Dissolved organic carbon concentrations in four Norway spruce stands of different ages

Boreal forests are increasing in age partly due to reduced logging and efficient wildfire control. As a result, they also stock more carbon. Whether increased forest C stock causes greater production of dissolved organic carbon (DOC) is uncertain. DOC in bulk precipitation, throughfall and soil water was studied in 10-, 30-, 60- and 120-year-old stands of Norway spruce (Picea abies (L.) Karst.) DOC concentrations in throughfall and O horizon soil water followed the order 10 < 30 < 60 = 120 and 10 = 30 < 120 < 60, respectively. DOC fluxes followed the order 10 = 30 < 60 = 120 in throughfall, while no significant difference between stands was found for O horizon soil water. Above-ground tree litter varied according to 10 < 30 < 60 = 120, a pattern identical to that for DOC concentrations in throughfall and resembling but not identical to that for DOC concentrations in O horizon soil water. This indicates additional sources for DOC in soil water. Seasonality in DOC concentrations was observed at the base of the O horizon, and seasonality in DOC fluxes in both throughfall and O horizon soil water. Our results suggest differences in the polarity of DOC between the 10-year stand and the others, which we interpret as reflecting the lack of grown trees and possibly the different vegetation on the 10-year stand.     

Effects of soil frost on soil respiration and its radiocarbon signature in a Norway spruce forest soil

Apart from a general increase of mean annual air temperature, climate models predict a regional increase of the frequency and intensity of soil frost with possibly strong effects on C cycling of soils. In this study, we induced mild soil frost (up to −5 °C in a depth of 5 cm below surface) in a Norway spruce forest soil by removing the natural snow cover in the winter of 2005/2006. Soil frost lasted from January to April 2006 and was detected down to 15 cm depth. Soil frost effectively reduced soil respiration in the snow removal plots in comparison to undisturbed control plots. On an annual basis 6.2 t C ha⁻¹ a⁻¹ were emitted in the control plots compared with 5.1 t C ha⁻¹ a⁻¹ in the snow removal plots. Only 14% of this difference was attributed to reduced soil respiration during the soil frost period itself, whereas 63% of this difference originated from differences during the summer of 2006. Radiocarbon (Δ¹⁴C) signature of CO₂ revealed a considerable reduction of heterotrophic respiration on the snow removal plots, only partly compensated for by a slight increase of rhizosphere respiration. Similar CO₂ concentrations in the uppermost mineral horizons of both treatments indicate that differences between the treatments originated from the organic horizons. Extremely low water contents between June and October of 2006 may have inhibited the recovery of the heterotrophic organisms from the frost period, thereby enhancing the differences between the control and snow removal plots. We conclude that soil frost triggered a change in the composition of the microbial community, leading to an increased sensitivity of heterotrophic respiration to summer drought. A CO₂ pulse during thawing, such as described for arable soils several times throughout the literature, with the potential to partly compensate for reduced soil respiration during soil frost, appears to be lacking for this soil. Our results from this experiment indicate that soil frost reduces C emission from forest soils, whereas mild winters may enhance C losses from forest soils.     

Seasonal and spatial variability of soil respiration in four Sitka spruce stands

We investigated the causes for the seasonal and spatial variation of soil respiration in a first rotation Sitka spruce chronosequence composed of four age classes (10, 15, 31, and 47 year old) in Central Ireland. The study aimed at identifying easily determinable environmental parameters that explained the variation in soil respiration rates. The variation in temperature and soil water content influenced the seasonal trend observed in the spatial variability of soil respiration. The highest coefficients of variation in soil respiration were observed during autumn drought, while lower coefficients were generally observed during periods with highest soil respiration rates. On average, the sampling strategy of 30 sampling points per stand was adequate to obtain an average rate of soil respiration within 20% of its actual value at the 95% confidence level. Significantly higher soil respiration rates were observed at locations with high accumulation of organic matter and in collars established in close vicinity to tree stems. The organic layer thickness was the only variable that yielded significant regressions for explaining spatial variation in soil respiration in all the stands. Correlation analyses between the studied variables and soil respiration suggested the relative importance of heterotrophic and autotrophic components differed in their annual contribution to total soil respiration at each forest stand. Multiple regression analyses were used to assess the relative importance of primary temporal and spatial controls over soil respiration. Soil temperature and organic layer thickness explained most of the variance of soil respiration for the different sampling periods, while soil water content had a weaker effect as well as a different influence on soil respiration depending on the time of the year. The strong linear correlation between forest floor carbon and soil carbon stock further confirmed organic layer thickness as an integrative factor encompassing the effect of soil carbon pools on soil respiration. Moreover, its inclusion in the multiple regression analyses overrode the influence of both distance and fine root biomass. Overall, a multiple linear regression model driven by easily determinable environmental variables such as soil temperature, organic thickness, soil water content, soil bulk density, and soil organic carbon concentration allowed us to explain 54% of total variance of soil respiration over the different stand ages for the entire year (P < 0.05). Our results show that the adoption of an adequate sampling strategy, and the determination of some key environmental variables may help to explain a large proportion of total variation of soil respiration over the entire rotation length of afforested ecosystems.     

The effect of nutrition on the seasonal course of needle respiration in Norway spruce stands

 Respiration of 1-year-old needles of 30-year-old Norway spruce trees [Picea abies (L.) Karst.] was studied in a nutrient optimisation experiment in northern Sweden. Respiration rates of detached needles, from ten control (C) and ten irrigated-fertilised (IL) trees, were measured on 16 occasions from June 1992 to June 1993. The aim of the study was to determine the influence of temperature on the seasonal course of needle maintenance respiration, and the effect of nitrogen concentration [N] and carbohydrate content on needle respiration in young Norway spruce trees subjected to long-term fertilisation. The IL treatment significantly affected needle size, in terms of dry mass and length, but not specific needle length (SNL). There was, however, a strong tree-specific effect on SNL (P<10^–9, R ² = 0.75). Needle starch content varied markedly with season (0–25% of total dry mass). This, unless accounted for, would cause erroneous estimates of nutrient concentrations, and of rates of needle respiration, within and between treatments. There was considerable seasonal variation in needle respiration, both in terms of maintenance respiration and temperature dependence (Q₁₀). Q₁₀ had its highest value (2.8) during winter and its lowest (2.0) in the middle of summer. In early autumn (August, September), respiration rate and needle [N] were significantly related (C: P = 0.001, IL: P<0.0005). There was no significant difference in the slope between the two regression lines, but a difference in intercept. At the same needle [N], needles from IL-plots always had a lower respiration rate than needles from control plots. No obvious explanation for the observed difference in intercept was found, but some plausible assumptions are put forward and discussed. Received: 24 January 1997 / Accepted: 1 July 1997     

Succession-driven changes in soil respiration following fire in black spruce stands of interior Alaska

Boreal forests are highly susceptible to wildfire, and post-fire changes in soil temperature and moisture have the potential to transform large areas of the landscape from a net sink to a net source of carbon (C). Understanding the ecological controls that regulate these disturbance effects is critical to developing models of ecosystem response to changes in fire frequency and severity. This paper combines laboratory and field measurements along a chronosequence of burned black spruce stands into regression analyses and models that assess relationships between moss succession, soil microclimate, decomposition, and C source-sink dynamics. Results indicate that post-fire changes in temperature and substrate quality increased decomposition in humic materials by a factor of 3.0 to 4.0 in the first 7 years after fire. Bryophyte species exhibited a distinct successional pattern in the first five decades after fire that corresponded to decreased soil temperature and increased C accumulation in organic soils. Potential rates of C exchange in mosses were greatest in early successional species and declined as the stand matured. Residual sources of CO₂ (those not attributed to moss respiration or humic decomposition) increased as a function of stand age, reflecting increased contributions from roots as the stand recovered from disturbance. Together, the field measurements, laboratory experiments, and models provide strong evidence that interactions between moss and plant succession, soil temperature, and soil moisture largely regulate C source-sink dynamics from black spruce systems in the first century following fire disturbance.     

Effects of soil warming during spring on photosynthetic recovery in boreal Norway spruce stands

The effect of soil thawing and soil temperature on postwinter recovery of photosynthetic capacity was studied, during late spring and early summer, in Norway spruce stands in northern Sweden. Soil temperature was manipulated by means of buried heating cables. The warming treatment was applied to stands with low (natural) and high (fertilized) availability of nutrients. Soil thawing, expressed as water availability, was followed by means of sapflow in stems, and shoot water potentials. The recovery of photosynthetic capacity was assessed by measuring the rate of light-saturated photosynthesis (A_max), and maximum photochemical efficiency of photosystem II in detached shoots, and chlorophyll a fluorescence. Accumulation of starch reserves in the needles was followed as an independent indicator of photosynthetic performance in situ. Snowmelt and soil thawing occurred more than one month earlier in heated than in unheated plots. This was expressed both as sapflow and as differences in shoot water potential between treatments. During May, the rates of A_max were significantly higher on heated than on control plots. The effect of soil warming on A_max was, however, not reflected in chlorophyll fluorescence or needle starch content. The time course of the recovery of photosynthetic capacity was mainly controlled by mean air temperature and by the frequency of severe night frosts, and to a lesser extent by earlier soil thawing and higher soil temperatures.     

Clean rain promotes fine root growth and soil respiration in a Norway spruce forest

Soil acidification and N saturation are considered to affect the decomposition of soil organic matter as well as growth and mortality of fine roots in many forest soils. Here we report from a field experiment where ‘clean rain’ has been applied to the soil for about 10 years under a roofed plot of a 71‐year‐old Norway spruce plantation at Solling, Central Germany. Reduced amounts of protons (?78%), sulphate (?53%), ammonium (?86%), and nitrate (?49%) were sprayed on the soil surface of the clean rain plot between 1992 and 2001. In an adjacent roofed control plot, throughfall was collected and immediately re‐sprinkled below the roof construction without any chemical manipulation. One year before the clean rain treatment started, live and dead fine root masses (≤2 mm) were determined from undisturbed soil cores down to 40 cm mineral soil depth. Total live fine root mass was significantly lower in the clean rain plot than in the control plot. After the first sampling, the soil holes were refilled with quartz sand and repeatedly sampled in June 1992, June 1996, and October 2001. There were no differences in live and dead fine root masses between the plots in 1992 and 1996. In 2001, both live and dead fine root masses of the clean rain plot were about twice as high as in the control plot, indicating that fine root growth recovered in the mineral soil following 10 years of clean rain treatment. Moreover, the clean rain treatment significantly reduced the total N concentrations of live fine roots and 1‐year‐old needles. Our results suggest that the reduced N input promoted fine root growth to compensate N deficiency. Reduced Al concentration in soil solution may have contributed to the recovery of fine root growth, however, the toxicity of Al species is largely unknown. Mean annual soil respiration rate was 24% higher in the period from 2000 to 2001, indicating that the clean rain treatment increased respiration of roots and heterotrophic microorganisms within the rhizosphere. Laboratory incubation of samples from the organic horizon and the top mineral soil revealed no differences between the plots in the decay rate of soil organic matter. Our results suggest that strong reductions in atmospheric N deposition from about 30 to 10 kg N ha^?1 yr^?1 and decreasing acid stress can have beneficial effects on growth of fine roots in the mineral soil within a decade. We conclude that biological recovery under reduced atmospheric loads can affect the nutrient and carbon budget of spruce soils in the long run.     

Revitalization experiments in magnesium deficient Norway spruce stands in Austria

Amelioration of degraded forest ecosystems on acidic substrates showing the new type of forest decline is a major goal of forest management. A number of experiments show positive effects of Mg-application to systems suffering from Mg-deficiencies. The current paper compares experiments conducted in the Austrian part of the Bohemian Massif, where both effects on soil solution chemistry and effects on plant nutrition, vitality and growth were investigated. It turned out that any type of Mg-source is able to improve Mg-nutrition of trees; both a neutral salt like KIESERITE as well as alkaline reacting magnesite and dolomite derived materials. A positive reaction of vitality and growth could however only be induced with dolomitic lime or magnesite. Using mineral NPK fertilizers, even with high Mg-content, induced Mg-deficiencies and led to nutritional imbalances. In addition significant NO₃ ^--leaching occured. On the other hand an organic slow release fertilizer (BACTOSOL*) amended with magnesite derived fertilizers (BIOMAG**) led to balanced nutrition and a fast recovery of tree health status, as judged by crown transparency, vitality index and growth rates. In both cases, when either magnesite derived compounds or combinations with the organic slow release fertilizer were applied, NO₃ ^--leaching occured only during the first three years after fertilization. The leaching rates declined afterwards to values comparable to unfertilized plots, while Mg-content of the soil solution could be elevated compared to the CONTROL, showing the sustainability of proper fertilization.     

The ectomycorrhizal community structure in high mountain Norway spruce stands

The species composition of ectomycorrhizal (ECM) fungal communities can be strongly influenced by abiotic and biotic factors, which determine interactions among the species such as resource partitioning, disturbance, competition, or relationships with other organisms. To verify whether ectomycorrhization of the root tips and composition of the ECM community in Norway spruce vary according to site features and if ECM species peculiar to these environmental variables can be detected, ten comparable stands differing in bedrock pH and exposure were selected and studied. The results demonstrated that tips vitality and ectomycorrhization degree do not change significantly either on the same tree, or among trees growing in the same stand, whereas they differ greatly with bedrock pH and exposure, even if no spatial or temporal trend were found. ECM species composition revealed instead a significant connection with the two environmental features, with a few species significantly associated to them. The results suggest that pH/exposure patterns play a primary role in the adaptive selection of ECM species constituting the consortium.     

Changes in soil mineralogy and exchangeable cation pools in stands of Norway spruce planted on former pasture land

Chemical and mineralogical properties of the soils in 35- and 70-year-old stands of Norway spruce (Picea abies (L.) Karst), planted on former pasture and were studied at Asa Experimental Forest, southern Sweden. Remnant deciduous forests bordering the spruce stands were used as controls to assess possible tree-species-related effects on soil development. All soils are acid with little difference in soil pH between the spruce and deciduous stands. However, the saturation of the exchange complex with Mg is lower beneath spruce and the total exchangeable Mg pool in the upper meter of these soils is one third of the Mg store beneath the deciduous stands. Amphibole, biotite and chlorite are the major sources of Mg in the parent soil. The clay fraction of the topsoil beneath spruce has been depleted of all these easily weatherable ferromagnesian minerals. Apart from weathering-resistant primary silicates, the clay fraction consists almost exclusively of expandable, smectitic mixed-layer minerals, which are believed to be the products of advanced stages of biotite weathering. In contrast, vermiculite is the dominant secondary mineral in the A-horizon in the deciduous stands, and some chlorite has survived. Moreover, a greater depth of in situ weathering is indicated for the soil of the old spruce stands where biotite/vermiculite mixed-layers have formed in the C-horizon as products of early stages of biotite weathering. Thus, differences between the paired sites in soil solution chemistry are supported by the qualitative differences in soil mineralogy, and are believed to reflect divergent biotic and/or abiotic processes in the different stand types. This revised version was published online in August 2006 with corrections to the Cover Date.     

Seasonal variation of maximum photochemical efficiency in boreal Norway spruce stands

 The seasonal variation in maximum photochemical efficiency of photosystem II (F_v/F_m) and the relationship between F_v/F_m and climatic factors such as irradiance, frost-nights and daily mean temperature was studied in young Norway spruce trees for 4 years in northern Sweden. As a result of night frost, the F_v/F_m-ratio gradually decreased during the autumn. There was between-year variation in the pattern of F_v/F_m in fully exposed shoots during autumn and spring, largely as an effect of differing temperature conditions. During spring, there was a strong apparent relationship between daily mean temperature and F_v/F_m within the temperature range –3 to 12°C. The light regime to which the needles were exposed during winter affected F_v/F_m, and moderately shaded shoots from the bottom of the canopy generally had a higher F_v/F_m-ratio than fully exposed shoots from the top of the canopy. Received: 1 October 1997 / Accepted: 16 June 1998     

Belowground ectomycorrhizal communities in three Norway spruce stands with different degrees of decline in the Czech Republic

We investigated the ectomycorrhizal communities on the roots of adult trees and seedlings associated with three Norway spruce stands in the Czech Republic using morphological and molecular tools. The stands had different degrees of forest decline due to air pollution. The aims of the study were to obtain information about the belowground ectomycorrhizal community in a heavily damaged spruce forest and to assess whether missing ectomycorrhizal fungal partners could be one of the reasons for the observed lack of regeneration. The ectomycorrhizal species richness on the roots of adult trees was significantly lower in the heavily damaged site Mumlavska hora than in the other two, but less drastically so than that found in a fruitbody survey. The roots of adult trees and seedlings were fully mycorrhizal at this site although they were less species-rich. The most abundant ectomycorrhizal species on the root system of adult trees in all three forest stands was Tylospora fibrillosa, a member of the athelioid clade. It made up over 60% of root tips in Mumlavska hora and its proportion was at least twice that in the other two sites. This species was also an efficient colonizer of roots from seedlings, in particular, in the most damaged site. The different soil properties in this site may have caused the observed differences in the ectomycorrhizal species richness and composition. For example, cation exchange capacity and soil base saturation were lower and the soil more often saturated. However, the number of living trees and their defoliation status may well directly impact the ectomycorrhizal species composition by presumably affecting the amount of carbon delivered to the symbiotic fungal partners. Athelioids and thelephoroids are an important component of the belowground ectomycorrhizal community in most temperate and boreal forests, but the role they play might even be more crucial in stressed forest ecosystems. Based on our results, we suggest that factors other than missing ectomycorrhizal inoculum constrain natural regeneration in the heavily damaged site Mumlavska hora.     

Fractionated extraction and speciation of macronutrients from Scots pine and Norway spruce needles of different ages

Summary The relationship between proteins and the macroelements potassium, magnesium, calcium, sulphur and phosphorus was studied in homogenates of needles of different ages from Scots pine (Pinus sylvestris L.) and Norway spruce [Picea abies (L.) Karst.]. Complete extractions by acid digestion, protein extractions by a buffer-detergent and non-protein extractions by a buffer alone showed that most of the potassium and magnesium of the needles was soluble independent of the proteins. Only a minor part (50–60 ppm, dry weight) of the magnesium could be referred to the chlorophyll content of the needles. Both potassium and magnesium appeared independent of the protein gel filtration. This was also valid for the minor fraction of the total calcium content, which was extractable in a buffer. Heterogeneous calcium deposits are also suggested by extraction using a chelator (EGTA). Part of the buffer-soluble sulphur and phosphorus compounds occurred independently of solubilized proteins, and had complex distribution patterns after gel filtration, even outside the high molecular separation range. It is suggested that further analyses along the present lines, compared with conventional analyses on total extracts, may extend the usefulness of mineral nutrient analyses in plants.     

The use of different soil nitrogen sources by young Norway spruce plants

 Three-year-old Norway spruce trees were planted into a low-nitrogen mineral forest soil and supplied either with two different levels of mineral nitrogen (NH₄NO₃) or with a slow-release form of organic nitrogen (keratin). Supply of mineral nitrogen increased the concentrations of ammonium and nitrate in the soil solution and in CaCl₂-extracts of the rhizosphere and bulk soil. In the soil solution, in all treatments nitrate concentrations were higher than ammonium concentrations, while in the soil extracts ammonium concentrations were often higher than nitrate concentrations. After 7 months of growth, ¹⁵N labelled ammonium or nitrate was added to the soil. Plants were harvested 2 weeks later. Keratin supply to the soil did not affect growth and nitrogen accumulation of the trees. In contrast, supply of mineral nitrogen increased shoot growth and increased the ratio of above-ground to below-ground growth. The proportion of needle biomass to total above-ground biomass was not increased by mineral N supply. The atom-% ¹⁵N was higher in younger needles than in older needles, and in younger needles higher in plants supplied with ¹⁵N-nitrate than in plants supplied with ¹⁵N-ammonium. The present data show that young Norway spruce plants take up nitrate even under conditions of high plant internal N levels. Received: 1 April 1998 / Accepted: 9 October 1998     

Nitrogen mineralisation in podzol soils under boreal Scots pine and Norway spruce stands

N mineralisation was investigated in the mor humus layer of a podzol at a forested catchment area of Saarejärve Lake in Eastern Estonia. The investigated areas were pine (Rhodococcumunderstorey) and spruce (Vaccinium understorey) stands, which are permanent sample plots of an integrated monitoring network. The seasonal pattern of net N mineralisation was studied by incubating undisturbed cores of mor humus (0–8 cm) in buried polyethylene bags in situ. Samples were collected and incubated between July 1996 and April 1998. The period of incubation was approximately 1 month, except for wintertime when incubation lasted till thawing of ground (5 months). The amounts of mineral nitrogen formed during monthly incubations in vegetation period vary considerably (0.4–8.7 kg ha^–1). About 70% of the variation of net ammonification could be explained by environmental factors - temperature, initial moisture and pH. Ammonium was the dominant form of mineral nitrogen, which is typical for mor humus. The rate of nitrification was very low, and most of the annual net nitrification occurred during just one or two months (May–June, October) depending on site and year. Measured annual net N mineralisation was 29.2 kg ha^–1 for the spruce stand and 23.6 kg ha^–1 for the pine stand. These measures were found to be in good accordance with other N-fluxes in the ecosystem.     

Needle analyses and graphic vector analyses of Norway spruce and Scots pine stands

 The aims of this study were to test the nutritional condition of young stands of Norway spruce and Scots pine in southernmost Norway, where atmospheric inputs of anthropogenic nitrogen and strong acids, as well as deposition of magnesium by sea spray, are relatively high. There has been general concern about forest health in this region, specifically about nutrient deficiencies and nitrogen saturation caused by the atmospheric pollution. To expand our knowledge about these subjects, needle analyses and graphic vector analyses, as well as tree vigour index and height growth response to fertiliser application, were used as diagnostic tools. The overall conclusion is that phosphorous limitation is probably more frequent than expected in the coastal zone of southern Norway, especially in spruce on shallow soils. It is hypothesised that phosphorous limitation might be a more general problem or will arise as one in a 30 – 60 km coastal zone stretching from southernmost Sweden along the coast to the north and then westwards along the south coast of Norway. Nitrogen saturation in southernmost Norway might already occur on some sites with shallow soils. Needle analyses and graphic vector analyses can yield erroneous results if stress factors other than nutrient supply, e. g. severe drought, determine the growth rate of trees. In addition, the graphic vector analyses require that nutrients applied to the test plots are available to roots early in the growing season, and that needle nutrient concentrations in late autumn reflect those when the needles were formed. Received: 30 May 1995/Accepted: 27 December 1995     

Biomass, morphology and nutrient contents of fine roots in four Norway spruce stands

Fine root systems may respond to soil chemical conditions, but contrasting results have been obtained from field studies in non-manipulated forests with distinct soil chemical properties. We investigated biomass, necromass, live/dead ratios, morphology and nutrient concentrations of fine roots (<2 mm) in four mature Norway spruce (Picea abies [L.] Karst.) stands of south-east Germany, encompassing variations in soil chemical properties and climate. All stands were established on acidic soils (pH (CaCl₂) range 2.8–3.8 in the humus layer), two of the four stands had molar ratios in soil solution below 1 and one of the four stands had received a liming treatment 22 years before the study. Soil cores down to 40 cm mineral soil depth were taken in autumn and separated into four fractions: humus layer, 0–10 cm, 10–20 cm and 20–40 cm. We found no indications of negative effects of N availability on fine root properties despite large variations in inorganic N seepage fluxes (4–34 kg N ha⁻¹ yr⁻¹), suggesting that the variation in N deposition between 17 and 26 kg N ha⁻¹ yr⁻¹ does not affect the fine root system of Norway spruce. Fine root biomass was largest in the humus layer and increased with the amount of organic matter stored in the humus layer, indicating that the vertical pattern of fine roots is largely affected by the thickness of this horizon. Only two stands showed significant differences in fine root biomass of the mineral soil which can be explained by differences in soil chemical conditions. The stand with the lowest total biomass had the lowest Ca/Al ratio of 0.1 in seepage, however, Al, Ca, Mg and K concentrations of fine roots were not different among the stands. The Ca/Al ratio in seepage might be a less reliable stress parameter because another stand also had Ca/Al ratios in seepage far below the critical value of 1.0 without any signs of fine root damages. Large differences in the live/dead ratio were positively correlated with the Mn concentration of live fine roots from the mineral soil. This relationship was attributed to faster decay of dead fine roots because Mn is known as an essential element of lignin degrading enzymes. It is questionable if the live/dead ratio can be used as a vitality parameter of fine roots since both longevity of fine roots and decay of root litter may affect this parameter. Morphological properties were different in the humus layer of one stand that was limed in 1983, indicating that a single lime dose of 3–4 Mg ha⁻¹ has a long-lasting effect on fine root architecture of Norway spruce. Almost no differences were found in morphological properties in the mineral soil among the stands, but vertical patterns were apparently different. Two stands with high base saturation in the subsoil showed a vertical decrease in specific root length and specific root tip density whereas the other two stands showed an opposite pattern or no effect. Our results suggest that proliferation of fine roots increased with decreasing base saturation in the subsoil of Norway spruce stands.