{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T21:39:59Z","timestamp":1791841199423,"version":"4.3.4"},"reference-count":76,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2026,4,28]],"date-time":"2026-04-28T00:00:00Z","timestamp":1777334400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100005046","name":"Heilongjiang Provincial Natural Science Foundation of China","doi-asserted-by":"crossref","award":["YQ2023H001"],"award-info":[{"award-number":["YQ2023H001"]}],"id":[{"id":"10.13039\/501100005046","id-type":"DOI","asserted-by":"crossref"}]},{"name":"the North Medicine and Functional Food Characteristic Subject Project in Heilongjiang Province","award":["HLJTSXK-2022-03"],"award-info":[{"award-number":["HLJTSXK-2022-03"]}]},{"name":"Longjiang Technology Talent Spring Goose Support Program","award":["CYQN24035"],"award-info":[{"award-number":["CYQN24035"]}]},{"name":"the Basic Research Project of the Fundamental Research Business Expenses of the Education Department in Heilongjiang Province","award":["2022-KYYWF-0641"],"award-info":[{"award-number":["2022-KYYWF-0641"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Molecules"],"abstract":"<jats:p>The immunomodulatory potential of natural polysaccharides is often limited by their structural complexity and high molecular weight. In this study, DFIP-A3-1 (Mw = 8.68 \u00d7 103 g\/mol) was obtained from the leaves of Isatis indigotica Fort. by ultrafiltration, DEAE-650M, and Sephadex G-100 chromatography, followed by acid degradation. Fortunately, DFIP-A3-1 exhibited the most potent immunostimulatory activity in vitro. HPGPC, HPSEC-MALLS-RID, GC-MS, FT-IR, Congo red tests, SEM, and AFM were used to characterize their structure, and 1D\/2D NMR was used for further investigation of DFIP-A3-1 for in-depth structural clarification. DFIP-A3-1 was primarily composed of Rha and Gal. Based on methylation and NMR analyses, the structure of DFIP-A3-1 was elucidated as follows: \u21921)-\u03b2-Galp-(4\u21921,4)-\u03b1-Rhap-(2\u21921,4)-\u03b1-Rhap-(2\u21921)-\u03b2-Galp-(6\u21921)-\u03b2-Galp-(6\u21921,6)-\u03b2-Galp-(3\u21921,6)-\u03b2-Galp-(3\u2192. Furthermore, DFIP-A3-1 was found to exhibit a triple-helix conformation. DFIP-A3-1 markedly upregulated the secretion of NO, IL-6, and TNF-\u03b1 and enhanced the mRNA expression levels of their related genes in RAW 264.7 cells. Moreover, DFIP-A3-1 activated p-I\u03baB\u03b1, p-p65, and TLR4, while co-treatment with TAK-242 markedly suppressed the expression of these pathway-related proteins. All of the aforementioned findings suggested that DFIP-A3-1 is a promising natural immunomodulatory drug deserving of additional research and use.<\/jats:p>","DOI":"10.3390\/molecules31091461","type":"journal-article","created":{"date-parts":[[2026,4,28]],"date-time":"2026-04-28T13:26:41Z","timestamp":1777382801000},"page":"1461","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Acid Degradation, Structure Characterization of a Novel Polysaccharide from Leaves of Isatis indigotica Fort. with Immunomodulatory Activity"],"prefix":"10.3390","volume":"31","author":[{"ORCID":"https:\/\/orcid.org\/0009-0007-2530-8473","authenticated-orcid":false,"given":"Yu","family":"Shen","sequence":"first","affiliation":[{"name":"College of Pharmacy, Jiamusi University, Jiamusi 154007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-9935-5358","authenticated-orcid":false,"given":"Xuefeng","family":"Wang","sequence":"additional","affiliation":[{"name":"College of Pharmacy, Jiamusi University, Jiamusi 154007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-5594-1405","authenticated-orcid":false,"given":"Huiming","family":"Zhang","sequence":"additional","affiliation":[{"name":"College of Basic Medical Sciences, Jiamusi University, Jiamusi 154007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5383-2079","authenticated-orcid":false,"given":"Yuliang","family":"Wang","sequence":"additional","affiliation":[{"name":"College of Pharmacy, Jiamusi University, Jiamusi 154007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-0061-2754","authenticated-orcid":false,"given":"Zheng","family":"Wang","sequence":"additional","affiliation":[{"name":"College of Pharmacy, Jiamusi University, Jiamusi 154007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4280-4811","authenticated-orcid":false,"given":"Yiying","family":"Zhang","sequence":"additional","affiliation":[{"name":"Department of Epidemiology and Biostatistics, School of Public Health, Jiamusi University, Jiamusi 154007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-2154-7981","authenticated-orcid":false,"given":"Hongbo","family":"Zhao","sequence":"additional","affiliation":[{"name":"College of Rehabilitation Medicine, Jiamusi University, Jiamusi 154007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,4,28]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"2448","DOI":"10.1111\/febs.13730","article-title":"Initiation of Innate Immune Responses by Surveillance of Homeostasis Perturbations","volume":"283","author":"Moita","year":"2016","journal-title":"FEBS J."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1007\/s12016-024-09020-3","article-title":"Clinical and Pathophysiological Tangles between Allergy and Autoimmunity: Deconstructing an Old Dichotomic Paradigm","volume":"68","author":"Ramirez","year":"2025","journal-title":"Clin. Rev. Allergy Immunol."},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Tang, J., and Zhao, X. (2024). Research Progress on Regulation of Immune Response by Tanshinones and Salvianolic Acids of Danshen (Salvia miltiorrhiza Bunge). Molecules, 29.","DOI":"10.3390\/molecules29061201"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"13440","DOI":"10.1038\/s41598-025-97423-3","article-title":"Immunostimulatory Effects Mechanism of Polysaccharide Extracted from Acanthopanax senticosus on RAW 264.7 Cells through Activating the TLR\/MAPK\/NF-\u03baB Signaling Pathway","volume":"15","author":"Wang","year":"2025","journal-title":"Sci. Rep."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"907","DOI":"10.1016\/j.ijbiomac.2021.07.024","article-title":"Molecular Mechanisms Underlying Macrophage Immunomodulatory Activity of Rubus Chingii Hu Polysaccharides","volume":"185","author":"Xu","year":"2021","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_6","first-page":"2686992","article-title":"Polygonatum Polysaccharide Regulates Macrophage Polarization and Improves LPS-Induced Acute Lung Injury through TLR4-MAPK\/NF-\u03baB Pathway","volume":"2022","author":"Zhou","year":"2022","journal-title":"Can. Respir. J."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Sotto, A.D., Vitalone, A., and Giacomo, S.D. (2020). Plant-Derived Nutraceuticals and Immune System Modulation: An Evidence-Based Overview. Vaccines, 8.","DOI":"10.3390\/vaccines8030468"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"114319","DOI":"10.1016\/j.indcrop.2021.114319","article-title":"Isolation, Structural Modification, Characterization, and Bioactivity of Polysaccharides from Folium Isatidis","volume":"176","author":"Abuduwaili","year":"2022","journal-title":"Ind. Crops Prod."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"114648","DOI":"10.1016\/j.jep.2021.114648","article-title":"Isatidis Radix and Isatidis Folium: A Systematic Review on Ethnopharmacology, Phytochemistry and Pharmacology","volume":"283","author":"Chen","year":"2022","journal-title":"J. Ethnopharmacol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1038\/s41438-020-0240-5","article-title":"A Chromosome-Scale Genome Assembly of Isatis Indigotica, an Important Medicinal Plant Used in Traditional Chinese Medicine: An Isatis Genome","volume":"7","author":"Kang","year":"2020","journal-title":"Hortic. Res."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Min, G.-Y., Kim, T.I., Kim, J.-H., Cho, W.-K., Yang, J.-H., and Ma, J.Y. (2023). Anti-Atopic Effect of Isatidis Folium Water Extract in TNF-\u03b1\/IFN-\u03b3-Induced HaCaT Cells and DNCB-Induced Atopic Dermatitis Mouse Model. Molecules, 28.","DOI":"10.3390\/molecules28093960"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1142\/S0192415X22500069","article-title":"A Systemic Review for Ethnopharmacological Studies on Isatis Indigotica Fortune: Bioactive Compounds and Their Therapeutic Insights","volume":"50","author":"Wong","year":"2022","journal-title":"Am. J. Chin. Med."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"121552","DOI":"10.1016\/j.indcrop.2025.121552","article-title":"Preparation, Structure-Activity Relationship, Biological Activities and Market Applications of Isatis Indigotica Polysaccharides: A Review","volume":"234","author":"Huang","year":"2025","journal-title":"Ind. Crops Prod."},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Yin, Y., Shi, X., Cai, X., Liu, F., Ni, W., Li, B., Wan, X., and Ren, M. (2024). Isolation Techniques, Structural Characteristics, and Pharmacological Effects of Phellinus Polysaccharides: A Review. Molecules, 29.","DOI":"10.3390\/molecules29133047"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"120965","DOI":"10.1016\/j.carbpol.2023.120965","article-title":"The Antioxidant Activity of Polysaccharides: A Structure-Function Relationship Overview","volume":"314","author":"Fernandes","year":"2023","journal-title":"Carbohydr. Polym."},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Liu, T., Ren, Q., Wang, S., Gao, J., Shen, C., Zhang, S., Wang, Y., and Guan, F. (2023). Chemical Modification of Polysaccharides: A Review of Synthetic Approaches, Biological Activity and the Structure\u2013Activity Relationship. Molecules, 28.","DOI":"10.3390\/molecules28166073"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"2359","DOI":"10.1007\/s10924-021-02052-2","article-title":"Polysaccharides; Classification, Chemical Properties, and Future Perspective Applications in Fields of Pharmacology and Biological Medicine (a Review of Current Applications and Upcoming Potentialities)","volume":"29","author":"Mohammed","year":"2021","journal-title":"J. Polym. Environ."},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Chatnarin, S., and Thirabunyanon, M. (2023). Potential Bioactivities via Anticancer, Antioxidant, and Immunomodulatory Properties of Cultured Mycelial Enriched \u03b2-D-Glucan Polysaccharides from a Novel Fungus Ophiocordyceps Sinensis OS8. Front. Immunol., 14.","DOI":"10.3389\/fimmu.2023.1150287"},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Casas-Arrojo, V., Arrojo Agudo, M.d.L.\u00c1., C\u00e1rdenas Garc\u00eda, C., Carrillo, P., P\u00e9rez Manr\u00edquez, C., Mart\u00ednez-Manzanares, E., and Abdala D\u00edaz, R.T. (2022). Antioxidant, Immunomodulatory and Potential Anticancer Capacity of Polysaccharides (Glucans) from Euglena gracilis G.A. Klebs. Pharmaceuticals, 15.","DOI":"10.3390\/ph15111379"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"125204","DOI":"10.1016\/j.ijbiomac.2023.125204","article-title":"Structure Characterization and Biological Activities Evaluation of Two Hetero-Polysaccharides from Lepista Nuda: Cell Antioxidant, Anticancer and Immune-Modulatory Activities","volume":"244","author":"Li","year":"2023","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"110913","DOI":"10.1016\/j.foodres.2021.110913","article-title":"Characterization, Antioxidant, Antineoplastic and Immune Activities of Selenium Modified Sagittaria sagittifolia L. Polysaccharides","volume":"153","author":"Feng","year":"2022","journal-title":"Food Res. Int."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"2276814","DOI":"10.1080\/19490976.2023.2276814","article-title":"Modulating the Gut Microbiota Is Involved in the Effect of Low-Molecular-Weight Glycyrrhiza Polysaccharide on Immune Function","volume":"15","author":"Song","year":"2023","journal-title":"Gut Microbes"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"136097","DOI":"10.1016\/j.ijbiomac.2024.136097","article-title":"Low Molecular Weight Polysaccharide of Tremella Fuciformis Exhibits Stronger Antioxidant and Immunomodulatory Activities than High Molecular Weight Polysaccharide","volume":"281","author":"Lee","year":"2024","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Lin, Y., Yang, J., Luo, L., Zhang, X., Deng, S., Chen, X., Li, Y., Bekhit, A.E.-D.A., Xu, B., and Huang, R. (2022). Ferroptosis Related Immunomodulatory Effect of a Novel Extracellular Polysaccharides from Marine Fungus Aureobasidium melanogenum. Mar. Drugs, 20.","DOI":"10.3390\/md20050332"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1016\/0003-2697(92)90349-C","article-title":"Carbohydrate Composition Analysis of Bacterial Polysaccharides: Optimized Acid Hydrolysis Conditions for HPAEC-PAD Analysis","volume":"201","author":"Ip","year":"1992","journal-title":"Anal. Biochem."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"109591","DOI":"10.1016\/j.carres.2025.109591","article-title":"Structural Basis of Immunomodulation by Edible Fungal Polysaccharides: From Molecular Characteristics to Action Mechanisms","volume":"555","author":"Gao","year":"2025","journal-title":"Carbohydr. Res."},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Pan, M., Zhang, X., Huang, S., Liu, M., Song, S., Ma, J., Li, C., Ma, W., and Liu, X. (2025). Preparation, Structural Characteristics and Immune Regulatory Effects of Codonopsis Pilosula Polysaccharides: A Review. Front. Immunol., 16.","DOI":"10.3389\/fimmu.2025.1641928"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"134117","DOI":"10.1016\/j.ijbiomac.2024.134117","article-title":"Efficient Degradation and Enhanced Anticomplementary Activity of Belamcanda chinensis (L.) DC. Polysaccharides via Trifluoroacetic Acid Treatment with Different Degrees","volume":"276","author":"Duan","year":"2024","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"052014","DOI":"10.1088\/1757-899X\/392\/5\/052014","article-title":"Studies on the Acid Degradation Process and in Vitro Immune Activity of the Polysaccharide H6PC20 in Hericium Erinaceus","volume":"392","author":"Li","year":"2018","journal-title":"IOP Conf. Ser. Mater. Sci. Eng."},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Gieroba, B., Kalisz, G., Krysa, M., Khalavka, M., and Przekora, A. (2023). Application of Vibrational Spectroscopic Techniques in the Study of the Natural Polysaccharides and Their Cross-Linking Process. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24032630"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"129244","DOI":"10.1016\/j.ijbiomac.2024.129244","article-title":"Polysaccharide from Thymelaea hirsuta L. Leaves: Structural Characterization, Functional Properties and Antioxidant Evaluation","volume":"262","author":"Ghamgui","year":"2024","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Chen, X., Yang, T., Huang, Q., Li, B., Ding, X., and Hou, Y. (2023). Comparative Studies on the Structure and Biological Activities of Two New Polysaccharides from Tricholoma sinoportentosum (TS-P) and Termitomyces albuminosus (TA-P). Polymers, 15.","DOI":"10.3390\/polym15092227"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"129591","DOI":"10.1016\/j.ijbiomac.2024.129591","article-title":"Effect of Subcritical Water Temperature on the Structure, Antioxidant Activity and Immune Activity of Polysaccharides from Glycyrrhiza Inflata Batalin","volume":"261","author":"Cheng","year":"2024","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Wu, Q., Liang, B., Wang, J., and Dai, Y. (2025). Ultrasound-Assisted Extraction of Polysaccharides from Lyophyllum decastes: Structural Analysis and Bioactivity Assessment. Molecules, 30.","DOI":"10.3390\/molecules30040961"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"527","DOI":"10.3390\/analytica4040036","article-title":"Physicochemical and Microstructural Characteristics of Sulfated Polysaccharide from Marine Microalga","volume":"4","year":"2023","journal-title":"Analytica"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"642","DOI":"10.1016\/j.ijbiomac.2022.02.187","article-title":"Comparative Study of Water-Soluble Polysaccharides Isolated from Leaves and Roots of Isatis Indigotica Fort","volume":"206","author":"Pak","year":"2022","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"e14362","DOI":"10.1111\/jfbc.14362","article-title":"Tumor Inhibitory Effect via Immunostimulating Activities of a Rhamnogalacturonan-I-Rich Polysaccharide Isolated from Turmeric (Curcuma longa L.)","volume":"46","author":"Choi","year":"2022","journal-title":"J. Food Biochem."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"280","DOI":"10.1016\/j.phytochem.2009.10.001","article-title":"Different Contributions of Side-Chains in Beta-D-(1\u21923,6)-Galactans on Intestinal Peyer\u2019s Patch-Immunomodulation by Polysaccharides from Astragalus Mongholics Bunge","volume":"71","author":"Kiyohara","year":"2010","journal-title":"Phytochemistry"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"123695","DOI":"10.1016\/j.carbpol.2025.123695","article-title":"Novel Galactose-Rich Polysaccharide from Ganoderma Lucidum: Structural Characterization and Immunomodulatory Activities","volume":"362","author":"Wu","year":"2025","journal-title":"Carbohydr. Polym."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"3307","DOI":"10.1016\/0031-9422(92)83678-R","article-title":"NMR Spectroscopy in the Structural Elucidation of Oligosaccharides and Glycosides","volume":"31","author":"Agrawal","year":"1992","journal-title":"Phytochemistry"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"156","DOI":"10.1016\/j.carres.2013.06.026","article-title":"Complete (1)H and (13)C NMR Chemical Shift Assignments of Mono- to Tetrasaccharides as Basis for NMR Chemical Shift Predictions of Oligosaccharides Using the Computer Program CASPER","volume":"380","author":"Pendrill","year":"2013","journal-title":"Carbohydr. Res."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1190","DOI":"10.1016\/j.carres.2010.04.004","article-title":"Isolation of Phosphorylated Polysaccharides from Algae: The Immunostimulatory Principle of Chlorella pyrenoidosa","volume":"345","author":"Kralovec","year":"2010","journal-title":"Carbohydr. Res."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"497","DOI":"10.1016\/j.ijbiomac.2020.06.275","article-title":"Purification, Structural Characterization, and Cognitive Improvement Activity of a Polysaccharides from Schisandra chinensis","volume":"163","author":"Xu","year":"2020","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"184","DOI":"10.1016\/j.ijbiomac.2013.04.052","article-title":"Structural Features, Antioxidant and Immunological Activity of a New Polysaccharide (SP1) from Sisal Residue","volume":"59","author":"Zhang","year":"2013","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"117326","DOI":"10.1016\/j.carbpol.2020.117326","article-title":"Ultrafiltration Isolation, Structures and Anti-Tumor Potentials of Two Arabinose- and Galactose-Rich Pectins from Leaves of Aralia elata","volume":"255","author":"Shen","year":"2021","journal-title":"Carbohydr. Polym."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"867","DOI":"10.1007\/s10541-005-0196-y","article-title":"Structural Studies on Pectin from Marsh Cinquefoil Comarum palustre L","volume":"70","author":"Ovodova","year":"2005","journal-title":"Biochem. Biokhimiia"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1016\/j.carbpol.2016.06.110","article-title":"Structure Analysis of a Heteropolysaccharide from Taraxacum mongolicum Hand.-Mazz. and Anticomplementary Activity of Its Sulfated Derivatives","volume":"152","author":"Chen","year":"2016","journal-title":"Carbohydr. Polym."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"923","DOI":"10.1016\/j.ijbiomac.2022.04.045","article-title":"Clarification of the Structural Features of Rhamnogalacturonan-I Type Polysaccharide Purified from Radish Leaves","volume":"209","author":"Son","year":"2022","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"117962","DOI":"10.1016\/j.carbpol.2021.117962","article-title":"Triple-Helix Polysaccharides: Formation Mechanisms and Analytical Methods","volume":"262","author":"Guo","year":"2021","journal-title":"Carbohydr. Polym."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"10627","DOI":"10.1021\/acs.jafc.5c01560","article-title":"Exploring the Structure-Activity Relationship of F. Velutipes Polysaccharide: A Novel Promising Anti-Inflammatory and Intestinal Barrier-Protective Molecule Extracted by Ultrasound","volume":"73","author":"Chen","year":"2025","journal-title":"J. Agric. Food Chem."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"e14023","DOI":"10.1111\/jfbc.14023","article-title":"Structural Characterization and Immunoregulatory Activity of Polysaccharides from Dendrobium officinale Leaves","volume":"46","author":"Zhong","year":"2022","journal-title":"J. Food Biochem."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"740","DOI":"10.1016\/j.ijbiomac.2020.03.235","article-title":"Purification, Structural Elucidation and Physicochemical Properties of a Polysaccharide from Abelmoschus esculentus L (Okra) Flowers","volume":"155","author":"Zhang","year":"2020","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Liu, J., Zhang, J., Feng, J., Tang, C., Yan, M., Zhou, S., Chen, W., Wang, W., and Liu, Y. (2023). Multiple Fingerprint-Activity Relationship Assessment of Immunomodulatory Polysaccharides from Ganoderma Lucidum Based on Chemometric Methods. Molecules, 28.","DOI":"10.3390\/molecules28072913"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"141973","DOI":"10.1016\/j.ijbiomac.2025.141973","article-title":"Structural Determination, Immunomodulatory Activity, and Antitumor Activity of a Low-Molecular-Weight Polysaccharide Extracted from Lepista sordida","volume":"307","author":"Li","year":"2025","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"e70013","DOI":"10.1111\/1750-3841.70013","article-title":"Structural Characterization of Two Low-Molecular-Weight Polysaccharides from the Roots of Aucklandia lappa Decne. and Their Bioactivity on the Gut Microbiota of Immunocompromised Mice","volume":"90","author":"Peng","year":"2025","journal-title":"J. Food Sci."},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Pang, G., Zhou, K., Gao, D., Wang, D., Xue, J., Lu, T., and Chai, H. (2026). Extraction, Purification, Structural Elucidation, and Immunomodulatory Activity of Bletilla striata Polysaccharides via the NF-\u03baB Signaling Pathway. Front. Immunol., 17.","DOI":"10.3389\/fimmu.2026.1749545"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"107072","DOI":"10.1016\/j.jff.2025.107072","article-title":"Research Progress on the Structure-Activity Relationship and Action Mechanisms of Polysaccharides with Immunomodulatory Activity","volume":"134","author":"Huang","year":"2025","journal-title":"J. Funct. Foods"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"133500","DOI":"10.1016\/j.ijbiomac.2024.133500","article-title":"Impact of Molecular Weight and Gastrointestinal Digestion on the Immunomodulatory Effects of Lycium barbarum Polysaccharides","volume":"274","author":"Guo","year":"2024","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"JCS263513","DOI":"10.1242\/jcs.263513","article-title":"Biochemical and Biophysical Mechanisms Macrophages Use to Tune Phagocytic Appetite","volume":"138","author":"Bond","year":"2025","journal-title":"J. Cell Sci."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"e70256","DOI":"10.1002\/mco2.70256","article-title":"Macrophage Signaling Pathways in Health and Disease: From Bench to Bedside Applications","volume":"6","author":"Chi","year":"2025","journal-title":"MedComm"},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Fu, Y.L., and Harrison, R.E. (2021). Microbial Phagocytic Receptors and Their Potential Involvement in Cytokine Induction in Macrophages. Front. Immunol., 12.","DOI":"10.3389\/fimmu.2021.662063"},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Arango Duque, G., and Descoteaux, A. (2014). Macrophage Cytokines: Involvement in Immunity and Infectious Diseases. Front. Immunol., 5.","DOI":"10.3389\/fimmu.2014.00491"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1159\/000550397","article-title":"Macrophages in Ulcerative Colitis: Immunomodulatory Roles, Phenotypic Switching, and Therapeutic Targeting","volume":"18","author":"Zhao","year":"2026","journal-title":"J. Innate Immun."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1016\/S0022-2836(02)01404-3","article-title":"TAK1 Is Critical for IkappaB Kinase-Mediated Activation of the NF-kappaB Pathway","volume":"326","author":"Takaesu","year":"2003","journal-title":"J. Mol. Biol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"218087","DOI":"10.1016\/j.canlet.2025.218087","article-title":"Celebrating the 40-Year Milestone: NF-\u0138B in Oncoimmunity","volume":"636","author":"Alipourgivi","year":"2026","journal-title":"Cancer Lett."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"122644","DOI":"10.1016\/j.carbpol.2024.122644","article-title":"Structural Identification, Rheological Properties and Immunological Receptor of a Complex Galacturonoglucan from Fruits of Schisandra chinensis (Turcz.) Baill","volume":"346","author":"Liang","year":"2024","journal-title":"Carbohydr. Polym."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"108709","DOI":"10.1016\/j.biopha.2019.108709","article-title":"Immunomodulatory Effects Exerted by Poria Cocos Polysaccharides via TLR4\/TRAF6\/NF-\u03baB Signaling in Vitro and in Vivo","volume":"112","author":"Tian","year":"2019","journal-title":"Biomed. Pharmacother."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"135808","DOI":"10.1016\/j.ijbiomac.2024.135808","article-title":"Structural Identification and Immunostimulatory Effect of Bacillus velezensis GV1 Polysaccharides via TLR4\/NF-\u03baB Signaling Pathway in RAW264.7 Macrophages","volume":"280","author":"Le","year":"2024","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Shen, Y., Wu, S., Song, M., Zhang, H., Zhao, H., Wu, L., Zhao, H., Qiu, H., and Zhang, Y. (2024). The Isolation, Structural Characterization and Anti-Inflammatory Potentials of Neutral Polysaccharides from the Roots of Isatis indigotica Fort. Molecules, 29.","DOI":"10.3390\/molecules29112683"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"135116","DOI":"10.1016\/j.ijbiomac.2024.135116","article-title":"Extraction, Purification, Chemical Characterization, and in Vitro Hypoglycemic Activity of Polysaccharides Derived from Rosa Laevigata Michx","volume":"279","author":"Chen","year":"2024","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"142853","DOI":"10.1016\/j.ijbiomac.2025.142853","article-title":"Arabinan and Galactooligosaccharide-Rich Pectin from the Stem Bark of Aralia Elata (Miq.) Seem.: Structural Elucidation and Immunomodulatory Activities","volume":"309","author":"Jiang","year":"2025","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1016\/j.carbpol.2018.06.066","article-title":"Gas Chromatography-Mass Spectrometry-Based Trimethylsilyl-Alditol Derivatives for Quantitation and Fingerprint Analysis of Anemarrhena asphodeloides Bunge Polysaccharides","volume":"198","author":"Xia","year":"2018","journal-title":"Carbohydr. Polym."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1016\/j.ijbiomac.2021.12.109","article-title":"Structure and Immunological Activity of an Arabinan-Rich Acidic Polysaccharide from Atractylodes lancea (Thunb.) DC","volume":"199","author":"Shen","year":"2022","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Zhao, M., Pang, S., Gao, Y., Li, T., and Jiang, H. (2025). Effect of Chitooligosaccharides on TLR2\/NF-\u03baB Signaling in LPS-Stimulated RAW 264.7 Macrophages. Molecules, 30.","DOI":"10.3390\/molecules30102226"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"1170","DOI":"10.1093\/jpp\/rgac027","article-title":"Protective Effects of Radix Isatidis Polysaccharide Ameliorates Obesity via Promotion AMPK Pathway in High-Fat-Diet-Induced Obese Rats and 3T3-L1 Adipocyte Cells","volume":"74","author":"Yuan","year":"2022","journal-title":"J. Pharm. Pharmacol."},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Lin, P., Yue, W., Xiang, H., Liu, J., and Chen, X. (2025). Investigating the Effects of Gardenia Polysaccharides on LPS-Induced Immune Injury in Mice and Exploring the Molecular Mechanisms Underlying Its Regulatory Effect on the Immune Function of Macrophages. Foods, 14.","DOI":"10.3390\/foods14203455"}],"container-title":["Molecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1420-3049\/31\/9\/1461\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,4,30]],"date-time":"2026-04-30T04:39:08Z","timestamp":1777523948000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1420-3049\/31\/9\/1461"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,4,28]]},"references-count":76,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,5]]}},"alternative-id":["molecules31091461"],"URL":"https:\/\/doi.org\/10.3390\/molecules31091461","relation":{},"ISSN":["1420-3049"],"issn-type":[{"value":"1420-3049","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,4,28]]}}}