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Effects of elevated pCO2 and/or pO3 on C-, N-, and S-metabolites in the leaves of juvenile beech and spruce differ between trees grown in monoculture and mixed culture.

Three and four-year-old saplings of beech (Fagus sylvatica L.) and spruce (Picea abies (L.) Karst.) grown in monoculture and mixed culture were exposed in phytotrons to (1) ambient air, (2) elevated pO(3), (3) elevated pCO(2), or (4) elevated pCO(2) plus elevated pO(3). After 5 months, the contents of soluble sugars, starch, soluble amino compounds, non-structural proteins (NSP), as well as reduced (GSH) and oxidized (GSSG) glutathione were determined in the leaves of both species in order to assess the effects of the gaseous regimes on primary metabolism. Elevated pO(3) did not affect sugar and starch levels in beech leaves in monoculture, but significantly increased sugar levels in beech leaves grown in mixed culture. In spruce needles, sugar levels tended to be enhanced in both culture types. Individual and combined exposure of elevated pCO(2) led to an increase in non-structural carbohydrate (soluble sugars plus starch) levels in beech and spruce leaves of both culture types. Differences in the responses of non-structural carbohydrate levels to elevated pCO(2) between beech and spruce were apparent from different contributions of sugars and starch to the increase in carbohydrate levels. Exposure to elevated pCO(2) and/or elevated pO(3) did not affect the levels of soluble amino compounds and NSP in beech leaves, but reduced amino compound levels in spruce needles of both culture types. Elevated pO(3) increased GSH levels in the leaves of both tree species in both culture types, while GSSG levels in monoculture were reduced in beech leaves, but significantly enhanced in spruce needles. Elevated pCO(2) reduced GSSG levels in beech and spruce leaves in monoculture, and GSH levels in spruce needles of both culture types. The combination of elevated pCO(2) and pO(3) increased GSSG levels in beech leaves of both culture types and in spruce needles in monoculture, but reduced GSH levels in spruce needles of both culture types. Apparently, under each gaseous regime, the culture type significantly altered primary metabolism of the leaves of beech and spruce.

Carbohydrate Metabolism↗

A comparison of lead isotope ratios in the bark pockets and annual rings of two beech trees collected in Derbyshire and South Yorkshire, UK.

ICP-MS analysis of the bark pockets and annual rings of two beech (Fagus sylvatica L.) trees collected from Longshaw, Derbyshire and Swinton, South Yorkshire in the UK recorded differences in the (206)Pb/(207)Pb isotope ratio. In the Longshaw sample, the (206)Pb/(207)Pb isotope ratio of the bark pockets ( approximately 1914-1998, 78-260 microg g(-1) Pb) declined from approximately 1.16 to 1.12, whilst the annual rings (1899-1998, 0.2-2.5 microg g(-1) Pb) had a (206)Pb/(207)Pb ratio of approximately 1.18. In the Swinton sample, the bark pockets (approximately 1919-1998, 7-78 microg g(-1) Pb) declined from 1.15 to 1.11 and the annual rings (1899-1998, 0.2-0.5 microg g(-1) Pb) from 1.18 to 1.15. The data implied that the bark pockets accumulated lead directly from the atmosphere through wet and dry deposition, whilst the annual rings accumulated lead from the soil via the roots. The bark pockets recorded a relative decline in the accumulation of lead from indigenous sources, such as lead smelting and coal combustion (1.17-1.19), and increase in imported sources such as the smelting of Australian ores (1.04) and leaded petrol usage (1.06-1.09). In contrast, the annual rings at Longshaw recorded ratios typical of indigenous lead, whilst the annual rings in Swinton recorded a relatively small decrease in (206)Pb/(207)Pb reflecting leaded petrol usage. The decline in (206)Pb/(207)Pb of the bark pockets was consistent with the historical decline in (206)Pb/(207)Pb of atmospheric lead recorded in peat, lake sediments and archival herbage at other UK locations.

Environmental Monitoring↗

Effects of stream acidification on fungal biomass in decaying beech leaves and leaf palatability.

We examined the effect of surface water acidification on rates of decomposition, ergosterol concentrations (as a measure of fungal biomass), and palatability to shredders of common beech leaves (Fagus sylvatica L.) in five mountain streams (pH 4.7-7.1). Leaf decomposition was significantly faster in the circumneutral streams (pH 6.4-7.1; k > or = 0.00175 d(-1)), when compared to acidic streams (pH 4.7-4.9; k < or = 0.00100 d(-1)). Fungal biomass showed no particular trend along the acidification gradient except that it peaked earlier in the stream closest to neutrality. Leaf palatability, measured as the feeding activity of the leaf-shredding amphipod Gammarus fossarum Koch, varied with the exposure time in the streams. Except for the higher palatability of leaves exposed during 6 weeks at the highest pH, patterns among streams were mostly similar. These results suggest that reduced processing rates in the most acidic streams were not related to differences in fungal biomass associated with decomposing leaves and that microbial conditioning was only slightly delayed by acidification. Possible effects of low pH and related variables (Ca, Al) on microbial decomposition and detritivorous macroinvertebrates are discussed to clarify the inhibition of beech leaf decomposition in the studied systems.

Acid Rain↗

Pb and 206Pb/207Pb isotopic analysis of a tree bark pocket near Sheffield, UK recording historical change in airborne pollution during the 20th century.

A section of tree trunk (beech, Fagus sylvatica) containing a bark pocket progressively enclosed at the junction of two branches was collected from a semi-rural location near Sheffield, UK. According to the annual growth rings, the bark pocket formed between ca. 1919 and 1998 (the date of felling). The bark pocket was divided into consecutive samples of differing radial depth (and thus age), that were analysed by ICP mass spectrometry. The Pb concentration varied from 7 to 78 mg kg (-1) and the 206Pb/207Pb isotope ratio from 1.11 to 1.15. In contrast, the current surface bark contained 46 mg kg(-1) Pb and recorded a 206Pb/207Pb ratio of 1.11. The changing elemental and isotopic composition of the bark pocket recorded historical change in the level and sources of airborne Pb pollution. An overall increase in Pb concentration with time was accompanied by a progressive reduction in 206Pb/207Pb from ca. 1935 to 1943. Mass balance calculations indicated that Pb additives in petrol contributed significantly to the rise in concentration, accounting for a maximum of 50% of the total Pb for ca. 1986-1998, but that other sources were generally dominant. The highest Pb concentrations were recorded from ca. 1951 to 1973, suggesting a high level of industrial pollution. A reduction in Pb concentration and reversal of the trend in 206Pb/207Pb was observed in the current bark.

Air Pollutants↗

Phytophthora pseudosyringae sp. nov., a new species causing root and collar rot of deciduous tree species in Europe.

In several studies of oak decline in Europe, a semi-papillate homothallic Phytophthora taxon was consistently isolated, together with other Phytophthora species, from rhizosphere soil samples. It was also found associated with necrotic fine roots and stem necroses of Fagus sylvatica and Alnus glutinosa. Due to morphological and physiological similarities, the semi-papillate isolates were previously identified as P. syringae by various authors. The morphology, physiology and pathogenicity against fine roots of Quercus robur, Q. petraea and F. sylvatica, bark of A. glutinosa, leaves of Ilex aquifolium and apple fruits of this Phytophthora species are described and compared with those of related and similar Phytophthora species, namely P. ilicis, P. psychrophila, P. quercina, P. citricola and P. syringae. The phylogenetic placement on the basis of ITS and mtDNA sequence data was also examined. Isolates of this taxon produce colonies with stellate to rosaceous growth patterns and limited aerial mycelium on various agar media. Antheridia are predominantly paragynous. In water culture catenulate hyphal swellings and semi-papillate caducous sporangia, that are usually limoniform, ellipsoid or ovoid, are formed abundandly, mostly in lax or dense sympodia. This taxon is a moderately slow growing, low temperature species with optimum and maximum temperatures around 20 and 25 degrees C, respectively. Tested isolates are moderately aggressive to fine roots of oaks and beech, highly aggressive to holly leaves and apple fruits, and slightly pathogenic to alder bark. Thirteen tested isolates had an identical and distinct ITS sequence which was more similar to that of P. ilicis and P. psychrophila than any other known taxa. On the basis of their unique combination of morphological characters, colony growth patterns, cardinal temperatures for growth, growth rates, pathogenicity to oaks, beech, alder, apple and holly, their host range, and ITS and mtDNA sequences the semi-papillate caducous Phytophthora isolates from oaks, beech and alder are clearly separated from related and similar Phytophthora spp., and described as a new species, P. pseudosyringae sp. nov.

Alnus↗

Historical records of airborne polycyclic aromatic hydrocarbons by analyzing dated corks of the bark pocket in a Longpetiole Beech tree.

Historical monitoring of airborne polycyclic aromatic hydrocarbons (PAHs) pollution levels was novelly demonstrated by analyzing the dated corks of a bark pocket formed from 1873 to 2003 in a Longpetiole Beech (Fagus longipetiolata) tree trunk sampled from southeastern China. The fundamental studies indicated that the PAHs of log K(oa) < 8.5 are primarily accumulated through interactions with lipid substances in cork and log K(oa) dependent, while the PAHs of log K(oa) > 8.5 existing as particle-phase dependent on log Vp are accumulated through stochastic entrapment by the lenticels on the surface of the cork. The translocation of PAHs by xylem flow and phloem stream as well as radial diffusion from the cork to the inner tissues was not significant, and the cork is most effective for accumulating airborne PAHs. The total concentrations of 16 EPA PAHs (sumPAHs) in the dated corks progressively increased from 43.5 ng/g recorded in the earliest available cork in 1873-1875 to the maximum 345.7 ng/g in 1956-1961, and then gradually decreased to 267.0 ng/g in 2003, while the concentration of perylene (PER) was slightly fluctuating at 0.178 +/- 0.033 ng/g. Moreover, the concentration ratio of sumPAHs to PER increased from 193 to 2431 from 1873 to 2003, indicating a progressive increase in PAH pollution in southeastern China.

Air Pollutants↗

Influence of odor from wood-decaying fungi on host selection behavior of deathwatch beetle, Xestobium rufovillosum.

Adult females of Xestobium rufovillosum de Geer demonstrated anemotactic orientation when exposed to an odor plume containing volatiles generated by wood-decaying fungi (Coriolus versicolor, Donkioporia expansa) and decayed oak wood (Quercus petraea, Quercus robur). They did not orient towards undecayed oak wood, beech (Fagus sylvatica), or pine wood (Pinus sylvestris). Although all insects tested showed anemotactic orientation, responses were nonlinear with respect to insect age. Adult females more readily oriented upwind when they were between 10 and 16 days old. Oviposition choice bioassays showed that ovipositing females would preferentially oviposit on extract-treated cellulose paper discs that had been treated with various strains of the wood-decaying fungus, Donkioporia expansa. HPLC-fractionated mycelial extracts were attractive to ovipositing deathwatch beetles, whereas HPLC-fractionated fungal broth extracts were repellent to ovipositing females. The implications of these results are discussed in the context of timber pest management in historically important buildings.

Animals↗

Crown condition surveys in Italian forests: issues in reporting findings.

This study examines defoliation and discoloration findings collected throughout the Italian Level I network between 1997 and 2000. Prior to this period no Quality Assurance procedures had been implemented, so that earlier findings cannot be considered reliable. The aim of the study is to compare different indices used in reporting findings, in order to determine the full potential and limitations of each. In international surveys, findings are normally expressed in terms of individual trees presenting a defoliation level greater than 25%; this criterion, however, has been challenged since it is not based on scientific evidence. By analysing the distribution of defoliationvalues (grouped in 5% classes) relating to the main species (Fagus sylvatica, Quercus pubescens, Quercus cerris, Picea abies), the study examines the behaviour of statistical indices such as the median, the mode and the mean. Instead of using the traditional 25% threshold to determine the part of the population with the highest defoliation values, a procedure is suggested whereby a different threshold can be determined for each species. It is established based on the 90 degrees percentile of cumulative defoliation values recorded over the four-year period. Each index used to report findings possesses its own information potential, and the findings may at times appear contradictory. The use of a combined defoliation and discoloration index (Ilce = Lacking Crown Equivalent Index) is also proposed, but the results obtained do not differ in substance from the results relating to defoliation alone. The functional limitations of this index are due mainly to the way the data are collected: for this reason suggestions are made on how to improve the procedure in future surveys.

Air Pollutants↗

On methods of spatial analysis for genotyped individuals.

Spatial autocorrelation methods have commonly been applied to individual-based spatial genetic studies, although their properties and the relations among the statistics have not been carefully examined. This paper first introduces a reformulation of widely used spatial statistics using point processes. When Moran's I statistics are applied to allele frequencies within an individual, the frequencies are no longer continuous variables but have only three discrete values and specific interpretations of Moran's I statistics and the number of alleles in common (NAC) can be expressed as the weighted sum of join-count statistics. The distributions of minor genotypes are amplified in Moran's I depending on the allele frequency in the population, while NAC uses a constant weighting system. Under the point process framework, spatial analysis can be conducted on the common theoretical base, from individual locations to genetic distributions of different levels, (for example, genotype and allele). The methodology is demonstrated by application to field data for molecular ecological studies of Fagus crenata population dynamics.

Biological Evolution↗

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.

Ascomycota↗

Physiological responses of beech and sessile oak in a natural mixed stand during a dry summer.

Responses of CO2 assimilation and stomatal conductance to decreasing leaf water potential, and to environmental factors, were analysed in a mixed natural stand of sessile oak (Quercus petraea ssp. medwediewii) and beech (Fagus svlvatica L.) in Greece during the exceptionally dry summer of 1998. Seasonal courses of leaf water potential were similar for both species, whereas mean net photosynthesis and stomatal conductance were always higher in sessile oak than in beech. The relationship between net photosynthesis and stomatal conductance was strong for both species. Sessile oak had high rates of photosynthesis even under very low leaf water potentials and high air temperatures, whereas the photosynthetic rate of beech decreased at low water potentials. Diurnal patterns were similar in both species but sessile oak had higher rates of CO2 assimilation than beech. Our results indicate that sessile oak is more tolerant of drought than beech, due, in part, to its maintenance of photosynthesis at low water potential.

Carbon Dioxide↗

Does canopy position affect wood specific gravity in temperate forest trees?

The radial increases in wood specific gravity known in many tree species have been interpreted as providing mechanical support in response to the stresses associated with wind loading. This interpretation leads to the hypothesis that individuals reaching the canopy should (1) be more likely to have radial increases in specific gravity and (2) exhibit greater increases than individuals in the subcanopy. Wood specific gravity was determined for three species of forest trees (Acer rubrum, Fagus grandifolia and Tsuga canadensis) growing in central Massachusetts, USA. Acer rubrum shows radial increases in specific gravity, but these increases are not more pronounced in canopy trees; the other two species show a pattern of radial decreases. The degree of radial increase or decrease is influenced by tree height and diameter. Of the dominant tree species for which we have data, A. rubrum, Betula papyrifera and Pinus strobus show radial increases in specific gravity, whereas F. grandifolia, T. canadensis and Quercus rubra show decreases. The occurrence of radial increases in B. papyrifera and P. strobus, which are often canopy emergents, suggests that it is overall adaptive strategy that is important rather than position (canopy vs. subcanopy) of any individual tree. It is suggested that radial increases in specific gravity are associated with early-successional status or characteristics and decreases with late-successional status or persistence in mature forest.

Acer↗

A new method to determine the oxygen concentration inside the sapwood of trees.

Research into the short-term fluctuations of oxygen concentrations in tree stems has been hampered by the difficulty of measuring oxygen inside tissues. A new method, which is based on fluorescence quenching of a ruthenium complex in the presence of oxygen, has been applied to measure changes of oxygen concentration in the sapwood of trees. During a field day-course oxygen increased with the radiation load and fell during the night (in Fagus orientalis from 20.3% in the afternoon to 17.5% in the morning next day). In a greenhouse experiment the sapwood oxygen concentration of Laurus nobilis could be influenced by flooding the root system. The very fast response, high resolution (better than 0.1%), easy calibration, and dependence only on oxygen and temperature make the technique well suited for measurements of oxygen concentrations in the sapwood.

Algorithms↗

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.

Acclimatization↗

Seasonal dynamics of soil carbon dioxide efflux and simulated rhizosphere respiration in a beech forest.

Respiration of the rhizosphere in a beech (Fagus sylvatica L.) forest was calculated by subtracting microbial respiration associated with organic matter decomposition from daily mean soil CO2 efflux. We used a semi-mechanistic soil organic matter model to simulate microbial respiration, which was validated against "no roots" data from trenched subplots. Rhizosphere respiration exhibited pronounced seasonal variation from 0.2 g C m(-2) day(-1) in January to 2.3 g C m(-2) day(-1) in July. Rhizosphere respiration accounted for 30 to 60% of total soil CO2 efflux, with an annual mean of 52%. The high Q10 (3.9) for in situ rhizosphere respiration was ascribed to the confounding effects of temperature and changes in root biomass and root and shoot activities. When data were normalized to the same soil temperature based on a physiologically relevant Q10 value of 2.2, the lowest values of temperature-normalized rhizosphere respiration were observed from January to March, whereas the highest value was observed in early July when fine root growth is thought to be maximal.

Carbon Dioxide↗

Fate of nitrogen released from 15N-labeled litter in European beech forests.

The decomposition and fate of 15N-labeled beech litter was monitored in three European beech (Fagus sylvatica L.) forests (Aubure, France; Ebrach, Germany; and Collelongo, Italy) for 3 years. Circular plots around single beech trees were isolated from roots of neighboring trees by soil trenching, and annual litterfall was replaced by 15N-labeled litter. Nitrogen was continuously released from the decomposing litter. However, over a 2-year period, this release was balanced by the incorporation of exogenous N. Released N accumulated mainly at the soil surface and in the topsoil. Microbial biomass remained almost constant during the experiment at all sites except for considerably lower values at Ebrach. The 15N enrichment of the microbial biomass increased strongly during the first year and then remained stable. The 15N released from the decomposing litter was rapidly detected in roots and leaves of the beech trees, increasing regularly and linearly over the course of the experiment. The uptake of litter-released 15N by the trees was reduced under conditions that reduced tree growth. Under these conditions, leaves and fine roots were the dominant N sinks, and little N was allocated to other plant parts. By contrast, N uptake and N allocation from leaves to stem and bark tissues increased when tree growth was enhanced. Budgets for 15N showed that 2 to 4% of litter-released N was incorporated into the trees, about 35% remained in the litter and about 50% reached the topsoil.

Biomass↗

Influence of tree internal N status on uptake and translocation of C and N in beech: a dual 13C and 15N labeling approach.

Influence of plant internal nitrogen (N) stocks on carbon (C) and N uptake and allocation in 3-year-old beech (Fagus sylvatica L.) was studied in two 15N- and 13C-labeling experiments. In the first experiment, trees were grown in sand and received either no N nutrition (-N treatment) or 4 mM unlabeled N (+N treatment) for 1 year. The -N- and +N-pretreated trees were then supplied with 4 mM 15N and grown in a 13CO2 atmosphere for 24 weeks. In the second experiment, trees were pretreated with 4 mM 15N for 1 year and then supplied with unlabeled N for 24 weeks and the remobilization of stored 15N was monitored. On the whole-plant level, uptake of new C was significantly reduced in -N-pretreated trees; however, partitioning of new C was not altered, although there was a trend toward increased belowground respiration. The amount of N taken up was not influenced by N nutrition in the previous year. In +N-pretreated trees, partitioning of new N was dominated by the fine roots (59.7% at Week 12), whereas in -N-pretreated trees, partitioning of new N favored stem, coarse roots and fine roots (24, 21 and 31.9%, respectively, at Week 12), indicating the formation of N stores. The contribution of previous-year N to leaf N was about 15%. The N remobilized for leaf formation had been stored in stem and coarse roots. We conclude that, within a growing season, the growth of beech is strongly determined by the availability of tree internal N stores, whereas the current N supply is of less importance.

Carbon↗

Contrasting distribution and seasonal dynamics of carbohydrate reserves in stem wood of adult ring-porous sessile oak and diffuse-porous beech trees.

We tested the hypothesis that broad-leaved forest species with contrasting wood anatomy and hydraulic system (ring-porous versus diffuse-porous) also differ in distribution and seasonal dynamics of carbohydrate reserves in stem wood. Total nonstructural carbohydrate (TNC) reserves (starch and sugars) were measured enzymatically in the 10 youngest stem xylem rings of adult oak (Quercus petraea (Matt.) Liebl.) and beech (Fagus sylvatica L.) trees during an annual cycle. Radial distribution of carbohydrates was investigated according to ring age. On all dates, oak trees had twofold higher TNC concentration than beech trees (41 versus 23 mg g(DM)(-1)), with starch accounting for the high TNC concentration in oak. Seasonal dynamics of TNC concentration were significantly (P < 0.05) more pronounced in oak (20-64 mg TNC g(DM)(-1)) than in beech (17-34 mg TNC g(DM)(-1)). A marked decrease in TNC concentration was observed in oak trees during bud burst and early wood growth, whereas seasonal fluctuations in TNC concentrations in beech trees were small. The radial distribution of TNC based on ring age differed between species: TNC was restricted to the sapwood rings in oak, whereas in beech, it was distributed throughout the wood from the outermost sapwood ring to the pith. Although the high TNC concentrations in the outermost rings accounted for most of the observed seasonal pattern, all of the 10 youngest xylem rings analyzed participated in the seasonal dynamics of TNC in beech trees. The innermost sapwood rings of oak trees had low TNC concentrations. Stem growth and accumulation of carbon reserves occurred concomitantly during the first part of the season, when there was no soil water deficit. When soil water content was depleted, stem growth ceased in both species, whereas TNC accumulation was negligibly affected and continued until leaf fall. The contrasting dynamics and distribution of carbohydrate reserves in oak and beech are discussed with reference to differences in phenology, early spring growth and hydraulic properties between ring-porous trees and diffuse-porous trees.

Carbohydrate Metabolism↗