Methanogenesis
Encyclopedia
Methanogenesis or biomethanation is the formation of methane
Methane
Methane is a chemical compound with the chemical formula . It is the simplest alkane, the principal component of natural gas, and probably the most abundant organic compound on earth. The relative abundance of methane makes it an attractive fuel...

 by microbes known as methanogen
Methanogen
Methanogens are microorganisms that produce methane as a metabolic byproduct in anoxic conditions. They are classified as archaea, a group quite distinct from bacteria...

s. Organisms capable of producing methane have been identified only from the domain
Domain (biology)
In biological taxonomy, a domain is the highest taxonomic rank of organisms, higher than a kingdom. According to the three-domain system of Carl Woese, introduced in 1990, the Tree of Life consists of three domains: Archaea, Bacteria and Eukarya...

 Archaea
Archaea
The Archaea are a group of single-celled microorganisms. A single individual or species from this domain is called an archaeon...

, a group phylogenetically
Phylogenetics
In biology, phylogenetics is the study of evolutionary relatedness among groups of organisms , which is discovered through molecular sequencing data and morphological data matrices...

 distinct from both eukaryote
Eukaryote
A eukaryote is an organism whose cells contain complex structures enclosed within membranes. Eukaryotes may more formally be referred to as the taxon Eukarya or Eukaryota. The defining membrane-bound structure that sets eukaryotic cells apart from prokaryotic cells is the nucleus, or nuclear...

s and bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...

, although many live in close association with anaerobic bacteria. The production of methane is an important and widespread form of microbial metabolism
Metabolism
Metabolism is the set of chemical reactions that happen in the cells of living organisms to sustain life. These processes allow organisms to grow and reproduce, maintain their structures, and respond to their environments. Metabolism is usually divided into two categories...

. In most environments, it is the final step in the decomposition of biomass
Biomass (ecology)
Biomass, in ecology, is the mass of living biological organisms in a given area or ecosystem at a given time. Biomass can refer to species biomass, which is the mass of one or more species, or to community biomass, which is the mass of all species in the community. It can include microorganisms,...

.

Recently, some experiments have suggested that leaf
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....

 tissues of living plants emit methane. Other research has indicated that the plants are not actually generating methane; they are just absorbing methane from the soil and then emitting it through their leaf tissues. There may still be some unknown mechanism by which plants produce methane, but that is by no means certain.

Biochemistry of methanogenesis

Methanogenesis in microbes is a form of anaerobic respiration
Anaerobic respiration
Anaerobic respiration is a form of respiration using electron acceptors other than oxygen. Although oxygen is not used as the final electron acceptor, the process still uses a respiratory electron transport chain; it is respiration without oxygen...

. Methanogens do not use oxygen to respire; in fact, oxygen inhibits the growth of methanogens. The terminal electron acceptor in methanogenesis is not oxygen, but carbon. The carbon can occur in a small number of organic compounds, all with low molecular weights. The two best described pathways involve the use of carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...

 and acetic acid
Acetic acid
Acetic acid is an organic compound with the chemical formula CH3CO2H . It is a colourless liquid that when undiluted is also called glacial acetic acid. Acetic acid is the main component of vinegar , and has a distinctive sour taste and pungent smell...

 as terminal electron acceptors:

CO2 + 4 H2 → CH4 + 2H2O

CH3COOH → CH4 + CO2

However, methanogenesis has been shown to use carbon from other small organic compounds, such as formic acid
Formic acid
Formic acid is the simplest carboxylic acid. Its chemical formula is HCOOH or HCO2H. It is an important intermediate in chemical synthesis and occurs naturally, most notably in the venom of bee and ant stings. In fact, its name comes from the Latin word for ant, formica, referring to its early...

 (formate), methanol
Methanol
Methanol, also known as methyl alcohol, wood alcohol, wood naphtha or wood spirits, is a chemical with the formula CH3OH . It is the simplest alcohol, and is a light, volatile, colorless, flammable liquid with a distinctive odor very similar to, but slightly sweeter than, ethanol...

, methylamines, dimethyl sulfide
Dimethyl sulfide
Dimethyl sulfide or methylthiomethane is an organosulfur compound with the formula 2S. Dimethyl sulfide is a water-insoluble flammable liquid that boils at and has a characteristic disagreeable odor. It is a component of the smell produced from cooking of certain vegetables, notably maize,...

, and methanethiol
Methanethiol
Methanethiol is a colorless gas with a smell like rotten cabbage. It is a natural substance found in the blood and brain of humans and other animal as well as plant tissues. It is disposed of through animal feces. It occurs naturally in certain foods, such as some nuts and cheese...

.

The biochemistry of methanogenesis is relatively complex, involving the following coenzymes and cofactors: F420
Coenzyme F420
Coenzyme F420 or 8-hydroxy-5-deazaflavin is a coenzyme involved in redox reactions in methanogens , in many Actinobacteria, and sporadically in other bacterial lineages. It is a flavin derivative...

, coenzyme B
Coenzyme B
Coenzyme B is a coenzyme required for redox reactions in methanogens. The full chemical name of coenzyme B is 7-mercaptoheptanoylthreoninephosphate...

, coenzyme M
Coenzyme M
Coenzyme M is a coenzyme required for methyl-transfer reactions in the metabolism of methanogens. The coenzyme is an anion with the formula . It is named 2-mercaptoethanesulfonate and abbreviated HS–CoM. The cation is unimportant, but the sodium salt is most available...

, methanofuran
Methanofuran
Methanofuran describes a family of chemical compounds found in methanogenic archaea. These species feature a 2-aminomethylfuran linked to phenoxy group...

, and methanopterin.

Importance in carbon cycle

Methanogenesis is the final step in the decay of organic matter. During the decay process, electron acceptors (such as oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...

, ferric iron, sulfate
Sulfate
In inorganic chemistry, a sulfate is a salt of sulfuric acid.-Chemical properties:...

, nitrate
Nitrate
The nitrate ion is a polyatomic ion with the molecular formula NO and a molecular mass of 62.0049 g/mol. It is the conjugate base of nitric acid, consisting of one central nitrogen atom surrounded by three identically-bonded oxygen atoms in a trigonal planar arrangement. The nitrate ion carries a...

, and manganese
Manganese
Manganese is a chemical element, designated by the symbol Mn. It has the atomic number 25. It is found as a free element in nature , and in many minerals...

) become depleted, while hydrogen
Hydrogen
Hydrogen is the chemical element with atomic number 1. It is represented by the symbol H. With an average atomic weight of , hydrogen is the lightest and most abundant chemical element, constituting roughly 75% of the Universe's chemical elemental mass. Stars in the main sequence are mainly...

 (H2) and carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...

 accumulate. Light organics produced by fermentation
Fermentation (biochemistry)
Fermentation is the process of extracting energy from the oxidation of organic compounds, such as carbohydrates, using an endogenous electron acceptor, which is usually an organic compound. In contrast, respiration is where electrons are donated to an exogenous electron acceptor, such as oxygen,...

 also accumulate. During advanced stages of organic decay, all electron acceptors become depleted except carbon dioxide. Carbon dioxide is a product of most catabolic processes, so it is not depleted like other potential electron acceptors.

Only methanogenesis and fermentation can occur in the absence of electron acceptors other than carbon. Fermentation only allows the breakdown of larger organic compounds, and produces small organic compounds. Methanogenesis effectively removes the semi-final products of decay: hydrogen, small organics, and carbon dioxide. Without methanogenesis, a great deal of carbon (in the form of fermentation products) would accumulate in anaerobic environments.

In ruminants

Methanogenesis occurs in the guts of humans and other animals, especially ruminants. In the rumen
Rumen
The rumen, also known as a paunch, forms the larger part of the reticulorumen, which is the first chamber in the alimentary canal of ruminant animals. It serves as the primary site for microbial fermentation of ingested feed...

, anaerobic organisms, including methanogens, digest cellulose into forms usable by the animal. Without these microorganisms, animals such as cattle would not be able to consume grass. The useful products of methanogenesis are absorbed by the gut, but methane is released from the animal mainly by belching (eructation). The average cow emits around 250 liters of methane per day.

In humans

Some humans produce flatus that contains methane. In one study of the feces
Feces
Feces, faeces, or fæces is a waste product from an animal's digestive tract expelled through the anus or cloaca during defecation.-Etymology:...

 of nine adults, only five of the samples contained archaea
Archaea
The Archaea are a group of single-celled microorganisms. A single individual or species from this domain is called an archaeon...

 capable of producing methane. Similar results are found in samples of gas obtained from within the rectum
Rectum
The rectum is the final straight portion of the large intestine in some mammals, and the gut in others, terminating in the anus. The human rectum is about 12 cm long...

.

Even among humans whose flatus does contain methane, the amount is only 0% - 10% of the total amount of gas.

Role in global warming

Methane in the Earth's atmosphere is an important greenhouse gas
Greenhouse gas
A greenhouse gas is a gas in an atmosphere that absorbs and emits radiation within the thermal infrared range. This process is the fundamental cause of the greenhouse effect. The primary greenhouse gases in the Earth's atmosphere are water vapor, carbon dioxide, methane, nitrous oxide, and ozone...

 with a global warming potential
Global warming potential
Global-warming potential is a relative measure of how much heat a greenhouse gas traps in the atmosphere. It compares the amount of heat trapped by a certain mass of the gas in question to the amount of heat trapped by a similar mass of carbon dioxide. A GWP is calculated over a specific time...

 21 times greater than carbon dioxide (averaged over 100 years), and methanogenesis in livestock and the decay of organic material is thus a considerable contributor to global warming. It may not be a net contributor in the sense that it works on organic material which used up atmospheric carbon dioxide when it was created, but its overall effect is to convert the carbon dioxide into methane which is a much more potent greenhouse gas.

Methanogenesis can also be beneficially exploited, to treat organic waste, to produce useful compounds, and the methane can be collected and used as biogas
Biogas
Biogas typically refers to a gas produced by the biological breakdown of organic matter in the absence of oxygen. Organic waste such as dead plant and animal material, animal dung, and kitchen waste can be converted into a gaseous fuel called biogas...

, a fuel.
It is the primary pathway whereby most organic matter disposed of via landfill
Landfill
A landfill site , is a site for the disposal of waste materials by burial and is the oldest form of waste treatment...


is broken down.

Methanogenesis and extra-terrestrial life

The presence of atmospheric methane
Atmospheric methane
Atmospheric methane levels are of interest due to its impact on climate change. Atmospheric methane is one of the most potent and influential greenhouse gases on Earth. The 100-year global warming potential of methane is 25, i.e...

 has a role in the scientific search for extra-terrestrial life. The argument being that methane in the atmosphere will eventually dissipate, unless something is replenishing it. This can be detected (by using a spectrometer
Spectrometer
A spectrometer is an instrument used to measure properties of light over a specific portion of the electromagnetic spectrum, typically used in spectroscopic analysis to identify materials. The variable measured is most often the light's intensity but could also, for instance, be the polarization...

 for example) then that means there is, or relatively recently was, life present.
This was debated when methane was discovered in the Martian atmosphere by M.J. Mumma of NASA's Goddard Flight Center, and verified by the Mars Express Orbiter (2004) and in Titan
Titan (moon)
Titan , or Saturn VI, is the largest moon of Saturn, the only natural satellite known to have a dense atmosphere, and the only object other than Earth for which clear evidence of stable bodies of surface liquid has been found....

's atmosphere by the Huygens probe
Huygens probe
The Huygens probe was an atmospheric entry probe carried to Saturn's moon Titan as part of the Cassini–Huygens mission. The probe was supplied by the European Space Agency and named after the Dutch 17th century astronomer Christiaan Huygens....

 (2005). It is also argued that atmospheric methane
Atmospheric methane
Atmospheric methane levels are of interest due to its impact on climate change. Atmospheric methane is one of the most potent and influential greenhouse gases on Earth. The 100-year global warming potential of methane is 25, i.e...

 can come from volcanoes or other fissures in the planet's crust and that without an Isotopic signature
Isotopic signature
An isotopic signature is a ratio of stable or unstable isotopes of particular elements found in an investigated material...

it is difficult to say what exactly was the origin.
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