Functionally discrete fine roots differ in microbial assembly, microbial functional potential, and produced metabolites.

Traditionally, fine roots were grouped using arbitrary size categories, rarely capturing the heterogeneity in physiology, morphology and functionality among different fine root orders. Fine roots with different functional roles are rarely separated in microbiome-focused studies and may result in confounding microbial signals and host-filtering across different root microbiome compartments. Using a 26-year-old common garden, we sampled fine roots from four temperate tree species that varied in root morphology and sorted them into absorptive and transportive fine roots. The rhizoplane and rhizosphere were characterized using 16S rRNA gene and internal transcribed spacer region amplicon sequencing and shotgun metagenomics for the rhizoplane to identify potential microbial functions. Fine roots were subject to metabolomics to spatially characterize resource availability. Both fungi and bacteria differed according to root functional type. We observed additional differences between the bacterial rhizoplane and rhizosphere compartments for absorptive but not transportive fine roots. Rhizoplane bacteria, as well as the root metabolome and potential microbial functions, differed between absorptive and transportive fine roots, but not the rhizosphere bacteria. Functional differences were driven by sugar transport, peptidases and urea transport. Our data highlights the importance of root function when examining root-microbial relationships, emphasizing different host selective pressures imparted on different root microbiome compartments.

Effects of long-term feeding by spotted lanternfly (Hemiptera: Fulgoridae) on ecophysiology of common hardwood host trees.

While the invasive spotted lanternfly, Lycorma delicatula (White) [Hemiptera: Fulgoridae], continues to expand its range in the United States, there remains a knowledge gap regarding the economic threat that this pest presents to forest ecosystems and production nurseries. L. delicatula uses several common hardwood trees as hosts and a previous study found that short-term feeding can reduce growth of young maple saplings. Herein, long-term feeding over 4 consecutive seasons significantly reduced diameter growth and below-ground starch storage in roots of young silver maples (Acer saccharinum L.), weeping willows (Salix babylonica L.), river birches (Betula nigra L.), and trees of heaven (Ailanthus altissima [Mill.] Swingle) in response to L. delicatula feeding pressure in a density-dependent manner. In Year 3 when feeding pressure was the lowest, silver maple and willow recovered with greater diameter growth than in Year 2. Nutrients essential for photosynthesis and growth (iron, sulfur, and phosphorus) were reduced in leaves of all tree species compared to controls in the second year. This 4-yr study represents a worst-case scenario in which L. delicatula fed on the same trees for 4 consecutive growing seasons. In the wild, population numbers can vary greatly from year to year on individual trees and they move frequently among hosts (until autumn when they settle on A. altissima or other late-season hosts that have not yet senesced). Thus, we would not expect negative impacts of unconfined L. delicatula in natural settings on forest or ornamental trees to be as marked as reported here.

Linkages between Plant Community Composition and Soil Microbial Diversity in Masson Pine Forests.

Plant species identity influences soil microbial communities directly by host specificity and root exudates, and indirectly by changing soil properties. As a native pioneer species common in early successional communities, Masson pine (Pinus massoniana) forests are widely distributed in subtropical China, and play a key role in improving ecosystem productivity. However, how pine forest composition, especially the dominance of plant functional groups, affects soil microbial diversity remains unclear. Here, we investigated linkages among woody plant composition, soil physicochemical properties, and microbial diversity in forests along a dominance gradient of Masson pine. Soil bacterial and fungal communities were mainly explained by woody plant community composition rather than by woody species alpha diversity, with the dominance of tree (without including shrub) species and ectomycorrhizal woody plant species accounting for more of the variation among microbial communities than pine dominance alone. Structural equation modeling revealed that bacterial diversity was associated with woody plant compositional variation via altered soil physicochemical properties, whereas fungal diversity was directly driven by woody plant composition. Bacterial functional groups involved in carbohydrate and amino acid metabolism were negatively correlated with the availability of soil nitrogen and phosphorus, whereas saprotrophic and pathogenic fungal groups showed negative correlations with the dominance of tree species. These findings indicate strong linkages between woody plant composition than soil microbial diversity; meanwhile, the high proportion of unexplained variability indicates great necessity of further definitive demonstration for better understanding of forest-microbe interactions and associated ecosystem processes.

Functionally-explicit sampling can answer key questions about the specificity of plant-microbe interactions.

The rhizosphere is a nexus for plant-microbe interactions and, as a host-structured environment, a location of high activity for distinct microbes and plant species. Although our insights into this habitat have exploded in recent years, we are still limited in our ability to answer key questions about the specificity of these root-microbial relationships. In particular, it can be difficult to confirm or reject microbiome heritability in many plant systems and to pinpoint which microbial taxa are key to plant functioning. Like other host-structured environments, the rhizosphere is structurally, chemically, and biologically complex, driven largely by differences in root anatomy, location, and function. In this Correspondence, we describe a review of 377 "rhizosphere microbiome" research papers and demonstrate how matching a sampling method to the biological question can advance our understanding of host-microbe interactions in a functionally heterogeneous environment. We found that the vast majority of studies (92%) pool all roots from a root system during sampling, ignoring variation in microbial composition between roots of different function and limiting insight into key root-microbial relationships. Furthermore, approaches for removing root-associated microbes are highly variable and non-standard, complicating multi-study analyses. Our understanding of the strength and nature of host-microbe relationships in heterogenous host-microbiome environments can be clarified by targeting sampling to locations of high interaction. While the high complexity of the rhizosphere creates logistical challenges, we suggest that unambiguous language and refined approaches will improve our ability to match methods to research questions and advance our understanding of the specificity of plant-microbial interactions.

Phylogenetic Signal, Root Morphology, Mycorrhizal Type, and Macroinvertebrate Exclusion: Exploring Wood Decomposition in Soils Conditioned by 13 Temperate Tree Species.

Woodlands are pivotal to carbon stocks, but the process of cycling C is slow and may be most effective in the biodiverse root zone. How the root zone impacts plants has been widely examined over the past few decades, but the role of the root zone in decomposition is understudied. Here, we examined how mycorrhizal association and macroinvertebrate activity influences wood decomposition across diverse tree species. Within the root zone of six predominantly arbuscular mycorrhizal (AM) (Acer negundo, Acer saccharum, Prunus serotina, Juglans nigra, Sassafras albidum, and Liriodendron tulipfera) and seven predominantly ectomycorrhizal (EM) tree species (Carya glabra, Quercus alba, Quercus rubra, Betula alleghaniensis, Picea rubens, Pinus virginiana, and Pinus strobus), woody litter was buried for 13 months. Macroinvertebrate access to woody substrate was either prevented or not using 0.22 mm mesh in a common garden site in central Pennsylvania. Decomposition was assessed as proportionate mass loss, as explained by root diameter, phylogenetic signal, mycorrhizal type, canopy tree trait, or macroinvertebrate exclusion. Macroinvertebrate exclusion significantly increased wood decomposition by 5.9%, while mycorrhizal type did not affect wood decomposition, nor did canopy traits (i.e., broad leaves versus pine needles). Interestingly, there was a phylogenetic signal for wood decomposition. Local indicators for phylogenetic associations (LIPA) determined high values of sensitivity value in Pinus and Picea genera, while Carya, Juglans, Betula, and Prunus yielded low values of sensitivity. Phylogenetic signals went undetected for tree root morphology. Despite this, roots greater than 0.35 mm significantly increased woody litter decomposition by 8%. In conclusion, the findings of this study suggest trees with larger root diameters can accelerate C cycling, as can trees associated with certain phylogenetic clades. In addition, root zone macroinvertebrates can potentially limit woody C cycling, while mycorrhizal type does not play a significant role.

Impacts of short-term feeding by spotted lanternfly (Lycorma delicatula) on ecophysiology of young hardwood trees in a common garden.

Spotted lanternfly (SLF; Lycorma delicatula White; Hemiptera: Fulgoridae) invaded the US from Asia and was first detected in 2014; currently, populations have established in 14 states primarily in the Northeast and Mid-Atlantic. It feeds voraciously on phloem sap from a broad range of host plants, with a preference for tree of heaven (Ailanthus altissima [Sapindales: Simaroubaceae]), grapevines (Vitis spp. [Vitales: Vitaceae]), and several common hardwood tree species. We evaluated the impacts of fourth instars and adults confined to a single branch or whole trees on gas exchange attributes (carbon assimilation [photosynthetic rate], transpiration and stomatal conductance), selected nutrients, and diameter growth using young saplings of four host tree species planted in a common garden. In general, the effects of adults on trees were greater than nymphs, although there was variation depending on tree species, pest density, and time post-infestation. Nymphs on a single branch of red maple (Acer rubrum [Sapindales: Sapindaceae]), or silver maple (Acer saccharinum [Sapindales: Sapindaceae]) at three densities (0, 15, or 30) had no significant effects on gas exchange. In contrast, 40 adults confined to a single branch of red or silver maple rapidly suppressed gas exchange and reduced nitrogen concentration in leaves; soluble sugars in branch wood were reduced in the fall for silver maple and in the following spring for red maple. Fourth instars confined to whole silver maple trees reduced soluble sugars in leaves and branch wood, and reduced tree diameter growth by >50% during the next growing season. In contrast, fourth instars in whole tree enclosures had no effects on black walnut (Juglans nigra [Fagales: Juglandaceae]). SLF enclosed on tree of heaven at 80 adults per tree suppressed gas exchange after two weeks of feeding, but did not alter non-structural carbohydrates, nitrogen concentrations, or tree growth. Results suggest that moderate to heavy feeding by SLF on young maple saplings may impair tree growth, which could have implications for production nurseries and forest managers.

Topographical shifts in fine root lifespan in a mixed, mesic temperate forest.

Root lifespan, often is estimated in landscape- and ecosystem-level carbon models using linear approximations. In water manipulation experiments, fine root lifespan can vary with soil water content. Soil water content is generally structured by complex topography, which is largely unaccounted for in landscape- and ecosystem-scale carbon models. Topography governs the range of soil water content experienced by roots which may impact their lifespan. We hypothesized that root lifespan varied nonlinearly across a temperate, mesic, forested catchment due to differences in soil water content associated with topographic position. We expected regions of the landscape that were too wet or too dry would have soils that were not optimal for roots and thus result in shorter root lifespans. Specifically, we hypothesized that root lifespan would be longest in areas that consistently had soil water content in the middle of the soil water content spectrum, while in soils at either very low or very high soil water content, root lifespan would be relatively short. We tested this hypothesis by collecting and analyzing two years of minirhizotron and soil moisture data in plots widely distributed in the Shale Hills catchment of the Susquehanna-Shale Hills Critical Zone Observatory in Pennsylvania. We found that fine root lifespans were longer in traditionally wetter topographic regions, but detected no short term (biweekly) effect of soil moisture on root lifespan. Additionally, depth in soil, soil series, slope face orientation, and season of birth strongly affected root lifespans across the catchment. In contrast, lifespan was unaffected by root diameter or mycorrhizal association. Failure to account for these variables could result in erroneous estimates of fine root lifespan and, consequentially, carbon flux in temperate forested regions.

Limited evidence of vertical fine-root segregation in a subtropical forest.

Vertical root segregation and the resulting niche partitioning can be a key underpinning of species coexistence. This could result from substantial interspecific variations in root profiles and rooting plasticity in response to soil heterogeneity and neighbours, but they remain largely untested in forest communities. In a diverse forest in subtropical China, we randomly sampled > 4000 root samples from 625 0-30 cm soil profiles. Using morphological and DNA-based methods, we identified 109 woody plant species, determined the degree of vertical fine-root segregation, and examined rooting plasticity in response to soil heterogeneity and neighbour structure. We found no evidence of vertical fine-root segregation among cooccurring species. By contrast, root abundance of different species tended to be positively correlated within soil zones. Underlying these findings was a lack of interspecific variation in fine-root profiles with over 90% of species concentrated in the 0-10 cm soil zone with only one species dominating in the 10-20 cm soil zone. Root profiles exhibited low responsiveness to root neighbours but tended to be shallow in soils with low phosphorus and copper content. These findings suggest that if there is niche differentiation leading to coexistence in this diverse forest, it would be occurring by mechanisms other than vertical fine-root segregation.

The hierarchy of root branching order determines bacterial composition, microbial carrying capacity and microbial filtering.

Fine roots vary dramatically in their functions, which range from resource absorption to within-plant resource transport. These differences should alter resource availability to root-associated microorganisms, yet most root microbiome studies involve fine root homogenization. We hypothesized that microbial filtering would be greatest in the most distal roots. To test this, we sampled roots of six temperate tree species from a 23-year-old common garden planting, separating by branching order. Rhizoplane bacterial composition was characterized with 16S rRNA gene sequencing, while bacterial abundance was determined on a subset of trees through flow cytometry. Root order strongly impacted composition across tree species, with absorptive lower order roots exerting the greatest selective pressure. Microbial carrying capacity was higher in absorptive roots in two of three tested tree species. This study indicates lower order roots as the main point of microbial interaction with fine roots, suggesting that root homogenization could mask microbial recruitment signatures.

Limiting similarity shapes the functional and phylogenetic structure of root neighborhoods in a subtropical forest.

Environmental filtering and limiting similarity mechanisms can simultaneously structure community assemblages. However, how they shape the functional and phylogenetic structure of root neighborhoods remains unclear, hindering the understanding of belowground community assembly processes and diversity maintenance. In a 50-ha plot in a subtropical forest, China, we randomly sampled > 2700 root clusters from 625 soil samples. Focusing on 10 root functional traits measured on 76 woody species, we examined the functional and phylogenetic structure of root neighborhoods and linked their distributions with environmental cues. Functional overdispersion was pervasive among individual root traits (50% of the traits) and accentuated when different traits were combined. Functional clustering (20% of the traits) seemed to be associated with a soil nutrient gradient with thick roots dominating fertile areas whereas thin roots dominated infertile soils. Nevertheless, such traits also were sorted along other environmental cues, showing multidimensional adaptive trait syndromes. Species relatedness also was an important factor defining root neighborhoods, resulting in significant phylogenetic overdispersion. These results suggest that limiting similarity may drive niche differentiation of coexisting species to reduce competition, and that alternative root strategies could be crucial in promoting root neighborhood resource use and species coexistence.

On characterizing root function in perennial horticultural crops.

PREMISE: While root-order approaches to fine-root classification have shown wide utility among wild plants, they have seen limited use for perennial crop plants. Moreover, inadequate characterization of fine roots across species of domesticated perennial crops has led to a knowledge gap in the understanding of evolutionary and functional patterns associated with different fine-root orders. METHODS: We examined fine-root traits of common horticultural fruit and nut crops: Malus ×domestica, Prunus persica, Vitus vinifera, Prunus dulcis, and Citrus ×clementina. Additional roots were sampled from 33 common perennial horticultural crops, native to tropical, subtropical, and temperate regions, to examine variation in 1st- and 2nd-order absorptive roots. RESULTS: First-order roots of grape and 1st- and 2nd-order roots of apple and peach were consistently thin, nonwoody, mycorrhizal, and had high N:C ratios. In contrast, 4th- and 5th-order roots of grape and 5th-order roots of apple and peach were woody, nonmycorrhizal, had low N:C ratios, and were thicker than lower-order roots. Among the 33 horticultural species, diameter of 1st- and 2nd-order roots varied about 15-fold, ranging from 0.04 to 0.60 mm and 0.05 to 0.89 mm respectively. This variation generally was phylogenetically conserved across plant lineages. CONCLUSIONS: Collectively, our research shows that root-order characterization has considerably more utility than an arbitrary diameter cutoff for identifying roots of different functions in perennial horticultural crops. In addition, much of the variation in root diameter among species can be predicted by evolutionary relationships.

Remarkable Similarity in Timing of Absorptive Fine-Root Production Across 11 Diverse Temperate Tree Species in a Common Garden.

Long-term minirhizotron observations of absorptive fine roots provide insights into seasonal patterns of belowground root production and carbon dynamics. Our objective was to compare root dynamics over time across mature individuals of 11 temperate trees species: five evergreen and six deciduous. We analyzed the timing and growth on 1st-and 2nd-order roots in minirhizotron images down to a vertical depth of 35 cm, as well as monthly and total annual length production. Production patterns were related to total annual precipitation of the actual and previous year of root production over 6 years. The main or largest peak of annual fine-root production occurred between June and September for almost all species and years. In most years, when peaks occurred, the timing of peak root production was synchronized across all species. A linear mixed model revealed significant differences in monthly fine-root length production across species in certain years (species x year, P < 0.0001), which was strongly influenced by three tree species. Total annual root production was much higher in 2000-2002, when there was above-average rainfall in the previous year, compared with production in 2005-2007, which followed years of lower-than-average rainfall (2003-2006). Compared to the wetter period all species experienced a decline of at least 75% in annual production in the drier years. Total annual root length production was more strongly associated with previous year's (P < 0.001) compared with the actual year's precipitation (P = 0.003). Remarkably similar timing of monthly absorptive fine-root growth can occur across multiple species of diverse phylogeny and leaf habit in a given year, suggesting a strong influence of extrinsic factors on absorptive fine-root growth. The influence of previous year precipitation on annual absorptive fine-root growth underscores the importance of legacy effects in biological responses and suggests that a growth response of temperate trees to extreme precipitation or drought events can be exacerbated across years.

Asymmetry in above- and belowground productivity responses to N addition in a semi-arid temperate steppe.

Nitrogen (N) enrichment often increases aboveground net primary productivity (ANPP) of the ecosystem, but it is unclear if belowground net primary productivity (BNPP) track responses of ANPP. Moreover, the frequency of N inputs may affect primary productivity but is rarely studied. To assess the response patterns of above- and belowground productivity to rates of N addition under different addition frequencies, we manipulated the rate (0-50 g N m-2  year-1 ) and frequency (twice vs. monthly additions per year) of NH4 NO3 inputs for six consecutive years in a temperate grassland in northern China and measured ANPP and BNPP from 2012 to 2014. In the low range of N addition rates, BNPP showed the greatest negative response and ANPP showed the greatest positive responses with increases in N addition (<10 g N m-2  year-1 ). As N addition increased beyond 10 g N m-2  year-1 , increases in ANPP dampened and decreases in BNPP ceased altogether. The response pattern of net primary productivity (combined above- and belowground; NPP) corresponded more closely to ANPP than to BNPP. The N effects on BNPP and BNPP/NPP (fBNPP ) were not dependent on N addition frequency in the range of N additions typically associated with N deposition. BNPP was more sensitive to N addition frequency than ANPP, especially at low rates of N addition. Our findings provide new insights into how plants regulate carbon allocation to different organs with increasing N rates and changing addition frequencies. These root response patterns, if incorporated into Earth system models, may improve the predictive power of C dynamics in dryland ecosystems in the face of global atmospheric N deposition.

Author Correction: Evolutionary history resolves global organization of root functional traits.

We thank reader Joseph Craine for pointing out three inadvertent errors in this Letter. First, 4 of the 71 divergence dates extracted from ref. 1 of this Amendment and used in Fig. 1b of the original Letter were overestimated. The correct values are 45 million years ago (Ma) for Apocynaceae, 51 Ma for Anacardiaceae, 40 Ma for Primulaceae, and 53 Ma for Amaryllidaceae. These errors had little influence on the overall trend of Fig. 1b (r2 is now 0.48 rather than 0.54, with no change to P < 0.001) and do not change our conclusion and inferences. Second, we neglected to note that since refs. 1 and 2 of this Amendment considered only angiosperms, our Fig. 1b necessarily did not include gymnosperm taxa. The in-text reference to Fig. 1b should therefore read "all major angiosperm plant families in our dataset" rather than "all major vascular plant families in our dataset". Third, in Fig. 1c the trait value of mycorrhizal colonization for Machilus kwangtungensis was erroneously given the value 0.25 instead of 1.0. This error had little influence on the overall Fig. 1c trend, reducing r2 from 0.64 to 0.63 (with no change to P < 0.001).

Soil organic carbon stability in forests: Distinct effects of tree species identity and traits.

Rising atmospheric CO2 concentrations have increased interest in the potential for forest ecosystems and soils to act as carbon (C) sinks. While soil organic C contents often vary with tree species identity, little is known about if, and how, tree species influence the stability of C in soil. Using a 40 year old common garden experiment with replicated plots of eleven temperate tree species, we investigated relationships between soil organic matter (SOM) stability in mineral soils and 17 ecological factors (including tree tissue chemistry, magnitude of organic matter inputs to the soil and their turnover, microbial community descriptors, and soil physicochemical properties). We measured five SOM stability indices, including heterotrophic respiration, C in aggregate occluded particulate organic matter (POM) and mineral associated SOM, and bulk SOM δ15 N and ∆14 C. The stability of SOM varied substantially among tree species, and this variability was independent of the amount of organic C in soils. Thus, when considering forest soils as C sinks, the stability of C stocks must be considered in addition to their size. Further, our results suggest tree species regulate soil C stability via the composition of their tissues, especially roots. Stability of SOM appeared to be greater (as indicated by higher δ15 N and reduced respiration) beneath species with higher concentrations of nitrogen and lower amounts of acid insoluble compounds in their roots, while SOM stability appeared to be lower (as indicated by higher respiration and lower proportions of C in aggregate occluded POM) beneath species with higher tissue calcium contents. The proportion of C in mineral associated SOM and bulk soil ∆14 C, though, were negligibly dependent on tree species traits, likely reflecting an insensitivity of some SOM pools to decadal scale shifts in ecological factors. Strategies aiming to increase soil C stocks may thus focus on particulate C pools, which can more easily be manipulated and are most sensitive to climate change.

Linking fine root morphology, hydraulic functioning and shade tolerance of trees.

Background and Aims: Understanding root traits and their trade-off with other plant processes is important for understanding plant functioning in natural ecosystems as well as agricultural systems. The aim of the present study was to determine the relationship between root morphology and the hydraulic characteristics of several orders of fine roots (<2 mm) for species differing in shade tolerance (low, moderate and high). Methods: The morphological, anatomical and hydraulic traits across five distal root orders were measured in species with different levels of shade tolerance and life history strategies. The species studied were Acer negundo, Acer rubrum, Acer saccharum, Betula alleghaniensis, Betula lenta, Quercus alba, Quercus rubra, Pinus strobus and Pinus virginiana. Key Results: Compared with shade-tolerant species, shade-intolerant species produced thinner absorptive roots with smaller xylem lumen diameters and underwent secondary development less frequently, suggesting that they had shorter life spans. Shade-tolerant species had greater root specific hydraulic conductance among these roots due to having larger diameter xylems, although these roots had a lower calculated critical tension for conduit collapse. In addition, shade-intolerant species exhibited greater variation in hydraulic conductance across different root growth rings in woody transport roots of the same root order as compared with shade-tolerant species. Conclusions: Plant growth strategies were extended to include root hydraulic properties. It was found that shade intolerance in trees was associated with conservative root hydraulics but greater plasticity in number of xylem conduits and hydraulic conductance. Root traits of shade-intolerant species were consistent with the ability to proliferate roots quickly for rapid water uptake needed to support rapid shoot growth, while minimizing risk in uncertain environments.

Erratum: Evolutionary history resolves global organization of root functional traits.

This corrects the article DOI: 10.1038/nature25783.

Evolutionary history resolves global organization of root functional traits.

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.

Building a better foundation: improving root-trait measurements to understand and model plant and ecosystem processes.

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.