Nitrogen cycling in microcosms of yellow birch exposed to elevated CO2: simultaneous positive and negative below-ground feedbacks

This study investigated simultaneous plant and soil feedbacks on growth enhancement with elevated [CO₂] within microcosms of yellow birch (Betula alleghaniensis Britt.) in the second year of growth. Understanding the integrated responses of model ecosystems may provide key insight into the potential net nutrient feedbacks on [CO₂] growth enhancements in temperate forests. We measured the net biomass production, C:N ratios, root architecture, and mycorrhizal responses of yellow birch, in situ rates gross nitrogen mineralization and the partitioning of available NH₄⁺ between yellow birch and soil microbes. Elevated atmospheric [CO₂] resulted in significant alterations in the cycling of N within the microcosms. Plant C/N ratios were significantly increased, gross mineralization and NH₄⁺ consumption rates were decreased, and relative microbial uptake of NH₄⁺ was increased, representing a suite of N cycling negative feedbacks on N availability. However, increased C/N ratios may also be a mechanism which allows plants to maintain higher growth with a constant or reduced N supply. Total plant N content was increased with elevated [CO₂], suggesting that yellow birch had successfully increased their ability to acquire nutrients during the first year of growth. However, plant uptake rates of NH₄⁺ had decreased in the second year. This discrepancy implies that, in this study, nitrogen uptake showed a trend through ontogeny of decreasing enhancement under elevated [CO₂]. The reduced N mineralization and relatively increased N immobilization are a potential feedback which may drive this ontogenetic trend. This study has demonstrated the importance of using an integrated approach to exploring potential nutrient-cycling feedbacks in elevated [CO₂].     

Elevated CO2 increases nitrogen fixation and decreases soil nitrogen mineralization in Florida scrub oak

We report changes in nitrogen cycling in Florida scrub oak in response to elevated atmospheric CO₂ during the first 14 months of experimental treatment. Elevated CO₂ stimulated above-ground growth, nitrogen mass, and root nodule production of the nitrogen-fixing vine, Galactia elliottii Nuttall. During this period, elevated CO₂ reduced rates of gross nitrogen mineralization in soil, and resulted in lower recovery of nitrate on resin lysimeters. Elevated CO₂ did not alter nitrogen in the soil microbial biomass, but increased the specific rate of ammonium immobilization (NH₄⁺ immobilized per unit microbial N) measured over a 24-h period. Increased carbon input to soil through greater root growth combined with a decrease in the quality of that carbon in elevated CO₂ best explains these changes. These results demonstrate that atmospheric CO₂ concentration influences both the internal cycling of nitrogen (mineralization, immobilization, and nitrification) as well as the processes that regulate total ecosystem nitrogen mass (nitrogen fixation and nitrate leaching) in Florida coastal scrub oak. If these changes in nitrogen cycling are sustained, they could cause long-term feedbacks to the growth responses of plants to elevated CO₂. Greater nitrogen fixation and reduced leaching could stimulate nitrogen-limited plant growth by increasing the mass of labile nitrogen in the ecosystem. By contrast, reduced nitrogen mineralization and increased immobilization will restrict the supply rate of plant-available nitrogen, potentially reducing plant growth. Thus, the net feedback to plant growth will depend on the balance of these effects through time.     

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.     

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

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

Influence of different irrigation regimes on crop yield and water use efficiency of olive

Despite increasing interest in the effects of climate change on soil processes, the response of nitrification to elevated CO₂ remains unclear. Responses may depend on soil nitrogen (N) status, and inferences may vary depending on the methodological approach used. We investigated the interactive effects of elevated CO₂ and inorganic N supply on gross nitrification (using ¹⁵N pool dilution) and potential nitrification (using nitrifying enzyme activity assays) in Dactylis glomerata mesocosms. We measured the responses of putative drivers of nitrification (NH ₄ ⁺ production, NH ₄ ⁺ consumption, and soil environmental conditions) and of potential denitrification, a process functionally linked to nitrification. Gross nitrification was insensitive to all treatments, whereas potential nitrification was higher in the high N treatment and was further stimulated by elevated CO₂ in the high N treatment. Gross mineralization and NH ₄ ⁺ consumption rates were also significantly increased in response to elevated CO₂ in the high N treatment, while potential denitrification showed a significant increase in response to N addition. The discrepancy between the responses of gross and potential nitrification to elevated CO₂ and inorganic N supply suggest that these measurements provide different information, and should be used as complementary approaches to understand nitrification response to global change.     

Elevated CO2 and O3 Alter Soil Nitrogen Transformations beneath Trembling Aspen, Paper Birch, and Sugar Maple

Nitrogen cycling in northern temperate forest ecosystems could change under increasing atmospheric CO₂ and tropospheric O₃ as a result of quantitative and qualitative changes in plant litter production. At the Aspen Free Air CO₂–O₃ Enrichment (FACE) experiment, we previously found that greater substrate inputs to soil under elevated CO₂ did not alter gross N transformation rates in the first 3 years of the experiment. We hypothesized that greater litter production under elevated CO₂ would eventually cause greater gross N transformation rates and that CO₂ effects would be nullified by elevated O₃. Following our original study, we continued measurement of gross N transformation rates for an additional four years. From 1999 to 2003, gross N mineralization doubled, N immobilization increased 4-fold, but changes in microbial biomass N and soil total N were not detected. We observed year-to-year variation in N transformation rates, which peaked during a period of foliar insect damage. Elevated CO₂ caused equivalent increases in gross rates of N mineralization (+34%) and NH ₄ ⁺ immobilization (+36%). These results indicate greater rates of N turnover under elevated CO₂, but do not indicate a negative feedback between elevated CO₂ and soil N availability. Elevated O₃ decreased gross N mineralization (−16%) and had no effect on NH ₄ ⁺ immobilization, indicating reduced N availability under elevated O₃. The effects of CO₂ and O₃ on N mineralization rates were mainly related to changes in litter production, whereas effects on N immobilization were likely influenced by changes in litter chemistry and production. Our findings also indicate that concomitant increases in atmospheric CO₂ and O₃ could lead to a negative feedback on N availability.     

Nitrate influx kinetic parameters of five potato cultivars during vegetative growth

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

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.     

Amino acid abundance and proteolytic potential in North American soils

Studies of nitrogen (N) cycling have traditionally focused on N mineralization as the primary process limiting plant assimilation of N. Recent evidence has shown that plants may assimilate amino acids (AAs) directly, circumventing the mineralization pathway. However, the general abundance of soil AAs and their relative importance in plant N uptake remains unclear in most ecosystems. We compared the concentrations and potential production rates of AAs and NH₄ ⁺, as well as the edaphic factors that influence AA dynamics, in 84 soils across the United States. Across all sites, NH₄ ⁺ and AA-N comprised similar proportions of the total bioavailable N pool (~20%), with NO₃ ⁻ being the dominant form of extractable N everywhere but in tundra and boreal forest soils. Potential rates of AA production were at least comparable to those of NH₄ ⁺ production in all ecosystems, particularly in semi-arid grasslands, where AA production rates were six times greater than for NH₄ ⁺ (P < 0.01). Potential rates of proteolytic enzyme activity were greatest in bacteria-dominated soils with low NH₄ ⁺ concentrations, including many grassland soils. Based on research performed under standardized laboratory conditions, our continental-scale analyses suggest that soil AA and NH₄ ⁺ concentrations are similar in most soils and that AAs may contribute to plant and microbial N demand in most ecosystems, particularly in ecosystems with N-poor soils.     

Effect of soil heterogeneity on gross nitrogen mineralization measured by 15N-pool dilution techniques

Pool dilution techniques, where the soil ammonium pool is labeled with ¹⁵NH₄ ⁺, are commonly used to estimate gross N mineralization rates in soil. To estimate the rates unbiased, it is assumed that the ¹⁵NH₄ ⁺ is distributed homogenously in ambient ¹⁴NH₄ ⁺ pool of the soil. However, completely homogeneous distribution of ¹⁵NH₄ ⁺ may commonly not be feasible. The objective of this paper was to examine the importance of the spatial distribution of ¹⁴NH₄ ⁺ and ¹⁵NH₄ ⁺ on the measured gross N mineralization rate. In a 2-day incubation experiment, gross N mineralization rates were measured in soil, where different distributions of ¹⁴NH₄ ⁺ and ¹⁵NH₄ ⁺ were combined. Generally, distribution of ¹⁵NH₄ ⁺ to 50% of the soil volume did not alter the measured gross mineralization rate however more heterogeneous distribution caused the rate to be underestimated. Certain combinations of ¹⁴NH₄ ⁺ and ¹⁵NH₄ ⁺ distributions caused the rate to be overestimated. Based on the experimental results, we developed a 2-dimensional model array of soil compartments, to estimate the gross N mineralization and gross NH₄ ⁺ consumption rates in local microsites in the soil. If one of the nitrogen-isotopes was more abundant in a compartment with high NH₄ ⁺-concentration, and the other nitrogen-isotope was more abundant in a compartment with low NH₄ ⁺-concentration, the nitrogen-isotopes would have different apparent bioavailability, hence the gross N mineralization rate would be erroneously estimated. On the other hand, in soil where all compartments had low NH₄ ⁺-concentrations, heterogeneous distribution of ¹⁴NH₄ ⁺, ¹⁵NH₄ ⁺ and microbial activity did not influence the measured gross N mineralization rate significantly.     

蚂蚁筑巢对西双版纳热带森林土壤有机氮矿化的影响

蚂蚁作为生态系统工程师能够调节土壤微生物及理化环境,进而对热带森林土壤有机氮矿化速率及其时间动态产生显著影响。以西双版纳白背桐热带森林群落为研究对象,采用室内需氧培养法测定土壤有机氮矿化速率,比较蚁巢和非蚁巢土壤有机氮矿化速率的时间动态,揭示蚂蚁筑巢活动引起土壤无机氮库、微生物生物量碳及化学性质改变对有机氮矿化速率时间动态的影响。结果表明：（1）蚂蚁筑巢显著影响土壤有机氮矿化速率（P<0.01）,相较于非蚁巢,蚁巢土壤有机氮矿化速率提高了261%;（2）土壤有机氮矿化速率随月份推移呈明显的单峰型变化趋势,即6月最大（蚁巢1.22 mg kg^-1 d^-1、非蚁巢0.41 mg kg^-1 d^-1）,12月最小（蚁巢0.82 mg kg^-1 d^-1、非蚁巢0.18 mg kg^-1 d^-1）;（3）两因素方差分析表明,不同月份及不同处理对土壤有机氮矿化速率、NH₄-N及NO₃-N产生显著影响（P<0.05）,但对NO₃-N的交互作用不显著;（4）蚂蚁筑巢显著提高了无机氮库（NH₄-N与NO₃-N）、微生物生物量碳、有机质、水解氮、全氮及易氧化有机碳等土壤养分含量,而降低了土壤pH值;（5）回归分析表明,铵态氮和硝态氮对土壤有机氮矿化速率产生显著影响,分别解释87.89%、61.84%的有机氮矿化速率变化;（6）主成份分析表明NH₄-N、微生物生物量碳及有机质是影响有机氮矿化速率时间动态的主要因素,而全氮、NO₃-N、易氧化有机碳、水解氮及pH对土壤有机氮矿化速率的影响次之,且pH与土壤有机氮矿化速率呈显著负相关。总之,蚂蚁筑巢活动主要通过影响土壤NH₄-N、微生物生物量碳及有机质的状况,进而调控西双版纳热带森林土壤有机氮矿化速率的时间动态。研究结果将有助于进一步提高对土壤氮矿化生物调控机制的认识。     

A 3-year exposure to CO2 and O3 induced minor changes in soil N cycling in a meadow ecosystem

We investigated the individual and interactive effects of moderately elevated CO₂ (ambient air + 100 ppm) and/or O₃ (40–50 ppb) on soil N cycling and microbial biomass N in a 3-year open-top chamber experiment conducted in meadow mesocosms. The results show that elevated O₃ decreased the concentrations of mineral N and NH₄⁺-N in the mesocosm soil in the last growing season (2004). Total N, NO₃⁻-N, microbial biomass N, decomposition rate, potential nitrification and denitrification were not affected by elevated O₃ and/or CO₂. It is thus concluded that the proposed future ambient O₃ and CO₂ levels, such as used in this experiment, may not induce major changes in the below-ground N processes in N-poor northern European hay meadow ecosystems.     

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     

Growth and nitrogen uptake in an experimental community of annuals exposed to elevated atmospheric CO2

Rising levels of atmospheric CO₂ may alter patterns of plant biomass production. These changes will be dependent on the ability of plants to acquire sufficient nutrients to maintain enhanced growth. Species-specific differences in responsiveness to CO₂ may lead to changes in plant community composition and biodiversity. Differences in species-level growth responses to CO₂ may be, in a large part, driven by differences in the ability to acquire nutrients. To understand the mechanisms of how elevated CO₂ leads to changes in community-level productivity, we need to study the growth responses and patterns of nutrient acquisition for each of the species that comprise the community. In this paper, we present a study of how elevated CO₂ affects community-level and species-level patterns of nitrogen uptake and biomass production. As an experimental system we use experimental communities of 11 co-occurring annuals common to disturbed seasonal grasslands in south-western U.S.A. We established experimental communities with approximately even numbers of each species in three different atmospheric CO₂ concentrations (375, 550, and 700 ppm). We maintained these communities for 1, 1.5, and 2 months at which times we applied a ¹⁵N tracer (¹⁵NH₄¹⁵NO₃) to quantify the nitrogen uptake and then measured plant biomass, nitrogen content, and nitrogen uptake rates for the entire communities as well as for each species. Overall, community-level responses to elevated CO₂ were consistent with the majority of other studies of individual- and multispecies assemblages, where elevated CO₂ leads to enhanced biomass production early on, but this enhancement declines through time. In contrast, the responses of the individual species within the communities was highly variable, showing the full range of responses from positive to negative. Due to the large variation in size between the different species, community-level responses were generally determined by the responses of only one or a few species. Thus, while several of the smaller species showed trends of increased biomass and nitrogen uptake in elevated CO₂ at the end of the experiment, community-level patterns showed a decrease in these parameters due to the significant reduction in biomass and nitrogen content in the single largest species. The relationship between enhancement of nitrogen uptake and biomass production in elevated CO₂ was highly significant for both 550 ppm and 700 ppm CO₂. This relationship strongly suggests that the ability of plants to increase nitrogen uptake (through changes in physiology, morphology, architecture, or mycorrhizal symbionts) may be an important determinant of which species in a community will be able to respond to increased CO₂ levels with increased biomass production. The fact that the most dominant species within the community showed reduced enhancement and the smaller species showed increased enhancement suggest that through time, elevated CO₂ may lead to significant changes in community composition. At the community level, nitrogen uptake rates relative to plant nitrogen content were invariable between the three different CO₂ levels at each harvest. This was in contrast to significant reductions in total plant nitrogen uptake and nitrogen uptake relative to total plant biomass. These patterns support the hypothesis that plant nitrogen uptake is largely regulated by physiological activity, assuming that physiological activity is controlled by nitrogen content and thus protein and enzyme content.     

Effect of elevated CO2 concentration and nitrate: ammonium ratios on gas exchange and growth of cassava (Manihot esculenta Crantz)

Aims

This study evaluated how different nitrogen forms affect growth and photosynthetic responses of cassava to CO₂ concentration.

Methods

Cassava was grown in 14-L pots in a greenhouse at 390 or 750 ppm of CO₂. Three nitrogen treatments were applied: (a) 12?mM NO₃ ^?, (b) 6?mM NO₃ ^??+?6?mM NH₄ ⁺, and (c) 12?mM NH₄ ⁺.

Results

Thirty-six days after treatments began, plants grown under elevated CO₂ and fertilized only with NO₃ ^? (750_NO₃ ^?) had photosynthetic rates similar to plants grown under 390_NO₃ ^?, indicating significant photosynthetic acclimation to CO₂. In contrast, photosynthetic rates at elevated CO₂ increased as NH₄ ⁺ increased in the nutrient solution, such that photosynthetic acclimation was reduced for plants fertilized with only NH₄ ⁺. However, this positive effect of NH₄ ⁺ on photosynthesis was not observed in more advanced growth stages, and the toxic effects of NH₄ ⁺ severely reduced total dry mass for these plants measured at the end of the experiment.

Conclusions

Our results indicate that cassava will respond with increased biomass accumulation in response to raising atmospheric CO₂ levels, and that N form can have an important impact on the photosynthetic response. However, the positive effect of NH₄ ⁺ fertilization on cassava photosynthetic CO₂ response eventually led to a toxicity problem that reduced biomass production. The challenge is to determine how to manage NH₄ ⁺ fertilization so that the photosynthetic benefit observed in the initial phase may persist throughout the crop cycle.     

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.     

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.     

Effects of elevated CO2 and nitrogen addition on soil organic carbon fractions in a subtropical forest

Aims

The aim of this study was to investigate the effects of elevated CO₂ concentration and nitrogen addition on soil organic carbon fractions in subtropical forests where the ambient N deposition was high.

Methods

Seedlings of typical subtropical forest ecosystems were transplanted in ten open-top chambers and grown under CO₂ and nitrogen treatments. The treatments included: 1) elevated CO₂ (700?μmol?mol^-1); 2) N addition of 100?kg NH₄NO₃ ha^-1?yr^-1; 3) combined elevated CO₂ and N addition; and 4) control. We measured soil total organic carbon (TOC), particulate organic carbon (POC), readily oxidizable organic carbon (ROC), and microbial biomass carbon (MBC).

Results

Results showed that elevated CO₂ alone did not significantly affect soil TOC, POC and ROC after 4?years of treatment, but increased soil MBC and soil respiration compared to the control. N addition alone had no significant effect neither on soil TOC, POC and ROC, but decreased MBC and soil respiration over time. However, the elevated CO₂ and N addition together significantly increased soil POC and ROC, and had no significant effect on soil MBC.

Conclusions

This study indicated that even in N-rich subtropical forest ecosystems, inputs of N are still needed in order to sustain soil C accumulation under elevated CO₂.     

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.     

Responses of communities of tropical tree species to elevated CO2 in a forest clearing

Communities of ten species of tropical forest tree seedlings from three successional classes were grown at ambient and elevated CO₂ in large open-top chambers on the edge of a forest in Panamá. Communities grew from 20?cm to approximately 2?m in height in 6 months. No enhancements in plant biomass accumulation occurred under elevated CO₂ either in the whole communities or in growth of individual species. Reductions in leaf area index under elevated CO₂ were observed, as were decreases in leaf nitrogen concentrations and increases in the C:N ratio of leaf tissue. Species tended to respond individualistically to elevated CO₂, but some generalizations of how successional groupings responded could be made. Early and mid-successional species generally showed greater responses to elevated CO₂ than late-successional species, particularly with respect to increases in photosynthetic rates and leaf starch concentrations, and reductions in leaf area ratio. Late-successional species showed greater increases in C:N ratios in response to elevated CO₂ than did other species. Our results indicate that there may not be an increase in the growth of regenerating tropical forest under elevated CO₂, but that there could be changes in soil nutrient availability because of reductions in leaf tissue quality, particularly in late-successional species.