Effects of soil pH and calcium on mycorrhizas of Picea abies

The effects of lime, increased soil pH and increased soil Ca concentration on the mycorrhizas of Norway spruce. [Picea abies (L.) Karst.] were studied independently of each other to elucidate the different mechanisms through which lime may influence mycorrhizas in acidic soil. In a field experiment (mature Norway spruce in podzol), lime was applied as CaCO₃; increased Ca concentration without an increase in pH was achieved with CaSO₄; and soil pH was increased without calcium by means of Na₂CO₃ and K₂CO₃ (Na+K treatment). Treatments were done in October, and mycorrhizas were counted from samples collected in the following June and September. All treatments increased the percentage of dead short root tips compared to controls in September, and Na+K already in June. Cenococcum geophilum Fr. increased in proportion in plots treated with Na+K.In a sand culture experiment, Norway spruce seedlings were grown from seed and inoculated with Cenococcum geophilum, or root inoculum, or left uninoculated. When mycorrhizas were beginning to form, CaCO₃ and CaSO₄ treatments were applied. Six weeks later, the percent of dead short root tips in both salt treatments was significantly increased from control, but formation of mycorrhizas was not inhibited by treatments.As all the treatments increased the proportion of dead short root tips, it is concluded that lime directly and adversely affected mycorrhizas of Norway spruce in sand culture and in mor humus. Both increased ionic strength and increased pH may be reasons for this rather than Ca²⁺ specifically.     

Effects of liming and boron fertilization on mycorrhizas of Picea abies

Effects of liming and B fertilization on Norway spruce [Picea abies (L.) Karst.] mycorrhizas were studied in factorial field experiments. The lime was applied twice, about 30 years and 12 years before sampling (2000 and 4000 kg ha^-1 dolomite). B was applied at the rate of 1.5 kg B ha^-1 two years before sampling.Boron fertilization doubled the number of root tips in the top 10 mm of the humus layer. The proportion of dead short root tips was increased from 10 % in control plots to 29 % in the limed plots. Numbers of dead root tips were increased when both lime and B were applied. The % of mycorrhizas with external mycelium was slightly increased and the % of Piloderma croceum Erikss. and Hjortst. was decreased by lime. In conclusion, adverse effects of lime on mycorrhizas were found, which were ameliorated by B fertilization, but lime-induced B deficiency alone was not the only reason for the effects of lime on root mortality.     

Abundance of two threatened woodpecker species in relation to the proportion of spruce plantations in native pine forests of western Norway

In a comparative study we investigated woodpecker abundance in forest landscapes with different proportion of native pine forest and spruce plantations in western Norway. In 100 circular study plots of 100ha each we recorded 38 white-backed –Dendrocopos leucotos, 22 grey-headed –Picus canus, 13 great spotted –Dendrocopos major, 6 green –Picus viridis, and 2 lesser spotted –Dendrocopos minor woodpeckers in the breeding season. The mean number of recorded woodpecker species peaked at 20–40% spruce plantations. The two most common species in the study, the white-backed and the grey-headed woodpeckers are both Red-listed species in Norway and among the rarest woodpeckers in Europe. The white-backed woodpecker preferred plots with higher than average proportions of standing dead trees and deciduous trees, and low proportions of spruce plantations in the plots. The grey-headed woodpecker preferred plots in the western (coastal) parts of the study area with presence of large aspen Populus tremula trees. Logistic regression models did not reveal any clear threshold values with respect to proportion of spruce plantations in plots, although both woodpecker species were extremely rare in plots with >60% spruce plantations. We recommend spruce plantations to be kept at moderate levels to ensure viable populations of woodpeckers in western Norway.     

Effects of liming and boron fertilization on boron uptake of Picea abies

The effects of liming on concentrations of boron and other elements in Norway spruce [Picea abies (L) Karst.] needles and in the mor humus layer were studied in long-term field experiments with and without B fertilizer on podzolic soils in Finland. Liming (2000+4000 kg ha^-1 last applied 12 years before sampling) decreased needle B concentrations in the four youngest needle age classes from 6–10 mg kg^-1 to 5 mg kg^-1. In boron fertilized plots the corresponding concentrations were 23–35 mg kg^-1 in control plots and 21–29 mg kg^-1 in limed plots. Both liming and B fertilizer decreased the Mn concentrations of needles. In the humus layer, total B concentration was increased by both lime and B fertilizer, and Ca and Mg concentrations and pH were still considerably higher in the limed plots than controls. Liming decreased the organic matter concentration in humus layer, whilst B fertilizer increased it.The results about B uptake were confirmed in a pot experiment, in which additionally the roles of increased soil pH and increased soil Ca concentration were separated by means of comparing the effects of CaCO₃ and CaSO₄. Two-year-old bare-rooted Norway spruce seedlings were grown in mor humus during the extension growth of the new shoot. The two doses of lime increased the pH of soil from 4.1 to 5.6 to 6.1, and correspondingly decreased the B concentrations in new needles from 22 to 12 to 9 mg kg^-1. However, CaSO₄ did not affect the pH of the soil or needle B concentrations. Hence the liming effect on boron availability in these soils appeared to be caused by the increased pH rather than increased calcium concentration.     

Does forest liming impact the enzymatic profiles of ectomycorrhizal communities through specialized fungal symbionts?

Liming (Ca–Mg soil amendment) is a forestry practice used to correct soil acidification and restore health and productivity in declining stands. Liming is known to modify tree mineral nutrition beyond the sole Ca and Mg. We hypothesized that liming also modifies the very functioning of the tree absorbing system (that is the ectomycorrhizal fine roots) in a way that facilitates the mobilization of mineral nutrients, particularly those entrapped in soil organic matter. This hypothesis has been tested here in beech and Norway spruce stands in North-Eastern France. In autumn, we compared the ectomycorrhizal community structure and the enzymatic profiles of ectomycorrhizal root tips in limed and untreated plots by measuring the activities of eight enzymes related to the degradation of soil organic matter. The results show that the ectomycorrhizal community responds to the Ca–Mg amendment and to the resulting soil modifications by modified enzyme activity profiles and ability to mobilize nutrients from soil organic matter. The effects of liming on the belowground functioning of the tree stands result essentially from specialized ECM fungal species such as Clavulina cristata (with strong glucuronidase activity), Lactarius subdulcis (with strong laccase activity) or Xerocomus pruinatus (with strong leucine aminopeptidase activity).     

Nitrogen transformations in limed and nitrogen fertilized soil in Norway spruce stands

Nitrogen transformations in the soil, and the resulting changes in carbon and nitrogen compounds in soil percolate water, were studied in two stands of Norway spruce (Picea abies L.). Over the last 30 years the stands were repeatedly limed (total 6000 kg ha^–1), fertilized with nitrogen (total about 900 kg ha^–1), or both treatments together. Both aerobic incubations of soil samples in the laboratory, and intact soil core incubations in the field showed that in control plots ammonification widely predominated over nitrification. In both experiments nitrogen addition increased the formation of mineral-N. In one experiment separate lime and nitrogen treatments increased nitrification, in the other, only lime and nitrogen addition together had this effect. In one experiment immobilization of nitrogen to soil microbial biomass was lower in soil only treated with nitrogen. Soil percolate water was collected by means of lysimeters placed under the humus layer and 10 cm below in the mineral soil. Total N, NH₄-N and NO₃-N were measured, and dissolved organic nitrogen was fractioned according to molecular weight. NO₃-N concentrations in percolate water, collected under the humus layer, were higher in plots treated with N-fertilizer, especially when lime was also added. The treatments had no effect on the N concentrations in mineral soil. A considerable proportion of nitrogen was leached in organic form.     

Four years continuous record of CH4-exchange between the atmosphere and untreated and limed soil of a N-saturated spruce and beech forest ecosystem in Germany

In order to gain information about seasonal and interannual variations of CH₄-fluxes at a spruce control site, a limed spruce site and a beech site of the Höglwald Forest, Bavaria, Germany, complete annual cycles of CH₄-exchange between the soil and the atmosphere with 2-hourly resolution were followed for 4 consecutive years. The ranges of CH₄ fluxes observed for the different sites were: +12.4 to –69.4 g CH₄ m^–2 h^–1 (spruce control site), +11.7 to –51.4 g CH₄ m^–2 h^–1 (limed spruce site), and –4.4 to –167.3 g CH₄ m^–2 h^–1 (beech site). Lowest rates of atmospheric CH₄-uptake or even a weak net-emission of CH₄ by the soils were observed during winter/spring times, whereas highest rates of CH₄-uptake were always found in summer/spring. Over the entire observation period of 4 years, mean CH₄-uptake rates were –1.82 kg CH₄-C ha^–1 yr^–1 at the spruce control site, –1.31 kg CH₄-C ha^–1 yr^–1 at the limed spruce site, and –4.84 kg CH₄-C ha^–1 yr^–1 at the beech site. The results obtained in this study demonstrate that in view of the huge interannual variations in CH₄-fluxes of approx. 1 kg CH₄-C ha^–1 yr^–1, multiple year measurements of CH₄-fluxes are necessary to accurately characterize the sink strength of temperate forest for atmospheric CH₄. By comparison of CH₄-fluxes measured at the spruce control site and the limed spruce site, a significant negative effect of forest floor liming on CH₄-uptake could be demonstrated. Compared to the spruce stand, the beech stand showed on average approx. 3 times higher rates of atmospheric CH₄-uptake, most likely due to pronounced differences between both sites with regard to the organic layer structure and bulk density of the mineral soil. Regression analysis between CH₄-fluxes and environmental parameters revealed that at all sites the dominating factors regulating temporal variations of CH₄ fluxes were soil moisture and soil temperature. Field measurements of CH₄ concentrations in the soil profile and laboratory measurements of CH₄-oxidation and CH₄-production activity on soil samples taken from different soil depths showed that the CH₄-flux at the Höglwald Forest sites is the net-result of simultaneous occurring production and consumption of CH₄ within the soil. Highest CH₄-oxidation activity was found in the uppermost centimeters of the mineral soil, whereas highest potential CH₄-production activity was found in the organic layer.     

Proportion of fungal mantle,cortex and stele of ectomycorrhizas in Picea abies (L.) Karst. in different soils and site conditions

Anatomical variability of ectomycorrhizal short roots in Norway spruce (Picea abies (L.) Karst) stands was investigated in five stands differing in site quality class (I^a–V) and soil type. Ten root samples per stand were randomly collected from the forest floor and the subsequent 20 cm soil layer. Thin transverse or axial sections (5 m) of randomly taken short roots were examined by light microscopy (AXIOPHOT; magnification 200–800×). All analyzed short root tips of Norway spruce in different stands were colonized by ectomycorrhizal fungi. Thickness of the mantle (T_mantle) and cortex (T_cortex) and diameter of the root (D_root) were measured in four crossing radial directions for transverse sections and in two radial directions for axial sections. The proportions of the mantle (PS_mantle), cortex (PS_cortex) and stele (PS_stele) of the root cross-sectional area (CSA) were calculated. Mean T_mantle and T_cortex varied from 16.5 ± 0.6 to 29 ± 1.3 m and 83.9 ± 1.7 to 108.4 ± 2.4 m, respectively; significant differences between stands were found. The number of cell rows in the cortex (4–6) did not vary between different stands, thus thickness of cortex depended on cell size. Mean PS_mantle, PS_cortex and PS_stele of the root CSA varied from 17.7 to 28.1%, from 58.9 to 66.9%, and from 13.4 to 15.8%, respectively. No differences between stands were revealed in mean CSA ratio of cortex and stele. It can be concluded that irrespective of big differences in soil and site conditions, including the influence of fungal symbionts, a 4:1 relation on a CSA basis between the cortex and stele is inherent to Norway spruce.     

Root and needle litter decomposition responses to enhanced supplies of N and S in a Norway spruce forest in southwest Sweden

The effects of additions of ammonium sulfate (NS) on the decomposition of litter derived from Norway spruce roots (< 2 and 2 - 5 mm in diameter) in the humus and mineral soil layers (0 - 15 cm) of a Norway spruce stand in southern Sweden were investigated over a 6-year period. To this purpose, litterbags were incubated in the humus layer and in the mineral soil in June 1996, with roots collected from NS and control (C) plots incubated in the NS and C plots, respectively. The N concentrations in fine roots (< 2 mm) in the NS- plots were higher than those in 2 - 5 mm roots in both humus and mineral soil layers. In the humus layer, N concentrations in the fine roots in the C- and NS- plots were 12.8 and 15.7 mg g ^{? 1}, respectively. By the end of the fifth year the < 2 mm roots in humus layer had lost 48.5 and 50% of their mass in the C and NS plots, respectively, while the corresponding values for the 2 - 5 mm diameter class were 44 and 54%. The fresh root litter may be a sensitive indicator to responses to enhanced N and S deposition, although decomposition rates of both litter types are affected.     

Chemical composition of soil solutions extracted from New Zealand beech forests and West German beech and spruce forests

Concentrations of ions were measured in soil solutions from beech (Nothofagus) forests in remote areas of New Zealand and in solutions from beech (Fagus sylvatica) and Norway spruce (Picea abies) forests in North-East Bavaria, West Germany, to compare the chemistry of soil solutions which are unaffected by acid deposition (New Zealand) with those that are affected (West Germany). In New Zealand, soil solution SO₄ ^2– concentrations ranged between <2 and 58 mol L^–1, and NO₃ ^– concentrations ranged between <1 and 3 mol L^–1. In West Germany, SO₄ ^2– concentrations ranged between 80 and 700 mol L^–1, and NO₃ ^– concentrations at three of six sites ranged between 39 and 3750 mol L^–1, but was not detected at the remaining three sites. At all sites in New Zealand, and at sites where the soil base status was moderately high in West Germany, pH levels increased, and total Al (Al_t) and inorganic monomeric Al (Al_i) levels decreased rapidly with increasing soil depth. In contrast, at sites on soils of low base status in West Germany, pH levels increased only slightly, and Al levels did not decline with increasing soil depth.Under a high-elevation Norway spruce stand showing severe Mg deficiency and dieback symptoms in West Germany, soil solution Mg²⁺ levels ranged between 20 and 60 mol L^–, and were only half those under a healthy stand. Al_t and Al_i levels were substantially higher the healthy stand than under the unhealthy stand, indicating that Al toxicity was not the main cause of spruce decline.     

The fate of 15N-labelled nitrate additions to a northern hardwood forest in eastern Maine,USA

We followed the movements of ¹⁵N-labelled nitrate additions into biomass and soil pools of experimental plots (15×15 m each) in a mid-successional beech-maple-birch-spruce forest in order to identify sinks for nitrate inputs to a forest ecosystem. Replicate plots (n=3) were spray-irrigated with either 28 or 56 kg N ha^–1 year^–1 using ¹⁵N-labelled nitric acid solutions (¹⁵N = 344 ) during four successive growing seasons (April–October). The ¹⁵N contents of foliage, bolewood, forests floor and mineral soil (0–5 cm) increased during the course of treatments. Mass balance calculations showed that one-fourth to one-third of the nitrate applied to forest plots was assimilated into and retained by above ground plant tissues and surface soil horizons at both rates of nitrate application. Plant and microbial assimilation were of approximately equal importance in retaining nitrate additions to this forest. Nitrate use among tree species varied, however, with red spruce showing lower rates of nitrate assimilation into foliage and bolewood than American beech and other deciduous species.     

The effect of acid rain,soil temperature and humidity on C-mineralization rates in organic soil layers under spruce

The biological activity in organic soil layers under spruce was determined by measuring rates of carbon dioxide emission. Under laboratory conditions, biological activity was found to be optimal at temperatures ranging from 20°C to 35°C and at water contents ranging from 40% to 60%. Weakly acidic to neutral pH values of organic materials stimulated microbial CO₂ formation whereas high acidity (pH<2.4) inhibited it. Low CO₂ emission rates were observed at pH<2.4 and therefore populations of microorganisms highly resistant to acid must be present in the organic materials. The O_L horizon was found to contribute 50% of the total potential C-mineralization with the lower horizons O_F and O_H contributing 25% each. In simulating acid rain experiments, analysis of water percolating through the organic layers was shown to cause constant leaching of cations at the pH 3 to 6.5 range. In this respect, the O_L horizon exerts a buffering effect. It can be postulated that the acidification of soil organic material under spruce by acid rain inhibits C-mineralization and decreases the release of mineral nutrients. In the long run this can be expected to affect the turnover of mineral nutrients in forest ecosystems.     

Acclimation of Two Distinct Plant Species,Spring Barley and Norway Spruce,to Combined Effect of Various Irradiance and CO2 Concentration During Cultivation in Controlled Environment

The short-term acclimation (10-d) of Norway spruce [Picea abies (L.) Karst] to elevated CO₂ concentration (EC) in combination with low irradiance (100 mol m^–2 s^–1) resulted in stimulation of CO₂ assimilation (by 61 %), increased total chlorophyll (Chl) content (by 17 %), significantly higher photosystem 2 (PS2) photochemical efficiency (F_v/F_m; by 4 %), and reduced demand on non-radiative dissipation of absorbed excitation energy corresponding with enhanced capacity of photon utilisation within PS2. On the other hand, at high cultivation irradiance (1 200 mol m^–2 s^–1) both Norway spruce and spring barley (Hordeum vulgare L. cv. Akcent) responded to EC by reduced photosynthetic capacity and prolonged inhibition of F_v/F_m accompanied with enhanced non-radiative dissipation of absorbed photon energy. Norway spruce needles revealed the expressive retention of zeaxanthin and antheraxanthin (Z+A) in darkness and higher violaxanthin (V) convertibility (yielding even 95 %) under all cultivation regimes in comparison with barley plants. In addition, the non-photochemical quenching of minimum Chl a fluorescence (SV₀), expressing the extent of non-radiative dissipation of absorbed photon energy within light-harvesting complexes (LHCs), linearly correlated with V conversion to Z+A very well in spruce, but not in barley plants. Finally, a key role of the Z+A-mediated non-radiative dissipation within LHCs in acclimation of spruce photosynthetic apparatus to high irradiance alone and in combination with EC was documented by extremely high SV₀ values, fast induction of non-radiative dissipation of absorbed photon energy, and its stability in darkness.     

Fine Root Production and Turnover in a Norway Spruce Stand in Northern Sweden: Effects of Nitrogen and Water Manipulation

Fine root length production, biomass production, and turnover in forest floor and mineral soil (0–30 cm) layers were studied in relation to irrigated (I) and irrigated-fertilized (IL) treatments in a Norway spruce stand in northern Sweden over a 2-year period. Fine roots (<1 mm) of both spruce and understory vegetation were studied. Minirhizotrons were used to estimate fine root length production and turnover, and soil cores were used to estimate standing biomass. Turnover was estimated as both the inverse of root longevity (RTL) and the ratio of annual root length production to observed root length (RTR). RTR values of spruce roots in the forest floor in I and IL plots were 0.6 and 0.5 y⁻¹, respectively, whereas the corresponding values for RTL were 0.8 and 0.9 y⁻¹. In mineral soil, corresponding values for I, IL, and control (C) plots were 1.2, 1.2, and 0.9 y⁻¹ (RTR) and 0.9, 1.1, and 1 y⁻¹ (RTL). RTR and RTL values of understory vegetation roots were 1 and 1.1 y⁻¹, respectively. Spruce root length production in both the forest floor and the mineral soil in I plots was higher than in IL plots. The IL-treated plots gave the highest estimates of spruce fine root biomass production in the forest floor, but, for the mineral soil, the estimates obtained for the I plots were the highest. The understory vegetation fine root production in the I and IL plots was similar for both the forest floor and the mineral soil and higher (for both layers) than in C plots. Nitrogen (N) turnover in the forest floor and mineral soil layers (summed) via spruce roots in IL, I, and C plots amounted to 2.4, 2.1, and 1.3 g N m⁻² y⁻¹, and the corresponding values for field vegetation roots were 0.6, 0.5, and 0.3 g N m⁻² y⁻¹. It was concluded that fertilization increases standing root biomass, root production, and N turnover of spruce roots in both the forest floor and mineral soil. Data on understory vegetation roots are required for estimating carbon budgets in model studies.     

Aluminum and iron chemistry in the O horizon changed by a shift in tree species composition

In the present study we characterized the buffering system including aluminum in the organic surface horizon (O) of field experiments with replicated plots of pure Norway spruce (Picea abies (L.) Karst) and replicated plots of spruce with a birch (Betula pendula Roth and B. pubescens Ehrh.) admixture. Our results show that a change in tree species composition may have a rather large, short-term (12 years) effect on the concentration of organically bound aluminum in the humus layer (H) of the O horizon. The pure spruce plots had a significantly higher concentration of exchangeable aluminum (Al_e) and organically bound (pyrophosphate extractable) aluminum (Al_P) in the H layer and a lower concentration of each base cation. Furthermore, our results confirm earlier findings that aluminum has base cation properties in acidic organic horizons. Thus, the change in tree species composition did not affect the pH in spite of a change in base saturation, because base cations were mainly compensated for by aluminum. The change in organically bound aluminum was accompanied by a similar change in organically bound pyrophosphate extractable iron (Fe_P). The observed differences between the mixed and pure spruce plots in the amounts of Al_P and Fe_P in the H layer could not be explained by estimated differences in biocycling of Al and Fe either by above-ground litterfall or by root turnover.     

Effect of clear-cutting on soil CO<Subscript>2</Subscript> emission

An experimental study to estimate the effect of clear-cutting on CO₂ emission from the soil surface was performed using the chamber method. For field measurements, several experimental plots within the clear-cut with different degrees of damage of the upper organic soil layer and different amounts of litter and logging residue on the surface were selected. Soil CO₂ fluxes were simultaneously measured both on the clear-cutting plots and on two plots within the spruce forest stand located close to the clear-cut area. The results show a significant seasonal and diurnal variability of soil CO₂ emission. It was found that the soil respiration rate varies significantly among plots and depends on the damage to the upper soil layer and the availability of litter and logging residue on the soil surface. It was found that the rate of CO₂ emission from soil surface is strongly dependent on the air and soil temperature and moisture of the upper soil layer. Different rates of soil respiration are also revealed on the plots located at different distances from tree trunks within the control forest stand.     

Aluminium effects on spruce and beech seedlings

Summary Natural spruce seedlings sampled on acid brown soil of the Belgian Ardennes, either on mineral soil, or on the litter layer, show higher Al content in the latter case, where substrate Al content is lower. Absorbing power of the root for Al organic chelates is probably higher than for Al⁺³ ions.Most soluble Al is chelated, especially in the holorganic and hemiorganic layers, under spruce and beech. In the mineral layers pH is above 3.8, and total Al in soil solutions is always found to be less than 2 eq/ml.     

The concept of succession in relation to the spread of sheathing mycorrhizal fungi on inoculated tree seedlings growing in unsterile soils

Summary Repeated annual assessments of the toadstools (fruitbodies) of mycorrhizal fungi associated with a mixed stand ofBetula spp. indicated that they were produced in a pattern ordered in time and space, suggesting a succession with identifiable early-and late-stage fungi. This concept is supported by below-ground observations of mycorrhizas which, however, need to be augmented.Both early- and late-stage mycorrhizal fungi form mycorrhizas on seedlings growing in axenic (aseptic) conditions. In contrast, only early-stage fungi seem able to trigger mycorrhizal formation on seedlings growing in unsterile soils.During axenic propagation, the early-stageHebeloma sacchariolens and the late-stageAmanita muscaria formed similar numbers of mycorrhizas per root system. After being transplanted to a range of unsterile field soils,A. muscaria failed to keep pace with the spread of the developing root system: no moreA. muscaria mycorrhizas were formed. On the other hand the continued development ofH. sacchariolens mycorrhizas precluded, during the first season after transplanting, the development of mycorrhizas by fungi naturally occurring in field soils. In the second season, however, the development ofH. sacchariolens mycorrhizas was restricted in acid peat but not in three other types of soil.The development ofLaccaria mycorrhizas after inoculating Sitka spruce with this fungus was associated, irrespective of soil type, with accelerated tree growth; with heights at the end of the first season being doubled.     

Changes in the extractability of cations (Ca,Mg and K) in the rhizosphere soil of Norway spruce (Picea abies) roots

Nutrient concentrations in the rhizosphere soil can be higher, lower or remain unchanged compared to the bulk soil, but relatively little is known about such changes for basic cations in the rhizosphere of tree roots. A modified root container technique of studying rhizosphere processes was employed. Plexiglas cylinders were horizontally split by a membrane with 30 M mesh size into an upper compartment for root growth and a root-free lower compartment, each with an inner diameter of 5 cm and a height of 10 cm. One 2-year-old Norway spruce (Picea abies) seedling was transplanted from a nursery into each cylinder. Plants were not specifically inoculated, but roots were colonised by a mix of ectomycorrhizal fungi originating from the nursery. The nutrient poor mineral soil used in the experiment was taken from a forest site in Bayerischer Wald, southern Germany. The soil was either supplied with a mix of Ca, Mg and K, or not supplied with these cations. Plants were harvested 30 weeks after transplanting. The nylon membrane between the root compartments restricted root growth to the upper compartment, so that by the end of the experiment a root mat was formed at the top side of the membrane. In the lower compartment, soil nearest to the root mat was regarded as rhizosphere soil while soil in a distance from the root mat was regarded as bulk soil. In the upper compartment, rhizosphere soil was obtained at the end of the experiment by gently shaking the roots. The soils were analysed for Ca, Mg and K contents following two different soil extraction methods. In the fertilised treatment, H₂O-extractable Ca and Mg were accumulated in the rhizosphere. In contrast, K (NH₄Cl-extraction) was depleted in the rhizosphere. In the bottom tube, the depletion of K (NH₄Cl-extraction) was restricted to 1 cm distance from the root mat. In unfertilised soil, Ca, Mg and K concentrations did not differ clearly between rhizosphere and bulk soils. The results indicated that the occurrence of cation gradients in the rhizosphere depended on the level of soil nutrient supply. Distinct rhizosphere effects were measured by conventional soil extraction methods only when the soil was freshly fertilised with mineral elements prior to the experiment. In this case, K depletion in the rhizosphere reflected higher K uptake by the fertilised Norway spruce plants. For low-nutrient soils, novel techniques are required to follow subtle changes in the rhizosphere.     

Fine root mass in relation to soil N supply in a cool temperate forest

Soil inorganic nitrogen supply and fine root mass in the top layers of mineral soil (0–5 and 5–10cm) were investigated at upper and lower sites of a cool temperate forest where Fagus crenata and Quercus crispula dominate. At both sites, soil inorganic nitrogen supply was greatest in the 0–5cm layer. The predominant forms of soil inorganic nitrogen supply were NH₄⁺-N at the upper site and NO₃^–-N at the lower site. Fine roots were concentrated in the 0–5cm layer at the upper site, but not at the lower site. The form of supplied soil inorganic nitrogen supply can be important in determining the vertical distribution of fine roots.