Relative Importance of Environmental Stress and Herbivory in Reducing Litter Fall in a Semiarid Woodland

We examined the impact of soil stress (low water and nutrient availabilities) and two keystone insect herbivores on pinyon pine (Pinus edulis) needle litterfall. We compared trees growing on two distinct soil types: volcanic cinders, which exhibit pronounced water and nutrient limitation, and sandy-loam soils, which have higher water-storage capacity and nutrient availability. Using two long-term herbivore removal experiments (15 and 18 years, respectively), we also examined the effects of the pinyon needle scale (Matsucoccus acalyptus, which attacks juvenile trees) and the stem-boring moth (Dioryctria albovittella, which attacks mature trees) on pinyon litterfall. These herbivores reach high densities on cinder soils but are absent or occur at much lower levels on sandy-loam soils. Four years of litterfall measurements showed four major patterns. First, independent of herbivory, needle litterfall was 20% lower under trees on high-stress cinder soils than on sandy-loam soils. Second, in agreement with the negative impact of scales on tree growth (that is, a 30% decline in stem growth), trees with scale infestations had 25% lower litterfall rates than trees resistant to scale; however, 15 years of scale-insect removal did not significantly increase needle litterfall. This implies possible intrinsic differences in litter production between scale-resistant and scale-susceptible trees. Third, in contrast with significant negative effects of moth herbivory on tree growth (that is, a 27% decline in stem growth), moth herbivory had no effect on needle litterfall. This, along with increased stem density in moth-susceptible trees, may be evidence of compensatory production. Fourth, there were strong year by soil type and year by scale herbivory interactions, such that in some years the effect on litterfall can be obscured or reversed by some other factor. In summary, soil stress has a strong and predictable effect on needle litterfall, whereas the relationship between insect herbivory and needle litterfall is weaker and depends on the individual herbivore. These effects, however, are mediated by other environmental factors that have considerable annual variation.     

Nutrient requirements of exotic tree species in Zimbabwe

Pot and field experiments were conducted in the greenhouse and at three field sites (Marondera, Domboshawa and Makoholi) in Zimbabwe to examine the effects of soils and fertilizers on nutrient uptake and growth of 6 exotic tree species (Eucalyptus camaldulensis, E. grandis, E. tereticornis, Leucaena leucocephala, Casuarina cunninghamiana, and Acacia holosericea). Plant growth, N and P contents of all species were increased by the application of N, P, K and micronutrient fertilizers. The effect of individual nutrients (N, P, K and micronutrients) and their combination on Eucalyptus species was further investigated in a pot experiment using soil from Domboshawa. Eucalyptus species responded only to N application and no significant interactions were found between N and the other elements. Nutrient uptake results showed that E. camaldulensis and E. tereticornis removed more cations than the N-fixing trees but only in the fertilized treatments. L. leucocephala and C. cunninghamiana were higher in P, but no clear trends were observed for N. Plant growth and nutrient uptake by E. camaldulensis. C. cunninghamiana and A. holosericea were assessed in the field at the three sites. Plant species grown in the Marondera site had greater height and diameter at breast height (DBH) than those in the two other sites. These results followed trends in soil nutrient contents. The analysis of foliage revealed differences in the nutrient concentration of leaves from different trees, but no effect of site was found. The interrelationships between plant characteristics, soil and foliage nutrients were examined. In a pot experiment, mineral N was the only variable correlated with growth response and nutrient uptake, while in the field the only significant correlation was obtained with soil pH.     

Nutrition ofPinus caribaea in its native savanna habitat

Summary Fertility levels in soils beneathPinus caribaea trees were examined in the Mountain Pine Ridge savannas, Belize, where fire control has precipitated the development of pine woodland. Slight surface soil enrichment was recorded beneath pine canopies, but to levels well below those found beneath associated hardwoods. Estimates of total nutrient pools beneath trees showed modest cation accumulation beneath a 73 year old tree but some defecits in Ca and Mg beneath a 24 year old tree. A tap root cutting experiment on trees of the same species revealed no significant declines in foliar nutrient levels after 19 months. It is concluded that no pronounced long-term deterioration in soil fertility levels is developing beneath stands ofP. caribaea in the savanna, although some temporary nutrient declines may exist beneath young pine stands. Atmospheric inputs are the most likely source of nutrient accretion and it is suggested that the establishment of hardwood associates with pine may enhance the rates of nutrient capture from this source.     

三种苦苣苔对石灰土和红壤的适应性分析

为探讨苦苣苔科植物对其岩溶生境的适应性,该研究选取黄花牛耳朵(Primulina lutea)、紫花报春苣苔(Pri.purpurea)和桂林蛛毛苣苔(Paraboea guilinensis)三种苦苣苔科植物,将其栽种在石灰土及红壤两种不同类型的土壤中,观测记录其生长性状并对其叶片元素含量进行测定和比较。植株采集过程中,同时采集自然生境中三种苦苣苔科植物叶片及取样植物基部土壤,并对叶片及土壤元素的含量进行测定,作为今后苗圃试验的参照。结果表明:三种苦苣苔科植物在两种土壤上的生长状况及适应性具有差异,其在石灰土上生长良好,在红壤上生长较差;在两种不同土壤中,除N外,桂林蛛毛苣苔的叶片其他元素(P、K、Mn、Mg、Ca、Zn、Cu)差异极显著(P0.01);除P外,紫花报春苣苔的叶片其他元素(N、K、Mn、Mg、Ca、Zn、Cu)差异极显著(P0.01);除N、Cu、Ca外,黄花牛耳朵的叶片元素(P、K、Mn、Mg、Zn)差异极显著(P0.01);三种植物的叶片元素比值,除少数值没有差异外,大部分指标差异都极显著;对叶片元素与栽培土壤元素的相关性分析,发现植物叶片Mn元素与土壤中N、Ca、Mg、Zn、Mn、有机质含量等呈正相关,土壤P元素与叶片中N、P元素呈正相关,而与叶片中Zn元素呈负相关关系。在其他栽培条件一致的条件下,土壤因素及物种差别是造成黄花牛耳朵、紫花报春苣苔和桂林蛛毛苣苔适应性产生差异的主要原因。     

Bioavailability index for quantitative evaluation of plant availability of extractable soil trace elements

A new index, Bioavailability Index (BI) and the corresponding experimental method were developed for quantitative evaluation of bioavailability of the extractable soil trace elements. Soils were first treated with various extractants (DTPA, HCl, NH₂OH·HCl+HCl) separately to remove the extractable elements. The soils after extraction were washed with deionised water to eliminate the extractant and its pH was adjusted with Ca0 and finally restored to its original pH level. Wheat (Triticum aestivum L.) and rape (Brassica chinensis) were planted in the untreated and treated soils for 8 weeks. The concentrations of the trace elements in plants were determined after harvest. Nutrient accumulation by plants is significantly reduced due to removal of extractable trace elements from the soil. BI of the extractable fraction was proportional to the ratio of plant accumulation reduction to trace element extractability. In the present study, BI value of the total content of soil trace elements was designated as 1. Though only a minor fraction of the total soil nutrient, generally less than 5%, was removed by DTPA, the nutrient accumulation by plants, especially for wheat, was reduced greatly, leading to relatively large BI values. For wheat, the average BI values of the eight nutrients Cu, Mn, Zn, Ni, Co, Pb, Cr, and V were found to be 22.7, 17.6 and 17.4 for the three testing soils, and for rape, the corresponding values of 8.9, 10.0 and 11.0 were obtained, indicating that the DTPA-extractable elements represent the highly available fraction of the total content. The BI values for HCl-extractable elements were much lower compared with those for DTPA. For wheat, the average BI values for the three soils are 2.0, 1.9 and 2.4, and for rape, the corresponding values are 4.8, 4.1 and 3.7. The high availability of DTPA-extractable trace elements and relatively low availability of HCl-extractable trace elements highlight the significant role that chelation action might play in plant nutrient acquisition. The different responses of wheat and rape to the soils previously subjected to the same extraction procedure could be explained by their genotypical differences in sensitivity to nutrient deficiencies. The quantitative nature of BI makes it valuable in the study of nutrient bioavailability and plant accumulation mechanisms.     

大仔买麻藤在两种栽培环境下叶片的营养成分比较

为探讨栽培环境对大仔买麻藤(Gnetum montanum Markgr.f.megalocarpa Markgr.)叶片营养成分的影响,将采自云南省景东县的大仔买麻藤分别在北京和昆明的温室内培养,对其1年生幼苗叶片的营养成分进行研究。结果表明,不同栽培环境对大仔买麻藤叶片的大部分营养成分无显著影响;叶片总淀粉、总糖、黄酮和生物碱含量无显著差异;但在二糖、莰菲醇、部分氨基酸和某些元素含量上有显著差异;并且大部分的元素含量受土壤的影响显著。因此,不同栽培环境对大仔买麻藤叶片营养成分的影响不大。     

Local adaptation to serpentine soils in Pinus ponderosa

Local adaptation to serpentine soils is studied using both transplant experiments and molecular genetic techniques. In long-lived species, such as pines, it is unclear how soon after germination local adaptation becomes detectable. Here I present results of a 36-year reciprocal transplant experiment using Pinus ponderosa, along with allozyme analyses from the same trees. Using a repeated measures analysis of variance, there is evidence for adaptation to serpentine soils; however, significant differences between source soil types do not become apparent until 20 years after the start of the experiment. Analysis of allozyme data showed no evidence for differentiation between the serpentine and non-serpentine populations. Comparing the performance of families over the course of the experiment found that there was little correlation between performance after 1 or 4 years of growth in the field and performance after 36 years. This suggests that short-term transplant experiments may not provide definitive evidence for adaptation to serpentine soils. A literature survey of all transplant studies using pine species growing on and off of serpentine soils found that studies that lasted fewer than 2 years showed no evidence for adaptation. However, in the two experiments (this one included) that lasted more than 2 years, both showed evidence for adaptation to serpentine soils. More long-term experiments are required to validate these results.     

Characterization of the tolerance to iron chlorosis in different peach rootstocks grown in nutrient solution

Iron chlorosis is an important problem in peach trees, but differences exist between peach rootstocks in their tolerance to Fe chlorosis in calcareous soils. The purpose of this investigation was to characterize the tolerance of different rootstocks to Fe chlorosis induced by bicarbonate in nutrient solution. The rootstocks studied included peach (Nemaguard), plums (Brompton, San Julian A and Puebla de Soto 101) and almond × peach hybrids (Adafuel and GF677). Young plants obtained from rooted cuttings or from in vitro culture techniques were grown individually, under controlled conditions, in flasks with 700 mL of aerated nutrient solution low in iron and with or without 10 mM bicarbonate or 10 mM phosphate. Susceptiblity to bicarbonate-induced chlorosis was inversely correlated with both the Fe content in young leaves and the reducing capacity of roots, but not with the phosphorus content in young leaves. The plum Puebla de Soto 101 and the hybrid GF677 showed the lowest degree of chlorosis and the highest reducing capacity. Phosphate did not induce chlorosis.     

Effects of Soil Burn Severity on Post-Fire Tree Recruitment in Boreal Forest

Fire, which is the dominant disturbance in the boreal forest, creates substantial heterogeneity in soil burn severity at patch and landscape scales. We present results from five field experiments in Yukon Territory, Canada, and Alaska, USA that document the effects of soil burn severity on the germination and establishment of four common boreal trees: Picea glauca, Picea mariana, Pinus contorta subsp. latifolia, and Populus tremuloides. Burn severity had strong positive effects on seed germination and net seedling establishment after 3 years. Growth of transplanted seedlings was also significantly higher on severely burned soils. Our data and a synthesis of the literature indicated a consistent, steep decline in conifer establishment on organic soils at depths greater than 2.5 cm. A meta-analysis of seedling responses found no difference in the magnitude of severity effects on germination versus net establishment. There were, however, significant differences in establishment but not germination responses among deciduous trees, spruce, and pine, suggesting that small-seeded species experience greater mortality on lightly burned, organic soils than large-seeded species. Together, our analyses indicate that variations in burn severity can influence multiple aspects of forest stand structure, by affecting the density and composition of tree seedlings that establish after fire. These effects are predicted to be most important in moderately-drained forest stands, where a high potential variability in soil burn severity is coupled with strong severity effects on tree recruitment.     

Nutrient status of NPK-treated coffee plots

Summary The chemical properties of 128 soil samples collected from the surface and subsurface layers of 64 NPK treated coffee (Coffea canephora P.) plots were analysed. The ensuing data were compared with those from other coffee, cocoa and tropical soils of West Africa and Brazil. After seven years of NPK application to the coffee soil in Southwestern Nigeria, the soil was considered deficient in base elements especially Ca and Mg, N, P and K were considered fairly marginal. NPK treatments were found to effect statistically significant differences in the nutrient contents of coffee plots.     

Increased moth herbivory associated with environmental stress of pinyon pine at local and regional levels

 Using 6 years of observational and experimental data, we examined the hypothesis that water and nutrient stress increase the susceptibility of pinyon pine (Pinus edulis) to the stem- and cone-boring moth (Dioryctria albovittella). At two geographic levels, a local scale of 550 km² and a regional scale of 10,000 km², moth herbivory was strongly correlated with an edaphic stress gradient. At a local scale, from the cinder soils of Sunset Crater to nearby sandy-loam soils, nine of ten soil macro- and micronutrients, and soil water content were lowest in cinder-dominated soils. Herbivore damage was six times greater on trees growing in the most water and nutrient deficient site at Sunset Crater compared to sites with well-developed soils. Percentage silt-clay content of soil, which was highly positively correlated with soil nutrient and soil moisture at a local scale, accounted for 56% of the variation in herbivory at a regional scale among 22 sites. Within and across sites, increased stem resin flow was positively associated with reduced moth attack. On the basis of moth distribution across a stress gradient, we predicted that pinyons growing in highly stressful environments would show increased resistance to herbivores if supplemented with water and/or nutrients. We conducted a 6-year experiment at a high-stress site where individual trees received water only, fertilizer only, and water + fertilizer. Relative to control trees, stem growth and resin flow increased in all three treatments, but only significantly in the water + fertilizer treatment. Although there was no significant difference in herbivore damage among these three treatments, there was an overall reduction in herbivore damage on all treatment trees combined, compared to control trees. This experiment suggests that release from stress leads to increased resistance to insect attack and is consistent with our observational data. While other studies have predicted that short-term stress will result in herbivore outbreaks, our studies extend this prediction to chronically stressed host populations. Finally, while flush-feeders are not predicted to respond positively to stressed host plants, we found a positive association between herbivore attack and stressed pinyon populations. Received: 25 December 1995 / Accepted: 15 August 1996     

Alternate bearing influences annual nutrient consumption and the total nutrient content of mature pistachio trees

The influence of alternate bearing on nutrient utilization and total tree nutrient content was investigated in mature pistachio (Pistacia vera L. cv Kerman trees). Removal of N, P and Zn in fruit and abscised leaves of cropping (‘on’) trees averaged 5, 6, and 2 times, respectively, the removal in abscised leaflets of the non-fruiting, ‘off’ year trees. One hundred and thirty-five kg N, 131 kg K, 86 kg Ca, 39 kg Mg and 18 kg P per hectare were removed in fruits and abscised leaves in ‘on’ year trees. Tree nutrient contents and, presumably, the size of nutrient storage pools in dormant trees varied between ‘on’ and ‘off’ years. There was 22% and 14% more N and P, respectively, in dormant trees following ‘off’ than ‘on’ years. The greater N and P accumulation in ‘off’ year trees is depleted in support of the large fruit demand for N and P during ‘on’ years. In contrast to N and P, there was greater K and Ca accumulation in perennial tree parts during ‘on’ years than during ‘off’ years. The greater K accumulation in perennial tree parts and approximately 30% greater removal of K in annual organs during ‘on’ than ‘off’ years suggests that K uptake could be 4 times higher in ‘on’ year trees than in (non-cropping), ‘off’ year trees.     

雷州半岛桉树人工林凋落物量和养分循环研究

为了解雷州半岛地区桉树人工林凋落物量和养分归还特征,对不同林龄人工林凋落物量和养分动态连续12个月进行监测。结果表明,桉树人工林的凋落物总量为5 a9 a7 a,5 a生桉树人工林的凋落物总量显著高于7和9 a生林分,且7和9 a生林分间没有显著差异。不同林龄桉树人工林凋落物总量具有明显的季节变化,均呈双峰型,峰值出现在雨季初期和末期。林分结构因子同凋落物量相关性不显著,但气候因子中的月均温与凋落物量存在显著正相关关系。不同林龄桉树人工林的养分年循环量由凋落物量和凋落物养分含量共同决定,氮、磷、钾元素的养分年归还总量为9 a7 a5 a,且彼此间达到显著差异。凋落物养分元素归还高峰期集中在雨季前后,以冬季养分归还量较低。因此,为增加桉树人工林产量且利于人工林地力恢复,桉树人工林应适当增加种植年限,且采伐季节选择在冬季。     

Nutrient accumulation byPinus caribaea in its native savanna habitat

Summary Concentrations of P, N, K, Ca, and Mg in above-ground tissues ofP. caribaea were sampled in the species'native savanna habitat. Concentrations were relatively low, but some evidence of higher consumption of K and Ca was found in trees grown on more fertile soils. Regressions were developed to predict the quantities of nutrients sequestered in above-ground tree tissues, and estimates made of the quantities stored in above-ground stands of this species in its native habitat and in several plantations elsewhere. Estimates were also made of the nutrient removals to be expected by harvesting these stands in different ways. Nutrient quantities stored in stands generally exceed those extractable from savanna surface soils, and it is suggested that inputs from the atmosphere are the most probable alternate nutrient source. A comparison of these inputs for tropical areas with the quantities required for stand growth in the savanna, and harvesting removals, suggests that an adequate supply of all elements except P exists, provided that capture by pine is effective. However, atmospheric inputs generally fall below the storage and harvest removal rates for fast growing exotic plantations of this species suggesting that multiple rotations of these plantations at current growth rates may not be feasible without artifical fertilization.     

Mycorrhizal inoculation in neem (Azadirachta indica) enhances azadirachtin content in seed kernels

In view of the high mycorrhizal dependency of neem trees (Azadirachta indica), an experiment was conducted to study if Arbuscular Mycorrhizal (AM) inoculation can enhance the azadirachtin content in seed kernels of trees grown in the field. Azadirachtin is an important active ingredient in neem seed kernels based on which a large biopesticide industry has emerged in India and few countries in Europe and the USA. Inoculation of neem seedlings in the nursery with Glomus fasciculatum and Glomus mosseae resulted in increased height, dry weight, root colonization and phosphorus (P) content. In a separate experiment, field-grown neem plants inoculated in the nursery and during transplantation with Glomus fasciculatum were evaluated after 5 years. No significant differences were found in the tree height, girth at breast height (GBH) and fruit yield but oil percentage, total triterpenoids and azadirachtin content in kernels increased significantly as a result of AM inoculation. A similar enhancement in azadirachtin was noted with P application. These results open up possibilities of producing quality neem seed with high bioactive ingredients through AM inoculation.     

Differential host utilization by two parasites in a population of ponderosa pine

We compared phloem characteristics of individual Pinus ponderosa attacked by the dwarf-mistletoe Arceuthobium vaginatum (Viscaceae) or by the beetle Dendroctonus ponderosae (Scolytidae) or by neither species. We quantified total nonstructural carbohydrates and a broad range of chemical elements for these three categories of trees. There were significant differences between trees parasitized by Arceuthobium, trees parasitized by Dendroctonus, and non-infected trees. Discriminant function analysis of trees attacked by either Arceuthobium or Dendroctonus correctly predicted group membership for 59 of 60 trees tested. Some of the differences detected may be induced, but many probably are not. Given that the accumulation of certain elements and compounds is under genetic control, and that both parasites often cause severe reductions in fitness, including death of their host, our results suggest that the two parasites may generate diversifying selection in ponderosa pine populations.     

Infidelity of leafcutting ants to host plants: resource heterogeneity or defense induction?

Summary Leafcutting ants have strong among- and within-plant preferences, and generally abandon plants long before they are completely defoliated. Two tropical deciduous forest tree species preferred by the leafcutting ant Atta colombica were studied to determine how variation in resource quality affects ant selectivity and partial defoliation of plants. Significant differences in palatability and leaf characteristics of Spondias mombin and Bursera simaruba were found among trees and among leaf types within trees, but not among branches within trees. No short-term responses to experimental defoliation of up to 50% of total canopy were found in either species. Leaf nutrient and poisture content were positively correlated, and phenolic content negatively correlated, with the palatability of Spondias mombin, a species containing hydrolyzable tannins. Leaf moisture and phenolic content were both positively correlated with the palatability of Bursera simaruba, which contains predominantly condensed tannins. The results suggest that variation in leaf quality among and within plants is at least a partial explanation for ant selectivity and partial defoliation of preferred species. There is no evidence that rapidly induced changes in plant chemistry affect ant decisions to abandon these plants. Instead, it appears likely that ants abandon plants once high-quality leaf patches are exhausted. Quantitative variation in leaf nutrients, moisture, and secondary chemicals all appear to contribute to ant preferences for individuals and tissues of highly palatable plants.     

An analysis of mineral uptake in apple rootstock seedlings

Summary Eight families from biparental crosses of apple rootstocks and 12 families from open pollinated Malus spp. were analysed in 2 years for N, P, K, Ca and Mg content of the foliage. Highly significant differences were found between the families for all elements. There were no significant differences between the means of the biparental group and the open pollinated group. Ca and K content were significantly more variable in the open pollinated families compared with the biparental families. It is suggested that this increased variation could prove useful in breeding for efficiency of mineral uptake by apple rootstocks.     

Influence of the wood ant, Formica polyctena, on soil nutrient and the spruce tree growth

The effect of Formica polyctena ant nests on the distribution of soil nutrients, soil pH and the growth of Norway spruce trees was studied in the southern part of the Czech Republic. Soil nutrient content (exchangeable P, N, K and pH) and growth of mature spruce trees were measured at four distances from the nearest ant hill (0–1, 3–5, 10–50 and >200 m). Trees at all distances were visited by ants, except for those >200 m from the nearest nest. Soil pH and of P, K and NO₃ concentrations were higher near ant nests (<1 m), and pH and K at distances of 3–5 m, when compared with distances of 10–50 and >200 m from the nests, where no significant differences in these variables were detected. In contrast, tree ring analyses (1974–2004) showed that trees >200 m from the ant nests grew significantly faster than trees at other distances, followed by trees within 1 m of the nests. No growth differences were found between the growth of trees at 3–5 and 10–50 m from ant nests. We postulate that nutrient and carbohydrate removal of honeydew collected by ant‐tended aphids are slowing growth of tree. However, trees may partly compensate for this depletion by having access to a larger supply of soil nutrients near ant nests.     

Carbon, nitrogen and phosphorus in volcanic soils following afforestation with native birch (Betula pubescens) and introduced larch (Larix sibirica) in Iceland

Afforestation has become an important tool for soil protection and land reclamation in Iceland. Nevertheless, the harsh climate and degraded soils are growth-limiting for trees, and little is know about changes in soil nutrients in maturing forests planted on the volcanic soils. In the present chronosequence study, changes in C, N and total P in soil (0–10 and 10–20 cm depth) and C and N in foliar tissue were investigated in stands of native Downy birch (Betula pubescens Enrh.) and the in Iceland introduced Siberian larch (Larix sibirica Ledeb.). The forest stands were between 14 and 97 years old and were established on heath land that had been treeless for centuries. Soils were Andosols derived from basaltic material and rhyolitic volcanic ash. A significant effect of tree species was only found for the N content in foliar tissue. Foliar N concentrations were significantly higher and foliar C/N ratios significantly lower in larch needles than in birch leaves. There was no effect of stand age. Changes in soil C and the soil nutrient status with time after afforestation were little significant. Soil C concentrations in 0–10 cm depth in forest stands older than 30 years were significantly higher than in heath land and forest stands younger than 30 years. This was attributed to a slow accumulation of organic matter. Soil N concentrations and soil P_tot were not affected by stand age. Nutrient pools in the two soil layers were calculated for an average weight of soil material (400 Mg soil ha⁻¹ in 0–10 cm depth and 600 Mg soil ha⁻¹ in 10–20 cm depth, respectively). Soil nutrient pools did not change significantly with time. Soil C pools were in average 23.6 Mg ha⁻¹ in the upper soil layer and 16.9 Mg ha⁻¹ in the lower soil layer. The highest annual increase in soil C under forest compared to heath land was 0.23 Mg C ha⁻¹ year⁻¹ in 0–10 cm depth calculated for the 53-year-old larch stand. Soil N pools were in average 1.0 Mg N ha⁻¹ in both soil layers and did not decrease with time despite a low N deposition and the uptake and accumulation of N in biomass of the growing trees. Soil P_tot pools were in average 220 and 320 kg P ha⁻¹ in the upper and lower soil layer, respectively. It was assumed that mycorrhizal fungi present in the stands had an influence on the availability of N and P to the trees. Responsible Editor: Hans Lambers.