Soil carbon sequestration in a pine forest after 9 years of atmospheric CO2 enrichment

The impact of anthropogenic CO₂ emissions on climate change may be mitigated in part by C sequestration in terrestrial ecosystems as rising atmospheric CO₂ concentrations stimulate primary productivity and ecosystem C storage. Carbon will be sequestered in forest soils if organic matter inputs to soil profiles increase without a matching increase in decomposition or leaching losses from the soil profile, or if the rate of decomposition decreases because of increased production of resistant humic substances or greater physical protection of organic matter in soil aggregates. To examine the response of a forest ecosystem to elevated atmospheric CO₂ concentrations, the Duke Forest Free‐Air CO₂ Enrichment (FACE) experiment in North Carolina, USA, has maintained atmospheric CO₂ concentrations 200 μL L^?1 above ambient in an aggrading loblolly pine (Pinus taeda) plantation over a 9‐year period (1996–2005). During the first 6 years of the experiment, forest‐floor C and N pools increased linearly under both elevated and ambient CO₂ conditions, with significantly greater accumulations under the elevated CO₂ treatment. Between the sixth and ninth year, forest‐floor organic matter accumulation stabilized and C and N pools appeared to reach their respective steady states. An additional C sink of ～30 g C m^?2 yr^?1 was sequestered in the forest floor of the elevated CO₂ treatment plots relative to the control plots maintained at ambient CO₂ owing to increased litterfall and root turnover during the first 9 years of the study. Because we did not detect any significant elevated CO₂ effects on the rate of decomposition or on the chemical composition of forest‐floor organic matter, this additional C sink was likely related to enhanced litterfall C inputs. We also failed to detect any statistically significant treatment effects on the C and N pools of surface and deep mineral soil horizons. However, a significant widening of the C : N ratio of soil organic matter (SOM) in the upper mineral soil under both elevated and ambient CO₂ suggests that N is being transferred from soil to plants in this aggrading forest. A significant treatment × time interaction indicates that N is being transferred at a higher rate under elevated CO₂ (P=0.037), suggesting that enhanced rates of SOM decomposition are increasing mineralization and uptake to provide the extra N required to support the observed increase in primary productivity under elevated CO₂.     

Annual burning of a tallgrass prairie inhibits C and N cycling in soil,increasing recalcitrant pyrogenic organic matter storage while reducing N availability

Grassland ecosystems store an estimated 30% of the world's total soil C and are frequently disturbed by wildfires or fire management. Aboveground litter decomposition is one of the main processes that form soil organic matter (SOM). However, during a fire biomass is removed or partially combusted and litter inputs to the soil are substituted with inputs of pyrogenic organic matter (py‐OM). Py‐OM accounts for a more recalcitrant plant input to SOM than fresh litter, and the historical frequency of burning may alter C and N retention of both fresh litter and py‐OM inputs to the soil. We compared the fate of these two forms of plant material by incubating ¹³C‐ and ¹⁵N‐labeled Andropogon gerardii litter and py‐OM at both an annually burned and an infrequently burned tallgrass prairie site for 11 months. We traced litter and py‐OM C and N into uncomplexed and organo‐mineral SOM fractions and CO₂ fluxes and determined how fire history affects the fate of these two forms of aboveground biomass. Evidence from CO₂ fluxes and SOM C:N ratios indicates that the litter was microbially transformed during decomposition while, besides an initial labile fraction, py‐OM added to SOM largely untransformed by soil microbes. Additionally, at the N‐limited annually burned site, litter N was tightly conserved. Together, these results demonstrate how, although py‐OM may contribute to C and N sequestration in the soil due to its resistance to microbial degradation, a long history of annual removal of fresh litter and input of py‐OM infers N limitation due to the inhibition of microbial decomposition of aboveground plant inputs to the soil. These results provide new insight into how fire may impact plant inputs to the soil, and the effects of py‐OM on SOM formation and ecosystem C and N cycling.     

Altered microbial community structure and organic matter composition under elevated CO2 and N fertilization in the duke forest

The dynamics and fate of terrestrial organic matter (OM) under elevated atmospheric CO₂ and nitrogen (N) fertilization are important aspects of long‐term carbon sequestration. Despite numerous studies, questions still remain as to whether the chemical composition of OM may alter with these environmental changes. In this study, we employed molecular‐level methods to investigate the composition and degradation of various OM components in the forest floor (O horizon) and mineral soil (0–15 cm) from the Duke forest free air CO₂ enrichment (FACE) experiment. We measured microbial responses to elevated CO₂ and N fertilization in the mineral soil using phospholipid fatty acid (PLFA) profiles. Increased fresh carbon inputs into the forest floor under elevated CO₂ were observed at the molecular‐level by two degradation parameters of plant‐derived steroids and cutin‐derived compounds. The ratios of fungal to bacterial PLFAs and Gram‐negative to Gram‐positive bacterial PLFAs decreased in the mineral soil with N fertilization, indicating an altered soil microbial community composition. Moreover, the acid to aldehyde ratios of lignin‐derived phenols increased with N fertilization, suggesting enhanced lignin degradation in the mineral soil. ¹H nuclear magnetic resonance (NMR) spectra of soil humic substances revealed an enrichment of leaf‐derived alkyl structures with both elevated CO₂ and N fertilization. We suggest that microbial decomposition of SOM constituents such as lignin and hydrolysable lipids was promoted under both elevated CO₂ and N fertilization, which led to the enrichment of plant‐derived recalcitrant structures (such as alkyl carbon) in the soil.     

Distinct responses of soil respiration to experimental litter manipulation in temperate woodland and tropical forest

Global change is affecting primary productivity in forests worldwide, and this, in turn, will alter long‐term carbon (C) sequestration in wooded ecosystems. On one hand, increased primary productivity, for example, in response to elevated atmospheric carbon dioxide (CO₂), can result in greater inputs of organic matter to the soil, which could increase C sequestration belowground. On other hand, many of the interactions between plants and microorganisms that determine soil C dynamics are poorly characterized, and additional inputs of plant material, such as leaf litter, can result in the mineralization of soil organic matter, and the release of soil C as CO₂ during so‐called “priming effects”. Until now, very few studies made direct comparison of changes in soil C dynamics in response to altered plant inputs in different wooded ecosystems. We addressed this with a cross‐continental study with litter removal and addition treatments in a temperate woodland (Wytham Woods) and lowland tropical forest (Gigante forest) to compare the consequences of increased litterfall on soil respiration in two distinct wooded ecosystems. Mean soil respiration was almost twice as high at Gigante (5.0 μmol CO₂ m^?2 s^?1) than at Wytham (2.7 μmol CO₂ m^?2 s^?1) but surprisingly, litter manipulation treatments had a greater and more immediate effect on soil respiration at Wytham. We measured a 30% increase in soil respiration in response to litter addition treatments at Wytham, compared to a 10% increase at Gigante. Importantly, despite higher soil respiration rates at Gigante, priming effects were stronger and more consistent at Wytham. Our results suggest that in situ priming effects in wooded ecosystems track seasonality in litterfall and soil respiration but the amount of soil C released by priming is not proportional to rates of soil respiration. Instead, priming effects may be promoted by larger inputs of organic matter combined with slower turnover rates.     

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

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

Indirect effects of soil moisture reverse soil C sequestration responses of a spring wheat agroecosystem to elevated CO2

Increased plant productivity under elevated atmospheric CO₂ concentrations might increase soil carbon (C) inputs and storage, which would constitute an important negative feedback on the ongoing atmospheric CO₂ rise. However, elevated CO₂ often also leads to increased soil moisture, which could accelerate the decomposition of soil organic matter, thus counteracting the positive effects via C cycling. We investigated soil C sequestration responses to 5 years of elevated CO₂ treatment in a temperate spring wheat agroecosystem. The application of ¹³C‐depleted CO₂ to the elevated CO₂ plots enabled us to partition soil C into recently fixed C (C_new) and pre‐experimental C (C_old) by ¹³C/¹²C mass balance. Gross C inputs to soils associated with C_new accumulation and the decomposition of C_old were then simulated using the Rothamsted C model ‘RothC.’ We also ran simulations with a modified RothC version that was driven directly by measured soil moisture and temperature data instead of the original water balance equation that required potential evaporation and precipitation as input. The model accurately reproduced the measured C_new in bulk soil and microbial biomass C. Assuming equal soil moisture in both ambient and elevated CO₂, simulation results indicated that elevated CO₂ soils accumulated an extra ～40–50 g C m^?2 relative to ambient CO₂ soils over the 5 year treatment period. However, when accounting for the increased soil moisture under elevated CO₂ that we observed, a faster decomposition of C_old resulted; this extra C loss under elevated CO₂ resulted in a negative net effect on total soil C of ～30 g C m^?2 relative to ambient conditions. The present study therefore demonstrates that positive effects of elevated CO₂ on soil C due to extra soil C inputs can be more than compensated by negative effects of elevated CO₂ via the hydrological cycle.     

Nonlinear root-derived carbon sequestration across a gradient of nitrogen and phosphorous deposition in experimental mesocosms

Enhanced sequestration of plant‐carbon (C) inputs to soil may mitigate rising atmospheric carbon dioxide (CO₂) concentrations and related climate change but how this sequestration will respond to anthropogenic nitrogen (N) and phosphorous (P) deposition is uncertain. We couple isotope, soil C fractionation and mesocosm techniques to assess the sequestration of plant‐C inputs, and their partitioning into C pools with different sink potentials, under an experimental gradient of N and P deposition (0, 10, 30, 60 and 100 kg N ha^?1 yr^?1; and 0, 2, 6, 12 and 20 kg P ha^?1 yr^?1). We hypothesized that N deposition would increase sequestration, with the majority of the C being sequestered in faster cycling soil pools because N deposition has been shown to accelerate the turnover of these pools while decelerating the turnover of slower cycling pools. In contrast to this hypothesis, sequestration into all soil C pools peaked at intermediate levels of N deposition. Given that P amendment has been shown to cause a net loss of soil C, we postulated that P deposition would decrease sequestration. This expectation was not supported by our data, with sequestration generally being greater under P deposition. When soils were amended simultaneously with N and P, neither the shape of the sequestration relationship across the deposition gradient, nor the observed sequestration at the majority of the deposition rates, was statistically predictable from the effects of N and P in isolation. The profound nonlinearities we observed, both for total sequestration responses and the partitioning of C into soil pools with different sink potentials, suggests that the rates of N and P deposition to ecosystems will be the critical determinant of whether they enhance or decrease the long‐term sequestration of fresh plant‐C inputs to soils.     

Chemistry and decomposition of litter from Populus tremuloides Michaux grown at elevated atmospheric CO2 and varying N availability

It has been hypothesized that greater production of total nonstructural carbohydrates (TNC) in foliage grown under elevated atmospheric carbon dioxide (CO₂) will result in higher concentrations of defensive compounds in tree leaf litter, possibly leading to reduced rates of decomposition and nutrient cycling in forest ecosystems of the future. To evaluate the effects of elevated atmospheric CO₂ on litter chemistry and decomposition, we performed a 111 day laboratory incubation with leaf litter of trembling aspen (Populus tremuloides Michaux) produced at 36 Pa and 56 Pa CO₂ and two levels of soil nitrogen (N) availability. Decomposition was quantified as microbially respired CO₂ and dissolved organic carbon (DOC) in soil solution, and concentrations of nonstructural carbohydrates, N, carbon (C), and condensed tannins were monitored throughout the incubation. Growth under elevated atmospheric CO₂ did not significantly affect initial litter concentrations of TNC, N, or condensed tannins. Rates of decomposition, measured as both microbially respired CO₂ and DOC did not differ between litter produced under ambient and elevated CO₂. Total C lost from the samples was 38 mg g^?1 litter as respired CO₂ and 138 mg g^?1 litter as DOC, suggesting short‐term pulses of dissolved C in soil solution are important components of the terrestrial C cycle. We conclude that litter chemistry and decomposition in trembling aspen are minimally affected by growth under higher concentrations of CO₂.     

The impact of elevated CO2, increased nitrogen availability and biodiversity on plant tissue quality and decomposition

Elevated CO₂, increased nitrogen (N) deposition and increasing species richness can increase net primary productivity (NPP). However, unless there are comparable changes in decomposition, increases in productivity will most likely be unsustainable. Without comparable increases in decomposition nutrients would accumulate in dead organic matter leading to nutrient limitations that could eventually prohibit additional increases in productivity. To address this issue, we measured aboveground plant and litter quality and belowground root quality, as well as decomposition of aboveground litter for one and 2‐year periods using in situ litterbags in response to a three‐way factorial manipulation of CO₂ (ambient vs. 560 ppm), N deposition (ambient vs. the addition of 4 g N m⁻² yr⁻¹) and plant species richness (one, four, nine and 16 species) in experimental grassland plots. Litter chemistry responded to the CO₂, N and plant diversity treatments, but decomposition was much less responsive. Elevated CO₂ induced decreases in % N and % lignin in plant tissues. N addition led to increases in % N and decreases in % lignin. Increasing plant diversity led to decreases in % N and % lignin and an increase in % cellulose. In contrast to the litter chemistry changes, elevated CO₂ had a much lower impact on decomposition and resulted in only a 2.5% decrease in carbon (C) loss. Detectable responses were not observed either to N addition or to species richness. These results suggest that global change factors such as biodiversity loss, elevated CO₂ and N deposition lead to significant changes in tissue quality; however, the response of decomposition is modest. Thus, the observed increases in productivity at higher diversity levels and with elevated CO₂ and N fertilization are not matched by an increase in decomposition rates. This lack of coupled responses between production and decomposition is likely to result in an accumulation of nutrients in the litter pool which will dampen the response of NPP to these factors over time.     

Interactive effects of elevated CO2 and nitrogen deposition on fatty acid molecular and isotope composition of above‐ and belowground tree biomass and forest soil fractions

Atmospheric carbon dioxide (CO₂) and reactive nitrogen (N) concentrations have been increasing due to human activities and impact the global carbon (C) cycle by affecting plant photosynthesis and decomposition processes in soil. Large amounts of C are stored in plants and soils, but the mechanisms behind the stabilization of plant‐ and microbial‐derived organic matter (OM) in soils are still under debate and it is not clear how N deposition affects soil OM dynamics. Here, we studied the effects of 4 years of elevated (¹³C‐depleted) CO₂ and N deposition in forest ecosystems established in open‐top chambers on composition and turnover of fatty acids (FAs) in plants and soils. FAs served as biomarkers for plant‐ and microbial‐derived OM in soil density fractions. We analyzed above‐ and belowground plant biomass of beech and spruce trees as well as soil density fractions for the total organic C and FA molecular and isotope (δ¹³C) composition. FAs did not accumulate relative to total organic C in fine mineral fractions, showing that FAs are not effectively stabilized by association with soil minerals. The δ¹³C values of FAs in plant biomass increased under high N deposition. However, the N effect was only apparent under elevated CO₂ suggesting a N limitation of the system. In soil fractions, only isotope compositions of short‐chain FAs (C₁₆₊₁₈) were affected. Fractions of ‘new’ (experimental‐derived) FAs were calculated using isotope depletion in elevated CO₂ plots and decreased from free light to fine mineral fractions. ‘New’ FAs were higher in short‐chain compared to long‐chain FAs (C_20?30), indicating a faster turnover of short‐chain compared to long‐chain FAs. Increased N deposition did not significantly affect the quantity of ‘new’ FAs in soil fractions, but showed a tendency of increased amounts of ‘old’ (pre‐experimental) C suggesting that decomposition of ‘old’ C is retarded by high N inputs.     

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

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

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.     

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.     

Decomposition of soil and plant carbon from pasture systems after 9 years of exposure to elevated CO2: impact on C cycling and modeling

Elevated atmospheric CO₂ may alter decomposition rates through changes in plant material quality and through its impact on soil microbial activity. This study examines whether plant material produced under elevated CO₂ decomposes differently from plant material produced under ambient CO₂. Moreover, a long‐term experiment offered a unique opportunity to evaluate assumptions about C cycling under elevated CO₂ made in coupled climate–soil organic matter (SOM) models. Trifolium repens and Lolium perenne plant materials, produced under elevated (60 Pa) and ambient CO₂ at two levels of N fertilizer (140 vs. 560 kg ha^?1 yr^?1), were incubated in soil for 90 days. Soils and plant materials used for the incubation had been exposed to ambient and elevated CO₂ under free air carbon dioxide enrichment conditions and had received the N fertilizer for 9 years. The rate of decomposition of L. perenne and T. repens plant materials was unaffected by elevated atmospheric CO₂ and rate of N fertilization. Increases in L. perenne plant material C : N ratio under elevated CO₂ did not affect decomposition rates of the plant material. If under prolonged elevated CO₂ changes in soil microbial dynamics had occurred, they were not reflected in the rate of decomposition of the plant material. Only soil respiration under L. perenne, with or without incorporation of plant material, from the low‐N fertilization treatment was enhanced after exposure to elevated CO₂. This increase in soil respiration was not reflected in an increase in the microbial biomass of the L. perenne soil. The contribution of old and newly sequestered C to soil respiration, as revealed by the ¹³C‐CO₂ signature, reflected the turnover times of SOM–C pools as described by multipool SOM models. The results do not confirm the assumption of a negative feedback induced in the C cycle following an increase in CO₂, as used in coupled climate–SOM models. Moreover, this study showed no evidence for a positive feedback in the C cycle following additional N fertilization.     

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.     

Faster turnover of new soil carbon inputs under increased atmospheric CO2

Rising levels of atmospheric CO₂ frequently stimulate plant inputs to soil, but the consequences of these changes for soil carbon (C) dynamics are poorly understood. Plant‐derived inputs can accumulate in the soil and become part of the soil C pool (“new soil C”), or accelerate losses of pre‐existing (“old”) soil C. The dynamics of the new and old pools will likely differ and alter the long‐term fate of soil C, but these separate pools, which can be distinguished through isotopic labeling, have not been considered in past syntheses. Using meta‐analysis, we found that while elevated CO₂ (ranging from 550 to 800 parts per million by volume) stimulates the accumulation of new soil C in the short term (<1 year), these effects do not persist in the longer term (1–4 years). Elevated CO₂ does not affect the decomposition or the size of the old soil C pool over either temporal scale. Our results are inconsistent with predictions of conventional soil C models and suggest that elevated CO₂ might increase turnover rates of new soil C. Because increased turnover rates of new soil C limit the potential for additional soil C sequestration, the capacity of land ecosystems to slow the rise in atmospheric CO₂ concentrations may be smaller than previously assumed.     

Effects of plant species diversity, atmospheric [CO2], and N addition on gross rates of inorganic N release from soil organic matter

A significant challenge in predicting terrestrial ecosystem response to global changes comes from the relatively poor understanding of the processes that control pools and fluxes of plant nutrients in soil. In addition, individual global changes are often studied in isolation, despite the potential for interactive effects among them on ecosystem processes. We studied the response of gross N mineralization and microbial respiration after 6 years of application of three global change factors in a grassland field experiment in central Minnesota (the BioCON experiment). BioCON is a factorial manipulation of plant species diversity (1, 4, 9 and 16 prairie species), atmospheric [CO₂] (ambient and elevated: 560 μmol mol^?1), and N inputs (ambient and ambient +4 g N m^?2 yr^?1). We hypothesized that gross N mineralization would increase with increasing levels of all factors because of stimulated plant productivity and thus greater organic inputs to soils. However, we also hypothesized that N addition would enhance, while elevated [CO₂] and greater diversity would temper, gross N mineralization responses because of increased and reduced plant tissue N concentrations, respectively. In partial support of our hypothesis, gross N mineralization increased with greater diversity and N addition, but not with elevated [CO₂]. The ratio of gross N mineralization to microbial respiration (i.e. the ‘yield’ of inorganic N mineralized per unit C respired) declined with greater diversity and [CO₂] suggesting increasing limitation of microbial processes by N relative to C in these treatments. Based on these results, we conclude that the plant supply of organic matter primarily controls gross N mineralization and microbial respiration, but that the concentration of N in organic matter input secondarily influences these processes. Thus, in systems where N limits plant productivity these global change factors could cause different long‐term ecosystem trajectories because of divergent effects on soil N and C cycling.     

Isopod effects on decomposition of litter produced under elevated CO2, N deposition and different soil types

The performance of Oniscus asellus (Isopoda) and its influence on litter mass loss and mineralization was assessed in a microcosm experiment, using beech (Fagus sylvatica) leaf litter that was produced on different soil types, contrasting atmospheric CO₂ concentrations, and different nitrogen deposition rates. Litter quality was significantly altered by these treatments, and many of the CO₂ and N effects differed between soil types. Litter quality affected subsequent litter mass loss rates, microbial respiration, and leaching of dissolved organic carbon (DOC) and nitrate. These effects were largely independent of the presence of isopods, even though isopods highly accelerated litter mass loss, stimulated microbial respiration by 37%, and increased nitrate leaching by 50%. Isopods did not change their relative rates of litter consumption and growth in response to litter quality. Isopod mortality, however, increased with increasing litter lignin/N ratios, and was significantly different between soil types, which may indicate long‐term effects on litter decomposition through altered isopod densities. Having the choice among the litter of three different species [maple (Acer pseudoplatanus), beech (Fagus sylvatica) and oak (Quercus robur)] grown at either ambient or elevated CO₂, isopods preferred maple to beech when all the litter was produced under elevated CO₂. This suggests that beyond changes in consumption, an altered food selection by isopods in a CO₂‐enriched atmosphere may affect the temporal and spatial composition of the litter layer in temperate forests. In contrast to previous findings, we conclude that isopods do not always increase their consumption rates, and hence do not differentially affect microbial decomposition in litter of poorer quality. Nevertheless changes in animal densities and/or shifts in their food preferences, could result in altered decomposition and carbon and nutrient turnover rates.     

Dual,differential isotope labeling shows the preferential movement of labile plant constituents into mineral‐bonded soil organic matter

The formation and stabilization of soil organic matter (SOM) are major concerns in the context of global change for carbon sequestration and soil health. It is presently believed that lignin is not selectively preserved in soil and that chemically labile compounds bonding to minerals comprise a large fraction of the SOM. Labile plant inputs have been suggested to be the main precursor of the mineral‐bonded SOM. Litter decomposition and SOM formation are expected to have temperature sensitivity varying with the lability of plant inputs. We tested this framework using dual ¹³C and ¹⁵N differentially labeled plant material to distinguish the metabolic and structural components within a single plant material. Big Bluestem (Andropogon gerardii) seedlings were grown in an enriched ¹³C and ¹⁵N environment and then prior to harvest, removed from the enriched environment and allowed to incorporate natural abundance ¹³C–CO₂ and ¹⁵N fertilizer into the metabolic plant components. This enabled us to achieve a greater than one atom % difference in ¹³C between the metabolic and structural components within the plant litter. This differentially labeled litter was incubated in soil at 15 and 35 °C, for 386 days with CO₂ measured throughout the incubation. After 14, 28, 147, and 386 days of incubation, the soil was subsequently fractionated. There was no difference in temperature sensitivity of the metabolic and structural components with regard to how much was respired or in the amount of litter biomass stabilized. Only the metabolic litter component was found in the sand, silt, or clay fraction while the structural component was exclusively found in the light fraction. These results support the stabilization framework that labile plant components are the main precursor of mineral‐associated organic matter.     

Short-term decomposition of litter produced by plants grown in ambient and elevated atmospheric CO2 concentrations

The effects of elevated atmospheric CO₂ (ambient + 340 μmol mol^–1) on above-ground litter decomposition were investigated over a 6-week period using a field-based mesocosm system. Soil respiratory activity in mesocosms incubated in ambient and elevated atmospheric CO₂ concentrations were not significantly different (t-test, P > 0.05) indicating that there were no direct effects of elevated atmospheric CO₂ on litter decomposition. A study of the indirect effects of CO₂ on soil respiration showed that soil mesocosms to which naturally senescent plant litter had been added (0.5% w/w) from the C₃ sedge Scirpus olneyi grown in elevated atmospheric CO₂ was reduced by an average of 17% throughout the study when compared to soil mesocosms to which litter from Scirpus olneyi grown in ambient conditions had been added. In contrast, similar experiments using senescent material from the C₄ grass Spartina patens showed no difference in soil respiration rates between mesocosms to which litter from plants grown in elevated or ambient CO₂ conditions had been added. Analysis of the C:N ratio and lignin content of the senescent material showed that, while the C:N ratio and lignin content of the Spartina patens litter did not vary with atmospheric CO₂ conditions, the C:N ratio (but not the lignin content) of the litter from Scirpus olneyi was significantly greater (t-test;P < 0.05) when derived from plants grown under elevated CO₂ (105:1 compared to 86:1 for litter derived from Scirpus olneyi grown under ambient conditions). The results suggest that the increased C:N ratio of the litter from the C₃ plant Scirpus olneyi grown under elevated CO₂ led to the lower rates of biodegradation observed as reduced soil respiration in the mesocosms. Further long-term experiments are now required to determine the effects of elevated CO₂ on C partitioning in terrestrial ecosystems.