Nutrient limitations induced by drought affect forage N and P differently in two permanent grasslands

Drought events can strongly affect ecosystem functioning by modifying relationship between plants, microbes and soil chemistry, with consequent impacts on nutrient cycling. However, the potential impacts of a soil moisture reduction on the nitrogen (N) and phosphorus (P) cycling in grasslands remain poorly understood, especially in regard to. forage production. To fill this knowledge gap, a drought experiment was carried out using rainout shelters in two permanent grasslands, characterized by similar vegetation communities but contrasted soil nutrient limitations. Drought treatments were applied during two months, either when plant growth was highest (Early-season drought) or after the peak of biomass production (Late-season drought). Dry matter production, forage N status (NNI) and P content as well as N and P contents in microbial biomass and soil were determined. Both early and late-season drought significantly reduced soil moisture during the vegetation growth period. Forage yield was also reduced by drought, but only when it occurred late in the season. Using a structural equation model, we showed that soil moisture reduction had a direct effect on forage N status, suggesting that water shortage induced lower transpiration and water fluxes. Soil moisture reduction also affected forage P by reducing the availability of soil P. However, other mechanisms played a larger role and were site-specific. At the more fertile site, reduction in soil moisture directly impaired forage P, suggesting that water stress mainly resulted in lower diffusion rates to roots, while at the less fertile site, an indirect reduction of forage P through a pathway implying microbes (decrease in microbial P) was detected. Our results suggest that the two grasslands suffered mainly from water shortage per se, but also from drought induced nutrient deficiency (mainly P), which amplified yield losses and further decreased forage quality. Overall, our findings emphasize the need for further research on the plant-soil-microbe system functioning, in order to secure a sustainable and resilient forage production in the context of climate change.

Effects of Subordinate Plant Species in Plant and Soil Community Structure and Ecosystem Functioning

Pierre Rémi Mariotte

The relevance of biodiversity to human health is an increasing international political issue as it causes concern for ethical and aesthetic reasons, but also has a strong impact on ecosystem properties and ecological goods and services utilised by humanity. However, humans have applied increasing pressure on worldwide biodiversity through pollution, land-use and climate change, over exploitation and the introduction of invasive species which can lead to major alterations of biological communities and a consequent decrease in biodiversity. When communities are assembled at random from a pool of species, more diverse mixtures have a higher probability to contain species or species-groups with high capacity to drive ecosystem processes. Therefore, it is relevant to ask "how may we classify species in a community in terms of functionality?", and "which species-group is important to maintain the productivity and stability of ecosystems?" In this thesis, I chose to classify species according their frequency and their cumulative relative cover in a plant community (i.e. plant hierarchy) and to differentiate three species-groups: dominant, subordinate and transient, which contribute towards biodiversity. Dominant species are clearly very important for ecosystem functioning, due to the large amount of biomass they produce, but there is growing evidence that subordinate species, which represent a low amount of plant biomass in grassland ecosystems, are also of considerable importance for ecosystem functioning. Given this recognition, a further objective was to explore the ecosystem level effects of these subordinate plant species, which is currently not well known. This research focused on two aspects: the persistence of subordinate species in the community, and their role in grassland ecosystems. The approach to satisfy these objectives encompassed four years of experimental field studies and glasshouse microcosm experiments. In order to explain the persistence of subordinate species in semi-natural grasslands, we performed two greenhouse experiments to test the effects of root competition exclusion and the inoculation of arbuscular mycorrhizal fungi (AMF) on the competitiveness of dominant and subordinate species, directly measured by their biomass production. The effects of subordinate species on ecosystem functioning were assessed through long-term field experiment in two sites in the Swiss Jura Mountains, where dominant, subordinate and transient species were preliminarily determined. At both site (Les Amburnex, La Frétaz), a removal experiment was carried out in randomly replicated plots with three different treatments: control without perturbations, removal of all subordinate species and partial removal of dominant biomass. In one site, a summer drought treatment was added using rainout shelters to simulate an extreme climatic event. Biomass production, litter decomposition and soil respiration were monitored during four years in each plot and complemented in some cases by measurements of soil dissolved inorganic nitrogen concentrations, soil microbial communities (bacteria and AMF) and carbon and nitrogen isotopes in plant leaves. The results of the different experiments conducted in this thesis highlight the importance of cattle activity and AMF on the persistence of subordinate plant species in semi-natural grasslands. Indeed, cattle activities (i.e. trampling) provide spatial heterogeneity, through gap creation (areas of root competition reduction), and thus favours the growth of less competitive subordinate species. While AMF acted as a parasite in the greenhouse experiment, the current findings suggest that the competitiveness of dominant species was reduced through the action of the fungus, which indirectly enhanced subordinate species. The interactions between subordinate species and soil organisms were confirmed in the field as bacterial and AMF communities shifted in plots where subordinate species were removed compared to the control treatments. Moreover, the absence of subordinate species had many negative effects on ecosystem functioning by reducing litter decomposition, soil respiration, nitrogen mineralization and community above-ground production. These findings suggest that plant-soil feedbacks explain the importance of subordinate species on ecosystem functioning despite only representing a very low quantity of biomass in plant communities. During the summer drought, subordinate species increased the resistance of the plant community and maintained productivity. Therefore, these outcomes confirm the role of drought-resistant subordinates in the functioning of grassland ecosystems threatened by climate change. The synthesis of this thesis has important implications for the way grasslands should be managed for biodiversity and confirms that extensive grazing must be maintained as it promotes species coexistence and the persistence of key subordinate species. Moreover, this work demonstrates the important role of subordinate species in ecosystem functioning, showing that not only dominant species determine ecosystem properties. This thesis serves as a contribution to the advances in our understanding of ecosystem functioning and above and below-ground linkages, and provides basis for future research in this domain.

EPFL2012

Land management trumps the effects of climate change and elevated CO2 on grassland functioning

Pierre Rémi Mariotte, Aurélie Thébault

Grasslands cover similar to 30% of the Earth's terrestrial surface and provide many ecosystem services. Many grasslands are heavily managed to maximize these services for human benefit, but the outcome of management is anticipated to be increasingly influenced by various aspects of climate change and elevated atmospheric CO2. The relative importance of global change vs. land management on grasslands is largely unknown. 2. A meta-analysis is used here to examine drivers at both scales primarily targeting services provided by grasslands relating to plant productivity (above- and below-ground biomass) and soil processes (nutrients and soil respiration) in 38 manipulative experiments published in the last decade. We specifically target effects of (i) single and combined land management practices (LMs), (ii) single and combined factors relating to broad-scale climate change and elevated CO2, and (iii) combined management practices and changes to climate and CO2. Collectively, this examines the general efficacy of global change models in predicting changes to grassland functioning. 3. We found that combinations of management practices had approximately double the explanatory power for variation in grassland services compared with individual or interactive effects of factors associated with climate change and CO2. These interacting management practices such as nutrient additions and defoliation predominantly influenced functions associated with productivity or biomass both below and above ground. The effects of interacting factors of climate and CO2 influenced a wider range of ecosystem functions, but the magnitude of these effects was relatively smaller. 4. Interactions between management practices or between climate change/CO2 factors always had higher explanatory power than any factor in isolation indicating that multivariate synergistic models of environmental change can better describe impacts on ecosystem function in plant communities (e.g. relative to univariate climate-based models). Given that the magnitude and direction (positive or negative) of the interactions varied widely, this also implies that the outcomes of these multivariate interactions can vary spatially, temporally or by immediate context (e.g. management prescriptions). 5. Synthesis. Although our work confirms how climate change and CO2 can affect many ecosystem-based functional attributes, it suggests that combinations of LMs remain the dominant set of factors in determining the performance of grassland plant communities. Land management may thus be critical for influencing projected responses to future climate change and elevated CO2 in models of grassland function at least for factors relating to primary production.

Wiley-Blackwell2014

Effects of Subordinate Plant Species in Plant and Soil Community Structure and Ecosystem Functioning

Pierre Rémi Mariotte

EPFL2012

Land management trumps the effects of climate change and elevated CO2 on grassland functioning

Pierre Rémi Mariotte, Aurélie Thébault

Grasslands cover similar to 30% of the Earth's terrestrial surface and provide many ecosystem services. Many grasslands are heavily managed to maximize these services for human benefit, but the outcome of management is anticipated to be increasingly influenced by various aspects of climate change and elevated atmospheric CO2. The relative importance of global change vs. land management on grasslands is largely unknown. 2. A meta-analysis is used here to examine drivers at both scales primarily targeting services provided by grasslands relating to plant productivity (above- and below-ground biomass) and soil processes (nutrients and soil respiration) in 38 manipulative experiments published in the last decade. We specifically target effects of (i) single and combined land management practices (LMs), (ii) single and combined factors relating to broad-scale climate change and elevated CO2, and (iii) combined management practices and changes to climate and CO2. Collectively, this examines the general efficacy of global change models in predicting changes to grassland functioning. 3. We found that combinations of management practices had approximately double the explanatory power for variation in grassland services compared with individual or interactive effects of factors associated with climate change and CO2. These interacting management practices such as nutrient additions and defoliation predominantly influenced functions associated with productivity or biomass both below and above ground. The effects of interacting factors of climate and CO2 influenced a wider range of ecosystem functions, but the magnitude of these effects was relatively smaller. 4. Interactions between management practices or between climate change/CO2 factors always had higher explanatory power than any factor in isolation indicating that multivariate synergistic models of environmental change can better describe impacts on ecosystem function in plant communities (e.g. relative to univariate climate-based models). Given that the magnitude and direction (positive or negative) of the interactions varied widely, this also implies that the outcomes of these multivariate interactions can vary spatially, temporally or by immediate context (e.g. management prescriptions). 5. Synthesis. Although our work confirms how climate change and CO2 can affect many ecosystem-based functional attributes, it suggests that combinations of LMs remain the dominant set of factors in determining the performance of grassland plant communities. Land management may thus be critical for influencing projected responses to future climate change and elevated CO2 in models of grassland function at least for factors relating to primary production.

Wiley-Blackwell2014