Bark phenolics of American beech (Fagus grandifolia) in relation to the beech bark disease1

The amount of total extractable phenols was determined for bark sections obtained from behind cankers naturally induced by Nectria coccinea var. faginata and from behind mechanically inflicted wounds on stems of American beech. Healthy bark from susceptible trees was found to contain a similar level of phenolics as bark from trees determined to be resistant to the beech bark disease. Six months after wounding, wound-altered bark from susceptible trees was found to be lower in phenols than woundaltered bark from resistant trees. Inoculation of wounds with N. coccinea var. faginata resulted in decreased phenolic levels in bark sections nearest the wound surface, and increased phenolic levels in sections nearest the vascular cambium, several mm distant. Phenolic levels in injured or infected bark appear to follow similar patterns as those resulting from injury or infection of xylem tissues.     

Amino nitrogen and phenolic constituents of bark of American beech,Fagus grandifolia,and infestation by beech scale,Cryptococcus fagisuga

Concentrations of amino acids, total amino nitrogen, and phenols, ratio of phenol to amino nitrogen, and pH in the bark of American beech, Fagus grandifolia, were determined to see if these chemical constituents were correlated with susceptibility of trees to infestation by the beech scale, Cryptococcus fagisuga. The relationship of these bark constituents with scale infestation levels (none, light, and moderate), tree size (large vs. small trees), and bark shading was determined. Levels of some individual amino acids and total amino acid content in moderately infested trees were significantly higher than in uninfested trees. Large uninfested beech trees, which tend to be more readily infested by scale, had significantly higher concentrations of aspartic acid. Concentrations of phenol in the outer bark of both infested and uninfested trees were significantly higher than in the inner bark. However in bark infected by Nectria, phenol levels in the inner bark were significantly higher than in the outer bark.     

Root sprouting in American beech (Fagus grandifolia): Effects of root injury,root exposure,and season

Superficial woody American beech roots were manipulated and left in situ for three growing seasons to evaluate seasonal influences of injury and exposure on root sprout formation. A total of 648 root segments were arranged in a 3² × 2 split-plot factorial experiment with three injury levels (scrape, cut, and control), two exposure levels (exposed and buried), and three seasons (summer, late fall, and spring). Analysis of variance revealed significant injury, season, exposure and interaction effects. Injured roots developed more callus, adventitious buds and sprouts than did controls. Exposure stimulated bud initiation, and spring injury resulted in higher levels of sprouting compared to late fall or summer injury.     

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.     

Sensitivity of red oak (Quercus rubra L.) and American beech (Fagus grandifolia Ehrh.) seedlings to sodium salts in solution culture

Sodium salt sensitivity of red oak (Quercus rubra L.) and American beech (Fagus grandifolia Ehrh.) was evaluated in solution culture. Both species showed symptoms of salt injury when grown in the presence of less than 10 mM Na. In red oak, leaf symptoms first appeared at a sodium concentration of 6.0 mM and leaf weight was significantly reduced at 7.5 mM Na. Leaf, stem and root dry weights of American beech were significantly reduced in the presence of 4.0 mM sodium. In both species, browning of leaf margins and necrosis were evident in the Na-treated plants. The observed symptoms were associated with high concentrations of sodium in the tissues. Neither species appears to have control over sodium uptake and translocation.     

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.     

Japanese beech (Fagus crenata) plantations established from seedlings of non-native genetic lineages

In order to examine whether seedlings of non-native genetic lineages had been planted, we compared chloroplast DNA haplotypes in plantations and neighboring natural populations of Fagus crenata in Nagano Prefecture, Japan. This region was chosen for study because there are abundant natural F. crenata populations with different haplotypes, and because the species is also frequently used for afforestation in the area. We sampled 159 trees from 30 populations of the species across most of its natural range in the region, and 136 trees from 20 plantations of the species, ranging in age from 3 to 17 years. Six single nucleotide polymorphisms and four insertions/deletions in two chloroplast DNA regions (trnL–trnF and trnK) were analyzed in each of the sampled individuals in order to determine their haplotypes. Four haplotypes (B, D, E and F) were detected in the natural populations, and these exhibited a clear geographical structure; in contrast, only two haplotypes (A and B) were found in the plantations. We found that 15 out of the 20 plantations—located in central and southern areas on the Pacific side of Honshu, where the natural populations contain haplotypes D, E, and F, and where the climate is characterized by dry, cold winters—had been established using seedlings with haplotypes A and B derived from the Sea of Japan side of the Japanese Islands, where it snows heavily. The risks associated with planting seedlings of inappropriate lineages are discussed.     

Spatial patterns and genetic structures within beech populations (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) of forked and non-forked individuals

Analyses of distribution patterns and genetic structures of forest stands can address distinct family structures and provide insights into the association of genetic and phenotypic variation patterns. In this study, point pattern analysis and spatial autocorrelation were used to examine the spatial and genetic structures in two naturally generated beech stands, which differ in age, trunk morphology, and stand management. Significant tree clumping was observed at distances up to 20 m in the young forest stand, whereas dispersion at distances under 10 m was observed in the old stand. The spatial analysis based on Ripley’s k function of the two different groups of trees showed that the non-forked trees match in both stands the spatial pattern of all trees while the forked were randomly distributed. Additionally, according to the bivariate analysis, forked trees in both stands were randomly distributed as related to non-forked tree positions. Finally, Moran’s I values were not very high, though significant genetic autocorrelation was identified at distances up to 20 m in the young stand, suggesting the existence of distinct family structures. However, no significant genetic structuring was observed in the old stand. Our findings suggest that spatial genetic patterns are impacted by stand age, environmental factors and human activities. The spatial distribution of forked trees was not clearly associated to family structures. Random effects and also micro-environmental variation could be additional factors explaining forking of beech individuals.     

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.     

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.     

Methods of inoculating Acer spp., Populus tremuloides, and Fagus grandifolia logs for commercial spalting applications

One of the most promising wood value-added processes currently under development is spalting, where pigment is added to wood via fungal colonization. Previous studies have shown laboratory level spalting to be achievable and highly predictable. However, large-scale spalting for potential commercial applications introduces a substantial number of additional variables which impact the spalting process. To test the potential of commercial-scale spalting, Acer saccharum, Fagus grandifolia, and Populus tremuloides logs were inoculated with multiple known spalting fungi utilizing both liquid spray cultures and live dowel pin cultures. Many of the fungi that successfully produce spalting in small, sterile cultures also produced significant amounts in large logs, with many spalting patterns identical to those found in small-scale testing. Pairings of Trametes versicolor/Scytalidium cuboideum and Xylaria polymorpha/Xylaria polymorpha (different isolates) produced significant amounts of zone lines. In addition, the method of inoculation impacted the amount of spalting: more zone lines were produced when fungi were introduced via plugs, while more stain was produced when liquid cultures were sprayed onto the logs. These results indicate that many of the standard spalting fungi are suitable for large-scale applications; however, the inoculation method appears to be a vital component for successful spalting under a restricted time schedule.     

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     

Over-mature beech trees (Fagus orientalis Lipsky) and close-to-nature forestry in northern Iran

Oriental Beech is the most important commercial tree species in northern Iran. In recent years wood production companies interested in felling large beech trees for profit have challenged advocates of close-to-nature silviculture who favor conservation. Our study objective was to assess the economic value of over-mature beech trees by relating tree diameter (DBH) to amount of decay. Based on the location of onset of decay, we categorized three types of decay as stump, stem, and crown decay. Trees of greater diameter (age) typically showed greater decay in the stem. Percent of decayed volume, diameter of decayed tissue, and length of decay in tree stems varied between 0.5%?64.3%, 15 cm?75 cm, and 2.0?19.5 m, respectively. With increasing trunk diameter, the proportion of truck decay increased. Red heart and dark red heart constituted 25% and 14.3% of sampled trees, respectively. However, we found no correlation between intensity of stem decay and morphological characteristics of trees. Seedlings were not abundant around the bases of over-mature trees, suggesting that the trees did not contribute to regeneration of the stand. Beech trees of diameter >1 m do not provide valuable round wood for industries and cause to raise wood production costs. We recommend that these trees >1 m DBH should be retained in forest stands because of their low commercial value but high ecological and conservational values such as maintaining biodiversity in forest ecosys-tems.     

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.     

Thioacidolytic analysis of residual lignins isolated from the chemical pulps of spruce (Picea abies) and beech (Fagus sylvatica) wood

Eight different residual lignins isolated from unbleached chemical pulps [sulfite, kraft, alkaline sulfite-anthraquinone-methanol (ASAM), soda/AQ/MeOH] of spruce and beech were characterized using gel permeation chromatography (GPC), thioacidolysis, and desulfurization to determine average molecular weight, amounts of uncondensed β-O-4 linkages, and dimeric linkage patterns, respectively. The total amounts of G-CHSEt-CHSEt-CH₂SEt and S-CHSEt-CHSEt-CH₂SEt were markedly reduced in residual lignins to 40% to 80% of the values for the corresponding milled wood lignins (MWLs). The number of dimeric units determined by thioacidolysis and desulfurization of the residual lignins was decreased by onehalf to one-fifth compared to the MWLs. Among the diverse types of dimeric units, reduction of β-1 and β-5 units was significant in most of the residual lignins, with the exception of beech ASAM residual lignin. Compared to beech MWL, 40% more C6–C3 main monomers were detected, whereas the relative composition of the dimeric units in the beech residual lignin was very similar to that in the beech MWL. The average molecular weights of residual lignins were less than those of the MWLs. However, the average molecular weights of the spruce kraft and soda residual lignins were determined to be higher than those of the corresponding MWLs.     

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.

     

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.     

Spatial and seasonal variations in photosynthetic properties within a beech (Fagus crenata Blume) crown

The light response curve and the intercellular CO₂ concentration response curve of CO₂ assimilation rate were investigated together with the light conditions at the four different heights within the beech crown from 1995 to 1997 on Mt. Fuji in Japan. On the seasonal fluctuation, the CO₂ assimilation rate at light saturated condition increased rapidly in May, and attained to the maximum between the end of June and July, thereafter, slightly decreased until the middle of August and rapidly decreased in September and October. The daily sum of photosynthetic photon flux density attenuated with deeping within the crown, and particularly, the relative value on 2nd position dropped to only 30%. TheA _max decreased from 10 to 5μmol m⁻² s⁻¹, approximately, with deeping within the crown. The light saturation point, quantum yield, light compensation point and dark respiration rate also varied with deeping. These results suggest that the photosynthetic properties vary gradually from sun to shade leaves along the light attenuation within a beech crown. At light saturated condition, the stomatal conductance and mesophyll conductance were strongly correlated withA _max among the four different heights (r > 0.96, respectively). TheC _i/C _a ratio was around 0.8, and there were no remarkable differences among the four different heights. These results suggest that the vertical gradient ofA _max depends on the variation of mesophyll conductance. The stomatal conductance may be also one of the major factor in the vertical gradient ofA _max. However the intercellular CO₂ concentration doesn’t influence the vertical gradient ofA _max within the crown. This work is supported by the Sasagawa Scientific Research Grant from The Japan Science Society and Grant-in-Aid for Scientific Research (C).     

Relationship between branch-scar parameters and knot features of oriental beech (Fagus orientalis Libsky)

The classification of roundwood is inextricably linked to the measurement of a particular single wood defect. The appearance, location, and number of defects are important in the quality evaluation of logs and sawn timber, and the most important defects are knots. The purpose of this study was to investigate the relationship between the appearance of branch scars and features of the related knot inside oriental beech logs, and to model the relationship between well-defined branch-scar and knot parameters. One hundred and fifty knots in 15 stems of oriental beech trees were studied. Image analysis software was used to measure the branch-scar and knot features. The results showed a significant positive correlation between the branch-scar parameter “moustache length” and the knot length. The ratio of branch-seal length to width was found to be a good estimator of the stem diameter at the time of knot occlusion and the amount of clear wood between the knot occlusion and the bark. The relationship obtained for the oriental beech stem radius at time of knot occlusion confirms relationship reported for European beech (Fagus sylvatica L.).