The concentration of atmospheric carbon dioxide (CO2) in the atmosphere has increased by 35% since pre-industrial levels. Projections for the next 100 years indicate an increase to levels between 490 and 1260 parts per million by volume (ppm) of CO2, equating to a 75 % to 350 % increase in concentration since the year 1750. Associated with this increase in [CO2] will be a 1.4 to 5.8?? C increase in lower atmospheric temperature. While past research has attempted to address the effects of such climatic changes on individual plant responses, predictions of plant responses at the ecosystem level are still highly uncertain. Difficulties lie in the enormous variation of plant responses to climate change variables among and within species, and between and within environmental conditions. Past research assumed that plants using either the C3 or C4 metabolic pathways would respond differently but predictably to climate-change variables based on their metabolic pathway. Recent evidence has suggested however, that the added interactions of external environmental variables and species-specific sensitivities to climate change make it difficult to predict plant and ecosystem responses to climate change. To investigate the mechanisms behind responses of Australian grasses to climate change, 2 pot experiments was conducted using growth cabinets to compare the effect of elevated CO2 and water-limitation on the invasive C4 grassland plant, Eragrostis curvula (E. curvula), native Australian C3 grassland plant, Austrodanthonia racemosa (A. racemosa), and wheat species, Triticum aestivum (T. aestivum). The experiment was run at ambient levels of CO2 maintained at 390 ppm compared to elevated levels of 740 ppm. Imposed restrictions to water supply consisted of gradually drying the soil down to 30 % available soil water (ASW) followed by re-wetting to 50 % ASW. Well-watered conditions for the experiment consisted of gradually drying the soil down to 50 % ASW, followed by rewetting to 95 % ASW. Plants were grown in mixtures and monocultures, consisting of 9 plants equally spaced in a grid design. The three significant findings of the thesis were that: 1) the metabolic pathway (C3 versus C4) was not always an accurate predictor of biomass accumulation under elevated CO2 in the plants studied. Previous research suggested that CO2-stimulation of photosynthesis in C3 plants would lead to greater increases in biomass under elevated CO2 compared to C4 plants, though both C3 and C4 plants could benefit from any reduction in stomatal conductance under dry conditions at elevated CO2. The results from the experiments in this thesis showed a strongly significant biomass response to elevated CO2 in both dry and wet conditions for C4 grass E. curvula. The C3 grass A. racemosa in dry conditions, did not. It was speculated that without the CO2-induced water conservation effect, the C3 grass experienced photosynthetic down-regulation and this precluded a positive biomass response under elevated CO2. 2) the magnitude and direction of biomass response to elevated CO2 was dependant on factors such as resource-availability and the phenotypic variability of the plants species. 3) critical analysis of results from this thesis, combined with past research on plant responses under elevated CO2 showed a tendency for researchers to repeatedly test plants from the Poaceae family, or close relatives of the Poaceae family. As a result, when past data were corrected for this lack of independence, there was no relationship between the evolution of the C3 and C4 metabolic pathway and biomass response to elevated CO2. Instead, other factors (such as growth rate, plant height, leaf number, etc) were presented as being more important in determining biomass response. These observations were supported by results found in this thesis.

The concentration of atmospheric carbon dioxide (CO2) in the atmosphere has increased by 35% since pre-industrial levels. Projections for the next 100 years indicate an increase to levels between 490 and 1260 parts per million by volume (ppm) of CO2, equating to a 75 % to 350 % increase in concentration since the year 1750. Associated with this increase in [CO2] will be a 1.4 to 5.8?? C increase in lower atmospheric temperature. While past research has attempted to address the effects of such climatic changes on individual plant responses, predictions of plant responses at the ecosystem level are still highly uncertain. Difficulties lie in the enormous variation of plant responses to climate change variables among and within species, and between and within environmental conditions. Past research assumed that plants using either the C3 or C4 metabolic pathways would respond differently but predictably to climate-change variables based on their metabolic pathway. Recent evidence has suggested however, that the added interactions of external environmental variables and species-specific sensitivities to climate change make it difficult to predict plant and ecosystem responses to climate change. To investigate the mechanisms behind responses of Australian grasses to climate change, 2 pot experiments was conducted using growth cabinets to compare the effect of elevated CO2 and water-limitation on the invasive C4 grassland plant, Eragrostis curvula (E. curvula), native Australian C3 grassland plant, Austrodanthonia racemosa (A. racemosa), and wheat species, Triticum aestivum (T. aestivum). The experiment was run at ambient levels of CO2 maintained at 390 ppm compared to elevated levels of 740 ppm. Imposed restrictions to water supply consisted of gradually drying the soil down to 30 % available soil water (ASW) followed by re-wetting to 50 % ASW. Well-watered conditions for the experiment consisted of gradually drying the soil down to 50 % ASW, followed by rewetting to 95 % ASW. Plants were grown in mixtures and monocultures, consisting of 9 plants equally spaced in a grid design. The three significant findings of the thesis were that: 1) the metabolic pathway (C3 versus C4) was not always an accurate predictor of biomass accumulation under elevated CO2 in the plants studied. Previous research suggested that CO2-stimulation of photosynthesis in C3 plants would lead to greater increases in biomass under elevated CO2 compared to C4 plants, though both C3 and C4 plants could benefit from any reduction in stomatal conductance under dry conditions at elevated CO2. The results from the experiments in this thesis showed a strongly significant biomass response to elevated CO2 in both dry and wet conditions for C4 grass E. curvula. The C3 grass A. racemosa in dry conditions, did not. It was speculated that without the CO2-induced water conservation effect, the C3 grass experienced photosynthetic down-regulation and this precluded a positive biomass response under elevated CO2. 2) the magnitude and direction of biomass response to elevated CO2 was dependant on factors such as resource-availability and the phenotypic variability of the plants species. 3) critical analysis of results from this thesis, combined with past research on plant responses under elevated CO2 showed a tendency for researchers to repeatedly test plants from the Poaceae family, or close relatives of the Poaceae family. As a result, when past data were corrected for this lack of independence, there was no relationship between the evolution of the C3 and C4 metabolic pathway and biomass response to elevated CO2. Instead, other factors (such as growth rate, plant height, leaf number, etc) were presented as being more important in determining biomass response. These observations were supported by results found in this thesis.

La croissance secondaire est à l'origine de l'accumulation de biomasse pérenne par les arbres et détermine en partie la capacité des écosystèmes forestiers à stocker du carbone. Cependant, les contraintes environnementales sur la croissance en milieu méditerranéen sont encore mal décrites et nous ne savons pas comment les changements climatiques futurs vont les modifier. L'objectif de la thèse est de déterminer, principalement à partir de l'étude de l'allocation du carbone à la croissance secondaire, les réponses fonctionnelles saisonnières et interannuelles du chêne vert (Quercus ilex L.) aux variations climatiques en région méditerranéenne. L'utilisation de différentes approches expérimentales, à des échelles spatiales allant du cerne à l'écosystème et à des échelles temporelles allant de la journée à plusieurs dizaines d'années, a permis de mettre en évidence l'effet de différentes contraintes environnementales (disponibilité en eau, réchauffement de la température, et densité du peuplement) sur la croissance secondaire et la composition isotopique du cerne. L'étude de la phénologie de la croissance montre que celle-ci est contrôlée directement par les températures hivernales et le déficit hydrique, plus que par la disponibilité en éléments carbonés issus de la photosynthèse. De 1968 à 2013, les changements climatiques ont entrainé une contrainte hydrique de plus en plus précoce qui s'est trouvée compensée par un début de croissance initié plus tôt dans l'année, sous l'effet du réchauffement des températures hivernales, et une meilleure efficacité d'utilisation de l'eau, sous l'effet de l'augmentation de la concentration en CO2 atmosphérique. La réduction de la mortalité et l'augmentation de la croissance observée dans des parcelles éclaircies montre que cette pratique sylvicole permet de préparer les taillis de chêne vert à l'intensification de la sècheresse prévue pour la région méditerranéenne. / Tree secondary growth is responsible for woody biomass accumulation and is a major component of carbon storage in forest ecosystems. Environmental constraints on secondary growth in Mediterranean ecosystems must, however, be described in more to details to better understand how they will be modified by climate change. This dissertation aims at studying the functional responses of Mediterranean holm oak (Quercus ilex) to seasonal and inter-annual climate variations through the study of carbon allocation to secondary growth. Different experimental approaches, at spatial scales ranging from tree rings to the ecosystem and at temporal scales from the day to several decades, were used to identify the main environmental constraints (water availability, temperature warming, competition) to secondary growth and carbon isotopic composition of tree rings. The phenology of stem growth shows evidence for a direct environmental control on annual growth by winter temperature and summer drought that is more limiting than the carbon supply from photosynthesis. Climate change from 1968 to 2013 resulted in earlier water limitation on secondary growth, which was compensated by earlier growth onset, due to warmer winter temperature, and higher water use efficiency, due to increased atmospheric CO2 concentration. Thinning reduced tree mortality and increased stem growth, so thinning management in old holm oak coppices could prepare the ecosystem to better withstand the increasing drought forecasted for the Mediterranean region.

Le contrôle biologique (C. -B. - l'utilisation d'ennemis naturels pour lutter contre les ravageurs) est durable, écologique et rentable pour contrer la résistance des ravageurs en augmentant l'utilisation des pesticides. Les parasitoïdes des pucerons sont des ennemis naturels communs des pucerons, les principaux ravageurs mondiaux dans l'agriculture. L'étude de la spécificité de l'hôte parasitoïde contribue à (1) comprendre les mécanismes écologiques et évolutifs de l'écosystème et (2) évaluer l'efficacité des agents de lutte biologique et les risques écologiques pour les espèces non ciblées. Cette étude porte sur la spécificité de l'hôte fondamental des parasitoïdes sur les niveaux individuels, en matière de besoins en ressources et dans le contexte des interactions multi trophiques sous stress abiotique environnemental, c'est-à-dire la limitation de l'eau. Aphidius ervi (Hymenoptera: Braconidés: Aphidiinae) a été choisi; ce parasitoïde puceron est largement utilisé comme modèle écologique et comme agent de lutte biologique commercial (BCA). D'une part, l'indice de spécificité de l'hôte A. ervi a été mesuré sur une large gamme d'espèces de pucerons. D'autre part, les impacts indirects de la limitation de l'eau ont été étudiés sur la spécificité de l'hôte du parasitoïde. En outre, les modifications induites par le stress hydrique dans la plante et les traits de vie des pucerons ont été mesurés. A. ervi s'est avéré être une espèce intermédiaire spécialisée qui a attaqué toutes les espèces de pucerons à des taux élevés, mais n'a pas pu se développer correctement sur toutes les espèces. Les quelques espèces qui se développaient bien étaient phylogénétiquement proches et appartenaient à la tribu des Macrosiphini. En outre, une corrélation positive de préférence - performance a été trouvée. Sous stress hydrique, la préférence et la performance des parasitoïdes ont été affectées, causant la perte de la corrélation. La limitation de l'eau a modifié négativement la qualité nutritionnelle de la plante, ce qui a entraîné une faible performance des pucerons sur les plantes hôtes. Ceci à son tour a diminué la convenance des hôtes pucerons pour le parasitoïde. Les effets de la limitation de l'eau n'étaient pas similaires pour toutes les combinaisons plantes-pucerons et dépendaient de plusieurs facteurs, à savoir les mécanismes végétaux adaptés au stress et la spécialisation de l'hôte des pucerons et des parasitoïdes. / Biological control (BC - the use of natural enemies to control pests) are sustainable, environmentally friendly and cost-effective methods to counteract pest resistance by increasing pesticide use. Aphid parasitoids are common natural enemies of aphids, the major worldwide pests in agriculture. The study of parasitoid host specificity contributes to (1) understanding ecological and evolutionary mechanisms driving the ecosystem and (2) evaluating the efficiency of biocontrol agents and the ecological risks for non-target species. This study focuses on the parasitoids fundamental host specificity on individual levels, in terms of resource requirements and in the context of multi-trophic interactions under environmental abiotic stress, i.e.water limitation. Aphidius ervi (Hymenoptera: Braconidae: Aphidiinae) was chosen; this aphid parasitoid is used widely as an ecological model and commercial biological control agent (BCA). On the one hand, A. ervi host specificity index was measured on a broad range of aphid species. On the other hand, the indirect impacts of water limitation were investigated on the host specificity of the parasitoid. Furthermore, water stress-induced modifications in the plant and the aphid life-history traits were measured. A. ervi was shown to be an intermediate specialist species who attacked all aphid species at high rates but was unable to develop well on all of them. The few that developed well were phylogenetically close and belong to the Macrosiphini tribe. Interestingly, a positive correlation preference – performance was found. Under water stress, both preference and performance of parasitoids were affected causing loss of the correlation. Water limitation negatively altered the plant nutritional quality resulting in low aphid performance on host plants. This in turn decreased the suitability of aphid hosts for the parasitoid. The impacts of water limitation were not similar across all plant-aphid combinations and depended on several factors, namely stress-adapted plant mechanisms and the host specialization of both aphids and parasitoids.

<p>Earth’s ecosystems emerge from interconnected biosphere, geosphere, and atmosphere processes. Changes to any one process ripple through the Earth system, affecting other processes. As global climate change continues, nitrogen deposition is anticipated to increase and precipitation is expected to have varied changes across the globe. These changes to the atmosphere and geosphere will have implications for the biosphere. Namely, vegetation will be impacted by changes to nutrient and precipitation regimes. Vegetation comprises the aggregate strategies of individual plants, which are also influenced by changes in nutrient and water availability. The responses of individual plants to nitrogen, water, and uncertainty are the main focus of this dissertation, as understanding those will be critical to scaling up to the aggregate.</p> <p> First, I describe a mathematical model that predicts grassland root and shoot biomass across carbon, nitrogen, and water gradients. The model simulates competition among plants by dynamically allocating carbon to either root or shoot growth depending on the growth strategy employed by the other plant. I show that the model accurately predicts root net primary productivity (NPP), but performs poorly for shoot and total NPP. At the biome scale, modeled NPP does not vary with water alone but rather water and nitrogen interact to influence NPP. Second, I conduct a greenhouse experiment using <em>Eragrostis capillaris</em> (L.) Nees to examine the predictions of the model mentioned above to answer the question: how do water and nitrogen affect fitness and biomass allocation in a drought-tolerant C4 grass? And ask: what is the nature of the relationship between water and nitrogen as resources? I show that water was important for increasing shoot and total biomass, but that root biomass and root:shoot ratio was influenced interactively by water and nitrogen as predicted by the model. I conclude that the nature of the relationship between water and nitrogen was that of either interacting or hemi-essential resources. That is, additional water was able to partially substitute for limited nitrogen to maintain biomass. Third, I explore how information theory can apply to plants that face uncertainty in resource availability and briefly review the types and sources of information and the mechanisms that plants use to perceive and respond to their environment. Overall, my framework posits that plants interpret information from their surroundings as an emergent property of distributed information processed by a network of cells. I end with a prospectus of directions for future research, including decoding signal from noise, storage of information, strategies to cope with information entropy, additional means of information transmission, and two-way information signaling with biotic partners. Finally, I use the information theory framework discussed above to answer the questions: can plants sense and respond to information entropy? I explore this question using data from an experiment which altered the temporal supply of nutrients and found no support that <em>P. sativum</em> can sense and respond to entropy. Understanding the relationships of water, nitrogen, and uncertainty is critical to predicting plant growth, especially as climate change continues to influence the global system.</p>