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

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

SummaryThe identification of plant functional traits that can be linked to ecosystem processes is of wide interest, especially for predicting vegetational responses to climate change. Root diameter of the finest absorptive roots may be one plant trait that has wide significance. Do species with relatively thick absorptive roots forage in nutrient-rich patches differently from species with relatively fine absorptive roots?We measured traits related to nutrient foraging (root morphology and architecture, root proliferation, and mycorrhizal colonization) across six coexisting arbuscular mycorrhizal (AM) temperate tree species with and without nutrient addition.Root traits such as root diameter and specific root length were highly correlated with root branching intensity, with thin-root species having higher branching intensity than thick-root species. In both fertilized and unfertilized soil, species with thin absorptive roots and high branching intensity showed much greater root length and mass proliferation but lower mycorrhizal colonization than species with thick absorptive roots. Across all species, fertilization led to increased root proliferation and reduced mycorrhizal colonization.These results suggest that thin-root species forage more by root proliferation, whereas thick-root species forage more by mycorrhizal fungi. In mineral nutrient-rich patches, AM trees seem to forage more by proliferating roots than by mycorrhizal fungi.

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

SummaryTrait-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 fineroot 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.

SummaryPatterns 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.

Root turnover is fastest in the finest roots of the root system (first root order). Additionally, tissue chemistry varies among even the finest root orders and between white roots and older, pigmented roots. Yet the effects of pigmentation and order on root decomposition have rarely been examined. We separated the first four root orders (all <1 mm) of four temperate tree species into three classes: white first‐ and second‐order roots; pigmented first‐ and second‐order roots; and pigmented third‐ and fourth‐order roots. Roots were enclosed in litterbags and buried under their own and under a common species canopy in a 34‐year‐old common garden in Poland. When comparing decomposition of different root orders over 36 months, pigmented third‐ and fourth‐order roots with a higher C:N ratio decomposed more rapidly, losing 20–40% of their mass, than pigmented first‐ and second‐order roots, which lost no more than 20%. When comparing decomposition of roots of different levels of pigmentation within the same root order over 14 months, pigmented (older) first‐ and second‐order roots lost ∼10% of their mass, while white (younger) first‐ and second‐order roots lost ∼30%. In contrast to root mass loss, root N content declined more rapidly in the first‐ and second‐order roots than in third‐ and fourth‐order roots. In higher‐order roots, N increased in the first 10 months from ∼110% to nearly 150% of initial N content, depending on species; by the end of the study N content had returned to initial levels. These findings suggest that, in plant communities where root mortality is primarily of pigmented first‐ and second‐order roots, microbial decomposition may be slower than estimates derived from bulk fine‐root litterbag experiments, which typically contain at least four root orders. Thus, a more mechanistic understanding of root decomposition and its contribution to ecosystem carbon and nutrient dynamics requires a fundamental shift in experimental methods that stratifies root samples for decomposition along more functionally based criteria such as root order and pigmentation, which parallel the markedly different longevities of these different root classes.

Summary• Not all roots born as first-order branches are the same and this has important consequences for overall function. We hypothesized that, compared with fibrous roots, pioneer roots are built to live longer at the expense of absorptive capacity.• We tested this hypothesis by investigating pioneer and fibrous roots in their first 14 d of life in the arbuscular mycorrhizal tree species: Acer negundo, Acer saccharum, Juglans nigra, Liriodendron tulipifera and Populus tremuloides. Root observations were made with root-access boxes that allowed roots to be sampled at known ages in field-grown trees.• Compared to fibrous roots, pioneer roots had larger diameter, lower specific root length, greater average length and a lack of mycorrhizal or nonmycorrhizal fungal colonization. Pioneer roots < 14 d old had more layers of hypodermis with a lower percentage of putative passage cells and more protoxylem groups than similar age fibrous roots.• Our results suggest that pioneer roots are constructed for defense against biotic and abiotic challenges, exploration of soil distal to the stem, high fibrous root branching and secondary development with high axial hydraulic conductivity at the expense of mycorrhizal colonization and high absorptive capacity for water and nutrients.