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     

Nitrate Nutrition and Temperature Effects on Wheat: a Synthesis of Plant Growth and Nitrogen Uptake in Relation to Metabolic and Physiological Processes

Lawlor, D. W., Boyle, F. A., Keys, A. J., Kendall, A. C. andYoung, A. T. 1988. Nitrate nutrition and temperature effectson wheat: a synthesis of plant growth and nitrogen uptake inrelation to metabolic and physiological processes.—J.exp. Bot. 39: 329-343. Growth of spring wheat was measured in cool (13°C day/10°Cnight) or warm (23°C/18°C) temperatures, combined withlarge and small amounts of nitrate fertilizer. The rate of growthof dry matter was less at cool temperatures but total growthover the same period of development was slightly greater inthe cool than in the warm. Main-shoot and tiller leaves grewslower and, despite growing for a longer period, were shorterin the cool than in the warm. They had greater fresh and drymass and content of starch and fructosans per unit area. Coolconditions increased root dry mass, root to shoot ratio andnitrogen content in dry matter. Additional nitrate increasedleaf area of main shoots slightly but of tillers greatly; itincreased leaf and tiller dry matter and total plant dry mass.Additional nitrate decreased the proportion of dry matter inroots and in stems and the N content of dry matter in all plantparts. Regulation of growth by temperature, nitrate supply andthe rôle of photosynthesis and nitrogen uptake, is consideredin relation to the mechanisms of incorporation of carbon andnitrogen into biochemical constituents. It is concluded thattemperature regulates the rate of protein synthesis, which determinesplant growth rate. Nitrogen flux into the plant is not directlylinked to protein synthesis so that the content of NO^–₃and of amino acids is related both to growth and to conditionsgoverning NO^–₃ uptake and its reduction. When nitrogensupply is large, growth is limited by temperature, not NO^–₃.Inadequate nitrate supply decreases protein synthesis (and thereforegrowth) more than it decreases carbon assimilation, so thatorgans such as roots and stems increase in dry matter relativeto shoots and all tissues have smaller proportions of nitrogenin dry matter. Cool conditions, although decreasing the rateof protein synthesis, increase its duration and decrease thesize of leaves, so that the content of protein per unit leafarea is greater in cool than in warm grown leaves. Consequencesof changes in the balance of N and C supply and growth ratefor dry matter distribution in plants are discussed. Key words: Wheat, nitrate nutrition, temperature     

N2 Fixation and Nitrate Uptake by White Clover Swards in Response to Root Temperature in Flowing Solution Culture

Nodulated white clover plants (Trifolium repens L. cv. Huia)were grown as simulated swards for 71 d in flowing nutrientsolutions with roots at 11 C and shoots at 20/15 C, day/night,under natural illumination. Root temperatures were then changedto 3, 5, 7, 11, 13, 17 or 25 C and the total N₂, fixation over21 d was measured in the absence of a supply mineral N. Alltreatments were subsequently supplied with 10 mmol m^–2NO₂ ^– in the flowing solutions for 14 d, and the relativeuptake of N by N₂, fixation and NO₃ ^– uptake was compared.Net uptake of K⁺ was measured on a daily basis. Root temperature had little effect on root d. wt over the 35-dexperimental period, but shoot d. wt increased by a factor of3.5 between 3 and 25 C, with the sharpest increase occurringat 7–11 C. Shoot: root d. wt ratios increased from 25to 68 with increasing temperature at 7–25 C. N₂-fixationper plant (in the absence of NO₂ ^–) increased with roottemperature at 3–13C, but showed little change above13 C. The ratios of N₂ fixation: NO₂ ^– uptake over 14d (mol N: mol N) were 0.47–0.77 at 3–7 C, 092–154at 11–17 C, and 046 at 25 C, reflecting the dominanceof NO₃ ^– uptake over N₂ fixation at extremes of high andlow root temperature. The total uptake of N varied only slightlyat 11–25 –C (095–110 mmol N plant^–1),the decline in N₂ fixation as root temperature increased above11 C was compensated for by the increase in NO^– ₃ uptake.The % N in shoot dry matter declined with decreasing root temperature,from 32% at 13 C to 15% at 3 C. In contrast, concentrationsof N expressed on a shoot water content basis showed a modestdecrease with increasing temperature, from 345 mol m^–3at 3 C to 290 mol m^–3 at 25 C. Trifolium repens L, white clover, root temperature, N₂ fixation, potassium uptake, nitrate uptake, flowing solution culture     

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 Effect of Root Temperature on Growth and Uptake of Ammonium and Nitrate by Brassica napus L. cv. Bien venu in Flowing Solution Culture: II. UPTAKE FROM SOLUTIONS CONTAINING NH4NO3

Macduff, J. H., Hopper, M. J. and Wild, A. 1987. The effectof root temperature on growth and uptake of ammonium and nitrateby Brassica napus L. CV. Bien venu in flowing solution culture.II. Uptake from solutions containing NH₄NO₃.—J. exp. Bot.38: 53–66 The effects of root temperature on uptake and assimilation ofNH₄⁺ and NO₃^– by oilseed rape (Brassica napus L. CV. Bienvenu) were examined. Plants were grown for 49 d in flowing nutrientsolution at pH 6?0 with root temperature decrementally reducedfrom 20?C to 5?C; and then exposed to different root temperatures(3, 5, 7, 9, 11, 13, 17 or 25?C) held constant for 14 d. Theair temperature was 20/15?C day/night and nitrogen was suppliedautomatically to maintain 10 mmol m^–3 NH₄NO₃ in solution.Total uptake of nitrogen over 14 d increased threefold between3–13?C but was constant above 13?C. Net uptake of NH₄⁺exceeded that of NO₃^– at all temperatures except 17?C,and represented 47–65% of the total uptake of nitrogen.Unit absorption rates of NH₄⁺ and of 1?5–2?7 for NO₃^–suggested that NO₃^– absorption was more sensitive thanNH₄⁺ absorption to temperature. Rates of absorption were relativelystable at 3?C and 5?C compared with those at 17?C and 25?C whichincreased sharply after 10 d. Tissue concentration of N in theshoot, expressed on a fresh weight basis, was independent ofroot temperature throughout, but doubled between 3–25?Cwhen expressed on a dry weight basis. The apparent proportionof net uptake of NO₃^– that was assimilated was inverselyrelated to root temperature. The results are used to examinethe relation between unit absorption rate adn shoot:root ratioin the context of short and long term responses to change ofroot temperature Key words: Brassica napus, oilseed rape, root temperature, nitrogen uptake     

The Effect of Root Temperature on Growth and Uptake of Ammonium and Nitrate by Brassica napus L. in Flowing Solution Culture: I. GROWTH

Macduff, J. H., Hopper, M. J. and Wild, A. 1987. The effectof root temperature on growth and uptake of ammonium and nitrateby Brassica napus L. in flowing solution culture. I. Growth.—J.exp. Bot. 38: 42–52 Oilseed rape (Brassica napus L. cv. Bien venu) was grown for49 d in flowing nutrient solution at pH 6?0 with root temperaturedecrementally reduced from 20?C to 5?C; and then exposed todifferent root temperatures (3, 5, 7, 9, 11, 13,17 or 25?C)held constant for 14 d. The air temperature was 20/15?C day/nightand nitrogen was supplied automatically to maintain 10 mmolm^–3 NH₄NO₃ in solution. Total dry matter production wasexponential with time and similar at all root temperatures givinga specific growth rate of 0?0784 g g^–1 d^–1. Partitioningof dry matter was influenced by root temperature; shoot: rootratios increased during treatment at 17?C and 25?C but decreasedafter 5 d at 3?C and 5?C. The ratio of shoot specific growthrate: root specific growth rate increased with the ratio ofwater soluble carbohydrates (shoot: root). Concentrations ofwater soluble carbohydrates in shoot and root were inverselyrelated to root temperature; at 3, 5 and 7?C they increasedin stem + petioles throughout treatment, coinciding with a decreasein the weight of tissue water per unit dry matter. These resultssuggest that the accumulation of soluble carbohydrates at lowtemperature is the result of metabolic imbalance and of osmoticadjustment to water stress. Key words: Brassica napus, oilseed rape, root temperature, specific growth rate     

Acclimation of NO-3 fluxes to low root temperature by Brassica napus in relation to NO-3 supply

Acclimation of NO^–₃ transport fluxes (influx, efflux)in roots of oilseed rape (Brassica napus L. cv. Bien venu) andtheir sensitivity to growth at low root temperature was studiedin relation to external NO^–₃ supply, defined by constantconcentrations ranging from sub- to supra-optimal with respectto plant growth rate. Plants were grown from seed in flowingnutrient solutions containing 250 mmol m^–3 NO^–₃at 17°C for 20d, and solution temperature in half the cultureunits was then lowered decrementally over 3 d to 7°C. Threedays later plants were supplied with NO^–₃ at 1, 10, 100or 1000 mmol m^–3 maintained for 18 d. Dry matter productionwas decreased more by low root zone temperature than low [NO^–₃]_e. Root specific growth rates were inversely related to [NO^–₃]_eand shoot:root ratios increased with time at [NO^–₃]_e between10–1000 mmol m^–3. Net uptake of NO^–₃ at 17°Cwas twice that at 7°C, and at both temperatures it doubledwith increasing [NO^–₃]_e between 1–10 mmol m^–3with further small increases at higher [NO^–₃]_e. Mean unitabsorption rates of NO^–₃ between 0–6 d and 6–14d were linearly related (r² of 0.79–0.99) to log₁₀[NO].Steady-state Q₁₀ (7–17°C) for uptake between 0–6d were 0.91, 1.62, 1.27, and 1.10, respectively, at [NO^–₃]_eof 1, 10, 100, and 1000 mmol m^–3, compared with correspondingvalues of 0.98, 1.38, 1.68, and 1.89 between 6–14 d. Thedata indicated that net uptake rates at 7 and 17°C divergedover time at high [NO^–₃]_e. Short-term uptake rates from1 mol m^–3 NO^–₃ measured at 17°C were higherin plants grown with roots at 7°C than at 17°C; for7°C plants there was a strong inverse linear relationship(r²=0.94) between uptake rate and treatment log₁₀ [NO^–₃]_ewhilst rates in 17°C plants were independent of prior [NO^–₃]_e. Rates of NO^–₃ influx and efflux under different steady-stateconditions of NO^–₃ supply and root temperature were calculatedfrom dilution of ¹⁵N added to culture solutions. Efflux wassubstantial relative to net uptake in all treatments, and wasinversely related to [NO^–₃]_e at 17°C but not at 7°C.Ratios of influx: efflux ranged from 1.6–2.9 at 17°Cand 1.3–1.8 at 7°C, indicating the proportionatelygreater impact of efflux at low root temperature. Ratios ofefflux: net uptake were 0.53–1.56 at 17°C and 1.21–3.58at 7°C. The apparent sensitivities of influx and effluxto steady-state root temperature varied with [NO^–₃]_e.Both fluxes were higher at 17°C than 7°C in the presenceof 100–1000 mmol m^–3 NO^–₃ but the trend wasreversed at 1–10 mmol m^–3 NO. Concentrations oftotal N measured in xylem exudate were at least 2-fold higherat 7°C compared with 17°C, attributable mainly to higherconcentrations of NO^–₃ glutamine and proline. The resultsare discussed in terms of acclimatory and other responses shownby the NO^–₃ transport system under conditions of limitingNO^–₃ supply and low root temperature. Key words: Brassica napus, nitrate supply, efflux, influx, root temperature, xylem exudate     

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     

Effects of nitrogen nutrition on salt-stressed Nicotiana tabacum var. Samsun in vitro plantlets

Tobacco shoots were grown in vitro for 35 d, in MS culture mediummodified to include various sources (nitrate-N, ammonium-N ora mixture) and levels (0–120 mM) of N, and in the presenceof 0–180 mM NaCI or iso-osmotic concentrations of mannitol.Growth of control plantlets was significantly inhibited whenNH₄⁺-N was the sole N source, and at high (120 mM) NO^–₃-N supply. Under conditions of salt stress (90 and 180 mM NaCI)growth was repressed, with roots being more severely affectedthan shoots. Salinity also inhibited root emergence in vitro.The only alleviation of the salt stress by nitrate nutritionobserved in this study was on shoot growth parameters of plantletsgrown on 60 mM NO^–₃-N and 90 mM NaCI. Although both weresignificantly inhibited by NaCI, nitrate reduc-tase activitywas more severely affected than nitrate uptake. When mannitolreplaced NaCI in the culture medium, similar Inhibition of growth,nutrient uptake and enzyme activity were recorded. These observations,together with the relatively low recorded values for Na⁺ andCI^– uptake, indicate that under in vitro salt stress conditionsthe negative effects of NaCI are primarily osmotic. Key words: Growth, nitrogen metabolism, osmotic stress, salinity     

Carbon and Nitrogen Assimilation by Vicia faba L. at Low Temperature: the Importance of Concentration and Form of Applied-N

Low temperature (6 C) growth was examined in two cultivarsof Vicia faba L. supplied with 4 and 20 mol m^–3 N as nitrateor urea. Both cultivars showed similar growth responses to increasedapplied-N concentration regardless of N-form. Total leaf areaincreased, as did root, stem and leaf dry weight, total carboncontent and total nitrogen content. In contrast to findingsat higher growth temperatures, 20 mol m^–3 urea-N gavesubstantially greater growth (all parameters measured) than20 mol m^–3 nitrate-N. The increased carbon content per plant associated with increasedapplied nitrate or urea concentration, or with urea in comparisonto nitrate, was due to a greater leaf area per plant for CO₂uptake and not an increased CO₂, uptake per unit area, carbon,chlorophyll or dry weight, all of which either remained constantor decreased. Nitrate reductase activity was substantial inplants given nitrate but negligible in plants given urea. Neitherfree nitrate nor free urea contributed greatly to nitrogen levelsin plant tissues. It is concluded that there is no evidence for a restrictionin nitrate reduction at 6 C, and it is likely that urea givesgreater growth than nitrate because of greater rates of uptake. Vicia faba, broad bean, low temperature growth, carbon assimilation, nitrogen assimilation     

Nitrate Effects on Pre-emergence Growth and Emergence Percentage of Wheat (Triticum aestivum L.) from Different Sowing Depths

The effects of a range of applied nitrate (NO₃^–) concentrations(0–20 mol m3) on germination and emergence percentageof Triticum aestivum L. cv. Otane were examined at 30, 60, 90and 120 mm sowing depths. Germination percentage was not affectedby either sowing depth or applied NO₃^– concentration whereasemergence percentage decreased with increased sowing depth regardlessof applied NO₃^– concentration. Nitrate did not affectemergence percentage at 30 mm sowing depth, but at 60 to 120mm depth, emergence percentage decreased sharply with an increasedapplied NO₃^– concentration of 0 to 1·0 mol m^–3then decreased only slightly with further increases in appliedNO₃^– of about 5·0 mol m^–3. Root and shoot growth, NO₃^– accumulation and nitrate reductaseactivity (NRA) of plants supplied with 0, 1·0 and 1·0mol m^–3 NO₃^– at a sowing depth of 60 mm were measuredprior to emergence. The coleoptile of all seedlings opened withinthe substrate. Prior to emergence from the substrate, shootextension growth was unaffected by additional NO₃^– butshoot fr. wt. and dry wt. were both greater at 1·0 and1·0 mol m^–3 NO₃^– than with zero NO₃^–.Root dry wt. was unaffected by NO₃^–. Nitrate concentrationand NRA in root and shoot were always low without NO₃^–.At 1·0 and 10 mol m3 NO₃^–, NO₃^– accumulatedin the root and shoot to concentrations substantially greaterthan that applied and caused the induction of NRA. Regardlessof the applied NO₃^– concentration, seedlings which failedto emerge still had substantial seed reserves one month afterplanting. Coleoptile length was substantially less for seedlingswhich did not emerge than for seedlings which emerged, but wasnot affected by NO₃^–. It is proposed that (a) decreasedemergence percentage with increased sowing depth was due tothe emergence of leaf I from the coleoptile within the substrateand (b) decreased emergence percentage with additional NO₃^–was due to the increased expansion of leaf 1 within the substrateresulting in greater folding and damage of the leaf. Key words: Triticum aestivwn L., nitrate, sowing depth, seedling growth, seedling emergence     

The Distribution of Nitrate Assimilation Between Root and Shoot in Vicia faba L.

Nitrate assimilation was examined in two cultivars (Banner Winterand Herz Freya) of Vicia faba L. supplied with a range of nitrateconcentrations. The distribution between root and shoot wasassessed. The cultivars showed responses to increased applied nitrateconcentration. Total plant dry weight and carbon content remainedconstant while shoot: root dry weight ratio, total plant nitrogen,total plant leaf area and specific leaf area (SLA) all increased.The proportion of total plant nitrate and nitrate reductase(NR) activity found in the shoot of both cultivars increasedwith applied nitrate concentrations as did NO₃^–: Kjeldahl-Nratios of xylem sap. The cultivars differed in that a greaterproportion of total plant NR activity occurred in the shootof cv. Herz Freya at all applied nitrate concentrations, andits xylem sap NO₃^–: Kjeldahl-N ratio and SLA were consistentlygreater. It is concluded that the distribution of nitrate assimilationbetween root and shoot of V. faba varies both with cultivarand with external nitrate concentration. Vicia faba L., field bean, nitrate assimilation, nitrate reductase, xylem sap analysis     

Root Zone Temperature Effects on Growth and Phosphate Absorption in Rape Brassica napus cv. Emerald

Solution culture experiments with fodder rape (Brassica napuscv. Emerald) show that reduced root temperatures appear to havelittle effect on phosphate inflow over a wide range of P concentration.At a cool root temperature (10 ?C) plant growth rate was reducedbut this was compensated for by a low root: shoot ratio, sothat inflow remained relatively steady. An increased inflowper unit length of root was only achieved at an elevated roottemperature of 35 ?C. The minimum phosphate concentration towhich plants could lower the culture solution (C_mln) rangedfrom 0.15 to 2.5 mmol m^–3 according to whether roots wereat a low (5 ?C) or high (35 ?C) temperature respectively. Thetotal phosphorus concentration in tissues was affected by rootzone temperature and at low root temperatures this could bea growth limiting factor. The organic (assimilated) fractionof P in shoot tissues was smaller in low temperature plants.These showed visual symptoms of apparent P deficiency. Levelsof inorganic P in roots may also be a factor in feedback ofcontrol of inflow. Key words: Temperature, Roots, Phosphate, Rape (Brassica napus)     

Nitrogen Utilization in N-limited Barley during Vegetative and Generative Growth: I. GROWTH AND NITRATE UPTAKE KINETICS IN VEGETATIVE CULTURES GROWN AT DIFFERENT RELATIVE ADDITION RATES OF NITRATE-N

Growth and nitrate uptake kinetics in vegetatively growing barley(Hordeum vulgare L., cvs Laevigatum, Golf, and Mette) were investigatedin solution culture under long-term limitations of externalnitrogen availability. Nitrate was fed to the cultures at relativeaddition rates (RA) ranging from 0.02 to 0.2 d^–1. Therelative growth rate (RG, calculated for total plant dry weight)correlated well with RA in the range 0.02 to 0.07 d^–1.In the RA range from 0.07 to 0.2 d^–1 RG continued to increase,but an increasing fraction of nitrogen, added and absorbed,was apparently stored rather than used for structural growth.The RG of the roots was less affected by RA. V_max, for net nitrateuptake increased with RA up to 0.11 d^–1, but decreasedat higher RA. The decline in V_max coincided with a build-upof nitrate stores in both roots and shoots. V_max, expressedper unit nitrogen in the plants (the relative V_max, was higherthan required for maintenance of growth (up to 30-fold) at lowRA, whereas at higher RA the relative V_max decreased. Kineticpredictions of steady-state external nitrate concentrationsduring N-limited growth ranged from 0.2 to 5.0 mmol m^–3over the RG range 0.02 to 0.11 d_–1. It is suggested thatthe nitrate uptake system is not under specific regulation atlow RA, but co-ordinated with root protein synthesis and growthin general. At RA higher than 0.11 d_–1, however, specificregulation of nitrate uptake, possibly via root nitrate pools,become important. The three cultivars showed very similar growthand nitrate uptake characteristics. Key words: Barley, growth, nitrogen limitation, nitrate uptake, kinetics     

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     

Growth and Partitioning in Pascopyrum smithii (C3) and Bouteloua gracilis(C4) as Influenced by Carbon Dioxide and Temperature

This study investigated how CO₂and temperature affect dry weight(d.wt) accumulation, total nonstructural carbohydrate (TNC)concentration, and partitioning of C and N among organs of twoimportant grasses of the shortgrass steppe,Pascopyrum smithiiRydb. (C₃) andBouteloua gracilis(H.B.K.) Lag. ex Steud. (C₄).Treatment combinations comprised two temperatures (20 and 35°C)at two concentrations of CO₂(380 and 750 µmol mol^-1),and two additional temperatures of 25 and 30°C at 750 µmolmol^-1CO₂. Plants were maintained under favourable nutrient andsoil moisture and harvested following 21, 35, and 49d of treatment.CO₂-induced growth enhancements were greatest at temperaturesconsidered favourable for growth of these grasses. Comparedto growth at 380 µmol mol^-1CO₂, final d.wt of CO₂-enrichedP.smithiiincreased 84% at 20°C, but only 4% at 35°C. Finald.wt ofB. graciliswas unaffected by CO₂at 20°C, but wasenhanced by 28% at 35°C. Root:shoot ratios remained relativelyconstant across CO₂levels, but increased inP. smithiiwith reductionin temperature. These partitioning results were adequately explainedby the theory of balanced root and shoot activity. Favourablegrowth temperatures led to CO₂-induced accumulations of TNCin leaves of both species, and in stems ofP. smithii, whichgenerally reflected responses of above-ground d.wt partitioningto CO₂. However, CO₂-induced decreases in plant tissue N concentrationswere more evident forP. smithii. Roots of CO₂-enrichedP. smithiihadgreater total N content at 20°C, an allocation of N below-groundthat may be an especially important adaptation for C₃plants.Tissue N contents ofB. graciliswere unaffected by CO₂. Resultssuggest CO₂enrichment may lead to reduced N requirements forgrowth in C₃plants and lower shoot N concentration, especiallyat favourable growth temperatures. Acclimation to CO₂; blue grama; Bouteloua gracilis ; carbohydrate; climate change; global change; grass; growth; growth temperature optima; nitrogen; N uptake; Pascopyrum smithii; western wheatgrass     

Growth of Phaseolus vulgaris on Various Nitrogen Sources: The Importance of Urease

Rhizobium-inoculatcd plants of Phaseolus vulgaris L. were grownwith different N-sources (nitrate, ammonium, urea) and differentconcentrations of urea. The distribution of growth between plantparts varied with N-sources. Nitrate and ammonium were moreinhibitory to nodulation than urea, which at 40 mol m^–3N had no effect. Urease activity varied in amount and locationover a range of urea concentrations. At higher concentrations,more urea was transported to and increased urease activity wasfound in the shoot Lower levels of activity in plants relianton N₂-fixation were consistent with a ureide-degradation pathwaynot involving urea. Moderate doses of urea could be assimilatedconcomitantly with N₂-fixation. At higher levels of appliedurea, nodulation and ureide transport to the shoots were reduced,although increased growth could not be maintained at concentrationsof applied urea greater than 6.0 mol m^–3 urea N. Key words: Phaseolus vulgaris, growth, nitrogen source, urease     

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     

The Combined Effects of Salinity and Root Anoxia on Growth and Net Na+ and K+-accumulation in Zea mays Grown in Solution Culture

The effects of excess salinity and oxygen deficiency on growthand solute relations in Zea mays L. cv. Pioneer 3906 were examinedin greenhouse experiments. The roots of plants 14 d old growingin nutrient solution containing additions of NaCl in the range1.0–200 mol m^–3 were either exposed to a severedeficiency of O₂ by bubbling with nitrogen gas (N₂ treatment),or maintained with a supply of air (controls), for a periodof 1–7 d. The threshold NaCl concentration resulting inappreciable inhibition of leaf extension, and shoot f. wt gainin controls was between 10 and 25 mol m^–3. At 25 mol m^–3NaCl the ratio of Na+/K+ transported to shoots was about 20times greater than in plants in 1.0 mol m^–3 NaCl. Theeffect of addition of NaCl to the nutrient solution was to enhanceNa+ movement but simultaneously depress the rate of K+ transportto shoots (per g f. wt roots). Interactions between NaCl levels and aeration treatment wereshown by analyses of variance to be statistically significantfor leaf extension, shoot and root f. wt gains, Na+ and K+ concentrationsin shoots and roots. When roots were N₂-treated, shoot and rootgrowth were depressed, the effect of aeration treatment beinggreatest at NaCl concentrations of 50 mol m^–3 or less.Additionally, N₂-treatment greatly accelerated Na- transportto shoots while depressing K+ transport still further, so thatat 10 mol m^–3 NaCl the ratio Na+/K+ acquired by the shootswas 230 times greater than in controls. Over the concentrationrange 1.0 to 50 mol m^–3 NaCl, the ratio Na+/K+ transportedto shoots by anoxic roots increased by a factor of 860. Mechanisms controlling changes in solute flux to the shoot,and the significance in relation to plant tolerance of excesssalts or oxygen deficiency are discussed. Anaerobic, corn, flooding, maize, oxygen-deficiency, salinity     

Regeneration from Rhizome Fragments of Agropyron repens (L.) Beauv. III. Effects of Nitrogen and Temperature on the Development of Dominance Amongst Shoots on Multi-node Fragments

Not all buds developed equally when 7-node rhizome fragmentsof Agropyron repens (L.) Beauv. were incubated in the dark at23 °C. Instead, after an initial flush of several shoots,buds were inhibited in a highly ordered sequence to leave onlyone dominant shoot growing. Applying an exogenous supply ofnitrogen KNO₃) early during this sequence increased the meanshoot lengths and delayed the onset of dominance. Additionally,the application of nitrogen after eight days incubation alteredthe sequence of shoot growth such that, in some instances, smallrapidly-growing basal shoots ‘dominated’ largerand more slowly-growing apical ones. Dominance (correlativeinhibition) was maintained in untreated fragments for up to383 days Numbers of active budsand shoot extensionrate weremaximal intherange 13°to 23 °C where dominance was establishedwithin 30 days. Incontrast only 6 per cent of rhizome fragmentskept at 33 °C had dominant shoots after 65 days. At 3 °Cshoot growth was so slow that dominance was not permanentlyestablished within 150 days. Numbers ofactive budsand shoot extensionrate weremaximal intherange 13° to 23°C where dominance was establishedwithin 30 days. Incontrast only 6 per cent of rhizome fragmentskept at 33 °C had dominant shoots after 65 days. At 3 °Cshoot growth was so slow that dominance was not permanentlyestablished within 150 days It is suggested that the effects of nitrogenand temperatureon dominance in multi-noderhizome fragments can be interpretedin terms of competition for nutrients between shoots, and theantagonistic effects of nitrogen on an auxin-mediated inhibitionby the dominant shoot.