Dissolved soil organic matter from surface organic horizons under birch and conifers: Degradation in relation to chemical characteristics

The degradability and chemical characteristics of dissolved organic carbon (DOC) and nitrogen (DON) from the litter, F and H layers of silver birch (Betula pendula Roth), Norway spruce (Picea abies (L.) Karst) and Scots pine (Pinus sylvestris L.) stands were studied in an incubation experiment. Soil dissolved organic matter (DOM) was collected by centrifugation. Degradability was assessed in an incubation experiment by measuring the loss of DOC and DON, the mineralization rate of DOC and the availability of DOM to both bacteria and fungi, and by estimating the proportion of labile DOC of the total DOC. The degradability of DOC was highest in the litter layer and in that layer under birch. In the F and H layers, however, the degradability was highest under spruce. The most degradable fractions were the hydrophilic neutral fraction of DOC, the hydrophilic base fraction of DON, and the phenol fraction, as well as the smallest (<1 kDa) and largest (>100 kDa) molecular size classes of both DOC and DON. The degradability of these fractions seemed to be related to their relatively low C-to-N ratios. The hydrophilic acid fraction and the molecular size class of 1-10 kDa were more abundant in the H layer than in the litter layer, and thus apparently indicating a more decomposed DOM. In general, the effect of tree species on DOM was more obvious in the litter layer than in the lower organic layers.     

Carbon and nitrogen turnover in response to warming and nitrogen addition during early stages of forest litter decomposition—an incubation experiment

Purpose

Little is known about the interactive effects of temperature, nitrogen (N) supply, litter quality, and decomposition time on the turnover of carbon (C) and N of forest litter. The objective of this study was to investigate the interactive effects of warming, N addition and tree species on the turnover of C and N during the early decomposition stage of litters in a temperate forest.

Materials and methods

A 12-week laboratory incubation experiment was carried out. The leaf litters including two types of broadleaf litters (Quercus mongolica and Tilia amurensis), a needle litter (Pinus koraiensis), and a mixed litter of them were collected from a broad-leaved Korean pine mixed forest ecosystem in northeastern China in September 2009. Nine treatments were conducted using three temperatures (15, 25, and 35 °C) combined with three doses of N addition (equal to 0, 75, and 150 kg?·?ha^?1?a^?1, respectively, as NH₄NO₃).

Results and discussion

After 12 weeks of incubation, the mass loss ranged between 12 and 35 %. The broadleaf litters had greater mass loss and cumulative CO₂–C emission than the needle litter. Temperature and N availability interacted to affect litter mass loss and decomposition rate. The dissolved organic carbon (DOC) and nitrogen (DON) concentrations in litter leachate varied widely with litter types. DOC increased significantly with increased temperature but decreased significantly with increased N availability. DON increased significantly with increased N availability but showed a higher level at the moderate decomposition temperature. The amounts of CO₂ and N₂O emission were significantly higher at 25 °C than those at 15 and 35 °C, and were significantly increased by the N addition.

Conclusions

The present study indicated relatively intricate temperature and N addition effects on C and N cycling during early stages of litter decomposition, implying that future increases in temperature and N deposition will directly affect C and N cycling in broad-leaved Korean pine mixed forest ecosystem, and may indirectly influence the ecosystem composition, productivity, and functioning in NE China. It is, therefore, important to understand the interactive effects of biotic and abiotic factors on litter decomposition in field conditions in order to assess and predict future ecosystem responses to environmental changes in NE China.     

Properties of dissolved organic matter derived from silver birch and Norway spruce stands: Degradability combined with chemical characteristics

Dissolved organic matter (DOM) derived from the humus layer under silver birch (Betula pendula Roth), Norway spruce (Picea abies (L.) Karst.), and mixed stands, and from senescent birch leaves and from spruce needles of the four oldest year-growth were characterized microbiologically and chemically. Samples were collected in the autumn and the solutions were obtained by centrifugation-drainage technique. The degradability of DOM, the availability of DOM to bacteria and fungi, concentrations of phenolic compounds and carbohydrates, and the distribution of carbon and nitrogen into fractions according to the chemical nature and the molecular size were studied. DOM derived from leaves and needles was clearly more labile than DOM derived from the humus layer indicating the importance of studying the DOM originating from fresh litter when assessing the turnover of DOM.DOM derived from spruce needles appeared to differ chemically greatly from all other samples. It had very high concentrations of carbohydrates, probably due to the sampling time, and phenolic compounds. The chemical composition of DOM derived from humus layer did not reflect the composition of DOM derived from needles and leaves. DOM derived from birch leaves degraded more than DOM derived from spruce needles and DOM derived from humus layer collected at the birch sites degraded more than DOM derived from humus layer collected at the spruce sites. The degradability of different compound groups of DOC and DON was studied in a short-term incubation (20 d) of DOM solutions by characterizing the solutions initially and after the incubation. Almost all compound groups appeared to degrade but weak hydrophobic acids, bases, hydrophilic neutrals, the smallest molecular size compounds, carbohydrates, and phenolic compounds degraded the most.     

Quality of soluble organic C,N, and P produced by different types and species of litter: Root litter versus leaf litter

In forested ecosystems, the quality of dissolved organic matter (DOM) produced by freshly senesced litter may differ by litter type and species, and these differences may influence the amount of DOM that is respired versus that which may either contribute to soil organic matter accumulation or be leached from the ecosystem. In this study, we investigated the effect of litter type (including freshly senesced fine root, leaf, fine woody, and reproductive litter) and species (5 species of leaf litter) on several measures of the quality of DOM produced at a site along a primary successional chronosequence at Mt. Shasta, California. We measured differences in solid litter chemistry (C, N, and P concentration) and differences in the concentration of dissolved organic C, N, and P (DOC, DON, and DOP, respectively), water-soluble monomeric carbohydrates, polyphenols, proteins, fractions of DOC, as well as UV absorbance. For several aspects of DOC quality, DOM from fine roots was less labile than DOM from leaf litter. In contrast to DOC quality, soluble material originating from fine roots was high in labile forms of dissolved N and P in comparison to leaf litter. We also found that leaf litter with greater total %N or %P in solid litter had higher DON or DOP concentration (and higher total soluble P concentration). A very high percentage, on average 72% (up to 89%) of the total P in leaf litter was water-soluble and mostly inorganic P. Concentrations of soluble polyphenols were strongly related to DOC, and concentrations of soluble proteins were significantly related to DON in leaf litter of different species. During primary succession at the Mt. Shasta site, an increasing ratio of root to leaf litter production and shifting species composition has been found to occur, and the results of this study suggest that some aspects of DOC quality reflect a decrease in labile forms of DOC originating from both above and belowground litter. In contrast, dissolved N and P reflect an increase in labile forms with increasing inputs of root litter. In particular, our study has demonstrated important differences in the quality of inputs of DOM from freshly senesced root and leaf litter, and these differences have implications for C and nutrient cycling.     

Warming effects on the early decomposition of three litter types,Eastern Tibetan Plateau,China

Temperature and litter quality are two of the key factors controlling litter decomposition. Predicted global warming and vegetation succession will therefore have profound impacts. This study was conducted to assess effects of experimental warming on litter decomposition and nutrient dynamics of two contrasting tree species (red birch, Betula albosinensis Burk., and dragon spruce, Picea asperata Mast.) and a mixture of the two with the heating cable method in the eastern Tibetan Plateau of China. This treatment raised surface soil temperature by 3.2°C and resulted in a 5.2% decline in soil moisture 10 cm below the soil surface. The water content of dragon spruce, red birch and mixed litter was decreased by 18, 11 and 13%, respectively. Marked differences between the two species in the decomposition rates and nutrient remaining percentages were detected. Moreover, we found positive, non‐additive effects of litter mixture. Experimental warming did not affect mass loss and nutrient release of dragon spruce litter but significantly increased mass loss and affected nutrient release of red birch and mixed litter during the early decomposition period. Overall, inter‐specific (red birch and dragon spruce) differences in decomposability were substantially larger than warming‐induced responses. Thus, a warming‐induced community succession towards dragon spruce forests in the Tibetan Plateau region could have a greater impact on early litter decomposition than warming itself.     

Organic matter characteristics and C and N transformations in the humus layer under two tree species, Betula pendula and Picea abies

The aim of this study was to compare the effects of silver birch (Betula pendula Roth) and Norway spruce (Picea abies (L.) Karst.) on soil C and N transformations and on the characteristics of organic matter. Soil samples were taken from the humus layer of a replicated 35-year-old birch-spruce field experiment growing on Vaccinium myrtillus site type in middle-eastern Finland. The soil was a podzol and humus type mor. Soil pH was higher under birch (4.7) than under spruce (4.1). The C-to-N ratio was lower under birch (17) than under spruce (23). Per unit organic matter, microbial biomass C and N, net N mineralization and net nitrification were all higher in birch soil than in spruce soil. The rate of C mineralization (CO₂ production) was, however, the same regardless of tree species. Water-extracts were analyzed for the concentrations of dissolved organic C (DOC) and N (DON) and characterized according to molecular size distribution by ultrafiltration and according to chemical composition using a resin fractionation technique. The concentration of DON, in particular, was higher in birch soil than in spruce soil. The distribution of DOC and DON into different fractions based on molecular size or chemical composition was rather similar in both soils. The concentration of total phenolics, expressed as tannic acid equivalents, was higher in the humus layer under birch than in the humus layer under spruce, because the birch humus layer contained significantly more low-molecular weight (about <0.5 kD) phenolics than the spruce humus layer did. The concentration of proanthocyanidins (condensed tannins) was higher in spruce soil than in birch soil. The concentrations of the five most abundant phenolic acids showed that ferulic and p-coumaric acids were more abundant in spruce soil. Birch soil tended to contain slightly more nonvolatile sesquiterpenes than the spruce soil. The concentration of diterpenes was similar in both soils; but birch soil contained significantly more triterpenes, mainly sterols, than spruce soil did.     

Non-additive effects of mixing different sources of dissolved organic matter on its biodegradation

To examine the potential impact of plant species richness on ecosystems, we studied non-additive effects of different plant litters on the biodegradation rate of dissolved organic matter (DOM) when mixing plant leaf-derived DOM derived from different plant species. A full factorial biodegradation experiment (31 possible singular and multiple combinations of five litter type-derived DOM sources) was conducted using plant litters from the five most abundant plant species in a subtropical watershed ecosystem, from which dissolved organic carbon (DOC) disappearance was measured. Loss of DOC over time was considered biodegradable DOC. We tested whether DOM diversity, measured as source species richness and composition, would affect biodegradation rates. Overall, we found significant non-additive (synergistic) effects of DOM diversity on biodegradation rates of DOM, which were explained both by plant species richness and composition. Across all treatments, a significantly higher biodegradation rate was correlated with the presence of DOM from higher nitrogen (N) containing plant litters; conversely, the presence of lower N decreased these rates. The N content and chemical characteristic of DOM might influence the magnitude of the synergistic effect. Our results suggest that loss of plant litter species diversity would not affect DOC biodegradation rate, provided that at least two species are conserved. However, the variability in DOC biodegradation rate across the treatments decreased with increased DOM diversity at three incubation time points. Our results also indicate that in an ecosystem with low plant biodiversity, loss of key species such as Lophostemon confertus could reduce the synergistic effects on DOC biodegradation rate.     

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

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

Responses of soil dissolved organic matter to long-term plantations of three coniferous tree species

Tree species have significant effects on the availability and dynamics of soil organic matter. In the present study, the pool sizes of soil dissolved organic matter (DOM), potential mineralizable N (PMN) and bio-available carbon (C) (measured as cumulative CO₂ evolution over 63 days) were compared in soils under three coniferous species — 73 year old slash (Pinus elliottii), hoop (Araucaria cunninghamii) and kauri (Agathis robusta) pines. Results have shown that dissolved organic N (DON) in hot water extracts was 1.5–1.7 times lower in soils under slash pine than under hoop and kauri pines, while soil dissolved organic C (DOC) in hot water extracts tended to be higher under slash pine than hoop and kauri pines but this was not statistically significant. This has led to the higher DOC:DON ratio in soils under slash pine (32) than under hoop and kauri pines (17). Soil DOC and DON in 2 M KCl extracts were not significantly different among the three tree species. The DOC:DON ratio (hot water extracts) was positively and significantly correlated with soil C:N (R² = 0.886, P < 0.01) and surface litter C:N ratios (R² = 0.768, P < 0.01), indicating that DOM was mainly derived from litter materials and soil organic matter through dissolution and decomposition. Soil pH was lower under slash pine (4.5) than under hoop (6.0) and kauri (6.2) pines, and negatively correlated with soil total C, C:N ratio, DOC and DOC:DON ratio (hot water extracts), indicating the soil acidity under slash pine favored the accumulation of soil C. Moreover, the amounts of dissolved inorganic N, PMN and bio-available C were also significantly lower in soils under slash pine than under hoop and kauri pines. It is concluded that changes in the quantity and quality of surface litters and soil pH induced by different tree species largely determined the size and quality of soil DOM, and plantations of hoop and kauri pine trees may be better in maintaining long-term soil N fertility than slash pine plantations.     

Comparison of tree species effects on microbial C and N transformations and dissolved organic matter properties in the organic layer of boreal forests

The aim of this study was to determine whether tree species consistently affects soil microbial activities related to C and N cycling and to compare these activities with the characteristics of soil dissolved organic matter (DOM). Samples were taken from the mor-type organic layer (Of+Oh) underlain by podzols of six 20–72-year-old tree-species experiments on different site types in different parts of Finland. Sampling plots were dominated by silver birch (Betula pendula Roth), Norway spruce (Picea abies (L.) Karst) or Scots pine (Pinus sylvestris L., only on four sites). Amounts of C and N in the microbial biomass and rates of C mineralization (CO₂ production) and net N mineralization were determined, and water extracts were analysed for concentrations of DOC and DON and characterized according to molecular size by ultrafiltration and according to chemical composition using a resin fractionation technique. In all older stands, birch, compared to spruce or pine, increased soil pH, NH₄-concentration and amounts of C and N in microbial biomass and decreased the C-to-N ratio and ratio of dissolved organic N (DON)-to-mineral N. Birch had similar effects also in part of the younger stands. Birch also increased the rates of both C and net N mineralization compared to spruce or pine but only on two sites. In all soils, net nitrification was low. The distribution of DOC into different fractions based on chemical composition and molecular size was rather similar in all soils. The most abundant chemical fraction was hydrophobic acids, and the most abundant molecular size fraction was 10–100 kDa. The C-to-N ratio varied but was lowest in hydrophilic bases and in the smallest molecular size class. Mineralization of C was highly and positively correlated with concentration of DOC (Pearson's correlation coefficient r = 0.9, P < 0.01). The results indicated close interactions between microbial processes and dissolved organic matter.     

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

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

Simulating soil freeze/thaw cycles typical of winter alpine conditions: Implications for N and P availability

Seasonally snow-covered alpine soils may be subjected to freeze/thaw cycles, particularly during years having little snow and during the late winter and early spring periods. Freeze/thaw cycles can stimulate soil mineralization and could therefore be one factor regulating nitrogen (N) and phosphorus (P) availability and cycling. In this study laboratory incubation experiments using four soils having contrasting properties have been used to characterize the change in N and P forms (microbial and soluble inorganic/organic) that occur after simulated freeze/thaw cycles.Soil samples were collected from locations representing extreme examples of either direct human management (grazed meadow (site M) and extensive grazing beneath larch (site L)) or those disturbed by more natural events (recent avalanche and colonisation by alder (site A)) and from beneath the expected forest climax vegetation beneath fir (site F). Topsoil from these sites, maintained at two different water contents (20 and 30%, w/w), were exposed to either a single (SF) or four sequential (4SF) freeze/thaw cycles. Each cycle consisted of 12 h at −9 °C and 12 h at +4 °C mimicking a diurnal pattern.A SF cycle reduced microbial N for soils from sites F and A and was accompanied by a significant increase in dissolved organic nitrogen (DON) at both moisture contents. In contrast, the microbial N of soils from M and L was not affected by the freeze/thaw cycles, suggesting a particular adaptation of soil microbes to these extremes in temperature. Freeze/thaw cycles resulted in a significant increase in the net ammonification in all soils.Extractable total dissolved N (TDN) and total dissolved P (TDP) increased in all soils after a SF cycle, however, the relative importance of the different N and P forms differed. At the lower soil moisture content, NO₃⁻ concentrations remained constant or slightly decreased in all soils, except that from site M. In all other soils DON appeared to replace NO₃⁻ as the potentially mobile N source after the freeze/thaw cycles. The relative contribution of dissolved organic P to TDP after freeze/thaw remained significant, and greater than 50% in all soils.Freeze/thaw cycles, in seasonally snow covered soils, are likely to have a selective effect on the microbial biomass. Freezing and thawing resulted in a pulse of net ammonification and DON release, which represent an important influence upon N cycling in these alpine systems.     

Soil organisms and carbon, nitrogen and phosphorus mineralisation in Norway spruce and mixed Norway spruce – Birch stands

 We examined how soil organisms and C, N and P mineralisation are affected by admixing deciduous tree species, silver birch (Betula pendula) and woollen birch (B. pubescens), in managed Norway spruce (Picea abies) stands. Pure spruce and mixed spruce–birch stands were examined at four sites in southern and central Sweden. Soil macroarthropods and enchytraeids were sampled in litter and soil. In the uppermost 5 cm of soil humus we determined microbial biomass and microbial respiration; we estimated the rate of C, N and P mineralisation under laboratory conditions. The densities of Coleoptera, Diptera and Collembola were larger in mixed stands than in spruce stands. Soil fauna composition differed between mixed and spruce stands (as revealed by redundancy analysis). Staphyliniidae, Elateridae, Cecidiomyidae larvae and Onychiuridae were the families that increased most strongly in mixed stands. There were no differences in microbial biomass and microbial respiration, nor in the C, N and P mineralisation rates, between mixed and spruce stands. However, within mixed stands microbial biomass, microbial activity and C mineralisation were approximately 15% higher under birch trees than under spruce trees. We propose that the presence of birch leaf litter was likely to be the most important factor causing differences in soil fauna composition. Birch may also influence the quality and the decomposition rate of humus in mixed stands. However, when the proportion of birch trees is low, the short-term (decades) effect of this species on decomposition is likely to be small in mixed stands on acid forest soils. Received: 20 February 1998     

Dissolved soil organic nitrogen and carbon in a Norway spruce stand and an adjacent clear-cut

 The aims of this study were to characterize dissolved soil organic N (DON) and C (DOC) in a coniferous stand and an adjacent clear-cut, and to evaluate the importance of DON in N leaching. The study was carried out in a Norway spruce stand and a clear-cutting treatment in the same forest stand. Concentrations of DON in soil solution were monitored for 5 years after clear-cutting with gravity lysimeters. In the Norway spruce stand DON comprised 62–83% of the total N in soil solution over the 5-year period. The concentrations of DON in the clear-cut were higher than in the forest stand, but the proportion of total N was lower. To characterize dissolved organic matter, soil samples were aerobically incubated for 6 weeks in the laboratory, and the quantity, molecular size distribution and chemical nature of both DON and DOC were determined from water extracts made before and after the incubation. In the soil samples from the Norway spruce stand, C-rich compounds with a high C/N ratio and large molecular size were formed. In contrast, after the incubation the major carriers of DON in soil samples from the clear-cut were N-rich organic compounds with a low C/N ratio and a small molecular size. The distribution of different chemical fractions of DOC in soil did not differ much whether recovered from the Norway spruce stand or the clear-cut. It was (from highest to lowest concentration): hydrophobic acids>hydrophilic acids>phenols>hydrophilic neutrals. A major part of DON was also carried by these fractions. During incubation the concentration of N-containing hydrophilic acids increased, especially in the soil from the clearcut. In soil samples from the Norway spruce stand, the rate of net N mineralization was low and no NO₃ was formed, whilst the rate of net N mineralization was high and net nitrification was intensive in soil from the clear-cut. Received: 12 June 2000     

Do late-successional tannin-rich plant communities occurring on highly acidic soils increase the DON/DIN ratio?

Previous studies suggested that late-successional tannin-rich plant communities increase the amount of dissolved organic N (DON) relative to dissolved inorganic N (DIN) in decomposing litter. We devised an experiment to test this hypothesis by adding varying proportions of black spruce (Picea mariana) and tannin-rich Kalmia angustifolia litter to forest floor samples collected on six black spruce cutovers. An increasing proportion of Kalmia litter increased condensed tannin and total phenolic concentrations over the course of a 46-week incubation period. Mineral N concentrations did not vary among treatments in spite of much higher total N concentrations in Kalmia litter. This was more likely due to the formation of protein–tannin complexes rather than microbial immobilization of N, as indicated by the decline in available C with increasing Kalmia litter on two of the five sampling dates. There was a significant positive linear trend between the proportion of Kalmia litter and the DON/DIN ratio on one sampling date (week 13) only. Results suggest that the DON/DIN ratio is controlled by confounding factors (e.g., tannins bonding with non-extractable humus particles) and has limited value for describing ecological succession.     

Effect of N addition,moisture, and temperature on soil microbial respiration and microbial biomass in forest soil at different stages of litter decomposition

Purpose

The objective of the present study was to investigate the interactive effects of nitrogen (N) addition, temperature, and moisture on soil microbial respiration, microbial biomass, and metabolic quotient (qCO₂) at different decomposition stages of different tree leaf litters.

Materials and methods

A laboratory incubation experiment with and without litter addition was conducted for 80 days at two temperatures (15 and 25 °C), two wetting intensities (35 and 50 % water-filled porosity space (WFPS)) and two doses of N addition (0 and 4.5 g N m^?2, as NH₄NO₃). The tree leaf litters included three types of broadleaf litters, a needle litter, and a mixed litter of them. Soil microbial respiration, microbial biomass, and qCO₂ along with other soil properties were measured at two decomposition stages of tree leaf litters.

Results and discussion

The increase in soil cumulative carbon dioxide (CO₂) flux and microbial biomass during the incubation depended on types of tree leaf litters, N addition, and hydrothermal conditions. Soil microbial biomass carbon (C) and N and qCO₂ were significantly greater in all litter-amended than in non-amended soils. However, the difference in the qCO₂ became smaller during the late period of incubation, especially at 25 °C. The interactive effect of temperature with soil moisture and N addition was significant for affecting the cumulative litter-derived CO₂-C flux at the early and late stages of litter decomposition. Furthermore, the interactive effect of soil moisture and N addition was significant for affecting the cumulative CO₂ flux at the late stage of litter decomposition but not early in the experiment.

Conclusions

This present study indicated that the effects of addition of N and hydrothermal conditions on soil microbial respiration, qCO₂, and concentrations of labile C and N depended on types of tree leaf litters and the development of litter decomposition. The results highlight the importance of N availability and hydrothermal conditions in interactively regulating soil microbial respiration and microbial C utilization during litter decomposition under forest ecosystems.

     

Ant-mediated effects on spruce litter decomposition, solution chemistry, and microbial activity

Forest management practices often generate clear-cut patches, which may be colonized by ants not present in the same densities in mature forests. In addition to the associated changes in abiotic conditions ants can initiate processes, which do not occur in old-growth stands. Here, we analyse the effects of ants and aphid honeydew on litter solution of Norway spruce, microbial enzyme activities, and needle decomposition in a field and greenhouse experiment during summer 2003. In the field, low ant densities had relatively little effects on litter solution 30 cm away from a tree trunk, but significantly increased organic carbon concentrations and decreased inorganic nitrogen concentrations next to a trunk where ants tend to build their nests. In a greenhouse experiment, the addition of ants to lysimeters containing spruce litter significantly increased dissolved organic carbon (DOC), dissolved organic nitrogen (DON), NH₄-N, NO₃-N and K concentrations in litter solutions compared to the control treatment, while the simulation of aphid infestation (addition of honeydew) significantly increased DOC as a direct result of honeydew leaching, and decreased inorganic N concentrations in leachates. The presence of ants resulted in a changed composition of dissolved organic matter (DOM) with more aromatic and complex compounds, and microbial enzyme activity was significantly higher in litter extracts from the ant treatment compared to the honeydew and control treatment. However, mass loss, litter %C and %N were not affected by ants or honeydew. Our results suggest that ants have a distinct and immediate effect on solution composition and microbial activity in the litter layer indicating accelerated litter decay whereas the effect of honeydew was insignificant.     

Dissolved organic carbon and nitrogen in precipitation,throughfall, soil solution,and stream water of the tropical highlands in northern Thailand

Dissolved organic matter (DOM) is important for the cycling and transport of carbon (C) and nitrogen (N) in soil. In temperate forest soils, dissolved organic N (DON) partly escapes mineralization and is mobile, promoting loss of N via leaching. Little information is available comparing DOC and DON dynamics under tropical conditions. Here, mineralization is more rapid, and the demand of the vegetation for nutrients is larger, thus, leaching of DON could be small. We studied concentrations of DOC and DON during the rainy seasons 1998–2001 in precipitation, canopy throughfall, pore water in the mineral soil at 5, 15, 30, and 80 cm depth, and stream water under different land‐use systems representative of the highlands of northern Thailand. In addition, we determined the distribution of organic C (OC) and N (ON) between two operationally defined fractions of DOM. Samples were collected in small water catchments including a cultivated cabbage field, a pine plantation, a secondary forest, and a primary forest. The mean concentrations of DOC and DON in bulk precipitation were 1.7 ± 0.2 and 0.2 ± 0.1 mg L^–1, respectively, dominated by the hydrophilic fraction. The throughfall of the three forest sites became enriched up to three times in DOC in the hydrophobic fraction, but not in DON. Maximum concentrations of DOC and DON (7.9–13.9 mg C L^–1 and 0.9–1.2 mg N L^–1, respectively) were found in samples from lysimeters at 5 cm soil depth. Hydrophobic OC and hydrophilic ON compounds were released from the O layer and the upper mineral soil. Concentrations of OC and ON in mineral‐soil solutions under the cabbage cultivation were elevated when compared with those under the forests. Similar to most temperate soils, the concentrations in the soil solution decreased with soil depth. The reduction of OC with depth was mainly due to the decrease of hydrophobic compounds. The changes in OC indicated the release of hydrophobic compounds poor in N in the forest canopy and the organic layers. These substances were removed from solution during passage through the mineral soil. In contrast, organic N related more to labile microbial‐derived hydrophilic compounds. At least at the cabbage‐cultivation site, mineralization seemed to contribute largely to the decrease of DOC and DON with depth, possibly because of increased microbial activity stimulated by the inorganic‐N fertilization. Similar concentrations and compositions of OC and ON in subsoils and streams draining the forested catchments suggest soil control on stream DOM. The contribution of DON to total dissolved N in those streams ranged between 50% and 73%, underscoring the importance of DOM for the leaching of nutrients from forested areas. In summary, OC and ON showed differences in their dynamics in forest as well as in agricultural ecosystems. This was mainly due to the differing distribution of OC and ON between the more immobile hydrophobic and the more easily degradable hydrophilic fraction.     

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.