The world\'s largest \"green tide\" (Ulva prolifera) has occurred every year since 2007 in the Yellow Sea. The Subei Shoal area is thought to be the origin of the green tide. Based on field data from 2016 to 2023, seasonal and interannual variations of dissolved nutrients and their ecological effects in the Subei Shoal were analyzed. Spatial distribution of dissolved inorganic nitrogen (DIN), dissolved inorganic phosphorus (DIP) and dissolved silicate (DSi) showed clear terrestrial sources, while ammonia (NH4-N) and dissolved organic nitrogen (DON) were not solely controlled by terrestrial sources. The seasonal variations of NH4-N, DIN, DON, DIP and DSi concentrations were significant, and the interannual variations of DIN, DON, DIP and DSi concentrations showed general decreasing trends from 2016 to 2023. The key factors affecting the seasonal and interannual variations of DIN and DIP concentrations were terrestrial input, aquaculture wastewater discharge, atmospheric deposition, submarine groundwater discharge and macroalgae absorption, while the dominant factor determining the variations of DSi concentrations was terrestrial input. NH4-N and DON concentrations were mainly influenced by aquaculture wastewater discharge and the absorption and release of macroalgae. The high nutrient concentrations in the Subei Shoal throughout the year provided sufficient material basis for the growth of Ulva prolifera in the source area of green tide outbreak.

Since 2007, persistent green tides in the Yellow Sea of China (YSC) have inflicted substantial economic and social losses. In response, the Chinese government has enacted various policies to mitigate these impacts. This study introduced an evolutionary-multiple streams framework and employed Social Network Analysis (SNA) and Text Analysis from 2007 to 2023, tracing three phases of policy development: growth (2007-2012), stability (2013-2017), and explosion (2018-2023). Findings indicated that during the growth phase, the most of the policy themes were \"monitoring\" and \"emergency\"; in the stability phase, policy issuance by provincial and municipal agencies began to increase; in the explosion phase, a basic consensus was reached on source control and intergovernmental cooperation in the management of green tides. Themes such as \"ecology\", \"extreme weather\", and \"green tide exploitation\" have been emphasized. This analysis provides insights for future policy formulation in green tide control and broader marine environmental governance.

The proliferation of large green macroalgae in marine environments has led to the occurrence of green tides, particularly in the South Yellow Sea region of China, where Ulva prolifera has been identified as the primary species responsible for the world\'s largest green tide events. Allelopathy among plants is a critical factor influencing the dynamics of green tides. This review synthesizes previous research on allelopathic interactions within green tides, categorizing four extensively studied allelochemicals: fatty acids, aldehydes, phenols, and terpenes. The mechanisms by which these compounds regulate the physiological processes of green tide algae are examined in depth. Additionally, recent advancements in the rapid detection of allelochemicals are summarized, and their potential applications in monitoring green tide events are discussed. The integration of advanced monitoring technologies, such as satellite observation and environmental DNA (eDNA) analysis, with allelopathic substance detection is also explored. This combined approach addresses gaps in understanding the dynamic processes of green tide formation and provides a more comprehensive insight into the mechanisms driving these phenomena. The findings and new perspectives presented in this review aim to offer valuable insights and inspiration for researchers and policymakers.

Micro-propagules (banks of microscopic forms) play important roles in the expansion of green tides, which are spreading on eutrophic coasts worldwide. In particular, large-scale green tides (Yellow Sea Green Tide, YSGTs) have persisted in the Yellow Sea for over 15 years, but the dynamics and functions of micro-propagules in their development remain unclear. In the present study, year-round field surveys were conducted to identify the reservoirs and investigate the persistence mechanisms and associated biotic and abiotic factors driving the temporal and spatial variations of micro-propagules. Micro-propagules in the southern Yellow Sea (SYS) showed evident spatial heterogeneity in terms of seasonal patterns and major influencing factors. Offshore of the SYS, the micro-propagule population underwent ephemeral expansion along with a large-scale bloom of floating Ulva algae in late spring and early summer. The Subei Shoal, particularly the sediments in the central raft region, had the highest micro-propagule abundance (MA) and was a major reservoir. The pronounced seasonal variation of MA in the Subei Shoal was primarily associated with the attached Ulva algae on Neopyropia aquaculture rafts. Vast aquaculture rafts provided essential substrates for micro-propagules to complete their life cycle and replenish the seed bank, thereby sustaining persistent YSGTs. It implied that habitat modification has pronounced ecological impacts on this intertidal muddy flat. The unique environmental conditions (enriched nutrients, esp. nitrate, favourable seawater temperatures in spring, and strong tidal mixing) facilitated the abundance, seasonal variation and recruitment of micro-propagules in the Subei Shoal. Given the current mitigation measures implemented in the raft region, further research is required to monitor and investigate the physiological and ecological responses of micro-propagule populations to the complex hydrobiological, geochemical, and physical matrices.

The recurring appearance of Ulva prolifera green tides has become a pressing environmental issue, especially for marine transportation, tourism, and aquaculture in the stage of decomposition. An abundance of decaying U. prolifera leads to water acidification, hypoxia and pathogenic microorganism proliferation, threatening marine germplasm resources, particularly benthic organisms with weak escape ability. Epigenetic modification is considered to be one of the molecular mechanisms involved in the plastic adaptive response to environmental changes. However, few studies concerning the specific impact of decaying green tide on benthic animals at the epigenetic level. In this study, decomposing algal effluents of U. prolifera, sediments containing uncorrupted U. prolifera, pathogenic microorganism were considered as impact factors, to reveal the effect of decaying U. prolifera on marine economic benthic species, Paralichthys olivaceus, using both field and laboratory simulation experiments. Field simulation experiment showed higher mortality rates and serious histopathological damage than the laboratory simulation experiment. And both the decaying U. prolifera and the sediment containing U. prolifera were harmful to P. olivaceus. Genome-wide DNA methylation and transcription correlation analyses showed that the response of P. olivaceus to green tide stress and bacterial infection was mainly mediated by immune signaling pathways such as PI3K-Akt signaling pathway. DNA methylation regulates the expression of immune-related genes involved in the PI3K-Akt signaling pathway, which enables P. olivaceus to adapt to the adverse environmental stresses by resisting apoptosis. In summary, this research analyzed the potential role of P. olivaceus in decaying U. prolifera, which is of great significance for understanding the impact of decaying green tide on marine commercial fish and also provides some theoretical guidance for the proliferation and release of fish seedlings.

We conducted continuous monitoring at 13 stations along the Jiangsu coast to study the spatiotemporal distribution, population succession of micropropagules of green algae, and their impact on the outbreak of Southern Yellow Sea green tide. The study discovered that: 1) Green algae micropropagules had obvious temporal and spatial distribution and population changes along the Jiangsu coast. The monthly average abundance of micropropagules of green algae at station BH1, which was the high-value area, was 1230 inds/L. Station XS2 had the second-highest value area. Green algae micropropagules had an average monthly abundance of 836 inds/L. Between stations XS2 and BH1, the amount of green algae micropropagules steadily declined in comparison to other stations. The abundance was greatest from spring to early summer, and Ulva prolifera micropropagules predominated; 2) Compared with salinity, temperature had a more obvious effect on the micropropagules of green algae along the Jiangsu coast; 3) Green algae micropropagules on the Jiangsu coast could be a potential additional source on the outbreak of Southern Yellow Sea green tide. More data are needed to corroborate this conclusion. For the purpose of preventing and managing green tide, it is crucial to investigate the Southern Yellow Sea\'s potential supplementary source. This study analyzes the spatiotemporal distribution and population changes of green algae micropropagules along the Jiangsu coast, as well as their impact on green tide outbreaks, providing scientific data support for the prevention and control of green tides in the Southern Yellow Sea.

In order to study the role of oil spills in the occurrence of green tide in the Yellow Sea, the physiological characteristics and photosynthetic activities of green tide causative-species Ulva prolifera was monitored under different conditions including two oil water-accommodated fractions (WAFs) of diesel oil and crude oil, dispersed water-accommodated fractions (DWAFs) and dispersant GM-2. The results showed that, the physiological parameters of U. prolifera including the growth, pigment, carbohydrate and protein contents decreased with the increased diesel oil WAF (WAFDO) concentration, while crude oil WAF (WAFCO) showed low concentration induction and high concentration inhibition effect. In addition, with the increase of WAFs concentration, two antioxidant activities were activated. However, compared with WAFDO alone and WAFCO alone, the mixture of oil and dispersant enhanced the toxicity on the above physiological characteristics of U. prolifera. On the other hand, the photosynthetic efficiency of U. prolifera showed a similar trend. Two WAFs showed significant concentration effects on the chlorophyll-a fluorescence transients and JIP-test. The addition of dispersant further blocked the electron flow beyond QA and from plastoquinone (PQ) to PSI acceptor side, damaged the active OEC centers at the PSII donor side, suppressed the pool size and the reduction rate of PSI acceptor side, and reduced the energy transfer efficiency between PSII functional units. These results implied that the crude oil spills may induce the formation of U. prolifera green tide, and the oil dispersant GM-2 used after the oil spills is unlikely to further stimulate the scale of bloom, while the diesel oil spills is always not conducive to the outbreak of green tide of U. prolifera.

We investigated spatial heterogeneity and diel variations in bacterioplankton and pico-nanoeukaryote communities, and potential biotic interactions at the extinction stage of the Ulva prolifera bloom in the Jiaozhou Bay, Yellow Sea. It was found that the presence of Ulva canopies significantly promoted the cell abundance of heterotrophic bacteria, raised evenness, and altered the community structure of bacterioplankton. A diel pattern was solely significant for pico-nanoeukaryote community structure. >50 % of variation in the heterotrophic bacterial abundance was accounted for by the ratio of Bacteroidota to Firmicutes, and dissolved organic nitrogen effectively explained the variations in cell abundances of phytoplankton populations. The factors representing biotic interactions frequently contributed substantially more than environmental factors in explaining the variations in diversity and community structure of both bacterioplankton and pico-nanoeukaryotes. There were higher proportions of eukaryotic pathogens compared to other marine systems, suggesting a higher ecological risk associated with the Ulva blooms.

Green tides, a globally prevalent marine ecological anomaly observed in coastal regions, have received substantial attention. However, there is limited research on the burial of Ulva prolifera in sediments during the late stages of green tide outbreaks. This study investigates the effect of temperature on U. prolifera buried in sediment over 30 days. The measurements included the length, biomass, relative growth rate, chlorophyll composition and maximum quantum yield (Fv/Fm) of PS II at different stages. The results indicate that at -20 °C, numerous seedlings emerged after 14 days of recovery culture, suggesting the release of spores or gametes; survival was possible from -2 °C to 15 °C; but at 20 °C and 30 °C, all U. prolifera died. The U. prolifera buried in sediment during the late stage of green tide outbreaks may serve as one of the sources for the subsequent year\'s green tide eruption. This research provides insights into the origins of green tide outbreaks in the southern Yellow Sea.

Environmental pollution is considered to lead to Ulva sp. green tides. Nevertheless, nutrients with high concentrations inhibit algae which may be damaged by antibiotics, such as OTC (oxytetracycline). Thus, Ulva sp. algae might pay a physiological cost under nutrient-OTC combined pressures. If this hypothesis is confirmed, Ulva sp. algae cannot easily form green tides, or green tides are difficult to maintain. To test this hypothesis, an uniform design experiment during which OTC, ammonia (NH4-N) and phosphate (PO4-P) were factors was set to simulate nutrient-OTC combined pressures, and Ulva lactuca was exposed to the pressures for 96 h. The TN (total nitrogen, CTN) or TP (total phosphorus, CTP) content in U. lactuca increased with increasing nutrient concentrations, as CTN = 21.206±1.000+ 1.227±0.418NH4-N × PO4-P (R2 = 0.282, p < 0.05) and CTP = 1.886±0.266+ 0.877±0.126PO4-P (R2 = 0.689, p < 0.05), respectively. The increase in dry weight of U. lactuca (Wdry) had a relationship with combined pressures, Wdry = 0.011±0.029 - 0.036±0.014PO4-P (R2 = 0.243, p < 0.05), i.e., the algal growth was inhibited by increasing PO4-P concentration. The SOD (Superoxide dismutase) activity (ASOD) was stimulated by OTC, as ASOD = 127.868±8.741+9.587±4.179 OTC (R2 = 0.193, p < 0.05). The contents of Chl a and b (Ca and Cb) were negatively affected by OTC or PO4-P with high concentration, as Ca = 0.566±0.042 - 0.024±0.022 OTC × PO4-P (R2 = 0.179, p < 0.05) and Cb = 0.512±0.043-0.044±0.020PO4-P (R2 = 0.180, p < 0.05). Thus, too high concentrations of PO4-P or OTC may hinder the formation and maintenance of Ulva sp. green tides.