Ulva fenestrata is an economically and ecologically important green algal species with a large potential in seaweed aquaculture due to its high productivity, wide environmental tolerance, as well as interesting functional and nutritional properties. Here, we performed a series of manipulative cultivation experiments in order to investigate the effects of irradiance (50, 100, and 160 μmol photons m−2 s−1), temperature (13 and 18 °C), nitrate (< 5, 150, and 500 μM), phosphate (< 1 and 50 μM), and pCO2 (200, 400, and 2500 ppm) on the relative growth rate and biochemical composition (fatty acid, protein, phenolic, ash, and biochar content) in indoor tank cultivation of Swedish U. fenestrata. High irradiance and low temperature were optimal for the growth of this northern hemisphere U. fenestrata strain, but addition of nutrients or changes in pCO2 levels were not necessary to increase growth. Low irradiance resulted in the highest fatty acid, protein, and phenolic content, while low temperature had a negative effect on the fatty acid content but a positive effect on the protein content. Addition of nutrients (especially nitrate) increased the fatty acid, protein, and phenolic content. High nitrate levels decreased the total ash content of the seaweeds. The char content of the seaweeds did not change in response to any of the manipulated factors, and the only significant effect of changes in pCO2 was a negative relationship with phenolic content. We conclude that the optimal cultivation conditions for Swedish U. fenestrata are dependent on the desired biomass traits (biomass yield or biochemical composition).
Climate change-related effects threaten species worldwide; within-species populations may react differently to climate-induced stress due to local adaptation and partial isolation, particularly in areas with steep environmental gradients. Populations of the marine foundation seaweed Fucus vesiculosus are established over a steep salinity gradient at the entrance of the brackish water in the Baltic Sea (NE Atlantic). First, we analyzed the genetic differentiation among populations using thousands of genetic markers. Second, we measured the physiological tolerance to reduced salinity, a predicted effect of climate change in the study area, by measuring growth, phlorotannin (defense compounds) content, and maximum photochemical yield in tissue of the same individuals exposed to both current and projected future salinities. Our results show that despite short geographic distances (max 100 km), most populations were genetically well separated. Furthermore, populations responded very differently to a salinity decrease of four practical salinity units (psu) corresponding to projected future salinity. At the high salinity end of the gradient, some populations maintained growth at the cost of reduced phlorotannin production. However, at the low salinity end, mortality increased and growth was strongly reduced in one population, while a second population from similar salinity instead maintained growth and phlorotannin production. Among genetic markers that appeared as outliers (showing more genetic differentiation than the majority of loci), we found that four were associated with genes that were potential candidates for being under selection. One of these, a calcium-binding protein gene, also showed a significant genotype-phenotype relationship in the population where this genetic marker was variable. We concluded that local selection pressure, genetic affinity, and possibly also population history could explain the very different responses to reduced salinity among these populations, despite being from the same geographic area. Our results highlight the importance of local perspective in the management of species.
In recent years, the interest in using seaweed for the sustainable production of commodities has been increasing as seaweeds contain many potentially worthwhile compounds. Thus, the extraction and refining processes of interesting compounds from seaweeds is a hot research topic but has been found to have problems with profitability for novel applications. To increase the economic potential of refining seaweed biomass, the content of the compounds of interest should be maximized, which can potentially be achieved through optimization of cultivation conditions. In this study, we studied how the monosaccharide composition of the green seaweed species Ulva fenestrata is influenced by the abiotic factors; irradiance, temperature, nitrate, phosphate, and pCO2. It was evident that lower nitrate concentration and cultivation at elevated temperature increased monosaccharide contents. A 70% increase in iduronic acid and a 26% increase in rhamnose content were seen under elevated irradiance and temperature conditions, though the absolute differences in monosaccharide concentration were small. Irradiance and nitrate impacted the ratio between iduronic and rhamnose, which is an indicator of the ulvan structure. These results could potentially be utilized to coax the ulvan towards specific bioactivities, and thus have a considerable impact on a potential biorefinery centered around Ulva.
Ocean acidification driven by anthropogenic climate change is causing a global decrease in pH, which is projected to be 0.4 units lower in coastal shallow waters by the year 2100. Previous studies have shown that seaweeds grown under such conditions may alter their growth and photosynthetic capacity. It is not clear how such alterations might impact interactions between seaweed and herbivores, e.g. through changes in feeding rates, nutritional value, or defense levels. Changes in seaweeds are particularly important for coastal food webs, as they are key primary producers and often habitat-forming species. We cultured the habitat-forming brown seaweed Fucus vesiculosus for 30 days in projected future pCO2 (1100 μatm) with genetically identical controls in ambient pCO2 (400 μatm). Thereafter the macroalgae were exposed to grazing by Littorina littorea, acclimated to the relevant pCO2-treatment. We found increased growth (measured as surface area increase), decreased tissue strength in a tensile strength test, and decreased chemical defense (phlorotannins) levels in seaweeds exposed to high pCO2-levels. The herbivores exposed to elevated pCO2-levels showed improved condition index, decreased consumption, but no significant change in feeding preference. Fucoid seaweeds such as F. vesiculosus play important ecological roles in coastal habitats and are often foundation species, with a key role for ecosystem structure and function. The change in surface area and associated decrease in breaking force, as demonstrated by our results, indicate that F. vesiculosus grown under elevated levels of pCO2 may acquire an altered morphology and reduced tissue strength. This, together with increased wave energy in coastal ecosystems due to climate change, could have detrimental effects by reducing both habitat and food availability for herbivores.
Climate change leads to multiple effects caused by simultaneous shifts in several physical factors which will interact with species and ecosystems in complex ways. In marine systems the effects of climate change include altered salinity, increased temperature, and elevated pCO2 which are currently affecting and will continue to affect marine species and ecosystems. Seaweeds are primary producers and foundation species in coastal ecosystems, which are particularly vulnerable to climate change. The brown seaweed Fucus vesiculosus (bladderwrack) is an important foundation species in nearshore ecosystems throughout its natural range in the North Atlantic Ocean and the Baltic Sea. This study investigates how individual and interactive effects of temperature, salinity, and pCO2 affect F. vesiculosus, using a fully crossed experimental design. We assessed the effects on F. vesiculosus in terms of growth, biochemical composition (phlorotannin content, C:N ratio, and ∂13C), and susceptibility to the specialized grazer Littorina obtusata. We observed that elevated pCO2 had a positive effect on seaweed growth in ambient temperature, but not in elevated temperature, while growth increased in low salinity at ambient but not high temperature, regardless of pCO2-level. In parallel to the statistically significant, but relatively small, positive effects on F. vesiculosus growth, we found that the seaweeds became much more susceptible to grazing in elevated pCO2 and reduced salinity, regardless of temperature. Furthermore, the ability of the seaweeds to induce chemical defenses (phlorotannins) was strongly reduced by all the climate stressors. Seaweeds exposed to ambient conditions more than doubled their phlorotannin content in the presence of grazers, while seaweeds exposed to any single or combined stress conditions showed only minor increases in phlorotannin content, or none at all. Despite the minor positive effects on seaweed growth, the results of this study imply that climate change can strongly affect the ability of fucoid seaweeds to induce chemical defenses and increase their susceptibility to grazers. This will likely lead to widespread consequences under future climate conditions, considering the important role of F. vesiculosus and other fucoids in many coastal ecosystems.
Adaptive phenotypic plasticity may improve the response of individuals when faced with new environmental conditions. Typically, empirical evidence for plasticity is based on phenotypic reaction norms obtained in reciprocal transplant experiments. In such experiments, individuals from their native environment are transplanted into a different environment, and a number of trait values, potentially implicated in individuals' response to the new environment, are measured. However, the interpretations of reaction norms may differ depending on the nature of the assessed traits, which may not be known beforehand. For example, for traits that contribute to local adaptation, adaptive plasticity implies non-zero slopes of reaction norms. By contrast, for traits that are correlated to fitness, high tolerance to different environments (possibly due to adaptive plasticity in traits that contribute to adaptation) may, instead, result in flat reaction norms. Here we investigate reaction norms for adaptive versus fitness-correlated traits, and how they may affect the conclusions regarding the contribution of plasticity. To this end, we first simulate range expansion along an environmental gradient where plasticity evolves to different values locally and then perform reciprocal transplant experiments in silico. We show that reaction norms alone cannot inform us whether the assessed trait exhibits locally adaptive, maladaptive, neutral or no plasticity, without any additional knowledge of the traits assessed and species' biology. We use the insights from the model to analyse and interpret empirical data from reciprocal transplant experiments involving the marine isopod Idotea balthica sampled from two geographical locations with different salinities, concluding that the low-salinity population likely has reduced adaptive plasticity relative to the high-salinity population. Overall, we conclude that, when interpreting results from reciprocal transplant experiments, it is necessary to consider whether traits assessed are locally adaptive with respect to the environmental variable accounted for in the experiments, or correlated to fitness.
Background In the brackish Baltic Sea, shedding of adventitious branches is central to asexual recruitment of new thalli in the brown algae Fucus vesiculosus and F. radicans . To test which factors influence the formation of adventitious branches in brackish and in more marine conditions, we sampled 29 Fucus sites in the Baltic Sea (salinity 3–11) and 18 sites from the Danish straits, Kattegat, Skagerrak, and the North Sea (salinity 15–35). Separately for each area, we used structural equation modelling to determine which of eight predictor factors (phosphate, nitrate, chlorophyll- a (as a proxy for turbidity), temperature, salinity, oxygen, grazing pressure, and thallus area) best explained observed numbers of adventitious branches. Results In more marine waters, high yearly average values of phosphate, salinity and turbidity had positive effects on the formation of adventitious branches. In brackish-waters, however, high numbers of adventitious branches were found in areas with low yearly average values of temperature, salinity and oxygen. Grazing intensity had no significant effect in either of the two study areas, contrasting findings from studies in other areas. In areas with both sexually and asexually reproducing Fucus individuals, clones had on average more adventitious branches than unique genotypes, although there was strong variation among clonal lineages. Conclusion This study is the first to investigate multiple potential drivers of formation of adventitious branches in natural populations of Fucus . Our results suggest that several different factors synergistically and antagonistically affect the growth of adventitious branches in a complex way, and that the same factor (salinity) can have opposing effects in different areas. Electronic supplementary material The online version of this article (10.1186/s12898-019-0239-7) contains supplementary material, which is available to authorized users.
Phytoplankton induce defensive traits in response to chemical alarm signals from grazing zooplankton. However, these signals are potentially vulnerable to changes in pH and it is not yet known how predator recognition may be affected by ocean acidification. We exposed four species of diatoms and one toxic dinoflagellate to future pCO2 levels, projected by the turn of the century, in factorial combinations with predatory cues from copepods (copepodamides). We measured the change in growth, chain length, silica content, and toxin content. Effects of increased pCO2 were highly species specific. The induction of defensive traits was accompanied by a significant reduction in growth rate in three out of five species. The reduction averaged 39% and we interpret this as an allocation cost associated with defensive traits. Copepodamides induced significant chain length reduction in three of the four diatom species. Under elevated pCO2Skeletonema marinoi reduced silica content by 30% and in Alexandrium minutum the toxin content was reduced by 30%. Using copepodamides to induce defensive traits in the absence of direct grazing provides a straightforward methodology to assess costs of defense in microplankton. We conclude that copepodamide signalling system is likely robust to ocean acidification. Moreover, the variable responses of different taxa to ocean acidification suggest that there will be winners and losers in a high pCO2 world, and that ocean acidification may have structuring effects on phytoplankton communities.
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