Radiation regime and nitrogen supply in modelled alley cropping systems ofErythrina poeppigiana with sequential maize-bean cultivation

A simulation study relating the effect of the spatial arrangement of trees on the photon exposure above an associated crop (PE_i) was carried out for an alley cropping system of the leguminous treeErythrina poeppigiana (Walpers) O. F. Cook with sequential maize-bean cultivation. These results were combined with estimates of the nitrogen supplied by prunings from different tree densities to rank the arrangements. A significant difference was found in the PE_i between the east-west and north-south-oriented alleys. The latter exhibited more areas of moderate shade, while the former has unshaded centres and heavily shaded parts next to the tree row. When the simulated PE_i data was combined with the N supply from tree residues, the wider between-row tree spacings (6 or 8 m, with 1 or 2 m within-row spacing) appeared to be best. In general, to minimize shading of the crop, yet maintain a sufficient supply of organic matter and nitrogen in prunings, alley cropping tree arrangements of high between-row and low within-row spacings seem to be most promising. The east-west orientation seems to be more promising than north-south.     

The effects of between-row (alley widths) and within-row spacings ofGliricidia sepium on alley-cropped maize in Sierra Leone

The effects of spacings between hedgerows (alley widths) and the spacings of trees within hedgerows ofGliricidia sepium on growth and grain yield of maize were investigated at Senehun in southern Sierra Leone. Four between-row spacings (2, 4, 6 and 8 m) were combined with three within-row spacings (0.25, 0.50 and 1.00 m) in a split block design. Maize, at densities of 20,000, 40,000 and 53,333 plants ha^–1, was established in the alleys and also as pure crops. N, P and K fertilizers were applied to all plots before pruning of the trees began. When pruning started, only the pure maize plots received fertilizer; prunings from the hedgerows were returned to the appropriate alleys in the other plots.Plots with the highest maize populations consistently gave the best yields before pruning started, but lower populations gave improved yields after pruning. Yields of maize increased with increasing alley widths before the start of pruning, after which the narrower alleys of 2 and 4 m outyielded the wider ones by almost double, probably because of the large amount of nutrients applied in prunings. Lack of light limited grain yields before the start of pruning, when there was some shading by the hedgerows. Alleys of 2–4 m wide, planted no closer than 0.50 m within rows, resulted in more than twice the yields of maize than in the 8-m alleys planted at 0.25 m within rows, once the hedgerows were well established and were being managed.     

栽植密度对黄檗苗期生物量的影响

以黄檗3 a生幼树为材料,对不同栽植密度的生物量进行研究,结果表明:栽植当年和翌年秋季的成活率与保存率分别为92%和91%。不同株距树高、地径生长量均达到极显著差异(P0.01),树高、地径生长量从大到小顺序依次为株距0.5 m1 m0.3 m1.5 m;而不同行距对树高、地径生长量均无影响。采用一元生物量模型计算生物量,总生物量最大的种植密度为33 300株·hm~(-2)(株行距0.3 m×1.0 m),可产生干物质3 930.06 kg·hm~(-2);最小的为3 300株·hm~(-2)(株行距1.5 m×2.0 m),干物质仅为344.08 kg·hm~(-2)。     

Interactive effects of nitrogen and water availabilities on gas exchange and whole-plant carbon allocation in poplar

Cuttings of balsam spire hybrid poplar (Populus trichocarpa var. Hastata Henry x Populus balsamifera var. Michauxii (Dode) Farwell) were grown in sand culture and irrigated every 2 (W) or 10 (w) days with a solution containing either 3.0 (N) or 0.5 (n) mol nitrogen m(-3) for 90 days. Trees in the WN (control) and wn treatments had stable leaf nitrogen concentrations averaging 19.4 and 8.4 mg g(-1), respectively, over the course of the experiment. Trees in the Wn and wN treatments had a similar leaf nitrogen concentration, which increased from 12.0 to 15.8 mg g(-1) during the experiment. By the final harvest, mean stomatal conductances of trees in the wN and wn treatments were less than those of trees in the Wn and WN treatments (1.8 versus 4.6 mm s(-1)). Compared to the WN treatment, biomass at the final harvest was reduced by 61, 72 and 75% in the Wn, wN and wn treatments, respectively. At the final harvest, WN trees had a mean total leaf area of 4750 +/- 380 cm(2) tree(-1) and carried 164 +/- 8 leaves tree(-1) with a specific leaf area of 181 +/- 16 cm(2) g(-1), whereas Wn trees had a smaller mean total leaf area (1310 +/- 30 cm(2) tree(-1)), because of the production of fewer leaves (41 +/- 6) with a smaller specific leaf area (154 +/- 2 cm(2) g(-1)). A greater proportion of biomass was allocated to roots in Wn trees than in WN trees, but component nitrogen concentrations adjusted such that there was no Wn treatment effect on nitrogen allocation. Compared with WN trees, rates of photosynthesis and respiration per unit weight of tissue of Wn trees decreased by 28 and 31%, respectively, but the rate of photosynthesis per unit leaf nitrogen remained unaltered. The wN and Wn trees had similar leaf nitrogen concentrations; however, compared with the Wn treatment, the wN treatment decreased mean total leaf area (750 +/- 50 cm(2) tree(-1)), number of leaves per tree (29 +/- 2) and specific leaf area (140 +/- 6 cm(2) g(-1)), but increased the allocation of biomass and nitrogen to roots. Net photosynthetic rate per unit leaf nitrogen was 45% lower in the wN treatment than in the other treatments. Rates of net photosynthesis and respiration per unit weight of tissue were 48 and 33% less, respectively, in wN trees than in Wn trees.     

Biomass production and allometric above- and below-ground relations for young birch stands planted at four spacings on abandoned farmland

The objective of the study was to quantify above- and below-stumpbiomass of silver (Betula pendula Roth) and downy (Betula pubescensEhrh.) birches planted at four spacing intervals and growingon two soil types on an area of farmland. The 12-year-old bircheshad been grown at four spacings (1.3, 1.5, 1.8 and 2.6 m) ontwo sites: one on medium clay soil and the other on fine sandsoil. The dry weight of the stem, branches, leaves, stumps androots was estimated by drying and weighing sub-samples. Theprojected leaf area (PLA) m^–2 of trees, leaf area indexof stands and basic density (kg m^–3)of stems were alsoestimated. A significant greater dry weight of stem, branches,stump and roots and species and spacing for pendula birch werefound. The root length of silver birch was significantly greaterthan that for downy birch and for both species the root lengthwas greatest at the widest spacing (2.6 m). There was also asignificant difference between leaf weights of birch of thesame species growing on the two soil types. Significant differenceswere also found between PLA and species, and for both species,between PLA spacing. Basic density of stems was significantlydifferent between soil types. Equations for estimating the above-groundbiomass and root biomass from diameter at breast height weredeveloped for birches growing on fine sand and on medium claysoils. The total biomass production per hectare on fine sandwas higher for silver birch (19.9–65.9 tonnes ha^–1),than for downy birch (13.0–48.3 tonnes ha^–1). Onmedium clay soil, total biomass production for silver and downybirches was 30.8–52.8 and 16.8–42.8 tonnes ha^–1,respectively.     

Effect of pollarding frequency on biomass of Erythrina poeppigiana as a coffee shade tree

The use of pollarded Erythrina poeppigiana as shade tree in coffee plantations is apparently an old practice in Costa Rica. The tree is not native to this country but was introduced between late 19th and 20th century and was rapidly dispersed in the coffee and cacao areas. Currently, the Erythrina tree is widespread in the Turrialba Valley (elevation 600—1300m) and in the Central Valley (elevation 1200m) where the species is always associated with present or past coffee crops. Pollarding carried out by Costa Rican farmers constitutes a long dated and functional practice, hence the objective of this study was to evaluate the amount of biomass produced by pollarding of Erythrina poeppigiana used as shade in coffee crop planted at a density of 280 trees/hectare under different pollarding frequencies. Results showed that by pollarding once a year, 18,470 kg of dry matter per hectare are produced; with two pollardings per year 11,800 kg/ha are produced and with three pollarding per year 7,850 kg/ha are produced. The total amount of nitrogen removed is very similar for pollarding once and twice a year, but is lower for three times a year. The amount of nitrogen removed was approximately 230 kg/ha/year in the first two cases and 170 kg/ha/year in the last one.The above observations suggest that a considerable supply of nutrients exist in the systems with shade trees, when they are periodically pollarded.Finally some conclusions and follow up activities related to research on the species are suggested, such as higher biomass production techniques, appropriate planting practices, selection of genetic material, nutrient depletion when biomass is harvested, conversion of leaves to marketable feed sources (flour, pellets), alley cropping and green manure production and restoration of degraded areas and improductive savannas by planting large cuttings that would improve the soil by adding biomass and shade out undesirable grasses.This work is part of a Ph.D. Dissertation submitted to the Southeastern University, New Orleans, Lousiana by R.O. Russo.     

Shading effects of alley cropped Leucaena leucocephala on weed biomass and maize yield at Mtwapa,Coast Province,Kenya

Reductions of up to 90% in weed biomass was observed under Leucaena leucocephala alley cropping with varying tree between (2,4 and 8 m) and within-row spacing combinations when compared to crop-only control. An increase of 24 to 76 % in maize yields of alley cropped plots compared to the crop-only control was also recorded. The 2m alley widths closed canopy faster than the 4 and 8m, and hence realized highest weed biomass reduction during the short-fallow period between two cropping seasons. At the end of the short fallow period, substantial fuelwood (up to 8 t ha ^–1) was realized.     

Effects of understorey vegetation on tree establishment and growth in a silvopastoral system in northern Greece

The interaction between understorey plants and trees in a young silvopastoral system was studied in a sub-humid Mediterranean environment with cold winters in northern Greece. The experimental design was a split-split plot with three replications and included: three understorey treatments (grass, legume and control), two tree species (Acer pseudoplatanus and Pinus sylvestris) and two spacings (2.5×2.5 m and 3.5×3.5 m). The competition between herbaceous plants and trees resulted in significant differentiation in tree growth while their spacing did not produce any significant differences. The influence of trees on herbage yield either by plant species or spacing was not significant. During the early stages of establishment, a significant positive correlation was observed between sycamore growth and soil moisture, leaf weight, leaf area and leaf number as well as nitrogen and potassium concentration in leaves. Six years after planting the height increase of the sycamore trees was largest in the control treatment (415%), lowest in the grass treatment (134%) and intermediate (192%) in the legume treatment. Much higher was the increase obtained for the diameter, 161%, 207% and 536% respectively for the grass, legume and control treatments. The Scots pine trees grew faster than sycamore over the course of the experiment. Height increased by 397%, 351% and 400% and diameter by 518%, 443% and 683% respectively for the grass, legume and control treatments. This revised version was published online in June 2006 with corrections to the Cover Date.     

Biomass production and root distribution of Gmelina arborea under an agrisilviculture system in subhumid tropics of Central India

A study of an agrisilviculture system comprising Gmelina arborea and soybean (Glycine max) was conducted in the subhumid region of Central India. Above- and below-ground biomass production and distribution of coarse and fine roots were studied in 4-year-old G. arborea, planted at a spacing of 2 × 2 m, 2 × 3 m, 2 × 4 m and 2 × 5 m. The total biomass varied from 10.89 Mg ha^–1 to 3.65 Mg ha^–1 depending on the tree density. Among the different tree components, stemwood contributed maximum biomass (54.3–79.4%), followed by branches and leaves. Root distribution pattern showed that most of the coarse roots were distributed in the top 40 cm of soil, whereas fine roots were concentrated in the top 20 cm. Coarse root biomass decreased with an increase in spacing. The spread of roots was asymmetrical in trees planted at 2 × 2 m and 2 × 3 m spacings, while it was symmetrical in trees planted at wide spacings. No significant difference was observed in the fine root biomass in different stands. The root:shoot ratio increased with an increase in spacing. Crop (soybean) growth and productivity varied significantly and it increased with a decrease in tree density. Soybean yield varied between 1.5 Mg ha^–1 to 2.1 Mg ha^–1. The role of root architecture of G. arborea trees on productivity of crops under agri-silviculture system is discussed.     

Growth and quality response of Bilinga (Nauclea Diderrichii Merril.) to spacing

The effect of spacing on growth and tree bole quality was examined in a seven year old Bilinga (Nauclea diderrichii Merril.) stand. Four spacings ranging from 2.5 m to 5.0 m, were planted in a complete randomozed block design with three replications. Survival rate was significantly affected by spacing only at age 7. Diameter at breast height (DBH), crown ratio and number of knotty trees increased with wider spacing, while basal area and number of dead limbs increased inversely with spacing. Spacing did not seem to have any significant effect on total height growth. Trends in this study confirm some commonly held beliefs about spacing's effect on tree growth and quality. The results are still of limited application due to the juvenile age of the stand.     

Influence of different Taxodium ascendens stands on the open ranges and system performance in Jiangsu Province, China

This paper reveals the results of an eight-year experiment set up to identify the effects of four tree spacing configurations (1.5 × 4 m, 1.2 × 5 m, 2 × 3 m and 1.5 × 2 × 6 m) on open ranges and the relationship between open ranges and species coexistence in Taxodium ascendens-intercrop systems. Relative interrow open ranges and relative interplant open ranges were significantly different among four tree spacings. All relative open ranges (relative interrow open ranges, relative interplant open ranges, relative canopy open ranges and relative space-volumed open ranges) decreased with increasing age. The wider interrow or interplant spacings of trees offered wider (relative) interrow or interplant open ranges for intercrops. All of the tree spacings, except spacing 2 × 3 m, had negative relative interplant open ranges in nine-year-old stands, suggesting that the canopies between a plant and its neighbouring plant were in direct competition, and no interplant open range existed between two individual trees. Up to age nine, spacing 1.5 × 2 × 6 m showed no marked disadvantage in growth performance as compared to the other three spacings, but provided wider interrow open ranges for intercrops. Starting with tree stands of age four years, the yields of interplanted wheat (Triticum aestivum), soybean (Glyciae max) and rape (Brassica napus) decreased significantly as a result of the decrease of open ranges in tree stands. The yields of wheat, soybean and rape were lower in tree spacing 2 × 3 m than in other spacings. There was a positive correlation between wheat yields and canopy open ranges. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effect ofAcacia nilotica on mustard crop

Performance of mustard (Brassica campestris) crop cultivated underAcacia nilotica single tree, grown under irrigated and rainfed situations was studied for plant population and biomass production. Root behaviour of trees and mustard crop under tree canopy and open areas was also studied. Plant density and biomass production of mustard crop declined with increasing tree canopy depth towards tree trunk. There was a 64.58% reduction in biomass production under an irrigated field upto 4 m and cent percent up to 3 m in rainfed fields as compared to per unit mean yield data recorded at 21 m distance from the tree base in an open field. The plant density and biomass production data and root behaviour of both trees and crop showed a close relationship. The plant density and yield of crop tend to increase with the increase in depth of tree roots. Irrigating the mustard field enhances crop yield and plant density in the affected area as compared with the rainfed field.     

Estimating stem maintenance respiration rates of dissimilar balsam fir stands

Stem maintenance respiration rates were measured in five contrasting balsam fir (Abies balsamea (L.) Mill.) stands. At 15 degrees C, average respiration rates for individual stands ranged from 120 to 235 micro mol m(-3) s(-1) when expressed per unit of sapwood volume, from 0.80 to 1.80 micro mol m(-2) s(-1) when expressed per unit of stem surface area, and from 0.50 to 1.00 micro mol g(-1) s(-1) when expressed per unit of nitrogen in the living stem biomass, but differences among stands were not statistically significant. Coefficients of variation ranged from 50 to 100% within stands and were similar for all bases used to express respiration rates. Coefficients of determination for regressions between chamber flux and chamber values of sapwood volume, stem surface area and nitrogen content varied between stands and no one base was consistently higher than the other bases. We conclude that the bases for expressing stem respiration are equally useful. Respiration rates were more closely correlated to stem temperature observed approximately 2 h earlier than to current stem temperature. Among stands, annual stem maintenance respiration per hectare varied from 0.1 to 0.4 Mmol ha(-1) year(-1), primarily because of large differences in sapwood volumes per hectare. Annual stem maintenance respiration per unit of leaf area ranged from 3 to 6 mol m(-2) year(-1), increasing as sapwood volume per hectare increased.     

Changes in photosynthesis and leaf characteristics with tree height in five dipterocarp species in a tropical rain forest

Variations in leaf photosynthetic, morphological and biochemical properties with increasing plant height from seedlings to emergent trees were investigated in five dipterocarp species in a Malaysian tropical rain forest. Canopy openness increased significantly with tree height. Photosynthetic properties, such as photosynthetic capacity at light saturation, light compensation point, maximum rate of carboxylation and maximum rate of photosynthetic electron transport, all increased significantly with tree height. Leaf morphological and biochemical traits, such as leaf mass per area, palisade layer thickness, nitrogen concentration per unit area, chlorophyll concentration per unit dry mass and chlorophyll to nitrogen ratio, also changed significantly with tree height. Leaf properties had simple and significant relationships with tree height, with few intra- and interspecies differences. Our results therefore suggest that the photosynthetic capacity of dipterocarp trees depends on tree height, and that the trees adapt to the light environment by adjusting their leaf morphological and biochemical properties. These results should aid in developing models that can accurately estimate carbon dioxide flux and biomass production in tropical rain forests.     

六个杨树无性系苗木生长、生物量和光合作用的研究*

六个杨树无性系苗木（２年生根１年生于）叶面积、苗高和地径的季生长用Ｒｉｃｈａｒｄｓ方程模拟，根据模拟参数分别计算最大绝对生长速率，平均绝对生长速率，生长期和拐点。结果为：六个无性系之间在生长速率、拐点和生长期方面均有明显的差异；最好的欧美杨６４号生长季末的总生物量是赤峰杨３４号的六倍多；六个无性系的总生物量与它们的总叶面积、平均单个叶片的面积、叶面积的平均生长速率和每株树的日净光合总量等呈正相关，而与单位叶面积的日净光合总量呈负相关；总生物量生产依赖于叶面积、苗高、直径的生长速率及其生长期。     

Ailanthus triphysa at different densities and fertiliser regimes in Kerala, India: growth, yield, nutrient use efficiency and nutrient export through harvest

A split plot trial involving Ailanthus triphysa (ailanthus) at four spacings (3 m×1 m, 2 m×2 m, 3 m×2 m and 3 m×3 m) and four fertiliser regimes (0:0:0, 50:25:25, 100:50:50 and 150:75:75 kg ha⁻¹ per year N, P₂O₅, K₂O) was initiated in June 1991. Objectives included evaluating the growth and yield potential of ailanthus grown under differing density and fertiliser regimes and to estimate the nutrient export through harvest. Ninety-six randomly selected average-sized trees were felled at 8.8 years of age for assessment. Results show that height, diameter, stand leaf area index, biomass production and volume yield were greater in the 2 m×2 m spacing. Repeated application of fertilisers at 1.2, 2.25 and 5.25 years after planting had little effect on biomass and volume yields, presumably because of weed competition (despite periodic weed control), higher pest incidence (in the heavily fertilised plots) and/or moderately adequate soil nutrient levels. Regarding partitioning of tree biomass, stem wood represented the principal component (>70%), while foliage contributed the least (<7%). Conversely, foliar N, P and K concentrations were the highest, followed by branch wood, coarse roots and stem wood. Denser stands showed greater accumulation of N, P and K with higher potential for nutrient export through harvest. However, as the bole fraction accounted for only about 56–64% of the total nutrients removed, leaving other biomass components (foliage and branches) at the site will reduce the associated nutrient export. Wider spacings (3 m×2 m and 3 m×3 m) were more efficient in N and K use, but P use efficiency was higher in 2 m×2 m. Likewise, trees in the no fertiliser plots exhibited greatest N, P and K use efficiencies. Available soil P, K and organic C levels declined with increasing tree density, while repeated fertilisation increased nutrient concentrations. Soil pH and available P levels declined in comparison to the pre-treatment values.     

Comparative water use by dryland trees in Parklands in Senegal

Despite the clear evidence of competition for water between trees and crops, there have been very few studies comparing simultaneous water use by differing tree species in drylands. Comparative water use by dryland trees was measured in Senegal using heat balance gauges at the end of the wet season and in the dry season. Significant differences between tree species were found for maximum rates of water use per unit leaf area. Indigenous species may be better adapted to the dry environment than exotic species but the indigenous species Acacia seyal Del. used more water per unit leaf area than all other species. The exotic species Azadirachta indica Adr. Juss. consistently used less water per unit leaf area than most other species. There were significant differences in amounts of water used per unit leaf area by differing provenances of the same tree species. Water use in the dry season varied by a factor of three between two provenances of Acacia aneura F. Muell ex Benth. indicating potential to select provenances for drylands based on their water use characteristics. Absolute rates of water use as well as differences in sapflow between species were greatest when soils were moist suggesting that comparative sapflow studies will be most informative when carried out during the wet season. Water use rankings of the differing tree species were broadly maintained irrespective of season.This revised version was published online in November 2005 with corrections to the Cover Date.     

Intercropping Acacia albida with maize (Zea mays) and green gram (Phaseolus aureus) at Mtwapa,Coast Province,Kenya

Long-term agroforestry demonstrations/trials using Acacia albida and other nitrogen fixing multipurpose trees/shrubs were initiated in mid-1982 to assess soil and crop productivity at a coastal lowland site characterized by low soil fertility, weed problems and consequent poor crop yields. Growth performance (height and diameter at breast height, dbh) of Acacia albida under eight densities rotationally intercropped with maize (Zea mays) and green gram (Phaseolus aureus), crop grain yields, soil fertility changes and weed control were assessed for a 5-year period (May 1982 to March 1987). A parallel-row systematic spacing field layout was used. Intercropped Acacia albida mean hight and dbh were 140 and 24% respectively higher than tree-only controls by the fifth year. Growth rate was low during the first year but increased in subsequent years to mean height and dbh of 9 m and 10 cm respectively by March 1987. While differences in dbh were significant, those between stand heights were not. Crop yields, especially under higher tree densities, declined considerably due to unexpected shade which also caused significant reductions in weed biomass. Soil fertility levels remained unchanged during the experimental period relative to the initial status, and differences between the intercropped Acacia albida plots and the tree — or crop — only control appeared not to be significant. We conclude that an understanding of the mechanism regulating leaf fall/retention phenomena of Acacia albida is crucial towards determining the intercropping potentials of the species.     

Variations in productivity and wood properties of Amazonian tachi-branco trees planted at different spacings for bioenergy purposes

Tachi-branco(Tachigali vulgaris,L.F.Gomes da Silva H.C.Lima) is a leguminous tree species native to the Amazon rainforest that has drawn attention for its remarkably fast growth,a required trait for biomass/bioenergy plantations.In evaluations of biomass production and wood properties of T.vulgaris planted in homogeneous plantations at different spacings in the Amazonian state of Para,Brazil,biomass of 7-year-old trees was quantified for individual trees and the entire population.Wood was also sampled to assess properties relevant to bioenergy applications.The choice for spacing dimension for planting nonclonal T.vulgaris should consider whether the priority is greater productivity per tree,achieved with greater spacings(9.0 m~2 and 12.0 m~2),or productivity per area,achieved with closer spacings(6.0 m~2 and 7.5 m~2).Genetic variability of the T.vulgaris seed stand and/or high heritability of wood traits overcame the effect of different spacing on all morphological,physical,chemical and energetic properties of T.vulgaris wood.This species has moderate basic density when cultivated at spacings larger than 6 m~2 and net heating value above 7.95 MJ/kg,which is suitable for bioenergy purposes.The high variation in wood properties within tree spacing is strongly indicative of great potential for genetic breeding.The fast growth and the suitable moderate wood basic density confirm the outstanding potential of homogeneous plantations of T.vulgaris for providing wood for bioenergy.     

Biomass production, foliar and root characteristics and nutrient accumulation in young silver birch (Betula pendula Roth.) stand growing on abandoned agricultural land

The above-ground biomass and production, below-ground biomass, nutrient (NPK) accumulation, fine roots and foliar characteristics of a 8-year-old silver birch (Betula pendula) natural stand, growing on abandoned agricultural land in Estonia, were investigated. Total above-ground biomass and current annual production after eight growing seasons was 31.2 and 11.9 t DM ha⁻¹, respectively. The production of stems accounted for 62.4% and below-ground biomass accounted for 19.2% of the total biomass of the stand. Carbon sequestration in tree biomass reaches roughly 17.5 t C ha⁻¹ during the first 8 years. The biomass of the fine roots (d < 2 mm) was 1.7 ± 0.2 t DM ha⁻¹ and 76.2% of it was located in the 20 cm topsoil layer. The leaf area index (LAI) of the birch stand was estimated as 3.7 m² m⁻² and specific leaf area (SLA) 15.0 ± 0.1 m² kg⁻¹. The impact of the crown layer on SLA was significant as the leaves are markedly thicker in the upper part of the crown compared with the lower part. The short-root specific area (SRA) in the 30 cm topsoil was 182.9 ± 9.5 m² kg⁻¹, specific root length (SRL), root tissue density (RTD) and the number of short-root tips (>95% ectomycorrhizal) per dry mass unit of short roots were 145.3 ± 8.6 m g⁻¹, 58.6 ± 3.0 kg m⁻³ and 103.7 ± 5.5 tips mg⁻¹, respectively. In August the amount of nitrogen, phosphorus and potassium, accumulated in above ground biomass, was 192.6, 25.0 and 56.6 kg ha⁻¹, respectively. The annual flux of N and P retranslocation from the leaves to the other tree parts was 57.2 and 3.7 kg ha⁻¹ yr⁻¹ (55 and 27%), respectively, of which 29.1 kg ha⁻¹ N and 2.8 kg ha⁻¹ P were accumulated in the above-ground part of the stand.