Acknowledgement
This research was supported by the Regional Innovation System & Education (RISE) program through the Jeju RISE Center, funded by the Ministry of Education (MOE) and the Jeju Special-Governing Province, Republic of Korea (2025-RISE-17-001).
References
- Aina, O., Bakare, O. O., Daniel, A. I., et al. 2022. Seaweed-derived phenolic compounds in growth promotion and stress alleviation in plants. Life 12:1548. doi.org/10.3390/life12101548
- Azab, A., Nassar, A. & Azab, A. N. 2016. Anti-inflammatory activity of natural products. Molecules 21:1321. doi.org/10.3390/molecules21101321
- Bao, Y., He, X., Wu, W., et al. 2020. Sulfated galactofucan from Sargassum thunbergii induces senescence in human lung cancer A549 cells. Food Funct. 11:4785–4792. doi.org/10.1039/D0FO00699H
- Bergé, J.-P. & Barnathan, G. 2005. Fatty acids from lipids of marine organisms: molecular biodiversity, roles as biomarkers, biologically active compounds, and economical aspects. Adv. Biochem. Eng. Biotechnol. 96:49–125. doi.org/10.1007/b135782
- Bhatti, H. N. & Khera, R. A. 2012. Biological transformations of steroidal compounds: a review. Steroids 77:1267–1290. doi.org/10.1016/j.steroids.2012.07.018
- Cai, Y.-P., Xie, C.-B., Wang, B.-C., Li, P.-L. & Li, B.-F. 2010. Two new resorcinols from Sargassum thunbergii. J. Asian Nat. Prod. Res. 12:1001–1004. doi.org/10.1080/10286020.2010.522179
- Cao, H. 2014. Adipocytokines in obesity and metabolic disease. J. Endocrinol. 220:T47–T59. doi.org/10.1530/JOE-13-0339
- Carina, D., Sharma, S., Jaiswal, A. K. & Jaiswal, S. 2021. Seaweeds polysaccharides in active food packaging: a review of recent progress. Trends Food Sci. Technol. 110:559–572. doi.org/10.1016/j.tifs.2021.02.022
- Catarino, M. D., Silva-Reis, R., Chouh, A., et al. 2023. Applications of antioxidant secondary metabolites of Sargassum spp. Mar. Drugs 21:172. doi.org/10.3390/md21030172
- Cha, S.-H., Hwang, Y., Heo, S.-J. & Jun, H.-S. 2019. Indole-4-carboxaldehyde isolated from seaweed, Sargassum thunbergii, attenuates methylglyoxal-induced hepatic inflammation. Mar. Drugs 17:486. doi.org/10.3390/md17090486
- Chen, B., Chen, H., Qu, H., et al. 2022. Photoprotective effects of Sargassum thunbergii on ultraviolet B-induced mouse L929 fibroblasts and zebrafish. BMC Complement. Med. Ther. 22:144. doi.org/10.1186/s12906-022-03609-x
- Chen, X., Wang, Z., Duan, N., Zhu, G., Schwarz, E. M. & Xie, C. 2018. Osteoblast–osteoclast interactions. Connect. Tissue Res. 59:99–107. doi.org/10.1080/03008207.2017.1290085
- Chen, Z., Xu, Y., Liu, T., Zhang, L., Liu, H. & Guan, H. 2016. Comparative studies on the characteristic fatty acid profiles of four different Chinese medicinal Sargassum seaweeds by GC-MS and chemometrics. Mar. Drugs 14:68. doi.org/10.3390/md14040068
- Chinese Pharmacopoeia Commission. 2025. Pharmacopoeia of the People's Republic of China (2025 Edition, Part I). China Medical Science and Technology Press, Beijing, 1331pp.
- Chu, S. H., Zhang, Q. S., Liu, S. K., et al. 2012. Tolerance of Sargassum thunbergii germlings to thermal, osmotic and desiccation stress. Aquat. Bot. 96:1–6. doi.org/10.1016/j.aquabot.2011.09.002
- Cotas, J., Leandro, A., Monteiro, P., et al. 2020. Seaweed phenolics: from extraction to applications. Mar. Drugs 18:384. doi.org/10.3390/md18080384
- Cui, L., Qu, H., Qiao, K., et al. 2023. Anti-skin aging potential of Sargassum thunbergii ethanolic extract: antioxidant, anti-inflammatory, and antiwrinkle effects on L929 fibroblast cells. J. Food Process. Preserv. 2023:2230456. doi.org/10.1155/2023/2230456
- Cushnie, T. P. T., Cushnie, B. & Lamb, A. J. 2014. Alkaloids: an overview of their antibacterial, antibiotic-enhancing and antivirulence activities. Int. J. Antimicrob. Agents 44:377–386. doi.org/10.1016/j.ijantimicag.2014.06.001
- Du, F.-Y., Li, X., Li, X.-M., Zhu, L.-W. & Wang, B.-G. 2017. Indolediketopiperazine alkaloids from Eurotium cristatum EN-220, an endophytic fungus isolated from the marine alga Sargassum thunbergii. Mar. Drugs 15:24. doi.org/10.3390/md15020024
- Endalifer, M. L. & Diress, G. 2020. Epidemiology, predisposing factors, biomarkers, and prevention mechanism of obesity: a systematic review. J. Obes. 2020:6134362. doi.org/10.1155/2020/6134362
- Fajas, L., Schoonjans, K., Gelman, L., et al. 1999. Regulation of peroxisome proliferator-activated receptor γ expression by adipocyte differentiation and determination factor 1/sterol regulatory element binding protein 1: implications for adipocyte differentiation and metabolism. Mol. Cell. Biol. 19:5495–5503. doi.org/10.1128/mcb.19.8.5495
- Fink, P. 2007. Ecological functions of volatile organic compounds in aquatic systems. Mar. Freshw. Behav. Physiol. 40:155–168. doi.org/10.1080/10236240701602218
- Gam, D.-H., Park, J.-H., Hong, J.-W., Jeon, S.-J., Kim, J.-H. & Kim, J.-W. 2021. Effects of Sargassum thunbergii extract on skin whitening and anti-wrinkling through inhibition of TRP-1 and MMPs. Molecules 26:7381. doi.org/10.3390/molecules26237381
- Ganesan, A. R., Tiwari, U. & Rajauria, G. 2019. Seaweed nutraceuticals and their therapeutic role in disease prevention. Food Sci. Hum. Wellness 8:252–263. doi.org/10.1016/j.fshw.2019.08.001
- Ghaliaoui, N., Hazzit, M. & Mokrane, H. 2024. Seaweeds as a potential source of bioactive compounds. Res. Biotechnol. Environ. Sci. 3:1–8. doi.org/10.58803/rbes.v3i1.19
- Gressler, V., Colepicolo, P. & Pinto, E. 2009. Useful strategies for algal volatile analysis. Curr. Anal. Chem. 5:271–292. doi.org/10.2174/157341109788680255
- Gutiérrez-Rodríguez, A. G., Juárez-Portilla, C., Olivares-Bañuelos, T. & Zepeda, R. C. 2018. Anticancer activity of seaweeds. Drug Discov. Today 23:434–447. doi.org/10.1016/j.drudis.2017.10.019
- Güven, K. C., Percot, A. & Sezik, E. 2010. Alkaloids in marine algae. Mar. Drugs 8:269–284. doi.org/10.3390/md8020269
- Haeggstrom, J. Z. & Funk, C. D. 2011. Lipoxygenase and leukotriene pathways: biochemistry, biology, and roles in disease. Chem. Rev. 111:5866–5898. doi.org/10.1021/cr200246d
- Han, X. D. & Li, F. P. 2005. Characteristics and utilization of Sargassum thunbergii. Spec. Econ. Anim. Plant 1:27.
- Harder, J. 2010. Isoprene, isoprenoids and sterols. In Timmis, K. N. (Ed.) Handbook of Hydrocarbon and Lipid Microbiology. Springer, Berlin, pp. 127–131.
- He, S., Zhang, Y., Yuan, Y., et al. 2022. Process optimization and antioxidative activity of polyphenols derived from different seaweed species Sargassum miyabei, Undaria pinnatifida Suringar, and Sargassum thunbergii. Food Sci. Nutr. 10:2021–2028. doi.org/10.1002/fsn3.2818
- He, W.-F., Yao, L.-G., Liu, H.-L. & Guo, Y.-W. 2014. Thunberol, a new sterol from the Chinese brown alga Sargassum thunbergii. J. Asian Nat. Prod. Res. 16:685–689. doi.org/10.1080/10286020.2014.924511
- Heo, J.-H., Je, J.-G., Sim, J.-H., Ryu, B., Heo, S.-J. & Jeon, Y.-J. 2024. Quantitative analysis of fucose in fucoidans from Sargassum spp. in Jeju Island, South Korea using 3-methyl-1-phenyl-5-pyrazolone derivatization and RP-HPLCUV method. Algal Res. 79:103441. doi.org/10.1016/j.algal.2024.103441
- Hu, X. M., Zhang, W. K. & Zhu, Q. S. 1998. Zhonghua Bencao. Shanghai Science and Technology Publications, Shanghai, pp. 225–226.
- Ito, K. & Hori, K. 1989. Seaweed: chemical composition and potential food uses. Food Rev. Int. 5:101–144. doi.org/10.1080/87559128909540845
- Javed, A., Alam, M. B., Naznin, M., Shafique, I., Kim, S. & Lee, S.-H. 2025. Tyrosinase inhibitory activity of Sargassum fusiforme and characterisation of bioactive compounds. Phytochem. Anal. 36:343–357. doi.org/10.1002/pca.3233
- Jiang, F. W. & Zhang, Y. S. 1993. China ocean medicine glossary. Ocean Press, Beijing.
- Jiang, Q. 2012. Chemical constituents from marine alga Sargassum thunbergii. Chin. Pharm. J. 24:948–952.
- Jimenez-Lopez, C., Pereira, A. G., Lourenço-Lopes, C., et al. 2021. Main bioactive phenolic compounds in marine algae and their mechanisms of action supporting potential health benefits. Food Chem. 341:128262. doi.org/10.1016/j.foodchem.2020.128262
- Jin, W., Chen, F., Fang, Q., Mao, G. & Bao, Y. 2023. Oligosaccharides from Sargassum thunbergii inhibit osteoclast differentiation via regulation of IRF-8 signaling. Exp. Gerontol. 172:112057. doi.org/10.1016/j.exger.2022.112057
- Jin, W., Liu, B., Li, S., et al. 2018. The structural features of the sulfated heteropolysaccharide (ST-1) from Sargassum thunbergii and its neuroprotective activities. Int. J. Biol. Macromol. 108:307–313. doi.org/10.1016/j.ijbiomac.2017.12.009
- Jin, W., Wu, W., Tang, H., et al. 2019. Structure analysis and anti-tumor and anti-angiogenic activities of sulfated galactofucan extracted from Sargassum thunbergii. Mar. Drugs 17:52. doi.org/10.3390/md17010052
- Jin, W., Zhang, W., Liu, G., et al. 2017. The structure-activity relationship between polysaccharides from Sargassum thunbergii and anti-tumor activity. Int. J. Biol. Macromol. 105:686–692. doi.org/10.1016/j.ijbiomac.2017.07.089
- Kang, J. Y., Khan, M. N. A., Park, N. H., et al. 2008. Antipyretic, analgesic, and anti-inflammatory activities of the seaweed Sargassum fulvellum and Sargassum thunbergii in mice. J. Ethnopharmacol. 116:187–190. doi.org/10.1016/j.jep.2007.10.032
- Kang, M.-C., Ding, Y., Kim, E.-A., et al. 2017. Indole derivatives isolated from brown alga Sargassum thunbergii inhibit adipogenesis through AMPK activation in 3T3-L1 preadipocytes. Mar. Drugs 15:119. doi.org/10.3390/md15040119
- Kang, M.-C., Lee, H., Choi, H.-D. & Jeon, Y.-J. 2019. Antioxidant properties of a sulfated polysaccharide isolated from an enzymatic digest of Sargassum thunbergii. Int. J. Biol. Macromol. 132:142–149. doi.org/10.1016/j.ijbiomac.2019.03.178
- Kang, M.-C., Lee, H.-G., Kim, H.-S., et al. 2020. Anti-obesity effects of Sargassum thunbergii via downregulation of adipogenesis gene and upregulation of thermogenic genes in high-fat diet-induced obese mice. Nutrients 12:3325. doi.org/10.3390/nu12113325
- Kang, N., Heo, S.-Y., Kim, E.-A., Cha, S.-H., Ryu, B. & Heo, S.-J. 2023. Antiviral effect of fucoxanthin obtained from Sargassum siliquastrum (Fucales, Phaeophyceae) against severe acute respiratory syndrome coronavirus 2. Algae 38:295–306. doi.org/10.4490/algae.2023.38.11.29
- Kim, D., Yan, J., Bak, J., Park, J., Lee, H. & Kim, H. 2022. Sargassum thunbergii extract attenuates high-fat diet-induced obesity in mice by modulating AMPK activation and the gut microbiota. Foods 11:2529. doi.org/10.3390/foods11162529
- Kim, J.-A., Karadeniz, F., Ahn, B.-N., et al. 2016. Bioactive quinone derivatives from the marine brown alga Sargassum thunbergii induce anti-adipogenic and pro-osteoblastogenic activities. J. Sci. Food Agric. 96:783–790. doi.org/10.1002/jsfa.7148
- Kim, J.-A., Kong, C.-S. & Kim, S.-K. 2010a. Effect of Sargassum thunbergii on ROS mediated oxidative damage and identification of polyunsaturated fatty acid components. Food Chem. Toxicol. 48:1243–1249. doi.org/10.1016/j.fct.2010.02.017
- Kim, J.-A., Kong, C.-S., Seo, Y.-W. & Kim, S.-K. 2010b. Sargassum thunbergii extract inhibits MMP-2 and -9 expressions related with ROS scavenging in HT1080 cells. Food Chem. 120:418–425. doi.org/10.1016/j.foodchem.2009.10.022
- Kim, J. K., Yarish, C., Hwang, E. K., Park, M. & Kim, Y. 2017. Seaweed aquaculture: cultivation technologies, challenges and its ecosystem services. Algae 32:1–13. doi.org/10.4490/algae.2017.32.3.3
- Kim, J.-M., Lin, C., Stavre, Z., Greenblatt, M. B. & Shim, J.-H. 2020. Osteoblast-osteoclast communication and bone homeostasis. Cells 9:2073. doi.org/10.3390/cells9092073
- Kim, J.-W., Kim, Y.-J. & Eom, A.-H. 2024. Diversity and community structure of endophytic fungi isolated from the brown seaweed Sargassum thunbergii in coastal regions of Korea. Mycobiology 52:317–323. doi.org/10.1080/12298093.2024.2416730
- Kim, K.-B.-W.-R., Kim, M.-J. & Ahn, D.-H. 2014. Lipase inhibitory activity of chlorophyll a, isofucosterol and saringosterol isolated from chloroform fraction of Sargassum thunbergii. Nat. Prod. Res. 28:1310–1312. doi.org/10.1080/14786419.2014.900769
- Kim, S. O. & Choi, Y. H. 2020. Indole-6-carboxaldehyde isolated from Sargassum thunbergii (Mertens) Kuntze prevents oxidative stress-induced cellular damage in V79-4 Chinese hamster lung fibroblasts through the activation of the Nrf2/HO-1 signaling pathway. Cell. Physiol. Biochem. 54:959–974. doi.org/10.33594/000000281
- Kim, T.-H., Heo, S.-J., Ko, S.-C., et al. 2019. Indole-6-carboxaldehyde isolated from Sargassum thunbergii inhibits the expression and secretion of matrix metalloproteinase-9. Int. J. Mol. Med. 44:1979–1987. doi.org/10.3892/ijmm.2019.4319
- Kim, Y. H., Kim, E.-H., Lee, C., Kim, M.-H. & Rho, J.-R. 2007. Two new monogalactosyl diacylglycerols from brown alga Sargassum thunbergii. Lipids 42:395–399. doi.org/10.1007/s11745-007-3035-7
- Kittakoop, P., Mahidol, C. & Ruchirawat, S. 2014. Alkaloids as important scaffolds in therapeutic drugs for the treatments of cancer, tuberculosis, and smoking cessation. Curr. Topics Med. Chem. 14:239–252. doi.org/10.2174/1568026613666131216105049
- Koh, C.-H., Kim, Y. & Kang, S.-G. 1993. Size distribution, growth and production of Sargassum thunbergii in an intertidal zone of Padori, west coast of Korea. Hydrobiologia 260:207–214. doi.org/10.1007/BF00049021
- Kubota, N., Terauchi, Y., Miki, H., et al. 1999. PPARγ mediates high-fat diet-induced adipocyte hypertrophy and insulin resistance. Mol. Cell 4:597–609. doi.org/10.1016/s1097-2765(00)80210-5
- Kuntze, O. 1881. Revision von Sargassum und das sogenannte Sargasso-Meer. Bot. Jahrb. 1:191–239.
- Kurihara, H., Kagawa, Y., Konno, R., Kim, S. M. & Takahashi, K. 2014. Lipoxygenase inhibitors derived from marine macroalgae. Bioorg. Med. Chem. Lett. 24:1383–1385. doi.org/10.1016/j.bmcl.2014.01.046
- Lee, H.-G., Jayawardhana, H. H. A. C. K., Yang, F., et al. 2024a. Anti-obesity effects of fucoidan from Sargassum thunbergii in adipocytes and high fat diet induced obese mice through inhibiting adipogenic specific transcription factor. Food Sci. Hum. Wellness 13:1608–1616. doi.org/10.26599/FSHW.2022.9250136
- Lee, H.-G., Nagahawatta, D. P., Kurera, M. J. M. S., et al. 2024b. Effects of low molecular weight polysaccharide from Sargassum thunbergii against palmitic acid-induced intracellular lipid accumulation in 3T3-L1 adipocyte and HepG2 cells. Food Sci. Hum. Wellness 13:2244–2252. doi.org/10.26599/FSHW.2022.9250187
- Lee, J.-H., Hwang, S.-J. & Kim, H.-R. 2025. Antioxidant profiles of Korean brown algae revealed by multivariate analysis. Algae 40:285–294. doi.org/10.4490/algae.2025.40.8.14
- Lee, M.-K., Ryu, H., Lee, J. Y., et al. 2022. Potential beneficial effects of Sargassum spp. in skin aging. Mar. Drugs 20:540. doi.org/10.3390/md20080540
- Lee, S.-G. & Kang, H. 2015. Neuroprotective effect of Sargassum thunbergii (Mertens ex Roth) Kuntze in activated murine microglial cells. Trop. J. Pharm. Res. 14:235–240. doi.org/10.4314/tjpr.v14i2.7
- Lee, S.-G. & Kang, H. 2025. Regional variation in antioxidant and anti-inflammatory activities of the brown alga Sargassum thunbergii and mechanistic role of fucosterol in inflammation modulation. Biomedicines 13:2808. doi.org/10.3390/biomedicines13112808
- Li, D., Chen, L., Chen, S., Zhang, X., Chen, F. & Ye, N. 2012. Comparative evaluation of the pyrolytic and kinetic characteristics of a macroalga (Sargassum thunbergii) and a freshwater plant (Potamogeton crispus). Fuel 96:185–191. doi.org/10.1016/j.fuel.2012.01.005
- Li, J.-J., Hu, Z.-M., Gao, X., et al. 2017a. Oceanic currents drove population genetic connectivity of the brown alga Sargassum thunbergii in the north-west Pacific. J. Biogeogr. 44:230–242. doi.org/10.1111/jbi.12856
- Li, J.-J., Hu, Z.-M., Sun, Z.-M., et al. 2017b. Historical isolation and contemporary gene flow drive population diversity of the brown alga Sargassum thunbergii along the coast of China. BMC Evol. Biol. 17:246. doi.org/10.1186/s12862-017-1089-6
- Li, S., Hu, M., Tong, Y., et al. 2023. A review of volatile compounds in edible macroalgae. Food Res. Int. 165:112559. doi.org/10.1016/j.foodres.2023.112559
- Li, X., Wang, Y., Jiang, X., et al. 2022. Utilization of different seaweeds (Sargassum polycystum, Sargassum thunbergii, Sargassum horneri, Enteromorpha prolifera, Macrocystis pyrifera, and the residue of M. pyrifera) in the diets of sea cucumber Apostichopus japonicus (Selenka, 1867). Algal Res. 61:102591. doi.org/10.1016/j.algal.2021.102591
- Liu, F., Hu, Z., Liu, W., et al. 2016a. Distribution, function and evolution characterization of microsatellite in Sargassum thunbergii (Fucales, Phaeophyta) transcriptome and their application in marker development. Sci. Rep. 6:18947. doi.org/10.1038/srep18947
- Liu, F.-L., Li, J.-J., Liang, Z.-R., et al. 2021. A concise review of the brown seaweed Sargassum thunbergii: a knowledge base to inform large-scale cultivation efforts. J. Appl. Phycol. 33:3469–3482. doi.org/10.1007/s10811-021-02557-2
- Liu, F., Sun, X., Wang, W., Liang, Z. & Wang, F. 2014. De novo transcriptome analysis-gained insights into physiological and metabolic characteristics of Sargassum thunbergii (Fucales, Phaeophyceae). J. Appl. Phycol. 26:1519–1526. doi.org/10.1007/s10811-013-0140-2
- Liu, J., Willför, S. & Xu, C. 2015. A review of bioactive plant polysaccharides: biological activities, functionalization, and biomedical applications. Bioact. Carbohydr. Diet. Fibre 5:31–61. doi.org/10.1016/j.bcdf.2014.12.001
- Liu, L., Heinrich, M., Myers, S. & Dworjanyn, S. A. 2012. Towards a better understanding of medicinal uses of the brown seaweed Sargassum in traditional Chinese medicine: a phytochemical and pharmacological review. J. Ethnopharmacol. 142:591–619. doi.org/10.1016/j.jep.2012.05.046
- Liu, W., Wu, H., Zhan, D. & Duan, D. 2016b. Phenological study of Sargassum thunbergii (Fucales, Phaeophyta) in Lidao Bay, Rongcheng, China. Chin. J. Oceanol. Limnol. 34:498–506. doi.org/10.1007/s00343-016-5046-9
- Liu, Y. & Tan, H. 2014. Changes of growth and nutrient-relating enzymatic activities of Sargassum thunbergii when exposed to different nutrient conditions. Aquat. Sci. Technol. 2:1–13. doi.org/10.5296/ast.v2i2.5103
- Lohr, M., Schwender, J. & Polle, J. E. W. 2012. Isoprenoid biosynthesis in eukaryotic phototrophs: a spotlight on algae. Plant Sci. 185:9–22. doi.org/10.1016/j.plantsci.2011.07.018
- Lomartire, S., Cotas, J., Pacheco, D., Marques, J. C., Pereira, L. & Gonçalves, A. M. 2021. Environmental impact on seaweed phenolic production and activity: an important step for compound exploitation. Mar. Drugs 19:245. doi.org/10.3390/md19050245
- López-Alarcón, C. & Denicola, A. 2013. Evaluating the antioxidant capacity of natural products: a review on chemical and cellular-based assays. Anal. Chim. Acta 763:1–10. doi.org/10.1016/j.aca.2012.11.051
- Luo, D., Wang, Z. & Nie, K. 2019. Structural characterization of a novel polysaccharide from Sargassum thunbergii and its antioxidant and anti-inflammation effects. PLoS ONE 14:e0223198. doi.org/10.1371/journal.pone.0223198
- Luo, D., Yuan, X., Zeng, Y., Nie, K., Li, Z. & Wang, Z. 2016. Structure elucidation of a major fucopyranose-rich heteropolysaccharide (STP-II) from Sargassum thunbergii. Carbohydr. Polym. 143:1–8. doi.org/10.1016/j.carbpol.2016.01.049
- Mattio, L. & Payri, C. E. 2011. 190 Years of Sargassum taxonomy, facing the advent of DNA phylogenies. Bot. Rev. 77:31–70. doi.org/10.1007/s12229-010-9060-x
- Medzhitov, R. 2008. Origin and physiological roles of inflammation. Nature 454:428–435. doi.org/10.1038/nature07201
- Miyashita K., Mikami N. & Hosokawa M. 2013. Chemical and nutritional characteristics of brown seaweed lipids: a review. J. Funct. Foods 5:1507–1517. doi.org/10.1016/j.jff.2013.09.019
- Mun, O.-J., Kwon, M. S., Karadeniz, F., et al. 2017. Fermentation of Sargassum thunbergii by kimchi-derived Lactobacillus sp. SH-1 attenuates LPS-stimulated inflammatory response via downregulation of JNK. J. Food Biochem. 41:e12306. doi.org/10.1111/jfbc.12306
- Nagahawatta, D. P., Liyanage, N. M., Jayawardena, T. U., et al. 2023. Functions and values of sulfated polysaccharides from seaweed. Algae 38:217–240. doi.org/10.4490/algae.2023.38.12.1
- Nathan, C. & Ding, A. 2010. Nonresolving inflammation. Cell 140:871–882. doi.org/10.1016/j.cell.2010.02.029
- Ou, M., Sun, X., Liang, J., et al. 2017. A polysaccharide from Sargassum thunbergii inhibits angiogenesis via downregulating MMP-2 activity and VEGF/HIF-1α signaling. Int. J. Biol. Macromol. 94:451–458. doi.org/10.1016/j.ijbiomac.2016.10.046
- Park, C., HwangBo, H., Lee, H., et al. 2020a. The immunostimulatory effect of indole-6-carboxaldehyde isolated from Sargassum thunbergii (Mertens) Kuntze in RAW 264.7 macrophages. Anim. Cells Syst. 24:233–241. doi.org/10.1080/19768354.2020.1808529
- Park, C., Lee, H., Park, S.-H., et al. 2020b. Indole-6-carboxaldehyde prevents oxidative stress-induced mitochondrial dysfunction, DNA damage and apoptosis in C2C12 skeletal myoblasts by regulating the ROS-AMPK signaling pathway. Mol. Cell. Toxicol. 16:455–467. doi.org/10.1007/s13273-020-00102-9
- Park, I.-H., Lee, S.-H., Kim, S.-K., Ngo, D.-N., Jeon, Y.-J. & Kim, M.-M. 2011. Effects of seaweeds on matrix metalloproteinases derived from normal human dermal fibroblasts and human fibrosarcoma cells. J. Life Sci. 21:1501–1510. doi.org/10.5352/JLS.2011.21.11.1501
- Park, J.-S., Han, J.-M., Shin, Y.-N., et al. 2023. Exploring bioactive compounds in brown seaweeds using subcritical water: a comprehensive analysis. Mar. Drugs 21:328. doi.org/10.3390/md21060328
- Park, J.-S., Han, J.-M., Surendhiran, D. & Chun, B.-S. 2022. Physicochemical and biofunctional properties of Sargassum thunbergii extracts obtained from subcritical water extraction and conventional solvent extraction. J. Supercrit. Fluids 182:105535. doi.org/10.1016/j.supflu.2022.105535
- Park, K.-E., Kim, Y. A., Jung, H. A., et al. 2004. Three norisoprenoids from the brown alga Sargassum thunbergii. J. Korean Chem. Soc. 48:394–398. doi.org/10.5012/jkcs.2004.48.4.394
- Park, P.-J., Heo, S.-J., Park, E.-J., et al. 2005. Reactive oxygen scavenging effect of enzymatic extracts from Sargassum thunbergii. J. Agric. Food Chem. 53:6666–6672. doi.org/10.1021/jf050582+
- Pereira, L. 2018. Seaweeds as source of bioactive substances and skin care therapy: cosmeceuticals, algotheraphy, and thalassotherapy. Cosmetics 5:68. doi.org/10.3390/cosmetics5040068
- Pérez, M. J., Falqué, E. & Domínguez, H. 2016. Antimicrobial action of compounds from marine seaweed. Mar. Drugs 14:52. doi.org/10.3390/md14030052
- Phillips, N. 1995. Biogeography of Sargassum (Phaeophyta) in the Pacific basin. In Abbott, I. A. (Ed.) Taxonomy of Economic Seaweeds. California Sea Grant College Program, La Jolla, CA, pp. 107–144.
- Pichersky, E., Noel, J. P. & Dudareva, N. 2006. Biosynthesis of plant volatiles: nature's diversity and ingenuity. Science 311:808–811. doi.org/10.1126/science.1118510
- Ping, T., Qingling, X., Jungang, Y. & Xu, G. 2001. An analysis of nutrient components of thirteen kinds of seaweeds for food in Dalian coastline. J. Liaoning Normal Univ. (Nat. Sci.) 24:406–410.
- Promyo, K., Cho, A.-R., Song, D., et al. 2025. Effects of brown algae (Sargassum thunbergii and Sargassum fusiforme) on the progression of some characteristics of Alzheimer's disease and microglial dysfunction in D-galactose-induced aging rat. Appl. Biol. Chem. 68:82. doi.org/10.1186/s13765-025-01054-1
- Qiao, H. J., Hu, D. X., Hu, W. J., et al. 2019. Effects of dietary Sargassum thunbergii powder on growth performance, body composition, antioxidation and non-specific immune parameters of juvenile turbot. J. Shanghai Ocean Univ. 28:109–116.
- Qiu, T., Sun, Y., Qu, T., et al. 2023. Study on gender differences between male and female Sargassum thunbergii based on metabolomic analysis and physiological functions. Algal Res. 75:103267. doi.org/10.1016/j.algal.2023.103267
- Rasheed, A. & Qasim, M. 2013. A review of natural steroids and their applications. Int. J. Pharm. Sci. Res. 4:520–531. doi.org/10.13040/IJPSR.0975-8232.4(2).520-31
- Ren, B., Chen, C., Li, C., Fu, X., You, L. & Liu, R. H. 2017. Optimization of microwave-assisted extraction of Sargassum thunbergii polysaccharides and its antioxidant and hypoglycemic activities. Carbohydr. Polym. 173:192–201. doi.org/10.1016/j.carbpol.2017.05.094
- Richard, D., Kefi, K., Barbe, U., Bausero, P. & Visioli, F. 2008. Polyunsaturated fatty acids as antioxidants. Pharmacol. Res. 57:451–455. doi.org/10.1016/j.phrs.2008.05.002
- Rosen, E. D., Hsu, C.-H., Wang, X., et al. 2002. C/EBPα induces adipogenesis through PPARγ: a unified pathway. Genes Dev. 16:22–26. doi.org/10.1101/gad.948702
- Rushdi, M. I., Abdel-Rahman, I. A. M., Saber, H., et al. 2020. Pharmacological and natural products diversity of the brown algae genus Sargassum. RSC Adv. 10:24951–24972. doi.org/10.1039/d0ra03576a
- Ryter, S. W., Kim, H. P., Hoetzel, A., et al. 2007. Mechanisms of cell death in oxidative stress. Antioxid. Redox Signal. 9:49–89. doi.org/10.1089/ars.2007.9.49
- Sánchez-Machado, D. I., López-Cervantes, J., López-Hernández, J. & Paseiro-Losada, P. 2004a. Fatty acids, total lipid, protein and ash contents of processed edible seaweeds. Food Chem. 85:439–444. doi.org/10.1016/j.foodchem.2003.08.001
- Sánchez-Machado, D. I., López-Hernández, J., Paseiro-Losada, P. & López-Cervantes, J. 2004b. An HPLC method for the quantification of sterols in edible seaweeds. Biomed. Chromatogr. 18:183–190. doi.org/10.1002/bmc.316
- Saraswati, Giriwono, P. E., Iskandriati, D., Tan, C. P. & Andarwulan, N. 2019. Sargassum seaweed as a source of anti-inflammatory substances and the potential insight of the tropical species: a review. Mar. Drugs 17:590. doi.org/10.3390/md17100590
- Senthilkumar, K., Manivasagan, P., Venkatesan, J. & Kim, S.-K. 2013. Brown seaweed fucoidan: biological activity and apoptosis, growth signaling mechanism in cancer. Int. J. Biol. Macromol. 60:366–374. doi.org/10.1016/j.ijbiomac.2013.06.030
- Seo, J.-Y., Shin, I.-S. & Lee, S.-M. 2011. Effect of dietary inclusion of various plant ingredients as an alternative for Sargassum thunbergii on growth and body composition of juvenile sea cucumber Apostichopus japonicus. Aquac. Nutr. 17:549–556. doi.org/10.1111/j.1365-2095.2010.00849.x
- Seo, Y., Park, K. E. & Nam, T. J. 2007. Isolation of a new chromene from the brown alga Sargassum thunbergii. Bull. Korean Chem. Soc. 28:1831–1833. doi.org/10.5012/bkcs.2007.28.10.1831
- Seo, Y., Lee, H.-J., Park, K. E., et al. 2004. Peroxynitrite-scavenging constituents from the brown alga Sargassum thunbergii. Biotechnol. Bioprocess Eng. 9:212–216. doi.org/10.1007/BF02942295
- Seo, Y., Park, K. E., Kim, Y. A., et al. 2006. Isolation of tetraprenyltoluquinols from the brown alga Sargassum thunbergii. Chem. Pharm. Bull. 54:1730–1733. doi.org/10.1248/cpb.54.1730
- Serhan, C. N. & Levy, B. D. 2018. Resolvins in inflammation: emergence of the pro-resolving superfamily of mediators. J. Clin. Invest. 128:2657–2669. doi.org/10.1172/JCI97943
- Shalaby, E. A. 2011. Algae as promising organisms for environment and health. Plant Signal. Behav. 6:1338–1350. doi.org/10.4161/psb.6.9.16779
- Shibata, Y. & Morita, M. 1988. A novel, trimethylated arseno-sugar isolated from the brown alga Sargassum thunbergii. Agric. Biol. Chem. 52:1087–1089. doi.org/10.1080/00021369.1988.10868781
- Sies, H. 1985. Oxidative stress: introductory remarks. In Sies, H. (Ed.) Oxidative Stress. Academic Press, London, pp. 1–8.
- Silberfeld, T., Leigh, J. W., Verbruggen, H., Cruaud, C., de Reviers, B. & Rousseau, F. 2010. A multi-locus time-calibrated phylogeny of the brown algae (Heterokonta, Ochrophyta, Phaeophyceae): investigating the evolutionary nature of the "brown algal crown radiation". Mol. Phylogenet. Evol. 56:659–674. doi.org/10.1016/j.ympev.2010.04.020
- Singh, S., Kumar, S., Singh, A. K., Varshney, M. & Roy, S. 2025. Exploring marine alkaloids: a natural approach to cancer treatment. Curr. Pharm. Biotechnol. 26:63–79. doi.org/10.2174/0113892010316791240611093022
- Sireesha, B., Reddy, B. V., Basha, S., Chandra, K., Anasuya, D. & Bhavani, M. 2019. A review on pharmacological activities of alkaloids. World J. Curr. Med. Pharm. Res. 1:230–234. doi.org/10.37022/WJCMPR.2019.01068
- Sohn, S.-I., Rathinapriya, P., Balaji, S., et al. 2021. Phytosterols in seaweeds: an overview on biosynthesis to biomedical applications. Int. J. Mol. Sci. 22:12691. doi.org/10.3390/ijms222312691
- Son, B.-W., Cho, Y.-J., Kim, N.-K. & Choi, H.-D. 1992. New glyceroglycolipids from the brown alga Sargassum thunbergii. Bull. Korean Chem. Soc. 13:584–586. doi.org/10.5012/bkcs.1992.13.6.584
- Song, X.-H., Assis, J., Zhang, J., et al. 2021. Climate-induced range shifts shaped the present and threaten the future genetic variability of a marine brown alga in the Northwest Pacific. Evol. Appl. 14:1867–1879. doi.org/10.1111/eva.13247
- Song, X., Zhang, J., Du, Y., et al. 2025. Historical climate change influenced the phylogeographical patterns of the brown alga Colpomenia sinuosa in the southern China. Algae 40:29–43. doi.org/10.4490/algae.2025.40.2.27
- Souza, B. W. S., Cerqueira, M. A., Bourbon, A. I., et al. 2012. Chemical characterization and antioxidant activity of sulfated polysaccharide from the red seaweed Gracilaria birdiae. Food Hydrocoll. 27:287–292. doi.org/10.1016/j.foodhyd.2011.10.005
- Sugimoto, M. A., Sousa, L. P., Pinho, V., Perretti, M. & Teixeira, M. M. 2016. Resolution of inflammation: what controls its onset? Front. Immunol. 7:160. doi.org/10.3389/fimmu.2016.00160
- Su, Y.-C. & Liu, C. 2007. Vibrio parahaemolyticus: a concern of seafood safety. Food Microbiol. 24:549–558. doi.org/10.1016/j.fm.2007.01.005
- Sun, Y., Liu, Q., Shang, S., et al. 2022. Physiological responses and metabonomics analysis of male and female Sargassum thunbergii macroalgae exposed to ultraviolet-B stress. Front. Plant Sci. 13:778602. doi.org/10.3389/fpls.2022.778602
- Tanna, B. & Mishra, A. 2019. Nutraceutical potential of seaweed polysaccharides: structure, bioactivity, safety, and toxicity. Compr. Rev. Food Sci. Food Saf. 18:817–831. doi.org/10.1111/1541-4337.12441
- Thakuri, L. S., Park, C. M., Park, J. W., Kim, H.-A. & Rhyu, D. Y. 2023. Subcritical water extraction of Gracilaria chorda abbreviates lipid accumulation and obesity-induced inflammation. Algae 38:81–92. doi.org/10.4490/algae.2023.38.12.26
- Tsukamoto, S., Hirota, H., Kato, H. & Fusetani, N. 1994. Phlorotannins and sulfoquinovosyl diacylglycerols: promoters of larval metamorphosis in ascidians, isolated from the brown alga Sargassum thunbergii. Fish. Sci. 60:319–321.
- Van Gaal, L. F., Mertens, I. L. & De Block, C. E. 2006. Mechanisms linking obesity with cardiovascular disease. Nature 444:875–880. doi.org/10.1038/nature05487
- Van Pham, P., Dang, L. T.-T., Dinh, U. T., et al. 2014. In vitro evaluation of the effects of human umbilical cord extracts on human fibroblasts, keratinocytes, and melanocytes. In Vitro Cell. Dev. Biol. Anim. 50:321–330. doi.org/10.1007/s11626-013-9706-1
- Vogt, T. 2010. Phenylpropanoid biosynthesis. Mol. Plant 3:2–20. doi.org/10.1093/mp/ssp106
- Wallace, G. & Fry, S. C. 1994. Phenolic components of the plant cell wall. Int. Rev. Cytol. 151:229–267. doi.org/10.1016/S0074-7696(08)62634-0
- Wang, H., Zhang, M., Yang, W., Zhuang, L. & Guo, L. 2025. Extraction, enrichment, characterization, and antioxidant activities of Sargassum fusiforme polyphenols. Foods 14:3317. doi.org/10.3390/foods14193317
- Wang, J., Yang, Z., Wang, G., Shang, S., Tang, X. & Xiao, H. 2022. Diversity of epiphytic bacterial communities on male and female Sargassum thunbergii. AMB Express 12:97. doi.org/10.1186/s13568-022-01439-1
- Wang, P., Chen, J., Chen, L., Shi, L. & Liu, H. 2021. Characteristic volatile composition of seven seaweeds from the Yellow Sea of China. Mar. Drugs 19:192. doi.org/10.3390/md19040192
- Wang, R., Wang, Y. & Tang, X. 2012. Identification of the toxic compounds produced by Sargassum thunbergii to red tide microalgae. Chin. J. Oceanol. Limnol. 30:778–785. doi.org/10.1007/s00343-012-1294-5
- Wang, Z. & Liu, J. 2007. The sexual reproduction of Sargassum thunbergii and its application in seedling breeding. Oceanol. Limnol. Sin. 38:457.
- Wei, Y., Liu, Q., Xu, C., Yu, J., Zhao, L. & Guo, Q. 2016. Damage to the membrane permeability and cell death of Vibrio parahaemolyticus caused by phlorotannins with low molecular weight from Sargassum thunbergii. J. Aquat. Food Prod. Technol. 25:323–333. doi.org/10.1080/10498850.2013.851757
- Wen, K.-C., Chang, C.-S., Chien, Y.-C., et al. 2013. Tyrosol and its analogues inhibit alpha-melanocyte-stimulating hormone induced melanogenesis. Int. J. Mol. Sci. 14:23420–23440. doi.org/10.3390/ijms141223420
- Wu, H., Zhan, D., Liu, H., Ding, G., Liu, W. & Li, M. 2010. Study on accumulation and degradation of heavy metals by the brown alga Sargassum thunbergii. Mar. Sci. 34:69–74.
- Xia, Z., Chen, B. J. W., Korpelainen, H., Niinemets, Ü. & Li, C. 2024. Belowground ecological interactions in dioecious plants: why do opposites attract but similar ones repel? Trends Plant Sci. 29:630–637. doi.org/10.1016/j.tplants.2024.02.009
- Xie, J.-H., Jin, M.-L., Morris, G. A., et al. 2016. Advances on bioactive polysaccharides from medicinal plants. Crit. Rev. Food Sci. Nutr. 56(Suppl 1):S60–S84. doi.org/10.1080/10408398.2015.1069255
- Xu, L.-C., Luo, W.-H., Liu, S., Ye, J.-X. & Chen, Y. 2025. Fusasolpolyol A, an unreported polyhydroxy compound isolated from the Sargassum thunbergii-derived endophytic fungus Fusarium solani 2024f-xx. Rec. Nat. Prod. 19:541–546. doi.org/10.25135/rnp.520.2504.3506
- Yamada, T., Kogure, H., Kataoka, M., Kikuchi, T. & Hirano, T. 2020. Halosmysin A, a novel 14-membered macrodiolide isolated from the marine-algae-derived fungus Halosphaeriaceae sp. Mar. Drugs 18:320. doi.org/10.3390/md18060320
- Yamada, T., Yoshida, K., Kikuchi, T. & Hirano, T. 2022. Isolation and structure elucidation of new cytotoxic macrolides halosmysins B and C from the fungus Halosphaeriaceae sp. associated with a marine alga. Mar. Drugs 20:226. doi.org/10.3390/md20040226
- Yang, E.-J., Moon, J.-Y., Kim, M.-J., et al. 2010. Inhibitory effect of Jeju endemic seaweeds on the production of pro-inflammatory mediators in mouse macrophage cell line RAW 264.7. J. Zhejiang Univ. Sci. B 11:315–322. doi.org/10.1631/jzus.B0900364
- Yang, F., Nagahawatta, D. P., Yang, H.-W., et al. 2023. In vitro and in vivo immuno-enhancing effect of fucoidan isolated from non-edible brown seaweed Sargassum thunbergii. Int. J. Biol. Macromol. 253:127212. doi.org/10.1016/j.ijbiomac.2023.127212
- Yang, J. H., Graf, L., Cho, C. H., Jeon, B. H., Kim, J. H. & Yoon, H. S. 2016. Complete plastid genome of an ecologically important brown alga Sargassum thunbergii (Fucales, Phaeophyceae). Mar. Genomics 28:17–20. doi.org/10.1016/j.margen.2016.03.003
- Yang, Y., Zhang, M., Alalawy, A. I., et al. 2021. Identification and characterization of marine seaweeds for biocompounds production. Environ. Technol. Innov. 24:101848. doi.org/10.1016/j.eti.2021.101848
- Yende, S. R., Harle, U. N. & Chaugule, B. B. 2014. Therapeutic potential and health benefits of Sargassum species. Pharmacognosy Rev. 8:1–7. doi.org/10.4103/0973-7847.125514
- Yoshida, T. 1983. Japanese species of Sargassum subgenus Bactrophycus (Phaeophyta, Fucales). J. Fac. Sci. Hokkaido Univ. Series 5 Bot. 13:99–246.
- Yu, Y., Shen, M., Song, Q. & Xie, J. 2018. Biological activities and pharmaceutical applications of polysaccharide from natural resources: a review. Carbohydr. Polym. 183:91–101. doi.org/10.1016/j.carbpol.2017.12.009
- Yu, Y. Q., Zhang, Q. S., Tang, Y. Z., et al. 2012. Establishment of intertidal seaweed beds of Sargassum thunbergii through habitat creation and germling seeding. Ecol. Eng. 44:10–17. doi.org/10.1016/j.ecoleng.2012.03.016
- Yuan, X., Zeng, Y., Nie, K., Luo, D. & Wang, Z. 2015. Extraction optimization, characterization and bioactivities of a major polysaccharide from Sargassum thunbergii. PLoS ONE 10:e0144773. doi.org/10.1371/journal.pone.0144773
- Zhang, Z., Sun, Y., Li, Y., Song, X., Wang, R. & Zhang, D. 2024. The potential of marine-derived piperazine alkaloids: sources, structures and bioactivities. Eur. J. Med. Chem. 265:116081. doi.org/10.1016/j.ejmech.2023.116081
- Zhao, L. & Funk, C. D. 2004. Lipoxygenase pathways in atherogenesis. Trends Cardiovasc. Med. 14:191–195. doi.org/10.1016/j.tcm.2004.04.003
- Zhao, Y., Sun, T., Li, Y., et al. 2024. The host sex contributes to the endophytic bacterial community in Sargassum thunbergii and their receptacles. Front. Microbiol. 15:1334918. doi.org/10.3389/fmicb.2024.1334918
- Zhou, Q.-L., Wang, Z., Chen, W.-T., et al. 2024. The structural characteristics, biological activities and mechanisms of bioactive brown seaweed polysaccharides: a review. J. Funct. Foods 119:106303. doi.org/10.1016/j.jff.2024.106303
- Zhu, K., Wang, X., Weng, Y., et al. 2024. Sulfated galactofucan from Sargassum thunbergii attenuates atherosclerosis by suppressing inflammation via the TLR4/MyD88/NF-κB signaling pathway. Cardiovasc. Drugs Ther. 38:69–78. doi.org/10.1007/s10557-022-07383-3
- Zhuang, C., Itoh, H., Mizuno, T. & Ito, H. 1995. Antitumor active fucoidan from the brown seaweed, umitoranoo (Sargassum thunbergii). Biosci. Biotechnol. Biochem. 59:563–567. doi.org/10.1271/bbb.59.563