Relationship between empty seed and genetic factors in European beech (Fagus sylvatica L.)

Abstract

Replacing Norway spruce with European beech: a comparison of biomass and Net Primary Production patterns in young stands

An alteration of species composition in temperate forests – both managed and natural - is one of the expected effects of environmental change. Present forest tree species ranges will be altered by changing environmental conditions. By a combination of continuous and destructive sampling, we compared biomass stocks and annual NPP in naturally regenerated stands of Norway spruce and European beech. We purposely selected a site where future environmental conditions are predicted to favour beech over presently dominant spruce. We found no difference in overall productivity, but biomass allocation differed significantly between the two species. Beech allocated more assimilates to stem and roots than spruce. There was no significant difference between the species in NPP of the fast turnover biomass pool comprising foliage and fine roots. Maximum height growth occurred about a month earlier than in spruce, potentially changing the timing of carbon (C) flow into the soil pools. We show that the replacement of spruce by beech will result in changes in forest biomass allocation and in alterations of belowground C cycle. Such changes will affect forest ecosystem function by modifying the magnitude and timing of certain C fluxes, but also by potentially changing the species composition of forest biota dependent on them.

Differences in Soil Fungal Communities between European Beech (Fagus sylvatica L.) Dominated Forests Are Related to Soil and Understory Vegetation

Fungi are important members of soil microbial communities with a crucial role in biogeochemical processes. Although soil fungi are known to be highly diverse, little is known about factors influencing variations in their diversity and community structure among forests dominated by the same tree species but spread over different regions and under different managements. We analyzed the soil fungal diversity and community composition of managed and unmanaged European beech dominated forests located in three German regions, the Schwäbische Alb in Southwestern, the Hainich-Dün in Central and the Schorfheide Chorin in the Northeastern Germany, using internal transcribed spacer (ITS) rDNA pyrotag sequencing. Multiple sequence quality filtering followed by sequence data normalization revealed 1655 fungal operational taxonomic units. Further analysis based on 722 abundant fungal OTUs revealed the phylum Basidiomycota to be dominant (54%) and its community to comprise 71.4% of ectomycorrhizal taxa. Fungal community structure differed significantly (p≤0.001) among the three regions and was characterized by non-random fungal OTUs co-occurrence. Soil parameters, herbaceous understory vegetation, and litter cover affected fungal community structure. However, within each study region we found no difference in fungal community structure between management types. Our results also showed region specific significant correlation patterns between the dominant ectomycorrhizal fungal genera. This suggests that soil fungal communities are region-specific but nevertheless composed of functionally diverse and complementary taxa.

Differential Responses of Herbivores and Herbivory to Management in Temperate European Beech

Forest management not only affects biodiversity but also might alter ecosystem processes mediated by the organisms, i.e. herbivory the removal of plant biomass by plant-eating insects and other arthropod groups. Aiming at revealing general relationships between forest management and herbivory we investigated aboveground arthropod herbivory in 105 plots dominated by European beech in three different regions in Germany in the sun-exposed canopy of mature beech trees and on beech saplings in the understorey. We separately assessed damage by different guilds of herbivores, i.e. chewing, sucking and scraping herbivores, gall-forming insects and mites, and leaf-mining insects. We asked whether herbivory differs among different forest management regimes (unmanaged, uneven-aged managed, even-aged managed) and among age-classes within even-aged forests. We further tested for consistency of relationships between regions, strata and herbivore guilds. On average, almost 80 of beech leaves showed herbivory damage, and about 6 of leaf area was consumed. Chewing damage was most common, whereas leaf sucking and scraping damage were very rare. Damage was generally greater in the canopy than in the understorey, in particular for chewing and scraping damage, and the occurrence of mines. There was little difference in herbivory among differently managed forests and the effects of management on damage differed among regions, strata and damage types. Covariates such as wood volume, tree density and plant diversity weakly influenced herbivory, and effects differed between herbivory types. We conclude that despite of the relatively low number of species attacking beech; arthropod herbivory on beech is generally high. We further conclude that responses of herbivory to forest management are multifaceted and environmental factors such as forest structure variables affecting in particular microclimatic conditions are more likely to explain the variability in herbivory among beech forest plots.

Drivers of nitrogen leaching from organic layers in Central European beech forests

Background and Aims: The response of forest ecosystems to continuous nitrogen (N) deposition is still uncertain. We investigated imports and exports of dissolved N from mull-type organic layers to identify the controls of N leaching in Central European beech forests under continuous N deposition. Methods: Dissolved N fluxes with throughfall and through mull-type organic layers (litter leachate) were measured continuously in 12 beech forests on calcareous soil in two regions in Germany over three consecutive growing seasons. Results Mean growing season net (i.e. litter leachate – throughfall flux) fluxes of total dissolved N (TDN) from the organic layer were low (2.3 ± 5.6 kg ha −1 ) but varied widely from 12.9 kg ha −1 to –8.3 kg ha −1 . The small increase of dissolved N fluxes during the water passage through mull-type organic layers suggested that high turnover rates coincided with high microbial N assimilation and plant N uptake. Stand basal area had a positive feedback on N fluxes by providing litter for soil organic matter forma- tion. Plant diversity, especially herb diversity, reduced dissolved N fluxes. Soil fauna biomass increased NO3−-N fluxes with litter leachate by stimulating mineralization. Microbial biomass measures were not related to dissolved N fluxes. Conclusions Our results show that dissolved N exports from organic layers contain significant amounts of throughfall-derived N (mainly NO3−-N) that flushes through the organic layer but also highlight that N leaching from organic layers is driven by the complex interplay of plants, animals and microbes. Furthermore, diverse understories reduce N leaching from Central European beech forests.

What drives European beech (Fagus sylvatica L.) mortality after forest fires of varying severity?

Predicting the timing and amount of tree mortality after a forest fire is of paramount importance for post-fire management decisions, such as salvage logging or reforestation. Such knowledge is particularly needed in mountainous regions where forest stands often serve as protection against natural hazards (e.g., snow avalanches, rockfalls, landslides). In this paper, we focus on the drivers and timing of mortality in fire-injured beech trees (Fagus sylvatica L.) in mountain regions. We studied beech forests in the southwestern European Alps, which burned between 1970 and 2012. The results show that beech trees, which lack fire-resistance traits, experience increased mortality within the first two decades post-fire with a timing and amount strongly related to the burn severity. Beech mortality is fast and ubiquitous in high severity sites, whereas small- (DBH <12 cm) and intermediate-diameter (DBH 12–36 cm) trees face a higher risk to die in moderate-severity sites. Large-diameter trees mostly survive, representing a crucial ecological legacy for beech regeneration. Mortality remains low and at a level similar to unburnt beech forests for low burn severity sites. Beech trees diameter, the presence of fungal infestation and elevation are the most significant drivers of mortality. The risk of beech to die increases toward higher elevation and is higher for small-diameter than for large-diameter trees. In case of secondary fungi infestation beech faces generally a higher risk to die. Interestingly, fungi that initiate post-fire tree mortality differ from fungi occurring after mechanical injury. From a management point of view, the insights about the controls of post-fire mortality provided by this study should help in planning post-fire silvicultural measures in montane beech forests.

Individual biomass factors for beech, oak and pine in Slovakia: a comparative study in young naturally regenerated stands

Biomass conversion and expansion factors (BCEF) which convert tree stem volume to whole tree biomass and biomass allocation patterns in young trees were studied in order to estimate tree and stand biomass in naturally regenerated forests. European beech (Fagus sylvatica L.), Sessile oak (Quercus petraea (Mattuschka) Liebl.) and Scots pine (Pinus sylvestris L.) stands were compared. Seven forest stands of each species were chosen to cover their natural distribution in Slovakia. Species specific BCEF are presented, generally showing a steep decrease in all species in the smallest trees, with the only exception in the case of branch BCEF in beech which grows with increasing tree size. The values of BCEF for all tree compartments stabilise in all species once trees reach about 60-70mm diameter at base. As they grow larger, all species increase their allocation to stem and branches, while decreasing the relative growth of roots and foliage. There are, however, clear differences between species and also between broadleaves and conifers in biomass allocation. This research shows that species specific coefficients must be used if we are to reduce uncertainties in estimates of carbon stock changes by afforestation and reforestation activities.

Biomass partitioning and growth efficiency in four naturally regenerated forest tree species

Current forest growth models and yield tables are almost exclusively based on data from mature trees, reducing their applicability to young and developing stands. To address this gap, young European beech, sessile oak, Scots pine and Norway spruce trees approximately 0 to 10 years old were destructively sampled in a range of naturally regenerated forest stands in Central Europe. Diameter at base and height were first measured in situ for up to 175 individuals per species. Subsequently, the trees were excavated and dry biomass of foliage, branches, stems and roots was measured. Allometric relations were then used to calculate biomass allocation coefficients (BAC) and growth efficiency (GE) patterns in young trees. We found large differences in BAC and GE between broadleaves and conifers, but also between species within these categories. Both BAC and GE are strongly age-specific in young trees, their rapidly changing values reflecting different growth strategies in the earliest stages of growth. We show that linear relationships describing biomass allocation in older trees are not applicable in young trees. To accurately predict forest biomass and carbon stocks, forest growth models need to include species and age specific parameters of biomass allocation patterns.

Specific leaf area and leaf area index in developing stands of Fagus sylvatica L. and Picea abies Karst

European beech (Fagus sylvatica L.) and Norway spruce (Picea abies Karst.) are two of the most ecologically and economically important forest tree species in Europe. These two species co-occur in many locations in Europe, leading to direct competition for canopy space. Foliage characteristics of two naturally regenerated pure stands of beech and spruce with fully closed canopies were contrasted to assess the dynamic relationship between foliage adaptability to shading, stand LAI and tree growth. We found that individual leaf size is far more conservative in spruce than in beech. Individual leaf and needle area was larger at the top than at the bottom of the canopy in both species. Inverse relationship was found for specific leaf area (SLA), highest SLA values were found at lowest light availability under the canopy. There was no difference in leaf area index (LAI) between the two stands, however LAI increased from 10.8 to 14.6 m2m-2 between 2009 and 2011. Dominant trees of both species were more efficient in converting foliage mass or area to produce stem biomass, although this relationship changed with age and was species-specific. Overall, we found larger foliage plasticity in beech than in spruce in relation to light conditions, indicating larger capacity to exploit niche openings.

Simulação numérica do processo de secagem de madeiras

Pós-graduação em Engenharia Mecânica - FEG

Vertical variation in autumn leaf phenology of Fagus sylvatica L. in southern Germany

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Ectomycorrhizae of sessile oak (Quercus petraea (Matt.) Liebl.)

Investigations were performed during the years 1999 to 2001 on a limed and unlimed plot within a high-elevated sessile oak forest. The oak forest (with 90 years old European beech at the understorey) was 170 to 197 years old. It is located at forest district Merzalben, location 04/0705, which is situated in the Palatinate Forest in south-west Germany. Liming was performed in December 1988 when 6 tons/ha of powdered Dolomite were brought up by the forestry department. Liming was performed to counteract the effects of soil acidification (pH(H2O) at Horizon A (0-10 cm): 3.9), which is induced by long-term (anthropogenic) acidic cloud cover and precipitation. Potentially toxic Al3+ ions, which become solubilized below pH 5, were suspected to be responsible for forest dieback and sudden death of the mature oaks. The most logical entry point for these toxic ions was suspected to occur in the highly absorptive region of the ectomycorrhizae (fungal covered root tips). However, the diversity and abundance of oak-ectomycorrhizal species and their actual roles in aluminum translocation (or blockage) were unknown. It was hypothesized that the ectomycorrhizae of sessile oaks in a limed forest would exhibit greater seasonal diversity and abundance with less evidence of incorporated aluminum than similar oak ectomycorrhizae from unlimed soils. To test this hypothesis, 12 oaks in the limed plot and 12 in an adjacent unlimed plot were selected. Each spring and fall for 2 years (1999 & 2000), 2 sets of soil cylinders (9.9 cm dia.) were extracted from Horizon A (0-10 cm), Horizon B (30-40 cm) and Horizon C (50-60 cm depth) at a distance of 1 meter from each tree base. Roots were extracted from each probe by gentle sieving and rinsing. Soil samples were retained for pH (H2O, CaCl2, and KCl) and moisture analysis. One set of roots was sorted by size and air-dried for biomass analysis. The finest mycorrhizal roots of this set were used for bound and unbound (cytosolic) mineral [Al, Ca, Mg, K, Na, Mn, S, Zn, Fe, Cd and Pb] analysis (by Landwirtschaftliche Untersuchungs- und Forschungsanstalt Rheinland Palatinate (LUFA)). Within 7 days of collection, the mycorrhizal tips from the second set of probes were excised, sorted, identified (using Agerer’s Color Atlas), counted and weighed. Seasonal diversity and abundance was characterized for 50 of the 93 isolates. The location and relative abundance of Al within the fungal and root cell walls was characterized for 68 species using 0.01% Morin dye and fluorescence microscopy. Morin complexes with Al to produce an intense yellow fluorescence. The 4 most common species (Cenococcum geophilum, Quercirhiza fibulocsytidiata, Lactarius subdulcis, Piceirhiza chordata) were prepared for bound Al, Ca, Fe and K mineral analysis by LUFA. The unlimed and limed plots were then compared. Only 46 of the 93 isolated ectomycorrhizal species had been previously associated with oaks in the literature. Mycorrhizal biomass was most abundant in Horizon A, declining with depth, drought and progressive soil acidification. Mycorrhizae were most diverse (32 species) in the limed plot, but individual species abundance was low (R Selection) in comparison to the unlimed plot, where there were fewer species (24) but each species present was abundant (K Selection). Liming increased diversity and altered dominance hierarchy, seasonal distributions and succession trends of ectomycorrhizae at all depths. Despite an expected reduction in Al content, the limed ectomycorrhizae both qualitatively (fluorescence analysis) and quantitatively (mineral analysis) contained more bound Al, especially so in Horizon A. The Al content qualitatively and quantitatively increased with depth in the unlimed and limed plots. The bound Al content fluctuated between 4000-and 20000 ppm while the unbound component was consistently lower (4 -14 ppm). The relative amount of unbound Al declined upon liming implying less availability for translocation to the crown area of the trees. This correspouds with the findings of good crown appearance and lower tree mortality in the limed zone. Each ectomycorrhizal species was unique in its ability to block, sequester (hold) or translocate Aluminum. In several species, Al uptake varied with changes in moisture, pH, depth and liming. According to the fluorescence study, about 48% of the isolated ectomycorrhizal species blocked and/or sequestered (held) Al in their mantle and/or Hartig net walls, qualitatively lowering bound Al in the adjacent root cell walls. Generally, if Al was more concentrated in the fungal walls, it was less evident in the cortex and xylem and conversely, if Al was low or absent from the fungal walls it was frequently more evident in the cortex and xylem.

Measurement of nonmethane organic carbon

The present dissertation focuses on the measurement of nonmethane organic carbon compounds (NMOC) and their exchange by biosphere-atmosphere interactions. To access the accuracy, precision, and reproducibility of NMOC analysis, two intercomparison experiments were carried out during the present study. These experiments comprised the sampling of NMOCs on graphitised carbon blacks, followed by gas-chromatographic analysis. Furthermore, they comprised the sampling of short chain carbonyl compounds on solid phase extraction cartridges and their analysis by high pressure liquid chromatography. To investigate the exchange of NMOCs between vegetation and the atmosphere, plant enclosure studies were performed on two European deciduous tree species. These measurements were conducted during two consecutive summer seasons by utilisation of the above specified techniques on sunlit and shaded leaves of European beech (Fagus sylvatica L., monoterpene emitter) and sunlit leaves of English oak (Quercus robur L., isoprene emitter). According to its broad geographical distribution, the impact of European beech on the European monoterpene budget was characterized by a model simulation. Complementary an instrument was developed, that is capable of measuring the amount of total NMOC that is exchanged by biosphere-atmosphere interactions. The instrument was tested under laboratory conditions and was evaluated versus an independent method performing branch enclosure measurements.

Methodical study of nitrous oxide eddy covariance measurements using quantum cascade laser spectrometery over a Swiss forest

Nitrous oxide fluxes were measured at the Lägeren CarboEurope IP flux site over the multi-species mixed forest dominated by European beech and Norway spruce. Measurements were carried out during a four-week period in October–November 2005 during leaf senescence. Fluxes were measured with a standard ultrasonic anemometer in combination with a quantum cascade laser absorption spectrometer that measured N2O, CO2, and H2O mixing ratios simultaneously at 5 Hz time resolution. To distinguish insignificant fluxes from significant ones it is proposed to use a new approach based on the significance of the correlation coefficient between vertical wind speed and mixing ratio fluctuations. This procedure eliminated roughly 56% of our half-hourly fluxes. Based on the remaining, quality checked N2O fluxes we quantified the mean efflux at 0.8±0.4 μmol m−2 h−1 (mean ± standard error). Most of the contribution to the N2O flux occurred during a 6.5-h period starting 4.5 h before each precipitation event. No relation with precipitation amount could be found. Visibility data representing fog density and duration at the site indicate that wetting of the canopy may have as strong an effect on N2O effluxes as does below-ground microbial activity. It is speculated that above-ground N2O production from the senescing leaves at high moisture (fog, drizzle, onset of precipitation event) may be responsible for part of the measured flux.