Effects of drought stress on photosynthesis,rhizosphere respiration,and fine‐root characteristics of beech saplings: A rhizotron field study

Soil drought influences the C turnover as well as the fine‐root system of tree saplings. Particularly during the period of establishment, the susceptibility to drought stress of saplings is increased because of incompletely developed root systems and reduced access to soil water. Here, we subjected beech saplings (Fagus sylvatica L.) to different levels of drought stress. Beech saplings were planted in rhizotrons, which were installed in the soil of a Norway spruce forest before bud burst. Soil moisture was manipulated in the following year during May to September. We measured photosynthetic net CO₂ uptake, volume production of fine roots, and rhizosphere respiration during the growing season. Biometric parameters of the fine‐root system, biomass, and nonstructural carbohydrates were analyzed upon harvest in October. Photosynthesis and rhizosphere respiration decreased with increasing drought‐stress dose (cumulated soil water potential), and cumulative rhizosphere respiration was significantly negatively correlated with drought‐stress dose. Fine‐root length and volume production were highest at moderate soil drought, but decreased at severe soil drought. The proportion of fine‐roots diameter < 0.2 mm and the root‐to‐shoot ratio increased whereas the live‐to‐dead ratio of fine roots decreased with increasing drought‐stress dose. We conclude that the belowground C allocation as well as the relative water‐uptake efficiency of beech saplings is increased under drought.     

Image analysis for non‐destructive and non‐invasive quantification of root growth and soil water content in rhizotrons

Studies aiming at quantification of roots growing in soil are often constrained by the lack of suitable methods for continuous, non‐destructive measurements. A system is presented in which maize (Zea mays L.) seedlings were grown in acrylic containers — rhizotrons — in a soil layer 6‐mm thick. These thin‐layer soil rhizotrons facilitate homogeneous soil preparation and non‐destructive observation of root growth. Rhizotrons with plants were placed in a growth chamber on a rack slanted to a 45° angle to promote growth of roots along the transparent acrylic sheet. At 2‐ to 3‐day intervals, rhizotrons were placed on a flatbed scanner to collect digital images from which root length and root diameters were measured using RMS software. Images taken during the course of the experiment were also analyzed with QUACOS software that measures average pixel color values. Color readings obtained were converted to soil water content using images of reference soils of known soil water contents. To verify that roots observed at the surface of the rhizotrons were representative of the total root system in the rhizotrons, they were compared with destructive samples of roots that were carefully washed from soil and analyzed for total root length and root diameter. A significant positive relation was found between visible and washed out roots. However, the influence of soil water content and soil bulk density was reflected on seminal roots rather than first order laterals that are responsible for more than 80 % of the total root length. Changes in soil water content during plant growth could be quantitifed in the range of 0.04 to 0.26 cm³ cm^—3 if image areas of 500 x 500 pixel were analyzed and averaged. With spatial resolution of 12 x 12 pixel, however, soil water contents could only be discriminated below 0.09 cm³ cm^—3 due to the spatial variation of color readings. Results show that this thin‐layer soil rhizotron system allows researchers to observe and quantify simultaneously the time courses of seedling root development and soil water content without disturbance to the soil or roots.     

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.     

Different effect of drying on the fluxes of dissolved organic carbon and nitrogen from a Norway spruce forest floor

The forest floor represents the major source of dissolved organic carbon (DOC) and nitrogen (DON) in forest soils. The release mechanisms of DOC and DON from forest floors and their environmental controls as well as the dynamics of concentrations and fluxes are still poorly understood. We investigated the effect of drying and rewetting on the release of DOC and DON from a Norway spruce forest floor. Undisturbed soil columns of 17 cm diameter and 15—20 cm height were taken with 7 replicates from the forest floor of a mature Norway spruce (Picea abies [L.] Karst.) site and established at 10°C in the laboratory. Columns were exposed to different periods of drying (3, 5, 10, 20 days). Each drying period was followed by a rewetting for 5 days at an irrigation rate of 10 mm d^—1 with a natural throughfall solution. The percolates from the forest floor were collected daily and analyzed for DOC, total N, NH₄, NO₃, pH, electrical conductivity and major ions. Drying for 10 and 20 days decreased the water content of the Oi horizon from 280% dry weight to about 30%. The water content of the Oe and the Oa horizon only changed from about 300% to 200%. The fluxes of DOC from the forest floor were moderately effected by drying and rewetting with an increase after 3 and 5 days of drying, but a decrease after 10 and 20 days. On the contrary, the drying for 10 and 20 days resulted in a drastic increase of the DON fluxes and a subsequent decrease of the DOC/DON ratios in the forest floor percolates from about 50 to 3.3. These results suggest that the mechanisms for DOC release in forest floors differ from those for DON and that drying and rewetting cause temporal variations in the DOC/DON ratios in forest floor percolates.     

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.     

Effects of experimental soil frost on the fine‐root system of mature Norway spruce

Soil‐frost events may influence the dynamics of fine roots and therefore affect root‐derived C fluxes to the soil. We studied the impact of soil frost on the fine‐root dynamics of Norway spruce (Picea abies [L.] Karst.) by a replicated snow‐removal experiment in a mature forest in SE Germany. Snow removal in the three treatment plots reduced soil temperature significantly with minima <–5.5°C in the O layer while the snow‐covered control plots never reached temperatures below the freezing point. Sequential soil coring in the O layer at the beginning and at the end of the soil‐frost period as well as after thawing revealed that the soil frost treatment increased fine‐root mortality by 29%. However, enhanced fine‐root production in the snow‐removal plots nearly compensated for the fine‐root losses caused by low temperatures. These findings were confirmed by minirhizotron observations in the O layer and the upper 25 cm of the mineral soil showing that relative fine‐root loss was by far higher in the snow‐removal plots than in the control plots. Compensatory fine‐root production in the snow‐removal plots exceeded fine‐root production in the control plots during a period of 8 weeks after the soil frost application by 39% in the O layer while it was similar in both plot types in the mineral soil. Sequential coring and minirhizotron observations led to substantially different fine root–longevity estimates for the soil frost period. However, in both cases, the snow‐removal treatment was characterized by a significant reduction in root longevity indicating a faster fine‐root turnover. As a consequence, experimental soil frost enhanced the C input to the soil via root death at our study site by approx. 42 g m^–2 and stimulated the C investment towards the root system of the spruce trees due to a higher sink activity.     

Short‐Term Recovery of Ammonium‐15Nitrogen Applied to a Temperate Forest Inceptisol and Ultisol in East Tennessee,USA

Abstract

The short‐term fate and retention of ammonium (NH₄)‐¹⁵nitrogen (N) applied to two types of forest soils in east Tennessee was investigated. Four ridgetop forests, predominantly oak (Quercus spp.), were studied. Five applications of NH₄‐¹⁵N tracer were made to the forest floor at 2‐ to 4‐week intervals over a 14‐week period in 2004. Nitrogen‐15 recovery in the forest floor, fine roots (<2 mm), and the mineral soil (0–20 cm) was calculated at 6, 21, and 42 weeks after the last application. Most of the ¹⁵N was retained in the forest floor and the mineral soil, with only small amounts (≤2%) found in roots from both soil layers. Recovery of NH₄‐¹⁵N was greater in Inceptisols, which had a wider carbon (C)‐to‐N ratio than Ultisols. For both soil types, higher NH₄‐¹⁵N recoveries and long retention times (half‐lives>100 weeks) indicated the forest floor is an effective filter for atmospheric N inputs.     

Effects of growing roots of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica L.) on rhizosphere soil solution chemistry

The chemical conditions of the rhizosphere can be very different from that of bulk soil. Up to now, little attention has been given to the problem of spatial heterogeneity and temporal dynamics of rhizosphere soil solution and little is known about the influence of different tree species on rhizosphere chemistry. In the present study, we used micro suction cups to collect soil solution from the rhizosphere of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica L.) seedlings in high spatial resolution and capillary electrophoresis for the determination of major cations and anions. The results indicate, that in a soil with a base saturation of about 20—25% and a pH of 6.5, growing roots of beech and spruce lower the concentrations of nutrient cations and nitrate in the rhizosphere soil solution and decrease significantly the pH. The H⁺ release leads to an enhanced mineral weathering as indicated by an increase of CEC and base saturation and to a mobilization of soluble Al, however, on a very low concentration level. In our experiment rhizosphere effects of spruce have been more pronounced than those of beech, indicating, that with respect to below ground activity young spruce trees have a better competitive power than beech.     

Long‐term development of nitrogen fluxes in a coniferous ecosystem: Does soil freezing trigger nitrate leaching?

In many forest ecosystems chronically large atmospheric deposition of N has caused considerable losses of inorganic N by seepage. Freezing and thawing of soil may alter the N turnover in soils and thereby the interannual variation of N seepage fluxes, which in turn makes it difficult to evaluate the N status of forest ecosystems. Here, we analyzed long‐term monitoring data of concentrations and fluxes of dissolved inorganic N (DIN) in throughfall and seepage from a Norway spruce stand at the Fichtelgebirge (SE Germany) between 1993 and 2004. Despite constant or even slightly increasing N inputs in throughfall, N losses with seepage at 90 cm declined from 15–32 kg N ha^–1 y^–1 in the first years of the study period (1993–1999) to 3–10 kg N ha^–1 y^–1 in 2000 to 2004. The large N losses in the first years coincided with extreme soil frost in the winter of 1995/96, ranging from –3.3°C to –1.0°C at 35 cm soil depth. Over the entire observation period, maximum fluxes of nitrate and ammonium were observed in the mineral soil following thawing of the soil. The elevated ammonium and nitrate fluxes resulted apparently from increased net ammonification and nitrification rates in the mineral soil, whereas mineral‐N fluxes in the O horizon were less affected by frost. Our data suggest that (1) extreme soil frost may cause substantial annual variations of nitrate losses with seepage and that (2) the assessment of the N status of forest ecosystems requires long periods of monitoring. Time series of biogeochemical data collected over the last 20–30 y include years with extreme cold winters and warm summers as well as unusual precipitation patterns. Analysis of such long‐term monitoring data should address climate extremes as a cause of variation in N outputs via leaching. The mean loss of 14.7 kg N with seepage water during 12 y of observation suggests that the forest ecosystem was saturated with N.     

Elemental patterns in roots and foliage of mature spruce across a gradient of soil aluminium

Tissue concentrations of Al in red and Norway spruce trees were compared across 5 sites in North America and Europe as part of an investigation of Al biogeochemistry in forested ecosystems (ALBIOS). Fine roots and foliage were sampled and analyzed for Al, Ca, Mg, and P, and the chemistry of soil and soil solutions was characterized at each plot by horizon. Sites exhibited a wide range in soil Al saturation and in concentrations of Al and sulfate in lysimeter solutions. Aluminium concentrations in roots were two orders of magnitude higher than those in foliage. Fine roots (<1.0 mm) from B horizons had the highest Al concentrations and appeared to be the best phytoindicators of plant-available Al. Aluminium concentrations in fine roots from B horizons were highly correlated with soil solution monomeric Al, and with Al in 0.01 M SrC₂. soil extracts. Stronger soil Al extractants were generally poor predictors of concentrations of Al in plant tissue. Sites with higher levels of plant-available Al supported spruce trees with correspondingly lower foliar levels of Ca and Mg. As such, these field sites provided circumstantial evidence that Al may be interfering with Ca and Mg uptake and transport. No evidence was found of Al interference with P uptake or transport at these sites.     

Effects of forest vegetation on spatial variability of surface mineral soil pH,soluble aluminum and carbon

The purpose of this study was to examine spatial variability in forest soils at several levels including variability due to soil structure, to the presence of individual trees and to populations of different species of trees. Both classic statistical and geostatistical methods were used. Soil chemical properties measured include pH and Al and C in solution which was in equilibrium with the surface mineral soil. Results indicated that soil cores 1.8 cm in diameter were as effective as larger cores in incorporating variability in surface mineral soil pH. There was no spatial correlation in soil pH in samples separated by a distance of 20 to 360 cm. The presence of individual Norway spruce and red pine trees affected soil in their vicinity. Soil pH was depressed and soluble Al elevated. in soil near the base of the tree compared to soil 120 cm from the tree, independent of direction. In addition, in soil sampled at least 100 cm from the base of trees, pH was lower in Norway spruce compared to sugar maple plantations and soluble C was greater in red pine compared to Norway spruce plantations. It is concluded that in less than 50 yr the presence of individual trees and populations of different tree species can affect chemical properties of surface mineral soils. These effects should be considered in the design and interpretation of experiments.     

Mapping water,oxygen, and pH dynamics in the rhizosphere of young maize roots

Rhizosphere processes are highly dynamic in time and space and strongly depend on each other. Key factors influencing pH changes in the rhizosphere are root exudation, respiration, and nutrient supply, which are influenced by soil water content levels. In this study, we measured the real‐time distribution of soil water, pH changes, and oxygen distribution in the rhizosphere of young maize plants using a recently developed imaging approach. Neutron radiography was used to capture the root system and soil water distribution, while fluorescence imaging was employed to map soil pH and soil oxygen changes. Germinated seeds of maize (Zea mays L.) were planted in glass rhizotrons equipped with pH and oxygen‐sensitive sensor foils. After 20 d, the rhizotrons were wetted from the bottom and time‐lapsed images via fluorescence and neutron imaging were taken during the subsequent day and night cycles for 5 d. We found higher water content and stronger acidification in the first 0.5 mm from the root surface compared to the bulk soil, which could be a consequence of root exudation. While lateral roots only slightly acidified their rhizosphere, crown roots induced stronger acidification of up to 1 pH unit. We observed changing oxygen patterns at different soil moisture conditions and increasing towards lateral as well as crown roots while extending laterally with ongoing water logging. Our work indicates that plants alter the rhizosphere pH and oxygen also depending on root type, which may indirectly arise also from differences in age and water content changes. The results presented here were possible only by combining different imaging techniques to examine profiles at the root‐soil interface in a comprehensive way during wetting and drying.     

Effects of drying and rewetting on soluble phosphorus and nitrogen in forest floors: An experiment with undisturbed columns

Drying and rewetting (D/W) of soils often resulted in the release of soluble phosphorus (P) and nitrogen (N), thereby changing the availability of both nutrients. Most experiments on D/W have been conducted with disturbed mineral soil samples and with rewetting of the soil samples by abrupt change in the water potential. Here, we studied the effect of D/W on the leaching of P and N from undisturbed forest floors of a European beech and a Norway spruce site under near field conditions of desiccation and rewetting. We hypothesized that even under realistic rewetting of undisturbed forest floors, the leaching of P and N is increased after D/W and that the effects are less pronounced for spruce than for beech because of the larger hydrophobicity of the spruce forest floor. Undisturbed forest floor columns were subjected to desiccation at 20°C until a matrix potential of –100 MPa (pF 6.0) was reached, while controls were kept at moist conditions. Columns were irrigated by 22 mm day^?1 from day 1–3 and by 10 mm day^?1 from day 4–14 given in automated short pulses. Leachates from the soil columns were analyzed for orthophosphate, total P, NH₄, NO₃, and total N. In the spruce forest floor the concentrations of total P in leachates and the leachate fluxes were strongly increased after D/W. The increase of solute P was less for beech than for spruce coinciding with less actual rewetting of the beech forest floor. Leaching of total N from the spruce forest floor was not affected by D/W, however, concentrations and leaching of NH₄ increased, while leaching of NO₃ decreased. For beech the leaching of total N and NH₄ increased after D/W, while NO₃ leaching decreased. The results indicate that also under realistic conditions, D/W of forest floors increases solute P and leads to changes in the ratio of NH₄/NO₃ in solution, thereby altering the availability of the nutrients.     

Comparing microbial biomass, denitrification enzyme activity, and numbers of nitrifiers in the rhizospheres of Pinus sylvestris, Picea abies and Betula pendula seedlings by microscale methods

 Flushes of C and N from fumigation-extraction (FE-C and FE-N, respectively), substrate-induced respiration (SIR), denitrification enzyme activity (DEA) and numbers of NH₄ ⁺ and NO₂ ^– oxidizers were studied in the rhizospheres of Scots pine (Pinus sylvestris L.), Norway spruce [(Picea abies (L.) Karsten] and silver birch (Betula pendula Roth) seedlings growing in soil from a field afforestation site. The rhizosphere was defined as the soil adhering to the roots when they were carefully separated from the rest of the soil in the pots, termed as "planted bulk soil". Soil in unplanted pots was used as control soil. All seedlings had been grown from seed and had been infected by the natural mycorrhizas of soil. Overall, roots of all tree species tended to increase FE-C, FE-N, SIR and DEA compared to the unplanted soil, and the increase was higher in the rhizosphere than in the planted bulk soil. In the rhizospheres tree species did not differ in their effect on FE-C, FE-N and DEA, but SIR was lowest under spruce. In the planted bulk soils FE-C and SIR were lowest under spruce. The planted bulk soils differed probably because the roots of spruce did not extend as far in the pot as those of pine and birch. The numbers of both NH₄ ⁺ and NO₂ ^–oxidizers, determined by the most probable number method, were either unaffected or decreased by roots, with the exception of the spruce rhizosphere, where numbers of both were increased. Received: 26 August 1998     

Effects of decreasing soil water content on seminal lateral roots of young maize plants

Soil micropores that contain water at or below field capacity cannot be invaded by seminal or first‐order lateral roots of maize plants because their root diameters are larger than 10 μm. Hence, at soil‐water levels below field capacity plant roots must establish a new pore system by displacement of soil particles in order to access soil water. We investigated how decreasing soil water content (SWC) influences growth and morphology of the root system of young maize plants. Plants were grown in rhizotrons 40 cm wide, 50 cm high, and approximately 0.7 cm thick. Five SWC treatments were established by addition of increasing amounts of water to soil and thorough mixing before filling the rhizotrons. No water was added to treatments 1–4 throughout the experiment. Treatment 5 was watered frequently throughout the experiment to serve as a control. Seminal‐root length and SWC in soil layers 0–10, 10–20, 20–30, 30–40, and 40–50 cm were measured at intervals of 2–3 d on scanner images by image analysis. At 15 d after planting, for treatments 1–4 shoot dry weight and total root length were directly related to the amount of water added to the soil, and for treatments 4 and 5, total root length and shoot dry weights were similar. Length of seminal roots visible at the transparent surface of the rhizotron for all treatments was highest in the uppermost soil layer and decreased with distance from the soil surface. For all layers, seminal‐root elongation rate was at maximum above a SWC of 0.17 cm³ cm^–3, corresponding to a matric potential of –30 kPa. With decreasing SWC, elongation rate decreased, and 20% of maximum seminal root elongation rate was observed below SWC of 0.05 cm³ cm^–3. After destructive harvest for treatment 1–4, number of (root‐) tips per unit length of seminal root was found uninfluenced over the range of initial SWC from 0.10 to 0.26 cm³ cm^–3. However, initial SWC close to the permanent wilting point strongly increased number of tips. Average root length of first‐order lateral (FOL) roots increased as initial SWC increased, and the highest length was found for the frequently watered treatment 5. The results of the study suggest that the ability to produce new FOL roots across a wide range of SWC may give maize an adaptive advantage, because FOL root growth can rapidly adapt to changing soil moisture conditions.     

Mycorrhizal colonization and lead distribution in root tissues of Norway spruce seedlings

Non-inoculated spruce seedlings (Picea abies Karst.) and spruce seedlings colonized with Lactarius rufus Fr., Pisolithus tinctorius Coker & Couch or Paxillus involutus Fr. were grown for 35 or 37 weeks in a microcosm system on two types of natural forest humus differing in Pb content. Using X-ray microanalysis, the distribution and content of Pb in the tissues of mycorrhizal and non-mycorrhizal root tips were compared. No significant difference in the Pb contents of root cortex cell walls of non-mycorrhizal and seedlings colonized by Lactarius rufus, Pisolithus tinctorius, or indigenous mycorrhizal fungi (mainly Tylospora sp.) was found. However, in root tips of seedlings colonized by Paxillus involutus, due to a higher binding capacity for cations, the Pb content in cell walls of the root cortex were higher than in non-mycorrhizal roots. Pb contents in cell walls of the cortex of mycorrhizal and non-mycorrhizal roots were 3 times higher in plants growing in humus with a high Pb content than in plants growing in humus with a low Pb content. It is concluded that increasing contents of Pb in the organic matter may lead to an increased loading of the apoplast with Pb. The mycobionts tested in this investigation did not exclude Pb from root tissues.     

Are Indicators for Critical Load Exceedance Related to Forest Condition?

The aim of this study was to evaluate the suitability of the (Ca?+?Mg?+?K)/Al and the Ca/Al ratios in soil solution as chemical criteria for forest condition in critical load calculations for forest ecosystems. The tree species Norway spruce, Sitka spruce and beech were studied in an area with high deposition of sea salt and nitrogen in the south-western part of Jutland, Denmark. Throughfall and soil water were collected monthly and analysed for pH, NO₃-N, NH₄-N, K, Ca, Mg, DOC and Al_tot. Organic Al was estimated using DOC concentrations. Increment and defoliation were determined annually, and foliar element concentrations were determined every other year. The throughfall deposition was highest in the Sitka spruce stand (maximum of 40 kg N ha^?1yr^?1) and lowest in the beech stand (maximum of 11 kg N ha^?1yr^?1). The Sitka spruce stand leached on average 12 kg N ha^?1yr^?1 during the period 1988–1997 and leaching increased throughout the period. Only small amounts of N were leached from the Norway spruce stand whereas almost no N was leached from the beech stand. For all tree species, both (Ca?+?Mg?+?K)/Al and Ca/Al ratios decreased in soil solution at 90 cm depth between 1989 and 1999, which was mainly caused by a decrease in concentrations of base cations. The toxic inorganic Al species were by far the most abundant Al species at 90 cm depth. At the end of the measurement period, the (Ca?+?Mg?+?K)/Al ratio was approximately 1 for all species while the Ca/Al ratio was approximately 0.2. The lack of a trend in the increment rates, a decrease in defoliation as well as sufficient levels of Mg and Ca in foliage suggested an unchanged or even slightly improved health condition, despite the decreasing and very low (Ca?+?Mg?+?K)/Al and Ca/Al ratios. The suitability of these soil solution element ratios is questioned as the chemical criteria for soil acidification under field conditions in areas with elevated deposition rates of sea salts, in particular Mg.     

Heavy Metal Concentrations in Soil Solution, Soil and Needles in a Norway Spruce Stand on an Acid Sulphate Forest Soil

The effects of soil processes, related to the oxidation of sulphide sediments, on heavy metal concentrations in the soil and soil solution were investigated in a Norway spruce stand on a fine-textured, acidic soil rich in sulphates located on the isostatic land-uplift western coast of Finland. The age of the soil is ca. 300–400 years, and the soil texture is silt and till. The chemical properties of the soil and soil solution clearly reflected the formation of acid sulphate (AS) soil. Compared to background reference values for podzolic coniferous forest soil, the pH of the soil solution in the mineral soil (20–40 cm depth) was very low, and the Al, Fe and S concentrations extremely high. The Zn and Ni concentrations in the soil solution were also strongly elevated, and similar to the concentrations reported close to anthropogenic heavy-metal emission sources. The concentrations of Cd and Cu were also frequently elevated. In contrast, the acidity and metal concentrations of the soil solution sampled in the organic layer were not elevated. Similarly, exchangeable Zn and Ni concentrations were also elevated in the mineral soil, but not in the organic layer. Because Norway spruce has a very superficial rooting system and the zone with exceptionally high metal concentrations did not extend up to the topmost soil layers, sulphide-oxidation derived soil acidification is not likely to pose a serious threat to forest ecosystems growing on this type of site. Despite the elevated concentrations of protons and many metals in the mineral soil and soil solution (20–40 cm), the nutrient status of the spruce stand was satisfactory and the general health of the stand has been reported to be relatively good.     

Conversion of cropland to forest increases soil CH4 oxidation and abundance of CH4 oxidizing bacteria with stand age

We investigated CH₄ oxidation in afforested soils over a 200-year chronosequence in Denmark including different tree species (Norway spruce, oak and larch) and ages. Samples of the top mineral soil (0–5 cm and 5–15 cm depth) were incubated and analyzed for the abundance of the soil methane-oxidizing bacteria (MOB) and ammonia-oxidizing bacteria (AOB) and archaea (AOA) based on quantitative PCR (qPCR) on pmoA and amoA genes. Our study showed that CH₄ oxidation rates and the abundance of MOB increased simultaneously with time since afforestation, suggesting that the methanotrophic activity is reflected in the abundance of this functional group.The development of forest soils resulted in increased soil organic carbon and reduced bulk density, and these were the two variables that most strongly related to CH₄ oxidation rates in the forest soils. For the top mineral soil layer (0–5 cm) CH₄ oxidation rates did not differ between even aged stands from oak and larch, and were significantly smaller under Norway spruce. Compared to the other tree species Norway spruce caused a decrease in the abundance of MOB over time that could explain the decreased oxidation rates. However, the cause for the lower abundance remains unclear. The abundance of ammonia-oxidizers along the chronosequence decreased over time, oppositely to the MOB. However, our study did not indicate a direct link between CH₄ oxidation rates and ammonia-oxidizers. Here, we provide evidence for a positive impact of afforestation of former cropland on CH₄ oxidation capacity in soils most likely caused by an increased population size and activity of MOB.     

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