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The nitrogen-fixing organisms

All the nitrogen-fixing organisms are prokaryotes (bacteria). Some of them live independently of other organisms - the so-called free-living nitrogen-fixing bacteria. Others live in intimate symbiotic associations with plants or with other organisms (e.g. protozoa). Examples are shown in the table below.


  Examples of nitrogen-fixing bacteria (* denotes a photosynthetic bacterium)
  Free living   Symbiotic with plants
  Aerobic Anaerobic (see Winogradsky column for details)     Legumes   Other plants
Azotobacter Beijerinckia Klebsiella(some) Cyanobacteria (some)* Clostridium (some) Desulfovibrio Purple sulphur bacteria* Purple non-sulphur bacteria* Green sulphur bacteria*     Rhizobium   Frankia Azospirillum


A point of special interest is that the nitrogenase enzyme complex is highly sensitive to oxygen. It is inactivated when exposed to oxygen, because this reacts with the iron component of the proteins. Although this is not a problem for anaerobic bacteria, it could be a major problem for the aerobic species such as cyanobacteria (which generate oxygen during photosynthesis) and the free-living aerobic bacteria of soils, such as Azotobacter and Beijerinckia. These organisms have various methods to overcome the problem. For example, Azotobacter species have the highest known rate of respiratory metabolism of any organism, so they might protect the enzyme by maintaining a very low level of oxygen in their cells. Azotobacter species also produce copious amounts of extracellular polysaccharide (as do Rhizobium species in culture - see Exopolysaccharides). By maintaining water within the polysaccharide slime layer, these bacteria can limit the diffusion rate of oxygen to the cells. In the symbiotic nitrogen-fixing organisms such as Rhizobium, the root nodules can contain oxygen-scavenging molecules such as leghaemoglobin, which shows as a pink colour when the active nitrogen-fixing nodules of legume roots are cut open. Leghaemoglobin may regulate the supply of oxygen to the nodule tissues in the same way as haemoglobin regulates the supply of oxygen to mammalian tissues. Some of the cyanobacteria have yet another mechanism for protecting nitrogenase: nitrogen fixation occurs in special cells (heterocysts) which possess only photosystem I (used to generate ATP by light-mediated reactions) whereas the other cells have both photosystem I and photosystem II (which generates oxygen when light energy is used to split water to supply H2 for synthesis of organic compounds).

Date: 2015-01-29; view: 1798

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