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At least 19 records

Anthropogenic N deposition alters soil organic matter biochemistry and microbial communities on decaying fine roots

Abstract Fine root litter is a primary source of soil organic matter (SOM), which is a globally important pool of C that is responsive to climate change. We previously established that ~20 years of experimental nitrogen (N) deposition has slowed fine root decay and increased the storage of soil carbon (C; +18%) across a widespread northern hardwood forest ecosystem. However, the microbial mechanisms that have directly slowed fine root decay are unknown. Here, we show that experimental N deposition has decreased the relative abundance of Agaricales fungi (−31%) and increased that of partially ligninolytic Actinobacteria (+24%) on decaying fine roots. Moreover, experimental N deposition has increased the relative abundance of lignin‐derived compounds residing in SOM (+53%), and this biochemical response is significantly related to shifts in both fungal and bacterial community composition. Specifically, the accumulation of lignin‐derived compounds in SOM is negatively related to the relative abundance of ligninolytic Mycena and Kuehneromyces fungi, and positively related to Microbacteriaceae. Our findings suggest that by altering the composition of microbial communities on decaying fine roots such that their capacity for lignin degradation is reduced, experimental N deposition has slowed fine root litter decay, and increased the contribution of lignin‐derived compounds from fine roots to SOM. The microbial responses we observed may explain widespread findings that anthropogenic N deposition increases soil C storage in terrestrial ecosystems. More broadly, our findings directly link composition to function in soil microbial communities, and implicate compositional shifts in mediating biogeochemical processes of global significance.

Argiroff, William A.↗

Stored carbon partly fuels fine-root respiration but is not used for production of new fine roots

The relative use of new photosynthate compared to stored C for the production and maintenance of fine roots, and the rate of C turnover in heterogeneous fine-root populations, are poorly understood. We followed the relaxation of a 13C tracer in fine roots in a Liquidambar styraciflua plantation at the conclusion of a free-air CO2 enrichment experiment. Goals included quantifying the relative fractions of new photosynthate versus stored C used in root growth and root respiration, as well as the turnover rate of fine-root C fixed during [CO2] fumigation. New fine-root growth was largely from recent photosynthate, while nearly one-quarter of respired C was from a storage pool. Changes in the isotopic composition of the fine-root population over two full growing seasons indicated heterogeneous C pools; less than 10% of root C had a residence time < 3 months, while a majority of root C had a residence time > 2 years. Compared to a 1-pool model, a 2-pool model for C turnover in fine roots (with 5 and 0.37 yr-1 turnover times) doubles the fine-root contribution to forest NPP (9-13%) and supports the 50% root-to-soil transfer rate often used in models.

13C↗

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↗

Fine Root and Soil Organic Carbon Depth Distributions are Inversely Related Across Fertility and Rainfall Gradients in Lowland Tropical Forests

Humid tropical forests contain some of the largest soil organic carbon (SOC) stocks on Earth. Much of this SOC is in subsoil, yet variation in the distribution of SOC through the soil profile remains poorly characterized across tropical forests. We used a correlative approach to quantify relationships among depth distributions of SOC, fine root biomass, nutrients and texture to 1 m depths across 43 lowland tropical forests in Panama. The sites span rainfall and soil fertility gradients, and these are largely uncorrelated for these sites. We used fitted b parameters to characterize depth distributions, where b is a numerical index based on an asymptotic relationship, such that larger b values indicate greater concentrations of root biomass or SOC at depth in the profile. Root b values ranged from 0.82 to 0.95 and were best predicted by soil pH and extractable potassium (K) stocks. For example, the three most acidic (pH < 4) and Kpoor (< 20 g K m -2 ) soils contained 76 ± 5% of fine root biomass from 0 to 10 cm depth, while the three least acidic (pH > 6.0) and most K-rich (> 50 g K m -2 ) soils contained only 41 ± 9% of fine root biomass at this depth. Root b and SOC b values were inversely related, such that a large fine root biomass in surface soils corresponded to large SOC stocks in subsoils (50–100 cm). SOC b values were best predicted by soil pH and base cation stocks, with the three most base-poor soils containing 34 ± 8% of SOC from 50 to 100 cm depth, and the three most base-rich soils containing just 9 ± 2% of SOC at this depth. Nutrient depth distributions were not related to Root b or SOC b values. Here, these data show that large surface root biomass stocks are associated with large subsoil C stocks in strongly weathered tropical soils. Further studies are required to evaluate why this occurs, and whether changes in surface root biomass, as may occur with global change, could in turn influence SOC storage in tropical forest subsoils.

54 ENVIRONMENTAL SCIENCES↗

Dynamics of fine roots in five Chinese temperate forests

Quantifying fine root production and mortality is crucially needed for modeling forest ecosystem carbon cycling, but the fine root dynamics are poorly understood in Chinese temperate forests. We used a minirhizotron method to investigate spatial and temporal dynamics of fine roots (diameter ≤ 2 mm) in five representative temperate forests in northeastern China. Our specific objectives were to: (1) compare standing crop, production and mortality of fine roots among the five stands; (2) examine fine root phenology for the stands; and (3) examine vertical distribution patterns of fine roots for the stands. Fine root dynamics were significantly affected by forest type, soil layer, sampling time and their interactions. The mean values of fine root standing crop varied from 8.0 to 12.8 mm cm-2; those of production varied from 0.027 to 0.046 mm cm-2 d-1; and those of mortality varied from 0.013 to 0.024 mm cm-2 d-1. All stands had a similar seasonal “sinusoidal” pattern of fine root standing crop, and a “unimodal” pattern of production. However, the seasonal dynamics of the mortality was unsynchronized with that of the production. The minimum values of standing crop, production and mortality occurred in March for all stands, while the maximum values and occurring time differed among forest types. The occurrence of the maximum standing crop varied from DOY (day of year) 222 for the oak stand to DOY 271 for the aspen-birch stand; that of the maximum production varied from DOY 188 for the pine and hardwood stands to DOY 239 for the larch stand; and that of the maximum mortality varied from DOY 222 for the oak and aspen-birch stands to DOY 287 for the larch stand. The standing crop, production and mortality of fine roots tended to decrease with soil depths, of which the relative contribution at 0 -10 cm depth averaged 38%, 46%, and 58% of total, respectively. The fact that the production was approximate twice as great as the mortality suggested a net carbon input to the soil of these forests.

54 ENVIRONMENTAL SCIENCES↗

Redefining fine roots improves understanding of belowground contributions to terrestrial biosphere processes

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.

58 GEOSCIENCES↗

Functionally discrete fine roots differ in microbial assembly, microbial functional potential, and produced metabolites

Traditionally, fine roots were grouped using arbitrary size categories, rarely capturing the heterogeneity in physiology, morphology and functionality among different fine root orders. Fine roots with different functional roles are rarely separated in microbiome-focused studies and may result in confounding microbial signals and host-filtering across different root microbiome compartments. Using a 26-year-old common garden, we sampled fine roots from four temperate tree species that varied in root morphology and sorted them into absorptive and transportive fine roots. The rhizoplane and rhizosphere were characterized using 16S rRNA gene and internal transcribed spacer region amplicon sequencing and shotgun metagenomics for the rhizoplane to identify potential microbial functions. Fine roots were subject to metabolomics to spatially characterize resource availability. Both fungi and bacteria differed according to root functional type. We observed additional differences between the bacterial rhizoplane and rhizosphere compartments for absorptive but not transportive fine roots. Rhizoplane bacteria, as well as the root metabolome and potential microbial functions, differed between absorptive and transportive fine roots, but not the rhizosphere bacteria. Functional differences were driven by sugar transport, peptidases and urea transport. Our data highlights the importance of root function when examining root-microbial relationships, emphasizing different host selective pressures imparted on different root microbiome compartments.

59 BASIC BIOLOGICAL SCIENCES↗

Effects of fine-root senescence upon soil communities and nutrient flux into soil pools (Final Report)

Fine roots represent an important flux of carbon into terrestrial soils. Much of this flux occurs through the exudation of polysaccharides and other carbon compounds during the lifetime of fine-roots, but little is known about their contributions to flux as roots senesce, die and decompose. This project was designed 1) to test approaches for inducing senescence in fine roots and 2) assess the effects of the treatments on fine roots and the associated rhizosphere community. We implemented two fine-root treatments, a steam girdling approach and full severing of fine root modules from the tree, and we contrasted them with a control where roots were handled but not girdled. Fine roots and their associated community subjected to each of these three treatments were subsequently analyzed for changes in physical structure, nutrient content, community structure and gene function. Both girdling manipulations resulted in tissue disruption in fine roots and changes in the soil community relative to controls. In particular, microbial diversity for most taxa declined in girdled treatments whereas fungal diversity increased. We also observed changes in overall plant and fungal gene expression associated with treatment with plant gene expression declining over time in girdled roots with a simultaneous increase in fungal gene expression in these same treatments. Finally, in girdled fine root modules, we observed changes in gene function associated with senescence and plant stress in pine-associated genes while at the same time observing upregulation of genes associated with growth and proliferation in fungi. In conclusion, we were able to successfully girdle fine-roots in a field setting and resolve plant-level and community level changes in response to these girdling treatments.

54 ENVIRONMENTAL SCIENCES↗

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

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 N 2 -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.

54 ENVIRONMENTAL SCIENCES↗

Effects of Slope Aspect and Rainfall on Belowground Deep Fine Root Traits and Aboveground Tree Height

The vertical root distribution and rooting depth are the main belowground plant functional traits used to indicate drought resistance in arid and semiarid regions. The effects of the slope aspect on the aboveground traits are visible but not the belowground deep root traits. We aimed to investigate the fine root traits of the locust tree ( Robinia pseudoacacia L.) planted on southerly and northerly aspects, and the variations in the rooting depth in regions with different rainfall, as well as assessing how deep rooting, might affect the response to drought in a loess region. We selected three study sites with different rainfall amounts, with six sampling plots at each site (three each with southerly and northerly aspects). Soil core samples were collected down to the depth where no roots were present. The locust trees tended to develop deeper fine roots rather than greater heights. The tree height and diameter were greater for locust trees on northerly aspects, whereas trees on southerly aspects had significantly deeper rooting depths. Fine root traits (root length, root area, and root dry weight density) were higher in the southerly aspect for both Changwu and Ansai, but lower in Suide. The ratio of the root front depth tree height ranged from 1.04 to 3.17, which was higher on southerly than northerly aspects, and it increased as the rainfall decreased. Locust tree growth traits (belowground fine root and aboveground tree height) were positively correlated with the mean annual rainfall. The soil moisture content of the topsoil decreased as the rainfall decreased, but the pattern varied in the deep layer. Our results suggest that the variations in the belowground rooting depth under different slope aspects may be related to plant survival strategies. The vertical extension of the rooting depth and tree height may be key functional traits that determine plant growth in drought-prone regions.

Lihui, Ma↗

Peatland warming strongly increases fine-root growth

Belowground climate change responses remain a key unknown in the Earth system. Plant fine-root response is especially important to understand because fine roots respond quickly to environmental change, are responsible for nutrient and water uptake, and influence carbon cycling. However, fine-root responses to climate change are poorly constrained, especially in northern peatlands, which contain up to two-thirds of the world’s soil carbon. We present fine-root responses to warming between +2 °C and 9 °C above ambient conditions in a whole-ecosystem peatland experiment. Warming strongly increased fine-root growth by over an order of magnitude in the warmest treatment, with stronger responses in shrubs than in trees or graminoids. In the first year of treatment, the control (+0 °C) shrub fine-root growth of 0.9 km m -2 y -1 increased linearly by 1.2 km m -2 y -1 (130%) for every degree increase in soil temperature. An extended belowground growing season accounted for 20% of this dramatic increase. In the second growing season of treatment, the shrub warming response rate increased to 2.54 km m -2 °C -1 . Soil moisture was negatively correlated with fine-root growth, highlighting that drying of these typically water-saturated ecosystems can fuel a surprising burst in shrub belowground productivity, one possible mechanism explaining the “shrubification” of northern peatlands in response to global change. This previously unrecognized mechanism sheds light on how peatland fine-root response to warming and drying could be strong and rapid, with consequences for the belowground growing season duration, microtopography, vegetation composition, and ultimately, carbon function of these globally relevant carbon sinks.

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↗

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. And 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 > 70 000 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. There has been a 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.

59 BASIC BIOLOGICAL SCIENCES↗

Proportion of fine roots, but not plant biomass allocation below ground, increases with elevation in arctic tundra

Questions: Roots represent a considerable proportion of biomass, primary production and litter input in arctic tundra, and plant allocation of biomass to above- or below- ground tissue in response to climate change is a key factor in the future C balance of these ecosystems. According to optimality theory plants allocate C to the above- or below-ground structure that captures the most limiting resource. We used an elevational gradient to test this theory and as a space-for-time substitution to inform on tundra carbon allocation patterns under a shifting climate, by exploring if increasing elevation was positively related to the root:shoot ratio, as well as a larger plant allocation to adsorptive over storage roots. Location: Arctic tundra heath dominated by Empetrum hermaphroditum close to Abisko, Sweden. Methods: We measured root:shoot and fine:coarse root ratios of the plant communities along an elevational gradient by sampling above- and below-ground biomass, further separating root biomass into fine (<1 mm) and coarse roots. Results: Plant biomass was higher at the lower elevations, but the root:shoot ratio did not vary with elevation. Resource allocation to fine relative to coarse roots increased with elevation, resulting in a fine:coarse root ratio that more than doubled with in- creasing elevation. Conclusions: Contrary to previous works, the root:shoot ratio along this elevational gradient remained stable. However, communities along our study system were dominated by the same species at each elevation, which suggests that when changes in the root:shoot ratio occur with elevation these changes may be driven by differences in allocation patterns among species and thus turnover in plant community structure. Our results further reveal that the allocation of biomass to fine relative to coarse roots can differ between locations along an elevational gradient, even when overall above- vs below-ground biomass allocation does not. Given the functionally different roles of fine vs coarse roots this could have large implications for below-ground C cycling. Thus. our results highlight the importance of direct effects vs indirect effects (such as changes in plant community composition and nutrient availability) of climate change for future C allocation above and below ground.

59 BASIC BIOLOGICAL SCIENCES↗

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↗