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.     

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.     

Genetic structure and variability of phenological forms in the European beech (Fagus sylvatica L.)

? Microsatellite markers were used to describe the genetic structure and variability of early, intermediate and late phenological forms of European beech (Fagus sylvatica L.). Two hundred and seventy individuals from three populations located in southern Poland were divided into three forms according to the phenological criterion — bud burst, and analyzed for allelic variation at five highly polymorphic microsatellite loci.

? Population differentiation was moderate and differed significantly among phenological forms. Average values of F _ST and R _ST decreased across phenological forms and amounted to F _ST values of 0.135, 0.110 and 0.108 and R _ST values of 0.365, 0.231 and 0.098 for early, intermediate and late forms of beech, respectively.

? Analysis of Molecular Variance (AMOVA) revealed different genetic structures characteristic of respective phenological forms of beech. The amount of within-population variability increased with the delay of the beginning of vegetation and amounted to 64%, 77% and 90% of total variability, depending on phenological form. A similar trend was found in average pairwise genetic distance between individuals belonging to a given phenological form (11.78, 11.85 and 12.22, from early to late forms).

? Our results demonstrate the importance of late spring frosts as a factor influencing the genetic structure of beech, and as a cause of the decrease in genetic variability as well as the increase in population differentiation proportional to the degree of phenological earliness.

     

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.     

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.     

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.     

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.     

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%.

     

Impacts of drought on mineral macro- and microelements in provenances of beech (Fagus sylvatica L.) seedlings

Beech seedlings originating from 11 German provenances with different climatic conditions were grown in pots and cultivated in a greenhouse. The composition of macro- and microelements in roots, axes and leaves was measured after half of the seedlings were subjected to a simulated summer drought. The recently described sensitivity of these provenances to drought was compared with drought-mediated changes in the elemental and ionic composition in organs of the seedlings; in addition, partitioning between roots and shoots was evaluated. A number of element concentrations were decreased in roots due to drought (K 94% of control, Mg 94%, Mn 75% and Zn 85%). However, chloride concentration increased in all organs (115-125%) and was the only element affected in leaves. Some changes in ionome can be related to sensitivity of provenances, but it is difficult to decide whether these changes are a result of, or a reason for, drought tolerance or sensitivity. Observed increases in chloride concentration in all plant parts of drought-treated beech seedlings can be explained by its function in charge balance, in particular since the level of phosphate was reduced. As a result of chloride accumulation, the sum of added charges of anions (and cations) in water extracts of leaf and root material was similar between drought and control plants. Since only the partitioning of Ca and Al (both only in axis) as well as Mn was affected and other elements (together with previously observed effects on C, N, S and P) remained unaffected by drought in all provenances, it can be concluded that direct effects by means of mass flow inhibition in xylem and phloem are unlikely. Secondary effects, for example on the pH of transport sap and the apoplastic space, cannot be excluded from the present study. These effects may affect partitioning between the apoplast and symplast and therefore may be significant for drought sensitivity.     

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...     

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.     

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.     

A sixteen-year record of regional and temporal variation in the fruiting of beech (Fagus sylvatica L.) in England (1980-;1995)

Annual production of mast by the beech is highly variable, yetthere are few long-term records of the extent of masting whichmight help to explain the masting phenomenon. In this 16-yearstudy, beech mast has been sampled from 100 trees by 7-min samplescollected from the ground and from low-growing branches at sitesranging from the south to the north of England, and varyingfrom closed canopy beechwood to shelter belts, avenues and isolatedtrees in parks. Variation in production of viable seed is much greater fromyear to year than from north to south. Good masting occurredin 1980, 1982, 1984, 1990 and 1995: in these years at leastone year of very bad masting normally followed. Otherwise, predictionof masting is unreliable. No full mast was found in 1981. Overallthere was very little full mast in five other years, but evenin these years, some sites or some trees produced an appreciableamount. All trees produced empty pericarps as well as full mast, especiallyon lower or shaded branches. Isolated trees produced more emptypericarps than trees in groups; this is likely to result froma lack of cross pollination. It is concluded that ground samplingfor a fixed period affords a rapid and effective method of assessingmasting despite the factors intervening between mast sheddingand collection.     

Ring width and element concentrations in beech (Fagus sylvatica L.) from a periurban forest in central Belgium

The Forêt de Soignes is a beech high forest located near Brussels (Belgium), established on a strongly acidic soil and subjected to atmospheric pollution and recreational pressure. We investigated variations in ring width and mineral element concentrations (N, P, K, Ca, Mg, Mn, Al) over the last 95 years in five 135-year-old trees, variations in ring width in four 40-year-old trees and tested associations with climatic parameters. Growth curves showed a striking increase in mean sensitivity in the last 20 years in the old trees but not in the young ones, starting with the 1976 summer drought. Mean sensitivity is a statistical measure of the mean relative variability between adjacent ring widths within a tree, which is correlated to susceptibility to climatic stress (Fritts, H.C., 1976. Tree Rings and Climate, Academic Press, London, 567pp). May rainfall and, to a lesser extent, soil water recharge (i.e. pooled rainfall from October to May) correlated significantly with ring width. However, two of the five growth depressions in the last 20 years could not readily be accounted for by adverse climatic conditions. Element concentration profiles were suggestive of a decrease in the availability of Mn, Mg and Ca, but no trend of increasing N was obvious. The possible causes of the recent increase in sensitivity of old beech trees are discussed in terms of interactions between ageing and extreme climatic events, aggravated by soil compaction due to logging traffic.     

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.     

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.     

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.     

Phenological phases of common beech (Fagus sylvatica L.) and their dependence on region and altitude in Southern Germany

Long-term observational data of the phenological network of the German Weather Service (DWD) were used to derive the altitudinal dependence of leaf unfolding (LU), leaf discolouration (LD) and the length of the vegetation period (LD-LU) of common beech (Fagus sylvatica L.) in different regions of Southern Germany. Depending on altitude, the vegetation period starts in Bavaria between the 2nd and 24th May with a delay of around 2 days per 100 m altitude increase (averaged over the last 30 years). In Southern Bavaria, LU starts a few days earlier as well as increases slightly faster into higher altitudes than in the northern regions. This can be explained by a more favourable supply of warmth and radiation in the south. The end of the vegetation period, defined here as the beginning of LD, showed no altitudinal dependence, which reflects an interaction of endogenous and exogenous factors as triggers for LD. In the Bavarian Forest region, altitudinal dependences are strongly influenced by sites which are affected by cold-air tailback (‘Kaltluftstau’) was obtained. Hence, phenological gradients should only be derived and applied above a phenological inversion zone. The calculated vertical gradients for LU can be used together with available observational data to estimate the beginning of the forest vegetation period of beech for different altitudes of Bavaria.