{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,2]],"date-time":"2026-10-02T10:18:53Z","timestamp":1790936333386,"version":"4.1.0"},"reference-count":103,"publisher":"Portland Press Ltd.","issue":"4","license":[{"start":{"date-parts":[[2026,2,2]],"date-time":"2026-02-02T00:00:00Z","timestamp":1769990400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["portlandpress.com"],"crossmark-restriction":true},"short-container-title":[],"abstract":"<jats:p>Microbial nanowires are appendages that Bacteria and Archaea use to transfer electrons to external surfaces, such as minerals, electrodes, or other microbes. While initial studies suggested that nanowires were modified pili, recent advancements in cryo-EM revealed that microbial nanowires are composed of multi-heme c-type cytochromes. In this review, we discuss the discovery of microbial nanowires, advancements that allowed elucidation of their near-atomic resolution cryo-EM structures, and the impact of heme arrangement on electron transfer. We also discuss how new structural information can be used to identify filaments in images from published literature. The structural insights gained from these studies provide a deeper understanding of the mechanisms underlying long-range electron transport in microbial nanowires and their potential applications in bioelectronics and energy-generating microbial fuel cells.<\/jats:p>","DOI":"10.1042\/etls20240008","type":"journal-article","created":{"date-parts":[[2026,2,2]],"date-time":"2026-02-02T14:05:35Z","timestamp":1770041135000},"update-policy":"https:\/\/doi.org\/10.1042\/crossmark_policy","source":"Crossref","is-referenced-by-count":3,"title":["Cytochrome nanowires under electron microscopy"],"prefix":"10.1042","volume":"8","author":[{"given":"Holly A.","family":"Petersen","sequence":"first","affiliation":[{"name":"1Department of Biochemistry and Molecular Genetics, University of Alabama at Birmingham, Birmingham, AL 35233, U.S.A."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Allon I.","family":"Hochbaum","sequence":"additional","affiliation":[{"name":"2Department of Chemical and Biomolecular Engineering, Materials Science and Engineering, Chemistry, and Molecular Biology and Biochemistry, University of California Irvine, Irvine, CA 92697, U.S.A."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daniel R.","family":"Bond","sequence":"additional","affiliation":[{"name":"3Department of Plant and Microbial Biology, and BioTechnology Institute, University of Minnesota, Saint Paul, MN 55455, U.S.A."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1008-663X","authenticated-orcid":false,"given":"Fengbin","family":"Wang","sequence":"additional","affiliation":[{"name":"1Department of Biochemistry and Molecular Genetics, University of Alabama at Birmingham, Birmingham, AL 35233, U.S.A."},{"name":"4Gregory Fleming James Cystic Fibrosis Research Center, University of Alabama at Birmingham, Birmingham, AL 35233, U.S.A."}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"288","published-online":{"date-parts":[[2026,2,2]]},"reference":[{"key":"2026020209053273400_ETLS-2024-0008C1","doi-asserted-by":"crossref","first-page":"651","DOI":"10.1038\/nrmicro.2016.93","article-title":"Extracellular electron transfer mechanisms between microorganisms and minerals","volume":"14","author":"Shi","year":"2016","journal-title":"Nat. Rev. Microbiol."},{"key":"2026020209053273400_ETLS-2024-0008C2","doi-asserted-by":"crossref","first-page":"1319","DOI":"10.1126\/science.240.4857.1319","article-title":"Bacterial manganese reduction and growth with manganese oxide as the sole electron acceptor","volume":"240","author":"Myers","year":"1988","journal-title":"Science"},{"key":"2026020209053273400_ETLS-2024-0008C3","doi-asserted-by":"crossref","first-page":"1472","DOI":"10.1128\/aem.54.6.1472-1480.1988","article-title":"Novel mode of microbial energy metabolism: organic carbon oxidation coupled to dissimilatory reduction of iron or manganese","volume":"54","author":"Lovley","year":"1988","journal-title":"Appl. Environ. Microbiol."},{"key":"2026020209053273400_ETLS-2024-0008C4","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1021\/es9026248","article-title":"Biogeochemical redox processes and their impact on contaminant dynamics","volume":"44","author":"Borch","year":"2010","journal-title":"Environ. Sci. Technol."},{"key":"2026020209053273400_ETLS-2024-0008C5","doi-asserted-by":"crossref","first-page":"4599","DOI":"10.1128\/AEM.00895-14","article-title":"Direct interspecies electron transfer between Geobacter metallireducens and Methanosarcina barkeri","volume":"80","author":"Rotaru","year":"2014","journal-title":"Appl. Environ. Microbiol."},{"key":"2026020209053273400_ETLS-2024-0008C6","doi-asserted-by":"crossref","first-page":"1413","DOI":"10.1126\/science.1196526","article-title":"Direct exchange of electrons within aggregates of an evolved syntrophic coculture of anaerobic bacteria","volume":"330","author":"Summers","year":"2010","journal-title":"Science"},{"key":"2026020209053273400_ETLS-2024-0008C7","doi-asserted-by":"crossref","first-page":"1548","DOI":"10.1128\/AEM.69.3.1548-1555.2003","article-title":"Electricity production by Geobacter sulfurreducens attached to electrodes","volume":"69","author":"Bond","year":"2003","journal-title":"Appl. Environ. Microbiol."},{"key":"2026020209053273400_ETLS-2024-0008C8","doi-asserted-by":"crossref","first-page":"1124","DOI":"10.1111\/mmi.14801","article-title":"Geobacter sulfurreducens inner membrane cytochrome CbcBA controls electron transfer and growth yield near the energetic limit of respiration","volume":"116","author":"Joshi","year":"2021","journal-title":"Mol. Microbiol."},{"key":"2026020209053273400_ETLS-2024-0008C9","doi-asserted-by":"crossref","first-page":"e02134-18","DOI":"10.1128\/AEM.02134-18","article-title":"Divergent Nrf Family Proteins and MtrCAB Homologs facilitate extracellular electron transfer in aeromonas hydrophila","volume":"84","author":"Conley","year":"2018","journal-title":"Appl. Environ. Microbiol."},{"key":"2026020209053273400_ETLS-2024-0008C10","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1016\/j.bioelechem.2015.08.003","article-title":"Reduction of low potential electron acceptors requires the CbcL inner membrane cytochrome of Geobacter sulfurreducens","volume":"107","author":"Zacharoff","year":"2016","journal-title":"Bioelectrochemistry"},{"key":"2026020209053273400_ETLS-2024-0008C11","doi-asserted-by":"crossref","first-page":"e02034-14","DOI":"10.1128\/mBio.02034-14","article-title":"An inner membrane cytochrome required only for reduction of high redox potential extracellular electron acceptors","volume":"5","author":"Levar","year":"2014","journal-title":"MBio"},{"key":"2026020209053273400_ETLS-2024-0008C12","doi-asserted-by":"crossref","first-page":"1143","DOI":"10.1128\/jb.179.4.1143-1152.1997","article-title":"Cloning and sequence of cymA, a gene encoding a tetraheme cytochrome c required for reduction of iron(III), fumarate, and nitrate by Shewanella putrefaciens MR-1","volume":"179","author":"Myers","year":"1997","journal-title":"J. Bacteriol."},{"key":"2026020209053273400_ETLS-2024-0008C13","doi-asserted-by":"crossref","first-page":"00347","DOI":"10.1128\/JB.00347-18","article-title":"Identification of different putative outer membrane electron conduits necessary for Fe(III) Citrate, Fe(III) Oxide, Mn(IV) Oxide, or electrode reduction by Geobacter sulfurreducens","volume":"200","author":"Jim\u00e9nez Otero","year":"2018","journal-title":"J. Bacteriol."},{"key":"2026020209053273400_ETLS-2024-0008C14","doi-asserted-by":"crossref","first-page":"776","DOI":"10.1111\/1758-2229.12204","article-title":"A trans-outer membrane porin-cytochrome protein complex for extracellular electron transfer by Geobacter sulfurreducens\u2005PCA","volume":"6","author":"Liu","year":"2014","journal-title":"Environ. Microbiol. Rep."},{"key":"2026020209053273400_ETLS-2024-0008C15","doi-asserted-by":"crossref","first-page":"22169","DOI":"10.1073\/pnas.0900086106","article-title":"Characterization of an electron conduit between bacteria and the extracellular environment","volume":"106","author":"Hartshorne","year":"2009","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"2026020209053273400_ETLS-2024-0008C16","doi-asserted-by":"crossref","first-page":"789","DOI":"10.1080\/08927014.2012.710324","article-title":"Electrochemically active biofilms: facts and fiction. A review","volume":"28","author":"Babauta","year":"2012","journal-title":"Biofouling"},{"key":"2026020209053273400_ETLS-2024-0008C17","doi-asserted-by":"crossref","first-page":"3968","DOI":"10.1073\/pnas.0710525105","article-title":"Shewanella secretes flavins that mediate extracellular electron transfer","volume":"105","author":"Marsili","year":"2008","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"2026020209053273400_ETLS-2024-0008C18","doi-asserted-by":"crossref","first-page":"1098","DOI":"10.1038\/nature03661","article-title":"Extracellular electron transfer via microbial nanowires","volume":"435","author":"Reguera","year":"2005","journal-title":"Nature"},{"key":"2026020209053273400_ETLS-2024-0008C19","doi-asserted-by":"crossref","first-page":"8354","DOI":"10.1039\/C5RA28092C","article-title":"Conductivity of individual Geobacter pili","volume":"6","author":"Adhikari","year":"2016","journal-title":"RSC Adv."},{"key":"2026020209053273400_ETLS-2024-0008C20","doi-asserted-by":"crossref","first-page":"1012","DOI":"10.1038\/nnano.2014.236","article-title":"Visualization of charge propagation along individual pili proteins using ambient electrostatic force microscopy","volume":"9","author":"Malvankar","year":"2014","journal-title":"Nat. Nanotechnol."},{"key":"2026020209053273400_ETLS-2024-0008C21","doi-asserted-by":"crossref","first-page":"573","DOI":"10.1038\/nnano.2011.119","article-title":"Tunable metallic-like conductivity in microbial nanowire networks","volume":"6","author":"Malvankar","year":"2011","journal-title":"Nat. Nanotechnol."},{"key":"2026020209053273400_ETLS-2024-0008C22","doi-asserted-by":"crossref","first-page":"786416","DOI":"10.3389\/fbioe.2021.786416","article-title":"Biofilm Biology and Engineering of Geobacter and Shewanella spp. for Energy Applications","volume":"9","author":"Hu","year":"2021","journal-title":"Front. Bioeng. Biotechnol."},{"key":"2026020209053273400_ETLS-2024-0008C23","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1016\/j.cbpa.2020.08.004","article-title":"The blind men and the filament: Understanding structures and functions of microbial nanowires","volume":"59","author":"Yalcin","year":"2020","journal-title":"Curr. Opin. Chem. Biol"},{"key":"2026020209053273400_ETLS-2024-0008C24","doi-asserted-by":"crossref","first-page":"1291","DOI":"10.1038\/s41564-022-01159-z","article-title":"Cryo-EM structure of an extracellular Geobacter OmcE cytochrome filament reveals tetrahaem packing","volume":"7","author":"Wang","year":"2022","journal-title":"Nat. Microbiol."},{"key":"2026020209053273400_ETLS-2024-0008C25","doi-asserted-by":"crossref","first-page":"e81551","DOI":"10.7554\/eLife.81551","article-title":"Structure of Geobacter OmcZ filaments suggests extracellular cytochrome polymers evolved independently multiple times","volume":"11","author":"Wang","year":"2022","journal-title":"Elife"},{"key":"2026020209053273400_ETLS-2024-0008C26","doi-asserted-by":"crossref","first-page":"361","DOI":"10.1016\/j.cell.2019.03.029","article-title":"Structure of microbial nanowires reveals stacked hemes that transport electrons over micrometers","volume":"177","author":"Wang","year":"2019","journal-title":"Cell"},{"key":"2026020209053273400_ETLS-2024-0008C27","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1038\/s41564-022-01315-5","article-title":"Structure of Geobacter cytochrome OmcZ identifies mechanism of nanowire assembly and conductivity","volume":"8","author":"Gu","year":"2023","journal-title":"Nat. Microbiol."},{"key":"2026020209053273400_ETLS-2024-0008C28","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1038\/s42003-019-0448-9","article-title":"Cryo-EM reveals the structural basis of long-range electron transport in a cytochrome-based bacterial nanowire","volume":"2","author":"Filman","year":"2019","journal-title":"Commun. Biol."},{"key":"2026020209053273400_ETLS-2024-0008C29","doi-asserted-by":"crossref","first-page":"430","DOI":"10.1038\/s41586-021-03857-w","article-title":"Structure of Geobacter pili reveals secretory rather than nanowire behaviour","volume":"597","author":"Gu","year":"2021","journal-title":"Nature"},{"key":"2026020209053273400_ETLS-2024-0008C30","doi-asserted-by":"crossref","first-page":"2142","DOI":"10.1038\/s41396-018-0193-y","article-title":"Syntrophic growth with direct interspecies electron transfer between pili-free Geobacter species","volume":"12","author":"Liu","year":"2018","journal-title":"ISME J."},{"key":"2026020209053273400_ETLS-2024-0008C31","doi-asserted-by":"crossref","first-page":"15248","DOI":"10.1073\/pnas.1108616108","article-title":"Extracellular reduction of uranium via Geobacter conductive pili as a protective cellular mechanism","volume":"108","author":"Cologgi","year":"2011","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"2026020209053273400_ETLS-2024-0008C32","doi-asserted-by":"crossref","first-page":"1967","DOI":"10.1126\/science.1088727","article-title":"Genome of Geobacter sulfurreducens: metal reduction in subsurface environments","volume":"302","author":"Meth\u00e9","year":"2003","journal-title":"Science"},{"key":"2026020209053273400_ETLS-2024-0008C33","doi-asserted-by":"crossref","first-page":"3180","DOI":"10.1128\/AEM.67.7.3180-3187.2001","article-title":"Development of a genetic system for Geobacter sulfurreducens","volume":"67","author":"Coppi","year":"2001","journal-title":"Appl. Environ. Microbiol."},{"key":"2026020209053273400_ETLS-2024-0008C34","doi-asserted-by":"crossref","first-page":"e00340-17","DOI":"10.1128\/JB.00340-17","article-title":"Genome scale mutational analysis of Geobacter sulfurreducens reveals distinct molecular mechanisms for respiration and sensing of poised electrodes versus Fe(III) Oxides","volume":"199","author":"Chan","year":"2017","journal-title":"J. Bacteriol."},{"key":"2026020209053273400_ETLS-2024-0008C35","doi-asserted-by":"crossref","first-page":"7178","DOI":"10.1128\/AEM.01967-15","article-title":"Scarless genome editing and stable inducible expression vectors for Geobacter sulfurreducens","volume":"81","author":"Chan","year":"2015","journal-title":"Appl. Environ. Microbiol."},{"key":"2026020209053273400_ETLS-2024-0008C36","doi-asserted-by":"crossref","first-page":"483","DOI":"10.1126\/science.1066771","article-title":"Electrode-reducing microorganisms that harvest energy from marine sediments","volume":"295","author":"Bond","year":"2002","journal-title":"Science"},{"key":"2026020209053273400_ETLS-2024-0008C37","doi-asserted-by":"crossref","first-page":"327","DOI":"10.1038\/ismej.2016.136","article-title":"Happy together: microbial communities that hook up to swap electrons","volume":"11","author":"Lovley","year":"2017","journal-title":"ISME J."},{"key":"2026020209053273400_ETLS-2024-0008C38","doi-asserted-by":"crossref","first-page":"497","DOI":"10.1038\/nrmicro1442","article-title":"Bug juice: harvesting electricity with microorganisms","volume":"4","author":"Lovley","year":"2006","journal-title":"Nat. Rev. Microbiol."},{"key":"2026020209053273400_ETLS-2024-0008C39","doi-asserted-by":"crossref","first-page":"e00369-25","DOI":"10.1128\/aem.00369-25","article-title":"Extracellular electron transfer proteins contribute to reduction of ferric minerals by Geobacter biofilms","volume":"91","author":"Xu","year":"2025","journal-title":"Appl. Environ. Microbiol."},{"key":"2026020209053273400_ETLS-2024-0008C40","doi-asserted-by":"crossref","first-page":"2248","DOI":"10.1128\/AEM.66.5.2248-2251.2000","article-title":"Lack of production of electron-shuttling compounds or solubilization of Fe(III) during reduction of insoluble Fe(III) oxide by Geobacter metallireducens","volume":"66","author":"Nevin","year":"2000","journal-title":"Appl. Environ. Microbiol."},{"key":"2026020209053273400_ETLS-2024-0008C41","doi-asserted-by":"crossref","first-page":"5790","DOI":"10.1039\/c2ee03388g","article-title":"Biofilm conductivity is a decisive variable for high-current-density Geobacter sulfurreducens microbial fuel cells","volume":"5","author":"Malvankar","year":"2012","journal-title":"Energy Environ. Sci."},{"key":"2026020209053273400_ETLS-2024-0008C42","doi-asserted-by":"crossref","first-page":"20716","DOI":"10.1073\/pnas.1912498116","article-title":"NanoSIMS imaging reveals metabolic stratification within current-producing biofilms","volume":"116","author":"Chadwick","year":"2019","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"2026020209053273400_ETLS-2024-0008C43","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1016\/j.jelechem.2016.11.005","article-title":"Proving Geobacter biofilm connectivity with confocal Raman microscopy","volume":"793","author":"Robuschi","year":"2017","journal-title":"J. Electroanal. Chem. (Lausanne)."},{"key":"2026020209053273400_ETLS-2024-0008C44","doi-asserted-by":"crossref","first-page":"598","DOI":"10.1002\/cssc.201300605","article-title":"Physiological stratification in electricity-producing biofilms of Geobacter sulfurreducens","volume":"7","author":"Schrott","year":"2014","journal-title":"ChemSusChem"},{"key":"2026020209053273400_ETLS-2024-0008C45","doi-asserted-by":"crossref","first-page":"1957","DOI":"10.1002\/celc.201402168","article-title":"Electron Transport through Early Exponential-Phase Anode-Grown Geobacter sulfurreducens Biofilms","volume":"1","author":"Strycharz-Glaven","year":"2014","journal-title":"ChemElectroChem"},{"key":"2026020209053273400_ETLS-2024-0008C46","doi-asserted-by":"crossref","first-page":"15467","DOI":"10.1073\/pnas.1209829109","article-title":"Long-range electron transport in Geobacter sulfurreducens biofilms is redox gradient-driven","volume":"109","author":"Snider","year":"2012","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"2026020209053273400_ETLS-2024-0008C47","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1016\/j.tibs.2014.10.005","article-title":"How cryo-EM is revolutionizing structural biology","volume":"40","author":"Bai","year":"2015","journal-title":"Trends Biochem. Sci."},{"key":"2026020209053273400_ETLS-2024-0008C48","doi-asserted-by":"crossref","first-page":"1443","DOI":"10.1126\/science.1251652","article-title":"The resolution revolution","volume":"343","author":"K\u00fchlbrandt","year":"2014","journal-title":"Science"},{"key":"2026020209053273400_ETLS-2024-0008C49","doi-asserted-by":"crossref","first-page":"584","DOI":"10.1038\/nmeth.2472","article-title":"Electron counting and beam-induced motion correction enable near-atomic-resolution single-particle cryo-EM","volume":"10","author":"Li","year":"2013","journal-title":"Nat. Methods"},{"key":"2026020209053273400_ETLS-2024-0008C50","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1016\/j.jsb.2012.09.006","article-title":"RELION: implementation of a Bayesian approach to cryo-EM structure determination","volume":"180","author":"Scheres","year":"2012","journal-title":"J. Struct. Biol"},{"key":"2026020209053273400_ETLS-2024-0008C51","doi-asserted-by":"crossref","first-page":"290","DOI":"10.1038\/nmeth.4169","article-title":"cryoSPARC: algorithms for rapid unsupervised cryo-EM structure determination","volume":"14","author":"Punjani","year":"2017","journal-title":"Nat. Methods"},{"key":"2026020209053273400_ETLS-2024-0008C52","doi-asserted-by":"crossref","first-page":"13915","DOI":"10.1021\/acs.chemrev.1c00850","article-title":"Emerging Themes in CryoEM\u2500Single Particle Analysis Image Processing","volume":"122","author":"Vilas","year":"2022","journal-title":"Chem. Rev."},{"key":"2026020209053273400_ETLS-2024-0008C53","doi-asserted-by":"crossref","first-page":"4169","DOI":"10.1042\/BCJ20210708","article-title":"New tools for automated cryo-EM single-particle analysis in RELION-4.0","volume":"478","author":"Kimanius","year":"2021","journal-title":"Biochem. J."},{"key":"2026020209053273400_ETLS-2024-0008C54","doi-asserted-by":"crossref","first-page":"5181","DOI":"10.1074\/jbc.REV118.005602","article-title":"Challenges and opportunities in cryo-EM single-particle analysis","volume":"294","author":"Lyumkis","year":"2019","journal-title":"J. Biol. Chem."},{"key":"2026020209053273400_ETLS-2024-0008C55","doi-asserted-by":"crossref","first-page":"450","DOI":"10.1016\/j.cell.2015.03.049","article-title":"Single-Particle Cryo-EM at Crystallographic Resolution","volume":"161","author":"Cheng","year":"2015","journal-title":"Cell"},{"key":"2026020209053273400_ETLS-2024-0008C56","doi-asserted-by":"crossref","first-page":"14055","DOI":"10.1021\/acs.chemrev.1c00753","article-title":"Cryo-EM of Helical Polymers","volume":"122","author":"Wang","year":"2022","journal-title":"Chem. Rev."},{"key":"2026020209053273400_ETLS-2024-0008C57","doi-asserted-by":"crossref","first-page":"1401","DOI":"10.1038\/s41564-019-0458-x","article-title":"An extensively glycosylated archaeal pilus survives extreme conditions","volume":"4","author":"Wang","year":"2019","journal-title":"Nat. Microbiol"},{"key":"2026020209053273400_ETLS-2024-0008C58","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1038\/nmeth.3287","article-title":"De novo protein structure determination from near-atomic-resolution cryo-EM maps","volume":"12","author":"Wang","year":"2015","journal-title":"Nat. Methods"},{"key":"2026020209053273400_ETLS-2024-0008C59","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1038\/s41592-019-0637-y","article-title":"Bottom-up structural proteomics: cryoEM of protein complexes enriched from the cellular milieu","volume":"17","author":"Ho","year":"2020","journal-title":"Nat. Methods"},{"key":"2026020209053273400_ETLS-2024-0008C60","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1038\/s41586-021-03819-2","article-title":"Highly accurate protein structure prediction with AlphaFold","volume":"596","author":"Jumper","year":"2021","journal-title":"Nature"},{"key":"2026020209053273400_ETLS-2024-0008C61","doi-asserted-by":"crossref","first-page":"2840","DOI":"10.1016\/j.bpj.2022.06.025","article-title":"DeepTracer-ID: De novo protein identification from cryo-EM maps","volume":"121","author":"Chang","year":"2022","journal-title":"Biophys. J"},{"key":"2026020209053273400_ETLS-2024-0008C62","doi-asserted-by":"crossref","first-page":"e2017525118","DOI":"10.1073\/pnas.2017525118","article-title":"DeepTracer for fast de novo cryo-EM protein structure modeling and special studies on CoV-related complexes","volume":"118","author":"Pfab","year":"2021","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"2026020209053273400_ETLS-2024-0008C63","doi-asserted-by":"crossref","first-page":"450","DOI":"10.1038\/s41586-024-07215-4","article-title":"Automated model building and protein identification in cryo-EM maps","volume":"628","author":"Jamali","year":"2024","journal-title":"Nature"},{"key":"2026020209053273400_ETLS-2024-0008C64","doi-asserted-by":"crossref","first-page":"721","DOI":"10.1016\/j.jmb.2003.07.013","article-title":"Optimal determination of particle orientation, absolute hand, and contrast loss in single-particle electron cryomicroscopy","volume":"333","author":"Rosenthal","year":"2003","journal-title":"J. Mol. Biol."},{"key":"2026020209053273400_ETLS-2024-0008C65","doi-asserted-by":"crossref","first-page":"1136","DOI":"10.1038\/s41589-020-0623-9","article-title":"Electric field stimulates production of highly conductive microbial OmcZ nanowires","volume":"16","author":"Yalcin","year":"2020","journal-title":"Nat. Chem. Biol."},{"key":"2026020209053273400_ETLS-2024-0008C66","doi-asserted-by":"crossref","first-page":"384","DOI":"10.1016\/j.tim.2022.11.004","article-title":"Microbial nanowires: type IV pili or cytochrome filaments?","volume":"31","author":"Wang","year":"2023","journal-title":"Trends Microbiol."},{"key":"2026020209053273400_ETLS-2024-0008C67","doi-asserted-by":"crossref","first-page":"2853","DOI":"10.1016\/j.cell.2023.05.012","article-title":"Extracellular cytochrome nanowires appear to be ubiquitous in prokaryotes","volume":"186","author":"Baquero","year":"2023","journal-title":"Cell"},{"key":"2026020209053273400_ETLS-2024-0008C68","doi-asserted-by":"crossref","first-page":"e2207037119","DOI":"10.1073\/pnas.2207037119","article-title":"Archaeal bundling pili of Pyrobaculum calidifontis reveal similarities between archaeal and bacterial biofilms","volume":"119","author":"Wang","year":"2022","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"2026020209053273400_ETLS-2024-0008C69","doi-asserted-by":"crossref","first-page":"e2304256120","DOI":"10.1073\/pnas.2304256120","article-title":"The evolution of archaeal flagellar filaments","volume":"120","author":"Kreutzberger","year":"2023","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"2026020209053273400_ETLS-2024-0008C70","doi-asserted-by":"crossref","first-page":"666","DOI":"10.1038\/s41467-023-36349-8","article-title":"Archaeal DNA-import apparatus is homologous to bacterial conjugation machinery","volume":"14","author":"Beltran","year":"2023","journal-title":"Nat. Commun."},{"key":"2026020209053273400_ETLS-2024-0008C71","doi-asserted-by":"crossref","first-page":"476","DOI":"10.1038\/22802","article-title":"Structure of cytochrome c nitrite reductase","volume":"400","author":"Einsle","year":"1999","journal-title":"Nature"},{"key":"2026020209053273400_ETLS-2024-0008C72","doi-asserted-by":"crossref","first-page":"276","DOI":"10.1038\/nsb0497-276","article-title":"The 2.8 A structure of hydroxylamine oxidoreductase from a nitrifying chemoautotrophic bacterium, Nitrosomonas europaea","volume":"4","author":"Igarashi","year":"1997","journal-title":"Nat. Struct. Biol."},{"key":"2026020209053273400_ETLS-2024-0008C73","doi-asserted-by":"crossref","first-page":"3381","DOI":"10.1039\/b505184c","article-title":"Multi-heme cytochromes--new structures, new chemistry","volume":"2005","author":"Mowat","year":"2005","journal-title":"Dalton Trans."},{"key":"2026020209053273400_ETLS-2024-0008C74","doi-asserted-by":"crossref","first-page":"551","DOI":"10.1099\/00221287-146-3-551","article-title":"Bacterial respiration: a flexible process for a changing environment","volume":"146 ( Pt 3)","author":"Richardson","year":"2000","journal-title":"Microbiology (Reading, Engl.)"},{"key":"2026020209053273400_ETLS-2024-0008C75","doi-asserted-by":"crossref","first-page":"1005","DOI":"10.1038\/2975","article-title":"Heme packing motifs revealed by the crystal structure of the tetra-heme cytochrome c554 from Nitrosomonas europaea","volume":"5","author":"Iverson","year":"1998","journal-title":"Nat. Struct. Biol."},{"key":"2026020209053273400_ETLS-2024-0008C76","doi-asserted-by":"crossref","first-page":"1113","DOI":"10.1038\/70051","article-title":"Structure and mechanism of the flavocytochrome c fumarate reductase of Shewanella putrefaciens MR-1","volume":"6","author":"Leys","year":"1999","journal-title":"Nat. Struct. Biol."},{"key":"2026020209053273400_ETLS-2024-0008C77","doi-asserted-by":"crossref","first-page":"35703","DOI":"10.1074\/jbc.M203866200","article-title":"crystal structures at atomic resolution reveal the novel concept of \u201cElectron-harvesting\u201d AS a Role for the Small Tetraheme Cytochrome c","volume":"277","author":"Leys","year":"2002","journal-title":"Journal of Biological Chemistry"},{"key":"2026020209053273400_ETLS-2024-0008C78","doi-asserted-by":"crossref","first-page":"849","DOI":"10.1021\/bi0301439","article-title":"Family of cytochrome c7-type proteins from Geobacter sulfurreducens: structure of one cytochrome c7 at 1.45 A resolution","volume":"43","author":"Pokkuluri","year":"2004","journal-title":"Biochemistry"},{"key":"2026020209053273400_ETLS-2024-0008C79","doi-asserted-by":"crossref","first-page":"2349","DOI":"10.1002\/prot.22747","article-title":"Heme proteins--diversity in structural characteristics, function, and folding","volume":"78","author":"Smith","year":"2010","journal-title":"Proteins"},{"key":"2026020209053273400_ETLS-2024-0008C80","doi-asserted-by":"crossref","first-page":"9384","DOI":"10.1073\/pnas.1017200108","article-title":"Structure of a bacterial cell surface decaheme electron conduit","volume":"108","author":"Clarke","year":"2011","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"2026020209053273400_ETLS-2024-0008C81","doi-asserted-by":"crossref","first-page":"665","DOI":"10.1016\/j.cell.2020.03.032","article-title":"The crystal structure of a biological insulated transmembrane molecular wire","volume":"181","author":"Edwards","year":"2020","journal-title":"Cell"},{"key":"2026020209053273400_ETLS-2024-0008C82","doi-asserted-by":"crossref","first-page":"2434","DOI":"10.1038\/s41467-024-46192-0","article-title":"Widespread extracellular electron transfer pathways for charging microbial cytochrome OmcS nanowires via periplasmic cytochromes PpcABCDE","volume":"15","author":"Portela","year":"2024","journal-title":"Nat. Commun."},{"key":"2026020209053273400_ETLS-2024-0008C83","doi-asserted-by":"crossref","first-page":"10083","DOI":"10.1021\/acs.jpcb.2c06822","article-title":"Assessing thermal response of redox conduction for Anti-Arrhenius Kinetics in a Microbial Cytochrome Nanowire","volume":"126","author":"Guberman-Pfeffer","year":"2022","journal-title":"J. Phys. Chem. B"},{"key":"2026020209053273400_ETLS-2024-0008C84","doi-asserted-by":"crossref","first-page":"1549441","DOI":"10.3389\/fchem.2025.1549441","article-title":"Cytochrome \u201cnanowires\u201d are physically limited to sub-picoamp currents that suffice for cellular respiration","volume":"13","author":"Guberman-Pfeffer","year":"2025","journal-title":"Front. Chem."},{"key":"2026020209053273400_ETLS-2024-0008C85","doi-asserted-by":"crossref","first-page":"9421","DOI":"10.1021\/acs.jpclett.0c02842","article-title":"Which multi-heme protein complex transfers electrons more efficiently? comparing MtrCAB from Shewanella with OmcS from Geobacter","volume":"11","author":"Jiang","year":"2020","journal-title":"J. Phys. Chem. Lett."},{"key":"2026020209053273400_ETLS-2024-0008C86","doi-asserted-by":"crossref","first-page":"eabm7193","DOI":"10.1126\/sciadv.abm7193","article-title":"A 300-fold conductivity increase in microbial cytochrome nanowires due to temperature-induced restructuring of hydrogen bonding networks","volume":"8","author":"Dahl","year":"2022","journal-title":"Sci. Adv."},{"key":"2026020209053273400_ETLS-2024-0008C87","doi-asserted-by":"crossref","first-page":"11191","DOI":"10.1021\/acs.chemrev.5b00298","article-title":"Recent advances in the theory and molecular simulation of biological electron transfer reactions","volume":"115","author":"Blumberger","year":"2015","journal-title":"Chem. Rev."},{"key":"2026020209053273400_ETLS-2024-0008C88","doi-asserted-by":"crossref","first-page":"3425","DOI":"10.1073\/pnas.1818003116","article-title":"Kinetics of trifurcated electron flow in the decaheme bacterial proteins MtrC and MtrF","volume":"116","author":"Jiang","year":"2019","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"2026020209053273400_ETLS-2024-0008C89","doi-asserted-by":"crossref","first-page":"17237","DOI":"10.1021\/jacs.7b08831","article-title":"Cysteine linkages accelerate electron flow through tetra-heme protein STC","volume":"139","author":"Jiang","year":"2017","journal-title":"J. Am. Chem. Soc"},{"key":"2026020209053273400_ETLS-2024-0008C90","doi-asserted-by":"crossref","first-page":"e2107939118","DOI":"10.1073\/pnas.2107939118","article-title":"Nanosecond heme-to-heme electron transfer rates in a multiheme cytochrome nanowire reported by a spectrally unique His\/Met-ligated heme","volume":"118","author":"Van Wonderen","year":"2021","journal-title":"Proc. Natl. Acad. Sci. U.S.A"},{"key":"2026020209053273400_ETLS-2024-0008C91","doi-asserted-by":"crossref","first-page":"101556","DOI":"10.1016\/j.coelec.2024.101556","article-title":"Do multiheme cytochromes containing close-packed heme groups have a band structure formed from the heme \u03c0 and \u03c0\u2217 orbitals?","volume":"47","author":"Van Wonderen","year":"2024","journal-title":"Current Opinion in Electrochemistry"},{"key":"2026020209053273400_ETLS-2024-0008C92","doi-asserted-by":"crossref","first-page":"8037","DOI":"10.1021\/acs.jpclett.5c01339","article-title":"Coherent electron transfer in cytochrome nanowires at 300 K","volume":"16","author":"Parson","year":"2025","journal-title":"J. Phys. Chem. Lett."},{"key":"2026020209053273400_ETLS-2024-0008C93","doi-asserted-by":"crossref","first-page":"11889","DOI":"10.1021\/acs.jpcb.5c04205","article-title":"Investigating electron conductivity regimes in the bacterial Cytochrome Wire OmcS","volume":"129","author":"Mohanam","year":"2025","journal-title":"J. Phys. Chem. B"},{"key":"2026020209053273400_ETLS-2024-0008C94","doi-asserted-by":"crossref","first-page":"19571","DOI":"10.1038\/s41598-024-70348-z","article-title":"Computation of biological conductance with Liouville quantum master equation","volume":"14","author":"Papp","year":"2024","journal-title":"Sci. Rep."},{"key":"2026020209053273400_ETLS-2024-0008C95","doi-asserted-by":"crossref","first-page":"314002","DOI":"10.1088\/1361-6528\/ab8767","article-title":"Coherence-assisted electron diffusion across the multi-heme protein-based bacterial nanowire","volume":"31","author":"Eshel","year":"2020","journal-title":"Nanotechnology"},{"key":"2026020209053273400_ETLS-2024-0008C96","doi-asserted-by":"crossref","first-page":"9868","DOI":"10.1021\/ja3027696","article-title":"Thermodynamics of electron flow in the bacterial deca-heme cytochrome MtrF","volume":"134","author":"Breuer","year":"2012","journal-title":"J. Am. Chem. Soc."},{"key":"2026020209053273400_ETLS-2024-0008C97","doi-asserted-by":"crossref","first-page":"15582","DOI":"10.1021\/jp057068r","article-title":"Electronic coupling between heme electron-transfer centers and its decay with distance depends strongly on relative orientation","volume":"110","author":"Smith","year":"2006","journal-title":"J. Phys. Chem. B"},{"key":"2026020209053273400_ETLS-2024-0008C98","doi-asserted-by":"crossref","first-page":"693","DOI":"10.1002\/ijch.201600021","article-title":"Going with the Electron Flow: Heme Electronic Structure and Electron Transfer in Cytochrome c","volume":"56","author":"Bren","year":"2016","journal-title":"Isr. J. Chem."},{"key":"2026020209053273400_ETLS-2024-0008C99","doi-asserted-by":"crossref","first-page":"e0085022","DOI":"10.1128\/mbio.00850-22","article-title":"Untangling Geobacter sulfurreducens Nanowires","volume":"13","author":"Lovley","year":"2022","journal-title":"MBio"},{"key":"2026020209053273400_ETLS-2024-0008C100","doi-asserted-by":"crossref","first-page":"e01041-22","DOI":"10.1128\/mbio.01041-22","article-title":"Reply to Lovley, \u201cUntangling Geobacter sulfurreducens Nanowires\u201d","volume":"13","author":"Liu","year":"2022","journal-title":"MBio"},{"key":"2026020209053273400_ETLS-2024-0008C101","doi-asserted-by":"crossref","first-page":"325","DOI":"10.1002\/jemt.1060090404","article-title":"Computer image processing of electron micrographs of biological structures with helical symmetry","volume":"9","author":"Stewart","year":"1988","journal-title":"J. Electron Microsc. Tech."},{"key":"2026020209053273400_ETLS-2024-0008C102","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1016\/j.jsb.2006.05.009","article-title":"EMAN2: an extensible image processing suite for electron microscopy","volume":"157","author":"Tang","year":"2007","journal-title":"J. Struct. Biol."},{"key":"2026020209053273400_ETLS-2024-0008C103","doi-asserted-by":"crossref","first-page":"1901","DOI":"10.1039\/c3ee40441b","article-title":"Engineering Geobacter sulfurreducens to produce a highly cohesive conductive matrix with enhanced capacity for current production","volume":"6","author":"Leang","year":"2013","journal-title":"Energy Environ. Sci."}],"container-title":["Emerging Topics in Life Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/portlandpress.com\/emergtoplifesci\/article-pdf\/doi\/10.1042\/ETLS20240008\/984281\/etls-2024-0008.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/portlandpress.com\/emergtoplifesci\/article-pdf\/doi\/10.1042\/ETLS20240008\/984281\/etls-2024-0008.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,2]],"date-time":"2026-02-02T14:05:47Z","timestamp":1770041147000},"score":1,"resource":{"primary":{"URL":"https:\/\/portlandpress.com\/emergtoplifesci\/article\/8\/4\/ETLS20240008\/237149\/Cytochrome-nanowires-under-electron-microscopy"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,2]]},"references-count":103,"journal-issue":{"issue":"4","published-online":{"date-parts":[[2026,2,2]]}},"URL":"https:\/\/doi.org\/10.1042\/etls20240008","relation":{},"ISSN":["2397-8554","2397-8562"],"issn-type":[{"value":"2397-8554","type":"print"},{"value":"2397-8562","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,2,2]]},"article-number":"ETLS20240008"}}