Model of apparent and real priming effects: Linking microbial activity with soil organic matter decomposition

The most frequently used models simulating soil organic matter (SOM) dynamics are based on first-order kinetics. These models fail to describe and predict such interactions as priming effects (PEs), which are short-term changes in SOM decomposition induced by easily available C or N sources. We hypothesized that if decomposition rate depends not only on size of the SOM pool, but also on microbial biomass and its activity, then PE can be simulated. A simple model that included these interactions and that consisted of three C pools - SOM, microbial biomass, and easily available C - was developed. The model was parameterized and evaluated using results of ¹²C-CO₂ and ¹⁴C-CO₂ efflux after adding ¹⁴C-labeled glucose to a loamy Haplic Luvisol. Experimentally measured PE, i.e., changes in SOM decomposition induced by glucose, was compared with simulated PE. The best agreement between measured and simulated CO₂ efflux was achieved by considering both the total amount of microbial biomass and its activity. Because it separately described microbial turnover and SOM decomposition, the model successfully simulated apparent and real PE.The proposed PE model was compared with three alternative approaches with similar complexity but lacking interactions between the pools and neglecting the activity of microbial biomass. The comparison showed that proposed new model best described typical PE dynamics in which the first peak of apparent PE lasted for 1 day and the subsequent real PE gradually increased during 60 days. This sequential decomposition scheme of the new model, with immediate microbial consumption only of soluble substrate, was superior to the parallel decomposition scheme with simultaneous microbial consumption of two substrates with different decomposability. Incorporating microbial activity function in the model improved the fit of simulation results with experimental data, by providing the flexibility necessary to properly describe PE dynamics. We conclude that microbial biomass should be considered in models of C and N dynamics in soil not only as a pool but also as an active driver of C and N turnover.     

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.     

Rhizosphere priming effect: Its functional relationships with microbial turnover, evapotranspiration, and C–N budgets

Understanding soil organic matter (SOM) decomposition and its interaction with rhizosphere processes is a crucial topic in soil biology and ecology. Using a natural ¹³C tracer method to separately measure SOM-derived CO₂ from root-derived CO₂, this study aims to connect the level of rhizosphere-dependent SOM decomposition with the C and N balance of the whole plant–soil system, and to mechanistically link the rhizosphere priming effect to soil microbial turnover and evapotranspiration. Results indicated that the magnitude of the rhizosphere priming effect on SOM decomposition varied widely, from zero to more than 380% of the unplanted control, and was largely influenced by plant species and phenology. Balancing the extra soil C loss from the strong rhizosphere priming effect in the planted treatments with C inputs from rhizodeposits and root biomass, the whole plant–soil system remained with a net carbon gain at the end of the experiment. The increased soil microbial biomass turnover rate and the enhanced evapotranspiration rate in the planted treatments had clear positive relationships with the level of the rhizosphere priming effect. The rhizosphere enhancement of soil carbon mineralization in the planted treatments did not result in a proportional increase in net N mineralization, suggesting a possible de-coupling of C cycling with N cycling in the rhizosphere.     

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.     

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.     

Turnover of grain legume N rhizodeposits and effect of rhizodeposition on the turnover of crop residues

The turnover of N derived from rhizodeposition of faba bean (Vicia faba L.), pea (Pisum sativum L.) and white lupin (Lupinus albus L.) and the effects of the rhizodeposition on the subsequent C and N turnover of its crop residues were investigated in an incubation experiment (168 days, 15 °C). A sandy loam soil for the experiment was either stored at 6 °C or planted with the respective grain legume in pots. Legumes were in situ ¹⁵N stem labelled during growth and visible roots were removed at maturity. The remaining plant-derived N in soil was defined as N rhizodeposition. In the experiment the turnover of C and N was compared in soils with and without previous growth of three legumes and with and without incorporation of crop residues. After 168 days, 21% (lupin), 26% (faba bean) and 27% (pea) of rhizodeposition N was mineralised in the treatments without crop residues. A smaller amount of 15–17% was present as microbial biomass and between 30 and 55% of mineralised rhizodeposition N was present as microbial residue pool, which consists of microbial exoenzymes, mucous substances and dead microbial biomass. The effect of rhizodeposition on the C and N turnover of crop residues was inconsistent. Rhizodeposition increased the crop residue C mineralisation only in the lupin treatment; a similar pattern was found for microbial C, whereas the microbial N was increased by rhizodeposition in all treatments. The recovery of residual ¹⁵N in the microbial and mineral N pool was similar between the treatments containing only labelled crop residues and labelled crop residues + labelled rhizodeposits. This indicates a similar decomposability of both rhizodeposition N and crop residue N and may be attributable to an immobilisation of both N sources (rhizodeposits and crop residues) as microbial residues and a subsequent remineralisation mainly from this pool.Abbreviations C or N_dec C or N decomposed from residues - C or N_mic microbial C or N - C or N_micres microbial residue C or N - C or N_min mineralised C or N - C or N_input added C or N as crop residues and/or rhizodeposits - dfr derived from residues - dfR derived from rhizodeposition - Ndfr N derived from residues - NdfR N derived from rhizodeposition - N_loss losses of N derived from residues - SOM soil organic matter - WHC water holding capacity     

Priming effect of the addition of maize to a Japanese volcanic ash soil and its temperature sensitivity: a short-term incubation study

ABSTRACT

The response of soil organic matter (SOM) to global warming is a crucial subject. However, the temperature sensitivity of SOM turnover remains largely uncertain. Changes in the mineralization of native SOM, i.e., priming effect (PE) may strongly affect the temperature sensitivity of SOM turnover in the presence of global warming. We investigated the direction and magnitude of the PE in a Japanese volcanic ash soil at different temperatures (15°C, 25°C, and 35°C) using a natural ¹³C tracer (C4-plant, maize leaf) in a short-term (25 days) incubation study. In addition, we evaluated the temperature sensitivity expressed as Q₁₀ value with and without the addition of maize to the soil and their relations to PE. We found that positive PE occurred at each temperature condition and tended to increase with decreased temperature, and these PE results were consistent with the microbial biomass at the end of the incubation period. CO₂ emission from control soil (without maize) increased with increasing temperature (Q₁₀ = 2.6), but CO₂ emission from the soil with added maize did not significantly change with increasing temperature (Q₁₀ = 1.0). This was caused by the suppression of CO₂ emission from the soil with increasing temperature (Q₁₀ = 0.9). On the other hand, soil-originated CO₂ emission clearly increased with increasing temperature (Q₁₀ = 3.4) when Q₁₀ values were calculated on the assumption that the temperature and substrate supply increase at the same time (from 25°C). These results suggest that not only the temperature increase but also the labile carbon supply may be important for the temperature sensitivity of Japanese volcanic ash soil.     

C and N turnover in structurally intact soils of different texture

The turnover of native and applied C and N in undisturbed soil samples of different texture but similar mineralogical composition, origin and cropping history was evaluated at −10 kPa water potential. Cores of structurally intact soil with 108, 224 and 337 g clay kg⁻¹ were horizontially sliced and ¹⁵N-labelled sheep faeces was placed between the two halves of the intact core. The cores together with unamended treatments were incubated in the dark at 20 °C and the evolution of CO₂-C determined continuously for 177 d. Inorganic and microbial biomass N and ¹⁵N were determined periodically. Net nitrification was less in soil amended with faeces compared with unamended soil. When adjusted for the NO₃-N present in soil before faeces was applied, net nitrification became negative indicating that NO₃-N had been immobilized or denitrified. The soil most rich in clay nitrified least N and ¹⁵N. The amounts of N retained in the microbial biomass in unamended soils increased with clay content. A maximum of 13% of the faeces ¹⁵N was recovered in the microbial biomass in the amended soils. CO₂-C evolution increased with clay content in amended and unamended soils. CO₂-C evolution from the most sandy soil was reduced due to a low content of potentially mineralizable native soil C whereas the rate constant of C mineralization rate peaked in this soil. When the pool of potentially mineralizable native soil C was assumed proportional to volumetric water content, the three soils contained similar proportions of potentially mineralizable native soil C but the rate constant of C mineralization remained highest in the soil with least clay. Thus although a similar availability of water in the three soils was ensured by their identical matric potential, the actual volume of water seemed to determine the proportion of total C that was potentially mineralizable. The proportion of mineralizable C in the faeces was similar in the three soils (70% of total C), again with a higher rate constant of C mineralization in the soil with least clay. It is hypothesized that the pool of potentially mineralizable C and C rate constants fluctuate with the soil water content.     

Soil Carbon Dynamics Under Changing Climate——A Research Transition from Absolute to Relative Roles of Inorganic Nitrogen Pools and Associated Microbial Processes: A Review

It is globally accepted that soil carbon (C) dynamics are at the core of interlinked environmental problems,deteriorating soil quality and changing climate.Its management remains a complex enigma for the scientific community due to its intricate relationship with soil nitrogen (N) availability and moisture-temperature interactions.This article reviews the management aspects of soil C dynamics in light of recent advances,particularly in relation to the availability of inorganic N pools and associated microbial processes under changing climate.Globally,drastic alterations in soil C dynamics under changing land use and management practices have been primarily attributed to the variation in soil N availability,resulting in a higher decomposition rate and a considerable decline in soil organic C (SOC) levels due to increased soil CO2 emissions,degraded soil quality,and increased atmospheric CO2 concentrations,leading to climate warming.Predicted climate warming is proposed to enhance SOC decomposition,which may further increase soil N availability,leading to higher soil CO2 efflux.However,a literature survey revealed that soil may also act as a potential C sink,if we could manage soil inorganic N pools and link microbial processes properly.Studies also indicated that the relative,rather than the absolute,availability of inorganic N pools might be of key importance under changing climate,as these N pools are variably affected by moisture-temperature interactions,and they have variable impacts on SOC turnover.Therefore,multi-factorial studies are required to understand how the relative availability of inorganic N pools and associated microbial processes may determine SOC dynamics for improved soil C management.     

Short-term dynamics of carbon and nitrogen after tillage in a freshwater marsh of northeast China

The Sanjiang Plain, one of the largest freshwater marshes in China, has experienced intensive cultivation over the past 50 years. However, there were few reports of short-term dynamics of soil carbon and nitrogen and CO₂ emission after tillage. In this paper, we studied the short-term dynamics of carbon and nitrogen after tillage in a freshwater marsh of northeast China. The results showed that response of carbon and nitrogen dynamic to tillage was different for intact wetland and soil cultivated for 10 years. Tillage was followed by immediate and significant increases in CO₂ efflux, which peaked at 0.25 h after tillage, four times higher than control in the wetland soils; while, only 2.5 times higher than control in the cultivated soils. Although, dissolved organic C (DOC) increased, the relative stability of microbial biomass C (MBC) pools together with the decreased respiration in the wetland soil suggested that the tillage did not lead to a burst in microbial activity and growth. Other factors such as moisture content before and after tillage may play an important role in determining microbial activity in the intact wetland. On the contrary, although dissolved organic C did not change, MBC pools, and soil respiration increase after tillage, suggesting tillage led to an increase in microbial activity and growth in the cultivated soil. Tillage initiated changes in soil aeration that was an important factor affecting soil microbiology in the long history of cultivation. Net N mineralization and nitrification occurred in both wetland and cultivated soils, but at different rates after tillage that in the intact wetland soil was higher than cultivated soil. Macroaggregates in the wetland soil would be expected to contain larger amounts of organic matter, and thus release a larger source of newly available substrate for microbes after tillage. In the intact wetland soil, ammonium, nitrate, and dissolved organic N (DON) concentrations were significantly negatively correlated to soil moisture (p < 0.01), suggesting high soil moisture in the natural wetland was not in favor of N mineralization.     

Carbon mineralization is promoted by phosphorus and reduced by nitrogen addition in the organic horizon of northern hardwood forests

Limitations to the respiratory activity of heterotrophic soil microorganisms exert important controls of CO₂ efflux from soils. In the northeastern US, ecosystem nutrient status varies across the landscape and changes with forest succession following disturbance, likely impacting soil microbial processes regulating the transformation and emission of carbon (C). We tested whether nitrogen (N) or phosphorus (P) limit the mineralization of soil organic C (SOC) or that of added C sources in the Oe horizon of successional and mature northern hardwood forests in three locations in central New Hampshire, USA. Added N reduced mineralization of C from SOC and from added leaf litter and cellulose. Added P did not affect mineralization from SOC; however, it did enhance mineralization of litter- and cellulose- C in organic horizons from all forest locations. Added N increased microbial biomass N and K₂SO₄-extractable DON pools, but added P had no effect. Microbial biomass C increased with litter addition but did not respond to either nutrient. The direction of responses to added nutrients was consistent among sites and between forest ages. We conclude that in these organic horizons limitation by N promotes mineralization of C from SOC, whereas limitation by P constrains mineralization of C from new organic inputs. We also suggest that N suppresses respiration in these organic horizons either by relieving the N limitation of microbial biomass synthesis, or by slowing turnover of C through the microbial pool; concurrent measures of microbial growth and turnover are needed to resolve this question.     

Microbial use of maize cellulose and sugarcane sucrose monitored by changes in the C/C ratio

An arable soil with organic matter formed from C₃-vegetation was amended initially with maize cellulose (C₄-cellulose) and sugarcane sucrose (C₄-sucrose) in a 67-day laboratory incubation experiment with microcosms at 25 °C. The amount and isotopic composition (¹³C/¹²C) of soil organic C, CO₂ evolved, microbial biomass C, and microbial residue C were determined to prove whether the formation of microbial residues depends on the quality of the added C source adjusted with NH₄NO₃ to the same C/N ratio of 15. In a subsequent step, C₃-cellulose (3 mg C g⁻¹ soil) was added without N to soil to determine whether the microbial residues formed initially from C₄-substrate are preferentially decomposed to maintain the N-demand of the soil microbial community. At the end of the experiment, 23% of the two C₄-substrates added was left in the soil, while 3% and 4% of the added C₄-cellulose and C₄-sucrose, respectively, were found in the microbial biomass. The addition of the two C₄-substrates caused a significant 100% increase in C₃-derived CO₂ evolution during the 5-33 day incubation period. The addition of C₃-cellulose caused a significant 50% increase in C₄-derived CO₂ evolution during the 38-67 day incubation period. The decrease in microbial biomass C₄-C accounted for roughly 60% of this increase. Cellulose addition promoted microorganisms strongly able to recycle N immediately from their own tissue by “cryptic growth” instead of incorporating NO₃⁻ from the soil solution. The differences in quality of the microbial residues produced by C₄-cellulose and C₄-sucrose decomposing microorganisms are also reflected by the difference in the rates of CO₂ evolution, but not in the rates of net N mineralization.     

Extractability of labelled microbial biomass N by electro-ultrafiltration and CaCl2 extraction

A laboratory soil incubation and a pot experiment with ryegrass were carried out in order to examine the extractability of microbial biomass N by using either 10-mM CaCl₂ extraction or the electro-ultrafiltration (EUF) method. The aim of the experiment was to test the hypothesis whether the organic N (Norg) extracted by EUF or CaCl₂ from dried soil samples represents a part of the microbial biomass. For the laboratory incubation a ¹⁵N-labelled Escherichia coli suspension was mixed with the soil. For the pot experiment a suspension of ¹⁵N-labelled bacteria was applied which had previously been isolated from the soil used. Soil samples of both treatments, with and without applied bacterial suspension, were extracted by EUF and CaCl₂. The extractability of applied microbial biomass was estimated from the difference in extractable Norg between the two treatments. In addition, the N isotopic composition in the upper plant matter, in the soil, and in organic and inorganic N fractions of EUF and CaCl₂ extracts was analysed. Both experiments showed that the applied microbial biomass was highly accessible to mineralization and thus represented potentially mineralizable N. However, this mineralizable N was not extractable by CaCl₂ or by the EUF method. It was, therefore, concluded that the organic N released on soil drying and which was thus extractable was derived from the non-biomass soil organic matter. The result suggests that both extraction methods may provide a suitable index for mineralizable N only in cases where the decomposable organic substrates are derived mainly from sources other than the living soil biota.Dedicated to Professor J. C. G. Ottow on the occasion of his 60th birthday     

Sources of CO2 efflux from soil and review of partitioning methods

Five main biogenic sources of CO₂ efflux from soils have been distinguished and described according to their turnover rates and the mean residence time of carbon. They are root respiration, rhizomicrobial respiration, decomposition of plant residues, the priming effect induced by root exudation or by addition of plant residues, and basal respiration by microbial decomposition of soil organic matter (SOM). These sources can be grouped in several combinations to summarize CO₂ efflux from the soil including: root-derived CO₂, plant-derived CO₂, SOM-derived CO₂, rhizosphere respiration, heterotrophic microbial respiration (respiration by heterotrophs), and respiration by autotrophs. These distinctions are important because without separation of SOM-derived CO₂ from plant-derived CO₂, measurements of total soil respiration have very limited value for evaluation of the soil as a source or sink of atmospheric CO₂ and for interpreting the sources of CO₂ and the fate of carbon within soils and ecosystems. Additionally, the processes linked to the five sources of CO₂ efflux from soil have various responses to environmental variables and consequently to global warming. This review describes the basic principles and assumptions of the following methods which allow SOM-derived and root-derived CO₂ efflux to be separated under laboratory and field conditions: root exclusion techniques, shading and clipping, tree girdling, regression, component integration, excised roots and insitu root respiration; continuous and pulse labeling, ¹³C natural abundance and FACE, and radiocarbon dating and bomb-¹⁴C. A short sections cover the separation of the respiration of autotrophs and that of heterotrophs, i.e. the separation of actual root respiration from microbial respiration, as well as methods allowing the amount of CO₂ evolved by decomposition of plant residues and by priming effects to be estimated. All these methods have been evaluated according to their inherent disturbance of the ecosystem and C fluxes, and their versatility under various conditions. The shortfalls of existing approaches and the need for further development and standardization of methods are highlighted.     

Decomposition of N-labelled maize leaves in soil affected by endogeic geophagous Aporrectodea caliginosa

A microcosm experiment was carried out for 56 days at 12 °C to evaluate the feeding effects of the endogeic geophagous earthworm species Aporrectodea caliginosa on the microbial use of ¹⁵N-labelled maize leaves (Zea mays) added as 5 mm particles equivalent to 1 mg C and 57 μg N g⁻¹ soil. The dry weight of A. caliginosa biomass decreased in the no-maize treatment by 10% during the incubation and increased in the maize leaf treatments by 18%. Roughly 5% and 10% of the added maize leaf-C and leaf-N, respectively, were incorporated into the biomass of A. caliginosa. About 29% and 33% of the added maize leaf-C were mineralised to CO₂ in the no-earthworm and earthworm treatments, respectively. The presence of A. caliginosa significantly increased soil-derived CO₂ production by 90 μg g⁻¹ soil in the no-maize and maize leaf treatments, but increased the maize-derived CO₂ production only by 40 μg g⁻¹ soil. About 10.5% of maize leaf-C and leaf-N was incorporated into the soil microbial biomass in the absence of earthworms, but only 6% of the maize leaf-C and 3% of the maize leaf-N in the presence of earthworms. A. caliginosa preferentially fed on N rich, maize leaf-colonizing microorganisms to meet its N demand. This led to a significantly increased C/N ratio of the unconsumed microbial biomass in soil. The ergosterol-to-microbial biomass C ratio was not significantly decreased by the presence of earthworms. A. caliginosa did not directly contribute to comminution of plant residues, as indicated by the absence of any effects on the contents of the different particulate organic matter fractions, but mainly to grazing of residue-colonizing microorganisms, increasing their turnover considerably.     

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.     

Dynamics of maize (Zea mays L.) leaf straw mineralization as affected by the presence of soil and the availability of nitrogen

An incubation experiment was carried out with maize (Zea mays L.) leaf straw to analyze the effects of mixing the residues with soil and N amendment on the decomposition process. In order to distinguish between soil effects and nitrogen effects for both the phyllospheric microorganisms already present on the surface of maize straw and soil microorganisms the N amendment was applied in two different placements: directly to the straw or to the soil. The experiment was performed in dynamic, automated microcosms for 22 days at 15 °C with 7 treatments: (1) untreated soil, (2) non-amended maize leaf straw without soil, (3) N amended maize leaf straw without soil, (4) soil mixed with maize leaf straw, (5) N amended soil, (6) N amended soil mixed with maize leaf straw, and (7) soil mixed with N amended maize leaf straw. ¹⁵NH₄¹⁵NO₃ (5 at%) was added. Gas emissions (CO₂, ¹³CO₂ and N₂O) were continuously recorded throughout the experiment. Microbial biomass C, biomass N, ergosterol, δ¹³C of soil organic C and of microbial biomass C as well as ¹⁵N in soil total N, mineral N and microbial biomass N were determined in soil samples at the end of the incubation. The CO₂ evolution rate showed a lag-phase of two days in the non-amended maize leaf straw treatment without soil, which was completely eliminated when mineral N was added. The addition of N generally increased the CO₂ evolution rate during the initial stages of maize leaf straw decomposition, but not the cumulative CO₂ production. The presence of soil caused roughly a 50% increase in cumulative CO₂ production within 22 days in the maize straw treatments due to a slower decrease of CO₂ evolution after the initial activity peak. Since there are no limitations of water or N, we suggest that soil provides a microbial community ensuring an effective succession of straw decomposing microorganisms. In the treatments where maize and soil was mixed, 75% of microbial biomass C was derived from maize. We concluded that this high contribution of maize using microbiota indicates a strong influence of organisms of phyllospheric origin to the microbial community in the soil after plant residues enter the soil.     

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.     

Three-source partitioning of CO2 efflux from soil planted with maize by C natural abundance fails due to inactive microbial biomass

A theoretical approach to the partitioning of carbon dioxide (CO₂) efflux from soil with a C₃ vegetation history planted with maize (Zea mays), a C₄ plant, into three sources, root respiration (RR), rhizomicrobial respiration (RMR), and microbial soil organic matter (SOM) decomposition (SOMD), was examined. The δ¹³C values of SOM, roots, microbial biomass, and total CO₂ efflux were measured during a 40-day growing period. A three-source isotopic mass balance based on the measured δ¹³C values and on assumptions made in other studies showed that RR, RMR, and SOMD amounted to 91%, 4%, and 5%, respectively. Two assumptions were thoroughly examined in a sensitivity analysis: the absence of ¹³C fractionation and the conformity of δ¹³C of microbial CO₂ and that of microbial biomass. This approach strongly overestimated RR and underestimated RMR and microbial SOMD. CO₂ efflux from unplanted soil was enriched in ¹³C by 2.0‰ compared to microbial biomass. The consideration of this ¹³C fractionation in the mass balance equation changed the proportions of RR and RMR by only 4% and did not affect SOMD. A calculated δ¹³C value of microbial CO₂ by a mass balance equation including active and inactive parts of microbial biomass was used to adjust a hypothetical below-ground CO₂ partitioning to the measured and literature data. The active microbial biomass in the rhizosphere amounted to 37% to achieve an appropriate ratio between RR and RMR compared to measured data. Therefore, the three-source partitioning approach failed due to a low active portion of microbial biomass, which is the main microbial CO₂ source controlling the δ¹³C value of total microbial biomass. Since fumigation-extraction reflects total microbial biomass, its δ¹³C value was unsuitable to predict δ¹³C of released microbial CO₂ after a C₃-C₄ vegetation change. The second adjustment to the CO₂ partitioning results in the literature showed that at least 71% of the active microbial biomass utilizing maize rhizodeposits would be necessary to achieve that proportion between RR and RMR observed by other approaches based on ¹⁴C labelling. The method for partitioning total below-ground CO₂ efflux into three sources using a natural ¹³C labelling technique failed due to the small proportion of active microbial biomass in the rhizosphere. This small active fraction led to a discrepancy between δ¹³C values of microbial biomass and of microbially respired CO₂.     

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.