Xiao, X. et al. CAS The mixture of the bacterial solution (5mL), Tris-HCl buffer (1mL), Na2SO3 (0.36%, 0.5mL) and TTC (0.4%, 1mL) was kept in a rotary incubator at 37C for 30min, followed by addition of methanal (37%, 1mL). Ye Sun, Yujie Feng, Flemming Besenbacher, Feng Zhao or Miao Yu. Thank you for visiting nature.com. eCollection 2021. 6f). For electricity generation in the bioelectrical systems of bacteria, the cellular adhesion and the cell surface electronegativity play a vital role in determining the junctions of the cells with the anode and the direct extracellular electron transfer rate. Copyright American Society for Microbiology. *p0.05, **p0.01, and ***p0.001 were considered to be statistically significant. To further explore whether the CDs were attached on the cell surface specifically or also being able to distribute inside the cells, cryo-electron tomography was employed. (2010), ArcS, the cognate sensor kinase in an atypical Arc system of, Gao H., Wang X., Yang Z.K., Palzkill T. and Zhou J. In this study, a novel green biosynthesis route for producing Cu-NPs using the metal-reducing bacterium, Shewanella oneidensis is demonstrated. Logan, B. E. & Rabaey, K. Conversion of wastes into bioelectricity and chemicals by using microbial electrochemical technologies. These emerging technologies would further increase the value of EAB, while further efforts are necessary for bringing these technologies close to practical applications. EG has recently been proposed as a technology to control gene expression using electrodes in BESs [15,59,60], and EG using EAB is based on the finding that the Arc global regulatory system regulates the expression of diverse catabolic genes in MR-1 by sensing electrode potentials [27] (Figure 4). (2007) Ecology and biotechnology of the genus Shewanella. 136, 277287 (1996). Article The increased intracellular electron generation, higher ATP level, more vigorous extracellular secretion, and enhanced transmembrane and extracellular electron transfer all emphasize the positive and effective contribution of the CDs to the elevated metabolism of S. oneidensis MR-1. Unlike the control group that only showed weak redox peaks, two pairs of pronounced redox peaks were observed in MEC-CD: (1) the 1st pair, i.e., the reduction peak current of 103.5A at 0.43V and oxidation peak current of 157.8A at 0.20V, where the enhancement is correlated with the mediated extracellular electron transfer increased by additional riboflavin excretion23 upon the CDs addition; and (2) the 2nd pair, i.e., the reduction peak current of 4.8A at 0.24V and oxidation peak current of 61.9 A at 0.02V, where the enhancement is associated with the improved transmembrane electron transfer29,30. This proteobacterium was first isolated from Lake Oneida, NY in 1988, hence its name.[1]. a TEM image and size distribution. S. oneidensis MR-1 fed by the CDs (100gmL1) were added into the electrochemical reactor to test the MECs performance. (3) The CDs can even restore charge transfer capability of OmcA/MtrC mutant that suffers from compromised electron-transfer path. Hazard. Carlin Geor Malar, Seenuvasan Muthulingam, Rakesh Sankar, Nature Communications In addition, studies have been focused on its ability as an electrochemically active bacterium that is capable of discharging electrons to and receiving electrons from electrodes in bioelectrochemical systems (BESs) for balancing intracellular redox states. The protein concentration was measured by BCA assay kit46. 6b). Superscript A indicates enzymes whose gene expression is regulated by the Arc system, superscript C indicates enzymes whose gene expression is regulated by the cAMP/CRP system, while superscript F indicates enzymes whose gene expression is regulated by the Fnr system. Adv. Would you like email updates of new search results? 2011 Feb 2;6(2):e16649. Having made its first sale in 2014, Cambrian now has industrial-scale projects at more than 10 different wineries and breweries. Mar. . 6e). Mater. In addition, fluorescence imaging was employed to demonstrate CDs-cytochromes interaction intuitively. Soc. (A) Heme-stained SDS-PAGE analysis of loosely attached proteins from cells grown anaerobically on fumarate. (2009), Electrochemical measurement of electron transfer kinetics by, Carmona-Martinez A.A., Harnisch F., Fitzgerald L.A., Biffinger J.C., Ringeisen B.R. Soc. Silver nanoparticles are considered to be a new generation of antimicrobial as they exhibit biocidal activity towards a broad range of bacteria, and are gaining importance with the increasing resistance in antibiotics by pathogenic bacteria. The TEM images of pristine S. oneidensis MR-1 cells showed a relatively smooth profile and uniform contrast (Fig. 2c) showed evident crystalline structure, with a lattice distance of 0.21nm corresponding to the in-plane spacing of graphene (100)17,18. Genomic and genetic studies have assembled catabolic and electron-transport pathways in S. oneidensis MR-1 (Figure 1), and enzymes involved in these pathways are detailed elsewhere [17,18]. Understanding of electrochemical characteristics is essential for utilizing and controlling EAB in BESs applied to biotechnology, since cell voltage is directly linked to power output from MFC, and charged voltage is related to electricity expense in MEC. Adv. Nitrogen-doped carbon dots with excitation-independent long-wavelength emission produced by a room-temperature reaction. In addition, since some BESs, e.g. Rev. Rev. An interesting feature in sugar catabolism in MR-1 is that this bacterium utilizes d-lactate as a temporal electron sink in the process to produce acetate as the catabolic end product under electron acceptor-limited conditions [20] (Figure 1). Researchers expect that EG will be used in combination with EF for developing efficient bioproduction processes; e.g., foreign genes that encode bioproduction pathways are expressed under the control of electrode potentials to synchronize gene expression (EG) and energy supply (EF) [15]. 73, 70037012 (2007). Clipboard, Search History, and several other advanced features are temporarily unavailable. Li, N. et al. Sci. EB, namely, electrochemical control of biological functions, is a new area of research with tremendous future applications. Okamoto, A. et al. Anodic and cathodic current generation by S. oneidensis MR-1 in BESs, Figure 3. Since such genes encode pivotal enzymes in the bioconversion and electron-transfer pathways, deep understanding of the regulatory cascades is necessary for developing efficient EB processes. Biotechnol. In addition to the cAMP/CRP and Arc systems, other global regulators, including homologs of ferric uptake regulator (Fur) and fumarate nitrate reduction regulator (Fnr) in E. coli, are also involved in the transcriptional regulation of catabolic genes [28,29]. 52, 39533957 (2013). sharing sensitive information, make sure youre on a federal Natl Acad. An important example is the synthesis of silver nanoparticle by S. oneidensis, where its antibacterial activity can be influenced by the expression of outer membrane c-type cytochromes. Wu, R. et al. Nevertheless, not only the pristine cells but also the CDs-fed S. oneidensis MR-1 and CDs-fed OmcA/MtrC mutant showed no fluorescence signal (Supplementary Fig. 2022 Jun 1;13:852527. doi: 10.3389/fmicb.2022.852527. A. et al. 2019 Oct 1;142:111571. doi: 10.1016/j.bios.2019.111571. Soc. MR-1 is however unable to grow on glucose, an abundantly present sugar in the natural environment and a widely used feedstock in biotechnology processes [17]. 2020 Feb 24;15(2):e0229359. wrote the manuscript; M.Y., F.Z., Y.S., Y.F., and F.B. The CDs possessed pronounced PL emission, with in vitro imaging potential (Supplementary Figs. Three-electrode BES designed for EG, Figure 4. Appl. These authors contributed equally: Chenhui Yang, Hsn Aslan, Peng Zhang. The genus Shewanella comprises over 70 species of heterotrophic bacteria with versatile respiratory capacities. Ross DE, Flynn JM, Baron DB, Gralnick JA, Bond DR. PLoS One. Shi L, Squier TC, Zachara JM, Fredrickson JK. Proc. Dual-chamber MFCs (100mL) with the electrodes connected via a 1k external resistor was used to record voltage output of every 10min using a data acquisition system (USB2801, ATD Co., China). Development of targeting lonidamine liposomes that circumvent drug-resistant cancer by acting on mitochondrial signaling pathways. DSFO+ could completely replace natural current-conducting proteins, boosting the power that the microbe generated. 2. C-type cytochromes are a family of iron-containing proteins (e.g., MtrA, MtrB, MtrC, OmcA, and CymA) in S. oneidensis MR-1, which work synergistically to enable electron transfer throughout the cell membrane31. Biol. Studies to accelerate production rates in EF are therefore necessary, and these studies will be conducted based on achievements of studies to gain deeper understanding of metabolic and electrochemical characteristics of MR-1. Google Scholar. The enhancement efficacy is found to be strongly dependent on the surface charge of the CDs. A comprehensive review of the importance of thermal activation in the production of carbon dots and the potential for their use in the bioenergy industry, http://creativecommons.org/licenses/by/4.0/. This combined view of results from the biomass and flow-cell experiments suggests the incremental cellular electronegativity by the CDs, consistent with the densely aligned CDs in the cell membrane revealed in the TEM images (Fig. e, f AFM images of dehydrated pristine and CDs-fed S. oneidensis MR-1, depicting altered roughness in the presence of CDs. HHS Vulnerability Disclosure, Help Biofilms formed by S. oneidensis, Figure 3. Consistently, the ATP level of the CDs-fed cells increased to 136% as that of the pristine cells (0.190.02 nmol/mg protein vs. 0.140.01 nmol/mg protein), while the influence of the CDs alone (without the cells) on the ATP measurement was negligible. 2022 Aug 21;27(16):5330. doi: 10.3390/molecules27165330. Sci. This engineered strain however does not grow on glucose under fermentative conditions (in the absence of electron acceptors), even though it has all enzymes necessary for completing the fermentative pathway from glucose to lactate [18]. Clipboard, Search History, and several other advanced features are temporarily unavailable. Chem. Nanostructured material-based biofuel cells: recent advances and future prospects. Ed. A dose of ionizing radiation that results in 17% survival of S. oneidensis is 20- and 200-fold lower than doses that give the same survival rate in E. coli and Deinococcusradiodurans, respectively. The cell population with the CDs were found to be evidently higher (32.943.30gcm2 vs. 16.924.35gcm2). doi: 10.1371/journal.pone.0149812. EDTA control and extensive cation presence/absence tests show that Ca(II), Mn(II), Cu(II) and Zn(II) are all essential in this process, probably functioning as a part of a coenzyme or prosthetic group. Careers, Monitoring Editor: Diethard Mattanovich and Pablo Ivan Nikel. To our knowledge, this is the first report of E. coli ccm genes being functional in another organism. Refer to [42] for experimental details. Chong GW, Pirbadian S, Zhao Y, Zacharoff LA, Pinaud F, El-Naggar MY. This capacity is supported by the presence of various terminal reductases and overlapping electron-transport pathways, facilitating them to survive in redox-stratified environments, where available electron acceptors repeatedly change in response to fluctuations in environmental conditions [5]. USA 105, 39683973 (2008). A S. oneidensis fur deletion mutant was defective in growth under iron-abundant or acidic environment. After 6h incubation with the CDs, extracellular secretion layer was abundantly growing into high-coverage continuous domains (Fig. In addition, the above-mentioned features support the idea that the EET pathway of MR-1 is applicable to EF for producing either oxidized or reduced products. Carbon quantum dots and their applications. (A) Three-electrode BES designed for EF, and possible major reactions in the presence of a WE poised at a low (blue) or high (red) potential. (2008), Electrochemical interrogations of the Mtr cytochromes from, Bacterial manganese reduction and growth with manganese oxide as the sole electron acceptor, Miyahara M., Hashimoto K. and Watanabe K. (2013), Use of cassette-electrode microbial fuel cell for wastewater treatment, Kitayama M., Koga R., Kasai T., Kouzuma A. and Watanabe K. (2017), Structures, compositions, and activities of live, Silva A.V., Edel M., Gescher J. and Paquete C.M. Li, C. et al. Scale bars in the inserts are 1m. Proc. Consistently, FTIR analysis (Fig. For the extracellular polymeric substances, many proteins and other bio-macromolecules are required. will also be available for a limited time. Microbiol. The addition of the CDs can also promote cellular adhesion, electronegativity, and biofilm formation. Prepare S. oneidensis overnight cultures in tryptic soy broth at 30C with continuous shaking (250 rpm). Atmosphere. Putative multidrug efflux transporters, detoxification proteins, extracytoplasmic sigma factors and PAS domain regulators are shown to have higher expression activity in presence of heavy metal. In this case, the concentration of sodium lactate was 5mmolL1, and the reactor volume was 50mL. Starwalt-Lee R, El-Naggar MY, Bond DR, Gralnick JA. Sci. carried out ATP assay; T.B. The involvement of cell-to-cell signals in the development of a bacterial biofilm. [15][16][17], "Polyphasic taxonomy of the genus Shewanella and description of Shewanella oneidensis sp. The excitation and emission wavelengths are 535 and 617nm for PI, 485 and 498nm for Syto 9. A pronounced difference in contrast was observed in the tomograms between the membranes of the CDs-fed cells (Fig. The protein concentration of the solution was measured by BCA assay kit. The genus Shewanella comprises over 70 species and includes bacterial strains isolated from diverse ecosystems, including freshwater sediment, deep-sea sediment, oil brine, fish bodies, and spoiled foods [1,2]. Print 2016 Sep 1. Different from conventional fermentation processes, in which the total redox equivalent must be balanced between substrates and products, bioprocesses using EAB are considered to be able to produce either oxidized or reduced products at high yields using electrodes as electron scavengers or suppliers, respectively, and such a technology has been termed electro-fermentation (EF) [13,14]. In addition, recent studies have proposed electro-genetics (EG), in which gene expression in EAB is regulated by intracellular redox-sensing systems whose function is controllable using electrodes in BESs [15]. Ed. Barchinger SE, Pirbadian S, Sambles C, Baker CS, Leung KM, Burroughs NJ, El-Naggar MY, Golbeck JH. Article Enhancing extracellular electron transfer of Shewanella oneidensis MR-1 through coupling improved flavin synthesis and metal-reducing conduit for pollutant degradation. Meanwhile, the CDs promote cellular adhesion, electronegativity, and biofilm formation. Furthermore, some catabolic genes are regulated by multiple global regulatory systems [30], suggesting the presence of complex regulatory cascades in MR-1. Publishers note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. and C.Y. Provided by the Springer Nature SharedIt content-sharing initiative, Journal of Thermal Analysis and Calorimetry (2022). However, the capability of the three CDs specimens to induce these enhancements was distinct from one another. The extracellular electron transfer activity is mainly divided into two mechanisms according to the transfer path: (1) direct electron transfer from the cells to extracellular electron acceptors via bacterial membrane-bounded cytochromes and/or conductive bacterial pili nanowires24,25,26; (2) mediated electron transfer via soluble redox mediators, e.g., flavins23,27,28. In this work, feeding with the CDs, S. oneidensis MR-1 showed an accelerated metabolic rate, as supported by the evidence of (1) more intracellular electrons, (2) higher ATP level, (3) quicker substrate utilization, (4) incremental extracellular excretion and accumulated biofilms, and (5) boosted transmembrane and extracellular electron transfer. J. The vitrified samples were loaded into a Titan Krios (Thermo Fischer Scientific, USA) cryo-transmission electron microscope equipped with image Cs corrector and a Gatan Quantum 967 Special energy filter with a K2 Summit direct electron detector. National Library of Medicine Deionized (DI) water with a resistivity of 18.2 Mcm was obtained from a Milli-Q Water Purification System and used in all the experiments. Among various bacteria applied for this purpose, Shewanella oneidensis (S. oneidensis) MR-1 is the most representative, due to their prominent competency on electron generation as well as electron transfer from the quinone and quinol pool in the cytoplasmic membrane to extracellular electron acceptors, e.g., minerals containing Fe(III) or electrodes of fuel cells4,5. Cyclic voltammetry (CV) curves in the MEC system of the CDs-fed cells (denoted as MEC-CD) were measured. silver nanoparticles), or the hydrogenase pathway[13] (e.g. Logan, B. E. Exoelectrogenic bacteria that power microbial fuel cells. Article A subsequent study has shown that the diguanylate synthase is involved in regulating the global level of cyclic-di-GMP (c-di-GMP) in MR-1 cells and facilitating biofilm formation on electrodes [52]. Anodic current was generated with lactate as an electron donor in the presence of WE at +0.6 V (vs. SHE), while cathodic current was generated with fumarate as an electron acceptor in the presence of WE at 0.4 V. Current density was calculated based on a projected area of WE. eCollection 2022. Angew. Subsequently, triethylamine (0.2mL) was added into the solution and stirred continuously for 4h. Afterwards, the product was evaporated by the rotary evaporation apparatus under low pressure at 30C, then re-dispersed and centrifuged at 3821g. +0.6 V vs. SHE), resulting in the activation of the NADH-dependent pathway at high potentials (see Figure 1). Membrane roughness as a sensitive parameter reflecting the status of neuronal cells in response to chemical and nanoparticle treatments. Bioelectrochemistry 81, 7480 (2011). Abbreviations: Ack, acetate kinase; Atp, ATP synthetase; cAMP/CRP, cyclic AMP/cAMP-receptor protein; Dld, In the presence of a low-potential electrode, ArcS is reduced (ArcS. Electrolocation? High power density from a miniature microbial fuel cell using Shewanella oneidensis DSP10. Adv. The promotion of metabolic activities and electron generation/transfer of S. oneidensis MR-1 by CDs addition holds tremendous potential to improve, diversify, and innovate the aforementioned technologies and related practices. This idea has been confirmed in a study that shows that the introduction of genes encoding sugar permease and hexose kinase in Escherichia coli facilitates MR-1 to grow on glucose in the presence of electron acceptors, e.g. Bacteria that can form electric circuits? Sci. Herein, we report on the augmentation of S. oneidensis MR-1 with carbon dots (CDs). The https:// ensures that you are connecting to the Google Scholar. or F.B. a, b TEM images and size distribution, c, d HRTEM images, and e, f AFM images and height profile of the linescan of CD-X1and CD-X2. However, it coped with iron depletion better than the wild-type strain MR-1. and N.K. Researchers are also attempting to utilize its electrochemical ability for controlling gene expression, for which electro-genetics (EG) has been coined. The authors declare no competing interests. 2004 Sep;70(9):5415-25. doi: 10.1128/AEM.70.9.5415-5425.2004. 1) presented two peaks corresponding to C and O, respectively. Rev. https://en.wikipedia.org/w/index.php?title=Shewanella_oneidensis&oldid=1093314379, This page was last edited on 15 June 2022, at 20:25. One such electron acceptor is Fe 2 O 3, which is commonly found in the clay hematite. Chem. PubMed Central S. oneidensis MR-1 belongs to a class of bacteria known as "Dissimilatory Metal-Reducing Bacteria (DMRB)" because of their ability to couple metal reduction with their metabolism. Superscript A indicates enzymes whose gene expression is regulated by the Arc system, superscript C indicates enzymes whose gene expression is regulated by the cAMP/CRP system, while superscript F indicates enzymes whose gene expression is regulated by the Fnr system. Breuer, M., Rosso, K. M. & Blumberger, J. Electron flow in multiheme bacterial cytochromes is a balancing act between heme electronic interaction and redox potentials. Adv. Among these, MFCs are expected to be applied to power generation from biomass wastes [10], while MECs are designed to generate hydrogen gas, CO2-free clean energy, at high efficiencies [11]. The extracellular electron transport chains responsible for transferring electrons across cell membranes are relatively well characterized, in particular outer membrane c-type cytochromes MtrC and OmcA. This is an open access article published by Portland Press Limited on behalf of the Biochemical Society and distributed under the. The transmembrane electron transfer refers to delivering electrons from redox carriers in the cytoplasm to the cell outer surface over the cell membranes and periplasm. bioelectrochemistry; bioenergetics; biofilm; bioproduction; fermentation. 2007 Jul;65(1):12-20. doi: 10.1111/j.1365-2958.2007.05783.x. For the MFC measurements, a double-chamber reactor (100mL of each chamber) was used at a constant resistant load of 1.0k. To explore the CDs surface charge on the efficacy of bioelectricity enhancement, two additional samples, i.e., CD-X1 and CD-X2, were synthesized, with ethylenediamine/propylamine as the precursor instead of triethylamine, respectively. Optical images of S. oneidensis MR-1 cells b, d without and c, e with the CDs on the neutral (b, c) and positively charged (d, e, +charged) glass surfaces, respectively. See this image and copyright information in PMC. Shewanella oneidensis MR-1 is a metal-reducing bacterium with the ability to utilize many different terminal electron acceptors, including oxygen and solid-metal oxides. (1) The surface oxygen related groups of CDs increase the electronegativity of the bacteria hence improve their static and dynamic adhesion and charge transfer at the anode. i, j TEM images of cross-section slices of the CDs-fed cells, where the dark clusters are assigned to accumulated LBDs. PubMedGoogle Scholar. 2a) and atomic force microscope (AFM) images (Fig. S. oneidensis MR-1 and OmcA/MtrC mutant were cultured at 30C in LB broth (100mL) aerobically at 150rpm for 12h until the logarithmic stationary phase was achieved. ADS Epub 2022 May 3. Shewanella oneidensis is a gram negative bacterium which was first discovered in the Lake Oneida in the New York state in the USA in the year 1988. Adv. Chem. The pristine/treated cells were added to C-flat 2/2 200 mesh copper grids (Protochips) and plunge frozen using a Leica GP2 plunge freezer. 27, 41694177 (2015). The boosted transmembrane electron transfer (Fig. PubMed Another mutant with disruption in the cytochrome c maturation gene ccmA was unable to produce any cytochrome c and to grow on vanadate. doi: 10.1128/AEM.02668-18. Kaim, G. & Dimroth, P. ATP synthesis by F-type ATP synthase is obligatorily dependent on the transmembrane voltage. Bioelectricity produced by electrochemically active bacteria has sparked substantial interests, showing considerable prospects for sustainable energy production, environmental pollution treatment, as well as fabrication of metal nanoparticles1,2,3. The authors declare that all data supporting the findings of this study are available within the paper [and its supplementary information files]. Technol. 61, 237258 10.1146/annurev.micro.61.080706.093257 In these studies, S. oneidensis MR-1 serves as a bacterial platform that provides researchers with novel interdisciplinary ideas that bridge over electrochemistry and biochemistry. Since efficiencies of biotechnology processes carried out in BESs (electro-biotechnology, EB) are determined by metabolic and electrochemical properties of EAB as well as their interaction with electrodes as biofilms, we review knowledge on these features of strain MR-1 and suggest future directions of studies for developing efficient EB processes. or F.Z. We will conclude that this bacterium serves as a bacterial platform for developing EB with broad applications. Atsushi Kouzuma reviewed all sections, and Kazuya Watanabe coordinated the contributions by co-authors and revised the manuscript. The biofilm needs bacterial cells to move in a certain manner, while flagella is the organelle which has locomotive function. Insights into the Biosynthesis of Nanoparticles by the Genus. [1] S. oneidensisis a facultative bacterium, capable of surviving and proliferating in both aerobicand anaerobicconditions. oxygen and fumarate [21]. In addition, it has also been shown that this engineered strain generates anodic current in glucose-supplemented BES via a working electrode (WE) poised at +0.6 V (vs. a standard hydrogen electrode, SHE) and produces lactate as an intermediate metabolite and acetate as the end product [21]. Natl Acad. This species is referred to as S. oneidensis MR-1, indicating "manganese reducing", a special feature of this organism. In contrast, in the presence of a high-potential electrode, ArcS is oxidized (ArcSO) by oxidized quinones (Q), resulting in the release of dephosphorylated ArcA from promoter regions. In 2002, its genomic sequence was published. In CV, WE potential is repeatedly sweeped between two values for measuring current under potential-changing conditions. 3a, b). Sci. The evidence for redox-mediated taxis in Shewanella oneidensis. palladium nanoparticles). PubMed Central Kumar, A. et al. Sci. [6], DSFO+ could possibly allow the bacteria to electrically communicate with an electrode and generate electricity in a wastewater application. g, h TEM image of a CDs-fed cell constructed using cryo-electron tomography, where the LBDs located at the different zones of the cell are marked by the solid-line circles in different colors. 4g, h) of the CDs-fed S. oneidensis MR-1 thus suggests the possible interaction between the CDs and these cytochromes. sharing sensitive information, make sure youre on a federal Hartshorne, R. et al. Biosci Biotechnol Biochem. Google Scholar. d C 1s XP, e O 1s XP, and f FTIR spectra of the CDs. The genus Shewanella comprises over 70 species of heterotrophic bacteria with versatile respiratory capacities. This work was financially supported by the National Key R&D Program of China (2017YFA0207201), National Natural Science Foundation of China (21473045, 51772066, 21777155, and 41471260), and State Key Laboratory of Urban Water Resource and Environment (2018DX04). [14] A re-annotation was made in 2003. 14, 9 (2016). . 4a, b), whilst obvious black spots were found from the images of the cells after 6h incubation with the CDs (Fig. OD485 of supernatant was measured by ultravioletvisible absorption spectroscopy. The CV test for S. oneidensis MR-1 and its OmcA/MtrC mutant with a scan rate of 10mVs1 were performed within the potential region from 0.60 to 0.20V and 0.60 to 0.40V, respectively. Rep. 3, 3307 (2013). Microbiol. Prokaryotes 6, 11331151 10.1007/0-387-30746-X_45 Sun, M., Zhai, L., Li, W. & Yu, H. 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