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Tree mycorrhizal type plays an important role in promoting plant species diversity and coexistence, via its mediating role in conspecific negative density dependence (CNDD), i.e., the process by which an individual's performance is impaired by the density of conspecific plants. Previous findings suggest that ectomycorrhizal (EM) tree species are generally less susceptible to CNDD than arbuscular mycorrhizal (AM) tree species, due to the chemical and physical protection that EM fungi provide their host with. We examined how CNDD effects on leaf herbivory, seedling growth, and survival differ between AM and EM seedlings of ten tree species collected over 3 years in an old-growth temperate forest in northeastern China. We found that AM and EM seedlings differed in how conspecific density affected their leaf herbivory, seedling growth, and survival. Specifically, AM seedlings leaf herbivory rates significantly increased with increasing conspecific seedling and adult density, and their growth and survival rates decreased with increasing conspecific adult density, these patterns were, however, absent in EM seedlings. Our work suggests that AM seedlings have a performance disadvantage relative to EM seedlings related to the negative effects from conspecific neighbors. We highlight the importance of integrating information on seedling leaf herbivory, seedling growth, to provide further understanding on potential mechanisms driving differences in CNDD between AM and EM tree seedlings.

Arbuscular mycorrhizal fungi (AMF) and ectomycorrhizal fungi (EMF) produce contrasting plant-soil feedbacks, but how these feedbacks are constrained by lithology is poorly understood. We investigated the hypothesis that lithological drivers of soil fertility filter plant resource economic strategies in ways that influence the relative fitness of trees with AMF or EMF symbioses in a Bornean rain forest containing species with both mycorrhizal strategies. Using forest inventory data on 1245 tree species, we found that although AMF-hosting trees had greater relative dominance on all soil types, with declining lithological soil fertility EMF-hosting trees became more dominant. Data on 13 leaf traits and wood density for a total of 150 species showed that variation was almost always associated with soil type, whereas for six leaf traits (structural properties; carbon, nitrogen, phosphorus ratios, nitrogen isotopes), variation was also associated with mycorrhizal strategy. EMF-hosting species had slower leaf economics than AMF-hosts, demonstrating the central role of mycorrhizal symbiosis in plant resource economies. At the global scale, climate has been shown to shape forest mycorrhizal composition, but here we show that in communities it depends on soil lithology, suggesting scale-dependent abiotic factors influence feedbacks underlying the relative fitness of different mycorrhizal strategies.

International audience; Microbial communities strongly influence ecological processes across space and time. However, the influence of abiotic and biotic factors on microbial functioning in a complex, heterogeneous soil environment is still poorly understood, especially in montane plant communities. Here, we ask if microbial activity and functional diversity can be influenced by plant species identity and diversity and associated root system morphological and chemical traits that also influence soil properties. We investigated microbial global catabolic activity (i.e. microbial activity) and catabolic diversity (i.e. functional diversity) in bulk and rhizospheric soil beneath three plant species (Vaccinium myrtillus, Juniperus communis and Picea abies) that shape local plant communities. To do this, we measured soil physical and chemical properties, and plant diversity along an elevational gradient (1400–2400 m a.s.l.) in the French Alps. Microbial global catabolic activity and diversity were assessed using multiple substrate-induced respiration. Morphological and chemical traits of roots in bulk soil (‘community’ level traits, where several plant species were pooled together) and of individual plants (‘species’ level, where roots of single species were excavated) were measured. Results showed that at lower elevations, global catabolic activity in the rhizosphere was higher than in bulk soil, but converged in the nutrient-poor, colder soils found at higher elevations, although changes in catabolic diversity were negligible. Variations in soil physical and chemical properties were the main drivers of global catabolic activity, especially texture, cation exchange capacity, carbon and nitrogen content and pH, but their effects on catabolic diversity were minimal. Plant root traits at the community level in bulk soil had less effect on global catabolic activity than abiotic factors, with thicker roots, high root lignin content and low cellulose content influencing microbial activity, but not altering catabolic diversity. At the species level, more dense root tissue decreased global catabolic activity, reflecting changes in chemical composition. Our results show that relationships between catabolic activity, mean annual temperature and soil properties differed between plant species.

International audience; With the aim to explore how plants acclimate to elevation changes in the understudied (sub)alpine tropics we tested two hypotheses along a 1000-m elevation gradient in Mexico: (H1) due to a severe increase in abiotic constraints at higher elevations, the functional traits of the plant species will converge toward more resource conservation, and (H2) the specific growth forms and biogeographic origins present in the (sub)alpine tropics may influence the interspecific trait variation along the gradient. We measured five aboveground functional traits: specific leaf area (SLA), leaf dry-matter content (LDMC), leaf thickness, leaf area and plant height, of 11 species representing four growth forms: rosette, tussock grass, shrub and tree the soil microclimate. Microclimatic data revealed a steep decrease in soil water content at higher elevations. Across all species and all individuals, SLA, plant height and leaf area decreased with elevation, whereas LDMC and leaf thickness increased, all of which revealing adjustments towards resource conservation in line with H1. Consistently with H2, the functional traits of the growth forms that were characteristic of tropical alpine regions (tussock grasses and erect shrubs) were less sensitive to changes in elevation compared to more generalist growth forms such as forbs. In addition, within the growth form “rosette” the functional traits of species of tropical biogeographic origin changed with elevation, whereas those of Holarctic origin did not. Our data indicate a convergence of plant traits toward improved resource conservation at higher elevations, which may be influenced partially by the growth form and the biogeographical origin of plant species.

International audience; 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.

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. ISSN:0028-646X ISSN:1469-8137

Root systems show a tremendous diversity both between and within species, suggesting a large variability in plant functioning and effects on ecosystem properties and processes. In recent decades, developments in many areas of root research have brought considerable advances in our understanding of root traits and their contribution to plant and ecosystem functioning. However, despite major progress, a comprehensive overview—bridging research fields—is lacking. Furthermore, considerable uncertainties exist in the identification of root entities, and the selection and standardized measurement of traits. Here, we provide a comprehensive overview on root entities, exemplify recent advances in our understanding of both theoretical and demonstrated relationships between root traits and plant or ecosystem functioning, discuss trait-trait relationships and hierarchies among traits, and critically assess current strengths and gaps in our knowledge.

1. The root economics spectrum (RES) hypothesis predicts that fast-growing tree species have short-lived roots with high specific root length (SRL) to allow rapid resource uptake, and opposite trait expressions for slow-growing species. Yet, the mixed support for this hypothesis suggests that trees can adopt alternative strategies to increase resource uptake, besides an increase in SRL. 2. We combined a novel mechanistic whole-tree model and empirical fine-root data of ten tree species to test the effects of one of these alternative strategies, notably increasing fine-root mass, on the tree’s net C gain (used here as a proxy for tree performance), and to assess how fine-root lifespan influences the relative importance of SRL and fine-root mass for the C balance of trees. 3. Our results indicate that accounting for the short lifespan of high-SRL roots has important implications for explaining tree performance and the role of roots herein. Without considering their faster turnover, high-SRL roots and low fine-root mass resulted in the highest performance as predicted from the RES. Yet, when their higher turnover rates were accounted for, a high fine-root mass and low SRL lead to the highest performance. Both our model outcomes and field data further show a negative relationship between SRL and fine-root mass through which species aim to realise a similar root length density. This trade-off further indicates how high a SRL and low fine-root mass as well as opposite trait values can both lead to a positive C balance in a similar environment. 4. Our study may explain why high-SRL roots do not necessarily lead to the fastest tree growth as often hypothesised and demonstrates the importance of fine-root mass in combination with fine-root lifespan for explaining interspecific differences in tree performance. More generally, our work demonstrates the value of identifying and investigating different belowground strategies across species from a whole-plant modelling perspective, and identifies the relationship between SRL, fine-root biomass and lifespan as an important functional dimension to variation in species’ performance.

The growth of trees depends on photosynthetic carbon gain by the leaves, which in turn relies on water and nutrient acquisition by the fine roots. Because the availability of carbon, water and nutrients fluctuates, trees can adjust their leaf and fine-root functional traits to maintain their resource uptake and growth rates. Aboveground, the variation in leaf traits is closely related to light availability, light uptake and tree growth. Within species, leaves show general, plastic responses to their light environment, so that trees can still intercept light when its availability changes. Across species, leaf traits are coordinated along a leaf economics spectrum (LES), which reflects species resource strategies. On the one end of this spectrum, acquisitive species have leaves that allow fast resource uptake and therefore fast tree growth. Conversely, species with a conservative strategy acquire resources more slowly, but retain them longer, so they can tolerate low resource availability. Belowground, the relationships between fine-root functional traits, water and nutrient availability and acquisition, and tree growth are expected to be similar to those aboveground, but are still poorly understood. Understanding these relationships is essential as tree growth results from the simultaneous uptake of above- and belowground resources. Therefore, this thesis examines how fine-root traits relate to growth, and focuses on across- and within-species variation in tree root traits. We first tested whether plant resource strategies can explain drought effects on tree growth across 10 common tree species that ranged from acquisitive to conservative species (Chapter 2). Based on tree-ring analyses, we found that the growth rates of all species were significantly lower in years with dry summers. Although the strength of these growth responses differed, these differences were not related to species resource strategies. However, when groundwater levels receded, acquisitive species grew slower but conservative species did not, which suggests root trait differences across these species. Drought effects on tree growth may thus not always be fully explained from an acquisitive or conservative resource strategy. We further evaluated whether a root economics spectrum (RES) parallel to a LES can explain variation in fine-root functional traits across species (Chapter 3). Our literature review shows no consistent evidence for an RES, due to three fundamental differences between fine roots and leaves. First, fine-root traits are not only aimed at increased resource uptake or conservation, but are also constrained by the soil environment. Second, the relationships between traits and function are far less clear for roots than for leaves. Third, the expected relationships between fine-root traits and resource uptake are obscured by mycorrhizal fungi. Revealing the links between fine-root traits, resource acquisition, and growth across species, therefore requires a multidimensional approach that incorporates these different interacting variables. Chapter 4 examines intraspecific variation in fine-root traits and mycorrhizal biomass in Fagus sylvatica L. and Picea abies L. forests on a poor, sandy soil and a resource-rich clay soil in the Netherlands. Both species increased their fine-root mass and fine-root growth rates on the sandy soils compared to the clay soils, but fine-root morphology did not differ between the soil types. In the P. abies stands, ectomycorrhizal biomass was larger on sand than on clay, possibly increasing tree resource uptake. Besides the strong increase in fine-root mass observed for both species, species may thus also differ in their fine-root plasticity strategies to cope with various soil environments. To understand tree growth from below- and aboveground trait integration, we explored the impacts of fine-root mass and morphology on nutrient acquisition and tree fitness using a whole-tree growth model (Chapter 5). More specifically, we tested which combination(s) of fine-root mass and specific root length (SRL) led to optimal fitness, based on the uptake benefits (i.e. increasing the belowground uptake area) and carbon costs (i.e. turnover and respiration) of these traits. Our results show that tree fitness increased with fine-root mass but especially through an increase in SRL. Furthermore, both a combination of high fine-root mass and low SRL, and of low fine-root mass and high SRL, resulted in similar net carbon gain, indicating that alternative strategies that may lead to similar fitness. To conclude, trees rely on various uptake strategies to ensure belowground resource uptake and tree growth in different environments. Specific root length is often expected to be tightly linked to tree growth, but this thesis shows that there is little support for this hypothesised relationship. Consequently, the functional meaning of SRL requires further study. Instead, fine-root mass and mycorrhizal symbiosis may present more important alternatives to enhance water and nutrient uptake, both across and within species. Moreover, to cope with the highly complex soil (resource) environment, species have adopted various other uptake strategies besides fine-root mass, morphology and mycorrhiza. This thesis stresses that a multidimensional root-trait framework is needed to link fine-root traits to tree growth, that can accommodate this variety of fine-root traits and the diversity of the soil environment.

AimThe fine roots of trees may show plastic responses to their resource environment. Several, contrasting hypotheses exist on this plasticity, but empirical evidence for these hypotheses is scattered. This study aims to enhance our understanding of tree root plasticity by examining intra-specific variation in fine-root mass and morphology, fine-root growth and decomposition, and associated mycorrhizal interactions in beech (Fagus sylvatica L.) and spruce (Picea abies (L.) Karst.) forests on soils that differ in resource availability.MethodsWe measured the mass and morphological traits of fine roots (i.e. ≤ 2 mm diameter) sampled to 50 cm depth. Fine-root growth was measured with ingrowth cores, and fine-root decomposition with litter bags. Mycorrhizal fungal biomass was determined using ingrowth mesh bags.ResultsBoth tree species showed more than three times higher fine-root mass, and a ten-fold higher fine-root growth rate on sand than on clay, but no or marginal differences in overall fine-root morphology. Within the fine-root category however, beech stands had relatively more root length of their finest roots on clay than on sand. In the spruce stands, ectomycorrhizal mycelium biomass was larger on sand than on clay.ConclusionsIn temperate beech and spruce forests, fine-root mass and mycorrhizal fungal biomass, rather than fine-root morphology, are changed to ensure uptake under different soil resource conditions. Yet enhancing our mechanistic understanding of fine-root trait plasticity and how it affects tree growth requires more attention to fine-root dynamics, the functional diversity within the fine-roots, and mycorrhizal symbiosis as an important belowground uptake strategy.

Studies on climate impacts on tree annual growth are mainly restricted to marginal sites. To date, the climate effects on annual growth of trees in favorable environments remain therefore unclear despite the importance of these sites in terms of forest productivity. Because species respond differently to climate, comparing a multitude of species further enhances our knowledge on climate impacts on tree growth and forest productivity. We present a first study that reveals to what extent radial growth is limited by climate and the groundwater table across 10 temperate tree species growing on a uniform, productive soil (i.e. high nutrient and water contents) in the Netherlands. We ranked our study species according to their shade tolerance, which is associated with species’ resource requirements and growth rates, and examined their annual growth using tree-ring analysis. This allowed us to investigate how these species with diverse ecological backgrounds differ in their growth response to precipitation, temperature, irradiance, potential evapotranspiration (PET), and the groundwater table, when growing under similar and favorable site conditions. Nine out of 10 species had strikingly similar radial growth rates (on average 3.9–4.8 mm yr−1), which contradicts the widely established trade-off between shade tolerance and growth. Populus trichocarpa, the least shade-tolerant of our species, however grew much faster (on average 6.8 mm yr−1). Trees of all species reduced their growth significantly during dry summers, driven by low rainfall and high PET. Yet, the magnitude of their growth responses to climate conditions and especially the groundwater table differed across species. Receding groundwater tables significantly and strongly reduced radial growth of the shade-intolerant species, but not of the shade-tolerant species. The climate impacts on growth were not associated with variation in shade tolerance across our species. Our study demonstrates that even on a productive forest site, summer droughts reduced radial growth across a multitude of common tree species, whereas lower groundwater tables only affected the shade-intolerant species significantly. Thus, the productivity of forests in favorable environments may be seriously affected when summers in north western Europe become hotter and drier as predicted by climate models.

Dead wood provides a huge terrestrial carbon stock and a habitat to wide-ranging organisms during its decay. Our brief review highlights that, in order to understand environmental change impacts on these functions, we need to quantify the contributions of different interacting biotic and abiotic drivers to wood decomposition. LOGLIFE is a new long-term 'common-garden' experiment to disentangle the effects of species' wood traits and siterelated environmental drivers on wood decomposition dynamics and its associated diversity of microbial and invertebrate communities. This experiment is firmly rooted in pioneering experiments under the directorship of Terry Callaghan at Abisko Research Station, Sweden. LOGLIFE features two contrasting forest sites in the Netherlands, each hosting a similar set of coarse logs and branches of 10 tree species. LOGLIFE welcomes other researchers to test further questions concerning coarse wood decay that will also help to optimise forest management in view of carbon sequestration and biodiversity conservation. Copyright © Royal Swedish Academy of Sciences 2012.

Leaf, xylem and phloem areas drive the water and carbon fluxes within branches and trees, but their mutual coordination is poorly understood. We test the hypothesis that xylem and phloem areas increase relative to leaf area when species are selected for, or branches are exposed to, higher levels of light intensity. Trees of 10 temperate, broadleaved and deciduous, tree species were selected. Fifty-centimetre-long branches were collected from shaded and exposed conditions at a height of 3-4 m. We measured the total leaf area, xylem area, phloem area and leaf traits, as well as the area of the constituent cell types, for a stem section at the branch base. Xylem area : leaf area and phloem area : leaf area ratios did not differ consistently between sun and shade branches, but, as expected, they decreased with species' shade tolerance. Similar trends were observed for conductive cell areas in xylem and phloem. Trees of light-demanding species maintain higher water loss and carbon gain rates per leaf area by producing more xylem area and phloem area than shade-tolerant species. We call for more comparative branch studies as they provide an integrated biological perspective on functional traits and their role in the ecology of tree species.

Contents 1159 I. 1159 II. 1161 III. 1164 IV. 1166 1167 References 1167 SUMMARY: The search for a root economics spectrum (RES) has been sparked by recent interest in trait-based plant ecology. By analogy with the one-dimensional leaf economics spectrum (LES), fine-root traits are hypothesised to match leaf traits which are coordinated along one axis from resource acquisitive to conservative traits. However, our literature review and meta-level analysis reveal no consistent evidence of an RES mirroring an LES. Instead the RES appears to be multidimensional. We discuss three fundamental differences contributing to the discrepancy between these spectra. First, root traits are simultaneously constrained by various environmental drivers not necessarily related to resource uptake. Second, above- and belowground traits cannot be considered analogues, because they function differently and might not be related to resource uptake in a similar manner. Third, mycorrhizal interactions may offset selection for an RES. Understanding and explaining the belowground mechanisms and trade-offs that drive variation in root traits, resource acquisition and plant performance across species, thus requires a fundamentally different approach than applied aboveground. We therefore call for studies that can functionally incorporate the root traits involved in resource uptake, the complex soil environment and the various soil resource uptake mechanisms - particularly the mycorrhizal pathway - in a multidimensional root trait framework.