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As steel production increases, large volumes of highly toxic and nitrogen-rich coking wastewater (CWW) are produced, prompting the development of a novel oxic-hydrolytic-oxic (OHO) biological treatment combination designed for highly efficient removal of nitrogen-contained contaminants. However, previous studies have not comprehensively explored the CWW biotreatment from the perspective of nitrogen metabolism functional genes and pathways. Based on the investigation of taking the full-scale OHO biotreatment combination as a case, it was found that the O1 and O2 bioreactors remove nitrogen through the ammonia assimilation accounting for 33.87% of the total nitrogen (TN) removal rate, while the H bioreactor removes nitrogen through the simultaneous nitrification-denitrification accounting for 61.11% of the TN removal rate. The major ammonia assimilation taxa include Thauera, Immundisolibacter and Thiobacillus; the major nitrifying taxa include Nitrospira and Nitrosomonas; and the major denitrifying taxa include Thiobacillus, Lautropia and Mesorhizobium. Additionally, the H bioreactor exhibits the potential to be optimized for simultaneous nitrification-denitrification coupled with anaerobic ammonium oxidation (Anammox). These understandings will guide the optimization of engineering design and operational practices, contributing to more effective and sustainable wastewater treatment strategies.

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Anammox is recognized as a prospective alternative for future biological nitrogen removal technologies. However, the nitrate by-products produced by anammox bacteria limit its overall nitrogen removal efficiency below 88 %. This study introduced Fe(III) into the anammox bioreactor to enhance the nitrogen removal efficiency to approximately 95 %, surpassing the biochemical limit of 88 % imposed by anammox stoichiometry. Anammox sludge was demonstrated to utilize extracellular polymeric substances to reduce Fe(III) into Fe(II), and this process promoted the dominance of Ca. Brocadia. The iron addition improved the abundance of narGHI genes and facilitated the partial dissimilatory nitrate reduction to ammonium, with nitrite as the end product. The accumulated nitrite was then eliminated through the anammox pathway, along with the excess ammonium (30 mg/L) in the influent. Overall, this study deepens our understanding of the enhanced nitrogen removal triggered by Fe(III) in anammox sludge and offers an effective approach to boost anammox process.

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This study investigates the performance, resilience and microbial community dynamics of two anaerobic processes, i.e. pure anammox (R1) and partial denitrification/anammox (PD/A) (R2), following a 30-day starvation period. The tolerance to starvation was assessed by comparing nitrogen removal efficiency and microbial activity across both reactors. Results show that the PD/A process recovery to pre-starvation performance levels within just one day, as compared to the pure anammox process. Notably, although the activity of anammox bacteria decreased in both processes during starvation, the decay rate in R1 was 69.59 % higher than in R2, potentially explaining the quicker recovery of R2. Furthermore, enhanced secretion of extracellular polymeric substance (EPS) during starvation served as a protective mechanism. The potential functions and genes in microorganisms, as well as the pathway of nitrogen cycling, were demonstrated through analyses using the KEGG database. This research reveals essential mechanistic insights and strategic guidance for the effective implementation of anammox-based biological nitrogen removal processes.

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Swine wastewater (SW) contains high levels of traditional pollutants, antibiotics, and antibiotic resistance genes (ARGs), necessitating effective elimination. Two parallel aerobic granular sludge (AGS) reactors, R1 and R2, were constructed and optimized for treating SW from two pig farms, identified as SW1 and SW2. R2 showed higher antibiotic removal efficiency, particularly in the removal of sulfonamides, while fluoroquinolones tended to adsorb onto the sludge. Process optimization by introducing an additional anoxic phase enhanced denitrification and reduced effluent ARG levels, also aiding in the improved removal of fluoroquinolones. The nitrite-oxidizing bacteria (NOB) Nitrospira accumulated after the treatment process, reaching 12.8 % in R1 and 14.1 % in R2, respectively. Mantel's test revealed that pH, NH4+-N, and Mg significantly affected ARGs and microbial community. Sulfadiazine and sulfamethazine were found to significantly impact ARGs and the microbial communities. This study provides innovative insights into the application of AGS for the treatment of real SW.

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Nitrification is highly crucial for both anammox systems and the global nitrogen cycle. The discovery of complete ammonia oxidation (comammox) challenges the inherent concept of nitrification as a two-step process. Its wide distribution, adaptability to low substrate environments, low sludge production, and low greenhouse gas emissions may make it a promising new nitrogen removal treatment process. Meanwhile, anammox technology is considered the most suitable process for future wastewater treatment. The diverse metabolic capabilities and similar ecological niches of comammox bacteria and anammox bacteria are expected to achieve synergistic nitrogen removal within a single system. However, previous studies have overlooked the existence of comammox, and it is necessary to re-evaluate the conclusions drawn. This paper outlined the ecophysiological characteristics of comammox bacteria and summarized the environmental factors affecting their growth. Furthermore, it focused on the enrichment, regulatory strategies, and nitrogen removal mechanisms of comammox and anammox, with a comparative analysis of hydroxylamine, a particular intermediate product. Overall, this is the first critical overview of the conclusions drawn from the last few years of research on comammox-anammox, highlighting possible next steps for research.

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The current work investigated the performance of an Integrated Fixed-Film Activated Sludge Sequencing Batch Reactor (IFAS-SBR) for Biological Nitrogen Removal (BNR) from mature landfill leachate through the nitritation-denitritation process. During the experimental period two IFAS-SBR configurations were examined using two different biocarrier types with the same filling ratio (50%). The dissolved oxygen (DO) concentration ranged between 2 and 3 mg/L and 4-6 mg/L in the first (baseline-IFAS) and the second (S8-IFAS) setup, respectively. Baseline-IFAS operated for 542 days and demonstrated a high and stable BNR performance maintaining a removal efficiency above 90% under a Nitrogen Loading Rate (NLR) up to 0.45 kg N/m3-d, while S8-IFAS, which operated for 230 days, was characterized by a limited and unstable BNR performance being unable to operate sufficiently under an NLR higher than 0.20 kg N/m3-d. It also experienced a severe inhibition period, when the BNR process was fully deteriorated. Moreover, S8-IFAS suffered from extensive biocarrier stagnant zones and a particularly poor sludge settleability. The attached biomass cultivated in both IFAS configurations had a negligible content of nitrifying bacteria, probably attributed to the insufficient DO diffusion through the biofilm, caused by the low DO concentration in the liquid in the baseline case and the extensive stagnant zones in the S8-IFAS case. As a result of the high biocarrier filling ratio, the S8-IFAS was unstable and low. This was probably attributed to the mass transfer limitations caused by the biocarrier stagnant zones, which hinder substrate and oxygen diffusion, thus reducing the biomass activity and increasing its vulnerability to inhibitory and toxic factors. Hence, the biocarrier filling fraction is a crucial parameter for the efficient operation of the IFAS-SBR and should be carefully selected taking into consideration both the media type and the overall reactor configuration.

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The nitrogen cycle plays a vital role in maintaining ecological health and biodiversity. In aquatic systems, nitrogen transformation genes significantly contribute to biological nitrogen cycling. Although the function of these genes is known to be influenced by environmental factors, there is limited research exploring the relationship between nitrogen transformation genes and environmental factors. Therefore, the correlations, between nitrogen transformation genes and environmental factors, were investigated at the estuaries of Chaohu lake (China) in different seasons. The results showed that the values of temperature, pH, organic compounds, nitrogen, and dissolved oxygen were higher in dry season, whereas the abundance of the genes was lower in dry season. In addition, the abundance of the anaerobic ammoxidation gene was much lower than the nitrification gene and denitrification gene. The results indicated that biological nitrification and denitrification were the primary mechanisms for nitrogen removal at estuaries in different seasons, and the reduction of nitric oxide may be a limiting step in the denitrification process. The Co-occurrence Network and Mantel test indicated that, during the dry season, the temperature was the primary driver of ammonification and nitrification functions, the NO3- and NO2- were the primary drivers of denitrification, and the total nitrogen (TN) and NH4+ were the main drivers of anaerobic ammonia oxidation. During the wet season, the dissolved oxygen was the primary driver of ammonification and nitrification functions, the chemical oxygen demand was the primary driver of denitrification, and the TN was the main driver of anaerobic ammonia oxidation. This study provides valuable insights into nitrogen cycling in surface water, contributing to a better understanding of this important process.

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The DEnitrifying AMmonium OXidation (DEAMOX) has been proven to be a promising process treating contaminated surface water containing ammonia and nitrate, while the enrichment of the slow-growing anammox bacteria (AnAOB) remains a challenge. In this study, a novel polyurethane-adhesion vermiculite/tourmaline (VTP) modified carrier was developed to achieve effective enrichment of AnAOB. The results demonstrated that the VTP-1 (vermiculite: tourmaline = 1:1) system exhibited the greatest performance with the total nitrogen removal efficiency reaching 87.6% and anammox contributing 63% to nitrogen removal. Scanning electron microscope analysis revealed the superior biofilm structure of the VTP-1 carrier, providing attachment for AnAOB. The addition of VTP-1 promoted the secretion of EPS (extracellular polymeric substances) by microorganisms, which increased to 85.34 mg/g VSS, contributing to the aggregation of anammox cells. The favorable substrate microenvironment created by NH4+ adsorption and NO2- supply via partial denitrification process facilitated the growth of AnAOB. The relative abundance of Candidatus Brocadia and Thauera increased from 0.04% to 0.3%-1.03% and 2.06% in the VTP-1 system, respectively. This study sheds new light on the anammox biofilm formation and provides a valid approach to initiate the DEAMOX process for low nitrogen polluted water treatment.

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Aerobic granular sludge (AGS) capable of nitrogen and phosphorus removal is mainly limited to the applications in sequencing batch reactors. This study introduced an innovative continuous self-circulating up-flow fluidized bed process (Zier process) using separate aeration. The process was composed of an anoxic column (Zier-A), aeration column (Zier-OO) and aerobic column (Zier-O), and was used to treat actual municipal sewage continuously for 170 days. The process achieved self-circulation of 20-32 times and an up-flow velocity within the reactor of 7-16 m/h which were accurately controlled with only separate aeration. The larger proportion of self-circulating multiple times contributed to particle formation and stability, providing hydraulic shear conditions, and screened the precipitation performance of the granular sludge (GS). Meanwhile, the dissolved oxygen (DO) of Zier-O was controlled at 0.1-0.3 mg/L, and the DO of Zier-A input water was zero. The accurate oxygen supply enhanced simultaneous nitrification and denitrification (SND) as well as short-cut nitrification and denitrification in Zier-O and improved the COD utilization rate and the nitrogen removal rate in Zier-A. The COD treatment capacity reached 2.46 kg-COD/(m³·d). With a hydraulic retention time of 10 h, the process consistently ensured that the average concentrations of ammonia nitrogen and total nitrogen in the effluent were maintained below 5 and 15 mg/L, respectively. Moreover, the process maintained the shape and stability of GS, the median diameter of GS ranged between 300-1210 µm, the percentage of mass with particle size distribution < 200 µm at a height of 150 cm within Zier-A and Zier-O accounted for as low as 0.04%-0.05%, and showed good settling performance. The suspended solids in effluent can be maintained at 50-80 mg/L. Overall, the unique structural setting and control method of the Zier process provide another approach for the application of continuous AGS treatment for municipal sewage.

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Sulfur-driven autotrophic denitrification (SAD) process is a cost-effective and sustainable method for nitrogen removal from wastewater. However, a higher concentration of zinc ions (Zn(II)) flowing into wastewater treatment plants poses a potential threat to the SAD process. This study examined that a half maximal inhibitory concentration (IC50) of Zn(II) was 7 mg·L-1 in the SAD process. Additionally, the addition of 20 mg·L-1 Zn(II) resulted in a severe accumulation of nitrite to 150.20 ± 6.00 mg·L-1 when the initial concentration of nitrate was 500 mg·L-1. Moreover, the activities of nitrate reductase, nitrite reductase, dehydrogenase and electron transport system were significantly inhibited under Zn(II) stress. The addition of Zn(II) inhibited EPS secretion and worsened electrochemical properties. The result was attributed to the spontaneous binding between EPS and Zn(II), with a ΔG of -17.50 KJ·mol-1 and a binding constant of 1.77 × 104 M-1, respectively. Meanwhile, the protein, fulvic acid, and humic-like substances occurred static quenching after Zn(II) addition, with -OH and -C=O groups providing binding sites. The binding sequence was fulvic acid→protein→humic acid and -OH â†’ -C=O. Zn(II) also reduced the content of α-helix, which was unfavorable for electron transfer. Additionally, the Zn(II) loosened protein structure, resulting in a 50 % decrease in α-helix/(ß-sheet+random coil). This study reveals the effect of Zn(II) on the SAD process and enhances our understanding of EPS behavior under metal ions stress.

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The substantial discharge of ferroferric oxide nanoparticles (Fe3O4 NPs) into sewage threatens the survival of functional microorganisms in wastewater treatment. This study elucidated responses of anaerobic ammonium oxidation (anammox) consortia to inhibition from high Fe3O4 NPs concentration and recovery mechanisms. The nitrogen removal efficiency decreased by 20.3 % and recovered after 55 days under 1000 mg/L Fe3O4 NPs concentration. Toxicity was attributed to reactive oxygen species (ROS) production. The excessive ROS damaged membrane integrity, nitrogen metabolism, and DNA synthesis, resulting in the inhibition of anammox bacteria activity. However, recovery mechanisms of anammox consortia activity were activated in response to 1000 mg/L Fe3O4 NPs. The increase of heme oxygenase-1, thioredoxin, and nicotinamide adenine dinucleotide-quinone oxidoreductase genes alleviated oxidative stress. Furthermore, the activation of metabolic processes associated with membrane and DNA repair promoted recovery of anammox bacteria activity. This study provided new insights into NPs contamination and control strategies during anammox process.

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N2O as a potent greenhouse gas often generates in the biological nitrogen removal (BNR) processes during wastewater treatment, which makes BNR become an important greenhouse gas emission source. The emerging pollutants (EPs) are ubiquitous in wastewater and they have shown to influence the BNR processes. However, the deep discussion on potential impacts of EPs on N2O emissions during BNR is rare. Moreover, the experimental parameters for EPs investigation in most of literatures are generally not in line with real-world BNR processes, which calls for deep elucidating the roles of EPs on N2O production and emission. In this work, a critical review summarizes the existing literature about influences of typical EPs on N2O emissions and associated mechanisms during BNR, and it discusses the impacts of some easily overlooked factors, such as real EPs environmental concentrations, EPs bioaccumulation, and multiple EPs coexistence on N2O emissions. This review will provide an insight into exploring and mitigating threats posed by typical EPs on N2O emissions.

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The partial denitrification (PD) coupled with anaerobic ammonium oxidation (Anammox) (PD/A) process is a unique biological denitrification method for sewage that concurrently removes nitrate (NO3--N) and ammonium (NH4+-N) in sewage. Comparing PD/A to conventional nitrification and denitrification technologies, noticeable improvements are shown in energy consumption, carbon source demand, sludge generation and emissions of greenhouse gasses. The PD is vital to obtaining nitrites (NO2--N) in the Anammox process. This paper provided valuable insight by introduced the basic principles and characteristics of the process and then summarized the strengthening strategies. The functional microorganisms and microbial competition have been discussed in details, the S-dependent denitrification-anammox has been analyzed in this review paper. Important factors affecting the PD/A process were examined from different aspects, and finally, the paper pointed out the shortcomings of the coupling process in experimental research and engineering applications. Thus, this research provided insightful information for the PD/A process's optimization technique in later treating many types of real and nitrate-based wastewater. The review paper also provided the prospective economic and environmental position for the actual design implementation of the PD/A process in the years to come.

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One-stage partial nitrification coupled with anammox (PN/A) technology effectively reduces the energy consumption of a biological nitrogen removal system. Inhibiting nitrite-oxidizing bacteria (NOB) is essential for this technology to maintain efficient nitrogen removal performance. Initial ammonium concentration (IAC) affects the degree of inhibited NOB. In this study, the effect of the IAC on a PN/A biofilm was investigated in a moving bed biofilm reactor. The results showed that nitrogen removal efficiency decreased from 82.49% ± 1.90% to 64.57% ± 3.96% after the IAC was reduced from 60 to 20 mg N/L, while the nitrate production ratio increased from 13.87% ± 0.90% to 26.50% ± 3.76%. NOB activity increased to 1,133.86 mg N/m2/day after the IAC decreased, approximately 4-fold, indicating that the IAC plays an important inhibitory role in NOB. The rate-limiting step in the mature biofilm of the PN/A system is the nitritation process and is not shifted by the IAC. The analysis of the microbial community structure in the biofilm indicates that the IAC was the dominant factor in changes in community structure. Ca. Brocadia and Ca. Jettenia were the main anammox bacteria, and Nitrosomonas and Nitrospira were the main AOB and NOB, respectively. IAC did not affect the difference in growth between Ca. Brocadia and Ca. Jettenia. Thus, modulating the IAC promoted the PN/A process with efficient nitrogen removal performance at medium to low ammonium concentrations.

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Chlorella sp. and Navicula sp. were separately used to construct an algal-bacterial symbiotic system in two identical sequencing batch reactors (R1 and R2) to explore the influence of algal species differences on nitrite accumulation. The Navicula-bacterial symbiotic system showed a higher nitrite accumulation efficiency of 85% and a stronger resistance to ammonia load. It secreted twice as many extracellular polymeric substances than the Chlorella-bacterial symbiotic system. Nitrospira and SM1A02 were the dominant functional genera of nitrite-oxidizing bacteria in R1. The dominant functional genus of ammonium-oxidizing bacteria and the dominant functional genus of denitrifying bacteria were Ellin6067 and unclassified_Saprospiraceae in R2, respectively. In general, this research provided some reference for the construction of an algal-bacterial symbiotic system and achieving nitrite accumulation through an algal-bacterial symbiotic system.

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Studies published recently proposed that ammonia-oxidizing archaea (AOA) may be beneficial for hypersaline (salinity > 50 g NaCl L-1) industrial wastewater treatment. However, knowledge of AOA activity in hypersaline bioreactors is limited. This study investigated the effects of salinity, organic matter, and practical pickled mustard tuber wastewater (PMTW) on AOA and ammonia-oxidizing bacteria (AOB) in two sequencing batch biofilm reactors (SBBRs). Results showed that despite observed salinity inhibition (p < 0.05), both AOA and AOB contributed to high ammonia removal efficiency at a salinity of 70 g NaCl L-1 in the two SBBRs. The ammonia removal efficiency of SBBR2 did not significantly differ from that of SBBR1 in the absence of organic matter (p > 0.05). Batch tests and quantitative real-time PCR (qPCR) reveal that salinity and organic matter inhibition resulted in a sharp decline in specific ammonia oxidation rates and amoA gene copy numbers of AOA and AOB (p < 0.05). AOA demonstrated higher abundance and more active ammonia oxidation activity in hypersaline and high organic matter environments. Salinity was positively correlated with the potential ammonia oxidation contribution of AOA (p < 0.05), resulting in a potential transition from AOB dominance to AOA dominance in SBBR1 as salinity levels rose. Moreover, autochthonous AOA in PMTW promoted the abundance and ammonia oxidation activities of AOA in SBBR2, further elevating the nitrification removal efficiency after feeding the practical PMTW. AOA demonstrates greater tolerance to the challenging hypersaline environment, making it a valuable candidate for the treatment of practical industrial wastewater with high salinity and organic content.

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Sludge alkaline fermentation liquid (SAFL) is a promising alternative to acetate for improving biological nitrogen removal (BNR) from wastewater. SAFL inevitably contains some refractory compounds, while the characteristics of dissolved organic matter (DOM) in effluent from SAFL-fed BNR process remain unclear. In this study, the molecular weight distribution, fluorescent composition and molecular profiles of DOM in effluent from SAFL and acetate-fed sequencing batch reactors (S-SBRs and A-SBRs, respectively) at different hydraulic retention time (12 h and 24 h) was comparatively investigated. Two carbon sources resulted in similar effluent TN, but a larger amount of DOM, which was bio-refractory or microorganisms-derived, was found in effluent of S-SBRs. Compared to acetate, SAFL increased the proportion of large molecular weight organics and humic-like substances in effluent DOM by 74.87%-101.3% and 37.52%-48.35%, respectively, suggesting their bio-refractory nature. Molecular profiles analysis revealed that effluent DOM of S-SBRs exhibited a more diverse composition and a higher proportion of lignin-like molecules. Microorganisms-derived molecules were found to be the dominant fraction (71.51%-72.70%) in effluent DOM (<800 Da) of S-SBRs. Additionally, a prolonged hydraulic retention time enriched Bacteroidota, Haliangium and unclassified_f_Comamonadaceae, which benefited the degradation of DOM in S-SBRs. The results help to develop strategies on reducing effluent DOM in SAFL-fed BNR process.

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Nitrous oxide (N2O) and nitrogen oxides (NOx) (i.e., nitric oxide (NO) and nitrogen dioxide (NO2)), which could be produced in wastewater treatment process and result in greenhouse effect and atmospheric pollution, respectively, have been studied limitedly in their emission characteristics and transformation mechanisms. In this study, intelligent oxygen regulation was applied in anoxic/oxic wastewater treatment process (I-A/O), and its effects on regulating NOx and N2O transformations were extensively explored by comparing it with conventional A/O process (C-A/O). Results showed that the average emission amounts of N2O and NOx in I-A/O were 7.45 ± 0.66 mg and 1.88 ± 0.10 mg, respectively. Satisfactory reduction of N2O by 29.28 %-45.08 % was achieved in I-A/O compared to that of C-A/O, but together with increased NOx emission by 83.19 %-120.57 %. Pearson correlation and transcriptional analysis suggested that NO2--N reduction in the anoxic phase dominated N2O production, while no significant N2O production in the oxic phase was found. Hence, the reduced N2O production in I-A/O was mainly attributed to its efficient denitrification process. On the other hand, both the anoxic and oxic phases played important roles in NO production. More importantly, sufficient oxygen in I-A/O promoted the ammonia oxidation process, resulting in higher NO emission in I-A/O in the oxic phase. The imbalance in NO and N2O emissions was then amplified by the NOR enzyme, which mediates the conversion of NO to N2O in both the anoxic and oxic phases. Besides, carbon emission reduction by 31.32 %-36.50 % was obtained in I-A/O due to aeration consumption savings and greenhouse gas emissions reduction compared to C-A/O. Overall, intelligent oxygen regulation optimized the nitrogen transformation and achieved carbon emission reduction in A/O process, but special attention should be paid to the associated risk caused by increased NO emissions.

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Anaerobic ammonia oxidation (anammox) of municipal wastewater is a research focus, especially the combined treatment with mature landfill leachate is a current research hotspot. In this study, municipal wastewater was treated by partial nitrification via sequencing batch reactor (SBR), and its effluent and mature landfill leachate were then mixed into an up-flow anaerobic sludge blanket (UASB) for simultaneous anammox and partial denitrification reaction. Through partial nitrification, a high nitrite accumulation rate (93.0 ± 3.8 %) was achieved by low dissolved oxygen (0.5-1.6 mg/L) and controlled aerobic time (3.5 h) in SBR. The UASB system was responsible for 78.8 ± 2.1 % nitrogen removal of the entire system with a hydraulic reaction time (HRT) of 3.8 h, accompanied by the anammox contribution up to 89.4 ± 6.0 %. The overall partial nitrification-simultaneous anammox and partial denitrification (PN-SAPD) system was controlled at a total COD/TIN of 2.8 ± 0.3 and a total HRT of only 10.2 h, achieving the nitrogen removal efficiency and effluent TIN were 95.2 ± 2.2 % and 3.4 ± 1.5 mg/L, respectively. The qPCR results showed functional genes (hzsA(B), hdh) associated with anaerobic ammonia-oxidizing bacteria (AnAOB), whose high gene copy abundance and transcription expression ensured the removal of major nitrogen from municipal wastewater and mature landfill leachate. 16S amplicon sequencing showed that the Ca. Brocadia (9.72-12.6 %) was further enrichment after sodium acetate was added, and the transcription expression of Thauera (0.5-7.0 %) caused nitrate to nitrite. The high abundance of related enzymes (hao, hzs, hdh, narGHI) involved in anammox and partial denitrification processes were found in the macrogenomic sequencing, and only Ca. Brocadia was involved in multi-pathway nitrogen metabolism in AnAOB. Based on the efficient nitrogen removal by AnAOB and denitrifying bacteria, this modified PN-SAPD process provides a new option for the co-treatment of mature landfill leachate in municipal wastewater treatment plants.

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Sulfur-driven autotrophic denitrification (SAD) granular process has significant advantages in treating low-carbon/nitrogen wastewater; however, the slow growth rate of sulfur-oxidizing bacteria (SOB) results in a prolonged start-up duration. In this study, the thiosulfate-driven autotrophic denitrification (TAD) was successfully initiated by inoculating anaerobic granular sludge on Day 7. Additionally, the electron donor was successfully transferred to the cheaper elemental sulfur from Day 32 to Day 54 at the nitrogen loading rate of 176.2 g N m-3 d-1. During long term experiment, the granules maintained compact structures with the α-helix/(ß-sheet + random coil) of 29.5-40.1 %. Extracellular electron transfer (EET) pathway shifted from indirect to direct when electron donors were switched thiosulfate to elemental sulfur. Microbial analysis suggested that thiosulfate improved EET involving enzymes activity. Thiobacillus and Sulfurimonas were dominant in TAD, whereas Longilinea was enriched in elemental sulfur-driven autotrophic denitrification. Overall, this strategy achieved in-situ enrichment of SOB in granules, thereby shortening start-up process.

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