Impact of various microplastics on the morphological characteristics and nutrition of the young generation of beech (Fagus sylvatica L.).

Microplastics have the capacity to accumulate in soil due to their high resistance to degradation, consequently altering soil properties and influencing plant growth. This study focused on assessing the impact of various types and doses of microplastics on beech seedling growth. In our experiment, we used polypropylene and styrene granules with diameter of 4.0 mm in quantities of 2.5% and 7%. The hypothesis was that microplastics significantly affect seedlings' nutritional status and growth characteristics. The research analysed seedlings' nutrition, root morphological features, above-ground growth, and enzymatic activity in the substrate. Results confirmed the importance of microplastics in shaping the nutritional status of young beech trees. Microplastic type significantly impacted N/P and Ca/Mg stoichiometry, while microplastic quantity influenced Ca/Al and Ca+K+Mg/Al stoichiometry. Notably, only in the case of root diameter were significantly thicker roots noted in the control variant, whereas microplastics played a role in shaping the leaves' characteristics of the species studied. The leaf area was significantly larger in the control variant compared to the variant with polypropylene in the amount of 2.5% and styrene in the amount of 7%. Additionally, the study indicates a significant impact of microplastics on enzyme activity. In the case of CB and SP, the activity was twice as high in the control variant compared to the variants with microplastics. In the case of BG, the activity in the control variant was higher in relation to the variants used in the experiment. Research on the impact of microplastics on the growth of beech seedlings is crucial for enhancing our understanding of the effects of environmental pollution on forest ecosystems. Such studies are integral in shaping forestry management practices and fostering a broader public understanding of the ecological implications of plastic pollution.

Quantifying the resistance of mixed-forests against natural hazards in the Pyrenees.

Mountain protection forests can prevent natural hazards by reducing their onset and propagation probabilities. In fact, individual trees act as natural barriers against hydrogeomorphic events. However, assessing the structural strength of trees against these hazards is challenging, especially in a context of climate change due to the intensification of extreme events and changes in forest dynamics. Here, we focus on the mechanical analyses of two of the most common tree species across the Pyrenees (Abies alba Mill. and Fagus sylvatica L.) growing in two different areas (Spain and France), and affected by recurrent snow avalanche and rockfall events. We first performed 53 pulling test on mature trees, where the root-plate stiffness and the modulus of elasticity of the stems were evaluated. To further analyse the impact of forest management and climate on protective forests, we yielded information on tree growth using dendroecology techniques. Then, we assessed structure and neighbourhood characteristics for each target tree to account for the surrounding forest structure. Finally, using linear and structured equation models we tested if the mechanical capacity of the trees is determined either by functional traits (e.g. species, tree growth, diameter and height) or forest structural traits (e.g. tree density, tree structure and slenderness) or both. Our results suggest that the forest neighbourhood influences tree mechanical capacity through two pathways, including both functional and structural traits. The individual stiffness parameter of trees is influenced by their functional traits, while their structural traits are more closely related with changes in the modulus of elasticity. Both species exhibit varying levels of dominance in different locations, which is related to their resilience to the diverse natural hazards they confront. Our findings provide relevant insights to anticipating management strategies for forests that serve as a protective barrier against natural hazards in the context of a changing climate.

Tail-dependence of masting synchrony results in continent-wide seed scarcity.

Spatial synchrony may be tail-dependent, meaning it is stronger for peaks rather than troughs, or vice versa. High interannual variation in seed production in perennial plants, called masting, can be synchronized at subcontinental scales, triggering extensive resource pulses or famines. We used data from 99 populations of European beech (Fagus sylvatica) to examine whether masting synchrony differs between mast peaks and years of seed scarcity. Our results revealed that seed scarcity occurs simultaneously across the majority of the species range, extending to populations separated by distances up to 1800 km. Mast peaks were spatially synchronized at distances up to 1000 km and synchrony was geographically concentrated in northeastern Europe. Extensive synchrony in the masting lower tail means that famines caused by beech seed scarcity are amplified by their extensive spatial synchrony, with diverse consequences for food web functioning and climate change biology.

Genotypic Identification of Trees Using DNA Barcodes and Microbiome Analysis of Rhizosphere Microbial Communities.

DNA barcodes can provide accurate identification of plants. We used previously reported DNA primers targeting the internal transcribed spacer (ITS1) region of the nuclear ribosomal cistron, internal transcribed spacer (ITS2), and chloroplast trnL (UAA) intron to identify four trees at Bergen Community College. Two of the four trees were identified as Acer rubrum and Fagus sylvatica. However, Quercus was only identified at the genus level, and the fourth tree did not show similar identification between barcodes. Next-generation sequencing of 16S rRNA genes showed that the predominant bacterial communities in the rhizosphere mainly consisted of the Pseudomonadota, Actinomycetota, Bacteroidota, and Acidobacteriota. A. rubrum showed the most diverse bacterial community while F. sylvatica was less diverse. The genus Rhodoplanes showed the highest relative bacterial abundance in all trees. Fungal ITS sequence analysis demonstrated that the communities predominantly consisted of the Ascomycota and Basidiomycota. Quercus showed the highest fungi diversity while F. sylvatica showed the lowest. Russula showed the highest abundance of fungi genera. Average similarity values in the rhizosphere for fungi communities at the phylum level were higher than for bacteria. However, at the genus level, bacterial communities showed higher similarities than fungi. Similarity values decreased at lower taxonomical levels for both bacteria and fungi, indicating each tree has selected for specific bacterial and fungal communities. This study confirmed the distinctiveness of the microbial communities in the rhizosphere of each tree and their importance in sustaining and supporting viability and growth but also demonstrating the limitations of DNA barcoding with the primers used in this study to identify genus and species for some of the trees. The optimization of DNA barcoding will require additional DNA sequences to enhance the resolution and identification of trees at the study site.

Coping with extremes: Responses of Quercus robur L. and Fagus sylvatica L. to soil drought and elevated vapour pressure deficit.

Climate change, particularly droughts and heat waves, significantly impacts global photosynthesis and forest ecosystem sustainability. To understand how trees respond to and recover from hydrological stress, we investigated the combined effects of soil moisture and atmospheric vapour pressure deficit (VPD) on seedlings of the two major European broadleaved tree species Fagus sylvatica (FS) and Quercus robur (QR). The experiment was conducted under natural forest gap conditions, while soil water availability was strictly manipulated. We monitored gas exchange (net photosynthesis, stomatal conductance and transpiration rates), nonstructural carbohydrates (NSC) concentration in roots and stomatal morphometry (size and density) during a drought period and recovery. Our comparative empirical study allowed us to distinguish and quantify the effects of soil drought and VPD on stomatal behavior, going beyond theoretical models. We found that QR conserved water more conservatively than FS by reducing transpiration and regulating stomatal conductance under drought. FS maintained higher stomatal conductance and transpiration at elevated VPD until soil moisture became critically low. QR showed higher intrinsic water use efficiency than FS. Stomata density and size also likely played a role in photosynthetic rate and speed of recovery, especially since QR with its seasonal adjustments in stomatal traits (smaller, more numerous stomata in summer leaves) responded and recovered faster compared to FS. Our focal species showed different responses in NSC content under drought stress and recovery, suggesting possible different evolutionary pathways in coping with stress. QR mobilized soluble sugars, while FS relied on starch mobilization to resist drought. Although our focal species often co-occur in mixed forests, our study showed that they have evolved different physiological, morphological and biochemical strategies to cope with drought stress. This suggests that ongoing climate change may alter their competitive ability and adaptive potential in favor of one of the species studied.

European beech spring phenological phase prediction with UAV-derived multispectral indices and machine learning regression.

Acquiring phenological event data is crucial for studying the impacts of climate change on forest dynamics and assessing the risks associated with the early onset of young leaves. Large-scale mapping of forest phenological timing using Earth observation (EO) data could enhance our understanding of these processes through an added spatial component. However, translating traditional ground-based phenological observations into reliable ground truthing for training and validating EO mapping applications remains challenging. This study explored the feasibility of predicting high-resolution phenological phase data for European beech (Fagus sylvatica) using unoccupied aerial vehicle (UAV)-based multispectral indices and machine learning. Employing a comprehensive feature selection process, we identified the most effective sensors, vegetation indices, training data partitions, and machine learning models for phenological phase prediction. The model that performed best and generalized well across various sites utilized Green Chromatic Coordinate (GCC) and Generalized Additive Model (GAM) boosting. The GCC training data, derived from the radiometrically calibrated visual bands of a multispectral sensor, were predicted using uncalibrated RGB sensor data. The final GCC/GAM boosting model demonstrated capability in predicting phenological phases on unseen datasets within a root mean squared error threshold of 0.5. This research highlights the potential interoperability among common UAV-mounted sensors, particularly the utility of readily available, low-cost RGB sensors. However, considerable limitations were observed with indices that implement the near-infrared band due to oversaturation. Future work will focus on adapting models to better align with the ICP Forests phenological flushing stages.

Forest genomics in the Caucasus through the lens of its dominant tree species - Fagus orientalis.

The last glacial period is known to have greatly influenced the demographic history of temperate forest trees, with important range contractions and post-glacial expansions that led to the formation of multiple genetic lineages and secondary contact zones in the Northern Hemisphere. These dynamics have been extensively studied for European and North American species but are still poorly understood in other temperate regions of rich biodiversity such as the Caucasus. Our study helps filling that gap by deciphering the genomic landscapes of F. orientalis across the South Caucasus. The use of genome-wide data confirmed a past demographic history strongly influenced by the Last Glacial Maximum, revealing two disjunct glacial refugia in the Colchis and Hyrcanian regions. The resulting patterns of genetic diversity, load and differentiation are not always concordant across the region, with genetic load pinpointing the location of the glacial refugia more efficiently than genetic diversity alone. The Hyrcanian forests show depleted genetic diversity and substantial isolation, even if long-distance gene flow is still present with the main centre of diversity in the Greater Caucasus. Finally, we characterize a strong heterogeneity of genetic diversity and differentiation along the species chromosomes, with noticeably a first chromosome showing low diversity and weak differentiation.

Collimonas rhizosphaerae sp. nov., a novel species isolated from the beech rhizosphere.

Bacterial strain H4R21T was isolated from beech rhizosphere soil sampled in the forest experimental site of Montiers (Meuse, France). It effectively weathers minerals, hydrolyses chitin and produces quorum sensing signal molecules. The strain is aerobic and Gram-stain-negative. Phylogenetic analysis based on its 16S rRNA gene sequence indicated that strain H4R21T belongs to the genus Collimonas with high sequence similarity to C. arenae Ter10T (99.38 %), C. fungivorans Ter6T(98.97 %), C. pratensis Ter91T (98.76 %), C. humicola RLT1W51T (98.46 %) and C. silvisoli RXD178 T (98.46 %), but less than 98 % similarity to other strains of the genus Collimonas. The predominant quinone in H4R21T is ubiquinone-8 (Q8). The major polar lipids are diphosphatidylglycerol, phosphatidylethanolamine, diphosphatidylglycerol, phosphatidylglycerol and lipid. The major fatty acids identified were C12 : 0, C12:0 3-OH, C16  :  0 and C17:0 cyclo. The digital DNA G+C content of the genomic DNA was 59.5 mol%. Furthermore, the strain could be clearly distinguished from its closely related type strains by a combination of phylogenomic and in silico DNA-DNA hybridization results, and phenotypic characteristics. Therefore, strain H4R21T represents a novel species within the genus Collimonas, for which the name Collimonas rhizosphaerae sp. nov. is proposed, with strain H4R21T (=CFBP 9203T=DSM 117599T) as the type strain.

Tip-to-base conduit widening remains consistent across cambial age and climates in Fagus sylvatica L.

Water transport, mechanical support and storage are the vital functions provided by the xylem. These functions are carried out by different cells, exhibiting significant anatomical variation not only within species but also within individual trees. In this study, we used a comprehensive dataset to investigate the consistency of predicted hydraulic vessel diameter widening values in relation to the distance from the tree apex, represented by the relationship Dh ∝ Lß (where Dh is the hydraulic vessel diameter, L the distance from the stem apex and ß the scaling exponent). Our analysis involved 10 Fagus sylvatica L. trees sampled at two distinct sites in the Italian Apennines. Our results strongly emphasize that vessel diameter follows a predictable pattern with the distance from the stem apex and ß ~ 0.20 remains consistent across cambial age and climates. This finding supports the hypothesis that trees do not alter their axial configuration represented by scaling of vessel diameter to compensate for hydraulic limitations imposed by tree height during growth. The study further indicates that within-tree variability significantly contributes to the overall variance of the vessel diameter-stem length exponent. Understanding the factors that contribute to the intraindividual variability in the widening exponent is essential, particularly in relation to interspecific responses and adaptations to drought stress.

Unravelling the thermodynamic properties of soil ecosystems in mature beech forests.

Thermodynamics is a vast area of knowledge with a debatable role in explaining the evolution of ecosystems. In the case of soil ecosystems, this role is still unclear due to difficulties in determining the thermodynamic functions that are involved in the survival and evolution of soils as living systems. The existing knowledge is largely based on theoretical approaches and has never been applied to soils using thermodynamic functions that have been experimentally determined. In this study, we present a method for the complete experimental thermodynamic characterization of soil organic matter. This method quantifies all the thermodynamic functions for combustion and formation reactions which are involved in the thermodynamic principles governing the evolution of the universe. We applied them to track the progress of soil organic matter with soil depth in mature beech forests. Our results show that soil organic matter evolves to a higher degree of reduction as it is mineralized, yielding products with lower carbon but higher energy content than the original organic matter used as reference. These products have higher entropy than the original one, demonstrating how the soil ecosystem evolves with depth, in accordance with the second law of thermodynamics. The results were sensitive to soil organic matter transformation in forests under different management, indicating potential applicability in elucidating the energy strategies for evolution and survival of soil systems as well as in settling their evolutionary states.

Ectomycorrhizal fungi of Douglas-fir retain newly assimilated carbon derived from neighboring European beech.

Ectomycorrhizal (ECM) fungi distribute tree-derived carbon (C) via belowground hyphal networks in forest ecosystems. Here, we asked the following: (1) Is C transferred belowground to a neighboring tree retained in fungal structures or transported within the recipient tree? (2) Is the overlap of ectomycorrhizal fungi in mycorrhizal networks related to the amount of belowground C transfer? We used potted sapling pairs of European beech (Fagus sylvatica) and North-American Douglas-fir (Pseudotsuga menziesii) for 13CO2 pulse-labeling. We compared 13C transfer from beech (donor) to either beech or Douglas-fir (recipient) and identified the ECM species. We measured the 13C enrichment in soil, plant tissues, and ECM fractions of fungal-containing parts and plant transport tissues. In recipients, only fungal-containing tissue of ectomycorrhizas was significantly enriched in 13C and not the plant tissue. Douglas-fir recipients shared on average one ECM species with donors and had a lower 13C enrichment than beech recipients, which shared on average three species with donors. Our results support that recently assimilated C transferred belowground is shared among fungi colonizing tree roots but not among trees. In mixed forests with beech and Douglas-fir, the links for C movement might be hampered due to low mycorrhizal overlap with consequences for soil C cycling.

Recover of Soil Microbial Community Functions in Beech and Turkey Oak Forests After Coppicing Interventions.

Forest management influences the occurrence of tree species, the organic matter input to the soil decomposer system, and hence, it can alter soil microbial community and key ecosystem functions it performs. In this study, we compared the potential effect of different forest management, coppice and high forest, on soil microbial functional diversity, enzyme activities and chemical-physical soil properties in two forests, turkey oak and beech, during summer and autumn. We hypothesized that coppicing influences soil microbial functional diversity with an overall decrease. Contrary to our hypothesis, in summer, the functional diversity of soil microbial community was higher in both coppice forests, suggesting a resilience response of the microbial communities in the soil after tree cutting, which occurred 15-20 years ago. In beech forest under coppice management, a higher content of soil organic matter (but also of soil recalcitrant and stable organic carbon) compared to high forest can explain the higher soil microbial functional diversity and metabolic activity. In turkey oak forest, although differences in functional diversity of soil microbial community between management were observed, for the other investigated parameters, the differences were mainly linked to seasonality. The findings highlight that the soil organic matter preservation depends on the type of forest, but the soil microbial community was able to recover after about 15 years from coppice intervention in both forest ecosystems. Thus, the type of management implemented in these forest ecosystems, not negatively affecting soil organic matter pool, preserving microbial community and potentially soil ecological functions, is sustainable in a scenario of climate change.

Coordination between degree of isohydricity and depth of root water uptake in temperate tree species.

In an increasingly dry environment, it is crucial to understand how tree species use soil water and cope with drought. However, there is still a knowledge gap regarding the relationships between species-specific stomatal behaviour, spatial root distribution, and root water uptake (RWU) dynamics. Our study aimed to investigate above- and below-ground aspects of water use during soil drying periods in four temperate tree species that differ in stomatal behaviour: two isohydric tracheid-bearing conifers, Scots pine and Norway spruce, and two more anisohydric deciduous species, the diffuse-porous European beech, and the ring-porous Downy oak. From 2015 to 2020, soil-tree-atmosphere-continuum parameters were measured for each species in monospecific forests where trees had no access to groundwater. The hourly time series included data on air temperature, vapor pressure deficit, soil water potential, soil hydraulic conductivity, and RWU to a depth of 2 m. Analysis of drought responses included data on stem radius, leaf water potential, estimated osmotically active compounds, and drought damage. Our study reveals an inherent coordination between stomatal regulation, fine root distribution and water uptake. Compared to conifers, the more anisohydric water use of oak and beech was associated with less strict stomatal closure, greater investment in deep roots, four times higher maximum RWU, a shift of RWU to deeper soil layers as the topsoil dried, and a more pronounced soil drying below 1 m depth. Soil hydraulic conductivity started to limit RWU when values fell below 10-3 to 10-5 cm/d, depending on the soil. As drought progressed, oak and beech may also have benefited from their leaf osmoregulatory capacity, but at the cost of xylem embolism with around 50 % loss of hydraulic conductivity when soil water potential dropped below -1.25 MPa. Consideration of species-specific water use is crucial for forest management and vegetation modelling to improve forest resilience to drought.

A warmer climate impairs the growth performance of Central Europe's major timber species in lowland regions.

Recent hot droughts have caused tree vitality decline and increased mortality in many forest regions on earth. Most of Central Europe's important timber species have suffered from the extreme 2018/2019 hot drought, confronting foresters with difficult questions about the choice of more drought- and heat-resistant tree species. We compared the growth dynamics of European beech, sessile oak, Scots pine and Douglas fir in a warmer and a cooler lowland region of Germany to explore the adaptive potential of the four species to climate warming (24 forest stands). The basal area increment (BAI) of the two conifers has declined since about 1990-2010 in both regions, and that of beech in the warmer region, while oak showed positive BAI trends. A 2 °C difference in mean temperatures and a higher frequency of hot days (temperature maximum >30 °C) resulted in greater sensitivity to a negative climatic water balance in beech and oak, and elevated sensitivity to summer heat in Douglas fir and pine. This suggests to include hot days in climate-growth analyses. Negative pointer years were closely related to dry years. Nevertheless, all species showed growth recovery within one to three years. We conclude that all four species are sensitive to a deteriorating climatic water balance and hot temperatures, and have so far not been able to successfully acclimate to the warmer climate, with especially Douglas and beech, but also Scots pine, being vulnerable to a warming and drying climate.

Mast seeding in European beech (Fagus sylvatica L.) is associated with reduced fungal sporocarp production and community diversity.

Mast seeding is a well-documented phenomenon across diverse forest ecosystems. While its effect on aboveground food webs has been thoroughly studied, how it impacts the soil fungi that drive soil carbon and nutrient cycling has not yet been explored. To evaluate the relationship between mast seeding and fungal resource availability, we paired a Swiss 29-year fungal sporocarp census with contemporaneous seed production for European beech (Fagus sylvatica L.). On average, mast seeding was associated with a 55% reduction in sporocarp production and a compositional community shift towards drought-tolerant taxa across both ectomycorrhizal and saprotrophic guilds. Among ectomycorrhizal fungi, traits associated with carbon cost did not explain species' sensitivity to seed production. Together, our results support a novel hypothesis that mast seeding limits annual resource availability and reproductive investment in soil fungi, creating an ecosystem 'rhythm' to forest processes that is synchronized above- and belowground.

Quantitative analysis of commercial coating penetration into Fagus crenata wood using X-ray microtomography.

Recent advances in wood treatment include the use of eco-friendly coatings to improve the wood's dimensional stability and appearance. Assessing coating performance during its service life is critical for establishing a knowledge base for product optimization. Numerous approaches, including microimaging, are available for analyzing coating behavior. In addition to conventional microscopic techniques, high-resolution X-ray microtomography is a tool that provides nondestructive imaging of coatings and their substrates. In this study, we performed two-dimensional (2D) and three-dimensional (3D) visualization of tomographic reconstruction images of two coating types, spray and brush, to observe and assess the distribution of several commercial Japanese coating materials in Fagus crenata. X-ray images and plot profiles were used to determine the penetration depths and thicknesses of coatings. Each coated sample was scanned using X-ray microtomography, which allowed successful visualization and quantification of the coating penetration depth. Chemical content and concentration of the coating materials influenced penetration depth and amount.

Rolling down that mountain: microgeographical adaptive divergence during a fast population expansion along a steep environmental gradient in European beech.

Forest tree populations harbour high genetic diversity thanks to large effective population sizes and strong gene flow, allowing them to diversify through adaptation to local environmental pressures within dispersal distance. Many tree populations also experienced historical demographic fluctuations, including spatial population contraction or expansions at various temporal scales, which may constrain their ability to adapt to environmental variations. Our aim is to investigate how recent contraction and expansion events interfere with local adaptation, by studying patterns of adaptive divergence between closely related stands undergoing environmentally contrasted conditions, and having or not recently expanded. To investigate genome-wide signatures of local adaptation while accounting for demography, we analysed divergence in a European beech population by testing pairwise differentiation among four tree stands at ~35k Single Nucleotide Polymorphisms from ~9k genomic regions. We applied three divergence outlier search methods resting on different assumptions and targeting either single SNPs or contiguous genomic regions, while accounting for the effect of population size variations on genetic divergence. We found 27 signals of selective signatures in 19 target regions. Putatively adaptive divergence involved all stand pairs. We retrieved signals both when comparing old-growth stands and recently colonised areas and when comparing stands within the old-growth area. Therefore, adaptive divergence processes have taken place both over short time spans, under strong environmental contrasts, and over short ecological gradients, in populations that have been stable in the long term. This suggests that standing genetic variation supports local, microgeographic divergence processes, which can maintain genetic diversity at the landscape level.

A genome-wide genetic association study reveals SNPs significantly associated with environmental variables and specific leaf area in European beech.

European beech is negatively affected by climate change and a further growth decline is predicted for large parts of its distribution range. Despite the importance of this species, little is known about its genetic adaptation and especially the genetic basis of its physiological traits. Here, we used genotyping by sequencing to identify SNPs in 43 German European beech populations growing under different environmental conditions. In total, 28 of these populations were located along a precipitation and temperature gradient in northern Germany, and single tree-based hydraulic and morphological traits were available. We obtained a set of 13,493 high-quality SNPs that were used for environmental and SNP-trait association analysis. In total, 22 SNPs were identified that were significantly associated with environmental variables or specific leaf area (SLA). Several SNPs were located in genes related to stress response. The majority of the significant SNPs were located in non-coding (intergenic and intronic) regions. These may be in linkage disequilibrium with the causative coding or regulatory regions. Our study gives insights into the genetic basis of abiotic adaptation in European beech, and provides genetic resources that can be used in future studies on this species. Besides clear patterns of local adaptation to environmental conditions of the investigated populations, the analyzed morphological and hydraulic traits explained most of the explainable genetic variation. Thus, they could successfully be altered in tree breeding programs, which may help to increase the adaptation of European beech to changing environmental conditions in the future.

Widespread breakdown in masting in European beech due to rising summer temperatures.

Climate change effects on tree reproduction are poorly understood, even though the resilience of populations relies on sufficient regeneration to balance increasing rates of mortality. Forest-forming tree species often mast, i.e. reproduce through synchronised year-to-year variation in seed production, which improves pollination and reduces seed predation. Recent observations in European beech show, however, that current climate change can dampen interannual variation and synchrony of seed production and that this masting breakdown drastically reduces the viability of seed crops. Importantly, it is unclear under which conditions masting breakdown occurs and how widespread breakdown is in this pan-European species. Here, we analysed 50 long-term datasets of population-level seed production, sampled across the distribution of European beech, and identified increasing summer temperatures as the general driver of masting breakdown. Specifically, increases in site-specific mean maximum temperatures during June and July were observed across most of the species range, while the interannual variability of population-level seed production (CVp) decreased. The declines in CVp were greatest, where temperatures increased most rapidly. Additionally, the occurrence of crop failures and low seed years has decreased during the last four decades, signalling altered starvation effects of masting on seed predators. Notably, CVp did not vary among sites according to site mean summer temperature. Instead, masting breakdown occurs in response to warming local temperatures (i.e. increasing relative temperatures), such that the risk is not restricted to populations growing in warm average conditions. As lowered CVp can reduce viable seed production despite the overall increase in seed count, our results warn that a covert mechanism is underway that may hinder the regeneration potential of European beech under climate change, with great potential to alter forest functioning and community dynamics.

Does long-term drought or repeated defoliation affect seasonal leaf N cycling in young beech trees?

Forest trees adopt effective strategies to optimize nitrogen (N) use through internal N recycling. In the context of more recurrent environmental stresses due to climate change, the question remains of whether increased frequency of drought or defoliation threatens this internal N recycling strategy. We submitted 8-year-old beech trees to 2 years of either severe drought (Dro) or manual defoliation (Def) to create a state of N starvation. At the end of the second year before leaf senescence, we labeled the foliage of the Dro and Def trees, as well as that of control (Co) trees, with 15N-urea. Leaf N resorption, winter tree N storage (total N, 15N, amino acids, soluble proteins) and N remobilization in spring were evaluated for the three treatments. Defoliation and drought did not significantly impact foliar N resorption or N concentrations in organs in winter. Total N amounts in Def tree remained close to those in Co tree, but winter N was stored more in the branches than in the trunk and roots. Total N amount in Dro trees was drastically reduced (-55%), especially at the trunk level, but soluble protein concentrations increased in the trunk and fine roots compared with Co trees. During spring, 15N was mobilized from the trunk, branches and twigs of both Co and Def trees to support leaf growth. It was only provided through twig 15N remobilization in the Dro trees, thus resulting in extremely reduced Dro leaf N amounts. Our results suggest that stress-induced changes occur in N metabolism but with varying severity depending on the constraints: within-tree 15N transport and storage strategy changed in response to defoliation, whereas a soil water deficit induced a drastic reduction of the N amounts in all the tree organs. Consequently, N dysfunction could be involved in drought-induced beech tree mortality under the future climate.