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Electroactive microorganisms (EAMs) play a vital role in biogeochemical cycles by facilitating extracellular electron transfer. They demonstrate remarkable adaptability to river sediments that are characterized by pollution and poor water quality, significantly contributing to the sustainability of river ecosystems. However, the distribution and diversity of EAMs remain poorly understood. In this study, 16S rRNA gene high-throughput sequencing and real-time fluorescence quantitative PCR were used to assess EAMs in 160 samples collected from eight rivers within the Pearl River Delta of Southern China. The results indicated that specialized EAMs communities in polluted sediments exhibited variations in response to water quality and sediment depth. Compared to clean sediment, polluted sediments showed a 4.5% increase in the relative abundances of EAMs communities (59 genera), with 45- and 17-times higher abundances of Geobacter and cable bacteria. Additionally, the abundance of cable bacteria decreased with increasing sediment depth in polluted sediments, while the abundance of L. varians GY32 exhibited an opposite trend. Finally, the abundances of Geobacter, cable bacteria, and L. varians GY32 were positively correlated with the abundance of filamentous microorganisms (FMs) across all samples, with stronger interactions in polluted sediments. These findings suggest that EAMs demonstrate heightened sensitivity to polluted environments, particularly at the genus (species) level, and exhibit strong adaptability to conditions characterized by high levels of acid volatile sulfide, low dissolved oxygen, and elevated nitrate nitrogen. Therefore, environmental factors could be manipulated to optimize the growth and efficiency of EAMs for environmental engineering and natural restoration applications.

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Electrobioremediation is one of the most innovative disciplines for treating organic pollutants and it is based on the ability of electroactive bacteria to exchange electrons with electroconductive materials. Electroactive biofilters have been demonstrated to be efficient for treating urban wastewater with a low footprint; however, their application can be expanded for treating industrial wastewater containing significant concentrations (2.4 %vol) of commercial surfactants (containing lauryl sulfate, lauryl ether sulfate, cocamydopropyl betaine, and dodecylbenzene sulfonate, among others). Our electroactive biofilter outperformed a conventional inert biofilter made of gravel for all tested conditions, reaching removal rates as high as 4.5 kg COD/m3bed·day and withstood Organic Loading Rates as high as 9 Kg COD/m3·d without significantly affecting removal efficiency. The biomass accumulation reduced available bed volume in the electroactive biofilter just by 39 %, while the gravel biofilter decreased by 80 %. Regarding microbial communities, anaerobic and electroactive bacteria represented a substantial proportion of the total population in the electroactive biofilter. Pseudomonas was the dominant genus, while Cupriavidus, Shewanella, Citrobacter, Desulfovibrio, and Arcobacter were potential electroactive strains found in relevant proportions. The microbial community's composition might be the key to understanding how high removal rates can coexist with limited biomass production, making electroactive biofilters a promising strategy to overcome classical biofilter limitations.

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By integrating electroactive genes into engineered sensing microorganisms, information about the object to be measured can be converted into the output of an electrical signal, omitting the process of converting the output of an electrical signal in conventional sensing strategies and simplifying the steps of biosensor development. By utilizing synthetic biology methods, we can not only create novel genetic circuits by using logic gate operations and integrating genes from other biological components, solving biosensing issues in living systems and enhancing sensor performance, but also convert various types of genetic circuits into electrical signals, broadening the application range of biosensors. Here, we describe an example of how to genetically engineer microorganisms with electroactive genes and the fabrication of an electrochemical microbial biosensor.

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Microorganisms are key players in the global biogeochemical sulfur cycle. Among them, some have garnered particular attention due to their electrical activity and ability to perform extracellular electron transfer. A growing body of research has highlighted their extensive phylogenetic and metabolic diversity, revealing their crucial roles in ecological processes. In this review, we delve into the electron transfer process between sulfate-reducing bacteria and anaerobic alkane-oxidizing archaea, which facilitates growth within syntrophic communities. Furthermore, we review the phenomenon of long-distance electron transfer and potential extracellular electron transfer in multicellular filamentous sulfur-oxidizing bacteria. These bacteria, with their vast application prospects and ecological significance, play a pivotal role in various ecological processes. Subsequently, we discuss the important role of the pili/cytochrome for electron transfer and presented cutting-edge approaches for exploring and studying electroactive microorganisms. This review provides a comprehensive overview of electroactive microorganisms participating in the biogeochemical sulfur cycle. By examining their electron transfer mechanisms, and the potential ecological and applied implications, we offer novel insights into microbial sulfur metabolism, thereby advancing applications in the development of sustainable bioelectronics materials and bioremediation technologies.

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A biological treatment is the core process for removing organic pollutants from industrial wastewater. However, industrial wastewater often contains large amounts of toxic and harmful pollutants, which can inhibit the activity of microorganisms in a treatment system, precipitate the deterioration of effluent quality, and threaten water ecological security from time to time. In most of the existing anaerobic biological treatment processes, toxic effects on microorganisms are determined according to the amounts of end-products of the biochemical reactions, and the evaluation results are relatively lacking. When microorganisms contact toxic substances, changes in biological metabolic activity precede the accumulation of reaction products. As sensitive units, electroactive microorganisms can generate electrical signals, a change in which can directly reflect the toxicity level. The applications of electroactive microorganisms for the toxicity monitoring of wastewater are very promising. Further attention needs to be paid to considering the appropriate evaluation index, the influence of the environment on test results, mechanisms, and other aspects. Therefore, we reviewed the literature regarding the above aspects in order to provide a research foundation for the practical application of electroactive microorganisms in toxicant monitoring.

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Bidirectional extracellular electron transfer (EET) is crucial to upholding microbial metabolism with insoluble electron acceptors or donors in anoxic environments. Investigating bidirectional EET-capable microorganisms is desired to understand the cell-cell and microbe-mineral interactions and their role in mineral cycling besides leveraging their energy generation and conversion, biosensing, and bio-battery applications. Here, we report on iron cycling by haloalkaliphilic Geoalkalibacter halelectricus via bidirectional EET under haloalkaline conditions. It efficiently reduces Fe3+ oxide (Fe2O3) to Fe0 at a 0.75 ± 0.08 mM/mgprotein/d rate linked to acetate oxidation via outward EET and oxidizes Fe0 to Fe3+ at a 0.24 ± 0.03 mM/mgprotein/d rate via inward EET to reduce fumarate. Bioelectrochemical cultivation confirmed its outward and inward EET capabilities. It produced 895 ± 23 µA/cm2 current by linking acetate oxidation to anode reduction via outward EET and reduced fumarate by drawing electrons from the cathode (‒2.5 ± 0.3 µA/cm2) via inward EET. The cyclic voltammograms of G. halelectricus biofilms revealed redox moieties with different formal potentials, suggesting the involvement of different membrane components in bidirectional EET. The cyclic voltammetry and GC-MS analysis of the cell-free spent medium revealed the lack of soluble redox mediators, suggesting direct electron transfer by G. halelecctricus in achieving bidirectional EET. By reporting on the first haloalkaliphilic bacterium capable of oxidizing and reducing insoluble Fe0 and Fe3+ oxide, respectively, this study advances the limited understanding of the metabolic capabilities of extremophiles to respire on insoluble electron acceptors or donors via bidirectional EET and invokes the possible role of G. halelectricus in iron cycling in barely studied haloalkaline environments. IMPORTANCE Bidirectional extracellular electron transfer (EET) appears to be a key microbial metabolic process in anoxic environments that are depleted in soluble electron donor and acceptor molecules. Though it is an ecologically important and applied microbial phenomenon, it has been reported with a few microorganisms, mostly from nonextreme environments. Moreover, direct electron transfer-based bidirectional EET is studied for very few microorganisms with electrodes in engineered systems and barely with the natural insoluble electron acceptor and donor molecules in anoxic conditions. This study advances the understanding of extremophilic microbial taxa capable of bidirectional EET and its role in barely investigated Fe cycling in highly saline-alkaline environments. It also offers research opportunities for understanding the membrane components involved in the bidirectional EET of G. halelectricus. The high rate of Fe3+ oxide reduction activity by G. halelectricus suggests its possible use as a biocatalyst in the anaerobic iron bioleaching process under neutral-alkaline pH conditions.

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Electroactive microorganisms are now understood to be abundant across nature, though many are categorized as "weak electricigens" not suitable for reasonable power generation. We report the use of weak electricigens from the natural environment for rapid, real-time water quality monitoring. Using a variety of pesticides as model chemical pollutants, the bioelectrochemical sensor was responsive within minutes at all concentrations tested (0.05-2 ppm) and could be repreatedly used long-term. Due to the prevalence of electroactive microorganisms in the natural environment, such sensors could work in tandem with conventional monitoring methods and may be useful for detecting emerging contaminants.

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Research on electroactive microorganisms (EAM) often focuses either on their physiology and the underlying mechanisms of extracellular electron transfer or on their application in microbial electrochemical technologies (MET). Thermodynamic understanding of energy conversions related to growth and activity of EAM has received only a little attention. In this study, we aimed to prove the hypothesized restricted energy harvest of EAM by determining biomass yields by monitoring growth of acetate-fed biofilms presumably enriched in Geobacter, using optical coherence tomography, at three anode potentials and four acetate concentrations. Experiments were concurrently simulated using a refined thermodynamic model for EAM. Neither clear correlations were observed between biomass yield and anode potential nor acetate concentration, albeit the statistical significances are limited, mainly due to the observed experimental variances. The experimental biomass yield based on acetate consumption (YX/ac = 37 ± 9 mgCODbiomass gCODac-1) was higher than estimated by modeling, indicating limitations of existing growth models to predict yields of EAM. In contrast, the modeled biomass yield based on catabolic energy harvest was higher than the biomass yield from experimental data (YX/cat = 25.9 ± 6.8 mgCODbiomass kJ-1), supporting restricted energy harvest of EAM and indicating a role of not considered energy sinks. This calls for an adjusted growth model for EAM, including, e.g., the microbial electrochemical Peltier heat to improve the understanding and modeling of their energy metabolism. Furthermore, the reported biomass yields are important parameters to design strategies for influencing the interactions between EAM and other microorganisms and allowing more realistic feasibility assessments of MET.

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The poor bacterial loading capacity and biocompatibility of the anode lead to weak electricity production performance of microbial fuel cells (MFCs). Inspired by kelp, we developed a double-layer hydrogel bioanode based on sodium alginate (SA). The inner hydrogel layer of encapsulated Fe3O4 and electroactive microorganisms (EAMs) was used as the bioelectrochemical catalytic layer. The outer hydrogel layer formed by cross-linking SA with polyvinyl alcohol (PVA) was used as the protective layer. The 3D porous structure of the inner hydrogel formed based on Fe3O4 facilitated the electroactive bacteria colonization and electron transfer, while the high structural toughness, salt-resistance and antibacterial properties of the outer highly cross-linked hydrogel served to protect the catalytic layer for stable electricity production. When high-salt waste leachate was used as the nutrient, the amazing open-circuit voltage (OCV) of 1.17 V and the operating voltage of 781 mV were brought by the double-layer hydrogel bioanode PVA@SA&Fe3O4/EAMs@SA.

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High energy consumption is impedimental for eliminating refractory organics in wastewater by current technologies. Herein, we develop an efficient self-purification process for actual non-biodegradable dyeing wastewater at pilot scale, using N-doped graphene-like (CN) complexed Cu-Al2O3 supported Al2O3 ceramics (HCLL-S8-M) fixed-bed reactor without additional input. About 36% chemical oxygen demand removal was achieved within 20 min empty bed retention time and maintained stability for almost one year. The HCLL-S8-M structure feature and its interface on microbial community structure, functions, and metabolic pathways were analyzed by density-functional theory calculation, X-ray photoelectron spectroscopy, multiomics analysis of metagenome, macrotranscriptome and macroproteome. On the surface of HCLL-S8-M, a strong microelectronic field (MEF) was formed by the electron-rich/poor area due to Cu-π interaction from the complexation between phenolic hydroxy of CN and Cu species, driving the electrons of the adsorbed dye pollutants to the microorganisms through extracellular polymeric substance and the direct transfer of extracellular electrons, causing their degradation into CO2 and intermediates, which was degraded partly via intracellular metabolism. The lower energy feeding for the microbiome produced less adenosine triphosphate, resulting in little sludge throughout reaction. The MEF from electronic polarization is greatly potential to develop low-energy wastewater treatment technology.

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Natural resources are in short supply, and the ecosystem is being damaged as a result of the overuse of fossil fuels. The creation of novel technology is greatly desired for investigating renewable and sustainable energy sources. Microorganisms have received a lot of interest recently for their potential to transform organic waste into sustainable energy and high-value goods. New exoelectrogens that can transmit electrons to electrodes and remove specific wastewater contaminants are expected to be studied. In this study, we examined three distinct samples (as determined by chemical oxygen demand and pH) that can be used as anolytes to generate power in single-chamber and double-chamber microbial fuel cells using graphite electrodes. Wastewater from poultry farms was studied as an exoelectrogenic anolyte for microbial fuel cell power generation. The study examined 10 different bacterial strains, numbered A1 through A10. Due to their highly anticipated capacity to metabolize organic/inorganic chemicals, the diverse range of microorganisms found in poultry wastewater inspired us to investigate the viability of generating electricity using microbial fuel cells. From the investigated bacterial strains, the highest voltage outputs were produced by strains A1 (Lysinibacillus sphaericus) and A2 (Bacillus cereus), respectively, at 402 mV and 350 mV. Among the 10 different bacterial strains, strain A6 generated the least amount of electricity, measuring 35.03 mV. Furthermore, a maximum power density of 16.16 1.02 mW/m2 was achieved by the microbial fuel cell using strain A1, significantly outperforming the microbial fuel cell using a sterile medium. The strain A2 showed significant current and power densities of 35 1.12 mA/m2 and 12.25 1.05 mW/m2, respectively. Moreover, in the two representative strains, chemical oxygen demand removal and Coulombic efficiency were noted. Samples from the effluent anode chamber were taken in order to gauge the effectiveness of chemical oxygen demand removal. Wastewater had an initial chemical oxygen demand content of 350 mg/L on average. Strains A1 and A2 decomposed 94.28% and 91.71%, respectively, of the organic substrate, according to the chemical oxygen demand removal efficiency values after 72 h. Strains A1 and A2 had electron donor oxidation efficiencies for 72 h of 54.1% and 60.67%, respectively. The Coulombic efficiency increased as the chemical oxygen demand decreased, indicating greater microbial electroactivity. With representative strains A1 and A2, Coulombic efficiencies of 10% and 3.5%, respectively, were obtained in the microbial fuel cell. The findings of this study greatly advance the field as a viable source of power technology for alternative energy in the future, which is important given the depletion of natural resources.

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Natural biomass is a promising candidate for manufacturing an efficient anode in the microbial electrochemical system (MES) for its abundance and low cost. However, the structure and performance of the electrode highly depend on the biomass species. A simple and sustainable method for creating a self-supporting electrode is proposed by freeze-drying and carbonizing a blend of cellulose, lignin, and hemicellulose. This strategy leads to a cork-like structure and improved mechanical strength of the lignocellulose carbon. A power density of 4780 ± 260 mW m-2 (CLX-800) was achieved, which was the highest record for unmodified lignocellulose-based anodes in the microbial fuel cells. The morphological as lamellar multilayer and rich in hydrophilic functional groups could facilitate the formation of thick electroactive biofilms and enrich Geobacter with the highest abundance of 92.3%. The CLX material is expected to be the ideal electrode for high performance and functionally controllability.

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Facing the increasingly severe energy shortage and environmental pollution, electrocatalytic processes using electroactive microorganisms provide a new alternative for achieving environmental-friendly production. Because of its unique respiratory mode and electron transfer ability, Shewanella oneidensis MR-1 has been widely used in the fields of microbial fuel cell, bioelectrosynthesis of value-added chemicals, metal waste treatment and environmental remediation system. The electrochemically active biofilm of S. oneidensis MR-1 is an excellent carrier for transferring the electrons of the electroactive microorganisms. The formation of electrochemically active biofilm is a dynamic and complex process, which is affected by many factors, such as electrode materials, culture conditions, strains and their metabolism. The electrochemically active biofilm plays a very important role in enhancing bacterial environmental stress resistance, improving nutrient uptake and electron transfer efficiency. This paper reviewed the formation process, influencing factors and applications of S. oneidensis MR-1 biofilm in bio-energy, bioremediation and biosensing, with the aim to facilitate and expand its further application.

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Various conductive materials and their dielectric counterparts were used to get deeper insights into contribution of direct interspecies electron transfer (DIET) in improving methanogenesis from highly concentrated volatile fatty acids (12.5 g/L). Potential CH4 yield, maximum CH4 production rate and lag phase were significantly (up to 1.4, 3.9 and 2.0 times, respectively) improved with addition of stainless-steel mesh (SM) and carbon felt (CF) compared to both control and dielectric counterparts (p < 0.05). kapp increased by 82% for SM and 63% for CF compared to control (p < 0.05). Short thick pili-like structures up to 150 nm in width were formed only in CF and SM biofilms, however, were more abundant for SM. Ureibacillus and Limnochordia specific for SM biofilms, and Coprothermobacter and Ca. Caldatribacterium for CF biofilms, were considered electrogenic. Promotion of DIET by conductive materials is governed by many factors, including specificity of electrogenic groups to material surface.

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Groundwater contamination has become increasingly prominent, therefore, the development of efficient remediation technology is crucial for improving groundwater quality. Bioremediation is cost-effective and environmentally friendly, while coexisting pollutant stress can affect microbial processes, and the heterogeneous character of groundwater medium can induce bioavailability limitations and electron donor/acceptor imbalances. Electroactive microorganisms (EAMs) are advantageous in contaminated groundwater because of their unique bidirectional electron transfer mechanism, which allows them to use solid electrodes as electron donors/acceptors. However, the relatively low-conductivity groundwater environment is unfavorable for electron transfer, which becomes a bottleneck problem that limits the remediation efficiency of EAMs. Therefore, this study reviews the recent advances and challenges of EAMs applied in the groundwater environment with complex coexisting ions, heterogeneity, and low conductivity and proposes corresponding future directions.

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Soil microbial fuel cells (MFCs) have been increasingly studied in recent years and have attracted significant attention as an environmentally sustainable bioelectrochemical technology. However, the poor conductivity of the soil matrix and the neglect of the cathodic function have limited its application. In this study, quartz sand and activated carbon were subjected to investigation on their influence on atrazine degradation. Atrazine was introduced in different layers (cathode, upper layer) to explore the cathodic effect on atrazine removal. The results revealed that activated carbon could reduce the internal resistance (693 Ω) and generate the highest power density (25.51 mW/m2) of the soil MFCs, and thus increase the removal efficiency (97.92%) of atrazine. The dynamic degradation profiles of atrazine were different for different adding layers. The cathode electrode acted as an electron donor could increase the distance of the effective influence of the soil MFCs' cathode from the middle to the cathode layer. The cathode (region) and the region close to the cathode could degrade atrazine with the atrazine removal efficiencies ranging from 60.67% to 92.79%, and the degradation ability of the cathode was stronger than that of other layers. The degradation effect followed the order: cathode > upper > lower > middle). Geobacter, Desulfobulbus, and Desulfuromonas belonging to the δ-Proteobacteria class were identified as the dominant electroactive microorganisms in the anode layer, while their relative abundances are quite low in the upper and cathode layers. Pseudomonas is an atrazine-degrading bacterium, but its relative abundance was only 0.13-0.51%. Thus, bioelectrochemistry rather than microbial degradation was the primary driving force.

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Microbial electrolysis cell-assisted thermophilic anaerobic digestion (MEC-TAD) is a promising method to improve anaerobic co-digestion efficiency; however, its application is restricted by high energy consumption. To improve the energy use efficiency of MEC-TAD, this study investigated the effect of different intermittent energization strategies on thermophilic co-digestion performance. Results revealed that an 18 h-ON/6h-OFF energization schedule resulted in the fastest electron transfer rate and the highest methane yield (364.3 mL/g VS). Mechanistic analysis revealed that 18 h-ON/6h-OFF resulted in the enrichment of electroactive microorganisms and increased abundance of enzyme-coding genes associated with energy metabolism (ntp, nuo, atp), electron transfer (pilA, nfrA2, ssuE), and the hydrogenotrophic methanogenic pathway. Finally, energy balance analysis revealed that 18 h-ON/6h-OFF had the highest net energy benefit (2.52 kJ) and energy conversion efficiency (110.76 %). Therefore, intermittent energization of MEC-TAD using an 18 h-ON/6h-OFF schedule can provide improved performance and more energy savings.

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Obtaining electroactive microbes capable of efficient extracellular electron transfer is a large undertaking for the scalability of bio-electrochemical systems. Inevitably, researchers need to pursue the co-modification of multiple genes rather than expecting that modification of a single gene would make a significant contribution to improving extracellular electron transfer rates. Base editing has enabled highly-efficient gene deactivation in model electroactive microbe Shewanella oneidensis MR-1. Since multiplexed application of base editing is still limited by its low throughput procedure, we thus here develop a rapid and efficient multiplex base editing system in S. oneidensis. Four approaches to express multiple gRNAs were assessed firstly, and transcription of each gRNA cassette into a monocistronic unit was validated as a more favorable option than transcription of multiple gRNAs into a polycistronic cluster. Then, a smart scheme was designed to deliver one-pot assembly of multiple gRNAs. 3, 5, and 8 genes were deactivated using this system with editing efficiency of 83.3%, 100% and 12.5%, respectively. To offer some nonrepetitive components as alternatives genetic parts of sgRNA cassette, different promoters, handles, and terminators were screened. This multiplex base editing tool was finally adopted to simultaneously deactivate eight genes that were identified as significantly downregulated targets in transcriptome analysis of riboflavin-overproducing strain and control strain. The maximum power density of the multiplex engineered strain HRF(8BE) in microbial fuel cells was 1108.1 mW/m2, which was 21.67 times higher than that of the wild-type strain. This highly efficient multiplexed base editing tool elevates our ability of genome manipulation and combinatorial engineering in Shewanella, and may provide valuable insights in fundamental and applied research of extracellular electron transfer.

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Facing the increasingly severe energy shortage and environmental pollution, electrocatalytic processes using electroactive microorganisms provide a new alternative for achieving environmental-friendly production. Because of its unique respiratory mode and electron transfer ability, Shewanella oneidensis MR-1 has been widely used in the fields of microbial fuel cell, bioelectrosynthesis of value-added chemicals, metal waste treatment and environmental remediation system. The electrochemically active biofilm of S. oneidensis MR-1 is an excellent carrier for transferring the electrons of the electroactive microorganisms. The formation of electrochemically active biofilm is a dynamic and complex process, which is affected by many factors, such as electrode materials, culture conditions, strains and their metabolism. The electrochemically active biofilm plays a very important role in enhancing bacterial environmental stress resistance, improving nutrient uptake and electron transfer efficiency. This paper reviewed the formation process, influencing factors and applications of S. oneidensis MR-1 biofilm in bio-energy, bioremediation and biosensing, with the aim to facilitate and expand its further application.

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The performance of peat biomethanation was investigated in bioelectrochemical anaerobic digestion at different applied voltages, and compared to conventional anaerobic digestion. The methane yield was stabilized at 16 mL/g peat in the conventional anaerobic digestion. However, in the bioelectrochemical anaerobic digestion, the methane yield was significantly increased to 264 mL/g peat at the applied voltage of 4 V, followed by 1 V, 2 V, 0.5 V and 0 V. The bioelectrochemical system could enrich more electroactive microorganisms on the electrode, as well as in the bulk solution, and further improve the direct interspecies electron transfer for methane production. The 16S rRNA analysis showed a significant increase in the abundance of specific microorganisms in the bulk solution, including Firmicutes phylum and Proteobacteria phylum, in addition to a gradual increase in acetoclastic methanogenesis with an increase in applied voltage. These results provide a solution to turn low-rank coal into a new alternative energy.

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