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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.

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.