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Summary Roots vary in anatomy, morphology and physiology, both spatially (different parts of the same root system) and temporally (plastic changes, root ageing), suggesting that root trait measurements are strongly affected by root sampling categories. In this context, it is urgent to clarify the functional significance of current root sampling categories (e.g. fine roots of the first order, the first three orders, ≤1 mm or ≤2 mm), establish guidelines for choosing between sampling methods and revise root ontology to account for functional differences between traits measured on distinct root categories. Here, we used a worldwide database of fine‐root traits to test the hypothesis that distinct fine‐root trait values – with link to fine‐root functions – were generally affected by different root sampling categories. We observed indeed a clear functional break between first‐order roots and roots of all three other sampling categories, and a smaller but substantial break between roots of the three first orders and the ≤2 mm category, demonstrating globally that different sampling methodologies capture different functional parts of roots. Our synthesis suggests that all current root sampling categories present both advantages and pitfalls and that no single method can appropriately tackle the main current challenge of root functional ecology: i.e. linking fine roots to plant and ecosystem functions in a truly comparable way across all plants. We argue instead that a small set of complementary standardized sampling methods is necessary to capture the linkages between root forms and functions. To assist experimenters selecting adequate sampling we developed a decision table following three logical questions: (i) what plant or ecosystem function must be addressed; (ii) what root categories are involved in this function and (iii) what traits should be measured on these root categories. Challenging, strengthening and expending such common reference framework would be a substantial step towards wider comparability of future functional trait datasets. A lay summary is available for this article. Lay Summary

Plant trait variation drives plant function, community composition, and ecosystem processes. However, our current understanding of trait variation disproportionately relies on aboveground observations. Here we integrate root traits into the global framework of plant form and function. We developed and tested an overarching conceptual framework that integrates two recently identified root trait gradients with a well-established aboveground plant trait framework. We confronted our novel framework with published relationships between above- and belowground trait analogues and with multivariate analyses of aboveground and belowground traits of 2510 species. Our traits represent the leaf- and root conservation gradients (specific leaf area, leaf and root nitrogen concentration and root tissue density), the root collaboration gradient (root diameter and specific root length), and the plant size gradient (plant height and rooting depth). We found that an integrated, whole-plant trait space required as much as four axes. The two main axes represented the fast-slow ‘conservation’ gradient on which leaf and fine-root traits were well aligned, and the ‘collaboration’ gradient in roots. The two additional axes were separate, orthogonal plant size axes for height and rooting depth. This perspective on the multi-dimensional nature of plant trait variation better encompasses plant function and influence on the surrounding environment.

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.

• Background and Aims Fine root decomposition is an important determinant of nutrient and carbon cycling in grasslands; however, little is known about the factors controlling root decomposition among species. Our aim was to investigate whether interspecific variation in the potential decomposition rate of fine roots could be accounted for by root chemical and morphological traits, life history and taxonomic affiliation. We also investigated the co-ordinated variation in root and leaf traits and potential decomposition rates. • Methods We analysed potential decomposition rates and the chemical and morphological traits of fine roots on 18 Mediterranean herbaceous species grown in controlled conditions. The results were compared with those obtained for leaves in a previous study conducted on similar species. • Key Results Differences in the potential decomposition rates of fine roots between species were accounted for by root chemical composition, but not by morphological traits. The root potential decomposition rate varied with taxonomy, but not with life history. Poaceae, with high cellulose concentration and low concentrations of soluble compounds and phosphorus, decomposed more slowly than Asteraceae and Fabaceae. Patterns of root traits, including decomposition rate, mirrored those of leaf traits, resulting in a similar species clustering. • Conclusions The highly co-ordinated variation of roots and leaves in terms of traits and potential decomposition rate suggests that changes in the functional composition of communities in response to anthropogenic changes will strongly affect biogeochemical cycles at the ecosystem level.

Summary Variation and tradeoffs within and among plant traits are increasingly being harnessed by empiricists and modelers to understand and predict ecosystem processes under changing environmental conditions. While fine roots play an important role in ecosystem functioning, fine‐root traits are underrepresented in global trait databases. This has hindered efforts to analyze fine‐root trait variation and link it with plant function and environmental conditions at a global scale. This Viewpoint addresses the need for a centralized fine‐root trait database, and introduces the Fine‐Root Ecology Database (FRED, http://roots.ornl.gov) which so far includes > 70 000 observations encompassing a broad range of root traits and also includes associated environmental data. FRED represents a critical step toward improving our understanding of below‐ground plant ecology. For example, FRED facilitates the quantification of variation in fine‐root traits across root orders, species, biomes, and environmental gradients while also providing a platform for assessments of covariation among root, leaf, and wood traits, the role of fine roots in ecosystem functioning, and the representation of fine roots in terrestrial biosphere models. Continued input of observations into FRED to fill gaps in trait coverage will improve our understanding of changes in fine‐root traits across space and time.

Although fine roots are important components of the global carbon cycle, there is limited understanding of root structure-function relationships among species. We determined whether root respiration rate and decomposability, two key processes driving carbon cycling but always studied separately, varied with root morphological and chemical traits, in a coordinated way that would demonstrate the existence of a root economics spectrum (RES). Twelve traits were measured on fine roots (diameter 2mm) of 74 species (31 graminoids and 43 herbaceous and dwarf shrub eudicots) collected in three biomes. The findings of this study support the existence of a RES representing an axis of trait variation in which root respiration was positively correlated to nitrogen concentration and specific root length and negatively correlated to the root dry matter content, lignin:nitrogen ratio and the remaining mass after decomposition. This pattern of traits was highly consistent within graminoids but less consistent within eudicots, as a result of an uncoupling between decomposability and morphology, and of heterogeneity of individual roots of eudicots within the fine-root pool. The positive relationship found between root respiration and decomposability is essential for a better understanding of vegetation-soil feedbacks and for improving terrestrial biosphere models predicting the consequences of plant community changes for carbon cycling.

Fil: Díaz, Sandra Myrna. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina

Aims Mineral-associated organic matter, mainly derived from microbial by-products, persists longer in soil compared to particulate organic matter (POM). POM is highly recalcitrant and originates largely from decomposing root and shoot litter. Theory suggests that root traits and growth dynamics should affect carbon (C) accumulation into these different pools, but the specific traits driving this accumulation are not clearly identified. Methods Twelve herbaceous species were grown for 37 weeks in monocultures. Root elongation rate (RER) was measured throughout the experiment. At the end of the experiment, we determined morphological and chemical root traits, as well as substrate induced respiration (SIR) as a proxy for microbial activity. Carbon was measured in four different soil fractions, following particle-size and density fractionation. Results Root biomass, RER, root diameter, hemicellulose content and SIR (characteristic of N 2 -fixing Fabaceae species), were all positively correlated with increased C in the coarse silt fraction. Root diameter and hemicellulose content were negatively correlated with C in the POM fraction, that was greater under non N 2 -fixing Poaceae species, characterized by lignin-rich roots with a high carbon:nitrogen ratio that grew slowly. The accumulation of C in different soil pools was mediated by microbial activity. Conclusions Our results show that root traits determine C input into different soil pools, mediated primarily by microbial activity, thus determining the fate of soil organic C. We also highlight that C in different soil pools, and not only total soil organic C, should be reported in future studies to better understand its origin, fate and dynamics.

Although the structure and composition of plant communities is known to influence the functioning of ecosystems, there is as yet no agreement as to how these should be described from a functional perspective. We tested the biomass ratio hypothesis, which postulates that ecosystem properties should depend on species traits and on species contribution to the total biomass of the community, in a successional sere following vineyard abandonment in the Mediterranean region of France. Ecosystem-specific net primary productivity, litter decomposition rate, and total soil carbon and nitrogen varied significantly with field age, and correlated with community-aggregated (i.e., weighed according to the relative abundance of species) functional leaf traits. The three easily measurable traits tested, specific leaf area, leaf dry matter content, and nitrogen concentration, provide a simple means to scale up from organ to ecosystem functioning in complex plant communities. We propose that they be called \"functional markers,\" and be used to assess the impacts of community changes on ecosystem properties induced, in particular, by global change drivers.

BACKGROUND AND AIMS: Leaf thickness plays an important role in leaf and plant functioning, and relates to a species' strategy of resource acquisition and use. As such, it has been widely used for screening purposes in crop science and community ecology. However, since its measurement is not straightforward, a number of estimates have been proposed. Here, the validity of the (SLA x LDMC)⁻¹ product is tested to estimate leaf thickness, where SLA is the specific leaf area (leaf area/dry mass) and LDMC is the leaf dry matter content (leaf dry mass/fresh mass). SLA and LDMC are two leaf traits that are both more easily measurable and often reported in the literature. METHODS: The relationship between leaf thickness (LT) and (SLA x LDMC)⁻¹ was tested in two analyses of covariance using 11 datasets (three original and eight published) for a total number of 1039 data points, corresponding to a wide range of growth forms growing in contrasted environments in four continents. KEY RESULTS AND CONCLUSIONS: The overall slope and intercept of the relationship were not significantly different from one and zero, respectively, and the residual standard error was 0·11. Only two of the eight datasets displayed a significant difference in the intercepts, and the only significant difference among the most represented growth forms was for trees. LT can therefore be estimated by (SLA x LDMC)⁻¹, allowing leaf thickness to be derived from easily and widely measured leaf traits.