Denitrification

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Nitrogen cycle.

Denitrification is a microbially facilitated process where nitrate (NO3−) is reduced

and ultimately produces molecular nitrogen (N2) through a series of intermediate
gaseous nitrogen oxide products. Facultative anaerobic bacteria perform
denitrification as a type of respiration that reduces oxidized forms of nitrogen in
response to the oxidation of an electron donor such as organic matter. The preferred
nitrogen electron acceptors in order of most to least thermodynamically favorable
include nitrate (NO3−), nitrite (NO2−), nitric oxide (NO), nitrous oxide (N2O) finally
resulting in the production of dinitrogen (N2) completing the nitrogen cycle.
Denitrifying microbes require a very low oxygen concentration of less than 10%, as
well as organic C for energy. Since denitrification can remove NO3−, reducing
its leaching to groundwater, it can be strategically used to treat sewage or animal
residues of high nitrogen content. Denitrification can leak N 2O, which is an ozone-
depleting substance and a greenhouse gas that can have a considerable influence
on global warming.
The process is performed primarily by heterotrophic bacteria (such as Paracoccus
denitrificans and various pseudomonads),[1] although autotrophic denitrifiers have
also been identified (e.g., Thiobacillus denitrificans).[2] Denitrifiers are represented in
all main phylogenetic groups.[3] Generally several species of bacteria are involved in
the complete reduction of nitrate to N2, and more than one enzymatic pathway has
been identified in the reduction process.[4] The denitrification process does not only
provide energy to the organism performing nitrate reduction to dinitrogen gas, but
also some anaerobic ciliates can use denitrifying endosymbionts to gain energy
similar to the use of mitochondria in oxygen respiring organisms. [5]
Direct reduction from nitrate to ammonium, a process known as dissimilatory
nitrate reduction to ammonium or DNRA,[6] is also possible for organisms that have
the nrf-gene.[7][8] This is less common than denitrification in most ecosystems as a
means of nitrate reduction. Other genes known in microorganisms which denitrify
 include nir (nitrite reductase) and nos (nitrous oxide reductase) among others;
[3]
 organisms identified as having these genes include Alcaligenes
faecalis, Alcaligenes xylosoxidans, many in the
genus Pseudomonas, Bradyrhizobium japonicum, and Blastobacter denitrificans.[9]

Contents

 1Overview
o 1.1Half reactions
o 1.2Conditions of denitrification
 2Use in wastewater treatment
 3See also
 4References

Overview[edit]
Half reactions[edit]
Denitrification generally proceeds through some combination of the following half
reactions, with the enzyme catalyzing the reaction in parentheses:

 NO3− + 2 H+ + 2 e−→ NO

2  + H2O (Nitrate reductase)
−

 NO
2  + 2 H  + e  → NO + H2O (Nitrite reductase)
− + −

 2 NO + 2 H+ + 2 e− → N
2O + H2O (Nitric oxide reductase)

 N
2O + 2 H  + 2 e  → N
+ −

2 + H2O (Nitrous oxide reductase)

The complete process can be expressed as a net balanced redox reaction,

where nitrate (NO3−) gets fully reduced to dinitrogen (N2):

 2 NO3− + 10 e− + 12 H+ → N2 + 6 H2O

Conditions of denitrification[edit]
In nature, denitrification can take place in both terrestrial and marine ecosystems.
[10]
 Typically, denitrification occurs in anoxic environments, where the concentration of
dissolved and freely available oxygen is depleted. In these areas, nitrate (NO 3−) or
nitrite (NO
2 ) can be used as a substitute terminal electron acceptor instead of oxygen (O2), a
−

more energetically favourable electron acceptor. Terminal electron acceptor is a

compound that gets reduced in the reaction by receiving electrons. Examples of
anoxic environments can include soils,[11] groundwater,[12] wetlands, oil reservoirs,
[13]
 poorly ventilated corners of the ocean and seafloor sediments.
Furthermore, denitrification can occur in oxic environments as well. High activity of
denitrifiers can be observed in the intertidal zones, where the tidal cycles cause
fluctuations of oxygen concentration in sandy coastal sediments. [14] For example, the
bacterial species Paracoccus denitrificans engages in denitrification under both oxic
and anoxic conditions simultaneously. Upon oxygen exposure, the bacteria is able to
utilize nitrous oxide reductase, an enzyme that catalyzes the last step of
denitrification.[15] Aerobic denitrifiers are mainly Gram-negative bacteria in the phylum
Proteobacteria. Enzymes NapAB, NirS, NirK and NosZ are located in the periplasm,
a wide space bordered by the cytoplasmic and the outer membrane in Gram-
negative bacteria.[16]
Denitrification can lead to a condition called isotopic fractionation in the soil
environment. The two stable isotopes of nitrogen, 14N and 15N are both found in the
sediment profiles. The lighter isotope of nitrogen, 14N, is preferred during
denitrification, leaving the heavier nitrogen isotope, 15N, in the residual matter. This
selectivity leads to the enrichment of 14N in the biomass compared to 15N.[17] Moreover,
the relative abundance of 14N can be analyzed to distinguish denitrification apart from
other processes in nature.

Use in wastewater treatment[edit]

Further information: Sewage treatment

Denitrification is commonly used to remove nitrogen from sewage and

municipal wastewater. It is also an instrumental process in constructed
wetlands[18] and riparian zones[19] for the prevention of groundwater pollution with
nitrate resulting from excessive agricultural or residential fertilizer usage.[20] Wood
chip bioreactors have been studied since the 2000s and are effective in removing
nitrate from agricultural run off[21] and even manure.[22]
Reduction under anoxic conditions can also occur through process called anaerobic
ammonium oxidation (anammox):[23]
NH4+ + NO2− → N2 + 2 H2O
In some wastewater treatment plants, compounds such
as methanol, ethanol, acetate, glycerin, or proprietary products are added to the
wastewater to provide a carbon and electron source for denitrifying bacteria.
[24]
 The microbial ecology of such engineered denitrification processes is
determined by the nature of the electron donor and the process operating
conditions.[25][26] Denitrification processes are also used in the treatment
of industrial wastewater.[27] Many denitrifying bioreactor types and designs are
available commercially for the industrial applications, including Electro-
Biochemical Reactors (EBRs), membrane bioreactors (MBRs), and moving bed
bioreactors (MBBRs).
Aerobic denitrification, conducted by aerobic denitrifiers, may offer the potential
to eliminate the need for separate tanks and reduce sludge yield. There are less
stringent alkalinity requirements because alkalinity generated during
denitrification can partly compensate for the alkalinity consumption in nitrification