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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↗

Acquisitive root exploration strategies help maintain higher peak sap flux rates during summer drought, but more root biomass does not

Roots are responsible for soil water uptake, yet little is known about how variation in fine-root traits relates to whole-tree water movement, particularly during periods of drought. By combining a 3-yr dataset monitoring sap flow rates with measures of fine-root biomass, length, and morphology across 10 tree species, we addressed hypotheses concerning the role of fine-root system size and morphology in determining tree responses to summer drought as well as potential changes in their relative importance under more severe droughts. Greater fine-root biomass and length did not enable trees to maintain high rates of sap flow during drought, whereas the morphological traits, specific root length and specific root area, were linked to sustained sap flow rates during drought. We found that all species, irrespective of root traits, progressively reduced their sap flow under more extreme drought conditions, although more acquisitive root morphology was still associated with smaller reductions. Our results run counter to long-standing assumptions that larger root systems are better able to access soil water and maintain photosynthetic activity during drought. Instead, we find evidence that root morphology at least partially determines the capacity for water uptake and movement as soil moisture declines.

drought↗

Tropical forests and global change: biogeochemical responses and opportunities for cross-site comparisons, an organized INSPIRE session at the 108th Annual Meeting, Ecological Society of America, Portland, Oregon, USA, August 2023

In this study, tropical forests play a critical role in the global carbon (C) cycle. These ecosystems maintain the highest rates of net primary production (NPP) on Earth, contain c. 30% of terrestrial C stocks, and have some of the largest stores of fine-root biomass globally, as well as higher fine-root production and turnover rates compared with other biomes. Tropical forest responses to projected warming, altered rainfall regimes, and elevated CO 2 concentrations are likely to be different from other ecosystems because of their unique characteristics (Box 1), making targeted research and model development important for understanding tropical forest–climate feedbacks. There is now a critical mass of long-term global change field experiments and modeling efforts in tropical forests, yet thus far there has been little synthesis, cross-site comparison, or multi-site standardized experimentation among tropical forests to help us understand how these biomes are changing. An organized INSPIRE session at the 108th Annual Meeting of the Ecological Society of America set out to tackle just this. Speakers covered large-scale tropical forest field experiments and modeling efforts, with an emphasis on changes in ecosystem biogeochemistry under warming, drying, elevated atmospheric CO 2 , and changing nutrient status. In this Meeting report, we provide an overview of the large-scale global change experiments presented and highlight the main objectives and opportunities for tropical forest research that emerged, including cross-site comparisons and integration with ecosystem-scale models (Fig. 1).

54 ENVIRONMENTAL SCIENCES↗

Stand development and other intrinsic factors largely control fine-root dynamics with only subtle modifications from resource availability

Forest productivity depends on resource acquisition by ephemeral roots and leaves. A combination of intrinsic and environmental factors influences ephemeral organs; however, difficulties in studying belowground organs impede mechanistic understanding of fine-root production and turnover. To quantify factors controlling fine-root dynamics, we grew a deciduous hardwood (Populus deltoides Bartr.) and an evergreen conifer (Pinus taeda L.) with distinct soil moisture and nutrient availability treatments. We monitored fine-root dynamics with minirhizotrons for 6 years during early stand development and expressed results on a root length, biomass and mortality-risk basis. Stand development and other intrinsic factors consistently influenced both species in the same direction and by similar magnitude. Live-root length increased to a peak during establishment and slowly declined after roots of neighboring trees overlapped. Root longevity was highest during establishment and decreased thereafter. Root longevity consistently increased with depth of appearance and initial root diameter. Season of appearance affected root longevity in the following order: spring > summer > fall > winter. The influence of soil resource availability on fine-root dynamics was inconsistent between species, and ranked below that of rooting depth, initial diameter, stand development and phenology. Fine-root biomass either increased or was unaffected by greater resource availability. Fine-root production and live root length decreased with irrigation for both species, and increased with fertilization only for poplar. Fine-root mortality risk both increased and decreased depending on species and amendment treatment. Differing responses to soil moisture and nutrient availability between species suggests we should carefully evaluate generalizations about the response of fine-root dynamics to resource availability. Finally, while attempting to describe and explain carbon allocation to fine-root production and turnover, modelers and physiologists should first consider consistent patterns of allocation caused by different depth, diameter, stand development, phenology and species before considering allocation due to soil resource availability.

59 BASIC BIOLOGICAL SCIENCES↗

Leaf Area, Vegetation Biomass and Nutrient Content, Utqiagvik (Barrow), Alaska, 2012 - 2013

This dataset consists of measurements of vegetation harvested from Areas A to D of Intensive Site 1 at the Next-Generation Ecosystem Experiments (NGEE) Arctic site near Barrow, Alaska. The dataset includes i) values of leaf area index, biomass, carbon (C), nitrogen (N) and phosphorus (P) content of aboveground plant parts from 0.25 m x 0.25 m clip-plots at peak growing season and ii) fine-root biomass from 5.08-cm diameter soil cores taken throughout the active layer in the same location as the clip plots in late July-early August 2012, and iii) values of aboveground biomass and nitrogen (N) content measured from 0.1 m x 0.1 m clip-plots harvested at 2-week intervals throughout the 2013 growing season. This dataset includes five csv files and one pdf file.The Next-Generation Ecosystem Experiments: Arctic (NGEE Arctic), was a research effort to reduce uncertainty in Earth System Models by developing a predictive understanding of carbon-rich Arctic ecosystems and feedbacks to climate. NGEE Arctic was supported by the Department of Energy's Office of Biological and Environmental Research.The NGEE Arctic project had two field research sites: 1) located within the Arctic polygonal tundra coastal region on the Barrow Environmental Observatory (BEO) and the North Slope near Utqiagvik (Barrow), Alaska and 2) multiple areas on the discontinuous permafrost region of the Seward Peninsula north of Nome, Alaska.Through observations, experiments, and synthesis with existing datasets, NGEE Arctic provided an enhanced knowledge base for multi-scale modeling and contributed to improved process representation at global pan-Arctic scales within the Department of Energy's Earth system Model (the Energy Exascale Earth System Model, or E3SM), and specifically within the E3SM Land Model component (ELM).

%_nitrogen↗

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↗

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↗

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↗

Root-soil depth profile in Luquillo Experimental Forest, Puerto Rico, February, 2019

With an objective of associating the distribution of roots in the soil profile with functional attributes related to phosphorus nutrition, 1-m cores were collected from sites in the Luquillo Experimental Forest, Puerto Rico, in February, 2019. The sampled sites were ridge and valley locations at El Verde and Icacos with soils that differ in parent material and phosphorus fertility. Depth profiles of soil physical characteristics (bulk density, moisture content, and particle size distribution), phosphorus content (total phosphorus, organic phosphorus, and resin or available phosphorus), soil and root-associated phosphatase activity, and fine-root traits (biomass, specific root length and area, and phosphorus content) were determined.

54 ENVIRONMENTAL SCIENCES↗

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↗

Processed sap flow and fine-root trait data associated with summer drought responses in temperate trees in Lisle, Illinois, USA (2019–2021)

These data support the manuscript “Acquisitive root exploration strategies help maintain higher peak sap flux rates during summer drought, but more root biomass does not”. The dataset includes processed sap flow measurements and fine-root trait data collected between 2019 and 2021 from temperate monodominant tree plots established in the 1920s to 1930s ranging in size from 0.05 to 0.8 ha at The Morton Arboretum in Lisle, IL. Sap flow was measured with ICT sap flow sensors using the heat ratio method. Fine-root traits were measured from soil cores which includes specific root length (SRL), specific root area (SRA), diameter, biomass, and length for diameter classes ≤1 mm and ≤2 mm. The package contains comma separated value (CSV) data files and associated metadata that can be viewed and analyzed using common software such as spreadsheet programs, R, and Python. These data are used to investigate how variation in fine-root traits relate to tree water use and drought response during summer drought linking belowground root traits and aboveground physiological responses.

drought↗

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↗

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↗

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↗