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The ability of treeline conifers in the Central European Alps to cope with recent climate warming and increasing CO2 concentration is still poorly understood. We determined basal area increment (BAI) and tree ring stable carbon isotope ratios (δ13C) of Pinus cembra trees from 1925 through 2013. Stable isotope ratios and BAI were compared with leaf level gas exchange measurements carried out in situ between 1934 and 2012, and thus, provided new insights into long-term trends of tree-ring derived intrinsic water-use efficiency (iWUE). Mature P. cembra trees at treeline responded to increasing Ca and air temperature with a parallel increase in maximum net CO2 uptake rate at ambient CO2 (Amax) and tree-ring-derived intercellular CO2 partial pressure (Ci). Amax tripled and was positively correlated to BAI and Ci. The latter increased in parallel with ambient CO2 concentration and stomatal conductance. In contrast to the instantaneous gas exchange parameters, δ13C derived iWUE informs about the long-term changes in the carbon water relations. These data showed three changes in the iWUE chronosequences, which could be identified with different long term gas exchange patterns: (1) from stomatal controlled functioning from 1925 to 1981, to a situation where (2) both net CO2 fixation (A) and leaf conductance for water vapour (gw), responded to the environment from 1982 to 1997, and (3) back to a stomata controlled pattern over iWUE from 1998 onwards. This temporal pattern was also mirrored in leaf level gas exchange assessments, suggesting a parallel increase of A and gw of P. cembra at treeline during the last nine decades.

Under drought, potential plant death from depletion of carbon (C) stores, referred to as carbon starvation, is thought to result from negative carbon balance during (partial) stomatal closure. As evidence for C starvation is rare for mature trees, we investigated the C dynamics in mature beech and spruce under drought, focusing on non-structural carbohydrates (NSC) as an indicator of their C balance. Trees were exposed to complete exclusion of precipitation throughfall over two consecutive growing seasons. We assessed NSC concentrations during the early dormant season in leaves, twigs, stem phloem/xylem, coarse and fine roots. Up-scaling to whole-tree NSC pools was achieved using a process-based single-tree model (“BALANCE”), estimating tree biomass increment. While there were distinct differences in NSC concentration among different tissues in both beech and spruce (root

Tree roots and mycorrhizal fungi form an important resource for soil microinvertebrates; however, there is little knowledge on the role of root feeders and microbial grazers on belowground carbon flux of forest trees. Using oak (Quercus robur) microcuttings (Clone DF159) inoculated with an ectomycorrhizal symbiont partner (Piloderma croceum), the present study investigated the impact of multitrophic interactions in the rhizosphere on the carbon flow to the soil food web in a pulse-chase labelling experiment. Changes in plant biomass, microbial communities (soil phospholipid fatty acids - PLFAs) and Collembola lipid pattern (total lipid fatty acids - TLFAs) as well as the 13C incorporation into fatty acids were assessed at 2, 5 and 20 days post labelling of oak microcuttings. Microcosms were inoculated with Pratylenchus penetrans (root-feeding nematode), Protaphorura armata (fungal-feeding Collembola) or their combination. The interaction of P. penetrans and P. armata synergistically reduced oak biomass. Both soil animals independently impacted growth of microorganisms in the rhizosphere, P. penetrans inhibited whilst P. armata promoted bacterial biomass. Irrespective of treatment and time, Gram-positive bacteria derived most of the recent photoassimilated oak carbon, with amounts ranging between 80 and 92% of the total 13C allocated in microbial PLFAs. Presence of either P. armata or P. penetrans enhanced the incorporation of plant carbon in bacteria, whereas collective effects of these two functional groups was scarce. Predominantly the impact of the root-feeding nematode on belowground carbon flow diminished with time. In sum, root feeders and detritivores played independent roles in carbon allocation patterns and community structure of microorganisms in the rhizosphere of oaks.

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.

As long-lived organisms, trees use resources to support both growth and below- and aboveground trophic interactions. Resources fluctuate in relation to periods of growth cease that are regulated by internal and external factors, and these fluctuations feed backs to trophic partners. Some major forest trees display an endogenous growth rhythm, and related pulses of variation in allocation of resources have been detected. As this trait makes it possible to separate growth into defined phases, it offers an opportunity to disentangle the intermingled complex regulation of growth and multitrophic interactions in trees. We present “TrophinOak”, a platform using microcuttings of pedunculated oak, a tree that displays endogenous rhythmic growth characterized by alternating shoot and root growth flushes. We select seven beneficial or detrimental above- and belowground partners including animals (Lymantria dispar, Pratylenchus penetrans, Protaphorura armata), fungi (Piloderma croceum, Microsphaera alphitoides, Phytophthora quercina) and bacteria (Streptomyces sp.), to synthesize bi- and tripartite trophic interactions, including ectomycorrhizal symbioses, and monitor fluctuations of carbon and nitrogen allocation as well as plant gene expression at distinct phases of oak growth. We use this model to identify and resolve the experimental challenges inherent in synthesizing diverse types of associations in a common microcosm system, in labeling plants with stable N and C isotopes and in analyzing transcripts in a non-model plant, a process which requires generating a specific contig library. We develop hypotheses and experimental design to test them in order to identify core mechanisms that help trees to modulate their own development and their multitrophic interactions for optimizing their long term performance in their environment. First results constitute a proof of concept that the platform works and enables us to test the hypotheses.

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 hypothesize that reductions in stand density through thinning improve the recovery of radial stem growth in Norway spruce trees ( Picea abies ) from severe drought. However, thinning may not lead to higher relative radial growth during drought. Annual stem growth and stable carbon and oxygen isotopes in early- and latewood were assessed in trees from heavily thinned (HT), moderately thinned (MT) and un-thinned control stands at two sites in southern Germany. Physiological performance of trees as inferred from stable isotope analysis was used to interpret annual stem growth in response to the drought events in 1976 and 2003. Only in recently thinned stands, trees maintained growth probably through higher soil water availability during the drought year when compared to controls. In contrast, thinning improved the growth recovery in the years following the drought irrespective of the time span between thinning and drought. We conclude that thinning improves drought recovery response in the short and long term and should be considered as an effective management strategy to increase drought tolerance of Norway spruce stands.

The effects of elevated O3 on photosynthetic properties in adult beech trees (Fagus sylvatica) were investigated in relation to leaf mass per area as a measure of the gradually changing, within-canopy light availability. Leaves under elevated O3 showed decreased stomatal conductance at unchanged carboxylation capacity of Rubisco, which was consistent with enhanced delta 13C of leaf organic matter, regardless of the light environment during growth. In parallel, increased energy demand for O3 detoxification and repair was suggested under elevated O3 owing to enhanced dark respiration. Only in shade-grown leaves,light-limited photosynthesis was reduced under elevated O3, this effect being accompanied by lowered F(v)/F(m). These results suggest that chronic O3 exposure primarily caused stomatal closure to adult beech trees in the field regardless of the within-canopy light gradient. However, light limitation apparently raised the O3 sensitivity of photosynthesis and accelerated senescence in shade leaves.

The current AOT40 concept for inferring risks in forest trees by ozone (O3) injury is based on an accumulated external O3 exposure rather than an internal O3 dose or uptake rate. AOT40 assumes O3 concentrations below 40 nl l−1 and night-time exposure to be negligible. Hence, this concept is rather inconsistent with observed forest conditions. In contrast, the flux concept of cumulative O3 uptake (CU) into the leaves has the potential of reflecting a physiologically meaningful internal O3 dose experienced by trees. In this paper, we relate AOT40 to cumulative O3 uptake into European beech (Fagus sylvatica), Norway spruce (Picea abies), European larch (Larix decidua) and cembran pine (Pinus cembra) trees differing in size, age and site conditions. We demonstrate that the flux concept can be extended to the tree and the stand level, making use of sap flow measurements through tree trunks. Although in both seedlings and adult trees AOT40 may show some linearity in correlations with average CU, the latter varies, at given AOT40, by 25±11% within and between species. This is because O3 flux is primarily influenced by stomatal aperture, the latter being affected by climate, canopy position, leaf and tree age while varying between species. In particular, if weighed by detoxification capacity, we suggest, therefore, O3 uptake related air quality indices to be promoted towards ecologically meaningful standards in forest protection, overcoming the shortcomings of exposure concepts. As O3 injury results from the balance between O3 uptake and detoxification in the leaf mesophyll, we conclude the flux concept in combination with measures of biochemical defence to have the capacity for predicting tree response to O3 stress.