Sibylle Steinbeiss

Plant diversity effects on soil microorganisms support the singular hypothesis

The global decline in biodiversity has generated concern over the consequences for ecosystem functioning and services. Although ecosystem functions driven by soil microorganisms such as plant productivity, decomposition, and nutrient cycling are of particular importance, interrelationships between plant diversity and soil microorganisms are poorly understood. We analyzed the response of soil microorganisms to variations in plant species richness (1-60) and plant functional group richness (1-4) in an experimental grassland system over a period of six years. Major abiotic and biotic factors were considered for exploring the mechanisms responsible for diversity effects. Further, microbial growth characteristics were assessed following the addition of macronutrients. Effects of plant diversity on soil microorganisms were most pronounced in the most diverse plant communities though differences only became established after a time lag of four years. Differences in microbial growth characteristics indicate successional changes from a disturbed (zymogeneous) to an established (autochthonous) microbial community four years after establishment of the experiment. Supporting the singular hypothesis for plant diversity, the results suggest that plant species are unique, each contributing to the functioning of the belowground system. The results reinforce the need for long-term biodiversity experiments to fully appreciate consequences of current biodiversity loss for ecosystem functioning.

Plant diversity increases soil microbial activity and soil carbon storage

Plant diversity strongly influences ecosystem functions and services, such as soil carbon storage. However, the mechanisms underlying the positive plant diversity effects on soil carbon storage are poorly understood. We explored this relationship using long-term data from a grassland biodiversity experiment (The Jena Experiment) and radiocarbon ((14)C) modelling. Here we show that higher plant diversity increases rhizosphere carbon inputs into the microbial community resulting in both increased microbial activity and carbon storage. Increases in soil carbon were related to the enhanced accumulation of recently fixed carbon in high-diversity plots, while plant diversity had less pronounced effects on the decomposition rate of existing carbon. The present study shows that elevated carbon storage at high plant diversity is a direct function of the soil microbial community, indicating that the increase in carbon storage is mainly limited by the integration of new carbon into soil and less by the decomposition of existing soil carbon.

Foliar and soil δ15N values reveal increased nitrogen partitioning among species in diverse grassland communities

Plant and soil nitrogen isotope ratios (δ¹⁵N) were studied in experimental grassland plots of varying species richness. We hypothesized that partitioning of different sources of soil nitrogen among four plant functional groups (legumes, grasses, small herbs, tall herbs) should increase with diversity. Four years after sowing, all soils were depleted in ¹⁵N in the top 5 cm whereas in non-legume plots soils were enriched in ¹⁵N at 5-25 cm depth. Decreasing foliar δ¹⁵N and Δδ¹⁵N (= foliar δ¹⁵N-soil δ¹⁵N) values in legumes indicated increasing symbiotic N₂ fixation with increasing diversity. In grasses, foliar Δδ¹⁵N also decreased with increasing diversity suggesting enhanced uptake of N depleted in ¹⁵N. Foliar Δδ¹⁵N values of small and tall herbs were unaffected by diversity. Foliar Δδ¹⁵N values of grasses were also reduced in plots containing legumes, indicating direct use of legume-derived N depleted in ¹⁵N. Increased foliar N concentrations of tall and small herbs in plots containing legumes without reduced foliar δ¹⁵N indicated that these species obtained additional mineral soil N that was not consumed by legumes. These functional group and species specific shifts in the uptake of different N sources with increasing diversity indicate complementary resource use in diverse communities.

Soil Environmental Conditions and Microbial Build-Up Mediate the Effect of Plant Diversity on Soil Nitrifying and Denitrifying Enzyme Activities in Temperate Grasslands

Random reductions in plant diversity can affect ecosystem functioning, but it is still unclear which components of plant diversity (species number – namely richness, presence of particular plant functional groups, or particular combinations of these) and associated biotic and abiotic drivers explain the observed relationships, particularly for soil processes. We assembled grassland communities including 1 to 16 plant species with a factorial separation of the effects of richness and functional group composition to analyze how plant diversity components influence soil nitrifying and denitrifying enzyme activities (NEA and DEA, respectively), the abundance of nitrifiers (bacterial and archaeal amoA gene number) and denitrifiers (nirK, nirS and nosZ gene number), and key soil environmental conditions. Plant diversity effects were largely due to differences in functional group composition between communities of identical richness (number of sown species), though richness also had an effect per se. NEA was positively related to the percentage of legumes in terms of sown species number, the additional effect of richness at any given legume percentage being negative. DEA was higher in plots with legumes, decreased with increasing percentage of grasses, and increased with richness. No correlation was observed between DEA and denitrifier abundance. NEA increased with the abundance of ammonia oxidizing bacteria. The effect of richness on NEA was entirely due to the build-up of nitrifying organisms, while legume effect was partly linked to modified ammonium availability and nitrifier abundance. Richness effect on DEA was entirely due to changes in soil moisture, while the effects of legumes and grasses were partly due to modified nitrate availability, which influenced the specific activity of denitrifiers. These results suggest that plant diversity-induced changes in microbial specific activity are important for facultative activities such as denitrification, whereas changes in microbial abundance play a major role for non-facultative activities such as nitrification.

Plant effects on soil N mineralization are mediated by the composition of multiple soil organic fractions

Despite the topic of soil nitrogen (N) mineralization being well-studied, very few studies have addressed the relative contribution of different plant and soil variables in influencing soil N mineralization rates, and thus the supply of inorganic N to plants. Here, we used data from a well-studied N-limited grassland to address the relative effects of six plant and soil variables on net and on gross rates of soil N mineralization. We also addressed whether plant effects on soil N mineralization were mediated by changes in C and N concentrations of multiple soil organic matter (SOM) fractions. Regression analyses show that key plant traits (i.e., plant C:N ratios and total root mass) were more important than total C and N concentrations of bulk soil in influencing N mineralization. This was mainly because plant traits influenced the C and N concentration (and C:N ratios) of different SOM fractions, which in turn were significantly associated with changes in net and gross N mineralization. In particular, C:N ratios of a labile soil fraction were negatively related to net soil N mineralization rates, whereas total soil C and N concentrations of more recalcitrant fractions were positively related to gross N mineralization. Our study suggests that changes in belowground N-cycling can be better predicted by simultaneously addressing how plant C:N ratios and root mass affect the composition and distribution of different SOM pools in N-limited grassland systems.