Decomposing ability of interior and surface fungal colonizers of beech leaves with reference to lignin decomposition

Fungi were isolated from interior and surface of beech (Fagus crenata Blume) leaf litter by surface sterilization and washing methods. Species composition differed between the interior and surface of the leaves. Xylaria sp. (anamorph) was a major interior colonizer, while Pestalotiopsis spp. and Trichoderma spp. were predominantly surface colonizers. Ascochyta sp. was isolated from both the interior and surface of leaves. Leaf litter decomposing abilities of all isolated species were assessed by pureculture decomposition tests. Changes in lignin, carbohydrate and polyphenol amounts in the litter were investigated. Percent loss of original weight of sterilized beech leaf litter ranged from 0.24 to 11.16 % in the decomposition test. Interior fungi such as Xylaria sp. (anamorph) had the highest abilities to bleach leaf litter and decompose lignin. Most surface fungi had limited ability to decompose leaf litter and lignin. The difference in the decomposing abilities between the interior and surface fungi is discussed in relation to the difference in organic-chemical compositions between interior and surface of 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.     

Effects of nitrogen addition and litter properties on litter decomposition and enzyme activities of individual fungi

Litter decomposition is an important process of C and N cycling in the soil. Variation in the response of litter decomposition to nitrogen (N) addition (positive, negative or neutral) has been observed in many field studies. However, mechanism about variability in individual fungal species response to N addition has not yet been well demonstrated in the literature. Therefore, the objective of this study was to investigate the effects of N addition and litter chemistry properties on litter decomposition and enzyme activities of individual fungi. Three fungal species (Penicillium, Aspergillus, and Trichoderma) were isolated from a subtropical mixed forest soil. An incubation experiment was conducted using the individual fungi with two types of litter (leaf of Pinus massoniana and needle of Cryptocarya chinensis) and different N addition levels (0, 50 and 100 for N-deficient treatments, and 500 and 1000 μg N for N-excessive treatments). Cumulative CO₂-C, enzyme activities, and lignin and cellulose loss were measured during the incubation period of 60 days. Litter decomposition and enzyme activities significantly varied with the fungal species, while the N addition and litter types greatly affected fungal enzyme activities. The N treatments significantly increased lignin-rich needle decomposition by lignocellulose decomposers (Penicillium and Aspergillus) but did not affect their leaf decomposition. On the contrary, The N treatments stimulated leaf decomposition by cellulolytic species (Trichoderma) but did not affect its needle decomposition. Correlation analysis showed that lignin in the litter was the key component to affect litter decomposition. Activities of N-acetyl-β-glucosaminidase and phenol oxidase were both positively correlated to litter decomposition. The fungi (Penicillium and Aspergillus) with higher production of N-acetyl-β-glucosaminidase showed higher litter decomposition ability. The low N addition levels stimulated Penicillium and Aspergillus litter decomposition, but they still required more N source (e.g., litter N source) to support decomposition. Depressed fungal litter N uptake (lower N-acetyl-β-glucosaminidase activities) only occurred at the highest N addition level. Litter decomposition of Trichoderma depended more on external N and its litter decomposition capability was the lowest among the three species.     

Effects of NH4+ and NO3− on litter and soil organic carbon decomposition in a Chinese fir plantation forest in South China

Soil organic carbon (SOC) dynamics and nutrient availability determine the soil quality and fertility in a Chinese fir plantation forest in subtropical China. Uniformly ¹³C-labeled Chinese fir (Cunninghamia lanceolata) and alder (Alnus cremastogyne) leaf litter with or without 100 mg NH₄⁺ or NO₃⁻ were added to the soil. The purpose was to investigate the influence of N availability on the decomposition of the litter and native SOC. The production of CO₂, the natural abundance of ¹³C–CO₂, and the inorganic N dynamics were monitored. The results showed that Chinese fir (with a high C:N ratio) and alder (with a low C:N ratio) leaf litter caused significant positive priming effects (PEs) of 24% and 42%, respectively, at the end of the experiment (235 d). The PE dynamics showed that positive PE can last for at least 87 d. However, the possible occurrence of a significant negative PE with a sufficient incubation period is difficult to confirm. The application of both NH₄⁺ and NO₃⁻ was found to have a stimulating effect on the decomposition of Chinese fir and alder leaf litter in the early stage (0–15 d) of incubation, but an adverse effect in the late stage. Compared with NO₃⁻, NH₄⁺ caused a greater decrease in the PE induced by both Chinese fir and alder leaf litter. The effects of NH₄⁺ and NO₃⁻ on the PE dynamics had different patterns for different incubation stages. This result may indicate that the stability or recalcitrance of SOC, especially in such plantation forest soils, strongly depends on available leaf litter and application of N to the soil.     

Decomposition of sun and shade leaves from three deciduous tree species,as affected by their chemical composition

Freshly fallen leaf litter from sweet chestnut (Castanea sativa Mill), oak (Quercus robur L.) and beech (Fagus sylvatica L.) trees were classified into sun, intermediate and shade leaf types and analysed for N, acid detergent fibre, holocellulose, and lignin. In addition, the sugar constituents of structural polysaccharides (mainly from hemicelluloses) were determined after trifluoracetic acid (TFA) hydrolysis, and the phenylpropanoid (PPD) derivatives of lignin after alkaline CuO oxidation. The litters were decomposed in laboratory microcosms for 2 years. Decomposition rates were initially rapid and then plateaued, but differences in mass losses for the leaf litter categories, and between the three species, were significant at 6, 12, 18 and 24 months. Mean mass losses after 24 months were 49.6% for chestnut, 40.4% for oak and 26.3% for beech. Mean losses for chestnut, oak and beech litter categories were 48.6%, 38.2% and 24.6%, respectively, for sun leaves, and 51.0%, 44.5% and 28.5%, respectively, for shade leaves. Initial lignin concentrations showed a negative correlation with mass losses over the first 6 months but initial acid detergent fibre was a better predictor of decomposition rates after 24 months. Within species, however, total extractable sugars and PPD concentrations reflected differences in decomposition rates between the different categories of leaf types. The analysis for specific carbohydrates and lignin derivatives improved the resolution of litter quality characterisation but did not explain the observed patterns of decomposition in long-term laboratory incubations. It is suggested that these may be affected by influence of the culture conditions on the composition of fungal communities.     

Impacts of anthropogenic N additions on nitrogen mineralization from plant litter in exotic annual grasslands

Urban regions of southern California receive up to 45 kg N ha^-1 y^-1 from nitrogen (N) deposition. A field decomposition study was done using ¹⁵N-labelled litter of the widespread exotic annual grass Bromus diandrus to determine whether elevated soil N is strictly from N deposition or whether N mineralization rates from litter are also increased under N deposition. Tissue N and lignin concentrations, which are inversely related in field sites with high and low N deposition, determine the rate at which N moves from plant litter to soil and becomes available to plants. The effect of soil N on N movement from litter to soil was tested by placing litter on high and low N soil in a factorial experiment with two levels of litter N and two levels of soil N. The litter quality changes associated with N deposition resulted in faster rates of N cycling from litter to soil. Concentrations of litter-derived N in total N, NH₄⁺, NO₃⁻, microbial N and organic N were all higher from high N/low lignin litter than from low N/high lignin litter. Litter contributed more N to soil NH₄⁺ and microbial N in high N than low N soil. At the end of the study, N mineralized from high N litter on high N soil accounted for 46% of soil NH₄⁺ and 11% of soil NO₃⁻, compared to 35% of soil NH₄⁺ and 6% of soil NO₃⁻ from low N litter on low N soil. The study showed that in high N deposition areas, elevated inorganic soil N concentrations at the end of the summer N deposition season are a result of N mineralized from plant litter as well as from N deposition.     

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.     

Elevated CO<Subscript>2</Subscript> concentration affected pine and oak litter chemistry and the respiration and microbial biomass of soils amended with these litters

Elevated atmospheric CO₂ concentration ([CO₂]) may change litter chemistry which affects litter decomposability. This study investigated respiration and microbial biomass of soils amended with litter of Pinus densiflora (a coniferous species; pine) and Quercus variabilis (a deciduous species; oak) that were grown under different atmospheric [CO₂] and thus had different chemistry. Elevated [CO₂] increased lignin/N through increased lignin concentration and decreased N concentration. The CO₂ emission from the soils amended with litter produced under the same [CO₂] regime was greater for oak than pine litter, confirming that broadleaf litter with lower lignin decomposes faster than needle leaf litter. Within each species, however, soils amended with high lignin/N litter grown under elevated [CO₂] emitted more CO₂ than those with low lignin/N litter grown under ambient [CO₂]. Such contrasting effects of lignin/N on inter- and intra-species variations in litter decomposition should be ascribed to the effects of other litter chemistry variables including nonstructural carbohydrate, calcium and manganese as well as inhibitory effect of N on lignin decomposition. The microbial biomass was also higher in the soils amended with high lignin/N litter than those with low lignin/N litter probably due to low substrate use efficiency of lignin by microbes. Our study suggests that elevated [CO₂] increases lignin/N for both species, but increased lignin/N does not always reduce soil respiration and microbial biomass. Further study investigating a variety of tree species is required for more comprehensive understanding of inter- and intra-species variations of litter decomposition under elevated [CO₂].     

Combined methods for the determination of lignin and cellulose in leaf litter

Precise and specific methods for the simultaneous quantitative determination of lignin and cellulose are discussed in this paper, enabling the monitoring of even slight changes in the content of lignocellulose in dead plant material during decomposition processes. Results from different leaf litter samples as well as a comparison of the lignocellulose content of freshly fallen leaf litter, leached and microbially inoculated leaf litter, and feces of saprophagous soil animals fed on this food source are presented. The suitability of the described methods for investigating decomposition processes is discussed.     

Veränderungen in der stofflichen Zusammensetzung von Buchenstreu und Gerstenstroh beim Abbau unter Laborbedingungen

Chemical changes of beech litter and barley straw during decomposition under laboratory conditions Beech litter and barley straw were incubated at 20°C and 70% of maximum water holding capacity in the presence and absence of artificial “soil” and earthworms (Eisenia fetida). Results show that beech litter biodegradation was enhanced by E. fetida during the first part, but delayed in later stages of the incubation period, as indicated by the changes of ash contents and C-to-N ratios with progressive decomposition. In the long run the organic matter (OM) of beech litter tended to be stabilised through the action of worms. In contrast, for barley straw a more intense biodegradation was observed in the presence of E. fetida throughout the experiment. Almost 80% of litter and straw OM could be identified by means of wet chemical degradation methods (70% polysaccharides, lignin; 10% lipids, protein). The proportion of the exclusively plant-derived constituents cellulose and lignin showed a partly strong decrease with progressive decomposition; simultaneously the contents of also microbially synthesised components such as non-cellulosic polysaccharides and protein increased. Changes of the non-cellulosic polysaccharide-cellulose-quotients (NCQ), lignin-cellulose-quotients (LCQ), and acid-to-aldehyde ratios (ac/al)_v and (ac/al)_s in the residual lignin reflected well the litter decomposition process. At the end of the experiment - irrespective of treatments - the degree of beech litter biodegradation was comparable to that of Of-Oh transitional layers in beech-derived forest humus profiles. In the case of barley straw only NCQ and LCQ, but not (ac/al)_v,s were valuable parameters for the characterisation of the decomposition process.     

Interactions between litter quality, decomposition and soil fertility: a laboratory study

Leaf litters from beech (Fagus sylvatica L.) and oak (Quercus robur L.) trees were collected from mixed, deciduous woodlands growing on three soil types that varied in mineral nutrient concentrations and N mineralisation potential. Litter quality, including %N, %Mn, %P, acid detergent fibre, cellulose, Klason lignin, phenylpropanoid constituents of lignin, hexose and pentose sugar (mainly from hemicelluloses) varied within species according to soil type. However, oak and beech showed the opposite responses to soil nutrient status for most of these variables. The litters were incubated in the laboratory for 12 months (at 18 °C and constant moisture) on beds of forest floor material from two soils of contrasting high nutrient material (HNM) or low nutrient material (LNM) nutrient status to investigate litter quality and substrate interactions. At 4, 8 and 12 months there were significant differences in mass losses from oak and beech litters from all sites, and for each litter type exposed to the HNM and LMN soils. At 12 months mean mass losses were higher for HNM treatment (38.7% oak, 27.8% beech) than for the LNM treatment (30.6% oak, 25.5% beech). However, the beech and oak litters from the different sites consistently responded in opposite ways on the same soil treatment reflecting site-related effects on litter quality. Initial concentration of Klason lignin was the best predictor for mass losses from litter species and litter types. Intra-specific variation in rates of litter decomposition of beech and oak litters from different sites, and differences in their interactions with the two forest floor materials, illustrate the complexities of proximate controls on decomposition that are often masked in system-level studies.     

Effect of earthworm addition on soil nitrogen availability,microbial biomass and litter decomposition in mesocosms

The aim of the study was to determine the effect of adding two tropical earthworm species, Rhinodrilus contortus and Pontoscolex corethrurus, to mesocosms on the availability of mineral N (NH₄ ⁺ and NO₃ ^– concentrations), soil microbial biomass (bio-N), and the decomposition rates of three contrasting leaf litter species, in a glasshouse experiment. The mesocosms were filled with forest soil and covered with a layer of leaf litter differing in nutritional quality: (1) Hevea brasiliensis (C/N=27); (2) Carapa guianensis (C/N=32); (3) Vismia sp., the dominant tree species in the second growth forest (control, C/N= 42); and, (4) a mixture of the former three leaf species, in equal proportions (C/N=34). At the end of the 97-day experiment, the soil mineral N concentrations, bio-N, and leaf litter weight loss were determined. Both earthworm species showed significant effects on the concentrations of soil NO₃ ^– (p<0.01) and NH₄ ⁺ (p<0.05). Bio-N was always greater in the mesocosms with earthworms (especially with R. contortus) and in the mesocosms with leaf litter of H. brasiliensis (6 µg N g^–1 soil), the faster decomposing species, than in the other treatments (0.1–1.6 µg N g^–1). Thus, earthworm activity increased soil mineral-N concentrations, possibly due to the consumption of soil microbial biomass, which can speed turnover and mineralization of microbial tissues. No significant differences in decomposition rate were found between the mesocosms with and without earthworms, suggesting that experiments lasting longer are needed to determine the effect of earthworms on litter decomposition rates.     

Translocation of 13C-labeled leaf or root litter carbon of beech (Fagus sylvatica L.) and ash (Fraxinus excelsior L.) during decomposition – A laboratory incubation experiment

The aim was to quantify medium term litter type and litter mixture effects on the translocation and transformation dynamics of root and leaf litter C during decomposition. Partitioning of ¹³C-labeled root or leaf litter C (beech – Fagus sylvatica L., ash – Fraxinus excelsior L.) to CO₂, water-extractable organic C (WEOC), microbial biomass C (C_MB) and light (LF) and heavy soil fraction (HF) was determined in a laboratory decomposition experiment of 206 days. The proportions of C mineralized from ash leaf (34%) and root litter (29%) were higher than those from beech leaf (24%) and root litter (23%). In mixture with beech, the mineralization of ash leaf litter was enhanced. Mineralization was positively correlated with litter-derived WEOC until day 29. Water-extractable organic C declined with time, until <0.1% of litter C remained in this fraction. Litter-C recovery in C_MB was higher for ash (0.7–1.0%) than for beech (0.2–0.4%). The litter C recovery in HF (4–12%) was positively correlated with that in WEOC (days 9 and 29) and C_MB, but did not differ between treatments. Ash leaf litter mineralization showed different behavior in mixed treatments from pure treatments. Thus, the ability to transfer results from pure to mixed treatments is limited. The litter differed in chemical composition and in mineralization dynamics, but differences in partitioning to HF, WEOC and MB were finally of minor importance.     

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.     

Feeding preferences of native terrestrial isopod species (Oniscoidea,Isopoda) for native and introduced leaf litter

Due to current predictions for Central Europe that forecast higher frequencies of hot and dry summers, Mediterranean drought-tolerant oak species are being evaluated as future forest trees for German forest sites that are becoming increasingly damaged by water deficit. As a result of planting foreign tree species, the leaf litter composition and thus the food resources of native saprophagous macroarthropods will change, possibly altering primary decomposition processes. Therefore, experiments concerning the acceptance and palatability of introduced versus native litter for native isopods were undertaken. Consumption rates of four native isopod species (Porcellio scaber, Oniscus asellus, Trachelipus rathkii, Trachelipus ratzeburgii) were investigated in laboratory choice tests with introduced (Quercus pubescens, Quercus frainetto, Quercus ilex) and comparable native (Fagus sylvatica, Quercus robur) leaf litter. Litter was characterized by measurement of C/N-ratios and lignin content. Although species-specific preferences of isopods could be observed in the experiments, Mediterranean oak litter was consumed by all investigated species. Furthermore, two isopod species even preferred the leaf litter of the introduced Q. ilex. Compared to native beech or oak litter, litter from these introduced tree species thus apparently do not negatively influence the consumption rates of terrestrial isopods. Possible reasons for the determined preferences are discussed.     

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.     

Effects of litter (beech and stinging nettle) and earthworms (Octolasion lacteum) on carbon and nutrient cycling in beech forests on a basalt-limestone gradient: A laboratory experiment

Effects of leaf litter of beech (Fagus sylvatica L.) and stinging nettles (Urtica dioica L.) and of the endogeic earthworm species Octolasion lacteum (Örley) on carbon turnover and nutrient dynamics in soil of three beechwood sites on a basalt hill (Hesse, Germany) were investigated in a laboratory experiment lasting for about 1 year. The sites were located along a gradient from basalt (upper part of the hill) to limestone (lower part of the hill) with an intermediate site in between (transition zone). At the intermediate site U. dioica dominated in the understory whereas at the other sites Mercurialis perennis L. was most abundant. The amount and composition of organic matter was similar in soil of the basalt (carbon content 5.9%, C/N ratio 13.8) and intermediate site (carbon content 5.6%, C/N ratio 14.3) but the soil of the intermediate site produced more CO₂ (in total +17.5%) and more nitrogen (as nitrate) was leached from this soil (in total +55.6%). It is concluded that the soil of the intermediate site contains a large mobile carbon and nitrogen pool and the formation of this pool is ascribed to the input of U. dioica litter. Leaf litter of U. dioica strongly increased NO₃ ^–-N leaching immediately after the litter had been added, whereas nitrogen was immobilized due to addition of beech litter. Despite the very fast initial decomposition of nettle litter, the increase in CO₂ production due to this litter material was only equivalent to 20.1% of the amount of carbon added with the nettle litter; the respective value for beech litter was 34.8%. Earthworms altered the time course of carbon and nitrogen mineralization in each of the treatments. In general, earthworms strongly increased mineralization of nitrogen but this effect was less pronounced in soil of the intermediate site (treatments without litter), which is ascribed to a depleted physically protected nitrogen and carbon pool. In contrast, their effect on the total amount of nitrogen mobilized from nettle litter was small. Earthworms significantly reduced CO₂ production from soil of the intermediate site (treatments without litter) and it is concluded that earthworm activity contributes to the restoration of the depleted physically protected carbon pool at this site.     

甘肃省兴隆山森林主要树种凋落叶分解速率与初始质量的关系

[目的]开展凋落叶分解速率研究,探讨凋落叶分解速率与初始质量的关系,为甘肃省兴隆山森林生态系统物质循环研究提供依据。[方法]采用凋落物分解袋法,以兴隆山青杄、山杨和白桦3种主要树种的凋落叶为研究对象,进行凋落叶分解速率及凋落叶初始质量的研究,明确凋落叶分解速率与初始质量的关系。[结果]青杄中龄林针叶分解速率为0.16,95%分解期为19.08a;青杄近熟林针叶分解速率为0.13,95%分解期为23.70a;山杨和白桦凋落叶分解速率均为0.11,95%分解期分别为28.57a和27.27a;山杨和白桦凋落叶分解速率明显要小于青杄针叶,这很可能是凋落叶分解主场效应和分解袋孔径较小所致。凋落叶分解速率与氮含量呈显著线性正相关,与木质素含量、碳/氮值、木质素/氮值和钾含量呈显著线性负相关,特别是与木质素含量、氮含量和木质素/氮值,相关系数均达0.700 0以上;钾含量、木质素含量、木质素/氮、碳/磷和纤维素含量是影响兴隆山森林凋落叶分解速率的重要指标。[结论]木质素/氮值是影响凋落叶分解速率的关键质量指标,凋落叶初始木质素/氮值越高,分解速率越低。     

Lignin and cellulose degradation and nitrogen dynamics during decomposition of three leaf litter species in a Mediterranean ecosystem

Cellulose and lignin degradation dynamics was monitored during the leaf litter decomposition of three typical species of the Mediterranean area, Cistus incanus L., Myrtus communis L. and Quercus ilex L., using the litter bag method. Total N and its distribution among lignin, cellulose and acid-detergent-soluble fractions were measured and related to the overall decay process. The litter organic substance of Cistus and Myrtus decomposed more rapidly than that of Quercus. The decay constants were 0.47 year⁻¹, 0.75 year⁻¹ and 0.30 year⁻¹ for Cistus, Myrtus and Quercus, respectively. Lignin and cellulose contents were different as were their relative amounts (34 and 18%, 15 and 37%, 37 and 39% of the overall litter organic matter before exposure, for Cistus, Myrtus and Quercus, respectively). Lignin began to decrease after 6 and 8 months of exposure in Cistus and Myrtus, respectively, while it did not change significantly during the entire study period in Quercus. The holocellulose, in contrast, began to decompose in Cistus after 1 year, while in Quercus and Myrtus immediately. Nitrogen was strongly immobilized in all the litters in the early period of decay. Its release began after the first year in Cistus and Myrtus and after 2 years of decomposition in Quercus. These litters still contained about 60, 20 and 90% of the initial nitrogen at the end of the experiment (3 years). Prior to litter exposure nitrogen associated with the lignin fraction was 65, 54 and 37% in Cistus, Myrtus and Quercus, while that associated with the cellulose fraction was 30, 24 and 28%. Although most of the nitrogen was not lost from litters, its distribution among the litter components changed significantly during decomposition. In Cistus and Myrtus the nitrogen associated with lignin began to decrease just 4 months after exposure. In Quercus this process was slowed and after 3 years of decomposition 8% of the nitrogen remained associated with lignin or lignin-like substances. The nitrogen associated with cellulose or cellulose-like substances, in contrast, began to decrease from the beginning of cellulose decomposition in all three species. At the end of the study period most of the nitrogen was not associated to the lignocellulose fraction but to the acid-detergent-soluble substance (87, 88 and 84% of the remaining litter nitrogen).     

Production of lignocellulose-degrading enzymes and degradation of leaf litter by saprotrophic basidiomycetes isolated from a Quercus petraea forest

Due to the production of lignocellulose-degrading enzymes, saprotrophic basidiomycetes can significantly contribute to the turnover of soil organic matter. The production of lignin- and polysaccharide-degrading enzymes and changes of the chemical composition of litter were studied with three isolates from a Quercus petraea forest. These isolates were capable of fresh litter degradation and were identified as Gymnopus sp., Hypholoma fasciculare and Rhodocollybia butyracea. Within 12 weeks of incubation, H. fasciculare decomposed 23%, R. butyracea 32% and Gymnopus sp. 38% of the substrate dry mass. All fungi produced laccase and Mn-peroxidase (MnP) and none of them produced lignin peroxidase or other Mn-independent peroxidases. There was a clear distinction in the enzyme production pattern between R. butyracea or H. fasciculare compared to Gymnopus sp. The two former species caused the fastest mass loss during the initial phase of litter degradation, accompanied by the temporary production of laccase (and MnP in H. fasciculare) and also high production of hydrolytic enzymes that later decreased. In contrast, Gymnopus sp. showed a stable rate of litter mass loss over the whole incubation period with a later onset of ligninolytic enzyme production and a longer lasting production of both lignin and cellulose-degrading enzymes. The activity of endo-cleaving polysaccharide hydrolases in this fungus was relatively low but it produced the most cellobiose hydrolase. All fungi decreased the C/N ratio of the litter from 24 to 15-19 and Gymnopus sp. also caused a substantial decrease in the lignin content. Analytical pyrolysis mass spectrometry of litter decomposed by this fungus showed changes in the litter composition similar to those caused by white-rot fungi during wood decay. These changes were less pronounced in the case of H. fasciculare and R. butyracea. All fungi also changed the mean masses of humic acid and fulvic acid fractions isolated from degraded litter. The humic acid fraction after degradation by all three fungi contained more lignin and less carbohydrates. Compared to the decomposition by saprotrophic basidiomycetes, litter degradation in situ on the site of fungal isolation resulted in the relative enrichment of lignin and differences in lignin composition revealed by analytical pyrolysis. It can most probably be explained by the participation of non-basidiomycetous fungi and bacteria during natural litter decomposition.