Nitrate assimilatory

Bowsher, C.G., Long, D.M., Oaks, A. Rothstein, S.J. (1991). Effect of light/dark cycles on expression of nitrate assimilatory genes in maize shoots and roots. Plant Physiology 95, 281-5. 

Assimilatory nitrate reduction Conversion of nitrate to reduced forms of nitrogen, generally ammonium, for the synthesis of amino acids and proteins. 

Assimilatory denitrifiers reduce nitrate to the amino acid level where it is incorporated into protein. Many plants and bacteria can do this and, therefore, use nitrate as a nitrogen source. 

Assimilatory nitrate reduction is the reduction of NOT, followed by uptake of the nitrogen by the organism as biomass. 

There are four different classes of nitrate reductases (234). The nitrate reductases from D. desulfuricans show a strong homology to the a-subunit of the class of periplasmic respiratory nitrate reductases, and also to some of the enzymes that are included on the class of cytoplasmic assimilatory nitrate reductases. Because of this fact, a proposal was made for a new class of monomeric NAP, which contains the minimal arrangement of metal centers to perform nitrate reduction one [4Fe-4S] cluster and a Mo bound to two MGD. 

Figure 3. The general nitrogen model for illustrating the bio geochemical cycling in Forest ecosystems. Explanations for the fluxes 1, ammonia volatilization 2, forest fertilization 3, N2-fixation 4, denitrification 5, nitrate respiration 6, nitrification 7, immobilization 8, mineralization 9, assimilatory and dissimilatory nitrate reduction to ammonium 10, leaching 11, plant uptake 12, deposition N input 13, residue composition, exudation 14, soil erosion 15, ammonium fixation and release by clay minerals 16, biomass combustion 17, forest harvesting 18, litterfall (Bashkin, 2002).

In the second step, 32 mol O2 are generated from the assimilatory reduction of nitrate, in the form of HNO3, amine nitrogen, represented in the organic molecule as (NHj) , ... 

Nitrogen uptake that results in the formation of new biomolecules is termed an assimilation process, such as assimilatory nitrogen reduction. The processes that result in the release of DIN into seawater are referred to as dissimilations, such as dissimi-latory nitrogen reduction. An example of the latter is denitrification, in which nitrate and nitrite obtained from seawater serve as electron acceptors to enable the oxidation of organic matter. This causes the nitrate and nitrite to be transformed into reduced species, such as N2O and N2, which are released back into seawater. 

This process is commonly referred to as assimilatory nitrogen (nitrate or nitrite) reduction. The electrons for these reductions are supplied by half-cell oxidations involving NADPH/NADP" and NADH/NAD" (Table 7.11). All of these reactions and membrane transport processes are mediated by enzymes that are specific to the DIN species. Considerable variation exists among the phytoplankton species in their ability to produce the necessary enzymes. Since marine phytoplankton are often nitrogen limited, the quantity and type of DIN available in the water column can greatly influence overall phytoplankton abundance and species diversity. 

Like assimilatory nitrogen reduction, denitrification proceeds through a series of steps with nitrate first being reduced to nitrite, followed by reduction of nitrite to N2(g). Under some conditions, N20(g) is also produced. 

Assimilatory nitrate reduetion The reduction of nitrate to organic nitrogen compounds that constitute the tissues of marine organisms. Plankton and some bacteria assimilate nitrogen via this process. 

Nitrate reductase (NADH) [EC 1.6.6.1], also known as assimilatory nitrate reduetase, eatalyzes the reaction of NADH with nitrate to produee NAD+, nitrite, and water. This enzyme uses FAD or FMN, heme, and a molybdenum ion as eofaetors. (2) Nitrate reductase (NAD(P)H) [EC 1.6.6.2], also known as assimilatory nitrate reduetase, eatalyzes the reaetion of NAD(P)H with nitrate to produee NAD(P)+, nitrite, and water. This enzyme uses FAD or FMN, heme, and a molybdenum ion as eofaetors. (3) Nitrate reductase (NADPH) [EC 1.6.6.3] eatalyzes the reaetion of NADPH with nitrate to produee NADP+, nitrite, and water. This enzyme uses FAD, heme, and a molybdenum ion as cofactors. (4) Nitrate reduetase (eytoehrome) [EC 1.9.6.1] catalyzes the reaetion of nitrate with ferroeytochrome to produce nitrite and ferrieytoehrome. (5) Nitrate reductase (ac-eeptor) [EC 1.7.99.4], also known as respiratory nitrate... 

Assembly-induced GTP hydrolysis, MICROTUBULE ASSEMBLY KINETICS Assimilatory nitrate reductase,... 

The fact that the cytochrome P-450 was induced even in the presence of NH3, which is the end product of assimilatory N-oxide reductions, suggested that it might funciton in dissimilatory N-oxide reductions. Anaerobic growth experiments with induced cells showed that reduction of nitrate to nitrite was energy yielding in F. oxysporum but reduction of nitrite to N2O was probably not (Shoun and Tanimoto, 1991). 

The assimilatory nitrate reductase (Eq. 16-61) of fungi and green plants (Chapter 24) also belongs to the sulfite oxidase family. 

The opposite sequence, reduction of nitrate and nitrite ions, provides a major route of acquisition of ammonia for incorporation into cells by bacteria, fungi, and green plants (Fig. 24-1). Assimilatory (biosynthetic) nitrate reductases catalyze the two-electron reduction of nitrate to nitrite (Eq. 16-61). This is thought to occur at the molybdenum atom of the large 900-residue highly regulated793 molybdopterin-dependent enzyme. In green plants the reductant is... 

Bacterial assimilatory nitrate reductases have similar properties.86/86a In addition, many bacteria, including E. coli, are able to use nitrate ions as an oxidant for nitrate respiration under anaerobic conditions (Chapter 18). Tire dissimilatory nitrate reductases involved also contain molybdenum as well as Fe-S centers.85 Tire E. coli enzyme receives electrons from reduced quinones in the plasma membrane, passing them through cytochrome b, Fe-S centers, and molybdopterin to nitrate. The three-subunit aPy enzyme contains cytochrome b in one subunit, an Fe3S4 center as well as three Fe4S4 clusters in another, and the molybdenum cofactor in the third.87 Nitrate reduction to nitrite is also on the pathway of denitrification, which can lead to release of nitrogen as NO, NzO, and N2 by the action of dissimi-latory nitrite reductases. These enzymes873 have been discussed in Chapters 16 and 18. 

Generally, the assimilatory nitrate and nitrite reductases are soluble enzymes that utilize reduced pyridine nucleotides or reduced ferrodoxin. In contrast, the dissimilatory nitrate reductases are membrane-bound terminal electron acceptors that are tightly linked to cytochrome by pigments. Such complexes allow one or more sites of energy conservation (ATP generation) coupled with electron transport. 

Nitrate reductase (assimilatory) Chloretta vulgaris 360000 4 4 heme, 4FAD... 

Nitrate reductases have been isolated from bacteria, plants and fungi and always contain molybdenum. Two types may be distinguished (a) the assimilatory nitrate reductases which catalyze the reduction of nitrate to nitrite, which ultimately is reduced to ammonia and used by... 

Nitrate reductase from Chlorella, an assimilatory enzyme, is a homotetramer of molecular weight 360 000 and contains one each of Mo, heme and FAD per subunit. The nitrate reductase from E. coli is a dissimilatory enzyme. EXAFS data are available on the molybdenum sites in both enzymes (Table 24).1050 The environment of the molybdenum in the assimilatory enzyme is similar to that found for sulfite oxidase, with at least two sulfur ligands near the molybdenum and a shuttle between monoxo and dioxo forms with redox change in the enzyme. This allows a similar mechanism to be put forward for the assimilatory nitrate reductase,1051 shown in equation (57), where an oxo group is transferred from nitrate to MoIV with production of nitrite and MoVI. 

The assimilatory nitrate reductase from Chlorella contains the molybdenum cofactor, as evidenced by the ability of the enzyme to donate the cofactor to the nitrate reductase of the mutant nit-1 of N. crassa. Reduction of the enzyme with NADH gives the Mov ESR signal, which is abolished on reoxidation with nitrate. Line shape and g values of the signal show a pH dependence similar to those observed previously for sulfite oxidase. The signal observed at pH 7.0 shows evidence for interaction with a single exchangeable proton.1053... 

Ammonia is oxidized in nature to nitrate via several intermediates in the process of nitrification. Nitrate may be reduced to nitrite by either a dissimilatory or an assimilatory process. Nitrite may be assimilated into the cell via reduction to ammonia, or it may be reduced by microorganisms to N20 and N2 in denitrification. A major part of the total nitrogen in this pathway is lost to the atmosphere. However, in turn, atmospheric dinitrogen is converted to ammonia by various bacteria in nitrogen fixation. 

As noted in Section 62.1.9.6, reduction of nitrate may occur by assimilatory or dissimilatory pathways. In the former case, the nitrate produced is reduced further to ammonia, which is incorporated into the cell. In the latter case, nitrate is reduced anaerobically to nitrite, serving as an electron acceptor in the respiration of facultative or a few obligate anaerobic bacteria. The example of Escherichia coli has been considered in Section 62.1.13.4.3. This process is usually terminated at nitrite, which accumulates around the cells, but may proceed further1511 as nitrite-linked respiration in the process of denitrification. 

Both assimilatory and dissimilatory nitrate reductases are molybdoenzymes, which bind nitrate at the molybdenum. EXAFS studies1050 have shown that there are structural differences between the assimilatory nitrate reductase from Chlorella vulgaris and the dissimilatory enzyme from E. coli. The Chlorella enzyme strongly resembles sulfite oxidase1050,1053 and shuttles between mon-and di-oxo forms, suggesting an oxo-transfer mechanism for reduction of nitrate. This does not appear to be the case for the E. coli enzyme, for which an oxo-transfer mechanism seems to be unlikely. The E. coli enzyme probably involves an electron transfer and protonation mechanism for the reduction of nitrate.1056 It is noteworthy that the EXAFS study on the E. coli nitrate reductase showed a long-distance interaction with what could be an electron-transfer subunit. 

Green plants, algae, fungi, cyanobacteria and bacteria that assimilate nitrate also produce assimilatory nitrite reductases, which catalyze the six-electron reduction of nitrite to ammonia (equation 89). The formation of heme-nitrosyl intermediates has been detected in several cases,1515 while hydroxylamine is commonly thought to be an intermediate. Added hydroxylamine is rapidly reduced to ammonia. However, no intermediates are released, and ammonia is the only product... 

Barber, M.J. Solomonson, L.P. (1986). The role of the essential sulfhydryl group in assimilatory NADH nitrate reductase of Chlorella. Journal of Biological Chemistry 261, 4562-7. 

Campbell, W.H. Kinghom, J.R. (1990). Functional domains of assimilatory nitrate reductases and nitrite reductases. Trends in Biochemistry 15, 315-19. 

Guerrero, M.G., Vega, J.M. Losada, M. (1981). The assimilatory nitrate-reducing system and its regulation. Annual Review of Plant Physiology 32, 169-201. 

LS, K.H.D. Lederer, F. (1983). On the presence of a heme-binding domain homologous to cytochrome b5 in Neurospora crassa assimilatory nitrate reductase. The EMBO Journal 2, 1909-14. 

Nason, A., Lee, K.Y., Pan, S.-S., Ketchum, P.A., Lamberti, A. Davies, J. (1971). In vitro formation of assimilatory reduced nicotinamide adenine dinucleotide phosphate nitrate reductase from a Neurospora mutant and a component of molybdenum-enzymes. Proceedings of the National Academy of Sciences (USA) 68, 3242-6. 

---