Reduction of decomposition rates of scots pine needle litter due to heavy-metal pollution

Decomposition of unpolluted Scots pine needle litter was studied in two heavy-metal-pollution gradients in Sweden; one near a brass mill and the other around a primary smelter. In the latter area locally collected polluted Scots pine needle litter was also incubated. Decomposition rates were strongly influenced by the metal pollution and a decrease in the rate of mass-loss occurred. In the brass-mill gradient this occurred until about 1 km from the pollution source which corresponded to about 500 µg Cu and 1 000 µg Zn g^?1 soil. Data are presented to indicate that lignin decomposition was more sensitive to pollution than decomposition of whole litter and affected further away from the pollution sources. At the smelter sites, the metal-polluted needle litter decomposed more slowly than the unpolluted needle litter, and this difference was enhanced close to the smelter. The results indicate that heavy metals accumulated in needles prior to shedding have a long-term impact on the subsequent decomposition of the litter. Both litter quality and soil factors thus contribute to the reduced litter decomposition rate in metal-polluted forests. A new non-linear model with decreasing decay rate was used in the statistical evaluation. The model can be used to characterize the effects of pollution on decomposition rate.     

Effects of fungal inocula on the decomposition of lignin and structural polysaccharides in Pinus sylvestris litter

 Litter bags containing sterile Scots pine (Pinus sylvestris) needles (19.8% lignin, 26.5% cellulose and 0.34% N) were inoculated with two species of fungi in the laboratory and then placed in the litter layer of a pine plantation. Marasmius androsaceus, which can degrade lignocellulose, was initially displaced by other fungal colonisers and was not detected in the litter after 2–3 months; but was re-isolated from the needles after 12 months. Trichoderma viride, which is a cellulolytic species and also antagonistic to other fungi, dominated the litter throughout the experiment. The control litter was naturally colonised by litter fungi. After 12 months, mass losses were similar at 52% for M. androsaceus and 48% for T. viride, compared with 36% for the control litter colonised by a more complex fungal community. Lignin concentrations increased with time in control litter and with T. viride because mass losses of carbohydrates were greater than those of lignin. Litter inoculated with M. androsaceus showed significant lignin decomposition throughout the experiment but cellulose concentrations showed a proportional increase in the first 6 months, suggesting that the fungus was preferentially exploiting hemicellulose and non-structural carbohydrates. Analysis of TFA-extractable sugars (mainly from hemicellulose) and CuO-derived phenylpropanoid moieties from lignin confirmed the differential patterns of resource decomposition which were not evident from total mass losses. During the initial stages of decomposition, T. viride was as effective in utilising structural polysaccharides as the complex fungal community in the control litter. Furthermore, M. androsaceus not only exhibited unexpectedly low cellulolytic activity but also facilitated lignin depolymerisation after the fungus was no longer detectable in the litter. The pre-inoculation of litter with these two fungal species therefore affected the overall dynamics of decomposition at a biochemical level. This study illustrates the importance of understanding the effects and interactions of specific fungi, rather than assumptions about the functional competence of diverse communities, on the processes of litter decomposition. Received: 5 July 2000     

Changes in the lignin fraction of spruce and pine needle litter during decomposition as studied by some chemical methods

Changes in the lignin fraction of spruce and pine needle litter were followed by four different methods: Klason lignin, phloroglucinol lignin, dioxane-water-HCl-lignin and alkaline CuO oxidation. The decomposition patterns of the lignins studied were different, the largest differences between the methods being obtained for the spruce needles. Depending on the method used, between 36 and 46% of the original amount of lignin remained in the pine needles and about 30–61% in the spruce needles after 3 yr of decomposition. The decomposition rates of the various lignin pools were highly correlated with the loss in mass of the litter. The phloroglucinol lignin was decomposed significantly faster (P < 0.001) in the spruce needles than in the pine needles for as long as the decomposition process was followed. During decomposition of the litter, the residual amount of Klason lignin was correlated with the residual amounts of the other lignins. The application of the different methods to litter decomposition studies is discussed.     

Soil carbon and nitrogen budget in Scots pine (Pinus sylvestris L.) stands along an air pollution gradient in eastern Germany

Litterfall, bio- and necromass of the forest floor vegetation, decomposition of recent organic material, soil respiration and humus stocks were examined in 3 Scots pine stands along an air pollution gradient in eastern Germany. High nitrogen loads and increased pH values due to Ca deposition caused shifts in the vegetation structure, and higher biomass production of the forest floor vegetation, whereas needle litter production was not impacted. Simultaneously, decomposition rates of the recently harvested forest floor vegetation decreased with increasing pollutant loads, but needle litter and soil organic matter decomposition rates did not differ between the sites. Consequently, soil carbon and nitrogen stocks increased with increasing pollutant input.     

Microbial biomass and activity in soil and litter underPinus sylvestris, Picea abies andBetula pendula at originally similar field afforestation sites

Microbial biomass C and N, and activities related to C and N cycles, were compared in needle and leaf litter, and in the uppermost 10 cm of soil under the litter layer in Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies L.) and silver birch (Betula pendula L.) stands, planted on originally similar field afforestation sites 23–24 years ago. The ground vegetation was differentiated under different tree species, consisting of grasses and herbs under birch and pine, and mosses or no vegetation with a thick layer of needles under spruce. The C:N ratio of the soils was 13–21 and the soil pH_{CaCl 2} 3.8–5.2. Both showed little variation under different tree species. Microbial biomass C and N, C mineralization, net ammonification, reduction) did not differ significantly in soil under different tree species either. Birch leaf litter had a higher pH_{CaCl 2} (5.9) than spruce and pine needle litter (pH 5.0 and 4.8, respectively). The C:N ratio of spruce needles was 30, and was considerably higher in pine needles (69) and birch leaves (54). Birch leaves tended to have the highest microbial biomass C and C mineralization. Spruce needles appeared to have the highest microbial biomass N and net formation of mineral N, whereas formation of mineral N in pine needles and birch leaves was negligible. Microbial biomass C and N were of the same order of magnitude in the soil and litter samples but C mineralization was tenfold higher in the litter samples.     

CPMAS 13C NMR and IR spectra of spruce and pine litter and of the Klason lignin fraction at different stages of decomposition

Fresh and decomposed spruce and pine litter and the Klason lignin fraction of spruce needles at different stages of decomposition were studied by CPMAS ¹³C NMR and IR spectroscopy as well as by chemical methods. It was shown that decomposition of needles is accompanied by an increase in aliphatic substances and carboxyl group content; the amount of polysaccharides is reduced. It is assumed that stable aliphatic compounds like cutin and lipids of microbial origin will accumulate during litter decomposition and humification. Aromaticity is low and does not alter drastically. The NMR spectra of the Klason lignin fraction show pronounced peaks at 30, 55, 115, 130, 150 and 175 ppm. Obviously, this fraction contains appreciable amounts of aliphatic and carboxyl carbon besides the typical aromatic units of lignin. During decomposition aromaticity decreases whereas the relative amounts of aliphatic substances and carboxyl groups increase. This is probably due to splitting of aromatic ring structures and side chains. The findings agree with the results from chemical analyses.     

Lignin degradation controls the production of dissolved organic matter in decomposing foliar litter

Lignin is considered to be a crucial component controlling litter decomposition but its role in the production of dissolved organic matter (DOM) from litter is not well understood. Our main objective therefore was to examine the amounts and properties of DOM produced in decomposing litter, with special emphasis on the role of lignin degradation. We exposed litter of five different tree species (Sycamore maple, Mountain ash, European beech, Norway spruce, Scots pine) in litterbags at the soil surface of two neighbouring sites to degradation under field conditions. Litterbags were sampled eight times during 27 months of exposure in the field. We determined mass loss and characterized the lignin fraction by two different methods (van Soest procedure, acid‐detergent lignin: ADL, CuO oxidation). Litter was irrigated in the laboratory and leachates were analysed for dissolved organic carbon (DOC) and characterized by UV and fluorescence spectroscopy. Litter decomposition followed a two‐stage model characterized by initially rapid and then decreasing degradation with time. In the initial phase of litter decomposition, leached amounts of DOM decreased with time and no effects of lignin degradation were found. The contents of ADL in the litter residues and CuO oxidation products suggest larger degradation and oxidation of lignin in beech, spruce and pine litter than in maple and ash litter. The production of DOM from litter with larger lignin degradation increased in the second phase of decomposition, when mass loss exceeded 10–20%. In contrast, DOM produced from litter showing weak lignin degradation (maple, ash) did not increase further in the second phase of decomposition. In the leachates of litter with large lignin degradation (beech, spruce, pine), UV absorbance and fluorescence spectroscopy indicated a larger increase in the contribution of lignin‐derived compounds to DOM with increasing mass loss than for litter species with relatively stable lignin. We conclude that degradation of lignin is an important control on DOM production during the second phase of litter decomposition.     

Microbial biomass and activity in soil and litter underPinus sylvestris, Picea abies andBetula pendula at originally similar field afforestation sites

Microbial biomass C and N, and activities related to C and N cycles, were compared in needle and leaf litter, and in the uppermost 10 cm of soil under the litter layer in Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies L.) and silver birch (Betula pendula L.) stands, planted on originally similar field afforestation sites 23–24 years ago. The ground vegetation was differentiated under different tree species, consisting of grasses and herbs under birch and pine, and mosses or no vegetation with a thick layer of needles under spruce. The C:N ratio of the soils was 13–21 and the soil pH_{CaCl 2} 3.8–5.2. Both showed little variation under different tree species. Microbial biomass C and N, C mineralization, net ammonification, reduction) did not differ significantly in soil under different tree species either. Birch leaf litter had a higher pH_{CaCl 2} (5.9) than spruce and pine needle litter (pH 5.0 and 4.8, respectively). The C:N ratio of spruce needles was 30, and was considerably higher in pine needles (69) and birch leaves (54). Birch leaves tended to have the highest microbial biomass C and C mineralization. Spruce needles appeared to have the highest microbial biomass N and net formation of mineral N, whereas formation of mineral N in pine needles and birch leaves was negligible. Microbial biomass C and N were of the same order of magnitude in the soil and litter samples but C mineralization was tenfold higher in the litter samples.     

Decomposition of root litter and some factors regulating the process: Long-term root litter decomposition in a scots pine forest

Decomposition of root litter was studied using Scots pine roots (six diameter classes) and rhizomes from heather (three diameter classes) and cowberry (one diameter class). For Scots pine roots, root diameter was correlated with initial concentrations of N, P, S and Mg but not with organic-chemical composition. The highest nutrient concentrations were found in Scots pine roots and the lowest in heather rhizomes, with cowberry rhizomes intermediate. The highest lignin concentrations were in heather and cowberry rhizomes. In the early decomposition stages diameter and nutrient concentration correlated with mass loss rate in Scots pine roots: in a comparison Scots pine roots were degraded faster than cowberry rhizomes which, in turn, were degraded faster than heather rhizomes. Root diameter, however, may not be important in decomposition of cowberry and heather rhizomes but nutrient and lignin concentrations appear important in all three species. In the late decomposition stages only Scots pine roots could be compared and it appeared that there was a negative correlation with lignin concentration and mass loss, and no correlation with any nutrient.     

Scots pine litter decomposition along drainage succession and soil nutrient gradients in peatland forests, and the effects of inter-annual weather variation

Peatlands form a large carbon (C) pool but their C sink is labile and susceptible to changes in climate and land-use. Some pristine peatlands are forested, and others have the potential: the amount of arboreal vegetation is likely to increase if soil water levels are lowered as a consequence of climate change. On those sites tree litter dynamics may be crucial for the C balance. We studied the decomposition of Scots pine (Pinus sylvestris L.) needle and root litter in boreal peatland sites representing gradients in drainage succession (succession following water level drawdown caused by forest drainage) and soil nutrient level during several years of varying weather conditions. Neither gradient had an unambiguous effect on litter mass loss. Mass loss over 2 years was faster in undrained versus drained sites for both needle litter, incubated in the moss layer, and fine root litter, incubated in 0-10 cm peat layer, suggesting moisture stress in the surface layers of the drained sites limited decomposition. Differences among the drained sites were not consistent. Among years, mass loss correlated positively with precipitation variables, and mostly negatively or not at all with temperature sum. We concluded that a long-term water level drawdown in peatlands does not necessarily enhance decay of fresh organic matter. Instead, the drained site may turn into a ‘large hummock-system’ where several factors, including litter quality, relative moisture deficiency, higher acidity, lower substrate temperature, and in deeper layers also oxygen deficiency, may interact to constrain organic matter decomposition. Further, the decomposition rates may not vary systematically among sites of different soil nutrient levels following water level drawdown. Our results emphasize the importance of annual weather variations on decomposition rates, and demonstrate that single-period incubation studies incorporate an indeterminable amount of temporal variation.     

A reciprocal decomposition experiment of Scots pine needles 19 yr after wood ash fertilization

Scots pine (Pinus sylvestris) needle litter originating from control plots and plots that had received a wood ash fertilization (3 t ha⁻¹) 19 yr earlier were allowed to decompose in a reciprocal experimental design to detect the effects of ash fertilization and needle litter origin on the decomposition rate. The experimental design was repeated in two Scots pine forest stands of different fertility and the litterbags were harvested after 4 and 16 months. Ash fertilization resulted in a higher needle litter decomposition rate but the needle origin did not influence the results. Stand fertility correlated positively to the decomposition rate.     

A comparison of litterbag and direct observation methods of Scots pine needle decomposition measurement

The litterbag method (LB) widely used in decomposition studies was compared to a direct observation method (DO) of the litter collected in the forest floor in three European Scots pine forests. In this last method, needles were sorted out of blocks of the forest floor and separated into 5 categories using morphological criteria representing different layers and decomposition levels. The residence time of the needles in each layer was calculated using the litterfall input, the size of the layer in the forest floor and its level of decomposition. The mass loss of each layer was determined using the linear mass density method. To estimate decomposition rates, the DO method might be more advantageous than the LB method because it is done without any manipulation or microclimatic artifact and because it integrates spatial and temporal variability. It is also convenient because the sampling does not have to be done simultaneously on different sites to allow site comparisons. Indeed, the collected litter blocks include the whole site history until the needle collapses which has been shown to take place after a period of up to 10 years in the present study. After 3 years, the mass loss was lower measured by the DO method than by the LB method. The decrease of litter mass measured by the DO method was best described by a simple asymptotic function of time. This model has a mechanistic signification because it underlines the possible long-term accumulation of recalcitrant products which is not the case with the exponential function. This accumulation of recalcitrant products is a key process in the storage of carbon in the organic matter reservoir of forest soils. Nevertheless this method is time consuming and cannot replace the LB method for large data set collection. This paper recommends when using the LB method to be aware of its inherent bias that can lead to misestimating decomposition in the long run.     

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.     

Decomposition of black locust and black pine leaf litter in two coeval forest stands on Mount Vesuvius and dynamics of organic components assessed through proximate analysis and NMR spectroscopy

Litter quality is an important determinant of soil organic matter formation. Changes of organic components were investigated along decomposition of black locust (Robinia pseudoacacia L.) leaf litter and black pine (Pinus nigra Arn.) needle litter in the native adjacent coeval forest stands. To this purpose, data from proximate analyses were compared with those from CPMAS ¹³C NMR. Newly shed leaf litter of black locust had significantly higher concentrations of ADSS (acid detergent soluble substances) as well as lower concentrations of cellulose and AUR (acid unhydrolyzable residues that include lignin) and higher AUR-to-Cellulose ratio than that of black pine. The ¹³C CPMAS NMR spectra of newly shed leaf litter of black locust and black pine revealed that O-Alkyl-C components (including cellulose and hemicelluloses) accounted, respectively, for 53.8% and 61.4% of the total area of the spectra. All other C fractions were relatively more abundant in black locust than in black pine. Within individual sampling periods, relationships between residual litter mass and concentrations of ADSS, cellulose and AUR were examined, as were relationships between residual litter C and NMR fractions. Four periods were defined based on the slopes of the decomposition curve, with the length of period I defined by the start of a net decrease of AUR. Proximate analyses and NMR data showed changes in chemical composition over the decomposition process, as well as changes in decay rates of the residues, following different paths in the two litters. ADSS decayed faster in black locust litter; in contrast cellulose and AUR decayed faster in that of black pine. AUR concentration increased in both litters during decomposition; however, compared to black pine, the remaining litter of black locust was richer in AUR, despite the lower initial concentration, and had a higher AUR-to-Cellulose ratio. Phenol-C and Aryl-C decayed faster in black locust litter, while Alkyl-C decayed faster in that of black pine. In both litters, mass loss in periods was negatively correlated to concentration of AUR at the start of the periods. C loss in periods was negatively correlated to the concentration at the start of the periods of MC-to-PC (an index of lignin content) in black locust litter and positively correlated to Alkyl-C and O-Alkyl-C in that of black pine. Phenol-C, O-Alkyl-C and Aryl-C were the most decomposable C fractions in black locust. O-Alkyl-C and Alkyl-C were the most decomposable C fractions in black pine. Limit value was lower in black pine than in black locust. Consequently the different pattern of litter decomposition can affect the size of C sequestration in the forest floor and the quality of accumulated organic carbon.     

Processes controlling the production of aromatic water-soluble organic matter during litter decomposition

Dissolved organic matter (DOM) plays a fundamental role for many soil processes. For instance, production, transport, and retention of DOM control properties and long-term storage of organic matter in mineral soils. Production of water-soluble compounds during the decomposition of plant litter is a major process providing DOM in soils. Herein, we examine processes causing the commonly observed increase in contribution of aromatic compounds to WSOM during litter decomposition, and unravel the relationship between lignin degradation and the production of aromatic WSOM. We analysed amounts and composition of water-soluble organic matter (WSOM) produced during 27 months of decomposition of leaves and needles (ash, beech, maple, spruce, pine). The contribution of aromatic compounds to WSOM, as indicated by the specific UV absorbance of WSOM, remained constant or increased during decomposition. However, the contribution of lignin-derived compounds to the total phenolic products of ¹³C-labelled tetramethylammonium hydroxide (¹³C-TMAH) thermochemolysis increased strongly (by >114%) within 27 months of decomposition. Simultaneous changes in contents of lignin phenols in solid litter residues (cupric oxide method as well as ¹³C-TMAH thermochemolysis) were comparably small (−39% to +21% within 27 months). This suggests that the increasing contribution of lignin-derived compounds to WSOM during decomposition does not reflect compositional changes of solid litter residues, but rather the course of decomposition processes. In the light of recently published findings, these processes include: (i) progressive oxidative alteration of lignin that results in increasing solubility of lignin, (ii) preferential degradation of soluble, non-lignin compounds that limits their contribution to WSOM during later phases of decomposition.     

Short-Term Influence of Clinker Dust and Wood Ash on Macronutrients and Growth in Norway Spruce (Picea abies) and Scots Pine (Pinus sylvestris) Seedlings

The effects of clinker dust and wood ash on Norway spruce and Scots pine seedlings were compared in buried pots. Clinker dust (0.5 kg m^?2) and wood ash (0.5 kg m^?2) were applied to the surface of a nutrient-poor mineral soil. In the second year, the increase in soil pH by the dust and ash were larger than in the first year. Both alkaline treatments caused a large increase in the needle potassium (K) concentration. An excess of soil K relative to magnesium (Mg) was observed by decreased Mg concentration in needles shortly after treatment. However, Mg concentration in needles stayed in the sufficiency range. Current results confirmed earlier findings that despite a positive effect on base cation nutrition, wood ash has a low potential for increasing the biomass of forest stands on mineral soils due to the N limitation in these soils.     

The effect of oxygen,pH and organic carbon on soil-layer specific denitrifying capacity in acid coniferous forest

Emissions of N₂O from acid coniferous forest soils are found to be low and considered to be due to nitrification rather than denitrification. Recently we have demonstrated soil-layer specific denitrification in a Scots pine forest in the Netherlands. N₂O production, in the presence of high concentrations of acetylene, was detected in the intact needle fraction but was absent in the fragmentation layer of this forest soil. To identify the factors regulating denitrification activity, in the present study the effects of oxygen, pH and organic carbon were investigated in the needle and fragmentation fraction of acid coniferous forest soils. Under natural circumstances denitrification in the Scots pine needles was higher than in Douglas fir needles and absent in fragmentation material. Under anaerobic conditions comparable N₂O production in the two soil types was found in needle suspensions of both forest types, indicating that differences in anaerobic microsites were responsible for different N₂O production under aerobic circumstances. Denitrifying capacity was absent in the fragmentation layer; under anaerobic circumstances little N₂O was produced. Neither an addition of available carbon (glucose and succinate) nor an increase in pH revealed a denitrifying capacity comparable to that observed in needles. The increase in pH, under anaerobic circumstances, was most effective on N₂O production in the fragmentation material. The denitrifying capacity in the fragmentation layer remained low during short-term incubation under optimal conditions. This indicates the presence of a low denitrifying population, most likely due to aerobic conditions, low pH and low available organic carbon. Although the significance of N₂O production under natural conditions remains speculative, this study seeks to clarify soil-layer specific denitrifying activity in acid coniferous forest soils.     

Response of soil C and N transformations to tannin fractions originating from Scots pine and Norway spruce needles

Tannins are polyphenolic compounds that may influence litter decomposition, humus formation, nutrient (especially N) cycling and ultimately, plant nutrition and growth. The aim of this study was to determine the response of C and N transformations in soil to tannins of different molecular weight from Norway spruce (Picea abies (L.) Karst) and Scots pine (Pinus sylvestris L.) needles, tannic acid and cellulose. Arginine was added to test whether the soil microbial community was limited by the amount of N, and arginine+tannin treatments were used to test whether the effects of tannins could be counteracted by adding N. Soil and needle samples were taken from adjacent 70-year-old Scots pine and Norway spruce stands located in Kivalo, northern Finland. Tannins were extracted from needles and fractioned based on molecular weight; the fractions were then characterized by LC-MS and GC-MS. Light fractions contained tannin monomers and dimers as well as many other compounds, whereas heavy fractions consisted predominantly of polymerized condensed tannins. Spruce needles contained more procyanidin than prodelphinidin units, while in pine needles prodelphinidin units seemed to be dominant. The fractions were added to soil samples, pine fractions to pine soil and spruce fractions to spruce soil, and incubated at 14 °C for 6 weeks. CO₂ evolution was followed throughout the experiment, and the rates of net mineralization of N and net nitrification, concentration of dissolved organic N (DON) and amounts of microbial biomass C and N were measured at the end of the experiment. The main effects of the fractions were similar in both soils. Light fractions strongly enhanced respiration and decreased net N mineralization, indicating higher immobilization of N in the microbial biomass. On the contrary, heavy fractions reduced respiration and slightly increased net N mineralization, suggesting toxic or protein-precipitating effects. The effects of tannic acid and cellulose resembled those of light fractions. DON concentrations generally decreased during incubation and were lower with heavy fractions than with light fractions. No clear differences were detected between the effects of light and heavy fractions on microbial biomass C and N. Treatments that included addition of arginine generally showed trends similar to treatments without it, although some differences between light and heavy fractions became more obvious with arginine than without it. Overall, light fractions seemed to act as a labile source of C for microbes, while heavy fractions were inhibitors.     

Amounts of carbon mineralised and leached as DOC during decomposition of Norway spruce needles and fine roots

Changes in climate or forest management practices leading to increased litter production will most likely cause increased leaching rates of dissolved organic carbon (DOC) from the O horizon. The rhizosphere is often assumed to have a large carbon flux associated with root turnover and exudation. However, little has been done to quantify the amount of DOC originating from root litter. We studied decomposition of fine root and needle litter of Norway spruce (Picea abies) through a combined incubation and leaching experiment in the laboratory using five different litter types: fresh needle litter, aged needles from the litter layer, fresh and dead roots from mineral soil samples, and seven-year-old roots from a previous litterbag study. After respiration measurements, the samples were percolated with artificial throughfall water and DOC and UV absorbance were measured in the leachate. Mineralisation of dissolved organic matter in the leachate and sorption of DOC to ferrihydrite were determined as a measure of DOC ability to be stabilised by iron (hydr)oxide surfaces.The mineralisation rate and DOC production rate of root samples were always lower than that of needle samples. However, root and needle derived dissolved organic matter (DOM) were similar in terms of aromaticity, as indicated by their specific UV absorbance, and ability to be sorbed by ferrihydrite. For seven-year-old roots, a significantly higher fraction of carbon was lost as DOC (30%) than for younger roots (20%). Furthermore, DOM from old roots bound more strongly to ferrihydrite and is mineralised at a lower rate than DOC from younger roots, suggesting that roots at late stages of decomposition, although a small fraction of total litter, significantly contribute to carbon build-up in mineral soils. The slower decomposition rate of roots compared with needles must be taken into account when modelling litter decomposition.     

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.