Effects of elevated atmospheric CO2 on root decomposition in a scrub oak ecosystem

The effects of elevated atmospheric CO₂ on fine root decomposition over a 828‐day period were investigated using open top chambers with both ambient and elevated (700 ppm) CO₂ treatments in an oak–palmetto scrub ecosystem at Kennedy Space Center, Florida. Carbon dioxide enrichment of the chambers began 15 May 1996. The experiment included roots grown in ambient and elevated carbon dioxide. Vertical litterbags installed in September 1996 in each elevated and ambient chamber incubated from December 1996 to December 1998 showed no significant treatment effect on fine root or rhizome mass loss. Initial fine root percentage mass loss varied from 10.3% to 13.5% after three months; 55.5% to 38.3% of original mass had been lost after 828 days. A period of nitrogen immobilization occurred in both fine roots and rhizomes in the elevated CO₂ incubation, which is a potential mechanism for nitrogen conservation for this system in an elevated CO₂ world .     

Assessment of soil nitrogen and phosphorous availability under elevated CO2 and N-fertilization in a short rotation poplar plantation

Photosynthetic stimulation by elevated [CO₂] is largely regulated by nitrogen and phosphorus availability in the soil. During a 6 year Free Air CO₂ Enrichment (FACE) experiment with poplar trees in two short rotations, inorganic forms of soil nitrogen, extractable phosphorus, microbial and total nitrogen were assessed. Moreover, in situ and potential nitrogen mineralization, as well as enzymatic activities, were determined as measures of nutrient cycling. The aim of this study was to evaluate the effects of elevated [CO₂] and fertilization on: (1) N mineralization and immobilization processes; (2) soil nutrient availability; and (3) soil enzyme activity, as an indication of microbial and plant nutrient acquisition activity. Independent of any treatment, total soil N increased by 23% in the plantation after 6 years due to afforestation. Nitrification was the main process influencing inorganic N availability in soil, while ammonification being null or even negative. Ammonium was mostly affected by microbial immobilization and positively related to total N and microbial biomass N. Elevated [CO₂] negatively influenced nitrification under unfertilised treatment by 44% and consequently nitrate availability by 30% on average. Microbial N immobilization was stimulated by [CO₂] enrichment and probably enhanced the transformation of large amounts of N into organic forms less accessible to plants. The significant enhancement of enzyme activities under elevated [CO₂] reflected an increase in nutrient acquisition activity in the soil, as well as an increase of fungal population. Nitrogen fertilization did not influence N availability and cycling, but acted as a negative feed-back on phosphorus availability under elevated CO₂.     

Does deep soil N availability sustain long‐term ecosystem responses to elevated CO2?

A scrub‐oak woodland has maintained higher aboveground biomass accumulation after 11 years of atmospheric CO₂ enrichment (ambient +350 μmol CO₂ mol^?1), despite the expectation of strong nitrogen (N) limitation at the site. We hypothesized that changes in plant available N and exploitation of deep sources of inorganic N in soils have sustained greater growth at elevated CO₂. We employed a suite of assays performed in the sixth and 11th year of a CO₂ enrichment experiment designed to assess soil N dynamics and N availability in the entire soil profile. In the 11th year, we found no differences in gross N flux, but significantly greater microbial respiration (P≤0.01) at elevated CO₂. Elevated CO₂ lowered extractable inorganic N concentrations (P=0.096) considering the whole soil profile (0–190 cm). Conversely, potential net N mineralization, although not significant in considering the entire profile (P=0.460), tended to be greater at elevated CO₂. Ion‐exchange resins placed in the soil profile for approximately 1 year revealed that potential N availability at the water table was almost 3 × greater than found elsewhere in the profile, and we found direct evidence using a ¹⁵N tracer study that plants took up N from the water table. Increased microbial respiration and shorter mean residence times of inorganic N at shallower depths suggests that enhanced SOM decomposition may promote a sustained supply of inorganic N at elevated CO₂. Deep soil N availability at the water table is considerable, and provides a readily available source of N for plant uptake. Increased plant growth at elevated CO₂ in this ecosystem may be sustained through greater inorganic N supply from shallow soils and N uptake from deep soil.     

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.     

The influence of plants grown under elevated CO2 and N fertilization on soil nitrogen dynamics

We investigated the effects of spring barley growth on nitrogen (N) transformations and rhizosphere microbial processes in a controlled system under elevated carbon dioxide (CO₂) at two levels of N fertilization (applied with ¹⁵N labelling). After 25 d, elevated CO₂ (twice ambient) increased plant growth (dry weight, DW) by 141% at low‐N fertilization and by 60% at high‐N fertilization, but its positive effect on the root‐to‐shoot ratio was only significant at low‐N input. As a result of this plant response, elevated CO₂ caused a greater soil CO₂ efflux, rhizosphere soil DW, and soil microbial biomass under N‐limiting conditions than under high N availability. Elevated CO₂ also caused a significant (P < 0.001) increase in the N recovered by the plant from both the labelled (N_f) and unlabelled (N_s + N_uf) N pools. The dynamics of N in the system as affected by elevated CO₂ were driven principally by mineralization–immobilization turnover, with little loss by denitrification. Under N‐limiting conditions, there is evidence to suggest enhanced nutrient release from soil organic matter (SOM) pools—a process which could be defined as priming. The results of our experiment did not indicate a direct plant‐mediated effect of elevated CO₂ on nitrous oxide (N₂O) fluxes or denitrification activity.     

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].     

Soil Nitrogen Conditions Approach Preinvasion Levels following Restoration of Nitrogen-Fixing Black Locust (Robinia pseudoacacia) Stands in a Pine–Oak Ecosystem

Restoring native plant communities on sites formerly occupied by invasive nitrogen‐fixing species poses unique problems due to elevated soil nitrogen availability. Mitigation practices that reduce available nitrogen may ameliorate this problem. We evaluated the effects of tree removal followed by soil preparation or mulching on native plant growth and soil nitrogen transformations in a pine–oak system formerly occupied by exotic nitrogen‐fixing Black locust (Robinia pseudoacacia) trees. Greenhouse growth experiments with native grasses, Andropogon gerardii and Sorghastrum nutans, showed elevated relative growth rates in soils from Black locust compared with pine–oak stands. Field soil nutrient concentrations and rates of net nitrification and total net N‐mineralization were compared 2 and 4 years since Black locust removal and in control sites. Although soil nitrogen concentrations and total net N‐mineralization rates in the restored sites were reduced to levels that were similar to paired pine–oak stands after only 2 years, net nitrification rates remained 3–34 times higher in the restored sites. Other nutrient ion concentrations (Ca, Mg) and organic matter content were reduced, whereas phosphorus levels remained elevated in restored sites. Thus, 2–4 years following Black locust tree removal and soil horizon mixing achieved through site preparation, the concentrations of many soil nutrients returned to preinvasion levels. However, net nitrification rates remained elevated; cover cropping or carbon addition during restoration of sites invaded by nitrogen fixers could increase nitrogen immobilization and/or reduce nitrate availability, making sites more amenable to native plant establishment.     

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.     

Laboratory incubations reveal potential responses of soil nitrogen cycling to changes in soil C and N availability in Mojave Desert soils exposed to elevated atmospheric CO2

Elevated atmospheric carbon dioxide (CO₂) has the potential to alter soil carbon (C) and nitrogen (N) cycling in arid ecosystems through changes in net primary productivity. However, an associated feedback exists because any sustained increases in plant productivity will depend upon the continued availability of soil N. We took soils from under the canopies of major shrubs, grasses, and plant interspaces in a Mojave Desert ecosystem exposed to elevated atmospheric CO₂ and incubated them in the laboratory with amendments of labile C and N to determine if elevated CO₂ altered the mechanistic controls of soil C and N on microbial N cycling. Net ammonification increased under shrubs exposed to elevated CO₂, while net nitrification decreased. Elevated CO₂ treatments exhibited greater fluxes of N₂O–N under Lycium spp., but not other microsites. The proportion of microbial/extractable organic N increased under shrubs exposed to elevated CO₂. Heterotrophic N₂‐fixation and C mineralization increased with C addition, while denitrification enzyme activity and N₂O–N fluxes increased when C and N were added in combination. Laboratory results demonstrated the potential for elevated CO₂ to affect soil N cycling under shrubs and supports the hypothesis that energy limited microbes may increase net inorganic N cycling rates as the amount of soil‐available C increases under elevated CO₂. The effect of CO₂ enrichment on N‐cycling processes is mediated by its effect on the plants, particularly shrubs. The potential for elevated atmospheric CO₂ to lead to accumulation of NH₄⁺ under shrubs and the subsequent volatilization of NH₃ may result in greater losses of N from this system, leading to changes in the form and amount of plant‐available inorganic N. This introduces the potential for a negative feedback mechanism that could act to constrain the degree to which plants can increase productivity in the face of elevated atmospheric CO₂.     

Combined effects of elevated CO2 and soil drought on carbon and nitrogen allocation of the desert shrub Caragana intermedia

Impacts of either elevated CO₂ or drought stress on plant growth have been studied extensively, but interactive effects of these on plant carbon and nitrogen allocation is inadequately understood yet. In this study the response of the dominant desert shrub, Caragana intermedia Kuanget H.c.Fu, to the interaction of elevated CO₂ (700 ± 20 μmol mol⁻¹) and soil drought were determined in two large environmental growth chambers (18 m²). Elevated CO₂ increased the allocation of biomass and carbon into roots and the ratio of carbon to nitrogen (C:N) as well as the leaf soluble sugar content, but decreased the allocation of biomass and carbon into leaves, leaf nitrogen and leaf soluble protein concentrations. Elevated CO₂ significantly decreased the partitioning of nitrogen into leaves, but increased that into roots, especially under soil drought. Elevated CO₂ significantly decreased the carbon isotope discrimination (Δ) in leaves, but increased them in roots, and the ratio of Δ values between root and leaf, indicating an increased allocation into below-ground parts. It is concluded that stimulation of plant growth by CO₂ enrichment may be negated under soil drought, and under the future environment, elevated CO₂ may partially offset the negative effects of enhanced drought by regulating the partitioning of carbon and nitrogen.     

13C and 15N in microarthropods reveal little response of Douglas-fir ecosystems to climate change

Understanding ecosystem carbon (C) and nitrogen (N) cycling under global change requires experiments maintaining natural interactions among soil structure, soil communities, nutrient availability, and plant growth. In model Douglas-fir ecosystems maintained for five growing seasons, elevated temperature and carbon dioxide (CO₂) increased photosynthesis and increased C storage belowground but not aboveground. We hypothesized that interactions between N cycling and C fluxes through two main groups of microbes, mycorrhizal fungi (symbiotic with plants) and saprotrophic fungi (free-living), mediated ecosystem C storage. To quantify proportions of mycorrhizal and saprotrophic fungi, we measured stable isotopes in fungivorous microarthropods that efficiently censused the fungal community. Fungivorous microarthropods consumed on average 35% mycorrhizal fungi and 65% saprotrophic fungi. Elevated temperature decreased C flux through mycorrhizal fungi by 7%, whereas elevated CO₂ increased it by 4%. The dietary proportion of mycorrhizal fungi correlated across treatments with total plant biomass (n= 4, r²= 0.96, P= 0.021), but not with root biomass. This suggests that belowground allocation increased with increasing plant biomass, but that mycorrhizal fungi were stronger sinks for recent photosynthate than roots. Low N content of needles (0.8–1.1%) and A horizon soil (0.11%) coupled with high C : N ratios of A horizon soil (25–26) and litter (36–48) indicated severe N limitation. Elevated temperature treatments increased the saprotrophic decomposition of litter and lowered litter C : N ratios. Because of low N availability of this litter, its decomposition presumably increased N immobilization belowground, thereby restricting soil N availability for both mycorrhizal fungi and plant growth. Although increased photosynthesis with elevated CO₂ increased allocation of C to ectomycorrhizal fungi, it did not benefit plant N status. Most N for plants and soil storage was derived from litter decomposition. N sequestration by mycorrhizal fungi and limited N release during litter decomposition by saprotrophic fungi restricted N supply to plants, thereby constraining plant growth response to the different treatments.     

Six years of in situ CO2 enrichment evoke changes in soil structure and soil biota of nutrient-poor grassland

Nutrient‐poor grassland on a silty clay loam overlying calcareous debris was exposed to elevated CO₂ for six growing seasons. The CO₂ exchange and productivity were persistently increased throughout the experiment, suggesting increases in soil C inputs. At the same time, elevated CO₂ lead to increased soil moisture due to reduced evapotransporation. Measurements related to soil microflora did not indicate increased soil C fluxes under elevated CO₂. Microbial biomass, soil basal respiration, and the metabolic quotient for CO₂ (qCO₂) were not altered significantly. PLFA analysis indicated no significant shift in the ratio of fungi to bacteria. 0.5 m KCl extractable organic C and N, indicators of changed DOC and DON concentrations, also remained unaltered. Microbial grazer populations (protozoa, bacterivorous and fungivorous nematodes, acari and collembola) and root feeding nematodes were not affected by elevated CO₂. However, total nematode numbers averaged slightly lower under elevated CO₂ (?16%, ns) and nematode mass was significantly reduced (?43%, P = 0.06). This reduction reflected a reduction in large‐diameter nematodes classified as omnivorous and predacious. Elevated CO₂ resulted in a shift towards smaller aggregate sizes at both micro‐ and macro‐aggregate scales; this was caused by higher soil moisture under elevated CO₂. Reduced aggregate sizes result in reduced pore neck diameters. Locomotion of large‐diameter nematodes depends on the presence of large enough pores; the reduction in aggregate sizes under elevated CO₂ may therefore account for the decrease in large nematodes. These animals are relatively high up the soil food web; this decline could therefore trigger top‐down effects on the soil food web. The CO₂ enrichment also affected the nitrogen cycle. The N stocks in living plants and surface litter increased at elevated CO₂, but N in soil organic matter and microbes remained unaltered. Nitrogen mineralization increased markedly, but microbial N did not differ between CO₂ treatments, indicating that net N immobilization rates were unaltered. In summary, this study did not provide evidence that soils and soil microbial communities are affected by increased soil C inputs under elevated CO₂. On the contrary, available data (¹³C tracer data, minirhizotron observations, root ingrowth cores) suggests that soil C inputs did not increase substantially. However, we provide first evidence that elevated CO₂ can reduce soil aggregation at the scale from µ m to mm scale, and that this can affect soil microfaunal populations.     

Effects of elevated atmospheric CO2 concentration on C and N pools and rhizosphere processes in a Florida scrub oak community

The effect of elevated atmospheric CO₂ concentration (C_a) on soil carbon and nitrogen accumulation and soil microbial biomass and activity in a native Florida scrub oak community was studied. The plant community, dominated by Quercus myrtifolia Willd. and Q. geminata Small, was exposed for 2 years to elevated C_a in open‐top chambers. Buried subsoil bags were retrieved after 1 year of exposure to elevated C_a. In addition, soil cores were taken twice from the chambers within two weeks in July 1998 (the first after a long dry spell and the second after 25 mm of rainfall) and divided into rhizosphere and bulk soil. Soil organic matter accumulation (excluding roots) into the buried subsoil bags was lower in elevated than in ambient C_a. Concentrations of soluble carbon and ninhydrin‐reactive nitrogen (N_ninh) in the rhizosphere soil were reduced by elevated C_a for the first sampling date and unaffected for the second sampling date. Microbial activity, measured as fluorescein diacetate (FDA) hydrolysis, decreased in elevated C_a for the first sampling date. Microbial biomass carbon and nitrogen in the bulk soil were unaffected by elevated C_a. There was no effect of elevated C_a on bacterial numbers in the rhizosphere.     

Regenerating temperate forest mesocosms in elevated CO2: belowground growth and nitrogen cycling

The response of temperate forest ecosystems to elevated atmospheric CO₂ concentrations is important because these ecosystems represent a significant component of the global carbon cycle. Two important but not well understood processes which elevated CO₂ may substantially alter in these systems are regeneration and nitrogen cycling. If elevated CO₂ leads to changes in species composition in regenerating forest communities then the structure and function of these ecosystems may be affected. In most temperate forests, nitrogen appears to be a limiting nutrient. If elevated CO₂ leads to reductions in nitrogen cycling through increased sequestration of nitrogen in plant biomass or reductions in mineralization rates, long-term forest productivity may be constrained. To study these processes, we established mesocosms of regenerating forest communities in controlled environments maintained at either ambient (375 ppm) or elevated (700 ppm) CO₂ concentrations. Mesocosms were constructed from intact monoliths of organic forest soil. We maintained these mesocosms for 2 years without any external inputs of nitrogen and allowed the plants naturally present as seeds and rhizomes to regenerate. We used ¹⁵N pool dilution techniques to quantify nitrogen fluxes within the mesocosms at the end of the 2 years. Elevated atmospheric CO₂ concentration significantly affected a number of plant and soil processes in the experimental regenerating forest mesocosms. These changes included increases in total plant biomass production, plant C/N ratios, ectomycorrhizal colonization of tree fine roots, changes in tree fine root architecture, and decreases in plant NH₄ ⁺ uptake rates, gross NH₄ ⁺ mineralization rates, and gross NH₄ ⁺ consumption rates. In addition, there was a shift in the relative biomass contribution of the two dominant regenerating tree species; the proportion of total biomass contributed by white birch (Betula papyrifera) decreased and the proportion of total biomass contributed by yellow birch (B. alleghaniensis) increased. However, elevated CO₂ had no significant effect on the total amount of nitrogen in plant and soil microbial biomass. In this study we observed a suite of effects due to elevated CO₂, some of which could lead to increases in potential long term growth responses to elevated CO₂, other to decreases. The reduced plant NH₄ ⁺ uptake rates we observed are consistent with reduced NH₄ ⁺ availability due to reduced gross mineralization rates. Reduced NH₄ ⁺ mineralization rates are consistent with the increases in C/N ratios we observed for leaf and fine root material. Together, these data suggest the positive increases in plant root architectural parameters and mycorrhizal colonization may not be as important as the potential negative effects of reduced nitrogen availability through decreased decomposition rates in a future atmosphere with elevated CO₂. Received: 10 January 1997 / Accepted: 25 July 1997     

Rapid root closure after fire limits fine root responses to elevated atmospheric CO2 in a scrub oak ecosystem in central Florida, USA

Elevated atmospheric carbon dioxide (CO₂) often stimulates the growth of fine roots, yet there are few reports of responses of intact root systems to long‐term CO₂ exposure. We investigated the effects of elevated CO₂ on fine root growth using open top chambers in a scrub oak ecosystem at Kennedy Space Center, Florida for more than 7 years. CO₂ enrichment began immediately after a controlled burn, which simulated the natural disturbance that occurs in this system every 10–15 years. We hypothesized that (1) root abundance would increase in both treatments as the system recovered from fire; (2) elevated CO₂ would stimulate root growth; and (3) elevated CO₂ would alter root distribution. Minirhizotron tubes were used to measure fine root length density (mm cm^?2) every three months. During the first 2 years after fire recovery, fine root abundance increased in all treatments and elevated CO₂ significantly enhanced root abundance, causing a maximum stimulation of 181% after 20 months. The CO₂ stimulation was initially more pronounced in the top 10 cm and 38–49 cm below the soil surface. However, these responses completely disappeared during the third year of experimental treatment: elevated CO₂ had no effect on root abundance or on the depth distribution of fine roots during years 3–7. The results suggest that, within a few years following fire, fine roots in this scrub oak ecosystem reach closure, defined here as a dynamic equilibrium between production and mortality. These results further suggest that elevated CO₂ hastens root closure but does not affect maximum root abundance. Limitation of fine root growth by belowground resources – particularly nutrients in this nutrient‐poor soil – may explain the transient response to elevated CO₂.     

Disentangling species and functional group richness effects on soil N cycling in a grassland ecosystem

Species richness (SR) and functional group richness (FGR) are often confounded in both observational and experimental field studies of biodiversity and ecosystem function. This precludes discernment of their separate influences on ecosystem processes, including nitrogen (N) cycling, and how those influences might be moderated by global change factors. In a 17‐year field study of grassland species, we used two full factorial experiments to independently vary SR (one or four species, with FGR = 1) and FGR (1–4 groups, with SR = 4) to assess SR and FGR effects on ecosystem N cycling and its response to elevated carbon dioxide (CO₂) and N addition. We hypothesized that increased plant diversity (either SR or FGR) and elevated CO₂ would enhance plant N pools because of greater plant N uptake, but decrease soil N cycling rates because of greater soil carbon inputs and microbial N immobilization. In partial support of these hypotheses, increasing SR or FGR (holding the other constant) enhanced total plant N pools and decreased soil nitrate pools, largely through higher root biomass, and increasing FGR strongly reduced mineralization rates, because of lower root N concentrations. In contrast, increasing SR (holding FGR constant and despite increasing total plant C and N pools) did not alter root N concentrations or net N mineralization rates. Elevated CO₂ had minimal effects on plant and soil N metrics and their responses to plant diversity, whereas enriched N increased plant and soil N pools, but not soil N fluxes. These results show that functional diversity had additional effects on both plant N pools and rates of soil N cycling that were independent of those of species richness.     

Seeing the forest for the trees: long-term exposure to elevated CO2 increases some herbivore densities

The effects of elevated CO₂ on plant growth and insect herbivory have been frequently investigated over the past 20 years. Most studies have shown an increase in plant growth, a decrease in plant nitrogen concentration, an increase in plant secondary metabolites and a decrease in herbivory. However, such studies have generally overlooked the fact that increases in plant production could cause increases of herbivores per unit area of habitat. Our study investigated leaf production, herbivory levels and herbivore abundance per unit area of leaf litter in a scrub‐oak system at Kennedy Space Center, Florida, under conditions of ambient and elevated CO₂, over an 11‐year period, from 1996 to 2007. In every year, herbivory, that is leafminer and leaftier abundance per 200 leaves, was lower under elevated CO₂ than ambient CO₂ for each of three species of oaks, Quercus myrtifolia, Quercus chapmanii and Quercus geminata. However, leaf litter production per 0.1143 m² was greater under elevated CO₂ than ambient CO₂ for Q. myrtifolia and Q. chapmanii, and this difference increased over the 11 years of the study. Leaf production of Q. geminata under elevated CO₂ did not increase. Leafminer densities per 0.1143 m² of litterfall for Q. myrtifolia and Q. chapmanii were initially lower under elevated CO₂. However, shortly after canopy closure in 2001, leafminer densities per 0.1143 m² of litter fall became higher under elevated CO₂ and remained higher for the remainder of the experiment. Leaftier densities per 0.1143 m² were also higher under elevated CO₂ for Q. myrtifolia and Q. chapmanii over the last 6 years of the experiment. There were no differences in leafminer or leaftier densities per 0.1143 m² of litter for Q. geminata. These results show three phenomena. First, they show that elevated CO₂ decreases herbivory on all oak species in the Florida scrub‐oak system. Second, despite lower numbers of herbivores per 200 leaves in elevated CO₂, increased leaf production resulted in higher herbivore densities per unit area of leaf litter for two oak species. Third, they corroborate other studies which suggest that the effects of elevated CO₂ on herbivores are species specific, meaning they depend on the particular plant species involved. Two oak species showed increases in leaf production and herbivore densities per 0.1143 m² in elevated CO₂ over time while another oak species did not. Our results point to a future world of elevated CO₂ where, despite lower plant herbivory, some insect herbivores may become more common.     

Ectomycorrhizal Colonization, Biomass, and Production in a Regenerating Scrub Oak Forest in Response to Elevated CO2

The effects of CO₂ elevation on the dynamics of fine root (FR) mass and ectomycorrhizal (EM) mass and colonization were studied in situ in a Florida scrub oak system over four years of postfire regeneration. Soil cores were taken at five dates and sorted to assess the standing crop of ectomycorrhizal and fine roots. We used ingrowth bags to estimate the effects of elevated CO₂ on production of EM roots and fine roots. Elevated CO₂ tended to increase EM colonization frequency but did not affect EM mass nor FR mass in soil cores (standing mass). However, elevated CO₂ strongly increased EM mass and FR mass in ingrowth bags (production), but it did not affect the EM colonization frequency therein. An increase in belowground production with unchanged biomass indicates that elevated CO₂ may stimulate root turnover. The CO₂-stimulated increase of belowground production was initially larger than that of aboveground production. The oaks may allocate a larger portion of resources to root/mycorrhizal production in this system in elevated rather than ambient CO₂.     

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.     

Stimulation of grassland nitrogen cycling under carbon dioxide enrichment

 Nitrogen (N) limits plant growth in many terrestrial ecosystems, potentially constraining terrestrial ecosystem response to elevated CO₂. In this study, elevated CO₂ stimulated gross N mineralization and plant N uptake in two annual grasslands. In contrast to other studies that have invoked increased C input to soil as the mechanism altering soil N cycling in response to elevated CO₂, increased soil moisture, due to decreased plant transpiration in elevated CO₂, best explains the changes we observed. This study suggests that atmospheric CO₂ concentration may influence ecosystem biogeochemistry through plant control of soil moisture. Received: 18 December 1995 / Accepted: 19 June 1996