Plant biomass influences rhizosphere priming effects on soil organic matter decomposition in two differently managed soils

We used a continuous labeling method of naturally ¹³C-depleted CO₂ in a growth chamber to test for rhizosphere effects on soil organic matter (SOM) decomposition. Two C3 plant species, soybean (Glycine max) and sunflower (Helianthus annus), were grown in two previously differently managed soils, an organically farmed soil and a soil from an annual grassland. We maintained a constant atmospheric CO₂ concentration at 400±5 ppm and δ¹³C signature at −24.4‰ by regulating the flow of naturally ¹³C-depleted CO₂ and CO₂-free air into the growth chamber, which allowed us to separate new plant-derived CO₂-C from original soil-derived CO₂-C in soil respiration. Rhizosphere priming effects on SOM decomposition, i.e., differences in soil-derived CO₂-C between planted and non-planted treatments, were significantly different between the two soils, but not between the two plant species. Soil-derived CO₂-C efflux in the organically farmed soil increased up to 61% compared to the no-plant control, while the annual grassland soil showed a negligible increase (up to 5% increase), despite an overall larger efflux of soil-derived CO₂-C and total soil C content. Differences in rhizosphere priming effects on SOM decomposition between the two soils could be largely explained by differences in plant biomass, and in particular leaf biomass, explaining 49% and 74% of the variation in primed soil C among soils and plant species, respectively. Nitrogen uptake rates by soybean and sunflower was relatively high compared to soil C respiration and associated N mineralization, while inorganic N pools were significantly depleted in the organic farm soil by the end of the experiment. Despite relatively large increases in SOM decomposition caused by rhizosphere effects in the organic farm soil, the fast-growing soybean and sunflower plants gained little extra N from the increase in SOM decomposition caused by rhizosphere effects. We conclude that rhizosphere priming effects of annual plants on SOM decomposition are largely driven by plant biomass, especially in soils of high fertility that can sustain high plant productivity.     

Tracing the source and fate of dissolved organic matter in soil after incorporation of a C labelled residue: A batch incubation study

While dissolved organic matter (DOM) in soil solution is a small but reactive fraction of soil organic matter, its source and dynamics are unclear. A laboratory incubation experiment was set up with an agricultural topsoil amended with ¹³C labelled maize straw. The dissolved organic carbon (DOC) concentration in soil solution increased sharply from 25 to 186 mg C L⁻¹ 4 h after maize amendment, but rapidly decreased to 42 mg C L⁻¹ and reached control values at and beyond 2 months. About 65% of DOM was straw derived after 4 h, decreasing to 29% after one day and only 1.3% after 240 days. A significant priming effect of the straw on the release of autochthonous DOM was found. The DOM fractionation with DAX-8 resin revealed that 98% of the straw derived DOM was hydrophilic in the initial pulse while this hydrophilic fraction was 20-30% in control samples. This was in line with the specific UV absorbance of the DOM which was significantly lower in the samples amended with maize residues than in the control samples. The δ¹³C of the respired CO₂ matched that of DOC in the first day after amendment but exceeded it in following days. The straw derived C fractions in respired CO₂ and in microbial biomass were similar between 57 and 240 days after amendment but were 3-10 fold above those in the DOM. This suggests that the solubilisation of C from the straw is in steady state with the DOM degradation or that part of the straw is directly mineralised without going into solution. This study shows that residue application releases a pulse of hydrophilic DOM that temporarily (<3 days) dominates the soil DOM pool and the degradable C. However, beyond that pulse the majority of DOM is derived from soil organic matter and its isotope signature differs from microbial biomass and respired C, casting doubt that the DOM pool in the soil solution is the major bioaccessible C pool in soil.     

Rhizosphere priming of soil organic matter by bacterial groups in a grassland soil

Plants often impact the rate of native soil organic matter turnover through root interactions with soil organisms; however the role of root-microbial interactions in mediation of the “priming effect” is not well understood. We examined the effects of living plant roots and N fertilization on belowground C dynamics in a California annual grassland soil (Haploxeralf) during a two-year greenhouse study. The fate of ¹³C-labeled belowground C (roots and organic matter) was followed under planted (Avena barbata) and unplanted conditions, and with and without supplemental N (20 kg N ha⁻¹ season⁻¹) over two periods of plant growth, each followed by a dry, fallow period of 120 d. Turnover of belowground ¹³C SOM was followed using ¹³C-phospholipid fatty acid (PLFA) biomarkers. Living roots increased the turnover and loss of belowground ¹³C compared with unplanted soils. Planted soils had 20% less belowground ¹³C present than in unplanted soils after 2 cycles of planting and fallow. After 2 treatment cycles, unlabeled soil C was 4.8% higher in planted soils than unplanted. The addition of N to soils decreased the turnover of enriched belowground ¹³C during the first treatment season in both planted and unplanted soils, however no effect of N was observed thereafter. Our findings suggest that A. barbata may increase soil C levels over time because root and exudate C inputs are significant, but that increase will be moderated by an overall faster C mineralization rate of belowground C. N addition may slow soil C losses; however, the effect was minor and transient in this system. The labeled root-derived ¹³C was initially recovered in gram negative (highest enrichment), gram positive, and fungal biomarkers. With successive growing seasons, the labeled C in the gram negative and fungal markers declined, while gram positive markers continued to accumulate labeled belowground C. The rhizosphere of A. barbata shifted the microbial community composition, resulting in greater abundances of gram negative markers and lower abundances of gram positive, actinobacteria and cyclopropyl PLFA markers compared to unplanted soil. However, the longer-term utilization of labeled belowground C by gram positive bacteria was enhanced in the rhizosphere microbial community compared with unplanted soils. We suggest that the activities of gram positive bacteria may be major controllers of multi-year rhizosphere-related priming of SOM decomposition.     

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.     

Microbial community abundance and structure are determinants of soil organic matter mineralisation in the presence of labile carbon

Altered rates of native soil organic matter (SOM) mineralisation in the presence of labile C substrate (‘priming’), is increasingly recognised as central to the coupling of plant and soil-biological productivity and potentially as a key process mediating the C-balance of soils. However, the mechanisms and controls of SOM-priming are not well understood. In this study we manipulated microbial biomass size and composition (chloroform fumigation) and mineral nutrient availability to investigate controls of SOM-priming. Effects of applied substrate (¹³C-glucose) on mineralisation of native SOM were quantified by isotopic partitioning of soil respiration. In addition, the respective contributions of SOM-C and substrate-derived C to microbial biomass carbon (MBC) were quantified to account for pool-substitution effects (‘apparent priming’). Phospholipid fatty acid (PLFA) profiles of the soils were determined to establish treatment effects on microbial community structure, while the ¹³C-enrichment of PLFA biomarkers was used to establish pathways of substrate-derived C-flux through the microbial communities. The results indicated that glucose additions increased SOM-mineralisation in all treatments (positive priming). The magnitude of priming was reduced in fumigated soils, concurrent with reduced substrate-derived C-flux through putative SOM-mineralising organisms (fungi and actinomycetes). Nutrient additions reduced the magnitude of positive priming in non-fumigated soils, but did not affect the distribution of substrate-derived C in microbial communities. The results support the view that microbial community composition is a determinant of SOM-mineralisation, with evidence that utilisation of labile substrate by fungal and actinomycete (but not Gram-negative) populations promotes positive SOM-priming.     

Sources and mechanisms of priming effect induced in two grassland soils amended with slurry and sugar

The mechanisms and specific sources of priming effects, i.e. short term changes of soil organic matter (SOM) decomposition after substance addition, are still not fully understood. These uncertainties are partly method related, i.e. until now only two C sources in released CO₂ could be identified. We used a novel approach separating three carbon (C) sources in CO₂ efflux from soil. The approach is based on combination of different substances originated from C₃ or C₄ plants in different treatments and identical transformation of substances like C₃ sugar (from sugar beet) and C₄ sugar (from sugar cane). We investigated the influence of the addition of two substances having different microbial utilizability, i.e. slurry and sugar on the SOM or/and slurry decomposition in two grassland soils with different levels of C_org (2.3 vs. 5.1% C). Application of slurry to the soil slightly accelerated the SOM decomposition. Addition of sugar lead to changes of SOM and slurry decomposition clearly characterized by two phases: immediately after sugar addition, the microorganisms switched from the decomposition of hardly utilizable SOM to the decomposition of easily utilizable sugar. This first phase was very short (2-3 days), hence was frequently missed in other experiments. The second phase showed a slightly increased slurry and SOM decomposition (compared to the soil without sugar). The separation of three sources in CO₂ efflux from grassland soils allowed us to conclude that the C will be utilized according to its utilizability: sugar>slurry>SOM. Additionally, decomposition of more inert C (here SOM) during the period of intensive sugar decomposition was strongly reduced (negative priming effect). We conclude that, priming effects involve a chain of mechanisms: (i) preferential substrate utilization, (ii) activation of microbial biomass by easily utilizable substrate (iii) subsequent increased utilization of following substrates according to their utilizability, and (iv) decline to initial state.     

A call to investigate drivers of soil organic matter retention vs. mineralization in a high CO2 world

Understanding how elevated atmospheric CO₂ alters the formation and decomposition of soil organic carbon (SOC) is important but challenging. If elevated CO₂ induces even small changes in rates of formation or decay of SOC, there could be substantial feedbacks on the atmosphere's concentration of CO₂. However, the long turnover times of many SOC pools - decades to centuries - make the detection of changes in the soil's pool size difficult. Long-term CO₂ enrichment experiments have offered unprecedented opportunities to explore these issues in intact ecosystems for more than a decade. Increased NPP with elevated CO₂ has prompted the hypothesis that SOC may increase at the same time that increased vegetation nitrogen (N) uptake and accumulation indicates probable declines in SON. Varying investigators thus have hypothesized that SOC will increase and SON will decline to explain increased NPP with elevated CO₂; researchers also invoke biogeochemical theory and stoichiometric constraints to argue for strong limitations on the co-occurrence of these phenomena. We call for researchers to investigate two broad research questions to elucidate the drivers of these processes. First, we ask how elevated CO₂ influences compound structure and stoichiometry of that proportion of NPP retained by soil profiles for relatively long time periods. We also call for investigations of the mechanisms underlying the decomposition of mineralizable organic matter with elevated CO₂. Specifically, we need to understand how elevated CO₂ influences microbial priming (driven by enhanced microbial energy needs associated with increases in biomass or activity) and microbial mining of N (driven by enhanced microbial N demand associated with greater vegetative N uptake), two processes that necessarily will be constrained by the stoichiometry of both substrates and microbial demands. Applying technologies such as nuclear magnetic resonance and the detection of biomarkers that reveal organic matter structure and origins, and studying microbial stoichiometric constraints, will dramatically improve our ability to predict future patterns of ecosystem C and N cycling.     

Priming effects: Interactions between living and dead organic matter

In this re-evaluation of our 10-year old paper on priming effects, I have considered the latest studies and tried to identify the most important needs for future research. Recent publications have shown that the increase or decrease in soil organic matter mineralization (measured as changes of CO₂ efflux and N mineralization) actually results from interactions between living (microbial biomass) and dead organic matter. The priming effect (PE) is not an artifact of incubation studies, as sometimes supposed, but is a natural process sequence in the rhizosphere and detritusphere that is induced by pulses or continuous inputs of fresh organics. The intensity of turnover processes in such hotspots is at least one order of magnitude higher than in the bulk soil. Various prerequisites for high-quality, informative PE studies are outlined: calculating the budget of labeled and total C; investigating the dynamics of released CO₂ and its sources; linking C and N dynamics with microbial biomass changes and enzyme activities; evaluating apparent and real PEs; and assessing PE sources as related to soil organic matter stabilization mechanisms. Different approaches for identifying priming, based on the assessment of more than two C sources in CO₂ and microbial biomass, are proposed and methodological and statistical uncertainties in PE estimation and approaches to eliminating them are discussed. Future studies should evaluate directions and magnitude of PEs according to expected climate and land-use changes and the increased rhizodeposition under elevated CO₂ as well as clarifying the ecological significance of PEs in natural and agricultural ecosystems. The conclusion is that PEs - the interactions between living and dead organic matter - should be incorporated in models of C and N dynamics, and that microbial biomass should regarded not only as a C pool but also as an active driver of C and N turnover.     

Near infrared reflectance spectroscopy for determination of organic matter fractions including microbial biomass in coniferous forest soils

The objective of this work was to investigate the usefulness of near infrared reflectance spectroscopy (NIRS) in determining some C and N fractions of soils: labile compounds, microbial biomass, compounds derived from added ¹³C- and ¹⁵N-labelled straw. Soil samples were obtained from a previous experiment where soils were labelled by addition of ¹³C- and ¹⁵N-labelled wheat straw and incubated in coniferous forests in northern Sweden (64-60°N) and south France (43°N). The incubation lasted three years with 7-9 samplings at regular time steps and four replicates at each sampling (204 samples). Samples were scanned using a near infrared reflectance spectrophotometer (NIRSystem 6500). Calibrations were obtained by using a modified partial least squares regression technique with reference data on total C and N, ¹³C, ¹⁵N, control extract-C, -N, -¹³C and -¹⁵N, fumigated extract-C, -N, -¹³C and -¹⁵N, biomass-C, -N, -¹³C and -¹⁵N contents. Mathematical treatments of the absorbance data were first or second derivative with a gap from 4 to 10 nm. The standard error of calibration (SEC)-to-standard deviation of the reference measurements ratio was ≤0.2 for 10 models, namely total C and N, ¹³C, ¹⁵N, control extract-C, fumigated extract-C and -N, biomass-C and -N and biomass-¹⁵N models and therefore considered as very good. With an R²=0.955, the fumigated extract-¹⁵N model is also good. The standard error of performance calculated on the independent set of data and SEC were within 20% of each other for all the best equations except for the biomass-¹⁵N model. The ability of NIRS to detect ¹³C and ¹⁵N in total C and N and in the extracts is noteworthy, not because of its predictive function that is not really of interest in this case, but because it indicates that the spectra kept the signature of the properties of the organic matter derived from the straw even after two- or three-year decomposition. The incorporation of the ¹³C in the biomass was less well predicted than that of the ¹⁵N. This could indicate that the biomass derived from the straw was characterised by a particular protein or amino acid composition compared to the total biomass that includes a large proportion of dormant micro-organisms. The predictive ability of NIRS for microbial biomass-C and -N is particularly interesting because the conventional analyses are time consuming. In addition, NIRS allows detecting analytical errors.     

Determination of the fate of C labelled maize and wheat rhizodeposit-C in two agricultural soils in a greenhouse experiment under C-CO2-enriched atmosphere

A deeper understanding of the contribution of carbon (C) released by plant roots (rhizodeposition) to soil organic matter (SOM) can help to increase our knowledge of global C-cycling. These insights can eventually lead to sustainable management of SOM especially in agricultural systems. This study was conducted to determine the fate of ¹³C labelled rhizodeposit-C of maize and wheat plants. They were grown in a greenhouse in permeable nylon bags filled with upper soil material from two agricultural soils of the same location, but with different crop yields. The bags were placed into pots, which were also filled with soil surrounding the bags. Soil inside the bags was considered as rhizosphere soil, wheras the one outside the bags represented bulk soil. The contributions of rhizodeposits to water extractable organic carbon (WEOC), microbial biomass-C (MB-C), CO₂-C evolution, and total organic carbon (C_org) were investigated during a 7-week growing period. The WEOC, MB-C, CO₂-C, C_org contents and the respective δ¹³C values were determined regularly, and a newly developed method for determining δ¹³C values in soil extracts was applied.In both soils, regardless of crop yield potential, significant incorporation of rhizodeposition-derived C was observed in the MB-C, CO₂-C, and C_org pool, but not in the WEOC. The pattern of C incorporation into the different pools was the same for both soils with both plants, and rhizodeposit-derived C was recovered in the order MB-C<C_org<CO₂-C. This showed that rhizodeposits were mainly respired, but since C_org was the second largest pool of the overall balances, they were also stabilized in the soils at least in the short term. It is suggested that the increased SOM mineralization observed in this study (positive priming effects) was probably induced by C exchange processes between the soil matrix and soluble rhizodeposits. Moreover, soluble rhizodeposit-C was detected in MB-C and CO₂-C evolved outside the direct root zone, showing the availability of these C-components in the bulk soil.     

Turnover of soil organic matter and of microbial biomass under C3-C4 vegetation change: Consideration of C fractionation and preferential substrate utilization

Two processes contribute to changes of the δ¹³C signature in soil pools: ¹³C fractionation per se and preferential microbial utilization of various substrates with different δ¹³C signature. These two processes were disentangled by simultaneously tracking δ¹³C in three pools - soil organic matter (SOM), microbial biomass, dissolved organic carbon (DOC) - and in CO₂ efflux during incubation of 1) soil after C₃-C₄ vegetation change, and 2) the reference C₃ soil.The study was done on the Ap horizon of a loamy Gleyic Cambisol developed under C₃ vegetation. Miscanthus giganteus - a perennial C₄ plant - was grown for 12 years, and the δ¹³C signature was used to distinguish between ‘old’ SOM (>12 years) and ‘recent’ Miscanthus-derived C (<12 years). The differences in δ¹³C signature of the three C pools and of CO₂ in the reference C₃ soil were less than 1‰, and only δ¹³C of microbial biomass was significantly different compared to other pools. Nontheless, the neglecting of isotopic fractionation can cause up to 10% of errors in calculations. In contrast to the reference soil, the δ¹³C of all pools in the soil after C₃-C₄ vegetation change was significantly different. Old C contributed only 20% to the microbial biomass but 60% to CO₂. This indicates that most of the old C was decomposed by microorganisms catabolically, without being utilized for growth. Based on δ¹³C changes in DOC, CO₂ and microbial biomass during 54 days of incubation in Miscanthus and reference soils, we concluded that the main process contributing to changes of the δ¹³C signature in soil pools was preferential utilization of recent versus old C (causing an up to 9.1‰ shift in δ¹³C values) and not ¹³C fractionation per se.Based on the δ¹³C changes in SOM, we showed that the estimated turnover time of old SOM increased by two years per year in 9 years after the vegetation change. The relative increase in the turnover rate of recent microbial C was 3 times faster than that of old C indicating preferential utilization of available recent C versus the old C.Combining long-term field observations with soil incubation reveals that the turnover time of C in microbial biomass was 200 times faster than in total SOM. Our study clearly showed that estimating the residence time of easily degradable microbial compounds and biomarkers should be done at time scales reflecting microbial turnover times (days) and not those of bulk SOM turnover (years and decades). This is necessary because the absence of C reutilization is a prerequisite for correct estimation of SOM turnover. We conclude that comparing the δ¹³C signature of linked pools helps calculate the relative turnover of old and recent pools.     

Soil organic matter dynamics in lands continuously cultivated with maize and soybean in Heilongjiang Province,Northeast China: Estimation from natural 13C abundance

Abstract

Northeast China is the main production area of maize and soybean in China. In the present study, the rates of decomposition and replacement of soil organic carbon (SOC) were estimated using the soil inventory collected since 1991 from long-term maize and soybean cultivation plots in Heilongjiang Province, Northeast China, to evaluate the sustainability of the present cultivation system. The total carbon (C) content in soil was stable without any significant changes in the plots (approximately 28.5 g C kg^?1). The δ¹³C value of soil organic matter under continuous maize cultivation increased linearly with an annual increment of 0.07 from ?23.9 in 1991, which indicated that approximately 13% of the initial SOC was decomposed during the 13-year period of maize cultivation, with a half-life of 65 years. Slow decomposition of SOC was considered to result from the low annual mean temperature (1.5°C) and long freezing period (170–180 days year^?1) in the study area. In contrast, the amount of organic C derived from maize increased in the soil with a very slow annual increment of 0.17 g C kg^?1, probably because of the removal of all the plant residues from the plots. Based on the soil organic matter dynamics observed in the study plots, intentional recycling/maintenance of plant residues was proposed as a way of increasing soil fertility in maize or soybean cultivation.     

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.     

Seasonal dynamics of leaf- and root-derived C in arctic tundra mesocosms

We investigated contributions of leaf litter, root litter and root-derived organic material to tundra soil carbon (C) storage and transformations. ¹⁴C-labeled materials were incubated for 32 weeks in moist tussock tundra soil cores under controlled climate conditions in growth chambers, which simulated arctic fall, winter, spring and summer temperatures and photoperiods. In addition, we tested whether the presence of living plants altered litter and soil organic matter (SOM) decomposition by planting shoots of the sedge Eriophorum vaginatum in half of the cores. Our results suggest that root litter accounted for the greatest C input and storage in these tundra soils, while leaf litter was rapidly decomposed and much of the C lost to respiration. We observed transformations of ¹⁴C between fractions even when total C appeared unchanged, allowing us to elucidate sources and sinks of C used by soil microorganisms. Initial sources of C included both water soluble (WS) and acid-soluble (AS) fractions, primarily comprised of carbohydrates and cellulose, respectively. The acid-insoluble (AIS) fraction appeared to be a sink for C when conditions were favorable for plant growth. However, decreases in ¹⁴C activity from the AIS fraction between the fall and spring harvests in all treatments indicated that microorganisms consumed recalcitrant C compounds when soil temperatures were below 0 °C. In planted leaf litter cores and in both planted and unplanted SOM cores, the greatest amounts of ¹⁴C at the end of the experiment were found in the AIS fraction, suggesting a high rate of humification or accumulation of decay-resistant plant tissues. In unplanted leaf litter cores and planted and unplanted root litter cores most of the ¹⁴C remaining at the end of the experiment was in the AS fraction suggesting less extensive humification of leaf and root detritus. Overall, the presence of living plants stimulated decomposition of leaf litter by creating favorable conditions for microbial activity at the soil surface. In contrast, plants appeared to inhibit decomposition of root litter and SOM, perhaps because of microbial preferences for newer, more labile inputs from live roots.     

In-situ warming does not change soil priming effect induced by glucose addition in a temperate forest

Priming effect (PE) on soil organic carbon (SOC) decomposition caused by the addition of organic carbon (C) is an important ecological process in regulating soil C cycle. Additionally, most priming studies are confined to laboratory trials, while the assessment of soil PE under field conditions with variable weather conditions is scarce. This study assessed the direct effects of glucose addition and in-situ warming on the extent of PE under field conditions in a temperate forest. We evaluated soil PE using ¹³C-glucose labelling, a simple and novel technique, based on the Keeling plot method. Glucose addition significantly enhanced native SOC decomposition and induced strong PE. However, the effect of in-situ warming on the extent of PE was not significant. This study confirms the importance of PE in regulating SOC turnover under field conditions.     

Carbon fluxes in soil food webs of increasing complexity revealed by C labelling and C natural abundance

Soil food webs are mainly based on three primary carbon (C) sources: root exudates, litter, and recalcitrant soil organic matter (SOM). These C sources vary in their availability and accessibility to soil organisms, which could lead to different pathways in soil food webs. The presence of three C isotopes (¹²C, ¹³C and ¹⁴C) offers an unique opportunity to investigate all three C sources simultaneously. In a microcosm experiment we studied the effect of food web complexity on the utilization of the three carbon sources. We choose an incomplete three factorial design with (i) living plants, (ii) litter and (iii) food web complexity. The most complex food web consisted of autochthonous microorganisms, nematodes, collembola, predatory mites, endogeic and anecic earthworms. We traced C from all three sources in soil, in CO₂ efflux and in individual organism groups by using maize grown on soil developed under C₃ vegetation and application of ¹⁴C labelled ryegrass shoots as a litter layer. The presence of living plants had a much greater effect on C pathways than food web complexity. Litter decomposition, measured as ¹⁴CO₂ efflux, was decreased in the presence of living plants from 71% to 33%. However, living plants increased the incorporation of litter C into microbial biomass and arrested carbon in the litter layer and in the upper soil layer. The only significant effect of food web complexity was on the litter C distribution in the soil layers. In treatments with fungivorous microarthropods (Collembola) the incorporation of litter carbon into mineral soil was reduced. Root exudates as C source were passed through rhizosphere microorganisms to the predator level (at least to the third trophic level). We conclude that living plants strongly affected C flows, directly by being a source of additional C, and indirectly by modifying the existing C flows within the food web including CO₂ efflux from the soil and litter decomposition.     

Comments on the paper by Kemmitt et al. (2008) ‘Mineralization of native soil organic matter is not regulated by the size, activity or composition of the soil microbial biomass - A new perspective’ [Soil Biology & Biochemistry 40, 61-73]: The biology of the Regulatory Gate

Kemmitt et al. (Kemmitt, S.J., Lanyon, C.V., Waite, I.S., Wen, Q., Addiscott, T.M., Bird, N.R.A., O'Donnell, A.G., Brookes, P.C., 2008. Mineralization of native soil organic matter is not regulated by the size, activity or composition of the soil microbial biomass - a new perspective. Soil Biology & Biochemistry 40, 61-73) recently proposed the “Regulatory Gate” hypothesis, which states that decomposition of soil organic matter (SOM) is regulated solely by abiotic factors. Without studying the mechanisms of such regulation, Kemmitt with coauthors challenged the classical Winogradsky theory of soil microbiology and questioned the concept of autochtonous and zymogenous microbial populations. In this letter, we revive the significance of microbial activity for SOM decomposition especially for the short-term (hours to weeks) processes and show that the “Regulatory Gate” is (micro)biologically driven.We explain the results of the three experiments in Kemmitt et al. (2008) from a microbiological point of view and suggest that SOM decomposition is mainly regulated by exoenzymes. We criticize the abiotic Regulatory Gate hypothesis based on bottleneck processes and pools limiting the SOM decomposition rate, comparison of constant and changing environmental conditions, as well as the connection between community structure and functions. We explain the results of Kemmitt et al. (2008) according to the properties of soil microbial community: functional redundancy and inconsistency between the excessive (but largely inactive) pool of total microbial biomass and the real mineralization activity. Finally, we suggest that to gain new perspectives on SOM decomposition and many other biochemical processes, future studies should focus on hot spots of (micro)biological activity (i.e., the rhizosphere, drillosphere, detritosphere, biopores, etc.) rather than on the bulk soil.     

Compound-specific δC and δH analyses of plant and soil organic matter: A preliminary assessment of the effects of vegetation change on ecosystem hydrology

Here we present δ¹³C and δ²H data of long-chained, even-numbered (C₂₇-C₃₁) n-alkanes from C3 (trees) and C4 (grasses) plants and from the corresponding soils from a grassland-woodland vegetation sequence in central Queensland, Australia. Our data show that δ¹³C values of the C4 grassland species were heavier relative to those of C3 tree species from the woodland (Acacia leaves) and woody grassland (Atalaya leaves). However, n-alkanes from the C4 grasses had lighter δ²H values relative to the Acacia leaves, but showed no significant difference in δ²H values when compared with C3 Atalaya leaves. These results differ from those of previous studies, showing that C4 grasses had heavier δ²H values relative to C3 grasses and trees. Those observations have been explained by C4 plants accessing the more evaporation-influenced and isotopically heavier surface water and tree roots sourcing deeper, isotopically lighter soil water (“Two-layered soil-water system”). By comparison, our data suggest that ecosystem changes (vegetation “thickening”) can significantly alter the soil hydrological characteristics. This is shown by the heavier δ²H values in the woodland soil compared with lighter δ²H values in the grassland soil, implying that the recent vegetation change (increased tree biomass) in the woodland had altered soil hydrological conditions. Estimated δ²H values of the source-water for vegetation in the grassland and woodland showed that both trees and grasses in open settings accessed water with lighter δ²H values (avg. −46‰) compared with water accessed by trees in the woodland vegetation (avg. −7‰). These data suggest that in semi-arid environments the “two-layer” soil water concept might not apply. Furthermore, our data indicate that compound-specific δ²H and δ¹³C analyses of n-alkanes from soil organic matter can be used to successfully differentiate between water sources of different vegetation types (grasses versus trees) in natural ecosystems.     

Factors controlling decomposition of soil organic matter in fallow systems of the high tropical Andes: A field simulation approach using C- and N-labelled plant material

N-rich (C:N=27) and N-poor (C:N=130) wheat straw, labelled with ¹⁴C and ¹⁵N, was incubated for 2 yr in two major ecosystems of the upper elevation belt of cultivation in the high Andes: the moist Paramo (precipitation=1329 mm, altitude=3400 m asl, Andes of Merida, Venezuela) and the dry Puna (precipitation=370 mm, altitude=3800 m asl, Central Altiplano, Bolivia). The experiment was installed in young (2 yr) and old (7 yr) fallow plots. The following soil analyses were performed at nine sampling occasions: soil moisture, total-¹⁴C and -¹⁵N, and Microbial Biomass (MB)-¹⁴C and -¹⁵N. The measured data were fitted by the MOMOS-6 model (a process based model, with five compartments: labile and stable plant material, MB, and labile (HL) and stable humus (HS)) coupled with the SAHEL model (soil moisture prediction) using daily measured and/or predicted meteorological data. The aim was to understand how (1) the climatic conditions, (2) the quality of plant material, (3) the fallow age and (4) the soil properties affect the cycling of C and N within the soil organic matter system.The fallow age (2 and 7 yr) did not affect the measured data or the model predictions, indicating that in these systems the decomposition potential is not affected by fallow length. During the short initial active decomposition phase, the labile plant material was quickly exhausted, enabling a build up of MB and of HL. During the low activity phase, that covered 4/5 of the time of exposure, the MB size decreased slowly and the HL pool was progressively exhausted as it was reused by the MB as substrate. The HL compartment was directly or indirectly the major source for the inorganic ¹⁵N production. If the C:N ratio of the added plant material increased, the model predicted (1) a reduction of the decomposition rates of the plant material (essentially the stable plant material) and (2) an increased mortality of the MB which increased the production of HL (microbial cadavers and metabolites). Thus the essential effect of the slower decomposition due to the N-poor plant material was a higher accumulation of C and N in the HL and its slower recycling by the MB during the low activity phase. The labelling experiment allows to understand the higher soil native organic matter content in Paramo soils compared to Puna. The large sequestration of organic matter generally observed in the Paramo soils can be explained by two abiotic factors: the unfavourable soil microstructure and the accumulation of free aluminium linked to the climatic and acid soil conditions, inhibiting the microbial activity physically and chemically.