Cutin and suberin biomarkers as tracers for the turnover of shoot and root derived organic matter along a chronosequence of Ecuadorian pasture soils

Forest‐to‐pasture conversion has been reported to increase soil organic matter (SOM) in mineral topsoils in the tropical mountain rainforest region of south Ecuador, with subsequent decreases following pasture abandonment. Until now the mechanisms behind these changes have not been fully understood. To elucidate their varied preservation patterns, we analysed root‐ and shoot‐derived organic matter and assessed their contribution to the formation of SOM in topsoils (0–5 cm) on a chronosequence of pastures (Setaria sphacelata (Schumach.); C₄) established after slash and burn of the natural forest (diverse C₃ plant species) and an abandoned pasture site invaded by bracken fern (Pteridium arachnoideum (Kaulf.) Maxon.; C₃). Cutin and suberin biomarkers of the two plant species (grass and bracken) and of forest litter were identified after saponification and their contribution to SOM was studied by compound‐specific stable carbon isotope analyses. Our results showed specific root and shoot biomarkers for the two plant species and for forest litter, which often did not correspond to the classification of root‐versus shoot‐specific monomers reported in the literature. This illustrates the importance of direct biomarker determination rather than using results from studies with different plants. Shoot‐ as well as root‐derived OM of forest and grass origin contributed to the stable SOM pool with decadal turnover times. Forest‐derived monomers contributed more to the stable SOM pool compared with grass‐derived monomers. ω‐hydroxy carboxylic acids and α,ω‐alkanedioic acids of forest origin may have been stabilized in these tropical soils by bonding to soil minerals. Rapid degradation of grass‐derived lipids from the same compound classes suggests a saturation of the mineral binding capacity. In pasture soils, the accumulation of SOM was mainly driven by large inputs of root OM. The accumulated SOM during pasture use is, however, lost rapidly after abandonment.     

Microbial use of lignite compared to recent plant litter as substrates in reclaimed coal mine soils

In the Lusatian mining district, rehabilitated mine soils contain substantial amounts of lignite in addition to recent carbon derived from plant litter. The aim of this study was to examine the importance of the two organic matter types as substrates for soil microbial biomass in mine soils containing organic matter with a contrasting degree of humification. Samples were taken from the lignite-containing overburden material, from a mine soil under 14-year-old black pine (Pinus nigra) and from a mine soil under 37-year-old red oak (Quercus rubra). Overburden material was ameliorated with alkaline ash and incubated in an identical manner as the 14-year-old and 37-year-old mine soils for 16 months. Carbon mineralisation was monitored throughout. After 0, 3, 6, 12 and 16 months, samples were removed and analysed for chemical parameters and for microbial biomass. In addition, ¹⁴C activity measurements in bulk soil and microbial biomass were used to estimate their lignite content.Despite the high content of organic carbon in lignite-rich overburden material, low contents of microbial biomass were recorded. Ash-amelioration led to high pH values in the overburden material, resulting in high concentrations of dissolved organic carbon most likely derived from lignite. Development of the microbial community was subsequently stimulated by presence of an easily available carbon source. In older mine soils, larger amounts of microbial biomass are most likely related to the presence of recent organic matter. Radiocarbon analysis of the microbial biomass extracted from the 14-year-old mine soil indicated higher lignite carbon contribution than recorded for microbial biomass of the 37-year-old mine soil. The highest concentration of lignite C present in microbial biomass as indicated by the C_mic/C_org ratio was, however, observed in the ameliorated overburden material. Therefore, we conclude that the importance of lignite as a carbon source for micro-organisms decreases when recent organic matter is present in the older stages of mine soil development.     

Transfer of N fixed by a legume tree to the associated grass in a tropical silvopastoral system

Below-ground transfer of nitrogen (N) fixed by legume trees to associated non-N₂-fixing crops has received little attention in agroforestry, although the importance of below-ground interactions is shown in other ecosystems. We used ¹⁵N natural abundance to estimate N transfer from the legume tree Gliricidia sepium (Jacq.) Kunth ex Walp. to C₄ grass Dichanthium aristatum (Poir.) C.E. Hubb. in a silvopastoral system, where N was recycled exclusively by below-ground processes and N₂ fixation by G. sepium was the sole N input to the system. Finding a suitable reference plant, a grass without contact with tree roots or litter, was problematic because tree roots invaded adjacent grass monocrop plots and soil isotopic signature in soil below distant grass monocrops differed significantly from the agroforestry plots. Thus, we used grass cultivated under greenhouse conditions in pots filled with agroforestry soil as the reference. A model of soil ¹⁵N fractionation during N mineralization was developed for testing the reliability of that estimate. Experimental and theoretical results indicated that 9 months after greenhouse transplanting, the percentage of fixed N in the grass decreased from 35% to <1%, due to N export in cut grass and dilution of fixed N with N taken up from the soil. The effect of soil ¹⁵N fractionation on the estimate of the reference value was negligible. This indicates that potted grass is a suitable reference N transfer studies using ¹⁵N natural abundance. About one third of N in field-grown grass was of atmospheric origin in agroforestry plots and in adjacent D. aristatum grassland invaded by G. sepium roots. The concentration of fixed N was correlated with fine root density of G. sepium but not with soil isotopic signature. This suggests a direct N transfer from trees to grass, e.g. via root exudates or common mycorrhizal networks.     

Turnover of organic matter in a Miscanthus field: effect of time in Miscanthus cultivation and inorganic nitrogen supply

To accurately predict the potential environmental benefits of energy crops, the sequestration of carbon in soil needs to be quantified. The aim of this study was to investigate the mineralisation rate of the perennial C₄ grass Miscanthus giganteus and Miscanthus-derived soil organic matter under contrasting nitrogen supply. Soils were collected from sites where Miscanthus had been grown for 11 and 18 years, respectively, and where a C₃-grass (Lolium spp.) had been grown for 7 years. The soils were incubated for 4 months at two levels of soil inorganic nitrogen with or without dead root material of Miscanthus.Addition of root material (residues) increased carbon mineralisation of indigenous organic matter when no nitrogen was added. Added inorganic nitrogen decreased carbon mineralisation in all soils. Nitrogen addition did not affect carbon mineralisation of the residues. Using the ¹³C fraction to calculate the proportion of respiratory CO₂ derived from Miscanthus showed that nitrogen addition decreased carbon mineralisation in soils, but it did not affect carbon mineralisation of the residues. Nitrogen mineralisation was highest in the C₃ grass soil without added residues. Nitrification decreased pH, especially in the treatments where nitrogen was added. The Miscanthus-derived organic matter is at least as stable as C₃ grassland-derived organic matter. Furthermore, the turnover time of the organic matter increases with time under Miscanthus cultivation.The CENTURY soil organic matter sub-model was used to simulate the organic matter decomposition in the experiment. Carbon mineralisation was accurately simulated but there were unexplained discrepancies in the simulation of the δ¹³C in the respiration from the treatment with residues. The δ¹³C in respiration did not decrease with time as predicted, indicating that lignin accumulation did not influence the measurements.     

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.     

Effects of addition of maize litter and earthworms on C mineralization and aggregate formation in single and mixed soils differing in soil organic carbon and clay content

C mineralization and aggregate stability directly depend upon organic matter and clay content, and both processes are influenced by the activity of microorganisms and soil fauna. However, quantitative data are scarce. To achieve a gradient in C and clay content, a topsoil was mixed with a subsoil. Single soils and the soil mixture were amended with 1.0 mg maize litter C g soil⁻¹ with and without endogeic earthworms (Aporrectodea caliginosa). The differently treated soils were incubated for 49 days at 15 °C and 40% water holding capacity. Cumulative C mineralization, microbial biomass, ergosterol content and aggregate fractions were investigated and litter derived C in bulk soil and aggregates were determined using isotope analyses. Results from the soil mixture were compared with the calculated mean values of the two single soils. Mixing of soil horizons differing in carbon and clay content stimulated C mineralization of added maize residues as well as of soil organic matter. Mixing also increased contents of macro-aggregate C and decreased contents of micro-aggregate C. Although A. caliginosa had a stimulating effect on C mineralization in all soils, decomposition of added litter by A. caliginosa was higher in the subsoil, whereas A. caliginosa decreased litter decomposition in the soil mixture and the topsoil. Litter derived C in macro-aggregates was higher with A. caliginosa than with litter only. In the C poor subsoil amended with litter, A. caliginosa stimulated the microbial community as indicated by the increase in microbial biomass. Furthermore, the decrease of ergosterol in the earthworm treated soils showed the influence of A. caliginosa on the microbial community, by reducing saprotrophic fungi. Overall, our data suggest both a decrease of saprotrophic fungi by selective grazing, burrowing and casting activity as well as a stimulation of the microbial community by A. caliginosa.     

Natural abundance δN and δC of DNA extracted from soil

We report the first simultaneous measurements of δ¹⁵N and δ¹³C of DNA extracted from surface soils. The isotopic composition of DNA differed significantly among nine different soils. The δ¹³C and δ¹⁵N of DNA was correlated with δ¹³C and δ¹⁵N of soil, respectively, suggesting that the isotopic composition of DNA is strongly influenced by the isotopic composition of soil organic matter. However, in all samples DNA was enriched in ¹³C relative to soil, indicating microorganisms fractionated C during assimilation or preferentially used ¹³C enriched substrates. Enrichment of DNA in ¹⁵N relative to soil was not consistently observed, but there were significant differences between δ¹⁵N of DNA and δ¹⁵N of soil for three different sites, suggesting microorganisms are fractionating N or preferentially using N substrates at different rates across these contrasting ecosystems. There was a strong linear correlation between δ¹⁵N of DNA and δ¹⁵N of the microbial biomass, which indicated DNA was depleted in ¹⁵N relative to the microbial biomass by approximately 3.4‰. Our results show that accurate and precise isotopic measurements of C and N in DNA extracted from the soil are feasible, and that these analyses may provide powerful tools for elucidating C and N cycling processes through soil microorganisms.     

The contribution of soil organic matter fractions to carbon and nitrogen mineralization and microbial community size and structure

The aims of this study were to: (i) assess the impact of hay and fertilizer application on organic matter (OM) fractions (dissolved organic matter (DOM), light fraction organic matter (LFOM, <1.0 g cm⁻³), heavy fraction OM (HFOM, <1.7 g cm⁻³)), carbon (C) and nitrogen (N) cycling processes and microbial community size and structure, and (ii) quantify the role of OM fractions to C and N cycling. Soil was collected in 2001 from a field experiment to which grass hay (1996) and/or fertilizer (1995 and 1999) had previously been applied. DOM-C (P<0.05) and DOM-N (P=0.07) were significantly higher in control and fertilized soil than hay and hay+fertilized soil. LFOM and HFOM C and N contents and C/N ratios were significantly (P<0.05) higher in hay+fertilized and hay amended soil than in control and fertilized soil. Potentially mineralizable-N (PMN), microbial biomass-C (MB-C), microbial biomass-N (MB-N) and microbial respiration (CO₂) were not affected by fertilizer and/or hay application. Gross N mineralization (Gross Min) and gross nitrification (Gross Nit) rates were significantly (P<0.05) higher in fertilized, hay, hay+fertilized soil than control soil. However, there was no significant difference between treatments in gross N immobilization rates. Results reported here highlight the importance of a labile fraction of the DOM pool to N and C cycling as its removal significantly (P<0.05) reduced PMN, MB-N, Gross Min and Gross Nit compared with whole soil in most or all treatments. In soil where DOM+LFOM were removed PMN was significantly (P<0.05) lower, but MB-C, Gross Min and Gross Nit was significantly (P<0.05) higher than in DOM removed soil. This suggests that LFOM plays an important role as a sink for mineral-N. Total soil phospholipid fatty acid (PLFA) concentration was significantly (P<0.05) higher in hay amended than control, fertilized and hay+fertilized soil. Principal components analysis was able to clearly discriminate between control, fertilized, hay+fertilized and hay amended soil. Soil amended with hay or fertilizer had a microbial community structure which differed from that of the control or hay+fertilized soils. Redundancy analysis with Monte Carlo permutation tests revealed that PLFA profiles were strongly correlated to differences in Gross Min, Gross Nit, MB-N, MB-C, MB-C/N ratio, total soil C and total soil C/N ratio. The results of this research suggest that changes in microbial structure are related to aspects of soil C and N pools and cycling.     

Quantification and bioavailability of scyllo-inositol hexakisphosphate in pasture soils

The recent identification of scyllo-inositol hexakisphosphate in alkaline soil extracts by solution ³¹P NMR spectroscopy allowed us to investigate this compound in soils by re-analyzing spectra from two previously published studies. Concentrations of scyllo-inositol hexakisphosphate in 29 temperate pasture soils from England and Wales ranged between 11 and 130 mg P kg⁻¹ soil and accounted for between 4 and 15% of the soil organic phosphorus. The ratio of scyllo-inositol hexakisphosphate to myo-inositol hexakisphosphate ranged between 0.29 and 0.79. In a 10 month pot experiment with six grassland soils from New Zealand, growth of pine seedlings (Pinus radiata D. Don) decreased scyllo-inositol hexakisphosphate concentrations by between 10 and 46%. Growth of ryegrass (Lolium perenne L.) decreased scyllo-inositol hexakisphosphate in three low-nutrient soils by 5-21%, but increased it in three other soils by 11-16%. We conclude that scyllo-inositol hexakisphosphate is an important component of soil organic phosphorus with potential ecological significance.     

Microbial community utilization of added carbon substrates in response to long-term carbon input manipulation

The chemical composition and quantity of plant inputs to soil are primary factors controlling the size and structure of the soil microbial community. Little is known about how changes in the composition of the soil microbial community affect decomposition rates and other ecosystem functions. This study examined the degradation of universally ¹³C-labeled glucose, glutamate, oxalate, and phenol in soil from an old-growth Douglas-fir (Pseudotsuga menziesii)—western hemlock (Tsuga heterophylla) forest in the Oregon Cascades that has experienced 7 y of chronic C input manipulation. The soils used in this experiment were part of a larger Detritus Input and Removal Treatment experiment and have received normal C inputs (control), doubled wood inputs, or root and litter input exclusion (no inputs). Soil from the doubled wood treatment had a higher fungal:bacterial ratio, and soil from the no inputs treatment had a lower fungal:bacterial ratio, than the control soil. Differences in the utilization of the compounds added to the field-manipulated soils were assessed by following the ¹³C tracer into microbial biomass and respiration. In addition, ¹³C-phospholipid fatty acids (PLFA) analysis was used to examine differential microbial utilization of the added substrates. Glucose and glutamate were metabolized similarly in soils of all three litter treatments. In contrast, the microbial community in the double wood soil respired more added phenol and oxalate, whereas microbes in the no inputs soil respired less added phenol and oxalate, than the control soil. Phenol was incorporated primarily into fungal PLFA, especially in soil of the double wood treatment. The addition of all four substrates led to enhanced degradation of soil organic matter (priming) in soils of all three litter treatments, and was greater following the addition of phenol and oxalate as compared to glucose and glutamate. Priming was greater in the no inputs soil as compared to the control or doubled wood soils. These results demonstrate that altering plant inputs to soil can lead to changes in microbial utilization of C compounds. It appears that many of these changes are the result of alteration in the size and composition of the microbial community.     

Changes in enzyme activities and microbial biomass after “in situ” remediation of a heavy metal-contaminated soil

Microbial properties such as microbial biomass carbon (MBC), arylsulfatase, β-glucosidase and dehydrogenase activities, and microbial heterotrophic potential, together with several chemical properties such as pH, CaCl₂ soluble heavy metal concentrations, total organic carbon and hydrosoluble carbon were measured to evaluate changes in soil quality, after “in situ” remediation of a heavy metal-contaminated soil from the Aznalcóllar mine accident (Southern Spain, 1998). The experiment was carried out using containers, filled with soil from the affected area. Four organic amendments (a municipal waste compost, a biosolid compost, a leonardite and a litter) and an inorganic amendment (sugarbeet lime) were mixed with the top soil at the rate of 100 Mg ha⁻¹. Unamended soil was used as control. Agrostis stolonifera L. was sown in the containers. The soil was sampled twice: one month and six months after amendment application. In general, these amendments improved the soil chemical properties: soil pH, total organic carbon and hydrosoluble carbon increased in the amended soils, while soluble heavy metal concentrations diminished. At the same time, higher MBC, enzyme activities and maximum rate of glucose mineralization values were found in the organically amended soils. Plant cover was also important in restoring the soil chemical and microbial properties in all the soils, but mainly in those that were not amended organically. As a rule, remediation measures improved soil quality in the contaminated soils.     

Role of soil drying in nitrogen mineralization and microbial community function in semi-arid grasslands of north-west Australia

We examined effects of wetting and then progressive drying on nitrogen (N) mineralization rates and microbial community composition, biomass and activity of soils from spinifex (Triodia R. Br.) grasslands of the semi-arid Pilbara region of northern Australia. We compared soils under and between spinifex hummocks and also examined impacts of fire history on soils over a 28 d laboratory incubation. Soil water potentials were initially adjusted to −100 kPa and monitored as soils dried. We estimated N mineralization by measuring changes in amounts of nitrate (NO₃⁻-N) and ammonium (NH₄⁺-N) over time and with change in soil water potential. Microbial activity was assessed by amounts of CO₂ respired. Phospholipid fatty acid (PLFA) analyses were used to characterize shifts in microbial community composition during soil drying. Net N mineralized under hummocks was twice that of open spaces between hummocks and mineralization rates followed first-order kinetics. An initial N mineralization flush following re-wetting accounted for more than 90% of the total amount of N mineralized during the incubation. Initial microbial biomass under hummocks was twice that of open areas between hummocks, but after 28 d microbial biomass was<2 μ g⁻¹ ninhydrin N regardless of position. Respiration of CO₂ from soils under hummocks was more than double that of soils from between hummocks. N mineralization, microbial biomass and microbial activity were negligible once soils had dried to −1000 kPa. Microbial community composition was also significantly different between 0 and 28 d of the incubation but was not influenced by burning treatment or position. Regression analysis showed that soil water potential, microbial biomass N, NO₃⁻-N, % C and δ¹⁵N all explained significant proportions of the variance in microbial community composition when modelled individually. However, sequential multiple regression analysis determined only microbial biomass was significant in explaining variance of microbial community compositions. Nitrogen mineralization rates and microbial biomass did not differ between burned and unburned sites suggesting that any effects of fire are mostly short-lived. We conclude that the highly labile nature of much of soil organic N in these semi-arid grasslands provides a ready substrate for N mineralization. However, process rates are likely to be primarily limited by the amount of substrate available as well as water availability and less so by substrate quality or microbial community composition.     

A mass-balance approach to measuring microbial uptake and pools of phosphorus in nutrient-amended soils

Soil microbial biomass P is usually determined through fumigation-extraction (FE), in which partially extractable P from lysed biomass is converted to biomass P using a conversion factor (K_p). Estimation of K_p has been usually based on cultured microorganisms, which may not adequately represent the soil microbial community in either nutrient-poor or in altered carbon and nutrient conditions following fertilisation. We report an alternative approach in which changes in microbial P storage are determined as the residual in a mass balance of extractable P before and after incubation. This approach was applied in three low-fertility sandy soils of southwestern Australia, to determine microbial P immobilisation during 5-day incubations in response to the amendment by 2.323 mg C g⁻¹, 100 μg N g⁻¹ and 20 μg P g⁻¹. The net P immobilisation during the amended incubations determined to be 18.1, 14.1 and 16.3 μg P g⁻¹ in the three soils, accounting for 70.6-90.5% of P added through amendment. Such estimates do not rely on fumigation and K_p values, but for comparison with the FE method we estimated ‘nominal’ K_p values to be 0.20-0.31 for the soils under the amended conditions. Our results showed that microbial P immobilisation was a dominant process regulating P concentration in soil water following the CNP amendment. The mass-balance approach provides information not only about changes in the microbial P compartment, but also about other major P-pools and their fluxes in regulating soil-water P concentrations under substrate- and nutrient-amended conditions.     

Decomposer animals (Lumbricidae, Collembola) and organic matter distribution affect the performance of Lolium perenne (Poaceae) and Trifolium repens (Fabaceae)

Decomposer animals stimulate plant growth by indirect effects such as increasing nutrient availability or by modifying microbial communities in the rhizosphere. In grasslands, the spatial distribution of organic matter (OM) rich in nutrients depends on agricultural practice and the bioturbation activities of large detritivores, such as earthworms. We hypothesized that plants of different functional groups with contrasting nutrient uptake and resource allocation strategies differentially benefit from sites in soil with OM accumulation and the presence of decomposer animals. In a greenhouse experiment we investigated effects of spatial distribution of ¹⁵N-labelled grass litter, earthworms and collembola on a simple grassland community consisting of Lolium perenne (grass) and Trifolium repens (legume). Litter aggregates (compared to homogeneous litter distribution) increased total shoot biomass, root biomass and ¹⁵N uptake by the plants. Earthworms and collembola did not affect total N uptake of T. repens; however, the presence of both increased ¹⁵N uptake by T. repens and L. perenne. Earthworms increased shoot biomass of T. repens 1.11-fold and that of L. perenne 2.50 fold. Biomass of L. perenne was at a maximum in the presence of earthworms, collembola and with litter concentrated in a single aggregate. Shoot biomass of T. repens increased in the presence of collembola, with L. perenne generally responding opposingly. The results indicate that the composition of the decomposer community and the distribution of OM in soil affect plant competition and therefore plant community composition.     

C and N natural abundance of the soil microbial biomass

Stable isotope analysis is a powerful tool in the study of soil organic matter formation. It is often observed that more decomposed soil organic matter is ¹³C, and especially ¹⁵N-enriched relative to fresh litter and recent organic matter. We investigated whether this shift in isotope composition relates to the isotope composition of the microbial biomass, an important source for soil organic matter. We developed a new approach to determine the natural abundance C and N isotope composition of the microbial biomass across a broad range of soil types, vegetation, and climates. We found consistently that the soil microbial biomass was ¹⁵N-enriched relative to the total (3.2 ‰) and extractable N pools (3.7 ‰), and ¹³C-enriched relative to the extractable C pool (2.5 ‰). The microbial biomass was also ¹³C-enriched relative to total C for soils that exhibited a C3-plant signature (1.6 ‰), but ¹³C-depleted for soils with a C4 signature (−1.1 ‰). The latter was probably associated with an increase of annual C3 forbs in C4 grasslands after an extreme drought. These findings are in agreement with the proposed contribution of microbial products to the stabilized soil organic matter and may help explain the shift in isotope composition during soil organic matter formation.     

Rhizosphere activity, grass species and N availability effects on the soil C and N cycles

We examined whether grass species and soil nitrogen (N) availability could enhance Carbon (C) and N turnover during root litter decay in grassland. Three species with increasing competitiveness (Festuca ovina, Dactylis glomerata and Lolium perenne) were grown at two N fertiliser levels in an undisturbed grassland soil, in which soil organic fractions derived for the last 9 years from Lolium root litter which was ¹³C-depleted. During the subsequent experimental year, the C turnover was calculated using the respective δ¹³C values of the old and new C in the root phytomass, in two Particulate Organic Matter (POM) fractions above 200 μm and in the lightest part of the aggregated soil fraction between 50 and 200 μm. Soil N availability was monitored during the regrowth periods with ion exchange resins (IER). The C decay rates of each particle size fraction were calculated with a simple mechanistic model of C dynamics. The N mineralisation immobilisation turnover (MIT) was characterised by dilution of ¹⁵N-labelled fertiliser in the N harvestThe C:N ratio and the residence time of C in the fractions decreased with particle size. The presence of a grass rhizosphere increased the decay rate of old C. Accumulation of new C in particle size fractions increased with species competitiveness and with N supply. Species competitiveness increased C turnover in the aggregated fraction, as a result of greater accumulation of new C and faster decay of old C. Fertiliser N increased N turnover and C mineralisation in the SOM. Species competitiveness decreased soil -N exchanged with the IER and increased dissolved organic C (DOC) content. The nature of the current rhizosphere is thus an important factor driving C and N transformations of the old root litter, in relation with grass species strategy. Plant competitiveness may stimulate the C and N turnover in the more evolved SOM fractions in a similar way to the mineral N supply.     

Carbon dioxide emissions of soils under pure and mixed stands of beech and spruce, affected by decomposing foliage litter mixtures

Soil respiration is the largest terrestrial source of CO₂ to the atmosphere. In forests, roughly half of the soil respiration is autotrophic (mainly root respiration) while the remainder is heterotrophic, originating from decomposition of soil organic matter. Decomposition is an important process for cycling of nutrients in forest ecosystems. Hence, tree species induced changes may have a great impact on atmospheric CO₂ concentrations. Since studies on the combined effects of beech-spruce mixtures are very rare, we firstly measured CO₂ emission rates in three adjacent stands of pure spruce (Picea abies), mixed spruce-beech and pure beech (Fagus sylvatica) on three base-rich sites (Flysch) and three base-poor sites (Molasse; yielding a total of 18 stands) during two summer periods using the closed chamber method. CO₂ emissions were higher on the well-aerated sandy soils on Molasse than on the clayey soils on Flysch, characterized by frequent water logging. Mean CO₂ effluxes increased from spruce (41) over the mixed (55) to the beech (59) stands on Molasse, while tree species effects were lower on Flysch (30-35, mixed > beech = spruce; all data in mg CO₂-C m⁻² h⁻¹). Secondly, we studied decomposition after fourfold litter manipulations at the 6 mixed species stands: the O_i - and O_e horizons were removed and replaced by additions of beech -, spruce - and mixed litter of the adjacent pure stands of known chemical quality and one zero addition (blank) in open rings (20 cm inner diameter), which were covered with meshes to exclude fresh litter fall. Mass loss within two years amounted to 61-68% on Flysch and 36-44% on Molasse, indicating non-additive mixed species effects (mixed litter showed highest mass loss). However, base cation release showed a linear response, increasing from the spruce - over the mixed - to the beech litter. The differences in N release (immobilization) resulted in a characteristic converging trend in C/N ratios for all litter compositions on both bedrocks during decomposition. In the summers 2006 and 2007 we measured CO₂ efflux from these manipulated areas (a closed chamber fits exactly over such a ring) as field indicator of the microbial activity. Net fluxes (subtracting the so-called blank values) are considered an indicator of litter induced changes only and increased on both bedrocks from the spruce - over the mixed - to the beech litter. According to these measurements, decomposing litter contributed between 22-32% (Flysch) and 11-28% (Molasse) to total soil respiration, strengthening its role within the global carbon cycle.     

Effects of annual grass senescence on NH4 and N-glycine uptake by blue oak (Quercus douglasii) seedlings and soil microorganisms in California oak woodland

Annual grasses are stronger competitors for available soil N than blue oak seedlings and soil microorganisms. However, little is known about the dynamics of N competition during annual grass senescence. We conducted a field experiment in a California oak woodland to study effects of annual grass senescence on N uptake by grasses, blue oak seedlings and soil microorganisms. Labeled N was applied at the beginning of April, May and of June in the form of ¹⁵NH₄⁺ or ¹⁵N-glycine. Plants and soils were harvested after 5 days (¹⁵NH₄⁺ and ¹⁵N-glycine treatments) and after 26 days (¹⁵NH₄⁺ treatment only). We evaluated effects of N form, season and labeling period on N competition among oak seedlings, annual grasses and soil microorganisms. N forms did not affect competition among grasses, oak seedlings and soil microorganisms, but more ¹⁵N was incorporated into the soil organic N pool in the ¹⁵N-glycine treatments than in the ¹⁵NH₄⁺ treatments. There were no seasonal (May vs June) effects on ¹⁵N recovery in blue oak seedlings and soil microorganisms. Plant samples from April harvest were lost. In June, when grasses were senescing, more ¹⁵N was found in the soil inorganic pool than in May. Extremely dry soils in June may have limited inorganic N availability to oak seedlings and soil microorganisms. After 26-day labeling period, ¹⁵N recovery in blue oak seedlings and the soil organic N pool significantly increased, while ¹⁵N recovery in both the soil microbial and inorganic N pools decreased compared to the 5-day labeling period. Although blue oak seedling biomass changed little from early May to late June, N concentrations in oak roots increased 53%. In contrast, annual grass biomass peaked in May, and then decreased rapidly. Our results suggest that blue oak seedlings and annual grasses have different temporal competitive abilities. Blue oak seedlings appear to have a long-term strategy for N competition. Blue oaks take up N slowly but steadily, increasing N uptake from 5 to 26 days. This extended time period has a greater positive effect on N uptake than does reduced grass uptake caused by senescence.     

Electron donors and acceptors influence anaerobic soil organic matter mineralization in tidal marshes

Anaerobic decomposition in wetland soils is carried out by several interacting microbial processes that influence carbon storage and greenhouse gas emissions. To understand the role of wetlands in the global carbon cycle, it is critical to understand how differences in both electron donor (i.e., organic carbon) and terminal electron acceptor (TEA) availability influence anaerobic mineralization of soil organic matter. In this study we manipulated electron donors and acceptors to examine how these factors influence total rates of carbon mineralization and the pathways of microbial respiration (e.g., sulfate reduction versus methanogenesis). Using a field-based reciprocal transplant of soils from brackish and freshwater tidal marshes, in conjunction with laboratory amendments of TEAs, we examined how rates of organic carbon mineralization changed when soils with different carbon contents were exposed to different TEAs. Total mineralization (the sum of CO₂ + CH₄ produced) on a per gram soil basis was greater in the brackish marsh soils, which had higher soil organic matter content; however, on a per gram carbon basis, mineralization was greater in the freshwater soils, suggesting that the quality of carbon inputs from the freshwater plants was higher. Overall anaerobic metabolism was higher for both soil types incubated at the brackish site where SO₄²⁻ was the dominant TEA. When soils were amended with TEAs in the laboratory, more thermodynamically favorable respiration pathways typically resulted in greater organic matter mineralization (Fe(III) respiration > SO₄²⁻ reduction > methanogenesis). These results suggest that both electron donors and acceptors play important roles in regulating anaerobic microbial mineralization of soil organic matter.     

Relative importance of substrate type and previous soil management in synthesis of microbial biomass and substrate mineralization

Our aim was to determine whether the soil microbial biomass, which has developed naturally over many years in a given ecosystem, is specially adapted to metabolize the plant‐derived substrate C of the ecosystem within which it developed or whether the nature of recently added substrate is the more important factor. To examine this, soils from three sites in close proximity (woodland, grassland and arable from the Broadbalk Experiment at Rothamsted Research, Harpenden, UK) were each amended with air‐dried wheat straw (Triticum aestivum), ryegrass leaves (Lolium perenne) or woodland leaf litter (mainly Quercus robur and Fagus sylvatica) in a fully replicated 3 × 3 factorial laboratory experiment. The initial mineralization rates (evolved CO₂‐C) were determined during the first 6.5 hours and again, together with the amount of microbial biomass synthesized (microbial biomass C), at 7, 14, 21, 30 and 49 days of incubation. The hourly rate of CO₂‐C production during the first 6.5 hours was slowest following leaf litter addition, while the added grass gave the fastest rates of CO₂‐C evolution both within and between soils. Ryegrass addition to the arable soil led to approximately four times more CO₂‐C being evolved than when it was added to the woodland soil, at an overall rate in the arable soils of 41 μg C g^?1 soil hour^?1. In each soil, the net amounts of CO₂‐C produced were in the order grass > straw > leaf litter. In each case, the amount produced by the added leaf litter was significantly less (P < 0.05) than either the added grass or straw. Overall, the trend was for much slower rates of mineralization of all substrates in the woodland soil than in either the arable or grassland soils. During 49 days of incubation in the woodland and grassland soils, the net total amounts of CO₂‐C evolved differed significantly (P < 0.01), with grass > straw > leaf litter, respectively. In the arable soil, the amounts of CO₂‐C evolved from added grass and straw were significantly larger (P < 0.01) than from the leaf litter treatment. Our findings indicated that the amounts of CO₂‐C evolved were not related to soil management or to the size of the original biomass but to the substrate type. The amount of biomass C synthesized was also in the order grass > straw > leaf litter, at all stages of incubation in the woodland and grassland soil. In the arable soil, the same effect was observed up to 14 days, and for the rest of the incubation the biomass C synthesized was in the order grass > straw > leaf litter. Up to three times more biomass C was synthesized from the added grass than from the other substrates in all soils throughout the incubation. The maximum biomass synthesis efficiency was obtained with grass (7% of added C). Overall, the woodland soil was most efficient at synthesizing biomass C and the arable soil the least. We conclude that substrate type was the overriding factor that determined the amount of new soil microbial biomass synthesized. Mineralization of substrate C by soil microorganisms was also influenced mainly by substrate type and less by soil management or size of original biomass.