Abstract Plants have evolved various adaptive strategies for balancing the benefits and costs of having a high affinity for resources, plasticity of growth allocation and mycorrhizal symbiosis. The relative growth rates of mycorrhizal and non-mycorrhizal plants were modelled for stable and variable nutrient availability. Mycorrhizal plants had higher growth rates at low and non-mycorrhizal plants at high nutrient availability. Variation in nutrient availability reduced the growth rate of mycorrhizal plants due to a high affinity for nutrients. However, mycorrhizal plants may be able to buffer against external fluctuations and therefore experience less environmental variation than non-mycorrhizal plants. Non-mycorrhizal plants may even benefit from variation. The optimal allocation of growth between shoot and roots depends on the availability of energy and nutrients. The optimisation model predicted that the requirement for phenotypic plasticity of shoot/root allocation is greatest in environments with low resource availability. Plants with a high affinity for resources required more plasticity in order to tolerate variation than plants with a low affinity. The model predicted a trade-off between the ability to deplete resources and the ability to tolerate resource fluctuations. Changes in the availability and ratio of resources lead to changes in the structure and composition of vegetation during primary succession. The field study of the forested phases of the land uplift island Hailuoto showed a successional change in the vegetation from the dominance of bryophytes and deciduous dwarf shrubs to dominance by lichens and evergreen dwarf shrubs. The humus layer became thinner and the availability of nutrients declined, while the C/N ratio of soil organic matter increased during succession indicating a decline in the quality of organic matter. The increased soil respiration rate indicates a successional increase in the energetic costs of decomposing organic matter. Nutrients mediate both direct and indirect trophic interactions. Indirect interactions of nutrient cycling are not explicit in continuous time models. A transformation to a discrete time model was shown to make the indirect interactions explicit as transition probabilities and allowed their dynamic contribution to be evaluated with an elasticity analysis. The importance of indirect interactions was greater in tundra than temperate forest and increased with the rate of nutrient cycling.

No description available.

Understanding the mechanisms that control the rate and trajectory of primary succession can lead to insights for ecosystem rehabilitation. Proposed mechanisms include life history traits and nutrient limitation. To explore how these mechanisms can drive successional dynamics, I devised a stoichiometric ecosystem-level model that considered the role of nitrogen and phosphorus limitation in plant primary succession in conjunction with life history traits. This model was applied to the plant community on Mount St. Helens to check the validity of the mechanisms. The results show the competitive hierarchy of plants at the local scale can be explained by nutrient limitation and plant stoichiometry. At regional scales, life history traits interact with local processes to shape community structure and successional dynamics. At all scales, the presence of Lupinus lepidus, a nitrogen-fixer, significantly altered community dynamics and succession. This study suggests that primary succession can be examined within the framework of ecological stoichiometry. / Ecology

Natural abundance of 15N varies greatly and unpredictably within and between environments. The unpredictable nature of 15N limits the use of N isotope natural abundance (d15N) in tracing the flow and fate of N in environments. Recent investigations have, however, revealed consistent and repeatable patterns of 15N in some ecosystem components. These patterns suggest that d15N may yet provide a tool to investigate and illuminate ecosystem N cycling processes. Identifying and quantifying the sources of isotopic variation must precede any significant advance in the application of this technique, and to this end an assessment of isotopic variation associated with major ecosystem components has been carried out in this thesis. d15N patterns have been established, hypotheses proposed and tested, and conclusions about the application of the technique are presented. 15N patterns in surface and groundwater were measured in a variety of different land-use catchments in an attempt to identify distinct isotopic 'fingerprints'. High levels of 15N variation were measured in both stream and groundwaters, resulting in strongly overlapping land-use 'fingerprints'. Environmental 15N variation in streams and groundwaters was found to be too great to differentiate between land-uses based on d15N alone. In contrast, the artificially 15N enriched signature of effluent N was used to trace its flow and fate, following irrigation, in a forested catchment. The effluent d15N signature allowed it to be traced into the major ecosystem components, permitting a first order N budget to be determined for effluent N storage and loss. N sources with significantly different 15N signatures to that of 'background ecosystem N' can therefore be used to trace the flow and fate of N in ecosystems. During the course of this work a number of higher and lower order plants were observed to have highly depleted (lt; -8 ) d15N signatures. Epiphytes and lithophytes, strongly reliant on atmospheric N sources, were consistently depleted in 15N, with signatures as low as -24 , measured in a range of environments. A similar level of depletion was measured in a wide range of plants growing in early primary succession sites (as low as -22.3 ), which could not be accounted for by any abiotic or biotic factor or significantly depleted N source. The absence of any measurable driver of depletion suggested a universal fractionating mechanism which acts in a wide range of environments and vegetation types. Diffusive uptake of atmospheric NH3(g) and the proportional uptake of a supplied N source were two proposed mechanisms that could theoretically account for the level and universal nature of depletion. Diffusive uptake of atmospheric NH3(g) was tested as a primary fractionating mechanism in plants. Strongly N deficient plants were capable of utilising NH3(g) as a nutritional source, but the level of 15N depletion measured in these plants closely approximated that of the inherent NH3(g) d15N signature. No significant additional fractionation is associated with NH3(g) diffusive uptake. Diffusive uptake of atmospheric NH3(g) by plants cannot alone account for the level of depletion measured in early primary succession plant communities. Proportional uptake of a N source as a primary fractionating mechanism was tested by growing plants in various concentrations and rates of applied N. Fractionation attributed to the proportional uptake of a supplied N source, as a consequence of P limitation or rapid flow over roots, resulted in a significant level of 15N depletion in plants. The level of depletion attributed to this mechanism was, however, not sufficient to account for the level measured in early primary succession plant communities. Individual 15N fractionating mechanisms cannot alone explain the level of depletion observed in early primary succession plants, however a combination of fractionating mechanisms can. Fractionation attributed to the proportional uptake of an already depleted N source, i.e., wet deposited N, largely accounts for the level of depletion measured in early succession plant communities. This two-step fractionation model can act on both higher and lower plants, independent of ecosystem biotic and abiotic factors. Additional, and less dramatic fractionations attributed to atmospheric NH3(g) uptake, mycorrhizal associations, internal remobilisation, and taxon-specific N acquisition strategies, will contribute to the level of d15N depletion. This thesis presents the first extensive survey of highly depleted d15N signatures in terrestrial vegetation. Furthermore, thorough testing of theoretically plausible mechanisms has resulted in a full account of the highly depleted d15N signatures measured in a wide range of vegetation types and environments.

Despite the wealth of research into strategies for microbiome modulation, studies of microbiome management in pig hosts have found mixed results. A refined understanding of the patterns of microbiome assembly during the host’s early life, when management strategies are most commonly applied, is necessary for the development of successful management practices. Here, we study the development of the pig gut microbial community in a monitoring experiment, sampling the microbiome of pigs in a commercial farm intensively during the first month of life. We found that the community’s taxonomic richness increased linearly with host age. Furthermore, rapid changes across communities occurred in stages, and non-linear patterns in relative abundance were commonly observed among dominant taxa across host age, consistent with primary succession. Our results highlight the importance of understanding the patterns of microbiome assembly during host development, and identify successional stages as windows of opportunity for future research.

Sand Pits have great potential to restore via spontaneous succession. There are not many studies of the development of soils in sand pits; therefore, this work focuses on the development of soil properties in sand pits comparing technically reclaimed sites and sites with primary succession. 120 samples were collected at fourteen sand pits in South Bohemia and selected soil properties were assessed. The surveyed plot were grouped according to age, and the successional and technically reclaimed sites were distinguished. The results were evaluated by analysis of variation and regression analysis. The differences in soil bulk density, pH, thickness of soil organic horizon, Cox, N and C/N ratio between reclaimed and spontaneously restored sites of different age were analysed. The bulk density and pH decrease significantly on successional sites; however, the reclaimed sites do not show signifiant differences. Organic horizon thickness increases with age on both types of sites.Cox, N and C/N ratio do not show significant influence age. Lower values of Cox, N and C/N were measured in initial stages of succession. On the reclaimed sites there is almost no change in values of Cox, N and C/N ratio, because of the occurrence of nutrient-richer substráte used during technical reclamation. Keywords: soil...

Les plantes, comme tous les êtres vivants, entretiennent un rapport double à leur environnement. L’environnement sélectionne quelles stratégies peuvent s’établir, et les stratégies ainsi sélectionnées façonnent en retour cet environnement. Cette boucle de rétroaction environnementale, lorsqu’elle est alimentée par une variabilité de formes, est le moteur de l’évolution, de l’assemblage des communautés et du développement écosystémique, et détermine en fin de compte les propriétés émergentes des écosystèmes.Les approches issues de l’écologie théorique reconnaissent depuis longtemps cette dualité, comme en témoignent les concepts de "niche de besoin" et "niche d’impact" au cœur de la théorie contemporaine de la niche. Similairement, les approches type « théorie des jeux » comme la dynamique adaptative reconnaissent le rôle central joué par la boucle de rétroaction environnementale en tant que moteur des dynamiques éco-évolutives.Dans cette thèse, j'unifie ces deux perspectives théoriques et les applique à des problèmes écologiques variés, dans le but de comprendre comment les interactions réciproques entre les plantes et leur environnement déterminent les traits adaptatifs des plantes et les propriétés émergentes des écosystèmes.Dans un premier temps, je propose un cadre mathématique général et rigoureux à la théorie contemporaine de la niche et la méthode graphique qui lui est associée. Après avoir étendu ce cadre à la prise en compte d’un continuum de stratégies en interaction à l’aide d’enveloppes géométriques, je montre comment appliquer la théorie contemporaine de la niche à deux perspectives, à savoir les dynamiques éco-évolutives et l’assemblage de communautés par remplacements successifs de stratégies.Dans un second temps, j’applique cette approche à l’étude de l’évolution des défenses des plantes contre les herbivores le long de gradients de nutriments, en considérant l’évolution des traits d’acquisition de la ressource, de tolérance et de résistance aux herbivores. Je montre que la prise en compte des transferts trophiques conduit à la sélection de stratégies compétitives mais sans défense dans les environnements pauvres, alors que ce sont toujours des stratégies défendues (résistantes, tolérantes, ou la coexistence des deux) qui dominent dans les environnements riches en nutriments. Mes résultats mettent en évidence le rôle central joué par la rétroaction plante-herbivores dans la détermination des patrons de défense des plantes.Dans un troisième temps, je montre comment la théorie contemporaine de la niche peut être étendue pour prendre en compte la facilitation. J’utilise ensuite cette approche pour montrer comment la colonisation d’un substrat nu par une communauté de plantes fixatrices d’azote couplée au recyclage des nutriments peut donner naissance à de la succession par facilitation. Contrairement aux modèles habituels de succession, je montre que la succession par facilitation donne lieu à un développement autogène de l’écosystème ainsi qu’un régime de bistabilité entre la végétation et le substrat nu en fin de succession. Enfin, je propose une nouvelle théorie de la succession basée sur les ratios de ressources.Pris dans leur ensemble, ces nouveaux développements démontrent que la théorie de la niche peut être adaptée à l’étude d’un large champ de situations écologiques, de la facilitation aux dynamiques éco-évolutives et à l’assemblage des communautés. Dans ce cadre conceptuel, mon approche basée sur les enveloppes s’avère être un outil efficace pour passer de l’échelle individuelle à l’échelle de l’écosystème, en assimilant le remplacement adaptatif d’espèces à une plasticité des propriétés écosystémiques. Cette approche permet alors de décrire l’émergence des boucles de régulation qui contrôlent le fonctionnement des écosystèmes, comme l’illustrent mes résultats le long de gradients de nutriments sur la transition entre régimes de succession ou encore l’émergence de culs-de-sac trophiques. / As living organisms, plants present a dual relationship with their biotic and abiotic environment. The environment selects plant strategies that can establish, and selected strategies in turn impact and shape the environment as they spread. When fueled by variation ,this environmental feedback loop drives evolution, community assembly and ecosystem development, and eventually determines the emergent properties of ecosystems.Theoretical ecology approaches have long recognized this duality, as it is at the core of contemporary niche theory through the concepts of requirement and impact niche. Similarly, game-theoretical approaches such as adaptive dynamics have emphasized the role played by the environmental feedback loop in driving eco-evolutionary dynamics. However, niche theory could benefit from a more individualistic, selection based perspective, while adaptive dynamics could benefit from niche theory’s duality and graphical approach.In my dissertation, I unify these theoretical perspectives and apply them to various ecological situations in an attempt to understand how the reciprocal interaction between plants and their environment determines plant adaptive traits and emergent ecosystem functions.First, I introduce a general and rigorous mathematical framework to contemporary niche theory and the associated graphical approach. By extending these ideas to a continuum of interacting strategies using geometrical envelopes, I show how contemporary niche theory enables the study of both eco-evolutionary dynamics and community assembly through species sorting. I show how these two perspectives only differ by the range of invaders considered, from infinitesimally similar mutants to any strategy from the species pool. My results also emphasize the fact that selection only acts on the requirement niche, evolution of the impact niche being just an indirect consequence of the former.Second, I use this approach to study the evolution of plant defenses against herbivores along a nutrient gradient, by considering the joint evolution of resource acquisition, tolerance and resistance to herbivores. I show that trophic transfers lead to the selection of very competitive, undefended strategies in nutrient-poor environments, while defended strategies -- either resistant, tolerant or the coexistence of both -- always dominate in nutrient-rich environments. My results highlight the central, and often underestimated, role played by plant-environment feedbacks in shaping plant defense patterns.Third, I extend contemporary niche theory to facilitation originating from positive environmental feedback loops. I use these new tools to show how colonization of a bare substrate by a community of nitrogen-fixing plants coupled with nutrient recycling can lead to facilitative succession. Contrarily to previous competition-based succession models, I point out that facilitative succession leads to autogenic ecosystem development, relatively ordered trajectories and late succession bistability between the vegetated ecosystem and the bare substrate. By showing how facilitative succession can turn into competition-based succession along an increasing nitrogen gradient, I derive a new resource-ratio theory of succession.Overall, these new theoretical developments demonstrate that niche theory can be adapted to study a broad range of ecological situations, from facilitation to eco-evolutionary dynamics and community assembly. Within this framework, my envelope-based approach provides a powerful tool to scale from the individual level to the ecosystem level, lumping selection-driven species turnover into plastic ecosystem properties. This, is turn, helps describing the emergence at the ecosystem scale of regulation feedback loops that drive ecosystem dynamics and functioning, as exemplified by my results along increasing resource gradients showing a transition from facilitation- to competition-based succession or the emergence of trophic dead-ends.

Diversity of the soil microbial community and its functional aspects were investigated in man-influenced environments, such as colliery spoil heaps in post mining sites and upland pasture used for outdoor cattle husbandry. The study was based on the cultivation of bacteria and streptomycetes as well as culture-independent approaches. Cultivated bacteria and streptomycetes were characterized by phenotypic and genotypic means. The culture-independent approaches were based on an analysis of environmental DNA in terms of both qualitative and quantitative parameters.

Abstract Succession is a compositional change of species and other ecosystem characteristics over time. Mire development, i.e., long-term mire succession is basically driven by an increase in peat layer height, promoting changes in hydrology, vegetation and nutrient status of a particular mire. Due to this, ecosystem processes, such as production and loss of carbon due to decomposition (i.e. carbon gas functions), change with increasing successional mire stage. An adequate method for studying the changes in ecosystem C functions is to measure CO2 and CH4 fluxes between the ecosystem and atmosphere. Succession and carbon dynamics of boreal pristine mires have been much studied. However the link between these phenomena is largely unknown. Further, if and how the C gas functions of mires change during mire succession it is rather poorly understood. The main objective of this thesis was to study how ecosystem functions, measured as CO2 and CH4 exchange, change during mire development. The study also aims to explore the drivers of succession in mire development, i.e., mire succession. Successional mire C dynamics were studied along an eight-kilometer-long successional sequence of primary paludified mires located in the land uplift coast of the Bothnian Bay. Due to the short distance between sites, they all have been under the same climatic control for most of their development. The gradual replacement of plant species with different photosynthetic potential, phenology and assimilating green area resulted in lower-level and temporal variation of CO2 exchange patterns at the later successional stages. Similar to this, CH4 also had the lowest interannual variation in the later stages. In general, CH4 emissions increased with mire age even though this trend did not emerge during the rainy season. Further, this study showed that the wintertime C function pattern was related to the C pattern during the previous summer confirming the important effect of growing season patterns on wintertime C dynamics. In addition to the fundamental effect of vegetation as a driver of succession which was also confirmed in this study, the role of hydrological conditions appeared to be equally important. More constant hydrological conditions at later successional stages resulted in lower temporal variation in CH4 and CO2 fluxes. The present results suggest that the stability of ecosystem C gas functions increases during mire development due to increasing autogenic control. / Tiivistelmä Sukkessio on ekosysteemin lajistossa ja sen muissa ominaisuuksissa ajan kuluessa tapahtuva muutos. Suon kehitystä eli pitkäaikaista suosukkessiota vie eteenpäin turpeen paksuuskasvu, joka saa aikaan muutoksia suoekosysteemin hydrologiassa, kasvillisuudessa ja ravinnetilassa. Tästä johtuen myös suoekosysteemin erilaiset prosessit, kuten tuotanto sekä hajoamisen kautta tapahtuva hiilen vapautuminen eli hiilikaasutoiminta muuttuu suon ikääntyessä. Ekosysteemin hiilikaasutoiminnassa tapahtuvia muutoksia voidaan tutkia muun muassa mittaamalla ekosysteemin ja ilmakehän välisiä hiilidioksidi- ja metaanivirtoja. Boreaalisten luonnontilaisten soiden sukkessiota ja hiilidynamiikkaa on tutkittu runsaasti, mutta niiden välistä yhteyttä ei sen sijaan juuri tunneta. Tämän vuoksi ei tiedetä kuinka soiden hiilikaasutoiminta mahdollisesti muuttuu suon kehityksen aikana eli suosukkession edetessä. Tämän tutkimuksen päätavoitteena oli tutkia kuinka hiilidioksidin ja metaanin vaihdolla mitattu ekosysteemitoiminta muuttuu suon kehityksen aikana. Tutkimus pyrki myös selvittämään suosukkessiota kontrolloivat tekijät. Eri-ikäisten soiden hiilikaasudynamiikkaa tutkittiin mittaamalla hiilikaasuja Perämeren maankohoamisrannikolla kahdeksan kilometrin pituisella sukkessiogradientilla, joka koostuu primaarisoistumisen kautta syntyneistä soista. Soiden lyhyestä keskinäisestä etäisyydestä johtuen ne ovat olleet saman ilmastollisen kontrollin alaisena suurimman osan kehityksestään. Vaiheittainen kasvilajien muutos sukkessiogradientilla yhdessä kasvilajien erilaisen yhteyttämispotentiaalin, fenologian ja yhteyttävän lehtipinta-alan kanssa johti hiilidioksidivaihdon alhaisempaan tasoon sekä pienempään ajalliseen vaihteluun vanhemmilla sukkessiovaiheilla. Myös metaanin vaihdolla oli alhaisimmat vuosien väliset vaihtelut vanhemmilla vaiheilla. Yleisesti ottaen metaanipäästöt kasvoivat suon iän myötä, vaikkakaan tätä trendiä ei havaittu sateisena kasvukautena. Lisäksi tutkimus osoitti, että talviaikaiset hiilivirrat (CO2, CH4) seurasivat kesäaikaisen hiilidynamiikan vaihtelua. Kasvillisuuden keskeinen rooli ekosysteemin sukkessiossa havaittiin myös tässä tutkimuksessa. Kasvillisuuden ohella merkittäväksi suosukkessiota sääteleväksi tekijäksi osoittautui hydrologisten olojen vaikutus. Tasaisemmat hydrologiset olot vanhemmilla sukkessiovaiheilla johtivat vähäisempään ajalliseen vaihteluun metaani- ja hiilidioksidivirroissa. Tutkimuksen tulokset viittaavat siihen, että ekosysteemin hiilidynamiikka stabilisoituu suon kehityksen aikana lisääntyvän autogeenisen kontrollin kautta.