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Global vegetation models use conceived relationships between functional traits to simulate ecosystem responses to environmental change. In this context, the concept of the leaf economics spectrum (LES) suggests coordinated leaf trait variation, and separates species which invest resources into short-lived leaves with a high expected energy return rate from species with longer-lived leaves and slower energy return. While it has been assumed that being fast (acquisitive) or slow (conservative) is a general feature for all organ systems, the translation of the LES into a root economics spectrum (RES) for tree species has been hitherto inconclusive. This may be partly due to the assumption that the bulk of tree fine roots have similar uptake functions as leaves, despite the heterogeneity of their environments and resources. In this study we investigated well-established functional leaf and stature traits as well as a high number of fine root traits (14 traits split by different root orders) of 13 dominant or subdominant temperate tree species of Central Europe, representing two phylogenetic groups (gymnosperms and angiosperms) and two mycorrhizal associations (arbuscular and ectomycorrhizal). We found reflected variation in leaf and lower-order root traits in some (surface areas and C:N) but not all (N content and longevity) traits central to the LES. Accordingly, the LES was not mirrored belowground. We identified significant phylogenetic signal in morphological lower-order root traits, i.e., in root tissue density, root diameter, and specific root length. By contrast, root architecture (root branching) was influenced by the mycorrhizal association type which developed independent from phylogeny of the host tree. In structural equation models we show that root branching significantly influences both belowground (direct influence on root C:N) and aboveground (indirect influences on specific leaf area and leaf longevity) traits which relate to resource investment and lifespan. We conclude that branching of lower order roots can be considered a leading root trait of the plant economics spectrum of temperate trees, since it relates to the mycorrhizal association type and belowground resource exploitation; while the dominance of the phylogenetic signal over environmental filtering makes morphological root traits less central for tree economics spectra across different environments.

Root exudation is a key plant function with a large influence on soil organic matter dynamics and plant–soil feedbacks in forest ecosystems. Yet despite its importance, the main ecological drivers of root exudation in mature forest trees remain to be identified. During two growing seasons, we analyzed the dependence of in situ collected root exudates on root morphology, soil chemistry and nutrient availability in six mature European beech (Fagus sylvatica L.) forests on a broad range of bedrock types. Root morphology was a major driver of root exudation across the nutrient availability gradient. A doubling of specific root length exponentially increased exudation rates of mature trees by c. 5-fold. Root exudation was also closely negatively related to soil pH and nitrogen (N) availability. At acidic and N-poor sites, where fungal biomass was reduced, exudation rates were c. 3-fold higher than at N- and base-richer sites and correlated negatively with the activity of enzymes degrading less bioavailable carbon (C) and N in the bulk soil. We conclude that root exudation increases on highly acidic, N-poor soils, in which fungal activity is reduced and a greater portion of the assimilated plant C is shifted to the external ecosystem C cycle.

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 (H3 33PO4 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.

Subsoils are known to harbor large amounts of soil organic carbon (SOC) and may represent key global carbon (C) sinks given appropriate management. Although rhizodeposition is a major input pathway of organic matter to subsoils, little knowledge exists on C dynamics, particularly stabilization mechanisms, such as soil aggregation, in the rhizosphere of different soil depths. The aim of this study was to investigate the influence of natural and elevated root exudation on C allocation and aggregation in the topsoil and subsoil of a mature European beech (Fagus sylvatica L.) forest. We experimentally added model root exudates to soil at two different concentrations using artificial roots and analyzed how these affect SOC, nitrogen, microbial community composition, and size distribution of water-stable aggregates. Based on the experimental data, a mathematical model was developed to describe the spatial distribution of the formation of soil aggregates and their binding strength. Our results demonstrate that greater exudate additions affect the microbial community composition in favor of fungi which promote the formation of macroaggregates. This effect was most pronounced in the C-poor subsoil, where macroaggregation increased by 86 % and SOC content by 10 %. Our modeling exercise reproduced the observed increase in subsoil SOC at high exudate additions. We conclude that elevated root exudation has the potential to increase biotic macroaggregation and thus the C sink strength in the rhizosphere of forest subsoils.

While multiple experiments have demonstrated that trees exposed to elevated CO₂can stimulate microbes to release nutrients from soil organic matter, the importance of root‐ versus mycorrhizal‐induced changes in soil processes are presently unknown. We analyzed the contribution of roots and mycorrhizal activities to carbon (C) and nitrogen (N) turnover in a loblolly pine (Pinus taeda) forest exposed to elevated CO₂by measuring extracellular enzyme activities at soil microsites accessed via root windows. Specifically, we quantified enzyme activity from soil adjacent to root tips (rhizosphere), soil adjacent to hyphal tips (hyphosphere), and bulk soil. During the peak growing season, CO₂enrichment induced a greater increase of N‐releasing enzymes in the rhizosphere (215% increase) than in the hyphosphere (36% increase), but a greater increase of recalcitrant C‐degrading enzymes in the hyphosphere (118%) than in the rhizosphere (19%). Nitrogen fertilization influenced the magnitude of CO₂effects on enzyme activities in the rhizosphere only. At the ecosystem scale, the rhizosphere accounted for c. 50% and 40% of the total activity of N‐ and C‐releasing enzymes, respectively. Collectively, our results suggest that root exudates may contribute more to accelerated N cycling under elevated CO₂at this site, while mycorrhizal fungi may contribute more to soil C degradation.

Precipitation as a key determinant of forest productivity influences forest ecosystems also indirectly through alteration of the nutrient status of the soil, but this interaction is not well understood. Along a steep precipitation gradient, we studied the consequences of reduced precipitation for the soil and biomass nutrient pools and dynamics in 14 mature European beech (Fagus sylvatica L.) forests on Triassic sandstone. We tested the hypotheses that lowered summer precipitation (1) is associated with less acid soils and (2) a reduced accumulation of organic matter on the forest floor, and (3) reduces nutrient supply from the soil and leads to decreasing foliar and root nutrient concentrations. Soil acidity, the amount of forest floor organic matter, and the associated organic matter N and P pools decreased to about a half from wet to dry sites; the C/P and N/P ratios, but not the C/N ratio, of forest floor organic matter were reduced as well. Net N mineralization and P and K pools in the mineral soil did not change with decreasing precipitation. Foliar P and K concentrations (beech sun leaves) increased while N remained constant, resulting in decreasing foliar N/P and N/K ratios. Estimated N resorption efficiency increased toward the dry sites. We conclude that a reduction in summer rainfall significantly reduces the soil C, N and P pools but does not result in decreasing foliar N and P contents in beech. However, the decreasing foliar N/P ratios towards the dry stands indicate that the importance of P limitation for tree growth declines with decreasing precipitation.

While a severe decrease in phosphorus (P) availability is already taking place in a large number of ecosystems, drought and nitrogen (N) deposition will likely further decrease the availability of P under global change. Plants have developed physiological strategies to cope with decreasing P resources, but it is unclear how these strategies respond to elevated N deposition and summer droughts. We investigated the influence of N and P availability and soil drought on P uptake (H PO feeding experiment) and use efficiencies in young Webb. trees. We hypothesized that (H1) the expected increases in soil N:P ratios will increase the efficiencies of P uptake and use of oak saplings but will decrease the efficiencies of N uptake and use, whereas (H2) drought will affect P uptake efficiency more than N uptake efficiency. In confirmation of (H1) we found that a sharp increase of the soil N:P ratio from 4 to 42 g g significantly increased the instantaneous P uptake efficiency ( PUptakeE) by five-fold and long-term P uptake efficiency (PUptakeE) by six-fold, while it decreased N uptake efficiency (NUptakeE) and N use efficiency (NUE). In contradiction to (H1), P use efficiency (PUE) did not respond to the simulated extended gradient of soil N:P ratios but remained relatively constant. (H2) was only partially confirmed as soil drought reduced PUptakeE by up to a fourth at high soil N:P ratios but had no significant effect on NUptakeE. As a consequence, increasing summer droughts may decrease the response of PUptakeE to increasing P limitation, which - in the absence of adjustments of the efficiency of P use - can aggravate growth reductions in this eastern Mediterranean tree species under global change.

Yet despite the importance of intact forest ecosystems for future generations of humankind (Costanza et al., 1997), notoriously more research funds are directed towards agricultural systems, or wood processing at most (Lovrić et al., 2020), than towards the preservation and sustainable management of forest ecosystems under global change. [...]Defrenne et al.shed light on the ectomycorrhizal fungal diversity in Pseudotsuga menziesii forests across regional gradients in Canada. [...]potential effects of shrub encroachment on groundwater recharge rates in a climate-change affected semi-arid savanna ecosystem were determined. [...]Martín-Sanz et al.investigated the effect of water availability on the resilience to forest fires in 19 Pinus halepensis provenances via absolute bark thickness (i.e. determining the degree of heat insulation).

African montane forests harbor some of the greatest biodiversity worldwide, with high levels of species endemism. However, the loss of these forests through fragmentation, deforestation and climate change has been rapidly increasing in recent years. Montane forests in Africa are more susceptible to changes in climate than their lowland counterparts, yet their ecological value is still underrepresented. These montane forests have recently been highlighted as a major aboveground carbon (C) stock. The estimated 149.4 Mg C ha −1 from aboveground live trees surpasses estimates from the Intergovernmental Panel on Climate Change for these endangered forests, and exceeds reported values for neotropical montane and lowland forests by up to 70%. Despite the tremendous implications of these findings, coordinated and available research on the C storage potential of the other share of African montane forest biomass, that is in tree roots, is largely missing. Broadly estimated from the allometry of aboveground C stocks and from root:shoot ratios in lowland forests, more than 60 Mg C ha −1 can be stored in African montane forest roots, about 40% more than previously determined. While this broad estimation points at the potential importance of root C stocks in African montane forests, it also unveils a far-reaching knowledge gap. Here, we advocate for a more quantitative representation of the root C stock from dominant forest tree species of African tropical montane forests and ultimately for a better grasp on tree C stocks from this endangered ecosystem.

AimsRoot exudation may have a large impact on soil biological activity and nutrient cycling. Recent advances in in situ-measurement techniques have enabled deeper insights into the impact of tree root exudation on rhizosphere processes, but the abiotic and biotic controls of exudation rate remain poorly understood. We explored the temperature dependence of root exudation in mature beech (Fagus sylvatica L.) trees.MethodsWe measured fine root exudation in seven beech forests along an elevational gradient (310–800 m a.s.l.) and related carbon (C)-flux rates to mean daily temperature, actual precipitation, mean summer temperature (MST) and precipitation (MAP), soil moisture (SWC), and stand structure.ResultsAverage mass-specific exudation (averaged over all sampling dates) ranged from 12.2 µg C g−1 h−1 to 21.6 µg C g−1 h−1 with lowest rates measured at highest elevations and peak rates at mid-elevation (490 m). Regression analyses showed a highly significant positive effect of site-specific daily air and soil temperature on exudation rates (p < 0.01) with an average increase by 2 µg C g−1 h−1 per 1 °C-temperature increase, while the relation to mean summer or annual temperature and mean temperature of the measuring year was less tight. Exudation decreased with increases in mean annual precipitation and soil moisture, but increased with increasing stem density.ConclusionsThe root exudation rate of beech trees roughly triples between 10 °C and 20 °C mean daily temperature, evidencing a large temperature influence on root-borne C flux to the soil.