Seasonal changes in the axial distribution of peroxidase activity in the xylem sap of beech (Fagus sylvatica L.) trees

Xylem sap was collected from trunk segments of adult beech (Fagus sylvatica L.) trees by water displacement. Peroxidase activity was analyzed in xylem saps collected in different phases of the yearly growth cycle and from different heights up the trunks (up to 14 m). The xylem saps contained two major peroxidase isozymes with acidic isoelectric points of 4.1 and 4.6, respectively. Mean peroxidase activity was low during the emergence of the new leaves and high in summer and in winter. In the cold season, peroxidase activity decreased from the stem base to the top, whereas significant gradients were not observed during the vegetative period.     

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.     

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.     

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.     

Xylem sap composition of beech (Fagus sylvatica L.) trees: seasonal changes in the axial distribution of sulfur compounds

During different phases of the annual growth cycle, xylem sap was collected from trunk segments of adult beech (Fagus sylvatica L.) trees by the water displacement technique. Irrespective of the height of the trunk, both sulfate and reduced sulfur compounds were detected in the xylem sap throughout the year. Sulfate was the predominant sulfur compound in all samples analyzed. Its concentration in the xylem sap varied between 10 and 350 micro mol l(-1), with highest concentrations in April, shortly before bud break. In contrast to other tree species, cysteine and not glutathione was the predominant thiol transported in the xylem sap of beech trees. The cysteine concentration ranged between 0.1 and 1 micro mol l(-1). As observed for sulfate, maximum cysteine concentrations were found in April. Apparently, both sulfate and cysteine transport contribute to the sulfur supply of the developing leaves. Seasonal changes in the axial distribution of cysteine and sulfate differed, indicating differences in the source-sink relations of these sulfur compounds. High, but uniform, xylem sap sulfate concentrations in April may originate from balanced sulfate uptake by the roots, whereas high cysteine concentrations in April, increasing with increasing height of the trunk, may originate in part from protein breakdown in the trunk. Reversal of the axial distribution of xylem sap cysteine in late summer-early fall to higher concentrations in the lower part of the trunk than in the upper part of the trunk suggests that the upper part of the trunk becomes a sink for cysteine as a result of the synthesis of storage proteins at this time of the year.     

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.     

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.

     

Investigations on viruses from declining beech trees (Fagus sylvatica L.) in Rhineland and Westfalia,Federal Republic of Germany

Viruses of the Poty-, Potex- and Tobamo-virus groups, and presumably Nepoviruses could be isolated from degenerating beech trees (Fagus sylvatica L.) in West-Germany. They are discussed as predisposing factors of tree decline.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.