Front. Immunol. Frontiers in Immunology Front. Immunol. 1664-3224 Frontiers Media S.A. 10.3389/fimmu.2023.1361355 Immunology Correction Corrigendum: Immunomodulatory function and anti-tumor mechanism of natural polysaccharides: a review Ying Yang 1 2 3 Hao Wu 1 2 3 * 1 Cancer Institute, The Second Affiliated Hospital, Zhejiang University School of Medicine, Hangzhou, Zhejiang, China 2 Key Laboratory of Cancer Prevention and Intervention, China National Ministry of Education, Key Laboratory of Molecular Biology in Medical Sciences, Hangzhou, Zhejiang, China 3 Zhejiang Provincial Clinical Research Center for Cancer, Cancer Center of Zhejiang University, Hangzhou, Zhejiang, China

Approved by: Frontiers Editorial Office, Frontiers Media SA, Switzerland

*Correspondence: Wu Hao, wu_hao@zju.edu.cn

09 01 2024 2023 14 1361355 25 12 2023 29 12 2023 Copyright © 2024 Ying and Hao 2024 Ying and Hao

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 A Corrigendum on Immunomodulatory function and anti-tumor mechanism of natural polysaccharides: a review By Ying Y and Hao W (2023) Front. Immunol. 14:1147641. doi: 10.3389/fimmu.2023.1147641 polysaccharides anti-tumor tumor immunomodulatory immune response section-in-acceptance Cancer Immunity and Immunotherapy

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 References Sung H Ferlay J Siegel RL Laversanne M Soerjomataram I Jemal A . Global cancer statistics 2020: GLOBOCAN estimates of incidence and mortality worldwide for 36 cancers in 185 countries. CA Cancer J Clin (2021) 71(3):209–49. doi: 10.3322/caac.21660 Xie L Shen M Hong Y Ye H Huang L Xie J . Chemical modifications of polysaccharides and their anti-tumor activities. Carbohydr Polym (2020) 229:115436. doi: 10.1016/j.carbpol.2019.115436 Yu Y Shen M Song Q Xie J . Biological activities and pharmaceutical applications of polysaccharide from natural resources: A review. Carbohydr Polym (2018) 183:91–101. doi: 10.1016/j.carbpol.2017.12.009 Manna DK Maity P Nandi AK Pattanayak M Panda BC Mandal AK . Structural elucidation and immunostimulating property of a novel polysaccharide extracted from an edible mushroom Lentinus fusipes. Carbohydr Polym (2017) 157:1657–65. doi: 10.1016/j.carbpol.2016.11.048 Fan S Zhang J Nie W Zhou W Jin L Chen X . Antitumor effects of polysaccharide from Sargassum fusiforme against human hepatocellular carcinoma HepG2 cells. Food Chem Toxicol (2017) 102:53–62. doi: 10.1016/j.fct.2017.01.020 He R Zhao Y Zhao R Sun P . Antioxidant and antitumor activities in vitro of polysaccharides from E. sipunculoides. Int J Biol Macromol (2015) 78:56–61. doi: 10.1016/j.ijbiomac.2015.03.030 Moradali MF Mostafavi H Ghods S Hedjaroude GA . Immunomodulating and anticancer agents in the realm of macromycetes fungi (macrofungi). Int Immunopharmacol (2007) 7(6):701–24. doi: 10.1016/j.intimp.2007.01.008 Zheng Z Pan X Luo L Zhang Q Huang X Liu Y . Advances in oral absorption of polysaccharides: Mechanism, affecting factors, and improvement strategies. Carbohydr Polym (2022) 282:119110. doi: 10.1016/j.carbpol.2022.119110 Wasser SP . Medicinal mushrooms as a source of antitumor and immunomodulating polysaccharides. Appl Microbiol Biotechnol (2002) 60(3):258–74. doi: 10.1007/s00253-002-1076-7 Yin M Zhang Y Li H . Advances in research on immunoregulation of macrophages by plant polysaccharides. Front Immunol (2019) 10:145. doi: 10.3389/fimmu.2019.00145 Li M Wang X Wang Y Bao S Chang Q Liu L . Strategies for remodeling the tumor microenvironment using active ingredients of ginseng-A promising approach for cancer therapy. Front Pharmacol (2021) 12:797634. doi: 10.3389/fphar.2021.797634 Wang D Cui Q Yang YJ Liu AQ Zhang G Yu JC . 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. BioMed Pharmacother (2022) 155:113541. doi: 10.1016/j.biopha.2022.113541 Zhang M Cui SW Cheung PCK Wang Q . Antitumor polysaccharides from mushrooms: a review on their isolation process, structural characteristics and antitumor activity. Trends Food Sci Technol (2007) 18(1):4–19. doi: 10.1016/j.tifs.2006.07.013 Maity P Sen IK Chakraborty I Mondal S Bar H Bhanja SK . Biologically active polysaccharide from edible mushrooms: A review. Int J Biol Macromol (2021) 172:408–17. doi: 10.1016/j.ijbiomac.2021.01.081 Yelithao K Surayot U Lee C Palanisamy S Prabhu NM Lee J . Studies on structural properties and immune-enhancing activities of glycomannans from Schizophyllum commune. Carbohydr Polym (2019) 218:37–45. doi: 10.1016/j.carbpol.2019.04.057 Xia S Zhai Y Wang X Fan Q Dong X Chen M . Phosphorylation of polysaccharides: A review on the synthesis and bioactivities. Int J Biol Macromol (2021) 184:946–54. doi: 10.1016/j.ijbiomac.2021.06.149 Chen F Huang G . Preparation and immunological activity of polysaccharides and their derivatives. Int J Biol Macromol (2018) 112:211–6. doi: 10.1016/j.ijbiomac.2018.01.169 Xu Y Wu YJ Sun PL Zhang FM Linhardt RJ Zhang AQ . Chemically modified polysaccharides: Synthesis, characterization, structure activity relationships of action. Int J Biol Macromol (2019) 132:970–7. doi: 10.1016/j.ijbiomac.2019.03.213 Guo MQ Hu X Wang C Ai L . "Polysaccharides: structure and solubility". In: Solubility of polysaccharides. London: INTECH (2017). p. 7–20. doi: 10.5772/intechopen.71570 Ren L Perera C Hemar Y . Antitumor activity of mushroom polysaccharides: a review. Food Funct (2012) 3(11):1118–30. doi: 10.1039/c2fo10279j Guo MZ Meng M Duan SQ Feng CC Wang CL . Structure characterization, physicochemical property and immunomodulatory activity on RAW264.7 cells of a novel triple-helix polysaccharide from Craterellus cornucopioides. Int J Biol Macromol (2019) 126:796–804. doi: 10.1016/j.ijbiomac.2018.12.246 Jimenez-Medina E Berruguilla E Romero I Algarra I Collado A Garrido F . The immunomodulator PSK induces in vitro cytotoxic activity in tumour cell lines via arrest of cell cycle and induction of apoptosis. BMC Cancer (2008) 8:78. doi: 10.1186/1471-2407-8-78 Liao W Luo Z Liu D Ning Z Yang J Ren J . Structure characterization of a novel polysaccharide from Dictyophora indusiata and its macrophage immunomodulatory activities. J Agric Food Chem (2015) 63(2):535–44. doi: 10.1021/jf504677r Maity P Sen IK Maji PK Paloi S Devi KS Acharya K . Structural, immunological, and antioxidant studies of beta-glucan from edible mushroom Entoloma lividoalbum. Carbohydr Polym (2015) 123:350–8. doi: 10.1016/j.carbpol.2015.01.051 Zhang T Ye J Xue C Wang Y Liao W Mao L . Structural characteristics and bioactive properties of a novel polysaccharide from Flammulina velutipes. Carbohydr Polym (2018) 197:147–56. doi: 10.1016/j.carbpol.2018.05.069 Li WJ Nie SP Xie MY Yu Q Chen Y He M . Ganoderma atrum polysaccharide attenuates oxidative stress induced by d-galactose in mouse brain. Life Sci (2011) 88(15-16):713–8. doi: 10.1016/j.lfs.2011.02.010 Li WJ Nie SP Peng XP Liu XZ Li C Chen Y . Ganoderma atrum polysaccharide improves age-related oxidative stress and immune impairment in mice. J Agric Food Chem (2012) 60(6):1413–8. doi: 10.1021/jf204748a Yu Q Nie SP Li WJ Zheng WY Yin PF Gong DM . Macrophage immunomodulatory activity of a purified polysaccharide isolated from Ganoderma atrum. Phytother Res (2013) 27(2):186–91. doi: 10.1002/ptr.4698 Yu Q Nie SP Wang JQ Yin PF Huang DF Li WJ . Toll-like receptor 4-mediated ROS signaling pathway involved in Ganoderma atrum polysaccharide-induced tumor necrosis factor-alpha secretion during macrophage activation. Food Chem Toxicol (2014) 66:14–22. doi: 10.1016/j.fct.2014.01.018 Yu Q Nie SP Wang JQ Huang DF Li WJ Xie MY . Signaling pathway involved in the immunomodulatory effect of Ganoderma atrum polysaccharide in spleen lymphocytes. J Agric Food Chem (2015) 63(10):2734–40. doi: 10.1021/acs.jafc.5b00028 Xiang QD Yu Q Wang H Zhao MM Liu SY Nie SP . Immunomodulatory Activity of Ganoderma atrum Polysaccharide on Purified T Lymphocytes through Ca(2+)/CaN and Mitogen-Activated Protein Kinase Pathway Based on RNA Sequencing. J Agric Food Chem (2017) 65(26):5306–15. doi: 10.1021/acs.jafc.7b01763 Zhang J Tang Q Zimmerman-Kordmann M Reutter W Fan H . Activation of B lymphocytes by GLIS, a bioactive proteoglycan from Ganoderma lucidum. Life Sci (2002) 71(6):623–38. doi: 10.1016/S0024-3205(02)01690-9 Liu MM Zeng P Li XT Shi LG . Antitumor and immunomodulation activities of polysaccharide from Phellinus baumii. Int J Biol Macromol (2016) 91:1199–205. doi: 10.1016/j.ijbiomac.2016.06.086 Liu Y Liu Y Jiang H Xu L Cheng Y Wang PG . Preparation, antiangiogenic and antitumoral activities of the chemically sulfated glucan from Phellinus ribis. Carbohydr Polym (2014) 106:42–8. doi: 10.1016/j.carbpol.2014.01.088 Wang Q Niu LL Liu HP Wu YR Li MY Jia Q . Structural characterization of a novel polysaccharide from Pleurotus citrinopileatus and its antitumor activity on H22 tumor-bearing mice. Int J Biol Macromol (2021) 168:251–60. doi: 10.1016/j.ijbiomac.2020.12.053 Zhang Y Zhang Z Liu H Wang J Wang Deng Z . Physicochemical characterization and antitumor activity in vitro of a selenium polysaccharide from Pleurotus ostreatus. Int J Biol Macromol (2020) 165(Pt B):2934–46. doi: 10.1016/j.ijbiomac.2020.10.168 Li X Xu W Chen J . Polysaccharide purified from Polyporus umbellatus (Per) Fr induces the activation and maturation of murine bone-derived dendritic cells via toll-like receptor 4. Cell Immunol (2010) 265(1):50–6. doi: 10.1016/j.cellimm.2010.07.002 Zhang GW Qin GF Han B Li CX Yang HG Nie PH . Efficacy of Zhuling polyporus polysaccharide with BCG to inhibit bladder carcinoma. Carbohydr Polym (2015) 118:30–5. doi: 10.1016/j.carbpol.2014.11.012 Guo Z Zang Y Zhang L . The efficacy of Polyporus Umbellatus polysaccharide in treating hepatitis B in China. Prog Mol Biol Transl Sci (2019) 163:329–60. doi: 10.1016/bs.pmbts.2019.03.012 Chen Y Li X-H Zhou L-Y Li W Liu L Wang D-D . Structural elucidation of three antioxidative polysaccharides from Tricholoma lobayense. Carbohydr Polymers (2017) 157:484–92. doi: 10.1016/j.carbpol.2016.10.011 Xie Y Wang L Sun H Shang Q Wang Y Zhang G . A polysaccharide extracted from alfalfa activates splenic B cells by TLR4 and acts primarily via the MAPK/p38 pathway. Food Funct (2020) 11(10):9035–47. doi: 10.1039/D0FO01711F Wang J Bao A Wang Q Guo H Zhang Y Liang J . Sulfation can enhance antitumor activities of Artemisia sphaerocephala polysaccharide in vitro and vivo . Int J Biol Macromol (2018) 107(Pt A):502–11. doi: 10.1016/j.ijbiomac.2017.09.018 Shi S Chang M Liu H Ding S Yan Z Si K . The structural characteristics of an acidic water-soluble polysaccharide from bupleurum chinense DC and its in vivo anti-tumor activity on H22 tumor-bearing mice. Polymers (Basel) (2022) 14(6):1119. doi: 10.3390/polym14061119 Park HB Lim SM Hwang J Zhang W You S Jin JO . Cancer immunotherapy using a polysaccharide from Codium fragile in a murine model. Oncoimmunology (2020) 9(1):1772663. doi: 10.1080/2162402X.2020.1772663 Zhang W Hwang J Park HB Lim SM Go S Kim J . Human peripheral blood dendritic cell and T cell activation by codium fragile polysaccharide. Mar Drugs (2020) 18(11):535. doi: 10.3390/md18110535 Zheng YS Wu ZS Ni HB Ke L Tong ZH Li WQ . Codonopsis pilosula polysaccharide attenuates cecal ligation and puncture sepsis via circuiting regulatory T cells in mice. Shock (2014) 41(3):250–5. doi: 10.1097/SHK.0000000000000091 Mazepa E Noseda MD Ferreira LG de Carvalho MM Goncalves AG Ducatti DRB . Chemical structure of native and modified sulfated heterorhamnans from the green seaweed Gayralia brasiliensis and their cytotoxic effect on U87MG human glioma cells. Int J Biol Macromol (2021) 187:710–21. doi: 10.1016/j.ijbiomac.2021.07.145 Wang H Bi H Gao T Zhao B Ni W Liu J . A homogalacturonan from Hippophae rhamnoides L. Berries enhance immunomodulatory activity through TLR4/MyD88 pathway mediated activation of macrophages. Int J Biol Macromol (2018) 107(Pt A):1039–45. doi: 10.1016/j.ijbiomac.2017.09.083 Fang Q Wang JF Zha XQ Cui SH Cao L Luo JP . Immunomodulatory activity on macrophage of a purified polysaccharide extracted from Laminaria japonica. Carbohydr Polym (2015) 134:66–73. doi: 10.1016/j.carbpol.2015.07.070 Perez-Recalde M Matulewicz MC Pujol CA Carlucci MJ . In vitro and in vivo immunomodulatory activity of sulfated polysaccharides from red seaweed Nemalion helminthoides. Int J Biol Macromol (2014) 63:38–42. doi: 10.1016/j.ijbiomac.2013.10.024 Sun W Hu W Meng K Yang L Zhang W Song X . Activation of macrophages by the ophiopogon polysaccharide liposome from the root tuber of Ophiopogon japonicus. Int J Biol Macromol (2016) 91:918–25. doi: 10.1016/j.ijbiomac.2016.06.037 Wang X Gao A Jiao Y Zhao Y Yang X . Antitumor effect and molecular mechanism of antioxidant polysaccharides from Salvia miltiorrhiza Bunge in human colorectal carcinoma LoVo cells. Int J Biol Macromol (2018) 108:625–34. doi: 10.1016/j.ijbiomac.2017.12.006 Chen X Yu G Fan S Bian M Ma H Lu J . Sargassum fusiforme polysaccharide activates nuclear factor kappa-B (NF-kappaB) and induces cytokine production via Toll-like receptors. Carbohydr Polym (2014) 105:113–20. doi: 10.1016/j.carbpol.2014.01.056 Chen H Zhang L Long X Li P Chen S Kuang W . Sargassum fusiforme polysaccharides inhibit VEGF-A-related angiogenesis and proliferation of lung cancer in vitro and in vivo. BioMed Pharmacother (2017) 85:22–7. doi: 10.1016/j.biopha.2016.11.131 Tang X Huang J Xiong H Zhang K Chen C Wei X . Anti-tumor effects of the polysaccharide isolated from tarphochlamys affinis in H22 tumor-bearing mice. Cell Physiol Biochem (2016) 39(3):1040–50. doi: 10.1159/000447811 Gupta PK Rajan MGR Kulkarni S . Activation of murine macrophages by G1-4A, a polysaccharide from Tinospora cordifolia, in TLR4/MyD88 dependent manner. Int Immunopharmacol (2017) 50:168–77. doi: 10.1016/j.intimp.2017.06.025 Wang Y Wang S Song R Cai J Xu J Tang X . Ginger polysaccharides induced cell cycle arrest and apoptosis in human hepatocellular carcinoma HepG2 cells. Int J Biol Macromol (2019) 123:81–90. doi: 10.1016/j.ijbiomac.2018.10.169 He R Ye J Zhao Y Su W . Partial characterization, antioxidant and antitumor activities of polysaccharides from Philomycusbilineatus. Int J Biol Macromol (2014) 65:573–80. doi: 10.1016/j.ijbiomac.2014.01.016 Zhao H Li Y Wang Y Zhang J Ouyang X Peng R . Antitumor and immunostimulatory activity of a polysaccharide-protein complex from Scolopendra subspinipes mutilans L. Koch in tumor-bearing mice. Food Chem Toxicol (2012) 50(8):2648–55. doi: 10.1016/j.fct.2012.05.018 Yuan P Fang F Shao T Li P Hu W Zhou Y . Structure and anti-tumor activities of exopolysaccharides from alternaria Mali roberts. Molecules (2019) 24(7):1345. doi: 10.3390/molecules24071345 Liu WB Xie F Sun HQ Meng M Zhu ZY . Anti-tumor effect of polysaccharide from Hirsutella sinensis on human non-small cell lung cancer and nude mice through intrinsic mitochondrial pathway. Int J Biol Macromol (2017) 99:258–64. doi: 10.1016/j.ijbiomac.2017.02.071 Li S Gao A Dong S Chen Y Sun S Lei Z . Purification, antitumor and immunomodulatory activity of polysaccharides from soybean residue fermented with Morchella esculenta. Int J Biol Macromol (2017) 96:26–34. doi: 10.1016/j.ijbiomac.2016.12.007 Zhang X Ding R Zhou Y Zhu R Liu W Jin L . Toll-like receptor 2 and Toll-like receptor 4-dependent activation of B cells by a polysaccharide from marine fungus Phoma herbarum YS4108. PloS One (2013) 8(3):e60781. doi: 10.1371/journal.pone.0060781 Huang T Lin J Cao J Zhang P Bai Y Chen G . An exopolysaccharide from Trichoderma pseudokoningii and its apoptotic activity on human leukemia K562 cells. Carbohydr Polym (2012) 89(2):701–8. doi: 10.1016/j.carbpol.2012.03.079 Falch BH Espevik T Ryan L Stokke BT . The cytokine stimulating activity of (1–>3)-beta-D-glucans is dependent on the triple helix conformation. Carbohydr Res (2000) 329(3):587–96. doi: 10.1016/S0008-6215(00)00222-6 Hanahan D Weinberg RA . Hallmarks of cancer: the next generation. Cell (2011) 144(5):646–74. doi: 10.1016/j.cell.2011.02.013 Leibovici J Itzhaki O Huszar M Sinai J . The tumor microenvironment: part 1. Immunotherapy (2011) 3(11):1367–84. doi: 10.2217/imt.11.111 Kellum JA Chawla LS . Cell-cycle arrest and acute kidney injury: the light and the dark sides. Nephrol Dial Transplant (2016) 31(1):16–22. doi: 10.1093/ndt/gfv130 Giono LE Manfredi JJ . The p53 tumor suppressor participates in multiple cell cycle checkpoints. J Cell Physiol (2006) 209(1):13–20. doi: 10.1002/jcp.20689 Zhou X Hao Q Lu H . Mutant p53 in cancer therapy-the barrier or the path. J Mol Cell Biol (2019) 11(4):293–305. doi: 10.1093/jmcb/mjy072 Elmore S . Apoptosis: a review of programmed cell death. Toxicologic Pathol (2007) 35(4):495–516. doi: 10.1080/01926230701320337 Reed JC . Mechanisms of apoptosis. Am J Pathol (2000) 157(5):1415–30. doi: 10.1016/S0002-9440(10)64779-7 Cory S Adams JM . The Bcl2 family: regulators of the cellular life-or-death switch. Nat Rev Cancer (2002) 2(9):647–56. doi: 10.1038/nrc883 Soares R Meireles M Rocha A Pirraco A Obiol D Alonso E . Maitake (D fraction) mushroom extract induces apoptosis in breast cancer cells by BAK-1 gene activation. J Medicinal Food (2011) 14(6):563–72. doi: 10.1089/jmf.2010.0095 Acharya A Das I Chandhok D Saha T . Redox regulation in cancer: a double-edged sword with therapeutic potential. Oxid Med Cell Longevity (2010) 3(1):23–34. doi: 10.4161/oxim.3.1.10095 Fuchs-Tarlovsky V . Role of antioxidants in cancer therapy. Nutrition (2013) 29(1):15–21. doi: 10.1016/j.nut.2012.02.014 Prasad S Gupta SC Tyagi AK . Reactive oxygen species (ROS) and cancer: Role of antioxidative nutraceuticals. Cancer Lett (2017) 387:95–105. doi: 10.1016/j.canlet.2016.03.042 Block KI Koch AC Mead MN Tothy PK Newman RA Gyllenhaal C . Impact of antioxidant supplementation on chemotherapeutic toxicity: a systematic review of the evidence from randomized controlled trials. Int J Cancer (2008) 123(6):1227–39. doi: 10.1002/ijc.23754 Sosa V Moline T Somoza R Paciucci R Kondoh H LLeonart ME . Oxidative stress and cancer: an overview. Ageing Res Rev (2013) 12(1):376–90. doi: 10.1016/j.arr.2012.10.004 Anderson NM Simon MC . The tumor microenvironment. Curr Biol (2020) 30(16):R921–5. doi: 10.1016/j.cub.2020.06.081 Liu L Nie S Xie M . Tumor microenvironment as a new target for tumor immunotherapy of polysaccharides. Crit Rev Food Sci Nutr (2016) 56 Suppl 1:S85–94. doi: 10.1080/10408398.2015.1077191 Wang WJ Wu YS Chen S Liu CF Chen SN . Mushroom beta-glucan may immunomodulate the tumor-associated macrophages in the lewis lung carcinoma. BioMed Res Int (2015) 2015:604385. doi: 10.1155/2015/604385 Nishimura T Nakui M Sato M Iwakabe K Kitamura H Sekimoto M . The critical role of Th1-dominant immunity in tumor immunology. Cancer Chemotherapy Pharmacol (2000) 46(1):S52–61. doi: 10.1007/PL00014051 Hu H-M Urba WJ Fox BA . Gene-modified tumor vaccine with therapeutic potential shifts tumor-specific T cell response from a type 2 to a type 1 cytokine profile. J Immunol (1998) 161(6):3033. doi: 10.4049/jimmunol.161.6.3033 Lim SM Park HB Jin JO . Polysaccharide from Astragalus membranaceus promotes the activation of human peripheral blood and mouse spleen dendritic cells. Chin J Nat Med (2021) 19(1):56–62. doi: 10.1016/S1875-5364(21)60006-7 Ren L Zhang J Zhang T . Immunomodulatory activities of polysaccharides from Ganoderma on immune effector cells. Food Chem (2021) 340:127933. doi: 10.1016/j.foodchem.2020.127933 Zhao L Dong Y Chen G Hu Q . Extraction, purification, characterization and antitumor activity of polysaccharides from Ganoderma lucidum. Carbohydr Polymers (2010) 80(3):783–9. doi: 10.1016/j.carbpol.2009.12.029 Wang S-Y Hsu M-L Hsu H-C Lee S-S Shiao M-S Ho C-K . The anti-tumor effect of Ganoderma Lucidum is mediated by cytokines released from activated macrophages and T lymphocytes. Int J Cancer (1997) 70(6):699–705. doi: 10.1002/(SICI)1097-0215(19970317)70:6<699::AID-IJC12>3.0.CO;2-5 Hsu JW Huang HC Chen ST Wong CH Juan HF . Ganoderma lucidum Polysaccharides Induce Macrophage-Like Differentiation in Human Leukemia THP-1 Cells via Caspase and p53 Activation. Evid Based Complement Alternat Med 2011 (2011) p:358717. doi: 10.1093/ecam/nep107 Hsu TL Cheng SC Yang WB Chin SW Chen BH Huang MT . Profiling carbohydrate-receptor interaction with recombinant innate immunity receptor-Fc fusion proteins. J Biol Chem (2009) 284(50):34479–89. doi: 10.1074/jbc.M109.065961 Brown GD Taylor PR Reid DM Willment JA Williams DL Martinez-Pomares L . Dectin-1 is a major beta-glucan receptor on macrophages. J Exp Med (2002) 196(3):407–12. doi: 10.1084/jem.20020470 Palma AS Feizi T Zhang Y Stoll MS Lawson AM Diaz-Rodriguez E . Ligands for the beta-glucan receptor, Dectin-1, assigned using "designer" microarrays of oligosaccharide probes (neoglycolipids) generated from glucan polysaccharides. J Biol Chem (2006) 281(9):5771–9. doi: 10.1074/jbc.M511461200 Lee JS Hong EK . Immunostimulating activity of the polysaccharides isolated from Cordyceps militaris. Int Immunopharmacol (2011) 11(9):1226–33. doi: 10.1016/j.intimp.2011.04.001 Li WJ Tang XF Shuai XX Jiang CJ Liu X Wang LF . Mannose receptor mediates the immune response to ganoderma atrum polysaccharides in macrophages. J Agric Food Chem (2017) 65(2):348–57. doi: 10.1021/acs.jafc.6b04888 Hsu HY Hua KF Lin CC Lin CH Hsu J Wong CH . Extract of Reishi polysaccharides induces cytokine expression via TLR4-modulated protein kinase signaling pathways. J Immunol (2004) 173(10):5989–99. doi: 10.4049/jimmunol.173.10.5989 Mazgaeen L Gurung P . Recent advances in lipopolysaccharide recognition systems. Int J Mol Sci (2020) 21(2):379. doi: 10.3390/ijms21020379 Liu GK Yang TX Wang JR . Polysaccharides from Polyporus umbellatus: A review on their extraction, modification, structure, and bioactivities. Int J Biol Macromol (2021) 189:124–34. doi: 10.1016/j.ijbiomac.2021.08.101 Zhou L Liu Z Wang Z Yu S Long T Zhou X . Astragalus polysaccharides exerts immunomodulatory effects via TLR4-mediated MyD88-dependent signaling pathway in vitro and in vivo. Sci Rep (2017) 17(7):44822. doi: 10.1038/srep44822 Li W Hu X Wang S Jiao Z Sun T Liu T . Characterization and anti-tumor bioactivity of astragalus polysaccharides by immunomodulation. Int J Biol Macromol (2020) 145:985–97. doi: 10.1016/j.ijbiomac.2019.09.189 Wang Y An E-K Kim S-J You S Jin J-O . Intranasal Administration of Codium fragile Polysaccharide Elicits Anti-Cancer Immunity against Lewis Lung Carcinoma. Int J Mol Sci (2021) 22(19):10608. doi: 10.3390/ijms221910608 Tanaka A Sakaguchi S . Regulatory T cells in cancer immunotherapy. Cell Res (2017) 27(1):109–18. doi: 10.1038/cr.2016.151 Bullard DC . "Cr3". In: The complement factsBook. Barnum S Schein T , editors. San Diego, CA: Academic Press (2018). p. 435–50. doi: 10.1016/B978-0-12-810420-0.00041-9 Xiao Z Zhou W Zhang Y . Chapter ten - fungal polysaccharides. In: Du G , editor. Advances in pharmacology. San Diego, CA: Academic Press (2020). p. 277–99. doi: 10.1016/bs.apha.2019.08.003 Lu CC Hsu YJ Chang CJ Lin CS Martel J Ojcius DM . Immunomodulatory properties of medicinal mushrooms: differential effects of water and ethanol extracts on NK cell-mediated cytotoxicity. Innate Immun (2016) 22(7):522–33. doi: 10.1177/1753425916661402 Huang Q Li L Chen H Liu Q Wang Z . GPP (Composition of ganoderma lucidum poly-saccharides and polyporus umbellatus poly-saccharides) enhances innate immune function in mice. Nutrients (2019) 11(7):1480. doi: 10.3390/nu11071480 Ma Y Wu X Yu J Zhu J Pen X Meng X . Can polysaccharide K improve therapeutic efficacy and safety in gastrointestinal cancer? a systematic review and network meta-analysis. Oncotarget (2017) 8(51):89108–18. doi: 10.18632/oncotarget.19059 Chan SL Yeung JH . Effects of polysaccharide peptide (PSP) from Coriolus versicolor on the pharmacokinetics of cyclophosphamide in the rat and cytotoxicity in HepG2 cells. Food Chem Toxicol (2006) 44(5):689–94. doi: 10.1016/j.fct.2005.10.001 Wan JM-F Sit W-H Louie JC-Y . Polysaccharopeptide enhances the anticancer activity of doxorubicin and etoposide on human breast cancer cells ZR-75-30. Int J Oncol (2008) 32(3):689–99. doi: 10.3892/ijo.32.3.689 Wan JM Sit WH Yang X Jiang P Wong LL . Polysaccharopeptides derived from Coriolus versicolor potentiate the S-phase specific cytotoxicity of Camptothecin (CPT) on human leukemia HL-60 cells. Chin Med (2010) 5:16. doi: 10.1186/1749-8546-5-16 Li J Bao Y Lam W Li W Lu F Zhu X . Immunoregulatory and anti-tumor effects of polysaccharopeptide and Astragalus polysaccharides on tumor-bearing mice. Immunopharmacol Immunotoxicol (2008) 30(4):771–82. doi: 10.1080/08923970802279183 Zong A Cao H Wang F . Anticancer polysaccharides from natural resources: a review of recent research. Carbohydr Polym (2012) 90(4):1395–410. doi: 10.1016/j.carbpol.2012.07.026 Fritz H Kennedy DA Ishii M Fergusson D Fernandes R Cooley K . Polysaccharide K and Coriolus versicolor extracts for lung cancer: a systematic review. Integr Cancer Ther (2015) 14(3):201–11. doi: 10.1177/1534735415572883 Yamasaki A Onishi H Imaizumi A Kawamoto M Fujimura A Oyama Y . Protein-bound polysaccharide-K inhibits hedgehog signaling through down-regulation of MAML3 and RBPJ transcription under hypoxia, suppressing the Malignant phenotype in pancreatic cancer. Anticancer Res (2016) 36(8):3945. Li X He Y Zeng P Liu Y Zhang M Hao C . Molecular basis for Poria cocos mushroom polysaccharide used as an antitumour drug in China. J Cell Mol Med (2019) 23(1):4–20. doi: 10.1111/jcmm.13564 Zeng P Guo Z Zeng X Hao C Zhang Y Zhang M . Chemical, biochemical, preclinical and clinical studies of Ganoderma lucidum polysaccharide as an approved drug for treating myopathy and other diseases in China. J Cell Mol Med (2018) 22(7):3278–97. doi: 10.1111/jcmm.13613 He Y Zhang L Wang H . The biological activities of the antitumor drug Grifola frondosa polysaccharide. Prog Mol Biol Transl Sci (2019) 163:221–61. doi: 10.1016/bs.pmbts.2019.02.010 Liu YH Qin HY Zhong YY Li S Wang HJ Wang H . Neutral polysaccharide from Panax notoginseng enhanced cyclophosphamide antitumor efficacy in hepatoma H22-bearing mice. BMC Cancer (2021) 21(1):37. doi: 10.1186/s12885-020-07742-z Chen J Pang W Kan Y Zhao L He Z Shi W . Structure of a pectic polysaccharide from Pseudostellaria heterophylla and stimulating insulin secretion of INS-1 cell and distributing in rats by oral. Int J Biol Macromol (2018) 106:456–63. doi: 10.1016/j.ijbiomac.2017.08.034 Opanasopit P Aumklad P Kowapradit J Ngawhiranpat T Apirakaramwong A Rojanarata T . Effect of salt forms and molecular weight of chitosans on in vitro permeability enhancement in intestinal epithelial cells (Caco-2). Pharm Dev Technol (2007) 12(5):447–55. doi: 10.1080/10837450701555901 Wang Y Bai X Hu B Xing M Cao Q Ji A . Transport mechanisms of polymannuronic acid and polyguluronic acid across caco-2 cell monolayers. Pharmaceutics (2020) 12(2):167. doi: 10.3390/pharmaceutics12020167 Hisada N Satsu H Mori A Totsuka M Kamei J Nozawa T . Low-molecular-weight hyaluronan permeates through human intestinal Caco-2 cell monolayers via the paracellular pathway. Biosci Biotechnol Biochem (2008) 72(4):1111–4. doi: 10.1271/bbb.70748 Lee DY Park K Kim SK Park RW Kwon IC Kim SY . Antimetastatic effect of an orally active heparin derivative on experimentally induced metastasis. Clin Cancer Res (2008) 14(9):2841–9. doi: 10.1158/1078-0432.CCR-07-0641 Gao Y He L Katsumi H Sakane T Fujita T Yamamoto A . Improvement of intestinal absorption of water-soluble macromolecules by various polyamines: intestinal mucosal toxicity and absorption-enhancing mechanism of spermine. Int J Pharm (2008) 354(1-2):126–34. doi: 10.1016/j.ijpharm.2007.11.061 Gao Y He L Katsumi H Sakane T Fujita T Yamamoto A . Improvement of intestinal absorption of insulin and water-soluble macromolecular compounds by chitosan oligomers in rats. Int J Pharm (2008) 359(1-2):70–8. doi: 10.1016/j.ijpharm.2008.03.016 Bernkop-Schnürch A Kast CE Guggi D . Permeation enhancing polymers in oral delivery of hydrophilic macromolecules: thiomer/GSH systems. J Control Release (2003) 93(2):95–103. doi: 10.1016/j.jconrel.2003.05.001