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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.

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.

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.