Effects of air pollutant-temperature interactions on mineral-N dynamics and cation leaching in reciplicate forest soil transplantation experiments

Increased emissions of nitrogen compounds have led to atmosphericdeposition to forest soils exceeding critical loads of N overlarge parts of Europe. To determine whether the chemistry offorest soils responds to changes in throughfall chemistry, intactsoil columns were reciprocally transplanted between sites, withdifferent physical conditions, across a gradient of N and Sdeposition in Europe.The transfer of a single soil to the various sites affected itsnet nitrification. This was not simply due to the nitrificationof different levels of N deposition but was explained bydifferences in physical climates which influenced mineralizationrates. Variation in the amount of net nitrification between soiltypes at a specific site were explained largely by soil pH.Within a site all soil types showed similar trends in netnitrification over time. Seasonal changes in net nitrificationcorresponds to oscillations in temperature but variable time lagshad to be introduced to explain the relationships. WithArrhenius law it was possible to approximate gross nitrificationas a function of temperature. Gross nitrification equalled netnitrification after adaptation of the microbial community oftransplanted soils to the new conditions. Time lags, andunderestimates of gross nitrification in autumn, were assumed tobe the result of increased NH ₄ ⁺ availability due either tochanges in the relative rates of gross and net N transformationsor to altered soil fauna-microbial interactions combined withimproved moisture conditions.Losses of NO ₃ ^- were associated with Ca²⁺and Mg²⁺ in non-acidified soil types and with losses ofAl³⁺ in the acidified soils. For single soils the ionequilibrium equation of Gaines-Thomas provided a useful approximationof Al³⁺ concentrations in the soil solution as a functionof the concentration of Ca²⁺. The between site deviationsfrom this predicted equilibrium, which existed for single soils, couldbe explained by differences in throughfall chemistry which affectedthe total ionic strength of the soil solution.The approach of reciprocally transferring soil columnshighlighted the importance of throughfall chemistry, interactingwith the effect of changes in physical climate on forest soilacidification through internal proton production, in determiningsoil solution chemistry. A framework outlining the etiology offorest die-back induced by nitrogen saturation is proposed.     

Quantification of soil carbon inputs under elevated CO2: C3 plants in a C4 soil

The objective of this investigation was to quantify the differences in soil carbon stores after exposure of birch seedlings (Betula pendula Roth.) over one growing season to ambient and elevated carbon dioxide concentrations. One-year-old seedling of birch were transplanted to pots containing C₄ soil derived from beneath a maize crop, and placed in ambient (350 L L^–1) and elevated (600 L L^–1) plots in a free-air carbon dioxide enrichment (FACE) experiment. After 186 days the plants and soils were destructively sampled, and analysed for differences in root and stem biomass, total plant tissue and soil C contents and ¹³C values. The trees showed a significant increase (+50%) in root biomass, but stem and leaf biomasses were not significantly affected by treatment. C isotope analyses of leaves and fine roots showed that the isotopic signal from the ambient and elevated CO₂ supply was sufficiently distinct from that of the C₄ soil to enable quantification of net root C input to the soil under both ambient and elevated CO₂. After 186 days, the pots under ambient conditions contained 3.5 g of C as intact root material, and had gained an additional 0.6 g C added to the soil through root exudation/turnover; comparable figures for the pots under elevated CO₂ were 5.9 g C and 1.5 g C, respectively. These data confirm the importance of soils as an enhanced sink for C under elevated atmospheric CO₂ concentrations. We propose the use of C₄ soils in elevated CO₂ experiments as an important technique for the quantification of root net C inputs under both ambient and elevated CO₂ treatments.     

Lipid content and carbon assimilation in Collembola: implications for the use of compound-specific carbon isotope analysis in animal dietary studies

In an effort to understand the relationships between both the lipid content and ¹³C values of Collembola and their diet, isotopically labelled (C₃ and C₄) bakers yeasts were cultured and fed to two Collembolan species, Folsomia candida and Proisotoma minuta. The fatty acid composition of Collembola generally reflected that of the diet with the addition of the polyunsaturated components 18:2(n-6), 20:4(n-6) and 20:5(n-3), which appeared to be biosynthesised by the Collembola. Whilst ergosterol was the only sterol detected in the yeast diets, only cholesterol was detected in Collembola, and although the ¹³C values of diet and consumer sterols differed by >2, the ¹³C values indicated that cholesterol was derived entirely from dietary sterol. The bulk ¹³C values of Collembola were similar to those of the diets, but fatty acid ¹³C values did not necessarily reflect those of the dietary fatty acids, indicating significant de novo biosynthesis of fatty acids within Collembola. Switching the Collembola from C₃ to C₄ yeast enabled the determination of the rates of incorporation of dietary carbon into Collembolan lipids, and showed that half-lives of the incorporation of dietary carbon varied between 1.5 and 5.8 days at 20°C. Cholesterol exhibited the slowest rate of incorporation in both species, while bulk carbon in F. candida possessed an intermediate rate. These results demonstrate that an understanding of the sources of isotopic fractionation and the role of biochemistry in regulating the ¹³C values of individual compounds is important in the application of compound-specific isotopic analysis to the study of animal trophic activities.     

Environmental and Vegetation Drivers of Seasonal CO2 Fluxes in a Sub-arctic Forest–Mire Ecotone

Unravelling the role of structural and environmental drivers of gross primary productivity (GPP) and ecosystem respiration (R _eco) in highly heterogeneous tundra is a major challenge for the upscaling of chamber-based CO₂ fluxes in Arctic landscapes. In a mountain birch woodland-mire ecotone, we investigated the role of LAI (and NDVI), environmental factors (microclimate, soil moisture), and microsite type across tundra shrub plots (wet hummocks, dry hummocks, dry hollows) and lichen hummocks, in controlling net ecosystem CO₂ exchange (NEE). During a growing season, we measured NEE fluxes continuously, with closed dynamic chambers, and performed multiple fits (one for each 3-day period) of a simple light and temperature response model to hourly NEE data. Tundra shrub plots were largely CO₂ sinks, as opposed to lichen plots, although fluxes were highly variable within microsite type. For tundra shrub plots, microsite type did not influence photosynthetic parameters but it affected basal (that is, temperature-normalized) ecosystem respiration (R ₀). PAR-normalized photosynthesis (P ₆₀₀) increased with air temperature and declined with increasing vapor pressure deficit. R ₀ declined with soil moisture and showed an apparent increase with temperature, which may underlie a tight link between GPP and R _eco. NDVI was a good proxy for LAI, maximum P ₆₀₀ and maximum R ₀ of shrub plots. Cumulative CO₂ fluxes were strongly correlated with LAI (NDVI) but we observed a comparatively low GPP/LAI in dry hummocks. Our results broadly agree with the reported functional convergence across tundra vegetation, but here we show that the role of decreased productivity in transition zones and the influence of temperature and water balance on seasonal CO₂ fluxes in sub-Arctic forest–mire ecotones cannot be overlooked.     

Acidifying processes and acid-base reactions in forest soils reciprocally transplanted along a European transect with increasing pollution

Forest ecosystems are currently beingexposed to changes in chemical inputs and it issuggested that physical climate is also changing. Anovel approach has been used to study the effects ofionic inputs and climatic conditions on forest soilsby reciprocally exchanging lysimeters containingundisturbed soil columns beween four forest sites inEurope. The soil columns contained no living roots andsimulated a clear cut situation. The soils chosenrepresented different stages of acidification and weretaken from sites along a transect of increasingexposure to acidic and nitrogen pollution. The purposeof the study was to quantify the reactions of soilswhen transferred to different environments. Elementbalances were used as an aggregated indicator todescribe the reaction of the soil. The input of protonsin local throughfall increased along the transect from0.01 kmol ha^-1 y^-1H⁺ at the unpolluted site up to 1.10 kmolha^-1 y^-1 at the most pollutedsite. Our results show that soil acidification always resultedfrom a combination of acid deposition and biologicaltransformations of nitrogen through nitrification ofimported ammonium, mineralized N, or stored N. Thebalances indicate that between 54% and 91%of the soil acidification resulted from nitrificationprocesses which were driven by a complex reaction whenclimatic and pollution conditions were changedsimultaneously. The combined changes in atmosphicinputs and climatic conditions, as expected withglobal change, may have serious consequences for soilacidfication and long term organic matter turnover.     

Respiration of the external mycelium in the arbuscular mycorrhizal symbiosis shows strong dependence on recent photosynthates and acclimation to temperature

* Although arbuscular mycorrhizal (AM) fungi are a major pathway in the global carbon cycle, their basic biology and, in particular, their respiratory response to temperature remain obscure. * A pulse label of the stable isotope (13)C was applied to Plantago lanceolata, either uninoculated or inoculated with the AM fungus Glomus mosseae. The extra-radical mycelium (ERM) of the fungus was allowed to grow into a separate hyphal compartment excluding roots. We determined the carbon costs of the ERM and tested for a direct temperature effect on its respiration by measuring total carbon and the (13)C:(12)C ratio of respired CO(2). With a second pulse we tested for acclimation of ERM respiration after 2 wk of soil warming. * Root colonization remained unchanged between the two pulses but warming the hyphal compartment increased ERM length. delta(13)C signals peaked within the first 10 h and were higher in mycorrhizal treatments. The concentration of CO(2) in the gas samples fluctuated diurnally and was highest in the mycorrhizal treatments but was unaffected by temperature. Heating increased ERM respiration only after the first pulse and reduced specific ERM respiration rates after the second pulse; however, both pulses strongly depended on radiation flux. * The results indicate a fast ERM acclimation to temperature, and that light is the key factor controlling carbon allocation to the fungus.     

Effects of enhanced atmospheric CO2 and nutrient supply on the quality and subsequent decomposition of fine roots of Betula pendula Roth. and Picea sitchensis (Bong.) Carr.

Fine root litter derived from birch (Betula pendula Roth.) and Sitka spruce (Picea sitchensis (Bong.) Carr.) plants grown under two CO₂ atmospheric concentrations (350 ppm and 600 ppm) and two nutrient regimes was used for decomposition studies in laboratory microcosms. Although there were interactions between litter type, CO₂/fertiliser treatments and decomposition rates, in general, an increase in the C/N ratio of the root tissue was observed for roots of both species grown under elevated CO₂ in unfertilized soil. Both weight loss and respiration of decomposing birch roots were significantly reduced in materials derived from enriched CO₂, whilst the decomposition of spruce roots showed no such effect. A parallel experiment was performed using Betula pendula root litter grown under different N regimes, in order to test the relationship between C/N ratio of litter and root decomposition rate. A highly significant (p<0.001) negative correlation between C/N ratio and root litter respiration was found, with an r²=0.97. The results suggest that the increased C/N ratio of plant tissues induced by elevated CO₂ can result in a reduction of decomposition rate, with a resulting increase in forest soil C stores.     

Decomposition of tree leaf litters grown under elevated CO2: Effect of litter quality

Ash (Fraxinus excelsior L.), birch (Betula pubescens Ehrh.), sycamore (Acer pseudoplatanus L.) and Sitka spruce (Picea sitchensis (Bong.) Carr.) leaf litters were monitored for decomposition rates and nutrient release in a laboratory microcosm experiment. Litters were derived from solar domes where plants had been exposed to two different CO₂ regimes: ambient (350 L L^-1 CO₂) and enriched (600 L L^-1 CO₂).Elevated CO₂ significantly affected some of the major litter quality parameters, with lower N, higher lignin concentrations and higher ratios of C/N and lignin/N for litters derived from enriched CO₂. Respiration rates of the deciduous species were significantly decreased for litters grown under elevated CO₂, and reductions in mass loss at the end of the experiment were generally observed in litters derived from the 600 ppm CO₂ treatment. Nutrient mineralization, dissolved organic carbon, and pH in microcosm leachates did not differ significantly between the two CO₂ treatments for any of the species studied. Litter quality parameters were examined for correlations with cumulative respiration and decomposition rates: N concentration, C/N and lignin/N ratios showed the highest correlations, with differences between litter types. The results indicate that higher C storage will occur in soil as a consequence of litter quality changes resulting from higher atmospheric concentrations of CO₂.Abbreviations CHO soluble carbohydrates - DOC dissolved organic carbon - HCel holocellulose - WTREM weight remaining