Topic: 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 respiration, electrons are shuttled to
an electron transport chain, and the final electron acceptor is oxygen. Molecular oxygen is a
high-energy oxidizing agent and, therefore, is an excellent electron acceptor. In anaerobes,
other less-oxidizing substances such as nitrate (NO−3), fumarate (C4H2O2−4), sulfate
(SO2−4), or sulfur (S) are used. These terminal electron acceptors have smaller reduction
potentials than O2, meaning that less energy is released per oxidized molecule. Therefore,
anaerobic respiration is less efficient than aerobic.

As compared with fermentation

Anaerobic cellular respiration and fermentation generate ATP in very different ways, and the
terms should not be treated as synonyms. Cellular respiration (both aerobic and anaerobic)
uses highly reduced chemical compounds such as NADH and FADH2 (for example produced
during glycolysis and the citric acid cycle) to establish an electrochemical gradient (often a
proton gradient) across a membrane. This results in an electrical potential or ion
concentration difference across the membrane. The reduced chemical compounds are
oxidized by a series of respiratory integral membrane proteins with sequentially increasing
reduction potentials, with the final electron acceptor being oxygen (in aerobic respiration) or
another chemical substance (in anaerobic respiration). A proton motive force drives protons
down the gradient (across the membrane) through the proton channel of ATP synthase. The
resulting current drives ATP synthesis from ADP and inorganic phosphate.

Fermentation, in contrast, does not use an electrochemical gradient. Fermentation instead

only uses substrate-level phosphorylation to produce ATP. The electron acceptor NAD+ is
regenerated from NADH formed in oxidative steps of the fermentation pathway by the
reduction of oxidized compounds. These oxidized compounds are often formed during the
fermentation pathway itself, but may also be external. For example, in homofermentative
lactic acid bacteria, NADH formed during the oxidation of glyceraldehyde-3-phosphate is
oxidized back to NAD+ by the reduction of pyruvate to lactic acid at a later stage in the
pathway. In yeast, acetaldehyde is reduced to ethanol to regenerate NAD+.

There are two important anaerobic microbial methane formation pathways, through carbon
dioxide / bicarbonate (HCO−3) reduction (respiration) or acetate fermentation.

Ecological importance
Anaerobic respiration is a critical component of the global nitrogen, iron, sulfur, and carbon
cycles through the reduction of the oxyanions of nitrogen, sulfur, and carbon to more-reduced
compounds. The biogeochemical cycling of these compounds, which depends upon anaerobic
respiration, significantly impacts the carbon cycle and global warming. Anaerobic respiration
occurs in many environments, including freshwater and marine sediments, soil, subsurface
aquifers, deep subsurface environments, and biofilms. Even environments, such as soil, that
contain oxygen also have micro-environments that lack oxygen due to the slow diffusion
characteristics of oxygen gas.
An example of the ecological importance of anaerobic respiration is the use of nitrate as a
terminal electron acceptor, or dissimilatory denitrification, which is the main route by which
fixed nitrogen is returned to the atmosphere as molecular nitrogen gas. The denitrification
process is also very important in host-microbe interactions. Similar to mitochondria in
oxygen-respiring microorganisms, some single-cellular anaerobic ciliates use denitrifying
endosymbionts to gain energy. Another example is methanogenesis, a form of carbon-dioxide
respiration, that is used to produce methane gas by anaerobic digestion. Biogenic methane is
used as a sustainable alternative to fossil fuels. On the negative side, uncontrolled
methanogenesis in landfill sites releases large volumes of methane into the atmosphere,
where it acts as a powerful greenhouse gas. Sulfate respiration produces hydrogen sulfide,
which is responsible for the characteristic 'rotten egg' smell of coastal wetlands and has the
capacity to precipitate heavy metal ions from solution, leading to the deposition of sulfidic
metal ores.

Economic relevance
Dissimilatory denitrification is widely used in the removal of nitrate and nitrite from
municipal wastewater. An excess of nitrate can lead to eutrophication of waterways into
which treated water is released. Elevated nitrite levels in drinking water can lead to problems
due to its toxicity. Denitrification converts both compounds into harmless nitrogen gas.

Specific types of anaerobic respiration are also critical in bioremediation, which uses
microorganisms to convert toxic chemicals into less-harmful molecules to clean up
contaminated beaches, aquifers, lakes, and oceans. For example, toxic arsenate or selenate
can be reduced to less toxic compounds by various anaerobic bacteria via anaerobic
respiration. The reduction of chlorinated chemical pollutants, such as vinyl chloride and
carbon tetrachloride, also occurs through anaerobic respiration.

Anaerobic respiration is useful in generating electricity in microbial fuel cells, which employ
bacteria that respire solid electron acceptors (such as oxidized iron) to transfer electrons from
reduced compounds to an electrode. This process can simultaneously degrade organic carbon
waste and generate electricity.

Examples of electron acceptors in respiration

See also
Hydrogenosomes and mitosomes

Anaerobic digestion

Microbial fuel cell

Standard electrode potential (data page)

Table of standard reduction potentials for half-reactions important in biochemistry

Lithotrophs
Further reading
Gregory, Kelvin B.; Bond, Daniel R.; Lovley, Derek R. (June 2004). "Graphite electrodes as
electron donors for anaerobic respiration". Environmental Microbiology. 6 (6): 596–604.
doi:10.1111/j.1462-2920.2004.00593.x. ISSN 1462-2912. PMID 15142248.

References