Cable bacteria

Long-distance electrical conductance in sediment was first observed in 2010 as a spatial separation of sulfide oxidation and oxygen reduction in marine sediment that was interrupted and re-established at a rate faster than could be explained by chemical diffusion.^[1] It was later found that this electrical conductance could be observed across a non-conductive layer of glass microspheres, where the only possible conductive structures were filamentous bacteria belonging to the family Desulfobulbaceae.^[2] The conductivity of single, live filaments was later demonstrated by observing the oxidation state of cytochromes using Raman microscopy.^[5] The same phenomenon was later observed in freshwater sediments^[6] and groundwater aquifers.^[7] Within a 15 cm thick top layer of sediment, cable bacteria densities providing total length of up to 2 km per square centimeter of surface have been observed.^[8]

Cable bacteria filaments have a diameter of 1–4 μm and lengths of over 1 cm.^[9] The individual cells in the filaments are rod-shaped with an average length of 3 μm.^[2] As Gram-negative bacteria, they have two cell-enveloping membranes, with each cell having its own individual inner cell membrane, but the outer cell membrane is shared by all cells in a filament.^[2] In the common periplasm there are around 15–60^[2][9] electron-conducting fibers with a diameter of around 50 nm, which are visible from the outside as parallel, longitudinal ribs. They consist of proteins that are rich in nickel and sulfur, are electrically insulated, and run the entire length of the cell filament.^[9]

Cable bacteria are generally found in reduced sediments.^[10] They can be present as a single filament or as an agglomeration of filaments.^[10] Cable bacteria have been identified as being intertwined with the root hairs of aquatic plants and are present in the rhizosphere.^[10] Their distribution ranges a gradient of salinities; they are present in freshwater, saltwater lakes, and marine habitats.^[11][12] Cable bacteria have been identified in a diverse range of climatic conditions worldwide,^[13] including Denmark,^[2][6] the Netherlands,^[14] Japan,^[15] Australia,^[16] and the United States.^[17]

Cable bacteria lack flagella, but are capable of motility in the form of gliding^[18] by propelling themselves forward through the excretion of substances.^[19] Cable bacteria have been observed to move as fast as 2.2 μm/s, with an average speed of 0.5 μm/s.^[18] Speed of motility in cable bacteria is not related to size of the bacteria.^[18] The average distance a cable bacterium glides is approximately 74 μm without interruption.^[18] Cable bacteria filaments tend to bend in half, and their movement is led by the apex of the bend as opposed to leading with one tip of the filament.^[18] Twisting to move through rotational gliding is rare, but does occur.^[18] Cable bacteria likely engage in oxygen chemotaxis, as they are observed to move when in anoxic or hypoxic environments, and cease gliding when contact with oxygen is made.^[18] Although motility is important for other microorganisms, once cable bacteria are located in a place that connects oxygen to sulfide, they no longer need to move.^[18] The reduced need for motility could explain why the cable bacteria genome contains fewer operons related to chemotaxis than other Desulfobulbaceae.^[19] Fewer operons related to chemotaxis results in limited motility.^[19]

The two candidate genera of cable bacteria until now (2016) described: Electrothrix containing four candidate species, found in marine or brackish sediments, and Electronema containing two candidate species, found in freshwater sediments, seem to be a monophyletic group.^[15] Freshwater and marine cable bacteria have been found to be 88% similar based on 16S ribosomal RNA comparisons.^[20] These genera are classified within the family Desulfobulbaceae.^[19] Cable bacteria are defined by their function rather than their phylogeny, and it is possible that further cable bacteria taxa will be discovered.

Cable bacteria strongly influence the geochemical properties of the surrounding environment. Their activity promotes the oxidation of iron at the surface of the sediment, and the resulting iron oxides bind phosphorus-containing compounds^[21] and hydrogen sulfide,^[22] limiting the amount of phosphorus and hydrogen sulfide in the water. Phosphorus can cause eutrophication, and hydrogen sulfide can be toxic to marine life, meaning that cable bacteria play an important role in maintaining marine ecosystems in coastal areas.

The presence of cable bacteria can lead to a decrease in methane emissions from saturated soils. The transfer of electrons through cable bacteria allows the sulfate reduction that occurs in inundated soils to be balanced by sulfide oxidation. Oxidation is possible because of the release of electrons through the cable bacteria filaments. Through this balance, sulfate remains readily available for sulfate reducing bacteria, which out compete methanogens. This causes a decrease in production of methane by methanogens.^[23]

Cable bacteria have been found associated with benthic microbial fuel cells, devices that convert chemical energy on the ocean floor to electrical energy.^[24] In the future, cable bacteria may play a role in increasing the efficiency of microbial fuel cells deployed in sedimentary environments. Cable bacteria have also been found associated with a bioelectrochemical system that enhances the degradation of marine sediment contaminated by hydrocarbons ^[25] and thus may play a role in future oil spill cleanup technologies.

• Biodegradable electronics

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