The seasonal dynamics of amino acids and other nutrients in Alaskan Arctic tundra soils

Past research strongly indicates the importance of amino acids in the N economy of the Arctic tundra, but little is known about the seasonal dynamics of amino acids in tundra soils. We repeatedly sampled soils from tussock, shrub, and wet sedge tundra communities in the summers of 2000 and 2001 and extracted them with water (H₂O) and potassium sulfate (K₂SO₄) to determine the seasonal dynamics of soil amino acids, ammonium (NH₄⁺), nitrate (NO₃^–), dissolved organic nitrogen (DON), dissolved organic carbon (DOC), and phosphate (PO₄^2–). In the H₂O extractions mean concentrations of total free amino acids (TFAA) were higher than NH₄⁺ in all soils but shrub. TFAA and NH₄⁺ were highest in wet sedge and tussock soils and lowest in shrub soil. The most predominant amino acids were alanine, arginine, glycine, serine, and threonine. None of the highest amino acids were significantly different than NH₄⁺ in any soil but shrub, in which NH₄⁺ was significantly higher than all of the highest individual amino acids. Mean NO₃^– concentrations were not significantly different from mean TFAA and NH₄⁺ concentrations in any soil but tussock, where NO₃^– was significantly higher than NH₄⁺. In all soils amino acid and NH₄⁺ concentrations dropped to barely detectable levels in the middle of July, suggesting intense competition for N at the height of the growing season. In all soils but tussock, amino acid and NH₄⁺ concentrations rebounded in August as the end of the Arctic growing season approached and plant N demand decreased. This pattern suggests that low N concentrations in tundra soils at the height of the growing season are likely the result of an increase in soil N uptake associated with the peak in plant growth, either directly by roots or indirectly by microbes fueled by increased root C inputs in mid-July. As N availability decreased in July, PO₄^2– concentrations in the K₂SO₄ extractions increased dramatically in all soils but shrub, where there was a comparable increase in PO₄^2– later in the growing season. Previous research suggests that these increases in PO₄^2– concentrations are due to the mineralization of organic phosphorus by phosphatase enzymes associated with soil microbes and plant roots, and that they may have been caused by an increase in organic P availability.     

Landscape patterns of free amino acids in arctic tundra soils

Concentrations of free amino acids were measured in soils from four major ecosystem types in arctic Alaska. Total free amino acid concentrations were several-fold higher than ammonium (the major form of inorganic nitrogen) in water extracts of soils. The dominant free amino acids in these soils were glycine, aspartic acid, glutamic acid, serine, and arginine. Concentrations of total amino acids ranged 5-fold across communities, being highest in tussock tundra and lowest in wet meadows. Incubation experiments indicate that the turnover of amino acids is rapid, which suggests high rates of gross nitrogen mineralization in these soils. The high concentrations and dynamic nature of soil free amino acids suggest that this nitrogen pool is a significant component of nitrogen cycling in these tundra ecosystems.     

Variability in the chemistry of estuarine plants and its effect on feeding by Canada geese

Summary We investigated the influence of interspecific and seasonal variations in plant chemistry on food choices by adult and gosling Canada Geese, Branta canadensis, on Cape Cod, Massachusetts. The geese fed primarily on the abundant marsh grasses, Spartina spp., and rushes, Juncus gerardi, early in the growing season and switched to a greater dependence on eelgrass, Zostera marina, later. Forbs were generally avoided all season even when growing within patches of abundant species. The avoidance of forbs was related to their low abundance and their high concentrations of deterrent secondary metabolites. Differences in plant chemistry also determined the switch from marsh graminoids to Z. marina during the growing season. Marsh grasses were higher than Z. marina in nitrogen, particularly in the spring when the nitrogen requirement of geese is especially high. Z. marina was a better source of soluble carbohydrates and was the preferred food during the summer when the need to build up energy reserves may be more critical to geese than protein intake. Goslings, which require a diet higher in nitrogen than do adults, fed on marsh graminoids later into the growing season than the adults. The nitrogen content of the diets of goslings was significantly higher than that available to them in the plants, indicating that they selected for introgen. The diets of non-breeding adults in the spring and all geese in mid summer closely reflected the nutrient content of the plants. The diet of breeding adults was more similar to that of their goslings than to that of non-breeding adults. The effects of plant chemistry and the nutritional needs of geese on food choices were modified by the need to select a safe feeding site.     

The role of lesser snow geese as nitrogen processors in a sub-arctic salt marsh

Summary Ammonia volatilization losses from faeces of Lesser Snow Geese were measured during the summer of 1987 on the salt-marsh flats at La Pérouse Bay. Amounts of ammonia volatilized increased with increasing ambient temperature, and ranged from 1.0 to 15.1 mg N per 100 mg of nitrogen present as soluble ammonium ions at the start of the 8-h experiment. Using estimates of faecal deposition reported previously, the annual loss via volatilization was estimated at 0.08 g N m^-2, or 7.9% of the nitrogen present in goose faeces. Percent change in soluble ammonium ions in fresh faeces after 8 h ranged from -51.1% to +41.1%, indicating that net mineralization of organic nitrogen occurred in some of the faeces. Microbial respiration of fresh goose faeces increased exponentially with temperature. However, variable rates of net mineralization per unit rate of respiration indicated that the substrate quality affected microbial immobilization and thus net nitrogen mineralization. In feeding experiments, captive goslings grazed different types of vegetation, each with distinctive nutritional qualities. Forage quality had significant effects on goose feeding behavior and subsequent rates of nitrogen mineralization in fresh faeces. Net nitrogen mineralization rates in faeces from geese which grazed the three vegetation types ranged from 1.31 to 4.97 mg NH ₄ ⁺ –N g_DW ^-1 24 h^-1. Because plant growth in this salt marsh is nitrogen-limited, where swards are grazed, mineralization of organic faecal nitrogen represents an essential link in the maintenance of the flow of nitrogen into the sediments and the sustained growth of vegetation at a time when most required by the geese.     

Interactions in the uptake of amino acids, ammonium and nitrate ions in the Arctic salt-marsh grass, Puccinellia phryganodes

The uptake of amino acids and inorganic nitrogen by roots of Puccinellia phryganodes was examined to assess the potential contribution of soluble organic nitrogen to plant nitrogen uptake in Arctic coastal marshes, where free amino acids constitute a substantial fraction of the soil‐soluble N pool. Short‐term excised root uptake experiments were performed using tillers grown hydroponically under controlled conditions in the field. The percentage reductions in ammonium uptake at moderate salinity (150 mm NaCl) compared with uptake at low salinity (50 mm NaCl) were double those of glycine, but glycine uptake was more adversely affected than ammonium uptake by low temperatures. Glycine uptake was higher at pH 5·7 than at pH 7·0 or 8·2. The glycine uptake was up‐regulated in response to glycine, whereas ammonium uptake was up‐regulated in response to ammonium starvation. Nitrate uptake was strongly down‐regulated when tillers were grown on either ammonium or glycine. In contrast to N‐starved roots, which absorbed ammonium ions more rapidly than glycine, the roots grown on glycine, ammonium and nitrate and not N‐starved prior to uptake absorbed glycine as rapidly as ammonium and nitrate ions combined. Overall, the results indicate that amino acids are probably an important source of nitrogen for P. phryganodes in Arctic coastal marshes.     

The nature of plant adaptations to salinity stress has trophic consequences

In different ecosystems herbivores highly prefer particular plant species. This is often explained in a stoichiometric framework of nutrient‐based plant adaptations to herbivory. We hypothesize that such super‐palatability can also arise as an evolutionary by‐product of osmoregulatory adaptations of plants to stressful environmental conditions, as salinity, drought and cold. Here, we investigate in a coastal salt marsh why some plant species are highly preferred by migratory brent geese Branta bernicla bernicla in spring while others are avoided. This salt marsh is an important spring staging site for the geese. Sufficient energy storage in a short period is critical to enable their northward migration to Siberia and subsequent reproduction. We test if geese prefer plants that balance their internal osmotic potential with the saline environment through energy‐rich soluble sugars over plant species that use (compartmentalized) salts for this. We find that plant nitrogen and acid detergent fiber content, classic predictors of herbivore preferences, poorly explain which plants the geese prefer. Instead, plant species that are highly preferred by the geese adapt to salinity by high soluble sugar concentrations while avoided species do this by high plant salt concentrations. Thus, the type of osmoregulatory adaptation to stress displayed by different plant species is a good predictor for the food preference of geese on this salt marsh. We suggest that variation in other types of osmoregulation‐based stress adaptations, as plant cold adaptations in tundras and plant drought adaptations in savannas, have similar important consequences for trophic interactions.     

不同典型地带性土壤氮素分布特征及其影响因素

在野外取样的基础上,研究中国不同典型地带性土壤各形态氮素分布特征及其影响因素.结果表明: 垂直地带性土壤中0.5 mol·L^-1 K₂SO₄提取的提取态总氮、提取态有机氮、吸附氨基酸随取样点海拔的增加而显著增加,且提取态总氮、提取态有机氮和吸附氨基酸平均值都大于水平地带性土壤;水平地带性土壤各形态氮含量随土壤类型的不同而差异显著.土壤吸附氨基酸含量是游离氨基酸的5倍,占提取态有机氮百分比为21.1%,表明吸附氨基酸可能作为土壤有机氮库的一种重要存在形态.相关分析结果表明,垂直地带性土壤中提取态总氮、提取态有机氮、铵态氮、氨基酸态氮均与有机质、全氮呈显著正相关(r=0.57～0.93,P<0.05),但与pH、硝态氮呈显著负相关(r=-0.37～-0.91,P<0.05);水平地带性土壤pH与提取态总氮、硝态氮、有机质、全氮、碱解氮及盐基离子(K⁺、Ca²⁺、Mg²⁺)呈显著正相关(r=0.36～0.85,P<0.05),与铵态氮、氨基酸态氮呈显著负相关(r=-0.39～-0.81,P<0.05).     

Vegetation cover regulates the quantity,quality and temporal dynamics of dissolved organic carbon and nitrogen in Antarctic soils

Populations of the two native Antarctic vascular plant species (Deschampsia antarctica and Colobanthus quitensis) have expanded rapidly in recent decades, yet little is known about the effects of these expansions on soil nutrient cycling. We measured the concentrations of dissolved organic carbon (DOC) and nitrogen (DON), amino acids and inorganic N in soils under these two vascular plant species, and under mosses and lichens, over a growing season at Signy Island in the maritime Antarctic. We recorded higher concentrations of nitrate, total dissolved nitrogen, DOC, DON and free amino acids in soil under D. antarctica and C. quitensis than in lichen or moss dominated soils. Each vegetation cover gave a unique profile of individual free amino acids in soil solution. Significant interactions between soil type and time were found for free amino acid concentrations and C/N ratios, indicating that vascular plants significantly change the temporal dynamics of N mineralization and immobilization. We conclude that D. antarctica and C. quitensis exert a significant influence over C and N cycling in the maritime Antarctic, and that their recent population expansion will have led to significant changes in the amount, type and rate of organic C and N cycling in soil.     

Salt marshes along the coast of The Netherlands

The area of salt marshes does no longer increase. The recent erosion coincides with a rise in MHT-level in the last 25 years. Despite the decrease in area, sedimentation continues, especially in the lower salt marsh, which acts as a sink of nitrogen. Assimilation and mineralization of nitrogen are in balance in most plant communities along the gradient from lower to higher salt marshes. Mineralization of nitrogen increases towards the higher salt marsh, whereas the above-ground production and the mean nitrogen content of plants decrease. There is a positive correlation between quality of food plants in salt marshes and breeding success of Brent geese in the arctic tundra. Sedimentation on mainland salt marshes can compensate for the expected sea level rise. This is not the case for island salt marshes, if the relative sea level rise is more than 0.5–1.0 cm yr⁻¹. The natural succession on salt marshes results in an accumulation of organic material, which is related to the dominance of single plant species. It is not clear to which extent this process is enhanced by eutrophication from acid deposition and seawater. Human exploitation of unprotected salt marshes is old and heavy in the system of mound settlements. Reclamation rates by dikes in the last centuries were higher than the rate of area increase. Grazing by cattle as a management practice results in both a higher plant species-richness and community diversity than abandoning; hay-making is intermediate, but shows less structural diversity than grazing with low stocking density. The invertebrate fauna is favoured by a short period of abandoning, but eventually characteristic salt marsh invertebrates are replaced by inland species. Many bird species prefer grazed salt marshes. The final section gives some perspectives. Provided that no further embankments take place the optimal nature management option for plants and animals is a vegetation pattern, which includes areas with a low canopy (grazed) and areas with a tall canopy.     

Dynamics of dissolved organic nitrogen in subalpine Castle Lake,California

Dissolved organic nitrogen (DON) concentrations were measured in meso-oligotrophic Castle Lake, California during the ice-free season in 1982, 1983 and 1984. No consistent relationships were found between DON and particulate-N, primary productivity rates or chlorophyll concentrations. However, increases in DON concentrations were observed in the early growing season of 1982 and 1984 when water temperature was rising and a diatom bloom was senescing. DON increased at a high rate (0.31 mg atom N m^–3 day^–1), and then rapidly disappeared. Sediment released appared to be the most important source of DON. Dissolved free amino acids were always less than 7.5% of the DON pool, and did not vary in the composition of specific amino acids during the growing season.     

Responses of soil inorganic nitrogen to increased temperature and plant removal during the growing season in a Sibiraea angustata scrub ecosystem of eastern Qinghai-Xizang Plateau北大核心CSCD

Aims Little information has been available on the soil nitrogen transformation process of alpine scrubland under global warming and changing climate. This study aimed at clarifying seasonal dynamics of the soil nitrate and ammonium contents and their responses to increased temperature under different plant treatments. Methods We conducted a field experiment including two plant treatments (removal- or unremoval-plant) subjected to two temperature conditions (increased temperature or control) in Sibiraea angustata scrub ecosystem on the eastern Qinghai-Xizang Plateau. The contents of soil nitrate and ammonium were measured at the early, middle and late growing seasons. Important findings The results showed that soil nitrate and ammonium contents exhibited obvious seasonal dynamics. Throughout the entire growing season, the soil nitrate contents increased firstly and then decreased, while the soil ammonium contents increased continually. Particularly, in the early and middle growing season, the soil nitrate contents were significantly higher than those of ammonium, regardless of increased temperature and plant treatments; however, in the late growing season, the soil nitrate contents were significantly lower than those of ammonium. These results implied that soil nitrification was the major process of soil nitrogen transformation in the early and middle growing season; soil ammonification contributed mostly to soil nitrogen transformation in the late growing season. Furthermore, different responses of soil nitrate and ammonium contents to increased temperature and plant removal treatments were observed at the different stages in the growing season. The effects of increased temperature on soil nitrate contents mainly occurred in the middle and late growing season, but the effects varied with plant treatments. Increased temperature only significantly increased soil ammonium contents in the unremoval-plant plots during the middle growing season. The effects of plant treatments on soil nitrate contents only occurred in the control plots (controlled temperature). Plant removal only increased soil nitrate contents in the early and middle growing season, but significantly decreased soil nitrate contents in the late growing season. Plant removal significantly decreased soil ammonium contents in the increased temperature plots during the middle growing season. Probably, in the early and middle growing season, scrub vegetation mainly absorbed soil nitrate and the absorption process was not affected by increased temperature. These results would increase our understanding of the soil nitrogen cycling process in these alpine scrub ecosystems under global warming and changing climate. © 2018 Editorial Office of Chinese Journal of Plant Ecology. All rights reserved.     

Variation in amino acid content and its relationship to nitrogen content and growth rate in Ulva ohnoi (Chlorophyta)

To evaluate the quantitative and qualitative changes in amino acids related to internal nitrogen content and growth rate of Ulva ohnoi, the supply of nitrogen to outdoor cultures of the seaweed was manipulated by simultaneously varying water nitrogen concentrations and renewal rate. Both internal nitrogen content and growth rate varied substantially, and the quantitative and qualitative changes in amino acids were described in the context of three internal nitrogen states: nitrogen‐limited, metabolic, and luxury. The nitrogen limited state was defined by increases in all amino acids with increasing nitrogen content and growth up until 1.2% internal nitrogen. The metabolic nitrogen state was defined by increases in all amino acids with increasing internal nitrogen content up to 2.6%, with no increases in growth rate. Luxury state was defined by internal nitrogen content above 2.6%, which occurred only when nitrogen availability was high but growth rates were reduced. In this luxury circumstance, excess nitrogen was accumulated as free amino acids, in two phases. The first phase was distinguished by a small increase in the majority of amino acids up to ≈3.3% internal nitrogen, and the second by a large increase in glutamic acid, glutamine, and arginine up to 4.2% internal nitrogen. These results demonstrate that the relationship between internal nitrogen content and amino acid quality is dynamic but predictable, and could be used for the selective culture of seaweeds.     

Morphogenesis and free amino acid composition of the ascomycete Sphaerostilbe repens as influenced by nitrogen and calcium

Abstract Sphaerostilbe repens utilizes nitrate and ammonium as nitrogen sources. Differentiation of mycelium into rhizomorphs and coremia was reduced in the presence of nitrate and completely inhibited in the absence of calcium. The most abundant free amino acids were, in decreasing order: alanine, glutamine, glutatomic acid, serine, aspartic acid, γ-aminobutyric acid, arginine and threonine. These compounds represented 90% of the total amino acid pool.
The free amino acid composition did not vary with cultural conditions although concentrations of individual amino acids differed. In ammonium-grown cells, γ-aminobutyric acid increased in concentration and glutamate, aspartate and alanine decreased. Calcium-deficient media reduced amino acid concentrations, especially of arginine and ornithine. Amino acid contents increased during the growth period and were higher in rhizomorphs than in vegetative mycelia.     

Bio-available amino acids and mineral nitrogen forms in soil of moderately mown and abandoned mountain meadows

Summary. The abandonment of traditional mowing methods of mountain meadows in the Czech Republic at the end of the last century has resulted in secondary re-colonization of these areas. Altered accumulation of plant biomass resulted in a deceleration of N turnover. A mountain meadow may be regarded as a N-limited ecosystem in which plant nutrition is dependent on direct uptake of soil amino acids. The composition and distribution of ammonium ions, nitrate ions and the 16 bio-available proteinaceuous amino acids were investigated in the top 7 cm of the Ah horizon of a Gleyic Luvisol in a long-term moderately mown meadow and an eleven year old, abandoned or uncut meadow. Ammonium N has a dominant role in both ecosystems. The moderately mown meadow showed accelerated N-turnover and higher net ammonization. The plant community showed a dependence on this form. Plant utilization of nitrates and amino acids appeared to be negligible. The uncut or abandoned meadow showed net ammonization from May (start of the experiment) through August, after which plant N-uptake consisted only of amino acids due to microbial immobilization. The release of bio-available nitrogen from spring until the beginning of summer in the Ah horizon was too low to explain total plant N-uptake. Glutamic acid, arginine and aspartic acids had the highest concentrations of any of the amino acids analyzed. Authors’ address: Pavel Formánek, Department of Geology and Pedology, Mendel University of Agriculture and Forestry, Zemedelska 3, 613 00 Brno, Czech Republic     

The effect of ammonium and nitrate nitrogen sources on copper uptake and amino acid status of cereals

Summary The effects of different nitrogen sources (NH₄, NO₃, and NH₄ NO₃) on the uptake of copper by wheat and barley growing in solution culture were compared in three experiments. Both the copper concentration and weight gain of shoots and roots were found to decrease in the order NO₃>NH₄ NO₃>NH₄ irrespective of the solution copper concentration. Ammonium nitrogen was also found to decrease the copper concentration of wheat grown on a copper deficient soil compared with a nitrate source of nitrogen. Increasing concentrations of ammonium ions in solution culture caused ammonium toxicity and reduced both plant copper concentrations and vegetative yield. Biochemical investigations using paper chromatography revealed that the amino acid asparagine was the major detoxification product of ammonia in wheat. Copper deficient plants were found to have elevated levels of amino acids compared with controls, irrespective of the nitrogen source.     

Assimilation of ammonium ions and reutilization of nitrogen in rice (Oryza sativa L.)

A major source of inorganic nitrogen for rice plants grown in paddy soil is ammonium ions. The ammonium ions are actively taken up by the roots via ammonium transporters and subsequently assimilated into the amide residue of glutamine (Gln) by the reaction of glutamine synthetase (GS) in the roots. The Gln is converted into glutamate (Glu), which is a central amino acid for the synthesis of a number of amino acids, by the reaction of glutamate synthase (GOGAT). Although a small gene family for both GS and GOGAT is present in rice, ammonium-dependent and cell type-specific expression suggest that cytosolic GS1;2 and plastidic NADH-GOGAT1 are responsible for the primary assimilation of ammonium ions in the roots. In the plant top, approximately 80% of the total nitrogen in the panicle is remobilized through the phloem from senescing organs. Since the major form of nitrogen in the phloem sap is Gln, GS in the senescing organs and GOGAT in developing organs are important for nitrogen remobilization and reutilization, respectively. Recent work with a knock-out mutant of rice clearly showed that GS1;1 is responsible for this process. Overexpression studies together with age- and cell type-specific expression strongly suggest that NADH-GOGAT1 is important for the reutilization of transported Gln in developing organs. The overall process of nitrogen utilization within the plant is discussed.     

Oligopeptide assimilation and transport by Oenococcus oeni

Aims: Oenococcus oeni is a slow‐growing wine bacterium with a low growth yield. It thrives better on complex nitrogen sources than on free amino‐acid medium. We aimed to characterize the oligopeptide use of this micro‐organism. Methods and Results: Several peptides of two to eight amino‐acid residues were able to provide essential amino acids. The disappearance of various peptides from extracellular medium was assessed with whole cells. Initial rates of utilization varied with the peptide, and free amino acids were released into the medium. Conclusions: Oenococcus oeni was able to transport the oligopeptides with two to five amino‐acid residues tested and to hydrolyse them further. Significance and Impact of the Study: This study has clear implications for the relationship between wine nitrogen composition and the ability of O. oeni to cope with its environment.     

Alternative criteria for assessing nutrient limitation of a wetland macrophyte (Peltandra virginica (L.) Kunth)

Various nutrient incorporation and plant production parameters were measured to assess their relative usefulness in determining possible nutrient limitation of the wetland plant Peltandra virginica (L.) Kunth. From four stations located along a transect in a tidal freshwater marsh, we documented spatial differences in peak standing biomass of plants. Plant biomass was positively correlated with porewater concentrations of both ammonium and phosphate, but not with sediment concentrations of total nitrogen and phosphorus. Tissue nitrogen and phosphorus concentrations decreased significantly over the growing season, but there were no differences among plants from the four stations, and correlations between plant biomass and ratios of carbon to nitrogen and carbon to phosphorus were weak. Because in situ fertilization of plants had no effect on either peak biomass or tissue concentrations of nitrogen and phosphorus, growth of Peltandra was probably not nutrient limited. Other criteria did predict nitrogen or nitrogen and phosphorus limitation, however, demonstrating that application of parameters used by ecologists to support contentions of nutrient limitation can yield conflicting results. Assessment of nutrient limitation of primary producers may be an ambiguous and unnecessary task in some environments where these criteria are utilized.     

AAP1 transports uncharged amino acids into roots of Arabidopsis

Amino acids are available to plants in some soils in significant amounts, and plants frequently make use of these nitrogen sources. The goal of this study was to identify transporters involved in the uptake of amino acids into root cells. Based on the fact that high concentrations of amino acids inhibit plant growth, we hypothesized that mutants tolerating toxic levels of amino acids might be deficient in the uptake of amino acids from the environment. To test this hypothesis, we employed a forward genetic screen for Arabidopsis thaliana mutants tolerating toxic concentrations of amino acids in the media. We identified an Arabidopsis mutant that is deficient in the amino acid permease 1 (AAP1, At1g58360) and resistant to 10 mm phenylalanine and a range of other amino acids. The transporter was localized to the plasma membrane of root epidermal cells, root hairs, and throughout the root tip of Arabidopsis. Feeding experiments with [(14)C]-labeled neutral, acidic and basic amino acids showed significantly reduced uptake of amino acids in the mutant, underscoring that increased tolerance of aap1 to high levels of amino acids is coupled with reduced uptake by the root. The growth and uptake studies identified glutamate, histidine and neutral amino acids, including phenylalanine, as physiological substrates for AAP1, whereas aspartate, lysine and arginine are not. We also demonstrate that AAP1 imports amino acids into root cells when these are supplied at ecologically relevant concentrations. Together, our data indicate an important role of AAP1 for efficient use of nitrogen sources present in the rhizosphere.