Heterotrophic Nitrification-aerobic Denitrification Bacteria Research Articles

Research on the application of heterotrophic nitrification-aerobic denitrification bacteria in membrane bioreactor (MBR).

Inoculating heterotrophic nitrification-aerobic denitrification bacteria (HN-AD) to enhance membrane bioreactor (MBR) efficiency may result in the loss of functional bacteria. Therefore, this study compares the application results of enhancing MBR with a self-designed biological amplifier coupled with HN-AD against the performance of conventional MBR. After enhancement, the MBR achieved a removal efficiency of 96.7% for NH4+-N (100mg/L) and 96.4% for COD (400mg/L) in synthetic wastewater. There was a 33% increase in TN (100mg/L) removal efficiency. The dominant bacteria in the MBR were Alcaligenes (48.4%) and Thauera (15.2%). Additionally, the abundance of denitrification genes (nirK, norB, nosZ) increased in the enhanced MBR, contributing to improved TN removal efficiency. The use of a biological amplifier effectively solved the problem of HN-AD loss in sewage treatment.

Outdoor tubular photobioreactor microalgae-microorganisms biofilm treatment of municipal wastewater: Enhanced heterotrophic assimilation and synergistic aerobic denitrogenation

This research evaluated a microalgae consortium (MC) in a pilot-scale tubular photobioreactor for municipal wastewater (MWW) treatment, compared with an aeration column photobioreactor. Transitioning from suspended MC to a microalgae-microbial biofilm (MMBF) maintained treatment performance despite increasing influent from 50 L to 150 L in a 260 L system. Carbon and nitrogen removal were effective, but phosphorus removal varied due to biofilm shading and the absence of phosphorus-accumulating organisms. High influent flow caused MMBF detachment due to shear stress. Stabilizing and re-establishing the MMBF showed that a stable phycosphere influenced microbial diversity and interactions, potentially destabilizing the MMBF. Heterotrophic nitrification-aerobic denitrification bacteria were crucial for MC equilibrium. Elevated gene expression related to nitrogen fixation, organic nitrogen metabolism, and nitrate reduction confirmed strong microalgal symbiosis, highlighting MMBF’s treatment potential. This study supports the practical application of microalgae in wastewater treatment.

Transcriptome analysis of the degradation process of organic nitrogen by two heterotrophic nitrifying and aerobic denitrifying bacteria.

The heterotrophic nitrification aerobic denitrification bacteria (HNDS) can perform nitrification and denitrification at the same time. Two HNDS strains, Achromobacter sp. HNDS-1 and Enterobacter sp. HNDS-6 which exhibited an amazing ability to solution nitrogen (N) removal have been successfully isolated from paddy soil in our lab. When peptone or ammonium sulfate as sole N source, no significant difference in gene expression related to nitrification and denitrification of the strains was found according to the transcriptome analysis. The expression of phosphomethylpyrimidine synthase (thiC), ABC transporter substrate-binding protein, branched-chain amino acid ABC transporter substrate-binding protein, and RNA polymerase (rpoE) in HNDS-1 were significantly upregulated when used peptone as N source, while the expression of exopolysaccharide production protein (yjbE), RNA polymerase (rpoC), glutamate synthase (gltD) and ABC-type branched-chain amino acid transport systems in HNDS-6 were significantly upregulated. This indicated that these two strains are capable of using organic N and converting it into NH4+-N, then utilizing NH4+-N to synthesize amino acids and proteins for their own growth, and strain HNDS-6 can also remove NH4+-N through nitrification and denitrification.

Characteristics of Novel Heterotrophic Nitrification–Aerobic Denitrification Bacteria Bacillus subtilis F4 and Alcaligenes faecalis P4 Isolated from Landfill Leachate Biochemical Treatment System

Heterotrophic nitrification-aerobic denitrification (HN-AD) bacteria are the key functional microorganisms needed to achieve simultaneous nitrification and denitrification (SND). In this study, 25 strains of HN-AD bacteria were successfully isolated from a stable landfill leachate biochemical treatment system, of which 10 strains belonged to Firmicutes and 15 strains belonged to Proteobacteria. Bacillus subtilis F4 and Alcaligenes faecalis P4 displayed good tolerance at a wide range of ammonia nitrogen (NH4+-N) concentrations. When the C/N ratio was 20, the removal rates of ammonia nitrogen were 90.1% and 89.5%, and the chemical oxygen demand (COD) removal rates were 92.4% and 93.9%, respectively. The napA gene encoding periplasmic nitrate reductase (Nap) and the nirS gene encoding nitrite reductase (Nir) were detected, and nitrogen balance showed assimilation and HN-AD was the main nitrogen metabolism mode in both strains. The use of immobilization materials could increase removal rate of ammonia nitrogen by 21.1% and 29.6%, respectively. The research results of this work can provide theoretical basis and technical support for the practical application of HN-AD bacteria to enhance the treatment of high ammonia nitrogen wastewater with high efficiency and low consumption.

Influence and mechanism of typical transition metal ions on the denitrification performance of heterotrophic nitrification-aerobic denitrification bacteria

To investigate the inhibitory effects of various transition metal ions on nitrogen removal and their underlying mechanisms, the single and combined effects of Cu2+ Ni2+ and Zn2+ on Heterotrophic nitrification-aerobic denitrification (HN-AD) bacteria Acinetobacter sp. TAC-1 were studied in a batch experiment system. The results revealed that increasing concentrations of Cu2+ and Ni2+ had a detrimental effect on the removal of ammonium nitrogen (NH4+-N) and total nitrogen (TN). Specifically, Cu2+ concentration of 10 mg/L, the TN degradation rate was 55.09%, compared to 77.60% in the control group. Cu2+ exhibited a pronounced inhibitory effect. In contrast, Zn2+ showed no apparent inhibitory effect on NH4+-N removal and even enhanced TN removal at lower concentrations. However, when the mixed ion concentration of Zn2++Ni2+ exceeded 5 mg/L, the removal rates of NH4+-N and TN were significantly reduced. Moreover, transition metal ions did not significantly impact the removal rates of chemical oxygen demand (COD). The inhibition model fitting results indicated that the inhibition sequence was Cu2+ > Zn2+ > Ni2+. Transcriptome analysis demonstrated that metal ions influence TAC-1 activity by modulating the expression of pivotal genes, including zinc ABC transporter substrate binding protein (znuA), ribosomal protein (rpsM), and chromosome replication initiation protein (dnaA) and DNA replication of TAC-1 under metal ion stress, leading to disruptions in transcription, translation, and cell membrane structure. Finally, a conceptual model was proposed by us to summarize the inhibition mechanism and possible response strategies of TAC-1 bacteria under metal ion stress, and to address the lack of understanding regarding the influence mechanism of TAC-1 on nitrogen removal in wastewater co-polluted by metal and ammonia nitrogen. The results provided practical guidance for the management of transition metal and ammonia nitrogen co-polluted water bodies, as well as the removal of high nitrogen.

Effects of Cd(II) on nitrogen removal by a heterotrophic nitrification aerobic denitrification bacterium Pseudomonas sp. XF-4

Simultaneous heterotrophic nitrification and aerobic denitrification (SND) is gaining tremendous attention due to its high efficiency and low cost in water treatment. However, SND on an industrial scale is still immature since effects of coexisting pollutants, for example, heavy metals, on nitrogen removal remains largely unresolved. In this study, a HNAD bacterium (Pseudomonas sp. XF-4) was isolated. It could almost completely remove ammonium and nitrate at pH 5–9 and temperature 20 ℃-35 ℃ within 10 h, and also showed excellently simultaneous nitrification and denitrification efficiency under the coexistence of any two of inorganic nitrogen sources with no intermediate accumulation. XF-4 could rapidly grow again after ammonium vanish when nitrite or nitrate existed. There was no significant effects on nitrification and denitrification when Cd(II) was lower than 10 mg/L, and 95 % of Cd(II) was removed by XF-4. However, electron carrier and electron transport system activity was inhibited, especially at high concentration of Cd(II). Overall, this study reported a novel strain capable of simultaneous nitrification and denitrification coupled with Cd(II) removal efficiently. The results provided new insights into treatment of groundwater or wastewater contaminated by heavy metals and nitrogen.

Effect of carbon source on carbon and nitrogen metabolism of common heterotrophic nitrification-aerobic denitrification pathway

Pseudomonas sp. ZHL02, removing nitrogen via ammonia nitrogen (NH4+) → hydroxylamine (HN2OH) → nitrite (NO2−) → nitrate (NO3−) → NO2− → nitric oxide (NO) → nitrous oxide (N2O) pathway was employed for getting in-depth information on the heterotrophic nitrification-aerobic denitrification (HNAD) pathway from carbon oxidation, nitrogen conversion, electron transport process, enzyme activity, as well as gene expression while sodium succinate, sodium citrate, and sodium acetate were utilized as the carbon sources. The nitrogen balance analysis results demonstrated that ZHL02 mainly removed NH4+-N through assimilation. The carbon source metabolism resulted in the discrepancies in electron transport chain and nitrogen removal between different HNAD bacteria. Moreover, the prokaryotic strand-specific transcriptome method showed that, amo and hao were absent in ZHL02, and unknown genes may be involved in ZHL02 during the HNAD process. As a fascinating process for removing nitrogen, the HNAD process is still puzzling, and the relationship between carbon metabolism and nitrogen metabolism among different HNAD pathways should be studied further.

Identification and Synergistic Denitrification of Two Heterotrophic Nitrification-Aerobic Denitrification Bacteria

Identification and Synergistic Denitrification of Two Heterotrophic Nitrification-Aerobic Denitrification Bacteria

Optimization of nitrogen removal by immobilized heterotrophic nitrification aerobic denitrifying bacterium using response surface methodology

An innovative bacterial carrier complex embedded with polyvinyl alcohol and sodium alginate complex (P.S.) was developed, using the heterotrophic nitrification-aerobic denitrification (HN-AD) strain Paracoccus sp. QD-21, which was carefully selected from activated sludge. The performance of the immobilized bacterium on ammonia nitrogen, nitrate nitrogen, and nitrite nitrogen was thoroughly analyzed, aiming to investigate the synergistic effects of the P.S.-encapsulated bacterial carrier on the HN-AD strain QD-21. To assess the adaptability of the immobilized bacterium, various single-factor analysis experiments were conducted, including the assessment of carbon source, C/N, temperature, pH, rotational speed, and salinity. Subsequently, response surface methodology was employed to conduct further optimization, leading to the identification of optimum conditions (147.31 rpm, 36°C, C/N 8.72, pH 8). Under these conditions, an impressive ammonia nitrogen removal rate of 94.82 ± 0.82% was achieved. Furthermore, practical wastewater treatment experiments were carried out, demonstrating a remarkable 63.31% increase in ammonia nitrogen removal when the carrier with HN-AD bacterium attached was employed. Notably, the addition of a specific carbon source remarkably enhanced the treatment efficiency, resulting in a complete removal rate of 100%. These significant findings lay a solid foundation for the engineering application of the HN-AD bacterium and highlight the promising potential of the P.S.-encapsulated bacterial carrier complex in various applications.

Conversion behavior of heterotrophic nitrification - aerobic denitrification bacterium Paracoccus denitrificans HY-1 in nitrogen and phosphorus removal

The heterotrophic nitrification-aerobic denitrification (HN-AD) bacteria harbored remarkable removal capacity in eliminating single nitrogen pollutant, but their comprehensive performance in dealing with mixed nitrogen sources remained unknown. In this study, the denitrification capacity of HN-AD bacteria Paracoccus denitrificans HY-1 was explored through whole-genome sequencing and nitrogen metabolic pathway analysis, and the complex nitrogen sources with gradient concentration were used to validate its nitrogen removal efficiency under both aerobic and anaerobic conditions. The results demonstrated that strain HY-1 harbored an efficient degradation capacity on three nitrogen sources, and the removal rates of NH4+-N, NO2−-N and NO3−-N were 10.67 mg·L−1·h−1, 27.81 mg·L−1·h−1, and 22.05 mg·L−1·h−1, respectively. The denitrification rate of 15.33 mg·L−1·h−1 was obtained under anaerobic condition with mixed nitrogen sources. Also, the simultaneous nitrification denitrification phosphorus removal (SNDPR) trial revealed that strain HY-1 could eliminate PO43−-P with a rate of 0.38 mg·L−1·h−1 under coexistence condition of the three nitrogen sources. Combined with the nitrogen balance analysis, the results indicated that strain HY-1 possesses a great potential for application in complex nitrogen and phosphorus contaminated environments.

Effects of sliver nanoparticles on nitrogen removal by the heterotrophic nitrification-aerobic denitrification bacteria Zobellella sp. B307 and their toxicity mechanisms

Due to the widespread use of sliver nanoparticles (AgNPs), a large amount of AgNPs has inevitably been released into the environment, and there is growing concern about the toxicity of AgNPs to nitrogen-functional bacteria. In addition to traditional anaerobic denitrifying bacteria, heterotrophic nitrification-aerobic denitrification (HNAD) bacteria are also important participants in the nitrogen cycle. However, the mechanisms by which AgNPs influence HNAD bacteria have yet to be explicitly demonstrated. In this study, the inhibitory effects of different concentrations of AgNPs on a HNAD bacteria Zobellella sp. B307 were investigated, and the underlying mechanism was explored by analyzing the antioxidant system and the activities of key denitrifying enzymes. Results showed that AgNPs could inhibit the growth and the HNAD ability of Zobellella sp. B307. AgNPs could accumulate on the surface of bacterial cells and significantly destroyed the cell membrane integrity. Further studies demonstrated that the presence of high concentration of AgNPs could result in the overproduction of reactive oxygen species (ROS) and related oxidative stress in the cells. Furthermore, the catalytic activities of key denitrifying enzymes (nitrate reductase (NAR), nitrite reductase (NIR), and nitrous oxide reductase (N2OR)) were significantly suppressed under exposure to a high concentration of AgNPs (20 mg·L−1), which might be responsible for the inhibited nitrogen removal performance of strain B307. This work could improve our understanding of the inhibitory effect and underlying mechanism of AgNPs on HNAD bacteria.

A novel derivative synchronous fluorescence method for the rapid, non-destructive and intuitive differentiation of denitrifying bacteria

It is challenging to differentiate bacteria residing in the same habitat by direct observation. This difficulty impedes the harvest, application and manipulation of functional bacteria in environmental engineering. In this study, we developed a novel method for rapid differentiation of living denitrifying bacteria based on derivative synchronous fluorescence spectroscopy, as exemplified by three heterotrophic nitrification-aerobic denitrification bacteria having the maximum nitrogen removal efficiencies greater than 90%. The intact bacteria and their living surroundings can be analyzed as an integrated target, which eliminates the need for the complex pre-processing of samples. Under the optimal synchronous scanning parameter (Δλ = 40 nm), each bacterium possesses a unique fluorescence spectral structure and the derivative synchronous fluorescence technique can significantly improve the spectral resolution compared to other conventional fluorescence methods, which enables the rapid differentiation of different bacteria through derivative synchronous fluorescence spectra as fast as 2 min per spectrum. Additionally, the derivative synchronous fluorescence technique can extract the spectral signals contributed by bacterial extracellular substances produced in the biological nitrogen removal process. Moreover, the results obtained from our method can reflect the real-time denitrification properties of bacteria in the biological nitrogen removal process of wastewater. All these merits highlight derivative synchronous fluorescence spectroscopy as a promising analytic method in the environmental field.

Enhanced ammonia removal in tidal flow constructed wetland by incorporating steel slag: Performance, microbial community, and heavy metal release

Utilizing alkaline solid wastes, such as steel slag, as substrates in tidal flow constructed wetlands (TFCWs) can effectively neutralize the acidity generated by nitrification. However, the impacts of steel slag on microbial communities and the potential risk of heavy metal release remain poorly understood. To address these knowledge gaps, this study compared the performance and microbial community structure of TFCWs filled with a mixture of steel slag and zeolite (TFCW-S) to those filled with zeolite alone (TFCW-Z). TFCW-S exhibited a much higher NH4+-N removal efficiency (98.35 %) than TFCW-Z (55.26 %). Additionally, TFCW-S also achieved better TN and TP removal. The steel slag addition helped maintain the TFCW-S effluent pH at around 7.5, while the TFCW-Z effluent pH varied from 3.74 to 6.25. The nitrification and denitrification intensities in TFCW-S substrates were significantly higher than those in TFCW-Z, consistent with the observed removal performance. Moreover, steel slag did not cause excessive heavy metal release, as the effluent concentrations were below the standard limits. Microbial community analysis revealed that ammonia-oxidizing bacteria, ammonia-oxidizing archaea, and complete ammonia-oxidizing bacteria coexisted in both TFCWs, albeit with different compositions. Furthermore, the enrichment of heterotrophic nitrification-aerobic denitrification bacteria in TFCW-S likely contributed to the high NH4+-N removal. In summary, these findings demonstrate that the combined use of steel slag and zeolite in TFCWs creates favorable pH conditions for ammonia-oxidizing microorganisms, leading to efficient ammonia removal in an environmentally friendly manner.

Factors affecting the enrichment of heterotrophic nitrifying aerobic denitrifying bacteria in municipal wastewater treatment systems and analysis of differences in nitrogen metabolism

In this study, a four-stage MPSR process was employed for the treatment of urban wastewater. Three different conditions were established by controlling dissolved oxygen (DO) concentration, organic load rate (OLR), and influent pH value to examine pollutant removal effects and the changes of the heterotrophic nitrification-aerobic denitrification (HN-AD) functional bacteria communities. The factors driving changes in the HN-AD bacterial community and differences in nitrogen metabolism were revealed in the urban wastewater treatment system. The results showed that by forming anoxic/aerobic alternate oxygen environments and increasing influent OLR and pH, various HN-AD bacteria (Pseudomonas, Aeromonas, Flavobacterium) were enriched in the system, leading to improved removal capacity for NH4+-N and TN. Under optimal condition, the removal rates of NH4+-N and TN were 94.0 % and 78.7 % respectively. The dominant species of HN-AD bacteria were influenced by OLR and pH. The HN-AD bacteria exhibited the capacity for heterotrophic nitrification and aerobic denitrification within the system. This study provides a technological basis for the application of HN-AD bacteria for wastewater treatment.

Nitrogen removal by heterotrophic nitrification-aerobic denitrification bacteria: A review

The rising urbanization and industrialization are increasing nitrogen pollution in water and wastewaters, calling for advanced remediation treatments. Traditional biological treatments are limited in area and efficiency. Therefore, heterotrophic nitrification-aerobic denitrification (HN-AD) in a single reactor has drawn significant attention and has shown higher efficiency. Here we review HN-AD bacteria with focus on bacterial species, nitrogen metabolism, and factors influencing denitrification. Nitrogen metabolism pathway includes nitrate-nitrite reduction denitrification, hydroxylamine reduction and assimilatory denitrification. Factors influencing denitrification comprise carbon source, C/N, temperature, pH, DO and so on. Compared to published literature reviews on HN-AD, our manuscript comprehensively summary the latest research on its nitrogen remove mechanisms and related enzymes for the first time.

Metabolomic insight into regulatory mechanism of heterotrophic bacteria nitrification-aerobic denitrification bacteria to high-strength ammonium wastewater treatment

This work aimed to elucidate the metabolic mechanism of heterotrophic nitrification-aerobic denitrification (HN-AD) bacteria influenced by varying concentrations of ammonium nitrogen (NH4+-N) in high-strength synthetic wastewater treatment. The results showed that the removal rates of NH4+-N and total nitrogen, along with enzymatic activities related to nitrification and denitrification, increased with rising NH4+-N concentrations (N500:500 mg/L, N1000:1000 mg/L and N2000:2000 mg/L). The relative abundances of HN-AD bacteria were 50 %, 62 % and 82 % in the three groups. In the N2000 group, the cAMP signaling pathway, glycerophospholipid metabolites, purines and pyrimidines related to DNA/RNA synthesis, electron donor NAD+-related energy, the tricarboxylic acid (TCA) cycle and glutamate metabolism were upregulated. Therefore, influent NH4+-N at 2000 mg/L promoted glutamate metabolism to accelerate the TCA cycle, and enhanced cellular energy and advanced denitrification activity of bacteria for HN-AD. This mechanism, in turn, enhanced microbial growth and the carbon and nitrogen metabolism of bacteria for HN-AD.

A novel pure biofilm system based on aerobic denitrification for nitrate wastewater treatment: Exploring the feasibility of high total nitrogen removal under low-carbon condition

To improve the poor denitrification efficiency of traditional biological treatment for low-carbon wastewater, a novel pure biofilm moving bed biofilm reactor (PH-MBBR) based on heterotrophic nitrification-aerobic denitrification (HN-AD) bacteria was proposed. The start-up of PH-MBBR inoculated with HN-AD bacteria and the operating performances and microbial community composition under various C/N ratios were studied. The results showed that at C/N ratio of only 4, the total nitrogen (TN) removal rate of 10.02 mg/(L·h) was achieved under aerobic conditions. Microbial analysis indicates that the synergistic effect of oligotrophic bacteria Thauera and Azoarcus was the main reason for the high pollutant removal at C/N ratio of 4. Furthermore, the analysis of functional genes and enzymes reveals that NorB and NosZ played important roles in nitrate metabolism in PH-MBBR, and aerobic denitrification pathway was NO3–-N → NO → N2O → N2. This study provides a practical and theoretical basis for low-carbon nitrate wastewater treatment.

Feast/famine ratio regulates the succession of heterotrophic nitrification-aerobic denitrification and autotrophic ammonia oxidizing bacteria in halophilic aerobic granular sludge treating saline wastewater

Heterotrophic nitrification-aerobic denitrification (HN-AD) shows innovation potential of wastewater treatment process in a single tank. However, how to enrich HN-AD bacteria in activated sludge to enhance their contribution remained unknown. This study explored the impact of the feast/famine (F/F) ratio on the succession of autotrophic ammonia oxidizing bacteria (AOB) and HN-AD bacteria in a halophilic aerobic granular sludge (HAGS) system. As the F/F ratio decreased from 1/9 to 1/15, the total inorganic nitrogen (TIN) removal performance significantly decreased. The proportion of heterotrophic bacteria was dropped from 79.0 % to 33 %. Accordingly, the relative abundance of Paracoccus decreased from 70.8 % to 25.4 %, and the copy number of the napA gene was reduced from 2.2 × 1010 copies/g HAGS to 8.1 × 109 copies/g HAGS. It found the F/F ratio regulated the population succession of autotrophic AOB and HN-AD bacteria, thereby providing a solution to achieve the enrichment of HN-AD bacteria in HAGS.

Indigenous mixotrophic aerobic denitrifiers stimulated by oxygen micro/nanobubble-loaded microporous biochar

The prevalence of hypoxia in surface sediment inhibits the growth of aerobic denitrifiers in natural waters. A novel oxygen micro/nanobubble-loaded microporous biochar (OMB) was developed to activate indigenous aerobic denitrifiers in this study. The results indicate a thin-layer OMB capping mitigates hypoxia effectively. Following a 30-day microcosm-based incubation, a 60 % decrease in total nitrogen concentration was observed, and the oxygen penetration depth in the sediment was increased from <4.0 mm to 38.4 mm. High-throughput sequencing revealed the stimulation of indigenous mixotrophic aerobic denitrifiers, including autotrophic denitrifiers such as Hydrogenophaga and Thiobacillus, heterotrophic denitrifiers like Limnobacter and unclassified_f_Methylophilaceae, and heterotrophic nitrification aerobic denitrification bacteria, including Shinella and Acidovorax, with total relative abundance reaching up to 38.1 %. Further analysis showed OMB enhanced the overall collaborative relationships among microorganisms and promoted the expression of nitrification- and denitrification-related genes. This study introduces an innovative strategy for stimulating indigenous aerobic denitrifiers in aquatic ecosystems.

Microalgae-assisted heterotrophic nitrification-aerobic denitrification process for cost-effective nitrogen and phosphorus removal from high-salinity wastewater: Performance, mechanism, and bacterial community

A microalgae-assisted heterotrophic nitrification-aerobic denitrification (HNAD) system for efficient nutrient removal from high-salinity wastewater was constructed for the first time as a cost-effective process in the present study. Excellent nutrient removal (∼100.0 %) was achieved through the symbiotic system. The biological removal process, biologically induced phosphate precipitation (BIPP), microalgae uptake, and ammonia stripping worked together for nutrient removal. Furthermore, the biological removal process achieved by biofilm contributed to approximately 55.3–71.8 % of nitrogen removal. BIPP undertook approximately 45.6–51.8 % of phosphorus removal. Batch activity tests confirmed that HNAD fulfilled an extremely critical role in nitrogen removal. Microalgal metabolism drove BIPP to achieve efficient phosphorus removal. Moreover, as the main HNAD bacteria, OLB13 and Thauera were enriched. The preliminary energy flow analysis demonstrated that the symbiotic system could achieve energy neutrality, theoretically. The findings provide novel insights into strategies of low-carbon and efficient nutrient removal from high-salinity wastewater.