Anaerobic Respiration

Electron Donors and Acceptors in Anaerobic Respiration

In anaerobic respiration, a molecule other than oxygen is used as the terminal electron acceptor in the
electron transport chain

Anaerobic respiration is the formation of ATP without oxygen. This method still incorporates the
respiratory electron transport chain, but without using oxygen as the terminal electron acceptor.
Instead, molecules such as sulfate (SO42-), nitrate (NO3–), or sulfur (S) are used as electron acceptors.
These molecules have a lower reduction potential than oxygen; thus, less energy is formed per molecule
of glucose in anaerobic versus aerobic conditions.

Many different types of electron acceptors may be used for anaerobic respiration. Denitrification is the
utilization of nitrate (NO3−) as the terminal electron acceptor. Nitrate, like oxygen, has a high reduction
potential. This process is widespread, and used by many members of Proteobacteria. Many denitrifying
bacteria can also use ferric iron (Fe3+) and different organic electron acceptors.

Sulfate reduction uses sulfate (SO2−4) as the electron acceptor, producing hydrogen sulfide (H2S) as a
metabolic end product. Sulfate reduction is a relatively energetically poor process, and is used by many
Gram negative bacteria found within the δ-Proteobacteria. It is also used in Gram-positive organisms
related to Desulfotomaculum or the archaeon Archaeoglobus.

Sulfate reduction requires the use of electron donors, such as the carbon compounds lactate and
pyruvate (organotrophic reducers), or hydrogen gas (lithotrophic reducers). Some unusual autotrophic
sulfate-reducing bacteria, such as Desulfotignum phosphitoxidans, can use phosphite (HPO3–) as an
electron donor. Others, such as certain Desulfovibrio species, are capable of sulfur disproportionation
(splitting one compound into an electron donor and an electron acceptor) using elemental sulfur (S0),
sulfite (SO3−2), and thiosulfate (S2O32-) to produce both hydrogen sulfide (H2S) and sulfate (SO2−).

Acetogenesis is a type of microbial metabolism that uses hydrogen (H2) as an electron donor and carbon
dioxide (CO2) as an electron acceptor to produce acetate, the same electron donors and acceptors used
in methanogenesis.

Ferric iron (Fe3+) is a widespread anaerobic terminal electron acceptor used by both autotrophic and
heterotrophic organisms. Electron flow in these organisms is similar to those in electron transport,
ending in oxygen or nitrate, except that in ferric iron-reducing organisms the final enzyme in this system
is a ferric iron reductase. Since some ferric iron-reducing bacteria (e.g.G. metallireducens) can use toxic
hydrocarbons (e.g. toluene) as a carbon source, there is significant interest in using these organisms as
bioremediation agents in ferric iron contaminated aquifers.

Other inorganic electron acceptors include the reduction of Manganic ion (Mn4+) to manganous
(Mn2+), Selenate (SeO42−) to selenite (SeO32−) to selenium (Se), Arsenate (AsO43−) to arsenite
(AsO33-), and Uranyl (UO22+) to uranium dioxide (UO2)

Organic compounds may also be used as electron acceptors in anaerobic respiration. These include the
reduction of fumarate to succinate, Trimethylamine N-oxide (TMAO) to trimethylamine (TMA), and
Dimethyl sulfoxide (DMSO) to Dimethyl sulfide (DMS).

Nitrate Reduction and Denitrification

Denitrification is a type of anaerobic respiration that uses nitrate as an electron acceptor

In anaerobic respiration, denitrification utilizes nitrate (NO3–) as a terminal electron acceptor in the
respiratory electron transport chain. Denitrification is a widely used process; many facultative anaerobes
use denitrification because nitrate, like oxygen, has a high reduction potential

Denitrification is a microbially facilitated process involving the stepwise reduction of nitrate to nitrite
(NO2–) nitric oxide (NO), nitrous oxide (N2O), and, eventually, to dinitrogen (N2) by the enzymes nitrate
reductase, nitrite reductase, nitric oxide reductase, and nitrous oxide reductase. The complete
denitrification process can be expressed as a redox reaction: 2 NO3− + 10 e− + 12 H+ → N2 + 6 H2O.

Protons are transported across the membrane by the initial NADH reductase, quinones and nitrous
oxide reductase to produce the electrochemical gradient critical for respiration. Some organisms (e.g. E.
coli) only produce nitrate reductase and therefore can accomplish only the first reduction leading to the
accumulation of nitrite. Others (e.g. Paracoccus denitrificans or Pseudomonas stutzeri) reduce nitrate
completely. Complete denitrification is an environmentally significant process because some
intermediates of denitrification (nitric oxide and nitrous oxide) are significant greenhouse gases that
react with sunlight and ozone to produce nitric acid, a component of acid rain. Denitrification is also
important in biological wastewater treatment, where it can be used to reduce the amount of nitrogen
released into the environment, thereby reducing eutrophication.
Denitrification takes place under special conditions in both terrestrial and marine ecosystems. In
general, it occurs where oxygen is depleted and bacteria respire nitrate as a substitute terminal electron
acceptor. Due to the high concentration of oxygen in our atmosphere, denitrification only takes place in
anaerobic environments where oxygen consumption exceeds the oxygen supply and where sufficient
quantities of nitrate are present. These environments may include certain soils and groundwater,
wetlands, oil reservoirs, poorly ventilated corners of the ocean, and in sea floor sediments.

Denitrification is performed primarily by heterotrophic bacteria (e.g. Paracoccus denitrificans), although

autotrophic denitrifiers have also been identified (e.g., Thiobacillus denitrificans). Generally, several
species of bacteria are involved in the complete reduction of nitrate to molecular nitrogen, and more
than one enzymatic pathway have been identified in the reduction process.

Rhizobia are soil bacteria with the unique ability to establish a N2-fixing symbiosis on legume roots.
When faced with a shortage of oxygen, some rhizobia species are able to switch from O2-respiration to
using nitrates to support respiration.

The direct reduction of nitrate to ammonium (dissimilatory nitrate reduction) can be performed by
organisms with the nrf- gene. This is a less common method of nitrate reduction than denitrification in
most ecosystems. Other genes involved in denitrification include nir (nitrite reductase) and nos (nitrous
oxide reductase), which are possessed by such organisms as Alcaligenes faecalis, Alcaligenes
xylosoxidans, Pseudomonas spp, Bradyrhizobium japonicum, and Blastobacter denitrificans.