Methane and nitrous oxide fluxes of soils in pure and mixed stands of European beech and Norway spruce

Tree species can affect the sink and source strength of soils for atmospheric methane and nitrous oxide. Here we report soil methane (CH₄) and nitrous oxide (N₂O) fluxes of adjacent pure and mixed stands of beech and spruce at Solling, Germany. Mean CH₄ uptake rates ranged between 18 and 48 μg C m^?2 hour^?1 during 2.5 years and were about twice as great in both mixed and the pure beech stand as in the pure spruce stand. CH₄ uptake was negatively correlated with the dry mass of the O horizon, suggesting that this diminishes the transport of atmospheric CH₄ into the mineral soil. Mean N₂O emission was rather small, ranging between 6 and 16 μg N m^?2 hour^?1 in all stands. Forest type had a significant effect on N₂O emission only in one mixed stand during the growing season. We removed the O horizon in additional plots to study its effect on gas fluxes over 1.5 years, but N₂O emissions were not altered by this treatment. Surprisingly, CH₄ uptake decreased in both mixed and the pure beech stands following the removal of the O horizon. The decrease in CH₄ uptake coincided with an increase in the soil moisture content of the mineral soil. Hence, O horizons may maintain the gas diffusivity within the mineral soil by storing water which cannot penetrate into the mineral soil after rainfall. Our results indicate that conversion of beech forests to beech–spruce and pure spruce forests could decrease soil CH₄ uptake, while the long‐term effect on N₂O emissions is expected to be rather small.     

Spatial variability of photosynthetically active radiation in European beech and Norway spruce

The vertical and horizontal variability of solar radiation within a mature European beech (Fagus sylvatica L.)-Norway spruce (Picea abies [L.] Karst) mixed stand in Southern Germany is investigated. A large dataset with more than one million spectral measurements of photon fluence rates at six vertical levels within the stand is analyzed with respect to tree species, meteorological sky conditions, and the influence of solar elevation angle on canopy penetration. Irradiance probability density functions of the photosynthetically active waveband are used to describe the three-dimensional radiation field. For a quantification of umbra, penumbra, and sunfleck frequencies, in-canopy fractions of photon fluence rates within the photosynthetically active waveband are investigated. Different phenological stages of beech and their effects on the in-canopy light climate are compared. The results show that during overcast conditions (OVC) fractions of photosynthetically active radiation (PAR) are higher at all canopy levels than during clear sky (CS) conditions due to their exclusively diffuse character. The lowest median PAR level of less than 1% of above-canopy PAR can be observed in the shade crown of beech and at ground level. More PAR can penetrate the canopy at a higher solar elevation under CS conditions. This effect is more pronounced for spruce than for beech due to the conical crown shape of the conifers that allows photons from higher angles to enter the gaps inbetween trees in contrast to the more homogeneously closed beech canopy. Solar elevation is not an important factor at uniformly overcast conditions. Differences in the vertical distribution of umbra and penumbra can be detected when comparing species or different sky conditions. The frequency of sunflecks differs more by species and by the vertical position within the canopy than by sky condition.     

Methane oxidation in boreal forest soils: kinetics and sensitivity to pH and ammonium

In soil incubation experiments we examined if there are differences in the kinetic parameters of atmospheric methane (CH₄) oxidation in soils of upland forests and forested peatlands. All soils showed net uptake of atmospheric CH₄. One of the upland forests included also managed (clear-cut with or without previous liming or N-fertilization) study plots. The CH₄ oxidation in the forested peat soil had a higher K_m (510 μl l⁻¹) and V_max (6.2 nmol CH₄ cm⁻³ h⁻¹) than the upland forest soils (K_m from 5 to 18 μl l⁻¹ and V_max from 0.15 to 1.7 nmol CH₄ cm⁻³ h⁻¹). The forest managements did not affect the K_m-values. At atmospheric CH₄ concentration, the upland forest soils had a higher CH₄ oxidation activity than the forested peat soil; at high CH₄ concentrations the reverse was true. Most of the soils oxidised CH₄ in the studied pH range from 3 to 7.5. The pH optimum for CH₄ oxidation varied from 4 to 7.5. Some of the soils had a pH optimum for CH₄ oxidation that was above their natural pH. The CH₄ oxidation in the upland forest soils and in the peat soil did not differ in their sensitivities to (NH₄)₂SO₄ or K₂SO₄ (used as a non-ammonium salt control). Inhibition of CH₄ oxidation by (NH₄)₂SO₄ resulted mainly from a general salt effect (osmotic stress) though NH₄⁺ did have some additional inhibitory properties. Both salts were better inhibitors of CH₄ oxidation than respiration. The differences in the CH₄ oxidation kinetics in the forested peat soil and in the upland forest soils reveal that there are differences in the physiologies of the CH₄ oxidisers in these soils.     

Vertical gradients of potential enzyme activities in soil profiles of European beech,Norway spruce and Scots pine dominated forest sites

Management of forest sites has the potential to modulate soil organic matter decomposition by changing the catalytic properties of soil microorganisms within a soil profile. In this study we examined the impact of forest management intensity and soil physico-chemical properties on the variation of enzyme activities (β-glucosidase, β-xylosidase, α-glucosidase, phenol oxidase, N-acetyl-glucosaminidase, l-leucine aminopeptidase, phosphatase) in the topsoil and two subsoil horizons in three German regions (Schorfheide-Chorin, Hainich-Dün, Schwäbische Alb). The sandy soils in the Schorfheide-Chorin (SCH) showed lower ratios of the activity of carbon (C) acquiring enzymes (β-glucosidase) relative to nitrogen (N) acquiring enzymes (N-acetyl-glucosaminidase + l-leucine aminopeptidase), and activity of C acquiring enzymes relative to phosphorous (P) acquiring enzymes (phosphatase) than the finer textured soils in the Hainich-Dün (HAI) and Schwäbische Alb (ALB), indicating a shift in investment to N and P acquisition in the SCH. All enzyme activities, except phenol oxidase activity, decreased in deeper soil horizons as concentrations of organic C and total N did, while the decrease was much stronger from the topsoil to the first subsoil horizon than from the first subsoil to the second subsoil horizon. In contrast, phenol oxidase activity showed no significant decrease towards deeper soil horizons. Additionally, enzyme activities responsible for the degradation of more recalcitrant C relative to labile C compounds increased in the two subsoil horizons. Subsoil horizons in all regions also indicate a shift to higher N acquisition, while the strength of the shift depended on the soil type. Further, our results clearly showed that soil properties explained most of the total variance of enzyme activities in all soil horizons followed by study region, while forest management intensity had no significant impact on enzyme activities. Among all included soil properties, the clay content was the variable that explained the highest proportion of variance in enzyme activities with higher enzyme activities in clay rich soils. Our results highlight the need for large scale studies including different regions and their environmental conditions in order to derive general conclusions on which factors (anthropogenic or environmental) are most influential on enzyme activities in the whole soil profile in the long term at the regional scale.     

A comparison of water soluble organic substances in acid soils under beech and spruce in NE-Bavaria

Water soluble organic substances extracted from the L, O_f, O_h, A, and B horizons of acid soils under Beech (Fagus sylvatica) and Spruce (Picea abies) were fractionated on Sephadex G-25 (medium). Infrared, ¹H NMR and ¹³C NMR spectra of the fractionated components were obtained and comparisons were made among fractions, among horizons, and between forest types. Usually, the spectra indicated that carbohydrates dominated the water soluble fractions. Substances originated from spruce exhibited more undissociated carboxylic acid character than the beech derived material. Despite many similarities, principal differences in the carbohydrate components were found among fractions, among horizons, and between vegetation types.     

Microbial biomass and C and N transformations in forest floors under European beech, sessile oak, Norway spruce and Douglas-fir at four temperate forest sites

The purpose of this research was to compare soil chemistry, microbially mediated carbon (C) and nitrogen (N) transformations and microbial biomass in forest floors under European beech (Fagus sylvatica L.), sessile oak (Quercus petraea (Mattuschka) Lieblein), Norway spruce (Picea abies (L.) Karst) and Douglas-fir (Pseudotsuga menziesii (Mirbel) Franco) at four study sites. We measured soil chemical characteristics, net N mineralization, potential and relative nitrification, basal respiration, microbial and metabolic quotient and microbial biomass C and N under monoculture stands at all sites (one mixed stand). Tree species affected soil chemistry, microbial activities and biomass, but these effects varied between sites. Our results indicated that the effect of tree species on net N mineralization was likely to be mediated through their effect on soil microbial biomass, reflecting their influence on organic matter content and carbon availability. Differences in potential nitrification and relative nitrification might be related to the presence of ground vegetation through its influence on soil NH₄ and labile C availability. Our findings highlight the need to study the effects of tree species on microbial activities at several sites to elucidate complex N cycle interactions between tree species, ground vegetation, soil characteristics and microbial processes.     

Methane oxidation in temperate soils: effects of inorganic N

Additions of inorganic nitrogen (N) to an oak soil with significant potential for methane (CH₄) oxidation resulted in differential reduction in CH₄ oxidation capacity depending on N species added. Nitrate, rather than nitrite or ammonium, proved to be the strongest inhibitor of CH₄ oxidation in oak soil. Both high (CH₄ at 10 μl l⁻¹) and low (CH₄ at 5 ml l⁻¹) affinity CH₄ oxidation in oak soil was completely inhibited at a nitrate concentration similar to that present in an alder soil from the same experimental site. The alder soil showed no capacity for low affinity CH₄ oxidation. A ‘low nitrate’ forest soil (oak) showed high affinity, low capacity CH₄ oxidation upto around 1 ml l⁻¹ CH₄, above which both high and low affinity CH₄ oxidation became apparent following a lag phase, indicating either an induced high affinity uptake mechanism or the existence of distinct low affinity and high affinity methanotroph populations. High affinity CH₄ oxidation became saturated at CH₄ concentrations >500 μl l⁻¹, while low affinity CH₄ oxidation became saturated at ∼30 ml l⁻¹ CH₄. In a ‘high nitrate’ forest soil (alder), CH₄ oxidation appeared to be due to high affinity CH₄ oxidation only and became undetectable at CH₄ concentrations >5 ml l⁻¹.     

Dynamics of dissolved organic nitrogen and carbon in a Central European Norway spruce ecosystem

Dissolved organic nitrogen and carbon (DOC) are significant in the C and N cycle in terrestrial ecosystems. Little is known about their dynamics in the field and the factors regulating their concentrations and fluxes. We followed the fluxes and concentrations of the two in a Norway spruce (Picea abies (L.) Karst.) forest ecosystem in Germany from 1995 to 1997 by sampling at fortnightly intervals. Bulk precipitation, throughfall, forest floor percolates from different horizons and soil solutions from different depths were analysed for major ions, dissolved organic N and DOC. The largest fluxes and concentrations were observed in percolates of the Oi layer, which contain amino N and amino sugar N as the major components. The average ratio of dissolved organic C to N in forest floor percolates corresponded to the C/N ratio of the solid phase. Concentrations and fluxes were highly dynamic with time and decreased with depth. The largest fluxes in forest floor percolates occurred when the snow melted. The concentrations and fluxes of dissolved organic N were significantly correlated with DOC, but the correlation was weak, indicating different mechanisms of release and consumption. The dynamics of dissolved organic N and DOC in forest floor percolates were not explained by pH and ionic strength of the soil solution nor by the water flux, despite large variations in these. Furthermore, the release of these fractions from the forest floor was not related to the quality and amount of throughfall. Concentrations of dissolved organic N in forest floor percolates increased with soil temperature, while temperature effects on DOC were less pronounced, but their fluxes from the forest floor were not correlated with temperature. In the growing season concentrations of both dissolved organic N and C in forest floor percolates decreased with increasing intensity of throughfall. Thus, the average throughfall intensity was more important than the amount of percolate in regulating their concentrations in forest floor percolates. Our data emphasize the role of dissolved organic N and DOC in the N and C cycle of forest ecosystems.     

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.     

Direct estimates of C:N ratios of ectomycorrhizal mycelia collected from Norway spruce forest soils

Direct estimates of C:N ratios of ectomycorrhizal (EM) mycelia growing in situ in forest soils have been obtained for the first time. The mycelial samples were collected from sand-filled mesh bags that were buried in the soil and incubated for 12-18 months in two Norway spruce forests in southern Sweden. At harvest the mesh bags were heavily colonized and the mycelia were extracted from the sand with water. The collected mycelia had earlier been identified as belonging to EM fungi based on their C isotopic composition. The mean value of the C:N ratio for mycelia was 20.2±0.8 (n=25). EM mycelia collected at different soil depths (5, 15 and 30 cm) had similar C:N ratios. C:N ratios of microbial biomass obtained by fumigation-extraction of similar soils have usually been lower (6-13) so possible differences in the extraction efficiency of C and N from bacteria and fungi are discussed.     

Carbon dioxide emissions of soils under pure and mixed stands of beech and spruce, affected by decomposing foliage litter mixtures

Soil respiration is the largest terrestrial source of CO₂ to the atmosphere. In forests, roughly half of the soil respiration is autotrophic (mainly root respiration) while the remainder is heterotrophic, originating from decomposition of soil organic matter. Decomposition is an important process for cycling of nutrients in forest ecosystems. Hence, tree species induced changes may have a great impact on atmospheric CO₂ concentrations. Since studies on the combined effects of beech-spruce mixtures are very rare, we firstly measured CO₂ emission rates in three adjacent stands of pure spruce (Picea abies), mixed spruce-beech and pure beech (Fagus sylvatica) on three base-rich sites (Flysch) and three base-poor sites (Molasse; yielding a total of 18 stands) during two summer periods using the closed chamber method. CO₂ emissions were higher on the well-aerated sandy soils on Molasse than on the clayey soils on Flysch, characterized by frequent water logging. Mean CO₂ effluxes increased from spruce (41) over the mixed (55) to the beech (59) stands on Molasse, while tree species effects were lower on Flysch (30-35, mixed > beech = spruce; all data in mg CO₂-C m⁻² h⁻¹). Secondly, we studied decomposition after fourfold litter manipulations at the 6 mixed species stands: the O_i - and O_e horizons were removed and replaced by additions of beech -, spruce - and mixed litter of the adjacent pure stands of known chemical quality and one zero addition (blank) in open rings (20 cm inner diameter), which were covered with meshes to exclude fresh litter fall. Mass loss within two years amounted to 61-68% on Flysch and 36-44% on Molasse, indicating non-additive mixed species effects (mixed litter showed highest mass loss). However, base cation release showed a linear response, increasing from the spruce - over the mixed - to the beech litter. The differences in N release (immobilization) resulted in a characteristic converging trend in C/N ratios for all litter compositions on both bedrocks during decomposition. In the summers 2006 and 2007 we measured CO₂ efflux from these manipulated areas (a closed chamber fits exactly over such a ring) as field indicator of the microbial activity. Net fluxes (subtracting the so-called blank values) are considered an indicator of litter induced changes only and increased on both bedrocks from the spruce - over the mixed - to the beech litter. According to these measurements, decomposing litter contributed between 22-32% (Flysch) and 11-28% (Molasse) to total soil respiration, strengthening its role within the global carbon cycle.     

Decomposition of beech leaves (Fagus sylvatica) and spruce needles (Picea abies) in pure and mixed stands of beech and spruce

The decomposition of spruce needles and beech leaves was investigated in a 30- and 120-yr-old beech, spruce and mixed (beech/spruce) forest using 1 mm mesh litterbags. The mass loss, content of C, N and water and microbial biomass, basal respiration and specific respiration of the litter materials were analyzed after exposure for 1.5, 3, 6, 9, 12, 18 and 24 months in the field. Decomposition of both types of litter was faster in beech than in spruce stands and after 24 months loss of C from litter materials was at a maximum in beech stands (>60%) and considerably less in the spruce and mixed stands (ca. 40%). Generally, spruce needles decomposed more rapidly than beech leaves, but the faster decay was not associated with higher N concentrations. Rather, N was accumulated more rapidly in beech leaves. Concomitantly, in beech stands microbial biomass of beech leaves exceeded that of spruce needles indicating that beech leaves consist of more favorable resources for microorganisms than spruce needles. Differences in decomposition between beech leaves and spruce needles were most pronounced in beech stands, intermediate in mixed stands and least pronounced in spruce stands. Decomposition, N content and microbial biomass in litter materials exposed in the 120-yr-old stand consistently exceeded that in the 30-yr-old stand indicating adverse conditions for litter decay in regrowing stands. Generally, mixed stands ranked intermediate between spruce and beech monocultures for most of the variables measured indicating that the adverse conditions for litter decay and microorganisms in spruce forest are effectively counteracted by admixture of beech to spruce monocultures. It is concluded that the accumulation of litter materials in spruce forests is not due to the recalcitrance of spruce needles to decay. Rather, adverse environmental conditions such as high polyphenol contents in the litter layer of spruce stands retard decomposition processes; spruce needles appear to be more sensitive to this retardation than beech leaves.     

Tree roots in canopy soils of old European beech trees—An ecological reassessment of a forgotten phenomenon

The formation of adventitious roots in humus accumulations in tree canopies is widely acknowledged from tropical and temperate rainforests, while the occurrence of those canopy roots in temperate tree species under mesic climates has been largely disregarded for ca. 100 years. Moreover, almost nothing is yet known of the ecological growth conditions or the structure or morphology of such canopy root systems. This study reports on the occurrence of tree fine roots in crown humus pockets of old European beech (Fagus sylvatica L.) trees. The aim was to compare these canopy roots with the fine roots in the terrestrial organic layer soil in terms of fine root biomass density, root morphological traits, ectomycorrhizal colonisation and chemical composition of the root tissue, and to relate these root traits to the chemical properties of the respective soils. Fine root biomass density in crown humus pockets was ca. 7 times higher than in the terrestrial organic layer, even though soil chemical properties of both rooting media were similar. Fine roots in the canopy differed from terrestrial fine roots by lower specific root tip abundance, specific root length, and specific root surface area, all of which points to a longer lifespan of the fine roots in the canopy. Moreover, canopy roots revealed a lower percentage of root tips colonised by ectomycorrhizal fungi than terrestrial roots (87% vs. 93%). Chemical composition of the root tissue in canopy and terrestrial soils was similar for most elements, but canopy roots showed lower P, Fe, and Al concentrations and a higher N/P ratio than terrestrial roots. Root P concentrations of both canopy and terrestrial fine roots were closely related to soil P concentration, but not to soil C/P or N/P ratios. On the other hand, tissue N of canopy roots, but not of terrestrial roots, revealed a clear dependence on soil N and C/N values, suggesting a more limited N availability in the canopy soil compared to the terrestrial organic layer. However, the overall small differences in soil chemical properties between canopy and terrestrial organic layer soil cannot explain the markedly higher volumetric root density in the crown humus and the differences in ecomorphological traits between canopy and terrestrial soil. Instead, it is speculated that these differences are more likely a result of temporarily high water availability in crown humus pockets due to high water flow along the surface of branches to the central crown parts of the beech trees.     

Phenology,photosynthesis, and phosphorus in European beech (Fagus sylvatica L.) in two forest soils with contrasting P contents

Phosphorus (P) is often a limiting macronutrient in temperate forests, but knowledge on the phenological and physiological responses of beech (Fagus sylvatica L.) to P deficiency is scarce. In this study, young beech trees were excavated with intact soil cores from two German forests, Unterlüss (LUE) with low soil P and Bad Brückenau (BBR) with high soil P concentrations. The trees were transferred to identical climatic conditions. In the subsequent growth phase phenological stages during bud burst and leaf unfolding were recorded; biomass production and total P concentrations in different tissues were measured. Seasonal fluctuations in photosynthesis and of soluble P in wood and bark exudates were determined. BBR beeches grew faster and produced more and larger leaves than the LUE beeches. Leaf extension and unfolding were delayed in LUE compared with BBR beeches, but not the time point of bud break. All plant tissues of BBR trees contained higher total P concentrations than those of LUE trees. Strong seasonal fluctuations for P in exudates of beech transport tissues, wood and bark, indicated higher P supply in BBR than in LUE plants, especially at the beginning of the growth phase until leaf maturity. Photosynthetic activity of LUE beeches was lower than that of BBR beeches due to stomatal limitations as the result of anatomically smaller stomatal pore widths, but not as the result of acute biochemical limitation of photosynthesis. Our results suggest that developmental retardation and lower photosynthesis under low P availability may be adaptation mechanisms that adjust the acquisition and recycling of P resources to seasonal growth demand.     

Evidence of plaggen soils in SW Norway

This study aimed at clarifying whether a notable group of soils of the Jæren region, SW Norway, with deep humus‐rich top soils support a man‐made genesis. Four sites were investigated. The soils are characterized by thick top soils of 45, 70, 80, and 90 cm, which are enriched in soil organic matter and often also in artifacts, like fragments of potter's clay, indicating an anthropogenic origin. Soil pH ranges from 5.4 to 6.2 (H₂O) and 4.4 to 5.3 (CaCl₂), respectively. Soil organic C (SOC) contents range from 6.4 to 51.6 g kg^?1 and N contents vary between 0 and 2.9 g kg^?1. Increased P contents of up to 2,924.3 mg kg^?1 total P (P_t) and 1,166.4 mg kg^?1 citric acid‐soluble phosphorus (P_c) in the humus‐rich top soils support the assumption of an anthropogenic influence. Although many characteristics indicate an anthropogenic genesis, one soil lacks the required depth of 50 cm of a plaggen horizon and cannot be classified as Plaggic Anthrosol (WRB) and Plagganthrept (US Soil Taxonomy). As the requirement is 40 cm in the German system, all soils can be classified as Plaggenesch. The formation of these soils is related to human activity aiming at increasing soil fertility and overcoming the need of bedding material, the basic aims of the plaggen management in Europe. Highest P contents ever found for this kind of soils and references from the literature indicate that the formation of the soils in Norway started at Viking time, hence, being older than most other Plaggic Anthrosols.     

Nitrogen transformations and pools in N-saturated mountain spruce forest soils

Nitrogen leaching persists in mountain forests of Europe even in the presence of decreasing N depositions. We have hypothesized that this leaching is linked to soil N transformations occurring over the whole year, even at 0°C temperatures. The aims were to estimate (1) the effect of temperature on N transformations and (2) N pools and fluxes. The study sites are situated in the Bohemian Forest (Czech Republic). Litter, humus, and 0–10-cm mineral layers were sampled in early spring, and the effect of temperature on net nitrification, net ammonification, and microbial N immobilization were measured in a short-term incubation experiment without substrate addition. Nitrogen pools were calculated from the concentrations of N forms in the soil and soil pool weights, while daily N fluxes were calculated from daily net rates of processes and soil pool weights. Relationships between temperature and net nitrification, net ammonification, and microbial N immobilization did not follow the Arrhenius type equation; all processes were active close to 0°C, indicating that microbial N transformations occur over the whole year. Microbial N immobilization rate was generally greater than N mineralization rate. The microbial N pool was significantly larger than mineral N pools. Organic layers containing tens of grams of available N per square meter contributed more than 70% to the available N in the soil profile. Daily N fluxes were related to N pools. On average, N fluxes represented daily mineral and microbial N pool changes of 1.14 and 1.95%, respectively. The effect of microbial composition on the C/N ratio of microbial biomass and respiration is discussed.     

Correlations between element concentrations in spruce foliage and forest soils

Abstract

The relationships between foliage element concentrations in red spruce and soil chemical properties were studied to determine if standard soil measurements of individual elements in soils were well correlated with the concentrations of these elements in foliage. Significant positive correlations between O horizon and foliage concentrations existed only for K, Mn and P. Significant negative correlations between the concentrations of the major divalent cations (i.e. Ca, Mg, Mn) and K in the foliage were found suggesting a possible antagonism between the mono‐ and divalent cations for uptake from the soil. Trees with the highest foliage concentrations of Ca also were determined to be growing on soils which were producing the best growth rates. Foliage concentrations of P, and to a lesser extent K and Mg, were below values considered to be adequate for optimum growth in red spruce.     

Environmental effects on soil NO concentrations and root N uptake in beech and spruce forests

This study aimed to investigate the shifts in net nitrogen (N) uptake and N compounds of fine roots over the vegetation period (i.e., spring, summer, autumn) and correlate this with NO concentration in the soil. Soil NO concentration was measured using gas lysimeters for collection and a chemiluminescence analyzer for quantification. Net N uptake by the roots was determined using the ¹⁵N enrichment technique. N pools were quantified using spectrophotometric techniques. Soil NO concentrations at beech and spruce forest sites were highest in spring (June), and lowest in winter (December). Total N of the roots was similar during the seasons and between the two years under study despite considerable variation of different N compounds. Net N uptake generally increased with higher N supply. Correlation analysis revealed a positive relationship between soil NO concentration and net N uptake only for spruce trees. This relationship seemed to be modulated by environmental factors and tree species.