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.     

How do coniferous needle tannins influence C and N transformations in birch humus layer?

The aim of this study was to explore the response of C and N transformations in the humus layer under silver birch (Betula pendula Roth) to compounds, especially condensed tannins, of different molecular weight extracted and fractioned from Norway spruce (Picea abies (L.) Karst) and Scots pine (Pinus sylvestris L.) needles. Lighter fractions containing tannin monomers and dimers as well as many other compounds, and heavier fractions consisting predominantly of polymerized condensed tannins, were added to samples taken from the humus layer of birch stand. The effects of the spruce and pine fractions were mostly similar, but some differences in magnitude were observed; our results indicated that lighter fractions of pine were easier for microbes to degrade and use than lighter fractions of spruce. Lighter fractions of both tree species increased soil respiration and decreased net N mineralization, while heavier fractions inhibited respiration and increased net N mineralization. Microbial biomass C was not clearly affected by any of the treatments, but with some of the pine fractions the amount of N in the microbial biomass was increased. Comparison of the effects of fractions in birch and in spruce and pine soils, which were studied earlier, showed no major differences between the effects of the fractions in birch and in their own soils, but gave some indication of adaptation.     

Protein precipitation by tannins in soil organic horizon and vegetation in relation to tree species

The aim of this study was to compare the concentration of tannins and their capacity to precipitate proteins in the dominant species of ground vegetation (Deschampsia flexuosa (L.) Trin., Pleurozium schreberi (Brid.) Mitt., Vaccinium myrtillus (L.), and Vaccinium vitis-idaea (L.)) and in different layers of the soil organic horizon (litter layer—L, fermentation layer—F, humified layer—H) under silver birch (Betula pendula Roth.), Norway spruce (Picea abies (L.) Karst.), and Scots pine (Pinus sylvestris L.). Total tannin concentrations were also measured in leaves or needles of birch, spruce, and pine. The study site is located in Kivalo, northern Finland, close to the Arctic Circle. Differences in total tannin concentrations in ground vegetation were due mainly to species, with Vaccinium species having the highest values. The influence of the dominant tree species was less important. Protein precipitating capacity was dependent on plant species; the highest values occurred in Vaccinium species and spruce. Because of their relatively high protein precipitating capacity but low total tannin concentration, D. flexuosa and P. schreberi seemed to have more astringent tannins. Concentrations of total tannin and hydrolyzable tannin in the soil organic horizon differed depending on the layer and tree species. In general, the highest concentrations of total tannins were found under birch and spruce in the L layer and the lowest concentrations under pine. Protein precipitating capacity was usually the lowest in the H layer and highest under birch and spruce in the F and H layers. We showed that lignin from rotted pine wood can also precipitate proteins but only small amounts; additionally, lignin can be an important source of error for soil total tannin measurements.     

Characteristics of dissolved organic matter and phenolic compounds in forest soils under silver birch (Betula pendula), Norway spruce (Picea abies) and Scots pine (Pinus sylvestris)

The aim was to characterize dissolved organic matter in soils under different tree species. Molecular size distribution and chemical composition of dissolved organic carbon and nitrogen were determined in water extracts from humus layers and mineral soils taken from silver birch ( Betula pendula Roth.), Norway spruce ( Picea abies (L.) Karst.) and Scots pine ( Pinus sylvestris L.) stands. Concentrations of tannins and 15 phenolic acids in the humus layers were measured. Per unit of organic matter, the concentrations of dissolved organic C and N were larger in birch and spruce humus layers than in the pine humus layer. In the underlying mineral soil, the concentrations of dissolved organic C were similar at all sites, but the concentration of dissolved organic N was greater in spruce and pine soils than in birch soil. In all soils, the 10–100 kDa fraction was the most abundant molecular size group and hydrophobic acids the most abundant chemical group of dissolved organic C. In all humus layers, hydrophobic acids and hydrophilic bases were the major components of dissolved organic N. There were only minor differences in the concentrations of total tannins in the humus layers under different tree species. Small-molecule tannins (about < 0.5 kDa) were most abundant in the birch humus, and large-molecule tannins in the pine humus. Coniferous humus contained more ferulic and p -coumaric acids than did the birch humus. The concentrations of 3,4 and 3,5-dihydroxybenzoic acid, vanillic acid and 4-hydroxybenzoic acid were similar in all soils.     

Potential activities of enzymes involved in N,C, P and S cycling in boreal forest soil under different tree species

The aim of this study was to compare the potential activity of enzymes involved in N, C, P and S cycling in the humus layer under three tree species: silver birch, Norway spruce and Scots pine. For arylsulphatase and protease the highest activities were found under birch, whereas beta-glucosidase activity was highest under pine. Beta-glucosaminidase and acid phosphatase showed similar activities regardless of tree species. Our studies show that soils under these species may differ enzymatically from each other. Enzyme activity studies under different tree species need more attention as the activity of different enzymes influences on soil nutrient availability in boreal forest soil.     

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.     

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.     

Phenolic compounds and terpenes in soil organic horizon layers under silver birch,Norway spruce and Scots pine

The aim of this study was to monitor the concentration of some plant secondary metabolites, such as low- and high-molecular-weight phenolics, condensed tannins (proanthocyanidins), and sesqui-, di- and triterpenes, in litter (L), fermentation (F) and humified (H) layers of the soil organic horizon in stands dominated by silver birch (Betula pendula Roth.), Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.), and from samples taken from birch leaves and spruce and pine needles. Concentrations of low- and high-molecular-weight phenolics and terpenes from the four most dominant species of ground vegetation taken from the stands were also determined. In general, the L layer showed higher concentrations of both phenolic compounds and terpenes than the F and H layers did. Concentrations of terpenes decreased relatively more with soil depth than did concentrations of total phenolics (=low + high) or condensed tannins. Of the total phenolics, the proportion of low-molecular-weight phenolics increased from the L to the H layer with all tree species. Concentrations of all terpenes were highest under pine and lowest under birch. Concentrations of the studied secondary metabolites in the ground vegetation species were similar under different tree species. Blueberry (Vaccinium myrtillus L.) and lingonberry (Vaccinium vitis-idaea L.) contained considerably higher concentrations of total phenolics than did feather moss (Pleurozium schreberi (Brid.) Mitt.) and wavy hair-grass (Deschampsia flexuosa (L.) Trin.). Concentration of total phenolics in soil correlated positively with soil respiration and microbial biomass C, and terpenes showed positive correlation with soil C-to-N ratio.     

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

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

Carbon and nitrogen dynamics in a forest soil amended with purified tannins from different plant species

Tannins are purported to be an important factor controlling nitrogen cycling in forest ecosystems, and the ability of tannins to bind proteins in protein-tannin complexes is thought to be the primary mechanism responsible for these effects. In this study, we examined the influence of well-characterized tannins purified from five different plant species on C and N dynamics of a forest soil A horizon. Tannic acid, a commonly used and commercially available hydrolyzable tannin (HT), and cellulose were also included for comparison. With the exception of tannins from huckleberry (Vaccinium ovatum), the amendments increased respiration 1.4-4.0 fold, indicating that they were acting as a microbial C source. Tannic acid was significantly more labile than the five purified tannins examined in this study. All treatments decreased net N mineralization substantially, through greater N immobilization and decreased mineralization. The six tannins inhibited gross ammonification rates significantly more than cellulose. This suggests that added tannins had effects in addition to serving as an alternative C source. Tannins purified from Bishop pine (Pinus muricata) were the only tannins that significantly inhibited potential gross nitrification rates, however, rates were low even in the control soil making it difficult to detect any inhibition. Differences in tannin structure such as condensed versus HTs and the hydroxylation pattern of the condensed tannin B-ring likely explain differences observed among the tannin treatments. Contrary to other studies, we did not find that condensed tannins were more labile and less inhibitory than HTs, nor that shorter chained tannins were more labile than longer chained tannins. In addition to supporting the hypothesis that reduced N availability in the presence of tannins is caused by complexation reactions, our data suggests tannins act as a labile C source leading to increased N immobilization.     

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     

Microbial enzyme activities in leaf litter, humus and mineral soil layers of European forests

The rationale of the study was to investigate microbial activity in different soil horizons in European forests. Hence, activities of chitinase and cellulase, microbial biomass carbon (C_mic) and basal respiration were measured in litter, fragmentation, humus and mineral soil layers collected several times from various beech and spruce forests. Sites were selected to form a gradient in N availability. Analyses were also performed on beech litter from a litterbag transplant experiment. Furthermore, microbiological parameters were measured in horizons of beech and spruce chronosequence sites with different stand age in order to investigate the influence of forest rotation, and hence changes in soil organic matter (SOM) dynamics, on microbial activity. Finally in horizons of one beech forest, the seasonal variation of selected microbiological parameters was measured more intensively. β-Glucosaminidase and cellobiohydrolase activities were measured using fluorogenic 4-methylumbelliferyl substrates to estimate chitinase and cellulase activities, respectively. On a spatial scale, chitinase and cellulase activities, C_mic determined by substrate induced respiration, and basal respiration ranged from 144 to 1924 and 6-177 nmol 4-MU g⁻¹ org-C h⁻¹, 8-48 mg C g⁻¹ org-C and 11-149 μg CO₂-C g⁻¹ org-C h⁻¹, respectively; in general values were significantly lower in layers of humus and mineral soil than of litter. Chitinase activity, C_mic and basal respiration from humus and mineral soil layers, together, correlated positively, while none correlated with cellulase activity. Similarly in the litter layer, no correlations were found between the microbiological parameters. On a seasonal scale, a time lag between a burst in basal respiration rate and activities of both enzymes were observed. In general, activities of cellulase and chitinase, C_mic and basal respiration, did not change with stand age, except in the humus layer in the spruce chronosequence, where C_mic decreased with stand age. In the litter layer, cellulase activity was significantly and positively related to the C:N ratio, while only a tendency for chitinase activity was shown, indicating that enzyme activities decreased with increasing N availability. In accordance, the enzyme activities and C_mic decreased significantly with increasing chronic N deposition in the humus layer, while basal respiration only tended to decrease with increasing N deposition. In contrast, enzyme activities in beech litter from litterbags after 2 years of incubation were generally higher at sites with higher N deposition. The results show different layer-specific responses of enzyme activities to changes in N availability, indicating different impacts of N availability on decomposition of SOM and stage of litter decomposition.     

Carbon respiration and nitrogen dynamics in Corsican pine litter amended with aluminium and tannins

We investigated the carbon (C) mineralisation and nitrogen (N) dynamics in litter from a Corsican pine forest in response to individual and combined additions of aluminium (Al), condensed tannin (extracted from fresh Corsican pine needles) and hydrolysable tannin (commercial tannic acid). Production rates of CO₂, NH₄⁺ and NO₃⁻ concentrations, tannin concentrations and Al speciation were determined at various time intervals during a 28-day incubation experiment. The addition of Al decreased CO₂ production and shut down nitrification. Exchangeable NH₄⁺ strongly increased in the Al-amended litter, likely due to (i) decreased microbial uptake of NH₄⁺, (ii) the inhibition of nitrification and (iii) competition for soil organic matter (SOM) binding sites by Al. Both tannin species affected C mineralisation and/or N dynamics, be it in different ways. Addition of tannic acid led to a strong increase of the C mineralisation rate and microbial uptake of N, caused by rapid degradation of this labile tannin and subsequent increased microbial nutrient demand. Net immobilisation of N occurred as long as one week after addition. Condensed tannin was not consumed but probably strongly bound to (nitrogenous) SOM compounds, forming recalcitrant complexes and decreasing net N mineralisation. Complexation of Al by tannins in solution before addition to the litter mitigated the Al-induced release of exchangeable NH₄⁺. In the case of condensed tannin with complexed Al, this was due to detoxification of Al through complexation. Increased microbial demand for N likely played a major role in decreased NH₄⁺ accumulation in the samples to which tannic acid with complexed Al was added. Nitrification was shut down despite of the complexation of Al by either condensed tannin or tannic acid.     

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.     

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.     

Microbial community structure and characteristics of the organic matter in soils under Pinus sylvestris, Picea abies and Betula pendula at two forest sites

 Microbial biomass C (C_mic), C mineralization rate, phospholipid fatty acid (PLFA) profiles and community level physiological profiles (CLPPs) using Biolog were determined from the humus and mineral soil layers in adjacent stands of Scots pine (Pinus sylvestris L.), Norway spruce [Picea abies (L.) Karst.] and silver birch (Betula pendula Roth) at two forest sites of different fertility. In addition, the Fourier-transformed infrared (FTIR) spectra were run on the samples for characterization of the organic matter. C_mic and C mineralization rate tended to be lowest under spruce and highest under birch, at the fertile site in all soil layers and at the less fertile site in the humus layer. There were also differences in microbial community structure in soils under different tree species. In the humus layer the PLFAs separated all tree species and in the mineral soil spruce was distinct from pine and birch. CLPPs did not distinguish microbial communities from the different tree species. The FTIR spectra did not separate the tree species, but clearly separated the two sites. Received: 3 December 1999     

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 degradation of hydrolysable and condensed tannin polyphenol-protein complexes in soils from different land-use histories

Polyphenols are capable of binding to proteins and form polyphenol-protein complexes thus reducing the release of N from decomposing plant materials. The objective of this work was to test if under polyphenol-rich vegetations adapted microbial communities had developed capable of breaking down recalcitrant polyphenol-protein complexes. Soils used for this investigation were from different 10-year-old tropical agricultural systems (maize, sugarcane plots and Gliricidia sepium or Peltophorum dasyrrachis woodlots) and natural systems (secondary forest and Imperata cylindrica grassland). TA (tannic acid, hydrolysable tannin), QUE (quebracho, condensed tannin), BSA (bovine serum albumin, protein) or TA/BSA and QUE/BSA polyphenol-protein complexes were incubated at 28 °C in these soils. CO₂-C and ¹³C evolution were periodically monitored and mineral N release, microbial biomass N and phospholipid fatty acid (PLFA) profiles measured at the end.QUE was able to bind about 25% more protein than TA. In all systems the individual uncomplexed substrates were more easily degraded than the complexes. On average, net cumulative CO₂-C evolution from TA/BSA complexes was more than 5 times higher than from QUE/BSA complexes, indicating higher C availability and/or lower protection capability of TA compared to QUE. However, net N release was higher from QUE/BSA than from TA/BSA probably due to their higher protein-binding capacity and associated larger degradation of partly unprotected protein as suggested by ¹³C-CO₂ signatures. Microbial respiration patterns indicated that polyphenol complexes were initially degraded more quickly in the maize cropping system than in soils from under polyphenol-rich communities (Peltophorum and natural forest) but this pattern reversed with time. Long-term incubation of QUE/BSA complexes even caused a negative effect on microbial respiration in agricultural soils with low polyphenol contents (e.g. maize and sugarcane).Incubation of polyphenol complexes in soil depressed microbial biomass N in maize, sugarcane, Imperata and forest systems and led to reduced soil pH. However, microbial biomass was increased under the polyphenol-rich vegetation of Peltophorum. The PLFA group 18:2w6,9 was highly enhanced by condensed tannin-protein complexes additions as compared to control and hydrolysable polyphenol-protein complexes in soils with high polyphenol contents. Polyphenol complexes increased the fungi:bacteria ratio in systems with a high polyphenol content, particularly with condensed tannin complexes. The results indicated that systems with a high polyphenol content favoured development of fungal communities that are highly adaptable to phenol-rich soil conditions and high acidity, particularly with regards to the more recalcitrant condensed tannin-protein complexes.     

Organic and inorganic soil phosphates and acid phosphatase activity in the rhizosphere of 80-year-old Norway spruce [Picea abies (L.) Karst.] trees

Summary Inorganic and organic phosphates (P) were measured in bulk soil, rhizosphere soil and mycorrhizal rhizoplane soil of Norway spruce. Various methods of P extraction and estimation were compared. In addition, acid phosphatase activity and mycelial hyphae length were determined. In soil solutions from various locations, about 50% (range 35%–65%) of the total P was present as organic P. Compared to the bulk soil, the concentrations of readily hydrolysable organic P were lower in the rhizosphere soil and in the rhizoplane soil; this difference was particularly marked in the humus layer. In contrast, the concentrations of inorganic P either remained unaffected or increased. A 2- to 2.5-fold increase was found in the activity of acid phosphatase in the rhizoplane soil in comparison to the bulk soil. There was a positive correlation (r = 0.83^***) between phosphatase activity and the length of mycelial hyphae. The results stress the role of organic P and of acid phosphatase in the rhizosphere in the P uptake by mycorrhizal roots of spruce trees grown on acid soils.     

Characterization of dissolved organic matter in decomposing Norway spruce and silver birch litter

Dissolved organic matter (DOM) plays an important role in transport, storage and cycling of carbon (C) and nitrogen (N) in forest soils where litter is one of the main sources. The aim was to study the amount and characteristics of DOM leached from freshly fallen litters of silver birch (Betula pendula Roth.), Norway spruce (Picea abies (L.) Karst.) and their mixture during decomposition. DOM was collected after irrigation on eight occasions during 252 days incubation in the laboratory at about 18°C, including one freeze‐thaw cycle. During the incubation about 33–35% of C from birch and spruce litter and 40% of C from their mixture was lost. The total cumulative flux of dissolved organic carbon (DOC) from the mixture of litters was approximately 40% larger than that from single litters. The flux of DOC, DON, phenolic compounds and proteins followed a two‐stage pattern during decomposition. In the first stage the initially large fluxes decreased gradually. In the second stage, after freezing and thawing, the fluxes tended to increase again. Mixing birch and spruce litters and a freeze‐thaw cycle seems to increase the decomposition of litter and result in the increased flux of DOC, DON and phenolic compounds. The flux of hemicelluloses and the degradability of DOM were large at the first leaching occasion and decreased during the incubation. Birch had a 40% larger total flux of easily degradable DOM than spruce, supporting the previous consistent signs of greater microbial biomass and activities related to C and N cycling in soil under birch than under spruce. It is known that recalcitrant DOM might be stabilized whereas labile DOM may promote microbial activity and nutrient cycling. We conclude that the storage and cycling of C and N is affected by both tree species and degradation stage of litter in forest soils.