the continuous flow of nitrogen through the biosphere by the processes of nitrogen fixation, ammonification (decay), nitrification, and denitrification. Nitrogen is vital to all living matter, both plant and animal; it is an essential constituent of amino acids, which form proteins of nucleic acids, and of many other organic materials.
Although the earth's atmosphere is 78% nitrogen, free gaseous nitrogen cannot be utilized by animals or by higher plants. They depend instead on nitrogen that is present in the soil. To enter living systems, nitrogen must be "fixed" (combined with oxygen or hydrogen) into compounds that plants can utilize, such as nitrates or ammonia. A certain amount of atmospheric nitrogen is fixed by lightning and by some cyanobacteria (blue-green algae). But the great bulk of nitrogen fixation is performed by soil bacteria of two kinds: those that live free in the soil and those that live enclosed in nodules in the roots of certain leguminous plants (e.g., alfalfa, peas, beans, clover, soybeans, and peanuts). Among the free-living forms are species of Clostridium, discovered c.1893 by Sergei Winogradsky, and Azotobacter, discovered c.1901 by M. W. Beijerinck. Both Clostridium and Azotobacter are generally present in agricultural soils, and both are saprophytes, i.e., they use the energy from decaying organic matter in the soil to fuel soil processes, including nitrogen fixation.
Bacteria that live in the roots of legumes are of the genus Rhizobium, first isolated c.1888 by Beijerinck. These rod-shaped bacteria enter the roots chiefly through the root hairs and then work their way to the inner root tissues. There they stimulate the growth of tumorlike nodules. Within the nodules the bacteria develop into forms called bacteroids, which live in a symbiotic (mutually beneficial) relationship with the green plant. The bacteroids take carbohydrates from the plant for energy to fix nitrogen and synthesize amino acids; the plants take the amino acids elaborated in the nodule to build plant tissue. Animals in turn consume the plants and convert plant protein into animal protein. Rhizobia can be found free-living in the soil, but they cannot fix nitrogen in the free state, nor can the legume root fix nitrogen without Rhizobia.
The exact biochemistry of nitrogen fixation within the nodule is not yet understood. It is estimated that more than 300 lbs of nitrogen per acre (340 kg per hectare) can be fixed by fields of alfalfa and other legumes. After a harvest legume roots left in the soil decay, returning organic nitrogen compounds to the soil for uptake by the next generation of plants. For this reason crop rotation in which a leguminous crop is rotated with a nonleguminous one is a common practice for maintaining soil fertility.
Decomposing animal remains and animal wastes also return organic nitrogen to the soil as ammonia. Many different kinds of decay microorganisms participate in ammonification. The nitrifying bacteria of the genus Nitrosomonas oxidize the ammonia to nitrites, and Nitrobacter oxidize the nitrites to nitrates. The nitrates can then be taken up again by the green plant. The cycle of fixation-decay-nitrification-fixation can proceed indefinitely without any nitrogen being returned to a gaseous state. But still another group of microorganisms, the denitrifying bacteria, can reduce nitrates all the way to molecular nitrogen. Denitrification occurs only in the absence of oxygen and is not common in well-cultivated soils.
Nitrogen fixation can also be accomplished artificially by various methods (see nitrogen). Humans annually fix vast amounts of nitrogen for industrial purposes and for use as fertilizer. Unfortunately, large-scale legume cultivation and artificial fixation may be upsetting the natural nitrogen cycle in the biosphere. There is some question whether natural denitrification can keep pace with fixation. For one thing, run-off of nitrate fertilizer can cause eutrophication of lakes and streams (see water pollution) and can foul drinking supplies. Another environmental problem is that inorganic fertilizers tend to depress legume fixation. As a consequence, root tissue remaining after harvest is poorer, and thus more fertilizer must be applied the following year.
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