Nitrogen (N) is an essential component of building blocks of life.

All

DNA, RNA, and proteins, the

organisms require nitrogen to live and grow.

Although the majority of the air we breathe is N 2, most of the nitrogen in the atmosphere is unavailable for use by organisms. This is because the strong triple bond between the N

atoms in N2 molecules makes it relatively inert.

In fact, in order for plants and animals to be able to use nitrogen, N2 gas must first be converted to more a chemically available form such as ammonium (NH4+), nitrate (NO3-), or organic nitrogen (e.g. urea - (NH2)2CO). The inert nature of N2 in natural

means that biologically available nitrogen is often in short supply

ecosystems, limiting plant growth and biomass accumulation. element, existing in both inorganic and oxidation states. The movement of biosphere, and geosphere in different

Nitrogen is an incredibly versatile

organic forms as well as many different nitrogen between the atmosphere,

forms is described by the nitrogen cycle (Figure 1), one of the major

biogeochemical cycles. Similar to the carbon cycle, the nitrogen cycle

consists of various storage pools of nitrogen and processes by which the pools exchange nitrogen (arrows) (see our information).

The Carbon Cycle module for more

Figure 1: The nitrogen cycle. Yellow arrows indicate human sources of nitrogen to the environment. Red arrows indicate microbial transformations of nitrogen. Blue arrows indicate physical forces acting on nitrogen. And green arrows indicate natural, non-microbial processes affecting the form and fate of nitrogen.

Five main processes cycle nitrogen through the geosphere:

biosphere, atmosphere, and

nitrogen fixation, nitrogen uptake (organismal growth), nitrogen

mineralization (decay), nitrification, and denitrification. Microorganisms,

particularly bacteria, play major roles in all of the principal nitrogen transformations. As microbially mediated processes, these nitrogen

transformations tend to occur faster than geological processes like plate motion, a very slow, purely physical process that is a part of the carbon cycle. Instead, rates are affected by environmental factors that influence microbial activity, such as temperature, moisture, and resource availability.

The Microbial World

Figure 2: Part of a clover root system bearing naturally occurring nodules of Rhizobium, bacteria that can fix atmospheric nitrogen. Each nodule is about 2-3 mm long. Image courtesy of http://helios.bto.ed.ac.uk/bto/microbes/nitrogen.htm.

Nitrogen fixation N2 NH4+

Nitrogen fixation is the process wherein N2 is converted to organisms can attain

ammonium, essential because it is the only way that

nitrogen directly from the atmosphere. Certain bacteria, for example those among the

genus Rhizobium, are the only organisms that fix nitrogen through symbiosis is well-known to occur in the legume family of

metabolic processes. Nitrogen fixing bacteria often form symbiotic relationships with host plants. This

plants (e.g. beans, peas, and clover). In this relationship, nitrogen fixing bacteria inhabit legume root nodules (Figure 2) and receive carbohydrates and a favorable environment from their host plant in exchange for some of the nitrogen they fix. There are also nitrogen fixing bacteria that exist without plant hosts, known as free-living nitrogen fixers. In aquatic environments, blue-green algae (really a bacteria called cyanobacteria) is an important free-living nitrogen fixer. In addition to nitrogen fixing bacteria, high-energy natural events such as lightning, forest fires, and even hot lava flows can cause the fixation of smaller, but significant amounts of nitrogen (Figure 3). The high natural phenomena can break the triple bonds of N2 making individual N

energy of these

molecules, thereby

atoms available for chemical transformation.

Within the last century, humans have become as important a source of fixed nitrogen as all natural sources combined. Burning fossil fuels, using synthetic nitrogen fertilizers, and cultivation of legumes all fix nitrogen. Through these activities, humans have more than doubled the amount of fixed nitrogen that is pumped into the

biosphere every year (Figure 3), the consequences of which

are discussed below.

Figure 3: Recent increases in anthropogenic N fixation in relation to natural N fixation. Modified from Vitousek, P. M. and P. A. Matson (1993). Agriculture, the global nitrogen cycle, and trace gas flux. The Biogeochemistry of Global Change: Radiative Trace Gases. R. S. Oremland. New York, Chapman and Hall: 193-208.

Nitrogen uptake NH4+ Organic N The ammonia produced by nitrogen fixing bacteria is usually

quickly incorporated into

protein and other organic nitrogen compounds, organism. When

either by a host plant, the bacteria itself, or another soil that has been fixed initially by nitrogen fixing bacteria. Nitrogen mineralization Organic N

organisms nearer the top of the food chain (like us!) eat, we are using nitrogen

NH4+ After nitrogen is incorporated into organic matter, it is often

converted back into inorganic nitrogen by a process called nitrogen

mineralization, otherwise known as decay. When organisms die,

decomposers (such as bacteria and fungi) consume the organic matter and lead

to the process of decomposition. During this process, a significant amount of the nitrogen contained within the dead organism is converted to ammonium. Once in the form of ammonium, nitrogen is available for use by plants or for further transformation into nitrate (NO3-) through the process called Nitrification NH4+ NO3- Some of the ammonium produced by decomposition is converted

nitrification.

to nitrate via a process called reaction gain

nitrification. The bacteria that carry out this

energy from it. Nitrification requires the presence of oxygen, so sediments. The process ions are

nitrification can happen only in oxygen-rich environments like circulating or flowing waters and the very surface layers of soils and

of nitrification has some important consequences. Ammonium

positively charged and therefore stick (are sorbed) to negatively charged clay particles and soil organic matter. The positive charge prevents ammonium nitrogen from being washed out of the soil (or leached) by rainfall. In contrast, the negatively charged nitrate ion is not held by soil particles and so can be washed down the soil profile, leading to decreased soil fertility and nitrate enrichment of downstream surface and Denitrification NO3N2+ N2O Through (NO2-)

groundwaters.

denitrification, oxidized forms of nitrogen such as

nitrate and nitrite

are converted to dinitrogen (N2) and, to a lesser

extent, nitrous oxide gas. Denitrification is an anaerobic process that is carried out by denitrifying bacteria, which convert nitrate to dinitrogen in the following sequence: NO3NO2NO N2O N2. Nitric oxide and nitrous oxide are both environmentally important gases. Nitric oxide (NO) contributes to

smog, and nitrous oxide (N2O) is an important

greenhouse gas, thereby contributing to global climate change.

Once converted to dinitrogen, nitrogen is unlikely to be reconverted to a biologically available form because it is a gas and is rapidly lost to the atmosphere.

Denitrification is the only nitrogen transformation that removes ecosystems (essentially irreversibly), and it roughly balances

nitrogen from

the amount of nitrogen fixed by the nitrogen fixers described above. Human alteration of the N cycle and its environmental consequences

Early in the 20th century, a German scientist named

Fritz Haber figured out

how to short circuit the nitrogen cycle by fixing nitrogen chemically at high temperatures and pressures, creating fertilizers that could be added directly to soil. This technology has spread rapidly over the past century, and, along with the advent of new crop varieties, the use of synthetic nitrogen fertilizers has led to an enormous boom in agricultural productivity. This agricultural productivity has helped us to feed a rapidly growing world population, but the increase in

nitrogen fixation has had some negative consequences as well. While the
consequences are perhaps not as obvious as an increase in global temperatures or a hole in the

ozone layer, they are just as serious and potentially harmful organisms.

for humans and other

Not all of the nitrogen fertilizer applied to agricultural fields stays to nourish crops. Some is washed off of agricultural fields by rain or irrigation water, where it leaches into surface or ground water and can accumulate. In

groundwater

that is used as a drinking water source, excess nitrogen can lead to cancer in humans and respiratory distress in infants. The U.S.

Environmental

Protection Agency has established a standard for nitrogen in drinking water

of 10 mg per liter nitrate-N. Unfortunately, many

systems (particularly in

agricultural areas) already exceed this level. By comparison, nitrate levels in waters that have not been altered by human activity are rarely greater than 1 mg/L. In surface waters, added nitrogen can lead to nutrient over-enrichment, particularly in coastal waters receiving the inflow from polluted rivers. This nutrient over-enrichment, also called eutrophication, has been blamed for increased frequencies of coastal fish-kill events, increased frequencies of harmful algal blooms, and

species shifts within coastal ecosystems.

Reactive nitrogen (like NO3- and NH4+) present in surface waters and soils, can also enter the atmosphere as the smog-component nitric oxide (NO) and the

greenhouse gas nitrous oxide (N2O). Eventually, this atmospheric nitrogen

can be blown into nitrogen-sensitive terrestrial environments, causing long-term changes. For example, nitrogen oxides comprise a significant portion of the acidity in

acid rain which has been blamed for forest death and decline in parts species

of Europe and the Northeast United States. Increases in atmospheric nitrogen deposition have also been blamed for more subtle shifts in dominant and

ecosystem function in some forest and grassland ecosystems. For serpentine soils of northern Californian grasslands,

example, on nitrogen-poor

plant assemblages have historically been limited to native species that can survive without a lot of nitrogen. There is now some evidence that elevated levels of atmospheric N input from nearby industrial and agricultural

development have paved the way for invasion by non-native plants. As noted earlier, NO is also a major factor in the formation of

smog, which is known to

cause respiratory illnesses like asthma in both children and adults. Currently, much research is devoted to understanding the effects of nitrogen enrichment in the air,

groundwater, and surface water. Scientists are also

exploring alternative agricultural practices that will sustain high productivity while decreasing the negative impacts caused by fertilizer use. These studies not only help us quantify how humans have altered the natural world, but increase our understanding of the processes involved in the nitrogen cycle as a whole.