Increased knowledge of plant interactions is important for our understanding of how ecosystems will respond to climate changes. Using four common low-herb and three tall- herb species as phytometers I measured the net outcome of plant interactions in an alpine environment by a neighbour removal experiment. Grazing and climate were tested as explanatory factors for differences in the outcome of plant interactions, with two altitudes representing different climates. The most important finding in this experiment is that competition is the dominating interaction among plants in this habitat, regardless of plant size, climate and grazing. Climatic exposure and grazing only influenced tall-herb species while low-herbs were mainly limited by competition, presumably for light. These results are important since facilitative interactions and net facilitation in plant communities are often reported to become more common in severe climates.

Alpine plants are assumed to be in particular danger as the climate changes rapidly worldwide. Specialist alpine species in Norrbotten County, northern Sweden have been surveyed over the last 20 years, providing insight to population dynamics and how the plants might respond to the changing climate. The main current threat to the species is habitat destruction as the climate changes. Variation in the number of plants among populations and years, and correlations with environmental variables were examined. Some species had increased while others had decreased over the years. No uniform relationship for all species and populations were discovered, but some of the species exhibited relationships between population size changes and temperature and precipitation. However, if the future climate in Norrbotten County changes according to the predictions, the habitats of the specialist alpine plants may be severely altered, leaving the species with no alternative places to establish and grow.

Alpine plants are assumed to be in particular danger as the climate changes rapidly worldwide. Specialist alpine species in Norrbotten County, northern Sweden have been surveyed over the last 20 years, providing insight to population dynamics and how the plants might respond to the changing climate. The main current threat to the species is habitat destruction as the climate changes. Variation in the number of plants among populations and years, and correlations with environmental variables were examined. Some species had increased while others had decreased over the years. No uniform relationship for all species and populations were discovered, but some of the species exhibited relationships between population size changes and temperature and precipitation. However, if the future climate in Norrbotten County changes according to the predictions, the habitats of the specialist alpine plants may be severely altered, leaving the species with no alternative places to establish and grow.

La niche climatique des espèces joue un rôle important dans la distribution spatiale de la biodiversité mais la manière dont les niches climatiques évoluent reste encore peu connue. Ce travail vise à révéler la manière dont les niches climatiques évoluent en général, et plus précisément à déterminer comment certaines plantes se sont adaptées aux environnements alpins. En étudiant de nombreux groupes de plantes, de poissons, de mammifères et d'oiseaux, nous avons montré que les niches climatiques évoluent le plus souvent par à-coups et non pas de manière graduelle. Les niches climatiques restent en effet stables pendant des périodes de plusieurs millions d'années puis évoluent de manière extrêmement rapide avant de se stabiliser à nouveau dans une autre gamme de climat. Des simulations ont permis de montrer que les phases de relative stabilité n'étaient pas forcément causées par une sélection stabilisante sur les niches climatiques mais pouvaient également résulter de la présence de barrières géographiques qui empêchent les espèces d'expérimenter de nouveaux climats. L'étude de l'histoire des plantes du genre Androsace a révélé que les changements rapides de niches correspondaient au contraire à l'apparition de nouveaux traits, comme la forme de vie en coussin. Ce travail montre que de nombreux facteurs influencent l'évolution des niches climatiques et souligne la nécessité de tous les étudier ensemble. / Species' climatic niches play an important role in the spatial distribution of biodiversity but the way climatic niches evolve remains poorly known. This works aims at determining the general mode of evolution of climatic niches, and more precisely at revealing how some plants have adapted to alpine environments. The study of many groups of plants, fishes, mammals and birds has shown that climatic niches usually evolve by fits and starts but not gradually. Climatic niches indeed remain stable over million years before evoving extremely quickly and stabilizing again in a new range of climates. Simulations have shown that these phases of relative stability need not be caused by stabilizing selection but can also be caused by geographic barriers that impede species from experiencing new climates. The study of the history of plants of the genus Androsace has revealed that rapid niche shifts on the contrary resulted from the appearing of novel traits, like the cushion life form. This work shows that numerous factors contribute to the evolution of climatic niches and emphasizes the necessity to study them together.

Le climat alpin à plus de 2400 mètres d’altitude montre des fortes variations de température, des intensités lumineuses très élevées (3000 µmol photons m-2 s-1) qui sont connues pour générer un état de réduction importante de la chaine de transport des électrons photosynthétique. Le bon fonctionnement du processus photosynthétique est primordial pour les quelques espèces de plantes vasculaires qui sont présentes à l’étage alpin et qui doivent terminer leur cycle de vie lors d’une très courte période de végétation.Soldanella alpina et Ranunculus glacialis sont deux espèces inféodées aux étages alpin et nival. Dans leur site naturel de croissance nous avons mesuré des températures faibles (0.7°C) et fortes (37°C) sous des lumières supérieures à 2500 µmol photons m-2 s-1. Chez les espèces non-alpines ces conditions induisent la photoinhibition du PSII, ce qui est évité chez S. alpina et R. glacialis, par des mécanismes très différents. Les systèmes antioxydants et le quenching non photochimique sont particulièrement importants chez S. alpina. Chez Ranunculus glacialis, la photorespiration reste très importante et un contenu élevé en PTOX est décrit. Le rôle des antioxydants et de la PTOX dans la photoprotection des deux espèces ont été étudiés. Dans une partie de thèse, nous avons montré qu’une diminution de la capacité antioxydante par une diminution de la concentration en glutathion n’affecte pas la tolérance vis-à-vis de la photoinhibition à basse température. Dans une deuxième partie les résultats supposent qu’une surexpression de la PTOX chez le tabac augmente la photoinhibition à lumière forte par production des espèces réactives d’oxygène. En utilisant différentes conditions environnementales de croissance pour Ranunculus glacialis, nous avons pu montrer que l’expression de la PTOX est induite par des fortes lumières et non par des basses températures. Grâce à une approche associant mesures d’échanges gazeux et mesures de la fluorescence de la chlorophylle, nous avons montré qu’un flux d’électrons conséquent vers l’oxygène, indépendant de la photorespiration, corrélait avec la présence de la PTOX mais que l’activité de la PTOX sous des conditions qui permettent l’assimilation du CO2 et la photorespiration n’est pas maximale. Grâce à des mesures de fluorescence chlorophyllienne en présence de différents inhibiteurs photosynthétiques, nous avons pu montrer que l’importance de ce flux d’électrons vers l’oxygène corrèle avec la quantité de PTOX présente dans les feuilles, dans des conditions réductrices. Ces résultats nous ont amenés à conclure que chez Ranunculus glacialis, la PTOX peut prendre en charge un flux significatif d’électrons, éviter ainsi l’apparition d’un état réduit de la chaine de transfert photosynthétique, et protéger la plante vis-à-vis de la photoinhibition en agissant comme une valve de sécurité. Ces travaux permettent d’apporter des précisions sur un modèle original de photoprotection, qui a été l’objet de nombreuses controverses. / The alpine climate above 2400 meters altitude shows large variations in temperature and very important light intensity (3000 µmol photons m-2 s-1), which are known to generate a state of significant reduction in the photosynthetic electron transport chain. The proper functioning of the photosynthetic process is essential for vascular plants species that are present in this alpine environment and must complete their life cycle within a very short growing season.Soldanella alpina and Ranunculus glacialis are two species restricted to alpine and snow floors. In their natural growth environment we measured very low (0.7 ° C) and high temperature (37 ° C) under lights above 2500 µmol photons m-2 s-1. Among non-alpine species such conditions induce photoinhibition of PSII, which is avoided in S. alpina and R. glacialis, by very different mechanisms. Antioxidant systems and non-photochemical quenching are particularly important in S. alpina. In Ranunculus glacialis, photorespiration remains very important and a high content of PTOX is described. The roles of antioxidants and PTOX in photoprotection of both species were studied.In one part of the thesis, we showed that a decrease in antioxidant capacity by reducing the concentration of glutathione does not affect tolerance to low-temperature photoinhibition. In the second part the results imply that overexpression of PTOX in tobacco enhances photoinhibition by strong light to produce reactive oxygen species.Using different environmental conditions for Ranunculus glacialis growth, we showed that expression of the PTOX is induced by strong light, but not by low temperatures. With an approach combining gas exchange measurements and chlorophyll fluorescence measurements, we showed that an electron flow to oxygen, independent of photorespiration, correlated with the presence of PTOX. Through measures of chlorophyll fluorescence in the presence of various inhibitors photosynthetic, we could show that the importance of this electron flow to oxygen correlates with the amount of PTOX in the leaves, under reducing conditions. These results led us to conclude that in Ranunculus glacialis, the PTOX may support a significant flow of electrons, thus avoiding the appearance of a reduced state of the photosynthetic chain transfer, and protect the plant from photoinhibition, acting as a safety valve. These studies are discussed to help clarify a new pathway of photoprotection, which was the subject of much controversy.

Hybrids are common, being found in about 25% of all plant species, but the isolating barriers which preserve species integrity are poorly studied. I investigated this question in the large New Zealand genus Celmisia Cass. (Asteraceae), which hybridises readily in cultivation, but wild hybrids are relatively rare. My study quantitatively tests four potential reproductive isolating barriers in 12 sympatric species of Celmisia found in the Craigieburn Range, inland Canterbury, New Zealand. I examined two potential prezygotic reproductive isolating barriers (flowering phenology and pollinator specialisation), and two potential postzygotic barriers (pre-dispersal seed predation and hybrid seed germination). I used null models to test whether Celmisia species had temporally segregated flowering times, and found that some Celmisia are temporally segregated and thus less likely to form hybrids. I used experimental pair-wise flowering arrays to observe insect visitation to six different Celmisia species pairs. While I found no difference in the overall pollinator community, several insect families showed preferences for some Celmisia species. Furthermore, I found that subtle floral character differences were driving these insect preferences. In particular, I found scape height to be positively associated with insect visitation with taller Celmisia being favoured over shorter species. Insect preferences did not translate into strong floral constancy, therefore indicating that Celmisia flower visitors are likely to be a weak barrier to hybridisation. I reared a range of insect seed predators from field-collected capitula of the hybrid C. x pseudolyallii and both parent species (C. lyallii and C. spectabilis). There was no overall difference in the number of seed-predators per capitulum between hybrid and parent Celmisia taxa. I collected and sowed seeds from three Celmisia hybrids and their parent species in order to test whether hybrids were less fertile than their parent species. I found no evidence to suggest that the seeds of hybrids had lower germination success than those of their parents. Overall I found evidence for only weak prezygotic reproductive isolation and no evidence for postzygotic isolation in the four barriers I examined in Celmisia.

Cette thèse a pour objectif principal de contribuer à la compréhension des facteurs environnementaux agissant sur la diversité et la productivité des communautés végétales alpine, et dans un deuxième temps de mettre en évidence des changements récents dans la structure de la végétation dans un contexte tempéré alpin, les Alpes françaises. L’approche adoptée se base sur des méthodes issues de la télédétection et également de l’écologie végétale, en combinant des relevés de végétation avec les données de climat de l’imagerie à haute résolution. Dans le premier chapitre, je démontre l’intérêt de quantifier la durée d’enneigement afin de pouvoir prédire les patrons de diversité taxonomique et fonctionnelle, en montrant aussi que la modélisation spatiale de la productivité primaire reste un défi important dans le contexte des paysages alpins. Dans le deuxième chapitre, je teste l’utilité de l’imagerie satellitaire pour quantifier les conditions environnementales au niveau des communautés végétales, et je mets en évidence que l’habitat des communautés dominantes peut être différencié à l’échelle régionale par la date de déneigement et le pic en productivité. J’essaie également de voir si la diversité fonctionnelle influe sur la réponse NDVI dans le cadre de trois années de durée d’enneigement contrastées. Le troisième chapitre marque un changement d’approche, dans lequel je passe à l’analyse de la dynamique observée de la végétation pendant les dernières décennies dans deux sites : la Réserve Naturelle des Hauts Plateaux du Vercors (RNHPV) et le Parc National des Ecrins (PNE). L’analyse diachronique de l’écotone forêt-prairie dans la RNHPV montre une forte expansion de la forêt à Pinus uncinata pendant les soixante dernières années, en réponse à un réchauffement global et également des changements spatio-temporels dans les usages des terres en lien avec le pâturage. Dans la deuxième partie, je montre qu’avec les satellites on observe une tendance de verdissement à l’échelle du PNE pendant les dernières décennies. Je propose qu’une réduction dans la durée d’enneigement et un réchauffement depuis les années 1980s ont contribué à une augmentation de la productivité des canopées dans des contextes alpins de haute altitude, et ceci indépendamment des effets d’utilisation des terres. Pris ensemble, au lieu d’avancer un aspect précis de l’écologie alpine, mon travail sert à compléter nos connaissances du fonctionnement des communautés végétales alpines et à confirmer certaines hypothèses basées sur l’observation terrain avec des analyses spatiales robustes. / The central aim of this thesis is to contribute to current understanding of environmental drivers of plant diversity and productivity as well as of recent changes in vegetation structure in a temperate alpine context, the French Alps. My approach draws on methods from remote sensing and plant ecology by combining plot-based measures of plant diversity and climate data with high-resolution imagery. Chapter I demonstrates the importance of quantifying snow cover duration for predicting patterns of plant taxonomic and functional diversity, and also highlights the ongoing challenge of modeling spatial gross primary productivity dynamics in alpine landscapes. In Chapter II, I explore the utility of satellite imagery for quantifying environmental conditions experienced by alpine plant communities, and further show how metrics of snow cover duration and peak productivity can be used to differentiate habitat for dominant alpine plant communities. I also explore how functional diversity mediates NDVI responses to highly contrasting snow years. Chapter III provides new evidence of recent shifts occurring in high-elevation plant communities in the French Alps in response to climate and land-use change. Analysis of the forest-grassland ecotone in the Vercors Regional Park shows a strong dynamic of forest expansion in response to overall climate warming and local shifts in grazing-related land-use practices, which supports findings from other studies conducted elsewhere in the Alps and Pyrenees. In the second part of Chapter III, for the first time I present evidence of greening dynamics in a protected area of the French Alps, the Ecrins National Park. I propose that a decrease in snow cover duration and pronounced warming occurring in the 1980s likely contributed to increased canopy productivity in high alpine contexts, and are driving long-term greening in the absence of land-use change. Taken collectively, rather than pushing a specific aspect alpine ecology forward, my work helps to fill out our working knowledge of alpine plant communities and serves to solidify a number of field-based observations by carrying out robust spatial analyses.

The westernmost spur of the Tibetan Plateau stretches to Eastern Ladakh in India. It is a region which remains poorly explored because of challenging conditions and long periods of political instability. At the same time, it is one of the highest places on earth supporting angiosperm life, which goes beyond 6000 m a.s.l. here. The whole region, due its remoteness, is practically unaffected by plant invasions and direct human activities. Thus, Ladakh represents a kind of "natural experiment", providing very long gradient of elevation suitable for comparative functional ecology as well as for testing various hypotheses concerning limitations of vascular plants. Arid climate and extreme elevations are the common factors. Our team pursued the goal of systematic botanical and ecological exploration of Ladakh, started by late Leoš Klimeš. This thesis provides insight into the main vegetation types, clonality in plants, plant-plant interactions and soil phototroph communities.