Soil nitrogen transformations under Populus tremuloides, Betula papyrifera and Acer saccharum following 3 years exposure to elevated CO2 and O3

Increases in atmospheric CO₂ and tropospheric O₃ may affect forest N cycling by altering plant litter production and the availability of substrates for microbial metabolism. Three years following the establishment of our free‐air CO₂–O₃ enrichment experiment, plant growth has been stimulated by elevated CO₂ resulting in greater substrate input to soil; elevated O₃ has counteracted this effect. We hypothesized that rates of soil N cycling would be enhanced by greater plant productivity under elevated CO₂, and that CO₂ effects would be dampened by O₃. We found that elevated CO₂ did not alter gross N transformation rates. Elevated O₃ significantly reduced gross N mineralization and microbial biomass N, and effects were consistent among species. We also observed significant interactions between CO₂ and O₃: (i) gross N mineralization was greater under elevated CO₂ (1.0 mg N kg^?1 day^?1) than in the presence of both CO₂ and O₃ (0.5 mg N kg^?1 day^?1) and (ii) gross NH₄⁺ immobilization was also greater under elevated CO₂ (0.8 mg N kg^?1 day^?1) than under CO₂ plus O₃ (0.4 mg N kg^?1 day^?1). We used a laboratory ¹⁵N tracer method to quantify transfer of inorganic N to organic pools. Elevated CO₂ led to greater recovery of NH₄⁺‐¹⁵N in microbial biomass and corresponding lower recovery in the extractable NO₃^? pool. Elevated CO₂ resulted in a substantial increase in NO₃^?‐¹⁵N recovery in soil organic matter. We observed no O₃ main effect and no CO₂ by O₃ interaction effect on ¹⁵N recovery in any soil pool. All of the above responses were most pronounced beneath Betula papyrifera and Populus tremuloides, which have grown more rapidly than Acer saccharum. Although elevated CO₂ has increased plant productivity, the resulting increase in plant litter production has yet to overcome the influence of the pre‐existing pool of soil organic matter on soil microbial activity and rates of N cycling. Ozone reduces plant litter inputs and also appears to affect the composition of plant litter in a way that reduces microbial biomass and activity.     

Carbon and nitrogen turnover in adjacent grassland and cropland ecosystems

The effects of cultivation and soil texture on net and gross N mineralization, CO₂ evolution and C and N turnover were investigated using paired grassland and cropped sites on soils of three textures. Gross N mineralization and immobilization were measured using¹⁵N-isotope dilution. Grassland soils had high CO₂ evolution and gross N mineralization rates, and low net N mineralization rates. Cropland soils had low CO₂ evolution rates but had high net and gross N mineralization rates. Grassland soils thus had high immobilization rates and cropland soils had low immobilization rates. Cultivation increased N turnover but reduced C turnover. The data suggest that the microflora in grassland soils are N limited, while those of cropland soils are limited by C availability. Increasing clay content reduced N turnover. C turnover was less clearly related to texture. Differences in the immobilization potential of substrates help explain why agricultural soils have higher N losses than do grassland soils.     

Net mineralization of N at deeper soil depths as a potential mechanism for sustained forest production under elevated [CO2]

Elevated atmospheric carbon dioxide concentrations [CO₂] is projected to increase forest production, which could increase ecosystem carbon (C) storage. This study contributes to our broad goal of understanding the causes and consequences of increased fine‐root production and mortality under elevated [CO₂] by examining potential gross nitrogen (N) cycling rates throughout the soil profile. Our study was conducted in a CO₂‐enriched sweetgum (Liquidambar styraciflua L.) plantation in Oak Ridge, TN, USA. We used ¹⁵N isotope pool dilution methodology to measure potential gross N cycling rates in laboratory incubations of soil from four depth increments to 60 cm. Our objectives were twofold: (1) to determine whether N is available for root acquisition in deeper soil and (2) to determine whether elevated [CO₂], which has increased inputs of labile C resulting from greater fine‐root mortality at depth, has altered N cycling rates. Although gross N fluxes declined with soil depth, we found that N is potentially available for roots to access, especially below 15 cm depth where rates of microbial consumption of mineral N were reduced relative to production. Overall, up to 60% of potential gross N mineralization and 100% of potential net N mineralization occurred below 15 cm depth at this site. This finding was supported by in situ measurements from ion‐exchange resins, where total inorganic N availability at 55 cm depth was equal to or greater than N availability at 15 cm depth. While it is likely that trees grown under elevated [CO₂] are accessing a larger pool of inorganic N by mining deeper soil, we found no effect of elevated [CO₂] on potential gross or net N cycling rates. Thus, increased root exploration of the soil volume under elevated [CO₂] may be more important than changes in potential gross N cycling rates in sustaining forest responses to rising atmospheric CO₂.     

Combined effects of atmospheric CO<Subscript>2</Subscript> and N availability on the belowground carbon and nitrogen dynamics of aspen mesocosms

It is uncertain whether elevated atmospheric CO₂ will increase C storage in terrestrial ecosystems without concomitant increases in plant access to N. Elevated CO₂ may alter microbial activities that regulate soil N availability by changing the amount or composition of organic substrates produced by roots. Our objective was to determine the potential for elevated CO₂ to change N availability in an experimental plant-soil system by affecting the acquisition of root-derived C by soil microbes. We grew Populus tremuloides (trembling aspen) cuttings for 2 years under two levels of atmospheric CO₂ (36.7 and 71.5 Pa) and at two levels of soil N (210 and 970 μg N g^–1). Ambient and twice-ambient CO₂ concentrations were applied using open-top chambers, and soil N availability was manipulated by mixing soils differing in organic N content. From June to October of the second growing season, we measured midday rates of soil respiration. In August, we pulse-labeled plants with ¹⁴CO₂ and measured soil ¹⁴CO₂ respiration and the ¹⁴C contents of plants, soils, and microorganisms after a 6-day chase period. In conjunction with the August radio-labeling and again in October, we used ¹⁵N pool dilution techniques to measure in situ rates of gross N mineralization, N immobilization by microbes, and plant N uptake. At both levels of soil N availability, elevated CO₂ significantly increased whole-plant and root biomass, and marginally increased whole-plant N capital. Significant increases in soil respiration were closely linked to increases in root biomass under elevated CO₂. CO₂ enrichment had no significant effect on the allometric distribution of biomass or ¹⁴C among plant components, total ¹⁴C allocation belowground, or cumulative (6-day) ¹⁴CO₂ soil respiration. Elevated CO₂ significantly increased microbial ¹⁴C contents, indicating greater availability of microbial substrates derived from roots. The near doubling of microbial ¹⁴C contents at elevated CO₂ was a relatively small quantitative change in the belowground C cycle of our experimental system, but represents an ecologically significant effect on the dynamics of microbial growth. Rates of plant N uptake during both 6-day periods in August and October were significantly greater at elevated CO₂, and were closely related to fine-root biomass. Gross N mineralization was not affected by elevated CO₂. Despite significantly greater rates of N immobilization under elevated CO₂, standing pools of microbial N were not affected by elevated CO₂, suggesting that N was cycling through microbes more rapidly. Our results contained elements of both positive and negative feedback hypotheses, and may be most relevant to young, aggrading ecosystems, where soil resources are not yet fully exploited by plant roots. If the turnover of microbial N increases, higher rates of N immobilization may not decrease N availability to plants under elevated CO₂. Received: 12 February 1999 / Accepted: 2 March 2000     

Influence of the phenolic compound bearing species Ledum palustre on soil N cycling in a boreal hardwood forest

The effects of the understory shrub Ledum palustre on soil N cycling were studied in a hardwood forest of Interior Alaska. This species releases high concentrations of phenolic compounds from green leaves and decomposing litter by rainfall. Organic and mineral soils sampled underneath L. palustre and at nearby non-Ledum sites were amended with L. palustre litter leachates and incubated at controlled conditions. We aimed to know (i) whether L. palustre presence and litter leachate addition changed net N cycling rates in organic and mineral soils, and (ii) what N cycling processes, including gross N mineralization, N immobilization and gross N nitrification, were affected in association with L. palustre. Our results indicate that N transformation rates in the surface organic horizon were not affected by L. palustre presence or leachate addition. However, mineral soils underneath L. palustre as well as soils amended with leachates had significantly higher C/N ratios and microbial respiration rates, and lower net N mineralization and N-to-C mineralization compared to no Ledum and no leachates soils. No nitrification was detected. Plant presence and leachate addition also tended to increase both gross N mineralization and immobilization. These results suggest that soluble C compounds present in L. palustre increased N immobilization in mineral soils when soil biota used them as a C source. Increases in gross N mineralization may have been caused by an enhanced microbial biomass due to C addition. Since both plant presence and leachate addition decreased soil C/N ratio and had similar effects on N transformation rates, our results suggest that litter leachates could be partially responsible for plant presence effects. The lower N availability under L. palustre canopy could exert negative interactions on the establishment and growth of other plant species.     

Leaf litter production and decomposition in a poplar short-rotation coppice exposed to free air CO2 enrichment (POPFACE)

The capacity of forest ecosystems to sequester C in the soil relies on the net balance between litter production above, as well as, below ground, and decomposition processes. Nitrogen mineralization and its availability for plant growth and microbial activity often control the speed of both processes. Litter production, decomposition and N mineralization are strongly interdependent. Thus, their responses to global environmental changes (i.e. elevated CO₂, climate, N deposition, etc.) cannot be fully understood if they are studied in isolation. In the present experiment, we investigated litter fall, litter decomposition and N dynamics in decomposing litter of three Populus spp., in the second and third growing season of a short rotation coppice under FACE. Elevated CO₂ did not affect annual litter production but slightly retarded litter fall in the third growing season. In all species, elevated CO₂ lowered N concentration, resulting in a reduction of N input to the soil via litter fall, but did not affect lignin concentrations. Litter decomposition was studied in bags incubated in situ both in control and FACE plots. Litter lost between 15% and 18% of the original mass during the eight months of field incubation. On average, litter produced under elevated CO₂ attained higher residual mass than control litter. On the other end, when litter was incubated in FACE plots it exhibited higher decay rates. These responses were strongly species‐specific. All litter increased their N content during decomposition, indicating immobilization of N from external sources. Independent of the initial quality, litter incubated on FACE soils immobilized less N, possibly as a result of lower N availability in the soil. Indeed, our results refer to a short‐term decomposition experiment. However, according to a longer‐term model extrapolation of our results, we anticipate that in Mediterranean climate, under elevated atmospheric CO₂, soil organic C pool of forest ecosystems may initially display faster turnover, but soil N availability will eventually limit the process.     

Correlation of foliage and litter chemistry of sugar maple, Acer saccharum, as affected by elevated CO2 and varying N availability, and effects on decomposition

Rising atmospheric carbon dioxide has the potential to alter leaf litter chemistry, potentially affecting decomposition and rates of carbon and nitrogen cycling in forest ecosystems. This study was conducted to determine whether growth under elevated atmospheric CO₂ altered the quality and microbial decomposition of leaf litter of a widely distributed northern hardwood species at sites of low and high soil nitrogen availability. In addition, we assessed whether the carbon–nutrient balance (CNB) and growth differentiation balance (GDB) hypotheses could be extended to predict changes in litter quality in response to resource availability. Sugar maple (Acer saccharum) was grown in the field in open‐top chambers at 36 and 55 Pa partial pressure CO₂, and initial soil mineralization rates of 45 and 348 μg N g^?1 d^?1. Naturally senesced leaf litter was assessed for chemical composition and incubated in the laboratory for 111 d. Microbial respiration and the production of dissolved organic carbon (DOC) were quantified as estimates of decomposition. Elevated CO₂ and low soil nitrogen resulted in higher litter concentrations of nonstructural carbohydrates and condensed tannins, higher C/N ratios and lower N concentrations. Soil N availability appears to have had a greater effect on litter quality than did atmospheric CO₂, although the treatments were additive, with highest concentrations of nonstructural carbohydrates and condensed tannins occurring under elevated CO₂–low soil N. Rates of microbial respiration and the production of DOC were insensitive to differences in litter quality. In general, concentrations of litter constituents, except for starch, were highly correlated to those in live foliage, and the CNB/GDB hypotheses proved useful in predicting changes in litter quality. We conclude the chemical composition of sugar maple litter will change in the future in response to rising atmospheric CO₂, and that soil N availability will exert a major control. It appears that microbial metabolism will not be directly affected by changes in litter quality, although conclusions regarding decomposition as a whole must consider the entire soil food web.     

Elevated CO2 and nutrient addition after soil N cycling and N trace gas fluxes with early season wet-up in a California annual grassland

We examined the effects of growth carbon dioxide (CO₂)concentration and soil nutrient availability on nitrogen (N)transformations and N trace gas fluxes in California grasslandmicrocosms during early-season wet-up, a time when rates of Ntransformation and N trace gas flux are high. After plant senescenceand summer drought, we simulated the first fall rains and examined Ncycling. Growth at elevated CO₂ increased root productionand root carbon:nitrogen ratio. Under nutrient enrichment, elevatedCO₂ increased microbial N immobilization during wet-up,leading to a 43% reduction in gross nitrification anda 55% reduction in NO emission from soil. ElevatedCO₂ increased microbial N immobilization at ambientnutrients, but did not alter nitrification or NO emission. ElevatedCO₂ did not alter soil emission of N₂O ateither nutrient level. Addition of NPK fertilizer (1:1:1) stimulatedN mineralization and nitrification, leading to increased N₂Oand NO emission from soil. The results of our study support a mechanisticmodel in which elevated CO₂ alters soil N cycling and NOemission: increased root production and increased C:N ratio in elevatedCO₂ stimulate N immobilization, thereby decreasingnitrification and associated NO emission when nutrients are abundant.This model is consistent with our basic understanding of how C availabilityinfluences soil N cycling and thus may apply to many terrestrial ecosystems.     

Independent, Interactive, and Species-Specific Responses of Leaf Litter Decomposition to Elevated CO2 and O3 in a Northern Hardwood Forest

The future capacity of forest ecosystems to sequester atmospheric carbon is likely to be influenced by CO₂-mediated shifts in nutrient cycling through changes in litter chemistry, and by interactions with pollutants like O₃. We evaluated the independent and interactive effects of elevated CO₂ (560 μl l⁻¹) and O₃ (55 nl l l⁻¹) on leaf litter decomposition in trembling aspen (Populus tremuloides) and paper birch (Betula papyrifera) at the Aspen free air CO₂ enrichment (FACE) site (Wisconsin, USA). Fumigation treatments consisted of replicated ambient, +CO₂, +O₃, and +CO₂ + O₃ FACE rings. We followed mass loss and litter chemistry over 23 months, using reciprocally transplanted litterbags to separate substrate quality from environment effects. Aspen decayed more slowly than birch across all treatment conditions, and changes in decomposition dynamics of both species were driven by shifts in substrate quality rather than by fumigation environment. Aspen litter produced under elevated CO₂ decayed more slowly than litter produced under ambient CO₂, and this effect was exacerbated by elevated O₃. Similarly, birch litter produced under elevated CO₂ also decayed more slowly than litter produced under ambient CO₂. In contrast to results for aspen, however, elevated O₃ accelerated birch decay under ambient CO₂, but decelerated decay under enriched CO₂. Changes in decomposition rates (k-values) were due to CO₂- and O₃-mediated shifts in litter quality, particularly levels of carbohydrates, nitrogen, and tannins. These results suggest that in early-successional forests of the future, elevated concentrations of CO₂ will likely reduce leaf litter decomposition, although the magnitude of effect will vary among species and in response to interactions with tropospheric O₃.     

Elevated atmospheric CO2 and feedback between carbon and nitrogen cycles

We tested a conceptual model describing the influence of elevated atmospheric CO₂ on plant production, soil microorganisms, and the cycling of C and N in the plant-soil system. Our model is based on the observation that in nutrient-poor soils, plants (C₃) grown in an elevated CO₂ atmosphere often increase production and allocation to belowground structures. We predicted that greater belowground C inputs at elevated CO₂ should elicit an increase in soil microbial biomass and increased rates of organic matter turnover and nitrogen availability. We measured photosynthesis, biomass production, and C allocation of Populus grandidentata Michx. grown in nutrient-poor soil for one field season at ambient and twice-ambient (i.e., elevated) atmospheric CO₂ concentrations. Plants were grown in a sandy subsurface soil i) at ambient CO₂ with no open top chamber, ii) at ambient CO₂ in an open top chamber, and iii) at twice-ambient CO₂ in an open top chamber. Plants were fertilized with 4.5 g N m⁻² over a 47 d period midway through the growing season. Following 152 d of growth, we quantified microbial biomass and the availabilities of C and N in rhizosphere and bulk soil. We tested for a significant CO₂ effect on plant growth and soil C and N dynamics by comparing the means of the chambered ambient and chambered elevated CO₂ treatments. Rates of photosynthesis in plants grown at elevated CO₂ were significantly greater than those measured under ambient conditions. The number of roots, root length, and root length increment were also substantially greater at elevated CO₂. Total and belowground biomass were significantly greater at elevated CO₂. Under N-limited conditions, plants allocated 50–70% of their biomass to roots. Labile C in the rhizosphere of elevated-grown plants was significantly greater than that measured in the ambient treatments; there were no significant differences between labile C pools in the bulk soil of ambient and elevated-grown plants. Microbial biomass C was significantly greater in the rhizosphere and bulk soil of plants grown at elevated CO₂ compared to that in the ambient treatment. Moreover, a short-term laboratory assay of N mineralization indicated that N availability was significantly greater in the bulk soil of the elevated-grown plants. Our results suggest that elevated atmospheric CO₂ concentrations can have a positive feedback effect on soil C and N dynamics producing greater N availability. Experiments conducted for longer periods of time will be necessary to test the potential for negative feedback due to altered leaf litter chemistry. ei]{gnH}{fnLambers} ei]{gnA C}{fnBorstlap}     

Reduced N cycling in response to elevated CO2, warming,and drought in a Danish heathland: Synthesizing results of the CLIMAITE project after two years of treatments

Field‐scale experiments simulating realistic future climate scenarios are important tools for investigating the effects of current and future climate changes on ecosystem functioning and biogeochemical cycling. We exposed a seminatural Danish heathland ecosystem to elevated atmospheric carbon dioxide (CO₂), warming, and extended summer drought in all combinations. Here, we report on the short‐term responses of the nitrogen (N) cycle after 2 years of treatments. Elevated CO₂ significantly affected aboveground stoichiometry by increasing the carbon to nitrogen (C/N) ratios in the leaves of both co‐dominant species (Calluna vulgaris and Deschampsia flexuosa), as well as the C/N ratios of Calluna flowers and by reducing the N concentration of Deschampsia litter. Belowground, elevated CO₂ had only minor effects, whereas warming increased N turnover, as indicated by increased rates of microbial NH₄⁺ consumption, gross mineralization, potential nitrification, denitrification and N₂O emissions. Drought reduced belowground gross N mineralization and decreased fauna N mass and fauna N mineralization. Leaching was unaffected by treatments but was significantly higher across all treatments in the second year than in the much drier first year indicating that ecosystem N loss is highly sensitive to changes and variability in amount and timing of precipitation. Interactions between treatments were common and although some synergistic effects were observed, antagonism dominated the interactive responses in treatment combinations, i.e. responses were smaller in combinations than in single treatments. Nonetheless, increased C/N ratios of photosynthetic tissue in response to elevated CO₂, as well as drought‐induced decreases in litter N production and fauna N mineralization prevailed in the full treatment combination. Overall, the simulated future climate scenario therefore lead to reduced N turnover, which could act to reduce the potential growth response of plants to elevated atmospheric CO₂ concentration.     

Interactions between plant growth and soil nutrient cycling under elevated CO2: a meta-analysis

free air carbon dioxide enrichment (FACE) and open top chamber (OTC) studies are valuable tools for evaluating the impact of elevated atmospheric CO₂ on nutrient cycling in terrestrial ecosystems. Using meta‐analytic techniques, we summarized the results of 117 studies on plant biomass production, soil organic matter dynamics and biological N₂ fixation in FACE and OTC experiments. The objective of the analysis was to determine whether elevated CO₂ alters nutrient cycling between plants and soil and if so, what the implications are for soil carbon (C) sequestration. Elevated CO₂ stimulated gross N immobilization by 22%, whereas gross and net N mineralization rates remained unaffected. In addition, the soil C : N ratio and microbial N contents increased under elevated CO₂ by 3.8% and 5.8%, respectively. Microbial C contents and soil respiration increased by 7.1% and 17.7%, respectively. Despite the stimulation of microbial activity, soil C input still caused soil C contents to increase by 1.2% yr^?1. Namely, elevated CO₂ stimulated overall above‐ and belowground plant biomass by 21.5% and 28.3%, respectively, thereby outweighing the increase in CO₂ respiration. In addition, when comparing experiments under both low and high N availability, soil C contents (+2.2% yr^?1) and above‐ and belowground plant growth (+20.1% and+33.7%) only increased under elevated CO₂ in experiments receiving the high N treatments. Under low N availability, above‐ and belowground plant growth increased by only 8.8% and 14.6%, and soil C contents did not increase. Nitrogen fixation was stimulated by elevated CO₂ only when additional nutrients were supplied. These results suggest that the main driver of soil C sequestration is soil C input through plant growth, which is strongly controlled by nutrient availability. In unfertilized ecosystems, microbial N immobilization enhances acclimation of plant growth to elevated CO₂ in the long‐term. Therefore, increased soil C input and soil C sequestration under elevated CO₂ can only be sustained in the long‐term when additional nutrients are supplied.     

Due to symbiotic N2 fixation,five years of elevated atmospheric pCO2 had no effect on the N concentration of plant litter in fertile,mixed grassland

Experimental findings indicate that, in terrestrial ecosystems, nitrogen cycling changes under elevated partial pressure of atmospheric CO₂ (pCO₂). It was suggested that the concentration of N in plant litter as well as the amount of litter are responsible for these changes. However, for grassland ecosystems, there have been no relevant data available to support this hypothesis. Data from five years of the Swiss FACE experiment show that, under fertile soil conditions in a binary plant community consisting of Lolium perenne L. and Trifolium repens L., the concentration of litter N does not change under elevated atmospheric pCO₂; this applies to harvest losses, stubble, stolons and roots as the sources of litter. This is in strong contrast to the CO₂ response of L. perenne swards without associated legumes; in this case the above-ground concentration of biomass N decreased substantially. Increased symbiotic N₂ fixation in T. repens nodules and a greater proportion of the N-rich T. repens in the community are regarded as the main mechanisms that buffer the increased C introduction into the ecosystem under elevated atmospheric pCO₂. Our data also suggest that elevated atmospheric pCO₂ results in greater amounts of litter, mainly due to increased root biomass production. This study indicates that, in a fertile grassland ecosystem with legumes, the concentration of N in plant litter is not affected by elevated atmospheric pCO₂ and, thus, cannot explain CO₂-induced changes in the cycling of N. This revised version was published online in June 2006 with corrections to the Cover Date.     

Microbial dynamics and carbon and nitrogen cycling following re-wetting of soils beneath two semi-arid plant species

Sporadic summer rainfall in semi-arid ecosystems can provide enough soil moisture to drastically increase CO₂ efflux and rates of soil N cycling. The magnitudes of C and N pulses are highly variable, however, and the factors regulating these pulses are poorly understood. We examined changes in soil respiration, bacterial, fungal and microfaunal populations, and gross rates of N mineralization, nitrification, and NH₄⁺ and NO₃^– immobilization during the 10 days following wetting of dry soils collected from stands of big sagebrush (Artemisia tridentata) and cheatgrass (Bromus tectorum) in central Utah. Soil CO₂ production increased more than tenfold during the 17 h immediately following wetting. The labile organic C pool released by wetting was almost completely respired within 2–3 days, and was nearly three times as large in sagebrush soil as in cheatgrass. In spite of larger labile C pools beneath sagebrush, microbial and microfaunal populations were nearly equal in the two soils. Bacterial and fungal growth coincided with depletion of labile C, and populations peaked in both soils 2 days after wetting. Protozoan populations, whose biomass was nearly 3,000-fold lower than bacteria and fungi, peaked after 2–4 days. Gross N mineralization and nitrification rates were both faster in cheatgrass soil than in sagebrush, and caused greater nitrate accumulation in cheatgrass soil. Grazing of bacteria and fungi by protozoans and nematodes could explain neither temporal trends in N mineralization rates nor differences between soil types. However, a mass balance model indicated that the initial N pulse was associated with degradation of microbial substrates that were rich in N (C:N <8.3), and that microbes had shifted to substrates with lower N contents (C:N =15–25) by day 7 of the incubation. The model also suggested that the labile organic matter in cheatgrass soil had a lower C:N ratio than in sagebrush, and this promoted faster N cycling rates and greater N availability. This study provides evidence that the high N availability often associated with wetting of cheatgrass soils is a result of cheatgrass supplying substrates to microbes that are of high decomposability and N content.     

Effects of elevated CO2, temperature and N fertilization on nitrogen fluxes in a temperate grassland ecosystem

The soil nitrogen cycle was investigated in a pre‐established Lolium perenne sward on a loamy soil and exposed to ambient and elevated atmospheric CO₂ concentrations (350 and 700 μL L^?1) and, at elevated [CO₂], to a 3 °C temperature increase. At two levels of mineral nitrogen supply, N– (150 kgN ha^?1 y^?1) and N+ (533 kgN ha^?1 y^?1), ¹⁵N‐labelled ammonium nitrate was supplied in split applications over a 2.5‐y period. The recovery of the labelled fertilizer N was measured in the harvests, in the stubble and roots, in the macro‐organic matter fractions above 200 μm in size (MOM) and in the aggregated organic matter below 200 μM (AOM). Elevated [CO₂] reduced the total amount of N harvested in the clipped parts of the sward. The harvested N derived from soil was reduced to a greater extent than that derived from fertilizer. At both N supplies, elevated [CO₂] modified the allocation of the fertilizer N in the sward, in favour of the stubble and roots and significantly increased the recovery of fertilizer N in the soil macro‐organic matter fractions. The increase of fertilizer N immobilization in the MOM was associated with a decline of fertilizer N uptake by the grass sward, which supported the hypothesis of a negative feedback of elevated [CO₂] on the sward N yield and uptake. Similar and even more pronounced effects were observed for the native N mineralized in the soil. At N–, a greater part of the fertilizer N organized in the root phytomass resulted in an underestimation of N immobilized in dead roots and, in turn, an underestimation of N immobilization in the MOM. The 3 °C temperature increase alleviated the [CO₂] effect throughout much of the N cycle, increasing soil N mineralization, N derived from soil in the harvests, and the partitioning of the assimilated fertilizer N to shoots. In conclusion, at ambient temperature, the N cycle was slowed down under elevated [CO₂], which restricted the increase in the aboveground production of the grass sward, and apparently contributed to the sequestration of carbon belowground. In contrast, a temperature increase under elevated [CO₂] stimulated the soil nitrogen cycle, improved the N nutrition of the sward and restricted the magnitude of the soil C sequestration.     

Elevated CO2 increases microbial carbon substrate use and nitrogen cycling in Mojave Desert soils

Identifying soil microbial responses to anthropogenically driven environmental changes is critically important as concerns intensify over the potential degradation of ecosystem function. We assessed the effects of elevated atmospheric CO₂ on microbial carbon (C) and nitrogen (N) cycling in Mojave Desert soils using extracellular enzyme activities (EEAs), community‐level physiological profiles (CLPPs), and gross N transformation rates. Soils were collected from unvegetated interspaces between plants and under the dominant shrub (Larrea tridentata) during the 2004–2005 growing season, an above‐average rainfall year. Because most measured variables responded strongly to soil water availability, all significant effects of soil water content were used as covariates to remove potential confounding effects of water availability on microbial responses to experimental treatment effects of cover type, CO₂, and sampling date. Microbial C and N activities were lower in interspace soils compared with soils under Larrea, and responses to date and CO₂ treatments were cover specific. Over the growing season, EEAs involved in cellulose (cellobiohydrolase) and orthophosphate (alkaline phosphatase) degradation decreased under ambient CO₂, but increased under elevated CO₂. Microbial C use and substrate use diversity in CLPPs decreased over time, and elevated CO₂ positively affected both. Elevated CO₂ also altered microbial C use patterns, suggesting changes in the quantity and/or quality of soil C inputs. In contrast, microbial biomass N was higher in interspace soils than soils under Larrea, and was lower in soils exposed to elevated CO₂. Gross rates of NH₄⁺ transformations increased over the growing season, and late‐season NH₄⁺ fluxes were negatively affected by elevated CO₂. Gross NO₃⁻ fluxes decreased over time, with early season interspace soils positively affected by elevated CO₂. General increases in microbial activities under elevated CO₂ are likely attributable to greater microbial biomass in interspace soils, and to increased microbial turnover rates and/or metabolic levels rather than pool size in soils under Larrea. Because soil water content and plant cover type dominates microbial C and N responses to CO₂, the ability of desert landscapes to mitigate or intensify the impacts of global change will ultimately depend on how changes in precipitation and increasing atmospheric CO₂ shift the spatial distribution of Mojave Desert plant communities.     

Soil–Nitrogen Cycling in a Pine Forest Exposed to 5 Years of Elevated Carbon Dioxide

Empirical and modeling studies have shown that the magnitude and duration of the primary production response to elevated carbon dioxide (CO₂) can be constrained by limiting supplies of soil nitrogen (N). We have studied the response of a southern US pine forest to elevated CO₂ for 5 years (1997–2001). Net primary production has increased significantly under elevated CO₂. We hypothesized that the increase in carbon (C) fluxes to the microbial community under elevated CO₂ would increase the rate of N immobilization over mineralization. We tested this hypothesis by quantifying the pool sizes and fluxes of inorganic and organic N in the forest floor and top 30 cm of mineral soil during the first 5 years of CO₂ fumigation. We observed no statistically significant change in the gross or net rate of inorganic N mineralization and immobilization in any soil horizon under elevated CO₂. Similarly, elevated CO₂ had no statistically significant effect on the concentration or flux of organic N, including amino acids. Microbial biomass N was not significantly different between CO₂ treatments. Thus, we reject our hypothesis that elevated CO₂ increases the rate of N immobilization. The quantity and chemistry of the litter inputs to the forest floor and mineral soil horizons can explain the limited range of microbially mediated soil–N cycling responses observed in this ecosystem. Nevertheless a comparative analysis of ecosystem development at this site and other loblolly pine forests suggests that rapid stand development and C sequestration under elevated CO₂ may be possible only in the early stages of stand development, prior to the onset of acute N limitation.     

Effect of Eucalyptus saligna and Albizia falcataria on soil processes and nitrogen supply in Hawaii

This study investigated the differences between two fast-growing tropical tree species on soil N flux and availability. The work was conducted in the island of Hawaii and included three sites located along the Hamakua coast on the northeastern side of the island. Within each site pure stands of Eucalyptus saligna (Sm.)␣and the N2-fixing Albizia falcataria (L.) Fosberg [=Paraserianthes falcataria (L.) Nielsen] were arranged in four randomized complete blocks. For most of the variables considered in this study, the species effects were usually strong and the site effects were significant in some cases. After 13 years, soils under the Albizia stand contained larger pools of total soil C and N, and larger pools of inorganic N. Soil N availability indexed by ion exchange resin bags revealed a strong pattern of species and site effect on N availability; soils under Albizia showed a 2.6–9 fold increase in N availability (P < 0.01). Potential net rates of N transformation (10- and 30-day aerobic incubations) were more than twice as high for soils under the Albizia than under the Eucalytus stands. Nitrogen mineralization during anaerobic incubations were about 10% greater on Albizia soils. Gross microbial mineralization and immobilization were determined by estimating the gross rates of N transformation by the ¹⁵N-isotope pool dilution techniques. Across species and sites, a strong linear positive relationship was obtained for gross immobilization and gross mineralization indicating faster gross immobilization as gross mineralization increases. Soil microbial biomass on Albizia soils contained larger proportion of it as bacterial biomass, while larger proportion of fungi biomass comprised the microbial biomass under Eucalyptus soils. This study clearly showed that the presence of Albizia increased total N pools and N supply to the ecosystem. The overall effect on soil fertility will need to be characterized by the effect of the N₂-fixer on other nutrients, especially the effect on phosphorus. Received: 28 February 1997 / Accepted: 22 September 1997     

Enhanced litter input rather than changes in litter chemistry drive soil carbon and nitrogen cycles under elevated CO2: a microcosm study

Elevated CO₂ has been shown to stimulate plant productivity and change litter chemistry. These changes in substrate availability may then alter soil microbial processes and possibly lead to feedback effects on N availability. However, the strength of this feedback, and even its direction, remains unknown. Further, uncertainty remains whether sustained increases in net primary productivity will lead to increased long‐term C storage in soil. To examine how changes in litter chemistry and productivity under elevated CO₂ influence microbial activity and soil C formation, we conducted a 230‐day microcosm incubation with five levels of litter addition rate that represented 0, 0.5, 1.0, 1.4 and 1.8 × litterfall rates observed in the field for aspen stand growing under control treatments at the Aspen FACE experiment in Rhinelander, WI, USA. Litter and soil samples were collected from the corresponding field control and elevated CO₂ treatment after trees were exposed to elevated CO₂ (560 ppm) for 7 years. We found that small decreases in litter [N] under elevated CO₂ had minor effects on microbial biomass carbon, microbial biomass nitrogen and dissolved inorganic nitrogen. Increasing litter addition rates resulted in linear increase in total C and new C (C from added litter) that accumulated in whole soil as well as in the high density soil fraction (HDF), despite higher cumulative C loss by respiration. Total N retained in whole soil and in HDF also increased with litter addition rate as did accumulation of new C per unit of accumulated N. Based on our microcosm comparisons and regression models, we expected that enhanced C inputs rather than changes in litter chemistry would be the dominant factor controlling soil C levels and turnover at the current level of litter production rate (230 g C m⁻² yr⁻¹ under ambient CO₂). However, our analysis also suggests that the effects of changes in biochemistry caused by elevated CO₂ could become significant at a higher level of litter production rate, with a trend of decreasing total C in HDF, new C in whole soil, as well as total N in whole soil and HDF.     

Ten years of elevated atmospheric carbon dioxide alters soil nitrogen transformations in a sheep‐grazed pasture

The increasing concentration of atmospheric carbon dioxide (CO₂) is expected to lead to enhanced competition between plants and microorganisms for the available nitrogen (N) in soil. Here, we present novel results from a ¹⁵N tracing study conducted with a sheep‐grazed pasture soil that had been under 10 years of CO₂ enrichment. Our study aimed to investigate changes in process‐specific gross N transformations in a soil previously exposed to an elevated atmospheric CO₂ (eCO₂) concentration and to examine indicators for the occurrence of progressive nitrogen limitation (PNL). Our results show that the mineralization–immobilization turnover (MIT) was enhanced under eCO₂, which was driven by the mineralization of recalcitrant organic N. The retention of N in the grassland was enhanced by increased dissimilatory NO₃^? reduction to NH₄⁺ (DNRA) and decreased NH₄⁺ oxidation. Our results indicate that heterotrophic processes become more important under eCO₂. We conclude that higher MIT of recalcitrant organic N and enhanced N retention are mechanisms that may alleviate PNL in grazed temperate grassland.