{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,13]],"date-time":"2026-10-13T00:19:30Z","timestamp":1791850770790,"version":"4.3.4"},"reference-count":117,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T00:00:00Z","timestamp":1788912000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Innovative Drug Research and Development-National Science and Technology Major Project","award":["2026ZD18069001"],"award-info":[{"award-number":["2026ZD18069001"]}]},{"DOI":"10.13039\/501100007129","name":"Shandong Provincial Natural Science Foundation","doi-asserted-by":"publisher","award":["ZR2023MH302"],"award-info":[{"award-number":["ZR2023MH302"]}],"id":[{"id":"10.13039\/501100007129","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012226","name":"Fundamental Research Funds for the Central Universities","doi-asserted-by":"publisher","award":["202561057"],"award-info":[{"award-number":["202561057"]}],"id":[{"id":"10.13039\/501100012226","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Mar. Drugs","Marine Drugs"],"abstract":"<jats:p>Hyperlipidemia is a chronic metabolic disorder in humans, contributing to the onset of cardiovascular diseases (CVDs) that remain the leading cause of death worldwide. Current clinical antihyperlipidemic agents are often accompanied by diverse adverse side effects. Therefore, developing safer novel lipid regulators is an urgent and challenging task. Marine polysaccharides exhibit remarkable antihyperlipidemic activity owing to their distinctive physicochemical properties, multi-target mechanisms, and system-level modulation. This review systematically summarizes recent progress regarding the antihyperlipidemic efficacy of marine polysaccharides and provides comprehensive insights into their sources, structural features, structure\u2013activity relationships, mechanisms of action, and application potential. Furthermore, this review also highlights the existing limitations and challenges in translational research of marine polysaccharides, and puts forward future research directions to advance marine polysaccharides into safe, effective, and sustainable natural therapeutics for hyperlipidemia.<\/jats:p>","DOI":"10.3390\/md24090315","type":"journal-article","created":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T08:05:43Z","timestamp":1788941143000},"page":"315","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Recent Advances in the Antihyperlipidemic Activity of Marine Polysaccharides"],"prefix":"10.3390","volume":"24","author":[{"given":"Yongxuan","family":"Liu","sequence":"first","affiliation":[{"name":"Key Laboratory of Marine Drugs of Ministry of Education, Shandong Key Laboratory of Glycoscience and Glycotherapeutics, School of Medicine and Pharmacy, Ocean University of China, Qingdao 266003, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Haowei","family":"Liu","sequence":"additional","affiliation":[{"name":"Key Laboratory of Marine Drugs of Ministry of Education, Shandong Key Laboratory of Glycoscience and Glycotherapeutics, School of Medicine and Pharmacy, Ocean University of China, Qingdao 266003, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dan","family":"Li","sequence":"additional","affiliation":[{"name":"Key Laboratory of Marine Drugs of Ministry of Education, Shandong Key Laboratory of Glycoscience and Glycotherapeutics, School of Medicine and Pharmacy, Ocean University of China, Qingdao 266003, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shengzhou","family":"Ma","sequence":"additional","affiliation":[{"name":"Key Laboratory of Marine Drugs of Ministry of Education, Shandong Key Laboratory of Glycoscience and Glycotherapeutics, School of Medicine and Pharmacy, Ocean University of China, Qingdao 266003, China"},{"name":"Laboratory for Marine Drugs and Bioproducts, Qingdao Marine Science and Technology Center, Qingdao 266237, China"},{"name":"Laboratory of Marine Glycodrugs Research and Development, Marine Biomedical Research Institute of Qingdao, Qingdao 266071, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8102-137X","authenticated-orcid":false,"given":"Chunxia","family":"Li","sequence":"additional","affiliation":[{"name":"Key Laboratory of Marine Drugs of Ministry of Education, Shandong Key Laboratory of Glycoscience and Glycotherapeutics, School of Medicine and Pharmacy, Ocean University of China, Qingdao 266003, China"},{"name":"Laboratory for Marine Drugs and Bioproducts, Qingdao Marine Science and Technology Center, Qingdao 266237, China"},{"name":"Laboratory of Marine Glycodrugs Research and Development, Marine Biomedical Research Institute of Qingdao, Qingdao 266071, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,9]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"4674","DOI":"10.1016\/j.bmc.2007.04.031","article-title":"The biology and chemistry of hyperlipidemia","volume":"15","author":"Jain","year":"2007","journal-title":"Bioorg. Med. Chem."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1016\/j.addr.2020.07.019","article-title":"Lipoproteins and lipids in cardiovascular disease: From mechanistic insights to therapeutic targeting","volume":"159","author":"Soppert","year":"2020","journal-title":"Adv. Drug Deliv. Rev."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1016\/j.pop.2012.11.003","article-title":"Hyperlipidemia as a Risk Factor for Cardiovascular Disease","volume":"40","author":"Nelson","year":"2013","journal-title":"Prim. Care"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"308","DOI":"10.1186\/s41043-025-01054-3","article-title":"Global prevalence of dyslipidemias in the general adult population: A systematic review and meta-analysis","volume":"44","year":"2025","journal-title":"J. Health Popul. Nutr."},{"key":"ref_5","first-page":"111","article-title":"Comprehensive Review of Hyperlipidemia: Pathophysiology, Diagnosis, and Management","volume":"12","author":"Rani","year":"2024","journal-title":"Int. J. All. Res. Educ. Sci. Methods"},{"key":"ref_6","first-page":"118","article-title":"Hyperlipidemia: Pathophysiology, causes, complications, and treatment. A review","volume":"1","author":"Naser","year":"2021","journal-title":"Kerbala J. Pharm. Pharm. Sci."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1038\/s41392-023-01690-3","article-title":"Targeting proprotein convertase subtilisin\/kexin type 9 (PCSK9): From bench to bedside","volume":"9","author":"Bao","year":"2024","journal-title":"Signal Transduct. Tar."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"503","DOI":"10.1038\/s41467-017-00568-7","article-title":"Heparan sulfate proteoglycans present PCSK9 to the LDL receptor","volume":"8","author":"Gustafsen","year":"2017","journal-title":"Nat. Commun."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"140585","DOI":"10.1016\/j.ijbiomac.2025.140585","article-title":"Polyguluronate sulfate has potential to mitigate hyperlipidemia by inhibiting the PCSK9-mediated degradation of LDLR","volume":"302","author":"Li","year":"2025","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"1125","DOI":"10.1172\/JCI0215593","article-title":"SREBPs: Activators of the complete program of cholesterol and fatty acid synthesis in the liver","volume":"109","author":"Horton","year":"2002","journal-title":"J. Clin. Invest."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1016\/j.ecl.2022.02.008","article-title":"Lipid and Lipoprotein Metabolism","volume":"51","author":"Feingold","year":"2022","journal-title":"Endocrin Metab. Clin."},{"key":"ref_12","first-page":"1997","article-title":"Statin Use for the Primary Prevention of Cardiovascular Disease in Adults","volume":"328","author":"Mangione","year":"2022","journal-title":"J. Am. Med. Assoc."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1943","DOI":"10.1161\/01.CIR.0000034044.95911.DC","article-title":"Inhibition of Intestinal Cholesterol Absorption by Ezetimibe in Humans","volume":"106","author":"Sudhop","year":"2002","journal-title":"Circulation"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1007\/s11739-011-0653-0","article-title":"Use and indications of cholestyramine and bile acid sequestrants","volume":"8","author":"Scaldaferri","year":"2013","journal-title":"Intern. Emerg. Med."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1186\/1475-2840-11-125","article-title":"Fibrates are an essential part of modern anti-dyslipidemic arsenal: Spotlight on atherogenic dyslipidemia and residual risk reduction","volume":"11","author":"Tenenbaum","year":"2012","journal-title":"Cardiovasc. Diabetol."},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Naranjo, M.C., Millan-Linares, M.C., and Montserrat-de la Paz, S. (2020). Niacin and hyperlipidemia. Molecular Nutrition, Elsevier.","DOI":"10.1016\/B978-0-12-811907-5.00008-7"},{"key":"ref_17","first-page":"277","article-title":"Evolution of Omega-3 Fatty Acid Therapy and Current and Future Role in the Management of Dyslipidemia","volume":"36","author":"Benes","year":"2018","journal-title":"Cardiol. Clin."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1038\/s41569-018-0107-8","article-title":"PCSK9 inhibitors: Clinical evidence and implementation","volume":"16","author":"Sabatine","year":"2019","journal-title":"Nat. Rev. Cardiol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"2178","DOI":"10.1016\/j.jacc.2013.07.081","article-title":"Mipomersen, an Apolipoprotein B Synthesis Inhibitor, Reduces Atherogenic Lipoproteins in Patients With Severe Hypercholesterolemia at High Cardiovascular Risk","volume":"62","author":"Thomas","year":"2013","journal-title":"J. Am. Coll. Cardiol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"40","DOI":"10.1016\/S0140-6736(12)61731-0","article-title":"Efficacy and safety of a microsomal triglyceride transfer protein inhibitor in patients with homozygous familial hypercholesterolaemia: A single-arm, open-label, phase 3 study","volume":"381","author":"Cuchel","year":"2013","journal-title":"Lancet"},{"key":"ref_21","first-page":"427","article-title":"Side effects of anti-lipid medications","volume":"Volume 46","author":"Mensah","year":"2023","journal-title":"Side Effects of Drugs Annual"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"100047","DOI":"10.1016\/j.glycos.2026.100047","article-title":"Development of marine organisms derived carbohydrate-based drugs: From bioactivity to clinical translation","volume":"2","author":"Pan","year":"2026","journal-title":"Glycosci. Ther."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Venugopal, V. (2016). Marine Polysaccharides, CRC Press.","DOI":"10.1201\/b10516"},{"key":"ref_24","first-page":"99","article-title":"Algal polysaccharides: Potential bioactive substances for cosmeceutical applications","volume":"39","author":"Kim","year":"2018","journal-title":"Crit. Rev. Biotechnol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"681","DOI":"10.1016\/j.ijbiomac.2022.02.189","article-title":"A comprehensive review on polysaccharides with hypolipidemic activity: Occurrence, chemistry and molecular mechanism","volume":"206","author":"Kalita","year":"2022","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"2106","DOI":"10.3390\/md9102106","article-title":"Important Determinants for Fucoidan Bioactivity: A Critical Review of Structure-Function Relations and Extraction Methods for Fucose-Containing Sulfated Polysaccharides from Brown Seaweeds","volume":"9","author":"Ale","year":"2011","journal-title":"Mar. Drugs"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"106303","DOI":"10.1016\/j.jff.2024.106303","article-title":"The structural characteristics, biological activities and mechanisms of bioactive brown seaweed polysaccharides: A review","volume":"119","author":"Zhou","year":"2024","journal-title":"J. Funct. Food"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1011","DOI":"10.1016\/j.foodres.2016.11.016","article-title":"Polysaccharides from macroalgae: Recent advances, innovative technologies and challenges in extraction and purification","volume":"99","author":"Rajauria","year":"2017","journal-title":"Food Res. Int."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"104142","DOI":"10.1016\/j.tifs.2023.104142","article-title":"Overview of alginate extraction processes: Impact on alginate molecular structure and techno-functional properties","volume":"140","author":"Bojorges","year":"2023","journal-title":"Trends Food Sci. Technol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"8131","DOI":"10.1039\/C3RA23373A","article-title":"Fucoidans from brown seaweeds: An update on structures, extraction techniques and use of enzymes as tools for structural elucidation","volume":"3","author":"Ale","year":"2013","journal-title":"RSC Adv."},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Xing, M.C., Cao, Q., Wang, Y., Xiao, H., Zhao, J.R., Zhang, Q., Ji, A.G., and Song, S.L. (2020). Advances in Research on the Bioactivity of Alginate Oligosaccharides. Mar. Drugs, 18.","DOI":"10.3390\/md18030144"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"102593","DOI":"10.1016\/j.algal.2021.102593","article-title":"Carrageenan oligosaccharides: A comprehensive review of preparation, isolation, purification, structure, biological activities and applications","volume":"61","author":"Guo","year":"2022","journal-title":"Algal Res."},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Cheong, K.L., Qiu, H.M., Du, H., Liu, Y., and Khan, B.M. (2018). Oligosaccharides Derived from Red Seaweed: Production, Properties, and Potential Health and Cosmetic Applications. Molecules, 23.","DOI":"10.3390\/molecules23102451"},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Chadwick, M., Carvalho, L.G., Vanegas, C., and Dimartino, S. (2025). A Comparative Review of Alternative Fucoidan Extraction Techniques from Seaweed. Mar. Drugs, 23.","DOI":"10.3390\/md23010027"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"1349","DOI":"10.1007\/s00396-020-04706-4","article-title":"Branched architecture of fucoidan characterized by dynamic and static light scattering","volume":"298","author":"Gasilova","year":"2020","journal-title":"Colloid. Polym. Sci."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1007\/s00253-011-3666-8","article-title":"Algal fucoidan: Structural and size-dependent bioactivities and their perspectives","volume":"93","author":"Morya","year":"2012","journal-title":"Appl. Microbiol. Biot."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"808","DOI":"10.1016\/j.carbpol.2015.06.088","article-title":"Identification of a highly sulfated fucoidan from sea cucumber with well-repeated tetrasaccharides units","volume":"134","author":"Hu","year":"2015","journal-title":"Carbohydr. Polym."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"129908","DOI":"10.1016\/j.foodchem.2021.129908","article-title":"A fucoidan from Sargassum fusiforme with novel structure and its regulatory effects on intestinal microbiota in high-fat diet-fed mice","volume":"358","author":"Liu","year":"2021","journal-title":"Food Chem."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"119411","DOI":"10.1016\/j.carbpol.2022.119411","article-title":"fucan suppresses high fat diet-induced obesity and enriches fucoidan-degrading gut bacteria","volume":"290","author":"Wei","year":"2022","journal-title":"Carbohydr. Polym."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"103755","DOI":"10.1016\/j.fbio.2024.103755","article-title":"The preparation and anti-atherosclerotic effects of different low-molecular weights fucoidan","volume":"58","author":"Lu","year":"2024","journal-title":"Food Biosci."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"e70310","DOI":"10.1111\/cbdd.70310","article-title":"Fucoidan: An Update on Function, Role in Human Health and Applications","volume":"107","author":"Critelli","year":"2026","journal-title":"Chem. Biol. Drug Des."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"4242","DOI":"10.1039\/D4FO05388E","article-title":"Therapeutic potential of fucoidan in atherosclerosis: A review","volume":"16","author":"Feng","year":"2025","journal-title":"Food Funct."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"188","DOI":"10.1016\/j.ijbiomac.2019.08.002","article-title":"Hypolipidemic effects of fucoidan fractions from Saccharina sculpera (Laminariales, Phaeophyceae)","volume":"140","author":"Ren","year":"2019","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"759","DOI":"10.1016\/j.ijbiomac.2019.05.070","article-title":"The fucoidan A3 from the seaweed Ascophyllum nodosum enhances RCT-related genes expression in hyperlipidemic C57BL\/6J mice","volume":"134","author":"Yang","year":"2019","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"118247","DOI":"10.1016\/j.carbpol.2021.118247","article-title":"Fucoidan reduces lipid accumulation by promoting foam cell autophagy via TFEB","volume":"268","author":"Zhao","year":"2021","journal-title":"Carbohydr. Polym."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"e0144525","DOI":"10.1128\/aem.01445-25","article-title":"Sargassum fusiforme polysaccharides modulate gut microbiota and metabolites to regulate hyperlipidemia in mice fed a high fat diet","volume":"91","author":"Su","year":"2025","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"135223","DOI":"10.1016\/j.ijbiomac.2024.135223","article-title":"Extraction, purification, structural characterization and anti-hyperlipidemia activity of fucoidan from Laminaria digitata","volume":"279","author":"Li","year":"2024","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"616","DOI":"10.1016\/j.ijbiomac.2022.01.102","article-title":"Fucoidan ameliorates glucose metabolism by the improvement of intestinal barrier and inflammatory damage in type 2 diabetic rats","volume":"201","author":"Liu","year":"2022","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_49","unstructured":"Kyodo Milk Industry Co., Ltd. (2001). Agent for Preventing or Treating Hyperlipidemia. (JP2001335491A)."},{"key":"ref_50","unstructured":"Hao, J., Guan, H., Dun, Y., Zhou, X., Yu, G., and Xia, X. (2015). Fucoidan Sulfate and Low-Molecular-Weight Fucoidan Sulfate in the Preparation of Drugs and Health Products for Lowering Blood Lipids and Weight Loss. (CN104523744A)."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"113435","DOI":"10.1016\/j.lwt.2022.113435","article-title":"Impact of alginate block type on the structure and physicochemical properties of curcumin-loaded complex biopolymer nanoparticles","volume":"162","author":"Li","year":"2022","journal-title":"LWT-Food Sci. Technol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"72","DOI":"10.2331\/suisan.69.72","article-title":"Effect of alginate viscosity on digestibility and lipid metabolism in rats","volume":"69","author":"Wang","year":"2003","journal-title":"Nippon. Suisan Gakk"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"304","DOI":"10.1016\/j.ejmech.2014.07.107","article-title":"Activated AMPK explains hypolipidemic effects of sulfated low molecular weight guluronate on HepG2 cells","volume":"85","author":"Liu","year":"2014","journal-title":"Eur. J. Med. Chem."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"625","DOI":"10.3724\/j.issn.1000-0518.2009.6.625628","article-title":"Study of alginate oxidative degradation by hydrogen peroxide","volume":"26","author":"Li","year":"2009","journal-title":"Chin. J. Appl. Chem."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1146\/annurev.micro.54.1.289","article-title":"Alginate lyase: Review of major sources and enzyme characteristics, structure-function analysis, biological roles, and applications","volume":"54","author":"Wong","year":"2000","journal-title":"Annu. Rev. Microbiol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1016\/bs.pmbts.2019.02.007","article-title":"Efficacy of heparinoid PSS in treating cardiovascular diseases and beyond\u2014A review of 31 years clinical experiences in China","volume":"Volume 163","author":"Zhang","year":"2019","journal-title":"Progress in Molecular Biology and Translational Science"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"330","DOI":"10.1016\/j.carbpol.2016.03.001","article-title":"Study on quality control of sulfated polysaccharide drug, propylene glycol alginate sodium sulfate (PSS)","volume":"144","author":"Xue","year":"2016","journal-title":"Carbohydr. Polym."},{"key":"ref_58","first-page":"687","article-title":"Effect of propylene glycol mannate sulfate on blood lipids and lipoprotein lipase in hyperlipidemic rat","volume":"37","author":"Gao","year":"2002","journal-title":"Yaoxue Xuebao"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"26","DOI":"10.1016\/j.ijbiomac.2018.03.068","article-title":"Propylene glycol guluronate sulfate (PGGS) reduces lipid accumulation via AMP-activated kinase activation in palmitate-induced HepG2 cells","volume":"114","author":"Xin","year":"2018","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"1393","DOI":"10.1016\/j.jnutbio.2015.07.009","article-title":"Alginate oligosaccharide enhances LDL uptake via regulation of LDLR and PCSK9 expression","volume":"26","author":"Yang","year":"2015","journal-title":"J. Nutr. Biochem."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1016\/j.ijbiomac.2022.06.073","article-title":"Sodium alginate and galactooligosaccharides ameliorate metabolic disorders and alter the composition of the gut microbiota in mice with high-fat diet-induced obesity","volume":"215","author":"Li","year":"2022","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_62","first-page":"100868","article-title":"Alginate oligosaccharides and glycolipid metabolism: Correlation with gut microbiota modulation in mice and clinical feces","volume":"11","author":"Zhuang","year":"2025","journal-title":"Carbohydr. Polym. Tech."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1007\/s40495-023-00348-6","article-title":"Biological Activities of Carrageenan from Red Algae: A Mini Review","volume":"10","author":"Udayakumar","year":"2023","journal-title":"Curr. Pharmacol. Rep."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1007\/s40495-025-00415-0","article-title":"Exploring Carrageenan: Biological Activities and Applications in Drug Delivery and Tissue Engineering","volume":"11","author":"Edo","year":"2025","journal-title":"Curr. Pharmacol. Rep."},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"du Preez, R., Paul, N., Mouatt, P., Majzoub, M.E., Thomas, T., Panchal, S.K., and Brown, L. (2020). Carrageenans from the Red Seaweed Sarconema filiforme Attenuate Symptoms of Diet-Induced Metabolic Syndrome in Rats. Mar. Drugs, 18.","DOI":"10.3390\/md18020097"},{"key":"ref_66","first-page":"209","article-title":"Blood cholesterol and lipid-lowering effects of carrageenan on human volunteers","volume":"12","author":"Panlasigui","year":"2003","journal-title":"Asia Pac. J. Clin. Nutr."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"115629","DOI":"10.1016\/j.carbpol.2019.115629","article-title":"Effect of carrageenans on some lipid metabolism components","volume":"230","author":"Sokolova","year":"2020","journal-title":"Carbohydr. Polym."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"118830","DOI":"10.1016\/j.carbpol.2021.118830","article-title":"Dietary \u03ba-carrageenan facilitates gut microbiota-mediated intestinal inflammation","volume":"277","author":"Wu","year":"2022","journal-title":"Carbohydr. Polym."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"126401","DOI":"10.1016\/j.ijbiomac.2023.126401","article-title":"Enzymatic hydrolysates of \u03ba-carrageenan by \u03ba-carrageenase-CLEA immobilized on amine-modified ZIF-8 confer hypolipidemic activity in HepG2 cells","volume":"252","author":"Zhu","year":"2023","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"926","DOI":"10.1039\/D0FO02768E","article-title":"Advances in the preparation and assessment of the biological activities of chitosan oligosaccharides with different structural characteristics","volume":"12","author":"Zhou","year":"2021","journal-title":"Food Funct."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"926","DOI":"10.3389\/fmars.2025.1592331","article-title":"Source-dependent variations in chitin: A comparative study on Antarctic krill, white shrimp and crayfish","volume":"12","author":"Yang","year":"2025","journal-title":"Front. Mar. Sci."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1016\/j.pharmthera.2016.10.013","article-title":"Chitosan oligosaccharide: Biological activities and potential therapeutic applications","volume":"170","author":"Muanprasat","year":"2017","journal-title":"Pharmacol. Ther."},{"key":"ref_73","doi-asserted-by":"crossref","unstructured":"Bai, Y., Zheng, J., Yuan, X., Jiao, S., Feng, C., Du, Y., Liu, H., and Zheng, L. (2018). Chitosan Oligosaccharides Improve Glucolipid Metabolism Disorder in Liver by Suppression of Obesity-Related Inflammation and Restoration of Peroxisome Proliferator-Activated Receptor Gamma (PPAR\u03b3). Mar. Drugs, 16.","DOI":"10.3390\/md16110455"},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Liu, S.H., Chiu, C.Y., Shi, C.M., and Chiang, M.T. (2018). Functional Comparison of High and Low Molecular Weight Chitosan on Lipid Metabolism and Signals in High-Fat Diet-Fed Rats. Mar. Drugs, 16.","DOI":"10.20944\/preprints201807.0044.v1"},{"key":"ref_75","doi-asserted-by":"crossref","unstructured":"Pan, H.T., Fu, C.H., Huang, L.L., Jiang, Y., Deng, X.Y., Guo, J., and Su, Z.Q. (2018). Anti-Obesity Effect of Chitosan Oligosaccharide Capsules (COSCs) in Obese Rats by Ameliorating Leptin Resistance and Adipogenesis. Mar. Drugs, 16.","DOI":"10.3390\/md16060198"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"137186","DOI":"10.1016\/j.ijbiomac.2024.137186","article-title":"Chitosan oligosaccharides: A potential therapeutic agent for inhibiting foam cell formation in atherosclerosis","volume":"282","author":"Le","year":"2024","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1016\/j.cca.2013.06.006","article-title":"Foam cells in atherosclerosis","volume":"424","author":"Yu","year":"2013","journal-title":"Clin. Chim. Acta"},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Yan, C., Zhang, C.L., Cao, X.J., Feng, B., and Li, X.L. (2020). Intestinal Population in Host with Metabolic Syndrome during Administration of Chitosan and Its Derivatives. Molecules, 25.","DOI":"10.3390\/molecules25245857"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"120","DOI":"10.1186\/s13098-025-01681-0","article-title":"Changes in gut microbiota following supplementation with chitosan in adolescents with overweight or obesity: A randomized, double-blind clinical trial","volume":"17","author":"Fatahi","year":"2025","journal-title":"Diabetol. Metab. Syndr."},{"key":"ref_80","unstructured":"Kawashima, S., and Murata, Y. (1997). Gel State Composition Having Compound Releasing and Adsorbing Property. (JPH09241169A)."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"117389","DOI":"10.1016\/j.carbpol.2020.117389","article-title":"Structure of a laminarin-type \u03b2-(1\u21923)-glucan from brown algae Sargassum henslowianum and its potential on regulating gut microbiota","volume":"255","author":"Cui","year":"2021","journal-title":"Carbohydr. Polym."},{"key":"ref_82","doi-asserted-by":"crossref","unstructured":"Karuppusamy, S., Rajauria, G., Fitzpatrick, S., Lyons, H., McMahon, H., Curtin, J., Tiwari, B.K., and O\u2019Donnell, C. (2022). Biological Properties and Health-Promoting Functions of Laminarin: A Comprehensive Review of Preclinical and Clinical Studies. Mar. Drugs, 20.","DOI":"10.3390\/md20120772"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"4193","DOI":"10.1039\/C6FO00929H","article-title":"Laminarin favorably modulates gut microbiota in mice fed a high-fat diet","volume":"7","author":"Nguyen","year":"2016","journal-title":"Food Funct."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"424","DOI":"10.1002\/(SICI)1097-0215(19991029)83:3<424::AID-IJC20>3.0.CO;2-L","article-title":"Inhibition of heparanase activity and tumor metastasis by laminarin sulfate and synthetic phosphorothioate oligodeoxynucleotides","volume":"83","author":"Miao","year":"1999","journal-title":"Int. J. Cancer"},{"key":"ref_85","first-page":"e56","article-title":"Macrophage Activation by Heparanase Is Mediated by TLR-2 and TLR-4 and Associates with Plaque Progression","volume":"33","author":"Blich","year":"2013","journal-title":"Arter. Throm. Vas."},{"key":"ref_86","doi-asserted-by":"crossref","unstructured":"He, Z.Q., Zhang, Z.Y., Xu, P.F., Dirsch, V.M., Wang, L.M., and Wang, K.W. (2023). Laminarin Reduces Cholesterol Uptake and NPC1L1 Protein Expression in High-Fat Diet (HFD)-Fed Mice. Mar. Drugs, 21.","DOI":"10.3390\/md21120624"},{"key":"ref_87","doi-asserted-by":"crossref","unstructured":"Alisi, A., Kondo, Y., Hasegawa, G., Okada, H., Senmaru, T., Fukui, M., Nakamura, N., Sawada, M., Kitawaki, J., and Okanoue, T. (2013). Leprdb\/db Mice with Senescence Marker Protein-30 Knockout (Leprdb\/dbSmp30Y\/\u2212) Exhibit Increases in Small Dense-LDL and Severe Fatty Liver Despite Being Fed a Standard Diet. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0065698"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"15691","DOI":"10.1038\/ncomms15691","article-title":"Cellular senescence drives age-dependent hepatic steatosis","volume":"8","author":"Ogrodnik","year":"2017","journal-title":"Nat. Commun."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"153359","DOI":"10.1016\/j.bbrc.2026.153359","article-title":"Laminarin ameliorates hepatic steatosis and upregulates hepatic SMP-30 expression in high-fat-diet-induced NAFLD mice","volume":"805","author":"Hwang","year":"2026","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"101422","DOI":"10.1016\/j.algal.2019.101422","article-title":"Ulvan: A systematic review of extraction, composition and function","volume":"39","author":"Kidgell","year":"2019","journal-title":"Algal Res."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"971","DOI":"10.1016\/j.ijbiomac.2018.02.104","article-title":"Antioxidant activity of purified ulvan in hyperlipidemic mice","volume":"113","author":"Li","year":"2018","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"1059","DOI":"10.1016\/j.ijbiomac.2019.09.198","article-title":"The antioxidant and antihyperlipidemic activities of phosphorylated polysaccharide from","volume":"145","author":"Jiang","year":"2020","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"120249","DOI":"10.1016\/j.carbpol.2022.120249","article-title":"Regulatory effects mediated by ulvan oligosaccharide and its zinc complex on lipid metabolism in high-fat diet-fed mice","volume":"300","author":"Chi","year":"2023","journal-title":"Carbohydr. Polym."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"cwae098","DOI":"10.1093\/glycob\/cwae098","article-title":"Fucosylated chondroitin sulfate, an intriguing polysaccharide from sea cucumber: Past, present, and future","volume":"35","author":"Felix","year":"2025","journal-title":"Glycobiology"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"8284","DOI":"10.1073\/pnas.1504229112","article-title":"Discovery of an intrinsic tenase complex inhibitor: Pure nonasaccharide from fucosylated glycosaminoglycan","volume":"112","author":"Zhao","year":"2015","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.carbpol.2016.03.013","article-title":"Sulfation pattern of fucose branches affects the anti-hyperlipidemic activities of fucosylated chondroitin sulfate","volume":"147","author":"Wu","year":"2016","journal-title":"Carbohydr. Polym."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"330","DOI":"10.1016\/j.carbpol.2017.05.063","article-title":"Macromolecular properties and hypolipidemic effects of four sulfated polysaccharides from sea cucumbers","volume":"173","author":"Li","year":"2017","journal-title":"Carbohydr. Polym."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"634","DOI":"10.1016\/j.carbpol.2018.08.020","article-title":"Fucosylated chondroitin sulfate oligosaccharides from Isostichopus badionotus regulates lipid disorder in C57BL\/6 mice fed a high-fat diet","volume":"201","author":"Li","year":"2018","journal-title":"Carbohydr. Polym."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"137371","DOI":"10.1016\/j.ijbiomac.2024.137371","article-title":"Fucosylated chondroitin sulfate alleviates diet-induced obesity by modulating intestinal lipid metabolism and colonic microflora","volume":"283","author":"Li","year":"2024","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_100","doi-asserted-by":"crossref","unstructured":"He, D., Yan, L., Hu, Y., Wu, Q., Wu, M., Choi, J.-i., and Tong, H. (2021). Optimization of Porphyran Extraction from Pyropia yezoensis by Response Surface Methodology and Its Lipid-Lowering Effects. Mar. Drugs, 19.","DOI":"10.3390\/md19020053"},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"116261","DOI":"10.1016\/j.carbpol.2020.116261","article-title":"Odd-numbered agaro-oligosaccharides alleviate type 2 diabetes mellitus and related colonic microbiota dysbiosis in mice","volume":"240","author":"Wang","year":"2020","journal-title":"Carbohydr. Polym."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"476","DOI":"10.1038\/s41586-025-08792-8","article-title":"Sensing ceramides by CYSLTR2 and P2RY6 to aggravate atherosclerosis","volume":"641","author":"Zhang","year":"2025","journal-title":"Nature"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"452","DOI":"10.1038\/s41586-026-10697-z","article-title":"Dietary cholesterol activates a Ral-dependent pathway driving LDLR turnover","volume":"656","author":"Feng","year":"2026","journal-title":"Nature"},{"key":"ref_104","doi-asserted-by":"crossref","unstructured":"Jim, E.L., Jim, E.L., Surya, R., Permatasari, H.K., and Nurkolis, F. (2025). Marine-Derived Polymers\u2013Polysaccharides as Promising Natural Therapeutics for Atherosclerotic Cardiovascular Disease. Mar. Drugs, 23.","DOI":"10.3390\/md23080325"},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"144797","DOI":"10.1016\/j.ijbiomac.2025.144797","article-title":"Structural characterization of polysaccharides of marine origin: A review","volume":"317","author":"Liu","year":"2025","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_106","doi-asserted-by":"crossref","unstructured":"Imran, M., and Ghadi, S.C. (2019). Role of carbohydrate active enzymes (CAZymes) in production of marine bioactive oligosaccharides and their pharmacological applications. Enzymatic Technologies for Marine Polysaccharides, CRC Press.","DOI":"10.1201\/9780429058653-16"},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"117695","DOI":"10.1016\/j.carbpol.2021.117695","article-title":"Biological strategies for oligo\/polysaccharide synthesis: Biocatalyst and microbial cell factory","volume":"258","author":"Zhao","year":"2021","journal-title":"Carbohydr. Polym."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"6229","DOI":"10.1039\/D4SC01466A","article-title":"Nanopore-based glycan sequencing: State of the art and future prospects","volume":"15","author":"Yao","year":"2024","journal-title":"Chem. Sci."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"cwaf048","DOI":"10.1093\/glycob\/cwaf048","article-title":"Modeling glycans with AlphaFold 3: Capabilities, caveats, and limitations","volume":"35","author":"Huang","year":"2025","journal-title":"Glycobiology"},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1007\/s12272-026-01603-2","article-title":"Mechanisms and structure\u2013activity relationship of hypolipidemic effects of polysaccharides from natural resources: A review","volume":"49","author":"Wu","year":"2026","journal-title":"Arch. Pharmacal Res."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1016\/j.ijbiomac.2015.10.081","article-title":"Marine polysaccharide-based nanomaterials as a novel source of nanobiotechnological applications","volume":"82","author":"Manivasagan","year":"2016","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"148236","DOI":"10.1016\/j.ijbiomac.2025.148236","article-title":"Polysaccharide-modified liposome-based drug delivery system for cancer targeted therapy: An updated review","volume":"330","author":"Zhu","year":"2025","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"4267","DOI":"10.3390\/md11114267","article-title":"Cytotoxic Effects of Fucoidan Nanoparticles against Osteosarcoma","volume":"11","author":"Kimura","year":"2013","journal-title":"Mar. Drugs"},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"1162","DOI":"10.1134\/S1068162024040125","article-title":"Methods for Introdusing Fluorescent Labels into Polysaccharides (A Review)","volume":"50","author":"Nokel","year":"2024","journal-title":"Russ. J. Bioorg. Chem."},{"key":"ref_115","doi-asserted-by":"crossref","unstructured":"Cresswell, T., Metian, M., Fisher, N.S., Charmasson, S., Hansman, R.L., Bam, W., Bock, C., and Swarzenski, P.W. (2020). Exploring New Frontiers in Marine Radioisotope Tracing\u2013Adapting to New Opportunities and Challenges. Front. Mar. Sci., 7.","DOI":"10.3389\/fmars.2020.00406"},{"key":"ref_116","doi-asserted-by":"crossref","unstructured":"Andr\u00e9, R., Pacheco, R., Bourbon, M., and Serralheiro, M.L. (2021). Brown Algae Potential as a Functional Food against Hypercholesterolemia: Review. Foods, 10.","DOI":"10.3390\/foods10020234"},{"key":"ref_117","first-page":"4668","article-title":"A Mechanistic Insight into the Effects of Marine Sources as Functional Foods Against Metabolic Syndrome: A Review Article","volume":"21","author":"Javad","year":"2024","journal-title":"Lett. Drug Des. Discov."}],"container-title":["Marine Drugs"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1660-3397\/24\/9\/315\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T08:08:07Z","timestamp":1788941287000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1660-3397\/24\/9\/315"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,9]]},"references-count":117,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["md24090315"],"URL":"https:\/\/doi.org\/10.3390\/md24090315","relation":{},"ISSN":["1660-3397"],"issn-type":[{"value":"1660-3397","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,9]]}}}