Nitrogen cycle

The nitrogen cycle is the biogeochemical cycle by

which nitrogen is converted into multiple chemical forms
as it circulates among atmosphere, terrestrial, and marine
ecosystems. The conversion of nitrogen can be carried
out through both biological and physical processes.
Important processes in the nitrogen cycle include
fixation, ammonification, nitrification, and denitrification.
The majority of Earth's atmosphere (78%) is atmosphere
nitrogen,[16] making it the largest source of nitrogen.
However, atmospheric nitrogen has limited availability
for biological use, leading to a scarcity of usable nitrogen
This figure summarizes the global cycling of
in many types of ecosystems.
reactive nitrogen.[1] includes industrial fertilizer
The nitrogen cycle is of particular interest to ecologists production,[2] nitrogen fixed by natural
because nitrogen availability can affect the rate of key ecosystems,[3] nitrogen fixed by oceans,[4]
ecosystem processes, including primary production and nitrogen fixed by agricultural crops,[5] NOx emitted
decomposition. Human activities such as fossil fuel by biomass burning,[6] NOx emitted from soil,[7]
combustion, use of artificial nitrogen fertilizers, and nitrogen fixed by lightning,[8] NH3 emitted by
release of nitrogen in wastewater have dramatically terrestrial ecosystems,[9] deposition of nitrogen to
altered the global nitrogen cycle.[17][18][19] Human terrestrial surfaces and oceans,[10][11] NH3 emitted
modification of the global nitrogen cycle can negatively from oceans,[12][13][11] ocean NO2 emissions from
affect the natural environment system and also human the atmosphere,[14] denitrification in
oceans,[4][15][11] and reactive nitrogen burial in
health.[20][21]
oceans.[5]

Contents
Processes
Nitrogen fixation
Assimilation
Ammonification
Nitrification
Denitrification
Dissimilatory nitrate reduction to ammonium
Anaerobic ammonia oxidation
Other processes
Marine nitrogen cycle Schematic representation of the flow of nitrogen
New vs. regenerated nitrogen through the ecosystem. The importance of bacteria
in the cycle is immediately recognized as being a
Human influences on the nitrogen cycle key element in the cycle, providing different forms
Consequence of human modification of the of nitrogen compounds able to be assimilated by
nitrogen cycle higher organisms
Impacts on natural systems
 Impacts on human health: nitrate
accumulation in drinking water
Impacts on human health: air quality
See also
References

Processes
Nitrogen is present in the environment in a wide variety of
+
chemical forms including organic nitrogen, ammonium (NH4 ),
− −
nitrite (NO2 ), nitrate (NO3 ), nitrous oxide (N2 O), nitric oxide
(NO) or inorganic nitrogen gas (N2 ). Organic nitrogen may be
in the form of a living organism, humus or in the intermediate
products of organic matter decomposition. The processes in the
nitrogen cycle is to transform nitrogen from one form to
another. Many of those processes are carried out by microbes,
either in their effort to harvest energy or to accumulate nitrogen
in a form needed for their growth. For example, the nitrogenous A simple diagram of the nitrogen cycle. The
wastes in animal urine are broken down by nitrifying bacteria in blue boxes represent stores of nitrogen, the
the soil to be used by plants. The diagram alongside shows how green writing is for processes that occur to
these processes fit together to form the nitrogen cycle. move the nitrogen from one place to another
and the red writing are all the bacteria
involved
Nitrogen fixation

The conversion of nitrogen gas (N2 ) into nitrates and nitrites

through atmospheric, industrial and biological processes is
called nitrogen fixation. Atmospheric nitrogen must be
processed, or "fixed", into a usable form to be taken up by
plants. Between 5 and 10 billion kg per year are fixed by
lightning strikes, but most fixation is done by free-living or
symbiotic bacteria known as diazotrophs. These bacteria have
the nitrogenase enzyme that combines gaseous nitrogen with
hydrogen to produce ammonia, which is converted by the
bacteria into other organic compounds. Most biological
nitrogen fixation occurs by the activity of Mo-nitrogenase,
found in a wide variety of bacteria and some Archaea. Mo-
nitrogenase is a complex two-component enzyme that has
multiple metal-containing prosthetic groups.[22] An example of
Classical representation of the nitrogen
free-living bacteria is Azotobacter. Symbiotic nitrogen-fixing
cycle
bacteria such as Rhizobium usually live in the root nodules of
legumes (such as peas, alfalfa, and locust trees). Here they form
a mutualistic relationship with the plant, producing ammonia in
exchange for carbohydrates. Because of this relationship, legumes will often increase the nitrogen content of
nitrogen-poor soils. A few non-legumes can also form such symbioses. Today, about 30% of the total fixed
nitrogen is produced industrially using the Haber-Bosch process,[23] which uses high temperatures and
pressures to convert nitrogen gas and a hydrogen source (natural gas or petroleum) into ammonia.[24]
Assimilation

Plants can absorb nitrate or ammonium from the soil by their root hairs. If nitrate is absorbed, it is first reduced
to nitrite ions and then ammonium ions for incorporation into amino acids, nucleic acids, and chlorophyll. In
plants that have a symbiotic relationship with rhizobia, some nitrogen is assimilated in the form of ammonium
ions directly from the nodules. It is now known that there is a more complex cycling of amino acids between
Rhizobia bacteroids and plants. The plant provides amino acids to the bacteroids so ammonia assimilation is
not required and the bacteroids pass amino acids (with the newly fixed nitrogen) back to the plant, thus
forming an interdependent relationship.[25] While many animals, fungi, and other heterotrophic organisms
obtain nitrogen by ingestion of amino acids, nucleotides, and other small organic molecules, other heterotrophs
(including many bacteria) are able to utilize inorganic compounds, such as ammonium as sole N sources.
Utilization of various N sources is carefully regulated in all organisms.

Ammonification

When a plant or animal dies or an animal expels waste, the initial form of nitrogen is organic. Bacteria or fungi
convert the organic nitrogen within the remains back into ammonium ( ), a process called
ammonification or mineralization. Enzymes involved are:

GS: Gln Synthetase (Cytosolic & Plastic)

GOGAT: Glu 2-oxoglutarate aminotransferase (Ferredoxin & NADH-dependent)
GDH: Glu Dehydrogenase:
Minor Role in ammonium assimilation.
Important in amino acid catabolism.

Nitrification

The conversion of ammonium to nitrate is performed

primarily by soil-living bacteria and other nitrifying
bacteria. In the primary stage of nitrification, the
+
oxidation of ammonium (NH4 ) is performed by
bacteria such as the Nitrosomonas species, which
−
converts ammonia to nitrites (NO2 ). Other bacterial
species such as Nitrobacter, are responsible for the
− −
oxidation of the nitrites (NO2 ) into nitrates (NO3 ). It
is important for the ammonia (NH3) to be converted
to nitrates or nitrites because ammonia gas is toxic to
plants.

Due to their very high solubility and because soils

are highly unable to retain anions, nitrates can enter
groundwater. Elevated nitrate in groundwater is a
concern for drinking water use because nitrate can A schematic representation of the microbial nitrogen
interfere with blood-oxygen levels in infants and cycle.[26][27] ANAMMOX is anaerobic ammonium
cause methemoglobinemia or blue-baby oxidation, DNRA is dissimilatory nitrate reduction to
[28] ammonium, and COMMAMOX is complete ammonium
syndrome. Where groundwater recharges stream
oxidation.
flow, nitrate-enriched groundwater can contribute to
eutrophication, a process that leads to high algal
population and growth, especially blue-green algal populations. While not directly toxic to fish life, like
ammonia, nitrate can have indirect effects on fish if it contributes to this eutrophication. Nitrogen has
contributed to severe eutrophication problems in some water bodies. Since 2006, the application of nitrogen
fertilizer has been increasingly controlled in Britain and the United States. This is occurring along the same
lines as control of phosphorus fertilizer, restriction of which is normally considered essential to the recovery of
eutrophied waterbodies.

Denitrification

Denitrification is the reduction of nitrates back into nitrogen gas (N2 ), completing the nitrogen cycle. This
process is performed by bacterial species such as Pseudomonas and Paracoccus, under anaerobic conditions.
They use the nitrate as an electron acceptor in the place of oxygen during respiration. These facultatively
(meaning optionally) anaerobic bacteria can also live in aerobic conditions. Denitrification happens in
anaerobic conditions e.g. waterlogged soils. The denitrifying bacteria use nitrates in the soil to carry out
respiration and consequently produce nitrogen gas, which is inert and unavailable to plants.

Dissimilatory nitrate reduction to ammonium

Dissimilatory nitrate reduction to ammonium (DNRA), or nitrate/nitrite ammonification, is an anaerobic

respiration process. Microbes which undertake DNRA oxidise organic matter and use nitrate as an electron
acceptor, reducing it to nitrite, then ammonium (NO3 −→NO2 −→NH4 +).[29] Both denitrifying and nitrate
ammonification bacteria will be competing for nitrate in the environment, although DNRA acts to conserve
bioavailable nitrogen as soluble ammonium rather than producing dinitrogen gas.[30]

Anaerobic ammonia oxidation

In this biological process, nitrite and ammonia are converted directly into molecular nitrogen (N2 ) gas. This
process makes up a major proportion of nitrogen conversion in the oceans. The balanced formula for this
+ −
"anammox" chemical reaction is: NH4 + NO2 → N2 + 2H2 O (ΔG° = −357 kJ⋅mol−1 ).[31]

Other processes

Though nitrogen fixation is the primary source of plant-available nitrogen in most ecosystems, in areas with
nitrogen-rich bedrock, the breakdown of this rock also serves as a nitrogen source.[32][33][34] Nitrate reduction
is also part of the iron cycle, under anoxic conditions Fe(II) can donate an electron to NO3 − and is oxidized to
Fe(III) while NO3 − is reduced to NO2 −, N2 O, N2 , and NH4 + depending on the conditions and microbial
species involved.[35]

Marine nitrogen cycle

The nitrogen cycle is an important process in the ocean as well. While the overall cycle is similar, there are
different players[36] and modes of transfer for nitrogen in the ocean. Nitrogen enters the water through the
precipitation, runoff, or as N2 from the atmosphere. Nitrogen cannot be utilized by phytoplankton as N2 so it
must undergo nitrogen fixation which is performed predominately by cyanobacteria.[37] Without supplies of
fixed nitrogen entering the marine cycle, the fixed nitrogen would be used up in about 2000 years.[38]
Phytoplankton need nitrogen in biologically available forms for the initial synthesis of organic matter.
Ammonia and urea are released into the water by excretion from plankton. Nitrogen sources are removed from
the euphotic zone by the downward movement of
the organic matter. This can occur from sinking of
phytoplankton, vertical mixing, or sinking of waste
of vertical migrators. The sinking results in ammonia
being introduced at lower depths below the euphotic
zone. Bacteria are able to convert ammonia to nitrite
and nitrate but they are inhibited by light so this
must occur below the euphotic zone.[39]
Ammonification or Mineralization is performed by
bacteria to convert organic nitrogen to ammonia.
A schematic representing the Marine Nitrogen Cycle
Nitrification can then occur to convert the
ammonium to nitrite and nitrate.[40] Nitrate can be
returned to the euphotic zone by vertical mixing and
upwelling where it can be taken up by phytoplankton to continue the cycle. N2 can be returned to the
atmosphere through denitrification.

Ammonium is thought to be the preferred source of fixed nitrogen for phytoplankton because its assimilation
does not involve a redox reaction and therefore requires little energy. Nitrate requires a redox reaction for
assimilation but is more abundant so most phytoplankton have adapted to have the enzymes necessary to
undertake this reduction (nitrate reductase). There are a few notable and well-known exceptions that include
most Prochlorococcus and some Synechococcus that can only take up nitrogen as ammonium.[38]

The nutrients in the ocean are not uniformly distributed. Areas of upwelling provide supplies of nitrogen from
below the euphotic zone. Coastal zones provide nitrogen from runoff and upwelling occurs readily along the
coast. However, the rate at which nitrogen can be taken up by phytoplankton is decreased in oligotrophic
waters year-round and temperate water in the summer resulting in lower primary production.[41] The
distribution of the different forms of nitrogen varies throughout the oceans as well.

Nitrate is depleted in near-surface water except in upwelling regions. Coastal upwelling regions usually have
high nitrate and chlorophyll levels as a result of the increased production. However, there are regions of high
surface nitrate but low chlorophyll that are referred to as HNLC (high nitrogen, low chlorophyll) regions. The
best explanation for HNLC regions relates to iron scarcity in the ocean, which may play an important part in
ocean dynamics and nutrient cycles. The input of iron varies by region and is delivered to the ocean by dust
(from dust storms) and leached out of rocks. Iron is under consideration as the true limiting element to
ecosystem productivity in the ocean.

Ammonium and nitrite show a maximum concentration at 50–80 m (lower end of the euphotic zone) with
decreasing concentration below that depth. This distribution can be accounted for by the fact that nitrite and
ammonium are intermediate species. They are both rapidly produced and consumed through the water
column.[38] The amount of ammonium in the ocean is about 3 orders of magnitude less than nitrate.[38]
Between ammonium, nitrite, and nitrate, nitrite has the fastest turnover rate. It can be produced during nitrate
assimilation, nitrification, and denitrification; however, it is immediately consumed again.

New vs. regenerated nitrogen

Nitrogen entering the euphotic zone is referred to as new nitrogen because it is newly arrived from outside the
productive layer.[37] The new nitrogen can come from below the euphotic zone or from outside sources.
Outside sources are upwelling from deep water and nitrogen fixation. If the organic matter is eaten, respired,
delivered to the water as ammonia, and re-incorporated into organic matter by phytoplankton it is considered
recycled/regenerated production.
New production is an important component of the marine environment. One reason is that only continual input
of new nitrogen can determine the total capacity of the ocean to produce a sustainable fish harvest.[41]
Harvesting fish from regenerated nitrogen areas will lead to a decrease in nitrogen and therefore a decrease in
primary production. This will have a negative effect on the system. However, if fish are harvested from areas
of new nitrogen the nitrogen will be replenished.

Human influences on the nitrogen cycle

As a result of extensive cultivation of legumes (particularly soy,
alfalfa, and clover), growing use of the Haber–Bosch process in the
creation of chemical fertilizers, and pollution emitted by vehicles and
industrial plants, human beings have more than doubled the annual
transfer of nitrogen into biologically available forms.[28] In addition,
humans have significantly contributed to the transfer of nitrogen trace
gases from Earth to the atmosphere and from the land to aquatic
systems. Human alterations to the global nitrogen cycle are most
intense in developed countries and in Asia, where vehicle emissions
and industrial agriculture are highest.[42] Nitrogen fertilizer application

Generation of Nr, reactive nitrogen, has increased over 10 fold in the

past century due to global industrialisation.[2][43] This form of
nitrogen follows a cascade through the biosphere via a variety of
mechanisms, and is accumulating as the rate of its generation is
greater than the rate of denitrification.[44]

Nitrous oxide (N2 O) has risen in the atmosphere as a result of

agricultural fertilization, biomass burning, cattle and feedlots, and
industrial sources.[45] N2 O has deleterious effects in the stratosphere,
where it breaks down and acts as a catalyst in the destruction of
atmospheric ozone. Nitrous oxide is also a greenhouse gas and is Nitrogen in manure production
currently the third largest contributor to global warming, after carbon
dioxide and methane. While not as abundant in the atmosphere as
carbon dioxide, it is, for an equivalent mass, nearly 300 times more potent in its ability to warm the planet.[46]

Ammonia (NH3 ) in the atmosphere has tripled as the result of human activities. It is a reactant in the
atmosphere, where it acts as an aerosol, decreasing air quality and clinging to water droplets, eventually
resulting in nitric acid (HNO3 ) that produces acid rain. Atmospheric ammonia and nitric acid also damage
respiratory systems.

The very high temperature of lightning naturally produces small amounts of NOx , NH3 , and HNO3 , but high-
temperature combustion has contributed to a 6- or 7-fold increase in the flux of NOx to the atmosphere. Its
production is a function of combustion temperature - the higher the temperature, the more NOx is produced.
Fossil fuel combustion is a primary contributor, but so are biofuels and even the burning of hydrogen.
However, the rate that hydrogen is directly injected into the combustion chambers of internal combustion
engines can be controlled to prevent the higher combustion temperatures that produce NOx .

Ammonia and nitrous oxides actively alter atmospheric chemistry. They are precursors of tropospheric (lower
atmosphere) ozone production, which contributes to smog and acid rain, damages plants and increases
nitrogen inputs to ecosystems. Ecosystem processes can increase with nitrogen fertilization, but anthropogenic
input can also result in nitrogen saturation, which weakens productivity and can damage the health of plants,
animals, fish, and humans.[28]
Decreases in biodiversity can also result if higher nitrogen availability increases nitrogen-demanding grasses,
causing a degradation of nitrogen-poor, species-diverse heathlands.[47]

Consequence of human modification of the nitrogen cycle

Impacts on natural systems

Increasing levels of nitrogen deposition are shown to have a number of negative effects on both terrestrial and
aquatic ecosystems.[48][49] Nitrogen gases and aerosols can be directly toxic to certain plant species, affecting
the aboveground physiology and growth of plants near large point sources of nitrogen pollution. Changes to
plant species may also occur, as accumulation of nitrogen compounds increase its availability in a given
ecosystem, eventually changing the species composition, plant diversity, and nitrogen cycling. Ammonia and
ammonium - two reduced forms of nitrogen - can be detrimental over time due to an increased toxicity toward
sensitive species of plants,[50] particularly those that are accustomed to using nitrate as their source of nitrogen,
causing poor development of their roots and shoots. Increased nitrogen deposition also leads to soil
acidification, which increases base cation leaching in the soil and amounts of aluminum and other potentially
toxic metals, along with decreasing the amount of nitrification occurring and increasing plant-derived litter.
Due to the ongoing changes caused by high nitrogen deposition, an environment's susceptibility to ecological
stress and disturbance - such as pests and pathogens - may increase, thus making it less resilient to situations
that otherwise would have little impact to its long-term vitality.

Additional risks posed by increased availability of inorganic nitrogen in aquatic ecosystems include water
acidification; eutrophication of fresh and saltwater systems; and toxicity issues for animals, including
humans.[51] Eutrophication often leads to lower dissolved oxygen levels in the water column, including
hypoxic and anoxic conditions, which can cause death of aquatic fauna. Relatively sessile benthos, or bottom-
dwelling creatures, are particularly vulnerable because of their lack of mobility, though large fish kills are not
uncommon. Oceanic dead zones near the mouth of the Mississippi in the Gulf of Mexico are a well-known
example of algal bloom-induced hypoxia.[52][53] The New York Adirondack Lakes, Catskills, Hudson
Highlands, Rensselaer Plateau and parts of Long Island display the impact of nitric acid rain deposition,
resulting in the killing of fish and many other aquatic species.[54]

Ammonia (NH3 ) is highly toxic to fish and the level of ammonia discharged from wastewater treatment
facilities must be closely monitored. To prevent fish deaths, nitrification via aeration prior to discharge is often
desirable. Land application can be an attractive alternative to the aeration.

Impacts on human health: nitrate accumulation in drinking water

Leakage of Nr (reactive nitrogen) from human activities can cause nitrate accumulation in the natural water
environment, which can create harmful impacts on human health. Excessive use of N-fertilizer in agriculture
has been one of the major sources of nitrate pollution in groundwater and surface water.[55][56] Due to its high
solubility and low retention by soil, nitrate can easily escape from the subsoil layer to the groundwater, causing
nitrate pollution. Some other non-point sources for nitrate pollution in groundwater are originated from
livestock feeding, animal and human contamination and municipal and industrial waste. Since groundwater
often serves as the primary domestic water supply, nitrate pollution can be extended from groundwater to
surface and drinking water in the process of potable water production, especially for small community water
supplies, where poorly regulated and unsanitary waters are used.[57]

The WHO standard for drinking water is 50 mg NO3 − L−1 for short-term exposure, and for 3 mg NO3 −
L−1 chronic effects.[58] Once it enters human body, nitrate can react with organic compounds through
nitrosation reactions in the stomach to form nitrosamines and nitrosamides, which are involved in some types
of cancers (e.g., oral cancer and gastric cancer).[59]

Impacts on human health: air quality

Human activities have also dramatically altered the global nitrogen cycle via production of nitrogenous gases,
associated with the global atmospheric nitrogen pollution. There are multiple sources of atmospheric reactive
nitrogen (Nr) fluxes. Agricultural sources of reactive nitrogen can produce atmospheric emission of ammonia
(NH3), nitrogen oxides (NOx ) and nitrous oxide (N2 O). Combustion processes in energy production,
transportation and industry can also result in the formation of new reactive nitrogen via the emission of NOx ,
an unintentional waste product. When those reactive nitrogens are released to the lower atmosphere, they can
induce the formation of smog, particulate matter and aerosols, all of which are major contributors to adverse
health effects on human health from air pollution.[60] In the atmosphere, NO2 can be oxidized to nitric acid
(HNO3 ), and it can further react with NH3 to form ammonium nitrate, which facilitates the formation of
particular nitrate. Moreover, NH3 can react with other acid gases (sulfuric and hydrochloric acids) to form
ammonium-containing particles, which are the precursors for the secondary organic aerosol particles in
photochemical smog.[61]

See also
Planetary boundaries
Phosphorus cycle

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