Diazótrofo
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Los diazótrofos son bacterias que hacen fijación de nitrógeno atmosférico en una forma más disponible como es el amonio (Postgate, 1998).
Un diazótrofo es un organismo que es capaz de crecer sin fuentes externas de nitrógeno fijado. Ejs. de organismos que hacen esto son los Bradyrhizobia, Frankia (en simbiosis) y Azospirillum (en vida libre). Todos los diazótrofos contienen sistemas hierro-molibdeno nitrogenasas. Dos de los más estudiados sistemas son sobre Klebsiella pneumoniae y Azotobacter vinlandii. Esos sistemas se han usado debido a su simpicidad genética (trazabilidad) y su rápido crecimiento y multiplicación (Dixon and Kahn 2004).
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[editar] Tipos de diazótrofos
Los diazótrofos se entremezclan entre grupos taxonómicos bacteriales (mayormente en ls Eubacteria pero también hay un par en Archaea). Inclusive dentro de una especie que puede fijar nitrógeno puede haber razas que no lo fijan (Postgate, 1998). La fijación se interrumpe cuando otras fuentes de nitrógeno están disponibles, y, en muchas especies, cuadno el oxígeno está a una presión parcial alta. Las bacterias tienen diferentes caminos de soportar los efectos debilitantes del oxígeno sobre las nitrogenasas, listadas más abajo.
[editar] Diazótrofos de vida libre
- Anaerobios -- son anaerobios obligados que no toleran oxígeno salvo si no están fijando nitrógeno. 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 (e.g. Desulfovibrio), and some Archean methanogens fix nitrogen in muds and animal intestines (Postgate 1998).
- Anaerobios facultativos -- Estas especies pueden crecer tanto con oxígeno como sin él, pero sólo fijan nitrógeno de forma anaerobia. A menudo, respiran el oxígeno tan pronto como lo consiguen manteniendo los niveles de oxígeno libre bajos. Algunos ejemplos son Klebsiella pneumoniae, Bacillus polymyxa, B. macerans y Escherichia intermedia (Postgate, 1998).
- 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 (Postgate 1998).
- Phototrophs -- 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 (heterocysts) 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 (e.g. Plectonema) (Postgate 1998).
[editar] Diazótrofos simbióticos
- Rhizobia -- these are the species that associate with legumes, plants of the Fabaceae family. 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 (Postgate 1998).
- Frankias -- much less is known about these 'actinorhizal' nitrogen fixers. The bacteria also infect the roots and form nodule-like structures. Frankia forms heterocyst-like structures in these nodules where N-fixation occurs (Vessey et al., 2005). Frankias also produce hemoglobins (Beckwith et al., 2002), but their role is less well established than for rhizobia (vessey et al., 2005). Although at first it appeared that they infect sets of unrelated plants (alders, Australian pines, California lilac, bog myrtle, bitter brush, Dryas), revisions to the phylogeny of angiosperms show a close relatedness of these species and the legumes (Soltis et al., 1995; Vessey et al. 2005).
- Cyanobacteria -- there are also symbiotic cyanobacteria. Some associate with fungi as lichens, with liverworts, with a fern, and with a cycad (Postgate, 1998). These do not form nodules (indeed most of the plants do not have roots). Heterocysts sequester the oxygen, as discussed above. The fern association is important agriculturally: the water fern Azolla harbouring Anabaena is an important green manure for rice culture (Postgate, 1998).
- Asociación con animales -- although diazotrophs have been found in many animal guts, there is usually sufficient ammonia present to suppress nitrogen fixation (Postgate 1998). 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 (Postgate 1998).
[editar] Importancia
En términos de generar nitrógeno disponible a otros organismos, las asociaciones simbióticas grandemente exceden a las spp. de vida libre con la excepción de cianobacterias (Postgate, 1998).
[editar] Referencias
Beckwith, J, Tjepkema, J D, Cashon, R E, Schwintzer, C R, Tisa, L S (2002). "Hemoglobin in five genetically diverse Frankia strains". Can J Microbiol 48: 1048-1055.
Dixon R and Kahn D (2004). "Genetic regulation of biological nitrogen fixation". Nat Rev Microbiol 2 (8): 621-31.
Postgate, J (1998). Nitrogen Fixation, 3rd Edition. Cambridge University Pres, Cambridge UK.
Soltis DE, Soltis PS, Morgan DR, Swensen SM, Mullin BC, Dowd JM, Martin PG (1995). "Chloroplast gene sequence data suggest a single origin of the predisposition for symbiotic nitrogen fixation in angiosperms". Proc Natl Acad Sci USA 92: 2647 – 2651.
Vessey JK, Pawlowski, K and Bergman B (2005). "Root-based N2-fixing symbioses: Legumes, actinorhizal plants, Parasponia sp and cycads". Plant and soil 274 (1-2): 51-78.