The biodiversity - N cycle relationship: a 15N tracer experiment with soil from plant mixtures of varying diversity to model N pool sizes and transformation rates

We conducted a 15N tracer experiment in laboratory microcosms with field-fresh soil samples from a biodiversity experiment to evaluate the relationship between grassland biodiversity and N cycling. To embrace the complexity of the N cycle, we determined N exchange between five soil N pools (labile and recalcitrant organic N, dissolved NH4+ and NO3− in soil solution, and exchangeable NH4+) and eight N transformations (gross N mineralization from labile and recalcitrant organic N, NH4+ immobilization into labile and recalcitrant organic N, autotrophic nitrification, heterotrophic nitrification, NO3− immobilization, adsorption of NH4+) expected in aerobic soils with the help of the N-cycle model Ntrace. We used grassland soil of the Jena Experiment, which includes plant mixtures with 1 to 60 species and 1 to 4 functional groups (legumes, grasses, tall herbs, small herbs). The 19 soil samples of one block of the Jena Experiment were labeled with either 15NH4+ or 15NO3- or both. In the presence of legumes, gross N mineralization and autotrophic nitrification increased significantly because of higher soil N concentrations in legume-containing plots and high microbial activity. Similarly, the presence of grasses significantly increased the soil NH4+ pool, gross N mineralization, and NH4+ immobilization, likely because of enhanced microbial biomass and activity by providing large amounts of rhizodeposits through their dense root systems. In our experiment, previously reported plant species richness effects on the N cycle, observed in a larger-scale field experiment within the Jena Experiment, were not seen. However, specific plant functional groups had a significant positive impact on the N cycling in the incubated soil samples.

Modeling How Community Assembly Alters the Functioning of Ecosystems

Although the effects of species richness on ecosystem functioning have been extensively studied, there is increased interest in understanding how community assembly in a more general sense can alter the functioning of ecosystems. We identify two complementary approaches for evaluating how community assembly influences biomass production. The first quantifies the relative importance of complementarity and selection by contrasting monocultures with polycultures. The second identifies the effects of species replacements (losses and/or gains of species relative to the baseline polyculture) as well as the indirect effects on other species’ productivity. We compare and contrast these two approaches using simulated communities where species compete for different resource types and experience different levels of environmental heterogeneity and find that these metrics can jointly diagnose the mechanisms of competition driving productivity. We apply our methods to data from a long-term biodiversity-ecosystem experiment (the Jena Experiment) and find that the data do not correspond to any of the competition scenarios we modeled. We address two additional possible complications: facilitation by nitrogen fixing plants, and non-equilibrium behavior during community assembly, and find that facilitation/competition by nitrogen fixing plants is the more likely explanation for the results obtained at Jena.

Biotic interactions, community assembly, and eco-evolutionary dynamics as drivers of long-term biodiversity–ecosystem functioning relationships

The functioning and service provisioning of ecosystems in the face of anthropogenic environmental and biodiversity change is a cornerstone of ecological research. The last three decades of biodiversity–ecosystem functioning (BEF) research have provided compelling evidence for the significant positive role of biodiversity in the functioning of many ecosystems. Despite broad consensus of this relationship, the underlying ecological and evolutionary mechanisms have not been well understood. This complicates the transition from a description of patterns to a predictive science. The proposed Research Unit aims at filling this gap of knowledge by applying novel experimental and analytical approaches in one of the longest-running biodiversity experiments in the world: the Jena Experiment. The central aim of the Research Unit is to uncover the mechanisms that determine BEF relationships in the short- and in the long-term. Increasing BEF relationships with time in long-term experiments do not only call for a paradigm shift in the appreciation of the relevance of biodiversity change, they likely are key to understanding the mechanisms of BEF relationships in general. The subprojects of the proposed Research Unit fall into two tightly linked main categories with two research areas each that aim at exploring variation in community assembly processes and resulting differences in biotic interactions as determinants of the long-term BEF relationship. Subprojects under “Microbial community assembly” and “Assembly and functions of animal communities” mostly focus on plant diversity effects on the assembly of communities and their feedback effects on biotic interactions and ecosystem functions. Subprojects under “Mediators of plant-biotic interactions” and “Intraspecific diversity and micro-evolutionary changes” mostly focus on plant diversity effects on plant trait expression and micro-evolutionary adaptation, and subsequent feedback effects on biotic interactions and ecosystem functions. This unification of evolutionary and ecosystem processes requires collaboration across the proposed subprojects in targeted plant and soil history experiments using cutting-edge technology and will produce significant synergies and novel mechanistic insights into BEF relationships. The Research Unit of the Jena Experiment is uniquely positioned in this context by taking an interdisciplinary and integrative approach to capture whole-ecosystem responses to changes in biodiversity and to advance a vibrant research field.