Mycorrhizal associations modify tree diversity-productivity relationships across experimental tree plantations.

Decades of studies have demonstrated links between biodiversity and ecosystem functioning, yet the generality of the relationships and the underlying mechanisms remain unclear, especially for forest ecosystems. Using 11 tree-diversity experiments, we tested tree species richness-community productivity relationships and the role of arbuscular (AM) or ectomycorrhizal (ECM) fungal-associated tree species in these relationships. Tree species richness had a positive effect on community productivity across experiments, modified by the diversity of tree mycorrhizal associations. In communities with both AM and ECM trees, species richness showed positive effects on community productivity, which could have resulted from complementarity between AM and ECM trees. Moreover, both AM and ECM trees were more productive in mixed communities with both AM and ECM trees than in communities assembled by their own mycorrhizal type of trees. In communities containing only ECM trees, species richness had a significant positive effect on productivity, whereas species richness did not show any significant effects on productivity in communities containing only AM trees. Our study provides novel explanations for variations in diversity-productivity relationships by suggesting that tree-mycorrhiza interactions can shape productivity in mixed-species forest ecosystems.

Automatic Root Length Estimation from Images Acquired In Situ without Segmentation.

Image-based root phenotyping technologies, including the minirhizotron (MR), have expanded our understanding of the in situ root responses to changing environmental conditions. The conventional manual methods used to analyze MR images are time-consuming, limiting their implementation. This study presents an adaptation of our previously developed convolutional neural network-based models to estimate the total (cumulative) root length (TRL) per MR image without requiring segmentation. Training data were derived from manual annotations in Rootfly, commonly used software for MR image analysis. We compared TRL estimation with 2 models, a regression-based model and a detection-based model that detects the annotated points along the roots. Notably, the detection-based model can assist in examining human annotations by providing a visual inspection of roots in MR images. The models were trained and tested with 4,015 images acquired using 2 MR system types (manual and automated) and from 4 crop species (corn, pepper, melon, and tomato) grown under various abiotic stresses. These datasets are made publicly available as part of this publication. The coefficients of determination (R2), between the measurements made using Rootfly and the suggested TRL estimation models were 0.929 to 0.986 for the main datasets, demonstrating that this tool is accurate and robust. Additional analyses were conducted to examine the effects of (a) the data acquisition system and thus the image quality on the models' performance, (b) automated differentiation between images with and without roots, and (c) the use of the transfer learning technique. These approaches can support precision agriculture by providing real-time root growth information.

Semantic segmentation of plant roots from RGB (mini-) rhizotron images-generalisation potential and false positives of established methods and advanced deep-learning models.

BACKGROUND: Manual analysis of (mini-)rhizotron (MR) images is tedious. Several methods have been proposed for semantic root segmentation based on homogeneous, single-source MR datasets. Recent advances in deep learning (DL) have enabled automated feature extraction, but comparisons of segmentation accuracy, false positives and transferability are virtually lacking. Here we compare six state-of-the-art methods and propose two improved DL models for semantic root segmentation using a large MR dataset with and without augmented data. We determine the performance of the methods on a homogeneous maize dataset, and a mixed dataset of > 8 species (mixtures), 6 soil types and 4 imaging systems. The generalisation potential of the derived DL models is determined on a distinct, unseen dataset. RESULTS: The best performance was achieved by the U-Net models; the more complex the encoder the better the accuracy and generalisation of the model. The heterogeneous mixed MR dataset was a particularly challenging for the non-U-Net techniques. Data augmentation enhanced model performance. We demonstrated the improved performance of deep meta-architectures and feature extractors, and a reduction in the number of false positives. CONCLUSIONS: Although correction factors are still required to match human labelled root lengths, neural network architectures greatly reduce the time required to compute the root length. The more complex architectures illustrate how future improvements in root segmentation within MR images can be achieved, particularly reaching higher segmentation accuracies and model generalisation when analysing real-world datasets with artefacts-limiting the need for model retraining.

Particulate matter accumulation by tree foliage is driven by leaf habit types, urbanization- and pollution levels.

Particulate matter (PM) pollution poses a significant threat to human health. Greenery, particularly trees, can act as effective filters for PM, reducing associated health risks. Previous studies have indicated that tree traits play a crucial role in determining the amount of PM accumulated on leaves, although findings have often been site-specific. To comprehensively investigate the key factors influencing PM binding to leaves across diverse tree species and geographical locations, we conducted an extensive analysis using data extracted from 57 publications. The data covers 11 countries and 190 tree species from 1996 to 2021. We categorized tree species into functional groups: evergreen conifers, deciduous conifers, deciduous broadleaves, and evergreen broadleaves based on leaf habit and phylogeny. Evergreen conifers exhibited the highest PM accumulation on leaves, and in general, evergreen leaves accumulated more PM compared to deciduous leaves across all PM size classes. Specific leaf traits, such as epicuticular wax, played a significant role. The highest PM loads on leaves were observed in peri-urban areas along the rural-peri-urban-urban gradient. However, the availability of global data was skewed, with most data originating from urban and peri-urban areas, primarily from China and Poland. Among different climate zones, substantial data were only available for warm temperate and cold steppe climate zones. Understanding the problem of PM pollution and the role of greenery in urban environments is crucial for monitoring and controlling PM pollution. Our systematic review of the literature highlights the variation on PM loading among different vegetation types with varying leaf characteristics. Notably, epicuticular wax emerged as a marker trait that exhibited variability across PM size fractions and different vegetation types. In conclusion, this review emphasizes the importance of greenery in mitigation PM pollution. Our findings underscore the significance of tree traits in PM binding. However, lack of data stresses the need for further research and data collection initiatives.

Root and branch hydraulic functioning and trait coordination across organs in drought-deciduous and evergreen tree species of a subtropical highland forest.

Vessel traits are key in understanding trees' hydraulic efficiency, and related characteristics like growth performance and drought tolerance. While most plant hydraulic studies have focused on aboveground organs, our understanding of root hydraulic functioning and trait coordination across organs remains limited. Furthermore, studies from seasonally dry (sub-)tropical ecosystems and mountain forests are virtually lacking and uncertainties remain regarding potentially different hydraulic strategies of plants differing in leaf habit. Here, we compared wood anatomical traits and specific hydraulic conductivities between coarse roots and small branches of five drought-deciduous and eight evergreen angiosperm tree species in a seasonally dry subtropical Afromontane forest in Ethiopia. We hypothesized that largest vessels and highest hydraulic conductivities are found in roots, with greater vessel tapering between roots and equally-sized branches in evergreen angiosperms due to their drought-tolerating strategy. We further hypothesized that the hydraulic efficiencies of root and branches cannot be predicted from wood density, but that wood densities across organs are generally related. Root-to-branch ratios of conduit diameters varied between 0.8 and 2.8, indicating considerable differences in tapering from coarse roots to small branches. While deciduous trees showed larger branch xylem vessels compared to evergreen angiosperms, root-to-branch ratios were highly variable within both leaf habit types, and evergreen species did not show a more pronounced degree of tapering. Empirically determined hydraulic conductivity and corresponding root-to-branch ratios were similar between both leaf habit types. Wood density of angiosperm roots was negatively related to hydraulic efficiency and vessel dimensions; weaker relationships were found in branches. Wood density of small branches was neither related to stem nor coarse root wood densities. We conclude that in seasonally dry subtropical forests, similar-sized coarse roots hold larger xylem vessels than small branches, but the degree of tapering from roots to branches is highly variable. Our results indicate that leaf habit does not necessarily influence the relationship between coarse root and branch hydraulic traits. However, larger conduits in branches and a low carbon investment in less dense wood may be a prerequisite for high growth rates of drought-deciduous trees during their shortened growing season. The correlation of stem and root wood densities with root hydraulic traits but not branch wood points toward large trade-offs in branch xylem towards mechanical properties.

Soil fertility determines whether ectomycorrhizal fungi accelerate or decelerate decomposition in a temperate forest.

Ectomycorrhizal (ECM) fungi can both accelerate and decelerate decomposition of organic matter in forest soils, but a mechanistic understanding of this differential influence is limited. Here, we tested how ECM fungi affect decomposition along a natural fertility gradient in a temperate forest of European beech. Trees were girdled to reduce belowground carbon supply to the soil. Girdling shifted soil fungal community composition and decreased hyphal biomass production and soil CO2 efflux, indicating a reduced ECM fungal activity. Girdling also affected decomposition processes, but the effects depended on fertility. Our results indicate that ECM fungi decelerate decomposition under conditions of low fertility while under conditions of high fertility ECM fungi and their host roots have an accelerating effect. We conclude that both acceleration and deceleration of decomposition of organic matter by ECM fungi can occur within a forest, with soil fertility determining the direction and magnitude of these effects. We suggest a positive feedback between fertility, stand productivity and soil carbon and nitrogen dynamics that is mediated to a large extent by ECM fungi.

Scions impact biomass allocation and root enzymatic activity of rootstocks in grafted melon and watermelon plants.

Vegetable grafting is increasingly recognized as an effective and sustainable plant production alternative. Grafted plants usually show increased uptake of water and minerals compared with self-rooted plants, mostly thought a consequence of the vigorous rootstocks selected. However, while studies frequently addressed the effects of rootstocks on the performance of scions, knowledge on the influences of scions on biomass allocation, morphology, and metabolic activity of roots is rare. In particular, the plasticity of root traits affecting resource acquisition and its efficiency remains poorly understood. Two different rootstock species, Cucurbita maxima × Cucurbita moschata and Lagenaria siceraria, were grafted in combination with melon (Cucumis melo) and watermelon (Citrullus lanatus). Self-grafted rootstocks were used as control. Plant biomass and root traits were determined after destructive harvesting 30 and/or 60 days after grafting. Traits included biomass allocation, leaf and root morphology, potential activities of four extracellular enzymes on root tips and basal root segments, and root respiration. Successfully grafted scions increase the ratio of root to whole plant dry matter (RMF), and increased ratios of root length to whole plant dry matter (RLR) and to plant leaf area (RL : LA). In contrast, morphological root traits such as diameter, tissue density, and specific root length remain surprisingly stable, and thus scion-induced changes of those traits may only play a minor role for the beneficial effects of grafting in Cucurbitaceae. Incompatibility in melon/L. siceraria grafts, however, was likely responsible for the reduced root growth in combination with clear changes in root morphological traits. Reduced root respiration rates seem to be the effects of a non-compatible rootstock-scion combination rather than an active, C-efficiency increasing acclimation. In contrast, heterografts with melon and watermelon frequently resulted in root-stock-specific, often enhanced potential enzymatic activities of acid phosphatase, ß-glucosidase, leucine-amino-peptidase, and N-acetyl-glucosaminidase both at root tips and basal parts of lateral roots-presenting a potential and complementary mechanism of grafted plants to enhance nutrient foraging. The studied melon and watermelon scions may thus increase the nutrient foraging capacity of grafted plants by fostering the relative allocation of C to the root system, and enhancing the extracellular enzymatic activities governed by roots or their rhizobiome.

Soil geochemistry - and not topography - as a major driver of carbon allocation, stocks, and dynamics in forests and soils of African tropical montane ecosystems.

The lack of field-based data in the tropics limits our mechanistic understanding of the drivers of net primary productivity (NPP) and allocation. Specifically, the role of local edaphic factors - such as soil parent material and topography controlling soil fertility as well as water and nutrient fluxes - remains unclear and introduces substantial uncertainty in understanding net ecosystem productivity and carbon (C) stocks. Using a combination of vegetation growth monitoring and soil geochemical properties, we found that soil fertility parameters reflecting the local parent material are the main drivers of NPP and C allocation patterns in tropical montane forests, resulting in significant differences in below- to aboveground biomass components across geochemical (soil) regions. Topography did not constrain the variability in C allocation and NPP. Soil organic C stocks showed no relation to C input in tropical forests. Instead, plant C input seemingly exceeded the maximum potential of these soils to stabilize C. We conclude that, even after many millennia of weathering and the presence of deeply developed soils, above- and belowground C allocation in tropical forests, as well as soil C stocks, vary substantially due to the geochemical properties that soils inherit from parent material.

Diversity and composition of farm plantation tree/shrub species along altitudinal gradients in North-eastern Ethiopia: implication for conservation.

On-farm tree plantation is a form of land use where trees are planted at the edge or interspersed with crops. It has been practiced in different parts of Ethiopia due to its contribution to the household economy and soil fertility. This study was carried out to evaluate the variation in tree/shrub plantations along altitudinal gradients and plantation niches, and farmers' on-farm tree plantation practices at Kobo and Guba Lafto districts, North-eastern Ethiopia. Transect walks and semi-structured questionnaire were administered to appraise farmers' tree/shrub plantation practices and compositions between August and December 2020. A total of 135 plots along altitudinal gradients (Forty-five sample plots per altitude) and 135 retrieved questionnaires (45 per altitude) were analyzed. At each plot, tree/shrub richness, diversity, stem density, and important value index (IVI) were computed. Multivariate analysis, descriptive statistics, and preference rankings were used to evaluate vegetation data and farmers' perceptions on tree/shrub plantations. The results showed that most farmers (78.5 %) integrate trees with their crops for household use and soil fertility maintenance. The multivariate analysis revealed a significant reduction in the number of taxa, stem density, richness, and diversity with increasing elevation, from homestead to the boundary and on-farm plantation niches. Ziziphus spina-christi and Cordia africana were the most preferred tree species; Fabaceae was the dominant family representing 18.9 % of the species. The results emphasized considerable variations in relative density, relative dominance, and important value index (IVI) across altitudinal gradients and plantation niches. Acacia seyal and Z. spina-christi contributed the highest IVI at lower and middle elevations, whereas Eucalyptus globulus had high IVI at a higher elevation. In the study districts, the distribution of multifunctional indigenous tree plantations gradually decreases with the entire altitudinal gradients compared to exotic trees/shrubs. This calls for substantial efforts on the propagation and conservation of native tree and shrub genetic resources.

The Who or the How? Species vs. Ecosystem Function Priorities in Conservation Ecology.

Current conservation strategies are targeted at preserving species, without explicitly aiming at the maintenance of ecosystem functions. In a physically highly connected world, the unintentional relocation of terrestrial, marine, and microbial life is therefore unavoidable and has been an integral part of human evolution for thousands of years. Here, we challenge the default perception often shared among conservation ecologists that preserving native species at all costs and reducing the number of exotic species and their abundance is the only way to conservation and restoration success. While this strategy is valuable in cases where exotic species disrupt ecological function, there are examples where exotic species have similar functional traits to the threatened or extinct native species and can in fact help maintain the overall or target function of an ecosystem. In the race to cope with global environmental change, we argue that ecosystem function and ecosystem services need to be viewed not only through a taxonomic lens, but increasingly also through a functional, trait-based one.

Diversity and Interrelations Among the Constitutive VOC Emission Blends of Four Broad-Leaved Tree Species at Seedling Stage.

Volatile organic compounds (VOCs) emitted by plants consist of a broad range of gasses which serve purposes such as protecting against herbivores, communicating with insects and neighboring plants, or increasing the tolerance to environmental stresses. Evidence is accumulating that the composition of VOC blends plays an important role in fulfilling these purposes. Constitutional emissions give insight into species-specific stress tolerance potentials and are an important first step in linking metabolism and function of co-occurring VOCs. Here, we investigate the blend composition and interrelations among co-emitted VOCs in unstressed seedlings of four broad-leaved tree species, Quercus robur, Fagus sylvatica, Betula pendula, and Carpinus betulus. VOCs of Q. robur and F. sylvatica mainly emitted isoprene and monoterpenes, respectively. B. pendula had relatively high sesquiterpene emission; however, it made up only 1.7% of its total emissions while the VOC spectrum was dominated by methanol (∼72%). C. betulus was emitting methanol and monoterpenes in similar amounts compared to other species, casting doubt on its frequent classification as a close-to-zero VOC emitter. Beside these major VOCs, a total of 22 VOCs could be identified, with emission rates and blend compositions varying drastically between species. A principal component analysis among species revealed co-release of multiple compounds. In particular, new links between pathways and catabolites were indicated, e.g., correlated emission rates of methanol, sesquiterpenes (mevalonate pathway), and green leaf volatiles (hexanal, hexenyl acetate, and hexenal; lipoxygenase pathway). Furthermore, acetone emissions correlated with eugenol from the Shikimate pathway, a relationship that has not been described before. Our results thus indicate that certain VOC emissions are highly interrelated, pointing toward the importance to improve our understanding of VOC blends rather than targeting dominant VOCs only.

A starting guide to root ecology: strengthening ecological concepts and standardising root classification, sampling, processing and trait measurements.

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.

Soil fertility relates to fungal-mediated decomposition and organic matter turnover in a temperate mountain forest.

Fungi are known to exert a significant influence over soil organic matter (SOM) turnover, however understanding of the effects of fungal community structure on SOM dynamics and its consequences for ecosystem fertility is fragmentary. Here we studied soil fungal guilds and SOM decomposition processes along a fertility gradient in a temperate mountain beech forest. High-throughput sequencing was used to investigate fungal communities. Carbon and nitrogen stocks, enzymatic activity and microbial respiration were measured. While ectomycorrhizal fungal abundance was not related to fertility, saprotrophic ascomycetes showed higher relative abundances under more fertile conditions. The activity of oxidising enzymes and respiration rates in mineral soil were related positively to fertility and saprotrophic fungi. In addition, organic layer carbon and nitrogen stocks were lower on the more fertile plots, although tree biomass and litter input were higher. Together, the results indicated a faster SOM turnover at the fertile end of the gradient. We suggest that there is a positive feedback mechanism between SOM turnover and fertility that is mediated by soil fungi to a significant extent. By underlining the importance of fungi for soil fertility and plant growth, these findings furthermore emphasise the dependency of carbon cycling on fungal communities below ground.

High Fungal Diversity but Low Seasonal Dynamics and Ectomycorrhizal Abundance in a Mountain Beech Forest.

Forests on steep slopes constitute a significant proportion of European mountain areas and are important as production and protection forests. This study describes the soil fungal community structure in a European beech-dominated mountain forest stands in the Northern Calcareous Alps and investigates how it is determined by season and soil properties. Samples were collected at high spatial resolution in an area of ca. 100 m × 700 m in May (spring) and August (summer). Illumina MiSeq high-throughput sequencing of the ITS2-region revealed distinct patterns for the soil fungal communities. In contrast to other studies from temperate European beech forest stands, Ascomycota dominated the highly diverse fungal community, while ectomycorrhizal fungi were of lower abundance. Russulaceae, which are often among the dominant ectomycorrhizal fungi associated with European beech, were absent from all samples. Potentially plant pathogenic fungi were more prevalent than previously reported. Only subtle seasonal differences were found between fungal communities in spring and summer. Especially, dominant saprotrophic taxa were largely unaffected by season, while slightly stronger effects were observed for ectomycorrhizal fungi. Soil characteristics like pH and organic carbon content, on the other hand, strongly shaped abundant taxa among the saprotrophic fungal community.

Root traits as drivers of plant and ecosystem functioning: current understanding, pitfalls and future research needs.

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.

Effects of Increased N Deposition on Leaf Functional Traits of Four Contrasting Tree Species in Northeast China.

Northeast China is persistently affected by heavy nitrogen (N) deposition. Studying the induced variation in leaf traits is pivotal to develop an understanding of the adaptive plasticity of affected species. This study thus assesses effects of increased N deposition on leaf morphological and anatomical traits and their correlation among and with biomass allocation patterns. A factorial experiment was conducted utilizing seedlings of two gymnosperms (Larix gmelinii, Pinus koraiensis) and two angiosperms (Fraxinus mandshurica, Tilia amurensis). Leaf mass per area and leaf density decreased and leaf thickness increased under high N deposition but trait interrelations remained stable. In gymnosperms, leaf mass per area was correlated to both leaf thickness and area, while being correlated to leaf density only in angiosperms. Epidermis, mesophyll thickness, conduit and vascular bundle diameter increased. Despite the differences in taxonomic groups and leaf habits, the common patterns of variation suggest that a certain degree of convergence exists between the species' reaction towards N deposition. However, stomata pore length increased in angiosperms, and decreased in gymnosperms under N deposition. Furthermore, biomass and leaf mass fraction were correlated to leaf traits in gymnosperms only, suggesting a differential coordination of leaf traits and biomass allocation patterns under high N deposition per taxonomic group.

Faba Bean Cultivation - Revealing Novel Managing Practices for More Sustainable and Competitive European Cropping Systems.

Faba beans are highly nutritious because of their high protein content: they are a good source of mineral nutrients, vitamins, and numerous bioactive compounds. Equally important is the contribution of faba bean in maintaining the sustainability of agricultural systems, as it is highly efficient in the symbiotic fixation of atmospheric nitrogen. This article provides an overview of factors influencing faba bean yield and quality, and addresses the main biotic and abiotic constraints. It also reviews the factors relating to the availability of genetic material and the agronomic features of faba bean production that contribute to high yield and the improvement of European cropping systems. Emphasis is to the importance of using new high-yielding cultivars that are characterized by a high protein content, low antinutritional compound content, and resistance to biotic and abiotic stresses. New cultivars should combine several of these characteristics if an increased and more stable production of faba bean in specific agroecological zones is to be achieved. Considering that climate change is also gradually affecting many European regions, it is imperative to breed elite cultivars that feature a higher abiotic-biotic stress resistance and nutritional value than currently used cultivars. Improved agronomical practices for faba bean crops, such as crop establishment and plant density, fertilization and irrigation regime, weed, pest and disease management, harvesting time, and harvesting practices are also addressed, since they play a crucial role in both the production and quality of faba bean.