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.     

Plant community responses to nitrogen addition and increased precipitation: the importance of water availability and species traits

Global nitrogen (N) enrichment and changing precipitation regimes are likely to alter plant community structure and composition, with consequent influences on biodiversity and ecosystem functioning. Responses of plant community structure and composition to N addition and increased precipitation were examined in a temperate steppe in northern China. Increased precipitation and N addition stimulated and suppressed community species richness, respectively, across 6 years (2005–2010) of the manipulative experiment. N addition and increased precipitation significantly altered plant community structure and composition at functional groups levels. The significant relationship between species richness and soil moisture (SM) suggests that plant community structure is mediated by water under changing environmental conditions. In addition, plant height played an important role in affecting the responses of plant communities to N addition, and the effects of increased precipitation on plant community were dependent on species rooting depth. Our results highlight the importance and complexity of both abiotic (SM) and biotic factors (species traits) in structuring plant community under changing environmental scenarios. These findings indicate that knowledge of species traits can contribute to mechanistic understanding and projection of vegetation dynamics in response to future environmental change.     

Available soil nitrogen in relation to fractions of soil nitrogen and other soil properties

The available N of 27 soils from England and Wales was assessed from the amounts of N taken up over a 6-month period by perennial ryegrass grown in pots under uniform environmental conditions. Relationships between availability and the distribution of soil N amongst various fractions were then examined using multiple regression. The relationship: available soil N (mg kg^–1 dry soil)=(N_min×0.672)+(N_inc×0.840)+(N_mom×0.227)–5.12 was found to account for 91% of the variance in available soil N, where N_min=mineral N, N_inc=N mineralized on incubation and N_mom=N in macro-organic matter. The N mineralized on incubation appeared to be derived largely from sources other than the macro-organic matter because these two fractions were poorly correlated. When availability was expressed in terms of available organic N as % of soil organic N (N_ao) the closest relationship with other soil characteristics was: N_ao=[N_inc×(1.395–0.0347×CN_mom]+[N_mom×0.1416], where CN_mom=CN ratio of the macro-organic matter. This relationship accounted for 81% of the variance in the availability of the soil organic N.The conclusion that the macro-organic matter may contribute substantially to the available N was confirmed by a subsidiary experiment in which the macro-organic fraction was separated from about 20 kg of a grassland soil. The uptake of N by ryegrass was then assessed on two subsamples of this soil, one without the macro-organic matter and the other with this fraction returned: uptake was appreciably increased by the macro-organic matter.     

Effects of nitrogen fertilization of grapefruit trees on soil acidification and nutrient availability in a Riviera fine sand

Nitrogen (N) fertilizer applied in the NH4+ form results in some degree of soil acidification, which could influence nutrient availability to plants and nutrient losses through leaching. Effects of various N rates (0 – 168 kg N ha-1 yr-1) on soil acidification and nutrient availability were investigated in a Riviera fine sand with 26-year-old white Marsh grapefruit (Citrus paradisi MacFadyen) trees. Soil pH significantly decreased with increasing NH4–N rates. Application of 112 kg N ha-1 yr-1 for four years decreased the pH by 0.7 to 1.7 unit. Soil acidification was greater when the NH4+ form of N fertilizer was applied as dry soluble granular material compared to fertigation or controlled release forms. The marked effect of NH4–N fertilization on the pH of the Riviera fine sand was due to its low buffering capacity. Soil acidification increased the concentration of extractable Fe and P but decreased that of K, Zn and Mn. Soil pH was positively correlated with concentration of Ca, but negatively with concentrations of Fe, Mn and Zn in six-month-old spring flush leaves of the grapefruit trees. Leaf P concentrations, however, were poorly correlated with soil pH. This study also demonstrated an increase in leaching of P and K below the grapefruit trees rootzone with a decrease in soil pH.     

Links between plant community composition,soil organic matter quality and microbial communities in contrasting tundra habitats

Plant communities, soil organic matter and microbial communities are predicted to be interlinked and to exhibit concordant patterns along major environmental gradients. We investigated the relationships between plant functional type composition, soil organic matter quality and decomposer community composition, and how these are related to major environmental variation in non-acid and acid soils derived from calcareous versus siliceous bedrocks, respectively. We analysed vegetation, organic matter and microbial community compositions from five non-acidic and five acidic heath sites in alpine tundra in northern Europe. Sequential organic matter fractionation was used to characterize organic matter quality and phospholipid fatty acid analysis to detect major variation in decomposer communities. Non-acidic and acidic heaths differed substantially in vegetation composition, and these disparities were associated with congruent shifts in soil organic matter and microbial communities. A high proportion of forbs in the vegetation was positively associated with low C:N and high soluble N:phenolics ratios in soil organic matter, and a high proportion of bacteria in the microbial community. On the contrary, dwarf shrub-rich vegetation was associated with high C:N and low soluble N:phenolics ratios, and a high proportion of fungi in the microbial community. Our study demonstrates a strong link between the plant community composition, soil organic matter quality, and microbial community composition, and that differences in one compartment are paralleled by changes in others. Variation in the forb-shrub gradient of vegetation may largely dictate variations in the chemical quality of organic matter and decomposer communities in tundra ecosystems. Soil pH, through its direct and indirect effects on plant and microbial communities, seems to function as an ultimate environmental driver that gives rise to and amplifies the interactions between above- and belowground systems. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Complex vegetation responses to soil disturbances in mountain grassland

We studied vegetation responses to disturbances originated by ants and voles in subalpine grasslands in the Eastern Pyrenees. We compared the effects of these small-scale disturbances with those of a large-scale disturbance caused by ploughing. We wanted to know if these soil disturbances promoted species richness through the existence of a specific guild of plants colonizing these areas, and if this guild was the same for all soil disturbances, independently of their extent. In general, grassland vegetation seemed to recover relatively quickly from soil-displacement disturbances, and the effects could be scaled up in time and space in terms of species richness and composition. Vole mound composition was similar to that in the surrounding grassland, suggesting that mounds were rapidly colonized by the neighbouring vegetation. Vegetation composition differed between the grassland and the ant mounds. Grasses and erect dicots coped well with repeated disturbance, while rosette-forming species and sedges were very sensitive to it. Landscape processes could be important to understanding recolonization. Species from xeric grasslands were found in mesic grasslands when disturbed by ploughing and on the tops of active ant mounds. Furrows in mesic grasslands recovered well, but decades after disturbance showed long persistence of some xeric species and increased species richness compared to terraces, while xeric grasslands showed decreased richness. This suggests that, because of those disturbances, within-habitat diversity was increased, although landscape diversity was not. However, specific disturbances showed idiosyncratic effects, which could enhance the species richness globally. In ant-affected areas, the grassland itself showed the highest plant species richness, partially associated to the presence of some species with elaiosomes not, or only rarely, found in adjacent grasslands without ant mounds. Therefore, soil disturbances occurring at different spatial scales contributed to complexity in vegetation patterns in addition to abiotic factors and grazing. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Nomenclature of the species follows Tutin et al. (1964–1980) and Bolòs et al. (1993).     

Plant responses to fertilization and exclusion of grazers on an arctic tundra heath

This study investigated the impacts of fertilization and grazing by Norwegian lemmings (Lemmus lemmus), grey‐sided voles (Clethrionomys rufocanus), and reindeer (Rangifer tarandus) on a diverse tundra plant community dominated by deciduous shrubs. Four out of eight study areas, having a size of 2500 m² each, were fertilized with a N‐P‐K fertilizer and four areas served as unfertilized controls. Two types of exclosures were used within each study area, one to exclude solely reindeer, and one to exclude both rodents and reindeer. Open, grazed plots served as controls. During 5 years following the fertilization event the changes in vegetation inside and outside the exclosures were monitored using a point frequency method. The densities of rodents on the fertilized and unfertilized areas were investigated by live trapping and by counting nests of overwintering individuals. Reindeer do not graze on the study area during the growing season but migrate through this area in autumn and spring. Fertilization increased the abundance of vascular plants while grazing by reindeer and rodents decreased the abundance of vascular plants significantly on both fertilized and unfertilized areas. Rodents preferred clearly the fertilized areas during winter, decreasing the abundance of Vaccinium myrtillus and Vaccinium vitis‐idaea, while very little grazing occurred during summer. Graminoids showed the strongest positive response to fertilization and dominated the plant community on ungrazed plots, while winter grazing by both reindeer and rodents significantly decreased the abundance of graminoids. Deciduous shrubs (Betula nana, Vaccinium myrtillus) increased slightly but significantly due to fertilization and evergreen dwarf shrubs showed no response to fertilization. However, the use of functional growth forms for predicting the responses of nutrient enrichment and grazing must be questioned, as responses to fertilization as well as preferences by herbivores were shown to be species‐specific rather than uniform within functional groups based on plant growth forms.     

Nutrient limitation of phytoplankton production in Alaskan Arctic foothill lakes

We used 54 enrichment bioassays to assess nutrient limitation (N, P) of ¹⁴C uptake by natural phytoplankton assemblages in 39 lakes and ponds in the Arctic Foothills region of Alaska. Our purpose was to categorize phytoplankton nutrient status in this under-represented region of North America and to improve our ability to predict the response of primary production to anticipated anthropogenically mediated increases in nutrient loading. Experiments were performed across several watersheds and included assays on terminal lakes and lakes occupying various positions in chains (lakes in series within a watershed and connected by streams). In total, 89% (48 of 54) of the bioassays showed significant stimulation of ¹⁴C primary production by some form of nutrient addition relative to unamended controls. A significant response was observed following enrichment with N and P, N alone and P alone in 83, 35 and 22% of the bioassays, respectively. In experiments where N and P proved stimulatory, the influence of N alone was significantly greater than the influence of P alone. Overall, the data point to a greater importance for N than P in regulating phytoplankton production in this region. The degree of response to N and P enrichment declined as the summer progressed and showed no relationship to irradiance or water temperature, suggesting secondary limitation by some micronutrient such as iron as the summer advanced. Phytoplankton nutrient status was often consistent across lakes within a watershed, suggesting that watershed characteristics influence nutrient availability. Lakes in this region will clearly show increased phytoplankton production in response to anthropogenic activities and anticipated changes in climate that will increase nutrient loading.     

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.     

Selecting appropriate forms of nitrogen fertilizer to enhance soil arsenic removal by Pteris vittata: a new approach in phytoremediation

Certain plant species have been shown to vigorously accumulate some metals from soil, and thus represent promising and effective remediation alternatives. In order to select the optimum forms of nitrogen (N) fertilizers for the arsenic (As) hyperaccumulator, Pteris vittata L., to maximize As extraction, five forms of N were added individually to different treatments to study the effect of N forms on As uptake of the plants under soil culture in a greenhouse. Although shoot As concentration tended to decrease and As translocation from root to shoot was inhibited, overall As accumulation was greater due to higher biomass when N fertilizer was added. Arsenic accumulation in plants with N fertilization was 100-300% more than in the plants without N fertilization. There were obvious differences in plant biomass and As accumulation among the N forms, i.e., NH4HCO3, (NH4)2S04, Ca(NO3)2, KNO3, urea. The total As accumulation in the plants grown in As-supplied soil, under different forms of N fertilizer, decreased as NH4HCO3>(NH4)2S04 > urea > Ca(NO3)2 >KNO3>CK. The plants treated with N and As accumulated up to 5.3-7.97 mg As/pot and removed 3.7-5.5% As from the soils, compared to approximately 2.3% of As removal in the control. NH4+ -N was apparently more effective than other N fertilizers in stimulating As removal when soil was supplied with As at initiation. No significant differences in available As were found among different forms of N fertilizer after phytoremediation. It is concluded that NH4+ -N was the preferable fertilizer for P. vittata to maximize As removal.     

Plant trait responses to grazing – a global synthesis

Herbivory by domestic and wild ungulates is a major driver of global vegetation dynamics. However, grazing is not considered in dynamic global vegetation models, or more generally in studies of the effects of environmental change on ecosystems at regional to global scale. An obstacle to this is a lack of empirical tests of several hypotheses linking plant traits with grazing. We, therefore, set out to test whether some widely recognized trait responses to grazing are consistent at the global level. We conducted a meta‐analysis of plant trait responses to grazing, based on 197 studies from all major regions of the world, and using six major conceptual models of trait response to grazing as a framework. Data were available for seven plant traits: life history, canopy height, habit, architecture, growth form (forb, graminoid, herbaceous legume, woody), palatability, and geographic origin. Covariates were precipitation and evolutionary history of herbivory. Overall, grazing favoured annual over perennial plants, short plants over tall plants, prostrate over erect plants, and stoloniferous and rosette architecture over tussock architecture. There was no consistent effect of grazing on growth form. Some response patterns were modified by particular combinations of precipitation and history of herbivory. Climatic and historical contexts are therefore essential for understanding plant trait responses to grazing. Our study identifies some key traits to be incorporated into plant functional classifications for the explicit consideration of grazing into global vegetation models used in global change research. Importantly, our results suggest that plant functional type classifications and response rules need to be specific to regions with different climate and herbivory history.     

Plant induced alteration in the rhizosphere and the utilisation of soil heterogeneity

Plant-soil interactions result in a special rhizosphere soil chemistry, differing from that of the bulk soil found only a few mm from the root. The aim of this study was to investigate adaptation mechanisms of herbs growing in acid soils through studying their rhizosphere chemistry in a greenhouse experiment and in a field study. Ten herbs were grown in acid soil (pH 4.2 in the soil solution) in the greenhouse. The concentrations of NO₃ ^-, SO₄ ^2-, phosphates, Ca²⁺, Mg²⁺, Mn²⁺, K⁺, Na⁺, NH₄ ⁺ and pH were analysed in soil solutions obtained by centrifugation. The general pattern found was a depletion of nutrients in the rhizosphere compared with their concentrations in the bulk soil. The pH increase (up to 0.7 units) in the rhizosphere soil appeared to be caused by plant uptake of NO₃ ^- (r²=0.88). The ion concentrations in the soil solution of the rhizosphere were dependent on plant species and biomass increase. Although species with a larger biomass and higher growth rates showed a higher degree of ion depletion (except for Na⁺, SO₄ ^2-) in the rhizosphere, there were also species specific responses. A field study of five herbs at five oak forest sites in Southern Sweden (Scania) was also carried out. In addition to the soil solution concentrations, the loss on ignition (LOI) and the concentrations of 0.1 M BaCl₂ extractable K⁺, Mg²⁺, Mn²⁺, Ca²⁺, and Al ions were measured. The amount of soil solution Al was determined as free ionic (quickly reacting) Al. For all species and sites, the LOI and the concentrations of exchangeable cations were higher in the rhizosphere than in the bulk soil, apparently due to the roots preferably growing at organic-rich microsites. The concentrations of the ions as measured in the centrifuged soil solution, were either higher in the rhizosphere than in the bulk soil or were the same in both, except for NO₃ ^- and quickly reacting Al. The lower concentrations of quickly reacting Al in the rhizosphere, compared with the bulk soil could indicate the uptake of Al by the plant or the exudation of complexing substances. The pH differences were only small and mostly non-significant. Plant-soil interactions and the ability of plants to utilise heterogeneity of the soil appear to be more important for plant growth in acid soils than recognised heretofore. Rhizosphere studies provide an important means of understanding plant strategies in acid soils. This revised version was published online in June 2006 with corrections to the Cover Date.     

Microbial biomass and nitrogen cycling responses to fertilization and litter removal in young northern hardwood forests

The influence of site fertility on soil microbial biomass and activity is not well understood but is likely to be complex because of interactions with plant responses to nutrient availability. We examined the effects of long-term (8 yr) fertilization and litter removal on forest floor microbial biomass and N and C transformations to test the hypothesis that higher soil resource availability stimulates microbial activity. Microbial biomass and respiration decreased by 20–30 % in response to fertilization. Microbial C averaged 3.8 mg C/g soil in fertilized, 5.8 mg C/g in control, and 5.5 mg C/g in litter removal plots. Microbial respiration was 200 µg CO₂-C g^–1 d^–1 in fertilized plots, compared to 270 µg CO₂-C g^–1 d^–1 in controls. Gross N mineralization and N immobilization did not differ among treatments, despite higher litter nutrient concentrations in fertilized plots and the removal of substantial quantities of C and N in litter removal plots. Net N mineralization was significantly reduced by fertilization. Gross nitrification and NO₃ ^– immobilization both were increased by fertilization. Nitrate thus became a more important part of microbial N cycling in fertilized plots even though NH₄ ⁺ availability was not stimulated by fertilization.Soil microorganisms did not mineralize more C or N in response to fertilization and higher litter quality; instead, results suggest a difference in the physiological status of microbial biomass in fertilized plots that influenced N transformations. Respiration quotients (qCO₂, respiration per unit biomass) were higher in fertilized plots (56 µg CO₂-C mg C^–1 d^–1) than control (48 µg CO₂-C mg C^–1 d ^–1) or litter removal (45 µg CO₂-C mg C^–1 d^–1), corresponding to higher microbial growth efficiency, higher proportions of gross mineralization immobilized, and lower net N mineralization in fertilized plots. While microbial biomass is an important labile nutrient pool, patterns of microbial growth and turnover were distinct from this pool and were more important to microbial function in nitrogen cycling.