{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T15:23:54Z","timestamp":1785425034517,"version":"3.56.0"},"reference-count":116,"publisher":"MDPI AG","issue":"15","license":[{"start":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T00:00:00Z","timestamp":1784764800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100002855","name":"National Key Research & Development Program of China","doi-asserted-by":"publisher","award":["2024YFC3109005"],"award-info":[{"award-number":["2024YFC3109005"]}],"id":[{"id":"10.13039\/501100002855","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100011427","name":"State Key Laboratory of Marine Geology","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100011427","id-type":"DOI","asserted-by":"crossref"}]},{"id":[{"id":"https:\/\/ror.org\/02bxj9z30","id-type":"ROR","asserted-by":"publisher"}]},{"name":"Young S&T Talents Cultivation Project, CAST"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Biology"],"abstract":"<jats:p>Green tide disasters have erupted consecutively in the Southern Yellow Sea, yet their driving mechanisms remain debated. The \u201craft-origin-hypothesis\u201d identifies the source as epiphytic Ulva macroalgae attached to laver aquaculture rafts in the Subei Shoal, whereas the \u201cmicropropagule-hypothesis\u201d speculates that Ulva micropropagules settling into open sea sediments constitute a cross-year seed reserve. The latter has hitherto relied primarily on laboratory culture simulations, with field-scale assessment lacking. This study established 46 surface sediment survey stations from 2020 to 2022, covering periods before, during, and after green tide outbreaks. Surface sediments were obtained using a box corer, and Ulva micropropagules were quantitatively analyzed through laboratory cultivation and morphological identification. Results showed that micropropagule abundance was generally low and exhibited spatial heterogeneity; high-value areas persistently existed in the Subei Shoal aquaculture zone, extended into the open sea during outbreaks, and generally disappeared in the open sea during and after the decline period. Monthly variations indicated that micropropagules released and settling into sediments (e.g., in the open sea) during large-scale outbreaks could not survive until the following year\u2019s floating period; the initial biomass of subsequent eruptions did not originate from the revival of the seafloor \u201cseed source\u201d in the open sea, which remains speculative.<\/jats:p>","DOI":"10.3390\/biology15151220","type":"journal-article","created":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T09:11:26Z","timestamp":1784797886000},"page":"1220","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Spatiotemporal Distribution of Ulva Micropropagules in Surface Sediments: Insights from Field Sampling Data in the Southern Yellow Sea (2020\u20132022)"],"prefix":"10.3390","volume":"15","author":[{"given":"Yutao","family":"Qin","sequence":"first","affiliation":[{"name":"Key Laboratory of Marine Ecological Monitoring and Restoration Technologies, East China Sea Ecological Center, Ministry of Natural Resources, Shanghai 201206, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lihua","family":"Xia","sequence":"additional","affiliation":[{"name":"Key Laboratory of Marine Ecological Monitoring and Restoration Technologies, East China Sea Ecological Center, Ministry of Natural Resources, Shanghai 201206, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Huanhong","family":"Ji","sequence":"additional","affiliation":[{"name":"Key Laboratory of Marine Ecological Monitoring and Restoration Technologies, East China Sea Ecological Center, Ministry of Natural Resources, Shanghai 201206, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaobo","family":"Wang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Marine Ecological Monitoring and Restoration Technologies, East China Sea Ecological Center, Ministry of Natural Resources, Shanghai 201206, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiaxing","family":"Cao","sequence":"additional","affiliation":[{"name":"Key Laboratory of Marine Ecological Monitoring and Restoration Technologies, East China Sea Ecological Center, Ministry of Natural Resources, Shanghai 201206, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuhan","family":"Zhang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Marine Ecological Monitoring and Restoration Technologies, East China Sea Ecological Center, Ministry of Natural Resources, Shanghai 201206, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jing","family":"Xia","sequence":"additional","affiliation":[{"name":"School of Oceanography, Shanghai Jiao Tong University, Shanghai 200030, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9916-6152","authenticated-orcid":false,"given":"Jinlin","family":"Liu","sequence":"additional","affiliation":[{"name":"Key Laboratory of Marine Ecological Monitoring and Restoration Technologies, East China Sea Ecological Center, Ministry of Natural Resources, Shanghai 201206, China"},{"name":"Project Management Office of China National Scientific Seafloor Observatory, Tongji University, Shanghai 200092, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,7,23]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"e43","DOI":"10.1017\/S0025315423000310","article-title":"Competition relations between selected microalgae and bloom-forming Ulva prolifera","volume":"103","author":"Liu","year":"2023","journal-title":"J. Mar. Biol. Assoc. UK"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"109625","DOI":"10.1016\/j.biombioe.2026.109625","article-title":"Identification of PHA synthesis potential and fermentation condition optimization of a Bacillus strain isolated from Ulva prolifera","volume":"215","author":"Zha","year":"2026","journal-title":"Biomass Bioenergy"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"115639","DOI":"10.1016\/j.jenvman.2022.115639","article-title":"The inhibitory effects of Ulva prolifera extracts on early growth of Spartina alterniflora and the underlying mechanisms","volume":"319","author":"Zhao","year":"2022","journal-title":"J. Environ. Manag."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1705","DOI":"10.1007\/s11802-025-6162-y","article-title":"Succession of Planktonic Crustaceans Responding to Ulva Green Tide in the Subei Shoal, Southwestern Yellow Sea","volume":"24","author":"Li","year":"2025","journal-title":"J. Ocean Univ. China"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"187","DOI":"10.1007\/s10201-020-00644-w","article-title":"Influences of green macroalgae blooms on nutrients and sulfide dynamics in hypereutrophic intertidal ecosystems","volume":"22","author":"Senga","year":"2021","journal-title":"Limnology"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1007\/s00343-011-0126-3","article-title":"Acute toxicity of live and decomposing green alga Ulva (Enteromorpha) prolifera to abalone Haliotis discus hannai","volume":"29","author":"Wang","year":"2011","journal-title":"Chin. J. Oceanol. Limnol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"107082","DOI":"10.1016\/j.marenvres.2025.107082","article-title":"Dissipation of Ulva prolifera green tides across various spatial and temporal scales and the short-term effects on marine environments","volume":"207","author":"Sun","year":"2025","journal-title":"Mar. Environ. Res."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"736378","DOI":"10.1016\/j.aquaculture.2021.736378","article-title":"Controlling the source of green tides in the Yellow Sea: NaClO treatment of Ulva attached on Pyropia aquaculture rafts","volume":"535","author":"Liu","year":"2021","journal-title":"Aquaculture"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"888","DOI":"10.1016\/j.marpolbul.2009.01.013","article-title":"World\u2019s largest macroalgal bloom caused by expansion of seaweed aquaculture in China","volume":"58","author":"Liu","year":"2009","journal-title":"Mar. Pollut. Bull."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"210","DOI":"10.1016\/j.marpolbul.2015.01.014","article-title":"Changes to the biomass and species composition of Ulva sp. on Porphyra aquaculture rafts, along the coastal radial sandbank of the Southern Yellow Sea","volume":"93","author":"Huo","year":"2015","journal-title":"Mar. Pollut. Bull."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1007\/s13131-019-1452-0","article-title":"The contribution of attached Ulva prolifera on Pyropia aquaculture rafts to green tides in the Yellow Sea","volume":"39","author":"Han","year":"2020","journal-title":"Acta Oceanol. Sin."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1016\/j.hal.2016.09.005","article-title":"Tracing the origin of green macroalgal blooms based on the large scale spatio-temporal distribution of Ulva microscopic propagules and settled mature Ulva vegetative thalli in coastal regions of the Yellow Sea, China","volume":"59","author":"Huo","year":"2016","journal-title":"Harmful Algae"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"825","DOI":"10.1093\/nsr\/nwz026","article-title":"Ulva prolifera green-tide outbreaks and their environmental impact in the Yellow Sea, China","volume":"6","author":"Zhang","year":"2019","journal-title":"Natl. Sci. Rev."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1169","DOI":"10.1016\/j.marpolbul.2011.03.040","article-title":"Inter- and intra-annual patterns of Ulva prolifera green tides in the Yellow Sea during 2007\u20132009, their origin and relationship to the expansion of coastal seaweed aquaculture in China","volume":"62","author":"Keesing","year":"2011","journal-title":"Mar. Pollut. Bull."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"106953","DOI":"10.1016\/j.marenvres.2025.106953","article-title":"Continuous monitoring and comparative genetic analysis of attached outbreak species of Ulva prolifera in the Southern Yellow Sea","volume":"204","author":"Xia","year":"2025","journal-title":"Mar. Environ. Res."},{"key":"ref_16","first-page":"103585","article-title":"Distribution of Ulva macroalgae before and after the outbreak of the Yellow Sea green tide in aquaculture ponds and rivers nearshore Jiangsu, China","volume":"76","author":"Li","year":"2024","journal-title":"Reg. Stud. Mar. Sci."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"114407","DOI":"10.1016\/j.marpolbul.2022.114407","article-title":"Biomass and species composition of green macroalgae in the Binhai Harbor intertidal zone of the southern Yellow Sea","volume":"186","author":"Xia","year":"2022","journal-title":"Mar. Pollut. Bull."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"121969","DOI":"10.1016\/j.envpol.2023.121969","article-title":"Attached Ulva meridionalis on nearshore dikes may pose a new ecological risk in the Yellow Sea","volume":"332","author":"Xia","year":"2023","journal-title":"Environ. Pollut."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1016\/j.marenvres.2009.10.007","article-title":"Tracking the algal origin of the Ulva bloom in the Yellow Sea by a combination of molecular, morphological and physiological analyses","volume":"69","author":"Pang","year":"2010","journal-title":"Mar. Environ. Res."},{"key":"ref_20","first-page":"291","article-title":"Analysis of relationships between onshore aquaculture ponds in Jiangsu and the source of green tide in the South Yellow Sea","volume":"3","author":"Liu","year":"2016","journal-title":"Mar. Fish."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"113646","DOI":"10.1016\/j.marpolbul.2022.113646","article-title":"Prevention strategies for green tides at source in the Southern Yellow Sea","volume":"178","author":"Sun","year":"2022","journal-title":"Mar. Pollut. Bull."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"115468","DOI":"10.1016\/j.marpolbul.2023.115468","article-title":"Morphological characteristics and genetic diversity of floating and attached Ulva prolifera\u2013\u2013A case study in the Yellow Sea, China","volume":"195","author":"Huan","year":"2023","journal-title":"Mar. Pollut. Bull."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"107504","DOI":"10.1016\/j.ecss.2021.107504","article-title":"Analysis of environmental factors affecting the large-scale long-term sequence of green tide outbreaks in the Yellow Sea","volume":"260","author":"Li","year":"2021","journal-title":"Estuar. Coast. Shelf Sci."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"105649","DOI":"10.1016\/j.marenvres.2022.105649","article-title":"Analysis of the reasons for the outbreak of Yellow Sea green tide in 2021 based on long-term multi-source data","volume":"178","author":"Li","year":"2022","journal-title":"Mar. Environ. Res."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"117989","DOI":"10.1016\/j.marpolbul.2025.117989","article-title":"Strong interannual variation of green tides in the southern Yellow Sea: Crucial factors and implications on management strategies","volume":"216","author":"Li","year":"2025","journal-title":"Mar. Pollut. Bull."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"108772","DOI":"10.1016\/j.ecss.2024.108772","article-title":"Review of the development of the green tide and the process of control in the southern Yellow Sea in 2022","volume":"302","author":"Xia","year":"2024","journal-title":"Estuar. Coast. Shelf Sci."},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Xia, L.H., Qin, Y.T., Ji, H.H., Cao, J.X., Wang, X.B., Zhang, Y.H., and Liu, J.L. (2026). Spatiotemporal Dynamics of Micropropagules in Seawater During the 2020 Green Tide Outbreak in the Southern Yellow Sea. Biology, 15.","DOI":"10.3390\/biology15070591"},{"key":"ref_28","unstructured":"CCTV.com (2026, April 05). Experts Discuss Ulva Control: Land-Sea Coordination Is Key to Reducing Marine Eutrophication. 2021. (In Chinese). Available online: https:\/\/ocean.cctv.com\/2021\/07\/27\/ARTIV2mdIMDxj7fVuq7trXYn210714.shtml."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Yao, W.M., Lei, Y.Y., Tan, S.L., Qin, Y.T., Ji, H.H., Sun, Y.Q., Zhang, J.H., and Liu, J.L. (2025). Distribution Characteristics of Suspended Macroalgae in the Southern Yellow Sea Before the Green Tide Outbreak. Biology, 14.","DOI":"10.3390\/biology14101347"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"114710","DOI":"10.1016\/j.marpolbul.2023.114710","article-title":"Advances in the research on micropropagules and their role in green tide outbreaks in the Southern Yellow Sea","volume":"188","author":"Cao","year":"2023","journal-title":"Mar. Pollut. Bull."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"116233","DOI":"10.1016\/j.marpolbul.2024.116233","article-title":"Overwintering and summer survival of Ulva prolifera in sediments: Indoor simulation of temperature impacts","volume":"201","author":"Li","year":"2024","journal-title":"Mar. Pollut. Bull."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Yu, F., Ren, Q., Diao, X.Y., Wei, C.J., and Hu, Y.B. (2022). The Sandwich Structure of the Southern Yellow Sea Cold Water Mass and Yellow Sea Warm Current. Front. Mar. Sci., 8.","DOI":"10.3389\/fmars.2021.767850"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1007\/s13131-022-2031-3","article-title":"Long-term nutrient variation trends and their potential impact on phytoplankton in the southern Yellow Sea, China","volume":"41","author":"Wang","year":"2022","journal-title":"Acta Oceanol. Sin."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"105635","DOI":"10.1016\/j.marenvres.2022.105635","article-title":"An evolving marine environment and its driving forces of algal blooms in the Southern Yellow Sea of China","volume":"178","author":"Xu","year":"2022","journal-title":"Mar. Environ. Res."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"2107","DOI":"10.1007\/s00343-021-1209-4","article-title":"Increased diversity and environmental threat of harmful algal blooms in the Southern Yellow Sea, China","volume":"40","author":"Song","year":"2022","journal-title":"J. Oceanol. Limnol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1007\/s00343-008-0023-6","article-title":"Petroleum geological framework and hydrocarbon potential in the Yellow Sea","volume":"26","author":"Wu","year":"2008","journal-title":"Chin. J. Oceanol. Limnol."},{"key":"ref_37","first-page":"801","article-title":"Tempo-spatial distribution of green algae micro-propagules in the Yellow Sea after the outbreak of the green tide in 2014","volume":"37","author":"Han","year":"2018","journal-title":"Mar. Environ. Sci."},{"key":"ref_38","first-page":"58","article-title":"The Current Situations of Micro-Propagules of Green Algae in Porphyra yezoensis Farming Areas of Southern Jiangsu in Autumn in 2012","volume":"45","author":"Liu","year":"2015","journal-title":"Period. Ocean Univ. China"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"2206","DOI":"10.4319\/lo.2013.58.6.2206","article-title":"Green algae blooms caused by Ulva prolifera in the Southern Yellow Sea: Identification of the original bloom location and evaluation of biological processes occurring during the early northward floating period","volume":"58","author":"Huo","year":"2013","journal-title":"Limnol. Oceanogr."},{"key":"ref_40","unstructured":"Ding, L.P., and Luan, R.X. (2013). Flora Algarum Marinarum Sinicarum. Tomus IV, Chlorophyta. No.I, Ulotrichales Chaetophorales Phaeophilales Ulvales Prasiolales Cladophorales Acrosiphoniales, Science Press. (In Chinese)."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"46","DOI":"10.1016\/j.hal.2013.05.018","article-title":"Seasonal variation of dominant free-floating and attached Ulva species in Rudong coastal area, China","volume":"28","author":"Han","year":"2013","journal-title":"Harmful Algae"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1016\/j.hal.2018.03.007","article-title":"Variations of dominant free-floating Ulva species in the source area for the world\u2019s largest macroalgal blooms, China: Differences of ecological tolerance","volume":"74","author":"Wang","year":"2018","journal-title":"Harmful Algae"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"549","DOI":"10.1016\/j.marpolbul.2017.09.060","article-title":"Effect of the large-scale green tide on the species succession of green macroalgal micro-propagules in the coastal waters of Qingdao, China","volume":"126","author":"Miao","year":"2018","journal-title":"Mar. Pollut. Bull."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1016\/j.ecss.2014.08.006","article-title":"Temporal and spatial distributions of green algae micro-propagules in the coastal waters of the Subei Shoal, China","volume":"163","author":"Song","year":"2015","journal-title":"Estuar. Coast. Shelf Sci."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"173658","DOI":"10.1016\/j.scitotenv.2024.173658","article-title":"Dynamics of green macroalgal micro-propagules and the influencing factors in the southern Yellow Sea, China","volume":"940","author":"Ma","year":"2024","journal-title":"Sci. Total Environ."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1016\/j.marpolbul.2018.03.036","article-title":"Rapid expansion of Ulva blooms in the Yellow Sea, China through sexual reproduction and vegetative growth","volume":"130","author":"Cui","year":"2018","journal-title":"Mar. Pollut. Bull."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"106671","DOI":"10.1016\/j.marenvres.2024.106671","article-title":"Why did the world\u2019s largest green tides occur exclusively in the southern Yellow Sea?","volume":"200","author":"Shao","year":"2024","journal-title":"Mar. Environ. Res."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"40","DOI":"10.1016\/j.hal.2014.05.013","article-title":"Tempo-spatial distribution and species diversity of green algae micro-propagules in the Yellow Sea during the large-scale green tide development","volume":"39","author":"Li","year":"2014","journal-title":"Harmful Algae"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"2462","DOI":"10.1007\/s00343-022-1365-1","article-title":"Tempo-spatial distribution of Ulva spp. micro-propagules in the Yellow Sea during and after green tide in 2019","volume":"40","author":"Zhao","year":"2022","journal-title":"J. Oceanol. Limnol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"e25220","DOI":"10.1016\/j.heliyon.2024.e25220","article-title":"The ever-lasting green tides: What can we do?","volume":"10","author":"Ren","year":"2024","journal-title":"Heliyon"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"102061","DOI":"10.1016\/j.hal.2021.102061","article-title":"Harmful macroalgal blooms (HMBs) in China\u2019s coastal water: Green and golden tides","volume":"107","author":"Xiao","year":"2021","journal-title":"Harmful Algae"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"17018","DOI":"10.1111\/gcb.17018","article-title":"Shift in algal blooms from micro- to macroalgae around China with increasing eutrophication and climate change","volume":"30","author":"Feng","year":"2024","journal-title":"Glob. Change Biol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1007\/s11802-024-5507-2","article-title":"Eutrophication of Jiangsu Coastal Water and Its Role in the Formation of Green Tide","volume":"23","author":"Xiao","year":"2024","journal-title":"J. Ocean Univ. China"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"111197","DOI":"10.1016\/j.marpolbul.2020.111197","article-title":"Role of nutrients in the development of floating green tides in the Southern Yellow Sea, China, in 2017","volume":"156","author":"Zhang","year":"2020","journal-title":"Mar. Pollut. Bull."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1016\/j.marenvres.2016.06.004","article-title":"A review of the green tides in the Yellow Sea, China","volume":"119","author":"Liu","year":"2016","journal-title":"Mar. Environ. Res."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"114789","DOI":"10.1016\/j.marpolbul.2023.114789","article-title":"The drifting and spreading mechanism of floating Ulva mass in the waterways of Subei shoal, the Yellow Sea of China-Application for abating the world\u2019s largest green tides","volume":"190","author":"Wang","year":"2023","journal-title":"Mar. Pollut. Bull."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"110678","DOI":"10.1016\/j.marpolbul.2019.110678","article-title":"Remote sensing methods for biomass estimation of green algae attached to nursery-nets and raft rope","volume":"150","author":"Jiang","year":"2020","journal-title":"Mar. Pollut. Bull."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"408","DOI":"10.1016\/j.marpolbul.2018.01.061","article-title":"A UAV and S2A data-based estimation of the initial biomass of green algae in the South Yellow Sea","volume":"128","author":"Xu","year":"2018","journal-title":"Mar. Pollut. Bull."},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Moulager, M., Corellou, F., Verg\u00e9, V., Escande, M.L., and Bouget, F.Y. (2010). Integration of Light Signals by the Retinoblastoma Pathway in the Control of S Phase Entry in the Picophytoplanktonic Cell Ostreococcus. PLoS Genet., 6.","DOI":"10.1371\/journal.pgen.1000957"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1081\/CBI-200062348","article-title":"Functional proteomics: A promising approach to find novel components of the circadian system","volume":"22","author":"Wagner","year":"2005","journal-title":"Chronobiol. Int."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1134\/S1063074007040013","article-title":"New rhythmic changes in mitosis and growth in low differentiated green and red marine macroalgae","volume":"33","author":"Kalita","year":"2007","journal-title":"Russ. J. Mar. Biol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"60","DOI":"10.1007\/BF00201624","article-title":"Circadian-rhythm and fast responses to blue-light of photosynthesis in ectocarpus (Phaeophyta, Ectocarpales). 2. Light and CO2 dependence of photosynthesis","volume":"187","author":"Schmid","year":"1992","journal-title":"Planta"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1007\/BF00201623","article-title":"Circadian-rhythm and fast responses to blue-light of photosynthesis in ectocarpus (Phaeophyta, Ectocarpales). 1. Characterization of the rhythm and the blue-light response","volume":"187","author":"Schmid","year":"1992","journal-title":"Planta"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"612","DOI":"10.1111\/j.0022-3646.1994.00612.x","article-title":"Kinetics of blue-light stimulation and circadian rhythmicity of light-saturated photosynthesis in brown-algae\u2014A species comparison","volume":"30","author":"Schmid","year":"1994","journal-title":"J. Phycol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1016\/S0006-3495(86)83471-3","article-title":"Magnetite and Magnetotaxis in Bacteria and Algae","volume":"50","author":"Dearaujo","year":"1986","journal-title":"Biophys. J."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"2879","DOI":"10.1111\/1462-2920.12186","article-title":"Marine biofilms on artificial surfaces: Structure and dynamics","volume":"15","author":"Salta","year":"2013","journal-title":"Environ. Microbiol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"RP104797","DOI":"10.7554\/eLife.104797.3","article-title":"Magnetotactic bacteria optimally navigate natural pore networks","volume":"14","author":"Petroff","year":"2025","journal-title":"eLife"},{"key":"ref_68","first-page":"35","article-title":"Sustainable approaches in microalgal biomass harvesting: Current advances and future perspectives","volume":"3","author":"Rahul","year":"2024","journal-title":"Int. J. Agric. Biosci."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"102436","DOI":"10.1016\/j.seares.2023.102436","article-title":"Distribution of Ulva prolifera, the dominant species in green tides along the Jiangsu Province coast in the Southern Yellow Sea, China","volume":"196","author":"Xia","year":"2023","journal-title":"J. Sea Res."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"2298","DOI":"10.1007\/s11434-012-5441-3","article-title":"The yellow sea green tides were dominated by one species, Ulva (Enteromorpha) prolifera, from 2007 to 2011","volume":"58","author":"Zhao","year":"2013","journal-title":"Chin. Sci. Bull."},{"key":"ref_71","first-page":"93","article-title":"Community structure and controlled factor of attached green algae on the Porphyra yezoensis aquaculture rafts in the Subei Shoal, China","volume":"34","author":"Li","year":"2015","journal-title":"Acta Oceanol. Sin."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"102","DOI":"10.1007\/s13131-018-1169-5","article-title":"Source of propagules of the fouling green macroalgae in the Subei Shoal, China","volume":"37","author":"Song","year":"2018","journal-title":"Acta Oceanol. Sin."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"2813","DOI":"10.1111\/are.14620","article-title":"Effects of Chaetomorpha valida bloom development on the structure and succession of the phytoplankton community in Apostichopus japonicus culture ponds","volume":"51","author":"Chen","year":"2020","journal-title":"Aquac. Res."},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Lin, G.R., Sun, F.L., Wang, C.Z., Zhang, L., and Zhang, X.Z. (2017). Assessment of the effect of Enteromorpha prolifera on bacterial community structures in aquaculture environment. PLoS ONE, 12.","DOI":"10.1371\/journal.pone.0179792"},{"key":"ref_75","first-page":"613","article-title":"Rapid attachment of spores of the fouling alga Ulva fasciata on biofilms","volume":"29","author":"Shin","year":"2008","journal-title":"J. Environ. Biol."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1016\/j.ecss.2014.12.026","article-title":"Comparative study of the germination of Ulva prolifera gametes on various substrates","volume":"163","author":"Geng","year":"2015","journal-title":"Estuar. Coast. Shelf Sci."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"515","DOI":"10.1007\/s10811-010-9620-9","article-title":"An investigation of the relationship between sediment particles size and the development of green algal mats (Ulva prolifera) on the intertidal flats of Muan, Korea","volume":"23","author":"Park","year":"2011","journal-title":"J. Appl. Phycol."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1515\/bot-2013-0024","article-title":"The importance of substratum and elevation in recruitment and persistence of ulvoid algal blooms on rocky intertidal shores of the southern Korean coast","volume":"57","author":"Park","year":"2014","journal-title":"Bot. Mar."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"136131","DOI":"10.1016\/j.jhazmat.2024.136131","article-title":"Synergistic effects of plastic debris and elevated nitrate concentrations on the proliferation of Ulva lactuca micro-propagules","volume":"480","author":"Thirumurugan","year":"2024","journal-title":"J. Hazard. Mater."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1016\/j.marenvres.2012.10.007","article-title":"Understanding the recurrent large-scale green tide in the Yellow Sea: Temporal and spatial correlations between multiple geographical, aquacultural and biological factors","volume":"83","author":"Liu","year":"2013","journal-title":"Mar. Environ. Res."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"2077","DOI":"10.1016\/j.marpolbul.2012.07.021","article-title":"Molecular identification of green algae from the rafts based infrastructure of Porphyra yezoensis","volume":"64","author":"Shen","year":"2012","journal-title":"Mar. Pollut. Bull."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1007\/s13131-014-0545-z","article-title":"Development and utility of EST-SSR markers in Ulva prolifera of the South Yellow Sea","volume":"33","author":"Zhang","year":"2014","journal-title":"Acta Oceanol. Sin."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"108","DOI":"10.1007\/s13131-022-1985-5","article-title":"Population differentiation in the dominant species (Ulva prolifera) of green tide in coastal waters of China","volume":"41","author":"Han","year":"2022","journal-title":"Acta Oceanol. Sin."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"533","DOI":"10.1007\/s00343-023-3060-2","article-title":"Genome-wide SNP markers provided insights into the reproductive strategy and genetic diversity of the green tide causative species Ulva prolifera in China","volume":"42","author":"Sun","year":"2024","journal-title":"J. Oceanol. Limnol."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"117834","DOI":"10.1016\/j.marpolbul.2025.117834","article-title":"Estimating windage coefficient of floating marine green tide of Ulva prolifera using UAV optical images","volume":"215","author":"Zheng","year":"2025","journal-title":"Mar. Pollut. Bull."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"105756","DOI":"10.1016\/j.marenvres.2022.105756","article-title":"Distribution characteristics of green tides and its impact on environment in the Yellow Sea","volume":"181","author":"Wang","year":"2022","journal-title":"Mar. Environ. Res."},{"key":"ref_87","doi-asserted-by":"crossref","unstructured":"Zhao, S., Liu, J.L., Xia, Z.Y., Sun, J.Y., Zhang, J.H., and He, P.M. (2025). Subtypes I and II of Ulva prolifera of M\u00fcller: Dominant Green Tide Species in the Southern Yellow Sea and Their Responses to Natural Light and Temperature Conditions. Biology, 14.","DOI":"10.3390\/biology14060702"},{"key":"ref_88","doi-asserted-by":"crossref","unstructured":"Jahan, K., Kristyanto, S., and Choi, K.H. (2025). Genome-Wide DNA Methylation and Transcription Analysis Reveal the Potential Epigenetic Mechanism of Heat-Light Stress Response in the Green Macro Algae Ulva prolifera. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms26136169"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"660","DOI":"10.1016\/j.marpolbul.2015.10.033","article-title":"Adaptability of free-floating green tide algae in the Yellow Sea to variable temperature and light intensity","volume":"101","author":"Cui","year":"2015","journal-title":"Mar. Pollut. Bull."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"468","DOI":"10.32615\/ps.2021.036","article-title":"Impacts of low pH and low salinity induced by acid rain on the photosynthetic activity of green tidal alga Ulva prolifera","volume":"59","author":"Zhao","year":"2021","journal-title":"Photosynthetica"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"105695","DOI":"10.1016\/j.marenvres.2022.105695","article-title":"Physiological acclimation of Ulva prolifera to seasonal environmental factors drives green tides in the Yellow Sea","volume":"179","author":"Wu","year":"2022","journal-title":"Mar. Environ. Res."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1016\/j.hal.2016.09.004","article-title":"Changes in morphological plasticity of Ulva prolifera under different environmental conditions: A laboratory experiment","volume":"59","author":"Gao","year":"2016","journal-title":"Harmful Algae"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"4385","DOI":"10.1007\/s10811-025-03704-9","article-title":"Interactive effects of CO2 concentration and temperature on photosynthesis and growth of cultivated Ulva prolifera germling clusters","volume":"37","author":"Yamaha","year":"2025","journal-title":"J. Appl. Phycol."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1007\/s11802-023-5260-y","article-title":"The Overwintering Capability of Ulva prolifera Spores and Gametes in the Yellow Sea, China","volume":"22","author":"Han","year":"2023","journal-title":"J. Ocean Univ. China"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1016\/j.ecss.2014.08.007","article-title":"Effects of temperature on the germination of green algae micro-propagules in coastal waters of the Subei Shoal, China","volume":"163","author":"Song","year":"2015","journal-title":"Estuar. Coast. Shelf Sci."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"117393","DOI":"10.1016\/j.marpolbul.2024.117393","article-title":"New insights on the impact of light, photoperiod and temperature on the reproduction of green algae Ulva prolifera via transcriptomics and physiological analyses","volume":"211","author":"Jiang","year":"2025","journal-title":"Mar. Pollut. Bull."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"108","DOI":"10.1007\/s13131-015-0654-3","article-title":"Altered seawater salinity levels affected growth and photosynthesis of Ulva fasciata (Ulvales, Chlorophyta) germlings","volume":"34","author":"Chen","year":"2015","journal-title":"Acta Oceanol. Sin."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"1971","DOI":"10.1007\/s10811-023-02960-x","article-title":"Salinity as a tool for strain selection in recirculating land-based production of Ulva spp. from germlings to adults","volume":"35","author":"Cardoso","year":"2023","journal-title":"J. Appl. Phycol."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1007\/s10811-010-9544-4","article-title":"Differential response of varying salinity and temperature on zoospore induction, regeneration and daily growth rate in Ulva fasciata (Chlorophyta, Ulvales)","volume":"23","author":"Mantri","year":"2011","journal-title":"J. Appl. Phycol."},{"key":"ref_100","doi-asserted-by":"crossref","unstructured":"Imchen, T. (2012). Recruitment Potential of a Green Alga Ulva flexuosa Wulfen Dark Preserved Zoospore and Its Development. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0032651"},{"key":"ref_101","doi-asserted-by":"crossref","unstructured":"Liu, J.L., Xia, Z.Y., Zeng, Y.Q., Xia, J., and He, P.M. (2025). Exploration and implication of green macroalgal proliferation in the Nanhui-east-tidal-flat: An investigation of post-reclamation mudflat wetlands. Front. Mar. Sci., 12.","DOI":"10.3389\/fmars.2025.1505586"},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"106719","DOI":"10.1016\/j.marenvres.2024.106719","article-title":"Temporal succession of micropropagules during accumulation and dissipation of green tide algae: A case study in Rudong coast, Jiangsu Province","volume":"202","author":"Xia","year":"2024","journal-title":"Mar. Environ. Res."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"84","DOI":"10.1016\/j.jembe.2005.11.003","article-title":"Small-scale burial of macroalgal detritus in marine sediments: Effects of Ulva spp. on the spatial distribution of macrofauna assemblages","volume":"332","author":"Rossi","year":"2006","journal-title":"J. Exp. Mar. Biol. Ecol."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"384","DOI":"10.1007\/s10021-020-00524-5","article-title":"Large Infaunal Bivalves Determine Community Uptake of Macroalgal Detritus and Food Web Pathways","volume":"24","author":"Karlson","year":"2021","journal-title":"Ecosystems"},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"116268","DOI":"10.1016\/j.watres.2020.116268","article-title":"DOC dynamics and bacterial community succession during long-term degradation of Ulva prolifera and their implications for the legacy effect of green tides on refractory DOC pool in seawater","volume":"185","author":"Chen","year":"2020","journal-title":"Water Res."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1016\/S0304-3770(02)00167-5","article-title":"Decomposition dynamics of the bloom forming macroalga Ulva rigida C. Agardh determined using a 14C-carbon radio-tracer technique","volume":"75","author":"Castaldelli","year":"2003","journal-title":"Aquat. Bot."},{"key":"ref_107","doi-asserted-by":"crossref","unstructured":"Liu, X.X., Zang, Y., Fan, S.L., Miao, X.X., Fu, M.Z., Ma, X.J., Li, M., Zhang, X.L., Wang, Z.L., and Xiao, J. (2024). Changes in the structure of the microbial community within the phycospheric microenvironment and potential biogeochemical effects induced in the demise stage of green tides caused by Ulva prolifera. Front. Microbiol., 15.","DOI":"10.3389\/fmicb.2024.1507660"},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"106842","DOI":"10.1016\/j.marenvres.2024.106842","article-title":"Herbivore grazing enhances macroalgal organic carbon release and alters their carbon sequestration fate in the ocean","volume":"203","author":"Li","year":"2025","journal-title":"Mar. Environ. Res."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"69","DOI":"10.3354\/meps09173","article-title":"Top-down and bottom-up regulation in a high nutrient-high herbivory coastal ecosystem","volume":"432","author":"Martinetto","year":"2011","journal-title":"Mar. Ecol. Prog. Ser."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"103343","DOI":"10.1016\/j.aquabot.2020.103343","article-title":"Responses of the germination and growth of Ulva prolifera parthenogametes, the causative species of green tides, to gradients of temperature and light","volume":"170","author":"Huo","year":"2021","journal-title":"Aquat. Bot."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1007\/s00367-012-0282-2","article-title":"Control of sediment supply, palaeoceanography and morphology on late Quaternary sediment dynamics at the Galician continental slope","volume":"32","author":"Bender","year":"2012","journal-title":"Geo-Mar. Lett."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"905","DOI":"10.1002\/lom3.10211","article-title":"The influence of coring method on the preservation of sedimentary and biogeochemical features when sampling soft-bottom, shallow coastal environments","volume":"15","author":"Mogg","year":"2017","journal-title":"Limnol. Oceanogr. Methods"},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"1021","DOI":"10.4319\/lo.1991.36.5.1021","article-title":"Core compression and surficial sediment loss of lake-sediments of high porosity caused by gravity coring","volume":"36","author":"Crusius","year":"1991","journal-title":"Limnol. Oceanogr."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"4951","DOI":"10.5194\/essd-13-4951-2021","article-title":"A worldwide meta-analysis (1977\u20132020) of sediment core dating using fallout radionuclides including 137Cs and 210Pbxs","volume":"13","author":"Foucher","year":"2021","journal-title":"Earth Syst. Sci. Data"},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"146522","DOI":"10.1016\/j.scitotenv.2021.146522","article-title":"A review of current and emerging approaches for Quaternary marine sediment dating","volume":"780","author":"Li","year":"2021","journal-title":"Sci. Total Environ."},{"key":"ref_116","first-page":"983","article-title":"Vertical distribution and species diversity of microscopic propagules of green macroalgae in sediments of Subei Shoal, China","volume":"49","author":"Liu","year":"2018","journal-title":"Oceanol. Limnol. Sin."}],"container-title":["Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2079-7737\/15\/15\/1220\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,25]],"date-time":"2026-07-25T04:19:51Z","timestamp":1784953191000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2079-7737\/15\/15\/1220"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,23]]},"references-count":116,"journal-issue":{"issue":"15","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["biology15151220"],"URL":"https:\/\/doi.org\/10.3390\/biology15151220","relation":{},"ISSN":["2079-7737"],"issn-type":[{"value":"2079-7737","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,23]]}}}