{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,5]],"date-time":"2026-11-05T00:37:08Z","timestamp":1793839028124,"version":"4.6.1"},"reference-count":135,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2025,7,22]],"date-time":"2025-07-22T00:00:00Z","timestamp":1753142400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Immunol."],"abstract":"<jats:p>Tonifying polysaccharides, as a class of natural medicines, have attracted considerable interest due to their low toxicity, high safety profile, and excellent stability. Investigations have highlighted the significant impact of polysaccharides derived from Chinese medicinal herbs on the modulation of immune responses, significantly enhancing the functionality of immune organs such as the spleen, thymus, bone marrow, and intestines, stimulating immune cell proliferation and activation of T and B lymphocytes, macrophages, natural killer cells, and dendritic cells, and regulating the secretion and release of immune factors, thereby enhancing overall immune function. Tonifying polysaccharides, such as those found in medicinal herbs like ginseng and Astragalus, have demonstrated significant therapeutic effects in treating immune-mediated diseases, including anti-tumor, anti-autoimmune, and anti-viral activities. Despite the extensive literature published on the pharmacological effects, mechanisms, and therapeutic applications of Tonifying polysaccharides, there remains a lack of systematic organization and summarization. This review summarizes recent research findings on Tonifying polysaccharides within the field of immunomodulation. The research explores the immunomodulatory mechanisms and therapeutic applications of polysaccharides, clarifying their roles in enhancing immune function and their potential in clinical treatments, and aims to establish a robust theoretical framework and scientific foundation for the investigation and practical application of Tonifying polysaccharides, drawing on the extensive research on their structural complexity, biological activities, and clinical applications, thereby promoting their development and application as immune-enhancing agents in traditional Chinese medicine and as vaccine adjuvants<\/jats:p>","DOI":"10.3389\/fimmu.2025.1640679","type":"journal-article","created":{"date-parts":[[2025,7,22]],"date-time":"2025-07-22T05:29:53Z","timestamp":1753162193000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":23,"title":["Immunomodulatory actions of tonifying polysaccharides: pharmacological effects, mechanisms and therapeutic applications"],"prefix":"10.3389","volume":"16","author":[{"given":"Jin-Yu","family":"Li","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chang","family":"Yi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Meng-Qin","family":"Zhu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yan-Feng","family":"Yuan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Guang","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ning-Ning","family":"Qiu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lei","family":"Shen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Li-Ya","family":"Song","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wen-Long","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xi-Li","family":"Zhang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2025,7,22]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1186\/s13223-024-00932-5","article-title":"Introduction to immunology and immune disorders","volume":"20","author":"Marshall","year":"2024","journal-title":"Allergy Asthma Clin Immunol"},{"key":"B2","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/978-3-030-38366-4_13-1","article-title":"Environmental Impact on Immune System","volume-title":"Environmental Endocrinology and Endocrine Disruptors","author":"Isidori","year":"2022"},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.1155\/2022\/1612348","article-title":"Protective effect of natural antioxidants on reducing cisplatin-induced nephrotoxicity","volume":"2022","author":"Zhou","year":"2022","journal-title":"Dis Markers"},{"key":"B4","doi-asserted-by":"publisher","DOI":"10.1016\/bs.pmbts.2019.03.003","article-title":"The structures and biological functions of polysaccharides from traditional Chinese herbs","volume":"163","author":"Zeng","year":"2019","journal-title":"Prog Mol Biol Transl Sci"},{"key":"B5","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2025.139845","article-title":"Application prospect of polysaccharide in the development of vaccine adjuvants","volume":"297","author":"Feng","year":"2025","journal-title":"Int J Biol Macromol"},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2024.1406213","article-title":"Unraveling the web of defense: the crucial role of polysaccharides in immunity","volume":"15","author":"Shen","year":"2024","journal-title":"Front Immunol"},{"key":"B7","doi-asserted-by":"publisher","DOI":"10.3389\/fmolb.2021.655570","article-title":"The adjuvant effects on vaccine and the immunomodulatory mechanisms of polysaccharides from traditional Chinese medicine","volume":"8","author":"Wang","year":"2021","journal-title":"Front Mol Biosci"},{"key":"B8","doi-asserted-by":"publisher","DOI":"10.1016\/j.fochx.2022.100321","article-title":"Immunoregulatory activity of a low-molecular-weight heteropolysaccharide from Ganoderma leucocontextum fruiting bodies in vitro and in vivo","volume":"14","author":"Gao","year":"2022","journal-title":"Food Chemistry: X"},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2024.135993","article-title":"Structural characterization of polysaccharides from Panax ginseng C. A. Meyer root and their triggered potential immunoregulatory and radioprotective activities","volume":"280","author":"Feng","year":"2024","journal-title":"Int J Biol Macromolecules"},{"key":"B10","doi-asserted-by":"publisher","DOI":"10.1007\/s12272-022-01393-3","article-title":"Astragalus polysaccharide: a review of its immunomodulatory effect","volume":"45","author":"Li","year":"2022","journal-title":"Arch Pharm Res"},{"key":"B11","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2019.09.189","article-title":"Characterization and anti-tumor bioactivity of astragalus polysaccharides by immunomodulation","volume":"145","author":"Li","year":"2020","journal-title":"Int J Biol Macromol"},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.1016\/j.biopha.2022.113015","article-title":"Extraction, structure, and activity of polysaccharide from Radix astragali","volume":"150","author":"Tang","year":"2022","journal-title":"BioMed Pharmacother"},{"key":"B13","doi-asserted-by":"publisher","DOI":"10.1016\/j.carres.2024.109230","article-title":"A review of Lycium barbarum polysaccharides: Extraction, purification, structural-property relationships, and bioactive molecular mechanisms","volume":"544","author":"Wang","year":"2024","journal-title":"Carbohydr Res"},{"key":"B14","doi-asserted-by":"publisher","DOI":"10.3390\/molecules23051170","article-title":"A Review: The Bioactivities and Pharmacological Applications of Polygonatum sibiricum polysaccharides","volume":"23","author":"Cui","year":"2018","journal-title":"Molecules"},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.1016\/j.carbpol.2021.118231","article-title":"Ganoderma lucidum polysaccharide modulates gut microbiota and immune cell function to inhibit inflammation and tumorigenesis in colon","volume":"267","author":"Guo","year":"2021","journal-title":"Carbohydr Polymers"},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.3390\/nu13082725","article-title":"Health-promoting of polysaccharides extracted from ganoderma lucidum","volume":"13","author":"Seweryn","year":"2021","journal-title":"Nutrients"},{"key":"B17","doi-asserted-by":"publisher","DOI":"10.1039\/c8fo02438c","article-title":"Lentinan modulates intestinal microbiota and enhances barrier integrity in a piglet model challenged with lipopolysaccharide","volume":"10","author":"Wang","year":"2019","journal-title":"Food Funct"},{"key":"B18","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2024.130621","article-title":"The effect of lentinan on dexamethasone-induced immunosuppression in mice","volume":"264","author":"Jiao","year":"2024","journal-title":"Int J Biol Macromol"},{"key":"B19","doi-asserted-by":"publisher","DOI":"10.3389\/fphar.2024.1468237","article-title":"Comparison of the composition, immunological activity and anti-fatigue effects of different parts in sika deer antler","volume":"15","author":"Chen","year":"2024","journal-title":"Front Pharmacol"},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.3390\/v15051217","article-title":"Viral co-infections and antiviral immunity in honey bees","volume":"15","author":"Durand","year":"2023","journal-title":"Viruses"},{"key":"B21","doi-asserted-by":"publisher","DOI":"10.2215\/CJN.10031014","article-title":"A brief journey through the immune system","volume":"10","author":"Yatim","year":"2015","journal-title":"Clin J Am Soc Nephrol"},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.1126\/sciimmunol.aau6085","article-title":"Structure and function of the immune system in the spleen","volume":"4","author":"Lewis","year":"2019","journal-title":"Sci Immunol"},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2022.926516","article-title":"Key factors for thymic function and development","volume":"13","author":"Shichkin","year":"2022","journal-title":"Front Immunol"},{"key":"B24","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2013.09.038","article-title":"Protective effect of chitooligosaccharides against cyclophosphamide-induced immunosuppression in mice","volume":"62","author":"Mei","year":"2013","journal-title":"Int J Biol Macromolecules"},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2020.05.245","article-title":"Purification, structural characterization and in vivo immunoregulatory activity of a novel polysaccharide from Polygonatum sibiricum","volume":"160","author":"Wang","year":"2020","journal-title":"Int J Biol Macromolecules"},{"key":"B26","doi-asserted-by":"publisher","DOI":"10.1016\/j.indcrop.2021.114309","article-title":"Extraction, purification, and biological activities in vivo of a novel fructose-rich polysaccharide from Codonopsis pilosula","volume":"176","author":"Yu","year":"2022","journal-title":"Ind Crops Products"},{"key":"B27","doi-asserted-by":"publisher","DOI":"10.3390\/nu17020225","article-title":"Glycyrrhiza uralensis polysaccharide modulates characteristic bacteria and metabolites, improving the immune function of healthy mice","volume":"17","author":"Song","year":"2025","journal-title":"Nutrients"},{"key":"B28","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2019.11.208","article-title":"Extraction, structural characterization, and immunoregulatory effect of a polysaccharide fraction from Radix Aconiti Lateralis Preparata (Fuzi)","volume":"143","author":"Yang","year":"2020","journal-title":"Int J Biol Macromolecules"},{"key":"B29","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2023.123386","article-title":"Structure of a Pueraria root polysaccharide and its immunoregulatory activity on T and B lymphocytes, macrophages, and immunosuppressive mice","volume":"230","author":"Cai","year":"2023","journal-title":"Int J Biol Macromolecules"},{"key":"B30","doi-asserted-by":"publisher","first-page":"48","DOI":"10.1161\/ATVBAHA.120.314215","article-title":"Trained immunity and cardiometabolic disease: the role of bone marrow","volume":"41","author":"Mitroulis","year":"2021","journal-title":"ATVB"},{"key":"B31","doi-asserted-by":"publisher","DOI":"10.1016\/j.biopha.2021.111338","article-title":"Polysaccharide-rich extract from Polygonatum sibiricum protects hematopoiesis in bone marrow suppressed by triple negative breast cancer","volume":"137","author":"Xie","year":"2021","journal-title":"Biomed Pharmacother"},{"key":"B32","doi-asserted-by":"publisher","DOI":"10.1080\/15384101.2020.1838793","article-title":"Astragalus polysaccharide inhibits radiation-induced bystander effects by regulating apoptosis in Bone Mesenchymal Stem Cells (BMSCs)","volume":"19","author":"Zhang","year":"2020","journal-title":"Cell Cycle"},{"key":"B33","doi-asserted-by":"publisher","DOI":"10.3389\/fendo.2022.965258","article-title":"Immunoporosis: Role of immune system in the pathophysiology of different types of osteoporosis","volume":"13","author":"Zhang","year":"2022","journal-title":"Front Endocrinol (Lausanne)"},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.1002\/jcb.26682","article-title":"Achyranthes bidentata polysaccharide suppresses osteoclastogenesis and bone resorption via inhibiting RANKL signaling","volume":"119","author":"Song","year":"2018","journal-title":"J Cell Biochem"},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.1302\/2046-3758.128.BJR-2022-0248.R2","article-title":"Astragalus polysaccharide promotes osteogenic differentiation of human bone marrow derived mesenchymal stem cells by facilitating ANKFY1 expression through miR-760 inhibition","volume":"12","author":"Hu","year":"2023","journal-title":"Bone Joint Res"},{"key":"B36","doi-asserted-by":"publisher","DOI":"10.1016\/j.phymed.2022.154322","article-title":"Coptis chinensis polysaccharides dynamically influence the paracellular absorption pathway in the small intestine by modulating the intestinal mucosal immunity microenvironment","volume":"104","author":"Yang","year":"2022","journal-title":"Phytomedicine"},{"key":"B37","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1186\/s40779-017-0122-9","article-title":"Interaction between the gut microbiome and mucosal immune system","volume":"4","author":"Shi","year":"2017","journal-title":"Military Med Res"},{"key":"B38","doi-asserted-by":"publisher","DOI":"10.1016\/j.carbpol.2021.117668","article-title":"Value added immunoregulatory polysaccharides of Hericium erinaceus and their effect on the gut microbiota","volume":"262","author":"Yang","year":"2021","journal-title":"Carbohydr Polymers"},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2024.135063","article-title":"Structural characteristics and intestinal flora metabolism mediated immunoregulatory effects of Lactarius deliciosus polysaccharide","volume":"278","author":"Dong","year":"2024","journal-title":"Int J Biol Macromolecules"},{"key":"B40","doi-asserted-by":"publisher","first-page":"129174","DOI":"10.1016\/j.ijbiomac.2023.129174","article-title":"In vitro and in vivo immunomodulatory activity of acetylated polysaccharides from Cyclocarya paliurus leaves","volume":"259","author":"Zhang","year":"2024","journal-title":"Int J Biol Macromolecules"},{"key":"B41","doi-asserted-by":"publisher","DOI":"10.1016\/j.carbpol.2024.122275","article-title":"A graminan type fructan from\u00a0Achyranthes bidentata prevents the kidney injury in diabetic mice by regulating gut microbiota","volume":"339","author":"Si","year":"2024","journal-title":"Carbohydr Polymers"},{"key":"B42","doi-asserted-by":"publisher","DOI":"10.1016\/j.coi.2018.01.002","article-title":"The regulation of lymphocyte activation and proliferation","volume":"51","author":"Heinzel","year":"2018","journal-title":"Curr Opin Immunol"},{"key":"B43","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1146\/annurev-immunol-061020-053702","article-title":"Cytokine regulation and function in T cells","volume":"39","author":"Dong","year":"2021","journal-title":"Annu Rev Immunol"},{"key":"B44","doi-asserted-by":"publisher","first-page":"795","DOI":"10.1016\/j.fshw.2022.03.010","article-title":"GPP (composition of Ganoderma lucidum polysaccharides and Polyporus umbellatus polysaccharides) protects against DSS-induced murine colitis by enhancing immune function and regulating intestinal flora","volume":"11","author":"Li","year":"2022","journal-title":"Food Sci Hum Wellness"},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2019.11.051","article-title":"Extraction, isolation, immunoregulatory activity, and characterization of Alpiniae oxyphyllae fructus polysaccharides","volume":"155","author":"Yang","year":"2020","journal-title":"Int J Biol Macromol"},{"key":"B46","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2024.130736","article-title":"Achyranthes bidentata polysaccharides improve cyclophosphamide-induced adverse reactions by regulating the balance of cytokines in helper T cells","volume":"265","author":"Lei","year":"2024","journal-title":"Int J Biol Macromol"},{"key":"B47","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2024.03.037","article-title":"Principles and therapeutic applications of adaptive immunity","volume":"187","author":"Chi","year":"2024","journal-title":"Cell"},{"key":"B48","doi-asserted-by":"publisher","DOI":"10.26599\/FSHW.2022.9250036","article-title":"Structure and immunomodulatory activity of Lentinus edodes polysaccharides modified by probiotic fermentation","volume":"13","author":"Liang","year":"2024","journal-title":"Food Sci Hum Wellness"},{"key":"B49","doi-asserted-by":"publisher","DOI":"10.1016\/j.indcrop.2021.114085","article-title":"The extraction, structure, and immunomodulation activities in vivo of polysaccharides from Salvia miltiorrhiza","volume":"173","author":"Ji","year":"2021","journal-title":"Ind\u00a0Crops Products"},{"key":"B50","doi-asserted-by":"publisher","DOI":"10.1016\/j.jep.2023.116351","article-title":"Effects of different steaming times on the composition, structure and immune activity of Polygonatum Polysaccharide","volume":"310","author":"Su","year":"2023","journal-title":"J Ethnopharmacol"},{"key":"B51","doi-asserted-by":"publisher","DOI":"10.3390\/ijms23010144","article-title":"Macrophage polarity and disease control","volume":"23","author":"Kadomoto","year":"2021","journal-title":"IJMS"},{"key":"B52","doi-asserted-by":"publisher","DOI":"10.1016\/j.carbpol.2020.116874","article-title":"Structural characterization and immunomodulatory activity of a water-soluble polysaccharide from Ganoderma leucocontextum fruiting bodies","volume":"249","author":"Gao","year":"2020","journal-title":"Carbohydr Polymers"},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.3390\/ph16060895","article-title":"Isolation, purification, and structural characterization of polysaccharides from codonopsis pilosula and their anti-tumor bioactivity by immunomodulation","volume":"16","author":"Li","year":"2023","journal-title":"Pharmaceuticals"},{"key":"B54","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1155\/2022\/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":"B55","doi-asserted-by":"publisher","first-page":"e1458","DOI":"10.1002\/wsbm.1458","article-title":"Neutrophils in innate immunity and systems biology-level approaches","volume":"12","author":"Rungelrath","year":"2020","journal-title":"WIREs Mech Dis"},{"key":"B56","doi-asserted-by":"publisher","DOI":"10.3390\/nu14050927","article-title":"Ultrasonic-assisted extraction of codonopsis pilosula glucofructan: optimization, structure, and immunoregulatory activity","volume":"14","author":"Ji","year":"2022","journal-title":"Nutrients"},{"key":"B57","doi-asserted-by":"publisher","first-page":"90","DOI":"10.1016\/j.ijbiomac.2021.04.077","article-title":"Structural characteristics and in vitro and in vivo immunoregulatory properties of a gluco-arabinan from Angelica dahurica","volume":"183","author":"Wang","year":"2021","journal-title":"Int J Biol Macromol"},{"key":"B58","doi-asserted-by":"publisher","DOI":"10.1016\/j.carbpol.2025.123502","article-title":"Structural characterization and immunomodulatory activity analysis of a novel pectic polysaccharide extracted from Tetrastigma hemsleyanum Diels et Gilg and its hydrolysis products","volume":"357","author":"Tian","year":"2025","journal-title":"Carbohydr Polymers"},{"key":"B59","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2018.01869","article-title":"Natural killer cells: development, maturation, and clinical utilization","volume":"9","author":"Abel","year":"2018","journal-title":"Front Immunol"},{"key":"B60","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2024.130320","article-title":"Angelica gigas polysaccharide induces CR3-mediated macrophage activation and the cytotoxicity of natural killer cells against HCT-116 cells via NF-\u03baB and MAPK signaling pathways","volume":"263","author":"Ge","year":"2024","journal-title":"Int J Biol Macromolecules"},{"key":"B61","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2023.127605","article-title":"Cnidium officinale polysaccharide enhanced RAW 264.7 cells activation and NK-92 cells cytotoxicity against colon cancer via NF-\u03baB and MAPKs signaling pathways","volume":"253","author":"Yang","year":"2023","journal-title":"Int J Biol Macromol"},{"key":"B62","doi-asserted-by":"publisher","DOI":"10.1016\/j.biopha.2022.113541","article-title":"Application of dendritic cells in tumor immunotherapy and progress in the mechanism of anti-tumor effect of Astragalus polysaccharide (APS) modulating dendritic cells: a review","volume":"155","author":"Wang","year":"2022","journal-title":"Biomed Pharmacother"},{"key":"B63","doi-asserted-by":"publisher","DOI":"10.1016\/j.intimp.2021.107513","article-title":"Immunoregulatory properties of a crude extraction fraction rich in polysaccharide from Chrysanthemum zawadskii Herbich var. latilobum and its potential role as a vaccine adjuvant","volume":"95","author":"Han","year":"2021","journal-title":"Int Immunopharmacol"},{"key":"B64","doi-asserted-by":"publisher","first-page":"301","DOI":"10.1186\/s12906-020-03103-2","article-title":"Isatis root polysaccharide promotes maturation and secretory function of monocyte-derived dendritic cells","volume":"20","author":"Wang","year":"2020","journal-title":"BMC Complement Med Ther"},{"key":"B65","doi-asserted-by":"publisher","DOI":"10.1021\/acs.molpharmaceut.4c00295","article-title":"Surface-engineered polygonatum sibiricum polysaccharide caCO3 microparticles as novel vaccine adjuvants to enhance immune response","volume":"21","author":"He","year":"2024","journal-title":"Mol Pharm"},{"key":"B66","doi-asserted-by":"publisher","DOI":"10.1016\/j.carbpol.2018.10.057","article-title":"Mechanism of Lycium barbarum polysaccharides liposomes on activating murine dendritic cells","volume":"205","author":"Bo","year":"2019","journal-title":"Carbohydr Polym"},{"key":"B67","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2024.132628","article-title":"Structural characterization and immune modulation activities of Chinese Angelica polysaccharide (CAP) and selenizing CAP (sCAP) on dendritic cells","volume":"277","author":"Qiao","year":"2024","journal-title":"Int J Biol Macromolecules"},{"key":"B68","doi-asserted-by":"publisher","DOI":"10.1016\/j.it.2022.07.004","article-title":"NF-\u03baB: blending metabolism, immunity, and inflammation","volume":"43","author":"Capece","year":"2022","journal-title":"Trends Immunol"},{"key":"B69","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2016.10.112","article-title":"Anti-tumor and immunomodulatory activities induced by an alkali-extracted polysaccharide BCAP-1 from Bupleurum chinense via NF-\u03baB signaling pathway","volume":"95","author":"Song","year":"2017","journal-title":"Int J Biol Macromol"},{"key":"B70","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2017.01.039","article-title":"Selenizing Hericium erinaceus polysaccharides induces dendritic cells maturation through MAPK and NF-\u03baB signaling pathways","volume":"97","author":"Qin","year":"2017","journal-title":"Int J Biol Macromol"},{"key":"B71","doi-asserted-by":"publisher","first-page":"391","DOI":"10.1016\/j.intimp.2018.03.002","article-title":"Toll-like receptors in immunity and inflammatory diseases: Past, present, and future","volume":"59","author":"Vijay","year":"2018","journal-title":"Int Immunopharmacol"},{"key":"B72","doi-asserted-by":"publisher","DOI":"10.1016\/j.biopha.2023.114471","article-title":"Immunoregulatory effects of Tetrastigma hemsleyanum polysaccharide via TLR4-mediated NF-\u03baB and MAPK signaling pathways in Raw264.7 macrophages","volume":"161","author":"Wu","year":"2023","journal-title":"Biomed Pharmacother"},{"key":"B73","doi-asserted-by":"publisher","DOI":"10.1016\/j.carbpol.2024.122896","article-title":"Structural characterization of two novel heteropolysaccharides from Catharanthus roseus and the evaluation of their immunological activities","volume":"348","author":"Zhang","year":"2025","journal-title":"Carbohydr Polymers"},{"key":"B74","doi-asserted-by":"publisher","DOI":"10.1111\/cns.14807","article-title":"Advances of the MAPK pathway in the treatment of spinal cord injury","volume":"30","author":"Huang","year":"2024","journal-title":"CNS Neurosci Ther"},{"key":"B75","doi-asserted-by":"publisher","first-page":"77","DOI":"10.24272\/j.issn.2095-8137.2018.066","article-title":"Involvement of ayu NOD2 in NF-\u03baB and MAPK signaling pathways: Insights into functional conservation of NOD2 in antibacterial innate immunity","volume":"40","author":"Ren","year":"2019","journal-title":"Zool Res"},{"key":"B76","doi-asserted-by":"publisher","DOI":"10.3390\/biom13010093","article-title":"PI3K\u2013AKT-targeting breast cancer treatments: natural products and synthetic compounds","volume":"13","author":"Yuan","year":"2023","journal-title":"Biomolecules"},{"key":"B77","doi-asserted-by":"publisher","first-page":"599","DOI":"10.1016\/j.ijbiomac.2022.09.195","article-title":"The variation on structure and immunomodulatory activity of polysaccharide during the longan pulp fermentation","volume":"222","author":"Hu","year":"2022","journal-title":"Int J Biol Macromolecules"},{"key":"B78","doi-asserted-by":"publisher","DOI":"10.1016\/j.phymed.2023.154859","article-title":"Ginseng polysaccharide reduces autoimmune hepatitis inflammatory response by inhibiting PI3K\/AKT and TLRs\/NF-\u03baB signaling pathways","volume":"116","author":"Qi","year":"2023","journal-title":"Phytomedicine"},{"key":"B79","doi-asserted-by":"publisher","DOI":"10.1016\/j.foodhyd.2019.105456","article-title":"Antidiabetic effects of polysaccharide from azuki bean (Vigna angularis) in type 2 diabetic rats via insulin\/PI3K\/AKT signaling pathway","volume":"101","author":"Wu","year":"2020","journal-title":"Food Hydrocolloids"},{"key":"B80","doi-asserted-by":"publisher","DOI":"10.3390\/molecules29102374","article-title":"Progress in the regulation of immune cells in the tumor microenvironment by bioactive compounds of traditional Chinese medicine","volume":"29","author":"Chen","year":"2024","journal-title":"Molecules"},{"key":"B81","doi-asserted-by":"publisher","first-page":"389","DOI":"10.1186\/s12935-021-02089-2","article-title":"New insights into M1\/M2 macrophages: key modulators in cancer progression","volume":"21","author":"Liu","year":"2021","journal-title":"Cancer Cell Int"},{"key":"B82","doi-asserted-by":"publisher","DOI":"10.3390\/nu11102264","article-title":"Astragalus polysaccharides (PG2) enhances the M1 polarization of macrophages, functional maturation of dendritic cells, and T cell-mediated anticancer immune responses in patients with lung cancer","volume":"11","author":"Bamodu","year":"2019","journal-title":"Nutrients"},{"key":"B83","doi-asserted-by":"publisher","first-page":"1534735421995256","DOI":"10.1177\/1534735421995256","article-title":"Astragalus polysaccharide injection (PG2) normalizes the neutrophil-to-lymphocyte ratio in patients with advanced lung cancer receiving immunotherapy","volume":"20","author":"Tsao","year":"2021","journal-title":"Integr Cancer therapies"},{"key":"B84","doi-asserted-by":"publisher","DOI":"10.1016\/j.biopha.2021.112016","article-title":"Codonopsis pilosula polysaccharide in synergy with dacarbazine inhibits mouse melanoma by repolarizing M2-like tumor-associated macrophages into M1-like tumor-associated macrophages","volume":"142","author":"Liu","year":"2021","journal-title":"Biomed Pharmacother"},{"key":"B85","doi-asserted-by":"publisher","first-page":"918","DOI":"10.1186\/s12885-022-10011-w","article-title":"Lentinan enhances the antitumor effects of Delta-like 1 via neutrophils","volume":"22","author":"Xu","year":"2022"},{"key":"B86","doi-asserted-by":"publisher","DOI":"10.1097\/MD.0000000000025220","article-title":"Lentinan combined with cisplatin for the treatment of non-small cell lung cancer","volume":"100","author":"Zhao","year":"2021","journal-title":"Medicine"},{"key":"B87","doi-asserted-by":"publisher","DOI":"10.1016\/j.it.2016.09.006","article-title":"Dendritic cells and cancer immunity","volume":"37","author":"Gardner","year":"2016","journal-title":"Trends Immunol"},{"key":"B88","doi-asserted-by":"publisher","DOI":"10.1016\/j.mtbio.2025.101559","article-title":"Lymph node-targeted delivery of Lonicera japonica thunb. polysaccharides for enhancing antitumor immunotherapy","volume":"31","author":"Zhang","year":"2025","journal-title":"Materials Today Bio"},{"key":"B89","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2024.136818","article-title":"Immunomodulatory effects of Fructus corni acidic polysaccharide and its selenium nanoparticles composites in hepatocellular carcinoma","volume":"282","author":"Lan","year":"2024","journal-title":"Int J Biol Macromolecules"},{"key":"B90","doi-asserted-by":"publisher","DOI":"10.1016\/j.fshw.2022.07.048","article-title":"Ginger polysaccharide UGP1 suppressed human colon cancer growth via p53, Bax\/Bcl-2, caspase-3 pathways and immunomodulation","volume":"12","author":"Qian","year":"2023","journal-title":"Food Sci Hum Wellness"},{"key":"B91","doi-asserted-by":"publisher","DOI":"10.3390\/ijms19030730","article-title":"The balance of th17 versus treg cells in autoimmunity","volume":"19","author":"Lee","year":"2018","journal-title":"Int J Mol Sci"},{"key":"B92","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2022.1021695","article-title":"Polysaccharide from Atractylodes macrocephala Koidz. ameliorates DSS-induced colitis in mice by regulating the Th17\/Treg cell balance","volume":"13","author":"Yang","year":"2022","journal-title":"Front Immunol"},{"key":"B93","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2022.08.032","article-title":"Ginger polysaccharides relieve ulcerative colitis via maintaining intestinal barrier integrity and gut microbiota modulation","volume":"219","author":"Hao","year":"2022","journal-title":"Int J Biol Macromolecules"},{"key":"B94","doi-asserted-by":"publisher","DOI":"10.1016\/j.carbpol.2020.117189","article-title":"Polysaccharides from natural resources exhibit great potential in the treatment of ulcerative colitis: A review","volume":"254","author":"Niu","year":"2021","journal-title":"Carbohydr Polymers"},{"key":"B95","first-page":"97","article-title":"Current developments in the immunology of psoriasis","volume":"93","author":"Gr\u00e4n","year":"2020","journal-title":"Yale J Biol Med"},{"key":"B96","doi-asserted-by":"publisher","DOI":"10.3389\/fphar.2021.806844","article-title":"P. granatum Peel Polysaccharides Ameliorate Imiquimod-Induced Psoriasis-Like Dermatitis in Mice via Suppression of NF-\u03baB and STAT3 Pathways","volume":"12","author":"Chen","year":"2022","journal-title":"Front Pharmacol"},{"key":"B97","doi-asserted-by":"publisher","DOI":"10.1016\/j.mam.2021.100995","article-title":"Cellular and molecular mechanisms of allergic asthma","volume":"85","author":"Koml\u00f3si","year":"2022","journal-title":"Mol Aspects Med"},{"key":"B98","doi-asserted-by":"publisher","DOI":"10.1016\/j.phymed.2024.155965","article-title":"Lentinan attenuates allergic airway inflammation and epithelial barrier dysfunction in asthma via inhibition of the PI3K\/AKT\/NF-\u03baB pathway","volume":"134","author":"Song","year":"2024","journal-title":"Phytomedicine"},{"key":"B99","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2021.11.176","article-title":"Structure, immunostimulatory activity, and the effect of ameliorating airway inflammation of polysaccharides from Pyrus sinkiangensis Yu","volume":"195","author":"Taoerdahong","year":"2022","journal-title":"Int J Biol Macromolecules"},{"key":"B100","doi-asserted-by":"publisher","DOI":"10.1016\/j.foodres.2021.110675","article-title":"Polysaccharides confer benefits in immune regulation and multiple sclerosis by interacting with gut microbiota","volume":"149","author":"Sun","year":"2021","journal-title":"Food Res Int"},{"key":"B101","doi-asserted-by":"publisher","DOI":"10.1007\/s10286-019-00605-z","article-title":"Immune and autonomic nervous system interactions in multiple sclerosis: clinical implications","volume":"29","author":"Habek","year":"2019","journal-title":"Clin Auton Res"},{"key":"B102","doi-asserted-by":"publisher","first-page":"78","DOI":"10.1016\/j.jneuroim.2019.03.019","article-title":"The PD-1\/PD-Ls pathway is up-regulated during the suppression of experimental autoimmune encephalomyelitis treated by Astragalus polysaccharides","volume":"332","author":"Sun","year":"2019","journal-title":"J Neuroimmunol"},{"key":"B103","doi-asserted-by":"publisher","DOI":"10.1142\/S0192415X16500610","article-title":"Protective effects on central nervous system by acidic polysaccharide of panax ginseng in relapse-remitting experimental autoimmune encephalomyelitis-induced SJL\/J mice","volume":"44","author":"Bing","year":"2016","journal-title":"Am J Chin Med"},{"key":"B104","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1155\/2022\/3002304","article-title":"The dectin-1 receptor signaling pathway mediates the remyelination effect of lentinan through suppression of neuroinflammation and conversion of microglia","volume":"2022","author":"Zhang","year":"2022","journal-title":"J Immunol Res"},{"key":"B105","doi-asserted-by":"publisher","DOI":"10.3390\/molecules28062483","article-title":"Potential anti-rheumatoid arthritis activities and mechanisms of ganoderma lucidum polysaccharides","volume":"28","author":"Meng","year":"2023","journal-title":"Molecules"},{"key":"B106","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2022.03.090","article-title":"Angelica sinensis polysaccharide improves rheumatoid arthritis by modifying the expression of intestinal Cldn5, Slit3 and Rgs18 through gut microbiota","volume":"209","author":"Hu","year":"2022","journal-title":"Int J Biol Macromolecules"},{"key":"B107","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1155\/2019\/8514247","article-title":"Cellulase-assisted extraction, characterization, and bioactivity against rheumatoid arthritis of astragalus polysaccharides","volume":"2019","author":"Cao","year":"2019","journal-title":"Int J Polymer Sci"},{"key":"B108","doi-asserted-by":"publisher","first-page":"45","DOI":"10.1186\/s13073-024-01300-z","article-title":"Systematic immune cell dysregulation and molecular subtypes revealed by single-cell RNA-seq of subjects with type 1 diabetes","volume":"16","author":"Honardoost","year":"2024","journal-title":"Genome Med"},{"key":"B109","doi-asserted-by":"publisher","DOI":"10.1038\/s41387-023-00233-7","article-title":"Lentinan confers protection against type 1 diabetes by inducing regulatory T cell in spontaneous non-obese diabetic mice","volume":"13","author":"Wu","year":"2023","journal-title":"Nutr Diabetes"},{"key":"B110","doi-asserted-by":"publisher","DOI":"10.1016\/j.psj.2024.103840","article-title":"Immunopotentiating effect of lentinan on chicks and its inhibitory effect on Marek\u2019s disease virus infection","volume":"103","author":"Gao","year":"2024","journal-title":"Poultry Sci"},{"key":"B111","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1186\/s40104-021-00562-6","article-title":"Lentinan administration alleviates diarrhea of rotavirus-infected weaned pigs via regulating intestinal immunity","volume":"12","author":"Fan","year":"2021","journal-title":"J Anim Sci Biotechnol"},{"key":"B112","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2022.999945","article-title":"Refined polysaccharide from Dendrobium devonianum resists H1N1 influenza viral infection in mice by activating immunity through the TLR4\/MyD88\/NF-\u03baB pathway","volume":"13","author":"Wei","year":"2022","journal-title":"Front Immunol"},{"key":"B113","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2017.10.171","article-title":"Hepatoprotective and inhibiting HBV effects of polysaccharides from roots of Sophora flavescens","volume":"108","author":"Yang","year":"2018","journal-title":"Int J Biol Macromolecules"},{"key":"B114","doi-asserted-by":"publisher","DOI":"10.1016\/j.carbpol.2021.118739","article-title":"Polysaccharides derived from Chinese medicinal herbs: A promising choice of vaccine adjuvants","volume":"276","author":"Wan","year":"2022","journal-title":"Carbohydr Polymers"},{"key":"B115","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2020.561758","article-title":"A polysaccharide purified from ganoderma lucidum acts as a potent mucosal adjuvant that promotes protective immunity against the lethal challenge with enterovirus A71","volume":"11","author":"Lin","year":"2020","journal-title":"Front Immunol"},{"key":"B116","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2023.123635","article-title":"Bidirectional and persistent immunomodulation of Astragalus polysaccharide as an adjuvant of influenza and recombinant SARS-CoV-2 vaccine","volume":"234","author":"Zhao","year":"2023","journal-title":"Int J Biol Macromolecules"},{"key":"B117","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2020.08.005","article-title":"Immune-adjuvant activity of lentinan-modified calcium carbonate microparticles on a H5N1 vaccine","volume":"163","author":"He","year":"2020","journal-title":"Int J Biol Macromolecules"},{"key":"B118","doi-asserted-by":"publisher","DOI":"10.1038\/ni.3580","article-title":"Identification of a role for TRIM29 in the control of innate immunity in the respiratory tract","volume":"17","author":"Xing","year":"2016","journal-title":"Nat Immunol"},{"key":"B119","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1701569","article-title":"TRIM29 negatively regulates the type I IFN production in response to RNA virus","volume":"201","author":"Xing","year":"2018","journal-title":"J Immunol"},{"key":"B120","doi-asserted-by":"publisher","first-page":"945","DOI":"10.1038\/s41467-017-00101-w","article-title":"TRIM29 promotes DNA virus infections by inhibiting innate immune response","volume":"8","author":"Xing","year":"2017","journal-title":"Nat Commun"},{"key":"B121","doi-asserted-by":"publisher","first-page":"3481","DOI":"10.1038\/s41467-024-44745-x","article-title":"Loss of TRIM29 mitigates viral myocarditis by attenuating PERK-driven ER stress response in male mice","volume":"15","author":"Wang","year":"2024","journal-title":"Nat Commun"},{"key":"B122","doi-asserted-by":"publisher","DOI":"10.1016\/j.mucimm.2024.10.004","article-title":"TRIM29 controls enteric RNA virus-induced intestinal inflammation by targeting NLRP6 and NLRP9b signaling pathways","volume":"18","author":"Wang","year":"2025","journal-title":"Mucosal Immunol"},{"key":"B123","doi-asserted-by":"publisher","DOI":"10.1038\/s44318-024-00126-0","article-title":"PARP14 and PARP9\/DTX3L regulate interferon-induced ADP-ribosylation","volume":"43","author":"Kar","year":"2024","journal-title":"EMBO J"},{"key":"B124","doi-asserted-by":"publisher","first-page":"2681","DOI":"10.1038\/s41467-021-23003-4","article-title":"Identification of poly(ADP-ribose) polymerase 9 (PARP9) as a noncanonical sensor for RNA virus in dendritic cells","volume":"12","author":"Xing","year":"2021","journal-title":"Nat Commun"},{"key":"B125","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2021.05.213","article-title":"Extraction, structure, pharmacological activities and drug carrier applications of Angelica sinensis polysaccharide","volume":"183","author":"Nai","year":"2021","journal-title":"Int J Biol Macromol"},{"key":"B126","doi-asserted-by":"publisher","first-page":"494","DOI":"10.1039\/D0FO01896A","article-title":"Recent progress in polysaccharides from Panax ginseng C. A. Meyer","volume":"12","author":"Guo","year":"2021","journal-title":"Food Funct"},{"key":"B127","doi-asserted-by":"publisher","DOI":"10.1016\/j.fochx.2024.101272","article-title":"Effects of UV\/H2O2 degradation on Moringa oleifera Lam. leaves polysaccharides: Composition, in vitro fermentation and prebiotic properties on gut microorganisms","volume":"22","author":"Yang","year":"2024","journal-title":"Food Chemistry: X"},{"key":"B128","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2020.11.018","article-title":"Isolation and structure elucidation of polysaccharides from fruiting bodies of mushroom Coriolus versicolor and evaluation of their immunomodulatory effects","volume":"166","author":"Zhang","year":"2021","journal-title":"Int J Biol Macromolecules"},{"key":"B129","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2020.04.141","article-title":"Modification and application of polysaccharide from traditional Chinese medicine such as Dendrobium officinale","volume":"157","author":"Huang","year":"2020","journal-title":"Int J Biol Macromolecules"},{"key":"B130","doi-asserted-by":"publisher","DOI":"10.3390\/foods13244073","article-title":"Immunoregulatory effects of codonopsis pilosula polysaccharide modified selenium nanoparticles on H22 tumor-bearing mice","volume":"13","author":"Long","year":"2024","journal-title":"Foods"},{"key":"B131","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2024.137529","article-title":"A glucose-rich heteropolysaccharide from Marsdenia tenacissima (Roxb.) Wight et Arn. and its zinc-modified complex enhance immunoregulation by regulating TLR4-Myd88-NF-\u03baB pathway","volume":"283","author":"Li","year":"2024","journal-title":"Int J Biol Macromolecules"},{"key":"B132","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2022.06.122","article-title":"Preparation, characterization and immunoregulatory activity of derivatives of polysaccharide from Atractylodes lancea (Thunb.) DC","volume":"216","author":"Liu","year":"2022","journal-title":"Int J Biol Macromolecules"},{"key":"B133","doi-asserted-by":"publisher","first-page":"94","DOI":"10.1016\/j.ijbiomac.2021.02.046","article-title":"Structure characterization and in vitro immunomodulatory activities of carboxymethyl pachymaran","volume":"178","author":"Liu","year":"2021","journal-title":"Int J Biol Macromol"},{"key":"B134","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2018.10.119","article-title":"Effect of sulfation and partial hydrolysis of polysaccharides from Polygonatum sibiricum on immune-enhancement","volume":"122","author":"Yelithao","year":"2019","journal-title":"Int J Biol Macromol"},{"key":"B135","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2023.1133050","article-title":"Active ingredients of traditional Chinese medicine for enhancing the effect of tumor immunotherapy","volume":"14","author":"Yang","year":"2023","journal-title":"Front Immunol"}],"container-title":["Frontiers in Immunology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2025.1640679\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,7,22]],"date-time":"2025-07-22T05:29:55Z","timestamp":1753162195000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2025.1640679\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,7,22]]},"references-count":135,"alternative-id":["10.3389\/fimmu.2025.1640679"],"URL":"https:\/\/doi.org\/10.3389\/fimmu.2025.1640679","relation":{},"ISSN":["1664-3224"],"issn-type":[{"value":"1664-3224","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,7,22]]},"article-number":"1640679"}}