Arthur Geßler

Potential risks for European beech (Fagus sylvatica L.) in a changing climate

Over large areas of Europe, coniferous monocultures are being transformed into mixed forests by the re-introduction of broadleaf tree species belonging to the potential natural vegetation. One important species of interest in this changing forest policy is European beech (Fagus sylvatica). However, at present, this forest management directive has ignored potential adverse effects of global climate change on wide-spread re-introduction of beech to these areas. Average global surface temperatures have risen by approx. 0.8°C in the period between 1861 and 2005 and are expected to continue to increase until the end of this century by 1.5–5.8°C above the 1990 value. To estimate the climate change in the southern part of central Europe in future, we reviewed calculations from regional climate models. Temperature increase for the southern part of central Europe is projected to be up to 2°C within the next 40 years. In contrast, the annual precipitation will most likely remain constant over the same time period, but will experience significant changes in seasonal patterns. Rising intensities of individual precipitation events may result in increasing number and intensities of flooding events and reduced precipitation during the growing season in a higher frequency of summer droughts. Growth and competitive ability of European beech will not, necessarily, respond to increasing CO2 concentrations but may be strongly impacted by intensive drought that occurs during the growing season. Seedlings as well as adult trees may suffer from xylem embolism, restricted nutrient uptake capacity and reduced growth under limited water availability. However, it remains uncertain to what extent other environmental factors (e.g. soil properties, competitive interactions) may modify the drought response of beech, thus either enhancing susceptibility or increasing drought tolerance and resilience potential. Water-logged soils, predicted during the spring for several regions due to higher than average precipitation, could negatively impact nutrient uptake and growth of beech. Whereas other dominant species as, e.g. oak are well adapted to that environmental stress, beech is known to be sensitive to water-logging and flooding. Thus, the competitive capacity of beech might—depending on the other environmental conditions—be reduced under the expected future climate conditions. Silvicultural practices must be aware today of the potential risks which a changing climate may impose on sustainable forest development.

Climate and forest management influence nitrogen balance of European beech forests: microbial N transformations and inorganic N net uptake capacity of mycorrhizal roots

The effects of local climate and silvicultural treatment on the inorganic N availability, net N uptake capacity of mycorrhizal beech roots and microbial N conversion were assessed in order to characterise changes in the partitioning of inorganic N between adult beech and soil microorganisms. Fine root dynamics, inorganic N in the soil solution and in soil extracts, nitrate and ammonium uptake kinetics of beech as well as gross ammonification, nitrification and denitrification rates were determined in a beech stand consisting of paired sites that mainly differed in aspect (SW vs. NE) and stand density (controls and thinning treatments). Nitrate was the only inorganic N form detectable in the soil water. Its concentration was high in control plots of the NE aspect, but only in canopy gaps and not influenced by thinning. Neither thinning nor aspect affected the abundance of root tips in the soil. Maximum nitrate net uptake by mycorrhizal fine roots of beech, however, differed with aspect, showing significantly lower values at the SW aspect with warm–dry local climate. There were no clear-cut significant effects of local climate or thinning on microbial N conversion, but a tendency towards higher ammonification and nitrification and lower denitrification rates on the untreated controls of the SW as compared to the NE aspect. Apparently, the observed sensitivity of beech towards reduced soil water availability is at least partially due to impaired N acquisition. This seems to be mainly a consequence of reduced N uptake capacity rather than of limited microbial re-supply of inorganic N or of changed patterns of inorganic N partitioning between soil bacteria and roots.

Climate and forest management affect 15N-uptake, N balance and biomass of European beech seedlings

We investigated the effect of (a) different local climate and (b) thinning of the forest canopy on growth and N status of naturally regenerated European beech seedlings in a beech forest on shallow rendzina soil in southern Germany. For this purpose, a 15N-tracing experiment was conducted during the growing season of the year 2000 with beech seedlings growing on a warm, dry SW-exposed site and a cooler, moist NE-exposed site, and in a thinned and a control stand at each site. Biomass, 15N uptake and partitioning and total N concentrations of beech seedlings were determined. Site and thinning produced clear differences, particularly at the end of the growing season. Biomass and cumulative 15N uptake of beech seedlings then increased due to thinning on the NE site and decreased on the SW site. Total N concentrations in leaves, roots and stems of beech seedlings responded similarly. Therefore, growth and N status of beech seedlings are found to be favoured by thinning under cool-moist conditions. However, under higher temperature and reduced water availability—conditions that are prognosticated in the near future—thinning reduces N uptake and plant N concentration and, thus, impairs N balance and growth of beech regeneration.

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.

Seasonal courses of key parameters of nitrogen, carbon and water balance in European beech (Fagus sylvatica L.) grown on four different study sites along a European North–South climate gradient during the 2003 drought

During the growing season of the exceptionally dry and warm year 2003, we assessed seasonal changes in nitrogen, carbon and water balance related parameters of mature naturally grown European beech (Fagus sylvatica L.) along a North–South transect in Europe that included a beech forest stand in central Germany, two in southern Germany and one in southern France. Indicators for N balance assessed at all four sites were foliar N contents and total soluble non-protein nitrogen compounds (TSNN) in xylem sap, leaves and phloem exudates; C and water balance related parameters determined were foliar C contents, δ13C and δ18O signatures. Tissue sampling was performed in May, July and September. The N related parameters displayed seasonal courses with highest concentrations during N remobilization in May. Decreased total foliar N contents as well as higher C/N ratios in the stands in central Germany and southern France compared to the other study sites point to an impaired N nutrition status due to lower soil N contents and precipitation perception. TSNN concentrations in leaves and phloem exudates of all study sites were in ranges previously reported, but xylem sap content of amino compounds in July was lower at all study sites when compared to literature data (c. 1 μmol N mL−1). In September, TSNN concentrations increased again at the two study sites in southern Germany after a rain event, whereas they remained constant at sites in central Germany and southern France which hardly perceived precipitation during that time. Thus, TSNN concentrations in the xylem sap might be indicative for water balance related N supply in the beech trees. TSNN profiles at all study sites, however, did not indicate drought stress. Foliar δ13C, but not foliar C and δ18O followed a seasonal trend at all study sites with highest values in May. Differences in foliar δ13C and δ18O did not reflect climatic differences between the sites, and are attributed to differences in altitude, photosynthesis and δ18O signatures of the water sources. Except of low TSNN concentrations in the xylem sap, no physiological indications of drought stress were detected in the trees analysed. We suppose that the other parameters assessed might not have been sensitive to the drought events because of efficient regulation mechanisms that provide a suitable physiological setting even under conditions of prolonged water limitation. The uniform performance of the trees from southern France and central Germany under comparably dry climate conditions denotes that the metabolic plasticity of mature beech from the different sites studied might be similar.

The Apoplast of Norway Spruce (Picea Abies) Needles as Habitat and Reaction Compartment for Autotrophic Nitrifiers

By the combination of both, the molecular biological fluorescence in-situ hybridization-technique with the technique of confocal laser scanning-microscopy (cLSM) it could be unequivocally demonstrated for the first time at the genetic level that autotrophic nitrifiers are present inside spruce needles of a spruce forest ecosystem (The “Hoglwald”) exposed to high levels of atmospheric nitrogen deposition and that they are located within the apoplastic space of the needle leaves (sub-stomatal cavity). In contrast, autotrophic nitrifers could not be detected in needles of adult trees at a spruce forest site (“Villingen”) exposed to low levels of atmospheric N input. When needles from adult spruce trees at the Hogwald site were exposed to 10 Pa acetylene – an inhibitor of ammonia monooxigenase of chemolithoautotrophic ammonia oxidisers (CAO) – the sink strength of the needles for NH3 decreased significantly. Since the reduction of NH3 deposition due to acetylene-induced inhibition of the ammonia monooxigenase was greatest when stomata were open and only minute when stomata were closed, it is concluded that physiologically active CAO are located inside the needles rather than on the needle surface. On the other hand, a reduction of NH3 uptake when applying acetylene was not observed with adult spruce trees from the nitrogen limited stand at Villingen. From the results obtained it is concluded that the observed NH3 flux from the atmosphere into the needle leaves in N-polluted forests is not exclusively a plant physiological process as has been assumed in the past, but is the result of both plant physiological plus microbial processes. A seasonal pattern of the colonization of the needles by nitrifiers at the Hoglwald site could not be demonstrated. For gaining first insights into the pathway by which needles might be colonized by autotrophic nitrifiers, sterile spruce seedlings fumigated with both ammonia and air were inoculated in the laboratory with nitrifier cultures. A successful establishment of the autotrophic nitrifiers in the phyllosphere of the spruce seedlings could not be achieved by a single inoculation event, however, was successful after multiple inoculations. It is concluded that autotrophic nitrifiers obviously are only able to colonize spruce needles in a later stage of spruce development.