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Flooding threatens the functioning of managed grasslands by decreasing primary productivity and increasing nitrogen losses, notably as the potent greenhouse gas nitrous oxide (N2O). Sowing species with traits that promote flood resilience and mitigate flood-induced N2O emissions within these grasslands could safeguard their productivity while mitigating nitrogen losses. We tested how plant traits and resource acquisition strategies could predict flood resilience and N2O emissions of 12 common grassland species (eight grasses and four legumes) grown in field soil in monocultures in a 14-week greenhouse experiment. We found that grasses were more resistant to flooding while legumes recovered better. Resource-conservative grass species had higher resistance while resource-acquisitive grasses species recovered better. Resilient grass and legume species lowered cumulative N2O emissions. Grasses with lower inherent leaf and root δ13C (and legumes with lower root δ13C) lowered cumulative N2O emissions during and after the flood. Our results highlight the differing responses of grasses with contrasting resource acquisition strategies, and of legumes to flooding. Combining grasses and legumes based on their traits and resource acquisition strategies could increase the flood resilience of managed grasslands, and their capability to mitigate flood-induced N2O emissions. A free Plain Language Summary can be found within the Supporting Information of this article.

Plant diversity is an important driver of below-ground ecosystem functions, such as root growth, soil organic matter (SOM) storage and microbial metabolism, mainly by influencing the interactions between plant roots and soil. Dissolved organic matter (DOM), as the most mobile form of SOM, plays a crucial role for a multitude of soil processes that are central for ecosystem functioning. Thus, DOM is likely to be an important mediator of plant diversity effects on soil processes. However, the relationships between plant diversity and DOM have not been studied so far. We investigated the mechanisms underlying plant diversity effects on concentrations of DOM using continuous soil water sampling across 6 years and 62 plant communities in a long-term grassland biodiversity experiment in Jena, Germany. Furthermore, we investigated plant diversity effects on the molecular properties of DOM in a subset of the samples. Although DOM concentrations were highly variable over the course of the year with highest concentrations in summer and autumn, we found that DOM concentrations consistently increased with plant diversity across seasons. The positive plant diversity effect on DOM concentrations was mainly mediated by increased microbial activity and newly sequestered carbon in topsoil. However, the effect of soil microbial activity on DOM concentrations differed between seasons, indicating DOM consumption in winter and spring, and DOM production in summer and autumn. Furthermore, we found increased contents of small and easily decomposable DOM molecules reaching deeper soil layers with high plant diversity. Synthesis. Our findings suggest that plant diversity enhances the continuous downward transport of DOM in multiple ways. On the one hand, higher plant diversity results in higher DOM concentrations, on the other hand, this DOM is less degraded. This study indicates, for the first time, that higher plant diversity enhances the downward transport of dissolved molecules that lik

Minimizing nitrogen (N) losses and increasing plant N uptake in agroecosystems is a major global challenge. Ecological concepts from (semi)natural grasslands suggest that manipulating plant community composition using plant species with different traits may represent a promising opportunity to face this challenge. Here, we translate these trait-based concepts to agricultural systems in a field experiment, aiming to reveal the main determinants of how plant community composition regulates N-cycling in intensively managed grasslands. We focused on key N pools (plant N from soil and from biological N-fixation, soil mineral N and N2O emissions) as well as on biological drivers of N-cycling in soil (abundance of N-cycling microbial communities, earthworm populations and arbuscular mycorrhizal fungi), using three common grass and one legume species in monoculture, two- and four-species mixtures. We hypothesized that: (a) plant species mixtures increase plant N uptake, reduce soil mineral N concentrations and N2O emissions and promote the abundance of biological N-cyclers; (b) legume presence stimulates N pools, fluxes and biological N-cycling activity, (c) but in combination with a grass with acquisitive traits, more N is retained in the plant community, while N2O emissions are reduced. We found that mixtures increased plant N and lowered the soil mineral N pool compared to monocultures. However, plant species identity played an overarching role: Legume presence increased N2O emissions, plant N pools, soil mineral N and the abundance of N-cycling microbes and earthworms. Combining the legume with a grass with low leaf dry matter content and high root length density (and with high root biomass) reduced the higher soil mineral N and N2O emissions induced by the legume, while harnessing positive effects on plant N pools and biological N-fixation. Synthesis and applications. Our results show the potential of plant community composition to steer N-cycling in fertilized agroeco

Plant diversity is an important driver of below-ground ecosystem functions, such as root growth, soil organic matter (SOM) storage and microbial metabolism, mainly by influencing the interactions between plant roots and soil. Dissolved organic matter (DOM), as the most mobile form of SOM, plays a crucial role for a multitude of soil processes that are central for ecosystem functioning. Thus, DOM is likely to be an important mediator of plant diversity effects on soil processes. However, the relationships between plant diversity and DOM have not been studied so far. We investigated the mechanisms underlying plant diversity effects on concentrations of DOM using continuous soil water sampling across 6 years and 62 plant communities in a long-term grassland biodiversity experiment in Jena, Germany. Furthermore, we investigated plant diversity effects on the molecular properties of DOM in a subset of the samples. Although DOM concentrations were highly variable over the course of the year with highest concentrations in summer and autumn, we found that DOM concentrations consistently increased with plant diversity across seasons. The positive plant diversity effect on DOM concentrations was mainly mediated by increased microbial activity and newly sequestered carbon in topsoil. However, the effect of soil microbial activity on DOM concentrations differed between seasons, indicating DOM consumption in winter and spring, and DOM production in summer and autumn. Furthermore, we found increased contents of small and easily decomposable DOM molecules reaching deeper soil layers with high plant diversity. Synthesis. Our findings suggest that plant diversity enhances the continuous downward transport of DOM in multiple ways. On the one hand, higher plant diversity results in higher DOM concentrations, on the other hand, this DOM is less degraded. This study indicates, for the first time, that higher plant diversity enhances the downward transport of dissolved molecules that lik

The increasing frequency of extreme weather events, such as floods, requires management strategies that promote resilience of grassland productivity. Mixtures of plant species may better resist and recover from flooding than monocultures, as they could combine species with stress‐coping and resource acquisition traits. This has not yet been tested in intensively managed grasslands despite its relevance for enhancing agroecosystem resilience. Using intact soil cores from an 18‐month‐old field experiment, we tested how 11 plant communities (Festuca arundinacea, Lolium perenne, Poa trivialis and Trifolium repens in monoculture, two‐ and four‐species mixtures) resist and recover from repeated flooding in a 4‐month greenhouse experiment. We found that plant community composition, not whether the community was a mixture or monoculture, influenced the community's resistance to flooding, although most communities were able to resist and recover from both floods. The plant community's position on the leaf economic spectrum in flooded conditions was related to its resistance to and recovery from flooding. Resistance to and recovery from a severe flood were related to flood‐induced intraspecific trait variation, causing a shift in the community's position on the leaf resource economic spectrum. In flooded conditions, resource‐conservative communities (characterized by low specific leaf area, low leaf nitrogen content and high leaf dry matter content) better resisted and recovered from flooding. The community's position on the root resource economic spectrum was less connected to the community's resistance and recovery. Synthesis and applications. Our study shows that in flooded conditions, resource‐conservative plant communities are more resilient to flooding than resource‐acquisitive communities in an intensively managed grassland. This suggests that plant community position on the leaf economic spectrum, as well as species’ flood‐induced intraspecific var

1. Plant diversity is an important driver of belowground ecosystem functions, such as root growth, soil organic matter (SOM) storage, and microbial metabolism, mainly by influencing the interactions between plant roots and soil. Dissolved organic matter (DOM), as the most mobile form of SOM, plays a crucial role for a multitude of soil processes that are central for ecosystem functioning. Thus, DOM is likely to be an important mediator of plant diversity effects on soil processes. However, the relationships between plant diversity and DOM have not been studied so far. 2. We investigated the mechanisms underlying plant diversity effects on concentrations of DOM using continuous soil water sampling across 6 years and 62 plant communities in a long‐term grassland biodiversity experiment in Jena, Germany. Furthermore, we investigated plant diversity effects on the molecular properties of DOM in a subset of the samples. 3. Although DOM concentrations were highly variable over the course of the year with highest concentrations in summer and autumn, we found that DOM concentrations consistently increased with plant diversity across seasons. The positive plant diversity effect on DOM concentrations was mainly mediated by increased microbial activity and newly sequestered carbon in topsoil. However, the effect of soil microbial activity on DOM concentrations differed between seasons, indicating DOM consumption in winter and spring, and DOM production in summer and autumn. Furthermore, we found increased contents of small and easily decomposable DOM molecules reaching deeper soil layers with high plant diversity. 4. Synthesis. Our findings suggest that plant diversity enhances the continuous downward transport of DOM in multiple ways. On the one hand, higher plant diversity results in higher DOM concentrations, on the other hand, this DOM is less degraded. The present study indicates, for the first time, that higher plant diversity enhances the downward transport of dissolved

Intensively managed grasslands are large sources of the potent greenhouse gas nitrous oxide (N2O) and important regulators of methane (CH4) consumption and production. The predicted increase in flooding frequency and severity due to climate change could increase N2O emissions and shift grasslands from a net CH4 sink to a source. Therefore, effective management strategies are critical for mitigating greenhouse gas emissions from flood-prone grasslands. We tested how repeated flooding affected the N2O and CH4 emissions from 11 different plant communities (Festuca arundinacea, Lolium perenne, Poa trivialis, and Trifolium repens in monoculture, 2- and 4-species mixtures), using intact soil cores from an 18-month old grassland field experiment in a 4-month greenhouse experiment. To elucidate potential underlying mechanisms, we related plant functional traits to cumulative N2O and CH4 emissions. We hypothesized that traits related with fast nitrogen uptake and growth would lower N2O and CH4 emissions in ambient (non-flooded) conditions, and that traits related to tissue toughness would lower N2O and CH4 emissions in flooded conditions. We found that flooding increased cumulative N2O emissions by 97 fold and cumulative CH4 emissions by 1.6 fold on average. Plant community composition mediated the flood-induced increase in N2O emissions. In flooded conditions, increasing abundance of the grass F. arundinacea was related with lower N2O emissions; whereas increases in abundance of the legume T. repens resulted in higher N2O emissions. In non-flooded conditions, N2O emissions were not clearly mediated by plant traits related with nitrogen uptake or biomass production. In flooded conditions, plant communities with high root carbon to nitrogen ratio were related with lower cumulative N2O emissions, and a lower global warming potential (CO2 equivalent of N2O and CH4). We conclude that plant functional traits related to slower decomposition and nitrogen mineralization could play a

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Dissolved organic matter affects fundamental biogeochemical processes in the soil such as nutrient cycling and organic matter storage. The current paradigm is that processing of dissolved organic matter converges to recalcitrant molecules (those that resist degradation) of low molecular mass and high molecular diversity through biotic and abiotic processes. Here we demonstrate that the molecular composition and properties of dissolved organic matter continuously change during soil passage and propose that this reflects a continual shifting of its sources. Using ultrahigh-resolution mass spectrometry and nuclear magnetic resonance spectroscopy, we studied the molecular changes of dissolved organic matter from the soil surface to 60 cm depth in 20 temperate grassland communities in soil type Eutric Fluvisol. Applying a semi-quantitative approach, we observed that plant-derived molecules were first broken down into molecules containing a large proportion of low-molecular-mass compounds. These low-molecular-mass compounds became less abundant during soil passage, whereas larger molecules, depleted in plant-related ligno-cellulosic structures, became more abundant. These findings indicate that the small plant-derived molecules were preferentially consumed by microorganisms and transformed into larger microbial-derived molecules. This suggests that dissolved organic matter is not intrinsically recalcitrant but instead persists in soil as a result of simultaneous consumption, transformation and formation.

Background Plant diversity influences ecosystem functioning. A positive relation between plant diversity and productivity above- and belowground has been established. Aboveground, this effect has been shown to be due to complementarity effects, interactions between species in a mixture that lead species to, on average, produce more biomass than expected based on their productivity in monoculture. The mechanisms underlying complementarity effects and the positive diversity-productivity relation are predicted to lie belowground, e.g. resource partitioning and/or facilitation. The relation between plant diversity and decomposition is less clear, and research on the diversity-decomposition relation belowground is limited. This is an important gap in biodiversity knowledge, as the decomposition of plant litter is the major source of nutrients and carbon in terrestrial ecosystems, and most plant litter in grasslands is belowground. Methods This thesis explored the effect of plant diversity on belowground productivity and decomposition. Belowground complementarity effects were quantified in the Jena Trait Based Experiment, and the diversity of or plasticity in species-specific vertical root distribution as underlying mechanism was tested. The plant diversity- root decomposition relation was quantified in the Jena Experiment and the Jena Trait Based Experiment. The role of root traits and the soil environment as mediating factors were tested. Major findings Plant diversity had a positive effect on root biomass production, and this relation was attributed to complementarity effects. The diversity in species-specific vertical root distribution did not explain complementarity effects, and thus, is not likely a major mechanism underlying the diversity-productivity relation. Species altered their vertical root distribution in response to inter-specific neighbours. The direction of this change differed between functional groups: grasses became shallower in mixture, forbs became d