Annual dynamics of phosphatase activities in an evergreen oak litter: influence of biotic and abiotic factors

As part of a study of the processes involved in litter biodegradation, we considered the variations over 1 year of the phosphatase activities in sclerophyllous evergreen oak litter (Quercus ilex L.). Evergreen oak is representative of tree species in the forests of the French Mediterranean area. Acid (E.C. 3.1.3.2.) and alkaline (E.C. 3.1.3.1.) phosphatases, were measured over 13 months in the forest litter, along with several biotic and abiotic variables, potentially involved in the regulation of these enzymes. These comprised moisture, temperature, pH, water-extractable inorganic P (P_I), fungi, culturable heterotrophic bacteria and protein concentrations. Moisture considerably affected the production of proteins and acid phosphatases, probably formed by litter microorganisms. This result corroborated the study of Criquet et al. [Soil Biology and Biochemistry 34 (2002) 1111] which indicated that rainfall was the most important factor regulating the production and the activity of numerous enzymes in sclerophyllous forest litter. However, it appeared that moisture cannot alone predict all of the variations in phosphatase activities and the mineralisation rate of organic P (P_O). Indeed, principal component analyses (PCA) and multiple regressions showed that temperature and bacterial communities were also implicated in phosphatase dynamics and P_O mineralisation. Acid phosphatases were negatively correlated with the temperature, whilst alkaline phosphatases were positively correlated with this variable. The significant correlation obtained between bacteria and P_I concentrations, and the lack of correlation between bacteria and both acid and alkaline phosphomonoesterases, suggest that other important phosphatase types, such as phosphodiesterases, must be strongly implicated in P_O mineralisation of the litter and in the regulation of P microbial metabolism.     

Spatial variations of chemical composition, microbial functional diversity, and enzyme activities in a Mediterranean litter (Quercus ilex L.) profile

Litter decomposition on the forest floor is an essential process in soil nutrient cycles and formation. These processes are controlled by abiotic factors such as climate and chemical litter quality, and by biotic factors such as microbial community diversity and activity. The aim of this study was to investigate the importance of litter depth with respect to (i) chemical litter quality as evaluated by solid-state ¹³C NMR, (ii) enzyme activities, and (iii) microbial functional diversity in four different litter layers (OLn, OLv, OF, and OH). A Mediterranean soil profile under an evergreen oak (Quercus ilex L.) forest was used as a model. The recalcitrant OM fraction, corresponding to the deepest layer, showed low enzyme activities. Peroxidases and fluorescein diacetate hydrolases (FDA) were more active in the OLn layer and probably originated largely from plants. High cellulase activity in the OLn and the OLv layers, which are rich in polysaccharides, corresponded with the high content of O-alkyl carbon compounds. Following polysaccharide degradation, laccases and lipases were much more evident in the intermediate layers. This spatial variation in nutrient demand reflected a preferential degradation of the specific plant polymers. Phosphatases were more active along the three upper layers and probably reflected a P limitation during litter degradation. Alkaline/acid (AcPAlP/AcP) ratio increased in the deepest layer, suggesting an increased participation of bacteria AlP in phosphatase pools. Results of Biolog^TM also indicated spatial variations in microbial functionality. Indeed, FF plates showed the highest functional diversity in the uppermost layer, while ECO plate functional diversity was highest in the intermediate layers. Finally, our results indicated that microbial activity and functional diversity of micro-organisms change with litter depth on a very small scale and vary with chemical organic matter (OM) composition. Thus, the observed increases in the biological variables studied were determined by the evolution of OM chemical structures, the nature and availability in C nutrients, and they ultimately resulted in a progressive accumulation of recalcitrant compounds.     

Dynamics of phosphatase activities in a cork oak litter (Quercus suber L.) following sewage sludge application

As part of a study of the processes involved in litter biodegradation following sewage sludge (SS) addition, the variations over 14 months of phosphatase activities in a cork oak litter (Quercus suber L.) were investigated. A field experimental design was carried out using the litter-bag method on both a fertilized plot receiving SS applications (twice ) and a control plot. Acid (APH) and alkaline (BPH) phosphatases were measured, along with several biotic and abiotic variables potentially involved in the regulation of these enzymes. These included moisture, temperature, pH, water-extractable inorganic P (P_W), culturable heterotrophic bacteria and fungi. Sludge addition had significant effects on all the variables measured. Indeed, sludge increased significantly BPH activities, available P_W, microbial densities (i.e. bacteria and fungi) and pH in the fertilized plot. In contrast, APH activities decreased significantly following sludge addition. As a consequence, the BPH/APH ratio increased markedly and immediately in the fertilized plot, but only after the 1st amendment. Following the 1st preconditioning SS amendment, the 2nd fertilization had fewer effects on biological variables, because of summer dryness. The different properties examined varied significantly with incubation time, and most were significantly related to the seasonal patterns of litter moisture in this Mediterranean forest ecosystem. Hence, sewage sludge application modified the intensity of microbial responses to environmental factors, but biological patterns regulating P turn-over were maintained.     

Organic amendments differ in their effect on microbial biomass and activity and on P pools in alkaline soils

Organic amendments could be used as alternative to inorganic P fertilisers, but a clear understanding of the relationship among type of P amendment, microbial activity and changes in soil P fractions is required to optimise their use. Two P-deficient soils were amended with farmyard manure (FYM), poultry litter (PL) and biogenic waste compost (BWC) at 10 g?dw?kg^?1 soil and incubated for 72 days. Soil samples were collected at days 0, 14, 28, 56 and 72 and analysed for microbial biomass C, N and P, 0.5 M NaHCO₃ extractable P and activity of dehydrogenase and alkaline phosphomonoesterase. Soil P fractions were sequentially extracted in soil samples collected at days 0 and 72. All three amendments increased cumulative CO₂ release, microbial biomass C, N and P and activity of dehydrogenase and alkaline phosphomonoesterase compared to unamended soils. The increase in microbial biomass C and N was highest with PL, whereas the greatest increase in microbial biomass P was induced with FYM. All three biomass indices showed the same temporal pattern, with the highest values on day 14 and the lowest on day 72. All amendments increased 0.5 M NaHCO₃ extractable P concentrations with the smallest increase with BWC and the greatest with FYM, although more P was added with PL than with FYM. Available P concentrations decreased over time in the amended soils. Organic amendments increased the concentration of the labile P pools (resin and NaHCO₃-P) and of NaOH-P, but had little effect on the concentrations of acid-soluble P pools and residual P except for increasing the concentration of organic P in the concentrated HCl pool. Resin P and NaHCO₃-P_i pools decreased over time whereas NaOH-P_i and all organic P pools increased. It is concluded that organic amendments can provide P to plants and can stimulate the build-up of organic P forms in soils which may provide a long-term slow-release P source for plants and soil organisms.     

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.     

Characterization of a Mediterranean litter by 13C CPMAS NMR: relationships between litter depth, enzyme activities and temperature

Organic matter mineralization of forest litter is catalysed by the action of different extracellular enzymes produced by microorganisms. Coupling enzyme activities with data on the general macromolecular structure of organic matter, provided by cross‐polarization magic angle spinning ¹³C nuclear magnetic resonance (¹³C CPMAS NMR), allows researchers new insights into organic matter degradation processes. In this paper, the effect of the temperature of incubation on the degradation processes was evaluated in three distinct layers (OhLn, OhLv and OhLf) of an evergreen oak litter (Quercus ilex L.), located in the Mediterranean area of south‐eastern France. We studied degradation phenomena by a combination of ¹³C CPMAS NMR and microbiological analysis. In order to determine the microbial activity of litter layers, three enzyme activities (laccase, cellulase and butyrate esterase) were measured in a 6‐month mesocosm study. Results showed an increase in the alkyl C to O‐alkyl‐C ratio and an increase of the phenolic C and carboxyl C regions, indicating a preferential degradation of polysaccharides. The aromaticity also increased with litter depth and degradation, and humification processes were more elevated at 30°C. anova showed significant effects (P < 0.001) of increased temperature, depth and time of degradation on microbiological variables. Further information is needed about the variations in temperature and temperature‐litter response and soil functions to link fundamental understanding of carbon stabilization, climate change and global C cycling.     

Non-enzymatic hydrolysis of fluorescein diacetate (FDA) in a Mediterranean oak (Quercus ilex L.) litter

We show the presence of interfering substances when the total microbial activity in litter samples is measured with fluorescein diacetate (FDA), and we propose some methodological modifications to avoid such interference. Three distinct litter layers (the OhLn, the OhLv and the OhLf) of evergreen oak (Quercus ilex L.) were characterized by ¹³C CPMAS NMR and the spectra show that the recalcitrant aromatic and phenolic compounds increase with the degree of degradation of litter. A wide range of sources of interference in the hydrolysis of FDA was found. To understand the origin of this interference, sterilized litter materials (i.e. γ‐irradiated or autoclaved) and a wide range of organic substances (i.e. amino acids, glucose, sorbitol and organic humic acids) were investigated. Insignificant differences on the FDA hydrolysis activity (FDA activity) were found in the γ‐irradiated and non‐irradiated OhLn litter, indicating that γ‐irradiation does not destroy enzymes. Conversely, after heat‐sterilization of litter, samples showed FDA activity corresponding to 60, 34.8 and 30.8% (in the OhLn, the OhLv and the OhLf layers, respectively) of that of control litters. This indicates the presence of non‐enzymatic interfering substances in the FDA assays. As the humification and litter depth increased, hydrolysis of FDA due to interferences decreased, indicating degradation and/or chelation of interfering substances. We hypothesize that lysine, arginine, histidine and cysteine are mainly responsible for the hydrolysis of FDA. We suggest that the use of phosphate buffer (50 mm , pH 7.0) with incubation < 30 minutes, in combination with a temperature between 30 and 40°C, produces insignificant interference in the determination of the final FDA activity in litter samples.     

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].     

Short-term effects of sewage sludge application on phosphatase activities and available P fractions in Mediterranean soils

The aim of this study was to investigate factors regulating phosphatase activities in Mediterranean soils subjected to sewage sludge applications. Soils originating from calcareous and siliceous mineral parent materials were amended with aerobically digested sewage sludge, with or without physico-chemical treatment by ferric chloride. Sludge amendments, ranging from 6.2 to 10 g kg⁻¹ soil, were carried out in order to provide soil with a P total quantity equivalent to 0.5 g P₂O₅ per kg of soil. Bacterial density, phosphatase activities (i.e. acid and alkaline phosphomonoesterases and phosphodiesterases) and available P (i.e. P Olsen and P water) were measured after 25 and 87 days of incubation. Results showed significant effects of sewage sludge application and incubation period. Sewage sludge effect resulted in an increase in phosphatase activities, microbial density and available P. Incubation period increased available P while decreasing phosphatase activities. This study also revealed that the origin of sludge and its chemical characteristics may show different effects on certain variables such as phosphodiesterases or bacterial density, whereas mineral parent materials of soils did not show any significant effects.     

Effects of organic and mineral amendments on available P and phosphatase activities in a degraded Mediterranean soil under short-term incubation experiment

The aim of this study was to determine the effects of mineral and organic-P-fertilizers on soil P availability, bacteria densities and phosphatase activities, in a degraded Mediterranean soil characterized by low level in soil organic matter and nutrients. A typical degraded Mediterranean soil, originating from a siliceous mineral parent material, was amended with different organic or mineral P-sources: aerobically digested sewage sludge (SS), with or without physico-chemical treatment by ferric chloride; sewage sludge compost (SSC); Na or K mineral P-salts (Pi-salts). All the amendments were carried out in order to provide soil with a P total quantity equivalent to 0.5 g P₂O₅/kg of soil. Bacterial density, phosphatase activities (i.e. acid (APH) and alkaline (BPH) phosphomonoesterases and phosphodiesterases), BPH/APH ratio, and available P (P Olsen) were measured after 25 and 87 days of incubation. Results showed that all the P-sources used to fertilize soil during this study resulted in significant increase in P concentration. However, different responses in phosphatase activities and bacterial densities were obtained with regards to the amendment applied to soil. Indeed, it appeared clearly that sewage sludge (SS) considerably stimulated soil biological activity, and more especially the different kinds of phosphatases involved in P mineralization and P turn-over. On the contrary, sewage sludge compost (SSC) as well as P-salts amendments did not affected these parameters in most cases. Results showed also that the incubation time influenced almost all the biological and chemical parameters investigated during this study. As a consequence, P availability was considerably improved in the amended soils between the two sampling dates.     

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.     

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.     

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.     

Successional changes in microbial biomass,respiration and nutrient status during litter decomposition in an aspen and pine forest

Microbial biomass, microbial respiration, metabolic quotient (qCO₂), C_mic/C_org ratio and nutrient status of the microflora was investigated in different layers of an aspen (Populus tremuloides Michx.) and pine forest (Pinus contorta Loud.) in southwest Alberta, Canada. Changes in these parameters with soil depth were assumed to reflect successional changes in aging litter materials. The microbial nutrient status was investigated by analysing the respiratory response of glucose and nutrient (N and P) supplemented microorganisms. A strong decline in qCO₂ with soil depth indicated a more efficient C use by microorganisms in later stages of decay in both forests. C_mic/C_org ratio also declined in the aspen forest with soil depth but in the pine forest it was at a maximum in the mineral soil layer. Microbial nutrient status in aspen leaf litter and pine needle litter indicated N limitation or high N demand, but changes in microbial nutrient status with soil depth differed strongly between both forests. In the aspen forest N deficiency appeared to decline in later stages of decay whereas P deficiency increased. In contrast, in the pine forest microbial growth was restricted mainly by N availability in each of the layers. Analysis of the respiratory response of CNP-supplemented microorganisms indicated that growth ability of microorganisms is related to the fungal-bacterial ratio.     

Differing nitrogen use strategies of two tropical rainforest late successional tree species in French Guiana: Evidence from 15N natural abundance and microbial activities

Previous studies in lowland tropical rainforests of French Guiana showed that, among non-N₂-fixing trees, two groups of late successional species contrasting in their leaf ¹⁵N natural abundance coexist, suggesting two different main ways of nitrogen acquisition. Two abundant late-successional species typically co-occurring in rainforests in French Guiana, namely Eperua falcata and Dicorynia guianensis, were chosen as representative of each group. Stable isotope techniques and measurements of potentials of microbial N transformation were performed to assess to what extent leaf ¹⁵N natural abundance of these species could be related to (i) δ¹⁵N signatures of soil mineral N sources and (ii) the capacity of soil to express nitrification and denitrification (both processes being directly involved in the balance between NH₄⁺ and NO₃^–). Soil δ¹⁵N-NH₄⁺ was roughly similar to leaf δ¹⁵N of D. guianensis (around 3.5‰), suggesting a preferential use of NH₄⁺, whereas in E. falcata, leaf δ¹⁵N values were closer to root δ¹⁵N-NO₃^? values (0.2 and ?2.0‰, respectively), suggesting a preferential use of NO₃^?. These differences in N source utilization were not accompanied by differences in availability in soil NO₃^? or in intensity of microbial functions responsible for soil N mineral evolution. However, (i) under both tree species, these functions showed clear spatial partitioning, with denitrification occurring potentially in soil and nitrification in the litter layer, and (ii) E. falcata fine roots colonized the litter layer much more strongly than D. guianensis fine roots. This strongly suggests that (i) the contrasted leaf δ¹⁵N values found in the two late-successional species reveal distinct N acquisition strategies and (ii) the ability of roots to predominantly exploit the litter layer (E. falcata) or the soil (D. guianensis) may constitute an important explanation of the observed differences. A complementarity between tree species, based on mineral N resource partitioning (itself resulting from a spatially structured location of the microbial functions responsible for the balance between NH₄⁺ and NO₃^–), can thus be supposed.     

Humus macro-morphology and soil microbial community changes along a 130-yr-old Fagus sylvatica chronosequence

We aimed to characterize humus macro-morphology and the associated soil microbial community within the unmodified litter (OL), the fragmented and humified layers (FH) and the organo-mineral (A) layer along a beech (Fagus sylvatica L.) forest chronosequence with four stand age-classes (15-, 65-, 95-, 130-yr-old) in Normandy, France. Humus macro-morphology was described with 36 quantitative and semi-quantitative variables. We measured microbial biomass N (N_mic), microbial N quotient (N_mic-to-N_t), fungal ergosterol, bacterial and fungal DNA using 16S and 18S rDNA real-time qPCR and evaluated the potential metabolic profile of heterotrophic bacteria within each soil layer and stand age-class. The log-transform ergosterol/fungal DNA ratio (EFR index) was used as an indicator related to active fungal biomass and the fungal/bacterial (F/B) ratio was calculated from qPCR results. There was a shift from mull (mainly dysmull) to moder humus forms along the chronosequence. While the N_mic did not change significantly, the N_mic-to-N_t decreased along the chronosequence in the OL layer. Ergosterol content increased in FH and A layers and the F/B ratio increased in the FH layer with increasing beech forest age. The EFR index was significantly higher in the OL and A layers of the oldest stands, whereas the highest EFR index in the FH layer occurred in the 15-yr-old stands. The functional diversity of heterotrophic bacteria was greater within OL and FH layers of 130-yr-old stands, but highest in the A layer of 15-yr-old stands while the Average Well Color Development remained stable for all soil layers. We found significant correlations between macro-morphology and microbial variables, especially between FH-based morphology and fungal biomass. Our main results are that beech forest maturation is accompanied by (1) an increase in fungal biomass in the FH layers and, (2) an increase in heterotrophic bacteria functional diversity in the organic layers. We have identified key macro-morphology variables that are good predictors of the structural and functional profile of the soil microbial community during beech forest development.     

Vertical stratification and trophic interactions among organisms of a soil decomposer food web – a field experiment using 15N as a tool

In this field study, we explored the spatial segregation between the litter- and humus-inhabiting organisms of the detrital food web using ¹⁵N-isotope technique. The study was established in 11 × 11 m plots fertilized with ¹⁵N-labelled urea. Ten years after urea application, soil samples were taken, both from the litter layer and the combined F+H layer. The samples were analysed for N content and the proportion of ¹⁵N in (i) the residual organic matter in the litter and F+H layer (excluding microbes), (ii) microbial biomass, and (iii) various feeding guilds of soil fauna. The basal resource, soil microbes, and the fauna were more enriched with ¹⁵N in the F+H layer than in the litter layer. In the litter layer, the ¹⁵N enrichment of the expected food source equalled the one of the consumers, whereas in the F+H layer all trophic groups, except microbes and small microbi-detritivores, showed a significantly lower ¹⁵N enrichment than their expected food source. The results indicate that large and mobile humus-inhabiting decomposers exploit the overlying litter layer as a feeding site, whereas the feeding of the more sedentary smaller organisms is restricted to the humus layer.     

Changes in soil chemical and microbial properties after a wildfire in a tropical rainforest in Sabah, Malaysia

Changes in soil caused by drought and wildfire in a Dipterocarp rainforest in Sabah, Malaysia were assessed by phosphorus fractionation, extractable nitrogen and nutrient limited respiration kinetics (after addition of glucose+N or P). Fire increased the concentration of total phosphorus (P) in the litter layer (per ha and per dry soil) by raising the 0.2 M NaOH extractable-P. In the soil organic layer, membrane exchangeable P was reduced by fire while 1.0 M HCl extractable-P, and 0.5 M NaHCO₃ extractable-P increased. Microbially available P increased after the fire and was most closely related to NaOH extractable-P that has been considered available to plants only over long time-scales. Total nitrogen (N) increased in the litter layer (per ha and per dry soil) due to post-fire litter fall, while the NO₃⁻ increased up to 10-fold down to the 10 cm mineral soil. In contrast, the microbially available N decreased by 50%. Basal respiration and substrate-induced respiration increased in the litter layer and decreased in the organic horizon (per dry soil and per organic matter). P limited microbial growth resulted in a slow and non-exponential increase in respiration, presumably reflecting the P-fixing nature of the soils, while N limitation resulted in a fast exponential increase. However, higher respiration rates were eventually achieved under P limitation than under N limitation.     

Acquisition of microbial communities and enhanced availability of soil nutrients by the isopod Porcellio scaber (Latr.) (Isopoda: Oniscidea)

 In microcosm experiments Porcellio scaber increased litter disappearance of oak and alder litter. Alder litter disappeared at more than twice the rate of oak litter. Soil texture did not influence the disappearance of oak litter; however, disappearance of alder litter was enhanced on silt rather than on sand. P. scaber enhanced microbial communities (i.e. microbial respiration and microbial biomass) on both silt and sand when feeding on either alder or oak. Overall, microbial respiration increased 10-fold when isopods fed on oak litter on sand and 20-fold when isopods fed on alder litter on sand. On silt, the initially high microbial respiration remained constant when isopods fed on oak and doubled when they fed on alder litter. In all treatments without P. scaber there was a decrease in microbial respiration over the 12-week experimental period. The availability of macronutrients (C_org, N_tot, P, K, Mg, Ca) in the topsoil was increased when P. scaber fed on alder litter but less pronounced when P. scaber fed on oak litter. Using sand as a substrate, there was an apparent increase only for C_org, Mg and Ca; on silt, increases in C_org, N_tot and P were measured. Under field conditions the contribution of P. scaber to nutrient fluxes will be higher on sand than on silt. Received: 1 July 1999     

Phosphatase activities in soil after repeated untreated and alum-treated poultry litter applications

Repeated additions of untreated and aluminum sulfate (alum)-treated poultry litter to soil affect ecology and consequent nutrient dynamics. The objective of this study was to determine how repeated annual poultry litter additions affected phosphatase activities in concert with changes in soil phosphorus (P). Field plots were amended annually since 1995 with either 2.24 or 8.96 Mg ha⁻¹ alum-treated (AL-1 or AL-4, respectively) or untreated poultry litter (PL-1 or PL-4, respectively) or equivalent rates by N content of ammonium nitrate (AN-1 or AN-4, respectively). Soil pH, total C (C_Tot), microbial biomass C, double-stranded deoxyribonucleic acid (dsDNA), Mehlich-III P (M3P), water-soluble P (WSP), and acid and alkaline phosphatase activities were measured before and 10 days, 1 month, and 6 months after fertilizer applications in 2003 and 2004. M3P and acid phosphatase activities were higher in AL-4 soil than in all other treatments. Higher WSP in the untreated compared to the alum-treated litter resulted in higher WSP in the soil amended with untreated litter. At the same time, alkaline phosphatase activities were significantly higher in PL-4 compared to AL-4. In contrast, alkaline phosphatase activities were significantly lower in AN-4, the treatment with the lowest pH. Additionally, alkaline phosphatase activities expressed per unit C_Tot or dsDNA remained significantly greater in PL-4 and significantly lower in AN-4, than all other treatments. Thus, some factor beyond an increase in soil C or microbial biomass contributed to elevated alkaline phosphatase activities in PL-4, despite increased WSP in the treatment receiving the high rate of untreated litter.