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Methanogenesis

Methanogenesis or biomethanation is the formation of methane by microbes known as methanogens. Organisms capable of
producing methane have been identified only from the domain Archaea, a group phylogenetically distinct from both eukaryotes and
bacteria, although many live in close association with anaerobic bacteria. The production of methane is an important and widespread
form of microbial metabolism. In most environments, it is the final step in the decomposition of biomass. Methanogenesis is
[1][2][3]
responsible for significant amounts of natural gas accumulations, the remainder being thermogenic.

Contents
Biochemistry
Proposed mechanism
Reverse methanogenesis
Importance in carbon cycle
Natural occurrence
In ruminants
In humans
In plants
Soils
Role in global warming
Extra-terrestrial life
See also
References

Biochemistry
Methanogenesis in microbes is a form of anaerobic respiration.[4]
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 acetic acid or inorganic carbon dioxide as
terminal electron acceptors:

CO2 + 4 H2 → CH4 + 2 H2O

CH3COOH → CH4 + CO2

During anaerobic respiration of carbohydrates, H2 and acetate are


Cycle for methanogenesis, showing intermediates.
formed in a ratio of 2:1 or lower, so H2 contributes only ca. 33%
to methanogenesis, with acetate contributing the greater
proportion. In some circumstances, for instance in the rumen, where acetate is largely absorbed into the bloodstream of the host, the
contribution of H2 to methanogenesis is greater.[5]
However, depending on pH and temperature, methanogenesis has been shown to use carbon from other small organic compounds,
such as formic acid (formate), methanol, methylamines, tetramethylammonium, dimethyl sulfide, and methanethiol. The catabolism
[4]
of the methyl compounds is mediated by methyl transferases to give methyl coenzyme M.

Proposed mechanism
The biochemistry of methanogenesis is relatively complex, involving the following coenzymes and cofactors: F420, coenzyme B,
coenzyme M, methanofuran, and methanopterin.

The mechanism for the conversion of CH3–S bond into methane involves a ternary complex of methyl coenzyme M and coenzyme B
fit into a channel terminated by the axial site on nickel of the cofactor F430. One proposed mechanism invokes electron transfer from
Ni(I) (to give Ni(II)), which initiates formation of CH4. Coupling of the coenzyme M thiyl radical (RS.) with HS coenzyme B
[6]
releases a proton and re-reduces Ni(II) by one-electron, regenerating Ni(I).

Reverse methanogenesis
Some organisms can oxidize methane, functionally reversing the process of methanogenesis, also referred to as the anaerobic
oxidation of methane (AOM). Organisms performing AOM have been found in multiple marine and freshwater environments
including methane seeps, hydrothermal vents, coastal sediments and sulfate-methane transition zones.[7] These organisms may
accomplish reverse methanogenesis using a nickel-containing protein similar to
methyl-coenzyme M reductaseused by methanogenic
archaea.[8] Reverse methanogenesis occurs according to the reaction:

SO42− + CH4 → HCO3− + HS− + H2O[9]

Importance in carbon cycle


Methanogenesis is the final step in the decay of organic matter. During the decay process, electron acceptors (such as oxygen, ferric
iron, sulfate, and nitrate) become depleted, while hydrogen (H2) and carbon dioxide accumulate. Light organics produced by
fermentation 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, agreat deal of carbon (in the form of
fermentation products) would accumulate in anaerobic environments.

Natural occurrence

In ruminants
Enteric fermentation occurs in the gut of some animals, especially ruminants. In the rumen, anaerobic organisms, including
methanogens, digest cellulose into forms nutritious to the animal. Without these microorganisms, animals such as cattle would not be
able to consume grasses. 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.[10] In this way, ruminants contribute
about 25% of anthropogenic methane emissions. One method of methane production control in ruminants is by feeding them 3-
nitrooxypropanol.[11]

In humans
Some humans produceflatus that contains methane. In one study of the feces of nine
adults, five of the samples contained archaea capable of producing methane.[12]
Similar results are found in samples of gas obtained from within therectum.

Even among humans whose flatus does contain methane, the amount is in the range
of 10% or less of the total amount of gas.[13]

In plants
Testing Australian sheep for exhaled
Many experiments have suggested that leaf tissues of living plants emit methane.[14] methane production (2001),CSIRO
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.[15]

Soils
Methanogens are observed in anoxic soil environments, contributing to the degradation of organic matter. This organic matter may be
placed by humans through landfill, buried as sediment on the bottom of lakes or oceans as sediments, and as residual organic matter
[16]
from sediments that have formed into sedimentary rocks.

Role in global warming


Atmospheric methane is an important greenhouse gas with a global warming potential25 times greater than carbon dioxide (averaged
over 100 years),[17] 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, a fuel.[18] It is the primary pathway whereby most organic matter disposed of via landfill is broken
down.[19]

Extra-terrestrial life
The presence of atmospheric methane has a role in the scientific search for extra-terrestrial life. The justification is that methane in
the atmosphere will eventually dissipate, unless something is replenishing it. If methane is detected (by using a spectrometer for
example) this may indicate that life is, or recently was, present. This was debated[20] 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)[21] and in Titan's
atmosphere by the Huygens probe (2005).[22] This debate was furthered with the discovery of 'transient', 'spikes of methane' on Mars
by the Curiosity Rover.[23]

It is also argued that atmospheric methane can come from volcanoes or other fissures in the planet's crust and that without an isotopic
signature, the origin or source may be difficult to identify.[24][25]

On 13 April 2017, NASA confirmed that the dive of the Cassini orbiter spacecraft on 28 October 2015 discovered the Enceladus
plume which has all the ingredients for methanogenesis-based life forms to feed from. Previous results, published in March 2015,
suggested hot water is interacting with rock beneath the sea; the new findings support that conclusion and add that the rock appears to
be reacting chemically. From these observations scientists have determined that nearly 98 percent of the gas in the plume is water,
[26]
about 1 percent is hydrogen and the rest is a mixture of other molecules including carbon dioxide, methane and ammonia.

See also
Aerobic methane production
Anaerobic digestion
Anaerobic oxidation of methane
Electromethanogenesis
Hydrogen cycle
Methanotroph
Mootral

References
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