Oxygen consumption in marine sediments is often coupled to the oxidation of sulphide generated by degradation of organic matter in deeper, oxygen-free layers. Geochemical observations have shown that this coupling can be mediated by electric currents carried by unidentified electron transporters across centimetre-wide zones. Here we present evidence that the native conductors are long, filamentous bacteria. They abounded in sediment zones with electric currents and along their length they contained strings with distinct properties in accordance with a function as electron transporters. Living, electrical cables add a new dimension to the understanding of interactions in nature and may find use in technology development.
Bacterial nanowires offer an extracellular electron transport (EET) pathway for linking the respiratory chain of bacteria to external surfaces, including oxidized metals in the environment and engineered electrodes in renewable energy devices. Despite the global, environmental, and technological consequences of this biotic-abiotic interaction, the composition, physiological relevance, and electron transport mechanisms of bacterial nanowires remain unclear. We report, to our knowledge, the first in vivo observations of the formation and respiratory impact of nanowires in the model metal-reducing microbe Shewanella oneidensis MR-1. Live fluorescence measurements, immunolabeling, and quantitative gene expression analysis point to S. oneidensis MR-1 nanowires as extensions of the outer membrane and periplasm that include the multiheme cytochromes responsible for EET, rather than pilin-based structures as previously thought. These membrane extensions are associated with outer membrane vesicles, structures ubiquitous in Gram-negative bacteria, and are consistent with bacterial nanowires that mediate long-range EET by the previously proposed multistep redox hopping mechanism. Redox-functionalized membrane and vesicular extensions may represent a general microbial strategy for electron transport and energy distribution. R eduction-oxidation (redox) reactions and electron transport are essential to the energy conversion pathways of living cells (1). Respiratory organisms generate ATP molecules-life's universal energy currency-by harnessing the free energy of electron transport from electron donors (fuels) to electron acceptors (oxidants) through biological redox chains. In contrast to most eukaryotes, which are limited to relatively few carbon compounds as electron donors and oxygen as the predominant electron acceptor, prokaryotes have evolved into versatile energy scavengers. Microbes can wield an astounding number of metabolic pathways to extract energy from diverse organic and inorganic electron donors and acceptors, which has significant consequences for global biogeochemical cycles (2-4).
Bacterial nanowires are extracellular appendages that have been suggested as pathways for electron transport in phylogenetically diverse microorganisms, including dissimilatory metal-reducing bacteria and photosynthetic cyanobacteria. However, there has been no evidence presented to demonstrate electron transport along the length of bacterial nanowires. Here we report electron transport measurements along individually addressed bacterial nanowires derived from electron-acceptor–limited cultures of the dissimilatory metal-reducing bacterium Shewanella oneidensis MR-1. Transport along the bacterial nanowires was independently evaluated by two techniques: ( i ) nanofabricated electrodes patterned on top of individual nanowires, and ( ii ) conducting probe atomic force microscopy at various points along a single nanowire bridging a metallic electrode and the conductive atomic force microscopy tip. The S. oneidensis MR-1 nanowires were found to be electrically conductive along micrometer-length scales with electron transport rates up to 10 9 /s at 100 mV of applied bias and a measured resistivity on the order of 1 Ω·cm. Mutants deficient in genes for c -type decaheme cytochromes MtrC and OmcA produce appendages that are morphologically consistent with bacterial nanowires, but were found to be nonconductive. The measurements reported here allow for bacterial nanowires to serve as a viable microbial strategy for extracellular electron transport.
TiO2 photocatalysts have been found to kill cancer cells, bacteria and viruses under mild UV illumination, which offers numerous potential applications. On the other hand, Ag has long been proved as a good antibacterial material as well. The advantage of Ag-TiO2 nanocomposite is to expand the nanomaterial's antibacterial function to a broader range of working conditions. In this study neat TiO2 and Ag-TiO2 composite nanofilms were successfully prepared on silicon wafer via the sol-gel method by the spin-coating technique. The as-prepared composite Ag-TiO2 and TiO2 films with different silver content were characterized by scanning electron microscopy (SEM), atomic force microscopy (AFM), x-ray diffraction (XRD) and x-ray photoelectron spectroscopy (XPS) to determine the topologies, microstructures and chemical compositions, respectively. It was found that the silver nanoparticles were uniformly distributed and strongly attached to the mesoporous TiO2 matrix. The morphology of the composite film could be controlled by simply tuning the molar ratio of the silver nitrate aqueous solution. XPS results confirmed that the Ag was in the Ag(0) state. The antimicrobial effect of the synthesized nanofilms was carried out against gram-negative bacteria (Escherichia coli ATCC 29425) by using an 8 W UV lamp with a constant relative intensity of 0.6 mW cm(-2) and in the dark respectively. The synthesized Ag-TiO2 thin films showed enhanced bactericidal activities compared to the neat TiO2 nanofilm both in the dark and under UV illumination.
The study of electrical transport in biomolecular materials is critical to our fundamental understanding of physiology and to the development of practical bioelectronics applications. In this study, we investigated the electronic transport characteristics of Shewanella oneidensis MR-1 nanowires by conducting-probe atomic force microscopy (CP-AFM) and by constructing field-effect transistors (FETs) based on individual S. oneidensis nanowires. Here we show that S. oneidensis nanowires exhibit p-type, tunable electronic behavior with a field-effect mobility on the order of 10(-1) cm(2)/(V s), comparable to devices based on synthetic organic semiconductors. This study opens up opportunities to use such bacterial nanowires as a new semiconducting biomaterial for making bioelectronics and to enhance the power output of microbial fuel cells through engineering the interfaces between metallic electrodes and bacterial nanowires.
Bacterial nanowires offer a pathway for extracellular electron transfer (EET) by linking the respiratory chain of bacteria to external surfaces, including oxidized metals in the environment and engineered electrodes in renewable energy devices. Specifically, nanowires of the model metal-reducing bacterium Shewanella oneidensis MR-1 were previously shown to be conductive under non-physiological conditions. Despite the global, environmental, and technological consequences of bacterial nanowire-mediated EET, the composition, electron transport mechanism, and physiological relevance of these appendages remain unclear. The nanowires of S. oneidensis MR-1 were previously thought, but never shown, to be bacterial pili. In addition, the transport mechanism through bacterial nanowires has been the subject of intense debate, with ''metallic-like'' band transport and multistep redox hopping between multiheme cytochromes as the two proposed mechanisms. Here we report the first in vivo observations of the formation and respiratory impact of nanowires in S. oneidensis MR-1. Using live fluorescence measurements and quantitative gene expression analysis, we demonstrate that S. oneidensis MR-1 nanowires are extensions of the outer membrane and periplasm, rather than pilin-based structures. We show, through immunolabeling, that multiheme cytochromes localize to nanowires, in turn supporting the multistep redox hopping model as the transport mechanism. Furthermore, these bacterial nanowires are associated with outer membrane vesicles, structures ubiquitous in Gram-negative bacteria, and occasionally appear as membrane vesicle chains that transition to smoother filaments. Redox-functionalized membrane and vesicular extensions may represent a general microbial strategy for electron transport and energy distribution.
In limiting oxygen as an electron acceptor, the dissimilatory metal-reducing bacterium Shewanella oneidensis MR-1 rapidly forms nanowires, extensions of its outer membrane containing the cytochromes MtrC and OmcA needed for extracellular electron transfer. RNA sequencing (RNA-Seq) analysis was employed to determine differential gene expression over time from triplicate chemostat cultures that were limited for oxygen. We identified 465 genes with decreased expression and 677 genes with increased expression. The coordinated increased expression of heme biosynthesis, cytochrome maturation, and transport pathways indicates that S. oneidensis MR-1 increases cytochrome production, including the transcription of genes encoding MtrA, MtrC, and OmcA, and transports these decaheme cytochromes across the cytoplasmic membrane during electron acceptor limitation and nanowire formation. In contrast, the expression of the mtrA and mtrC homologs mtrF and mtrD either remains unaffected or decreases under these conditions. The ompW gene, encoding a small outer membrane porin, has 40-fold higher expression during oxygen limitation, and it is proposed that OmpW plays a role in cation transport to maintain electrical neutrality during electron transfer. The genes encoding the anaerobic respiration regulator cyclic AMP receptor protein (CRP) and the extracytoplasmic function sigma factor RpoE are among the transcription factor genes with increased expression. RpoE might function by signaling the initial response to oxygen limitation. Our results show that RpoE activates transcription from promoters upstream of mtrC and omcA. The transcriptome and mutant analyses of S. oneidensis MR-1 nanowire production are consistent with independent regulatory mechanisms for extending the outer membrane into tubular structures and for ensuring the electron transfer function of the nanowires. IMPORTANCEShewanella oneidensis MR-1 has the capacity to transfer electrons to its external surface using extensions of the outer membrane called bacterial nanowires. These bacterial nanowires link the cell's respiratory chain to external surfaces, including oxidized metals important in bioremediation, and explain why S. oneidensis can be utilized as a component of microbial fuel cells, a form of renewable energy. In this work, we use differential gene expression analysis to focus on which genes function to produce the nanowires and promote extracellular electron transfer during oxygen limitation. Among the genes that are expressed at high levels are those encoding cytochrome proteins necessary for electron transfer. Shewanella coordinates the increased expression of regulators, metabolic pathways, and transport pathways to ensure that cytochromes efficiently transfer electrons along the nanowires.
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