▪ Abstract  We explore empirical and theoretical evidence for the functional significance of plant-litter diversity and the extraordinary high diversity of decomposer organisms in the process of litter decomposition and the consequences for biogeochemical cycles. Potential mechanisms for the frequently observed litter-diversity effects on mass loss and nitrogen dynamics include fungi-driven nutrient transfer among litter species, inhibition or stimulation of microorganisms by specific litter compounds, and positive feedback of soil fauna due to greater habitat and food diversity. Theory predicts positive effects of microbial diversity that result from functional niche complementarity, but the few existing experiments provide conflicting results. Microbial succession with shifting enzymatic capabilities enhances decomposition, whereas antagonistic interactions among fungi that compete for similar resources slow litter decay. Soil-fauna diversity manipulations indicate that the number of trophic levels, species identity, and the presence of keystone species have a strong impact on decomposition, whereas the importance of diversity within functional groups is not clear at present. In conclusion, litter species and decomposer diversity can significantly influence carbon and nutrient turnover rates; however, no general or predictable pattern has emerged. Proposed mechanisms for diversity effects need confirmation and a link to functional traits for a comprehensive understanding of how biodiversity interacts with decomposition processes and the consequences of ongoing biodiversity loss for ecosystem functioning.

A; et al., (2010). Bottom-up effects of plant diversity on multitrophic interactions in a biodiversity experiment. Nature, (468) Bottom-up effects of plant diversity on multitrophic interactions in a biodiversity experiment Abstract Biodiversity is rapidly declining1, and this may negatively affect ecosystem processes, including economically important ecosystem services. Previous studies have shown that biodiversity has positive effects on organisms and processes4 across trophic levels. However, only a few studies have so far incorporated an explicit food-web perspective. In an eight-year biodiversity experiment, we studied an unprecedented range of above-and below-ground organisms and multitrophic interactions. A multitrophic data set originating from a single long-term experiment allows mechanistic insights that would not be gained from meta-analysis of different experiments. Here we show that plant diversity effects dampen with increasing trophic level and degree of omnivory. This was true both for abundance and species richness of organisms. Furthermore, we present comprehensive above-ground/below-ground biodiversity food webs. Both above ground and below ground, herbivores responded more strongly to changes in plant diversity than did carnivores or omnivores. Density and richness of carnivorous taxa was independent of vegetation structure. Below-ground responses to plant diversity were consistently weaker than above-ground responses. Responses to increasing plant diversity were generally positive, but were negative for biological invasion, pathogen infestation and hyperparasitism. Our results suggest that plant diversity has strong bottom-up effects on multitrophic interaction networks, with particularly strong effects on lower trophic levels. Effects on higher trophic levels are indirectly mediated through bottom-up trophic cascades. However, only few studies so far incorporated an explicit food-web perspective 6 . In an 8-50 year biodiversity experiment, we studied an unprecedented range of above-and 51 belowground organisms and biotic interactions. A multitrophic dataset originating from 52 a single long-term experiment allows mechanistic insights that otherwise would not be 53 possible using meta-analysis of different experiments. We show that plant diversity 54 effects dampen with increasing trophic level and degree of omnivory. This finding was 55 consistent both for abundance and species richness of organisms. Further, we present 56 comprehensive above-belowground biodiversity food webs. Both above-and 57 belowground, herbivores responded more strongly to changes in plant diversity than did 58 carnivores or omnivores. Density and richness of carnivorous taxa was independent of 59 vegetation structure. Belowground responses to plant diversity were consistently weaker 60 than aboveground responses. Responses to increasing plant diversity were generally 61 positive, but negative for biological invasion, pathogen infestation and hyperparasitism. 62Our results suggest that plant diversity exhibits stron...

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 (14C) 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

The decomposition of dead organic matter is a major determinant of carbon and nutrient cycling in ecosystems, and of carbon fluxes between the biosphere and the atmosphere. Decomposition is driven by a vast diversity of organisms that are structured in complex food webs. Identifying the mechanisms underlying the effects of biodiversity on decomposition is critical given the rapid loss of species worldwide and the effects of this loss on human well-being. Yet despite comprehensive syntheses of studies on how biodiversity affects litter decomposition, key questions remain, including when, where and how biodiversity has a role and whether general patterns and mechanisms occur across ecosystems and different functional types of organism. Here, in field experiments across five terrestrial and aquatic locations, ranging from the subarctic to the tropics, we show that reducing the functional diversity of decomposer organisms and plant litter types slowed the cycling of litter carbon and nitrogen. Moreover, we found evidence of nitrogen transfer from the litter of nitrogen-fixing plants to that of rapidly decomposing plants, but not between other plant functional types, highlighting that specific interactions in litter mixtures control carbon and nitrogen cycling during decomposition. The emergence of this general mechanism and the coherence of patterns across contrasting terrestrial and aquatic ecosystems suggest that biodiversity loss has consistent consequences for litter decomposition and the cycling of major elements on broad spatial scales. Main textBiological diversity that directly influences litter decomposition exists at multiple trophic levels 4 . This diversity includes plants producing litter mixtures of varying quality, microbial decomposers, and invertebrate consumers of varying body size, which selectively exploit the heterogeneous resources provided by litter mixtures 4,13 . Efforts to derive generalities about biodiversity effects on litter decomposition have been elusive, since both pioneering work 14 and recent syntheses have highlighted contrasting effects of litter species richness on 3 decomposition [4][5][6]15,16 . In part, this variation appears to be due to site-specific conditions, including contrasts between aquatic and terrestrial ecosystems as well as geographic settings.Further differences may arise from variation in experimental protocols, selected plant species, and the types of decomposers included in a given experiment. Such methodological discrepancies have complicated syntheses across studies, hindering the emergence of common patterns and mechanisms.Here we report on the results from the first concerted biodiversity experiments on decomposition by manipulating diversity across trophic levels and distinct biomes in both forest floor and stream habitats (Extended Data Table 1). We hypothesised that functional diversity of decomposers (variation in body size) and leaf litter (variation in litter quality) promote C and N cycling across contrasting locations (subarctic to tr...

We summarize progress with respect to (1) different approaches to isolate, extract, and quantify organo‐mineral compounds from soils, (2) types of mineral surfaces and associated interactions, (3) the distribution and function of soil biota at organo‐mineral surfaces, (4) the distribution and content of organo‐mineral associations, and (5) the factors controlling the turnover of organic matter (OM) in organo‐mineral associations from temperate soils. Physical fractionation achieves a rough separation between plant residues and mineral‐associated OM, which makes density or particle‐size fractionation a useful pretreatment for further differentiation of functional fractions. A part of the OM in organo‐mineral associations resists different chemical treatments, but the data obtained cannot readily be compared among each other, and more research is necessary on the processes underlying resistance to treatments for certain OM components. Studies using physical‐fractionation procedures followed by soil‐microbiological analyses revealed that organo‐mineral associations spatially isolate C sources from soil biota, making quantity and quality of OM in microhabitats an important factor controlling community composition. The distribution and activity of soil microorganisms at organo‐mineral surfaces can additionally be modified by faunal activities. Composition of OM in organo‐mineral associations is highly variable, with loamy soils having generally a higher contribution of polysaccharides, whereas mineral‐associated OM in sandy soils is often more aliphatic. Though highly reactive towards Fe oxide surfaces, lignin and phenolic components are usually depleted in organo‐mineral associations. Charred OM associated with the mineral surface contributes to a higher aromaticity in heavy fractions. The relative proportion of OC bound in organo‐mineral fractions increases with soil depth. Likewise does the strength of the bonding. Organic molecules sorbed to the mineral surfaces or precipitated by Al are effectively stabilized, indicated by reduced susceptibility towards oxidative attack, higher thermal stability, and lower bioavailability. At higher surface loading, organic C is much better bioavailable, also indicated by little 14C age. In the subsurface horizons of the soils investigated in this study, Fe oxides seem to be the most important sorbents, whereas phyllosilicate surfaces may be comparatively more important in topsoils. Specific surface area of soil minerals is not always a good predictor for C‐stabilization potentials because surface coverage is discontinuous. Recalcitrance and accessibility/aggregation seem to determine the turnover dynamics in fast and intermediate cycling OM pools, but for long‐term OC preservation the interactions with mineral surfaces, and especially with Fe oxide surfaces, are a major control in all soils investigated here.

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.

Soil organisms are a crucial part of the terrestrial biosphere. Despite their importance for ecosystem functioning, no quantitative, spatially-explicit models of the active belowground community currently exist. In particular, nematodes are the most abundant animals on Earth, filling all trophic levels in the soil food web. Here, we use 6,579 georeferenced samples to generate a mechanistic understanding of the patterns of global soil nematode abundance and functional group composition. The resulting maps show that 4.4 ± 0.64  10 20 nematodes (total biomass ~0.3 Gt) inhabit surface soils across the world, with higher abundances in sub-arctic regions (38% of total), than in temperate (24%), or tropical regions (21%). Regional variations in these global trends also provide insights into local patterns of soil fertility and functioning. These high-resolution models provide the first steps towards representing soil ecological processes into global biogeochemical models, to predict elemental cycling under current and future climate scenarios.

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...