Commamox
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Comammox (COMplete AMMonia OXidation) is the name attributed to an organism that can convert
ammonia Ammonia is an inorganic chemical compound of nitrogen and hydrogen with the chemical formula, formula . A Binary compounds of hydrogen, stable binary hydride and the simplest pnictogen hydride, ammonia is a colourless gas with a distinctive pu ...
into
nitrite The nitrite polyatomic ion, ion has the chemical formula . Nitrite (mostly sodium nitrite) is widely used throughout chemical and pharmaceutical industries. The nitrite anion is a pervasive intermediate in the nitrogen cycle in nature. The name ...
and then into
nitrate Nitrate is a polyatomic ion with the chemical formula . salt (chemistry), Salts containing this ion are called nitrates. Nitrates are common components of fertilizers and explosives. Almost all inorganic nitrates are solubility, soluble in wa ...
through the process of
nitrification ''Nitrification'' is the biological oxidation of ammonia to nitrate via the intermediary nitrite. Nitrification is an important step in the nitrogen cycle in soil. The process of complete nitrification may occur through separate organisms or ent ...
. Nitrification has traditionally been thought to be a two-step process, where ammonia-oxidizing bacteria and archaea oxidize ammonia to nitrite and then nitrite-oxidizing bacteria convert to nitrate. Complete conversion of ammonia into nitrate by a single
microorganism A microorganism, or microbe, is an organism of microscopic scale, microscopic size, which may exist in its unicellular organism, single-celled form or as a Colony (biology)#Microbial colonies, colony of cells. The possible existence of unseen ...
was first predicted in 2006. In 2015 the presence of microorganisms that could carry out both conversion processes was discovered within the genus ''
Nitrospira ''Nitrospira'' (from Latin: nitro, meaning "nitrate" and Greek: spira, meaning "spiral") is a genus of bacteria within the monophyletic clade of the Nitrospirota phylum. The first member of this genus was described 1986 by Watson et al., isola ...
,'' and the
nitrogen cycle The nitrogen cycle is the biogeochemical cycle by which nitrogen is converted into multiple chemical forms as it circulates among atmosphere, atmospheric, terrestrial ecosystem, terrestrial, and marine ecosystems. The conversion of nitrogen can ...
was updated. Within the genus ''Nitrospira,'' the major ecosystems comammox are primarily found in are natural aquifers and engineered ecosystems. Complete nitrification step yield more energy (∆G°′ = −349 kJ mol−1 NH3) than either single oxidation alone (∆G°′ = −275 kJ mol−1 NH3 for ammonia oxidation to nitrite and ∆G°′ = −74 kJ mol−1 NO2 for nitrite oxidation to nitrate).


Comammox ''Nitrospira'' bacteria

Complete nitrification of oxidizing ammonia to nitrate is energetically advantageous for ''
Nitrospira ''Nitrospira'' (from Latin: nitro, meaning "nitrate" and Greek: spira, meaning "spiral") is a genus of bacteria within the monophyletic clade of the Nitrospirota phylum. The first member of this genus was described 1986 by Watson et al., isola ...
''. Due to the previous research done on ''Nitrospira,'' it was thought that all ''Nitrospira'' use nitrite as their energy source. Therefore, comammox ''Nitrospira'' were not discovered until 2015. All discovered nitrifiers belong to sublineage II of the genus ''Nitrospira.'' The
genome A genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as ...
of the nitrifying
chemolithoautotrophic A lithoautotroph is an organism that derives energy from reactions of reduced compounds of mineral (inorganic) origin. Two types of lithoautotrophs are distinguished by their energy source; photolithoautotrophs derive their energy from light, whil ...
bacterium from the genus ''Nitrospira'' encodes for both ammonia and nitrite oxidation. The genes associated with the growth by ammonia oxidation to nitrate are
ammonia monooxygenase Ammonia monooxygenase (, ''AMO'') is an enzyme, which catalysis, catalyses the following chemical reaction : ammonia + AH2 + O2 \rightleftharpoons NH2OH + A + H2O Ammonia monooxygenase contains copper and possibly nonheme iron. AMO is the first ...
and hydroxylamine dehydrogenases genes (e.g. amoA gene and hao cluster). This shows that complete nitrifying ''Nitrospira'' serve as cornerstones of the nitrogen-cycling microbial communities found in the environment. Nearly two years after the discovery of comammox organisms, ''
Nitrospira inopinata ''Nitrospira inopinata'' is a bacterium from the phylum Nitrospirota. This phylum contains nitrite-oxidizing bacteria playing role in nitrification. However ''N. inopinata'' was shown to perform complete ammonia oxidation to nitrate thus being th ...
'' was the first complete nitrifier to be isolated in pure culture. Kinetic and physiological analysis of ''Nitrospira inopinata'' demonstrated that this complete nitrifier has a high affinity for ammonia, slow growth rate, low maximum rate of ammonia oxidation, and high yield. The discovery of comammox ''Nitrospira'' provides a view into the modular evolution of the
nitrogen cycle The nitrogen cycle is the biogeochemical cycle by which nitrogen is converted into multiple chemical forms as it circulates among atmosphere, atmospheric, terrestrial ecosystem, terrestrial, and marine ecosystems. The conversion of nitrogen can ...
and expands upon the complexity of the evolutionary history of
nitrification ''Nitrification'' is the biological oxidation of ammonia to nitrate via the intermediary nitrite. Nitrification is an important step in the nitrogen cycle in soil. The process of complete nitrification may occur through separate organisms or ent ...
.


Ecosystem of comammox

Comammox have been identified in many ecosystems, including natural
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. The term excludes seawater and brackish water, but it does include non-salty mi ...
and terrestrial ecosystems. Notably commamox genes were not found to be abundant in oceans. Additionally, the use of engineered ecosystems for comammox could be used for
ammonium Ammonium is a modified form of ammonia that has an extra hydrogen atom. It is a positively charged (cationic) polyatomic ion, molecular ion with the chemical formula or . It is formed by the protonation, addition of a proton (a hydrogen nucleu ...
removal during water and
wastewater treatment Wastewater treatment is a process which removes and eliminates contaminants from wastewater. It thus converts it into an effluent that can be returned to the water cycle. Once back in the water cycle, the effluent creates an acceptable impact on ...
. Comammox have been found in many engineered systems including
aquaculture Aquaculture (less commonly spelled aquiculture), also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants (e.g. Nelu ...
biofiltration Biofiltration is a pollution control technique using a bioreactor containing living material to capture and biologically degrade pollutants. Common uses include processing waste water, capturing harmful chemicals or silt from surface runoff, and ...
units, drinking water treatment and distribution systems, and wastewater treatment plants. The growth of comammox in these engineered ecosystems co-occur with
ammonia-oxidizing bacteria Nitrifying bacteria are chemolithotrophic organisms that include species of genera such as ''Nitrosomonas'', ''Nitrosococcus'', ''Nitrobacter'', '' Nitrospina'', ''Nitrospira'' and '' Nitrococcus''. These bacteria get their energy from the oxidati ...
and/or archaea, and in some cases outnumber other ammonia-oxidizing prokaryotes. The ecosystem of comammox is currently unknown in terms of
biogeography Biogeography is the study of the species distribution, distribution of species and ecosystems in geography, geographic space and through evolutionary history of life, geological time. Organisms and biological community (ecology), communities o ...
, including their distribution and abundance, due to the influences of process configuration and chemical composition of the treated wastewater. Following these findings, it was determined that comammox may out-select canonical nitrite oxidizing bacteria in the genus ''
Nitrospira ''Nitrospira'' (from Latin: nitro, meaning "nitrate" and Greek: spira, meaning "spiral") is a genus of bacteria within the monophyletic clade of the Nitrospirota phylum. The first member of this genus was described 1986 by Watson et al., isola ...
'' in some engineered environments, suggesting the potentially important role for comammox in efficient biological nitrogen removal in wastewater treatment processes.


See also

*
Anammox Anammox, an abbreviation for "anaerobic ammonium oxidation", is a globally important microbial process of the nitrogen cycle that takes place in many natural environments. The bacteria mediating this process were identified in 1999, and were a gr ...


References

{{reflist Nitrogen cycle