Les interactions entre organismes ont des conséquences majeures sur la composition des communautés et le fonctionnement des écosystèmes. En écologie l'étude des interactions négatives, telles que la prédation et la compétition, a largement dominé la littérature. Des travaux récents ont souligné l’importance des interactions positives dans la nature, telles que la facilitation, mais ces interactions restent malgré tout peu intégrées dans les théories contemporaines en écologie. Cette thèse s’inscrit dans cette démarche et aborde deux questions centrales : (i) Comment la facilitation indirecte (via la protection contre le pâturage) affecte-t-elle la structure fonctionnelle (caractéristiques des distributions de traits) des communautés de plantes ? (ii) L'effet net des interactions facilitatrice-bénéficiaire identifiées à l’échelle d’une paire d’espèces restent-elles valides en présence d’une communauté entière d'espèces bénéficiaires potentielles ? Deux expérimentations in situ dans des environnements contrastés ont été mises en place : l’exclusion du pâturage dans des tourbières tropicales alpines et une transplantation de communautés de plantules sous des plantes adultes en milieu méditerranéen semi-aride. Nous avons montré que la facilitation indirecte affectait les caractéristiques du filtre environnemental, la dominance des espèces et la différenciation de niche au sein de la communauté. Nos travaux suggèrent également que la composition des communautés de plantules modifie les interactions adultes-plantules, remettant ainsi en cause la possibilité d’extrapolation des résultats entre paires d'individus à l’échelle de la communauté. / Interactions between organisms are key drivers of community composition and ecosystem functioning. Ecology has a long history of studies on negative interactions, such as predation and competition. Recent studies have highlighted the importance of positive interactions, such as facilitation in nature. The integration of these interactions into modern ecological theory has nonetheless lagged behind. This thesis aims at contributing to this research effort and addresses two core questions : (i) How does indirect facilitation (through protection against grazing) affect the functional structure (characteristics of trait distributions) of plant communities? (ii) Does the net effect of the interactions between a species pair (benefactor-beneficiary) remain valid in the presence of several beneficiary species at community level? We set up two insitu experiments in contrasted environments: a grazing exclusion experiment in tropical alpine peatlands and a transplantation experiment of sapling communities beneath adult plants in a mediterranean environment. Our results showed that indirect facilitation affects the characteristics of the environmental filter, species dominance and niche differentiation in the community. Our results also suggested that the composition of sapling communities modifies adult-sapling interactions, thereby questioning the possibility of extrapolating results from pairs of individuals to the community scale.

The object of this thesis was to study plant community structure, especially in relation to vertebrate herbivory, in an altitudinal gradient in the Fennoscandian mountain chain. A sowing experiment in a high alpine Ranunculus glacialis population showed that seeds germinated better in cleared microsites than under established individuals. This is contrasted with a hypothesis that predicts positive plant-plant interactions in high alpine environments. It was concluded that plant-plant interactions in die studied population varied from neutral to negative, whereas no indications for positive interactions were found. An exclosure experiment in a snow-bed showed that a lemming population consumed 33 % of the available graminoids and 66 % of the mosses from August to June during a population peak. The results shows that grazing needs to be considered as a structuring factor in snow-bed vegetation. The vegetation in exclosures in another snow-bed changed from a graminoid-dominated to a herb-dominated plant community during a long-term (six years) experiment No changes of the same magnitude were seen in a tall herb meadow on a lower altitude. Survival of transplanted adult shoots from the tall herb meadow was equally high in the snow-bed as on the meadow, and germination was also high on bare ground in the snow-bed. Grazing seemed to be a more important structuring factor in the snow-bed than in the more productive tall herb meadow. Raising the grazing pressure during one growing season by introducing microtine rodents into enclosures did not cause any large short-term effects on plant community structure in a tall hob meadow or in a snow-bed. Marked shoots showed that some preferred plant species had a high shoot mortality, but biomass for pooled categories of plants was not significantly affected. It was predicted that the tall herb meadow would be more grazing sensitive than die snow-bed, but productivity on the meadow seemed to be sufficiently high for the plants to compensate for the grazing during the growing season. A greenhouse experiment showed that voles, when grazing freely, have the potential to deplete productive field layer vegetation contrary to predictions from plant defence theories. A nitrogen-based defence did not prevent heavy shoot mortality for toxic tall herbs. / <p>Diss. (sammanfattning) Umeå : Umeå universitet, 1993, härtill 5 uppsatser.</p> / digitalisering@umu.se

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.

Les plantes vivent en association avec une grande diversité de microorganismes qui forment son microbiota. Ce microbiote fournit des fonctions clés qui influencent tous les aspects de la vie d'une plante, de l'établissement à la croissance jusqu'à la production. Cette thèse a pour intention de déterminer les règlent d'assemblage du microbiote et ses conséquences pour le phénotypel l'adaptation et l'évolution des plantes. Pour atteindre cet objectif nous avont utilisé différentes approches expérimentales comprenant des plantes clonales comme organismes modèles ainsi que des mésocosmes prairiaux pour analyses à l'échelle des communautés. Nos résultats ont démontré i) que les Champignons Mycohiziens à Arbuscules induisent d'important es variations phénotypiques pour les traits des plantes clonales impliqués dans l'exploration de l'espace et l'exploitation des ressources. Ces changements dépendent de l'identité des symbiontes et altèrent les capacités des plantes à développer des réponses plastiques à l'hétérogénéité environnementale. ii) Les plantes ont évolué un méchanisme permettant la transmission d'une partie de leur microbiote a leur descendance, assurant la qualité de leur habitat. iii) Le contexte spécifique des communautés de plantes est un facteur majeur structurant l'assemblage du microbiota des plantes à échelle locale. L'abondance de certaines espèces de plante dans le voisinage d'une plante cible augmente ou diminue la diversité de son microbiote, déterminant in fine ses performances. De manière générale, cette thèse démontre l'importance des organismes symbiotiques dans la compréhension de l'adaptation et de l'évolution des plantes. / Plants live in association with a wide diversity of microorganisms forming the microbiota. The plant microbiota provides a variety of key functions that influence many aspects of plant's life comprising establishment, growth and reproduction. The present thesis aims at determining the assembly rules of the plant microbiota and its consequences for plant phenotype, adaptation and evolution. To fulfill this objective, we used different experimental approaches using either clonal plants as model organisms or grassland mesocosms for community-wide analyses. Our results demonstrated i) that Arbuscular Mycorrhizal Fungi induce important phenotypic variations in clonal plants traits involved in space exploration and resources exploitation. These changes depended on the identity of the symbionts and altered the plants ability to produce plastic responses to environmental heterogeneity. ii) Plants have evolved a mechanism allowing the transmission of a part of their microbiota to their progeny, ensuring thus their habitat quality. iii) The plant community context is a major factor structuring local plant microbiota assembly. Particular plant species identity in the neighborhood increase or decrease the microbiota diversity and ultimately determine the focal plant performance. This thesis overall demonstrates the importance of symbiotic microorganisms in the understanding of the plant adaptation and evolution. From the knowledges acquired we developed a novel understanding of symbiotic interactions in clonal plants by extending the holobiont theory to the meta-holobiont theory.

Background: Pollen transfer via animals is necessary for reproduction by ~80 % of flowering plants, and most of these plants live in multispecies communities where they can share pollinators. While diffuse plant-pollinator interactions are increasingly recognized as the rule rather than the exception, their fitness consequences cannot be deduced from flower visitation alone, so other proxies, functionally closer to seed production and amenable for use in a broad variety of diverse communities, are necessary. Scope: We conceptually summarize how the study of pollen on stigmas of spent flowers can reflect key drivers and functional aspects of the plant-pollinator interaction (e.g. competition, facilitation or commensalism). We critically evaluate how variable visitation rates and other factors (pollinator pool and floral avoidance) can give rise to different relationships between heterospecific pollen and (1) conspecific pollen on the stigma and (2) conspecific tubes/grain in the style, revealing the complexity of potential interpretations. We advise on best practices for using these proxies, noting the assumptions and caveats involved in their use, and explicate what additional data are required to verify interpretation of given patterns. Conclusions: We conclude that characterizing pollen on stigmas of spent flowers provides an attainable indirect measure of pollination interactions, but given the complex processes of pollen transfer that generate patterns of conspecific-heterospecific pollen on stigmas these cannot alone determine whether competition or facilitation are the underlying drivers. Thus, functional tests are also needed to validate these hypotheses.

<p>Poslednjih godina je prisutan trend povratka prirodi i upotrebi biljnih lekova, kako u prevenciji tako i u lecenju razlicitih bolesti. Timijan (Thymus vulgaris L.) se u narodnoj medicini koristio u lecenju respiratornih oboljenja kao &scaron;to su ka&scaron;alj, bronhitis i astma. Rezultati novijih istraživanja pokazuju da timijan poseduje i druga potencijalno korisna farmakolo&scaron;ka svojstva (antimikrobna, antiinflamatorna, antioksidativna, spazmoliticka, antidijabetesna i anksioliticka). Ciljevi ovog istraživanja su bili da se ispitaju farmakodinamske osobine preparata timijana, njihove interakcije sa lekovima koji deluju na centralni nervni sistem, uticaj na funkciju jetre i parametrem oksidativnog stresa kod životinja izloženih ugljentetrahloridu, kao sadržaj karvakrola i timola u sirupu timijna, pri razlicitim uslovima cuvanja. U farmakodinamskim ispitivanjima kao eksperimentalne životinje kori&scaron;ceni su mi&scaron;evi soja NMRI, a u svim drugim ispitivanjima pacovi soja Wistar. Tinktura timijana je primenjena u dozi od 0,4mk/kg, a sirup u dozi od 12,08 ml/kg, na mi&scaron;evima. Primenjene doze na pacovima su bile 0,18 ml/kg za tinkturu i 5,6 ml/kg za sirup timijana. Za ispitivanje analgetickog dejstva kori&scaron;ceni su metod vrele ploce i test sircetne kiseline. Za procenu motorne koordinacije kori&scaron;cen je test rotirajuceg &scaron;tapa, a za procenu hipnotickog delovanja mereno je vreme spavanja. Prilikom ispitivanja uticaja preparata timijana na farmakokinetiku paracetamola, odre_ivana je koncentracija ovog leka HPLC metodom, a nakon toga su odreeni farmakokinetski parametri paracetamola. Antioksidantna aktivnost preparata timijana odre_ivana je pomocu in vitro i in vivo testova. Nakon žrtvovanja životinja ra_ena je histopatolo&scaron;ka analiza jetrenog tkiva, a u serumu su odre_ivani biohemijski parametri, kao i pokazatelji bubrežene i jetrene funkcije. Sadržaj timola i karvakrola i sirupu timijana odre_en je GC/MS metodom. Sirup i tinktura timijana su pokazali analgeticki efekat u testu vrele ploce, kao i smanjenje broja grceva izazvano primenom sircetne kiseline. Sedmodnevna primena preparata timijana smanjila je analgeticko dejstvo kodeina, a pojacala analgeticki efekat paracetamola. Sirup timijana je potencirao diazepamom izazvan poremecaj motorne koordinacije. Ispitivanjem uticaja preparata timijana na hipnoticko delovanje pentobarbitala, postignuti su razliciti rezultati u zavisnosti od dužine trajanja pretremana. Sedmodnevna primena timijana je produžila vreme trajanja spavanja, dok je jednokratna primena timijana skratila vreme trajanja spavanja. Nakon i intravenske i peroralne primene paracetamola, grupe životinja koje su bile pretretirane preparatima timijana imale su krace poluvreme eliminacije i vecu konstantu eliminacije. Upotreba samo preparata timijana nije imala uticaj na biohemijske i histolo&scaron;ke promene jetrene funkcije. S druge strane, upotreba tincture timijana u kombinaciji sa ugljen-tetrahloridom dovela je do porasta vrednosti AST i ALT enzima u serumu, dok je sirup timijana u kombinaciji sa ugljentetrahloridom smanjio aktivnost aminotransferaza. Najvece odstupanje u koncentracijama aktivnih komponenti timola i karavkrola, pokazali su sirupi cuvani na sobnoj temperaturi (20&deg;C), u sekundarnoj ambalaži i na svetlom mestu. Rezultati dobijeni u toku ovog istraživanja ukazuju da preparati timijana uticu na farmakodinamske osobine kodeina, paracetamola, diazepama i pentobarbitala, kao i na farmakokinetiku paracetamola. Upotreba preparata timijana ispoljila je analgeticki efekat i umanjila posledice izloženosti oksidativnom stresu. Uslovi cuvanja sirupa timijana uticali su na njegovu stabilnost.</p> / <p>In recent years is present trend of return to nature and the use of herbal medicines in prevention and treatment of different diseases. Thyme (Thymus vulgaris L.) was used in folk medicine in the treatment of respiratory diseases such as cough, bronchitis and asthma. The new research results have demonstrated that thyme has many others potentially useful pharmacological properties (antimicrobial, antiinflammatory, antioxidant, antispasmodic, antidiabetic and anxiolytic). The aims of this research were to determine the pharmacodynamic properties of thyme preparations and their interactions with central nervous system drugs, influence on liver function and oxidative stress parameters of animals exposed to carbon tetrachloride, as well as concentration of thymol and carvacrol in thyme syrup, at different storage conditions. In pharmacodynamics examination as experimental animals were used NMRI mice, while in all other test were used Wistar rats. Applied dose of thyme tincture was 0.4 ml/kg and of syrup 12.08 ml/kg, for mice. For rats, applied doses of tincture and syrup were 0.18 ml/kg and 5.6 ml/kg, respectively. The analgesic activity was examined by the hot plate test and acetic acid test. The Rotarod test was used to evaluate the motor coordination and to evaluate hypnotic activity sleeping time was mesaured. In order to examine the influence of thyme preparations on pharmacokinetics of paracetamol, the concentracion of this drug was measured by HPLC metods, and after that pharmocokinetic parameters of paracetamol were determined.The antioxidant acivity of thyme preparations was determined by using in vitro and in vivo tests. After animals sacrificing, histopathological analysis of liver tissue were peroformed, in serum were determined biochemical parameters and renal and hepatic function parameters. Quantification of thymol and carvacrol in syrup was carried out by GC/MS method. Thyme syrup and thyme tincture exhibited analgesic activity in hot plate test and reduced the number of writhes induced by acetic acid. Seven-day pretreatment with thyme preparations reduced analgesic activity of codeine and increased analgesic effect of paracetamol. Thyme syrup potentiated diazepam induced motor coordination impairment. Examining the impact of thyme preparations on hypnotic effect induced by pentobarbital, different results were achieved depending on the duration of pretreatment. Seven-day pretreatment with thyme had prolonged the sleeping time, while after single dose of thyme the sleeping time was decreased. After intravenous and after oral administration of paracetamol, groups pretreated with thyme preparations had decreased elimination half-life and increased elimination constant rate. Administration of thyme preparations alone did not change biochemical nor histological markers of hepatic function. On the other hand, co-administration of thyme tincture and carbon tetrachloride resulted in exacerbation of AST and ALT values in serum, while thyme syrup in coadministration with carbon tetrachloride managed to reduce activities of aminotransferases. The concentration of major active compounds, thymol and carvacrol, was mostly changed when syrups were stored at room temperature (20&deg;C), in secondary containers and in light place. Results obtained in this study demonstrated that thyme preparations do affect pharmacodynamic properties of codeine, paracetamol, diazepam and pentobarbital and pharmacokinetics of paracetamol. Administration of thyme preparations exhibited analgesic activity and reduced the effects of exposure to oxidative stress. Storage conditions of thyme syrup did affect its stability.</p>

Les plantes en coussins sont connues pour leur capacité fondatrice de nouvel habitat pour les autres espèces. Par ailleurs, des études ont montré que la variation morphologique au sein d’espèces fondatrices peut induire une variation des effets compétiteurs sur les espèces subordonnées, ce qui peut affecter la composition des communautés. Cette thèse a pour objectif d’étudier les conséquences d’une variation intraspécifique de deux espèces fondatrices (Festuca gautieri et Onobrychis cornuta) occupant des micro-environnements contrastés dans deux systèmes montagneux [les Pyrénées (France) et le Mont-Liban (Liban),respectivement] sur les communautés végétales associées (subalpines etoroméditerranéennes, respectivement). Nous avons évalué par des approches descriptives et/ou expérimentales (jardins expérimentaux, transplantations réciproques) les contributions de la génétique et de la plasticité à la variation morphologique entre deux phénotypes de coussins (dense et lâche) pour chaque espèce fondatrice, et à leurs effets contrastés sur les espèces subordonnées. Nous avons également quantifié les effets rétroactifs de la communauté pour la reproduction des espèces fondatrices. Nos résultats montrent une contribution à la fois de la génétique et de la plasticité à la variation phénotypique. La base génétique des variations morphologiques entre les phénotypes a induit des différences héréditaires d’effets compétiteurs sur les espèces subordonnées, tout en contrecarrant l’augmentation de la compétition avec la diminution du stress – le résultat dominant dans la littérature sur les systèmes subalpins. Nous avons aussi trouvé des effets rétroactifs négatifs des espèces subordonnées pour l’espèce fondatrice, avec une diminution de la production de fleurs (par les coussins) due au nombre croissant d’espèces subordonnées. La diversité des espèces subordonnées était plus élevée dans les conditions environnementales favorables que dans les stressantes. Par conséquent, nous avons conclu que les effets génétiques surmontent les effets environnementaux, limitant la compétition dans les milieux favorables, maintenant ainsi une plus grande diversité dans ces milieux que dans les milieux stressants. / Alpine cushion plants are foundation species known for their nursingability. Moreover, studies have shown that morphological variation in foundationspecies can trigger variation in competitive effects on subordinate species, likely toaffect community composition. We investigated the consequences of intraspecificvariation within two alpine cushion species (Festuca gautieri and Onobrychis cornuta)across heterogeneous environments in two mountain ranges [the Pyrenees (France)and Mount-Lebanon (Lebanon), respectively] for the associated plant communities(subalpine and oromediterranean, respectively). We assessed with observationaland/or experimental (common-gardens, reciprocal transplantation experiments)approaches the relative contribution of genetics and plasticity to the morphologicalvariation between two cushion phenotypes (tight and loose) of the foundationspecies, and to their differential effects on subordinate species. Communityfeedbacks were also quantified. Our results show that both genetics and plasticitycontributed to the phenotypic variation. The genetic basis of the morphologicaldifferences between phenotypes induced heritable differences in competitive effectson subordinate species, but however counteracted the general increase incompetition with decreasing stress dominantly found in the literature on subalpinesystems. We also found negative feedbacks of subordinates on foundation speciesfitness, with higher cover of subordinate species reducing the cushions flowerproduction. Subordinate species diversity was higher in benign than in stressedenvironmental conditions. Consequently, we concluded that genetic effects overcomethe environmental effects by limiting competition in benign physical conditions, thusmaintaining a higher diversity in benign than stressed conditions.

Au cours de son assemblage, une communauté végétale va subir de nombreux changements : immigration de nouvelles espèces de plantes possédant de nouveaux traits, disparition de certaines espèces de plantes avec d’autres traits, immigration de nouveaux organismes associés aux plantes (insectes, champignons…), changements de traits chez les espèces présentes… Tous ces changements sont susceptibles de modifier la décomposition de la litière produite par la communauté végétale. En effet, la décomposition de la litière est gouvernée par les traits des espèces végétales, par l’activité des organismes décomposeurs, et par le degré d’adaptation de ces organismes aux traits des espèces végétales. Cependant, les conséquences de l’assemblage de la communauté végétale pour la décomposition de la litière demeurent inconnues. L’objectif de cette thèse est de déterminer les conséquences de l’assemblage des communautés végétales prairiales sur la décomposition de leur litière, et ce à différentes échelles. Tout d’abord, nous avons étudié, très localement, les conséquences des plantes voisines que possèdent un individu pour la décomposition de sa litière (i.e. échelle intraspécifique). Nous avons distingué le cas où la litière de l’individu était seule, du cas où sa litière était mélangée à de la litière provenant d’autres espèces végétales. Puis, nous avons étudié les conséquences de l’assemblage sur la décomposition de la litière au niveau plus global de l’ensemble de la communauté végétale (i.e. échelle interspécifique). Enfin, nous avons exploré la rétroaction de la décomposition sur l’assemblage de la communauté. Deux grandes démarches expérimentales ont été développées, la première utilisant un dispositif de mésocosmes permettant de manipuler le voisinage local des individus, la seconde utilisant un dispositif Long Term Ecological Research (LTER) impliquant un vaste réseau de prairies avec différentes durées d’assemblage. À l’échelle locale, nos résultats indiquent qu’un individu qui possède des plantes voisines fonctionnellement dissemblables produit une litière plus décomposable et peut également abriter des décomposeurs plus efficaces. Lorsque la litière de cet individu est mélangée avec de la litière d’autres espèces, la décomposition du mélange est accélérée par des effets synergiques lorsque les plantes voisines sont évolutivement dissemblables et fonctionnellement éloignées du mélange. À l’échelle globale de l’ensemble de la communauté, nos résultats indiquent que tout au long de l’assemblage, de nombreux changements de traits fonctionnels des espèces végétales ont lieu (ratio C:N foliaire, teneur en matière sèche des feuilles, etc.) ainsi que des changements dans la composition de la communauté de décomposeurs (ratio C:N microbien). Ces changements impactent fortement la décomposition de la litière de la communauté prairiale mais s’annulent, maintenant le même taux global de décomposition. Enfin, nos résultats indiquent que plus la litière de couples d’espèces se décompose vite, notamment via des effets synergiques, plus ces espèces coexistent entre elles. Cette thèse met en évidence l’influence majeure de l’assemblage des communautés végétales prairiales sur la décomposition de leur litière, de l’invidu jusqu’à la communauté végétale toute entière. L’assemblage des communautés végétales peut donc influencer les processus écosystémiques d’après-vie tels que la décomposition de la litière. Cette influence se produit via les traits des plantes et l’activité de leurs décomposeurs. En retour, la décomposition de la litière impacte l’assemblage de la communauté végétale. La décomposition de la litière ne semble donc pas une conséquence collatérale des traits des espèces végétales, mais bien un élément important de leur stratégie écologique et de leurs interactions biotiques, situé au coeur d'une boucle de rétroaction avec les processus d'assemblage des communautés. / During its assembly, a plant community will be strongly modified: immigration of new plant species with new traits, disappearance of particular species with other traits, immigration of new plant-associated organisms (insects, fungi…), trait changes in existing species… All these changes are likely to drive the decomposition of litter produced by the plant community. Litter decomposition is indeed controlled by plant traits, activity of decomposer community, and adaptation of decomposer organisms to plant traits. However, the consequences of plant-community assembly on plant litter decomposition remain entirely unknown. This thesis aims at determining the consequences of plant-community assembly on plant litter decomposition, at distinct scales. First of all, we studied, locally, the consequences of neighboring plants on litter decomposition of plant individuals (i.e. intraspecific scale). We distinguished the case where litter of plant individuals was alone from the case where litter of plant individuals was mixed with litter from other species. Then we studied, more globally, the consequences of plant-community assembly on decomposition at the scale of the entire plant community (i.e. interspecific scale). Finally, we investigated whether plant litter decomposition feedbacks on plant-community assembly. We used two experimental approaches, the first one using a long-term mesocosm experiment for manipulating the local plant neighborhood of plant individuals, and the second one using of Long Term Ecological Research network involving grasslands with different time for assembly. At the local scale, our results indicate that plant individuals grown in functionally dissimilar neighborhood produce a more decomposable litter, and can also harbor more efficient decomposers. When the litter of these individuals is mixed with litter from other species, the decomposition of the litter mixture is accelerated by synergistic effects when neighboring plants are phylogenetically diverse, and functionnally dissimilar to the litter mixture. At the scale of whole plant community, our results show that numerous trait changes occur during assembly (leaf C:N ratio, leaf dry matter content…), as well as changes in the composition of the decomposer community (soil microbial C:N ratio). These changes strongly affect litter decomposition but offset each other, maintaining litter decomposition constant. Finally, our result show that the faster the decomposition of mixed-litter from two species is, the more both species coexist. This thesis demonstrates the major influence of plant-community assembly on plant litter decomposition in grassland ecosystems, from the scale of plant individuals to the scale of entire plant community. Plant-community assembly hence affects after-life ecosystem processes like litter decomposition. This influence occurs through plant traits and decomposer activity. In turn, litter decomposition feedbacks on plant-community assembly. Consequently, litter decomposition does not seem to be a collateral consequence of plant traits, but rather an important part of their ecological strategies and biotic interactions, participating to a feedback loop involving community assembly processes.

En aquesta tesi presento quatre capítols, en els quals es discuteix com diferents espècies subarbustives interactuen amb plantes coexistents, sota règims variables d’estrès. Aquesta recerca ha estat centrada en ecosistemes de gran valor ambiental, ecològic i de conservació, i alhora sensibles als canvis ambientals, en quatre regions distintes situades en zones de latitud o altitud elevades. Per ordre latitudinal, els ecosistemes estudiats han estat: 1) el límit supraforestal dels Pirineus Centrals, situats en una zona temperada; 2) el gradient de successió primària d’un ecosistema situat en una zona boreal, a Finlàndia,; 3) el límit supraforestal situat a la zona subàrtica de Lapònia; i 4) la tundra situada en una zona de l’alt àrtic, al nord‐est de Grenlàndia . Els dos primers capítols es basen en una aproximació experimental i se centren en investigar com els subarbustos més comuns trobats prop del límit supraforestal interactuen amb plançons d’arbres de les espècies que formen el límit del bosc en dos ambients contrastats, i.e. el límit supraforestal subàrtic, i el límit supraforestal dels Pirineus Centrals. A més, donat que els arbres que viuen prop del límit de distribució són molt sensibles als canvis ambientals , especialment a l’augment de temperatures durant l’estació de creixement (Körner 2003), també s’ha investigat com podrien afectar el creixement dels plançons d’arbre i la seva supervivència al llarg de l’ecotò, diferents escenaris de canvis ambientals. El primer capítol es titula ‘Les interaccions arbre‐arbust i els canvis ambientals dirigeixen la dinàmica supraforestal a la zona subàrtica’, i s’hi exposa l’experiment de tres anys de durada que es va executar al ‘Parc Nacional d’Abisko’, en els Scandes subàrtics, al nord de Suècia; les plantes estudiades varen ser plançons de Betula pubescens i el subarbust Vaccinium myrtillus. En aquest estudi es va certificar la importància de les interaccions arbust‐arbre, tant facilitatives com competitives, com a elements clau en la dinàmica supraforestal subàrtica. A més, vàrem demostrar que la gran sensibilitat dels plançons a l’escalfament tenia fortes implicacions per la dinàmica supraforestal, tenint en compte l’escenari previst d’augment de temperatures en latituds elevades. També es va observar que les interaccions complexes entre arbusts i herbívors són claus per preveure canvis futurs. El segon capítol es titula ‘Els plançons d’arbres situats als límits supraforestals Pirinencs i subàrtics mostren respostes semblants a la presència d'arbustos i a les simulacions de canvis ambientals’. Aquí es presenta una comparació entre els resultats de l’experiment presentat en el primer capítol i els obtinguts en un experiment paral∙lel dut a terme durant un període de temps semblant prop del límit supraforestal en el ‘Parc Natural de l’Alt Pirineu’, als Pirineus Centrals catalans. Aquest experiment es va basar en el mateix disseny factorial que l’anterior estudi, però amb diferents espècies (i.e. plançons de l’arbre Pinus uncinata i l’arbust Rhododendron ferrugineum). Segons ens consta, és el primer estudi que avalua experimentalment les respostes de les plantes vers diferents escenaris ambientals en un ecosistema d’alta muntanya als Pirineus. En aquest capítol es presenten alguns mecanismes que ajudaran a comprendre la variabilitat recentment observada de les respostes locals de límits supraforestals de zones temperades i subàrtiques com a resultat del clima canviant, i també identifiquem alguns paral∙lelismes que poden utilitzar‐se per generalitzar les respostes a gran escala dels límits supraforestals al canvi climàtic. El tercer capítol se centra en els efectes d’un subarbust dominant (i.e. Empetrum nigrum) en plançons de Pinus sylvestris al llarg del gradient de successió primària en un ecosistema boreal en una illa emergent de la badia de Bòtnia, a Finlàndia. Aquest capítol s’anomena ‘Un arbust ericoide exerceix la doble funció de reclutar pins i els seus simbionts fúngics al llarg d’un gradient de successió primària’. Aquí hem mostrat que els efectes facilitadors i competidors dels subarbustos determinen fortament l’establiment de plançons i la seva colonització fúngica al llarg d’aquest gradient de successió. Segons ens consta, aquests són els primers resultats que demostren que un arbust ericoide micorrízic pot millorar tant el desenvolupament de l’arbre hoste ectomicorrízic com el dels simbionts fúngics de l’arbre. L’estudi presentat al quart capítol es va realitzar al llarg d’un gradient de nivositat en un ecosistema extrem de tundra àrtica al ‘Parc Nacional del nord‐est de Grenlàndia’, el Parc Nacional més gran del món. El capítol es titula ‘Interaccions vegetals i composició de la vegetació àrtica al llarg d’un gradient de nivositat al nord‐est de Grenlàndia’. Aquest ecosistema és probablement el més sensible i fràgil de tots els ecosistemes estudiats en aquesta tesi, donat que s’espera que a la costa est de Grenlàndia s’hi esdevinguin canvis substancials en el clima com a resultat de canvis destacables en els règims de precipitació de neu i de les temperatures (Brown i Mote 2009). Aquí es va avaluar la riquesa d’espècies de plantes, així com els patrons d’establiment i composició de diverses formes de creixement existents en comunitats vegetals àrtiques associades a una variació de la cobertura de neu durant els mesos d’hivern. Aquest estudi ajudarà a preveure la diversitat potencial i els canvis en la vegetació a la zona de l’alt Àrtic, si els règims de precipitació de neu canvien en el futur com es preveu. / In this thesis I present four chapters, and in all of them I discuss how dwarf shrubs interact with co‐occurring plants under varying regimes of stress. This research involved ecosystems of great environmental, nature conservation and ecological value, yet highly sensitive to environmental changes, in four contrasting cold regions at high altitude or high latitude. Following a latitudinal order, the selected ecosystems were: 1) a temperate alpine treeline in the Central Pyrenees; 2) a primary succession gradient in a boreal ecosystem in Finland; 3) a subarctic alpine treeline in Lapland; and 4) a high‐arctic tundra in north‐eastern Greenland. The first two chapters are based on an experimental approach and focus on how shrubs commonly found near the treeline interact with tree seedlings of treelineforming species in two contrasting environments, i.e. in a subarctic forest‐tundra ecotone in Lapland, northern Sweden, and in a more southern, temperate forestalpine pasture ecotone in the Central Pyrenees. In addition, since trees living near their limit of distribution are very sensitive to environmental changes, especially to increased temperature during the growing season (Körner 2003), we also assessed how distinct environmental change scenarios may affect tree seedling growth and survival across the ecotone. The first chapter is entitled ‘Shrub‐tree interactions and environmental changes drive treeline dynamics in the Subarctic’, where we explain the three‐year‐long experiment performed in the Abisko National Park, in the subarctic Scandes, Northern Sweden; the species studied were Betula pubescens tree seedlings and the shrub Vaccinium myrtillus. In this study we showed the importance of facilitative and competitive shrub‐tree interactions as drivers of subarctic treeline dynamics. Furthermore, we demonstrated that the great sensitivity of tree seedlings to warming had strong implications for treeline dynamics under the predicted warmer scenario at high latitudes, and we identified that complex interactions between shrubs and herbivores are critical to predicting future changes. The second chapter is entitled ‘Similar tree seedling responses to shrubs and to simulated environmental changes at Pyrenean and subarctic treelines’. Here we presented a comparison between the results obtained in the experiment presented in the first chapter and those obtained in a parallel experiment performed during a similar period near the treeline in the Alt Pirineu Natural Park, in the Central Pyrenees, Catalonia. This experiment was based on the same factorial design but with different species (i.e. Pinus uncinata tree seedlings and the shrub Rhododendron ferrugineum). To our knowledge, it is the first study which experimentally tests the responses of plants to distinct environmental scenarios in a high mountain ecosystem in the Pyrenees. In this chapter we presented some mechanisms for understanding the recently observed variability of local responses of both subarctic and alpine treelines to currently changing climate while identifying some commonalities that can be used to generalise large scale response of treelines to climate warming. The third chapter focuses on the effects of a dominant dwarf shrub (i.e. Empetrum nigrum) on Pinus sylvestris tree seedlings along a primary succession within a boreal ecosystem on an uplifting island in Bothnian Bay, Finland. This chapter is called ‘An ericoid shrub plays a dual role in recruiting both pines and their fungal symbionts along primary succession gradients’. Here we showed that facilitative and competitive effects of shrubs markedly determined tree seedling establishment and their fungal colonisation along this succession gradient, but in this chapter we did not relate these findings to any environmental changes. As far as we know, we presented the first finding that an ericoid mycorrhizal shrub may enhance both the performance of the ectomycorrhizal host tree and the tree’s fungal symbionts. The study presented in the fourth chapter was performed along a snow‐depth gradient in an extreme arctic tundra ecosystem in the Northeast Greenland National Park, the largest national park in the world. The chapter is entitled ‘Plant interactions and higharctic vegetation composition along a snow‐depth gradient in NE Greenland’. This ecosystem is probably the most sensitive and fragile among the ecosystems studied in this thesis as the eastern coast of Greenland is expected to experience substantial changes in climate due to marked changes in snow precipitation and temperature regimes (Brown and Mote 2009). Here we assessed plant species richness, establishment and composition patterns in distinct growth forms occurring in common arctic plant communities associated with varying snow‐depth during the winter season. This study will help to predict potential diversity and vegetation changes in the high Arctic if snow precipitation regime changes in the future as anticipated.

In recent decades, it was shown that belowground competition for some plants may take form of the tragedy of the commons (TOC). In these plants, the competing neighbours invest more in root systems than would be appropriate for optimal nutrient uptake for the group and also more than they do when grown alone. However, there is also strong evidence that other species do not follow TOC, and tailor their root system to best nutrient exploitation irrespectively of competitor presence. The root investment strategy of these plants should correspond to the ideal free distribution (IFD). In my thesis I focus on two aspects:  I use game theoretical models to explore, whether those strategies can coexist within species and also whether different species can coexist if they have different strategy. From this model I draw predictions, which I test by meta-analysis.  Using Agrostis stolonifera as a model, I test assumptions on nutrient and neighbour perception, which underlie TOC and IFD models. I show that according to mathematical models, those two strategies can coexist in different species in a community, but cannot coexist within a species. Within a species, the TOC strategy should always dominate, once it appears. This can be extrapolated to macroevolutionary scale - once TOC occurs in certain clade, it should...