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1. Root, stem and leaf traits are thought to be functionally coordinated to maximize the efficiency of acquiring and using limited resources. However, evidence is mixed for consistent whole-plant trait coordination among woody plants, and we lack a clear understanding of the adaptive value of root traits along soil resource gradients. If fine roots are the below-ground analogue to leaves, then low specific root length (SRL) and high tissue density should be common on infertile soil. Here, we test the prediction that root, stem and leaf traits and relative growth rate respond in unison with soil fertility gradients. 2. We measured fine root, stem and leaf traits and relative growth rate on individual seedlings of 66 tree species grown in controlled conditions. Our objectives were (i) to determine whether multiple root traits align with growth rate, leaf and stem traits and with each other and (ii) to quantify the relationships between community-weighted mean root traits and two strong soil fertility gradients that differed in spatial extent and community composition. 3. At the species level, fast growth rates were associated with low root and stem tissue density and high specific leaf area. SRL and root diameter were not clearly related to growth rate and loaded on a separate principal component from the plant economic spectmm. 4. At the community level, growth rate was positively related to soil fertility, and root tissue density (RTD) and branching were negatively related to soil fertility. SRL was negatively related and root diameter was positively related to soil fertility on the large-scale gradient that included ectomycorrhizal angiosperms. 5. Synthesis. Root, stem and leaf tissue traits of tree seedlings are coordinated and influence fitness along soil fertility gradients. RTD responds in unison with above-ground traits to soil fertility gradients; however, root traits are multidimensional because SRL is orthogonal to the plant economic spectrum. In contrast to leaves, trees are not constrained in the way they construct fine roots: plants can construct high or low SRL roots of any tissue density. High RTD is the most consistent below-ground trait that reflects adaptation to infertile soil.

Plants respond to resource stress by changing multiple aspects of their biomass allocation, morphology, physiology and architecture. To date, we lack an integrated view of the relative importance of these plastic responses in alleviating resource stress and of the consistency/variability of these responses among species. We subjected nine species (legumes, forbs and graminoids) to nitrogen and/or light shortages and measured 11 above-ground and below-ground trait adjustments critical in the alleviation of these stresses (plus several underlying traits). Nine traits out of 11 showed adjustments that improved plants’ potential capacity to acquire the limiting resource at a given time. Above ground, aspects of plasticity in allocation, morphology, physiology and architecture all appeared important in improving light capture, whereas below ground, plasticity in allocation and physiology were most critical to improving nitrogen acquisition. Six traits out of 11 showed substantial heterogeneity in species plasticity, with little structuration of these differences within trait covariation syndromes. Such comprehensive assessment of the complex nature of phenotypic responses of plants to multiple stress factors, and the comparison of plant responses across multiple species, makes a clear case for the high (but largely overlooked) diversity of potential plastic responses of plants, and for the need to explore the potential rules structuring them.

The identification of plant functional traits that can be linked to ecosystem processes is of wide interest, especially for predicting vegetational responses to climate change. Root diameter of the finest absorptive roots may be one plant trait that has wide significance. Do species with relatively thick absorptive roots forage in nutrient-rich patches differently from species with relatively fine absorptive roots? We measured traits related to nutrient foraging (root morphology and architecture, root proliferation, and mycorrhizal colonization) across six coexisting arbuscular mycorrhizal (AM) temperate tree species with and without nutrient addition. Root traits such as root diameter and specific root length were highly correlated with root branching intensity, with thin-root species having higher branching intensity than thick-root species. In both fertilized and unfertilized soil, species with thin absorptive roots and high branching intensity showed much greater root length and mass proliferation but lower mycorrhizal colonization than species with thick absorptive roots. Across all species, fertilization led to increased root proliferation and reduced mycorrhizal colonization. These results suggest that thin-root species forage more by root proliferation, whereas thick-root species forage more by mycorrhizal fungi. In mineral nutrient-rich patches, AM trees seem to forage more by proliferating roots than by mycorrhizal fungi.

Fine-root traits play key roles in ecosystem processes, but the drivers of fine-root trait diversity remain poorly understood. The plant economic spectrum (PES) hypothesis predicts that leaf and root traits evolved in coordination. Mycorrhizal association type, plant growth form and climate may also affect root traits. However, the extent to which these controls are confounded with phylogenetic structuring remains unclear. Here we compiled information about root and leaf traits for > 600 species. Using phylogenetic relatedness, climatic ranges, growth form and mycorrhizal associations, we quantified the importance of these factors in the global distribution of fine-root traits. Phylogenetic structuring accounts for most of the variation for all traits excepting root tissue density, with root diameter and nitrogen concentration showing the strongest phylogenetic signal and specific root length showing intermediate values. Climate was the second most important factor, whereas mycorrhizal type had little effect. Substantial trait coordination occurred between leaves and roots, but the strength varied between growth forms and clades. Our analyses provide evidence that the integration of roots and leaves in the PES requires better accounting of the variation in traits across phylogenetic clades. Inclusion of phylogenetic information provides a powerful framework for predictions of belowground functional traits at global scales.

Plants adapt phenotypically to different conditions of light and nutrient supply, supposedly in order to achieve colimitation of these resources. Their key variable of adjustment is the ratio of leaf area to root length, which relies on plant biomass allocation and organ morphology. We recorded phenotypic differences in leaf and root mass fractions (LMF, RMF), specific leaf area (SLA) and specific root length (SRL) of 12 herbaceous species grown in factorial combinations of high/low irradiance and fertilization treatments. Leaf area and root length ratios, and their components, were influenced by nonadditive effects between light and nutrient supply, and differences in the strength of plant responses were partly explained by Ellenberg's species values representing ecological optima. Changes in allocation were critical in plant responses to nutrient availability, as the RMF contribution to changes in root length was 2.5× that of the SRL. Contrastingly, morphological adjustments (SLA rather than LMF) made up the bulk of plant response to light availability. Our results suggest largely predictable differences in responses of species and groups of species to environmental change. Nevertheless, they stress the critical need to account for adjustments in below‐ground mass allocation to understand the assembly and responses of communities in changing 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.

Background and aims Characterising the spatial distribution of tree fine roots (diameter≤2 mm) is fundamental for a better understanding of belowground functioning when tree are grown with associated crops in agroforestry systems. Our aim was to compare fine root distributions and orientations in trees grown in an alley cropping agroforestry stand with those in a tree monoculture. Methods Fieldwork was conducted in two adjacent 17 year old hybrid walnut (Juglans regia×nigra L.) stands in southern France: the agroforestry stand was intercropped with durum wheat (Triticum turgidum L. subsp. durum) whereas the tree monoculture had a natural understorey. Root intercepts were mapped to a depth of 150 cm on trench walls in both stands, and to a depth of 400 cm in the agroforestry stand in order to characterise tree root distribution below the crop's maximum rooting depth. Soil cubes were then extracted to assess three dimensional root orientation and to establish a predictive model of root length densities (RLD) derived from root intersection densities (RID). Results In the tree monoculture, root mapping demonstrated a very high tree RID in the top 50 cm and a slight decrease in RID with increasing soil depth. However, in the agroforestry stand, RID was significantly lower at 50 cm, tree roots colonized deeper soil layers and were more vertically oriented. In the agroforestry stand, RID and RLD were greater within the tree row than in the inter-row. Conclusions Fine roots of intercropped walnut trees grew significantly deeper, indicating a strong plasticity in root distribution. This plasticity reduced direct root competition from the crop, enabling trees to access deeper water tables not available to crop roots.

• Plant biomass allocation may be optimized to acquire and conserve resources. How trade-offs in the allocation of tropical tree seedlings depend on different stressors remains poorly understood. Here we test whether above- and below-ground traits of tropical tree seedlings could explain observed occurrence along gradients of resources (light, water) and defoliation (fire, herbivory). • We grew 24 tree species occurring in five African vegetation types, varying from dry savanna to moist forest, in a glasshouse for 6 months, and measured traits associated with biomass allocation. • Classification based on above-ground traits resulted in clusters representing savanna and forest species, with low and high shoot investment, respectively. Classification based on root traits resulted in four clusters representing dry savanna, humid savanna, dry forest and moist forest, characterized by a deep mean rooting depth, root starch investment, high specific root length in deeper soil layers, and high specific root length in the top soil layer, respectively. • In conclusion, tree seedlings in this study show root trait syndromes, which vary along gradients of resources and defoliation: seedlings from dry areas invest in deep roots, seedlings from shaded environments optimize shoot investment, and seedlings experiencing frequent defoliation store resources in the roots.

Root trait variation and plasticity could be key factors differentiating plant performance under drought. However, water manipulation and root measurements are rarely coupled empirically across growth forms to identify whether belowground strategies are generalizable across species. We measured seedling root traits across three moisture levels in 18 Mediterranean forbs, grasses, and woody species. Drought increased the root mass fraction (RMF) and decreased the relative proportion of thin roots (indicated by increased root diameters and decreased specific root length (SRL)), rates of root elongation and growth, plant nitrogen uptake, and plant growth. Although responses varied across species, plasticity was not associated with growth form. Woody species differed from forbs and grasses in many traits, but herbaceous groups were similar. Across water treatments, trait correlations suggested a single spectrum of belowground trade-offs related to resource acquisition and plant growth. While effects of SRL and RMF on plant growth shifted with drought, root elongation rate consistently represented this spectrum. We demonstrate that general patterns of root morphology and plasticity are identifiable across diverse species. Root trait measurements should enhance our understanding of belowground strategy and performance across growth forms, but it will be critical to incorporate plasticity and additional aspects of root function into these efforts.

Despite the importance of fine roots for the acquisition of soil resources such as nitrogen and water, the study of linkages between traits and both population and community dynamics remains focused on aboveground traits. We address this gap by investigating associations between belowground traits and metrics of species dynamics. Our analysis included 85 species from a long-term data set on the transition from old field to forest in eastern North America (the Buell-Small Succession Study) and the new Fine-Root Ecology Database. Given the prominent roles of life form (woody vs. non-woody) and species origin (native vs. exotic) in defining functional relationships, we also assessed whether traits or their relationships with species dynamics differed for these groups. Species that reached their peak abundance early in succession had fine-root traits corresponding to resource acquisitive strategies (i.e., they were thinner, less dense, and had higher nitrogen concentrations) while species that peaked progressively later had increasingly conservative strategies. In addition to having more acquisitive root traits than native species, exotics diverged from the above successional trend, having consistently thinner fine roots regardless of the community context. Species with more acquisitive fine-root morphologies typically had faster rates of abundance increase and achieved their maximal rates in fewer years. Decreasing soil nutrient availability and increasing belowground competition may become increasingly strong filters in successional communities, acting on root traits to promote a transition from acquisitive to conservative foraging. However, disturbances that increase light and soil resource availability at local scales may allow acquisitive species, especially invasive exotics, to continue colonizing late into the community transition to forest.