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.     

Nitrogen cycling in two Norway spruce (Picea abies) ecosystems and effects of a (NH4)2SO4 addition

Nitrogen cycling in two Norway spruce (Picea abies (L.) Karst.) ecosystems in the ARINUS experimental watershed areas Schluchsee and Villingen (Black Forest, SW Germany) and initial effects of a (NH₄)₂SO₄ treatment are discussed. Although N reserves in the soils are similar and atmospheric N input is the same low to moderate level characteristic for many forested areas in SW Germany, N export by both seepage and streamwater differs considerably. At Villingen, deposited N is almost totally retained in the ecosystem, whereas at Schluchsee N export is the order of the input. This is explained by differences in forest management history. The Villingen site had been subject to excessive biomass export (e.g., litter raking) leading to unfavorable microbial transformations in the soil. In contrast, as a ‘relic’ of the former beech stand, the Schluchsee site is characterized by high biological activity in the soil with vigorous nitrification despite low pH values. Accordingly, the two ecosystems responded differently to the additional N input (150 kg NH₄ ⁺ -N ha^?1 as (NH₄)₂SO₄). Nitrification starting immediately in the Schluchsee soils led to continued Al mobilization and leaching of basic cations and NO₃ ^?. The availability of Mg, already deficient before treatment, further decreased due to Mg leaching and marked N uptake by the stand. In contrast, most of the added N in Villingen was immobilized in the soil. Hence, uptake by the stand and leaching of NO₃ ^? and cations was correspondingly lower than at Schluchsee. The results emphasize the problems associated with the definition of generally applicable values for ‘critical loads’ of N deposition.     

Aluminium-induced callose formation in root tips of Norway spruce (Picea abies (L.) Karst.)

The objective of this study was to examine whether aluminium (Al) induces callose formation in roots of Norway spruce as it does in soybean. Spruce seedlings were grown in Al-free nutrient solution under controlled conditions in a growth chamber at pH 3.8. After 21 days 170 μM Al was added or not (controls) to the complete nutrient solution (molar Ca/Al ratio: 0.75). Callose could be detected in outer root-tip cells of Al-treated plants within 3 h, using fluorescence microscopy after staining with aniline blue. Prolonged Al treatment up to 24 h increased both the density of the callose deposits and the number of affected cell layers. Control plants showed no comparable callose deposits. Ultrastructural examinations showed cell-wall appositions in Al-treated root cells but not in controls. The possible implications of Al-induced callose formation for nutrient and water uptake by roots are discussed.     

On the causes of decline of Norway spruce (Picea abies Karst.) in Central Europe

Abstract. A careful study of the etiology and symptoms of the decline phenomena in stands of silver fir ( Abies alba Mill.), Scots pine ( Pinus sylv L.), European beech ( Fagus silv. L.) and Norway spruce ( Picea abies Karst.) in southern Germany leads to the conclusion that all these diseases, although exhibiting some common features (e.g. premature senescence and shedding of leaves, formation of transparent crowns), vary considerably between species and, within one particular species, between forest regions. It therefore seems plausible to assume, as a first approach, that we have to deal with different types of disease or decline, and consequently also with varying sets of causes or stress factors. This approach can be demonstrated best by reviewing the present knowledge of diseases in Norway spruce.     

Ozone and drought stress — Interactive effects on the growth and physiology of Norway spruce (Picea abies (L.) Karst.)

Young trees of Norway spruce (Picea abies (L.) Karst.) were grown in 120 litre pots under two different ozone levels in open-top chambers for three seasons, 1992–1994. The ozone treatments were charcoal filtered air (CF, average 24 h seasonal mean 6.5 ppb) and non-filtered air with extra ozone aiming to track 1.5 times ambient (NF+, average seasonal mean 34 ppb). In addition, half of the spruce trees in Aug – Sep each season recieved a drought period of between five and seven weeks. The remaining half were kept well-watered. The soil water content, the needle water potentials, and the gas exchange as well as the chamber micro climate were measured before, during and after the drought period. Furthermore, the growth of the trees was measured as biomass increase. During the 1993 drought period, where the trees experienced a moderate drought stress, the trees grown in NF+ consumed soil water faster and showed a higher needle conductance compared to CF. However, no negative effects were found on needle water potential or growth. During the more severe 1994 drought stress period we did not find any differences between the two ozone treatments in soil water consumption, needle conductance or needle water potential. There was a significant negative effect of the high ozone treatment on tree biomass of the well-watered trees. Total plant biomass was reduced 18 % and stem biomass was reduced as much as 28 %. The negative effect of ozone on tree biomass was much smaller for the droughted trees.     

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.     

The influence of stand density and canopy position on sulfur content in needles of Norway spruce (Picea abies L. Karst.)

Analysis of total S in tree foliage has become widely accepted as a means of assessing S pollution of forests when air quality data are lacking. An investigation of total S content of mixed needle samples of different age and canopy height from two stands of young Norway spruce with different densities showed that S-content exceeded the maximum natural S-content (i.e. in the absence of SO₂-pollution) for spruce. Significantly higher average needle S contents were encountered with decreasing stand density. Changed aerodynamic roughness and physiological status of thinned trees indicated combined action of atmospheric and tree physiological variables on S deposition and uptake in forest ecosystems. A marked increase in S content from 1 yr old to older needles and from the lower to the upper canopy was observed.     

The effect of lime on microbial activity in spruce (Picea abies L.) forests

Summary The use of lime increased heat output and decreased the C:N ratio (global indicators of biological activity) in the organic horizons of a spruce forest (Picea abies L.) soil. These changes were still present after 18 years. During the same period, the muramic acid content increased slightly, while the concentration of both ergosterol and glucosamine decreases. The ratios of ergosterol or glucosamine to muramic acid decreased significantly after 3 years in the plots that had been irrigated and limed, and after 8 years in the limed non-irrigated plots.     

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.     

Amelioration of Magnesium Deficiency in a Norway Spruce Stand (Picea abies) with Calcined Magnesite

Norway spruce has shown needle yellowing and defoliation symptoms on manysites of the Bohemian Forest in north central Austria since theearly 1980s. A forest amelioration experiment was set up toevaluate the response of spruce to magnesite based fertilizerswith different solubilities. Response variables were a crownvigor index, diameter growth, nutrient contents in needles,soil solution chemistry, and soil chemistry. Soil solution atfertilized plots showed an immediate response in base cationconcentrations, nitrate concentrations and pH-values.Fertilization increased the pool of exchangeable cations in theupper 15 cm of the soil and reduced levels of exchangeable Al. Themagnesium and calcium content of the needles of dominant trees wasgreatly improved. Fertilization has increased the mineralizationrate of soil organic matter. Diameter growth, as a measure of treevigor, showed a pronounced positive response with a time delay of5 years. Our results demonstratethat magnesite based fertilizers are a viable option to improvethe nutritional status of spruce forests on granitic bedrock.     

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.     

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.     

Nutrient response to diagnostic fertilization of Norway spruce Picea aries (L.) karst plantations in Western Quebec,Canada

Maple forests in southern Quebec are suffering in the 1980s from a serious decline thought to be associated, in part, with acid deposition effects on their nutrient status. Research was undertaken to determine if conifers were subject to similar phenomena. The nutrient status of three Norway spruce plantations in southwestern Quebec were studied through foliar and soil analyses. In all locations low N and K and marginal Mg foliar concentrations were identified. Diagnostic fertilization resulted in improved K and P and in some cases Mg and N levels. The foliage of the treated trees has an improved dark green color as compared with the controls. From this research it is evident that nutrient deficiencies of N, K and Mg, in particular, are showing up in spruce stands in the areas where air pollution and acid rain are relatively severe. Proper diagnostic fertilization helped to correct the nutrient disturbances.     

Foliar symptoms,ethylene biosynthesis and water use of young Norway spruce (Picea abies (L.) Karst.) exposed to drought and ozone

Four-year old Norway spruce (Picea abies (L.) Karst.) trees were pretreated at low and high water supply, and then placed into a growth chamber containing four compartments so that two levels of ozone exposure, 0.02 and 0.4 Μl l^?1, could be replicated. They were exposed to ozone and drought stress for 59 days, and water use was determined by periodic weighing. Small effects of ozone treatment were detected on new shoot dry weight, and water use by trees receiving the high ozone treatment appeared higher. Both visual symptoms and ethylene biosynthesis showed that drought stress reduced damage to trees exposed to high ozone. Ethylene emission and 1-aminocyclopropane-1-carboxylic acid (ACC) levels increased after 18 days of exposure to 0.4 Μl l^?1 ozone, while visual symptoms were seen at 30 days. After 59 days of exposure to the combined stresses, ethylene and ACC levels were lower, but showed an ozone x water interaction. Most ethylene and ACC were produced by wet trees at high ozone concentration, but dry trees also had high ethylene and ACC levels at low ozone. Levels of needle malonyl-ACC (MACC) were not significantly affected by treatment, and did not change with time, but root MACC levels, which were twice needle levels, were high in wet trees at high ozone concentration, but also high in dry trees at low ozone concentration. These results suggest that drought stress occurring during ozone exposure could be expected to reduce damage to young Norway spruce, and that this damage may be related to ethylene biosynthesis.     

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.     

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.     

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.     

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.     

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.     

Uptake and assimilation of atmospheric NO2 — N by spruce needles (Picea abies): A field study

NO₂ enters spruce needles by gas exchange through the slomata. Nitrate formed from NO₂ is reduced in the cytosol by nitrate reductase (NR), the rate limiting enzyme of the nitrogen assimilatory pathway. A linear relationship was found between the nitrate reductase activity (NRA), NO₂ concentration and the amount of N incorporated into amino acids and proteins, so that NRA was suggested as an estimate of NO₂-uptake. In the present field study, 50 spruce trees (Picea abies) have been selected, which grow in a natural habitat in a NO₂ concentration gradient in a forest crossed by a highway which is a major NO source. At part of the sites, the microclimatic conditions have been recorded, so that common models of local gas exchange of the needles could be used to estimate stomatal uptake of NO₂. NRA was investigated as a function of radiation and stomatal uptake on the day before needle sampling. Close to the highway NRA was permanently elevated with a maximum in summer. As with the laboratory results, a linear relationship between stomatal uptake and NRA was found. Total N — content of current year shoots was not affected by the additional N-source provided by airborne NO₂. The present study shows that the gas exchange models are consistent with the physiological reactions of spruce needles on a local level and therefore contribute to the validation of calculations of NO₂ dry deposition to spruce forests.