SummaryAlthough 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 ≤ 2 mm) 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.

The effects of plants on the biosphere, atmosphere, and geosphere are key determinants of terrestrial ecosystem functioning. However, despite substantial progress made regarding plant belowground components, we are still only beginning to explore the complex relationships between root traits and functions. Drawing on literature in plant physiology, ecophysiology, ecology, agronomy and soil science, we review 24 aspects of plant and ecosystem functioning and their relationships with a number of traits of root systems, including aspects of architecture, physiology, morphology, anatomy, chemistry, biomechanics and biotic interactions. Based on this assessment, we critically evaluate the current strengths and gaps in our knowledge, and identify future research challenges in the field of root ecology. Most importantly, we found that below-ground traits with widest importance in plant and ecosystem Accepted Article This article is protected by copyright. All rights reserved functioning are not those most commonly measured. Also, the fair estimation of trait relative importance for functioning requires us to consider a more comprehensive range of functionally-relevant traits from a diverse range of species, across environments and over time series. We also advocate that establishing causal hierarchical links among root traits will provide a hypothesis-based framework to identify the most parsimonious sets of traits with strongest influence on the functions, and to link genotypes to plant and ecosystem functioning.

International audience* Ecosystem functioning relies heavily on below-ground processes, which are largely regulated by plant fine-roots and their functional traits. However, our knowledge of fine-root trait distribution relies to date on local- and regional-scale studies with limited numbers of species, growth forms and environmental variation. * We compiled a world-wide fine-root trait dataset, featuring 1115 species from contrasting climatic areas, phylogeny and growth forms to test a series of hypotheses pertaining to the influence of plant functional types, soil and climate variables, and the degree of manipulation of plant growing conditions on species fine-root trait variation. Most particularly, we tested the competing hypotheses that fine-root traits typical of faster return on investment would be most strongly associated with conditions of limiting versus favourable soil resource availability. We accounted for both data source and species phylogenetic relatedness. * We demonstrate that: (i) Climate conditions promoting soil fertility relate negatively to fine-root traits favouring fast soil resource acquisition, with a particularly strong positive effect of temperature on fine-root diameter and negative effect on specific root length (SRL), and a negative effect of rainfall on root nitrogen concentration; (ii) Soil bulk density strongly influences species fine-root morphology, by favouring thicker, denser fine-roots; (iii) Fine-roots from herbaceous species are on average finer and have higher SRL than those of woody species, and N2-fixing capacity positively relates to root nitrogen; and (iv) Plants growing in pots have higher SRL than those grown in the field. * Synthesis. This study reveals both the large variation in fine-root traits encountered globally and the relevance of several key plant functional types and soil and climate variables for explaining a substantial part of this variation. Climate, particularly temperature, and plant functional types were the two strongest predictors of fine-root trait variation. High trait variation occurred at local scales, suggesting that wide-ranging below-ground resource economics strategies are viable within most climatic areas and soil conditions

I.Introduction: continuing to face up to root ecology's challenges 975 II.Semantics: defining concepts for better understanding and communication 977III. Species-level vs ecosystem-level measurements 978

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Summary 1.Although plant nitrogen (N) strategies may play an important role for community structure and ecosystem functioning, there is not a clear understanding of the link between N acquisition by roots and N utilization by shoots. Particularly, it is unclear how the co-variations between size-and physiology-related traits determine N acquisition and N utilization at the plant scale. 2. We used 13 co-occurring temperate pasture grasses to study inter-specific variations in above-ground N yield and in root N acquisition and shoot N utilization traits. N acquisition traits concerned root influx capacities for and , root mass and specific root area in ingrowth cores. N utilization traits concerned leaf life span, leaf N content, leaf N resorption, mean residence time of N and leaf N use efficiency. 3. We found evidence for three trade-offs across species concerning root N acquisition: (i) root mass increased when specific root area declined; (ii) an increase in root area was observed when total N influx capacity decreased; and (iii) root influx capacity increased when capacity declined. 4. High total root uptake capacity gave rise to high leaf N content and was associated across species to low leaf N use efficiency. Tall grasses were characterized by high shoot N yield, high root biomass and high leaf N use efficiency. Physiology-related traits and size-related traits were generally found independent. 5. Our study demonstrates how size and N uptake related root traits are associated to major axes of plant specialization ((i) plant size and (ii) conservation vs. exploitation of N) which were previously identified based on shoot traits. Contrasted N strategies were segregated across species according to four combinations along these two axes.

The plant economics spectrum is increasingly recognized as a major determinant of plant species effects on terrestrial ecosystem functioning related to carbon cycling. However, the role of plant economic strategies in the effects of living root activity on soil organic carbon (SOC) dynamics through rhizodeposition remains unexplored, despite SOC being the largest terrestrial carbon pool. Using a continuous 13C‐labelling method allowing partitioning of plant and soil sources to carbon fluxes and pools, we studied here the linkages between plant economic strategies and SOC cycling processes in a ‘common garden’ greenhouse experiment. It includes a panel of 12 grassland species selected along a gradient of economic traits and belonging to three functionnal groups (C3 grasses, forbs and legumes). All species induced an acceleration of native SOC mineralization but this rhizosphere priming effect (RPE) substantially differed across species and varied eleven‐fold by the end of the experiment (from +26% to +295% relative to unplanted soil). Interspecific variation in RPE was primarily linked to plant photosynthetic activity associated to species economic strategies of light and CO2 resource acquisition and processing. Fast‐growing acquisitive species, such as legumes, featured large RPE, in relation with their high canopy photosynthesis coupled to high leaf photosynthetic capacity and large net primary productivity allocated above‐ground. This large RPE was further associated with high root metabolic activity, rhizodeposition and soil microbial activity. In contrast, fine‐root growth and economic traits related to soil resource foraging ability were poor predictors of RPE. The formation of new root‐derived SOC varied nine‐fold across species and was similarly positively related to the net primary productivity allocated above‐ground. Fast‐growing acquisitive species with a high photosynthetic activity induced a disproportionately large RPE relative to SOC formation. Synthesis. Overall, our study demonstrates that rhizodeposition is a major mechanism through which plant economic strategies of grassland species control soil carbon dynamics. Acquisitive versus conservative species were associated with high versus low rates of photosynthesis and rhizodeposition, in turn leading to fast versus slow SOC turnover. This emphasizes the importance of considering rhizosphere processes for understanding plant species effects on soil biogeochemistry.

Atmospheric carbon dioxide (CO2), global mean temperature and interannual variability in temperature and rainfall are expected to increase significantly by the end of the 21st century. To review the effects of these factors on forage quality, we carried out a meta‐analysis of climate manipulation experiments. The first notable result was a lack of effect of elevated CO2 on structural carbohydrates and digestibility. Elevated CO2 increased the total non‐structural carbohydrates of forage tissues by an average of 25% and decreased forage nitrogen (N) content by 8%. Increased legume abundance in multispecies swards can, however, maintain N concentration in the harvested biomass. There were no consistent effects of warming on contents of N, water‐soluble and structural carbohydrates, or on digestibility. We highlight the continuum in the effect of water availability, from drought to irrigation, with a curvilinear increase of forage N as water availability decreased. Digestibility increased, on average, by 7% with drought, but with strong experimental variations. The review places special emphasis on discussion of the specificities of mountain and Mediterranean grasslands, the former being limited by low temperature, the latter by drought and heat. Elevated CO2 decreased forage N content in mountain areas and in temperate plains alike. It increased N content by an average of 3% in Mediterranean areas; this could be due to shifts in vegetation communities under elevated CO2 or to a greater concentration of N in plant tissues under drought conditions. Further experiments are needed to investigate the effects of combined factors, including extreme climatic events.