NO3−-N））[4]. 降雨和灌溉引起的地表径流和水分入渗极易造成可溶性无机氮流失. 径流初期NH4+-N易随表层土壤颗粒迁移，随后NO3−-N流失量显著上升约占氮流失量的70%－90%[5 − 6]. NO3−-N不仅是径流氮素流失的主要形态，还易随水分入渗向深层土壤迁移并污染地下水. Song等[7]指出，土壤年氮损失中约90%的氮素通过淋溶途径流失，NO3−-N流失占溶解氮损失的65%－90%. 除土壤含水量、降水量等自然因素外，施加化肥将导致土壤氮流失量提高约105%－123%[8 − 9]、农业生产成本显著上升和面源污染加重[10]. 研究表明，造成水体污染的氮素约77%来自农业，远高于畜牧养殖和生活排污[11]，农田氮素流失对水质恶化和水体富营养化的贡献逐年上升. 当前，我国农田氮流失情况严重[12]，氮流失高风险和极高风险区约占总调查面积的20%以上[13 − 14]，我国将面对农业土壤总氮流失显著增加的巨大挑战.

    国内外学者常将秸秆还田以缓解农田土壤氮流失，但秸秆还田可能会抑制播种期作物生长、加重作物病虫害、促进一氧化二氮（N2O）、甲烷（CH4）、二氧化碳（CO2）等温室气体的排放[15 − 18]，其中N2O累计排放量最高可增加1421%[19]. 此外秸秆还田减少NO3−-N淋溶流失的能力有限，秸秆内的氮素溶于径流还会导致土壤氮流失总量上升[20]. 但将秸秆制成生物炭,可以降低秸秆还田引发农田周边水体富营养化的风险[21]. 近年，大量研究证实，施加生物炭在减少氮素流失[22 − 24]、提升土壤肥力[25 − 27]等方面具有很高的工程应用价值和广阔的发展前景. 我国每年产生超过40多亿吨的农业废弃物，以农业废弃物为原料热解制成的生物炭既解决了废弃物随意处理造成的环境问题，又实现了农业废弃物资源化利用[28].

    因此，本文将从生物炭吸附NH4+-N和NO3−-N的能力和施加生物炭对土壤理化性质、微生物群落结构、氮循环相关基因表达、植物氮摄取能力的影响等方面阐述施加生物炭缓解土壤氮素流失的机理，以期为今后的研究提供理论基础.

    • 1.   生物炭对NH4+-N和NO3−-N的吸附（Adsorption of NH4+-N and NO3−-N by biochar）

    • 农田土壤中的氮素主要通过径流、淋溶和气体逸散等途径流失. 研究表明，农业生产过程中约5%－42%的氮肥通过淋溶途径流失，流失的氮肥中约19%为NO3−-N[29]. 施加生物炭可以有效阻止NO3－-N随水分入渗向深层土壤迁移，淋溶实验结果表明施加生物炭的土壤中NO3−-N主要分布由0－300 cm缩短为0－40 cm[30]. 陈灿等[31]实验结果指出，施加生物炭显著减少土壤NO3−-N淋溶损失约14%－76%. 土壤中氨气（NH3）和N2O的逸散不仅影响氮肥利用率还会造成雾霾、温室效应等环境问题[32]，抑制农田土壤NH3和N2O的逸散是减少农田土壤氮素流失的另一有效手段[33]. 土壤中施加生物炭不仅能显著降低NH3的挥发损失，还对N2O的逸散具有优秀的“削峰”能力[34 − 36]，例如家禽粪便生物炭减少土壤NH3挥发量约53%[37]，竹炭减少茶园土壤约79%的NH3逸出[38]，秸秆生物炭降低N2O峰值排放通量约70%[39].

      对Web of Science核心合集和CNKI数据库中关于生物炭的NH4+-N、NO3−-N吸附容量研究的数据进行提取和分析. 英文文献数据的检索条件为“TS=（biochar AND ammonium nitrogen AND adsorption）”和“TS=（biochar AND nitrate nitrogen AND adsorption）”；中文文献数据的检索条件为主题=“生物炭”并且“铵态氮”并且“吸附”和“生物炭”并且“硝态氮”并且“吸附”. 筛选后共得到NH4+-N吸附容量有效数据131个，NO3−-N吸附容量有效数据89个. 利用Origin 2022软件对筛选所得数据进行分析，结果如图1所示. 生物炭NH4+-N吸附容量大多分布在2.2－40.0 mg·g−1范围内，生物炭NO3−-N吸附容量分布范围较小，约为0.1－1.3 mg·g−1，生物炭NH4+-N和NO3−-N吸附容量的中位数分别为8.2 mg·g−1和0.2 mg·g−1，生物炭NH4+-N和NO3−-N吸附容量的均值分别约为26.1 mg·g−1和2.4 mg·g−1. 结果表明，生物炭对NH4+-N具有较强的吸附能力.

      Yadav等[40]实验结果指出，土壤中NH4+-N、NO3−-N的浓度与生物炭表面官能团数量具有良好的正相关性，生物炭吸附NH4+-N和NO3−-N的能力可能是缓解土壤氮素流失的关键. 现有研究证实，生物炭的NH4+-N吸附量分别与比表面积和表面官能团具有相关性[41]. 对筛选所得文献中生物炭理化性质和吸附容量的数据进行皮尔逊相关性分析，结果如图2所示. 生物炭的NH4+-N吸附容量与比表面积显著正相关（P＜0.05）. 生物炭巨大的比表面积和发达的孔隙结构为NH4+-N提供了充足的吸附位点，生物炭比表面积增加16%－32%，NH4+-N吸附容量提高103%－214%[42]，因此比表面积更大的生物炭NH4+N吸附容量可能更高[43]. 酸性官能团亦显著影响生物炭NH4+N吸附容量，其赋予生物炭阳离子交换能力[44 − 45]，因此生物炭吸附NH4+-N的能力与酸性官能团数量呈正比[43]. 生物炭对NO3−-N的吸附能力较弱，其可能通过羟基（—OH）、芳香环羰基（—C═O）、脂肪族醚类（—O—）等基团形成氢键或π键[46]以及叔胺基（-NH2）、Fe—O等带正电荷的基团静电吸附等途径吸附NO3−N [47 − 48]. 并且有研究显示，施加老化生物炭的促进效果优于新鲜生物炭[49]，证明生物炭减少土壤氮淋失的效应具有长期性，且冻融循环老化生物炭>高温老化生物炭>自然老化生物炭>新鲜生物炭[50]. 综上，生物炭对NH4+-N的吸附容量更大、吸附强度更高，因此生物炭对NH4+-N的吸附能力在缓解土壤NH4+-N和N2O流失方面发挥重要作用，而生物炭缓解土壤NO3−-N流失则更依赖于生物炭对土壤理化性质和微生物活动的影响.

    2.   施加生物炭改变土壤理化性质和微生物群落结构（Alteration of soil physicochemical properties and microbial community structure by biochar application）

    • 生物炭缓解土壤氮素流失的机理十分复杂，现有研究认为生物炭减少土壤氮素流失主要通过吸附无机氮（NH4+-N和NO3−-N）、改变土壤理化性质、提高植物的氮素吸收能力等途径抑制NH4+-N和NO3−-N的流失；通过影响土壤微生物群落结构、调控土壤氮循环等途径减少N2O的逸散，以此达到抑制土壤氮素流失、提升土壤肥力的目的[33,51].

      • 2.1.   施加生物炭改变土壤理化性质

    • 施加生物炭能显著改变土壤理化性质，Singh等[52]meta分析结果证实，施加生物炭使土壤pH值、阳离子交换量、总有机碳（SOC）含量和孔隙率分别提高46%、20%、27%和59%. 改变土壤理化性质是生物炭减少土壤氮流失、提高土壤总氮含量的关键[53]. 如表1所示，施加生物炭有效提高土壤SOC含量、pH值和含水量，降低土壤容重.

      土壤碳含量被认为是土壤总氮和无机氮含量的主要预测因子[68]. Gao等[69]的meta分析结果证实，施加生物炭可以提高土壤中SOC含量，土壤中无机氮含量受SOC含量影响最大. 并与生物炭添加量呈正比[70]. 土壤SOC含量还对amoA、nirK、nirS和nosZ等基因的丰度有显著影响，这些基因的表达参与调控氨氧化、硝化、反硝化等过程[71]，因此生物炭提高土壤SOC含量是生物炭减少土壤中氮流失的重要途径之一[72 − 73]. Phillips等[74]实验结果指出，施加生物炭显著降低土壤中无机氮的生成量约36%. 这是由于生物炭提高土壤C/N比，降低土壤中氮肥矿化速率[75]，减少土壤中NH4+-N的含量[76]，抑制土壤中氨化作用从而降低硝化速率[77]. 土壤中有机碳的分解还可以为异化硝酸盐还原成铵（DNRA）过程提供电子受体和易被微生物所利用的碳源[78]. 土壤硝化速率下降和DNRA反应速率上升导致土壤中NO2−-N和NO3−-N的生成量下降，N2O和N2的逸散量减少，以此缓解土壤氮素流失、提高土壤总氮含量.

      土壤在连续施加氮肥后会出现明显的酸化现象，土壤pH值下降10%－24%，酸化产生的H+会影响NH4+-N的吸附[79]，施加生物炭提高土壤pH值不仅可以提高土壤对NH4+-N的吸附能力，还可以刺激酸性土壤中氨氧化基因的表达和硝化细菌的富集[71, 80]. 氨氧化和硝化速率的提升促进土壤中NO2−-N氧化为NO3−-N，研究表明，施加生物炭后土壤中NO3−-N含量提高36%－89%[81]，而NO2−-N浓度降低导致反硝化反应速率降低和N2O的生成量减少[82]. 因此，土壤pH值在控制土壤 N2O 排放方面发挥着核心作用.

      施加生物炭影响土壤容重和孔隙特征，进而影响土壤的持水能力和导水性能. 研究指出，土壤饱和导水率与土壤中NH4+-N和NO3−-N含量线性相关[83]，生物炭添加可以减少风砂土、黄绵土和砂质潮土等土壤的饱和导水率42%－89%[84]，从而减少NH4+-N和NO3−-N因水分入渗造成的淋溶流失[85]. 生物炭添加还可以改变土壤孔隙分布，提高土壤含水率[86]，以此达到降低NO3−-N淋溶损失[87 − 88]，减少N2O逸散[89]，促进NH3的溶解[90]的目的. 此外，土壤中氮素主要被大团聚体吸附和贮存，生物炭能促进土壤中大型团聚体形成（粒径＞2 mm），减少土壤中黏粒所占百分比25%－40%[91]，提升土壤有机氮贮存和供给的能力[92].

      综上，施加生物炭通过改变土壤理化特性可以有效抑制NH4+-N和NO3−-N淋溶流失，影响土壤微生物生理活动，调控土壤中氨氧化反应、硝化反应、反硝化反应和DNRA反应速率，降低土壤中NO2−-N浓度和N2O的生成量.

      • 2.2.   施加生物炭改变土壤微生物群落结构，调控氮循环功能基因表达

    • 土壤中的硝化反应和反硝化反应是N2O产生的主要来源，N2O逸散受温度、降雨量、土壤pH值、土壤有机碳含量、微生物活动等因素影响，因此减少施肥土壤中N2O的逸散十分具有挑战性. 生物炭通过改变土壤微生物群落结构、影响酶活性、调控基因表达等途径减少土壤中N2O的逸散量约54%[93]. 研究表明，施加生物炭对土壤中氨氧化古菌（AOA）、氨氧化细菌（AOB）和硝化细菌等微生物丰度有显著影响，土壤中总氮、NO3−-N、NH4+-N的含量和反硝化基因丰度与生物炭施加量显著正相关[94 − 95]. 土壤中NH4+-N、NO3−-N、NO2−-N、N2O和NH3参与的主要反应过程如图3所示，NO2−-N在多个氮循环过程中都发挥中间底物的作用，且NO2－-N浓度与N2O产量的相关性最强，因此NO2−-N浓度的下降和N2O消耗量的上升是N2O减排的关键.

      土壤中NO2−-N的浓度受硝化反应、反硝化反应和DNRA反应共同调控. Venterea等[96]的实验结果指出，土壤硝化反应速率的提高促进NO2−-N转化为NO3−-N，导致N2O产量减少200%以上. 生物炭可能通过提高土壤pH值促进AOA和AOB的生长繁殖、吸附硝化抑制剂等途径提高土壤硝化反应速率[97]. AOA、AOB丰度的提高促进NH4+-N转化为NO2−-N，为NO2−-N转化为NO3−-N提供充足底物. 研究表明，土壤中NO2−-N浓度和N2O产量均与nxrA/amoA呈极强的负相关关系（皮尔逊相关系数分别为—0.79和—0.68）[98]，生物炭作为潜在碳源能提高土壤中nxrA基因的丰度，提高亚硝酸氧化还原酶活性，推动硝化反应的进程，促进NO2−-N转化为NO3−-N [99 − 100]. 近年来，有学者发现一条NH4+-N转化为NO3−-N的硝化反应新途径，完全氨氧化（Comammox）微生物可以将NH4+-N直接氧化为NO3−-N[101]，其在酸性土壤的硝化反应中发挥重要作用[102]. 该微生物的发现打破了硝化反应由NH4+-N氧化为NO2−-N再转化为NO3−-N的传统认知，间接减少了土壤中NO2−-N和N2O的产量. 现有研究指出，生物炭显著改变了土壤中硝化细菌的群落结构，提高了Comammox微生物丰度，Comammox微生物的丰度与土壤硝化速率呈正相关关系[103].

      生物炭可以直接参与电子转移过程，生物炭较高的芳香性会增加微生物与生物炭表面离域π电子系统相互作用的可能性，从而促进电子交换[104 − 105]. 因此Cayuela等[106]指出，生物炭提高土壤pH值不是减少N2O逸散量的唯一途径，其还可以促进电子向反硝化微生物转移，提高土壤反硝化速率. nirK、norB和nirS基因是主导NO2−-N转化为N2O的重要功能基因，nosZ基因是抑制N2O产生的关键基因. 现有研究指出，施加生物炭可以提高土壤中narG和nirK基因的丰度，推动反硝化反应的进程，促进N2O的产生[107]，而nosZ基因丰度的提高能加速N2O还原为N2 [108 − 110]. 因此，生物炭对土壤N2O逸散量的影响是动态的. 大量研究结果证实，施加生物炭的土壤中nosZ基因丰度的增幅显著高于nirK和nirS基因15.8%－182.9%，综合效应表现为土壤N2O逸散量降低14%－96%[111 − 113].

      DNRA是一种以NO2−-N作为中间产物，将NO3−-N还原为NH4+-N的生物过程，其在水体和土壤氮循环中发挥重要作用[114]，经由DNRA过程还原的NO3−-N总量约占总NO3−-N还原量的93%[115]. 生物炭能为土壤中DNRA过程提供电子受体和易被微生物所利用的碳源[78]，提高napAB、nrfA等DNRA反应相关基因的丰度约5%－89%[68, 116]，提升土壤DNRA反应速率约418%[116]. 推动DNRA反应进程不仅可以与反硝化过程竞争反应底物以减少N2O逸散量[117 − 118]，还可以将NO3−-N还原为植物更易吸收利用的NH4+-N[119].

      综上，降低土壤中NO2−-N的浓度和促进N2O转化为N2是生物炭减少土壤中N2O逸散的两个主要机理. 施加生物炭通过促进土壤中AOA、AOB、Comammox菌等微生物的富集、提高nxrA、napAB和nrfA等基因的丰度、提升亚硝酸氧化还原酶活性等途径降低土壤中NO2−-N浓度，抑制N2O的逸散. 当前关于生物炭施加对土壤中Comammox微生物和DNRA反应影响的研究较少，但其是硝化过程和反硝化过程外特殊的氮循环途径. Comammox微生物主导的氨氧化过程，会减少NO2−-N和N2O的产生[120]，降低环境污染风险和土壤氮流失量. Hu等[121]的研究显示在70%的土壤样品中，Comammox微生物占优势，其含有的amoA基因转录水平中位数超过了AOA和AOB，在原位土壤中，Comammox微生物的丰度与硝化潜力呈显著正相关. DNRA是陆地氮循环中几乎被遗忘的过程，它可以通过将易流失的NO3−-N转化为NH4+-N来保存氮，减少硝酸盐通过反硝化、淋溶和径流等途径的流失[122]. 未来应重视施加生物炭促进土壤Comammox和DNRA反应的机理研究，深入探讨生物炭对土壤氮循环的影响. 施加生物炭引起的反硝化速率提升极易造成土壤中氮素以N2O的形式流失，但生物炭能显著提高土壤中nosZ基因的丰度，促进反硝化反应产生的N2O转化为N2，调控农田土壤温室气体排放并降低农田土壤氮流失带来的危害. 但也有部分研究指出生物炭施加量过多导致土壤N2O逸散量增加17.3%－37.4% [123]，因此农田土壤生物炭施加量和生物炭对土壤N2O减排机制的影响值得进一步探究.

    3.   施加生物炭提高植物的氮素吸收能力（Improves nitrogen uptake in plants by biochar application）

    • 根是植物吸收养分和水分的主要器官，当植物根系发育良好时，施加生物炭的土壤中丰富的NH4+-N和NO3−-N将显著提高植物摄取氮素的能力[124]. 研究表明，生物炭改善土壤理化性质不仅可以抑制土壤氮素流失，还可以降低根系生长阻力，促进植物根系生长发育[125]. Xiang等[126]的meta分析结果表明，施加生物炭显著提高植物根生物量、根体积、根表面积、根长和根尖数量约32%、29%、39%、52%和17%. 为探究施加生物炭对植物生长的影响，以主题=“生物炭”并且“株高或生物量”为检索条件，对CNKI数据库中相关文献进行筛选、数据提取，共得到有效数据90个，分析结果如图4所示.

      向土壤中施加生物炭分别提高植物株高、生物量和作物产量约12%、24%和21%. 李婷等[127]研究证实，提高植物氮吸收能力可以降低土壤总氮流失量约30%－51%. 但部分研究指出，施加生物炭可能会导致土壤NH3逸散量激增和作物产量下降，例如Zhao等[128]实验结果证实，施加生物炭导致土壤中NH3的挥发量提高约292%；Calamai等[129]研究表明，生物炭施加可能会导致玉米生物量和产量分别降低约2%－6%和16%－20%. 高pH[130]、生物炭表面积和孔隙体积、孔结构分别通过增加NH4+-N向NH3转化，加快NH3扩散[131 − 132]，生物炭吸附酶反应底物，提高脲酶活性增加了土壤NH3的挥发 [133]，从而导致土壤中氮素流失. 另有研究表明由于土壤pH值和生物炭吸附位点的增加，生物炭施加量的增加也会增加NH3的产生[134]. 以高挥发性物质含量为特征的生物炭可能通过减少氮的吸收来影响土壤系统中的氮生化循环，从而对植物生长产生负面影响[135]. 生物炭可以通过抑制NO3−-N随水分入渗、促进土壤中硝化反应过程等途径提高植物根尖区土壤NO3−-N的浓度. 大多数植物更易吸收和利用土壤中的NO3−-N，植物氮吸收能力的提高不仅促进植物摄取更多NO3−-N，还可以进一步降低土壤溶液中NO3−-N的浓度并抑制反硝化反应. 此外，生物炭还在农业生产施肥过程中发挥“削峰填谷”的作用，其吸附土壤溶液中游离NH4+-N的能力有助于缓解作物生长受土壤中快速上升的NH4+-N浓度的抑制[136-137]. 因此施加生物炭显著提升植物的氮吸收能力不仅可以提高作物产量，还可以抑制NO3−-N的流失和N2O的逸散[138].

    4.   结论与展望（Conclusion and prospect）

    • 生物炭通过改良土壤理化性质、减少NH4+-N径流流失量和NO3−-N淋溶流失量、提高narG、nirK、nxrA、napAB、nrfA和nosZ等基因丰度、提升硝化反应、DNRA反应和N2O还原反应速率、促进植物摄取土壤中氮素等途径，达到抑制农田土壤氮素流失、提高作物产量和调控农田土壤温室气体排放的目的.

      近年，新型生物炭材料层出不穷，负载NH4+-N生物炭、改性生物炭、生物炭基肥等材料陆续用于减少农田土壤氮素流失. 相较于传统生物炭材料，负载NH4+-N生物炭和生物炭基肥抑制NH4+-N流失和NH3挥发、提升土壤氮素利用率的能力更强[139 − 142]；改性生物炭拥有更大的NH4+-N和NO3−-N吸附容量并能有效减少NO3−-N淋溶流失[143 − 144]. 因此，未来关于施加生物炭缓解农田土壤氮流失领域的研究建议从以下两个方向展开：（1）根据我国常见农业废弃物种类和主要土壤类型开展大田实验，因地制宜选择生物炭热解的原材料，进一步探讨生物炭热解条件、粒径、pH值、含碳量、施加量及施加方式等因素对农田土壤氮流失的影响；（2）减少温室气体N2O排放的Comammox反应、将NO3−-N转化为不易流失的NH4+-N的DNRA反应的发现改变了对传统氮素循环的认知，但目前关于生物炭对此过程研究较少，未来应深入研究生物炭材料调控土壤氮循环的机理，探究施加生物炭对土壤微生物群落结构和氮循环相关基因表达的影响，关注生物炭促进土壤中DNRA反应和Comammox反应对减少N2O逸散量的贡献，重视改性生物炭材料中金属离子对土壤中铁氨氧化反应（Feammox）、厌氧氨氧化反应（Anammox）等氮循环途径的促进作用.

    参考文献 (144)

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