The induction of sun and shade leaves of the European beech (Fagus sylvatica L.): anatomical studies

Summary Primordia from buds of sun and shade twigs of European beech (Fagus sylvatica L.) were collected six times a year for anatomical investigations. Differentiation into sun-leaf and shade-leaf primordia was first observed in early August. Sun-leaf primordia had five, and shade-leaf primordia four layers of mesophyll meristem cells. With potted graft unions of beeches possible structural changes of leaf primordia were investigated. Trees adapted to shade develop sun-leaf primordia when put into full daylight, provided the transfer happened before July. Trees adapted to full daylight developed leaf primordia which remained structurally sun-leaf primordia when the plant was kept under shade conditions. Shadeleaf branches of young beech trees cut in February in order to expose the shade buds to full daylight developed either shade leaves or intermediate shade/sun leaves. These experiments show that the subtending leaf may provide the developing axillary bud with photoassimilates, but its character, whether sun or shade leaf, has no influence on the character of the developing leaf primordia.     

Climate-tree-growth relationships of European beech (Fagus sylvatica L.) in the French Permanent Plot Network (RENECOFOR)

The influence of climate on the radial growth of Fagus sylvatica was investigated using 15 chronologies developed from mature stands of the French Permanent Plot Network (RENECOFOR) growing under different climatic and soil conditions. The relationships between climate and ring widths were analyzed using extreme growth years, simple correlations and response functions analysis. Monthly climatic regressors were derived by a physiological water balance model that used daily climatic data and stand parameters to estimate soil water deficits. The three most frequent negative pointer years (1959, 1989, 1976) result from a particularly intense and durable drought, whereas positive years (1977, 1958) coincide with wet conditions. The total ring chronology variance attributable to climate averages 34.1% (15.8% –57%). Current early-summer soil water deficit enters in 10 models and the deficit in June explains alone a large part of the radial growth variability (mean value: 26.6%). Temperature or soil water deficit for the other months and weather conditions during the previous season were of little consistency across stands. The response pattern of earlywood is very similar and the percentage of variance explained is higher (16.2% –57.8%). Latewood widths present a different response pattern. High minimum temperature in August and/or September often favour wide latewood widths and monthly water deficits play a secondary role. The percentage of variance explained ranges from 8.8% to 67.4%. Soil water capacity strongly modulates ring characteristics and climate-growth relationships. Mean sensitivity, expressed population signal, signal-to-noise ratio and the strength of growth-climate correlations increase with decreasing soil water capacity.     

The response of European beech (Fagus sylvatica L.) seedlings from two Italian populations to drought and recovery

The response of gas exchange, leaf chlorophyll content, relative fluorescence to decreasing water potential and recovery was followed in European beech seedlings from two Italian populations, differing in their native precipitation amounts. A population from Sicily (southern Italy) was selected as representative of a xeric population while a population from central Italy, Abetone, represented a mesic one. Dry-matter partitioning, leaf area, hydraulic sufficiency and xylem embolism were evaluated in both well-watered control plants as well as in plants subjected to drought. With the onset of water stress, values of water potential, leaf relative water content, net photosynthesis, leaf conductance and leaf chlorophyll concentration decreased concurrently while relative fluorescence remained unchanged. The population from Sicily showed a delay in effects of the imposed drought. Within 5 days of rewatering, leaf conductance was not fully recovered while all of the other parameters recovered to control levels, in both populations. Total, shoot, stem and root dry weight tended to be higher in seedlings from Abetone, even though both populations had similar photosynthetic rates. The population from Sicily exhibited about 3% greater (even if not significant) allocation to roots than the population from Abetone. Seedlings from Abetone had higher, but not significant, leaf specific conductivity and per cent loss in hydraulic conductivity than seedlings from Sicily. Drought resulted in a reduction of hydraulic conductivity and hydraulic sufficiency in seedlings from both populations. Photosynthesis of water stressed plants from both populations appeared to be reduced primarily by carbon dioxide diffusion through stomata and perhaps secondarily by changes in chlorophyll concentration rather than by efficiency of photosystem II. The effect of hydraulic factors on gas exchange during drought and recovery was not clearly evident.     

Root respiration and its importance for the carbon balance of beech saplings (Fagus sylvatica L.) in a montane beech forest

Root respiration of 10-year-old beech saplings (Fagus sylvatica L.) grown in the understorey (UND) and in a natural gap (GAP) of a mature beech forest in the Solling mountains, FRG, was investigated from April until December, 1990. Respiration rates of fine, medium and coarse roots were measured in situ by a PC-controlled cuvette system. Fine root respiration rates were in the range of 0.5–9.8 nmol CO₂ gDW^–1 s^–1 at both sites, but respiration rates of UND saplings were higher, compared to those of GAP saplings. The dependence of respiratory activity on soil temperature proved to be highly significant (p<0.001) for both plots, following a quasi-Arrhenius type curve. Fine root respiration rates of UND saplings were highly significantly, negatively correlated with the water content of the attached organic material, whereas respiration rates of GAP saplings did not show such a correlation. Further, a significant correlation (p<0.01) between mycorrhizal biomass and respiration rate was detected at the UND site, but not at the GAP site. Medium and coarse root respiration rates were very similar and no significant differences between the two sites were detected. Maximum respiration rates of 3.1 nmol CO₂ gDW^–1 s^–1 were reached in the middle of July. Due to low light intensities in the under storey, daily net CO₂ assimilation rates of UND saplings were much smaller than those of GAP saplings. At both sites, net CO₂ assimilation rates varied more than respiration rates and thus the carbon balance of beech saplings was more affected by the rate of carbon fixation than by the rate of respiratory carbon loss.     

Factors toxic to beech (Fagus sylvatica L.) seedlings in acid soils

The effects of highly and moderately acid soils on total biomass, biomass partitioning, fine root characteristics and nutritional status of beech seedlings (Fagus sylvatica L.) were studied in a growth chamber experiment. In Haplic Arenosols seedlings grew slowly but equally well without damage symptoms in a highly acid and a moderately acid soil horizon. The moderately acid A_h+B_w-horizon of a Eutric Cambisol was favourable to seedling growth. The fine root development was reduced in the highly acid A+B_w-horizon of a Dystric Cambisol and in the A_h+E-horizon of a Haplic Podzol, the latter of which also caused increased mortality. Seedling growth in the B₂-horizon of the Haplic Podzol was vigorous, in spite of a higher level of extractable Al and lower base saturation as compared with the A_h+E-horizon. These results are interpreted in relation to soil acidity, soil Al and nutritional status of the seedlings. We conclude that neither Al-toxicity nor nutrient deficiency cause the damage symptoms observed in the A_h+E-horizon of a Haplic Podzol and the fine root reduction in the A+B_w-horizon of a Dystric Cambisol. The damage symptoms of the PZh_A treatment seems to be more the result of H-toxicity or H-related factors other than nutrient shortage or Al-toxicity. Other pH-related toxic factors are discussed.     

Root growth inhibition effects of holorganic moder humus layer under spruce (Picea abies KARST.) and beech (Fagus sylvatica L.)

Field observations demonstrated inhibition of root growth by moder humus material of the Of₂ and Oh-layers under beech and spruce. Growth chamber experiments with spruce seedlings showed that root growth on a natural Of₂-substrate is ten times lower than on resin-quartz mixtures. Added phenolic acid (protocatechic a.) inhibited root growth only in sterilized substrate. Leachates of Of₂-material inhibited root growth in a mineral substrate devoid of absorption complex (pure quartz sand), but not in a resin-quartz mixture. It is concluded that inhibiting substances are probably ionized molecules that can withstand biodegradation.     

Gene Flow in European Beech (Fagus sylvatica L.)

Three relatively isolated stands were used to study gene flow in European beech (Fagus sylvatica L.) in Northern Germany. Nine allozyme loci (Got-B, Idh-A, Lap-A, Mdh-B, Mdh-C, Mnr-A, 6-pgdh-A, Pgi-B and Pgm-A) were utilized for multilocus-genotyping adult trees and seeds. Expected heterozygosity (He) ranged from 0.325 to 0.351 for the three stands. F(ST) revealed that there was small differentiation among stands (mean F(ST) = 0.013). The indirect estimates of gene flow (Nm) based on the mean F(ST) were high and the average Nm was 19.14. External gene flow by pollen ranged from 0.7 to 1.2% inferred from new alleles in seed samples. Moreover, paternity analysis was used to assess effective pollen dispersal by inferring paternity of offspring. The weighted mean distances of pollen dispersal for these three stands were 36.8 and 37.1 m based on simple exclusion procedure and most-likely method, respectively. Two of the trees in one stand had rare allozyme alleles (Lap-A1 and Idh-A4, respectively), which were used to directly measure pollen movement away from those trees. The frequency of the rare Lap and Idh alleles in seeds declines as the distance from the source tree increases. The weighted mean distance of pollen dispersal with rare allele Lap-A1 or Idh-A4 was 26.3 m.     

A contribution to understanding the internal nitrogen budget of beech (Fagus sylvatica L.)

Summary The seasonal variation in the total nitrogen content of the xylem sap of the lower trunk section was investigated for two middle aged beech tree stands in northern Hessen each containing 130 trees. In addition seasonal changes in the percentage of nitrate in the total nitrogen content are described. The median values of the total nitrogen content of the xylem sap during the spring mobilization period reached 175 and 250 mg/l. During the summer about 35% of the total nitrogen in the xylem sap is in the form of nitrate. Finally, the distribution of NO₃ in the xylem sap along the trunk height was studied for two sample trees for each of the four seasons (n = 8).     

A study of the cambial zone and conductive phloem of common beech (Fagus sylvatica L.) using an image analysis method

Quantification is a major problem when using histology to study the influence of ecological factors on tree structure. This paper presents a method to prepare and to analyse transverse sections of cambial zone and of conductive phloem in bark samples. The following paper (II) presents the automated measurement procedure. Part I here describes and discusses the preparation method, and the influence of tree age on the observed structure. Highly contrasted images of samples extracted at breast height during dormancy were analysed with an automatic image analyser. Between three young (38 years) and three old (147 years) trees, age-related differences were identified by size and shape parameters, at both cell and tissue levels. In the cambial zone, older trees had larger and more rectangular fusiform initials. In the phloem, sieve tubes were also larger, but their shape did not change and the area for sap conduction was similar in both categories. Nevertheless, alterations were limited, and demanded statistical analysis to be identified and ascertained. The physiological implications of the structural changes are discussed.     

Crown alterations induced by decline: a study of relationships between growth rate and crown morphology in beech (Fagus sylvatica L.)

One of the first symptoms expressed by declining trees is reduced growth in stem diameter and length increment. The possibility of a relationship between length increment and crown thinning in beech (Fagus sylvatica L.) was investigated by developing a computer model to simulate first order branching patterns of the apical 2 m of monopodially branching beech trees, 70–100 years old, for a range of length increment rates. The model was based on values for branching angle, main axis and branch length increment, number of branches produced per year and branch mortality rates for six healthy and declining trees. Shoot growth rates in the apical 2 m of the sample trees ranged from about 5 cm/year (decline class 3) to 43 cm/ year (healthy). Simulations of branching patterns in the apical 2 m of trees growing at different rates indicated that, when growth rate exceeded about 20 cm/year, total first order branch length and area explored were independent of growth rate. When growth rates fell below this value there was a reduction in total area explored and first order branch length due primarily to the formation of fewer branches. More acute branching angles contributed to a reduction in the area explored. Growth rate-related crown thinning could increase the risk of bark necrosis and secondary pathogen infection during dry and/or hot spells.     

Carbon,nitrogen, phosphorus,and sulphur concentration and partitioning in beech ecotypes (Fagus sylvatica L.): phosphorus most affected by drought

Beech seedlings of different drought sensitivity originating from 11 German provenances were grown in pots and cultivated in a greenhouse. The present paper aims to give insights on uptake, transport and use of macronutrients, since the knowledge of drought effects on the nutrition of trees is low compared to water relations. Therefore, the elemental composition, the ratio of inorganic to total content, and the partitioning between roots and shoots of carbon, nitrogen, phosphorus, and sulphur were investigated as affected by provenance and drought treatment. Phosphorus and phosphate concentrations decreased in all tissues after a 3 week drought treatment simulating a summer drought period. In roots carbon increased and nitrate decreased, in stems nitrogen decreased but nitrate increased following drought. The observed effects on phosphorus and phosphate are discussed in terms of lower phosphate mobility in the substrate due to lower water availability. The decrease in the ratio of phosphate to phosphorus in the tissues suggests the use of vacuolar phosphate pools for maintaining organic phosphorus homeostasis. The partitioning of all macronutrients was not affected by drought, although phosphorus and phosphate were significantly lowered in tissues. In most of the parameters studied significant differences between provenances were found. The recently observed drought sensitivity of provenances was not reflected in the strength of concentration changes or partitioning of macronutrients by drought over provenances.     

The variability of vessel size in beech (Fagus sylvatica L.) and its ecophysiological interpretation

The vessel areas of ten beech trees growing on a dry site were measured separately for all tree rings using automatic image analysis. These data were correlated with the monthly amount of precipitation from the July prior to the growing season until the August of the current growing season. It is evident that vessel formation at the beginning of cambial activity is mainly controlled by internal factors. The rainfall in the previous summer and autumn and in the contemporary May had only a slight influence. Vessel formation towards the end of the cambial activity is strongly influenced by the July rainfall and is thus determined to a greater degree by external factors. These results are discussed on the basis of hypotheses of tree physiology.     

Detailed analysis of the geometric relationship between external traits and the shape of red heartwood in beech trees (Fagus sylvatica L.)

Frequently occurring red heartwood decisively restricts the volume of light-coloured beech wood, which can be processed to high-grade appearance products. Forestry and wood industry lack suitable means for maximising the yield of light timber, since the intra-tree extent of red heartwood varies considerably. The present study characterised in detail the intra-tree shape of red heartwood and its relationships to tree-external traits (dead branches, branch scars), considered as possible initiation points of red heartwood formation. An experimental method based on log scanning and image analysis was developed and applied. Using its output for three-dimensional visualisation and data analyses, external traits being linked to the local red heartwood shape were identified. Furthermore geometric relationships were established for characterising these external traits and for deriving discriminating criteria.

Physiological responses of Fagus sylvatica L. exposed to low levels of ozone in open-top chambers

Four-year-old beech seedlings were fumigated with three levels of ozone for 2 consecutive years in open-top chambers. During the second growth season different physiological measurements were conducted before and during daily fumigation. A 25–40% decrease in net photosynthesis was seen during fumigation, whereas no differences were detected before fumigation in July. In August lasting effects in net photosynthesis were seen. The apparent quantum yield was decreased after fumigation. Stomatal conductance was generally decreased during fumigation, but transpiration was reduced relatively less than net photosynthesis indicating a lower water use efficiency of the trees exposed to ozone. Chlorophyll fluorescence (F_v/F_m) showed additive reductions in relation to ozone and light.     

Optimized nonradioisotopic amplified fragment length polymorphism method for European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.)

The amplified fragment length polymorphism (AFLP) technique has increasingly been used for the study of forest tree species. A nonradioisotopic AFLP method was optimized for European beech (Fagus sylvatica L.) and found reproducible. However, type ofTaq DNA polymerase and choice of primers must be considered for a consistent AFLP pattern.     

Displacement of frequently occurring heacy metals in autumn leaves of beech (Fagus sylvatica)

Summary By utilizing energy-dispersive X-ray microanalysis the heavy metals manganese, iron, zinc and lead were determined in different tissues of beech (Fagus sylvatica L.) autumn leaves. It was observed that all four metals were accumulated in sieve element-companion cell complexes of the leaf veins in the period from the green (2 October) to the brown (23 October) leaf coloration. Concomitantly, the concentration of these metals decreased in certain tissues of the stems subtending the leaves. The major reservoirs of heavy metals in the stems were periderm, cortex, pith and xylem rays. Since sieve element-companion cell complexes of the stems showed increasing metal concentrations during the transition from the green to the brown leaf stage, and since this temporary increase also occurred in the petiole phloem, it is inferred that symplastic transport can be used in addition to apoplastic transport for the displacement of heavy metals. It is assumed that the accumulation of heavy metals in the veins of autumn leaves indicates the deposition of surplus ions which are excluded when leaves are cast off.Supported by Deutsche Forschungsgemeinschaft and by Bundesministerium für Forschung und Technologie, Federal Republic of Germany, Grant PBE 373607.     

Changes in photosynthetic activity of buds and stem tissues of Fagus sylvatica during winter

Summary The photosynthetic activity of leafless twigs and buds of Fagus sylvatica was determined by in vivo chlorophyll fluorescence from November to May. Measurements were made on the day of sampling, and during exposure to warm temperatures until reactivation was attained. Under the same conditions, bud development and growth were forced by exposure of cut twigs to 25/18° C at long-day conditions, and bud swelling and bud burst were monitored. Winter inactivation of photosynthesis results in a reduction of the photochemical efficiency of PS II, as indicated by lowering of F_V/F_M, from January through March. The greatest reduction is in cortical chlorenchyma, the least in folded leaflets and primordia of buds. Restoration of photosynthetic activity, brought about by warming, needed 3–4 weeks in cortex and 1–2 weeks in buds during the coldest period of winter. Frequency distributions based on three types of chlorophyll fluorescence transients, defined by quantitative fluorescence parameters, have proved to be a valuable method for a differentiated expression of the unequal functional activation states of parallel samples. The seasonal course of winter inactivation of photosynthesis did not correspond entirely with the depth of bud dormancy as revealed by the forcing treatments; inactivation of photosynthesis may be more closely synchronized with changes in frost hardiness; possible causes are discussed. We suggest, therefore, that a distinction should be made between inactivation of metabolic processes and depth of dormancy, even though these processes are inherently interrelated.Dedicated to Professor Otto Härtel on the occasion of his 80th birthday     

Fine-root seasonal pattern,production and turnover rate of European beech (Fagus sylvatica L.) stands in Italy Prealps: Possible implications of coppice conversion to high forest

Abstract

The aim of this study was to investigate the possible effects of coppice conversion to high forest on the beech fine-root systems. We compared the seasonal pattern of live and dead fine-root mass (d < 2 mm), production and turnover in three beech stands that differed in management practices. Tree density was higher in the 40-year-old coppice stand than in the stands that were converted from coppice to high forest in 1994 and 2004, respectively. We found that a reduction in tree density reduced the total fine-root biomass (Coppice stand, 353.8 g m^?2; Conversion 1994 stand, 203.6 g m^?2; Conversion 2004 stand, 176.2 g m^?2) which continued to be characterised by a bimodal pattern with two major peaks, one in spring and one in early fall. Conversion to high forest may also affect the fine-root soil depth distribution. Both fine-root production and turnover rate were sensitive to management practices. They were lower in the Coppice stand (production 131.5 g m^?2 year^?1; turnover rate 0.41 year^?1) than in the converted stands (1994 Conversion stand: production 232 g m^?2 year^?1, turnover rate 1.06 year^?1; 2004 Conversion stand: production 164.2 g m^?2 year^?1, turnover rate 0.79 year^?1).