Eränen, J. 2003. Induction of phlorotannin production in a brown alga: defense or resource dynamics? -Oikos 103: 640-650.Increase of phenolic secondary metabolites, phlorotannins, in brown algae due to gastropod grazing has been interpreted as an anti-herbivore adaptation. Here we tested whether such a response could be due to changes in truly available resources for the alga, not by the grazing activity of snails as such. We allowed two species of snails, Theodoxus flu6iatilis and Physa fontinalis to graze on Fucus 6esiculosus. These species feed on epibiota and particulate matter on the thallus but do not eat the thallus of F. 6esiculosus. We further simulated snail grazing by nutrient enhancement, removal of epibiota and by a combination of the two. Manipulations of nutrient and light availability revealed the crucial role of epibiota in mediating resource availability for F. 6esiculosus. Nutrient enhancement alone increased epibiota and decreased phlorotannins. Cleaning the thallus resulted in increased growth, and together with nutrient enhancement also in a trade-off with phlorotannins. Presence of T. flu6iatilis on the thallus induced phlorotannin production, a response differing from the simulations of snail grazing. However, we suggest that the increase in phlorotannins may not be an induced defense but rather a consequence of a specific way of resource manipulation by this snail species. T. flu6iatilis removes hyaline hairs that facilitate nutrient uptake. P. fontinalis did not remove hyaline hairs and the response of the alga to its grazing was similar to the treatment where we mechanically removed epibiota suggesting that cleaning of the thallus is the major mechanism how this snail species affects F. 6esiculosus. Genetic variation in phlorotannin concentrations highly exceeded the induced responses of simulated or real snail grazing. This casts doubt for the efficiency of induced phlorotannin production to act as a defense, but is not contradictory with the interpretation of phlorotannins responding to variation in resource availability.

In ecological studies, phlorotannins have conventionally been quantified as a group with similar functionality. Since this group consists of oligo- and polymers, the quantification of their pooled contents alone may not sufficiently describe the variation of these metabolites. Genetic variation, plastic responses to environment, and the ecological functions of separate phlorotannin oligo- and polymers may differ. Two analyses, i.e., the colorimetric Folin-Ciocalteu assay and a normal-phase high-performance liquid chromatographic (HPLC) method were used to study genetic and environmental variation in phlorotannins of the brown alga Fucus vesiculosus (L.). The colorimetric method provides the total phlorotannin content, the latter a profile of 14 separate traces from the phenolic extract that represent an individual or groups of phlorotannins. We reared the algae that originated from three separate populations in a common garden for 3 months under ambient and enriched-nutrient availability and found that they differed in both their total phlorotannin content and in phlorotannin profiles. Some individual traces of the profiles separated the populations more clearly than the colorimetric assay. Although nutrient enrichment decreased total phlorotannin content, it did not show a significant influence on the phlorotannin profile. This implies that plastic responses of compounds other than phlorotannins may interfere with the determination of total phlorotannins. However, the phlorotannin profile and the total content showed genetic variation among local populations of F. vesiculosus; therefore, phlorotannins may respond to natural selection and evolve both quantitatively and qualitatively.

The stress-gradient hypothesis predicts that the importance of facilitation relative to competition should increase with increasing stress. The hypothesis has received support from several environments, but multi-gradient studies on the generality of the hypothesis are exceptionally rare. A within-species experiment with mountain birch Betula pubescens subsp. czerepanovii was conducted to test the hypothesis in the extreme ends of two subarctic stress gradients (elevation and seashore) in the Kola Peninsula, northwestern Russia. The high stress sites were characterized by strong winds, temperature extremes and potentially drought. The negative effects of abiotic stress on the study seedlings were verified from performance characteristics. Effects of adult hosts as well as seedlingÁseedlings interactions were studied. Positive hostÁseedling interactions dominated in each study site, and three out of four performance variables indicated stronger positive net effects in the high stress sites. In the seashore gradient also seedling survival gave similar interpretations. Also a temporal shift towards hostÁseedling competition was detected in a low stress site after two study years. In seedlingÁ seedling interactions competition dominated, but the effects were weak, likely due to the 'noise' caused by genetic and environmental factors. Our results support the stress-gradient hypothesis and its generality in subarctic environments, as the interpretations were similar for both stress gradients and several fitness-related variables. The temporal variation in hostÁseedling interactions and the difference between hostÁseedling and seedlingÁseedling effects hint on size-dependency of plant interactions: facilitation might dominate when the benefactor is substantially larger than the beneficiary, while competition may be stronger when the plants are of similar size and developmental status.

Adaptations to pollution among long‐lived trees have rarely been documented, possibly because of their long reproductive cycles and the evolutionarily short timescales of anthropogenic pollution. Here, I present the results of a greenhouse experiment that suggest rapid evolutionary adaptation of mountain birch [Betula pubescens subsp. czerepanovii (Orlova) Hämet‐Ahti] to heavy metal (HM) stress around two copper–nickel smelters in NW Russia. The adaptation incurs a cost with reduced performance of adapted seedlings in pristine conditions. The industrial barrens around the studied smelters are extremely high‐stress sites with low seed germination and survival. It is likely that strong natural selection has eliminated all sensitive genotypes within one or two generations, with only the most tolerant individuals persisting and producing adapted seeds in the individual barrens. The results were similar from around both smelters, suggesting parallel evolution towards HM resistance.

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