Diazotrophs are bacteria and archaea that fix atmospheric nitrogen gas into a more usable form such as ammonia. A diazotroph is a microorganism that is able to grow without external sources of fixed nitrogen. Examples of organisms that do this are rhizobia and Frankia and Azospirillum. All diazotrophs contain iron-molybdenum or -vanadium nitrogenase systems. Two of the most studied systems are those of Klebsiella pneumoniae and Azotobacter vinelandii. These systems are used because of their genetic tractability and their fast growth.
Etymology
The word diazotroph is derived from the words diazo meaning "dinitrogen " and troph meaning "pertaining to food or nourishment", in summary dinitrogen utilizing. The word azote means nitrogen in French and was named by French chemist and biologist Antoine Lavoisier, who saw it as the part of air which cannot sustain life.
Types of diazotrophs
Diazotrophs are scattered across Bacteriataxonomic groups. Even within a species that can fix nitrogen there may be strains that do not fix nitrogen. Fixation is shut off when other sources of nitrogen are available, and, for many species, when oxygen is at high partial pressure. Bacteria have different ways of dealing with the debilitating effects of oxygen on nitrogenases, listed below.
Free-living diazotrophs
Anaerobes—these are obligate anaerobes that cannot tolerate oxygen even if they are not fixing nitrogen. They live in habitats low in oxygen, such as soils and decaying vegetable matter. Clostridium is an example. Sulphate-reducing bacteria are important in ocean sediments, and some Archean methanogens, like Methanococcus, fix nitrogen in muds, animal intestines and anoxic soils.
Facultative anaerobes—these species can grow either with or without oxygen, but they only fix nitrogen anaerobically. Often, they respire oxygen as rapidly as it is supplied, keeping the amount of free oxygen low. Examples include Klebsiella pneumoniae, Paenibacillus polymyxa, Bacillus macerans, and Escherichia intermedia.
Aerobes—these species require oxygen to grow, yet their nitrogenase is still debilitated if exposed to oxygen. Azotobacter vinelandii is the most studied of these organisms. It uses very high respiration rates, and protective compounds, to prevent oxygen damage. Many other species also reduce the oxygen levels in this way, but with lower respiration rates and lower oxygen tolerance.
Oxygenic photosynthetic bacteria generate oxygen as a by-product of photosynthesis, yet some are able to fix nitrogen as well. These are colonial bacteria that have specialized cells that lack the oxygen generating steps of photosynthesis. Examples are Anabaena cylindrica and Nostoc commune. Other cyanobacteria lack heterocysts and can fix nitrogen only in low light and oxygen levels. Some cyanobacteria, including the highly abundant marine taxa Prochlorococcus and Synechococcus do not fix nitrogen, whilst other marine cyanobacteria, such as Trichodesmium and Cyanothece, are major contributors to oceanic nitrogen fixation.
Anoxygenic photosynthetic bacteria do not generate oxygen during photosynthesis, having only a single photosystem which cannot split water. Nitrogenase is expressed under nitrogen limitation. Normally, the expression is regulated via negative feedback from the produced ammonium ion but in the absence of N2, the product is not formed, and the by-product H2 continues unabated . Example species: Rhodobacter sphaeroides, Rhodopseudomonas palustris, Rhodobacter capsulatus.
Symbiotic diazotrophs
Rhizobia—these are the species that associate with legumes, plants of the family Fabaceae. Oxygen is bound to leghemoglobin in the root nodules that house the bacterial symbionts, and supplied at a rate that will not harm the nitrogenase.
Frankias—much less is known about these 'actinorhizal' nitrogen fixers. The bacteria also infect the roots leading to the formation of nodules. Actinorhizal nodules consist of several lobes, each lobe has a similar structure as a lateral root. Frankia is able to colonize in the cortical tissue of nodules where it fixes nitrogen. Actinorhizal plants and Frankias also produce haemoglobins, but their role is less well established than for rhizobia. Although at first it appeared that they inhabit sets of unrelated plants, revisions to the phylogeny of angiosperms show a close relatedness of these species and the legumes. These footnotes suggest the ontogeny of these replicates rather than the phylogeny. In other words, an ancient gene that is unused in most species was reawakened and reused in these species.
Cyanobacteria—there are also symbiotic cyanobacteria. Some associate with fungi as lichens, with liverworts, with a fern, and with a cycad. These do not form nodules. Heterocysts exclude the oxygen, as discussed above. The fern association is important agriculturally: the water fernAzolla harbouring Anabaena is an important green manure for rice culture.
Association with animals—although diazotrophs have been found in many animal guts, there is usually sufficient ammonia present to suppress nitrogen fixation. Termites on a low nitrogen diet allow for some fixation, but the contribution to the termite's nitrogen supply is negligible. Shipworms may be the only species that derive significant benefit from their gut symbionts.
Importance
In terms of generating nitrogen available to all organisms, the symbiotic associations greatly exceed the free-living species with the exception of cyanobacteria.
