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Dissimilatory metal-reducing microorganisms are a group of
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
s (both
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
and
archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...
) that can perform
anaerobic respiration Anaerobic respiration is respiration using electron acceptors other than molecular oxygen (O2). Although oxygen is not the final electron acceptor, the process still uses a respiratory electron transport chain. In aerobic organisms undergoing re ...
utilizing a
metal A metal (from Greek μέταλλον ''métallon'', "mine, quarry, metal") is a material that, when freshly prepared, polished, or fractured, shows a lustrous appearance, and conducts electricity and heat relatively well. Metals are typicall ...
as terminal
electron acceptor An electron acceptor is a chemical entity that accepts electrons transferred to it from another compound. It is an oxidizing agent that, by virtue of its accepting electrons, is itself reduced in the process. Electron acceptors are sometimes mista ...
rather than
molecular oxygen There are several known allotropes of oxygen. The most familiar is molecular oxygen (O2), present at significant levels in Earth's atmosphere and also known as dioxygen or triplet oxygen. Another is the highly reactive ozone (O3). Others are: *A ...
(O2), which is the terminal electron acceptor reduced to
water Water (chemical formula ) is an inorganic, transparent, tasteless, odorless, and nearly colorless chemical substance, which is the main constituent of Earth's hydrosphere and the fluids of all known living organisms (in which it acts as a ...
(H2O) in
aerobic respiration Cellular respiration is the process by which biological fuels are oxidised in the presence of an inorganic electron acceptor such as oxygen to produce large amounts of energy, to drive the bulk production of ATP. Cellular respiration may be des ...
. The most common metals used for this end are
iron Iron () is a chemical element with symbol Fe (from la, ferrum) and atomic number 26. It is a metal that belongs to the first transition series and group 8 of the periodic table. It is, by mass, the most common element on Earth, right in f ...
e(III)and
manganese Manganese is a chemical element with the symbol Mn and atomic number 25. It is a hard, brittle, silvery metal, often found in minerals in combination with iron. Manganese is a transition metal with a multifaceted array of industrial alloy use ...
n(IV) which are reduced to Fe(II) and Mn(II) respectively, and most microorganisms that reduce Fe(III) can reduce Mn(IV) as well. But other metals and metalloids are also used as terminal electron acceptors, such as
vanadium Vanadium is a chemical element with the symbol V and atomic number 23. It is a hard, silvery-grey, malleable transition metal. The elemental metal is rarely found in nature, but once isolated artificially, the formation of an oxide layer ( pas ...
(V)
chromium Chromium is a chemical element with the symbol Cr and atomic number 24. It is the first element in group 6. It is a steely-grey, lustrous, hard, and brittle transition metal. Chromium metal is valued for its high corrosion resistance and hardne ...
r(VI)
molybdenum Molybdenum is a chemical element with the symbol Mo and atomic number 42 which is located in period 5 and group 6. The name is from Neo-Latin ''molybdaenum'', which is based on Ancient Greek ', meaning lead, since its ores were confused with lea ...
o(VI)
cobalt Cobalt is a chemical element with the symbol Co and atomic number 27. As with nickel, cobalt is found in the Earth's crust only in a chemically combined form, save for small deposits found in alloys of natural meteoric iron. The free element, pr ...
o(III)
palladium Palladium is a chemical element with the symbol Pd and atomic number 46. It is a rare and lustrous silvery-white metal discovered in 1803 by the English chemist William Hyde Wollaston. He named it after the asteroid Pallas, which was itself na ...
d(II)
gold Gold is a chemical element with the symbol Au (from la, aurum) and atomic number 79. This makes it one of the higher atomic number elements that occur naturally. It is a bright, slightly orange-yellow, dense, soft, malleable, and ductile met ...
u(III) and
mercury Mercury commonly refers to: * Mercury (planet), the nearest planet to the Sun * Mercury (element), a metallic chemical element with the symbol Hg * Mercury (mythology), a Roman god Mercury or The Mercury may also refer to: Companies * Merc ...
g(II)


Conditions and mechanisms for dissimilatory metal reduction

Dissimilatory metal reducers are a diverse group of microorganisms, which is reflected in the factors that affect the different forms of metal reduction. The process of dissimilatory metal reduction occurs in the absence of oxygen (O2), but dissimilatory metal reducers include both obligate (strict) anaerobes, such as the family
Geobacter ''Geobacter'' is a genus of bacteria. ''Geobacter'' species are anaerobic respiration bacterial species which have capabilities that make them useful in bioremediation. ''Geobacter'' was found to be the first organism with the ability to oxidiz ...
aceae, and
facultative anaerobe A facultative anaerobic organism is an organism that makes ATP by aerobic respiration if oxygen is present, but is capable of switching to fermentation if oxygen is absent. Some examples of facultatively anaerobic bacteria are ''Staphylococcus' ...
s, such as '' Shewanella'' spp. As well, across the dissimilatory metal reducers species, various electron donors are used in the oxidative reaction that is coupled to metal reduction. For instance, some species are limited to small organic acids and hydrogen (H2), whereas others may oxidize aromatic compounds. In certain instances, such as Cr(VI) reduction, the use of small organic compounds can optimize the rate of metal reduction. Another factor that influences metal respiration is environmental acidity. Although acidophilic and alkaliphilic dissimilatory metal reducers exist, the neutrophilic metal reducers group contains the most well-characterized genera. In soil and sediment environments, where the pH is often neutral, metals like iron are found in their solid oxidized forms, and exhibit variable reduction potential, which can affect their use by microorganisms. Due to the impermeability of the cell wall to minerals and the insolubility of metal oxides, dissimilatory metal reducers have developed ways to reduce metals extracellularly via electron transfer. Cytochromes ''c'', which are transmembrane proteins, play an important role in transporting electrons from the cytosol to enzymes attached to the outside of the cell. The electrons are then further transported to the terminal electron acceptor via direct interaction between the enzymes and the metal oxide. In addition to establishing direct contact, dissimilatory metal reducers also display the ability to perform ranged metal reduction. For instance, some species of dissimilatory metal reducers produce compounds that can dissolve insoluble minerals or act as electron shuttles, enabling them to perform metal reduction from a distance. Other organic compounds frequently found in soils and sediments, such as humic acids, may also act as electron shuttles.Lovley, D. R., Fraga, J. L., Blunt-Harris, E. L., Hayes, L. A., Phillips, E. J. P., & Coates, J. D. (1998). Humic substances as a mediator for microbially catalyzed metal reduction. Acta hydrochimica et hydrobiologica, 26(3), 152-157. In
biofilm A biofilm comprises any syntrophic consortium of microorganisms in which cells stick to each other and often also to a surface. These adherent cells become embedded within a slimy extracellular matrix that is composed of extracellular ...
s, nanowires and multistep electron hopping (in which electrons jump from cell to cell towards the mineral) have also been suggested as methods for reducing metals without requiring direct cell contact. It has been proposed that cytochromes c are involved in both of these mechanisms. In nanowires, for instance, cytochromes c function as the final component that transfers electrons to the metal oxide.


Terminal electron acceptors

A wide range of Fe(III)-bearing minerals have been observed to function as terminal electron acceptors, including magnetite, hematite, goethite, lepidocrocite, ferrihydrite, hydrous ferric oxide, smectite, illite, jarosite, among others.


Secondary mineral formation

In natural systems, secondary minerals may form as a byproduct of bacterial metal reduction. Commonly observed secondary minerals produced during experimental bio-reduction by dissimilatory metal reducers include magnetite, siderite, green rust, vivianite, and hydrous Fe(II)-carbonate.


Genera that include dissimilatory metal reducers

* '' Albidiferax'' (Betaproteobacteria) * '' Shewanella'' (Gammaproteobacteria) * ''
Geobacter ''Geobacter'' is a genus of bacteria. ''Geobacter'' species are anaerobic respiration bacterial species which have capabilities that make them useful in bioremediation. ''Geobacter'' was found to be the first organism with the ability to oxidiz ...
'' (Deltaproteobacteria) * ''
Geothrix fermentans ''Geothrix fermentans'' is a rod-shaped, anaerobic bacterium. It is about 0.1 µm in diameter and ranges from 2-3 µm in length. Cell arrangement occurs singly and in chains. ''Geothrix fermentans'' can normally be found in aquatic sed ...
'' (Acidobacteria) * ''
Deferribacter ''Deferribacter'' is a genus in the phylum Deferribacterota (Bacteria). Etymology The name ''Deferribacter'' derives from:Latin pref. ''de''-, from; Latin noun ''ferrum'', iron; New Latin grammatical gender, masculine gender noun, a rod''-bacter ...
'' (Deferribacteres) * '' Thermoanaerobacter'' (Firmicutes)


References

{{reflist Bacteria Metabolism Extremophiles