Extracellular Electron Transfer Process Research Articles

Semiconducting minerals participated extracellular electron transfer in red soil of Ningxia Plain, China

In recent years, exploring interactions between sunlight, semiconducting minerals and microorganisms in nature has attracted great attention. However, relatively little research has been conducted on the interaction between minerals and microorganisms in the plain areas of the Yellow River Basin under sunlight. In this study, the mineral composition and microbial community of red soil from the Ningxia Plain, China were analyzed. X–ray diffraction (XRD) results showed that red soil contained semiconducting minerals such as birnessite, hematite and goethite. 16S rRNA analysis found that electroactive microorganisms such as Proteobacteria, Firmicutes and Actinobacteriota were enriched in in–situ microbial community of red soil. Afterwards, electrochemical tests such as linear sweep voltammetry (LSV) and I–t curves, indicated that red soil exhibited good semiconducting properties under light irradiation. Finally, a dual chamber system and electrochemical techniques were used to explore the electron transfer relationship between red soil and microorganisms. The open circuit voltage (350 mV) and maximum power density (1242.72 mW/m2) of red soil cathode system were significantly improved compared to blank graphite electrode, indicating that red soil could serve as electron acceptors for microbial extracellular respiration. Red soil electrode in live bacteria exhibited the highest photocurrent density (0.29 μA/cm2) and the lowest charge transfer resistance (Rct) (223 Ω) under light conditions, indicating that red soil accelerated the rapid transfer of electrons, reduced polarization loss, and enhanced extracellular electron transfer (EET) process under light conditions. This study demonstrated that the participation of semiconducting minerals in microbial EET processes under sunlight in the Ningxia Plain, China.

Biology-Based Synthesis of Nickel Single Atoms on the Surface of Geobacter sulfurreducens as an Efficient Electrocatalyst for Alkaline Water Electrolysis.

Single-atom metal catalysts are promising electrocatalysts for water electrolysis. Nickel-based electrocatalysts have shown attractive application prospects for water electrolysis. However, synthesizing stable Ni single atoms using chemical and physical approaches remains a practical challenge. Here, a facile and precise method for synthesizing stable nickel single atoms on the surface of Geobacter sulfurreducens using a microbial-mediated extracellular electron transfer (EET) process is demonstrated. It is shown that G. sulfurreducens can effectively anchor nickel single atoms on their surface. X-ray absorption near-edge structure and Fourier-transformed extended X-ray absorption fine structure spectroscopy confirm that the nickel single atom is coordinated to nitrogen in the cytochromes. The as-synthesized nickel single atoms on G. sulfurreducens exhibit excellent bifunctional catalytic properties for alkaline water electrolysis with low overpotential (η) to achieve current density (10mAcm-2) for both hydrogen evolution reactions (η = 80mV) and oxygen evolution reaction (η = 330mV) with minimal catalyst loading of 0.0015mgNicm-2. The nickel single-atom catalyst shows long-term stability at a constant electrode potential. This synthesis method based on the EET capability of electroactive bacteria provides a simple and scalable approach for producing low-cost and highly efficient nonnoble transition metal single-atom catalysts for practical applications.

Facilitating Intracellular Electron Bifurcation by Mediating Flavin-Based Extracellular and Transmembrane Electron Transfer: A Novel Role of Pyrogenic Carbon in Dark Fermentation for Hydrogen Production.

Pyrogenic carbon is considered an enhancer to H2-yielding dark fermentation (DF), but little is known about how it regulates extracellular electron transfer (EET) and influences transmembrane respiratory chains and intracellular metabolisms. This study addressed these knowledge gaps and demonstrated that wood waste pyrogenic carbon (biochar) could significantly improve the DF performance; e.g., addition of pyrogenic carbon produced by pyrolysis at 800 °C (PC800) increased H2 yield by 369.7%. Biochemical quantification, electrochemical analysis, and electron respiratory chain inhibition tests revealed that PC800 promoted the extracellular flavin-based electron transfer process and further activated the acceleration of the transmembrane electron transfer. Comparative metagenome/metatranscriptome analyses indicated that the flavin-containing Rnf complex was the potential transmembrane respiratory enzyme associated with PC800-mediated EET. Based on NADH/NAD+ circulation, the promoted Rnf complex could stimulate the functions of the electron bifurcating Etf/Bcd complex and startup of glycolysis. The promoted Etf/Bcd could further contribute to balance the NADH/NAD+ level for glycolytic reactions and meanwhile provide reduced ferredoxin for group A1 [FeFe]-hydrogenases. This proton-energy-linked mechanism could achieve coupling production of ATP and H2. This study verified the important roles of pyrogenic carbon in mediating EET and transmembrane/intracellular pathways and revealed the crucial roles of electron bifurcation in DF for hydrogen production.

Regulation of extracellular electron transfer by sustained existence of Fe²⁺/Fe³⁺ redox couples on iron oxide-functionalized woody biochar anode surfaces in bioelectrochemical systems

Sluggish extracellular electron transfer between the exoelectrogens and anode interface limits the application of bioelectrochemical systems (BECs) for energy generation. In this study, an innovative anode coated with oxidized biochar and co-functionalized with iron nanoparticles (FBC/Fe2O3) was developed to enhance microbial reactions through tuned physicochemical and electrical properties. The designed anode features a highly porous, heterogeneous structure with a large accessible surface area of 10.141 m²/g, promoting better habitation and metabolism of exoelectrogens. The synergistic effect of iron nanoparticles and oxidized functional groups increased the hydrophilicity of the anode surface, augmenting its affinity for bacterial outer membrane c-cysts and facilitating microbial adhesion at a density of 3.106 × 10⁹ CFU/cm². The iron moieties on the FBC/Fe2O3 electrode surface act as electron mediators, utilizing Fe²⁺/Fe³⁺ redox couples, as evidenced by X-ray photoelectron spectroscopy (XPS) data. This enhancement improved extracellular electron transfer (EET) between bacterial cells and the anode surface, resulting in a faster and bifurcated EET process through both direct transfer via outer membrane c-cysts and mediated transfer via the redox couples of Fe moieties. The assembled double-chamber microbial fuel cell (DC-MFC) with the FBC/Fe2O3 anode achieved a maximum power density of 528.75 mW/m² and a chemical oxygen demand (COD) removal efficiency of 47.5 % within 24 h of operation.

3D Bioprinting of Engineered Living Materials with Extracellular Electron Transfer Capability for Water Purification.

Attention is widely drawn to the extracellular electron transfer (EET) process of electroactive bacteria (EAB) for water purification, but its efficacy is often hindered in complex environmental matrices. In this study, the engineered living materials with EET capability (e-ELMs) were for the first time created with customized geometric configurations for pollutant removal using three-dimensional (3D) bioprinting platform. By combining EAB and tailored viscoelastic matrix, a biocompatible and tunable electroactive bioink for 3D bioprinting was initially developed with tuned rheological properties, enabling meticulous manipulation of microbial spatial arrangement and density. e-ELMs with different spatial microstructures were then designed and constructed by adjusting the filament diameter and orientation during the 3D printing process. Simulations of diffusion and fluid dynamics collectively showcase internal mass transfer rates and EET efficiency of e-ELMs with different spatial microstructures, contributing to the outstanding decontamination performances. Our research propels 3D bioprinting technology into the environmental realm, enabling the creation of intricately designed e-ELMs and providing promising routes to address the emerging water pollution concerns.

Insights into the role of electrochemical stimulation on sulfur-driven biodegradation of antibiotics in wastewater treatment

The presence of antibiotics in wastewater poses significant threat to our ecosystems and health. Traditional biological wastewater treatment technologies have several limitations in treating antibiotic-contaminated wastewaters, such as low removal efficiency and poor process resilience. Here, a novel electrochemical-coupled sulfur-mediated biological system was developed for treating wastewater co-contaminated with several antibiotics (e.g., ciprofloxacin (CIP), sulfamethoxazole (SMX), chloramphenicol (CAP)). Superior removal of CIP, SMX, and CAP with efficiencies ranging from 40.6 ± 2.6 % to 98.4 ± 1.6 % was achieved at high concentrations of 1000 μg/L in the electrochemical-coupled sulfur-mediated biological system, whereas the efficiencies ranged from 30.4 ± 2.3 % to 98.2 ± 1.4 % in the control system (without electrochemical stimulation). The biodegradation rates of CIP, SMX, and CAP increased by 1.5∼1.9-folds under electrochemical stimulation compared to the control. The insights into the role of electrochemical stimulation for multiple antibiotics biodegradation enhancement was elucidated through a combination of metagenomic and electrochemical analyses. Results showed that sustained electrochemical stimulation significantly enriched the sulfate-reducing and electroactive bacteria (e.g., Desulfobulbus, Longilinea, and Lentimicrobiumin on biocathode and Geobactor on bioanode), and boosted the secretion of electron transport mediators (e.g., cytochrome c and extracellular polymeric substances), which facilitated the microbial extracellular electron transfer processes and subsequent antibiotics removal in the sulfur-mediated biological system. Furthermore, under electrochemical stimulation, functional genes associated with sulfur and carbon metabolism and electron transfer were more abundant, and the microbial metabolic processes were enhanced, contributing to antibiotics biodegradation. Our study for the first time demonstrated that the synergistic effects of electrochemical-coupled sulfur-mediated biological system was capable of overcoming the limitations of conventional biological treatment processes. This study shed light on the mechanism of enhanced antibiotics biodegradation via electrochemical stimulation, which could be employed in sulfur-mediated bioprocess for treating antibiotic-contaminated wastewaters.

The effect of dissolved oxygen and Shewanella algae on the corrosion mechanism of titanium in a simulated marine environment

The influence of dissolved oxygen (DO) and Shewanella algae on the corrosion behavior of pure titanium were systematically studied in this work. The formation of S. algae biofilms on titanium surface was facilitated by the anaerobic environment, which accelerated titanium corrosion. Upon the breakdown of passive film, S. algae acquired electrons from the titanium base substrate through extracellular electron transfer (EET) processes. Various EET-related genes were overexpressed by the low DO condition, leading to enhanced EET kinetics. High concentration DO increased TiO2 content and the thickness of passive film, which enhanced the protective effect and mitigated microbiologically influenced corrosion.

Novel Insights on Extracellular Electron Transfer Networks in the Desulfovibrionaceae Family: Unveiling the Potential Significance of Horizontal Gene Transfer.

Some sulfate-reducing bacteria (SRB), mainly belonging to the Desulfovibrionaceae family, have evolved the capability to conserve energy through microbial extracellular electron transfer (EET), suggesting that this process may be more widespread than previously believed. While previous evidence has shown that mobile genetic elements drive the plasticity and evolution of SRB and iron-reducing bacteria (FeRB), few have investigated the shared molecular mechanisms related to EET. To address this, we analyzed the prevalence and abundance of EET elements and how they contributed to their differentiation among 42 members of the Desulfovibrionaceae family and 23 and 59 members of Geobacteraceae and Shewanellaceae, respectively. Proteins involved in EET, such as the cytochromes PpcA and CymA, the outer membrane protein OmpJ, and the iron-sulfur cluster-binding CbcT, exhibited widespread distribution within Desulfovibrionaceae. Some of these showed modular diversification. Additional evidence revealed that horizontal gene transfer was involved in the acquiring and losing of critical genes, increasing the diversification and plasticity between the three families. The results suggest that specific EET genes were widely disseminated through horizontal transfer, where some changes reflected environmental adaptations. These findings enhance our comprehension of the evolution and distribution of proteins involved in EET processes, shedding light on their role in iron and sulfur biogeochemical cycling.

Enhancing anaerobic ammonium oxidation via polymeric ferric sulfate (PFS)-Mediated dual-electron Acceptor: A solution to nitrite deficiency

In addressing the issue of the unstable supply of electron acceptors (nitrite) during Anammox, researchers have often overlooked that the electron transfer process in anammox bacteria (AnAOB) is not restricted to the intracellular, extracellular electron transfer can also serve as an alternative pathway. This study utilized Fe3+ as an adjunctive electron acceptor, coupling Anammox with ferric iron reduction (Feammox) by introducing polymeric ferric sulfate (PFS) in the Anammox system. This approach achieved effective nitrogen removal even when nitrite was deficient. At a NO2–-N to NH4+-N ratio of 0.5, the Feammox-Anammox combined system maintained stable nitrogen removal under the dual electron acceptor, with total nitrogen and NH4+-N removal efficiency of 85 % and 91 %, respectively. The contribution of Feammox and Anammox to NH4+-N removal was 45 % and 55 %, respectively. TEM-EDS confirmed the presence of iron elements across the periphery, periplasm, and cytoplasm of AnAOB, suggesting the involvement of PFS in the metabolic and electron transfer processes. Notably, the introduction of Fe3+ and the limitation of nitrite availability resulted in increased relative abundances of FeOB (Thermomonas) and FeRB (Ignavibacterium) to 30 % and 3.6 %, respectively, indicating that the electron transfer process with Fe3+ as the electron acceptor was continuously intensified, as well as a cycle of Fe3+ / Fe2+ was established. Due to unique nitrogen metabolism mode and high Fe3+ tolerance, the abundance of Candidatus_Brocadia was enriched up to 7.2 %, which alternately dominated the Anammox process with Candidatus_Kuenenia, providing favorable conditions for AnAOB to carry out the extracellular electron transfer process. These findings provide a novel strategy for improving the stability and efficiency of the Anammox process under deficient nitrite supply.

Riboflavin derivatives as a novel electron transfer mediator for enhancing Cr(VI) removal by Shewanella oneidensis MR-1

Bioremediation has garnered considerable interest due to its advantages of economy and no secondary pollution. However, the direct electron transfer rate between microorganisms and Cr(VI) is low. The bioreduction of Cr(VI) by Shewanella oneidensis MR-1 (S. oneidensis) in the presence of formylmethylflavin (FMF) was conducted to understand how FMF mediated the extracellular electron transfer process to improve Cr(VI) removal. In this study, FMF was firstly synthesized by modifing RF to increase its solubility and redox activity. The findings indicate that S. oneidensis/FMF exhibited better Cr(VI) removal performance compared to that of S. oneidensis. Under optimum conditions, 40 mg/L Cr(VI) could be completely removed by S. oneidensis/FMF within 120 h, while only 48.6% of Cr(VI) was removed by S. oneidensis, and the first order rate constant (k) for the Cr(VI) elimination by S. oneidensis/FMF (0.033 h−1) was about 4.1-fold greater than that of S. oneidensis (0.008 h−1). Moreover, the removal of Cr(VI) by S. oneidensis/FMF were dominated by reduction, and supplemented by adsorption and complexation. FMF enhance the extracellular electron transfer rate (EETR) was confirmed by electrochemical cyclic voltammetry (CV) and linear scanning voltammetry (LSV) experiments. This study emphasizes the potential important role of FMF in environmental bioremediation.

Boosting bioelectricity generation in bioelectrochemical systems with nitrogen-doped three-dimensional graphene aerogel anode

Bioelectricity generation by electrochemically active bacteria has become particularly appealing due to its vast potential in energy production, pollution treatment, and biosynthesis. However, developing high-performance anodes for bioelectricity generation remains a significant challenge. In this study, a highly efficient three-dimensional nitrogen-doped macroporous graphene aerogel anode with a nitrogen content of approximately 4.38 ± 0.50 at% was fabricated using hydrothermal method. The anode was successfully implemented in bioelectrochemical systems inoculated with Shewanella oneidensis MR-1, resulting in a significantly higher anodic current density (1.0 A/m2) compared to the control one. This enhancement was attributed to the greater biocapacity and improved extracellular electron transfer efficiency of the anode. Additionally, the N-doped aerogel anode demonstrated excellent performance in mixed-culture inoculated bioelectrochemical systems, achieving a high power density of 4.2 ± 0.2 W/m², one of the highest reported for three-dimensional carbon-based bioelectrochemical systems to date. Such improvements are likely due to the good biocompatibility of the N-doped aerogel anode, increased extracellular electron transfer efficiency at the bacteria/anode interface, and selectively enrichment of electroactive Geobacter soli within the NGA anode. Furthermore, based on gene-level Picrust2 prediction results, N-doping significantly upregulated the conductive pili-related genes of Geobacter in the three-dimensional anode, increasing the physical connection channels of bacteria, and thus strengthening the extracellular electron transfer process in Geobacter.

Promotion of direct electron transfer between Shewanella putrefaciens CN32 and carbon fiber electrodes via in situ growth of α-Fe2O3 nanoarray

The iron transport system plays a crucial role in the extracellular electron transfer process of Shewanella sp. In this study, we fabricated a vertically oriented α-Fe2O3 nanoarray on carbon cloth to enhance interfacial electron transfer in Shewanella putrefaciens CN32 microbial fuel cells. The incorporation of the α-Fe2O3 nanoarray not only resulted in a slight increase in flavin content but also significantly enhanced biofilm loading, leading to an eight-fold higher maximum power density compared to plain carbon cloth. Through expression level analyses of electron transfer-related genes in the outer membrane and core genes in the iron transport system, we propose that the α-Fe2O3 nanoarray can serve as an electron mediator, facilitating direct electron transfer between the bacteria and electrodes. This finding provides important insights into the potential application of iron-containing oxide electrodes in the design of microbial fuel cells and other bioelectrochemical systems, highlighting the role of α-Fe2O3 in promoting direct electron transfer.

Boosting power output in microbial fuel cell with sulfur-doped MXene/polypyrrole hydrogel anode for enhanced extracellular electron transfer process

Boosting power output in microbial fuel cell with sulfur-doped MXene/polypyrrole hydrogel anode for enhanced extracellular electron transfer process

Significance of a minor pilin PilV in biofilm cohesion of Geobacter sulfurreducens

Bacterial adhesion plays a vital role in forming and shaping the structure of electroactive biofilms that are essential for the performance of bioelectrochemical systems (BESs). Type IV pili are known to mediate cell adhesion in many Gram-negative bacteria, but the mechanism of pili-mediated cell adhesion of Geobacter species on anode surface remains unclear. Herein, a minor pilin PilV2 was found to be essential for cell adhesion ability of Geobacter sulfurreducens since the lack of pilV2 gene depressed the cell adhesion capability by 81.2% in microplate and the anodic biofilm density by 23.1 % at −0.1 V and 37.7 % at −0.3 V in BESs. The less cohesiveness of mutant biofilms increased the charge transfer resistance and biofilm resistance, which correspondingly lowered current generation of the pilV2-deficient strain by up to 63.2 % compared with that of the wild-type strain in BESs. The deletion of pilV2 posed an insignificant effect on the production of extracellular polysaccharides, pili, extracellular cytochromes and electron shuttles that are involved in biofilm formation or extracellular electron transfer (EET) process. This study demonstrated the significance of pilV2 gene in cell adhesion and biofilm formation of G. sulfurreducens, as well as the importance of pili-mediated adhesion for EET of electroactive biofilm.

Hierarchical modulation of extracellular electron transfer processes in microbial fuel cell anodes for enhanced power output through improved Geobacter adhesion

Although microbial fuel cell (MFC) technology is nearing practical realization, the challenge of attaining higher power output continues to impede its widespread implementation. In this study, we successfully developed free-standing three-dimensional high-performance anode materials for MFCs using a straightforward "one-step" process based on garlic. The resulting anode not only promotes the accumulation of biomass and the enrichment of exoelectrogen Geobacter but also facilitates interfacial extracellular electron transfer. This led to rapid start-up within just three days, a robust power density of 12.37 W/m3, and an impressive 24.04 % coulombic efficiency in mixed-culture MFC reactors. Our investigation into the underlying mechanisms of exocellular electron transfer revealed improvements in both direct electron transfer and mediated electron transfer processes. Notably, the fabrication of carbonized garlic MFC anodes proves to be technically competitive and economically relevant. It not only paves the way for the development of high-performance MFCs but also provides valuable insights for the rational design of materials related to microbial electrochemical technologies.

An integrated constructed wetland-Microbial fuel cell system with sewage sludge-biochar to enhance treatment and energy recovery efficiencies

Constructed wetlands and microbial fuel cells (CW-MFCs) are a promising green and sustainable wastewater treatment technology. However, the power production performance of CW-MFCs has long been limited by low-performance cathodes. In this study, we designed two CW-MFC systems using cheap and readily available sewage sludge biochar (SSB) as the substrate for the experimental units. The integration of SSB significantly facilitated the removal of chemical oxygen demand, ammonium nitrogen, total nitrogen, and total phosphorus. Moreover, SSB enhanced the electrochemical performance of the air cathode, resulting in higher power density (44.64 mW/m2) and lower internal resistance (900.00 Ω) compared to gravel-integrated CW-MFC (0.88 mW/m2, 2057.14 Ω). In addition, a typical electroactive bacterium---Geobacter was enriched in the SSB-integrated CW-MFC system. The iron oxides and graphic nitrogen of the SSB favor the growth of Geobacter, and the redox-active groups on the surface of SSB accelerate the extracellular electron transfer process, contributing to the enhancement of treatment performance. Meanwhile, the metal and nitrogen doping improves the oxygen reduction reaction activity and conductivity of SSB, enhancing the cathode area of CW-MFC and improving the energy recovery. Overall, these results demonstrated that SSB-integrated CW-MFC can be a competitive technology for effective wastewater treatment and energy recovery.

Iron-doped cross-linked puffed carbon as capacitive anode for the enhancement of bioelectron storage capacity in microbial fuel cells

Microbial fuel cell (MFC) can utilize the metabolism of electroactive microorganisms to remove organic pollutants from wastewater, and the application of capacitive anodes can enhance the power generation capacity of MFC and the energy storage capacity of bioelectronics. This research aimed to create a capacitive anode with a large specific surface area and excellent electrical conductivity. To achieve this aim, iron-doped cross-linked puffed carbon (CPC–Fe) was prepared for the modification of carbon cloth (CC) substrates by a simple one-step expansion carbonization method using FeCl3·6H2O and maltose as raw materials. The surface structure characterization confirms the successful incorporation of Fe atoms, with Fe3O4 accounting for 79.61%. Fe3O4 exhibits excellent specific capacitance characteristics and can also serve as a good conductor for the transmission or reception of electrons generated by electroactive bacteria. Furthermore, the mesoporous volume of CPC-Fe significantly increases, which contributes to the enhancement of the specific capacitance of material and the improvement of charge storage capacity. Based on these advantages, the maximum power density of MFC equipped with CPC-Fe/CC anode reaches 1850.50 mW m−2. The CPC-Fe/CC bioanode has a cumulative total charge of 1076.02 C m−2, showing good capacitive behaviour and exhibiting a significant enrichment effect on Geobacter species. This study demonstrates that the regulation of extracellular electron transfer (EET) process based on bioelectronic reception steps can not only enhance the power generation performance of MFC, but also effectively improve the energy storage capacity of MFC, which offers significant knowledge for the development of electrode design in other bioelectrochemical systems.

Bioelectricity facilitates carbon dioxide fixation by Alcaligenes faecalis ZS-1 in a biocathodic microbial fuel cell (MFC)

The CO2 fixation mechanism by Alcaligenes faecalis ZS-1 in a biocathode microbial fuel cell (MFC) was investigated. The closed-circuit MFC (CM) exhibited a significantly higher CO2 fixation rate (10.7%) compared to the open-circuit MFC (OC) (2.0%), indicating that bioelectricity enhances CO2 capture efficiency. During the inward extracellular electron transfer (EET) process, riboflavin concentration increased in the supernatant while cytochrome levels decreased. Genome sequencing revealed diverse metabolic pathways for CO2 fixation in strain ZS-1, with potential dominance of rTCA and C4 pathways under electrotrophic conditions as evidenced by significant upregulation of the ppc gene. Differential metabolite analysis using LC-MS demonstrated that CM promoted upregulation of various lipid metabolites. These findings collectively highlight that ZS-1 simultaneously generated electricity and fixed CO2 and that the ppc associated with bioelectricity played a critical role in CO2 capture. In conclusion, bioelectricity resulted in a significant enhancement in the efficiency of CO2 fixation and lipid production.

Retraction for Yadav et al., "The bidirectional extracellular electron transfer process aids iron cycling by Geoalkalibacter halelectricus in a highly saline-alkaline condition".

In our paper, we reported the bidirectional extracellular electron transfer capability of Geoalkalibacter halelectricus based on biochemical (i.e., with insoluble Fe-oxide and Fe(0)) and bioelectrochemical (i.e., with electrodes) experimental approaches. We noticed some issues and limitations of the methods and techniques that were used to analyze Fe species, particularly the reduced Fe ions in insoluble and precipitated Fe(II) minerals that led to incorrect interpretation of the results, specifically the reduction of Fe(III) to Fe(0) in this study. We were made aware of thermodynamic constraints that would make the biological reduction of Fe(III) to Fe(0) implausible. Hence, the conclusion about microbial Fe(III)-oxide reduction to Fe(0) is invalid. We also noticed errors in estimating the protein-based iron reduction/oxidation rates and faradic efficiencies, besides the limitations of the methods used for Fe analysis. For these reasons, we retract this article and sincerely apologize for the inconvenience it may have caused to the readers.

Response of biocurrent conduction to soil microenvironment

The biocurrent generated by soil extracellular electron transfer (EET) partly drives biogeochemical cycles and controls soil quality. However, it is unclear how the soil abiotic and biotic conditions affect the biocurrent conduction. In this study, the response relationship of soil microenvironment and in-situ biocurrent was studied. The results showed that red soil exhibited the optimal electron transfer efficiency, as evidenced by the maximum current density and accumulated charge output, with increments of 56–93 % and 80–2800 %, respectively, compared with the other five types of soils. Soil physicochemical properties were the most important factor on the biocurrent generation, and further the quantity and bioavailability of dissolved organic matter, NH4+-N content, and lower pH were predictive indicators for the exoelectrogenic processes of soils. In addition, the high soil biocurrent was likely determined by a complex synergistic network of the transformation of carbon and nitrogen, electroactive bacteria involving the functions of cell wall/membrane and cytochrome enzyme metabolism and transport related EET process. Overall, we provide an insight into the relationship among soil biocurrent conduction, physicochemical properties, bacteria community and metabolic function, and a support for bioelectrochemical technology application.