Tree drought-mortality risk depends more on intrinsic species resistance than on stand species diversity.

Increasing tree diversity is considered a key management option to adapt forests to climate change. However, the effect of species diversity on a forest's ability to cope with extreme drought remains elusive. In this study, we assessed drought tolerance (xylem vulnerability to cavitation) and water stress (water potential), and combined them into a metric of drought-mortality risk (hydraulic safety margin) during extreme 2021 or 2022 summer droughts in five European tree diversity experiments encompassing different biomes. Overall, we found that drought-mortality risk was primarily driven by species identity (56.7% of the total variability), while tree diversity had a much lower effect (8% of the total variability). This result remained valid at the local scale (i.e within experiment) and across the studied European biomes. Tree diversity effect on drought-mortality risk was mediated by changes in water stress intensity, not by changes in xylem vulnerability to cavitation. Significant diversity effects were observed in all experiments, but those effects often varied from positive to negative across mixtures for a given species. Indeed, we found that the composition of the mixtures (i.e., the identities of the species mixed), but not the species richness of the mixture per se, is a driver of tree drought-mortality risk. This calls for a better understanding of the underlying mechanisms before tree diversity can be considered an operational adaption tool to extreme drought. Forest diversification should be considered jointly with management strategies focussed on favouring drought-tolerant species.

Early overyielding in a mixed deciduous forest is driven by both above- and below ground species-specific acclimation.

BACKGROUND AND AIMS: Mixed forest plantations are increasingly recognised for their role in mitigating the impacts of climate change and enhancing ecosystem resilience. Yet, there remains a significant gap in understanding the early-stage dynamics of species trait diversity and interspecies interactions, particularly in pure deciduous mixtures. This study aims to explore the timing and mechanisms by which trait diversity of deciduous species and competitive interactions influence yield, carbon allocation, and space occupation in mixed forests, both above- and belowground. METHODS: A forest inventory was conducted in planted monocultures, 2-species, and 4-species mixtures of European Acer, Tilia, Carpinus, and Quercus, representing a spectrum from acquisitive to conservative tree species. Competition effects were assessed with linear mixed-effects models at the level of biomass and space acquisition, including leaf, canopy, stem, and fine root traits. KEY RESULTS: Early aboveground growth effects were observed six years post-planting, with significant biomass accumulation after eight years, strongly influenced by species composition. Mixtures, especially with acquisitive species, exhibited aboveground overyielding, 1.5- to 1.9-times higher than monocultures. Fine roots showed substantial overyielding in high diversity stands. Biomass allocation was species-specific and varied markedly by tree size, the level of diversity, and between acquisitive Acer and the more conservative species. No root segregation was found. CONCLUSIONS: Our findings underscore the critical role of species trait diversity in enhancing productivity in mixed deciduous forest plantations. Allometric changes highlight the need to differentiate between (active) acclimations and (passive) tree size-related changes, but illustrate major consequences of competitive interactions for the functional relation between leaves, stem, and roots. This study points towards the significant contributions of both above- and belowground components to overall productivity of planted mixed-species forests.

Soil fertility determines whether ectomycorrhizal fungi accelerate or decelerate decomposition in a temperate forest.

Ectomycorrhizal (ECM) fungi can both accelerate and decelerate decomposition of organic matter in forest soils, but a mechanistic understanding of this differential influence is limited. Here, we tested how ECM fungi affect decomposition along a natural fertility gradient in a temperate forest of European beech. Trees were girdled to reduce belowground carbon supply to the soil. Girdling shifted soil fungal community composition and decreased hyphal biomass production and soil CO2 efflux, indicating a reduced ECM fungal activity. Girdling also affected decomposition processes, but the effects depended on fertility. Our results indicate that ECM fungi decelerate decomposition under conditions of low fertility while under conditions of high fertility ECM fungi and their host roots have an accelerating effect. We conclude that both acceleration and deceleration of decomposition of organic matter by ECM fungi can occur within a forest, with soil fertility determining the direction and magnitude of these effects. We suggest a positive feedback between fertility, stand productivity and soil carbon and nitrogen dynamics that is mediated to a large extent by ECM fungi.

Soil fertility relates to fungal-mediated decomposition and organic matter turnover in a temperate mountain forest.

Fungi are known to exert a significant influence over soil organic matter (SOM) turnover, however understanding of the effects of fungal community structure on SOM dynamics and its consequences for ecosystem fertility is fragmentary. Here we studied soil fungal guilds and SOM decomposition processes along a fertility gradient in a temperate mountain beech forest. High-throughput sequencing was used to investigate fungal communities. Carbon and nitrogen stocks, enzymatic activity and microbial respiration were measured. While ectomycorrhizal fungal abundance was not related to fertility, saprotrophic ascomycetes showed higher relative abundances under more fertile conditions. The activity of oxidising enzymes and respiration rates in mineral soil were related positively to fertility and saprotrophic fungi. In addition, organic layer carbon and nitrogen stocks were lower on the more fertile plots, although tree biomass and litter input were higher. Together, the results indicated a faster SOM turnover at the fertile end of the gradient. We suggest that there is a positive feedback mechanism between SOM turnover and fertility that is mediated by soil fungi to a significant extent. By underlining the importance of fungi for soil fertility and plant growth, these findings furthermore emphasise the dependency of carbon cycling on fungal communities below ground.

High Fungal Diversity but Low Seasonal Dynamics and Ectomycorrhizal Abundance in a Mountain Beech Forest.

Forests on steep slopes constitute a significant proportion of European mountain areas and are important as production and protection forests. This study describes the soil fungal community structure in a European beech-dominated mountain forest stands in the Northern Calcareous Alps and investigates how it is determined by season and soil properties. Samples were collected at high spatial resolution in an area of ca. 100 m × 700 m in May (spring) and August (summer). Illumina MiSeq high-throughput sequencing of the ITS2-region revealed distinct patterns for the soil fungal communities. In contrast to other studies from temperate European beech forest stands, Ascomycota dominated the highly diverse fungal community, while ectomycorrhizal fungi were of lower abundance. Russulaceae, which are often among the dominant ectomycorrhizal fungi associated with European beech, were absent from all samples. Potentially plant pathogenic fungi were more prevalent than previously reported. Only subtle seasonal differences were found between fungal communities in spring and summer. Especially, dominant saprotrophic taxa were largely unaffected by season, while slightly stronger effects were observed for ectomycorrhizal fungi. Soil characteristics like pH and organic carbon content, on the other hand, strongly shaped abundant taxa among the saprotrophic fungal community.

Non-structural carbohydrate concentrations of Fagus sylvatica and Pinus sylvestris fine roots are linked to ectomycorrhizal enzymatic activity during spring reactivation.

We evaluated whether changes in fine root non-structural carbohydrate reserves of Fagus sylvatica and Pinus sylvestris trees influence potential enzymatic activities of their ectomycorrhizal symbionts from winter towards spring reactivation, and whether these changes influence potential soil enzymatic activities. We analyzed sugar and starch concentrations in the fine roots of Fagus sylvatica and Pinus sylvestris and potential activities of ß-glucosidase, ß-xylosidase, and cellobiohydrolase (as proxies for carbon-degrading enzymes) as well as leucine aminopeptidase and chitinase (as proxies for nitrogen-degrading enzymes) of their dominant ectomycorrhizal symbionts as well as in the soil. Sugar concentrations in the fine roots were significantly positively correlated with enzymatic activities of the ectomycorrhizal symbionts. In Pinus sylvestris, both carbon- and nitrogen-degrading enzyme activities showed significant positive correlations with fine root sugar concentrations. In Fagus sylvatica, fine root sugar concentrations were explicitly positively correlated with the activity of nitrogen-degrading enzymes. The chitinase activity in the soil was found to be strongly positively correlated with the enzymatic activity of the ectomycorrhizal symbionts as well as with fine root sugar concentrations. Fine root carbohydrate concentrations of Fagus sylvatica and Pinus sylvestris trees and enzymatic activities of their associated ectomycorrhizal fungi are connected. The specific nutrient demand of the tree species during spring reactivation may affect ectomycorrhizal enzymatic activity via carbon mobilization in the fine roots of Fagus sylvatica and Pinus sylvestris. Moreover, our results suggest that trees indirectly contribute to the degradation of fungal necromass by stimulating ectomycorrhizal chitinase activity in the soil.

Transition of Ethiopian highland forests to agriculture-dominated landscapes shifts the soil microbial community composition.

BACKGROUND: Land use changes and related land management practices significantly alter soil physicochemical properties; however, their effects on the soil microbial community structure are still unclear. In this study, we used automated ribosomal intergenic spacer analysis to determine the fungal and bacterial community composition in soils from different land use areas in the Ethiopian highlands. Soil samples were collected from five areas with different land uses, natural forest, eucalyptus plantation, exclosure, grassland and cropland, which had all historically been natural forest. RESULTS: Our results showed a significant shift in the soil bacterial and fungal community composition in response to land use change. We also identified soil physicochemical factors corresponding to the changes in bacterial and fungal communities. Although most soil attributes, including soil organic carbon, total soil nitrogen, labile P, soil pH and soil aggregate stability, were related to the change in bacterial community composition, the total soil nitrogen and soil organic carbon had the strongest relationships. The change in fungal community composition was correlated with soil nutrients, organic carbon, soil nitrogen and particularly the labile P concentration. CONCLUSIONS: The fungal community composition was likely affected by the alteration of vegetation cover in response to land use change, whereas the bacterial communities were mainly sensitive to changes in soil attributes. The study highlights the higher sensitivity of fungal communities than bacterial communities to land use changes.

Adaptive root foraging strategies along a boreal-temperate forest gradient.

The tree root-mycorhizosphere plays a key role in resource uptake, but also in the adaptation of forests to changing environments. The adaptive foraging mechanisms of ectomycorrhizal (EcM) and fine roots of Picea abies, Pinus sylvestris and Betula pendula were evaluated along a gradient from temperate to subarctic boreal forest (38 sites between latitudes 48°N and 69°N) in Europe. Variables describing tree resource uptake structures and processes (absorptive fine root biomass and morphology, nitrogen (N) concentration in absorptive roots, extramatrical mycelium (EMM) biomass, community structure of root-associated EcM fungi, soil and rhizosphere bacteria) were used to analyse relationships between root system functional traits and climate, soil and stand characteristics. Absorptive fine root biomass per stand basal area increased significantly from temperate to boreal forests, coinciding with longer and thinner root tips with higher tissue density, smaller EMM biomass per root length and a shift in soil microbial community structure. The soil carbon (C) : N ratio was found to explain most of the variability in absorptive fine root and EMM biomass, root tissue density, N concentration and rhizosphere bacterial community structure. We suggest a concept of absorptive fine root foraging strategies involving both qualitative and quantitative changes in the root-mycorrhiza-bacteria continuum along climate and soil C : N gradients.

It's complicated: intraroot system variability of respiration and morphological traits in four deciduous tree species.

Within branched root systems, a distinct heterogeneity of traits exists. Knowledge about the ecophysiology of different root types is critical to understand root system functioning. Classification schemes have to match functional root types as closely as possible to be used for sampling and modeling. Among ecophysiological root traits, respiration is of particular importance, consuming a great amount of carbon allocated. Root architecture differs between the four deciduous tree seedlings. However, two types of terminal root segments (i.e. first and second orders), white colored and brown colored, can be distinguished in all four species but vary in frequency, their morphology differing widely from each other and higher coarse root orders. Root respiration is related to diameter and tissue density. The use of extended root ordering (i.e. order and color) explains the variance of respiration two times as well as root diameter or root order classes alone. White terminal roots respire significantly more than brown ones; both possess respiration rates that are greater than those of higher orders in regard to dry weight and lower in regard to surface area. The correlation of root tissue density to respiration will allow us to use this continuous parameter (or easier to determine dry matter content) to model the respiration within woody root systems without having to determine nitrogen contents. In addition, this study evidenced that extended root orders are better suited than root diameter classes to picture the differences between root functional types. Together with information on root order class frequencies, these data allow us to calculate realistic, species-specific respiration rates of root branches.

Elevated CO2 enrichment induces a differential biomass response in a mixed species temperate forest plantation.

In a free-air carbon dioxide (CO(2)) enrichment study (BangorFACE), Alnus glutinosa, Betula pendula and Fagus sylvatica were planted in areas of one-, two- and three-species mixtures (n = 4). The trees were exposed to ambient or elevated CO(2) (580 µmol mol(-1)) for 4 yr, and aboveground growth characteristics were measured. In monoculture, the mean effect of CO(2) enrichment on aboveground woody biomass was + 29, + 22 and + 16% for A. glutinosa, F. sylvatica and B. pendula, respectively. When the same species were grown in polyculture, the response to CO(2) switched to + 10, + 7 and 0% for A. glutinosa, B. pendula and F. sylvatica, respectively. In ambient atmosphere, our species grown in polyculture increased aboveground woody biomass from 12.9 ± 1.4 to 18.9 ± 1.0 kg m(-2), whereas, in an elevated CO(2) atmosphere, aboveground woody biomass increased from 15.2 ± 0.6 to 20.2 ± 0.6 kg m(-2). The overyielding effect of polyculture was smaller (+ 7%) in elevated CO(2) than in an ambient atmosphere (+ 18%). Our results show that the aboveground response to elevated CO(2) is affected significantly by intra- and interspecific competition, and that the elevated CO(2) response may be reduced in forest communities comprising tree species with contrasting functional traits.

Particulate matter accumulation by tree foliage is driven by leaf habit types, urbanization- and pollution levels.

Particulate matter (PM) pollution poses a significant threat to human health. Greenery, particularly trees, can act as effective filters for PM, reducing associated health risks. Previous studies have indicated that tree traits play a crucial role in determining the amount of PM accumulated on leaves, although findings have often been site-specific. To comprehensively investigate the key factors influencing PM binding to leaves across diverse tree species and geographical locations, we conducted an extensive analysis using data extracted from 57 publications. The data covers 11 countries and 190 tree species from 1996 to 2021. We categorized tree species into functional groups: evergreen conifers, deciduous conifers, deciduous broadleaves, and evergreen broadleaves based on leaf habit and phylogeny. Evergreen conifers exhibited the highest PM accumulation on leaves, and in general, evergreen leaves accumulated more PM compared to deciduous leaves across all PM size classes. Specific leaf traits, such as epicuticular wax, played a significant role. The highest PM loads on leaves were observed in peri-urban areas along the rural-peri-urban-urban gradient. However, the availability of global data was skewed, with most data originating from urban and peri-urban areas, primarily from China and Poland. Among different climate zones, substantial data were only available for warm temperate and cold steppe climate zones. Understanding the problem of PM pollution and the role of greenery in urban environments is crucial for monitoring and controlling PM pollution. Our systematic review of the literature highlights the variation on PM loading among different vegetation types with varying leaf characteristics. Notably, epicuticular wax emerged as a marker trait that exhibited variability across PM size fractions and different vegetation types. In conclusion, this review emphasizes the importance of greenery in mitigation PM pollution. Our findings underscore the significance of tree traits in PM binding. However, lack of data stresses the need for further research and data collection initiatives.

Improving models of fine root carbon stocks and fluxes in European forests.

Fine roots and above-ground litterfall play a pivotal role in carbon dynamics in forests. Nonetheless, direct estimation of stocks of fine roots remains methodologically challenging. Models are thus widely used to estimate these stocks and help elucidate drivers of fine root growth and turnover, at a range of scales.We updated a database of fine root biomass, necromass and production derived from 454 plots across European forests. We then compared fine root biomass and production to estimates obtained from 19 different models. Typical input variables used for the models included climate, net primary production, foliage and above-ground biomass, leaf area index (LAI), latitude and/or land cover type. We tested whether performance could be improved by fitting new multiple regression models, and explored effects of species composition and sampling method on estimated fine root biomass.Average fine root biomass was 332 g/m2, and necromass 379 g/m2, for European forests where the average fine root production was 250 g m-2 year-1. Carbon fraction in fine roots averaged 48.4%, and was 1.5% greater in broadleaved species than conifers.Available models were poor predictors of fine root biomass and production. The best performing models assumed proportionality between above- and below-ground compartments, and used remotely sensed LAI or foliage biomass as key inputs. Model performance was improved by use of multiple regressions, which revealed consistently greater biomass and production in stands dominated by broadleaved species as well as in mixed stands even after accounting for climatic differences. Synthesis. We assessed the potential of existing models to estimate fine root biomass and production in European forests. We show that recalibration reduces by about 40% errors in estimates currently produced by the best available models, and increases three-fold explained variation. Our results underline the quantitative significance of fine roots (live and dead) to the global carbon cycle.

Ectomycorrhizal community structure in a healthy and a Phytophthora-infected chestnut (Castanea sativa Mill.) stand in central Italy.

Ink disease caused by Phytophthora cambivora is a major disease of sweet chestnut (Castanea sativa). In two C. sativa stands in central Italy, one (Montesanti) that is infected with P. cambivora and the trees showing symptoms of ink disease and another healthy stand (Puzzella), the ectomycorrhizal (ECM) community structure was investigated. On the roots of the surviving trees of the diseased stand, 29 different ECM species were determined compared to 23 in the healthy stand. Eleven ECM species were common to both stands; however, a number of species were unique to one of the stands. Cenococcum geophilum was dominant at both sites, but the percentage colonisation was much higher at Montesanti (40.8%) compared to Puzzella (27.2%). There was a switch in species from Russula vesca, Russula lepida and Russula azurea at Puzzella to Russula nigricans, R. lepida and Russula delica at Montesanti. Both R. vesca and R. azurea were found only at the Puzzella site. At the diseased site, the ECMs formed had a smaller root tip diameter, and the ECM at the healthy site had more abundant extramatrical hyphae.

Next generation of elevated [CO2] experiments with crops: a critical investment for feeding the future world.

A rising global population and demand for protein-rich diets are increasing pressure to maximize agricultural productivity. Rising atmospheric [CO(2)] is altering global temperature and precipitation patterns, which challenges agricultural productivity. While rising [CO(2)] provides a unique opportunity to increase the productivity of C(3) crops, average yield stimulation observed to date is well below potential gains. Thus, there is room for improving productivity. However, only a fraction of available germplasm of crops has been tested for CO(2) responsiveness. Yield is a complex phenotypic trait determined by the interactions of a genotype with the environment. Selection of promising genotypes and characterization of response mechanisms will only be effective if crop improvement and systems biology approaches are closely linked to production environments, that is, on the farm within major growing regions. Free air CO(2) enrichment (FACE) experiments can provide the platform upon which to conduct genetic screening and elucidate the inheritance and mechanisms that underlie genotypic differences in productivity under elevated [CO(2)]. We propose a new generation of large-scale, low-cost per unit area FACE experiments to identify the most CO(2)-responsive genotypes and provide starting lines for future breeding programmes. This is necessary if we are to realize the potential for yield gains in the future.

Mycorrhizas and soil ecosystem function of co-existing woody vegetation islands at the alpine tree line.

BACKGROUND AND AIMS: Picea abies, Pinus mugo and Rhododendron ferrugineum co-exist at the alpine tree line, and can have different mycorrhizal communities. The activity and diversity of mycorrhizal fungi are considered to be important factors in regulation of soil function. METHODS: At a tree line site and a lower elevation site in the Austrian Alps, the community structure of ectomycorrhiza on Picea abies and Pinus mugo was determined. The activity of surface enzymes was determined on ectomycorrhizal and ericoid mycorrhizal roots. In soils, the activity of a range of enzymes, nitrogen (N) mineralization and biomass decomposition were determined. RESULTS: The community structure of the ectomycorrhizal community of Picea abies and Pinus mugo differed strongly, but the average activity of surface enzymes of the ectomycorrhizal communities was similar. A lower root surface enzyme activity was determined on Rhododendron ferrugineum. Soil N-mineralization under Rhododendron ferrugineum was significantly lower than under Picea abies and Pinus mugo. In soil, the activity of a range of enzymes did not differ at the tree line but differed between the tree line and the lower elevation sites. CONCLUSION: The different ectomycorrhizal communities on Picea abies and Pinus mugo and ericoid mycorrhizas on Rhododendron ferrugineum support similar ecosystem functions in soil.

Changes in land use alter soil quality and aggregate stability in the highlands of northern Ethiopia.

Land use change alters biodiversity and soil quality and thus affects ecosystem functions. This study investigated the effects of changes in land use on major soil quality indicators. Soil samples were taken from a depth of 0-10 cm (top soil) under four major land uses (cropland, grassland, area exclosure, eucalyptus plantation) with similar land use change histories for analysis, and soil from a nearby natural forest was used as a reference. Land use change from natural forest to cropland and grassland significantly decreased major soil quality indicators such as soil organic C (SOC), total soil N (TSN), molybdate-reactive bicarbonate-extractable P, and arbuscular mycorrhizal fungi (AMF) spore density, but compared to the cropland, change to area exclosure and eucalyptus plantation significantly improved SOC, TSN and soil aggregate stability (SAS). In addition, we assessed the correlation among indicators and found that SOC, TSN and SAS significantly correlate with many other soil quality indicators. The study highlights that the conversion of natural forest to cropland results in decline of soil quality and aggregate stability. However, compared to cropland, application of area exclosure and afforestation on degraded lands restores soil quality and aggregate stability.

Growth and mycorrhizal colonization of three North American tree species under elevated atmospheric CO2.

We investigated the effect of elevated CO2 on the growth and mycorrhizal colonization of three tree species native to north-eastern American forests (Betula papyrifera Marsh., Pinus strobus L. and Tsuga Canadensis L. Carr). Saplings of the tree species were collected from Harvard Forest, Massachusetts, and grown in forest soil under ambient (c. 375 ppm) and elevated (700 ppm) atmospheric CO., concentrations for 27-35 wk. In all three species there was a trend to increasing whole-plant, total-root and fine-root biomass in elevated CO2 , and a significant increase in the degree of ectomycorrhizal colonization in B. papyrifera and P. strobus, but not in T. Canadensis. However, in T, Canadensis the degree of colonization with arbuscular mycorrhizas increased significantly. In both the ambient and elevated environments, on the roots of B. papyrifera and P. strobus 12 distinct ectomycorrhizal morphotypes were identified. Distinct changes in the ectomycorrhizal morphotype assemblage of B. papvrifera were observed under CO2 enrichment. This change resulted in an increase in the frequency of ectomycorrhizas with a higher incidence of emanating hyphae and rhizomorphs, and resulted in a higher density of fungal hyphae in a root exclusion chamber.

Nitrogen and phosphorus acquisition by the mycelium of the ectomycorrhizal fungus Paxillus involutus and its effect on host nutrition.

The contribution of the extramatrical mycelium to N and P nutrition of mycorrhizal Norway spruce (Picea abies (L.) Karst.) was investigated. Seedlings either inoculated with Paxillus involutus (Batsch) Fr. or non-mycorrhizal were grown in a two compartment sand culture system where hyphae were separated from roots by a 45 µm nylon net. Nutrient solution of the hyphal compartment contained either 1.8 mm NH4 + and 0.18 mm H2 PO4 - or no N and P. Aluminium added to the hyphal compartment as a tracer of mass flow was not detected in the plant compartment, indicating that measurements of N and P transfer by the mycelium were not biased by solute movement across the nylon net. The addition of N and P to the hyphal compartment markedly increased dry weight, N and P concentration and N and P content of mycorrhizal plants. Calculating uptake from the difference in input and output of nutrient in solution confirmed a hyphal contribution of 73% and 76% to total N and P uptake, respectively. Hyphal growth was increased at the site of nutrient solution input.

Interdependence of phosphorus, nitrogen, potassium and magnesium translocation by the ectomycorrhizal fungus Paxillus involutus.

• Translocation is shown of phosphorus, nitrogen, potassium and magnesium to a P-deficient host from ectomycorrhizal fungal hyphae. • Mycorrhizal (with Paxillus involutus) and nonmycorrhizal P-deficient spruce (P. abies) seedlings were grown in a two-compartment sand-culture system. Hyphal translocation of nutrients from the inner compartment (penetrated only by hyphae) to the host was measured using mass balance (for N, P and K) or stable isotope (15 N and 25 Mg) methods. • Addition of P to the hyphal compartment strongly stimulated hyphal growth, and this also increased both seedling P status and growth. Hyphae translocated nonlimiting elements in addition to P, contributing 52, 17, 5 and 3-4%, respectively, to total P, N, K or Mg plant uptake. The potential role of the ectomycorrhizal mycelium in K acquisition was demonstrated. Translocation to mycorrhizal seedings of N, K and Mg was strongly reduced when hyphal P-fluxes ceased; this translocation of nonlimiting nutrients depended on simultaneous translocation of P. • The ectomycorrhizal mycelium has an active role in P acquisition from sources not available to roots. Nutrient fluxes within fungal hyphae are interdependent and strong coupling of N, K and Mg fluxes with long-distance P translocation in the mycorrhizal mycelium occurs.