Search

Decomposition of dead fine roots contributes significantly to nutrient cycling and soil organic matter stabilization. Most knowledge of tree fine-root decomposition stems from studies in monospecific stands or single-species litter, although most forests are mixed. Therefore, we assessed how tree species mixing affects fine-root litter mass loss and which role initial litter quality and environmental factors play. For this purpose, we determined fine-root decomposition of 13 common tree species in four European forest types ranging from boreal to Mediterranean climates. Litter incubations in 315 tree neighborhoods allowed for separating the effects of litter species from environmental influences and litter mixing (direct) from tree diversity (indirect). On average, mass loss of mixed-species litter was higher than those of single-species litter in monospecific neighborhoods. This was mainly attributable to indirect diversity effects, that is, alterations in microenvironmental conditions as a result of tree species mixing, rather than direct diversity effects, that is, litter mixing itself. Tree species mixing effects were relatively weak, and initial litter quality and environmental conditions were more important predictors of fine-root litter mass loss than tree diversity. We showed that tree species mixing can alter fine-root litter mass loss across large environmental gradients, but these effects are context-dependent and of moderate importance compared to environmental influences. Interactions between species identity and site conditions need to be considered to explain diversity effects on fine-root decomposition. [ABSTRACT FROM AUTHOR]

Summary: The root economics space (RES) is multidimensional and largely shaped by belowground biotic and abiotic influences. However, how root–fungal symbioses and edaphic fertility drive this complexity remains unclear.Here, we measured absorptive root traits of 112 tree species in temperate and subtropical forests of China, including traits linked to functional differences between arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) hosts.Our data, from known mycorrhizal tree species, revealed a 'fungal‐symbiosis' dimension distinguishing AM from ECM species. This divergence likely resulted from the contrasting mycorrhizal evolutionary development of AM vs ECM associations. Increased root tissue cortical space facilitates AM symbiosis, whereas increased root branching favours ECM symbiosis. Irrespective of mycorrhizal type, a 'root‐lifespan' dimension reflecting aspects of root construction cost and defence was controlled by variation in specific root length and root tissue density, which was fully independent of root nitrogen content. Within this function‐based RES, we observed a substantial covariation of axes with soil phosphorus and nitrate levels, highlighting the role played by these two axes in nutrient acquisition and conservation.Overall, our findings demonstrate the importance of evolved mycorrhizal symbiosis pathway and edaphic fertility in framing the RES, and provide theoretical and mechanistic insights into the complexity of root economics. [ABSTRACT FROM AUTHOR]

Summary: 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. [ABSTRACT FROM AUTHOR]

Summary: 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. [ABSTRACT FROM AUTHOR]

Mixed‐species forests have often been shown to enhance above‐ground ecosystem properties and processes. Despite the significance of fine roots for tree and ecosystem functioning, the role of tree species diversity for below‐ground processes driven by fine roots remains largely unknown. Previously, an underyielding of fine‐root biomass (FRB) in tree mixtures across four major European forest types has been reported. To explain this phenomenon, we tested here the effect of tree species mixing on fine‐root traits related to soil exploitation efficiency, including biotic feedbacks from ectomycorrhizal fungi (EcM), and assessed the role of root trait dissimilarity.We analysed morphological and chemical traits as well as ectomycorrhizal colonisation intensity of absorptive fine roots (i.e. first three most distal orders) in soil samples from 315 mixed and mono‐specific tree neighbourhoods in mainly mature, semi‐natural forest stands across Europe. Additionally, we quantified mycorrhizal abundance and diversity in soil samples from the same stands.At the community level, fine roots in tree mixtures were characterised by higher specific root lengths and root nitrogen concentrations, lower diameters and root tissue densities indicating a faster resource acquisition strategy compared to mono‐specific stands. The higher root EcM colonisation intensity and soil EcM diversity in mixtures compared to mono‐specific stands may further provide evidence for positive biotic feedbacks. Moreover, the diversity of fine‐root traits influenced FRB, as mixtures characterised by a higher trait dissimilarity were linked to a lower reduction in FRB. At the level of phylogenetic groups, thin‐rooted angiosperm species showed stronger responses to mixing than thick‐rooted gymnosperms, especially in terms of root morphology and EcM colonisation, indicating different strategies of response to tree mixing.Our results indicate that a lower FRB can reflect a shift in soil resource acquisition strategies, rather than a lower performance of trees in mixtures. They show that several non‐exclusive mechanisms can simultaneously explain negative net effects of mixing on FRB. This study sheds new light on the importance of using integrative approaches including both above‐ and below‐ground biomass and traits to study diversity effects on plant productivity. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Intraspecific trait variation is an important driver of plant performance in different environments. Although roots acquire essential resources that vary with the environment, most studies have focused on intraspecific variation in leaf traits, and research on roots is often restricted to a few species. It remains largely unclear how and to what extent root traits vary with the environment and whether general intraspecific patterns exist across species.We compared intraspecific variation in specific root length (SRL), root diameter, root tissue density (RTD) and root branching density of 11 species along a 1,000 m elevation gradient in the French Alps. We tested (a) the extent of intraspecific versus interspecific root trait variation along the gradient, (b) whether intraspecific trait patterns with elevation were consistent among species and (c) whether environmental variables better explained intraspecific variation in root traits than elevation. Specifically, we hypothesised that within a species, root trait values would adjust to enhance resource acquisition (either through an increase in SRL or root diameter, and/or in branching density) and/or conservation (increased RTD) at higher elevations.Species identity explained most of the overall variation in root traits. Elevation explained only a minor proportion of intraspecific root trait variation, which was larger within than between elevations. Also, trait relationships with elevation rarely agreed with our hypotheses, varied strongly across species and were often differently related to environmental variation. Generally, climate, soil and vegetation properties better explained intraspecific root variation than elevation, but these relationships were highly species‐dependent.Along complex environmental gradients where multiple properties simultaneously change, roots of different species vary in different ways, leading to species‐specific patterns in intraspecific root trait variation. The lack of support for our hypotheses may be caused by the multiple interactions between environmental properties, small‐scale soil heterogeneity, species phylogeny and changing plant–plant interactions. Our findings suggest that, to enhance our understanding of the effects of environmental change on plant performance, we need to better integrate the multiple dimensions of plant responses to change and measure a broader set of root traits and environmental variables. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Fine roots play key roles in the capacity of plants to face environmental constraints and their traits reflect adaptations to the environment, including soil structure, resource availability and climate. However, the inaccuracy of global soil and climate databases to account for the large environmental variation occurring at small spatial scale prevents accurate estimations of the linkages between environmental variables and fine‐root strategies.Here, using two global databases on fine‐root traits (Rhizopolis‐db) and species phylogenetic relatedness, and a regional database of species ecological indicator values (Baseflor), we quantified the predictive value of ecological indicator values, as an alternative to classical coarse soil and climate indicators, on the variation in four major fine‐root traits.A strong phylogenetic signal was found among species for fine‐root mean diameter, specific root length (SRL) and root tissue density (RTD), but less so for root nitrogen concentration (RNC). After accounting for this relatedness, ecological indicators still explained a large part of trait variation in our dataset for SRL, RTD and RNC. Multi‐indicators best model R2 reached.40 for SRL and RTD, and.44 for RNC, whereas it was only 0.10 for diameter. Ecological indicators of nutrient availability and soil texture were those that most strongly related to SRL, RTD and RNC. Specifically, plant fast resources use strategies characterized by high SRL, RNC and low RTD occurred more frequently in nutrient‐rich soils and in soils with light sandy textures. Additionally, light availability and atmospheric temperature were negatively related with SRL and continentality negatively influenced RNC.With respect to both nutrient and water availability ecological indicator values, opposite adaptations were observed between growth forms, particularly between woody and herbaceous species, limiting our ability to define simple, widely applicable patterns of trait–environment relationships.Synthesis. Our analysis demonstrates that species ecological indicator values are valuable predictors of plant below‐ground strategies. It provides original evidence that herbaceous species with fine‐root traits representative of fast resource use strategies typically occur in more favourable soil habitats (high nutrient and water availability); meanwhile, woody species may show the opposite trend. Other important environmental parameters concomitantly influence fine‐root trait variation in contrasting ways. [ABSTRACT FROM AUTHOR]

Summary: Plants respond to resource stress by changing multiple aspects of their biomass allocation, morphology, physiology and architecture. To date, we lack an integrated view of the relative importance of these plastic responses in alleviating resource stress and of the consistency/variability of these responses among species. We subjected nine species (legumes, forbs and graminoids) to nitrogen and/or light shortages and measured 11 above‐ground and below‐ground trait adjustments critical in the alleviation of these stresses (plus several underlying traits). Nine traits out of 11 showed adjustments that improved plants’ potential capacity to acquire the limiting resource at a given time. Above ground, aspects of plasticity in allocation, morphology, physiology and architecture all appeared important in improving light capture, whereas below ground, plasticity in allocation and physiology were most critical to improving nitrogen acquisition. Six traits out of 11 showed substantial heterogeneity in species plasticity, with little structuration of these differences within trait covariation syndromes. Such comprehensive assessment of the complex nature of phenotypic responses of plants to multiple stress factors, and the comparison of plant responses across multiple species, makes a clear case for the high (but largely overlooked) diversity of potential plastic responses of plants, and for the need to explore the potential rules structuring them. [ABSTRACT FROM AUTHOR]

Litter chemical traits are one of the dominant controls on litter decomposition. Increasing atmospheric nitrogen (N) deposition is expected to alter litter chemical traits at the community level in both direct (altering intraspecific chemistry) and indirect ways (changing species abundance and composition). Compared to intraspecific changes, the role of changes in species composition in driving the responses of litter chemical traits to N enrichment has been seldom quantitatively addressed. We quantified the relative contribution of intraspecific changes versus changes in community composition on litter traits and how this would be influenced by the magnitude of N deposition by taking advantage of a long-term field N addition experiment in a semi-arid grassland with a wide range of N addition rates. Nitrogen deposition altered plant species abundance by facilitating the dominance of one species with a nutrient acquisitive strategy, producing higher quality litter and being more responsive to N addition at the intraspecific level. Overall, changes in species composition, intraspecific changes and their interaction all led to higher litter quality (higher N and lower lignin, cellulose and hemicellulose concentrations) under N deposition treatments. The relative contribution of species composition on the responses of litter chemical traits to N deposition also increased with N addition rate, ranging from 5 to 40% for litter N, and from 2 to ~ 30% for the three structural carbon components. Our results demonstrate the positive impacts of increasing N deposition on litter quality through changing intraspecific C and N chemistry and species turnover, which has potential consequences for litter decomposition and nutrient cycling in ecosystems. Further, we highlight the important contribution of shifts in species abundance to the plant-mediated biogeochemical responses to N deposition. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Roots vary in anatomy, morphology and physiology, both spatially (different parts of the same root system) and temporally (plastic changes, root ageing), suggesting that root trait measurements are strongly affected by root sampling categories., In this context, it is urgent to clarify the functional significance of current root sampling categories (e.g. fine roots of the first order, the first three orders, ≤1 mm or ≤2 mm), establish guidelines for choosing between sampling methods and revise root ontology to account for functional differences between traits measured on distinct root categories., Here, we used a worldwide database of fine-root traits to test the hypothesis that distinct fine-root trait values - with link to fine-root functions - were generally affected by different root sampling categories. We observed indeed a clear functional break between first-order roots and roots of all three other sampling categories, and a smaller but substantial break between roots of the three first orders and the ≤2 mm category, demonstrating globally that different sampling methodologies capture different functional parts of roots., Our synthesis suggests that all current root sampling categories present both advantages and pitfalls and that no single method can appropriately tackle the main current challenge of root functional ecology: i.e. linking fine roots to plant and ecosystem functions in a truly comparable way across all plants. We argue instead that a small set of complementary standardized sampling methods is necessary to capture the linkages between root forms and functions., To assist experimenters selecting adequate sampling we developed a decision table following three logical questions: (i) what plant or ecosystem function must be addressed; (ii) what root categories are involved in this function and (iii) what traits should be measured on these root categories. Challenging, strengthening and expending such common reference framework would be a substantial step towards wider comparability of future functional trait datasets., A is available for this article. [ABSTRACT FROM AUTHOR]

Plants adapt phenotypically to different conditions of light and nutrient supply, supposedly in order to achieve colimitation of these resources. Their key variable of adjustment is the ratio of leaf area to root length, which relies on plant biomass allocation and organ morphology., We recorded phenotypic differences in leaf and root mass fractions ( LMF, RMF), specific leaf area ( SLA) and specific root length ( SRL) of 12 herbaceous species grown in factorial combinations of high/low irradiance and fertilization treatments., Leaf area and root length ratios, and their components, were influenced by nonadditive effects between light and nutrient supply, and differences in the strength of plant responses were partly explained by Ellenberg's species values representing ecological optima. Changes in allocation were critical in plant responses to nutrient availability, as the RMF contribution to changes in root length was 2.5× that of the SRL. Contrastingly, morphological adjustments ( SLA rather than LMF) made up the bulk of plant response to light availability., Our results suggest largely predictable differences in responses of species and groups of species to environmental change. Nevertheless, they stress the critical need to account for adjustments in below-ground mass allocation to understand the assembly and responses of communities in changing environments. [ABSTRACT FROM AUTHOR]

Questions It remains unresolved why, despite the obvious functional importance of leaves and roots, co-existing plant species can display highly contrasting biomass distributions of these organs. Building on the 'functional equilibrium' hypothesis, we hypothesize that co-existing species can each achieve balanced resource acquisition above vs below ground by trading off the biomass vs morphology of structures responsible for resource acquisition, i.e. leaves and fine roots. Methods We tested this hypothesis in a natural field setting by measuring plant above- and below-ground biomass and morphological traits associated with resource uptake - specific leaf area ( SLA) and specific root length ( SRL) - of 18 dominant angiosperm species from a sub-alpine plant community. Location New Zealand South Island. Results We found a significant negative relationship between the species leaf mass to fine root mass ratio and the SLA to SRL ratio when we considered eudicot species only. The SLA to SRL ratio and plant size explained 31% and 34% of the total variation in the species leaf to fine root mass ratio respectively, and 45% when used in combination ( P < 0.05 in all cases). Within a given plant size, 90% of the variation among species in total leaf area was due to differences in SLA, whereas variation in the fine root mass fraction was responsible for 71% of the variation among species in fine root length. Conclusions In support of our hypothesis, part of the difference between co-occurring species in leaf and fine root biomass distribution could be explained by the variable morphologies of these organs as well as variation in plant size, independent of the plant economic strategy. We expect that this outcome may result from environmental and evolutionary constraints on plant species-average traits, as well as plastic responses to local environmental conditions. These findings help explain why a diversity of strategies for achieving balanced resource acquisition can co-exist within a single plant community. [ABSTRACT FROM AUTHOR]

The 'home-field advantage (HFA) hypothesis' predicts that plant litter is decomposed faster than expected in the vicinity of the plant where it originates from (i.e. its 'home') relative to some other location (i.e. 'away') because of the presence of specialized decomposers. Despite growing evidence for the widespread occurrence HFA effects, what drives HFA is not understood as its strength appears highly variable and context-dependent. Our work advances current knowledge about HFA effects by testing under what conditions HFA is most important. Using published data on mass loss from 125 reciprocal litter transplants from 35 studies, we evaluated if HFA effects were modulated by macroclimate, litter quality traits, and the dissimilarity between 'home' and 'away' of both the quality of reciprocally exchanged litters and plant community type. Our results confirmed the occurrence of an overall, worldwide, HFA effect on decomposition with on average 7.5% faster decomposition at home. However, there was considerable variation in the strength and direction (sometimes opposite to expectations) of these effects. While macroclimate and average litter quality had weak or no impact on HFA effects, home-field effects became stronger (regardless of the direction) when the quality of 'home' and 'away' litters became more dissimilar (e.g. had a greater dissimilarity in N:P ratio; F1,42 = 6.39, p = 0.015). Further, home-field effects were determined by the degree of difference between the types of dominant plant species in the 'home' versus 'away' communities (F2,105 = 4.03, p = 0.021). We conclude that home-field advantage is not restricted to particular litter types or climate zones, and that the dissimilarity in plant communities and litter quality between the 'home' and 'away' locations, are the most significant drivers of home-field effects. [ABSTRACT FROM AUTHOR]

Human activities that involve land-use change often cause major transforma-tions to community and ecosystem properties both aboveground and belowground, and when land use is abandoned, these modifications can persist for extended periods. However, the mechanisms responsible for rapid recovery vs. long-term maintenance of ecosystem changes following abandonment remain poorly understood. Here, we examined the long-term ecological effects of two remote former settlements, regularly visited for --300 years by reindeer-herding Sami and abandoned ~100 years ago, within an old-growth boreal forest that is considered one of the most pristine regions in northern Scandinavia. These human legacies were assessed through measurements of abiotic and biotic soil properties and vegetation characteristics at the settlement sites and at varying distances from them. Low-intensity land use by Sami is characterized by the transfer of organic matter towards the settlements by humans and reindeer herds, compaction of soil through trampling, disappearance of understory vegetation, and selective cutting of pine trees for fuel and construction. As a consequence, we found a shift towards early successional plant species and a threefold increase in soil microbial activity and nutrient availability close to the settlements relative to away from them. These changes in soil fertility and vegetation contributed to 83% greater total vegetation productivity, 35% greater plant biomass, and 23% and 16% greater concentrations of foliar N and P nearer the settlements, leading to a greater quantity and quality of litter inputs. Because decomposer activity was also 40% greater towards the settlements, soil organic matter cycling and nutrient availability were further increased, leading to likely positive feedbacks between the aboveground and belowground components resulting from historic land use. Although not all of the activities typical of Sami have left visible residual traces on the ecosystem after 100 years, their low-intensity but long-term land use at settlement sites has triggered a rejuvenation of the ecosystem that is still present. Our data demonstrates that aboveground-belowground interactions strongly control ecosystem responses to historical human land use and that medium- to long-term consequences of even low-intensity human activities must be better accounted for if we are to predict and manage ecosystems succession following land-use abandonment. [ABSTRACT FROM AUTHOR]

Conceptual frameworks relating plant traits to ecosystem processes such as organic matter dynamics are progressively moving from a leaf-centred to a whole-plant perspective. Through the use of meta-analysis and global literature data, we quantified the relative roles of litters from above- and below-ground plant organs in ecosystem labile organic matter dynamics., We found that decomposition rates of leaves, fine roots and fine stems were coordinated across species worldwide although less strongly within ecosystems. We also show that fine roots and stems had lower decomposition rates relative to leaves, with large differences between woody and herbaceous species. Further, we estimated that on average below-ground litter represents approximately 33 and 48% of annual litter inputs in grasslands and forests, respectively., These results suggest a major role for below-ground litter as a driver of ecosystem organic matter dynamics. We also suggest that, given that fine stem and fine root litters decompose approximately 1.5 and 2.8 times slower, respectively, than leaf litter derived from the same species, cycling of labile organic matter is likely to be much slower than predicted by data from leaf litter decomposition only., Synthesis. Our results provide evidence that within ecosystems, the relative inputs of above- versus below-ground litter strongly control the overall quality of the litter entering the decomposition system. This in turn determines soil labile organic matter dynamics and associated nutrient release in the ecosystem, which potentially feeds back to the mineral nutrition of plants and therefore plant trait values and plant community composition. [ABSTRACT FROM AUTHOR]

Coevolution has driven speciation and evolutionary novelty in functional traits across the Tree of Life. Classic coevolutionary syndromes such as plant–pollinator, plant–herbivore, and host–parasite have focused strongly on the fitness consequences during the lifetime of the interacting partners. Less is known about the consequences of coevolved traits for ecosystem-level processes, in particular their 'afterlife' legacies for litter decomposition, nutrient cycling, and the functional ecology of decomposers. We review the mechanisms by which traits resulting from coevolution between plants and their consumers, microbial symbionts, or humans, and between microbial decomposers and invertebrates, drive plant litter decomposition pathways and rates. This supports the idea that much of current global variation in the decomposition of plant material is a legacy of coevolution. Plant litter decomposition adds a different process and ecological context to the coevolution literature, which has thus far focused on the ecology of symbionts during their lifetimes. This context integrates the literature on how arms races between plants and their consumers (herbivores, pathogens) or mutualists (nitrogen-fixing bacteria, mycorrhizal fungi) drive the quality of plant tissue, with an important legacy for litter decomposability. We know little about recent coevolutionary decomposition legacies involving plant domestication by humans via farming, plant breeding, and modified landscapes with feedback to evolution of human brain, digestion, and dentition. How myriad coevolutionary links between invertebrates and microbial decomposers affect global carbon cycling needs further investigation. Overall, plant decomposition rates are largely the legacy of wide-ranging coevolutionary relationships. [ABSTRACT FROM AUTHOR]

1. Evidence is growing that leaf litter generally decomposes faster than expected in its environment of origin, owing to specialization of litter and topsoil decomposer communities to break down litter encountered most often. Nevertheless, this home-field advantage (HFA) in decomposition is inconsistently supported by experimental data and fails to account for situations where contrasting qualities of litter coexist within the same litter matrix. 2. In contrast to the HFA hypothesis, which expects a positive interaction between every litter species produced locally and the local decomposer communities irrespective of litter species quality, we define here an alternative substrate quality-matrix quality interaction (SMI) hypothesis that expects a continuum from positive to negative interaction between specific litters (substrates) and decomposer communities as specific litters and the ecosystem litter layer (i.e. the matrix, which drives local decomposer community composition) become increasingly dissimilar in quality. 3. To test this hypothesis, we conducted a reciprocal transplant decomposition experiment of eight leaf, six fine-stem and nine fine-root litter species from three neighbouring ecosystems of the subarctic biome: dry forest, riparian forest and forest-surrounded pond; and characterized the quality (represented by lignin content and an integrated measure of carbon/nutrient economics) of each litter species and each ecosystem litter layer. 4. We found substantial overall effects of SMI on decomposition rates of leaf (20% explained variance), stem (14%) and root (15%) litters, although this effect was lower than the single effects of litter quality and microclimate (remaining explained variance). Despite being partly inconsistent across litter species, likely due to the complexity of litter quality-decomposer community relationships, the SMI hypothesis appeared more broadly applicable than the HFA hypothesis. 5. Synthesis. We demonstrate here that plant traits, likely via their control on litter and topsoil decomposer community composition, have indirect effects on litter breakdown rates, not only at the interface between ecosystems but also within ecosystems, with likely implications for many other ecosystems world-wide. These results suggest functional variation in decomposer communities between ecosystems with respect to their efficiency to degrade litters with contrasting qualities, such as different lignolytic and detoxification activities but also contrasting efficiencies to degrade non-recalcitrant tissues. [ABSTRACT FROM AUTHOR]

Summary 1. Recent evidence indicates tight control of plant resource economics over interspecific trait variation amongst species, both within and across organs, referred to as 'plant economics spectrum' (PES). Whether and how these coordinated whole-plant economics strategies can influence the decomposition system and thereby impact on ecosystem carbon and nutrient cycling are yet an open question. More specifically, it is yet unknown whether plant functional traits have consistent afterlife effects across different plant organs. 2. To answer those questions, we conducted a common-garden decomposition experiment bringing together leaves, fine stems, coarse stems, fine roots and reproductive parts from a wide range of subarctic plant types, clades and environments. We measured all plant parts for the same (green and litter) plant economics traits and identified a whole-plant axis of carbon and nutrient economics. 3. We demonstrated that our local 'PES' has important afterlife effects on carbon turnover by driving coordinated decomposition rates of different organs across species. All organ decomposabilities were consistently controlled by the same structure-related traits (lignin, C and dry matter content) whilst nutrient-related traits (N, P, pH, phenols) had more variable influence, likely due to their contrasting functions across organs. Nevertheless, consistent shifts in elevation of parallel trait-decomposition relationships between organs indicate that other variables, potentially related to organ dimensions, configuration or chemical contents, codetermine litter decomposition rates. 4. Whilst the coordinated litter decomposabilities across species organs imply a coordinated impact of plant above-ground and below-ground litters on plant-soil feedbacks, the contrasting decomposabilities between plant parts suggest a major role for the relative inputs of organ litter as driver of soil properties and ecosystem biogeochemistry. These relationships, underpinning the afterlife effects of the PES on whole-plant litter decomposability, will provide comprehensive input of vegetation composition feedback to soil carbon turnover. [ABSTRACT FROM AUTHOR]

Summary 1. The potential resorption of substantial amounts of nutrients from all vegetative organs of plants has large implications for the plant nutrient economy and for biogeochemical cycles. So far, most studies have focused on leaf nutrient resorption only. Besides, while evidence is growing that soil fertility changes impact on leaf nutrient resorption at a large spatial scale, hardly anything is known of such coupling at a small spatial scale. 2. Here we show that nitrogen (N) in culms of four dominant grasses of northern Chinese steppes contributed from 17% to 36% to the total pool of N resorbed from above-ground senescing parts and accounted for 25-52% of above-ground litter N. These results demonstrate the tremendous importance of non-leaf organs for plant nutrient economy and ecosystem nutrient cycling. 3. More importantly, we found that even microscale variations in resource availability (soil inorganic N; soil moisture) can strongly impact on both leaf and culm N resorption proficiencies (RP) and absolute leaf N resorption of grasses. Moreover, plasticity was responsible for 86% and 43% of within-site variance in leaf and culm RP, respectively, with the remainder owing to interspecific differences between the four grasses. These results imply a much larger role of plant plasticity in driving ecosystem functioning than previously assumed. 4. Synthesis. Our results suggest that plant litter quality varies even at the microscale with heterogeneity in soil resource availability, thereby potentially feeding back on soil properties and sustaining microscale soil fertility patchiness. In parallel, plants of more fertile patches resorbed a greater absolute amount of N, likely benefiting their competitive and reproductive abilities. [ABSTRACT FROM AUTHOR]

Summary 1. While the importance of wood decay for the global carbon balance is widely recognized, surprisingly little is known about its long-term dynamics and its abiotic and biotic drivers. Progress in this field is hindered by the long time-scales inherent to the low decay rates of wood and the lack of short-term methods to assess long-term decomposition dynamics in standardized field conditions. 2. Here, we present such a method, which relies on the sampling and short-term incubation of wood from several decay stages covering the entire decay process. Together these short-term decay steps are used to model and discriminate between three potential decay dynamics (linear, exponential and sigmoid) using an iterative optimization procedure. We applied this method to analyse long-term wood decay of six subarctic tree species (six stems and two roots) and test the hypotheses that (i) different wood species follow distinct decay dynamics and (ii) interspecific variation in wood traits controls variation in wood decay rates in a standardized environment. 3. We found interspecific variation in long-term wood decay dynamics: decay of Alnus and Salix stems was best described by exponential models, whereas decay of Sorbus stems and Betula and Pinus roots was best fitted by linear models and Betula, Pinus and Populus stems each displayed a sigmoid decay dynamics (up to 5-year initial lag phase). A six-fold variation was observed between the decomposition half-lives of all eight wood types, from 6.8 years (6.1-7.5, 95% C.I.) for Alnus stems to 41.3 years (34.5-51.8) for Pinus roots. Initial wood traits such as pH ( R2 = 0.92), dry matter content ( R2 = 0.79) and lignin ( R2 = 0.73) were good predictors of long-term wood decay rates. 4. Synthesis. Our findings suggest changing decay dynamics across wood species and types that are likely to arise from changing underlying wood decay processes (i.e. varying wood functional traits/decomposer community interactions). Our new method, which combines advantages of direct observations and the chronosequence approach, allows reliable comparisons of species contributions to long-term wood decay rates and provides future opportunities to experimentally disentangle intrinsic and external abiotic and biotic drivers of long-term wood decay processes. [ABSTRACT FROM AUTHOR]

ABSTRACT Aim The drivers of species assembly, by limiting the possible range of functional trait values, can lead to either convergent or divergent distributions of traits in realized assemblages. Here, to evaluate the strengths of these species assembly drivers, we partition trait variance across global, regional and community scales. We then test the hypothesis that, from global to community scales, the outcome of co-occurring trait convergence and divergence is highly variable across biomes and communities. Location Global: nine biomes ranging from subarctic highland to tropical rain forest. Methods We analysed functional trait diversity at progressively finer spatial scales using a global, balanced, hierarchically structured dataset from 9 biomes, 58 communities and 652 species. Analyses were based on two key leaf traits (foliar nitrogen content and specific leaf area) that are known to drive biogeochemical cycling. Results While 35% of the global variance in these traits was between biomes, only 15% was between communities within biomes and as much as 50% occurred within communities. Despite this relatively high within-community variance in trait values, we found that trait convergence dominated over divergence at both global and regional scales through comparisons of functional trait diversity in regional and community assemblages against random (null) models of species assembly. Main conclusions We demonstrate that the convergence of traits occurring from global to regional assemblages can be twice as strong as that from regional to community assemblages, and argue that large differences in the nature and strength of abiotic and biotic drivers of dominant species assembly can, at least partly, explain the variable outcome of simultaneous trait convergence and divergence across sites. Ultimately, these findings stress the urgent need to extend species assembly research to address those scales where trait variance is the highest, i.e. between biomes and within communities. [ABSTRACT FROM AUTHOR]

Variation in plant functional traits is the product of evolutionary and environmental drivers operating at different scales. Little is known about whether, or how, this variation is coordinated between aboveground and belowground organs across and within spatial scales., We address these questions using a hierarchically designed dataset of pairwise leaf and root traits related to carbon and nutrient economy of 64 species belonging to 14 plant communities in northern Chinese semi-arid and arid regions., While both root and leaf traits showed most of their variance among (individuals and) species within communities, leaf trait variance tended to be relatively higher at coarser spatial scales than root trait variance. While leaf nitrogen (N) per area to root N per length ratio increased and specific leaf area to specific root length and leaf [N] to root [N] ratios decreased from semi-arid to arid environments owing to climatic/edaphic shifts, the matching pairs showed a strong pattern of positive correlation that was upheld across spatial scales and geographic areas., Thus, trade-offs in plant resource investment across organs within individual vascular plants are constrained within a rather narrow range of variation. A new challenge will be to test whether and how such trait coordination is also seen within and across other biomes of the world. [ABSTRACT FROM AUTHOR]

•Nutrient resorption and leaching resistance, through their roles in reducing nutrient losses, are important determinants of plant nutrient economy. However, the contributions of fine-stem and fine-root resorption, as well as leaf leaching resistance, have largely been overlooked. •We quantified the relative contributions of these processes to nutrient depletion of these organs during their senescence using 40 subarctic vascular species from aquatic, riparian and terrestrial environments. We hypothesized that interspecific variation in organ nutrient resorption and leaf leaching would be linked to the species’ nutrient acquisitive-conservative strategies, as quantified for a set of common-organ nutrient/carbon economics traits. •The subarctic flora generally had both high resistance to leaching and high internal nutrient recycling. Average nutrient resorption efficiencies were substantial for leaves (nitrogen (N), 66 ± 3% SE; phosphorus (P), 63 ± 4%), fine stems (N, 48 ± 4%; P, 56 ± 4%) and fine roots (N, 27 ± 7%; P, 57 ± 6%). The link between nutrient resorption and other nutrient/carbon economics traits was very weak across species, for all three organs. •These results emphasize the potential importance of resorption processes for the plant nutrient budget. They also highlight the idiosyncrasies of the relationship between resorption processes and plant economics, which is potentially influenced by several plant physiological and structural adaptations to environmental factors other than nutrient stress. [ABSTRACT FROM AUTHOR]

1. A fundamental trade-off among vascular plants between traits inferring rapid resource acquisition and those leading to conservation of resources has now been accepted broadly, but is based on empirical data with a strong bias towards leaf traits. Here, we test whether interspecific variation in traits of different plant organs obeys this same trade-off and whether within-plant trade-offs are consistent between organs. 2. Thereto, we measured suites of the same chemical and structural traits from the main vegetative organs for a species set representing aquatic, riparian and terrestrial environments including the main vascular higher taxa and growth forms of a subarctic flora. The traits were chosen to have consistent relevance for plant defence and growth across organs and environments: carbon, nitrogen, phosphorus, lignin, dry matter content, pH. 3. Our analysis shows several new trait correlations across leaves, stems and roots and a striking pattern of whole-plant integrative resource economy, leading to tight correspondence between the local leaf economics spectrum and the root ( r = 0.64), stem ( r = 0.78) and whole-plant ( r = 0.93) economics spectra. 4. Synthesis. Our findings strongly suggest that plant resource economics is consistent across species’ organs in a subarctic flora. We provide thus the first evidence for a ‘plant economics spectrum’ closely related to the local subarctic ‘leaf economics spectrum’. Extending that concept to other biomes is, however, necessary before any generalization might be made. In a world facing rapid vegetation change, these results nevertheless bear considerable prospects of predicting below-ground plant functions from the above-ground components alone. [ABSTRACT FROM AUTHOR]

Low available soil nitrogen (N) limits plant productivity in alpine regions, and alpine plants thus resorb and reallocate N from senescing tissues to conserve this limited N during the non‐growing season. However, the destination and extent of N redistribution during plant senescence among above‐ and below‐ground organs, let alone other processes of translocation outside of plants and into the soil components, remain poorly understood.Utilizing 15N stable isotope as a tracer, we quantified N redistribution among above‐ and below‐ground plant organs and different soil components during senescence in an alpine meadow ecosystem, and explored the relationship between 15N partitioning among plant–soil N pools with seasonal fluctuations of plant functional traits.We found a substantial depletion of 15N in fine roots (−40% ± 2.8%) and above‐ground tissues (−51% ± 5.1%), and an enhanced 15N retention primarily in coarse roots (+79% ± 27%) and soil organic matter (+37% ± 10%) during plant senescence, indicating a dual role of roots with coarse roots acting as an N sink and fine roots as a source of N recycling during senescence. In parallel, we observed a temporal variation in plant functional traits, representing a shift from more acquisitive to more conservative strategies as the growing season ends, such as higher coarse root N and coarse root to fine root ratio. The seasonal trait variations were highly correlated with the 15N retention in coarse roots and soil organic matter. Particularly, 15N retention in particulate and mineral‐associated organic matter increased by 30% ± 12% and 24% ± 9%, respectively, suggesting a potential pathway through which fine root and microbial mortality contribute to 15N redistribution into soil N pools during senescence.Synthesis. N redistribution and seasonal plant trait fluctuation facilitate plant N conservation and ecosystem N retention in the alpine system. This study suggests a coupled above‐ground‐below‐ground N conservation strategy that may optimize the temporal coupling between plant N demand and ecosystem N supply in N‐limited alpine ecosystems. [ABSTRACT FROM AUTHOR]

Summary: Plants often associate with specialized decomposer communities that increase plant litter breakdown, a phenomenon that is known as the 'home‐field advantage' (HFA). Although the concept of HFA has long considered only the role of the soil microbial community, explicit consideration of the role of the microbial community on the foliage before litter fall (i.e. the phyllosphere community) may help us to better understand HFA. We investigated the occurrence of HFA in the presence vs absence of phyllosphere communities and found that HFA effects were smaller when phyllosphere communities were removed. We propose that priority effects and interactions between phyllosphere and soil organisms can help explain the positive effects of the phyllosphere at home, and suggest a path forward for further investigation. [ABSTRACT FROM AUTHOR]

Mixed‐species forests often enhance the provision of ecosystem functions, both above‐ and below‐ground. Several of these effects are mediated by the amount and spatial distribution of tree tissues. However, previous studies on tree diversity effects on fine‐root biomass (FRB) have returned inconsistent results and did not distinguish between absorptive and transport fine roots. Furthermore, owing to the lack of species‐specific data, it is not well understood whether complementarity or selection effects contribute more to these mixing effects.Here, we analysed tree species mixing effects on fine‐root traits while considering the respective tree species contributions and root functional types. Specifically, we tested whether tree species mixing increases FRB and root length density (RLD) and results in vertical root stratification. We quantified FRB and RLD in 30‐cm deep soil profiles for 13 tree species in mixed and pure stands across four widespread European forest types. The differentiation of different fine‐root species in mixtures allowed us to disentangle complementarity and selection effects.Across all sites, mixtures supported on average less FRB than pure stands, which was reflected in negative complementarity and selection effects. RLD of absorptive fine roots did not change across the soil profile and even increased in the topsoil, which was associated with positive complementarity effects. There was no evidence for vertical root stratification. Conifer proportion, which was mainly associated with selection effects, dampened net diversity effects. Root functional type further influenced tree species mixing effects.Synthesis. Despite the underyielding of FRB in mixtures, overall soil occupation by absorptive fine roots (RLD) did not decrease in mixtures, pointing to morphological root trait adaptations associated with higher resource‐use efficiency. Increased RLD in the most nutrient‐rich layer in mixtures further indicates complementary interactions among species and a greater resource uptake capacity. This work illustrates that considering only one aspect of trait‐functioning relationships, for example, root biomass, may not capture the full effect of plant diversity on ecosystem functioning. The integration of a larger range of relevant traits is required. Moreover, traditional classification of fine roots based on the 2‐mm diameter cut‐off may obscure responses of roots to environmental changes. [ABSTRACT FROM AUTHOR]

Motivation: Trait data are fundamental to the quantitative description of plant form and function. Although root traits capture key dimensions related to plant responses to changing environmental conditions and effects on ecosystem processes, they have rarely been included in large‐scale comparative studies and global models. For instance, root traits remain absent from nearly all studies that define the global spectrum of plant form and function. Thus, to overcome conceptual and methodological roadblocks preventing a widespread integration of root trait data into large‐scale analyses we created the Global Root Trait (GRooT) Database. GRooT provides ready‐to‐use data by combining the expertise of root ecologists with data mobilization and curation. Specifically, we (a) determined a set of core root traits relevant to the description of plant form and function based on an assessment by experts, (b) maximized species coverage through data standardization within and among traits, and (c) implemented data quality checks. Main types of variables contained: GRooT contains 114,222 trait records on 38 continuous root traits. Spatial location and grain: Global coverage with data from arid, continental, polar, temperate and tropical biomes. Data on root traits were derived from experimental studies and field studies. Time period and grain: Data were recorded between 1911 and 2019. Major taxa and level of measurement: GRooT includes root trait data for which taxonomic information is available. Trait records vary in their taxonomic resolution, with subspecies or varieties being the highest and genera the lowest taxonomic resolution available. It contains information for 184 subspecies or varieties, 6,214 species, 1,967 genera and 254 families. Owing to variation in data sources, trait records in the database include both individual observations and mean values. Software format: GRooT includes two csv files. A GitHub repository contains the csv files and a script in R to query the database. [ABSTRACT FROM AUTHOR]

Manipulating plant functional diversity to improve agroecosystem multifunctionality is a central challenge of agricultural systems world‐wide. In cocoa agroforestry systems (cAFS), shade trees are used to supply many services to farmers, yet their impact on soil functioning and cocoa yields is likely to vary substantially among tree species.Here we compared the impact of five shade tree species (Canarium schweinfurthii (Canarium), Dacryodes edulis (Safou), Milicia excelsa (Iroko), Ceiba pentandra (Kapok tree), Albizia adianthifolia (Albizia)) and unshaded conditions on the functioning of poor sandy savanna soils within eight cocoa farms in Central Cameroon. We assessed the effects of plant functional traits, leaf litterfall and fine root biomass on a range of soil functions and on cocoa yield.Shade trees generally improved soil pH, NH4+, NO3- and Olsen P content, biomass production of bioassays and soil total C and N content, while leaving cocoa yields unchanged. However, these effects varied largely among species. Improvements of soil functions were low under the two fruit trees (Canarium and Dacryodes), medium under the legume tree Albizia and high under the two timber trees (Milicia and Ceiba). Low litter recalcitrance was most strongly associated with increases in soil fertility indicators such as N and P availability, whereas soil C and N content increased with litter Ca restitution.Synthesis and applications. We demonstrate that cocoa agroforest multifunctionality is substantially influenced by the functional traits of shade tree species. Shade tree species with the most dissimilar traits to cocoa (cocoa showing the lowest leaf litter quality) showed the largest improvement of soil functions. Therefore, selection of shade trees based on their functional traits appears as a promising practice to adequately manage soil functioning. In order to fully assess the beneficial role of shade trees in these agroecosystems. Future research will need to extend this approach to other below‐ground traits and other aspects of multifunctionality such as long‐term cocoa health and yield. [ABSTRACT FROM AUTHOR]

Plant litter inputs can promote the decomposition of soil organic matter (OM) through the priming effect (PE). However, whereas leaf litter chemistry has long been identified as the primary driver of litter decomposition within biomes world‐wide, little is known about how litter chemical traits influence the occurrence and strength of the PE.Here, we studied the effects of 15 co‐occurring C3 leaf litters of contrasting chemistry on C4 soil respiration by analysing changes in 13C natural abundance during early and later stages of litter decomposition (up to 125 days).Besides an apparent PE of 16% in the first 3 days, soil C respiration was increased by 24% on average with leaf litter addition in the initial stage of decomposition (4–26 days) and by 8% at later stages (27–125 days). Most interestingly, soil PE related well to initial litter chemistry and the dominant factors influencing the magnitude of the PE changed with decomposition stage. In the early stage of decomposition, litter leachate C content and litter hemicellulose concentration were positively correlated with the strength of the PE, whereas tannin concentration was negatively associated with soil PE. Together, tannin and hemicellulose explained half of the observed variation in the PE (R2 = 0.58). In the later phase of decomposition, lignin and lignin:N ratios were negatively related to the PE, whereas Ca, K and Mg concentrations were positively related to the PE; lignin alone gave the best prediction of the PE (R2 = 0.58) at later decomposition stages.Our findings provide evidence that the magnitude and direction of the PE is influenced by the chemistry of OM inputs and suggest that, as decomposition proceeds differently among litter of contrasting chemistry, litters can also have variable effect on soil PE through time. The predictive power of litter chemical traits on soil PE opens new perspectives for improving our mechanistic understanding of soil PE and improving our abilities to model soil C dynamics at variable scales. A plain language summary is available for this article. [ABSTRACT FROM AUTHOR]

Agricultural fields are commonly characterized by high nutrient and water availabilities, which are favourable for plant growth. Such conditions might promote the evolution of resource‐acquisitive strategies. We asked whether crop plants show root traits typical of resource‐acquisitive strategies and whether this strategy is primarily a result of their evolution under domestication or of the early selection of successful candidates for domestication.We studied a set of 30 crop species and their wild progenitors. We set up a greenhouse experiment to measure five root traits: root thickness, root tissue density, specific root length (SRL), root mass fraction (RMF) and root length ratio. In addition, we compiled data from other wild herbaceous species, growth in similar conditions to this experiment, to place the root traits of our crops in the context of wider botanical variation.Wild progenitors had thicker and less dense roots, with higher RMF and lower SRL, than other wild herbs. Thicker and less dense roots are indicative of fertile soils, which suggest that wild progenitors could have been adapted for success in agricultural conditions. Additionally, we found that domestication generally increased total plant dry mass, but none of the root traits evolved consistently towards a more resource‐acquisitive strategy after domestication across all species. Root trait values differed between progenitors and crop species for most pairs surveyed, but this occurred in diverse directions depending on crop species. Such differences were independent of phylogeny, functional group or variability in the domestication processes, such as timing of the domestication event or organ under focal artificial selection.Our comparative study revealed that the root phenotype exhibited by wild progenitors (thick roots with low density and SRL), when compared with other wild herbs, was in accordance with plants typical of fertile habitats. However, none of the root traits reacted to domestication in accordance with evolution towards faster growth strategies. Thus, the adaptation of crop root phenotypes to the fertile conditions of agricultural fields might be largely determined by early choices of wild species, rather than by further evolution under domestication. A plain language summary is available for this article. Plain Language Summary [ABSTRACT FROM AUTHOR]

Although microbial communities play an important role in explaining plant litter decomposition rates, whether and how litter chemistry may alter catabolic capacities of soil microbial communities remains poorly studied.During a 1‐year litter decomposition experiment of 12 herbaceous species with contrasting litter chemistry, we examined the effect of plant litter type (roots vs. leaves) and litter chemical traits on the resulting capacity of soil microbial communities to degrade a wide range of carbon substrates of variable complexity (MicroResp™ method).Litter chemistry impacted both the total catabolic activity as well as specific catabolic capacities of microbial communities. In the early stages of litter decomposition total catabolic activity was mainly influenced by the amount of C and N in litter leachates, and litter N, P and Mg, then, later, by lignin concentrations. Some specific catabolic capacities could also be related to litter initial chemistry. Overall, litter trait effects on soil microbial communities decreased over time and the relative importance of traits shifted during the decomposition process.Our results highlight that litter chemistry is a strong driver of catabolic capacities of microbial decomposers and, while its effect fades with time, it remains substantial throughout the litter decomposition process. These long‐lasting effects of litter chemistry suggest a persistent control on microbial catabolic capacities in ecosystems with recurrent litter production. Soil microbial catabolic activities were driven by broadly the same chemical traits across leaf and root litters.Synthesis. Such long‐lasting effects of litter chemistry on catabolic capacities of microbial communities may represent a substantial indirect driver of the decomposition process. Disentangling the relative importance of this overlooked effect of litter chemistry on decomposition represents the next challenge. We argue that such research line should open ground‐breaking perspectives for reconsidering our current understanding of the mechanistic links between litter traits and decomposition rate. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Living roots and their rhizodeposits can accelerate or decelerate the decomposition of soil organic matter which refers to the rhizosphere priming effect (RPE). However, whereas plant traits are thought to be key factors controlling the RPE, little is known about how root traits representative of plant biomass allocation, morphology, architecture, or physiology influence the magnitude of the RPE. Using a natural abundance 13C tracer method allowing partitioning of native soil organic carbon (SOC) decomposition and plant rhizosphere respiration, we studied here the effects of eight C3 tree species featuring contrasting functional traits on C4 soil respiration over a 204‐day period in a microcosm experiment. All tree species enhanced the rate of SOC decomposition, by 82% on average, but the strength of the rhizosphere priming significantly differed among species. Mean diameter of first‐order roots and root exudate‐derived respiration were positively correlated with the RPE, together explaining a large part of observed variation in the RPE (R2 = 0.72), whereas root branching density was negatively associated with the RPE. Path analyses further suggested that mean diameter of first‐order roots was the main driver of the RPE owing to its positive direct effect on the RPE and its indirect effects via root exudate‐derived respiration and root branching density. Our study demonstrates that the magnitude of the RPE is regulated by complementary aspects of root morphology, architecture and physiology, implying that comprehensive approaches are needed to reveal the multiple mechanisms driving plant effects on the RPE. Overall, our results emphasize the relevance of integrating root traits in biogeochemical cycling models to improve model performance for predicting soil C dynamics. [ABSTRACT FROM AUTHOR]

As soils store more carbon (C) than the Earth's atmosphere and terrestrial biomass together, the balance between soil C uptake in the form of soil organic matter (SOC) and release as CO2 upon its decomposition is a critical determinant in the global C cycle regulating our planet's climate. Although plant litter is the predominant source of C fuelling both soil C build‐up and losses, the issue of how litter chemistry influences this balance remains unresolved.As a contribution to solving that issue, we traced the fate of C during near‐complete decomposition of 13C‐labelled leaf and root litters from 12 plant species in a coarse‐textured soil. We separated the soil organic carbon into mineral‐associated organic matter (MAOM) and particulate organic matter (POM) pools, and investigated how 14 litter chemical traits affected novel SOC formation and native SOC mineralization (i.e. the priming effect) in these soil fractions.We observed an overall net increase in SOC due to the addition of litter, which was stronger for root than for leaf litters. The presumed stable MAOM‐C pool underwent both substantial stabilization and mineralization, whereas the presumably less stable POM‐C pool showed substantial stabilization and reduced mineralization. Overall, the initial increase in soil C mineralization was fully counterbalanced by a later decrease in native soil C mineralization. POM‐C formation as well as MAOM‐C formation and mineralization were positively related to the initial litter lignin concentration and negatively to that of the nitrogen leachates, whereas the opposite was observed for POM‐C mineralization.Synthesis. Our results highlight the importance of litter chemical traits for SOC formation, and stabilization, destabilization and mineralization. In our coarse‐textured soil, the amount of MAOM‐C did not change despite large C fluxes through this pool. The litter chemical traits that drove these processes differed from those frequently reported for fine‐textured soils far from mineral‐associated C saturation. To account for these discrepancies, we propose an integrative perspective in which litter quality and soil texture interactively control soil C fluxes by modulating several SOC stabilization and destabilization mechanisms. Irrespective, our results open new critical perspectives for managing soil C pools globally. [ABSTRACT FROM AUTHOR]

Determining the changes in within- and between-species functional diversity in plant communities, and their contribution to overall species trait overlap, can enhance efforts at understanding mechanisms of species coexistence. However, little is known about how variation in species functional diversity influences variation in species trait overlap among contrasting environments., Here, we studied the understorey vegetation in a well-characterized 5000-year-old chronosequence involving 30 forested islands that differ greatly in size, soil fertility and species diversity. Across this chronosequence, we expected consistent changes in both within- and between-species functional diversity that would lead to decreasing overall species trait overlap with increasing successional age, species richness, understorey vegetation density and spatial heterogeneity of soil resources., For each island, we measured specific leaf area ( SLA) of each of ten individuals of each plant species present. Using a variance decomposition method, we partitioned the total community functional diversity of SLA on each island into within- and between-species functional diversity. Further, we estimated overall species trait overlap as the ratio of within-species functional diversity to total functional diversity. Using regression analyses, we then explored relationships of within- and between-species functional diversity, and of overall species trait overlap, with several environmental variables across the 30 islands., Consistent with our hypotheses, overall species trait overlap decreased with successional age due to a statistically significant decrease in within-species functional diversity, and decreased with species richness due to a simultaneous decrease in within-species functional diversity and increase in between-species functional diversity. Against our predictions, overall species trait overlap increased in more competitive environments and did not change with increasing spatial heterogeneity of soil N or P., Synthesis. Our study suggests niche packing as a key mechanism for species coexistence in plant communities. Using SLA as an integrator of plant ecological strategy, we show that community successional age and species richness are significantly linked to trait space distribution of plant individuals of boreal forest understorey vegetation and therefore to local species coexistence. Our results also suggest that the trait space of dominant and subordinate species may respond differently to local environmental variables. [ABSTRACT FROM AUTHOR]

Questions We investigated some commonly held assumptions of community assembly theory needed to provide accurate predictions of changes in plant species assemblages across environmental gradients or following environmental change. Do (1) dominant and subordinate species respond in the same way to changes in environmental variables; (2) plant species assemblages show higher interspecific than intraspecific trait responses; and (3) co-existing dominant species differ in their responses to the same environmental variables? Location Islands in Lakes Uddjaure and Hornavan, northern Sweden. Methods We explored the responses of forest understorey vegetation assemblages to variation in environmental resources across a chronosequence of 30 lake islands that differ in fire history, above-ground and below-ground resource availability and species diversity. For one plot on each island, we measured specific leaf area, leaf dry matter content and foliar N and P of all dominant and subordinate understorey plant species to assess species-specific and weighted and non-weighted community-level trait responses to variation across islands in all major local environmental drivers. Results Consistent with our expectations, we found that species responses to environmental conditions were not homogenous within assemblages, and that responses of dominant and subordinate species differed. Further, intraspecific variation was often an important component of local-scale plant community-level responses. Responses were often relatively consistent across species, but dominant species sometimes showed contrasting responses of the same trait to the same environmental factor. Finally, environmental factors that influenced community average trait values also affected functional diversity. Conclusions This study has shown that several common assumptions that underpin community assembly theory do not necessarily hold, and this can cause inaccuracies in predicting plant functional composition responses to changes in environmental variables. Because these assumptions are central to current models that predict vegetation responses to environmental change, it is crucial to further test in which particular environmental context and to what extent these assumptions are critical for model accuracy. [ABSTRACT FROM AUTHOR]

Climate warming is shifting the distributions of mountain plant species to higher elevations. Cold‐adapted plant species are under increasing pressure from novel competitors that are encroaching from lower elevations. Plant capacity to adjust to these pressures may be measurable as variation in trait values within a species. In particular, the strength and patterns of intraspecific trait variation along abiotic and biotic gradients can inform us whether and how species can adjust their anatomy and morphology to persist in a changing environment.Here, we tested whether species specialized to high elevations or with narrow elevational ranges show more conservative (i.e. less variable) trait responses across their elevational distribution, or in response to neighbours, than species from lower elevations or with wider elevational ranges. We did so by studying intraspecific trait variation of 66 species along 40 elevational gradients in four countries in both hemispheres. As an indication of potential neighbour interactions that could drive trait variation, we also analysed plant species' height ratio, its height relative to its nearest neighbour.Variation in alpine plant trait values over elevation differed depending on a species' median elevation and the breadth of its elevational range, with species with lower median elevations and larger elevational range sizes showing greater trait variation, i.e. a steeper slope in trait values, over their elevational distributions. These effects were evidenced by significant interactions between species' elevation and their elevational preference or range for several traits: vegetative height, generative height, specific leaf area and patch area. The height ratio of focal alpine species and their neighbours decreased in the lower part of their distribution because neighbours became relatively taller at lower elevations. In contrast, species with lower elevational optima maintained a similar height ratio with neighbours throughout their range.Synthesis. We provide evidence that species from lower elevations and those with larger range sizes show greater intraspecific trait variation, which may indicate a greater ability to respond to environmental changes. Also, larger trait variation of species from lower elevations may indicate stronger competitive ability of upslope shifting species, posing one further threat to species from higher ranges. [ABSTRACT FROM AUTHOR]

Maintaining yield when reducing inputs is one prime objective of sustainable agriculture. In this context, cereal-legume intercropping is a practice that can achieve increased yield under low-input conditions through the complementary use of abiotic resources and facilitation mechanisms. Many management options exist to design cereal-legume intercropping systems, among which the choice of the species intercropped and the level of nitrogen (N) fertilization are essential. In this study, we collected the results of 35 field experiments across Europe of cereal-grain legume intercrops that combined various intercropped species and N fertilization levels. We first assessed the intensity of the biodiversity effect and its components in unfertilized intercrops. Then, we focused on a subset of systems to analyze how N fertilization influenced biodiversity effects on three intercrops (durum wheat/pea, soft wheat/pea, and durum wheat/faba bean). The biodiversity effect represents the gap between the observed and expected yields of a mixture. The complementarity effect is the performance of mixtures relative to the performance of the component monocultures. The selection effect captures the extent to which a species with a high monoculture yield dominates a mixture at the expense of the other intercropped species. Our results confirmed an overall positive biodiversity effect under unfertilized conditions and various climate conditions (0.86 ± 0.04 t.ha−1). Complementarity effect was the main driver as it represented 76% of the biodiversity effect, confirming intercropping as a useful practice in low-input systems. N fertilization lowered the complementarity effect in durum wheat/pea intercrops, did not influence these effects in soft wheat/pea intercrops, and increased only the selection effect in durum wheat/faba bean intercrops. These results highlight the need for a sufficiently competitive legume in intercrops when N fertilizers are applied in order to avoid too much disruption of plant–plant interactions. [ABSTRACT FROM AUTHOR]

Summary: 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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The importance of species richness to ecosystem functioning and services is a central tenet of biological conservation. However, most of our theory and mechanistic understanding is based on diversity found aboveground. Our study sought to better understand the relationship between diversity and belowground function by studying root biomass across a plant diversity gradient. We collected soil cores from 91 plots with between 1 and 12 aboveground tree species in three natural secondary forests to measure fine root (≤ 2 mm in diameter) biomass. Molecular methods were used to identify the tree species of fine roots and to estimate fine root biomass for each species. This study tested whether the spatial root partitioning (species differ by belowground territory) and symmetric growth (the capacity to colonize nutrient‐rich hotspots) underpin the relationship between aboveground species richness and fine root biomass. All species preferred to grow in nutrient‐rich areas and symmetric growth could explain the positive relationship between aboveground species richness and fine root biomass. However, symmetric growth only appeared in the nutrient‐rich upper soil layer (0–10 cm). Structural equation modelling indicated that aboveground species richness and stand density significantly affected fine root biomass. Specifically, fine root biomass depended on the interaction between aboveground species richness and stand density, with fine root biomass increasing with species richness at lower stand density, but not at higher stand density. Overall, evidence for spatial (i.e. vertical) root partitioning was inconsistent; assumingly any roots growing into deeper unexplored soil layers were not sufficient contributors to the positive diversity–function relationship. Alternatively, density‐dependent biotic interactions affecting tree recruitment are an important driver affecting productivity in diverse subtropical forests but the usual root distribution patterns in line with the spatial root partitioning hypothesis are unrealistic in contexts where soil nutrients are heterogeneously distributed. [ABSTRACT FROM AUTHOR]

The article provides a solid foundation for predictive understanding of belowground responses to changing environmental conditions. It shows that root-mycorrhizal collaboration can short circuit a one-dimensional economic spectrum, providing an entire space of economic possibilities. It mentions that symbiotic partnerships and different plant growth forms.

Deciphering the mechanisms that drive variation in biomass production across plant communities of contrasting species composition and diversity is a main challenge of biodiversity–ecosystem functioning research. Niche complementarity and selection effect have been widely investigated to address biodiversity–productivity relationships. However, the overlooking of the specific role played by key species has limited so far our capacity to comprehensively assess the relative importance of other potential drivers of biodiversity effects.Here, we conducted a grassland diversity–productivity experiment to test how four potential facets of biodiversity effects, namely species richness, functional diversity, species identity and the relaxation of intraspecific competition, account for variations in above and root biomass production.We grew six plant species in monoculture, as well as in every combination of two, three and six species. Plant density was kept constant across the richness gradient but we additionally grew each species in half‐density monoculture to estimate the strength of intraspecific competition for each studied species. We characterized eight functional traits, including root traits, related to nutrient and light acquisition and computed both the functional dissimilarity and the community‐weighted mean (CWM) of each trait. We further partitioned above‐ground biodiversity effect into complementarity and selection effects.We observed strong positive biodiversity effects on both above‐ground and root biomass as well as strong positive complementarity effect. These arose largely from the presence of a particular species (Plantago lanceolata) and from CWM trait values more than from a higher functional dissimilarity in plant mixtures. P. lanceolata displayed the highest intraspecific competition, which was strongly relaxed in species mixtures. By contrast, the presence of Sanguisorba minor negatively affected the productivity of plant mixtures, this species suffering more from interspecific than intraspecific competition.This study provides strong evidences that the search for key species is critical to understand the role of species diversity on ecosystem functioning and demonstrates the major role that the balance between intraspecific and interspecific competition plays in biodiversity–ecosystem functioning relationships. Developing more integrative approaches in community and ecosystem ecology can offer opportunities to better understand the role that species diversity plays on ecosystem functioning. A free Plain Language Summary can be found within the Supporting Information of this article. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Concern about the functional consequences of unprecedented loss in biodiversity has prompted biodiversity-ecosystem functioning (BEF) research to become one of the most active fields of ecological research in the past 25 years. Hundreds of experiments have manipulated biodiversity as an independent variable and found compelling support that the functioning of ecosystems increases with the diversity of their ecological communities. This research has also identified some of the mechanisms underlying BEF relationships, some context-dependencies of the strength of relationships, as well as implications for various ecosystem services that humankind depends upon. In this chapter, we argue that a multitrophic perspective of biotic interactions in random and non-random biodiversity change scenarios is key to advance future BEF research and to address some of its most important remaining challenges. We discuss that the study and the quantification of multitrophic interactions in space and time facilitates scaling up from small-scale biodiversity manipulations and ecosystem function assessments to management-relevant spatial scales across ecosystem boundaries. We specifically consider multitrophic conceptual frameworks to understand and predict the context-dependency of BEF relationships. Moreover, we highlight the importance of the eco-evolutionary underpinnings of multitrophic BEF relationships. We outline that FAIR data (meeting the standards of findability, accessibility, interoperability, and reusability) and reproducible processing will be key to advance this field of research by making it more integrative. Finally, we show how these BEF insights may be implemented for ecosystem management, society, and policy. Given that human well-being critically depends on the multiple services provided by diverse, multitrophic communities, integrating the approaches of evolutionary ecology, community ecology, and ecosystem ecology in future BEF research will be key to refine conservation targets and develop sustainable management strategies. [ABSTRACT FROM AUTHOR]

Questions: Mycorrhizae may be a key element of plant nutritional strategies and of carbon and nutrient cycling. Recent research suggests that in natural conditions, intensity of mycorrhizal colonization should be considered an important plant feature. How are inter-specific variations in mycorrhizal colonization rate, plant relative growth rate (RGR) and leaf litter decomposability related? Is (arbuscular) mycorrhizal colonization linked to the dominance of plant species in nutrient-stressed ecosystems? Location: Teberda State Biosphere Reserve, northwest Caucasus, Russia. Methods: We measured plant RGR under mycorrhizal limitation and under natural nutrition conditions, together with leaf litter decomposability and field intensity of mycorrhizal colonization across a wide range of plant species, typical for alpine communities of European mountains. We applied regression analysis to test whether the intensity of mycorrhizal colonization is a good predictor of RGR and decomposition rate, and tested how these traits predict plant dominance in communities. Results: Forb species with a high level of field mycorrhizal colonization had lower RGR under nutritional and mycorrhizal limitation, while grasses were unaffected. Litter decomposition rate was not related to the intensity of mycorrhizal colonization. Dominant species mostly had a higher level of mycorrhizal colonization and lower RGR without mycorrhizal colonization than subordinate species, implying that they were more dependent on mycorrhizal symbionts. There were no differences in litter decomposability. Conclusions: In alpine herbaceous plant communities dominated by arbuscular mycorrhizae, nutrient dynamics are to a large extent controlled by mycorrhizal symbiosis. Intensity of mycorrhizal colonization is a negative predictor for whole plant RGR. Our study highlights the importance of mycorrhizal colonization as a key trait underpinning the role of plant species in carbon and nutrient dynamics in nutrient-limited herbaceous plant communities. [ABSTRACT FROM AUTHOR]

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, ) 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. [ABSTRACT FROM AUTHOR]

Recent studies have shown that accounting for intraspecific trait variation ( ITV) may better address major questions in community ecology. However, a general picture of the relative extent of ITV compared to interspecific trait variation in plant communities is still missing. Here, we conducted a meta-analysis of the relative extent of ITV within and among plant communities worldwide, using a data set encompassing 629 communities (plots) and 36 functional traits. Overall, ITV accounted for 25% of the total trait variation within communities and 32% of the total trait variation among communities on average. The relative extent of ITV tended to be greater for whole-plant (e.g. plant height) vs. organ-level traits and for leaf chemical (e.g. leaf N and P concentration) vs. leaf morphological (e.g. leaf area and thickness) traits. The relative amount of ITV decreased with increasing species richness and spatial extent, but did not vary with plant growth form or climate. These results highlight global patterns in the relative importance of ITV in plant communities, providing practical guidelines for when researchers should include ITV in trait-based community and ecosystem studies. [ABSTRACT FROM AUTHOR]