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.     

A shift of paradigm is needed in plant science

The generally accepted norm for controlled experiments with plants has traditionally been restricted to a simplified maintenance of a constant state of the surrounding medium. The ‘treatments’ and a ‘control’, as a reference for comparisons, have been applied without a clear specification of causality and function. However, for pure mathematical reasons, plants are in a constant physiological state (steady-state) only when the internal concentrations of carbon and nutrients and the ratios between them remain constant. Consequently, experimental conditions are controlled in a meaningful sense only when carbon and nutrients are taken up with a constant relative uptake rate, R_u, that is equal to the relative growth rate, R_G. The successful application of these fundamental rules to experimental procedures has been verified in numerous experiments. As for acclimatized plants in steady-state, responses to limiting factors have been graded relative to maximum performance, which constitutes the ultimate reference value. Despite the comprehensive and straightforward material of evidence, the methodology is still a matter of argument. It is concluded that the driving variable commonly used for uptake, external concentration, should be abandoned since its form (lack of time dimension) makes it incompatible with the uptake rate. This calls for a shift of paradigm since the failure of recognition both of fundamental relationships and of experimental shortcomings has been harmful to the development of a sound scientific basis for specification of plant properties. Although fragmented knowledge of basic mechanisms may be good, understanding of causes and effects is poor in relationships between environmental, internal and inherited conditions on the one side, and plant performance on the other. A pseudo-science has in an empirical ‘trial-and-error’ research been based upon statistical evaluation of data of low quality. This is a problem of major significance that needs open discussion and public attention.     

Theories and methods on plant nutrition and growth

A theory comprising two basic concepts relating nutrition and growth is presented. The first concept is a nutrient flux model and is based upon studies of plants at constant internal nutrient concentrations, where a formal mathematical derivation shows that the relative uptake rate (R_U) and the relative growth rate (R_G) are equal. Deviations from equality are results of experimental insufficiencies and errors. The second concept is based on the observation that R_G is linearly related to the internal nutrient concentration. The slope represents nutrient productivity (P_n), an important parameter expressing growth rate per unit of nutrient. Light and the plant genome, for example, influence the value of the proportionality factor, P_n, but not the formal relationship between the internal nutrient concentration and R_G Not only the theory itself but many results and conclusions are very different from those obtained with traditional methods. In experiments where R_U was controlled during the exponential period of growth, the relationships between treatment (the relative addition rate, R_A), nutrient uptake (R_U) and growth (R_G) were reproduced with extremely low variability. In agreement with theory, internal nutrient concentration and R_G remained stable over time (steady-state). An extension of the theory is based upon the empirical assumption that after exponential growth, self-shading and ageing reduce P_n in proportion to biomass. This assumption has been successfully applied in predicting growth of forest stands, but the nature of the growth reduction is little understood. The generalized model has few parameters and can easily be rewritten to suit different experimental aims, for example to establish reference values and to model changes in soil fertility. Further extension and understanding of the model and different interpretations of the parameters are discussed.     

Nutrient uptake and allocation at steady-state nutrition

Ingestad, T. and Ågren, G. I. 1988. Nutrient uptake and allocation at steady-state nutrition. - Physiol. Plant. 72: 450–459. Net nutrient uptake and translocation rates are discussed for conditions of steady-state nutrition and growth. Under these conditions, the relative uptake rate is equal to the relative growth rate, for whole plants as well as for plant parts, since the root/shoot ratio and internal concentrations remain stable. The nutrient productivity and the minimum internal concentration are parameters characteristic for the plant and the nutrient. A conceptual, mathematical model, based on these two fundamental parameters is used for calculation and prediction of the net nutrient uptake rate, which is required to maintain steady-state nutrition at a specified internal nutrient concentration or relative growth rate. When uptake rate is expressed on the basis of the root growth rate, there is, up to optimum, a strong linear relationship between uptake rate and the internal concentration of the limiting nutrient. More complicated and less consistent relationships are obtained when uptake rate is related to root biomass. The limiting factor for suboptimum uptake is the amount of nutrients becoming available at the root surface. When replenishment is efficient, e.g. with vigorous stirring, the concentration requirement at the root surface appears to be extremely low, even at optimum. In the suboptimum range of nutrition, the effect of nutrient status on root growth rate is a critical factor with a strong feed-back on nutrition, growth and allocation. At supraoptimum conditions, the uptake mechanism is interpreted as a protection against too high uptake rates and internal concentrations at high external concentration. In birch (Betula pendula Roth.), the allocation of nitrogen to the shoots is high compared to that of potassium and also to that of phosphorus at low nitrogen or phosphorus status. With decreasing stress, phosphorus allocation becomes more and more similar to nitrogen allocation. The formulation of a mathematical model for calculation of allocation of biomass and nutrients requires more exact information on the quantitative dependence of the growth-regulating processes on nutrition.     

Cultivar differences in the rate of nitrate uptake by intact wheat plants as related to growth rate

Six Argentinian wheat ( Triticum aestivum L.) cultivars grown in nutrient solutions in controlled environment were compared for their nitrate uptake rates on a root dry weight basis. Up to 3-fold differences were observed among the cultivars at 16, 20 and 24 days from germination, either when measured by depletion from the nutrient solution in short-term experiments, or by total N accumulation in the tissue during 8 days.
No differences in total N concentration in root or shoots were found among cultivars. Although the different cultivars showed significant differences in shoot/root ratio and nitrate reductase activity (EC 1.6.6.1) in the roots, none of these parameters was correlated with the nitrate uptake rate. However, nitrate uptake was found to be positively correlated (r = 0.99) with the shoot relative growth rate of the cultivars. The three cultivars with the highest nitrate uptake rates and relative growth rates showed a positive correlation between root nitrate concentration and uptake. However, this correlation was not found in the cultivars with the lowest growth and uptake rates.
Our results indicate that the difference in nitrate uptake rate among these cultivars may only be a consequence of their differences in growth rate, and it is suggested that at least two mechanisms regulate nitrate uptake, one working when plant demand is low and another when plant demand is high.     

Relationships between nitrogen uptake and carbon assimilation in whole plants of tall fescue

Abstract. The present study investigates the relationships between nitrogen uptake, transpiration, and carbon assimilation. Plants growing on nutrient solution were enclosed for 10–16 d in a growth chamber, where temperature, photon flux density, vapour saturation deficit and CO₂ concentration were controlled. One of these factors was modified every 4 to 5 d. Shoot photosynthesis and root and shoot respiration were recorded every half-hour. Nitrogen uptake from the root medium and plant transpiration were measured daily. In most cases, an increase in photon flux density led to increases in transpiration, net daily carbon assimilation, and nitrogen uptake. By modifying transpiration rate without changing photosynthesis (varying vapour saturation deficit), or by modifying transpiration and carbon assimilation in opposite ways (varying CO₂ air concentration), it was shown that nitrogen uptake does not follow transpiration, but is linked to the carbon uptake of the plant. When light was increased from low to intermediate levels, the N uptake/C assimilation ratio remained constant. At higher photon flux density, this ratio declined markedly. It is proposed that in the first case, growth is limited by carbohydrate availability, thus any increase in carbon assimilation leads to a proportional increase in nitrogen uptake, in contrast to the second situation where carbohydrates may accumulate in the plant without further nitrogen requirement.     

Interaction between nitrogen and photon flux density in birch seedlings at steady-state nutrition

Birch ( Betula pendula Roth.) was investigated under steady-state nutrition and growth at different incident photon flux densities (PFD) and different relative addition rates of nitrogen. PFD had a strong influence on the relative growth rate at optimum nutrition and on the nitrogen productivity (growth rate per unit of nitrogen) but little effect on the formal relationships between nitrogen and growth, i.e. PFD and nitrogen nutrition are orthogonal growth factors. At a given suboptimum nitrogen (the same distance from optimum), increased PFD increased the relative growth rate and, therefore, the relative uptake rate and the required relative addition rate in accordance with the theoretical equality between these three parameters at steady-state nutrition. Correspondingly, at a given suboptimum relative addition rate, increased PFD decreased nitrogen status (larger distance from optimum) at an unchanged relative growth rate. Nutrient uptake rate, dry matter content, and partitioning of biomass and nutrients are strongly influenced by nitrogen status. PFD influences these characteristics, but only to an extent corresponding to its effect on the nitrogen status. The influence of PDF on the relative growth rate at optimum and on nitrogen productivity is well described by hyperbolic relationships, similar to reported PFD/photosynthesis relationships. These expressions for plant growth as well as the productivities of leaf area and quantum appear to be valuable characteristics of plant responses to light and nutrition. Although the calculated PFD/growth relationships indicate saturation at high values of PFD, a more realistic estimate of PFD at which saturation occurs is about 30 mol m⁻² day⁻¹, where the highest relative growth rate and nitrogen productivity were experimentally determined. No significant effect was observed because of day length differences between the present and previous experiments.     

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.     

Nutrient requirements and stress response of Populus simonii and Paulownia tomentosa

The aim of this investigation was to estimate the optimum nutrient requirements and responses to low relative nutrient addition rates of seedlings of two important broadleaf tree species in China, Populus simonii Carr. and Paulownia tomentosa (Thunb.) Steud. In preliminary experiments the optimum nutrient proportions were estimated under high concentration conditions. The nutrients consumed were replaced by means of daily additions determined by pH and conductivity titrations without changing the nutrient solutions. A relatively high K level was needed in relation to nitrogen; higher than in birch or grey alder seedlings. To obtain a high relative growth rate, suitable proportions by weight were 100 N:70 K:14 P:7 Ca:7 Mg for the Populus seedlings and 100 N:75 K:20 P:8 Ca:9 Mg for the Paulownia seedlings.
In studies of nutrient stress responses the relative nutrient addition rate was used as the treatment variable under low conductivity conditions. The responses and relationships were similar to those described for birch, grey alder and Salix . The relative addition rate, and there was also a strong linear regression between relative growth rate and nitrogen status. Relative growth rates were high and the maximum weight increase was about 19% day⁻¹ in Populus and over 25% day⁻¹ in Paulownia . The nitrogen productivity of Paulownia was very high, 0.26 g dry weight (g N)⁻¹ h⁻¹, and for Populus it was 0.16 g dry weight (g N)⁻¹ h⁻¹.     

Elevated CO2 induces physiological, biochemical and structural changes in leaves of Arabidopsis thaliana

Leaves of Arabidopsis thaliana grown under elevated or ambient CO2 (700 or 370 micromol mol(-1), respectively) were examined for physiological, biochemical and structural changes. Stomatal characters, carbohydrate and mineral nutrient concentrations, leaf ultrastructure and plant hormone content were investigated using atomic absorption spectrophotometry, transmission electron microscopy and enzyme-linked immunosorbent assay (ELISA). Elevated CO2 reduced the stomatal density and stomatal index of leaves, and also reduced stomatal conductance and transpiration rate. Elevated CO2 increased chloroplast number, width and profile area, and starch grain size and number, but reduced the number of grana thylakoid membranes. Under elevated CO2, the concentrations of carbohydrates and plant hormones, with the exception of abscisic acid, increased whereas mineral nutrient concentrations declined. These results suggest that the changes in chloroplast ultrastructure may primarily be a consequence of increased starch accumulation. Accelerated A. thaliana growth and development in elevated CO2 could in part be attributed to increased foliar concentrations of plant hormones. The reductions in mineral nutrient concentrations may be a result of dilution by increased concentrations of carbohydrates and also of decreases in stomatal conductance and transpiration rate.     

Photosynthetic acclimation to elevated CO2 in Phaseolus vulgaris L. is altered by growth response to nitrogen supply

Abstract Long‐term exposure of plants to elevated CO₂ often leads to downward photosynthetic acclimation. Nitrogen (N) deficiency could potentially exacerbate this response by reducing growth rate and the sink for photosynthates, but this has not always been observed. Experimentally, the interpretation of N effects on CO₂ responses can be confounded by increasing severity of tissue N deficiency over time when N supply is not adjusted as demand increases. In this study, N supply ranged from sub‐ to supra‐optimal (20–540 kgN ha^–l equivalent), and relatively stable levels of tissue N concentration were obtained in all treatments by varying twice‐weekly application rates in proportion to plant growth. The effects of N on photosynthesis and growth of beans (Phaseolus vulgaris L.) raised at ambient (35 Pa) and three elevated (70, 105, 140 Pa) CO₂ partial pressures (pCO₂) were evaluated. Averaging across N treatments, leaf total non‐structural carbohydrates (TNC) were 2.5‐ to 3‐fold higher and leaf N concentrations were 31–35% lower at elevated compared to ambient pCO₂. Light‐saturated net CO₂ assimilation rates measured at growth pCO₂ (A_satg) were significantly higher (26–40% depending on N supply) in plants grown at elevated compared to ambient pCO₂. When measured at a common pCO₂ of 35 Pa, the A_sat of plants grown at elevated CO₂ was 15–29% less than that of plants grown at 35 Pa, indicative of downward photosynthetic acclimation. The magnitude of downward photosynthetic acclimation to elevated CO₂ was greater in plants grown at high (180 and 540 kgN ha^–l) compared to low (20 and 60 kgN ha^–l) N supply, and this was associated with a higher A_sat at growth pCO₂, higher leaf area ratio (leaf area/total biomass), and higher TNC in leaves of high‐N plants. Our results indicate that the effect of N on acclimation to CO₂ will depend on the balance between supply and demand for N during the growing period, and the effect this has on biomass allocation and source‐sink C balance at the whole‐plant level.     

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。     

大气CO2浓度上升和氮添加对南亚热带模拟森林生态系统土壤碳稳定性的影响

大气CO₂浓度升高和氮(N)添加对土壤碳库的影响是当前国际生态学界关注的一个热点。为阐述土壤不同形态有机碳的抗干扰能力, 运用大型开顶箱, 研究了4种处理((1)高CO₂浓度(700 µmol·mol^-1)和高氮添加(100 kg N·hm^-2·a^-1) (CN); (2)高CO₂浓度和背景氮添加(CC); (3)高氮添加和背景CO₂浓度(NN); (4)背景CO₂和背景氮添加(CK))对南亚热带模拟森林生态系统土壤有机碳库稳定性的影响。近5年的试验研究表明: (1) CN处理能明显地促进各土层中土壤总有机碳含量的增加, 其中, 下层土壤(5-60 cm土层)中的响应达到统计学水平。(2)活性有机碳库各组分对处理的响应有所差异: 不同土层中微生物生物量碳(MBC)的含量对各处理的响应趋势基本一致, 各土层中的MBC含量均为CN > CC > NN > CK, 其中0-5 cm、5-10 cm、10-20 cm 3个土层的处理间差异都达到了显著水平; 10-20 cm与20-40 cm两个土层中的易氧化有机碳处理间有显著差异; 而对于各土层中水溶性有机碳, 处理间差异均不明显。(3)各团聚体组分中的有机碳含量的响应也有所差异: 20-40 cm与40-60 cm土层中250-2000 μm组分的有机碳含量存在处理间差异; 40-60 cm土层中53-250 μm组分的有机碳对各处理响应敏感, CC处理和NN处理都有利于该组分碳的深层积累, 尤其CN处理下的效果最为明显; 在各处理10-20 cm、20-40 cm及40-60 cm土壤中, < 53 μm组分中的碳含量间差异显著。大气CO₂浓度上升和N添加促进了森林生态系统中土壤有机碳的增加, 尤其有利于深层土壤中微团聚体与粉粒、黏粒团聚体等较稳定组分中有机碳的积累, 增加了土壤有机碳库的稳定性。     

Respiratory energy requirements of roots vary with the potential growth rate of a plant species

The rates of growth, net rate of nitrate uptake and root respiration of 24 wild species were compared under conditions of optimum nutrient supply. The relative growth rate (RGR)of the roots of these species varied between 110 and 370 mg g^-1 day^-1 and the net rate of nitrate uptake between 1 and 7 mmol (g root dry weight)^-1 day^-1. The rate of root respiration was positively correlated with the RGR of the roots. Root respiration was also calculated from the measured rate of growth and nitrate uptake, using previously determined values for the costs of maintenance, growth and ion uptake of two slow-growing species. The calculated rate of respiration was slightly lower than the measured one for slow-growing species, but twice as high as measured rates for rapid-growing species. This discrepancy was not due to a relatively smaller electron flow through the alternative pathway and, consequently, a more efficient ATP production in the fast-growing species. Neither could variation in specific costs for root growth or maintenance explain these differences. Therefore, we conclude that fast-growing species have lower specific respiratory costs for ion uptake than slow-growing ones. Due partly to these lower specific costs of nutrient uptake, the fraction of respiration that rapid-growing species spend on anion uptake is lower than that of slow-growing species, in spite of the much higher rate of ion uptake of the fast-growing ones.     

氮素添加和CO2浓度升高对白羊草根际和非根际土壤水溶性有机碳、氮的影响

采用盆栽控制试验对黄土丘陵区白羊草在不同CO₂浓度(400和800 μmol·mol^-1)和施氮水平(0、2.5、5.0 g N·m^-2·a^-1)条件下根际和非根际土壤水溶性有机碳(DOC)和水溶性有机氮(DON)的变化特征进行研究.结果表明: CO₂浓度升高对白羊草根际和非根际土壤DOC、水溶性总氮(DTN)、DON、水溶性铵态氮(NH₄⁺-N)、水溶性硝态氮(NO₃^--N)含量均无显著影响.施氮显著提高了根际和非根际土壤DTN、NO₃^--N含量和根际土壤DON含量,显著降低了根际土壤DOC/DON.在各处理条件下,根际土壤DTN、NO₃^--N和DON含量均显著低于非根际土壤,根际土壤DOC/DON显著高于非根际土壤.短期CO₂浓度升高对黄土丘陵区土壤水溶性有机碳、氮含量无显著影响,而氮沉降的增加在一定程度上改善了土壤中水溶性氮素缺乏的状况,但并不足以满足植被对水溶性氮素的需求.     

Control of relative growth rate by application of the relative addition rate technique to a traditional solution culture system

The relative addition rate (RAR) technique allows the nutritional control of plant relative growth rate (RGR) by the provision of nutrients at exponential supply rates. The technique, however, was developed with technologically sophisticated aeroponic systems. In this paper, we report on experiments used to adapt the RAR technique to a conventional solution culture system. A background concentration requirement of 36 μM nitrogen (N), with other nutrients supplied in proportion to N, was necessary to produce a constant RGR of Triticum aestivum L. (wheat) at a low RAR. Solution pH changes were reduced by increasing the percentage of NH₄ in the nitrogen supply, but the plants exhibited dry weight reductions and symptoms of toxicity above 30% NH₄. For wheat, a ratio of 25/75 NH₄/NO₃ was optimum for minimizing pH changes within the nontoxic range. A test of the effectiveness of the RAR technique using this background concentration and NH₄/NO₃ ratio showed that RGR increased with RAR with a linear slope of 0.55 and an intercept of 0.07 d^-1. Although the relationship between growth rate and nutrient supply was less than the one-to-one dependence of RGR on RAR that has been obtained with more sophisticated apparatus, application of the RAR technique to a conventional solution culture system still affords considerable control of RGR and presents a simple method for growing plants at different levels of nutrient stress and at distinct RGRs.     

Acclimation of nitrogen uptake capacity of rice to elevated atmospheric CO2 concentration

Background and Aims: Nitrogen (N) is a major factor affecting yield gain of cropsunder elevated atmospheric carbon dioxide concentrations [CO₂].It is well established that elevated [CO₂] increases root mass,but there are inconsistent reports on the effects on N uptakecapacity per root mass. In the present study, it was hypothesizedthat the responses of N uptake capacity would change with theduration of exposure to elevated [CO₂]. Methods: The hypothesis was tested by measuring N uptake capacity inrice plants exposed to long-term and short-term [CO₂] treatmentsat different growth stages in plants grown under non-limitingN conditions in hydroponic culture. Seasonal changes in photosynthesisrate and transpiration rate were also measured. Key Results: In the long-term [CO₂] study, leaf photosynthetic responsesto intercellular CO₂ concentration (Ci) were not affected byelevated [CO₂] before the heading stage, but the initial slopein this response was decreased by elevated [CO₂] at the grain-fillingstage. Nitrate and ammonium uptake capacities per root dry weightwere not affected by elevated [CO₂] at panicle initiation, butthereafter they were reduced by elevated [CO₂] by 31–41% at the full heading and mid-ripening growth stages. In theshort-term study (24 h exposures), elevated [CO₂] enhanced nitrateand ammonium uptake capacities at the early vegetative growthstage, but elevated [CO₂] decreased the uptake capacities atthe mid-reproductive stage. Conclusions: This study showed that N uptake capacity was downregulated underlong-term exposure to elevated [CO₂] and its response to elevated[CO₂] varied greatly with growth stage.     

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.     

The response of fast- and slow-growing Acacia species to elevated atmospheric CO2: an analysis of the underlying components of relative growth rate

In this study we assessed the impact of elevated CO₂ with unlimited water and complete nutrient on the growth and nitrogen economy of ten woody Acacia species that differ in relative growth rate (RGR). Specifically, we asked whether fast- and slow-growing species systematically differ in their response to elevated CO₂. Four slow-growing species from semi-arid environments (Acacia aneura, A. colei, A. coriacea and A. tetragonophylla) and six fast-growing species from mesic environments (Acacia dealbata, A. implexa, A. mearnsii, A. melanoxylon, A. irrorata and A. saligna) were grown in glasshouses with either ambient (˜350 ppm) or elevated (˜700 ppm) atmospheric CO₂. All species reached greater final plant mass with the exception of A. aneura, and RGR, averaged across all species, increased by 10% over a 12-week period when plants were exposed to elevated CO₂. The stimulation of RGR was evident throughout the 12-week growth period. Elevated CO₂ resulted in less foliage area per unit foliage dry mass, which was mainly the result of an increase in foliage thickness with a smaller contribution from greater dry matter content per unit fresh mass. The net assimilation rate (NAR, increase in plant mass per unit foliage area and time) of the plants grown at elevated CO₂ was higher in all species (on average 30% higher than plants in ambient CO₂) and was responsible for the increase in RGR. The higher NAR was associated with a substantial increase in foliar nitrogen productivity in all ten Acacia species. Plant nitrogen concentration was unaltered by growth at elevated CO₂ for the slow-growing Acacia species, but declined by 10% for faster-growing species. The rate of nitrogen uptake per unit root mass was higher in seven of the species when grown under elevated CO₂, and leaf area per unit root mass was reduced by elevated CO₂ in seven of the species. The absolute increase in RGR due to growth under elevated CO₂ was greater for fast- than for slow-growing Acacia species. Received: 21 December 1998 / Accepted: 31 May 1999