Leonardo Bassi

A system's functioning is influenced not only by resource availability but also, likely, by resource inequality. In ecological systems most experimental manipulations of resource availability typically focused on homogeneous resource addition, or spatial heterogeneity of resources in a qualitative sense. Here, we test whether resource inequality per se influences the diversity and functioning of model grassland communities in an Ecotron experiment. Specifically, we experimentally manipulated resource area and concentration (two quantitative components of heterogeneity) to create an inequality gradient quantified by the Gini coefficient. Moreover, we examine whether the association with mycorrhizal fungi can mediate the influence of resource inequality on nutrient dynamics between soils, plants and leachate. We show that resource inequality significantly increased phosphorus leaching and reduced plant phosphorus pools aboveground. Some plant species were more sensitive to inequality than others, as there were shifts in the relative dominance of the two species that produced the most biomass (Plantago lanceolata and Holcus lanatus). Taken together, shifts in plant community composition may provide cues that indicate changes in nutrient dynamics; even though overall plant biomass was not affected, and plant diversity showed a significant decline only during the first phase of the experiment. The enrichment with mycorrhiza did not significantly influence plant biomass and diversity or nutrient concentrations above‐ and belowground. These results demonstrate that inequality is an essential component in ecological systems determining nutrient availability for plants as well as nutrient use efficiency of plants. An important implication is that spatial inequality of resources in natural landscapes may alter community compositions and predictions of phosphorus uptake that are based on resource input alone.

The global loss of biodiversity has motivated many studies that experimentally vary plant species richness and examine the consequences for ecosystem functioning. Such experiments generally show a positive relationship between above- and below-ground biodiversity and the functioning of terrestrial ecosystems. Moreover, this relationship tends to strengthen over time, seen as enhanced functioning of diverse plant communities and reduced functioning of low-diversity plant communities. Differences in multitrophic community assembly and biotic interactions in high- versus low-diversity plant communities are hypothesized to affect plant performance by altering consumer community structure and function and driving plastic or micro-evolutionary responses of plant species in the plant communities. To resolve this complex interplay of community history, we separated these effects into plant and soil history. Plant history refers to all plant-level responses to past abiotic and biotic selection pressures experienced in their communities, while soil history relates to all abiotic and biotic soil properties developed as a legacy of plant-soil interactions under variable plant diversity. We set up a biodiversity experiment in an Ecotron, a terrestrial mesocosm facility that allows controlling environmental conditions above- and below-ground, to test whether the strengthening biodiversity-ecosystem functioning relationship is due to soil history, plant history, or a combination of both. We established a plant diversity gradient consisting of 1, 2, 3, and 6 grassland plant species and factorially nested with soil history and plant history treatments for each level of plant species richness. Representative results demonstrate the successful establishment of target treatments in the Ecotron experiment, observing the effects of plant and soil history on initial plant development and final plant growth. Additionally, we provide a case study for data analysis of individual response variables. We outline research objectives and methods to comprehensively assess the multifunctional responses to the experimental treatments necessary to ultimately address the overarching hypothesis.

Summary: The increased positive impact of plant diversity on ecosystem functioning is often attributed to the accumulation of mutualists and dilution of antagonists in diverse plant communities. While increased plant diversity alters traits related to resource acquisition, it remains unclear whether it reduces defence allocation, whether this reduction differs between roots and leaves, or varies among species. To answer these questions, we assessed the effect of plant species richness, plant species identity and their interaction on the expression of 23 physical and chemical leaf and fine root defence traits of 16 plant species in a 19‐yr‐old biodiversity experiment. Only leaf mass per area, leaf and root dry matter content and root nitrogen, traits associated with both, resource acquisition and defence, responded consistently to species richness. However, species richness promoted a decoupling of these defences in leaves and fine roots, possibly in response to resource limitations in diverse communities. Species‐specific responses were rare and related to chemical defence and mutualist collaboration, likely responding to species‐specific antagonists' dilution and mutualists' accumulation. Overall, our study suggests that resource limitation in diverse communities might mediate the relationship between plant defence traits and antagonist dilution.

The relationship of plant diversity and several ecosystem functions strengthens over time. This suggests that the restructuring of biotic interactions in the process of a community's assembly and the associated changes in function differ between species‐rich and species‐poor communities. An important component of these changes is the feedback between plant and soil community history. In this study, we examined the interactive effects of plant richness and community history on the trophic functions of the soil fauna community. We hypothesized that experimental removal of either soil or plant community history would diminish the positive effects of plant richness on the multitrophic functions of the soil food web, compared to mature communities. We tested this hypothesis in a long‐term grassland biodiversity experiment by comparing plots across three treatments (without plant history, without plant and soil history, controls with ~20 years of plot‐specific community history). We found that the relationship between plant richness and below‐ground multitrophic functionality is indeed stronger in communities with shared plant and soil community history. Our findings indicate that anthropogenic disturbance can impact the functioning of the soil community through the loss of plant species but also by preventing feedbacks that develop in the process of community assembly.

The enhancement of biodiversity's positive impact on ecosystem functioning (BEF) over time is commonly observed and attributed to the accumulation of mutualists and dilution of antagonists in more diverse communities. If antagonists play a role in the BEF relationship, the reduction of plant antagonists in more diverse communities, could allow plants to reduce allocation to defence. This study aimed to assess the influence of plant diversity on the expression of defence traits in 16 plant species. Our hypotheses were: (1) increased plant diversity reduces allocation to defence, (2) this reduction is more pronounced in roots than in leaves, and (3) this effect varies among species.We measured both physical and chemical defence traits in leaves and fine roots across communities with varying plant species richness in a 19-year-old biodiversity experiment. Using established methods and an innovative metabolome approach, we explored the interactive effects of plant diversity and species identity on defence traits through linear mixed models.Our findings were mixed concerning the first hypothesis, with only some leaf defence traits (leaf mass per area, leaf dry matter content, and hair length) showing reduction along the diversity gradient. Unexpectedly, the values of some root traits, root tissue density and nitrogen content, suggested increased allocation to defence along the same gradient. This might be attributed to these traits serving other functions, e.g. in resource acquisition and competition, which potentially overruled the impact of declining antagonists on plant defences. Our results did not support the third hypothesis, suggesting an overall convergence responses to biotic and abiotic factors related to plant diversity after two decades.While evidence for a consistent reduction in defence trait expression along the diversity gradient was limited, our findings underscore the complex nature of BEF relationships. Further experiments, possibly controlling confounding factors on trait expression or manipulating antagonist pressures along diversity gradients, are needed to elucidate the underlying mechanisms.

Biodiversity experiments revealed that plant diversity loss can decrease ecosystem functions across trophic levels. To address why such biodiversity-function relationships strengthen over time, we established experimental mesocosms replicating a gradient in plant species richness across treatments of shared versus non-shared history of (1) the plant community and (2) the soil fauna community. After 4 months, we assessed the multitrophic functioning of soil fauna via biomass stocks and energy fluxes across the food webs. We find that soil community history significantly enhanced belowground multitrophic function via changes in biomass stocks and community-average body masses across the food webs. However, variation in plant diversity and plant community history had unclear effects. Our findings underscore the importance of long-term community assembly processes for soil fauna-driven ecosystem function, with species richness and short-term plant adaptations playing a minimal role. Disturbances that disrupt soil community stability may hinder fauna-driven ecosystem functions, while recovery may require several years.

The relationship of plant diversity and several ecosystem functions strengthens over time. This suggests that the restructuring of biotic interactions in the process of a community’s assembly and the associated changes in function differ between species-rich and species-poor communities. An important component of these changes is the feedback between plant and soil community history. In this study, we examined the interactive effects of plant richness and community history on the trophic functions of the soil fauna community. We hypothesized that experimental removal of either soil or plant community history would diminish the positive effects of plant richness on the multitrophic functions of the soil food-web, compared to mature communities. We tested this hypothesis in a long-term grassland biodiversity experiment by comparing plots across three treatments (without plant history, without plant and soil history, controls with ~20 years of plot specific community history). We found that the relationship between plant richness and belowground multitrophic functionality is indeed stronger in communities with shared plant and soil community history. Our findings indicate that anthropogenic disturbance can impact the functioning of the soil community through the loss of plant species but also by preventing feedbacks that develop in the process of community assembly.

The relationship between biodiversity and ecosystem functioning strengthens with ecosystem age. However, the interplay between the plant diversity - ecosystem functioning relationship and Glomeromycotinian arbuscular mycorrhizal fungi (AMF) community assembly has not yet been scrutinized in this context, despite AMF’s role in plant survival and niche exploration. We study the development of AMF communities by disentangling soil- and plant-driven effects from calendar year effects. Within a long-term grassland biodiversity experiment, the pre-existing plant communities of varying plant diversity were re-established as split plots with combinations of common plant and soil histories: split plots with neither common plant nor soil history, with only soil but no plant history, and with both common plant and soil history. We found that bulk soil AMF communities were primarily shaped by common soil history, and additional common plant history had little effect. Further, the steepness of AMF diversity and plant diversity relationship did not strengthen over time, but AMF community evenness increased with common history. Specialisation of AMF towards plant species was low throughout, giving no indication of AMF communities specialising or diversifying over time. The potential of bulk soil AMF as mediators of variation in plant and microbial biomass over time and hence as drivers of biodiversity and ecosystem relationships was low. Our results suggest that soil processes may be key for the build-up of plant community-specific mycorrhizal communities with likely feedback effects on ecosystem productivity, but the plant-available mycorrhizal pool in bulk soil itself does not explain the strengthening of biodiversity and ecosystem relationships over time.

Trait-based approaches have been increasingly used to relate plants to soil microbial communities. However, the plant organs mediating this plant-microbe interaction – the roots – have been largely overlooked. The recent discovery of the root economics space offers a predictive framework for the structure of soil microbial communities, and specifically soil-borne fungal communities. Applying this novel approach, our study in a grassland plant diversity experiment reveals distinct root trait strategies at the level of the plant community. In addition to significant effects of plant species richness, we show that both axes of the root economics space – the collaboration and conservation gradient – are strong drivers of the composition of the different guilds of soil fungi, including saprotrophic, plant pathogenic, and mycorrhizal fungi. Our results illustrate that the root economics space and plant species richness jointly determine the effects of plants on fungal communities and their potential role in plant health and ecosystem functioning.

The relationship between biodiversity and ecosystem functioning strengthens with ecosystem age. However, the interplay between the plant diversity - ecosystem functioning relationship and Glomeromycotinian arbuscular mycorrhizal fungi (AMF) community assembly has not yet been scrutinized in this context, despite AMF’s role in plant survival and niche exploration. We study the development of AMF communities by disentangling soil- and plant-driven effects from year effects. Within a long-term grassland biodiversity experiment, the pre-existing plant communities of varying plant diversity were re-established as split plots with combinations of common plant and soil histories: split plots with neither common plant nor soil history, with only soil but no plant history, and with both common plant and soil history. We found that bulk soil AMF communities were primarily shaped by common soil history and additional common plant history had little effect. Further, the steepness of AMF diversity and plant diversity relationship did not strengthen over time, but AMF community evenness increased with common history. Specialisation of AMF towards plant species was low throughout giving no indication of AMF communities specialising or diversifying over time. The potential of bulk soil AMF as mediators of variation in plant and microbial biomass over time and hence as drivers of BEF relationships was low. Our results suggest that soil processes may be key for the build-up of plant community-specific mycorrhizal communities with likely feedback effects on ecosystem productivity, but the plant-available mycorrhizal pool in bulk soil itself does not explain the strengthening of BEF relationships over time.