Leaf gene expression trajectories during the growing season are consistent between sites and years in American beech.

Transcriptomics provides a versatile tool for ecological monitoring. Here, through genome-guided profiling of transcripts mapping to 33 042 gene models, expression differences can be discerned among multi-year and seasonal leaf samples collected from American beech trees at two latitudinally separated sites. Despite a bottleneck due to post-Columbian deforestation, the single nucleotide polymorphism-based population genetic background analysis has yielded sufficient variation to account for differences between populations and among individuals. Our expression analyses during spring-summer and summer-autumn transitions for two consecutive years involved 4197 differentially expressed protein coding genes. Using Populus orthologues we reconstructed a protein-protein interactome representing leaf physiological states of trees during the seasonal transitions. Gene set enrichment analysis revealed gene ontology terms that highlight molecular functions and biological processes possibly influenced by abiotic forcings such as recovery from drought and response to excess precipitation. Further, based on 324 co-regulated transcripts, we focused on a subset of GO terms that could be putatively attributed to late spring phenological shifts. Our conservative results indicate that extended transcriptome-based monitoring of forests can capture diverse ranges of responses including air quality, chronic disease, as well as herbivore outbreaks that require activation and/or downregulation of genes collectively tuning reaction norms maintaining the survival of long living trees such as the American beech.

Adaptive monitoring in action-what drives arthropod diversity and composition in central European beech forests?

Recent studies suggest that arthropod diversity in German forests is declining. Currently, different national programs are being developed to monitor arthropod trends and to unravel the effects of forest management on biodiversity in forests. To establish effective long-term monitoring programs, a set of drivers of arthropod diversity and composition as well as suitable species groups have to be identified. To aid in answering these questions, we investigated arthropod data collected in four Hessian forest reserves (FR) in the 1990s. To fully utilize this data set, we combined it with results from a retrospective structural sampling design applied at the original trap locations in central European beech (Fagus sylvatica) forests. As expected, the importance of the different forest structural, vegetation, and site attributes differed largely between the investigated arthropod groups: beetles, spiders, Aculeata, and true bugs. Measures related to light availability and temperature such as canopy cover or potential radiation were important to all groups affecting either richness, composition, or both. Spiders and true bugs were affected by the broadest range of explanatory variables, which makes them a good choice for monitoring general trends. For targeted monitoring focused on forestry-related effects on biodiversity, rove and ground beetles seem more suitable. Both groups were driven by a narrower, more management-related set of variables. Most importantly, our study approach shows that it is possible to utilize older biodiversity survey data. Although, in our case, there are strong restrictions due to the long time between species and structural attribute sampling.

Contrasting patterns of water use efficiency and annual radial growth among European beech forests along the Italian peninsula.

Tree mortality and forest dieback episodes are increasing due to drought and heat stress. Nevertheless, a comprehensive understanding of mechanisms enabling trees to withstand and survive droughts remains lacking. Our study investigated basal area increment (BAI), and δ13C-derived intrinsic water-use-efficiency (iWUE), to elucidate beech resilience across four healthy stands in Italy with varying climates and soil water availability. Additionally, fist-order autocorrelation (AR1) analysis was performed to detect early warning signals for potential tree dieback risks during extreme drought events. Results reveal a negative link between BAI and vapour pressure deficit (VPD), especially in southern latitudes. After the 2003 drought, BAI decreased at the northern site, with an increase in δ13C and iWUE, indicating conservative water-use. Conversely, the southern sites showed increased BAI and iWUE, likely influenced by rising CO2 and improved water availability. In contrast, the central site sustained higher transpiration rates due to higher soil water holding capacity (SWHC). Despite varied responses, most sites exhibited reduced resilience to future extreme events, indicated by increased AR1. Temperature significantly affected beech iWUE and BAI in northern Italy, while VPD strongly influenced the southern latitudes. The observed increase in BAI and iWUE in southern regions might be attributed to an acclimation response.

Beech Mushroom (Hypsizygus marmoreus, Agaricomycetes) Cultivation and Outstanding Health-Promoting Properties: A Review.

Currently, in many Asian and European countries, a group of mushrooms has been distinguished, which, due to their medicinal and/or dietary properties, are referred to as medicinal mushrooms. Hypsizygus marmoreus is a species valued for its proven nutritional and healing properties. In the scientific literature available so far, mainly from Asian countries, it is possible to find research on the therapeutic effect of H. marmoreus. Biologically active substances, their presence, and in particular, their level in this species have not been the subject of extensive research. The aim of the present review is to describe and summarize the dietary and therapeutic potential of this increasingly popular species among consumers in European countries. Because H. marmoreus is a species with known dietary and health-promoting properties, this species of wood fungi could be commonly obtained from cultivation in European countries.

Summer solstice orchestrates the subcontinental-scale synchrony of mast seeding.

High interannual variation in seed production in perennial plants can be synchronized at subcontinental scales with wide consequences for ecosystem functioning, but how such synchrony is generated is unclear1-3. We investigated the factors contributing to masting synchrony in European beech (Fagus sylvatica), which extends to a geographic range of 2,000 km. Maximizing masting synchrony via spatial weather coordination, known as the Moran effect, requires a simultaneous response to weather conditions across distant populations. A celestial cue that occurs simultaneously across the entire hemisphere is the longest day (the summer solstice). We show that European beech abruptly opens its temperature-sensing window on the solstice, and hence widely separated populations all start responding to weather signals in the same week. This celestial 'starting gun' generates ecological events with high spatial synchrony across the continent.

A framework for tracing timber following the Ukraine invasion.

Scientific testing including stable isotope ratio analysis (SIRA) and trace element analysis (TEA) is critical for establishing plant origin, tackling deforestation and enforcing economic sanctions. Yet methods combining SIRA and TEA into robust models for origin verification and determination are lacking. Here we report a (1) large Eastern European timber reference database (Betula, Fagus, Pinus, Quercus) tailored to sanctioned products following the Ukraine invasion; (2) statistical test to verify samples against a claimed origin; (3) probabilistic model of SIRA, TEA and genus distribution data, using Gaussian processes, to determine timber harvest location. Our verification method rejects 40-60% of simulated false claims, depending on the spatial scale of the claim, and maintains a low probability of rejecting correct origin claims. Our determination method predicts harvest location within 180 to 230 km of true location. Our results showcase the power of combining data types with probabilistic modelling to identify and scrutinize timber harvest location claims.

Drought alters aboveground biomass production efficiency: Insights from two European beech forests.

The fraction of photosynthetically assimilated carbon that trees allocate to long-lasting woody biomass pools (biomass production efficiency - BPE), is a key metric of the forest carbon balance. Its apparent simplicity belies the complex interplay between underlying processes of photosynthesis, respiration, litter and fruit production, and tree growth that respond differently to climate variability. Whereas the magnitude of BPE has been routinely quantified in ecological studies, its temporal dynamics and responses to extreme events such as drought remain less well understood. Here, we combine long-term records of aboveground carbon increment (ACI) obtained from tree rings with stand-level gross primary productivity (GPP) from eddy covariance (EC) records to empirically quantify aboveground BPE (= ACI/GPP) and its interannual variability in two European beech forests (Hainich, DE-Hai, Germany; Sorø, DK-Sor, Denmark). We found significant negative correlations between BPE and a daily-resolved drought index at both sites, indicating that woody growth is de-prioritized under water limitation. During identified extreme years, early-season drought reduced same-year BPE by 29 % (Hainich, 2011), 31 % (Sorø, 2006), and 14 % (Sorø, 2013). By contrast, the 2003 late-summer drought resulted in a 17 % reduction of post-drought year BPE at Hainich. Across the entire EC period, the daily-to-seasonal drought response of BPE resembled that of ACI, rather than that of GPP. This indicates that BPE follows sink dynamics more closely than source dynamics, which appear to be decoupled given the distinctive climate response patterns of GPP and ACI. Based on our observations, we caution against estimating the magnitude and variability of the carbon sink in European beech (and likely other temperate forests) based on carbon fluxes alone. We also encourage comparable studies at other long-term EC measurement sites from different ecosystems to further constrain the BPE response to rare climatic events.

Bonding performance and surface characterization of cold-bonded acetylated beech (Fagus sylvatica L.) laminated veneer lumber.

Acetylation of wood with acetic anhydride reduces the wood-moisture interaction, improves the dimensional stability and resistance against biodegradation. However, the adhesive bonding is affected by the modification, which is crucial to manufacture engineered wood products, such as laminated veneer lumber (LVL). In this study we report the bonding of 8-layered acetylated beech (Fagus sylvatica L.) LVL boards to 2-layered LVL beams. The beams were glued together at room temperature adding three common load-bearing construction adhesives: melamine-urea-formaldehyde (MUF), phenol-resorcinol-formaldehyde (PRF), and one-component polyurethane (PUR). The bonding performance was tested by assessing its dry and wet tensile shear strength (TSS) and wood failure percentage (WF). Also evaluated were the material's density and moisture content (MC). The surface was characterized prior to bonding by its pH, roughness, and contact angle (CA). The adhesive penetration was observed by fluorescence microscopy. Aside from MUF, applying PRF and PUR adhesives achieved good bonding performance on acetylated LVL and references. Acetylated LVL displayed a more hydrophobic behaviour, a higher pH, a somewhat smoother surface, and an increased density.

Temporal and spatial variability of phloem structure in Picea abies and Fagus sylvatica and its link to climate.

Using a unique 8-year data set (2010-2017) of phloem data, we studied the effect of temperature and precipitation on the phloem anatomy (conduit area, widths of ring, early and late phloem) and xylem-ring width in two coexisting temperate tree species, Picea abies and Fagus sylvatica, from three contrasting European temperate forest sites. Histometric analyses were performed on microcores taken from tree stems in autumn. We found high interannual variability and sensitivity of phloem anatomy and xylem-ring widths to precipitation and temperature; however, the responses were species- and site-specific. The contrasting response of xylem and phloem-ring widths of the same tree species to weather conditions was found at the two Slovenian sites generally well supplied with precipitation, while at the driest Czech site, the influence of weather factors on xylem and phloem ring widths was synchronised. Since widths of mean annual xylem and phloem increments were narrowest at the Czech site, this site is suggested to be most restrictive for the radial growth of both species. By influencing the seasonal patterns of xylem and phloem development, water availability appears to be the most important determinant of tissue- and species-specific responses to local weather conditions.

The impact of root systems and their exudates in different tree species on soil properties and microorganisms in a temperate forest ecosystem.

BACKGROUND: The species composition of tree stands plays an important role in shaping the properties of forest soils. The aim of our research was to determine the influence on soil properties of the root systems of six species of trees which form forest stands in the temperate climatic zone. The research covered areas including six tree species - Scots pine (Pinus sylvestris L.), European larch (Larix deciduas Mill.), English oak (Quercus robur L.), English ash (Fraxinus excelsior L.), European beech (Fagus sylvatica L.) and European hornbeam (Carpinus betulus L.). In our study, we determined the characteristics of the roots and the amount of carbon excreted alongside their exudates. Enzymatic activity, and the composition and diversity of the fungi and bacteria, were also determined in addition to the basic physicochemical properties of the soil samples. RESULTS: A strong relationship between the root characteristics and soil properties, including the pH, basic cation content and phosphorus content, was confirmed. In addition, the enzymatic activity of phosphatase, ß-glucosidase, N-acetyl-ß-D-glucosaminidase and ß-D-cellobiosidase were positively correlated with the root characteristics. The study on soil bacteria across different tree species revealed Proteobacteria and Actinobacteriota to be the most abundant phylum. Fungal analysis showed Basidiomycota and Ascomycota as the dominant phyla. Ascomycota dominated in hornbeam and oak soils. Mortierellomycota was remarkably more present in pine soil. CONCLUSIONS: This analysis of root systems and soil properties confirmed the distinctness of ash stands, which were also more abundant in various microorganisms. It was also found that soils affected by different tree species were characterised by varied fungal and bacterial composition. The ash had particularly beneficial impact on soil microbiota.

Major tree species of Central European forests differ in their proportion of positive, negative, and nonstationary growth trends.

Temperate forests are undergoing significant transformations due to the influence of climate change, including varying responses of different tree species to increasing temperature and drought severity. To comprehensively understand the full range of growth responses, representative datasets spanning extensive site and climatic gradients are essential. This study utilizes tree-ring data from 550 sites from the temperate forests of Czechia to assess growth trends of six dominant Central European tree species (European beech, Norway spruce, Scots pine, silver fir, sessile and pedunculate oak) over 1990-2014. By modeling mean growth series for each species and site, and employing principal component analysis, we identified the predominant growth trends. Over the study period, linear growth trends were evident across most sites (56% increasing, 32% decreasing, and 10% neutral). The proportion of sites with stationary positive trends increased from low toward high elevations, whereas the opposite was true for the stationary negative trends. Notably, within the middle range of their distribution (between 500 and 700 m a.s.l.), Norway spruce and European beech exhibited a mix of positive and negative growth trends. While Scots pine growth trends showed no clear elevation-based pattern, silver fir and oaks displayed consistent positive growth trends regardless of site elevation, indicating resilience to the ongoing warming. We demonstrate divergent growth trajectories across space and among species. These findings are particularly important as recent warming has triggered a gradual shift in the elevation range of optimal growth conditions for most tree species and has also led to a decoupling of growth trends between lowlands and mountain areas. As a result, further future shifts in the elevation range and changes in species diversity of European temperate forests can be expected.

Interactions between beech and oak seedlings can modify the effects of hotter droughts and the onset of hydraulic failure.

Mixing species with contrasting resource use strategies could reduce forest vulnerability to extreme events. Yet, how species diversity affects seedling hydraulic responses to heat and drought, including mortality risk, is largely unknown. Using open-top chambers, we assessed how, over several years, species interactions (monocultures vs mixtures) modulate heat and drought impacts on the hydraulic traits of juvenile European beech and pubescent oak. Using modeling, we estimated species interaction effects on timing to drought-induced mortality and the underlying mechanisms driving these impacts. We show that mixtures mitigate adverse heat and drought impacts for oak (less negative leaf water potential, higher stomatal conductance, and delayed stomatal closure) but enhance them for beech (lower water potential and stomatal conductance, narrower leaf safety margins, faster tree mortality). Potential underlying mechanisms include oak's larger canopy and higher transpiration, allowing for quicker exhaustion of soil water in mixtures. Our findings highlight that diversity has the potential to alter the effects of extreme events, which would ensure that some species persist even if others remain sensitive. Among the many processes driving diversity effects, differences in canopy size and transpiration associated with the stomatal regulation strategy seem the primary mechanisms driving mortality vulnerability in mixed seedling plantations.

Granivore abundance shapes mutualism quality in plant-scatterhoarder interactions.

Conditional mutualisms involve costs and benefits that vary with environmental factors, but mechanisms driving these dynamics remain poorly understood. Scatterhoarder-plant interactions are a prime example of this phenomenon, as scatterhoarders can either increase or reduce plant recruitment depending on the balance between seed dispersal and predation. We explored factors that drive the magnitude of net benefits for plants in this interaction using a mathematical model, with parameter values based on European beech (Fagus sylvatica) and yellow-necked mice (Apodemus flavicollis). We measured benefits as the percentage of germinating seeds, and examined how varying rodent survival (reflecting, e.g. changes in predation pressure), the rate of seed loss to other granivores, the abundance of alternative food resources, and changes in masting patterns affect the quality of mutualism. We found that increasing granivore abundance can degrade the quality of plant-scatterhoarder mutualism due to increased cache pilferage. Scatterhoarders are predicted to respond by increasing immediate consumption of gathered seeds, leading to higher costs and reduced benefits for plants. Thus, biotic changes that are detrimental to rodent populations can be beneficial for tree recruitment due to adaptive behavior of rodents. When scatterhoarder populations decline too drastically (< 5 individuals ha-1 ); however, tree recruitment may also suffer.

Slower growth prior to the 2018 drought and a high growth sensitivity to previous year summer conditions predisposed European beech to crown dieback.

The record-breaking drought in 2018 caused premature leaf discoloration and shedding (early browning) in many beech (Fagus sylvatica L.) dominated forests in Central Europe. However, a high degree of variability in drought response among individual beech trees was observed. While some trees were severely impacted by the prolonged water deficits and high temperatures, others remained vital with no or only minor signs of crown vitality loss. Why some beech trees were more susceptible to drought-induced crown damage than others and whether growth recovery is possible are poorly understood. Here, we aimed to identify growth characteristics associated with the variability in drought response between individual beech trees based on a sample of 470 trees in northern Switzerland. By combining tree growth measurements and crown condition assessments, we also investigated the possible link between crown dieback and growth recovery after drought. Beech trees with early browning exhibited an overall lower growth vigor before the 2018 drought than co-occurring vital beech trees. This lower vigor is mainly indicated by lower overall growth rates, stronger growth declines in the past decades, and higher growth-climate sensitivity. Particularly, warm previous year summer conditions negatively affected current growth of the early-browning trees. These findings suggest that the affected trees had less access to critical resources and were physiologically limited in their growth predisposing them to early browning. Following the 2018 drought, observed growth recovery potential corresponded to the amount of crown dieback and the local climatic water balance. Overall, our findings emphasize that beech-dominated forests in Central Europe are under increasing pressure from severe droughts, ultimately reducing the competitive ability of this species, especially on lowland sites with shallow soils and low water holding capacity.

Chronic warming and dry soils limit carbon uptake and growth despite a longer growing season in beech and oak.

Progressively warmer and drier climatic conditions impact tree phenology and carbon cycling with large consequences for forest carbon balance. However, it remains unclear how individual impacts of warming and drier soils differ from their combined effects and how species interactions modulate tree responses. Using mesocosms, we assessed the multiyear impact of continuous air warming and lower soil moisture alone or in combination on phenology, leaf-level photosynthesis, nonstructural carbohydrate concentrations, and aboveground growth of young European beech (Fagus sylvatica L.) and Downy oak (Quercus pubescens Willd.) trees. We further tested how species interactions (in monocultures and in mixtures) modulated these effects. Warming prolonged the growing season of both species but reduced growth in oak. In contrast, lower moisture did not impact phenology but reduced carbon assimilation and growth in both species. Combined impacts of warming and drier soils did not differ from their single effects. Under warmer and drier conditions, performances of both species were enhanced in mixtures compared to monocultures. Our work revealed that higher temperature and lower soil moisture have contrasting impacts on phenology vs. leaf-level assimilation and growth, with the former being driven by temperature and the latter by moisture. Furthermore, we showed a compensation in the negative impacts of chronic heat and drought by tree species interactions.

Interaction between beech and spruce trees in temperate forests affects water use, root water uptake pattern and canopy structure.

Beneficial and negative effects of species interactions can strongly influence water fluxes in forest ecosystems. However, little is known about how trees dynamically adjust their water use when growing with interspecific neighbours. Therefore, we investigated the interaction effects between Fagus sylvatica (European beech) and Picea abies (Norway spruce) on water-use strategies and aboveground structural characteristics. We used continuous in situ isotope spectroscopy of xylem and soil water to investigate source water dynamics and root water uptake depths. Picea abies exhibited a reduced sun-exposed crown area in equally mixed compared with spruce-dominated sites, which was further correlated to a reduction in sap flow of -14.5 ± 8.2%. Contrarily, F. sylvatica trees showed +13.3 ± 33.3% higher water fluxes in equally mixed compared with beech-dominated forest sites. Although a significantly higher crown interference by neighbouring trees was observed, no correlation of water fluxes and crown structure was found. High time-resolved xylem δ2H values showed a large plasticity of tree water use (-74.1 to -28.5‰), reflecting the δ2H dynamics of soil and especially precipitation water sources. Fagus sylvatica in equally mixed sites shifted water uptake to deeper soil layers, while uptake of fresh precipitation was faster in beech-dominated sites. Our continuous in situ water stable isotope measurements traced root water uptake dynamics at unprecedented temporal resolution, indicating highly dynamic use of water sources in response to precipitation and to neighbouring species competition. Understanding this plasticity may be highly relevant in the context of increasing water scarcity and precipitation variability under climate change.

New Tricholomalides D-G from the Mushroom Tricholoma ustaloides Grown in an Italian Beech Wood.

Four novel seconeodolastane diterpenoids, named tricholomalides D-G, were isolated, together with the known tricholomalide C, from the fruiting bodies of Tricholoma ustaloides Romagn., a species belonging to the large Tricholoma genus of higher mushrooms (Basidiomycota, family Tricholomataceae). They were isolated through multiple chromatographic separations, and the structures, including the absolute configuration, were established through a detailed analysis of MS, NMR, and CD spectral data and comparison with related compounds reported in the literature, which has been thoroughly revised.

Development of polymorphic microsatellite markers for Fagus pashanica (Fagaceae) using next-generation sequencing.

Fagus pashanica is an endangered and endemic tree species in China. To understand its genetic diversity and structure for effective conservation, we used next-generation sequencing data to develop a set of microsatellite markers. Twenty-three of the 68 designed loci were successfully amplified. Fifteen polymorphic loci with clear peaks were selected for further analyses in three F. pashanica populations sampled from Nanjiang, Wangcang and Pingwu counties in Sichuan Province, China. The number of alleles per locus ranged from two to 11. The levels of observed and expected heterozygosity ranged from 0.033-0.852 and 0.033-0.787, respectively. All 23 loci were also successfully amplified in F. longipetiolata and F. lucida, and 19 were successfully amplified in F. engleriana. These microsatellite markers will be useful for population genetic studies of F. pashanica and other Fagus species.

Cellular insights of beech leaf disease reveal abnormal ectopic cell division of symptomatic interveinal leaf areas.

The beech leaf disease nematode, Litylenchus crenatae subsp. mccannii, is recognized as a newly emergent nematode species that causes beech leaf disease (BLD) in beech trees (Fagus spp.) in North America. Changes of leaf morphology before emergence from the bud induced by BLD can provoke dramatic effects on the leaf architecture and consequently to tree performance and development. The initial symptoms of BLD appear as dark green, interveinal banding patterns of the leaf. Despite the fast progression of this disease, the cellular mechanisms leading to the formation of such aberrant leaf phenotype remains totally unknown. To understand the cellular basis of BLD, we employed several types of microscopy to provide an exhaustive characterization of nematode-infected buds and leaves. Histological sections revealed a dramatic cell change composition of these nematode-infected tissues. Diseased bud scale cells were typically hypertrophied and showed a high variability of size. Moreover, while altered cell division had no influence on leaf organogenesis, induction of cell proliferation on young leaf primordia led to a dramatic change in cell layer architecture. Hyperplasia and hypertrophy of the different leaf cell layers, coupled with an abnormal proliferation of chloroplasts especially in the mesophyll cell layers, resulted in the typical interveinal leaf banding. These discrepancies in leaf cell structure were depicted by an abnormal rate of cellular division of the leaf interveinal areas infected by the nematode, promoting significant increase of cell size and leaf thickness. The formation of symptomatic BLD leaves is therefore orchestrated by distinct cellular processes, to enhance the value of these feeding sites and to improve their nutrition status for the nematode. Our findings thus uncover relevant cellular events and provide a structural framework to understand this important disease.