Sep 29, 2009

Nitrogen Cycle

• All life requires nitrogen-compounds, e.g., proteins and nucleic acids.
• Air, which is 78% nitrogen gas (N2), is the major reservoir of nitrogen.
• But most organisms cannot use nitrogen in this form.
• Plants must secure their nitrogen in "fixed" form, i.e., incorporated in compounds such as:
  • nitrate ions (NO3−)
  • ammonia (NH3)
  • Animals secure their nitrogen (and all other) compounds from plants (or animals that have fed on plants).
Four processes participate in the cycling of nitrogen through the biosphere:

  • nitrogen fixation
  • decay
  • nitrification
  • denitrification
Microorganisms play major roles in all four of these.

Nitrogen Fixation
The nitrogen molecule (N2) is quite inert. To break it apart so that its atoms can combine with other atoms requires the input of substantial amounts of energy.

Three processes are responsible for most of the nitrogen fixation in the biosphere:
  • atmospheric fixation by lightning
  • biological fixation by certain microbes — alone or in a symbiotic relationship with some plants and animals
  • industrial fixation
Atmospheric Fixation
The enormous energy of lightning breaks nitrogen molecules and enables their atoms to combine with oxygen in the air forming nitrogen oxides. These dissolve in rain, forming nitrates that are carried to the earth. Atmospheric nitrogen fixation probably contributes some 5– 8% of the total nitrogen fixed.

Industrial Fixation
Under great pressure, at a temperature of 600°C, and with the use of a catalyst, atmospheric nitrogen and hydrogen (usually derived from natural gas or petroleum) can be combined to form ammonia (NH3). Ammonia can be used directly as fertiliser, but most of it is further processed to urea and ammonium nitrate (NH4NO3).

Biological Fixation
The ability to fix nitrogen is found only in certain bacteria and archaea.
  • Some live in a symbiotic relationship with plants of the legume family (e.g., soybeans, alfalfa).
  • Some establish symbiotic relationships with plants other than legumes (e.g., alders).
  • Some establish symbiotic relationships with animals, e.g., termites and "shipworms" (wood-eating bivalves).
  • Some nitrogen-fixing bacteria live free in the soil.
  • Nitrogen-fixing cyanobacteria are essential to maintaining the fertility of semi-aquatic environments like rice paddies.
Biological nitrogen fixation requires a complex set of enzymes and a huge expenditure of ATP.

Although the first stable product of the process is ammonia, this is quickly incorporated into protein and other organic nitrogen compounds.

The proteins made by plants enter and pass through food webs just as carbohydrates do. At each trophic level, their metabolism produces organic nitrogen compounds that return to the environment, chiefly in excretions. The final beneficiaries of these materials are microorganisms of decay. They break down the molecules in excretions and dead organisms into ammonia.

Ammonia can be taken up directly by plants — usually through their roots. However, most of the ammonia produced by decay is converted into nitrates. This is accomplished in two steps:

  • Bacteria of the genus Nitrosomonas oxidise NH3 to nitrites (NO2−).
  • Bacteria of the genus Nitrobacter oxidise the nitrites to nitrates (NO3−).
These two groups of autotrophic bacteria are called nitrifying bacteria. Through their activities (which supply them with all their energy needs), nitrogen is made available to the roots of plants.

Both soil and the ocean contain archaeal microbes, assigned to the Crenarchaeota, that convert ammonia to nitrites. They are more abundant than the nitrifying bacteria and may turn out to play an important role in the nitrogen cycle.

Many legumes, in addition to fixing atmospheric nitrogen, also perform nitrification — converting some of their organic nitrogen to nitrites and nitrates. These reach the soil when they shed their leaves.

The three processes above remove nitrogen from the atmosphere and pass it through ecosystems.
Denitrification reduces nitrates to nitrogen gas, thus replenishing the atmosphere.
Once again, bacteria are the agents. They live deep in soil and in aquatic sediments where conditions are anaerobic. They use nitrates as an alternative to oxygen for the final electron acceptor in their respiration.

Thus they close the nitrogen cycle.

Are the denitrifiers keeping up?
Agriculture may now be responsible for one-half of the nitrogen fixation on earth through
  • the use of fertilisers produced by industrial fixation
  • the growing of legumes like soybeans and alfalfa.
This is a remarkable influence on a natural cycle.

Are the denitrifiers keeping up the nitrogen cycle in balance? Probably not. Certainly, there are examples of nitrogen enrichment in ecosystems. One troubling example: the "blooms" of algae in lakes and rivers as nitrogen fertilisers leach from the soil of adjacent farms (and lawns). The accumulation of dissolved nutrients in a body of water is called eutrophication.