Age effects on Norway spruce (Picea abies) susceptibility to ozone uptake: a novel approach relating stress avoidance to defense

Cumulative ozone (O3) uptake and O3 flux were related to physiological, morphological and biochemical characteristics of Norway spruce (Picea abies (L.) Karst.) trees of different ages. Under ambient CO2 conditions, photosynthetic capacity (Amax) declined in mature trees when cumulative O3 uptake into needles, which provides a measure of effective O3 dose, exceeded 21 mmol m-2 of total needle surface area. A comparable decline in Amax of seedlings occurred when cumulative O(3) uptake was only 4.5 mmol m-2. The threshold O3 flux causing a significant decline in Amax ranged between 2.14 and 2.45 nmol m-2 s-1 in mature trees and seedlings subjected to exposure periods of > or = 70 and > or = 23 days, respectively. The greater O3 sensitivity of young trees compared with mature trees was associated with needle morphology. Biomass of a 100-needle sample increased significantly with tree age, whereas a negative correlation was found for specific leaf area, these changes parallel those observed during differentiation from shade-type to sun-type needles with tree ontogeny. Age-dependent changes in leaf morphology were related to changes in detoxification capacity, with area-based concentrations of ascorbate increasing during tree ontogeny. These findings indicate that the extent of O3-induced injury is related to the ratio of potentially available antioxidants to O3 influx. Because this ratio, when calculated for ascorbate, increased with tree age, we conclude that the ratio may serve as an empirical basis for characterizing age-related differences in tree responses to O3.     

Competition modifies effects of enhanced ozone/carbon dioxide concentrations on carbohydrate and biomass accumulation in juvenile Norway spruce and European beech

Elevated concentrations of carbon dioxide ([CO2]) and ozone ([O3]) affect primary metabolism of trees in opposite ways. We studied their potential interactions on carbohydrate concentrations and contents. Two hypotheses currently under debate were tested. (1) Stimulation of primary metabolism by prolonged exposure to elevated [CO2] does not compensate for the adverse effects of O3 on carbohydrate accumulation and biomass partitioning to the root. (2) Growth in a mixed-species planting will repress plant responses to elevated [O3] and [CO2] relative to conditions in a monoculture. To this end, European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst.) saplings grown under conditions of intra- and interspecific competition were pre-acclimated for 1 year to ambient or elevated [CO2]. In the following 2-year phytotron study, trees were exposed to factorial combinations of ambient and elevated [O3] and [CO2]. The total carbohydrate content (sugar and starch) of spruce was greater in plants exposed to elevated [CO2] than in plants exposed to ambient [CO2]. In beech, the opposite response was observed, especially when this species was grown in combination with spruce. Overall, the data did not support Hypothesis 1, because the adverse effects of O3 were counteracted by elevated [CO2]. Support for Hypothesis 2 was species-dependent. In beech saplings, reduction of carbohydrates by elevated [O3] and stimulation by elevated [CO2] were repressed by competitive interaction with spruce. In contrast, in spruce, stimulation of carbohydrates by elevated [CO2] was similar in mono- and mixed cultures. Thus Hypothesis 2 was supported for beech but not spruce. We conclude that, in juvenile beech and spruce, a 3-year exposure to elevated [CO2] counteracts the adverse effects of O3 on carbohydrate concentrations and contents. For beech, sensitivity to elevated [CO2] and [O3] was high in monoculture but was largely repressed by interspecific competition with spruce. In contrast, the response of spruce to perturbations of atmospheric chemistry was not significantly affected by either intra- or interspecific competition.     

Growth recovery of mature Norway spruce and European beech from chronic O<Subscript>3</Subscript> stress

Elevated O₃ levels can strongly impair the health and vitality of forest ecosystems. Free-air exposure systems reveal that forest tree and stand growth can be reduced strongly under chronic O₃ stress. Detailed knowledge of the effect of O₃ exposure on photosynthesis, carbon sequestration, allometry and growth during chronic stress is available. However, knowledge of growth response after O₃ reduction is scarce. Here, we analyse the growth of mature Norway spruce (Picea abies (L.) Karst.) and European beech (Fagus sylvatica L.) in the free-air O₃ fumigation experiment at Kranzberg Forest. We compare tree growth over a 9-year period (2008–2016) after exposure to O₃ (2000–2007). During 2?×?O₃ exposure, the annual basal area growth of Norway spruce and European beech decreased by 24 and 32%, respectively. After cessation of 2?×?O₃ exposure, the annual basal area growth of Norway spruce and European beech not only recovered but exceeded the growth of the trees in the control condition by 14 and 24%, respectively. The growth resilience and resistance of trees previously exposed to 2?×?O₃ towards drought stress and late frost was hardly lower than that of the trees in the control condition. The capacity for growth recovery even after long-term chronic O₃ stress emphasizes the strong beneficial effect of air pollution control on the health of forest ecosystems and on the global land carbon sink.     

Nutritional effects triggered by the extreme summer 2003 in the free air ozone fumigation experiment at the Kranzberger Forst

Since the year 2000 mature beech and spruce trees were treated in a field experiment with double ambient ozone concentrations. Elevated ozone had no influence on average single leaf biomass and there were also no ozone effects on leaf nutrient concentrations in climatic normal years. However, the extraordinary dry summer 2003 triggered significant differences between the fumigated and control trees. For beech in the year after the drought event the control trees surprisingly had significantly lower foliar levels of K and P than in former years, whereas the ozone exposed trees showed no significant nutritional effects. There are indications, that the trees exposed to double ambient ozone were already adapted to higher ozone values, whereas the control trees experienced extraordinary high ambient ozone concentrations in the dry and sunny summer 2003. For spruce in autumn 2003 and 2004 ozone treated trees had significantly higher foliar levels of K in current year needles than control trees, an effect which cannot be thoroughly interpreted yet on the basis of the dataset available. This article belongs to the special issue „Growth and defence of Norway spruce and European beech in pure and mixed stands“.     

Economic evaluation of cyclic regimes in beech (fagus sylvatica L.)

A general model for comparing the profitability of natural regeneration in beech (Fagus sylvatica L.) with planting of Norway spruce (Picea abies L.) is developed. The model is basically a method of comparing a series of net present values. An example of using the model is given: BEECH (90–110) denotes a cyclic regime in which regeneration is initiated at age 90 of a stand, and the canopy trees are cleared at age 110. This regime is compared with the alternative of planting Norway spruce. Economic data used for the comparison are representative of the southeastern regions of Denmark. The conclusion of the comparison is as follows: BEECH (90–110) is superior to planting of Norway spruce for required real rates of return up to 3–4%. For higher required rates of return planting of Norway spruce is preferable.     

Gap dynamics in the Western Carpathian mixed beech old-growth forests affected by spruce bark beetle outbreak

Understanding how species-specific disturbances affect the dynamics of mixed forests is becoming increasingly important due to rapidly changing disturbance regimes. This study estimated the effect of Norway spruce (Picea abies (L.) Karst.) mortality on the disturbance processes in two mixed beech stands of the Western Carpathians that were affected by a bark beetle outbreak. We evaluated the size distribution, fraction of canopy and expanded gaps, the characteristics of gapmakers and the contribution of different species to gap size. The measured canopy gap fraction was <5%, and most canopy gaps were small (<100 m²). Spruce was the most abundant gapmaker, and its share among gapmakers was 3–6 times higher than its share in the canopy. We found that the increase in spruce mortality due to the outbreak resulted in a fine-scale mortality pattern. However, spruce gapmakers did not contribute much to gap area size, as shown by a weak correlation between the number of spruce gapmakers and the area of expanded gaps. Diameter distribution of living versus recently dead trees showed that beech mortality occurred disproportionately in large size classes. However, dead spruce trees were equally frequent in all diameter classes, which means beetles did not exclusively attack larger trees in these stands during the outbreak. We conclude that spruce mortality may have influenced successional processes by giving a competitive advantage to two other species that were not affected by the outbreak, provided that a high deer browsing intensity does not hinder the regeneration of new seedlings.

     

Root distribution of under-planted European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) below the canopy of a mature Norway spruce stand as a function of light

Aboveground and belowground biomass of 15-year-old under-planted European beech seedlings (Fagus sylvatica L.) in Norway spruce stand were studied along a light gradient in three plots, in the northern part of Slovenia. Differences in soil water content, aboveground and fine root biomass distribution were confirmed between studied plots. Light had significant effect on the total biomass, root-shoot ratio (0.388 ± 0.076 under canopy, 0.549 ± 0.042 in the edge, 0.656 ± 0.047 in the open), specific root length (SRL) of fine beech roots (561.9 ± 42.2 under canopy, 664.3 ± 51.2 in the edge, 618.2 ± 72.8 in the open) and specific leaf area in beech, indicating morphological adjustment to shade. However, SRL of beech fine roots indicated no change between plots. The correlation between total aboveground and root biomass and light below the mature stand canopy was higher in the case of diffuse light intensity. Most fine roots of spruce were concentrated in the top (0–20 cm) soil layer. Beech fine roots under canopy and edge conditions were also concentrated in top (0–20 cm) soil layer and exhibited shift downwards to deeper soil horizons in open plot. Root proportion between beech and spruce changed with light toward beech with increasing light intensity for both fine and coarse roots.     

Photosynthetic characteristics of Fagus sylvatica and Quercus robur established for stand conversion from Picea abies

Efforts in Europe to convert Norway spruce (Picea abies) plantations to broadleaf or mixed broadleaf-conifer forests could be bolstered by an increased understanding of how artificial regeneration acclimates and functions under a range of Norway spruce stand conditions. We studied foliage characteristics and leaf-level photosynthesis on 7-year-old European beech (Fagus sylvatica) and pedunculate oak (Quercus robur) regeneration established in open patches and shelterwoods of a partially harvested Norway spruce plantation in southwestern Sweden. Both species exhibited morphological plasticity at the leaf level by developing leaf blades in patches with an average mass per unit area (LMA) 54% greater than of those in shelterwoods, and at the plant level by maintaining a leaf area ratio (LAR) in shelterwoods that was 78% greater than in patches. However, we observed interspecific differences in photosynthetic capacity relative to spruce canopy openness. Photosynthetic capacity (A₁₆₀₀, net photosynthesis at a photosynthetic photon flux density of 1600 μmol photons m⁻² s⁻¹) of beech in respect to the canopy gradient was best related to leaf mass, and declined substantially with increasing canopy openness primarily because leaf nitrogen (N) in this species decreased about 0.9 mg g⁻¹ with each 10% rise in canopy openness. In contrast, A₁₆₀₀ of oak showed a weak response to mass-based N, and furthermore the percentage of N remained constant in oak leaf tissues across the canopy gradient. Therefore, oak photosynthetic capacity along the canopy gradient was best related to leaf area, and increased as the spruce canopy thinned primarily because LMA rose 8.6 g m⁻² for each 10% increase in canopy openness. These findings support the premise that spruce stand structure regulates photosynthetic capacity of beech through processes that determine N status of this species; leaf N (mass basis) was greatest under relatively closed spruce canopies where leaves apparently acclimate by enhancing light harvesting mechanisms. Spruce stand structure regulates photosynthetic capacity of oak through processes that control LMA; LMA was greatest under open spruce canopies of high light availability where leaves apparently acclimate by enhancing CO₂ fixation mechanisms.     

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.     

Net photosynthetic response to light intensity of shoots from different crown positions and age in picea abies (L.) karst

Carbon dioxide uptake response to light intensity of detached shoots of open‐ and forest‐grown Norway spruces was investigated in controlled standard conditions. The initial slope of the CO₂ uptake light response curve, calculated on leaf area basis and the internal conductance, calculated on leaf weight basis were almost constant for shoots from different positions in the forest canopy. In open‐grown trees the maximum photosynthesis and internal conductance (on leaf weight basis) decreased in relation to the tree age, so that in the 8‐year‐old tree it was about two times of that in the 66‐year‐old tree. These results suggest that 1) the enhanced efficiency of “shade” shoots is caused by morphologic adaptation, i.e. the more sparsely packed photosynthetic apparatus in needles and shoots, and 2) adaptational possibilities of Norway spruce photosynthetic apparatus, its ecological plasticity diminish during tree ontogenesis.     

Photosynthesis in Norway spruce seedlings infected by the needle rust Chrysomyxa rhododendri

Chrysomyxa rhododendri (DC.) De Bary is a needle rust with a host shift between Rhododendron sp. and Norway spruce (Picea abies (L.) Karst.), penetrating only the new developing flushes of the conifer. Because little is known about its effects on trees, we investigated several parameters related to photosynthesis in artificially infected 3-year-old Norway spruce seedlings. The potential efficiency of photosystem II (PSII; derived from chlorophyll fluorescence measurements) was reduced in infected current-year needles as soon as disease symptoms were visible, about three weeks after inoculation. Two weeks later, photosynthetic O(2) evolution (P(max)) of infected needles was less than 20% of control needles, whereas respiratory O(2) uptake (R(D)) was about three times higher than that of control needles. Nonstructural carbohydrate concentrations were about 60% of control values in all parts of the shoots of infected trees. Photosynthetic inhibition was associated with marked decreases in chlorophyll concentration and chlorophyll a/b ratio but only a small reduction in carotenoid concentration. In infected trees, P(max) of noninfected 1-year-old and 2-year-old needles was 50 and 80% higher than in the corresponding age class of needles of control trees. Estimation of potential daily net dry mass production, based on P(max), R(D), specific leaf area, carbon content and needle biomass, indicated that seedlings infected once were able to produce 60%, and those infected twice only 25%, of the dry mass of controls. We conclude that afforestation and regeneration of Norway spruce is seriously impaired in regions where seedlings are frequently attacked by Chrysomyxa.     

Litter impair spruce seedling emergence in beech forests: a litter manipulation experiment

European beech Fagus sylvatica and Norway spruce Picea abies are economically and ecologically important forest trees in large parts of Europe. Today, the beech forest reaches its northern distribution limit in south-eastern Norway and it is expected to expand northwards due to climate warming. This expansion will likely result in fundamental ecosystem changes. To increase our knowledge about the competitive balance between spruce and beech, we have investigated how beech and spruce litter affect spruce seedling emergence, growth and uptake of C and N. We did this in a seed-sowing experiment that included litter layer removal as well as reciprocal transplantations of litter layers between spruce and beech forests. Our results show that spruce seedling emergence was significantly impaired by both litter layer types, and especially so by the beech litter layer in the beech forest. The low seedling emergence in beech forests is concurrent with their lower light availability.     

Stand density and growth of Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> (L.) Karst.) and European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.): evidence from long-term experimental plots

On the basis of nine Norway spruce (Picea abies (L.) Karst.) and ten European beech (Fagus sylvatica L.) thinning experiments in Germany, for which both residual and removed stock had been registered first during 1870, I scrutinize how moderate and heavy thinning from below (B-, C-grade) affects the production of merchantable volume compared with light thinning (A-grade). In relation to A-grade, cumulative merchantable volume (CV) of B- and C-grade amounts in average to 103–107% in juvenile and to 97–102% in mature Norway spruce stands. The corresponding findings for European beech are 101–106% and 94–102%. CV of individual stands varies between 89% and 130% for Norway spruce and 73% and 155% for European beech (CV of A-grade = 100%). These findings are substantiated by the relation between stand density (SDI) and periodic annual increment (PAI). On the B- and C-grade plots of spruce and beech, respectively, SDI was reduced down to 41–91% and 31–83% of the A-grade. When SDI is reduced in young stands, PAI follows a unimodal curve. Norway spruce’s PAI culminates in 109% if SDI is reduced to 59%; European beech’s PAI culminates in 123% when density is reduced to 50%. Whereas Norway spruce’s growth reacts most positively on thinning under poor site conditions and with increment reduction on favourable sites, European beech behaves oppositely. With stand development the culmination point of the unimodal relation moves towards maximum density, so that in older stands PAI follows the increasing pattern, which is the left portion of a unimodal curve. A model is presented which apparently unifies contradictory patterns of stand density–growth reactions by integrating relative stand density, average tree size and site fertility effects, and makes the findings operable for forest management.     

Regeneration growth in different light environments of mixed species, multiaged, mountainous forests of Romania

Growth of regenerating trees in different light environments was studied for the mountainous, mixed-species forests in the Carpathian Mountains of Romania. The primary species in these mixtures were silver fir (Abies alba Mill.), European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst). Seedlings/saplings of these species were selected and measured in different stands from two different geographical locations. Regenerating trees were measured for height and diameter growth during the summer of 2002. For each seedling/sapling, percentage of above canopy light (PACL) and stand basal area (BA) were used to assess available and occupied growing space respectively. Regeneration growth was compared against these two variables and regression relationships were developed. Using these models, we predicted the dynamics of regeneration as both growth and species composition. Our results showed that in low-light environments (PACL<20–35%; BA>30 m²/ha), shade tolerant fir and beech clearly outcompeted the spruce. Therefore, in dense stands, spruce could be eliminated by the shade tolerant species. For intermediate levels of cover (PACL=35–70%; BA=15–35 m²/ha) the spruce grew at comparable rates as the beech and fir. All three species showed similar growth rates in open conditions (PACL>80–90%; BA<15–20 m²/ha) with the spruce having a tendency to outgrow the others. However, in terms of establishment, such conditions favor spruce and inhibit fir and beech.     

Nutrient contents and concentrations in relation to growth of Picea abies and Fagus sylvatica along a European transect

Mineral nutrition of Norway spruce (Picea abies (L.) Karst.) and beech (Fagus sylvatica L.) was investigated along a transect extending from northern Sweden to central Italy. Nitrogen (N) concentrations of needles and leaves in stands growing on acid soils did not differ significantly between central Italy and southern Sweden (1.0 +/- 0.1 mmol N g(-1) for needles and 1.9 +/- 0.14 mmol N g(-1) for leaves). In both species, foliar N concentrations were highest in Germany (1.2 mmol N g(-1) for needles and 2.0 mmol N g(-1) for leaves) and decreased by 50% toward northern Sweden (0.5 mmol N g(-1)). Both species showed constant S/N and P/N ratios along the transect. Calcium, K and Mg concentrations generally reflected local soil conditions; however, Mg concentrations reached deficiency values in Germany. Leaf area per unit dry weight varied significantly along the transect with lowest values for Norway spruce recorded in northern Sweden and Italy (3.4 m(2) kg(-1)) and a maximum in central Europe (4.7 m(2) kg(-1)). A similar pattern was observed for beech. Despite the low variation in foliar N concentrations on the large geographic scale, local and regional variations in N concentrations equalled or exceeded the variation along the entire continental transect. Furthermore, nutrient contents (i.e., nutrient concentration x dry weight per needle or leaf) showed a greater variation than nutrient concentrations along the transect. Nitrogen contents of Norway spruce needles reached minimum values in northern Sweden (2.4 micro mol N needle(-1)) and maximum values in Denmark (5.0 micro mol N needle(-1)). The N content of beech leaves was highest in Denmark (242 micro mol N leaf(-1)). At the German site, foliar N content rather than N concentration reflected the seasonal dynamics of foliar growth and N storage of the two species. During foliage expansion, there was an initial rapid increase in N content and a decrease in N concentration. This pattern lasted for about 2 weeks after bud break and was followed by 6 weeks during which dry weight and N content of the foliage increased, resulting in a further decrease in N concentration. During summer, dry weight and N content of mature needles of Norway spruce increased further to reach a maximum in autumn, whereas N concentration remained constant. In spring, reallocation of N from 1- and 2-year-old needles was 1.5 and 1.0 micro mol N needle(-1), respectively. This remobilized N was a major source of N for the development of new needles, which had an N content of 1.5 micro mol N needle(-1) after bud break. The seasonal remobilization of N from old foliage decreased with increasing needle age. Needle N content and dry weight decreased progressively with age (1 micro mol N needle(-1) between age classes 2 and 5), whereas N concentrations remained constant. For Norway spruce, annual stemwood production was correlated with needle N content but not with foliar N concentration or with the total amount of N in the canopy. Interspecific and geographical differences in plant nutrition are discussed on the basis of competitive demands for C and N between growth of foliage and wood.     

Climate change impacts on stand structure and competitive interactions in a southern Swedish spruce–beech forest

It is believed that European beech (Fagus sylvatica L.) will increase its competitive ability at its northern range margin in Scandinavia due to climate change. In mixed old-growth forests of beech and Norway spruce (Picea abies (L.) Karst.) at Siggaboda nature reserve (southern Sweden), stand structure characteristics were sequentially recorded in the years 2004, 2005 and 2007 as well as growth in stem diameter using tree-coring analyses. Using these measurements, we studied the effects on stand dynamics of an extreme storm event (2005 “Gudrun” hurricane), drought and heat (mid-summer 2006, spring 2007) and subsequent bark beetle attacks on spruce (growing season 2007), overlaid with warming tendencies. The storm, which caused disastrous damage in many stands nearby, had comparatively little impact on the structure of the spruce–beech stand. All together, only 32 trees (19 spruces, 10 beeches, 3 other species) per hectare were thrown or broken mainly in the leeward direction (NE) or impacted by secondary damage by uprooted neighbour trees; this represents 7% of the total tree number and 11% of the growing stock. Diameter and height structure did not change significantly. However, the 2006 drought and the 2007 attack of biotic agents changed the stand structure and composition strongly due to the death of about 19% of the dominating older spruce trees that accounted for 35% of total stand volume. This resulted in a considerable increase in beech’s contribution to stem number (4% increase) and wood volume of the living stand (7% increase). A comparison of diameter growth of beech and spruce during the periods 1894–1949 and 1950–2005 showed a distinct decrease in growth superiority of spruce during the last 50 years. These results support the idea of a northward migration of European beech as a nemoral tree species in Sweden, due to a higher tolerance to the abiotic and biotic threats accompanying climate change and an increased competitive ability compared to boreal tree species Norway spruce.     

Canopy composition as a measure to identify patterns of nutrient input in a mixed European beech and Norway spruce forest in central Europe

The influence of canopy composition on litterfall and throughfall was investigated in a mixed spruce beech forest in central Germany. We hypothesised that different parts of the mixed canopy created distinct patterns of element inputs via litterfall and throughfall. The investigation was carried out in two plots, representing the most contrasting cases of mixed forests: a stand greatly dominated by spruce (SDP) and a stand greatly dominated by beech (BDP). The canopies of the two plots were classified in four categories: pure beech, pure spruce, mixed canopy and gap. Amounts of throughfall water were lower and major element fluxes were higher under spruce than under beech in both plots, indicating that the nutrient inputs under the canopies of individual trees are driven by species-specific properties of the canopies and are quite independent of the degree of admixture. With the exception of K⁺, mixed canopies showed intermediate element inputs via throughfall, compared with pure canopy classes. The K⁺ input was significantly greater under mixed canopies, and these differences were more pronounced in the SDP than in the BDP. Results suggest that individual spruce trees in the BDP induce greater spatial heterogeneity of throughfall input than individual beech trees in the SDP. Nutrient inputs via foliar litterfall were similar among the different canopy classes, but the Mg input was lower under spruce canopy. This effect was balanced by higher Mg input via spruce throughfall. In our study, throughfall was the main source of heterogeneity in nutrient inputs, while foliar litterfall had a homogenising effect.     

The effects of Douglas-fir on tree-specific arthropod communities in mixed species stands with European beech and Norway spruce

The ecological effects of planting exotic Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] in Central Europe are still poorly understood. The aim of this study was to answer the question of whether Douglas-fir affects tree specific arthropod communities in different mature forest types (Douglas-fir, spruce and beech dominated) in Southern Germany. Therefore, arthropod communities of stem and tree crown strata of Douglas-fir and spruce (Picea abies L.) were sampled in the years 1999–2001 using arboreal photo-eclectors and flight interception traps. Statistical analysis was conducted for all species and focused on conifer specialists at three levels: (1) species diversity, (2) guild structure and (3) community structure. Within the stem stratum, species diversity was significantly higher on spruce than on Douglas-fir independent of year and stand composition. This could not be explained by a single feeding guild, rather by species changing strata during the vegetation period. In contrast, species diversity in tree crowns was approximately the same for both conifer species. However, communities in Douglas-fir crowns were conspicuously different from those in spruce crowns, especially in the Douglas-fir dominated stand type. While zoophagous insects exhibited higher activity on Douglas-fir in 2000, xylophagous beetles were more abundant on spruce in 2001. In European beech stands with widely spaced Douglas-fir trees, the site specific and broad-leaved tree related fauna might be maintained. In addition, Douglas-fir with its resource of Adelges cooleyi and crowns that overtop the broad-leaved tree canopy, offer additional resources for several aphidophagous and thermophile species.     

Why does leaf nitrogen decline within tree canopies less rapidly than light? An explanation from optimization subject to a lower bound on leaf mass per area

A long-established theoretical result states that, for a given total canopy nitrogen (N) content, canopy photosynthesis is maximized when the within-canopy gradient in leaf N per unit area (N(a)) is equal to the light gradient. However, it is widely observed that N(a) declines less rapidly than light in real plant canopies. Here we show that this general observation can be explained by optimal leaf acclimation to light subject to a lower-bound constraint on the leaf mass per area (m(a)). Using a simple model of the carbon-nitrogen (C-N) balance of trees with a steady-state canopy, we implement this constraint within the framework of the MAXX optimization hypothesis that maximizes net canopy C export. Virtually all canopy traits predicted by MAXX (leaf N gradient, leaf N concentration, leaf photosynthetic capacity, canopy N content, leaf-area index) are in close agreement with the values observed in a mature stand of Norway spruce trees (Picea abies L. Karst.). An alternative upper-bound constraint on leaf photosynthetic capacity (A(sat)) does not reproduce the canopy traits of this stand. MAXX subject to a lower bound on m(a) is also qualitatively consistent with co-variations in leaf N gradient, m(a) and A(sat) observed across a range of temperate and tropical tree species. Our study highlights the key role of constraints in optimization models of plant function.     

The Effect of Landfill Leachate Spraying on Foliar Nutrient Concentrations and Leaf Transpiration in a Northern Hardwood Forest, Canada

The effect of municipal solid waste (MSW) leachate spray irrigationon a mature northern hardwood forest was investigated. Canopyfoliar samples and stem increment cores were collected fromtwo indicative species, sugar maple (Acer saccharum Marsh.)and American beech (Fagus grandifolia Ehrh.), within each ofa heavily sprayed, lightly sprayed and control area. Foliarconcentrations of N and P were significantly higher in bothmaple and beech foliage within the sprayed areas when comparedto an unsprayed area (control). Levels of Mg and K were markedlyhigher in maple but not beech foliar samples within the heavilysprayed areas when compared to foliage sampled within the unsprayedcontrol. While no significant trends were observed within themaple foliage, both Fe and B levels increased significantlyin beech foliar samples obtained from within the heavily sprayedarea in comparison to foliage samples from the control. Directporometric measurements of the transpiration rate and diffusive(stomatal) resistance of canopy and understory plant leavesrevealed a significant increase in diffusive resistance anda decline in transpiration rate with leachate spraying. Afterfour years of spraying a significant effect of leachate applicationon radial stem growth of both maple and beech trees has notbeen observed.