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Mixed stands of European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst.) frequently over-yield, when compared to respective monospecific stands. Over-yielding is attributed to enhanced resource uptake efficiency through niche complementarity alleviating species competition, for example through enhanced root stratification in mixture. Under severe and frequent summer drought, however, water limitation may become crucial in modifying the prevailing competitive interaction in mixed beech-spruce forests. We hypothesize, therefore, that under drought (H I) inter-specific interaction with beech reduces water accessibility for spruce more than intra-specific conditions, thus (H II) exacerbating drought susceptibility of spruce in terms of reduced photosynthesis and stem growth. Reactions at the organ (leaf, fine root), tree and stand scale were analysed in a mature forest with beech-spruce group mixture. Under inter-specific conditions spruce’s fine-root production and depth of water uptake (assessed via δ18O of xylem water) shifted to shallow, drought-prone soil horizons, in agreement with H I. Overall, lowered fine root production and ramification along with a reduction in long-distance explorative ectomycorrhizal types resulted in decreased soil exploitation in spruce when growing together with beech. Spruce’s drought sensitivity was exemplified by a distinct decrease in stomatal conductance, net CO2 uptake rate and stem growth during periods of water limitation. Notwithstanding, species interaction effects were absent in leaf gas exchange and stem diameter growth, during a six-week summer drought period in 2013 as well as in the extremely dry year of 2003, hence rejecting H II. Based on results from soil moisture measurements and water uptake depth, we interpret the conflicting findings for H I and H II to result from: (i) seasonal shifts between positive (during spring drought) and negative (during summer drought) effects of beech neighbourhood on soil water availability for spruce, possibly overriding each other in their effect on annual stem diameter growth and (ii) the group-wise mixture pattern, where spruce is exposed to competition with beech only along group edges, i.e. laterally only, so that the putatively adverse beech effect on water accessibility stays limited. Our results suggest, compared to single tree mixture, group-wise mixture of beech and spruce to be a favourable silvicultural option in the face of climate change.

Forest ecosystems in central Europe are predicted to face an increasing frequency and severity of summer droughts because of global climate change. European beech and Norway spruce often coexist in these forests with mostly positive effects on their growth. However, their different below-ground responses to drought may lead to differences in ectomycorrhizal (ECM) fungal community composition and functions which we examined at the individual root and ecosystem levels. We installed retractable roofs over plots in Kranzberg Forest (11 degrees 39'42 '' E, 48 degrees 25'12 '' N; 490 m a.s.l.) to impose repeated summer drought conditions and assigned zones within each plot where trees neighboured the same or different species to study mixed species effects. We found that ECM fungal community composition changed and the numbers of vital mycorrhizae decreased for both tree species over 3 drought years (2014-2016), with the ECM fungal community diversity of beech exhibiting a faster and of spruce a stronger decline. Mixed stands had a positive effect on the ECM fungal community diversity of both tree species after the third drought year. Ectomycorrhizae with long rhizomorphs increased in both species under drought, indicating long-distance water transport. However, there was a progressive decline in the number of vital fine roots during the experiment, resulting in a strong reduction in enzyme activity per unit volume of soil. Hydrolytic enzyme activities of the surviving ectomycorrhizae were stable or stimulated upon drought, but there was a large decline in ECM fungal species with laccase activity, indicating a decreased potential to exploit nutrients bound to phenolic compounds. Thus, the ectomycorrhizae responded to repeated drought by maintaining or increasing their functionality at the individual root level, but were unable to compensate for quantitative losses at the ecosystem level. These findings demonstrate a strong below-ground impact of recurrent drought events in forests.

In the present study, we performed a large-scale protein analysis based on 2-DE DIGE to examine the effects of ozone on the leaves of juvenile European beech ( Fagus sylvatica L.), one of the most important deciduous tree species in Central Europe. To this end, beech trees were grown under field conditions and subjected to ambient and twice ambient ozone concentrations during the vegetation periods of four consecutive years. The twice ambient ozone concentration altered the abundance of 237 protein spots, which showed relative ratios higher than 30% compared to the ambient control trees. A total of 74 protein spots were subjected to mass spectrometry identification (LC–MS/MS), followed by homology-driven searches. The differentially expressed proteins participate in key biological processes including the Calvin cycle and photosynthesis, carbon metabolism, defense- and stress-related responses, detoxification mechanisms, protein folding and degradation, and mechanisms involved in senescence. The ozone-induced responses provide evidence of a changing carbon metabolism and counteraction against increased levels of reactive oxygen species. Biological significance This study provides useful information on how European beech, an economically and ecologically important tree species, reacts on the molecular level to increased ozone concentrations expected in the near future. The main emphasis in the present study was placed on identifying differentially abundant proteins after long-term ozone exposure under climatically realistic settings, rather than short-term responses or reactions under laboratory conditions. Additionally, using nursery-grown beech trees, we took into account the natural genotypic variation of this species. As such, the results presented here provide information on molecular responses to ozone in an experimental plant system at very close to natural conditions. Furthermore, this proteomic approach was supported by previous studies on the present experiment. Ultimately, the combination of this proteomic approach with several approaches including transcriptomics, analysis of non-structural carbohydrates, and morphological effects contributes to a more global picture of how beech trees react under increased ozone concentrations.

We review causes of synergies in mixed-species stands, develop guiding hypotheses for revealing their mechanisms and present a rainfall exclusion experiment along with a transect (KROOF) for exploring drought effects. While monocultures have dominated forest research and practice in the past, in face of growing resource scarcity and climate change, mixed-species stands are on the advance. Long-term observations show that mixed-species stands frequently over-yield monocultures, and they further suggest that the over-yielding is often higher on poor than on fertile sites and in low-growth than in high-growth years. However, the underlying causes have not yet been clarified. We start with a review of, among others, hydraulic redistribution, complementary eco-physiological traits, and ectomycorrhizal networks as possible causes behind the observed productivity gains in mixed-species stands. Then, we develop guiding hypotheses for further exploration of synergies in mixed-species stands. Finally, we introduce into the concept and model approach of the rainfall exclusion experiment for examining the role of water supply in mixed versus pure forest stands of spruce and beech. At the Kranzberg site, six plots are equipped with water retention roofs, which only close during rain events. The remaining six plots serve as non-roofed control. Together with the rainfall exclusion experiment, an ecological gradient with five sites extending through SE-Germany forms the “Kranzberg Roof Experiment” (in short KROOF). Kranzberg Forest is a part of this gradient from moist to dry conditions, with each site providing mixed and pure associations each of spruce and beech. The ecophysiological approach will be complemented by tree ring analysis and modelling of productivity of the tree associations under intense drought.

Climate projections predict higher precipitation variability with more frequent dry extremes(1). CO2 assimilation of forests decreases during drought, either by stomatal closure(2) or by direct environmental control of sink tissue activities(3). Ultimately, drought effects on forests depend on the ability of forests to recover, but the mechanisms controlling ecosystem resilience are uncertain(4). Here, we have investigated the effects of drought and drought release on the carbon balances in beech trees by combining CO2 flux measurements, metabolomics and (13)CO2 pulse labelling. During drought, net photosynthesis (AN), soil respiration (RS) and the allocation of recent assimilates below ground were reduced. Carbohydrates accumulated in metabolically resting roots but not in leaves, indicating sink control of the tree carbon balance. After drought release, RS recovered faster than AN and CO2 fluxes exceeded those in continuously watered trees for months. This stimulation was related to greater assimilate allocation to and metabolization in the rhizosphere. These findings show that trees prioritize the investment of assimilates below ground, probably to regain root functions after drought. We propose that root restoration plays a key role in ecosystem resilience to drought, in that the increased sink activity controls the recovery of carbon balances.

Tropospheric ozone, one of the most phytotoxic air pollutants, may specially impose in long-lived forest trees substantial reduction in productivity and biomass. European beech saplings grown in lysimeter around areas were used to monitor proteomic changes upon elevated ozone concentrations following four vegetation periods of exposure. A proteome study based on highly sensitive two-dimensional fluorescence difference gel electrophoresis (2-D DIGE) was performed to identify protein changes in European beech, the most important deciduous tree in Central Europe. Main emphasis was on identifying differentially expressed proteins after long-time period of ozone exposure under natural conditions rather than short-term responses or reactions under controlled conditions. Our results clearly demonstrate a response of European beech saplings to long-term ozone fumigation at the protein level. We indicate changes in the protein abundance of 142 protein spots; among them 59 were increased and 83 decreased following three years of elevated ozone exposure. As the first step, 40 proteins were identified by a homology driven mass spectrometric approach. Some of the identified proteins have been previously described in the context of short-term ozone responses in plants, indicating, at least for certain cellular functions, the congruence of plant reactions following short- and long-term ozone exposure. Under elevated ozone exposure, abundance of proteins related to the Calvin cycle and photosynthetic electron transport chain were decreased whereas the abundance of proteins regarding the carbon metabolism/catabolism were increased.

Cenococcum geophilum is a widely distributed ectomycorrhizal fungus potentially playing a significant role in resistance and resilience mechanisms of its tree hosts exposed to drought stress. In this study, we performed a large scale protein analysis in pure cultures of C. geophilum in order to gain first global insights into the proteome assembly of this fungus. Using 1-D gel electrophoresis coupled with ESI-MS/MS, we indentified 638 unique proteins. Most of these proteins were related to the metabolic and cellular processes, and the transport machinery of cells. In a second step, we examined the influence of water deprivation on the proteome of C. geophilum pure cultures at three time points of gradually imposed drought. The results indicated that 12 proteins were differentially abundant in mycelia subjected to drought compared to controls. The induced responses in C. geophilum point towards regulation of osmotic stress, maintainance of cell integrity, and counteracting increased levels of reactive oxygen species formed during water deprivation. (c) 2012 Elsevier B.V. All rights reserved.

International audience; The contribution of proteomics to the knowledge of forest tree (the most recalcitrant and almost forgotten plant species) biology is being reviewed and discussed, based on the author’s own research work and papers published up to November 2010. This review is organized in four introductory sections starting with the definition of forest trees (1), the description of the environmental and economic importance (2) and its derived current priorities and research lines for breeding and conservation (3) including forest tree genomics (4). These precede the main body of this review: a general overview to proteomics (5) for introducing the forest tree proteomics section (6). Proteomics, defined as scientific discipline or experimental approach, it will be discussed both from a conceptual and methodological point of view, commenting on realities, challenges and limitations. Proteomics research in woody plants is limited to a reduced number of genera, including Pinus, Picea, Populus, Eucalyptus, and Fagus, mainly using first-generation approaches, e.g., those based on two-dimensional electrophoresis coupled to mass spectrometry. This area joins the own limitations of the technique and the difficulty and recalcitrance of the plant species as an experimental system. Furthermore, it contributes to a deeper knowledge of some biological processes, namely growth, development, organogenesis, and responses to stresses, as it is also used in the characterization and cataloguing of natural populations and biodiversity (proteotyping) and in assisting breeding programmes.