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{{Short description|Electrically conductive appendages produced by a number of bacteria}}
[[File:Proposal of catalyzing bio-voltage memristors.webp|thumb|350px|''[[Geobacter sulfurreducens]]'' and its nanowires]]
[[File:Proposal of catalyzing bio-voltage memristors.webp|thumb|350px|''[[Geobacter sulfurreducens]]'' and its nanowires]]
'''Bacterial nanowires''' (also known as microbial nanowires) are electrically [[conductive]] [[appendages]] produced by a number of [[bacteria]] most notably from (but not exclusive to) the ''[[Geobacter]]'' and ''[[Shewanella]]'' genera.<ref name=Reguera>{{cite journal | vauthors = Reguera G, McCarthy KD, Mehta T, Nicoll JS, Tuominen MT, Lovley DR | title = Extracellular electron transfer via microbial nanowires | journal = Nature | volume = 435 | issue = 7045 | pages = 1098–101 | date = June 2005 | pmid = 15973408 | doi = 10.1038/nature03661 }}</ref><ref name="Gorby2006">{{cite journal | vauthors = Gorby YA, Yanina S, McLean JS, Rosso KM, Moyles D, Dohnalkova A, Beveridge TJ, Chang IS, Kim BH, Kim KS, Culley DE, Reed SB, Romine MF, Saffarini DA, Hill EA, Shi L, Elias DA, Kennedy DW, Pinchuk G, Watanabe K, Ishii S, Logan B, Nealson KH, Fredrickson JK | display-authors = 6 | title = Electrically conductive bacterial nanowires produced by Shewanella oneidensis strain MR-1 and other microorganisms | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 103 | issue = 30 | pages = 11358–63 | date = July 2006 | pmid = 16849424 | pmc = 1544091 | doi = 10.1073/pnas.0604517103 | name-list-style = amp }}</ref> Conductive nanowires have also been confirmed in the oxygenic cyanobacterium ''Synechocystis'' PCC6803 and a thermophilic, methanogenic coculture consisting of ''Pelotomaculum thermopropionicum'' and ''Methanothermobacter thermoautotrophicus''.<ref name="Gorby2006" />
'''Bacterial nanowires''' (also known as microbial nanowires) are electrically [[conductive]] [[appendages]] produced by a number of [[bacteria]] most notably from the ''[[Geobacter]]'' and ''[[Shewanella]]'' genera.<ref name="Reguera 2005">{{cite journal | vauthors = Reguera G, McCarthy KD, Mehta T, Nicoll JS, Tuominen MT, Lovley DR | title = Extracellular electron transfer via microbial nanowires | journal = Nature | volume = 435 | issue = 7045 | pages = 1098–101 | date = June 2005 | pmid = 15973408 | doi = 10.1038/nature03661 | bibcode = 2005Natur.435.1098R | s2cid = 4425287 }}</ref><ref name="Gorby2006">{{cite journal | vauthors = Gorby YA, Yanina S, McLean JS, Rosso KM, Moyles D, Dohnalkova A, Beveridge TJ, Chang IS, Kim BH, Kim KS, Culley DE, Reed SB, Romine MF, Saffarini DA, Hill EA, Shi L, Elias DA, Kennedy DW, Pinchuk G, Watanabe K, Ishii S, Logan B, Nealson KH, Fredrickson JK | display-authors = 6 | title = Electrically conductive bacterial nanowires produced by Shewanella oneidensis strain MR-1 and other microorganisms | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 103 | issue = 30 | pages = 11358–63 | date = July 2006 | pmid = 16849424 | pmc = 1544091 | doi = 10.1073/pnas.0604517103 | bibcode = 2006PNAS..10311358G | name-list-style = amp | doi-access = free }}</ref> Conductive nanowires have also been confirmed in the oxygenic [[Cyanobacteria|cyanobacterium]] [[Synechocystis sp. PCC 6803|''Synechocystis'' PCC6803]] and a [[Thermophile|thermophilic]], [[Methanogenesis|methanogenic]] coculture consisting of ''[[Pelotomaculum thermopropionicum]]'' and ''[[Methanothermobacter|Methanothermobacter thermoautotrophicus]]''.<ref name="Gorby2006" /> From physiological and functional perspectives, bacterial nanowires are diverse.<ref name=":2" /><ref name=":0" /><ref name=":3" /> The precise role microbial nanowires play in their biological systems has not been fully realized, but several proposed functions exist.<ref name=":2" /> Outside of a naturally occurring environment, bacterial nanowires have shown potential to be useful in several fields, notably the [[bioenergy]] and [[bioremediation]] industries.<ref name=":4" /><ref name="Sure1">{{cite journal|vauthors=Sure S, Ackland ML, Torriero AA, Adholeya A, Kochar M|date=December 2016|title=Microbial nanowires: an electrifying tale|journal=Microbiology|volume=162|issue=12|pages=2017–2028|doi=10.1099/mic.0.000382|pmid=27902405|doi-access=free}}</ref>


==Physiology==
==Physiology==
''[[Geobacter]]'' nanowires were originally thought to be modified pili, which are used to establish connections to terminal electron receptors. Further research has shown that ''Geobacter'' nanowires are composed of stacked [[cytochromes]], namely [[OmcS oxidoreductase|OmcS]] and OmcZ.<ref>{{cite journal | vauthors = Yalcin SE, O'Brien JP, Gu Y, Reiss K, Yi SM, Jain R, Srikanth V, Dahl PJ, Huynh W, Vu D, Acharya A, Chaudhuri S, Varga T, Batista VS, Malvankar NS | display-authors = 6 | title = Electric field stimulates production of highly conductive microbial OmcZ nanowires | journal = Nature Chemical Biology | volume = 16 | issue = 10 | pages = 1136–1142 | date = October 2020 | pmid = 32807967 | pmc = 7502555 | doi = 10.1038/s41589-020-0623-9 }}</ref> These stacked cytochrome nanowires form a seamless array of [[heme]]s which stabilize the nanowire via pi-stacking and provide a path for electron transport.<ref>{{cite journal | vauthors = Wang F, Gu Y, O'Brien JP, Yi SM, Yalcin SE, Srikanth V, Shen C, Vu D, Ing NL, Hochbaum AI, Egelman EH, Malvankar NS | display-authors = 6 | title = Structure of Microbial Nanowires Reveals Stacked Hemes that Transport Electrons over Micrometers | journal = Cell | volume = 177 | issue = 2 | pages = 361–369.e10 | date = April 2019 | pmid = 30951668 | doi = 10.1016/j.cell.2019.03.029 | doi-access = free }}</ref> Species of the genus ''[[Geobacter]]'' use nanowires to transfer electrons to extracellular electron acceptors (such as Fe(III) oxides).<ref name="Reguera 2005">{{cite journal | vauthors = Reguera G, McCarthy KD, Mehta T, Nicoll JS, Tuominen MT, Lovley DR | title = Extracellular electron transfer via microbial nanowires | journal = Nature | volume = 435 | issue = 7045 | pages = 1098–101 | date = June 2005 | pmid = 15973408 | doi = 10.1038/nature03661 }}</ref> This function was discovered through the examination of mutants, whose nanowires could attach to the iron, but would not reduce it.<ref name="Reguera 2005"/>
''[[Geobacter]]'' nanowires were originally thought to be modified [[Pilus|pili]], which are used to establish connections to [[Electron acceptor|terminal electron acceptors]] during some types of [[anaerobic respiration]]. Further research has shown that ''Geobacter'' nanowires are composed of stacked [[cytochromes]], namely [[OmcS oxidoreductase|OmcS]] and OmcZ. Despite being physiologically distinct from pili, bacterial nanowires are often described as pili anyway due to the initial misconception upon their discovery.<ref name=":3">{{cite journal | vauthors = Yalcin SE, O'Brien JP, Gu Y, Reiss K, Yi SM, Jain R, Srikanth V, Dahl PJ, Huynh W, Vu D, Acharya A, Chaudhuri S, Varga T, Batista VS, Malvankar NS | display-authors = 6 | title = Electric field stimulates production of highly conductive microbial OmcZ nanowires | journal = Nature Chemical Biology | volume = 16 | issue = 10 | pages = 1136–1142 | date = October 2020 | pmid = 32807967 | pmc = 7502555 | doi = 10.1038/s41589-020-0623-9 }}</ref> These stacked cytochrome nanowires form a seamless array of [[heme]]s which stabilize the nanowire via [[Pi-Stacking (chemistry)|pi-stacking]] and provide a path for [[Electron transport chain|electron transport]].<ref>{{cite journal | vauthors = Wang F, Gu Y, O'Brien JP, Yi SM, Yalcin SE, Srikanth V, Shen C, Vu D, Ing NL, Hochbaum AI, Egelman EH, Malvankar NS | display-authors = 6 | title = Structure of Microbial Nanowires Reveals Stacked Hemes that Transport Electrons over Micrometers | journal = Cell | volume = 177 | issue = 2 | pages = 361–369.e10 | date = April 2019 | pmid = 30951668 | doi = 10.1016/j.cell.2019.03.029 | pmc = 6720112 | doi-access = free }}</ref> Species of the genus ''[[Geobacter]]'' use nanowires to transfer electrons to extracellular electron acceptors (such as Fe(III) oxides).<ref name="Reguera 2005"/> This function was discovered through the examination of mutants, whose nanowires could attach to the iron, but would not reduce it.<ref name="Reguera 2005"/>


However, ''[[Shewanella]]'' nanowires are not pili, but extensions of the outer membrane that contain the decaheme outer membrane cytochromes MtrC and OmcA.<ref>{{cite journal | vauthors = Pirbadian S, Barchinger SE, Leung KM, Byun HS, Jangir Y, Bouhenni RA, Reed SB, Romine MF, Saffarini DA, Shi L, Gorby YA, Golbeck JH, El-Naggar MY | display-authors = 6 | title = Shewanella oneidensis MR-1 nanowires are outer membrane and periplasmic extensions of the extracellular electron transport components | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 111 | issue = 35 | pages = 12883–8 | date = September 2014 | pmid = 25143589 | pmc = 4156777 | doi = 10.1073/pnas.1410551111 }}</ref> The reported presence of outer membrane cytochromes, and lack of conductivity in nanowires from the MtrC and OmcA-deficient mutant<ref>{{cite journal | vauthors = El-Naggar MY, Wanger G, Leung KM, Yuzvinsky TD, Southam G, Yang J, Lau WM, Nealson KH, Gorby YA | display-authors = 6 | title = Electrical transport along bacterial nanowires from Shewanella oneidensis MR-1 | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 107 | issue = 42 | pages = 18127–31 | date = October 2010 | pmid = 20937892 | pmc = 2964190 | doi = 10.1073/pnas.1004880107 }}</ref> directly support the proposed multistep hopping mechanism for electron transport through ''[[Shewanella]]'' nanowires.<ref>{{cite journal | vauthors = Pirbadian S, El-Naggar MY | title = Multistep hopping and extracellular charge transfer in microbial redox chains | journal = Physical Chemistry Chemical Physics | volume = 14 | issue = 40 | pages = 13802–8 | date = October 2012 | pmid = 22797729 | doi = 10.1039/C2CP41185G }}</ref><ref>{{cite journal | vauthors = Polizzi NF, Skourtis SS, Beratan DN | title = Physical constraints on charge transport through bacterial nanowires | journal = Faraday Discussions | volume = 155 | pages = 43–62; discussion 103-14 | year = 2012 | pmid = 22470966 | pmc = 3392031 | doi = 10.1039/C1FD00098E }}</ref><ref>{{cite journal | vauthors = Strycharz-Glaven SM, Snider RM, Guiseppi-Elie A, Tender LM | year = 2011 | title = On the electrical conductivity of microbial nanowires and biofilms | journal = Energy Environ Sci | volume = 4 | issue = 11| pages = 4366–4379 | doi = 10.1039/C1EE01753E }}</ref>
''[[Shewanella]]'' nanowires are also not technically pili, but extensions of the outer membrane that contain the decaheme [[Bacterial outer membrane|outer membrane]] [[cytochrome]]s MtrC and OmcA.<ref name=":0">{{cite journal | vauthors = Pirbadian S, Barchinger SE, Leung KM, Byun HS, Jangir Y, Bouhenni RA, Reed SB, Romine MF, Saffarini DA, Shi L, Gorby YA, Golbeck JH, El-Naggar MY | display-authors = 6 | title = Shewanella oneidensis MR-1 nanowires are outer membrane and periplasmic extensions of the extracellular electron transport components | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 111 | issue = 35 | pages = 12883–8 | date = September 2014 | pmid = 25143589 | pmc = 4156777 | doi = 10.1073/pnas.1410551111 | bibcode = 2014PNAS..11112883P | doi-access = free }}</ref> The reported presence of outer membrane cytochromes, and lack of conductivity in nanowires from the MtrC and OmcA-deficient mutant<ref>{{cite journal | vauthors = El-Naggar MY, Wanger G, Leung KM, Yuzvinsky TD, Southam G, Yang J, Lau WM, Nealson KH, Gorby YA | display-authors = 6 | title = Electrical transport along bacterial nanowires from Shewanella oneidensis MR-1 | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 107 | issue = 42 | pages = 18127–31 | date = October 2010 | pmid = 20937892 | pmc = 2964190 | doi = 10.1073/pnas.1004880107 | bibcode = 2010PNAS..10718127E | doi-access = free }}</ref> directly support the proposed multistep hopping mechanism for electron transport through ''[[Shewanella]]'' nanowires.<ref>{{cite journal | vauthors = Pirbadian S, El-Naggar MY | title = Multistep hopping and extracellular charge transfer in microbial redox chains | journal = Physical Chemistry Chemical Physics | volume = 14 | issue = 40 | pages = 13802–8 | date = October 2012 | pmid = 22797729 | doi = 10.1039/C2CP41185G | bibcode = 2012PCCP...1413802P }}</ref><ref>{{cite journal | vauthors = Polizzi NF, Skourtis SS, Beratan DN | title = Physical constraints on charge transport through bacterial nanowires | journal = Faraday Discussions | volume = 155 | pages = 43–62; discussion 103–14 | year = 2012 | pmid = 22470966 | pmc = 3392031 | doi = 10.1039/C1FD00098E | bibcode = 2012FaDi..155...43P }}</ref><ref>{{cite journal | vauthors = Strycharz-Glaven SM, Snider RM, Guiseppi-Elie A, Tender LM | year = 2011 | title = On the electrical conductivity of microbial nanowires and biofilms | journal = Energy Environ Sci | volume = 4 | issue = 11| pages = 4366–4379 | doi = 10.1039/C1EE01753E }}</ref>


Additionally, nanowires can facilitate long-range electron transfer across thick ''[[biofilm]]'' layers.<ref>{{cite journal | vauthors = Reguera G, Nevin KP, Nicoll JS, Covalla SF, Woodard TL, Lovley DR | title = Biofilm and nanowire production leads to increased current in Geobacter sulfurreducens fuel cells | journal = Applied and Environmental Microbiology | volume = 72 | issue = 11 | pages = 7345–8 | date = November 2006 | pmid = 16936064 | pmc = 1636155 | doi = 10.1128/aem.01444-06 }}</ref> By connecting to other cells above them, nanowires allow bacteria located in anoxic conditions to still use oxygen as their terminal electron acceptor. For example, organisms in the genus ''[[Shewanella]]'' have been observed to form electrically conductive nanowires in response to electron-acceptor limitation.<ref name="Gorby2006" />
Additionally, nanowires can facilitate long-range electron transfer across thick ''[[biofilm]]'' layers.<ref name=":4">{{cite journal | vauthors = Reguera G, Nevin KP, Nicoll JS, Covalla SF, Woodard TL, Lovley DR | title = Biofilm and nanowire production leads to increased current in Geobacter sulfurreducens fuel cells | journal = Applied and Environmental Microbiology | volume = 72 | issue = 11 | pages = 7345–8 | date = November 2006 | pmid = 16936064 | pmc = 1636155 | doi = 10.1128/aem.01444-06 | bibcode = 2006ApEnM..72.7345R }}</ref> By connecting to other cells around them, nanowires allow bacteria located in [[Hypoxia (environmental)|anoxic]] conditions to still use oxygen as their terminal electron acceptor. For example, organisms in the genus ''[[Shewanella]]'' have been observed to form electrically conductive nanowires in response to electron-acceptor limitation.<ref name="Gorby2006" />


==History==
==History==
The concept of electromicrobiology has been around since the early 1900s when a series of discoveries found cells capable of producing electricity. It was demonstrated for the first time in 1911 by [[British Mycological Society|Michael Cressé Potter]] that cells could convert chemical energy to electrical energy.<ref name=":2" /><ref>{{Cite journal| vauthors = Potter MC, Waller AD |date=1911-09-14|title=Electrical effects accompanying the decomposition of organic compounds|url=https://royalsocietypublishing.org/doi/10.1098/rspb.1911.0073|journal=Proceedings of the Royal Society of London. Series B, Containing Papers of a Biological Character|volume=84|issue=571|pages=260–276|doi=10.1098/rspb.1911.0073|doi-access=free}}</ref> It wasn't until 1988 that extracellular electron transport (EET) was observed for the first time with the independent discoveries of ''Geobacter'' and ''Shewanella'' bacteria and their respective nanowires. Since their discoveries, other nanowire containing microbes have been identified, but they remain the most intensively studied.<ref name=":2" /><ref>{{cite journal | vauthors = Myers CR, Nealson KH | title = Bacterial manganese reduction and growth with manganese oxide as the sole electron acceptor | journal = Science | volume = 240 | issue = 4857 | pages = 1319–21 | date = June 1988 | pmid = 17815852 | doi = 10.1126/science.240.4857.1319 | bibcode = 1988Sci...240.1319M | s2cid = 9662366 }}</ref><ref>{{cite journal | vauthors = Lovley DR, Phillips EJ | title = Novel mode of microbial energy metabolism: organic carbon oxidation coupled to dissimilatory reduction of iron or manganese | journal = Applied and Environmental Microbiology | volume = 54 | issue = 6 | pages = 1472–80 | date = June 1988 | pmid = 16347658 | doi = 10.1128/aem.54.6.1472-1480.1988 | pmc = 202682 | bibcode = 1988ApEnM..54.1472L }}</ref> In 1998, EET was observed in a [[microbial fuel cell]] setting for the first time using ''Shewanella'' bacteria to reduce an Fe(III) electrode.<ref name=":2" /><ref>{{cite journal| vauthors = Kim B |date=1999|title=Dynamic effects of learning capabilities and profit structures on the innovation competition |journal=Optimal Control Applications and Methods|language=en|volume=20|issue=3|pages=127–144|doi=10.1002/(SICI)1099-1514(199905/06)20:3<127::AID-OCA650>3.0.CO;2-I|issn=1099-1514}}</ref> In 2010, bacterial nanowires were shown to have facilitated the flow of electricity into [[Sporomusa ovata|''Sporomusa'']] bacteria. This was the first observed instance of EET used to draw electrons from the environment into a cell.<ref name=":2" /><ref name="Microbial electrosynthesis - revisi">{{cite journal | vauthors = Rabaey K, Rozendal RA | title = Microbial electrosynthesis - revisiting the electrical route for microbial production | journal = Nature Reviews. Microbiology | volume = 8 | issue = 10 | pages = 706–16 | date = October 2010 | pmid = 20844557 | doi = 10.1038/nrmicro2422 | s2cid = 11417035 }}</ref> Research persists to date to explore the mechanisms, implications, and potential applications of nanowires and the biological systems they are a part of.
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==Implications and potential applications==
==Implications and potential applications==


=== Biological implications ===
===Biological implications===
Biologically, the extent of the implications brought on by the existence of bacterial nanowires is not fully realized. Predictions exist in the microbiological field, but few definitive conclusions have been made. It has been speculated nanowires may function as conduits for electron transport between different members of a microbial community. This has potential to allow for regulatory feedback or other communication between members of the same or even different microbial species.<ref>{{cite journal | vauthors = Rabaey K, Rozendal RA | title = Microbial electrosynthesis - revisiting the electrical route for microbial production | journal = Nature Reviews. Microbiology | volume = 8 | issue = 10 | pages = 706–16 | date = October 2010 | pmid = 20844557 | doi = 10.1038/nrmicro2422 }}</ref><ref name=":1">{{cite journal | vauthors = Shi L, Dong H, Reguera G, Beyenal H, Lu A, Liu J, Yu HQ, Fredrickson JK | display-authors = 6 | title = Extracellular electron transfer mechanisms between microorganisms and minerals | journal = Nature Reviews. Microbiology | volume = 14 | issue = 10 | pages = 651–62 | date = October 2016 | pmid = 27573579 | doi = 10.1038/nrmicro.2016.93 }}</ref>  Microorganisms could potentially use nanowires to facilitate the use of extracellular metals as terminal electron acceptors in an [[electron transport chain]]. The high reduction potential of the metals receiving electrons could drive a considerable [[Adenosine triphosphate|ATP]] production.<ref name=":1" /><ref name=":2">{{cite journal | vauthors = Nealson KH, Rowe AR | title = Electromicrobiology: realities, grand challenges, goals and predictions | journal = Microbial Biotechnology | volume = 9 | issue = 5 | pages = 595–600 | date = September 2016 | pmid = 27506517 | pmc = 4993177 | doi = 10.1111/1751-7915.12400 }}</ref> Some organisms are capable of both expelling and taking in electrons through nanowires.<ref name=":2" /> Those species could potentially oxidize extracellular metals by using them as an electron or energy source to facilitate energy consuming cellular processes.<ref name=":1" /> Microbes also could potentially be using nanowires to temporarily store electrons on metals. Building up an electron concentration on a metal anode of sorts would create a battery of sorts that the cells could later use to fuel metabolic activity.<ref name=":1" /> While these potential implications provide a reasonable hypothesis towards the role of the bacterial nanowire in a biological system, more research is needed to fully understand the extent of how cellular species benefit from nanowire use.<ref name=":2" />
Microorganisms have shown to use nanowires to facilitate the use of extracellular metals as terminal electron acceptors in an [[electron transport chain]]. The high reduction potential of the metals receiving electrons is capable of driving a considerable [[Adenosine triphosphate|ATP]] production.<ref name=":1" /><ref name=":2">{{cite journal | vauthors = Nealson KH, Rowe AR | title = Electromicrobiology: realities, grand challenges, goals and predictions | journal = Microbial Biotechnology | volume = 9 | issue = 5 | pages = 595–600 | date = September 2016 | pmid = 27506517 | pmc = 4993177 | doi = 10.1111/1751-7915.12400 }}</ref> Aside from that, the extent of the implications brought on by the existence of bacterial nanowires is not fully realized. It has been speculated nanowires may function as conduits for electron transport between different members of a microbial community. This has potential to allow for regulatory feedback or other communication between members of the same or even different microbial species.<ref name="Microbial electrosynthesis - revisi"/><ref name=":1">{{cite journal | vauthors = Shi L, Dong H, Reguera G, Beyenal H, Lu A, Liu J, Yu HQ, Fredrickson JK | display-authors = 6 | title = Extracellular electron transfer mechanisms between microorganisms and minerals | journal = Nature Reviews. Microbiology | volume = 14 | issue = 10 | pages = 651–62 | date = October 2016 | pmid = 27573579 | doi = 10.1038/nrmicro.2016.93 | s2cid = 20626915 }}</ref> Some organisms are capable of both expelling and taking in electrons through nanowires.<ref name=":2" /> Those species would likely be able to oxidize extracellular metals by using them as an electron or energy source to facilitate energy consuming cellular processes.<ref name=":1" /> Microbes also could potentially use nanowires to temporarily store electrons on metals. Building up an electron concentration on a metal [[anode]] would create a battery of sorts that the cells could later use to fuel [[Metabolism|metabolic activity]].<ref name=":1" /> While these potential implications provide a reasonable hypothesis towards the role of the bacterial nanowire in a biological system, more research is needed to fully understand the extent of how cellular species benefit from nanowire use.<ref name=":2" />


=== Bioenergy applications in microbial fuel cells ===
===Bioenergy applications in microbial fuel cells===
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In [[microbial fuel cell]]s (MFCs), bacterial nanowires generate electricity via extracellular electron transport to the MFC's anode.<ref>Kodesia, A.; Ghosh, M.; Chatterjee, A. (September 5, 2017). [[hdl:10266/4826|"Development of Biofilm Nanowires and Electrode for Efficient Microbial Fuel Cells (MFCs)"]]. ''Thapar University Digital Repository (TuDR)''.</ref> Nanowire networks have been shown to enhance the electricity output of MFCs with efficient and long-range conductivity. In particular, bacterial nanowires of ''Geobacter sulfurreducens'' possess metallic-like conductivity, producing electricity at levels comparable to those of synthetic metallic nanostructures.<ref name="pmid21822253">{{cite journal | vauthors = Malvankar NS, Vargas M, Nevin KP, Franks AE, Leang C, Kim BC, Inoue K, Mester T, Covalla SF, Johnson JP, Rotello VM, Tuominen MT, Lovley DR | title = Tunable metallic-like conductivity in microbial nanowire networks | journal = Nature Nanotechnology | volume = 6 | issue = 9 | pages = 573–9 | date = August 2011 | pmid = 21822253 | doi = 10.1038/nnano.2011.119 | bibcode = 2011NatNa...6..573M }}</ref> When bacterial strains are [[Genetically modified organism|genetically manipulated]] to boost nanowire formation, higher electricity yields are generally observed.<ref name="Malvankar2012" /> Coating the nanowires with [[Oxide|metal oxides]] also further promotes electrical conductivity.<ref>{{cite journal | vauthors = Maruthupandy M, Anand M, Maduraiveeran G, Beevi AS, Priya RJ | title = Fabrication of CuO nanoparticles coated bacterial nanowire film for a high-performance electrochemical conductivity. | journal = Journal of Materials Science | date = September 2017 | volume = 52 | issue = 18 | pages = 10766–78 | doi = 10.1007/s10853-017-1248-6 | bibcode = 2017JMatS..5210766M | s2cid = 103105219 }}</ref> Additionally, these nanowires can transport electrons up to centimeter-scale distances.<ref name="Malvankar2012" /> Long-range electron transfer via microbial nanowire networks allows viable cells that are not in direct contact with an anode to contribute to electron flow.<ref name=":4"/>


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To date, the currency produced by bacterial nanowires is very low. Across a biofilm 7 micrometers thick, a current density of around 17 microamperes per square centimeter and a voltage of around 0.5 volts was reported.<ref name="Xiaomeng et al., 2020">{{cite journal | vauthors = Liu X, Gao H, Ward JE, Liu X, Yin B, Fu T, Chen J, Lovley DR, Yao J | display-authors = 6 | title = Power generation from ambient humidity using protein nanowires | journal = Nature | volume = 578 | issue = 7796 | pages = 550–554 | date = February 2020 | pmid = 32066937 | doi = 10.1038/s41586-020-2010-9 | bibcode = 2020Natur.578..550L | doi-access = free }}</ref>
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In [[microbial fuel cell]]s (MFCs), bacterial nanowires generate electricity via extracellular electron transport to the MFC's anode.<ref>Kodesia, A.; Ghosh, M.; Chatterjee, A. (September 5, 2017). [[hdl:10266/4826|"Development of Biofilm Nanowires and Electrode for Efficient Microbial Fuel Cells (MFCs)"]]. ''Thapar University Digital Repository (TuDR)''.</ref> Nanowire networks have been shown to enhance the electricity output of MFCs with efficient and long-range conductivity. In particular, pili of ''Geobacter sulfurreducens'' possess metallic-like conductivity, producing electricity at levels comparable to those of synthetic metallic nanostructures.<ref>Malvankar, N.S.; Vargas, M.; Nevin, K.P.; Franks, A.E.; Leang, C.; Kim, B.C.; Inoue, K. (August 7, 2011). [https://www.nature.com/articles/nnano.2011.119 "Tunable metallic-like conductivity in microbialnanowire networks"]. ''Nature Nanotechnology''. '''6''': 573–579. [[Digital object identifier|doi]]:[https://doi.org/10.1038%2FNNANO.2011.119 10.1038/NNANO.2011.119].</ref> When bacterial strains are genetically manipulated to boost nanowire formation, higher electricity yields are generally observed.<ref name=":0" /> Coating the nanowires with metal oxides also further promotes electrical conductivity.<ref>Maruthupandy, M.; Anand, M.; Maduraiveeran, G. (June 5, 2017). [https://link.springer.com/article/10.1007%2Fs10853-017-1248-6 "Fabrication of CuO nanoparticles coated bacterial nanowire film for a high-performance electrochemical conductivity"]. ''J Mater Sci''. '''52''': 10766–10778. [[Digital object identifier|doi]]:[https://doi.org/10.1007%2Fs10853-017-1248-6 10.1007/s10853-017-1248-6].</ref> Additionally, these nanowires can transport electrons up to centimeter-scale distances.<ref name=":0">Malvankar, N.S.; Lovley, D.R. (May 21, 2012). [http://onlinelibrary.wiley.com/doi/10.1002/cssc.201100733/abstract;jsessionid=81A61046E8B6898C5FFC37DBC5B1EC7A.f01t03 "Microbial Nanowires: A New Paradigm for Biological Electron Transfer and Bioelectronics"]. ''ChemSusChem''. '''5''' (6): 1039–1046. [[Digital object identifier|doi]]:[https://doi.org/10.1002%2Fcssc.201100733 10.1002/cssc.201100733] – via Wiley Online Library.</ref> Long-range electron transfer via pili networks allows viable cells that are not in direct contact with an anode to contribute to electron flow.<ref>{{cite journal | vauthors = Reguera G, Nevin KP, Nicoll JS, Covalla SF, Woodard TL, Lovley DR | title = Biofilm and nanowire production leads to increased current in Geobacter sulfurreducens fuel cells | journal = Applied and Environmental Microbiology | volume = 72 | issue = 11 | pages = 7345–8 | date = November 2006 | pmid = 16936064 | pmc = 1636155 | doi = 10.1128/aem.01444-06 }}</ref> Thus, increased current production in MFCs is observed in thicker biofilms.

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The currency produced by bacterial nanowires very low. Current density of around 17 microamperes per square centimeter, voltage of around 0.5 volts across a 7-micrometre-thick film.<ref name="Xiaomeng et al., 2020">{{cite journal | vauthors = Liu X, Gao H, Ward JE, Liu X, Yin B, Fu T, Chen J, Lovley DR, Yao J | display-authors = 6 | title = Power generation from ambient humidity using protein nanowires | journal = Nature | volume = 578 | issue = 7796 | pages = 550–554 | date = February 2020 | pmid = 32066937 | doi = 10.1038/s41586-020-2010-9 | doi-access = free }}</ref>


===Other significant applications===
===Other significant applications===
⚫
Microbial nanowires of ''Shewanella'' and ''Geobacter'' have been shown to aid in [[bioremediation of radioactive waste|bioremediation of uranium contaminated groundwater]].<ref>{{cite journal | vauthors = Jiang S, Kim MG, Kim SJ, Jung HS, Lee SW, Noh DY, Sadowsky MJ, Hur HG | display-authors = 6 | title = Bacterial formation of extracellular U(VI) nanowires | journal = Chemical Communications | volume = 47 | issue = 28 | pages = 8076–8 | date = July 2011 | pmid = 21681306 | doi = 10.1039/C1CC12554K }}</ref> To demonstrate this, scientists compared and observed the concentration of uranium removed by piliated and nonpiliated strains of ''Geobacter.'' Through a series of controlled experiments, they were able to deduce that nanowire present strains were more effective at the [[Biomineralization|mineralization]] of uranium as compared to nanowire absent mutants.<ref name="Cologgi">{{cite journal | vauthors = Cologgi DL, Lampa-Pastirk S, Speers AM, Kelly SD, Reguera G | title = Extracellular reduction of uranium via Geobacter conductive pili as a protective cellular mechanism | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 108 | issue = 37 | pages = 15248–52 | date = September 2011 | pmid = 21896750 | pmc = 3174638 | doi = 10.1073/pnas.1108616108 | bibcode = 2011PNAS..10815248C | doi-access = free }}</ref>


⚫
Further significant application of bacterial nanowires can be seen in the bioelectronics industry.<ref name="Sure1"/> With sustainable resources in mind, scientists have proposed the future use of biofilms of ''Geobacter'' as a platform for functional under water [[transistor]]s and [[supercapacitor]]s, capable of self-renewing energy.<ref name="Malvankar2012">{{cite journal | vauthors = Malvankar NS, Lovley DR | title = Microbial nanowires: a new paradigm for biological electron transfer and bioelectronics | journal = ChemSusChem | volume = 5 | issue = 6 | pages = 1039–46 | date = June 2012 | pmid = 22614997 | doi = 10.1002/cssc.201100733 }}</ref>
Bacterial nanowires have been shown to have significant potential applications in the fields of bioenergy and bioremediation.<ref name="Sure1">{{cite journal | vauthors = Sure S, Ackland ML, Torriero AA, Adholeya A, Kochar M | title = Microbial nanowires: an electrifying tale | journal = Microbiology | volume = 162 | issue = 12 | pages = 2017–2028 | date = December 2016 | pmid = 27902405 | doi = 10.1099/mic.0.000382 | doi-access = free }}</ref> Electron transfer between the pili of ''Geobacter'', a dissimilatory metal-reducing bacterium, generates conductivity that drives the conversion of organic compounds to electricity in microbial fuel cells.<ref name="Malvankar">{{cite journal | vauthors = Malvankar NS, Lovley DR | title = Microbial nanowires for bioenergy applications | journal = Current Opinion in Biotechnology | volume = 27 | pages = 88–95 | date = June 2014 | pmid = 24863901 | doi = 10.1016/j.copbio.2013.12.003 }}</ref> Biofilms produced by ''Geobacter'' colonies contribute greatly to the overall production of bioenergy. They allow the transfer of electrons via conductive pili over a greater distance from the anode.<ref name="Sure1"/> In fact, Bioenergy output can be further enhanced by inducing the expression of additional nanowire genes. ''Geobacter'' strains with heightened expression of conductive pili have been shown to produce more conductive biofilms, thus increasing overall electricity output.<ref name="Malvankar"/>

⚫
Microbial nanowires of ''Shewanella'' and ''Geobacter'' have also been shown to aid in bioremediation of uranium contaminated groundwater.<ref>{{cite journal | vauthors = Jiang S, Kim MG, Kim SJ, Jung HS, Lee SW, Noh DY, Sadowsky MJ, Hur HG | display-authors = 6 | title = Bacterial formation of extracellular U(VI) nanowires | journal = Chemical Communications | volume = 47 | issue = 28 | pages = 8076–8 | date = July 2011 | pmid = 21681306 | doi = 10.1039/C1CC12554K }}</ref> To demonstrate this, scientists compared and observed the concentration of uranium removed by piliated and nonpiliated strains of ''Geobacter.'' Through a series of controlled experiments, they were able to deduce that nanowire present strains were more effective at the mineralization of uranium as compared to nanowire absent mutants.<ref name="Cologgi">{{cite journal | vauthors = Cologgi DL, Lampa-Pastirk S, Speers AM, Kelly SD, Reguera G | title = Extracellular reduction of uranium via Geobacter conductive pili as a protective cellular mechanism | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 108 | issue = 37 | pages = 15248–52 | date = September 2011 | pmid = 21896750 | pmc = 3174638 | doi = 10.1073/pnas.1108616108 }}</ref>

⚫
Further application significance of bacterial nanowires include bioelectronics.<ref name="Sure1"/> With sustainable resources in mind, scientists have proposed the future use of biofilms of ''Geobacter'' as a platform for functional under water transistors and supercapacitors, capable of self-renewing energy.<ref name="Malvankar2012">{{cite journal | vauthors = Malvankar NS, Lovley DR | title = Microbial nanowires: a new paradigm for biological electron transfer and bioelectronics | journal = ChemSusChem | volume = 5 | issue = 6 | pages = 1039–46 | date = June 2012 | pmid = 22614997 | doi = 10.1002/cssc.201100733 }}</ref>


On 20 April 2020 researchers demonstrated a diffusive [[memristor]] fabricated from protein nanowires of the bacterium ''[[Geobacter sulfurreducens]]'' which functions at substantially lower voltages than the ones previously described and may allow the construction of [[artificial neuron]]s which function at voltages of biological [[action potential]]s. Bacterial nanowires vary from traditionally utilized silicon nanowires by showing an increased degree of [[biocompatibility]]. More research is needed, but the memristors may eventually be used to directly process [[Biosensor|biosensing signals]], for [[neuromorphic computing]] and/or [[Brain-computer interface|direct communication with biological neurons]].<ref>{{cite journal | vauthors = Fu T, Liu X, Gao H, Ward JE, Liu X, Yin B, Wang Z, Zhuo Y, Walker DJ, Joshua Yang J, Chen J, Lovley DR, Yao J | display-authors = 6 | title = Bioinspired bio-voltage memristors | journal = Nature Communications | volume = 11 | issue = 1 | pages = 1861 | date = April 2020 | pmid = 32313096 | doi = 10.1038/s41467-020-15759-y | doi-access = free }}</ref><ref>{{Cite web|title=Researchers Unveil Electronics that Mimic the Human Brain in Efficient, Biological Learning|url=https://www.umass.edu/newsoffice/article/researchers-unveil-electronics-mimic-human|access-date=2021-04-20|website=Office of News & Media Relations {{!}} UMass Amherst|language=en}}</ref>
On 20 April 2020, researchers demonstrated a diffusive [[memristor]] fabricated from protein nanowires of the bacterium ''[[Geobacter sulfurreducens]]'' which functions at substantially lower voltages than the ones previously described and may allow the construction of [[artificial neuron]]s which function at voltages of biological [[action potential]]s. Bacterial nanowires vary from traditionally utilized silicon nanowires by showing an increased degree of [[biocompatibility]]. More research is needed, but the memristors may eventually be used to directly process [[biosensor|biosensing signals]], for [[neuromorphic computing]] and/or [[Brain–computer interface|direct communication with biological neurons]].<ref>{{cite journal | vauthors = Fu T, Liu X, Gao H, Ward JE, Liu X, Yin B, Wang Z, Zhuo Y, Walker DJ, Joshua Yang J, Chen J, Lovley DR, Yao J | display-authors = 6 | title = Bioinspired bio-voltage memristors | journal = Nature Communications | volume = 11 | issue = 1 | pages = 1861 | date = April 2020 | pmid = 32313096 | doi = 10.1038/s41467-020-15759-y | pmc = 7171104 | bibcode = 2020NatCo..11.1861F | doi-access = free }}</ref><ref>{{Cite web|title=Researchers Unveil Electronics that Mimic the Human Brain in Efficient, Biological Learning|url=https://www.umass.edu/newsoffice/article/researchers-unveil-electronics-mimic-human|access-date=2021-04-20|website=Office of News & Media Relations {{!}} UMass Amherst|language=en}}</ref>


== References ==
==References==
{{Reflist}}
{{Reflist}}


Latest revision as of 23:22, 2 January 2024

Geobacter sulfurreducens and its nanowires

Bacterial nanowires (also known as microbial nanowires) are electrically conductive appendages produced by a number of bacteria most notably from the Geobacter and Shewanella genera.[1][2] Conductive nanowires have also been confirmed in the oxygenic cyanobacterium Synechocystis PCC6803 and a thermophilic, methanogenic coculture consisting of Pelotomaculum thermopropionicum and Methanothermobacter thermoautotrophicus.[2] From physiological and functional perspectives, bacterial nanowires are diverse.[3][4][5] The precise role microbial nanowires play in their biological systems has not been fully realized, but several proposed functions exist.[3] Outside of a naturally occurring environment, bacterial nanowires have shown potential to be useful in several fields, notably the bioenergy and bioremediation industries.[6][7]

Physiology

[edit]

Geobacter nanowires were originally thought to be modified pili, which are used to establish connections to terminal electron acceptors during some types of anaerobic respiration. Further research has shown that Geobacter nanowires are composed of stacked cytochromes, namely OmcS and OmcZ. Despite being physiologically distinct from pili, bacterial nanowires are often described as pili anyway due to the initial misconception upon their discovery.[5] These stacked cytochrome nanowires form a seamless array of hemes which stabilize the nanowire via pi-stacking and provide a path for electron transport.[8] Species of the genus Geobacter use nanowires to transfer electrons to extracellular electron acceptors (such as Fe(III) oxides).[1] This function was discovered through the examination of mutants, whose nanowires could attach to the iron, but would not reduce it.[1]

Shewanella nanowires are also not technically pili, but extensions of the outer membrane that contain the decaheme outer membrane cytochromes MtrC and OmcA.[4] The reported presence of outer membrane cytochromes, and lack of conductivity in nanowires from the MtrC and OmcA-deficient mutant[9] directly support the proposed multistep hopping mechanism for electron transport through Shewanella nanowires.[10][11][12]

Additionally, nanowires can facilitate long-range electron transfer across thick biofilm layers.[6] By connecting to other cells around them, nanowires allow bacteria located in anoxic conditions to still use oxygen as their terminal electron acceptor. For example, organisms in the genus Shewanella have been observed to form electrically conductive nanowires in response to electron-acceptor limitation.[2]

History

[edit]

The concept of electromicrobiology has been around since the early 1900s when a series of discoveries found cells capable of producing electricity. It was demonstrated for the first time in 1911 by Michael Cressé Potter that cells could convert chemical energy to electrical energy.[3][13] It wasn't until 1988 that extracellular electron transport (EET) was observed for the first time with the independent discoveries of Geobacter and Shewanella bacteria and their respective nanowires. Since their discoveries, other nanowire containing microbes have been identified, but they remain the most intensively studied.[3][14][15] In 1998, EET was observed in a microbial fuel cell setting for the first time using Shewanella bacteria to reduce an Fe(III) electrode.[3][16] In 2010, bacterial nanowires were shown to have facilitated the flow of electricity into Sporomusa bacteria. This was the first observed instance of EET used to draw electrons from the environment into a cell.[3][17] Research persists to date to explore the mechanisms, implications, and potential applications of nanowires and the biological systems they are a part of.

Implications and potential applications

[edit]

Biological implications

[edit]

Microorganisms have shown to use nanowires to facilitate the use of extracellular metals as terminal electron acceptors in an electron transport chain. The high reduction potential of the metals receiving electrons is capable of driving a considerable ATP production.[18][3] Aside from that, the extent of the implications brought on by the existence of bacterial nanowires is not fully realized. It has been speculated nanowires may function as conduits for electron transport between different members of a microbial community. This has potential to allow for regulatory feedback or other communication between members of the same or even different microbial species.[17][18] Some organisms are capable of both expelling and taking in electrons through nanowires.[3] Those species would likely be able to oxidize extracellular metals by using them as an electron or energy source to facilitate energy consuming cellular processes.[18] Microbes also could potentially use nanowires to temporarily store electrons on metals. Building up an electron concentration on a metal anode would create a battery of sorts that the cells could later use to fuel metabolic activity.[18] While these potential implications provide a reasonable hypothesis towards the role of the bacterial nanowire in a biological system, more research is needed to fully understand the extent of how cellular species benefit from nanowire use.[3]

Bioenergy applications in microbial fuel cells

[edit]

In microbial fuel cells (MFCs), bacterial nanowires generate electricity via extracellular electron transport to the MFC's anode.[19] Nanowire networks have been shown to enhance the electricity output of MFCs with efficient and long-range conductivity. In particular, bacterial nanowires of Geobacter sulfurreducens possess metallic-like conductivity, producing electricity at levels comparable to those of synthetic metallic nanostructures.[20] When bacterial strains are genetically manipulated to boost nanowire formation, higher electricity yields are generally observed.[21] Coating the nanowires with metal oxides also further promotes electrical conductivity.[22] Additionally, these nanowires can transport electrons up to centimeter-scale distances.[21] Long-range electron transfer via microbial nanowire networks allows viable cells that are not in direct contact with an anode to contribute to electron flow.[6]

To date, the currency produced by bacterial nanowires is very low. Across a biofilm 7 micrometers thick, a current density of around 17 microamperes per square centimeter and a voltage of around 0.5 volts was reported.[23]

Other significant applications

[edit]

Microbial nanowires of Shewanella and Geobacter have been shown to aid in bioremediation of uranium contaminated groundwater.[24] To demonstrate this, scientists compared and observed the concentration of uranium removed by piliated and nonpiliated strains of Geobacter. Through a series of controlled experiments, they were able to deduce that nanowire present strains were more effective at the mineralization of uranium as compared to nanowire absent mutants.[25]

Further significant application of bacterial nanowires can be seen in the bioelectronics industry.[7] With sustainable resources in mind, scientists have proposed the future use of biofilms of Geobacter as a platform for functional under water transistors and supercapacitors, capable of self-renewing energy.[21]

On 20 April 2020, researchers demonstrated a diffusive memristor fabricated from protein nanowires of the bacterium Geobacter sulfurreducens which functions at substantially lower voltages than the ones previously described and may allow the construction of artificial neurons which function at voltages of biological action potentials. Bacterial nanowires vary from traditionally utilized silicon nanowires by showing an increased degree of biocompatibility. More research is needed, but the memristors may eventually be used to directly process biosensing signals, for neuromorphic computing and/or direct communication with biological neurons.[26][27]

References

[edit]
  1. ^ a b c Reguera G, McCarthy KD, Mehta T, Nicoll JS, Tuominen MT, Lovley DR (June 2005). "Extracellular electron transfer via microbial nanowires". Nature. 435 (7045): 1098–101. Bibcode:2005Natur.435.1098R. doi:10.1038/nature03661. PMID 15973408. S2CID 4425287.
  2. ^ a b c Gorby YA, Yanina S, McLean JS, Rosso KM, Moyles D, Dohnalkova A, et al. (July 2006). "Electrically conductive bacterial nanowires produced by Shewanella oneidensis strain MR-1 and other microorganisms". Proceedings of the National Academy of Sciences of the United States of America. 103 (30): 11358–63. Bibcode:2006PNAS..10311358G. doi:10.1073/pnas.0604517103. PMC 1544091. PMID 16849424.
  3. ^ a b c d e f g h i Nealson KH, Rowe AR (September 2016). "Electromicrobiology: realities, grand challenges, goals and predictions". Microbial Biotechnology. 9 (5): 595–600. doi:10.1111/1751-7915.12400. PMC 4993177. PMID 27506517.
  4. ^ a b Pirbadian S, Barchinger SE, Leung KM, Byun HS, Jangir Y, Bouhenni RA, et al. (September 2014). "Shewanella oneidensis MR-1 nanowires are outer membrane and periplasmic extensions of the extracellular electron transport components". Proceedings of the National Academy of Sciences of the United States of America. 111 (35): 12883–8. Bibcode:2014PNAS..11112883P. doi:10.1073/pnas.1410551111. PMC 4156777. PMID 25143589.
  5. ^ a b Yalcin SE, O'Brien JP, Gu Y, Reiss K, Yi SM, Jain R, et al. (October 2020). "Electric field stimulates production of highly conductive microbial OmcZ nanowires". Nature Chemical Biology. 16 (10): 1136–1142. doi:10.1038/s41589-020-0623-9. PMC 7502555. PMID 32807967.
  6. ^ a b c Reguera G, Nevin KP, Nicoll JS, Covalla SF, Woodard TL, Lovley DR (November 2006). "Biofilm and nanowire production leads to increased current in Geobacter sulfurreducens fuel cells". Applied and Environmental Microbiology. 72 (11): 7345–8. Bibcode:2006ApEnM..72.7345R. doi:10.1128/aem.01444-06. PMC 1636155. PMID 16936064.
  7. ^ a b Sure S, Ackland ML, Torriero AA, Adholeya A, Kochar M (December 2016). "Microbial nanowires: an electrifying tale". Microbiology. 162 (12): 2017–2028. doi:10.1099/mic.0.000382. PMID 27902405.
  8. ^ Wang F, Gu Y, O'Brien JP, Yi SM, Yalcin SE, Srikanth V, et al. (April 2019). "Structure of Microbial Nanowires Reveals Stacked Hemes that Transport Electrons over Micrometers". Cell. 177 (2): 361–369.e10. doi:10.1016/j.cell.2019.03.029. PMC 6720112. PMID 30951668.
  9. ^ El-Naggar MY, Wanger G, Leung KM, Yuzvinsky TD, Southam G, Yang J, et al. (October 2010). "Electrical transport along bacterial nanowires from Shewanella oneidensis MR-1". Proceedings of the National Academy of Sciences of the United States of America. 107 (42): 18127–31. Bibcode:2010PNAS..10718127E. doi:10.1073/pnas.1004880107. PMC 2964190. PMID 20937892.
  10. ^ Pirbadian S, El-Naggar MY (October 2012). "Multistep hopping and extracellular charge transfer in microbial redox chains". Physical Chemistry Chemical Physics. 14 (40): 13802–8. Bibcode:2012PCCP...1413802P. doi:10.1039/C2CP41185G. PMID 22797729.
  11. ^ Polizzi NF, Skourtis SS, Beratan DN (2012). "Physical constraints on charge transport through bacterial nanowires". Faraday Discussions. 155: 43–62, discussion 103–14. Bibcode:2012FaDi..155...43P. doi:10.1039/C1FD00098E. PMC 3392031. PMID 22470966.
  12. ^ Strycharz-Glaven SM, Snider RM, Guiseppi-Elie A, Tender LM (2011). "On the electrical conductivity of microbial nanowires and biofilms". Energy Environ Sci. 4 (11): 4366–4379. doi:10.1039/C1EE01753E.
  13. ^ Potter MC, Waller AD (1911-09-14). "Electrical effects accompanying the decomposition of organic compounds". Proceedings of the Royal Society of London. Series B, Containing Papers of a Biological Character. 84 (571): 260–276. doi:10.1098/rspb.1911.0073.
  14. ^ Myers CR, Nealson KH (June 1988). "Bacterial manganese reduction and growth with manganese oxide as the sole electron acceptor". Science. 240 (4857): 1319–21. Bibcode:1988Sci...240.1319M. doi:10.1126/science.240.4857.1319. PMID 17815852. S2CID 9662366.
  15. ^ Lovley DR, Phillips EJ (June 1988). "Novel mode of microbial energy metabolism: organic carbon oxidation coupled to dissimilatory reduction of iron or manganese". Applied and Environmental Microbiology. 54 (6): 1472–80. Bibcode:1988ApEnM..54.1472L. doi:10.1128/aem.54.6.1472-1480.1988. PMC 202682. PMID 16347658.
  16. ^ Kim B (1999). "Dynamic effects of learning capabilities and profit structures on the innovation competition". Optimal Control Applications and Methods. 20 (3): 127–144. doi:10.1002/(SICI)1099-1514(199905/06)20:3<127::AID-OCA650>3.0.CO;2-I. ISSN 1099-1514.
  17. ^ a b Rabaey K, Rozendal RA (October 2010). "Microbial electrosynthesis - revisiting the electrical route for microbial production". Nature Reviews. Microbiology. 8 (10): 706–16. doi:10.1038/nrmicro2422. PMID 20844557. S2CID 11417035.
  18. ^ a b c d Shi L, Dong H, Reguera G, Beyenal H, Lu A, Liu J, et al. (October 2016). "Extracellular electron transfer mechanisms between microorganisms and minerals". Nature Reviews. Microbiology. 14 (10): 651–62. doi:10.1038/nrmicro.2016.93. PMID 27573579. S2CID 20626915.
  19. ^ Kodesia, A.; Ghosh, M.; Chatterjee, A. (September 5, 2017). "Development of Biofilm Nanowires and Electrode for Efficient Microbial Fuel Cells (MFCs)". Thapar University Digital Repository (TuDR).
  20. ^ Malvankar NS, Vargas M, Nevin KP, Franks AE, Leang C, Kim BC, Inoue K, Mester T, Covalla SF, Johnson JP, Rotello VM, Tuominen MT, Lovley DR (August 2011). "Tunable metallic-like conductivity in microbial nanowire networks". Nature Nanotechnology. 6 (9): 573–9. Bibcode:2011NatNa...6..573M. doi:10.1038/nnano.2011.119. PMID 21822253.
  21. ^ a b c Malvankar NS, Lovley DR (June 2012). "Microbial nanowires: a new paradigm for biological electron transfer and bioelectronics". ChemSusChem. 5 (6): 1039–46. doi:10.1002/cssc.201100733. PMID 22614997.
  22. ^ Maruthupandy M, Anand M, Maduraiveeran G, Beevi AS, Priya RJ (September 2017). "Fabrication of CuO nanoparticles coated bacterial nanowire film for a high-performance electrochemical conductivity". Journal of Materials Science. 52 (18): 10766–78. Bibcode:2017JMatS..5210766M. doi:10.1007/s10853-017-1248-6. S2CID 103105219.
  23. ^ Liu X, Gao H, Ward JE, Liu X, Yin B, Fu T, et al. (February 2020). "Power generation from ambient humidity using protein nanowires". Nature. 578 (7796): 550–554. Bibcode:2020Natur.578..550L. doi:10.1038/s41586-020-2010-9. PMID 32066937.
  24. ^ Jiang S, Kim MG, Kim SJ, Jung HS, Lee SW, Noh DY, et al. (July 2011). "Bacterial formation of extracellular U(VI) nanowires". Chemical Communications. 47 (28): 8076–8. doi:10.1039/C1CC12554K. PMID 21681306.
  25. ^ Cologgi DL, Lampa-Pastirk S, Speers AM, Kelly SD, Reguera G (September 2011). "Extracellular reduction of uranium via Geobacter conductive pili as a protective cellular mechanism". Proceedings of the National Academy of Sciences of the United States of America. 108 (37): 15248–52. Bibcode:2011PNAS..10815248C. doi:10.1073/pnas.1108616108. PMC 3174638. PMID 21896750.
  26. ^ Fu T, Liu X, Gao H, Ward JE, Liu X, Yin B, et al. (April 2020). "Bioinspired bio-voltage memristors". Nature Communications. 11 (1): 1861. Bibcode:2020NatCo..11.1861F. doi:10.1038/s41467-020-15759-y. PMC 7171104. PMID 32313096.
  27. ^ "Researchers Unveil Electronics that Mimic the Human Brain in Efficient, Biological Learning". Office of News & Media Relations | UMass Amherst. Retrieved 2021-04-20.
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