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.     

Experimental fire increases soil carbon dioxide efflux in a grassland long‐term multifactor global change experiment

Numerous studies have demonstrated that soil respiration rates increase under experimental warming, although the long‐term, multiyear dynamics of this feedback are not well constrained. Less is known about the effects of single, punctuated events in combination with other longer‐duration anthropogenic influences on the dynamics of soil carbon (C) loss. In 2012 and 2013, we assessed the effects of decadal‐scale anthropogenic global change – warming, increased nitrogen (N) deposition, elevated carbon dioxide (CO₂), and increased precipitation – on soil respiration rates in an annual‐dominated Mediterranean grassland. We also investigated how controlled fire and an artificial wet‐up event, in combination with exposure to the longer‐duration anthropogenic global change factors, influenced the dynamics of C cycling in this system. Decade‐duration surface soil warming (1–2 °C) had no effect on soil respiration rates, while +N addition and elevated CO₂ concentrations increased growing‐season soil CO₂ efflux rates by increasing annual aboveground net primary production (NPP) and belowground fine root production, respectively. Low‐intensity experimental fire significantly elevated soil CO₂ efflux rates in the next growing season. Based on mixed‐effects modeling and structural equation modeling, low‐intensity fire increased growing‐season soil respiration rates through a combination of three mechanisms: large increases in soil temperature (3–5 °C), significant increases in fine root production, and elevated aboveground NPP. Our study shows that in ecosystems where soil respiration has acclimated to moderate warming, further increases in soil temperature can stimulate greater soil CO₂ efflux. We also demonstrate that punctuated short‐duration events such as fire can influence soil C dynamics with implications for both the parameterization of earth system models (ESMs) and the implementation of climate change mitigation policies that involve land‐sector C accounting.     

Increased soil release of greenhouse gases shrinks terrestrial carbon uptake enhancement under warming

Warming can accelerate the decomposition of soil organic matter and stimulate the release of soil greenhouse gases (GHGs), but to what extent soil release of methane (CH₄) and nitrous oxide (N₂O) may contribute to soil C loss for driving climate change under warming remains unresolved. By synthesizing 1,845 measurements from 164 peer‐reviewed publications, we show that around 1.5°C (1.16–2.01°C) of experimental warming significantly stimulates soil respiration by 12.9%, N₂O emissions by 35.2%, CH₄ emissions by 23.4% from rice paddies, and by 37.5% from natural wetlands. Rising temperature increases CH₄ uptake of upland soils by 13.8%. Warming‐enhanced emission of soil CH₄ and N₂O corresponds to an overall source strength of 1.19, 1.84, and 3.12 Pg CO₂‐equivalent/year under 1°C, 1.5°C, and 2°C warming scenarios, respectively, interacting with soil C loss of 1.60 Pg CO₂/year in terms of contribution to climate change. The warming‐induced rise in soil CH₄ and N₂O emissions (1.84 Pg CO₂‐equivalent/year) could reduce mitigation potential of terrestrial net ecosystem production by 8.3% (NEP, 22.25 Pg CO₂/year) under warming. Soil respiration and CH₄ release are intensified following the mean warming threshold of 1.5°C scenario, as compared to soil CH₄ uptake and N₂O release with a reduced and less positive response, respectively. Soil C loss increases to a larger extent under soil warming than under canopy air warming. Warming‐raised emission of soil GHG increases with the intensity of temperature rise but decreases with the extension of experimental duration. This synthesis takes the lead to quantify the ecosystem C and N cycling in response to warming and advances our capacity to predict terrestrial feedback to climate change under projected warming scenarios.     

Individual and interactive effects of warming and nitrogen supply on CO2 fluxes and carbon allocation in subarctic grassland

Climate warming has been suggested to impact high latitude grasslands severely, potentially causing considerable carbon (C) losses from soil. Warming can also stimulate nitrogen (N) turnover, but it is largely unclear whether and how altered N availability impacts belowground C dynamics. Even less is known about the individual and interactive effects of warming and N availability on the fate of recently photosynthesized C in soil. On a 10-year geothermal warming gradient in Iceland, we studied the effects of soil warming and N addition on CO₂ fluxes and the fate of recently photosynthesized C through CO₂ flux measurements and a ¹³CO₂ pulse-labeling experiment. Under warming, ecosystem respiration exceeded maximum gross primary productivity, causing increased net CO₂ emissions. N addition treatments revealed that, surprisingly, the plants in the warmed soil were N limited, which constrained primary productivity and decreased recently assimilated C in shoots and roots. In soil, microbes were increasingly C limited under warming and increased microbial uptake of recent C. Soil respiration was increased by warming and was fueled by increased belowground inputs and turnover of recently photosynthesized C. Our findings suggest that a decade of warming seemed to have induced a N limitation in plants and a C limitation by soil microbes. This caused a decrease in net ecosystem CO₂ uptake and accelerated the respiratory release of photosynthesized C, which decreased the C sequestration potential of the grassland. Our study highlights the importance of belowground C allocation and C-N interactions in the C dynamics of subarctic ecosystems in a warmer world.     

Predominant role of water in regulating soil and microbial respiration and their responses to climate change in a semiarid grassland

Climate change can profoundly impact carbon (C) cycling of terrestrial ecosystems. A field experiment was conducted to examine responses of total soil and microbial respiration, and microbial biomass to experimental warming and increased precipitation in a semiarid temperate steppe in northern China since April 2005. We measured soil respiration twice a month over the growing seasons, soil microbial biomass C (MBC) and N (MBN), microbial respiration (MR) once a year in the middle growing season from 2005 to 2007. The results showed that interannual variations in soil respiration, MR, and microbial biomass were positively related to interannual fluctuations in precipitation. Laboratory incubation with a soil moisture gradient revealed a constraint of the temperature responses of MR by low soil moisture contents. Across the 3 years, experimental warming decreased soil moisture, and consequently caused significant reductions in total and microbial respiration, and microbial biomass, suggesting stronger negatively indirect effects through warming‐induced water stress than the positively direct effects of elevated temperature. Increased evapotranspiration under experimental warming could have reduced soil water availability below a stress threshold, thus leading to suppression of plant growth, root and microbial activities. Increased precipitation significantly stimulated total soil and microbial respiration and all other microbial parameters and the positive precipitation effects increased over time. Our results suggest that soil water availability is more important than temperature in regulating soil and microbial respiratory processes, microbial biomass and their responses to climate change in the semiarid temperate steppe. Experimental warming caused greater reductions in soil respiration than in gross ecosystem productivity (GEP). In contrast, increased precipitation stimulated GEP more than soil respiration. Our observations suggest that climate warming may cause net C losses, whereas increased precipitation may lead to net C gains in the semiarid temperate steppe. Our findings highlight that unless there is concurrent increase in precipitation, the temperate steppe in the arid and semiarid regions of northern China may act as a net C source under climate warming.     

Effects of Warming and Nitrogen Addition on Plant Photosynthate Partitioning in an Alpine Meadow on the Tibetan Plateau

Quantifying plant carbon (C) allocation among different pools is critical for understanding and predicting how C turnover responds to global climate change in terrestrial ecosystems. A field experiment with increasing warming and nitrogen (N) was established to investigate interactive effects on plant C allocation in alpine meadows. Open-top chambers (OTCs) were used to simulate warming. In OTCs, daytime air and soil temperature at 5 cm depth increased by 2.0 and 1.6 °C, respectively, compared with ambient conditions, but soil moisture at 5 cm depth decreased by 4.95% (v/v) from 2012 to 2014. Warming reduced aboveground biomass by 38, 36, and 43% in 2012, 2013, and 2014, respectively, and increased belowground biomass by 64% and 29% in 2013 and 2014, respectively, and the root-to-shoot ratio was significantly increased. Specifically, warming increased the proportion of plant roots in the deep layers (10–20 cm). Both N addition and its combination with warming substantially enhanced belowground biomass. Pulse-labeling experiments for ¹³C revealed that warming reduced the translocation of assimilated C to shoots by 8.8% (38.7% in warming, and 47.5% in the control [CK]), and increased the allocation to root by 12.2% (55.5% in warming, and 43.3% in CK) after 28 days labeling. However, N addition increased the proportion of assimilated C allocated to shoots by 6.5% (54.0% in N addition, and 47.5% in CK), whereas warming combined with N addition reduced this proportion by 10.9%. A decline in soil water content in the surface layer may be the main cause of plants allocating more newly fixed photosynthate to roots. Therefore, plants promoted root growth to draw water from deeper soil layers (10–20 cm). We concluded that climate warming will change the allocation patterns of plant photosynthates by affecting soil water availability, whereas N addition will increase plant photosynthates aboveground in alpine meadows and thus will significantly affect C turnover under future climate change scenarios.

     

Short‐term soil carbon sink potential of oil palm plantations

Oil palm plantations cover ≈14.6 million ha worldwide and the total area under cultivation is expected to increase during the 21st century . Indonesia and Malaysia together account for 87% of global palm oil production and the combined harvested area in these countries has expanded by 6.5 million ha since 1990. Despite this, soil C cycling in oil palm systems is not well quantified but such information is needed for C budget inventories. We quantified soil C storage (root biomass, soil organic matter (SOM) and microbial biomass) and losses [potential soil respiration (R_s) and soil surface CO₂ flux (F_s)] in mineral soils from an oil palm plantation chronosequence (11–34 years since planting) in Selangor, Malaysia. There were no significant effects of plantation age on SOM, microbial biomass, R_s or F_s, implying soil C was in dynamic equilibrium over the chronosequence. However, there was a significant increase in root biomass with plantation age, indicating a short‐term C sink. Across the chronosequence, R_s was driven by soil moisture, soil particle size, root biomass and soil microbial biomass N but not microbial biomass C. This suggests that the nutrient status of the microbial community may be of equal or greater importance for soil CO₂ losses than substrate availability and also raises particular concerns regarding the addition of nitrogenous fertilizer, i.e. increased yields will be associated with increased soil CO₂ emissions. To fully assess the impact of oil palm plantations on soil C storage, initial soil C losses following land conversion (e.g. from native forest or other previous plantations) must be accounted for. If initial soil C losses are large, our data show that there is no accumulation of stable C in the soil as the plantation matures and hence the conversion to oil palm would probably represent a net loss of soil C.     

Soil [N] modulates soil C cycling in CO2‐fumigated tree stands: a meta‐analysis

Under elevated atmospheric CO₂ concentrations, soil carbon (C) inputs are typically enhanced, suggesting larger soil C sequestration potential. However, soil C losses also increase and progressive nitrogen (N) limitation to plant growth may reduce the CO₂ effect on soil C inputs with time. We compiled a data set from 131 manipulation experiments, and used meta‐analysis to test the hypotheses that: (1) elevated atmospheric CO₂ stimulates soil C inputs more than C losses, resulting in increasing soil C stocks; and (2) that these responses are modulated by N. Our results confirm that elevated CO₂ induces a C allocation shift towards below‐ground biomass compartments. However, the increased soil C inputs were offset by increased heterotrophic respiration (Rh), such that soil C content was not affected by elevated CO₂. Soil N concentration strongly interacted with CO₂ fumigation: the effect of elevated CO₂ on fine root biomass and –production and on microbial activity increased with increasing soil N concentration, while the effect on soil C content decreased with increasing soil N concentration. These results suggest that both plant growth and microbial activity responses to elevated CO₂ are modulated by N availability, and that it is essential to account for soil N concentration in C cycling analyses.     

Experimental warming and burn severity alter soil CO2 flux and soil functional groups in a recently burned boreal forest

Global warming is projected to be greatest in northern regions, where forest fires are also increasing in frequency. Thus, interactions between fire and temperature on soil respiration at high latitudes should be considered in determining feedbacks to climate. We tested the hypothesis that experimental warming will augment soil CO₂ flux in a recently burned boreal forest by promoting microbial and root growth, but that this increase will be less apparent in more severely burned areas. We used open‐top chambers to raise temperatures 0.4–0.9°C across two levels of burn severity in a fire scar in Alaskan black spruce forest. After 3 consecutive years of warming, soil respiration was measured through a portable gas exchange system. Abundance of active microbes was determined by using Biolog EcoPlates^? for bacteria and ergosterol analysis for fungi. Elevated temperatures increased soil CO₂ flux by 20% and reduced root biomass, but had no effect on bacterial or fungal abundance or soil organic matter (SOM) content. Soil respiration, fungal abundance, SOM, and root biomass decreased with increasing burn severity. There were no significant interactions between temperature and burn severity with respect to any measurement. Higher soil respiration rates in the warmed plots may be because of higher metabolic activity of microbes or roots. All together, we found that postfire soils are a greater source of CO₂ to the atmosphere under elevated temperatures even in severely burned areas, suggesting that global warming may produce a positive feedback to atmospheric CO₂, even in young boreal ecosystems.     

Simple additive effects are rare: a quantitative review of plant biomass and soil process responses to combined manipulations of CO2 and temperature

In recent years, increased awareness of the potential interactions between rising atmospheric CO₂ concentrations ([ CO₂ ]) and temperature has illustrated the importance of multifactorial ecosystem manipulation experiments for validating Earth System models. To address the urgent need for increased understanding of responses in multifactorial experiments, this article synthesizes how ecosystem productivity and soil processes respond to combined warming and [ CO₂ ] manipulation, and compares it with those obtained in single factor [ CO₂ ] and temperature manipulation experiments. Across all combined elevated [ CO₂ ] and warming experiments, biomass production and soil respiration were typically enhanced. Responses to the combined treatment were more similar to those in the [ CO₂ ]‐only treatment than to those in the warming‐only treatment. In contrast to warming‐only experiments, both the combined and the [ CO₂ ]‐only treatments elicited larger stimulation of fine root biomass than of aboveground biomass, consistently stimulated soil respiration, and decreased foliar nitrogen (N) concentration. Nonetheless, mineral N availability declined less in the combined treatment than in the [ CO₂ ]‐only treatment, possibly due to the warming‐induced acceleration of decomposition, implying that progressive nitrogen limitation (PNL) may not occur as commonly as anticipated from single factor [ CO₂ ] treatment studies. Responses of total plant biomass, especially of aboveground biomass, revealed antagonistic interactions between elevated [ CO₂ ] and warming, i.e. the response to the combined treatment was usually less‐than‐additive. This implies that productivity projections might be overestimated when models are parameterized based on single factor responses. Our results highlight the need for more (and especially more long‐term) multifactor manipulation experiments. Because single factor CO₂ responses often dominated over warming responses in the combined treatments, our results also suggest that projected responses to future global warming in Earth System models should not be parameterized using single factor warming experiments.     

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.     

Differential effects of warming and nitrogen fertilization on soil respiration and microbial dynamics in switchgrass croplands

The mechanistic understanding of warming and nitrogen (N) fertilization, alone or in combination, on microbially mediated decomposition is limited. In this study, soil samples were collected from previously harvested switchgrass (Panicum virgatum L.) plots that had been treated with high N fertilizer (HN: 67 kg N ha^?1) and those that had received no N fertilizer (NN) over a 3‐year period. The samples were incubated for 180 days at 15 °C and 20 °C, during which heterotrophic respiration, δ¹³C of CO₂, microbial biomass (MB), specific soil respiration rate (R_s: respiration per unit of microbial biomass), and exoenzyme activities were quantified at 10 different collections time. Employing switchgrass tissues (referred to as litter) with naturally abundant ¹³C allowed us to partition CO₂ respiration derived from soil and amended litter. Cumulative soil respiration increased significantly by 16.4% and 4.2% under warming and N fertilization, respectively. Respiration derived from soil was elevated significantly with warming, while oxidase, the agent for recalcitrant soil substrate decomposition, was not significantly affected by warming. Warming, however, significantly enhanced MB and R_s indicating a decrease in microbial growth efficiency (MGE). On the contrary, respiration derived from amended litter was elevated with N fertilization, which was consistent with the significantly elevated hydrolase. N fertilization, however, had little effect on MB and R_s, suggesting little change in microbial physiology. Temperature and N fertilization showed minimal interactive effects likely due to little differences in soil N availability between NN and HN samples, which is partly attributable to switchgrass biomass N accumulation (equivalent to ~53% of fertilizer N). Overall, the differential individual effects of warming and N fertilization may be driven by physiological adaptation and stimulated exoenzyme kinetics, respectively. The study shed insights on distinct microbial acquisition of different substrates under global temperature increase and N enrichment.     

Combined effects of atmospheric CO<Subscript>2</Subscript> and N availability on the belowground carbon and nitrogen dynamics of aspen mesocosms

It is uncertain whether elevated atmospheric CO₂ will increase C storage in terrestrial ecosystems without concomitant increases in plant access to N. Elevated CO₂ may alter microbial activities that regulate soil N availability by changing the amount or composition of organic substrates produced by roots. Our objective was to determine the potential for elevated CO₂ to change N availability in an experimental plant-soil system by affecting the acquisition of root-derived C by soil microbes. We grew Populus tremuloides (trembling aspen) cuttings for 2 years under two levels of atmospheric CO₂ (36.7 and 71.5 Pa) and at two levels of soil N (210 and 970 μg N g^–1). Ambient and twice-ambient CO₂ concentrations were applied using open-top chambers, and soil N availability was manipulated by mixing soils differing in organic N content. From June to October of the second growing season, we measured midday rates of soil respiration. In August, we pulse-labeled plants with ¹⁴CO₂ and measured soil ¹⁴CO₂ respiration and the ¹⁴C contents of plants, soils, and microorganisms after a 6-day chase period. In conjunction with the August radio-labeling and again in October, we used ¹⁵N pool dilution techniques to measure in situ rates of gross N mineralization, N immobilization by microbes, and plant N uptake. At both levels of soil N availability, elevated CO₂ significantly increased whole-plant and root biomass, and marginally increased whole-plant N capital. Significant increases in soil respiration were closely linked to increases in root biomass under elevated CO₂. CO₂ enrichment had no significant effect on the allometric distribution of biomass or ¹⁴C among plant components, total ¹⁴C allocation belowground, or cumulative (6-day) ¹⁴CO₂ soil respiration. Elevated CO₂ significantly increased microbial ¹⁴C contents, indicating greater availability of microbial substrates derived from roots. The near doubling of microbial ¹⁴C contents at elevated CO₂ was a relatively small quantitative change in the belowground C cycle of our experimental system, but represents an ecologically significant effect on the dynamics of microbial growth. Rates of plant N uptake during both 6-day periods in August and October were significantly greater at elevated CO₂, and were closely related to fine-root biomass. Gross N mineralization was not affected by elevated CO₂. Despite significantly greater rates of N immobilization under elevated CO₂, standing pools of microbial N were not affected by elevated CO₂, suggesting that N was cycling through microbes more rapidly. Our results contained elements of both positive and negative feedback hypotheses, and may be most relevant to young, aggrading ecosystems, where soil resources are not yet fully exploited by plant roots. If the turnover of microbial N increases, higher rates of N immobilization may not decrease N availability to plants under elevated CO₂. Received: 12 February 1999 / Accepted: 2 March 2000     

Disentangling root responses to climate change in a semiarid grassland

Future ecosystem properties of grasslands will be driven largely by belowground biomass responses to climate change, which are challenging to understand due to experimental and technical constraints. We used a multi-faceted approach to explore single and combined impacts of elevated CO₂ and warming on root carbon (C) and nitrogen (N) dynamics in a temperate, semiarid, native grassland at the Prairie Heating and CO₂ Enrichment experiment. To investigate the indirect, moisture mediated effects of elevated CO₂, we included an irrigation treatment. We assessed root standing mass, morphology, residence time and seasonal appearance/disappearance of community-aggregated roots, as well as mass and N losses during decomposition of two dominant grass species (a C₃ and a C₄). In contrast to what is common in mesic grasslands, greater root standing mass under elevated CO₂ resulted from increased production, unmatched by disappearance. Elevated CO₂ plus warming produced roots that were longer, thinner and had greater surface area, which, together with greater standing biomass, could potentially alter root function and dynamics. Decomposition increased under environmental conditions generated by elevated CO₂, but not those generated by warming, likely due to soil desiccation with warming. Elevated CO₂, particularly under warming, slowed N release from C₄—but not C₃—roots, and consequently could indirectly affect N availability through treatment effects on species composition. Elevated CO₂ and warming effects on root morphology and decomposition could offset increased C inputs from greater root biomass, thereby limiting future net C accrual in this semiarid grassland.     

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.     

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}     

Vegetation feedbacks of nutrient addition lead to a weaker carbon sink in an ombrotrophic bog

To study vegetation feedbacks of nutrient addition on carbon sequestration capacity, we investigated vegetation and ecosystem CO₂ exchange at Mer Bleue Bog, Canada in plots that had been fertilized with nitrogen (N) or with N plus phosphorus (P) and potassium (K) for 7–12 years. Gross photosynthesis, ecosystem respiration, and net CO₂ exchange were measured weekly during May–September 2011 using climate‐controlled chambers. A substrate‐induced respiration technique was used to determine the functional ability of the microbial community. The highest N and NPK additions were associated with 40% less net CO₂ uptake than the control. In the NPK additions, a diminished C sink potential was due to a 20–30% increase in ecosystem respiration, while gross photosynthesis rates did not change as greater vascular plant biomass compensated for the decrease in Sphagnum mosses. In the highest N‐only treatment, small reductions in gross photosynthesis and no change in ecosystem respiration led to the reduced C sink. Substrate‐induced microbial respiration was significantly higher in all levels of NPK additions compared with control. The temperature sensitivity of respiration in the plots was lower with increasing cumulative N load, suggesting more labile sources of respired CO₂. The weaker C sink potential could be explained by changes in nutrient availability, higher woody : foliar ratio, moss loss, and enhanced decomposition. Stronger responses to NPK fertilization than to N‐only fertilization for both shrub biomass production and decomposition suggest that the bog ecosystem is N‐P/K colimited rather than N‐limited. Negative effects of further N‐only deposition were indicated by delayed spring CO₂ uptake. In contrast to forests, increased wood formation and surface litter accumulation in bogs seem to reduce the C sink potential owing to the loss of peat‐forming Sphagnum.     

Intensified plant N and C pool with more available nitrogen under experimental warming in an alpine meadow ecosystem

Nitrogen (N) availability is projected to increase in a warming climate. But whether the more available N is immobilized by microbes (thus stimulates soil carbon (C) decomposition), or is absorbed by plants (thus intensifies C uptake) remains unknown in the alpine meadow ecosystem. Infrared heaters were used to simulate climate warming with a paired experimental design. Soil ammonification, nitrification, and net mineralization were obtained by in situ incubation in a permafrost region of the Qinghai‐Tibet Plateau (QTP). Available N significantly increased due to the stimulation of net nitrification and mineralization in 0–30 cm soil layer. Microbes immobilized N in the end of growing season in both warming and control plots. The magnitude of immobilized N was lower in the warming plots. The root N concentration significantly reduced, but root N pool intensified due to the significant increase in root biomass in the warming treatment. Our results suggest that a warming‐induced increase in biomass is the major N sink and will continue to stimulate plant growth until plant N saturation, which could sustain the positive warming effect on ecosystem productivity.     

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.