The nitrogen cycle is the process by which nitrogen is converted between its various
chemical forms. This transformation 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 nitrogen, making it the largest pool of nitrogen. However, atmospheric nitrogen
has limited availability for biological use, leading to a scarcity of usable nitrogen in many
types of ecosystems.
The processes of the nitrogen cycle
Nitrogen is present in the environment in a wide variety of chemical forms including
organic nitrogen, ammonium (NH4+), nitrite (NO2-), nitrate (NO3-), nitrous oxide (N2O),
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 of the nitrogen cycle 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. The diagram
above shows how these processes fit together to form the nitrogen cycle.
Nitrogen fixation
Atmospheric nitrogen must be processed, or "fixed", to be used by plants. Some fixation
occurs in lightning strikes, but most fixations are done by free-living or symbiotic
bacteria. These bacteria have the nitrogenase enzyme that combines gaseous nitrogen
with hydrogen to produce ammonia, which is then further converted by the bacteria to
make their own 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. Some nitrogen fixing bacteria, such as Rhizobium, live in the root
nodules of legumes (such as peas or beans). Here they form a mutualistic relationship
with the plant, producing ammonia in exchange for carbohydrates. Nutrient-poor soils
can be planted with legumes to enrich them with nitrogen. A few other plants can form
such symbioses. Today, about 30% of the total fixed nitrogen is manufactured in
ammonia chemical plants.
Conversion of N2
The conversion of nitrogen (N2) from the atmosphere into a form readily available to
plants and hence to animals is an important step in the nitrogen cycle, which distributes
the supply of this essential nutrient. There are four ways to convert N2 (atmospheric
nitrogen gas) into more chemically reactive forms:
1. Biological fixation: some symbiotic bacteria (most often associated with
leguminous plants) and some free-living bacteria are able to fix nitrogen as
organic nitrogen. An example of mutualistic nitrogen fixing bacteria are the
Rhizobium bacteria, which live in legume root nodules. These species are
diazotrophs. An example of the free-living bacteria is Azotobacter.
2. Industrial N-fixation: Under great pressure, at a temperature of 600 C, and with
the use of an iron catalyst, hydrogen (usually derived from natural gas or
petroleum) and atmospheric nitrogen can be combined to form ammonia (NH3) in
the Haber-Bosch process which is used to make fertilizer and explosives.
3. Combustion of fossil fuels: automobile engines and thermal power plants, which
release various nitrogen oxides (NOx).
4. Other processes: In addition, the formation of NO from N2 and O2 due to photons
and especially lightning, can fix nitrogen.
Assimilation
Plants take nitrogen from the soil by absorption through their roots in the form of either
nitrate ions or ammonium ions. All nitrogen obtained by animals can be traced back to
the eating of plants at some stage of the food chain.
Plants can absorb nitrate or ammonium ions from the soil via 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.
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 in some cases, convert the organic nitrogen within the remains
back into ammonium (NH4+), a process called ammonification or mineralization.
Enzymes Involved are:
GS: Gln Synthetase (Cytosolic & PLastid)
GOGAT: Glu 2-oxoglutarate aminotransferase (Ferredoxin & NADH dependent)
GDH: Glu Dehydrogenase:
o Minor Role in ammonium assimilation.
o Important in amino acid catabolism.
Nitrification
The conversion of ammonia 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 the Nitrobacter, are
responsible for the oxidation of the nitrites into nitrates (NO3-). It is important for the
ammonia to be converted to nitrates because accumulated nitrites are toxic to plant life.
Due to their very high solubility, and because soils are largely unable to retain anions,
nitrates can enter groundwater. Elevated nitrate in groundwater is a concern for drinking
water use because nitrate can interfere with blood-oxygen levels in infants and cause
methemoglobinemia or blue-baby syndrome. Where groundwater recharges stream flow,
nitrate-enriched groundwater can contribute to eutrophication, a process that leads to high
algal, 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 the largely inert nitrogen gas (N 2),
completing the nitrogen cycle. This process is performed by bacterial species such as
Pseudomonas and Clostridium in anaerobic conditions. They use the nitrate as an
electron acceptor in the place of oxygen during respiration. These facultatively anaerobic
bacteria can also live in aerobic conditions.
Anaerobic ammonium oxidation
In this biological process, nitrite and ammonium are converted directly into molecular
nitrogen (N2) gas. This process makes up a major proportion of nitrogen conversion in the
oceans.