Roots from beech (Fagus sylvatica L.) and ash (Fraxinus excelsior L.) differentially affect soil microorganisms and carbon dynamics

Knowledge about the influence of living roots on decomposition processes in soil is scarce but is needed to understand carbon dynamics in soil. We investigated the effect of dominant deciduous tree species of the Central European forest vegetation, European beech (Fagus sylvatica L.) and European ash (Fraxinus excelsior L.), on soil biota and carbon dynamics differentiating between root- and leaf litter-mediated effects. The influence of beech and ash seedlings on carbon and nitrogen flow was investigated using leaf litter enriched in ¹³C and ¹⁵N in double split-root rhizotrons planted with beech and ash seedlings as well as a mixture of both tree species and a control without plants. Stable isotope and compound-specific fatty acid analysis (¹³C-PLFA) were used to follow the incorporation of stable isotopes into microorganisms, soil animals and plants. Further, the bacterial community composition was analyzed using pyrosequencing of 16S rRNA gene amplicons. Although beech root biomass was significantly lower than that of ash only beech significantly decreased soil carbon and nitrogen concentrations after 475 days of incubation. In addition, beech significantly decreased microbial carbon use efficiency as indicated by higher specific respiration. Low soil pH probably increased specific respiration of bacteria suggesting that rhizodeposits of beech roots induced increased microbial respiration and therefore carbon loss from soil. Compared to beech δ¹³C and δ¹⁵N signatures of gamasid mites in ash rhizotrons were significantly higher indicating higher amounts of litter-derived carbon and nitrogen to reach higher trophic levels. Similar δ¹³C signatures of bacteria and fine roots indicate that mainly bacteria incorporated root-derived carbon in beech rhizotrons. The results suggest that beech and ash differentially impact soil processes with beech more strongly affecting the belowground system via root exudates and associated changes in rhizosphere microorganisms and carbon dynamics than ash.     

Effects of organic chemical quality and mineral nitrogen addition on lignin and holocellulose decomposition of beech leaf litter by Xylaria sp.

Mycobiota and chemical composition of bleached and non-bleached portions were studied on leaf litter of beech (Fagus crenata Blume). By surface sterilization method, two xylariaceous species Xylaria sp. and Geniculosporium sp.1 were dominantly isolated in both portions. Frequency of occurrence of Xylaria sp. was significantly higher in the bleached portion than in the non-bleached portion. In the bleached portion, lignin concentration was lower than in the non-bleached portion, indicating that Xylaria sp. and Geniculosporium sp.1 took part in lignocellulose decomposition in the study site. Effects of organic chemical quality of litters and exogenous mineral nitrogen (NH₄ and NO₃) addition were then investigated on in vitro lignin decomposition by Xylaria sp. Weight loss of lignin was significantly related to lignocellulose index (LCI) for four litter types tested. In NH₄ and NO₃ addition treatments, lignin decomposition was completely and partially suppressed, respectively. Xylaria sp. produced bleaching spots on beech leaf litter in vitro in which lignin concentration was lower than in the non-bleached portion. These results suggest that heterogeneous distribution of carbon and nitrogen resources may control lignin decomposition on the litter by the fungus.     

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.     

Litter quality effects on soil carbon and nitrogen dynamics in temperate alley cropping systems

Microcosm and litterbag experiments were conducted to determine the effects of litter quality, soil properties and microclimate differences on soil carbon (C) and nitrogen (N) mineralization in alley cropping systems. Bulk soils were collected from 0 to 20 cm depth at three sites: a 21-year old pecan (Carya illinoinensis)/bluegrass (Poa trivialis) intercrop (Pecan site) in north-central Missouri, a 12-year old silver maple (Acer saccharinum)/soybean (Glycine max)–maize (Zea mays) rotation (Maple site) in northeastern Missouri and a restored prairie site (MDC site) in southwestern Missouri. Seven tree and crop litters with varying composition were collected, including pecan, silver maple, chestnut and walnut leaf litter (tree litter) and maize, soybean and bluegrass residues (crop litter). Aerobic microcosm incubations were maintained at 25 °C and a soil water potential of −47 kPa. Unamended MDC soil mineralized 24 and 18% more CO₂ than the Pecan and Maple soils, respectively. Soil amended with crop litter mineralized on average 32% more CO₂ than when amended with tree litter. Net N mineralization from soybean litter was 40 mg kg⁻¹, while all other litter immobilized N for various durations. A double pool and a single pool model best described C and N mineralization from amended soils, respectively. Cumulative CO₂ mineralized, labile C fraction (C₁) and potentially mineralizable C (C₀) were correlated to litter total N and lignin contents and to (lignin + polyphenol):N ratio. In the field, bluegrass litter decomposed and released N twice as fast as pecan leaf litter. Soybean, maize and silver maple litter released 84, 75 and 63% of initial N, respectively, 308 days after field placement, while no differences in mass loss was observed among the three litter materials. At the Maple site, mass and N remaining, 308 days after field placement was lower at the middle of the alley, corresponding to higher soil temperature and water content. No differences in mass loss and N release patterns were observed at the Pecan site. Microclimate and litter quality effects can lead to differences in nutrient availability in alley cropping systems.     

Effects of acid rain on leaf-litter decomposition in a beech forest on calcareous soil

Summary The effects of simulated acid rain on litter decomposition in a calcareous soil (pH_{H 2 O} 5.8) were studied. Litterbags (45 m and 1 mm mesh size) containing freshly fallen beech leaf litter were exposed to different concentrations of acid in a beech forest on limestone (Göttinger Wald. Germany) for 1 year. Loss of C, the ash content, and CO₂–C production were measured at the end of the experiment. Further tests measured the ability of the litter-colonizing microflora to metabolize ¹⁴C-labelled beech leaf litter and hyphae. The simulated acid rain strongly reduced CO₂–C and ¹⁴CO₂–C production in the litter. This depression in production was very strong when the input of protons was 1.5 times greater than the normal acid deposition, but comparatively low when the input was 32 times greater. acid deposition may thus cause a very strong accumulation of primary and secondary C compounds in the litter layer of base-rich soils, even with a moderate increase in proton input. The presence of mesofauna significantly reduced the ability of the acid rain to inhibit C mineralization. The ash content to the 1-mm litterbags indicated that this was largely due to transport of base-rich mineral soil into the litter.     

Partitioning of carbon and nitrogen during decomposition of 13C15N‐labeled beech and ash leaf litter

The aim of this study was to determine the influence of leaf‐litter type (i.e., European beech—Fagus sylvatica L. and European ash—Fraxinus excelsior L.) and leaf‐litter mixture on the partitioning of leaf‐litter C and N between the O horizon, the topsoil, the soil microbial biomass, and the CO₂ emission during decomposition. In a mature beech stand of Hainich National Park, Thuringia, Germany, undisturbed soil cores (?? 24 cm) were transferred to plastic cylinders and the original leaf litter was either replaced by ¹³C¹⁵N‐labeled beech or ash leaf litter, or leaf‐litter‐mixture treatments in which only one of the two leaf‐litter types was labeled. Leaf‐litter‐derived CO₂‐C flux was measured every second week over a period of one year. Partitioning of leaf‐litter C and N to the soil and microbial biomass was measured 5 and 10 months after the start of the experiment. Ash leaf litter decomposed faster than beech leaf litter. The decomposition rate was negatively related to initial leaf‐litter lignin and positively to initial Ca concentrations. The mixture of both leaf‐litter types led to enhanced decomposition of ash leaf litter. However, it did not affect beech leaf‐litter decomposition. After 5 and 10 months of in situ incubation, recoveries of leaf‐litter‐derived C and N in the O horizon (7%–20% and 9%–35%, respectively) were higher than in the mineral soil (1%–5% and 3%–8%, respectively) showing no leaf‐litter‐type or leaf‐litter‐mixture effect. Partitioning of leaf‐litter‐derived C and N to microbial biomass in the upper mineral soil (< 1% of total leaf‐litter C and 2%–3% of total leaf‐litter N) did not differ between beech and ash. The results show that short‐term partitioning of leaf‐litter C and N to the soil after 10 months was similar for ash and beech leaf litter under standardized field conditions, even though mineralization was faster for ash leaf litter than for beech leaf litter.     

Soil nutrient supply and biomass production in a mixed forest on a skeleton‐rich soil and an adjacent beech forest

In the natural forest communities of Central Europe, beech (Fagus sylvatica L.) predominates in the tree layer over a wide range of soil conditions. An exception with respect to the dominance of beech are skeleton‐rich soils such as screes where up to 10 broad‐leaved trees co‐exist. In such a Tilia‐Fagus‐Fraxinus‐Acer‐Ulmus forest and an adjacent mono‐specific beech forest we compared (1) soil nutrient pools and net nitrogen mineralization rates, (2) leaf nutrient levels, and (3) leaf litter production and stem increment rates in order to evaluate the relationship between soil conditions and tree species composition. In the mixed forest only a small quantity of fine earth was present (35 g l^—1) which was distributed in patches between basalt stones; whereas a significantly higher (P < 0.05) soil quantity (182 g l^—1) was found in the beech forest. In the soil patches of the mixed forest C and N concentrations and also concentrations of exchangeable nutrients (K, Ca, Mg) were significantly higher than in the beech forest. Net N mineralization rates on soil dry weight basis in the mixed forest exceeded those in the beech forest by a factor of 2.6. Due to differences in fine earth and stone contents, the volume related soil K pool and the N mineralization rate were lower in the mixed forest (52 kg N ha^—1 yr^—1, 0—10 cm depth) than in the beech forest (105 kg N ha^—1 yr^—1). The leaf N and K concentrations of the beech trees did not differ significantly between the stands, which suggests that plant nutrition was not impaired. In the mixed forest leaf litter fall (11 %) and the increment rate of stem basal area (52 %) were lower than in the beech forest. Thus, compared with the adjacent beech forest, the mixed forest stand was characterized by a low volume of patchy distributed nutrient‐rich soil, a lower volume related K pool and N mineralization rate, and low rates of stem increment. Together with other factors such as water availability these patterns may contribute to an explanation of the diverse tree species composition on Central European screes.     

Effects of Aporrectodea caliginosa (Savigny) on nitrogen mobilization and decomposition of elevated‐CO2 Charlock mustard litter

This study was conducted to improve our understanding of how earthworms and microorganisms interact in the decomposition of litter of low quality (high C : N ratio) grown under elevated atmospheric [CO₂]. A microcosm approach was used to investigate the influence of endogeic earthworm (Aporrectodea caliginosa Savigny) activity on the decomposition of senescent Charlock mustard (Sinapis arvensis L.) litter produced under ambient and elevated [CO₂]. Earthworms and microorganisms were exposed to litter which had changed in quality (C : N ratio) while growing under elevated [CO₂]. After 50 d of incubation in microcosms, C mineralization (CO₂ production) in the treatment with elevated‐[CO₂] litter was significantly lower in comparison to the ambient‐[CO₂] litter treatment. The input of Charlock mustard litter into the soil generally induced N immobilization and reduced N₂O‐emission rates from soil. Earthworm activity enhanced CO₂ production, but there was no relationship to litter quality. Although earthworm biomass was not affected by the lower quality of the elevated‐[CO₂] litter, soil microbial biomass (C_mic, N_mic) was significantly decreased. Earthworms reduced C_mic and fungal biomass, the latter only in treatments without litter. Our study clearly showed that A. caliginosa used the litter grown under different [CO₂] independent of its quality and that their effect on the litter‐decomposition process was also independent of litter quality. Soil microorganisms were shown to negatively react to small changes in Charlock mustard litter quality; therefore we expect that microbially mediated C and N cycling may change under future atmospheric [CO₂].     

Changes in the earthworm community of an acidophilous lowland beech forest during a stand rotation

The study examines humus profile development during a chronosequence consisting of four beech forest growth stages, and particularly the role of some components of soil fauna (lumbricid oligochaetes) on humus morphogenesis. An experimental site was set up in a lowland beech forest (Fougères state forest, eastern Brittany, France) to carry out a chronosequence analysis. This lowland beech forest is acidophilous, managed as an even-aged beech stand. The dominant tree species is beech (Fagus sylvatica L.). Samples were taken twice a year, from autumn 1997 to spring 2000 inclusive. Earthworms were caught after watering with formalin solutions. Only five species were found: three epigeic (litter-dwellers): Dendrobaena octaedra, D. rubida tenuis, Eisenia eiseni; one epi-anecic (litter/soil-dweller): Lumbricus rubellus; one endogeic (soil-dweller): Allolobophora caliginosa. D. octaedra is dominant in the four plots at densities ranging from 41 ind·m^–2 (88 % of total earthworm population) to 12.4 ind·m^–2 (99 % of total). Species richness and abundance are low in accordance with a moder humus form typical of acid soil conditions. Results are discussed according to plot heterogeneity, stand age, seasonal variations and functional diversity.     

Origin of the nitrogen assimilated by soil fauna living in decomposing beech litter

We investigated the nitrogen source for main taxa of soil fauna in two beech forests of contrasted humus type using ¹⁵N-labelled beech litter and ¹⁵N analysis of soil fauna. ¹⁵N-labelled beech litter was deposited on the topsoil in December 2000 in four stands of different ages at Leinefelde (Germany) with mull humus and in one mature stand at Sorø (Denmark) with moder humus. The fate of the tracer isotope was measured in litter and soil, as well as in the soil fauna, and for each taxa, we calculated the proportion of N in the animal derived from the labelled substrate. Of the original N contained in the litter, 20-41% was lost after 9 months at Leinefelde, and only 10% at Sorø. This loss was counterbalanced by the incorporation of 24-31% external N at Leinefelde, and 31% at Sorø, partly originating from fungal colonisation of the added litter. The proportion of N assimilated from the labelled litter by the different soil animals varied in relation to their mobility and feeding preferences. Large and mobile soil animals, especially predators, derived on average less ¹⁵N because they were also able to feed outside the labelled litter boxes. Detritivores assimilated at most 15% of their nitrogen content at Leinefelde and 11% at Sorø from the decomposing labelled litter. The most labelled taxa at Leinefelde were small fungivorous and coprophagous species, mainly isotomid Collembola such as Isotomiella and Folsomia. At Sorø, best labelled taxa were saprophagous species such as Enchytraeidae, Glomeridae and Phthiracaroidea. These low rates of ¹⁵N assimilation indicate that fresh litter is not directly the main N source for soil animals. The results obtained suggest that soil fauna fed preferentially upon microorganisms colonising the litter at Leinefelde (mull) and from litter itself at Sorø (moder).     

The occurrence of terrestrial algae on two beech sites on a Cambisol

The occurrence of terrestrial algae on two beech sites was determined qualitatively and quantitatively in the litter, organic and mineral layer. Both sites are beech stands of different age and the soil type is a Cambisol. On the site with young beech the maximal value determined was 1.0 · 10⁸ algae g^?1 dw in the litter layer. With increasing soil depth the number of algae decreased on both sites, and at the young beech stand site no algae were found in the mineral soil. Chlorophyceae were the dominant algae group and at the young beech stand Bacillariophyceae were present.     

The influence of slug (Arion rufus) mucus and cast material addition on microbial biomass,respiration, and nutrient cycling in beech leaf litter

We investigated the effects of slug (Arion rufus L.) mucus and cast material on litter decomposition, nutrient mobilization, and microbial activity in two laboratory experiments: (1) Slug mucus and cast material was added to beech leaf litter (Fagus sylvatica L.), and leaching of N and P and CO₂ production in microcosm systems were measured during 77 days of incubation; (2) mucus was added to beech leaf litter, and basal respiration, microbial biomass (substrate-induced respiration), specific respiration (qO₂), microbial growth ability after C, CN, CP, and CNP amendment, and lag time (time between CNP addition and start of exponential increase in respiration rate) were measured during 120 days of incubation. Leaching of N and P from beech leaf litter was significantly increased in treatments with mucus or faecal material of A. rufus. Following day 3, slug mucus increased nitrification processes. Mucus addition to beech leaf litter also increased basal respiration and microbial biomass significantly. In contrast, specific respiration was not significantly affected by mucus addition, and generally declined until day 60 but then increased until day 120. Nutrient amendments indicated that between days 1 and 30, N was available for microbial growth in litter with mucus but not in control litter. Generally, the lag time in beech leaf litter with added mucus was shorter than in control litter. Lag times generally increased with age, indicating dominance of slow-growing microbial populations at later stages as a consequence of depletion of easily available C resources and nutrients. We conclude that C, N, and P cycling is accelerated by slug activity.     

Forest Litters as a Link in the Carbon Cycle in Coniferous–Broadleaved Forests of the Southern Far East of Russia

Forest litter as a component of the carbon cycle in pine–broadleaved forests of different ages was characterized. Field studies of the forest site of the Primorskaya State Agricultural Academy in the south of the Sikhote Alin Range continued for three years. Multiple sampling of forest litter and plant litter fall and measurements of the CO₂ emission from the litter and underlying soil horizons were performed on test plots. The maximum litter pool (14.44 ± 0.86 t/ha) was found in the mature stand of Pinus koraiensis; the minimum litter pool (11.52 ± 0.65 t/ha), in the 80-year-old stand. The carbon stock in the litters amounted to 3.7% of the phytomass carbon. The rate of carbon turnover in the forest litters was relatively low in comparison with that in other regions: the ratio of carbon pools in the litter horizons and annual plant falloff reached 3.5. The winter season provided about 10–25% of the annual plant litter fall. The data obtained in this study describe a part of the carbon cycle and contribute to our understanding of the ecosystem function of climate regulation by valuable forest massifs in the south of the Far East of Russia. In this region, the decomposition of forest litters generates the CO₂ flux amounting up to 16% of the total CO₂ emission from the soil.     

Effects of tree species and topography on soil chemistry, litter quality, and decomposition in Northeast Turkey

Leaf litters from beech (Fagus orientalis Lipsky.) and oak (Quercus robur L.), and needle litters from fir (Abies nordmanniana Spach.) and pine (Pinus sylvestris L.) trees were collected from north-facing site and south-facing site and at three slope positions (top, middle and bottom) on each aspect that varied in soil chemical characteristics (soil pH, cation exchange capacity and base saturation). The litters were analysed for initial total carbon, nitrogen, acid detergent fibre, lignin and cellulose concentrations. Nitrogen, acid detergent fibre and lignin concentrations and carbon:nitrogen and lignin:nitrogen ratios varied significantly within and between species according to soil chemical characteristics on aspects and slope positions. Litter decomposition was studied in the field using the litterbag technique. The litters were placed on two aspects and at three slopes on each aspect in October 2001, and were sampled every 6-month for 2 years. The main effects of aspect, species and slope position on decomposition rates were all statistically significant. Oak leaf litter showed highest decomposition rates, followed by pine, fir and beech litter, and the litters placed on north-facing site decomposed faster than those on the south-facing site. The litters placed at the top slope position decomposed slower than at those at either the bottom or middle positions. Initial lignin concentrations explained most of the variation in decomposition rates between species, and within species for the aspects and the slope positions, but the explained variance showed differences between aspects and slope positions. This result illustrates the important point that litter quality may define the potential rates of microbial decomposition but these are significantly influenced by the biotic and abiotic environment in which decomposition takes place.     

Competition between invasive earthworms (Amynthas corticis, Megascolecidae) and native North American millipedes (Pseudopolydesmus erasus, Polydesmidae): Effects on carbon cycling and soil structure

Invasive earthworms can have significant impacts on C dynamics through their feeding, burrowing, and casting activities, including the protection of C in microaggregates and alteration of soil respiration. European earthworm invasion is known to affect soil micro- and mesofauna, but little is known about impacts of invasive earthworms on other soil macrofauna. Asian earthworms (Amynthas spp.) are increasingly being reported in the southern Appalachian Mountains in southeastern North America. This region is home to a diverse assemblage of native millipedes, many of which share niches with earthworm species. This situation indicates potential for earthworm-millipede competition in areas subject to Amynthas invasion.In a laboratory microcosm experiment, we used two ¹³C enriched food sources (red oak, Quercus rubra, and eastern hemlock, Tsuga canadensis) to assess food preferences of millipedes (Pseudopolydesmus erasus), to determine the effects of millipedes and earthworms (Amynthas corticis) on soil structure, and to ascertain the nature and extent of the interactions between earthworms and millipedes. Millipedes consumed both litter species and preferred red oak litter over eastern hemlock litter. Mortality and growth of millipedes were not affected by earthworm presence during the course of the experiment, but millipedes assimilated much less litter-derived C when earthworms were present.Fauna and litter treatments had significant effects on soil respiration. Millipedes alone reduced CO₂ efflux from microcosms relative to no fauna controls, whereas earthworms alone and together with millipedes increased respiration, relative to the no fauna treatment. CO₂ derived from fresh litter was repressed by the presence of macrofauna. The presence of red oak litter increased CO₂ efflux considerably, compared to hemlock litter treatments.Millipedes, earthworms, and both together reduced particulate organic matter. Additionally, earthworms created significant shifts in soil aggregates from the 2000-250 and 250-53 μm fractions to the >2000 μm size class. Earthworm-induced soil aggregation was lessened in the 0-2 cm layer in the presence of millipedes. Earthworms translocated litter-derived C to soil throughout the microcosm.Our results suggest that invasion of ecosystems by A. corticis in the southern Appalachian Mountains is unlikely to be limited by litter species and these earthworms are likely to compete directly for food resources with native millipedes. Widespread invasion could cause a net loss of C due to increased respiration rates, but this may be offset by C protected in water-stable soil aggregates.     

Inter- and intra-specific interactions of Lumbricus rubellus (Hoffmeister, 1843) and Octolasion lacteum (Örley, 1881) (Lumbricidae) and the implication for C cycling

Earthworms are important engineering species of many terrestrial ecosystems as they play a significant role in regulating C turnover. The effects of earthworms on moderating C decomposition processes differ across species and with interactions between species, which is not fully understood. We carried out an experiment to study the interactions of Lumbricus rubellus and Octolasion lacteum, and their effects on soil respiration. Laboratory mesocosms were set up using tulip poplar (Liriodendron tulipifera) leaf litter and varying densities of earthworms in single and combined species treatments. CO₂ efflux rate was used as an indicator of C decomposition rates, and measured with CO₂ sensors every five days over one month. L. rubellus induced higher leaf consumption rate and higher CO₂ efflux than O. lacteum; meanwhile O. lacteum grew more than L. rubellus. Both litter consumption rate and growth rate of earthworms decreased with increasing earthworm density. Soil CO₂ efflux increased with increasing earthworm density (from ∼1-2 μg CO₂ g⁻¹ hr⁻¹ with no earthworms to ∼ 4 μg CO₂ g⁻¹ hr⁻¹ with 8 earthworms). Combining the two species had a synergistic effect on leaf litter consumption, and neutralizing effects on soil respiration. The data suggest that the strength of intra- and inter-specific interactions among earthworm ecological groups varies at different absolute and relative densities, leading to altered leaf litter decomposition and C cycling.     

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

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

Litter decomposition and nitrogen mineralization in newly formed gaps in a Danish beech (Fagus sylvatica) forest

Gap formation is suggested as an alternative forest management approach to avoid extreme changes in the N cycle of forest ecosystems caused by traditional management practises. The present study aimed to investigate the effect of gap formation on N availability in beech litter and mineral soil on sites, which experienced only little soil disturbance during tree harvest. N pools, litter decomposition, and N mineralization rates in mineral soil were studied in two gaps (17 and 30 m in diameter) in a 75-year-old managed European beech (Fagus sylvatica L.) forest in Denmark and related to soil temperature (5 cm depth) and soil moisture (15 cm depth). Investigations were carried out during the first 2 years after gap formation in measurement plots located along the north-south transect running through the centre of each gap and into the surrounding forest.An effect of gap size was found only for soil temperatures and litter mass loss: soil temperatures were significantly increased in the northern part of the large gap during the first year after gap formation, and litter mass loss was significantly higher in the smaller gap. All other parameters investigated revealed no effect of gap size. Nitrification, net mineralization, and soil N concentrations tended to be increased in the gaps. Cumulative rates of net mineralization were two fold higher in the gaps during the growing season (June-October), but a statistically significant increase was found only for soil NH₄-N concentrations during this period. Forest floor parameters (C:N ratios, mass loss, N release) were not significantly modified during the first year after gap formation, neither were the total C content nor the C:N ratio in mineral soil at 0-10 cm depth.     

Decomposition of beech leaves (Fagus sylvatica) and spruce needles (Picea abies) in pure and mixed stands of beech and spruce

The decomposition of spruce needles and beech leaves was investigated in a 30- and 120-yr-old beech, spruce and mixed (beech/spruce) forest using 1 mm mesh litterbags. The mass loss, content of C, N and water and microbial biomass, basal respiration and specific respiration of the litter materials were analyzed after exposure for 1.5, 3, 6, 9, 12, 18 and 24 months in the field. Decomposition of both types of litter was faster in beech than in spruce stands and after 24 months loss of C from litter materials was at a maximum in beech stands (>60%) and considerably less in the spruce and mixed stands (ca. 40%). Generally, spruce needles decomposed more rapidly than beech leaves, but the faster decay was not associated with higher N concentrations. Rather, N was accumulated more rapidly in beech leaves. Concomitantly, in beech stands microbial biomass of beech leaves exceeded that of spruce needles indicating that beech leaves consist of more favorable resources for microorganisms than spruce needles. Differences in decomposition between beech leaves and spruce needles were most pronounced in beech stands, intermediate in mixed stands and least pronounced in spruce stands. Decomposition, N content and microbial biomass in litter materials exposed in the 120-yr-old stand consistently exceeded that in the 30-yr-old stand indicating adverse conditions for litter decay in regrowing stands. Generally, mixed stands ranked intermediate between spruce and beech monocultures for most of the variables measured indicating that the adverse conditions for litter decay and microorganisms in spruce forest are effectively counteracted by admixture of beech to spruce monocultures. It is concluded that the accumulation of litter materials in spruce forests is not due to the recalcitrance of spruce needles to decay. Rather, adverse environmental conditions such as high polyphenol contents in the litter layer of spruce stands retard decomposition processes; spruce needles appear to be more sensitive to this retardation than beech leaves.     

Carbon dynamics in a temperate grassland soil after 9 years exposure to elevated CO2 (Swiss FACE)

Elevated pCO₂ increases the net primary production, C/N ratio, and C input to the soil and hence provides opportunities to sequester CO₂-C in soils to mitigate anthropogenic CO₂. The Swiss 9 y grassland FACE (free air carbon-dioxide enrichment) experiment enabled us to explore the potential of elevated pCO₂ (60 Pa), plant species (Lolium perenne L. and Trifolium repens L.) and nitrogen fertilization (140 and 540 kg ha⁻¹ y⁻¹) on carbon sequestration and mineralization by a temperate grassland soil. Use of ¹³C in combination with respired CO₂ enabled the identification of the origins of active fractions of soil organic carbon. Elevated pCO₂ had no significant effect on total soil carbon, and total soil carbon was also independent of plant species and nitrogen fertilization. However, new (FACE-derived depleted ¹³C) input of carbon into the soil in the elevated pCO₂ treatments was dependent on nitrogen fertilization and plant species. New carbon input into the top 15 cm of soil from L. perennne high nitrogen (LPH), L. perenne low nitrogen (LPL) and T. repens low nitrogen (TRL) treatments during the 9 y elevated pCO₂ experiment was 9.3±2.0, 12.1±1.8 and 6.8±2.7 Mg C ha⁻¹, respectively. Fractions of FACE-derived carbon in less protected soil particles >53 μm in size were higher than in <53 μm particles. In addition, elevated pCO₂ increased CO₂ emission over the 118 d incubation by 55, 61 and 13% from undisturbed soil from LPH, LPL and TRL treatments, respectively; but only by 13, 36, and 18%, respectively, from disturbed soil (without roots). Higher input of new carbon led to increased decomposition of older soil organic matter (priming effect), which was driven by the quantity (mainly roots) of newly input carbon (L. perenne) as well as the quality of old soil carbon (e.g. higher recalcitrance in T. repens). Based on these results, the potential of well managed and established temperate grassland soils to sequester carbon under continued increasing concentrations of atmospheric CO₂ appears to be rather limited.