有机污染物胞外作用机理及微生物群体感应调控特征

doi:10.11894/iwt.2021-0280

摘要/Abstract

摘要：

胞外聚合物（EPS）是微生物产生的胞外多糖、蛋白、脂质等高分子聚合物，其在吸附和降解外源有机物过程中起着重要作用。长期以来，环境系统中采用微生物法处理有机污染物主要聚焦于筛选降解菌、优化代谢条件、外源添加氧化还原材料等，而鲜有涉及微生物对外源有机污染物的胞外感应识别、功能基因调控及其降解关系的深入研究。综述了EPS的组成和性质、去除有机物的胞外作用机理及微生物的群体感应调控特征，以期对强化微生物EPS合成，实现高效去除有机污染物提供指导作用。

关键词: 胞外聚合物(EPS), 有机污染物, 群体感应, 生物吸附与降解

Abstract:

Extracellular polymer substance(EPS) is a polymer matrix consisting of extracellular polysaccharides, proteins and lipids excreted by microorganisms, which plays an important role in the process of adsorbing and degrading exogenous organics. For a long time, the microbial technologies for treating organic pollutants in environmental systems has focused on the isolation of pollutant-degrading bacteria, the optimization of metabolic condition, and the addition of exogenous redox materials. An in-depth understanding of microbial extracellular sensation and recognition of exogenous organic pollutants, its functional gene regulation and the degradation relationship between microbes and the pollutants still lacks. The composition and properties of EPS, extracellular mechanisms of organic pollutant removal and microbial quorum sensing regulation characteristics were reviewed. It was expected to provide a guidance for enhancing synthesis of microbial EPS so as to achieve highly efficient removal of organic pollutants.

Key words: extracellular polymer substance(EPS), organic pollutants, quorum sensing, biosorption and biodegradation

中图分类号:

X703

刘亚南,朱颖楠,王瑾丰,丁丽丽,任洪强. 有机污染物胞外作用机理及微生物群体感应调控特征[J]. 工业水处理, 2021, 41(6): 1-13.

Ya'nan Liu,Yingnan Zhu,Jinfeng Wang,Lili Ding,Hongqiang Ren. The extracellular removal mechanism of organic pollutants and the regulation characteristics of microbial quorum sensing[J]. Industrial Water Treatment, 2021, 41(6): 1-13.

https://www.iwt.cn/CN/Y2021/V41/I6/1

图/表 2

参考文献 126

1	Cai Fangrui , Lei Lirong , Li Youming , et al. A review of aerobic granular sludge(AGS) treating recalcitrant wastewater: Refractory organics removal mechanism, application and prospect[J]. Science of The Total Environment, 2021, 782, 146852. doi: 10.1016/j.scitotenv.2021.146852
2	Costa F , Lago A , Rocha V , et al. A review on biological processes for pharmaceuticals wastes abatement-A growing threat to modern society[J]. Environmental Science & Technology, 2019, 53 (13): 7185. 7202. URL
3	Aziz A , Basheer F , Sengar A , et al. Biological wastewater treatment (anaerobic-aerobic) technologies for safe discharge of treated slaughterhouse and meat processing wastewater[J]. Science of the Total Environment, 2019, 686, 681- 708. doi: 10.1016/j.scitotenv.2019.05.295
4	Feng Yan , Yang Shumin , Xia Liu , et al. In-situ ion exchange electro-catalysis biological coupling(i-IEEBC) for simultaneously enhanced degradation of organic pollutants and heavy metals in electroplating wastewater[J]. Journal of Hazardous Materials, 2019, 364, 562- 570. doi: 10.1016/j.jhazmat.2018.10.068
5	Kishor R , Purchase D , Saratale G D , et al. Ecotoxicological and health concerns of persistent coloring pollutants of textile industry wastewater and treatment approaches for environmental safety[J]. Journal of Environmental Chemical Engineering, 2021, 9 (2): 105012. doi: 10.1016/j.jece.2020.105012
6	Kong Zhe , Li Lu , Xue Yi , et al. Challenges and prospects for the anaerobic treatment of chemical-industrial organic wastewater: A review[J]. Journal of Cleaner Production, 2019, 231, 913- 927. doi: 10.1016/j.jclepro.2019.05.233
7	Flemming H C , Wingender J , Szewzyk U , et al. Biofilms: An emergent form of bacterial life[J]. Nature Reviews Microbiology, 2016, 14 (9): 563- 575. doi: 10.1038/nrmicro.2016.94
8	Xu Juan , Sheng Guoping . Microbial extracellular polymeric substances(EPS) acted as a potential reservoir in responding to high concentrations of sulfonamides shocks during biological wastewater treatment[J]. Bioresource Technology, 2020, 313, 123654. doi: 10.1016/j.biortech.2020.123654
9	Li Yiyu , Zheng Ping , Zhang Meng , et al. Hydrophilicity/hydrophobicity of anaerobic granular sludge surface and their causes: An in situ research[J]. Bioresource Technology, 2016, 220, 117- 123. doi: 10.1016/j.biortech.2016.08.012
10	Li Wenwei , Yu Hanqing . Insight into the roles of microbial extracellular polymer substances in metal biosorption[J]. Bioresource Technology, 2014, 160, 15- 23. doi: 10.1016/j.biortech.2013.11.074
11	Wang Weigang , Yan Yuan , Wang Junjie , et al. Comparison and optimization of extraction methods of extracellular polymeric substances in anammox granules: From maintaining protein secondary structure perspective[J]. Chemosphere, 2020, 259, e127539. doi: 10.1016/j.chemosphere.2020.127539
12	Felz S , Vermeulen P , Van Loosdrecht M C M , et al. Chemical characterization methods for the analysis of structural extracellular polymeric substances(EPS)[J]. Water Research, 2019, 157, 201- 208. doi: 10.1016/j.watres.2019.03.068
13	Wang Binbin , Liu Xueting , Chen Jianmeng , et al. Composition and functional group characterization of extracellular polymeric substances(EPS) in activated sludge: The impacts of polymerization degree of proteinaceous substrates[J]. Water Research, 2018, 129, 133- 142. doi: 10.1016/j.watres.2017.11.008
14	Tian Xiangmiao , Shen Zhiqiang , Han Zhenfeng , et al. The effect of extracellular polymeric substances on exogenous highly toxic compounds in biological wastewater treatment: An overview[J]. Bioresource Technology Reports, 2019, 5, 28- 42. doi: 10.1016/j.biteb.2018.11.009
15	钱勇兴, 李中坚, 杨彬, 等. 胞外聚合物去除废水中污染物的机理探讨[J]. 工业水处理, 2016, 36 (7): 16- 21. URL
16	Adnan L A , Sathishkumar P , Mohd Yusoff A R , et al. Metabolites characterisation of laccase mediated Reactive Black 5 biodegradation by fast growing ascomycete fungus Trichoderma atroviride F03[J]. International Biodeterioration & Biodegradation, 2015, 104, 274- 282. URL
17	Kang Fuxing , Zhu Dongqiang . Abiotic reduction of 1, 3-dinitrobenzene by aqueous dissolved extracellular polymeric substances produced by microorganisms[J]. Journal of Environmental Quality, 2013, 42 (5): 1441- 1448. doi: 10.2134/jeq2012.0499
18	Wang Jinfeng , Liu Qiuju , Hu Haidong , et al. Insight into mature biofilm quorum sensing in full-scale wastewater treatment plants[J]. Chemosphere, 2019, 234, 310- 317. doi: 10.1016/j.chemosphere.2019.06.007
19	Mangwani N , Kumari S , Das S . Bacterial biofilms and quorum sensing: Fidelity in bioremediation technology[J]. Biotechnology and Genetic Engineering Reviews, 2016, 32 (1/2): 43- 73. URL
20	Maddela N R , Sheng B B , Yuan S S , et al. Roles of quorum sensing in biological wastewater treatment: A critical review[J]. Chemosphere, 2019, 221, 616- 629. doi: 10.1016/j.chemosphere.2019.01.064
21	Jia Fangxu , Yang Qing , Liu Xiuhong , et al. Stratification of extracellular polymeric substances(EPS) for aggregated anammox microorganisms[J]. Environmental Science & Technology, 2017, 51 (6): 3260- 3268. URL
22	Bi S Y , Sourjik V . Stimulus sensing and signal processing in bacterial chemotaxis[J]. Current Opinion in Microbiology, 2018, 45, 22- 29. URL
23	Jiang Ruhan , Wu Xiaoxiong , Xiao Yaqian , et al. Tween 20 regulate the function and structure of transmembrane proteins of Bacillus cereus: Promoting transmembrane transport of fluoranthene[J]. Journal of Hazardous Materials, 2021, 403, e123707. doi: 10.1016/j.jhazmat.2020.123707
24	Yi Li , Dong Xiao , Grenier D , et al. Research progress of bacterial quorum sensing receptors: Classification, structure, function and characteristics[J]. Science of The Total Environment, 2021, 763, 143031. doi: 10.1016/j.scitotenv.2020.143031
25	More T T , Yadav J S S , Yan S , et al. Extracellular polymeric substances of bacteria and their potential environmental applications[J]. Journal of Environmental Management, 2014, 144, 1- 25. URL
26	Shi Yahui , Huang Jinhui , Zeng Guangming , et al. Exploiting extracellular polymeric substances(EPS) controlling strategies for performance enhancement of biological wastewater treatments: An overview[J]. Chemosphere, 2017, 180, 396- 411. doi: 10.1016/j.chemosphere.2017.04.042
27	Le C C , Stuckey D C . Influence of feed composition on the monomeric structure free bacterial extracellular polysaccharides in anaerobic digestion[J]. Environmental Science & Technology, 2017, 51 (12): 7009- 7017.
28	Bolei J M , Pabst M , Neu T R , et al. Identification of glycoproteins isolated from extracellular polymeric substances of full-scale anammox granular sludge[J]. Environmental Science & Technology, 2018, 52 (22): 13127- 13135.
29	Xu Juan , Sheng Guoping , Ma Ying , et al. Roles of extracellular polymeric substances(EPS) in the migration and removal of sulfamethazine in activated sludge system[J]. Water Research, 2013, 47 (14): 5298- 5306. doi: 10.1016/j.watres.2013.06.009
30	Guo Hongxiao , Felz S , Lin Yuemei , et al. Structural extracellular polymeric substances determine the difference in digestibility between waste activated sludge and aerobic granules[J]. Water Research, 2020, 181, 115924. doi: 10.1016/j.watres.2020.115924
31	Wang Jinfeng , Ding Lili , Li Kan , et al. Estimation of spatial distribution of quorum sensing signaling in sequencing batch biofilm reactor(SBBR) biofilms[J]. Science of The Total Environment, 2018, 612, 405- 414. doi: 10.1016/j.scitotenv.2017.07.277
32	Sheng Guoping , Yu Hanqing , Li Xiaoyan . Extracellular polymeric substances(EPS) of microbial aggregates in biological wastewater treatment systems: A review[J]. Biotechnology Advances, 2010, 28 (6): 882- 894. doi: 10.1016/j.biotechadv.2010.08.001
33	Ye Jinshao , Yin Hua , Qiang Jing , et al. Biodegradation of anthracene by Aspergillus fumigatus[J]. Journal of Hazardous Materials, 2011, 185 (1): 174- 181. doi: 10.1016/j.jhazmat.2010.09.015
34	Haroune L , Saibi S , Bellenger J P , et al. Evaluation of the efficiency of Trametes hirsuta for the removal of multiple pharmaceutical compounds under low concentrations relevant to the environment[J]. Bioresource Technology, 2014, 171, 199- 202. doi: 10.1016/j.biortech.2014.08.036
35	Shou Weijun , Kang Fuxing , Huang Shuhan , et al. Substituted aromatic-facilitated dissemination of mobile antibiotic resistance genes via an antihydrolysis mechanism across an extracellular polymeric substance permeable barrier[J]. Environmental Science & Technology, 2019, 53 (2): 604- 613. URL
36	Zhu Yanxia , Cheng Jun , Zhang Ze , et al. Promoting extracellular polymeric substances to alleviate phenol toxicity in Arthrospira platensis at high carbon dioxide concentrations[J]. Journal of Cleaner Production, 2021, 290, 125167. doi: 10.1016/j.jclepro.2020.125167
37	Sourjik V , Wingreen N S . Responding to chemical gradients: Bacterial chemotaxis[J]. Current Opinion in Cell Biology, 2012, 24 (2): 262- 268. doi: 10.1016/j.ceb.2011.11.008
38	王慧, 胡金星, 秦智慧, 等. 细菌对有机污染物的趋化性及其对降解的影响[J]. 浙江大学学报(农业与生命科学版), 2017, 43 (6): 676- 684. URL
39	Yong Yangchun , Zhong Jianjiang . Regulation of aromatics biodegradation by rhl quorum sensing system through induction of catechol meta-cleavage pathway[J]. Bioresource Technology, 2013, 136, 761- 765. doi: 10.1016/j.biortech.2013.03.134
40	Parkinson J S , Hazelbauer G L , Falke J J . Signaling and sensory adaptation in Escherichia coli chemoreceptors: 2015 update[J]. Trends in Microbiology, 2015, 23 (5): 257- 266. doi: 10.1016/j.tim.2015.03.003
41	Ni Bin , Huang Zhou , Fan Zheng , et al. Comamonas testosteroni uses a chemoreceptor for tricarboxylic acid cycle intermediates to trigger chemotactic responses towards aromatic compounds[J]. Molecular Microbiology, 2013, 90 (4): 813- 823. doi: 10.1111/mmi.12400
42	Arora P K , Srivastava A , Singh V P . Degradation of 4-chloro-3-nitrophenol via a novel intermediate, 4-chlororesorcinol by Pseudomonas sp. JHN[J]. Scientific Reports, 2014, 4, 1- 6. URL
43	Gordillo F , Chavez F P , Jerez C A . Motility and chemotaxis of Pseudomonas sp. B4 towards polychlorobiphenyls and chlorobenzoates[J]. FEMS Microbiology Ecology, 2007, 60 (2): 322- 328. doi: 10.1111/j.1574-6941.2007.00293.x
44	Zusman D R , Scott A E , Yang Zhaomin , et al. Chemosensory pathways, motility and development in Myxococcus xanthus[J]. Nature Reviews Microbiology, 2007, 5 (11): 862- 872. doi: 10.1038/nrmicro1770
45	Kim J , Kang Yongsung , Choi O , et al. Regulation of polar flagellum genes is mediated by quorum sensing and FlhDC in Burkholderia glumae[J]. Molecular Microbiology, 2007, 64 (1): 165- 179. doi: 10.1111/j.1365-2958.2007.05646.x
46	Ringgaard S , Hubbard T , Mandlik A , et al. RpoS and quorum sensing control expression and polar localization of Vibrio cholerae chemotaxis cluster Ⅲ proteins in vitro and in vivo[J]. Molecular Microbiology, 2015, 97 (4): 660- 675. doi: 10.1111/mmi.13053
47	Fernandez N L , Hsueh B Y , Nhu N T Q , et al. Vibrio cholerae adapts to sessile and motile lifestyles by cyclic di-GMP regulation of cell shape[J]. Proceedings of the National Academy of Sciences of the United States of America, 2020, 117 (46): 29046- 29054. doi: 10.1073/pnas.2010199117
48	Krell T , Lacal J , Reyes-Darias J A , et al. Bioavailability of pollutants and chemotaxis[J]. Current Opinion in Biotechnology, 2013, 24 (3): 451- 456. doi: 10.1016/j.copbio.2012.08.011
49	Yang Jingyun , Chawla R , Rhee K Y , et al. Biphasic chemotaxis of Escherichia coli to the microbiota metabolite indole[J]. Proceedings of the National Academy of Sciences of the United States of America, 2020, 117 (11): 6114- 6120. URL
50	Wiener M C , Horanyi P S . How hydrophobic molecules traverse the outer membranes of gram-negative bacteria[J]. Proceedings of the National Academy of Sciences of the United States of America, 2011, 108 (27): 10929- 10930. doi: 10.1073/pnas.1106927108
51	Nickzad A , Lepine F , Deziel E . Quorum sensing controls swarming motility of Burkholderia glumae through regulation of rhamnolipids[J]. PloS one, 2015, 10 (6): 0128509. URL
52	Gorna H , Lawniczak L , Zgola-Grzeskowiak A , et al. Differences and dynamic changes in the cell surface properties of three Pseudomonas aeruginosa strains isolated from petroleum-polluted soil as a response to various carbon sources and the external addition of rhamnolipids[J]. Bioresource Technology, 2011, 102 (3): 3028- 3033. doi: 10.1016/j.biortech.2010.09.124
53	Pamp S J , Tolker-Nielsen T . Multiple roles of biosurfactants in structural biofilm development by Pseudomonas aeruginosa[J]. Journal of Bacteriology, 2007, 189 (6): 2531- 2539. doi: 10.1128/JB.01515-06
54	Libis V , Delepine B , Faulon J L . Sensing new chemicals with bacterial transcription factors[J]. Current Opinion in Microbiology, 2016, 33, 105- 112. doi: 10.1016/j.mib.2016.07.006
55	Khomenkov V G , Shevelev A B , Zhukov V G , et al. Organization of metabolic pathways and molecular-genetic mechanisms of xenobiotic biodegradation in microorganisms: A review[J]. Applied Biochemistry and Microbiology, 2008, 44 (2): 117- 135. doi: 10.1134/S0003683808020014
56	Aldrich T L , Chakrabarty A M . Transcriptional regulation, nucleotide sequence, and localization of the promoter of the catBC operon in Pseudomonas putida[J]. Journal of Bacteriology, 1988, 170 (3): 1297- 1304. doi: 10.1128/jb.170.3.1297-1304.1988
57	Yubero P , Poyatos J F . The impact of global transcriptional regulation on bacterial gene order[J]. iScience, 2020, 23 (4): e101029. doi: 10.1016/j.isci.2020.101029
58	Zhang Yinping , Wang Fang , Zhu Xiaoshu , et al. Extracellular polymeric substances govern the development of biofilm and mass transfer of polycyclic aromatic hydrocarbons for improved biodegradation[J]. Bioresource Technology, 2015, 193, 274- 280. doi: 10.1016/j.biortech.2015.06.110
59	Wang Zhikang , Hessler C M , Xue Zheng , et al. The role of extracellular polymeric substances on the sorption of natural organic matter[J]. Water Research, 2012, 46 (4): 1052- 1060. doi: 10.1016/j.watres.2011.11.077
60	Bai Leilei , Xu Huacheng , Wang Changhui , et al. Extracellular polymeric substances facilitate the biosorption of phenanthrene on cyanobacteria Microcystis aeruginosa[J]. Chemosphere, 2016, 162, 172- 180. doi: 10.1016/j.chemosphere.2016.07.063
61	Yan Zirun , Meng Huishan , Yang Xueyuan , et al. Insights into the interactions between triclosan(TCS) and extracellular polymeric substance(EPS) of activated sludge[J]. Journal of Environmental Management, 2019, 232, 219- 225. URL
62	王静, 张程程, 吉芳英, 等. 磺胺类抗生素与胞外聚合物作用的热化学机制[J]. 中国环境科学, 2017, (10): 160- 165. URL
63	Pi Shanshan , Li Ang , Cui Di , et al. Biosorption behavior and mechanism of sulfonamide antibiotics in aqueous solution on extracellular polymeric substances extracted from Klebsiella sp. J1[J]. Bioresource Technology, 2019, 272, 346- 350. doi: 10.1016/j.biortech.2018.10.054
64	Zhang Zhiqiang , Xia Siqing , Wang Xuejiang , et al. A novel biosorbent for dye removal: Extracellular polymeric substance(EPS) of Proteus mirabilis TJ-1[J]. Journal of Hazardous Materials, 2009, 163 (1): 279- 284. doi: 10.1016/j.jhazmat.2008.06.096
65	Li Na , Wei Dong , Sun Qunqun , et al. Fluorescent component and complexation mechanism of extracellular polymeric substances during dye wastewater biotreatment by anaerobic granular sludge[J]. Royal Society Open Science, 2018, 5 (2): 171445. doi: 10.1098/rsos.171445
66	包宜俊, 杨存满, 李颖, 等. 光谱法研究胞外聚合物与四溴双酚A的相互作用[J]. 中国环境科学, 2016, 36 (6): 1773- 1779. doi: 10.3969/j.issn.1000-6923.2016.06.026
67	刘鑫彤, 尹华, 彭辉, 等. 胞外聚合物对活性污泥吸附去除全氟辛烷磺酸(PFOS)的影响[J]. 环境科学, 2017, 38 (8): 3435- 3441. URL
68	姜春阳. 微生物胞外聚合物在多环芳烃降解酶释放过程中的作用[D]. 沈阳: 沈阳理工大学, 2015.
69	Wong Yuxing , Yu Jian . Laccase-catalyzed decolorization of synthetic dyes[J]. Water Research, 1999, 33 (16): 3512- 3520. doi: 10.1016/S0043-1354(99)00066-4
70	Liu Yinan , Zhang Feng , Li Jie , et al. Exclusive extracellular bioreduction of methyl orange by azo reductase-free Geobacter sulfurreducens[J]. Environmental Science & Technology, 2017, 51 (15): 8616- 8623. URL
71	Sun Shengli , Wang Yuxiao , Zang Tingting , et al. A biosurfactantproducing Pseudomonas aeruginosa S5 isolated from coking wastewater and its application for bioremediation of polycyclic aromatic hydrocarbons[J]. Bioresource Technology, 2019, 281, 421- 428. doi: 10.1016/j.biortech.2019.02.087
72	Hua Xiufu , Wu Zuojun , Zhang Hongxing , et al. Degradation of hexadecane by Enterobacter cloacae strain TU that secretes an exopolysaccharide as a bioemulsifier[J]. Chemosphere, 2010, 80 (8): 951- 956. doi: 10.1016/j.chemosphere.2010.05.002
73	Ross P D , Subramanian S . Thermodynamics of protein association reactions: Forces contributing to stability[J]. Biochemistry, 1981, 20 (11): 3096- 3102. doi: 10.1021/bi00514a017
74	Liu Qiuju , Wang Jinfeng , He Ruonan , et al. Bacterial assembly during the initial adhesion phase in wastewater treatment biofilms[J]. Water Research, 2020, 184, 116147. doi: 10.1016/j.watres.2020.116147
75	骆庆群, 杨洁明. 有气体溶解和没有气体溶解的水中石墨烯间的疏水引力[J]. 力学学报, 2016, (3): 714- 719. doi: 10.3969/j.issn.1006-6616.2016.03.025
76	Chen Yiqun , Wang Minli , Zhou Xinwei , et al. Sorption fractionation of bacterial extracellular polymeric substances(EPS) on mineral surfaces and associated effects on phenanthrene sorption to EPS-mineral complexes[J]. Chemosphere, 2021, 263, 128264. doi: 10.1016/j.chemosphere.2020.128264
77	Sheng Guoping , Zhang Menglin , Yu Hanqing . Characterization of adsorption properties of extracellular polymeric substances(EPS) extracted from sludge[J]. Colloids and Surfaces B: Biointerfaces, 2008, 62 (1): 83- 90. doi: 10.1016/j.colsurfb.2007.09.024
78	Yan Zirun , Zhu Yuying , Meng Huishan , et al. Insights into thermodynamic mechanisms driving bisphenol A(BPA) binding to extracellular polymeric substances(EPS) of activated sludge[J]. Science of The Total Environment, 2019, 677, 502- 510. doi: 10.1016/j.scitotenv.2019.04.413
79	Pimentel G C , Mcclellan A L . Hydrogen Bond[J]. Annual Review of Physical Chemistry, 2003, 22 (1): 347- 385.
80	Li Zhengwen , Wan Chunli , Liu Xiang , et al. Understanding of the mechanism of extracellular polymeric substances of aerobic granular sludge against tetracycline from the perspective of fluorescence properties[J]. Science of The Total Environment, 2021, 756, 144054. doi: 10.1016/j.scitotenv.2020.144054
81	Zhang Huiqun , Jia Yanyan , Khanal S K , et al. Understanding the role of extracellular polymeric substances on ciprofloxacin adsorption in aerobic sludge, anaerobic sludge, and sulfate-reducing bacteria sludge systems[J]. Environmental Science & Technology, 2018, 52 (11): 6476- 6486. URL
82	Xiao Yong , Zhang Enhua , Zhang Jingdong , et al. Extracellular polymeric substances are transient media for microbial extracellular electron transfer[J]. Science Advances, 2017, 3 (7): e1700623. doi: 10.1126/sciadv.1700623
83	Cao Danming , Xiao Xiang , Wu Yongmin , et al. Role of electricity production in the anaerobic decolorization of dye mixture by exoelectrogenic bacterium Shewanella oneidensis MR-1[J]. Bioresource Technology, 2013, 136, 176- 181. doi: 10.1016/j.biortech.2013.02.083
84	Stern N , Mejia J , He Shaomei , et al. Dual role of humic substances as electron donor and shuttle for dissimilatory iron reduction[J]. Environmental Science & Technology, 2018, 52 (10): 5691- 5699. URL
85	Yan Zaisheng, He Yuhong, Cai Haiyuan, et al. Interconnection of key microbial functional genes for enhanced benzo[a]pyrene biodegradation in sediments by microbial electrochemistry[J]. Environmental Science & Technology, 2017, 51(15): 8519-8529.
86	Zhang Chunfang , Katayama A . Humin as an electron mediator for microbial reductive dehalogenation[J]. Environmental Science & Technology, 2012, 46 (12): 6575- 6583. URL
87	Zhou Xinwei , Kang Fuxing , Qu Xiaolei , et al. Probing extracellular reduction mechanisms of Bacillus subtilis and Escherichia coli with nitroaromatic compounds[J]. Science of The Total Environment, 2020, 724, 138291. doi: 10.1016/j.scitotenv.2020.138291
88	Gianfreda L , Rao M A . Potential of extra cellular enzymes in remediation of polluted soils: A review[J]. Enzyme and Microbial Technology, 2004, 35, 339- 354. doi: 10.1016/j.enzmictec.2004.05.006
89	Zumstein M T , Helbling D E . Biotransformation of antibiotics: Exploring the activity of extracellular and intracellular enzymes derived from wastewater microbial communities[J]. Water Research, 2019, 155, 115- 123. doi: 10.1016/j.watres.2019.02.024
90	Haernvall K , Zitzenbacher S , Wallig K , et al. Hydrolysis of ionic phthalic acid based polyesters by wastewater microorganisms and their enzymes[J]. Environmental Science & Technology, 2017, 51 (8): 4596- 4605. URL
91	Karich A , Ullrich R , Scheibner K , et al. Fungal unspecific peroxygenases oxidize the majority of organic EPA priority pollutants[J]. Frontiers in Microbiology, 2017, 8, 1- 15. URL
92	Jia Chunyun , Li Peijun , Li Xiaojun , et al. Degradation of pyrene in soils by extracellular polymeric substances(EPS) extracted from liquid cultures[J]. Process Biochemistry, 2011, 46 (8): 1627- 1631. doi: 10.1016/j.procbio.2011.05.005
93	Zerva A , Zervakis G I , Christakopoulos P , et al. Degradation of olive mill wastewater by the induced extracellular ligninolytic enzymes of two wood-rot fungi[J]. Journal of Environmental Management, 2017, 203, 791- 798. doi: 10.1016/j.jenvman.2016.02.042
94	Xu Li , Chen Xiong , Li Huixin , et al. Characterization of the biosorption and biodegradation properties of Ensifer adhaerens: A potential agent to remove polychlorinated biphenyls from contaminated water[J]. Journal of Hazardous Materials, 2016, 302, 314- 322. doi: 10.1016/j.jhazmat.2015.09.066
95	Wang Jinfeng , Liu Qiuju , Li Xianhui , et al. In-situ monitoring AHLmediated quorum-sensing regulation of the initial phase of wastewater biofilm formation[J]. Environment International, 2020, 135, 105326. doi: 10.1016/j.envint.2019.105326
96	Huang Jinhui , Shi Yahui , Zeng Guangming , et al. Acyl-homoserine lactone-based quorum sensing and quorum quenching hold promise to determine the performance of biological wastewater treatments: An overview[J]. Chemosphere, 2016, 157, 137- 151. doi: 10.1016/j.chemosphere.2016.05.032
97	Huang Yili , Zeng Yanhua , Yu Zhiliang , et al. In silico and experimental methods revealed highly diverse bacteria with quorum sensing and aromatics biodegradation systems-A potential broad application on bioremediation[J]. Bioresource Technology, 2013, 148, 311- 316. doi: 10.1016/j.biortech.2013.08.155
98	Yong Yangchun , Zhong Jianjiang . N-Acylated homoserine lactone production and involvement in the biodegradation of aromatics by an environmental isolate of Pseudomonas aeruginosa[J]. Process Biochemistry, 2010, 45 (12): 1944- 1948. doi: 10.1016/j.procbio.2010.05.006
99	Chugani S , Greenberg E P . LuxR homolog-independent gene regulation by acyl-homoserine lactones in Pseudomonas aeruginosa[J]. Proceedings of the National Academy of Sciences of the United States of America, 2010, 107 (23): 10673- 10678. doi: 10.1073/pnas.1005909107
100	Zeng Yanhua , Cheng Keke , Cai Zhonghua , et al. Transcriptome analysis expands the potential roles of quorum sensing in biodegradation and physiological responses to microcystin[J]. Science of The Total Environment, 2021, 771, 145437. doi: 10.1016/j.scitotenv.2021.145437
101	Yong Yangchun , Wu Xiangyang , Sun Jianzhong , et al. Engineering quorum sensing signaling of Pseudomonas for enhanced wastewater treatment and electricity harvest: A review[J]. Chemosphere, 2015, 140, 18- 25. doi: 10.1016/j.chemosphere.2014.10.020
102	Valle A , Bailey M J , Whiteley A S , et al. N-acyl-l-homoserine lactones(AHLs) affect microbial community composition and function in activated sludge[J]. Environmental Microbiology, 2010, 6 (4): 424- 433. URL
103	Mangwani N , Kumari S , Das S . Effect of synthetic N-acylhomoserine lactones on cell-cell interactions in marine Pseudomonas and biofilm mediated degradation of polycyclic aromatic hydrocarbons[J]. Chemical Engineering Journal, 2016, 302, 172- 186. doi: 10.1016/j.cej.2016.05.042
104	Mangwani N , Kumari S , Das S . Involvement of quorum sensing genes in biofilm development and degradation of polycyclic aromatic hydrocarbons by a marine bacterium Pseudomonas aeruginosa N6P6[J]. Applied Microbiology and Biotechnology, 2015, 99 (23): 10283- 10297. doi: 10.1007/s00253-015-6868-7
105	Kang Y S , Park W . Contribution of quorum-sensing system to hexadecane degradation and biofilm formation in Acinetobacter sp. strain DR1[J]. Journal of Applied Microbiology, 2010, 109 (5): 1650- 1659. URL
106	Tang Xi , Guo Yongzhao , Wu Shanshan , et al. Metabolomics uncovers the regulatory pathway of acyl-homoserine lactones based quorum sensing in anammox consortia[J]. Environmental Science & Technology, 2018, 52, 2206- 2216. URL
107	Gu Yue , Tian Jianjun , Zhang Yue , et al. Dissecting signal molecule AI-2 mediated biofilm formation and environmental tolerance in Lactobacillus plantarum[J]. Journal of Bioscience and Bioengineering, 2021, 131 (2): 153- 160. doi: 10.1016/j.jbiosc.2020.09.015
108	Jatt A N , Tang Kaihao , Liu Jiwen , et al. Quorum sensing in marine snow and its possible influence on production of extracellular hydrolytic enzymes in marine snow bacterium Pantoea ananatis B9[J]. FEMS Microbiology Ecology, 2015, 91 (2): 1- 13. URL
109	Nasser W , Bouillant M L , Salmond G , et al. Characterization of the Erwinia chrysanthemi expI-expR locus directing the synthesis of two N-acyl-homoserine lactone signal molecules[J]. Molecular Microbiology, 2010, 29 (6): 1391- 1405. URL
110	Swift S , Lynch M J , Fish L , et al. Quorum sensing-dependent regulation and blockade of exoprotease production in Aeromonas hydrophila[J]. Infection and Immunity, 1999, 67 (10): 5192- 5199. doi: 10.1128/IAI.67.10.5192-5199.1999
111	Reis R S , Pereira A G , Neves B C , et al. Gene regulation of rhamnolipid production in Pseudomonas aeruginosa-A review[J]. Bioresource Technology, 2011, 102 (11): 6377- 6384. doi: 10.1016/j.biortech.2011.03.074
112	Chen Aina , Huang Yili . Acyl homoserine lactone based quorum sensing affects phenanthrene removal by Novosphingobium pentaromativorans US6-1 through altering cell surface properties[J]. International Biodeterioration & Biodegradation, 2020, 147, 104841. URL
113	Laganenka L , Colin R , Sourjik V . Chemotaxis towards autoinducer 2 mediates autoaggregation in Escherichia coli[J]. Nature Communications, 2016, 7 (1): e12984. doi: 10.1038/ncomms12984
114	张祥武. 大肠杆菌密度感应调节子C(QseC)在鞭毛调控基因flhDC mRNA表达中的作用研究[D]. 昆明: 昆明医科大学, 2015.
115	Ghosh S, Qureshi A, Purohit H J. Aromatic compounds and biofilms: Regulation and interlinking of metabolic pathways in bacteria[M]∥Arora P K. Microbial metabolism of xenobiotic compounds. Singapore: Springer Nature Singapore Pte Ltd., 2019, 10: 145-164.
116	Mangwani N , Dash H R , Chauhan A , et al. Bacterial quorum sensing: Functional features and potential applications in biotechnology[J]. Journal of Molecular Microbiology and Biotechnology, 2012, 22 (4): 215- 227. doi: 10.1159/000341847
117	Mangwani N , Shukla S K , Kumari S , et al. Characterization of Stenotrophomonas acidaminiphila NCW-702 biofilm for implication in the degradation of polycyclic aromatic hydrocarbons[J]. Journal of Applied Microbiology, 2015, 117 (4): 1012- 1024. URL
118	Molin S , Tolker-Nielsen T . Gene transfer occurs with enhanced efficiency in biofilms and induces enhanced stabilisation of the biofilm structure[J]. Current Opinion in Biotechnology, 2003, 14 (3): 255- 261. doi: 10.1016/S0958-1669(03)00036-3
119	Rui Bin , Shen Tie , Zhou Hong , et al. A systematic investigation of Escherichia coli central carbon metabolism in response to superoxide stress[J]. Bmc Systems Biology, 2010, 4 (122): 1- 12. URL
120	卢航. 两种鞘酯单胞菌的rsh基因缺失对群体感应信号分子积累的影响[D]. 杭州: 浙江大学, 2018.
121	Karpinets T V , Pelletier D , Pan C , et al. Phenotype fingerprinting suggests the involvement of single-genotype consortia in degradation of aromatic compounds by Rhodopseudomonas palustris[J]. Plos One, 2009, 4 (2): e4615. doi: 10.1371/journal.pone.0004615
122	Yang Liang , Barken K B , Skindersoe M E , et al. Effects of iron on DNA release and biofilm development by Pseudomonas aeruginosa[J]. Microbiology, 2007, 153 (5): 1318- 1328. doi: 10.1099/mic.0.2006/004911-0
123	Wang Jinfeng , Liu Qiuju , Dong Deyuan , et al. AHLs-mediated quorum sensing threshold and its response towards initial adhesion of wastewater biofilms[J]. Water Research, 2021, 194, 116925. doi: 10.1016/j.watres.2021.116925
124	Sadowsky M J , Tong Z K , Souza M , et al. AtzC is a new member of the amidohydrolase protein superfamily and is homologous to other atrazine-metabolizing enzymes[J]. Journal of Bacteriology, 1998, 180 (1): 152- 158. doi: 10.1128/JB.180.1.152-158.1998
125	徐苹, 杨晶, 陆丽兰, 等. 密度感应系统对弗氏志贺菌生长竞争能力的影响[J]. 遗传, 2015, 37 (5): 487- 493. URL
126	Zhang Bing , Li Wei , Guo Yuan , et al. A sustainable strategy for effective regulation of aerobic granulation: Augmentation of the signaling molecule content by cultivating AHL-producing strains[J]. Water Research, 2020, 169, 115193. doi: 10.1016/j.watres.2019.115193

污染物	微生物来源	EPS与污染物作用机制	研究方法/手段	参考文献
菲、芘（PAHs）	Micrococcus sp. PHE9、Mycobacterium sp. NJS-P	吸附：疏水作用；胞外PS、PN水解高分子有机物	通过促进生物膜形成促进PAHs降解	〔58〕
菲、芘（PAHs）	Microcystis aeruginosa	吸附：化学吸附；静电+疏水作用；自发放热	吸附动力学；3D-EEM （三维荧光光谱）	〔60〕
三氯生（TCS）	活性污泥	吸附：放热过程；PN通过氢键和疏水作用主导	热力学；3D-EEM	〔61〕
磺胺二甲基嘧啶（SMZ）	枯草芽孢杆菌	吸附：PN疏水性作用	等温滴定微量热	〔62〕
磺胺甲恶唑（SMX）、SM1、磺胺嘧啶（SDZ）	Klebsiella sp. J1	吸附：化学吸附（色氨酸与酪氨酸的疏水作用）	等温线，动力学；3D-EEM	〔63〕
BB54（染料）	Proteus mirabilis TJ-1	吸附：内输运机制；羧基、羟基和氨基等起作用	吸附动力学，等温线；FTIR；扫描电子显微镜	〔64〕
亚甲基蓝（MB）	厌氧颗粒污泥	吸附：类PN和类HS；羟基和氨基等起作用	吸附动力学；3D-EEM；FTIR	〔65〕
四溴双酚A（TBBPA）	活性污泥	吸附：结合后PN结构变化	3D-EEM；FTIR；同步荧光光谱	〔66〕
全氟辛烷磺酸（PFOS）	活性污泥	吸附：化学吸附；静电力；羟基、羧基和氨基	等温线，吸附动力学；FTIR；X射线光电子能谱	〔67〕
天然有机物（NOM）：HS、HA	Pseudomonas aeruginosa、P. putida	吸附：架桥>疏水；二价离子桥接NOM和EPS上负电子官能团	FTIR；疏水性，Zeta势	〔59〕
芘和苯并[a]芘	毛霉和分支杆菌	生物降解：胞外环羟基化双加氧酶	十二烷基硫酸钠聚丙烯酰胺凝胶电泳；蛋白质谱；DNA测序；3D-EEM	〔68〕
活性黑5（RB5）	Trichoderma atroviride F03	生物降解：胞外漆酶，活性最大达到5.8 U/mL	紫外-可见分光光度法（UV-Vis）；电化学；木质素酶活性；薄层色谱，GC-MS，FTIR	〔16〕
蒽醌、偶氮和靛蓝	Trametes versicolor	生物降解：胞外漆酶直接氧化蒽醌，与偶氮相互作用	UV-Vis；漆酶活性	〔69〕
1，3-二硝基苯	E. coli DH5a	还原：鼠李糖半缩醛、脂多糖不饱和脂肪链和还原酚醛官能团-还原剂；醌类-电子传导	FTIR、EEM、13C NMR、Tollen’s测试	〔17〕
偶氮染料甲基橙（MO）	Geobacter sulfurreducens PCA	电化学还原：关键外膜蛋白OmcB、OmcC和OmcE参与胞外还原过程	动力学；质量与电子平衡、荧光探针和蛋白酶K处理实验	〔70〕

污染物	微生物来源	EPS与污染物作用机制	研究方法/手段	参考文献
菲、芘（PAHs）	Micrococcus sp. PHE9、Mycobacterium sp. NJS-P	吸附：疏水作用；胞外PS、PN水解高分子有机物	通过促进生物膜形成促进PAHs降解	〔58〕
菲、芘（PAHs）	Microcystis aeruginosa	吸附：化学吸附；静电+疏水作用；自发放热	吸附动力学；3D-EEM （三维荧光光谱）	〔60〕
三氯生（TCS）	活性污泥	吸附：放热过程；PN通过氢键和疏水作用主导	热力学；3D-EEM	〔61〕
磺胺二甲基嘧啶（SMZ）	枯草芽孢杆菌	吸附：PN疏水性作用	等温滴定微量热	〔62〕
磺胺甲恶唑（SMX）、SM1、磺胺嘧啶（SDZ）	Klebsiella sp. J1	吸附：化学吸附（色氨酸与酪氨酸的疏水作用）	等温线，动力学；3D-EEM	〔63〕
BB54（染料）	Proteus mirabilis TJ-1	吸附：内输运机制；羧基、羟基和氨基等起作用	吸附动力学，等温线；FTIR；扫描电子显微镜	〔64〕
亚甲基蓝（MB）	厌氧颗粒污泥	吸附：类PN和类HS；羟基和氨基等起作用	吸附动力学；3D-EEM；FTIR	〔65〕
四溴双酚A（TBBPA）	活性污泥	吸附：结合后PN结构变化	3D-EEM；FTIR；同步荧光光谱	〔66〕
全氟辛烷磺酸（PFOS）	活性污泥	吸附：化学吸附；静电力；羟基、羧基和氨基	等温线，吸附动力学；FTIR；X射线光电子能谱	〔67〕
天然有机物（NOM）：HS、HA	Pseudomonas aeruginosa、P. putida	吸附：架桥>疏水；二价离子桥接NOM和EPS上负电子官能团	FTIR；疏水性，Zeta势	〔59〕
芘和苯并[a]芘	毛霉和分支杆菌	生物降解：胞外环羟基化双加氧酶	十二烷基硫酸钠聚丙烯酰胺凝胶电泳；蛋白质谱；DNA测序；3D-EEM	〔68〕
活性黑5（RB5）	Trichoderma atroviride F03	生物降解：胞外漆酶，活性最大达到5.8 U/mL	紫外-可见分光光度法（UV-Vis）；电化学；木质素酶活性；薄层色谱，GC-MS，FTIR	〔16〕
蒽醌、偶氮和靛蓝	Trametes versicolor	生物降解：胞外漆酶直接氧化蒽醌，与偶氮相互作用	UV-Vis；漆酶活性	〔69〕
1，3-二硝基苯	E. coli DH5a	还原：鼠李糖半缩醛、脂多糖不饱和脂肪链和还原酚醛官能团-还原剂；醌类-电子传导	FTIR、EEM、13C NMR、Tollen’s测试	〔17〕
偶氮染料甲基橙（MO）	Geobacter sulfurreducens PCA	电化学还原：关键外膜蛋白OmcB、OmcC和OmcE参与胞外还原过程	动力学；质量与电子平衡、荧光探针和蛋白酶K处理实验	〔70〕

污染物	AHLs种类	群感系统	作用主体	AHLs调控EPS	AHLs调控污染物去除	参考文献
苯酚	外源：C6-HSL、3O-C6-HSL 内源：BHL、HHL	LuxI/LuxR RhlI/RhlR	反应器污泥/铜绿假单胞菌	促进表面活性剂产生	苯酚去除率提高27%	〔102〕；〔98〕
苯酚、苯甲酸、菲	内源+外源：BHL	RhlI/RhlR	铜绿假单菌CGMCC	促进nahR、nahH转录→胞外酶（邻苯二酚-2，3-双加氧酶）产生	提高污染物去除率20%	〔39〕
邻氨基苯甲酸盐	内源：C4-HSL、3OC12-HSL；外源：C10-HSL	LasI/LasR RhlI/RhlR QscR	铜绿假单胞菌	促进antR、ant和cat操纵子转录→胞外酶（降解邻氨基苯甲酸盐）产生	促进邻氨基苯甲酸盐降解	〔99〕
菲、芘	外源：3OC8-HSL、3OC12-HSL		假产碱假单胞菌	促进鼠李糖脂产生	提高污染物去除率6%~16%	〔103〕
菲、芘	内源：C4-HSL、3OC12-HSL	LasI/LasR RhlI/RhlR	铜绿假单胞菌	促进胞外多糖产生	促进菲、芘生物降解	〔104〕
十六烷	外源：C12-AHLs	aqsI/aqsR	不动杆菌DR1	调控胞外PN产生	促进十六烷降解	〔105〕

污染物	AHLs种类	群感系统	作用主体	AHLs调控EPS	AHLs调控污染物去除	参考文献
苯酚	外源：C6-HSL、3O-C6-HSL 内源：BHL、HHL	LuxI/LuxR RhlI/RhlR	反应器污泥/铜绿假单胞菌	促进表面活性剂产生	苯酚去除率提高27%	〔102〕；〔98〕
苯酚、苯甲酸、菲	内源+外源：BHL	RhlI/RhlR	铜绿假单菌CGMCC	促进nahR、nahH转录→胞外酶（邻苯二酚-2，3-双加氧酶）产生	提高污染物去除率20%	〔39〕
邻氨基苯甲酸盐	内源：C4-HSL、3OC12-HSL；外源：C10-HSL	LasI/LasR RhlI/RhlR QscR	铜绿假单胞菌	促进antR、ant和cat操纵子转录→胞外酶（降解邻氨基苯甲酸盐）产生	促进邻氨基苯甲酸盐降解	〔99〕
菲、芘	外源：3OC8-HSL、3OC12-HSL		假产碱假单胞菌	促进鼠李糖脂产生	提高污染物去除率6%~16%	〔103〕
菲、芘	内源：C4-HSL、3OC12-HSL	LasI/LasR RhlI/RhlR	铜绿假单胞菌	促进胞外多糖产生	促进菲、芘生物降解	〔104〕
十六烷	外源：C12-AHLs	aqsI/aqsR	不动杆菌DR1	调控胞外PN产生	促进十六烷降解	〔105〕

[1]	靳亚斌, 徐甜甜, 赵德中, 张乐, 万振杰, 张高明. Bi₂O₃基复合光催化剂的制备及其在水处理中的应用进展[J]. 工业水处理, 2025, 45(3): 10-21.
[2]	赵杰, 吴俊峰, 窦艳艳, VIKTOROVICH Fedorov Svyatoslav, 王现丽, 刘彪, 朱新锋, 毛艳丽, 高宏斌. 单原子活化过硫酸盐降解有机污染物研究进展[J]. 工业水处理, 2024, 44(7): 38-46.
[3]	贾仲琪, 卫金秋, 宿辉, 刘宪斌. 纳米零价铁耦合高级氧化技术处理废水的研究进展[J]. 工业水处理, 2024, 44(7): 9-18.
[4]	杨震, 李阳, 杨世迎. 零价铝还原联合过硫酸盐氧化高效处理DDNP工业废水[J]. 工业水处理, 2024, 44(4): 98-104.
[5]	李棒, 田腾飞, 吴俊峰, 朱新锋, 韩昌睿, 崔璐雪, 张霞, 贠延滨. 膜蒸馏处理脱硫废水过程中污染物组成、分布及形成机理[J]. 工业水处理, 2024, 44(2): 139-146.
[6]	解鹤, 唐彬, 陈兵兵, 李佳利, 詹海鹃. 掺杂型钴基钙钛矿的合成及光催化降解有机废水[J]. 工业水处理, 2024, 44(12): 131-137.
[7]	姚有智, 华飞, 吴康, 叶舟, 张忠亮, 钱程. 石墨烯基复合材料光催化降解废水有机污染物的研究进展[J]. 工业水处理, 2024, 44(11): 1-8.
[8]	刘国煜, 曹向禹, 郑建华. 钴基氧化物光催化剂在降解有机污染物中的研究进展[J]. 工业水处理, 2024, 44(11): 27-34.
[9]	于会娟, 张英杰, 权泓, 高翠萍, 田春梅, 朱丹, 杨荣彬. 氮化碳改性光催化材料在水污染治理中的应用[J]. 工业水处理, 2023, 43(8): 38-47.
[10]	代朝猛,贾辰炎,胡佳俊,刘曙光,张亚雷,游学极. 过氧乙酸高级氧化技术机理及影响因素研究进展[J]. 工业水处理, 2023, 43(1): 1-9.
[11]	冉春秋,葛辉,赵钰,张倩,冯志静. 余热余碱活化过硫酸盐处理精制棉黑液[J]. 工业水处理, 2022, 42(3): 106-113.
[12]	孙家宁, 孙韶华, 宋娜, 辛晓东, 赵清华, 贾瑞宝. 高级氧化技术去除水中溴系阻燃剂的研究进展[J]. 工业水处理, 2022, 42(2): 19-26.
[13]	张楠, 麻微微, 黄浩勇, 牛越, 施雪卿. 好氧颗粒污泥技术处理石化废水的研究进展[J]. 工业水处理, 2022, 42(2): 35-44.
[14]	李莹, 刘强, 项玮, 李倩囡. 基于群体感应相关菌群分析的HMBR膜污染控制研究[J]. 工业水处理, 2022, 42(11): 122-126.
[15]	雷少庭,耿金菊,吴刚,于清淼,许柯,张徐祥,任洪强. 基于荧光预测污水中痕量有机污染物衰减研究进展[J]. 工业水处理, 2021, 41(6): 35-47.

选择文件类型/文献管理软件名称

选择包含的内容