Submerged fermentation of mycelial biomass and exopolysaccharide of Philippine Lentinus tigrinus: FTIR spectroscopy and bioactivity profiling
DOI:
https://doi.org/10.60923/issn.2531-7342/24237Keywords:
Antibacterial, Antioxidant, Cytotoxicity, Enzyme-based assay, TeratogenicityAbstract
Lentinus tigrinus (tiger sawgill mushroom) is an edible mushroom known for its high nutritional value. This study highlights the optimized submerged fermentation conditions for L. tigrinus to enhance mycelial biomass and exopolysaccharide (EPS) production, identifying mango puree as the most suitable culture medium. Optimal parameters were established at pH 7.0, 100 rpm agitation, and 10–15 days of incubation. Fourier transform infrared (FTIR) analysis revealed major absorption peaks at 3291.66, 2927.90, 1614.83, 1411.92, 1257.60, and 1043.35–416.27 cm⁻¹, indicating the presence of hydroxyl, carbonyl, amino, and ether groups typical of β-D-glucans and protein–polysaccharide conjugates. Enzyme-based assays showed low inhibitory activity against α-glucosidase, acetylcholinesterase, and cyclooxygenase 1 and 2. The 2,2-diphenyl-1-picrylhydrazyl (DPPH) radical scavenging assay demonstrated strong radical scavenging activity in the mycelial (62.69%) and EPS (50.76%) extracts. Brine shrimp lethality tests revealed dose-dependent toxicity in the mycelial extract but benign toxicity in EPS. Zebrafish embryo assays confirmed concentration-dependent embryotoxicity in the mycelial extract, while EPS exhibited minimal developmental effects, underscoring its biocompatibility. The mycelial extract also showed antibacterial activity against Staphylococcus aureus (8.83 mm). Overall, L. tigrinus demonstrates promise as a sustainable Philippine source of bioactive metabolites and polysaccharides with potential nutraceutical and pharmaceutical applications.
References
Acanto RB, Cuaderes VHS, Gomoto PH (2022) Phytochemical screening, cytotoxic activity, and proximate analysis of split gill mushroom (Schizophyllum commune). JPAIR Multidisciplinary Research 47(1):15–29. https://doi.org/10.7719/jpair.v47i1.542
Agatonovic-Kustrin S, Doyle E, Gegechkori V, Morton DW (2020) High-performance thin-layer chromatography linked with (bio)assays and FTIR-ATR spectroscopy as a method for discovery and quantification of bioactive components in native Australian plants. Journal of Pharmaceutical and Biomedical Analysis 184:113208. https://doi.org/10.1016/j.jpba.2020.113208
Agatonovic-Kustrin S, Ristivojevic P, Gegechkori V, Litvinova TM, Morton DW (2020) Essential oil quality and purity evaluation via FT-IR spectroscopy and pattern recognition techniques. Applied Sciences 10(20):7294. https://doi.org/10.3390/app10207294
Agudelo-Escobar LM, Gutiérrez-López Y, Urrego-Restrepo S (2017) Effects of aeration, agitation and pH on the production of mycelial biomass and exopolysaccharide from the filamentous fungus Ganoderma lucidum. DYNA 84(200):72–79. https://doi.org/10.15446/dyna.v84n200.57126
Alves MJ, Ferreira ICFR, Dias J, Teixeira V, Martins A, Pintado M (2012) A review on antimicrobial activity of mushroom (Basidiomycetes) extracts and isolated compounds. Planta Medica 78(16):1707–1718. https://doi.org/10.1055/s-0032-1315370
Angelova G, Brazkova M, Mihaylova D, Slavov A, Petkova N, Blazheva D, Deseva I, Gotova I, Dimitrov Z, Krastanov A (2022) Bioactivity of biomass and crude exopolysaccharides obtained by controlled submerged cultivation of medicinal mushroom Trametes versicolor. Journal of Fungi 8(7):738. https://doi.org/10.3390/jof8070738
Angelova G, Govedarova E, Brazkova M, Kostov G, Krastanov A (2021) Optimization of exopolysaccharide synthesis by medicinal fungus Trametes versicolor in submerged culture. Agricultural Sciences 13:84–93. https://doi.org/10.22620/agrisci.2021.30.012
Austria AB, Dulay RMR, Pambid RC (2021) Mycochemicals, antioxidant and antidiabetic properties of Philippine sawgill mushroom Lentinus swartzii (Higher Basidiomycetes). Asian Journal of Agriculture and Biology 2021(2):202006365. https://doi.org/10.35495/ajab.2020.06.365
Azuma S, Kitamoto Y (1994) Nutritional environment for mycelial growth and fruit-body formation in Lentinus edodes. Mushroom Science and Biotechnology 1:7–13. https://doi.org/10.24465/kinoko.1.1_7
Bajpai VK, Rahman A, Kang SC (2008) Chemical composition and inhibitory parameters of essential oil and extracts of Nandina domestica Thunb. to control food-borne pathogenic and spoilage bacteria. International Journal of Food Microbiology 125(2):117–122. https://doi.org/10.1016/j.ijfoodmicro.2008.03.011
Baravalia Y, Vaghasiya Y, Chanda S (2012) Brine shrimp cytotoxicity, anti-inflammatory and analgesic properties of Woodfordia fruticosa Kurz flowers. Iranian Journal of Pharmaceutical Research 11:851–861.
Baraza LD, Joseph CC, Moshi MJ, Nkunya MHH (2009) Chemical constituents and biological activity of three Tanzanian wild mushroom species. Tanzania Journal of Science 33:1–8. https://doi.org/10.4314/tjs.v33i1.44280
Barros L, Ferreira M, Queirós B, Ferreira IC, Baptista P (2006) Total phenols, ascorbic acid, β-carotene and lycopene in Portuguese wild edible mushrooms and their antioxidant activities. Food Chemistry 103(2):413–419. https://doi.org/10.1016/j.foodchem.2006.07.038
Batbayar S, Kim MJ, Kim HW (2011) Medicinal mushroom Lingzhi or Reishi, Ganoderma lucidum (W. Curt.: Fr.) P. Karst., β-glucan induces Toll-like receptors and fails to induce inflammatory cytokines in NF-κB inhibitor-treated macrophages. International Journal of Medicinal Mushrooms 13(3):213–225. https://doi.org/10.1615/intjmedmushr.v13.i3.10
Batbayar S, Lee DH, Kim HW (2012) Immunomodulation of fungal β-glucan in host defense signaling by dectin-1. Biomolecules and Therapeutics 20:433–445. https://doi.org/10.4062/biomolther.2012.20.5.433
Bauer AW, Kirby WMM, Sherris JC, Turck M (1966) Antibiotic susceptibility testing by a standardized single disk method. American Journal of Clinical Pathology 45:493–496. https://doi.org/10.1093/ajcp/45.4_ts.493
Bello-Pérez LA, García-Suárez FJL, Agama-Acevedo E (2007) Mango carbohydrates. Food 1(1):36–40.
Bhat R, Sharanabasava VG, Deshpande R, Shetti U, Sanjeev G, Venkataraman A (2013) Photo-bio-synthesis of irregular shaped functionalized gold nanoparticles using edible mushroom Pleurotus florida and its anticancer evaluation. Journal of Photochemistry and Photobiology B: Biology 125:63–69. https://doi.org/10.1016/j.jphotobiol.2013.05.002
Bischoff H (1994) Pharmacology of alpha-glucosidase inhibition. European Journal of Clinical Investigation 24(53):3-10. https://doi.org/10.1111/j.1365-2362.1994.tb02249.x
Boukes GJ, Koekemoer TC, van de Venter M, Govender S (2017) Cytotoxicity of thirteen South African macrofungal species against five cancer cell lines. South African Journal of Botany 113:62–67. https://doi.org/10.1016/j.sajb.2017.07.010
Carballo JL, Hernández-Inda ZL, Pérez P, García-Grávalos MD (2002) A comparison between two brine shrimp assays to detect in vitro cytotoxicity in marine natural products. BMC Biotechnology 2(1):17. https://doi.org/10.1186/1472-6750-2-17
Da Costa ECS, Santos LLD, Grazziotti PH, Da Costa LS, Silva CS, De Vasconcelos Ramires R, Da Costa MR, De Abreu CM (2020) Inoculant production of Pisolithus sp. in submerged culture under agitation. African Journal of Agricultural Research 16(5):746–751. https://doi.org/10.5897/ajar2019.14654
De Castro MEG, Dulay RMR (2015) Toxic and teratogenic effects of Lentinus sajor-caju and Pleurotus ostreatus ethanolic extracts in Danio rerio embryo model. International Journal of Biology, Pharmacy and Allied Sciences 4(4):2261–2269.
De Castro MEG, Dulay RMR, Enriquez MLD (2016) Toxic and teratogenic effects of medicinal and culinary mushroom Termitomyces clypeatus, collected from the termite mound in Mt. Makiling Forest Reserve, Los Baños, Laguna, Philippines, on developing embryos of zebrafish (Danio rerio). Der Pharmacia Lettre 8(5):237–242.
De Leon AM, Reyes RG, Cruz TD (2013) Enriched cultivation of three wild strains of Lentinus tigrinus (Bull.) Fr. using agricultural wastes. International Journal of Agricultural Technology 9:1199–1214.
De Silva DD, Rapior S, Hyde KD, Bahkali AH (2012) Medicinal mushrooms in prevention and control of diabetes mellitus. Fungal Diversity 56:1–29. https://doi.org/10.1007/s13225-012-0187-4
De Souza Kirsch L, De Macedo AJP, Teixeira MFS (2016) Production of mycelial biomass by the Amazonian edible mushroom Pleurotus albidus. Brazilian Journal of Microbiology 47(3):658–664. https://doi.org/10.1016/j.bjm.2016.04.007
Deacon JW (2006) Fungal Biology. Blackwell Publishing Ltd., New York.
Dulay RMR (2011) Optimization of culture conditions, nutritional characterization, toxicity and teratogenicity of Lentinus tigrinus–A newly recorded domesticated wild mushroom in the Philippines. Master’s thesis, De La Salle University.
Dulay RMR (2021) Molecular identification, submerged culture conditions, and cytotoxicity of Lentinus species from Luzon Island, Philippines. PhD dissertation, De La Salle University
Dulay RMR, Arenas MC, Kalaw SP, Reyes RG, Cabrera EC (2014a) Proximate composition and functionality of the culinary-medicinal tiger sawgill mushroom, Lentinus tigrinus (higher basidiomycetes), from the Philippines. International Journal of Medicinal Mushrooms 16:85–94. https://doi.org/10.1615/intjmedmushr.v16.i1.80
Dulay RMR, Cabrera EC, Kalaw SP, Reyes RG, Alfonso NF (2012a) Optimization of culture conditions for mycelial growth and basidiocarp production of Lentinus tigrinus (Bull.) Fr., a new record of domesticated wild edible mushroom in the Philippines. Philippine Agricultural Scientist 95:278–285.
Dulay RMR, Cabrera EC, Kalaw SP, Reyes RG (2012b) Optimal growth conditions for basidiospore germination and morphogenesis of Philippine wild strain of Lentinus tigrinus (Bull.) Fr. Mycosphere 3:926–933. https://doi.org/10.5943/mycosphere/3/6/6
Dulay RMR, Cabrera EC, Kalaw SP, Reyes RG (2021) Optimization of submerged culture conditions for mycelial biomass production of fourteen Lentinus isolates from Luzon Island, Philippines. Biocatalysis and Agricultural Biotechnology 38:102226. https://doi.org/10.1016/j.bcab.2021.102226
Dulay RMR, Cabrera EC, Kalaw SP, Reyes RG, Hou CT (2020) Nutritional requirements for mycelial growth of three Lentinus species from the Philippines. Biocatalysis and Agricultural Biotechnology 23:101506. https://doi.org/10.1016/j.bcab.2020.101506
Dulay RMR, Cardona EMG, Kalaw SP, Reyes RG (2016b) Optimization of liquid culture conditions of Coprinopsis cinerea as natural source of bioactive compounds. Der Pharma Chemica 8(18):313–319.
Dulay RMR, Flores KS, Tiniola RC, Marquez DHH, Dela Cruz AG, Kalaw SP, Reyes RG (2015a) Mycelial biomass production and antioxidant activity of Lentinus tigrinus and Lentinus sajor-caju in indigenous liquid culture. Mycosphere 6(6):659–666. https://doi.org/10.5943/mycosphere/6/6/2
Dulay RMR, Kalaw SP, Reyes RG, Alfonso NF, Eguchi F (2012c) Teratogenic and toxic effects of Lingzhi or Reishi medicinal mushroom, Ganoderma lucidum (W. Curt.: Fr.) P. Karst. (higher basidiomycetes), on zebrafish embryo as model. International Journal of Medicinal Mushrooms 14:507–512. https://doi.org/10.1615/intjmedmushr.v14.i5.90
Dulay RMR, Kalaw SP, Reyes RG, Cabrera EC (2014b) Embryo-toxic and teratogenic effects of Philippine strain of Lentinus tigrinus (Tiger sawgill basidiomycetes) extract on zebrafish (Danio rerio) embryos. Annals of Biological Research 5(6):9–14.
Dulay RMR, Miranda LA, Malasaga JS, Kalaw SP, Reyes RG, Hou CT (2017) Antioxidant and antibacterial activities of acetonitrile and hexane extracts of Lentinus tigrinus and Pleurotus djamour. Biocatalysis and Agricultural Biotechnology 9:141–144. https://doi.org/10.1016/j.bcab.2016.12.003
Dulay RMR, Pamiloza DG, Ramirez RL (2018) Toxic and teratogenic effects of mycelia and fruiting body extracts of Lentinus strigosus (BIL 1324) in zebrafish (Danio rerio) embryo. International Journal of Biosciences 13(5):205–211.
Dulay RMR, Ray K, Hou CT (2015b) Optimization of liquid culture conditions of Philippine wild edible mushrooms as potential source of bioactive lipids. Biocatalysis and Agricultural Biotechnology 4(3):409–415. https://doi.org/10.1016/j.bcab.2015.04.003
Dytkiewitz E, Morlock GE (2008) Analytical strategy for rapid identification and quantification of lubricant additives in mineral oil by high-performance thin-layer chromatography with UV absorption and fluorescence detection combined with mass spectrometry and infrared spectroscopy. Journal of AOAC International 91(5):1237–1244.
Elisashvili V (2012) Submerged cultivation of medicinal mushrooms: bioprocesses and products (review). International Journal of Medicinal Mushrooms 14(3):211–239. https://doi.org/10.1615/intjmedmushr.v14.i3.10
Elsacker E, Vandelook S, Van Wylick A, Ruytinx J, De laet L, Peeters E (2020) A comprehensive framework for the production of mycelium-based lignocellulosic composites. Science of The Total Environment 725:138431. https://doi.org/10.1016/j.scitotenv.2020.138431
Eskandari-Nojedehi M, Jafarizadeh-Malmiri H, Rahbar-Shahrouzi J (2017) Hydrothermal green synthesis of gold nanoparticles using mushroom (Agaricus bisporus) extract: physico-chemical characteristics and antifungal activity studies. Green Processing and Synthesis 7(1):38–47. https://doi.org/10.1515/gps-2017-0004
Fabros JA, Dulay RMR, De Leon AM, Kalaw SP, Reyes RG (2022) Distribution, cultivation, nutritional composition, and bioactivities of Lentinus (Polyporaceae, Basidiomycetes): a review. Current Research in Environmental & Applied Mycology 12:170–219. https://doi.org/10.5943/cream/12/1/13
Ferreira ICFR, Barros L, Abreu RMV (2009) Antioxidants in wild mushrooms. Current Medicinal Chemistry 16(12):1543–1560. https://doi.org/10.2174/092986709787909587
Ferreira ICFR, Heleno SA, Reis FS, Stojkovic D, Queiroz MJRP, Vasconcelos MH, Sokovic M (2014) Chemical features of Ganoderma polysaccharides with antioxidant, antitumor and antimicrobial activities. Phytochemistry 114:38–55. https://doi.org/10.1016/j.phytochem.2014.10.011
Ferreira SS, Passos CP, Madureira P, Vilanova M, Coimbra MA (2015) Structure–function relationships of immunostimulatory polysaccharides: a review. Carbohydrate Polymers 132:378–396. https://doi.org/10.1016/j.carbpol.2015.05.079
Heleno SA, Barros L, Sousa MJ, Martins A, Ferreira ICFR (2009) Tocopherols composition of Portuguese wild mushrooms with antioxidant capacity. Food Chemistry 119(4):1443–1450. https://doi.org/10.1016/j.foodchem.2009.09.025
Heleno SA, Ferreira ICFR, Esteves AP, Ćirić A, Glamočlija J, Martins A, Soković M, Queiroz MJRP (2013) Antimicrobial and demelanizing activity of Ganoderma lucidum extract, p-hydroxybenzoic and cinnamic acids and their synthetic acetylated glucuronide methyl esters. Food and Chemical Toxicology 58:95–100. https://doi.org/10.1016/j.fct.2013.04.025
Howe K, Clark MD, Torroja CF, Torrance J, Berthelot C, Muffato M, Collins JE, Humphray S, McLaren K, Matthews L, et al. (2013) The zebrafish reference genome sequence and its relationship to the human genome. Nature 496(7446):498–503. https://doi.org/10.1038/nature12111
Hwang HJ, Kim SW, Choi JW, Yun JW (2003) Production and characterization of exopolysaccharides from submerged culture of Phellinus linteus KCTC 6190. Enzyme and Microbial Technology 33(2–3):309–319. https://doi.org/10.1016/s0141-0229(03)00131-5
ICBF - Instituto Colombiano de Bienestar Familiar (2015) Tabla de composición de alimentos Colombia. Bogotá SAS, editor. Bogotá: Universidad Nacional de Colombia, 1–321.
Joshi K, Meher MK, Poluri KM (2020) Fabrication and characterization of bioblocks from agricultural waste using fungal mycelium for renewable and sustainable applications. ACS Applied Bio Materials 3:1884–1892. https://doi.org/10.1021/acsabm.9b01047
Joshi M, Patel H, Gupte S, Gupte A (2012) Nutrient improvement for simultaneous production of exopolysaccharide and mycelial biomass by submerged cultivation of Schizophyllum commune AGMJ-1 using statistical optimization. 3 Biotech 3(4):307–318. https://doi.org/10.1007/s13205-012-0103-3
Kadiri M, Fasidi IO (1994) Growth requirements of Lentinus subnudus Berk, a Nigerian edible mushroom. Chemie, Mikrobiologie, Technologie der Lebensmittel 16(3–4):80–84.
Kalaw SP, De Leon AM, Damaso EJ Jr, Ramos JC, Del Rosario MAG, Abon MD, Undan JR, Dulay RMR, Reyes RG (2021) Cultivation of different strains of Lentinus tigrinus from selected areas of Luzon Island, Philippines. Studies in Fungi 6:299–306. https://doi.org/10.5943/sif/6/1/20
Kim HG, Yoon DH, Lee WH, Han SK, Shrestha B, Kim CH, Lim MH, Chang W, Lim S, Choi S, Song WO, Sung JM, Hwang KC, Kim TW (2007) Phellinus linteus inhibits inflammatory mediators by suppressing redox-based NF-κB and MAPKs activation in lipopolysaccharide-induced RAW 264.7 macrophage. Journal of Ethnopharmacology 114(3):307–315. https://doi.org/10.1016/j.jep.2007.08.011
Kim HM, Park MK, Yun JW (2006) Culture pH affects exopolysaccharide production in submerged mycelial culture of Ganoderma lucidum. Applied Biochemistry and Biotechnology 134(3):249–262. https://doi.org/10.1385/abab:134:3:249
Kim HS, Kim JY, Lee HK, Kim MS, Lee SR, Kang JS, Kim HM, Lee KA, Hong JT, Kim Y, Han SB (2010) Dendritic cell activation by glucan isolated from Umbilicaria esculenta. Immune Network 10(6):188. https://doi.org/10.4110/in.2010.10.6.188
Kim SS, Lee JS, Cho JY, Kim YE, Hong EK (2010b) Process development for mycelial growth and polysaccharide production in Tricholoma matsutake liquid culture. Journal of Bioscience and Bioengineering 109(4):351–355. https://doi.org/10.1016/j.jbiosc.2009.10.010
Ko HH, Hung CF, Wang JP, Lin CN (2007) Anti-inflammatory triterpenoids and steroids from Ganoderma lucidum and G. tsugae. Phytochemistry 69(1):234–239. https://doi.org/10.1016/j.phytochem.2007.06.008
Kolak U, Öztürk M, Özgökçe F, Ulubelen A (2006) Norditerpene alkaloids from Delphinium linearilobum and antioxidant activity. Phytochemistry 67:2170–2175. https://doi.org/10.1016/j.phytochem.2006.06.006
Kour H, Kour D, Kour S, Singh S, Hashmi SAJ, Yadav AN, Kumar K, Sharma YP, Ahluwalia AS (2022) Bioactive compounds from mushrooms: emerging bioresources of food and nutraceuticals. Food Bioscience 50:102124. https://doi.org/10.1016/j.fbio.2022.102124
Kozarski M, Klaus A, Jakovljevic D, Todorovic N, Vunduk J, Petrović P, Niksic M, Vrvic M, Van Griensven L (2015) Antioxidants of edible mushrooms. Molecules 20(10):19489–19525. https://doi.org/10.3390/molecules201019489
Kumar H, Bhardwaj K, Sharma R, Nepovimova E, Cruz-Martins N, Dhanjal DS, Sing R, Chopra C, Verma R, Abd-Elsalam KA, Tapwal A, Musilek K, Kumar D, Kuča K (2021) Potential usage of edible mushrooms and their residues to retrieve valuable supplies for industrial applications. Journal of Fungi 7:427. https://doi.org/10.3390/jof7060427
Li N, Li H, Liu Z, Feng G, Shi C, Wu Y (2023) Unveiling the therapeutic potentials of mushroom bioactive compounds in Alzheimer’s disease. Foods 12(15):2972. https://doi.org/10.3390/foods12152972
Lian W, Yang X, Duan Q, Li J, Zhao Y, Yu C, He T, Sun T, Zhao Y, Wang W (2024) The biological activity of Ganoderma lucidum on neurodegenerative diseases: the interplay between different active compounds and the pathological hallmarks. Molecules 29(11):2516. https://doi.org/10.3390/molecules29112516
Liwanag EJ, Dulay RMR, Kalaw SP (2020) Mycelial growth of Philippine mushroom Lentinus tigrinus in selected cucurbit-based media and its antioxidant activity. Asian Journal of Agriculture and Biology 8:323–329.
Manjunathan J, Kaviyarasan V (2010) Studies on the growth requirements of Lentinus tuberregium (Fr.), an edible mushroom. Middle-East Journal of Scientific Research 5(2):81–85.
Manzano EM, Magat GCM, Dulay RMR, De Leon AMD (2025). Effect of Pseudomonas fluorescens and Bacillus cereus on the biomass production and bioactivity of Lentinus tigrinus and Lentinus strigosus in submerged culture. International Journal of Agricultural Biology 34:340506. https://doi.org/10.17957/IJAB/15.2391
Meyer BN, Ferrigni NR, Putnam JE, Jacobsen LB, Nichols DE, McLaughlin JL (1982). Brine shrimp: a convenient general bioassay for active plant constituents. Planta Medica 45(05):31–34. https://doi.org/10.1055/s-2007-971236
Movasaghi Z, Rehman S, Rehman IU (2008). Fourier Transform infrared (FTIR) spectroscopy of biological tissues. Applied Spectroscopy Reviews 43(2):134–179. https://doi.org/10.1080/05704920701829043
Nagel R (2002) DarT: The embryo test with the zebrafish Danio rerio – a general model in ecotoxicology and toxicology. Altex 19(Suppl 1):38–48.
Naing M, Yerro J, Fernandez P, Amor E (2019) Alpha-glucosidase inhibition assay. In: Tuklas Lunas® Protocols for Drug Discovery and Development Volume IIB: Primary Bioactivity Assays (Guevarra A, Alvero R. eds). Philippine Council for Health Research and Development, Taguig, pp. 2–16.
Nguyen TK, Im KH, Choi J, Shin PG, Lee TS (2016) Evaluation of antioxidant, anti-cholinesterase, and anti-inflammatory effects of culinary mushroom Pleurotus pulmonarius. Mycobiology 44(4):291–301. https://doi.org/10.5941/myco.2016.44.4.291
Niego AGT, Rapior S, Thongklang N, Raspé O, Hyde KD, Mortimer P (2022) Reviewing the contributions of macrofungi to forest ecosystem processes and services. Fungal Biology Reviews 44:100294. https://doi.org/10.1016/j.fbr.2022.11.002
Nikaido H (2003) Molecular basis of bacterial outer membrane permeability revisited. Microbiology and Molecular Biology Reviews 67(4):593–656. https://doi.org/10.1128/mmbr.67.4.593-656.2003
Nour AAM, Khalid KSM, Osman GAM (2011) Suitability of some Sudanese mango varieties for jam making. American Journal of Scientific and Industrial Research 2(1):17–23. https://doi.org/10.5251/ajsir.2011.2.1.17.23
Ooi VEC, Liu F (2000) Immunomodulation and anti-cancer activity of polysaccharide-protein complexes. Current Medicinal Chemistry 7:715–729. https://doi.org/10.2174/0929867003374705
Opog AC, Amor E (2019) Cyclooxygenase (COX) inhibition assay: fluorometric method. In: Tuklas Lunas® Protocols for Drug Discovery and Development Volume IIB: Primary Bioactivity Assays (Guevarra A, Alvero R, eds). Philippine Council for Health Research and Development, Taguig, pp 130–150.
Othman OC, Mbogo, GP (2009) Physico-chemical characteristics of storage-ripened Mango (Mangifera indica L.) fruits varieties of Eastern Tanzania. Tanzania Journal of Science 35(1):6. https://doi.org/10.65085/2507-7961.1904
Patocka J (2012) Natural cholinesterase inhibitors from mushrooms. Military Medical Science Letters 81(1):40–44. https://doi.org/10.31482/mmsl.2012.005
Petrova RD, Reznick AZ, Wasser SP, Denchev CM, Nevo E, Mahajna J (2008) Fungal metabolites modulating NF-κB activity: an approach to cancer therapy and chemoprevention (review). Oncology Reports 19(2):299–308. https://doi.org/10.3892/or.19.2.299
Randhawa K, Shri R (2018) Comparison of antioxidant and anticholinesterase activities of selected Pleurotus species (Agaricomycetes) from India. International Journal of Medicinal Mushrooms 20(8):739–748. https://doi.org/10.1615/intjmedmushrooms.2018027033
Rosak C, Mertes G (2012) Critical evaluation of the role of acarbose in the treatment of diabetes: patient considerations. Diabetes Metabolic Syndrome and Obesity 357. https://doi.org/10.2147/dmso.s28340
Roseiro LB, Rauter AP, Serralheiro MLM (2012) Polyphenols as acetylcholinesterase inhibitors: structural specificity and impact on human disease. Nutrition and Aging 1(2):99–111. https://doi.org/10.3233/nua-2012-0006
Ruthes AC, Smiderle FR, Iacomini M (2015) Mushroom heteropolysaccharides: a review on their sources, structure and biological effects. Carbohydrate Polymers 136:358–375. https://doi.org/10.1016/j.carbpol.2015.08.061
Sevindik M (2018) Investigation of antioxidant/oxidant status and antimicrobial activities of Lentinus tigrinus. Advances in Pharmacological Sciences 2018:1–4. https://doi.org/10.1155/2018/1718025
Shen Y, Zhao H, Wang X, Wu S, Wang Y, Wang C, Zhang Y, Zhao H (2024) Unraveling the web of defense: the crucial role of polysaccharides in immunity. Frontiers in Immunology 15:1406213. https://doi.org/10.3389/fimmu.2024.1406213
Siu KC, Chen X, Wu JY (2014) Constituents actually responsible for the antioxidant activities of crude polysaccharides isolated from mushrooms. Journal of Functional Foods 11:548–556. https://doi.org/10.1016/j.jff.2014.08.012
Sivanandhan S, Khusro A, Paulraj M, Ignacimuthu S, Al-Dhabi N (2017) Biocontrol properties of basidiomycetes: an overview. Journal of Fungi 3:2. https://doi.org/10.3390/jof3010002
Tabibzadeh F, Alvandi H, Hatamian-Zarmi A, Kalitukha L, Aghajani H, Ebrahimi-Hosseinzadeh B (2022) Antioxidant activity and cytotoxicity of exopolysaccharide from mushroom Hericium coralloides in submerged fermentation. Biomass Conversion and Biorefinery 14(21):26953–26963. https://doi.org/10.1007/s13399-022-03386-0
Tamfu AN, Kucukaydin S, Yeskaliyeva B, Ozturk M, Dinica RM (2021) Non-alkaloid cholinesterase inhibitory compounds from natural sources. Molecules 26(18):5582. https://doi.org/10.3390/molecules26185582
Taufek NM, Harith HH, Rahim MHA, Ilham Z, Rowan N, Wan-Mohtar WAAQI (2020) Performance of mycelial biomass and exopolysaccharide from Malaysian Ganoderma lucidum for the fungivore red hybrid tilapia (Oreochromis sp.) in zebrafish embryo. Aquaculture Reports 17:100322. https://doi.org/10.1016/j.aqrep.2020.100322
Thornton BP, Vetvicka V, Pitman M, Goldman RC, Ross GD (1996) Analysis of the sugar specificity and molecular location of the β-glucan-binding lectin site of complement receptor type 3 (CD11b/CD18). Journal of Immunology 156(3):1235–1246. https://doi.org/10.4049/jimmunol.156.3.1235
United States Department of Agriculture (2018) USDA Nutrient Lists from Standard Reference Legacy. Human Nutrition and Food Safety, https://www.nal.usda.gov/human-nutrition-and-food-safety/nutrient-lists-standard-reference-legacy-2018
Ványolós A, Muszyńska B, Chuluunbaatar B, Gdula-Argasińska J, Kała K, Hohmann J (2020) Extracts and steroids from the edible mushroom Hypholoma lateritium exhibit anti-inflammatory properties by inhibition of COX-2 and activation of Nrf2. Chemistry & Biodiversity 17(9). https://doi.org/10.1002/cbdv.202000391
Wahyudi P, Mangunwardoyo W, Sumaryono W, Gandjar I (2015) Optimization of submerged culture for biomass and polysaccharide of Pleurotus ostreatus BPPTCC 6017 using response surface methodology. Malaysian Journal of Microbiology 11(1):27–39. https://doi.org/10.21161/mjm.62014
Wang XM, Zhang J, Wu LH, Zhao YL, Li T, Li JQ, Liu HG (2014) A mini-review of chemical composition and nutritional value of edible wild-grown mushrooms from China. Food Chemistry 151:279–285. https://doi.org/10.1016/j.foodchem.2013.11.062
Wan-Mohtar WAAQI, Ilham Z, Jamaludin AA, Rowan N (2021) Use of zebrafish embryo assay to evaluate toxicity and safety of bioreactor-grown exopolysaccharides and endopolysaccharides from European Ganoderma applanatum mycelium for future aquaculture applications. International Journal of Molecular Sciences 22(4):1675. https://doi.org/10.3390/ijms22041675
Wasser S (2002) Medicinal mushrooms as a source of antitumor and immunomodulating polysaccharides. Applied Microbiology and Biotechnology 60:258–274. https://doi.org/10.1007/s00253-002-1076-7
Wasser S (2014) Medicinal mushroom science: Current perspectives, advances, evidences, and challenges. Biomedical Journal 37(6):345. https://doi.org/10.4103/2319-4170.138318
Wasser SP (2010) Medicinal mushroom science: history, current status, future trends, and unsolved problems. International Journal of Medicinal Mushrooms 12:1–16. https://doi.org/10.1615/intjmedmushr.v12.i1.10
Weigt S, Huebler N, Strecker R, Braunbeck T, Broschard TH (2011) Zebrafish (Danio rerio) embryos as a model for testing proteratogens. Toxicology 281(1–3):25–36. https://doi.org/10.1016/j.tox.2011.01.004
Wu J, Ding ZY, Zhang KC (2006) Improvement of exopolysaccharide production by macro-fungus Auricularia auricula in submerged culture. Enzyme and Microbial Technology 39(4):743–749. https://doi.org/10.1016/j.enzmictec.2005.12.012
Wu YL, Han F, Luan SS, Ai R, Zhang P, Li H, Chen LX (2019) Triterpenoids from Ganoderma lucidum and their potential anti-inflammatory effects. Journal of Agricultural and Food Chemistry 67(18):5147–5158. https://doi.org/10.1021/acs.jafc.9b01195
Wu ZW, Zhao XF, Quan CX, Liu XC, Tao XY, Li YJ, Peng XR, Qiu MH (2025) Structure–function insights of natural Ganoderma polysaccharides: Advances in biosynthesis and functional food applications. Natural Products and Bioprospecting 15(1):15. https://doi.org/10.1007/s13659-025-00496-w
Xiao C, Wu Q, Xie Y, Zhang J, Tan J (2015) Hypoglycemic effects of Grifola frondosa (Maitake) polysaccharides F2 and F3 through improvement of insulin resistance in diabetic rats. Food & Function 6(11):3567–3575. https://doi.org/10.1039/c5fo00497g
Yao S, Li T, Li JQ, Liu HG, Wang YZ (2018) Geographic identification of Boletus mushrooms by data fusion of FT-IR and UV spectroscopies combined with multivariate statistical analysis. Spectrochimica Acta Part A: Molecular and Biomolecular Spectroscopy 198:257–263. https://doi.org/10.1016/j.saa.2018.03.018
Yin Cm, Fan X, Liu C, Fan Z, Shi DF, Yao F, Cheng W, Gao H (2019) The antioxidant properties, tyrosinase and α-glucosidase inhibitory activities of phenolic compounds in different extracts from the golden oyster mushroom, Pleurotus citrinopileatus (Agaricomycetes). International Journal of Medicinal Mushrooms 21(9):865–874. https://doi.org/10.1615/intjmedmushrooms.2019031857
Ying YM, Zhang LY, Zhang X, Bai HB, Liang DE, Ma LF, Shan WG, Zhan ZJ (2014) Terpenoids with α-glucosidase inhibitory activity from the submerged culture of Inonotus obliquus. Phytochemistry 108:171–176. https://doi.org/10.1016/j.phytochem.2014.09.022
Zaidman BZ, Yassin M, Mahajna J, Wasser SP (2005) Medicinal mushroom modulators of molecular targets as cancer therapeutics. Applied Microbiology and Biotechnology 67:453–468. https://doi.org/10.1007/s00253-004-1787-z
Zhang M, Cui SW, Cheung PCK, Wang Q (2006) Antitumor polysaccharides from mushrooms: a review on their isolation process, structural characteristics and antitumor activity. Trends in Food Science & Technology 18(1):4–19. https://doi.org/10.1016/j.tifs.2006.07.013
Zhu ZY, Guo MZ, Liu F, Luo Y, Chen L, Meng M, Wang XT, Zhang YM (2016) Preparation and inhibition on α-D-glucosidase of low molecular weight polysaccharide from Cordyceps militaris. International Journal of Biological Macromolecules 93:27–33. https://doi.org/10.1016/j.ijbiomac.2016.08.058
Downloads
Additional Files
Published
How to Cite
Issue
Section
License
Copyright (c) 2026 Rich Milton R. Dulay, Eduard M. Manzano

This work is licensed under a Creative Commons Attribution 4.0 International License.