Frédérique Louault

Plant traits and functional types in response to reduced disturbance in a semi-natural grassland

Abstract Question: How do functional types respond to contrasting levels of herbage use in temperate and fertile grasslands? Location: Central France (3°1′ E, 45°43′ N), 870 m a.s.l. Methods: Community structure and the traits of dominant plant species were evaluated after 12 years of contrasted grazing and mowing regimes in a grazing trial, comparing three levels of herbage use (high, medium and low). Results and Conclusions: Of 22 measured traits (including leaf traits, shoot morphology and composition, phenology), seven were significantly affected by the herbage use treatment. A decline in herbage use reduced individual leaf mass, specific leaf area and shoot digestibility, but increased leaf C and dry matter contents. Plants were taller, produced larger seeds and flowered later under low than high herbage use. Nine plant functional response types were identified by multivariate optimization analysis; they were based on four optimal traits: leaf dry matter content, individual leaf area, mature plant height and time of flowering. In the high-use plots, two short and early flowering types were co-dominant, one competitive, grazing-tolerant and moderately grazing-avoiding, and one grazing-avoiding but not -tolerant. Low-use plots were dominated by one type, neither hardly grazing-avoiding nor grazing-tolerant, but strongly competitive for light. Nomenclature: Tutin et al. (1993). Abbreviations: BE = Beginning of flowering period; DI = Digestibility; IT = Height at top of inflorescence; LA = Individual leaf area; LCC = Leaf carbon concentration; LDM = Leaf dry mass; LDMC = Leaf dry matter content; LFM = Leaf lamina fresh mass; LNC = Leaf nitrogen concentration; ME = Flowering plant height, highest leaf elongated; MH = Flowering plant height, highest leaf not elongated; NG = Number of growing green leaves; NM = Number of mature green leaves; PRT = Plant functional response type; RA = Leaf:Shoot dry matter ratio; SLAF = Specific leaf area (fresh mass based); SLAD = Specific leaf area (dry mass based); SM = Seed mass.

EFFECTS OF GRAZING ON MICROBIAL FUNCTIONAL GROUPS INVOLVED IN SOIL N DYNAMICS

Enhancement of soil nitrogen (N) cycling by grazing has been observed in many grassland ecosystems. However, whether grazing affects the activity only of the key microbial functional groups driving soil N dynamics or also affects the size (cell number) and/or composition of these groups remains largely unknown. We studied the enzyme activity, size, and composition of five soil microbial communities (total microbial and total bacterial communities, and three functional groups driving N dynamics: nitrifiers, denitrifiers, and free N2 fixers) in grassland sites experiencing contrasting sheep grazing regimes (one light grazing [LG] site and one intensive grazing [IG] site) at two topographical locations. Enzyme activity was determined by potential carbon mineralization, nitrification, denitrification, and N2 fixation assays. The size of each community (except N2 fixers) was measured by the most-probable-number technique. The composition of the total soil microbial community was characterized by phospholipid f...

Habitat filtering and niche differentiation jointly explain species relative abundance within grassland communities along fertility and disturbance gradients

Deterministic niche-based processes have been proposed to explain species relative abundance within communities but lead to different predictions: habitat filtering (HF) predicts dominant species to exhibit similar traits while niche differentiation (ND) requires that species have dissimilar traits to coexist. Using a multiple trait-based approach, we evaluated the relative roles of HF and ND in determining species abundances in productive grasslands. Four dimensions of the functional niche of 12 co-occurring grass species were identified using 28 plant functional traits. Using this description of the species niche, we investigated patterns of functional similarity and dissimilarity and linked them to abundance in randomly assembled six-species communities subjected to fertilization/disturbance treatments. Our results suggest that HF and ND jointly determined species abundance by acting on contrasting niche dimensions. The effect of HF decreased relative to ND with increasing disturbance and decreasing fertilization. Dominant species exhibited similar traits in communities whereas dissimilarity favored the coexistence of rare species with dominants by decreasing inter-specific competition. This stabilizing effect on diversity was suggested by a negative relationship between species over-yielding and relative abundance. We discuss the importance of considering independent dimensions of functional niche to better understand species abundance and coexistence within communities.

Stimulation of soil nitrification and denitrification by grazing in grasslands: do changes in plant species composition matter?

Stimulation of nitrification and denitrification by long term (from years to decades) grazing has commonly been reported in different grassland ecosystems. However, grazing generally induces important changes in plant species composition, and whether changes in nitrification and denitrification are primarily due to changes in vegetation composition has never been tested. We compared soil nitrification- and denitrification-enzyme activities (NEA and DEA, respectively) between semi-natural grassland sites experiencing intensive (IG) and light (LG) grazing/mowing regimes for 13 years. Mean NEA and DEA (i.e. observed from random soil sampling) were higher in IG than LG sites. The NEA/DEA ratio was higher in IG than LG sites, indicating a higher stimulation of nitrification. Marked changes in plant species composition were observed in response to the grazing/mowing regime. In particular, the specific phytomass volume of Elymus repens was lower in IG than LG sites, whereas the specific volume of Lolium perenne was higher in IG than LG sites. In contrast, the specific volume of Holcus lanatus, Poa trivialis and Arrhenatherum elatius were not significantly different between treatments. Soils sampled beneath grass tussocks of the last three species exhibited higher DEA, NEA and NEA/DEA ratio in IG than LG sites. For a given grazing regime, plant species did not affect significantly soil DEA, NEA and NEA/DEA ratio. The modification of plant species composition is thus not the primary factor driving changes in nitrification and denitrification in semi-natural grassland ecosystems experiencing long term intensive grazing. Factors such as trampling, N returned in animal excreta, and/or modification of N uptake and C exudation by frequently defoliated plants could be responsible for the enhanced microbial activities.

Priming effect and microbial diversity in ecosystem functioning and response to global change: a modeling approach using the SYMPHONY model.

Integration of the priming effect (PE) in ecosystem models is crucial to better predict the consequences of global change on ecosystem carbon (C) dynamics and its feedbacks on climate. Over the last decade, many attempts have been made to model PE in soil. However, PE has not yet been incorporated into any ecosystem models. Here, we build plant/soil models to explore how PE and microbial diversity influence soil/plant interactions and ecosystem C and nitrogen (N) dynamics in response to global change (elevated CO2 and atmospheric N depositions). Our results show that plant persistence, soil organic matter (SOM) accumulation, and low N leaching in undisturbed ecosystems relies on a fine adjustment of microbial N mineralization to plant N uptake. This adjustment can be modeled in the SYMPHONY model by considering the destruction of SOM through PE, and the interactions between two microbial functional groups: SOM decomposers and SOM builders. After estimation of parameters, SYMPHONY provided realistic predictions on forage production, soil C storage and N leaching for a permanent grassland. Consistent with recent observations, SYMPHONY predicted a CO2 -induced modification of soil microbial communities leading to an intensification of SOM mineralization and a decrease in the soil C stock. SYMPHONY also indicated that atmospheric N deposition may promote SOM accumulation via changes in the structure and metabolic activities of microbial communities. Collectively, these results suggest that the PE and functional role of microbial diversity may be incorporated in ecosystem models with a few additional parameters, improving accuracy of predictions.

Plant trait–digestibility relationships across management and climate gradients in permanent grasslands

1. Dry matter digestibility is a critical component of herbage nutritive value, a major service delivered by grasslands. The aim of this study was to test whether the dominance hypothesis applies to assess the impacts of environmental gradients and management regimes on thiscomponent of herbage nutritive value in permanent grasslands. 2. At the plant level, digestibility has been related to a number of functional traits, but whether this can be scaled up to the community level in species-rich grasslands and how such relationships are modulated by environmental conditions and management regimes remainunknown. Our primary objective was to test whether community-weighted means – species trait values weighted by the species abundance – of morphological, phenological and chemical traits could be used to explain variations in digestibility over a large range of climatic contexts,soil resource levels and management regimes. Our second objective was to explain variations in community digestibility within and among nine contrasting sites along large natural and man-induced environmental gradients.3. Over the whole data set, digestibility and most community-weighted means of traits responded to climatic factors and management regimes, but relations were not always significant when each site was considered separately. Community digestibility was significantly related to one or more plant traits within each site and to all of the measured traits when considering all the sites. Leaf dry matter content (LDMC) had the most consistent effects on digestibility, with a strikingly similar negative effect within each site. Potential evapotranspiration was negatively related to digestibility and contributed to explain a large part of the among-site variance. In addition, a low return interval of disturbance and a high disturbance intensity (biomass removal) were both associated with a high digestibility.4. Synthesis and applications. Disturbance regime, plant traits and local climate impacted dry matter digestibility roughly equally in grasslands. The effects of community composition on digestibility and its response to abiotic factors could be successfully captured by community weightedmeans of leaf dry matter content. This functional marker can be used to develop indicators and grassland management rules to support farmers in the refinement of their practices towards specific needs, such as target production outputs.

Grass strategies and grassland community responses to environmental drivers: a review

Grassland covers about one quarter of the Earth’s land area and is currently estimated to contribute to the livelihoods of over 800 million people. Grassland provides ecosystem goods and services, mainly through the provisioning of milk and meat. Therefore, the proper use of grasslands will be essential for feeding the nine billion people that will inhabit planet Earth by 2050. In the context of a changing climate, we should better understand the interactions of environment, management and grass crop at individual, community and ecosystem levels. Functional ecology focuses on the roles and functions that species play in the community or ecosystem in which they occur. Functional ecology thus aims to understand how plant species adapt to environmental conditions and how management can alter this adaptation. Here, we review the latest advances in plant functional traits research and on species strategies to the main environmental factors occurring in grassland ecosystems: nutrient availability, grazing, cutting and shading. Functional ecology also provides a framework to better understand how species strategies interact with the species composition at the community level. Therefore, the literature on community assembling theories in relation to ecosystem processes most relevant to grassland management and services is also reviewed. Finally, future research questions and some new orientations for grassland experts are offered in order to meet the challenge of maintaining productivity and preservation of these semi-natural environments in the face of global change.

Recasting the dynamic equilibrium model through a functional lens: the interplay of trait‐based community assembly and climate

1. According to the dynamic equilibrium hypothesis (DEH), plant species richness is locally controlled by productivity and disturbance. Given that regional conditions widely affect local environmental variables such as soil nutrient availability, the DEH predictions could be improved by considering how climate influences local controls of species richness. Further, a trait-based approach to community assembly has the potential to reveal a deeper, mechanistic understanding of species richness variation across environments. Here, we bring together DEH and trait-based community assembly expectations to examine whether and how local relationships between diversity, disturbance and productivity are affected by habitat filtering and regional climate. 2. We specifically tested how gradients of local nutrient availability and disturbance intensity interact with climatic conditions to drive the species richness of grassland communities. Further, we recast the DEH through a functional lens by exploring how disturbance–diversity and nutrient availability–diversity relationships are shaped by the functional space occupied by species in a community and species packing within this functional space. 3. The functional space occupied by co-occurring species and the way they are functionally packed are quantified using multi-trait indices calculated with five core plant functional traits. Working with grassland communities spread across differing regional climatic conditions, we used mixed models to test whether the variation in taxonomic and functional metrics corresponded to the DEH predictions as well as to determine the relationship between those metrics. 4. Contrary to the expectations based on the relation between species richness and the functional components considered, taxonomic and functional metrics did not vary in accordance along environmental gradients. Climate strongly interacted with the local environment to modulate local diversity patterns, sometimes even inversing a given trend and falsifying the DEH predictions. 5. Synthesis. Our findings quantitatively highlight the interplay between regional and local environmental gradients in driving community assembly. We demonstrate that, depending on climatic conditions, observed patterns of both taxonomic and functional community composition can be opposite to expected productivity–diversity and disturbance–diversity relationships. This emphasizes the relevance of multifaceted studies of biodiversity and the need for a more systematic quantification of regional controls in community assembly studies.

Habitat filtering determines the functional niche occupancy of plant communities worldwide

How the patterns of niche occupancy vary from species-poor to species-rich communities is a fundamental question in ecology that has a central bearing on the processes that drive patterns of biodiversity. As species richness increases, habitat filtering should constrain the expansion of total niche volume, while limiting similarity should restrict the degree of niche overlap between species. Here, by explicitly incorporating intraspecific trait variability, we investigate the relationship between functional niche occupancy and species richness at the global scale. 2.We assembled 21 datasets worldwide, spanning tropical to temperate biomes and consisting of 313 plant communities representing different growth forms. We quantified three key niche occupancy components (the total functional volume, the functional overlap between species and the average functional volume per species) for each community, related each component to species richness, and compared each component to the null expectations. 3.As species richness increased, communities were more functionally diverse (an increase in total functional volume), and species overlapped more within the community (an increase in functional overlap) but did not more finely divide the functional space (no decline in average functional volume). Null model analyses provided evidence for habitat filtering (smaller total functional volume than expectation), but not for limiting similarity (larger functional overlap and larger average functional volume than expectation) as a process driving the pattern of functional niche occupancy. 4.Synthesis. Habitat filtering is a widespread process driving the pattern of functional niche occupancy across plant communities and coexisting species tend to be more functionally similar rather than more functionally specialized. Our results indicate that including intraspecific trait variability will contribute to a better understanding of the processes driving patterns of functional niche occupancy

Intervalo e intensidade de desfolhação nas taxas de crescimento, senescência e desfolhação e no equilíbrio de gramíneas em associação

The experiment was carried in INRA (Institut National de la Recherche Agronomique) at Theix, France. Two grasses (perennial ryegrass and tall fescue) were grown in sward boxes (0,13 m2) and submitted to three defoliation intervals (3,5, 7 and 14 days between two successive defoliation) and three defoliation intensities (160, 320 and 640 bites.m-2 at each defoliation). Four months after sowing, at defoliation date, sward boxes were offered to four individual dry ewes and removed after a given number of bites had been taken. For each grass species, the growth, senescence and defoliation fluxes were studied. The results showed that the interval and the intensity of grazing had distinct effects on the growth, senescence and defoliation fluxes as well as on the equilibrium of grass mixtures.