Dynamics of labile and recalcitrant soil carbon pools in a sorghum free-air CO2 enrichment (FACE) agroecosystem

Experimentation with dynamics of soil carbon pools as affected by elevated CO₂ can better define the ability of terrestrial ecosystems to sequester global carbon. In the present study, 6 N HCl hydrolysis and stable-carbon isotopic analysis (δ¹³C) were used to investigate labile and recalcitrant soil carbon pools and the translocation among these pools of sorghum residues isotopically labeled in the 1998-1999 Arizona Maricopa free air CO₂ enrichment (FACE) experiment, in which elevated CO₂ (FACE: 560 μmol mol⁻¹) and ambient CO₂ (Control: 360 μmol mol⁻¹) interact with water-adequate (wet) and water-deficient (dry) treatments. We found that on average 53% of the final soil organic carbon (SOC) in the FACE plot was in the recalcitrant carbon pool and 47% in the labile pool, whereas in the Control plot 46% and 54% of carbon were in recalcitrant and labile pools, respectively, indicating that elevated CO₂ transferred more SOC into the slow-decay carbon pool. Also, isotopic mixing models revealed that increased new sorghum residue input to the recalcitrant pool mainly accounts for this change, especially for the upper soil horizon (0-30 cm) where new carbon in recalcitrant soil pools of FACE wet and dry treatments was 1.7 and 2.8 times as large as that in respective Control recalcitrant pools. Similarly, old C in the recalcitrant pool under elevated CO₂ was higher than that under ambient CO₂, indicating that elevated CO₂ reduces the decay of the old C in recalcitrant pool. Mean residence time (MRT) of bulk soil carbon at the depth of 0-30 cm was significantly longer in FACE plot than Control plot by the averages of 12 and 13 yr under the dry and wet conditions, respectively. The MRT was positively correlated to the ratio of carbon content in the recalcitrant pool to total SOC and negatively correlated to the ratio of carbon content in the labile pool to total SOC. Influence of water alone on the bulk SOC or the labile and recalcitrant pools was not significant. However, water stress interacting with CO₂ enhanced the shift of the carbon from labile pool to recalcitrant pool. Our results imply that terrestrial agroecosystems may play a critical role in sequestrating atmospheric CO₂ and mitigating harmful CO₂ under future atmospheric conditions.     

Short and long-term effects of elevated CO2 on Lolium perenne rhizodeposition and its consequences on soil organic matter turnover and plant N yield

It is still unclear whether elevated CO₂ increases plant root exudation and consequently affects the soil microbial biomass. The effects of elevated CO₂ on the fate of the C and nitrogen (N) contained in old soil organic matter pools is also unclear. In this study the short and long-term effects of elevated CO₂ on C and N pools and fluxes were assessed by growing isolated plants of ryegrass (Lolium perenne) in glasshouses at elevated and ambient atmospheric CO₂ and using soil from the New Zealand FACE site that had >4 years exposure to CO₂ enrichment. Using ¹⁴CO₂ pulse labelling, the effects of elevated CO₂ on C allocation within the plant-soil system were studied. Under elevated CO₂ more root derived C was found in the soil and in the microbial biomass 48 h after labelling. The increased availability of substrate significantly stimulated soil microbial growth and acted as priming effect, enhancing native soil organic matter decomposition regardless of the mineral N supply. Despite indications of faster N cycling in soil under elevated CO₂, N availability to plants stayed unchanged. Soil previously exposed to elevated CO₂ exhibited a higher N cycling rate but again there was no effect on plant N uptake. With respect to the difficulties of extrapolating glasshouse experiment results to the field, we concluded that the accumulation of coarse organic matter observed in the field under elevated CO₂ was probably not created by an imbalance between C and N but was likely to be due to more complex phenomena involving soil mesofauna and/or other nutrients limitations.     

Interactive effects of elevated CO2 and nitrogen fertilization levels on photosynthesized carbon allocation in a temperate spring wheat and soil system

Increasing atmospheric CO2 concentration impacts the terrestrial carbon(C) cycle by affecting plant photosynthesis, the flow of photosynthetically fixed C belowground, and soil C pool turnover. For managed agroecosystems, how and to what extent the interactions between elevated CO2 and N fertilization levels influence the accumulation of photosynthesized C in crops and the incorporation of photosynthesized C into arable soil are in urgent need of exploration.We conducted an experiment simulating elevated CO2 with spring wheat(Triticum aestivum L.) planted in growth chambers.13C-enriched CO2 with an identical 13C abundance was continuously supplied at ambient and elevated CO2 concentrations(350 and 600 μmol mol-1, respectively) until wheat harvest.Three levels of N fertilizer application(equivalent to 80, 120, and 180 kg N ha-1 soil) were supplied for wheat growth at both CO2 concentrations. During the continuous 62-d 13CO2 labeling period, elevated CO2 and increased N fertilizer application increased photosynthesized C accumulation in wheat by 14%–24% and 11%–20%, respectively, as indicated by increased biomass production, whereas the C/N ratio in the roots increased under elevated CO2 but declined with increasing N fertilizer application levels. Wheat root deposition induced 1%–2.5% renewal of soil C after 62 d of 13CO2 labeling. Compared to ambient CO2, elevated CO2 increased the amount of photosynthesized C incorporated into soil by 20%–44%. However, higher application rates of N fertilizer reduced the net input of root-derived C in soil by approximately 8% under elevated CO2. For the wheat-soil system, elevated CO2 and increased N fertilizer application levels synergistically increased the amount of photosynthesized C. The pivotal role of plants in photosynthesized C accumulation under elevated CO2 was thereby enhanced in the short term by the increased N application. Therefore, robust N management could mediate C cycling and sequestration by influencing the interactions between plants and soil in agroecosystems under elevated CO2.     

Changes in forest soil organic matter pools after a decade of elevated CO2 and O3

The impact of rising atmospheric carbon dioxide (CO₂) may be mitigated, in part, by enhanced rates of net primary production and greater C storage in plant biomass and soil organic matter (SOM). However, C sequestration in forest soils may be offset by other environmental changes such as increasing tropospheric ozone (O₃) or vary based on species-specific growth responses to elevated CO₂. To understand how projected increases in atmospheric CO₂ and O₃ alter SOM formation, we used physical fractionation to characterize soil C and N at the Rhinelander Free Air CO₂-O₃ Enrichment (FACE) experiment. Tracer amounts of ¹⁵NH₄⁺ were applied to the forest floor of Populus tremuloides, P. tremuloides-Betula papyrifera and P. tremuloides-Acer saccharum communities exposed to factorial CO₂ and O₃ treatments. The ¹⁵N tracer and strongly depleted ¹³C-CO₂ were traced into SOM fractions over four years. Over time, C and N increased in coarse particulate organic matter (cPOM) and decreased in mineral-associated organic matter (MAOM) under elevated CO₂ relative to ambient CO₂. As main effects, neither CO₂ nor O₃ significantly altered ¹⁵N recovery in SOM. Elevated CO₂ significantly increased new C in all SOM fractions, and significantly decreased old C in fine POM (fPOM) and MAOM over the duration of our study. Overall, our observations indicate that elevated CO₂ has altered SOM cycling at this site to favor C and N accumulation in less stable pools, with more rapid turnover. Elevated O₃ had the opposite effect, significantly reducing cPOM N by 15% and significantly increasing the C:N ratio by 7%. Our results demonstrate that CO₂ can enhance SOM turnover, potentially limiting long-term C sequestration in terrestrial ecosystems; plant community composition is an important determinant of the magnitude of this response.     

Natural N abundances in a tallgrass prairie ecosystem exposed to 8-y of elevated atmospheric CO2

After 8-y of elevated CO₂, we previously detected greater amounts of total soil nitrogen, suggesting that rates of ecosystem N flux into or out of tallgrass prairie had been altered. Denitrification and associative N fixation rates are the two primary biological processes that are known to control N loss and accumulation in tallgrass prairie soil. Therefore, our objective was to assess the natural abundance of plant and soil ¹⁵N isotopes as a cumulative index of potential change in efflux or influx of N into and out of the tallgrass prairie after 8-y of exposure to elevated CO₂. Aboveground plant delta ¹⁵N values of Andropogon gerardii were close to zero and more positive as a result of elevated CO₂, but whole-soil values at the 5-30 cm depth were significantly reduced (6.8 vs 7.3; P<0.05) under elevated CO₂-chamber (EC) relative to ambient CO₂- chamber (AC). Total, aboveground plant biomass, root-in-growth, extractable N, microbial biomass N, and soil pools collectively exhibited a range of delta ¹⁵N values from −2.8 to 7.3. Measurements of surface soil ¹⁵N indicate that a change in N inputs and outputs has occurred as a result of elevated atmospheric CO₂. In addition to possible changes in denitrification and N₂ fixation, other sources of N such as the re-translocation of N to the surface from deeper soil layers are needed to explain how soil N accrues in surface soils as a consequence of elevated CO₂. Our results support the notion that C accrual may promote N accrual, possibly driven by high plant and microbial N demand amplified by soil N limitation.     

Elevated CO2, but not defoliation, enhances N cycling and increases short-term soil N immobilization regardless of N addition in a semiarid grassland

Elevated CO₂ and defoliation effects on nitrogen (N) cycling in rangeland soils remain poorly understood. Here we tested whether effects of elevated CO₂ (720 μl L⁻¹) and defoliation (clipping to 2.5 cm height) on N cycling depended on soil N availability (addition of 1 vs. 11 g N m⁻²) in intact mesocosms extracted from a semiarid grassland. Mesocosms were kept inside growth chambers for one growing season, and the experiment was repeated the next year. We added ¹⁵N (1 g m⁻²) to all mesocosms at the start of the growing season. We measured total N and ¹⁵N in plant, soil inorganic, microbial and soil organic pools at different times of the growing season. We combined the plant, soil inorganic, and microbial N pools into one pool (PIM-N pool) to separate biotic + inorganic from abiotic N residing in soil organic matter (SOM). With the ¹⁵N measurements we were then able to calculate transfer rates of N from the active PIM-N pool into SOM (soil N immobilization) and vice versa (soil N mobilization) throughout the growing season. We observed significant interactive effects of elevated CO₂ with N addition and defoliation with N addition on soil N mobilization and immobilization. However, no interactive effects were observed for net transfer rates. Net N transfer from the PIM-N pool into SOM increased under elevated CO₂, but was unaffected by defoliation. Elevated CO₂ and defoliation effects on the net transfer of N into SOM may not depend on soil N availability in semiarid grasslands, but may depend on the balance of root litter production affecting soil N immobilization and root exudation affecting soil N mobilization. We observed no interactive effects of elevated CO₂ with defoliation. We conclude that elevated CO₂, but not defoliation, may limit plant productivity in the long-term through increased soil N immobilization.     

Increased N deposition retards mineralization of old soil organic matter

The aim of this study was to investigate the effects of increased N deposition on new and old pools of soil organic matter (SOM). We made use of a 4-yr experiment, where spruce and beech growing on an acidic loam and a calcareous sand were exposed to increased N deposition (7 vs. 70 kg N ha⁻¹ yr⁻¹) and to elevated atmospheric CO₂. The added CO₂ was depleted in ¹³C, which enabled us to distinguish between old and new C in SOM-pools fractionated into particle sizes. Elevated N deposition for 4 yr increased significantly the contents of total SOM in 0-10 cm depth of the acidic loam (+9%), but not in the calcareous sand. Down to 25 cm soil depth, C storage in the acidic loam was between 100 and 300 g C m⁻² larger under high than under low N additions. However, this increase was small as compared with the SOM losses of 600-700 g C g C 0.25 m⁻¹ m⁻² from the calcareous sand resulting from the disturbance of soils during setting up of the experiment. The amounts of new, less than 4 yr old SOM in the sand fractions of both soils were greater under high N deposition, showing that C inputs from trees into soils increased. Root biomass in the acidic loam was larger under N additions (+25%). Contents of old, more than 4 yr old C in the clay and silt fractions of both soils were significantly greater under high than under low N deposition. Since clay- and silt-bound SOM consists of humified compounds, this indicates that N additions retarded mineralization of old and humified SOM. The retardation of C mineralization in the clay and silt fraction accounted for 60-80 g C m⁻² 4 yr⁻¹, which corresponds to about 40% of the old SOM mineralized in these fraction. As a consequence, preservation of old and humified SOM under elevated N deposition might be a process that could lead to an increased soil C storage in the long-term.     

Effect of elevated CO2 on soil N dynamics in a temperate grassland soil

The response of terrestrial ecosystems to elevated atmospheric CO₂ is related to the availability of other nutrients and in particular to nitrogen (N). Here we present results on soil N transformation dynamics from a N-limited temperate grassland that had been under Free Air CO₂ Enrichment (FACE) for six years. A ¹⁵N labelling laboratory study (i.e. in absence of plant N uptake) was carried out to identify the effect of elevated CO₂ on gross soil N transformations. The simultaneous gross N transformation rates in the soil were analyzed with a ¹⁵N tracing model which considered mineralization of two soil organic matter (SOM) pools, included nitrification from NH₄⁺ and from organic-N to NO₃⁻ and analysed the rate of dissimilatory NO₃⁻ reduction to NH₄⁺ (DNRA). Results indicate that the mineralization of labile organic-N became more important under elevated CO₂. At the same time the gross rate of NH₄⁺ immobilization increased by 20%, while NH₄⁺ oxidation to NO₃⁻ was reduced by 25% under elevated CO₂. The NO₃⁻ dynamics under elevated CO₂ were characterized by a 52% increase in NO₃⁻ immobilization and a 141% increase in the DNRA rate, while NO₃⁻ production via heterotrophic nitrification was reduced to almost zero. The increased turnover of the NH₄⁺ pool, combined with the increased DNRA rate provided an indication that the available N in the grassland soil may gradually shift towards NH₄⁺ under elevated CO₂. The advantage of such a shift is that NH₄⁺ is less prone to N losses, which may increase the N retention and N use efficiency in the grassland ecosystem under elevated CO₂.     

Microbial immobilisation of C rhizodeposits in rhizosphere and root-free soil under continuous C labelling of oats

A greenhouse experiment was conducted by growing oats (Avenasativa L.) in a continuously ¹³CO₂ labeled atmosphere. The allocation of ¹³C-labeled photosynthates in plants, microbial biomass in rhizosphere and root-free soil, pools of soil organic C, and CO₂ emissions were examined over the plant's life cycle. To isolate rhizosphere from root-free soil, plant seedlings were placed into bags made of nylon monofilament screen tissue (16 μm mesh) filled with soil. Two peaks of ¹³C in rhizosphere pools of microbial biomass and dissolved organic carbon (DOC), as well as in CO₂ emissions at the earing and ripeness stages were revealed. These ¹³C maxima corresponded to: (i) the end of rapid root growth and (ii) beginning of root decomposition, respectively. The δ¹³C values of microbial biomass were higher than those of DOC and of soil organic matter (SOM). The microbial biomass C accounted for up to 56 and 39% of ¹³C recovered in the rhizosphere and root-free soil, respectively. Between 4 and 28% of ¹³C assimilated was recovered in the root-free soil. Depending on the phenological stage, the contribution of root-derived C to total CO₂ emission from soil varied from 61 to 92% of total CO₂ evolved, including 4-23% attributed to rhizomicrobial respiration. While 81-91% of C substrates used for microbial growth in the root-free soil and rhizosphere came from SOM, the remaining 9-19% of C substrates utilized by the microbial biomass was attributable to rhizodeposition. The use of continuous isotopic labelling and physical separation of root-free and rhizosphere soil, combined with natural ¹³C abundance were effective in gaining new insight on soil and rhizosphere C-cycling.     

Rice root-derived carbon input and its effect on decomposition of old soil carbon pool under elevated CO2

Rice (Oryza sativa) was grown in sunlit, semi-closed growth chambers (4×3×2 m, L×W×H) at 650 μl l⁻¹ CO₂ (elevated CO₂) to determine: (1) rice root-derived carbon (C) input into the soil under elevated CO₂ in one growing season, and (2) the effect of the newly input C on decomposition of the more recalcitrant native soil organic C. The initial δ¹³C value of the experimental soil was −25.8‰, which was 6‰ less depleted in ¹³C than the plants grown under elevated CO₂. Significant changes in δ¹³C of the soil organic C were detected after one growing season. The amount of new soil C input was estimated to be 0.9 t ha⁻¹ (or 2.1%) at 30 kg N ha⁻¹ and 1.8 t ha⁻¹ (4.1%) at 90 kg N ha⁻¹. Changes in soil δ¹³C suggested that the surface 5 cm of soil received more C input from plants than soils below. Laboratory incubation (25 °C) of soils from different horizons indicated that increased availability of the labile plant-derived C in the soil reduced decomposition of the native soil organic C. Provided the retardant effect of the new C on old soil organic C holds in the field in the longer-term, paddy soils will likely sequester more C from the atmosphere if more plant C enters the soil under elevated atmospheric CO₂.     

Elevation of atmospheric CO2 and N-nutritional status modify nodulation, nodule-carbon supply, and root exudation of Phaseolus vulgaris L.

Increased root exudation and a related stimulation of rhizosphere-microbial growth have been hypothesised as possible explanations for a lower nitrogen- (N-) nutritional status of plants grown under elevated atmospheric CO₂ concentrations, due to enhanced plant-microbial N competition in the rhizosphere. Leguminous plants may be able to counterbalance the enhanced N requirement by increased symbiotic N₂ fixation. Only limited information is available about the factors determining the stimulation of symbiotic N₂ fixation in response to elevated CO₂.In this study, short-term effects of elevated CO₂ on quality and quantity of root exudation, and on carbon supply to the nodules were assessed in Phaseolus vulgaris, grown in soil culture with limited (30 mg N kg⁻¹ soil) and sufficient N supply (200 mg N kg⁻¹ soil), at ambient (400 μmol mol⁻¹) and elevated (800 μmol mol⁻¹) atmospheric CO₂ concentrations.Elevated CO₂ reduced N tissue concentrations in both N treatments, accelerated the expression of N deficiency symptoms in the N-limited variant, but did not affect plant biomass production. ¹⁴CO₂ pulse-chase labelling revealed no indication for a general increase in root exudation with subsequent stimulation of rhizosphere microbial growth, resulting in increased N-competition in the rhizosphere at elevated CO₂. However, a CO₂-induced stimulation in root exudation of sugars and malate as a chemo-attractant for rhizobia was detected in 0.5-1.5 cm apical root zones as potential infection sites. Particularly in nodules, elevated CO₂ increased the accumulation of malate as a major carbon source for the microsymbiont and of malonate with essential functions for nodule development. Nodule number, biomass and the proportion of leghaemoglobin-producing nodules were also enhanced. The release of nod-gene-inducing flavonoids (genistein, daidzein and coumestrol) was stimulated under elevated CO₂, independent of the N supply, and was already detectable at early stages of seedling development at 6 days after sowing.     

Soil organic matter dynamics in grassland soils under elevated CO2: Insights from long-term incubations and stable isotopes

Elevated atmospheric carbon dioxide (CO₂) levels generally stimulate carbon (C) uptake by plants, but the fate of this additional C largely remains unknown. This uncertainty is due in part to the difficulty in detecting small changes in soil carbon pools. We conducted a series of long-term (170-330 days) laboratory incubation experiments to examine changes in soil organic matter pool sizes and turnover rates in soil collected from an open-top chamber (OTC) elevated CO₂ study in Colorado shortgrass steppe. We measured concentration and isotopic composition of respired CO₂ and applied a two-pool exponential decay model to estimate pool sizes and turnover rates of active and slow C pools. The active and slow C pools of surface soils (5-10 cm depth) were increased by elevated CO₂, but turnover rates of these pools were not consistently altered. These findings indicate a potential for C accumulation in near-surface soil C pools under elevated CO₂. Stable isotopes provided evidence that elevated CO₂ did not alter the decomposition rate of new C inputs. Temporal variations in measured δ¹³C of respired CO₂ during incubation probably resulted mainly from the decomposition of changing mixtures of fresh residue and older organic matter. Lignin decomposition may have contributed to declining δ¹³C values late in the experiments. Isotopic dynamics during decomposition should be taken into account when interpreting δ¹³C measurements of soil respiration. Our study provides new understanding of soil C dynamics under elevated CO₂ through the use of stable C isotope measurements during microbial organic matter mineralization.     

C and N in soil organic matter density fractions under elevated atmospheric CO2: Turnover vs. stabilization

Turnover of C and N in an arable soil under Free Air Carbon Dioxide (FACE) experiment was studied by the use of ¹³C natural abundance and ¹⁵N-labeled fertilizers. Wheat was kept four growing seasons under ambient and elevated CO₂ concentrations and fertilized for three growing seasons. Density fractionation of soil organic matter (SOM) allowed to track ¹³C and ¹⁵N in free particulate organic matter (fPOM; <1.6 g cm⁻³), particulate organic matter occluded within aggregates with two densities (oPOM 1.6, oPOM 1.6-2.0 g cm⁻³), and in mineral-associated organic matter (>2.0 g cm⁻³) fractions. Elevated CO₂ and N fertilization did not significantly affect C and N contents in the bulk soil. Calculated mean residence time (MRT) of C and N revealed the qualitative differences of SOM density fractions: (i) the shortest MRT_C and MRT_N in fPOM confirmed high availability of this fraction to decomposition. Larger C/N ratio of fPOM under elevated vs. ambient CO₂ indicated an increasing recalcitrance of FACE-derived plant residues. (ii) There was no difference in MRT of C and N between lighter and heavier oPOMs probably due to short turnover time of soil aggregates which led to oPOM mixing. The increase of MRT_C and MRT_N in both oPOMs during the experiment confirmed the progressive degradation of organic material within aggregates. (iii) Constant turnover rates of C in the mineral fraction neither confirmed nor rejected the assumed stabilization of SOM to take place in the mineral fraction. Moreover, a trend of decreasing of C and N amounts in the Min fraction throughout the experiment was especially pronounced for C under elevated CO₂. Hence, along with the progressive increase of C_FACE in the Min fraction the overall losses of C under elevated CO₂ may occur at the expense of older “pre-FACE” C.     

Temporal and spatial dynamics of soil solution C and N concentrations during Lolium perenne L. sward establishment and the effects of elevated CO2 and N additions

There is now clear evidence for a prolonged increase in atmospheric CO₂ concentrations and enrichment of the biosphere with N. Understanding the fate of C in the plant-soil system under different CO₂ and N regimes is therefore of considerable importance in predicting the environmental effects of climate change and in predicting the sustainability of ecosystems. Swards of Lolium perenne were grown from seed in a Eutric Cambisol at either ambient (ca. 350 μmol mol⁻¹) or elevated (700 μmol mol⁻¹) atmospheric pCO₂ and subjected to two inorganic N fertilizer regimes (no added N and 70 kg N ha⁻¹ month⁻¹). After germination, soil solution concentrations of dissolved organic C (DOC), dissolved inorganic N (DIN), dissolved organic N (DON), phenolics and H⁺ were measured at five depths down the soil profile over 3 months. The exploration of soil layers down the soil profile by roots caused transient increases in soil solution DOC, DON and phenolic concentrations, which then subsequently returned to lower quasi-stable concentrations. In general, the addition of N tended to increase DOC and DON concentrations while exposure to elevated pCO₂ had the opposite effect. These treatment effects, however, gradually diminished over the duration of the experiment from the top of the soil profile downwards. The ambient pCO₂ plus added N regime was the only treatment to maintain a notable difference in soil solution solute concentration, relative to other treatments. This effect on soil solution chemistry appeared to be largely indirect resulting from increased plant growth and a decrease in soil moisture content. Our results show that although plant growth responses to elevated pCO₂ are critically dependent upon N availability, the organic chemistry of the soil solution is relatively insensitive to changes in plant growth once the plants have become established.     

Release of C and N from roots of peas and oats and their availability to soil microorganisms

Nutrient mobilisation in the rhizosphere is driven by soil microorganisms and controlled by the release of available C compounds from roots. It is not known how the quality of release influences this process in situ. Therefore, the present study was conducted to investigate the amount and turnover of rhizodeposition, in this study defined as root-derived C or N present in the soil after removal of roots and root fragments, released at different growth stages of peas (Pisum sativum L.) and oats (Avena sativa L.). Plants were grown in soil columns placed in a raised bed under outdoor conditions and simultaneously pulse labelled in situ with a ¹³C-glucose-¹⁵N-urea solution using a stem feeding method. After harvest, ¹³C and ¹⁵N was recovered in plant parts and soil pools, including the microbial biomass. Net rhizodeposition of C and N as a percentage of total plant C and N was higher in peas than in oats. Moreover, the C-to-N ratio of the rhizodeposits was lower in peas, and a higher proportion of the microbial biomass and inorganic N was derived from rhizodeposition. These results suggest a positive plant-soil feedback shaping nutrient mobilisation. This process is driven by the C and N supply of roots, which has a higher availability in peas than in oats.     

Nitrogen rhizodeposition of young wheat plants under elevated CO<Subscript>2</Subscript> and drought stress

The objective of this study was to determine the effect of drought stress and elevated CO₂ concentrations around the shoots on N rhizodeposition of young wheat plants. In a pot experiment, the plant N pool was labeled through ¹⁵NH₃ application to shoots at nontoxic NH₃ concentrations, and the impact of low water supply (40% field capacity), elevated CO₂ (720 μmol mol⁻¹ CO₂), and the combination of both factors on the ¹⁵N distribution was studied. Total ¹⁵N rhizodeposition ranged from 5 to 11% of the total ¹⁵N recovered in the plant/soil system. Elevated CO₂ concentration as well as drought stress increased the belowground transport of N and increased the relative portion of N rhizodeposition on total ¹⁵N in the plant/soil system. However, while the increased N rhizodeposition with elevated CO₂ was the result of increased total belowground N transport, drought stress additionally increased the portion of ¹⁵N found in rhizodeposition vs roots. Elevated CO₂ intensified the effect of drought stress. The percentage of water soluble ¹⁵N in the ¹⁵N rhizodeposition was very low under all treatments, and it was significantly decreased by the drought-stressed treatments.     

CO2 and inorganic N supply modify competition for N between co-occurring grass plants, tree seedlings and soil microorganisms

Plant-plant and plant-soil interactions play a key role in determining plant community structure and ecosystem function. However, the effects of global change on the interplay between co-occurring plants and soil microbes in successional communities are poorly understood. In this study, we investigated competition for nitrogen (N) between soil microorganisms, grass plants and establishing tree seedlings under factorial carbon dioxide (CO₂) and N treatments. Fraxinus excelsior seedlings were germinated in the presence or absence of grass competition (Dactylis glomerata) at low (380 μmol mol⁻¹) or high (645 μmol mol⁻¹) CO₂ and at two levels of N nutrition in a mesocosm experiment. Pulse ¹⁵N labelling was used to examine N partitioning among plant and soil compartments. Dactylis exerted a strong negative effect on Fraxinus biomass, N capture and ¹⁵N recovery irrespective of N and CO₂ treatment. In contrast, the presence of Dactylis had a positive effect on the microbial N pool. Plant and soil responses to N treatment were of a greater magnitude compared with responses to elevated CO₂, but the pattern of Fraxinus- and microbial-N pool response to N and CO₂ varied depending on grass competition treatment. Within the Dactylis competition treatment, decreases in Fraxinus biomass in response to N were not mirrored by decreases in tree seedling N content, suggesting a shift from below- to above-ground competition. In the Dactylis-sown pots, ¹⁵N recovery could be ranked Dactylis > microbial pool > Fraxinus in all N and CO₂ treatment combinations. Inequalities between Fraxinus and soil microorganisms in terms of ¹⁵N recovery were exacerbated by N addition. Contrary to expectations, elevated CO₂ did not increase plant-microbe competition. Nevertheless, microbial ¹⁵N recovery showed a small positive increase in the high CO₂ treatment. Overall, elevated CO₂ and N supply did not interact on plant/soil N partitioning. Our data suggest that the competitive balance between establishing tree seedlings and grass plants in an undisturbed sward is relatively insensitive to CO₂ or N-induced modifications in N competition between plant and soil compartments.     

Changes in the fungal-to-bacterial respiratory ratio and microbial biomass in agriculturally managed soils under free-air CO2 enrichment (FACE) - A six-year survey of a field study

In soil ecology, microbial parameters have been identified as sensitive indicators of changes in the soil environment. The Braunschweig FACE project provided the opportunity to study the effects of elevated CO₂ (550 μmol mol⁻¹) as compared to ambient CO₂ (370 μmol mol⁻¹) on total microbial biomass (C_mic), C_mic-to-C_org ratio and the fungal-to-bacterial respiratory ratio together with total C_org, N_t, C:N ratio and pH over a six-year period. Field management followed a typical crop rotation system of this region with either a crop-related full nitrogen supply (N100) or 50% reduced N supply (N50). The soil microbial parameters responded to the elevated CO₂ treatment in varying intensities and time spans. The fungal-to-bacterial respiratory ratio was the most sensitive parameter in responding to an elevated CO₂ treatment with highly significant differences to ambient CO₂-treated control plots in the third year of CO₂ fumigation. After six years bacterial respiratory activity had increased in ascending order to 34% in FACE-treated plots (N50 and N100) as compared to control plots. Soil microbial biomass (C_mic) responded more slowly to the FACE treatment with highly significant increases of >12% after the fourth year of CO₂ fumigation. The C_mic-to-C_org ratio responded very late in the last two years of the CO₂ treatment with a significant increase of >7.0% only in the N100 variant. Total C_org and N_t were slightly but significantly increased under FACE around 10.0% with ascending tendency over time starting with the second year of CO₂ treatment. No significant FACE effects could be recorded for the C:N ratio or pH.These results suggest that under FACE treatment changes in the soil microbial community will occur. In our study the fungal-to-bacterial respiratory ratio was superior to total C_mic as microbial bioindicators in reflecting changes in the soil organic matter composition.     

Fate of C- and N-labelled rhizodeposition of Lolium perenne as function of the distance to the root surface

A greenhouse rhizobox experiment was carried out to quantify the incorporation of ¹³C- and ¹⁵N-labelled rhizodeposits into different soil pools, especially into the rhizosphere microbial biomass, with increasing distances to the root surface of Lolium perenne. Five layers were analysed over 0-4.2 mm distance to an artificial root surface. C and N derived from rhizodeposition were 4.2% of total C and 2.8% of total N in soil at 0-1.0 mm distance and decreased rapidly with increasing distance. Microbial biomass C and N increased significantly towards the roots. At 0-1.0 mm distance microbial biomass C and N accounted for 66% and 29% of C and N derived from rhizodeposition, respectively. These percentages declined with increasing distance to the roots, but were still traceable up to 4.2 mm distance. Only small amounts of root released C and N were found in the 0.05 M K₂SO₄-extractable fraction. Extractable C and N derived from rhizodeposition varied around means of 4% of total C and N derived from rhizodeposition and increased only marginally with increasing distance to the roots. C derived from rhizodeposition in the non-extractable soil organic matter increased from 65 to 89% of total C derived from rhizodeposition at 0-3.4 mm distance. Conversely, microbial biomass C derived from rhizodeposition decreased from 33 to 4%. N derived from rhizodeposition in the non-extractable soil organic matter increased from 61 to 79% of total N derived from rhizodeposition at 0-2.6 mm distance, followed by a decline to roughly 55% in the two outer layers. Microbial biomass N decreased from 37 to 16% at 0-2.6 mm distance, followed by an increase to roughly 41% in the two outer layers. The C/N ratio of total C and N derived from rhizodeposition as well as that of extractable C and N derived from rhizodeposition increased with increasing distance to the roots to values above 30. In contrast, the C/N ratio of incorporated rhizodeposition C and N into the microbial biomass decreased to values less than 5 at 2.6-4.2 mm distance. The data indicate differential microbial response to C and N derived from rhizodeposition at a high spatial resolution from the root surface. The turnover of C and N derived from rhizodeposition in the rhizosphere as a function of the distance to the root surface is discussed.     

Rice rhizodeposition and its utilization by microbial groups depends on N fertilization

Rhizodeposits have received considerable attention, as they play an important role in the regulation of soil carbon (C) sequestration and global C cycling and represent an important C and energy source for soil microorganisms. However, the utilization of rhizodeposits by microbial groups, their role in the turnover of soil organic matter (SOM) pools in rice paddies, and the effects of nitrogen (N) fertilization on rhizodeposition are nearly unknown. Rice (Oryza sativa L.) plants were grown in soil at five N fertilization rates (0, 10, 20, 40, or 60 mg N kg^?1 soil) and continuously labeled in a ¹³CO₂ atmosphere for 18 days during tillering. The utilization of root-derived C by microbial groups was assessed by ¹³C incorporation into phospholipid fatty acids. Rice shoot and root biomass strongly increased with N fertilization. Rhizodeposition increased with N fertilization, whereas the total ¹³C incorporation into microorganisms, as indicated by the percentage of ¹³C recovered in microbial biomass, decreased. The contribution of root-derived ¹³C to SOM formation increased with root biomass. The ratio of ¹³C in soil pools (SOM and microbial biomass) to ¹³C in roots decreased with N fertilization showing less incorporation and faster turnover with N. The ¹³C incorporation into fungi (18:2ω6,9c and 18:1ω9c), arbuscular mycorrhizal fungi (16:1ω5c), and actinomycetes (10Me 16:0 and 10Me 18:0) increased with N fertilization, whereas the ¹³C incorporation into gram-positive (i14:0, i15:0, a15:0, i16:0, i17:0, and a17:0) and gram-negative (16:1ω7c, 18:1ω7c, cy17:0, and cy19:0) bacteria decreased with N fertilization. Thus, the uptake and microbial processing of root-derived C was affected by N availability in soil. Compared with the unfertilized soil, the contribution of rhizodeposits to SOM and microorganisms increased at low to intermediate N fertilization rates but decreased at the maximum N input. We conclude that belowground C allocation and rhizodeposition by rice, microbial utilization of rhizodeposited C, and its stabilization within SOM pools are strongly affected by N availability: N fertilization adequate to the plant demand increases C incorporation in all these polls, but excessive N fertilization has negative effects not only on environmental pollution but also on C sequestration in soil.