Climate and forest management influence nitrogen balance of European beech forests: microbial N transformations and inorganic N net uptake capacity of mycorrhizal roots

The effects of local climate and silvicultural treatment on the inorganic N availability, net N uptake capacity of mycorrhizal beech roots and microbial N conversion were assessed in order to characterise changes in the partitioning of inorganic N between adult beech and soil microorganisms. Fine root dynamics, inorganic N in the soil solution and in soil extracts, nitrate and ammonium uptake kinetics of beech as well as gross ammonification, nitrification and denitrification rates were determined in a beech stand consisting of paired sites that mainly differed in aspect (SW vs. NE) and stand density (controls and thinning treatments). Nitrate was the only inorganic N form detectable in the soil water. Its concentration was high in control plots of the NE aspect, but only in canopy gaps and not influenced by thinning. Neither thinning nor aspect affected the abundance of root tips in the soil. Maximum nitrate net uptake by mycorrhizal fine roots of beech, however, differed with aspect, showing significantly lower values at the SW aspect with warm–dry local climate. There were no clear-cut significant effects of local climate or thinning on microbial N conversion, but a tendency towards higher ammonification and nitrification and lower denitrification rates on the untreated controls of the SW as compared to the NE aspect. Apparently, the observed sensitivity of beech towards reduced soil water availability is at least partially due to impaired N acquisition. This seems to be mainly a consequence of reduced N uptake capacity rather than of limited microbial re-supply of inorganic N or of changed patterns of inorganic N partitioning between soil bacteria and roots.     

Effects of vegetation control on nutrient removal and Fagus orientalis, Lipsky regeneration in the western Black Sea Region of Turkey

Dense Rhododendron ponticum (L.) understories of eastern beech (Fagus orientalis Lipsky) stands in the Black Sea Region (BSR) of Turkey create challenging forest vegetation management problems relative to beech regeneration. Rhododendron traditionally is controlled in Turkish forests with grubbing and bulldozing. The effects of these practices on nutrient removal and natural beech regeneration have not been quantitatively studied. Two woody vegetation control treatments (bulldozing and hand-grubbing) were installed during late summer, 2002 in three mature beech stands with dense rhododendron understories in the Düzce Forest Management Directorate, in the Turkish western BSR. Aboveground biomass of each vegetation component, total aboveground vegetation biomass, nutrient concentrations, organic matter (OM) removal, and total amount of OM nutrients were determined for each woody vegetation control treatment. Soil bulk density and nutrient content, and beech seedling biomass, nutrient content, and natural regeneration also were studied. One year after treatment, the machine site preparation by bulldozing (MSP) that removed understory vegetation and attached roots, reduced mean forest floor OM content by about 84%, when compared to hand-grubbing. Mean soil C, N, K and Mg concentrations on the bulldozed sites were 36, 27, 50 and 55% less, respectively, than those on the grubbed sites. Total C and Mg amounts at the 5–10 cm soil depth were 24 and 47% lower, respectively, for mechanical site preparation (MSP) sites, when compared to grubbed sites. Overall, soil bulk density did not differ significantly between the grubbing and MSP treatments. Frequent passes on designated transects on MSP sites resulted in a significant (P < 0.01) one-third increase in bulk density of the first 20 cm soil depth, when compared to grubbed sites. No natural beech regeneration occurred on untreated control sites. Mineral soil exposure on the MSP sites increased beech seedling germination substantially. One year after treatment, the mean number of naturally regenerated seedlings for MSP sites was 2.5 times greater than for grubbed sites. Woody control treatments had significant effects upon beech seedling chemistry, with N, P, K, Ca and Mg concentrations averaging 35, 47, 12, 33 and 25% lower, respectively, for MSP sites, when compared to grubbed sites. However, mean seedling biomass and nutrient content were significantly greater (P < 0.05) on MSP sites. Long-term effects from windrowing on MSP site residues, associated topsoil and nutrients need to be evaluated.     

Light,soil, and seedling characteristics associated with varying levels of competition in a red pine plantation

Red pine (Pinus resinosa Ait.) seedlings growing under differing levels of competition were evaluated during the fifth growing season following planting, and placed into low, moderate, or high tree classes, as a function of levels of competing vegetation. Tree growth, moisture status, and nutrition were monitored over the growing season. Additionally, site characteristics such as soil temperature and moisture, inorganic nitrogen concentration, mineralized soil nitrogen, and light were measured. Red pine seedlings growing under low competition had an absolute volume growth increase of 795% over the seedlings growing under the heaviest competition. The associated relative volume growth increase during the fifth growing season was 44%. Discriminant analysis was used to describe three classes of trees representing low, moderate, and high levels of competition. Trees growing under low competition had longer and heavier needles, but generally lower nutrient concentrations. Pre-dawn plant moisture status did not vary among competition levels. Soil variables indicated that, in general, the trees growing under low competition occupied warmer, drier, and less fertile microsites, as inorganic soil N and mean monthly mineralized NO₃-N and NH₄>-N tended to be lower on these microsites. The suite of independent variables was effective in classifying the model data into three tree competition classes, with percent correct classifications of 92, 75, and 100 for low (tree class one), moderate (tree class two), and heavy (tree class three) competition, respectively.     

Morphological and physiological responses of beech (Fagus sylvatica) seedlings to grass-induced below ground competition

We examined morphological and physiological responses of beech (Fagus sylvatica L.) seedlings to grass-induced below ground competition in full-light conditions. Two-year-old beech seedlings were grown during two growing seasons in 160-l containers in bare soil or with a mixture of five grass species widely represented in semi-natural meadows of central France. At the end of the second growing season, beech seedlings in the presence of grass showed significant reductions in diameter and height growth, annual shoot elongation, and stem, root and leaf biomass, but an increase in root to shoot biomass ratio. Grasses greatly reduced soil water availability, which was positively correlated with daily seedling diameter increment. Beech seedlings seemed to respond to water deficit by anticipating stomatal closure. There was evidence of competition for nitrogen (N) by grasses, but its effect on seedling development could not be separated from that of competition for water. By labeling the plants with 15N, we showed that beech seedlings absorbed little N when grasses were present, whereas grasses took up more than 97% of the total N absorbed in the container. We conclude that, even if beech seedlings display morphological and physiological adaptation to below ground competition, their development in full-light conditions may be strongly restricted by competition from grass species.     

Effects of different soil treatments on growth and net nitrogen uptake of newly planted Picea abies (L.) Karst. seedlings

Five soil treatments in a 4-year-old clearcut in southern Sweden affected biomass increase and net nitrogen uptake by planted Norway spruce (Picea abies (L.) Karst.) seedlings through their on net mineralisation and root growth. The patch soil treatments studied were: (i) soil inversion in an old clearcut; (ii) mineral soil from the clearcut remaining, (iii) mineral soil from the clearcut with fertiliser application during the first season; (iv) mineral soil from a nearby uncut forest replacing the clearcut mineral soil; and (v) an untreated control. Growth increased in seedlings in treatments (i) and (iii), but growth in soil treatments with humus removal was not better than that of seedlings in untreated soil. High N uptake early in the first growing season resulted in increased growth during this season in contrast to late N uptake that resulted in a high N concentration in the seedlings after the first growing season. This in turn led to a high growth rate during the next growing season. Generally, both root growth and net N mineralisation were positively correlated to N uptake in the soil treatments. Therefore, a combination of low net N mineralisation and poor root growth as a result of high soil density appears to explain the low N uptake in seedlings in undisturbed soil. The importance of competition with field vegetation for N and water was not clear. Net mineralisation was larger in soil treatments where the humus layer was retained than where it was removed. Net N mineralisation in soil from old clearcuts was the same as in soil from fresh clearcuts.     

Nutrient concentrations in roots, leaves and wood of seedling and mature sugar maple and American beech at two contrasting sites

Differences in sensitivity to soil conditions across tree species and developmental stage are important to predicting forest response to environmental change. This study was conducted to compare elemental concentrations in leaves, stems, and roots of (1) sugar maple (Acer saccharum Marsh.) seedlings vs. mature trees and (2) mature sugar maple vs. mature American beech (Fagus grandifolia Ehrh.) in two sites that differ in soil base saturation and pH. Both sites are located in Huntington Forest, NY, USA; one site (hereafter ‘H’) has higher soil pH and Ca, Mg, and Mn concentrations than the other site (hereafter ‘L’). Sugar maple growth at H (14.8 cm² year⁻¹ per tree) was much greater than at L (8.6 cm² year⁻¹ per tree), but the growth of beech was not different between the two sites. Leaves, roots, and stem wood of mature beech trees and sugar maple seedlings and mature trees were sampled for nutrient analysis. Foliar Ca, K, and Al concentrations were positively correlated with soil elements, but Mn concentrations were negatively correlated. Sugar maple differed more than beech between sites in foliar K and Mn concentrations. Root Mg and P concentrations reflected soil chemistry differences, in contrast to foliar concentrations of Mg and P, which were indistinguishable between the sites. In sugar maple, seedlings differed more than in mature trees in nutrient concentrations in roots, especially for Mg and Mn. Although beech was not as responsive to nutrient availability as sugar maple in foliar and root nutrient concentrations, Ca and Mg concentrations in beech wood were higher in H (52% higher for Ca and 68% higher for Mg), while sugar maple did not differ between sites. Sugar maple regeneration failure on acidic soils in the same region is consistent with our finding that sugar maple seedlings were very sensitive to nutrient availability. This sensitivity could ultimately contribute to the replacement of sugar maple by American beech in regions of low pH and base cations if base cation leaching by anthropogenic deposition and tree harvesting continues.     

Fate of nitrogen released from 15N-labeled litter in European beech forests

The decomposition and fate of 15N-labeled beech litter was monitored in three European beech (Fagus sylvatica L.) forests (Aubure, France; Ebrach, Germany; and Collelongo, Italy) for 3 years. Circular plots around single beech trees were isolated from roots of neighboring trees by soil trenching, and annual litterfall was replaced by 15N-labeled litter. Nitrogen was continuously released from the decomposing litter. However, over a 2-year period, this release was balanced by the incorporation of exogenous N. Released N accumulated mainly at the soil surface and in the topsoil. Microbial biomass remained almost constant during the experiment at all sites except for considerably lower values at Ebrach. The 15N enrichment of the microbial biomass increased strongly during the first year and then remained stable. The 15N released from the decomposing litter was rapidly detected in roots and leaves of the beech trees, increasing regularly and linearly over the course of the experiment. The uptake of litter-released 15N by the trees was reduced under conditions that reduced tree growth. Under these conditions, leaves and fine roots were the dominant N sinks, and little N was allocated to other plant parts. By contrast, N uptake and N allocation from leaves to stem and bark tissues increased when tree growth was enhanced. Budgets for 15N showed that 2 to 4% of litter-released N was incorporated into the trees, about 35% remained in the litter and about 50% reached the topsoil.     

Influence of nurse birch and Scots pine seedlings on early aerial development of European beech seedlings in an open-field plantation of Central France

Beech woodland can be restored by direct planting of beech (Fagussylvatica L.) seedlings in abandoned areas, but this methodis generally avoided in forestry because of the growth difficultiesof beech in full-light conditions. This study tested a methodthat consists of planting beech seedlings in full-light conditionswith silver birch (Betula pendula Roth.) or Scots pine (Pinussylvestris L.) as nurse trees. A total of 65 two-year-old beechseedlings surrounded by either 3, 4, 5 or 6 pine or birch seedlingsor without competitors were planted in bare-soil open-fieldconditions in Central France. Tree growth and light availabilitywere monitored at the end of each of the following 3 years,and soil moisture was also measured the last year. At the endof the experiment, aerial biomass measurements were made onthe beeches. Results showed that relative beech growth in termsof diameter was significantly reduced by the local neighbouringtrees and that this reduction was particularly noticeable inthe pine treatments where there was extensive pine seedlingdevelopment. In contrast, relative beech height did not exhibitany significant variations among the treatments. Light availabilitywas decreased by the neighbouring trees, especially in the pinetreatments. Light reduction was more pronounced in the last2 years and in the middle or lower parts of the canopy of thesubject beech. Soil water content was lower under the pine canopiesthan under the birch canopies, and was positively correlatedto beech relative diameter growth. Specific leaf area, height-to-diameterratio and crown length-to-crown width ratio weakly but significantlyincreased with competition from the neighbours. Beech biomasspartitioning was only weakly affected by the treatments. Thisstudy showed that nurse trees tend to reduce beech growth butimprove form although effects remained weak due to the insufficientlength of the experiment. Further studies are also needed toquantify the effects of the neighbourhood on the growth of competitiveherbaceous vegetation and on changes in the microclimatic conditions.     

Water and element fluxes during the regeneration of Norway spruce with European beech: Effects of shelterwood-cut and clear-cut

The regeneration of mature Norway spruce with European beech using the shelterwood silvicultural system is a good example of continuous cover forestry. In contrast, the regeneration may also start with clear-cut plots, which often occur after calamities like wind-throw or bark beetle attack. During regeneration the forest ecosystem becomes a highly dynamic system. Nutrient losses from the soil may occur as the element turnover is affected by the reduced nutrient uptake of forest trees as well as the enhanced mineralisation and nitrification due to higher soil temperature and soil moisture. Continuous cover forestry may help to reduce these nutrient losses. In order to test this, we investigated water and element fluxes of two chronosequences. The first investigated regeneration in the shelterwood system, while the second concerned itself with regeneration on clear-cut plots. In a shelterwood-cut about 30% of the mature spruce trees are removed and young beech trees are planted. Some 10 years later a secondary felling is carried out and at age 20 of the beech regeneration the final harvest of the mature trees occurs. Thus, the studied time steps were (a) the first 5 years after the initial cut and planting, (b) 10-year-old beech regeneration after the second shelterwood cut and (c) 20-year-old beech regeneration after the final harvest.Our results indicate that nutrient losses with seepage water – especially nitrogen, calcium and magnesium – occur during the first years after the clear cut and, to a lesser extent, after secondary felling on the selective-cut plot. This may temporarily affect seepage water quality due to elevated nitrate concentrations, which reached values of more than 100 mg l⁻¹. In the time span between planting and an age 20 of the beech regeneration, total losses of nitrogen from the main rooting zone reach 230 kg ha⁻¹ after clear cut. Preliminary estimates of the total nitrogen loss in the shelterwood system range between 150 and 230 kg ha⁻¹ indicating either significantly lower or equal losses of nutrients. In the second case, however, element output is distributed more equally over the 20-year-period than after clear felling where 85% of the nitrate leaching occurs during the first 3 years.     

Carbon partitioning and allocation in northern red oak seedlings and mature trees in response to ozone

Northern red oak (Quercus rubra L.) seedlings and trees differ in their response to ozone. Previous work reported reductions in net photosynthesis, carboxylation efficiency and quantum yield of mature tree leaves, whereas seedling processes were unaffected by the same ozone exposure. To further characterize differences in ozone response between seedlings and mature trees, we examined carbon partitioning and allocation in 32-year-old trees and 4-year-old seedlings of northern red oak after exposure to subambient (seasonal SUM00 dose (sum of all hourly ozone exposures) = 31 ppm-h), ambient (SUM00 dose = 85 ppm-h) and twice ambient (SUM00 dose = 151 ppm-h) ozone concentrations for three growing seasons. For mature trees, ozone exposure decreased foliar starch partitioning, increased starch partitioning in branches and increased (14)C retention in leaves. In contrast, starch partitioning in leaves and branches, and foliar (14)C retention in seedlings were unaffected by ozone exposure, but soluble carbohydrate concentrations in coarse and fine roots of seedlings were reduced. Differences in carbohydrate demand between seedlings and mature trees may underlie the higher leaf ozone uptake rates and greater physiological response to ozone in mature northern red oak trees compared with seedlings.     

氮肥对盆栽欧洲水青冈生物量和营养分配的影响(英文)

调查了施加氮肥(15NH4和15NO3)处理后在两个连续生长季内欧洲水青冈(Fagus sylvatica L.)幼苗地上部分和地下部分的生物量和营养元素分配。盆栽欧洲水青冈幼苗培养于温室大棚内,培养土样取自相邻的三种林分:欧洲水青冈,挪威赤松,欧洲水青冈-赤松混交林。结果表明,氮肥(15N)处理对欧洲水青冈营养元素分配没有显著影响,施加氮素形式决定自身流入植物库的情况。欧洲水青獭收氮素主要以硝态氮的形式,因此,尽管植物体内保存的硝态氮和氨态氮并没有统计差异,但是叶片中硝态氮明显减少。施加硝态氮对欧洲水青冈氮素恢复的影响要大于施加氨态氮。与欧洲水青冈茎、粗根相比,优质根系对氮素(15N)固定是一个缓慢过程。表8图1参40。     

Effects of root trenching of overstorey Norway spruce (Piceaabies) on growth and biomass of underplanted beech (Fagussylvatica) and Douglas fir (Pseudotsugamenziesii) saplings

In Central Europe, the conversion of pure Norway spruce stands (Picea abies [L.] Karst.) into mixed stands with beech (Fagus silvatica L.) and other species like e.g. Douglas fir (Pseudotsuga menziesii [Mirb.] Franco) is accomplished mainly by underplanting of seedlings beneath the canopy of overstorey spruce trees after partial cutting treatments what means exposure to shade and below-ground root competition by the overstorey to the seedlings. Particularly about the second factor, our knowledge is limited. Therefore, we carried out a below-ground competition exclusion experiment by root trenching and investigated the effects on soil resources, growth, and biomass partitioning of underplanted beech and Douglas fir saplings under target diameter and strip cutting treatments. The exclusion of overstorey root competition by trenching increased the soil water potential in the second year that had a fairly dry growing season and led to significantly higher foliar concentrations of most nutrients, particularly in Douglas fir, indicating an amended nutrient supply. Both improvements were accompanied by an increase in length and diameter increment of the underplanted saplings, appearing in both species only after having surpassed a species-specific threshold light value (Douglas fir 16% of above canopy radiation, beech 22%). We also found significant interactions between trenching and light for specific fine root length and further biomass and morphological parameters. Judged by the much steeper increase in height and diameter growth with increasing light after release from below-ground competition, Douglas fir saplings appeared to be more sensitive to root competition than beech saplings what conforms to older findings for beech. According to our results, a strip cutting seems to be more appropriate than a target diameter cutting treatment to replace a pure spruce stand by a mixed stand with beech and Douglas fir.     

Growth and physiological response of eastern white pine seedlings to partial cutting and site preparation

We examined how white pine (Pinus strobus L.) seedlings planted under a mature cover of white and red (Pinus resinosa Ait.) pine in eastern Ontario (Canada) responded to treatments aimed at improving light and soil conditions for seedling growth. The treatments were: (a) three levels of partial cutting (no cut or CS0, cut to one-crown spacing between residual trees or CS1, cut to two-crown spacing or CS2); (b) two levels of vegetation control (without herbicide or H0, with herbicide or H1); and (c) two levels of soil scarification (S0 and S1). On the third growing season after planting, total growth of seedlings was lowest in CS0 treatment and similar in CS1 and CS2 treatments. The CS2 created better growing light conditions than the CS1, with and average of 50% of full light at seedling height, which corresponded to the maximum height and diameter growth rates of seedlings. However, CS2 also stimulated the growth of competing woody vegetation (both understory trees and shrubs), and resulted in greater microsite heterogeneity of light availability. Scarification warmed the soil (approximately 1–3 °C in the middle of the growing season), decreased the density of competing trees, but increased the shrub density, with no impact on white pine seedling growth. The treatments had no effect on light-saturated photosynthetic rate (A) of current-year foliage of seedlings, nor on their midday shoot water potential. Leaf N was higher in partial cuts and with vegetation control, but the relationship between N and A was weak to non-existent for the different foliage classes. Measures of the proportion of aboveground biomass allocated to foliage (leaf-mass ratio) suggest an acclimation response of young white pine that improves growth under moderate light availability and compensates for the lack of leaf-level photosynthetic plasticity. We suggest a combination of soil scarification under a one-crown spacing partial cut (corresponding to 14 m² ha⁻¹ of residual basal area, or an average of 32% of available light at seedling height) as an establishment cut. This should provide optimum growth conditions for planted understory white pine, while also favoring natural regeneration and providing some protection against damage from insects and disease.     

Patterns of regeneration and ground vegetation associated with canopy gaps in a managed beechwood in southern England

In autumn 2001, 15 canopy gaps were selected for study in RumerhedgeWood, a semi-natural, mesotrophic beechwood in southern England.The gaps were located in mature, beech-dominated stands, andhad originated from openings created during a thinning in theearly 1980s and wind damage in 1987/1990 and/or the consequentsalvage operations. The extent of each gap and surrounding treeswere mapped. Tree/shrub regeneration, ground vegetation, bareearth, leaf litter and canopy openness (using a canopy-scope)were measured within and around the gaps using a 5 x 5-m gridand placing a 1 x 1-m quadrat at each grid intersection (totalnumber of quadrats = 400). Most of the gaps were <75 m² inarea. The largest was 241 m². They were generally irregularin shape and there was little or no understorey present. Mostsurrounding trees were beech Fagus sylvatica L. Bramble Rubusfruticosus L. formed a moderate to dense ground vegetation belowmost gaps and declined around the edges only once the gap openingwas substantially obscured. Apart from a few larger saplings,most regeneration was small and of beech. Most of the latterappeared to be in their fifth or sixth growing season, were10–35 cm tall, had an erect base and flat top, had increasedby <5 cm in height during 2001 and were not browsed by deer.Their height and growth form was related to (1) their positionwithin gaps, (2) the degree of canopy openness and (3) the coverof ground vegetation. This was translated into the followingzonation—(1) around the centre of larger gaps: canopyopenness increased to >15 per cent; bramble cover was nearcomplete; litter depth was low; many places had no beech seedlings,but some of the few present were among the tallest, most uprightand fastest growing; (2) towards the edges of the large gapsor directly below smaller gaps: canopy openness was about 4–10per cent; bramble cover was slightly less; beech seedlings weremoderately abundant but patchy, generally shorter, more flattopped and slow growing than in the gap centre, albeit somewere still among the tallest, most upright and fastest growing;(3) beyond the edge of the large/medium gaps (with the gap onlypartially visible) or directly below very small gaps: canopyopenness was only about 2–3 per cent; bramble was muchreduced; beech seedlings were at their most abundant but stillpatchy in distribution and even shorter, more flat topped andslower growing than in the above zone; (4) in an outer zonebeyond or almost beyond the sight of the gaps: canopy opennesswas <2 per cent; bramble was weak and sparse; beech seedlingswere mostly at low densities and predominately short, slow growingand flat topped. This ring pattern of beech regeneration appearedto relate mainly to (1) differences in light availability affectingthe survival, growth rate and form of seedlings; (2) competitionfrom bramble and possibly (3) limited dispersal of beech seedinto gaps. Recommendations are given for managers who wish touse natural regeneration to restock beech woodland.     

Changes in fine root production and longevity in relation to water and nutrient availability in a Norway spruce stand in northern Sweden

The PEACH computer simulation model of reproductive and vegetative growth of peach trees (Grossman and DeJong 1994) was adapted to estimate seasonal nitrogen (N) dynamics in organs of mature peach (Prunus persica (L.) Batsch cv. O'Henry) trees grown with high and low soil N availability. Seasonal N accumulation patterns of fruits, leaves, stems, branches, trunk and roots of mature, cropping peach trees were modeled by combining model predictions of organ dry mass accumulation from the PEACH model with measured seasonal organ N concentrations of trees that had been fertilized with either zero or 200 kg N ha(-1) in April. The results provided a comparison of the N use of perennial and annual organs during the growing season for trees growing under both low and high N availability. Nitrogen fertilization increased tree N content by increasing organ dry masses and N concentrations during the fruit growing season. Dry mass of current-year vegetative growth was most affected by N fertilization. Whole-tree N content of fertilized trees was almost twice that of non-fertilized trees. Although N use was higher in fertilized trees, calculated seasonal N accumulation patterns were similar for trees in both treatments. Annual organs exhibited greater responses to N fertilization than perennial organs. Estimated mean daily N use per tree remained nearly constant from 40 days after anthesis to harvest. The calculations indicated that fertilized trees accumulated about 1 g N tree(-1) day(-1), twice that accumulated by non-fertilized trees. Daily N use by the fertilized orchard was calculated to be approximately 1 kg N ha(-1), whereas it was approximately 0.5 kg N ha(-1) for the non-fertilized trees. During the first 25-30 days of the growing season, all N use by growing tissues was apparently supplied by storage organs. Nitrogen release from storage organs for current growth continued until about 75 days after anthesis in both N treatments.     

Comparison of soil nitrogen dynamics under beech,Norway spruce and Scots pine in central Germany

To investigate the effect of tree species on soil N dynamics in temperate forest ecosystems, total N (Nt), microbial N (Nmic), net N mineralization, net nitrification, and other soil chemical properties were comparatively examined in beech (64–68 years old) and Norway spruce (53–55 years old) on sites 1 and 2, and beech and Scots pine (45 years old) on site 3. The initial soil conditions of the two corresponding stands at each site were similar; soil types were dystric Planosol (site 1), stagnic Gleysols (site 2), and Podzols (site 3). In organic layers (LOf₁, Of₂, Oh), Nmic and Nmic/Nt, averaged over three sampling times (Aug., Nov., Apr.), were higher under the beech stands than under the corresponding coniferous ones. However, the Nmic in the organic layers under beech had a greater temporal variation. Incubation (10 weeks, 22 °C, samples from November) results showed that the net N mineralization rates in organic layers were relatively high with values of 8.1 to 24.8 mg N kg^–1 d^–1. Between the two corresponding stands, the differences in net N mineralization rates in most of the organic layers were very small. In contrast, initial net nitrification rates (0.2–17.1 mg N kg^–1 day^–1) were considerably lower in most of the organic layers under the conifer than under the beech. In the mineral soil (0–10 cm), Nmic values ranged from 4.1–72.7 mg kg^–1, following a clear sequence: August>November>April. Nmic values under the beech stands were significantly higher than those under the corresponding coniferous stands for samples from August and April, but not from November. The net N mineralization rates were very low in all the mineral soils studied (0.05–0.33 mg N kg^–1 day^–1), and no significant difference appeared between the two contrasting tree species.     

Influence of woody and herbaceous competition on microclimate and growth of eastern white pine (Pinus strobus L.) seedlings planted in a central Ontario clearcut

The influence of woody and herbaceous plant competition, either alone or in combination, on microclimate and growth of planted eastern white pine (Pinus strobus L.) seedlings was examined over four consecutive growing seasons in a central Ontario clearcut. Treatments that manipulated the comparative abundance of these two plant functional groups significantly affected light availability, soil moisture, and air and soil temperature regimes. These microclimate alterations, coupled with the relative competitiveness of herbaceous and woody vegetation, corresponded to temporal changes in vegetation cover and dominance. The more rapid colonization and growth of the herbaceous plant community, dominated by bracken fern (Pteridium aquilinum) and ericaceous shrubs (Kalmia sp., Vaccinium sp.), resulted in this form of vegetation being a comparatively important early competitor for soil moisture. As the woody plant community, dominated by naturally regenerated trembling aspen (Populus tremuloides Michx.), grew in height and leaf area, it became a comparatively strong competitor for both light and soil moisture. For all vegetation treatments combined, white pine seedling growth responses were strongly correlated with total cover of competing vegetation and its relative influence on above- and belowground microclimatic variables. Higher total cover of competing vegetation was generally associated with lower light and soil moisture availability and cooler soil temperatures. Multiple regression analyses indicated that pine seedling relative height growth increased with soil moisture content and growing season soil heat sum, while seedling relative diameter and relative volume growth increased with light availability.     

Soluble N compound profiles and concentrations in European beech (Fagus sylvatica L.) are influenced by local climate and thinning

We assessed seasonal changes of total soluble nonprotein nitrogen compounds (TSNN) in adult European beech trees (Fagus sylvatica, L.) growing under different local climate during the growing season immediately following a thinning treatment and 3 years later. In both years, samples of leaves, xylem sap and phloem exudates from beech trees growing in thinned and unthinned (control) stands on a dry, warm SW exposed and a cooler, moist NE exposed site were collected in May, July and September. In May of both years, asparagine (Asn) and glutamine (Gln) were most abundant in leaves and xylem, respectively, whereas arginine (Arg) dominated in the phloem. In July, TSNN concentrations decreased in all tissues and sites, but differences in water availability between aspects were reflected in TSNN concentrations. In September, differences in the increase of Arg concentration in the phloem were related to differences in the onset of senescence between treatments. Thinning treatment increased amino compound concentrations of beech tissues in July on both aspects, particularly at the NE thinned site. It is supposed that, the N balance of adult beech is favoured by both, the thinning treatments as well as the cool-moist climate prevailing at the NE aspect.     

Impact of slash removal,drag scarification,and mounding on lodgepole pine cone distribution and seedling regeneration after cut-to-length harvesting on high elevation sites

Excessive slash loading could pose a problem for the regeneration of the serotinous lodgpole pine especially in forests at higher elevation where soil temperature is limiting. In the past, these forests have commonly been harvested using full-tree harvesting where trees are processed at roadside; however, recently cut-to-length harvesting has become a more frequent harvesting method. In cut-to-length harvesting the harvested trees are processed in the block, as a result slash accumulation is much higher on these cutblocks. In an experimental field trial, the cone distribution, natural lodgepole pine regeneration, and the growth and establishment of planted lodgepole pine were evaluated in response to slash load, drag scarification, and mounding after cut-to-length harvesting of high elevation lodgepole pine stands in the Rocky Mountains. Twelve sites were established, each contained six plots which were randomly assigned to six treatment combinations of two slash removal (slash and slash removed) and three mechanical soil preparation treatments (no soil preparation, drag scarifying, and mounding). The slash removal reduced slash volume by more than 50% but also reduced the number of lodgepole pine cones available for regeneration by over 33%. However, soil mechanical treatments offset this effect as fewer cones were necessary to achieve high natural pine regeneration densities. Drag scarification of plots resulted in 12 times the number of pine seedlings compared to the non-prepared plots. Although slash removal did not have an effect on the number of naturally regenerated lodgepole pine seedlings, it had a positive effect on their growth performance. Conversely, planted pine seedlings had lower mortality and better growth in soils that had been mechanically prepared and had the slash removed; however, the growth effects became only apparent 4 years after planting. While slash removal and mechanical soil preparation did increase soil temperatures; the slash removal treatment had a more transient effect on soil temperatures than soil preparation. Differences in soil temperature decreased over time which appeared to be mostly driven by a warming of the soils in the plots with no soil preparation, likely a result of the decomposition of the finer slash and feathermosses. Overall, it appears that surface disturbance on these high elevation sites had a far greater effect on lodgepole pine regeneration and growth than the increased accumulation of slash as a result of cut-to-length harvesting.     

Indices of soil nitrogen availability for an ecological site classification of British forests

An adequate supply of nitrogen (N) is essential for the successful establishment and sustainable productivity of forest stands. N deficits may necessitate the use of artificial fertilisers. Availability of N in the inorganic forms, and the relative abundance of the NO₃-N and NH₄-N components, influences the species composition of natural forest vegetation. Hence it is essential to use reliable measures of soil N supply that fully reflect its ecological significance. The new Ecological Site Classification (ESC) used in British forestry employs a multi-factorial definition of soil nutrient regime (SNR), including soil N. To develop this, a soil and vegetation study was made at 89 forest sites throughout Great Britain covering the major soil types used for forestry. “Total N” levels were compared with separate pre- and post-incubation measures of the two inorganic N components as potential indices of soil N supply. Multivariate statistical analysis showed that the major discriminant chemical variables for the sampled soils were pH, calcium and NO₃-N and that these were also the main variables influencing the species composition of the ground vegetation. Total N and NH₄-N were less effective discriminant variables for these sites. In some infertile soils the levels of NH₄-N or total N may be of greater importance, as NO₃-N is usually in very limited supply. A multivariate gradient of SNR, which incorporates the NO₃-N measures, has been adopted for use within the ESC system. The position of a site on this gradient can be estimated quantitatively from soil type, ground vegetation species composition and humus type. This enables soil N supply and overall SNR to be assessed in a simple but effective way that guides the operational management of British forest soils for sustainable productivity. It will also be possible to use these techniques to monitor the nutritional status of forest sites over time.