At present, there are 15,000 species of large fungi in the world, of which more than 2,000 are edible fungi, and more than 1,000 species of edible fungi have been clearly recorded in China [1]. Mushrooms are rich in nutrients and functional active substances, including polysaccharides, active proteins and peptides, phenols, ketones, terpenes and steroid compounds [2]. Among them, polysaccharides are one of the main active ingredients of edible fungi. Modern research shows that edible fungi polysaccharides have a variety of biological activities, including antioxidant, anti-tumor, anti-inflammatory, immunomodulatory, hypoglycemic, intestinal flora regulation, antiviral and other activities [3]. Edible fungi polysaccharides are used as functional active ingredients to prevent and treat various chronic diseases and are very beneficial to human health. The author systematically reviews the efficacy of various typical edible fungi polysaccharides and the research progress of their application in functional foods, biomedicine and cosmetics, in order to provide reference and scientific basis for the further development and utilization of edible fungi and their polysaccharides.
Typical edible fungus polysaccharides and their effects
Ganoderma lucidum polysaccharides
Ganoderma lucidum is a famous edible and medicinal fungus with a history of thousands of years of application. The Pharmacopoeia of the People’s Republic of China [4] records that Ganoderma lucidum is “sweet and neutral. It enters the heart, lung, liver and kidney meridians” and has the effects of “tonifying qi and calming the mind, relieving cough and relieving asthma”. Polysaccharides are one of the main components of Ganoderma lucidum to exert biological activity. At present, Ganoderma lucidum polysaccharides are mainly derived from spore powder, fruiting bodies, mycelium and Ganoderma lucidum fermentation liquid. Ganoderma lucidum polysaccharides have antioxidant, anti-tumor, hypoglycemic, anti-inflammatory, liver protection, immunomodulatory and neuroprotective effects. Ganoderma lucidum polysaccharides show good antioxidant activity by scavenging excess free radicals, increasing antioxidant enzyme activity and reducing peroxides [5]. Liu Yang et al. [6] found that Ganoderma lucidum polysaccharides can significantly promote the cellular immune response of 4T1 tumor-bearing mice to HER2/neu plasmid DNA vaccine and have a significant immune-promoting effect on mice. In addition, Wu Ruiting et al. [7] found that black Ganoderma lucidum polysaccharide can significantly regulate blood glucose, blood lipids and insulin in diabetic rats induced by streptozotocin (STZ), and its mechanism may be through changing the structural composition of microorganis Ganoderma lucidumms in the rat intestine. Dilisoyer Yalekunmu et al. [8] pointed out that Ganoderma lucidum polysaccharide may play a radiation protection role by improving the expression of relevant potential biomarkers and their related metabolic pathways in the thymus of radiation model mice.
Cordyceps polysaccharide
Cordyceps polysaccharide is a highly branched galactomannan. Studies have found that Cordyceps polysaccharide plays an important role in improving immune function, anti-cancer, sterilization, lung protection, and kidney benefit. Jiang Yuzhen et al. [9] optimized the enzyme-assisted hot water extraction of Cordyceps militaris polysaccharide by response surface methodology and found that the DPPH free radical scavenging rate of Cordyceps militaris polysaccharide reached 88.06%, and the hydroxyl free radical scavenging rate was 78.80%, both of which were dose-dependent, proving that Cordyceps militaris polysaccharide has good antioxidant activity. Yao Yanting et al. [10] obtained Cordyceps militaris polysaccharide fraction (SeCMP0.2) by water extraction and alcohol precipitation combined with anion exchange chromatography column separation and purification. The results showed that SeCMP0.2 can activate mitogen-activated protein kinase and nuclear factor-κB signaling pathways, upregulate the expression of inducible nitric oxide synthase, tumor necrosis factor-α, interleukin-6, and interleukin-1β mRNA, and exert its immunomodulatory effect. Liu Guangqing et al. [11] extracted Cordyceps flower polysaccharide by water extraction and alcohol precipitation combined with Sevag method and found that Cordyceps flower polysaccharide can significantly reduce the expression levels of tumor necrosis factor α, nuclear factor κB, etc. in CCl4-induced mouse liver, significantly downregulate serum ALT and AST levels, alleviate CCl4-induced liver damage, inhibit hepatocyte apoptosis, and provide a reference for the treatment of liver fibrosis.
Phellinus igniarius polysaccharide
Phellinus igniarius can promote diuresis and stop bleeding, improve internal organs. “Compendium of Materia Medica” [12] records that “Mulberry parasite benefits the five internal organs, promotes intestinal and stomach qi, and removes toxic qi.” As one of the main bioactive components of Phellinus igniarius, polysaccharides have been verified to have multiple biological activities such as anti-tumor, immunomodulatory, anti-inflammatory, and antioxidant. Zhang Kunlin et al. [13] found that Phellinus igniarius polysaccharides can enhance the activity of embryonic alkaline phosphatase secreted by RAW-BlueTM cells and their macrophage phagocytosis, proving that Phellinus igniarius polysaccharides have immune activity. Miao Yunping et al. [14] used Phellinus igniarius polysaccharides to treat rat ulcerative colitis induced by 2,4,6-trinitrobenzenesulfonic acid (TNBS). The results showed that Phellinus igniarius polysaccharides could significantly reduce the expression levels of NF-κB p65 protein and IL-6 and TNF-α mRNA in the rat colon. The therapeutic effect on rat ulcerative colitis was obvious, which may be related to regulating the secretion of inflammatory cytokines, anti-oxidation and inhibiting the expression of NF-κB. Li Yuanyuan et al. [15] prepared crude polysaccharide from mycelium of Phellinus igniarius by water extraction and alcohol precipitation, obtained fractions (IPS45-W) by distilled water elution, and found that it could inhibit the growth of HepG2 cells (the inhibition rate was 57% at a concentration of 50 μg/mL) through in vitro cytotoxicity and cell viability tests, proving the anti-tumor activity of crude polysaccharide from mycelium of Phellinus igniarius. Yu Qiuju et al. [16] found that mulberry linterus polysaccharide has good antioxidant and tumor inhibition activity through in vitro activity tests.
Grifola frondosa polysaccharide
The polysaccharide components extracted from Grifola frondosa fruiting bodies have biological activities such as antioxidant, anti-tumor, radiation protection and immunomodulatory. Xu Yingying [17] found that Grifola frondosa polysaccharide component (GFP1) has a significant inhibitory effect on the proliferation of lung cancer cells H1975, and GFP1 may achieve anti-tumor effects by inhibiting the Ras/Raf/MEK/ERK signaling pathway. Tang Mengting [18] used the Illumina system to analyze the structure and composition of the intestinal flora after fermentation of the culture medium and found that the components of Grifola frondosa polysaccharide (GFP-H, GFP-L) can significantly promote the proliferation of Bifidobacterium and Bacteroides in the intestine, and have a beneficial effect on the health of the intestinal flora. Quan Shulin et al. [19] found that Grifola frondosa polysaccharide GFP-A1 can significantly improve the exercise endurance of exercise mice and has an anti-fatigue effect. Liu Yonghua et al. [20] found that Grifola frondosa polysaccharide can significantly increase the content of superoxide dismutase (SOD) and glutathione peroxidase (GSH-Px) in rats with gastric ulcer model, and reduce the levels of gastrin (GAS), motilin (MTL), somatostatin (SS), and malondialdehyde (MDA), further proving the role of Grifola frondosa polysaccharide in reducing gastrointestinal hormone secretion and improving antioxidant levels.
Agaricus blazei polysaccharide
Studies have found that Agaricus blazei polysaccharide has biological activities such as anti-cancer, antioxidant, hypoglycemic, diabetes improvement, and cholesterol reduction. Pu Chao et al. [21] treated CHON-001 cells with 10 ng/mL IL-1β to establish a chondrocyte inflammation model and explored the protective effect of Agaricus blazei polysaccharide on interleukin 1β (IL-1β)-induced chondrocyte CHON-001 injury. The results showed that Agaricus blazei polysaccharide could significantly reduce the apoptosis rate and Bax protein level of the chondrocyte inflammation model, and pointed out that IL-1β-induced chondrocyte inflammation and apoptosis were inhibited by upregulating the expression level of miR-382-3p. In addition, Zhong Pupeng et al. [22] found that Agaricus blazei polysaccharide also has a certain effect on intestinal barrier protection. They pointed out that Agaricus blazei polysaccharide can significantly increase the relative expression of Faecalibacterium prausnitzii, Lactobacillus and Bifidobacterium 16SrRNA genes in rats with ulcerative colitis induced by sodium dextran sulfate (DSS), and significantly reduce the relative expression of Escherichia coli 16SrRNA genes, indicating its important role in preventing acute colitis.
Hericium erinaceus polysaccharide
Hericium erinaceus is flat in nature, sweet in taste, rich in nutrition, helps digestion and benefits the five internal organs. Hericium erinaceus is mostly used to improve peptic ulcers, chronic gastroenteritis, loss of appetite and other diseases, and also has a certain inhibitory effect on cancer. Hericium erinaceus contains polysaccharide, which is called “hericium erinaceus polysaccharide”, which has multiple biological activities such as antioxidant, immunomodulatory, regulating intestinal microbiota, lowering blood sugar, anti-tumor, and gastric protection. Ma Zheng et al. [23] studied the effects of Hericium erinaceus polysaccharide on the recovery of physical activity-induced fatigue and antioxidant capacity in the elderly and found that supplementing Hericium erinaceus polysaccharide in the elderly can significantly improve their aerobic capacity and blood rheology, promote fatigue recovery, and improve antioxidant capacity. Niu Meilan et al. [24] showed that Hericium erinaceus polysaccharide can significantly reduce the levels of TNF-α, IL-6, and IFN-r in the serum of rats with chronic non-bacterial prostatitis, reduce the levels of immunoglobulins IgA and IgG in the serum, further alleviate chronic non-bacterial prostatitis in rats, and delay the development of the disease. Li Xinghai et al. [25] explored the anti-tumor activity and mechanism of Hericium erinaceus polysaccharide, mainly by culturing gastric adenocarcinoma cells and setting up a Hericium erinaceus polysaccharide addition group. It was found that Hericium erinaceus polysaccharide combined with transforming growth factor-β1 can effectively inhibit the proliferation of gastric adenocarcinoma cells BGC-823 and SGC7901 by negatively regulating p-AKT and p-PI3K, thereby further inhibiting the number of cancer cell clones.
Poria cocos polysaccharide
Poria cocos is the dried sclerotium of the fungus Poria cocos of the Polyporaceae family. It is “neutral in nature, sweet and light in taste, and enters the heart, spleen, and kidney meridians. It is diuretic and dehumidifying, strengthens the spleen, and calms the mind. It is used for edema, oliguria, phlegm and fluid, palpitations, spleen deficiency, poor appetite, loose stools, restlessness, palpitations, and insomnia.” A large number of studies have confirmed that Poria cocos polysaccharide has antiviral and anti-tumor effects, and reduces the side effects of radiotherapy and chemotherapy. Liu Jiali et al. [26] found that Poria cocos polysaccharide can significantly reduce the release of NO and the rate of apoptosis in lipopolysaccharide-induced cat kidney cells, and believed that it may achieve the effect of cell inflammation protection by inhibiting the TLR4-NF-κB signaling pathway. This research result provides a new target and experimental basis for the verification of Poria cocos polysaccharide in the treatment of cat kidney cells. Zheng et al. [27] used hot water extraction to obtain Poria cocos polysaccharide (PCPP-1A) and found that PCPP-1A has excellent potential in stimulating the immune response of RAW264.7 cells and protecting AML-12 cells from radiation.
Tremella polysaccharide
Tremella is sweet in taste, neutral in nature, and belongs to the lung meridian. It has the effects of nourishing yin and moistening the lungs, replenishing qi and promoting fluid, and beautifying the skin. The tremella polysaccharide it contains has the effects of enhancing immunity, lowering blood sugar, and preventing and treating cardiovascular diseases. Xing Youzhe et al. [28] found that tremella polysaccharide can significantly promote the proliferation of mouse spleen lymphocytes, promote the secretion of IL-6, IL-10, IL-12 and IFN-γ, and exert its immunomodulatory effect by activating the PI3K/Akt and NF-κB signaling pathways of mouse spleen lymphocytes. Huang Xiaolan et al. [29] found that tremella polysaccharide can significantly reduce the levels of TNF-α and Il-6 in the serum of mice undergoing exhaustive exercise by regulating the Nrf2/HO-1 signaling pathway, significantly increase the levels of SOD and GSH-Px, and prolong the exhaustive swimming time of mice, proving that tremella polysaccharide improves oxidative damage caused by exercise by inhibiting oxidative stress. Tan Minying et al. [30] found that Tremella polysaccharide has the effect of inhibiting osteoarthritis, protecting cartilage tissue to a certain extent, and resisting cell apoptosis.
Black fungus polysaccharide
Black fungus is “neutral, sweet, smooth, and enters the stomach and large intestine meridians”. It is known as the “intestinal scavenger”. It mainly relies on its detoxification, lipid-lowering, intestinal motility promotion, and metabolism enhancement. Black fungus is rich in antioxidants and has rich biological activities. Black fungus polysaccharide has multiple biological activities such as antioxidant, lipid-lowering, immune enhancement and anti-tumor. Liu Nannan [31] extracted and purified two black fungus polysaccharide components (AAP-1 and AAP-2) by hot water extraction-ethanol precipitation method, and used C57BL/6J male mice to construct a type 2 diabetes model for in vivo hypoglycemic activity research. It was found that both black fungus polysaccharide components can activate the PI3K/AKT signaling pathway, reduce insulin resistance, reduce oxidative stress damage, and improve glucose and lipid metabolism to achieve the effect of lowering blood sugar. Li Xue et al. [32] prepared phosphorylated black fungus polysaccharide liposomes by phosphorylation modification of black fungus polysaccharide, which significantly improved its scavenging rate of DPPH free radicals, proving its antioxidant capacity in vitro. It was also found that the encapsulation of liposomes could effectively prevent the oxidation of phosphorylated black fungus polysaccharide. Zhang Tingting et al. [33] found that black fungus polysaccharide could significantly increase the abundance of Rhodococcus or non-digestive saccharobacterium in the fecal flora of SD rats, significantly reduce the abundance of pro-inflammatory bacteria Prevotella, and alleviate hepatitis and other diseases caused by a high-fat diet. In addition, black fungus polysaccharide could significantly increase the content of short-chain fatty acids in the cecum of SD rats. This study provides ideas for black fungus polysaccharide to improve blood lipid metabolism.
Application Status of Typical Edible Fungi Polysaccharides
Application in Functional Foods
Edible fungi have unique texture and flavor. By adding edible fungi or edible fungi polysaccharides, the flavor and taste of food can be improved. At the same time, due to the multiple biological activities of edible fungi and their polysaccharides, they can be used as food raw materials to develop new foods, which can not only affect the texture and nutrition of food, but also give new functional characteristics to new products. my country’s national standard GB 16740-2014 General Standard for Health (Functional) Foods points out that polysaccharides can be used as the effective ingredients of functional health foods. Among the health foods with polysaccharides as the main ingredients that have been launched in my country, only 14 types are recorded by the National Special Food Information Query Platform, including three dosage forms: capsules, tablets, and oral liquids, covering a variety of Chinese herbal polysaccharides such as astragalus polysaccharides and ganoderma lucidum polysaccharides. For example, Qingfu Ning capsule, Yangpeptide Lingzhi capsule, Tangzhi capsule, and Propolis Lingzhi capsule are developed with Lingzhi polysaccharide as the active ingredient, Seabuckthorn Tianbao soft capsule is developed with Poria polysaccharide as the main biologically active ingredient, and Cordyceps militaris Lingzhi Milk thistle capsule is developed with Cordyceps militaris polysaccharide as the main active ingredient. In 2001, the National Health Commission (formerly the Ministry of Health) released a list of 11 fungal species that can be used in health foods [34]. It can be seen that the potential efficacy of edible fungi polysaccharides and functional foods made from edible fungi polysaccharides need to be further developed.
Foods with a high glycemic index (GI) can cause a sharp increase in blood sugar levels and lead to the occurrence of acute metabolic diseases. The digestion characteristics of starch are closely related to the postprandial blood sugar response. Therefore, controlling the absorption rate of starch to control the blood sugar level of the human body has become an effective clinical nutrition approach. It is reported that edible fungi polysaccharides can improve insulin resistance and effectively reduce starch content [35-36]. Su Jing[37] used Tremella fuciformis polysaccharide as an active ingredient (added at 0.4%), maltitol and isomalt as raw materials, and gelatin as a gelling agent to develop a gel soft candy. Liu et al.[38] studied the effects of Pleurotus eryngii and lemongrass powder on the physical and chemical properties, sensory quality and digestibility of bread. The results showed that Pleurotus eryngii powder, as a high-value ingredient, can improve the nutritional quality of bread and reduce its GI. Sun Fan[39] developed Pleurotus eryngii immune yogurt with the activity of improving human immunity based on the immune-enhancing activity of Pleurotus eryngii polysaccharide. The formula is 0.1% Lactobacillus bulgaricus and Streptococcus thermophilus, 0.7% Pleurotus eryngii polysaccharide, 7% white sugar, and 0.2% pectin. This product has the inherent flavor of yogurt and the appropriate Pleurotus eryngii flavor, and the texture and appearance meet the product quality standards. Li Guangfu[40] added ganoderma lucidum polysaccharide, ganoderma acid, tea mushroom polysaccharide and enoki mushroom polysaccharide to yogurt to develop edible fungus polysaccharide yogurt, and found that the in vitro antioxidant and anti-aging effects of edible fungus polysaccharide yogurt were significant. In addition, edible fungus polysaccharides are also used to produce functional foods such as health bread[41], noodles[42], and beverages[43], playing an important role in improving product flavor and nutrition.
Application in the field of biomedicine
Due to the potential biological activity of edible fungus polysaccharides, they are often used to develop anti-cancer drugs, anti-diabetic drugs, antiviral drugs, prebiotics, immunosuppressive (regulatory) agents, etc.[44].
Atpresent, many edible fungus polysaccharides have been developed into injections, tablets, granules, oral liquids, decoction pieces, syrups, etc. suitable for clinical application. The edible fungus polysaccharide medical products approved by the National Medicine Standards involve a wide range of types. The drug products developed with edible fungus polysaccharides that have been used in clinical practice include Ganoderma lucidum injection, carboxymethyl Poria cocos polysaccharide injection, Polyporus umbellatus polysaccharide injection, Lentinula edodes polysaccharide tablets, Hericium erinaceus tablets, polysaccharide protein decoction pieces, Ganoderma lucidum dew, compound Ganoderma lucidum dry syrup, Yunzhi Gantai granules, Liangjun tablets, Ganbi Fu, Cordyceps essence, etc.
In addition to being used alone, edible fungus polysaccharides can be compounded with other active ingredients. Chen Shuping et al. [45] compounded Tremella fuciformis, Poria cocos, Hericium erinaceus, Lentinula edodes and Dictyophora indica polysaccharides and found that the compound polysaccharides can regulate the immune activity of macrophage RAW264.7, which provides ideas for the development of compound polysaccharide products. Yang Yingge et al. [46] produced polysaccharides by solid bidirectional fermentation of Ganoderma lucidum and Astragalus membranaceus residues and found that when the polysaccharide dose was 25 mL/kg, the tumor inhibition rate of human colon cancer HCT116 transplanted into nude mice was 55.99%.
Application in the field of cosmetics
According to the ancient medical book “Imperial Medicine Prescriptions” [47], edible fungus polysaccharides play an important role in beauty. It is recorded in the ancient Chinese medicine book “Taiping Jing Hui Fang” [48] that Tremella fuciformis was used for bath therapy in the Tang Dynasty. A large number of studies have confirmed that edible fungi are popular raw materials that can be used in both cosmetics and oral beauty products. Taking Ganoderma lucidum as an example, modern scientific tests have proved that Ganoderma lucidum contains ingredients such as glucosamine and polysaccharides, is rich in antioxidants, and can play a good role in skin care products [49].
Tremella fuciformis polysaccharides have the unique effect of removing freckles, improving skin and increasing moisture, and are widely used in cosmetics. The herbal skin care products (Jungwang Cream, Essence Oil) launched by the East Beast in 2021 mainly contain Ganoderma lucidum triterpenes and Ganoderma lucidum polysaccharides, which play their anti-inflammatory and antioxidant effects. The medicinal lamellae series developed by Ximuyuan for oily and sensitive skin is added with medicinal lamellae skin care extracts. The polysaccharides contained in it mainly play their antioxidant, soothing and repairing effects, and are more suitable for sensitive skin. The pine mushroom series skin care products first recommended by the Xiwuji brand are added with pine mushroom extracts. The polysaccharides contained in them mainly play antioxidant and repair effects.
Outlook
Currently, most of the research on edible fungus polysaccharides focuses on biological activity. The knowledge about the structure-activity relationship and molecular mechanism of different edible fungus polysaccharides is still limited. Therefore, it is still necessary to broaden the research on the biological activity of different types of edible fungi and their polysaccharides, and to explore the molecular structure and potential biological activity of edible fungus polysaccharides in depth. At the same time, through chemical modification of edible fungus polysaccharides, such as carboxylation and methylation, its scope of use can be broadened and its biological activity can be enhanced.
Different edible fungus polysaccharides have different biological activities. Therefore, according to the mechanism of action of their biological activities, polysaccharides can be compounded in different proportions, mixed into mixtures, or combined with drugs to exert synergistic effects and double their efficacy. In order to make edible fungus polysaccharides more efficient in medicinal treatment, their dosage forms can be enriched, and the solubility and bioavailability of polysaccharides can be significantly improved.
Combining different technologies and disciplines, such as computational biology, bioinformatics, and omics, to explore the potential value of different edible fungus polysaccharides, provide reference and guidance for understanding and broadening the biological activity of edible fungus polysaccharides. In specific applications, optimize and upgrade extraction and purification technologies to convert nutrients and polysaccharides contained in edible fungi into active ingredients that can be used in cosmetics, medicine and food, and truly become high-efficiency active raw materials.
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