Nitrifying bacteria

 Nitrifying_bacteria

Nitrifying_bacteriaNitrifying bacteria are chemolithotrophic organisms that include species of the genera e.g. Nitrosomonas, Nitrosococcus, Nitrobacter, Nitrospina, Nitrospira and Nitrococcus. These bacteria get their energy by the oxidation of inorganic nitrogen compounds.[1] Types include ammonia-oxidizing bacteria (AOB) and nitrite-oxidizing bacteria (NOB). Many species of nitrifying bacteria have complex internal membrane systems that are the location for key enzymes in nitrification: ammonia monooxygenase (which oxidizes ammonia to hydroxylamine), hydroxylamine oxidoreductase (which oxidizes hydroxylamine to nitric oxide - which is oxidized to nitrite by a currently unidentified enzyme), and nitrite oxidoreductase (which oxidizes nitrite to nitrate).[2]

EcologyEdit

Nitrifying bacteria are present in distinct taxonomical groups and are found in highest numbers where considerable amounts of ammonia are present (areas with extensive protein decomposition, and sewage treatment plants).^[3] Nitrifying bacteria thrive in lakes and rivers streams with high inputs and outputs of sewage and wastewater and freshwater because of the high ammonia content.

Oxidation of ammonia to nitrateEdit

Nitrification in nature is a two-step oxidation process of ammonium (NH₄⁺) or ammonia (NH₃) to nitrite (NO₂⁻) and then to nitrate (NO₃⁻) catalyzed by two ubiquitous bacterial groups growing together. The first reaction is oxidation of ammonium to nitrite by ammonia oxidizing bacteria (AOB) represented by members of Betaproteobacteria and Gammaproetobacteria. Further organisms able to oxidize ammonia are Archaea (AOA).^[4]

The second reaction is oxidation of nitrite (NO₂⁻) to nitrate by nitrite-oxidizing bacteria (NOB), represented by the members of Nitrospinae, Nitrospirae, Proteobacteria and Chloroflexi.^[5][6]

This two-step process was described already in 1890 by the Russian microbiologist Sergei Winogradsky.

Ammonia can be also oxidized completely to nitrate by one comammox bacterium.

First step nitrification - molecular mechanismEdit

Figure 1. Molecular mechanism of ammonium oxidation by AOB

Ammonia oxidation in autotrophic nitrification is a complex process that requires several enzymes, proteins and presence of oxygen. The key enzymes, necessary to obtaining energy during oxidation of ammonia to nitrite are ammonia monooxygenase (AMO) and hydroxylamine oxidoreductase (HAO). First is a transmembrane copper protein which catalyzes the oxidation of ammonia to hydroxylamine (1.1) taking two electrons directly from the quinone pool. This reaction requires O₂.

The second step of this process has recently fallen into question.^[7]

For the past few decades, the common view was that a trimeric multiheme c-type HAO converts hydroxylamine into nitrite in the periplasm with production of four electrons (1.2). The stream of four electron are channelled through cytochrome c₅₅₄ to a membrane-bound cytochrome c₅₅₂. Two of the electrons are routed back to AMO, where they are used for the oxidation of ammonia (quinol pool). The remaining two electrons are used to generate a proton motive force and reduce NAD(P) through reverse electron transport.^[8]

Recent results, however, show that HAO does not produce nitrite as a direct product of catalysis. This enzyme instead produces nitric oxide and three electrons. Nitric oxide can then be oxidized by other enzymes (or oxygen) to nitrite. In this paradigm, the electron balance for overall metabolism needs to be reconsidered.^[7]

NH₃ + O₂ → NO−
2 + 3H⁺ + 2e⁻ (1)

NH₃ + O₂ + 2H⁺ + 2e⁻ → NH₂OH + H
2O (1.1)

NH₂OH + H
2O → NO−
2 + 5H⁺ + 4e⁻ (1.2)

Second step nitrification - molecular mechanismEdit

Nitrite produced in the first step of autotrophic nitrification is oxidized to nitrate by nitrite oxidoreductase (NXR)(2). It is a membrane-associated iron-sulfur molybdoprotein, and is part of an electron transfer chain which channels electrons from nitrite to molecular oxygen.^{[citation needed]} The molecular mechanism of oxidation nitrite is less described than oxidation ammonium. In new research (e.g. Woźnica A. et al., 2013)^[9] proposed new hypothetical model of NOB electron transport chain and NXR mechanism (Figure 2.). In contrast to earlier models ^[10] the NXR acts on the outside of the plasma membrane, directly contributing to postulated by Spieck ^[11] and coworkers mechanism of proton gradient generation. Nevertheless, the molecular mechanism of nitrite oxidation is an open question.

NO−
2 + H
2O → NO−
3 + 2H⁺ + 2e⁻ (2)

Comammox bacteriaEdit

Main page: commamox

Characteristic of nitrifying bacteriaEdit

Nitrifying bacteria that oxidize ammonia ^[5][12]Edit

Genus	Phylogenetic group	DNA (mol% GC)	Habitats	Characteristics
Nitrosomonas	Beta	45-53	Soil, Sewage, freshwater, Marine	Gram-negative short to long rods, motile (polar flagella)or nonmotile; peripheral membrane systems
Nitrosococcus	Gamma	49-50	Freshwater, Marine	Large cocci, motile, vesicular or peripheral membranes
Nitrosospira	Beta	54	Soil	Spirals, motile (peritrichous flagella); no obvious membrane system

Nitrifying bacteria that oxidize nitrite ^[5][12]Edit

Genus	Phylogenetic group	DNA (mol% GC)	Habitats	Characteristics
Nitrobacter	Alpha	59-62	Soil, Freshwater, Marine	Short rods, reproduce by budding, occasionally motile (single subterminal flagella) or non-motile; membrane system arranged as a polar cap
Nitrospina	Delta	58	Marine	Long, slender rods, nonmotile, no obvious membrane system
Nitrococcus	Gamma	61	Marine	Large Cocci, motile (one or two subterminal flagellum) membrane system randomly arranged in tubes
Nitrospira	Nitrospirae	50	Marine, Soil	Helical to vibroid-shaped cells; nonmotile; no internal membranes

Comammox bacteria^[13][14][15]Edit

Species	Phylogenetic group	DNA (mol% GC)	Habitats	Characteristics
Nitrospira inopinata	Nitrospirae	59.23	microbial mat in hot water pipes (56 °C, pH 7.5)	Rods

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