Nitrogen is an essential component of all living organisms and the main nutrient limiting life on our planet By far, the largest inventory of freely accessible nitrogen is atmospheric dinitrogen, but most organisms rely on more bioavailable forms of nitrogen, such as ammonium and nitrate, for growth The availability of these substrates depends on diverse nitrogen-transforming reactions that are carried out by complex networks of metabolically versatile microorganisms In this Review, we summarize our current understanding of the microbial nitrogen-cycling network, including novel processes, their underlying biochemical pathways, the involved microorganisms, their environmental importance and industrial applications

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The microbial nitrogen-cycling network

Biological N2 fixation represents the major source of N input in agricultural soils including those in arid regions. The major N2-fixing systems are the symbiotic systems, which can play a significant role in improving the fertility and productivity of low-N soils. The Rhizobium-legume symbioses have received most attention and have been examined extensively. The behavior of some N2-fixing systems under severe environmental conditions such as salt stress, drought stress, acidity, alkalinity, nutrient deficiency, fertilizers, heavy metals, and pesticides is reviewed. These major stress factors suppress the growth and symbiotic characteristics of most rhizobia; however, several strains, distributed among various species of rhizobia, are tolerant to stress effects. Some strains of rhizobia form effective (N2-fixing) symbioses with their host legumes under salt, heat, and acid stresses, and can sometimes do so under the effect of heavy metals. Reclamation and improvement of the fertility of arid lands by application of organic (manure and sewage sludge) and inorganic (synthetic) fertilizers are expensive and can be a source of pollution. The Rhizobium-legume (herb or tree) symbiosis is suggested to be the ideal solution to the improvement of soil fertility and the rehabilitation of arid lands and is an important direction for future research.

Rhizobium-Legume Symbiosis and Nitrogen Fixation under Severe Conditions and in an Arid Climate

Legumes, broadly defined by their unusual flower structure, podded fruit, and the ability of 88% of the species examined to date to form nodules with rhizobia ([de Faria et al., 1989][1]), are second only to the Graminiae in their importance to humans. The 670 to 750 genera and 18,000 to 19,000

Legumes: Importance and Constraints to Greater Use

Because of their ability to transform atmospheric N2 into ammonia that can be used by the plant, researchers were originally very optimistic about the potential of associative diazotrophic bacteria to promote the growth of many cereals and grasses. However, multiple inoculation experiments during recent decades failed to show a substantial contribution of Biological Nitrogen Fixation (BNF) to plant growth in most cases. It is now clear that associative diazotrophs exert their positive effects on plant growth directly or indirectly through (a combination of) different mechanisms. Apart from fixing N2, diazotrophs can affect plant growth directly by the synthesis of phytohormones and vitamins, inhibition of plant ethylene synthesis, improved nutrient uptake, enhanced stress resistance, solubilization of inorganic phosphate and mineralization of organic phosphate. Indirectly, diazotrophs are able to decrease or prevent the deleterious effects of pathogenic microorganisms, mostly through the synthesis of anti...

Plant Growth-Promoting Effects of Diazotrophs in the Rhizosphere

This paper reviews a growing body of literature on the use of variations in the natural abundance of 15N to estimate the fractional contribution of N2-fixation to N2-fixing systems. This method is based on the small difference in 15NN abundance which frequently occurs between N derived from N2-fixation and N derived from other sources. The requirement of the method is that this difference be significant. Whether this requirement is met is site specific and must be empirically established at each site of interest. Advantages and disadvantages of this method are compared with those of more conventional methods. Sources of error, including heterogeneity of 15NN abundance of non-atmospheric N sources are considered. Tests of the method, under both greenhouse and field conditions, are described. Estimates based on this method compare favourably with other methods for field evaluation of N2-fixation, provided that the site and the sampling strategy are appropriate for application of the method. Applications of the method in several ecosystems are described.

N2-Fixation in Field Settings: Estimations Based on Natural 15N Abundance

Biological nitrogen fixation (BNF) is the process of the reduction of dinitrogen from the air to ammonia carried out by a large number of species of free-living and symbiotic microbes called diazotrophs. BNF presents an inexpensive and environmentally sound, sustainable approach to crop production and constitutes one of the most important Plant Growth Promotion (PGP) scenarios. Here I will summarize various aspects of BNF, including the dinitrogen reduction catalysed reaction carried out by “nitrogenase” and the enzymes/genes involved and their regulation, the inherent “oxygen paradox” , the identification of diazotrophs, sustainable agricultural uses of BNF, symbiotic plant-diazotroph interactions and endophytic diazotrophs, data from the field, and future prospects in BNF.

Biological Nitrogen Fixation

In the photosynthetic bacterium Rhodospirillum rubrum the regulation of nitrogen fixation occurs at both transcriptional and posttranslational levels. Recently, three PII homologs, named GlnB, GlnK, and GlnJ, have been identified in R. rubrum, and they play very important roles in the regulation of nitrogen fixation. Similar to that seen in Azospirillum brasilense and other N2-fixation bacteria, the expression of the nif operon is controlled by posttranslational regulation of NifA activity in R. rubrum. Only the uridylylated form of GlnB is required for the activation of NifA activity, and GlnK and GlnJ do not appear to be involved in that process. The posttranslational regulation of nitrogenase activity involves the DRAT/DRAG regulatory system, which responds to fixed nitrogen and to energy limitations. Both GlnB and GlnJ, but not GlnK, are involved in the regulation of DRAT/DRAG activities. All three P II homologs can support proper regulation of glutamine synthetase activity. The discovery through genome analysis of both DRAT/DRAG and P II homologs in many organisms, including those incapable of nitrogen fixation, suggests that these proteins might interact to serve a variety of biological roles.

Gary P. Roberts

Papers

Biological Nitrogen Fixation

Review article Regulation of Nitrogen Fixation by Multiple PII Homologs in the Photosynthetic Bacterium Rhodospirillum rubrum

Mutations inthedraTanddraGGenesofRhodospirillum rubrum Result inLossofRegulation ofNitrogenase by Reversible ADP-Ribosylation