Search

Endophytic fungi are known to be present in roots of salt marsh plants, but their ecological role in this symbiosis is still largely unknown. Generally considered parasitic or saprophytic, they may still be mutualistic, at least under certain circumstances. Among salt marsh plants,Salicorniaspp. are recognized as particularly salt-tolerant and their frequent colonization by root endophytes has also been reported. This study aimed to investigate whether the inoculation ofSalicorniawith different root endophytes isolated from field-collectedSalicorniaaffects biomass production, nutrient uptake and photosynthesis (assessed via chlorophyll fluorescence). In addition, we investigated whether fungal inoculation confers tolerance to salt stress given that endophytes are suggested to increase salt tolerance and improve plant fitness in other less salt-tolerant plants. The inoculation ofSalicorniawith an isolate of the genusStemphyliumpositively influenced total biomass production and nitrogen concentration in roots at optimum salinity condition (150 mM NaCl). However, under salt stress (650 mM NaCl), no significant effects of fungal inoculation on biomass production and photosynthesis were observed. Further, positive and negative effects of fungal inoculation on nutrient concentrations were observed in roots and shoots, respectively. Our results indicate that different endophytic fungi and their interaction result in distinct fungal species-specific plant growth responses ofSalicorniaunder different growth conditions.

Root-associated fungi (RAF) link nutrient fluxes between soil and roots and thus play important roles in ecosystem functioning. To enhance our understanding of the factors that control RAF, we fitted statistical models to explain variation in RAF community structure using data from 150 temperate forest sites covering a broad range of environmental conditions and chemical root traits. We found that variation in RAF communities was related to both root traits (e.g., cations, carbohydrates, NO3 - ) and soil properties (pH, cations, moisture, C/N). The identified drivers were the combined result of distinct response patterns of fungal taxa (determined at the rank of orders) to biotic and abiotic factors. Our results support that RAF community variation is related to evolutionary adaptedness of fungal lineages and consequently, drivers of RAF communities are context-dependent.

Polyploidy is recognized as a mechanism of speciation in plants with cascading effects on biotic interactions. However, a limited number of studies have investigated the effects of polyploidy on the association of plants and microorganisms. Herein, we investigated whether two Salicornia cytotypes (S. europaea – 2x and S. procumbens – 4x) show different root-associated fungal communities. Additionally, we explored the existence of cytotype-specific root anatomical traits, which could influence fungal recruitment and establishment. Salicornia spp. were identified based on their ploidy level. The root-associated fungal microbiome of Salicornia was analyzed using high throughput amplicon sequencing (ITS1 of rDNA) in spring and summer. The following root anatomical traits were investigated: maximum root diameter, periderma thickness, parenchyma thickness, diameter of the vascular cylinder and maximum diameter of parenchyma cells. Our results showed that Shannon diversity and evenness indices were higher in samples of Salicornia procumbens (4x) compared to those of S. europaea (2x), and in summer the root-associated fungal community of S. procumbens (4x) was significantly different from that of S. europaea (2x). The orders Xylariales, Malasseziales and Pleosporales were the most frequent root colonizers in both cytotypes and most of the taxa associated with Salicornia were functionally classified as saprophytes or plant pathogens. Finally, we observed larger periderma and parenchyma layers in S. procumbens (4x) than S. europaea (2x) that may contribute to the observed differences in community composition between the two cytotypes. Our results suggest that differences in ploidy may modulate plant interaction with fungi by affecting species recruitment and microbiome structure. In addition, cytotype-specific root traits may also have the potential to affect differently community assembly in the two cytotypes.

Pongamia pinnata (L.) Pierre (Pongamia) is a tree native to Southeast Asia. Recently, interest in Pongamia focused on its potential as a biofuel source as its seeds contain around 40% oil. However, Pongamia has multiple applications beyond biofuel production. It is a legume, can form symbiotic associations with mycorrhizal fungi, has been shown to be tolerant to drought, salinity, and heavy metals in soil, and has potential to mitigate climate change. Additionally, Pongamia oil has medicinal properties, can be used as biopesticide, insect repellent, to produce soap, and as a source of edible grade vegetable oil. The seed cake can be used as a source of bioenergy, food and feed protein, and organic fertiliser, and the flowers are a good source of pollen and nectar. Pongamia can also bring socio-economic benefits as its ability to restore degraded and contaminated land provides opportunities for local communities through novel valorisation pathways. These multiple applications have potential to form part of a circular bioeconomy in line with sustainable development goals. Although research on the multiple applications of Pongamia has grown considerably, knowledge gaps remain and these need to be addressed so that the full potential of Pongamia can be achieved. Further understanding of the mechanisms underlying its resilience to abiotic stresses, phytoremediation potential and biotic interactions should be a priority, and co-ordinated breeding efforts will be key. Here, we critically review the available literature on Pongamia and highlight gaps in knowledge in which future research should focus on to ensure that the full potential of this versatile tree can be achieved. We conclude that Pongamia can potentially form part of a circular bioeconomy and that harnessing the multiple applications of Pongamia in a holistic manner, with collaboration among key stakeholders, is crucial for the successful application of its benefits far beyond biofuel production.

Functionally, ectomycorrhizal (ECM) and saprotrophic (SAP) fungi belong to different guilds, and they play contrasting roles in forest ecosystem C-cycling. SAP fungi acquire C by degrading the soil organic material, which precipitates massive CO2 release, whereas, as plant symbionts, ECM fungi receive C from plants representing a channel of recently assimilated C to the soil. In this study, we aim to measure the amounts and identify the drivers of ECM and SAP fungal biomass in temperate forest topsoil. To this end, we measured ECM and SAP fungal biomass in mineral topsoils (0–12 cm depth) of different forest types (pure European beech, pure conifers, and mixed European beech with other broadleaf trees or conifers) in a range of about 800 km across Germany; moreover, we conducted multi-model inference analyses using variables for forest and vegetation, nutritive resources from soil and roots, and soil conditions as potential drivers of fungal biomass. Total fungal biomass ranged from 2.4 ± 0.3 mg g−1 (soil dry weight) in pure European beech to 5.2 ± 0.8 mg g−1 in pure conifer forests. Forest type, particularly the conifer presence, had a strong effect on SAP biomass, which ranged from a mean value of 1.5 ± 0.1 mg g−1 in broadleaf to 3.3 ± 0.6 mg g−1 in conifer forests. The European beech forests had the lowest ECM fungal biomass (1.1 ± 0.3 mg g−1), but in mixtures with other broadleaf species, ECM biomass had the highest value (2.3 ± 0.2 mg g−1) among other forest types. Resources from soil and roots such as N and C concentrations or C:N ratios were the most influential variables for both SAP and ECM biomass. Furthermore, SAP biomass were driven by factors related to forest structure and vegetation, whereas ECM biomass was mainly influenced by factors related to soil conditions, such as soil temperature, moisture, and pH. Our results show that we need to consider a complex of factors differentially affecting biomass of soil fungal functional groups and highlight the potential of forest management to control forest C-storage and the consequences of changes in soil fungal biomass.

Nutrient imbalances cause the deterioration of tree health in European forests, but the underlying physiological mechanisms are unknown. Here, we investigated the consequences of decreasing root carbohydrate reserves for phosphorus (P) mobilization and uptake by forest trees. In P-rich and P-poor beech (Fagus sylvatica) forests, naturally grown, young trees were girdled and used to determine root, ectomycorrhizal and microbial activities related to P mobilization in the organic layer and mineral topsoil in comparison with those in non-girdled trees. After girdling, root carbohydrate reserves decreased. Root phosphoenolpyruvate carboxylase activities linking carbon and P metabolism increased. Root and ectomycorrhizal phosphatase activities and the abundances of bacterial genes catalysing major steps in P turnover increased, but soil enzymes involved in P mobilization were unaffected. The physiological responses to girdling were stronger in P-poor than in P-rich forests. P uptake was decreased after girdling. The soluble and total P concentrations in roots were stable, but fine root biomass declined after girdling. Our results support that carbohydrate depletion results in reduced P uptake, enhanced internal P remobilization and root biomass trade-off to compensate for the P shortage. Since reductions in root biomass render trees more susceptible to drought, our results link tree deterioration with disturbances in the P supply as a consequence of decreased belowground carbohydrate allocation.

While forest management strongly influences biodiversity, it remains unclear how the structural and compositional changes caused by management affect different community dimensions (e.g. richness, specialisation, abundance or completeness) and how this differs between taxa. We assessed the effects of nine forest features (representing stand structure, heterogeneity and tree composition) on thirteen above‐ and belowground trophic groups of plants, animals, fungi and bacteria in 150 temperate forest plots differing in their management type. Canopy cover decreased light resources, which increased community specialisation but reduced overall diversity and abundance. Features increasing resource types and diversifying microhabitats (admixing of oaks and conifers) were important and mostly affected richness. Belowground groups responded differently to those aboveground and had weaker responses to most forest features. Our results show that we need to consider forest features rather than broad management types and highlight the importance of considering several groups and community dimensions to better inform conservation. The work was partly funded by the DFG Priority Program 1374 ‘Infrastructure-Biodiversity-Exploratories’ (DFG-Refno. Po362/18-3). SSo was supported by the Spanish Government under a Ramón y Cajal contract (RYC-2016-20604).

Root-associated mycobiomes (RAMs) link plant and soil ecological processes, thereby supporting ecosystem functions. Understanding the forces that govern the assembly of RAMs is key to sustainable ecosystem management. Here, we dissected RAMs according to functional guilds and combined phylogenetic and multivariate analyses to distinguish and quantify the forces driving RAM assembly processes. Across large biogeographic scales (>1,000 km) in temperate forests (>100 plots), RAMs were taxonomically highly distinct but composed of a stable trophic structure encompassing symbiotrophic, ectomycorrhizal (55%), saprotrophic (7%), endotrophic (3%) and pathotrophic fungi (

There is growing interest in the application of plant functional trait-based approaches for development of sustainable land-use strategies. In this context, one crucial task is to identify and measure plant traits, which respond to land-use intensity (response traits) and simultaneously have an impact on ecosystem functions (effect traits). We hypothesized that species-specific leaf chemical composition, which may function both as response and effect trait, can be derived from Attenuated Total Reflection Fourier Transform Infrared (ATR-FTIR) spectroscopy tools in combination with multivariate statistical methods We investigated leaf ATR-FTIR spectra of two grasses, Poa pratensis L. and Dactylis glomerata L., and one forb, Achillea millefolium L. collected in grassland plots along a land-use intensity gradient in three regions of Germany. ATR-FTIR spectra appear to function as biochemical fingerprints unique to each species. The spectral response to land-use intensity was not consistent among species and less apparent in the two grasses than in the forb species. Whereas land-use intensification enhanced protein and cellulose content in A. millefolium , giving rise to changes in six spectral bands in the frequency range of 1088–1699 cm −1 , only cellulose content increased in D. glomerata , affecting the bands of 1385–1394 cm −1 . Poa pratensis spectra exhibited minimal changes under the influence of land-use, only in the spectral bands of 1373–1375 cm −1 associated with suberin-like aliphatic compounds. Our findings suggest that some species’ leaf chemical composition is responsive to land-use intensity, and thus, may have a predictive value for ecosystem services provided by those species within grassland vegetation (i.e., herbage yield quality).

Beech in original forest soil are colonized by higher ectomycorrhizal diversity and show better performance than those in γ-irradiated soil but no provenances-related differences. Knowledge on the effects of the intra-specific diversity of beech and inter-specific diversity of ectomycorrhizal fungal species (EMF) on the performance and nitrogen (N) uptake of beech (Fagus sylvatica, L) provenances are scarce. Here, we investigated EMF diversity and performance (growth, photosynthesis, N nutrition) of beech provenances from three different geographic regions: Schwabische Alb, Hainich (HAI) and Schorfheide. The seedlings were grown in a common environment in either original HAI forest soil or in γ-irradiated HAI soil to which soil bacteria had been added. The soil treatments resulted in high (up to four fungi) or low EMF diversity (one fungus) at the root tips, but not in differences among the provenances or in mycorrhizal colonization. The provenances exhibited better performance in original than in treated soil, with the HAI provenance showing highest biomass in original soil. The effect was lost in treated soil, where EMF diversity was low suggesting that the positive interaction between soil and provenances was mediated by soil microbes but not by genetic adaptation of the beech provenances. Short-term labelling with 15NH4 + or 15NO3 − did not show provenance-related differences, but higher aboveground allocation of 15NO3 − than of 15NH4 +-derived N. Ordination of root tip community composition with N uptake and performance parameters revealed correspondence of low fungal diversity in treated soil with 15N uptake and better performance of plants with higher fungal diversity in original soil. These results suggest control of plant performance and N supply by EMF diversity.

For 15+ years, a beech (Fagus sylvatica L.) dominated forest on calcareous soil was studied on two opposing slopes with contrasting microclimate in Tuttlingen, Swabian Alb, Germany. The cool-humid NE aspect of these slopes represents the majority of beech forests under current climate, the warmer and drier SW aspect represents beech forests under future climate conditions. The field studies were supplemented by investigations under controlled conditions. The research program aimed to provide a comprehensive understanding of plant-soil-microbe water, carbon and nitrogen feedbacks in a changing climate and a holistic view of the sensitivity of beech to climate change. The results of comparative and experimental studies underpin the high vulnerability of adult beech and its natural regeneration on calcareous soil to both direct climate change effects on plant physiology and indirect effects mediated by soil biogeochemical cycles. Mechanisms contributing to this vulnerability at the ecosystem and organismic level indicate a high significance of competitive interactions of beech with other vegetation components and soil microbial communities. Obvious forest management practices such as selective felling did not necessarily counteract negative effects of climate change.

European beech presents intraspecific variations in drought resistance strategies that are mediated by the amount of precipitation in the habitat. Climate change predictions forecast extended drought periods, which are expected to pose an enhanced risk to forest trees. Here, we investigated the drought response and fitness traits in European beech (Fagus sylvatica L.) in response to a severe progressive drought. The capability of three beech provenances from habitats differing in annual precipitation (544, 665, and 766 mm year−1) to cope with drought was compared in a common garden experiment using beech seedlings. Soil and plant water status, maximum quantum yield of PSII, growth and biomass partitioning, stomatal conductance, and transcript abundance pattern of the kinase, Open Stomata 1 (OST1), of control (well-watered) and drought-treated (water withheld) plants from each provenance were repeatedly measured during a 60-day drought experiment. The lowest precipitation provenance displayed a more isohydric phenotype with a prompt stomatal closure, increased OST1 levels, high water potential and leaf water content, and a decrement in the maximum quantum yield of PSII. The other two provenances showed a more anisohydric stomatal regulation with a slow and delayed stomatal closure and a decrease in the water status. These findings suggest that intraspecific variations in beech for diverging drought resistance strategies exist and might be mediated by differences in the abscisic acid signaling pathway. The higher precipitation provenance maintained high quantum yield of PSII, and water potentials above −2.0 MPa for a longer period of time than the other two provenances, and consequently, mortality was delayed in this provenance. We concluded that lower precipitation adapted plants employ a drought resistance strategy suitable for the moderate drought, whereas the higher precipitation habitat plants revealed mechanisms, which could be better suited to cope with more severe drought events.

Phosphorus (P) is an important nutrient, whose concentrations are declining in many European forest ecosystems. Here, we selected five old-aged temperate beech ( Fagus sylvatica ) forests that represented a sequence of decreasing soil P resources. We addressed the following hypotheses: (i) root P concentrations correspond to soil P concentrations, when P availability is suboptimal for tree nutrition, (ii) decreasing soil P concentrations, and increasing host P demand foster increasing ectomycorrhizal fungal (EMF) species richness and lead to a shift in the EMF community structure towards increasing soil exploration. We found that the decrease in P concentrations along the geosequence was less steep in the organic layer than that in the mineral topsoil. P concentrations in roots showed a positive relationship with P concentrations in soil, with a stronger correlation in coarse than in fine roots. This finding indicates that low P availability mainly affected P storage of the host. The root tips were completely colonized with EMF. In the organic layer EMF biomass was higher than that of saprophytic fungi, and correlated with inorganic P (P i ). In the mineral topsoil EMF biomass was about 10-fold lower than in the organic layer and biomass of saprophytes and microbial P, but not that of EMF, was correlated with P i and phosphatase activities. Based on these results, we propose that beech P nutrition was mainly achieved by EMF in the organic layer. Variation in EMF species richness was unrelated to P in soil and decreased with increasing N in the organic layer. The EMF community structures were taxonomically divergent and filtered by habitat soil chemistry in the mineral layer and P i in the organic layer between the P-rich forest and the P-poor forest. Changes in the taxonomic structures of the EMF did not result in corresponding changes in soil exploration. In conclusion, our results support a relationship between soil P concentrations and P storage in roots, but do not support mono-causal relationships between soil P and EMF species richness or hyphal soil exploration. Our results suggest that the taxonomic dissimilarities of the EMF along the P gradient were mainly driven by P i concentrations in the organic layer and by the nutrient resources in the mineral layer.

In forest ecosystems, ectomycorrhizal (ECM) fungi are important for plant growth and soil biogeochemical processes. The biochemical composition of ECM mycelium is an important fungal effect trait with consequences for its decomposition rate, and consequently on soil carbon pools and plant nutrition. Although the link between ECM fungi and leaf litter–released nutrients is well known, the response of ECM fungal biochemical composition to different leaf litter species remains poorly understood. To determine how leaf litter quality influences ECM fungi’s biochemical profiles, we planted young beech trees in an oak forest and replaced the natural leaf litter with that of European beech (Fagus sylvatica), ash (Fraxinus excelsior), maple (Acer pseudoplatanus), or lime (Tilia cordata). We assessed the biochemical profiles of ECM root tips colonized by common fungal taxa in temperate forests (i.e., Cenococcum geophilum, Inocybe sp., and Lactarius subdulcis), using attenuated total reflectance Fourier transform infrared spectroscopy (ATR-FTIR). ECM fungal biochemical composition changed with leaf litter species. Changes were apparent in the infrared absorption bands assigned to functional groups of lipids, amides, and carbohydrates. C. geophilum and L. subdulcis exhibited large spectral differences corresponding to the initial pattern of leaf litter chemical composition between samples collected in the beech and ash leaf litter treatments. In contrast, Inocybe sp. was influenced by lime, but with no differences between samples from ash or beech leaf litter treatments. Although the spectral bands affected by leaf litter type differed among ECM fungi, they were mainly related to amides, indicating a dynamic response of the fungal proteome to soil nutritional changes. Overall, the results indicate that the biochemical response of ECM fungi to leaf litter species varies among ECM fungal species and suggests that the biochemical composition of ECM mycelium is a fungal response trait, sensitive to environmental changes such as shifts in leaf litter species.

Phosphorus (P) is often a limiting macronutrient in temperate forests, but knowledge on the phenological and physiological responses of beech (Fagus sylvatica L.) to P deficiency is scarce. In this study, young beech trees were excavated with intact soil cores from two German forests, Unterluss (LUE) with low soil P and Bad Bruckenau (BBR) with high soil P concentrations. The trees were transferred to identical climatic conditions. In the subsequent growth phase phenological stages during bud burst and leaf unfolding were recorded; biomass production and total P concentrations in different tissues were measured. Seasonal fluctuations in photosynthesis and of soluble P in wood and bark exudates were determined. BBR beeches grew faster and produced more and larger leaves than the LUE beeches. Leaf extension and unfolding were delayed in LUE compared with BBR beeches, but not the time point of bud break. All plant tissues of BBR trees contained higher total P concentrations than those of LUE trees. Strong seasonal fluctuations for P in exudates of beech transport tissues, wood and bark, indicated higher P supply in BBR than in LUE plants, especially at the beginning of the growth phase until leaf maturity. Photosynthetic activity of LUE beeches was lower than that of BBR beeches due to stomatal limitations as the result of anatomically smaller stomatal pore widths, but not as the result of acute biochemical limitation of photosynthesis. Our results suggest that developmental retardation and lower photosynthesis under low P availability may be adaptation mechanisms that adjust the acquisition and recycling of P resources to seasonal growth demand.

Earthworms (EWs) and mycorrhizal fungi are key components of soil biota participating in N recycling from leaf litter. The extent to which their interactions impact leaf litter-derived N uptake by tree species colonized by different mycorrhizal types and the variation of these interactions with leaf litter quality is unknown. We used a greenhouse experiment to investigate the effects of EWs on plant acquisition of N from 15 N enriched ash ( Fraxinus excelsior ) and beech ( Fagus sylvatica ) leaf litter. We employed arbuscular mycorrhizal (AM) ash and ectomycorrhizal (EM) beech trees. EW presence did not affect the overall root mycorrhizal colonization, but in the beech leaf litter treatment it caused a shift in the EM fungal community, favoring species with a ruderal life strategy at the expense of more specialized species able to degrade organic compounds. No differences in 15 N capture were found between EM and AM plants. Generally, 2–7% of the N released from leaf litter was taken up by the plants. EWs uniformly increased plant acquisition of N from leaf litter. This effect was more pronounced in treatments with ash than in those with beech leaf litter, presumably because of higher degradability of ash than of beech litter. The results suggest that leaf litter quality dominates the impact of EWs on plant N acquisition from leaf litter while the mycorrhizal type is of minor importance.

Research highlights: Interaction effects of coniferous on deciduous species have been investigated before the background of climate change. Background and objectives: The cultivation of European beech (Fagus sylvatica L.) in mixed stands has currently received attention, since the future performance of beech in mid-European forest monocultures in a changing climate is under debate. We investigated water relations and nitrogen (N) nutrition of beech in monocultures and mixed with silver-fir (Abies alba Mill.) in the Black Forest at different environmental conditions, and in the Croatian Velebit at the southern distribution limit of beech, over a seasonal course at sufficient water availability. Material and methods: Water relations were analyzed via &delta, 13C signatures, as integrative measures of water supply assuming that photosynthesis processes were not impaired. N nutrition was characterized by N partitioning between soluble N fractions and structural N. Results: In the relatively wet year 2016, water relations of beech leaves, fir needles and roots differed by season, but generally not between beech monocultures and mixed cultivation. At all sites, previous and current year fir needles revealed significantly lower total N contents over the entire season than beech leaves. Fir fine roots exhibited higher or similar amounts of total N compared to needles. Correlation analysis revealed a strong relationship of leaf and root &delta, 13C signatures with soil parameters at the mixed beech stands, but not at pure beech stands. While glutamine (Gln) uptake capacity of beech roots was strongly related to soil N in the monoculture beech stands, arginine (Arg) uptake capacities of beech roots were strongly related to soil N in mixed stands. Conclusions: Leaf N contents indicated a facilitative effect of silver-fir on beech on sites where soil total N concentrations where low, but an indication of competition effect where it was high. This improvement could be partially attributed to protein contents, but not to differences in uptake capacity of an individual N source. From these results it is concluded that despite similar performance of beech trees at the three field sites investigated, the association with silver-fir mediated interactive effects between species association, climate and soil parameters even at sufficient water supply.

Increases in summer droughts and nitrogen (N) deposition have raised concerns of widespread biodiversity loss and nutrient imbalances, but our understanding of the ecological role of ectomycorrhizal fungal (ECMF) diversity in mediating root functions remains a major knowledge gap. We used different global change scenarios to experimentally alter the composition of ECMF communities colonizing European beech saplings and examined the consequences for phosphorus (P) uptake (H333 PO4 feeding experiment) and use efficiencies of trees. Specifically, we simulated increases in temperature and N deposition and decreases in soil moisture and P availability in a factorial experiment. Here, we show that ECMF α diversity is a major factor contributing to root functioning under global change. P uptake efficiency of beech significantly increased with increasing ECMF species richness and diversity, as well as with decreasing P availability. As a consequence of decreases in ECMF diversity, P uptake efficiency decreased when soil moisture was limiting. By contrast, P use efficiencies were a direct (negative) function of P availability and not of ECMF diversity. We conclude that increasing summer droughts may reduce ECMF diversity and the complementarity of P uptake by ECMF species, which will add to negative growth effects expected from nutrient imbalances under global change.

International audience; The spatiotemporal dynamics of, and interspecific differences in, the acquisition of litter-derived nitrogen (N) by natural assemblages of ectomycorrhizal root tips are poorly understood. Small cylindrical mesh bags containing 15N-labelled beech (Fagus sylvatica) leaf litter that permit hyphal but not root ingrowth were inserted vertically into the top soil layer of an old-growth beech forest. The lateral transfer of 15N into the circumjacent soil, roots, microbes and ectomycorrhizas was measured during an 18-month exposure period. Ectomycorrhial fungi (EMF) showed large interspecific variation in the temporal pattern and extent of 15N accumulation. Initially, when N was mainly available from the leachate, microbes were more efficient at N immobilization than the majority of EMF, but distinct fungal species also showed significant 15N accumulation. During later phases, the enrichment of 15N in Tomentella badia was higher than in microbes and other EMF species. Roots and soil accumulated 15N with a large delay compared with microbes and EMF. Because approximately half of the studied fungal species had direct access to N from leaf litter and the remainder to N from leached compounds, we suggest that EMF diversity facilitates the N utilization of the host by capturing N originating from early-released solutes and late degradation products from a recalcitrant source.

The aim of this study was to investigate the influence of ectomycorrhizal fungi (EMF) on the architecture of and nitrogen (N) partitioning in young beech (Fagus sylvatica) plants in response to different light regimes and water deprivation. We hypothesized that EMF modify biomass partitioning and architecture of young beech plants by increased N uptake in comparison with non-mycorrhizal (NM) plants and that therefore, the drought responses of EM and NM plants diverge. We anticipated that full light-exposed plants were more drought tolerant due to improved water status and nutrition, whereas shade-acclimated EM plants were more drought susceptible because of decreased mycorrhizal colonization. To test these hypotheses seedlings were grown in native or sterilized forest soil. To avoid effects of soil pretreatment NM and EM plants were transplanted into sand-peat culture systems and exposed to shade, drought or the combination of both factors. Shade resulted in reduced root biomass production decreasing the root-to-shoot ratio. Mild drought stress (pre-dawn water potential [Ψpd] = −1.3 MPa) did not affect biomass partitioning. EMF colonization did not increase plant biomass, but had strong effects on root architecture: the numbers of root tips as well as the absolute and specific root lengths were increased because of formation of thin roots, especially in the diameter classes from 0.2 to 0.8 mm. In contrast to our expectation N uptake of well irrigated EM plants was not increased despite their larger potential for soil exploitation. Overall, EM plants exhibited higher amounts of carbon fixed per unit of N taken up than NM plants and shifted N partitioning towards the roots. Beneficial effects of EMFs were apparent under mild drought but the responses differed depending on the light availability: shaded EM plants showed a delay in the decrease of Ψpd; light exposed EM plants showed increased N uptake compared with NM beeches. These results indicate that EMFs are involved in mediating divergent responses of beech to drought depending on the light availability.

Phosphorus availability may shape plant–microorganism–soil interactions in forest ecosystems. Our aim was to quantify the interactions between soil P availability and P nutrition strategies of European beech (Fagus sylvatica) forests. We assumed that plants and microorganisms of P-rich forests carry over mineral-bound P into the biogeochemical P cycle (acquiring strategy). In contrast, P-poor ecosystems establish tight P cycles to sustain their P demand (recycling strategy). We tested if this conceptual model on supply-controlled P nutrition strategies was consistent with data from five European beech forest ecosystems with different parent materials (geosequence), covering a wide range of total soil P stocks (160–900 g P m−2

Ectomycorrhizal (EM) fungal taxonomic, phylogenetic, and trait diversity (exploration types) were analyzed in beech and conifer forests along a north-to-south gradient in three biogeographic regions in Germany. The taxonomic community structures of the ectomycorrhizal assemblages in top soil were influenced by stand density and forest type, by biogeographic environmental factors (soil physical properties, temperature, and precipitation), and by nitrogen forms (amino acids, ammonium, and nitrate). While α-diversity did not differ between forest types, β-diversity increased, leading to higher γ-diversity on the landscape level when both forest types were present. The highest taxonomic diversity of EM was found in forests in cool, moist climate on clay and silty soils and the lowest in the forests in warm, dry climate on sandy soils. In the region with higher taxonomic diversity, phylogenetic clustering was found, but not trait clustering. In the warm region, trait clustering occurred despite neutral phylogenetic effects. These results suggest that different forest types and favorable environmental conditions in forests promote high EM species richness in top soil presumably with both high functional diversity and phylogenetic redundancy, while stressful environmental conditions lead to lower species richness and functional redundancy.

Dannenmann, M. et al.-- 10 pages, 1 figure, 2 tables, 60 references., Nitrogen (N) cycling in terrestrial ecosystems is complex since it involves the closely interwoven processes of both N uptake by plants and microbial turnover of a variety of N metabolites. Major interactions between plants and microorganisms involve competition for the same N species, provision of plant nutrients by microorganisms and labile carbon (C) supply to microorganisms by plants via root exudation. Despite these close links between microbial N metabolism and plant N uptake, only a few studies have tried to overcome isolated views of plant N acquisition or microbial N fluxes. In this study we studied competitive patterns of N fluxes in a mountainous beech forest ecosystem between both plants and microorganisms by reducing rhizodeposition by tree girdling. Besides labile C and N pools in soil, we investigated total microbial biomass in soil, microbial N turnover (N mineralization, nitrification, denitrification, microbial immobilization) as well as microbial community structure using denitrifiers and mycorrhizal fungi as model organisms for important functional groups. Furthermore, plant uptake of organic and inorganic N and N metabolite profiles in roots were determined. Surprisingly plants preferred organic N over inorganic N and nitrate (NO3 ) over ammonium (NH4 þ ) in all treatments. Microbial N turnover and microbial biomass were in general negatively correlated to plant N acquisition and plant N pools, thus indicating strong competition for N between plants and free living microorganisms. The abundance of the dominant mycorrhizal fungi Cenococcum geophilum was negatively correlated to total soil microbial biomass but positively correlated to glutamine uptake by beech and amino acid concentration in fine roots indicating a significant role of this mycorrhizal fungus in the acquisition of organic N by beech. Tree girdling in general resulted in a decrease of dissolved organic carbon and total microbial biomass in soil while the abundance of C. geophilum remained unaffected, and N uptake by plants was increased. Overall, the girdling-induced decline of rhizodeposition altered the competitive balance of N partitioning in favour of beech and its most abundant mycorrhizal symbiont and at the expense of heterotrophic N turnover by free living microorganisms in soil. Similar to tree girdling, drought periods followed by intensive drying/rewetting events seemed to have favoured N acquisition by plants at the expense of free living microorganisms., Funding of this work by the German Research Foundation/ Deutsche Forschungsgemeinschaft (DFG) within the framework of the Beech Research Group under contract numbers FOR 788/1, MA 749/21-1, KO 1035/29-1, RE 515/27-1 and PO 362/17-1 is gratefully acknowledged.

In most ecosystems roots and their associated mycorrhizal fungi are a significant component of the soil food web where many bacteria, fungi, and animals directly or indirectly depend on them for much of their nutrition. This dependency is to a large extent mutual: decomposers (saprotrophs) in soil are the primary agents in releasing vital nutrients locked up in organic matter, which in many cases neither plant roots nor mycorrhizal fungi can effectively release. Therefore plants and mycorrhizal fungi engage in an intriguing relationship with saprotrophs in which they act as a main substrate for saprotrophs and simultaneously depend on them and compete with them for soil nutrients. These multiple interactions present us with difficulties in generalizing the effect of mycorrhizal fungi on saprotrophs. In some cases their interests are aligned and saprotrophs and mycorrhizal fungi can act in concert in releasing and taking up essential nutrients; in others competition may be more important. In this chapter we discuss the ways in which different types of mycorrhizal fungi (arbuscular, ectomycorrhizal, and ericoid) can prime soil organic matter decomposition, compete with saprotrophs for substrate, and engage in other ways with members of the soil food web to affect soil organic carbon stocks.

Distance decay, the general reduction in similarity of community composition with increasing geographical distance, is known as predictor of spatial variation and distribution patterns of organisms. However, changes in fungal communities along environmental gradients are little known. Here we show that distance decays of soil-inhabiting and root-associated fungal assemblages differ and identify explanatory environmental variables. High-throughput sequencing analysis of fungal communities of beech-dominated forests at three study sites across Germany shows that root-associated fungi are recruited from the soil fungal community. However, distance decay is substantially weaker in the root-associated than in the soil community. Variance partitioning of factors contributing to the observed distance decay patterns support the hypothesis that host trees stabilize the composition of root-associated fungi communities, relative to soil communities. Thus, they not only have selective impacts on associated communities, but also buffer effects of changes in microclimatic and environmental variables that directly influence fungal community composition.

Mycorrhizal fungi have a key role in nitrogen (N) cycling, particularly in boreal and temperate ecosystems. However, the significance of ectomycorrhizal fungal (EMF) diversity for this important ecosystem function is unknown. Here, EMF taxon-specific N uptake was analyzed via (15)N isotope enrichment in complex root-associated assemblages and non-mycorrhizal root tips in controlled experiments. Specific (15)N enrichment in ectomycorrhizas, which represents the N influx and export, as well as the exchange of (15)N with the N pool of the root tip, was dependent on the fungal identity. Light or water deprivation revealed interspecific response diversity for N uptake. Partial taxon-specific N fluxes for ectomycorrhizas were assessed, and the benefits of EMF assemblages for plant N nutrition were estimated. We demonstrated that ectomycorrhizal assemblages provide advantages for inorganic N uptake compared with non-mycorrhizal roots under environmental constraints but not for unstressed plants. These benefits were realized via stress activation of distinct EMF taxa, which suggests significant functional diversity within EMF assemblages. We developed and validated a model that predicts net N flux into the plant based on taxon-specific (15)N enrichment in ectomycorrhizal root tips. These results open a new avenue to characterize the functional traits of EMF taxa in complex communities.

International audience; Our aims were to characterize the fate of leaf-litter-derived nitrogen in the plant-soil-microbe system of a temperate beech forest of Southern Germany and to identify its importance for N nutrition of beech seedlings. N-15-labelled leaf litter was traced in situ into abiotic and biotic N pools in mineral soil as well as into beech seedlings and mycorrhizal root tips over three growing seasons. There was a rapid transfer of N-15 into the mineral soil already 21 days after tracer application with soil microbial biomass initially representing the dominant litter-N sink. However, N-15 recovery in non-extractable soil N pools strongly increased over time and subsequently became the dominant N-15 sink. Recovery in plant biomass accounted for only 0.025 % of N-15 excess after 876 days. After three growing seasons, N-15 excess recovery was characterized by the following sequence: non-extractable soil N >> extractable soil N including microbial biomass >> plant biomass > ectomycorrhizal root tips. After quick vertical dislocation and cycling through microbial N pools, there was a rapid stabilization of leaf-litter-derived N in non-extractable N pools of the mineral soil. Very low N-15 recovery in beech seedlings suggests a high importance of other N sources such as root litter for N nutrition of beech understorey.

The relationships between plant carbon resources, soil carbon and nitrogen content, and ectomycorrhizal fungal (EMF) diversity in a monospecific, old-growth beech ( Fagus sylvatica ) forest were investigated by manipulating carbon flux by girdling. We hypothesized that disruption of the carbon supply would not affect diversity and EMF species numbers if EM fungi can be supplied by plant internal carbohydrate resources or would result in selective disappearance of EMF taxa because of differences in carbon demand of different fungi. Tree carbohydrate status, root demography, EMF colonization, and EMF taxon abundance were measured repeatedly during 1 year after girdling. Girdling did not affect root colonization but decreased EMF species richness of an estimated 79 to 90 taxa to about 40 taxa. Cenococcum geophilum , Lactarius blennius , and Tomentella lapida were dominant, colonizing about 70% of the root tips, and remained unaffected by girdling. Mainly cryptic EMF species disappeared. Therefore, the Shannon-Wiener index ( H ′) decreased but evenness was unaffected. H ′ was positively correlated with glucose, fructose, and starch concentrations of fine roots and also with the ratio of dissolved organic carbon to dissolved organic nitrogen (DOC/DON), suggesting that both H ′ and DOC/DON were governed by changes in belowground carbon allocation. Our results suggest that beech maintains numerous rare EMF species by recent photosynthate. These EM fungi may constitute biological insurance for adaptation to changing environmental conditions. The preservation of taxa previously not known to colonize beech may, thus, form an important reservoir for future forest development.

The goal of this work was to increase the understanding of factors regulating nitrogen (N) competition between roots and soil microbes. For this purpose, root assimilate supply was diminished or abolished in beech (Fagus sylvatica L.) seedlings by girdling, drought stress or a combination of both factors. This was revealed by 13C tracer abundance in root tips after 13CO2 pulse labelling of the shoots. Analysis of different root tip fractions revealed that only 6% were ectomycorrhizal. Carbon (C) allocation to ectomycorrhizal and vital non-mycorrhizal root tips was ~26% higher than to distorted root tips. Drought resulted in ~30% increased ammonium (NH4+) and amino acid concentrations in roots and ~65% increased soil NH4+ concentrations, probably because of lower consumption of NH4+ by free-living microorganisms. Root uptake of glutamine of 13 nmol g–1 fresh mass h–1 decreased 2-fold with drought, although the number of vital root tips did not decrease. Carbon content in biomass of free-living microbes increased with glucose application regardless of drought, resulting in significant depletion in soil nitrate (NO3–), root NH4+ and amino acid concentrations. Our results suggest that the root–soil system of young beech trees was C-limited, and this prevented amino acid metabolism in roots and microbial NO3– consumption in the soil, thereby exerting feedback inhibition on uptake of inorganic N by roots. We suggest that rhizodeposition is a key link in regulating the plant–microbial N balance.