Anaerobic Methane Oxidation
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Anaerobic oxidation of methane (AOM) is a methane-consuming microbial process occurring in
anoxic Anoxia means a total depletion in the level of oxygen, an extreme form of hypoxia or "low oxygen". The terms anoxia and hypoxia are used in various contexts: * Anoxic waters, sea water, fresh water or groundwater that are depleted of dissolved ox ...
marine and freshwater
sediments Sediment is a solid material that is transported to a new location where it is deposited. It occurs naturally and, through the processes of weathering and erosion, is broken down and subsequently sediment transport, transported by the action of ...
. AOM is known to occur among
mesophiles A mesophile is an organism that grows best in moderate temperature, neither too hot nor too cold, with an optimum growth range from . The optimum growth temperature for these organisms is 37 °C (about 99 °F). The term is mainly applied ...
, but also in
psychrophiles Psychrophiles or cryophiles (adj. ''psychrophilic'' or ''cryophilic'') are extremophilic organisms that are capable of growth and reproduction in low temperatures, ranging from to . They are found in places that are permanently cold, such as th ...
,
thermophiles A thermophile is a type of extremophile that thrives at relatively high temperatures, between . Many thermophiles are archaea, though some of them are bacteria and fungi. Thermophilic eubacteria are suggested to have been among the earliest bacte ...
,
halophiles A halophile (from the Greek word for 'salt-loving') is an extremophile that thrives in high salt concentrations. In chemical terms, halophile refers to a Lewis acidic species that has some ability to extract halides from other chemical species. ...
,
acidophiles Acidophiles or acidophilic organisms are those that thrive under highly acidic conditions (usually at pH 5.0 or below). These organisms can be found in different branches of the tree of life, including Archaea, Bacteria,Becker, A.Types of Bacteri ...
, and alkophiles. During AOM,
methane Methane ( , ) is a chemical compound with the chemical formula (one carbon atom bonded to four hydrogen atoms). It is a group-14 hydride, the simplest alkane, and the main constituent of natural gas. The abundance of methane on Earth makes ...
is oxidized with different terminal
electron acceptors An oxidizing agent (also known as an oxidant, oxidizer, electron recipient, or electron acceptor) is a substance in a redox chemical reaction that gains or " accepts"/"receives" an electron from a (called the , , or ''electron donor''). In ot ...
such as
sulfate The sulfate or sulphate ion is a polyatomic anion with the empirical formula . Salts, acid derivatives, and peroxides of sulfate are widely used in industry. Sulfates occur widely in everyday life. Sulfates are salts of sulfuric acid and many ...
,
nitrate Nitrate is a polyatomic ion with the chemical formula . salt (chemistry), Salts containing this ion are called nitrates. Nitrates are common components of fertilizers and explosives. Almost all inorganic nitrates are solubility, soluble in wa ...
,
nitrite The nitrite polyatomic ion, ion has the chemical formula . Nitrite (mostly sodium nitrite) is widely used throughout chemical and pharmaceutical industries. The nitrite anion is a pervasive intermediate in the nitrogen cycle in nature. The name ...
and
metals A metal () is a material that, when polished or fractured, shows a lustrous appearance, and conducts electricity and heat relatively well. These properties are all associated with having electrons available at the Fermi level, as against no ...
, either alone or in
syntrophy In biology, syntrophy, syntrophism, or cross-feeding () is the cooperative interaction between at least two microbial species to degrade a single substrate. This type of biological interaction typically involves the transfer of one or more met ...
with a partner organism.


Coupled to sulfate reduction

The overall reaction is: :CH4 + SO42− → HCO3 + HS + H2O Sulfate-driven AOM is mediated by a syntrophic consortium of methanotrophic
archaea Archaea ( ) is a Domain (biology), domain of organisms. Traditionally, Archaea only included its Prokaryote, prokaryotic members, but this has since been found to be paraphyletic, as eukaryotes are known to have evolved from archaea. Even thou ...
and
sulfate-reducing bacteria Sulfate-reducing microorganisms (SRM) or sulfate-reducing prokaryotes (SRP) are a group composed of sulfate-reducing bacteria (SRB) and sulfate-reducing archaea (SRA), both of which can perform anaerobic respiration utilizing sulfate () as termina ...
. They often form small aggregates or sometimes voluminous mats. The archaeal partner is abbreviated ANME, which stands for "anaerobic
methanotroph Methanotrophs (sometimes called methanophiles) are prokaryotes that metabolize methane as their source of carbon and chemical energy. They are bacteria or archaea, can grow aerobically or anaerobically, and require single-carbon compounds to ...
". ANME's are very closely related to methanogenic archaea and recent investigations suggest that AOM is an enzymatic reversal of
methanogenesis Methanogenesis or biomethanation is the formation of methane coupled to energy conservation by microbes known as methanogens. It is the fourth and final stage of anaerobic digestion. Organisms capable of producing methane for energy conservation h ...
. It is still poorly understood how the
syntrophic In biology, syntrophy, syntrophism, or cross-feeding () is the cooperative interaction between at least two microbial species to degrade a single substrate. This type of biological interaction typically involves the transfer of one or more metabol ...
partners interact and which intermediates are exchanged between the archaeal and bacterial cell. The research on AOM is hindered by the fact that the responsible organisms have not been isolated. This is because these organisms show very slow growth rates with a minimum doubling time of a few months. Countless isolation efforts have not been able to isolate one of the anaerobic methanotrophs, a possible explanation can be that the ANME archaea and the SRB have an obligate syntrophic interaction and can therefore not be isolated individually. In
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...
marine areas with strong methane releases from fossil reservoirs (e.g. at
cold seeps A cold seep (sometimes called a cold vent) is an area of the ocean floor where seepage of fluids rich in hydrogen sulfide, methane, and other hydrocarbons occurs, often in the form of a brine pool. ''Cold'' does not mean that the temperature o ...
,
mud volcano A mud volcano or mud dome is a landform created by the eruption of mud or Slurry, slurries, water and gases. Several geological processes may cause the formation of mud volcanoes. Mud volcanoes are not true Igneous rock, igneous volcanoes as th ...
es or
gas hydrate Clathrate hydrates, or gas hydrates, clathrates, or hydrates, are crystalline water-based solids physically resembling ice, in which small non-polar molecules (typically gases) or polar molecules with large hydrophobic moieties are trapped inside ...
deposits) AOM can be so high that
chemosynthetic In biochemistry, chemosynthesis is the biological conversion of one or more carbon-containing molecules (usually carbon dioxide or methane) and nutrients into organic matter using the oxidation of inorganic compounds (e.g., hydrogen gas, hydrog ...
organisms like filamentous sulfur bacteria (see
Beggiatoa ''Beggiatoa'' is a genus of ''Gammaproteobacteria'' belonging to the order '' Thiotrichales'', in the ''Pseudomonadota'' phylum. These bacteria form colorless filaments composed of cells that can be up to 200 μm in diameter, and are one of ...
) or animals (clams, tube worms) with
symbiont Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biological interaction, between two organisms of different species. The two organisms, termed symbionts, can fo ...
sulfide-oxidizing bacteria can thrive on the large amounts of
hydrogen sulfide Hydrogen sulfide is a chemical compound with the formula . It is a colorless chalcogen-hydride gas, and is toxic, corrosive, and flammable. Trace amounts in ambient atmosphere have a characteristic foul odor of rotten eggs. Swedish chemist ...
that are produced during AOM. The
bicarbonate In inorganic chemistry, bicarbonate (IUPAC-recommended nomenclature: hydrogencarbonate) is an intermediate form in the deprotonation of carbonic acid. It is a polyatomic anion with the chemical formula . Bicarbonate serves a crucial bioche ...
(HCO3) produced from AOM can (i) get sequestered in the sediments by the precipitation of
calcium carbonate Calcium carbonate is a chemical compound with the chemical formula . It is a common substance found in Rock (geology), rocks as the minerals calcite and aragonite, most notably in chalk and limestone, eggshells, gastropod shells, shellfish skel ...
or so-called methane-derived
authigenic Authigenesis is the process whereby a mineral or sedimentary rock deposit is generated where it is found or observed. Such deposits are described as authigenic. Authigenic sedimentary Mineral, minerals form during or after sedimentation by Precipita ...
carbonates and (ii) get released to the overlying water column. Methane-derived
authigenic Authigenesis is the process whereby a mineral or sedimentary rock deposit is generated where it is found or observed. Such deposits are described as authigenic. Authigenic sedimentary Mineral, minerals form during or after sedimentation by Precipita ...
carbonates are known to be the most 13C depleted carbonates on Earth, with δ13C values as low as -125 per mil PDB reported.


Coupled to nitrate and nitrite reduction

The overall reactions are: :CH4 + 4 NO3 → CO2 + 4 NO2 + 2 H2O :3 CH4 + 8 NO2 + 8 H+ → 3 CO2 + 4 N2 + 10 H2O ANME-2d is shown to be responsible nitrate-driven AOM. The ANME-2d, named '' Methanoperedens nitroreducens'', is able to perform nitrate-driven AOM without a partner organism via reverse methanogenesis with nitrate as the terminal electron acceptor, using genes for nitrate reduction that have been laterally transferred from a bacterial donor. This was also the first complete reverse methanogenesis pathway including the ''mcr'' and ''mer'' genes. In 2010,
omics Omics is the collective characterization and quantification of entire sets of biological molecules and the investigation of how they translate into the structure, function, and dynamics of an organism or group of organisms. The branches of scien ...
, especially
metagenomics Metagenomics is the study of all genetics, genetic material from all organisms in a particular environment, providing insights into their composition, diversity, and functional potential. Metagenomics has allowed researchers to profile the mic ...
, analysis showed that nitrite reduction can be coupled to methane oxidation by a single bacterial species ''Candidatus'' Methylomirabilis oxyfera ( phylum NC10), without the need for an
archaeal Archaea ( ) is a domain of organisms. Traditionally, Archaea only included its prokaryotic members, but this has since been found to be paraphyletic, as eukaryotes are known to have evolved from archaea. Even though the domain Archaea cladis ...
partner.


Environmental relevance

AOM is considered an important process reducing the emission of the
greenhouse gas Greenhouse gases (GHGs) are the gases in the atmosphere that raise the surface temperature of planets such as the Earth. Unlike other gases, greenhouse gases absorb the radiations that a planet emits, resulting in the greenhouse effect. T ...
methane from the ocean into the atmosphere. It is estimated that almost 80% of all the methane that arises from marine sediments is oxidized anaerobically by this process.


See also

*
Borg (microbiology) Borgs are large (up to ~1 Mbp) extrachromosomal linear DNA elements found in methanotrophic archaea (specifically '' Methanoperedens'' spp.) that live in oxygen-starved environments such as deep mud. They have been found in organisms isolated fro ...


References


Bibliography

* Dennis D. Coleman; J. Bruno Risatti; Martin Schoell (1981) ''Fractionation of carbon and hydrogen isotopes by methane-oxidizing bacteria'' , Geochimica et Cosmochimica Acta , Volume 45, Issue 7, July 1981, Pages 1033-1037 , https://doi.org/10.1016/0016-7037(81)90129-0
abstract


External links

{{wikispecies Methane Limnology