Ecophysiological traits explain species dominance patterns in macroalgal blooms

Macroalgal blooms are a growing environmental problem in eutrophic coastal ecosystems world wide. These blooms are dominated typically by only one out of several co-occurring opportunistic species, which are all favored by increased nutrient loads. We asked whether pronounced dominance of filamentous Pilayella littoralis Kjellm. (Phaeophyceae) over foliose Enteromorpha intestinalis L. (Chlorophyceae) in the Baltic Sea can be explained by interspecific physiological differences. In laboratory experiments, we analyzed uptake kinetics of nitrate, ammonium, and phosphate and the time dependency of uptake rates for both species. We further examined growth rates and nutrient assimilation in relation to single and combined enrichment with nitrate and phosphate, and three different nitrogen sources. Overall, we did not detect distinct differences in uptake, growth, and assimilation rates between P. littoralis and E. intestinalis. Minor differences and the related advantages for single species are discussed. Highest maximal uptake rates were found for ammonium, followed by nitrate and phosphate. Strong time dependency of uptake occurred, with the highest rates during the first 15 to 30 min. Nitrate enrichment had far more of an effect on growth than phosphate. Enrichment with urea, ammonium, and nitrate significantly increased growth rates without interspecific differences. A larger surface area to volume (SA/V) ratio in Pilayella compared with Enteromorpha did not translate into greater physiological capacity. We conclude that species dominance patterns in macroalgal blooms are not always a direct result of different ecophysiological traits among species. Ecological traits such as susceptibility to herbivory are important factors in determining species distribution in the field.     

Mineral Nutrient Requirements of Pinus silvestris and Picea abies Seedlings

The mineral nutrient requirements of Pinus silvestris L. and Picea abies Karst. were studied according to previously published methods applied to a series of various plant species. The optimum nutrient proportions are similar to those of Vaccinium, with a lower relative potassium requirement than birch and other broad-leaf species. Various ratios between ammonium and nitrate nitrogen were about equally efficient except for a minor growth reduction with pure nitrate, which gave a comparatively low nitrogen content and a high cation/nitrogen ratio. The rate of ammonium uptake was much higher than that of nitrate when both sources were supplied. The required total concentration in the nutrient solution for maximum growth is lower in pine than in spruce, but both fall within the low salt range. Both species, especially pine, are sensitive to high salt concentrations. Although pine and spruce grow on the same type of soils as Vaccinium— leached soils with low base saturation – accumulation of calcium or other cations is not as pronounced as in Vaccinium, especially not in pine. The results are compared with results from similar experiments with a series of other conifers. All the conifers have more flexible cation uptake mechanisms than Vaccinium but the results indicate tendencies to accumulation of anions, nitrate and phosphate. Recommended fertilizer compositions for forest fertilization and nurseries are discussed.     

Relationship of ion absorption to growth rate in taiga trees

Summary Rates of nutrient absorption were measured on excised roots of taiga tree seedlings grown in the laboratory. Phosphate and to a lesser extent ammonium (relatively immobile ions in the soil) were absorbed most rapidly by poplar and aspen, two species with rapid growth rates and most slowly by alder and/or black spruce, species with slow growth rates. In contrast, potassium (which is more mobile in soil) was absorbed most rapidly by slowly growing species. All species had low rates of nitrate and chloride absorption. Absorption rate of each ion was most temperature sensitive in those species that typically occupy the warmest soils (i.e. poplar and aspen). We suggest that in infertile soils a high capacity for uptake is an important component of root competition only in the case of mobile ions (e.g. potassium, nitrate), because only for these ions do diffusion shells of adjacent roots overlap; in contrast plants compete for immobile ions (e.g. phosphate) only by increasing absorptive surface via root growth or mycorrhizal association.     

Mineral Nutrient Requirements of Vaccinium vitis idaea and V. myrtillus

The mineral nutrient requirements of Vaccinium vitis idaea L. and V. myrtillus L. have been studied. It is found that the optimum nutrient proportions are similar to those of birch seedlings except for a relatively low potassium requirement. The nitrogen sources ammonium and nitrate are both usable at low concentrations. A system with both sources present seems preferable. The ammonium uptake is then much more rapid than the nitrate uptake. The optimum total concentration falls within the low salt range and is about the same as found for birch seedlings, but the sensitivity to high salt concentrations is more pronounced in the Vaccinium species. A very marked characteristic of the nutrient absorption is the rapid calcium uptake. The results are discussed in relation to similar experiments with birch and cucumber seedlings, representing species with a wide or calcareous ecological adaptation.     

Nutrient uptake and growth responses of three intertidal macroalgae with perennial, opportunistic and summer-annual strategies

An important life history trait of macroalgae species is the physiological ability to cope with nutrient limiting conditions, which seasonally occur in temperate coasts while other environmental factors are adequate (e.g., sufficient light). Nitrogen (N) and Phosphorus (P) uptake kinetics and field growth limitation were investigated in the perennial Bifucaria bifurcata, the opportunistic Ulva intestinalis, and the summer-annual Nemalion helminthoides from Asturian coasts (N Spain). We performed 4 nutrient uptake experiments (ammonium, nitrate, nitrate + ammonium, and phosphate) and monitored the growth and N content of field individuals in the presence/absence of artificial nutrient supply to assess potential growth limitations. B. bifurcata was not actively growing during summer thus low nutrient demands probably relied on stored pools and/or the low background nutrient levels in seawater, as generally observed for perennials. Corresponding N content and uptake rates in this species were the lowest. The opportunistic U. intestinalis showed kinetics suitable for assimilating N quickly at high external concentrations in order to fulfill the high nutrient demands that support its fast-growing strategy. This response is well adapted to seasons and sites of high nutrient loading but signs of nutrient starvation during summer (decreasing growth and N content) were found in the pristine studied area. N. helminthoides showed an intermediate response in terms of thallus N content and uptake affinity, together with an inducible activation of nitrate uptake. This response assured the uptake of transient nutrient pulses without the nutrient storage response of perennials, or the costly enzymatic machinery of opportunistics. This allows N. helminthoides to effectively exploit low background nutrient conditions interrupted by transient peaks during spring-summer, when most ephemerals found difficulties to survive and perennials suspend their active growth. P uptake did not differ greatly between species suggesting its secondary importance compared to N in the tested algae.     

The effect of constant moisture and aeration levels in soil on Fusarium wilt of tomatoes

Soil patches on opposite sides of Pseudoroegneria spicata plants in the field were treated with either distilled water or a nutrient solution containing N, P, or K. Roots from these enriched and control patches were tested three days later for their capacities of ammonium, phosphate, and potassium uptake. When phosphate was augmented in the enriched patches, rates of phosphate uptake increased significantly, but not rates of ammonium or potassium uptake. When the enriched patches were augmented with nitrogen, uptake capacities of both ammonium and potassium increased significantly (mean increases of up to 88% and 71% for ammonium and potassium, respectively). Potassium augmentation did not lead to increased soil-available K and, correspondingly, did not induce changes in the capacity for uptake of K, N, or P. The potential importance of nutrient uptake kinetics in the exploitation of nutrient-rich soil patches is discussed.     

Control of Net Uptake of Nutrients by Regulation of Influx in Barley Plants Recovering from Nutrient Deficiency

Rates of influx and net uptake of nitrate, phosphate and sulphatewere measured in intact barley plants, and concurrent effluxwas obtained by difference. Net uptake of these anions variedwidely depending on the nutrient status of the plants, and thedifferences in net uptake could be accounted for almost entirelyby changes in influx. Efflux played only a minor role in regulatingnet uptake of nitrate, phosphate or sulphate during recoveryfrom N-, P-, or S-deficiency. Nitrate influx and short-term ammonium absorption by N-deficientbarley plants were closely correlated, and varied in parallelwith rates of net uptake of nitrate or ammonium by similar plants.Again, it would seem that net uptake of ammonium is controlledpredominantly by changes in the rate of influx.Copyright 1993,1999 Academic Press Hordeum vulgare, barley, nutrient absorption, influx, nitrate, phosphate, sulphate, ammonium     

NITRATE AND PHOSPHATE UPTAKE INTERACTIONS IN A MARINE PRYMNESIOPHYTE1

The short- and long-term uptake of nitrate and phosphate ions, and their interactions, were studied as functions of the preconditioning of Pavlova lutheri (Droop) Green. Populations were preconditioned in continuous culture at a variety of growth rates and N:P supply ratios. The maximum uptake rates cell^?1 for nitrate and phosphate were of similar magnitudes, in spite of the forty-fold smaller requirement for phosphorus. Short-term phosphate uptake was independent of the nitrate concentration, but the short-term nitrate uptake rate was reduced in the presence of phosphate. The severity of inhibition of nitrate uptake by phosphate was positively correlated with the preconditioning N:P supply ratio and the preconditioning growth rate. In response to large additions of nutrients, P. lutheri was able to increase its phosphorus content sixty-fold, but was only able to take up enough nitrate to double its nitrogen content. The high rate of phosphate uptake relative to its requirement, the inhibition of nitrate uptake by phosphate, and the large capacity for phosphorus storage relative to its requirement, all of which were observed even under N limitation, may imply that even where nitrogen is limiting there can be interspecific competition for available phosphate.     

Effects of Atrazine,Metolachlor, Carbaryl and Chlorothalonil on Benthic Microbes and Their Nutrient Dynamics

Atrazine, metolachlor, carbaryl, and chlorothalonil are detected in streams throughout the U.S. at concentrations that may have adverse effects on benthic microbes. Sediment samples were exposed to these pesticides to quantify responses of ammonium, nitrate, and phosphate uptake by the benthic microbial community. Control uptake rates of sediments had net remineralization of nitrate (−1.58 NO₃ µg gdm⁻¹ h⁻¹), and net assimilation of phosphate (1.34 PO₄ µg gdm⁻¹ h⁻¹) and ammonium (0.03 NH₄ µg gdm⁻¹ h⁻¹). Metolachlor decreased ammonium and phosphate uptake. Chlorothalonil decreased nitrate remineralization and phosphate uptake. Nitrate, ammonium, and phosphate uptake rates are more pronounced in the presence of these pesticides due to microbial adaptations to toxicants. Our interpretation of pesticide availability based on their water/solid affinities supports no effects for atrazine and carbaryl, decreasing nitrate remineralization, and phosphate assimilation in response to chlorothalonil. Further, decreased ammonium and phosphate uptake in response to metolachlor is likely due to affinity. Because atrazine target autotrophs, and carbaryl synaptic activity, effects on benthic microbes were not hypothesized, consistent with results. Metolachlor and chlorothalonil (non-specific modes of action) had significant effects on sediment microbial nutrient dynamics. Thus, pesticides with a higher affinity to sediments and/or broad modes of action are likely to affect sediment microbes'' nutrient dynamics than pesticides dissolved in water or specific modes of action. Predicted nutrient uptake rates were calculated at mean and peak concentrations of metolachlor and chlorothalonil in freshwaters using polynomial equations generated in this experiment. We concluded that in natural ecosystems, peak chlorothalonil and metolachlor concentrations could affect phosphate and ammonium by decreasing net assimilation, and nitrate uptake rates by decreasing remineralization, relative to mean concentrations of metolachlor and chlorothalonil. Our regression equations can complement models of nitrogen and phosphorus availability in streams to predict potential changes in nutrient dynamics in response to pesticides in freshwaters.     

Nitrogen acquisition in four co-existing species from an upland acidic grassland

The concentration of both nitrate and ammonium nitrogen was measured in soil taken from an upland acidic (pH 4.5) grassland habitat, containing four co-existing species, Deschampsia flexuosa (L.) Trin., Festuca ovina L., Juncus squarrosus L. and Nardus stricta L. Both nitrate and ammonium nitrogen were found to be present in the soil, in similarly small quantities. The effect of both sources of nitrogen on relative growth rate was studied, and an attempt was made to determine whether nitrate or ammonium nitrogen is the immediate source of nitrogen for these plants using assays of nitrate reductase (EC 1.6.6.2) and ammonium uptake. All four species showed larger growth rates on the same concentration of ammonium nitrogen compared to nitrate nitrogen. All species showed low activities of leaf nitrate reductase, even in plants grown on 18 mol nitrate m⁻³. Ammonium uptake activity appeared to be higher in species which showed the lowest nitrate reductase activity and least response to increasing nitrate concentration in the growth medium.     

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.     

Solubilization of rock phosphate by rape

Rape and sunflower were compared with respect to their rock phosphate mobilizing capacities, cationanion balance and uptake of Ca and NO₃ at P-starvation. Rape was able to mobilize P from rock phosphate, whereas sunflower was not.When grown on a complete nutrient solution with NO₃ as the only nitrogen source, both species took up more nutrient anions than cations. Withholding phosphate from the nutrient solution did not change the uptake pattern of rape, but sunflower took up more nutrient cations than anions at P-starvation, due to a strong decline in NO₃ uptake. With both species, Ca uptake was not affected by phosphate in the nutrient solution.In split pot experiments, with rock phosphate supplied spatially separated from other nutrients, rape was still able to mobilize rock phosphate. A high Ca concentration had no effect on this capacity.The results indicated that in our experiments rock phosphate mobilization by rape was not associated with an excess of cation over anion uptake and neither with a high Ca uptake rate.     

Physiological responses and biofilter potential of <Emphasis Type="Italic">Hypnea aspera</Emphasis> (Rhodophyta,Gigartinales) cultivated in different availabilities of nitrate,ammonium, and phosphate

Along with the search for new species of seaweeds with biofilter capacity, it is also necessary to understand the physiological and biochemical responses of these seaweeds cultivated in different availabilities of nitrate, ammonium, and phosphate. To accomplish this, a laboratory study was performed to evaluate the ability of Hypnea aspera Kützing (Gigartinales, Rhodophyta), to growth under different nitrate, ammonium, and phosphate availabilities and to uptake, assimilate, and remove these nutrients from seawater. Treatments were composed of sterilized seawater enriched with quarter-strength von Stosch’s nutrient solution modified (without any nitrogen and phosphorus compounds). Nitrate or ammonium, together with phosphate, was added in combined N/P ratios of 100:1 and 10:1. Nitrate concentrations varied from 0 to 150 μM, and ammonium concentrations varied from 0 to 70 μM. Growth rates of H. aspera increased with nitrate addition, and the highest value was 4.68 ± 0.76 % day^?1 in 150 μM, but the addition of high ammonium concentration (70 μM) in N/P ratio of 10:1 inhibited the growth rates (?3.89 ± 1.03 % day^?1). Nitrogen was accumulated as proteins and phycobiliproteins, mainly phycoerythrin, at higher phosphate availability (N/P ratio of 10:1) for nitrate addition (150 μM for proteins and 50, 100, and 150 μM for phycoerythrin), and lower phosphate availability (N/P ratio of 100:1) for ammonium addition (50 and 70 μM for proteins and 50 μM for phycoerythrin). Nitrogen and phosphate were assimilated into thallus in all treatments with nutrient addition. Hypnea aspera showed high removal efficiency (higher than 90 %) of nitrate, nitrite, ammonium, and phosphate present in the seawater. These results suggest that H. aspera could be cultivated in integrated multitrophic aquaculture systems to reduce nutrient loading in eutrophic seawater.     

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.     

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.     

Measurement of short-term nutrient uptake rates in cranberry by aeroponics

Aeroponics, a soil-less plant culture system in which fresh nutrient solutions are intermittently or continuously misted on to plant roots, is capable of sustaining plant growth for extended periods of time while maintaining a constantly refreshed nutrient solution. Although used relatively extensively in commercial installations and in root physiology research, use of aeroponics in nutrient studies is rare. The object of this study was to examine whether nutrient uptake rates could be calculated for aeroponic systems by difference using measurements of concentrations and volumes of input and efflux solutions. Data were collected from an experiment with cranberry plants (Vaccinium macrocarpon Ait. cv. Stevens) cultured aeroponically with nutrient solutions containing various concentrations of ammonium-N and isotopically labelled nitrate-N. Validation of the calculated uptake rates was sought by: (1) evaluating charge balance of the solutions and total ion uptake (including proton efflux) and (2) comparison with N-isotope measurements. Charge balance and proton efflux calculations required use of chemical modelling of the solutions to determine speciation of dissolved phosphate and acid-neutralizing capacity (ANC). The results show that charge balance requirements were acceptably satisfied for individual solution analyses and for total ion uptake when proton efflux was included. Relative rates of nitrate/ammonium uptake determined by difference were in agreement with those determined by isotopic techniques. Additional information was easily obtained from this experimental technique, including evidence of diurnal variation in nutrient uptake, correlation between ammonium uptake and proton efflux, and the relationship between ion concentration and uptake. Use of aeroponic systems for non-destructive measurement of water and ion uptake rates for numerous other species and nutrients appears promising.     

Partitioning of inorganic nitrogen assimilation between the roots and shoots of cerrado and forest trees of contrasting plant communities of South East Brasil

Summary Woody plants growing in cerrado and forest communities of south-east Brasil were found to have low levels of nitrate reductase activity in their leaves suggesting that nitrate ions are not an important nitrogen source in these communities. Only in the leaves of species growing in areas of disturbance, such as gaps and forest margins, were high levels of nitrate reductase present. When pot-grown plants were supplied with nitrate, leaves and roots of almost all species responded by inducing increased levels of nitrate reductase. Pioneer or colonizing species exhibited highest levels of nitrate reductase and high shoot: root nitrate reductase activities. Glutamine synthetase, glutamate synthase and glutamate dehydrogenase were present in leaves and roots of the species examined.¹⁵N-labelled nitrate and ammonium were used to compare the assimilatory characteristics of two species:Enterolobium contortisiliquum, with a high capacity to reduce nitrate, andCalophyllum brasiliense, of low capacity. The rate of nitrate assimilation in the former was five times that of the latter. Both species had similar rates of ammonium assimilation. Results for eight species of contrasting habitats showed that leaf nitrogen content increased in parallel with xylem sap nitrogen concentrations, suggesting that the ability of the root system to acquire, assimilate or export nitrate determines shoot nitrogen status. These results emphasise the importance of nitrogen transport and metabolism in roots as determinants of whole plant nitrogen status.     

Uptake of Nitrate by Mutants of Arabidopsis thaliana, Disturbed in Uptake or Reduction of Nitrate

A study of nitrate and chlorate uptake by Arabidopsis thaliana was made with a wildtype and two mutant types, both mutants having been selected by resistance to high chlorate concentrations. All plants were grown on a nutrient solution with nitrate and/or ammonium as the nitrogen source. Uptake was determined from depletion in the ambient solution. Nitrate and chlorate were able to induce their own uptake mechanisms. Plants grown on ammonium nitrate showed a higher subsequent uptake rate of nitrate and chlorate than plants grown on ammonium alone. Mutant B25, which has no nitrate reductase activity, showed higher rates of nitrate and chlorate uptake than the wildtype, when both types were grown on ammonium nitrate. Therefore, the uptake of nitrate is not dependent on the presence of nitrate reductase. Nitrate has a stimulating effect on nitrate and chlorate uptake, whereas some product of nitrate and ammonium assimilation inhibits uptake of both ions by negative feedback. Mutant B 1, which was supposed to have a low chlorate uptake rate, also has disturbed uptake characteristics for nitrate.     

Growth rate and phosphate utilization of some Carex species from a range of oligotrophic to eutrophic swamp habitats

In a growth experiment at phosphate levels varying between 0,0005 and 0.1 mM phosphate, relative growth rates and other growth parameters were determined in Carex species (C. rostrata Stokes, C. limosa L., C. lasiocarpa Ehrh., C. diandra Schrank, and C. acutiformis Ehrh., listed in order of increasing nutrient availability of their natural habitats). In all species, more efficient utilization of the phosphate was observed with decreasing phosphate levels, together with reduced fresh shoot ratio. In addition, each species shows characteristics which may help it to grow under low phosphate conditions: relatively high phosphate level in the plant (C. rostrata), a low fresh shoot ratio (C. limosa) and regulation of uptake and translocation of phosphate in such a way that relative growth rate during the first two weeks is maintained or even increased for a longer period (C. rostrata, C. limosa, and C. diandra). In the studied Carex species, the contribution of fresh shoot ratio to relative growth rate is much larger than that of net assimilation rate. C. acutiformis, a species of eutrophic areas, had the highest relative growth rate.     

The use of mean pool abundances to interpret 15N tracer experiments

A pulse dilution ¹⁵N technique was used in the field to determine the effect of the ammonium to nitrate ratio in a fertilizer application on the uptake of ammonium and nitrate by ryegrass and on gross rates of mineralization and nitrification. Two experiments were performed, corresponding approximately to the first and second cuts of grass. Where no substantial recent immobilization of inorganic nitrogen had occurred, mineralization was insensitive to the form of nitrogen applied, ranging from 2.1–2.6 kg N ha^-1 d^-1. The immobilization of ammonium increased as the proportion of ammonium in the application increased. In the second experiment there was evidence that high rates of immobilization in the first experiment were associated with high rates of mineralization in the second. The implication was that some nitrogen immobilized in the first experiment was re-mineralized during the second. Whether this was nitrogen taken up, stored in roots and released following defoliation was not clear. Nitrification rates in this soil were low (0.1–0.63 kg N ha^-1 d^-1), and as a result, varying the ratio of ammonium to nitrate applied markedly altered the relative uptake of ammonium and nitrate. In the first experiment, where temperatures were low, preferential uptake of ammonium occurred, but where >90% of the uptake was as ammonium, a reduction in yield and nitrogen uptake was observed. In the second experiment, where temperatures and growth rates were higher, the proportion of ammonium to nitrate taken up had no effect on yield or nitrogen uptake.