The Ulva prolifera bloom is considered one of the most serious ecological disasters in the Yellow Sea in the past decade, forming a carbon sink in its source area within a short period but becoming a carbon source at its destination. To explore the effects of different environmental changes on seawater dissolved carbon pools faced by living U. prolifera in its originating area, U. prolifera were cultured in three sets with different light intensity (54, 108, and 162 μmol m-2 s-1), temperature (12, 20, and 28 °C) and nitrate concentration gradients (25, 50, and 100 μmol L-1). The results showed that moderate light (108 μmol m-2 s-1), temperature (20 °C), and continuous addition of exogenous nitrate significantly enhanced the absorption of dissolved inorganic carbon (DIC) in seawater by U. prolifera and most promoted its growth. Under the most suitable environment, the changes in the seawater carbonate system were mainly dominated by biological production and denitrification, with less influence from aerobic respiration. Facing different environmental changes, U. prolifera continuously changed its carbon fixation mode according to tissue δ13C results, with the changes in the concentrations of various components of DIC in seawater, especially the fluctuation of HCO3- and CO2 concentrations. Enhanced light intensity of 108 μmol m-2 s-1 could shift the carbon fixation pathway of U. prolifera towards the C4 pathway compared to temperature and nitrate stimulation. Environmental conditions at the origin determined the amount of dissolved carbon fixed by U. prolifera. Therefore, more attention should be paid to the changes in marine environmental conditions at the origin of U. prolifera, providing a basis for scientific management of U. prolifera.

The world\'s largest green tide, caused by the nuisance green algae Ulva prolifera, has occurred in the southern Yellow Sea for 16 consecutive years. It is puzzling why the extensive floating green tide occurs exclusively in the southern Yellow Sea, rather than other waters. We speculate that the transition of U. prolifera from a sessile state to a surface-floating one is the underlying cause of the floating green tide. Here we founded that the floating of U. prolifera was attributed to detachment from substrata and appropriate desiccation. The convergence of unreasonable green algae disposal, geographical features and farming patterns of Porphyra (economic red algae) in Subei Shoal contributed to mass production of floating U. prolifera, resulting in the exclusive occurrence of the floating green tides. Inducing the natural inactivation of green algae to prevent the floating of U. prolifera may effectively mitigate the extensive Ulva bloom at zero cost.

Flavobacteriia are the dominant and active bacteria during algal blooms and play an important role in polysaccharide degradation. However, little is known about phages infecting Flavobacteriia, especially during green tide. In this study, a novel virus, vB_TgeS_JQ, infecting Flavobacteriia was isolated from the surface water of the Golden Beach of Qingdao, China. Transmission electron microscopy demonstrated that vB_TgeS_JQ had the morphology of siphovirus. The experiments showed that it was stable from -20°C to 45°C and pH 5 to pH 8, with latent and burst periods both lasting for 20 min. Genomic analysis showed that the phage vB_TgeS_JQ contained a 40,712-bp dsDNA genome with a GC content of 30.70%, encoding 74 open-reading frames. Four putative auxiliary metabolic genes were identified, encoding electron transfer-flavoprotein dehydrogenase, calcineurin-like phosphoesterase, phosphoribosyl-ATP pyrophosphohydrolase, and TOPRIM nucleotidyl hydrolase. The abundance of phage vB_TgeS_JQ was higher during Ulva prolifera (U. prolifera) blooms compared with other marine environments. The phylogenetic and comparative genomic analyses revealed that vB_TgeS_JQ exhibited significant differences from all other phage isolates in the databases and therefore was classified as an undiscovered viral family, named Zblingviridae. In summary, this study expands the knowledge about the genomic, phylogenetic diversity and distribution of flavobacterial phages (flavophages), especially their roles during U. prolifera blooms.
OBJECTIVE: The phage vB_TgeS_JQ was the first flavobacterial phage isolated during green tide, representing a new family in Caudoviricetes and named Zblingviridae. The abundance of phage vB_TgeS_JQ was higher during the Ulva prolifera blooms. This study provides insights into the genomic, phylogenetic diversity, and distribution of flavophages, especially their roles during U. prolifera blooms.

The bacterial communities related to seaweed can vary considerably across different locations, and these variations influence the seaweed\'s nutrition, growth, and development. To study this further, we evaluated the bacteria found on the green marine seaweed Ulva prolifera from Garorim Bay and Muan Bay, two key locations on Republic of Korea\'s west coast. Our analysis found notable differences in the bacterial communities between the two locations. Garorim Bay hosted a more diverse bacterial population, with the highest number of ASVs (871) compared to Muan Bay\'s 156 ASVs. In Muan Bay, more than 50% of the bacterial community was dominated by Pseudomonadota. On the other hand, Garorim Bay had a more balanced distribution between Bacteroidota and Pseudomonadota (37% and 35.5%, respectively). Additionally, Cyanobacteria, particularly Cyanothece aeruginosa, were found in significant numbers in Garorim Bay, making up 8% of the community. Mineral analysis indicated that Garorim Bay had higher levels of S, Na, Mg, Ca, and Fe. Function-wise, both locations exhibited bacterial enrichment in amino acid production, nucleosides, and nucleotide pathways. In conclusion, this study broadens our understanding of the bacterial communities associated with Ulva prolifera in Korean waters and provides a foundation for future research on the relationships between U. prolifera and its bacteria.

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.

Photoperiod and temperature are two main factors in the growth of macroalgae, and changes in photoperiod and diurnal temperature difference exist in natural condition. In order to study the effects of photoperiod and diurnal temperature difference on the growth of green algae Ulva prolifera, we cultured this species under three light/dark cycles (light: dark = 10:14, 12:12 and 16:08) with constant (22 °C for light and dark period, noted as 22-22 °C) and diurnal temperature difference (22 °C and 16 °C for light and dark period, respectively, noted as 22-16 °C) conditions. The results showed that: 1) Compared with 10:14 light/dark cycle, the growth of U. prolifera under 12:12 light/dark cycle was significantly enhanced by 39% and 16% for 22-22 °C and 22-16 °C treatments, respectively, while the increase proportion decreased when the daylength increase from 12 h to 16 h. 2) The enhancement in growth induced by diurnal temperature difference was observed under 10:14 light/dark cycle, but not for 12:12 and 16:08 light/dark cycle treatments. 3) The Chl a content and photosynthetic rate increased under short light period and 22-22 °C conditions, while under 22-16 °C conditions, higher photosynthetic rate was observed under 12:12 light/dark cycle and no significant difference in Chl a content was observed. 4) Under 22-22 °C conditions, compared with 10:14 (L:D) treatment, the expression levels of proteins in light-harvesting complexes, PSII and carbon fixation were down regulated, while the photorespiration and pentose phosphate pathway (PPP) were up regulated by 16:08 light dark cycle. Then we speculate that the higher photosynthetic rate may be one compensation mechanism in short photoperiod, and under long light period condition the up regulations of photorespiration and PPP can be in charge of the decrease in enhancement growth induced by longer daylength.

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.