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Whether and how warming alters functional traits of absorptive plant roots remains to be answered across the globe. Tackling this question is crucial to better understanding terrestrial responses to climate change as fine-root traits drive many ecosystem processes. We carried out a detailed synthesis of fine-root trait responses to experimental warming by performing a meta-analysis of 964 paired observations from 177 publications. Warming increased fine-root biomass, production, respiration and nitrogen concentration as well as decreased root carbon : nitrogen ratio and nonstructural carbohydrates. Warming effects on fine-root biomass decreased with greater warming magnitude, especially in short-term experiments. Furthermore, the positive effect of warming on fine-root biomass was strongest in deeper soil horizons and in colder and drier regions. Total fine-root length, morphology, mortality, life span and turnover were unresponsive to warming. Our results highlight the significant changes in fine-root traits in response to warming as well as the importance of warming magnitude and duration in understanding fine-root responses. These changes have strong implications for global soil carbon stocks in a warmer world associated with increased root-derived carbon inputs into deeper soil horizons and increases in fine-root respiration.

Rock fragments are widespread in soil profiles. Despite direct effects of rock fragment content (RFC) on vegetation and soil properties, how plants respond to variations in RFC remains poorly understood. In this work, we investigated responses of two contrasting xerophytic species to varying RFC. Root biomass allocation, vertical distribution and above-ground growth were measured in Artemisia vestita and Bauhinia brachycarpa after 2 years of growth in an experiment with four levels of RFC (0, 25, 50 and 75% ν ν−1). The responses of above-ground growth and total biomass of both species showed a unimodal curve with values increasing up to intermediate RFC (25% and 50%) and then declining. Both species increased relative biomass allocation to roots at the highest RFC level (75%). A. vestita had a shallow rooting profile and greater declines in plant growth with high RFC compared with B. brachycarpa which had a deeper rooting profile. We found that intermediate RFCs were beneficial for growth of both species and both species increased root-to-shoot ratios to compensate for high RFC. The higher overall root fraction and deeper rooting profile may make B. brachycarpa more suitable than A. vestita for areas with high RFC, enabling greater extraction of increasingly limited soil resources.

Vertical fine-root distribution determines the potential for acquisition of resources throughout soil profiles; yet, variation among forest types and changes in vertical distribution with environments are poorly understood. We examined vertical root distributions of different forest communities to determine how belowground strategies shift across different forest types and along edaphic gradients. Specific root length and diameter of fine roots as well as fine-root biomass, length and area densities were measured in sequential soil layers at 10 cm depth increments across 118 forest plots representing five subalpine forest types. Evergreen forest types, including evergreen oaks, were more deeply rooted than birch forests. Differences in rooting depth were due to the dominant tree species identity, not to variations in shrub or herbaceous components. Within forest types, soil nutrients and physical properties contributed to shifts rooting depth but not root morphology. Vertical distributions of fine roots represent critical inputs of plant carbon into soils as well as different capacities for the acquisition of soil resources. Our findings identify consistent patterns of rooting distributions among forest types that may be predictable based on more easily measured root and soil properties and can improve efforts to model rooting depth profiles in forest communities.

Convergent patterns in morphological and genetic traits of mycorrhizas have been well-documented and reflect common selection forces that define mycorrhizas. However, generalizable patterns of mycorrhiza-associated chemical alterations, which are immediately linked to plant and fungal strategies for successful symbiosis, have yet to be emerged. Comparing root metabolomes of phylogenetically-diverse plants inoculated by mycorrhizal fungi across two major lifestyles (arbuscular- vs. ecto-mycorrhizas), our study uncovers metabolite changes unique to each mycorrhizal lifestyle and those common across plant-mycorrhizal combinations irrespective of lifestyles. Arbuscular and ecto- mycorrhizal colonized roots accumulated different sets of carbohydrates, indicating unique carbon partitioning strategies: particularly, arbuscular mycorrhizal roots accumulated cyclic polyols inaccessible for symbionts, suggesting tighter regulation of plants in carbon partitioning. Mycorrhizas also altered specialized metabolism, featuring frequent increases of flavan-3-ols and decreases of flavanols irrespective of mycorrhizal lifestyles, suggesting tactical reconfiguration of specialized metabolites to facilitate/contain symbiosis. Our data show for the first time, to our knowledge, that part of the root metabolite alterations by mycorrhizas were relatively common across plant-mycorrhizal systems, highlighting their potentially critical regulatory and evolutionary role for successful symbiosis. This commonality appears robust to phylogenetic diversity of host plants and thus may be widespread in land plants. Our findings offer future research venues to elucidate the finer roles of these common traits of mycorrhiza-associated metabolite alterations and thus help to eventually develop a comprehensive understanding of this omnipresent plant-fungus partnership.

International audience; 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.

Absorptive roots and arbuscular mycorrhizal fungi (AMF) constitute two related pathways for plant nutrient acquisition. However, if and how soil nutrient availability can regulate the integrated pattern of roots and AMF remain unclear. We analyzed the abundance and morphology of absorptive roots and AMF as well as the intraradical AMF community composition in response to four-year nitrogen (N) and phosphorus (P) additions in a Chinese fir (Cunninghamia lanceolata) plantation. Absorptive root biomass, length density and tissue density significantly decreased, and specific root length significantly increased with P addition. However, none of these root characteristics were significantly affected by N addition. In contrast, extraradical hyphal length density in the rhizosphere soil and the ratio of hyphal length density to mycorrhizal colonization rate significantly decreased with N addition, but remained unchanged under P addition while extraradical hyphal length density in the ingrowth mesh bags significantly decreased with both N and P additions. The relative abundance of Acaulosporaceae and Gigasporaceae increased and that of Glomeraceae decreased with P, while N addition did not significantly shift AMF community composition. Our findings indicate that the responses of absorptive roots and AMF to N addition in the Chinese fir plantation differ from their responses to P addition, suggesting diverse strategies for a single tree species to adjust to multiple soil nutrient conditions.

21 Pág. Instituto de Ciencias Forestales (ICIFOR), Ecological theory is built on trade-offs, where trait differences among species evolved as adaptations to different environments. Trade-offs are often assumed to be bidirectional, where opposite ends of a gradient in trait values confer advantages in different environments. However, unidirectional benefits could be widespread if extreme trait values confer advantages at one end of an environmental gradient, whereas a wide range of trait values are equally beneficial at the other end. Here, we show that root traits explain species occurrences along broad gradients of temperature and water availability, but model predictions only resembled trade-offs in two out of 24 models. Forest species with low specific root length and high root tissue density (RTD) were more likely to occur in warm climates but species with high specific root length and low RTD were more likely to occur in cold climates. Unidirectional benefits were more prevalent than trade-offs: for example, species with large-diameter roots and high RTD were more commonly associated with dry climates, but species with the opposite trait values were not associated with wet climates. Directional selection for traits consistently occurred in cold or dry climates, whereas a diversity of root trait values were equally viable in warm or wet climates. Explicit integration of unidirectional benefits into ecological theory is needed to advance our understanding of the consequences of trait variation on species responses to environmental change., sPlot was initiated by sDiv and funded by the German Research Foundation (FZT 118) and is now a platform of iDiv. The sRoot workshops and L.M. were also supported by NWO-Vidi grant 864.14.006. C.M.I. and the Fine-Root Ecology Database were supported by the Biological and Environmental Research program in the US Department of Energy’s Office of Science. J.B. was supported by Deutsche Forschungsgemeinschaft (DFG) project 432975993. N.R.G.-R. thanks the Dorothea Schlözer Postdoctoral Programme of the Georg-August-Universität.

Harsh environmental conditions affect both leaf structure and root traits. However, shoot growth in high-latitude systems is predominately under photoperiod control while root growth may occur for as long as thermal conditions are favorable. The different sensitivities of these organs may alter functional relationships above- and belowground along environmental gradients. We examined the relationship between absorptive root and foliar traits of Scots pine trees growing in situ along a temperate-boreal transect and in trees grown in a long-term common garden at a temperate latitude. We related changes in foliar nitrogen, phosphorus, specific leaf area, needle mass and 13 C signatures to geographic trends in absorptive root biomass to better understand patterns of altered tree nutrition and water balance. Increased allocation to absorptive fine roots was associated with greater uptake of soil nutrients and subsequently higher needle nutrient contents in the northern provenances compared with more southern provenances when grown together in a common garden setting. In contrast, the leaf δ13 C in northern and southern provenances were similar within the common garden suggesting that higher absorptive root biomass fractions could not adequately increase water supply in warmer climates. These results highlight the importance of allocation within the fine-root system and its impacts on needle nutrition while also suggesting increasing stomatal limitation of photosynthesis in the context of anticipated climatic changes.

International audience; 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 the literature in plant physiology, ecophysiology, ecology, agronomy and soil science, we reviewed 24 aspects of plant and ecosystem functioning and their relationships with a number of root system traits, including aspects of architecture, physiology, morphology, anatomy, chemistry, biomechanics and biotic interactions. Based on this assessment, we critically evaluated the current strengths and gaps in our knowledge, and identify future research challenges in the field of root ecology. Most importantly, we found that belowground traits with the broadest importance in plant and ecosystem functioning are not those most commonly measured. Also, the 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 the strongest links on functions, and to link genotypes to plant and ecosystem functioning.

In the context of a recent massive increase in research on plant root functions and their impact on the environment, root ecologists currently face many important challenges to keep on generating cutting-edge, meaningful and integrated knowledge. Consideration of the below-ground components in plant and ecosystem studies has been consistently called for in recent decades, but methodology is disparate and sometimes inappropriate. This handbook, based on the collective effort of a large team of experts, will improve trait comparisons across studies and integration of information across databases by providing standardised methods and controlled vocabularies. It is meant to be used not only as starting point by students and scientists who desire working on below-ground ecosystems, but also by experts for consolidating and broadening their views on multiple aspects of root ecology. Beyond the classical compilation of measurement protocols, we have synthesised recommendations from the literature to provide key background knowledge useful for: (1) defining below-ground plant entities and giving keys for their meaningful dissection, classification and naming beyond the classical fine-root vs coarse-root approach; (2) considering the specificity of root research to produce sound laboratory and field data; (3) describing typical, but overlooked steps for studying roots (e.g. root handling, cleaning and storage); and (4) gathering metadata necessary for the interpretation of results and their reuse. Most importantly, all root traits have been introduced with some degree of ecological context that will be a foundation for understanding their ecological meaning, their typical use and uncertainties, and some methodological and conceptual perspectives for future research. Considering all of this, we urge readers not to solely extract protocol recommendations for trait measurements from this work, but to take a moment to read and reflect on the extensive information contained in this broader guide to root ecology, including sections I–VII and the many introductions to each section and root trait description. Finally, it is critical to understand that a major aim of this guide is to help break down barriers between the many subdisciplines of root ecology and ecophysiology, broaden researchers’ views on the multiple aspects of root study and create favourable conditions for the inception of comprehensive experiments on the role of roots in plant and ecosystem functioning. ISSN:0028-646X ISSN:1469-8137

Root systems show a tremendous diversity both between and within species, suggesting a large variability in plant functioning and effects on ecosystem properties and processes. In recent decades, developments in many areas of root research have brought considerable advances in our understanding of root traits and their contribution to plant and ecosystem functioning. However, despite major progress, a comprehensive overview—bridging research fields—is lacking. Furthermore, considerable uncertainties exist in the identification of root entities, and the selection and standardized measurement of traits. Here, we provide a comprehensive overview on root entities, exemplify recent advances in our understanding of both theoretical and demonstrated relationships between root traits and plant or ecosystem functioning, discuss trait-trait relationships and hierarchies among traits, and critically assess current strengths and gaps in our knowledge.

[Motivation]: Trait data are fundamental to the quantitative description of plant form and function. Although root traits capture key dimensions related to plant responses to changing environmental conditions and effects on ecosystem processes, they have rarely been included in large-scale comparative studies and global models. For instance, root traits remain absent from nearly all studies that define the global spectrum of plant form and function. Thus, to overcome conceptual and methodological roadblocks preventing a widespread integration of root trait data into large-scale analyses we created the Global Root Trait (GRooT) Database. GRooT provides ready to use data by combining the expertise of root ecologists with data mobilization and curation. Specifically, we (a) determined a set of core root traits relevant to the description of plant form and function based on an assessment by experts, (b) maximized species coverage through data standardization within and among traits, and (c) implemented data quality checks., [Main types of variables contained]: GRooT contains 114,222 trait records on 38 continuous root traits.Spatial location and grain: Global coverage with data from arid, continental, polar, temperate and tropical biomes. Data on root traits were derived from experimental studies and field studies., [Time period and grain]: Data were recorded between 1911 and 2019., [Major taxa and level of measurement]: GRooT includes root trait data for which taxonomic information is available. Trait records vary in their taxonomic resolution, with subspecies or varieties being the highest and genera the lowest taxonomic resolution available. It contains information for 184 subspecies or varieties, 6,214 species, 1,967 genera and 254 families. Owing to variation in data sources, trait records in the database include both individual observations and mean values., [Software format]: GRooT includes two csv files. A GitHub repository contains the csv files and a script in R to query the database., The study has been supported by the TRY initiative on plant traits (http://www.try-db.org). The TRY initiative and database are hosted, developed and maintained at the Max Planck Institute for Biogeochemistry, Jena, Germany. TRY is currently supported by DIVERSITAS/Future Earth and the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig. Open access funding enabled and organized by Projekt DEAL.

Collaboration broadens the “root economics space” ranging from “do-it-yourself” to “outsourcing” to mycorrhizal partners., 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.

Fine roots mediate below-ground resource acquisition, yet understanding of how fine-root functional traits vary along environmental gradients, within branching orders and across phylogenetic scales remains limited. Morphological and architectural fine-root traits were measured on individual root orders of 20 oak species (genus Quercus) from divergent climates of origin that were harvested after three growing seasons in a glasshouse. These were then compared with similar measurements obtained from a phylogenetically diverse dataset of woody species from the Fine-Root Ecology Database (FRED). For the oaks, only precipitation seasonality and growing season moisture availability were correlated to aspects of root diameter and branching. Strong correlations among root diameters and architecture of different branch orders were common, while correlations between diameter and length were weakly negative. By contrast, the FRED dataset showed strong positive correlations between diameter and length and fewer correlations between root diameter and architectural traits. Our findings suggest that seasonal patterns of water availability are more important drivers of root adaptation in oaks than annual averages in precipitation and temperature. Furthermore, contrasting patterns of trait relationships between the oak and FRED datasets suggest that branching patterns are differentially constrained at narrow vs broad phylogenetic scales.

International audience; Background and aims: Plant phenology is a sensitive indicator of plant response to climate change. Observations of phenological events belowground for most ecosystems are difficult to obtain and very little is known about the relationship between tree shoot and root phenology. We examined the influence of environmental factors on fine root production and mortality in relation with shoot phenology in hybrid walnut trees (Juglans sp.) growing in three different climates (oceanic, continental and Mediterranean) along a latitudinal gradient in France.Methods : Eight rhizotrons were installed at each site for 21 months to monitor tree root dynamics. Root elongation rate (RER), root initiation quantity (RIQ) and root mortality quantity (RMQ) were recorded frequently using a scanner and time-lapse camera. Leaf phenology and stem radial growth were also measured. Fine roots were classified by topological order and 0–1 mm, 1–2 mm and 2–5 mm diameter classes and fine root longevity and risk of mortality were calculated during different periods over the year.Results : Root growth was not synchronous with leaf phenology in any climate or either year, but was synchronous with stem growth during the late growing season. A distinct bimodal pattern of root growth was observed during the aerial growing season. Mean RER was driven by soil temperature measured in the month preceding root growth in the oceanic climate site only. However, mean RER was significantly correlated with mean soil water potential measured in the month preceding root growth at both Mediterranean (positive relationship) and oceanic (negative relationship) sites. Mean RIQ was significantly higher at both continental and Mediterranean sites compared to the oceanic site. Soil temperature was a driver of mean RIQ during the late growing season at continental and Mediterranean sites only. Mean RMQ increased significantly with decreasing soil water potential during the late aerial growing season at the continental site only. Mean root longevity at the continental site was significantly greater than for roots at the oceanic and Mediterranean sites. Roots in the 0–1 mm and 1–2 mm diameter classes lived for significantly shorter periods compared to those in the 2–5 mm diameter class. First order roots (i.e. the primary or parents roots) lived longer than lateral branch roots at the Mediterranean site only and first order roots in the 0–1 mm diameter class had 44.5% less risk of mortality than that of lateral roots for the same class of diameter.Conclusions: We conclude that factors driving root RER were not the same between climates. Soil temperature was the best predictor of root initiation at continental and Mediterranean sites only, but drivers of root mortality remained largely undetermined

Ectomycorrhizal fungi are important in many forest ecosystems, yet their production dynamics and responses to environmental changes are poorly understood. Cenococcum geophilum is a common ectomycorrhizal fungus important to plant and forest soil biogeochemical cycles. The seasonal and inter-annual patterns of production and persistence of mycorrhizas formed by C. geophilum in a pine forest exposed to elevated atmospheric CO2 and nitrogen fertilization were monitored using a 12 y minirhizotron dataset. Production of C. geophilum mycorrhizas was distinctly seasonal and peaked in late summer/autumn. Elevated CO2 generally increased production while nitrogen fertilization strongly decreased production. Persistence times of C. geophilum mycorrhizas was ca. 2.7 y and was unaffected by CO2 and nitrogen addition. Total production was greater in shallow soil (0–16 cm) but persistence was longer in deeper soil (17–32 cm). These observations provide insights into the autecology of C. geophilum and suggest that its tissues may be slow to decompose compared to other ectomycorrhizal species.

Vegetation impacts on ecosystem functioning are mediated by mycorrhizas, plant–fungal associations formed by most plant species. Ecosystems dominated by distinct mycorrhizal types differ strongly in their biogeochemistry. Quantitative analyses of mycorrhizal impacts on ecosystem functioning are hindered by the scarcity of information on mycorrhizal distributions. Here we present global, high-resolution maps of vegetation biomass distribution by dominant mycorrhizal associations. Arbuscular, ectomycorrhizal, and ericoid mycorrhizal vegetation store, respectively, 241 ± 15, 100 ± 17, and 7 ± 1.8 GT carbon in aboveground biomass, whereas non-mycorrhizal vegetation stores 29 ± 5.5 GT carbon. Soil carbon stocks in both topsoil and subsoil are positively related to the community-level biomass fraction of ectomycorrhizal plants, though the strength of this relationship varies across biomes. We show that human-induced transformations of Earth’s ecosystems have reduced ectomycorrhizal vegetation, with potential ramifications to terrestrial carbon stocks. Our work provides a benchmark for spatially explicit and globally quantitative assessments of mycorrhizal impacts on ecosystem functioning and biogeochemical cycling., Mycorrhizas—mutualistic relationships formed between fungi and most plant species—are functionally linked to soil carbon stocks. Here the authors map the global distribution of mycorrhizal plants and quantify links between mycorrhizal vegetation patterns and terrestrial carbon stocks.

Variation in resource acquisition strategies enables plants to adapt to different environments and may partly determine their responses to climate change. However, little is known about how belowground plant traits vary across climate and soil gradients. Focusing on interior Douglas-fir (Pseudotsuga menziesii var. glauca) in western Canada, we tested whether fine-root traits relate to the environment at the intraspecific level. We quantified the variation in commonly measured functional root traits (morphological, chemical, and architectural traits) among the first three fine-root orders (i.e., absorptive fine roots) and across biogeographic gradients in climate and soil factors. Moderate but consistent trait-environment linkages occurred across populations of Douglas-fir, despite high levels of within-site variation. Shifts in morphological traits across regions were decoupled from those in chemical traits. Fine roots in colder/drier climates were characterized by a lower tissue density, higher specific area, larger diameter, and lower carbon-to-nitrogen ratio than those in warmer/wetter climates. Our results showed that Douglas-fir fine roots do not rely on adjustments in architectural traits to adapt rooting strategies in different environments. Intraspecific fine-root adjustments at the regional scale do not fit along a single axis of root economic strategy and are concordant with an increase in root acquisitive potential in colder/drier environments.

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.

Fine-root traits mediate the capacity of plants to acquire soil resources in different environments. This study aimed to examine the changes of fine-root traits when roots proliferate into the litter layer vs. mineral soils, and to determine fine-root trait plasticity of these roots in response to nitrogen (N) addition. A one-year N addition experiment was conducted in a 22-year-old broadleaf Mytilaria laosensis (Hamamelidaceae) plantation in subtropical China. Newly produced fine roots were collected monthly from the litter layer and upper mineral soil (0–10 cm) layer to measure root morphological traits and nutrient concentrations. Fine-root production was determined using ingrowth mesh screens in the litter layer. Fine-root production in the litter layer in the Mytilaria laosensis plantation was 2.6 g m−2 yr.−1 but increased 3- to 5-fold with N addition. Significant differences in fine-root morphological traits and nutrient concentrations were found between the litter layer and 0–10 cm mineral soil layer. Fine roots in the litter layer were thinner, with higher specific root length (SRL), higher specific root area (SRA), a higher proportion of fine-root biomass in lower, more absorptive root orders, and lower root tissue density (RTD) than those in 0–10 cm mineral soil layer. Higher carbon (C), N, phosphorus (P), potassium (K), calcium (Ca), magnesium (Mg) concentrations and lower C:N ratio (C/N) were also observed in fine roots in the litter layer, compared to the 0–10 cm mineral soil layer. Nitrogen addition significantly increased root P, K, and Ca concentrations, but had no effect on Mg concentration. Nitrogen addition did not affect most fine-root morphological traits but did result in decreased root diameter. Compared with the mineral soil, roots produced in the litter layer generally reflected a more absorptive strategy with smaller root diameter and lower RTD and with higher SRL, SRA, and nutrient concentrations which together are generally associated with more metabolically active, but shorter lived roots. Strong responses of fine-root production and nutrient concentrations to N addition also suggest that N may be a driving factor for fine-root growth into the litter layer. Further studies are required to identify the effect of fine-root growth into the litter layer on microbial activity.

Plant functional traits may be altered as plants adapt to various environmental constraints. Cold, low fertility growing conditions are often associated with root adjustments to increase acquisition of limiting nutrient resources, but they may also result in construction of roots with reduced uptake potential but higher tissue persistence. It is ultimately unclear whether plants produce fine roots of different structure in response to decreasing temperatures and whether these changes represent a trade-off between root function or potential root persistence. We assessed patterns of root construction based on various root morphological, biochemical and defense traits including root diameter, specific root length (SRL), root tissue density (RTD), C:N ratio, phenolic compounds, and number of phellem layers across up to 10 root orders in diverse populations of Scots pine along a 2000-km climatic gradient in Europe. Our results showed that different root traits are related to mean annual temperature (MAT) and expressed a pattern of higher root diameter and lower SRL and RTD in northern sites with lower MAT. Among absorptive roots, we observed a gradual decline in chemical defenses (phenolic compounds) with decreasing MAT. In contrast, decreasing MAT resulted in an increase of structural protection (number of phellem layers) in transport fine roots. This indicated that absorptive roots with high capacity for nutrient uptake, and transport roots with low uptake capacity, were characterized by distinct and contrasting trade-offs. Our observations suggest that diminishing structural and chemical investments into the more distal, absorptive roots in colder climates is consistent with building roots of higher absorptive capacity. At the same time, roots that play a more prominent role in transport of nutrients and water within the root system saw an increase in structural investment, which can increase persistence and reduce long-term costs associated with their frequent replacement.

Compared with ectomycorrhizal (ECM) forests, arbuscular mycorrhizal (AM) forests are hypothesized to have higher carbon (C) cycling rates and a more open nitrogen (N) cycle. To test this hypothesis, we synthesized 645 observations, including 22 variables related to below-ground C and N dynamics from 100 sites, where AM and ECM forests co-occurred at the same site. Leaf litter quality was lower in ECM than in AM trees, leading to greater forest floor C stocks in ECM forests. By contrast, AM forests had significantly higher mineral soil C concentrations, and this result was strongly mediated by plant traits and climate. No significant differences were found between AM and ECM forests in C fluxes and labile C concentrations. Furthermore, inorganic N concentrations, net N mineralization and nitrification rates were all higher in AM than in ECM forests, indicating 'mineral' N economy in AM but 'organic' N economy in ECM trees. AM and ECM forests show systematic differences in mineral vs organic N cycling, and thus mycorrhizal type may be useful in predicting how different tree species respond to multiple environmental change factors. By contrast, mycorrhizal type alone cannot reliably predict below-ground C dynamics without considering plant traits and climate.

Patterns of plant biomass allocation and functional adjustments along climatic gradients are poorly understood, particularly belowground. Generally, low temperatures suppress nutrient release and uptake, and forests under such conditions have a greater proportion of their biomass in roots. However, it is not clear whether 'more roots' means better capacity to acquire soil resources. Herein we quantified patterns of fine-root anatomy and their biomass distribution across Scots pine (Pinus sylvestris) populations both along a 2000-km latitudinal gradient and within a common garden experiment with a similar range of populations. We found that with decreasing mean temperature, a greater percentage of Scots pine root biomass was allocated to roots with higher potential absorptive capacity. Similar results were seen in the common experimental site, where cold-adapted populations produced roots with greater absorptive capacity than populations originating from warmer climates. These results demonstrate that plants growing in or originated from colder climates have more acquisitive roots, a trait that is likely adaptive in the face of the low resource availability typical of cold soils.

Plant phenology is one of the strongest indicators of ecological responses to climate change, and altered phenology can have pronounced effects on net primary production, species composition in local communities, greenhouse gas fluxes, and ecosystem processes. Although many studies have shown that aboveground plant phenology advances with warmer temperatures, demonstration of a comparable association for belowground phenology has been lacking because the factors that influence root phenology are poorly understood. Because roots can constitute a large fraction of plant biomass, and root phenology may not respond to warming in the same way as shoots, this represents an important knowledge gap in our understanding of how climate change will influence phenology and plant performance. We review studies of root phenology and provide suggestions to direct future research. Only 29% of examined studies approached root phenology quantitatively, strongly limiting interpretation of results across studies. Therefore, we suggest that researchers emphasize quantitative analyses in future phenological studies. We suggest that root initiation, peak growth, and root cessation may be under different controls. Root initiation and cessation may be more constrained by soil temperature and the timing of carbon availability, whereas the timing of peak root growth may represent trade-offs among competing plant sinks. Roots probably do not experience winter dormancy in the same way as shoots: 89% of the studies that examined winter phenology found evidence of growth during winter months. More research is needed to observe root phenology, and future studies should be careful to capture winter and early season phenology. This should be done quantitatively, with direct observations of root growth utilizing rhizotrons or minirhizotrons.

The turnover of ectomycorrhizal (EM) fungal biomass represents a significant input into forest carbon (C) and nutrient cycles. Given the size of these fluxes, understanding the factors that control the decomposition of this necromass will greatly improve understanding of C and nutrient cycling in ecosystems. Recent research has highlighted the considerable variation in the decomposition rates of EM fungal necromass, and patterns from this research are beginning to emerge. In this article we review the research that has examined both intrinsic and extrinsic factors that control the decomposition of EM fungal necromass and propose additional factors that may strongly influence EM fungal necromass decomposition and ecosystem properties. We argue that, as with most plant litters, the stoichiometry (C:N) of EM necromass is an important factor governing decomposition, but its role is modulated by the nature of the C and N in the tissue. In particular, melanin concentration appears to negatively influence the quality of EM fungal necromass much as lignin does in plant litters. Other intrinsic factors such as the morphology of the mycelium may also play a large role and suggest this as a focus for future research. Extrinsic factors, such as decomposer community activity and physical protection by soil, are also likely to be important in governing the decomposition of ectomycorrhizal necromass in situ. Finally, we highlight the potential importance of EM fungal necromass diversity and abundance in influencing terrestrial biogeochemical cycles. Understanding the factors that control the decomposition of EM necromass will then improve the predictive power of next-generation terrestrial biosphere models.

Understanding the effects of global change in terrestrial communities requires an understanding of how limiting resources interact with plant traits to affect productivity. Here, we focus on nitrogen and ask whether plant community nitrogen uptake rate is determined (a) by nitrogen availability alone or (b) by the product of nitrogen availability and fine-root mass. Surprisingly, this is not empirically resolved. We performed controlled microcosm experiments and reanalyzed published pot experiments and field data to determine the relationship between community-level nitrogen uptake rate, nitrogen availability, and fine-root mass for 46 unique combinations of species, nitrogen levels, and growing conditions. We found that plant community nitrogen uptake rate was unaffected by fine-root mass in 63% of cases and saturated with fine-root mass in 29% of cases (92% in total). In contrast, plant community nitrogen uptake rate was clearly affected by nitrogen availability. The results support the idea that although plants may over-proliferate fine roots for individual-level competition, it comes without an increase in community-level nitrogen uptake. The results have implications for the mechanisms included in coupled carbon-nitrogen terrestrial biosphere models (CN-TBMs) and are consistent with CN-TBMs that operate above the individual scale and omit fine-root mass in equations of nitrogen uptake rate but inconsistent with the majority of CN-TBMs, which operate above the individual scale and include fine-root mass in equations of nitrogen uptake rate. For the much smaller number of CN-TBMs that explicitly model individual-based belowground competition for nitrogen, the results suggest that the relative (not absolute) fine-root mass of competing individuals should be included in the equations that determine individual-level nitrogen uptake rates. By providing empirical data to support the assumptions used in CN-TBMs, we put their global climate change predictions on firmer ground.

Leaf economics and hydraulic traits are critical to leaf photosynthesis, yet it is debated whether these two sets of traits vary in a fully coordinated manner or there is room for independent variation. Here, we tested the relationship between leaf economics traits, including leaf nitrogen concentration and leaf dry mass per area, and leaf hydraulic traits including stomatal density and vein density in five tropical-subtropical forests. Surprisingly, these two suites of traits were statistically decoupled. This decoupling suggests that independent trait dimensions exist within a leaf, with leaf economics dimension corresponding to light capture and tissue longevity, and the hydraulic dimension to water-use and leaf temperature maintenance. Clearly, leaf economics and hydraulic traits can vary independently, thus allowing for more possible plant trait combinations. Compared with a single trait dimension, multiple trait dimensions may better enable species adaptations to multifarious niche dimensions, promote diverse plant strategies and facilitate species coexistence.

Arbuscular mycorrhizal fungi (AMF) mediate plant interspecific competition and community structure. However, the magnitude and direction of AMF effects and underlying mechanisms are not clear. Here, we synthesized the results of 304 studies to evaluate how AMF affect plant competition and community structure and which abiotic and biotic conditions in experimental design modify these AMF effects. The magnitude and direction of AMF effects on plant competitive ability (in terms of competitive response) differed markedly among plant functional groups. When AMF inoculum was added, competitive ability was strongly enhanced in N-fixing forbs and was significantly suppressed in C-3 grasses, whereas no effect was observed in C-4 grasses, non-N-fixing forbs and woody species. Furthermore, AMF inoculation increased competitive ability of perennial species when their competitors were annual species. AMF inoculation differentially influenced separate aspects of plant community structure and species composition. AMF inoculation significantly increased plant diversity but had no effects on plant productivity. Response of dominant plant species to AMF inoculation was the determining factor in explaining variations in how and to what degree plant diversity was influenced by AMF inoculation. When dominant species derived strong benefits from AMF, their dominance level was increased by AMF inoculation, which consequently decreased plant diversity. We did not find stronger AMF effects on plant diversity and productivity when greater numbers of AMF species were used in the inoculation.Synthesis. Despite large variations in AMF effects among studies, a unifying mechanism was observed that the mycorrhizal responsiveness (differences in plant growth between AMF and non-AMF colonization treatments) of target and neighbouring plant species can determine AMF effects on the competitive outcome among plant species, which in turn influenced plant species diversity and community composition. Given that plant traits, soil nutrient conditions and probably mycorrhizal fungal traits are all factors determining the degree of mycorrhizal response of plant species, future studies should explicitly consider each of these factors in experimental design to better understand AMF effects on plant coexistence, plant community dynamics and ecosystem processes.

Patterns of root traits among different root orders and their variation across seasons are of considerable importance for soil resource acquisition and partitioning in forest ecosystems. We evaluated whether morphological, anatomical and biochemical traits varied among root orders of Quercus robur (L.) sampled across spring, summer and fall seasons and growing in two different soil types with contrasting site fertility. We found no consistent differences in root diameter and specific root length in relation to soil type or growing season. There was, however, a strong seasonal variation in patterns of nitrogen (N) concentration among root orders. During spring and summer, N concentration was highest in the most distal, absorptive portion of the root system. At the end of the growing season, we observed a sharp decline in the N concentration of these lower-order, absorptive roots and an increase in N concentration of the higher-order, transport roots. The specific mechanisms driving the seasonally changing N concentration remain unclear but are likely related to different functions of lower-order roots for absorption and higher-order roots for structure and storage. Future work should identify how common the observed seasonal changes in N concentration are across species and determine what specific environmental cues plants or roots use to trigger shifts in resource allocation within the root branching hierarchy.

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.

Background and Aims Root mechanical traits, including tensile strength (T-r), tensile strain (epsilon(r)) and modulus of elasticity (E-r), are key functional traits that help characterize plant anchorage and the physical contribution of vegetation to landslides and erosion. The variability in these traits is high among tree fine roots and is poorly understood. Here, we explore the variation in root mechanical traits as well as their underlying links with morphological (diameter), architectural (topological order) and anatomical (stele and cortex sizes) traits. Methods We investigated the four tropical tree species Pometia tomentosa, Barringtonia fusicarpa, Baccaurea ramiflora and Pittosporopsis kerrii in Xishuangbanna, Yunnan, China. For each species, we excavated intact, fresh, fine roots and measured mechanical and anatomical traits for each branching order. Key Results Mechanical traits varied enormously among the four species within a narrow range of diameters (

Plant roots have greatly diversified in form and function since the emergence of the first land plants1,2, but the global organization of functional traits in roots remains poorly understood3,4. 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 studies5,6 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.

Functional traits and their variation mediate plant species coexistence and spatial distribution. Yet, how patterns of variation in belowground traits influence resource acquisition across species and plant communities remains obscure. To characterize diverse belowground strategies in relation to species coexistence and abundance, we assessed four key belowground traits - root diameter, root branching intensity, first-order root length and mycorrhizal colonization - in 27 coexisting species from three grassland communities along a precipitation gradient. Species with thinner roots had higher root branching intensity, but shorter first-order root length and consistently low mycorrhizal colonization, whereas species with thicker roots enhanced their capacity for resource acquisition by producing longer first-order roots and maintaining high mycorrhizal colonization. Plant species observed across multiple sites consistently decreased root branching and/or mycorrhizal colonization, but increased lateral root length with decreasing precipitation. Additionally, the degree of intraspecific trait variation was positively correlated with species abundance across the gradient, indicating that high intraspecific trait variation belowground may facilitate greater fitness and chances of survival across multiple habitats. These results suggest that a small set of critical belowground traits can effectively define diverse resource acquisition strategies in different environments and may forecast species survival and range shifts under climate change.

Trait-based approaches provide a useful framework to investigate plant strategies for resource acquisition, growth, and competition, as well as plant impacts on ecosystem processes. Despite significant progress capturing trait variation within and among stems and leaves, identification of trait syndromes within fine-root systems and between fine roots and other plant organs is limited. Here we discuss three underappreciated areas where focused measurements of fine-root traits can make significant contributions to ecosystem science. These include assessment of spatiotemporal variation in fine-root traits, integration of mycorrhizal fungi into fine-root-trait frameworks, and the need for improved scaling of traits measured on individual roots to ecosystem-level processes. Progress in each of these areas is providing opportunities to revisit how below-ground processes are represented in terrestrial biosphere models. Targeted measurements of fine-root traits with clear linkages to ecosystem processes and plant responses to environmental change are strongly needed to reduce empirical and model uncertainties. Further identifying how and when suites of root and whole-plant traits are coordinated or decoupled will ultimately provide a powerful tool for modeling plant form and function at local and global scales.

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.

Leaf and root phenology play important roles controlling plant productivity and ecosystem function, yet, few studies link patterns of leaf and root phenology across woody species. Trees with diffuse-porous wood anatomy tend to leaf-out before ring-porous species and we expected that increases in transpiration with spring leaf-out would be coupled with initiation of root production to support uptake of soil resources. Therefore, we hypothesized that the timing of root production would follow patterns of leaf production and wood anatomy. Root production was observed using minirhizotrons and related to leaf phenology across six temperate tree species with different wood anatomy in a common garden. As expected, leaves of diffuse-porous species emerged before ring-porous, followed by tracheid species. Root production peaked before bud break in five of the six species and before maximum leaf area index in all species, but did not follow expected patterns with leaf production. Our observations did not indicate tight linkages between root and leaf phenology but do highlight the potential for very early season root production and greater variation in the phenology of roots than leaves. Future work should identify the environmental factors and species traits that best explain variation in root phenology.

Root biomass, root production and lifespan, and root-mycorrhizal interactions govern soil carbon fluxes and resource uptake and are critical components of terrestrial models. However, limitations in data and confusions over terminology, together with a strong dependence on a small set of conceptual frameworks, have limited the exploration of root function in terrestrial models. We review the key root processes of interest to both field ecologists and modelers including root classification, production, turnover, biomass, resource uptake, and depth distribution to ask (1) what are contemporary approaches for modeling roots in terrestrial models? and (2) can these approaches be improved via recent advancements in field research methods? We isolate several emerging themes that are ready for collaboration among field scientists and modelers: (1) alternatives to size-class based root classifications based on function and the inclusion of fungal symbioses, (2) dynamic root allocation and phenology as a function of root environment, rather than leaf demand alone, (3) improved understanding of the treatment of root turnover in models, including the role of root tissue chemistry on root lifespan, (4) better estimates of root stocks across sites and species to parameterize or validate models, and (5) dynamic interplay among rooting depth, resource availability and resource uptake. Greater attention to model parameterization and structural representation of roots will lead to greater appreciation for belowground processes in terrestrial models and improve estimates of ecosystem resilience to global change drivers. (C) 2014 Elsevier B.V. All rights reserved.