Species control variation in litter decomposition in a pine forest exposed to elevated CO2

Net primary production and the flux of dry matter and nutrients from vegetation to soils has increased following four years of exposure to elevated CO₂ in a southern pine forest in NC, USA. This has increased the demand for nutrients to support enhanced rates of NPP and altered the conditions for litter decomposition on the forest floor. We quantified the chemistry and decomposition dynamics of leaf litter produced by five of the most abundant tree species in this ecosystem during the third and fourth growing seasons under elevated CO₂. The objectives of this study were to determine (i) if there were systemic or species‐specific changes in leaf litter chemistry associated with a sustained enhancement of plant growth under elevated CO₂; and (ii) whether the process of litter decomposition was altered by increased inputs of energy and nutrients to the forest floor in the plots under elevated CO₂. Leaf litter chemistry, including various C fractions and N concentration, was virtually unchanged by elevated CO₂. With few exceptions, plant litter produced under elevated CO₂ lost mass or N at the same relative rate as that produced under ambient CO₂. The relationship between initial litter chemistry and decomposition was not altered by elevated CO₂. The greater forest floor mass and nutrient content in the plots under elevated CO₂ had no consistent or long‐term effect on litter decomposition. Thus, we found no evidence that plant and microbial processes under elevated CO₂ resulted in systemic changes in mass loss or N dynamics during decomposition. In contrast to the limited effects of elevated CO₂ on litter chemistry and decomposition, there were large differences among species in initial litter chemistry, mass loss and N dynamics during decomposition. If the species composition of this forest community is altered by elevated CO₂, the indirect effect of a change in species composition will exert greater control over the long‐term rate of nutrient cycling than the direct effect of elevated CO₂ on litter chemistry and decomposition dynamics alone.     

Limitations to CO2-induced growth enhancement in pot studies

Recently, it has been suggested that small pots may reduce or eliminate plant responses to enriched CO₂ atmospheres due to root restriction. While smaller pot volumes provide less physical space available for root growth, they also provide less nutrients. Reduced nutrient availability alone may reduce growth enhancement under elevated CO₂. To investigate the relative importance of limited physical rooting space separate from and in conjunction with soil nutrients, we grew plants at ambient and double-ambient CO₂ levels in growth containers of varied volume, shape, nutrient concentration, and total nutrient content. Two species (Abutilon theophrasti, a C₃ dicot with a deep tap root andSetaria faberii, a C₄ monocot with a shallow diffuse root system) were selected for their contrasting physiology and root architecture. Shoot demography was determined weekly and biomass was determined after eight and ten weeks of growth. Increasing total nutrients, either by increasing nutrient concentration or by increasing pot size, increased plant growth. Further, increasing pot size while maintaining equal total nutrients per pot resulted in increased total biomass for both species. CO₂-induced growth and reproductive yield enhancements were greatest in pots with high nutrient concentrations, regardless of total nutrient content or pot size, and were also mediated by the shape of the pot. CO₂-induced growth and reproductive yield enhancements were unaffected by pot size (growth) or were greater in small pots (reproductive yield), regardless of total nutrient content, contrary to predictions based on earlier studies. These results suggest that several aspects of growth conditions within pots may influence the CO₂ responses of plants; pot size, pot shape, the concentration and total amount of nutrient additions to pots may lead to over-or underestimates of the CO₂ responses of real-world plants.     

Tree mineral nutrition is deteriorating in Europe

The response of forest ecosystems to increased atmospheric CO₂ is constrained by nutrient availability. It is thus crucial to account for nutrient limitation when studying the forest response to climate change. The objectives of this study were to describe the nutritional status of the main European tree species, to identify growth‐limiting nutrients and to assess changes in tree nutrition during the past two decades. We analysed the foliar nutrition data collected during 1992–2009 on the intensive forest monitoring plots of the ICP Forests programme. Of the 22 significant temporal trends that were observed in foliar nutrient concentrations, 20 were decreasing and two were increasing. Some of these trends were alarming, among which the foliar P concentration in F. sylvatica, Q. Petraea and P. sylvestris that significantly deteriorated during 1992–2009. In Q. Petraea and P. sylvestris, the decrease in foliar P concentration was more pronounced on plots with low foliar P status, meaning that trees with latent P deficiency could become deficient in the near future. Increased tree productivity, possibly resulting from high N deposition and from the global increase in atmospheric CO₂, has led to higher nutrient demand by trees. As the soil nutrient supply was not always sufficient to meet the demands of faster growing trees, this could partly explain the deterioration of tree mineral nutrition. The results suggest that when evaluating forest carbon storage capacity and when planning to reduce CO₂ emissions by increasing use of wood biomass for bioenergy, it is crucial that nutrient limitations for forest growth are considered.     

System-level adjustments to elevated CO2 in model spruce ecosystems

Atmospheric carbon dioxide enrichment and increasing nitrogen deposition are often predicted to increase forest productivity based on currently available data for isolated forest tree seedlings or their leaves. However, it is highly uncertain whether such seedling responses will scale to the stand level. Therefore, we studied the effects of increasing CO₂ (280, 420 and 560 μL L^-1) and increasing rates of wet N deposition (0, 30 and 90 kg ha^-1 y^-1) on whole stands of 4-year-old spruce trees (Picea abies). One tree from each of six clones, together with two herbaceous understory species, were established in each of nine 0.7 m² model ecosystems in nutrient poor forest soil and grown in a simulated montane climate for two years. Shoot level light-saturated net photosynthesis measured at growth CO₂ concentrations increased with increasing CO₂, as well as with increasing N deposition. However, predawn shoot respiration was unaffected by treatments. When measured at a common CO₂ concentration of 420 μL L^-1 37% down-regulation of photosynthesis was observed in plants grown at 560 μL CO₂ L^-1. Length growth of shoots and stem diameter were not affected by CO₂ or N deposition. Bud burst was delayed, leaf area index (LAI) was lower, needle litter fall increased and soil CO₂ efflux increased with increasing CO₂. N deposition had no effect on these traits. At the ecosystem level the rate of net CO₂ exchange was not significantly different between CO₂ and N treatments. Most of the responses to CO₂ studied here were nonlinear with the most significant differences between 280 and 420 μL CO₂ L^-1 and relatively small changes between 420 and 560 μL CO₂ L^-1. Our results suggest that the lack of above-ground growth responses to elevated CO₂ is due to the combined effects of physiological down-regulation of photosynthesis at the leaf level, allometric adjustment at the canopy level (reduced LAI), and increasing strength of below-ground carbon sinks. The non-linearity of treatment effects further suggests that major responses of coniferous forests to atmospheric CO₂ enrichment might already be under way and that future responses may be comparatively smaller.     

Effect of nitrogen and phosphorus availability on the growth response of Eucalyptus grandis to high CO2

The response of Eucalyptus grandis seedlings to elevated atmospheric CO₂ concentrations was examined by growing seedlings at either 340 or 660 n mol CO₂ mol^-1 for 6 weeks. Graded increments of phosphorus and nitrogen fertilizers were added to a soil deficient in these nutrients to establish if the growth response to increasing nutrient availability was affected by CO₂ concentration. At 660 μmol CO₂ mol^-1, seedling dry weight was up to five times greater than at 340 μmol CO₂ mol^-1. The absolute response was largest when both nitrogen and phosphorus availability was high but the relative increase in dry weight was greatest at low phosphorus availability. At 340 μmol CO₂ mol^-1 and high nitrogen availability, growth was stimulated by addition of phosphorus up to 76 mg kg 1 soil. Further additions of phosphorus had little effect. However, at 660 μmol CO₂ mol^-1, growth only began to plateau at a phosphorus addition rate of 920mg kg^-1 soil. At 340 μmol CO₂ mol^-1 and high phosphorus availability, increasing nitrogen from 40 to 160mg kg^-1 soil had little effect on plant growth. At high CO₂, growth reached a maximum at between 80 and 160mg nitrogen kg^-1 soil. Total uptake of phosphorus was greater at high CO₂ concentration at all fertilizer addition rates, but nitrogen uptake was either lower or unchanged at high CO₂ concentration except at the highest nitrogen fertilizer rate. The shoot to root ratio was increased by CO₂ enrichment, primarily because the specific leaf weight was greater. The nitrogen and phosphorus concentration in the foliage was lower at elevated CO₂ concentration partly because of the higher specific leaf weight. These results indicate that critical foliar concentrations currently used to define nutritional status and fertilizer management may need to be reassessed as the atmospheric CO₂ concentration rises.     

Tree ring responses to elevated CO2 and increased N deposition in Picea abies

Four- to seven-year-old spruce trees (Picea abies) were exposed to three CO₂ concentrations (280, 420 and 560 cm³ m^?3) and three rates of wet N deposition (0, 30 and 90 kg ha^?1 year^?1) for 3 years in a simulated montane forest climate. Six trees from each of six clones were grown in competition in each of nine 100 × 70 × 36 cm model ecosystems with nutrient-poor natural forest soil. Stem dises were analysed using X-ray densitometry. The radial stem increment was not affected by [CO₂] but increased with increasing rates of N deposition. Wood density was increased by [CO₂], but decreased by N deposition. Wood-starch concentration increased, and wood nitrogen concentration decreased with increasing [CO₂], but neither was affected by N deposition. The lignin concentration in wood was affected by neither [CO₂] nor N deposition. Our results suggest that, under natural growth conditions, rising atmospheric [CO₂] will not lead to enhanced radial stem growth of spruce, but atmospheric N deposition will, and in some regions is probably already doing so. Elevated [CO₂], however, will lead to denser wood unless this effect is compensated by massive atmospheric N deposition. If can be speculated that greater wood density under elevated [CO₂] may alter the mechanical properties of wood, and higher ratios of C/N and lignin/N in wood grown at elevated [CO₂] may affect nutrient cycles of forest ecosystems.     

Elevated CO2 enhances resprouting of a tropical savanna tree

The savannas (cerrado) of south-central Brazil are currently subjected to frequent anthropogenic burning, causing widespread reduction in tree density. Increasing concentrations of atmospheric CO₂ could reduce the impact of such frequent burning by increasing the availability of nonstructural carbohydrate, which is necessary for resprouting. We tested the hypotheses that elevated CO₂ stimulates resprouting and accelerates replenishment of carbohydrate reserves. Using a factorial experiment, seedlings of a common Brazilian savanna tree, Keilmeyera coriacea, were grown at 350 ppm and 700 ppm CO₂ and at two nutrient levels. To simulate burning, the plants were either clipped at 15 weeks or were left unclipped. Among unclipped plants, CO₂ and nutrients both stimulated growth, with no significant interaction between nutrient and CO₂ effects. Among clipped plants, both CO₂ and nutrients stimulated resprouting. However, there was a strong interaction between CO₂ and nutrient effects, with CO₂ having a significant effect only in the presence of high nutrient availability. Under elevated CO₂, carbohydrate reserves remained at higher levels following clipping. Root total nonstructural carbohydrate remained above 36% in all treatments, so carbohydrate reserves did not limit regrowth. These results indicate that under elevated CO₂ this species may be better able to endure the high frequency of anthropogenic burning in the Brazilian savannas. Received: 19 May 1999 / Accepted: 8 November 1999     

Mineralization of carbon and nitrogen in acid forest soil treated with fast and slow-release nutrients

The effects of slow (apatite, biotite) and fast-release nutrients (P, K, Mg) on C and N mineralization in acid forest soil were studied. These nutrients were applied alone or together with urea or urea and limestone. The production of CO₂ in the soil samples taken one and three growing seasons after the application was lower in the soils treated with the fast-release nutrients than in the untreated soils. Similar reduction of microbial activity was not seen after the apatite and apatite+biotite treatments. In the first growing season, urea and urea+limestone enhanced CO₂ production, but after three growing seasons, the opposite was true. Apatite and apatite+biotite added together with urea did not compensate for the decreasing effect of urea on the CO₂ production. The addition of fast-release salts increased somewhat the concentration of NH _inf4 ^sup+ in the soil and more NH₄ ⁺ accumulated during laboratory incubation in the soil samples taken one growing season after the application. The urea addition immediately increased the concentrations of NH₄ ⁺ and of NO₃ ⁻ in the soil, but, three growing seasons after application, urea had only a slight increasing effect on mineral N content of the soil. Slow-release nutrients seem to have a more favourable effect than fast-release salts on nutrient turnover in acid forest soil.     

Simulated mechanisms of soil N feedback on the forest CO2 response

An improved understanding of the response of forest ecosystems to elevated levels of CO₂ in the atmosphere is crucial because atmospheric CO₂ concentration continues to increase at an accelerating rate and forests are an important sink in the global carbon cycle. Several CO₂‐enrichment experiments have now been running for more than 10 years, with highly variable short‐term results after the first decade. Responses to rising [CO₂] over the next few decades will depend on several plant and ecosystem feedbacks that are inadequately understood. In this study, we conduct a sensitivity analysis, within the context of the simulated CO₂ response, using a new version of the G'DAY ecosystem model, with an improved decomposition submodel, applied to a nitrogen‐limited Norway spruce forest site in the north of Sweden. The new decomposition model incorporates important modifications to soil processes, including some that constitute negative feedbacks on an ecosystem's growth response to elevated [CO₂]. The sensitivity analysis reveals key parameters and processes that are important for the simulated CO₂ response on the short term and others that are more important on the long term. A process that has a strong impact on the short‐term response is a change in decomposer composition, potentially in response to altered litter quality. Parameters that become increasingly important in the long term are carbon allocation to root exudates that are directly or indirectly associated with atmospheric N₂ fixation, and the rate of humification of soil organic matter. We identify factors intrinsic to species and site (microbes and resources) and ecosystem nutrient supply that determine the duration of the enhanced simulated growth response to elevated [CO₂].     

The response of a natural phytoplankton community from the Godavari River Estuary to increasing CO2 concentration during the pre-monsoon period

This paper reports for the first time upon the effects of increasing CO₂ concentrations on a natural phytoplankton assemblage in a tropical estuary (the Godavari River Estuary in India). Two short-term (5-day) bottle experiments were conducted (with and without nutrient addition) during the pre-monsoon season when the partial pressure of CO₂ in the surface water is quite low. The results reveal that the concentrations of total chlorophyll, the phytoplankton growth rate, the concentrations of particulate organic matter, the photosynthetic oxygen evolution rates, and the total bacterial count were higher under elevated CO₂ treatments, as compared to ambient conditions (control). δ¹³C of particulate organic matter (POM) varied inversely with respect to CO₂, indicating a clear signature of higher CO₂ influx under the elevated CO₂ levels. Whereas, δ¹³C_POM in the controls indicated the existence of an active bicarbonate transport system under limited CO₂ supply. A considerable change in phytoplankton community structure was noticed, with marker pigment analysis by HPLC revealing that cyanobacteria were dominant over diatoms as CO₂ concentrations increased. A mass balance calculation indicated that insufficient nutrients (N, P and Si) might have inhibited diatom growth compared to cyanobacteria, regardless of increased CO₂ supply. The present study suggests that CO₂ concentration and nutrient supply could have significant effects on phytoplankton physiology and community composition for natural phytoplankton communities in this region. However, this work was conducted during a non-discharge period (nutrient-limited conditions) and the responses of phytoplankton to increasing CO₂ might not necessarily be the same during other seasons with high physicochemical variability. Further investigation is therefore needed.     

Enhanced CO2 uptake is marginally offset by altered fluxes of non-CO2 greenhouse gases in global forests and grasslands under N deposition

Despite the increasing impact of atmospheric nitrogen (N) deposition on terrestrial greenhouse gas (GHG) budget, through driving both the net atmospheric CO₂ exchange and the emission or uptake of non-CO₂ GHGs (CH₄ and N₂O), few studies have assessed the climatic impact of forests and grasslands under N deposition globally based on different bottom-up approaches. Here, we quantify the effects of N deposition on biomass C increment, soil organic C (SOC), CH₄ and N₂O fluxes and, ultimately, the net ecosystem GHG balance of forests and grasslands using a global comprehensive dataset. We showed that N addition significantly increased plant C uptake (net primary production) in forests and grasslands, to a larger extent for the aboveground C (aboveground net primary production), whereas it only caused a small or insignificant enhancement of SOC pool in both upland systems. Nitrogen addition had no significant effect on soil heterotrophic respiration (R_H) in both forests and grasslands, while a significant N-induced increase in soil CO₂ fluxes (R_S, soil respiration) was observed in grasslands. Nitrogen addition significantly stimulated soil N₂O fluxes in forests (76%), to a larger extent in grasslands (87%), but showed a consistent trend to decrease soil uptake of CH₄, suggesting a declined sink capacity of forests and grasslands for atmospheric CH₄ under N enrichment. Overall, the net GHG balance estimated by the net ecosystem production-based method (forest, 1.28 Pg CO₂-eq year⁻¹ vs. grassland, 0.58 Pg CO₂-eq year⁻¹) was greater than those estimated using the SOC-based method (forest, 0.32 Pg CO₂-eq year⁻¹ vs. grassland, 0.18 Pg CO₂-eq year⁻¹) caused by N addition. Our findings revealed that the enhanced soil C sequestration by N addition in global forests and grasslands could be only marginally offset (1.5%–4.8%) by the combined effects of its stimulation of N₂O emissions together with the reduced soil uptake of CH₄.     

Effects of carbon dioxide enrichment on response of mycorrhizal pitch pine (Pinus rigida) to aluminum: growth and mineral nutrition

 Carbon dioxide enrichment may increase the Al tolerance of trees by increasing root growth, root exudation and/or mycorrhizal colonization. The effect of elevated CO₂ on the response of mycorrhizal pitch pine (Pinus rigida Mill.) seedlings to Al was determined in two experiments with different levels of nutrients, 0.1- or 0.2-strength Clark solution. During each experiment, seedlings inoculated with the ectomycorrhizal fungus Pisolithus tinctorius (Pers.) Coker & Couch were grown 13 weeks in sand irrigated with nutrient solution (pH 3.8) containing 0, 6.25, 12.5, or 25 mg/l Al (0, 232, 463, or 927 μM Al) in growth chambers fumigated with 350 (ambient) or 700 (elevated) μl/l CO₂. At ambient CO₂, in the absence of Al, mean total dry weights (DW) of seedlings at the high nutrient level were 164% higher than those at the low level. Total DW at elevated CO₂, in the absence of Al, was significantly greater than that in ambient CO₂ at the low (+34%) and high (+16%) nutrient levels. Root and shoot DW at both nutrient levels decreased with increasing Al concentrations with Al reducing root growth more than shoot growth. Although visible symptoms of Al toxicity in roots and needles were reduced by CO₂ enrichment, there were no significant CO₂ × Al interactions for shoot or root DW. The percentage of seedling roots that became mycorrhizal was negatively related to nutrient level and was greater at elevated than at ambient CO₂ levels. Generally, elevated CO₂ had little effect on concentration of mineral nutrients in roots and needles. Aluminum reduced concentrations of most nutrients by inhibiting uptake. Received: 18 June 1997 / Accepted: 8 December 1997     

Effects of elevated CO2 on leaf gas exchange in beech and oak at two levels of nutrient supply: consequences for sensitivity to drought in beech

Beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.) were grown from seed for two whole seasons at two CO₂ concentrations (ambient and ambient + 250 μmol mol^?1) with two levels of soil nutrient supply. Measurements of net leaf photosynthetic rate (A) and stomatal conductance (g_s) of well-watered plants were taken over both seasons; a drought treatment was applied in the middle of the second growing season to a separate sample of beech drawn from the same population. The net leaf photosynthetic rate of well-watered plants was stimulated in elevated CO₂ by an average of 75% in beech and 33% in oak; the effect continued through both growing seasons at both nutrient levels. There were no interactive effects of CO₂ concentration and nutrient level on A or g_s in beech or oak. Stomatal conductance was reduced in elevated CO₂ by an average of 34% in oak, but in beech there were no significant reductions in g_s except under cloudy conditions (–22% in elevated CO₂). During drought, there was no effect of CO₂ concentration on g_s in beech grown with high nutrients, but for beech grown with low nutrients, g_s was significantly higher in elevated CO₂, causing more rapid soil drying. With high nutrient supply, soil drying was more rapid at elevated CO₂ due to increased leaf area. It appears that beech may substantially increase whole-plant water consumption in elevated CO₂, especially under conditions of high temperature and irradiance when damage due to high evaporative demand is most likely to occur, thereby putting itself at risk during periods of drought.     

Growth,nutrient dynamics,and efficiency responses to carbon dioxide and phosphorus nutrition in soybean

Plant mineral nutrients such as phosphorus may exert major control on crop responses to the rising atmospheric carbon dioxide (CO₂) concentrations. To evaluate the growth, nutrient dynamics, and efficiency responses to CO₂ and phosphorus nutrition, soybean (Glycine max (L.) Merr.) was grown in controlled environment growth chambers with sufficient (0.50 mM) and deficient (0.10 and 0.01 mM) phosphate (Pi) supply under ambient and elevated CO₂ (aCO₂, 400 and eCO₂, 800 µmol mol^?1, respectively). The CO₂ × Pi interaction was detected for leaf area, leaf and stem dry weight, and total plant biomass. The severe decrease in plant biomass in Pi-deficient plants (10–76%) was associated with reduced leaf area and photosynthesis (P_net). The degree of growth stimulation (0–55% total biomass) by eCO₂ was dependent upon the severity of Pi deficiency and was closely associated with the increased phosphorus utilization efficiency. With the exception of leaf and root biomass, Pi deficiency decreased the biomass partitioning to other plant organs with the maximum decrease observed in seed weight (8–42%) across CO₂ levels. The increased tissue nitrogen (N) concentration in Pi-deficient plants was accredited to the lower biomass and increased nutrient uptake due to the larger root to shoot ratio. The tissue P and N concentration tended to be lower at eCO₂ versus aCO₂ and did not appear to be the main cause of the lack of CO₂ response of growth and P_net under severe Pi deficiency. The leaf N/P ratio of >16 was detrimental to soybean growth. The tissue P concentration needed to attain the maximum productivity for biomass and seed yield tended to be higher at eCO₂ versus aCO₂. Therefore, the eCO₂ is likely to increase the leaf critical P concentration for maximum biomass productivity and yield in soybean.     

Temperate forest responses to carbon dioxide, temperature and nitrogen: a model analysis

The ITE Edinburgh Forest Model, which describes diurnal and seasonal changes in the pools and fluxes of C, N and water in a fully coupled forest–soil system, was parametrized to simulate a managed conifer plantation in upland Britain. The model was used to examine (i) the transient effects on forest growth of an IS92a scenario of increasing [CO₂] and temperature over two future rotations, and (ii) the equilibrium (sustainable) effects of all combinations of increases in [CO₂] from 350 to 550 and 750 μmol mol^?1, mean annual temperature from 7.5 to 8.5 and 9.5°C and annual inputs of 20 or 40 kg N ha^?1. Changes in underlying processes represented in the model were then used to explain the responses. Eight conclusions were supported by the model for this forest type and climate.

• 1 Increasing temperatures above 3°C alone may cause forest decline owing to water stress.
• 2 Elevated [CO₂] can protect trees from water stress that they may otherwise suffer in response to increased temperature.
• 3 In N-limiting conditions, elevated [CO₂] can increase allocation to roots with little increase in leaf area, whereas in N-rich conditions elevated [CO₂] may not increase allocation to roots and generally increases leaf area.
• 4 Elevated [CO₂] can decrease water use by forests in N-limited conditions and increase water use in N-rich conditions.
• 5 Elevated [CO₂] can increase forest productivity even in N-limiting conditions owing to increased N acquisition and use efficiency.
• 6 Rising temperatures (along with rising [CO₂]) may increase or decrease forest productivity depending on the supply of N and changes in water stress.
• 7 Gaseous losses of N from the soil can increase or decrease in response to elevated [CO₂] and temperature.
• 8 Projected increases in [CO₂] and temperature (IS92a) are likely to increase net ecosystem productivity and hence C sequestration in temperate forests.

     

Species-specific responses of plant communities to altered carbon and nutrient availability

In a field microcosm experiment, species‐specific responses of aboveground biomass of two California annual grassland communities to elevated CO₂ and nutrient availability were investigated. One community grows on shallow, nutrient‐poor serpentine‐derived soil whereas the other occurs on deeper, modestly fertile sandstone/greenstone‐derived substrate. In most species, CO₂ effects did not appear until late in the growing season, probably because the elevated CO₂ increased water‐use‐efficiency easing, the onset of the summer drought. Responses of aboveground biomass to elevated CO₂ differed depending on nutrient availability. Similarly, biomass responses to nutrient treatments differed depending on the CO₂ status. For the majority of the species, production increased most under elevated CO₂ with added nutrients (N,P,K, and micro nutrients). Some species were losers under conditions that increased overall community production, including Bromus hordeaceus in the serpentine community (negative biomass response under elevated CO₂) and Lotus wrangelianus in both communities (negative biomass response with added nitrogen). Treatment and competitive effects on species‐specific biomass varied in both magnitude and direction, especially in the serpentine community, significantly affecting community structure. Individual resource environments are likely to be affected by neighbouring plants, and these competitive interactions complicate predictions of species' responses to elevated CO₂.     

Nutrient Input and Carbon and Microbial Dynamics in an Ombrotrophic Bog

Slow rates of plant production and decomposition in ombrotrophic bogs are believed to be partially the result of low nutrient availability. To test the effect of nutrient availability on decomposition, carbon dioxide (CO₂) flux dynamics, microbial biomass, and nutrients, we added nitrogen (N) with phosphorus (P) and potassium (K), to prevent limitation of the latter 2 nutrients, over 2 growing seasons to plots at Mer Bleue peatland, Ontario, Canada. After the first growing season, increasing N fertilization (with constant P and K) decreased in vitro CO₂ production potential and increased microbial biomass measured with a chloroform fumigation-extraction technique in the upper peat profile, while by the end of the second season, CO₂ production potential was increased in response to N plus PK treatment, presumably due to more easily decomposable newly formed plant material. In situ CO₂ fluxes measured using chamber-techniques over the second year corroborated this presumption, with greater photosynthetic CO₂ uptake and ecosystem respiration (ER) during high N plus PK treatments. The more efficient microbial community, with slower CO₂ production potential and larger biomass, after the first year was characterized by larger fungal biomass measured with signature phospholipid fatty acids. The majority of N was likely quickly sequestered by the vegetation and transferred to dissolved organic forms and microbial biomass in the upper parts of the peat profile, while additional P relative to controls was distributed throughout the profile, implying that the vegetation at the site was N limited. However, in situ CO₂ flux data suggested the possibility of P or NPK limitation. We hypothesize that nutrient deposition may lead to enhanced C uptake by altering the microbial community and decomposition, however this pattern disappears through subsequent changes in the vegetation and production of more readily decomposable plant tissues.     

Effects of Atmospheric CO(2) Enrichment on the Growth and Mineral Nutrition of Quercus alba Seedlings in Nutrient-Poor Soil

One-year-old dormant white oak (Quercus alba L.) seedlings were planted in a nutrient-deficient forest soil and grown for 40 weeks in growth chambers at ambient (362 microliters per liter) or elevated (690 microliters per liter) levels of CO₂. Although all of the seedlings became severely N deficient, CO₂ enrichment enhanced growth by 85%, with the greatest enhancement in root systems. The growth enhancement did not increase the total water use per plant, so water-use efficiency was significantly greater in elevated CO₂. Total uptake of N, S, and B was not affected by CO₂, therefore, tissue concentrations of these nutrients were significantly lower in elevated CO₂. An increase in nutrient-use efficiency with respect to N was apparent in that a greater proportion of the limited N pool in the CO₂-enriched plants was in fine roots and leaves. The uptake of other nutrients increased with CO₂ concentration, and P and K uptake increased in proportion to growth. Increased uptake of P by plants in elevated CO₂ may have been a result of greater proliferation of fine roots and associated mycorrhizae and rhizosphere bacteria stimulating P mineralization. The results demonstrate that a growth response to CO₂ enrichment is possible in nutrient-limited systems, and that the mechanisms of response may include either increased nutrient supply or decreased physiological demand.     

Rising concentrations of atmospheric CO2 have increased growth in natural stands of quaking aspen (Populus tremuloides)

As atmospheric CO₂ levels rise, temperate and boreal forests in the Northern Hemisphere are gaining importance as carbon sinks. Quantification of that role, however, has been difficult due to the confounding effects of climate change. Recent large‐scale experiments with quaking aspen (Populus tremuloides), a dominant species in many northern forest ecosystems, indicate that elevated CO₂ levels can enhance net primary production. Field studies also reveal that droughts contribute to extensive aspen mortality. To complement this work, we analyzed how the growth of wild aspen clones in Wisconsin has responded to historical shifts in CO₂ and climate, accounting for age, genotype (microsatellite heterozygosity), and other factors. Aspen growth has increased an average of 53% over the past five decades, primarily in response to the 19.2% rise in ambient CO₂ levels. CO₂‐induced growth is particularly enhanced during periods of high moisture availability. The analysis accounts for the highly nonlinear changes in growth rate with age, and is unaffected by sex or location sampled. Growth also increases with individual heterozygosity, but this heterozygote advantage has not changed with rising levels of CO₂ or moisture. Thus, increases in future growth predicted from previous large‐scale, common‐garden work are already evident in this abundant and ecologically important tree species. Owing to aspen's role as a foundation species in many North American forest ecosystems, CO₂‐stimulated growth is likely to have repercussions for numerous associated species and ecosystem processes.