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.     

Accelerated belowground C cycling in a managed agriforest ecosystem exposed to elevated carbon dioxide concentrations

We investigated the effects of three elevated atmospheric CO₂ levels on a Populus deltoides plantation at Biosphere 2 Laboratory in Oracle Arizona. Stable isotopes of carbon have been used as tracers to separate the carbon present before the CO₂ treatments started (old C), from that fixed after CO₂ treatments began (new C). Tree growth at elevated [CO₂] increased inputs to soil organic matter (SOM) by increasing the production of fine roots and accelerating the rate of root C turnover. However, soil carbon content decreased as [CO₂] in the atmosphere increased and inputs of new C were not found in SOM. Consequently, the rates of soil respiration increased by 141% and 176% in the 800 and 1200 μL L^?1 plantations, respectively, when compared with ambient [CO₂] after 4 years of exposure. However, the increase in decomposition of old SOM (i.e. already present when CO₂ treatments began) accounted for 72% and 69% of the increase in soil respiration seen under elevated [CO₂]. This resulted in a net loss of soil C at a rate that was between 10 and 20 times faster at elevated [CO₂] than at ambient conditions. The inability to retain new and old C in the soil may stem from the lack of stabilization of SOM, allowing for its rapid decomposition by soil heterotrophs.     

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.     

Soil carbon and nitrogen cycling and storage throughout the soil profile in a sweetgum plantation after 11 years of CO2‐enrichment

Increased partitioning of carbon (C) to fine roots under elevated [CO₂], especially deep in the soil profile, could alter soil C and nitrogen (N) cycling in forests. After more than 11 years of free‐air CO₂ enrichment in a Liquidambar styraciflua L. (sweetgum) plantation in Oak Ridge, TN, USA, greater inputs of fine roots resulted in the incorporation of new C (i.e., C with a depleted δ¹³C) into root‐derived particulate organic matter (POM) pools to 90‐cm depth. Even though production in the sweetgum stand was limited by soil N availability, soil C and N contents were greater throughout the soil profile under elevated [CO₂] at the conclusion of the experiment. Greater C inputs from fine‐root detritus under elevated [CO₂] did not result in increased net N immobilization or C mineralization rates in long‐term laboratory incubations, possibly because microbial biomass was lower in the CO₂‐enriched plots. Furthermore, the δ¹³CO₂ of the C mineralized from the incubated soil closely tracked the δ¹³C of the labile POM pool in the elevated [CO₂] treatment, especially in shallower soil, and did not indicate significant priming of the decomposition of pre‐experiment soil organic matter (SOM). Although potential C mineralization rates were positively and linearly related to total SOM C content in the top 30 cm of soil, this relationship did not hold in deeper soil. Taken together with an increased mean residence time of C in deeper soil pools, these findings indicate that C inputs from relatively deep roots under elevated [CO₂] may increase the potential for long‐term soil C storage. However, C in deeper soil is likely to take many years to accrue to a significant fraction of total soil C given relatively smaller root inputs at depth. Expanded representation of biogeochemical cycling throughout the soil profile may improve model projections of future forest responses to rising atmospheric [CO₂].     

Soil carbon storage under simulated climate change is mediated by plant functional type

The stability of soil organic matter (SOM) pools exposed to elevated CO₂ and warming has not been evaluated adequately in long‐term experiments and represents a substantial source of uncertainty in predicting ecosystem feedbacks to climate change. We conducted a 6‐year experiment combining free‐air CO₂ enrichment (FACE, 550 ppm) and warming (+2 °C) to evaluate changes in SOM accumulation in native Australian grassland. In this system, competitive interactions appear to favor C₄ over C₃ species under FACE and warming. We therefore investigated the role of plant functional type (FT) on biomass and SOM responses to the long‐term treatments by carefully sampling soil under patches of C₃‐ and C₄‐dominated vegetation. We used physical fractionation to quantify particulate organic matter (POM) and long‐term incubation to assess potential decomposition rates. Aboveground production of C₄ grasses increased in response to FACE, but total root biomass declined. Across treatments, C : N ratios were higher in leaves, roots and POM of C₄ vegetation. CO₂ and temperature treatments interacted with FT to influence SOM, and especially POM, such that soil carbon was increased by warming under C₄ vegetation, but not in combination with elevated CO₂. Potential decomposition rates increased in response to FACE and decreased with warming, possibly owing to treatment effects on soil moisture and microbial community composition. Decomposition was also inversely correlated with root N concentration, suggesting increased microbial demand for older, N‐rich SOM in treatments with low root N inputs. This research suggests that C₃–C₄ vegetation responses to future climate conditions will strongly influence SOM storage in temperate grasslands.     

An invasive wetland grass primes deep soil carbon pools

Understanding the processes that control deep soil carbon (C) dynamics and accumulation is of key importance, given the relevance of soil organic matter (SOM) as a vast C pool and climate change buffer. Methodological constraints of measuring SOM decomposition in the field prevent the addressing of real‐time rhizosphere effects that regulate nutrient cycling and SOM decomposition. An invasive lineage of Phragmites australis roots deeper than native vegetation (Schoenoplectus americanus and Spartina patens) in coastal marshes of North America and has potential to dramatically alter C cycling and accumulation in these ecosystems. To evaluate the effect of deep rooting on SOM decomposition we designed a mesocosm experiment that differentiates between plant‐derived, surface SOM‐derived (0–40 cm, active root zone of native marsh vegetation), and deep SOM‐derived mineralization (40–80 cm, below active root zone of native vegetation). We found invasive P. australis allocated the highest proportion of roots in deeper soils, differing significantly from the native vegetation in root : shoot ratio and belowground biomass allocation. About half of the CO₂ produced came from plant tissue mineralization in invasive and native communities; the rest of the CO₂ was produced from SOM mineralization (priming). Under P. australis, 35% of the CO₂ was produced from deep SOM priming and 9% from surface SOM. In the native community, 9% was produced from deep SOM priming and 44% from surface SOM. SOM priming in the native community was proportional to belowground biomass, while P. australis showed much higher priming with less belowground biomass. If P. australis deep rooting favors the decomposition of deep‐buried SOM accumulated under native vegetation, P. australis invasion into a wetland could fundamentally change SOM dynamics and lead to the loss of the C pool that was previously sequestered at depth under the native vegetation, thereby altering the function of a wetland as a long‐term C sink.     

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.     

Carbon budgeting in plant–soil mesocosms under elevated CO2: locally missing carbon?

Studies have suggested that more carbon is fixed due to a large increase in photosynthesis in plant–soil systems exposed to elevated CO₂ than could subsequently be found in plant biomass and soils –‐ the locally missing carbon phenomenon. To further understand this phenomenon, an experiment was carried out using EcoCELLs which are open‐flow, mass‐balance systems at the mesocosm scale. Naturally occurring ¹³C tracers were also used to separately measure plant‐derived carbon and soil‐derived carbon. The experiment included two EcoCELLs, one under ambient atmospheric CO₂ and the other under elevated CO₂ (ambient plus 350 μL L^{? 1}). By matching carbon fluxes with carbon pools, the issue of locally missing carbon was investigated. Flux‐based net primary production (NPP_f) was similar to pool‐based primary production (NPP_p) under ambient CO₂, and the discrepancy between the two carbon budgets (12 g C m^{? 2}, or 4% of NPP_f) was less than measurement errors. Therefore, virtually all carbon entering the system under ambient CO₂ was accounted for at the end of the experiment. Under elevated CO₂, however, the amount of NPP_f was much higher than NPP_p, resulting in missing carbon of approximately 80 g C m^{? 2} or 19% of NPP_f which was much higher than measurement errors. This was additional to the 96% increase in rhizosphere respiration and the 50% increase in root growth, two important components of locally missing carbon. The mystery of locally missing carbon under elevated CO₂ remains to be further investigated. Volatile organic carbon, carbon loss due to root washing, and measurement errors are discussed as some of the potential contributing factors.     

Elevated atmospheric CO2 and feedback between carbon and nitrogen cycles

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

Fine root chemistry and decomposition in model communities of north-temperate tree species show little response to elevated atmospheric CO<Subscript>2</Subscript> and varying soil resource availability

Rising atmospheric [CO₂] has the potential to alter soil carbon (C) cycling by increasing the content of recalcitrant constituents in plant litter, thereby decreasing rates of decomposition. Because fine root turnover constitutes a large fraction of annual NPP, changes in fine root decomposition are especially important. These responses will likely be affected by soil resource availability and the life history characteristics of the dominant tree species. We evaluated the effects of elevated atmospheric [CO₂] and soil resource availability on the production and chemistry, mycorrhizal colonization, and decomposition of fine roots in an early- and late-successional tree species that are economically and ecologically important in north temperate forests. Open-top chambers were used to expose young trembling aspen (Populus tremuloides) and sugar maple (Acer saccharum) trees to ambient (36 Pa) and elevated (56 Pa) atmospheric CO₂. Soil resource availability was composed of two treatments that bracketed the range found in the Upper Lake States, USA. After 2.5 years of growth, sugar maple had greater fine root standing crop due to relatively greater allocation to fine roots (30% of total root biomass) relative to aspen (7% total root biomass). Relative to the low soil resources treatment, aspen fine root biomass increased 76% with increased soil resource availability, but only under elevated [CO₂]. Sugar maple fine root biomass increased 26% with increased soil resource availability (relative to the low soil resources treatment), and showed little response to elevated [CO₂]. Concentrations of N and soluble phenolics, and C/N ratio in roots were similar for the two species, but aspen had slightly higher lignin and lower condensed tannins contents compared to sugar maple. As predicted by source-sink models of carbon allocation, pooled constituents (C/N ratio, soluble phenolics) increased in response to increased relative carbon availability (elevated [CO₂]/low soil resource availability), however, biosynthetically distinct compounds (lignin, starch, condensed tannins) did not always respond as predicted. We found that mycorrhizal colonization of fine roots was not strongly affected by atmospheric [CO₂] or soil resource availability, as indicated by root ergosterol contents. Overall, absolute changes in root chemical composition in response to increases in C and soil resource availability were small and had no effect on soil fungal biomass or specific rates of fine root decomposition. We conclude that root contributions to soil carbon cycling will mainly be influenced by fine root production and turnover responses to rising atmospheric [CO₂], rather than changes in substrate chemistry.     

Salinity and sea level mediate elevated CO2 effects on C3–C4 plant interactions and tissue nitrogen in a Chesapeake Bay tidal wetland

Elevated atmospheric carbon dioxide concentrations ([CO₂]) generally increase plant photosynthesis in C₃ species, but not in C₄ species, and reduce stomatal conductance in both C₃ and C₄ plants. In addition, tissue nitrogen concentration ([N]) often fails to keep pace with enhanced carbon gain under elevated CO₂, particularly in C₃ species. While these responses are well documented in many species, implications for plant growth and nutrient cycling in native ecosystems are not clear. Here we present data on 18 years of measurement of above and belowground biomass, tissue [N] and total standing crop of N for a Scirpus olneyi‐dominated (C₃ sedge) community, a Spartina patens‐dominated (C₄ grass) community and a C₃–C₄‐mixed species community exposed to ambient and elevated (ambient +340 ppm) atmospheric [CO₂] in natural salinity and sea level conditions of a Chesapeake Bay wetland. Increased biomass production (shoots plus roots) under elevated [CO₂] in the S. olneyi‐dominated community was sustained throughout the study, averaging approximately 35%, while no significant effect of elevated [CO₂] was found for total biomass in the C₄‐dominated community. We found a significant decline in C₄ biomass (correlated with rising sea level) and a concomitant increase in C₃ biomass in the mixed community. This shift from C₄ to C₃ was accelerated by the elevated [CO₂] treatment. The elevated [CO₂] stimulation of total biomass accumulation was greatest during rainy, low salinity years: the average increase above the ambient treatment during the three wettest years (1994, 1996, 2003) was 2.9 t ha⁻¹ but in the three driest years (1995, 1999, 2002), it was 1.2 t ha⁻¹. Elevated [CO₂] depressed tissue [N] in both species, but especially in the S. olneyi where the relative depression was positively correlated with salinity and negatively related with the relative enhancement of total biomass production. Thus, the greatest amount of carbon was added to the S. olneyi‐dominated community during years when shoot [N] was reduced the most, suggesting that the availability of N was not the most or even the main limitation to elevated [CO₂] stimulation of carbon accumulation in this ecosystem.     

Mycorrhizal Hyphal Turnover as a Dominant Process for Carbon Input into Soil Organic Matter

The atmospheric concentration of CO₂ is predicted to reach double current levels by 2075. Detritus from aboveground and belowground plant parts constitutes the primary source of C for soil organic matter (SOM), and accumulation of SOM in forests may provide a significant mechanism to mitigate increasing atmospheric CO₂ concentrations. In a poplar (three species) plantation exposed to ambient (380 ppm) and elevated (580 ppm) atmospheric CO₂ concentrations using a Free Air Carbon Dioxide Enrichment (FACE) system, the relative importance of leaf litter decomposition, fine root and fungal turnover for C incorporation into SOM was investigated. A technique using cores of soil in which a C₄ crop has been grown (δ¹³C −18.1‰) inserted into the plantation and detritus from C₃ trees (δ¹³C −27 to −30‰) was used to distinguish between old (native soil) and new (tree derived) soil C. In-growth cores using a fine mesh (39 μm) to prevent in-growth of roots, but allow in-growth of fungal hyphae were used to assess contribution of fine roots and the mycorrhizal external mycelium to soil C during a period of three growing seasons (1999–2001). Across all species and treatments, the mycorrhizal external mycelium was the dominant pathway (62%) through which carbon entered the SOM pool, exceeding the input via leaf litter and fine root turnover. The input via the mycorrhizal external mycelium was not influenced by elevated CO₂, but elevated atmospheric CO₂ enhanced soil C inputs via fine root turnover. The turnover of the mycorrhizal external mycelium may be a fundamental mechanism for the transfer of root-derived C to SOM.     

Plants mediate soil organic matter decomposition in response to sea level rise

Tidal marshes have a large capacity for producing and storing organic matter, making their role in the global carbon budget disproportionate to land area. Most of the organic matter stored in these systems is in soils where it contributes 2–5 times more to surface accretion than an equal mass of minerals. Soil organic matter (SOM) sequestration is the primary process by which tidal marshes become perched high in the tidal frame, decreasing their vulnerability to accelerated relative sea level rise (RSLR). Plant growth responses to RSLR are well understood and represented in century‐scale forecast models of soil surface elevation change. We understand far less about the response of SOM decomposition to accelerated RSLR. Here we quantified the effects of flooding depth and duration on SOM decomposition by exposing planted and unplanted field‐based mesocosms to experimentally manipulated relative sea level over two consecutive growing seasons. SOM decomposition was quantified as CO₂ efflux, with plant‐ and SOM‐derived CO₂ separated via δ¹³CO₂. Despite the dominant paradigm that decomposition rates are inversely related to flooding, SOM decomposition in the absence of plants was not sensitive to flooding depth and duration. The presence of plants had a dramatic effect on SOM decomposition, increasing SOM‐derived CO₂ flux by up to 267% and 125% (in 2012 and 2013, respectively) compared to unplanted controls in the two growing seasons. Furthermore, plant stimulation of SOM decomposition was strongly and positively related to plant biomass and in particular aboveground biomass. We conclude that SOM decomposition rates are not directly driven by relative sea level and its effect on oxygen diffusion through soil, but indirectly by plant responses to relative sea level. If this result applies more generally to tidal wetlands, it has important implications for models of SOM accumulation and surface elevation change in response to accelerated RSLR.     

Quantification of soil carbon inputs under elevated CO2: C3 plants in a C4 soil

The objective of this investigation was to quantify the differences in soil carbon stores after exposure of birch seedlings (Betula pendula Roth.) over one growing season to ambient and elevated carbon dioxide concentrations. One-year-old seedling of birch were transplanted to pots containing C₄ soil derived from beneath a maize crop, and placed in ambient (350 L L^–1) and elevated (600 L L^–1) plots in a free-air carbon dioxide enrichment (FACE) experiment. After 186 days the plants and soils were destructively sampled, and analysed for differences in root and stem biomass, total plant tissue and soil C contents and ¹³C values. The trees showed a significant increase (+50%) in root biomass, but stem and leaf biomasses were not significantly affected by treatment. C isotope analyses of leaves and fine roots showed that the isotopic signal from the ambient and elevated CO₂ supply was sufficiently distinct from that of the C₄ soil to enable quantification of net root C input to the soil under both ambient and elevated CO₂. After 186 days, the pots under ambient conditions contained 3.5 g of C as intact root material, and had gained an additional 0.6 g C added to the soil through root exudation/turnover; comparable figures for the pots under elevated CO₂ were 5.9 g C and 1.5 g C, respectively. These data confirm the importance of soils as an enhanced sink for C under elevated atmospheric CO₂ concentrations. We propose the use of C₄ soils in elevated CO₂ experiments as an important technique for the quantification of root net C inputs under both ambient and elevated CO₂ treatments.     

Estimating soil carbon sequestration under elevated CO2 by combining carbon isotope labelling with soil carbon cycle modelling

Elevated CO₂ concentrations generally stimulate grassland productivity, but herbaceous plants have only a limited capacity to sequester extra carbon (C) in biomass. However, increased primary productivity under elevated CO₂ could result in increased transfer of C into soils where it could be stored for prolonged periods and exercise a negative feedback on the rise in atmospheric CO₂. Measuring soil C sequestration directly is notoriously difficult for a number of methodological reasons. Here, we present a method that combines C isotope labelling with soil C cycle modelling to partition net soil sequestration into changes in new C fixed over the experimental duration (C_new) and pre‐experimental C (C_old). This partitioning is advantageous because the C_new accumulates whereas C_old is lost in the course of time (ΔC_new>0 whereas ΔC_old<0). We applied this method to calcareous grassland exposed to 600 μL CO₂ L^?1 for 6 years. The CO₂ used for atmospheric enrichment was depleted in ¹³C relative to the background atmosphere, and this distinct isotopic signature was used to quantify net soil C_new fluxes under elevated CO₂. Using ¹³C/¹²C mass balance and inverse modelling, the Rothamsted model ‘RothC’ predicted gross soil C_new inputs under elevated CO₂ and the decomposition of C_old. The modelled soil C pools and fluxes were in good agreement with experimental data. C isotope data indicated a net sequestration of ≈90 g C_new m^?2 yr^?1 in elevated CO₂. Accounting for C_old‐losses, this figure was reduced to ≈30 g C m^?2 yr^?1 at elevated CO₂; the elevated CO₂‐effect on net C sequestration was in the range of≈10 g C m^?2 yr^?1. A sensitivity and error analysis suggests that the modelled data are relatively robust. However, elevated CO₂‐specific mechanisms may necessitate a separate parameterization at ambient and elevated CO₂; these include increased soil moisture due to reduced leaf conductance, soil disaggregation as a consequence of increased soil moisture, and priming effects. These effects could accelerate decomposition of C_old in elevated CO₂ so that the CO₂ enrichment effect may be zero or even negative. Overall, our findings suggest that the C sequestration potential of this grassland under elevated CO₂ is rather limited.     

Seasonal soil‐surface carbon fluxes from the root systems of young Pinus radiata trees growing at ambient and elevated CO2 concentration

Elevated atmospheric CO₂ concentration may result in increased below‐ground carbon allocation by trees, thereby altering soil carbon cycling. Seasonal estimates of soil surface carbon flux were made to determine whether carbon losses from Pinus radiata trees growing at elevated CO₂ concentration were higher than those at ambient CO₂ concentration, and whether this was related to increased fine root growth. Monthly soil surface carbon flux density (f) measurements were made on plots with trees growing at ambient (350) and elevated (650 μmol mol^?1) CO₂ concentration in large open‐top chambers. Prior to planting the soil carbon concentration (0.1%) and f (0.28 μmol m^?2 s^?1 at 15 °C) were low. A function describing the radial pattern of f with distance from tree stems was used to estimate the annual carbon flux from tree plots. Seasonal estimates of fine root production were made from minirhizotrons and the radial distribution of roots compared with radial measurements of f. A one‐dimensional gas diffusion model was used to estimate f from soil CO₂ concentrations at four depths. For the second year of growth, the annual carbon flux from the plots was 1671 g y^?1 and 1895 g y^?1 at ambient and elevated CO₂ concentrations, respectively, although this was not a significant difference. Higher f at elevated CO₂ concentration was largely explained by increased fine root biomass. Fine root biomass and stem production were both positively related to f. Both root length density and f declined exponentially with distance from the stem, and had similar length scales. Diurnal changes in f were largely explained by changes in soil temperature at a depth of 0.05 m. Ignoring the change of f with increasing distance from tree stems when scaling to a unit ground area basis from measurements with individual trees could result in under‐ or overestimates of soil‐surface carbon fluxes, especially in young stands when fine roots are unevenly distributed.     

Root production and mortality under elevated atmospheric carbon dioxide

An essential component of an understanding of carbon flux is the quantification of movement through the root carbon pool. Although estimates have been made using radiocarbon, the use of minirhizotrons provides a direct measurement of rates of root birth and death. We have measured root demographic parameters under a semi-natural grassland and for wheat. The grassland was studied along a natural altitudinal gradient in northern England, and similar turf from the site was grown in elevated CO₂ in solardomes. Root biomass was enhanced under elevated CO₂. Root birth and death rates were both increased to a similar extent in elevated CO₂, so that the throughput of carbon was greater than in ambient CO₂, but root half-lives were shorter under elevated CO₂ only under a Juncus/Nardus sward on a peaty gley soil, and not under a Festuca turf on a brown earth soil. In a separate experiment, wheat also responded to elevated CO₂ by increased root production, and there was a marked shift towards surface rooting: root development at a depth of 80–85 cm was both reduced and delayed. In conjunction with published results for trees, these data suggest that the impact of elevated CO₂ will be system-dependent, affecting the spatio-temporal pattern of root growth in some ecosystems and the rate of turnover in others. Turrnover is also sensitive to temperature, soil fertility and other environmental variables, all of which are likely to change in tandem with atmospheric CO₂ concentrations. Differences in turnover and time and location of rhizodeposition may have a large effect on rates of carbon cycling.     

The influence of elevated atmospheric CO2 on fine root dynamics in an intact temperate forest

Root dynamics are important for plant, ecosystem and global carbon cycling. Changes in root dynamics caused by rising atmospheric CO₂ not only have the potential to moderate further CO₂ increases, but will likely affect forest function. We used FACE (Free‐Air CO₂ Enrichment) to expose three 30‐m diameter plots in a 13‐year‐old loblolly pine (Pinus taeda) forest to elevated (ambient + 200 µL L^?1) atmospheric CO₂. Three identical fully instrumented plots were implemented as controls (ambient air only). We quantified root dynamics from October 1998 to October 1999 using minirhizotrons. In spite of 16% greater root lengths and 24% more roots per minirhizotron tube, the effects of elevated atmospheric CO₂ on root lengths and numbers were not statistically significant. Similarly, production and mortality were also unaffected by the CO₂ treatment, even though annual root production and mortality were 26% and 46% greater in elevated compared to ambient CO₂ plots. Average diameters of live roots present at the shallowest soil depth were, however, significantly enhanced in CO₂‐enriched plots. Mortality decreased with increasing soil depth and the slopes of linear regression lines (mortality vs. depth) differed between elevated and ambient CO₂ treatments, reflecting the significant CO₂ by depth interaction. Relative root turnover (root flux/live root pool) was unchanged by exposure to elevated atmospheric CO₂. Results from this study suggest modest, if any, increases in ecosystem‐level root productivity in CO₂‐enriched environments.     

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.     

Influence of rhizodeposition under elevated CO2 on plant nutrition and soil organic matter

Atmospheric CO₂ concentrations can influence ecosystem carbon storage through net primary production (NPP), soil carbon storage, or both. In assessing the potential for carbon storage in terrestrial ecosystems under elevated CO₂, both NPP and processing of soil organic matter (SOM), as well as the multiple links between them, must be examined. Within this context, both the quantity and quality of carbon flux from roots to soil are important, since roots produce specialized compounds that enhance nutrient acquisition (affecting NPP), and since the flux of organic compounds from roots to soil fuels soil microbial activity (affecting processing of SOM).From the perspective of root physiology, a technique is described which uses genetically engineered bacteria to detect the distribution and amount of flux of particular compounds from single roots to non-sterile soils. Other experiments from several labs are noted which explore effects of elevated CO₂ on root acid phosphatase, phosphomonoesterase, and citrate production, all associated with phosphorus nutrition. From a soil perspective, effects of elevated CO₂ on the processing of SOM developed under a C4 grassland but planted with C3 California grassland species were examined under low (unamended) and high (amended with 20 g m^–2 NPK) nutrients; measurements of soil atmosphere 13C combined with soil respiration rates show that during vegetative growth in February, elevated CO₂ decreased respiration of carbon from C4 SOM in high nutrient soils but not in unamended soils.This emphasis on the impacts of carbon loss from roots on both NPP and SOM processing will be essential to understanding terrestrial ecosystem carbon storage under changing atmospheric CO₂ concentrations.Abbreviations SOM soil organic matter - NPP net primary productivity - NEP net ecosystem productivity - PNPP p-nitrophenyl phosphate