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Since their emergence onto land, terrestrial plants have developed diverse strategies to acquire soil resources. However, we lack a framework that adequately captures how these strategies vary among species. Observations from around the world now allow us to quantify the variation observed in commonly-measured fine-root traits but it is unclear how root traits are interrelated and whether they fall along an “economic” spectrum of acquisitive to conservative strategies. We assessed root trait variation and mycorrhizal colonization rates by leveraging the largest global database of fine-root traits (the Fine-Root Ecology Database; FRED). We also developed a heuristic model to explore the role of mycorrhizal fungi in defining belowground exploration efficiency across a gradient of thin- to thick-diameter roots. In support of the expectations of the “root economic spectrum,” we found that root diameter was negatively related to specific root length (Pearson’s r =-0.76). However, we found an unexpected negative relationship between root diameter and root tissue density (Pearson’s r = -0.40), and we further observed that root nitrogen content was largely unrelated to other economic traits. Mycorrhizal colonization was most closely associated with root diameter (Pearson’s r = 0.62) and was unrelated to root tissue density and root nitrogen. The heuristic model demonstrated that while thinner roots have inherently greater capacity to encounter soil resources based on higher surface area per unit mass, the potential for increased associations with mycorrhizal fungi in thicker roots, combined with greater hyphal growth, can result in equally acquisitive strategies for both thin- and thick roots. Taken together, our assessments of root trait variation, trade-offs with mycorrhizal fungi, and broader connections to root longevity allowed us to propose a series of fundamental constraints on belowground resource acquisition strategies. Physical tradeoffs based on root construction (i.e., economic traits) and functional limitations related to the capacity of a root to encounter and acquire soil resources combine to limit the two-dimensional belowground trait space. Within this trait space there remains a diversity of additional variation in root traits that facilitates a wide range of belowground resource acquisition strategies.

Experimental evidence from a diverse set of forested ecosystems indicates that CO₂ enrichment may lead to deeper rooting distributions. While the causes of greater root production at deeper soil depths under elevated CO₂ concentration ([CO₂]) require further investigation, altered rooting distributions are expected to affect important ecosystem processes. The depth at which fine roots are produced may influence root chemistry, physiological function, and mycorrhizal infection, leading to altered nitrogen (N) uptake rates and slower turnover. Also, soil processes such as microbial decomposition are slowed at depth in the soil, potentially affecting the rate at which root detritus becomes incorporated into soil organic matter. Deeper rooting distributions under elevated [CO₂] provide exciting opportunities to use novel sensors and chemical analyses throughout the soil profile to track the effects of root proliferation on carbon (C) and N cycling. Models do not currently incorporate information on root turnover and C and N cycling at depth in the soil, and modification is necessary to accurately represent processes associated with altered rooting depth distributions. Progress in understanding and modeling the interface between deeper rooting distributions under elevated [CO₂] and soil C and N cycling will be critical in projecting the sustainability of forest responses to rising atmospheric [CO₂].

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 above- 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 multidimensional nature of plant trait variation better encompasses plant function and influence on the surrounding environment.

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.

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

Summary 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.

59 I. 59 II. 62 III. 69 IV. 73 73 References 73 SUMMARY: There is wide breadth of root function within ecosystems that should be considered when modeling the terrestrial biosphere. Root structure and function are closely associated with control of plant water and nutrient uptake from the soil, plant carbon (C) assimilation, partitioning and release to the soils, and control of biogeochemical cycles through interactions within the rhizosphere. Root function is extremely dynamic and dependent on internal plant signals, root traits and morphology, and the physical, chemical and biotic soil environment. While plant roots have significant structural and functional plasticity to changing environmental conditions, their dynamics are noticeably absent from the land component of process‐based Earth system models used to simulate global biogeochemical cycling. Their dynamic representation in large‐scale models should improve model veracity. Here, we describe current root inclusion in models across scales, ranging from mechanistic processes of single roots to parameterized root processes operating at the landscape scale. With this foundation we discuss how existing and future root functional knowledge, new data compilation efforts, and novel modeling platforms can be leveraged to enhance root functionality in large‐scale terrestrial biosphere models by improving parameterization within models, and introducing new components such as dynamic root distribution and root functional traits linked to resource extraction.

Stimulation of terrestrial plant production by rising CO₂ concentration is projected to reduce the airborne fraction of anthropogenic CO₂ emissions. Coupled climate–carbon cycle models are sensitive to this negative feedback on atmospheric CO₂, but model projections are uncertain because of the expectation that feedbacks through the nitrogen (N) cycle will reduce this so-called CO₂ fertilization effect. We assessed whether N limitation caused a reduced stimulation of net primary productivity (NPP) by elevated atmospheric CO₂ concentration over 11 y in a free-air CO₂ enrichment (FACE) experiment in a deciduous Liquidambar styraciflua (sweetgum) forest stand in Tennessee. During the first 6 y of the experiment, NPP was significantly enhanced in forest plots exposed to 550 ppm CO₂ compared with NPP in plots in current ambient CO₂, and this was a consistent and sustained response. However, the enhancement of NPP under elevated CO₂ declined from 24% in 2001–2003 to 9% in 2008. Global analyses that assume a sustained CO₂ fertilization effect are no longer supported by this FACE experiment. N budget analysis supports the premise that N availability was limiting to tree growth and declining over time—an expected consequence of stand development, which was exacerbated by elevated CO₂. Leaf- and stand-level observations provide mechanistic evidence that declining N availability constrained the tree response to elevated CO₂; these observations are consistent with stand-level model projections. This FACE experiment provides strong rationale and process understanding for incorporating N limitation and N feedback effects in ecosystem and global models used in climate change assessments.

Peatlands are an important carbon (C) reservoir storing one-third of global soil organic carbon (SOC), but little is known about the fate of these C stocks under climate change. Here, we examine the impact of warming and elevated atmospheric CO 2 concentration (eCO 2 ) on the molecular composition of SOC to infer SOC sources (microbe-, plant- and fire-derived) and stability in a boreal peatland. We show that while warming alone decreased plant- and microbe-derived SOC due to enhanced decomposition, warming combined with eCO 2 increased plant-derived SOC compounds. We further observed increasing root-derived inputs (suberin) and declining leaf/needle-derived inputs (cutin) into SOC under warming and eCO 2 . The decline in SOC compounds with warming and gains from new root-derived C under eCO 2 , suggest that warming and eCO 2 may shift peatland C budget towards pools with faster turnover. Together, our results indicate that climate change may increase inputs and enhance decomposition of SOC potentially destabilising C storage in peatlands. No inherently stable peat soil carbon. Researchers found that all molecular components of peatland soil organic carbon responded to warming and eCO2, including the components presumed to be slow cycling and stable.