Nitrogen uptake and metabolism in Populus x canescens as affected by salinity

External salinization can affect different steps of nitrogen (N) metabolism (ion uptake, N assimilation, and amino acid and protein synthesis) depending on the inorganic N source. Here, we assessed the net uptake of N supplied as nitrate or ammonium and N assimilation (combining metabolite analyses with molecular biological approaches) in grey poplar (Populus x canescens) plants grown under saline (75 mM NaCl) and control conditions. The specific (micromol N g(-1) dry weight fine roots h(-1)) and total plant (micromol N per plant h(-1)) N net uptake rates, total plant N content, total plant biomass and total leaf protein concentration were reduced under saline conditions when plants were supplied with ammonium. In both nutritional groups, salt treatment caused pronounced accumulation of soluble N compounds in the leaves. The mRNAs of genes coding for enzymes catalyzing rate-limiting steps of both proline synthesis and degradation (delta-1-pyrroline-5-carboxylate synthase and proline dehydrogenase) as well as for NADH-dependent glutamate synthase were accumulated under saline conditions. Whereas under control conditions the plant N status seemed to be superior when ammonium was supplied, the N balance of ammonium-fed plants was more severely affected by salt stress than that of plants supplied with nitrate. Possible metabolic implications of stress-related accumulation of particular amino acids are discussed.     

Nitrogen Use Efficiency of Crop Plants: Physiological Constraints upon Nitrogen Absorption

Current global nitrogen fertilizer use has reached approximately one hundred billion kg per annum. In many agricultural systems, a very substantial portion of this applied nitrogen fertilizer is lost from soil to groundwaters, rivers and oceans. While soil physicochemical properties play a significant part in these losses, there are several characteristic features of plant nitrogen transporter function that facilitate N losses. Nitrate and ammonium efflux from roots result in a reduction of net nitrogen uptake. As external nitrate and ammonium concentrations, respectively, are increased, particularly into the range of concentrations that are typical of agricultural soils, elevated rates of nitrate and ammonium efflux result. The rapid down-regulation of high-affinity influx as plants become nitrogen replete further reduces the root's capacity to acquire external nitrogen; only nitrogen-starved roots absorb with both high capacity and high affinity. The results of studies using molecular biology methods demonstrate that genes encoding nitrate and ammonium transporters are rapidly down-regulated when nitrogen is resupplied to nitrogen-starved plants. Provision of ammonium to roots of plants actively absorbing nitrate imposes a block on nitrate uptake, the extent of which depends on the ammonium concentration, thus further reducing the efficient utilization of soil nitrate. During the daily variation of incoming light and during periods of low incident irradiation (i.e. heavy cloud cover) the expression levels of genes encoding nitrate and ammonium transporters, and rates of nitrate and ammonium uptake, are substantially reduced. Low temperatures reduce growth and nitrogen demand, and appear to discriminate against high-affinity nitrogen influx. In sum, these several factors conspire to limit rates of plant nitrogen uptake to values that are well below capacity. These characteristics of the plant's nitrogen uptake systems facilitate nitrogen losses from soils.     

硝态氮对小白菜铬污染毒性的调控作用

通过盆栽和水培试验,探讨了硝态氮对小白菜铬污染毒性的调控作用。结果表明:外源Cr6 对小白菜铬吸收和积累具有明显的促进效应,抑制了小白菜对铁养分的吸收并降低了小白菜的硝酸还原酶活性;硝态氮可有效缓减Cr6 对小白菜吸收铁和硝酸还原酶活性的抑制作用,促进小白菜碳氮代谢和Vc的生物合成,并刺激小白菜生长。在相同铬污染条件下,土壤硝态氮的增加促进了外源Cr6 向有机态转化,且硝态氮有协同强化小白菜吸收Cr6 的效应。表明硝态氮在促进小白菜生长的同时,也促进了小白菜对Cr6 的吸收,提高了小白菜的铬累积水平。     

Diurnal nitrate uptake in young tomato (Lycopersicon esculentum Mill.) plants: test of a feedback-based model

A simple model is proposed to describe diurnal net nitrate uptake rate patterns observed experimentally on young plants grown under constant non-limiting nutrition. It rests on two hypotheses: net uptake rate is under negative feedback control by internal plant nitrate content, and nitrogen metabolism occurs only during the light period. The model parameters were determined from the results of three independent experiments performed under non-disturbing conditions in a growth room at constant air and solution temperatures. Net hourly nitrate uptake rate was measured through a diurnal cycle and after an extended 28 h period of darkness. It increased continuously during the light period and decreased during the dark period. Under prolonged darkness, net uptake declined to an asymptotic positive uptake rate of about 10^-5 mol h^-1 g^-1 total plant dry weight. The measured hourly nitrate uptake rate values were consistent with independent determinations of long-term nitrate and total N accumulations in the plant. Realistic simulations of experimental data are achieved with the proposed model. Furthermore, the maintenance of a positive net uptake rate, measured in non-growing plants subjected to prolonged darkness, is explained in the model by the continuous increase of plant water content. The importance of the diurnal variations of plant water content for nitrate uptake rate is emphasized and gives consistency to the homeostasis hypothesis of the model. The diurnal changes in nitrate uptake predicted by the model are strongly dependent on the assumption made for diurnal changes in nitrate assimilation. While the purely photosynthetic assumption is convenient, a more realistic metabolism sub-model is needed.     

Effects of rhizospheric bicarbonate on net nitrate uptake and partitioning between the main nitrate utilising processes in Populus canescens and Sambucus nigra

Increased levels of rhizospheric dissolved inorganic carbon have repeatedly been demonstrated to enhance plant growth by up to 80%, although carbon from dark fixation accounts for only 1–3% of total plant carbon gain. This study, therefore, aimed at investigating the effects of bicarbonate on nitrate uptake, assimilation and translocation to shoots. Clonal saplings of poplar (Populus canescens(Ait.) Sm.) and elder (Sambucus nigraL.) were grown hydroponically for 35 days in a nutrient solution containing 0, 0.5 and 1 mM bicarbonate and 2 mM nitrate as the sole nitrogen source at pH 7.0. Net nitrate uptake, root nitrate accumulation and reduction, and export of nitrogenous solutes to shoots were measured after incubating plants with ¹⁵N-labelled nitrate for 24 h. Net nitrate uptake increased non-significantly in plant species (19–61% compared to control plants) in response to 1 mM bicarbonate. Root nitrate reduction and nitrogen export to shoots increased by 80 and 95% and 15 and 44% in poplar and elder, respectively. With enhanced root zone bicarbonate, both species also exhibited a marked shift between the main nitrate utilising processes. Poplar plants increasingly utilised nitrate via nitrate reduction (73–88% of net nitrate uptake), whereas the proportions of export (20–9%) and storage in roots (7–3%) declined as plants were exposed to 1 mM external bicarbonate. On the other hand, elder plants exhibited a significant increase of root nitrate reduction (44–66%) and root nitrate accumulation (6–25%). Nitrate translocation to elder shoots decreased from 50 to 8% of net nitrate uptake. The improved supply of nitrogen to shoots did not translate into a significant stimulation of growth, relative growth rates increased by only 16% in poplar saplings and by 7% in elder plants. This revised version was published online in June 2006 with corrections to the Cover Date.     

木霉对盐渍逆境下枸杞根系氮素吸收和氮素利用效率的影响

本研究采集滨海盐渍土开展盆栽试验,分析施加有机肥、木霉菌剂及菌肥对枸杞氮素吸收、同化、积累和利用效率的影响,以揭示木霉对盐渍逆境下枸杞的促生机理。有机肥为木霉菌肥的灭菌物,不含木霉活菌,但两者氮、磷、钾等养分含量无显著差异。结果表明: 施加有机肥、木霉菌剂和菌肥处理较对照均显著提高了根系分生区NO₃^-、NH₄⁺内流速率和成熟区NO₃^-内流速率,且施加菌肥的提升幅度高于施加有机肥。与对照相比,盐渍土壤施加木霉菌剂及菌肥显著增加了根、茎、叶生物量和氮含量以及植株氮累积量,增强了枸杞根和叶中硝酸还原酶、亚硝酸还原酶和谷氨酰胺合成酶活性,提高了枸杞氮素吸收效率、光合速率、稳定碳同位素丰度值和光合氮素利用效率,而且施加菌肥的效果明显优于施加有机肥。综上,木霉能增强盐渍逆境下枸杞氮素吸收、同化和积累,提升光合固碳能力和氮素利用效率,最终促进植株生长。     

Molecular Components of Nitrate and Nitrite Efflux in Yeast

Some eukaryotes, such as plant and fungi, are capable of utilizing nitrate as the sole nitrogen source. Once transported into the cell, nitrate is reduced to ammonium by the consecutive action of nitrate and nitrite reductase. How nitrate assimilation is balanced with nitrate and nitrite efflux is unknown, as are the proteins involved. The nitrate assimilatory yeast Hansenula polymorpha was used as a model to dissect these efflux systems. We identified the sulfite transporters Ssu1 and Ssu2 as effective nitrate exporters, Ssu2 being quantitatively more important, and we characterize the Nar1 protein as a nitrate/nitrite exporter. The use of strains lacking either SSU2 or NAR1 along with the nitrate reductase gene YNR1 showed that nitrate reductase activity is not required for net nitrate uptake. Growth test experiments indicated that Ssu2 and Nar1 exporters allow yeast to cope with nitrite toxicity. We also have shown that the well-known Saccharomyces cerevisiae sulfite efflux permease Ssu1 is also able to excrete nitrite and nitrate. These results characterize for the first time essential components of the nitrate/nitrite efflux system and their impact on net nitrate uptake and its regulation.     

The interaction between elevated carbon dioxide and nitrogen nutrition: the physiological and molecular background

AGPase, ADP glucose pyrophosphorylase
GS, glutamine synthetase
GOGAT, glutamate : oxoglutarate amino transferase
NADP-ICDH, NADP-dependent isocitrate dehydrogenase
NR, nitrate reductase
OPPP, oxidative pentose phosphate pathway
3PGA, glycerate-3-phosphate
PEPCase, phosphoenolpyruvate carboxylase
Rubisco, ribulose-1,5-bisphosphate carboxylase/oxygenase
SPS, sucrose phosphate-synthase

This review first summarizes the numerous studies that have described the interaction between the nitrogen supply and the response of photosynthesis, metabolism and growth to elevated [CO₂]. The initial stimulation of photosynthesis in elevated [CO₂] is often followed by a decline of photosynthesis, that is typically accompanied by a decrease of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), an accumulation of carbohydrate especially starch, and a decrease of the nitrogen concentration in the plant. These changes are particularly marked when the nitrogen supply is low, whereas when the nitrogen supply is adequate there is no acclimation of photosynthesis, no major decrease in the internal concentration of nitrogen or the levels of nitrogen metabolites, and growth is stimulated markedly. Second, emerging evidence is discussed that signals derived from nitrate and nitrogen metabolites such as glutamine act to regulate the expression of genes involved in nitrate and ammonium uptake and assimilation, organic acid synthesis and starch accumulation, to modulate the sugar-mediated repression of the expression of genes involved in photosynthesis, and to modulate whole plant events including shoot–root allocation, root architecture and flowering. Third, increased rates of growth in elevated [CO₂] will require higher rates of inorganic nitrogen uptake and assimilation. Recent evidence is discussed that an increased supply of sugars can increase the rates of nitrate and ammonium uptake and assimilation, the synthesis of organic acid acceptors, and the synthesis of amino acids. Fourth, interpretation of experiments in elevated [CO₂] requires that the nitrogen status of the plants is monitored. The suitability of different criteria to assess the plant nitrogen status is critically discussed. Finally the review returns to experiments with elevated [CO₂] and discusses the following topics: is, and if so how, are nitrate and ammonium uptake and metabolism stimulated in elevated [CO₂], and does the result depend on the nitrogen supply? Is acclimation of photosynthesis the result of sugar-mediated repression of gene expression, end-product feedback of photosynthesis, nitrogen-induced senescence, or ontogenetic drift? Is the accumulation of starch a passive response to increased carbohydrate formation, or is it triggered by changes in the nutrient status? How do changes in sugar production and inorganic nitrogen assimilation interact in different conditions and at different stages of the life history to determine the response of whole plant growth and allocation to elevated [CO₂]?     

The Effects of Irradiance on Uptake and Assimilation of Nitrate by Young Barley Seedlings

The nitrogen economy of barley plants growing in a range ofirradiances from full shade (less than 0·5 W m^–2)to 119 W ^m–2 has been examined by analysing levels oftotal, organic and nitrate nitrogen, and by determining nitratereductase activity in leaf extracts. It has been confirmed thatroot growth is reduced in low irradiances which are also associatedwith a lower level of total nitrogen in the plant, and hencewith a lower uptake of nitrate. In all parts of the plant thelevel of organic nitrogen is higher in high light intensitybut nitrate-nitrogen as a proportion of the total is greatestin low irradiances. In the first leaf accumulation of free nitrateis substantially greater in low irradiances. The data indicate a higher level of nitrate assimilation inhigh irradiances and nitrate reductase activity in leaf extractsis higher in such conditions. When the first leaf is shadednitrate reductase activity falls to undetectable levels afterabout 4 days, but in the case of the second leaf, where thisis shaded, some reductase activity is always found, althoughthis is substantially less than that in unshaded conditions. It is concluded that in vitro rates of nitrate reduction mayover-estimate nitrate assimilation determined as increase inorganic nitrogen.     

Metabolism of carbon and nitrogen by soybean seedlings in response to vegetative apex removal

Short-term (31-hour diurnal) growth-chamber studies were conducted to determine the effects of removing the vegetative apex (meristem and developing trifoliolate leaves) on net photosynthesis (changes in plant dry weight), on distribution of metabolites among plant parts, and on nitrate metabolism and reduced-N accumulation by soybean [Glycine max (L.) Merr.] seedlings. Roots and stems served as alternate sinks for dry matter accumulation in the absence of the vegetative apex. Sugar concentration in roots increased (42%) within 4 hours of vegetative apex removal, and remained higher than for the controls during the 31-hour experimental period. Nitrate assimilation (nitrate reductase activity and total accumulation of reduced-N) was also enhanced in response to vegetative apex removal. Although dry matter accumulation was similar between treated and control plants (113 versus 116 milligrams per plant) over the 31-hour sampling period, more nitrate (1.31 versus 0.79 milligrams per plant) and more reduced-N (3.96 versus 3.45 milligrams per plant) accumulated in treated plants during the same interval. It was concluded that vegetative apex removal had little effect on overall net photosynthesis of soybean seedlings during the 31-hour treatment period, but did alter partitioning of photosynthate and enhanced uptake, transport, and reduction of nitrate. Implications are that uptake and metabolism of nitrate by soybeans may be limited by flux of carbohydrate to the roots, although hormonal effects due to vegetative apex removal cannot be ruled out.     

Plant assimilation and nitrogen cycling

Nitrogen, an abundant and yet limiting nutrient for crop and food production, enters the plant as nitrate or ammonium, or as dinitrogen through biological fixation by procaryotes associated with the plant. Nitrogen incorporation into the soil-plant-animal system is ultimately restricted by rates of biological and industrial fixation. Biological fixation conserves fossil fuel, but fertilization is preferred in most present agriculture. Nitrogen-metabolism research has the practical objectives of allowing more efficient N-fertilizer utilization by plants, including those that fix N₂ but benefit from fertilizer_N supplements. Nitrogen accumulation by harvested crops results in changes in soil acidity, with the direction of change depending on the N-source. There is little information on long-term effects of crop N-nutrition on acidity, and acidity is a critical factor that affects agricultural productivity in many tropical soils. Thus, plant control of pH and the acid/base balance in the soil as a consequence of nitrogen uptake and assimilation are important areas of future research.     

土壤易矿化有机态氮和微生物态氮作为土壤氮素生物有效性指标的评价

通过盆栽试验研究了土壤易矿化有机态氮和土壤微生物态氮与土壤净矿化氮及植物吸氮量之间的关系。结果表明,种植前土壤易矿化有机态氮和土壤微生物态氮以及种植前后土壤易矿化有机态氮的变化量均与土壤氮素净矿化量和植物吸氮量之间存在显著的相关性。在盆栽试验条件下,土壤易矿化有机态氮和种植前土壤微生物态氮能够较好地反映土壤氮素的矿化和供应能力,可以作为土壤氮素的生物有效性指标。     

Interactions between nitrate and ammonium during uptake by carob seedlings and the effect of the form of earlier nitrogen nutrition

Seedlings of carob ( Ceratonia siliqua L. cv. Mulata) were used in two sets of experiments in order to evaluate; (1) the reciprocal effects of each nitrogen form on net uptake of nitrate and ammonium, and (2) the effect of earlier nitrogen nutrition on ammonium versus nitrate uptake. In the former group of experiments we studied the kinetics of nitrate and ammonium uptake as well as the interference of each of the two forms with net uptake of ammonium and nitrate by both nitrogen depleted and nitrogen fed carob seedlings. On the whole, nitrogen depletion led to increase in both affinity and V_max of the system for both forms of nitrogen, at the same time as the effects of nitrate on uptake of ammonium and vice versa were concentration dependent. In the second group of experiments the effects of earlier nitrogen nutrition on nitrate and ammonium uptake were characterized, and in this case we observed that: (a) if only one form of N was supplied, ammonium was taken up in greater amounts than nitrate; (b) the presence of ammonium enhanced nitrate uptake; (c) ammonium uptake was inhibited by nitrate; (d) there was a significant effect of the earlier nitrogen nutrition on the response of the plants to a different nitrogen source. The latter was evident mainly as regards ammonium uptake by plants grown in ammonium nitrate. The interactions between nitrate and ammonium uptake systems are discussed on the basis of the adaptation to the nitrogen source during early growth.     

Accumulation of Oxalate in Tissues of Sugar-beet, and the Effect of Nitrogen Supply

In field-grown sugar-beet concentration of insoluble oxalatewas low in roots and high (about 12 per cent of ethanol insolublematerial) in leaves, and for a particular leaf the concentrationincreased continuously during its life. Of the insoluble oxalate,15–30 per cent was present as the magnesium salt and theremainder as the calcium salt. Oxalate contents of plants grownin culture solutions with nitrate as nitrogen source were similarto those of plants grown in soil, but when nitrogen was suppliedas ammonium sulphate or ammonium nitrate both soluble and insolubleoxalate were low. Plants grown in soil with regular additionsof ammonium sulphate or ammonium nitrate also had very low concentrationsof soluble oxalate although insoluble oxalate was only slightlylower than with nitrate nitrogen. Disks of root or leaf tissuewashed for several days in distilled water lost insoluble oxalatebut when washed in tap water insoluble oxalate increased morethan twofold. Addition of calcium and nitrate to the distilledwater caused an increase of insoluble oxalate, while additionof potassium caused a decrease. Use of ¹⁴C labelled oxalateand washing experiments showed that oxalate can be metabolizedby tissue disks and so is not necessarily a final product ofmetabolism. The accumulation of oxalate appears to be connectedwith the assimilation of nitrate and the preservation of thecation-anion balance of the plant.     

局部根区水分胁迫下氮形态与供给部位对玉米幼苗生长的影响

通过向玉米幼苗分根装置一侧根室的营养液中加入聚乙二醇(PEG 6000)来模拟植物水分胁迫,并设3种供氮形态(硝态氮、铵态氮、两者各占50%的混合氮),且只加入到一侧根室(当氮加入到和PEG同侧时为水氮异区,加入到无PEG一侧时为水氮同区),测定各处理的光合、生理指标,以研究局部根区水分胁迫下氮形态与供给部位对玉米幼苗生长的影响.结果表明:同一氮形态供给下水氮同区植株的光合速率(Pn)、最大净光合速率(Pmax)、光饱和点(LSP)、CO2饱和点(CSP)、叶绿素a、b及叶绿素总含量、根系活力、氮含量和生物量高于水氮异区,光呼吸速率(Rp)、CO2补偿点(CCP)、木质部汁液脱落酸(ABA)浓度、氮利用效率、水分利用效率低于水氮异区;供混合氮和硝态氮的植株Pn、Pmax、LSP、CSP、氮含量和生物量高于供铵态氮的植株,而CCP、Rp、木质部汁液ABA浓度、氮利用效率、水分利用效率变化趋势则相反.可见,同一供氮形态下,水氮同区比水氮异区更利于植物生长,而水氮利用效率在水氮异区下较高;混合氮和硝态氮对植物生长的促进作用优于单一供给铵态氮,但铵态氮更有利于提高水氮利用效率.     

Root zone solute dynamics under drip irrigation: A review

Infiltration and subsequent distribution of water and solutes under cropped conditions is strongly dependent on the irrigation method, soil type, crop root distribution, and uptake patterns and rates of water and solutes. This review discusses aspects of soil water and solute dynamics as affected by the irrigation and fertigation methods, in the presence of active plant uptake of water and solutes. Fertigation with poor quality water can lead to accumulation of salts in the root zone to toxic levels, potentially causing deterioration of soil hydraulic and physical properties. The high frequency of application under drip irrigation enables maintenance of salts at tolerable levels within the rooting zone. Plant roots play a major role in soil water and solute dynamics by modifying the water and solute uptake patterns in the rooting zone. Modeling of root uptake of water and solutes is commonly based on incorporating spatial root distribution and root length or density. Other models attempt to construct root architecture. Corn uptake rate and pattern of nitrate nitrogen was determined from field studies of nitrate dynamics under drip irrigation using TDR monitoring. The determined nitrate nitrogen uptake rates are within literature values for corn. This revised version was published online in June 2006 with corrections to the Cover Date.     

Role of Arbuscular Mycorrhizal Fungi in Uptake of Phosphorus and Nitrogen From Soil

Abstract

Colonization of plant roots by arbuscular mycorrhizal fungi can greatly increase the plant uptake of phosphorus and nitrogen. The most prominent contribution of arbuscular mycorrhizal fungi to plant growth is due to uptake of nutrients by extraradical mycorrhizal hyphae. Quantification of hyphal nutrient uptake has become possible by the use of soil boxes with separated growing zones for roots and hyphae. Many (but not all) tested fungal isolates increased phosphorus and nitrogen uptake of the plant by absorbing phosphate, ammonium, and nitrate from soil. However, compared with the nutrient demand of the plant for growth, the contribution of arbuscular mycorrhizal fungi to plant phosphorus uptake is usually much larger than the contribution to plant nitrogen uptake. The utilization of soil nutrients may depend more on efficient uptake of phosphate, nitrate, and ammonium from the soil solution even at low supply concentrations than on mobilization processes in the hyphosphere. In contrast to ectomycorrhizal fungi, nonsoluble nutrient sources in soil are used only to a limited extent by hyphae of arbuscular mycorrhizal fungi. Side effects of mycorrhizal colonization on, for example, plant health or root activity may also influence plant nutrient uptake.