Efficiency of nitrate uptake in spinach: impact of external nitrate concentration and relative growth rate on nitrate influx and efflux

Regulation of nitrate influx and efflux in spinach (Spinacia oleracea L., cv. Subito), was studied in short-term label experiments with 13N- and 15N-nitrate. Nitrate fluxes were examined in relation to the N demand for growth, defined as relative growth rate (RGR) times plant N concentration. Plants were grown at different nitrate concentrations (0.8 and 4 mM), with mineral composition of growth and uptake solutions identical. Nitrate influx, efflux and net nitrate uptake rate (NNUR) were independent of the external nitrate concentration, despite differences in internal nitrate concentration. At both N regimes, NNUR was adequate to meet the N demand for growth. RGR-related signals predominantly determined the nitrate fluxes. At high RGR (0.25 g g-1 day-1), nitrate influx was 20 to 40% lower and nitrate efflux was 50 to 70% lower than at lower RGR (0.17 g g-1 day-1); efflux:influx ratio (E:I) declined from 0.5 at low RGR to 0.2 at higher RGR. Thus, the efficiency of NNUR substantially increased with increasing RGR. Differences in nitrate translocation between morning and afternoon coincided with differences in nitrate efflux, which is in accordance with the suggested regulation of nitrate efflux by the root cytoplasmic nitrate concentration. This revised version was published online in July 2006 with corrections to the Cover Date.     

Relationship of nitrate supply with growth rate, plasma membrane-bound and cytosolic nitrate reductase, and tissue nitrate content in tobacco plants

cNR, cytosolic nitrate reductase
PM-NR, plasma membrane-bound nitrate reductase

Activities of plasma membrane-bound nitrate reductase (PM-NR) and cytosolic nitrate reductase (cNR) in tobacco (Nicotiana tabacum L. cv. Samsun) are regulated differently, depending upon the nitrate supply to the culture medium (in sand culture). The cNR activity of roots was higher at low nitrate concentrations with the maximum at 5 mM nitrate supply and declined to low values beyond 5 mM . In contrast, the PM-NR activity of roots increased with higher nitrate concentrations with the maximum at 25 mM nitrate and clearly decreased only at 40 mM . This high PM-NR activity correlated with a low growth rate and might be one of the responses to excess nitrate. Internal nitrate and total nitrogen content of the tissues, however, showed a relative minimum in shoots and in roots of between 15 and 25 mM external nitrate. With declining PM-NR activities beyond 25 mM external nitrate, the nitrate content in the tissue increased indicating an inverse relationship between tissue nitrate content and root PM-NR activity. In leaves both NR activities (cNR and PM-NR) correlated with the internal nitrate content, but with a different response at low nitrate.     

Nitrite in the root zone and its effects on ion uptake and growth of wheat seedlings

The immediate and posteffects of various concentrations of NaNO₂ on ion uptake of wheat ( Triticum aestivum L. cv. GK Öthalom) seedlings were studied at different pH values. Without pretreatment, the higher the concentration of NaNO₂ the greater was the decrease in uptake of K⁺ into the roots, both at pH 4 and pH 6. At pH 6 but not at pH 4 the reverse was true when the seedlings were pretreated with NaNO₂. Due to the high Na⁺ content of the roots, an effect of Na⁺ in this process cannot be excluded. Nitrite was taken up by the roots more rapidly than nitrate. Nitrite at 0.1 m M in the medium induced the development of an uptake system for both NO⁻₂ and NO⁻₃ in wheat roots. At higher concentrations pretreatment with NO⁻₂ decreased NO⁻₃ uptake by the roots, but NO₃ did not inhibit the uptake of NO₂. The toxic effect of NO⁻₂ was strongly pH dependent. Lower pH of the external solution led to an increased inhibition by NO⁻₂ of both ion uptake and growth of seedlings. The inhibitory effect of NO⁻₂ differed considerably for roots and shoots. The roots and especially the root hairs were particularly sensitive to NO⁻₂ treatment.     

Effects of carbon and nitrate additions to soil upon leaching of nitrate,microbial predators and nitrogen uptake by plants

Amendments with glucose significantly reduced the amount of nitrate leached from a sandy soil amended with nitrate. The decrease was most likely caused by immobilisation of the nitrate into microbial cells. Populations of ciliates and flagellates and amoebae, but not nematodes, increased 7–14 days following glucose amendments. Mineralisation of the immobilised nitrate occurred during this period. Some of the mineralised nitrogen appeared to be available to ryegrass plants only if the roots exploited most of the soil during the period of maximum predator activity. After 28 days, 44% of the organic N remaining in the soil after leaching was taken up by the plants. The difference developed over the last 2 weeks when amoebal populations were large.     

氮过量吸收对地肤暗呼吸速率及相对生长速率的影响

为研究N过量吸收对植物生长的作用,以耐盐植物地肤(Kochia scoparia)作为研究对象,设置3个不同的施N处理,测量了不同生长时期的N含量、暗呼吸速率、生物量和相对生长速率(RGR)。结果表明:在N过量吸收的情况下,多余的N对暗呼吸速率并没有显著的影响,导致了暗呼吸中N的利用效率变低;单位质量暗呼吸速率与相对生长速率(RGR)有很好的线性相关,并且直线的斜率和截距并不受氮素过量吸收的影响,表明单位质量暗呼吸速率与RGR的关系不受施氮水平的影响;暗呼吸速率与总N的异速关系中,幂指数的大小与施N量相关,施N量越大对应的幂指数越小。     

Methods and applications to control the uptake rate of carbon

Methods to control carbon and nutrient uptake at different availability of carbon were tested on plants of birch (Betula pendula Roth.) and tomato (Lycopersicon esculentum Mill. cv. Solentos). The present paper accounts for the methods and the possibility to maintain steady-state, i.e., a long-term and stable physiological state of acclimated plants. Steady-state comprises, by definition, equality between constant relative growth rates, and relative uptake rates of carbon and nutrients. Two methods were tested. The first, not previously applied, method (a), was based on a constant relative addition rate of carbon, R_AC. In the second method (b), a constant concentration of CO₂ in the air, c_a, was used to attain non-limiting conditions. The methods are analogous to those used by us to control plant nutrition, and the generality of fluxes to quantify supply as well as uptake and growth was verified. Thus, different R_AC resulted in clear-cut responses, from strong reduction to non-limitation of uptake and growth, whereas different c_a levels in the range 100 to 700 ppm had comparatively small effects, with an unclear causality. Non-limiting conditions were achieved at c_a≥ 200 ppm. Effects reported in the literature have been based upon the control of c_a, similarly to method (b), whereas results comparable to those obtained with method (a) are lacking. Transpiration rate increased rapidly at c_a < 200 ppm CO₂, and at low R_AC levels, ≤ 0.1 day^?1, wilting tendencies were observed. Elevated c_a, 500 or 700 ppm, did not increase the relative growth rate (R_G) but reduced transpiration and increased both nitrogen productivity (growth rate per unit of nitrogen in the plant) and transpiration productivity (growth rate per unit of water transpired by the plant). Obviously, effects of c_a may be due to changed transpiration rate rather than to changed quantitative availability of CO₂. Relative uptake (R_UC) and growth (R_G) rates were closely equal to the R_AC applied (R_AC? R_UC? R_G); i.e., the purely mathematical conditions defining steady-state were fulfilled. This unambiguous and straightforward test of reliability confirms that experimental artefacts did not produce uncontrolled or unintended effects, so that the new technique allows an accurate control of CO₂ uptake and plant growth. The results add to previous databases and reference systems, where limiting conditions grade and classify plant performance as deviations from maximum growth. Evidently, methodology in experimentation and in evaluation of plant responses, can be based upon unifying concepts and general theories.     

Research note: Comparison of growth and nitrate uptake by New England Porphyra species from different tidal elevations in relation to desiccation

Desiccation stress can determine the upper distribution limits and may enhance the uptake of nitrate and ammonium of eulittoral algal species. Upper shore species may exhibit greater stimulation of nitrate uptake following desiccation and achieve maximum uptake at higher desiccation levels. The objective of this study was to determine whether Porphyra species from different vertical elevations respond differently to the desiccation stress, in terms of growth and nitrate uptake. A eulittoral species ( Porphyra umbilicalis) and a sublittoral species ( P. amplissima ) were compared in the present study. Samples were exposed to air for 0, 30 min (40 ± 10% water loss) and 2 h (90 ± 5% water loss), after an initial 4 h light period every day. Desiccation was more stressful to the sublittoral species, Porphyra amplissima, than to the eulittoral species, P. umbilicalis . When tissues were exposed for 2 h daily, P. amplissima lost weight over a 24 h day, while the growth rate of P. umbilicalis dropped by only 30% compared with that of continuously submerged blades. Nitrate uptake rate of sublittoral P. amplissima was only 73% (40 ± 10% water loss) and 62% (90 ± 5% water loss) of that of continuously submerged tissue. Nitrate uptake rates of P. umbilicalis were not significantly affected by desiccation. These results suggest that species in the eulittoral zone, which have longer exposure times, have a higher time-use efficiency than the sublittoral species in terms of nitrate uptake. This indicates a possible correlation between nitrate uptake and observed vertical distribution patterns.     

Short- and long-term effects of a change in the spatial distribution of nitrate in the root zone on N uptake, growth and root development of young lettuce plants

Abstract. The effects of a change in the distribution of nitrate within the root zone on N uptake and growth were studied using young lettuce plants after reducing the proportion of their root systems supplied with nitrate from 100 to ca 10% in split-root experiments in the glasshouse. The main effects of the localized nitrate supply were concentrated in a 2-week period immediately after the treatment was imposed, when a temporary reduction in nitrate uptake caused the gradual development of N deficiency and a decline in plant growth rate. The plants adapted to the change in nitrate distribution, initially by increasing unit absorption rates (uptake rates per unit weight of root) and more gradually by increasing production of new roots in the high-nitrate zone. As a result, relative N uptake rates and relative growth rates were restored to the same levels as for control plants (given a spatially uniform N supply throughout) after ca 12d, even though only ca 12–15% of their roots were exposed to nitrate at this time. Thereafter, the plants continued to adapt by concentrating new root growth in the nitrate-containing zone, ultimately allowing unit absorption rates to return to normal. There was no evidence of any significant N deficiency in the plants after the initial adaptive response was complete, even though the total-N concentrations of the plants given the localized supply were consistently less than those given the uniform N treatment, and nitrate concentrations in the petiole sap were generally lower in leaves on one side of the plant (because of limited lateral movement of nitrate between xylem vessels during its transport to the shoot). The delay in the initiation of an adaptive response caused a significant check in growth, and the resulting relative weight differences were maintained throughout the subsequent life of the plant. Plants in all treatments matured on the same date, so yields for those grown with the localized supply were less than those of the control, and could not be recovered by delaying final harvest without unacceptable loss of quality. The pattern of the changes in N uptake and plant growth, and the effect on final yield, were similar to those exhibited by young lettuce plants subjected to a temporary interruption in nitrate supply, suggesting that the reduction in final yield for plants grown with the localized supply was largely the effect of the check in growth which occurred whilst the Plants were adapting to the change in nitrate distribution during the early part of the experiment. This implies that the rate of dry matter production of young lettuce plants can be altered by N treatment without affecting their rate of physiological development.     

Relative growth rate, biomass allocation pattern and water use efficiency of three wheat cultivars during early ontogeny as dependent on water availability

We have investigated the water use efficiency of whole plants and selected leaves and allocation patterns of three wheat cultivars (Mexipak, Nesser and Katya) to explore how variation in these traits can contribute to the ability to grow in dry environments. The cultivars exhibited considerable differences in biomass allocation and water use efficiency. Cultivars with higher growth rates of roots and higher proportions of biomass in roots (Nesser and Katya) also had higher leaf growth rates, higher proportions of their biomass as leaves and higher leaf area ratios. These same cultivars had lower rates of transpiration per unit leaf area or unit root weight and higher biomass production per unit water use. They also had higher ratios of photosynthesis to transpiration, and lower ratios of intercellular to external CO₂ partial pressure. The latter resulted from large differences in stomatal conductance associated with relatively small differences in rates of photosynthesis. There was little variation between cultivars in response to drought, and differences in allocation pattern and plant water use efficiency between cultivars as found under well-watered conditions persisted under dry conditions. At the end of the non-watered treatment, relative growth rates and transpiration rates decreased to similar values for all cultivars. High ratios of photosynthesis to transpiration, and accordingly high biomass production per unit of transpiration, is regarded as a favourable trait for dry environments, since more efficient use of water postpones the decrease in plant water status.     

Long-term comparative study of soil nitrate test,Gilat plant indicator method and wheat nitrogen uptake

A long-term comparison between two routine soil nitrogen tests, soil nitrate versus plant indicator method, was performed on the Negev Desert loessial soil in Israel. The Gilat plant indicator method was found to be a better method to reflect the soil nitrogen availability for wheat under field conditions. It was found that 15 to 38 kg ha^-1 of NO₃-N, measured by nitrate soil test, for each 30 cm soil increment, is not available for plant uptake. This plant unavailable NO₃-N background in the soil cannot be leached by repeated irrigation cycles of 100 mm each, or by heavy rains.     

Relative growth rate, biomass allocation pattern and water use efficiency of three wheat cultivars during early ontogeny as dependent on water availability

We have investigated the water use efficiency of whole plants and selected leaves and allocation patterns of three wheat cultivars (Mexipak, Nesser and Katya) to explore how variation in these traits can contribute to the ability to grow in dry environments. The cultivars exhibited considerable differences in biomass allocation and water use efficiency. Cultivars with higher growth rates of roots and higher proportions of biomass in roots (Nesser and Katya) also had higher leaf growth rates, higher proportions of their biomass as leaves and higher leaf area ratios. These same cultivars had lower rates of transpiration per unit leaf area or unit root weight and higher biomass production per unit water use. They also had higher ratios of photosynthesis to transpiration, and lower ratios of intercellular to external CO₂ partial pressure. The latter resulted from large differences in stomatal conductance associated with relatively small differences in rates of photosynthesis. There was little variation between cultivars in response to drought, and differences in allocation pattern and plant water use efficiency between cultivars as found under well-watered conditions persisted under dry conditions. At the end of the non-watered treatment, relative growth rates and transpiration rates decreased to similar values for all cultivars. High ratios of photosynthesis to transpiration, and accordingly high biomass production per unit of transpiration, is regarded as a favourable trait for dry environments, since more efficient use of water postpones the decrease in plant water status.     

Nutrition and growth of coniferous seedlings at varied relative nitrogen addition rate

The growth of two provenances of Pinus sylvestris L. were compared with two provenances of Picea abies (L.) Karst. and with Pinus contorta Dougl. when grown in solution cultures with low nutrient concentrations. Nitrogen was added at different exponentially increasing rates, and the other nutrients were added at a rate high enough to ensure free access of them to the seedlings. During an initial period of the culture (a lag phase), when the internal nutrient status was changing from optimum to the level of the treatment, deficiency symptoms appeared. The needles yellowed and the root/shoot ratio increased. The initial phase was followed by a period of exponential growth and steady-state nutrition. The needles turned green again, and the root/shoot ratio stabilized at a level characteristic of the treatment. These patterns were the same as previously reported for other tree species. The relative growth rate during exponential growth was numerically closely equal to the relative nitrogen addition rate. The maximum relative growth rates were about 6 to 7.5% dry weight increase day^-1. This is a much lower maximum than for broad-leaved species (about 20 to 30% day^-1) under similar growth conditions. The internal nitrogen concentrations of the seedlings and the relative growth rates were stable during the exponential period. Close linear relationships were found between these parameters and the relative addition rate up to maximum growth. During steady state the relative growth rates of the different plant parts were equal. However, there were large differences between genotypes in absolute root growth rate at the same seedling size because of differences in root/shoot ratio. Lodgepole pine had the highest root growth rate, whereas that of Norway spruce, especially the southern provenance, was remarkably low. Yet, Norway spruce had a high ability to utilize available nutrients. In treatments with free nutrient access, growth allocation to the shoot had a high priority in all genotypes, but there was still a marked tendency for luxury uptake of nutrients. Nitrogen productivity (growth rate per unit of nitrogen) was lower than in broadleaved species and highest in lodgepole pine. The relevance of the dynamic factors, i.e. maximum relative growth rate, nutrient uptake rate, nitrogen productivity, growth allocation and root growth rate, are discussed with regard to conifer characteristics and selection value.     

Varietal differences in root growth as related to nitrogen uptake by sorghum plants in low-nitrogen environment

Hybrid (Sorghum bicolor (L.) Moench cv CSH 9) and local variety (S. bicolor cv. FSRP) of sorghum were grown in an Alfisol field and in pots of Alfisol in semi-arid tropical India, and the productivity of sorghum in low N condition was studied in relation to N absorption by plant. The grain yield of the hybrid (CSH 9) was higher than that of the local variety (FSRP) irrespective of the rate of N application. The reduction in hybrid grain yield was less than that for the local variety when N was not applied. After booting stage hybrid absorbed more N than the local variety. To estimate the role of roots in N absorption, the characteristics of N absorption were parameterized. Relative growth rate was highly correlated with relative N absorption rate in plant. N absorption was regulated by root activity (specific N absorption rate per unit root mass and length). Specific N absorption rate in roots at physiological maturity was higher in the hybrid than in local variety in low N conditions. Therefore, the smaller reduction of hybrid yield in low-N conditions was ascribed to high-N absorption rate in its roots, and the high ability of roots to absorb N after booting stage.     

The contribution of different regions of the seminal roots of wheat to uptake of nitrate from soil

A technique was developed to determine the physiological activity of defined sections of seminal roots of wheat grown in sand. Wheat plants were grown for 2 weeks in narrow columns of N-deficient sand to which all other nutrients had been added. The columns were split longitudinally and ¹⁵N-labelled nitrate, in an agar medium, supplied to 2 cm sections of root. Shoots and roots were analysed after 24 h to determine the uptake of ¹⁵N. Three sections were examined on either the secondary or tertiary seminal root: 1 cm from the seed (basal segment), 35 cm from the seed (middle segment) and 4 cm from the root apex (apical segment). Total uptake was greatest from the basal and middle segments, declining by 50% from the apical segment. However, uptake per unit root length, including exposed sections of lateral roots, was not significantly different along the root.     

Modelling the uptake of nitrate by a growing plant with an adjustable root nitrate uptake capacity

A new model is presented to predict the plant uptake of nitrate supplied by diffusion and mass flow to its roots. Plant growth, root-shoot ratio and the plant's nitrate uptake capacity are all set dependent on the plant's N nutrition state. By thoroughly integrating processes occurring in both plant and soil, the model enables to control the relative importance of both under a wide range of different nutritional scenarios.Soil parameters D₀ diffusion coefficient in water (m² day^-1) - D_e diffusion coefficient in soil (m² day^-1) - C nitrate concentration in soil (mol m^-3) - f tortuosity (-) - volumetric moisture content (-) - R radial distance from root axis (m) Plant parameters b₁, b₂ parameters of biomass partitioning Equation (10) - IR interroot distance (m) - K_mU Michaelis-Menten constant of the uptake system (mol m^-3) - K_mNRA Michaelis-Menten constant of nitrogen reduction system (mol g^-1) - k₁, k₂, k₃ parameters of growth model Equation (9) - L_v Root length density (m m^-3) - NO_{3 set} ^- Set point of the cytoplasmatic nitrate pool (mol g^-1 dw) - NO_{3 c} ^- cytoplasmatic nitrate concentration (mol g^-1 dw) - NO_{3 v} ^- vacuolar nitrate concentration (mol g^-1 dw) - NRA_max maximum nitrate reductase activity (mol g^-1 dw day^-1) - N_re reduced nitrogen content (mol) - N_remax maximum reduced N concentration in the plant (mol g^-1 dw) - P partitioning coefficient of nitrate between cyplasm and vacuole - R(1) root radius (m) - RGR relative growth rate (day^-1) - U uptake rate (mol day^-1 m^-2) - U_max maximum uptake rate (Eq. 6) (day^-1 m^-2) - Vo water flux at root surface (m day^-1) - W_r root dry weight (g) - W_sh shoot dry weight (g) - X model parameter: number of root compartments - Y model parameter: number of nodes     

Effects of a localized supply of nitrate on NO3 - uptake rate and growth of roots in Lolium multiflorum Lam.

Roots of higher plants are usually exposed to varying spatial and temporal changes in concentrations of soil mineral nitrogen. A split root system was used to see how Lolium multiflorum Lam. roots adapt to such variations to cope with their N requirements. Plants were grown in hydroponic culture with their root system split in two spatially separated compartments allowing them to be fed with or without KNO₃. Net NO₃ ^- uptake, ¹⁵NO₃ ^- influx and root growth were studied in relation to time. Within less than 24 h following deprivation of KNO₃ to half the roots, the influx in NO₃ ^- fed roots was observed to increase (about 200% of the influx measured in plant uniformly NO₃ ^- supplied control plant) thereby compensating the whole plant for the lack of uptake by the N deprived roots. Due to the large NO₃ ^- concentrations in the roots, the NO₃ ^- efflux was also increased so that the net uptake rate increased only slightly (35% maximum) compared with the values obtained for control plants uniformly supplied with NO₃ ^-. This increase in net NO₃ ^- uptake rate was not sufficient to compensate the deficit in N uptake rate of the NO₃ ^- deprived split root in the short term. Over a longer period (>1 wk), root growth of the part of the root system locally supplied with NO₃ ^- was stimulated. An increase in root growth was mainly responsable for the greater uptake of nitrate in Lolium multiflorum so that it was able to fully compensate the deficit in N uptake rate of the NO₃ ^- deprived split root.     

The regulation of nitrogen accumulation in the grain of wheat plants (Triticum aestivum)

Nitrogen accumulation in the ear of wheat plants ( Triticum aestivum L. cv. Klein Chamaco) during ear growth was studied under 4 experimental conditions. Plants were grown in pots with Perlite or soil, and fertilized with nutrient solutions. In one experiment the plants were grown in a greenhouse and supplied with high (16m M ) or low (1.6 m M ) N in the nutrient solutions until anthesis, and then with or without nitrogen supply until ripening. In a second experiment the plants were grown with high N supply until anthesis, and then for half of the plants light intensity was decreased by 50%, and at the same time. N supply was terminated for half of the plants within each light treatment. A third experiment was similar to the previous one, but was carried out in a growth cabinet under 20% of the maximal irradiance in the greenhouse. In a fourth experiment half the ear was excised at anthesis in half of the plants, and these plants were then supplied with or without nitrogen.
In all experiments there was a linear relation between the rate of N accumulation and the rate of ear growth. A wide range of final individual grain weights and N concentration was observed among the experiments. The same maximum N concentration was observed for all grain sizes, although the N concentration could be different between grains of the same size. The grain N concentration correlated with the rate of N accumulation per unit of ear weight increase during ear growth. It is suggested that in wheat plants there is a dependence of nitrogen transport on carbon transport to the ear, and to the ear, and that the final grain N concentration is determined by the N/C ratio exported from the vegetative tissues.