Reindeer can have strong effects on the plant species composition and functioning of tundra ecosystems, and often promote a transition towards a graminoid-dominated vegetation type. As a result, they influence many ecological processes, such as nutrient dynamics, soil biotic composition and functioning, and carbon storage. Several studies suggest that the effect of reindeer on vegetation may follow predictable patterns and could induce an alternative stable vegetation state. However, little empirical data on the long-term stability of reindeer effects on vegetation exist, as it is inherently challenging to study these ecological processes experimentally on a sufficiently long timescale. The main objective of this thesis was therefore to gain a better understanding of the long-term ecological processes following reindeer-induced vegetation shifts. In order to gain a more mechanistic insight in what initially drives this transition, I used a field-based grazing simulation experiment in which I separated defoliation, trampling, moss removal and the addition of feces. This allowed me to test the relative contribution of reindeer-related activities to initiating the shift from moss and heath- dominated tundra towards a graminoid-dominated vegetation state. Additionally, I studied the long-term ecological stability following such a vegetation shift. I did this by addressing historical milking grounds (HMGs): sites where high reindeer concentrations associated with historical traditional reindeer herding practices induced a vegetation transition from shrubs towards graminoids several centuries earlier, but which were abandoned a century ago. Studying HMGs allowed me to address: 1. The potential stability of reindeer-induced vegetation shifts; 2. The ecological mechanisms contributing to the long-term stability of these vegetation shifts; and 3. How such long-lasting vegetation changes influence soil carbon- and nutrient cycling. I found that trampling by reindeer is an important mechanism by which reindeer cause vegetation change. Addressing HMGs further revealed that this vegetation change can be hightly persistent, as the studied HMGs showed only a low encroachment at the surrounding borders in the last 50 years. The vegetation in the core areas of all studied HMGs had remained strikingly stable, and were hardly invaded by surrounding shrubs. Interestingly, soil nutrient concentrations and microbial activities were still different from the surrounding area as well, and even comparable to actively grazed areas. Even after many centuries of changed vegetation composition and soil processes, there was no difference in total carbon sequestration. This suggests that the environmental conditions for microbial decomposition were more important than vegetation composition for the soil carbon stocks, in our study site. After studying the contemporary habitat use of HMGs by reindeer and other herbivores, investigating the potential plant-soil feedbacks mechanisms and detailed soil analyses, I concluded that several ecological mechanisms contribute to the long-term stability of HMGs: first, the altered soil biotic and abiotic conditions appear to have a stronger advantage for HMG vegetation than for the surrounding tundra vegetation. Furthermore, I found a clear browsing preference of small rodents on single shrubs proliferating in HMGs, causing a strong limitation on shrub expansion. Moreover, the dense established sward of graminoids likely poses a strong direct competition for space and nutrients, hindering seedling establishment. Finally, I conclude that HMGs are highly stable on relevant ecological timescales, and propose how the concepts of historical contingency and ASS can be applied to understand stability of these reindeer-induced vegetation transitions.

La nutrition minérale des plantes dépend à la fois du développement et du fonctionnement de leur appareil racinaire, incluant l'absorption mais aussi la capacité des plantes à influencer les cycles des nutriments, notamment par l'exsudation. Le but de cette thèse est de lier les rétroactions plantes-sol impliquant les cycles des nutriments aux stratégies d'exploration racinaire. Dans la revue bibliographique, je recense des mécanismes d'interaction plantes-sol et leurs échelles spatiales et temporelles. En considérant, à l'échelle de la rhizosphère, les interactions directes entre racines et sol, je propose que la combinaison entre exsudation et absorption des nutriments mène à des synergies entre racines d'une même plante. Ma seconde hypothèse est celle de l'existence d'un compromis entre l'exploration du sol et son occupation (défini comme la capacité des plantes à influencer efficacement le cycle des nutriments). Dans un premier chapitre, je développe un modèle général de recyclage des nutriments afin de déterminer sous quelles conditions les plantes auraient intérêt à limiter leur exploration du sol. Je montre qu'une exploration limitée est une stratégie de nutrition efficace sous certaines conditions, dont l'existence de synergies entre racines et le fait d'être dans un sol pauvre en nutriment. Dans un deuxième chapitre, je mesure le patron d'exploration racinaire et évalue le recyclage de l'azote à l'aide des outils isotopiques, chez trois espèces de Poacées pérennes de la savane de Hwange (Zimbabwe). Cette étude de terrain montre un gradient d'hétérogénéité racinaire entre ces trois espèces. Les Poacées exprimant le patron d'exploration le plus hétérogène ont un cycle de l'azote plus lent, mais potentiellement plus efficace. Dans un dernier chapitre, je développe un modèle mécaniste à l'échelle de la rhizosphère, pour une plante absorbant le phosphore et contrôlant sa disponibilité par l'exsudation de citrate. Je montre que, selon l'échelle d'influence des racines en terme d'exsudation et d'abaissement de la concentration en phosphore, la combinaison de l'exsudation et de l'absorption mène soit à une compétition, soit à une facilitation entre les racines d'une même plante. En me plaçant à l'échelle du système racinaire, je montre que les pertes en phosphore sont limitées par une exploration limitée du sol. Ce dernier résultat va dans le sens du compromis exploration/occupation. Au cours de cette thèse, j'ai donc développé des approches complémentaires, mettant en jeu différents mécanismes et échelles d'interactions plantes-sol. Le fait que les racines ne se limitent pas à un rôle d'absorption, mais agissent activement sur les cycles de nutriments a mené à deux résultats originaux : la facilitation inter-racinaire et intra-plante, et le fait qu'une exploration limitée puisse être considérée comme une stratégie efficace de nutrition. Enfin, ce travail souligne l'importance d'intégrer les divers mécanismes d'interaction plantes-sol pour comprendre les stratégies de nutrition des plantes et mieux prédire leur impact sur les cycles de nutriments à l'échelle des écosystèmes. / Plant nutrition depends on complementary mechanisms : the development of root systems, root uptake and plant ability to control nutrient cycling, e.g. through exudation. The aim of this thesis is to link plant-soil feedbacks involving the cycling of nutrients and root foraging strategies. I first review the different mechanisms of plant influence on nutrient cycling within the soil and assess their respective scales. Considering the direct effect of roots on the soil at the scale of the rhizosphere, I hypothesize that the combination of absorption and exudation may lead to synergies between the roots of a plant. At the scale of the whole root system, I propose a second, heuristic hypothesis: the existence of a trade-off between soil exploration and soil occupation (defined as the ability of plants to influence efficiently nutrient cycling). In a first chapter, I develop a general model of nutrient cycling, to determine under which condition plants should limit the exploration of soil by their roots. I show that limited exploration is an efficient strategy under specific conditions, especially nutrient-poor soils and the existence of synergies between roots. In a second chapter, I characterize soil occupation and nitrogen cycling, by the use of isotopes ratios, in the plant-soil system of three perennial grasses of the savanna of Hwange (Zimbabwe). This field study shows a gradient of root heterogeneity among these grass species. Those showing the more heterogeneous root pattern have a slower but potentially more efficient nitrogen cycling. In a last chapter, I develop a numerical mechanistic model at the rhizosphere scale for a plant taking up phosphorus and increasing its availability through exudation of citrate. I show that, depending on the extent of root influence on soil by exudation and nutrient depletion, competition between roots as well as facilitation arise from the combination of root uptake and exudation. By upscaling rhizosphere processes to the root system, I show that phosphorus losses are minimized by a restricted soil exploration, which backs the hypothesis of a trade-off between soil exploration and occupation. Overall, I developed complementary approaches that took into account several mechanisms and scales of plant-soil interactions. Considering that root functions are not limited to nutrient uptake, but also involve their influence on nutrient cycling, lead to two novel results: the potential existence of intra-plant and inter-root facilitation, and limited soil exploration as an efficient foraging strategy. This work underlines the importance of accurately integrating the mechanisms of plant-soil interaction to assess their nutrient strategies and to predict their impact on nutrient cycling within ecosystems.