Cross-Ecosystem Comparisons of In Situ Plant Uptake of Amino Acid-N and NH4 +

Plant and microbial use of nitrogen (N) can be simultaneously mutualistic and competitive, particularly in ecosystems dominated by mycorrhizal fungi. Our goal was to quantify plant uptake of organic and inorganic N across a broad latitudinal gradient of forest ecosystems that varied with respect to overstory taxon, edaphic characteristics, and dominant mycorrhizal association. Using ¹³C and ¹⁵N, we observed in situ the cycling dynamics of NH₄ ⁺ and glycine through various soil pools and fine roots over 14 days. Recovery of ¹⁵N as soil N varied with respect to N form, forest type, and sampling period; however, there were similarities in the cycling dynamics of glycine and NH₄ ⁺ among all forest types. Microbial immobilization of ¹⁵N was immediately apparent for both treatments and represented the largest sink (~25%) for ¹⁵N among extractable soil N pools during the first 24 h. In contrast, fine roots were a relatively small sink (<10%) for both N forms, but fine root ¹³C enrichment indicated that plants in all forest types absorbed glycine intact, suggesting that plants and microbes effectively target the same labile soil N pools. Relative uptake of amino acid-N versus NH₄ ⁺ varied significantly among sites and approximately half of this variation was explained by mycorrhizal association. Estimates of plant uptake of amino acid-N relative to NH₄ ⁺ were 3× higher in ectomycorrhizal-dominated stands (1.6 ± 0.2) than arbuscular mycorrhizae-dominated stands (0.5 ± 0.1). We conclude that free amino acids are an important component of the N economy in all stands studied; however, in these natural environments plant uptake of organic N relative to inorganic N is explained as much by mycorrhizal association as by the availability of N forms per se.     

Intact amino acid uptake by northern hardwood and conifer trees

Empirical and modeling studies of the N cycle in temperate forests of eastern North America have focused on the mechanisms regulating the production of inorganic N, and assumed that only inorganic forms of N are available for plant growth. Recent isotope studies in field conditions suggest that amino acid capture is a widespread ecological phenomenon, although northern temperate forests have yet to be studied. We quantified fine root biomass and applied tracer-level quantities of U–¹³C₂–¹⁵N-glycine, ¹⁵NH₄ ⁺ and ¹⁵NO₃ ⁻ in two stands, one dominated by sugar maple and white ash, the other dominated by red oak, beech, and hemlock, to assess the importance of amino acids to the N nutrition of northeastern US forests. Significant enrichment of ¹³C in fine roots 2 and 5 h following tracer application indicated intact glycine uptake in both stands. Glycine accounted for up to 77% of total N uptake in the oak–beech–hemlock stand, a stand that produces recalcitrant litter, cycles N slowly and has a thick, amino acid-rich organic horizon. By contrast, glycine accounted for only 20% of total N uptake in the sugar maple and white ash stand, a stand characterized by labile litter and rapid rates of amino acid production and turnover resulting in high rates of mineralization and nitrification. This study shows that amino acid uptake is an important process occurring in two widespread, northeastern US temperate forest types with widely differing rates of N cycling.     

Amino acid uptake by temperate tree species characteristic of low- and high-fertility habitats

The relationship between inorganic nitrogen (N) cycling and plant productivity is well established. However, recent research has demonstrated the ability of plants to take up low molecular weight organic N compounds (i.e., amino acids) at rates that often rival those of inorganic N forms. In this study, we hypothesize that temperate forest tree species characteristic of low-fertility habitats will prefer amino acids over species characteristic of high-fertility habitats. We measured the uptake of ¹⁵N-labeled amino acids (glycine, glutamine, arginine, serine), ammonium (NH₄ ⁺), and nitrate (NO₃ ⁻) by four tree species that commonly occur in eastern North America, where their abundances have been correlated with inorganic N availability. Specific uptake rates of amino acids were largely similar for all tree species; however, high-fertility species took up NH₄ ⁺ at rates more than double those of low-fertility species, rendering amino acid N relatively more important to the N nutrition of low-fertility species. Low-fertility species acquired over four times more total N from arginine compared to NH₄ ⁺ and NO₃ ⁻; high-fertility species acquired the most N from NH₄ ⁺. Arginine had the highest uptake rates of any amino acid by all species; there were no significant differences in uptake rates of the remaining amino acids. Our results support the idea that the dominant species in a particular habitat are those best able to utilize the most available N resources.     

Effects of elevated atmospheric carbon dioxide on amino acid and NH4+-N cycling in a temperate pine ecosystem

Rising atmospheric carbon dioxide (CO₂) is expected to increase forest productivity, resulting in greater carbon (C) storage in forest ecosystems. Because elevated atmospheric CO₂ does not increase nitrogen (N) use efficiency in many forest tree species, additional N inputs will be required to sustain increased net primary productivity (NPP) under elevated atmospheric CO₂. We investigated the importance of free amino acids (AAs) as a source for forest N uptake at the Duke Forest Free Air CO₂ Enrichment (FACE) site, comparing its importance with that of better‐studied inorganic N sources. Potential proteolytic enzyme activity was monitored seasonally, and individual AA concentrations were measured in organic horizon extracts. Potential free AA production in soils ranged from 190 to 690 nmol N g⁻¹ h⁻¹ and was greater than potential rates of soil NH₄⁺ production. Because of this high potential rate of organic N production, we determined (1) whether intact AA uptake occurs by Pinus taeda L., the dominant tree species at the FACE site, (2) if the rate of cycling of AAs is comparable with that of ammonium (NH₄⁺), and (3) if atmospheric CO₂ concentration alters the aforementioned N cycling processes. A field experiment using universally labeled ammonium (¹⁵NH₄⁺) and alanine (¹³C₃H₇¹⁵NO₂) demonstrated that ¹⁵N is more readily taken up by plants and heterotrophic microorganisms as NH₄⁺. Pine roots and microbes take up on average 2.4 and two times as much NH₄^{+ 15}N compared with alanine ¹⁵N 1 week after tracer application. N cycling through soil pools was similar for alanine and NH₄⁺, with the greatest ¹⁵N tracer recovery in soil organic matter, followed by microbial biomass, dissolved organic N, extractable NH₄⁺, and fine roots. Stoichiometric analyses of ¹³C and ¹⁵N uptake demonstrated that both plants and soil microorganisms take up alanine directly, with a ¹³C : ¹⁵N ratio of 3.3 : 1 in fine roots and 1.5 : 1 in microbial biomass. Our results suggest that intact AA (alanine) uptake contributes substantially to plant N uptake in loblolly pine forests. However, we found no evidence supporting increased recovery of free AAs in fine roots under elevated CO₂, suggesting plants will need to acquire additional N via other mechanisms, such as increased root exploration or increased N use efficiency.     

Amino acid uptake in deciduous and coniferous taiga ecosystems

We measured in situ uptake of amino acids and ammonium across deciduous and coniferous taiga forest ecosystems in interior Alaska to examine the idea that late successional (coniferous) forests rely more heavily on dissolved organic nitrogen (DON), than do early successional (deciduous) ecosystems. We traced ¹⁵N-NH₄⁺ and ¹³C-¹⁵N-amino acids from the soil solution into plant roots and soil pools over a 24 h period in stands of early successional willow and late successional black spruce. Late successional soils have much higher concentrations of amino acid in soil solution and a greater ratio of DON to dissolved inorganic N (DIN) (ammonium plus nitrate) than do early successional soils. Moreover, late successional coniferous forests exhibit higher rates of soil proteolytic activity, but lower rates of inorganic N turnover. Differences in ammonium and amino acid uptake by early successional willow stands were insignificant. By contrast, the in situ uptake of amino acid by late successional black spruce forests were approximately 4-fold greater than ammonium uptake. The relative difference in uptake of ammonium and amino acids in these forests was approximately proportional to the relative difference of these N forms in the soil solution. Thus, we suggest that differences in uptake of different N forms across succession in these boreal forests largely reflect edaphic variation in available soil N (composition), rather than any apparent physiological specialization to absorb particular forms of N. These finding are relevant to our understanding of how taiga ecosystems may respond to increases in temperature, fire frequency, N deposition, and other potential consequences of global change.     

Fertilization rates and clay fixed ammonium in two Quebec soils

Clay fixed NH₄ ⁺ can provide a significant sink for fertilizer N, as well as a source of N for plant uptake. Knowledge or soil NH₄ ⁺ fixing capacity and release for crops is necessary to develop long-term fertilizer programs. Field experiments with corn (Zea mays L.) were carried out to investigate soil NH₄ ⁺ fixing capacity and subsequent release as influenced by fertilizer rates using ¹⁵N in a Ste. Rosalie clay (fine, mixed, frigid, Typic Humaquept) and a Chicot sandy clay loam (fine-loamy, mixed, frigid, Typic Hapludalf). With high N rates increased NH₄ ⁺ fixation occurred only in the Ste. Rosalie soil. At the end of the first growing season, fertilizer N recovery as clay fixed NH₄ ⁺ for high and normal rates of fertilizer in the Ste. Rosalie soil was 17.8% and 28.7%, respectively and the recovery for the high and normal rates in the Chicot soil was 4.6 and 10.5%, respectively. Significant amounts of clay fixed NH₄ ⁺-N were released in the soil profile in the second year after ¹⁵N application on the Chicot soil. Recently clay fixed fertilizer NH₄ ⁺N was released more rapidly than that of the native fixed NH₄ ⁺, from the surface layer of the Ste. Rosalie soil. The fertilizer fixed NH₄ ⁺ seems to be in a more labile N pool than the native fixed NH₄ ⁺-N in the Chicot soil.     

Variability in Soil Nitrogen Retention Across Forest,Urban, and Agricultural Land Uses

In regions of mixed land use, some ecosystems are sinks for N pollution and others are sources. Yet, beyond this gross characterization, we have little understanding of how adjacent land-use types vary in mechanisms of N cycling and retention. This study assessed the rate and magnitude of soil N retention pathways in forest, urban, and agricultural ecosystems. Soil plots in each land use were labeled with inorganic ¹⁵N and cored at 15 min, 2 days, and 20 days following injection. Subsamples were biologically fractionated to differentiate labile and stable pools, while gross N transformations were assessed via the ¹⁵N isotope dilution method. Stable soil organic ¹⁵N formed rapidly (within 15 min) in all land uses when added as ¹⁵NH₄ ⁺, and became a proportionally larger sink for inorganic ¹⁵N over time. Forests had the lowest gross immobilization rates, but the greatest amount of stable N formation. Rapid retention of NH₄ ⁺ in forests may be driven by abiotic processes, with root uptake becoming a more important mechanism of retention over time. Urban sites, on the other hand, had the highest gross microbial immobilization rates and highest root N uptake, suggesting that high short-term N retention may be due to rapid biological processing. Agricultural systems, with low root uptake and the lowest stable N formation, had little capacity for retention of added N. These apparently distinct land-use cases can be understood by synthesizing several emerging aspects of N retention theory that (1) distinguish kinetic and capacity N saturation, (2) recognize links between soil C saturation on minerals and N retention, and (3) account for rapid transfers of NH₄ ⁺ to stable organic pools.     

Mineralization-immobilization and plant uptake of nitrogen as influenced by the spatial distribution of cattle slurry in soils of different texture

The effect of incorporating cattle slurry in soil, either by mixing or by simulated injection into a hollow in soil, on the ryegrass uptake of total N and ¹⁵NH₄ ⁺-N was determined in three soils of different texture. The N accumulation in Italian ryegrass (Lolium multiflorum L.) from slurry N and from an equivalent amount of NH₄ ⁺-N in (¹⁵NH₄) SO₄ (control) was measured during 6 months of growth in pots. After this period the total recovery of labelled N in the top soil plus herbage was similar in the slurry and the control treatments. This indicated that gaseous losses from slurry NH₄ ⁺-N were insignificant. Consequently, the availability of slurry N to plants was mainly influenced by the mineralization-immobilization processes. The apparent utilization of slurry NH₄ ⁺-N mixed into soil was 7%, 14% and 24% lower than the utilization of (NH₄)₂SO₄-N in a sand soil, a sandy loam soil and a loam soil, respectively. Thus, the net immobilization of N due to slurry application increased with increasing soil clay content, whereas the recovery in plants of ¹⁵N-labelled NH₄ ⁺-N from slurry was similar on the three soils. A parallel incubation experiment showed that the immobilization of slurry N occurred within the first week after slurry application. The incorporation of slurry N by simulated injection increased the plant uptake of both total and labelled N compared to mixing the slurry into the soil. The apparent utilization of injected slurry NH₄ ⁺-N was 7% higher, 8% lower and 4% higher than the utilization of (NH₄)₂SO₄-N in the sand, the sandy loam and the loam soil, respectively. It is concluded that the spatial distribution of slurry in soil influenced the net mineralization of N to the same degree as did the soil type.     

Challenging the paradigm of nitrogen cycling: no evidence of in situ resource partitioning by coexisting plant species in grasslands of contrasting fertility

In monoculture, certain plant species are able to preferentially utilize different nitrogen (N) forms, both inorganic and organic, including amino acids and peptides, thus forming fundamental niches based on the chemical form of N. Results from field studies, however, are inconsistent: Some showing that coexisting plant species predominantly utilize inorganic N, while others reveal distinct interspecies preferences for different N forms. As a result, the extent to which hypothetical niches are realized in nature remains unclear. Here, we used in situ stable isotope tracer techniques to test the idea, in temperate grassland, that niche partitioning of N based on chemical form is related to plant productivity and the relative availability of organic and inorganic N. We also tested in situ whether grassland plants vary in their ability to compete for, and utilize peptides, which have recently been shown to act as an N source for plants in strongly N-limited ecosystems. We hypothesized that plants would preferentially use NO₃⁻-N and NH₄⁺-N over dissolved organic N in high-productivity grassland where inorganic N availability is high. On the other hand, in low-productivity grasslands, where the availability of dissolved inorganic N is low, and soil availability of dissolved organic N is greater, we predicted that plants would preferentially use N from amino acids and peptides, prior to microbial mineralization. Turves from two well-characterized grasslands of contrasting productivity and soil N availability were injected, in situ, with mixtures of ¹⁵N-labeled inorganic N (NO₃⁻ and NH₄⁺) and ¹³C¹⁵N labeled amino acid (l-alanine) and peptide (l-tri-alanine). In order to measure rapid assimilation of these N forms by soil microbes and plants, the uptake of these substrates was traced within 2.5 hours into the shoots of the most abundant plant species, as well as roots and the soil microbial biomass. We found that, contrary to our hypothesis, the majority of plant species across both grasslands took up most N in the form of NH₄⁺, suggesting that inorganic N is their predominant N source. However, we did find that organic N was a source of N which could be utilized by plant species at both sites, and in the low-productivity grassland, plants were able to capture some tri-alanine-N directly. Although our findings did not support the hypothesis that differences in the availability of inorganic and organic N facilitate resource partitioning in grassland, they do support the emerging view that peptides represent a significant, but until now neglected, component of the terrestrial N cycle.     

氨基酸添加对亚热带森林红壤氮素转化的影响

为探究氨基酸氮形态对亚热带土壤氮素含量及转化的影响,选择建瓯市万木林保护区的山地红壤为对象,采用室内培养实验法,通过设计60%和90%WHC两种土壤含水量并添加不同性质氨基酸,测定了土壤中铵态氮、硝态氮、可溶性有机氮的含量和氧化亚氮的释放量,分析了可溶性有机碳、土壤p H值的大小变化及其与氮素的相互关系。结果表明:与对照处理相比,氨基酸添加显著增加了土壤NH_4~+-N含量并使土壤p H值升高,且在一定程度上解除了高含水量(90%WHC)对NH_4~+-N产生的抑制,其中甲硫氨基酸的效果最为明显。酸性、碱性、中性氨基酸对土壤NO_3~--N含量和N_2O释放影响不显著,但甲硫氨基酸可显著抑制土壤硝化从而导致NH_4~+-N的积累,并在培养前期抑制土壤N_2O产生而在培养后期促进N_2O释放,总体上促进N_2O释放。60%WHC的氨基酸添加处理较90%WHC条件下降低土壤可溶性有机氮的幅度更大。氨基酸对土壤氮素转化的影响与带电性关系较小,而可能与其分解产物密切相关。可见,不同性质氨基酸处理对森林土壤氮素含量及转化存在不同程度的影响,且甲硫氨基酸对土壤氮素转化的影响机理值得深入研究。     

Recovery of 15N-labeled urea and soil nitrogen dynamics as affected by irrigation management and nitrogen application rate in a double rice cropping system

A sophisticated soil microcosm system and ¹⁵N-labeled urea were used to investigate nitrogen (N) use efficiency and soil N dynamics in a rice monoculture system in two successive seasons. Topsoil (0 cm?C20 cm) and subsoil (20 cm?C50 cm) samples were collected from a traditional double rice cropping field in the Jiangxi Province, China, and these soil samples were derived from Quaternary red clay. Treatments were randomly assigned with two irrigation regimes and three N application rates (no application control, 80% traditional rate and 100% traditional rate noted as N0, N1 and N2, respectively). The levels of ¹⁵N recovery of plants, ¹⁵N and N remaining in soil were determined. Moreover, the N dynamic of soil solution from different layers of the soil profile was surveyed. The results showed that the effects of irrigation management and N application rate varied in different rice growing seasons. Irrigation regimes had remarkable effects on grain yield and chemical ¹⁵N fertilizers (CF-¹⁵N) uptake. When compared to flood irrigation (FI), the shallow water depth with wetting and drying (WD) increased grain yield up to 5.7%?C20%. Although the highest grain yield was obtained with reduced N application level, both N apparent recovery (NAR) and ¹⁵N use efficiency (the percentage of plant N uptake derived from applied N, %Ndfan) significantly decreased with increasing N inputs. However, the interaction between irrigation management and N application rate on grain yield and N use efficiency (NUE) of CF-¹⁵N were not significant. A survey of soil solutions every 5 days indicated that NH ₄ ⁺ -N was the main residual form of N, and high NH ₄ ⁺ -N leaching was observed. When compared to FI, WD decreased vertical NH ₄ ⁺ -N and TN leaching, especially at 10?C50 cm depths of soil profile in the second season. NH ₄ ⁺ -N was the main N residual form in the soil profile. Therefore, in this study, the WD irrigation regime and reduced rate (N1) was the optimal irrigation and fertilizer management strategy to increase the NUE of CF-N, increase the after effects of CF-¹⁵N, decrease leaching loss of CF-¹⁵N and minimize the shallow groundwater pollution risk, which were all beneficial for the ecological environment.     

Low sedimentary accumulation of lead caused by weak downward export of organic matter in Hudson Bay,northern Canada

Subtropical forests receive increasing amounts of atmogenic nitrogen (N), both as ammonium (NH₄ ⁺) and nitrate (NO₃ ^?). Previous long-term studies indicate efficient turnover of atmogenic NH₄ ⁺ to NO₃ ^? in weathered, acidic soils of the subtropics, leading to excessive NO₃ ^? leaching. To clarify the mechanism governing the fate of atmogenic inputs in these soils, we conducted an in situ ¹⁵N tracing experiment in the TieShanPing (TSP) forested catchment, SW China. ¹⁵NH₄NO₃, NH ₄ ¹⁵ NO₃ and ¹⁵N-glutamic acid were applied to an upland hillslope soil and inorganic N, total soil N and nitrous oxide (N₂O) were monitored for nine days. Incorporation of ¹⁵NO₃ ^? into soil organic N was negligible and 80% of the applied label was lost from the top soil (0–15 cm) primarily by leaching within 9 days. In contrast, ¹⁵NH₄ ⁺ was largely retained in soil organic N. However, instant production of ¹⁵NO₃ ^? in the ¹⁵NH₄ ⁺ treatment suggested active nitrification. In both the ¹⁵NH₄ ⁺ and ¹⁵N-glutamic acid treatments, the ¹⁵N enrichment in the NO₃ ^? pool exceeded that in the NH₄ ⁺ pool one day after ¹⁵N application, suggesting preferential nitrification of added ¹⁵NH₄ ⁺ with subsequent dilution of the NH₄ ⁺ pool and/or immobilization of ¹⁵NH₄ ⁺ followed by heterotrophic nitrification. The cumulative recovery of ¹⁵N in N₂O after 9 days ranged from 2.5 to 6.0% in the ¹⁵NO₃ ^? treatment, confirming the previously reported significant response of N₂O emission to N deposition. Source partitioning of ¹⁵N₂O demonstrated a measurable contribution of nitrification to N₂O emissions, particularly at low soil moistures. Our study emphasizes the role of a fast-cycling organic N pool (including microbial N) for retention and transformation of atmogenic NH₄ ⁺ in subtropical, acid forest soils. Thus, it explains the near-quantitative leaching of deposited N (as NO₃ ^? and NH₄ ⁺) common to subtropical forest soils with chronic, elevated atmogenic N inputs by (i) negligible retention of NO₃ ^? in the soil and (ii) rapid immobilization-mineralization of NH₄ ⁺ followed by nitrification. Our findings point to a leaky N cycle in N-saturated Chinese subtropical forests with consequences for regional soil acidification, N pollution of fresh waters and N₂O emission.     

The partitioning of fertilizer-N between soil and crop: Comparison of ammonium and nitrate applications

Field experiments were carried out in 1987 on winter wheat crops grown on three types of soil. ¹⁵N-labelled urea, ¹⁵NH₄NO₃ or NH₄ ¹⁵NO₃ (80 kg N ha^-1) was applied at tillering. The soils (chalky soil, hydromorphic loamy soil, sandy clay soil) were chosen to obtain a range of nitrogen dynamics, particularly nitrification. Soil microbial N immobilization and crop N uptake were measured at five dates. Shortly after fertilizer application (0–26 days), the amount of N immobilized in soil were markedly higher with labelled urea or ammonium than that with nitrate in all soils. During the same period, crop ¹⁵N uptake occurred preferentially at the expense of nitrate. Nitrification differed little between soils, the rates were 2.0 to 4.7 kg N ha^-1 day^-1 at 9°C daily mean temperature. The differences in immobilization and uptake had almost disappeared at flowering and harvest. ¹⁵N recovery in soil and crop varied between 50 and 100%. Gaseous losses probably occurred by volatilization in the chalky soil and denitrification in the hydromorphic loamy soil. These losses affected the NH₄ ⁺ and NO₃ ^- pools differently and determined the partitioning of fertilizer-N between immobilization and absorption.     

Flow of Deposited Inorganic N in Two Gleysol-dominated Mountain Catchments Traced with <Superscript>15</Superscript>NO<Subscript>3</Subscript><Superscript>−</Superscript> and <Superscript>15</Superscript>NH<Subscript>4</Subscript><Superscript>+</Superscript>

In two mountain ecosystems at the Alptal research site in central Switzerland, pulses of ¹⁵NO₃ and ¹⁵NH₄ were separately applied to trace deposited inorganic N. One forested and one litter meadow catchment, each approximately 1600 m², were delimited by trenches in the Gleysols. K¹⁵NO₃ was applied weekly or fortnightly over one year with a backpack sprayer, thus labelling the atmospheric nitrate deposition. After the sampling and a one-year break, ¹⁵NH₄Cl was applied as a second one-year pulse, followed by a second sampling campaign. Trees (needles, branches and bole wood), ground vegetation, litter layer and soil (LF, A and B horizon) were sampled at the end of each labelling period. Extractable inorganic N, microbial N, and immobilised soil N were analysed in the LF and A horizons. During the whole labelling period, the runoff water was sampled as well. Most of the added tracer remained in both ecosystems. More NO₃⁻ than NH₄⁺ tracer was retained, especially in the forest. The highest recovery was in the soil, mainly in the organic horizon, and in the ground vegetation, especially in the mosses. Event-based runoff analyses showed an immediate response of ¹⁵NO₃⁻ in runoff, with sharp ¹⁵N peaks corresponding to discharge peaks. NO₃⁻ leaching showed a clear seasonal pattern, being highest in spring during snowmelt. The high capacity of N retention in these ecosystems leads to the assumption that deposited N accumulates in the soil organic matter, causing a progressive decline of its C:N ratio.     

Changes in the C/N balance caused by increasing external ammonium concentrations are driven by carbon and energy availabilities during ammonium nutrition in pea plants: the key roles of asparagine synthetase and anaplerotic enzymes

An understanding of the mechanisms underlying ammonium (NH₄⁺) toxicity in plants requires prior knowledge of the metabolic uses for nitrogen (N) and carbon (C). We have recently shown that pea plants grown at high NH₄⁺ concentrations suffer an energy deficiency associated with a disruption of ionic homeostasis. Furthermore, these plants are unable to adequately regulate internal NH₄⁺ levels and the cell‐charge balance associated with cation uptake. Herein we show a role for an extra‐C application in the regulation of C–N metabolism in NH₄⁺‐fed plants. Thus, pea plants (Pisum sativum) were grown at a range of NH₄⁺ concentrations as sole N source, and two light intensities were applied to vary the C supply to the plants. Control plants grown at high NH₄⁺ concentration triggered a toxicity response with the characteristic pattern of C‐starvation conditions. This toxicity response resulted in the redistribution of N from amino acids, mostly asparagine, and lower C/N ratios. The C/N imbalance at high NH₄⁺ concentration under control conditions induced a strong activation of root C metabolism and the upregulation of anaplerotic enzymes to provide C intermediates for the tricarboxylic acid cycle. A high light intensity partially reverted these C‐starvation symptoms by providing higher C availability to the plants. The extra‐C contributed to a lower C4/C5 amino acid ratio while maintaining the relative contents of some minor amino acids involved in key pathways regulating the C/N status of the plants unchanged. C availability can therefore be considered to be a determinant factor in the tolerance/sensitivity mechanisms to NH₄⁺ nutrition in plants.     

Nitrate influx kinetic parameters of five potato cultivars during vegetative growth

This study examines the effect of elevated CO₂ on short-term partitioning of inorganic N between a grass and soil micro-organisms. ¹⁵N-labelled NH₄⁺ was injected in the soil of mesocosms of Holcus lanatus (L.) that had been grown for more than 15 months at ambient or elevated CO₂ in reconstituted grassland soil. After 48 h, the percentage recovery of added ¹⁵N was increased in soil microbial biomass N at elevated CO₂, was unchanged in total plant N and was decreased in soil extractable N. However, plant N content and microbial biomass N were not significantly affected by elevated CO₂. These results and literature data from plant–microbial ¹⁵N partitioning experiments at elevated CO₂ suggest that the mechanisms controlling the effects of CO₂ on short- vs. long-term N uptake and turnover differ. In particular, short-term immobilisation of added N by soil micro-organisms at elevated CO₂ does not appear to lead to long-term increases in N in soil microbial biomass. In addition, the increased soil microbial C:N ratios that we observed at elevated CO₂ suggest that long-term exposure to CO₂ alters either the functioning or structure of these microbial communities.     

Mycorrhizas alter nitrogen acquisition by the terrestrial orchid Cymbidium goeringii

Background and Aims

Orchid mycorrhizas exhibit a unique type of mycorrhizal symbiosis that occurs between fungi and plants of the family Orchidaceae. In general, the roots of orchids are typically coarse compared with those of other plant species, leading to a considerably low surface area to volume ratio. As a result, orchids are often ill-adapted for direct nutrient acquisition from the soil and so mycorrhizal assocaitions are important. However, the role of the fungal partners in the acquisition of inorganic and organic N by terrestrial orchids has yet to be clarified.

Methods

Inorganic and amino acid N uptake by non-mycorrhizal and mycorrhizal Cymbidium goeringii seedlings, which were grown in pots in a greenhouse, was investigated using a ¹⁵N-labelling technique in which the tracer was injected at two different soil depths, 2·5 cm or 7·5 cm. Mycorrhizal C. goeringii seedlings were obtained by inoculation with three different mycorrhizal strains isolated from the roots of wild terrestrial orchids (two C. goeringii and one C. sinense).

Key Results

Non-mycorrhizal C. goeringii primarily took up NO₃⁻ from tracers injected at 2·5-cm soil depth, whereas C. goeringii inoculated with all three mycorrhiza primarily took up NH₄⁺ injected at the same depth. Inoculation of the mycorrhizal strain MLX102 (isolated from adult C. sinense) on C. goeringii roots only significantly increased the below-ground biomass of the C. goeringii; however, it enhanced ¹⁵NH₄⁺ uptake by C. goeringii at 2·5-cm soil depth. Compared to the uptake of tracers injected at 2·5-cm soil depth, the MLX102 fungal strain strongly enhanced glycine-N uptake by C. goeringii from tracers injected at 7·5-cm soil depth. Cymbidium goeringii inoculated with CLB113 and MLX102 fungal strains demonstrated a similar N uptake pattern to tracers injected at 2·5-cm soil depth.

Conclusions

These findings demonstrate that mycorrhizal fungi are able to switch the primary N source uptake of a terrestrial orchid, in this case C. goeringii, from NO₃⁻ to NH₄⁺. The reasons for variation in N uptake in the different soil layers may be due to possible differentiation in the mycorrhizal hyphae of the C. goeringii fungal partner.     

Spatiotemporal variations affect uptake of inorganic and organic nitrogen by dominant plant species in an alpine wetland

Aims

Dominant plant species may coexist and maintain high productivity in alpine wetland through available nitrogen (N) niche differentiation over time and space. We tested the hypotheses that dominant plant species differ in uptake of inorganic and organic N and that such differences depend on soil depth and season.

Methods

We conducted a short-term ¹⁵N-labeling experiment in an alpine wetland on the Tibetan Plateau. The experiment used a factorial design with three N forms (nitrate, ammonium and glycine), three soil depths (0–5, 5–10 and 10–15 cm), two seasons (May and July) and three dominant species (Carex muliensis, C. lasiocarpa and Potentilla anserina).

Results

All three species took up organic N (glycine), but showed different patterns over seasons and depths. ¹⁵N uptake rate was higher in May than in July in C. muliensis and C. lasiocarpa, but lower in May than in July in P. anserina. C. muliensis took up more ¹⁵NH₄ ⁺ and ¹⁵NO₃ ^? than glycine-¹⁵N at all soil depths. C. lasiocarpa took up more glycine-¹⁵N than ¹⁵NH₄ ⁺ or ¹⁵NO₃ at 5–10 cm depth. P. anserina showed little difference in uptake at any soil depths.

Conclusions

Dominant species in alpine wetland are able to take up both organic and inorganic N, but show different patterns depending on N form, soil depth, season and their interactions.     

Uptake and inhibition kinetics of nitrogen in Microcystis aeruginosa: Results from cultures and field assemblages collected in the San Francisco Bay Delta,CA

The nitrogen (N) uptake kinetic parameters for Microcystis field assemblages collected from the San Francisco Bay Delta (Delta) in 2012 and non-toxic and toxic laboratory culture strains of M. aeruginosa were assessed. The ¹⁵N tracer technique was used to investigate uptake of ammonium (NH₄⁺), nitrate (NO₃⁻), urea and glutamic acid over short-term incubations (0.5–1 h), and to study inhibition of NO₃⁻, NH₄⁺ and urea uptake by NH₄⁺, NO₃⁻ and NH₄⁺, respectively. This study demonstrates that Delta Microcystis can utilize different forms of inorganic and organic N, with the greatest capacity for NH₄⁺ uptake and the least for glutamic acid uptake, although N uptake did not always follow the classic Michaelis–Menten hyperbolic relationship at substrate concentrations up to 67 μmol N L⁻¹. Current ambient N concentrations in the Delta may be at sub-saturating levels for N uptake, indicating that if N loading (especially NH₄⁺) were to increase, Delta Microcystis assemblages have the potential for increased N uptake rates. Delta Microcystis had the highest specific affinity, α, for NH₄⁺ and the lowest for NO₃⁻. In culture, N uptake by non-toxic and toxic M. aeruginosa strains was much higher than from the field, but followed similar N utilization trends to those in the field. Neither strain showed severe inhibition of NO₃⁻ uptake by NH₄⁺ or inhibition of NH₄⁺ uptake on NO₃⁻, but both strains showed some inhibition of urea uptake by NH₄⁺.     

Segregation of nitrogen use between ammonium and nitrate of ectomycorrhizas and beech trees

Here, we characterized nitrogen (N) uptake of beech (Fagus sylvatica) and their associated ectomycorrhizal (EM) communities from NH₄⁺ and NO₃^?. We hypothesized that a proportional fraction of ectomycorrhizal N uptake is transferred to the host, thereby resulting in the same uptake patterns of plants and their associated mycorrhizal communities. ¹⁵N uptake was studied under various field conditions after short‐term and long‐term exposure to a pulse of equimolar NH₄⁺ and NO₃^? concentrations, where one compound was replaced by ¹⁵N. In native EM assemblages, long‐term and short‐term ¹⁵N uptake from NH₄⁺ was higher than that from NO₃^?, regardless of season, water availability and site exposure, whereas in beech long‐term ¹⁵N uptake from NO₃^? was higher than that from NH₄⁺. The transfer rates from the EM to beech were lower for ¹⁵N from NH₄⁺ than from NO₃^?. ¹⁵N content in EM was correlated with ¹⁵N uptake of the host for ¹⁵NH₄⁺, but not for ¹⁵NO₃^?‐derived N. These findings suggest stronger control of the EM assemblage on N provision to the host from NH₄⁺ than from NO₃^?. Different host and EM accumulation patterns for inorganic N will result in complementary resource use, which might be advantageous in forest ecosystems with limited N availability.