Plant litter quality influences the contribution of soil fauna to litter decomposition in humid tropical forests, southwestern China

The aim of this field experiment was to quantify the contribution of soil fauna to plant litter decomposition in three forest sites differing in C/N ratio under natural conditions in Xishuangbanna, southwestern China. We conducted a survey of soil fauna communities, the forest floor litter and investigated mass loss of mixed tree species leaf litter for two years in a tropical secondary forest, an evergreen broad-leaf forest and a tropical rain forest. Exclusion treatments of different sized soil fauna from the leaf litter by using varying mesh size litter bags (2 mm and 0.15 mm) were also performed. Mass loss and C and N concentrations in litter bag leaf materials were determined at monthly intervals. We found that: (1) the three forests differed in floor litter biomass and nutrient contents but not in soil fauna richness and abundance; (2) litter mass loss and decomposition rate were slower when soil macrofauna and most of mesofauna were excluded; and (3) greatest soil fauna contribution to plant litter decomposition occurred in the rain forest, where leaf litter C/N ratio was also highest (41.5% contribution: 54.8 C/N ratio), in comparison to 8.69% in the broad-leaf forest and 19.52% in the secondary forest, both with low leaf litter C/N ratios (<32). Our results suggested that, soil fauna played a more pronounced role in the decomposition of mixed leaf litter in tropical rain forest, and significantly bigger effects from fauna were ascribed to the enhancement of N concentration and decrease of C concentration of the initially high C/N ratio litter in this forest site.     

Soil fauna alter the effects of litter composition on nitrogen cycling in a mineral soil

Plant chemical composition and the soil community are known to influence litter and soil organic matter decomposition. Although these two factors are likely to interact, their mechanisms and outcomes of interaction are not well understood. Studies of their interactive effects are rare and usually focus on carbon dynamics of litter, while nutrient dynamics in the underlying soil have been ignored. A potential mechanism of interaction stems from the role fauna plays in regulating availability of litter-derived materials in the mineral soil. We investigated the role of soil fauna (meso, macro) in determining the effect of surface-litter chemical composition on nitrogen mineralization and on the micro-food web in mineral soils. In a field setting we exposed mineral soil to six types of surface-applied litter spanning wide ranges of multiple quality parameters and restricted the access of larger soil animals to the soils underlying these litters. Over six months we assessed litter mass and nitrogen loss, nitrogen mineralization rates in the mineral soils, and soil microbes and microfauna. We found evidence that the structure of the soil community can alter the effect of surface-litter chemical composition on nitrogen dynamics in the mineral soil. In particular, we found that the presence of members of the meso- and macrofauna can magnify the control of nitrogen mineralization by litter quality and that this effect is time dependent. While fauna were able to affect the size of the micro-food web they did not impact the effect of litter composition on the abundance of the members of the micro-food web. By enhancing the strength of the impact of litter quality on nitrogen dynamics, the larger fauna can alter nitrogen availability and its temporal dynamics which, in turn, can have important implications for ecosystem productivity. These findings contribute to evidence demonstrating that soil fauna shape plant litter effects on ecosystem function.     

Predicting soil fauna effect on plant litter decomposition by using boosted regression trees

Extensive studies have been conducted to evaluate the effect of soil fauna on plant litter decomposition in terrestrial ecosystems. However, scholars have reported inconsistent results on the direction and magnitude of the soil fauna effect. We present a global synthesis of 75 papers that cover 197 plant species with 543 cases of plant litter decomposition experiments and soil fauna effects on plant litter decomposition. By using a boosted regression tree model (BRT), we aim to provide a synthesis of existing data that comprehensively and quantitatively evaluates how climate, plant litter quality, and study methods interact with soil fauna in affecting plant litter decomposition. Global average effect size (ES) is −0.426, which indicates a 35% lower decomposition rate when soil fauna is excluded by physical or chemical exclusion techniques. The final model explains 32.3% of the variation in ES. The predictors that substantially account for the explained variation include mean annual temperature (MAT, 37.1%), mean annual precipitation (MAP, 9.7%), phosphorus (12.4%), nitrogen (5.6%), and lignin content (5.5%). By contrast, the heterogeneity of the study duration and soil fauna exclusion technique have negligible contributions (each <5%). Log effect size strongly decreases with both MAT and MAP. Plant litters with high quality have stronger soil fauna effect because the log effect size is negatively related to nitrogen and phosphorus content and positively related to lignin content. Our analysis demonstrates the critical role of climate and plant litter quality in determining the soil fauna effect on plant litter decomposition in terrestrial ecosystems. However, the large unexplained variation (67.7%) in ES in the BRT model indicates undiscovered mechanisms underlying the soil fauna effect in our analysis. We call for further studies on this topic in the future.     

Effects of litter quality and microarthropods on N dynamics and retention of exogenous 15N in decomposing litter

Summary Surface additions of (¹⁵NH₄)₂SO₄ were used to measure the immobilization and subsequent movement of exogenous N added to two litter types of contrasting quality (Cornus florida and Quercus prinus). Litterbaskets were used to measure the litter mass loss and N dynamics and to follow the movement of the ¹⁵N label through litter, F layer, and soil pools. Half of the litterbaskets of each species were treated with naphthalene to reduce microarthropod densities. The faster decomposing C. florida litter maintained a higher excess atom % ¹⁵N, and a greater relative concentration of the labeled input (g ¹⁵N g^–1) than did Q. prinus litter. In both litter types the excess atom % ¹⁵N, relative concentration (g ¹⁵N g^–1), and absolute amount of label recovered in the litter declined over time. This occurred during a period of net accumulation of total litter N, implying simultaneous release of the initial input and immobilization of N from other sources. The concentration of ¹⁵N in the soil increased over time, while the F layer apparently acted as an intermediary in the transfer of ¹⁵N from litter to soil. Naphthalene effectively reduced microarthropod numbers in all horizons of the litterbaskets and significantly reduced the decay rates of Q. prinus, but not C. florida litter. Naphthalene did not appear to affect total N dynamics in the litter. However, with all horizons taken together, the naphthalene-treated litterbaskets retained more total ¹⁵N than the control litterbaskets. Naphthalene also changed the vertical distribution of ¹⁵N within litterbaskets, so that the litter retained less of the ¹⁵N-labeled input and the F layer and soil horizons retained more of the labeled input than in control litterbaskets. Our major conclusions are: (1) the N pool of decomposing litter is dynamic, with simultaneous N release and immobilization activating N turnover even during the net accumulation phase; (2) litter quality is an important determinant of immobilization and retention of exogenous N inputs and, therefore, turnover of the litter N pool; and (3) microarthropod activity can significantly affect the incorporation and retention of exogenous N inputs in decomposing litter, although these changes are apparently not reflected in net N accumulation or release during the 1st year of decomposition. However, the naphthalene may have affected microbially mediated N dynamics and this possibility needs to be considered in interpreting the results.     

Effect of chemical composition on the release of nitrogen from agricultural plant materials decomposing in soil under field conditions

Summary In two field experiments, plant materials labelled with ¹⁵N were buried separately within mesh bags in soil, which was subsequently sown with barley. In the first experiment, different parts of white clover (Trifolium repens), red clover (T. pratense), subterranean clover (T. subterraneum), field bean (Vicia faba), and timothy (Phleum pratense) were used, and in the second, parts of subterranean clover of different maturity. The plant materials were analysed for their initial concentrations of total N, ¹⁵N, C, ethanol-soluble compounds, starch, hemicellulose, cellulose, lignin, and ash. After the barley had been harvested, the bags were collected and analysed for their total N and ¹⁵N. In the first experiment the release of N was highest from white clover stems + petioles (86%) and lowest from field bean roots (20%). In stepwise regression analysis, the release of N was explained best by the initial concentrations of lignin, cellulose, hemicellulose, and N (listed according to decreasing partial correlations). Although the C/N ratio of the plant materials varied widely (11–46), statistically the release of N was not significantly correlated with this variable. The results of the second experiment using subterranean clover of different maturity confirmed those of the first experiment.     

土壤C/N对苹果植株生长及氮素利用的影响

土壤C/N是土壤氮素循环的重要影响因素。本研究以2年生"富士"/平邑甜茶为试验材料, 应用¹⁵N示踪技术研究了不同土壤C/N[6.21(CK)、10、15、20、25、30、35和40]对苹果植株生长及氮素利用和损失的影响。结果表明: 随着土壤C/N比值的逐渐增大, 苹果新梢长度和植株鲜重均呈先升高后降低的变化趋势, C/N=15、20和25的3个处理苹果新梢长度和植株鲜重最大, 三者间无显著差异, 但均显著高于其他处理。不同C/N处理间植株15N利用率存在差异, 土壤C/N=25时, 植株¹⁵N利用率最大, 为22.87%, 与C/N=20的处理间无显著差异, 但两者均显著高于其他处理; 土壤C/N=40时, 植株¹⁵N利用率最低, 仅为15.43%, 低于CK处理的16.65%。土壤C/N处于15~25时, 植株吸收的氮素来自于肥料氮的比例较高; 而土壤C/N较低(<15)或太高(>25)时, 植株吸收的氮素来自于土壤氮的比例较高。土壤氮素残留量随土壤C/N的增大逐渐增加, C/N=40处理的土壤氮素残留量是CK的1.32倍。随着土壤C/N比值的逐渐增大, 肥料氮损失量呈先减少后增加的变化趋势, 以C/N=25时最少, 仅为施氮量的49.87%, 而对照最大, 为61.54%。因此, 综合土壤C/N对苹果植株生长及氮素平衡状况来看, 土壤C/N为15~25时, 能促进植株的生长发育, 降低氮肥损失, 提高肥料利用率。     

A N tracing model to analyse N transformations in old grassland soil

A new ¹⁵N tracing model was developed to analyse nitrogen (N) transformations in old grassland soil. There was a need to develop a new model because existing models such as FLUAZ were not able to simulate the observed N dynamics. The new features of the model are: (a) simulation of heterotrophic nitrification, (b) simulation of dissimilatory nitrate (NO₃⁻) reduction to ammonium (NH₄⁺) (DNRA), (c) release of adsorbed or stored fertiliser N into the available mineral N pools and (d) immobilisation of NH₄⁺ and NO₃⁻ into two separate organic N pools with different re-mineralisation characteristics. The tracing model contains six N pools and nine simultaneous N transformations either at zero- or first-order kinetics. The model is set up in the modelling software ModelMaker which contains non-linear optimisation routines based on the Marquardt-Levenberg algorithm. The model is able to simulate data obtained from triple labelling studies where either the NH₄⁺, the NO₃⁻ or both pools were labelled with ¹⁵N. The flexible modelling environment allows the user to develop the model further.     

Relative significance of soil and nitrogenous fertilizer in nitrogen nutrition and growth of wetland rice (Oryza sativa L.)

Summary A pot experiment was conducted to compare the yields from five commercially cultivated varieties (Bas-198, Bas-370, Bas-Pak, Bas-385, and IR-6) of rice (Oryza sativa L.) and to establish the relative significance of soil N and fertilizer N (¹⁵N-labelled ammonium sulphate) in affecting crop performance. Another aim was to study the interaction of fertilizer N and soil N as influenced by different rice varieties. Among the five varieties tested, Bas-Pak gave the maximum dry matter and N yield. The N-use efficiency (percentage of applied N taken up by the plants) of different varieties ranged between 33.7 and 43.7%, Bas-Pak being the most efficient. Significant losses of fertilizer N occurred from the soil-plant system. The maximum N loss (52.1% of applied N) was observed with IR-6 and the minimum loss (39.2%) with Bas-Pak. A substantial increase in the uptake of soil N following the application of fertilizer and an interaction between the two N sources were observed with all varieties except Bas-385. The interaction was attributed to greater root proliferation following the application of fertilizer. It was concluded that a varietey with greater potential to use soil N is likely to give a better yield and that, of the two N sources, the availability of soil N was more important in determining the yield performance of different varieties of rice.     

Coarse particulate organic matter is the primary source of mineral N in the topsoil of three beech forests

Forest soils contain a variable amount of organic N roughly repartitioned among particles of different size, microbial biomass and associated with mineral compounds. All pools are alimented by annual litter fall as main input of organic N to the forest floor. Litter N is further subject to mineralization/stabilization recognized as the crucial process for the turnover of litter N. Although it is well documented that different soil types have different soil N stocks, it is presently unknown how different soil types affect the turnover of recent litter N. Here, we compared the potential mineralization of the total soil organic N with that of recent litter-released N in three beech forests varying in their soil properties. Highly ¹⁵N-labelled beech litter was applied to stands located at Aubure, Ebrach, Collelongo, which differ in humus type, soil type and soil chemistry. After 4-5 years of litter decomposition, the upper 3 cm of the organo-mineral A horizon was sampled and the net N mineralization was measured over 112 days under controlled conditions. The origin of mineralized N (litter N versus soil organic N) was calculated using ¹⁵N labeling. In addition, soils were fractionated according to their particle size (>2000 μm, 200-2000 μm, 50-200 μm, <50 μm) and particulate organic matter (POM) was separated from the mineral fraction in size classes, except the <50 μm fraction. Between 41 and 69% of soil organic N was recovered as POM. Litter-released ¹⁵N was mainly to be found in the coarse POM fractions >200 μm. On a soil mass basis, N mineralization was two-fold higher at Aubure and Collelongo than at Ebrach, but, on a soil N basis, N mineralization was the lowest at Collelongo and the highest at Ebrach. On a soil N (or ¹⁵N) basis, mineralization of litter ¹⁵N was two to four-fold higher than mineralization of the average soil N. Furthermore, the δ¹⁵N of the mineral N produced was closer to that of POM than to that of the mineral-bound fraction (<50 μm). Highest rates of ¹⁵N mineralization happened in the soil with the lowest N content, and we found a negative relationship between accumulations of N in the upper A horizon and the mineralization of ¹⁵N from the litter. Our results show that mineral N is preferentially mineralized from POM in the upper organo-mineral soil irrespective of the soil chemistry and that the turnover rate of litter N is faster in soils with a low N content.     

Contribution of red alder to soil nitrogen input in a silvopastoral system

Studies on Alnus rubra Bong. (red alder) were carried out to assess its potential for use as a component of a silvopastoral system. Comparison was between two treatments: red alder at 400 stems ha⁻¹ (silvopasture) and red alder at 2,500 stems ha⁻¹ (forestry control). δ¹⁵N values close to zero were recorded in all red alder plant parts except for root nodules, indicating that a large proportion of N in red alder was fixed from the atmosphere. Overall, it was estimated that there was 63.45 kg N ha⁻¹ fixed N accumulated in red alder trees, and the rate of N fixation was estimated at 30.95 kg ha⁻¹ year⁻¹ in the silvopasture treatment. The total amount of N that could potentially be added to the soil in the silvopasture treatment as a result of decomposition of senescent leaves, roots, and dead nodules was estimated at 40.56 kg ha⁻¹ year⁻¹. Of the total N added to the soil, 27.1 kg ha⁻¹ year⁻¹ was due to N fixation from the atmosphere. These results show that red alder has a potential to improve and maintain soil fertility in a silvopastoral system.     

Litter-type specific microbial responses to the transformation of leaf litter into millipede feces

The transformation of leaf litter into fecal pellets by saprophagous macroarthropods has long been suggested to play an important role in litter decomposition by altering microbial processes. However, conflicting results are reported in the literature, and it is currently not clear to what extent varying initial litter quality contributes to distinct microbial responses to the transformation of litter into feces. Here we performed a screening test using a wide range of distinct leaf litter from 26 tree species. We fed these litters to the macroarthropod species Glomeris marginata during one week under controlled conditions, and compared microbial responses in uningested leaf litter with that of feces produced from the 26 different leaf litter types. We assessed substrate induced respiration (SIR) as an integrative measure of microbial responses. We found that litter SIR was highly variable across species and well related to initial litter quality. However, variability in feces SIR was strongly reduced and only weakly related to initial litter quality. Moreover, the difference between feces and litter SIR decreased with increasing litter SIR as a result of higher microbial stimulation in litter with low associated litter SIR. Our data clearly showed that the direction and magnitude of microbial stimulation in feces depend strongly on the litter type. Therefore, the consequence of litter transformation into macroarthropod fecal pellets for microbial decomposers and possibly for subsequent decomposition of feces is specific to litter species.     

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.     

Mineralization of soil organic nitrogen solubilized by aqua ammonia fertilizer

Organic N solubilized by NH_3(aq) was extracted from ¹⁵N-labelled or unlabelled soil, concentrated and added to non-extracted soil, which was incubated under aerobic conditions at 27±1°C. Gross N mineralization, gross N immobilization, and nitrification in soils with or without addition of unlabelled soluble organic N were estimated by models based on the dilution of the NH ₄ ⁺ or NO _inf3 ^sup- pools, which were labelled with ¹⁵N at the beginning of incubation. Mineralization of labelled organic N was measured by the appearance of label in the mineral N pool. Although gross N mineralization and gross N immobilization were increased in two soils between day 0 and day 7 following addition of unlabelled organic N solubilized by NH_3(aq), there was no increase in net N mineralization. Solubilization of ¹⁵N-labelled organic N increased and the ¹⁵N enrichment of the soluble organic N decereased as the concentration of NH_3(aq) added increased. A constant proportion of approximately one-quarter of the labelled organic N added at different rates to non-extracted soil was recovered in the mineral N pool after an incubation period of 14 days, and the availability ratios calculated from net N mineralization data were 1.1:1 and 2.1:1 for 111 and 186 mg added organic-N kg^-1 soil, respectively, indicating that the mineralization of organic N was increased by solubilization.     

Nutrient release from decomposing leaf litter of temperate deciduous forest trees along a gradient of increasing tree species diversity

In the litter of six deciduous tree species (Fagus sylvatica, Tilia spp., Fraxinus excelsior, Carpinus betulus, Acer pseudoplatanus and Acer platanoides) and in stand-specific litter mixtures, we compared mass loss and nutrient release across diversity levels along a gradient of decreasing proportion of Fagus in stands with similar environmental and physical soil conditions. The litterbag studies ran over 22 months. The decomposition rate constants (k) of the temperate forest species ranged from k = 0.5 for Fagus to k = 1.5-2 for all other tree species. In Fagus, k was closely negatively correlated with the thickness of the litter layer and positively correlated with soil pH and isopod abundance. k was significantly higher in the mixed species stands (except for Carpinus and Fraxinus) and was positively correlated with earthworm abundance. Over the whole incubation time, nutrient amount and release rates of N, P, K, Ca and Mg as well as C-related ratios showed significant differences between tree species but no consistent differences among the diversity levels. Initial C-related nutrient ratios of the leaf litter and abundance of mesofauna and macrofauna were correlated with the length of time lag before nutrient release. We conclude that the mere number of tree species is not the main driver of nutrient release and decomposition processes, but that key groups of the decomposer fauna as well as the characteristic traits of the individual tree species are decisive.     

土壤腐殖质氮结构的X-光电能谱研究

X－ray photoelectron spectroscopy(XPS) was applied to examine the N structures of soil humic substances and some of their analogues.It was found that for soil humic substances XPS method gave similar results as those obtained by ^15N CPMAS NMR (cross-polarization magic-angle spinning nuclear magnetic resonance) method.70%-86% of total N in soil humic substances was in the form of amide,and 6%-13% was presented as ammes,with the remaining part as heterocyclic N.There was no difference in the distribution of the forms of N between the humic substances from soils formed over hundreds or thousands of years and the newly formed ones.For fulvic acid from weathered coal and benzoquinone-(NH4)2SO4 polymer the XPS results deviated significantly from the ^15N CPMAS NMR data.     

Availability of soil and fertilizer nitrogen to wheat (Triticum aestivum L.) following rice-straw amendment

Summary A pot experiment was conducted to study the N availability to wheat and the loss of ¹⁵N-labelled fertilizer N as affected by the rate of rice-straw applied. The availability of soil N was also studied. The straw was incorporated in the soil 2 or 4 weeks before a sowing of wheat and allowed to decompose at a moisture content of 60% or 200% of the water-holding capacity. The wheat plants were harvested at maturity and the roots, straw, and grains were analysed for total N and ¹⁵N. The soil was analysed for total N and ¹⁵N after the harvest to determine the recovery of fertilizer N in the soil-plant system and assess its loss. The dry matter and N yields of wheat were significantly retarded in the soil amended with rice straw. The availability of soil N to wheat was significantly reduced due to the straw application, particularly at high moisture levels during pre-incubation, and was assumed to cause a reduction in the dry matter and N yields of wheat. A significant correlation (r=0.89) was observed between the uptake of soil N and the dry matter yield of wheat with different treatments. In unamended soil 31.44% of the fertilizer N was taken up by the wheat plants while 41.08% of fertilizer N was lost. The plant recovery of fertilizer N from the amended soil averaged 30.78% and the losses averaged 45.55%     

Dynamics of dissolved and extractable organic nitrogen upon soil amendment with crop residues

Dissolved organic nitrogen (DON) is increasingly recognized as a pivotal pool in the soil nitrogen (N) cycle. Numerous devices and sampling procedures have been used to estimate its size, varying from in situ collection of soil solution to extraction of dried soil with salt solutions. Extractable organic N (EON) not only consists of DON but contains also compounds released from soil biomass and desorbed organic matter. There is no consensus whether DON or EON primarily regulates N mineralisation in soil, and their contribution to N mineralisation has not been quantified simultaneously. We evaluated three sampling procedures on their ability to determine the dynamic of dissolved organic N pools. The three procedures were the determination of DON in 1) soil solution collected by centrifugation, and the determination of EON in 2) a 0.01 M CaCl₂ extract of field moist or 3) dried soil. We added unlabeled leek and ¹⁵N-labeled ryegrass residues to a loamy sandy soil to create a temporarily increase in DON and EON, to stimulate microbial activity, and to test whether the source and dynamics of the three pools differ. We also tested whether the flow of N through DON or EON was associated with the production of inorganic N using ¹⁵N isotope tracing. Sampling procedures significantly affected the amount, but not the dynamics and origin of the three organic N pools. DON and EON (determined on field-moist and dried soils) showed all a significant increase upon crop amendment and returned to their background concentrations within 10 to 30 days. The fraction of DON and EON originating from the crop residue slightly decreased over 138 days and was not different for DON and EON. Field moist extraction of a loamy sandy soil with 0.01 M CaCl₂ gave a reliable estimate of the concentration of in situ dissolved organic N. In contrast, extraction of dried soil significantly increased EON compared to DON. The agreement in dynamics, ¹⁵N enrichment and C-to-N ratio’s indicate that dissolved and extracted organic N have a similar role in N mineralisation. Our results also suggest that they make a minor contribution to N mineralisation; changes in the turnover rate of EON were not associated with changes in the net N mineralisation rate.     

Effect of extracellular-enzyme activities on solubilization rate of soil organic nitrogen

In a sandy soil containing ¹⁵N-labeled active (soluble and easily degradable) and non-labelled passive (recalcitrant) fractions of soil organic matter, the rate of net N mineralization (solubilization) was determined during a 55-day incubation at 25°C, 63% water-holding capacity and different levels of soil extracellular-enzyme activities. The active fraction of soil N was labelled by preincubation (at 5°C and 74% water-holding capacity for 6 months) of soil amended with ¹⁵N-labeled plant material. Increases in the activity of extracellular-enzymes in soil were induced by the addition of glucose and KH₂PO₄ at the beginning of the incubation. The results show that the contents of total soluble N (NO ₃ ^– –N+NH ₄ ⁺ –N + soluble organic N) were significantly higher in glucose-amended soil compared to the unamended soil. The increases in soluble N in soil amended with 1 and 2 mg glucose g^-1 dry soil corresponded to a mean rate of net solubilization of 7.9±1.4 and 18.8±0.7 nmol N g^-1 dry soil day^-1, respectively. The mean rate of net N solubilization (3.6±1.0 nmol N g^-1 dry soil day^-1) in unamended soil was significantly lower than those of glucose amended soils. The content of ¹⁵N in total soluble N in soil amended with 2 mg glucose, for example, was diluted from 3.11±0.08 atom% before the incubation to 2.77±0.03 atom% after 55 days. This indicates that 89% of soluble-N accumulated in soil by the end of the incubation originated from the active fraction of soil N and the rest, estimated at 11%, originated from the passive fraction. The activities of soluble and total proteases as well as the rate of N solubilization in the soil increased with the application of glucose. The activity of these extracellular enzymes was highly correlated with the rates of net N solubilization. Thus, increases in extracellular-enzyme activities in glucose-amended soils had a priming effect on the solubilization of ¹⁵N-labeled active and non-labeled passive fractions of soil organic N. It seems that the activity of extracellular-enzymes expressed in terms of total and soluble protease activities could be a rate-limiting factor in the processes of soil organic N solubilization.     

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

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