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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.

Here, in the letter published in this issue of New Phytologist (pp. 310-312), 'Fine roots - functional definition expanded to crop species?' Dr. Zobel emphasizes the importance of heterogeneity within crop-root systems.

Aims Arbuscular mycorrhizal (AM) fungi are intimately associated with fine roots and are involved in nutrient acquisition. However, little information is available on the links between roots of individual orders and fungus colonization and community structure. Our aim was to elucidate AM fungal communities in the fine root systems of the temperate coniferous tree species, Cryptomeria japonica (Cupressaceae). Methods We characterized the morphological traits of AM fungi microscopically and determined the community structure of AM fungi using metabarcoding with an Ion Torrent Personal Genome Machine (PGM) focusing on lower-order roots from first to third order roots. Results Paris-type and Arum-type AM morphologies were both generally more prevalent on first-order roots than on second- or third-order roots, but the colonization rates by the Paris type were higher than those by the Arum type. We found a total of 48 fungal operational taxonomic units dominated by Glomeraceae, and all the AM taxa detected on third order roots were also found on first and/or second order roots. In the case of the second and third orders, AM fungal communities were affected by soil conditions: electrical conductivity, pH, and N concentration. Conclusion These results suggest that the abundance and species richness of AM fungi vary among lower root order systems, and that the AM community is sensitive to soil conditions and turns over as roots age.

Aims The phylogenetic variations of fine root traits, which are related to plant growth and development as well as to physiological and ecological processes, are not fully understood. This study aimed to: (1) examine how tree species and sampling methodology affect the anatomical, morphological and nutrient traits of fine roots; and (2) determine whether phylogenetic signals affect fine root trait relationships and influence comparison of root traits between the branch order-based and diameter-based cut-off sampling categories. Methods Fine root samples of 16 subtropical forest tree species were obtained and their anatomical, morphological and nutrient traits were studied. The phylogenetic signals of trait variations were calculated to determine trait relationships. Results Tree species and sampling methodology significantly affected fine root traits ( p  < 0.05). Mean root diameters, root tissue density (RTD) and carbon-to-nitrogen ratio were the lowest in the first-order category and highest in the ≤2 mm category. The reverse pattern was found for specific root length, specific root area and nitrogen concentration. Morphological traits showed significant phylogenetic signals; however, nutrient traits did not reflect phylogenetic conservatism. Phylogenetic factors influenced correlations between traits for the first-order root economics spectrum. Root traits were multidimensional and RTD was loaded on a novel phylogenetic principal component analysis. Conclusions Functional traits of fine roots are multidimensional for subtropical tree species and closely linked to phylogeny. Morphological traits of first order roots showed a much stronger phylogenetic signal than those of roots ≤2 mm (traditionally defined fine roots). The findings improve understanding of root trait strategies in response to environmental changes.

Stand density directly affects the distribution of ecological factors such as light, heat, and water in forest communities and changes the diversity and structure of undergrowth species, thereby affecting soil health. Fine roots can provide water and nutrients to plants rapidly in the fierce competition of soil resources, so as to get rid of environmental factors. This study examined the fine-root responses of the Populus tomentosa clone S86 to three stand densities (plant × row spacing: 2 × 2 m, 4 × 3 m, 4 × 5 m). We measured the biomass, morphology, and nitrogen content of lower- (1–3 order) and higher-order (>3 order) fine roots, and analyzed soil chemical properties in 10–30 cm. The soil from the density (2 × 2 m) stands showed lower soil organic matter content, available nitrogen, available phosphorous, and available potassium than others. Obviously, lower and higher-order fine roots were different: biomass of the >3 order accounted for 77–87% of the total biomass, 1–3-order fine-root diameter around 0.28–0.38 mm, while >3-order fine root were 1.28–1.69 mm; the length of 1–3-order fine root was longer than the >3 order, and root length density, specific root length, and nutrient content between the 1–3 and >3 orders were different. At 2 × 2 m, 1–3-order fine-root biomass was the highest, 132.5 g/m3, and the 1–3-order fine-root length, diameter, surface, root length density was also the highest; at the same time, the 1–3-order fine-root total nitrogen and organic matter content was also the highest, while the >3 order was highest under 4 × 3 m or 4 × 5 m. The findings of this study show that stand density affected the available nutrient content of the soil. When soil resources were poor, the biomass, morphology, and chemical content of fine roots were adjusted to increase the nutrient absorption rate, particularly in the lower-order roots.

Trees can build fine-root systems with high variation in root size (e.g., fine-root diameter) and root number (e.g., branching pattern) to optimize belowground resource acquisition in forest ecosystems. Compared with leaves, which are visible above ground, information about the distribution and inequality of fine-root size and about key associations between fine-root size and number is still limited. We collected 27,573 first-order fine-roots growing out of 3,848 second-order fine-roots, covering 51 tree species in three temperate forests (Changbai Mountain, CBS; Xianrendong, XRD; and Maoershan, MES) in Northeastern China. We investigated the distribution and inequality of fine-root length, diameter and area (fine-root size), and their trade-off with fine-root branching intensity and ratio (fine-root number). Our results showed a strong right-skewed distribution in first-order fine-root size across various tree species. Unimodal frequency distributions were observed in all three of the sampled forests for first-order fine-root length and area and in CBS and XRD for first-order fine-root diameter, whereas a marked bimodal frequency distribution of first-order fine-root diameter appeared in MES. Moreover, XRD had the highest and MES had the lowest inequality values (Gini coefficients) in first-order fine-root diameter. First-order fine-root size showed a consistently linear decline with increasing root number. Our findings suggest a common right-skewed distribution with unimodality or bimodality of fine-root size and a generalized trade-off between fine-root size and number across the temperate tree species. Our results will greatly improve our thorough understanding of the belowground resource acquisition strategies of temperate trees and forests.

To explore the adaptation of the fine root morphology and chemical characteristics of Tamarix chinensis to water–salt heterogeneity in the groundwater–soil system of a coastal wetland zone, T. chinensis forests at different groundwater levels (high: GW1 0.54 m and GW2 0.83 m; medium: GW3 1.18 m; low: GW4 1.62 m and GW5 2.04 m) in the coastal wetland of the Yellow River Delta were researched, and the fine roots of T. chinensis standard trees were excavated. The fine roots were classified by the Pregitzer method, and the morphology, nutrients, and nonstructural carbohydrate characteristics of each order were determined. The results showed that the groundwater level had a significant indigenous effect on the soil water and salt conditions and affected the fine roots of T. chinensis . At high groundwater levels, the specific root length and specific surface area of fine roots were small, the root tissue density was high, the fine root growth rate was slow, the nutrient use efficiency was higher than at low groundwater levels, and the absorption of water increased with increasing specific surface area. With decreasing groundwater level, the N content and C/N ratio of fine roots first decreased and then increased, and the soluble sugar, starch, and nonstructural carbohydrate content of fine roots first increased and then decreased. At high and low groundwater levels, the metabolism of fine roots of T. chinensis was enhanced, and their adaptability to high salt content and low water content soil environments improved. The first- and second-order fine roots of T. chinensis were mainly responsible for water and nutrient absorption, while the higher-order (from the third to fifth orders) fine roots were primarily responsible for the transportation and storage of carbohydrates. The fine root morphology, nutrients, nonstructural carbohydrate characteristics, and other aspects of the water and salt environment heterogeneity cooperated in a synergistic response and trade-off adjustment.

Understanding root decomposition dynamics is essential to address declining carbon sequestration and nutrient imbalances in monoculture plantations. This study elucidates how forest gaps regulate Pinus massoniana root decomposition through comparative methodological analysis, providing theoretical foundations for near-natural forest management and carbon–nitrogen cycle optimization in plantations. The results showed the following: (1) Root decomposition was significantly accelerated by the in situ soil litterbag method (ISLM) versus the traditional litterbag method (LM) (decomposition rate (k) = 0.459 vs. 0.188), reducing the 95% decomposition time (T0.95) by nearly nine years (6.53 years vs. 15.95 years). ISLM concurrently elevated the root potassium concentration and reconfigured the relationships between root decomposition and soil nutrients. (2) Lower-order roots (orders 1–3) decomposed significantly faster than higher-order roots (orders 4–5) (k = 0.455 vs. 0.193). This disparity was amplified under ISLM (lower-/higher-order root k ratio = 4.1) but diminished or reversed under LM (lower-/higher-order root k ratio = 0.8). (3) Forest gaps regulated decomposition through temporal phase interactions, accelerating decomposition initially (0–360 days) while inhibiting it later (360–720 days), particularly for higher-order roots. Notably, forest gap effects fundamentally reversed between methodologies (slight promotion under LM vs. significant inhibition under ISLM). Our study reveals that conventional LM may obscure genuine ecological interactions during root decomposition, confirms lower-order roots as rapid nutrient-cycling pathways, provides crucial methodological corrections for plantation nutrient models, and advances theoretical foundations for precision management of P. massoniana plantations.

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 due to challenges in consistent measurement and interpretation of fine-root systems. We define fine roots as all roots less than or equal to 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. 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 separated into either individual root orders or functionally defined into a shorter-lived absorptive pool and a longer-lived transport fine root pool. Furthermore, using these frameworks, we estimate that fine-root production and turnover represent 22% of terrestrial net primary production globally a ca. 30% reduction from previous estimates assuming a single fine-root pool. In the future we hope to develop tools to rapidly differentiate functional fine-root classes, explicit incorporation of mycorrhizal fungi in fine-root studies, and wider adoption of a two-pool approach to model fine roots provide opportunities to better understand belowground processes in the terrestrial biosphere.

Summary Roots vary in anatomy, morphology and physiology, both spatially (different parts of the same root system) and temporally (plastic changes, root ageing), suggesting that root trait measurements are strongly affected by root sampling categories. In this context, it is urgent to clarify the functional significance of current root sampling categories (e.g. fine roots of the first order, the first three orders, ≤1 mm or ≤2 mm), establish guidelines for choosing between sampling methods and revise root ontology to account for functional differences between traits measured on distinct root categories. Here, we used a worldwide database of fine‐root traits to test the hypothesis that distinct fine‐root trait values – with link to fine‐root functions – were generally affected by different root sampling categories. We observed indeed a clear functional break between first‐order roots and roots of all three other sampling categories, and a smaller but substantial break between roots of the three first orders and the ≤2 mm category, demonstrating globally that different sampling methodologies capture different functional parts of roots. Our synthesis suggests that all current root sampling categories present both advantages and pitfalls and that no single method can appropriately tackle the main current challenge of root functional ecology: i.e. linking fine roots to plant and ecosystem functions in a truly comparable way across all plants. We argue instead that a small set of complementary standardized sampling methods is necessary to capture the linkages between root forms and functions. To assist experimenters selecting adequate sampling we developed a decision table following three logical questions: (i) what plant or ecosystem function must be addressed; (ii) what root categories are involved in this function and (iii) what traits should be measured on these root categories. Challenging, strengthening and expending such common reference framework would be a substantial step towards wider comparability of future functional trait datasets. A lay summary is available for this article. Lay Summary