Molecular genetic diversity of seaweeds morphologically related to Ulva rigida at three sites along the French Atlantic coast

Foliose species of the genus Ulva are notoriously difficult to identify due to their variable morphological characteristics and high phenotypic plasticity. We reassessed the taxonomic status of several distromatic foliose Ulva spp., morphologically related to Ulva rigida, using DNA barcoding with the chloroplastic tufA and rbcL (for a subset of taxa) genes for 339 selected attached Ulva specimens collected from three intertidal rocky sites. Two of the collection sites were in Brittany and one site was in Vendée, along the Atlantic coast of France. Molecular analyses included several museum specimens and the holotype of Ulva armoricana Dion, Reviers & Coat. We identified five different tufA haplotypes using a combination of phylogenetic analysis, with the support of several recently sequenced holotypes and lectotypes, and a species delimitation method based on hierarchical clustering. Four haplotypes were supported by validly named species: Ulva australis Areschoug, Ulva fenestrata Postels & Ruprecht, Ulva lacinulata (Kützing) Wittrock and U. rigida C. Agardh. The later was additionally investigated using rbcL. The fifth haplotype represented exact sequence matches to an unnamed species from European Atlantic coasts. Our results support: (1) the synonymy of both U. rigida sensu Bliding non C. Agardh and U. armoricana with U. lacinulata. This finding is based on current genetic analysis of tufA from the U. armoricana holotype and recent molecular characterization of the lectotype of U. laetevirens, which is synonymous to U. australis, (2) the presence of U. australis as a misidentified introduced species in Brittany, and (3) the presence of U. fenestrata and U. rigida in southern Brittany. The taxonomic history of each species is discussed, highlighting issues within distromatic foliose taxa of the genus Ulva and the need to genetically characterize all its available type specimens.

Sustainable Large-Scale Aquaculture of the Northern Hemisphere Sea Lettuce, Ulva fenestrata, in an Off-Shore Seafarm

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.

Effects of irradiance, temperature, nutrients, and pCO2 on the growth and biochemical composition of cultivated Ulva fenestrata

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).

Cultivation conditions affect the monosaccharide composition in Ulva fenestrata

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.

Inducible versus constitutive antioxidant defenses in algae along an environmental stress gradient

Optimal defense theory (ODT) predicts that antiherbivore defenses should be constitutive when plants are frequently attacked and inducible when the probability of attack is low. Like antiherbivore defenses, antioxidant defenses can be inducible or constitutive. We hypothesized the ODT predictions should apply to antioxidant defenses; thus, species inhabiting environments where oxidative stresses occur frequently should produce constitutive antioxidant defenses, whereas species in environments where stresses occur less frequently should produce inducible defenses. We tested this hypothesis by attempting to induce production of the antioxidant precursor dimethylsulfoniopropionate (DMSP) in 4 ulvoid algae species that experience different levels of environmental stress because they are zoned along a tidal gradient. The 2 lower intertidal species Ulvaria obscura and Ulva fenestrata, which experience oxidative stresses less frequently, induced DMSP production in response to applications of the chemical oxidant hydrogen peroxide within 7 d, whereas the higher intertidal species Ulva linza and Ulva intestinalis, which regularly experience oxidative stress, did not have increased DMSP concentrations. This study demonstrates a novel waterborne signaling mechanism for DMSP induction in marine macroalgae and provides evidence of selection for inducible antioxidant defenses in organisms experiencing less frequent environmental stresses.