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

Effects of irrigation and liquid fertilization on fine root (< 1 mm) production and longevity, and fine root (< 0.5-2 mm) biomass were studied in a Norway spruce (Picea abies (L.) Karst.) stand in northern Sweden. Fine root length production and longevity were measured by the minirhizotron technique at 0-10 cm depth in the following treatments: irrigation (I), liquid fertilization (IL) and control (C). Standing root biomass and root length density (RLD) were studied in the litter-fermented humus (LFH) layer and at depths of 0-10, 10-20 and 20-30 cm using soil cores in solid fertilized (F) and C plots. Minirhizotrons were installed in October 1994 and measurements recorded monthly from July to September 1995 and during the growing season in 1996. Soil cores were sampled in 1996. Fine root production increased significantly in IL plots compared with C plots, but the I treatment did not increase root production. Root mortality increased significantly in IL plots compared with C plots. Fine root longevity in IL plots was significantly lower compared with C and I plots. No significant difference was found between longevity of fine roots in I and C plots. Compared with C, F treatment increased fine root biomass in the LFH and mineral soil layers, and increased the amount of fine roots in mineral soil layers relative to the LFH layer. Furthermore, F increased RLD and the number of mycorrhizal root tips significantly.     

Effects of nutrition and soil water availability on water use in a Norway spruce stand

We investigated effects of nutrition and soil water availability on sap flux density, transpiration per unit leaf area (EL), and canopy stomatal conductance (GS) of Norway spruce (Picea abies L. (Karst.)) in northern Sweden during the 1996 growing season. Our objectives were to determine (1) if artificially imposed drought (65% rain diversion) reduces soil water sufficiently to cause physiological limitations to whole-tree and plot-scale water transport, and (2) whether increased capacity for water transport resulting from fertilization-induced increases in leaf (> 3-fold) and sapwood areas (> 2.3-fold) deplete soil water sufficiently to cause a negative feedback on GS and EL. We monitored soil water content (theta) and soil water potential (PsiS) in control (C), drought (D), fertilized (F) and irrigated + fertilized (IL) treatment plots, along with site meteorological conditions. Ten trees per plot were monitored for sap flow. Although there were significant treatment differences in mean daily EL (C > D > F; P < 0.01) and GS (C > D > F; P < 0.05), variation in absolute magnitudes was small. Therefore, transpiration differences on a unit ground area basis (EC) were nearly proportional to leaf area differences. Precipitation was well distributed throughout the study period and so PsiS remained high, except during short dry periods in Plot F when it declined rapidly. Thus, although soil water was not limiting to GS, EL or EC when precipitation was uniformly distributed throughout the growing season, we cannot conclude that water availability would not limit GS in fertilized stands if the seasonal distribution of precipitation were altered.     

Effects of slash retention and wood ash addition on fine root biomass and production and fungal mycelium in a Norway spruce stand in SW Sweden

In the study reported here we examined the short-term effects (1–3 years) of slash retention (SR) and the long-term effects (13–15 years) of wood-ash application (A) on fine roots and mycorrhizae in a 40-year-old Norway spruce forest in southwest Sweden. Soil cores were used to obtain estimates of the biomass (g m⁻²) of roots in three diameter classes (<0.5, 0.5–1 and 1–2 mm), root length density (RLD), specific root length (SRL) and mycorrhizal root tip density (RTD). Fine root (<1 mm) length production and mortality, and mycelium production, were estimated using minirhizotron and mesh bag techniques, respectively. Compared with the control plots (C), the biomass of fine roots in diameter classes <0.5 mm and 0.5–1 mm was significantly higher in A plots, but lower in SR plots. In addition, RLD was significantly lower in the humus layer of SR plots than in the humus layers of C and A plots, but not in the other layers. None of the treatments affected the SRL. In all soil layers, the SR treatment resulted in significant reductions in the number of ectomycorrhizal root tips, and the mycelia production of fungi in mesh bags, relative to the C treatment, but the C and A treatments induced no significant changes in these variables. Fine root length production in the C, A and SR plots amounted to 94, 87 and 70 mm tube⁻¹ during the 2003 growing season, respectively. Fine root mortality in treated plots did not change over the course of the study. We suggest that leaving logging residues on fertile sites may result in nitrogen mineralisation, which may in turn induce reductions in root biomass, and both root and mycelium production, and consequently affect nutrient uptake and the accumulation of organic carbon in soil derived from roots and mycorrhizae.     

Fine root biomass in relation to site and stand characteristics in Norway spruce and Scots pine stands

Variations in fine root biomass of trees and understory in 16 stands throughout Finland were examined and relationships to site and stand characteristics determined. Norway spruce fine root biomass varied between 184 and 370 g m(-2), and that of Scots pine ranged between 149 and 386 g m(-2). In northern Finland, understory roots and rhizomes (< 2 mm diameter) accounted for up to 50% of the stand total fine root biomass. Therefore, the fine root biomass of trees plus understory was larger in northern Finland in stands of both tree species, resulting in a negative relationship between fine root biomass and the temperature sum and a positive relationship between fine root biomass and the carbon:nitrogen ratio of the soil organic layer. The foliage:fine root ratio varied between 2.1 and 6.4 for Norway spruce and between 0.8 and 2.2 for Scots pine. The ratio decreased for both Norway spruce and Scots pine from south to north, as well as from fertile to more infertile site types. The foliage:fine root ratio of Norway spruce was related to basal area and stem surface area. The strong positive correlations of these three parameters with fine root nitrogen concentration implies that more fine roots are needed to maintain a certain amount of foliage when nutrient availability is low. No significant relationships were found between stand parameters and fine root biomass at the stand level, but the relationships considerably improved when both fine root biomass and stand parameters were calculated for the mean tree in the stand. When the northern and southern sites were analyzed separately, fine root biomass per tree of both species was significantly correlated with basal area and stem surface area per tree. Basal area, stem surface area and stand density can be estimated accurately and easily. Thus, our results may have value in predicting fine root biomass at the tree and stand level in boreal Norway spruce and Scots pine forests.     

Potential production of Norway spruce in Sweden

In this paper, the potential stem wood production for Norway spruce was estimated for different regions in Sweden. This was done by using basic physiological relationships of intercepted radiation versus biomass production and knowledge of how a water deficit reduces the potential production, derived from results of field experiments on nutrient optimisation. To scale these relationships up to regional and national levels, data of incident radiation and humidity during the growing season for all of Sweden were used. The figures for potential and attainable production indicate that the temperate to boreal climate allows considerably higher production than the current production, if availability of water and nutrients is non-limiting. In northern Sweden, the attainable production is ca. 300% higher than the current production and in southern Sweden, the yield can be increased by ca. 100%. In absolute numbers, as a mean for a whole rotation period, it is possible to achieve an annual stem-wood dry mass production of 7–9 Mg ha⁻¹ in southern and central and ca. 5–6 Mg in northern Sweden. This increased production would mean that rotation periods can be shorter than they are now by 20–30 years in southern Sweden and by ca. 50–60 years in northern Sweden.     

Stand age and fine root biomass, distribution and morphology in a Norway spruce chronosequence in southeast Norway

We assessed the influence of stand age on fine root biomass and morphology of trees and understory vegetation in 10-, 30-, 60- and 120-year-old Norway spruce stands growing in sandy soil in southeast Norway. Fine root (< 1, 1-2 and 2-5 mm in diameter) biomass of trees and understory vegetation (< 2 mm in diameter) was sampled by soil coring to a depth of 60 cm. Fine root morphological characteristics, such as specific root length (SRL), root length density (RLD), root surface area (RSA), root tip number and branching frequency (per unit root length or mass), were determined based on digitized root data. Fine root biomass and morphological characteristics related to biomass (RLD and RSA) followed the same tendency with chronosequence and were significantly higher in the 30-year-old stand and lower in the 10-year-old stand than in the other stands. Among stands, mean fine root (< 2 mm) biomass ranged from 49 to 398 g m(-2), SLR from 13.4 to 19.8 m g(-1), RLD from 980 to 11,650 m m(-3) and RSA from 2.4 to 35.4 m(2) m(-3). Most fine root biomass of trees was concentrated in the upper 20 cm of the mineral soil and in the humus layer (0-5 cm) in all stands. Understory fine roots accounted for 67 and 25% of total fine root biomass in the 10- and 120-year-old stands, respectively. Stand age had no affect on root tip number or branching frequency, but both parameters changed with soil depth, with increasing number of root tips and decreasing branching frequency with increasing soil depth for root fractions < 2 mm in diameter. Specific (mass based) root tip number and branching density were highest for the finest roots (< 1 mm) in the humus layer. Season (spring or fall) had no effect on tree fine root biomass, but there was a small and significant increase in understory fine root biomass in fall relative to spring. All morphological characteristics showed strong seasonal variation, especially the finest root fraction, with consistently and significantly higher values in spring than in fall. We conclude that fine root biomass, especially in the finest fraction (< 1 mm in diameter), is strongly dependent on stand age. Among stands, carbon concentration in fine root biomass was highest in the 30-year-old stand, and appeared to be associated with the high tree and canopy density during the early stage of stand development. Values of RLD and RSA, morphological features indicative of stand nutrient-uptake efficiency, were higher in the 30-year-old stand than in the other stands.     

Effects of repeated urea doses on soil chemistry and nutrient pools in a Norway spruce stand

Pools of macro-nutrients in soil and vegetation were studied in an old fertilization experiment with a large previous input of N. Different doses of N, in the form of urea, had been added four times during a 20-year period. In total, between 480 and 2400 kg N ha⁻¹ had been given. The experiment was established in a relatively productive Norway spruce stand and the expectation was that the large N input would cause an accelerated leaching of N, especially nitrate, accompanied by soil acidification and losses of several nutrients. The aim was to test for possible residual effects. Thirteen years after the last N addition, samples from the aboveground part of trees, field layer, S-layer, humus layer and mineral soil (0–10 cm) were analyzed for concentrations of most major nutrients. Nutrient pools were calculated. In the humus layer, the concentration of N increased and the C/N-ratio decreased with increasing N dose. The calculated recovery of added N in soil including ground vegetation was complete for the lowest N dose, while it was 25–50% for higher doses. The amount of N retained was unaffected by the N dose. The amount of extractable P in the upper part of the mineral soil was negatively correlated with N dose, as was also the concentration of total P in the S-layer. Neither soil pH, nor concentrations or amounts of Ca, Mg and K were affected by the previous fertilization. The calculated total soil-plant pool was only influenced by N dose in the case of P, which was 20% lower at the highest N dose compared with unfertilized conditions. Despite the large extra N input, the nutritional changes in plants and soil of the actual study site seemed surprisingly small.     

Impacts of changing climate on the productivity of Norway spruce dominant stands with a mixture of Scots pine and birch in relation to water availability in southern and northern Finland

A process-based ecosystem model was used to assess the impacts of changing climate on net photosynthesis and total stem wood growth in relation to water availability in two unmanaged Norway spruce (Picea abies) dominant stands with a mixture of Scots pine (Pinus sylvestris) and birch (Betula sp.). The mixed stands were grown over a 100-year rotation (2000-99) in southern and northern Finland with initial species shares of 50, 25 and 25% for Norway spruce, Scots pine and birch, respectively. In addition, pure Norway spruce, Scots pine and birch stands were used as a comparison to identify whether species' response is different in mixed and pure stands. Soil type and moisture conditions (moderate drought) were expected to be the same at the beginning of the simulations irrespective of site location. Regardless of tree species, both annual net canopy photosynthesis (P(nc)) and total stem wood growth (V(s)) were, on average, lower on the southern site under the changing climate compared with the current climate (difference increasing toward the end of the rotation); the opposite was the case for the northern site. Regarding the stand water budget, evapotranspiration (E(T)) was higher under the changing climate regardless of site location. Transpiration and evaporation from the canopy affected water depletion the most. Norway spruce and birch accounted for most of the water depletion in mixed stands on both sites regardless of climatic condition. The annual soil water deficit (W(d)) was higher on the southern site under the changing climate. On the northern site, the situation was the opposite. According to our results, the growth of pure Norway spruce stands in southern Finland could be even lower than the growth of Norway spruce in mixed stands under the changing climate. The opposite was found for pure Scots pine and birch stands due to lower water depletion. This indicates that in the future the management should be properly adapted to climate change in order to sustain the productivity of mixed stands dominated by Norway spruce.     

Effect of removal of logging residue on nutrient leaching and nutrient pools in the soil after clearcutting in a Norway spruce stand

Following clearcutting applying the conventional stem-only harvesting method in a Norway spruce (Picea abies (L.) Karst.) stand and different levels of removal of logging residue, the nutrient fluxes from the heaps of logging residue and from the O horizon were monitored over four growing seasons and the soil nutrient pools were determined. Three levels of removal of logging residue were carried out using (i) conventional stem-only harvesting (no residues removed); (ii) residues removed; and (iii) removal of branches (foliage left on site). The heaps of logging residue were a minor source of inorganic N entering the soil in the water percolating through the heaps, but they were a significant source of organic N, P, Ca, Mg, and especially K. Nutrient fluxes from the O horizon were in general greater under the heaps of logging residue as compared to soils without overlying logging residue. The leaching of inorganic N from the O horizon under the heaps of logging residue resulted in a net loss of these compounds, while the O horizon without overlying logging residue gained N. The removal of logging residue significantly decreased the extractable K pools in the soil while it or conversely, the presence of residue heaps had no significant effect on the pools of organic matter and the pools of N, P, Ca, and Mg in the O horizon and in the 0–10 cm soil layer. The results show that the short-term effects of logging residue on nutrient dynamics in the soil can be complex and difficult to interpret in terms of site productivity as there are changes in the nutrient fluxes, which imply the opposite effects on site productivity. However, the results do indicate that, in the short-term, the removal of logging residue does not impair pools of N in the soil nor site productivity on sites where the availability of N limits productivity.     

Fine root respiration in northern hardwood forests in relation to temperature and nitrogen availability

We examined fine-root (< 2.0 mm diameter) respiration throughout one growing season in four northern hardwood stands dominated by sugar maple (Acer saccharum Marsh.), located along soil temperature and nitrogen (N) availability gradients. In each stand, we fertilized three 50 x 50 m plots with 30 kg NO(3) (-)-N ha(-1) year(-1) and an additional three plots received no N and served as controls. We predicted that root respiration rates would increase with increasing soil temperature and N availability. We reasoned that respiration would be greater for trees using NO(3) (-) as an N source than for trees using NH(4) (+) as an N source because of the greater carbon (C) costs associated with NO(3) (-) versus NH(4) (+) uptake and assimilation. Within stands, seasonal patterns of fine-root respiration rates followed temporal changes in soil temperature, ranging from a low of 2.1 micro mol O(2) kg(-1) s(-1) at 6 degrees C to a high of 7.0 micro mol O(2) kg(-1) s(-1) at 18 degrees C. Differences in respiration rates among stands at a given soil temperature were related to variability in total net N mineralized (48-90 micro g N g(-1)) throughout the growing season and associated changes in mean root tissue N concentration (1.18-1.36 mol N kg(-1)). The hypothesized increases in respiration in response to NO(3) (-) fertilization were not observed. The best-fit model describing patterns within and among stands had root respiration rates increasing exponentially with soil temperature and increasing linearly with increasing tissue N concentration: R = 1.347Ne(0.072T) (r(2) = 0.63, P < 0.01), where R is root respiration rate ( micro mol O(2) kg(-1) s(-1)), N is root tissue N concentration (mol N kg(-1)), and T is soil temperature ( degrees C). We conclude that, in northern hardwood forests dominated by sugar maple, root respiration is responsive to changes in both soil temperature and N availability, and that both factors should be considered in models of forest C dynamics.     

Fine root biomass and production in Scots pine stands in relation to stand age

We determined fine root biomass and production of 15-, 35- and 100-year-old Scots pine (Pinus sylvestris L.) stands during three growing seasons. Fine roots were sampled by the soil core method. Mean (+/- SE) annual fine root biomass of Scots pine in the 15-, 35- and 100-year-old stands was 220 +/- 25, 357 +/- 21 and 259 +/- 26 g m(-2), respectively. Fine root biomass of the understory vegetation was 159 +/- 54 g m(-2), 244 +/- 30 and 408 +/- 81 g m(-2), and fine root necromass was 500 +/- 112, 1,047 +/- 452 and 1,895 +/- 607 g m(-2) in the sapling, pole stage and mature stands, respectively. Both understory and Scots pine fine root production increased with stand age. Mean annual Scots pine fine root production was 165 +/- 131, 775 +/- 339 and 860 +/- 348 g m(-2) year(-1) in the sapling, pole stage and mature stand, respectively. The respective mean annual production of all fine roots (Scots pine and understory) was 181 +/- 129, 1,039 +/- 497 and 1,360 +/- 869 g m(-2) year(-1). The Scots pine and understory fine root biomass, necromass and production varied in relation to stand age, although the variation was not statistically significant.     

Fine-root distribution and morphology in an acidic Norway spruce (Picea abies (L.) Karst.) stand in SW Sweden in relation to granulated wood ash application

Wood ash is recommended as a compensatory fertiliser to counteract the effects of acidic deposition on forest ecosystems. Spatial distribution of biomass, necromass and morphology parameters of the fine roots (diameter classes <1, 1–2, <2 mm) of Norway spruce (Picea abies (L.) Karst.) were analysed in response to fertilisation with granulated wood ash (GWA) in a long-term field experiment in SW Sweden. GWA was applied as a single dose of 3200 kg ha⁻¹ and the fine roots were sampled 9 years later by soil coring. Soil cores were divided into 1-cm strata within the top 0–2.5 cm humus limits, the lower humus below 2.5 cm (with varying thickness) and the mineral soil to 50 cm depth (from ground surface). Total fine-root biomass in the control (C) and GWA treatment, 256 ± 20 and 258 ± 25 g m⁻², respectively, and length 2072 ± 182 and 1800 ± 198 m m⁻², respectively, did not differ statistically from each other. Total fine-root necromass in the 1–2 mm fraction was significantly higher in C than in the GWA treatment, 130 ± 12 and 80 ± 10 g m⁻², respectively. Fine-root biomass in the <1 mm fraction was significantly greater in the lower humus in the GWA treatment, but this did not affect the total biomass in the <1 mm fraction in the whole soil profile. The biomass-to-necromass ratio (1–2 mm) was significantly higher in the GWA treatment in the 0–30 cm soil layer than in the corresponding layer of the control. Specific root length (SRL) was lower in the GWA treatment than in the control in the 0–5 cm soil layer. The lower necromass and SRL were more clearly related to the GWA treatment, whereas the difference in the vertical distribution of biomass may have been related to the thicker humus layer in the GWA plots.     

Fine root production and turnover in forest ecosystems in relation to stand and environmental characteristics

The production and turnover of fine roots (diameter ?2 mm) contributes significantly to carbon cycling in forest ecosystems. We compiled an up-to-date global database covering 186 stands from the literature and estimated fine root production (FRP) and fine root turnover (FRT) for boreal, temperate and tropical forests in order to study the relationships between FRP or FRT and environmental and stand variables. FRP for all plants (trees + understorey) was 311 ± 259 (n = 39), 428 ± 375 (n = 71) and 596 ± 478 g m⁻² a⁻¹ (n = 32) in the boreal, temperate and tropical forests, respectively, and the corresponding annual FRT rates were 0.77 ± 0.70, 1.21 ± 1.04 and 1.44 ± 0.76, respectively. When the FRP and FRT of trees were estimated separately for boreal and temperate forests the differences between the two biomes were insignificant. The mean FRP of trees for the two biomes combined was 306 ± 240 g m⁻² a⁻¹ (n = 86) and the annual FRT was 1.31 ± 1.43. Fine root biomass (FRB) was the most significant factor explaining the variation in FRP, and more so at the tree level than at the stand level, explaining 53% of the variation in FRP for trees at the tree level. The corresponding proportions at the stand level were 21% for all plants and 12% for trees. Latitude, mean annual temperature and annual precipitation each explained <20% of the variation in FRP or FRT. Fine root production and FRT estimates are highly dependent on the species included in the sampling, the sampling depth and the methods used for estimating FRP or calculating FRT. The results indicate that the variation in FRP on a global scale can be explained to a higher degree if we focus on tree roots separately from the roots of the understorey vegetation and on FRP at the tree level instead of FRP at the stand level or on FRT.     

White spruce foliar nutrient concentrations in relation to tree growth and soil nutrient amounts

Concentrations of foliar N, P, S, K, Ca, and Mg were studied in relation to stand age, tree growth, site index, and soil nutrient amounts for natural white spruce stands on a wide range of site conditions in the sub-boreal spruce zone of British Columbia, Canada. While Ca was sufficient in every sampled stand, relatively widespread deficiency in N was diagnosed. Deficiencies of other nutrients were diagnosed only on wet to very wet sites. Foliar N, P and K were negatively correlated with stand age and positively correlated with height and diameter growth. White spruce site index was positively correlated with foliar nutrients, and their relationships were quantified using a quadratic function. Foliar nutrients, except Ca, are positively correlated with soil nutrients measured in routine chemical analysis, and their relationships were quantified using Mitscherlich's function. It is recommended that the existing standards need to be modified should they be applied to nutrient diagnosis in natural white spruce stands. These standards appear too high for N and too low for P, K, and Ca.     

Effects of growth rates, tree morphology and site conditions on longevity of Norway spruce in the northern Swiss Alps

Longevity of trees is known to be associated with growth rates, but also with tree morphology and spatial influences. However, very little quantitative information is available on the effects of these biotic and abiotic influences on maximum ages of trees. The objectives of this study were to investigate the trade-off between longevity and growth rates of Norway spruce (Picea abies) and to quantify the effects of tree morphology and abiotic site conditions on longevity of this species. Data were collected along different topographical and climatic gradients in a 20?×?25?km study area in the northern part of the Swiss Alps (Glarus). The ages of the more than 100 sampled dead Norway spruces ranged between 50 and 367?years. Longevity of these trees was negatively related to tree growth, i.e. slow-growing trees tended to grow older than fast-growing trees. Tree height was positively associated with longevity for both upper and lower storey trees. Longevity of lower storey trees was increased with large crown diameter, but decreased with long crown length. Upper storey trees growing at higher altitude tended to get older than at lower altitude. We conclude that the combined effects of growth rates, variability in site conditions and different traits of tree morphology determine tree longevity of Norway spruce in the Swiss Alps. Because longevity is tightly linked to mortality rates of tree populations, our study may improve our understanding of long-term processes of forest dynamics under current and future climate.     

Effects of soil temperature on shoot and root growth and nutrient uptake of 5-year-old Norway spruce seedlings

Soil temperature is a main factor limiting root growth in the boreal forest. To simulate the possible soil-warming effect of future climate change, 5-year-old Norway spruce (Picea abies (L.) Karst.) seedlings were subjected to three simulated growing seasons in controlled environment rooms. The seedlings were acclimated to a soil temperature of 16 degrees C during the first (GS I) and third growing seasons (GS III), but were assigned to random soil-temperature treatments of 9, 13, 18 and 21 degrees C during the second growing season (GS II). In GS II, shoot diameter growth was lowest in the 21 degrees C treatment and root growth was lowest in the 9 degrees C treatment. In GS III, shoot height and root length growth improved in seedlings that had been kept at 9 degrees C during GS II, indicating compensatory growth in response to increased soil temperature. The temporary decrease in soil temperature had no long-lasting significant effect on seedling biomass or total nutrient uptake. At the end of GS III, fine roots of seedlings exposed to a soil temperature of 21 degrees C in GS II were distributed more evenly between the organic and mineral soil layers than roots of seedlings in the other treatments. During GS II and GS III, root growth started earlier than shoot growth, decreased during the rapid shoot elongation phase and increased again as shoot growth decreased.     

Leaf physiology and sugar concentrations of transplanted Quercus rubra seedlings in relation to nutrient and water availability

Newly planted seedlings incur transplant stress resulting from poor root-soil contact, which limits access to soil moisture and nutrients and reduces growth for one or more growing seasons. Controlled release fertilizer (CRF) applied at planting may reduce transplant stress by augmenting rhizosphere nutrient availability yet with potential risk of root system damage due to elevated fertilizer salt concentrations, which may be further exacerbated by drought. Under controlled conditions, we examined northern red oak (Quercus rubra L.) leaf physiological parameters and soluble sugar concentrations in response to varying nutrient levels (via CRF application) and moisture availability gradients ranging from drought to flooding. Net photosynthetic rates, transpiration rates, and chlorophyll fluorescence parameters responded positively to CRF application, and no interactions were observed between CRF and moisture availability; however, CRF did not increase soluble sugar concentrations. No effects of short-term drought were observed, but flooding exerted a rapid negative influence on net photosynthetic rates, transpiration rates, and chlorophyll fluorescence parameters; flooding also elevated soluble sugar concentrations, indicative of disrupted carbon partitioning and a much greater sensitivity to root-zone hypoxia than to drought in this species. Lack of interactions between CRF application and soil moisture availability indicates relatively similar responses of fertilized seedlings across moisture gradients.     

Diachronic analysis of individual-tree mortality in a Norway spruce stand in the eastern Italian Alps

? Understanding tree mortality processes across time requires long term studies. Spatiotemporal patterns of mortality in a 200 years-old mono-layered Norway spruce stand were evaluated to determine what factors affected individual-tree mortality.

? We performed an analysis on two surveys (1993 and 2005) in a 1-ha permanent plot in the Paneveggio forest (Eastern Italian Alps). Tree diameter and age distribution between surveys were compared. We examined spatial patterns of living and dead trees before 1993, in 1993 and in 2005 using univariate and bivariate Ripley’s K(d) function, and a kernel estimator of local crowding. A logistic model was used to assess the effects of diameter, age, recent growth and competitive pressure on tree mortality.

? Spatial pattern analysis indicated mortality was associated to tree neighbourhood (neighbour effect at 2–5 m). An increment of regularization of tree spatial pattern occurred due to density-dependent mortality. Logistic regression showed tree diameter and recent growth were determinant on mortality risk during the monitoring period.

? Even if the stand is relatively aged, mortality dynamics are those typical of stem exclusion stage. Mortality was related to competitive dynamics, and small suppressed trees with slow growth rate had higher probability to die.

     

The nutrient status of Norway spruce in pure and in mixed-species stands

Atmospheric deposition of N and S appears to have caused nutrient imbalance in Norway spruce stands in southern Sweden. This calls for a change of forest management to procedures that promote nutrient balance. Studies have shown lower soil acidity in Norway spruce/deciduous mixed stands than in spruce monocultures, but the tree nutrient status in such mixtures has not been much investigated so far.

The nutrient status of Norway spruce foliage and top mineral soil chemistry in monocultures and in stands mixed with beech, birch, or oak was investigated through paired comparisons on 30 sites in southern Sweden (27 sites) and eastern Denmark (three sites). In total, 45 mixed stands and 34 pure stands were included in the study.

Spruce needles from mixed stands had higher concentrations and ratios to N of K, P, and Zn than needles from pure spruce stands. Among the mixed stands, the K status appeared to be positively correlated with the percentage of deciduous tree basal area. Soil samples from mixed stands had a higher Mg concentration, base saturation, and BC/Al ratio than soil samples from pure stands. The spruce needle nutrient status was comparable in pure stands on fertile sites and in mixed stands on poor sites. We did not detect any differences in spruce tree growth between pure and mixed stands.

This paper discusses possible reasons for a positive effect on the tree nutrient status in mixed-species stands and the possibility of using mixed-species stands as a forest management procedure to avoid nutrient imbalance.