Earthworm and microbe response to litter and soils of tropical forest plantations with contrasting C:N:P stoichiometric ratios

1

The stoichiometry of resources is increasingly acknowledged as a major control of consumer activity and abundance. Chemical properties of litter, the main source of food for decomposers, are likely to be important drivers of decomposer activity.

2

Theory predicts a high control of resource stoichiometry on the abundance of consumer organisms that maintain strict homeostasis, due to costs associated with the regulation of nutrient balance in their body tissue. Decomposer efforts in nutrient acquisition should be related to imbalances in resource stoichiometry.

3

A 21 year old experimental plantation of monospecific plots of trees with leaves of contrasting chemistry was used to test four hypotheses: (i) soil and litter nutrient stoichiometry (C, N, P) are linked; (ii); soil enzyme activity ratios and stoichiometry are linked; (iii) earthworms’ tissue stoichiometry does not depend on soil and litter stoichiometry (homeostasis); (iv) earthworm density is dependent upon phosphorus availability, the most limiting nutrient in soils at this site, and, to a lesser extent, to nitrogen availability.

4

We found (i) no relationship between litter and soil stoichiometry, (ii) microbial activity was linked to soil stoichiometry, (iii) earthworms showed strict homeostasis in their tissue and (iv) earthworm abundance increased with P availability.

5

We discuss the mechanisms that might lead to these patterns.

     

Cost-effectiveness of silvicultural measures to increase substrate availability for red-listed wood-living organisms in Norway spruce forests

It is important that measures to maintain biodiversity are taken in a way that is cost-effective for the landowner. We analyzed the cost-effectiveness of silvicultural measures that aim at increasing the substrate availability for red-listed (species that are threatened, near threatened or where species probably are threatened but data is deficient) saproxylic (wood-inhabiting) organisms. We modelled stands of Norway spruce (Picea abies) in three regions of Sweden by using computer simulations and a database with substrate requirements of saproxylic beetles and cryptogams on the Swedish Red-List. Conclusions concerning cost-effectiveness of silvicultural measures depend on the extinction thresholds of the species they are intended to conserve; measures that generate only small amounts of coarse woody debris (CWD) may provide too little substrate to be useful for species with high extinction thresholds. In northern Sweden, forestland is relatively inexpensive, so a cost-effective strategy to increase the amount of spruce CWD was to set aside more forests as reserves. In central and southern Sweden, more emphasis should instead be given to increasing the amount of CWD in the managed forest. The regulations by the Forest Stewardship Council (FSC) could be made more cost-effective by prescribing creation of more high stumps and retention of larger amounts of naturally dying trees. Large-sized CWD, CWD from slow-growing trees, and CWD in late decay stages are substrate types that were particularly rare in managed forest in relation to unmanaged forests. Manual soil scarification and retention of living trees are measures that can increase the proportion of these underrepresented CWD types.     

Uptake of Sr from microcline and biotite by ectomycorrhizal fungi in a Norway spruce forest

Ectomycorrhizal (EM) fungi growing in symbiosis with tree seedlings have been found in laboratory experiments to stimulate weathering and the uptake of nutrients from silicate minerals. In the present study, we used the natural abundance of strontium isotopes to confirm that these fungi obtain strontium from biotite and microcline under field conditions. Minerals enriched in radiogenic ⁸⁷Sr were introduced into fungal in-growth mesh bags and placed under a 5-10 cm thick humus layer developed on boulders in a Norway spruce (Picea abies (L.) Karst) forest in south-western Sweden. EM root tips were sampled above the mesh bags and the ⁸⁷Sr/⁸⁶Sr ratio was used to calculate the fraction of Sr in the root tips that originated from the minerals. In the EM root tips sampled above the mesh bags containing the different minerals, 1.5% of the Sr originated from biotite and 6.4% from microcline. The amount of Sr was more than 300 times higher in the mesh bags containing microcline than in those containing biotite, indicating that proportionally more Sr was released from the biotite. This study demonstrates that EM fungi have the potential to take up measurable amounts of nutrients, such as Ca and K, from microcline and biotite in the field.     

Effect of liming on some chemical,biochemical, and microbiological properties of acid soils under spruce (Picea abies L.)

Summary Soil pH, total organic C, total N, exchangeable Al, available P, CO₂ evolution, microbial biomass C and N, phosphatase and dehydrogenase activities were determined in acid soils sampled under spruce subjected to acid deposition, before and after liming. A slight decrease in pH values was observed from the edge of a tree canopy to the base of the trunk in acid soils. Liming drastically reduced exchangeable Al and increased CO₂ evolution, microbial biomass, and the metabolic quotient. The microbial biomass C to total organic C ratio increased after liming but did not reach 2%, the average value considered valid in soils where the C content is in equilibrium, that is when C inputs are equal to C outputs. The microbial biomass C:N ratio decreased after liming, thus indicating that bacteria became predominant over fungi when soil acidity decreased. Dehydrogenase activity but not phosphatase activity was increased by liming. The decrease in phosphatase activity was not completely related to the increase in available P, but was also dependent on microbial growth and the decrease in acid phosphatase, the predominant component of acid soils.     

Arbuscular mycorrhizal fungi contribute to C and N enrichment of soil organic matter in forest soils

Increasing evidence suggests that accretion of microbial turnover products is an important driver for isotopic carbon (C) and nitrogen (N) enrichment of soil organic matter (SOM). However, the exact contribution of arbuscular mycorrhizal fungi (AMF) to soil isotopic patterns remains unknown. In this study, we compared ¹³C and ¹⁵N patterns of glomalin-related soil protein (GRSP), which includes a main fraction derived from AMF, litter, and bulk soil in four temperate rainforests. GRSP was an abundant C and N pool in these forest soils, showing significant ¹³C and ¹⁵N enrichment relative to litter and bulk soil. Hence, cumulative accumulation of recalcitrant AMF turnover products in the soil profile likely contributes to ¹³C and ¹⁵N enrichment in forest soils. Further research on the relationship between GRSP and AMF should clarify the exact extent of this process.     

Effects of acidification and liming on feeding groups of nematodes in coniferous forest soils

Summary Nematodes were sampled in untreated, acidified, and limed plots in a Norway spruce (Fexboda) and a Scots pine (Norrliden) stand. At Fexboda, the total number of nematodes was significantly reduced after the acidification. This reduction was probably due to a shock effect, because the samples were taken only 5 months after an application of 200 kg H₂SO₄ ha^-1 to the forest floor. However, the root/fungal-feeding Aphelenchoides was not reduced, probably because it is more tolerant of high acid concentrations than most other nematodes. At Norrliden, where the samples were taken 7 years after the last application of H₂SO₄, no significant differences were found between the acidified and untreated plots. If the treatment with H₂SO₄ caused similar effects as at Fexboda, the results indicate a recovery of the nematode populations. Decreased predation from lumbricids rather than a recovery of microfloral populations probably allowed this recovery. No marked effect of lime, spread 2 (Fexboda) and 12 years (Norrliden) before the sampling on the numbers of any of the nematode feeding groups was found. This correlated with almost no change in bacterial biomass after liming, while the active fraction of fungal hyphae was unaffected by liming at Fexboda and reduced by liming at Norrliden. A tendency for decreasing numbers of all nematode feeding groups in the limed plots at Norrliden coincided with increasing numbers of lumbricids.     

Saprotrophic fungi transform organic phosphorus from spruce needle litter

Fungal decomposition of and phosphorus transformation from spruce litter needles (Picea abies) were simulated in systems containing litter needles inoculated with individual saprotrophic fungal strains and their mixtures. Fungal strains of Setulipes androsaceus (L.) Antonín, Chalara longipes (Preus) Cooke, Ceuthospora pinastri (Fr.) Höhn., Mollisia minutella (Sacc.) Rehm, Scleroconidioma sphagnicola Tsuneda, Currah & Thormann and an unknown strain NK11 were used as representatives of autochthonous mycoflora. Systems were incubated for 5.5 months in laboratory conditions. Fungal colonization in systems and competition among strains were assessed using the reisolation of fungi from individual needles. After incubation, needles were extracted with NaOH and extracts were analysed using ³¹P nuclear magnetic resonance spectroscopy (NMR). Needle decomposition was determined based on the decrease in C:N ratio. Systems inoculated with the basidiomycete S. androsaceus revealed substantial decrease in C:N ratio (from 25.8 to 11.3) while the effect of ascomycetes on the C:N ratio was negligible. We suppose that tested strains of saprotrophic ascomycetes did not participate substantially in litter decomposition, but were directly involved in phosphorus transformation and together with S. androsaceus could transform orthophosphate monoesters and diesters from spruce litter needles into diphosphates, polyphosphates and phosphonates. These transformations seem to be typical for saprotrophic fungi involved in litter needle decomposition, although the proportion of individual phosphorus forms differed among studied fungal strains. Phosphonate presence in needles after fungal inoculation is of special interest because no previous investigation recorded phosphonate synthesis and accumulation by fungi. Our results confirmed that the ³¹P NMR spectroscopy is an excellent instrumental method for studying transformations of soil organic phosphorus during plant litter decomposition. We suggest that polyphosphate production by S. androsaceus may contribute to the phosphorus cycle in forest ecosystems because this fungus is a frequent litter colonizer that substantially participates in decomposition.     

Impact of faunal complexity on microbial biomass and N turnover in field mesocosms from a spruce forest soil

In a field study using soil mesocosms in an acid spruce forest soil we investigated the effects of mesofauna and macrofauna on microbial biomass, dissolved organic matter, and N cycling. Intact soil monoliths were taken from the ground, defaunated by deep-freezing, and wrapped in nets of various mesh-sizes to control re-immigration of different faunal size-classes. The monoliths were then replanted in the field. Three treatments of mesocosms were prepared: (1) with only microbiota, (2) microbiota and mesofauna, and (3) microbiota, mesofauna, and macrofauna (= complex fauna). After 8 months of exposure the mesocosms and the unmanipulated control plots (treatment 4) were destructively sampled. We estimated microbial biomass by substrate-induced respiration and the chloroform fumigation-extraction method. N cycling was measured by monitoring microbial N mineralization, the NH _inf4 ^sup+ content, and selected amino acids and the activities of protease, urease, and deaminase. The results from the L/F layer showed that the pool of the microbial biomass was not changed by the activity of the mesofauna. However, the mesofauna and macrofauna together enhanced SIR. An increase in microbial N mineralization was only observed in treatment 3 (microbiota + complex fauna). Protease activity and NH _inf4 ^sup+ content increased in treatments 2 (microbiota + mesofauna) and 3 (microbiota + complex fauna). The complex fauna induced a soil pH increase in treatment 3 as opposed to treatment 1 and the control. This increase was presumably due to excretory NH _inf4 ^sup+ . Principal component analysis revealed that the complex fauna in treatment 3 caused a significantly higher N turnover per unit of microbial biomass.     

Microbial biomass and C and N transformations in forest floors under European beech, sessile oak, Norway spruce and Douglas-fir at four temperate forest sites

The purpose of this research was to compare soil chemistry, microbially mediated carbon (C) and nitrogen (N) transformations and microbial biomass in forest floors under European beech (Fagus sylvatica L.), sessile oak (Quercus petraea (Mattuschka) Lieblein), Norway spruce (Picea abies (L.) Karst) and Douglas-fir (Pseudotsuga menziesii (Mirbel) Franco) at four study sites. We measured soil chemical characteristics, net N mineralization, potential and relative nitrification, basal respiration, microbial and metabolic quotient and microbial biomass C and N under monoculture stands at all sites (one mixed stand). Tree species affected soil chemistry, microbial activities and biomass, but these effects varied between sites. Our results indicated that the effect of tree species on net N mineralization was likely to be mediated through their effect on soil microbial biomass, reflecting their influence on organic matter content and carbon availability. Differences in potential nitrification and relative nitrification might be related to the presence of ground vegetation through its influence on soil NH₄ and labile C availability. Our findings highlight the need to study the effects of tree species on microbial activities at several sites to elucidate complex N cycle interactions between tree species, ground vegetation, soil characteristics and microbial processes.     

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.     

A possible role for saprotrophic microfungi in the N nutrition of ectomycorrhizal Pinus resinosa

We determined whether Pinus resinosa, selected ectomycorrhizal and saprotrophic microfungi have access to various organic nitrogen sources commonly found in the forest. Vector analysis demonstrated nitrogen limitation of the P. resinosa in the plantation from which most of the fungi were isolated, establishing this study's relevance. Nonmycorrhizal P. resinosa seedlings did not absorb significant N from amino acids. The ectomycorrhizal fungi, including Pisolithus tinctorius, Suillus intermedius and Tylopilus felleus, obtained substantial N from amino acids, a limited amount of N from glucosamine, and essentially no N from protein-tannin complex. In contrast, Penicillium and Trichoderma readily acquired N from protein-tannin and glucosamine. Thus, there was an increasing ability to obtain N from complex organic N sources from plant to ectomycorrhizal fungi to saprotrophic fungi. Furthermore, N mineralization from an organic N source by Penicillium depended on the C:N ratio. We conclude that acquisition of relatively simple organic N sources by P. resinosa is likely to be largely indirect via ectomycorrhizal fungi, and that more complex organic N sources may become accessible to ectomycorrhizal fungi (and thus possibly their host plants) following mineralization by saprotrophic fungi such as Penicillium or Trichoderma when C:N ratios are sufficiently low.     

Soil microbial biomass and activity under a potato crop fertilised with N with and without C

Summary A range of soil microbiological parameters were measured at intervals throughout the growing season of a potato crop. Treatments applied to the soil at sowing were zero N fertilisation of N fertilisation at 120 kg N ha^–1, either alone or supplemented with straw or sucrose at 1200 kg C ha^–1. C and N flushes determined by fumigation-incubation and fumigation-extraction, and substrate-induced respiration, were measured as indicators of microbial biomass. Microbial activity was measured as respiration (CO₂ production) and dehydrogenase activity (formazan production). The greatest effects were obtained from the addition of N plus sucrose. Both biomass size and activity were significantly stimulated for up to 25 days after incorporation, with the magnitude of the effects consistently diminishing over time. By 125 days after planting, there was no detectable legacy from any of the treatmentson any of the biomass parameters that were measured, and all values had reverted to those prevalent at planting. There was no consistent effect from adding N, either alone or supplemented with straw, on any of the biomass parameters. There was no evidence for crop-induced stimulation of the biomass. The experiment demonstrates that biomass is only influenced where the quantity, quality, and rate of incorporation of C into the soil is appropriate, in this case, only by adding C as a pulse of sucrose.     

Explaining temporal variation in soil CO2 efflux in a mature spruce forest in Southern Germany

An open dynamic chamber system was used to measure the soil CO₂ efflux intensively and continuously throughout a growing season in a mature spruce forest (Picea abies) in Southern Germany. The resulting data set contained a large amount of temporally highly resolved information on the variation in soil CO₂ efflux together with environmental variables. Based on this background, the dependencies of the soil CO₂ efflux rate on the controlling environmental factors were analysed in-depth. Of the abiotic factors, soil temperature alone explained 72% of the variation in the efflux rate, and including soil water content (SWC) as an additional variable increased the explained variance to about 83%. Between April and December, average rates ranged from 0.43 to 5.15 μmol CO₂ m⁻² s⁻¹ (in November and July, respectively) with diurnal variations of up to 50% throughout the experiment. The variability in wind speed above the forest floor influenced the CO₂ efflux rates for measuring locations with a litter layer of relatively low bulk density (and hence relatively high proportions of pore spaces). For the temporal integration of flux rates for time scales of hours to days, however, wind velocities were of no effect, reflecting the fact that wind forcing acts on the transport, but not the production of CO₂ in the soil. The variation in both the magnitude of the basal respiration rate and the temperature sensitivity throughout the growing season was only moderate (coefficient of variation of 15 and 25%, respectively). Soil water limitation of the CO₂ production in the soil could be best explained by a reduction in the temperature-insensitive basal respiration rate, with no discernible effect on the temperature sensitivity. Using a soil CO₂ efflux model with soil temperature and SWC as driving variables, it was possible to calculate the annual soil CO₂ efflux for four consecutive years for which meteorological data were available. These simulations indicate an average efflux sum of 560 g C m⁻² yr⁻¹ (SE=22 g C m⁻² yr⁻¹). An alternative model derived from the same data but using temperature alone as a driver over-estimated the annual flux sum by about 7% and showed less inter-annual variability. Given a likely shift in precipitation patterns alongside temperature changes under projected global change scenarios, these results demonstrate the necessity to include soil moisture in models that calculate the evolution of CO₂ from temperate forest soils.     

Hot water extractable C and N in relation to microbiological properties of soils under beech forests

Hot water extraction is sometimes recommended as an easy method to estimate the readily mineralizable fractions of total C (C_t) and total N (N_t) in arable soils. However, the usefulness of this method for forest soils has not been adequately studied. The objectives of this study were to relate the hot water extractable C (C_hw) and N (N_hw) to microbiological and chemical properties of the forest soils under beech (Fagus sylvatica L.) stands and to test the ability of near infrared spectroscopy (NIRS) to predict chemical and microbial properties of these soils. Soils differing in humus type, soil type and soil texture were collected from five locations and five depths. In all soils the amount of C_hw was higher than the microbial biomass C (C_mic) indicating that a considerable part of C_hw was of non-microbial origin. The amount of C_hw in mineral soil correlated significantly (r =–0.30–0.53) with C_mic, basal respiration (BAS) and C_t/N_t ratio but was not related to C_mic/C_t ratio. The amount of N_hw was correlated with C_mic, BAS, C_mic/C_t ratio, and C_t/N_t ratio (r =–0.59–0.78). However, C_t and N_t values showed better relationships (r =–0.42–0.88) with all the parameters, indicating no advantage in using C_hw and N_hw in forest soils. NIRS predicted satisfactorily C_t, N_t, C_hw, N_hw, C_mic, C_mic/C_t ratio and BAS in the mineral soils [the regression coefficients (a) of linear regression (measured against predicted values) ranged from 0.84 to 1.17 and the correlation coefficients (r) ranged from 0.86 to 0.94] indicating the applicability of NIRS to estimate these properties.     

Influence of ectomycorrhizal mat soils on lignin and cellulose degradation

Summary The ectomycorrhizal fungus Hysterangium setchellii (Fisher) forms extensive hyphal mats at the soillitter interface with the roots of the host tree Douglas fir Pseudotsuga menziesii [(Mirb.) Franco]. Microbial biomass, and lignin and cellulose decomposition rates were measured seasonally for 1 year, using ¹⁴C techniques in ectomycorrhizal mat soils and adjacent non-mat soils in a second-growth Douglas fir forest. The microbial biomass and cellulose degradation rates were 3–6 times higher in ectomycorrhizal mat soils than in adjacent nonmat soils. Lignin degradation rates were higher in ectomycorrhizal mat soils than adjacent non-mat soils. Our data suggest that the ectomycorrhizal fungus H. setchellii provides a microenvironment with increased microbiological activity which results in faster lignin and cellulose decomposition.     

Turnover of grain legume N rhizodeposits and effect of rhizodeposition on the turnover of crop residues

The turnover of N derived from rhizodeposition of faba bean (Vicia faba L.), pea (Pisum sativum L.) and white lupin (Lupinus albus L.) and the effects of the rhizodeposition on the subsequent C and N turnover of its crop residues were investigated in an incubation experiment (168 days, 15 °C). A sandy loam soil for the experiment was either stored at 6 °C or planted with the respective grain legume in pots. Legumes were in situ ¹⁵N stem labelled during growth and visible roots were removed at maturity. The remaining plant-derived N in soil was defined as N rhizodeposition. In the experiment the turnover of C and N was compared in soils with and without previous growth of three legumes and with and without incorporation of crop residues. After 168 days, 21% (lupin), 26% (faba bean) and 27% (pea) of rhizodeposition N was mineralised in the treatments without crop residues. A smaller amount of 15–17% was present as microbial biomass and between 30 and 55% of mineralised rhizodeposition N was present as microbial residue pool, which consists of microbial exoenzymes, mucous substances and dead microbial biomass. The effect of rhizodeposition on the C and N turnover of crop residues was inconsistent. Rhizodeposition increased the crop residue C mineralisation only in the lupin treatment; a similar pattern was found for microbial C, whereas the microbial N was increased by rhizodeposition in all treatments. The recovery of residual ¹⁵N in the microbial and mineral N pool was similar between the treatments containing only labelled crop residues and labelled crop residues + labelled rhizodeposits. This indicates a similar decomposability of both rhizodeposition N and crop residue N and may be attributable to an immobilisation of both N sources (rhizodeposits and crop residues) as microbial residues and a subsequent remineralisation mainly from this pool.Abbreviations C or N_dec C or N decomposed from residues - C or N_mic microbial C or N - C or N_micres microbial residue C or N - C or N_min mineralised C or N - C or N_input added C or N as crop residues and/or rhizodeposits - dfr derived from residues - dfR derived from rhizodeposition - Ndfr N derived from residues - NdfR N derived from rhizodeposition - N_loss losses of N derived from residues - SOM soil organic matter - WHC water holding capacity     

Drying and rewetting effects on C and N mineralization and microbial activity in surface and subsurface California grassland soils

Rewetting a dry soil has long been known to cause a burst of respiration (the “Birch Effect”). Hypothesized mechanisms for this involve: (1) release of cellular materials as a result of the rapid increase in water potential stress and (2) stimulating C-supply to microbes via physical processes. The balance of these factors is still not well understood, particularly in the contexts of multiple dry/wet cycles and of how resource and stress patterns vary through the soil profile. We evaluated the effects of multiple dry/wet cycles on surface and subsurface soils from a California annual grassland. Treatments included 4, 6, and 12 cycles that varied the length of the drying period between rewetting events. Respiration was monitored after each wetting event while extractable C and N, microbial biomass, and microbial activity were assayed initially, after the first rewetting event, and at the end of the experiment. Initially, microbial biomass and activity (respiration, dehydrogenase, and N mineralization) in subsurface soils were ca. 10% and 20% of surface soil levels. After multiple cycles, however, subsurface soil microbial biomass and activity were enhanced by up to 8-fold, even in comparison to the constantly moist treatment. By comparison, surface soil microbial biomass and activity were either moderately (i.e. 1.5 times increase) or not affected by wetting and drying. Drying and rewetting led to a cascade of responses (soluble C release, biomass growth, and enhanced activity) that mobilized and metabolized otherwise unavailable soil carbon, particularly in subsurface soils.     

Microbial activities in forest soils exposed to chronic depositions from a lignite power plant

Atmospheric emissions of fly ash and SO₂ from lignite-fired power plants strongly affect large forest areas in Germany. The impact of different deposition loads on the microbial biomass and enzyme activities was studied at three forest sites (Picea abies (L.) Karst.) along an emission gradient of 3, 6, and 15 km downwind of a coal-fired power plant (sites Ia, II, and III, respectively), representing high, moderate and low emission rates. An additional site (site Ib) at a distance of 3 km from the power plant was chosen to study the influence of forest type on microbial parameters in coniferous forest soils under fly ash and SO₂ emissions. Soil microbial biomass C and N, CO₂ evolved and activities of l-asparaginase, l-glutaminase, β -glucosidase, acid phosphatase and arylsulfatase (expressed on dry soil and organic C basis) were determined in the forest floor (L, Of and Oh horizon) and mineral top soil (0-10 cm). The emission-induced increases in ferromagnetic susceptibility, soil pH, concentrations of mobile (NH₄NO₃ extractable) Cd, Cr, and Ni, effective cation exchange capacity and base saturation in the humus layer along the 15 km long transect significantly (P<0.05) reflected the effect of past depositions of alkaline fly ash. Soil microbial and biochemical parameters were significantly (P<0.05) affected by chronic fly ash depositions. The effect of forest type (i.e. comparison of sites Ia and Ib) on the studied parameters was generally dominated by the deposition effect. Alkaline depositions significantly (P<0.05) decreased the microbial biomass C and N, microbial biomass C-to-N ratios and microbial biomass C-to-organic C ratios. Microbial respiration, metabolic quotient (qCO₂) and the activities of l-asparaginase, l-glutaminase, β-glucosidase, acid phosphatase and arylsulfatase were increased by long-term depositions from the power plants. Acid phosphatase had the highest specific (enzyme activities expressed per unit organic C) activity values among the enzymes studied and arylsulfatase the lowest. The responses of the microbial biomass and soil respiration data to different atmospheric deposition loads were mainly controlled by the content of organic C and cation exchange capacity, while those of enzyme activities were governed by the soil pH and concentrations of mobile heavy metals. We concluded that chronic fly ash depositions decrease litter decomposition by influencing specific microbial and enzymatic processes in forest soils.     

Nitrous oxide emissions as influenced by amendment of plant residues with different C:N ratios

To investigate the influence of plant residues decomposition on N₂O emission, laboratory incubations were carried out for a period of 21 days using urea and five plant residues with a wide range of C:N ratios from 8 to 118. Incorporation of plant residues enhanced N₂O and CO₂ emissions. The two gas fluxes were significantly correlated (R²=0.775, p<0.001). Cumulative emissions of N₂O and CO₂ were negatively correlated with the C:N ratio in plant residues (R²=0.783 and 0.986 for N₂O, and 0.854 for CO₂, respectively). A negative relationship between the N₂O-N/NO₃⁻-N ratio and the C:N ratio was observed (R²=0.867) when residue plus urea was added. We calculated the changes in dissolved organic C (DOC) and the relevant changes in N₂O emission. The incorporation of residues increased DOC when compared with the control, while the incorporation of residue plus urea decreased DOC. Cumulative emissions of N₂O and CO₂ were positively correlated with DOC concentration measured at the end of the incubation. In addition, the N₂O emission fraction, defined as N₂O-N emissions per unit N input, was not found to be a constant for either residue-N or urea-N amendment but dependent on C:N ratio when plant residue was incorporated.     

Vertical structure of evapotranspiration at a forest site (a case study)

The components of ecosystem evapotranspiration of a Norway spruce forest (Picea abies L.) as well as the vertical structure of canopy evapotranspiration were analyzed with a combination of measurements and models for a case study of 5 days in September 2007. Eddy-covariance and sap flux measurements were performed at several heights within the canopy at the FLUXNET site Waldstein-Weidenbrunnen (DE-Bay) in the Fichtelgebirge mountains in Germany. Within and above canopy fluxes were simulated with two stand-scale models, the 1D multilayer model ACASA that includes a third-order turbulence closure and the 3D model STANDFLUX. The soil and understory evapotranspiration captured with the eddy-covariance system in the trunk space constituted 10% of ecosystem evapotranspiration measured with the eddy-covariance system above the canopy. A comparison of transpiration measured with the sap flux technique and inferred from below and above canopy eddy-covariance systems revealed higher estimates from eddy-covariance measurements than for sap flux measurements. The relative influences of possible sources of this mismatch, such as the assumption of negligible contribution of evaporation from intercepted water, and differences between the eddy-covariance flux footprint and the area used for scaling sap flux measurements, were discussed. Ecosystem evapotranspiration as well as canopy transpiration simulated with the two models captured the dynamics of the measurements well, but slightly underestimated eddy-covariance values. Profile measurements and models also gave us the chance to assess in-canopy profiles of canopy evapotranspiration and the contributions of in-canopy layers. For daytime and a coupled or partly coupled canopy, mean simulated profiles of both models agreed well with eddy-covariance measurements, with a similar performance of the ACASA and the STANDFLUX model. Both models underestimated profiles for nighttime and decoupled conditions. During daytime, the upper half of the canopy contributed approximately 80% to canopy evapotranspiration, whereas during nighttime the contribution shifted to lower parts of the canopy.