微生物学报

Origin	Strains	Gram
G⁺: Gram-positive; G⁻: Gram-negative; − : Fungi without Gram-differentiation.
Wastewater	Coking wastewater	Cupriavidus sp. S1^[11]	G⁻
Wastewater treatment plants	Ochrobactrum anthropi HND19^[12]	G⁻
Aquaculture wastewater	Pseudomonas mendocina S16^[13]	G⁻
Pig farm wastewater	Acinetobacter sp. T1^[14]	G⁻
Eutrophic lake	Pseudomonas sp. N31942^[15]	G⁻
The Songhua River	Microbacterium esteraromaticum SFA13^[16]	G⁺
Artificial reservoir	Acinetobacter sp. H36^[17]	G⁻
Activated sludge	Acinetobacter calcoaceticus TY1^[18]	G⁻
Soil	Paddy field soil	Arthrobacter arilaitensis Y10^[19]	G⁺
Soil	Pseudomonas sp. PHR6^[20]	G⁻
Soil	Basidiomycetes^[21]	−
Soil	Penicillium^[22]	−
Paddy soil	Pseudomonas tolaasii Y-11^[23]	G⁻
Soil	Bacillus cereus PK5^[24]	G⁺
Soil	Bacillus subtilis PK15^[24]	G⁺

Origin	Strains	Gram
G⁺: Gram-positive; G⁻: Gram-negative; − : Fungi without Gram-differentiation.
Wastewater	Coking wastewater	Cupriavidus sp. S1^[11]	G⁻
Wastewater treatment plants	Ochrobactrum anthropi HND19^[12]	G⁻
Aquaculture wastewater	Pseudomonas mendocina S16^[13]	G⁻
Pig farm wastewater	Acinetobacter sp. T1^[14]	G⁻
Eutrophic lake	Pseudomonas sp. N31942^[15]	G⁻
The Songhua River	Microbacterium esteraromaticum SFA13^[16]	G⁺
Artificial reservoir	Acinetobacter sp. H36^[17]	G⁻
Activated sludge	Acinetobacter calcoaceticus TY1^[18]	G⁻
Soil	Paddy field soil	Arthrobacter arilaitensis Y10^[19]	G⁺
Soil	Pseudomonas sp. PHR6^[20]	G⁻
Soil	Basidiomycetes^[21]	−
Soil	Penicillium^[22]	−
Paddy soil	Pseudomonas tolaasii Y-11^[23]	G⁻
Soil	Bacillus cereus PK5^[24]	G⁺
Soil	Bacillus subtilis PK15^[24]	G⁺

Number	Metabolic pathway	References
1	NH₄⁺-N→NO₂^–-N→NO₃⁻-N→NO₂⁻-N→NO→N₂O→N₂	[4]
2	NH₄⁺-N→NH₂OH→NO₂⁻-N→NO₃⁻-N→NO₂⁻-N→N₂	[48]
3	NH₄⁺-N→NH₂OH→NO₂⁻-N→N₂O→N₂	[49]
4	NH₄⁺-N→NH₂OH→NO₂⁻-N→NO₃⁻-N→N₂O→N₂	[50]
5	NH₄⁺-N→NO→N₂O	[51]
6	NH₄⁺-N→N₂O→N₂	[52]

Number	Metabolic pathway	References
1	NH₄⁺-N→NO₂^–-N→NO₃⁻-N→NO₂⁻-N→NO→N₂O→N₂	[4]
2	NH₄⁺-N→NH₂OH→NO₂⁻-N→NO₃⁻-N→NO₂⁻-N→N₂	[48]
3	NH₄⁺-N→NH₂OH→NO₂⁻-N→N₂O→N₂	[49]
4	NH₄⁺-N→NH₂OH→NO₂⁻-N→NO₃⁻-N→N₂O→N₂	[50]
5	NH₄⁺-N→NO→N₂O	[51]
6	NH₄⁺-N→N₂O→N₂	[52]

Strain	T/℃	NH₄⁺-N removal	NO₃⁻-N removal
−: It is not mentioned in the literature.
Acinetobacter sp. TAC-1^[4]	5	94.6%	40%
Microbacterium sp. SFA13^[71]	5	1.6 mg/(L·h)	0.24 mg/(L·h)
Pseudomonas sp. M-11^[66]	10	98%	−
Pseudomonas putida Y-9^[5]	15	99.36%	82%
Acinetobacter sp. HA2^[37]	10	100%	−
Pseudomonas migulae AN-1^[72]	10	1.6 mg/(L·h)	1.5 mg/(L·h)
Acinetobacter TY1^[73]	8	97.4%	81.85%
Acinetobacter sp. Z1^[74]	10	96%	97.5%
Chelatococcus daeguensis TAD1^[63]	50	100%	100%
Anoxybacillus contaminans HA^[64]	55	71%	74.7%

Strain	T/℃	NH₄⁺-N removal	NO₃⁻-N removal
−: It is not mentioned in the literature.
Acinetobacter sp. TAC-1^[4]	5	94.6%	40%
Microbacterium sp. SFA13^[71]	5	1.6 mg/(L·h)	0.24 mg/(L·h)
Pseudomonas sp. M-11^[66]	10	98%	−
Pseudomonas putida Y-9^[5]	15	99.36%	82%
Acinetobacter sp. HA2^[37]	10	100%	−
Pseudomonas migulae AN-1^[72]	10	1.6 mg/(L·h)	1.5 mg/(L·h)
Acinetobacter TY1^[73]	8	97.4%	81.85%
Acinetobacter sp. Z1^[74]	10	96%	97.5%
Chelatococcus daeguensis TAD1^[63]	50	100%	100%
Anoxybacillus contaminans HA^[64]	55	71%	74.7%

HNADMs	Wastewater type	Scale
Chelatococcus daeguensis TAD1^[63]	Synthetic wastewater	Lab
Zobellella B307^[106]	Mariculture wastewater	Lab
Acinetobacter sp. TN-14^[109]	Pig farm wastewater	Lab
Pseudomonas sp. HJ3^[110]	Coal gasification wastewater	Lab
HNADMs flora^[111]	Livestock wastewater	Lab
Bacillus subtilis GHSP10^[112]	Aquaculture sewage	Lab
Acinetobacter sp. Z1^[75]	Wastewater treatment plants	Lab
pseudomonas sp. DM02^[76]	Aquaculture wastewater	Lab

HNADMs	Wastewater type	Scale
Chelatococcus daeguensis TAD1^[63]	Synthetic wastewater	Lab
Zobellella B307^[106]	Mariculture wastewater	Lab
Acinetobacter sp. TN-14^[109]	Pig farm wastewater	Lab
Pseudomonas sp. HJ3^[110]	Coal gasification wastewater	Lab
HNADMs flora^[111]	Livestock wastewater	Lab
Bacillus subtilis GHSP10^[112]	Aquaculture sewage	Lab
Acinetobacter sp. Z1^[75]	Wastewater treatment plants	Lab
pseudomonas sp. DM02^[76]	Aquaculture wastewater	Lab

极端环境中异养硝化-好氧反硝化微生物的研究进展

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吕鹏翼 ¹ , 李小曼 ² , 罗晓 ¹^,^* , 刘云琦 ² , 林冬庆 ²

微生物学报 | 综述 2024,64(12): 4681-4700

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微生物学报 | 综述 2024, 64(12): 4681-4700

极端环境中异养硝化-好氧反硝化微生物的研究进展

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吕鹏翼¹, 李小曼², 罗晓¹^,^*, 刘云琦², 林冬庆²

作者信息

1 河北科技大学环境科学与工程学院, 河北省污染防治生物技术重点实验室, 河北石家庄 050018

2 河北科技大学建筑工程学院, 河北石家庄 050018

Heterotrophic nitrifying and aerobic denitrifying microorganisms in extreme environments

Pengyi LYU¹, Xiaoman LI², Xiao LUO¹^,^*, Yunqi LIU², Dongqing LIN²

Affiliations

1 Hebei Key Laboratory of Pollution Prevention Biotechnology, College of Environmental Science and Engineering, Hebei University of Science and Technology, Shijiazhuang 050018, Hebei, China

2 School of Civil Engineering, Hebei University of Science and Technology, Shijiazhuang 050018, Hebei, China

出版时间: 2024-09-23 doi: 10.13343/j.cnki.wsxb.20240395

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摘要

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异养硝化-好氧反硝化微生物(heterotrophic nitrification and aerobic denitrification microorganisms, HNADMs)在有氧条件下可以同时完成硝化和反硝化过程，为生物脱氮技术开发提供了新思路。研究发现部分HNADMs在极端环境下仍具有良好的脱氮效能，其在非典型污水脱氮领域表现出较高的应用价值，然而，当前关于极端环境中HNADMs的研究仍处于起步阶段。本文系统概述了HNADMs的多样性、生理生化特性和代谢路径的复杂性；重点对具有极端温度、pH、高盐、寡营养、重金属耐受性，以及同步脱氮除磷、降解复杂有机物、抗生素耐药性HNADMs的脱氮效能和耐受机理进行了综述；同时对极端环境耐受HNADMs的应用现状进行了总结；并基于当前极端环境耐受HNADMs研究中的瓶颈问题进行了展望，以期为HNADMs在复杂污水处理中的应用提供基础资料。

关键词

异养硝化-好氧反硝化 / 极端环境 / 脱氮效能 / 耐受机理 / 应用现状

Abstract

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Heterotrophic nitrification and aerobic denitrification microorganisms (HNADMs) can complete both nitrification and denitrification under aerobic conditions, which paves a new way for biological nitrogen removal. Researchers have discovered that some HNADMs maintain high nitrogen removal efficiency in extreme environments, exhibiting high application potential in the nitrogen removal of atypical wastewater. However, the available studies on HNADMs in extreme conditions remain in the initial stage. This article introduces the diversity, physiological and biochemical characteristics, and the complexity of metabolic pathways of HNADMs. Particularly, it reviews the research progress in HNADMs regarding the tolerance mechanisms to extreme temperature, pH, high salt, oligotrophic, and heavy metal stress conditions, nitrogen removal efficiency of simultaneous nitrogen and phosphorus removal, degradation of complex organic matter, and antibiotic resistance. Furthermore, the application status of HNADMs in extreme environments is summarized. Finally, the bottlenecks in the application of HNADMs in extreme environments are prospected. The review is expected to provide basic data for the application of HNADMs in complex wastewater treatment.

Key words

heterotrophic nitrification and aerobic denitrification / extreme environments / nitrogen removal efficiency / tolerance mechanism / application status

引用本文

吕鹏翼, 李小曼, 罗晓, 刘云琦, 林冬庆. 极端环境中异养硝化-好氧反硝化微生物的研究进展. 微生物学报, 2024 , 64 (12) : 4681 -4700 . DOI: 10.13343/j.cnki.wsxb.20240395

Pengyi LYU, Xiaoman LI, Xiao LUO, Yunqi LIU, Dongqing LIN. Heterotrophic nitrifying and aerobic denitrifying microorganisms in extreme environments[J]. Acta Microbiologica Sinica, 2024 , 64 (12) : 4681 -4700 . DOI: 10.13343/j.cnki.wsxb.20240395

正文

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传统生物脱氮(biological nitrogen removal, BNR)是当前国内外污水处理厂(wastewater treatment plants, WWTPs)主流的污水脱氮工艺。鉴于硝化过程与反硝化过程对溶解氧(dissolved oxygen, DO)及有机物需求的差异性，WWTPs需要将两者有机结合实现完整的生物脱氮。硝化过程主要由氨氧化细菌(ammonia-oxidizing bacteria, AOB)和亚硝酸盐氧化菌(nitrite-oxidizing bacteria, NOB)驱动完成，此外，研究表明氨氧化古菌在污水处理系统中的作用同样不可小觑^[1]。由于AOB及NOB自养菌的代谢属性，其缓慢的生长速率使硝化过程成为BNR的限速步骤。传统的反硝化过程主要由异养反硝化微生物在厌氧/缺氧条件下将NO₃⁻-N/NO₂⁻-N还原为N₂。传统的BNR基于其效果显著、操作简单及技术成熟等优点在含氮废水的处理中应用最为广泛，然而在WWTPs迫切需要“提质增效”和“高效低耗”的大背景下，其占地面积大、设备和运营成本较高、硝化菌世代时间长以及对极端环境[高氨、高化学需氧量(chemical oxygen demand, COD)、高盐、极端温度、极端pH]的耐受性差等缺点亟待解决。

近年来，越来越多的氮代谢过程被发现，如厌氧氨氧化(anammox)、短程硝化-反硝化(short-cut nitrification denitrification)、全程硝化-反硝化(complete nitrification and denitrification)、硝酸盐异化还原成氨(dissimilatory nitrate reduction to ammonia, DNRA)、异养硝化-好氧反硝化(heterotrophic nitrification and aerobic denitrification, HNAD)等，其中，HNAD基于其在污水处理系统中的普遍存在性以及不可忽略的脱氮潜能日益被关注。与自养硝化菌和缺氧反硝化细菌相比，异养硝化-好氧反硝化微生物(heterotrophic nitrification and aerobic denitrification microorganisms, HNADMs)生长代谢快，有氧条件下可同时完成硝化和反硝化过程，并实现同步去除有机物等^[2]，具有环境适应能力强和耐受多种污染物胁迫等优势。当前，关于HNADMs的研究主要聚焦于高效菌株的筛选、最优作用参数(碳源、碳氮比、DO浓度、温度及pH值等)的确定、应用模式的研究以及代谢机制的探究等。随着研究的深入，部分HNADMs表现出了较强的极端环境耐受性，如耐低温菌株谷氨酸杆菌(Glutamicibacter sp.) WS1^[3]、不动杆菌属(Acinetobacter sp.) TAC-1^[4]和恶臭假单胞菌(Pseudomonas putida) Y-9^[5]等在5−15 ℃条件下对NO₃⁻-N的去除率均可达到80%以上；耐极端pH菌株Acinetobacter sp. JR1^[6]在pH为4.5−10.0时，其总氮(total nitrogen, TN)去除率均能达到96%以上；菌株巨大芽孢杆菌(Bacillus megatherium) N07^[7]在NaCl为8.00%的高盐环境中对NO₃⁻-N (初始浓度150 mg/L)的去除率可达到94.94%；Pseudomonas putida ZN1^[8]对重金属Cu²⁺ (浓度800 mg/L)具有较强耐受性；节杆菌属(Arthrobacter sp.) HHEP5^[9]可去除海水养殖废水中99%的PO₄³⁻-P和95%的TN。

耐受极端环境的HNADMs在极端环境含氮废水的处理中表现出了较高的应用潜能。然而，目前针对耐受极端环境的HNADMs的研究仍处于起步阶段。本文系统概述了HNADMs的多样性、生态分布特征及其生理生化特征，总结了其代谢路径，重点综述了耐受极端环境的HNADMs的类型及其研究现状，讨论了其应用模式并对其前景进行了展望。

1 HNADMs的研究概述

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1.1 HNADMs多样性

自1983年从活性污泥中分离鉴定出第1株HNAD菌泛养硫球菌(Thiosphaera pantotropha)^[10] (现更名为Paracoccus denitrifican)以来，鉴于HNADMs独特的脱氮方式，近40年来引起了全球研究人员的极大关注。利用Web of Science、Google Scholar和ScienceDirect平台对自1983年以来报道的185株HNAD菌株进行统计，分析发现HNADMs广泛存在于细菌域中，少数为真菌及其他菌属，占比分别为79.1%、18.6%和2.3% (图1)；细菌中变形菌门(Proteobacteria)为绝对优势菌属，占比为76.5%，其后依次为放线菌门(Actinobacteria)、拟杆菌门(Bacteroidetes)和厚壁菌门(Firmicutes)，占比分别为14.7%、5.9%和2.9%；真菌中的优势菌门主要有子囊菌门(Ascomycota)、真菌门(Eumycophyta)及担子菌(Basidiomycota)，占比分别为50.0%、33.3%和16.7%；变形菌门在纲水平的优势纲主要为γ-变形菌纲(59.26%)、β-变形菌纲(22.2%)以及α-变形菌纲(18.15%)。将绝对优势的变形菌门在属水平进一步统计分析(图1)，当前分离株主要集中于假单胞菌属(Pseudomonas) (26.7%)、不动杆菌属(Acinetobacter) (24.7%)、芽孢杆菌属(Bacillus) (13.7%)和盐单胞菌属(Halomonas) (5.5%)。

氮元素作为最主要的生命元素及环境元素在自然界中无处不在，由微生物驱动的氮循环是地球生物化学循环的重要组成部分，由微生物主导的脱氮过程是氮循环中不可或缺的重要环节。研究表明，HNADMs在水生生态系统、陆地生态系统及大气生态系统中均有分布，尤其WWTPs、水库、河流、海水等水生系统为HNADMs的主要生态位，不同类型水体及土壤中HNADMs的分布情况见表1，明确HNADMs在上述不同类型水体中的分布情况、在氮循环中的贡献及作用是当前的研究热点。在土壤环境中HNAD过程主要由真菌驱动，常见于担子菌(Basidiomycetes)^[21]、曲霉菌(Aspergillus)^[21]、青霉菌(Penicillium)^[22]等。大量研究^[25-26]报道，鉴于真菌比细菌具有更好的抗酸性和异养硝化性能，它们通常在氧化土壤中有机氮和铵态氮方面发挥重要作用。

1.2 生理生化特性

HNADMs种类繁多，多数为革兰氏阴性菌(表1)，如Pseudomonas^[27]、粪产碱菌(Alcaligenes faecalis)^[28]、无色杆菌属(Achromobacter)^[29]等；少数为革兰氏阳性菌，例如枯草芽孢杆菌(Bacillus subtilis)^[26]、甲基营养芽孢杆菌(Bacillus methylotrophicus)^[30]等。研究表明，碳源、C/N、温度、pH和DO是影响HNAD细菌生长和代谢的主要因素。(1) HNADMs为异养型，在脱氮过程中能够以有机碳为基质用于细胞骨架的合成以及电子供体，进而实现有机物的同步去除。HNADMs可以利用多种碳源作为反应底物，例如乙酸钠、琥珀酸钠、柠檬酸钠、草酸、蔗糖和葡萄糖等，不同HNAD菌株的最适碳源类型不尽相同，如以柠檬酸钠为碳源时，食聚羟基丁酸酯有益杆菌(Diaphorobacter polyhydroxybutyrativorans) SL-20^[31]的NH₄⁺-N去除效率仅为2.9%，显著低于碳源为乙酸钠(90.8%)、琥珀酸钠(79.8%)和聚(3-羟基丁酸酯-co-3-羟基戊酸酯) [poly (3-hydroxybutyrate-co-3-hydroxyvalerate), PHBV] (90.5%)。较宽的基质利用范围使其在不同类型有机物废水中具有很高的竞争力，成为维持反硝化反应稳定进程的优势菌株。不同的HNADMs对特定的环境表现出不同的适应性。(2) 大多数HNADMs的C/N作用范围为8−15，然而，部分HNADMs在寡营养条件下表现出很强的生存及脱氮能力，如Acinetobacter sp. Y16^[32]、Pseudomonas sp. M-33^[33]等；除此之外，一些耐高COD浓度的HNAD菌株如Pseudomonas aeruginosa KDQ4^[34]等也被分离。(3) 温度会影响酶的活性和细胞膜的特性，大多数HNADMs的最适温度范围为25−35 ℃，如施氏假单胞菌(Pseudomonas stutzeri) ADP-19^[35]和魔鬼弧菌(Vibrio diabolicus) SF16^[36]等。近年来，越来越多的耐低温菌株陆续被发现，例如Acinetobacter sp. HA2^[37]、Acinetobacter sp. Y16^[32]等。(4) pH主要通过影响细胞表面电荷和酶活性影响微生物代谢，大多数菌株最适pH范围为5.0−10.0，代表菌株如Bacillus methylotrophicus strain L7^[38]、红球菌属(Rhodococcus sp.) CPZ24^[39]和Pseudomonas sp. N31942^[19]等。随着HNADMs的关注度日益提高，一些在极端环境中能够生存同时具有较高脱氮性能的HNADMs菌株，如具备耐低(高)温、pH、高盐、寡营养、重金属以及同步脱氮除磷、去除难降解有机物及微塑料等的菌株陆续被分离筛选。

1.3 氮代谢路径

代谢过程的复杂性和氮代谢路径的多样性是HNAD研究中的又一热点和难点。当前关于HNADMs氮代谢路径的构建多是基于PCR技术对功能基因的定性结果、氮平衡理论的化学计量学方法或功能基因抑制剂的添加效果来进行的，具有一定局限性。大量研究表明，不同的HNADMs具有不同的氮代谢路径，例如能够执行完全硝化反硝化过程^[40]的菌株Pseudomonas aeruginosa^[41]、蜡样芽胞杆菌(Bacillus cereus) GS-5^[42]和Arthrobacter sp. HHEP5^[9]具备完整的硝化及反硝化酶系，当以NH₄⁺-N为氮源时，菌株能够利用氨单加氧酶(ammonia monooxygenase, AMO)、羟胺氧化酶(hydroxylamine oxidase, HAO)及亚硝酸盐氧化还原酶(nitrite oxidoreductase, NXR)依次将NH₄⁺-N氧化为NH₂OH、NO₂⁻-N和NO₃⁻-N，硝化过程的终产物NO₃⁻-N在反硝化酶周质硝酸盐还原酶(nitrate reductase, NAP)、亚硝酸盐还原酶(nitrite reductase, NIR)、一氧化氮还原酶(nitric oxide reductase, NOR)及一氧化二氮还原酶(nitrous oxide reductase, NOS)的催化下最终还原生成N₂释放(图2中Pathway 1)；执行羟胺氧化途径^[43] (图2中Pathway 2)氮代谢途径的菌株Halomonas sp. DN3^[44]、Alcaligenes faecalis strain NR^[45]和Alcaligenes faecalis TF-1^[46]等能够利用AMO将NH₄⁺-N氧化为NH₂OH，随后转化为NO，在反硝化酶NOR和NOS的催化下还原生成N₂ (图2中Pathway 2)，该路径未检测到NO₂⁻-N与NO₃⁻-N的相互转化过程，一定程度上提高了氮转化效率；部分HNAD菌株不具有完全的HNAD代谢途径，例如通过基因组学分析发现Pseudomonas sp. N31942^[19]不具备通过反硝化作用生成气态氮的能力，如图2中Pathway 3所示，当以NH₄⁺-N为氮源时，其代谢路径为NH₄⁺-N→NH₂OH→NO₂⁻-N→NO₃⁻- N→NO₂⁻-N→NH₄⁺-N→Glutamate，菌株首先利用AMO、HAO及NXR依次将NH₄⁺-N氧化为NH₂OH、NO₂⁻-N和NO₃⁻-N，生成的NO₃⁻-N通过硝酸盐还原酶(nitratereductase, NR)转化为NO₂⁻-N，还原产物NO₂⁻-N被NIR转化为NH₄⁺-N，最终，新生成的NH₄⁺-N被谷氨酸还原酶(glutamate dehydrogenase, GDH)转化为谷氨酸用"于细胞生长，在该氮代谢路径中同化作用是NH₄⁺-N代谢的最主要途径。此外，HNAD过程中可能存在的其他氮代谢路径详见表2。

同一菌株在不同的条件下可能驱动不同的氮代谢过程。研究表明，不同氮源类型下菌株会选择不同的氮代谢过程，NO₂⁻-N或NO₃⁻-N的存在会降低菌株的同化率，例如将脱氮副球菌(Paracoccus denitrificans) HY-1^[53]的氮源由NH₄⁺-N更换为NO₃⁻-N或NO₂⁻-N时，同化率从80.7%分别降至28.91%和30%；当以NH₄⁺-N为唯一氮源时，根瘤菌(Rhizobium sp.) WS7^[54]通过羟胺还原途径代谢NH₄⁺-N，以NO₃⁻-N或NO₂⁻-N为氮源时，菌株执行硝酸盐和亚硝酸盐反硝化途径；Pseudomonas putida Y-9^[5]以NO₃⁻-N为唯一氮源时，通过同化以及DNRA代谢NO₃⁻-N，以NH₄⁺-N及NO₃⁻-N为混合氮源时，硝化、DNRA和反硝化共存；由此可知，对于HNAD过程而言，“异养硝化”环节既可能发生硝化反应，也可能将外源氮源合成自身生物体氮。不同HNADMs发生好氧反硝化的DO不尽相同：Pseudomonas sp. G16^[55]的反硝化能力随DO升高而增强，在0 r/min时NO₃⁻-N去除效果仅为10%，NH₄⁺-N去除率为98.26%，当摇床转速提高到150 r/min或更高时，G16对NH₄⁺-N和NO₃⁻-N去除效率均趋于100%；Pseudomonas sp. AND-42^[56]的最佳DO为3.0 mg/L，过高或过低的DO浓度均会导致NH₄⁺-N去除率降低；当将振荡速度从160 r/min降低到50 r/min时，Pseudomonas stutzeri T13^[57]对TN的去除效率从23.47%显著提高到99.21%；因此，严格来说，“好氧反硝化”更准确地表述应为“有氧反硝化”。越来越多的研究证实，HNADMs为兼性厌氧菌，Thiosphaera pantotropha在厌氧和有氧条件分别通过催化膜结合硝酸盐还原酶(respiratory nitrate reuctase, NAR)和周质硝酸盐还原酶NAP进行硝酸盐还原^[58]；Huan等^[59]利用全基因组测序证实菌株Paracoccus denitrificans HY-1同时具有napA (周质硝酸盐还原酶NAP编码基因)及narG (编码膜结合硝酸盐还原酶NAR)，并在有氧和厌氧条件下均具有良好的反硝化能力；由此推测大部分HNADMs在不同的DO环境下可通过驱动不同的酶系进行硝酸盐还原过程。

上述研究表明，HNADMs的氮代谢途径是高度动态且受环境条件调控的。氮源类型及DO是驱动HNADMs选择氮代谢路径的关键环境因子。代谢过程的多样性及灵活性使得HNADMs能够在复杂氮源环境中繁殖并进行脱氮，该特性也导致了HNADMs代谢机理研究的复杂性。当前基于氮源利用、氮平衡来推测HNAD途径的方法有一定的局限性，随着分子生物学技术的发展，基因组学、转录组学、代谢组学和蛋白质组学在HNAD代谢路径的研究中提供了更多可能性，同时同位素法在氮代谢路径的构建中提高产物的辨识度。

2 极端环境中的HNADMs

收起

近年来，一些耐受极端环境的HNADMs如耐低温/高温、耐酸/碱、耐盐、耐重金属、同步脱氮除磷菌株等陆续被分离筛选，以上特殊优势使HNADMs在非典型废水BNR领域的应用展现出了巨大的潜力。

2.1 极端温度耐受HNADMs

温度是影响微生物新陈代谢的最主要因素之一。温度通过影响酶活性、基因的表达影响其生长及代谢，高温会导致核酸蛋白质组分变性，破坏菌体生理结构，而低温会抑制微生物酶的活性^[60-61]，降低微生物世代时间。大多数HNADMs最佳的生长及脱氮温度范围为25−37 ℃^[62]，然而部分HNADMs对温度表现出较高耐受性：如耐高温菌株大邱螯合球菌(Chelatococcus daeguensis) TAD1^[63]，将其与生物滤池相结合，在最优运行参数下50 ℃时其对NH₄⁺-N及TN的去除效率均达到了100%；55 ℃时污染无氧芽孢杆菌(Anoxybacillus contaminans) HA^[64]对NH₄⁺-N和NO₃⁻-N的去除率分别为71.00%和74.70%；大多数研究将其归因于高温下与HNAD过程相关的酶活性的增加^[65]。同时，越来越多的耐低温菌陆续被发现，耐低温菌株Acinetobacter sp. Y16在2 ℃下对NH₄⁺-N及NO₂⁻-N的去除率仍可达到66.00%和56.73%；杨墨等^[33]从冬季松花江底泥中分离出能够在10 ℃条件下实现高效脱氮的菌株Pseudomonas sp. M-33，该菌株在中性偏碱性的条件对NH₄⁺-N的去除率高达95%；2 ℃条件下菌株Pseudomonas sp. M-11^[66]产生低温应激反应并合成能维持其低温反应的特有酶，如冷激蛋白。上述耐低温菌株在低温下可正常生长同时可实现高效脱氮，关于其耐冷机制的解释总结如下：(1) 细胞膜脂肪酸的调控机制^[67]，Acinetobacter sp. TAC-1^[4]的膜转运蛋白和不饱和脂肪酸脱氢酶相关基因在5 ℃时显著上调，使其能够在低温下存活；(2) 内含抗冻蛋白和耐低温酶的编码基因^[68]，例如拉恩氏菌属(Rahnella sp.) R3产生β-半乳糖苷酶，使其在4 ℃时依然能保持最适宜温度下的80%的酶活性^[69]；(3) 冷休克蛋白(cold shock proteins, CSPs)^[70]的表达在低温下显著上调。然而，当前相关功能基因的类型尚不明确，未来可以通过基因敲除和全基因组分析等生信手段确定。更多极端环境耐受HNADMs详见表3。以上研究表明，部分耐极端温度的HNAD功能菌株有相对较广的温度适应范围，在低温或高温胁迫下均表现出了优秀的脱氮效能，该类微生物在冬季低温废水(10−15 ℃^[75])及工业废水(高于40 ℃)等污水处理领域中具有广泛的应用前景。

2.2 极端pH耐受HNADMs

脱氮微生物是一类pH敏感菌，pH波动会改变细胞表面的电荷，进而影响营养物质的吸收，其次，细菌中相关酶的活性也会受到pH的影响。强酸(pH < 5.0)或强碱(pH > 10.0)条件对HNAD细菌的生长有负面影响^[76-79]。基于AMO对氨的利用率大于铵根的理论，在微碱性环境中，较多的游离氨可能更有利于异养硝化作用^[80]，因此，大多数HNADMs一般适应于中性或弱碱条件(pH 7.0−9.0)。然而，一些HNADMs表现出了较高的pH耐受性：如陈茂霞等^[81]从活性污泥中分离出的气单胞菌属(Aeromonas) HN-02，其最佳pH范围为6.0−9.0，24 h时对NH₄⁺-N的去除率为82.93%−90.20%，当pH为2.3和4.0，24 h时对NH₄⁺-N的去除率仍能达到61%和73%；Acinetobacter sp. JR1^[6]在pH 4.5−10.0时，对TN的最大去除效率均能达到96%以上。综上可知，部分HNADMs在强酸或强碱环境中仍表现出了较高的脱氮能力，该类菌株在酸性含氮废水(如食品废水)及碱性含氮废水(如海产养殖废水)处理中具有广阔的应用潜能。

2.3 耐盐HNADMs

理论上，高盐环境会导致菌体内渗透压升高，影响细胞的结构、酶活性和代谢活性^[36]。高盐会影响营养物质进入细胞内的运输效率，导致生物脱氮效率降低，难以达到排放标准。然而令人惊讶的是，许多HNADMs如墨西哥微小杆菌(Exiguobacterium mexicanum) SND-01^[82]和门多萨假单胞菌(Pseudomonas mendocina) TJPU04^[83]在高盐(盐度大于1%)含氮废水中仍然可以进行高效的氮代谢和氮去除。Bacillus methylotrophicus L7^[38]进行异养硝化过程的最适盐度为0.0 g/L NaCl，而在好氧反硝化过程中，其最适盐度为10.0 g/L NaCl。从海洋沉积物中分离出的嗜盐Vibrio diabolicus SF16^[36]在高盐条件下具有良好的异养硝化-好氧反硝化能力，盐度为1%−5%时能够去除91.82%的NH₄⁺-N (119.77 mg/L)和99.71%的NO₃⁻-N (136.43 mg/L)。Aeromonas sp. HN-02^[84]的抗氧化酶体系活性上升以减少高盐对菌体造成的损伤，使其具有一定的耐盐性。鞘氨醇盒菌(Sphingopyxis sp.) CY-10^[85]去除NH₄⁺-N的最佳盐度在0−5%之间，最大脱氮速率为6.25 mg/(L·h)，代谢组学数据显示，在5%盐度条件下，蔗糖和d-塔格糖(d-tagatose)的代谢水平显著升高，使菌株能够调节渗透压并在高盐环境中生存。Pseudomonas aeruginosa P-1^[86]表现出了较强的高盐度耐受性，当盐度低于40.0 g/L时，在24 h内可去除高达90%的NH₄⁺-N。当前，关于耐盐HNADMs的研究仍聚焦于实验室规模的分离和性能验证，对于该类菌株的耐盐机制及在实际废水处理中的应用仍需进一步研究。

2.4 寡营养耐受HNADMs

在传统的污水处理厂中，较低的碳氮比(C/N)有利于氨氧化过程，因为较高的有机负荷可能对自养硝化菌(autotrophic nitrifying bacteria, ANB)产生不利影响。与ANB相比，HNADMs需要更高的C/N，且适当的C/N可以显著加快硝化过程中的硝化速率。不同HNADMs的最佳C/N各不相同，大多数研究表明，较高的C/N更有利于铵的去除。一般情况而言，大多数HNADMs的最佳C/N比在8−20^[87]之间，菌株可获得较好的生长速率和反硝化效率。对于HNAD而言，碳源一部分作为电子供体，一部分作为生物体碳来源，匮乏的碳源环境会显著抑制反硝化作用并降低反硝化微生物的生物量。C/N为5时，Pseudomonas sp. M-33^[33]对NH₄⁺-N的去除率达90%。C/N为4时，Pseudomonas stutzeri ADP-19^[35]的NH₄⁺-N去除率为96.5%。Acinetobacter sp. Y16^[32]的最适C/N为2，在寡营养环境下其对NH₄⁺-N的去除效率最高可达63%，是目前发现的HNADMs进行氮代谢的最低C/N。寡营养菌株的发现扩大了HNADMs在饮用水水源地、寡营养湖泊等水体中的应用范围。

2.5 重金属耐受HNADMs

Fe、Mg、Cu等重金属是全酶的辅助因子，在催化生化反应中非常重要，然而金属含量过高会抑制微生物活性^[88]。例如当Cu²⁺浓度为1.5 mg/L时，气单胞菌属(Aeromonas sp.) HN-02^[84]对NH₄⁺-N的去除几乎停止。工业废水中通常含有Cu²⁺、Zn²⁺等重金属离子，因此，应更加重视含氮工业废水系统中重金属的影响。部分HNADMs表现出了较强的重金属耐受特性：添加Fe²⁺ (11.2 mg/L)可显著减少多能副球菌(Paracoccus versutus) LYM^[89]反硝化过程中NO₂⁻-N的积累。贪铜菌属(Cupriavidus sp.) S1^[15]对分别含有Cu²⁺ (8.0 mg/L)和Zn²⁺ (20.0 mg/L)的废水可实现90%以上NH₄⁺-N的去除；0.5 mmol Cu²⁺ (31.8 mg/L)和0.5 mmol Zn²⁺ (32.5 mg/L)对Bacillus sp. PK15^[26]的生长以及NH₄⁺-N的去除未产生影响；80.0 mg/L的Cu²⁺和Zn²⁺单独存在时对Pseudomonas putida ZN1的生长和NH₄⁺-N的去除几乎未产生不利影响，copB-I成功表达是ZN1耐受较高Cu²⁺的原因^[8]；铜绿假单胞菌(Pseudomonas aeruginosa) PCN-2^[90]在好氧及pH 8.0−9.0条件下，9 h内能同时去除400 mg/L的NO₃⁻-N并还原5.0 mg/L的Cr⁶⁺；ChrR (铬酸还原酶基因)和反硝化基因的表达使PCN-2具备了同时还原Cr⁶⁺和NO₃⁻-N的能力。当Cr³⁺浓度低于60 mg/L时，褪色沙雷氏菌(Serratia marcescens) CL1502^[91]的异养硝化能力不受影响，当浓度增加到150 mg/L时，异养硝化能力仅降低了10%；Pseudomonas aeruginosa P-1^[87]表现出了较强的Cu²⁺及Cr⁶⁺耐受性，当Cu²⁺为30.0 mg/L和50.0 mg/L时，菌株P-1对NH₄⁺-N的去除率分别为100%和52.18%，在50 mg/L的高Cr⁶⁺环境下，80%以上的NH₄⁺-N能够被P-1去除；在60 mg/L的Cr⁶⁺条件下，Pseudomonas putida ZN1^[92]可削减94.86%的NH₄⁺-N。一方面，低浓度的重金属如Co²⁺、Mn²⁺、Fe²⁺等会以辅助因子的身份参与到酶的催化过程进而促进生物反应的进行，另一方面，重金属抗性基因的表达使其对相应的重金属具有抗性；然而，当其浓度超过安全阈值时会通过损伤微生物细胞膜，阻碍代谢和分子合成从而抑制生物过程；有毒有害重金属如Cr⁶⁺及Cd等在生物体中可以导致自由基的形成并损伤DNA^[93]。耐重金属的HNADMs基于其经济、高效和环境友好的优势，在处理含Cr、Cd等重金属的污染水体领域引起了广泛关注，关于其应用模式有待进一步完善。

2.6 同步脱氮除磷的HNADMs

近年来，部分HNADMs在有氧条件下进行硝化反硝化的同时也可利用O₂、NO₃^–-N或NO₂^–-N作为电子受体氧化有机物获得能量促进磷的吸收，即在硝化反硝化脱氮的同时实现过量吸磷作用^[94]，例如Pseudomonas stutzeri YG-24^[79]、Bacillus cereus GS-5^[42]、阴沟肠杆菌(Enterobacter cloacae) HW-15^[95]和Acinetobacter sp. strain C-13^[79]等菌株具有同步脱氮除磷的功能，打破了传统聚磷菌(phosphorus accumulating organisms, PAOs)和反硝化聚磷(denitrifying P-accumulating organisms, DPAOs)需要厌氧、缺氧/好氧交替的环境条件限制，该类微生物被归类为异养硝化-好氧反硝化脱氮除磷菌(heterotrophic nitrification-aerobic denitrification and aerobic P removal, HNADPR)。菌株Pseudomonas putida strain NP5^[96]可将水体中的NH₄⁺-N和PO₄³⁻-P几乎全部去除，催化除磷的关键酶多聚磷酸激酶(polyphosphate kinase, PPK)、外切聚磷酸酶(polyphosphate esterase, PPX)的酶活性显著升高，进一步说明了该菌株同步脱氮除磷的代谢机制。PPK可催化三磷酸腺苷(adenosine triphosphate, ATP)的末端磷酸转化为长链多聚磷酸(polyphosphate, Poly-p)，PPX是催化多磷酸盐释放磷酸的主要酶^[35]，在菌株Pseudomonas stutzeri ADP-19^[35]中，PPK的酶活性远高于PPX，表明聚磷为ADP-19除磷的主要方式。ppk基因已在Pseudomonas putida strain NP5^[96]、Bacillus cereus GS-5^[42]、Enterobacter cloacae HW-15^[95]和Acinetobacter sp. strain C-13^[79]等HNADMs中被成功扩增。虽然生物脱磷基因ppk和ppx已被成功扩增，但上述基因究竟是如何发挥功能的尚不清楚。因此，需要进一步深入探究。

传统的聚磷菌PAOs储存的细胞内磷高达90%，并且Poly-p [约占总磷(total P, TP)的50%]是生物除磷(biological phosphorus removal, BPR)过程微生物细胞中磷的主要形式^[97] (图3 Pathway 2)；好氧除磷菌如希瓦氏菌属(Shewanella sp.) CF8-6^[98]、皱褶念珠菌(Diutina rugosa) BL3^[99]等吸收磷酸盐主要存在于胞外多聚物(extracellular polymeric substance, EPS) (60%−70%)中(图3 Pathway 3)，例如菌株Diutina rugosa BL3的EPS提取物中正磷酸盐(orthophosphate, Ortho-P)和Poly-p的TP占比分别为41.5%和38.6%。当前关于HNADPR菌的磷代谢路径研究相对匮乏，Pseudomonas stutzeri ADP-19主要依赖EPS中过量P积累以及磷酸盐同化^[35]。Huang等^[41]通过P平衡分析推测出HNADPR菌株Pseudomonas aeruginosa strain SNDPR-01吸收PO₄³⁻-P后，14%−41%转化为了细胞膜磷脂(图3 Pathway 1)、10%−14%转化为了细胞质脂类以及37%−46%转化为EPS，其中磷酸单酯是EPS中P的主要存在形式；氮源会影响HNADMs的磷代谢，例如SNDPR-01以NH₄⁺-N为唯一氮源时，细胞膜中的P含量(41%)略高于EPS (37%)，以NO₂⁻-N或NO₃⁻-N为唯一氮源时，细胞膜中的P含量(14%或17%)显著低于EPS (45%和46%)。上述研究表明，细胞膜、细胞体和EPS在P代谢中起着重要作用，HNADPR、DPAOs和PAOs之间的磷代谢可能存在差异，并且氮源类型会影响HNADPR菌株的磷代谢途径，但是他们的磷代谢机制目前尚未明确。尽管近年来关于HNADMs的分离和脱氮除磷能力的研究较多，但对生物除磷机制的研究较少，目前针对磷代谢的研究大多是通过对HNADPR磷酸盐代谢过程中的磷分布、磷种类和相关功能基因的表达来推断，但该基因究竟是如何发挥功能的尚不清楚，亟须进一步研究。

2.7 复杂有机物降解HNADMs

在反硝化过程中，有机物是反硝化的重要电子供体来源。可利用碳源类型的多样性使HNADMs在生物脱氮体系中具有很高的竞争力及优势。除了琥珀酸钠、柠檬酸钠、乙酸钠和葡萄糖等常规碳源外，研究发现部分异养硝化菌、好氧反硝化菌还能利用其他难降解有机物作为碳源，例如苯酚、喹啉、芳香族化合物、抗生素和全氟化合物等。动胶菌样申氏菌(Shinella zoogloeoides) BC026^[100]能够以难降解的吡啶为唯一碳源、氮源和能量供给自身的生长，在最佳条件下该菌株能够在45.5 h内完全降解1 806 mg/L的吡啶，产物检测和遗传分析表明有58%的吡啶转化为铵。食吡啶红球菌(Rhodococcus pyridinivorans) RL-GZ01^[101]可以通过利用外部碳源，如邻苯二甲酸酯(di-2-ethylhexyl phthalate, DEHP)等有机化合物，同时去除氮和磷，84 h内对TN (71.2 mg/L)、PO₄³⁻-P (10.89 mg/L)和DEHP (5.0 mg/L)的去除率分别达到64.45%、76.30%和98.94%，DEHP中间体和代谢途径的鉴定表明该菌株将DEHP转化为邻苯二酚作为碳源进行细胞生长，DEHP提供的碳源可作为碳骨架用于合成谷氨酸。HNADMs Pseudomonas aeruginosa KDQ4^[34]在未外加碳源的条件下能够去除800 mg/L的喹啉以及400 mg/L的苯酚。HNADMs的发现，扩大了HNADMs在难降解有机废水中的应用，HNADMs在含有复杂有机物废水领域具有良好的应用潜能。

2.8 抗生素耐药性HNADMs

抗生素作为四大新污染物之一，其过量排放问题受到了世界范围内的广泛关注。研究表明，HNADMs在环境中广泛存在，抗生素对HNADMs的影响与抗生素的类型和浓度有关。残留的抗生素通过阻碍硝化或反硝化细菌来影响生物脱氮。在序列批反应器接种肺炎克雷伯氏菌(Klebsiella pneumoniae) y6，当氨苄西林浓度为5 mg/L时，反应器脱氮速率和有机物去除率分别为0.79 kg/(m³·d)和7.17 kg/(m³·d)，当提高氨苄西林浓度为25 mg/L后，反应器脱氮率和有机物去除率分别下降了23.8%和13.4%，进一步增加抗生素浓度至50 mg/L时去除率分别下降48.6%和50.9%^[65]。与外源分离株相比，土著HNADMs具有更高的抗生素耐药性，Huang等^[102]对从水产养殖废水中分离的HNADMs的耐药性进行评估，发现由普遍海单胞菌(Marinomonas communis)和泰坦尼克号盐单胞菌(Halomonas titanicae)组成的MCH、由海水海单胞菌(Marinomonas aquimarina)和Halomonas titanicae组成的MAH微生物复合菌群以及单菌株在最适条件下对NH₄⁺-N和NO₂⁻-N的去除率均高于95%，同时对多肽类、四环素类、磺胺类、呋喃类和大环内酯类抗生素均具有较强耐药性。在地表水体环境中，磺胺甲恶唑(sulfamethoxazole, SMX)因其高检测频率、强稳定性和高流动性成为最受关注的抗生素之一，研究表明，在2 mg/L的SMX环境下，与糖酵解、三羧酸循环、反硝化相关基因(narG、narH、narI和nirK)和电子转移链相关基因(nuoA、nuoB、nuoD、nuoF、nuoG、nuoI和nuoL)均发生了下调^[103]。部分HNADMs可以通过调节抗生素抗性基因(antibiotic resistance genes, ARGs)的表达来缓解抗生素的负面影响^[103]。Zhao等^[104]通过转录组分析将好氧反硝化微生物Pseudomonas aeruginosa PCN-2对SMX的抗性归结为以下原因：(1) 多药物外排系统的表达，在较高的SMX条件下，更多的SMX可以被泵出细胞膜；(2) 生物膜的形成；(3) 其他机制，如群体感应(quorum sensing, QS)以及脓青素的产生，QS系统和脓青素合成相关基因(如lasB、lasR、lasI、rhlR和rhlI)均显著上调，表明它们参与了菌株PCN-2对SMX的抗性。目前，尽管少数对抗生素具有抗性的HNADMs已被筛选，但抗生素对HNADMs的影响并未受到广泛关注，关于HNADMs与抗生素的互作影响及互作机制有待进一步揭示。

3 极端环境耐受HNADMs的应用现状

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不同于传统含氮废水，冬季低温废水、高氨氮废水、养殖废水、高浓度有机废水、抗生素废水、重金属废水等由于其特殊性，采用传统的生物脱氮方式很难达到良好的处理效果。将具有极端环境耐受性的HNADMs进行富集、驯化并应用于非典型废水处理中可大幅度降低运行成本并提高污染物去除效率。例如Yao等^[35]为了解决低温条件下污水处理厂脱氮效率低下问题，将耐冷的HNAD菌群投加到反应器中进行生物强化并增加DO浓度，10 ℃下每升污水对NH₄⁺-N和NO₃⁻-N的去除率分别高达2.44 mg/h和7.68 mg/h。将Acinetobacter sp. Z1^[71]接种于经处理的新疆某污水处理厂水样中，10 ℃ (中国北部冬季典型污水温度)条件下NH₄⁺-N和PO₄³⁻-P的去除率分别达到70%和40%以上，证明其在冬季污水处理中有良好的应用潜力；当前关于耐低温HNADMs的应用研究仍以实验室小试为主，仍需进一步加快高效耐低温HNADMs的研发及应用方式的探究。传统脱氮微生物对10‰以上的盐度高度敏感^[105]，海水养殖废水由于其高盐度抑制了自养硝化菌的活性，硝化速率显著降低，因此，亟须开发耐盐、高活性的脱氮微生物。Xiang等^[106]筛选出一株新型嗜盐菌佐贝尔菌(Zobellella) B307，盐度的增加对B307的生长无抑制作用，菌株B307在盐度为65‰时可去除90%以上的NH₄⁺-N和97.1%的NO₃⁻-N，将其投加于移动床生物膜反应器(moving bed biofilm reactor, MBBR)中处理人工合成海水养殖废水(盐度30‰)，投加前后MBBR系统对COD、NH₄⁺-N、NO₃⁻-N和NO₂⁻-N的去除率由83.01%、83.10%、76.68%和80.98%分别提高至95.57%、94.44%、85.67%和87.44%，且接种菌株后HNADMs的丰度及参与氮代谢的功能基因的丰度均有所提高。Zhang等^[107]利用HNADMs混合菌与小球藻建立共生体系处理高氨沼液，TN和TP的去除率分别达到了79.4%和90.4%，相较于对照组(HNADMs混合菌单独处理)分别提高了28.9%和67.6%。Wang等^[108]利用Pseudomonas mendocina K0、布鲁氏菌属(Brucella sp.) K1、Pseudomonas putida T4和Paracoccus sp. T9构建高效HNADMs复合菌群处理石油废水，固定化的复合菌群显著增强了好氧颗粒污泥(aerobic granular sludge, AGS)的脱氮效果，NH₄⁺-N和TN的去除率分别为92.4%和79.8%，与对照组相比AGS的形成量提高了16.7%。其他HNADMs应用案例见表4。

综上所述，虽然部分HNADMs在一些极端污染水环境中表现出了巨大的应用优势，但仍以小试规模为主，且小试实验中微生物的作用环境是人为可控的，从理论研究到实际工程应用需要综合考虑污水类型的多样性、水质条件的复杂性以及在实际污水处理过程中如何保持功能微生物稳定且长期地发挥作用等现实问题。未来研究可集中于极端HNADMs的资源开发(高效单菌株、复合功能菌群)、HNADMs代谢路径的明晰以及应用模式的开发及工艺参数的优化等方向。

4 结论与展望

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HNADMs鉴于其高效的脱氮效能及特殊的氮转化特征引起了广泛的关注，大量研究表明，HNADMs在水体、土壤等不同环境介质中具有普遍存在性，HNADMs表现出了系统发育多样性、底物基质类型多样、环境适应性强和氮代谢路径复杂的特征。随着研究的深入，部分HNADMs表现出了较强的极端温度、pH、盐度、寡营养、重金属等环境胁迫耐受性，同时表现出了高效同步脱氮除磷、复杂有机物降解和抗生素耐药性，以上新功能的发现不仅丰富了微生物资源，同时为HNADMs在非典型含氮废水处理中的应用提供了更多的可能性。当前关于极端HNADMs的研究多集中在单一环境胁迫下功能菌株的分离筛选、胁迫因子耐受范围的确定及耐受机理的初步探究等方面，其应用研究主要集中于实验室规模，尚无关于其中试及全规模应用的报道。综上所述，HNADMs在污水脱氮尤其是复杂水体生物脱氮领域展示出了较高的研究价值和应用潜能，然而在基础理论及应用技术层面仍存在一定瓶颈，建议后续研究可聚焦于以下方向：(1) 目前极端环境耐受HNADMs的研究刚刚起步，HNADMs功能菌种资源相对匮乏，且由于污染水体中的污染物类型愈加复杂多样，应进一步加大不同极端环境下HNADMs的筛选，驯化更多环境适应性好且污染物去除效能高的菌株以丰富HNADMs菌种库；(2) 鉴于HNADMs代谢过程复杂且研究基础相对薄弱的现状，应积极利用基因组学、转录组学等平台并与反应动力学、质谱技术等相结合，多角度、全方位绘制HNAD模式菌株的氮代谢通路，同时结合基因敲除、基因重组及异源表达技术破译功能菌株对环境胁迫因子的耐受机制；(3) 继续探究HNADMs的应用模式，深入研究HNADMs与当前主流污水处理工艺如厌氧-缺氧-好氧工艺(anaerobic-anoxic-oxic, A²O)、MBBR、序批式反应器(sequencing batch reactor, SBR)的兼容性、确定运行参数并制定工艺优化调控策略，开发极端HNADMs在复杂污染水体中定殖并长期起主导作用的应用方式，重点关注基于生物膜的生物强化技术与极端HNADMs相结合的应用可行性。

基金

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河北省重点研发计划(21373904D)
河北科技大学引进人才科研启动基金(1181374)

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2024年第64卷第12期

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doi: 10.13343/j.cnki.wsxb.20240395

接收时间：2024-06-28
首发时间：2026-03-21
出版时间：2024-09-23

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出版历史

收稿日期：2024-06-28
录用日期：2024-09-20

基金

Key Research and Development Project of Hebei Province(21373904D)

河北省重点研发计划(21373904D)

Doctoral Starting up Foundation of Hebei University of Science and Technology(1181374)

河北科技大学引进人才科研启动基金(1181374)

作者信息

1 河北科技大学环境科学与工程学院, 河北省污染防治生物技术重点实验室, 河北石家庄 050018

2 河北科技大学建筑工程学院, 河北石家庄 050018

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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Origin	Strains	Gram
G⁺: Gram-positive; G⁻: Gram-negative; − : Fungi without Gram-differentiation.
Wastewater	Coking wastewater	Cupriavidus sp. S1^[11]	G⁻
Wastewater treatment plants	Ochrobactrum anthropi HND19^[12]	G⁻
Aquaculture wastewater	Pseudomonas mendocina S16^[13]	G⁻
Pig farm wastewater	Acinetobacter sp. T1^[14]	G⁻
Eutrophic lake	Pseudomonas sp. N31942^[15]	G⁻
The Songhua River	Microbacterium esteraromaticum SFA13^[16]	G⁺
Artificial reservoir	Acinetobacter sp. H36^[17]	G⁻
Activated sludge	Acinetobacter calcoaceticus TY1^[18]	G⁻
Soil	Paddy field soil	Arthrobacter arilaitensis Y10^[19]	G⁺
Soil	Pseudomonas sp. PHR6^[20]	G⁻
Soil	Basidiomycetes^[21]	−
Soil	Penicillium^[22]	−
Paddy soil	Pseudomonas tolaasii Y-11^[23]	G⁻
Soil	Bacillus cereus PK5^[24]	G⁺
Soil	Bacillus subtilis PK15^[24]	G⁺

Origin

Strains

Gram

G⁺: Gram-positive; G⁻: Gram-negative; − : Fungi without Gram-differentiation.

Wastewater

Coking wastewater

Cupriavidus sp. S1^[11]

G⁻

Wastewater treatment plants

Ochrobactrum anthropi HND19^[12]

G⁻

Aquaculture wastewater

Pseudomonas mendocina S16^[13]

G⁻

Pig farm wastewater

Acinetobacter sp. T1^[14]

G⁻

Eutrophic lake

Pseudomonas sp. N31942^[15]

G⁻

The Songhua River

Microbacterium esteraromaticum SFA13^[16]

G⁺

Artificial reservoir

Acinetobacter sp. H36^[17]

G⁻

Activated sludge

Acinetobacter calcoaceticus TY1^[18]

G⁻

Soil

Paddy field soil

Arthrobacter arilaitensis Y10^[19]

G⁺

Soil

Pseudomonas sp. PHR6^[20]

G⁻

Soil

Basidiomycetes^[21]

−

Soil

Penicillium^[22]

−

Paddy soil

Pseudomonas tolaasii Y-11^[23]

G⁻

Soil

Bacillus cereus PK5^[24]

G⁺

Soil

Bacillus subtilis PK15^[24]

G⁺

Number	Metabolic pathway	References
1	NH₄⁺-N→NO₂^–-N→NO₃⁻-N→NO₂⁻-N→NO→N₂O→N₂	[4]
2	NH₄⁺-N→NH₂OH→NO₂⁻-N→NO₃⁻-N→NO₂⁻-N→N₂	[48]
3	NH₄⁺-N→NH₂OH→NO₂⁻-N→N₂O→N₂	[49]
4	NH₄⁺-N→NH₂OH→NO₂⁻-N→NO₃⁻-N→N₂O→N₂	[50]
5	NH₄⁺-N→NO→N₂O	[51]
6	NH₄⁺-N→N₂O→N₂	[52]

Number

Metabolic pathway

References

NH₄⁺-N→NO₂^–-N→NO₃⁻-N→NO₂⁻-N→NO→N₂O→N₂

[4]

NH₄⁺-N→NH₂OH→NO₂⁻-N→NO₃⁻-N→NO₂⁻-N→N₂

[48]

NH₄⁺-N→NH₂OH→NO₂⁻-N→N₂O→N₂

[49]

NH₄⁺-N→NH₂OH→NO₂⁻-N→NO₃⁻-N→N₂O→N₂

[50]

NH₄⁺-N→NO→N₂O

[51]

NH₄⁺-N→N₂O→N₂

[52]

Strain	T/℃	NH₄⁺-N removal	NO₃⁻-N removal
−: It is not mentioned in the literature.
Acinetobacter sp. TAC-1^[4]	5	94.6%	40%
Microbacterium sp. SFA13^[71]	5	1.6 mg/(L·h)	0.24 mg/(L·h)
Pseudomonas sp. M-11^[66]	10	98%	−
Pseudomonas putida Y-9^[5]	15	99.36%	82%
Acinetobacter sp. HA2^[37]	10	100%	−
Pseudomonas migulae AN-1^[72]	10	1.6 mg/(L·h)	1.5 mg/(L·h)
Acinetobacter TY1^[73]	8	97.4%	81.85%
Acinetobacter sp. Z1^[74]	10	96%	97.5%
Chelatococcus daeguensis TAD1^[63]	50	100%	100%
Anoxybacillus contaminans HA^[64]	55	71%	74.7%

Strain

T/℃

NH₄⁺-N removal

NO₃⁻-N removal

−: It is not mentioned in the literature.

Acinetobacter sp. TAC-1^[4]

94.6%

40%

Microbacterium sp. SFA13^[71]

1.6 mg/(L·h)

0.24 mg/(L·h)

Pseudomonas sp. M-11^[66]

98%

−

Pseudomonas putida Y-9^[5]

99.36%

82%

Acinetobacter sp. HA2^[37]

100%

−

Pseudomonas migulae AN-1^[72]

1.6 mg/(L·h)

1.5 mg/(L·h)

Acinetobacter TY1^[73]

97.4%

81.85%

Acinetobacter sp. Z1^[74]

96%

97.5%

Chelatococcus daeguensis TAD1^[63]

100%

Anoxybacillus contaminans HA^[64]

71%

74.7%

HNADMs	Wastewater type	Scale
Chelatococcus daeguensis TAD1^[63]	Synthetic wastewater	Lab
Zobellella B307^[106]	Mariculture wastewater	Lab
Acinetobacter sp. TN-14^[109]	Pig farm wastewater	Lab
Pseudomonas sp. HJ3^[110]	Coal gasification wastewater	Lab
HNADMs flora^[111]	Livestock wastewater	Lab
Bacillus subtilis GHSP10^[112]	Aquaculture sewage	Lab
Acinetobacter sp. Z1^[75]	Wastewater treatment plants	Lab
pseudomonas sp. DM02^[76]	Aquaculture wastewater	Lab

HNADMs

Wastewater type

Scale

Chelatococcus daeguensis TAD1^[63]

Synthetic wastewater

Lab

Zobellella B307^[106]

Mariculture wastewater

Lab

Acinetobacter sp. TN-14^[109]

Pig farm wastewater

Lab

Pseudomonas sp. HJ3^[110]

Coal gasification wastewater

Lab

HNADMs flora^[111]

Livestock wastewater

Lab

Bacillus subtilis GHSP10^[112]

Aquaculture sewage

Lab

Acinetobacter sp. Z1^[75]

Wastewater treatment plants

Lab

pseudomonas sp. DM02^[76]

Aquaculture wastewater

Lab