Emerging Sources of Glycosaminoglycans: Can Fungi Be the New Alternative?
Keywords:
Bioactive compounds, Fungal glycosaminoglycans, Medicinal mushrooms, PolysaccharidesAbstract
Glycosaminoglycans are polysaccharides, widely studied for their medicinal properties such as anticoagulant, antitumor and immunomodulatory as well as for their applications in cosmetic products. Although GAGs have traditionally been sourced from animals such as cattle, pigs, sheep, and marine organisms, evidence suggests that some fungi can also produce GAGs or GAG-like compounds, expanding the potential resources. Given the high market value of GAGs and their wide industrial applications, the limited supply from current sources is insufficient to meet global demand including non-animal, vegetarian or vegan products. This review provides a comprehensive overview of the occurrence, structural diversity, and health benefits of both animal and fungal GAGs. It examines their potential uses in medicine, nutraceuticals, and biotechnology, while highlighting market trends, future prospects, and research gaps. By highlighting the role of fungi as renewable and versatile GAG producers, this work offers a new perspective on sustainable alternatives to traditional animal-based production systems.
References
Abdallah MM, Fernández N, Matias AA and Bronze MdR. 2020 – Hyaluronic acid and chondroitin sulfate from marine and terrestrial sources: Extraction and purification methods. Carbohydrate Polymers 243, 116441. Doi 10.1016/j.carbpol.2020.116441
Acquisto NM. 2014 – Heparin. In: Wexler P (ed) Encyclopedia of Toxicology: Third Edition, Academic Press, p. 5220.
Afratis N, Gialeli C, Nikitovic D, Tsegenidis T, et. al. 2012 – Glycosaminoglycans: key players in cancer cell biology and treatment. The FEBS Journal 279, 1177–1197. Doi 10.1111/j.1742-4658.2012.08529.x
Ajisaka K, Oyanagi Y, Miyazaki T, Suzuki Y. 2016 – Effect of the chelation of metal cation on the antioxidant activity of chondroitin sulfates. Bioscience, Biotechnology, and Biochemistry 80, 1179–1185. Doi 10.1080/09168451.2016.1141036
Al-Hakim A. 2021 – General considerations for diversifying heparin drug products by improving the current heparin manufacturing process and reintroducing bovine sourced heparin to the US market. Clinical and Applied Thrombosis/Hemostasis 27, 10760296211052292. Doi 10.1177/10760296211052293
American Board of Cosmetic Surgery. 2025 – Injectible filler guide. Available at: https://www.americanboardcosmeticsurgery.org/procedure-learning-center/non-surgical/ injectable-fillers-guide/#:~:text=It helps keep skin plump,discomfort during and after treatment (Accessed on January 23, 2025).
Anderegg U, Halfter N, Schnabelrauch M, Hintze V. 2021 – Collagen/glycosaminoglycan-based matrices for controlling skin cell responses. Biological Chemistry 402, 1325–1335. Doi 10.1515/hsz-2021-0176
Andrade GP V, Lima MA, de Souza Junior AA, Fareed J, et al. 2013 – A heparin-like compound isolated from a marine crab rich in glucuronic acid 2-O-sulfate presents low anticoagulant activity. Carbohydrate Polymers 94, 647–654. Doi 10.1016/j.carbpol.2013.01.069
Andrgie AT, Birhan YS, Mekonnen TW, Hanurry EY, et al. 2019 – Redox-responsive heparin–chlorambucil conjugate polymeric prodrug for improved anti-tumor activity. Polymers 12, 43. Doi 10.3390/polym12010043
Arumugam M, Garg H, Ajithkumar T, Shanmugam A. 2009 – Antiproliferative heparin (glycosaminoglycans) isolated from giant clam (Tridacna maxima) and green mussel (Perna viridis). African Journal of Biotechnology 8, 2394–2396.
Aya KL, Stern R. 2014 – Hyaluronan in wound healing: Rediscovering a major player. Wound Repair and Regeneration 22, 579–593. Doi 10.1111/wrr.12214
Badalyan SM, Barkhudaryan A, Rapior S. 2022 – Medicinal macrofungi as cosmeceuticals: A review. International Journal of Medicinal Mushrooms 24, 1–13. Doi 10.1615/IntJMedMushrooms.2022043124
Badri A, Williams A, Awofiranye A, Datta P, et al. 2021 – Complete biosynthesis of a sulfated chondroitin in Escherichia coli. Nature Communications 12, 1389. Doi 10.1038/s41467-021-21692-5
Badri A, Williams A, Linhardt RJ, Koffas MA. 2018 – The road to animal-free glycosaminoglycan production: current efforts and bottlenecks. Current Opinion in Biotechnology 53, 85–92. Doi 10.1016/j.copbio.2017.12.018
Baig N, Kouta A, Jeske W, Hoppensteadt D, et al. 2020 – Validation of the bioequivalence of USP potency adjusted porcine, ovine, and bovine heparins. Blood 136, 6. Doi 10.1182/blood-2020-142861
Balazs EA, Laurent TC, Jeanloz RW. 1986 – Nomenclature of hyaluronic acid. The Biochemical journal 235, 903. Doi 10.1042/bj2350903
Bandara AR, Rapior S, Bhat DJ, Kakumyan P, et al. 2015 – Polyporus umbellatus, an edible-medicinal cultivated mushroom with multiple developed health-care products as food, medicine and cosmetics: A review. Cryptogamie, Mycologie 36, 3–42. Doi 10.7872/crym.v36.iss1.2015.3
Banik N, Yang SB, Kang TB, Lim JH, et al. 2021 – Heparin and its derivatives: Challenges and advances in therapeutic biomolecules. International Journal of Molecular Sciences 22, 10524. Doi 10.3390/ijms221910524
Baytas SN, Linhardt RJ. 2020 – Advances in the preparation and synthesis of heparin and related products. Drug Discovery Today 25, 2095–2109. Doi 10.1016/j.drudis.2020.09.011
Berbari P, Thibodeau A, Germain L, Saint-Cyr M, et al. 1999 – Antiangiogenic effects of the oral administration of liquid cartilage extract in humans. Journal of Surgical Research 87, 108–113. Doi 10.1006/jsre.1999.5698
Bhambri A, Srivastava M, Mahale VG, Mahale S, et al. 2022 – Mushrooms as potential sources of active metabolites and medicines. Frontiers in Microbiology 13, 837266. Doi 10.3389/fmicb.2022.837266
Bhaskar U, Sterner E, Hickey AM, Onishi A, et al. 2012 – Engineering of routes to heparin and related polysaccharides. Applied Microbiology and Biotechnology 93, 1–16. Doi 10.1007/s00253-011-3641-4
Bhaskar U, Yang B, Fu L, Takieddin M, et al. 2013 – Pharmaceutical discovery, development and manufacturing forum. AIChE Annual Meeting: Global Challenges for Engineering a Sustainable Future. San Francisco, CA
Bhilegaonkar KN, Rawat S, Agarwal R. 2014 – Food safety assurance systems: Good animal husbandry practice. Encyclopedia of Food Safety, Elsevier, pp. 168–173. Doi 10.1016/B978-0-12-378612-8.00344-9
Bian C, Wang Z, Shi J. 2020 – Extraction optimization, structural characterization, and anticoagulant activity of acidic polysaccharides from Auricularia auricula-judae. Molecules 25, 710. Doi 10.3390/molecules25030710
Birchfield AS, McIntosh CA. 2020 – Metabolic engineering and synthetic biology of plant natural products – A minireview. Current Plant Biology 24, 100163. Doi 10.1016/j.cpb.2020.100163
Borkertas S, Viskelis J, Viskelis P, Streimikyte P, et al. 2025 – Fungal biomass fermentation: Valorizing the food industry’s waste. Fermentation 11, 351. Doi 10.3390/fermentation11060351
Böttger A, Doxey AC, Hess MW, Pfaller K, et al. 2012 – Horizontal gene transfer contributed to the evolution of extracellular surface structures: the freshwater polyp Hydra is covered by a complex fibrous cuticle containing glycosaminoglycans and proteins of the PPOD and SWT (sweet tooth) families. PloS One 7, e52278. Doi 10.1371/journal.pone.0052278
Bourgeois P, Chales G, Dehais J, Delcambre B, et al. 1998 – Efficacy and tolerability of chondroitin sulfate 1200 mg/day vs chondroitin sulfate 3×400 mg/day vs placebo. Osteoarthritis and Cartilage 6, 25–30. Doi 10.1016/S1063-4584(98)80008-3
Branco S, Schauster A, Liao H, Ruytinx J. 2022 – Mechanisms of stress tolerance and their effects on the ecology and evolution of mycorrhizal fungi. New Phytologist 235, 2158–2175. Doi 10.1111/nph.18308
Brandt F, Cazzaniga A. 2008 – Hyaluronic acid gel fillers in the management of facial aging. Clinical Interventions Aging 3, 153–159. Doi 10.2147/CIA.S2135
Brito R, Costa D, Dias C, Cruz P, et al. 2023 – Chondroitin sulfate supplements for osteoarthritis: A critical review. Cureus 15, e40192. Doi 10.7759/cureus.40192
Bucsi L, Poór G. 1998 – Efficacy and tolerability of oral chondroitin sulfate as a symptomatic slow-acting drug for osteoarthritis (SYSADOA) in the treatment of knee osteoarthritis. Osteoarthritis and Cartilage 6, 31–36. Doi 10.1016/S1063-4584(98)80009-5
Bui T, Wiesolek H, Sumagin R. 2020 – ICAM-1: A master regulator of cellular responses in inflammation, injury resolution, and tumorigenesis. Journal of Leukocyte Biology 108, 787– 799. Doi 10.1002/JLB.2MR0220-549R
Bukhari SNA, Roswandi NL, Waqas M, Habib H, et al. 2018 – Hyaluronic acid, a promising skin rejuvenating biomedicine: A review of recent updates and pre-clinical and clinical investigations on cosmetic and nutricosmetic effects. International Journal of Biological Macromolecules 120, 1682–1695. Doi 10.1016/j.ijbiomac.2018.09.188
Bumbu MG, Niculae M, Ielciu I, Hanganu D, et al. 2024 – Comprehensive review of functional and nutraceutical properties of Craterellus cornucopioides (L.) Pers. Nutrients 16, 831. Doi 10.3390/nu16060831
Burdick JA, Prestwich GD. 2011 – Hyaluronic acid hydrogels for biomedical applications. Advanced Materials 23, H41–H56. Doi 10.1002/adma.201003963
Casale J, Crane JS. 2019 – Biochemistry, Glycosaminoglycans. StatPearls. StatPearls Publishing, Treasure Island, Florida
Cássaro CM, Dietrich CP. 1977 – Distribution of sulfated mucopolysaccharides in invertebrates. Journal of Biological Chemistry 252, 2254–2261. Doi 10.1016/S0021-9258(17)40548-5
Casu B. 1985 – Structure and biological activity of heparin. Advances in Carbohydrate Chemistry and Biochemistry 43, 51–134. Doi 10.1016/S0065-2318(08)60067-0
Casu B, Naggi A, Torri G. 2015 – Re-visiting the structure of heparin. Carbohydrate Research 403, 60–68. Doi 10.1016/j.carres.2014.06.023
Cerana GS, Bos P. 2021 – Process for extracting a hyaluronic acid from a fungus, a hyaluronic acid of plant origin and use thereof. Patent no. WO2021250566A1, 1–45. Available at: https://patents.google.com/patent/WO2021250566A1/en (Accessed on November 18, 2025).
Cesaretti M, Luppi E, Maccari F, Volpi N. 2004 – Isolation and characterization of a heparin with high anticoagulant activity from the clam Tapes phylippinarum: evidence for the presence of a high content of antithrombin III binding site. Glycobiology 14, 1275–1284. DOI 10.1093/glycob/cwh128
Chang-yu C, Yan-yan W, Zhen-hua D, Hong C, et al. 2011 – Optimization of microwave-assisted extraction of chondroitin sulfate from tilapia byproduct by response surface methodology. 2011 International Conference on Consumer Electronics, Communications and Networks (CECNet, Xianning, China, 2011, pp. 1462–1465. Doi 10.1109/CECNET.2011.5768447
Chaplin M. 1981 – The biochemistry of glycoprotein and proteoglycans. Biochemical Education 9, 38. Doi 10.1016/0307-4412(81)90087-X
Cheng F, Gong Q, Yu H, Stephanopoulos G. 2016 – High‐titer biosynthesis of hyaluronic acid by recombinant Corynebacterium glutamicum. Biotechnology Journal 11, 574–584. Doi 10.1002/biot.201500404
Cheng JJ, Chang CC, Chao CH, Lu MK. 2012 – Characterization of fungal sulfated polysaccharides and their synergistic anticancer effects with doxorubicin. Carbohydrate Polymers 90, 134–139. Doi 10.1016/j.carbpol.2012.05.005
Cho H-J, Oh J, Choo M-K, Ha J-I, et al. 2014 – Chondroitin sulfate-capped gold nanoparticles for the oral delivery of insulin. International Journal of Biological Macromolecules 63, 15–20. Doi 10.1016/j.ijbiomac.2013.10.026
Choi E, Oh J, Sung G-H. 2020 – Antithrombotic and antiplatelet effects of Cordyceps militaris. Mycobiology 48, 228–232. Doi 10.1080/12298093.2020.1763115
Choocheep K, Nathip N. 2018 – Detection of a non-animal source of glycosaminoglycans from edible mushrooms in Northern Thailand. Chiang Mai University Journal of Natural Sciences 17, 203–218. Doi 10.12982/CMUJNS.2018.0015
Cimini D, Carlino E, Giovane A, Argenzio O, et al. 2015 – Engineering a branch of the UDP‐precursor biosynthesis pathway enhances the production of capsular polysaccharide in Escherichia coli O5: K4: H4. Biotechnology Journal 10, 1307–1315. Doi 10.1002/biot.201400602
Cobbinah-Sam E, Ekaette I. 2025 – Upcycling eggshell matrix for sustainable production of glycosaminoglycans. Biopolymers 116, e70040. Doi 10.1002/bip.70040
Cohen AT, Dobromirski M, Gurwith MMP. 2014 – Managing pulmonary embolism from presentation to extended treatment. Thrombosis Research 133, 139–148. Doi 10.1016/j.thromres.2013.09.040
Coherent Market Insights. 2024 – Chondrotin sulfate market analysis. Available at: https://www.coherentmarketinsights.com/market-insight/chondroitin-sulfate-market-6000; Accessed on March 25, 2025.
Couto MR, Rodrigues JL, Dias O, Rodrigues LR. 2024 – Saccharomyces cerevisiae as a host for chondroitin production. SynBio 2, 125–141. Doi 10.3390/synbio2020008
Credence Research. 2025 – Glycosaminoglycans (GAGs) market by application (pharmaceuticals, cosmetics, medical devices); by type (heparin, hyaluronic acid, chondroitin sulfate, dermatan sulfate, keratan sulfate); by region- growth, share, opportunities & competitive analysis, 2024. Available at: https://www.credenceresearch.com/report/glycosaminoglycans-market; (Accessed on March 25, 2025).
Crognale S, Russo C, Petruccioli M, D’Annibale A. 2022 – Chitosan production by fungi: Current state of knowledge, future opportunities and constraints. Fermentation 8, 76. Doi 10.3390/fermentation8020076
Davis DAS, Parish CR. 2013 – Heparan sulfate: A ubiquitous glycosaminoglycan with multiple roles in immunity. Frontiers in Immunology 4, 470. Doi 10.3389/fimmu.2013.00470
Deng J-Q, Li Y, Wang Y-J, Cao Y-L, et al. 2024 – Biosynthetic production of anticoagulant heparin polysaccharides through metabolic and sulfotransferases engineering strategies. Nature Communications 15, 3755. Doi 10.1038/s41467-024-48193-5
Dewanjee D, Ghosh S, Khatua S, Rapior S. 2024 – Ganoderma in skin health care: A state-of-the-art review. Ganoderma: Cultivation, Chemistry and Medicinal Applications. Krishnendu Acharya and Somanjana Khatua (eds), CRC Press (Taylor and Francis Group), Boca Raton, pp. 79–101.
Dicker KT, Gurski LA, Pradhan-Bhatt S, Witt RL, et al. 2014 – Hyaluronan: A simple polysaccharide with diverse biological functions. Acta Biomaterialia 10, 1558–1570. Doi 10.1016/j.actbio.2013.12.019
Dovedytis M, Liu ZJ, Bartlett S. 2020 – Hyaluronic acid and its biomedical applications: A review. Engineered Regeneration 1, 102–113. Doi 10.1016/j.engreg.2020.10.001
DrugBank. 2025a – Hyaluronic acid. Available at: https://go.drugbank.com/drugs/DB08818 (Accessed on March 25, 2025).
DrugBank. 2025b – Glucosamine. Available at: https://go.drugbank.com/drugs/DB01296 (Accessed on January 15, 2026).
Du Z, Jia X, Chen J, Zhou S, Chen J, et al. 2019 – Isolation and characterization of a heparin-like compound with potent anticoagulant and fibrinolytic activity from the clam Coelomactra antiquata. Marine Drugs 18, 6. Doi 10.3390/md18010006
DuPont. 2023 Heparin extraction. Available at:https://www.dupont.com/content/dam/dupont/amer/us/en/water-solutions/public/documents/ en/IER-AmberLite-FPA98-Cl-IER-Heparin- Extraction-Br -45-D04430-en.pdf (Accessed on September 22, 2025).
Elkhateeb W, Daba G, Thomas P, Wen TC. 2019 – Medicinal mushrooms as a new source of natural therapeutic bioactive compounds. Egyptian Pharmaceutical Journal 18, 88–101. Doi 10.4103/epj.epj_17_19
Esfandiari H, Loewen NA. 2019 – Effect of glucosamine on intraocular pressure. Handbook of Nutrition, Diet, and the Eye, Elsevier, pp. 333–338.
Espíndola RF, Castro EFS, Santhiago MR, Kara-Junior N. 2012 – A clinical comparison between DisCoVisc and 2% hydroxypropylmethylcellulose in phacoemulsification: a fellow eye study. Clinics 67, 1059–1062. Doi 10.6061/clinics/2012(09)13
Ewald CY. 2021 – Drug screening implicates chondroitin sulfate as a potential longevity pill. Frontiers in Aging 2, 741843. Doi 10.3389/fragi.2021.741843
Fallacara A, Baldini E, Manfredini S, Vertuani S. 2018 – Hyaluronic acid in the third millennium. Polymers 10, 701. Doi 10.3390/polym10070701
Fardellone P, Zaim M, Saurel A-S, Maheu E. 2013 – Comparative efficacy and safety study of two chondroitin sulfate preparations from different origin (avian and bovine) in symptomatic osteoarthritis of the knee. The Open Rheumatology Journal 7, 1–12. Doi 10.2174/1874312901307010001
Fareed J, Jeske W, Ramacciotti E. 2019 – Porcine mucosal heparin shortage crisis! What are the options? Clinical and Applied Thrombosis/Hemostasis 25, 1076029619878786. Doi 10.1177/1076029619878786
Farndale RW, Sayers CA, Barrett AJ. 1982 – A direct spectrophotometric microassay for sulfated glycosaminoglycans in cartilage cultures. Connective Tissue Research 9, 247–248. Doi 10.3109/03008208209160269
Fonseca R, Mourão P. 2006 – Fucosylated chondroitin sulfate as a new oral antithrombotic agent. Thrombosis and Haemostasis 96, 822–829. Doi 10.1160/TH06-06-0304
Fortune Business Insight. 2025 – Heparin market size, share & industry analysis. Available at: https://www.fortunebusinessinsights.com/heparin-market-104447 (Accessed on January 18, 2026).
Fransen M, Agaliotis M, Nairn L, Votrubec M, et al. 2015 – Glucosamine and chondroitin for knee osteoarthritis: a double-blind randomised placebo-controlled clinical trial evaluating single and combination regimens. Annals of the Rheumatic Diseases 74, 851–858. Doi 10.1136/annrheumdis-2013-203954
Fransson F, Kyrk T, Skagerlind M, Stegmayr B. 2013 – Rinsing the extra corporeal circuit with a heparin and albumin solution reduces the need for systemic anticoagulant in hemodialysis. The International Journal of Artificial Organs 36, 725–729. Doi 10.5301/ijao.5000253
Fraser JRE, Laurent TC, Laurent UBG. 1997 – Hyaluronan: its nature, distribution, functions and turnover. Journal of Internal Medicine 242, 27–33. Doi 10.1046/j.1365-2796.1997.00170.x
Frenkel JS. 2014 – The role of hyaluronan in wound healing. International Wound Journal 11, 159–163. Doi 10.1111/j.1742-481X.2012.01057.x
FRWRD Skincare. 2025 – Tremella mushroom: The hyaluronic acid alternative for superior skin hydration. Available at: https://www.frwrdskincare.com/blogs/our-blogs/tremella-mushroom-the-hyaluronic-acid-alternative-for-superior-skinhydration?srsltid=AfmBOorIZKCpygL19pY MNxtUQMfam-eKEVZhfbSxmddjEyJWW1xdzOgy (Accessed on January 18, 2026).
Fu L, Li G, Yang B, Onishi A, et al. 2013 – Structural characterization of pharmaceutical heparins prepared from different animal tissues. Journal of Pharmaceutical Sciences 102, 1447–1457. Doi 10.1002/jps.23501
Fuentes EP, Diaz VB. 1998 – Oligosaccharide mapping of chondroitin sulfate obtained from different animal sources. Acta Farmaceutica Bonaerense 17, 135–142.
Fujisawa S, Takagi K, Yamaguchi-Tanaka M, Sato A, et al. 2024 – Receptor for hyaluronan mediated motility (RHAMM)/hyaluronan axis in breast cancer chemoresistance. Cancers 16, 3600. Doi 10.3390/cancers16213600
Fuster MM, Esko JD. 2005 – The sweet and sour of cancer: Glycans as novel therapeutic targets. Nature Reviews Cancer 5, 526–542. Doi 10.1038/nrc1649
Gabay C, Medinger‐Sadowski C, Gascon D, Kolo F, Finckh A. 2011 – Symptomatic effects of chondroitin 4 and chondroitin 6 sulfate on hand osteoarthritis: A randomized, double‐blind, placebo‐controlled clinical trial at a single center. Arthritis & Rheumatism 63, 3383–3391. Doi 10.1002/art.30574
Galla R, Mulè S, Ferrari S, Parini F, et al. 2025 – Non-animal hyaluronic acid from Tremella fuciformis: A new source with a structure and chemical profile comparable to hyaluronic acid. Foods 14, 1362. Doi 10.3390/foods14081362
Gao Z, Wu C, Wu J, Zhu L, et al. 2022 – Antioxidant and anti-inflammatory properties of an aminoglycan-rich exopolysaccharide from the submerged fermentation of Bacillus thuringiensis. International Journal of Biological Macromolecules 220, 1010–1020. Doi 10.1016/j.ijbiomac.2022.08.116
Garnjanagoonchorn W, Wongekalak L, Engkagul A. 2007 – Determination of chondroitin sulfate from different sources of cartilage. Chemical Engineering and Processing: Process Intensification 46, 465–471. Doi 10.1016/j.cep.2006.05.019
Ghiselli G. 2017 – Drug‐mediated regulation of glycosaminoglycan biosynthesis. Medicinal Research Reviews 37, 1051–1094. Doi 10.1002/med.21429
Glasser DB. 1989 – Protective effects of viscous solutions in phacoemulsification and traumatic lens implantation. Archives of Ophthalmology 107, 1047. Doi 10.1001/archopht.1989.01070020109041
Gomes AM, Kozlowski EO, Pomin VH, de Barros CM, et al. 2010 – Unique extracellular matrix heparan sulfate from the bivalve Nodipecten nodosus (Linnaeus, 1758) safely inhibitsaArterial thrombosis after photochemically induced endothelial lesion. Journal of Biological Chemistry 285, 7312–7323. Doi 10.1074/jbc.M109.091546
González RP, Soares FS, Farias RF, Pessoa C, et al. 2001 – Demonstration of inhibitory effect of oral shark cartilage on basic fibroblast growth factor-induced angiogenesis in the rabbit cornea. Biological & Pharmaceutical Bulletin 24, 151–154. Doi 10.1248/bpb.24.151
Gow NAR, Latge J-P, Munro CA. 2017 – The fungal cell wall: structure, biosynthesis, and function. Microbiology Spectrum 5, FUNK-0035-2016. Doi 10.1128/microbiolspec.FUNK-0035-2016
Grand View Research. 2024 – Hyaluronic acid market size, share & trends analysis report by application (dermal fillers, osteoarthritis, ophthalmic, vesicoureteral reflux), and segment forecasts, 2024 - 2030. Available at: https://www.grandviewresearch.com/industry-analysis/hyaluronic-acid-market#:~:text=Hyaluronic Acid Market Size %26 Trends,contributing to the market growth (Accessed on July 2, 2025).
Grand View Research. 2025 – Mushroom market size, share & trends analysis report by product (button, oyster), by form (fresh, processed), by application (food, pharma), by distribution channel (online, grocery stores), and segment forecasts, 2021 - 2028. Available at: https://www.grandviewresearch.com/industry-analysis/mushroom-market; (Accessed on January 8, 2026).
Gu YJ. 2015 – Extraction technology of crude heparin sodium. Patent no CN105061638A.
Guan Y, Xu X, Liu X, Sheng A, et al. 2016 – Comparison of low-molecular-weight heparins prepared from bovine lung heparin and porcine intestine heparin. Journal of Pharmaceutical Sciences 105, 1843–1850. Doi 10.1016/j.xphs.2016.03.037
Gupta RC, Lall R, Srivastava A, Sinha A. 2019 – Hyaluronic acid: Molecular mechanisms and therapeutic trajectory. Frontiers in Veterinary Science 6, 192. Doi 10.3389/fvets.2019.00192
Hampton JO, Hyndman TH, Allen BL, Fischer B. 2021 – Animal harms and food production: Informing ethical choices. Animals 11, 1225. Doi 10.3390/ani11051225
Hansen SU, Miller GJ, Cliff MJ, Jayson GC, et al. 2015 – Making the longest sugars: a chemical synthesis of heparin-related [4]n oligosaccharides from 16-mer to 40-mer. Chemical Science 6, 6158–6164. Doi 10.1039/C5SC02091C
He W, Zhu Y, Shirke A, Sun X, et al. 2017 – Expression of chondroitin-4-O-sulfotransferase in Escherichia coli and Pichia pastoris. Applied Microbiology and Biotechnology 101, 6919–6928. Doi 10.1007/s00253-017-8411-5
Hedlund KD, Coyne DP, Sanford DM, Huddelson J. 2013 – The heparin recall of 2008. Perfusion 28, 61–65. Doi 10.1177/0267659112462274
Henrotin Y, Marty M, Mobasheri A. 2014 – What is the current status of chondroitin sulfate and glucosamine for the treatment of knee osteoarthritis? Maturitas 78, 184–187. Doi 10.1016/j.maturitas.2014.04.015
Higashi K, Okamoto Y, Mukuno A, Wakai J, et al. 2015 – Functional chondroitin sulfate from Enteroctopus dofleini containing a 3-O-sulfo glucuronic acid residue. Carbohydrate Polymers 134, 557–565. Doi 10.1016/j.carbpol.2015.07.082
Hinneh J, Gilis J, Moore N, Butler L, et al. 2022 – The role of RHAMM in cancer: Exposing novel therapeutic vulnerabilities. Frontiers in Oncology 12, 982231. Doi 10.3389/fonc.2022.982231
Hou Z, Wang W, Wang Y, Chen S, et al. 2025 – Rapid characterization of polysaccharides from marine animals using chemical degradation combined with liquid chromatography mass spectrometry. International Journal of Biological Macromolecules 291, 138535. Doi 10.1016/j.ijbiomac.2024.138535
Hsiao CW, Cheng H, Ghafouri R, Ferko NC, et al. 2023 – Corneal outcomes following cataract surgery using ophthalmic viscosurgical devices composed of chondroitin sulfate-hyaluronic acid: A systematic review and meta-analysis. Clinical Ophthalmology 17, 2083–2096. Doi 10.2147/OPTH.S419863
Huang G, Huang H. 2018 – Application of hyaluronic acid as carriers in drug delivery. Drug Delivery 25, 766–772. Doi 10.1080/10717544.2018.1450910
Hui L, He L. 2012 – Comparison of the moisture retention capacity of Tremella polysaccharides and hyaluronic acid. Journal of Anhui Agricultural Sciences 2012, 13093–13094.
Hussain Z, Thu HE, Katas H, Bukhari SNA. 2017 – Hyaluronic acid-based biomaterials: a versatile and smart approach to tissue regeneration and treating traumatic, surgical, and chronic wounds. Polymer Reviews 57, 594–630. Doi 10.1080/15583724.2017.1315433
Hyde KD, Bahkali AH, Moslem MA. 2010 – Fungi—an unusual source for cosmetics. Fungal Diversity 43, 1–9. Doi 10.1007/s13225-010-0043-3
Hyde KD, Baldrian P, Chen Y, Chethana KWT, et al. 2024 – Current trends, limitations and future research in the fungi? Fungal Diversity 125, 1–71. Doi 10.1007/s13225-023-00532-5
Hyde KD, Xu J, Rapior S, Jeewon R, et al. 2019 – The amazing potential of fungi: 50 ways we can exploit fungi industrially. Fungal Diversity 97, 1–136. Doi 10.1007/s13225-019-00430-9
Hynes W. 2000 – Hyaluronidases of Gram-positive bacteria. FEMS Microbiology Letters 183, 201–207. Doi 10.1016/S0378-1097(99)00669-2
Iaconisi GN, Lunetti P, Gallo N, Cappello AR, et al. 2023 – Hyaluronic acid: a powerful biomolecule with wide-ranging applications-a comprehensive review. International Journal of Molecular Sciences 24, 10296. Doi 10.3390/ijms241210296
INCIDecoder. 2024 – Sodium chondroitin sulfate. Available at: https://incidecoder.com/ingredients/sodium-chondroitin-sulfate#:~:text=As for skincare%2C Sodium Chondroitin,affinity for the skin surface (Accessed on January 8, 2026).
Iozzo RV, Schaefer L. 2015 – Proteoglycan form and function: A comprehensive nomenclature of proteoglycans. Matrix Biology 42, 11–55. Doi 10.1016/j.matbio.2015.02.003
Jaques LB, Waters ET, Charles AF. 1942 – A comparison of the heparins of various mammalian species. Journal of Biological Chemistry 144, 229–235. Doi 10.1016/S0021-9258(18)72576-3
Jeong E, Shim WY, Kim JH. 2014 – Metabolic engineering of Pichia pastoris for production of hyaluronic acid with high molecular weight. Journal of Biotechnology 185, 28–36. Doi 10.1016/j.jbiotec.2014.05.018
Jeong S-C, Cho S-P, Yang B-K, Gu Y-A, et al. 2004 – Production of an anti-complement exo-polymer produced by Auricularia auricula-judae in submerged culture. Biotechnology Letters 26, 923–927. Doi 10.1023/B:bile.0000025904.21519.3a
Jia Y, Zhu J, Chen X, Tang D, et al. 2013 – Metabolic engineering of Bacillus subtilis for the efficient biosynthesis of uniform hyaluronic acid with controlled molecular weights. Bioresource Technology 132, 427–431. Doi 10.1016/j.biortech.2012.12.150
Jin P, Kang Z, Yuan P, Du G, et al. 2016 – Production of specific-molecular-weight hyaluronan by metabolically engineered Bacillus subtilis 168. Metabolic Engineering 35, 21–30. Doi 10.1016/j.ymben.2016.01.008
Jin W, Zhang F, Linhardt RJ. 2021a – Bioengineered production of glycosaminoglycans and their analogues. Systems Microbiology and Biomanufacturing 1, 123–130. Doi 10.1007/s43393-020-00011-x
Jin X, Zhang W, Wang Y, Sheng J, et al. 2021b – Biosynthesis of non-animal chondroitin sulfate from methanol using genetically engineered Pichia pastoris. Green Chemistry 23, 4365–4374. Doi 10.1039/D1GC00260K
Jong A, Wu C-H, Chen H-M, Luo F, et al. 2007 – Identification and characterization of CPS1 as a Hyaluronic Acid synthase contributing to the pathogenesis of Cryptococcus neoformans infection. Eukaryotic Cell 6, 1486–1496. Doi 10.1128/EC.00120-07
Juncan AM, Moisă DG, Santini A, Morgovan C, et al. 2021 – Advantages of hyaluronic acid and its combination with other bioactive ingredients in cosmeceuticals. Molecules 26, 4429. Doi 10.3390/molecules26154429
Kahan A, Uebelhart D, De Vathaire F, Delmas PD, et al. 2009 – Long-term effects of chondroitins 4 and 6 sulfate on knee osteoarthritis: The study on osteoarthritis progression prevention, a two-year, randomized, double‐blind, placebo-controlled trial. Arthritis & Rheumatism 60, 524–533. Doi 10.1002/art.24255
Kalantarmahdavi M, Salari A, Pasdar Z, Amiryousefi MR. 2022 – Edible hyaluronic acid-rich burger separator discs prepared from slaughterhouse waste. Food Science & Nutrition 10, 3515–3526. Doi 10.1002/fsn3.2954
Kang DY, Kim W-S, Heo IS, Park YH, et al. 2010 – Extraction of hyaluronic acid (HA) from rooster comb and characterization using flow field-flow fractionation (FlFFF) coupled with multiangle light scattering (MALS). Journal of Separation Science 33, 3530–3536. Doi 10.1002/jssc.201000478
Kang X, Kirui A, Muszyński A, Widanage MCD, et al. 2018a – Molecular architecture of fungal cell walls revealed by solid-state NMR. Nature Communications 9, 2747. Doi 10.1038/s41467-018-05199-0
Kang Z, Zhou Z, Wang Y, Huang H, et al. 2018b – Bio-based strategies for producing glycosaminoglycans and their oligosaccharides. Trends in Biotechnology 36, 806–818. Doi 10.1016/j.tibtech.2018.03.010
Kantor ED, Zhang X, Wu K, Signorello LB, et al. 2016 – Use of glucosamine and chondroitin supplements in relation to risk of colorectal cancer: Results from the nurses’ health study and health professionals follow-up study. International Journal of Cancer 139, 1949–1957. Doi 10.1002/ijc.30250
Kappes A, Tozooneyi T, Shakil G, Railey AF, et al. 2023 – Livestock health and disease economics: a scoping review of selected literature. Frontiers in Veterinary Science 10, 1168649. Doi 10.3389/fvets.2023.1168649
Karunarathna SC, Patabendige NM, Hapuarachchi KK, Promputtha I. 2025 – Exploring the health benefits of Ganoderma: antimicrobial properties and mechanisms of action. Frontiers in Cellular and Infection Microbiology 15, 1535246. Doi 10.3389/fcimb.2025.1535246
Kaul A, Short WD, Keswani SG, Wang X. 2021 – Immunologic roles of hyaluronan in dermal wound healing. Biomolecules 11, 1234. Doi 10.3390/biom11081234
Khanna JM, Malone MH, Euler KL, Brady LR. 1965 – Atromentin anticoagulant from Hydnellum diabolus. Journal of Pharmaceutical Sciences 54, 1016–1020. Doi 10.1002/jps.2600540714
Kiani AK, Pheby D, Henehan G, Brown R, et al. 2022 – Ethical considerations regarding animal experimentation. Journal of Preventive Medicine and Hygiene 63, E255–E266. Doi 10.15167/2421-4248/jpmh2022.63.2S3.2768
Konstantinides S V., Torbicki A, Agnelli G, Danchin N, et al. 2014 – 2014 ESC Guidelines on the diagnosis and management of acute pulmonary embolism. European Heart Journal 35, 3033–3080. Doi 10.1093/eurheartj/ehu283
Korner JD. 2022 – Methods of production of heparosan, heparan sulfate and heparin in yeast (Patent No. WO2023110854A1). Genhtis Fine Chemicals GmbH, Germany
Kouta A, Jeske W, Hoppensteadt D, Iqbal O, et al. 2019 – Comparative pharmacological profiles of various bovine, ovine, and porcine heparins. Clinical and Applied Thrombosis/Hemostasis 25, 1076029619889406. Doi 10.1177/1076029619889406
Kragh M, Binderup L, Hjarnaa PJV, Bramm E, et al. 2005 – Non-anti-coagulant heparin inhibits metastasis but not primary tumor growth. Oncology Reports 14, 99-104.
Krichen F, Bougatef H, Capitani F, Ben Amor I, et al. 2018 – Purification and structural elucidation of chondroitin sulfate/dermatan sulfate from Atlantic bluefin tuna (Thunnus thynnus) skins and their anticoagulant and ACE inhibitory activities. RSC Advances 8, 37965–37975. Doi 10.1039/C8RA06704J
Krishnan S, Chakraborty K, Dhara S. 2025 – Sulphated glycosaminoglycan isolated from the edible slipper oyster Magallana bilineata (Röding, 1798) attenuates inflammatory cytokines on lipopolysaccharide-prompted macrophages. Natural Product Research 39, 6492–6503. Doi 10.1080/14786419.2024.2377311
Krüger-Genge A, Blocki A, Franke R, Jung F. 2019 – Vascular endothelial cell biology: An update. International Journal of Molecular Sciences 20, 4411. Doi 10.3390/ijms20184411
Lapierre F, Holme K, Lam L, Tressler RJ, et al. 1996 – Chemical modifications of heparin that diminish its anticoagulant but preserve its heparanase-inhibitory, angiostatic, anti-tumor and anti-metastatic properties. Glycobiology 6, 355–366. Doi 10.1093/glycob/6.3.355
Latimer A. 2025 – Natural hyaluronic acid alternative (from Tremella mushroom). Available at: https://www.gcimagazine.com/ingredients/regulatory/news/21869055/natural-hyaluronic-acid-alternative-from-tremella-mushroom-from-alpha-environmental (Accessed on January 18, 2026).
Lee DY, Lee SY, Kang HJ, Park Y, et al. 2020 – Development of effective heparin extraction method from pig by-products and analysis of their bioavailability. Journal of Animal Science and Technology 62, 933–947. Doi 10.5187/jast.2020.62.6.933
Li J, Kusche-Gullberg M. 2016 – Heparan sulfate: Biosynthesis, structure, and function. International Review of Cell and Molecular Biology 325, 215–273. Doi 10.1016/bs.ircmb.2016.02.009
Li L-D, Mao P-W, Shao K-D, Bai X-H, Zhou X-W. 2019a – Ganoderma proteins and their potential applications in cosmetics. Applied Microbiology and Biotechnology 103, 9239–9250. Doi 10.1007/s00253-019-10171-z
Li S, Ma F, Pang X, Tang B, Lin L. 2019b – Synthesis of chondroitin sulfate magnesium for osteoarthritis treatment. Carbohydrate Polymers 212, 387–394. Doi 10.1016/j.carbpol.2019.02.061
Li X, Qian K, Han W. 2021 – Prediction of hyaluronic acid target on sucrase-isomaltase (SI) with reverse docking and molecular dynamics simulations for inhibitors binding to SI. PLoS One 16, e0255351. Doi 10.1371/journal.pone.0255351
Lim S, Park J, Shim MK, Um W, et al. 2019 – Recent advances and challenges of repurposing nanoparticle-based drug delivery systems to enhance cancer immunotherapy. Theranostics 9, 7906–7923. Doi 10.7150/thno.38425
Linhardt RJ. 2003 – 2003 Claude S. Hudson Award address in carbohydrate chemistry. Heparin: structure and activity. Journal of Medicinal Chemistry 46, 2551–2564. Doi 10.1021/jm030176m
Linhardt RJ, Toida T. 2004 – Role of glycosaminoglycans in cellular communication. Accounts of Chemical Research 37, 431–438. Doi 10.1021/ar030138x
Liu H, He L. 2012 – Comparison of moisturizing properties of polysaccharides from Tremella fuciformis and hyaluronic acid. Journal of Anhui Agricultural Sciences 40, 13093–13094.
Liu J, Wang Y, Li Z, Ren Y, et al. 2018 – Efficient production of high-molecular-weight hyaluronic acid with a two-stage fermentation. RSC Advances 8, 36167–36171. Doi 10.1039/c8ra07349j
Liu K, Guo L, Chen X, Liu L, et al. 2023 – Microbial synthesis of glycosaminoglycans and their oligosaccharides. Trends in Microbiology 31, 369–383. Doi 10.1016/j.tim.2022.11.003
Liu L, Liu Y, Li J, Du G, Chen J. 2011 – Microbial production of hyaluronic acid: current state, challenges, and perspectives. Microbial Cell Factories 10, 99. Doi 10.1186/1475-2859-10-99
Liu S, Xiang Y, Xu C, Sun J, et al. 2025 – Systematic preparation of animal-derived glycosaminoglycans: Research progress and industrial significance. Food Chemistry 464, 141565. Doi 10.1016/j.foodchem.2024.141565
Llanaj X, Törős G, Hajdú P, Abdalla N, et al. 2023 – Biotechnological applications of mushrooms under the water-energy-food nexus: Crucial aspects and prospects from farm to pharmacy. Foods 12, 2671. Doi 10.3390/foods12142671
Luo X, Fosmire G, Leach R. 2002 – Chicken keel cartilage as a source of chondroitin sulfate. Poultry Science 81, 1086–1089. Doi 10.1093/ps/81.7.1086
Luppi E, Cesaretti M, Volpi N. 2005 – Purification and characterization of heparin from the Italian clam Callista chione. Biomacromolecules 6, 1672–1678. Doi 10.1021/bm049196b
Ma X, Yang M, He Y, Zhai C, et al. 2021 – A review on the production, structure, bioactivities and applications of Tremella polysaccharides. International Journal of Immunopathology and Pharmacology 35, 20587384211000541. Doi 10.1177/20587384211000541
Mahmoud Abd-Elaty N, Elprince M, El-salam MA. 2007 – Oral Abstract Session : Treatment of asthma; Oral communication. World Allergy Organization Journal 129, S42–S43. Doi 10.1097/01.WOX.0000301200.96305.00
Marinho A, Nunes C, Reis S. 2021 – Hyaluronic acid: A key ingredient in the therapy of inflammation. Biomolecules 11, 1518. Doi 10.3390/biom11101518
Martin RJ, Chu HW, Honour JM, Harbeck RJ. 2001 – Airway inflammation and bronchial hyperresponsiveness after Mycoplasma pneumoniae Infection in a murine model. American Journal of Respiratory Cell and Molecular Biology 24, 577–582. Doi 10.1165/ajrcmb.24.5.4315
Martínez-Duncker I, Díaz-Jímenez DF, Mora-Montes HM. 2014 – Comparative analysis of protein glycosylation pathways in humans and the fungal pathogen Candida albicans. International Journal of Microbiology 2014, 267497. Doi 10.1155/2014/267497
Martins-Santana L, Nora LC, Sanches-Medeiros A, Lovate GL, Cassiano MHA, Silva-Rocha R. 2018 – Systems and synthetic biology approaches to engineer fungi for fine chemical production. Frontiers in Bioengineering and Biotechnology 6, 117. Doi 10.3389/fbioe.2018.00117
Matarasso SL, Carruthers JD, Jewell ML. 2006 – Consensus recommendations for soft-tissue augmentation with nonanimal stabilized hyaluronic acid (Restylane). Plastic and Reconstructive Surgery 117, 3S–34S. Doi 10.1097/01.prs.0000204759.76865.39
Matsuzaka Y, Yashiro R. 2024 – Classification and molecular functions of heparan sulfate proteoglycans and their molecular mechanisms with the receptor. Biologics 4(2), 105–129. Doi 10.3390/biologics4020008
Mazières B, Hucher M, Zaïm M, Garnero P. 2007 – Effect of chondroitin sulphate in symptomatic knee osteoarthritis: a multicentre, randomised, double-blind, placebo-controlled study. Annals of the Rheumatic Diseases 66, 639–645. Doi 10.1136/ard.2006.059899
Medeiros LHC, Vasconcelos BMF, Silva MB, Souza-Junior AA, et al. 2021 – Chondroitin sulfate from fish waste exhibits strong intracellular antioxidant potential. Brazilian Journal of Medical and Biological Research 54, e10730. Doi 10.1590/1414-431X2020E10730
Meher MK, Naidu G, Mishra A, Poluri KM. 2024 – A review on multifaceted biomedical applications of heparin nanocomposites: Progress and prospects. International Journal of Biological Macromolecules 260, 129379. Doi 10.1016/j.ijbiomac.2024.129379
Mehta R. 2020 – Anti-thrombotic attribute of different type of mushrooms. International Journal of Current Microbiology and Applied Sciences 9, 815–820. Doi 10.20546/ijcmas.2020.908.087
Menaa F, Meena A, Meena B. 2011 – Hyaluronic acid and derivatives for tissue engineering. Journal of Biotechnology & Biomaterials S3, 001. Doi 10.4172/2155-952X.S3-001
Metoree. 2025 – 13 Chondroitin Sulfate Manufacturers in 2025. Available at: https://us.metoree.com/categories/6262/#manufacturers (Accessed on January 18, 2026).
Middeldorp S. 2008 – Heparin: From animal organ extract to designer drug. Thrombosis Research 122, 753–762. Doi 10.1016/j.thromres.2007.07.004
Min D, Park S, Kim H, Lee SH, et al. 2020 – Potential anti‐ageing effect of chondroitin sulphate through skin regeneration. International Journal of Cosmetic Science 42, 520–527. Doi 10.1111/ics.12645
Misra S, Hascall V, Markwald R, Ghatak S. 2015 – Interactions between hyaluronan and its receptors (CD44, RHAMM) regulate the activities of inflammation and cancer. Frontiers in Immunology 6, 201. Doi 10.3389/fimmu.2015.00201
Miyazaki T, Nishijima M. 1981 – A novel glycosaminoglycan from the fungus Omphalia lapidescence. Carbohydrate Research 96, 105–111. Doi 10.1016/S0008-6215(00)84700-X
Mol PC, Wessels JGH. 1987 – Linkages between glucosaminoglycan and glucan determine alkali-insolubility of the glucan in walls of Saccharomyces cerevisiae. FEMS Microbiology Letters 41, 95–99. Doi 10.1111/j.1574-6968.1987.tb02148.x
Möller I, Gharbi M, Martinez Serrano H, Herrero Barbero M, et al. 2016 – Effect of chondroitin sulfate on soluble biomarkers of osteoarthritis: a method to analyze and interpret the results from an open-label trial in unilateral knee osteoarthritis patients. BMC Musculoskeletal Disorders 17, 416. Doi 10.1186/s12891-016-1268-4
Monfort J, Pujol J, Contreras-Rodríguez O, Llorente-Onaindia J, López-Solà M, et al. 2017 – Effects of chondroitin sulfate on brain response to painful stimulation in knee osteoarthritis patients. A randomized, double-blind, placebo-controlled functional magnetic resonance imaging study. Medicina Clínica (English Edition) 148, 539–547. Doi 10.1016/j.medcle.2017.05.012
Moreira TD, Martins VB, da Silva Júnior AH, Sayer C, et al. 2024 – New insights into biomaterials for wound dressings and care: Challenges and trends. Progress in Organic Coatings 187, 108118. Doi 10.1016/j.porgcoat.2023.108118
Moschos MM, Chatziralli IP, Sergentanis TN. 2011 – Viscoat versus Visthesia during phacoemulsification cataract surgery: corneal and foveal changes. BMC Ophthalmology 11, 9. Doi 10.1186/1471-2415-11-9
Muran AC, Schaffler BC, Wong A, Neufeld E, et al. 2023 – Effect of increasing hyaluronic acid content in collagen scaffolds on the maintenance of chondrogenic phenotype in chondrocytes and mesenchymal stem cells. Journal of Cartilage & Joint Preservation 3, 100099. Doi 10.1016/j.jcjp.2023.100099
MyAlcon Professionals. 2025 – Ophthalmic Viscosurgical Devices (OVDs). Available at: https://www.myalcon.com/professional/cataract-surgery/disposables/ovds/#link2 (Accessed on January 18, 2026).
Nagasawa K, Uchiyama H. 1984 – Anticoagulant properties of heparin preparations from different animal sources with equivalent high affinity for antithrombin III. The Journal of Biochemistry 95, 619–626. Doi 10.1093/oxfordjournals.jbchem.a134650
Naggi A, Casu B, Perez M, Torri G, et al. 2005 – Modulation of the heparanase-inhibiting activity of heparin through selective desulfation, graded N-acetylation, and glycol splitting. Journal of Biological Chemistry 280, 12103–12113. Doi 10.1074/jbc.M414217200
Nahain A Al, Ignjatovic V, Monagle P, Tsanaktsidis J, et al. 2018 – Heparin mimetics with anticoagulant activity. Medicinal Research Reviews 38, 1582–1613. Doi 10.1002/med.21489
Narayanan K, Sivagurunathan N, Subrahmanyam VM, Venkata Rao J. 2017 – Fungal chondroitinase: Production and prospects for therapeutic application. In: Recent advances in Applied Microbiology, Springer Singapore, Singapore, pp. 275–290.
Necas J, Bartosikova L, Brauner P, Kolar J. 2008 – Hyaluronic acid (hyaluronan): a review. Veterinární Medicína 53, 397–411. Doi 10.17221/1930-VETMED
Niego AGT, Rapior S, Thongklang N, Raspé O, et al. 2023a – Reviewing the contributions of macrofungi to forest ecosystem processes and services. Fungal Biology Reviews 44, 100294. Doi 10.1016/j.fbr.2022.11.002
Niego AGT, Lambert C, Mortimer P, Thongklang N, et al. 2023b – The contribution of fungi to the global economy. Fungal Diversity 121, 95–137. Doi 10.1007/s13225-023-00520-9
Nikitovic D, Pérez S. 2021 – Preface for the special issue on the exploration of the multifaceted roles of glycosaminoglycans: GAGs. Biomolecules 11, 1630. Doi 10.3390/biom11111630
Nurunnabi M, Khatun Z, Moon W-C, Lee G, et al. 2012 – Heparin based nanoparticles for cancer targeting and noninvasive imaging. Quantitative Imaging in Medicine and Surgery 2, 219–226. Doi 10.3978/j.issn.2223-4292.2012.09.01
Oduah E, Linhardt R, Sharfstein S. 2016 – Heparin: Past, present, and future. Pharmaceuticals 9, 38. Doi 10.3390/ph9030038
Oertli B, Beck-Schimmer B, Fan X, Wüthrich RP. 1998 – Mechanisms of hyaluronan-induced up-regulation of ICAM-1 and VCAM-1 expression by murine kidney tubular epithelial cells: hyaluronan triggers cell adhesion molecule expression through a mechanism involving activation of nuclear factor-B and activating protein-1. Journal of Immunology 161, 3431–3437. Doi 10.4049/jimmunol.161.7.3431
Oliveira SNMCG, Bezerra FF, Pereira MS, Tovar AMF, Aquino RS, Mourão PAS. 2025 – Global use of bovine heparin: Challenges and opportunities. Proteoglycan Research 3, e70025. Doi 10.1002/pgr2.70025
Oliveira SNMCG, Tovar AMF, Bezerra FF, Piquet AA, et al. 2022 – Anticoagulant activity of heparins from different animal sources are driven by a synergistic combination of physical-chemical factors. TH Open 6, e309–e322. Doi 10.1055/a-1946-0325
Onishi A, St Ange K, Dordick JS, Linhardt RJ. 2016 – Heparin and anticoagulation. Frontiers in Bioscience (Landmark ed) 21, 1372–1392. Doi 10.2741/4462
Ordiales H, Alcalde I, Vázquez F, Merayo-Lloves J, et al. 2022a – Cell surface glycosaminoglycans as receptors for adhesion of Candida spp. to corneal cells. Polish Journal of Microbiology 71, 55–62. Doi 10.33073/pjm-2022-008
Ordiales H, Vázquez-López F, Pevida M, Vázquez-Losada B, et al. 2022b – La unión de Candida albicans y Malassezia spp. a células de piel promueve cambios de expresión en los genes responsables de la síntesis de las cadenas de heparán y condroitín sulfato. Actas Dermo-Sifiliográficas 113, 712–716. Doi 10.1016/j.ad.2021.11.010
Pai AA, Chakraborty K, Dhara S, Raj A, et al. 2024 – Therapeutic potential of sulfated glycosaminoglycan from seafood Asian green mussel (Perna viridis): Insights from an in vivo study. Food Bioscience 61, 104837. Doi 10.1016/j.fbio.2024.104837
Palmieri B, Merighi A, Corbascio D, Rottigni V, et al. 2013 – Fixed combination of hyaluronic acid and chondroitin-sulphate oral formulation in a randomized double blind, placebo controlled study for the treatment of symptoms in patients with non-erosive gastroesophageal reflux. European Review for Medical and Pharmacological Sciences 17, 3272–3278.
Paluck SJ, Nguyen TH, Maynard HD. 2016 – Heparin-mimicking polymers: Synthesis and biological applications. Biomacromolecules 17, 3417–3440. Doi 10.1021/acs.biomac.6b01147
Pan NC, Biz G, Baldo C, Celligoi MAPC. 2019 – Factorial design in fermentation medium development for hyaluronic acid production by Streptococcus zooepidemicus. Acta Scientiarum. Technology 42, e42729. Doi 10.4025/actascitechnol.v42i1.42729
Pang H, Lu H, Liu P, Zhang YT, et al. 2024 – A chondroitin sulfate purified from shark cartilage and bovine serum albumin interaction activity. International Journal of Biological Macromolecules 260, 129499. Doi 10.1016/j.ijbiomac.2024.129499
Papaconstantinou D, Karmiris T, Diagourtas A, Koutsandrea C, et al. 2014 – Clinical trial evaluating Viscoat and Visthesia ophthalmic viscosurgical devices in corneal endothelial loss after cataract extraction and intraocular lens implantation. Cutaneous and Ocular Toxicology 33, 173–180. Doi 10.3109/15569527.2013.845835
Parapouli M, Vasileiadis A, Afendra A-S, Hatziloukas E. 2020 – Saccharomyces cerevisiae and its industrial applications. AIMS Microbiology 6, 1–31. Doi 10.3934/microbiol.2020001
Park J, Choi Y, Chang H, Um W, et al. 2019 – Alliance with EPR effect: Combined strategies to improve the EPR effect in the tumor microenvironment. Theranostics 9, 8073–8090. Doi 10.7150/thno.37198
Park JW, Jeon OC, Kim SK, Al-Hilal TA, et al. 2010 – High antiangiogenic and low anticoagulant efficacy of orally active low molecular weight heparin derivatives. Journal of Controlled Release 148, 317–326. Doi 10.1016/j.jconrel.2010.09.014
Park K, Lee GY, Kim Y-S, Yu M, et al. 2006 – Heparin–deoxycholic acid chemical conjugate as an anticancer drug carrier and its antitumor activity. Journal of Controlled Release 114, 300–306. Doi 10.1016/j.jconrel.2006.05.017
Park PW. 2018 – Isolation and functional analysis of syndecans. Methods in Cell Biology 143, 317–333. Doi 10.1016/bs.mcb.2017.08.019
Patel PK, Free SJ. 2019 – The genetics and biochemistry of cell wall structure and synthesis in Neurospora crassa, a model filamentous fungus. Frontiers in Microbiology 10, 2294. Doi 10.3389/fmicb.2019.02294
Paterska M, Czerny B, Cielecka-Piontek J. 2024 – Macrofungal extracts as a source of bioactive compounds for cosmetical anti-aging therapy: a comprehensive review. Nutrients 16, 2810. Doi 10.3390/nu16162810
Pavão MSG, Mourão PAS, Mulloy B, Tollefsen DM. 1995 – A unique dermatan sulfate-like glycosaminoglycan from ascidian: Its structure and the effect of its unusual sulfation pattern on anticoagulant activity. Journal of Biological Chemistry 270, 31027–31036. Doi 10.1074/jbc.270.52.31027
Pérez S, Bonnardel F, Lisacek F, Imberty A, et al. 2020 – GAG-DB, the new interface of the three-dimensional landscape of glycosaminoglycans. Biomolecules 10, 1660. Doi 10.3390/biom10121660
Perez S, Makshakova O, Angulo J, Bedini E, et al. 2023 – Glycosaminoglycans: What remains to be deciphered? Journal of the American Chemical Society Au 3, 628–656. Doi 10.1021/jacsau.2c00569
Person P, Mathews MB. 1967 – Endoskeletal cartilage in a marine polychaete, Eudistylia polymorpha. The Biological Bulletin 132, 244 –252. Doi 10.2307/1539892
De Pourcq K, De Schutter K, Callewaert N. 2010 – Engineering of glycosylation in yeast and other fungi: current state and perspectives. Applied Microbiology and Biotechnology 87, 1617–1631. Doi 10.1007/s00253-010-2721-1
Prydz K. 2015 – Determinants of glycosaminoglycan (GAG) structure. Biomolecules 5, 2003–2022. Doi 10.3390/biom5032003
PubChem. 2025 – Atromentin. Available at: https://pubchem.ncbi.nlm.nih.gov/compound/Atromentin (Accessed on January 18, 2026).
Qiu XL, Fan ZR, Liu YY, Wang DF, et al. 2021 – Preparation and evaluation of a self-nanoemulsifying drug delivery system loaded with heparin phospholipid complex. International Journal of Molecular Sciences 22, 4077. Doi 10.3390/ijms22084077
Railhac JJ, Zaim M, Saurel A-S, Vial J, Fournie B. 2012 – Effect of 12 months treatment with chondroitin sulfate on cartilage volume in knee osteoarthritis patients: a randomized, double-blind, placebo-controlled pilot study using MRI. Clinical Rheumatology 31, 1347–1357. Doi 10.1007/s10067-012-2022-4
Rajora AK, Ravishankar D, Zhang H, Rosenholm JM. 2020 – Recent advances and impact of chemotherapeutic and antiangiogenic nanoformulations for combination cancer therapy. Pharmaceutics 12, 592. Doi 10.3390/pharmaceutics12060592
Ravikumar M, Smith RAA, Nurcombe V, Cool SM. 2020 – Heparan sulfate proteoglycans: Key mediators of stem cell function. Frontiers in Cell and Developmental Biology 8, 581213. Doi 10.3389/fcell.2020.581213
Ray P, Kundu S, Paul D. 2024 – Exploring the therapeutic properties of chinese mushrooms with a focus on their anti-cancer effects: A systemic review. Pharmacological Research - Modern Chinese Medicine 11, 100433. Doi 10.1016/j.prmcm.2024.100433
Reginster JY, Veronese N. 2021 – Highly purified chondroitin sulfate: a literature review on clinical efficacy and pharmacoeconomic aspects in osteoarthritis treatment. Aging Clinical and Experimental Research 33, 37–47. Doi 10.1007/s40520-020-01643-8
Rodriguez-Marquez CD, Arteaga-Marin S, Rivas-Sánchez A, Autrique-Hernández R, et al. 2022 – A review on current strategies for extraction and purification of hyaluronic acid. International Journal of Molecular Sciences 23, 6038. Doi 10.3390/ijms23116038
Roughley PJ, Mason RM. 1975 – Proteins and hyaluronic acid associated with proteoglycans extracted from bovine nasal cartilage with low-ionic-strength salt solution. Biochemical Society Transactions 3, 140–142. Doi 10.1042/bst0030140
Saha SK, Zhu Y, Murray P, Madden L. 2024 – Future proofing of chondroitin sulphate production: Importance of sustainability and quality for the end-applications. International Journal of Biological Macromolecules 267, 131577. Doi 10.1016/j.ijbiomac.2024.131577
Sahu B, Sharma DD, Jayakumar GC, Madhan B, et al. 2023 – A review on an imperative by-product: glycosaminoglycans - a holistic approach. Carbohydrate Polymer Technologies and Applications 5, 100275. Doi 10.1016/j.carpta.2022.100275
Salih ARC, Farooqi HMU, Amin H, Karn PR, et al. 2024 – Hyaluronic acid: comprehensive review of a multifunctional biopolymer. Future Journal of Pharmaceutical Sciences 10, 63. Doi 10.1186/s43094-024-00636-y
Sandargo B, Chepkirui C, Cheng T, Chaverra-Muñoz L, et al. 2019 – Biological and chemical diversity go hand in hand: Basidiomycota as source of new pharmaceuticals and agrochemicals. Biotechnology Advances 37, 107344. Doi 10.1016/j.biotechadv.2019.01.011
Sangthong S, Pintathong P, Pongsua P, Jirarat A, et al. 2022 – Polysaccharides from Volvariella volvacea mushroom: Extraction, biological activities and cosmetic efficacy. Journal of Fungi 8, 572. Doi 10.3390/jof8060572
Saribaş AS, Mobasseri A, Pristatsky P, Chen X, et al. 2004 – Production of N-sulfated polysaccharides using yeast-expressed N-deacetylase/N-sulfotransferase-1 (NDST-1). Glycobiology 14, 1217–1228. Doi 10.1093/glycob/cwh129
Sarrazin S, Lamanna WC, Esko JD. 2011 – Heparan sulfate proteoglycans. Cold Spring Harbor Perspectives in Biology 3(7), a004952. Doi 10.1101/cshperspect.a004952
Schneider H. 2012 – Symptom-modifying effect of chondroitin sulfate in knee osteoarthritis: A meta-analysis of randomized placebo-controlled trials performed with Structum®. The Open Rheumatology Journal 6, 183–189. Doi 10.2174/1874312901206010183
Serra M, Casas A, Toubarro D, Barros AN, et al. 2023 – Microbial hyaluronic acid production: A review. Molecules 28, 2084. Doi 10.3390/molecules28052084
Shan G, Meihe L, Minchao K, Rui Z, et al. 2022 – Identification and validation of Osteopontin and receptor for hyaluronic acid-mediated motility (RHAMM, CD168) for potential immunotherapeutic significance of in lung squamous cell carcinoma. International Immunopharmacology 107, 108715. Doi 10.1016/j.intimp.2022.108715
Sharma R, Kuche K, Thakor P, Bhavana V, et al. 2022 – Chondroitin Sulfate: Emerging biomaterial for biopharmaceutical purpose and tissue engineering. Carbohydrate Polymers 286, 119305. Doi 10.1016/j.carbpol.2022.119305
Shen Q, Guo Y, Wang K, Zhang C, et al. 2023 – A review of chondroitin sulfate’s preparation, properties, functions, and applications. Molecules 28, 7093. Doi 10.3390/molecules28207093
Shen Q, Zhang C, Mo H, Zhang H, Qin X, Li J, Zhang Z, Richel A. 2021 – Fabrication of chondroitin sulfate calcium complex and its chondrocyte proliferation in vitro. Carbohydrate Polymers 254, 117282. Doi 10.1016/j.carbpol.2020.117282
Sheng J, Ling P, Wang F. 2015 – Constructing a recombinant hyaluronic acid biosynthesis operon and producing food-grade hyaluronic acid in Lactococcus lactis. Journal of Industrial Microbiology and Biotechnology 42, 197–206. Doi 10.1007/s10295-014-1555-8
Shi D, Sheng A, Chi L. 2021 – Glycosaminoglycan-protein interactions and their roles in human disease. Frontiers in Molecular Biosciences 8, 639666. Doi 10.3389/fmolb.2021.639666
Shikina E V, Kovalevsky RA, Shirkovskaya AI, Toukach PV. 2022 – Prospective bacterial and fungal sources of hyaluronic acid: A review. Computational and Structural Biotechnology Journal 20, 6214–6236. Doi 10.1016/j.csbj.2022.11.013
Shimizu MT, Jorge AOC, Unterkircher CS, Fantinato V, et al. 1995 – Hyaluronidase and chondroitin sulphatase production by different species of Candida. Medical Mycology 33, 27–31. Doi 10.1080/02681219580000061
Shriver Z, Capila I, Venkataraman G, Sasisekharan R. 2011 – Heparin and heparan sulfate: Analyzing structure and microheterogeneity. Handbook of Experimental Pharmacology 207, 159–176. Doi 10.1007/978-3-642-23056-1_8
Siddiqui SA, Erol Z, Rugji J, Taşçı F, et al. 2023 – An overview of fermentation in the food industry - looking back from a new perspective. Bioresources and Bioprocessing 10, 85. Doi 10.1186/s40643-023-00702-y
Sim JS, Im AR, Cho SM, Jang HJ, et al. 2007 – Evaluation of chondroitin sulfate in shark cartilage powder as a dietary supplement: Raw materials and finished products. Food Chemistry 101, 532–539. Doi 10.1016/j.foodchem.2006.02.011
Ślusarczyk J, Adamska E, Czerwik-Marcinkowska J. 2021 – Fungi and algae as sources of medicinal and other biologically active compounds: A review. Nutrients 13, 3178. Doi 10.3390/nu13093178
Smorenburg SM, Van Noorden CJF. 2001 – The complex effects of heparins on cancer progression and metastasis in experimental studies. Pharmacological Reviews 53, 93–105. Doi 10.1016/S0031-6997(24)01481-9
Song Y, Zhang F, Linhardt RJ. 2021 – Glycosaminoglycans BT - The Role of Glycosylation in Health and Disease. In: Lauc G and Trbojević-Akmačić I (eds) pp. 103–116. Springer International Publishing, Cham
de Sousa GF, Palmero CY, de Souza-Menezes J, Araujo AK, et al. 2020 – Dermatan sulfate obtained from the Phallusia nigra marine organism is responsible for antioxidant activity and neuroprotection in the neuroblastoma-2A cell lineage. International Journal of Biological Macromolecules 164, 1099–1111. Doi 10.1016/j.ijbiomac.2020.06.285
Srimasorn S, Souter L, Green DE, Djerbal L, et al. 2022 – A quartz crystal microbalance method to quantify the size of hyaluronan and other glycosaminoglycans on surfaces. Scientific Reports 12, 10980. Doi 10.1038/s41598-022-14948-7
Staples GO, Shi X, Zaia J. 2010 – Extended N-sulfated domains reside at the nonreducing end of heparan sulfate chains. Journal of Biological Chemistry 285, 18336–18343. Doi 10.1074/jbc.M110.101592
Stern R. 2008 – Hyaluronan in cancer biology. Seminars in Cancer Biology 18, 237. Doi 10.1016/j.semcancer.2008.04.001
Sudha PN, Rose MH. 2014 – Beneficial effects of hyaluronic acid. Advances in Food and Nutrition Research 72, 137–176. Doi 10.1016/B978-0-12-800269-8.00009-9
Suleimani YM Al, Dong Y, Walker MJA. 2008 – Differential responses to various classes of drugs in a model of allergic rhinitis in guinea pigs. Pulmonary Pharmacology & Therapeutics 21, 340–348. Doi 10.1016/j.pupt.2007.08.004
Sullivan G, Garrett RD, Lenehan RF. 1971 – Occurrence of atromentin and thelephoric acid in cultures of Clitocybe Subilludens. Journal of Pharmaceutical Sciences 60, 1727–1729. Doi 10.1002/jps.2600601134
Sullivan G, Guess WL. 1969 – Atromentin: a smooth muscle stimulant in Clitocybe subilludens. Lloydia 32, 72–75.
Sultana R, Kamihira M. 2024 – Bioengineered heparin: Advances in production technology. Biotechnology Advances 77, 108456. Doi 10.1016/j.biotechadv.2024.108456
Sun H, Cao D, Liu Y, Wang H, et al. 2018 – Low molecular weight heparin-based reduction-sensitive nanoparticles for antitumor and anti-metastasis of orthotopic breast cancer. Biomaterials Science 6, 2172–2188. Doi 10.1039/C8BM00486B
Sun L, Wang X, Deng T, Luo L, et al. 2024 – Bionic sulfated glycosaminoglycan-based hydrogel inspired by snail mucus promotes diabetic chronic wound healing via regulating macrophage polarization. International Journal of Biological Macromolecules 281, 135708. Doi 10.1016/j.ijbiomac.2024.135708
Surarit R, Gopal PK, Shepherd MG. 1988 – Evidence for a glycosidic linkage between chitin and glucan in the cell wall of Candida albicans. Microbiology 134, 1723–1730. Doi 10.1099/00221287-134-6-1723
Szychowski KA, Skóra B, Pomianek T, Gmiński J. 2021 – Inonotus obliquus – from folk medicine to clinical use. Journal of Traditional and Complementary Medicine 11, 293–302. Doi 10.1016/j.jtcme.2020.08.003
Tandon R, Sharp JS, Zhang F, Pomin VH, et al. 2021 – Effective inhibition of SARS-CoV-2 entry by heparin and enoxaparin derivatives. Journal of Virology 95, e01987-20. Doi 10.1128/JVI.01987-20
Tang Y, Wang X, Li Z, He Z, et al. 2021 – Heparin prevents caspase-11-dependent septic lethality independent of anticoagulant properties. Immunity 54, 454–467. Doi 10.1016/j.immuni.2021.01.007
Teymoorian SK, Nouri H, Moghimi H. 2024 – In-vivo and in-vitro wound healing and tissue repair effect of Trametes versicolor polysaccharide extract. Scientific Reports 14, 3796. Doi 10.1038/s41598-024-54565-0
Thacker BE, Xu D, Lawrence R, Esko JD. 2014 – Heparan sulfate 3-O-sulfation: A rare modification in search of a function. Matrix Biology 35, 60–72. Doi 10.1016/j.matbio.2013.12.001
Three Ships. 2025 – Tremella mushroom benefits. Available at: https://www.threeshipsbeauty.ca/blogs/news/tremella-mushroom-benefits?srsltid=AfmBOoodRjhNuQdx_nVsOf6YzeW90mr6OAZTV_ia46Z5PgKZwqSi21zl (Accessed on January 18, 2026).
Ticar BF, Rohmah Z, Ambut C V., Choi YJ, et al. 2015 – Enzyme-assisted extraction of anticoagulant polysaccharide from Liparis tessellatus eggs. International Journal of Biological Macromolecules 74, 601–607. Doi 10.1016/j.ijbiomac.2015.01.002
Ticar BF, Rohmah Z, Neri TAN, Pahila IG, et al. 2020 – Biocompatibility and structural characterization of glycosaminoglycans isolated from heads of silver-banded whiting (Sillago argentifasciata Martin & Montalban 1935). International Journal of Biological Macromolecules 151, 663–676. Doi 10.1016/j.ijbiomac.2020.02.160
Tithi AD, Song Y, Paskaleva E, Koffas M. 2024 – Biosynthesis of animal-free recombinant chondroitin sulfate E using a functional chondroitin sulfotransferase in E. coli. Applied Microbiology and Biotechnology 108, 440. Doi 10.1007/s00253-024-13275-3
Tiwari P, Park K-I. 2024 – Advanced fungal biotechnologies in accomplishing sustainable development goals (SDGs): What do we know and what comes next? Journal of Fungi 10, 506. Doi 10.3390/jof10070506
Tovar AMF, Santos GRC, Capillé N V, Piquet AA, et al. 2016 – Structural and haemostatic features of pharmaceutical heparins from different animal sources: challenges to define thresholds separating distinct drugs. Scientific Reports 6, 35619. Doi 10.1038/srep35619
Tsai M-F, Huang C-Y, Nargotra P, Tang W-R, et al. 2023 – Green extraction and purification of chondroitin sulfate from jumbo squid cartilage by a novel procedure combined with enzyme, ultrasound and hollow fiber dialysis. Journal of Food Science and Technology 60, 1711–1722. Doi 10.1007/s13197-023-05701-7
U.S. Food and Drugs Administration. 2025 – Letter regarding the relationship between the consumption of glucosamine and/or chondroitin sulfate and a reduced risk of: osteoarthritis; osteoarthritis-related joint pain, joint tenderness, and joint swelling; joint degeneration; and cartilage deteriora. Available at: https://wayback.archive-it.org/7993/20171115122116/; https://www.fda.gov/Food/IngredientsPackagingLabeling/LabelingNutrition/ucm073400.htm (Accessed on January 22, 2026).
Uebelhart D, Malaise M, Marcolongo R, DeVathaire F, Piperno M, Mailleux E, Fioravanti A, Matoso L, Vignon E. 2004 – Intermittent treatment of knee osteoarthritis with oral chondroitin sulfate: a one-year, randomized, double-blind, multicenter study versus placebo. Osteoarthritis and Cartilage 12, 269–276. Doi 10.1016/j.joca.2004.01.004
Ultrus Prospectus. 2025 – Tremella polysaccharide hyaluronic acid. Available at: https://www.ulprospector.com/en/asia/PersonalCare/Detail/99919/1001535/Tremella-Polysaccharide-Hyaluronic-Acid#:~:text=Hyaluronic Acid is an extracted,%2C ECOCERT%2C and COSMOS certified (Accessed on January 22, 2026).
Urbi Z, Azmi NS, Ming LC, Hossain MS. 2022 – A concise review of extraction and characterization of chondroitin sulphate from fish and fish wastes for pharmacological application. Current Issues in Molecular Biology 44, 3905–3922. Doi 10.3390/cimb44090268
Vaidyanathan D, Williams A, Dordick JS, Koffas MAG, Linhardt RJ. 2017 – Engineered heparins as new anticoagulant drugs. Bioengineering & Translational Medicine 2, 17–30. Doi 10.1002/btm2.10042
Valachová K, Volpi N, Stern R, Soltes L. 2016 – Hyaluronan in medical practice. Current Medicinal Chemistry 23, 3607–3617. Doi 10.2174/0929867323666160824162133
Valcarcel J, Novoa-Carballal R, Pérez-Martín RI, Reis RL, et al. 2017 – Glycosaminoglycans from marine sources as therapeutic agents. Biotechnology Advances 35, 711–725. Doi 10.1016/j.biotechadv.2017.07.008
Varghese R, Dalvi YB, Lamrood PY, Shinde BP, et al. 2019 – Historical and current perspectives on therapeutic potential of higher basidiomycetes: an overview. 3 Biotech 9, 362. Doi 10.1007/s13205-019-1886-2
Vessella G, Traboni S, Laezza A, Iadonisi A, et al. 2020 – (Semi)-synthetic fucosylated chondroitin sulfate oligo- and polysaccharides. Marine Drugs 18, 293. Doi 10.3390/md18060293
Vessella G, Vázquez JA, Valcárcel J, Lagartera L, et al. 2021 – Deciphering structural determinants in chondroitin sulfate binding to FGF-2: Paving the way to enhanced predictability of their biological functions. Polymers 13, 313. Doi 10.3390/polym13020313
Volpi N. 2004 – Disaccharide mapping of chondroitin sulfate of different origins by high-performance capillary electrophoresis and high-performance liquid chromatography. Carbohydrate Polymers 55, 273–281. Doi 10.1016/j.carbpol.2003.09.010
Volpi N. 2007 – Analytical aspects of pharmaceutical grade chondroitin sulfates. Journal of Pharmaceutical Sciences 96, 3168–3180. Doi 10.1002/jps.20997
Wang L, Lian J, Zheng Q, Wang L, et al. 2023 – Composition analysis and prebiotics properties of polysaccharides extracted from Lepista sordida submerged cultivation mycelium. Frontiers in Microbiology 13, 1077322. Doi 10.3389/fmicb.2022.1077322
Wang P, Tang J. 2009 – Solvent-free mechanochemical extraction of chondroitin sulfate from shark cartilage. Chemical Engineering and Processing: Process Intensification 48, 1187–1191. Doi 10.1016/j.cep.2009.04.003
Wang Z, Arnold K, Dhurandhare VM, Xu Y, et al. 2021 – Investigation of the biological functions of heparan sulfate using a chemoenzymatic synthetic approach. RSC Chemical Biology 2, 702–712. Doi 10.1039/D0CB00199F
Watson RDS. 2002 – ABC of antithrombotic therapy: Antithrombotic therapy in acute coronary syndromes. BMJ give the full name here 325, 1348–1351. Doi 10.1136/bmj.325.7376.1348
Wildi LM, Raynauld JP, Martel-Pelletier J, Beaulieu A, et al. 2011 – Chondroitin sulphate reduces both cartilage volume loss and bone marrow lesions in knee osteoarthritis patients starting as early as 6 months after initiation of therapy: a randomised, double-blind, placebo-controlled pilot study using MRI. Annals of the Rheumatic Diseases 70, 982–989. Doi 10.1136/ard.2010.140848
Wise J, Greco T. 2006 – Injectable treatments for the aging face. Facial Plastic Surgery 22, 140–146. Doi 10.1055/s-2006-947720
Wolf KJ, Shukla P, Springer K, Lee S, et al. 2020 – A mode of cell adhesion and migration facilitated by CD44-dependent microtentacles. Proceedings of the National Academy of Sciences 117, 11432–11443. Doi 10.1073/pnas.1914294117
Wu M, Huang R, Wen D, Gao N, He J, Li Z, Zhao J. 2012 – Structure and effect of sulfated fucose branches on anticoagulant activity of the fucosylated chondroitin sulfate from sea cucumber Thelenata ananas. Carbohydrate Polymers 87, 862–868. Doi 10.1016/j.carbpol.2011.08.082
Wu Y, Choi M-H, Li J, Yang H, et al. 2016 – Mushroom cosmetics: The present and future. Cosmetics 3, 22. Doi 10.3390/cosmetics3030022
Wu Y, Li F, Zhang X, Li Z, et al. 2021 – Tumor microenvironment-responsive PEGylated heparin-pyropheophorbide-a nanoconjugates for photodynamic therapy. Carbohydrate Polymers 255, 117490. Doi 10.1016/j.carbpol.2020.117490
Xu Z, Chen S, Feng D, Liu Y, et al. 2021 – Biological role of heparan sulfate in osteogenesis: A review. Carbohydrate Polymers 272, 118490. Doi 10.1016/j.carbpol.2021.118490
Yamada S, Morimoto H, Fujisawa T, Sugahara K. 2007 – Glycosaminoglycans in Hydra magnipapillata (Hydrozoa, Cnidaria): demonstration of chondroitin in the developing nematocyst, the sting organelle, and structural characterization of glycosaminoglycans. Glycobiology 17, 886–894. Doi 10.1093/glycob/cwm051
Yang P, Lu Y, Gou W, Qin Y, et al. 2024 – Glycosaminoglycans’ ability to promote wound healing: From native living macromolecules to artificial biomaterials. Advanced Science 11, e2305918. Doi 10.1002/advs.202305918
Ye H, Zhang R, Zhang C, Xia Y, et al. 2025 – Advances in hyaluronic acid: Bioactivity, complexed biomaterials and biological application: A review. Asian Journal of Surgery 48, 49–61. Doi 10.1016/j.asjsur.2024.08.100
Yoon SJ, Yu MA, Pyun YR, Hwang JK, et al. 2003 – The nontoxic mushroom Auricularia auricula contains a polysaccharide with anticoagulant activity mediated by antithrombin. Thrombosis Research 112, 151–158. Doi 10.1016/j.thromres.2003.10.022
Yuvashri U, Kanchana S, Abirami A, Naidu K, et al. 2020 – Chondroitin sulfate from marine invertebrates. Encyclopedia of Marine Biotechnology, Wiley, pp. 1051–1063.
Zang L, Zhu H, Wang K, Liu Y, et al. 2022 – Not just anticoagulation—New and old applications of heparin. Molecules 27, 6968. Doi 10.3390/molecules27206968
Zare EN, Khorsandi D, Zarepour A, Yilmaz H, et al. 2024 – Biomedical applications of engineered heparin-based materials. Bioactive Materials 31, 87–118. Doi 10.1016/j.bioactmat.2023.08.002
Zegels B, Crozes P, Uebelhart D, Bruyère O, et al. 2013 – Equivalence of a single dose (1200 mg) compared to a three-time a day dose (400 mg) of chondroitin 4&6 sulfate in patients with knee osteoarthritis. Results of a randomized double blind placebo controlled study. Osteoarthritis and Cartilage 21, 22–27. Doi 10.1016/j.joca.2012.09.017
Zhang B, Shi D, Li M, Shi F, Chi L. 2023a – A quantitative mass spectrometry method to differentiate bovine and ovine heparins from pharmaceutical porcine heparin. Carbohydrate Polymers 301, 120303. Doi 10.1016/j.carbpol.2022.120303
Zhang W, Xu R, Chen J, Xiong H, et al. 2023b – Advances and challenges in biotechnological production of chondroitin sulfate and its oligosaccharides. International Journal of Biological Macromolecules 253, 126551. Doi 10.1016/j.ijbiomac.2023.126551
Zhang Y, Dong J, Xu G, Han R, et al. 2023c – Efficient production of hyaluronic acid by Streptococcus zooepidemicus using two-stage semi-continuous fermentation. Bioresource Technology 377, 128896. Doi 10.1016/j.biortech.2023.128896
Zhang Y, Wang Y, Zhou Z, Wang P, et al. 2022 – Synthesis of bioengineered heparin by recombinant yeast Pichia pastoris. Green Chemistry 24, 3180–3192. Doi 10.1039/D1GC04672A
Zhou H, Ni J, Huang W, Zhang J. 2006 – Separation of hyaluronic acid from fermentation broth by tangential flow microfiltration and ultrafiltration. Separation and Purification Technology 52, 29–38. Doi 10.1016/j.seppur.2006.03.011
Zhou J, Ge W, Zhang X, Wu J, Chen Q, et al. 2020 – Effects of spent mushroom substrate on the dissipation of polycyclic aromatic hydrocarbons in agricultural soil. Chemosphere 259, 127462. Doi 10.1016/j.chemosphere.2020.127462
Zhou X, Chandarajoti K, Pham TQ, Liu R, et al. 2011 – Expression of heparan sulfate sulfotransferases in Kluyveromyces lactis and preparation of 3’-phosphoadenosine-5’-phosphosulfate. Glycobiology 21, 771–780. Doi 10.1093/glycob/cwr001
Zhu S, Li J, Loka RS, Song Z, et al. 2020 – Modulating heparanase activity: Tuning sulfation pattern and glycosidic linkage of oligosaccharides. Journal of Medicinal Chemistry 63, 4227–4255. Doi 10.1021/acs.jmedchem.0c00156
Zou G, Nielsen JB, Wei Y. 2023 – Harnessing synthetic biology for mushroom farming. Trends in Biotechnology 41, 480–483. Doi 10.1016/j.tibtech.2022.10.001
Zulueta MML, Chyan C, Hung S. 2018 – Structural analysis of synthetic heparan sulfate oligosaccharides with fibroblast growth factors and heparin-binding hemagglutinin. Current Opinion in Structural Biology 50, 126–133. Doi 10.1016/j.sbi.2018.03.003
