供水管网生物膜中微生物种间相互作用及其影响因素综述

祝泽兵; 裴云燕; 单莉莉; 徐思扬; 许琳燕; 袁一星

doi:10.13205/j.hjgc.202306028

供水管网生物膜中微生物种间相互作用及其影响因素综述

doi: 10.13205/j.hjgc.202306028

1. 华东交通大学土木建筑学院, 南昌 330013;
2. 哈尔滨工业大学城市水资源与水环境国家重点实验室, 哈尔滨 150090

基金项目:

国家自然科学基金青年科学基金（51608198）；江西省青年科学基金（20212BAB214006）；江西省教育厅面上基金（GJJ2200652，GJJ2200645）

详细信息

作者简介:
祝泽兵(1983-),男,工学博士,副教授,主要研究方向为水质安全保障技术。hitzzb@126.com

通讯作者:
祝泽兵(1983-),男,工学博士,副教授,主要研究方向为水质安全保障技术。hitzzb@126.com

计量
- 文章访问数: 196
- HTML全文浏览量: 19
- PDF下载量: 6
- 被引次数: 0
出版历程
- 收稿日期: 2022-04-21
- 网络出版日期: 2023-09-02

MICROBIAL INTERSPECIFIC INTERACTION AND ITS INFLUENCING FACTORS IN BIOFILM OF DRINKING WATER DISTRIBUTION SYSTEMS: A REVIEW

1. School of Civil Engineering and Architecture, East China Jiao Tong University, Nanchang 330013, China;
2. State Key Laboratory of Urban Water Resource and Environment, Harbin Institute of Technology, Harbin 150090, China

摘要

摘要: 微生物生长显著影响供水管网中的水质安全，供水管网中微生物主要存在于生物膜中。近年来，随着高通量基因测序技术的广泛使用，多物种生物膜的研究越来越受到重视。微生物种间相互作用等因素会影响多物种生物膜的形成，而控制供水管网生物膜中微生物的生长对于限制水传播疾病至关重要。从胞外聚合物、生物膜调节基因、抗生素和环境适应性等方面讨论了多物种生物膜形成的初步机制，综述了供水管网多物种生物膜中微生物之间的种间相互作用（竞争、协作及中性作用），并从群体感应信号分子、代谢共生和交互共生等方面探讨了种间相互作用机制及其对生物膜形成的影响，从生物学及非生物学角度总结了微生物种间相互作用的影响因素。基于上述分析，提出研究微生物种间相互作用机制以及多物种生物膜形成因素，为限制生物膜生长、保障饮用水水质安全提供新思路。
- 供水管网 /
- 多物种 /
- 生物膜 /
- 种间相互作用
Abstract: Microbial growth significantly affects water safety in drinking water distribution systems, where microorganisms are mainly found in biofilms. In recent years, with the development of high-throughput genetic sequencing, more and more attention has been paid to multi-species biofilms. Factors such as microbial interspecific interactions can affect the formation of multi-species biofilms, and controlling the growth of microorganisms in the biofilms of the drinking water distribution systems is essential to limit water-borne diseases. In this paper, the preliminary mechanisms of multi-species biofilms formation are discussed from the aspects of extracellular polymeric substances, biofilm-regulated genes, antibiotics and environmental adaptability, and then the interspecific interactions (competition, synergism and neutralism) among microorganisms in multi-species biofilms in drinking water distribution systems are reviewed. The mechanisms of interspecific interactions and their effects on biofilm formation are discussed from the aspects of quorum sensing signal molecules, metabolic commensalism and cross-feeding. Finally, the influencing factors of microbial interspecific interactions are summarized from the biological and non-biological perspectives. Based on the above analysis, this paper proposes to study the mechanisms of microbial interspecific interactions and the factors of multi-species biofilms formation, which provides a new idea for limiting biofilm growth and ensuring the safety of drinking water quality.
- drinking water distribution system /
- multi-species /
- biofilm /
- interspecific interaction

HTML全文

参考文献(107)

[1]	COLLIER S A, DENG L, ADAM E A, et al. Estimate of burden and direct healthcare cost of infectious waterborne disease in the united states[J]. Emerging Infectious Diseases, 2021, 27(1):140-149.
[2]	WANG T, SUN D L, ZHANG Q, et al. China's drinking water sanitation from 2007 to 2018:a systematic review[J]. Science of the Total Environment, 2021, 757:143923.
[3]	ZHU Z B, SHAN L L, LI X S, et al. Effects of interspecific interactions on biofilm formation potential and chlorine resistance:evaluation of dual-species biofilm observed in drinking water distribution systems[J]. Journal of Water Process Engineering, 2020, 38:101564.
[4]	BIMAKR F, GINIGE M P, KAKSONEN A H, et al. Assessing graphite and stainless-steel for electrochemical sensing of biofilm growth in chlorinated drinking water systems[J]. Sensors and Actuators B-Chemical, 2018, 277:526-534.
[5]	景双艳, 魏莲花. 金黄色葡萄球菌生物膜形成及其与持留菌关系研究进展[J]. 中国生物制品学杂志, 2021, 34(1):102-105.
[6]	CHAN Y, WU X H, CHIENG B W, et al. Superhydrophobic nanocoatings as intervention against biofilm-associated bacterial infections[J]. Nanomaterials, 2021, 11(4):1046.
[7]	REUBEN R C, ROY P C, SARKAR S L, et al. Multispecies interactions in biofilms and implications to safety of drinking water distribution system[J]. Microbiology and Biotechnology Letters, 2019, 47(4):473-486.
[8]	MAKOVCOVA J, BABAK V, KULICH P, et al. Dynamics of mono- and dual-species biofilm formation and interactions between Staphylococcus aureus and Gram-negative bacteria[J]. Microbial Biotechnology, 2017, 10(4):819-832.
[9]	WINGENDER J, FLEMMING H C. Biofilms in drinking water and their role as reservoir for pathogens[J]. International Journal of Hygiene and Environmental Health, 2011, 214(6):417-423.
[10]	SAXENA P, JOSHI Y, RAWAT K, et al. Biofilms:architecture, resistance, quorum sensing and control mechanisms[J]. Indian Journal of Microbiology, 2019, 59(1):3-12.
[11]	KVICH L, BURMOLLE M, BJARNSHOLT T, et al. Do mixed-species biofilms dominate in chronic infections?-need forin situ visualization of bacterial organization[J]. Frontiers in Cellular and Infection Microbiology, 2020, 10:396.
[12]	ZHU Z B, SHAN L L, ZHANG X Y, et al. Effects of bacterial community composition and structure in drinking water distribution systems on biofilm formation and chlorine resistance[J]. Chemosphere, 2021, 264:128410.
[13]	SAMROT A V, MOHAMED A A, FARADJEVA E, et al. Mechanisms and impact of biofilms and targeting of biofilms using bioactive compounds:a review[J]. Medicina-Lithuania, 2021, 57(8):839.
[14]	PREST E I, SCHAAP P G, BESMER M D, et al. Dynamic hydraulics in a drinking water distribution system influence suspended particles and turbidity, but not microbiology[J]. Water, 2021, 13(1):109.
[15]	MAKRIS K C, ANDRA S S, BOTSARIS G. Pipe scales and biofilms in drinking-water distribution systems:undermining finished water quality[J]. Critical Reviews in Environmental Science and Technology, 2014, 44(13):1477-1523.
[16]	AFONSO A C, GOMES I B, SAAVEDRA M J, et al. Bacterial coaggregation in aquatic systems[J]. Water Research, 2021, 196:117037.
[17]	ELIAS S, BANIN E. Multi-species biofilms:living with friendly neighbors[J]. Fems Microbiology Reviews, 2012, 36(5):990-1004.
[18]	RICE S A, WUERTZ S, KJELLEBERG S. Next-generation studies of microbial biofilm communities[J]. Microbial Biotechnology, 2016, 9(5):677-680.
[19]	ZHANG W, SUN J, DING W, et al. Extracellular matrix-associated proteins form an integral and dynamic system during Pseudomonas aeruginosa biofilm development[J]. Frontiers in Cellular and Infection Microbiology, 2015, 5:40.
[20]	PRATT L A, KOLTER R. Genetic analysis of Escherichia coli biofilm formation:roles of flagella, motility, chemotaxis and type Ⅰ pili[J]. Molecular Microbiology, 1998, 30(2):285-293.
[21]	CASEY A, FOX E M, SCHMITZ-ESSER S, et al. Transcriptome analysis of Listeria monocytogenes exposed to biocide stress reveals a multi-system response involving cell wall synthesis, sugar uptake, and motility[J]. Frontiers in Microbiology, 2014, 5:68.
[22]	MADSEN J S, RODER H L, RUSSEL J, et al. Coexistence facilitates interspecific biofilm formation in complex microbial communities[J]. Environmental Microbiology, 2016, 18(8):2565-2574.
[23]	KARYGIANNI L, REN Z, KOO H, et al. Biofilm matrixome:extracellular components in structured microbial communities[j]. trends in microbiology, 2020, 28(8):668-681.
[24]	WANG R. Biofilms and meat safety:a mini-review[J]. Journal of Food Protection, 2019, 82(1):120-127.
[25]	DUANIS-ASSAF D, DUANIS-ASSAF T, ZENG G H, et al. Cell wall associated protein TasA provides an initial binding component to extracellular polysaccharides in dual-species biofilm[J]. Scientific Reports, 2018, 8:9350.
[26]	MARINCOLA G, JASCHKOWITZ G, KIENINGER A K, et al. Plasmid-chromosome crosstalk in Staphylococcus aureus:a horizontally acquired transcription regulator controls polysaccharide intercellular adhesin-mediated biofilm formation[J]. Frontiers in Cellular and Infection Microbiology, 2021, 11:660702.
[27]	DOMKA J, LEE J T, BANSAL T, et al. Temporal gene-expression in Escherichia coli K-12 biofilms[J]. Environmental Microbiology, 2007, 9(2):332-346.
[28]	WANG M J, ZHAO L, WU H, et al. Cladodionen is a potential quorum sensing inhibitor against pseudomonas aeruginosa[J]. Marine Drugs, 2020, 18(4):205.
[29]	WIEDENBECK J, COHAN F M. Origins of bacterial diversity through horizontal genetic transfer and adaptation to new ecological niches[J]. Fems Microbiology Reviews, 2011, 35(5):957-976.
[30]	MADSEN J S, BURMOLLE M, HANSEN L H, et al. The interconnection between biofilm formation and horizontal gene transfer[J]. Fems Immunology and Medical Microbiology, 2012, 65(2):183-195.
[31]	LUO A, WANG F, SUN D, et al. Formation, development, and cross-species interactions in biofilms[J]. Frontiers in Microbiology, 2022, 12:757327.
[32]	ZHANG Y J, ZHANG Y Y, LIU L N, et al. Impacts of antibiotics on biofilm bacterial community and disinfection performance on simulated drinking water supply pipe wall[J]. Environmental Pollution, 2021, 288:117736.
[33]	PENESYAN A, GILLINGS M, PAULSEN I T. Antibiotic discovery:combatting bacterial resistance in cells and in biofilm communities[J]. Molecules, 2015, 20(4):5286-5298.
[34]	HUO L X, ZHAO S H, SHI B Y, et al. Bacterial community change and antibiotic resistance promotion after exposure to sulfadiazine and the role of UV/H₂O₂-GAC treatment[J]. Chemosphere, 2021, 283:131214.
[35]	ABISADO R G, BENOMAR S, KLAUS J R, et al. Bacterial quorum sensing and microbial community interactions[J]. Mbio, 2018, 9(3):e02331-17.
[36]	DI SOMMA A, MORETTA A, CANE C, et al. Antimicrobial and antibiofilm peptides[J]. Biomolecules, 2020, 10(4):652.
[37]	SHAHROUR H, FERRER-ESPADA R, DANDACHE I, et al. AMPs as anti-biofilm agents for human therapy and prophylaxis[M]. Switzerland:Springer International Publishing Ag, 2019:257-279.
[38]	MAGALHAES A P, JORGE P, PEREIRA M O. Pseudomonas aeruginosa and Staphylococcus aureus communication in biofilm infections:insights through network and database construction[J]. Critical Reviews in Microbiology, 2019, 45(5/6):712-728.
[39]	LIU W Z, RUSSEL J, BURMOLLE M, et al. Micro-scale intermixing:a requisite for stable and synergistic co-establishment in a four-species biofilm[J]. Isme Journal, 2018, 12(8):1940-1951.
[40]	HABIMANA O, HEIR E, LANGSRUD S, et al. Enhanced surface colonization by Escherichia coli O157:H7 in biofilms formed by an acinetobacter calcoaceticus isolate from meat-processing environments[J]. Applied and Environmental Microbiology, 2010, 76(13):4557-4559.
[41]	DAI D, RASKIN L, XI C. The effect of interactions between a bacterial strain isolated from drinking water and a pathogen surrogate on biofilms formation diverged under static vs flow conditions[J]. Journal of Applied Microbiology, 2017, 123(6):1614-1627.
[42]	ZHANG W, SILEIKA T, PACKMAN A I. Effects of fluid flow conditions on interactions between species in biofilms[J]. Fems Microbiology Ecology, 2013, 84(2):344-354.
[43]	CORRE M H, DELAFONT V, LEGRAND A, et al. Exploiting the richness of environmental waterborne bacteria species to find natural legionella pneumophila competitors[J]. Frontiers in Microbiology, 2019, 9:3360.
[44]	MAES S, de REU K, van WEYENBERG S, et al. Pseudomonas putida as a potential biocontrol agent against Salmonella Java biofilm formation in the drinking water system of broiler houses[J]. Bmc Microbiology, 2020, 20(1):373.
[45]	SIMOES L C, SIMOES M, VIEIRA M J. Biofilm interactions between distinct bacterial genera isolated from drinking water[J]. Applied and Environmental Microbiology, 2007, 73(19):6192-6200.
[46]	LI J, CHEN X J, LIN J F, et al. Antibiotic intervention redisposes bacterial interspecific interacting dynamics in competitive environments[J]. Environmental Microbiology, 2021, 23(12):7432-7444.
[47]	SADIQ F A, BURMOLLE M, HEYNDRICKX M, et al. Community-wide changes reflecting bacterial interspecific interactions in multispecies biofilms[J]. Critical Reviews in Microbiology, 2021, 47(3):338-358.
[48]	GHOUL M, MITRI S. The ecology and evolution of microbial competition[J]. Trends in Microbiology, 2016, 24(10):833-845.
[49]	祝泽兵. 供水管网中的耐氯菌群及其耐氯机制研究[D]. 哈尔滨:哈尔滨工业大学, 2015.
[50]	LI Q, LIU L, GUO A L, et al. Formation of multispecies biofilms and their resistance to disinfectants in food processing environments:a review[J]. Journal of Food Protection, 2021, 84(12):2071-2083.
[51]	de GRANDI A Z, PINTO U M, DESTRO M T. Dual-species biofilm of Listeria monocytogenes and Escherichia coli on stainless steel surface[J]. World Journal of Microbiology & Biotechnology, 2018, 34(4):61.
[52]	EVANS C R, KEMPES C P, PRICE-WHELAN A, et al. Metabolic heterogeneity and cross-Feeding in bacterial multicellular systems[J]. Trends in Microbiology, 2020, 28(9):732-743.
[53]	ESTRELA S, TRISOS C H, BROWN S P. From metabolism to ecology:cross-feeding interactions shape the balance between polymicrobial conflict and mutualism[J]. American Naturalist, 2012, 180(5):566-576.
[54]	YI L, DONG X, 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.
[55]	ZHAO X, YU Z, DING T. Quorum-sensing regulation of antimicrobial resistance in bacteria[J]. Microorganisms, 2020, 8(3):425.
[56]	SHARMA A, SINGH P, SARMAH B K, et al. Quorum sensing:its role in microbial social networking[J]. Research in Microbiology, 2020, 171(5/6):159-164.
[57]	ZHOU L, ZHANG Y, GE Y, et al. Regulatory mechanisms and promising applications of quorum sensing-inhibiting agents in control of bacterial biofilm formation[J]. Frontiers in Microbiology, 2020, 11:589640.
[58]	RAJU MADDELA N, SHENG B, YUAN S, et al. Roles of quorum sensing in biological wastewater treatment:a critical review[J]. Chemosphere, 2019, 221:616-629.
[59]	成婷婷, 冯媛, 王崇刚. 鲍曼不动杆菌群体感应抑制剂与生物膜形成的研究进展[J]. 中国感染与化疗杂志, 2022, 22(1):125-128.
[60]	DIXON E F, HALL R A. Noisy neighbourhoods:quorum sensing in fungal-polymicrobial infections[J]. Cellular Microbiology, 2015, 17(10):1431-1441.
[61]	MUKHERJEE S, BOSSIER B L. Bacterial quorum sensing in complex and dynamically changing environments[J]. Nature Reviews Microbiology, 2019, 17(6):371-382.
[62]	KAUR A, CAPALASH N, SHARMA P. Communication mechanisms in extremophiles:exploring their existence and industrial applications[J]. Microbiological Research, 2019, 221:15-27.
[63]	KOLENBRANDER P E, ANDERSEN R N, BLEHERT D S, et al. Communication among oral bacteria[J]. Microbiology and molecular biology reviews:MMBR, 2002, 66(3):486-505.
[64]	PANDE S, KAFTAN F, LANG S, et al. Privatization of cooperative benefits stabilizes mutualistic cross-feeding interactions in spatially structured environments[J]. Isme Journal, 2016, 10(6):1413-1423.
[65]	SUN Z P, KOFFEL T, STUMP S M, et al. Microbial cross-feeding promotes multiple stable states and species coexistence, but also susceptibility to cheaters[J]. Journal of Theoretical Biology, 2019, 465:63-77.
[66]	WINTERMUTE E H, SILVER P A. Emergent cooperation in microbial metabolism[J]. Molecular Systems Biology, 2010, 6:407.
[67]	SHOU W, RAM S, VILAR J M G. Synthetic cooperation in engineered yeast populations[J]. Proceedings of the National Academy of Sciences of the United States of America, 2007, 104(6):1877-1882.
[68]	YUAN L, SADIQ F A, WANG N, et al. Recent advances in understanding the control of disinfectant-resistant biofilms by hurdle technology in the food industry[J]. Critical Reviews in Food Science and Nutrition, 2021, 61(22):3876-3891.
[69]	SALGAR-CHAPARRO S J, LEPKOVA K, POJTANABUNTOENG T, et al. Nutrient level determines biofilm characteristics and subsequent impact on microbial corrosion and biocide effectiveness[J]. Applied and Environmental Microbiology, 2020, 86(7):e02885-19.
[70]	JAMAL M, AHMAD W, ANDLEEB S, et al. Bacterial biofilm and associated infections[J]. Journal of the Chinese Medical Association, 2018, 81(1):7-11.
[71]	da CRUZ NIZER W S, INKOVSKIY V, OVERHAGE J. Surviving reactive chlorine stress:responses of gram-negative bacteria to hypochlorous acid[J]. Microorganisms, 2020, 8(8):1220.
[72]	BANSAL M, DHOWLAGHAR N, NANNAPANENI R, et al. Decreased biofilm formation by planktonic cells of Listeria monocytogenes in the presence of sodium hypochlorite[J]. Food Microbiology, 2021, 96:103714.
[73]	BYUN K H, HAN S H, YOON J W, et al. Efficacy of chlorine-based disinfectants (sodium hypochlorite and chlorine dioxide) on Salmonella enteritidis planktonic cells, biofilms on food contact surfaces and chicken skin[J]. Food Control, 2021, 123:107838.
[74]	LUO X R, ZHANG B P, LU Y H, et al. Advances in application of ultraviolet irradiation for biofilm control in water and wastewater infrastructure[J]. Journal of Hazardous Materials, 2022, 421:126682.
[75]	WAAK M B, HOZALSKI R M, HALLE C, et al. Comparison of the microbiomes of two drinking water distribution systemswith and without residual chloramine disinfection[J]. Microbiome, 2019, 7:87.
[76]	XUE Z, LEE W H, COBURN K M, et al. Selective reactivity of monochloramine with extracellular matrix components affects the disinfection of biofilm and detached clusters[J]. Environmental Science & Technology, 2014, 48(7):3832-3839.
[77]	罗志逢. 供水管道生物膜生长的影响因素与控制方法研究[D]. 杭州:浙江大学, 2016.
[78]	周玲玲. 给水管网中生物膜及硝化作用控制[D]. 哈尔滨:哈尔滨工业大学, 2010.
[79]	LIPPONEN M T T, SUUTARI M H, MARTIKAINEN P J. Occurrence of nitrifying bacteria and nitrification in Finnish drinking water distribution systems[J]. Water Research, 2002, 36(17):4319-4329.
[80]	LI N, LI X, YANG Y L, et al. Secondary disinfection ensures biosafety of domestic hot water and its impact on biofilm bacterial community[J]. Desalination and Water Treatment, 2020, 173:186-196.
[81]	FENG C, ZHU N, LI Y, et al. Microbial characteristics of the combined ozone and tea polyphenols or sodium hypochlorite disinfection in the pipe network[J]. Water, 2021, 13(13):1835.
[82]	ZHANG K W, HU H, CHEN G. Mechanisms of microbial disinfectant resistance[J]. Progress in Biochemistry and Biophysics, 2022, 49(1):34-47.
[83]	LIU G, ZHANG Y, KNIBBE W J, et al. Potential impacts of changing supply-water quality on drinking water distribution:a review[J]. Water Research, 2017, 116:135-148.
[84]	LIU G, LUT M C, VERBERK J, et al. A comparison of additional treatment processes to limit particle accumulation and microbial growth during drinking water distribution[J]. Water Research, 2013, 47(8):2719-2728.
[85]	van DER WIELEN P W J J, VAN DER KOOIJ D. Effect of water composition, distance and season on the adenosine triphosphate concentration in unchlorinated drinking water in the Netherlands[J]. Water Research, 2010, 44(17):4860-4867.
[86]	RATZKE C, BARRERE J, GORE J. Strength of species interactions determines biodiversity and stability in microbial communities[J]. Nature Ecology & Evolution, 2020, 4(3):376-397.
[87]	LIU Y Y, SHAN R R, CHEN G W, et al. Linking flow velocity-regulated EPS production with early-stage biofilm formation in drinking water distribution systems[J]. Water Supply, 2020, 20(4):1253-1263.
[88]	逯清清. 给水管网环境条件对生物膜形成和水质的影响[D]. 合肥:合肥工业大学, 2016.
[89]	LAUTENSCHLAGER K, HWANG C, LIU W T, et al. A microbiology-based multi-parametric approach towards assessing biological stability in drinking water distribution networks[J]. Water Research, 2013, 47(9):3015-3025.
[90]	WANG H, MASTERS S, HONG Y J, et al. Effect of disinfectant, water age, and pipe material on occurrence and persistence of legionella, mycobacteria, pseudomonas aeruginosa, and two amoebas[J]. Environmental Science & Technology, 2012, 46(21):11566-11574.
[91]	YE C S, XIAN X X, BAO R H, et al. Recovery of microbiological quality of long-term stagnant tap water in university buildings during the COVID-19 pandemic[J]. Science of the Total Environment, 2022, 806:150616.
[92]	黄保国. 给水管网流速对水质和生物膜种群结构的影响[D]. 合肥:合肥工业大学, 2018.
[93]	LEARBUCH K L G, SMIDT H, van DER WIELEN P W J J. Influence of pipe materials on the microbial community in unchlorinated drinking water and biofilm[J]. Water Research, 2021, 194:116922.
[94]	池年平, 董秉直, 姚若虚. 给水管网中微生物研究进展[J]. 水处理技术, 2010, 36(2):29-32.
[95]	ZHANG X Y, LIN T, JIANG F C, et al. Impact of pipe material and chlorination on the biofilm structure and microbial communities[J]. Chemosphere, 2022, 289:133218.
[96]	李相宜, 赵蓓, 游晓旭, 等. 供水管道管材的特性及应用综述[J]. 净水技术, 2021, 40(7):52-59.
[97]	XU X X, LIU S M, SMITH K, et al. An overview on corrosion of iron and steel components in reclaimed water supply systems and the mechanisms involved[J]. Journal of Cleaner Production, 2020, 276:124079.
[98]	LIU L, HU Q Y, LE Y, et al. Chlorination-mediated EPS excretion shapes early-stage biofilm formation in drinking water systems[J]. Process Biochemistry, 2017, 55:41-48.
[99]	XU H J, LIU Y. Reduced microbial attachment by D-amino acid-inhibited AI-2 and EPS production[J]. Water Research, 2011, 45(17):5796-5804.
[100]	JAYATHILAKE P G, JANA S, RUSHTON S, et al. Extracellular polymeric substance production and aggregated bacteria colonization influence the competition of microbes in biofilms[J]. Frontiers in Microbiology, 2017, 8:1865.
[101]	WANG Z, KIM J, SEO Y. Influence of bacterial extracellular polymeric substances on the formation of carbonaceous and nitrogenous disinfection byproducts[J]. Environmental Science & Technology, 2012, 46(20):11361-11369.
[102]	MANDAL A, MANDAL R K, YANG Y, et al. In vitro characterization of chicken gut bacterial isolates for probiotic potentials[J]. Poultry Science, 2021, 100(2):1083-1092.
[103]	马晓春, 代军, 徐磊, 等. 鲍曼不动杆菌生物膜形成机制研究进展[J]. 中国感染与化疗杂志, 2018, 18(1):124-128.
[104]	WANG F, DENG L, HUANG F, et al. Flagellar motility is critical for salmonella entericaserovar typhimurium biofilm development[J]. Frontiers in Microbiology, 2020, 11:1695.
[105]	WOOD T K, BARRIOS A F G, HERZBERG M, et al. Motility influences biofilm architecture in Escherichia coli[J]. Applied Microbiology and Biotechnology, 2006, 72(2):361-367.
[106]	PANG X, YUK H G. Effects of the colonization sequence of Listeria monocytogenes and Pseudomonas fluorescens on survival of biofilm cells under food-related stresses and transfer to salmon[J]. Food Microbiology, 2019, 82:142-150.
[107]	GUTTENPLAN S B, KEARNS D B. Regulation of flagellar motility during biofilm formation[J]. Fems Microbiology Reviews, 2013, 37(6):849-871.