Effects of shade on growth and mortality of maple (Acer pseudoplatanus), ash (Fraxinus excelsior) and beech (Fagus sylvatica) saplings

We compared shade tolerance of maple, ash and beech in the saplingstage from two sites with rich soils differing in water supply,growing in dense thickets underneath a beech shelterwood ofvarying canopy densities. Shade tolerance was described by twocomponents: mortality in shade and height growth in high light.At low light, beech showed the least mortality, maple the highestand ash in between on both sites. The decline with increasinglight was steepest in beech and more gradual with ash and maple.At 15 per cent above canopy light, all three species approachedzero mortality. Beech as the most shade-tolerant species hadthe highest survival rate under low light and the least lengthgrowth rate under high light (>17 per cent). Ash had a lowersurvival rate at low light than beech and a highest growth rateat high light. Maple showed a bit weaker trade-off with thelowest survival rate but a growth rate inferior to ash. On thebetter water-supplied site, height growth was significantlysuperior in all three species only under high light. On thebasis of these results, silvicultural conclusions are drawnwith respect to appropriate light levels and cutting types.     

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

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

Leaf photosynthetic characteristics of beech (Fagus sylvatica) saplings during three years of exposure to elevated CO(2) concentration

Beech (Fagus sylvatica L.) seedlings were cultivated from seeds sown in pots or directly in the ground in outdoor chambers that were transparent to solar radiation, and provided either ambient air or CO(2)-enriched air (ambient + 350 &mgr;mol mol(-1)). The rooting volume was high in all experiments. In the short-term experiment, potted plants were assigned to a factorial CO(2) x nutrient treatment (optimal nutrient supply and severe nutrient shortage) for 1 year. In the long-term experiment, plants were grown directly in the ground and received an optimal supply of water and nutrients in both CO(2) treatments for 3 years. Nutrient stress caused carboxylation capacity (V(m)) to decrease in the potted seedlings exposed to CO(2)-enriched air during their first growing season. In the long-term experiment with optimal nutrient supply, CO(2)-enriched air did not affect V(m), but caused an upward acclimation of maximum electron transport rate (J(m)). Consequently, there was a 14% increase in the J(m)/V(m) ratio, indicating nitrogen reallocation to maintain an equilibrium between RuBP consumption and RuBP regeneration. Both V(m) and J(m) decreased during the growing season in both CO(2) treatments. Although upward acclimation of J(m) was no longer apparent at the end of the third growing season, plants in CO(2)-enriched air maintained a higher J(m)/V(m) ratio than plants in ambient air, indicating that photosynthetic acclimation always occurred. Second flush leaves appeared during each growing season. When expressed on the basis of foliar nitrogen concentration, their photosynthetic characteristics (V(m) and J(m)) were enhanced compared with other leaves. Because the number of second flush leaves was also increased in the elevated CO(2) treatment, this response should be taken into account when modeling the effects of elevated CO(2) concentration on canopy photosynthesis. Stomatal conductance decreased in response to atmospheric CO(2) enrichment; however, the stomatal response to irradiance followed a single relationship based on two stomatal conductance models.     

Hydraulic architecture correlates with bud organogenesis and primary shoot growth in beech (Fagus sylvatica)

In beech (Fagus sylvatica L.), the number of leaf primordia preformed in the buds determines the length and the type (long versus short) of annual growth units, and thus, branch growth and architecture. We analyzed the correlation between the number of leaf primordia and the hydraulic conductance of the vascular system connected to the buds. Terminal buds of short growth units and axillary buds of long growth units on lower branches of mature trees were examined. Buds with less than four and more than five leaf primordia formed short and long growth units, respectively. Irrespective of the type of growth unit the bud was formed on, the occurrence of a large number of leaf primordia was associated with high xylem hydraulic conductance. Xylem conductance was correlated to the area of the outermost annual ring. These results suggest that organogenesis and primary growth in buds correlates with secondary growth of the growth units and thus with their hydraulic architecture. Possible causal relationships between the variables are discussed.     

Six-year time course of light-use efficiency, carbon gain and growth of beech saplings (Fagus sylvatica) planted under a Scots pine (Pinus sylvestris) shelterwood

Two-year-old Fagus sylvatica L. saplings were planted under the cover of a Pinus sylvestris L. stand in the French Massif Central. The stand was differentially thinned to obtain a gradient of transmitted photosynthetically active radiation (PAR(t); 0-0.35). Eighteen Fagus saplings were sampled in this gradient, and their growth (basal stem diameter increment) was recorded over six years. Over the same period, morphological parameters (leaf area, number and arrangement in space) were monitored by 3D-digitization. Photosynthetic parameters were estimated with a portable gas-exchange analyzer. Photosynthesis was mainly related to light availability, whereas sapling morphology was mainly driven by sapling size. Annual stem diameter increment was related to the amount of light-intercepting foliage (silhouette to total leaf area ratio (STAR) x total sapling leaf area (LA)) and light availability above the saplings (PAR(t)). However, light-use efficiency, i.e., the slope of the relationship between STAR x LA x PAR(t) and stem diameter increment, decreased over time as a result of a relative decrease in the proportion of photosynthetic tissues to total sapling biomass.     

Thermally initiated solvent-free radical modification of beech (Fagus sylvatica) wood

A procedure for rapidly modifying beech wood using a thermally initiated solvent-free grafting system was examined. In the modification, butyl acrylate and butyl methacrylate were used as vinyl monomers. Free radicals were generated from 2,2′-azobis(2-methylpropionitrile) or benzoyl peroxide at 103 and 180 °C by contact heating of the modified material. Chemical changes in the material were investigated by FTIR and X-ray photoelectron spectroscopies. The modification resulted in decreased surface wetting of the material manifested by increased water contact angles. The hardness of the resultant material decreased, while its color changed by the effect of temperature. It was shown that the approach allowed for efficient thermal-initiated modification of wood with rapid contact heating.     

Response of planted beech (Fagus sylvatica L.) and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) saplings to herbaceous and small shrubs control on clearcuts

The reaction of young beech (Fagus sylvatica L.) and Douglas fir (Pseudotsuga menziesii (Mirb.) Franco) saplings on competition of two types of vegetation—(1) gramineous with mainly Agrostis capillaries, Calamagrostis epigejos, Deschampsia flexuosa, and (2) small shrubs with mainly Rubus fruticosus and R. idaeus—on clear cuts on two sites was studied for 2 years. Half the sample saplings were released from competing vegetation by repeated herbicide applications. This treatment significantly raised the diameter increment in both species at the site with higher competition intensity, and more strongly after the removal of small shrubs than after the removal of grasses. Sapling length increment was not significantly affected. After being released from small shrubs, saplings of both species developed a smaller specific fine root length (cm g⁻¹ fine root biomass) than unreleased saplings during the second year which was characterized by low rainfall. Root nitrogen concentration significantly increased after weed control in both vegetation types. Sapling foliar content of main nutritional elements was negatively related to dry mass and total chemical content of surrounding ground vegetation. Based on these results, a release from ground vegetation could be a useful tool to improve growth of planted beech and Douglas-fir saplings on sites with well-developed small shrubs competition (mainly by Rubus fruticosus and R. idaeus), or under fairly dry conditions.     

Interaction of lignin and polysaccharides in beech wood (Fagus sylvatica) during drying processes

Summary ¹³C CP MAS NMR spectroscopy was used to characterize the structural changes of cell wall polymers in beech wood Fagus sylvatica during drying processes. The analysis of five wood samples, namely, untreated, untreated dried, pre-treated by steam and/or NaOH subjected to drying showed partial depolymerization of lignin component as well as the change of the ratio of the crystalline and of the amorphous parts of cellulose as the consequence of wood pre-treatment. In addition, T_1ρ(¹H) relaxation times were determined in beech wood sample pre-treated with steam at 135 °C and the lignin isolated from this sample. The magnitudes of the relaxation times were found comparable in both samples as well as in the lignin-cellulose model compound. These unique T_1ρ (¹H) values indicate that spin diffusion is complete and homogeneous due to spatial proximity of spins and confirmed the formation of lignin-cellulose complex during thermal treatment of wood. Received 30 June 1997     

A twig blight of understorey European beech (Fagus sylvatica) caused by soilborne Phytophthora spp.

During and after prolonged periods of rainfall in late spring, blighted young twigs of European beech (Fagus sylvatica) were frequently observed in several beech stands in south‐western and southern Germany. Long and short shoots of young understorey trees or lower branches up to 1.5 m above the soil level were affected. Symptoms also occurred regularly on twigs in heights up to 2 m and more above the ground. Necroses usually expanded within the current year’s tissue and often also reached into the previous year’s wood. Ponding rain water in the stands or along forest roads or open soil seemed to promote the disease. Of a total of 54 symptomatic twigs collected in four stands, 37 revealed Phytophthora isolates, of which 33 were P. plurivora and four were P. cambivora. Both species caused extensive lesions on beech twigs in laboratory pathogenicity tests. Patterns of the disease indicated that these pathogens, generally considered soilborne species, in most cases are transmitted from the soil to above‐ground parts of the trees via rain splash. In larger heights, however, other vectors such as snails might be responsible for transmission. Although Phytophthora spp. are well known as causal agents of seedling blight as well as root and cambium rot and aerial bleeding cankers of mature beech, to our knowledge this is the first report of a twig blight in beech associated with soilborne Phytophthora spp. In particular in periods of high precipitation, this disease might pose an additional threat to Central European beech forests, especially endangering the success of artificial and natural regeneration of beech in affected stands.     

Uptake and distribution of calcium and phosphorus in beech (Fagus sylvatica) as influenced by aluminum and nitrogen

We studied the effects of excess nitrogen added as nitrate (NO(3) (-)) or ammonium (NH(4) (+)), or both, on mineral nutrition and growth of beech (Fagus sylvatica L.) plants grown at pH 4.2 in Al-free nutrient solution or in solutions containing 0.1 or 1.0 mM AlCl(3). A high external concentration of NH(4) (+) increased the concentration of nitrogen in roots, stems and leaves. The root/shoot dry weight ratio was less in plants grown in the presence of NH(4) (+) than in plants grown in the presence of NO(3) (-). The concentration of phosphorus in the roots was increased and the concentration of potassium in all parts of the plant was decreased by NH(4) (+). A high external concentration of NO(3) (-) caused a decrease in phosphorus concentrations of the root, stem and leaf. Uptake of (45)Ca(2+) by roots was reduced in the presence of high concentrations of NH(4) (+) or NO(3) (-), and a combination of high concentrations of nitrogen and aluminum further reduced the uptake of (45)Ca(2+). Uptake of phosphate ((32)P) and concentrations of phosphorus in root and shoot were increased when plants were grown in the presence of 0.1 mM Al. Exposure to 1.0 mM Al, however, reduced the concentration of phosphorus in roots and shoots and the reduction was greater when plants were grown in the presence of a high external NO(3) (-) concentration. Aluminum binds to roots, and plants grown in the presence of 1.0 mM Al had a slightly higher concentration of aluminum in roots than plants grown in the presence of 0.1 mM Al, whereas the concentration of Al in the shoot was increased 2 to 3 times in plants exposed to 1.0 mm Al. Furthermore, the effects of 1.0 mM Al on uptake of other macronutrients were quite different from the effects of 0.1 mM Al. We conclude that 0.1 mM Al facilitates uptake and transport of phosphorus in beech and that between 0.1 and 1.0 mM Al there is a dramatic change in the effects of Al on uptake and transport of divalent cations and phosphorus.     

Effects of selective thinning on growth and development of beech (Fagus sylvatica L.) forest stands in south-eastern Slovenia

We studied the effects of two types of selective thinning on beech stands formed by a shelterwood cut in 1910 — with lower number of crop trees and higher thinning intensity (T1) and higher number of crop trees with lower thinning intensity (T2). The stands were thinned in 1980, 1991 and 2001. Despite a lower stand density after thinning, the annual basal area increments of thinned stands in both thinning periods (1980–1991 and 1991–2002) were around 20% higher compared to those of the control (unthinned) stands. The mean annual basal area increment of dominant trees was 30–56% larger in the thinned plots compared to the control plots. Of 176 initial crop trees in the T1, 72% were chosen again during the last thinning. In the T2, 258 crop trees were chosen in the first thinning, and only 62% of these trees were chosen again during the last thinning. Only crown suppression and diameter classes of crop trees significantly influenced their basal area increment when diameter classes, crown size, crown suppression, and social status were tested. In the thinned stands, the dominant trees are more uniformly distributed if compared to the dominant trees in the control plots. Finally, the herbaceous cover and the species diversity were higher in the thinned plots.     

Precommercial thinning of beech (Fagus sylvatica L.): Early effects of stump height on growth and natural pruning of potential crop trees

Abstract

This paper reports the early effects of stump height on the growth and natural pruning of potential crop trees after precommercial thinning of a young stand of naturally regenerated beech (Fagus sylvatica L.) in Denmark. The experiment comprises five treatments based on combinations of three grades of precommercial thinning and three stump heights. Treatments include the unthinned control, thinning only of whips and wolf trees (using low stumps), and thinning for potential crop trees using low, medium or high stumps. All treatments were replicated three times. Stump heights averaged 10, 90 or 230 cm, respectively. Pretreatment stem number (live trees) varied from 17,500 to 41,000 ha^?1. In the potential crop tree treatment, post-treatment stem number ranged from 4750 to 9500 ha^?1. Following two growth seasons, the quantity of stump regrowth increased with increasing stump height, the rate of stump regrowth increased with increasing stump height, the diameter growth of potential crop trees increased with decreasing stump height, the increase in stand height did not depend on stump height or post-treatment stem number, and the natural pruning of potential crop trees increased with increasing stump height. It remains to be seen whether these trends hold in the long run, and whether additional economic return from the increase in wood quality with increasing stump height compensates for the reduction in diameter growth.     

Impact of late frost events on radial growth of common beech (Fagus sylvatica L.) in Southern Germany

Phenological, temperature, and tree-ring data were used in order to identify and quantify the impact of late frosts on common beech (Fagus sylvatica L.) at different altitudes in Southern Germany during the last century. For this intention, dendroecological investigations were made upon trees at the Meteorological Observatory Hohenpeißenberg as well as from seven stands in the Bavarian Forest and 17 stands at the northern fringe of the Alps. From these locations, a considerable number of severe growth minima in the tree-ring series could be related to late frost in the days of or immediately after leaf unfolding. The frequency of frost-related growth minima increases with altitude. In individual years, radial growth can be reduced by more than 90% (stand mean) in relation to the average growth of the ten previous years. Hence, late frosts are considered as important ecological events that strongly affect beech vitality and competitiveness especially at high altitudes. Evidence of significant impacts on radial growth by late frosts distinct before leaf unfolding or with temperatures above ?3°C was not found. Also, increasing frequency and intensity of late frosts during recent decades were not ascertained. Hence, the recently observed decreased vitality of common beech accompanied by growth depressions especially at high altitude sites in Central Europe cannot be explained as a consequence of late frost damage.     

Bud dormancy in beech (Fagus sylvatica L.). Effect of chilling and photoperiod on dormancy release of beech seedlings

Two-year-old Fagus sylvatica L. seedlings were subjected to natural winter chilling or were overwintered in a heated greenhouse. Plants were then grown in controlled environment chambers with photoperiods of 9 or 13 h. Renewal of bud growth was found to be mainly determined by winter chilling. There was a slight interaction between chilling and photoperiod. Sprouting of apical buds took two to three times as long in unchilled plants as in chilled plants. Shoot elongation was influenced by chilling and was also greater in the 13-h photoperiod than in the 9-h photoperiod, but this may have been due at least in part to the higher irradiance. Chilling resulted in rapid dormancy loss and changed the growth pattern from basitonal to acrotonal.     

Biscogniauxia nummularia infecting beech (Fagus sylvatica) trees and sympatric plants of the sedge Carex brevicollis

Biscogniauxia nummularia is known for its association with beech (Fagus sylvatica), on which it occurs as a saprotroph and a pathogen causing strip cankers following water stress. This fungus has also been reported as a dominant endophytic species in plants of the sedge Carex brevicollis growing in the understory of beech forests and adjacent grasslands in Sierra de Urbasa (Navarre, Spain). In this area, stromata of B. nummularia were observed in dead and living wood of beech trees at several locations where plants of C. brevicollis also contained the fungus as an endophyte. Pure cultures obtained from stromata of B. nummularia on F. sylvatica trees were compared to endophytic isolates from symptomless C. brevicollis. Culture morphology and micromorphology as well as rDNA sequences of the internal transcribed spacer (ITS) regions were identical, suggesting that B. nummularia from beech can also live endophytically in C. brevicollis. It is unknown whether the endophytic strains of Carex might have a role as an inoculum source for the infection of beech trees, or whether they represent a dead end in the life cycle of the fungus.     

Genetic diversity and bark physiology of the European beech (Fagus sylvatica): a coevolutionary relationship with the beech scale (Cryptococcus fagisuga)

In 1994 and 1995, the degree of infestation by the beech scale (Cryptococcus fagisuga Lind.) was recorded on 120 beech trees (Fagus sylvatica L.) in the Pless Forest near G?ttingen, Germany. Simultaneously, the trees were characterized genetically and compounds of primary and secondary metabolism of beech bark were analyzed. A correlation was established between beech scale infestation and the genotype of the host trees, based on gene locus A of isocitrate dehydrogenase (IDH-A). The fraction of infested beech trees was higher in the heterozygous genotype A2A3 group than in the homozygous genotype A2A2 and A3A3 groups, whereas the fraction of beech trees with decreasing infestation from one year to the next was lower in the heterozygous genotype A2A3 group than in the homozygous genotype A2A2 and A3A3 groups. Concentrations of soluble carbohydrates, protein amino acids and proanthocyanidins were determined in the inner and outer bark of trees with differing degrees of infestation. The results indicate that the defense reaction of beech against infestation by beech scale comprises multiple processes in which nutrient availability to the beech scale is reduced by concentration shifts and by the formation of inhibiting compounds.     

Vulnerability of spruce (Picea abies) and beech (Fagus sylvatica) forest stands to storms and consequences for silviculture

The form and magnitude of storm damage and stand disclosure patterns were assessed in 332 randomly chosen pure and regular stands of spruce (Picea abies L.) and beech (Fagus sylvatica L.) after storm lothar, within a region of the Swiss Midlands. This data was analysed in relation to maximal wind speed, measured with Doppler radar techniques and other influential factors such as relief, allometric characteristics, silvicultural history, and neighbourhood. In addition, storm damage, assessed from aerial photographs over an extended perimeter (about 70,000 ha) was considered. A storm of the magnitude of lothar (December 26 1999), with an average maximal wind speed of 45 m s⁻¹ (160 km h⁻¹) appears to have a highly chaotic wind field structure, with great spatial and temporal variation of wind gusts. Wind speeds were not a significant predictor for damage in spruce stands and only weakly influential for beech. The consequences of this high randomness were analysed to estimate the return time of such a storm at the stand level. It lies between 86 and 113 years for spruce, 357 and 408 for beech. Only a few independent variables were significant and the overall explanatory strength of the model was unexpectedly low (R ²=0.07 for spruce and 0.30 for beech). Among the more reliable predisposing factors were mixture and aspect combined with gradient. An admixture of 10% or more broadleaved tree species or wind-firm conifers like Douglas fir [Pseudotsuga menziesii (Mirb.) Franco] significantly reduced the vulnerability of spruce stands (by a factor of more than three). On wind-exposed aspects, damage was more than twice the average. Steeper slopes caused a significant reduction in susceptibility (by a factor of six for slopes over 50%, in comparison to gentle slopes <20%). Other factors such as height to diameter ratio of trees or time since last thinning did not appear to be significant predictors.     

Transformation of even-aged European beech (Fagus sylvatica L.) to uneven-aged management under changing growth conditions caused by climate change

Transformation from even-aged to uneven-aged forest management is currently taking place throughout Europe. Climate change is, however, expected to change growth conditions—possibly quite radically. Using a deterministic approach, it was the objective of this study to investigate the influence of such changes on optimal transformation strategies for an even-aged stand of European Beech in Denmark. For a range of growth change scenarios, represented by changes in site index, optimal harvest policies were determined using a matrix modelling approach and a differential evolution algorithm. Transition probabilities were updated continuously based on stand level variables and the transition matrix was thus dynamic. With optimal transformation policies, stand development followed similar pathways during the transformation phase irrespective of climate change scenario. Optimal transformation policies were thus robust, suggesting that a good policy would work well under different outcomes of climate change, i.e., acting under erroneous assumptions about change would not lead to major economic loss. For the chosen case stand, the net present value (NPV) of the transformation phase (first 100 years) contributed 80–90 % of the total expectation value at a 2 % discount rate. To assess the robustness of the optimisation procedure and understand the nature of the response surface, 100 replications per scenario were carried out. Variation between replications peaked during the later stages of the transformation phase indicating several pathways for transformation, which were characterised by almost identical levels of profitability.