Sewage fungus

Sewage fungus^[1] (also known as undesirable river biofilms, URBs) is a polymicrobial biofilm (a microbial mat) that proliferates in saprobic rivers^[2] and has been frequently used as a bioindicator^[3]^[4] of organic river pollution for the past century.^[5] Its presence has been strongly associated with discharges of untreated or inadequately treated sewage,^[6]^[7]^[8]^[9] yet its presence extends beyond these areas, with contributors including airport de-ice fluid runoff,^[10]^[11]^[12] papermill effluents,^[13] and agricultural runoff.^[14]^[15]

The name "sewage fungus" is somewhat of a misnomer,^[5] as these growths are not primarily fungal in nature. Instead, they are complex polymicrobial mats bound within a matrix of extracellular polymeric substances. The bacterial taxa most frequently associated with this phenomenon include Sphaerotilus natans, Zoogloea spp., Beggiatoa spp., and Rhodoferax spp.^[11]^[16]^[17]

Environmental Impacts

In addition to being a bioindicator of organic pollution in rivers and playing a vital role utilizing excess organic carbon in fluvial systems, sewage fungus causes significant ecological impacts through direct and indirect ecological pathways.

Sewage fungus thrives in the low dissolved oxygen (DO) environment of an organically polluted river.^[4]^[17]^[18]^[19] Whilst DO is required for sewage fungus growth, it readily outcompetes other benthic organisms at low DO,^[20]^[21]^[22] quickly smothering riverbeds, greatly altering the benthic habitat for invertebrates^[23]^[24] and fish spawning.^[2]^[25]^[26] The dominating growth of sewage fungus also reduces hyporheic exchange flows, an important part of a rivers self-cleaning system.^[27]^[28] Similar river biofilms are also reported to accumulate heavy metals^[29]^[30] and other toxic substances.^[31] within their matrix causing ecological impacts throughout the food web.^[32]^[33] As a heterotroph, sewage fungus uses considerably higher DO than an aquatic macrophyte of the same mass,^[34] it can maintain DO concentrations below thresholds required for other organisms. Once sewage fungus becomes established, it is difficult to remove,^[35] unless all sources of organic nutrients (pollution) are addressed, causing a further loss in biodiversity^[36] and other flora and fauna^[37]^[38] in the river. These ecological impacts and the striking visible presence of sewage fungus growth on a riverbed further affects people's perceptions and use of rivers.^[7]^[39]

Microbial composition

Sewage fungus is a type of microbial mat, the specific composition of which is affected by the available nutrients (especially organic carbon sources) and the environmental drivers of each unique occurrence. However, several key taxa are reported as highly frequent and dominant within sewage fungus.

Sphaerotilus natans, has been strongly associated with this phenomenon since it was first studied ^[5] and continues to be regarded as a key sewage fungus organism.^[11]^[12] Consequently, Sphaerotilus has been used seemingly synonymously with sewage fungus and a series of laboratory studies use S. natans as sewage fungus.^[25]^[40]^[41]

Other key taxa include the bacteria Zoogloea spp., Beggiatoa spp., Thiothrix spp., Flavobacterium spp., and Flexibacter spp..^[16]^[17] However, fungi (e.g., Leptomitus lacteus, Geotrichum candidum, and Fusarium aquaeductuum), algae (e.g., Cladophora glomerata) along with archaea and protozoa (e.g., Carchesium polypinum) also form integral and important pasts of the biofilm.

Recent genomic studies of sewage fungus composition have identified some of these taxa within airport de-icer implicated occurrences but have also identified new taxa not previously associated with sewage fungus: Rhodoferax as a dominant component of sewage fungus,^[11] and the presence of Thiothrix.^[12]

Drivers of sewage fungus

Alongside the complex nutrient utilisation requirements of sewage fungus, there are several key environmental drivers including substrate type, flow velocity, temperature, shading/sunlight, and water chemistry (e.g. pH).

Flowing water is a requirement for sewage fungus growth, to provide a constant replenishment of nutrients.^[2]^[4]^[42] However, if the velocity of the river is too fast, then growths are scoured away, especially on more readily mobilised substrates. In turn, the specific flow of the river shapes the morphotype and structure of the biofilm.^[43] Intrinsically the substrate affects the upper limit of flow as more stable riverbeds are less readily mobilised in periods of higher flows. Surfaces such as large cobbles, anthropogenic litter (e.g., bricks), and concrete channels facilitate excellent sewage fungus growth, whereas fine sediments and gravel provide a less stable substrate.

References

^ Exton, Ben; Hassard, Francis; Medina-Vaya, Angel; Grabowski, Robert C. (April 2024). "Undesirable river biofilms: The composition, environmental drivers, and occurrence of sewage fungus". Ecological Indicators. 161: 111949. doi:10.1016/j.ecolind.2024.111949. ISSN 1470-160X.
^ ^a ^b ^c Curtis, E.J.C. (May 1969). "Sewage fungus: Its nature and effects". Water Research. 3 (5): 289–311. Bibcode:1969WatRe...3..289C. doi:10.1016/0043-1354(69)90084-0.
^ "Freshwater Biology and Ecology Handbook". Foundation for Water Research. Retrieved 2022-08-15.
^ ^a ^b ^c Quinn, McFarlane (1985). "Sewage fungus as a monitor of water quality". Biological Monitoring in Freshwaters: Proceedings of a Seminar.
^ ^a ^b ^c Butcher, R.W. (August 1932). "Contribution to our knowledge of the ecology of sewage fungus". Transactions of the British Mycological Society. 17 (1–2): 112–IN4. doi:10.1016/S0007-1536(32)80029-X.
^ Chonova, Teofana; Labanowski, Jérôme; Cournoyer, Benoit; Chardon, Cécile; Keck, François; Laurent, Élodie; Mondamert, Leslie; Vasselon, Valentin; Wiest, Laure; Bouchez, Agnès (April 2018). "River biofilm community changes related to pharmaceutical loads emitted by a wastewater treatment plant". Environmental Science and Pollution Research. 25 (10): 9254–9264. doi:10.1007/s11356-017-0024-0. ISSN 0944-1344. PMID 28884270. S2CID 3997467.
^ ^a ^b Curtis, E.J.C.; Harrington, D.W. (June 1971). "The occurrence of sewage fungus in rivers in the United Kingdom". Water Research. 5 (6): 281–290. Bibcode:1971WatRe...5..281C. doi:10.1016/0043-1354(71)90173-4.
^ Harrison, Heukelekian (1958). "Slime Infestation: Literature Review". Sewage and Industrial Wastes. 30 (10): 1278–1302. JSTOR 25033719 – via JSTOR.
^ Hammond, Peter; Suttie, Michael; Lewis, Vaughan T.; Smith, Ashley P.; Singer, Andrew C. (2021-03-11). "Detection of untreated sewage discharges to watercourses using machine learning". npj Clean Water. 4 (1). doi:10.1038/s41545-021-00108-3. ISSN 2059-7037. S2CID 232173162.
^ Mericas, Dean; Sturman, Paul; Lutz, Michelle; Corsi, Steve; Cieciek, Chris; Boltz, Josh; Morgenroth, Eberhard; Airport Cooperative Research Program; Transportation Research Board (2014-11-03). Understanding Microbial Biofilms in Receiving Waters Impacted by Airport Deicing Activities. Washington, D.C.: Transportation Research Board. doi:10.17226/22262. ISBN 978-0-309-30809-0.
^ ^a ^b ^c ^d Exton, Benjamin; Hassard, Francis; Medina Vaya, Angel; Grabowski, Robert C. (2023-03-01). "Polybacterial shift in benthic river biofilms attributed to organic pollution – a prospect of a new biosentinel?". Hydrology Research. 54 (3): 348–359. doi:10.2166/nh.2023.114. ISSN 0029-1277. S2CID 257347315.
^ ^a ^b ^c Nott, Michelle A.; Driscoll, Heather E.; Takeda, Minoru; Vangala, Mahesh; Corsi, Steven R.; Tighe, Scott W. (2020-01-22). Loiselle, Steven Arthur (ed.). "Advanced biofilm analysis in streams receiving organic deicer runoff". PLOS ONE. 15 (1): e0227567. Bibcode:2020PLoSO..1527567N. doi:10.1371/journal.pone.0227567. ISSN 1932-6203. PMC 6975536. PMID 31968006.
^ Roberts, J.C. (1978). "Sewage fungus growth in rivers below papermill discharges". New Processes for Wastewater Treatment and Recovery: 140–158.
^ Rutt, G.P.; Pickering, T.D.; Reynolds, N.R.M. (1993). "The impact of livestock-farming on welsh streams: The development and testing of a rapid biological method for use in the assessment and control of organic pollution from farms". Environmental Pollution. 81 (3): 217–228. doi:10.1016/0269-7491(93)90205-3. PMID 15091808.
^ Seager, J.; Jones, F.; Rutt, G. (February 1992). "Assessment and Control of Farm Pollution". Water and Environment Journal. 6 (1): 48–53. doi:10.1111/j.1747-6593.1992.tb00737.x. ISSN 1747-6585.
^ ^a ^b Geatches, Gething, Rutt (2014). 'Sewage fungus': A field and microscopic guide. Environment Agency and Natural Resources Wales.{{cite book}}: CS1 maint: multiple names: authors list (link)
^ ^a ^b ^c Curtis, E.J.C.; Curds, C.R. (December 1971). "Sewage fungus in rivers in the United Kingdom: The slime community and its constituent organisms". Water Research. 5 (12): 1147–1159. Bibcode:1971WatRe...5.1147C. doi:10.1016/0043-1354(71)90080-7.
^ Gray, N.F.; Hunter, Christine A. (January 1985). "Heterotrophic slimes in Irish rivers". Water Research. 19 (6): 685–691. doi:10.1016/0043-1354(85)90113-7.
^ Hickey, Christopher W. (November 1988). "River oxygen uptake and respiratory decay of sewage fungus biofilms". Water Research. 22 (11): 1375–1380. Bibcode:1988WatRe..22.1375H. doi:10.1016/0043-1354(88)90093-0.
^ Adeola, Samuel; Revitt, Michael; Shutes, Brian; Garelick, Hemda; Jones, Huw; Jones, Clive (2009-01-12). "Constructed Wetland Control of BOD Levels in Airport Runoff". International Journal of Phytoremediation. 11 (1): 1–10. doi:10.1080/15226510802363220. ISSN 1522-6514. S2CID 85164686.
^ Turnbull, D.A.; Bevan, J.R. (1995). "The impact of airport de-icing on a river: The case of the Ouseburn, Newcastle upon Tyne". Environmental Pollution. 88 (3): 321–332. doi:10.1016/0269-7491(95)93446-7. PMID 15091545.
^ United States Environmental Protection Agency (US EPA) (2000). "Preliminary Data Summary - Airport Deicing Operations (Revised)" (PDF).
^ Hirsch (1958). "Biological evaluation of organic pollution of New Zealand streams". New Zealand Journal of Science. 1 (4): 500–553.
^ Hynes (1960). The biology of polluted waters. Liverpool University Press.
^ ^a ^b Curtis, E.J.C.; Delves-Broughton, J.; Harrington, D.W. (June 1971). "Sewage fungus: studies of Sphaerotilus slimes using laboratory recirculating channels". Water Research. 5 (6): 267–279. Bibcode:1971WatRe...5..267C. doi:10.1016/0043-1354(71)90172-2.
^ Smith, Lloyd L.; Kramer, Robert H. (July 1963). "Survival of Walleye Eggs in Relation to Wood Fibers and Sphaerotilus natans in the Rainy River, Minnesota". Transactions of the American Fisheries Society. 92 (3): 220–234. doi:10.1577/1548-8659(1963)92[220:SOWEIR]2.0.CO;2. ISSN 0002-8487.
^ Magliozzi, Chiara; Grabowski, Robert C.; Packman, Aaron I.; Krause, Stefan (2018-11-30). "Toward a conceptual framework of hyporheic exchange across spatial scales". Hydrology and Earth System Sciences. 22 (12): 6163–6185. doi:10.5194/hess-22-6163-2018. ISSN 1607-7938.
^ Lewandowski, Jörg; Arnon, Shai; Banks, Eddie; Batelaan, Okke; Betterle, Andrea; Broecker, Tabea; Coll, Claudia; Drummond, Jennifer; Gaona Garcia, Jaime; Galloway, Jason; Gomez-Velez, Jesus; Grabowski, Robert; Herzog, Skuyler; Hinkelmann, Reinhard; Höhne, Anja (2019-10-25). "Is the Hyporheic Zone Relevant beyond the Scientific Community?". Water. 11 (11): 2230. doi:10.3390/w11112230. hdl:20.500.11850/382125. ISSN 2073-4441.
^ Decho, Alan W.; Visscher, Pieter T.; Reid, R. Pamela (2005), "Production and cycling of natural microbial exopolymers (EPS) within a marine stromatolite", Geobiology: Objectives, Concepts, Perspectives, Elsevier, pp. 71–86, doi:10.1016/b978-0-444-52019-7.50008-5, ISBN 978-0-444-52019-7, retrieved 2023-10-10
^ Flemming, Hans-Curt; Wingender, Jost (September 2010). "The biofilm matrix". Nature Reviews Microbiology. 8 (9): 623–633. doi:10.1038/nrmicro2415. ISSN 1740-1526. PMID 20676145. S2CID 28850938.
^ Flemming, Hans-Curt; Wingender, Jost; Szewzyk, Ulrich; Steinberg, Peter; Rice, Scott A.; Kjelleberg, Staffan (September 2016). "Biofilms: an emergent form of bacterial life". Nature Reviews Microbiology. 14 (9): 563–575. doi:10.1038/nrmicro.2016.94. ISSN 1740-1526. PMID 27510863. S2CID 4384131.
^ Friedman, Barry A.; Dugan, Patrick R. (May 1968). "Identification of Zoogloea species and the Relationship to Zoogloeal Matrix and Floc Formation". Journal of Bacteriology. 95 (5): 1903–1909. doi:10.1128/jb.95.5.1903-1909.1968. ISSN 0021-9193. PMC 252226. PMID 5650090.
^ Patrick, F; Loutit, M (1976). "Passage of metals in effluents, through bacteria to higher organisms". Water Research. 10 (4): 333–335. Bibcode:1976WatRe..10..333P. doi:10.1016/0043-1354(76)90176-7.
^ Gray, N (1987). Sewage Fungus in Irish Rivers: A Guide to Identification, Evaluation and Control. Trinity College, University of Dublin.
^ McKinney, Ross E. (2004-03-11). Environmental Pollution Control Microbiology (0 ed.). CRC Press. doi:10.1201/9780203025697. ISBN 978-1-135-52187-5.
^ Hartwell, S. Ian; Jordahl, David M.; Evans, Joyce E.; May, Eric B. (August 1995). "Toxicity of aircraft de-icer and anti-icer solutions to aquatic organisms". Environmental Toxicology and Chemistry. 14 (8): 1375–1386. doi:10.1002/etc.5620140813.
^ Pillard, David A (February 1995). "Comparative toxicity of formulated glycol deicers and pure ethylene and propylene glycol to Ceriodaphnia dubia and Pimephales promelas". Environmental Toxicology and Chemistry. 14 (2): 311–315. doi:10.1002/etc.5620140217.
^ Pillard, D. A.; DuFresne, D. L. (1999-07-01). "Toxicity of Formulated Glycol Deicers and Ethylene and Propylene Glycol to Lactuca sativa, Lolium perenne, Selenastrum capricornutum, and Lemna minor". Archives of Environmental Contamination and Toxicology. 37 (1): 29–35. doi:10.1007/s002449900486. ISSN 0090-4341. PMID 10341039. S2CID 12747431.
^ Gray, N. F. (November 1985). "Heterotrophic Slimes in Flowing Waters". Biological Reviews. 60 (4): 499–548. doi:10.1111/j.1469-185X.1985.tb00621.x. ISSN 1464-7931. S2CID 83810613.
^ Dias, FF; Dondero, NC; Finstein, MS (August 1968). "Attached Growth of Sphaerotilus and Mixed Populations in a Continuous-flow Apparatus". Applied Microbiology. 16 (8): 1191–99. doi:10.1128/am.16.8.1191-1199.1968. ISSN 0003-6919. PMC 547617. PMID 4877499.
^ Dias, FF; Heukelekian, H (September 1967). "Utilization of Inorganic Nitrogen Compounds by Sphaerotilus natans Growing in a Continuous-Flow Apparatus". Applied Microbiology. 15 (5): 1083–86. doi:10.1128/am.15.5.1083-1086.1967. ISSN 0003-6919. PMC 547145. PMID 16349722.
^ Phaup, JD (October 1968). "The biology of Sphaerotilus species". Water Research. 2 (9): 597–614. Bibcode:1968WatRe...2..597P. doi:10.1016/0043-1354(68)90065-1.
^ Besemer, K; Singer, G; Limberger, R; Chlup, AK; Hochedlinger, G; Hödl, I; Baranyi, C; Battin, TJ (August 2007). "Biophysical Controls on Community Succession in Stream Biofilms". Applied and Environmental Microbiology. 73 (15): 4966–74. Bibcode:2007ApEnM..73.4966B. doi:10.1128/AEM.00588-07. ISSN 0099-2240. PMC 1951047. PMID 17557861.

[1] Exton, Ben; Hassard, Francis; Medina-Vaya, Angel; Grabowski, Robert C. (April 2024). "Undesirable river biofilms: The composition, environmental drivers, and occurrence of sewage fungus". Ecological Indicators. 161: 111949. doi:10.1016/j.ecolind.2024.111949. ISSN 1470-160X.

[:0-2] Curtis, E.J.C. (May 1969). "Sewage fungus: Its nature and effects". Water Research. 3 (5): 289–311. Bibcode:1969WatRe...3..289C. doi:10.1016/0043-1354(69)90084-0.

[3] "Freshwater Biology and Ecology Handbook". Foundation for Water Research. Retrieved 2022-08-15.

[:1-4] Quinn, McFarlane (1985). "Sewage fungus as a monitor of water quality". Biological Monitoring in Freshwaters: Proceedings of a Seminar.

[:2-5] Butcher, R.W. (August 1932). "Contribution to our knowledge of the ecology of sewage fungus". Transactions of the British Mycological Society. 17 (1–2): 112–IN4. doi:10.1016/S0007-1536(32)80029-X.

[6] Chonova, Teofana; Labanowski, Jérôme; Cournoyer, Benoit; Chardon, Cécile; Keck, François; Laurent, Élodie; Mondamert, Leslie; Vasselon, Valentin; Wiest, Laure; Bouchez, Agnès (April 2018). "River biofilm community changes related to pharmaceutical loads emitted by a wastewater treatment plant". Environmental Science and Pollution Research. 25 (10): 9254–9264. doi:10.1007/s11356-017-0024-0. ISSN 0944-1344. PMID 28884270. S2CID 3997467.

[:3-7] Curtis, E.J.C.; Harrington, D.W. (June 1971). "The occurrence of sewage fungus in rivers in the United Kingdom". Water Research. 5 (6): 281–290. Bibcode:1971WatRe...5..281C. doi:10.1016/0043-1354(71)90173-4.

[8] Harrison, Heukelekian (1958). "Slime Infestation: Literature Review". Sewage and Industrial Wastes. 30 (10): 1278–1302. JSTOR 25033719 – via JSTOR.

[9] Hammond, Peter; Suttie, Michael; Lewis, Vaughan T.; Smith, Ashley P.; Singer, Andrew C. (2021-03-11). "Detection of untreated sewage discharges to watercourses using machine learning". npj Clean Water. 4 (1). doi:10.1038/s41545-021-00108-3. ISSN 2059-7037. S2CID 232173162.

[10] Mericas, Dean; Sturman, Paul; Lutz, Michelle; Corsi, Steve; Cieciek, Chris; Boltz, Josh; Morgenroth, Eberhard; Airport Cooperative Research Program; Transportation Research Board (2014-11-03). Understanding Microbial Biofilms in Receiving Waters Impacted by Airport Deicing Activities. Washington, D.C.: Transportation Research Board. doi:10.17226/22262. ISBN 978-0-309-30809-0.

[:4-11] Exton, Benjamin; Hassard, Francis; Medina Vaya, Angel; Grabowski, Robert C. (2023-03-01). "Polybacterial shift in benthic river biofilms attributed to organic pollution – a prospect of a new biosentinel?". Hydrology Research. 54 (3): 348–359. doi:10.2166/nh.2023.114. ISSN 0029-1277. S2CID 257347315.

[:5-12] Nott, Michelle A.; Driscoll, Heather E.; Takeda, Minoru; Vangala, Mahesh; Corsi, Steven R.; Tighe, Scott W. (2020-01-22). Loiselle, Steven Arthur (ed.). "Advanced biofilm analysis in streams receiving organic deicer runoff". PLOS ONE. 15 (1): e0227567. Bibcode:2020PLoSO..1527567N. doi:10.1371/journal.pone.0227567. ISSN 1932-6203. PMC 6975536. PMID 31968006.

[13] Roberts, J.C. (1978). "Sewage fungus growth in rivers below papermill discharges". New Processes for Wastewater Treatment and Recovery: 140–158.

[14] Rutt, G.P.; Pickering, T.D.; Reynolds, N.R.M. (1993). "The impact of livestock-farming on welsh streams: The development and testing of a rapid biological method for use in the assessment and control of organic pollution from farms". Environmental Pollution. 81 (3): 217–228. doi:10.1016/0269-7491(93)90205-3. PMID 15091808.

[15] Seager, J.; Jones, F.; Rutt, G. (February 1992). "Assessment and Control of Farm Pollution". Water and Environment Journal. 6 (1): 48–53. doi:10.1111/j.1747-6593.1992.tb00737.x. ISSN 1747-6585.

[:6-16] Geatches, Gething, Rutt (2014). 'Sewage fungus': A field and microscopic guide. Environment Agency and Natural Resources Wales.{{cite book}}: CS1 maint: multiple names: authors list (link)

[:7-17] Curtis, E.J.C.; Curds, C.R. (December 1971). "Sewage fungus in rivers in the United Kingdom: The slime community and its constituent organisms". Water Research. 5 (12): 1147–1159. Bibcode:1971WatRe...5.1147C. doi:10.1016/0043-1354(71)90080-7.

[18] Gray, N.F.; Hunter, Christine A. (January 1985). "Heterotrophic slimes in Irish rivers". Water Research. 19 (6): 685–691. doi:10.1016/0043-1354(85)90113-7.

[19] Hickey, Christopher W. (November 1988). "River oxygen uptake and respiratory decay of sewage fungus biofilms". Water Research. 22 (11): 1375–1380. Bibcode:1988WatRe..22.1375H. doi:10.1016/0043-1354(88)90093-0.

[20] Adeola, Samuel; Revitt, Michael; Shutes, Brian; Garelick, Hemda; Jones, Huw; Jones, Clive (2009-01-12). "Constructed Wetland Control of BOD Levels in Airport Runoff". International Journal of Phytoremediation. 11 (1): 1–10. doi:10.1080/15226510802363220. ISSN 1522-6514. S2CID 85164686.

[21] Turnbull, D.A.; Bevan, J.R. (1995). "The impact of airport de-icing on a river: The case of the Ouseburn, Newcastle upon Tyne". Environmental Pollution. 88 (3): 321–332. doi:10.1016/0269-7491(95)93446-7. PMID 15091545.

[22] United States Environmental Protection Agency (US EPA) (2000). "Preliminary Data Summary - Airport Deicing Operations (Revised)" (PDF).

[23] Hirsch (1958). "Biological evaluation of organic pollution of New Zealand streams". New Zealand Journal of Science. 1 (4): 500–553.

[24] Hynes (1960). The biology of polluted waters. Liverpool University Press.

[:8-25] Curtis, E.J.C.; Delves-Broughton, J.; Harrington, D.W. (June 1971). "Sewage fungus: studies of Sphaerotilus slimes using laboratory recirculating channels". Water Research. 5 (6): 267–279. Bibcode:1971WatRe...5..267C. doi:10.1016/0043-1354(71)90172-2.

[26] Smith, Lloyd L.; Kramer, Robert H. (July 1963). "Survival of Walleye Eggs in Relation to Wood Fibers and Sphaerotilus natans in the Rainy River, Minnesota". Transactions of the American Fisheries Society. 92 (3): 220–234. doi:10.1577/1548-8659(1963)92[220:SOWEIR]2.0.CO;2. ISSN 0002-8487.

[27] Magliozzi, Chiara; Grabowski, Robert C.; Packman, Aaron I.; Krause, Stefan (2018-11-30). "Toward a conceptual framework of hyporheic exchange across spatial scales". Hydrology and Earth System Sciences. 22 (12): 6163–6185. doi:10.5194/hess-22-6163-2018. ISSN 1607-7938.

[28] Lewandowski, Jörg; Arnon, Shai; Banks, Eddie; Batelaan, Okke; Betterle, Andrea; Broecker, Tabea; Coll, Claudia; Drummond, Jennifer; Gaona Garcia, Jaime; Galloway, Jason; Gomez-Velez, Jesus; Grabowski, Robert; Herzog, Skuyler; Hinkelmann, Reinhard; Höhne, Anja (2019-10-25). "Is the Hyporheic Zone Relevant beyond the Scientific Community?". Water. 11 (11): 2230. doi:10.3390/w11112230. hdl:20.500.11850/382125. ISSN 2073-4441.

[29] Decho, Alan W.; Visscher, Pieter T.; Reid, R. Pamela (2005), "Production and cycling of natural microbial exopolymers (EPS) within a marine stromatolite", Geobiology: Objectives, Concepts, Perspectives, Elsevier, pp. 71–86, doi:10.1016/b978-0-444-52019-7.50008-5, ISBN 978-0-444-52019-7, retrieved 2023-10-10

[30] Flemming, Hans-Curt; Wingender, Jost (September 2010). "The biofilm matrix". Nature Reviews Microbiology. 8 (9): 623–633. doi:10.1038/nrmicro2415. ISSN 1740-1526. PMID 20676145. S2CID 28850938.

[31] Flemming, Hans-Curt; Wingender, Jost; Szewzyk, Ulrich; Steinberg, Peter; Rice, Scott A.; Kjelleberg, Staffan (September 2016). "Biofilms: an emergent form of bacterial life". Nature Reviews Microbiology. 14 (9): 563–575. doi:10.1038/nrmicro.2016.94. ISSN 1740-1526. PMID 27510863. S2CID 4384131.

[32] Friedman, Barry A.; Dugan, Patrick R. (May 1968). "Identification of Zoogloea species and the Relationship to Zoogloeal Matrix and Floc Formation". Journal of Bacteriology. 95 (5): 1903–1909. doi:10.1128/jb.95.5.1903-1909.1968. ISSN 0021-9193. PMC 252226. PMID 5650090.

[33] Patrick, F; Loutit, M (1976). "Passage of metals in effluents, through bacteria to higher organisms". Water Research. 10 (4): 333–335. Bibcode:1976WatRe..10..333P. doi:10.1016/0043-1354(76)90176-7.

[34] Gray, N (1987). Sewage Fungus in Irish Rivers: A Guide to Identification, Evaluation and Control. Trinity College, University of Dublin.

[35] McKinney, Ross E. (2004-03-11). Environmental Pollution Control Microbiology (0 ed.). CRC Press. doi:10.1201/9780203025697. ISBN 978-1-135-52187-5.

[36] Hartwell, S. Ian; Jordahl, David M.; Evans, Joyce E.; May, Eric B. (August 1995). "Toxicity of aircraft de-icer and anti-icer solutions to aquatic organisms". Environmental Toxicology and Chemistry. 14 (8): 1375–1386. doi:10.1002/etc.5620140813.

[37] Pillard, David A (February 1995). "Comparative toxicity of formulated glycol deicers and pure ethylene and propylene glycol to Ceriodaphnia dubia and Pimephales promelas". Environmental Toxicology and Chemistry. 14 (2): 311–315. doi:10.1002/etc.5620140217.

[38] Pillard, D. A.; DuFresne, D. L. (1999-07-01). "Toxicity of Formulated Glycol Deicers and Ethylene and Propylene Glycol to Lactuca sativa, Lolium perenne, Selenastrum capricornutum, and Lemna minor". Archives of Environmental Contamination and Toxicology. 37 (1): 29–35. doi:10.1007/s002449900486. ISSN 0090-4341. PMID 10341039. S2CID 12747431.

[39] Gray, N. F. (November 1985). "Heterotrophic Slimes in Flowing Waters". Biological Reviews. 60 (4): 499–548. doi:10.1111/j.1469-185X.1985.tb00621.x. ISSN 1464-7931. S2CID 83810613.

[40] Dias, FF; Dondero, NC; Finstein, MS (August 1968). "Attached Growth of Sphaerotilus and Mixed Populations in a Continuous-flow Apparatus". Applied Microbiology. 16 (8): 1191–99. doi:10.1128/am.16.8.1191-1199.1968. ISSN 0003-6919. PMC 547617. PMID 4877499.

[41] Dias, FF; Heukelekian, H (September 1967). "Utilization of Inorganic Nitrogen Compounds by Sphaerotilus natans Growing in a Continuous-Flow Apparatus". Applied Microbiology. 15 (5): 1083–86. doi:10.1128/am.15.5.1083-1086.1967. ISSN 0003-6919. PMC 547145. PMID 16349722.

[42] Phaup, JD (October 1968). "The biology of Sphaerotilus species". Water Research. 2 (9): 597–614. Bibcode:1968WatRe...2..597P. doi:10.1016/0043-1354(68)90065-1.

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Sewage fungus

Environmental Impacts

Microbial composition

Drivers of sewage fungus

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