Adult Beech Trees Research Articles

Gas Exchange and Antioxidative Compounds in Young Beech Trees under Free‐Air Ozone Exposure and Comparisons to Adult Trees

Three-year-old beech (Fagus sylvatica) seedlings growing in containers were placed into the sun and shade crown of a mature beech stand exposed to ambient (1 x O(3)) and double ambient (2 x O(3)) ozone concentrations at a free-air exposure system ("Kranzberg Forst", Germany). Pigments, alpha-tocopherol, glutathione, ascorbate, and gas exchange were measured in leaves during 2003 (a drought year) and 2004 (an average year). Sun-exposed seedlings showed higher contents of antioxidants, xanthophylls, and beta-carotene and lower contents of chlorophyll, alpha-carotene, and neoxanthin than shade-exposed seedlings. In 2003 sun-exposed seedlings showed higher contents of carotenoids and total glutathione and lower net photosynthesis rates (A(max)) compared to 2004. O(3) exposure generally affected the content of chlorophyll, the xanthophyll cycle, and the intercellular CO(2) concentration (c(i)). Seedlings differed from the adjacent adult trees in most biochemical and physiological parameters investigated: Sun exposed seedlings showed higher contents of alpha-tocopherol and xanthophylls and lower contents of ascorbate, chlorophyll, neoxanthin, and alpha-carotene compared to adult trees. Shade exposed seedlings had lower contents of xanthophylls, alpha-carotene, and alpha-tocopherol than shade leaves of old-growth trees. In 2003, seedlings had higher A(max), stomatal conductance (g(s)), and c(i) under 2 x O(3) than adult trees. The results showed that shade acclimated beech seedlings are more sensitive to O(3), possibly due to a lower antioxidative capacity per O(3) uptake. We conclude that beech seedlings are uncertain surrogates for adult beech trees.

Effects of Long‐Term Free‐Air Ozone Fumigation on δ15N and Total N in Fagus sylvatica and Associated Mycorrhizal Fungi

Patterns of nitrogen (N) isotope composition (delta(15)N) and total N contents were determined in leaves, fine roots, root-associated ectomycorrhizal fungi (ECM) of adult beech trees (FAGUS SYLVATICA), and soil material under ambient (1 x O(3)) and double ambient (2 x O(3)) atmospheric ozone concentrations over a period of two years. From fine root to leaf material delta(15)N decreased consecutively. Under enhanced ozone concentrations total N was reduced in fine roots and delta(15)N showed a decrease in roots and leaves. In the soil and in most types of mycorrhizae, delta(15)N and total N were not altered due to ozone fumigation. The number of vital ectomycorrhizal root tips increased and the mycorrhizal community structure changed in 2 x O(3). Simultaneously, the specific rate of inorganic N-uptake by the roots was reduced under the double ozone regime. From these results it is assumed that 2 x O(3) changes N-nutrition of the trees at the level of N-acquisition, as indicated by enhanced mycorrhizal root tip density, altered mycorrhizal species composition, and reduced specific N-uptake rates.

Chronic ozone exposure affects leaf senescence of adult beech trees: a chlorophyll fluorescence approach

Accelerated leaf senescence is one of the harmful effects of elevated tropospheric ozone concentrations ([O(3)]) on plants. The number of studies dealing with mature forest trees is scarce however. Therefore, five 66-year-old beech trees (Fagus sylvatica L.) have been exposed to twice-ambient (2xambient) [O(3)] levels by means of a free-air canopy O(3) exposure system. During the sixth year of exposure, the hypothesis of accelerated leaf senescence in 2xambient [O(3)] compared with ambient [O(3)] trees was tested for both sun and shade leaves. Chlorophyll (chl) fluorescence was used to assess the photosynthetic quantum yield, and chl fluorescence images were processed to compare functional leaf homogeneity and the proportion of O(3)-injured leaf area (stipples) under ambient and 2xambient [O(3)] regimes. Based on the analysis of chl fluorescence images, sun leaves of both ambient and 2xambient [O(3)] trees had apparently developed typical necrotic O(3) stipples during high O(3) episodes in summer, while accelerated senescence was only observed with sun leaves of 2xambient [O(3)] trees. This latter effect was indicated along with a faster decrease of photosynthetic quantum yield, but without evidence of changes in non-photochemical quenching. Overall, treatment effects were small and varied among trees. Therefore, compared with ambient [O(3)], the consequence of the observed O(3)-induced accelerated leaf senescence for the carbon budget is likely limited.

Carbon isotopic composition and oxygen isotopic enrichment in phloem and total leaf organic matter of European beech (Fagus sylvatica L.) along a climate gradient

This study investigated the influence of climate on the carbon isotopic composition (sigma13C) and oxygen isotopic enrichment (delta18O) above the source water of different organic matter pools in European beech. In July and September 2002, sigma13C and delta18O were determined in phloem carbohydrates and in bulk foliage of adult beech trees along a transect from central Germany to southern France, where beech reaches its southernmost distributional limit. The data were related to meteorological and physiological parameters. The climate along the transect stretches from temperate [subcontinental (SC)] to submediterranean (SM). Both sigma13Cleaf and delta18Oleaf were representative of site-specific long-term environmental conditions. sigma13C of leaves collected in September was indicative of stomatal conductance, vapour pressure deficit (VPD) and radiation availability of the current growing season. delta18O was mainly correlated to mean growing season relative humidity (RH) and VPD. In contrast to the leaves, sigma13Cphloem varied considerably between July and September and was well correlated with canopy stomatal conductance (Gs) in a 2 d integral prior to phloem sampling. The relationship between sigma13C and delta18O in both leaves and phloem sap points, however, to a combined influence of stomatal conductance and photosynthetic capacity on the variation of sigma13C along the transect. delta18Ophloem could be described by applying a model that included 18O fractionation associated with water exchange between the leaf and the atmosphere and with the production of organic matter. Hence, isotope signatures can be used as effective tools to assess the water balance of beech, and thus, help predict the effects of climatic change on one of the ecologically and economically most important tree species in Central Europe.

Extraordinary drought of 2003 overrules ozone impact on adult beech trees (Fagus sylvatica)

The extraordinary drought during the summer of 2003 in Central Europe allowed to examine responses of adult beech trees (Fagus sylvatica) to co-occurring stress by soil moisture deficit and elevated O3 levels under forest conditions in southern Germany. The study comprised tree exposure to the ambient O3 regime at the site and to a twice-ambient O3 regime as released into the canopy through a free-air O3 fumigation system. Annual courses of photosynthesis (Amax), stomatal conductance (gs), electron transport rate (ETR) and chlorophyll levels were compared between 2003 and 2004, the latter year representing the humid long-term climate at the site. ETR, Amax and gs were lowered during 2003 by drought rather than ozone, whereas chlorophyll levels did not differ between the years. Radial stem increment was reduced in 2003 by drought but fully recovered during the subsequent, humid year. Comparison of AOT40, an O3 exposure-based risk index of O3 stress, and cumulative ozone uptake (COU) yielded a linear relationship throughout humid growth conditions, but a changing slope during 2003. Our findings support the hypothesis that drought protects plants from O3 injury by stomatal closure, which restricts O3 influx into leaves and decouples COU from high external ozone levels. High AOT40 erroneously suggested high O3 risk under drought. Enhanced ozone levels did not aggravate drought effects in leaves and stem.

Soluble N compound profiles and concentrations in European beech (Fagus sylvatica L.) are influenced by local climate and thinning

We assessed seasonal changes of total soluble nonprotein nitrogen compounds (TSNN) in adult European beech trees (Fagus sylvatica, L.) growing under different local climate during the growing season immediately following a thinning treatment and 3 years later. In both years, samples of leaves, xylem sap and phloem exudates from beech trees growing in thinned and unthinned (control) stands on a dry, warm SW exposed and a cooler, moist NE exposed site were collected in May, July and September. In May of both years, asparagine (Asn) and glutamine (Gln) were most abundant in leaves and xylem, respectively, whereas arginine (Arg) dominated in the phloem. In July, TSNN concentrations decreased in all tissues and sites, but differences in water availability between aspects were reflected in TSNN concentrations. In September, differences in the increase of Arg concentration in the phloem were related to differences in the onset of senescence between treatments. Thinning treatment increased amino compound concentrations of beech tissues in July on both aspects, particularly at the NE thinned site. It is supposed that, the N balance of adult beech is favoured by both, the thinning treatments as well as the cool-moist climate prevailing at the NE aspect.

Beech Leaf Colonization by the Endophyte Apiognomonia errabunda Dramatically Depends on Light Exposure and Climatic Conditions

Ozone and light effects on endophytic colonization by Apiognomonia errabunda of adult beech trees (Fagus sylvatica) and their putative mediation by internal defence compounds were studied at the Kranzberg Forest free-air ozone fumigation site. A. errabunda colonization was quantified by "real-time PCR" (QPCR). A. errabunda-specific primers allowed detection without interference by DNA from European beech and several species of common genera of plant pathogenic fungi, such as Mycosphaerella, Alternaria, Botrytis, and Fusarium. Colonization levels of sun and shade leaves of European beech trees exposed either to ambient or twice ambient ozone regimes were determined. Colonization was significantly higher in shade compared to sun leaves. Ozone exhibited a marginally inhibitory effect on fungal colonization only in young leaves in 2002. The hot and dry summer of 2003 reduced fungal colonization dramatically, being more pronounced than ozone treatment or sun exposure. Levels of soluble and cell wall-bound phenolic compounds were approximately twice as high in sun than in shade leaves. Acylated flavonol 3- O-glycosides with putatively high UV-B shielding effect were very low in shade canopy leaves. Ozone had only a minor influence on secondary metabolites in sun leaves. It slightly increased kaempferol 3- O-glucoside levels exclusively in shade leaves. The frequently prominent hydroxycinnamic acid derivative, chlorogenic acid, was tested for its growth inhibiting activity against Apiognomonia and showed an IC50 of approximately 8 mM. Appearance of Apiognomonia-related necroses strongly correlated with the occurrence of the stress metabolite, 3,3',4,4'-tetramethoxybiphenyl. Infection success of Apiognomonia was highly dependent on light exposure, presumably affected by the endogenous levels of constitutive phenolic compounds. Ozone exerted only minor modulating effects, whereas climatic factors, such as pronounced heat periods and drought, were dramatically overriding.

Role of ethylene in the regulation of cell death and leaf loss in ozone‐exposed European beech

ABSTRACTTo test the involvement of ethylene in mediating ozone‐induced cell death and leaf loss in European beech (Fagus sylvatica L.), tree seedlings were exposed to proportionally increased or decreased field ozone levels for up to 6 months. Ozone treatment caused cell death and accelerated leaf loss at higher than ambient levels, but had only minor effects at ambient and no effects at subambient ozone levels. The emission of ethylene, the levels of its precursor, 1‐aminocyclopropane‐1‐carboxylate (ACC), and mRNA levels of specific ACC synthase (FS‐ACS2) and ACC oxidase (FS‐ACO1) isoforms showed a persistent increase and preceded cell death by approximately 2 weeks. Inhibition of ethylene biosynthesis led to reduced lesion formation whereas application of ACC accelerated ozone‐induced cell death and leaf loss. Similar results were obtained when adult beech trees were exposed to 2 × ozone by a whole tree free‐air canopy exposure system. The results suggest a role of ethylene in amplifying ozone effects under field conditions in this major European broad‐leaved tree species.

Response patterns in adult forest trees to chronic ozone stress: identification of variations and consistencies

The responsiveness of adult beech and spruce trees to chronic O 3 stress was studied at a free-air O 3 exposure experiment in Freising/Germany. Over three growing seasons, gas exchange characteristics, biochemical parameters, macroscopic O 3 injury and the phenology of leaf organs were investigated, along with assessments of branch and stem growth as indications of tree performance. To assess response pattern to chronic O 3 stress in adult forest trees, we introduce a new evaluation approach, which provides a comprehensive, readily accomplishable overview across several tree-internal scaling levels, different canopy regions and growing seasons. This new approach, based on a three-grade colour coding, combines statistical analysis and the proficient ability of the “human eye” in pattern recognition.

Water shortage affects the water and nitrogen balance in Central European beech forests.

Whilst forest policy promotes cultivation and regeneration of beech dominated forest ecosystems, beech itself is a highly drought sensitive tree species likely to suffer from the climatic conditions prognosticated for the current century. Taking advantage of model ecosystems with cool-moist and warm-dry local climate, the latter assumed to be representative for future climatic conditions, the effects of climate and silvicultural treatment (different thinning regimes) on water status, nitrogen balance and growth parameters of adult beech trees and beech regeneration in the understorey were assessed. In addition, validation experiments with beech seedlings were carried out under controlled conditions, mainly in order to assess the effect of drought on the competitive abilities of beech. As measures of water availability xylem flow, shoot water potential, stomatal conductance as well as delta (13)C and delta (18)O in different tissues (leaves, phloem, wood) were analysed. For the assessment of nitrogen balance we determined the uptake of inorganic nitrogen by the roots as well as total N content and soluble N compounds in different tissues of adult and young trees. Retrospective and current analysis of delta (13)C, growth and meteorological parameters revealed that beech growing under warm-dry climatic conditions were impaired in growth and water balance during periods with low rain-fall. Thinning affected water, N balance and growth mostly of young beech, but in a different way under different local climatic conditions. Under cool, moist conditions, representative for the current climatic and edaphic conditions in beech forests of Central Europe, thinning improves nutrient and water status consistent to published literature and long-term experience of forest practitioners. However, beech regeneration was impaired as a result of thinning at higher temperatures and under reduced water availability, as expected in future climate.

Carbon and oxygen isotope composition of organic compounds in the phloem sap provides a short‐term measure for stomatal conductance of European beech (Fagus sylvatica L.)

ABSTRACTAt eight different dates during the 2000 growing season, δ13C and δ18O were determined in the phloem of adult beech trees growing in natural beech stands in south‐west Germany differing in stand density and local climate. In addition, stand transpiration, precipitation, photosynthetic active radiation, relative air humidity, water pressure deficit of the air, air and soil temperature, soil water potential, and sugar concentration of the phloem sap were determined directly and evaporation and canopy stomatal conductance were modelled. All parameters were related to δ13C. The study aimed to identify the time integral within which the δ13C of organic compounds transported in the phloem is an indicative measure of these environmental influences. δ13C of soluble carbon transported in the phloem was well correlated with mean stomatal conductance in a two‐day integral prior to phloem sampling but did not depend on either light intensity or soil water availability. A strong positive relationship between δ13C and δ18O pointed to observed variation in δ13C of phloem sap being a result of variation in stomatal conductance. Bulk leaf δ13C was a poor indicator of changes in environmental conditions during the growing season. From these results we conclude that the analysis of δ13C in soluble carbon transported in the phloem is a reliable indicator of short‐term changes in Ci/Ca. In contrast, the δ13C of structural carbon in beech foliage represents an integration of a range of factors that mask short‐term influences responsible for Ci/Ca.

Spatial and seasonal variations in stem respiration of beech trees (Fagus sylvatica)

Stem respiration of adult beech (Fagus sylvatica L.) trees was measured in the field in eastern France at several levels in the crown and along the stem. Strong variations in respiration rates throughout the season and within the trees were mainly caused by gradients in stem tem- perature, growth rates and distribution of living cells. The higher respiration rates, were measured in the upper crown. During the non-growing season, maintenance respiration ranged between 7.2 and 528 µmol m -3 s -1 at breast height and in the upper crown, respectively. Q10 increased along the stem from 1.3 at breast height to 2.0 in the upper crown. There was a linear relationship between (N) and the percentage of living cells in the wood, but respiration increased strongly with (N). Growth respiration accounted for 45-76% of annual stem respiration, and the growth respiration coefficient was close to 0.2 g C respired g -1 C fixed.

Diurnal courses of ammonium net uptake by the roots of adult beech (Fagus sylvatica) and spruce (Picea abies) trees

Differences in ammonium net uptake by the roots of beech (Fagus sylvatica L.) and spruce (Picea abies (L.) Karst) trees between day and night were examined during the growing seasons in 1995 and 1996 using the depletion technique. In addition, diurnal courses of ammonium net uptake of both species were analysed in five sets of uptake experiments in May and September 1997 and were related (1) to the content of carbohydrates, organic acids and total soluble non protein N (TSNN) in the fine roots, and (2) to xylem flow densities and soil temperature. During the growing seasons 1995 and 1996, ammonium net uptake of beech was significantly lower during the night than during the day at 5 of 8 dates of measurement. On average, uptake rates during the night amounted to 50% of the uptake rates during the day. In spruce, the mean values of ammonium net uptake rates determined were similar between day and night during both growing seasons. In beech, the assessment of diurnal courses showed highest ammonium uptake rates during noon and in the afternoon and minima at midnight. In May 1997, comparable, but less pronounced diurnal patterns of ammonium uptake were observed in spruce, whereas in September 1997, ammonium uptake by spruce was constant during the day. Since no distinct differences in carbohydrate and organic acid contents in fine roots were observed during the diurnal courses and since the addition of sucrose into the artificial soil solutions root tips were exposed to did not alter ammonium uptake, depression of uptake by C- and/or energy limitation during night could be excluded. The TSNN contents in the fine roots of beech (May and September 1997) and spruce (May 1997) showed a diurnal pattern inverse to ammonium uptake. It is concluded that the enrichment of TSNN compounds during night that is apparently caused by a reduction of xylem transport is responsible for the down-regulation of ammonium net-uptake.

Acclimation of leaf characteristics of Fagus species to previous-year and current-year solar irradiances.

To examine the effects of different solar irradiances on leaf characteristics at the leaf primordium and expansion stages, we shaded parts of branches in the upper canopies of two adult beech trees, Fagus crenata Blume and Fagus japonica Maxim., for 4 years. The treatments during the leaf primordium and leaf expansion stages, respectively, were: (1) high light and high light (H, control), (2) high light and low light (HL), (3) low light and low light (LL), and (4) low light and high light (LH). Both number of cell layers in palisade tissue and individual leaf area were affected by the previous-year irradiance, whereas cell length of palisade tissue was larger in LH leaves than in LL leaves, suggesting determination by current-year irradiance. Lamina chlorophyll/nitrogen ratio was higher in HL and LL leaves than in LH leaves, suggesting determination by current-year irradiance. Diurnal minimum values of leaf water potential measured under sunlit conditions were lower in H and LH leaves than in HL and LL leaves. Effective osmotic adjustment was found in H and LH leaves, suggesting that leaf water relations were affected by current-year irradiance. Net photosynthetic rate and stomatal conductance measured under sunlight conditions were higher in H and LH leaves than in HL and LL leaves. Thus, effects of current-year irradiance had a greater effect on leaf-area-based daily carbon gain than previous-year irradiance.

Interaction of phloem-translocated amino compounds with nitrate net uptake by the roots of beech (Fagus sylvatica) seedlings

From these results it is concluded that phloem transport of amino compounds, especially of Gln and Asp, In the present study two experimental approaches from the shoot to the roots mediates regulation of were used to investigate the influence of changes in nitrate net uptake by the roots of beech trees in order the allocation of amino compounds in the phloem of to adapt this process to the nitrogen demand of the beech (Fagus sylvatica L.) seedlings on nitrate net whole plant. uptake by the roots. In a first set of experiments Gln or Asp were directly fed into the phloem of the epicotyl Key words: Amino compounds, phloem transport, nitrate via bark flaps. These compounds were previously net uptake, NH 3 fumigation, regulation, Gln, Asp, Arg. found to be allocated in the phloem of adult beech trees and were shown to inhibit nitrate net uptake

Relationships between transpiration, stomatla damage and leaf wettability in declining beech trees

In order to determine the relationship between transpiration, stomatal damage and leaf wettability in adult beech trees ( Fagus sylvatica L.) in situ, sample leaves were gathered from healthy trees and from declining trees, both at the beginning and at the end of the growing season. Four parameters were assessed in each leaf: transpiration in the field, transpiration in the laboratory after rehydration, stomatal damage index, and leaf-water drop contact angle. The results showed that the transpiration was linked to the health status of the tree only when the external conditions were least favourable, i.e. in the month of June in the field. During the season the index of stomatal damage increased considerably in declining trees, while the drop contact angle decreased. Linear regression between the transpiration and the stomatal damage indix did not show any significant correlation, thus suggesting - at least in the environmental conditions prevailing in thi experiment - structural alterations in the stomata were not capable of modifying the transpiration capacity. Lastly, contact angle data did not correlate significantly with either the transpiration or the stomatal damage index.

Interpretation of seasonal changes of xylem embolism and plant hydraulic resistance in Fagus sylvatica

ABSTRACTThe annual course of xylem embolism in twigs of adult beech trees was monitored, and compared to concurrent changes of tree water status and hydraulic resistances. Xylem embolism was quantified in 1‐year‐old apical twigs by the hydraulic conductivity as a percentage of the maximum measured after removal of air emboli. Tree and root hydraulic resistances were estimated from water potential differences and sap flux measurements. The considerable degree of twig embolism found in winter (up to 90% loss of hydraulic conductivity) may be attributed to the effect of freeze‐thaw cycles in the xylem. A partial recovery from winter embolism occurred in spring, probably because of the production of new functional xylem. Xylem embolism fluctuated around 50% throughout the summer, without significant changes. Almost complete refilling of apical twigs was observed early in autumn. A significant negative correlation was found between xylem embolism and precipitation; thus, an active role of rainfall in embolism reversion is hypothesized. Tree and root hydraulic resistances were found to change throughout the growing period. A marked decrease of hydraulic resistance preceded the refilling of apical twigs in the autumn. Most of the decrease in total tree resistance was estimated to be located in the root compartment.

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.

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.

Part 4. Composition of the Beech-Podocarp Forest

Abstract Unmodified beech-podocarp forest was sampled by the quarter method near sea level and at an elevation of 1,000–1,300 ft. These samples indicated that: (a) there were about 250 trees per acre near sea level and 400 trees per acre at 1,000–1,300 ft. (b) about half the total tree population was kamahi (Weinmannia racemosa). (c) adult podocarps approximately equalled adult beeches in numbers at both altitudes. (d) totara (Podocarpus ballii) was the least important of the adult podocarps near sea level, but replaced rimu (Dacrydium cupressinum) substantially and miro (Podocarpus ferrugineus) completely at 1,000–1,300 ft. (e) at both altitudes two-thirds or more of the adult beech trees were mountain beech (Nothofagus solandri var. cliffortioides) and the remainder were silver beech (N. menziesii). (f) rata (Metrosideros umbellata) was rare near sea level and less important than either beeches or podocarps at 1,000–1,300 ft. (g) totara was smaller in diameter at 1,000–1,300 ft than near sea level but s...