Factors affecting cation-anion uptake balance and iron acquisition in peanut plants grown on calcareous soils

Dicotyledonous plants subjected to Fe-deficiency stress can decrease pH in the rhizosphere by proton excretion and reduce ferric iron by an activated reduction system in the plasma membranes of the root or by reductants released from the roots. The efficiency by which these plants take up Fe may strongly depend on their cation-anion balance. This study presents results of two experiments conducted to evaluate the effect of K, growth stage and cultivar on ionic balance and Fe acquisition of peanut (Arachis hypogaea L.) plants.Potassium applications to the high calcareous soil (30.3% CaCO₃) favoured proton release, but did not ameliorate plant Fe acquisition. At the earliest stages of plant growth, anion uptake exceeded cation uptake due to intensive N uptake. With time, a shift in the ionic balance was observed as a result of predominant cation uptake. It appears that the relationship between H/OH-ion release and Fe nutrition of peanut plants is actually a complex phenomenon under soil conditions and depends on some soil parameters, such as CaCO₃ content. Even by enhanced H-ion release Fe nutrition of plants can be impaired if soil CaCO₃ is too high.     

Characterization of ammonium and nitrate uptake and assimilation in roots of tea plants

It has been pointed out that tea (Camellia sinensis (L.) O. Kuntze) prefers ammonium (NH ₄ ⁺ ) over nitrate (NO ₃ ^? ) as an inorganic nitrogen (N) source. ¹⁵N studies were conducted using hydroponically grown tea plants to clarify the characteristics of uptake and assimilation of NH ₄ ⁺ and NO ₃ ^? by tea roots. The total ¹⁵N was detected, and kinetic parameters were calculated after feeding ¹⁵NH ₄ ⁺ or ¹⁵NO ₃ ^? to tea plants. The process of N assimilation was studied by monitoring the dynamic ¹⁵N abundance in the free amino acids of tea plant roots by GC-MS. Tea plants supplied with ¹⁵NH ₄ ⁺ absorbed significantly more ¹⁵N than those supplied with ¹⁵NO ₃ ^? . The kinetics of ¹⁵NH ₄ ⁺ and ¹⁵NO ₃ ^? influx into tea plants followed a classic biphasic pattern, demonstrating the action of a high affinity transport system (HATS) and a low affinity transport system (LATS). The V _max value for NH ₄ ⁺ uptake was 54.5 nmol/(g dry wt min), which was higher than that observed for NO ₃ ^? (39.3 nmol/(g dry wt min)). K_M estimates were approximately 0.06 mM for NH ₄ ⁺ and 0.16 mM for NO ₃ ^? , indicating a higher rate of NH ₄ ⁺ absorption by tea plant roots. Tea plants fed with ¹⁵NH ₄ ⁺ accumulated larger amounts of assimilated N, especially glutamine (Gln), compared with those fed with ¹⁵NO ₃ ^? . Gln, Glu, theanine (Thea), Ser, and Asp were the main free amino acids that were labeled with ¹⁵N under both conditions. The rate of N assimilation into Thea in the roots of NO ₃ ^? -supplied tea plants was quicker than in NH ₄ ⁺ -supplied tea plants. NO ₃ ^? uptake by roots, rather than reduction or transport within the plant, seems to be the main factor limiting the growth of tea plants supplied with NO ₃ ^? as the sole N source. The NH ₄ ⁺ absorbed by tea plants directly, as well as that produced by NO ₃ ^? reduction, was assimilated through the glutamine synthetase-glutamine oxoglutarate aminotransferase pathway in tea plant roots. The ¹⁵N labeling experiments showed that there was no direct relationship between the Thea synthesis and the preference of tea plants for NH ₄ ⁺ .     

Relationships between nitrogen uptake and carbon assimilation in whole plants of tall fescue

Abstract. The present study investigates the relationships between nitrogen uptake, transpiration, and carbon assimilation. Plants growing on nutrient solution were enclosed for 10–16 d in a growth chamber, where temperature, photon flux density, vapour saturation deficit and CO₂ concentration were controlled. One of these factors was modified every 4 to 5 d. Shoot photosynthesis and root and shoot respiration were recorded every half-hour. Nitrogen uptake from the root medium and plant transpiration were measured daily. In most cases, an increase in photon flux density led to increases in transpiration, net daily carbon assimilation, and nitrogen uptake. By modifying transpiration rate without changing photosynthesis (varying vapour saturation deficit), or by modifying transpiration and carbon assimilation in opposite ways (varying CO₂ air concentration), it was shown that nitrogen uptake does not follow transpiration, but is linked to the carbon uptake of the plant. When light was increased from low to intermediate levels, the N uptake/C assimilation ratio remained constant. At higher photon flux density, this ratio declined markedly. It is proposed that in the first case, growth is limited by carbohydrate availability, thus any increase in carbon assimilation leads to a proportional increase in nitrogen uptake, in contrast to the second situation where carbohydrates may accumulate in the plant without further nitrogen requirement.     

Effects of aluminium on nitrate uptake and assimilation

A study was conducted to examine the hypothesis that the effects of external Al on NO₃^? uptake and assimilation depend upon the concentration of Al present. Young soybean seedlings [Glycine max (L.) Merrill, cv. Essex], growing under moderate acidity stress at pH 4-2, were exposed to a range of {A1³⁺} in solution for 3d, and to labelled 99 atom %¹⁵NO₃^? during the final hour of Al exposure. Uptake of ¹⁵NO₃^?g^?1 root dry weight was increased by about 28% in the presence of Al at {A1³⁺} below 10 mmolm^?3, and NO₃^? uptake was decreased by about 12% when the {A1³⁺} increased to 44mmoln^?3. The stimulation phase closely paralleled stimulation of root elongation. At higher {A1³⁺}, the inhibition of root elongation was much more severe than that of NO₃^? uptake. There was no indication of a separate effect of Al on root ¹⁵NO₃^? reduction in situ, as the accumulation of reduced ¹⁵N in the root remained a similar percentage of ¹⁵NO₃^? uptake at all {A1³⁺}. At higher {A1³⁺}, the atom %¹⁵N enrichment of the insoluble reduced-N (protein) fraction of root tips increased. This suggests that the Al inhibition of root elongation did not result from disruption of the N supply to the root apex.     

Effects of increasing inorganic carbon supply to roots on net nitrate uptake and assimilation in tomato seedlings

We investigated the influence of an increased inorganic carbon supply in the root medium on NO^?₃ uptake and assimilation in seedlings of Lycopersicon esculentum (L.) Mill. cv. F144. The seedlings were pre-grown for 4 to 7 days with 0 or 100 mM NaCl in hydroponic culture using 0.2 mM NO^?₃ (group A) or 0.2 mM NH⁺₄ (group B) as nitrogen source. The nutrient solution for group A plants was aerated with air or with air containing 4 800 μumol mol^?1 CO₂. Nitrate uptake rate and root and leaf malate contents in these plants were determined. The plants of group B were subdivided into two sets. Plants of one set were transferred either to N-free solution containing 0 or 5 mM NaHCO₃, or to a medium containing 2 mM NO^?₃ and 5 mM NaHCO₃. Both sets of group B plants were grown for 12 h in darkness prior to 2 h of illumination, and were assayed for malate content and NO^?₃ uptake rate (only for plants grown in N-free solution). The second set of group B plants was labeled with ¹⁴C by a 1-h pulse of H¹⁴CO^?₃ which was added to a 5 mM NaHCO₃ solution containing 0 or 100 mM NaCl and 0 or 2 mM NO^?₃, and ¹⁴C-assimilates were extracted and fractionated. The roots of group B plants growing in carbonated medium accumulated twice as much malate as did control plants. This malate was accumulated only when NO^?₃ was absent from the root medium. Both a high level of root malate and aeration with CO₂-enriched air stimulated NO^?₃ uptake. Analysis of ¹⁴C-assimilates indicated that with no NO^?₃ in the medium, the ¹⁴C was present mainly in organic acids, whereas with NO^?₃, a large proportion of ¹⁴C was incorporated into amino acids. Transport of root-incorporated ¹⁴C to the shoot was enhanced by NO^?₃, while the amino acid fraction was the major ¹⁴C-assimilates in the shoot. It is concluded that inorganic carbon fixed through phosphoenolpyruvate carboxylase (EC 4.1.1.31) in roots of tomato plants may have two fates: (a) as a carbon skeleton for amino acid synthesis; and (b) to accumulate, mainly as malate, in the roots, in the absence of a demand for the carbon skeleton. Inorganic carbon fixation in the root provides carbon skeletons for the assimilation of the NH⁺₄ resulting from NO₃ reduction, and the subsequent removal of amino acids through the xylem. This ‘removal’ of NO^?₃ from the cytoplasm of the root cells may in turn increase NO^?₃ uptake.     

Peptide alcohols as promoters of nitrate and ammonium ion uptake in plants

Several aryl-carbamoyl dipeptide alcohols increased the uptake of nitrate and ammonium ions into corn root segments by up to 50% and 90%, respectively. The most effective one was N-carbobenzoxy-l-prolyl-l-valinol. Foliar application of this compound to underfertilized corn plants caused an increase in the rate of plant growth in the greenhouse and provided modest (6-10%) corn yield increases in field tests in Delaware.     

Ionic balance in different tissues of the tomato plant in relation to nitrate, urea, or ammonium nutrition

An investigation was carried out to study the cation-anion balance in different tissues of tomato plants supplied with nitrate, urea, or ammonium nitrogen in water culture.

Irrespective of the form of nutrition, a very close balance was found in the tissues investigated (leaves, petioles, stems, and roots) between total cations (Ca, Mg, K and Na), and total anions (NO₃⁻, H₂PO₄⁻, SO₄⁻⁻, Cl⁻) total non-volatile organic acids, oxalate, and uronic acids. In comparison with the tissues of the nitrate fed plants, the corresponding ammonium tissues contained lower concentrations of inorganic cations, and organic acids and a correspondingly higher proportion of inorganic anions. Tissues from the urea plants were intermediate between the other 2 treatments. These results were independent of concentration or dilution effects, caused by growth. In all tissues approximately equivalent amounts of diffusible cations (Ca⁺⁺, Mg⁺⁺, K⁺ and Na⁺), and diffusible anions (No₃⁻, SO₄⁻⁻, H₂PO₄⁻, Cl⁻) and non-volatile organic acids were found. An almost 1:1 ratio occurred between the levels of bound calcium and magnesium, and oxalate and uronic acids. This points to the fact that in the tomato plant the indiffusible anions are mainly oxalate and pectate. Approximately equivalent values were found for the alkalinity of the ash, and organic anions (total organic acids including oxalate, and uronic acids).

The influence of nitrate, urea, and ammonium nitrogen nutrition on the cation-anion balance and the organic acid content of the plant has been considered and the effects of these different nitrogen forms on both the pH of the plant and the nutrient medium and its consequences discussed.

     

Light and dark assimilation of nitrate in plants

Abstract. Heterotrophic assimilation of nitrate in roots and leaves in darkness is closely linked with the oxidative pentose phosphate pathway. The supply of glucose-6-phosphate to roots and chloroplasts in leaves in darkness is essential for assimilation of nitrite into amino acids. When green leaves are exposed to light, the key enzyme, glucoses-phosphate dehydrogenase, is inhibited by reduction with thioredoxin. Hence the dark nitrate assimilatory pathway is inhibited under photoautotrophic conditions and replaced by regulatory reactions functioning in light. On account of direct photo-synthetic reduction of nitrite in chloroplasts and availability of excess NADH for nitrate reduclase, the rate of nitrate assimilation is extremely rapid in light. Under dark anaerobic conditions also nitrate is equally rapidly reduced to nitrite on account of abolition of competition for NADH between nitrate reductase and mitochondrial oxidation.     

Modulation of nitrate uptake in Anacystis nidulans by the balance between ammonium assimilation and CO2 fixation

Gradual inhibition of ammonium assimilation in Anacystis nidulans cells by increasing concentrations of 5-hydroxylysine resulted in a progressive enhancement of nitrate uptake. For 5-hydroxylysine-treated cells, the magnitude of the inhibition of nitrate uptake promoted by added ammonium was dependent on the ammonium assimilation capacity. In cells with a moderate ammonium assimilation activity, acceleration of CO2 fixation induced by bicarbonate addition antagonized the negative effect of ammonium, allowing full nitrate uptake activity. The results support the contention that nitrate utilization is under the feed-back control exerted by products of its own assimilation via ammonium, the inhibitory effect being potentiated by ammonium addition and alleviated by enhanced CO2 fixation. Results of amino acid analysis in cells exhibiting different capacities to utilize nitrate speak against these compounds as direct effectors of nitrate uptake.     

Electrophysiological factors in the influence of nitrate and ammonium ions on calcium uptake and translocation in tomato plants

Abstract Tomato plants (Lycopersicon esculentum Mill. cv. San Marzano), grown in dilute nutrient solutions containing (in meq ˙ 1^-1) 0.5 NaNO₃, 0.5 NH₄NO₃ or 0.25 (NH₄)₂ SO₄ as the nitrogen source, were detopped for collection of xylem sap and measurement of trans-root electrical potentials. The plant parts and the xylem exudate were subsequently analysed for mineral content. The commonly observed effects of NH₄⁺ were noted, including reduction of calcium concentration in the xylem sap, and of calcium content in stems and leaves, compared with NO₃^-fed plants. This effect was attributed principally to the less negative trans-root electrical potential measured in NH₄⁺-fed plants, and the resultant reduction of inward driving force on passively moving divalent cations.     

Effect of enriched rhizosphere carbon dioxide on nitrate and ammonium uptake in hydroponically grown tomato plants

Previous reports have indicated positive effects of enriched rhizosphere dissolved inorganic carbon on the growth of salinity-stressed tomato (Lycopersicon esculentum L. Mill. cv. F144) plants. In the present work we tested whether a supply of CO₂ enriched air to the roots of hydroponically grown tomato plants had an effect on nitrogen uptake in these plants. Uptake was followed over periods of 6 to 12 hours and measured as the depletion of nitrogen from the nutrient solution aerated with either ambient or CO₂ enriched air. Enriched rhizosphere CO₂ treatments (5000 μmol mol^-1) increased NO₃ ^- uptake up to 30% at pH 5.8 in hydroponically grown tomato plants compared to control treatments aerated with ambient CO₂ (360 μmol mol^-1). Enriched rhizosphere CO₂ treatments had no effect on NH₃ ⁺ uptake. Acetazolamide, an inhibitor of apoplastic carbonic anhydrase, increased NO₃ ^- uptake in ambient rhizosphere CO₂ treatments, but had no effect on NO₃ ^- uptake in enriched rhizosphere CO₂ treatments. Ethoxyzolamide, an inhibitor of both cytoplasmic and extracellular carbonic anhydrase, decreased NO₃ ^- uptake in ambient rhizosphere CO₂ treatments. In contrast, a CO₂ enriched rhizosphere increased NO₃ ^- uptake with ethoxyzolamide, although not to the same extent as in plants without ethoxyzolamide. It is suggested that a supply of enriched CO₂ to the rhizosphere influenced NO₃ ^- uptake through the formation of increased amounts of HCO₃ ^- in the cytosol. This revised version was published online in June 2006 with corrections to the Cover Date.     

Nitrate uptake by roots as regulated by nitrate assimilation in the shoot of castor oil plants

Ricinus communis was used to test the Ben Zioni-Dijkshoorn hypothesis that NO₃ uptake by roots can be regulated by NO₃ assimilation in the shoot. The rate of the anion charge from assimilated NO₃⁻ (and SO₄²⁻) was followed in its distribution between organic acid anion accumulation and HCO₃⁻ efflux into the nutrient solution. In plants adequately supplied with NO₃⁻, HCO₃⁻ efflux accounted for between 56 and 63% of the anion charge. When the plants were subjected to a low NO₃ regime HCO₃⁻ excretion accounted for only 23% of the charge. A comparison of mature plants growing for a 10-day period at the two levels of NO₃ nutrition revealed that the uptake of NO₃⁻ at the higher level was increased 3-fold, whereas K uptake was unaltered. To trace ion movement within the plant, the ionic constituents of xylem and phloem sap were determined. In xylem sap these constituents were found to be predominantly K⁺, Ca²⁺, and NO₃⁻, whereas in the phloem sap they were mainly K⁺ and organic acid anions. Results have been obtained which may be interpreted as providing direct evidence of NO₃ uptake by roots regulated by NO₃ reduction in the tops, the process being facilitated by the recirculation of K⁺ in the plant.     

Effects of cadmium on the uptake,distribution and assimilation of nitrate in Pisum sativum

The net uptake, distribution and assimilation of NO ₃ ^– were studied in pea plants subjected to either long-term continuous Cd treatment for 10 d (10 or 50 M Cd) or short-term treatment (72 h) with 50 M Cd. In the latter treatment, the effects of transferring the plants to a Cd-free nutrient solution for a 'recovery period' of 96 h were also studied. All these treatments were compared with 'controls', plants which received no Cd. In both experiments, the reduction in fresh weight was associated with a decrease in the content (%) of shoot and root water and in transpiration rate as Cd concentration increased. The concentration of ₃ ^– in the shoots and sap decreased dramatically and net ₃ ^– uptake was severely inhibited, effects associated with a loss of shoot nitrate reductase (NR) activity. In the short-term Cd treatment, net ₃ ^– uptake was almost completely inhibited after 24 h, but recovered after the transfer of plants to a Cd-free nutrient solution. Similarly, a dramatic decrease in the shoot NR activity was observed. The uptake, distribution and tissue partitioning of K was also studied, which is considered to be the major counterion of ₃ ^– . Potassium uptake was similarly affected by Cd, as inferred from the ratio ₃ ^– /K uptake, which was ca. 10. The ratio K/ ₃ ^– tissue content increased in the shoot concomitantly to Cd in both long-term and short-term metal supply. These parameters showed a tendency of K similar to that observed for ₃ ^– , although its relative tissue distribution was not affected by Cd.     

Effects of nitrogen dioxide and nitrate nutrition on growth and nitrate assimilation in bean leaves

The influence of nutrient nitrate level (0-20 millimolar) on the effects of NO₂ (0-0.5 parts per million) on growth, K, photosynthetic pigment, N contents, and the activities of enzymes of N assimilation was studied in bean (Phaseolus vulgaris L. cv Kinghorn Wax) leaves. Exposing 7-day old bean seedlings for 5 days continuously to 0.02 to 0.5 parts per million NO₂ increased plant height, fresh weight, chlorophyll, carotenoid, organic N and nitrate contents, and nitrate reductase and glutamate synthase activities in the leaves of seedlings supplied with no external N. At 20 millimolar nitrate, most of the parameters examined were inhibited except for organic N and nitrate contents and glutamate synthase activity which increased in most cases. Generally, with an increase in NO₂ concentration, the stimulatory effect declined and/or the inhibitory effect increased. A 3-hour exposure of 12-day-old bean seedlings to 0.1 to 2.0 parts per million NO₂ increased nitrate content and nitrate reductase activity at each nutrient nitrate level except for a slight inhibition of enzyme activity during exposure to 2.0 parts per million NO₂ at 20 millimolar nitrate. The experiments demonstrated that the effect of NO₂ is strongly influenced by nutrient N level and that NO₂ is assimilated into organic nitrogenous compounds to serve as a source of N, only to a limited extent.     

Effect of nodulation on the nitrate assimilation in vegetative soybean plants

Summary Nitrate assimilation in the first trifoliate leaf of vegetative soybean plants (Glycine max L. Merr, cv Hodgson) was studied in relation to nodulation. Nodulated and non-nodulated plants were grown in a nitrate medium (4 mM). As a control nodulated plants were grown in a nutrient medium without combined nitrogen. This study included measurements of the acetylene reduction activity of the whole plant and of thein vitro nitrate reductase, glutamine synthetase and glutamate dehydrogenase activities in the first leaf and of the nitrate concentration. Nitrate accumulation and nitrate reductase activity were depressed in nodulated plants; root growth was decreased in the presence of nitrate. The relationships between nitrate assimilation and nodulation are discussed.     

Selenium uptake,translocation, assimilation and metabolic fate in plants

The chemical and physical resemblance between selenium (Se) and sulfur (S) establishes that both these elements share common metabolic pathways in plants. The presence of isologous Se and S compounds indicates that these elements compete in biochemical processes that affect uptake, translocation and assimilation throughout plant development. Yet, minor but crucial differences in reactivity and other metabolic interactions infer that some biochemical processes involving Se may be excluded from those relating to S. This review examines the current understanding of physiological and biochemical relationships between S and Se metabolism by highlighting their similarities and differences in relation to uptake, transport and assimilation pathways as observed in Se hyperaccumulator and non-accumulator plant species. The exploitation of genetic resources used in bioengineering strategies of plants is illuminating the function of sulfate transporters and key enzymes of the S assimilatory pathway in relation to Se accumulation and final metabolic fate. These strategies are providing the basic framework by which to resolve questions relating to the essentiality of Se in plants and the mechanisms utilized by Se hyperaccumulators to circumvent toxicity. In addition, such approaches may assist in the future application of genetically engineered Se accumulating plants for environmental renewal and human health objectives.     

Active ion uptake and maintenance of cation-anion balance: A critical examination of their role in regulating rhizosphere pH

The processes responsible for maintenance of cation-anion balance in plants and their relation to active ion accumulation and changes in rhizosphere pH are outlined and discussed. The major processes involved are: (1) accumulation and degradation of organic acids which occur in the plant mainly as organic acid anions (and their transfer within the plant) and (2) extrusion of H⁺ or OH^– into the rhizosphere. The relative importance of the two processes is determined by the size of the excess anion or cation uptake. Indeed, plants typically absorb unequal quantities of nutritive cations (NH₄ ⁺+Ca²⁺+ Mg²⁺+K⁺+Na⁺) and anions (NO₃ ^–+Cl^–+SO₄ ^2–+H₂PO₄ ^–) and charge balance is maintained by excretion of an amount of H⁺ or OH^– which is stoichiometrically equal to the respective excess cation or anion uptake. The mechanisms and processes by which H⁺ and in particular OH^– ions are excreted in response to unequal cation-anion uptake are, however, poorly understood.The contemporary view is that primary active extrusion of H⁺, catalyzed by a membrane-located ATPase, is the major driving force for secondary transport of cations and anions across the plasma membrane. However, the fact that net OH^– extrusion often occurs (since excess anion absorption commonly takes place) implies there is a yet-to-be characterized OH^– ion efflux mechanism at the plasma membrane that is associated with anion uptake. There is, therefore, a need for future studies of the uptake mechanisms and stoichiometry of anion uptake; particularly that of NO₃ ^– which is often the predominant anion absorbed. Another related phenonenon which requires detailed study in terms of cation-anion balance is localized rhizosphere acidification which can occur in response to deficiencies of Fe and P.