Seasonal effects of liming,irrigation, and acid precipitation on microbial biomass N in a spruce (Picea abies L.) forest soil

Summary Seasonal effects of liming, irrigation, and acid precipitation on microbial biomass N and some physicochemical properties of different topsoil horizons in a spruce forest (Picea abies L.) were measured throughout one growing season. The highest biomass N was recorded in autumn and spring in the upper soil horizons, while the lowest values were obtained in summer and in deeper horizons. The clearest differences between the different soil treatments were apparent in autumn and in the upper horizons. Liming increased the microbial biomass N from 1.7% of the total N content to 6.8% (Olf₁ layer) and from 1% to 2% of the total N content in the Of₂ layer. The main inorganic-N fraction in the deeper horizons was NO _inf3 ^sup- . An increase in cation exchange capacity was observed down to the Oh layer, while soil pH was only slightly higher in the Olf₁ and Of₂ layers after liming. The effects of irrigation were less marked. The microbial biomass N increased from 1.7% of total N to 4.8% in the Olf₁ layer and from 1% to 2% of total N in the Of₂ layer. In the Olf₁ layer an increase in C mineralization was observed. Acid precipitation decreased the microbial biomass N in the upper horizons from 4.8% of total N to 1.8% in the Olf₁ layer and from 2% to 0.5% in the Of₂ layer. No significant changes in soil pH were observed, but the decrease in cation exchange capacity may result in a decrease in the proton buffering capacity in the near future.     

Long-term experiments in selectively cut Norway spruce (Picea abies) forest

Selective cutting is one of many harvesting methods alternative to clear-cutting. Both the influence on different ecological systems and the effect on forest productivity of this harvesting method are important. In this study, the productivity in 16 long-term experimental plots in selectively cut Norway spruce (Picea abies) forest is analysed.     

Impact of ectomycorrhizas on the concentration and biodegradation of simple organic acids in a forest soil

Simple organic acids seem to be important in many soil processes including nutrient uptake in the rhizosphere and long‐term pedogenic processes such as podzolization. The factors regulating the concentrations of these acids in soil, however, remain poorly understood. We have investigated one set of factors, namely the impact of ectomycorrhizal (Paxillus involutus) and non‐mycorrhizal Picea abies seedlings and humic acid on the concentration and dynamics of organic acids in soil solution. We did so over 10 months in laboratory columns containing soil from the E horizon of a sandy forest soil. Several organic acids were identified in the solution extracted from the root zone including oxalic, citric, malonic, succinic, acetic, formic and lactic acids at concentrations ranging from <0.1 to 2.3 µm . Both plants and ectomycorrhizas had significant effects on the concentration of organic acids in soil solution. In contrast, omitting P from the irrigation water appeared to have little effect on the concentrations. The microbial mineralization kinetics of oxalate conformed well to a single Michaelis–Menten equation. Further, the soil with the mycorrhizas had a significantly faster mineralization of oxalic acid over a wide concentration range than did the soil without ectomycorrhizas and without plants. We conclude that the oxalate flux through the soil with both trees and mycorrhizas is much faster than is evident from measurements of solution concentration at steady state. Humic acid had little effect on the concentrations of organic acids or dynamics in the soil solution. Oxalic acid concentrations in the soil solution were correlated with hyphal length, rate of microbial mineralization, soil respiration, and shoot to root weight ratio. We conclude that both mycorrhizas and plants have a large impact on organic acid cycling in forest soils.     

Effects of acids on forest trees as measured by titration in vitro,inheritance of buffering capacity in Picea abies

The effect of acid precipitation on vegetation is the result of an interaction between acid and plant. The metabolism of plants is dependent on optimal pH-values, which are maintained by regulation. There are differences in the effectiveness of regulation under such exogenous influences as acidic precipitation. These differences can be related to the resistance of plants to acidic precipitation. Such differences were measured as buffering capacity of homogenized leaves during titration with acid. There are significant differences in buffering capacity between clones in Pinus spp. and Picea abies. A highly significant variance in buffering capacity also was found among families of P. abies. Calculations of genetical parameters show that the phenotypical variance of buffering capacity is governed mainly by genetical factors.     

Mobilization of dissolved organic matter, aluminium and iron in podzol eluvial horizons as affected by formation of metal-organic complexes and interactions with solid soil material

Interactions with dissolved organic matter (DOM) are generally believed to play a crucial role in the translocation of Al and Fe in acid sandy soils. Binding of Al and Fe to DOM affects their mobility in soils by altering sorption equilibria of charged sites on solid soil material, inducing precipitation of organo‐metallic complexes and preventing the formation of inorganic Al and Fe phases. The relative importance of the different processes, especially with respect to the translocation of Al, Fe and organic matter in podzols, remains unresolved. We determined the effect of the presence of solid soil material from the eluvial (AhE and AE, respectively) horizons of a Fimic Anthrosol and a Haplic Podzol on the metal‐to‐organic carbon (M/C) ratio in solution and the formation of dissolved organic Al and Fe complexes. Furthermore, we assessed the resulting influence on the mobilization of Al, Fe and DOM. Even under considerable metal loading, the M/C ratios and ‘free’ metal fractions in solution remained low and relatively constant, due to an apparent buffering by the solid phase and the formation of organo‐metal complexes in solution. The M/C ratios remained so low that significant precipitation of organo‐metal complexes due to saturation with metals was not found. The apparent buffering by the solid phase can be explained by a strong release of organic matter from solid soil material and adsorption of non‐complexed Al and Fe on solid organic matter upon metal addition. Adsorption of organo‐metal complexes most likely played only a minor role. The observations confirm the expected mobilization of Al, Fe and DOM in eluvial horizons and seem to indicate that even under fluctuating input of Al, Fe and DOM the soil solution will have a constant composition with respect to M/C ratios and percentage of Al and Fe present in dissolved organo‐metal complexes.     

云冷杉林土壤酶活性与土壤养分的研究

对不同生长状况云冷杉林土壤酶活性在不同土层进行比较,并对土壤酶与土壤养分及与林下植物的关系进行分析。结果表明：不同土层的土壤酶活性对云冷杉林死亡程度响应不同,0-15 cm土壤酶活性变化最为显著,随土层深度的增加,酶活性减小,且对云冷杉的死亡响应程度减弱;5种土壤酶活性对云冷杉死亡程度响应表现不一,脲酶和过氧化氢酶活性随死亡程度的加大对云冷杉死亡率响应较最大,而转化酶、磷酸酶及多酚氧化酶活性随云冷杉死亡程度的增加下降较小;土壤有机质的含量对云冷杉林死亡程度的响应较大,对照样地有机质含量与重度及极重度死亡样地间呈极显著相关关系（P〈0.01）,5种酶活性均与有机质、速效K含量极显著正相关（P〈0.01）;云冷杉林死亡率增加,其林下植物多样性指数亦呈增加趋势,随着云冷杉林死亡程度的增加林下灌木层多样性增加,而林下草本层的多样性先增加后减少;林下植物Shannon-Wiener指数和Pielou指数与土壤多酚氧化酶活性的相关性最强,其次是过氧化氢酶。     

Mobilization of aluminium in soil by acid deposition and its uptake by grass cut for hay – a Chemical Time Bomb

Abstract. Analyses of soil and hay samples collected from the Park Grass Experiment at Rothamsted during the last 137 years indicate slow but significant increases in KCl- and EDTA-extractable aluminium in soil and a sudden and very large recent increase in the concentration of aluminium in the herbage. The latter is associated with a sudden increase in the rate of acidification of the soil over the last 10–15 years and the mobilization of aluminium as the soil enters the aluminium buffer range -a Chemical Time Bomb. Such severe acidification from atmospheric inputs on a well-buffered soil illustrates how quickly an apparently stable situation can change as a result of acid deposition. It highlights the need to protect soils and plants from the effects of acidification by decreasing acid inputs or by liming.     

Soil biochemical and chemical changes in relation to mature spruce (Picea abies) forest conversion and regeneration

To investigate soil changes from forest conversion and regeneration, soil net N mineralization, potential nitrification, microbial biomass N, L‐asparaginase, L‐glutaminase, and other chemical and biological properties were examined in three adjacent stands: mature pure and dense Norway spruce (Picea abies (L.) Karst) (110 yr) (stand I), mature Norway spruce mixed with young beech (Fagus sylvatica) (5 yr) (stand II), and young Norway spruce (16 yr) (stand III). The latter two stands were converted or regenerated from the mature Norway spruce stand as former. The studied soils were characterized as having a very low pH value (2.9 – 3.5 in 0.01 M CaCl₂), a high total N content (1.06 – 1.94 %), a high metabolic quotient (qCO₂) (6.7 – 16.9 g CO₂ kg^–1 h^–1), a low microbial biomass N (1.1 – 3.3 % of total N, except LOf₁ at stand III), and a relatively high net N mineralization (175 – 1213 mg N kg^–1 in LOf₁ and Of₂, 4 weeks incubation). In the converted forest (stand II), C : N ratio and qCO₂ values in the LOf₁ layer decreased significantly, and base saturation and exchangeable Ca showed a somewhat increment in mineral soil. In the regenerated forest (stand III), the total N storage in the surface layers decreased by 30 %. The surface organic layers (LOf₁, Of₂) possessed a very high net N mineralization (1.5 – 3 times higher than those in other two stands), high microbial biomass (C, N), and high basal respiration and qCO₂ values. Meanwhile, in the Oh layer, the base saturation and the exchangeable Ca decreased. All studied substrates showed little net nitrification after the first period of incubation (2 weeks). In the later period of incubation (7 – 11 weeks), a considerable amount of NO₃‐N accumulated (20 – 100 % of total cumulative mineral N) in the soils from the two pure spruce stands (I, III). In contrast, there was almost no net NO₃‐N accumulation in the soils from the converted mixed stand (II) indicating that there was a difference in microorganisms in the two types of forest ecosystems. Soil microbial biomass N, mineral N, net N mineralization, L‐asparaginase, and L‐glutaminase were correlated and associated with forest management.     

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.     

Chemistry of soil organic matter as related to C : N in Norway spruce forest (Picea abies(L.) Karst.) floors and mineral soils

Due to high nitrogen deposition in central Europe, the C : N ratio of litter and the forest floor has narrowed in the past. This may cause changes in the chemical composition of the soil organic matter. Here we investigate the composition of organic matter in Oh and A horizons of 15 Norway spruce soils with a wide range of C : N ratios. Samples are analyzed with solid‐state ¹³C nuclear magnetic resonance (NMR) spectroscopy, along with chemolytic analyses of lignin, polysaccharides, and amino acid‐N. The data are investigated for functional relationships between C, N contents and C : N ratios by structural analysis. With increasing N content, the concentration of lignin decreases in the Oh horizons, but increases in the A horizons. A negative effect of N on lignin degradation is observed in the mineral soil, but not in the humus layer. In the A horizons non‐phenolic aromatic C compounds accumulate, especially at low N values. At high N levels, N is preferentially incorporated into the amino acid fraction and only to a smaller extent into the non‐hydrolyzable N fraction. High total N concentrations are associated with a higher relative contribution of organic matter of microbial origin.     

Spatial heterogeneity of soil solution chemistry in a mature Norway spruce (Picea abies (L.) Karst.) stand

The determination of the average soil solution concentrations in forest soils is hindered by the spatial heterogeneity of the soil conditions and the stand structure on all scales. The aim of this paper is to investigate the spatial heterogeneity of the soil solution chemistry within a mature stand of Norway spruce and to evaluate the implication of this heterogeneity for the sampling design for soil solutions. The site is a 140 years old Norway spruce stand of 2.5 ha located in the German Fichtelgebirge at 800 m elevation on granitic, deeply weathered bedrock. At 35 cm soil depth, 59 ceramic suction lysimeters (5 cm length, 2 cm diameter) were installed in a systematic grid of 25 · 25 m and soil solution was sampled at 3 dates in June and July 1994. The solutions were analysed for major cations and anions. Semi-variance of the concentrations at a given date revealed no systematic spatial patterns. The coefficients of variance of the element concentrations were between 36 and 298% with highest values for NH₄ ⁺-N. The implications of the observed heterogeneity for the appropriate number of replicates was investigated by Monte Carlo simulations. As an example, the probability that the measured average concentration of SO₄ ^2?-S is outside a ±10% range (related to the ‘true’ 59 lysimeter average) is about 68% if only 3 replicates would have been used, 41% with 10 replicates and 25% with 20 replicates. Due to the generally large spatial heterogeneity of the soil solution chemistry in forest soils the number of lysimeters used must be carefully adjusted to site conditions and the specific question.     

Organic acids in root exudates from Picea abies seedlings influenced by mycorrhiza and aluminum

Organic acid anions exuded from roots of Picea abies (Norway spruce) seedlings grown on glass beads in the presence and absence of mycorrhiza (Laccaria bicolor) and aluminum (Al) at pH 3.9 were measured. We wanted to test if the roots exuded more organic acid anions when exposed to Al and if mycorrhization influenced the exudation. Oxalate was exuded in far higher amounts than any other organic acid anion, with a maximum rate of 1.7 nmol (mg root DW)^–1 d^–1. Mycorrhizal roots exuded significantly more oxalate than nonmycorrhizal roots. The presence of Al did not enhance oxalate exudation. We conclude that the oxalate exuded constitutively by Picea abies/Laccaria bicolor may lead to rhizosphere oxalate concentrations that are relevant for Al resistance.     

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     

The effect of spruce (Picea abies Karst.) on soil development: an analytical and experimental approach

Changes in soil and biogeochemical processes following the replacement, 60 years ago, of part of a deciduous forest by a coniferous stand were evaluated by (i) a balance-sheet approach using soil and biomass element content analyses and the mineral flux, and (ii) the inclusion of test-mineral and resin bags in an integrative experimental approach. The comparison of soils under different stands shows that the change to spruce is inducing physical, chemical and biological modifications in soil properties. Changes in the humus layers lead to a reduction in mineralization and the mobilization of active organic acids in the system. Under spruce, the soil structural stability is reduced as fine clay is dispersed. Soil acidification is increased, linked to a decrease in soil pH and desaturation of the soil exchange complex. Soil mineralogy also indicates this tendency, particularly with an increase in the stability of Al intergrades. The biogeochemical cycle is modified especially for N, S and Al. with large changes in the internal fluxes between the ecosystem components, but the modification of the input-output balance due to spruce introduction is large only for S. The acidification is partly due to a change in litter quality, which inhibits biological activity, and partly because, during the dormant season, evergreen species intercept ‘dry and occult’ depositions from the atmosphere containing acidic or potentially acidic components. Use of test-mineral bags improves understanding and interpretation of the current soil mechanisms. The test mineral, a vermiculite, introduced into the soil or placed in the lysimeter flux very quickly undergoes measurable transformation characterized by desaturation and fixation of non-exchangeable Al in the interlayer zone, easily identified by Al speciation. The acidifying effect of spruce is clearly demonstrated by the experimental approach used, including the characterization of soil solutions by resin bags. All the techniques are complementary and could be used together or individually, depending on the aim of the study.     

Long-term development of element budgets in a Norway spruce (Picea abies (L.) Karst.) forest of the German solling area

To evaluate ecosystem response to changing atmospheric deposition, element budgets were established over the period from 1973 to 1991 for a Norway Spruce (Picea abies (L.) Karst.) site. Budgets for Na⁺, Cl^?, Ca²⁺, Mg²⁺, N, S and H⁺ were based on total deposition and seepage water fluxes. The deposition of Ca²⁺, Mg²⁺, particularly, of S and H⁺ decreased with time, while calculated N deposition remained constant at a high level. The decrease in Ca²⁺ deposition led to a reduction of Ca²⁺ fluxes with seepage water. The decrease of Mg²⁺ deposition did not have an effect on the output fluxes of Mg²⁺. The reversibility of soil and seepage water acidification by reduced S deposition was delayed by the release of previously accumulated soil SO ₄ ^2? . The highest NO ₃ ^? fluxes were observed during the period of 1986 to 1988; NO ₃ ^? fluxes in general demonstrated a considerable annual and periodic variation. Total N accumulation in the ecosystem amounted to nearly 590 kg ha^?1 yr^?1 during the observation period. The major sink of N in the spruce site is the aggrading humus layer. The results emphasize the need for measurements over several years to make conclusions regarding the function of ecosystems in response to atmospheric deposition.     

Accumulation of,and interactions between,calcium and heavy metals in wood and bark of Picea abies

Spreading of wood ashes from the pulp and paper industry will change the content and proportions of calcium (Ca), copper (Cu), cadmium (Cd), and zinc (Zn) in forest soils and thus also in the forest trees. The accumulation and distributions of, and interaction between, Ca and heavy metals in wood and bark of two‐year‐old Norway spruce (Picea abies [L.] Karst.) were investigated in this study. The treatment was carried out for 3 months in nutrient solutions, and there was a low or a high addition of Ca, Cd, Cu or Zn. The metal accumulation in, and distribution between, the bark, the wood formed during the treatment period (new wood), and the wood formed before the treatment period (old wood) was analyzed with AAS. The contents of the metals in the stems (i.e., bark, new wood, old wood) increased with elevated addition of the metal in question, also at the low addition of Ca, Cu, and Zn. Interactions between Ca and the heavy metals were found. Elevated Ca additions decreased the Cd content of the bark and the Zn content of the old wood, and tended to decrease the Cu content of the bark and the Cd content of the old wood. The Ca content decreased in both, wood and bark after Cu addition and the high Cd addition. Thus, even small changes in metal availability and proportions in forest soil, such as after spreading of wood ashes in the forest, will be reflected in the content of the metals in the wood and bark of forest trees.     

Mobilization of X‐ray amorphous and crystalline aluminum and iron phosphates by common soil extraction procedures

Soil extraction with 0.1 M NaOH completely dissolves X‐ray amorphous as well as crystalline AlPO₄ and FePO₄. Extraction with acidic NH₄ oxalate completely mobilizes X‐ray amorphous AlPO₄ and FePO₄ in addition to phosphate adsorbed to poorly‐crystallized Al and Fe minerals, but only a minor portion of crystalline AlPO₄ and FePO₄. Extraction with dithionite‐citrate‐bicarbonate mobilizes X‐ray amorphous AlPO₄ and FePO₄ as well as crystalline FePO₄, but only a minor portion of crystalline AlPO₄.     

Molybdate sorption by oxides of aluminium and iron

The sorption of molybdate by goethite, hematite, bayerite and α-Al₂O₃ was studied as a function of molybdate concentration and pH. The sorption isotherms exhibited double sorption plateaus in which the amount of Mo sorbed in the second plateau was double that in the first. This was attributed to the polymerization of molybdates as the concentration increases. Furthermore Mo sorption was found to be associated with a cosorption of Na cations, probably present to maintain electroneutrality in the surface zone. The sorption showed high sensitivity towards the pH, attaining a maximum at pH 4. For the iron oxides the decrease in sorption was much less pronounced on the acid side of the pH maximum and occured at lower pH values than that of the Al oxides. Molybdate sorption is explained in terms of a ligand exchange reaction between molybdate and surface hydroxyls.