Nitrogen utilization by plant species from acid heathland soils: I. Comparison between nitrate and ammonium nutrition at constant low pH

Seven heathland species, four herbaceous plants and three dwarfshrubs, were tested for their capacity to utilize NH₄⁺ or NO₃^–. When cultured in solution at pH 4.0 with 2mol m^–3 N,all species showed similar growth responses with respect toN source. Nitrate was assimilated almost equally well as ammonium,with relative growth rate generally averaging 5–8% lowerfor NO₃^– grown plants, albeit not always significantly.However, N source was significantly and consistently correlatedwith biomass partitioning, as NH₄⁺-fed plants allocated moredry matter to shoots and less to roots when compared to NO₃^–-fed plants. The strong difference in biomass partitioning mayrelate to the relative surplus of carbon per unit plant N (or,alternatively, the relatively suboptimal rate of N assimilationper unit plantC) in NO₃^–-fed plants Inherently slow-growing dwarf shrubs accumulated virtually nofree nitrate in their tissues and reduction of nitrate was strictlyroot-based. Faster-growing herbaceous plants, however, partitionedthe assimilation of nitrate over both shoots and roots, therebyaccumulating relatively high tissue NO₃^– levels. Ion uptakerates depended clearly on the ‘relative shoot demand’.At similar shoot demands, especially in the herbaceous species,specific uptake rates for N and total inorganic (non-N) anionswere higher in NH₄⁺ -fed plants, whereas the uptake rate fortotal (non-N) cations was higher in NO₃^–-fed plants. Rateof P uptake was enhanced with increasing plant demand, but wasindependent of the N source. Net H⁺ extrusions ranged from 1.00to 1.34 H⁺ per NH₄⁺, and from –0.48 to –0.77 H⁺per NO₃^– taken up. Key words: Ammonium, biomass partitioning, heathland plants, low pH, nitrate, nitrate reductase activity, relative shoot demand, specific absorption rate     

Ammonium and nitrate uptake by soybean during recovery from nitrogen deprivation

Soybean [Glycine max (L.) Merrill] plants that had been subjectedto 15 d of nitrogen deprivation were resupplied for 10 d with1.0 mol m^–3 nitrogen provided as NO₃^–, NH₄⁺, orNH₄⁺+NO₃^– in flowing hydroponic culture. Plants in a fourthhydroponic system received 1.0 mol m^–3 NO₃^– duringboth stress and resupply periods. Concentrations of solublecarbohydrates and organic acids in roots increased 210 and 370%,respectively, during stress. For the first day of resupply,however, specific uptake rates of nitrogen, determined by ionchromatography as depletion from solution, were lower for stressedthan for non-stressed plants by 43% for NO₃^- resupply, by 32%for NH₄⁺ + NO₃^– resupply, and 86% for NH₄⁺ resupply. Whenspecific uptake of nitrogen for stressed plants recovered torates for non-stressed plants at 6 to 8 d after nitrogen resupply,carbohydrates and organic acids in their roots had declinedto concentrations lower than those of non-stressed plants. Recoveryof nitrogen uptake capacity of roots thus does not appear tobe regulated simply by the content of soluble carbon compoundswithin roots. Solution concentrations of NH₄⁺ and NO₃^– were monitoredat 62.5 min intervals during the first 3 d of resupply. Intermittent‘hourly’ intervals of net influx and net effluxoccurred. Rates of uptake during influx intervals were greaterfor the NH₄⁺ -resupplied than for the NO₃^– -resuppliedplants. For NH₄⁺ -resupplied plants, however, the hourly intervalsof efflux were more numerous than for NO₃^– -resuppliedplants. It thus is possible that, instead of repressing NH₄⁺influx, increased accumulation of amino acids and NH₄⁺ in NH₄⁺-resupplled plants inhibited net uptake by stimulation of effluxof NH₄⁺ absorbed in excess of availability of carbon skeletonsfor assimilation. Entry of NH₄⁺ into root cytoplasm appearedto be less restricted than translocation of amino acids fromthe cytoplasm into the xylem. Key words: Ammonium, nitrate, nitrogen-nutrition, nitrogen-stress, soybean     

Influence of nitrate on uptake of ammonium by nitrogen-depleted soybean: is the effect located in roots or shoots?

In non-nodulated soybean [Glycine max (L.) Merrill cv. Ransom]plants that were subjected to 15 d of nitrogen deprivation inflowing hydroponic culture, concentrations of nitrogen declinedto 1.0 and 1.4mmol Ng^–1 dry weight in shoots and roots,respectively, and the concentration of soluble amino acids (determinedas primary amines) declined to 40µmol g^–1 dry weightin both shoots and roots. In one experiment, nitrogen was resuppliedfor 10 d to one set of nitrogen-depleted plants as 1.0 mol m^–3NH₄⁺ to the whole root system, to a second set as 0.5 mol m^–3NH₄⁺ plus 0.5 mol m^–3 NO₃^– to the whole root system,and to a third set as 1.0 mol m^–3 NH₄⁺ to one-half ofa split-root system and 1.0 mol m^–3 NO₃^– to theother half. In a second experiment, 1.0 mol m^–3 of nitrogenwas resupplied for 4 d to whole root systems in NH₄⁺ : NO₃^–ratios of 1:0, 9:1, and 1:1. Nutrient solutions were maintainedat pH 6.0. When NH₄⁺ was resupplied in combination with NO₃^– to thewhole root system in Experiment I, cumulative uptake of NH₄⁺for the 10 d of resupply was about twice as great as when NH₄⁺was resupplied alone. Also, about twice as much NH₄⁺ as NO₃^–was taken up when both ions were resupplied to the whole rootsystem. When NH₄⁺ and NO₃^– were resupplied to separatehalves of a split-root system, however, cumulative uptake ofNH₄⁺ was about half that of NO₃^–. The uptake of NH₄⁺,which is inhibited in nitrogen-depleted plants, thus is facilitatedby the presence of exogenous NO₃^–, and the stimulatingeffect of NO₃^– on uptake of NH₄⁺ appears to be confinedto processes within root tissues. In Experiment II, resupplyof nitrogen as both NH₄⁺ and NO₃^– in a ratio of either1:1 or 9:1 enhanced the uptake of NH₄⁺. The enhancement of NH₄⁺uptake was 1.8-fold greater when the NH₄⁺: NO₃^–-resupplyratio was 1:1 than when it was 9:1; however, only 1.3 timesas much NO₃^– was taken up by plants resupplied with the1 :1 exogenous ratio. The effect of NO₃^– on enhancementof uptake of NH₄⁺ apparently involves more than net uptake ofNO₃^– itself and perhaps entails an effect of NO₃^–uptake on maintenance of K⁺ availability within the plant. Theconcentration of K⁺ in plants declined slightly during nitrogendeprivation and continued to decline following resupply of nitrogen.The greatest decline in K⁺ concentration occurred when nitrogenwas resupplied as NH₄⁺ alone. It is proposed that decreasedavailability of K⁺ within the NH₄⁺-resup-plied plants inhibitedNH₄⁺ uptake through restricted transfer of amino acids fromthe root symplasm into the xylem. Key words: Ammonium, Glycine max, nitrate, nitrogen-nutrition, nitrogen stress, split-root cultures     

The Influence of NO-3 and NH+4 Nutrition on the Gas Exchange Characteristics of the Roots of Wheat (Triticum aestivum) and Maize (Zea mays) Plants

Respiratory oxygen consumption by roots was 1·4- and1·6-fold larger in NH⁺₄-fed than in NO^-₃-fed wheat (Triticumaestivum L.) and maize (Zea mays L.) plants respectively. Higherroot oxygen consumption in NH⁺₄-fed plants than in NO^-₃-fedplants was associated with higher total nitrogen contents inNH⁺4-fed plants. Root oxygen consumption was, however, not correlatedwith growth rates or shoot:root ratios. Carbon dioxide releasewas 1·4- and 1·2-fold larger in NO⁺3-fed thanin NH⁺₄-fed wheat and maize plants respectively. Differencesin oxygen and carbon dioxide gas exchange rates resulted inthe gas exchange quotients of NH^-₄-fed plants (wheat, 0·5;maize, 0·6) being greatly reduced compared with thoseof NO^-₃-fed plants (wheat, 1·0; maize, 1·1). Measuredrates of HCO^-₃ assimilation by PEPc in roots were considerablylarger in 4 mM NH⁺₄-fed than in 4 NO^-₃ plants (wheat, 2·6-fold;maize, 8·3-fold). These differences were, however, insufficientto account for the observed differences in root carbon dioxideflux and it is probable that HCO^-₃ uptake is also importantin determining carbon dioxide fluxes. Thus reduced root extension in NH⁺₄-fed compared with NO^-₃-fedwheat plants could not be ascribed to differences in carbondioxide losses from roots.Copyright 1993, 1999 Academic Press Triticum aestivum, wheat, Zea mays, maize assimilation, ammonium assimilation, root respiration     

Growth of Actinorhizal Plants as Influenced by the Form of N with Special Reference to Myrica gale and Alnus incana

Growth of two actinorhizal species was studied in relation tothe form of N supply in water culture. Non-nodulated bog myrtle(Myrica gale) and grey alder (Alnus incana) were grown withNH₄⁺, NH₄NO₃^– or NO₃^– (4 mol m^–3 N). A nodulatedseries of bog myrtle was also cultivated in N-free medium. Relative growth rate (RGR), utilization rate of N, and shoot/rootratio were highest for the two species with the N completelysupplied as NH₄⁺. In both species, nitrate was largely reducedin the roots and the presence of NO₃^– in combined-N supplyalways affected the RGR and N utilization rate negatively. BothN₂ fixation and complete NO₃^– nutrition represented conditionsof relative N-deficiency resulting in relatively low tissue-Nconcentrations and a greater allocation of dry mass to the roots.The physiological N status of nodulated M. gale plants was comparativelygood, as indicated by a normal nodule weight ratio and a relativelyhigh N₂-fixing rate per unit nodule mass. However, whole-plantN₂-fixing capacity remained relatively low in comparison withacquisition rates of N in combined-N plants. The anion charge from the nitrate reduction was largely directlyexcreted as an OH^– efflux. H ⁺ /N ratios generally agreedwith the theory. In comparison with NH₄⁺ nutrition, carboxylateconcentrations were higher in N₂-fixing M. gale plants and theH ⁺ /N ratio in nodulated plants was less than unity below thevalue for ammonium plants as previously found for other actinorhizalspecies. Therefore, NH₄⁺ should be an energetically more attractiveN source for actinorhizal plants than N₂. The results agree with commonly accepted views on energeticsof N uptake and assimilation in higher plants and support theconcept of a basically similar physiological behaviour betweennon-legumes and legumes. Key words: Actinorhizal symbioses, ammonium, H⁺/OH^– efflux, nitrate, N₂ fixation, NRA     

Concurrent Rates of N2 Fixation, Nitrate and Ammonium Uptake by White Clover in Response to Growth at Different Root Temperatures

Nodulated white clover plants (Trifolium repens L. cv. Huia)were grown for 71 d in flowing nutrient solutions containingN as 10 mmol m_–3 NH₄NO₃, under artificial illumination,with shoots at 20/15°C day/night temperatures and root temperaturereduced decrementally from 20 to 5°C. Root temperatureswere then changed to 3, 7, 9, 11, 13, 17 or 25°C, and theacquisition of N by N₂ fixation, NH₄+ and NO₃^– uptakewas measured over 14 d. Shoot specific growth rates (d. wt)doubled with increasing temperature between 7 and 17°C,whilst root specific growth rates showed little response; shoot:root ratios increased with root temperature, and over time at11°C. Net uptake of total N per plant (N₂ fixation + NH₄++ NO₃^–) over 14 d increased three-fold between 3 and 17°C.The proportion contributed by N₂ fixation decreased with increasingtemperature from 51% at 5°C to 18% at 25°C. Uptake ofNH₄+ as a proportion of NH₄+ + NO₃^– uptake over 14 d variedlittle (55–62%) with root temperature between 3 and 25°C,although it increased with time at most temperatures. Mean ratesof total N uptake per unit shoot f. wt over 14 d changed littlebetween 9 and 25°C, but decreased progressively with temperaturebelow 9°C, due to the decline in the rates of NH₄+ and NO₃^–uptake, even though N₂ fixation increased. The results suggestthat N₂ fixation in the presence of sustained low concentrationsof NH₄+ and NO₄^– is less sensitive to low root temperaturethan are either NH₄+ or NO₃^– uptake systems. White clover, Trifolium repens L. cv. Huia, root temperature, nitrogen fixation, ammonium, nitrate     

Inhibition of Nitrate Assimilation in Roots in the Presence of Ammonium: The Moderating Influence of Potassium

Experiments were conducted to investigate the effect of concentrationof NH₄⁺ in nutrient solution on root assimilation of NO₃^–and to determine whether the NH₄⁺NO₃^– interaction wasmodified in the presence of K⁺. Dark-grown, detopped corn seedlings(cv. Pioneer 3369A) were exposed for 8 h to 0.15 mM Ca(NO₃)₂and varying concentrations of (NH₄)₂SO₄ in the absence or presenceof 0.15 mM K₂SO₄. The accelerated phase of NO₃^– uptakeappeared most sensitive to restriction by additions of 0.15mM (NH₄)₂SO₄. In the absence of K⁺, the restriction increasedonly slightly even when solution (NH₄)₂SO₄, was increased from0.15 mM to 12.5 mM which was accompanied by an increase of NH₄⁺in the tissue from about 7.0 to 35 µmol g^–1 fr.wt. of root. Increasing concentrations of solution NH₄⁺ progressivelyinhibited net K+ uptake. At the highest solution NH₄⁺ concentrations,there was an initial net efflux of K⁺ and no net influx occurredduring the treatment period. The severity of the NH₄)SO₄ restrictionof NO₃^– uptake was moderated considerably in the presenceof K⁺ as long as a net influx of K⁺ occurred. However, net influxof K⁺ was not associated with alteration of NH₄⁺ uptake, assimilation,or accumulation in the root tissue. The lack of correlationbetween the severity of restriction of NO₃^– uptake andendogenous NHJ suggested the restriction resulted from an effectexerted by exogenous NH₄⁺ which tended to saturate at lowersolution NHJ concentrations or by inhibitory factors generatedduring assimilation of NH₄⁺. Several mechanisms were postulatedto account for the moderating influence of K⁺. In all experiments,root NO₃^– reduction was restricted by the presence ofambient NH₄⁺. The quantitative decreases in reduction tendedto be less than decreases in NO₃^– uptake and therefore,could result from inhibition solely of uptake with subsequentlimitation in availability of substrate for the reduction process,but the possibility of a direct effect on reduction could notbe excluded.     

Ammonium Nutrition in Ricinus communis: Its Effect on Plant Growth and the Chemical Composition of the Whole Plant, Xylem and Phloem Saps

Allen, S. and Smith, J A. C. 1986. Ammonium nutrition in Ricinuscommunis: its effect on plantgrowth and the chemical compositionof the whole plant, xylem and phloem saps.—J. exp. Bot.37: 1599–1610. The growth and chemical composition of Ricinus communis cultivatedhydroponically on 12 mol m ^{– 3} NO₃^–-N were comparedwith plants raised on a range of NH₄⁺-N concentrations. At NH₄⁺-Nconcentrations between 0·5 and 4·0 mol m^–3,fresh- and dry-weight yields of 62-d-old plants were not significantlydifferent from those of the NO₃^–-N controls. Growth wasreduced at 0·2 mol m^–3 NH₄⁺-N and was associatedwith increased root. shoot and C: organic N ratios, suggestingthat the plants were N-limited. At 8·0 mol m^–3NH₄⁺-N, growth was greatly restricted and the plants exhibitedsymptoms of severe ‘NH₄⁺ toxicity’. Plants growingon NH₄⁺-N showed marked acidification of the rooting medium,this effect being greatest on media supporting the highest growthrates. Shoot carboxylate content per unit dry weight was lower at mostNH₄⁺-N concentrations than in the NO₃^–-N controls, althoughit increased at the lowest NH₄⁺-N levels. Root carboxylate contentwas comparable on the two N sources, but also increased substantiallyat the lowest NH₄⁺-N levels. N source had little effect on inorganic-cationcontent at the whole-plant level, while NO₃^– and carboxylatewere replaced by Cl as the dominant anion in the NH₄⁺-N plants.This was reflected in the ionic composition of the xylem andleaf-cell saps, the latter containing about 100 mol m^–3Cl^– in plants on 8·0 mol m^–3 NH₄⁺. Xylem-saporganic-N concentration increased more than threefold with NH₄⁺-N(with glutamine being the dominant compound irrespective ofN source) while in leaf-cell sap it increased more than 12-foldon NH₄⁺-N media (with arginine becoming the dominant species).In the phloem, N source had little or no effect on inorganic-cation,sucrose or organic-N concentrations or sap pH, but sap fromNH₄⁺-N plants contained high levels of Cl^– and serine. Collectively, the results suggested that the toxic effects ofhigh NH₄⁺ concentrations were not the result of medium acidification,reduced inorganic-cation or carboxylate levels, or restrictedcarbohydrate availability, as is commonly supposed. Rather,NH₄⁺ toxicity in R. communis is probably the result of changesin protein N turnover and impairment of the photorespiratoryN cycle. Key words: Ricinus, ammonium nutrition, nitrate, whole-plant composition, xylem, Phloem, amino acids, carboxylate     

The Effects of Temperature and Form of Nitrogen Supply on the Relative Contribution of Root Uptake and Remobilization in Supplying Nitrogen for Laminae Regrowth of Lolium perenne L.

Plants of Lolium perenne L. cv. S23 were grown in sand culturesupplied with either ammonium (NH₄⁺) or nitrate (NO₃^–)in an otherwise complete nutrient solution at 12°C or 20°C.Three weeks after germination, plants were clipped weekly tosimulate grazing. After 10 weeks growth all nitrogen (N) wassupplied enriched with ¹⁵N to quantify the effects of form ofN supply and temperature on the relative ability of currentroot uptake and remobilization to supply N for laminae regrowth. The form of N supply had no effect on the dry matter partitioning,while at 20°C more dry weight was allocated to laminae regrowthand less to the remaining plant material. The current root uptakeof N, which subsequently appeared in the laminae regrowth, wassimilar for plants supplied with NH₄⁺ or NO₃^–, and bothwere equally reduced at the lower temperature of growth. Remobilizationof N to laminae regrowth was greater for plants receiving NH₄⁺than NO₃^–; remobilization with either form of N supplywas reduced at the lower temperature of growth. Remobilizationwas reduced to a lesser extent at 12°C than current rootuptake. It was concluded that remobilization became relativelymore important in supplying N for regrowth of laminae at lowertemperatures. Key words: Lolium perenne, ammonium, nitrate, temperature, remobilization     

Interactions of NH4+ and L-glutamate with NO3- transport processes of non-mycorrhizal Fagus sylvatica roots

The processes of NO₃^– uptake and transport and the effectsof NH₄⁺ or L-glutamate on these processes were investigatedwith excised non-mycorrhizal beech (Fagus sylvatica L.) roots.NO₃^– net uptake followed uniphasic Michaelis-Menten kineticsin a concentration range of 10µM to 1 mM with an apparentK_m of 9.2 µM and a V_max of 366 nmol g^–1 FW h^–1.NH₄⁺, when present in excess to NO₃^–, or 10 mM L-glutamateinhibited the net uptake of NO₃^– Apparently, part of NO₃^–taken up was loaded into the xylem. Relative xylem loading ofNO₃^– ranged from 3.21.6 to 6.45.1% of NO₃^– netuptake. It was not affected by treatment with NH₄⁺ or L-glutamate.¹⁶N/¹³N double labelling experiments showed that NO₃^–efflux from roots increased with increasing influx of NO₃^–and, therefore, declined if influx was reduced by NH₄⁺ or L-glutamateexposure. From these results it is concluded that NO₃^–net uptake by non-mycorrhizal beech roots is reduced by NH₄⁺or L-glutamate at the level of influx and not at the level ofefflux. Key words: Nitrate transport, net uptake, influx, efflux, ammonium, Fagus, Fagaceae     

Metabolism of Glycollate by Lemna minor L. Grown on Nitrate or Ammonium as Nitrogen Source

Marques, I. A., Oberholzer, M. J. and Erismann, K. H. 1985.Metabolism of glycollate by Lemna minor L. grown on nitrateor ammonium as nitrogen source.—J. exp. Bot. 36: 1685–1697. Duckweed, Lemna minor L., grown on inorganic nutrient solutionscontaining either NH₄⁺ or NO₃^– as nitrogen source wasallowed to assimilate [1-¹⁴C]- or [2-¹⁴C]glycollate during a20 min period in darkness or in light. The incorporation ofradioactivity into water-soluble metabolites, the insolublefraction, and into the CO₂ released was measured. In additionthe extractable activity of phosphoenolpyruvate carboxylasewas determined. During the metabolism of [2-¹⁴C]glycollate in darkness, as wellas in the light, NH₄⁺ grown plants evolved more ¹⁴CO₂ than NO₃^–grown plants. Formate was labelled only from [2-¹⁴C]glycollateand in NH₄⁺ grown plants it was significantly less labelledin light than in darkness. In NO₃^– grown plants formateshowed similar radioactivity after dark and light labelling.The radioactivity in glycine was little influenced by the nitrogensource. Amounts of radioactivity in serine implied that thefurther metabolism of serine was reduced in darkness comparedwith its metabolism in the light under both nitrogen regimes.In illuminated NH₄⁺ plants, serine was labelled through a pathwaystarting from phosphoglycerate. After [1-¹⁴C]glycollate feedingNH₄⁺ grown plants contained markedly more radioactive aspartateand malate than NO₃^– plants indicating a stimulated phosphoenolpyruvatecarboxylation in plants grown on NH₄⁺. Key words: Photorespiration, glycollate, nitrogen, Lemna     

Cortical Cell Fluxes of Ammonium and Nitrate in Excised Root Segments ofAllium cepa L; Studies using15N

From compartmental analysis of ¹⁵N elution measurements, concentrationsand fluxes of NH⁺₄ and NO₃^– were estimated for corticalcells in excised root segments, when bathed in a complete nutrientsolution, in which 20 mol m^–3 NH₄⁺ or NO₃^– werethe single N sources. The results were compared with those fornutrient solution containing 20 mol m^–3 NH₄NO₃. No nitratereductase activity was detected in the roots but rapid assimilationof NH₄⁺ occurred, due to glutamine synthetase activity. Theefflux curves for NH₄⁺, on a 'µg ¹⁵N remaining againsttime' basis, deviated from the criteria determining conformityto first order kinetics, since the slowest rate constant wasan order of magnitude lower than that exhibited by the curvefor efflux versus time. The data were transformed to conformto the appropriate criteria, revealing a large slowly exchangingpool equated with assimilated NH₄⁺. The presence of NO₃^–had little effect on NH₄⁺ uptake and exchange, but NH₄⁺ suppressedNOj uptake and reduced exchange across plasmalemma and tonoplast.It was established that NH₄⁺ absorption was an active process.However, NH₄⁺ entering and leaving the vacuole was overestimated,since the flux equation used did not differentiate between total¹⁵NH₄ influx at the plasmalemma and that at the tonoplast, afterassimilation. The only active NO₃^– transfer was influxat the plasmalemma. The results were compared with those ofothers using¹³N and ³⁶C1O₃ analogues to measure NH₄⁺ and NO₃^–fluxes in cereal roots. Key words: Ammonium, nitrate, compartmental analysis, ¹⁵N, active transport     

Influx and Efflux of Nitrate and Ammonium in Italian Ryegrass and White Clover Roots: Comparisons Between Effects of Darkness and Defoliation

Seedlings of Italian ryegrass (Lolium multiflorum Lam. cv. RVP)and clonal stolon cuttings of white clover (Trifolium repensL. cv. Blanca) were grown for 19 d in flowing solution culture,with N supplied as either 250 mmol m^–3 NO₃^– or NH₃⁺.Rates of net uptake, influx and translocation of NO₃^–and NH₄⁺ were then determined using ¹⁵N and ¹³N labelling techniques:between 3–5 h into the photoperiod following 8 h darknessfor white clover (CL), and for ryegrass plants that were eitherentire (IL) or with shoots excised 90 min prior to ¹³N influx(IC); and 75 min into the photoperiod following 37–39h darkness for ryegrass (ID). Rates of net uptake, influx andefflux of NH₄⁺ exceeded those of NO₃^– in IL and IC ryegrassplants: the opposite occurred in white clover (CL). The decreasein net uptake following defoliation of ryegrass was greaterfor NH₄⁺ (62%) than NO₃^– (40%). For NH₄⁺ this was associatedwith a large decrease in influx from 110 to 6.0µmol h^–1g^–1 root fr. wt; but for NO₃^–, influx only decreasedfrom 42 to 37 µmol h^–1 g^–1. Prolonged exposureto darkness (ID plants) also lowered net uptake of NO₃^–and NH₄⁺ by, respectively, 86% and 95% of IL levels. For NH₄⁺this was characterized by a large decrease in influx and a smalldecrease in efflux; whilst for NO₃^– the effect of a largedecrease in influx was reinforced by a smaller increase in efflux. The data were used to estimate the translocatory fluxes of NO₃^–(03–20µmol h^–1 g^–1) and NH₄⁺ (003–0.4µmolh^–1 g^–1), assimilation in the roots of NO₃^–(02–26µmol h^–1 g^–1) and NH⁺⁴ (05–89 µmolh^–1 g^–1), and the concentrations of NO₃^– (9–15mol m^–3) in the cytoplasmic compartment of the roots.The relevance of variable influx and efflux to models for theregulation of N uptake is discussed. Key words: Lolium multiflorum, Trifolium repens, influx, efflux, nitrate, ammonium, ¹³N     

Promotion of Assimilation of Ammonium Ions by Simultaneous Application of Nitrate and Ammonium Ions in Radish Plants

When radish plants were grown in nutrient solutions that containedammonium ions (NH₄⁺) as the sole source of nitrogen, they grewpoorly and accumulated high levels of NH₄⁺ in their leaves.However, radish plants cultured in 5 mM NH₄⁺ plus 1 mM NO₃^–(a ratio of 5 : 1 in forms of nitrogen; referred to as 5:lmix-N)grew well and accumulated very low levels of NH₄⁺ in their leaves.After radish plants were cultured in solutions that containedNO₃^–, or NH₄⁺, or 5: lmix-N for a week, they were thensupplied with the same nitrogen source labeled with ¹⁵N forone day. The uptake of ¹⁵N from labeled NH₄⁺ into total nitrogenwas the highest in plants supplied with 5:1mix-N. These plantsconverted far fewer labeled NH₄⁺ into free NH₄⁺ than did NH₄⁺-fedplants, but converted many more labeled NH₄⁺ into the insolublefraction than did NH₄⁺- or NO₃^–-fed plants. The presence of a small amount of nitrate was shown to stimulatethe assimilation of ammonium ions and the synthesis of proteins. (Received October 26, 1988; Accepted January 24, 1989)     

Nitrogen utilization by plant species from acid heathland soils: II. Growth and shoot/root partitioning of NO3- assimilation at constant low pH and varying NO3-/NH4+ ratio

The growth of four heathland species, two grasses (D. flexuosa,M. caerulea) and two dwarf shrubs (C. vulgaris, E. tetralix),was tested in solution culture at pH 4.0 with 2 mol m^–3N, varying the N0₃^–/NH₄⁺ ratio up to 40% nitrate. In addition,measurements of NRA, plant chemical composition, and biomassallocation were carried out on a complete N0₃^–/NH₄⁺ replacementseries up to 100% nitrate. With the exception of M. caerulea, the partial replacement ofNH₄⁺ by NO₃^– tended to enhance the plant's growth ratewhen compared to NH₄⁺ only. In contrast to the other species,D. flexuosa showed a very flexible response in biomass allocation:a gradual increase in the root weight ratio (RWR) with NO₃^–increasing from 0 to 100%. In the presence of NH₄⁺, grassesreduced nitrate in the shoot only; roots did not become involvedin the reduction of nitrate until zero ambient NH₄⁺. The dwarfshrubs, being species that assimilate N exclusively in theirroots, displayed an enhanced root NRA in the presence of nitrate;in contrast to the steady increase with increasing NO₃^–in Calluna roots, enzyme activity in Erica roots followed arather irregular pattern. Free nitrate accumulated in the tissuesof grasses only, and particularly in D. flexuosa. The relative uptake ratio for NO₃^– [(proportion of nitratein N uptake)/(proportion of nitrate in N supply)] was lowestin M. caerulea and highest in D. flexuosa. Whereas M. caeruleaand the dwarf shrubs always absorbed ammonium highly preferentially(relative uptake ratio for NO₃^– <0.20), D. flexuosashowed a strong preference for NO₃^– at low external nitrate(the relative uptake ratio for N0₃^– reaching a value of2.0 at 10% NO₃^–). The ecological significance of thisprominent high preference for NO₃^– at low NO₃^–/NH₄⁺ratio by D. flexuosa and its consequences for soil acidificationare briefly discussed. Key words: Ammonium, heathland lants, N0₃^–/NH₄⁺ ratio, nitrate, nitrate reductase activity, soil acidification, specific absorption rate     

Growth and Preferences for Ammonium or Nitrate Uptake by Barley in Relation to Root Termperature

Barley plants (Hordewn vulgare L. cv. Atem) were grown fromseed for 28 d in flowing solution culture, during which timeroot temperature was lowered decrementally to 5?C. Plants werethen subjected to root temperatures of 3, 5, 7, 9, 11, 13, 17or 25 ?C, with common air temperature of 25/15 ?C (day/night).Changes in growth, plant total N, and NO₃^– levels, andnet uptake of NH₄⁺ and NO₃^– from a maintained concentrationof 10 mmol m^–3 NH₄NO₃ were measured over 14 d. Dry matterproduction increased 6-fold with increasing root temperaturebetween 3–25 ?C. The growth response was biphasic followingan increase in root temperature. Phase I, lasting about 5 d,was characterized by high root specific growth rates relativeto those of the shoot, particularly on a fresh weight basis.During Phase I the shoot dry weight specific growth rates wereinversely related to root temperature between 3–13 ?C.Phase 2, from 5–14 d, was characterized by the approachtowards, and/or attainment of, balanced exponential growth betweenshoots and roots. Concentrations of total N in plant dry matterincreased with root temperature between 3–25 ?C, moreso in the shoots than roots and most acutely in the youngestfully expanded leaf (2?l–6?9% N). When N contents wereexpressed on a tissue fresh weight basis the variation withtemperature lessened and the highest concentration in the shootwas at 11 ?C. Uptake of N increased with root temperature, andat all temperatures uptake of NH₄⁺, exceeded that of NO₃^–,irrespective of time. The proportions of total N uptake over14 d absorbed in the form of NH₄⁺ were (%): 86, 91, 75, 77,76, 73, 77, and 80, respectively, at 3, 5, 7, 9, Il, 13, 17,and 25 ?C. At all temperatures the preference for NH₄⁺ overNO₃^– uptake increased with time. An inverse relationshipbetween root temperature (3–11 ?C) and the uptake of NH₄⁺as a proportion of total N uptake was apparent during PhaseI. The possible mechanisms by which root temperature limitsgrowth and influences N uptake are discussed. Key words: Hordeum vulgare, root temperature, ammonium, nitrate, ion uptake, growth rate     

The Composition of Phloem Exudate and Xylem Sap from Broccoli (Brassica oleracea var. italica) Supplied with NH4+, NO""3 or NH4NO3

Shelp, B. J. 1987. The composition of phloem exudate and xylemsap from broccoli (Brassica oleracea var. italica) suppliedwith NH⁺₄, NO₃ or NH₄NO₃.—J. exp. Bot. 38: 1619–1636. The detailed composition of xylem sap and exudate from stemincisions of attached inflorescences of broccoli (Brassica oleraceavar. italica) was compared in plants supplied with NH⁺₄, NO₃or NH₄NO₃. A phloem origin for the exudate was suggested fromthe high levels of sugars (71–133 mg cm^-3), amino acids(8·1-26·7 mg cm³) and K. (2·3–3·8mg cm³), the low levels of NO₃ and Ca, the high C: N (w/w) ratios(8·3–33), and the alkaline pH (7·2–7·3).In contrast, the xylem sap was mildly acidic (pH 5·6–6·0),and possessed lower levels of all organic and inorganic solutesbut NO₃ and Ca, and lower ratios of K: Ca, Mg: Ca and C: N (0·6–4·4). Glutamine was the predominant o-phthalaldehyde-reactive aminocompound in both transport fluids with the next most abundantamino acids dependent on sap type and N-form. Together witharginine, -aminobutyric acid, which was found only in the xylemstream, was enhanced by NH⁺₄compared to NO₃ -nutrition suggestingthat glutamate metabolism was stimulated in the roots. Underlimiting N the amino acid concentrations in the transport fluidswere greater with NH⁺₄ than with NO₃. NO₃ reduction occurredin both the root and shoot with the latter site predominatingover the entire N range (0-300 mol m³). Even though the compositionof nitrogenous solutes in the xylem was dependent on cultivarand N source, the composition of the phloem streams supplyingthe developing inflorescence was relatively unaffected. The data on the element composition of organs and phloem sapare interpreted to suggest that, in spite of the restrictedmobility of some elements such as B and Mn, a significant proportionof their total supply to developing sinks is carried in thephloem stream. Key words: Transport fluid composition, plant nutrition, phloem mobility.     

The Uptake and Flow of C, N and lons between Roots and Shoots in Ricinus communis L.

Seedlings of Ricinus communis L. cultivated in quartz sand weresupplied with a nutrient solution containing either 1 mol m^–3NO^–₃ or 1 mol m^–3 NH⁺₄ as the nitrogen source. Duringthe period between 41 and 51 d after sowing, the flows of N,C and inorganic ions between root and shoot were modelled andexpressed on a fresh weight basis. Plant growth was clearlyinhibited in the presence of NH⁺₄. In the xylem sap the majornitrogenous solutes were nitrate (74%) or glutamine (78%) innitrate or ammonium-fed plants, respectively. The pattern ofamino acids was not markedly influenced by nitrogen nutrition;glutamine was the dominant compound in both cases. NH⁺₄ wasnot transported in significant amounts in both treatments. Inthe phloem, nitrogen was transported almost exclusively in organicform, glutamine being the dominant nitrogenous solute, but theN-source affected the amino acids transported. Uptake of nitrogenand carbon per unit fresh weight was only slightly decreasedby ammonium. The partitioning of nitrogen was independent ofthe form of N-nutrition, although the flow of nitrogen and carbonin the phloem was enhanced in ammonium-fed plants. Cation uptakerates were halved in the presence of ammonium and lower quantitiesof K⁺, Na⁺ and Ca²⁺ but not of Mg²⁺ were transported to theshoot. As NH⁺₄ was balanced by a 30-fold increase in chloride in thesolution, chloride uptake was increased 6-fold under ammoniumnutrition. We concluded that ammonium was predominantly assimilated inthe root. Nitrate reduction and assimilation occurred in bothshoot and root. The assimilation of ammonium in roots of ammonium-fedplants was associated with a higher respiration rate. Key words: Ricinus communis, nitrogen nutrition (nitrate/ammonium), phloem, xylem, transport, partitioning, nitrogen, carbon, potassium, sodium, magnesium, calcium, chloride     

Plant Characteristics and Nutrient Composition and Mobility of Broccoli (Brassica oleracea var. italica) Supplied with NH4+, NO3"" or NH4NO3

Shelp, B. J. 1987. Plant characteristics and nutrient compositionand mobility of broccoli (Brassica oleracea var. italica) suppliedwith NH⁺₄, NO₃ or NH₄NO₃.—J. exp. Bot. 38: 1603–1618. The effects of varying NH⁺₄, NO₃ or NH₄NO₃ concentration onthe final plant characteristics, element composition, and accumulationof NO₃-N, NH⁺₄-N and organic-N were evaluated in broccoli (Brassicaoleracea var. italicacv. Futura and Premium Crop) plants culturedin vermiculite under greenhouse conditions supplemented withlight. NH⁺₄-grown plants were stunted and exhibited signs ofmarginal necrosis on the old leaves, accompanied by an accumulationof NH4. The tissue levels of N, P, Mn, Cu, Zn and B were generallyincreased by NH⁺₄ versus NO₃ nutrition whereas the reverse wastrue for Ca; Mg and K were only slightly affected, if at all.These results are attributed to: changes in element availabilityresulting from reduced rhizosphere pH due to NH⁺₄uptake ratherthan NO ₃uptake; competition of Ca uptake by NH⁺₄; and dilutionof N by increased vegetative growth with NO₃-nutrition. Theelement concentrations of N, P or K were similar in all tissueswhereas Ca, B and Mn were markedly less in the florets and youngleaves compared to mature leaves; this supports literature indicatingthat the former elements are phloem-mobile whereas the latterare not. Assuming that the nutrient supply for mature leavesis delivered principally via the xylem stream, the data suggestthat nutrients for developing leaves and florets are suppliedpredominantly in the phloem. If so, under our experimental conditions.Zn and Cu were also readily mobile in the phloem whereas Mgmovement was restricted. NH₄⁺ versus NO₄⁺ J nutrition alteredthe distribution of these elements. The two broccoli cultivarstested under the greenhouse environment differed in NH⁺₄ toleranceand in the distribution of K and Cu suggesting there was a geneticbasis for cultivar variation in mineral acquisition and redistribution. Key words: Plant nutrition, phloem mobility, elemental composition.     

Dynamic Interactions between Root NH4+ Influx and Long-Distance N Translocation in Rice: Insights into Feedback Processes

Ammonium influx into roots and N translocation to the shootswere measured in 3-week-old hydroponically grown rice seedlings(Oryza sativa L., cv. IR72) under conditions of N deprivationand NH₄⁺ resupply, using ¹³NH₄⁺as a tracer. Root NH₄⁺ influxwas repressed in plants continuously supplied with NH₄⁺ (at0.1 mM), but a high proportion of absorbed N (20 to 30%) wastranslocated to the shoot in the form of N assimilates duringthe 13-min loading and desorption periods. Interruption of exogenousNH₄⁺ supply for periods of 1 to 3 d caused NH₄⁺ influx to bede-repressed. This same treatment caused N translocation tothe shoot to decline rapidly, until, by 24 h, less than 5% ofthe absorbed ¹³N was translocated to the shoot, illustratinga clear priority of root over shoot N demand under conditionsof N deprivation. Upon resupplying 1 mM NH₄⁺, root NH₄⁺ influxresponded in a distinct four-phase pattern, exhibiting periodsin which NH₄⁺ influx was first enhanced and subsequently reduced.Notably, a 25 to 40% increase in root influx, peaking at {smalltilde}2 h following re-exposure was correlated with a 4- to5-fold enhancement in shoot translocation and a repression ofroot GS activity. The transient increase of NH₄⁺ influx wasalso observed in seedlings continuously supplied with NO₃^–and subsequently transferred to NH₄⁺. Extended exposure to NH₄⁺caused root NH₄⁺ influx to decrease progressively, while shoottranslocation was restored to {small tilde}30% of incoming NH₄⁺.The nature of the feedback control of NH₄⁺ influx as well asthe question of its inducibility are discussed. (Received August 7, 1998; Accepted September 21, 1998)