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.     

Litter- and ecosystem-driven decomposition under elevated CO2 and enhanced N deposition in a Sphagnum peatland

Peatlands represent massive global C pools and sinks. Carbon accumulation depends on the ratio between net primary production and decomposition, both of which can change under projected increases of atmospheric CO₂ and N deposition. The decomposition of litter is influenced by 1) the quality of the litter, and 2) the microenvironmental conditions in which the litter decomposes. This study aims at experimentally testing the effects of these two drivers in the context of global change. We studied the in situ litter decomposition from three common peatland species (Eriophorum vaginatum, Polytrichum strictum and Sphagnum fallax) collected after one year of litter production under pre-treatment conditions (elevated CO₂: 560 ppm or enhanced N: 3 g m⁻² y⁻¹ NH₄NO₃) and decomposed the following year under treatment conditions (same as pre-treatment). By considering the cross-effects between pre-treatments and treatments, we distinguished between the effects on mass loss of 1) the pre-treatment-induced litter quality and 2) the treatment conditions under which the litters were decomposing. The combination between CO₂ pre-treatment and CO₂ treatment reduced Polytrichum decomposition by −24% and this can be explained by litter quality-driven decomposition changes brought by the pre-treatment. CO₂ pre-treatment reduced Eriophorum litter quality, although this was not sufficient to predict decomposition. The N addition pre-treatment reduced the decomposition of Eriophorum, due to enhanced lignin and soluble phenols concentrations in the initial litter, and reduced litter-driven losses of starch and enhanced litter-driven losses of soluble phenols. While decomposition indices based on initial litter quality provide a broad explanation of quantitative and qualitative decomposition, they can only be taken as first approximations. Indeed, the microbial ATP activity, the litter N loss and resulting litter quality, were strongly altered irrespective of the compounds' initial concentration and by means of processes that occurred independently of the initial litter-qualitative changes. The experimental design was valuable to assess litter- and ecosystem-driven decomposition pathways simultaneously or independently. The ability to separate these two drivers makes it possible to attest the presence of litter-qualitative changes even without any litter biochemical determinations, and shows the screening potential of this approach for future experiments dealing with multiple plant species.     

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.     

Temperature sensitivity of respiration differs among forest floor layers in a Pinus resinosa plantation

Decomposer microorganisms contribute to carbon loss from the forest floor as they metabolize organic substances and respire CO₂. In temperate and boreal forest ecosystems, the temperature of the forest floor can fluctuate significantly on a day-to-night or day-to-day basis. In order to estimate total respiratory CO₂ loss over even relatively short durations, therefore, we need to know the temperature sensitivity (Q₁₀) of microbial respiration. Temperature sensitivity has been calculated for microbes in different soil horizons, soil fractions, and at different depths, but we would suggest that for some forests, other ecologically relative soil portions should be considered to accurately predict the contribution of soil to respiration under warming. The floor of many forests is heterogeneous, consisting of an organic horizon comprising a few more-or-less distinct layers varying in decomposition status. We therefore determined at various measurement temperatures the respiration rates of litter, F-layer, and H-layer collected from a Pinus resinosa plantation, and calculated Q₁₀ values for each layer. Q₁₀ depended on measurement temperature, and was significantly greater in H-layer than in litter or F-layer between 5 and 17 °C. Our results indicate, therefore, that as the temperature of the forest floor rises, the increase in respiration by the H-layer will be disproportionate to the increase by other layers. However, change in respiration by the H-layer associated with change in temperature may contribute minimally or significantly to changes of total forest floor respiration in response to changes in temperature depending on the depth and thickness of the layer in different forest ecosystems.     

Plant canopy effects on litter accumulation and soil microbial biomass in two temperate forests

The objective of this study was to determine whether differences in canopy structure and litter composition affect soil characteristics and microbial activity in oak versus mixed fir-beech stands. Mean litter biomass was greater in mixed fir-beech stands (51.9t ha⁻¹) compared to oak stands (15.7t ha⁻¹). Canopy leaf area was also significantly larger in mixed stands (1.96m² m⁻²) than in oak stands (1.73m² m⁻²). Soil organic carbon (C _org) and moisture were greater in mixed fir-beech stands, probably as a result of increased cover. Soil microbial biomass carbon (C _mic), nitrogen (N _mic), and total soil nitrogen (N _tot) increased slightly in the mixed stand, although this difference was not significant. Overall, mixed stands showed a higher mean C _org/N _tot ratio (22.73) compared to oak stands (16.39), indicating relatively low rate of carbon mineralization. In addition, the percentage of organic C present as C _mic in the surface soil decreased from 3.17% in the oak stand to 2.26% in the mixed stand, suggesting that fir-beech litter may be less suitable as a microbial substrate than oak litter.     

Influence of temperature on CO2 evolution,microbial biomass C and metabolic quotient during the decomposition of two humic forest horizons

The decomposition of the litter layer and the humic mineral horizon from a beech forest site was studied at temperatures of 5, 12, and 22°C for both substrates and additionally at 32°C for beech litter. Weight losses, basal and substrate-induced CO₂ production, and the extractable biomass C were monitored periodically during a 2-year incubation period. Weight losses and microbial activity were controlled by substrate quality and temperature. No significant differences were found between 5 and 12°C in decomposition, biomass C, and the metabolic quotient in the humic mineral horizon. The decay of beech litter and the humic mineral horizon was highest at 22°C but was faster in the litter material by a factor of 2.9 on average. In the glucose-amended samples, the relationship among the CO₂-C fluxes was 1:1:2:3 at temperatures of 5, 12, 22, and 32°C in the litter layer, and 1: 2: 2.4 at 5, 12, and 22°C in the A horizon, respectively. The microbial activity in the humic mineral horizon was only 2–11% of that in the litter layer. The level of biomass C remained constant over 1 year and no significant differences were obtained from the 12 and 22°C treatments in the litter layer.     

Microbial enzyme activities in leaf litter, humus and mineral soil layers of European forests

The rationale of the study was to investigate microbial activity in different soil horizons in European forests. Hence, activities of chitinase and cellulase, microbial biomass carbon (C_mic) and basal respiration were measured in litter, fragmentation, humus and mineral soil layers collected several times from various beech and spruce forests. Sites were selected to form a gradient in N availability. Analyses were also performed on beech litter from a litterbag transplant experiment. Furthermore, microbiological parameters were measured in horizons of beech and spruce chronosequence sites with different stand age in order to investigate the influence of forest rotation, and hence changes in soil organic matter (SOM) dynamics, on microbial activity. Finally in horizons of one beech forest, the seasonal variation of selected microbiological parameters was measured more intensively. β-Glucosaminidase and cellobiohydrolase activities were measured using fluorogenic 4-methylumbelliferyl substrates to estimate chitinase and cellulase activities, respectively. On a spatial scale, chitinase and cellulase activities, C_mic determined by substrate induced respiration, and basal respiration ranged from 144 to 1924 and 6-177 nmol 4-MU g⁻¹ org-C h⁻¹, 8-48 mg C g⁻¹ org-C and 11-149 μg CO₂-C g⁻¹ org-C h⁻¹, respectively; in general values were significantly lower in layers of humus and mineral soil than of litter. Chitinase activity, C_mic and basal respiration from humus and mineral soil layers, together, correlated positively, while none correlated with cellulase activity. Similarly in the litter layer, no correlations were found between the microbiological parameters. On a seasonal scale, a time lag between a burst in basal respiration rate and activities of both enzymes were observed. In general, activities of cellulase and chitinase, C_mic and basal respiration, did not change with stand age, except in the humus layer in the spruce chronosequence, where C_mic decreased with stand age. In the litter layer, cellulase activity was significantly and positively related to the C:N ratio, while only a tendency for chitinase activity was shown, indicating that enzyme activities decreased with increasing N availability. In accordance, the enzyme activities and C_mic decreased significantly with increasing chronic N deposition in the humus layer, while basal respiration only tended to decrease with increasing N deposition. In contrast, enzyme activities in beech litter from litterbags after 2 years of incubation were generally higher at sites with higher N deposition. The results show different layer-specific responses of enzyme activities to changes in N availability, indicating different impacts of N availability on decomposition of SOM and stage of litter decomposition.     

Winter soil CO2 efflux and its contribution to annual soil respiration in different ecosystems of a forest-steppe ecotone, north China

Most soil respiration measurements are conducted during the growing season. In tundra and boreal forest ecosystems, cumulative winter soil CO₂ fluxes are reported to be a significant component of their annual carbon budgets. However, little information on winter soil CO₂ efflux is known from mid-latitude ecosystems. Therefore, comparing measurements of soil respiration taken annually versus during the growing season will improve the accuracy of ecosystem carbon budgets and the response of soil CO₂ efflux to climate changes. In this study we measured winter soil CO₂ efflux and its contribution to annual soil respiration for seven ecosystems (three forests: Pinus sylvestris var. mongolica plantation, Larix principis-rupprechtii plantation and Betula platyphylla forest; two shrubs: Rosa bella and Malus baccata; and two meadow grasslands) in a forest-steppe ecotone, north China. Overall mean winter and growing season soil CO₂ effluxes were 0.15-0.26 μmol m⁻² s⁻¹ and 2.65-4.61 μmol m⁻² s⁻¹, respectively, with significant differences in the growing season among the different ecosystems. Annual Q₁₀ (increased soil respiration rate per 10 °C increase in temperature) was generally higher than the growing season Q₁₀. Soil water content accounted for 84% of the variations in growing season Q₁₀ and soil temperature range explained 88% of the variation in annual Q₁₀. Soil organic carbon density to 30 cm depth was a good surrogate for SR₁₀ (basal soil respiration at a reference temperature of 10 °C). Annual soil CO₂ efflux ranged from 394.76 g C m⁻² to 973.18 g C m⁻² using observed ecosystem-specific response equations between soil respiration and soil temperature. Estimates ranged from 424.90 g C m⁻² to 784.73 g C m⁻² by interpolating measured soil respiration between sampling dates for every day of the year and then computing the sum to obtain the annual value. The contributions of winter soil CO₂ efflux to annual soil respiration were 3.48-7.30% and 4.92-7.83% using interpolated and modeled methods, respectively. Our results indicate that in mid-latitude ecosystems, soil CO₂ efflux continues throughout the winter and winter soil respiration is an important component of annual CO₂ efflux.     

Soil organisms and carbon, nitrogen and phosphorus mineralisation in Norway spruce and mixed Norway spruce – Birch stands

 We examined how soil organisms and C, N and P mineralisation are affected by admixing deciduous tree species, silver birch (Betula pendula) and woollen birch (B. pubescens), in managed Norway spruce (Picea abies) stands. Pure spruce and mixed spruce–birch stands were examined at four sites in southern and central Sweden. Soil macroarthropods and enchytraeids were sampled in litter and soil. In the uppermost 5 cm of soil humus we determined microbial biomass and microbial respiration; we estimated the rate of C, N and P mineralisation under laboratory conditions. The densities of Coleoptera, Diptera and Collembola were larger in mixed stands than in spruce stands. Soil fauna composition differed between mixed and spruce stands (as revealed by redundancy analysis). Staphyliniidae, Elateridae, Cecidiomyidae larvae and Onychiuridae were the families that increased most strongly in mixed stands. There were no differences in microbial biomass and microbial respiration, nor in the C, N and P mineralisation rates, between mixed and spruce stands. However, within mixed stands microbial biomass, microbial activity and C mineralisation were approximately 15% higher under birch trees than under spruce trees. We propose that the presence of birch leaf litter was likely to be the most important factor causing differences in soil fauna composition. Birch may also influence the quality and the decomposition rate of humus in mixed stands. However, when the proportion of birch trees is low, the short-term (decades) effect of this species on decomposition is likely to be small in mixed stands on acid forest soils. Received: 20 February 1998     

A microcosm study on the respiration and weight loss in birch litter and raw humus as influenced by soil fauna

Summary The effect of diverse soil fauna (Collembola, Acari, Enchytraeidae, Nematoda) on decomposition of dead organic matter was studied in microcosms containing (1) birch leaf litter, (2) raw humus of coniferous forest and (3) litter on humus. Total respiration (CO₂ evolution) was monitored weekly, and mass loss, length of fungal hyphae (total and metabolically active) and survival of animal populations were checked at the end of weeks 12 and 21–22 from the start of experiment. Animal populations established themselves well during the incubation. At the end of the experiment some replicates containing litter had microarthropod densities of up to 500 specimens per microcosm, corresponding to a field population of 200 000 m^–2. The soil animals had a positive influence on total respiration in all substrates. By the end of experiment 32.0%, 22.6% and 14.6% more CO₂ had evolved in the presence of animals in litter, litter + humus and humus alone, respectively. There was clear trend towards a higher mass loss in the presence of animals, though it was significant in litter only. Our results showed that a diverse soil animal community enhances the activity of soil microbes, and may thereby accelerate decomposition in raw coniferous forest soil.