摘要
近年来，随着氧化亚氮(nitrous oxide, N2O)在大气中浓度的逐年上升，微生物介导的N2O产生机制日益引起学界的关注。近期研究表明，好氧氨氧化微生物(aerobic ammonia oxidizing microorganisms, AOMs)参与的氨氧化过程是全球N2O的主要来源之一。本文从AOMs的种群分类、各类群的生态分布特点及影响其分布的环境因素、AOMs介导N2O产生的热点地区、AOMs产生N2O的途径及其影响因素进行归纳总结，并对未来的研究方向进行了展望。本综述有助于进一步理解AOMs类群及其产N2O机制。
好氧氨氧化是在好氧条件下将铵盐转化为亚硝酸盐或硝酸盐的过程，是氮循环的基本过程之一，对大多数陆地、海洋以及淡水生态系统的元素循环至关重
N2O是除二氧化碳和甲烷外最受关注的温室气体之一。自1750年以来，大气中的N2O浓度增长了近23
1 好氧氨氧化微生物类群
1.1 好氧氨氧化微生物的种群分类
Winogradsky是最早从环境中分离出氨氧化细菌的学者，他于1892年发布了亚硝化单胞菌(Nitrosomonas)和亚硝化球菌(Nitrosococcus) 2个新属，分别属于β变形菌纲(Betaproteobacteria)和γ变形菌纲(Gammaproteobacteria)；1933年，Winogradsky又发布了亚硝化螺菌(Nitrosospira)这一新属，其也属于β变形菌纲。为研究AOB的生物多样性并对新分离的未知AOB菌株进行分类鉴定，Head等发布了第一个基于16S rRNA基因序列的AOB类群系统发育树，此后还有学者利用AOB的其他功能基因进行建
与AOB相比，学者们对AOA种群的认识相对有限。2005年，第一株AOA从海洋中分离出来，当时被归类于泉古菌门(Crenarchaeota)，并被证实具有氨氧化作用，此后AOA的相关研究也备受学者关
学界一般认为硝化过程的两步反应分别由氨氧化微生物和亚硝酸盐氧化微生物(nitrite-oxidizing microbes, NOM)催化。然而，2015年，Daims
1.2 好氧氨氧化微生物的分布特征
本文选取了29条分离自各种不同生境的AOMs的16S rRNA基因进行分子系统学分析(
图1 好氧氨氧化菌基于16S rRNA基因序列的系统发育树
Figure 1 Phylogenetic evolutionary tree based on 16S rRNA gene sequences of aerobic ammonia oxidizing microorganisms
AOA的生态分布规律与AOB不同。例如，维也纳亚硝化球形菌(Nitrososphaera viennensis) EN76分离自花园土壤，Nitrosarchaeum koreense MY1分离自农业土壤，Nitrosopumilus zosterae NM25分离自海岸带沉积物；同样分离自土壤的Nitrososphaera viennensis具有三者中最高的最适温度(42 ℃)，Nitrosarchaeum koreense具有高达10 mmol/L的铵耐受能力，Nitrosopumilus zosterae则有着最宽泛的盐度耐受能
上述AOA与AOB的生态分布特征主要基于其分离环境进行总结归纳。尽管目前Comammox的纯培养物还很有限，但宏基因组测序技术以及Comammox所携带的标记基因PCR使其在各种环境中的检测成为可
1.3 影响好氧氨氧化微生物分布的环境因素
影响AOMs分布的环境因素众多，主要包括溶解氧(dissolved oxygen, DO)、pH值、微量元素、铵浓度与水文异质性、氨亲和力等。
一般认为AOA比AOB更适应缺氧生态位。例如，在一项对热带东北太平洋氧限制区(DO<2 μmol/L)的调查研究中，发现主要的氨氧化类群是AOA而非AO
大部分AOMs是嗜中性微生物，但学者对一些酸性环境(包括农业土壤、森林土壤、火山土壤等)和碱性环境(包括盐碱地、碱性湖泊等)的调查发现，仍有可以生存于酸性或碱性环境的AOM
AOMs在大洋中的分布还会受到游离的铁离子和铜离子的影响。游离的铁离子可能使AOB分布在较浅层的透光水域，那里铵供应较高，但也存在对铁离子的竞争；而AOA则对海洋中游离的铜离子有更强的亲和力和更高的耐受
铵浓度与水文异质性同样影响着AOMs的分布。在一项对地下含水层的调查中，发现以Nitrosopumilus主导的AOA类群分布在地下水补给区(0.1 mg/L NH-N)，而以Nitrosospira主导的AOB则分布在地下水排泄区(3.8 mg/L NH-N
在最近一项对4组AOA类群的氨亲和力进行定量的研究中发现，AOA对氨的亲和力比之前学界所认为的更宽泛，甚至在某些情况下与非寡营养AOB的氨亲和力重叠，这可能会改变原有AOMs生态位分化的认
2 好氧氨氧化微生物介导N2O的产生
2.1 产生N2O的热点区域
随着大气中N2O浓度的逐年上升，不少N2O产生的热点区域被发现，这些热点区域包括各种类型的土
| Hotspots | Samplinng environments | N2O flux or concentration | Reference |
|---|---|---|---|
| Various types of soil | Alkaline, neutral purple soil | 103.71 ng/(g·d) |
[ |
| Oil palm soil | 408.57 ng/(g·h) |
[ | |
| Forest soil | 18.86 ng/(g·h) |
[ | |
| Agricultural soil |
226.60 μg/( |
[ | |
| Marine | The eastern tropical south Pacific | 13.73 ng/(L·d) |
[ |
| The eastern tropical north Pacific | 13.20 ng/d |
[ | |
| Various freshwater systems | River |
580.80 μg/( |
[ |
| Stream |
264.00 μg/( |
[ | |
| Lake |
69.41 g/( |
[ | |
| Sewage treatment system | 7.00 μg/(g·min) |
[ | |
| Various types of sediments | Plateau wetland sediment | 50.29 ng/(g·d) |
[ |
| Thawing Yedoma permafrost |
1.72 mg/( |
[ | |
| Estuarine sediment | 9.24 ng/(g·h) |
[ |
2.2 产生N2O的途径
目前发现AOB产生N2O的途径主要有2类(
图2 好氧氨氧化菌N2O产生途径示意图
Figure 2 Schematic diagram of N2O production pathway of aerobic ammonia oxidizing microorganisms.A: Three pathways of AOB-mediated N2O production; B: Two hypotheses for AOA-mediated N2O production. AMO: Ammonia monooxygenase; HAO: Hydroxylamine oxidoreductase; NOO: Nitric oxide oxidase; NIR: Nitrite reductase; NOR: Nitric oxide reductase; Cyt P460: Cytochrome P460; NXOR: Nitroxyl oxidoreductase; Cu-HAO: Copper hydroxylamine oxidoreductase; Cu-NIR: Copper nitrite reductase.
有学者认为AOB可以进行硝化反硝化，是因为其具有与经典反硝化菌同源的2种酶：一是含铜亚硝酸盐还原酶(nitrite reductase, NIR)，二是一氧化氮还原酶(NOR)。其中，NOR被认为在NO还原为N2O过程中是必需的，而NIR被认为可能参与了羟胺的氧化而非亚硝酸盐还原，因为有研究证明在缺乏NIR的AOB中仍能检测到N2O的产
与AOB类似，AOA的第一步反应也是铵根或氨在氨单加氧酶的作用下发生转化，但是中间产物可能发生变化，证据有：(1) 在AOA的基因组中缺乏可以直接催化羟胺发生反应的酶的基因；(2) 在海洋AOA极其活跃的区域，有大量的N2O产生；(3) 有研究检测到了AOA的中间产物包括N
在对农业溪流以及碱性耕地N2O产生的主要贡献者的野外调查中，Ta
2.3 影响产生N2O的因素
随着从野外调查到室内研究的不断深入，众多因素被发现可以影响AOMs的N2O产生，主要包括溶解氧、酸碱度、温度和底物浓度。
溶解氧、酸碱度和底物浓度均可通过控制代谢途径或影响某一途径的酶活性来改变AOMs产生N2O的量。在多项溶解氧对N2O产生影响的研究中发现，低氧或氧限制条件(DO<1 mg/L)会使AOMs通过硝化反硝化途径产生N2O的量增加，但随着DO的增加(DO>2 mg/L)，AOMs通过羟胺氧化途径产生N2O的比例也会增
同时，溶解氧、酸碱度和温度还可以通过改变体系的群落结构，进而影响AOMs对N2O的产生。在高铵废水和硝化活性污泥的研究中分别发现，高氧条件[DO=(3.75±0.49) mg/L]会使Nitrosomonas这一类群占比增加33.43%，导致亚硝酸盐累积，为N2O产生创造了前体物
3 研究不足与展望
近年来，关于AOMs类群及其N2O产生机制的研究，学界已取得诸多重要进展。然而，随着科学技术的不断革新，更多机遇和挑战主要体现在以下几个方面。
目前，基于高通量测序技术从更多生境中挖掘出了AOMs，但由于AOMs培养的难度，被纯培养的物种还很有限，亟待加强对AOMs菌株的分离、培养及其功能验证。将流式细胞
随着科学技术的发展，将(宏)基因组、(宏)转录组、(宏)蛋白组、代谢组等多组学技术与稳定同位素核酸探针技术(DNA-based stable isotope probing, DNA-SIP
厘清不同生境下各种因素对AOMs类群产生N2O的影响，并准确评估其贡献度，可以为农业生产、生活污水处理、畜禽养殖废水处理等与人类生产生活息息相关的行业，提供温室气体减排以及微生物调控技术更加科学和有效的指导。
作者贡献声明
马睿：论文撰写并修改，图表制作；王亚琦：论文撰写并修改；王和林：论文修改与校稿；李平：文章选题设计、指导撰写与修改、课题支持。
利益冲突
作者声明绝无任何可能会影响本文所报告工作的已知经济利益或个人关系。
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