In the past two decades, a large number of studies have investigated the relationship between biodiversity and ecosystem functioning, most of which focussed on a limited set of ecosystem variables. The Jena Experiment was set up in 2002 to investigate the effects of plant diversity on element cycling and trophic interactions, using a multi-disciplinary approach. Here, we review the results of 15 years of research in the Jena Experiment, focussing on the effects of manipulating plant species richness and plant functional richness. With more than 85,000 measures taken from the plant diversity plots, the Jena Experiment has allowed answering fundamental questions important for functional biodiversity research. First, the question was how general the effect of plant species richness is, regarding the many different processes that take place in an ecosystem. About 45% of different types of ecosystem processes measured in the ‘main experiment’, where plant species richness ranged from 1 to 60 species, were significantly affected by plant species richness, providing strong support for the view that biodiversity is a significant driver of ecosystem functioning. Many measures were not saturating at the 60-species level, but increased linearly with the logarithm of species richness. There was, however, great variability in the strength of response among different processes. One striking pattern was that many processes, in particular belowground processes, took several years to respond to the manipulation of plant species richness, showing that biodiversity experiments have to be long-term, to distinguish trends from transitory patterns. In addition, the results from the Jena Experiment provide further evidence that diversity begets stability, for example stability against invasion of plant species, but unexpectedly some results also suggested the opposite, e.g. when plant communities experience severe perturbations or elevated resource availability. This highlights the need to revisit diversity–stability theory. Second, we explored whether individual plant species or individual plant functional groups, or biodiversity itself is more important for ecosystem functioning, in particular biomass production. We found strong effects of individual species and plant functional groups on biomass production, yet these effects mostly occurred in addition to, but not instead of, effects of plant species richness. Third, the Jena Experiment assessed the effect of diversity on multitrophic interactions. The diversity of most organisms responded positively to increases in plant species richness, and the effect was stronger for above- than for belowground organisms, and stronger for herbivores than for carnivores or detritivores. Thus, diversity begets diversity. In addition, the effect on organismic diversity was stronger than the effect on species abundances. Fourth, the Jena Experiment aimed to assess the effect of diversity on N, P and C cycling and the water balance of the plots, separating between element input into the ecosystem, el...
The widely observed positive relationship between plant diversity and ecosystem functioning is thought to be substantially driven by complementary resource use of plant species. Recent work suggests that biotic interactions among plants and between plants and soil organisms drive key aspects of resource use complementarity. Here, we provide a conceptual framework for integrating positive biotic interactions across guilds of organisms, more specifically between plants and mycorrhizal types, to explain resource use complementarity in plants and its consequences for plant competition. Our overarching hypothesis is that ecosystem functioning increases when more plant species associate with functionally dissimilar mycorrhizal fungi because differing mycorrhizal types will increase coverage of habitat space for and reduce competition among plants. We introduce a recently established field experiment (MyDiv) that uses different pools of tree species that associate with either arbuscular or ectomycorrhizal fungi to create orthogonal experimental gradients in tree species richness and mycorrhizal associations and present initial results. Finally, we discuss options for future mechanistic studies on resource use complementarity within MyDiv. We show how mycorrhizal types and biotic interactions in MyDiv can be used in the future to test novel questions regarding the mechanisms underlying biodiversity–ecosystem function relationships.
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 mankind depends upon. In this paper, we argue that a multitrophic perspective of biotic interactions in random and nonrandom 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 Eisenhauer et al.
Research on mycorrhizal interactions has traditionally developed into separate disciplines addressing different organizational levels. This separation has led to an incomplete understanding of mycorrhizal functioning. Integration of mycorrhiza research at different scales is needed to understand the mechanisms underlying the context dependency of mycorrhizal associations, and to use mycorrhizae for solving environmental issues. Here, we provide a road map for the integration of mycorrhiza research into a unique framework that spans genes to ecosystems. Using two key topics, we identify parallels in mycorrhiza research at different organizational levels. Based on two current projects, we show how scientific integration creates synergies, and discuss future directions. Only by overcoming disciplinary boundaries, we will achieve a more comprehensive understanding of the functioning of mycorrhizal associations.
The extensive use of traits in ecological studies over the last few decades to predict community functions has revealed that plant traits are plastic and respond to various environmental factors. These plant traits are assumed to predict how plants compete and capture resources. Variation in stoichiometric ratios both within and across species reflects resource capture dynamics under competition. However, the impact of local plant diversity on species‐specific stoichiometry remains poorly studied. Here, we analyze how spatial and temporal diversity in resource‐acquisition traits affects leaf elemental stoichiometry of plants (i.e. the result of resource capture) and how flexible this stoichiometry is depending on the functional composition of the surrounding community. Therefore, we assessed inter‐ and intraspecific variations of leaf carbon (C), nitrogen (N), and phosphorus (P) (and their ratios) of 20 grassland species in a large trait‐based plant diversity experiment located in Jena (Germany) by measuring leaf elemental concentrations at the species‐level along a gradient in plant trait dissimilarity. Our results show that plants showed large intra‐ and interspecific variation in leaf stoichiometry, which was only partly explained by the functional group identity (grass or herb) of the species. Elemental concentrations (N, P, but not C) decreased with plant species richness, and species tended to become more deviant from their monoculture stoichiometry with increasing trait dissimilarity in the community. These responses differed among species, some consistently increased or decreased in P and N concentrations; for other species, the negative or positive change in P and N concentrations increased with increasing trait difference between the target species and the remaining community. The strength of this relationship was significantly associated to the relative position of the species along trait gradients related to resource acquisition. Trait‐difference and trait‐diversity thus were important predictors of how species’ resource capture changed in competitive neighbourhoods.
Abstract.Research on the functional importance of biodiversity, motivated by global species loss, has documented that plant species richness affects many plant-related ecosystem functions. Less is known about the effects of plant species richness on functions related to higher trophic levels, such as the consumption of biomass by animals, that is, herbivory. Previous studies have shown positive, neutral, or negative effects of plant species richness on herbivory. In the framework of a grassland biodiversity experiment (the Jena Experiment), we investigated herbivory (the proportion of leaf area damaged and the amount of leaf biomass consumed by arthropod herbivores) along two experimental gradients of plant species richness ranging from 1 to 60 species (Main Experiment) and from 1 to 8 species (Trait-Based Experiment) biannually for five and three years, respectively. Additionally, plant functional diversity, based on traits related to plant growth, was manipulated as the number of functional groups in a community (Main Experiment) or a gradient of functional trait dissimilarity (Trait-Based Experiment). Herbivory at the level of plant communities ranged from 0% to 31% (0 and 33.8 g/m 2 ) in the Main Experiment and 0% to 8% (0 and 13.7 g/m 2 ) in the Trait-Based Experiment, and it was on average higher in summer than in spring. For both experimental gradients and all years investigated, we found a consistent increase in damaged leaf area and consumed biomass with increasing plant species richness. As mechanistic explanations for effects of plant species richness, we propose changes in plant quality and herbivore communities. The presence of specific plant functional groups significantly affected herbivory, likely related to traits affecting plant defense and nutritional value, but we found little evidence for effects of plant functional diversity. The general positive relationship between plant species richness and herbivory might contribute to effects of plant species richness on other ecosystem functions such as productivity and nutrient mineralization and can cascade up the food web also affecting higher trophic levels.
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