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.     

欧洲山毛榉干燥工艺初探

通过试验研究探索出在国产蒸汽加热干燥窑上干燥欧洲山毛榉的干燥工艺,该工艺具有处理后干燥周期短、开裂和变形小的特点,达到了国家干燥锯材一级标准.     

Response of Fagus sylvatica Seedlings to Root Trenching of Overstorey Picea abies

In a trenching experiment the impact of root competition by overstorey Norway spruce on growth and allocation of above-ground woody biomass of advanced planted beech was investigated in two stands over two growing seasons after trenching. Measurements of soil water potential were taken inside and outside trenched areas by tensiometers at 16 measuring points in one stand from May to September 1999. Height and diameter of the seedlings, as well as branch length and diameter, were measured of beech growing in trenched and untrenched conditions in September before trenching (1998) and afterwards (1999 and 2000). Tensiometers inside the trenches reached the expiring value of -0.085 Mpa 30-40 days later, or never, compared with those in untrenched areas. Whereas relative growth rate (RGR) in height was not affected by trenching, RGR both in diameter and in estimated dry weight of the main stem were substantially enhanced in the first growing season after trenching (1999). A different result was obtained in 2000, when RGR in seedling diameter and estimated dry weight of the main stem was slightly reduced by trenching. The only variable that showed increased RGR of the trenched seedlings in both years was the estimated dry weight of the branches, indicating different above-ground biomass allocation patterns of seedlings with and without root competition.     

Effects of drought and canopy ozone exposure on antioxidants in fine roots of mature European beech (Fagus sylvatica)

We quantified ascorbate, glutathione and alpha-tocopherol in fine roots of mature Fagus sylvatica L. under free-air canopy ozone (O(3)) exposure (twice ambient O(3) concentration, 2x[O(3)]) during two growing seasons that differed in the extent of summer drought (exceptional drought year 2003, average year 2004). This design allowed us to test whether O(3) exposure or drought, or both, affected root antioxidants during the growing season. In both years, root ascorbate and alpha-tocopherol showed a similar relationship with volumetric soil water content (SWC): ascorbate concentrations on a root dry mass basis increased from about 6 to 12 micromol g(-1) when SWC dropped from 25 to 20%, and a-tocopherol increased from 100 to 150 nmol g(-1) at SWC values below 20%. Root glutathione showed no relationship with SWC or differences between the dry and the average year, but it was significantly and consistently diminished by 2x[O(3)]. Our results were inconclusive as to whether shoot-root translocation of glutathione or glutathione production in the roots was diminished. Phloem glutathione concentrations in the canopy remained constant, but reduced transport velocity in the phloem and, as a consequence, reduced mass flow of glutathione cannot be ruled out.     

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.     

Early infection of Fagus sylvatica by Heterobasidion annosum sensu stricto

Heterobasidion annosum sensu stricto is the most important damaging agent in Scots pine stands planted on the former agricultural lands in Poland. The routine action in pine stands which have lost stability because of H. annosum root rot is to change stand management, including species conversion. In many cases, the Fagus sylvatica is used for this purpose. This study was the first assessment of widespread infection by H. annosum in young F. sylvatica plantations. Disease symptoms included atrophy and yellowing of leaves, wilting and the presence of pathogen sporocarps around the root collars of young trees. Heterobasidion annosum s. s. was observed on both 4‐ and 17‐year‐old beech. Based on annual increments, the disease could be present for 3–4 years before tree death. A high incidence of H. annosum in pine stumps of previous stands (80–100%) and dry periods in recent years may be the main reasons for such common infection of F. sylvatica. This work also showed that mice and frost were not the main killing factors F. sylvatica in plantations.     

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.     

Mechanisms of xylem recovery from winter embolism in Fagus sylvatica

Hydraulic conductivity in the terminal branches of mature beech trees (Fagus sylvatica L.) decreased progressively during winter and recovered in the spring. The objective of this study was to determine the mechanisms involved in recovery. Two periods of recovery were identified. The first recovery of hydraulic conductivity occurred early in the spring, before bud break, and was correlated with the occurrence of positive xylem pressure at the base of the tree trunk. Active refilling of the embolized vessels caused the recovery. The second recovery of hydraulic conductivity occurred after bud break and was correlated with the onset of cambial activity. Formation of new functional vessels, leading to an increase in xylem diameter, was largely responsible for the increase in xylem conductivity. The two mechanisms were complementary: active refilling of embolized vessels occurred mostly in the root and the trunk, whereas formation of new functional vessels occurred mainly in young terminal shoots.     

Dormancy breaking in Fagus sylvatica seeds is linked to formation of abscisic acid-glucosyl ester

New Forests - Seed dormancy is an adaptive mechanism that allows seed germination under suitable environmental conditions. Germination of stored dormant seeds proceeds after dormancy breaking...     

Growth and survival of Fagus sylvatica seedlings in relation to light intensity and soil water content

Previous investigations have shown different growth and root/shoot ratio increases of beech seedlings (Fagus sylvatica) with increased light intensity. In the present investigation both light intensity and soil water content were regulated on four levels in a factorial experiment. At the highest level of soil water content the seedling growth increased linearly with light intensity while the root/shoot ratio decreased with light intensity. At low levels of soil water content both the growth and root/shoot ratio slightly increased with increased light. The soil water content was also affected by the light intensity indicating that the soil water content may have caused unobserved effects in responding to increased light in previous investigations. It is concluded that both light intensity and soil water content should be quantified or manipulated even if only the effect of one of the two growth factors are investigated.     

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.     

Root distribution of Fagus sylvatica in a chronosequence in western France

The distribution of fine (<2 mm diameter) and small roots (2–20 mm diameter) was investigated in a chronosequence consisting of 9-year-old, 26-year-old, 82-year-old and 146-year-old European beech (Fagus sylvatica) stands. A combination of trench wall observations and destructive root sampling was used to establish whether root distribution and total biomass of fine and small roots varied with stand age. Root density decreased with soil depth in all stands, and variability appeared to be highest in subsoil horizons, especially where compacted soil layers occurred. Roots clustered in patches in the top 0–50 cm of the soil or were present as root channels at greater depths. Cluster number, cluster size and number of root channels were comparable in all stands, and high values of soil exploitation occurred throughout the entire chronosequence. Overall fine root biomass at depths of 0–120 cm ranged from 7.4 Mg ha⁻¹ to 9.8 Mg ha⁻¹, being highest in the two youngest stands. Small root biomass ranged from 3.6 Mg ha⁻¹ to 13.3 Mg ha⁻¹. Use of trench wall observations combined with destructive root samples reduced the variability of these estimates. These records showed that variability in fine root distribution depended more on soil depth and edaphic conditions than on stand age, and suggest that trench wall studies provide a useful tool to improve estimates of fine root biomass.     

Morphological and physiological responses to drought stress of European provenances of Scots pine

Increased frequency and intensity of drought episodes as a consequence of current and predicted climatic changes require an understanding of the intra-specific variability in structural and physiological characteristics of forest trees. Adaptive plasticity and genotypic variability are considered two of the main processes by which trees can either be selected or can acclimate to changing conditions. We tested for the relative importance of genotypic variability, phenotypic plasticity and their interaction by comparing the growth and physiological performance of 15 provenances of Scots pine (Pinus sylvestris L.), under two contrasting irrigation regimes. Selected provenances representing the distribution range of the species in Anatolia, Turkey, were contrasted with seed sources spanning the range from Spain to the UK, in Europe. We found a strong latitudinal differentiation among the 15 provenances for survival after drought, largely the result of the higher mortality of some western and central European provenances. Differentiation in diameter and height growth was also clear with the worst provenance coming from Western Europe (UK). Among the Turkish provenances, the more extreme southern high-elevation populations showed greater survival and lower growth rates overall. Differences in growth and survival were related to differences in photosynthetic pigment and nutrient contents and in the photosynthetic efficiency of photosystem II. Plasticity was strongest for growth characters and pigment contents.     

Detection of the endophyte Discula umbrinella in buds and twigs of Fagus sylvatica

Endophytic fungi have been isolated from buds and twigs of beech trees collected at four different sites in Switzerland. Discula umbrinella. was recovered at high frequencies from the bud scales and the twig pieces contiguous to the buds, but was virtually absent from the rolled up leaves enclosed by the scales. In addition to infection by air-borne inoculum, thalli of D. umbrinella may grow from the twigs into the leaf tissues.     

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

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

Nutrient contents and concentrations in relation to growth of Picea abies and Fagus sylvatica along a European transect

Mineral nutrition of Norway spruce (Picea abies (L.) Karst.) and beech (Fagus sylvatica L.) was investigated along a transect extending from northern Sweden to central Italy. Nitrogen (N) concentrations of needles and leaves in stands growing on acid soils did not differ significantly between central Italy and southern Sweden (1.0 +/- 0.1 mmol N g(-1) for needles and 1.9 +/- 0.14 mmol N g(-1) for leaves). In both species, foliar N concentrations were highest in Germany (1.2 mmol N g(-1) for needles and 2.0 mmol N g(-1) for leaves) and decreased by 50% toward northern Sweden (0.5 mmol N g(-1)). Both species showed constant S/N and P/N ratios along the transect. Calcium, K and Mg concentrations generally reflected local soil conditions; however, Mg concentrations reached deficiency values in Germany. Leaf area per unit dry weight varied significantly along the transect with lowest values for Norway spruce recorded in northern Sweden and Italy (3.4 m(2) kg(-1)) and a maximum in central Europe (4.7 m(2) kg(-1)). A similar pattern was observed for beech. Despite the low variation in foliar N concentrations on the large geographic scale, local and regional variations in N concentrations equalled or exceeded the variation along the entire continental transect. Furthermore, nutrient contents (i.e., nutrient concentration x dry weight per needle or leaf) showed a greater variation than nutrient concentrations along the transect. Nitrogen contents of Norway spruce needles reached minimum values in northern Sweden (2.4 micro mol N needle(-1)) and maximum values in Denmark (5.0 micro mol N needle(-1)). The N content of beech leaves was highest in Denmark (242 micro mol N leaf(-1)). At the German site, foliar N content rather than N concentration reflected the seasonal dynamics of foliar growth and N storage of the two species. During foliage expansion, there was an initial rapid increase in N content and a decrease in N concentration. This pattern lasted for about 2 weeks after bud break and was followed by 6 weeks during which dry weight and N content of the foliage increased, resulting in a further decrease in N concentration. During summer, dry weight and N content of mature needles of Norway spruce increased further to reach a maximum in autumn, whereas N concentration remained constant. In spring, reallocation of N from 1- and 2-year-old needles was 1.5 and 1.0 micro mol N needle(-1), respectively. This remobilized N was a major source of N for the development of new needles, which had an N content of 1.5 micro mol N needle(-1) after bud break. The seasonal remobilization of N from old foliage decreased with increasing needle age. Needle N content and dry weight decreased progressively with age (1 micro mol N needle(-1) between age classes 2 and 5), whereas N concentrations remained constant. For Norway spruce, annual stemwood production was correlated with needle N content but not with foliar N concentration or with the total amount of N in the canopy. Interspecific and geographical differences in plant nutrition are discussed on the basis of competitive demands for C and N between growth of foliage and wood.     

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.