Genetic structure of populations of beech (Fagus sylvatica L.) in Denmark

The genetic structure in Danish indigenous and introduced provenances of beech (Fagus sylvatica L.) is described. Ten stands are analysed based on variation revealed by seven enzyme systems. The present study reveals significant allele frequency differences between populations with a small absolute amount of genetic differentiation, with no pattern related to geographic distance. The genetic differentiation between indigenous populations measured by Wright's F _ST‐statistic does not deviate significantly from zero, and only 2.9% of the total genetic variation in the foreign populations was due to interpopulation differences, The allele frequencies in relatively small and isolated populations do not differ from the frequencies found in larger areas with beech forest. The low genetic differentiation of isozyme level between indigenous and foreign populations was opposed by significant differences in foliation performance measured on three pairs of neighbouring stands. Foliation performance in adjacent stands of indigenous and introduced beech indicates that quantitative characteristics must be considered separately from neutral variation at single loci.     

Radial distribution of carbohydrate reserves in the trunk of declining European beech trees (Fagus sylvatica L.)

Context

European beech (Fagus sylvatica L.) is considered threatened by anticipated climate change, but the physiological causes of potential beech decline or mortality remains poorly understood.

Aims

The purpose of the present study was to fuel debate about the assumption that carbohydrate depletion is involved in the decline of mature European beech.

Methods

The health status of beech trees from a severely declining stand was visually assessed by examining their crown condition. Content and radial distribution of non-structural carbohydrates (starch and soluble carbohydrate) were analyzed in the trunks and compared to those reported earlier in trunks of healthy beech trees.

Results and discussion

The distribution of carbohydrate in the beech trunks recorded here seemed affected by decline. We found a stronger radial decrease of starch content than those reported earlier for healthy beech trees. Carbohydrate reserves appear partially maintained in the outermost rings while starch depletion occurred in older wood rings in declining trees that may be able to mobilize carbohydrate reserves from older wood rings in response to successive climatic constraints.     

Growth response of advanced planted European beech (Fagus sylvatica L.) after storm-caused loss of shelterwood

European Journal of Forest Research - Outcomes after an abrupt, storm-caused loss of spruce shelterwood (Picea abies L. Karst.) overstories on advanced planted beech (Fagus sylvatica L.) were...     

Genetic variation of beech (Fagus sylvatica L.) in Rodopi (N.E. Greece)

The recent taxonomic classification of beech in Europe considers existence of one species (Fagus sylvatica L.) with two subspecies: F. sylvatica ssp. sylvatica and F. sylvatica ssp. orientalis. Four beech populations growing on the Greek part of the Rodopi Mountains were studied using morphological traits as well as DNA molecular markers (AFLPs and chloroplast DNA SSR). The aim of the study was to describe the variation patterns of beech in the Rodopi Mountains and to test the hypothesis of possible introgression between the beech subspecies’ sylvatica and orientalis in this area. Both morphological traits and gene markers revealed a possible influence of F. orientalis on the east side of Rodopi and at the low elevations, while characters resembling F. sylvatica were observed mainly on the western part of the mountains and in higher altitudes. There was a clinal increase of genetic diversity from the west to the east, reaching a level firstly reported for beech populations. These results can be explained either by the existence of a main refugial area for beech during the last glaciation or by the occurrence of a recent hybridization among the subspecies, which were spatially isolated during the last glaciation and came into reproductive contact during their postglacial remigration. These two scenarios are not necessarily mutually exclusive.     

Identification of viruses from European beech (Fagus sylvatica L.) of declining forests in Northrhine-Westfalia (FRG)

Five different viruses isolated from degenerating European beech of different forest sites of West Germany could be identified as cherry leaf roll virus, brome mosaic virus, potex and potyviruses.     

Biomass, basic density and biomass expansion factor functions for European beech (Fagus sylvatica L.) in Denmark

The objective of this study was to analyse the within-tree allocation of biomass and to develop biomass functions for above- and below-ground components of European beech in Denmark. Separate functions were developed for stem, branches, below-ground stump and root system, total above-ground biomass and total tree biomass. For each of these components or aggregate components, models were also developed for the average basic density of wood and bark. To enhance the versatility of the models, a function for estimating the biomass expansion factor (BEF) was also developed. The functions were based on 66 trees measured for total biomass. Model performance was evaluated based on 74 trees measured only for above-ground biomass. The trees were sampled in 18 different forest stands covering a wide range of tree sizes and stand treatments. Models were estimated using a linear mixed-effects procedure to account for within-stand correlations. The functions for biomass and BEFs included only diameter at breast height and total tree height for individual trees as predictor variables. Inclusion of additional variables reflecting site quality or stand density did not improve model performance. The functions for basic density included individual tree diameter, tree height and quadratic mean diameter as predictor variables, indicating an effect of stand density on the basic density of wood and bark.     

Modelling natural drying of European beech (Fagus sylvatica L.) logs for energy based on meteorological data

A key factor for improving the whole fuel wood supply chain is moisture management. Moisture affects material quality, transport costs and revenue. The aim of this study was to develop a drying model for piled European beech (Fagus sylvatica L.) logs based on meteorological data, employing the continuous weighing approach. Two drying cycles with three piles altogether were conducted. In the first one (February to October 2013) the moisture content of the pile dropped from 43.5% to 22.5%, whereas in the second one (December 2013 to March 2015), the moisture content of the 2 piles dropped from 42.1% and 40.5% to 20.6% and 19.8%, respectively. Model performance (R² adj?=?0.41) was validated by applying the model developed from the second cycle's data on the first cycle's data. Model's mean deviation in moisture content from the observed curve was ?0.41%?±?0.72%. Economic benefits and environmental impact were highlighted. Calorific value increased from 2.65 to 3.57?kWh?kg^–1 from February to October 2013. Truckload volume utilization increased by 25.1% and transported calorific value by 48.0%. The number of truck trips for transportation was reduced from nine to seven.     

Xylem traits in European beech (Fagus sylvatica L.) display a large plasticity in response to canopy release

Key message

The position of trees in the canopy impacts xylem structure and its inter-annual variation. After canopy release, the increase in the hydraulic conductivity of growth rings was driven by an increase in radial growth in large trees, and by both an increase in radial growth and changes in xylem structure in saplings.

Context

Forest canopies are frequently subjected to disturbances that allow understory trees to access the upper canopy. The effect of canopy release on xylem anatomy has been assessed in juvenile trees and saplings, while the potential acclimation of larger trees remains poorly documented.

Aims

We estimated the potential hydraulic conductivity of growth rings in large understory trees compared to overstory trees, and evaluated the responses to canopy release in large trees and in saplings.

Methods

We recorded radial growth, wood density, and vessel structure in beech trees according to their position within the canopy and their size. Xylem traits were followed during 6 years after canopy release for large trees, and during 2 years for saplings. Vessel diameter and frequency as well as ring area were used to compute the potential annual ring hydraulic conductivity.

Results

Large understory trees displayed lower radial growth increments and lower potential annual ring hydraulic conductivity than overstory trees. After canopy release, potential annual ring hydraulic conductivity increased in large trees, due exclusively to increased radial growth without any change in specific hydraulic conductivity. It increased in saplings due to both increased radial growth and increased specific conductivity.

Conclusion

Tree size impacted xylem structure and resulted in plasticity of the potential hydraulic conductivity of the annual tree ring following canopy release.

     

DNA sequence variation and development of SNP markers in beech (Fagus sylvatica L.)

European beech (Fagus sylvatica L.) is one of the most important deciduous tree species in Central Europe. The potential of beech to adapt to climate change, higher temperatures, and less precipitation in the summer months is still unknown. Most studies in beech used microsatellite, AFLP (amplified fragment length polymorphism), or isozyme markers, which have only a restricted potential to analyze adaptation. Only few studies investigated genes probably involved in the adaptation to drought stress and bud phenology in beech. In this study, SNP (single nucleotide polymorphisms) markers were developed in order to analyze adaptation and their technical advantages compared to microsatellites and AFLPs were discussed. Partial sequences of ten candidate genes probably involved in drought stress and/or bud phenology were identified at the genomic level, and SNPs and indels (insertions/deletions) in coding and non-coding regions were analyzed. Plant material was sampled along a precipitation gradient in Germany. In total, 8,145?bp were sequenced and analyzed, 4,038?bp were located in exon and 4,107?bp in intron regions. 63 SNPs and 11 indels were detected, which are differently distributed over the studied gene regions. The nucleotide diversity ranged from 0 to 6.62 (π?×?10^?3) and is comparable to other tree species, whereas the mean nucleotide diversity (2.64) for F. sylvatica is comparatively low. These results will help to investigate the genetic basis of drought stress and bud burst and to conduct association mapping in natural populations. Furthermore, the detected SNPs can also be used for population genetic studies.     

Interspecific competition impacts on the morphology and distribution of fine roots in European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst.)

Morphology and vertical distribution patterns of spruce and beech live fine roots (diameter ≤2 mm) were studied using a soil core method in three comparable mature stands in the Solling: (1) pure beech, (2) pure spruce and (3) mixed spruce–beech. This study was aimed at determining the effects of interspecific competition on fine root structure and spatial fine root distribution of both species. A vertical stratification of beech and spruce fine root systems was found in the mixed stand due to a shift in beech fine roots from upper to lower soil layers. Moreover, compared to pure beech, a significantly higher specific root length (SRL, P<0.05) and specific surface area (SSA, P<0.05) were found for beech admixed with spruce (pure beech/mixed beech SRL 16.1–23.4 m g⁻¹, SSA 286–367 cm² g⁻¹). Both indicate a flexible ‘foraging’ strategy of beech tending to increase soil exploitation and space sequestration efficiency in soil layers less occupied by competitors. Spruce, in contrast, followed a more conservative strategy keeping the shallow vertical rooting and the root morphology quite constant in both pure and mixed stands (pure spruce/mixed spruce SRL 9.6/7.7 m g⁻¹, P>0.10; SSA 225/212 cm² g⁻¹, P>0.10). Symmetric competition belowground between mixed beech and spruce was observed since live fine roots of both species were under-represented compared to pure stand. However, the higher space sequestration efficiency suggests a higher competitive ability of beech belowground.     

Soluble N compound profiles and concentrations in European beech (Fagus sylvatica L.) are influenced by local climate and thinning

We assessed seasonal changes of total soluble nonprotein nitrogen compounds (TSNN) in adult European beech trees (Fagus sylvatica, L.) growing under different local climate during the growing season immediately following a thinning treatment and 3 years later. In both years, samples of leaves, xylem sap and phloem exudates from beech trees growing in thinned and unthinned (control) stands on a dry, warm SW exposed and a cooler, moist NE exposed site were collected in May, July and September. In May of both years, asparagine (Asn) and glutamine (Gln) were most abundant in leaves and xylem, respectively, whereas arginine (Arg) dominated in the phloem. In July, TSNN concentrations decreased in all tissues and sites, but differences in water availability between aspects were reflected in TSNN concentrations. In September, differences in the increase of Arg concentration in the phloem were related to differences in the onset of senescence between treatments. Thinning treatment increased amino compound concentrations of beech tissues in July on both aspects, particularly at the NE thinned site. It is supposed that, the N balance of adult beech is favoured by both, the thinning treatments as well as the cool-moist climate prevailing at the NE aspect.     

Exploratory study on the impregnation of Scots pine sapwood (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) with different hot melting waxes

Scots pine sapwood (Pinus sylvestris L.) and beech (Fagus sylvatica L.) were impregnated with five waxes. The experiments indicate deep penetration into pine sapwood. Besides the viscosity, an influence of the wax polarity is presumed. Wax penetrates pine wood deeply via the cross-section, but not sufficiently enough to impregnate longer construction elements. However, the radial wax uptake exceeds the uptake via the tangential orientation and guarantees complete soaking of the sapwood tissue. The lateral wax penetration within beech is quite low and irregular. In addition to the temperature, a prolonged process procedure is decisive for an increasing wax uptake. As such, beech wood vessel elements seem to be fully impregnable via the longitudinal surface after a longer process procedure.     

A recent growth increase of European beech (Fagus sylvatica L.) at its Mediterranean distribution limit contradicts drought stress

Future changes in tree growth, associated with a warmer and drier climate, are predicted for many species and locations across the European Mediterranean Basin. However, quantification of the intensity and severity of related consequences for forest ecosystem functioning and productivity remains challenging. Species-specific distribution limits that are particularly sensitive to small changes in the ambient climate may provide an ideal test bed to assess the nature of past growth trends and extremes and their responsible controls. Here, we seek to understand how twentieth century climate change affected the growth of European beech (Fagus sylvatica L.) nearby its south-eastern distribution limit in Albania and Macedonia on the Balkan Peninsula. We sampled 93 living trees from undisturbed mixed forest stands at ~1,450 m a.s.l. and 29 timbers from nearby historical buildings. Application of different tree-ring detrending techniques allowed robust composite chronologies with varying degrees of high- to low-frequency variability to be developed back to 1648 ad. Comparison with local meteorological station measurements and continental grid-box climate indices revealed spatiotemporal instability in growth–climate response patterns. Nevertheless, year-to-year and decadal-long fluctuations in radial beech growth were significantly (P < 0.001) negatively correlated at ?0.61 with June–September temperature over the 1951–1995 period. This (inverse) relationship between increased beech growth and decreased summer temperature is somewhat indicative for the importance of plant-available soil moisture, which likely controls ring width formation near the species-specific south-eastern distribution limit. Significant positive correlations between beech growth and drought (scPDSI; r = 0.57) confirm metabolistic drought constraints. However, an unexpected late twentieth century growth increase not only contradicts the previously observed growth dependency to summer soil moisture, but also denies any putative drought-induced forest ecosystem suppression in this part of the Mediterranean Basin.     

Identification of moisture-induced stresses in cross-laminated wood panels from beech wood (Fagus sylvatica L.)

The crosswise bonding of the layers in laminated solid wood panels results in internal stresses when the humidity varies. The layers hinder one another as a result of the anisotropy of wood. The purpose of this study was to determine the internal stress state in free and constrained swelling. The expansion properties in the three panel directions were measured. Furthermore, the swelling of samples was constrained while the resulting forces were recorded. Hygroscopic warping experiments were carried out inducing a climate gradient within the panels. Afterwards the stresses were calculated from released deformations and non-destructive measurements of the Young’s modulus. The materials used were untreated and heat-treated beech wood, the latter modified in two levels. In addition to homogenously structured panels, treated top layers were combined with an untreated middle layer. Swelling, swelling pressure, warping and internal stresses considerably decreased from untreated to treated wood. If layers from treated and untreated material were combined, stresses and deformations increased as compared to the variants produced only from treated wood. It was concluded that the lower equilibrium moisture content of heat-treated beech wood improves its dimensional stability, which results in smaller deformation differences between the layers. Hence, the stresses were less distinctive.     

Seasonal time-course of the above ground biomass production efficiency in beech trees (Fagus sylvatica L.)

Key message

In order to record the seasonal changes in aboveground biomass production (trunk and branches) in a forest, changes in wood density must be taken into account. A 60-year-old beech forest displayed a large intra-annual variability in its aboveground woody biomass production efficiency. This variation followed a seasonal trend with a maximum during the summer while gross primary production was rather low.

Context

In the current context of land use and climate change, there is a need to precisely quantify the carbon (C) balance of forest ecosystems, and more specifically, of C allocation to tree compartments.

Aims

We quantified the seasonal changes in the aboveground biomass production (aBP) of a beech forest growing on two different soils: an alocrisol and a calci-brunisol. In addition, for the alocrisol ecosystem, we assessed the existence and degree of intra-annual variability in the ratio of wood aBP to gross primary production (GPP), i.e., the wood aBP efficiency.

Methods

The study site is a 60-year-old beech forest in northeastern France. An eddy covariance tower records continuously net ecosystem exchange. To investigate the temporal changes in aBP, mini-cores were drilled and diameter at breast height measurements were taken on a monthly basis from 45 trees for both stands studied over 2014.

Results

A clear difference in aBP was observed between the two soils with the alocrisol being more productive than the calci-brunisol. For the alocrisol, both woody aBP and GPP changed over the course of the year, reaching peak values during June (6 and 12.5 gC m^?2 day^?1, respectively). Wood applied bias photon-to-current efficiency aboveground Biomass Production Efficiency (aBPE) also showed important intra-annual variations, ranging from 0.09 in September to 0.58 in July. Wood density varied throughout the year, and not taking it into account would have led to an overestimation of aBP by as much as 20% in April and May.

Conclusion

Our study highlights the importance of taking wood density into account for intra-annual studies of aBP. Wood aBPE cannot be considered as constant as it fluctuated from 0.09 to 0.58 throughout the year for an annual value of 0.34. The potential error in wood aBPE stemming from not taking these changes into account amounts to 15%.

     

Effects of weather conditions on mast year frequency in beech (Fagus sylvatica L.) in Sweden

During the last two decades, an increase of the frequency ofnaturally regenerated beech seedlings has been reported. Thismay be due to an increased masting. In this investigation insouthern Sweden, mast year frequency, mast crop size and therelationships between mast year and climatic variables werestudied. Our analysis shows that while the average mast yearinterval was 4–6 years from the end of the seventeenthcentury up to the 1960s, the mean interval has decreased to2.5 years during the most recent 30 years, and there have beentwo consecutive mast years on two occasions during this laterperiod. Mast years have often followed years in which the temperaturein July and September was higher than the 30-year mean. However,there were significant variations in the amounts of beechnutsproduced between different mast years, and beechnut productionincreased with increasing site index. Climatic changes, especiallyincreases in temperature, may have been responsible for thehigher frequency of mast years, but increased atmospheric nitrogendeposition may also have been a contributory factor.     

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.     

Climate-sensitive modelling of site-productivity relationships for Norway spruce (Picea abies (L.) Karst.) and common beech (Fagus sylvatica L.)

The aim of the presented research project is to fit a site index model capable for predicting changes in site-productivity in a changing climate. A generalized additive model is used to predict site index as a function of soil and climate variables. The climate parameter values are estimated using the regional climate model WETTREG, based on global climate simulations with the global circulation model ECHAM5/MPI-OM for the reference period from 1961 to 1990. The climate values are further regionalized on a 200 m × 200 m grid. The generalized additive model quantifies the partial linear and non-linear effects of the predictor variables on site index. The model is parameterized for Norway spruce (Picea abies (L.) Karst.) and common beech (Fagus sylvatica L.) in Lower Saxony, Germany. Two case studies investigate the model's ability to generate information in order to support forest management planning decisions under a changing climate. One example analyzes the possible shift in site index of spruce along a precipitation gradient under the International Panel on Climate Change (IPCC) emission scenario A1B in the period from 2041 to 2050. The other case study shows possible future changes in site index of beech along a temperature gradient.