Dynamic and steady-state responses of inorganic nitrogen pools and NH(3) exchange in leaves of Lolium perenne and Bromus erectus to changes in root nitrogen supply

Short- and long-term responses of inorganic N pools and plant-atmosphere NH(3) exchange to changes in external N supply were investigated in 11-week-old plants of two grass species, Lolium perenne and Bromus erectus, characteristic of N-rich and N-poor grassland ecosystems, respectively. A switch of root N source from NO(-)(3)to NH(4)(+) caused within 3 h a 3- to 6-fold increase in leaf apoplastic NH(4)(+) concentration and a simultaneous decrease in apoplastic pH of about 0.4 pH units in both species. The concentration of total extractable leaf tissue NH(4)(+) also increased two to three times within 3 h after the switch. Removal of exogenous NH(4)(+) caused the apoplastic NH(4)(+) concentration to decline back to the original level within 24 h, whereas the leaf tissue NH(4)(+)concentration decreased more slowly and did not reach the original level in 48 h. After growing for 5 weeks with a steady-state supply of NO(-)(3)or NH(4)(+), L. perenne were in all cases larger, contained more N, and utilized the absorbed N more efficiently for growth than B. erectus, whereas the two species behaved oppositely with respect to tissue concentrations of NO(-)(3), NH(4)(+), and total N. Ammonia compensation points were higher for B. erectus than for L. perenne and were in both species higher for NH(4)(+)- than for NO(-)(3)-grown plants. Steady-state levels of apoplastic NH(4)(+), tissue NH(4)(+), and NH(3) emission were significantly correlated. It is concluded that leaf apoplastic NH(4)(+) is a highly dynamic pool, closely reflecting changes in the external N supply. This rapid response may constitute a signaling system coordinating leaf N metabolism with the actual N uptake by the roots and the external N availability.     

Diurnal variation in uptake and xylem contents of inorganic and assimilated N under continuous and interrupted N supply to Phleum pratense and Festuca pratensis

Compensation by dark-period uptake of NH(4)(+) and NO(3)(-) in the grasses Phleum pratense L. and Festuca pratensis Huds. following N deprivation during the preceding light period was investigated in flowing solution culture under an artificial 10/14 h light/dark cycle. N was supplied as either NO(3)(-), NH(4)(+) or NH(4)NO(3) at 20+/-5 mmol m(-3), available continuously or only during the dark period, for 5-10 d. Intermittent N supply did not affect total daily N uptake, growth rate or net partitioning of dry matter. Net uptake and influx of NO(3)(-) varied similarly throughout the diurnal cycle when NO(3)(-) was supplied continuously, with a marginal contribution by NO(3)(-) efflux. Influx was significantly higher and efflux slightly higher following interruption of NO(3)(-) supply during the light period. Nitrate accounted for 80% of N in xylem exudate except between hours 6-9 of the light period when the amino acid concentration increased 3-fold, primarily as glutamine. Diurnal variation in relative NO(3)(-) uptake exhibited five phases of constant acceleration/deceleration, described reasonably well assuming NO(3)(-) influx was subject to metabolic co-regulation by NO(3)(-) and amino acid levels in the cytoplasmic compartment of the roots. Accordingly, influx is determined by variation in root NO(3)(-) levels throughout the dark period and the first half of the light period, but is down-regulated by increased amino acid levels during the second half of the light period. The sharp light/dark transitions affect transpiration rate and hence xylem N flux which, in turn, affect NO(3)(-) levels in the cytoplasmic compartment of the roots and the rate of NO(3)(-) assimilation in the shoot.     

Abscisic acid (ABA) flows from Hordeum vulgare to the hemiparasite Rhinanthus minor and the influence of infection on host and parasite abscisic acid relations

Using the facultative root hemiparasite Rhinanthus minor and Hordeum vulgare as a host, the flows, depositions, and metabolism of abscisic acid (ABA) within the host, within the parasite, and between host and parasite have been studied. When the plants were supplied with 5 mM NO(3)(-), there were weak or no effects of parasitism on ABA flows, biosynthesis, and ABA degradation in barley. However, ABA deposition was significantly affected in the leaf laminae (3-fold) and in the leaf sheath (2.4-fold), but not in roots. Dramatic changes in ABA flows, metabolism, and deposition on a per plant basis, however, have been observed in Rhinanthus. Biosynthesis in the roots was 12-fold higher after attachment, resulting in 14-fold higher ABA flows in the xylem. A large portion of this ABA was metabolized, a small portion was deposited. Phloem flows of ABA were increased 13-fold after attachment. The concentrations of ABA in tissues and transport fluids were higher in attached Rhinanthus by an order of magnitude than in host tissues and xylem sap. The same tendency was also found in a comparison between single Rhinanthus and unparasitized barley. As compared with 5 mM NO(3)(-), lower NO(3)(-) or 1 mM NH(4)(+) supply doubled the ABA concentrations in barley leaf laminae, while having only small or non-significant effects in the other organs. The possible function of ABA for the parasite is discussed.     

Deposition of ammonium and nitrate in the roots of maize seedlings supplied with different nitrogen salts

This study measured total osmolarity and concentrations of NH(4)(+), NO(3)(-), K(+), soluble carbohydrates, and organic acids in maize seminal roots as a function of distance from the apex, and NH(4)(+) and NO(3)(-) in xylem sap for plants receiving NH(4)(+) or NO(3)(-) as a sole N-source, NH(4)(+) plus NO(3)(-), or no nitrogen at all. The disparity between net deposition rates and net exogenous influx of NH(4)(+) indicated that growing cells imported NH(4)(+) from more mature tissue, whereas more mature root tissues assimilated or translocated a portion of the NH(4)(+) absorbed. Net root NO(3)(-) influx under Ca(NO(3))(2) nutrition was adequate to account for pools found in the growth zone and provided twice as much as was deposited locally throughout the non-growing tissue. In contrast, net root NO(3)(-) influx under NH(4)NO(3) was less than the local deposition rate in the growth zone, indicating that additional NO(3)(-) was imported or metabolically produced. The profile of NO(3)(-) deposition rate in the growth zone, however, was similar for the plants receiving Ca(NO(3))(2) or NH(4)NO(3). These results suggest that NO(3)(-) may serve a major role as an osmoticant for supporting root elongation in the basal part of the growth zone and maintaining root function in the young mature tissues.     

Growth response of barley and tomato to nitrogen stress and its control by abscisic acid,water relations and photosynthesis

Barley (Hordeum vulgare L.) and tomato Lycopersicon esculentum Mill.) were grown hydroponically and examined 2, 5, and 10 d after being deprived of nitrogen (N) supply. Leaf elongation rate declined in both species in response to N stress before there was any reduction in rate of dryweight accumulation. Changes in water transport to the shoot could not explain reduced leaf elongation in tomato because leaf water content and water potential were unaffected by N stress at the time leaf elongation began to decline. Tomato maintained its shoot water status in N-stressed plants, despite reduced water absorption per gram root, because the decline in root hydraulic conductance with N stress was matched by a decline in stomatal conductance. In barley the decline in leaf elongation coincided with a small (8%) decline in water content per unit area of young leaves; this decline occurred because root hydraulic conductance was reduced more strongly by N stress than was stomatal conductance. Nitrogen stress caused a rapid decline in tissue NO ₃ ^- pools and in NO ₃ ^- flux to the xylem, particularly in tomato which had smaller tissue NO ₃ ^- reserves. Even in barley, tissue NO ₃ ^- reserves were too small and were mobilized too slowly (60% in 2 d) to support maximal growth for more than a few hours. Organic N mobilized from old leaves provided an additional N source to support continued growth of N-stressed plants. Abscisic acid (ABA) levels increased in leaves of both species within 2 d in response to N stress. Addition of ABA to roots caused an increase in volume of xylem exudate but had no effect upon NO ₃ ^- flux to the xylem. After leaf-elongation rate had been reduced by N stress, photosynthesis declined in both barley and tomato. This decline was associated with increased leaf ABA content, reduced stomatal conductance and a decrease in organic N content. We suggest that N stress reduces growth by several mechanisms operating on different time scales: (1) increased leaf ABA content causing reduced cell-wall extensibility and leaf elongation and (2) a more gradual decline in photosynthesis caused by ABA-induced stomatal closure and by a decrease in leaf organic N.Abbreviation and symbols ABA abscisic acid - c_i leaf internal CO₂ concentration - L_p root hydraulic conductance     

Bacterial immobilization and remineralization of N at different growth rates and N concentrations

An experiment was designed to resolve two largely unaddressed questions about the turnover of N in soils. One is the influence of microbial growth rate on mobilization and remineralization of cellular N. The other is to what extent heterotrophic immobilization of NO(3)(-) is controlled by the soil concentration of NH(4)(+). Bacteria were extracted from a deciduous forest soil and inoculated into an aqueous medium. Various N pool dilution/enrichment experiments were carried out to: (1) calculate the gross N immobilization and remineralization rates; (2) investigate their dependence on NH(4)(+)and NO(3)(-) concentrations; (3) establish the microbial preference for NH(4)(+)and NO(3)(-) depending on the NH(4)(+)/NO(3)(-) concentration ratio. Remineralization of microbial N occurred mainly at high growth rates and NH(4)(+) concentrations. There was a positive correlation between NH(4)(+) immobilization and remineralization rates, and intracellular recycling of N seemed to be an efficient way for bacteria to withstand low inorganic N concentrations. Thus, extensive remineralization of microbial N is likely to occur only when environmental conditions promote high growth rates. The results support previous observations of high NO(3)(-) immobilization rates, especially at low NH(4)(+) concentrations, but NO(3)(-) was also immobilized at high NH(4) concentrations. The latter can be understood if part of the microbial community has a preference for NO(3)(-) over NH(4)(+).     

NH4+-stimulated and -inhibited components of K+ transport in rice (Oryza sativa L.)

The disruption of K(+) transport and accumulation is symptomatic of NH(4)(+) toxicity in plants. In this study, the influence of K(+) supply (0.02-40 mM) and nitrogen source (10 mM NH(4)(+) or NO(3)(-)) on root plasma membrane K(+) fluxes and cytosolic K(+) pools, plant growth, and whole-plant K(+) distribution in the NH(4)(+)-tolerant plant species rice (Oryza sativa L.) was examined. Using the radiotracer (42)K(+), tissue mineral analysis, and growth data, it is shown that rice is affected by NH(4)(+) toxicity under high-affinity K(+) transport conditions. Substantial recovery of growth was seen as [K(+)](ext) was increased from 0.02 mM to 0.1 mM, and, at 1.5 mM, growth was superior on NH(4)(+). Growth recovery at these concentrations was accompanied by greater influx of K(+) into root cells, translocation of K(+) to the shoot, and tissue K(+). Elevating the K(+) supply also resulted in a significant reduction of NH(4)(+) influx, as measured by (13)N radiotracing. In the low-affinity K(+) transport range, NH(4)(+) stimulated K(+) influx relative to NO(3)(-) controls. It is concluded that rice, despite its well-known tolerance to NH(4)(+), nevertheless displays considerable growth suppression and disruption of K(+) homeostasis under this N regime at low [K(+)](ext), but displays efficient recovery from NH(4)(+) inhibition, and indeed a stimulation of K(+) acquisition, when [K(+)](ext) is increased in the presence of NH(4)(+).     

Root-zone acidity and nitrogen source affects Typha latifolia L. growth and uptake kinetics of ammonium and nitrate

The NH(4)(+) and NO(3)(-) uptake kinetics by Typha latifolia L. were studied after prolonged hydroponics growth at constant pH 3.5, 5.0, 6.5 or 7.0 and with NH(4)(+) or NO(3)(-) as the sole N-source. In addition, the effects of pH and N source on H(+) extrusion and adenine nucleotide content were examined. Typha latifolia was able to grow with both N sources at near neutral pH levels, but the plants had higher relative growth rates, higher tissue concentrations of the major nutrients, higher contents of adenine nucleotides, and higher affinity for uptake of inorganic nitrogen when grown on NH(4)(+). Growth almost completely stopped at pH 3.5, irrespective of N source, probably as a consequence of pH effects on plasma membrane integrity and H(+) influx into the root cells. Tissue concentrations of the major nutrients and adenine nucleotides were severely reduced at low pH, and the uptake capacity for inorganic nitrogen was low, and more so for NO(3)(-)-fed than for NH(4)(+)-fed plants. The maximum uptake rate, V(max), was highest for NH(4)(+) at pH 6.5 (30.9 micro mol h(-1) g(-1) root dry weight) and for NO(3)(-) at pH 5.0 (31.7 micro mol h(-1) g(-1) root dry weight), and less than 10% of these values at pH 3.5. The affinity for uptake as estimated by the half saturation constant, K((1/2)), was lowest at low pH for NH(4)(+) and at high pH for NO(3)(-). The changes in V(max) and K((1/2)) were thus consistent with the theory of increasing competition between cations and H(+) at low pH and between anions and OH(-) at high pH. C(min) was independent of pH, but slightly higher for NO(3)(-) than for NH(4)(+) (C(min)(NH(4)(+)) approximately 0.8 mmol m(-3); C(min)(NO(3)(-)) approximately 2.8 mmol m(-3)). The growth inhibition at low pH was probably due to a reduced nutrient uptake and a consequential limitation of growth by nutrient stress. Typha latifolia seems to be well adapted to growth in wetland soils where NH(4)(+) is the prevailing nitrogen compound, but very low pH levels around the roots are very stressful for the plant. The common occurrence of T. latifolia in very acidic areas is probably only possible because of the plant's ability to modify pH-conditions in the rhizosphere.     

Nitrate does not result in iron inactivation in the apoplast of sunflower leaves

It has been hypothesized that nitrate (NO(3)(-)) nutrition might induce iron (Fe) deficiency chlorosis by inactivation of Fe in the leaf apoplast (H.U. Kosegarten, B. Hoffmann, K. Mengel [1999] Plant Physiol 121: 1069-1079). To test this hypothesis, sunflower (Helianthus annuus L. cv Farnkasol) plants were grown in nutrient solutions supplied with various nitrogen (N) forms (NO(3)(-), NH(4)(+) and NH(4)NO(3)), with or without pH control by using pH buffers [2-(N-morpholino)ethanesulfonic acid or 4-(2-hydroxyethyl)-1-piperazineethanesulfonic acid]. It was shown that high pH in the nutrient solution restricted uptake and shoot translocation of Fe independently of N form and, therefore, induced Fe deficiency chlorosis at low Fe supply [1 micro M ferric ethylenediaminedi(O-hydroxyphenylacetic acid)]. Root NO(3)(-) supply (up to 40 mM) did not affect the relative distribution of Fe between leaf apoplast and symplast at constant low external pH of the root medium. Although perfusion of high pH-buffered solution (7.0) into the leaf apoplast restricted (59)Fe uptake rate as compared with low apoplastic solution pH (5.0 and 6.0, respectively), loading of NO(3)(-) (6 mM) showed no effect on (59)Fe uptake by the symplast of leaf cells. However, high light intensity strongly increased (59)Fe uptake, independently of apoplastic pH or of the presence of NO(3)(-) in the apoplastic solution. Finally, there are no indications in the present study that NO(3)(-) supply to roots results in the postulated inactivation of Fe in the leaf apoplast. It is concluded that NO(3)(-) nutrition results in Fe deficiency chlorosis exclusively by inhibited Fe acquisition by roots due to high pH at the root surface.     

Response to ammonium and nitrate by a mycorrhizal annual invasive grass and native shrub in southern California

The goal of this study was to determine the interaction of mycorrhizae and two N sources, ammonium (NH(4)(+)) and nitrate (NO(3)(-)), on the growth of a coastal sage scrub (CSS) species, Artemisia californica, and an exotic annual grass, Bromus madritensis ssp. rubens. Anthropogenic nitrogen deposition may be influencing the decline of CSS and replacement by exotic grasses, but the extent to which mycorrhizae are involved in shrubland decline is unknown. NO(3)(-) is the dominant form of deposition in southern California, although the native, uneutrophied soils have a greater concentration of NH(4)(+). Seeds of each species were germinated in pots of sterile soil, inoculated with native soil containing mycorrhizal spores and infective root fragments, and fertilized with 50 μg/g of either NO(3)(-) or NH(4)(+). NH(4)(+) enhanced the growth of both mycorrhizal species, while NO(3)(-) did not. Control plants of B. madritensis under low N had a significant response to mycorrhizae, but A. californica did not. Nitrate increased the growth of nonmycorrhizal A. californica as much as the mycorrhizal NH(4)(+)-treated plants. There is no evidence in this study to suggest that the decline of A. californica or increase in B. madritensis is due to a mycorrhizal response to NO(3)(-). Other life history traits of the two species must be used to explain the invasive behavior of the annual grass. Mycorrhizae may be more important in controlling plant growth in native uneutrophied soils dominated by NH(4)(+) rather than NO(3)(-).     

Transpiration, potassium uptake and flow in tobacco as affected by nitrogen forms and nutrient levels

BACKGROUND AND AIMS: Ammonium can result in toxicity symptoms in many plants when it is supplied as the sole source of N. In this work, influences of different nitrogen forms at two levels (2 and 15 mm N) on growth, water relations and uptake and flow of potassium were studied in plants of Nicotiana tabacum 'K 326'. METHODS: Xylem sap from different leaves was collected from 106-d-old tobacco plants cultured in quartz sand by application of pressure to the root system. Whole-shoot transpiration for each of the treatments was measured on a daily basis by weight determination. KEY RESULTS: Total replacement of NO(3)(-)N by NH(4)(+)-N caused a substantial decrease in dry weight gain, even when plants grew under nutrient deficiency. Increasing nutrient concentration resulted in a greater net dry weight gain when nitrogen was supplied as NO(3)(-) or NH(4)NO(3), but resulted in little change when nitrogen was supplied as NH(4)(+). NH(4)(+)-N as the sole N-source also caused reduction in transpiration rate, changes in plant WUE (which depended on the nutrient levels) and a decrease in potassium uptake. However, the amount of xylem-transported potassium in the plants fed with NH(4)(+) was not reduced: it was 457 % or 596 % of the potassium currently taken up at low or high nutrient level, respectively, indicating a massive export from leaves and cycling of potassium in the phloem. CONCLUSIONS: Ammonium reduces leaf stomatal conductance of tobacco plants. The flow and partitioning of potassium in tobacco plants can be changed, depending on the nitrogen forms and nutrient levels.     

The regulation of ammonium translocation in plants

Much controversy exists about whether or not NH(+)(4) is translocated in the xylem from roots to shoots. In this paper it is shown that such translocation can indeed take place, but that interference from other metabolites such as amino acids and amines may give rise to large uncertainties about the magnitude of xylem NH(+)(4) concentrations. Elimination of interference requires sample stabilization by, for instance, formic acid or methanol. Subsequent quantification of NH(+)(4) should be done by the OPA-fluorometric method at neutral pH with 2-mercaptoethanol as the reducing agent since this method is sensitive and reliable. Colorimetric methods based on the Berthelot reaction should never be used, as they are prone to give erroneous results. Significant concentrations of NH(+)(4), exceeding 1 mM, were measured in both xylem sap and leaf apoplastic solution of oilseed rape and tomato plants growing with NO(-)(3) as the sole N source. When NO(-)(3) was replaced by NH(+)(4), xylem sap NH(+)(4) concentrations increased with increasing external concentrations and with time of exposure to NH(+)(4). Up to 11% of the translocated N was constituted by NH(+)(4). Glutamine synthetase (GS) incorporates NH(+)(4) into glutamine, but root GS activity and expression were repressed when high levels of NH(+)(4) were supplied. Ammonium concentrations measured in xylem sap sampled just above the stem base were highly correlated with NH(+)(4) concentrations in apoplastic solution from the leaves. Young leaves tended to have higher apoplastic NH(+)(4) concentrations than older non-senescing leaves. The flux of NH(+)(4) (concentration multiplied by transpirational water flow) increased with temperature despite a decline in xylem NH(+)(4) concentration. Retrieval of leaf apoplastic NH(+)(4) involves both high and low affinity transporters in the plasma membrane of mesophyll cells. Current knowledge about these transporters and their regulation is discussed.     

Effect of nitrogen stress and abscisic acid on nitrate absorption and transport in barley and tomato

The potential of barley (Hordeum vulgare L.) and tomato (Lycopersicon esculentum Mill.) roots for net NO ₃ ^- absorption increased two-to five fold within 2 d of being deprived of NO ₃ ^- supply. Nitrogen-starved barley roots continued to maintain a high potential for NO ₃ ^- absorption, whereas NO ₃ ^- absorption by tomato roots declined below control levels after 10 d of N starvation. When placed in a 0.2 mM NO ₃ ^- solution, roots of both species transported more NO ₃ ^- and total solutes to the xylem after 2 d of N starvation than did N-sufficient controls. However, replenishment of root NO ₃ ^- stores took precedence over NO ₃ ^- transport to the xylem. Consequently, as N stress became more severe, transport of NO ₃ ^- and total solutes to the xylem declined, relative to controls. Nitrogen stress caused an increase in hydraulic conductance (L _p) and exudate volume (J _v) in barley but decrased these parameters in tomato. Nitrogen stress had no significant effect upon abscisic acid (ABA) levels in roots of barley or flacca (a low-ABA mutant) tomato, but prevented an agerelated decline in ABA in wild-type tomato roots. Applied ABA had the same effect upon barley and upon the wild type and flacca tomatoes: L _p and J _v were increased, but NO ₃ ^- absorption and NO ₃ ^- flux to the xylem were either unaffected or sometimes inhibited. We conclude that ABA is not directly involved in the normal changes in NO ₃ ^- absorption and transport that occur with N stress in barley and tomato, because (1) the root ABA level was either unaffected by N stress (barley and flacca tomato) or changed, after the greatest changes in NO ₃ ^- absorption and transport and L _p had been observed (wild-type tomato); (2) changes in NO ₃ ^- absorption/transport characteristics either did not respond to applied ABA, or, if they did, they changed in the direction opposite to that predicted from changes in root ABA with N stress; and (3) the flacca tomato (which produces very little ABA in response to N stress) responded to N stress with very similar changes in NO ₃ ^- transport to those observed in the wild type.Abbreviation and symbols ABA abscisic acid - J_v exudate volume - L_p root hydraulic conductance     

Apoplastic pH and Fe(3+) reduction in intact sunflower leaves

It has been hypothesized that under NO(3)(-) nutrition a high apoplastic pH in leaves depresses Fe(3+) reductase activity and thus the subsequent Fe(2+) transport across the plasmalemma, inducing Fe chlorosis. The apoplastic pH in young green leaves of sunflower (Helianthus annuus L.) was measured by fluorescence ratio after xylem sap infiltration. It was shown that NO(3)(-) nutrition significantly increased apoplastic pH at distinct interveinal sites (pH >/= 6.3) and was confined to about 10% of the whole interveinal leaf apoplast. These apoplastic pH increases presumably derive from NO(3)(-)/proton cotransport and are supposed to be related to growing cells of a young leaf; they were not found in the case of sole NH(4)(+) or NH(4)NO(3) nutrition. Complementary to pH measurements, the formation of Fe(2+)-ferrozine from Fe(3+)-citrate was monitored in the xylem apoplast of intact leaves in the presence of buffers at different xylem apoplastic pH by means of image analysis. This analysis revealed that Fe(3+) reduction increased with decreasing apoplastic pH, with the highest rates at around pH 5. 0. In analogy to the monitoring of Fe(3+) reduction in the leaf xylem, we suggest that under alkaline nutritional conditions at interveinal microsites of increased apoplastic pH, Fe(3+) reduction is depressed, inducing leaf chlorosis. The apoplastic pH in the xylem vessels remained low in the still-green veins of leaves with intercostal chlorosis.     

The effect of competition from neighbours on stomatal conductance in lettuce and tomato plants

Competition decreased transpiration from young lettuce plants after 2 days, before any reductions in leaf area became apparent, and stomatal conductance (g(s) ) of lettuce and tomato plants was also reduced. Stomatal closure was not due to hydraulic signals or competition for nutrients, as soil water content, leaf water status and leaf nitrate concentrations were unaffected by neighbours. Competition-induced stomatal closure was absent in an abscisic acid (ABA)-deficient tomato mutant, flacca, indicating a fundamental involvement of ABA. Although tomato xylem sap ABA concentrations were unaffected by the presence of neighbours, ABA/pH-based stomatal modulation is still likely to underlie the response to competition, as soil and xylem sap alkalization was observed in competing plants. Competition also modulated leaf ethylene production, and treatment of lettuce plants with an ethylene perception inhibitor (1-methylcyclopropene) diminished the difference in g(s) between single and competing plants grown in a controlled environment room, but increased it in plants grown in the greenhouse: ethylene altered the extent of the stomatal response to competition. Effects of competition on g(s) are discussed in terms of the detection of the absence of neighbours: increases in g(s) and carbon fixation may allow faster initial space occupancy within an emerging community/crop.     

Partial substitution of NO(3)(-) by NH(4)(+) fertilization increases ammonium assimilating enzyme activities and reduces the deleterious effects of salinity on the growth of barley

Productivity of cereal crops is restricted in saline soils but may be improved by nitrogen nutrition. In this study, the effect of ionic nitrogen form on growth, mineral content, protein content and ammonium assimilation enzyme activities of barley (Hordeum vulgare cv. Alexis L.) irrigated with saline water, was determined. Leaf and tiller number as well as plant fresh and dry weights declined under salinity (120 mM NaCl). In non-saline conditions, growth parameters were increased by application of NH(4)(+)/NO(3)(-) (25:75) compared to NO(3)(-) alone. Under saline conditions, application of NH(4)(+)/NO(3)(-) led to a reduction of the detrimental effects of salt on growth. Differences in growth between the two nitrogen regimes were not due to differences in photosynthesis. The NH(4)(+)/NO(3)(-) regime led to an increase in total N in control and saline treatments, but did not cause a large decrease in plant Na(+) content under salinity. Activities of GS (EC 6.3.1.2), GOGAT (EC 1.4.1.14), PEPC (EC 4.1.1.31) and AAT (EC 2.6.1.1) increased with salinity in roots, whereas there was decreased activity of the alternative ammonium assimilation enzyme GDH (EC 1.4.1.2). The most striking effect of nitrogen regime was observed on GDH whose salinity-induced decrease in activity was reduced from 34% with NO(3)(-) alone to only 14% with the mixed regime. The results suggest that the detrimental effects of salinity can be reduced by partial substitution of NO(3)(-) with NH(4)(+) and that this is due to the lower energy cost of N assimilation with NH(4)(+) as opposed to NO(3)(-) nutrition.     

Single-step nitrification models erroneously describe batch ammonia oxidation profiles when nitrite oxidation becomes rate limiting

Nitrification involves the sequential biological oxidation of reduced nitrogen species such as ammonium-nitrogen (NH(4)(+)-N) to nitrite-nitrogen (NO(2)(-)-N) and nitrate-nitrogen (NO(3)(-)-N). The adequacy of modeling NH(4)(+)-N to NO(3)(-)-N oxidation as one composite biochemical reaction was examined at different relative dynamics of NH(4)(+)-N to NO(2)(-)-N and NO(2)(-)-N to NO(3)(-)-N oxidation. NH(4)(+)-N to NO(2)(-)-N oxidation and NO(2)(-)-N to NO(3)(-)-N oxidation by a mixed nitrifying consortium were uncoupled using selective inhibitors allylthiourea and sodium azide. The kinetic parameters of NH(4)(+)-N to NO(2)(-)-N oxidation (q(max,ns) and K(S,ns)) and NO(2)(-)-N to NO(3)(-)-N oxidation (q(max,nb) and K(S,nb)) were determined by a rapid extant respirometric technique. The stoichiometric coefficients relating nitrogen removal, oxygen uptake and biomass synthesis were derived from an electron balanced equation. NH(4)(+)-N to NO(2)(-)-N oxidation was not affected by NO(2)(-)-N concentrations up to 100 mg NO(2)(-)-N L(-1). NO(2)(-)-N to NO(3)(-)-N oxidation was noncompetitively inhibited by NH(4)(+)-N but was not inhibited by NO(3)(-)-N concentrations up to 250 mg NO(3)(-)-N L(-1). When NH(4)(+)-N to NO(2)(-)-N oxidation was the sole rate-limiting step, complete NH(4)(+)-N to NO(3)(-)-N oxidation was adequately modeled as one composite process. However, when NH(4)(+)-N to NO(2)(-)-N oxidation and NO(2)(-)-N to NO(3)(-)-N oxidation were both rate limiting, the estimated lumped kinetic parameter estimates describing NH(4)(+)-N to NO(3)(-)-N oxidation were unrealistically high and correlated. These findings indicate that the use of single-step models to describe batch NH(4)(+) oxidation yields erroneous kinetic parameters when NH(4)(+)-to-NO(2)(-) oxidation is not the sole rate-limiting process throughout the assay. Under such circumstances, it is necessary to quantify NH(4)(+)-N to NO(2)(-)-N oxidation and NO(2)(-)-N to NO(3)(-)-N oxidation, independently.     

Transport and accumulation rates of abscisic acid and aldehyde oxidase activity in Pisum sativum L. in response to suboptimal growth conditions.

Pea plants (Pisum sativum L.) grown initially in nutrient solutions with adequate nitrogen supply (4 mM NO3-) were transferred to solutions containing salt (50 or 100 mM NaCl), ammonium (4 mM) or a low nitrogen supply (0.4 mM NO3-). No changes of abscisic acid (ABA) content were found in roots of stressed pea plants 9 d after the beginning of the treatments; however, accumulation of ABA in the leaves was observed. Old leaves accumulated ABA to a higher extent than young leaves. Accumulation of ABA in leaves of ammonium-fed plants and plants grown under low nitrogen supply occurred in the absence of both increased ABA xylem loading rate and enhanced aldehyde oxidase (AO, EC 1.2.3.1) activity in roots. Enhanced leaf AO activity was observed in all treatments, with the highest increase in old leaves. Among the three AO isoforms (AO-1, AO-2 and AO-3) detected in extracts of pea leaves, the lowest one AO-3 (highest mobility in the gel) correlated with ABA production and showed the highest increment in response to the treatments. The increase of AO activity detected in leaves after 2 weeks of stress application was less prominent than after 9 d, suggesting a transient enhancement of ABA production following the onset of stress. An increase of ABA xylem loading rate as well as AO root activity 4 d and 9 d after application of the treatments was observed only in salt-treated plants followed by a decrease after 14 d in 100 mM NaCl. Decreased cytokinin (trans-zeatin riboside) delivery rate into the xylem sap was observed in all treatments. The role of abscisic acid and cytokinins as positive and negative growth signals, as well as the involvement of root-generated ABA on ABA accumulation in leaves is discussed.     

Complementarity in mineral nitrogen use among dominant plant species in a subalpine community

The underlying mechanisms that enable plant species to coexist are poorly understood. Complementarity in resource use is among the major mechanisms proposed that could favor species coexistence but is insufficiently documented. In alpine soil, low temperatures are a major constraint for the supply of plant nitrogen. We carried out (15)N labeling of soil mineral N to determine to what extent four major species of a subalpine community compete for N, or develop ionic (NH(4)(+) vs. NO(3)(-)) or temporal complementarity. The Poaceae took up much more (15)N per soil area unit than the ericaceous species, and all species displayed three major strategies in exploiting (15)N: (1) uptake mainly early in the growing season (Vaccinium myrtillus), (2) uptake at a slow and similar rate throughout the growing season (Rhododendron ferrugineum), and (3) uptake at high rates over the growing season (Festuca eskia and Nardus stricta). However, while F. eskia used (15)NH(4)(+) mainly early and (15)NO(3)(-) mainly late in the growing season, the reverse was observed for N. stricta. Taking into account (15)N dilution in soil NH(4)(+) and NO(3)(-) pools, we calculated that NH(4)(+) provided more than 80% of the mineral N uptake in Ericaceae and about 60% in grasses. Together, such ionic and temporal complementarity would reduce competition between species and could be a major mechanism promoting species diversity.