Rhizodeposition-induced decomposition increases N availability to wild and cultivated wheat genotypes under elevated CO2

Elevated CO₂ may increase nutrient availability in the rhizosphere by stimulating N release from recalcitrant soil organic matter (SOM) pools through enhanced rhizodeposition. We aimed to elucidate how CO₂-induced increases in rhizodeposition affect N release from recalcitrant SOM, and how wild versus cultivated genotypes of wheat mediated differential responses in soil N cycling under elevated CO₂. To quantify root-derived soil carbon (C) input and release of N from stable SOM pools, plants were grown for 1 month in microcosms, exposed to ¹³C labeling at ambient (392 μmol mol⁻¹) and elevated (792 μmol mol⁻¹) CO₂ concentrations, in soil containing ¹⁵N predominantly incorporated into recalcitrant SOM pools. Decomposition of stable soil C increased by 43%, root-derived soil C increased by 59%, and microbial-¹³C was enhanced by 50% under elevated compared to ambient CO₂. Concurrently, plant ¹⁵N uptake increased (+7%) under elevated CO₂ while ¹⁵N contents in the microbial biomass and mineral N pool decreased. Wild genotypes allocated more C to their roots, while cultivated genotypes allocated more C to their shoots under ambient and elevated CO₂. This led to increased stable C decomposition, but not to increased N acquisition for the wild genotypes. Data suggest that increased rhizodeposition under elevated CO₂ can stimulate mineralization of N from recalcitrant SOM pools and that contrasting C allocation patterns cannot fully explain plant mediated differential responses in soil N cycling to elevated CO₂.     

Effects of elevated CO2 concentration on rhizodeposition from Lolium perenne grown on soil exposed to 9 years of CO2 enrichment

The effects of enriched CO₂ atmosphere on partitioning of recently assimilated carbon were investigated in a plant-soil-microorganism system in which Lolium perenne seedlings were planted into cores inserted into the resident soil within a sward that had been treated with elevated CO₂ for 9 consecutive years, under two N fertilisation levels (Swiss FACE experiment). The planted cores were excavated from the ambient (35 Pa pCO₂) and enriched (60 Pa pCO₂) rings at two dates, in spring and autumn, during the growing season. The cores were brought back to the laboratory for ¹⁴C labelling of shoots in order to trace the transfer of recently assimilated C both within the plant and to the soil and microbial biomass. At the spring sampling, high N supply stimulated shoot and total dry matter production. Consistently, high N enhanced the allocation of recently fixed C to shoots, and reduced it to belowground compartments. Elevated CO₂ had no consequences for DM or the pattern of C allocation. At the autumn sampling, at high N plot, yield of L. perenne was stimulated by elevated CO₂. Consistently, ¹⁴C was preferentially allocated aboveground and, consequently belowground recent C allocation was depressed and rhizodeposition reduced. At both experimental periods, total soil C content was similar in all treatments, providing no evidence for soil carbon sequestration in the Swiss Free Air CO₂ Enrichment experiment (FACE) after 9 years of enrichment. Recently assimilated C and soil C were mineralised faster in soils from enriched rings, suggesting a CO₂-induced shift in the microbial biomass characteristics (structure, diversity, activity) and/or in the quality of the root-released organic compounds.     

Carbon allocation in grassland communities under drought stress followed by 14C pulse labeling

Although extreme climatic events such as drought have important consequences for belowground carbon (C) cycling, their impact on the plant-soil system of mixed plant communities is poorly understood. Our objective was to study the effect of drought on C allocation and rhizosphere-mediated CO₂ fluxes under three plant species: Lolium perenne, Festuca arundinacea and Medicago sativa grown in monocultures or mixture. The conceptual approach included ¹⁴CO₂ pulse labeling of plants grown under drought and optimum water conditions in order to be able to follow above- and belowground C allocation. After ¹⁴C pulse labeling, we traced ¹⁴C allocation to shoots and roots, soil and rhizospheric CO₂, dissolved organic carbon (DOC) and microbial biomass.Drought and plant community composition significantly affected assimilate allocation in the plant-soil system. Drought conditions changed the source sink relationship of monocultures, which transferred a relatively larger portion of assimilates to their roots compared to water sufficient plants. In contrast, plant mixture showed an increase in ¹⁴C allocation to shoots when exposed to drought.Under drought stress, root respiration was reduced for all monocultures except under the legume species. Microbial respiration remained similar in all cases showing that microbial activity was less affected by drought than root activity. This may be explained by strongly increased assimilate allocation to easily available exudates or rhizodeposits under drought. In conclusion, plant community composition may modify the impact of climatic changes on carbon allocation and belowground carbon fluxes. The presence of legume species attenuates drought effects on rhizosphere processes.     

Elevated CO2 differentially alters belowground plant and soil microbial community structure in reed canary grass-invaded experimental wetlands

Several recent studies have indicated that an enriched atmosphere of carbon dioxide (CO₂) could exacerbate the intensity of plant invasions within natural ecosystems, but little is known of how rising CO₂ impacts the belowground characteristics of these invaded systems. In this study, we examined the effects of elevated CO₂ and nitrogen (N) inputs on plant and soil microbial community characteristics of plant communities invaded by reed canary grass, Phalaris arundinacea L. We grew the invasive grass under two levels of invasion: the invader was either dominant (high invasion) at >90% plant cover or sub-dominant (low invasion) at <50% plant cover. Experimental wetland communities were grown for four months in greenhouses that received either 600 or 365 μl l⁻¹ (ambient) CO₂. Within each of three replicate rooms per CO₂ treatment, the plant communities were grown under high (30 mg l⁻¹) or low (5 mg l⁻¹) N. In contrast to what is often predicted under N limitation, we found that elevated CO₂ increased native graminoid biomass at low N, but not at high N. The aboveground biomass of reed canary grass did not respond to elevated CO₂, despite it being a fast-growing C3 species. Although elevated CO₂ had no impact on the plant biomass of heavily invaded communities, the relative abundance of several soil microbial indicators increased. In contrast, the moderately invaded plant communities displayed increased total root biomass under elevated CO_2, while little impact occurred on the relative abundance of soil microbial indicators. Principal components analysis indicated that overall soil microbial community structure was distinct by CO₂ level for the varying N and invasion treatments. This study demonstrates that even when elevated CO₂ does not have visible effects on aboveground plant biomass, it can have large impacts belowground.     

二氧化碳升高与干旱对植物生理、土壤碳及土壤酶活的影响

Global climate models have indicated high probability of drought occurrences in the coming future decades due to the impacts of climate change caused by a mass release of CO2.Thus,climate change regarding elevated ambient CO2 and drought may consequently affect the growth of crops.In this study,plant physiology,soil carbon,and soil enzyme activities were measured to investigate the impacts of elevated CO2 and drought stress on a Stagnic Anthrosol planted with soybean (Glycine max).Treatments of two CO2 levels,three soil moisture levels,and two soil cover types were established.The results indicated that elevated CO2 and drought stress significantly affected plant physiology.The inhibition of plant physiology by drought stress was mediated via prompted photosynthesis and water use efficiency under elevated CO2 conditions.Elevated CO2 resulted in a longer retention time of dissolved organic carbon (DOC) in soil,probably by improving the soil water effectiveness for organic decomposition and mineralization.Drought stress significantly decreased C:N ratio and microbial biomass carbon (MBC),but the interactive effects of drought stress and CO2 on them were not significant.Elevated CO2 induced an increase in invertase and catalase activities through stimulated plant root exudation.These results suggested that drought stress had significant negative impacts on plant physiology,soil carbon,and soil enzyme activities,whereas elevated CO2 and plant physiological feedbacks indirectly ameliorated these impacts.     

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.     

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.     

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.     

Does labelling frequency affect N rhizodeposition assessment using the cotton-wick method?

The aim of the present study was to test and improve the reliability of the ¹⁵N cotton-wick method for measuring soil N derived from plant rhizodeposition, a critical value for assessing belowground nitrogen input in field-grown legumes. The effects of the concentration of the ¹⁵N labelling solution and the feeding frequency on assessment of nitrogen rhizodeposition were studied in two greenhouse experiments using the field pea (Pisum sativum L.). Neither the method nor the feeding frequency altered plant biomass and N partitioning, and the method appeared well adapted for assessing the belowground contribution of field-grown legumes to the soil N pool. However, nitrogen rhizodeposition assessment was strongly influenced by the feeding frequency and the concentration of labelling solution. At pod-filling and maturity, despite similar root ¹⁵N enrichment, the fraction of plants' belowground nitrogen allocated to rhizodeposition in both Frisson pea and the non-nodulating isoline P2 was 20 to more than 50% higher when plants were labelled continuously than when they were labelled using fortnightly pulses. Our results suggest that when ¹⁵N root enrichment was high, nitrogen rhizodeposition was overestimated only for plants that were ¹⁵N-fed by fortnightly pulses, and not in plants ¹⁵N-fed continuously. This phenomenon was especially observed for plants that rely on symbiotic N₂ fixation for N acquisition, and it may be linked to the concentration of the labelling solution. In conclusion, the assessment of nitrogen rhizodeposition was more reliable when plants were labelled continuously with a dilute solution of ¹⁵N urea.     

Effect of CO<Subscript>2</Subscript> on nitrification and immobilization of NH<Subscript>4</Subscript><Superscript>+</Superscript>-N

A laboratory incubation experiment was conducted to demonstrate that reduced availability of CO₂ may be an important factor limiting nitrification. Soil samples amended with wheat straw (0%, 0.1% and 0.2%) and (¹⁵NH₄)₂SO₄ (200 mg N kg^–1 soil, 2.213 atom% ¹⁵N excess) were incubated at 30±2°C for 20 days with or without the arrangement for trapping CO₂ resulting from the decomposition of organic matter. Nitrification (as determined by the disappearance of NH₄⁺ and accumulation of NO₃^–) was found to be highly sensitive to available CO₂ decreasing significantly when CO₂ was trapped in alkali solution and increasing substantially when the amount of CO₂ in the soil atmosphere increased due to the decomposition of added wheat straw. The co-efficient of correlation between NH₄⁺-N and NO₃^–-N content of soil was highly significant (r =0.99). During incubation, 0.1–78% of the applied NH₄⁺ was recovered as NO₃^– at different incubation intervals. Amendment of soil with wheat straw significantly increased NH₄⁺ immobilization. From 1.6% to 4.5% of the applied N was unaccounted for and was due to N losses. The results of the study suggest that decreased availability of CO₂ will limit the process of nitrification during soil incubations involving trapping of CO₂ (in closed vessels) or its removal from the stream of air passing over the incubated soil (in open-ended systems).     

Effects of elevated CO2 and O3 on soil respiration under ponderosa pine

Soil respiration represents the integrated response of plant roots and soil organisms to environmental conditions and the availability of C in the soil. A multi-year study was conducted in outdoor sun-lit controlled-environment chambers containing a reconstructed ponderosa pine/soil-litter system. The study used a 2×2 factorial design with two levels of CO₂ and two levels of O₃ and three replicates of each treatment. The objectives of our study were to assess the effects of long-term exposure to elevated CO₂ and O₃, singly and in combination, on soil respiration, fine root growth and soil organisms. Fine root growth and soil organisms were included in the study as indicators of the autotrophic and heterotrophic components of soil respiration. The study evaluated three hypotheses: (1) elevated CO₂ will increase C assimilation and allocation belowground increasing soil respiration; (2) elevated O₃ will decrease C assimilation and allocation belowground decreasing soil respiration and (3) as elevated CO₂ and O₃ have opposing effects on C assimilation and allocation, elevated CO₂ will eliminate or reduce the negative effects of elevated O₃ on soil respiration. A mixed-model covariance analysis was used to remove the influences of soil temperature, soil moisture and days from planting when testing for the effects of CO₂ and O₃ on soil respiration. The covariance analysis showed that elevated CO₂ significantly reduced the soil respiration while elevated O₃ had no significant effect. Despite the lack of a direct CO₂ stimulation of soil respiration, there were significant interactions between CO₂ and soil temperature, soil moisture and days from planting indicating that elevated CO₂ altered soil respiration indirectly. In elevated CO₂, soil respiration was more sensitive to soil temperature changes and less sensitive to soil moisture changes than in ambient CO₂. Soil respiration increased more with days from planting in elevated than in ambient CO₂. Elevated CO₂ had no effect on fine root biomass but increased abundance of culturable bacteria and fungi suggesting that these increases were associated with increased C allocation belowground. Elevated CO₂ had no significant effect on microarthropod and nematode abundance. Elevated O₃ had no significant effects on any parameter except it reduced the sensitivity of soil respiration to changes in temperature.     

Denitrification in grass swards is increased under elevated atmospheric CO2

Emissions of N₂O and N₂ were measured from Lolium perenne L. swards under ambient (36 Pa) and elevated (60 Pa) atmospheric CO₂ at the Swiss free air carbon dioxide enrichment experiment following application of 11.2 g N m⁻² as ¹⁵NH₄¹⁵NO₃ or ¹⁴NH₄¹⁵NO₃ (1 at.% excess ¹⁵N). Total denitrification (N₂O+N₂) was increased under elevated pCO₂ with emissions of 6.2 and 19.5 mg ¹⁵N m⁻² measured over 22 d from ambient and elevated pCO₂ swards, respectively, supporting the hypothesis that increased belowground C allocation under elevated pCO₂ provides the energy for denitrification. Nitrification was the predominant N₂O producing process under ambient pCO₂ whereas denitrification was predominant under elevated pCO₂. The N₂-to-N₂O ratio was often higher under elevated pCO₂ suggesting that previous estimates of gaseous N losses based only on N₂O emissions have greatly underestimated the loss of N by denitrification.     

Gross fluxes of nitrogen in grassland soil exposed to elevated atmospheric pCO2 for seven years

Plant response to increasing atmospheric CO₂ partial pressure (pCO₂) depends on several factors, one of which is mineral nitrogen availability facilitated by the mineralisation of organic N. Gross rates of N mineralisation were examined in grassland soils exposed to ambient (36 Pa) and elevated (60 Pa) atmospheric pCO₂ for 7 years in the Swiss Free Air Carbon dioxide Enrichment experiment. It was hypothesized that increased below-ground translocation of photoassimilates at elevated pCO₂ would lead to an increase in immobilisation of N due to an excess supply of energy to the roots and rhizosphere. Intact soil cores were sampled from Lolium perenne and Trifolium repens swards in May and September, 2000. The rates of gross N mineralisation (m) and NH₄⁺ consumption (c) were determined using ¹⁵N isotopic dilution during a 51-h period of incubation. The rates of N immobilisation were estimated either as the difference between m and the net N mineralisation rate or as the amount of ¹⁵N released from the microbial biomass after chloroform fumigation. Soil samples from both swards showed that the rates of gross N mineralisation and NH₄⁺ consumption did not change significantly under elevated pCO₂. The lack of a significant effect of elevated pCO₂ on organic N turnover was consistent with the similar size of the microbial biomass and similar immobilisation of applied ¹⁵N in the microbial N pool under ambient and elevated pCO₂. Rates of m and c, and microbial ¹⁵N did not differ significantly between the two sward types although a weak (p<0.1) pCO₂ by sward interaction occurred. A significantly larger amount of NO₃⁻ was recovered at the end of the incubation in soil taken from T. repens swards compared to that from L. perenne swards. Eleven percent of the added ¹⁵N were recovered in the roots in the cores sampled under L. perenne, while only 5% were recovered in roots of T. repens. These results demonstrate that roots remained a considerable sink despite the shoots being cut at ground level prior to incubation and suggest that the calculation of N immobilisation from gross and net rates of mineralisation in soils with a high root biomass does not reflect the actual immobilisation of N in the microbial biomass. The results of this study did not support the initial hypothesis and indicate that below-ground turnover of N, as well as N availability, measured in short-term experiments are not strongly affected by long-term exposure to elevated pCO₂. It is suggested that differences in plant N demand, rather than major changes in soil N mineralisation/immobilisation, are the long-term driving factors for N dynamics in these grassland systems.     

Direct and indirect effects of elevated CO2 and nitrogen addition on soil microbial communities in the rhizosphere of Bothriochloa ischaemum

Purpose

Elevated CO₂ and nitrogen (N) addition both affect soil microbial communities, which significantly influence soil processes and plant growth. Here, we evaluated the combined effects of elevated CO₂ and N addition on the soil–microbe–plant system of the Chinese Loess Plateau.

Materials and methods

A pot cultivation experiment with two CO₂ treatment levels (400 and 800 μmol mol^?1) and three N addition levels (0, 2.5, and 5 g N m^?2 year^?1) was conducted in climate-controlled chambers to evaluate the effects of elevated CO₂ and N addition on microbial community structure in the rhizosphere of Bothriochloa ischaemum using phospholipid fatty acid (PLFA) profiles and associated soil and plant properties. Structural equation modeling (SEM) was used to identify the direct and indirect effects of the experimental treatments on the structure of microbial communities.

Results and discussion

Elevated CO₂ and N addition both increased total and fungal PLFAs. N addition alone increased bacterial, Gram-positive, and Gram-negative PLFAs. However, elevated CO₂ interacting with N addition had no significant effects on the microbial community. The SEM indicated that N addition directly affected the soil microbial community structure. Elevated CO₂ and N addition both indirectly affected the microbial communities by affecting plant and soil variables. N addition exerted a stronger total effect than elevated CO₂.

Conclusions

The results highlighted the importance of comprehensively studying soil–microbe–plant systems to deeply reveal how characteristics of terrestrial ecosystems may respond under global change.

     

Rhizodeposition: Its contribution to microbial growth and carbon and nitrogen turnover within the rhizosphere

A greenhouse rhizobox experiment was carried out to investigate the fate and turnover of ¹³C‐ and ¹⁵N‐labeled rhizodeposits within a rhizosphere gradient from 0–8 mm distance to the roots of wheat. Rhizosphere soil layers from 0–1, 1–2, 2–3, 3–4, 4–6, and 6–8 mm distance to separated roots were investigated in an incubation experiment (42 d, 15°C) for changes in total C and N and that derived from rhizodeposition in total soil, in soil microbial biomass, and in the 0.05 M K₂SO₄–extractable soil fraction. CO₂‐C respiration in total and that derived from rhizodeposition were measured from the incubated rhizosphere soil samples. Rhizodeposition C was detected in rhizosphere soil up to 4–6 mm distance from the separated roots. Rhizodeposition N was only detected in the rhizosphere soils up to 3–4 mm distance from the roots. Microbial biomass C and N was increased with increasing proximity to the separated roots. Beside ¹³C and ¹⁵N derived from rhizodeposits, unlabeled soil C and N (native SOM) were incorporated into the growing microbial biomass towards the roots, indicating a distinct acceleration of soil organic matter (SOM) decomposition and N immobilization into the growing microbial biomass, even under the competition of plant growth. During the soil incubation, microbial biomass C and N decreased in all samples. Any decrease in microbial biomass C and N in the incubated rhizosphere soil layers is attributed mainly to a decrease of unlabeled (native) C and N, whereas the main portion of previously incorporated rhizodeposition C and N during the plant growth period remained immobilized in the microbial biomass during the incubation. Mineralization of native SOM C and N was enhanced within the entire investigated rhizosphere gradient. The results indicate complex interactions between substrate input derived from rhizodeposition, microbial growth, and accelerated C and N turnover, including the decomposition of native SOM (i.e., rhizosphere priming effects) at a high spatial resolution from the roots.     

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.     

Potassium deficiency impedes elevated carbon dioxide‐induced biomass enhancement in well watered or drought‐stressed bread wheat

Potassium (K) deficiency reduces photosynthesis and biomass production of crop plants and also renders them vulnerable to drought stress, whereas elevated carbon dioxide (CO₂) has a positive effect on photosynthesis and yield and ameliorates the adverse effects of drought stress. This study aimed to characterize the physiological responses of wheat (Triticum aestivum L.) stressed with K deficiency under elevated CO₂ and drought conditions. Increased biomass production caused by elevated CO₂ as a consequence of increased photosynthesis and water use efficiency was absent in young K‐deficient wheat plants. Shoot K concentration was negatively affected by elevated CO₂ particularly under K‐deficient conditions, whereas K content per plant was greatest in plants supplied with adequate K and adequate water. Specific leaf weight was increased as a consequence of carbohydrate accumulation in the source leaves of K‐deficient plants particularly under elevated CO₂ and drought stress. Potassium deficiency clearly impeded the impact of elevated CO₂ in both well watered as well as drought‐stressed plants. Adequate K fertilization is a prerequisite for efficient harvesting of atmospheric CO₂ through increased photosynthesis, decreased transpiration, and increased biomass production under changing atmospheric CO₂ and soil moisture conditions.     

Atmospheric CO2 enrichment and nutrient additions to planted soil increase mineralisation of soil organic matter, but do not alter microbial utilisation of plant- and soil C-sources

Plants link atmospheric and soil carbon pools through CO₂ fixation, carbon translocation, respiration and rhizodeposition. Within soil, microbial communities both mediate carbon-sequestration and return to the atmosphere through respiration. The balance of microbial use of plant-derived and soil organic matter (SOM) carbon sources and the influence of plant-derived inputs on microbial activity are key determinants of soil carbon-balance, but are difficult to quantify. In this study we applied continuous ¹³C-labelling to soil-grown Lolium perenne, imposing atmospheric CO₂ concentrations and nutrient additions as experimental treatments. The relative use of plant- and SOM-carbon by microbial communities was quantified by compound-specific ¹³C-analysis of phospholipid fatty acids (PLFAs). An isotopic mass-balance approach was applied to partition the substrate sources to soil respiration (i.e. plant- and SOM-derived), allowing direct quantification of SOM-mineralisation. Increased CO₂ concentration and nutrient amendment each increased plant growth and rhizodeposition, but did not greatly alter microbial substrate use in soil. However, the increased root growth and rhizosphere volume with elevated CO₂ and nutrient amendment resulted in increased rates of SOM-mineralisation per experimental unit. As rhizosphere microbial communities utilise both plant- and SOM C-sources, the results demonstrate that plant-induced priming of SOM-mineralisation can be driven by factors increasing plant growth. That the balance of microbial C-use was not affected on a specific basis may suggest that the treatments did not affect soil C-balance in this study.     

Effects of nitrogen fertilization on pot‐grown wheat photosynthate partitioning within intensively farmed soil determined by 13C pulse‐labeling

Understanding rhizodeposited carbon (C) dynamics of winter wheat (Triticum aestivum L.) is important for improving soil fertility and increasing soil C stocks. However, the effects of nitrogen (N) fertilization on photosynthate C allocation to rhizodeposition of wheat grown in an intensively farmed alkaline soil remain elusive. In this study, pot‐grown winter wheat under N fertilization of 250 kg N ha^?1 was pulse‐labeled with ¹³CO₂ at tillering, elongation, anthesis, and grain‐filling stages. The ¹³C in shoots, roots, soil organic carbon (SOC), and rhizosphere‐respired CO₂ was measured 28 d after each ¹³C labeling. The proportion of net‐photosynthesized ¹³C recovered (shoots + roots + soil + soil respired CO₂) in the shoots increased from 58–64% at the tillering to 86–91% at the grain‐filling stage. Likewise, the proportion in the roots decreased from 21–28% to 2–3%, and that in the SOC pool increased from 1–2% to 6–7%. However, the ¹³C respired CO₂ allocated to soil peaked (17–18%) at the elongation stage and decreased to 6–8% at the grain‐filling stage. Over the entire growth season of wheat, N fertilization decreased the proportion of net photosynthate C translocated to the below‐ground pool by about 20%, but increased the total amount of fixed photosynthate C, and therefore increased the below‐ground photosynthate C input. We found that the chase period of about 4 weeks is sufficient to accurately monitor the recovery of ¹³C after pulse labeling in a wheat–soil system. We conclude that N fertilization increased the deposition of photoassimilate C into SOC pools over the entire growth season of wheat compared to the control treatment.     

Release of carbon and nitrogen compounds by plant roots and their possible ecological importance+

The root-borne C- and N-flux in the plant/soil system was studied by determining the ¹⁴C- or ¹⁵N-balances in pot trials with soil as a substrate (¹⁴CO₂- or ¹⁵NH₃-application to the shoots, comparison of sterile and nonsterile treatments for quantification of root-borne substances). The following results were obtained: 1. The amount of (primary) root-borne carbon compounds released into soil was (besides root respiration) 11—20% of net-CO₂-assimilation or 13—32% of the ¹⁴C incorporated into the plants (= 1 t C · ha^—1). 5—6% of ¹⁵N assimilated by the plants were released as root-borne N compounds (= 15 kg N · ha^—1). 2. A considerable portion of the root-borne C (about 6% = 600 kg C · ha^—1) was found in the rooted soil zone at the end of the experiments (rhizodeposition). 3. (Primary) root-borne C and N compounds found in immediate vicinity of the roots (about 60—80%) were mainly water soluble, whereas most of the C and N compounds found in a greater distance were water insoluble. The water soluble exudates consisted mainly of neutral (carbohydrates) and acid fractions (organic acids). The basic fraction (amino acids) made up a small portion only. 4. The root-borne C and N compounds influenced the nutrient balance of soil and plant directly and/or indirectly via microbes (depending on species, variety and nutritional status of plants). 5. Microbes stimulated the release of C- and N-compounds, but rapidly respired 65—85% of the root-borne C-compounds, thereby putting a burden on the C-budget of the “host” plant. 6. It could be shown by the example of hup⁺ Rhizobium meliloti strains (tested by ³H₂-incorporation) and the wheat-Serratia-association, that energy efficient microbenplant systems can improve plant performance.