水体富营养化是当前水环境保护工作的重点关注问题,微生物修复富营养化水体具有高效、低耗且不产生二次污染等特点,已经成为富营养化水体生态修复的一种重要方式。近年来,对反硝化聚磷菌的研究及其在污水处理工艺中的应用越来越广泛。不同于传统的反硝化细菌联合聚磷菌去除氮磷工艺,反硝化聚磷菌在交替厌氧、缺氧/好氧条件下能同时进行脱氮除磷而被广泛关注与研究。值得注意的是,近几年报道的部分微生物仅在好氧条件下就可进行同时脱氮除磷,但是其脱氮除磷机理仍未理清。基于此,文中总结了目前发现的反硝化聚磷菌和同时硝化反硝化聚磷微生物的种类及特点,并对其脱氮与除磷的关系及其机理进行了系统性分析,对目前反硝化除磷存在的问题进行了梳理,最后对今后的研究方向进行了展望,以期为完善反硝化聚磷菌的脱氮除磷机理及工艺改进提供参考。
Water eutrophication poses great threats to protection of water environment. Microbial remediation of water eutrophication has shown high efficiency, low consumption and no secondary pollution, thus becoming an important approach for ecological remediation. In recent years, researches on denitrifying phosphate accumulating organisms and their application in wastewater treatment processes have received increasing attention. Different from the traditional nitrogen and phosphorus removal process conducted by denitrifying bacteria and phosphate accumulating organisms, the denitrifying phosphate accumulating organisms can simultaneously remove nitrogen and phosphorus under alternated anaerobic and anoxic/aerobic conditions. It is worth noting that microorganisms capable of simultaneously removing nitrogen and phosphorus absolutely under aerobic conditions have been reported in recent years, but the mechanisms remain unclear. This review summarizes the species and characteristics of denitrifying phosphate accumulating organisms and the microorganisms capable of performing simultaneous nitrification-denitrification and phosphorous removal. Moreover, this review analyzes the relationship between nitrogen removal and phosphorus removal and the underlying mechanisms, discusses the challenges of denitrifying phosphorus removal, and prospects future research directions, with the aim to facilitate process improvement of denitrifying phosphate accumulating organisms.
随着工农业的迅速发展,大量未经处理或处理不完全的废水被排入天然水体中,导致水体富营养化程度日趋严重[
传统的生物脱氮除磷工艺中,活性污泥成分复杂,存在着不同微生物类群,聚磷菌(phosphate accumulating organisms, PAOs)和反硝化细菌之间存在碳源竞争和泥龄差异,致使氮与磷的去除不能同时高效率[
传统的生物脱氮除磷是通过脱氮细菌与PAOs联合进行,且脱氮与除磷过程分开进行。脱氮过程主要由氨化、硝化和反硝化等细菌通过氨化、硝化和反硝化3个过程实现[
生物除磷主要依据强化生物除磷(enhanced biological phosphorus removal, EBPR)理论,即PAOs能够进行厌氧释磷和好氧吸磷过程[
不同于传统PAOs仅能利用氧气作为电子受体,DPAOs可在没有氧气的条件下以NO3–-N/ NO2–-N作为末端电子受体,利用细胞内的聚羟基烷酸酯(poly-β-hydroxyalkanoates, PHA)在缺氧条件下完成脱氮与除磷的双过程,从而高效、低耗地实现对污水脱氮除磷的双重目的[
DPAOS及其脱氮除磷特点
DPAOs and their characteristics of nitrogen and phosphorus removal
Strain | Genus | Nitrogen source | Carbon source | Temperature (℃) | pH | C/N | Nitrogen removal efficiency (%) | Phosphorus removal efficiency (%) | References |
–: Indicates no relevant data in the literature. | |||||||||
RT1901 | NO3–-N | Sodium acetate | 30 | 8.00 | 10 | 77.71 | 98.28 | [ |
|
qdcs18 | NH4+-N+NO3–-N | Sodium acetate | 28 | 8.00 | ‒ | 90.00 | 85.00 | [ |
|
ZK-1 | Ammonium sulfate+peptone | Sodium acetate | ‒ | 7.85 | ‒ | 98.70 | 90.70 | [ |
|
j16 | NO3–-N | Sodium acetate | 30 | 7.20–8.00 | ‒ | 96.03 | 94.55 | [ |
|
K14 | NO3–-N | Sodium pyruvate | 27 | 7.50 | 10 | 99.78 | 98.00 | [ |
|
ZK-2 | NO3–-N+NO2–-N | Sodium acetate | ‒ | 7.00 | 10 | 79.62 | 87.22 | [ |
|
C-17 | NO3–-N | Sodium acetate | 30 | ‒ | ‒ | 87.00 | 75.00 | [ |
|
N-8 | NO3–-N | Sodium acetate | ‒ | 7.00–9.00 | ‒ | 82.69 | 69.20 | [ |
|
Q-hrb05 | NO3–-N | Sodium acetate | 30 | 7.00 | ‒ | 81.00 | 88.00 | [ |
据前人研究可知,影响DPAOs脱氮除磷的因子有很多,研究较多的有碳源类型、温度、C/N比、pH等,这些因子与细菌细胞生长、酶的活性以及物质能量代谢等密切相关[
1983年,Robertson等[
SNDPR菌株及其脱氮除磷特点
SNDPR strains and their characteristics of nitrogen and phosphorus removal
Strain | Genus | Optimum nitrogen source | Optimum carbon source | Optimum temperature (℃) | pH | C/N | The initial concentraion and removal efficiency of N ((mg/L)/%) | The initial concentraion and removal efficiency of P ((mg/L)/%) | References |
‒: Indicates no relevant data in the literature. | |||||||||
NP5 | NH4+-N | Succinate | 30 | ‒ | 10.00 | 100.00/99.65 | 70.00/99.55 | [ |
|
SND5 | NO3–-N | Lactic acid | ‒ | ‒ | 10.00 | 80.00/99.20 | 20.00/90.00 | [ |
|
HHEP5 | NO3–-N+ NO2–-N | Glucose | 28 | 7.50 | 5.00– 20.00 | 42.80/95.00 | 2.56/99.00 | [ |
|
ADP-19 | NH4+-N | Sodium acetate | 30 | 7.00– 8.00 | 10.00 | 100.00/96.50 | 20.00/73.30 | [ |
|
C-13 | NH4+-N | Sodium acetate | 30 | 7.50 | 10.00 | 181.00/92.10 | 134.80/81.70 | [ |
|
GHSP10 | NH4+-N+ NO3–-N | Sodium acetate | 28 | 7.50 | 10.00– 20.00 | 50.00/95.84 | 10/.0089.88 | [ |
|
YG-24 | NH4+-N | Sodium citrate | 30 | ‒ | 8.00 | 35.72/85.28 | 2.28/51.21 | [ |
|
HW-15 | NH4+-N | Sodium acetate | 30 | ‒ | 8.00 | 99.76/99.16 | 36.63/73.00 | [ |
|
GS-5 | NO3–-N | Sodium acetate | 35 | 7.50 | 7.50 | 34.50/96.00 | 5.70/84.00 | [ |
|
YP1 | NO3–-N | Sodium citrate | ‒ | 8.00 | ‒ | 30.00/58.80 | 8.00/75.30 | [ |
|
ISTOD1 | NO3–-N | Sugarcane molasses | ‒ | ‒ | ‒ | 47.53/97.30 | 13.10/93.00 | [ |
在传统的生物脱氮除磷工艺中,脱氮过程与除磷过程分别在不同微生物内完成,由于两个过程对于氧的需求不同,且不同微生物之间存在碳源竞争和生长周期的差异,因此,传统的生物脱氮除磷工艺需要较多的设施才能实现。脱氮与除磷两个过程的联合作用是生物脱氮除磷的前提,DPR工艺将两个过程耦合在一起,可同时进行。有研究表明,将硝化和反硝化耦合后可加速氮的去除率,从而缓解水体生态系统的富营养化,且NO3–-N和NO2–-N不仅能够诱导反硝化基因的转录,还能够激活聚磷基因的表达[
微生物的脱氮过程可分为硝化和反硝化过程。其中,硝化过程的氮转化途径为NH4+-N→NH2OH→NO2–-N→NO3–-N。NH4+-N经氨单加氧酶(ammonia monooxygenase, AMO)催化产生NH2OH,受
无论是在好氧还是缺氧条件下,微生物的聚磷过程都是吸取环境中的磷酸盐,然后以poly-P的形式存于细胞内,此过程是由多聚磷酸盐激酶(polyphosphate kinase, PPK)催化完成的[
由上述反应可知,微生物除磷过程与物质代谢和能量代谢过程密切相关。
厌氧条件下,DPAOs胞内的poly-P水解,产生的无机磷酸盐(PO43–-P)会被释放到水中。同时,DPAOs可利用poly-P水解产生的ATP和糖原分解提供的还原力NADH2将易降解的低分子脂肪酸(volatile fatty acids, VFA),如乙酸、丙酸、正丁酸、戊酸和异戊酸等吸收到体内,被活化生成乙酰辅酶A (acetyl-CoA, AcCoA),最终以PHA的形式储存在体内,以上即为DPAOs的厌氧释磷过程。值得注意的是,PHA主要包括PHB和PHV,是一类可生物降解的碳聚合物,既能在有氧条件下作为生化反应的碳源为细胞生长和代谢活动提供ATP,又能在缺氧条件下直接作为能源分解,形成质子驱动力及为电子传递链提供电子[
DPAOs的脱氮除磷机理图
Mechanistic diagram of nitrogen and phosphorus removal by DPAOs.
与厌氧-缺氧/好氧交替条件下完成脱氮除磷的DPAOs不同,SNDPR菌的脱氮除磷过程均发生在好氧条件下,因而能够在有氧环境中实现同步脱氮除磷[
SNDPR菌株脱氮除磷机理模型
Mechanistic model of nitrogen and phosphorus removal by SNDPR strain.
反硝化脱氮除磷不仅会受碳源类型、温度、pH和C/N比的影响,还会受重金属离子、污泥龄和抑制剂等的影响。微量金属离子是微生物生命活动所需的营养物质,是生物酶活性的辅助因子,如镁、铜、锌、铁、镍、钴等。然而,对于大多数微生物而言,当金属离子质量浓度达到一定水平时,会降低微生物生长速率和活性,削弱其降解有机物和脱氮除磷的能力,甚至导致死亡[
综上所述,选择合适的环境条件对DPAOs的生长增殖与脱氮除磷能力有促进作用,但目前对于DPAOs脱氮除磷的影响因子研究多处于生理水平,对于这些环境因素是通过何种代谢通路来进行调控DPAOs脱氮除磷功能、蛋白结构是否会受到重金属离子的影响、受到影响后的蛋白质会发生何种改变以及涉及哪些基因参与其中还处于探索阶段,且各因素之间也可能相互作用,因此,目前还无法建立较为全面的响应模型。
反硝化过程是按照NO3–-N→NO2–-N→ NO→N2O→N2的过程进行的,大部分已报道的反硝化细菌在此过程中均会出现NO2–-N积累的现象,这可能是由于电子竞争、微生物种类和碳源类型等因素而导致的。(1) 电子竞争:NO2–-N积累的主要原因是NO2–-N还原速率远低于NO3–-N还原速率,NIR对电子竞争的能力低于NR,且无论碳源足够与否,均会发生电子竞争[
N2O被认为是一种强烈的温室气体,其温室效应潜力是二氧化碳的265倍[
随着对DPAOs和DPR的广泛研究,DPR工艺取得了突破性的进展。DPR技术的研究已经从实验室模拟到工程应用阶段,并取得了一些成果,如单污泥工艺和双污泥工艺。在单污泥系统中,DPAOs和硝化细菌共存于同一活性污泥中,硝化细菌需要较长的好氧时间,这会抑制DPAOs的生长和活性,使得DPR效能难以大幅度提高[
分离并鉴定兼备SND和EBPR潜能的新型细菌,可为积累在富营养化湖泊、水产养殖系统及工业和生活污水中氮磷化合物的同时降解开辟新的道路。如菌株HEPP5在海水养殖和生活污水处理中均具有99%的除磷和95%的除氮能力[
综上所述,无论是DPAOs还是SNDPR在废水处理中的工程应用都还有待继续研究,应加强对连续运行的反硝化脱氮除磷工艺菌种性能的研究,同时结合现有的城市污水厂处理工艺,进行工艺改进和创新,结合先进的自动化控制技术,对工艺过程参数进行优化,使工艺朝着经济高效、低能耗和资源化的方向发展。
DPAOs在修复富营养化水体的应用中,已经成为一种高效低耗与节能减排的重要微生物资源。理清DPAOs的脱氮除磷机理不仅能促进对不同代谢途径间相互作用的理解,也有助于现有脱氮除磷工艺的改进,对富营养化水体的生物修复以及其他含氮或磷废水中的治理具有重要意义。然而,基于前人研究报道和文献分析,本文系统分析了现有DPAOs的分布、脱氮除磷之间的关系及机理,但仍存在诸多尚未明确的地方,未来还可以从以下几个方面开展研究:(1) 对DPAOs的研究多集中于生化水平上的氮磷代谢过程,调控DPAOs氮磷代谢过程的关键基因和酶类还未明确,可从基因组、转录组、蛋白组和代谢组等多组学协同分析,更加全面地理解DPAOs的脱氮除磷机理;(2) 目前在控制N2O的释放方面的研究较少,分离筛选出既不产N2O又不积累亚硝酸盐的生物脱氮除磷菌株资源是未来氮磷污染废水治理理想的方式之一;(3) 反硝化脱氮除磷过程影响因子的响应机制尚未阐明,可结合反应动力学、光谱扫描分析等技术方法研究一些关键调控因子,构建影响因子响应模型;(4) 可通过构建高效脱氮除磷的基因工程菌来提高脱氮除磷效率,其显著优势在于能够通过基因重组或基因编辑等技术手段调控微生物的氮磷代谢过程。但构建高效脱氮除磷的基因工程DPAOs还有赖于挖掘野生菌株的高效功能基因。
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