Variation In Fine Root Traits Research Articles

Available nitrogen and enzyme activity in rhizosphere soil dominate the changes in fine-root nutrient foraging strategies during plantation development

The variation in fine-root traits in response to soil resources (i.e., fine-root nutrient foraging strategy) is critical for plants to adapt to environmental changes and even win in intra- or interspecific resource competition. However, the patterns and driving mechanisms that change in fine-root traits and nutrient foraging strategies during the development of plantations remain unclear. We analyzed the relationships among fine-root traits, rhizosphere soil nutrient variables, and enzyme activities at four stages of Pinus massoniana plantations: 18 years (young), 30 years (middle-aged), 43 years (mature), and 63 years (overmature). We found that specific root area (SRA) and specific root length (SRL), which represent the speed of nutrient acquisition, decreased with stand development. Root tissue density (RTD) and specific root tip number (SRT), which represent the cost of nutrient foraging, decreased with stand development after peaking in the mature and middle-aged forests, respectively. Compared with young stands, fine-root organic carbon and total phosphorus contents decreased by 23 % in the overmature forest, but total nitrogen increased by nearly 50 %. Principal component analysis (PCA) showed that those fine-root traits significantly varied from young to overmature stands on PC1, and such a shift was consistent with changes in the growth period from fast (in the young forest) to slow (in the overmature forest) for P. massoniana. We suggest that those changes in fine-root traits along the PC1 represents a change in the fine-root foraging strategy of P. massoniana. This strategy (i.e., PC1) was negatively correlated with rhizosphere soil total nutrients (i.e., soil organic carbon and total nitrogen) and carbon (C)-nitrogen (N)-phosphorus (P) stoichiometry but positively correlated with available carbon, nitrogen, and C and N cycling-related enzyme activities. Specifically, the variation in rhizosphere soil NH4+ had the highest amount of explication (72 %) for the variation in the root nutrient foraging strategy. These results demonstrate that the intensity of the fine-root response to soil resources is related to the structure of rhizosphere soil available nutrients (especially soil available nitrogen) as well as carbon and nitrogen conversion enzyme activities during plantation development. Such relationships can reshape fine-root traits and change the fine-root nutrient foraging strategies during the development of plantations

Intraspecific variation in fine-root traits is larger than in aboveground traits in European herbaceous species regardless of drought.

Differences within species (Intraspecific trait variation - ITV) contribute substantially to overall trait variability and environmental harshness can reduce among-species variation. While aboveground traits have received considerable attention, knowledge about ITV in fine-root traits and how it differs from ITV in aboveground traits remains limited. This study examined the partitioning of trait variation aboveground and fine-root traits in 52 European herbaceous species and how such proportions change in response to drought, offering valuable insights for accurate functional species characterization and inter-species comparisons. We studied seven morphological aboveground and fine-root traits under drought and well-watered conditions in a greenhouse experiment. Linear mixed effect models and permutational multivariate analysis of variance (PERMANOVA) were employed to decompose trait variation, ensuring the robustness of our results. We also calculated variance partitioning for the combination of aboveground traits and the combination of fine-root traits, as well as pairs of analogous leaf and fine-root traits (i.e., traits that fulfill similar functions) for each treatment (control and drought). Among-species trait differences explained a greater proportion of overall variance than within-species variation, except for root dry matter content (RDMC). Height and leaf area stood out, with species' identity accounting for 87-90% of total trait variation. Drought had no significant effect on the proportions of variation in any of the traits. However, the combination of fine-root traits exhibited higher intraspecific variability (44-44%) than aboveground traits (19-21%) under both drought and control. Analogous root traits also showed higher ITV (51-50%) than analogous leaf traits (27-31%). Our findings highlight substantial within-species variation and the nuanced responses of fine-root traits, particularly RDMC, suggesting root traits' flexibility to soil heterogeneity that fosters less differentiation among species. Among-species trait differences, especially aboveground, may underscore distinct strategies and competitive abilities for resource acquisition and utilization. This study contributes to elucidate the mechanisms underlying the multifunctionality of the above- and belowground plants compartments.

Responses of vascular plant fine roots and associated microbial communities to whole-ecosystem warming and elevated CO2 in northern peatlands.

Warming and elevated CO2 (eCO2) are expected to facilitate vascular plant encroachment in peatlands. The rhizosphere, where microbial activity is fueled by root turnover and exudates, plays a crucial role in biogeochemical cycling, and will likely at least partially dictate the response of the belowground carbon cycle to climate changes. We leveraged the Spruce and Peatland Responses Under Changing Environments (SPRUCE) experiment, to explore the effects of a whole-ecosystem warming gradient (+0°C to 9°C) and eCO2 on vascular plant fine roots and their associated microbes. We combined trait-based approaches with the profiling of fungal and prokaryote communities in plant roots and rhizospheres, through amplicon sequencing. Warming promoted self-reliance for resource uptake in trees and shrubs, while saprophytic fungi and putative chemoorganoheterotrophic bacteria utilizing plant-derived carbon substrates were favored in the root zone. Conversely, eCO2 promoted associations between trees and ectomycorrhizal fungi. Trees mostly associated with short-distance exploration-type fungi that preferentially use labile soil N. Additionally, eCO2 decreased the relative abundance of saprotrophs in tree roots. Our results indicate that plant fine-root trait variation is a crucial mechanism through which vascular plants in peatlands respond to climate change via their influence on microbial communities that regulate biogeochemical cycles.

Differential Variation in Non-structural Carbohydrates in Root Branch Orders of Fraxinus mandshurica Rupr. Seedlings Across Different Drought Intensities and Soil Substrates.

Non-structural carbohydrates (NSCs) facilitate plant adaptation to drought stress, characterize tree growth and survival ability, and buffer against external disturbances. Previous studies have focused on the distribution and dynamics of NSCs among different plant organs under drought conditions. However, discussion about the NSC levels of fine roots in different root branch orders is limited, especially the relationship between fine root trait variation and NSC content. The objective of the study was to shed light on the synergistic variation in fine root traits and NSC content in different root branch orders under different drought and soil substrate conditions. The 2-year-old Fraxinus mandshurica Rupr. potted seedlings were planted in three different soil substrates (humus, loam, and sandy–loam soil) and subjected to four drought intensities (CK, mild drought, moderate drought, and severe drought) for 2 months. With increasing drought intensity, the biomass of fine roots decreased significantly. Under the same drought intensity, seedlings in sandy–loam soil had higher root biomass, and the coefficient of variation of 5th-order roots (37.4, 44.5, and 53% in humus, loam, and sandy–loam soil, respectively) was higher than that of lower-order roots. All branch order roots of seedlings in humus soil had the largest specific root length (SRL) and specific root surface area (SRA), in addition to the lowest diameter. With increasing drought intensity, the SRL and average diameter (AD) of all root branch orders increased and decreased, respectively. The fine roots in humus soil had a higher soluble sugar (SS) content and lower starch (ST) content compared to the loam and sandy–loam soil. Additionally, the SS and ST contents of fine roots showed decreasing and increasing tendencies with increasing drought intensities, respectively. SS and ST explained the highest degree of the total variation in fine root traits, which were 32 and 32.1%, respectively. With increasing root order, the explanation of the variation in root traits by ST decreased (only 6.8% for 5th-order roots). The observed response in terms of morphological traits of different fine root branch orders of F. mandshurica seedlings to resource fluctuations ensures the maintenance of a low cost-benefit ratio in the root system development.

The Economics Spectrum Drives Root Trait Strategies in Mediterranean Vegetation.

Extensive research efforts are devoted to understand fine root trait variation and to confirm the existence of a belowground root economics spectrum (RES) from acquisitive to conservative root strategies that is analogous to the leaf economics spectrum (LES). The economics spectrum implies a trade-off between maximizing resource acquisition and productivity or maximizing resource conservation and longevity; however, this theoretical framework still remains controversial for roots. We compiled a database of 320 Mediterranean woody and herbaceous species to critically assess if the classic economics spectrum theory can be broadly extended to roots. Fine roots displayed a wide diversity of forms and properties in Mediterranean vegetation, resulting in a multidimensional trait space. The main trend of variation in this multidimensional root space is analogous to the main axis of LES, while the second trend of variation is partially determined by an anatomical trade-off between tissue density and diameter. Specific root area (SRA) is the main trait explaining species distribution along the RES, regardless of the selected traits. We advocate for the need to unify and standardize the criteria and approaches used within the economics framework between leaves and roots, for the sake of theoretical consistency.

Variation in fine root traits with thinning intensity in a Chinese fir plantation insights from branching order and functional groups

Thinning is a widely used practice in forest management, but the acclimation mechanisms of fine roots to forest thinning are still unclear. We examined the variations in fine root traits of different branching orders and functional groups along a thinning intensity gradient in a 26-year-old Chinese fir (Cunninghamia lanceolata) plantation. With increasing thinning intensity, the root C concentration (RCC), root N concentration (RNC), specific root area (SRA), and specific root length (SRL) of the absorptive roots (the first two orders) significantly decreased, while root abundance (root biomass and root length density) and root tissue density (RTD) significantly increased. Fifty-four percent of the variation in the absorptive root traits could be explained by the soil N concentration and the biomass and diversity of the understorey vegetation. Conversely, transport root (third- and higher-order) traits did not vary significantly among different thinning intensities. The covariation of absorptive root traits across thinning intensities regarding two dimensions was as follows: the first dimension (46% of the total variation) represented changes in root abundance and chemical traits (related to RCC, RNC), belonging to an extensive foraging strategy; the second dimension (41% of the total variation) represented variations in root morphological traits (related to RTD, SRL and SRA), which is an intensive foraging strategy (i.e., root economic spectrum). These results suggested that the absorptive roots of Chinese fir adopt two-dimensional strategies to acclimate to the altered surroundings after thinning.

Topography as a factor driving small-scale variation in tree fine root traits and root functional diversity in a species-rich tropical montane forest.

We investigated the variation in tree fine root traits and their functional diversity along a local topographic gradient in a Neotropical montane forest to test if fine root trait variation along the gradient is consistent with the predictions of the root economics spectrum on a shift from acquisitive to conservative traits with decreasing resource supply. We measured five fine root functional traits in 179 randomly selected tree individuals of 100 species and analysed the variation of single traits (using Bayesian phylogenetic multilevel models) and of functional trait diversity with small-scale topography. Fine roots exhibited more conservative traits (thicker diameters, lower specific root length and nitrogen concentration) at upper slope compared with lower slope positions, but the largest proportion of variation (40-80%) was explained by species identity and phylogeny. Fine root functional diversity decreased towards the upper slopes. Our results suggest that local topography and the related soil fertility and moisture gradients cause considerable small-scale variation in fine root traits and functional diversity along tropical mountain slopes, with conservative root traits and greater trait convergence being associated with less favourable soil conditions due to environmental filtering. We provide evidence of a high degree of phylogenetic conservation in fine root traits.

The response of fine root morphological and physiological traits to added nitrogen in Schrenk's spruce (Picea schrenkiana) of the Tianshan mountains, China.

Fine roots are essential for water and nutrient uptake in plants, but little is known about the variation in fine root traits and the underlying mechanisms that drive it. Understanding the responses of fine root function traits to changing environmental conditions and the role of fine root traits as drivers of forest ecosystem processes are critical for informing physiological and ecological theory as well as ecosystem management. We measured morphological and physiological traits of fine roots from six soil layers and three diameter classes in Schrenk’s spruce (Picea shrenkiana) forests of the Tianshan mountains, China. We found significant effects of nitrogen addition on these morphological and physiological traits, which varied by soil layer and root diameter. Specifically, specific root length (SRL) was higher in medium N addition group (N2) than in control group (N0). Specific root area (SRA) was higher in the control group (N0) than fertilized groups (N1, N2 and N3). Root tissue density (RTD) was higher in low N addition group (N1) than in the other group. Root dry matter content had no significant difference among four treatment groups. SRL, SRA, and RTD of fine roots in different diameter classes were all significantly different between high N addition (N3) and the control (N0) groups. The physiological characteristics of fine roots showed that soluble sugar (SS), fine root vitality (FRV), and tissue water content (TWC) in different soil layers were higher in the control group than in the fertilized groups. While soluble protein (SP), malondialdehyde (MDA) and free proline (FP) were lower in the control group (N0) than in the fertilized groups. In addition, SS, FRV, SP, TWC, FP, and MDA in all N addition treatments groups were significantly different from the control group. Fine root morphological traits were closely related to physiological traits, and added nitrogen inputs change these correlations. Our study confirms that nitrogen addition has specific effects on the morphological and physiological traits of fine roots of Schrenk’s spruce, and the effects of N addition vary according to the amount added.

Variation in fine root traits reveals nutrient-specific acquisition strategies in agroforestry systems

Changes in root functional traits reveal important nutrient acquisition strategies, with well documented patterns in root trait expression within complex communities or along gradients of singular nutrients. In this field study, we investigate intra-root functional trait expression with six soil macro- and micro-nutrients in Theobroma cacao agroforestry systems. Using image, chemical, and spatial analysis, the fine root distribution, architecture, and morphology of T. cacao were compared to localized soil nutrients on two-dimensional soil profiles with conspecific and heterospecific neighbours. Fine-scale variation in soil nutrients was observed within the range of T. cacao root systems. Higher NH4+ and Ca2+ was associated with greater root length and biomass densities, coupled with greater investment to individual roots, expressed as increased fine root tissue density and diameter and lower specific root length. Conversely, NO3− had the opposite effect. Overall, roots tended towards higher acquisitive trait values when next to a shade tree. Plants generally employ several concomitant and at times opposing strategies for nutrient acquisition in heterogeneous soils. We show that fine-scale root plasticity is highly linked to localized nutrient-specific and neighbour-specific effects, driving patterns of nutrient acquisition in agroforestry systems.

Six-Year Nitrogen-Water Interaction Shifts the Frequency Distribution and Size Inequality of the First-Order Roots of Fraxinus mandschurica in a Mixed Mature Pinus koraiensis Forest.

The variation in fine root traits in terms of size inequality at the individual root level can be identified as a strategy for adapting to the drastic changes in soil water and nutrient availabilities. The Gini and Lorenz asymmetry coefficients have been applied to describe the overall degree of size inequality, which, however, are neglected when conventional statistical means are calculated. Here, we used the Gini coefficient, Lorenz asymmetry coefficient and statistical mean in an investigation of Fraxinus mandschurica roots in a mixed mature Pinus koraiensis forest on Changbai Mountain, China. We analyzed 967 individual roots to determine the responses of length, diameter and area of the first-order roots and of branching intensity to 6 years of nitrogen addition (N), rainfall reduction (W) and their combination (NW). We found that first-order roots had a significantly greater average length and area but had smaller Gini coefficients in NW plots compared to in control plots (CK). Furthermore, the relationship between first-order root length and branching intensity was negative in CK, N, and W plots but positive in NW plots. The Lorenz asymmetry coefficient was >1 for the first-order root diameter in NW and W plots as well as for branching intensity in N plots. The bimodal frequency distribution of the first-order root length in NW plots differed clearly from the unimodal one in CK, N, and W plots. These results demonstrate that not only the mean but also the variation and the distribution mode of the first-order roots of F. mandschurica respond to soil nitrogen and water availability. The changes in size inequality of the first-order root traits suggest that Gini and Lorenz asymmetry coefficients can serve as informative parameters in ecological investigations of roots to improve our ability to predict how trees will respond to a changing climate at the individual root level.

Global drivers of fine root trait variation: Commentary on Freschet et al. (2017)

Global drivers of fine root trait variation: Commentary on Freschet <i>et al</i>. (2017)

Climate, soil and plant functional types as drivers of global fine‐root trait variation

Summary Ecosystem functioning relies heavily on below‐ground processes, which are largely regulated by plant fine‐roots and their functional traits. However, our knowledge of fine‐root trait distribution relies to date on local‐ and regional‐scale studies with limited numbers of species, growth forms and environmental variation. We compiled a world‐wide fine‐root trait dataset, featuring 1115 species from contrasting climatic areas, phylogeny and growth forms to test a series of hypotheses pertaining to the influence of plant functional types, soil and climate variables, and the degree of manipulation of plant growing conditions on species fine‐root trait variation. Most particularly, we tested the competing hypotheses that fine‐root traits typical of faster return on investment would be most strongly associated with conditions of limiting versus favourable soil resource availability. We accounted for both data source and species phylogenetic relatedness. We demonstrate that: (i) Climate conditions promoting soil fertility relate negatively to fine‐root traits favouring fast soil resource acquisition, with a particularly strong positive effect of temperature on fine‐root diameter and negative effect on specific root length (SRL), and a negative effect of rainfall on root nitrogen concentration; (ii) Soil bulk density strongly influences species fine‐root morphology, by favouring thicker, denser fine‐roots; (iii) Fine‐roots from herbaceous species are on average finer and have higher SRL than those of woody species, and N2‐fixing capacity positively relates to root nitrogen; and (iv) Plants growing in pots have higher SRL than those grown in the field. Synthesis. This study reveals both the large variation in fine‐root traits encountered globally and the relevance of several key plant functional types and soil and climate variables for explaining a substantial part of this variation. Climate, particularly temperature, and plant functional types were the two strongest predictors of fine‐root trait variation. High trait variation occurred at local scales, suggesting that wide‐ranging below‐ground resource economics strategies are viable within most climatic areas and soil conditions.

Building a better foundation: improving root-trait measurements to understand and model plant and ecosystem processes.

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.

A global Fine-Root Ecology Database to address below-ground challenges in plant ecology.

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 >70000 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.

Fine root foraging strategies in Norway spruce forests across a European climate gradient

AbstractFine root acclimation to different environmental conditions is crucial for growth and sustainability of forest trees. Relatively small changes in fine root standing biomass (FRB), morphology or mycorrhizal symbiosis may result in a large change in forest carbon, nutrient and water cycles. We elucidated the changes in fine root traits and associated ectomycorrhizal (EcM) fungi in 12 Norway spruce stands across a climatic and N deposition gradient from subarctic‐boreal to temperate regions in Europe (68°N–48°N). We analysed the standing FRB and the ectomycorrhizal root tip biomass (EcMB, g m−2) simultaneously with measurements of the EcM root morphological traits (e.g. mean root length, root tissue density (RTD), N% in EcM roots) and frequency of dominating EcM fungi in different stands in relation to climate, soil and site characteristics. Latitude and N deposition explained the greatest proportion of variation in fine root traits. EcMB per stand basal area (BA) increased exponentially with latitude: by about 12.7 kg m−2 with an increase of 10° latitude from southern Germany to Estonia and southern Finland and by about 44.7 kg m−2 with next latitudinal 10° from southern to northern Finland. Boreal Norway spruce forests had 4.5 to 11 times more EcM root tips per stand BA, and the tips were 2.1 times longer, with 1.5 times higher RTD and about 1/3 lower N concentration. There was 19% higher proportion of root tips colonized by long‐distance exploration type forming EcM fungi in the southern forests indicating importance of EcM symbiont foraging strategy in fine root nutrient acquisition. In the boreal zone, we predict ca. 50% decrease in EcMB per stand BA with an increase of 2 °C annual mean temperature. Different fine root foraging strategies in boreal and temperate forests highlight the importance of complex studies on respective regulatory mechanisms in changing climate.