Understanding the differential impact of PFOS and F-53B pollution on reed-mediated soil organic matter decomposition: Insights from rhizosphere priming effect.

Plants affect soil microorganisms through the release of root exudates under pollution stress. This process may affect rhizosphere priming effect (RPE) and alter the rate of soil organic matter decomposition. However, the influence of plants on the decomposition of organic matter in soil subjected to pollution stress remains unclear. We studied the effects of exposure to perfluorooctanesulfonic (PFOS) and its alternative, chlorinated polyfluoroalkyl ether sulfonic (F-53B), at concentrations of 0.1 mg/kg and 50 mg/kg on the RPE of reed. We conducted our experiments in an artificial climate chamber and used the natural 13C tracer method to determine RPE. In the PFOS-exposed groups, the RPE was negative, with values of -11.45 mg C kg-1 soil d-1 in the low PFOS group and -8.04 mg C kg-1 soil d-1 in the high PFOS group. In contrast, in the F-53B-exposed groups, the RPE was positive, with values of 8.26 mg C kg-1 soil d-1 in the low F-53B group and 12.18 mg C kg-1 soil d-1 in the high F-53B group. Exposure of reeds to PFOS/F-53B stress resulted in differential effects on extracellular enzyme activities. The observed positive and negative RPE phenomena could be attributed to variations in extracellular enzyme activities. In conclusion, RPE responded differently under PFOS/F-53B exposure.

Diverse factors influence the amounts of carbon input to soils via rhizodeposition in plants: A review.

Rhizodeposition encompasses the intricate processes through which plants generate organic compounds via photosynthesis, store these compounds within aboveground biomass and roots through top-down transport, and subsequently release this organic matter into the soil. Rhizodeposition represents one of the carbon (C) cycle in soils that can achieve long-term organic C sequestration. This function holds significant implications for mitigating the climate change that partly stems from the greenhouse effect associated with increased atmospheric carbon dioxide levels. Therefore, it is essential to further understand how the process of rhizodeposition allocates the photosynthetic C that plants create via photosynthesis. While many studies have explored the basic principles of rhizodeposition, along with the associated impact on soil C storage, there is a palpable absence of comprehensive reviews that summarize the various factors influencing this process. This paper compiles and analyzes the literature on plant rhizodeposition to describe how rhizodeposition influences soil C storage. Moreover, the review summarizes the impacts of soil physicochemical, microbial, and environmental characteristics on plant rhizodeposition and priming effects, and concludes with recommendations for future research.

Effects of nitrogen addition on rhizosphere priming: The role of stoichiometric imbalance.

Nitrogen (N) input has a significant impact on the availability of carbon (C), nitrogen (N), and phosphorus (P) in the rhizosphere, leading to an imbalanced stoichiometry in microbial demands. This imbalance can result in energy or nutrient limitations, which, in turn, affect C dynamics during plant growth. However, the precise influence of N addition on the C:N:P imbalance ratio and its subsequent effects on rhizosphere priming effects (RPEs) remain unclear. To address this gap, we conducted a 75-day microcosm experiment, varying N addition rates (0, 150, 300 kg N ha-1), to examine how microbes regulate RPE by adapting to stoichiometry and maintaining homeostasis in response to N addition, using the 13C natural method. Our result showed that N input induced a stoichiometric imbalance in C:N:P, leading to P or C limitation for microbes during plant growth. Microbes responded by adjusting enzymatic stoichiometry and functional taxa to preserve homeostasis, thereby modifying the threshold element ratios (TERs) to cope with the C:N:P imbalance. Microbes adapted to the stoichiometric imbalance by reducing TER, which was attributed to a reduction in carbon use efficiency. Consequently, we observed higher RPE under P limitation, whereas the opposite trend was observed under C or N limitation. These results offer novel insights into the microbial regulation of RPE variation under different soil nutrient conditions and contribute to a better understanding of soil C dynamics.

Deciphering the dual role of bacterial communities in stabilizing rhizosphere priming effect under intra-annual change of growing seasons.

The rhizosphere priming effect (RPE) is a widely observed phenomenon affecting carbon (C) turnover in plant-soil systems. While multiple cropping and seasonal changes can have significant impacts on RPE, the mechanisms driving these processes are complex and not yet fully understood. Here, we planted maize in paddy soil during two growing seasons having substantial temperature differences [May-August (warm season, 26.6 °C) and September-November (cool season, 23.1 °C)] within the same calendar year in southern China to examine how seasonal changes affect RPEs and soil C. We identified sources of C emissions by quantifying the natural abundance of 13C and determined microbial metabolic limitations or efficiency and functional genes related to C cycling using an enzyme-based biogeochemical equilibrium model and high-throughput quantitative PCR-based chip technology, respectively. Results showed that microbial metabolism was mainly limited by phosphorus in the warm season, but by C in the cool season, resulting in positive RPEs in both growing seasons, but no significant differences (9.02 vs. 6.27 mg C kg-1 soil day-1). The RPE intensity remained stable as temperature increased (warm season compared to a cool season), which can be largely explained by the simultaneous increase in the abundance of functional genes related to both C degradation and fixation. Our study highlights the simultaneous response and adaptation of microbial communities to seasonal changes and hence contributes to an understanding and prediction of microbially mediated soil C turnover under multiple cropping systems.

Arctic rooting depth distribution influences modelled carbon emissions but cannot be inferred from aboveground vegetation type.

The distribution of roots throughout the soil drives depth-dependent plant-soil interactions and ecosystem processes, particularly in arctic tundra where plant biomass, is predominantly belowground. Vegetation is usually classified from aboveground, but it is unclear whether such classifications are suitable to estimate belowground attributes and their consequences, such as rooting depth distribution and its influence on carbon cycling. We performed a meta-analysis of 55 published arctic rooting depth profiles, testing for differences both between distributions based on aboveground vegetation types (Graminoid, Wetland, Erect-shrub, and Prostrate-shrub tundra) and between 'Root Profile Types' for which we defined three representative and contrasting clusters. We further analyzed potential impacts of these different rooting depth distributions on rhizosphere priming-induced carbon losses from tundra soils. Rooting depth distribution hardly differed between aboveground vegetation types but varied between Root Profile Types. Accordingly, modelled priming-induced carbon emissions were similar between aboveground vegetation types when they were applied to the entire tundra, but ranged from 7.2 to 17.6 Pg C cumulative emissions until 2100 between individual Root Profile Types. Variations in rooting depth distribution are important for the circumpolar tundra carbon-climate feedback but can currently not be inferred adequately from aboveground vegetation type classifications.

Soil microbial community structure dynamics shape the rhizosphere priming effect patterns in the paddy soil.

Microbial community structure plays a crucial part in soil organic carbon (SOC) decomposition and variation of rhizosphere priming effects (RPEs) during plant growth. However, it is still uncertain how bacterial community structure regulates RPEs in soil and how RPE patterns respond to plant growth. Therefore, we conducted an experiment to examine the RPE response to plant growth and nitrogen (N) addition (0 (N0), 150 (N150), and 300 (N300) kg N ha-1) using the 13C natural abundance method in a C3 soil (paddy soil) - C4 plant (maize, Zea mays L.) system; we then explored the underlying biotic mechanisms using 16S rRNA sequencing techniques. Networks were constructed to identify keystone taxa and to analyze the correlations between network functional modules of bacterial community and C decomposition. The results indicated that negative and positive RPEs occurred on Day 30 and Day 75 after maize planting, respectively. Bacterial community structure significantly changed and tended to shift from r-strategists toward K-strategists with changing labile C: N stoichiometry and soil pH during plant growth stages. The different network modules of bacterial community were aggregated in response to RPE pattern variation. Caulobacteraceae, Bacillus, and Chitinophagaceae were keystone taxa on Day 30, while Gemmatimonas, Candidatus Koribacter, and Xanthobacteraceae were keystone taxa on Day 75. Moreover, keystone taxa with different C utilization strategies were significantly different between the two growth stages and related closely to different RPE patterns. This study provides deeper insights into the network structure of bacterial communities corresponding to RPE patterns and emphasizes the significance of keystone taxa in RPE variation.

A distinct sensitivity to the priming effect between labile and stable soil organic carbon.

Soil organic carbon (SOC) is a mixture of various carbon (C) compounds with different stability, which can be distinctly affected by the priming effect (PE). However, little is known about how the PE changes with SOC stability. We address this issue by combining results from two experiments and a metaanalysis. We found that the PE increased with the prolongation of soil preincubation, suggesting that higher PE occurred for more stable SOC than for labile SOC. This was further supported by the metaanalysis of 42 observations. There were significant negative relationships between the difference in PE (ΔPE) between labile and more stable SOC and their differences in SOC, microbial biomass C and soil C : N ratio, indicating that soil C availability exerts a vital control on ΔPE. We conclude that, compared with labile SOC, stable SOC can be more vulnerable to priming once microbes are provided with exogenous C substrates. This high vulnerability of stable SOC to priming warrants more attention in future studies on SOC cycling and global change.

Root litter decomposition is suppressed in species mixtures and in the presence of living roots.

Plant species diversity and identity can significantly modify litter decomposition, but the underlying mechanisms remain elusive, particularly for root litter. Here, we aimed to disentangle the mechanisms by which plant species diversity alters root litter decomposition. We hypothesised that (1) interactions between species in mixed communities result in litter that decomposes faster than litter produced in monocultures; (2) litter decomposition is accelerated in the presence of living plants, especially when the litter and living plant identities are matched (known as home-field advantage).Monocultures and a mixture of four common grassland species were established to obtain individual litter and a 'natural' root litter mixture. An 'artificial' mixed litter was created using litter from monocultures, mixed in the same proportions as the species composition in the natural litter mixtures based on qPCR measurements. These six root litter types were incubated in four monocultures, a four-species mixture and an unplanted soil.Root decomposition was strongly affected by root litter identity and the presence, but not diversity, of living roots. Mixed-species litter decomposed slower than expected based on the decomposition of single-species litters. In addition, the presence of living roots suppressed decomposition independent of the match between litter and living plant identities. Decomposition was not significantly different between the 'natural' and 'artificial' root litter mixtures, indicating that root-root interactions in species mixtures did not affect root chemical quality. Synthesis. Suppressed decomposition in the presence of living roots indicates that interactions between microbial communities associated with living roots and root litter control root litter decomposition. As we found no support for the importance of home-field advantage or interspecific root interactions in modifying decomposition, suppressed decomposition of mixed-species litter seems to be primarily driven by chemical rather than biotic interactions.

Belowground Carbon Efficiency for Nitrogen and Phosphorus Acquisition Varies Between Lolium perenne and Trifolium repens and Depends on Phosphorus Fertilization.

Photosynthetically derived carbon (C) is allocated belowground, allowing plants to obtain nutrients. However, less is known about the amount of nutrients acquired relative to the C allocated belowground, which is referred to as C efficiency for nutrient acquisition (CENA). Here, we examined how C efficiency for nitrogen (N) and phosphorus (P) acquisition varied between ryegrass (Lolium perenne) and clover (Trifolium repens) with and without P fertilization. A continuous 13C-labeling method was applied to track belowground C allocation. Both species allocated nearly half of belowground C to rhizosphere respiration (49%), followed by root biomass (37%), and rhizodeposition (14%). With regard to N and P, CENA was higher for clover than for ryegrass, which remained higher after accounting for relatively low C costs associated with biological N2 fixation. Phosphorus fertilization increased the C efficiency for P acquisition but decreased the C efficiency for N acquisition. A higher CENA for N and P in clover may be attributed to the greater rhizosphere priming on soil organic matter decomposition. Increased P availability with P fertilization could induce lower C allocation for P uptake but exacerbate soil N limitation, thereby making N uptake less C efficient. Overall, our study revealed that species-specific belowground C allocation and nutrient uptake efficiency depend on which nutrient is limited.

Shift of microbial turnover time and metabolic efficiency strongly regulates rhizosphere priming effect under nitrogen fertilization in paddy soil.

Microbial turnover and the decomposition of soil organic matter can be stimulated by living roots in a phenomenon known as the rhizosphere priming effect (RPE). Both the microbial turnover time (MTT) and metabolic efficiency are closely related to RPE. However, changes in MTT, metabolic efficiency and RPE in response to nitrogen (N) fertilization at different levels and the associations between these factors during plant growth are unknown. The effects of N fertilization at different levels (0, 150 and 300 kg N ha-1) on RPE and the underlying mechanisms were investigated in maize (Zea mays L.) grown in paddy soil using a 13Carbon (C) natural abundance method. The RPE varied from -1.49 to 15.93 mg C kg-1 soil day-1, with significant effects at different levels of N fertilization, growth stages and interactions between these factors. Nitrogen fertilization reduced microbial C:N imbalance and soil pH. During the plant growth periods, the RPE was initially low because the microbes preferentially utilized plant-derived C, but later increased due to trade-offs between microbial N acquisition and acidity stress alleviation under N fertilization. The soil microbes altered their MTT and metabolic efficiency with changes in the microbial community structure to maintain stoichiometric homeostasis and adapt to acidity stress. RPE was lowest whereas MTT and metabolic efficiency were highest with N fertilization at 150 kg N ha-1. Changes in MTT and metabolic efficiency explained 84.5% of the variations in the RPE, and the latter had greater impact (55.8%) than the former (28.7%). Changes in MTT and metabolic efficiency to cope with microbial resource acquisition and acidity stress under N fertilization represent an important pathway for RPE regulation in paddy soil. These findings highlight the significance of MTT and metabolic efficiency in RPE regulation for optimization of the N fertilization level to mitigate soil C losses.

[Effect of Water Management on Rice Growth and Rhizosphere Priming Effect in Paddy Soils].

The rhizosphere priming effect (RPE) caused by carbon inputs from crop rhizodeposits plays a key role in regulating the carbon emission flux and carbon balance of farmland soils. Due to frequent alternations between dry and wet conditions, CO2 and CH4 emissions and the RPE in paddy field ecosystems are significantly different to those of other ecosystems. Therefore, it is of great significance to determine the direction and intensity of the rice RPE under alternations of dry and wet to limit greenhouse gas emissions. In this study, using a 13C-CO2 continuous labeling method combined with a pot-based experiment, the response of rice growth and the RPE under alternating dry and wet and continuous flooding conditions was examined. The results showed that, compared with the continuous flooding treatment, the alternating dry and wet treatments significantly increased aboveground and root biomass and the root-to-root ratio, and also increased soil microbial biomass. Under continuous flooding conditions, fluxes of 13CO2 and 13CH4 increased with rice growth from 10.2 µg·(kg·h)-1 and 2.8 µg·(kg·h)-1 (63 d) to 16.0 µg·(kg·h)-1 and 3.2 µg·(kg·h)-1 (75 d), respectively. During the 12-day drying process, the emissions of 13CO2 and 13CH4 derived from rhizosphere deposited C decreased by 57.5% and 88.1%. Under continuous flooding conditions, the RPE for CO2 and CH4 were positive and increased with the growth of rice. Under the alternating dry and wet treatment, after 12 days of drying, the RPE for CO2 and CH4 was reduced from 0.29 mg·(kg·h)-1 and 12.3 µg·(kg·h)-1 (63 d) to -0.39 mg·(kg·h)-1 and 0.07 µg·(kg·h)-1 (75 d). Thus, alternating wet and dry treatment can effectively promote rice growth and reduce the cumulative emissions of CH4. Therefore, adopting appropriate field water management is of great significance for increasing rice yields and mitigating greenhouse gas emissions.

Root effects on soil organic carbon: a double-edged sword.

From recent developments on how roots affect soil organic carbon (SOC) an apparent paradox has emerged where roots drive SOC stabilization causing SOC accrual, but also SOC destabilization causing SOC loss. We synthesize current results and propose the new Rhizo-Engine framework consisting of two linked components: microbial turnover and the soil physicochemical matrix. The Rhizo-Engine is driven by rhizodeposition, root turnover, and plant uptake of nutrients and water, thereby accelerating SOC turnover through both stabilization and destabilization mechanisms. This Rhizo-Engine framework emphasizes the need for a more holistic approach to study root-driven SOC dynamics. This framework would provide better understanding of plant root effects on soil carbon sequestration and the sensitivity of SOC stocks to climate and land-use changes.

Harnessing PGPR inoculation through exogenous foliar application of salicylic acid and microbial extracts for improving rice growth.

Plant-growth-promoting rhizobacteria (PGPR) should effectively colonize along the plant root to enhance the plant and soil health. The present investigation aims to improve the PGPR-mediated plant health benefits through above-ground foliar management. A green fluorescent protein-tagged PGPR strain, Pseudomonas chlororaphis (ZSB15-M2) was inoculated in a nonautoclaved agricultural soil before rice culturing. Salicylic acid and cell extracts of Corynebacterium glutamicum and Saccharomyces cerevisiae as a supply of hormonal and inducer compounds were applied on the foliage of the 10-days-old rice plants and subsequently observed the colonizing ability of ZSB15-M2. The cell extracts of Corynebacteria and yeast showed a 100-fold increase in the ZSB15-M2 population in the rhizosphere of rice, whereas salicylic acid had a 10-fold increase in relation to mock control. The rice root exudates collected after the spraying of salicylic acid and microbial extracts showed significantly enhanced release of total carbon, total protein, total sugar, total amino nitrogen, total nitrogen, and phenol content. In vitro assays revealed that these root exudates collected after exogenous spray of these chemicals enhanced the chemotactic motility and biofilm formation of ZSB15-M2 compared to the control plant's root exudate. Metabolomic analysis of root exudates collected from these rice plants by gas chromatography-mass spectrometry revealed that the Corynebacteria and yeast cell extracts enhanced the divergence of metabolites of rice root exudate. Further, due to these cumulative effects in the rice rhizosphere, the total chlorophyll, total protein, total nitrogen, and total phosphorus of rice were significantly improved. These observations provide insights into the rhizosphere functioning of rice plants as modulated by above-ground treatments with improved colonization of inoculant strains as well as the plant growth.

Rhizosphere control of soil nitrogen cycling: a key component of plant economic strategies.

Understanding how plant species influence soil nutrient cycling is a major theme in terrestrial ecosystem ecology. However, the prevailing paradigm has mostly focused on litter decomposition, while rhizosphere effects on soil organic matter (SOM) decomposition have attracted little attention. Using a dual 13 C/15 N labeling approach in a 'common garden' glasshouse experiment, we investigated how the economic strategies of 12 grassland plant species (graminoids, forbs and legumes) drive soil nitrogen (N) cycling via rhizosphere processes, and how this in turn affects plant N acquisition and growth. Acquisitive species with higher photosynthesis, carbon rhizodeposition and N uptake than conservative species induced a stronger acceleration of soil N cycling through rhizosphere priming of SOM decomposition. This allowed them to take up larger amounts of N and allocate it above ground to promote photosynthesis, thereby sustaining their faster growth. The N2 -fixation ability of legumes enhanced rhizosphere priming by promoting photosynthesis and rhizodeposition. Our study demonstrates that the economic strategies of plant species regulate a plant-soil carbon-nitrogen feedback operating through the rhizosphere. These findings provide novel mechanistic insights into how plant species with contrasting economic strategies sustain their nutrition and growth through regulating the cycling of nutrients by soil microbes in their rhizosphere.

Rhizosphere priming of two near-isogenic wheat lines varying in citrate efflux under different levels of phosphorus supply.

BACKGROUNDS AND AIMS: The rhizosphere priming effect (RPE) has been explained from the perspective of microbial responses to root exudates and nutrient availability. This study introduced a chemical process that could also contribute to RPE: root exudates (organic acid ligands) could liberate mineral-protected carbon (C) in soil for microbial degradation. METHODS: Wheat (Triticum aestivum L.) near-isogenic lines varying in citrate efflux were grown for 6 weeks in a C4 soil supplied with either low (10 µg g-1) or high P (40 µg g-1). Total below-ground CO2 was trapped and partitioned for determination of soil organic C decomposition and RPE using a stable isotopic tracing technique. Mineral dissolution was examined by incubating soil with citric ligand at a series of concentrations. KEY RESULTS: High P increased RPE (81 %), shoot (32 %) and root biomass (57 %), root-derived CO2-C (20 %), microbial biomass C (28 %) and N (100%), soil respiration (20 %) and concentrations of water-extractable P (30 %), Fe (43 %) and Al (190 %), but decreased inorganic N in the rhizosphere. Compared with Egret-Burke, wheat line Egret-Burke TaMATE1B with citrate efflux had lower inorganic N, microbial biomass C (16 %) and N (30 %) in the rhizosphere but greater RPE (18 %), shoot biomass (12 %) and root-derived CO2-C (low P 36 %, high P 13 %). Egret-Burke TaMATE1B also had higher concentrations of water-extractable P, Fe and Al in the rhizosphere, indicating the release of mineral-protected C. In addition, citrate ligand facilitated Fe and Al release from soil, with their concentrations rising with increasing ligand concentration and incubation time. CONCLUSIONS: While high P supply increased microbial growth and RPE possibly due to higher total root exudation, citrate efflux from the root might have facilitated the liberation of mineral-bound C, leading to the higher RPE under Egret-Burke TaMATE1B. Mineral dissolution may be an important process that regulates RPE and should be considered in future RPE research.

Vascular plant-mediated controls on atmospheric carbon assimilation and peat carbon decomposition under climate change.

Climate change can alter peatland plant community composition by promoting the growth of vascular plants. How such vegetation change affects peatland carbon dynamics remains, however, unclear. In order to assess the effect of vegetation change on carbon uptake and release, we performed a vascular plant-removal experiment in two Sphagnum-dominated peatlands that represent contrasting stages of natural vegetation succession along a climatic gradient. Periodic measurements of net ecosystem CO2 exchange revealed that vascular plants play a crucial role in assuring the potential for net carbon uptake, particularly with a warmer climate. The presence of vascular plants, however, also increased ecosystem respiration, and by using the seasonal variation of respired CO2 radiocarbon (bomb-14 C) signature we demonstrate an enhanced heterotrophic decomposition of peat carbon due to rhizosphere priming. The observed rhizosphere priming of peat carbon decomposition was matched by more advanced humification of dissolved organic matter, which remained apparent beyond the plant growing season. Our results underline the relevance of rhizosphere priming in peatlands, especially when assessing the future carbon sink function of peatlands undergoing a shift in vegetation community composition in association with climate change.

Relationship between fine-root exudation and respiration of two Quercus species in a Japanese temperate forest.

Plants allocate a considerable amount of carbon (C) to fine roots as respiration and exudation. Fine-root exudation could stimulate microbial activity, which further contributes to soil heterotrophic respiration. Although both root respiration and exudation are important components of belowground C cycling, how they relate to each other is less well known. In this study, we aimed to explore this relationship on mature trees growing in the field. The measurements were performed on two canopy species, Quercus serrata Thunb. and Quercus glauca, in a warm temperate forest. The respiration and exudation rates of the same fine-root segment were measured in parallel with a syringe-basis incubation and a closed static chamber, respectively. We also measured root traits and ectomycorrhizal colonization ratio because these indexes commonly relate to root respiration and reflect root physiology. The microbial activity enhanced by root exudation was investigated by comparing the dissolved organic carbon (DOC) and microbial biomass carbon (MBC) between rhizosphere soils and bulk soils. Mean DOC concentration and MBC were ca two times higher in the rhizosphere soils and positively related to exudation rates, indicating that exudation further relates to the C dynamics in the soils. Flux rates of exudation and respiration were positively correlated with each other. Both root exudation and respiration rates positively related to ectomycorrhizal colonization and root tissue nitrogen, and therefore the relationship between the two fluxes may be attributed to fine-root activity. The flux rates of root respiration were 8.7 and 10.5 times as much as those of exudation on a root-length basis and a root-weight basis, respectively. In spite of the fact that flux rates of respiration and exudation varied enormously among the fine-root segments of the two Quercus species, exudation was in proportion to respiration. This result gives new insight into the fine-root C-allocation strategy and the belowground C dynamics.

The rhizosphere and hyphosphere differ in their impacts on carbon and nitrogen cycling in forests exposed to elevated CO2.

While multiple experiments have demonstrated that trees exposed to elevated CO2 can stimulate microbes to release nutrients from soil organic matter, the importance of root- versus mycorrhizal-induced changes in soil processes are presently unknown. We analyzed the contribution of roots and mycorrhizal activities to carbon (C) and nitrogen (N) turnover in a loblolly pine (Pinus taeda) forest exposed to elevated CO2 by measuring extracellular enzyme activities at soil microsites accessed via root windows. Specifically, we quantified enzyme activity from soil adjacent to root tips (rhizosphere), soil adjacent to hyphal tips (hyphosphere), and bulk soil. During the peak growing season, CO2 enrichment induced a greater increase of N-releasing enzymes in the rhizosphere (215% increase) than in the hyphosphere (36% increase), but a greater increase of recalcitrant C-degrading enzymes in the hyphosphere (118%) than in the rhizosphere (19%). Nitrogen fertilization influenced the magnitude of CO2 effects on enzyme activities in the rhizosphere only. At the ecosystem scale, the rhizosphere accounted for c. 50% and 40% of the total activity of N- and C-releasing enzymes, respectively. Collectively, our results suggest that root exudates may contribute more to accelerated N cycling under elevated CO2 at this site, while mycorrhizal fungi may contribute more to soil C degradation.