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Fine-root dynamics vary with soil depth and precipitation in a lown-utrient tropical forest in the Central Amazonia

A common assumption in tropical ecology is that root systems respond rapidly to climatic cues but that most of that response is limited to the uppermost layer of the soil, with relatively limited changes in deeper layers. However, this assumption has not been tested directly, preventing models from accurately predicting the response of tropical forests to environmental change. We measured seasonal dynamics of fine roots in a Central Amazonia mature forest using minirhizotrons to 90 cm depth, which were calibrated with fine roots extracted from soil cores. Root productivity and mortality in superficial soil layers were positively correlated with precipitation, whereas root standing length was greater during the dry periods at the deeper layers. Contrary to historical assumptions, a large fraction of fine-root standing biomass (46%) and productivity (41%) were found in soil layers deeper than 30 cm. Furthermore, root turnover decreased linearly with soil depth. Our findings demonstrate a strong relationship between fine-root dynamics and precipitation regimes in Central Amazonia. Our results also emphasize the importance of deep roots for accurate estimates of primary productivity and the interaction between roots and carbon, water, and nutrients.

54 ENVIRONMENTAL SCIENCES↗

Temporal patterns of fine‐root dynamics have little influence on seasonal soil CO 2 efflux in a mixed, mesic forest

Among the contributors to soil CO 2 efflux, there remains uncertainty about the contribution of root activity to the overall soil efflux. Soil water and temperature frequently have been used to predict a large portion of the variation in soil CO 2 efflux. We hypothesized that fine-root dynamics explain most of the remaining variability in soil CO 2 efflux that cannot be explained by soil temperature and water content. We anticipated that seasonal increases in root production, mortality via decomposition, and standing crop would result in corresponding increases in soil CO 2 efflux. We tested our hypotheses by collecting and analyzing two years of minirhizotron and soil chamber CO 2 flux data from plots distributed throughout the Shale Hills Catchment of the Susquehanna-Shale Hills Critical Zone Observatory in Central Pennsylvania, USA. Here we showed that: (1) seasonal fluctuations in fine-root dynamics yielded only a very small increase in the predictability of soil CO 2 efflux; (2) fine-root mortality effects on soil CO 2 efflux were strongly tied to soil temperature; (3) fluctuations in fine-root presence or standing mass independent of temperature and moisture had little effect on soil CO 2 efflux; and (4) new fine-root length and root length mortality had limited impacts on soil CO 2 efflux rates. We conclude that, at least in temperate forests on rocky soils, characterizing fine-root dynamics may provide only limited improvement in the estimation of soil CO 2 efflux.

54 ENVIRONMENTAL SCIENCES↗

Tradeoffs and Synergies in Tropical Forest Root Traits and Dynamics for Nutrient and Water Acquisition: Field and Modeling Advances

Vegetation processes are fundamentally limited by nutrient and water availability, the uptake of which is mediated by plant roots in terrestrial ecosystems. While tropical forests play a central role in global water, carbon, and nutrient cycling, we know very little about tradeoffs and synergies in root traits that respond to resource scarcity. Tropical trees face a unique set of resource limitations, with rock-derived nutrients and moisture seasonality governing many ecosystem functions, and nutrient versus water availability often separated spatially and temporally. Root traits that characterize biomass, depth distributions, production and phenology, morphology, physiology, chemistry, and symbiotic relationships can be predictive of plants’ capacities to access and acquire nutrients and water, with links to aboveground processes like transpiration, wood productivity, and leaf phenology. In this review, we identify an emerging trend in the literature that tropical fine root biomass and production in surface soils are greatest in infertile or sufficiently moist soils. We also identify interesting paradoxes in tropical forest root responses to changing resources that merit further exploration. For example, specific root length, which typically increases under resource scarcity to expand the volume of soil explored, instead can increase with greater base cation availability, both across natural tropical forest gradients and in fertilization experiments. Also, nutrient additions, rather than reducing mycorrhizal colonization of fine roots as might be expected, increased colonization rates under scenarios of water scarcity in some forests. Efforts to include fine root traits and functions in vegetation models have grown more sophisticated over time, yet there is a disconnect between the emphasis in models characterizing nutrient and water uptake rates and carbon costs versus the emphasis in field experiments on measuring root biomass, production, and morphology in response to changes in resource availability. Closer integration of field and modeling efforts could connect mechanistic investigation of fine-root dynamics to ecosystem-scale understanding of nutrient and water cycling, allowing us to better predict tropical forest-climate feedbacks.

54 ENVIRONMENTAL SCIENCES↗

Fine‐root dynamics in deeper soils: a critical but overlooked component of ecosystem responses to climate warming

Climate warming is predicted to strongly affect the functioning of terrestrial ecosystems. The plant root system is a critical component of these ecosystems, with fine roots, in particular, playing a key role in plant water and nutrient uptake and transport. In addition, root litter and exudation represent the dominant plant carbon inputs into soil. Consequently, understanding fine‐root responses to warming is essential for predicting how the growth, resilience, and carbon storage of terrestrial ecosystems will respond to future climate change. Despite the growing literature on fine‐root responses to warming, most studies have focused on topsoil (0–30 cm). However, a significant portion of the fine‐root mass occurs below this depth. For instance, c. 40% of fine‐root mass is found below 30 cm in temperate and tropical ecosystems. Due to the importance of fine roots for plants and belowground carbon cycling, focusing solely on surface soils overlooks the critical need for insights into how roots in deeper soil layers (e.g. below 30 cm) respond to warming. We argue that studying the entire soil profile is necessary to comprehensively understand fine‐root dynamics under climate warming and the implications for soil organic carbon cycling, water, and nutrient uptake.

climate warming↗

SPRUCE High-Resolution Minirhizotrons in an Experimentally-Warmed Peatland Provide an Unprecedented Glimpse at Fine Roots and their Fungal Partners: Supporting Data

Images were collected using first of their kind, non-destructive, high-resolution automated minirhizotrons (RhizoSystems, LLC) to assess the response of plant fine-root and fungal mycelium dynamics to elevated temperatures after 4-6 years of whole-ecosystem warming and exposure to elevated carbon dioxide concentrations (e[CO2]) in a peat bog where the SPRUCE (Spruce and Peatland Responses Under Changing Environments) experiment is located. We focused on two SPRUCE experimental plots: Plot 10 has elevated temperature (+9°C) and plot 19 is a control (+0°C). Both have elevated carbon dioxide (e[CO2]). Changes in root and fungal abundance with warming were estimated from a timeseries of landscape-level mosaiced images for each plot by measuring the proportional abundance of five belowground classes: fine roots of vascular plants, ectomycorrhizas, fungal hyphae, fungal rhizomorphs, and fungal sporocarps. To examine root and fungal phenology responses to warming, the length per individual root or fungal structure (except fungal hyphae that did not grow linearly but rather increased in areal coverage) were measured per image area of a set of timeseries patch-level mosaiced images for each plot. The experimental work was conducted in a Picea mariana [black spruce] – Sphagnum spp. bog forest in northern Minnesota, 40 km north of Grand Rapids, in the USDA Forest Service Marcell Experimental Forest (MEF). This ecosystem, which is located at the southern margin of the boreal forest, is considered especially vulnerable to climate change and anticipated to be near its tipping point. These data were used in analyses published in Defrenne et al (2021). This dataset contains 5 data files in comma-separate values (*.csv) format and a compressed folder (*.zip) containing 383 JPEG (*.jpg) images. Data files contain landscape-level assessment of belowground class abundance, patch-level growth phenology, and environmental variables (originally published in Hanson et al., 2016 and Hanson et al 2020). Images were collected with automated minirhizotrons and analyzed for phenology. Additional metadata are provided: 5 data dictionaries and a file-level metadata file in comma-separate values (.csv) format and a user guide in PDF (*.pdf) format.

54 ENVIRONMENTAL SCIENCES↗

Embracing fine-root system complexity in terrestrial ecosystem modeling

Projecting the dynamics and functioning of the biosphere requires a holistic consideration of whole-ecosystem processes. However, biases toward leaf, canopy, and soil modeling since the 1970s have constantly left fine-root systems being rudimentarily treated. As accelerated empirical advances in the last two decades establish clearly functional differentiation conferred by the hierarchical structure of fine-root orders and associations with mycorrhizal fungi, a need emerges to embrace this complexity to bridge the data-model gap in still extremely uncertain models. In this work, we propose a three-pool structure comprising transport and absorptive fine roots with mycorrhizal fungi (TAM) to model vertically resolved fine-root systems across organizational and spatial–temporal scales. Emerging from a conceptual shift away from arbitrary homogenization, TAM builds upon theoretical and empirical foundations as an effective and efficient approximation that balances realism and simplicity. A proof-of-concept demonstration of TAM in a big-leaf model both conservatively and radically shows robust impacts of differentiation within fine-root systems on simulating carbon cycling in temperate forests. Theoretical and quantitative support warrants exploiting its rich potentials across ecosystems and models to confront uncertainties and challenges for a predictive understanding of the biosphere. Echoing a broad trend of embracing ecological complexity in integrative ecosystem modeling, TAM may offer a consistent framework where modelers and empiricists can work together toward this grand goal.

59 BASIC BIOLOGICAL SCIENCES↗

Soil Carbon Stocks Not Linked to Aboveground Litter Input and Chemistry of Old-Growth Forest and Adjacent Prairie

The long-standing assumption that aboveground plant litter inputs have a substantial influence on soil organic carbon storage (SOC) and dynamics has been challenged by a new paradigm for SOC formation and persistence. We tested the importance of plant litter chemistry on SOC storage, distribution, composition, and age by comparing two highly contrasting ecosystems: an old-growth coast redwood (Sequoia sempervirens) forest, with highly aromatic litter, and an adjacent coastal prairie, with more easily decomposed litter. We hypothesized that if plant litter chemistry was the primary driver, redwood would store more and older SOC that was less microbially processed than prairie. Total soil carbon stocks to 110 cm depth were higher in prairie (35 kg C m —2 ) than redwood (28 kg C m —2 ). Radiocarbon values indicated shorter SOC residence times in redwood than prairie throughout the profile. Higher amounts of pyrogenic carbon and a higher degree of microbial processing of SOC appear to be instrumental for soil carbon storage and persistence in prairie, while differences in fine-root carbon inputs likely contribute to younger SOC in redwood. We conclude that at these sites fire residues, root inputs, and soil properties influence soil carbon dynamics to a greater degree than the properties of aboveground litter.

13C-NMR spectroscopy↗

TropiRoot 1.0: Database of tropical root characteristics across environments

Tropical ecosystems contain the world's largest biodiversity of vascular plants. Yet, our understanding of tropical functional diversity and its contribution to global diversity patterns is constrained by data availability. This discrepancy underscores an urgent need to bridge data gaps by incorporating comprehensive tropical root data into global datasets. Here, we provide a database of tropical root characteristics. This new database, TropiRoot 1.0, will be instrumental in evaluating an array of hypotheses pertaining to root functional ecology and plant biogeography, both within the tropics and relative to other global biomes. The data compilation was conducted by the TropiRoot Initiative, in partnership with the Fine-Root Ecology Database (FRED) and the Global Root Trait (GRooT) database, Colorado State University (CSU) and the Smithsonian Tropical Research Institute (STRI). Literature search and data extraction were conducted between 2020 and 2024. Literature was identified using Web of Science, Scopus, and complemented using the expert knowledge of members of TropiRoot. To provide broad environmental and geographical distributions, literature searches included root characteristics (traits) across global change drivers, natural gradients, and from different continents. We adopted FRED standardized data columns and streamlined the format to enhance accessibility for data extraction across various user groups. This optimized framework resulted in a smaller, yet comprehensive datasheet. To make the database compatible with other global root trait initiatives, column identification was standardized following the codes provided by FRED. These efforts culminated in data extracted from 104 new sources, resulting in more than 8000 rows of data (either species or community data). Most of the data in TropiRoot 1.0 include root characteristics such as root biomass, morphology, root dynamics, mass fraction, architecture, anatomy, physiology, and root chemistry. This initiative represents a 30% increase in the currently available data for tropical roots in FRED. TropiRoot 1.0 contains root characteristics from 25 different countries, where seven are located in Asia, six in South America, five in Central America and the Caribbean, four in Africa, two in North America, and 1 in Oceania. Due to the volume of data, when ancillary data were available, including soil data, these data were either extracted and included in the database or its availability was recorded in an additional column. Multiple contributors checked the entries for outliers during the collation process to ensure data quality. For text-based observations, we examined all cells to ensure that their content relates to their specific categories. For numerical observations, we ordered each numerical value from least to greatest and plotted the values, checking apparent outliers against the data in their respective sources and correcting or removing incorrect or impossible values. Some data (soil and aboveground) have different columns for the same variable presented in different units, including originally published units, but root characteristics data had units converted to match those reported in FRED. By filling a gap from global databases, TropiRoot 1.0 expands our knowledge of otherwise so far underrepresented regions and our ability to assess global trends. This advancement can be used to improve tropical forest representation in vegetation models. The data are freely available and should be cited when used.

FRED↗

Agile Allocation in the Tundra: A Single Growing Season of Warming Increases Nutrient Availability While Decreasing Fine-Root Length

The majority of plant biomass is located belowground in Arctic ecosystems and plant roots are responsible for the uptake of the nutrients that constrain plant growth in these infertile ecosystems. Despite performing a crucial role connecting primary producers to the soil, roots are relatively understudied in the Arctic and their functional response to a rapidly warming and increasingly variable climate is unknown. Here, we assessed whether one growing season with elevated temperatures would have an impact on nutrient uptake and allocation by applying a warming technique that increased daily air temperatures by 3.2 °C. Destructive sampling was performed at the peak of the growing season to quantify biomass pools of carbon (C) and nitrogen (N), root traits, and uptake of a 15 N tracer ( 15 NH 4 + ) for the dominant plant species, Arctagrostis latifolia. We found that soil nutrient availability increased with short-term warming, but A. latifolia NH 4 + uptake remained unchanged. Fine-root length density and root biomass within the soil profile, however, were both reduced by warming. N allocation patterns across plant tissues were also altered by warming. NH 4 + uptake was best fit with a logistic model that captured the spatial relationship between roots and soil (NH 4 + uptake expressed per length fine root and NH 4 + availability expressed per unit soil volume) rather than a traditional Michaelis–Menten model. Our results indicate that short-term experimental warming can shift plant–soil interactions, suggesting that the tundra’s belowground response to elevated temperatures may be more dynamic than previously recognized.

15N tracer↗

Tundra Vegetation Community Type, Not Microclimate, Controls Asynchrony of Above‐ and Below‐Ground Phenology

The below-ground growing season often extends beyond the above-ground growing season in tundra ecosystems and as the climate warms, shifts in growing seasons are expected. However, we do not yet know to what extent, when and where asynchrony in above- and below-ground phenology occurs and whether variation is driven by local vegetation communities or spatial variation in microclimate. Here, we combined above- and below-ground plant phenology metrics to compare the relative timings and magnitudes of leaf and fine-root growth and senescence across microclimates and plant communities at five sites across the Arctic and alpine tundra biome. We observed asynchronous growth between above- and below-ground plant tissue, with the below-ground season extending up to 74% (~56 days) beyond the onset of above-ground leaf senescence. Plant community type, rather than microclimate, was a key factor controlling the timing, productivity, and growth rates of fine roots, with graminoid roots exhibiting a distinct ‘pulse’ of growth later into the growing season than shrub roots. Our findings indicate the potential of vegetation change to influence below-ground carbon storage as the climate warms and roots remain active in unfrozen soils for longer. Taken together, our findings of increased root growth in soils that remain thawed later into the growing season, in combination with ongoing tundra vegetation change including increased shrub and graminoid abundance, indicate increased below-ground productivity and altered carbon cycling in the tundra biome.

below-ground↗

Fine-root functional trait responses to experimental warming: a global meta-analysis

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. In this work, 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.

59 BASIC BIOLOGICAL SCIENCES↗

Fine-Root Ecology Database (FRED): A Global Collection of Root Trait Data with Coincident Site, Vegetation, Edaphic, and Climatic Data, Version 4.

To address the need for a centralized root trait database, we compiled the Fine-Root Ecology Database (FRED) from published and unpublished data sources. We have continued to add to the FRED database since the release of FRED 1.0 in 2017, followed by 2.0 in 2018, and 3.0 in 2021. This new release of FRED 4.0 now has 213,941 observations of 238 root traits, for a combined total of roughly 3.4 million data fields for root traits and ancillary data together. FRED 4.0 has 39.8% more root trait observations than FRED 3.0 and a 34.4% increase in unique data sources. This release of FRED 4.0 also includes significant increases in geographic regions that have long been underrepresented in global datasets, notably in the tropical low latitudes. Ancillary data on associated site, vegetation, edaphic, and climatic conditions from across the globe have also increased concurrently with root trait observations. FRED is focused on fine roots (traditionally defined as roots less than 2 mm in diameter), as coarse roots are studied using different methodology, often at very different scales, and have different traits and trait interpretations. Despite this fine-root focus, FRED accepts data collected from roots of all sizes and contains observations of many root classes including coarse roots. Data collection will continue for the foreseeable future. The FRED4_Entire_Database_2026.csv file is the flat csv data file for FRED 4.0, and the FRED4_dd.csv file is the data dictionary of all columns available in FRED, including column IDs, column names, definitions, and unit (where applicable).

54 ENVIRONMENTAL SCIENCES↗

Data from a throughfall exclusion experiment: Fine root dynamics, morphology, chemistry, and AMF colonization across four lowland Panamanian forests

Fine roots regulate forest nutrient, carbon, and water cycling, yet their variation within and among tropical forests remains under-characterized. We quantified root productivity, disappearance, and stocks to 1 m using minirhizotron imaging, and we measured morphology, elemental composition [root carbon (C), root nitrogen (N), root phosphorus (P)], and arbuscular mycorrhizal fungi (AMF) colonization to 20 cm using ingrowth cores and sequential coring. Sampling took place in four distinct lowland Panamanian forests (32 plots; 8 per forest) from 2018 through 2022 under control and throughfall-exclusion (drought) treatments in the Panama Rainforest Changes with Experimental Drying (PARCHED) experiment.The dataset is presented as an Excel workbook with six tabs. The first tab is the data dictionary. Tab S1 contains ingrowth-core production and mortality, morphology and soil moisture. Tab S2 contains sequential-coring standing stocks with associated morphology and soil moisture. Tab S3 contains minirhizotron row data records to 1 m depth, including per-frame root length and diameter, normalized length metrics, and session timing. Tab S4 contains AMF colonization. Tab S5 contains fine-root chemistry at 0–10 cm, reporting %P, %C, %N, and C:N for samples collected via ingrowth cores and sequential-coring standing stocks. CSV mirrors for each tab are provided, and a KML file supplies coordinates for all 32 plots.Key variables span live and dead fine-root biomass (and coarse fractions where applicable), specific root length (SRL) and area (SRA), diameter, root tissue density (RTD), soil moisture, AMF colonization, root %N, %C, %P, and C:N, along with minirhizotron root length and diameter. Depth, season, treatment, and plot/site identifiers are included to support cross-tab integration and analysis from 0–100 cm (minirhizotron) and 0–20 cm (cores).Units are reported in-column and missing values are coded as NA. No special software is required to open or use the files (Excel, CSV, and KML compatible).

54 ENVIRONMENTAL SCIENCES↗