505I.506II.506III.508IV.512V.513VI.514515References515 Summary Fine roots acquire essential soil resources and mediate biogeochemical cycling in terrestrial ecosystems. Estimates of carbon and nutrient allocation to build and maintain these structures remain uncertain because of the challenges of consistently measuring and interpreting fine‐root systems. Traditionally, fine roots have been defined as all roots ≤ 2 mm in diameter, yet it is now recognized that this approach fails to capture the diversity of form and function observed among fine‐root orders. Here, we demonstrate how order‐based and functional classification frameworks improve our understanding of dynamic root processes in ecosystems dominated by perennial plants. In these frameworks, fine roots are either separated into individual root orders or functionally defined into a shorter‐lived absorptive pool and a longer‐lived transport fine‐root pool. Using these frameworks, we estimate that fine‐root production and turnover represent 22% of terrestrial net primary production globally – a c. 30% reduction from previous estimates assuming a single fine‐root pool. Future work developing tools to rapidly differentiate functional fine‐root classes, explicit incorporation of mycorrhizal fungi into fine‐root studies, and wider adoption of a two‐pool approach to model fine roots provide opportunities to better understand below‐ground processes in the terrestrial biosphere.

Plant traits-the morphological, anatomical, physiological, biochemical and phenological characteristics of plants-determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait-based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits-almost complete coverage for 'plant growth form'. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait-environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects.We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives. Geosphere-Biosphere Program (IGBP) and DIVERSITAS, the TRY database (TRY-not an acronym, rather a statement of sentiment; https ://www.try-db.org; Kattge et al., 2011) was proposed with the explicit assignment to improve the availability and accessibility of plant trait data for ecology and earth system sciences. The Max Planck Institute for Biogeochemistry (MPI-BGC) offered to host the database and the different groups joined forces for this community-driven program. Two factors were key to the success of TRY: the support and trust of leaders in the field of functional plant ecology submitting large databases and the long-term funding by the Max Planck Society, the MPI-BGC and the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, which has enabled the continuous development of the TRY database.

Plant economics run on carbon and nutrients instead of money. Leaf strategies aboveground span an economic spectrum from “live fast and die young” to “slow and steady,” but the economy defined by root strategies belowground remains unclear. Here, we take a holistic view of the belowground economy and show that root-mycorrhizal collaboration can short circuit a one-dimensional economic spectrum, providing an entire space of economic possibilities. Root trait data from 1810 species across the globe confirm a classical fast-slow “conservation” gradient but show that most variation is explained by an orthogonal “collaboration” gradient, ranging from “do-it-yourself” resource uptake to “outsourcing” of resource uptake to mycorrhizal fungi. This broadened “root economics space” provides a solid foundation for predictive understanding of belowground responses to changing environmental conditions.

Plant roots have greatly diversified in form and function since the emergence of the first land plants, but the global organization of functional traits in roots remains poorly understood. Here we analyse a global dataset of 10 functionally important root traits in metabolically active first-order roots, collected from 369 species distributed across the natural plant communities of 7 biomes. Our results identify a high degree of organization of root traits across species and biomes, and reveal a pattern that differs from expectations based on previous studies of leaf traits. Root diameter exerts the strongest influence on root trait variation across plant species, growth forms and biomes. Our analysis suggests that plants have evolved thinner roots since they first emerged in land ecosystems, which has enabled them to markedly improve their efficiency of soil exploration per unit of carbon invested and to reduce their dependence on symbiotic mycorrhizal fungi. We also found that diversity in root morphological traits is greatest in the tropics, where plant diversity is highest and many ancestral phylogenetic groups are preserved. Diversity in root morphology declines sharply across the sequence of tropical, temperate and desert biomes, presumably owing to changes in resource supply caused by seasonally inhospitable abiotic conditions. Our results suggest that root traits have evolved along a spectrum bounded by two contrasting strategies of root life: an ancestral 'conservative' strategy in which plants with thick roots depend on symbiosis with mycorrhizal fungi for soil resources and a more-derived 'opportunistic' strategy in which thin roots enable plants to more efficiently leverage photosynthetic carbon for soil exploration. These findings imply that innovations of belowground traits have had an important role in preparing plants to colonize new habitats, and in generating biodiversity within and across biomes.

The earth's future climate state is highly dependent upon changes in terrestrial C storage in response to rising concentrations of atmospheric CO₂. Here we show that consistently enhanced rates of net primary production (NPP) are sustained by a C-cascade through the root-microbe-soil system; increases in the flux of C belowground under elevated CO₂ stimulated microbial activity, accelerated the rate of soil organic matter decomposition and stimulated tree uptake of N bound to this SOM. This process set into motion a positive feedback maintaining greater C gain under elevated CO₂ as a result of increases in canopy N content and higher photosynthetic N-use efficiency. The ecosystem-level consequence of the enhanced requirement for N and the exchange of plant C for N belowground is the dominance of C storage in tree biomass but the preclusion of a large C sink in the soil.

Summary• Although linkages of leaf and whole-plant traits to leaf lifespan have been rigorously investigated, there is a limited understanding of similar linkages of whole-plant and fine root traits to root lifespan. In comparisons across species, do suites of traits found in leaves also exist for roots, and can these traits be used to predict root lifespan?• We observed the fine root lifespan of 12 temperate tree species using minirhizotrons in a common garden and compared their median lifespans with fine-root and whole-plant traits. We then determined which set of combined traits would be most useful in predicting patterns of root lifespan.• Median root lifespan ranged widely among species (95-336 d). Root diameter, calcium content, and tree wood density were positively related to root lifespan, whereas specific root length, nitrogen (N) : carbon (C) ratio, and plant growth rate were negatively related to root lifespan. Root diameter and plant growth rate, together (R 2 = 0.62) or in combination with root N : C ratio (R 2 = 0.76), were useful predictors of root lifespan across the 12 species.• Our results highlight linkages between fine root lifespan in temperate trees and plant functional traits that may reduce uncertainty in predictions of root lifespan or turnover across species at broader spatial scales.

Plant economics run on carbon and nutrients instead of money. Leaf strategies aboveground span an economic spectrum from 'live fast and die young' to 'slow and steady', but the economy defined by root strategies belowground remains unclear. Here we take a holistic view of the belowground economy, and show that root-mycorrhizal collaboration can short circuit a one-dimensional economic spectrum, providing an entire space of economic 45 possibilities. Root trait data from 1,781 species across the globe confirm a classical fast-slow 'conservation' gradient but show that most variation is explained by an orthogonal 'collaboration' gradient, ranging from 'do-it-yourself' resource uptake to 'outsourcing' of resource uptake to mycorrhizal fungi. This broadened 'root economics space' provides a solid foundation for predictive understanding of belowground responses to changing environmental 50 conditions. One Sentence Summary: Collaboration broadens the 'root economics space' ranging from 'doit-yourself' resource acquisition to 'outsourcing' to mycorrhizal partners. 55

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.