Evolutionary trends and analysis of the driving factors of Ulva prolifera green tides: A study based on the random forest algorithm and multisource remote sensing images.

Understanding the prolonged spatiotemporal evolution and identifying the underlying causes of Ulva prolifera green tides play pivotal roles in managing such occurrences, restoring water ecology, and fostering sustainable development in marine ecosystems. Satellite remote sensing represents the primary choice for monitoring Ulva prolifera green tides due to its capability for extensive, long-term ocean monitoring. Based on multi-source remote sensing images, ecological and environmental datasets, and machine learning algorithms, therefore, this study focused on "remote sensing modelling - evolution history - change trends - mechanism analysis" to elucidate both the remote sensing monitoring models and the underlying driving factors governing the spatiotemporal evolution of Ulva prolifera green tides in the highly impacted South Yellow Sea of China. With the use of GOCI Ⅰ/Ⅱ images, an hybrid remote sensing extraction model merging the robustness of the random forest (RF) model and the optical algae cloud index (ACI) was established to map Ulva prolifera distribution patterns. The ACI-RF method exhibited exceptional performance, with an F1 score surpassing 0.95, outperforming alternative methods such as the support vector machine (SVM) and K-nearest neighbour (KNN) methods. On the basis, we analysed the evolutionary trends and the driving factors determining these distribution patterns using meteorological data, runoff data, and data on various water quality parameters (SST, ocean current speed, wind speed, precipitation, DO, PAR, Si, NO3-, PO43-and N/P). Over the period from 2011 to 2022, excluding 2021, there was a notable decline in the area of Ulva prolifera green tides, varying between 397 and 2689.9 km2, with an average annual reduction rate of 3%. The maximum annual biomass varied between 0.12 and 15.9 kt. Notably, more than 75% of the area of Ulva prolifera green tides exhibited northward drift, which was significantly influenced by northern currents and wind fields. The analysis of driving factors indicates that factors such as average sea surface temperature, eastward wind speed, northward wind speed, precipitation, PO43- and N/P/Si significantly influence the biological growth rate of Ulva prolifera. Furthermore, coastal land use change and surface runoff, particularly surface runoff in June, significantly impacted the growth rate of Ulva prolifera, with Pearson correlation coefficients of 0.74 and 0.67, respectively. Against the background of global warming and severe deterioration in the marine environment, Ulva prolifera blooms persist. Consequently, two distinct management strategies were proposed based on the distribution patterns and cause analysis results for addressing Ulva prolifera green tides: establishing a continuous protection framework for rivers, lakes, and nearshore areas to mitigate pollutant inputs and implementing precise environmental monitoring measures in urban expansion areas and farmlands to combat overgrowth-induced green tides. This methodology could be applied in other regions affected by marine ecological disasters, and the criteria for selecting influencing factors offer a valuable reference for designing tailored and proactive measures aimed at controlling Ulva prolifera green tides.

Attached Ulva meridionalis on nearshore dikes may pose a new ecological risk in the Yellow Sea.

Green tides have been reported to occur in many sea areas worldwide. In China, most of them are caused by Ulva spp., such as Ulva prolifera and Ulva meridionalis. Green tide algae shed are frequently the initial biomass for the formation of green tide. Human activities and seawater eutrophication are the fundamental causes of the formation of the green tides in the Bohai Sea, Yellow Sea, and South China Sea, but other environmental factors may also have an impact on the shedding of green tide algae, such as typhoons and currents. Algae shedding is divided into artificial shedding and natural shedding. However, few studies have explored the relationship between algal natural shedding and environmental factors. pH, sea surface temperature, and salinity are critical environmental factors affecting the physiological state of algae. Therefore, based on field observations of the shedding of attached green macroalgae in Binhai Harbor, this study assessed the correlation between the shedding rate and environmental factors (pH, sea surface temperature, and salinity). The green algae that shed from Binhai Harbor in August 2022 were all identified as U. meridionalis. The shedding rate range was 0.88% ± 0.11% d-1 to 4.78% ± 1.76% d-1, and was not correlated with pH, sea surface temperature, or salinity; however, the environmental conditions were very suitable for the proliferation of U. meridionalis. This study provided a reference for the shedding mechanism of green tide algae and revealed that with the frequent human activities along the coast, U. meridionalis may pose a new ecological risk in the Yellow Sea.

Combined effects of ocean deoxygenation, acidification, and phosphorus limitation on green tide macroalga, Ulva prolifera.

Ocean deoxygenation, acidification, and decreased phosphorus availability are predicted to increase in coastal ecosystems under future climate change. However, little is known regarding the combined effects of such environmental variables on the green tide macroalga Ulva prolifera. Here, we provide quantitative and mechanistic understanding of the acclimation mechanisms of U. prolifera to ocean deoxygenation, acidification, and phosphorus limitation under both laboratory and semi-natural (mesocosms) conditions. We found that there were significant interactions between these global environmental conditions on algal physiological performance. Although algal growth rate and photosynthesis reduced when the nitrogen-to­phosphorus (N/P) ratio increased from 16:1 to 35:1 under ambient CO2 and O2 condition, they remained constant with further increasing N/P ratios of 105:1, 350:1, and 1050:1. However, the increasing alkaline phosphatase activities at high N/P ratios suggests that U. prolifera could use organic P to support its growth under phosphorus limitation. Deoxygenation had no effect on specific growth rate (SGR) but decreased photosynthesis under low N/P ratios of 16:1, 35:1, and 105:1, with reduced activities of several enzymes involved in N assimilation pathway being observed. Elevated CO2 promoted algal growth and alleviated the negative effect of deoxygenation on algal photosynthesis. The patterns of responses to high CO2 and low O2 treatments in in situ experiments were generally consistent with those observed in laboratory experiments. Our results generally found that the strong physiological acclimation capacity to elevated CO2, low O2, and high N/P could contribute to its large-scale blooming in coastal ecosystem.

Temperature and high nutrients enhance hypo-salinity tolerance of the bloom forming green alga, Ulva prolifera.

The response of seaweeds to environmental stressors can be population-specific, and be related to the regime of their habitats. To explore the growth and physiological responses of Ulva prolifera, two strains of this alga (Korean and Chinese strains) were studied under an interaction of temperature (20 and 25 °C), nutrients (low nutrients: 50 µM of nitrate and 5 µM of phosphate; high nutrients: 500 µM of nitrate and 50 µM of phosphate) and salinity (20, 30 and 40 psu). The lowest growth rates of both strains were observed at 40 psu of salinity, independent of temperature and nutrient levels. At 20 °C and low nutrients condition, the carbon: nitrogen (C: N) ratio and growth rate in the Chinese strain were increased by 31.1% and 21.1% at a salinity of 20 psu in comparison to the salinity of 30 psu, respectively. High nutrients decreased the ratio of C:N in both strains with increasing tissue N content. At the same time, high nutrients also increased soluble protein and pigments contents, as well as photosynthetic and growth rates in both strains at the same salinity levels at 20 °C. Under 20 °C and high nutrients conditions, the growth rates and C:N ratio of both strains were significantly decreased with increasing salinity. The pigment, soluble protein and tissue N showed an inverse trend with the growth rate at all conditions. Moreover, the higher temperature of 25 °C inhibited the growth in both strains regardless of nutrients levels. The temperature of 25 °C enhanced the contents of tissue N and pigments in the Chinese strain only at the low nutrients level. The interaction of high nutrients and 25 °C led to the accumulation of tissue N and pigment contents in both strains under all salinity conditions compared to the 20 °C and high nutrients level. The temperature of 25 °C and high nutrients decreased the growth rate in the Chinese strain at both salinities of 30 and 40 psu more than the 20 °C, and low nutrients level at the same salinity. These results suggest that the Ulva blooms caused by the Chinese strain were more impacted at hypo-salinity levels compared to the Korean strain. Eutrophic or high nutrients level enhanced the salinity tolerance in both strains of U. prolifera. There will be a decline of U. prolifera blooms of the Chinese strain at hyper-salinity levels.

Physiological acclimation of Ulva prolifera to seasonal environmental factors drives green tides in the Yellow Sea.

To understand how seasonal factors could drive the formation of green tide blooms and their flotation and decay, we cultured the green tide algal species Ulva prolifera at various temperatures (5, 10, 15, 20, 25, 30, and 35 °C) and light intensities (40, 80, 140, 240, and 400 µmol photons m-2 s-1). The results showed that the ratio of floating U. prolifera increased with increasing light and temperature, which was accompanied by morphological changes. The net photosynthetic rate and the proportion of floating U. prolifera thalli showed a nonlinear relationship. Furthermore, an enclosure experiment confirmed that the flotation of U. prolifera was influenced by light intensity via the regulation of photosynthesis of the thalli. These results suggest that seasonal factors control the photosynthesis of U. prolifera, which drives the flotation and decay of green tide algae in the Yellow Sea.

Green Tides in the Yellow Sea Promoted the Proliferation of Pelagophyte Aureococcus anophagefferens.

Harmful algal blooms formed by fast-growing, ephemeral macroalgae have expanded worldwide, yet there is limited knowledge of their potential ecological consequences. Here, we select intense green tides formed by Ulva prolifera in the Yellow Sea, China, to examine the ecological consequences of these blooms. Using 28-isofucosterol in the surface sediment as a biomarker of green algae, we identified the settlement region of massive floating green algae in the area southeast of the Shandong Peninsula in the southern Yellow Sea. The responses of the phytoplankton assemblage from the deep chlorophyll-a maximum layer were then resolved using high-throughput sequencing. We found striking changes in the phytoplankton community in the settlement region after an intensive green tide in 2016, characterized by a remarkable increase in the abundance of the pelagophyte Aureococcus anophagefferens, the causative species of ecosystem disruptive brown tides. Our study strongly suggests that the occurrence of massive macroalgal blooms may promote blooms of specific groups of microalgae through alteration of the marine environment.

Amphiphilic Nitroxide-Bearing Siloxane-Based Block Copolymer Coatings for Enhanced Marine Fouling Release.

The buildup of organic matter and organisms on surfaces exposed to marine environments, known as biofouling, is a disruptive and costly process affecting maritime operations. Previous research has identified some of the surface characteristics particularly suited to the creation of antifouling and fouling-release surfaces, but there remains room for improvement against both macrofouling and microfouling organisms. Characterization of their adhesives has shown that many rely on oxidative chemistries. In this work, we explore the incorporation of the stable radical 2,2,6,6-tetramethylpipiderin-1-oxyl (TEMPO) as a component in an amphiphilic block copolymer system to act as an inhibitor for marine cements, disrupting adhesion of macrofouling organisms. Using polystyrene-b-poly(dimethylsiloxane-r-vinylmethysiloxane) block copolymers, pendent vinyl groups were functionalized with TEMPO and poly(ethylene glycol) to construct an amphiphilic material with redox active character. The antifouling and fouling-release performance of these materials was investigated through settlement and removal assays of three model fouling organisms and correlated to surface structure and chemistry. Surfaces showed significant antifouling character and fouling-release performance was increased substantially toward barnacles by the incorporation of stable radicals, indicating their potential for marine antifouling applications.

How Ulva lactuca can influence the impacts induced by the rare earth element Gadolinium in Mytilus galloprovincialis? The role of macroalgae in water safety towards marine wildlife.

Rare earth elements (REEs) are gaining growing attention in environmental and ecotoxicological studies due to their economic relevance, wide range of applications and increasing environmental concentrations. Among REEs, special consideration should be given to Gadolinium (Gd), whose wide exploitation as a magnetic resonance imaging (MRI) contrast agent is enhancing the risk of its occurrence in aquatic environments and impacts on aquatic organisms. A promising approach for water decontamination from REEs is sorption, namely through the use of macroalgae and in particular Ulva lactuca that already proved to be an efficient biosorbent for several chemical elements. Therefore, the present study aimed to evaluate the toxicity of Gd, comparing the biochemical effects induced by this element in the presence or absence of algae. Using the bivalve species Mytilus galloprovincialis, Gd toxicity was evaluated by assessing changes on mussels' metabolic capacity and oxidative status. Results clearly showed the toxicity of Gd but further revealed the capacity of U. lactuca to prevent injuries to M. galloprovincialis, mainly reducing the levels of Gd in water and thus the bioaccumulation and toxicity of this element by the mussels. The results will advance the state of the art not only regarding the effects of REEs but also with regard to the role of algae in accumulation of metals and protection of aquatic organisms, generating new insights on water safety towards aquatic wildlife and highlighting the possibility for resources recovery.

Variation in microbial populations and antibiotic resistance genes in mariculture sediments in the present of the seaweed Ulva fasciata and under selective pressure of oxytetracycline.

The widely distributed seaweed Ulva fasciata has nutrient absorption abilities and can be used in the bioremediation of polluted maricultural environments. This study explored microbial community and antibiotic resistance gene (ARG) variation in mariculture sediments in response to different trace levels (10, 100, and 500 µg L-1) of oxytetracycline (OTC) and the presence of Ulva fasciata. The increase in OTC level promoted nutrient (NO3_-N and PO43--P) removal mainly due to Ulva fasciata adsorption. The abundances of the Euryarchaeota and Planctomycetes phyla in sediments were positively related to the increase in OTC stress, while a negative correlation occurred for the Proteobacteria phylum via metagenomic analysis. Compared with the control system, the increase rates of total ARGs were 3.90%, 7.36% and 13.42% at the OTC levels of 10, 100 and 500 µg L-1, respectively. OTC stress mainly favoured the collateral enrichment of non-corresponding polypeptide and MLS ARGs, mainly due to the enrichment of the phyla Planctomycetes and Euryarchaeota by the synergistic effect of OTC and nutrients. The results of quantitative PCR with tetracycline resistance genes (TRGs) (tetO, tetT, tetPB, tetW and otrA) and a horizontal transfer gene (intl1) demonstrated that all of genes had much higher gene numbers in sediments after 3 months of OTC stress than in those without OTC stress, which was strongly related to the variation in the phyla Bacteroidetes, Gemmatimonadetes and Acidobacteria. The significant correlation between intl1 and the target TRGs is indicative of the important role of the horizontal transfer of integron-resistant genes in the spread of TRGs.

Physiological and metabolic responses to hypersalinity reveal interpopulation tolerance in the green macroalga Ulva compressa with different pollution histories.

There is scarce investigation addressing interpopulation tolerance responses to address the influence of a history of chronic stress exposure, as that occurring in polluted environments, in photoautotrophs. We evaluated ecophysiological (photosynthetic activity) and metabolic (oxidative stress and damage) responses of two populations of green macroalga Ulva compressa from polluted (Ventanas) and non-polluted (Cachagua) localions of central Chile, and exposed to controlled hypersalinity conditions of 32 (control), 42, 62 and 82 psu (practical salinity units) for 6 h, 48 h and 6 d. Both primary production (ETRmax) and photosynthetic efficiency (αETR) were generally higher in the population from Cachagua compared to Ventanas at all times and salinities. Moreover, at most experimental times and salinities the population from Ventanas had greater levels of H2O2 and lipid peroxidation that individuals from Cachagua. Total ascorbate was higher in the population of Cachagua than Ventanas at 42 and 82 psu after 6 and 48 h, respectively, while at 6 d concentrations were similar between both populations at all salinities. Total glutathione was greater in both populations after 6 h at all salinities, but at 48 h its concentrations were higher only in the population from Cachagua, a trend that was maintained at 6 d under 82 psu only. Reduced and oxidized ascorbate (ASC and DHA, respectively) and glutathione (GSH and GSSG, respectively) demonstrated similar patterns between U. compressa populations, with an increase oxidation with greater salinities but efficient recycling to maintain sufficient batch of ASC and GSH. When assessing the expression of antioxidant enzymes catalase (CAT), superoxide dismutase (SOD) and dehydroascorbate reductase (DHAR), while the population of Ventanas displayed a general trend of upregulation with increasing salinities along the experiments, U. compressa from Cachagua revealed patterns of downregulation. Results demonstrated that although both populations were still viable after the applied hypersalinities during all experimental times, biological performance was usually more affected in the population from the Ventanas than Cachagua, likely due to a depressed baseline metabolism after a long history of exposition to environmental pollution.

Copper-induced concomitant increases in photosynthesis, respiration, and C, N and S assimilation revealed by transcriptomic analyses in Ulva compressa (Chlorophyta).

BACKGROUND: The marine alga Ulva compressa is the dominant species in copper-polluted coastal areas in northern Chile. It has been shown that the alga tolerates micromolar concentrations of copper and accumulates copper at the intracellular level. Transcriptomic analyses were performed using total RNA of the alga cultivated with 10 µ M copper for 0, 1, 3 and 5 days using RNA-seq in order to identify processes involved in copper tolerance. RESULTS: The levels of transcripts encoding proteins belonging to Light Harvesting Complex II (LHCII), photosystem II (PSII), cytochrome b6f, PSI, LHCI, ATP synthase and proteins involved in repair of PSII and protection of PSI were increased in the alga cultivated with copper. In addition, the level of transcripts encoding proteins of mitochondrial electron transport chain, ATP synthase, and enzymes involved in C, N and S assimilation were also enhanced. The higher percentages of increase in the level of transcripts were mainly observed at days 3 and 5. In contrast, transcripts involved protein synthesis and degradation, signal transduction, and replication and DNA repair, were decreased. In addition, net photosynthesis and respiration increased in the alga cultivated with copper, mainly at days 1 to 3. Furthermore, the activities of enzymes involved in C, N and S assimilation, rubisco, glutamine synthase and cysteine synthase, respectively, were also increased, mainly at days 1 and 3. CONCLUSIONS: The marine alga U. compressa tolerates copper excess through a concomitant increase in expression of proteins involved in photosynthesis, respiration, and C, N and S assimilation, which represents an exceptional mechanism of copper tolerance.

Hazardous effects of silver nanoparticles for primary producers in transitional water systems: The case of the seaweed Ulva rigida C. Agardh.

The acute toxicity of citrate capped silver nanoparticles (AgNP) and silver nitrate was evaluated on the marine macroalga Ulva rigida C. Agardh (1823). Silver bioaccumulation, ultrastructural chloroplast damages verified by TEM microscopy, inhibition of primary production, neutral lipid production and oxidative stress were observed after 24 h of exposure to AgNP. The toxic effects of silver nitrate in artificial seawater started from a concentration of 0.05 ppm and was more toxic than AgNP that produced effects from a concentration of 0.1 ppm. However only AgNP induced lipid peroxidation in U. rigida. The addition of natural organic and inorganic ligands, represented by transparent exopolymer particles (TEP) and clay, drastically reduced AgNP acute toxicity in a ratio AgNP:ligand of 1:100 and 1:200, respectively. The findings suggest a marked toxicity of Ag on marine macroalgae which however should be mitigated by the high natural ligand concentrations of the transitional environments.

Reproductive strategy of the floating alga Ulva prolifera in blooms in the Yellow Sea based on a combination of zoid and chromosome analysis.

Green algal blooms have occurred in the Yellow Sea for 13 consecutive years since 2007. However, little is known about the reproductive strategy of the dominant species Ulva prolifera in the field. In particular, it is not clear whether the floating Ulva species are sporophytes or gametophytes, and if their life history is sexual or asexual. In this study, the life history type was determined based on the size, phototactic response, and flagella number for the zoids in at least two successive generations. In addition, chromosome observations were conducted to distinguish the gametophytes and sporophytes in the floating Ulva species. The results showed that the floating Ulva species were all sporophytes with sexual reproductive patterns, thereby indicating that this Ulva species always maintains vegetative growth from April to June during the early stage of the blooms. In addition, we found that the chromosome numbers were 18 for the diploid sporophytes and nine for the haploid male and female gametophytes. These results provide useful information to help understand the explosive growth of these green algal blooms.

Darkness and low nighttime temperature modulate the growth and photosynthetic performance of Ulva prolifera under lower salinity.

In order to understand how darkness/irradiance and low nighttime temperature might alter physiology of Ulva prolifera under lower salinity conditions, we analyzed the growth rates, water content, superoxide dismutase (SOD) activity, total soluble proteins (SPs) and carbohydrates content at the end of dark and light period under three temperature levels (25-25 °C treatment: 25 °C for day and night; 15-15 °C treatment: 15 °C for day and night; 25-15 °C treatment: 25 °C for day with 15 °C for night) and two salinity conditions (15, 25), meanwhile, the pigment content (chlorophyll a and b), chlorophyll fluorescence and photosynthetic oxygen evolution also were determined during light phase. We found that the U. prolifera showed higher growth rate and SOD activity during dark phase at 25 °C, but this dark-induced increase could not be observed at 15 °C. The reasons for this increase varied, however, maybe not included water content and SPs for no significant difference in water content observed under all the treatments, as well as lower SPs content for dark period aside that at 15 °C and salinity 15. Compared to other two temperature treatments, the thalli grown at 25-15 °C showed higher growth rate and the photosynthetic oxygen evolution rate in light phase under salinity 15 conditions, although the maximum relative electron transport rate (rETRmax) showed higher value under 25 °C treatment. These results indicate that the darkness and the lower nighttime temperature maybe responsible reason for the rapid growth of these green tide algae.

Stress-localized durable anti-biofouling surfaces.

Growing demands for bio-friendly antifouling surfaces have stimulated the development of new and ever-improving material paradigms. Despite notable progress in bio-friendly coatings, the biofouling problem remains a critical challenge. In addition to biofouling characteristics, mechanically stressed surfaces such as ship hulls, piping systems, and heat exchangers require long-term durability in marine environments. Here, we introduce a new generation of anti-biofouling coatings with superior characteristics and high mechanical, chemical and environmental durability. In these surfaces, we have implemented the new physics of stress localization to minimize the adhesion of bio-species on the coatings. This polymeric material contains dispersed organogels in a high shear modulus matrix. Interfacial cavitation induced at the interface of bio-species and organogel particles leads to stress localization and detachment of bio-species from these surfaces with minimal shear stress. In a comprehensive study, the performance of these surfaces is assessed for both soft and hard biofouling including Ulva, bacteria, diatoms, barnacles and mussels, and is compared with that of state-of-the-art surfaces. These surfaces show Ulva accumulation of less than 1%, minimal bacterial biofilm growth, diatom attachment of 2%, barnacle adhesion of 0.02 MPa and mussel adhesion of 7.5 N. These surfaces promise a new physics-based route to address the biofouling problem and avoid adverse effects of biofouling on the environment and relevant technologies.

Salinity mediates the effects of nitrogen enrichment on the growth, photosynthesis, and biochemical composition of Ulva prolifera.

To study the combined effects of multiple nitrogen (N) sources and salinity on the growth and physiology on macroalgae, we cultured Ulva prolifera under three N levels (N0, 0.1235 mg L-1; N1, 0.6 mg L-1; and N2, 4.4 mg L-1; the ratios were 18:74:8 for NH4-N, NO3-N, and NO2-N, respectively) and three salinity conditions (15, 25, and 35). Then, the growth, pigment content, photosynthetic performance, superoxide dismutase (SOD) activity, and contents of soluble protein and carbohydrates were measured. The results showed the following: (1) Compared to that grown at salinity 25, the growth of U. prolifera decreased under salinity 35, especially under the N0 and N2 levels, but there were no significant effects of salinity 15 under any of the N levels. (2) There were no significant effects of salinity on the chlorophyll a (Chla) content, but compared to the content at salinity 25, the chlorophyll b (Chlb) content was enhanced by salinity 15 and 35; lower ratio values between Chla and carotenoids (Car) occurred under the salinity 25 treatment. Under each salinity condition, the pigments were enhanced by a high N level. (3) A relatively higher salinity level decreased the photosynthetic oxygen evolution rate, while a higher N level increased this value. Compared to the rate at salinity 25, the dark respiration rate (Rd) significantly increased at salinity 15 under the N0 condition. (4) SOD activity was enhanced by a high N level, but no significant effects of salinity were observed. (5) The carbohydrate content was enhanced at salinity 35 under the N0 and N1 levels, and under salinity 15, this value increased with increasing N levels. In conclusion, although the growth of U. prolifera decreased at high N levels under high salinity conditions, a high N level induced an increase in photosynthesis, while no significant decrease in growth occurred. These findings indicate that low salinity and high N levels may be nonnegligible reasons why this species thrives, and low salinity was the better choice when this species was used for wastewater treatment.

Dynamic metabolic profiles of the marine macroalga Ulva prolifera during fragmentation-induced proliferation.

Ulva prolifera, a type of marine macroalgae, is the causative species behind green tides mainly in the Yellow Sea and adjacent regions. Nevertheless, it can be used as food or animal feed in South China. The vegetative fragments of U. prolifera are an important seed source for successive green tide blooms. Fragmentation shortens the transition time from the vegetative state to the reproductive state. However, the translation of the algal metabolites during gametogenesis is far from well understood. In this study, the dynamic metabolic profiles of U. prolifera thallus during fragmentation-induced proliferation were investigated using non-targeted metabolomics approach via a series of time course experiments in June 2017. After a 30 min low temperature shock, fragmentation induced a reproductive response of 91.57% of U. prolifera in 48 h, whereas the value was only 21.43% in the control group. A total of 156 chromatographic peaks were detected, and 63 metabolites were significantly changed in U. prolifera during reproduction. Aanlysis of the kinetic metabolic pattern showed that the fragments not only induced the formation of sporangia, but also led to complex metabolite accumulation. During fragmentation-induced proliferation, U. prolifera consumed different sugars at different time points. γ-Aminobutyric acid (GABA), glutamic acid, gallic acid, and malic acid may play important roles in germ cell formation and in the release of U. prolifera, whereas n-hexanol, 2-methyl-3-phenylindole, and 3-indoleacetonitrile may be beneficial for biotic stress resistance. Compared with the control group, in the treatment group, metabolites such as alcohols and organic acids also showed significant difference with the photoperiod at the initial stage of proliferation (before 60 h). In conclusion, changes in the levels of metabolites, including sugars, organic acids, and alcohol with photoperiod may be the strategy adopted by U. prolifera to cope with fragmentation in nature.

Adsorptive removal of polycyclic aromatic hydrocarbons by detritus of green tide algae deposited in coastal sediment.

Rare information is available on the adsorptive removal of polycyclic aromatic hydrocarbons (PAHs) in the presence of algal detritus deposited in the coastal sediment during the outbreak of the green tide. The adsorptive removal of typical PAHs by Ulva prolifera (U. prolifera) detritus was firstly investigated since the algal detritus was of great importance for the biogeochemical cycle of coastal contaminants. The results showed that equilibrium adsorptive capacities of naphthalene, phenanthrene and benzo[a] pyrene on the U. prolifera detritus were 1.27, 1.97, and 2.49 mg kg-1, respectively, at the initial concentration of 10 µg L-1. The in situ monitoring using laser confocal scanning microscopy confirmed the adsorptive removal of PAHs by U. prolifera detritus. The adsorption of these PAHs was highly pH-dependent. The increase in salinity led to the increase in naphthalene removal rate, while the salinity showed scarce influence on the removal of phenanthrene and benzo[a] pyrene. There was a good linear relationship (R2 ≥ 0.9892) between the removal efficiency of PAHs and the initial concentration of PAHs. Slow desorption kinetics and low desorption rate (<16%) indicated that the adsorptive removal of PAHs could be benign to the environment. These findings demonstrated that the occurrence of green tide could provide a new natural remediation approach for contamination of PAHs through the adsorptive removal by the detritus of green tidal algae deposited in the coastal sediment.

Buoyancy potential of dominant green macroalgal species in the Yellow Sea's green tides, China.

Large-scale green tides caused by Ulva prolifera, occurred for 12 consecutive years in the Yellow Sea of China. To resolve the abrupt shift in species composition between attached and floating macroalgal assemblages, field experiments were conducted from May to July 2017 to quantify the net buoyancy force and compare the floating potential of the common green macroalgae from the red algal seaweed Pyropia yezoensis rafts. At the same time, U. prolifera from different sampling locations were tested to study variable buoyancy of this species and the associated influencing factors. Our results illustrated a stronger positive buoyant force and a proportionally greater buoyancy capacity of U. prolifera, compared to the other co-occurring species. Buoyancy is a dynamic trait and is closely correlated with light intensity, morphology and physiological status. The positive buoyancy of U. prolifera is an important factor that helps explain its predominance in the Yellow Sea's large-scale green tides.

Differential Photosynthetic Response of a Green Tide Alga Ulva linza to Ultraviolet Radiation, Under Short- and Long-term Ocean Acidification Regimes.

Both ocean acidification (OA) and solar ultraviolet (UV) radiation can bring about changes in macroalgal physiological performance. However, macroalgal responses to UV radiation when acclimatized to OA under different time scales are rare. Here, we investigate the response of Ulva linza, a green tide alga, to UV radiation in the form of photosynthetically active radiation (PAR) or PAB (PAR+UVA+UVB) radiation. Radiation exposures were assessed following long-term (from spore to adult stage, 1 month) and short-term (adult stage, 1 week) OA treatments. Results showed that increased CO2 decreased the damage rate (k) and repair rate (r) of thalli grown under short-term OA conditions with PAB treatment, the ratio of r:k was not altered. Following long-term OA conditions, r was not affected, although k was increased in thalli following PAB treatment, resulting in a reduced ratio of r:k. The relative level of UV inhibition increased and UV-absorbing compounds decreased when algae were cultured under long-term OA conditions. The recovery rate of thalli was enhanced when grown under long-term OA after UV radiation treatment. These results show that blooming algae may be more sensitive to UV radiation in marine environments, but it can develop effective mechanisms to offset the negative effects, reflecting acclimation to long-term OA conditions.