Modelling of the responses to nitrogen availability of two Plantago species grown at a range of exponential nutrient addition rates

Abstract. Plantago major ssp. major and P. lanceolata were grown in solution culture with exponential nutrient addition rates. Compared with P. lanceolata, P. major major showed a higher shoot weight ratio (SWR, fraction of plant dry weight in the shoot) and a higher net assimilation rate (NAR, expressed on a leaf dry weight basis) at equal plant (PNC) and shoot (SNC) nitrogen concentration, respectively. No difference was observed in shoot nitrogen ratio (SNR, fraction of plant nitrogen in the shoot) against PNC between the two species. The effect of these differences in matter partitioning and NAR on plant growth was examined by using a growth model. The model assumed (1) that the SWR and SNR are a linear function of PNC and (2) that the NAR is a rectangular hyperbolic function of SNC. Curvilinear relationships were observed between relative growth rate (RGR) and PNC. P. major major had a higher RGR at equal PNC and, thus, a higher nitrogen productivity (NP) than P. lanceolata. Steady-stale exponential growth was simulated for different nitrogen availability in the environment. P. major major had a higher RGR over the whole range of nitrogen availability but the difference attenuated with decreasing availability of nitrogen. The simulation also showed P. lanccolata having higher plasticity in the shoot/root ratio, which resulted from its higher variability in PNC.     

Differences in nitrate uptake of species from habitats rich or poor in nitrogen when grown at low nitrate concentrations,using a new growth technique

Summary A method is described for culturing plants at extremely low nutrient concentrations. Using a Braun infusion pump, a fixed amount of nitrate or ammonium was supplied continuously to plants growing in a culture vessel at a rate limiting the uptake of the plants. At a very low nitrogen concentration an equilibrium was established where uptake rate of the plants is equal to the rate of supply by the infusion pump. The nitrogen concentrations reached appeared to be in the order of 1 μM. The method compared the nitrate uptake byHypochaeris radicata L.ssp.radicata, H. radicata ssp.ericetorum Van Soest andUrtica dioica L. and ammonium uptake byH. radicata ssp.radicata andH. radicata ssp.ericetorum. Plants were cultivated in monocultures or in mixed cultures (two species per culture vessel). For the mixed cultures competition for nitrate (or ammonium) between the species was maintained for long periods. The capacities of the uptake systems of two subspecies ofH. radicata from places different in nitrogen supply and pH were adapted equally well to both low nitrate and low ammonium concentrations. Apparently factors other than nitrogen uptake play a part in the distribution of the subspecies. The capacity of the uptake system ofU. dioica, a nitrophilous species, was lower than that ofH. radicata ssp.radicata, a species from places poorer in nitrogen. This difference is related to the different distribution of the two species in the field. The present results are compared with those of previous experiments where K_m and V_max were measured and the significance of both parameters is discussed.     

The Effect of Nitrogen Supply to Urtica dioica L. Plants on the Distribution of Assimilate Between Shoot and Roots

In this study the influence of nitrogen nutrition on the patterns of carbon distribution was investigated with Urtica dioica. The nettles were grown in sand culture at 3 levels of NO^?₃, namely 3 (low), 15 (medium) and 22 (high) mM. These levels encompassed a range within which nitrogen did not affect total biomass production. The ratio of root: shoot biomass of the low nitrogen plants was, however, significantly higher than that of the nettles grown at medium and high N supply. Carbon allocation from one leaf of each pair of leaves was examined after a ¹⁴CO₂-pulse and a subsequent ¹⁴C distribution period of one night. Only the youngest two leaf pairs did not export assimilates. Carbon (¹⁴C) export to the shoot apex and to the roots, as measured at the individual nodes responded to the nitrogen status: At medium and high nitrogen supply the 3rd, 4th and 5th leaf pairs exported to the shoot apex, while lower leaves exported to the root. At low nitrogen supply only the 3rd leaf exported towards the shoot apex. The results illustrate the plastic response of carbon distribution patterns to the nitrogen supply, even when net photosynthesis, carbon export from the source leaves and biomass production were not affected by the nitrogen supply to the plant.     

Size and morphology of root systems of perennial grasses from contrasting habitats as affected by nitrogen supply

This paper discusses interspecific differences and phenotypic responses to nitrogen supply in various root parameters of five perennial grasses from contrasting habitats. The following root parameters were studied: root:shoot ratio, specific root length, specific root area, mean root diameter, frequency of fine roots, and the length and density of root hairs. Significant between-species variation was found in all of these features. Species from fertile sites had higher root:shoot ratios at high nitrogen supply than species from infertile habitats. All species growing at low nitrogen supply showed a significant increase in root:shoot ratio. Specific root length, specific root area, mean root diameter and frequency of fine roots were not affected significantly by nitrogen supply. Species from infertile sites responded to low nitrogen supply by a significant increase in root hair length and root hair density.     

Biomass partitioning,architecture and turnover of six herbaceous species from habitats with different nutrient supply

Three grasses (Holcus lanatus, Anthoxanthum odoratum and Festuca ovina) and three herbs (Rumex obtusifolius, Plantago lanceolata and Hieracium pilosella) were grown in a greenhouse at 3 nutrient levels in order to evaluate plant allocation, architecture and biomass turnover in relation to fertility level of their habitats.Four harvests were done at intervals of 4 weeks. Various plant traits related to biomass partitioning, plant architecture, biomass turnover and performance were determined. Differences in nutrient supply induced a strong functional response in the species shoot:root allocation, but architecture and turnover showed little or no response. Architectural parameters like specific leaf area and specific root length, however, in general decreased during plant development.Species from more nutrient-rich successional stages were characterized by a larger specific leaf area and longer specific shoot height (height/shoot biomass), resulting in a higher RGR and total biomass in all nutrient conditions. There was no evidence that species from nutrient-poor environments had a longer specific root length or any other superior growth characteristic. The only advantage displayed by these species was a lower leaf turnover when expressed as the fraction of dead leaves and a shorter specific shoot height (SSH) which might prevent herbivory and mowing losses.The dead leaf fraction, which is a good indicator for biomass and nutrient loss, appeared to be not only determined by the leaf longevity, but was also found to be directly related to the RGR of the species. This new fact might explain the slow relative growth rates in species from a nutrient-poor habitat and should be considered in future discussions about turnover.     

Shoot and root growth and nutrients uptake of wheat as affected by soil layers

The effect of soil layering on the growth and nutrient content of wheat shoots and roots was studied. PVC containers (120 cm long and 25 cm inside diameter) were filled with layers of loam and loamy sand. Both roots and shoots dry weight increased as the thickness of loam layer increased. The root:shoot ratios decreased throughout the growing season. The N, P and K content of the shoots peaked at two weeks before anthesis, while shoot dry weight peaked at anthesis. The ranges of shoot content of N, P and K at anthesis for the different treatments were 6–25, 8–25 and 5–25% of the total plant nutrients, respectively. Late in the season the translocation rate of nutrients from the shoots to the seeds were in the following order N>P>K.     

Simulation study of nutrient uptake by plants from soilless cultures as affected by salinity buildup and transpiration

Soilless plant growth systems are widely used as a means to save irrigation water and to reduce groundwater contamination. While nutrient concentrations in the growth medium are depleted due to uptake by the plants, salinity and toxic substances accumulate due to transpiration. A theoretical model is suggested, to simulate nutrient uptake by plants grown in soilless cultures with recycled solutions. The model accounts for salinity accumulation with time and plant growth, and its effects on uptake of the different nutrients by means of interaction with Na and Cl ions. The sink term occurs due to uptake by a growing root system. Influx as a function of the ion concentration is according to Michaelis–Menten active mechanisms for K⁺, NO₃ ^–-N, NH₄ ⁺-N, PO₄-P, Ca²⁺, Mg²⁺ and SO₄ ^2-, whose influx parameters are affected by Na and Cl^–, but not with time (age). Sodium influx is passive above a critical concentration. Sum of cations–anions concentrations is balanced by Cl^– to maintain electro-neutrality of the growth solution. Salinity (by means of Na concentration) suppresses root and leaf growth, which further effect uptake and transpiration. The model accounts for instantaneous transpiration losses, during daytime only and its effect on uptake of nutrients and plant development due to salt accumulation. The model was tested against NO₃ ^– and K⁺ uptake by plants associated with cumulative transpiration and with different NaCl salinity levels. Deviations from observed K⁺ uptake should be attributed to the salinity tolerance of the plants. In a study with data obtained from published literature, the model indicated that nutrient depletion and salinity buildup might be completely different with fully grown-up plants (that do not grow) and plants that grow with time. Depletion of different nutrients are according to their initial concentration and plant uptake rate, but also affected by their interactions with Na and Cl ions.     

Nitrate reductase activity and uptake of nitrate and oxygen as affected by nitrate supply to detached maize organs

Supply of 1, 2, 5, 10 or 20 mM nitrate to detached roots, scutella or shoots from 5- to 6-d-old Zea mays L. seedlings increased in vitro nitrate reductase (NR) activity in all the organs and NADPH specific NR (NADPH:NR) activity in roots and scutella but not in the shoots. Usually 2 to 5 mM nitrate supported maximum enzyme activity, the higher concentration did not increase it further. The protein content in the roots, scutella and shoots increased up to 5, 2 and 20 mM medium nitrate, respectively. Nitrate uptake also increased with increasing nitrate concentration in roots and shoots, but it increased only slightly in the scutella. In both roots and scutella, methionine sulfoximine had no effect, while cycloheximide and tungstate abolished nitrate induced NADH:NR activity completely and NADPH:NR partially. Methionine sulfoximine increased nitrate uptake by roots and scutella slightly, but other inhibitors had no effect. The depletion of dissolved oxygen from the medium was lower in the presence of nitrate than in its absence or in the presence of ammonium, especially in the scutella. This revised version was published online in July 2006 with corrections to the Cover Date.     

Root and leaf attributes accounting for the performance of fast- and slow-growing grasses at different nutrient supply

Despite their difference in potential growth rate, the slow-growing Brachypodium pinnatum and the fast-growing Dactylis glomerata co-occur in many nutrient-poor calcareous grasslands. They are known to respond differently to increasing levels of N and P. An experiment was designed to measure which characteristics are affected by nutrient supply and contribute to the ecological performance of these species. Nutrient acquisition and root and shoot traits of these grasses were studied in a garden experiment with nine nutrient treatments in a factorial design of 3 N and 3 P levels each. D. glomerata was superior to B. pinnatum in nutrient acquisition and growth in all treatments. B. pinnatum was especially poor in P acquisition. Both species responded to increasing N supply and to a lesser extent to increasing P supply by decreasing their root length and increasing their leaf area per total plant weight. D. glomerata showed a higher plasticity. In most treatments, the root length ratio (RLR) and the leaf area ratio (LAR) were higher for D. glomerata. A factorization of these parameters into components expressing biomass allocation, form (root fineness or leaf thickness) and density (dry matter content) shows that the low density of the biomass of D. glomerata was the main cause for the higher RLR and LAR. The biomass allocation to the roots showed a considerable plasticity but did not differ between the species. B. pinnatum had the highest leaf weight ratio. Root fineness was highly plastic in D. glomerata, the difference with B. pinnatum being mainly due to the thick roots of D. glomerata at high nutrient supply. The leaf area/leaf fresh weight ratio did not show any plasticity and was slightly higher for B. pinnatum. It is concluded, that the low density of the biomass of D. glomerata is the pivotal trait responsible for its faster growth at all nutrient levels. It enables simultaneously a good nutrient acquisition capacity by the roots as well as a superior carbon acquisition by the leaves. The high biomass density of B. pinnatum will then result in a lower nutrient requirement due to a slower turnover, which in the long term is advantageous under nutrient-poor conditions.     

Regulation of shoot/root ratio by cytokinins from roots inUrtica dioica: Opinion

According to current knowledge, cytokinins are predominantly root-born phytohormones which are transported into the shoot by the transpiration stream. In the hormone message concept they are considered the root signals, which mediate the flux of the photosynthates to the various sinks of the plant. In this review, experiments are assessed, in which changes of the shoot to root ratio of biomass, caused by different levels of nitrogen supply to a model plant,Urtica dioica, could be traced to the natural cytokinin relations of the plant. Disturbance of the internal cytokinin balance of the plant resulted in a disproportionate distribution of the assimilates in favour of the cytokinin-enriched shoot. Inspite of some shortcomings of the hormone message concept, the presented work corroborates the significance of root-sourced cytokinins in the regulation of biomass partitioning between shoot and root.     

Photosynthesis and growth of Erythrina variegata as affected by water stress and triacontanol

Erythrina variegata Lam. seedlings were grown under water stress (Ψ = -3.2 MPa) and subsequently sprayed with triacontanol (Tria). Water stress significantly reduced shoot growth rate, while roots continued to grow. Content of chlorophyll (Chl) a decreased more than that of Chl b. Water stress also reduced photosynthetic activity of chloroplasts as measured by Chl fluorescence induction. Stress effect was identified at the oxidation site of photosystem (PS) 2 prior to the hydroxylamine donating site and perhaps close to or after the diphenylcarbazide donating site. The loss of O₂ evolving thylakoid polypeptides (33, 23, 17 kDa) and the large (55 kDa) and small (15 kDa) subunits of ribulose-1,5-bisphosphate carboxylase (RuBPC) were found in water stressed seedlings. The reduction in RuBPC activity was accompanied by reduction of CO₂ fixation and stomatal conductance. All photosynthetic parameters were improved by Tria. This revised version was published online in August 2006 with corrections to the Cover Date.     

Closely related allopatric Podalyria species from the Core Cape Subregion differ in their mechanisms for acquisition of phosphorus,growth and ecological niche

Aims In the Core Cape Subregion (CCR), a Mediterranean-climate ecosystem with infertile soils, the legume species Podalyria calyptrata and P. burchellii are in a separate clade to P. leipoldtii and P. myrtillifolia. The closely related species are allopatric, and with the west-east climate gradient and variation in soil nutrient availability in the CCR, it was hypothesized that the two closely related allopatric species would differ in their ecological niche and root:shoot ratio, specific root length (SRL) and organic acid exudation responses to phosphorus (P) supply.Methods With increasing P supply in the glasshouse, we measured plant biomass, leaf nitrogen ([N]), [P], root morphology and release of organic acids. We determined species soil and leaf [N] and [P] and climate in field sites.Important findings At low P supply, P. calyptrata roots exuded more organic acids than P. burchellii which instead produced roots with a greater SRL, and P. myrtillifolia allocated more biomass to roots than P. leipoldtii. In the field, leaf [P] and climate suggested that P. leipoldtii occupied the most oligotrophic niche followed by P. burchellii and then P. calyptrata and P. myrtillifolia. Closely related allopatric species differed in their mechanisms for P-acquisition and ecological niche, indicating that the environment overrides phylogeny in determining P-acquisition traits for these species, and suggesting that climate regulates nutrient availability, driving distribution and speciation.     

青藏高原三种优势植物生物量分配的变化规律

该研究利用4个由高到低不同海拔的同质园实验,以青藏高原高寒草地优势植物垂穗披碱草(Elymus nutans)、矮嵩草(Kobresia humilis)和珠芽蓼(Polygonum viviparum)为对象,分析了植物个体根、茎、叶生物量分配及根冠比的变化规律及影响因素。结果表明:(1)植物个体根、茎、叶质量比和根冠比具有显著的种间差异;与垂穗披碱草和珠芽蓼相比,矮嵩草具有显著较高的根质量比而叶、茎质量比较低,所以其根冠比较高。(2)在向低海拔移栽的过程中,珠芽蓼叶质量比保持不变,茎质量比显著降低而根质量比显著升高,根冠比表现出显著上升的趋势;垂穗披碱草则相反,即叶、茎质量比显著升高而根质量比显著降低,根冠比表现出显著下降的趋势;矮嵩草根、茎、叶质量比和根冠比则无显著变化。(3)随着海拔降低,年均气温明显升高而年均降雨量明显降低,且在植物个体种源地和土壤基质保持一致的条件下,向低海拔移栽过程中温度是导致珠芽蓼根、茎、叶生物量分配及根冠比变化的重要因素,而水分是垂穗披碱草根、茎、叶生物量分配及根冠比变化的重要驱动因素;矮嵩草根、茎、叶生物量分配及根冠比受其遗传因素影响较大。因此,在将来暖干化的背景下,青藏高原高寒草地植物生物量的分配将会发生改变,导致它们对资源(光照、水分和土壤养分)获取和利用的变化而改变它们的种间关系,从而影响群落的物种多样性与组成,最终可能导致生态系统功能的变化。     

Development of Frost Tolerance in Winter Wheat as Modulated by Differential Root and Shoot Temperature

Abstract: Winter wheat plants ( Triticum aestivum L. cv. Urban), grown in nutrient solution, were exposed to differential shoot/root temperatures (i.e., 4/4, 4/20, 20/4 and 20/20 C) for six weeks. Leaves grown at 4C showed an increase in frost tolerance from - 4C down to - 11 C, irrespective of root temperature. In 4/20 C plants, high root temperature decreased the rate of hardening of the leaves, but did not influence the final level of frost tolerance. In roots grown at 4C frost tolerance increased from - 3 C down to - 4 C, independently of shoot temperature. An accumulation of soluble sugars in the leaves was only observed when both shoot and root were grown at 4C and was not correlated with final frost tolerance achieved. However, the rate of hardening was correlated with the soluble sugar concentration. An increase in root soluble sugar concentration was exclusively observed in roots exposed to a temperature of 4C, irrespective of shoot temperature. Proline concentration only increased in plant parts exposed to a temperature of 4C. The present results indicate that the importance of root temperature in low-temperature hardening of winter wheat is limited, even though exposure to differential root and shoot temperatures brought about pronounced changes in growth, soluble sugar concentration, insoluble sugar concentration and proline concentration in roots and leaves.     

Growth and the efficiency of root respiration of Pisum sativum as dependent on the source of nitrogen

Growth and efficiency of root respiration were investigated in Pisum sativum L. cv. Alaska and cv. Rondo. Plants were grown in culture solutions, either in symbiosis with Rhizobium leguminosanm , or with an abundant supply of nitrate or ammonium and completely lacking nodules. In comparison with plants utilizing nitrate or ammonium, Ni-fixing plants showed lower rates of dry matter and nitrogen accumulation, as well as lower rates of total and cytochrome-mediated root respiration. Rates of shoot dry matter accumulation and root respiration in plants utilizing ammonium were lower than in plants utilizing nitrate. The efficiency of root respiration was high in N₂-fixing plants, as indicated by a low activity of the SHAM-sensitive, alternative, non-phosphorylating pathway. In nitrate and ammonium grown plants of cv. Alaska, the efficiency of root respiration was about the same, and in both cases lower than in N₂-fixing plants. The efficiency of root respiration in non-symbiotically grown pea plants was generally higher than in many non-legumes. Comparison of the ATP costs of synthesis of root dry matter for different N-sources was complicated by large differences in relative growth rate of the root and in shoot to root ratio between N-treatments. A quantitative correction of the ATP production during synthesis of root dry matter for differences in shoot to root ratio and root maintenance respiration has been made. It is concluded that ATP costs of root dry matter production are highest in the case of N₂-fixing plants. In plants utilizing ammonium, ATP costs of synthesis of root dry matter were slightly lower than in plants utilizing nitrate. The physiological significance of the alternative pathway in root metabolism is discussed in relation to the assimilation of different sources of nitrogen.     

The effects of temperature and nutrient concentrations on nitrate and phosphate uptake in different species of Porphyra from Long Island Sound (USA)

Uptake rates of nitrate and phosphate were measured for four species and one variety of Porphyra from Long Island Sound (USA) at two temperatures and two nutrient medium concentrations at increasing intervals over a 24- or 48-h period. Maximum uptake rates found were: V_{30 μM}^{0-1 h}=73.8 μmol NO₃ g⁻¹ DW h⁻¹ and V_{3 μM}^{0-1 h}=16.7 μmol PO₄ g⁻¹ DW h⁻¹, in the two thinnest Porphyra. We found that the nitrate uptake rates were significantly greater at 30 μM than 3 μM NO₃ concentration, and that the uptake rates decreased with time of exposure. Temperature (5, 15, and 25 °C) did not have as strong an effect on nitrate uptake rates as did nutrient concentration. Q₁₀ values and uptake rates at four different nitrate concentrations indicated that nutrient uptake at 5 °C was initially an active process. After 24 h, the processes involved appeared passive as Q₁₀ values were between 1.0 and 1.3 and nitrate uptake curves were linear. Nitrate uptake rates correlated positively with the surface area/volume (SA/V) ratio. No coherent trends were found for uptake of phosphate, except that the uptake rates were significantly higher in 30 μM NO₃ medium as opposed to 3 μM NO₃. We did not find any significant difference in uptake rate and pattern between the summer species Porphyra purpurea (Roth.) C. Agardh, the eurythermic Porphyra suborbiculata Kjellm., the winter species Porphyra rosengurttii J. Coll and J. Cox, and the two varieties of Porphyra leucosticta Thur. Le Jol. (both winter species).     

Oxygen uptake by roots of Rumex species at different temperatures: the relative importance of diffusive resistance and enzyme kinetics

Abstract. Oxygen uptake characteristics of the roots of three Rumex species were compared, and related to kinetics of the respiratory system and to root anatomy. The observed differences could not be explained by differences in fundamental characteristics of the oxygen uptake system: with all three species, cytochrome-mediated respiration contributed 70% and cyanide-insensitive (alternative) respiration 30% of the total respiration rate, and apparent K_m values of cytochrome oxidase were lower than those obtained for the alternative oxidase in all cases. However, differences in critical oxygen pressure for respiration (COPR) and in apparent K_m for oxygen, were strongly correlated with differences in root porosity and root diameter. K_m(O₂) values at high and low temperatures were determined, and from Arrhenius plots of oxygen uptake rates between 11 and 32°C, the role of diffusional impedance could be estimated. Root respiration of Rumex maritimus and R. crispus , both with high root porosity, but differing in root diameter, had a low K_m for oxygen (3–7 mmol m⁻³). In contrast with this were the responses of R. thvrsiflorus , which has thin roots but low root porosity: a high K_m (10-20 mmol m⁻³) was found at all temperatures. The role of diffusional impedance as a function of temperature in oxygen uptake rate by the three species is discussed and related to the differential resistance of the species towards flooding.     

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.