Ammonia oxidizing bacteria community dynamics in a pilot-scale wastewater treatment plant

Background

Chemoautotrophic ammonia oxidizing bacteria (AOB) have the metabolic ability to oxidize ammonia to nitrite aerobically. This metabolic feature has been widely used, in combination with denitrification, to remove nitrogen from wastewater in wastewater treatment plants (WWTPs). However, the relative influence of specific deterministic environmental factors to AOB community dynamics in WWTP is uncertain. The ecological principles underlying AOB community dynamics and nitrification stability and how they are related are also poorly understood.

Methodology/Principal Findings

The community dynamics of ammonia oxidizing bacteria (AOB) in a pilot-scale WWTP were monitored over a one-year period by Terminal Restriction Fragment Length Polymorphism (T-RFLP). During the study period, the effluent ammonia concentrations were almost below 2 mg/L, except for the first 60 days, indicting stable nitrification. T-RFLP results showed that, during the test period with stable nitrification, the AOB community structures were not stable, and the average change rate (every 15 days) of AOB community structures was 10%±8%. The correlations between T-RFLP profiles and 10 operational and environmental parameters were tested by Canonical Correlation Analysis (CCA) and Mantel test. The results indicated that the dynamics of AOB community correlated most strongly with Dissolved Oxygen (DO), effluent ammonia, effluent Biochemical Oxygen Demand (BOD) and temperature.

Conclusions/Significance

This study suggests that nitrification stability is not necessarily accompanied by a stable AOB community, and provides insight into parameters controlling the AOB community dynamics within bioreactors with stable nitrification.     

Bacterial community dynamics in two full‐scale wastewater treatment systems with functional stability

Aims: To characterize the bacterial community dynamics over 1 year in two full‐scale wastewater treatment systems operated under constant conditions and exhibiting stable performance. Methods and Results: Functional stability was defined and quantified by the effluent concentration of biological oxygen demand, total nitrogen and ammonia. Community dynamics were investigated using specific PCR followed by terminal restriction fragment length polymorphism (T‐RFLP) of the 16S rRNA gene. The T‐RFLP results indicated that during the period of functional stability, the bacterial community structures in two full‐scale wastewater treatment systems were not stable, and the average change rates every 15 days of the two systems were 22·6 ± 6·9 and 21·6 ± 7·3%, respectively. The corresponding species with dominant T‐RFs were determined by clonal sequencing and T‐RFLP. Based on Pareto–Lorenz distribution curves, it was observed that only a small number of micro‐organisms were numerically dominant in the two systems. Conclusions: The results of this study showed that, throughout the period of the study, the bacterial community structure changed significantly in two full‐scale wastewater treatment systems despite the stable function. Significance and Impact of the Study: The findings enrich the theory involving the relation between bacterial community dynamics and functional stability in full‐scale wastewater treatment plants.     

Performance of a pilot-scale sewage treatment: an up-flow anaerobic sludge blanket (UASB) and a down-flow hanging sponge (DHS) reactors combined system by sulfur-redox reaction process under low-temperature conditions

Performance of a wastewater treatment system utilizing a sulfur-redox reaction of microbes was investigated using a pilot-scale reactor that was fed with actual sewage. The system consisted of an up-flow anaerobic sludge blanket (UASB) reactor and a down-flow hanging sponge (DHS) reactor with a recirculation line. Consequently, the total COD_Cr (465 ± 147 mg L⁻¹; total BOD of 207 ± 68 mg L⁻¹) at the influent was reduced (70 ± 14 mg L⁻¹; total BOD of 9 ± 2 mg L⁻¹) at the DHS effluent under the conditions of an overall hydraulic retention time of 12 h, a recirculation ratio of 2, and a low-sewage temperature of 7.0 ± 2.8 °C. A microbial analysis revealed that sulfate-reducing bacteria contributed to the degradation of organic matter in the UASB reactor even in low temperatures. The utilized sulfur-redox reaction is applicable for low-strength wastewater treatment under low-temperature conditions.     

Links between Plant and Rhizoplane Bacterial Communities in Grassland Soils, Characterized Using Molecular Techniques

Molecular analysis of grassland rhizosphere soil has demonstrated complex and diverse bacterial communities, with resultant difficulties in detecting links between plant and bacterial communities. These studies have, however, analyzed “bulk” rhizosphere soil, rather than rhizoplane communities, which interact most closely with plants through utilization of root exudates. The aim of this study was to test the hypothesis that plant species was a major driver for bacterial rhizoplane community composition on individual plant roots. DNA extracted from individual roots was used to determine plant identity, by analysis of the plastid tRNA leucine (trnL) UAA gene intron, and plant-related bacterial communities. Bacterial communities were characterized by analysis of PCR-amplified 16S rRNA genes using two fingerprinting methods: terminal restriction fragment length polymorphisms (T-RFLP) and denaturing gradient gel electrophoresis (DGGE). Links between plant and bacterial rhizoplane communities could not be detected by visual examination of T-RFLP patterns or DGGE banding profiles. Statistical analysis of fingerprint patterns did not reveal a relationship between bacterial community composition and plant species but did demonstrate an influence of plant community composition. The data also indicated that topography and other, uncharacterized, environmental factors are important in driving bacterial community composition in grassland soils. T-RFLP had greater potential resolving power than DGGE, but findings from the two methods were not significantly different.     

Removal of nitrate and phosphorus from hydroponic wastewater using a hybrid denitrification filter (HDF)

A laboratory-scale hybrid-denitrification filter (HDF) was designed by combining a plant material digester and a denitrification filter into a single unit for the removal of nitrate and phosphorus from glasshouse hydroponic wastewater. The carbon to nitrate (C:N) ratio for efficient operation of the HDF was calculated to be 1.93:1 and the COD/BOD₅ ratio was 1.2:1. When the HDF was continuously operated with the plant material replaced every 2 days and 100% internal recirculation of the effluent, a high level of nitrate removal (320–5 mg N/L, >95% removal) combined with a low effluent sBOD₅ concentration (<5 mg/L) was consistently achieved. Moreover, phosphate concentrations in the effluent were maintained below 7.5 mg P/L (>81% reduction). This study demonstrates the potential to combine a digester and a denitrification filter in a single unit to efficiently remove nitrate and phosphate from hydroponic wastewater in a single unit.     

Bacterial community responses to increasing ammonia concentrations in model recirculating vertical flow saline biofilters

This study represents the first analysis of ammonia removal and bacterial communities in gravel biofilters treating saline wastewater and is of relevance to the growing inland marine aquaculture industry. As part of a study to gain greater understanding of the microbial processes occurring in a newly constructed limestone gravel wetland at a commercial marine fish farm, this study was designed to establish the ammonia removal capacity of model biofilters treating saline aquaculture wastewater and to investigate changes to total bacterial communities and ammonia-oxidizing bacterial communities as the biofilters are exposed to increasing ammonia concentrations. Three replicate laboratory-scale gravel biofilters were constructed and the limits of nitrification capacity were tested by dosing with aquaculture wastewater supplemented with increasing amounts of ammonium chloride. The experiment was run over a 12-week period with the water temperature between 24.5 and 28 °C and salinity between 28 and 38 ppt. Greater than 97% ammonia removal in each weekly treatment period was observed with ammonia concentrations of up to 600 ppm. At higher concentrations of ammonia, a lower percentage of ammonia was removed, and on occasion nitrite accumulation was observed. A drop in the number of operational taxonomic units (OTUs) detected in the bacterial community as measured by 16s rRNA T-RFLP was observed concurrent with the decrease in percentage ammonia removal. T-RFLP of the amoA gene showed the experimental biofilters to be dominated by three different OTUs of ammonia-oxidizing bacteria. A synchronous successional pattern among these three ammonia oxidizers was observed. The three OTUs were identified as belonging to three different nitrosomonad clusters. This study demonstrates that the vertical flow gravel biofilters have the ability to treat saline aquaculture wastewater that has a high ammonia concentration and that the microbial community within saline biofilters has the capacity to adapt to changing ammonia levels while maintaining nitrification activity.     

High reproducibility of ammonia-oxidizing bacterial communities in parallel sequential batch reactors

Aims:  To investigate whether the ammonia-oxidizing bacterial (AOB) communities of replicate nitrifying bioreactors (i) co-evolve or diverge over time and (ii) are stable or dynamic during periods of complete nitrification.
Methods and Results:  Three sequential batch reactors (SBR) were inoculated with sludge from a municipal wastewater treatment plant, fed with ammonium-enriched tap water and operated in parallel for 134 days. Polymerase chain reaction-denaturing gradient gel electrophoresis (PCR-DGGE) demonstrated co-evolvement of the AOB communities over time. During start-up, temporary decreases in nitrification were noticed, and the AOB community rate of change values (Δ _{t (week)}) were medium to high (12–22%). During the adjacent period of complete nitrification, low AOB community dynamics were observed (Δ _{t (week)} < 5%). Further pragmatic processing of the DGGE profiles revealed a high range-weighted richness and a medium functional organization of the AOB communities.
Conclusions:  After a start-up period, high functional stability and low dynamics of the AOB communities were observed. Deterministic rather than stochastic driving forces led to AOB community co-evolvement in the replicate SBR.
Significance and Impact of the Study:  Replicates in identical set-ups are reproducible, and pragmatic processing of DGGE patterns is a straightforward tool to score and compare the functionality of the bacterial communities.     

Treatment of olive mill wastewater in pilot-scale vertical flow constructed wetlands

Pilot-scale constructed wetlands (CW) were constructed and operated to treat pre-treated olive mill wastewater. Pilot-scale units comprising three identical series with four pilot-scale vertical flow CWs were operated for one harvest season in a Greek olive mill plant. The pilot-scale CWs were filled with various porous media (i.e., cobble, gravel, and sand) of different gradations. Two series of pilot-scale units were planted with common reeds and the third (control) was unplanted. Mean influent concentrations were 14,120 mg/L, 2841 mg/L, 95 mg/L, 123 mg/L and 506 mg/L for COD, phenols, ortho-phosphate, ammonia and TKN, respectively. Despite the rather high influent concentrations, the performance of the CW units was very effective since it achieved removals of about 70%, 70%, 75% and 87% for COD, phenols, TKN and ortho-phosphate, respectively. COD, phenol and TKN removal seems to be significantly higher in the planted series, while ortho-phosphate removal shows no significant differences among the three series. Temperature and pollutant surface load seem to affect the removal efficiency of all pollutants. Compared to previous studies, pollutant surface loads applied here were higher (by one or two orders of magnitude). Even though high removal efficiencies were achieved, effluent pollutant concentrations remained high, thus preventing their use for irrigation or immediate disposal into the environment.     

Functionally stable and phylogenetically diverse microbial enrichments from microbial fuel cells during wastewater treatment

Microbial fuel cells (MFCs) are devices that exploit microorganisms as biocatalysts to recover energy from organic matter in the form of electricity. One of the goals of MFC research is to develop the technology for cost-effective wastewater treatment. However, before practical MFC applications are implemented it is important to gain fundamental knowledge about long-term system performance, reproducibility, and the formation and maintenance of functionally-stable microbial communities. Here we report findings from a MFC operated for over 300 days using only primary clarifier effluent collected from a municipal wastewater treatment plant as the microbial resource and substrate. The system was operated in a repeat-batch mode, where the reactor solution was replaced once every two weeks with new primary effluent that consisted of different microbial and chemical compositions with every batch exchange. The turbidity of the primary clarifier effluent solution notably decreased, and 97% of biological oxygen demand (BOD) was removed after an 8-13 day residence time for each batch cycle. On average, the limiting current density was 1000 mA/m(2), the maximum power density was 13 mW/m(2), and coulombic efficiency was 25%. Interestingly, the electrochemical performance and BOD removal rates were very reproducible throughout MFC operation regardless of the sample variability associated with each wastewater exchange. While MFC performance was very reproducible, the phylogenetic analyses of anode-associated electricity-generating biofilms showed that the microbial populations temporally fluctuated and maintained a high biodiversity throughout the year-long experiment. These results suggest that MFC communities are both self-selecting and self-optimizing, thereby able to develop and maintain functional stability regardless of fluctuations in carbon source(s) and regular introduction of microbial competitors. These results contribute significantly toward the practical application of MFC systems for long-term wastewater treatment as well as demonstrating MFC technology as a useful device to enrich for functionally stable microbial populations.     

Links between plant and rhizoplane bacterial communities in grassland soils, characterized using molecular techniques

Molecular analysis of grassland rhizosphere soil has demonstrated complex and diverse bacterial communities, with resultant difficulties in detecting links between plant and bacterial communities. These studies have, however, analyzed "bulk" rhizosphere soil, rather than rhizoplane communities, which interact most closely with plants through utilization of root exudates. The aim of this study was to test the hypothesis that plant species was a major driver for bacterial rhizoplane community composition on individual plant roots. DNA extracted from individual roots was used to determine plant identity, by analysis of the plastid tRNA leucine (trnL) UAA gene intron, and plant-related bacterial communities. Bacterial communities were characterized by analysis of PCR-amplified 16S rRNA genes using two fingerprinting methods: terminal restriction fragment length polymorphisms (T-RFLP) and denaturing gradient gel electrophoresis (DGGE). Links between plant and bacterial rhizoplane communities could not be detected by visual examination of T-RFLP patterns or DGGE banding profiles. Statistical analysis of fingerprint patterns did not reveal a relationship between bacterial community composition and plant species but did demonstrate an influence of plant community composition. The data also indicated that topography and other, uncharacterized, environmental factors are important in driving bacterial community composition in grassland soils. T-RFLP had greater potential resolving power than DGGE, but findings from the two methods were not significantly different.     

Correlations between molecular and operational parameters in continuous lab-scale anaerobic reactors

In this study, the microbial community characteristics in continuous lab-scale anaerobic reactors were correlated to reactor functionality using the microbial resource management (MRM) approach. Two molecular techniques, denaturing gradient gel electrophoresis (DGGE) and terminal-restriction fragment length polymorphism (T-RFLP), were applied to analyze the bacterial and archaeal communities, and the results obtained have been compared. Clustering analyses showed a similar discrimination of samples with DGGE and T-RFLP data, with a clear separation between the meso- and thermophilic communities. Both techniques indicate that bacterial and mesophilic communities were richer and more even than archaeal and thermophilic communities, respectively. Remarkably, the community composition was highly dynamic for both Bacteria and Archaea, with a rate of change between 30% and 75% per 18 days, also in stable performing periods. A hypothesis to explain the latter in the context of the converging metabolism in anaerobic processes is proposed. Finally, a more even and diverse bacterial community was found to be statistically representative for a well-functioning reactor as evidenced by a low Ripley index and high biogas production.     

Bacterial community composition of a wastewater treatment system reliant on N2 fixation

The temporal stability and change of the dominant phylogenetic groups of the domain bacteria were studied in a model plant-based industrial wastewater treatment system showing high levels of organic carbon removal supported by high levels of N₂ fixation. Community profiles were obtained through terminal restriction fragment length polymorphism analysis and cloning of 16S rRNA amplicons followed by sequencing. Bacterial community profiles showed that ten common terminal restriction fragments made up approximately 50% of the measured bacterial community. As much as 42% of the measured bacterial community could be monitored by using quantitative PCR and primers that targeted three dominant operational taxonomic units. Despite changes in wastewater composition and dissolved oxygen levels, the bacterial community composition appeared stable and was dominated by α-Proteobacteria and β-Proteobacteria, with a lesser amount of the highly diverse bacterial phylum Bacteroidetes. A short period of considerable change in the bacterial community composition did not appear to affect treatment performance indicating functional redundancy in this treatment system.     

Seasonal Fluctuations of Bacterial Community Diversity in Agricultural Soil and Experimental Validation by Laboratory Disturbance Experiments

Natural fluctuations in soil microbial communities are poorly documented because of the inherent difficulty to perform a simultaneous analysis of the relative abundances of multiple populations over a long time period. Yet, it is important to understand the magnitudes of community composition variability as a function of natural influences (e.g., temperature, plant growth, or rainfall) because this forms the reference or baseline against which external disturbances (e.g., anthropogenic emissions) can be judged. Second, definition of baseline fluctuations in complex microbial communities may help to understand at which point the systems become unbalanced and cannot return to their original composition. In this paper, we examined the seasonal fluctuations in the bacterial community of an agricultural soil used for regular plant crop production by using terminal restriction fragment length polymorphism profiling (T-RFLP) of the amplified 16S ribosomal ribonucleic acid (rRNA) gene diversity. Cluster and statistical analysis of T-RFLP data showed that soil bacterial communities fluctuated very little during the seasons (similarity indices between 0.835 and 0.997) with insignificant variations in 16S rRNA gene richness and diversity indices. Despite overall insignificant fluctuations, between 8 and 30% of all terminal restriction fragments changed their relative intensity in a significant manner among consecutive time samples. To determine the magnitude of community variations induced by external factors, soil samples were subjected to either inoculation with a pure bacterial culture, addition of the herbicide mecoprop, or addition of nutrients. All treatments resulted in statistically measurable changes of T-RFLP profiles of the communities. Addition of nutrients or bacteria plus mecoprop resulted in bacteria composition, which did not return to the original profile within 14 days. We propose that at less than 70% similarity in T-RFLP, the bacterial communities risk to drift apart to inherently different states.     

Ethanol degradation and the benefits of incremental priming in pilot-scale constructed wetlands

There has been significant global growth in the use of constructed wetlands for wastewater treatment. The fundamental microbial processes involved in the biodegradation of organic wastewater pollutants determine the range of design and operational parameters relevant to individual constructed wetlands. In this study, the biodegradation and mineralization of ethanol by acclimated and non-acclimated microbial populations in pilot-scale constructed wetlands were compared. By increasing the pollutant concentration at incremental intervals (incremental priming), the biodegradative capacity of a sand-filled constructed wetland was significantly enhanced. At an influent COD concentration of 15,800 mg L⁻¹, no volatile fatty acids were detected in the effluent of an incrementally primed system and the maximum effluent COD concentration was 180 mg L⁻¹. In contrast, an identical, unprimed system, amended with a lower concentration of COD (7587 mg L⁻¹), exhibited a maximum effluent COD concentration of 1400 mg L⁻¹, with the anaerobic metabolites, butyrate and propionate accounting for up to 83% of the effluent COD. It was demonstrated that the use of incremental priming, together with a vertical subsurface flow mode of operation enhanced long-term function of constructed wetlands. Future research should focus on determining the concentration gradients and incremental intervals necessary for optimal microbial acclimation to a range of organic pollutants and/or wastewaters, in order to minimize start-up times without significantly impairing the benefits derived from incremental priming.     

Diversity of abundant bacteria in subsurface vertical flow constructed wetlands

Microorganisms are mainly responsible for the transformation and mineralization of degradable organic pollutants within constructed wetlands (CWs). There is still a lack of knowledge concerning microbial community composition within CWs. In order to elucidate the diversity of bacteria inhabiting subsurface vertical flow CWs, the molecular fingerprint technique “terminal restriction fragment length polymorphism” (T-RFLP) derived from total community DNA, was applied.A comparison of the bacterial communities from a full-scale outdoor vertical flow CW with planted and unplanted indoor pilot-scale vertical flow CWs, operated under similar conditions, revealed that both systems are colonized by similar populations showing only little variation in their composition over filter depth. A comparison of bulk soil from an unplanted CW with the rhizosphere soil from the outdoor and indoor CWs showed differences in the bacterial composition, demonstrating the influence of the plants on the rhizosphere community. A comparison of the wastewater before and after the CW passage demonstrated that the bacterial diversity was clearly reduced within the planted outdoor system only.     

Performance Assessment of Full-Scale Wastewater Treatment Plants Based on Seasonal Variability of Microbial Communities via High-Throughput Sequencing

Microbial communities of activated sludge (AS) play a key role in the performance of wastewater treatment processes. However, seasonal variability of microbial population in varying AS-based processes has been poorly correlated with operation of full-scale wastewater treatment systems (WWTSs). In this paper, significant seasonal variability of AS microbial communities in eight WWTSs located in the city of Guangzhou were revealed in terms of 16S rRNA-based Miseq sequencing. Furthermore, variation redundancy analysis (RDA) demonstrated that the microbial community compositions closely correlated with WWTS operation parameters such as temperature, BOD, NH₄⁺-N and TN. Consequently, support vector regression models which reasonably predicted effluent BOD, SS and TN in WWTSs were established based on microbial community compositions. This work provided an alternative tool for rapid assessment on performance of full-scale wastewater treatment plants.     

Long term operation of pilot-scale biological nutrient removal process in treating municipal wastewater

The performance of a pilot-scale biological nutrient removal process has been evaluated for 336 days, receiving the real municipal wastewater with a flowrate of 6.8 m³/d. The process incorporated an intermittent aeration reactor for enhancing the effluent quality, and a nitrification reactor packed with the porous polyurethane foam media for supporting the attached-growth of microorganism responsible for nitrification. The observation shows that the process enabled a relatively stable and high performance in both organics and nutrient removals. When the SRT was maintained at 12 days, COD, nitrogen, and phosphorus removals averaged as high as 89% at a loading rate of 0.42–3.95 kg COD/m³ d (corresponding to average influent concentration of 304 mg COD/L), 76% at the loading rate of 0.03–0.27 kg N/m³ d (with 37.1 mg TN/L on average), and 95% at the loading rate of 0.01–0.07 kg TP/m³ d (with 5.4 mg TP/L on average), respectively.     

Bacterial population changes in a membrane bioreactor for graywater treatment monitored by denaturing gradient gel electrophoretic analysis of 16S rRNA gene fragments

The bacterial population of a graywater treatment system was monitored over the course of 100 days, along with several wastewater biochemical parameters. The graywater treatment system employed an 1,800-liter membrane bioreactor (MBR) to process the waste, with essentially 100% recycling of the biomass. Graywater feed consisting of 10% galley water and 90% laundry water, selected to approximate the graywater composition on board U.S. Navy ships, was collected offsite. Five-day biological oxygen demand (BOD(5)), oils and greases (O/G), nitrogen, and phosphorus were monitored in the feed and were found to vary greatly day to day. Changes in the bacterial population were monitored by PCR amplification of region 332 to 518 (Escherichia coli numbering) of the 16S rRNA gene and denaturing gradient gel electrophoresis (DGGE) analysis of the resultant PCR products. DGGE analysis indicated a diverse and unstable bacterial population throughout the 100-day period, with spikes in feed strength causing significant changes in community structure. Long-term similarity between the communities was 0 to 25%, depending on the method of analysis. In spite of the unstable bacterial population, the MBR system was able to meet effluent quality parameters approximately 90% of the time.     

Nutrient recovery from domestic wastewater using a UASB-duckweed ponds system

The pilot-scale wastewater treatment system used in this study comprised a 40-l UASB reactor (6-h HRT) followed by three duckweed ponds in series (total HRT 15 days). During the warm season, the treatment system achieved removal values of 93%, 96% and 91% for COD, BOD and TSS, respectively. Residual values of ammonia, TKN and total phosphorus were 0.41 mg N/l, 4.4 mg N/l and 1.11 mg P/l, with removal efficiencies of 98%, 85% and 78%, respectively. The system achieved 99.998% faecal coliform removal during the warm season with final effluent containing 4 x 10(3) cfu/100 ml. During the winter, the system was efficient in removing COD, BOD and TSS but not nutrients. The system was deficient in the removal of faecal coliforms during the winter, producing effluent with 4.7 x 10(5) cfu/100 ml. During the warm season, the N removal consisted of 80% by plant uptake, 5% by sedimentation and 15% unaccounted for. A duckweed production rate of 33 t dry matter per hectare per 8 months was achieved.     

Performance evaluation and phylogenic analysis of a full-scale subsurface cobble-bed biofilm system

The purpose of this study is to evaluate the efficiency of municipal wastewater treatment by a subsurface cobble-bed biofilm system (SCBS) in Taipei, Taiwan. In contrast to traditional wastewater treatment facilities, SCBS uses cobbles as the contact media in the biofilm treatment unit. In this study, the SCBS consists of a series of underground treatment units, including a sedimentation tank, a grit chamber, two bar screens, a pumping station, a distribution tank, a collection tank and an effluent tank. At the flowrate of 4000 m³/day, the average influent concentrations for biochemical oxygen demand, suspended solid, ammonium nitrogen, and total phosphorus were 66.99 mg/L, 26.14 mg/L, 17.33 mg/L, and 1.96 mg/L, respectively. After 39 months of operation, the measured influent and effluent results show that the treatment efficiencies obtained from the SCBS for biochemical oxygen demand, suspended solid, ammonium nitrogen, and total phosphorus are 91.3%, 84.0%, 84.0%, and 26.0%, respectively. The result of a first-order kinetic analysis shows that the NH₃-N degradation constant is greater than the BOD degradation constant in this cobble-bed biofilm unit. Probability analysis revealed that the SCBS may be an attractive alternative from the perspectives of treatment efficiency for municipal wastewater treatment. Klebsiella spp. were found to be the predominant species in the biofilm system in the SCBS.