<p>Tolerance to herbivory is the ability of plants to maintain fitness in spite of damage. The goal of this thesis is to investigate the genetic variation and expression of tolerance within species, determine whether and in what conditions tolerance has negative side-effects, and how tolerance is affected by different ecological factors. Tolerance is investigated with special focus on the effects of different damage types, competitive regimes, history of herbivory, and polyploidization in plants. Studies are conducted as a literature review and three experiments on two cruciferous species Raphanus raphanistrum and Cardamine pratensis.</p><p>In the tolerance experiments, plants are subjected to artificial damage solely, or in a combination with natural damage. A literature review was conducted in order to investigate the effects of damage method. We found that traits related to tolerance, such as growth and fitness were not as sensitive in regard to damage method as measures of induced chemical traits, or measures of secondary herbivory.</p><p>Genetic variation of tolerance was demonstrated within populations of R. raphanistrum and between subspecies of C. pratensis. In R. raphanistrum, traits involved in floral display and male fitness were positively associated with plant tolerance to herbivore damage. A potential cost of tolerance was demonstrated as a negative correlation between levels of tolerance in high and low competitive regimes. I found no evidence of other proposed costs of tolerance in terms of highly tolerant plants suffering of reduced fitness in the absence of herbivores or trade-offs in terms of a negative association between tolerance to apical and leaf damage, or between tolerance and competitive ability. In C. pratensis, higher ploidy level in plants involved higher levels of tolerance measured as clonal reproduction. Furthermore, populations exposed to higher levels of herbivory had better tolerance than populations exposed to lower levels of herbivory. In this thesis, I demonstrate evidence of different components for the evolution of tolerance in plants: genotypic variation, selective factors in terms of costs and ploidization, and selective agents in terms of changing environment or herbivore pressure.</p>

Tolerance to herbivory is the ability of plants to maintain fitness in spite of damage. The goal of this thesis is to investigate the genetic variation and expression of tolerance within species, determine whether and in what conditions tolerance has negative side-effects, and how tolerance is affected by different ecological factors. Tolerance is investigated with special focus on the effects of different damage types, competitive regimes, history of herbivory, and polyploidization in plants. Studies are conducted as a literature review and three experiments on two cruciferous species Raphanus raphanistrum and Cardamine pratensis. In the tolerance experiments, plants are subjected to artificial damage solely, or in a combination with natural damage. A literature review was conducted in order to investigate the effects of damage method. We found that traits related to tolerance, such as growth and fitness were not as sensitive in regard to damage method as measures of induced chemical traits, or measures of secondary herbivory. Genetic variation of tolerance was demonstrated within populations of R. raphanistrum and between subspecies of C. pratensis. In R. raphanistrum, traits involved in floral display and male fitness were positively associated with plant tolerance to herbivore damage. A potential cost of tolerance was demonstrated as a negative correlation between levels of tolerance in high and low competitive regimes. I found no evidence of other proposed costs of tolerance in terms of highly tolerant plants suffering of reduced fitness in the absence of herbivores or trade-offs in terms of a negative association between tolerance to apical and leaf damage, or between tolerance and competitive ability. In C. pratensis, higher ploidy level in plants involved higher levels of tolerance measured as clonal reproduction. Furthermore, populations exposed to higher levels of herbivory had better tolerance than populations exposed to lower levels of herbivory. In this thesis, I demonstrate evidence of different components for the evolution of tolerance in plants: genotypic variation, selective factors in terms of costs and ploidization, and selective agents in terms of changing environment or herbivore pressure.

About 70% of the plant species in the temperate zone are characterised by clonal growth, clonal species are also in majority in the Arctic and Subarctic where they affect the structure and composition of the vegetation. It is therefore of great importance to increase our knowledge about clonal plants and their growth and life histories. I have investigated how ramets of the stoloniferous plant Linnaea borealis are affected by the naturally occurring variation in environmental factors, such as: light, nutrient and water availability. Moreover, I examined the seed set and how supplemental hand pollination affects seed set in L. borealis, and also investigated the significance of the apical meristem for shoot population fitness. All studies were performed under field conditions in northern Sweden in a Subarctic environment and most are experimental. The results show that nutrient resorption from senescing leaves is not significantly affecting the growth and nutrient pools of the ramet. This implies that the growth of L. borealis ramets is not governed by micro-site resource availability. However, removal of light competition resulted in increased branching and number of lateral meristems produced, reduced growth, and decreased root:shoot ratio on a per ramet basis. Thus, ramets of L. borealis can efficiently exploit favourable light patches through plastic growth. Apical dominance exerts a significant effect on shoot population fitness and can be lost through rodent grazing. However, loss of apical dominance is dependent on the timing of grazing, if the apical meristem is removed early in the autumn the ramet can repair the loss until the next summer. If grazing occur during spring the dry weight and leaf area production is affected negatively. Seed production in L. borealis in the Abisko area varies between years and sites, and was unaffected by supplemental hand pollination treatment, implying that there is no lack of pollinator activity.

<p>About 70% of the plant species in the temperate zone are characterised by clonal growth, clonal species are also in majority in the Arctic and Subarctic where they affect the structure and composition of the vegetation. It is therefore of great importance to increase our knowledge about clonal plants and their growth and life histories. I have investigated how ramets of the stoloniferous plant <i>Linnaea borealis</i> are affected by the naturally occurring variation in environmental factors, such as: light, nutrient and water availability. Moreover, I examined the seed set and how supplemental hand pollination affects seed set in <i>L. borealis</i>, and also investigated the significance of the apical meristem for shoot population fitness. All studies were performed under field conditions in northern Sweden in a Subarctic environment and most are experimental.</p><p>The results show that nutrient resorption from senescing leaves is not significantly affecting the growth and nutrient pools of the ramet. This implies that the growth of<i> L. borealis </i>ramets is not governed by micro-site resource availability. However, removal of light competition resulted in increased branching and number of lateral meristems produced, reduced growth, and decreased root:shoot ratio on a per ramet basis. Thus, ramets of <i>L. borealis </i>can efficiently exploit favourable light patches through plastic growth. Apical dominance exerts a significant effect on shoot population fitness and can be lost through rodent grazing. However, loss of apical dominance is dependent on the timing of grazing, if the apical meristem is removed early in the autumn the ramet can repair the loss until the next summer. If grazing occur during spring the dry weight and leaf area production is affected negatively. Seed production in <i>L. borealis</i> in the Abisko area varies between years and sites, and was unaffected by supplemental hand pollination treatment, implying that there is no lack of pollinator activity.</p>

About 70% of the plant species in the temperate zone are characterised by clonal growth, clonal species are also in majority in the Arctic and Subarctic where they affect the structure and composition of the vegetation. It is therefore of great importance to increase our knowledge about clonal plants and their growth and life histories. I have investigated how ramets of the stoloniferous plant Linnaea borealis are affected by the naturally occurring variation in environmental factors, such as: light, nutrient and water availability. Moreover, I examined the seed set and how supplemental hand pollination affects seed set in L. borealis, and also investigated the significance of the apical meristem for shoot population fitness. All studies were performed under field conditions in northern Sweden in a Subarctic environment and most are experimental. The results show that nutrient resorption from senescing leaves is not significantly affecting the growth and nutrient pools of the ramet. This implies that the growth of L. borealis ramets is not governed by micro-site resource availability. However, removal of light competition resulted in increased branching and number of lateral meristems produced, reduced growth, and decreased root:shoot ratio on a per ramet basis. Thus, ramets of L. borealis can efficiently exploit favourable light patches through plastic growth. Apical dominance exerts a significant effect on shoot population fitness and can be lost through rodent grazing. However, loss of apical dominance is dependent on the timing of grazing, if the apical meristem is removed early in the autumn the ramet can repair the loss until the next summer. If grazing occur during spring the dry weight and leaf area production is affected negatively. Seed production in L. borealis in the Abisko area varies between years and sites, and was unaffected by supplemental hand pollination treatment, implying that there is no lack of pollinator activity.