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Macroalgal nitrogen isotope analysis (δ 15 N) is a reliable method for the identification of nitrogen pollutant sources. Understanding δ 15 N geospatial variation within small bays and/or harbour environments can help identify point sources of nitrogen pollution. This study sampled over 300 Fucus vesiculosus and Ulva sp. specimens in September 2022 and May 2023 from Staithes Harbour, North Yorkshire, England. δ 15 N values for Staithes Beck were elevated when compared to sites in Staithes Harbour and the North Sea: this is attributed to sewage effluent and/or agricultural manure. Few sites within Staithes Harbour were significantly different from one another in terms of δ 15 N, suggesting a relatively homogenous nitrogen isotope record of the harbour. Simple harbour environments like Staithes may be relatively well mixed, and thus, sampling one harbour site may be enough to represent the entire harbour. Of course, more complex harbours may require more sample locations to ascertain point sources and mixing in the harbour., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Ulva are hardy green seaweeds that contain the sulfated polysaccharide ulvan and grow in two distinct morphologies: foliose and tubular. The authors hypothesise that ulvan from tubular species are more structurally complex than ulvans from foliose species. Herein, using standardised methods, the glycosyl linkage positions and sulfate ester substitutions of constituent monosaccharides of ulvan isolated from foliose (U. lacinulata and U. stenophylloides) and tubular (U. prolifera and U. ralfsii) species of Ulva were investigated. Comparison of native ulvans with 80 and 100 °C desulfated counterparts indicated that 4-linked rhamnose is predominantly 3-O-sulfated in all four ulvans. Ulvans from the foliose species predominantly contained →3,4)-Rhap-(1→, →4)-GlcAp-(1→ and →4)-IdoAp-(1→, collectively accounting for 67 to 81 mol% of the total linkages. In contrast, these same linkages in ulvans from the tubular species only collectively accounted for 29 to 36 mol%. Instead, ulvan from tubular species contained a combination of →2,3,4)-Rhap-(1→, terminal Rhap-(1→, →4)-GlcAp-(1→, →4)-Xylp-(1→, and/or →4)-Galp-(1→ in high proportions; some of the latter three residues were also likely O-2 sulfated. The results presented here suggest that ulvan from foliose species are predominantly unbranched polysaccharides composed of repeat disaccharides while ulvans from tubular species contain a greater diversity of branch and sulfate substitution locations., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Non-antibiotic substances have been found to contribute to the spread of antibiotic resistance. Bromophenols (BPs) are special anti-bacterial substances obtained from seaweed. This study explored the modulatory effect of trace BPs from a live seaweed on the antibiotic resistance of pathogenic Vibrio (V.) strains. A hydroponic solution of Ulva fasciata was found to contain trace levels (9-333 μg L -1 ) of 2,4,6-tribromophenol (TBP), a typical BP. TBP at a concentration of 165 μg L -1 significantly increased the inhibition zone diameter of widely used β-lactam antibiotics (amoxicillin and ampicillin) against V. alginolyticus M7 (Va. M7) and V. parahaemolyticus M3 (Vp. M3) as well as reduced the minimum inhibitory concentration by 2-4 fold against Va. M7. Whole genome re-sequencing analysis demonstrated that Va. M3 (53-60) had more mutant genes than Vp. M7 (44) in β-lactam resistance pathway. Transcriptome sequencing analysis, along with verification through RT-qPCR, further showed that oligopeptide permease (opp) was the only differentially expressed gene (DEG) among the mutated genes in the β-lactam resistance pathway. The opp transport activity and membrane permeability of Vibrio were both enhanced at 165 μg L -1 of TBP, and the ability of biofilm formation was also decreased. Thus, antibiotics resistance improvement of Vibrio by TBP was potentially related with the promoted opp transport activity, membrane permeability and inhibited biofilm formation., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Microplastics and other microscopic debris are a concern in the coastal environment but measurements in the water column and sediment are often problematic and rely on non-standardised and highly variable methodologies. To this end, we explore the potential of different species of temperate-cold marine macroalgae as passive biomonitors of anthropogenic microparticles at three contrasting locations in southwest England. Specifically, fronds from samples of fucoids and Ulva lactuca (n = 9 in total, and three from each location) have been sectioned and analysed directly under a microscope and anthropogenic microparticles counted and subsequently characterised for chemical composition. Microparticles were heterogeneously distributed throughout sections from the same sample. However, on a dry weight basis, combined microparticle concentrations for each sample ranged from about 7.5 g -1 to 110 g -1 , and from about 0.2 cm -2 to 0.9 cm -2 , and for a given species were higher in samples from a semi-enclosed harbour and urban beach than in samples from a protected beach facing the open sea. These values compare with published concentrations of microplastics and microfibres reported for the regional water column on the order of 0.1 m -3 . Most particles were cellulosic (e.g., rayon) and petroleum-based (mainly polyester and polyethylene terephthalate) fibres but plastic fragments were also present on most samples. Glass retroreflective beads derived from road markings were also present at up to 18 g -1 on fucoids from the urban beach because of its proximity to a stormwater effluent. Most microparticles were adhered to the smooth parts of the macroalgal surface but some displayed wrapping around edges and creases or entrapment by appendages. The practical and environmental implications of macroalgae passively capturing significant quantities of anthropogenic microparticles are discussed., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Main Conclusion: The combined photoinhibitory and PSII-reaction centre quenching against light stress is an important mechanism that allows the green macroalga Ulva rigida to proliferate and form green tides in coastal ecosystems. Eutrophication of coastal ecosystems often stimulates massive and uncontrolled growth of green macroalgae, causing serious ecological problems. These green tides are frequently exposed to light intensities that can reduce their growth via the production of reactive oxygen species (ROS). To understand the physiological and biochemical mechanisms leading to the formation and maintenance of green tides, the interaction between inorganic nitrogen (N i ) and light was studied. In a bi-factorial physiological experiment simulating eutrophication under different light levels, the bloom-forming green macroalga Ulva rigida was exposed to a combination of ecologically relevant nitrate concentrations (3.8-44.7 µM) and light intensities (50-1100 µmol photons m -2  s -1 ) over three days. Although artificial eutrophication (≥ 21.7 µM) stimulated nitrate reductase activity, which regulated both nitrate uptake and vacuolar storage by a feedback mechanism, nitrogen assimilation remained constant. Growth was solely controlled by the light intensity because U. rigida was N i -replete under oligotrophic conditions (3.8 µM), which requires an effective photoprotective mechanism. Fast declining Fv/Fm and non-photochemical quenching (NPQ) under excess light indicate that the combined photoinhibitory and PSII-reaction centre quenching avoided ROS production effectively. Thus, these mechanisms seem to be key to maintaining high photosynthetic activities and growth rates without producing ROS. Nevertheless, these photoprotective mechanisms allowed U. rigida to thrive under the contrasting experimental conditions with high daily growth rates (12-20%). This study helps understand the physiological mechanisms facilitating the formation and persistence of ecologically problematic green tides in coastal areas., (© 2024. The Author(s).)

This study assesses the health risk due to heavy metals/metalloids (HMs/Ms) in edible seaweeds (Caulerpa racemosa, Kappaphycus alvarezii, and Ulva lactuca) through an in vitro bioaccessibility study. The percentage of bioabsorbed HMs/Ms in unprocessed and processed C. racemosa, U. lactuca, and K. alvarezii ranged from 3 % to 46 %, 3 % to 42 %, and 3 % to 40 %, respectively. The levels of HMs/Ms in seawater, sediment, and seaweeds were below the levels recommended by the European Commission (EC) and World Health Organization/Food and Agriculture Organization (WHO/FAO). The maximum accumulation of HMs/Ms was found during monsoons and post-monsoon seasons, and Cd, Pb, Hg, Cr, As, and Pb were predominant in all the samples. Tukey's post hoc test and t-test confirmed that thermal processing significantly reduced HMs/Ms in seaweeds. On the basis of the bioabsorption of HMs/Ms, the TTHQ values were found to be < 1, and the LCR values were within the acceptable limit (10 -06 to 10 -04 ), indicating no carcinogenic risks through seaweeds., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

The sea lettuce Ulva spp is becoming an increasingly important macroalgae for aquaculture. Sea lettuce can be grown on- and off-shore, displays high growth rates, and its biomass possesses attractive nutritional benefits. Among those are their fatty acids (FA) and lipid profiles, rich in omega 3 Polyunsaturated Fatty Acids (PUFAs) as well as bioactive lipids. In order to tailor those properties for food applications, we explored the use of a short-term (seven days) low salinity treatment to modulate the lipid profile of two species of Ulva. We found large quantitative differences between species, and while a low-salinity treatment negatively affected growth, Ulva australis' lipid profile was positively impacted. Total FA particularly ɷ-3 PUFAs, increased three-fold, as well as most polar lipid species including known bioactive compounds. This study highlights profound differences between species and describes a simple method to increase the nutritional properties of Ulva biomass for food applications., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

A fully enantioselective, catalytic synthesis of the algal morphogen (-)-thallusin using polyene cyclization chemistry is reported. The synthesis features dedicated precursor design, introduction of a TMS-substituted arene as a regioselective terminator, very high enantiomer excess (ee) on gram scale, and productive scaffold functionalization. Furthermore, an ee determination methodology of thallusin samples was developed, and the ee of biosynthesized thallusin was determined. Fe(III)-uptake studies demonstrated that the cellular uptake of iron facilitated by thallusin derivatives was independent of their morphogenic activity, suggesting their active import via siderophore transporters as a shuttle system., (© 2024 The Authors. Chemistry - A European Journal published by Wiley-VCH GmbH.)

Green tides occurrence has increased in coral reefs, yet few reference values have been documented to support bloom management in these ecosystems. Here, we took advantage of recent Ulva green tides that occurred in New Caledonia to (i) identify the elements limiting the growth of Ulva spp. during these blooms; and (ii) validate the use of isotopic markers for identifying sources of nutrients that generated blooms. N/P ratios highlighted a stronger limitation of algae by phosphorus than by nitrogen on sites under oceanic influence, while the proportions of N and P were optimal for algal growth at sites where green tides occurred. Macroalgae highly exposed to sewage water was characterized by higher δ 15 N than macroalgae collected in areas exposed to synthetic inorganic fertilizers. From these results, we established a new set of threshold values for using δ 15 N in Ulva species as an indicator of nitrogen source type in coral reefs., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024. Published by Elsevier Ltd.)

Background: Solar radiation is primarily composed of ultraviolet radiation (UVR, 200 - 400 nm) and photosynthetically active radiation (PAR, 400 - 700 nm). Ultraviolet-B (UVB) radiation accounts for only a small proportion of sunlight, and it is the primary cause of plant photodamage. The use of chlorofluorocarbons (CFCs) as refrigerants caused serious ozone depletion in the 1980s, and this had led to an increase in UVB. Although CFC emissions have significantly decreased in recent years, UVB radiation still remains at a high intensity. UVB radiation increase is an important factor that influences plant physiological processes. Ulva prolifera, a type of macroalga found in the intertidal zone, is intermittently exposed to UVB. Alternative oxidase (AOX) plays an important role in plants under stresses. This research examines the changes in AOX activity and the relationships among AOX, photosynthesis, and reactive oxygen species (ROS) homeostasis in U. prolifera under changes in UVB and photosynthetically active radiation (PAR)., Results: UVB was the main component of solar radiation impacting the typical intertidal green macroalgae U. prolifera. AOX was found to be important during the process of photosynthesis optimization of U. prolifera due to a synergistic effect with non-photochemical quenching (NPQ) under UVB radiation. AOX and glycolate oxidase (GO) worked together to achieve NADPH homeostasis to achieve photosynthesis optimization under changes in PAR + UVB. The synergism of AOX with superoxide dismutase (SOD) and catalase (CAT) was important during the process of ROS homeostasis under PAR + UVB., Conclusions: AOX plays an important role in the process of photosynthesis optimization and ROS homeostasis in U. prolifera under UVB radiation. This study provides further insights into the response of intertidal macroalgae to solar light changes., (© 2024. The Author(s).)

Seaweeds, including the green Ulva lactuca, can potentially reduce competition between feed, food, and fuel. They can also contribute to the improved development of weaned piglets. However, their indigestible polysaccharides of the cell wall pose a challenge. This can be addressed through carbohydrase supplementation, such as the recombinant ulvan lyase. The objective of our study was to assess the muscle metabolism of weaned piglets fed with 7% U. lactuca and 0.01% ulvan lyase supplementation, using an integrated transcriptomics (RNA-seq) and proteomics (LC-MS) approach. Feeding piglets with seaweed and enzyme supplementation resulted in reduced macronutrient availability, leading to protein degradation through the proteasome (PSMD2), with resulting amino acids being utilized as an energy source (GOT2, IDH3B). Moreover, mineral element accumulation may have contributed to increased oxidative stress, evident from elevated levels of antioxidant proteins like catalase, as a response to maintaining tissue homeostasis. The upregulation of the gene AQP7, associated with the osmotic stress response, further supports these findings. Consequently, an increase in chaperone activity, including HSP90, was required to repair damaged proteins. Our results suggest that enzymatic supplementation may exacerbate the effects observed from feeding U. lactuca alone, potentially due to side effects of cell wall degradation during digestion., (© 2024. The Author(s).)

In this study, effective biomaterials were prepared from marine macroalgae, namely Fucus spiralis (F.S), Ulva intestinalis (U.I), and Corallina officinalis (C.O). The ability to adsorb the hazardous organic dye crystal violet (CV) was examined, revealing different adsorptive properties for the three algae. The removal of CV dye occurred onto only a homogeneous monolayer for F.S, and both a homogeneous monolayer and a heterogeneous multilayer for U.I and C.O algae. The predicted monolayer capacities at 25 °C were approximately 53 mg/g, 55 mg/g, and 97 mg/g for F.S, C.O, and U.I, respectively. The adsorption of CV dye on all the algae was found to follow a pseudo-second-order rate. Ulva intestinalis algae, as a potential adsorbent of CV dye, were also tested in the adsorption of inorganic substances and demonstrated significant efficiency in the removal of chromium (VI). The findings highlight various adsorption properties and the relevance of macroalgae for wastewater treatment applications., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Marine macroalgae (seaweeds) are important primary global producers, with a wide distribution in oceans around the world from polar to tropical regions. Most of these species are exposed to variable environmental conditions, such as abiotic (e.g., light irradiance, temperature variations, nutrient availability, salinity levels) and biotic factors (e.g., grazing and pathogen exposure). As a result, macroalgae developed numerous important strategies to increase their adaptability, including synthesizing secondary metabolites, which have promising biotechnological applications, such as UV-absorbing Mycosporine-Like Amino Acid (MAAs). MAAs are small, water-soluble, UV-absorbing compounds that are commonly found in many marine organisms and are characterized by promising antioxidative, anti-inflammatory and photoprotective properties. However, the widespread use of MAAs by humans is often restricted by their limited bioavailability, limited success in heterologous expression systems, and low quantities recovered from the natural environment. In contrast, bloom-forming macroalgal species from all three major macroalgal clades (Chlorophyta, Phaeophyceae, and Rhodophyta) occasionally form algal blooms, resulting in a rapid increase in algal abundance and high biomass production. This review focuses on the bloom-forming species capable of producing pharmacologically important compounds, including MAAs, and the application of proteomics in facilitating macroalgal use in overcoming current environmental and biotechnological challenges.

Macroalgal biomass being rich in carbohydrates, proteins and lipids in their cell wall has been considered as the most efficient organic rich sources for biofuel (biohydrogen) production. In this study, Pluronic P-123-induced microwave pretreatment was applied to disintegrate the marine macroalgae biomass, Ulva reticulata. Microwave disintegration was done by varying the treatment time and microwave power from 0 to 40 min and 0.09 KW to 0.63 KW. A maximum chemical oxygen demand (COD) solubilization of 22.33% was achieved at a microwave power and time duration of 0.36 kW and 15 min. Chemical (Pluronic P-123, a mild surfactant) was combined with optimum microwave disintegration conditions to increase the solubilization efficiency and this combined pretreatment achieved a maximum COD solubilization of 31.02% at 10 min pretreatment time and 0.06 g per g TS of Pluronic P-123 dosage. The present study indicated that combination of surfactant with microwave pretreatment substantially improves the COD solubilization with reduced pretreatment -time than mono microwave pretreatment. An optimal hydrogen yield of 98.37 mL was achieved through this combined pretreatment. The biohydrogen data was modelled with Gompertz model and the kinetic parameters derived through this model implies that the calculated adjusted R squared values for all the samples lies between 0.95 and 0.99. This shows that the model fitted biohydrogen experimental values accurately. In addition, Pluronic P-123-induced microwave pretreatment was regarded as energy efficient and cost effective than microwave pretreatment alone with net energy production and a greater energy ratio of 504.38 kWh/Ton macroalgae and 1.2 when compared to microwave pretreatment alone (-2975.6 kWh/Ton macroalgae and 0.5)., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Inc. All rights reserved.)

Offshore macroalgae production could provide an alternative source of biomass for food, materials and energy. However, the offshore environment in general, specifically the Eastern Mediterranean Sea (EMS) offshore, is a high energy and low nutrients environment, thus challenging for macroalgae farming. In this study, we experimentally investigated the impact of season, depth, and pre-cultivation fertilization duration on the growth rates and chemical composition of offshore Ulva biomass, and developed a predictive model tailored to offshore conditions, capable of estimating both biomass growth rate and nitrogen content. Specifically, we measured Ulva biomass growth rate and internal nitrogen in the nitrogen-poor EMS a few kilometers offshore the Israeli coast at various depths and on-shore pre-cultivation fertilization schedules. Based on these data, we constructed a predictive cultivation model of Ulva offshore growth, which allows for the optimization of fertilization requirements for offshore cultivation. This study provides new insights on the effects of seasonality, depth, and pre-cultivation fertilization duration on growth rates and chemical composition of offshore Ulva sp. biomass production., (© 2023. Springer Nature Limited.)

In this study, response surface methodology (RSM) was applied with a Box-Behnken design to optimize the biosorption (removal and bioconcentration) of rare earth elements (REEs) (Y, La, Ce Eu, Gd, Tb) by living Ulva sp. from diluted industrial wastewaters (also containing Pt and the classic contaminants Hg, Pb, Zn, Cu, Co, and Cd). Element concentration (A: 10-190 μg/L), wastewater salinity (B: 15-35), and Ulva sp. dosage (C: 1.0-5.0 g/L) were the operating parameters chosen for optimization. Analysis of the Box-Behnken central point confirmed the reproducibility of the methodology and p-values below 0.0001 validated the developed mathematical models. The largest inter-element differences were observed at 24 h, with most REEs, Cu, Pb and Hg showing removals ≥ 50 %. The factor with the greatest impact (positive) on element removal was the initial seaweed dosage (ANOVA, p < 0.05). The optimal conditions for REEs removal were an initial REEs concentration of 10 μg/L, at a wastewater salinity of 15, and an Ulva sp. dosage of 5.0 g/L, attaining removals up to 88 % in 24 h. Extending the time to 96 h allowed seaweed dosage to be reduced to 4.2 g/L while achieving removals ≥ 90 %. The high concentrations in REE-enriched biomass (∑REEs of 3222 μg/g), which are up to 3000 times higher than those originally found in water and exceed those in common ores, support their use as an alternative source of these critical raw materials., (© 2023. The Author(s).)

Optical remote sensing provides optimal technical support for the detection and quantification of floating macroalgae. Although the spatial scale effect on optical estimation of floating macroalgae coverage or biomass from different images has been clarified, the directional effect on them has not been investigated until now. In this study, synchronous multi-angle imaging spectroradiometer (MISR) and MODIS images were collected to investigate the multi-angle remote sensing of green tides. A dual thresholding method, based on the difference vegetation index (DVI) and scaled algae index, was employed to determine algae pixels. In addition, piecewise empirical models were developed for MISR and MODIS images to estimate the total biomass of green tides based on laboratory measurements and DVI values. Comparative analysis of DVI histograms and total biomass shows that the sensor zenith angle has a significant impact on the quantification of green tides. Under the same solar conditions, as the sensor zenith angle increases, the optical signals received from algae pixels weaken, resulting in a decrease in the quantification of green tides. In future research, the observation geometry (including the solar/sensor zenith angle and the solar/sensor azimuth angle) needs to be considered to improve the accuracy of optical remote detection and quantification of floating macroalgae.

A bioaccumulation study in red (Palmaria palmata) and green (Ulva sp.) seaweed has been carried out after exposure to different concentrations of citrate-coated titanium dioxide nanoparticles (5 and 25 nm) for 28 days. The concentration of total titanium and the number and size of accumulated nanoparticles in the seaweeds has been determined throughout the study by inductively coupled plasma mass spectrometry (ICP-MS) and single particle-ICP-MS (SP-ICP-MS), respectively. Ammonia was used as a reaction gas to minimize the effect of the interferences in the 48 Ti determination by ICP-MS. Titanium concentrations measured in Ulva sp. were higher than those found in Palmaria palmata for the same exposure conditions. The maximum concentration of titanium (61.96 ± 15.49 μg g -1 ) was found in Ulva sp. after 28 days of exposure to 1.0 mg L -1 of 5 nm TiO 2 NPs. The concentration and sizes of TiO 2 NPs determined by SP-ICP-MS in alkaline seaweed extracts were similar for both seaweeds exposed to 5 and 25 nm TiO 2 NPs, which indicates that probably the element is accumulated in Ulva sp. mainly as ionic titanium or nanoparticles smaller than the limit of detection in size (27 nm). The implementation of TiO 2 NPs in Ulva sp. was confirmed by electron microscopy (TEM/STEM) in combination with energy dispersive X-Ray analysis (EDX)., (© 2023. The Author(s).)

Ulva lactuca is a marine green seaweed. Its bloom based biomass accumulated in the İzmir bay and is collected by local authorities. In this investigation, an alternative solution was proposed to utilize the biomass of U. lactuca to produce biohydrogen via green synthesized silver nanoparticles. According to the results, the optimum conditions related to silver nanoparticle production such as pH, temperature, biomass concentration, silver nitrate concentrations, and incubation time were determined to be 11, 25 °C, 10 mg/mL, 4 mM, and 3 days, respectively. Effective conditions for biohydrogen production such as pH, temperature, agitation rate, and sodium borohydride concentration were found to be 7, 50 °C, 250 rpm and 150 mM, respectively. These parameters are also modelled with an artificial neural network. The data presented here provide recommendations for producing biohydrogen from waste algae and helping reduce carbon emissions for better environment and future., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Green seaweeds exhibit a wide range of morphologies and occupy various ecological niches, spanning from freshwater to marine and terrestrial habitats. These organisms, which predominantly belong to the class Ulvophyceae, showcase a remarkable instance of parallel evolution toward complex multicellularity and macroscopic thalli in the Viridiplantae lineage. Within the green seaweeds, several Ulva species ("sea lettuce") are model organisms for studying carbon assimilation, interactions with bacteria, life cycle progression, and morphogenesis. Ulva species are also notorious for their fast growth and capacity to dominate nutrient-rich, anthropogenically disturbed coastal ecosystems during "green tide" blooms. From an economic perspective, Ulva has garnered increasing attention as a promising feedstock for the production of food, feed, and biobased products, also as a means of removing excess nutrients from the environment. We propose that Ulva is poised to further develop as a model in green seaweed research. In this perspective, we focus explicitly on Ulva mutabilis/compressa as a model species and highlight the molecular data and tools that are currently available or in development. We discuss several areas that will benefit from future research or where exciting new developments have been reported in other Ulva species., (© 2023 Phycological Society of America.)

Background: Macroalgal epiphytic microbial communities constitute a rich resource for novel enzymes and compounds, but studies so far largely focused on tag-based microbial diversity analyses or limited metagenome sequencing of single macroalgal species., Results: We sampled epiphytic bacteria from specimens of Ulva sp. (green algae), Saccharina sp. (brown algae), Grateloupia sp. and Gelidium sp. (both red algae) together with seawater and sediment controls from a coastal reef in Weihai, China, during all seasons. Using 16S rRNA amplicon sequencing, we identified 14 core genera (consistently present on all macroalgae), and 14 dominant genera (consistently present on three of the macroalgae). Core genera represented ~ 0.7% of all genera, yet accounted for on average 51.1% of the bacterial abundances. Plate cultivation from all samples yielded 5,527 strains (macroalgae: 4,426) representing 1,235 species (685 potentially novel). Sequencing of selected strains yielded 820 non-redundant draft genomes (506 potentially novel), and sequencing of 23 sampled metagenomes yielded 1,619 metagenome-assembled genomes (MAGs), representing further 1,183 non-redundant genomes. 230 isolates and 153 genomes were obtained from the 28 core/dominant genera. We analyzed the genomic potential of phycosphere bacteria to degrade algal polysaccharides and to produce bioactive secondary metabolites. We predicted 4,451 polysaccharide utilization loci (PULs) and 8,810 biosynthetic gene clusters (BGCs). These were particularly prevalent in core/dominant genera., Conclusions: Our metabolic annotations and analyses of MAGs and genomes provide new insights into novel species of phycosphere bacteria and their ecological niches for an improved understanding of the macroalgal phycosphere microbiome. Video Abstract., (© 2023. The Author(s).)

Preservative is of importance to retard fruit deterioration and prolong the shelf-life. The suitability of using water-soluble polysaccharide extracted from waste macroalgae Enteromorpha prolifera (EPP) for cherry tomato preservation was evaluated. As compared with the control, the EPP-treated cherry tomatoes exhibited better fruit appearance, lower disease index and rot index during storage. Around 47 % EPP-treated cherry tomatoes were commercially acceptable after 36 days, which was however only 15.6 % for untreated cherry tomatoes, indicating the satisfactory preservation effectiveness of EPP-rich solution for cherry tomatoes. The post-extraction residue was commonly underutilized, we herein attempted to employ an emerging thermochemical conversion technique, hydrothermal liquefaction, to produce crude bio-oil (biocrude) from post-extraction E. prolifera. A biocrude yield of ∼23 wt% (dry-ash-free, daf) was obtained, and fatty acids and phenolics were identified to be the two main components in biocrude. The biocrude contained ∼70 % carbon and the higher heating value was ∼30 MJ/kg., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Ulva prolifera macroalgae blooming caused by water eutrophication seriously affects the marine ecological environment. Exploring an efficient approach to turning algae biomass waste into high-value-added products is significant. The present work aimed to demonstrate the feasibility of the bioactive polysaccharide extraction from Ulva prolifera and to evaluate its potential biomedical application. A short autoclave process was proposed and optimized using the response surface methodology to extract Ulva polysaccharides (UP) with high molar mass. Our results indicated that UP with high molar mass (9.17 × 10 5  g/mol) and competitive radical scavenging activity (up to 53.4 %) could be effectively extracted with the assistance of Na 2 CO 3 (1.3 %, wt.) at a solid-liquid ratio of 1/10 in 26 min. The obtained UP mainly composes of galactose (9.4 %), glucose (73.1 %), xylose (9.6 %), and mannose (4.7 %). The biocompatibility of the UP and its potential application as a bioactive ingredient in 3D cell culture has been evaluated and confirmed by confocal laser scanning microscopy and fluorescence microscope imaging inspection. This work demonstrated the feasibility of extracting bioactive sulfated polysaccharides with potential applications in biomedicine from biomass waste. Meanwhile, this work also provided an alternative solution to deal with the environmental challenges incurred by algae blooming worldwide., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Characterization of biochar-derived dissolved organic matter (DOM) can provide deep insight into potential applications of biochar. Herein, biochar from six macroalgae (Phaeophyta-Sargassum fusiforme, Sargassum thunbergii, and Sargassum vachellianum; Rhodophyta-Grateloupia turuturu and Chondria crassicaulis; and Chlorophyta-Ulva pertusa) were subjected to pyrolysis at different temperatures (200 °C-500 °C). The effects of pyrolysis temperature and extraction solution pH on the characteristics of the macroalgal biochar-derived DOM (MBDOM) were investigated via fluorescence excitation-emission matrix spectroscopy with parallel factor (PARAFAC) analysis. Five humic-like substances and one protein-like substance were identified. The distributions of the six PARAFAC components depended on the macroalgae species, pyrolysis temperature, and extraction solution pH. The proportion of the protein-like substance (0 %-46.77 %) was less than that of the humic-like substances (100 %-53.23 %) in a given MBDOM regardless of the extraction solution pH values. Fluorescence spectral indicators show that DOM from macroalgal biochar is more autochthonous and humified than that from the corresponding biomass. Hierarchical cluster analysis and redundancy analysis results further show that the macroalgae species, pyrolysis temperature, and extraction solution pH jointly affect DOM characteristics with varying contribution levels., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Metal pollution in the marine coastal environment is an important topical issue. In this study, the water quality in five locations along the Alexandria coast (Eastern Harbor, El-Tabia pumping station, El Mex Bay, Sidi Bishir, and Abu Talat) was assessed by measuring physicochemical parameters from water samples. Depending on the morphological classification of macroalgae, the collected morphotypes were related to Ulva fasciata, Ulva compressa, Corallina officinalis, Corallina elongata, and Petrocladia capillaceae. Corallina officinalis and Corallina elongata demonstrated a high capacity for Cd, Pb, and Ni accumulation, and the highest values of Fe, Cu, and Mn were reported in Ulva fasciata and Ulva compressa. Two standard markers were applied, and results showed that the morphological classification matched the molecular data. Moreover, the analysis of algae can only reflect the accumulation of metals. The conclusion is that Ulva compressa and Corallina officinalis are potentially suitable indicators of localized short-term heavy metal pollution., Competing Interests: Declaration of competing interest The authors declare that they have no conflict of interest., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Mercury (Hg) is a global and top priority contaminant, toxic at low concentrations. Although it has been progressively eliminated from processes, this metal continues to circulate in the atmosphere, soil, and water. In this work, the Response Surface Methodology (RSM) combined with a Box-Behnken Design (3 factors - 3 levels) was used to optimize key operational conditions that influence the removal and uptake of Hg by living macroalga Ulva sp. in a complex mixture containing several elements used in industry (potentially toxic elements, rare earth elements, and platinum-group elements) (initial concentration 10, 100 and 190 µg/L, salinity 15, 25 and 35, seaweed stock density 1.0, 3.0 and 5.0 g/L). Results evidenced the great capability of Ulva sp. to remove Hg, with removal efficiencies between 69 % and 97 %. 3-D surfaces showed that the most impactful variable was seaweed stock density, with higher densities leading to higher removal. Regarding the uptake, a positive correlation between initial concentration and q t values was observed. The appliance of RSM made possible to obtain optimal operating conditions for removing virtually 100 % of Hg from waters with high ionic strength, which is a pivotal step in the direction of the application of this remediation biotechnology at large scale., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 The Authors. Published by Elsevier B.V. All rights reserved.)

As a result of the wide use of Titanium (Ti) compounds in various products, Ti and Ti nanoparticles (nTi) are released into aquatic environments, inducing varying degrees of toxicity on aquatic fauna. Ulva lactuca, green macroalgae commonly found in coastal areas, has been extensively studied due to its worldwide distribution and capacity to accumulate trace elements under toxic conditions, which makes it a good universal sorbent. The present study aimed to establish the remediation properties of U. lactuca by evaluating the toxicity of Ti and nTi in bivalves, in the presence and absence of algae. Using the bivalve species Mytilus galloprovincialis, Ti toxicity was evaluated by assessing changes in mussel's metabolic capacity and oxidative status. Results evidenced cellular damage in M. galloprovincialis exposed to Ti and nTi. This was a result of the inactivation of antioxidant defences. The presence of U. lactuca limited cellular damage, however, this was not a result of the previously demonstrated bioremediation capacity, as no accumulation of Ti was verified in algal tissues. As a metabolic depression was verified for mussels exposed to Ti/nTi in the presence of algae, we hypothesise that U. lactuca may have been responsible for changes to the water quality which induced this response., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 The Author(s). Published by Elsevier B.V. All rights reserved.)

Unravelling the structure-function variation of phycospheric microorganisms and its ecological correlation with harmful macroalgal blooms (HMBs) is a challenging research topic that remains unclear in the natural dynamic process of HMBs. During the world's largest green tide bloom, causative macroalgae Ulva prolifera experienced dramatic changes in growth state and environmental conditions, providing ideal scenarios for this investment. Here, we assess the phycospheric physicochemical characteristics, the algal host's biology, the phycospheric bacterial constitutive patterns, and the functional potential during the U . prolifera green tide. Our results indicated that (i) variation in the phycosphere nutrient structure was closely related to the growth state of U . prolifera ; (ii) stochastic processes govern phycospheric bacterial assembly, and the contribution of deterministic processes to assembly varied among phycospheric seawater bacteria and epiphytic bacteria; (iii) phycospheric seawater bacteria and epiphytic bacteria exhibited significant heterogeneity variation patterns in community composition, structure, and metabolic potential; and (iv) phycospheric bacteria with carbon or nitrogen metabolic functions potentially influenced the nutrient utilization of U . prolifera . Furthermore, the keystone genera play a decisive role in the structure-function covariation of phycospheric bacterial communities. Our study reveals complex interactions and linkages among environment-algae-bacterial communities which existed in the macroalgal phycosphere and highlights the fact that phycospheric microorganisms are closely related to the fate of the HMBs represented by the green tide. IMPORTANCE Harmful macroalgal blooms represented by green tides have become a worldwide marine ecological problem. Unraveling the structure-function variation of phycospheric microorganisms and their ecological correlation with HMBs is challenging. This issue is still unclear in the natural dynamics of HMBs. Here, we revealed the complex interactions and linkages among environment-algae-bacterial communities in the phycosphere of the green macroalgae Ulva prolifera , which causes the world's largest green tides. Our study provides new ideas to increase our understanding of the variation patterns of macroalgal phycospheric bacterial communities and the formation mechanisms and ecological effects of green tides and highlights the importance of phycospheric microorganisms as a robust tool to help understand the fate of HMBs.

Green tides have occurred in the Southern Yellow Sea (SYS) for 16 consecutive years, causing widespread concern. Attached and floating green macroalgae have been observed in Binhai Harbor, Jiangsu Province, China, in the SYS. This study used morphological analysis, and internal transcribed spacers and rps2-trnL molecular identification methods, to analyze the species composition and biomass of green macroalgae along the Binhai Harbor coast. Six species of green tide algae (Ulva prolifera, Ulva meridionalis, Ulva linza, Ulva flexuosa, Ulva californica, and Ulva intestinalis) were identified, in addition to Blidingia sp. The discovery of U. californica is the first report of this species off the coast of Jiangsu Province. The floating green macroalgae along the Binhai Harbor coast originated from attached green macroalgae in Binhai Harbor, and a small number of the attached algae were closely related to the large-scale floating U. prolifera in the SYS. Moreover, in December 2021, February 2022, and April 2022, the total biomass of attached green macroalgae in Binhai Harbor was 25.600, 10.767, and 25.867 t, respectively, of which the U. prolifera biomass was 10.697, 8.709, and 4.185 t, respectively. This study proved Binhai Harbor may not be an important source of green tide in the SYS., Competing Interests: Declaration of competing interest We declare that there is no conflict of interest in the article., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Chronic, low-grade inflammation is linked to the development of non-communicable diseases, including cancer, cardiovascular disease, obesity, insulin resistance, diabetes, and others which together contribute to more than 50% of deaths globally. Modulation of inflammatory responses may be a promising strategy, and n-3 long chain polyunsaturated fatty acids (n-3 LC-PUFA) may offer a new therapeutic option in inflammatory conditions. Seaweeds are characterised by high nutritional quality and are a good source of many bioactive compounds, including n-3 LC-PUFA. This review addresses the potential anti-inflammatory properties of seaweed derived lipids, and their immunomodulating mechanisms in order to identify the possible applications of seaweed as an anti-inflammatory functional food ingredient or dietary supplement. A few studies have evaluated the anti-inflammatory activity of seaweed lipids using crude lipid extracts, lipid fractions and isolated complex lipids from several seaweeds belonging to the Ochrophyta and Rhodophyta phyla, with only three Ulva rigida, Ulva sp. and Codium tomentosum within the Chlorophyta phylum. It was reported that seaweed derived lipids suppress inducible nitric oxide synthase and cyclooxygenase-2 expression and reduce nuclear factor κB p100 and myeloid differentiation primary response 88 protein levels leading to the downregulation of the production of several pro-inflammatory cytokines and nitric oxide. Further investigations are required to unravel the complex mechanisms underlying their preventive action against chronic inflammation and their potential use as a new functional food ingredient and/or health supplement.

Managing diabetes is challenging due to the complex physiology of the disease and the numerous complications associated with it. As part of the ongoing search for antidiabetic chemicals, marine algae have been demonstrated to be an excellent source due to their medicinal properties. In this study, Ulva reticulata extracts were investigated for their anti-diabetic effect by examining its inhibitory effects on α-amylase, α-glucosidase, and DPP-IV and antioxidant (DPPH) potential in vitro and its purified fraction using animal models. Among the various solvents used, the Methanolic extract of Ulva reticulata (MEUR) displayed the highest antidiabetic activity in both in vitro and in vivo; it showed no cytotoxicity and hence was subjected to bioassay-guided chromatographic separation. Among the seven isolated fractions (F1 to F7), the F4 (chloroform) fraction exhibited substantial total phenolic content (65.19 μg mL -1 ) and total flavonoid content (20.33 μg mL -1 ), which showed the promising inhibition against α-amylase (71.67%) and α-glucosidase (38.01%). Active fraction (F4) was further purified using column chromatography, subjected to thin-layer chromatography (TLC), and characterized by spectroscopy techniques. Upon structural elucidation, five distinct compounds, namely, Nonane, Hexadecanoic acid, 1-dodecanol, Cyclodecane methyl, and phenol, phenol, 3,5-bis(1,1-dimethylethyl) were identified. The antidiabetic mechanism of active fraction (F4) was further investigated using various in vitro and in vivo models. The results displayed that in in vitro both 1 and 24 h in vitro cultures, the active fraction (F4) at a concentration of 100 μg mL -1 demonstrated maximum glucose-induced insulin secretion at 4 mM (0.357 and 0.582 μg mL -1 ) and 20 mM (0.848 and 1.032 μg mL -1 ). The active fraction (F4) reduces blood glucose levels in normoglycaemic animals and produces effects similar to that of standard acarbose. Active fraction (F4) also demonstrated outstanding hypoglycaemic activity in hyperglycemic animals at a dose of 10 mg/kg B.wt. In the STZ-induced diabetic rat model, the active fraction (F4) showed a (61%) reduction in blood glucose level when compared to the standard drug glibenclamide (68%). The results indicate that the marine algae Ulva reticulata is a promising candidate for managing diabetes by inhibiting carbohydrate metabolizing enzymes and promoting insulin secretion.

Marine macroalgae are considered an untapped source of healthy natural metabolites and their market demand is rapidly increasing. Intertidal macroalgae present chemical defense mechanisms that enable them to thrive under changing environmental conditions. These intracellular chemicals include compounds that can be used for human benefit. The aim of this study was to test cultivation protocols that direct seaweed metabolic responses to enhance the production of target antioxidant and photoprotective biomaterials. We present an original integrated multi-trophic aquaculture (IMTA) design, based on a two-phase cultivation plan, in which three seaweed species were initially fed by fish effluents, and subsequently exposed to various abiotic stresses, namely, high irradiance, nutrient starvation, and high salinity. The combined effect of the IMTA's high nutrient concentrations and/or followed by the abiotic stressors enhanced the seaweeds' content of mycosporine-like amino acids (MAAs) by 2.3-fold, phenolic compounds by 1.4-fold, and their antioxidant capacity by 1.8-fold. The Sun Protection Factor (SPF) rose by 2.7-fold, and the chlorophyll and phycobiliprotein synthesis was stimulated dramatically by an order of magnitude. Our integrated cultivation system design offers a sustainable approach, with the potential to be adopted by emerging industries for food and health applications.

This study examined the antiproliferative and apoptotic-inducing effects of Ecklonia maxima (KP) and Ulva rigida (URL) extracts in the human liver cancer (HepG2) cell line model. HepG2 cells were cultured and grown in an incubator (5% CO 2 ) at 37°C. Cell viability was determined, while the effect of the extracts on apoptosis, ROS production, mitochondria membrane potential, and antioxidant enzymes were also assessed. KP and URL induced cytotoxic effects on HepG2 cells at the concentrations tested (0-1000 μg/ml). The morphological characteristics of the cells after treatment with KP and URL revealed cell shrinkage of the nucleus, cell injury, and damage compared to the control. The fluorescent micrographs from the apoptotic assay revealed induction of apoptosis and necrosis in HepG2 cells after treatment with KP and URL (200 and 400 μg/ml). The extracts also induced ROS production and reduced mitochondria membrane potential in HepG2 cells. The apoptotic-inducing effects, activation of ROS generation, and disruption of antioxidant enzymes are associated with the cytotoxic effects of the seaweed extracts. KP and URL showed good anticancer properties and could be explored as a good source of nutraceuticals, food additives, and dietary supplements to prevent uncontrolled proliferation of HepG2 cells. PRACTICAL APPLICATIONS: Seaweeds are reservoirs of nutrients and naturally occurring biologically active compounds, including sterols, phlorotannins, and polyunsaturated fatty acids. Due to the presence of these compounds, they are used as emulsifying agents, nutraceuticals, and additives in functional foods. Evidence suggests that seaweed bioactives may inhibit uncontrolled cell proliferation and induce apoptosis in cancer cells. Hence, exploring the antiproliferative and apoptotic-inducing effects of Ecklonia maxima and Ulva rigida will provide insights into their anticancer potentials as functional foods and nutraceuticals., (© 2022 The Authors. Journal of Food Biochemistry published by Wiley Periodicals LLC.)

In this study, the volatile compounds fingerprinting as well as the relationship between phytochemical composition and antioxidant bioactivity of U. prolifera, U. linza and U. clathrata were investigated. The significant differences of volatile compounds in three dehydrated seaweeds were observed by headspace gas chromatography-ion mobility spectrometry (HS-GC-IMS), and the type of volatile compounds in U. clathrata (83 compounds) was highest, followed by U. linza (75 compounds) and U. prolifera (57 compounds). Furthermore, a variety of polarity solvents were used to extract Ulva spp. in order to determine the influence of total phenolics (TPC), flavonoids (TFC) and carbohydrates content (TCC) on antioxidant activities. The results showed that ethyl acetate extracts of U. prolifera had the highest antioxidant activity (IC 50  = 143.18 μg/mL) in ABTS assay, and methanol extracts of U. clathrata revealed the highest scavenging activity (IC 50  = 95.32 μg/mL) against OH radicals as well as the aqueous extracts of U. prolifera exhibited the strongest radical scavenging capacity (IC 50  = 527.75 μg/mL) against DPPH radicals. Pearson correlation analysis revealed that the antioxidant capacity of Ulva spp. was significantly impacted by TFC and TPC. The findings presented in this thesis add to our understanding of volatilization characteristics and biological properties in edible marine green seaweeds., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Ocean acidification (OA) is a dramatic perturbation of seawater environments due to increasing anthropogenic emissions of CO 2 . Several studies indicated that OA frequently induces marine biota stress and a reduction of biodiversity. Here, we adopted the macroalga Ulva prolifera as a model and applied a complementary multi-omics approach to investigate the metabolic profiles under normal and acidified conditions. Our results show that U. prolifera grows at higher rates in acidified environments. Consistently, we observed lower sucrose and phosphocreatine concentrations in response to a higher demand of energy for growth and a higher availability of essential amino acids, likely related to increased protein biosynthesis. In addition, pathways leading to signaling and deterrent compounds appeared perturbed. Finally, a remarkable shift was observed here for the first time in the fatty acid composition of triglycerides, with a decrease in the relative abundance of PUFAs towards an appreciable increase of palmitic acid, thus suggesting a remodeling in lipid biosynthesis. Overall, our studies revealed modulation of several biosynthetic pathways under OA conditions in which, besides the possible effects on the marine ecosystem, the metabolic changes of the alga should be taken into account considering its potential nutraceutical applications., Competing Interests: The authors declare no conflict of interest.

The sessile green macroalgae in the Neopyropia aquaculture areas of the Subei Shoal are a confirmed source of green tides in the Southern Yellow Sea (SYS) of China. The green tides have a significant impact on local economic development (tourism, aquaculture, etc.) and ecological stability. In order to develop an effective method for eliminating the green macroalgae attached to Neopyropia aquaculture rafts, this study investigated the effects of mixed acid solutions (0.0475 % hydrochloric acid [HCl] and pH 2.0 citric acid) on cell damage, chlorophyll composition, phycobiliprotein content, and the photosynthetic fluorescence characteristics of Ulva prolifera and Neopyropia yezoensis. The chlorophyll contents of U. prolifera and N. yezoensis were significantly affected by the mixed acid solutions. After treatment with a mixed acid solution (4:3 volume ratio of HCl to citric acid) for 5 s, the photosynthetic pigment content of U. prolifera was significantly different from that of normal U. prolifera. However, photosynthetic pigment content in the treated N. yezoensis increased significantly. In addition, mixed acid solution treatment had a significant effect on the Fv/Fm of U. prolifera and N. yezoensis. After mixed acid treatment (4:3 HCl to citric acid), U. prolifera completely died, but the Fv/Fm of N. yezoensis was restored after 3 d. Therefore, a mixed acid solution comprising 0.0475 % HCl and pH 2.0 citric acid (at a volume ratio of 4:3) can be used as an agent for the removal of green macroalgae from Neopyropia aquaculture areas., Competing Interests: Declaration of competing interest We declare that there is no conflict of interest in this study., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

The global population is expected to reach 11 billion people by the year 2100 and will require sustainable sources of dietary protein. Most dietary protein originates from animal and terrestrial plant agriculture, which leads to deforestation, water pollution, and greenhouse gas emissions. Discovering alternative protein sources that are nutritionally adequate for the human diet without harmful environmental effects is imperative. Seaweeds are a promising option as they produce abundant protein with a low carbon footprint. Experimental evidence shows that seaweeds contain high concentrations of the essential amino acids (EAAs) necessary for human consumption, but seaweeds have yet to be evaluated with standardized metrics to compare their nutritional value to other protein sources. In this technical note, independent literature describing the EAA content and protein digestibility of 3 commonly consumed species of seaweeds was evaluated alongside traditional protein sources using a novel hybrid protein quality (HPQ) metric. HPQ is derived from the protein digestibility-corrected amino acid score and digestibility indispensable amino acid score but includes modifications to address the lack of in vivo digestibility data for seaweeds. Seaweed proteins are similar in quality to common plant protein sources such as peas, soy, and tree nuts. Furthermore, seaweed proteins from different species have complementary EAA profiles and can be mixed to form protein blends that are nutritionally on par with animal products such as milk and whey. Thus, seaweeds may be viable protein sources with a reduced footprint that provide beneficial ecosystem services., (© The Author(s) 2022. Published by Oxford University Press on behalf of the American Society for Nutrition.)

Around the world, green tides are happening with increasing frequency because of the dual effects of increasingly intense human activity and climate change; this leads to significant impacts on marine ecology and economies. In the last decade, the world's largest green tide, which is formed by Ulva/Enteromorpha porifera, has become a recurrent phenomenon every year in the southern Yellow Sea (China), and it has been getting worse. To alleviate the impacts of such green tide outbreaks, multiple measures need to be developed. Among these approaches, biotechnology plays important roles in revealing the outbreak mechanism (e.g., molecular identification technology for algal genotypes), controlling and preventing outbreaks at the origin sites (e.g., technology to inhibit propagation), and utilizing valuable algal biomass. This review focuses on the various previously used biotechnological approaches that may be applicable to worldwide seaweed blooms that result from global climate change and environmental degradation., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Copper ion (Cu 2+ ) and copper oxide (CuO) nanoparticle (NP) ecotoxicity are of increasing concern as they are considered to be a potential risk to marine systems. This study represents the first attempt to evaluate CuO NP impacts on the seaweeds and Cu 2+ on the chlorophyte Ulva rigida. Effects on oxidative stress, antioxidant defence markers, photosystem II function, thalli growth, and cell viability in U. rigida exposed for 4 up 72 h to1 and 5 mg L -1 Cu 2+ and CuO NPs were examined. Hydrogen peroxide (H 2 O 2 ) generation, superoxide dismutase (SOD) activity, malondialdehyde (MDA) content, and growth inhibition seemed to be reliable and early warning markers of toxicity. The most important variables of the principal component analysis (PCA): H 2 O 2 generation, antioxidant stress markers, and growth-based toxicity index, were higher at 1 mg L -1 CuO NPs compared to CuSO 4 and at 5 mg L -1 CuSO 4 compared to CuO NPs. Intracellular uptake kinetics fit well to the Michaelis-Menten equation. The higher toxicity at 5 mg L -1 CuSO 4 compared to 1 mg L -1 was due to the higher Cu uptake with increasing concentration, suggesting and higher accumulation ability. On the contrary, 1 mg L -1 CuO NPs induced more strongly toxicity effects than 5 mg L -1 . The relatively stronger effect of CuO NPs at 1 mg L -1 than the respective CuSO 4 concentration could be attributed to the higher rate of initial uptake (V c ) and the mean rate of Cu uptake [C max /(2 × K m )] at CuO NP treatment. The intracellular seaweed experimental threshold of Cu, which coincided with the onset of oxidative stress, was within the Cu concentration range recorded in Mediterranean Ulva spp., indicating that it may pose a substantial risk to marine environments., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Green tides dominated by Ulva prolifera have be present in the Southern Yellow Sea for 15 consecutive years. They not only damage the marine environment, but also cause economic losses to coastal cities. However, there is still no fully effective approach for preventing green tides. In this article, approaches for the prevention of U. prolifera taken over recent years are reviewed. They can be generally divided into physical, chemical, and biological approaches. Physical approaches have been used to control the overwhelming green macroalgae bloom and inhibit the germination of U. prolifera, including physical salvage approach, refrigeration net technology, improved farming methods and raft technology, and modified clay method. These approaches require significant labor and material resources. Many chemical reagents have been used to eliminate U. prolifera early germination and growth, such as oxidative algaecide, acid treatment, heavy metal compounds, antifouling coating, and alkaloids. Chemical approaches have high efficiency, high economic benefit, and simple operation. Presently, biological control approaches remain in the exploratory stage. The verification of pilot and large-scale experiment results in sea areas is lacking, including the application of large organisms and microorganisms to control U. prolifera, and some of the mechanisms have not been thoroughly studied. This article introduces the three types of approaches, and evaluates the advantages and disadvantages of different methods to facilitate the reduction of the green tide bloom scale in the Southern Yellow Sea., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Nowadays, algae are becoming more and more popular as a food group rich in nutrients, cosmetic raw materials full of antioxidants or valuable dietary supplements. They are of interest for the industry because they are found almost all over the world, in all climatic zones, both in fresh and salt waters. The aim of this study was to take a broad look at green algae (Chlorophyta) and to show how large the variability of the content of active compounds may depend on the species and the place and time of sample collection. Particular attention was paid to compounds with antioxidant activity, whose simplified profiles were created on the basis of complementary, semi-quantitative methods. Additionally, time-yield extraction optimizations were performed. Three different specimens of Ulva lactuca were compared: from the coastal zone of the Baltic Sea, from the open Baltic Sea area around Bornholm and Ulva spiralis ( Ulva lactuca polymorph) from the Atlantic Ocean. The studied algae of the Cladophora genera were three different species of freshwater algae from various habitats: a lake ( Cladophora glomerata ), a river ( Cladophora rivularis ) and aquarium farming ( Cladophora aegagropila, syn. Aegagropila linnaei ). The content of antioxidants and the extraction efficiency varied significantly depending on the species.

The impact of addition of an Ulva laetevirens (previously Ulva rigida) co-product treated with a broad-spectrum endo-protease when added to a standard corn-soy (S) based diet and a diet based on European protein sources (EU) on performance, in vivo digestibility and multiple gastrointestinal characteristics was investigated in broilers. In total, 624 Ross 308 one-day-old male broilers were fed one of 6 experimental diets (8 replicates) consisting of a basal diet (S or EU), or a basal diet including the U. laetevirens co-product (U) treated without (U-) or with (U+) a proteolytic enzyme. Starter diets contained 0 (wk 1) and 2.5 (wk 2), and the grower diets (wk 3 and 4) 5% seaweed co-product. In the last 2 wk, birds fed the S vs. EU grower diets showed a higher BW, BWG, and FI, as well as a lower FCR (-0.05 g/g) in wk 3 (P < 0.05). Heavier gizzards (+13%; P < 0.001) and heavier gizzard contents (+92%; P < 0.001) were observed in birds fed the EU vs. S diets, as well as longer villi (+8%; P = 0.010). U diets had a higher water holding capacity than the basal diets (+19%). In wk 4, U inclusion resulted in increased FCR (+0.06 g/g; P < 0.001), water intake (+7%; P < 0.001), and duodenal cross section (+5%; P = 0.033). Enzyme treatment did not affect digestibility of any nutrients, except for ash which was increased in birds fed U+ vs. U- diets (+60%; P < 0.001). U in S diets led to higher, and U in EU diets led to lower apparent pre-cecal digestibility of all nutrients (P < 0.001 for all nutrients). Although for both diet types performance was decreased, dietary U. laetevirens inclusion had different effects when added to a standard corn-soy diet and a diet based on European protein sources. No obvious health effects were observed, leading to the conclusion of the absence of performance of health promoting bioactive components in the U. laetevirens co-product, or of diminishing of these effects due to the proteolytic enzyme treatment., (Copyright © 2022 The Authors. Published by Elsevier Inc. All rights reserved.)

As global ecological disasters, green tide outbreaks have been observed in the Southern Yellow Sea (SYS) of China since 2007, resulting in considerable economic losses and environmental damage to the coastal cities of Jiangsu and Shandong Provinces. Therefore, prevention of green tides is crucial. Previous studies have revealed that a relatively small green tide outbreak scale in the SYS was observed in 2018 and 2020, with the green tides covering areas of 193 km 2 and 192 km 2 and durations of 91 days and 64 days, respectively. Killing green macroalgae attached to cultivation ropes in Neopyropia aquaculture areas, which has been considered a primary source of the blooms, early removal of Neopyropia aquaculture rafts, and green tide prevention in the SYS are the key reasons for the decrease in green tides in 2018 and 2020. Furthermore, to address the challenges associated with the current green tide source prevention measures, we proposed a comprehensive control method that combines ecological farming, early green tide prevention, and resource utilization. Potential secondary pollution caused by the chemicals used to control Ulva prolifera can be minimized. Conversely, Neopyropia yezoensis quality may be enhanced through continuous improvement of its culturing process, which in turn, could reduce the green tide blooming scale., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Green tide in the sea is an abnormal ecological phenomenon caused by the explosive proliferation or high aggregation of some green macroalgae under particular environmental conditions. Since 2007, green tides have occurred in the Yellow Sea for 15 consecutive years, resulting in extremely serious damage to coastal ecological environment, social development, and economic construction. Therefore, it is urgent to solve the green tide problem, which is driven by external and internal factors. Global researchers have scientifically recognized the external causes (environmental factors) of the green tide blooms, and have carried out a series of studies on the physiological response of green tide algae to some environmental factors and obtained some achievements. However, the internal causes of green tide blooms, which is the intense ecophysiological adaptability of green tide algae in response to drastic fluctuation of environmental factors, has not yet been sufficiently addressed. From the perspective of algae ecophysiology, we reviewed the response mechanisms of green tide forming species to the fluctuations of various ecological factors, including light intensity, carbon dioxide, temperature, salinity, desiccation, nutrient, heavy metals, and biotic factors. Moreover, we summarized the adaptive regulation mechanisms of green tide algae dealing with fluctuating environmental factors from the aspects of photosynthetic and growth physiology, nutritional physio-logy, and reproductive physiology, which help reveal the internal mechanisms of green tide blooming.

To explore the potential use of seaweed co-products for broiler diets, this study investigates whether an enzyme treatment of seaweed co-products improves performance, in vivo digestibility and health in broilers. In total, 360 Ross 308 male broilers were fed one of 5 experimental diets: a basal diet, or a basal diet including the U. laetevirens or S. chordalis co-product, with or without proteolytic enzyme treatment of the seaweed, using 6 replicate pens of 12 birds each. The starter (d 0-13) and grower (d 14-21) diet contained 5 and 10% (w/w) seaweed product, respectively. A general linear model with contrast statements was used after model assumptions and goodness of fit were evaluated through normal distribution of residuals. Inclusion of seaweed in the broiler diets increased body weight gain (+14%; P = 0.002), and feed intake (+12%; P = 0.001) in the third week of the experiment. Birds fed the U. laetevirens compared to the S. chordalis diets had a higher body weight gain (+11%; P = 0.007), and a lower feed conversion ratio (FCR; -7%; P < 0.001). Seaweed inclusion reduced apparent pre-cecal digestibility of all nutrients (P < 0.05). Birds fed U. laetevirens vs. S. chordalis diets had a 10% reduced villus length (P < 0.001). Enzymatic treatment reduced the digestibility of most nutrients, and increased crypt depth in birds fed the U. laetevirens diets, whereas the opposite was observed for the birds fed the S. chordalis diets (Seaweed × Enzyme P = 0.035). Untreated vs. treated seaweed in the diets led to lower (-60%) plasma Interleukin-13 levels (P = 0.035). In conclusion, the proteolytic enzyme treatment of the seaweed co-products did not improve performance nor health-related parameters, and reduced digestibility of the diets. Dietary inclusion of U. laetevirens co-products did improve performance based on growth and FCR, whereas inclusion of S. chordalis did not. Inclusion of U. laetevirens in broiler diets slightly reduced duodenal villus length and crypt depth. The inflammation response was strongly reduced, specifically in birds fed the untreated U. laetevirens diet, making the U. laetevirens co-product of interest for future research., (Copyright © 2022 The Authors. Published by Elsevier Inc. All rights reserved.)

Macroalgal blooms have become a serious threat to public health, fisheries, ecosystems, and global economies. Since 2007, in the Yellow Sea, China, Ulva green tides have occurred for 15 consecutive years. However, effective control methods are limited. Ulva prolifera attached to Neopyropia aquaculture rafts are believed to be the main source of blooms, therefore eliminating Ulva from rafts could effectively prevent and control blooms. We investigated this phenomenon and showed that macroalgae germination was significantly inhibited by dried Neopyropia yezoensis at concentrations of 1.2, 2.4, and 4.8 g DW -1 . Also, the inhibitory effects of dried N. yezoensis toward U. prolifera gametes at 2.4 and 4.8 g DW -1 were >90% at day 21. N. yezoensis culture filtrates and thalli were also used to determine dose-dependent inhibition effects on U. prolifera gamete germination. Both were potent and significantly inhibited germination at 1.75-7 g FW -1 ; the inhibitory effect 7 g FW -1 was >90% at day 21. As N. yezoensis thalli exhibited high inhibitory effects in laboratory experiments, we also performed field studies. N. yezoensis on ropes displayed high inhibitory effects on Ulva attachment and growth. Thus N. yezoensis powder, culture filtrates, and thalli displayed strong inhibitory effects on U. prolifera gametes, suggesting N. yezoensis attachment to ropes could be used to control green tides at the source., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

The high-valued utilization of Ulva (previously known as Enteromorpha ) bioresources has drawn increasing attention due to the periodic blooms of world-wide green tide. The polysaccharide is the main functional component of Ulva and exhibits various physiological activities. The Ulva oligosaccharide as the degradation product of polysaccharide not only possesses some obvious activities, but also possesses excellent solubility and bioavailability. Both Ulva polysaccharides and oligosaccharides hold promising potential in the food industry as new functional foods or food additives. Studies on Ulva polysaccharides and oligosaccharides are increasing and have been the focus of the marine bioresources field. However, the comprehensive review of this topic is still rare and do not cover the recent advances of the structure, isolation, preparation, activity and applications of Ulva polysaccharides and oligosaccharides. This review systematically summarizes and discusses the recent advances of chemical composition, extraction, purification, structure, and activity of Ulva polysaccharides as well as oligosaccharides. In addition, the potential applications as new functional food and food additives have also been considered, and these will definitely expand the applications of Ulva oligosaccharides in the food and medical fields.

Macroalgae and their rocky substrates both support diverse and abundant microbiota, performing essential ecological functions in marine ecosystem. However, the differences in the epiphytic microbial communities on macroalgae and rocky substrate are still poorly understood. In this study, the epiphytic microbial communities on four macroalgae (Corallina officinalis, Rhodomela confervoides, Sargassum thunbergii, and Ulva linza) and their rocky substrates from Weihai coast zone were characterized using high-throughput sequencing technology. The results showed that the alpha diversity indices were greater in rocky substrates than that in macroalgae. The microbial similarities among macroalgae and rocky substrate groups tended to decrease from the high taxonomic ranks to lower ranks, only 22.69% of the total amplicon sequence variants (ASVs) were shared between them. The functional analysis revealed that the microbiotas were mainly involved in metabolic activities. This study would provide the theoretical foundation for macroalgal cultivation and algal reef applications., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Biodegradation of polyvinyl chloride (PVC) by marine bacteria is a sustainable approach that leads to the production of different by-products but their toxicity needs to be evaluated. In the present study, polyvinyl chloride degradation products (PVCDP) produced by three marine bacterial isolates (T-1.3, BP-4.3 and S-237) in the culture supernatant were evaluated for toxicity on the germination of Vigna radiata and growth of Ulva lactuca. A total of 24 compounds comprising of benzene, fatty acid, ether, ester and plastic stabilizer (tris (2, 4-di-tert-butylphenyl) phosphate) were identified by GC-MS using diethyl ether solvent extraction from the supernatant. The per cent germination rate of the seed treated with PVCDP showed no significant effect but germination index and elongation inhibition rate were influenced significantly by PVCDP treatments. In seaweed (U. lactuca), PVCDP showed improvement in the daily growth rate. After ten days of treatment with PVCDP, pigment contents were improved in seaweed and PVCDP (2%) of isolate T-1.3 recorded the highest chlorophyll-a and chlorophyll-b., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Currently identification of alternate fuel is the key area of research under progress to overcome the depletion of fossil fuels, meet the domestic and industrial requirements. Generation of hydrogen, which is a clean fuel gas can solve various environmental related problems. Extensive research is being carried out to increase production of hydrogen through different substrates. This study aims to increase the production of hydrogen from Ulva reticulata (a macroalgal biomass). Initially, the biomass is pretreated mechanically with disperser and a biosurfactant, namely rhamnolipid in order to increase the solubilization of the biomass. The rate of COD liquefaction increased from 14% to 25% with the addition of biosurfactant to the macroalgal biomass, which is further treated mechanically using a disperser. The disperser rotor speed of 12,000 rpm and the specific energy input of 1175 kJ/kg TS (Total Solids) with the disintegration time of 30 min and biosurfactant dosage of 0.075 g/g TS were considered as the optimum parameters for the effective liquefaction of the macroalgal biomass. Approximately 3500 mg/L of the biopolymers were released after the combinative pretreatment (using disperser and biosurfactant). About 80 mL biohydrogen/g COD (Chemical Oxygen Demand) was generated when the biomass was pretreated with both the disperser and biosurfactant while the biomass pretreated with the disperser alone generated just 30 mL biohydrogen/g COD and the untreated biomass generated 5 mL biohydrogen/g COD. Thus, it can be concluded that Ulva reticulata can be utilized effectively to generate biohydrogen., (Copyright © 2021 Elsevier Ltd. All rights reserved.)