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The green seaweed Ulva fenestrata is a future food candidate, however, both compositional, e.g. protein content, and sensory qualities depend on cultivation and harvest conditions. The aim was to explore sensory qualities of, and consumer attitudes to, U. fenestrata derived from three different cultivation conditions and three harvest times. U. fenestrata was cultivated at sea and in two different land-based tank settings, one mimicking seawater and one with added process water from fishing industry to increase protein content. The seaweed was subjected to sensory analysis performed by an analytical sensory panel and a focus group consisting of consumers. The former assessed the seaweed as dried whole blades (9 samples) and as emulsions (9 samples). Consumers assessed whole blades and a vegan spread enriched with powdered U. fenstrata. All seaweed samples were intensely green; had tastes of umami, salt, bitter, and sour; odours and flavours of grass and fresh seaweed; and had a crispy texture. Cultivation in tanks resulted in a crispier texture than sea cultivation (p

IntroductionCampylobacter jejuni gastroenteritis is the most commonly reported zoonosis within the EU, with poultry products regarded as the primary source of transmission to humans. Therefore, finding strategies to reduce Campylobacter colonization in broilers holds importance for public health. Recent studies suggest that supplementation of broiler feed with brown algal extracts, particularly laminarin, can provide beneficial effects on broiler gut health, growth performance, and gut microbiota. However, its effect on gut microbiota development and subsequent reduction of Campylobacter loads in broiler caeca during the later stages of the birds' lives remains unclear.MethodsExperimental colonization of Ross 308 broilers with two different strains of C. jejuni was conducted, with groups fed either a basal diet or the same basal diet supplemented with 725 ppm algal extract from Saccharina latissima to provide 290 ppm laminarin. Fecal samples were collected for bacterial enumeration, and caecal samples were obtained before and after the C. jejuni challenge for the determination of microbiota development.Results and discussionNo significant differences in fecal C. jejuni concentrations between the groups fed different diets or exposed to different C. jejuni strains were observed. This suggests that both strains colonized the birds equally well and that the laminarin rich algal extract did not have any inhibitory effect on C. jejuni colonization. Notably, 16S rRNA amplicon sequencing revealed detailed data on the caecal microbiota development, likely influenced by both bird age and C. jejuni colonization, which can be valuable for further development of broiler feed formulations aimed at promoting gut health.

Sea lettuce (Ulva) species have been identified as a future protein source, and post-harvest techniques, including the sustainable integration of liquid food side streams, have been developed to further increase their protein content and commercial market value. This study explores the post-harvest treatment of commercially produced seaweed biomass from large-scale, sea-based cultivations of U. fenestrata with residual water streams emerging from industrial storage of herring – so called herring production tub water (TUB). Growth rates of U. fenestrata were significantly higher in TUB treated seaweeds compared to controls. Further, the crude protein content was 71.26 % higher when cultivated in TUB, compared to controls, reaching a crude protein content of 37.37 ± 1.83 % dry weight. Notably, there were no limiting amino acids, nor fish-related allergenic activity in the seaweed biomass. Our study demonstrates a new nutrient loop turning food waste into protein-rich biomass by applying sustainable seaweed cultivation.

A nutritional bottleneck in salmonid aquaculture is the procurement of marine-derived compounds, such as essential amino and fatty acids, including omega-3 fatty acids, lysine, and methionine. Therefore, insects containing these compounds are highly promising as feed ingredients. The present study evaluates larvae of a “marine” insect (Coelopa frigida, the bristly-legged seaweed fly larvae, SWFL) reared on brown algae side streams as a feed ingredient for rainbow trout (Oncorhynchus mykiss). SWFL contained, on a dry matter basis, 60% crude protein (CP), 3.5% lysine, and 1.5% methionine, as well as 17% lipids, including 4% eicosapentaenoic acid and docosahexaenoic acid. Four isoenergetic (ca. 23 MJ kg−1 gross energy) and isoproteic (ca. 45% CP) feeds were evaluated in a 10-week feeding trial. The diets included a control containing 25% fish meal, a commercial reference, and two diets substituting 40% fish meal with either SWFL or partially defatted black soldier fly larvae (BSFL) meal (Hermetia illucens). SWFL-fed fish displayed higher specific growth rates (SGR) compared to BSFL-fed fish and similar SGR compared to control and reference diet-fed fish. Feed intake in SWFL-fed fish was higher than for the control and BSFL diets and similar to the reference diet. The feed conversion ratio of fish fed the BSFL and SWFL diets was higher compared to the control, suggesting lower bioavailability of both insect meals compared to fish meals. No difference in intestinal health parameters was noted between the insect diets and the control diet, indicating good intestinal health across all treatments. However, changes in electrogenic intestinal transport were observed between the SWFL and BSFL diets, illustrating the heterogeneous effect of different insect products. Overall, SWFL meal is a promising alternative marine feed ingredient, compatible with circular production systems, as it can be efficiently cultivated using marine side streams.

Abstract The Centre for Marine Evolutionary Biology (CeMEB) at the University of Gothenburg, Sweden, was established in 2008 through a 10‐year research grant of 8.7 m€ to a team of senior researchers. Today, CeMEB members have contributed >500 scientific publications, 30 PhD theses and have organised 75 meetings and courses, including 18 three‐day meetings and four conferences. What are the footprints of CeMEB, and how will the centre continue to play a national and international role as an important node of marine evolutionary research? In this perspective article, we first look back over the 10 years of CeMEB activities and briefly survey some of the many achievements of CeMEB. We furthermore compare the initial goals, as formulated in the grant application, with what has been achieved, and discuss challenges and milestones along the way. Finally, we bring forward some general lessons that can be learnt from a research funding of this type, and we also look ahead, discussing how CeMEB’s achievements and lessons can be used as a springboard to the future of marine evolutionary biology.

Abstract Inversions often underlie complex adaptive traits, but the genic targets inside them are largely unknown. Gene expression profiling provides a powerful way to link inversions with their phenotypic consequences. We examined the effects of the Cf‐Inv(1) inversion in the seaweed fly Coelopa frigida on gene expression variation across sexes and life stages. Our analyses revealed that Cf‐Inv(1) shapes global expression patterns, most likely via linked variation, but the extent of this effect is variable, with much stronger effects in adults than larvae. Furthermore, within adults, both common as well as sex‐specific patterns were found. The vast majority of these differentially expressed genes mapped to Cf‐Inv(1). However, genes that were differentially expressed in a single context (i.e., in males, females, or larvae) were more likely to be located outside of Cf‐Inv(1). By combining our findings with genomic scans for environmentally associated SNPs, we were able to pinpoint candidate variants in the inversion that may underlie mechanistic pathways that determine phenotypes. Together the results of this study, combined with previous findings, support the notion that the polymorphic Cf‐Inv(1) inversion in this species is a major factor shaping both coding and regulatory variation resulting in highly complex adaptive effects.

There is an increasing interest in the cultivation of seaweeds for food and feed, and the seaweed aquaculture industry is rapidly developing. The nutritional status of the seaweeds is important to ensure a good quality crop. Cost-efficient and straightforward methods for farmers to analyze their crop are essential for the successful development of the industry. In this study, we developed non-destructive, labor- and cost-efficient models to estimate the nitrogen content in the crop seaweed Ulva fenestrata by color image analysis. We quantified tissue nitrogen content and thallus color in sea-farmed seaweed every week throughout a whole cultivation season (15 consecutive weeks) and analyzed data with linear regression models. We showed that color image analysis accurately estimated the nitrogen content in the seaweed (R2 = 0.944 and 0.827 for fresh tissue and dried powder, respectively), and through tenfold cross validation we showed that the developed models were robust and precise. Based on these models, we developed a web-based application that automatically analyzes the nitrogen content of U. fenestrata. Furthermore, we produced a color guide that can easily be brought to the farm for onsite crude estimation of the nitrogen content of U. fenestrata. Our results demonstrate that color can be a powerful tool for seaweed farmers (and researchers) to estimate seaweeds’ nutritional status. We anticipate that similar models can be developed for other commercially interesting seaweed species.

Seaweed aquaculture can provide the growing human population with a sustainable source of proteins. Sea-based cultivation is an effective method for farming seaweeds on a large scale and can yield high biomass output. However, the quality and biochemical composition of the biomass is seasonally dependent, which limits the harvests to certain periods of the year. Here we show the possibility to extend the sea-based cultivation season of Ulva fenestrata when aiming for high protein levels, by post-harvest treatment in herring production process waters. We harvested U. fenestrata at an optimal period in terms of yield, but suboptimal in terms of protein content. We then cultivated the seaweed in onshore tank systems with the nutrient-rich process waters for 14 days. We monitored biomass yield, crude protein content, amino acid composition, and content of the health concerning metals arsenic, mercury, lead, and cadmium, as well as the sensory properties of the dried biomass. After cultivation in the process waters, biomass yields were 30 - 40% higher (210 – 230 g fresh weight) compared to in seawater (160 g fresh weight). Also, the crude protein and amino acid content increased three to five times in the process waters, reaching 12 - 17 and 15 – 21% dry weight, respectively. The protein enriched biomass followed food graded standards for heavy metal content, and consumption of the biomass does not exceed health based reference points. Additionally, no sensory attributes regarded as negative were found. This rapid, post-harvest treatment can help extend the cultivation season of sea-based seaweed farms, maximizing their output of sustainable proteins.

Ulva spp. (sea lettuce) has recently gained attention as a sustainable protein source due to its high productivity and many nutritional properties interesting for the food industry. In this study, we explored a possible industrial symbiosis between herring production processing industries and Ulva fenestrata cultivation. We show that U. fenestrata cultivated in herring production process waters had four to six times higher biomass yields (27.17 - 37.07 g fresh weight vs. 6.18 g fresh weight) and three times higher crude protein content (> 30% dry weight vs. 10% dry weight) compared to U. fenestrata cultivated in seawater. Along with the elevation of protein, the herring production process waters also significantly increased levels of all essential amino acids in the seaweed biomass. The content of some heavy metals (arsenic, mercury, lead, and cadmium) was well below the maximum allowed levels in foodstuff. Therefore, quantities of biomass around 100 g dry weight could be consumed daily following the US Environmental Protection Agency’s reference doses. Combined, the results show that cultivation of U. fenestrata in herring production process waters has great potential to produce sustainable proteins for the growing world population. At the same time, nutrients of currently discarded process waters are circulated back to the food chain.

Sea lettuce (Ulva) aquaculture has increased the last decade due to high productivity, wide environmental tolerance, and interesting functional and nutritional properties of the crop. Research focus has mainly been on adult biomass production, but knowledge of performance and biochemical content of early developmental stages – which are the basis to any large-scale production - is still limited. The life-history of Ulva alternates between a diploid sporophytic life-stage and a haplontic gametophytic life-stage. Whereas the sporophyte give raise to recombinant gametophytes through zoids, gametophytes can give raise to parthenogenetically developing, clonal gametes in absence of a mating partner. This study shows that recombinant gametophytes have a faster ontogenetic development, higher growth rate, as well as higher protein, fatty acid, and pigment contents compared to clonal gametophytes of the crop Ulva fenestrata. Nutrient addition is required for a normal development, but temperature and swarmer density have relatively small effects on the hatchery success, relative growth rate and biochemical profile of the juvenile biomass. Our study reveals that the selection of the life-history-phase in novel sea lettuce crop strains could largely contribute to the emerging seaweed aquaculture sector.

Hatching concepts such as on-farm hatching provide an opportunity to supply newly hatched chickens with optimal nutrition that support growth and development of a healthy gut. Brown algae contain bioactive compounds, especially laminarin and fucoidan that may improve intestinal health and immune responses. This study aimed to examine the effects of early access to feed and water posthatch and feed supplementation with algal extract rich in laminarin from Laminaria digitata, on growth performance, organ and microbiota development and antibody production. A total of 432 Ross 308 chicks were allotted to 36 rearing pens in a 2 × 3 factorial design with two hatching treatments and three dietary treatments. During chick placement, half of the pens were directly provided access to feed and water (Early) while half of the pens were deprived of feed and water for 38 h (Late). The chicks were fed three different starter diets until day 6; a wheat-soybean meal-based control diet, a diet with low inclusion of algal extract (0.057%) and a diet with high inclusion of algal extract (0.114%). Feed intake and BW were registered on pen basis at placement, days 1, 6, 12, 19, 26, 33 and 40. To induce antibody responses, all chicks were vaccinated against avian pneumovirus on day 10. Three chicks per pen were selected as focal animals and used for blood sampling on days 10 and 39. On days 6, 19, and 40, two birds per pen were killed and used for organ measurement and caecal digesta sampling for gut microbiota analysis using the Illumina Miseq PE 250 sequencing platform. Results showed that algal extract did not influence gut microbiota, gut development or vaccine-induced antibody responses. However, during the first 38 h, early-fed chicks consumed on average 19.6 g of feed and gained 27% in BW, while late-fed chicks lost 9.1% in BW which lowered BW and feed intake throughout the study (P

Seaweed biomass is a renewable resource with multiple applications. Sea-based cultivation of seaweeds can provide high biomass yields, low construction, operation, and maintenance costs and could offer an environmentally and economically sustainable alternative to land-based cultivations. The biochemical profile of sea-grown biomass depends on seasonal variation in environmental factors, and the optimization of harvest time is important for the quality of the produced biomass. To identify optimal harvest times of Swedish sea-based cultivated sea lettuce (Ulva fenestrata), this study monitored biomass yield, morphology, chemical composition, fertility, and biofouling at five different harvesting times in April – June 2020. The highest biomass yields (approximately 1.2 kg fw [m rope]–1) were observed in late spring (May). The number and size of holes in the thalli and the amount of fertile and fouled tissue increased with prolonged growth season, which together led to a significant decline in both biomass yield and quality during summer (June). Early spring (April) conditions were optimal for obtaining high fatty acid, protein, biochar, phenolic, and pigment contents in the biomass, whereas carbohydrate and ash content, as well as essential and non-essential elements, increased later in the growth season. Our study results show that the optimal harvest time of sea-based cultivated U. fenestrata depends on the downstream application of the biomass and must be carefully selected to balance yield, quality, and desired biochemical contents to maximize the output of future sea-based algal cultivations in the European Northern Hemisphere.

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.

Abstract Cuticular hydrocarbons (CHCs) form the boundary between insects and their environments and often act as essential cues for species, mate, and kin recognition. This complex polygenic trait can be highly variable both among and within species, but the causes of this variation, especially the genetic basis, are largely unknown. In this study, we investigated phenotypic and genetic variation of CHCs in the seaweed fly, Coelopa frigida, and found that composition was affected by both genetic (sex and population) and environmental (larval diet) factors. We subsequently conducted behavioral trials that show CHCs are likely used as a sexual signal. We identified general shifts in CHC chemistry as well as individual compounds and found that the methylated compounds, mean chain length, proportion of alkenes, and normalized total CHCs differed between sexes and populations. We combined these data with whole genome resequencing data to examine the genetic underpinnings of these differences. We identified 11 genes related to CHC synthesis and found population‐level outlier SNPs in 5 that are concordant with phenotypic differences. Together these results reveal that the CHC composition of C. frigida is dynamic, strongly affected by the larval environment, and likely under natural and sexual selection.

Abstract In the course of the ongoing global intensification and diversification of human pressures, the study of variation patterns of biological traits along environmental gradients can provide relevant information on the performance of species under shifting conditions. The pronounced salinity gradient, co‐occurrence of multiple stressors, and accelerated rates of change make the Baltic Sea and its transition to North Sea a suitable region for this type of study. Focusing on the bladderwrack Fucus vesiculosus, one of the main foundation species on hard‐bottoms of the Baltic Sea, we analyzed the phenotypic variation among populations occurring along 2,000 km of coasts subjected to salinities from 4 to >30 and a variety of other stressors. Morphological and biochemical traits, including palatability for grazers, were recorded at 20 stations along the Baltic Sea and four stations in the North Sea. We evaluated in a common modeling framework the relative contribution of multiple environmental drivers to the observed trait patterns. Salinity was the main and, in some cases, the only environmental driver of the geographic trait variation in F. vesiculosus. The decrease in salinity from North Sea to Baltic Sea stations was accompanied by a decline in thallus size, photosynthetic pigments, and energy storage compounds, and affected the interaction of the alga with herbivores and epibiota. For some traits, drivers that vary locally such as wave exposure, light availability or nutrient enrichment were also important. The strong genetic population structure in this macroalgae might play a role in the generation and maintenance of phenotypic patterns across geographic scales. In light of our results, the desalination process projected for the Baltic Sea could have detrimental impacts on F. vesiculosus in areas close to its tolerance limit, affecting ecosystem functions such as habitat formation, primary production, and food supply.

Abstract 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.

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.

The temperate seagrass species eelgrass Zostera marina can be infected by the wasting disease pathogen Labyrinthula zosterae, which is believed to have killed about 90% of the seagrass in the Atlantic Ocean in the 1930s. It is not known why this opportunistic pathogen sometimes becomes virulent, but the recurrent outbreaks may be due to a weakening of the Z. marina plants from adverse environmental changes. This study investigated the individual and interactive effects of multiple extrinsic factors (temperature, light, and tissue damage) on the host-pathogen interaction between Z. marina and L. zosterae in a fully crossed infection experiment. The degree of infection was measured as both lesion coverage and L. zosterae cell concentration. We also investigated if the treatment factors affect the chemical defense of the host, measured as the inhibitory capacity of seagrass extracts in bioassays with L. zosterae. Finally, gene expression of a set of targeted genes was quantified in order to investigate how the treatments change Z. marina’s response to infection. Light had a pronounced effect on L. zosterae infection measured as lesion coverage, where reduced light conditions increased lesions by 35%. The response to light on L. zosterae cell concentration was more complex and showed significant interaction with the temperature treatment. Cell concentration was also significantly affected by physical damage, where damage surprisingly resulted in a reduced cell concentration of the pathogen. No treatment factor caused detectable decrease in the inhibitory capacity of the seagrass extracts. There were several interactive effects between L. zosterae infection and the treatment factors on Z. marina growth, and on the expression of genes associated with immune defense, phenol synthesis and primary metabolism, showing that the molecular reaction toward L. zosterae infection depends on prevailing environmental conditions. Our study shows that individual or interactive effects of light, temperature and tissue damage can affect multiple aspects of host-pathogen interactions in seagrasses. These results highlight the complexity of marine host-pathogen systems, showing that more multi-factorial investigations are needed to gain a better understanding of disease in marine plants under different environmental conditions.

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.

The growing world population demands an increase in sustainable resources for biorefining. The opening of new farm grounds and the cultivation of extractive species, such as marine seaweeds, increases worldwide, aiming to provide renewable biomass for food and non-food applications. The potential for European large-scale open ocean farming of the commercial green seaweed crop Ulva is not yet fully realized. Here we conducted manipulative cultivation experiments in order to investigate the effects of hatchery temperature (10 and 15 °C), nutrient addition (PES and 3xPES) and swarmer density (500 and 10,000 swarmers ml−1) on the biomass yield and biochemical composition (fatty acid, protein, carbohydrate, pigment and phenolic content) of off-shore cultivated Ulva fenestrata in a Swedish seafarm. High seedling densities were optimal for the growth of this northern hemisphere crop strain and significantly increased the mean biomass yield by ~84% compared to low seedling densities. Variations of nutrients or changes in temperature levels during the hatchery phase were not necessary to increase the subsequent growth in an open-water seafarm, however effects of the factors on the thallus habitus (thallus length/width) were observed. We found no significant effect of the environmental factors applied in the hatchery on the total fatty acid or crude protein content in the off-shore cultivated Ulva. However, low seedling density and low temperature increased the total carbohydrate content and furthermore, high temperature in combination with high nutrient levels decreased the pigment content (chlorophyll a, b, carotenoids). Low temperature in combination with high nutrient levels increased the phenolic content. Our study confirms the successful and sustainable potential for large-scale off-shore cultivation of the Scandinavian crop U. fenestrata. We conclude that high seedling density in the hatchery is most important for increasing the total biomass yield of sea-farmed U. fenestrata, and that changing temperature or addition of nutrients overall does not have a large effect on the biochemical composition. To summarize, our study contributes novel insights into the large-scale off-shore cultivation potential of northern hemisphere U. fenestrata and underpins suitable pre-treatments during the hatchery phase of seedlings to facilitate a successful and cost-efficient large-scale rope cultivation.

Copepods are ubiquitous in aquatic habitats. They exude bioactive compounds that mediate mate finding or induce defensive traits in prey organisms. However, little is known about the chemical nature of the copepod exometabolome that contributes to the chemical landscape in pelagic habitats. Here we describe the development of a closed loop solid phase extraction setup that allows for extraction of exuded metabolites from live copepods. We captured exudates from male and female Temora longicornis and analyzed the content with high resolution LC-MS. Chemometric methods revealed 87 compounds that constitute a specific chemical pattern either qualitatively or quantitatively indicating copepod presence. The majority of the compounds were present in both female and male exudates, but nine compounds were mainly or exclusively present in female exudates and hence potential pheromone candidates. Copepodamide G, known to induce defensive responses in phytoplankton, was among the ten compounds of highest relative abundance in both male and female extracts. The presence of copepodamide G shows that the method can be used to capture and analyze chemical signals from living source organisms. We conclude that solid phase extraction in combination with metabolic profiling of exudates is a useful tool to develop our understanding of the chemical interplay between pelagic organisms.

Photosynthesis is a fundamental process sustaining heterotrophic organisms at all trophic levels. Some mixotrophs can retain functional chloroplasts from food (kleptoplasty), and it is hypothesized that carbon acquired through kleptoplasty may enhance trophic energy transfer through increased host growth efficiency. Sacoglossan sea slugs are the only known metazoans capable of kleptoplasty, but the relative fitness contributions of heterotrophy through grazing, and phototrophy via kleptoplasts, are not well understood. Fitness benefits (i.e. increased survival or growth) of kleptoplasty in sacoglossans are commonly studied in ecologically unrealistic conditions under extended periods of complete darkness and/or starvation. We compared the growth efficiency of the sacoglossan Elysia viridis with access to algal diets providing kleptoplasts of differing functionality under ecologically relevant light conditions. Individuals fed Codium fragile, which provide highly functional kleptoplasts, nearly doubled their growth efficiency under high compared to low light. In contrast, individuals fed Cladophora rupestris, which provided kleptoplasts of limited functionality, showed no difference in growth efficiency between light treatments. Slugs feeding on Codium, but not on Cladophora, showed higher relative electron transport rates (rETR) in high compared to low light. Furthermore, there were no differences in the consumption rates of the slugs between different light treatments, and only small differences in nutritional traits of algal diets, indicating that the increased growth efficiency of E. viridis feeding on Codium was due to retention of functional kleptoplasts. Our results show that functional kleptoplasts from Codium can provide sacoglossan sea slugs with fitness advantages through photosynthesis.

Factors determining the degree of dietary generalism versus specialism are central in ecology. Species that are generalists at the population level may in fact be composed of specialized individuals. The optimal diet theory assumes that individuals choose diets that maximize fitness, and individual specialization may occur if individuals' ability to locate, recognize, and handle different food types differ. We investigate if individuals of the marine herbivorous slug Elysia viridis, which co-occur at different densities on several green macroalgal species in the field, are specialized to different algal hosts. Individual slugs were collected from three original algal host species (Cladophora sericea, Cladophora rupestris and Codium fragile) in the field, and short-term habitat choice and consumption, as well as long-term growth (proxy for fitness), on four algal diet species (the original algal host species and Chaetomorpha melagonium) were studied in laboratory experiments. Nutritional (protein, nitrogen, and carbon content) and morphological (dry weight, and cell/utricle volume) algal traits were also measured to investigate if they correlated with the growth value of the different algal diets. E. viridis individuals tended to choose and consume algal species that were similar to their original algal host. Long-term growth of E. viridis, however, was mostly independent of original algal host, as all individuals reached a larger size on the non-host C. melagonium. E. viridis growth was positively correlated to algal cell/utricle volume but not to any of the other measured algal traits. Because E. viridis feeds by piercing individual algal cells, the results indicate that slugs may receive more cytoplasm, and thus more energy per unit time, on algal species with large cells/utricles. We conclude that E. viridis individuals are specialized on different hosts, but host choice in natural E. viridis populations is not determined by the energetic value of seaweed diets as predicted by the ODT.

A number of studies have shown that the production of chemical defences is costly in terrestrial vascular plants. However, these studies do not necessarily reflect the costs of defence production in macroalgae, due to structural and functional differences between vascular plants and macroalgae. Using a specific culturing technique, we experimentally manipulated the defence production in the red alga Bonnemaisonia hamifera to examine if the defence is costly in terms of growth. Furthermore, we tested if the defence provides fitness benefits by reducing harmful bacterial colonisation of the alga. Costly defences should provide benefits to the producer in order to be maintained in natural populations, but such benefits through protection against harmful bacterial colonisation have rarely been documented in macroalgae. We found that algae with experimentally impaired defence production, but with an externally controlled epibacterial load, grew significantly better than algae with normal defence production. We also found that undefended algae exposed to a natural epibacterial load experienced a substantial reduction in growth and a 6-fold increase in cell bleaching, compared to controls. Thus, this study provides experimental evidence that chemical defence production in macroalgae is costly, but that the cost is outweighed by fitness benefits provided through protection against harmful bacterial colonisation.