Brominated aromatic compounds (BACs) are widely distributed in the marine environment. Some of these compounds are highly toxic, such as certain hydroxylated polybrominated diphenyl ethers (OH-PBDEs). In addition to anthropogenic emissions through use of BACs as e.g. flame retardants, BACs are natural products formed by marine organisms such as algae, sponges, and cyanobacteria. Little is known of the transfer of BACs from natural producers and further up in the trophic food chain. In this study it was observed that total sum of methoxylated polybrominated diphenyl ethers (MeO-PBDEs) and OH-PBDEs increased in concentration from the filamentous red alga Ceramium tenuicorne, via Gammarus sp. and three-spined stickleback (Gasterosteus aculeatus) to perch (Perca fluviatilis). The MeO-PBDEs, which were expected to bioaccumulate, increased in concentration accordingly up to perch, where the levels suddenly dropped dramatically. The opposite pattern was observed for OH-PBDEs, where the concentration exhibited a general trend of decline up the food web, but increased in perch, indicating metabolic demethylation of MeO-PBDEs. Debromination was also indicated to occur when progressing through the food chain resulting in high levels of tetra-brominated MeO-PBDE and OH-PBDE congeners in fish, while some penta- and hexa-brominated congeners were observed to be the dominant products in the alga. As it has been shown that OH-PBDEs are potent disruptors of oxidative phosphorylation and that mixtures of different congener may act synergistically in terms of this toxic mode of action, the high levels of OH-PBDEs detected in perch in this study warrants further investigation into potential effects of these compounds on Baltic wildlife, and monitoring of their levels.

Tolerance to herbivory is an adaptation that promotes regrowth and maintains fitness in plants after herbivore damage. Here, we hypothesized that the effect of competition on tolerance can be different for different genotypes within a species and we tested how tolerance is affected by competitive regime and damage type. We inflicted apical or leaf damage in siblings of 29 families of an annual plant Raphanus raphanistrum (Brassicaceae) grown at high or low competition. There was a negative correlation of family tolerance levels between competition treatments: plant families with high tolerance to apical damage in the low competition treatment had low tolerance to apical damage in the high competition treatment and vice versa. We found no costs of tolerance, in terms of a trade-off between tolerance to apical and leaf damage or between tolerance and competitive ability, or an allocation cost in terms of reduced fitness of highly tolerant families in the undamaged state. High tolerance bound to a specific competitive regime may entail a cost in terms of low tolerance if competitive regime changes. This could act as a factor maintaining genetic variation for tolerance.

We tested whether differences in ploidy level and previous exposure to herbivory can affect plant tolerance to herbivory. We conducted a common garden experiment with 12 populations of two ploidy levels of the perennial herb Cardamine pratensis (five populations of tetraploid ssp. pratensis and seven populations of octoploid ssp. paludosa). Earlier studies have shown that attack rates by the main herbivore, the orange tip butterfly Anthocharis cardamines, are lower in populations of octoploids than in populations of tetraploids, and vary among populations. In the common garden experiment, a combination of natural and artificial damage significantly reduced seed and flower production. We measured tolerance based on four plant-performance metrics: survival, growth, seed production and clonal reproduction. For three of these measurements, tolerance of damage did not differ between ploidy levels. For clonal reproduction, the octoploids had a higher tolerance than the tetraploids, although they experience lower herbivore attack rates in natural populations. Populations from sites with high levels of herbivory had higher tolerance, measured by seed production, than populations with low levels of herbivory. We did not detect any significant costs of tolerance. We conclude that high intensity of herbivory has selected for high tolerance measured by seed production in C. pratensis.

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.

To study mechanisms underlying plant tolerance to herbivore damage, we used apical and foliar damage as experimental treatments to study whether there are similar tolerance mechanisms to different types of damage. We also studied whether tolerance to different types of damage are associated, and whether there is a cost involved in plant tolerance to different types of herbivore damage. Our greenhouse experiment involved 480 plants from 30 full-sib families of an annual weed Raphanus raphanistrum, wild radish, which were subjected to control and two different simulated herbivore damage treatments, apex removal and foliar damage of 30% of leaf area. Apical damage significantly decreased seed production, whereas foliar damage had no effect. There was a significant genetic variation for tolerance to foliar, but not apical damage. No costs were observed in terms of negative correlation between tolerance to either damage type and fitness of undamaged plants. Tolerances to apical and foliar damage were not significantly correlated with each other. We observed a larger number of significant associations between tolerance and reproductive traits than between tolerance and vegetative traits. Plant height and leaf size of damaged plants interacted in their association to tolerance to foliar damage. Inflorescence number and pollen quantity per flower of damaged plants were positively associated with tolerance to apical damage. In late-flowering genotypes, petal size of undamaged plants and pollen quantity of damaged plants were positively associated with tolerance to foliar damage. In summary, traits involved in floral display and male fitness were associated with plant tolerance to herbivore damage.