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 called physolmere; it is respiration without oxygen.[1]

In aerobic organisms undergoing respiration, electrons are shuttled to an electron transport chain, and the final electron acceptor is oxygen. Molecular oxygen is a highly oxidizing agent and, therefore, is an excellent acceptor. In anaerobes, other less-oxidizing substances such as sulfate (SO42−), nitrate (NO3−), sulphur (S), or fumarate are used. These terminal electron acceptors have smaller reduction potentials than O2, meaning that less energy is released per oxidized molecule. Anaerobic respiration is, therefore, in general energetically less efficient than aerobic respiration.

Anaerobic respiration is used mainly by bacteria and archaea that live in environments devoid of oxygen. Many anaerobic organisms are obligate anaerobes meaning that they can respire only using anaerobic compounds and will die in the presence of oxygen.

Anaerobic respiration as compared with fermentation

Cellular respiration (both aerobic and anaerobic) utilizes 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, resulting 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 or pmf 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 utilize 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+. The two processes thus generate ATP in very different ways, and the terms should not be treated as synonyms.

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.[2] Another example is methanogenesis, a form of carbonate 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.[3] Sulfate respiration produces hydrogen sulfide, which is responsible for the characteristic 'rotten egg' smell of coast wetlands and has the capacity to precipitate heavy metal ions from solution, leading to the deposition of sulfidic metal ores.[4]

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.[5]

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.[6]

Examples of respiration

Type Lifestyle Electron acceptor Products Eo' [V] Example organisms
aerobic respiration obligate aerobes and facultative anaerobes O2 H2O + 0.82 eukaryotes and aerobic prokaryotes
iron reduction facultative anaerobes and obligate anaerobes Fe(III) Fe(II) + 0.75 Organisms within the order Desulfuromonadales (such as Geobacter, Geothermobacter, Geopsychrobacter, Pelobacter) and Shewanella species [7]
manganese facultative anaerobes and obligate anaerobes Mn(IV) Mn(II) Desulfuromonadales and Shewanella species [7]
cobalt reduction facultative anaerobes and obligate anaerobes Co(III) Co(II) Geobacter sulfurreducens
uranium reduction facultative anaerobes and obligate anaerobes U(VI) U(IV) Geobacter metallireducens, Shewanella oneidensis
nitrate reduction (denitrification) facultative anaerobes nitrate NO3− nitrite NO2− + 0.40 Paracoccus denitrificans, Escherichia coli
fumarate respiration facultative anaerobes fumarate succinate + 0.03 Escherichia coli
sulfate respiration obligate anaerobes sulfate SO42− sulfide HS− - 0.22 Many Gammaproteobacteria species in the orders Desulfobacterales, Desulfovibrionales, and Syntrophobacterales
methanogenesis (carbonate reduction) methanogens carbon dioxide CO2 methane CH4 - 0.25 Methanosarcina barkeri
sulfur respiration (sulfur reduction) facultative anaerobes and obligate anaerobes sulfur S0 sulfide HS− - 0.27 Desulfuromonadales
acetogenesis (carbonate reduction) obligate anaerobes carbon dioxide CO2 acetate - 0.30 Acetobacterium woodii
dehalorespiration facultative anaerobes and obligate anaerobes halogenated organic compounds R-X Halide ions and dehalogenated compound X− + R-H + 0.25–+ 0.60[8] Dehalococcoides and Dehalobacter species

See also

References

  1. ↑ Slonczewski, Joan L.; Foster, John W. (2011). Microbiology : An Evolving Science (2nd ed.). New York: W.W. Norton. p. 166. ISBN 9780393934472.
  2. ↑ Simon, Jörg; Klotz, Martin G. (2013-02-01). "Diversity and evolution of bioenergetic systems involved in microbial nitrogen compound transformations". Biochimica et Biophysica Acta (BBA) - Bioenergetics. The evolutionary aspects of bioenergetic systems. 1827 (2): 114–135. doi:10.1016/j.bbabio.2012.07.005.
  3. ↑ Bogner, Jean; Pipatti, Riitta; Hashimoto, Seiji; Diaz, Cristobal; Mareckova, Katarina; Diaz, Luis; Kjeldsen, Peter; Monni, Suvi; Faaij, Andre (2008-02-01). "Mitigation of global greenhouse gas emissions from waste: conclusions and strategies from the Intergovernmental Panel on Climate Change (IPCC) Fourth Assessment Report. Working Group III (Mitigation)". Waste management & research: the journal of the International Solid Wastes and Public Cleansing Association, ISWA. 26 (1): 11–32. doi:10.1177/0734242x07088433. ISSN 0734-242X. PMID 18338699.
  4. ↑ Pester, Michael; Knorr, Klaus-Holger; Friedrich, Michael W.; Wagner, Michael; Loy, Alexander (2012-01-01). "Sulfate-reducing microorganisms in wetlands - fameless actors in carbon cycling and climate change". Frontiers in Microbiology. 3: 72. doi:10.3389/fmicb.2012.00072. ISSN 1664-302X. PMC 3289269Freely accessible. PMID 22403575.
  5. ↑ Nancharaiah, Y. V.; Venkata Mohan, S.; Lens, P. N. L. (2016-09-01). "Recent advances in nutrient removal and recovery in biological and bioelectrochemical systems". Bioresource Technology. 215: 173–185. doi:10.1016/j.biortech.2016.03.129. ISSN 1873-2976. PMID 27053446.
  6. ↑ Xu, Bojun; Ge, Zheng; He, Zhen (2015-05-15). "Sediment microbial fuel cells for wastewater treatment: challenges and opportunities". Environ. Sci.: Water Res. Technol. 1 (3): 279–284. doi:10.1039/c5ew00020c. ISSN 2053-1419.
  7. 1 2 Richter, Katrin; Schicklberger, Marcus; Gescher, Johannes (2012-02-01). "Dissimilatory reduction of extracellular electron acceptors in anaerobic respiration". Applied and Environmental Microbiology. 78 (4): 913–921. doi:10.1128/AEM.06803-11. ISSN 1098-5336. PMC 3273014Freely accessible. PMID 22179232.
  8. ↑ Holliger, C.; Wohlfarth, G.; Diekert, G. (1998). "Reductive dechlorination in the energy metabolism of anaerobic bacteria". FEMS Microbiology Reviews. 22 (5): 383. doi:10.1111/j.1574-6976.1998.tb00377.x.
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