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Using 11 closely related grass species, we tested the capacity of physiological criteria to predict the growth response to elevated CO2 and to categorize the species with regard to their CO2 response. A growth analysis was conducted under productive conditions both at ambient (350 μmol mol-1) and elevated (700 μmol mol-1) CO2. The relative growth rate stimulation was regressed against each of the growth rate components measured at ambient CO2. Growth response to CO2 was positively correlated with specific leaf area (SLA, the leaf surface area per unit of leaf weight), leaf area ratio (the leaf area per unit of total plant dry weight) and negatively correlated with net assimilation rate and leaf nitrogen concentration, both per unit of leaf area. We suggest that SLA has a predominant role in these relationships. Different hypotheses are proposed and discussed in order to explain why species with low SLA are less responsive to elevated CO2. Neither biomass allocation, relative growth rate, shoot or root specific activities per unit of mass, nor chemical composition were significantly correlated with growth response to CO2. The four predictive criteria mentioned above coherently differentiate the five wild annual species (higher SLA, stronger growth response to CO2) from the four wild perennials. The two perennial crop species, with the highest SLA, were more responsive than the wild species. [ABSTRACT FROM AUTHOR]

The effect of CO2 enrichment on the growth and the economy of carbon and nitrogen of 11 Mediterranean grass species was investigated in order to determine the underlying causes of the large variation observed between species in their responses to elevated CO2. Plants were grown for 26-43 d (depending on species growth rate) under productive conditions at ambient (350 μmol mol-1) and elevated (700 μmol mol-1) concentrations of CO2. Plant parameters were determined at a common biomass of 0.15 g to determine the CO2 effect independent of ontogenic effects. The effect of CO2 on RGR ranged from -6.7 to 22.5% with a mean stimulation of 10.3%. Averaged over the 11 species, the growth enhancement resulted from an increase in net assimilation rate per unit leaf d. wt. (NARW) of 10.6%. This was the result of a large increase (18.7%) in NAR per unit leaf area (NARa) associated with a 8.1% decrease in the specific leaf area (SLA). This decrease in SLA was due to a large increase of the non-structural carbohydrates. The increase in shoot activity was balanced by a 7.6% increase in the specific absorption rate of nitrogen (SAR). AS a result, plant nitrogen content was not modified. Leaf nitrogen productivity was significantly increased (14.9%). Shoot vs. root allocation of biomass and nitrogen was not modified. An analysis across the 11 species of the relationships between the stimulation of RGR and the alteration in RGR components showed a significant correlation only with increases in NARW, SAR and nitrogen productivity. The coordinated increase in these three parameters constitutes a single response syndrome, whose intensity is responsible for most of the species variability. [ABSTRACT FROM AUTHOR]

We determined the proximate chemical composition as well as the construction costs of leaves of 27 species, grown at ambient and at a twice-ambient partial pressure of atmospheric CO2. These species comprised wild and agricultural herbaceous plants as well as tree seedlings. Both average responses across species and the range in response were considered. Expressed on a total dry weight basis, the main change in chemical composition due to CO2 was the accumulation of total non-structural carbohydrates (TNC). To a lesser extent, decreases were found for organic N compounds and minerals. Hardlyany change was observed for total structural carbohydrates (cellulose plus hemicellulose), lignin and lipids. When expressed on a TNC-free basis decreases in organic N compounds and minerals were still present. On this basis, there was also an increase in the concentration of soluble phenolics. In terms of glucose required for biosynthesis, the increase in costs for one chemical compound -- TNC -- was balancedby a decrease in the costs for organic N compounds. Therefore, the construction costs, the total amount of glucose required to produce 1 g of leaf, were rather similar for the two CO2 treatments;on average a small decrease of 3% was found. This decrease was attributable to a decrease of up to 30% in the growth respiration coefficient, the total CO2 respired [mainly for NAD(P)H and ATP] in the process of constructing 1 g of biomass. The main reasons for this reduction were the decrease in organic N compounds and the increase in TNC. [ABSTRACT FROM AUTHOR]

In the context of a recent massive increase in research on plant root functions and their impact on the environment, root ecologists currently face many important challenges to keep on generating cutting-edge, meaningful and integrated knowledge. Consideration of the below-ground components in plant and ecosystem studies has been consistently called for in recent decades, but methodology is disparate and sometimes inappropriate. This handbook, based on the collective effort of a large team of experts, will improve trait comparisons across studies and integration of information across databases by providing standardised methods and controlled vocabularies. It is meant to be used not only as starting point by students and scientists who desire working on below-ground ecosystems, but also by experts for consolidating and broadening their views on multiple aspects of root ecology. Beyond the classical compilation of measurement protocols, we have synthesised recommendations from the literature to provide key background knowledge useful for: (1) defining below-ground plant entities and giving keys for their meaningful dissection, classification and naming beyond the classical fine-root vs coarse-root approach; (2) considering the specificity of root research to produce sound laboratory and field data; (3) describing typical, but overlooked steps for studying roots (e.g. root handling, cleaning and storage); and (4) gathering metadata necessary for the interpretation of results and their reuse. Most importantly, all root traits have been introduced with some degree of ecological context that will be a foundation for understanding their ecological meaning, their typical use and uncertainties, and some methodological and conceptual perspectives for future research. Considering all of this, we urge readers not to solely extract protocol recommendations for trait measurements from this work, but to take a moment to read and reflect on the extensive information contained in this broader guide to root ecology, including sections I-VII and the many introductions to each section and root trait description. Finally, it is critical to understand that a major aim of this guide is to help break down barriers between the many subdisciplines of root ecology and ecophysiology, broaden researchers' views on the multiple aspects of root study and create favourable conditions for the inception of comprehensive experiments on the role of roots in plant and ecosystem functioning., (© 2021 The Authors. New Phytologist © 2021 New Phytologist Foundation.)

The effects of plants on the biosphere, atmosphere and geosphere are key determinants of terrestrial ecosystem functioning. However, despite substantial progress made regarding plant belowground components, we are still only beginning to explore the complex relationships between root traits and functions. Drawing on the literature in plant physiology, ecophysiology, ecology, agronomy and soil science, we reviewed 24 aspects of plant and ecosystem functioning and their relationships with a number of root system traits, including aspects of architecture, physiology, morphology, anatomy, chemistry, biomechanics and biotic interactions. Based on this assessment, we critically evaluated the current strengths and gaps in our knowledge, and identify future research challenges in the field of root ecology. Most importantly, we found that belowground traits with the broadest importance in plant and ecosystem functioning are not those most commonly measured. Also, the estimation of trait relative importance for functioning requires us to consider a more comprehensive range of functionally relevant traits from a diverse range of species, across environments and over time series. We also advocate that establishing causal hierarchical links among root traits will provide a hypothesis-based framework to identify the most parsimonious sets of traits with the strongest links on functions, and to link genotypes to plant and ecosystem functioning., (© 2021 The Authors. New Phytologist © 2021 New Phytologist Foundation.)

Plant trait variation drives plant function, community composition and ecosystem processes. However, our current understanding of trait variation disproportionately relies on aboveground observations. Here we integrate root traits into the global framework of plant form and function. We developed and tested an overarching conceptual framework that integrates two recently identified root trait gradients with a well-established aboveground plant trait framework. We confronted our novel framework with published relationships between above- and belowground trait analogues and with multivariate analyses of above- and belowground traits of 2510 species. Our traits represent the leaf and root conservation gradients (specific leaf area, leaf and root nitrogen concentration, and root tissue density), the root collaboration gradient (root diameter and specific root length) and the plant size gradient (plant height and rooting depth). We found that an integrated, whole-plant trait space required as much as four axes. The two main axes represented the fast-slow 'conservation' gradient on which leaf and fine-root traits were well aligned, and the 'collaboration' gradient in roots. The two additional axes were separate, orthogonal plant size axes for height and rooting depth. This perspective on the multidimensional nature of plant trait variation better encompasses plant function and influence on the surrounding environment., (© 2021 The Authors. New Phytologist © 2021 New Phytologist Foundation.)

Ecological theory is built on trade-offs, where trait differences among species evolved as adaptations to different environments. Trade-offs are often assumed to be bidirectional, where opposite ends of a gradient in trait values confer advantages in different environments. However, unidirectional benefits could be widespread if extreme trait values confer advantages at one end of an environmental gradient, whereas a wide range of trait values are equally beneficial at the other end. Here, we show that root traits explain species occurrences along broad gradients of temperature and water availability, but model predictions only resembled trade-offs in two out of 24 models. Forest species with low specific root length and high root tissue density (RTD) were more likely to occur in warm climates but species with high specific root length and low RTD were more likely to occur in cold climates. Unidirectional benefits were more prevalent than trade-offs: for example, species with large-diameter roots and high RTD were more commonly associated with dry climates, but species with the opposite trait values were not associated with wet climates. Directional selection for traits consistently occurred in cold or dry climates, whereas a diversity of root trait values were equally viable in warm or wet climates. Explicit integration of unidirectional benefits into ecological theory is needed to advance our understanding of the consequences of trait variation on species responses to environmental change., (© 2021. The Author(s), under exclusive licence to Springer Nature Limited.)

Objectives: Altitude integrates changes in environmental conditions that determine shifts in vegetation, including temperature, precipitation, solar radiation and edaphogenetic processes. In turn, vegetation alters soil biophysical properties through litter input, root growth, microbial and macrofaunal interactions. The belowground traits of plant communities modify soil processes in different ways, but it is not known how root traits influence soil biota at the community level. We collected data to investigate how elevation affects belowground community traits and soil microbial and faunal communities. This dataset comprises data from a temperate climate in France and a twin study was performed in a tropical zone in Mexico., Data Description: The paper describes soil physical and chemical properties, climatic variables, plant community composition and species abundance, plant community traits, soil microbial functional diversity and macrofaunal abundance and diversity. Data are provided for six elevations (1400-2400 m) ranging from montane forest to alpine prairie. We focused on soil biophysical properties beneath three dominant plant species that structure local vegetation. These data are useful for understanding how shifts in vegetation communities affect belowground processes, such as water infiltration, soil aggregation and carbon storage. Data will also help researchers understand how plant communities adjust to a changing climate/environment.

Plant economics run on carbon and nutrients instead of money. Leaf strategies aboveground span an economic spectrum from "live fast and die young" to "slow and steady," but the economy defined by root strategies belowground remains unclear. Here, we take a holistic view of the belowground economy and show that root-mycorrhizal collaboration can short circuit a one-dimensional economic spectrum, providing an entire space of economic possibilities. Root trait data from 1810 species across the globe confirm a classical fast-slow "conservation" gradient but show that most variation is explained by an orthogonal "collaboration" gradient, ranging from "do-it-yourself" resource uptake to "outsourcing" of resource uptake to mycorrhizal fungi. This broadened "root economics space" provides a solid foundation for predictive understanding of belowground responses to changing environmental conditions., (Copyright © 2020 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works. Distributed under a Creative Commons Attribution NonCommercial License 4.0 (CC BY-NC).)

Fine-root traits play key roles in ecosystem processes, but the drivers of fine-root trait diversity remain poorly understood. The plant economic spectrum (PES) hypothesis predicts that leaf and root traits evolved in coordination. Mycorrhizal association type, plant growth form and climate may also affect root traits. However, the extent to which these controls are confounded with phylogenetic structuring remains unclear. Here we compiled information about root and leaf traits for > 600 species. Using phylogenetic relatedness, climatic ranges, growth form and mycorrhizal associations, we quantified the importance of these factors in the global distribution of fine-root traits. Phylogenetic structuring accounts for most of the variation for all traits excepting root tissue density, with root diameter and nitrogen concentration showing the strongest phylogenetic signal and specific root length showing intermediate values. Climate was the second most important factor, whereas mycorrhizal type had little effect. Substantial trait coordination occurred between leaves and roots, but the strength varied between growth forms and clades. Our analyses provide evidence that the integration of roots and leaves in the PES requires better accounting of the variation in traits across phylogenetic clades. Inclusion of phylogenetic information provides a powerful framework for predictions of belowground functional traits at global scales., (© 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.)

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 > 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. 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., (© 2017 UT-Battelle LLC. New Phytologist © 2017 New Phytologist Trust.)

Background and Aims In Costa Rica, coffee (Coffea arabica) plants are often grown in agroforests. However, it is not known if shade-inducing trees reduce coffee plant biomass through root competition, and hence alter overall net primary productivity (NPP). We estimated biomass and NPP at the stand level, taking into account deep roots and the position of plants with regard to trees. Methods Stem growth and root biomass, turnover and decomposition were measured in mixed coffee/tree (Erythrina poeppigiana) plantations. Growth ring width and number at the stem base were estimated along with stem basal area on a range of plant sizes. Root biomass and fine root density were measured in trenches to a depth of 4 m. To take into account the below-ground heterogeneity of the agroforestry system, fine root turnover was measured by sequential soil coring (to a depth of 30 cm) over 1 year and at different locations (in full sun or under trees and in rows/inter-rows). Allometric relationships were used to calculate NPP of perennial components, which was then scaled up to the stand level. Key Results Annual ring width at the stem base increased up to 2·5 mm yr-1 with plant age (over a 44-year period). Nearly all (92 %) coffee root biomass was located in the top 1·5 m, and only 8 % from 1·5 m to a depth of 4 m. Perennial woody root biomass was 16 t ha-1 and NPP of perennial roots was 1·3 t ha-1 yr-1. Fine root biomass (0-30 cm) was two-fold higher in the row compared with between rows. Fine root biomass was 2·29 t ha-1 (12 % of total root biomass) and NPP of fine roots was 2·96 t ha-1 yr-1 (69 % of total root NPP). Fine root turnover was 1·3 yr-1 and lifespan was 0·8 years. Conclusions Coffee root systems comprised 49 % of the total plant biomass; such a high ratio is possibly a consequence of shoot pruning. There was no significant effect of trees on coffee fine root biomass, suggesting that coffee root systems are very competitive in the topsoil., (© The Author 2016. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Although fine roots are important components of the global carbon cycle, there is limited understanding of root structure-function relationships among species. We determined whether root respiration rate and decomposability, two key processes driving carbon cycling but always studied separately, varied with root morphological and chemical traits, in a coordinated way that would demonstrate the existence of a root economics spectrum (RES). Twelve traits were measured on fine roots (diameter ≤ 2 mm) of 74 species (31 graminoids and 43 herbaceous and dwarf shrub eudicots) collected in three biomes. The findings of this study support the existence of a RES representing an axis of trait variation in which root respiration was positively correlated to nitrogen concentration and specific root length and negatively correlated to the root dry matter content, lignin : nitrogen ratio and the remaining mass after decomposition. This pattern of traits was highly consistent within graminoids but less consistent within eudicots, as a result of an uncoupling between decomposability and morphology, and of heterogeneity of individual roots of eudicots within the fine-root pool. The positive relationship found between root respiration and decomposability is essential for a better understanding of vegetation-soil feedbacks and for improving terrestrial biosphere models predicting the consequences of plant community changes for carbon cycling., (© 2016 CNRS. New Phytologist © 2016 New Phytologist Trust.)

Background and Aims: Fine root decomposition is an important determinant of nutrient and carbon cycling in grasslands; however, little is known about the factors controlling root decomposition among species. Our aim was to investigate whether interspecific variation in the potential decomposition rate of fine roots could be accounted for by root chemical and morphological traits, life history and taxonomic affiliation. We also investigated the co-ordinated variation in root and leaf traits and potential decomposition rates., Methods: We analysed potential decomposition rates and the chemical and morphological traits of fine roots on 18 Mediterranean herbaceous species grown in controlled conditions. The results were compared with those obtained for leaves in a previous study conducted on similar species., Key Results: Differences in the potential decomposition rates of fine roots between species were accounted for by root chemical composition, but not by morphological traits. The root potential decomposition rate varied with taxonomy, but not with life history. Poaceae, with high cellulose concentration and low concentrations of soluble compounds and phosphorus, decomposed more slowly than Asteraceae and Fabaceae. Patterns of root traits, including decomposition rate, mirrored those of leaf traits, resulting in a similar species clustering., Conclusions: The highly co-ordinated variation of roots and leaves in terms of traits and potential decomposition rate suggests that changes in the functional composition of communities in response to anthropogenic changes will strongly affect biogeochemical cycles at the ecosystem level.