Research advances in fungal polysaccharides: production, extraction, characterization, properties, and their multifaceted applications

Abstract

Fungal polysaccharides have demonstrated significant biological potential, including immune stimulation, antioxidant activity, anticancer properties, and anti-inflammatory effects. These properties hold considerable promise for innovative applications across various fields. This study provides a brief review of current findings, based on literature published over the past 15 years on fungal polysaccharides. This includes the production process and various extraction methods, highlighting their distinct advantages and limitations. Additionally, we summarize techniques for purification and characterization, elucidating their biological properties and practical applications in medicine, pharmacology, the food industry, agriculture, and environment. Global patent trends related to fungal polysaccharides are also reviewed. Finally, we discuss challenges and future perspectives related fungal polysaccharides. This article offers valuable insights and enhances the understanding of fungal polysaccharides for researchers, paving the way for further research and applications.

1 Introduction

Fungi have played an important role in human history for millions of years. Terrestrial fossil evidence indicates that fungi first appeared around 400 million years ago (Brundrett, 2002). An updated estimation of the global fungi species has been revised and raised to 2.5 million worldwide according to updated molecular data, but only about 155,000 species have been officially identified and described, highlighting the ongoing research in this field (Antonelli et al., 2023). Belonging to the kingdom of Fungi, they are vital in decomposing organic matter within nutrient cycles and are associated with various organisms that help maintain ecosystems (Niego et al., 2023). Certain edible fungi serve as a crucial food source, providing high nutritional value with proteins, vitamins, and essential minerals, making them suitable for consumption to enhance health. Additionally, some of them exhibit medicinal properties, including antimicrobial, anticancer, and immunomodulatory effects (Valverde et al., 2015). In the industrial sector, extracts from edible fungi are utilized in food and beverage production, such as functional foods, brewing beer, fermenting wine, and making bread (Lorenzo et al., 2018). The cultivation of edible fungi holds significant economic value, providing farmers with income opportunities due to the high market demand (Zhang et al., 2014). Edible fungi can also serve as meat substitutes and are a rich protein source in various dishes, making them an attractive option for those reducing meat consumption or following a vegetarian diet (Ayimbila and Keawsompong, 2023). Interestingly, many edible fungi produce bioactive compounds, which are chemicals that exert biological effects on the human body or other living organisms. Examples of bioactive compounds derived from fungi include polysaccharides, triterpenoids, ergosterol, polyphenols, flavonoids, vitamins, minerals, and antibiotic compounds (Valverde et al., 2015; Antonelli et al., 2023).

One of the most popular and widely recognized fungal bioactive compounds is polysaccharides. Fungal polysaccharides are complex carbohydrates composed of multiple monosaccharide units linked by glycosidic bonds, and they possess biological properties (Hamidi et al., 2022; Stoica et al., 2023). From 2000 to 2024, a search using the keyword “fungal polysaccharide” retrieved 8,138 titles of documents published over the last 25 years in the Scopus database (https://www.scopus.com, accessed 20 March 2025). It was found that the trend in research on fungal polysaccharides is expected to increase in the future (Figure 1A).

Figure 1

Additionally, the majority of applications for fungal polysaccharides have been reported in the fields of biochemistry, genetics, and molecular biology (25%), followed by immunology and microbiology (17%), medicine (11%), agricultural and biological sciences (10%), and chemistry (7%) (Figure 1B). In this study, we summarize current findings on fungal polysaccharides, including type, production processes, extraction methods, characterization, biological properties, and their applications in various fields from 2010 to 2024 (a 15-year period). Moreover, we summarize trends in patents related to fungal polysaccharides. The overview of this review article is presented in Figure 2.

Figure 2

2 Source of fungal polysaccharides

Fungal polysaccharides can be produced by yeasts (single cells), filamentous fungi (mycelia), and macrofungi (mycelia and fruiting bodies). Notably, basidiomycetous fungi have attracted significant research attention, particularly for their polysaccharide production and diverse industrial applications. Additionally, the diversity of fungal species and the types of polysaccharides they produce highlight the significant potential of fungi as a source of bioactive compounds for various applications. Examples of fungal species reported to produce polysaccharides are compiled and presented in Table 1.

3 Types of fungal polysaccharides

Fungal polysaccharides are categorized primarily into two types based on their composition: homopolysaccharides, composed of a single type of monosaccharide, and heteropolysaccharides, which include various monosaccharides (Murphy et al., 2023a). Homopolysaccharides are polysaccharides composed of only one type of monosaccharide unit repeated in long chains, e.g., glucans (polymers of glucose), chitin (a polymer of N-acetylglucosamine), and mannans (polymers of mannose) (De Felice et al., 2020). Conversely, heteropolysaccharides are polysaccharides composed of two or more different types of monosaccharide units, e.g., galactomannans (composed of mannose and galactose), pectins (composed of galacturonic acid and rhamnose), and glycosaminoglycans (e.g., hyaluronic acid) (Kadooka et al., 2023). Additionally, fungal polysaccharides are broadly divided into two forms based on their cellular location: endopolysaccharides (EnPs) or intracellular polysaccharides, and exopolysaccharides (ExPs) or extracellular polysaccharides (Zhang et al., 2016). These types differ considerably in terms of structure, biological activities, and extraction methods, offering distinct benefits and applications in biotechnological fields.

Both fungal EnPs and ExPs are crucial for fungal growth and adaptation to environmental conditions. Fungal EnPs are located within cells and often have complex, branched structures. They are crucial components of cell walls or are stored in the cytoplasm, providing structural support to the cell. They are typically composed of chitin and monosaccharides such as glucose, mannose, and galactose (Wang et al., 2017b; Fernando et al., 2021). β-glucan is well-known fungal EnPs, particularly in fungi belonging to the basidiomycetes and ascomycetes groups. In contrast, fungal ExPs are secreted by cells into their surrounding environment, where they generally form linear or mildly branched structures. These structures are typically simpler than those of EnPs. Fungal ExPs contribute to the extracellular matrix, forming gels that facilitate adhesion and biofilm formation, and providing structural support and protection to the fungi under adverse conditions (Mohamed et al., 2024). Primarily composed of glucose, fructose, and other simple monosaccharides, these polysaccharides typically lack complex functional groups and are valued for their gelling and viscosity-enhancing properties (Wang et al., 2020a). Fungal ExPs are often characterized by favorable physical properties such as high-water solubility and significant viscosity, though some types may be insoluble or display unique physical traits based on their structural makeup (Guo et al., 2017). Furthermore, extraction methods for fungal ExPs are usually more straightforward than those for fungal EnPs, as they do not require cell disruption (Jeong et al., 2013; Miao et al., 2020). The most well-known fungal polysaccharides, including both EnPs and ExPs (e.g., botryosphaeran, grifolan, lasiodiplodan, lentinan, pleuran, pullulan, schizophyllan, and scleroglucan), are listed in Table 2, and their chemical structures are presented in Figure 3.

Figure 3

4 Production process of fungal polysaccharides

The industrial-scale production of fungal polysaccharides from fungi is a process that requires precise control and high efficiency to ensure the desired quantity and quality of the product. The production process begins with the selection of fungal strains capable of producing large amounts of polysaccharides (Sánchez et al., 2015; Junior Letti et al., 2018). Additionally, inoculum preparation under optimal conditions is crucial to maximize fungal biomass before transferring to the fermentation system. The choice of fermentation method is critical, with the primary approaches being submerged fermentation and solid state fermentation (Bakratsas et al., 2021). Solid state fermentation occurs on solid substrates or non-soluble support materials and is particularly suitable for certain fungal fruit bodies (Junior Letti et al., 2018), utilizing agricultural residues as growing substrates (Pérez-Chávez et al., 2019). In contrast, submerged fermentation, a common industrial method, involves liquid-based fermentation requiring the careful control of parameters such as pH, temperature, agitation, and aeration (Sánchez et al., 2015). The enhancement of polysaccharide production from both fermentation processes requires optimal conditions for fungal growth specific to each fungal species and strain (Montoya et al., 2013).

4.1 Solid state fermentation

Solid state fermentation (SSF) involves the cultivation of fungi on solid substrates or low-moisture materials, typically derived from agricultural or industrial residues such as wheat straw, rice straw, coffee pulp, wood logs, sawdust, bran, husks, and sugarcane bagasse (Junior Letti et al., 2018; Yafetto, 2022), due to the ability of fungi to break down and utilize lignocellulosic materials as a source of nutrients to support their growth and development. Additionally, the selection and utilization of agricultural or industrial residues in SSF is contingent on their availability in each country. SSF is particularly suitable for cultivating fungi to produce their fruiting bodies, commonly known as mushrooms (Junior Letti et al., 2018). The cultivation of mushrooms using SSF has been studied worldwide because it allows the use of various agricultural or industrial residues as substrates, promotes the recycling of these residues into materials for fungal growth, and facilitates their conversion into fruiting bodies (Grimm and Wösten, 2018; Junior Letti et al., 2018). For example, Sánchez and Montoya (2020) found that the cultivation of Pl. ostreatus, Le. edodes, and Tr. versicolor under SSF conditions using oak sawdust revealed that the highest polysaccharide production was recorded for the growing substrate of Tr. versicolor (96.09 mg/g of solid substrate), followed by Pl. ostreatus (90.78 mg/g) and Le. edodes (87.45 mg/g). The research conducted by Xu et al. (2019) found that the optimal conditions for polysaccharide production from Co. militaris using rice under SSF resulted in a maximum yield of 68.3 mg/g of dry substrate. Shi et al. (2012) used soybean curd residue as a substrate in SSF to produce polysaccharides from the fruiting bodies of F. velutipes, yielding 106.74 mg/g of dried fruiting bodies. Additionally, Le. edodes fruiting bodies cultivated under SSF using beechwood sawdust yielded a maximum polysaccharide production of 67.33 mg/g of dry weight (Reza et al., 2024). SSF of Gr. frondosa using corn bran and oak sawdust resulted in a polysaccharide yield of 60.5 mg/g of dry weight (Montoya Barreto et al., 2011). Thus, SSF can be used for the production of fungal polysaccharides, demonstrating its economic viability as well as its potential for further pilot research and large-scale industrial applications. Remarkably, the carbon/nitrogen (C/N) ratio and the composition of cellulose, hemicellulose, and lignin in agricultural or industrial substrates vary across different types, making them crucial factors in SSF. Large-scale SSF, particularly in industrial settings, remains challenging due to the need to carefully control factors such as pH, heat and mass transfer, water activity, fungal strain selection, substrate heterogeneity, C/N ratio, and optimal moisture levels for fungal growth.

4.2 Submerged fermentation

Submerged fermentation involves the cultivation of fungal mycelium in a liquid medium, where nutrients are dissolved, and agitation is used to facilitate aeration within the fermenter or bioreactor. The primary advantages of submerged fermentation include efficient oxygen transfer and homogeneous distribution of the liquid medium. However, this technique requires meticulous control of various factors such as nutrient (carbon and nitrogen sources), temperature, aeration, agitation, pH, medium composition, and the type and amount of inoculum. Consequently, submerged fermentation is a reproducible technique for the continuous cultivation of fungal mycelium, enabling controlled production of metabolites (Elisashvili, 2012). Nonetheless, prolonged cultivation in liquid media can lead to increased medium viscosity due to the growth and accumulation of fungal mycelium, potentially impacting oxygen distribution, carbon dioxide removal, and product dilution (Bentil et al., 2018). Recently, edible fungi, particularly those from the basidiomycetes and ascomycetes genera, including Agaricus, Cordyceps, Ganoderma, Lentinus, and Pleurotus, have been successfully cultivated using bioreactors (Bakratsas et al., 2021). For example, Wang et al. (2019a) achieved the highest extracellular polysaccharide yield (5.713 g/L) of Co. militaris in a 5-L bioreactor using glucose and yeast extract as carbon and nitrogen sources, with conditions of 25°C, 150 rpm shaking speed, and 1.5 vvm aeration. The most suitable submerged fermentation conditions for Le. crinitus to achieve the highest yield of ExPs at 0.65 g/L in a 5-L bioreactor, using Kirk’s liquid medium, were 30°C, pH 4.5, 300 rpm stirring, and a 1.5 vvm aeration rate for four days (López-Legarda et al., 2020). The fed-batch submerged fermentation process for EnP production from G. lucidum was successfully scaled up in stages, from 7.5 L to 20 L, and finally to a 200-L stirred-tank reactor, where maintaining a low impeller tip speed of 1.234 m/s resulted in a maximum EnP production of 4.74 g/L (Tang et al., 2011). Moreover, Wu et al. (2013) studied the effect of different light wavelengths on ExPs production from Pl. eryngii in submerged cultivation and found that ExPs production was highest under blue light (455 mg/L) condition, compared to green (425 mg/L), red (217 mg/L), yellow (314 mg/L), white (50 mg/L), and dark (59 mg/L) conditions. In a recent study by Bakratsas et al. (2024), the cultivation of Pl. ostreatus was successfully scaled up in a 3.5 L bioreactor under optimal conditions, resulting in a maximum biomass of 12.6 g/L, the highest ExPs production of 3.7 g/L.

Overall, previous studies highlight several key factors that significantly enhance the production efficiency of polysaccharides from selected fungi. Among these factors, the selection of suitable carbon sources, such as glucose and yeast extract, plays a crucial role in stimulating fungal growth and the production of desired compounds. Optimal concentrations of nutrients should be maintained to support maximum biomass development and polysaccharide yield. Additionally, controlling fermentation parameters such as temperature, pH, agitation, and aeration is crucial for optimizing polysaccharide production. Furthermore, the use of specific light wavelengths during cultivation has been shown to enhance the production of fungal ExPs.

5 Extraction techniques for fungal polysaccharides

The processes for obtaining fungal EnPs and ExPs involve several steps, as shown in Figure 4. The extraction of polysaccharides from fungi primarily differs in the locations of the polysaccharides and the methods employed for extraction. For EnP extraction, the process begins with harvesting the fungal mycelia or fruiting bodies, followed by cell disruption, which can be achieved either chemically or mechanically to break the cell walls (Wang et al., 2015b; Liu et al., 2022b). On the other hand, ExP is secreted outside the fungal cells and accumulates in the liquid culture or surrounding fluid. The extraction of ExPs typically involves filtering or separating the liquid culture from the fungal biomass. The ExP extraction is simpler because it does not require cell disruption. As a result, ExPs tend to be purer since they are directly extracted from the fungal supernatant (Jeong et al., 2013). However, EnP extraction could be contaminated by other intracellular substances, requiring further purifying processes.

Figure 4

5.1 Extraction of fungal exopolysaccharides

Generally, submerged fermentation is used for fungal ExP production. The extraction of ExP typically involves straightforward methods. Fungal ExP is separated from the liquid culture medium by first removing the fungal mycelia and cells through filtration or centrifugation and then collecting the supernatant (Stoica et al., 2023). The ExP present in the supernatant is then precipitated with organic solvents (e.g., acetone, ethanol, propanol, or isopropyl alcohol) under cold conditions (Maziero et al., 1999; Elisashvili et al., 2009). After precipitation, centrifugation or filtration is used to collect the crude fungal ExP.

5.2 Extraction of fungal endopolysaccharides

The EnP extraction from fungi is a more complex procedure than ExP extraction and requires specific methods to isolate EnP from the fungal cells (Jeong et al., 2013). After the fungus cells or fruiting bodies have been collected, the cell walls must be broken down in order to release the EnP. This can be achieved through chemical techniques, such as using lysozyme enzymes or solvents that degrade the fungal cell walls, or mechanical techniques like grinding or ultrasonic waves to break down the cell walls (Ferraboschi et al., 2021; Larrañaga-Ordaz et al., 2022). After the cell walls are disrupted, the extraction process can be carried out using a variety of techniques, including hot water extraction, alkaline- or acid-extraction, ultrasonic-assisted extraction, microwave-assisted extraction, enzyme-assisted extraction, or subcritical liquid extraction. Following extraction, EnP is collected, purified, and freeze-dried for subsequent application (Leong et al., 2021; Zhao et al., 2023b). Examples of fungi used for EnP extraction through various techniques are shown in Table 3.

5.2.1 Hot water extraction

The hot water extraction method is a popular technique for EnP extraction from fungi due to its simplicity, safety, and the availability of common materials (Dai et al., 2023). The fungal cells or fruiting bodies are collected, dried, and ground into a fine powder to enhance extraction efficiency. By boiling the powdered fungal samples in hot water at temperatures between 50°C and 121°C for 30 min to 10 h, the cell walls are broken down, releasing the EnPs (Saetang et al., 2022). The resulting solution is then filtered to remove impurities, and alcohol is added to concentrate and precipitate the EnP. The EnP is subsequently collected and dried (Leong et al., 2021). This method is suitable for various fungi and can be carried out in a standard laboratory, provided that temperature and extraction time are carefully controlled to achieve optimal results (Parniakov et al., 2014; Leong et al., 2021). The advantages of this method include the absence of toxic chemicals and low cost; however, some loss of EnP may occur due to heat, and the extraction may be incomplete if the cell walls are not fully broken down.

5.2.2 Alkaline or acid extraction

The alkaline or acid extraction technique involves using alkaline or acidic solutions to break down the cell walls of fungi and release the EnP. In alkaline extraction, an alkaline solution, such as sodium hydroxide (NaOH) or potassium hydroxide (KOH), is used to break down the fungal cells. In contrast, an acidic solution, such as hydrochloric acid (HCl) or ammonium oxalate [(NH₄)₂C₂O₄], is used in acid extraction. After the extraction process, the extracted samples must be pH-neutralized before filtration and precipitation of the polysaccharide (Yi et al., 2020; Leong et al., 2021). Alkaline extraction, using NaOH as the extracting agent, is more commonly employed than acid extraction for isolating fungal polysaccharides. NaOH solutions are typically used in concentrations ranging from 0.1 M to 1 M, with extraction ratios varying from 1:5 to 1:20 (w/v) (Palacios et al., 2012; Yang et al., 2019). Nevertheless, acid extraction methods remain an alternative, with previous reports highlighting the use of 0.1 M HCl (1:10 w/v) and 1% (w/v) (NH₄)₂C₂O₄ for extraction (Sermwittayawong et al., 2018; Zhang et al., 2022a). This technique is highly effective, especially for extracting EnPs from fungi with robust cell walls. However, careful control of extraction conditions is necessary to prevent EnP degradation and avoid side effects from the use of highly concentrated alkaline or acidic solutions (Wang et al., 2015b, c).

5.2.3 Ultrasonic-assisted extraction

Ultrasonic-assisted extraction is a method that uses ultrasonic waves to enhance the extraction of EnP from fungal cells. This technique involves using high-frequency sound waves to create cavitation, a phenomenon where small bubbles in the liquid rapidly collapse, producing energy that can break down the cell walls of mycelia or mushrooms and facilitate the release of EnP into the extraction solvent. The ultrasonic technique can be divided into two types: low-intensity and high-intensity ultrasound. The low-intensity ultrasonic technique uses frequency waves ranging from 5 to 10 MHz and energy levels of less than 1 W/cm². On the other hand, the high-intensity ultrasonic technique uses frequencies ranging from 20 to 100 kHz and higher energy levels from 10 to 1,000 W/cm². In comparison to the low-intensity technique, the high-intensity ultrasonic method has a higher destructive capability (Charoux et al., 2017; Li et al., 2024a). In general, EnP extraction using this technique typically involves ultrasonic treatment for 14 to 180 min at temperatures ranging from approximately 25°C to 95°C (Ke, 2015). The process begins by preparing the fungal mycelia or mushroom samples, grinding or cutting them into small pieces, immersing them in an appropriate solvent, and applying ultrasonic waves to facilitate the extraction. Subsequently, the extracted sample is filtered to separate the solid residues, and the solution containing EnP is then precipitated and the EnP is harvested. The advantages of this technique include increased extraction efficiency and reduced solvent usage (Leong et al., 2021; Shen et al., 2023).

5.2.4 Microwave-assisted extraction

Microwave-assisted extraction is a technique that uses microwave energy to accelerate the extraction process. Microwave energy causes water molecules or solvents within the sample to vibrate, generating heat that helps break down the cell structure and release the desired compounds more efficiently (Hu et al., 2021). Key factors influencing EnP yield using this method include microwave power, typically ranging from 400 to 1200 W, extraction temperature, maintained between 85°C and 180°C, and extraction time, which ranges from 1 to 30 min (Zhao et al., 2014; Gil-Ramírez et al., 2019; Chuensun et al., 2021). This technique is a highly efficient and fast technique that reduces solvent and energy usage (Chen et al., 2013). However, it has limitations, including high initial costs, restrictions on solvent selection, risks of unwanted reactions, and challenges in controlling various parameters (Xu et al., 2018).

5.2.5 Enzyme-assisted extraction

Enzyme-assisted extraction is a technique that uses enzymes to extract EnP from fungi. This method leverages the ability of enzymes to break down the cell walls of fungi (Leong et al., 2021; Bains et al., 2023). Several cellulolytic enzymes and proteolytic enzymes have been used, including cellulase, papain, pectinase, protease, and trypsin (Yin et al., 2015; Bhotmange et al., 2017; Bains et al., 2023). The process begins with sample preparation, where the fungal materials are ground or milled into a fine powder. The powder is then mixed with an enzyme solution under optimized conditions to maximize enzyme activity, typically at a pH of 4 to 6, a temperature of 40°C to 60°C, and an incubation time of 30 to 180 min (Zhu et al., 2014; Yin et al., 2015; Ashfaque et al., 2017; Bhotmange et al., 2017). The enzyme is allowed to act for a period during which the cell walls are degraded, and the polysaccharides are released into the solution. After extraction, the solution is filtered, and the polysaccharides are precipitated and harvested (Qin et al., 2017; Bains et al., 2023). This technique offers advantages such as increased extraction efficiency and reduced use of harsh chemicals, making it an environmentally friendly method (Chen et al., 2013; Zhao et al., 2016b). However, it also has disadvantages, including high costs, enzyme stability that depends on environmental conditions, challenges in selecting appropriate enzymes, potentially longer extraction times, and the complexity of process optimization (Yin et al., 2011; You et al., 2013).

5.2.6 Subcritical water extraction

Subcritical water extraction is a technique that uses subcritical fluids to extract polysaccharides from fungi. A subcritical fluid is a liquid heated above its boiling point (100 to 374°C) but remains in liquid form due to the increased pressure (1 to 22.1 MPa) (Gbashi et al., 2017). This technique typically uses water, ethanol-water mixtures, or other solvents (ethanol, methanol, or acetone) under subcritical conditions for extraction (Herrero et al., 2006). The process begins by preparing the fungal material, which is then mixed with the subcritical fluid in a closed system where both temperature and pressure are carefully controlled (Morales et al., 2019; Leong et al., 2021). In subcritical conditions, the physical properties of the fluid, such as its solubility and diffusivity, change, which enhances the efficiency of polysaccharide extraction from fungal cells. After extraction, the solution is cooled and separated to isolate the desired polysaccharides, while the remaining fluid can be reused in subsequent processes (Zhang et al., 2022a). IPS extraction using this technique requires optimization of conditions, typically involving pressures ranging from 0.15 to 5 MPa, temperatures between 115°C and 210°C, and extraction durations of approximately 13 to 80 min (Luo et al., 2017; Li et al., 2018a; Zhang et al., 2019). This technique offers advantages, including shorter extraction times, reduced solvent usage, and better control over the quality of the extracted polysaccharides (Huber et al., 2021). Despite its benefits, this technique has some limitations. One challenge is the need for equipment that can withstand high pressure and temperature, resulting in higher installation costs. Furthermore, operating at high pressure requires specialized expertise to manage the process effectively (Yabalak et al., 2024).

Several previous studies have compared different extraction techniques to obtain the highest yield of fungal EnPs. For example, Sangthong et al. (2022) compared ultrasonic-assisted extraction (sonication at 95°C for 5 h), and microwave-assisted extraction (860 W for 25 min) with hot water extraction (95°C for 5 h) and found that hot water extraction yielded higher EnP levels from V. volvacea (15.58% of dried fruiting bodies) compared to microwave-assisted extraction (11.05%) and ultrasonic-assisted extraction (9.06%). Similarly, Chen et al. (2020) compared ultrasonic-assisted extraction (450 W for 20 min) and microwave-assisted extraction (550 W for 5 min) with hot water extraction (100°C for 2 h) and found that hot water extraction yielded the highest polysaccharide content from Sch. commune (8.26% of dried fruiting bodies), higher than both ultrasonic-assisted extraction (5.07%) and microwave-assisted extraction (4.98%). Additionally, Zhang et al. (2022b) conducted a comparative study on five extraction methods for isolating EnP from fruiting bodies of Tri. mongolicum, including hot water extraction (85°C for 3 h), ultrasonic-assisted extraction (300 W power at 60°C for 35 min), enzyme-assisted extraction (using 5% cellulase and 2% pectinase at pH 5 and 50°C for 100 min), alkaline extraction (0.1 M NaOH at 90°C for 1 h), and acid extraction (0.1 M HCl at 90°C for 1 h). Among these five methods, alkaline extraction yielded the highest polysaccharide content at 13.16% of dried fruiting body, followed by acid extraction (10.83%), hot water extraction (6.64%), enzyme-assisted extraction (5.87%), and ultrasonic-assisted extraction (4.41%). Therefore, the selection of an extraction method for polysaccharides from fungi depends on the characteristics of the fungal samples, the equipment, and the limitations of each extraction technique. Each extraction method has its own distinct advantages and limitations, as summarized in Table 4. Hot water extraction is a convenient and safe method, although it may result in partial loss of polysaccharides due to the heat and may not efficiently extract them if the fungal cell walls are not adequately disrupted. For fungi with hard cell walls, alkaline or acid extraction works quite well. However, extreme acidity or alkalinity must be avoided to prevent polysaccharide degradation. Ultrasonic-assisted and microwave-assisted extractions enhance efficiency by utilizing energy from sound waves or microwaves; however, these methods require careful control of energy and extraction time, as they are costly and may limit solvent selection. In addition, enzyme-assisted extraction is effective for breaking down fungal cell walls, but it requires costly enzymes and may involve longer process optimization times. Lastly, subcritical water extraction, which operates at temperatures below the boiling point of water, is efficient and fast; however, it is costly because it requires specialized equipment capable of withstanding high temperatures and pressures.

Interestingly, the combination of extraction techniques can enhance polysaccharide yields, resulting in higher yields compared to using a single technique. For instance, Zhang et al. (2023) demonstrated that the combination of microwave-assisted and ultrasonic-assisted extraction was the most effective method for extracting polysaccharides from D. indusiata, yielding 12.66%, compared to other methods, including ultrasonic extraction (11%), microwave extraction (10%), and hot water extraction (8.5%). A combination of pressure and hot water extraction was used to extract β-glucan from Len. sajor-caju under conditions of 140°C and 0.92 MPa for 40 min, resulting in a high yield of 3.20 g per 100 g of dry fruiting body (Sakdasri et al., 2022). Similarly, Tepsongkroh et al. (2023) applied high-pressure processing at 600 MPa for 10 min, combined with hot water extraction at 60°C for 2 h to extract polysaccharides from V. volvacea, resulting in a 12% increase in crude polysaccharide yield and a 20% increase in β-glucan yield. The combined extraction of polysaccharides from G. lucidum under optimal conditions (ultrasonic power of 240 W, enzyme concentration of 0.5 mg/mL, pH 7.9, solvent-to-material ratio of 50:1 mL/g, temperature of 55°C, and extraction time of 144 min) resulted in a yield of 3.72% of dried fruiting bodies (Hoa, 2018). The combination of enzyme-assisted extraction (1:1:1 ratio of papain, pectinase, and cellulase, a solvent-to-material ratio of 1:30, pH 5, a temperature of 48°C), and microwave-assisted extraction (440 W for 10 min) for the extraction of EnPs from Le. edodes resulted in a high yield of 9.79% of dried fruiting bodies (Yin et al., 2018). Recently, Yang et al. (2024d) found that the enzyme-assisted extraction technique, combined with ultrasonic-assisted extraction, effectively extracted polysaccharides from F. velutipes under optimal conditions (ultrasonic power of 150 W, snailase enzyme concentration of 1%, solvent-to-material ratio of 10:1 mL/g, temperature of 55°C, and extraction time of 90 min), resulting in a polysaccharide yield of 109.54 mg/g of dried fruiting bodies.

Thus, the selection of an extraction method of fungal polysaccharide must consider multiple factors, including the type of sample, extraction efficiency, cost, time, and environmental impact, in order to achieve the highest yield of fungal polysaccharides. Additionally, treatment temperature, solvent-to-sample ratio, and energy used in the extraction process all significantly affect polysaccharide yields. It is essential to optimize these variables to efficiently extract fungal polysaccharides and achieve high yields, along with the combination of extraction techniques.

6 Precipitation techniques for fungal polysaccharides

The precipitation of fungal polysaccharides is a process used to isolate these polysaccharides from a solution by adding a chemical agent, such as ethanol, propanol, or isopropyl alcohol, in an appropriate ratio. Ethanol is typically used to directly precipitate crude polysaccharides from the fungal extract solution (Moradi and Kalanpour, 2019; Miao et al., 2020). The use of ethanol for precipitation reduces the solubility and hydrophilic properties of polysaccharides in water (Monroy et al., 2016; Meng et al., 2023). Furthermore, precipitation at lower temperatures, such as incubation at 4°C, enhances polysaccharide precipitation (Wang et al., 2022b, 2023a, b). The precipitation of fungal polysaccharides using ethanol involves varying concentrations and volumes, depending on the type of crude polysaccharide and the extraction method employed. Previous studies reported that ethanol concentrations used for fungal polysaccharide precipitation typically range from 80% to absolute ethanol, with ethanol-to-sample ratios varying from 1:1 (v/v) to 1:5 (v/v), usually under cold conditions (4°C) and over an overnight period (Liu et al., 2018; Gao et al., 2022; Sakdasri et al., 2022; Liu et al., 2024a; Panya et al., 2024; Sahib et al., 2024). After precipitation, the polysaccharides can be separated by filtration or centrifugation and then dried by either conventional drying or lyophilization to obtain a crude solid form. The crude polysaccharides are usually kept dry to prevent moisture and microbial contamination and are used for further studies.

7 Purification techniques for fungal polysaccharides

For research and biological applications, it is essential to purify crude fungal polysaccharides in order to maintain the greatest biological properties and achieve high purity. Crude fungal polysaccharides contain a variety of contaminants, including pigments, proteins, monosaccharides, and other compounds, as a result of the limitations of extraction techniques (Wang et al., 2022b). Protein removal and decolorization of crude polysaccharides are commonly achieved using the sevage method and hydrogen peroxide treatment, respectively (Li et al., 2019b; Hu et al., 2022b). Generally, purification methods for polysaccharides include anion exchange chromatography and gel permeation chromatography. Following purification, the polysaccharides are concentrated, dialyzed, and freeze-dried. Finally, the polysaccharide and protein contents in the purified polysaccharides are quantified using the dinitrosalicylic acid, Folin-Ciocalteu, or phenol-sulfuric acid methods (Wood et al., 2012; Saravanakumar et al., 2021; Gao et al., 2022).

7.1 Anion exchange chromatography

Anion exchange chromatography is a crucial technique for purifying fungal polysaccharides, particularly those with a negative charge, such as acidic polysaccharides. This method relies on the principle of ion exchange, where the sample is passed through a column containing a medium capable of binding anions, such as DEAE-cellulose 52 or DEAE-Sephadex A-25, which have positively charged functional groups on their surfaces. When the polysaccharide is introduced into the column, negatively charged polysaccharides are adsorbed onto the medium via ion exchange. Subsequently, the bound polysaccharides are eluted from the column using a buffer solution with increasing salt concentration, which helps release the retained polysaccharides. This technique is well-suited for separating polysaccharides with varying structures and charges, enhancing extraction efficiency, and significantly reducing impurities in fungal polysaccharides (Ren et al., 2019; Wang et al., 2022b).

7.2 Gel permeation chromatography

Gel permeation chromatography is a widely used technique for purifying fungal polysaccharides, particularly for separating substances according to molecular size. The principle of gel permeation chromatography involves the use of a gel with a porous structure within the column, which facilitates the separation of sample molecules based on size. The polysaccharides are injected into a column packed with gel, such as Sephadex G-100 or Sephadex G-200. As the solution flows through the column, larger molecules are unable to penetrate the pores of the gel effectively, allowing them to move through the column more quickly and elute first. In contrast, smaller molecules can penetrate the pores, causing them to migrate more slowly and elute later. Gel permeation chromatography is particularly suitable for isolating polysaccharides with varying molecular weights, effectively reducing contaminants and resulting in purer, more homogeneous polysaccharides (Ren et al., 2019; Wang et al., 2022b).

8 Techniques for analyzing the structural and compositional characteristics of fungal polysaccharides

Fungal polysaccharides are complex compounds with diverse molecular weights, monosaccharide compositions, types of glycosidic bonds, and backbone structures, all of which vary significantly across species and growth conditions. These polysaccharides typically consist of monosaccharide units, such as glucose, fructose, mannose, and galactose, linked by glycosidic bonds, and may adopt either linear or branched structural configurations (Sun et al., 2022b; Yang et al., 2022a). Advanced analytical techniques, including gel permeation chromatography (GPC), high-pressure gel permeation chromatography (HPGPC), gas chromatography-mass spectrometry (GC-MS), high-performance liquid chromatography (HPLC), Fourier transform infrared (FT-IR) spectroscopy, nuclear magnetic resonance (NMR), and scanning electron microscopy (SEM), are employed to investigate the specific structural characteristics of fungal polysaccharides (Wang et al., 2022c; Zhao et al., 2023a). Thus, understanding these structural and functional properties is critical for advancing their potential applications and can lead to innovative product development that maximizes the benefits of fungal polysaccharides. Examples of monosaccharide composition, structure, and characterization methods of fungal polysaccharides are compiled and presented in Table 5.

8.1 GPC and HPGPC

GPC and HPGPC have been used to separate and analyze molecular sizes based on the principle of size exclusion. Larger molecules elute earlier from the column, while smaller molecules, which can penetrate the pores of the gel more effectively, elute later. GPC is particularly suited for determining average molecular weights and molecular weight distributions in polymers, proteins, and polysaccharides. In contrast, HPGPC is an advanced version of GPC, utilizing high-pressure systems and columns with smaller particle sizes to enhance resolution, speed, and accuracy (Ren et al., 2019). Both GPC and HPGPC are techniques that have been used to analyze polysaccharides from fungi with diverse molecular weights. For example, previous studies on the molecular weight of EnPs and ExPs from Co. militaris found that the HPGPC separates low-molecular-weight polysaccharides more effectively than the GPC, which can only obtain molecular weights lower than 100 kDa (Wang et al., 2015a; Huang et al., 2018; Kang et al., 2022; Liu et al., 2024; Wang et al., 2024). Similarly, studies on the molecular weight of EnPs and ExPs from Le. edodes found that HPGPC is more efficient than GPC at separating low-molecular-weight polysaccharides (Li et al., 2018a; Ren et al., 2018; Zhang et al., 2022a). Moreover, molecular weight analysis of both EnPs and ExPs from He. erinaceus revealed that HPGPC is more effective at separating low-molecular-weight polysaccharides below 10 kDa compared to GPC (Wang et al., 2015b; Wu et al., 2017, 2018). Based on the previous studies mentioned, HPGPC demonstrates the capability to separate smaller fungal polysaccharides with higher effectiveness than GPC.

8.2 GC-MS

GC is a technique used to separate components of a mixture in the gas phase based on differences in solubility and adsorption properties of each substance. This technique involves of two phases: a mobile phase, which is a carrier gas, and a stationary phase, which can be a liquid, solid, or semi-solid material contained within a column. Substances with different chemical characteristics move through the column at varying speeds, depending on their properties, such as solubility or interaction with the stationary phase. This technique allows for detailed analysis of monosaccharide composition in fungal polysaccharides with exceptional accuracy (Millette et al., 2023). For example, GC analysis revealed that arabinose, galactose, glucose, mannose, and xylose are the monosaccharides present in the composition of EnPs extracted from Co. militaris (Luo et al., 2017; He et al., 2020a; Liu et al., 2024c). Studies on the monosaccharide profile of EnPs from Le. edodes revealed the composition of rhamnose, arabinose, xylose, mannose, galactose, and glucose through GC analysis (Li et al., 2018a; Zhang et al., 2022a). The monosaccharide composition of the EnPs from G. resinaceum consists of glucose, galactose, mannose, fucose, and xylose, as determined by GC analysis (Bleha et al., 2022). Additionally, GC techniques have also been employed to determine the monosaccharide composition in ExPs. For instance, in the study by Vanin et al. (2023), the monosaccharide composition of ExPs from Sch. commune was found to be primarily composed of glucose, followed by xylose, mannose, galactose, and arabinose in smaller amounts. Furthermore, the study by Ren et al. (2018) investigating lentinan, an ExPs from Le. edodes, using GC analysis revealed that its monosaccharide composition consists of glucose, mannose, and galactose.

8.3 HPLC

HPLC is used to analyze and separate the components of fungal polysaccharides through precise chromatographic techniques. Prepared samples are introduced into an HPLC column packed with specialized separation media. As the solution flows through the column under high pressure, molecules interact with the column material according to their chemical characteristics and molecular size, enabling efficient separation. Upon elution, a detector captures the concentration of each component, yielding data for detailed analysis of monosaccharide composition, concentration quantification, and the assessment of polysaccharide degradation across various processes (Ingale et al., 2023). Using HPLC, Liu et al. (2018) examined the monosaccharide composition of EnPs isolated from Pax. involutus and found that glucose was the main component, followed by galactose, mannose, and fucose. Wang et al. (2024) investigated the monosaccharide composition of four EnPs isolated from Co. militaris using HPLC, finding that glucose and galactose were present, with glucose being the predominant component. In the study by Chen et al. (2020), HPLC was used to analyze the monosaccharide composition of four different EnPs extracted from Sch. commune obtained through various extraction methods. The results revealed that mannose, ribose, rhamnose, glucuronic acid, galacturonic acid, glucose, galactose, xylose, arabinose, and fucose were present in varying ratios, with glucuronic acid being the main component. Additionally, Ayimbila et al. (2021) analyzed the monosaccharide composition of EnPs extracted from Len. squarrosulus using HPLC, revealing that the polysaccharides were primarily composed of glucose, followed by galactose, mannose, and fucose. The HPLC technique has also been employed to analyze the monosaccharide composition of ExPs. For instance, a study reported that lentinan, an ExP derived from Le. edodes, predominantly consists of glucose as the main monosaccharide, with minor amounts of galactose and mannose detected (Xu et al., 2023). Additionally, the study by Kang et al. (2022) also revealed that the monosaccharide composition of ExPs from Co. militaris consists primarily of glucose, with smaller amounts of galactose and mannose. Comparative studies using HPLC and GC to analyze the monosaccharide composition of fungal polysaccharides have shown no significant differences, with the primary components being glucose, mannose, and galactose. However, arabinose, fucose, ribose, rhamnose, and xylose can also be found in some fungal polysaccharides.

8.4 FT-IR spectroscopy

FT-IR spectroscopy is an essential analytical method for studying fungal polysaccharides. This technique functions by inducing vibrational modes in molecular bonds via infrared radiation, which is then absorbed by the sample. The resulting absorption data is processed into a spectrum, presenting the frequency and absorption intensity relationship. This enables detailed structural analysis and functional group identification, such as glucose and fructose residues, allowing for precise assessments of sample purity and detection of potential impurities within the polysaccharide material (Hong et al., 2021). The study by He et al. (2020a) analyzed EnPs from Co. militaris using FT-IR spectroscopy, revealing a strong and broad absorption peak at 3392.8 cm⁻¹, attributed to O-H stretching vibrations of hydroxyl groups due to intramolecular or intermolecular interactions, along with a weaker absorption band at 2930.6 cm⁻¹, corresponding to C-H stretching vibrations in -CH₃ or -CH₂ groups. Similar to the study by Wang et al. (2024), which analyzed EnPs from Co. militaris to examine organic functional groups, the broad absorption peak at 3400 cm⁻¹ was attributed to O-H stretching vibrations from intermolecular or intramolecular interactions within the polysaccharide, while the weaker absorption band at 2925 to 2931 cm⁻¹ corresponded to asymmetric C-H stretching vibrations in sugar groups. In the study by Zhang et al. (2022a), the functional groups in Le. edodes EnPs were observed, with the strong absorption peak near 3300 cm^-1 attributed to O-H stretching vibrations, and the signal at 2922 cm^-1 resulting from C-H stretching vibrations; strong absorptions around 1600 cm^-1 indicated C=O stretching, while the absorption near 1400 cm^-1 was due to C-H deformation vibrations; C-O-C group signals between 1000–1100 cm^-1 confirmed pyranose ring linkages, and the weaker signal at 890 cm^-1 reflected β-glycosidic bonds. Ascencio et al. (2021) investigated the functional groups of lasiodiplodan from L. theobromae and revealed distinct spectral characteristics, including broadband with high intensity at 3274 cm^-1 corresponding to the stretching vibration of R-OH groups, a band at 1650 cm^-1 associated with the glucose ring structure, symmetric vibrations of the C-O-C bond characteristic of carbohydrates exhibiting absorption at 1075 cm^-1, and a low-intensity band at 890 cm^-1 indicative of glycosidic bonds representing the β-configuration arrangement. Based on the previous research examples provided, it can be observed that the strong and broad absorption peak is commonly attributed to the O-H hydroxyl group, which is frequently found in fungal polysaccharides. However, other functional groups, including C-H, C=O, and C-O-C, also appear, emphasizing the chemical diversity in fungal polysaccharides and reflecting the unique properties and structures found across different fungal species (He et al., 2020a; Zhang et al., 2022a).

8.5 NMR spectroscopy

NMR spectroscopy is a valuable tool for analyzing the structural properties of fungal polysaccharides by exploiting the behavior of atomic nuclei in a powerful magnetic field. Nuclei such as hydrogen (¹H) and carbon (¹³C) resonate at characteristic frequencies, absorbing energy in ways that reveal precise structural details. This technology allows for accurate determination of polysaccharide structure, including the sequence of monosaccharide units and the types of glycosidic linkages present (Yao et al., 2021). NMR spectroscopy was used to investigate the glycosidic bond structure of EnPs derived from Mor. sextelata, revealing that the EnPs contain glycosidic linkages similar to (1→4)-linked-α-Glcp suggesting a specific type of linkage in its structure Li et al. (2022). In the study by Tepsongkroh et al. (2023), NMR spectroscopy was used to examine the glycosidic bonds of polysaccharides from V. volvacea, revealing that the polysaccharide consists of (1→3)-linked-β-D-Glcp glycosidic linkages, forming a β-glucan (1,3/1,6) structure. Some studies have revealed the complexity of glycosidic bond structures, such as a research by Ayimbila et al. (2021), where NMR spectroscopy was used to analyze the glycosidic bond structure of EnPs derived from Len. squarrosulus. The analysis identified a complex structure composed of (1→4,6)-linked-β-D-Glcp, (1→4)-linked-β-D-Glcp, (1→6)-linked-β-D-Glcp, (1→4)-linked-β-D-Manp, α-L-fucose, and (1→6)-linked-α-galactosyl, highlighting the structural diversity of these EnPs. Zhang et al. (2023) found that glycosidic bond structure of EnPs derived from D. indusiata revealed a complex structure consisting of (1→6)-linked-β-Glcp, (1→2,6)-linked-α-Glcp, (1→3)-linked-α-Manp, (1→6)-linked-β-Manp, and (1→6)-linked-β-Galp. Additionally, lentinan from Le. edodes, analyzed for glycosidic bond structure using NMR spectroscopy, revealed that its backbone consists of β-(1,3)-glucan with β-(1,6) branching (Sahib et al., 2024). The NMR spectroscopy technique aids in identifying the types of glycosidic linkages, such as β-D-glucan and α-D-glucan, by detecting specific signals in the spectra. The most commonly observed glycosidic linkages include (1→3)-linked-β-Glcp, (1→6)-linked-β-Glcp, (1→3)-linked-α-Glcp, and (1→6)-linked-α-Glcp (Ayimbila et al., 2021; Gao et al., 2022; Li et al., 2022; Tepsongkroh et al., 2023; Zhang et al., 2023). This technique is crucial for understanding the types of glycosidic bonds and the backbone structures of fungal polysaccharides, which in turn influence the potential applications.

8.6 SEM

SEM is employed to examine the physical characteristics of fungal polysaccharides by using a high-energy electron beam to capture highly detailed surface images at the nanometer scale. This process begins with sample preparation, which typically includes applying a metallic coating to enhance conductivity. When the electron beam interacts with the sample surface, it generates scattered electrons and photons that are detected to produce a three-dimensional representation. SEM allows for precise analysis of structural features, including particle size, surface morphology, and distribution within the polysaccharide matrix (Fellak et al., 2022). In the study by Li et al. (2022), SEM analysis of EnPs from Mor. sextelata revealed the sponge-like porous structure with a smooth surface. Similarly, the study by Qiu et al. (2024) reported that the surface morphology of EnPs extracted from Pl. eryngii exhibited a sponge-like porous structure. A comparable sponge-like porous structure was also observed in pullulan, an exopolysaccharide extracted from Au. pullulans (Maia et al., 2023). Additionally, different extraction methods influence the surface morphology and structure of fungal polysaccharides. Each method affects the integrity and arrangement of molecules within the polysaccharide structure (Deng et al., 2015; Chemat et al., 2017). In the study by Zhang et al. (2022a), SEM was used to examine the surface morphology of EnPs from Le. edodes extracted using different methods. The polysaccharide extracted via subcritical water extraction with a deep eutectic solvent showed a loose, rough, and porous structure. The polysaccharide from subcritical water extraction alone exhibited a leaf-like surface with uneven distribution but a smooth and fine texture. The polysaccharide extracted by hot water extraction displayed a sheet-like structure with irregular aggregation. Moreover, in the study by Rosyida et al. (2024), EnPs from Pl. ostreatus were extracted using various techniques. The results revealed that polysaccharides extracted through kinetic-assisted hot extraction exhibited fewer pores and a smoother surface. In contrast, those extracted by microwave and ultrasound-assisted extraction displayed puffed structures, numerous open cavities, and collapsed surfaces. Furthermore, SEM has been employed to examine the surface morphology of exopolysaccharides, such as pullulan derived from Au. pullulans, which exhibits a smooth surface structure (Tagne et al., 2024), and lentinan from Le. edodes, characterized by a rough surface and irregular pores resembling a honeycomb-like porous structure (Xu et al., 2023).

9 Properties of fungal polysaccharides

Fungal polysaccharides have garnered significant interest in recent years due to their diverse and potent properties. These polysaccharides exhibit a wide range of beneficial effects, including antidiabetic, antioxidant, antiviral, antilipidemic, antitumor, and immunomodulating properties. An example of the biological properties of fungal polysaccharides is shown in Table 6.

9.1 Antidiabetic property

Diabetes, a non-communicable disease characterized by chronic hyperglycemia, is classified into Type 1 and Type 2. Type 1 occurs when the pancreas fails to produce insulin, while Type 2, the most common form, results from the body’s inability to effectively use insulin, often linked to factors like being overweight and lack of physical activity (American Diabetes Association, 2009). Natural supplements derived from plants and fungi, along with medications used to treat diabetes, can help control blood glucose levels (Liu et al., 2022a). According to previous research, fungal polysaccharides could beneficially affect people with diabetes by inhibiting glucose absorption efficacy, gastrointestinal viscosity, inhibition of α-amylase and α-glucosidase activity to control hyperglycemia, improving pancreatic β-cell mass, and enhancing insulin signaling (Jovanović et al., 2021; Fu et al., 2022; Liu et al., 2022a; Figueroa et al., 2023; Ji et al., 2023; Li et al., 2023; Yu et al., 2024). Generally, Streptozotocin-induced diabetic mice are frequently used to assess a compound’s ability to reduce blood glucose levels. This method yields consistent results, demonstrating the compound’s capability to lower blood glucose. Several previous studies demonstrated polysaccharides from Aur. auricula, Aur. polytricha, Co. taii, D. indusiate, G. frondose, Pl. eryngii, Pl. geesterani, Pl. ostreatus, and Su. luridus can effectively reduce blood glucose levels in Streptozotocin-induced diabetic mice (Duobin et al., 2013; Zhang et al., 2018a; Chen et al., 2019; Liu et al., 2019; Xiang et al., 2021) (Table 6). These findings indicate that fungal polysaccharides could be used as supplements for diabetes. However, future studies must evaluate the effectiveness of these fungal polysaccharides in diabetic patients to assess their true efficacy, safety, and potential side effects. If these polysaccharides can be developed into a treatment for diabetes, they could greatly benefit both medical practice and pharmacology by offering patients a more effective and safer treatment option. Moreover, the development of drugs based on fungal polysaccharides could offer new opportunities for treating diabetes.

9.2 Antilipidemic property

Hyperlipidemia is a condition characterized by elevated levels of total cholesterol (TC), triglycerides (TG), and low-density lipoprotein cholesterol (LDL-C) in the blood, exceeding established thresholds. It is a significant risk factor for various disorders, including diabetes, hypertension, stroke, and cardiovascular disease. In addition to conventional medications, dietary supplements provide an alternative for individuals with hyperlipidemia, with fungal polysaccharides emerging as a promising option due to their antilipidemic properties (De Costa and Park, 2017). Prior research employing mouse models has frequently shown that fungal polysaccharides can successfully reduce blood levels of LDL-C, TC, and TG (Table 6) (Miranda-Nantes et al., 2011; Zeng et al., 2013; Wang et al., 2019b; Ge et al., 2022). Although animal studies have demonstrated the efficacy of fungal polysaccharides in significantly reducing TC, TG, and LDL-C, further clinical trials in patients with hyperlipidemia are required to confirm their effectiveness.

9.3 Antibacterial property

The antibacterial effects refer to the ability of substances or materials to inhibit the growth of various bacteria through mechanisms such as disrupting the bacterial cell wall, inhibiting enzymes essential for bacterial growth, or preventing bacteria from adhering to host cells (Di Martino, 2022). Previous studies have focused on exploring the antibacterial properties of fungal polysaccharides, revealing that these polysaccharides have the potential to inhibit the growth of human pathogenic bacteria, including genera Bacillus, Escherichia, Klebsiella, Listeria, Micrococcus, Pseudomonas, Proteus, Shigella, Salmonella, and Staphylococcus, as well as methicillin-susceptible Staphylococcus aureus (Skalicka-Woźniak et al., 2012; Mahendran et al., 2013; Osińska-Jaroszuk et al., 2014; Wan-Mohtar et al., 2016; Ayimbila et al., 2021) (Table 6). The effectiveness of these polysaccharides varies depending on the type of fungal polysaccharide used. Thus, fungal polysaccharides exhibit promising bacterial properties and show potential for development as components in products for preventing and treating bacterial infections. However, further research is required to confirm their efficacy and safety for human use.

9.4 Antioxidant property

Free radicals, generated in the body through normal metabolism or external factors like UV radiation, pollution, or chemicals, are unstable and stabilize by stealing electrons from other molecules. They can lead to oxidative stress, which damages cells and tissues, contributing to various diseases such as heart disease, cancer, and accelerated aging (Phaniendra et al., 2015). However, the human body has compounds that protect against oxidative stress, called antioxidants, which help neutralize free radicals and reduce the damage they may cause. Antioxidants can be obtained from external sources such as food and supplements (Pham-Huy et al., 2008). Interestingly, certain fungi can produce polysaccharides with antioxidant properties, adding another potential source of these beneficial compounds (Bai et al., 2022). Previous studies have reported that fungal polysaccharides possess antioxidant properties, as demonstrated using various methods, including ABTS, DPPH, hydroxyl radical, and superoxide radical assays (Table 6) (Liu et al., 2018; Si et al., 2018; Ayimbila et al., 2021; Deng et al., 2021; Lu et al., 2022; Tabibzadeh et al., 2022). These assays help evaluate the potential of fungal polysaccharides in protecting cells from damage caused by free radicals, providing researchers with deeper insights into their role in mitigating cellular damage. However, one concern with fungal polysaccharides is the variability in their antioxidant efficacy. Thus, optimizing cultivation conditions, standardizing extraction methods, using specific fungal strains, and implementing rigorous testing and quality control can help control and reduce the variability in the antioxidant efficacy of fungal polysaccharides.

9.5 Anticancer property

Cancer is currently one of the leading causes of death worldwide. Despite significant advancements in research and treatment, including immunotherapy, targeted therapy, and proton therapy, challenges persist. Certain types of cancer remain complex and show limited response to treatment. In addition, side effects from therapies such as chemotherapy and radiation can negatively impact a patient’s quality of life (Christyani et al., 2024; Tufail et al., 2024). The alternative therapeutic strategies are being explored, including the potential of polysaccharides derived from fungi. Research has increasingly focused on the anticancer properties of fungal polysaccharides, which have shown promise in combating cancer through various mechanisms. Current studies have shown that fungal polysaccharides possess anticancer properties, with experiments conducted both in vitro using cell lines and in vivo in animal models (Table 6). However, while they induce apoptosis in cancer cells and inhibit cancer cell growth, the results obtained are still in the preliminary stages and have limitations (Mao et al., 2016; Su et al., 2018; Geraldelli et al., 2020; Udchumpisai and Bangyeekhun, 2020; Wu et al., 2022b; Abdala-Díaz et al., 2024). Although initial findings show promising potential, further long-term studies and clinical trials in humans are necessary to confirm the effectiveness and safety of using fungal polysaccharides as a therapeutic approach to cancer treatment.

9.6 Antiviral property

Developing antiviral substances holds significant importance for advancements in medicine and public health (Gonçalves et al., 2021). Antiviral properties are one of the key characteristics of fungal polysaccharides that have been extensively studied. Research has demonstrated that fungal polysaccharides can inhibit viral replication, including against herpes simplex virus type I (Sacchelli et al., 2019), hepatitis B virus (Jiao et al., 2018), and influenza A virus (Vlasenko et al., 2020) (Table 6). Although current research has yet to definitively demonstrate that fungal polysaccharides inhibit several viruses, ongoing studies are required to fully realize their potential in antiviral therapeutics. Further studies should address important constraints, including their narrow spectrum, unknown mechanisms, low bioavailability, and the need for additional clinical evidence.

9.7 Immunomodulating property

Fungal polysaccharides are known for their immunomodulatory properties, with low toxicity, high molecular mass, and diverse branching structures that may trigger appropriate immune responses in humans (Zhao et al., 2020; Yin et al., 2021). Fungal polysaccharides possess notable properties in modulating and enhancing immune system function. The key mechanism involves activating immune cells, such as macrophages, and stimulating cytokine secretion, which further supports immune system activity (Table 6). These polysaccharides hold the potential for development into immune-boosting supplements or therapeutic agents to prevent and alleviate the effects of infections, chronic diseases, or immune deficiencies (Murphy et al., 2023a, b). However, most of the research to date has been limited to in vitro and in vivo studies, which may not be sufficient for the development of clinical-grade therapeutics. Therefore, further clinical studies in humans are necessary to confirm the effectiveness of fungal polysaccharides in immune modulation, as well as to evaluate any potential side effects, ensuring safety and reliability for practical use.

9.8 Prebiotic property

Prebiotic property is another important characteristic of polysaccharides, as previous research has shown that polysaccharides from edible fungi are resistant to human digestive enzymes and serve as a crucial energy source for the gut microbiome, promoting the growth of beneficial bacteria (Uthan et al., 2021; Zhao et al., 2023b). Bifidobacterium, Lacticaseibacillus, and Lactobacillus are key probiotic bacteria in the human gastrointestinal system, playing an essential role in promoting intestinal health, supporting immune function, and maintaining the balance of gut microbiota (Wang et al., 2022b; Panya et al., 2024). Recent studies have highlighted the potent prebiotic properties of fungal polysaccharides, which can enhance the growth of probiotics both in vitro and in animal models (Table 6). These findings suggest that fungal polysaccharides hold significant potential for development as functional ingredients in prebiotic products that support gastrointestinal health (Barcan et al., 2024). Despite their potential as prebiotic agents, fungal polysaccharides have a number of limitations that require more further research and development. Unlocking their full potential as functional components in prebiotic products will require increasing their bioavailability, understanding their mechanisms of action, establishing clinical evidence, and resolving issues with cost and production scalability.

10 Applications for fungal polysaccharides

This research presents the application of polysaccharides derived from fungi in the fields of medicine, pharmacology, food, agriculture, animal, and environmental applications as shown in Figure 5.

Figure 5

10.1 Applications in medicine and pharmacology

10.1.1 Drug delivery

Considerable advancements have been made in drug delivery systems, enhancing the precision of drug release, stability against degradation, and extending the duration of drug action. Innovations such as microcapsules, nanoparticles, and polysaccharide-based coatings have been developed to improve the accuracy of drug targeting (Ezike et al., 2023). The incorporation of polysaccharide-based coatings in drug delivery systems offers several advantages, including safety, stability, controlled release, biodegradability, and natural origin, making these materials highly compatible. This versatility, especially when combined with other delivery platforms, broadens the potential applications of polysaccharides in drug administration (Visan and Cristescu, 2023). Fungal polysaccharides, particularly pullulan, play a crucial role in drug delivery due to their unique beneficial properties. As noted by Thakur et al. (2023), pullulan-based polymeric drug delivery systems offer a pioneering approach for respiratory disease treatment, featuring a sustained release profile, biocompatibility, non-immunogenicity, chemical modifiability, and superior permeability through lung mucosa compared to other polysaccharides. Gehrcke et al. (2022) have developed a bilayer film composed of pullulan, encapsulating silibin-loaded nanocapsules for the treatment of atopic dermatitis. In vitro, results demonstrated that pullulan enhances the film’s adhesion to skin tissue, and the film possesses antioxidant properties without inducing hemolysis. Interestingly, in vivo experiments, the film regulated inflammation and oxidative parameters as effectively as, or even better than, silibinin solution and hydrocortisone, which are conventional treatments for atopic dermatitis. In addition, Balasso et al. (2017) used pullulan in a bioconjugate specifically designed for targeted drug delivery to hepatocellular carcinoma. The results indicated that the PreS1-Pullulan-Doxorubicin polymer enhanced anticancer efficacy against HepG2/SERPINB3 cells twofold compared to the control pullulan-Doxorubicin. This finding highlights the potential of enhancing polysaccharides as selective therapeutic agents for hepatocellular carcinoma by promoting targeted accumulation within cancerous liver cells. Furthermore, research has been conducted on drug delivery using schizophyllan, which forms stearic acid-schizophyllan micelles, a novel type of drug carrier (Negahban et al., 2021). Paclitaxel loaded into these micelles exhibited higher anticancer activity against MCF-7 cells compared to free paclitaxel. Notably, schizophyllan also possesses unique anti-inflammatory and immune-enhancing properties, which could potentially synergize with the therapeutic effects of the loaded drug.

10.1.2 Gene delivery

Gene delivery refers to the process of delivering genetic materials, such as DNA or RNA, to various cells or tissues in the body to modify, enhance, or alter gene function. This technique is widely used in genetic research, vaccine development, gene therapy, and studies related to various diseases (Sung and Kim, 2019). Gene delivery can be achieved through several methods, including the use of viruses that have been modified to be non-pathogenic, such as adenoviruses, lentiviruses, and adeno-associated viruses (AAV), to deliver genetic materials into cells. Additionally, non-viral approaches utilize nanocarriers made from synthetic materials or natural substances to facilitate gene delivery (Pan et al., 2021). These include polymeric nanoparticles or polysaccharides, providing alternative methods for introducing genetic material without the use of viruses (Li et al., 2018b). Previous research by Kang et al. (2010) incorporated the polysaccharide pullulan into polyethyleneimine (PEI) and small interfering RNA (siRNA) to create a pullulan-modified PEI/siRNA complex for targeted delivery to the liver, comparing it with the PEI/siRNA complex. After the injection into mice, siRNA was detected using fluorescence techniques. The results revealed that the PEI-pullulan/siRNA complex exhibited higher fluorescence in the liver compared to the PEI/siRNA complex, while the PEI/siRNA complex demonstrated greater fluorescence in the lungs. The PEI-pullulan/siRNA complex significantly reduced mouse mortality and could serve as an effective, low-toxicity strategy for delivering siRNA to the liver. Similar to other targeted delivery systems for plasmid DNA (pDNA) and siRNA directed at folate receptors on cancer cells, Wang et al. (2014a) synthesized an enhanced gene delivery carrier using folate-polyethyleneimine-modified pullulan for the delivery of pDNA and siRNA. The results indicated that the folate-polyethyleneimine-modified pullulan effectively encapsulates pDNA and siRNA, with efficient delivery occurring through the folate receptor. This suggests that folate-polyethyleneimine-modified pullulan is a promising, safe, and precise gene delivery system suitable for targeting cancer cells. Additionally, the study by Liu et al. (2014) developed an innovative nanoparticle system with a core-shell structure, utilizing pullulan and poly(β-amino) ester (PBAE) conjugated to methotrexate (anti-cancer drug) and green fluorescent protein (pEGFP). This methotrexate-pullulan/PBAE/pEGFP nanoparticle was specifically designed for the targeted delivery of gene and chemotherapeutic agents to liver cancer cells. This nanoparticle exhibited strong hepatoma-targeting capability, primarily accumulating in liver cancer cells within 24 hours post-intravenous injection. It successfully facilitated the co-delivery of gene and chemotherapy agents to tumor sites at both the cellular and animal levels, offering a promising approach to liver cancer treatment.

10.1.3 Tissue engineering

Tissue engineering is an emerging field that plays a crucial role in promoting the regeneration of damaged tissues that are unable to self-repair. It enhances the body’s natural healing potential, supporting recovery in a way that complements the patient’s healing process. Injured tissue repair can be stimulated through the use of synthetic polymer scaffolds, which serve as structural supports that facilitate efficient tissue regeneration (Singh et al., 2016). In a prior study by Schlaubitz et al. (2014), pullulan combined with dextran and reinforced with nanocrystalline hydroxyapatite was developed into highly porous scaffold beads to evaluate cellular growth and new bone formation in a rat model. The findings indicated effective cellular penetration into the scaffold spaces and new tissue formation around the beads. Additionally, calcium and mineral deposition within the structure increased over time, with no inflammation observed at the implantation site. Thangavel et al. (2020) used pullulan gel to evaluate its wound-healing efficacy on open excision wounds made on the dorsum of rats. The study found that pullulan gel significantly stimulated collagen, hexosamine, protein, and DNA synthesis compared to the untreated and povidone-iodine ointment-treated groups, demonstrating its potential as a wound-healing agent. The effects of β-glucan from S. cerevisiae on wound healing in venous ulcers in humans were evaluated by Medeiros et al. (2012) through histopathological analysis. The findings revealed that β-glucan significantly accelerated wound healing by promoting epithelial hyperplasia, stimulating angiogenesis, increasing plasmocyte numbers, and enhancing fibroblast proliferation.

10.1.4 Vaccine adjuvants

Vaccine adjuvants are substances or components added to vaccines to enhance the immune response’s effectiveness. Typically, vaccines contain immunogenic agents such as antigens; however, the incorporation of adjuvants intensifies and prolongs the immune stimulation. Therefore, the use of adjuvants in vaccines enables the development of more effective vaccines and enhances their ability to prevent various diseases more effectively (Zhao et al., 2023c). A study by Zhang et al. (2017) found that lentinan, tremella, pachymaran, and their combination with the H1N1 influenza vaccine in mice enhanced the vaccine’s efficacy. These findings suggest that fungal polysaccharides could help improve the effectiveness of influenza vaccines. He et al. (2020b) used calcium carbonate-lentinan loaded with the H5N1 antigen to develop an adjuvant for the H5N1 vaccine, aimed at preventing the avian influenza virus. After injection into mice, the calcium carbonate-lentinan/H5N1 complex significantly enhanced the expression of MHC-II and CD86 in dendritic cells from lymph nodes. It also unexpectedly led to elevated hemagglutination inhibition (HI) titers and stimulated the secretion of IgG subtypes (IgG1 and IgG2b), along with T-helper-associated cytokines (TNF-α, IFN-γ, and IL-4) in vaccinated mice. Additionally, Liu et al. (2019) developed a vaccine by encapsulating polysaccharides from G. lucidum and inactivated porcine circovirus type II (PCV-II) into liposomes. When administered to mice, this vaccine induced stronger PCV-II-specific immune responses, including higher titers of PCV-II-specific IgG antibodies, increased cytokine levels, and splenocyte activation, compared to other single-component formulations.

10.1.5 Cosmeceuticals

Cosmeceuticals are products that combine the properties of cosmetics and pharmaceuticals to enhance the effectiveness of skincare and beauty treatments. These products typically contain active ingredients that nourish or treat the skin, such as vitamins, minerals, antioxidants, and natural substances. They are designed to improve skin health, prevent deterioration, and effectively reduce the appearance of wrinkles (Singh et al., 2024). Kanlayavattanakul and Lourith (2023) found that the addition of polysaccharide extract from Co. militaris to a skincare cream formulation increased the cream’s stability. A study by Woźniak et al. (2023) revealed that polysaccharide extract from Tre. fuciformis serves as a natural alternative to hyaluronic acid in cosmetic formulations, significantly improving skin hydration without causing irritation or erythema, as confirmed by dermatological evaluations. A study by Sangthong et al. (2022) demonstrated that a cosmetic gel cream containing polysaccharides from V. volvacea could enhance skin hydration, elasticity, and firmness, while reducing skin roughness, dryness, wrinkles, and melanin content, with no cytotoxic effects on human dermal fibroblasts. Additionally, a study conducted by Jesenak et al. (2016) demonstrated that the application of Imunoglukan P4H^® cream, containing β-glucan (pleuran isolated from Pl. ostreatus), significantly reduced the frequency and severity of atopic dermatitis in patients.

10.2 Applications in functional food and food industry

10.2.1 Food additives

Fungal polysaccharides can be applied in the food industry as additives to improve food quality, such as gelling agents, stabilizers, thickeners, or texture modifiers. These polysaccharides do not affect the taste of food, which is a critical aspect to consider in food products, while also enhancing sensory properties. Furthermore, polysaccharides help extend the shelf life of food (Jindal and Singh Khattar, 2018). Pullulan has been used in the food industry for over 20 years and has been recognized as Generally Recognized as Safe (GRAS) in the USA (Oğuzhan and Yangılar, 2013). This is due to its characteristics as a colorless, odorless, edible polysaccharide that is non-toxic and non-carcinogenic (Prajapati et al., 2013). Pullulan has been utilized as a thickening agent in foods such as soups, sauces, and various beverages (Yatmaz and Turhan, 2012). According to Singh et al. (2019), pullulan has been used as a stabilizer and texture modifier in mayonnaise, and it has also been employed to enhance the adhesion of nuts in cookies. Furthermore, pullulan has low viscosity and remains stable when exposed to sodium chloride and heat, making it an ideal thickening agent for food products with high salt concentrations, such as barbecue sauces, soy sauces, pickled fruits, and vegetables. In addition, scleroglucan can serve as a gelling agent, stabilizer, or thickener, offering an alternative to xanthan gum in the food industry. It improves the quality of heat-treated and frozen foods, such as steamed foods, rice crackers, Japanese cakes, and bread (Jindal and Singh Khattar, 2018). Pleuran and lentinan can be used to produce hydrogels for yogurt products without affecting the flavor or texture while enhancing health benefits (Vetter, 2023).

10.2.2 Edible films and biodegradable packaging

Fungal polysaccharides can be utilized in the production of edible or biodegradable films (Cazón et al., 2017). Pullulan is widely used in the fabrication of edible films due to its strength, flexibility, and excellent oxygen barrier properties, which prevent food oxidation from air exposure (Oğuzhan and Yangılar, 2013). Edible films made from pullulan have also been employed in film formation and packaging for various foods, including vegetables, fruits, grains, snacks, and dried goods (Singh et al., 2016, 2019). Previous studies have shown that pullulan, when combined with other substances such as chitosan, alginate, or shellac, can create films with oxygen, water, and UV protection, making them suitable for food packaging (Zhou et al., 2021; Mugnaini et al., 2024; Tang et al., 2025). Additionally, pullulan has been incorporated with propolis to develop antifungal food packaging, which protects against pathogenic fungi, thus preventing fungal growth in food and extends food shelf life (Gniewosz et al., 2022). A study by Viñarta et al. (2013) revealed that scleroglucan derived from Scl. rolfsii demonstrated superior performance, particularly in enhancing pseudoplastic behavior, and showed excellent compatibility with corn starch, xanthan, pectin, and carboxymethylcellulose, making it highly suitable for applications in the food industry. Beyond pullulan, scleroglucan, and schizophyllan can be used to form edible films for dietary supplement packaging due to their chemical stability and natural biodegradability (Rahman et al., 2021; Ghosh et al., 2022).

10.2.3 Functional foods

Edible mushrooms are recognized as health-promoting foods for humans. In recent years, bioactive polysaccharides extracted from edible mushrooms have been developed into functional foods that are more easily and rapidly absorbed by the body, resulting in faster bioactivity. In addition to functional foods for human health, polysaccharides from mushrooms can also be used as animal feed (Osińska-Jaroszuk et al., 2020). β-glucan, a polysaccharide derived from mushrooms, is well-known for its diverse biological and pharmacological properties, including immunomodulatory, antioxidant, antimicrobial, anticancer, cardioprotective, and hepatoprotective activities. β-glucan from mushrooms can enhance both innate and cell-mediated immune responses. It has demonstrated varying degrees of antitumor activity in humans due to differences in structure, water solubility, size, and molecular mass (Khan et al., 2018). Furthermore, previous studies have shown that polysaccharides from several mushrooms, e.g. G. lucidum, Le. edodes, Tre. fuciformis, Au. pullulans, Pleurotus spp., and Ag. bisporus (Cheng et al., 2011; Giannenas et al., 2011; Delzenne and Bindels, 2015) exhibit prebiotic properties, promoting the growth of beneficial bifidobacteria, a probiotic in the human gut.

Currently, polysaccharide extracts from mushrooms are commercially available as dietary supplements and functional foods. For instance, Immune-Assist™ Critical Care Formula, produced by Aloha Medicinals Inc., contains extracts from various mushrooms, including Ag. blazei (58.5% β‐glucan), Co. sinensis (30% β‐glucan), Gr. frondosa (28% β‐glucan), Le. edodes (40% β‐glucan, lentinan, and α‐glucan), Tr. versicolor (40% β‐glucan), and G. lucidum (40% β‐glucan and triterpenoids). This product has been shown to reduce the side effects of chemotherapy and radiotherapy in cancer patients. Super Reishi^®, a β‐glucan extract from G. lucidum produced by Mushroom Wisdom Inc., is known for modulating and supporting multiple bodily systems, including the heart, lungs, liver, nervous system, and brain. Maitake^®, developed by Pharmaceutical Mushrooms Inc., is a β‐glucan-based product derived from Gr. frondosa that demonstrates strong immunomodulatory effects, notably enhancing T-cell production, and is specifically recommended for managing immunodeficiency conditions. Fine‐Agaricus^® Gold, produced by FineCo. Ltd. is a 100% polysaccharide extract from Agaricus that has demonstrated efficacy against various cancers through immune system enhancement, while also balancing physiological functions and providing benefits for the treatment of chronic diseases. Additionally, Transfer Factor Plus^® Tri‐Factor^® Formula, produced by Product 4life Inc., contains extracts from Le. edodes, Gr. frondosa, and Cordyceps sp., comprising β‐glucans, hexaphosphate inositol, β‐sitosterol, and an extract of olive leaves. This formula has been shown to stimulate the immune system by enhancing the activity of NK cells in the body (Morris et al., 2016).

10.3 Applications in agriculture, animals, and environment

10.3.1 Animal feed ingredients

Polysaccharides from fungi, in addition to being used as functional food for humans, can also be applied as animal feed ingredients to enhance the nutritional value of livestock (Osińska-Jaroszuk et al., 2020). Research by Muthusamy et al. (2013) demonstrated that β-glucan derived from Pl. floridanus could be incorporated into poultry feed, significantly enhancing the immune response in broiler chickens. Similar to the study by Gao et al. (2024), which found that supplementation of polysaccharides from G. lucidum increased HDL-C levels in the serum, reduced TG levels, enhanced antioxidant activity, increased antioxidant enzyme levels in broiler chickens, and contributed to the improvement of gut microbiota composition. In addition, β-glucan from Pl. floridanus and Pl. ostreatus has been used in fish feed to enhance immune function in fish (Dobšíková et al., 2013). Furthermore, β-glucan has been combined with milk to serve as a dietary supplement in newly weaned piglets, promoting growth performance and gut health (Mukhopadhya et al., 2019).

10.3.2 Soil improvement and plant growth stimulation

Fungal polysaccharides can be applied in agriculture, particularly ExP, which are mucilaginous and carry ionic charges, allowing them to bind soil particles together and enhance soil fertility (Costa et al., 2018). Previous research introduced the use of ExPs in soil, revealing that ExPs improve the aggregate stability of soil particle aggregation in water, thereby enhancing water retention and increasing soil porosity (Costa et al., 2018; Angulo et al., 2024). ExPs produced by basidiomycetes and Trichocomacea can bind soil particles, promoting soil aggregate formation and stability (Daynes et al., 2012; Angulo et al., 2024). Polysaccharides from G. lucidum have been shown to increase seed germination rates and seedling heights when combined with chemical fungicides in seed-coating formulations, effectively controlling soil-borne diseases. Furthermore, polysaccharides from G. lucidum enhance the expression of genes associated with disease resistance in maize and wheat, enabling their use alongside chemical fungicides in the prevention of maize root rot, wheat root rot, and sharp eyespot diseases (Yang et al., 2022a, b). Application of polysaccharides isolated from the fruiting body of Pl. ferulae exhibited antifungal activity against Rhizoctonia solani, promoted cucumber plant growth by enhancing root length and fresh weight of cucumber seedlings, and stimulated the activities of enzymes such as superoxide dismutase, peroxidase, and polyphenol oxidase for disease resistance (Yang et al., 2024c).

10.3.3 Oil recovery

Oil recovery refers to the process of increasing the extraction rate of crude oil from underground reservoirs (Jafarinejad, 2017). After conventional extraction methods, substantial amounts of oil often remain trapped within the rock formations, necessitating enhanced techniques such as enhanced oil recovery (EOR). EOR methods utilize chemical agents, thermal injections, or biotechnological solutions to improve the recovery of this residual oil (Rizvi, 2024). Fungal polysaccharides are particularly promising for EOR applications. These polysaccharides can enhance the viscosity of injected solutions, effectively displacing oil trapped within the pore spaces of rock formations, thus facilitating oil movement toward the extraction wellbore. Additionally, fungal polysaccharides exhibit high stability under extreme temperature and pressure conditions, making them well-suited for EOR in the challenging underground environments of oil reservoirs (Xia et al., 2020). For example, scleroglucan can be modified to acquire hydrophobic properties by grafting stearate groups, along with the addition of ionic-sulfonic groups to enhance the viscosity of the compound. This modification alters the adsorption behavior of scleroglucan on oil reservoir rock surfaces, with greater stearate grafting density resulting in increased adsorption (Bakhshi et al., 2017). Schizophyllan is used in EOR processes and has been shown to increase the recovery of crude oil by up to 28% over the residual oil saturation (Joshi et al., 2016). Additionally, the use of pullulan in microbial-enhanced oil recovery experiments with Berea sandstone core samples resulted in a 9.4% increase in the recovery of medium-heavy oil (Elshafie et al., 2017).

11 Patent search

A search for patents using the keyword “fungal polysaccharides” in the Espacenet database (https://worldwide.espacenet.com, accessed 25 March 2025) reveals significant changes in patent filing trends between 2010 and 2024. The data can be categorized into three periods, as presented in Figure 6A. The number of patent filings increased significantly from approximately 9,972 between 2010 and 2014 to around 12,550 from 2015 to 2019, indicating substantial growth in interest. However, this trend reversed from 2020 to 2024, with filings decreasing to 9,488. This decline may be influenced by factors such as economic conditions, policy shifts, and evolving technological advancements during that period, as well as delays in research and patent applications resulting from the COVID-19 pandemic.

Figure 6

Based on the International Patent Classification (IPC) codes, the top ten patent classifications for fungal polysaccharides revealed that category A61K31, which covers organic compounds for medicinal and cosmetic preparations, had the highest total with 6,685 patents (Figure 6B). This was followed by category C12N15 (genetic modification, vectors, and genetic material manipulation) with 6,183 patents, C12N1 (microorganisms and biotechnological processes) with 4,880 patents, A61K9 (pharmaceutical formulations) with 4,635 patents, and A61K47 (drug carriers and excipients) with 4,382 patents. Other notable categories included A61P31 (anti-infectives), C12N9 (enzymes and proenzymes), A61P35 (antineoplastic agents), and A61K39 (medicinal preparations containing antigens or antibodies). This indicates that categories involving organic compounds for pharmaceutical and cosmetic formulations were particularly popular, reflecting the strong potential of fungal polysaccharides to meet competitive market demands with value-added products. The top ten countries for fungal polysaccharide-related patent filings are shown in Figure 6C. The majority of patent filings were made through the World Intellectual Property Organization (WIPO) with 18,520 patents, followed by China with 18,137 patents, the United States with 16,038 patents, and the European Patent Office (EU) with 13,325 patents. Japan ranked fifth with 8,912 patents, followed by Canada (7,740 patents), Australia (6,488 patents), South Korea (6,150 patents), Brazil (5,070 patents), and Mexico (4,321 patents), rounding out the list.

12 Conclusions and future perspectives

Fungal polysaccharides are derived from a wide variety of fungal species, including yeasts, filamentous fungi, and mushrooms. The types of polysaccharides vary in composition and structure. To obtain the highest yield of fungal polysaccharides, the processes of fermentation, extraction, precipitation, and purification must be optimized for each fungal species and strain. Each step requires careful consideration.

In recent years, significant advancements have been made in the study of fungal polysaccharides, particularly in their structural characterization, bioactivities, and biotechnological applications. Despite this progress, notable deficiencies remain in our fundamental understanding, especially regarding structure–function and size–function relationships, as well as biosynthetic pathways. Fungal polysaccharides possess diverse biological properties and potential therapeutic benefits. These polysaccharides are used in agriculture, medicine, food, cosmetics, and biotechnology industries. The trend in research on fungal polysaccharides has been growing significantly in recent years. Future research may focus on discovering new fungal species and strains to enhance the optimization of production processes, reduce costs, and support large-scale industrial applications, along with the integration of advanced technologies. Advances in extraction, purification and characterization techniques will further enhance the quality and functional properties of fungal polysaccharides, enabling more precise applications in pharmaceuticals, food, cosmetics, and biotechnology. Moreover, challenges persist in the standardization of extraction and purification methods, which continues to be a major limitation in the field. Although fungal polysaccharides exhibit great potential in medicine and the food industry, their practical applications are still in the experimental and clinical testing stages. Furthermore, continued further research into the biomedical applications of fungal polysaccharides, such as drug delivery systems, immune modulation, and tissue engineering, will expand their impact in healthcare. The biological responses of the human body to fungal polysaccharides are not yet fully understood, highlighting the need for further studies on their safety and health impacts. Finally, to advance both the fundamental understanding and practical applications of fungal polysaccharides, these gaps must be addressed through advanced integrative techniques, including omics (genomics, transcriptomics, proteomics, and metabolomics), CRISPR-based genome editing, bioinformatics, and systems biology, combined with interdisciplinary collaboration.

References

• 1

  Abdala-DíazR. T.Casas-ArrojoV.Castro-VarelaP.RiquelmeC.CarrilloP.MedinaM.Á.et al. (2024). Immunomodulatory, antioxidant, and potential anticancer activity of the polysaccharides of the fungus Fomitiporia chilensis. Molecules29, e3628. doi: 10.3390/molecules29153628

  • CrossRef
  • Google Scholar
  • View reference in article

• 2

  AbreuH.ZavadinackM.SmiderleF. R.CiprianiT. R.CordeiroL. M. C.IacominiM. (2021). Polysaccharides from Pleurotus eryngii: Selective extraction methodologies and their modulatory effects on THP-1 macrophages. Carbohyd. Polym.15, e117177. doi: 10.1016/j.carbpol.2020.117177

  • CrossRef
  • Google Scholar
  • View reference in article

• 3

  Aguiló-AguayoI.WaltonJ.ViñasI.TiwariB. K. (2017). Ultrasound assisted extraction of polysaccharides from mushroom by-products. LWT-Food Sci. Technol.77, 92–99. doi: 10.1016/j.lwt.2016.11.043

  • CrossRef
  • Google Scholar
  • View reference in article

• 4

  AlitongbiekeG.ZhangX.ZhuF.WuQ.LinZ.LiX.et al. (2024). Glucan from Oudemansiella raphanipes suppresses breast cancer proliferation and metastasis by regulating macrophage polarization and the WNT/β-catenin signaling pathway. J. Cancer15, 1169–1181. doi: 10.7150/jca.89873

  • CrossRef
  • Google Scholar
  • View reference in article

• 5

  Alves da CunhaM. A.TurminaJ. A.IvanovR. C.BarrosoR. R.MarquesP. T.FonsecaE. A. I.et al. (2012). Lasiodiplodan, an exocellular (1→6)-β-d-glucan from Lasiodiplodia theobromae MMPI: production on glucose, fermentation kinetics, rheology and anti-proliferative activity. J. Ind. Microbiol. Biot.39, 1179–1188. doi: 10.1007/s10295-012-1112-2

  • CrossRef
  • Google Scholar
  • View reference in article

• 6

  AmerM. S.ZaghloulE. H.IbrahimM. I. A. (2020). Characterization of exopolysaccharide produced from marine-derived Aspergillus terreus SEI with prominent biological activities. Egypt. J. Aquat. Res.46, 363–369. doi: 10.1016/j.ejar.2020.08.008

  • CrossRef
  • Google Scholar
  • View reference in article

• 7

  American Diabetes Association (2009). Diagnosis and classification of diabetes mellitus. Diabetes Care32, 62–67. doi: 10.2337/dc09-s062

  • CrossRef
  • Google Scholar
  • View reference in article

• 8

  AnchundiaM.León-ReveloG.SantacruzS.TorresF. (2024). Polyphasic identification of Rhizopus oryzae and evaluation of physical fermentation parameters in potato starch processing liquid waste for β-glucan production. Sci. Rep.14, e14913. doi: 10.1038/s41598-024-66000-5

  • CrossRef
  • Google Scholar
  • View reference in article

• 9

  AngelovaG.BrazkovaM.MihaylovaD.SlavovA.PetkovaN.BlazhevaD.et al. (2022). Bioactivity of biomass and crude exopolysaccharides obtained by controlled submerged cultivation of medicinal mushroom Trametes versicolor. J. Fungi8, e738. doi: 10.3390/jof8070738

  • CrossRef
  • Google Scholar

• 10

  AnguloV.BleichrodtR.DijksterhuisJ.ErktanA.HeftingM. M.KraakB.et al. (2024). Enhancement of soil aggregation and physical properties through fungal amendments under varying moisture conditions. Environ. Microbiol26, e16627. doi: 10.1111/1462-2920.16627

  • CrossRef
  • Google Scholar
  • View reference in article

• 11

  AntonelliA.FryC.SmithR. J.EdenJ.GovaertsR. H. A.KerseyP.et al. (2023). State of the world’s plants and fungi 2023 (Royal Botanic Gardens, Kew). doi: 10.34885/wnwn-6s63

  • CrossRef
  • Google Scholar
  • View reference in article

• 12

  AscencioJ. J.PhilippiniR. R.GomesF. M.PereiraF. M.da SilvaS. S.KumarV.et al. (2021). Comparative highly efficient production of β-glucan by Lasiodiplodia theobromae CCT 3966 and its multiscale characterization. Fermentation7, e108. doi: 10.3390/fermentation7030108

  • CrossRef
  • Google Scholar
  • View reference in article

• 13

  AshfaqueA. K.ShahzorG. K.YingL.SaghirA. S.Yan-FengW.AijazH. S.et al. (2017). Optimization of enzyme assisted extraction of polysaccharides from Poria cocos. J. Med. Plant Res.11, 331–337. doi: 10.5897/jmpr2017.6366

  • CrossRef
  • Google Scholar
  • View reference in article

• 14

  AyimbilaF.KeawsompongS. (2023). Nutritional quality and biological application of mushroom protein as a novel protein alternative. Curr. Nutr. Rep.12, 290–307. doi: 10.1007/s13668-023-00468-x

  • CrossRef
  • Google Scholar
  • View reference in article

• 15

  AyimbilaF.SiriwongS.KeawsompongS. (2021). Structural characteristics and bioactive properties of water-soluble polysaccharide from Lentinus squarrosulus. Bioact. Carbohydr. Diet. Fibre.26, e100266. doi: 10.1016/j.bcdf.2021.100266

  • CrossRef
  • Google Scholar
  • View reference in article

• 16

  Azuar MohamadS.Abd HalimN. H.KeongC. Y.Mohd DzomirA. Z.YazidH.HamzahM. Y.et al. (2018). Local β-glucan source from oyster mushroom (Pleurotus flabellatus) as potential usage in medical applications. Front. Pharmacol.9, e112.

  • Google Scholar
  • View reference in article

• 17

  BackhausK.HeilmannC. J.SorgoA. G.PurschkeG.de KosterC. G.KlisF. M.et al. (2010). A systematic study of the cell wall composition of Kluyveromyces lactis. Yeast27, 647–660. doi: 10.1002/yea.1781

  • CrossRef
  • Google Scholar
  • View reference in article

• 18

  BaiL.XuD.ZhouY. M.ZhangY.-B.ZhangH.ChenY.-B.et al. (2022). Antioxidant activities of natural polysaccharides and their derivatives for biomedical and medicinal applications. Antioxidants11, e2491. doi: 10.3390/antiox11122491

  • CrossRef
  • Google Scholar
  • View reference in article

• 19

  BainsA.SridharK.KaushikR.ChawlaP.SharmaM. (2023). Enzyme-assisted polysaccharides extraction from Calocybe indica: Synergistic antibiofilm and oxidative stability of essential oil nanoemulsion. Int. J. Biol. Macromol.242, e124843. doi: 10.1016/j.ijbiomac.2023.124843

  • CrossRef
  • Google Scholar
  • View reference in article

• 20

  BakhshiM.OzeiriM.SharifA.AalaieJ. (2017). Effect of hydrophobic modification on the structure and rheology of aqueous and brine solutions of scleroglucan polymer. Korean J. Chem. Eng.34, 903–912. doi: 10.1007/s11814-016-0322-0

  • CrossRef
  • Google Scholar
  • View reference in article

• 21

  BakratsasG.PolyderaA.KatapodisP.StamatisH. (2021). Recent trends in submerged cultivation of mushrooms and their application as a source of nutraceuticals and food additives. Future Foods.4, e100086. doi: 10.1016/j.fufo.2021.100086

  • CrossRef
  • Google Scholar
  • View reference in article

• 22

  BakratsasG.TsoumanisC.StamatisH.KatapodisP. (2024). Exopolysaccharide production in submerged fermentation of Pleurotus ostreatus under red and green light. Fermentation10, e313. doi: 10.3390/fermentation10060313

  • CrossRef
  • Google Scholar
  • View reference in article

• 23

  BalassoA.SalmasoS.PontissoP.RosatoA.QuartaS.MalfantiA.et al. (2017). Re-programming pullulan for targeting and controlled release of doxorubicin to the hepatocellular carcinoma cells. Eur. J. Pharm. Sci.103, 104–115. doi: 10.1016/j.ejps.2017.02.016

  • CrossRef
  • Google Scholar
  • View reference in article

• 24

  BarcanA. S.BarcanR. A.VamanuE. (2024). Therapeutic potential of fungal polysaccharides in gut microbiota regulation: implications for diabetes, neurodegeneration, and oncology. J. Fungi10, e394. doi: 10.3390/jof10060394

  • CrossRef
  • Google Scholar
  • View reference in article

• 25

  BentilJ. A.ThygesenA.MensahM.LangeL.MeyerA. S. (2018). Cellulase production by white-rot basidiomycetous fungi: solid-state versus submerged cultivation. Appl. Microbiol. Biotechnol.102, 5827–5839. doi: 10.1007/s00253-018-9072-8

  • CrossRef
  • Google Scholar
  • View reference in article

• 26

  BhotmangeD. U.WalleniusJ. H.SinghalR. S.ShamekhS. S. (2017). Enzymatic extraction and characterization of polysaccharide from Tuber aestivum. Bioact. Carbohydr. Diet. Fibre.10, 1–9. doi: 10.1016/j.bcdf.2017.02.001

  • CrossRef
  • Google Scholar
  • View reference in article

• 27

  BlehaR.TřešnákováL.SushytskyiL.CapekP.ČopíkováJ.KloučekP.et al. (2022). Polysaccharides from basidiocarps of the polypore fungus Ganoderma resinaceum: isolation and structure. Polym.14, 255. doi: 10.3390/polym14020255

  • CrossRef
  • Google Scholar
  • View reference in article

• 28

  BrundrettM. C. (2002). Coevolution of roots and mycorrhizas of land plants. New Phytal.154, 275–304. doi: 10.1046/j.1469-8137.2002.00397.x

  • CrossRef
  • Google Scholar
  • View reference in article

• 29

  Bzducha-WróbelA.FarkašP.BielikováS.ČížováA.Sujkowska-RybkowskaM. (2024). How do the carbon and nitrogen sources affect the synthesis of β-(1,3/1,6)-glucan, its structure and the susceptibility of Candida utilis yeast cells to immunolabelling with β-(1,3)-glucan monoclonal antibodies? Microb. Cell Fact.23, 28. doi: 10.1186/s12934-024-02305-4

  • CrossRef
  • Google Scholar
  • View reference in article

• 30

  CazónP.VelazquezG.RamírezJ. A.VázquezM. (2017). Polysaccharide-based films and coatings for food packaging: A review. Food Hydrocoll.68, 136–148. doi: 10.1016/j.foodhyd.2016.09.009

  • CrossRef
  • Google Scholar

• 31

  CerlettiC.EspositoS.IacovielloL. (2021). Edible mushrooms and beta-glucans: impact on human health. Nutrients.13, e2195. doi: 10.3390/nu13072195

  • CrossRef
  • Google Scholar

• 32

  CharouxC. M. G.OjhaK. S.O’DonnellC. P.CardoniA.TiwariB. K. (2017). Applications of airborne ultrasonic technology in the food industry. J. Food Eng.208, 28–36. doi: 10.1016/j.jfoodeng.2017.03.030

  • CrossRef
  • Google Scholar

• 33

  ChematF.RombautN.SicaireA.-G.MeullemiestreA.Fabiano-TixierA.-S.Abert-VianM. (2017). Ultrasound assisted extraction of food and natural products. Mechanisms, techniques, combinations, protocols and applications. A review. Ultrason. Sonochem.34, 540–560. doi: 10.1016/j.ultsonch.2016.06.035

  • CrossRef
  • Google Scholar

• 34

  ChenY.HuangY.CuiZ.LiuJ. (2015). Purification, characterization and biological activity of a novel polysaccharide from Inonotus obliquus. Int. J. Biol. Macromol.79, 587–594. doi: 10.1016/j.ijbiomac.2015.05.016

  • CrossRef
  • Google Scholar

• 35

  ChenY. F.JiangW. W.ZhangS. Q.KanJ. Q.LiangY. (2016). Antioxidant activity and characterization of one new polysaccharide obtained from Perigord Truffle (Tuber huidongense). Evid. Based Complementary Altern. Med.2016, 3537193. doi: 10.1155/2016/3537193

  • CrossRef
  • Google Scholar

• 36

  ChenJ.LaiP.ShenH.ZhenH.FangR. (2013). Effect of extraction methods on polysaccharide of Clitocybe maxima Stipe. Adv. J. Food Sci. Technol.5, 370–373. doi: 10.19026/ajfst.5.3273

  • CrossRef
  • Google Scholar

• 37

  ChenY.LiuY.SarkerM. M. R.YanX.YangC.ZhaoL.et al. (2018). Structural characterization and antidiabetic potential of a novel heteropolysaccharide from Grifola frondosa via IRS1/PI3K-JNK signaling pathways. Carbohydr. Polym.198, 452–461. doi: 10.1016/j.carbpol.2018.06.077

  • CrossRef
  • Google Scholar

• 38

  ChenY.LiuD.WangD.LaiS.ZhongR.LiuY.et al. (2019). Hypoglycemic activity and gut microbiota regulation of a novel polysaccharide from Grifola frondosa in type 2 diabetic mice. Food Chem. Toxicol.126, 295–302. doi: 10.1016/j.fct.2019.02.034

  • CrossRef
  • Google Scholar

• 39

  ChenZ.YinC.FanX.MaK.YaoF.ZhouR.et al. (2020). Characterization of physicochemical and biological properties of Schizophyllum commune polysaccharide extracted with different methods. Int. J. Biol. Macromol.156, 1425–1434. doi: 10.1016/j.ijbiomac.2019.11.183

  • CrossRef
  • Google Scholar

• 40

  ChenJ.YuM.YangC.HuangZ.HeL.BianJ.et al. (2024). Study on the nutritional relationships in mycelia and fruiting bodies of Hypsizygus marmoreus under defined nutrient conditions. Food Chem.467, 142323. doi: 10.1016/j.foodchem.2024.142323

  • CrossRef
  • Google Scholar

• 41

  ChengK. C.DemirciA.CatchmarkJ. M. (2011). Pullulan: biosynthesis, production, and applications. Appl. Microbiol. Biotechnol.92, 29–44. doi: 10.1007/s00253-011-3477-y

  • CrossRef
  • Google Scholar

• 42

  ChengB.YuK.WengX.LiuZ.HuangX.JiangY.et al. (2024). Impact of cell wall polysaccharide modifications on the performance of Pichia pastoris: novel mutants with enhanced fitness and functionality for bioproduction applications. Microb. Cell Fact.23, e55. doi: 10.1186/s12934-024-02333-0

  • CrossRef
  • Google Scholar

• 43

  ChioruA.ChirsanovaA. (2023). β-Glucans: characterization, extraction methods, and valorization. Food Nutr. Sci.14, 963–983. doi: 10.4236/fns.2023.1410061

  • CrossRef
  • Google Scholar

• 44

  ChristyaniG.CarswellM.QinS.KimW. (2024). An overview of advances in rare cancer diagnosis and treatment. Int. J. Mol. Sci.25, e1201. doi: 10.3390/ijms25021201

  • CrossRef
  • Google Scholar

• 45

  ChuensunT.ChewonarinT.LaopajonW.Kawee-aiA.PinpartP.Utama-angN. (2021). Comparative evaluation of physicochemical properties of Lingzhi (Ganoderma lucidum) as affected by drying conditions and extraction methods. Int. J. Food Sci. Technol.56, 2751–2759. doi: 10.1111/ijfs.14906

  • CrossRef
  • Google Scholar

• 46

  CostaO. Y. A.RaaijmakersJ. M.KuramaeE. E. (2018). Microbial extracellular polymeric substances: Ecological function and impact on soil aggregation. Front. Microbiol.9, e1636. doi: 10.3389/fmicb.2018.01636

  • CrossRef
  • Google Scholar

• 47

  CuiF. J.QianL. S.SunW. J.ZhangJ. S.YangY.LiN.et al. (2018). Ultrasound-assisted extraction of polysaccharides from Volvariella volvacea: Process optimization and structural characterization. Molecules23, e1706. doi: 10.3390/molecules23071706

  • CrossRef
  • Google Scholar

• 48

  DaiY.WangL.ChenX.SongA.HeL.WangL.et al. (2023). Lentinula edodes Sing polysaccharide: extraction, characterization, bioactivities, and emulsifying applications. Foods12, e3289. doi: 10.3390/foods12173289

  • CrossRef
  • Google Scholar

• 49

  DaynesC. N.ZhangN.SaleebaJ. A.McGeeP. A. (2012). Soil aggregates formed in vitro by saprotrophic Trichocomaceae have transient water-stability. Soil Biol. Biochem.48, 151–161. doi: 10.1016/j.soilbio.2012.01.010

  • CrossRef
  • Google Scholar

• 50

  De CostaG.ParkA. (2017). Hyperlipidaemia. Medicine45, 579–582. doi: 10.1016/j.mpmed.2017.06.002

  • CrossRef
  • Google Scholar

• 51

  De FeliceB.DamianoS.MontaninoC.Del BuonoA.La RosaG.GuidaB.et al. (2020). Effect of beta- and alpha-glucans on immune modulating factors expression in enterocyte-like Caco-2 and goblet-like LS 174T cells. Int. J. Biol. Macromol.153, 600–607. doi: 10.1016/j.ijbiomac.2020.03.046

  • CrossRef
  • Google Scholar

• 52

  DelzenneN. M.BindelsL. B. (2015). Gut microbiota: Ganoderma lucidum, a new prebiotic agent to treat obesity?: Gut microbiota. Nat. Rev. Gastroenterol. Hepatol.12, 553–554. doi: 10.1038/nrgastro.2015.137

  • CrossRef
  • Google Scholar

• 53

  DengY.HuangQ.HuL.LiuT.ZhengB.LuD.et al. (2021). Enhanced exopolysaccharide yield and antioxidant activities of Schizophyllum commune fermented products by the addition of Radix Puerariae. RSC Adv.11, 38219–38234. doi: 10.1039/d1ra06314f

  • CrossRef
  • Google Scholar

• 54

  DengQ.ZinoviadouK. G.GalanakisC. M.OrlienV.GrimiN.VorobievE.et al. (2015). The effects of conventional and non-conventional processing on glucosinolates and its derived forms, isothiocyanates: extraction, degradation, and applications. Food Eng. Rev.7, 357–381. doi: 10.1007/s12393-014-9104-9

  • CrossRef
  • Google Scholar

• 55

  DeveciE.Tel-CayanG.AltinokB. Y.AktasS. (2024). Characterization of polysaccharide extracts of four edible mushrooms and determination of in vitro antioxidant, enzyme inhibition and anticancer activities. ACS Omega9, 25887–25901. doi: 10.1021/acsomega.4c00322

  • CrossRef
  • Google Scholar

• 56

  Di MartinoP. (2022). Antimicrobial agents and microbial ecology. AIMS Microbiol.8, 1–4. doi: 10.3934/microbiol.2022001

  • CrossRef
  • Google Scholar

• 57

  DobšíkováR.BlahováJ.MikulíkováI.ModráH.PráškováE.SvobodováZ.et al. (2013). The effect of oyster mushroom β-1.3/1.6-D-glucan and oxytetracycline antibiotic on biometrical, haematological, biochemical, and immunological indices, and histopathological changes in common carp (Cyprinus carpio L.). Fish Shellfish Immunol.35, 1813–1823. doi: 10.1016/j.fsi.2013.09.006

  • CrossRef
  • Google Scholar

• 58

  DongX.SunS.WangX.YuH.DaiK.JiaoJ.et al. (2024). Structural characteristics and intestinal flora metabolism mediated immunoregulatory effects of Lactarius deliciosus polysaccharide. Int. J. Biol. Macromol.278, e135063. doi: 10.1016/j.ijbiomac.2024.135063

  • CrossRef
  • Google Scholar

• 59

  DongY.ZhangJ.GaoZ.ZhaoH.SunG.WangX.et al. (2019). Characterization and anti-hyperlipidemia effects of enzymatic residue polysaccharides from Pleurotus ostreatus. Int. J. Biol. Macromol.129, 316–325. doi: 10.1016/j.ijbiomac.2019.01.164

  • CrossRef
  • Google Scholar

• 60

  DornellesG.AraújoG. R.deS.RodriguesM.AlvesV.Almeida-PaesR.et al. (2023). Comparative analysis of capsular and secreted polysaccharides produced by Rhodotorula mucilaginosa and Cryptococcus neoformans. J. Fungi9, e1124. doi: 10.3390/jof9111124

  • CrossRef
  • Google Scholar

• 61

  DuobinM.YupingM.LujingG.AijingZ.JianqiangZ.ChunpingX. (2013). Fermentation characteristics in stirred-tank reactor of exopolysaccharides with hypolipidemic activity produced by Pleurotus geesteranus 5#. An. Acad. Bras. Cienc.85, 1473–1481. doi: 10.1590/0001-3765201320130037

  • CrossRef
  • Google Scholar

• 62

  ElisashviliV. (2012). Submerged cultivation of medicinal mushrooms: Bioprocesses and products (review). Int. J. Med. Mushrooms14, 211–239. doi: 10.1615/intjmedmushr.v14.i3.10

  • CrossRef
  • Google Scholar

• 63

  ElisashviliV. I.KachlishviliE. T.WasserS. P. (2009). Carbon and nitrogen source effects on basidiomycetes exopolysaccharide production. Appl. Biochem. Microbiol.45, 531–535. doi: 10.1134/s0003683809050135

  • CrossRef
  • Google Scholar

• 64

  El-MahdyO. M.MohamedH. I.El-AnsaryA. E. (2023). Optimizations of exopolysaccharide production by Fusarium nygamai strain AJTYC1 and its potential applications as an antioxidant, antimicrobial, anticancer, and emulsifier. BMC Microbiol.23, e345. doi: 10.1186/s12866-023-03100-8

  • CrossRef
  • Google Scholar

• 65

  ElshafieA.JoshiS. J.Al-WahaibiY. M.Al-BahryS. N.Al-BemaniA. S.Al-HashmiA.et al. (2017). Isolation and characterization of biopolymer producing Omani Aureobasidium pullulans strains and its potential applications in microbial enhanced oil recovery. Soc Pet. Eng.2017, 583–593. doi: 10.2118/185326-MS

  • CrossRef
  • Google Scholar

• 66

  EzikeT. C.OkpalaU. S.OnojaU. L.NwikeC. P.EzeakoE. C.OkparaO. J.et al. (2023). Advances in drug delivery systems, challenges and future directions. Heliyon9, e17488. doi: 10.1016/j.heliyon.2023.e17488

  • CrossRef
  • Google Scholar

• 67

  FellakS.RafikM.HaidaraH.TaybiH.BoukirA.LhassaniA. (2022). Scanning electron microscopy examination of the surface of softwood attacked by fungus. MATEC Web Conf.360, e00007. doi: 10.1051/matecconf/202236000007

  • CrossRef
  • Google Scholar

• 68

  FernandoL. D.Dickwella WidanageM. C.PenfieldJ.LiptonA. S.WashtonN.LatgéJ. P.et al. (2021). Structural polymorphism of chitin and chitosan in fungal cell walls from solid-state NMR and principal component analysis. Front. Mol. Biosci.8, e727053. doi: 10.3389/fmolb.2021.727053

  • CrossRef
  • Google Scholar

• 69

  FerraboschiP.CiceriS.GrisentiP. (2021). Applications of lysozyme, an innate immune defense factor, as an alternative antibiotic. Antibiotics10, e1534. doi: 10.3390/antibiotics10121534

  • CrossRef
  • Google Scholar

• 70

  FigueroaG. M.CostaD. N.MirandaA. C. A.JuniorG. L. V.MendesT. P. S.CedroP. E. P. (2023). Fungal polysaccharides and their hypoglycemic potential to act as an adjuvant in the treatment of diabetes mellitus. Evidência23, 47–62. doi: 10.18593/evid.32561

  • CrossRef
  • Google Scholar

• 71

  FonsecaP. R. M. S.DekkerR. F. H.BarbosaA. M.SilveiraJ. L. M.VasconcelosA. F. D.MonteiroN. K.et al. (2011). Thermal and rheological properties of a family of botryosphaerans produced by Botryosphaeria rhodina MAMB-05. Molecules16, 7488–7501. doi: 10.3390/molecules16097488

  • CrossRef
  • Google Scholar

• 72

  FosmerA.GibbonsW. (2011). Separation of scleroglucan and cell biomass from Sclerotium glucanicum grown in an inexpensive, by-product based medium. Int. J. Agric. Biol. Eng.4, e52. doi: 10.3965/j.issn.1934-6344.2011.01.0-0

  • CrossRef
  • Google Scholar

• 73

  FuY.ShiL.DingK. (2019). Structure elucidation and anti-tumor activity in vivo of a polysaccharide from spores of Ganoderma lucidum (Fr.) Karst. Int. J. Biol. Macromol.141, 693–699. doi: 10.1016/j.ijbiomac.2019.09.046

  • CrossRef
  • Google Scholar

• 74

  FuY.WangL.JiangG.RenL.WangL.LiuX. (2022). Anti-diabetic activity of polysaccharides from Auricularia cornea var. Li. Food11, e1464. doi: 10.3390/foods11101464

  • CrossRef
  • Google Scholar

• 75

  GaoL.WangY.ZhangF.LiS.ZhaoJ.ZhangQ.et al. (2022). A standardized method for the quantification of polysaccharides: An example of polysaccharides from Tremella fuciformis. LWT- Food Sci. Technol.167, e113860. doi: 10.1016/j.lwt.2022.113860

  • CrossRef
  • Google Scholar

• 76

  GaoY. Y.ZhouY. H.LiuX. P.DiB.HeJ. Y.WangY. T.et al. (2024). Ganoderma lucidum polysaccharide promotes broiler health by regulating lipid metabolism, antioxidants, and intestinal microflora. Int. J. Biol. Macromol.280, e135918. doi: 10.1016/j.ijbiomac.2024.135918

  • CrossRef
  • Google Scholar

• 77

  GbashiS.AdeboO. A.PiaterL.MadalaN. E.NjobehP. B. (2017). Subcritical water extraction of biological materials. Sep. Purif. Rev.46, 21–34. doi: 10.1080/15422119.2016.1170035

  • CrossRef
  • Google Scholar

• 78

  GeY.QiuH.ZhengJ. (2022). Physicochemical characteristics and anti-hyperlipidemic effect of polysaccharide from BaChu mushroom (Helvella leucopus). Food Chem.15, e100443. doi: 10.1016/j.fochx.2022.100443

  • CrossRef
  • Google Scholar

• 79

  GehrckeM.MartinsC. C.de Bastos BrumT.da RosaL. S.LucheseC.WilhelmE. A.et al. (2022). Novel pullulan/gellan gum bilayer film as a vehicle for silibinin-loaded nanocapsules in the topical treatment of atopic dermatitis. Pharmaceutics14, e2352. doi: 10.3390/pharmaceutics14112352

  • CrossRef
  • Google Scholar

• 80

  GeraldelliD.RibeiroM. C.MedeirosT. C.ComiranP. K.MartinsK. O.OliveiraM. F.et al. (2020). Botryosphaeran, a (1→3)(1→6)-β-D-glucan, reduces tumor development and cachexia syndrome in obese male rats by increasing insulin sensitivity and FOXO3a activity. Int. J. Biol. Macromol.165, 985–994. doi: 10.1016/j.ijbiomac.2020.09.168

  • CrossRef
  • Google Scholar

• 81

  GhoshT.PriyadarshiR.Krebs de SouzaC.AngiolettiB. L.RhimJ. W. (2022). Advances in pullulan utilization for sustainable applications in food packaging and preservation: A mini-review. Trends Food Sci. Technol.125, 43–53. doi: 10.1016/j.tifs.2022.05.001

  • CrossRef
  • Google Scholar

• 82

  GiannenasI.TsalieE.ChronisE.MavridisS.TontisD.KyriazakisI. (2011). Consumption of Agaricus bisporus mushroom affects the performance, intestinal microbiota composition and morphology, and antioxidant status of Turkey poults. Anim. Feed Sci. Technol.165, 218–229. doi: 10.1016/j.anifeedsci.2011.03.002

  • CrossRef
  • Google Scholar

• 83

  GientkaI.Bzducha-WróbelA.Stasiak-RóżańskaL.BednarskaA. A.BłażejakS. (2016). The exopolysaccharides biosynthesis by Candida yeast depends on carbon sources. Electron. J. Biotechnol.22, 31–37. doi: 10.1016/j.ejbt.2016.02.008

  • CrossRef
  • Google Scholar

• 84

  Gil-RamírezA.SmiderleF. R.MoralesD.IacominiM.Soler-RivasC. (2019). Strengths and weaknesses of the aniline-blue method used to test mushroom (1→3)-β-d-glucans obtained by microwave-assisted extractions. Carbohydr. Polym.217, 135–143. doi: 10.1016/j.carbpol.2019.04.051

  • CrossRef
  • Google Scholar

• 85

  GniewoszM.PobiegaK.KraśniewskaK.SynowiecA.ChaberekM.GalusS. (2022). Characterization and antifungal activity of pullulan edible films enriched with Propolis extract for active packaging. Foods11, 2319. doi: 10.3390/foods11152319

  • CrossRef
  • Google Scholar

• 86

  GolovchenkoV. V.NaranmandakhS.GanbaatarJ.PrilepskiiA. Y.BuryginG. L.ChizhovA. O.et al. (2020). Structural investigation and comparative cytotoxic activity of water-soluble polysaccharides from fruit bodies of the medicinal fungus quinine conk. Phytochemistry175, e112313. doi: 10.1016/j.phytochem.2020.112313

  • CrossRef
  • Google Scholar

• 87

  GonçalvesB. C.Lopes BarbosaM. G.Silva OlakA. P.Belebecha TerezoN.NishiL.WatanabeM. A.et al. (2021). Antiviral therapies: advances and perspectives. Fundam. Clin. Pharmacol.35, 305–320. doi: 10.1111/fcp.12609

  • CrossRef
  • Google Scholar

• 88

  GongP.LongH.GuoY.WangS.ChenF.ChenX. (2022). Isolation, structural characterization, and hypoglycemic activities in vitro of polysaccharides from Pleurotus eryngii. Molecules27, e7140. doi: 10.3390/molecules27207140

  • CrossRef
  • Google Scholar

• 89

  GovindanS.ShanmugamJ.RajendranG.RamaniP.UnniD.VenkatachalamB.et al. (2023). Antidiabetic activity of polysaccharide from Hypsizygus ulmarius in streptozotocin-nicotinamide induced diabetic rats. Bioact. Carbohydr. Diet. Fibre29, 100350. doi: 10.1016/j.bcdf.2023.100350

  • CrossRef
  • Google Scholar

• 90

  GrimmD.WöstenH. A. B. (2018). Mushroom cultivation in the circular economy. Appl. Microbiol. Biotechnol.102, 7795–7803. doi: 10.1007/s00253-018-9226-8

  • CrossRef
  • Google Scholar

• 91

  GuoC.GuoD.FangL.SangT.WuJ.GuoC.et al. (2021). Ganoderma lucidum polysaccharide modulates gut microbiota and immune cell function to inhibit inflammation and tumorigenesis in colon. Carbohydr. Polym.267, e118231. doi: 10.1016/j.carbpol.2021.118231

  • CrossRef
  • Google Scholar

• 92

  GuoM. Q.HuX.WangC.AiL. (2017). “Polysaccharides: structure and solubility,” in Solubility of polysaccharides. Ed. XuZ. (InTech, London, UK). doi: 10.5772/intechopen.71570

  • CrossRef
  • Google Scholar

• 93

  GuoL.YaoQ.LvJ.LiZ.WangL. A.ZhangJ. (2024a). Anti-hyperglycemic effect of the brown slime cap mushroom Chroogomphus rutilus (Agaricomycetes) crude polysaccharide in vitro and in vivo. Int. J. Med. Mushrooms26, 1–12. doi: 10.1615/IntJMedMushrooms.2024053173

  • CrossRef
  • Google Scholar

• 94

  GuoW.YunJ.WangB.XuS.YeC.WangX.et al. (2024b). Comparative study on physicochemical properties and hypoglycemic activities of intracellular and extracellular polysaccharides from submerged fermentation of Morchella esculenta. Int. J. Biol. Macromol.278, e134759. doi: 10.1016/j.ijbiomac.2024.134759

  • CrossRef
  • Google Scholar

• 95

  HabibiE.HemmatiP.ArabnozariH.KhaliliH. A.SharifianjaziF.EnderamiS. E.et al. (2024). Phytochemical analysis and immune-modulatory potential of Trichaptum biforme polysaccharides: Implications for cancer. Int. J. Biol. Macromol.280, e135691. doi: 10.1016/j.ijbiomac.2024.135691

  • CrossRef
  • Google Scholar

• 96

  HamidiM.OkoroO. V.MilanP. B.KhaliliM. R.SamadianH.NieL.et al. (2022). Fungal exopolysaccharides: properties, sources, modifications, and biomedical applications. Carbohydr. Polym.284, e119152. doi: 10.1016/j.carbpol.2022.119152

  • CrossRef
  • Google Scholar

• 97

  HaoH.CuiC.XingY.JiaX.MaB.KangW.et al. (2023a). Sulfation of the extracellular polysaccharide from the edible fungus Stropharia rugosoannulata with its antioxidant activity. J. Future Foods3, 37–42. doi: 10.1016/j.jfutfo.2022.09.006

  • CrossRef
  • Google Scholar

• 98

  HaoL.ShengZ.LuJ.TaoR.JiaS. (2016). Characterization and antioxidant activities of extracellular and intracellular polysaccharides from Fomitopsis pinicola. Carbohydr. Polym141, 54–59. doi: 10.1016/j.carbpol.2015.11.048

  • CrossRef
  • Google Scholar

• 99

  HaoR.ZhouX.ZhaoX.LvX.ZhuX.GaoN.et al. (2023b). Flammulina velutipes polysaccharide counteracts cadmium-induced gut injury in mice via modulating gut inflammation, gut microbiota and intestinal barrier. Sci. Total Environ.877, e162910. doi: 10.1016/j.scitotenv.2023.162910

  • CrossRef
  • Google Scholar

• 100

  HeJ.LiuZ.JiangW.ZhuT.WusimanA.GuP.et al. (2020b). Immune-adjuvant activity of lentinan-modified calcium carbonate microparticles on a H5N1 vaccine. Int. J. Biol. Macromol.163, 1384–1392. doi: 10.1016/j.ijbiomac.2020.08.005

  • CrossRef
  • Google Scholar

• 101

  HeB. L.ZhengQ. W.GuoL. Q.HuangJ. Y.YunF.HuangS. S.et al. (2020a). Structural characterization and immune-enhancing activity of a novel high-molecular-weight polysaccharide from Cordyceps militaris. Int. J. Biol. Macromol.145, 11–20. doi: 10.1016/j.ijbiomac.2019.12.115

  • CrossRef
  • Google Scholar

• 102

  HerreroM.CifuentesA.IbanezE. (2006). Sub- and supercritical fluid extraction of functional ingredients from different natural sources: Plants, food-by-products, algae and microalgae A review. Food Chem.98, 136–148. doi: 10.1016/j.foodchem.2005.05.058

  • CrossRef
  • Google Scholar

• 103

  HoaT. Q. (2018). Extraction of polysaccharides from Lingzhi by ultrasonic-assisted enzymatic method. Vietnam J. Sci. Technol.56, e171. doi: 10.15625/2525-2518/56/4a/12879

  • CrossRef
  • Google Scholar

• 104

  HongT.YinJ. Y.NieS. P.XieM. Y. (2021). Applications of infrared spectroscopy in polysaccharide structural analysis: Progress, challenge and perspective. Food Chem.: X12, e100168. doi: 10.1016/j.fochx.2021.100168

  • CrossRef
  • Google Scholar

• 105

  HsuK. D.WuS. P.LinS. P.LumC. C.ChengK. C. (2017). Enhanced active extracellular polysaccharide production from Ganoderma formosanum using computational modeling. J. Food Drug Anal.25, 804–811. doi: 10.1016/j.jfda.2016.12.006

  • CrossRef
  • Google Scholar

• 106

  HuQ.HeY.WangF.WuJ.CiZ.ChenL.et al. (2021). Microwave technology: a novel approach to the transformation of natural metabolites. Chin. Med.16, e87. doi: 10.1186/s13020-021-00500-8

  • CrossRef
  • Google Scholar

• 107

  HuY.XuJ.ShengY.LiuJ.LiH.GuoM.et al. (2022a). Pleurotus ostreatus ameliorates obesity by modulating the gut microbiota in obese mice induced by high-fat diet. Nutrients14, e1868. doi: 10.3390/nu14091868

  • CrossRef
  • Google Scholar

• 108

  HuZ.YuR.SunJ.DuanY.ZhouH.ZhouW.et al. (2022b). Static decolorization of polysaccharides from the leaves of Rhododendron dauricum: Process optimization, characterization and antioxidant activities. Proc. Biochem.121, 113–125. doi: 10.1016/j.procbio.2022.06.025

  • CrossRef
  • Google Scholar

• 109

  HuangZ. F.ZhangM. L.ZhangS.WangY. H.JiangX. W. (2018). Structural characterization of polysaccharides from Cordyceps militaris and their hypolipidemic effects in high fat diet fed mice. RSC Adv.8, 41012–41022. doi: 10.1039/c8ra09068h

  • CrossRef
  • Google Scholar

• 110

  HuberV.MullerL.DegotP.TouraudD.KunzW. (2021). NADES-based surfactant-free microemulsions for solubilization and extraction of curcumin from Curcuma Longa. Food Chem.355, e129624. doi: 10.1016/j.foodchem.2021.129624

  • CrossRef
  • Google Scholar

• 111

  HwangA. Y.YangS. C.KimJ.LimT.ChoH.HwangK. T. (2019). Effects of non-traditional extraction methods on extracting bioactive compounds from chaga mushroom (Inonotus obliquus) compared with hot water extraction. LWT-Food Sci. Technol.110, 80–84. doi: 10.1016/j.lwt.2019.04.073

  • CrossRef
  • Google Scholar

• 112

  InaK.FurutaR.KataokaT.KayukawaS.YoshidaT.MiwaT.et al. (2011). Lentinan prolonged survival in patients with gastric cancer receiving S-1-based chemotherapy. World J. Clin. Oncol.2, 339–343. doi: 10.5306/wjco.v2.i10.339

  • CrossRef
  • Google Scholar

• 113

  IngaleA.WadherD. S. J.BagulS.GujratiP.GovandeA. (2023). High performance liquid chromatography: An overview. Int. J. Pharm. Sci. Rev. Res.3, 18–24. doi: 10.47583/ijpsrr.2023.v82i02.003

  • CrossRef
  • Google Scholar

• 114

  JafarinejadS. (2017). “Chapter 7,” in Petroleum waste treatment and pollution control (Butterworth Heinemann, Amsterdam, Netherlands), 269–270.

  • Google Scholar

• 115

  JayusJ.AkromanR.NugrahaA. S.PiluhartoB.SeviourR. J. (2021). Structural elucidation of the exopolysaccharide produced by Curvularia lunata isolate RJ01. Biodiversitas22, 2699–2705. doi: 10.13057/biodiv/d220530

  • CrossRef
  • Google Scholar

• 116

  JenC. I.LuM. K.LaiM. N.NgL. T. (2024). Sulfated polysaccharides of Laetiporus sulphureus fruiting bodies exhibit anti-breast cancer activity through cell cycle arrest, apoptosis induction, and inhibiting cell migration. J. Ethnopharmacol.321, e117546. doi: 10.1016/j.jep.2023.117546

  • CrossRef
  • Google Scholar

• 117

  JeongS. C.KoyyalamudiS. R.HughesJ.KhooC.BaileyT.MarripudiK.et al. (2013). Antioxidant and immunomodulating activities of exo-and endopolysaccharide fractions from submerged mycelia cultures of culinary-medicinal mushrooms. Int. J. Med. Mushrooms15, 251–266. doi: 10.1615/intjmedmushr.v15.i3.30

  • CrossRef
  • Google Scholar

• 118

  JesenakM.UrbancekS.MajtanJ.BanovcinP.HercogovaJ. (2016). β-Glucan-based cream (containing pleuran isolated from Pleurotus ostreatus) in supportive treatment of mild-to-moderate atopic dermatitis. J. Dermatolog. Treat.27, 351–354. doi: 10.3109/09546634.2015.1117565

  • CrossRef
  • Google Scholar

• 119

  JiX.GuoJ.CaoT.ZhangT.LiuY.YanY. (2023). Review on mechanisms and structure-activity relationship of hypoglycemic effects of polysaccharides from natural resources. Food Sci. Hum. Wellness12, 1969–1980. doi: 10.1016/j.fshw.2023.03.017

  • CrossRef
  • Google Scholar

• 120

  JiangX.HaoJ.ZhuY.LiuZ.LiL.ZhouY.et al. (2022). The anti-obesity effects of a water-soluble glucan from Grifola frondosa via the modulation of chronic inflammation. Front. Immunol.3, e962341. doi: 10.3389/fimmu.2022.962341

  • CrossRef
  • Google Scholar

• 121

  JiaoF.LiD.YangS.ZhangJ.ZhangC.JiaL. (2018). Inhibition effects of polysaccharides on HBV replication and cell proliferation from Lentinus edodes waste material. Micro. Pathog.123, 461–466. doi: 10.1016/j.micpath.2018.08.004

  • CrossRef
  • Google Scholar

• 122

  JinJ.NguyenT. T. H.KimC.KimD. (2019). Antimelanogenesis effects of fungal exopolysaccharides prepared from submerged culture of Fomitopsis castanea mycelia. J. Microbiol. Biotechnol.29, 1204–1211. doi: 10.4014/jmb.1905.05037

  • CrossRef
  • Google Scholar

• 123

  JinX.ZhaoS. (2014). Extraction optimization and bioactivities of an extracellular polysaccharide produced by Aspergillus fumigatus. Int. J. Biol. Macromol.68, 13–17. doi: 10.1016/j.ijbiomac.2014.04.024

  • CrossRef
  • Google Scholar

• 124

  JindalN.Singh KhattarJ. (2018). “Microbial polysaccharides in food industry”, in Biopolymers for Food Design, e.d. GrumezescuA. M.HolbanA. M. (Cambridge: Elsevier Inc.), 95–123. doi: 10.1016/B978-0-12-811449-0.00004-9

  • CrossRef
  • Google Scholar

• 125

  JoshiS. J.Al-WahaibiY. M.Al-BahryS.ElshafieA.Al-BemaniA. S.Al-HashmiA.et al. (2016). Production and application of schizophyllan in microbial enhanced heavy oil recovery. SPE EOR Conference at Oil and Gas West Asia. Oman: Society of Petroleum Engineers.

  • Google Scholar

• 126

  JovanovićJ. A.MihailovićM.UskokovićA.GrdovićN.DicnićS.VidakovićM. (2021). The effects of major mushroom bioactive compounds on mechanisms that control blood glucose level. J. Fungi7, e58. doi: 10.3390/jof7010058

  • CrossRef
  • Google Scholar

• 127

  Junior LettiL. A.Destéfanis VítolaF. M.Vinícius de Melo PereiraG.KarpS. G.Pedroni MedeirosA. B.Ferreira da CostaE. S.et al. (2018). “Solid-state fermentation for the production of mushrooms,” in Current developments in biotechnology and bioengineering (Elsevier, Amsterdam, The Netherlands), 285–318. doi: 10.1016/B978-0-444-63990-5.00014-1

  • CrossRef
  • Google Scholar

• 128

  KadookaC.TanakaY.HiraD.MaruyamaJ.-I.GotoM.OkaT. (2023). Identification of galactofuranose antigens such as galactomannoproteins and fungal-type galactomannan from the yellow koji fungus (Aspergillus oryzae). Front. Microbiol.14, e1110996. doi: 10.3389/fmicb.2023.1110996

  • CrossRef
  • Google Scholar

• 129

  KangJ. Y.LeeB.KimC. H.ChoiJ. H.KimM. S. (2022). Enhancing the prebiotic and antioxidant effects of exopolysaccharides derived from Cordyceps militaris by enzyme-digestion. LWT-Food Sci. Technol.167, e113830. doi: 10.1016/j.lwt.2022.113830

  • CrossRef
  • Google Scholar

• 130

  KangJ. H.TachibanaY.KamataW.MaharaA.Harada-ShibaM.YamaokaT. (2010). Liver-targeted siRNA delivery by polyethylenimine (PEI)-pullulan carrier. Bioorg. Med. Chem.18, 3946–3950. doi: 10.1016/j.bmc.2010.04.031

  • CrossRef
  • Google Scholar

• 131

  KanlayavattanakulM.LourithN. (2023). Cordyceps militaris polysaccharides: preparation and topical product application. Fungal Biol. Biotechnol.10, e3. doi: 10.1186/s40694-023-00150-5

  • CrossRef
  • Google Scholar

• 132

  KanwalS.AliyaS.XinY. (2020). Anti-obesity effect of Dictyophora indusiate mushroom polysaccharide (DIP) in high fat diet-induced obesity via regulating inflammatory cascades and intestinal microbiome. Front. Endocrinol.11, e558874. doi: 10.3389/fendo.2020.558874

  • CrossRef
  • Google Scholar

• 133

  KarimA.GerlianiN.AïderM. (2020). Kluyveromyces marxianus: An emerging yeast cell factory for applications in food and biotechnology. Int. J. Food Microbiol.333, e108818. doi: 10.1016/j.ijfoodmicro.2020.108818

  • CrossRef
  • Google Scholar

• 134

  KawaharaH.MatsudaY.SakaguchiT.AraiN.KoideY. (2016). Antifreeze activity of xylomannan from the mycelium and fruit body of Flammulina velutipes. Biocontrol Sci.21, 153–159. doi: 10.4265/bio.21.153

  • CrossRef
  • Google Scholar

• 135

  KeL. Q. (2015). Optimization of ultrasonic extraction of polysaccharides from Lentinus edodes based on enzymatic treatment: Extraction of Lentinus edodes polysaccharides. J. Food Proces. Preserv.39, 254–259. doi: 10.1111/jfpp.12228

  • CrossRef
  • Google Scholar

• 136

  KhanA. A.GaniA.KhandayF. A.MasoodiF. A. (2018). Biological and pharmaceutical activities of mushroom β-glucan discussed as a potential functional food ingredient. Bioact. Carbohydr. Diet. Fibre16, 1–13. doi: 10.1016/j.bcdf.2017.12.002

  • CrossRef
  • Google Scholar

• 137

  KhatuaS.AcharyaK. (2017). Alkaline extractive crude polysaccharide from Russula senecis possesses antioxidant potential and stimulates innate immunity response. J. Pharm. Pharmacol.69, 1817–1828. doi: 10.1111/jphp.12813

  • CrossRef
  • Google Scholar

• 138

  KhouryZ. H.VilaT.PuthranT. R.SultanA. S.Montelongo-JaureguiD.MeloM. A. S.et al. (2020). The role of Candida albicans secreted polysaccharides in augmenting streptococcus mutans adherence and mixed biofilm formation: in vitro and in vivo studies. Front. Microbiol.11, e307. doi: 10.3389/fmicb.2020.00307

  • CrossRef
  • Google Scholar

• 139

  KizitskaT.BarshteynV.SevindikM.KrupodorovaT. (2024). Evaluation of Fomitopsis betulina strains for growth on different media and exopolysaccharide production. Arch. Biol. Sci.76, 257–265. doi: 10.2298/ABS240523018K

  • CrossRef
  • Google Scholar

• 140

  Larrañaga-OrdazD.Martínez-MaldonadoM.Millán-ChiuB. E.FernándezF.Castaño-TostadoE.Gómez-LimM.Á.et al. (2022). Effect of shock waves on the growth of Aspergillus niger conidia: Evaluation of germination and preliminary study on gene expression. J. Fungi8, e1117. doi: 10.20944/preprints202208.0455.v1

  • CrossRef
  • Google Scholar

• 141

  LazićV.KlausA.KozarskiM.DoroškiA.TostiT.SimićS.et al. (2024). The effect of green extraction technologies on the chemical composition of medicinal Chaga mushroom extracts. J. Fungi10, e225. doi: 10.3390/jof10030225

  • CrossRef
  • Google Scholar

• 142

  LeongY. K.YangF.-C.ChangJ.-S. (2021). Extraction of polysaccharides from edible mushrooms: Emerging technologies and recent advances. Carbohydr. Polym.251, e117006. doi: 10.1016/j.carbpol.2020.117006

  • CrossRef
  • Google Scholar

• 143

  LeungP. H.ZhaoS.HoK. P.WuJ. Y. (2009). Chemical properties and antioxidant activity of exopolysaccharides from mycelial culture of Cordyceps sinensis fungus Cs-HK1. Food Chem.114, 1251–1256. doi: 10.1016/j.foodchem.2008.10.081

  • CrossRef
  • Google Scholar

• 144

  LiJ.CaiC.ZhengM.HaoJ.WangY.HuM.et al. (2019a). Alkaline extraction, structural characterization, and bioactivities of (1→6)-β-d-glucan from Lentinus edodes. Molecules24, e1610. doi: 10.3390/molecules24081610

  • CrossRef
  • Google Scholar

• 145

  LiJ.GuF.CaiC.HuM.FanL.HaoJ.et al. (2020a). Purification, structural characterization, and immunomodulatory activity of the polysaccharides from Ganoderma lucidum. Int. J. Biol. Macromol.143, 806–813. doi: 10.1016/j.ijbiomac.2019.09.141

  • CrossRef
  • Google Scholar

• 146

  LiF.LeiH.XuH. (2022). Influences of subcritical water extraction on the characterization and biological properties of polysaccharides from Morchella sextelata. J. Food Process. Preserv.46, e9. doi: 10.1111/jfpp.16024

  • CrossRef
  • Google Scholar

• 147

  LiS.LiJ.ZhangJ.WangW.WangX.JingH.et al. (2017a). The antioxidative, antiaging, and hepatoprotective effects of alkali-extractable polysaccharides by Agaricus bisporus. Evid. Based Complement. Alternat. Med.2017, e7298683. doi: 10.1155/2017/7298683

  • CrossRef
  • Google Scholar

• 148

  LiW. J.LiL.ZhenW. Y.WangL. F.PanM.LvJ. Q.et al. (2017b). Ganoderma atrum polysaccharide ameliorates ROS generation and apoptosis in spleen and thymus of immunosuppressed mice. Food Chem. Toxicol.99, 199–208. doi: 10.1016/j.fct.2016.11.033

  • CrossRef
  • Google Scholar

• 149

  LiY.ShengY.LuX.GuoX.XuG.HanX.et al. (2020b). Isolation and purification of acidic polysaccharides from Agaricus blazei Murill and evaluation of their lipid-lowering mechanism. Int. J. Biol. Macromol.157, 276–287. doi: 10.1016/j.ijbiomac.2020.04.190

  • CrossRef
  • Google Scholar

• 150

  LiL.SuZ.HeY.ZhongX.FuC.ZouL.et al. (2024b). Physicochemical characterization and anti-angiogenesis activity of polysaccharides from Amauroderma rugosum, a medicinal and edible mushroom. Int. J. Biol. Macromol.274, e133478. doi: 10.1016/j.ijbiomac.2024.133478

  • CrossRef
  • Google Scholar

• 151

  LiY.ThambiT.LeeD. S. (2018b). Co-delivery of drugs and genes using polymeric nanoparticles for synergistic cancer therapeutic effects. Adv. Healthc. Mater.7, 1. doi: 10.1002/adhm.201700886

  • CrossRef
  • Google Scholar

• 152

  LiW.WangJ.ChenZ.GaoX.ChenY.XueZ.et al. (2018a). Physicochemical properties of polysaccharides from Lentinus edodes under high pressure cooking treatment and its enhanced anticancer effects. Int. J. Biol. Macromol.115, 994–1001. doi: 10.1016/j.ijbiomac.2018.04.094

  • CrossRef
  • Google Scholar

• 153

  LiS.WangA.LiuL.TianG.XuF. (2019b). Effect of deproteinization methods on the antioxidant activity of polysaccharides extracted from Lentinus edodes stipe. J. Food Meas. Charact.13, 1382–1389. doi: 10.1007/s11694-019-00054-2

  • CrossRef
  • Google Scholar

• 154

  LiB.ZhongM.SunY.LiangQ.ShenL.QayumA.et al. (2024a). Recent advancements in the utilization of ultrasonic technology for the curing of processed meat products: A comprehensive review. Ultrason. Sonochem.103, e106796. doi: 10.1016/j.ultsonch.2024.106796

  • CrossRef
  • Google Scholar

• 155

  LiX.ZhuJ.WangT.SunJ.GuoT.ZhangL.et al. (2023). Antidiabetic activity of Armillaria mellea polysaccharides: Joint ultrasonic and enzyme assisted extraction. Ultrason. Sonochem.95, e106370. doi: 10.1016/j.ultsonch.2023.106370

  • CrossRef
  • Google Scholar

• 156

  LiaoT.ShenF.ZhuH.MuW.QianH.LiuY. (2024). Extracellular polysaccharides from Sporidiobolus pararoseus alleviates rheumatoid through ameliorating gut barrier function and gut microbiota. Int. J. Biol. Macromol.260, e129436. doi: 10.1016/j.ijbiomac.2024.129436

  • CrossRef
  • Google Scholar

• 157

  LinN.LiM.HanZ.LiuX.QuJ.WangX.et al. (2024). Polysaccharides from Auricularia cornea var. Li.: Structural alterations during in vitro digestion and its potent immunomodulatory properties on macrophages. Food Biosci.59, e104014. doi: 10.1016/j.fbio.2024.104014

  • CrossRef
  • Google Scholar

• 158

  LiuX.ChenS.LiuH.XieJ.HasanK. M. F.ZengQ.et al. (2023b). Structural properties and anti-inflammatory activity of purified polysaccharides from Hen-of-the-woods mushrooms (Grifola frondosa). Front. Nutr.10, e1078868. doi: 10.3389/fnut.2023.1078868

  • CrossRef
  • Google Scholar

• 159

  LiuJ. J.ChenS. K.WangX.HeW. W.SongX. X.HuangX. J.et al. (2024a). Changes of the physicochemical properties and structural characteristics of alkali-extracted polysaccharides from Agrocybe cylindracea across the growth process. J. Agric. Food Chem.72, 12810–12821. doi: 10.1021/acs.jafc.4c02218

  • CrossRef
  • Google Scholar

• 160

  LiuY.ChenS.ZhangJ.GaoM.LiL. (2023c). Purification of polysaccharide produced by the haploid yeast strain of Tremella sanGuinea and its antioxidant and prebiotic activities. Molecules28, e5391. doi: 10.3390/molecules28145391

  • CrossRef
  • Google Scholar

• 161

  LiuR. M.DaiR.LuoY.XiaoJ. H. (2019). Glucose-lowering and hypolipidemic activities of polysaccharides from Cordyceps taii in streptozotocin-induced diabetic mice. BMC Complement. Altern. Med.19, e230. doi: 10.1186/s12906-019-2646-x

  • CrossRef
  • Google Scholar

• 162

  LiuY.LiuX.CuiY.YuanW. (2022b). Ultrasound for microalgal cell disruption and product extraction: A review. Ultrason. Sonochem.87, e106054. doi: 10.1016/j.ultsonch.2022.106054

  • CrossRef
  • Google Scholar

• 163

  LiuX.LuoD.GuanJ.ChenJ.XuX. (2022a). Mushroom polysaccharides with potential in anti-diabetes: Biological mechanisms, extraction, and future perspectives: A review. Front. Nutr.9, e1087826. doi: 10.3389/fnut.2022.1087826

  • CrossRef
  • Google Scholar

• 164

  LiuX.WangQ.WangJ.GuoL.ChuY.MaC.et al. (2024b). Structural characterization, chain conformation and immunomodulatory activity of a heteropolysaccharide from Inonotus hispidus. Int. J. Biol. Macromol.260, e129187. doi: 10.1016/j.ijbiomac.2023.129187

  • CrossRef
  • Google Scholar

• 165

  LiuY.WangY.ZhangC.ZhouP.LiuY.AnT.et al. (2014). Core-shell nanoparticles based on pullulan and poly(β-amino) ester for hepatoma-targeted codelivery of gene and chemotherapy agent. ACS Appl. Mater. Interfaces6, 18712–18720. doi: 10.1021/am504203x

  • CrossRef
  • Google Scholar

• 166

  LiuJ.WuD.LengY.LiY.LiN. (2023a). Dietary supplementation with selenium polysaccharide from selenium-enriched Phellinus linteus improves antioxidant capacity, immunity and production performance of laying hens. J. Trace Elem. Med. Biol.77, e127140. doi: 10.1016/j.jtemb.2023.127140

  • CrossRef
  • Google Scholar

• 167

  LiuY.YangJ.GuoZ.LiQ.ZhangL.ZhaoL.et al. (2024c). Immunomodulatory effect of Cordyceps militaris polysaccharide on RAW 264.7 macrophages by regulating MAPK signaling pathways. Molecules29, e3408. doi: 10.3390/molecules29143408

  • CrossRef
  • Google Scholar

• 168

  LiuY.ZhouY.LiuM.WangQ.LiY. (2018). Extraction optimization, characterization, antioxidant and immunomodulatory activities of a novel polysaccharide from the wild mushroom Paxillus involutus. Int. J. Biol. Macromol.112, 326–332. doi: 10.1016/j.ijbiomac.2018.01.132

  • CrossRef
  • Google Scholar

• 169

  LoiraI.MorataA.PalomeroF.GonzálezC.Suárez-LepeJ. A. (2018). Schizosaccharomyces pombe: A promising biotechnology for modulating wine composition. Fermentation4, e70. doi: 10.3390/fermentation4030070

  • CrossRef
  • Google Scholar

• 170

  LongZ.XueY.NingZ.SunJ.LiJ.SuZ.et al. (2021). Production, characterization, and bioactivities of exopolysaccharides from the submerged culture of Ganoderma cantharelloideum M. H. Liu. Biotech.11, e145. doi: 10.1007/s13205-021-02696-w

  • CrossRef
  • Google Scholar

• 171

  López-LegardaX.Arboleda-EchavarríaC.Parra-SaldívarR.Rostro-AlanisM.AlzateJ. F.Villa-PulgarínJ. A.et al. (2020). Biotechnological production, characterization and in vitro antitumor activity of polysaccharides from a native strain of Lentinus crinitus. Int. J. Biol. Macromol.164, 3133–3144. doi: 10.1016/j.ijbiomac.2020.08.191

  • CrossRef
  • Google Scholar

• 172

  LorenzoJ. M.MunekataP. E.DominguezR.PateiroM.SaraivaJ. A.FrancoD.et al. (2018). “Main groups of microorganisms of relevance for food safety and stability,” in Innovative technologies for food preservation. Eds. BarbaF. J.Sant’AnaA. S.OrlienV. (Academic Press, Cambridge, Massachusetts, United States), 53–107. doi: 10.1016/B978-0-12-811031-7.00003-0

  • CrossRef
  • Google Scholar

• 173

  LuM. K.LeeM. H.ChaoC. H.HsuY. C. (2024a). Sodium sulfate addition increases the bioresource of biologically active sulfated polysaccharides from Antrodia cinnamomea. Int. J. Biol. Macromol.257, e128699. doi: 10.1016/j.ijbiomac.2023.128699

  • CrossRef
  • Google Scholar

• 174

  LuY.LiuD. (2024). Optimization of polysaccharide conditions and analysis of antioxidant capacity in the co-culture of Sanghuangporus vaninii and Pleurotus sapidus. PeerJ12, e17571. doi: 10.7717/peerj.17571

  • CrossRef
  • Google Scholar

• 175

  LuX.WangC.LiY.LiuP. (2022). Improved production and antioxidant activity of exopolysaccharides by submerged culture of Lentinula edodes by the addition of lignocellulose. J. Biosci. Bioeng.134, 162–166. doi: 10.1016/j.jbiosc.2022.05.003

  • CrossRef
  • Google Scholar

• 176

  LuX.WuS.AiH.WuR.ChengY.YunS.et al. (2024b). Sparassis latifolia polysaccharide alleviated lipid metabolism abnormalities in kidney of lead-exposed mice by regulating oxidative stress-mediated inflammation and autophagy based on multi-omics. Int. J. Biol. Macromol.278, e134662. doi: 10.1016/j.ijbiomac.2024.134662

  • CrossRef
  • Google Scholar

• 177

  LuoY.CaoN.HuangL.TangL.LiuX.ZhangW.et al. (2024). Structural characterization, and antioxidant, hypoglycemic and immunomodulatory activity of exopolysaccharide from Sanghuangporus sanghuang JM-1. Molecules29, e4564. doi: 10.3390/molecules29194564

  • CrossRef
  • Google Scholar

• 178

  LuoX.DuanY.YangW.ZhangH.LiC.ZhangJ. (2017). Structural elucidation and immunostimulatory activity of polysaccharide isolated by subcritical water extraction from Cordyceps militaris. Carbohydr. Polym.157, 794–802. doi: 10.1016/j.carbpol.2016.10.066

  • CrossRef
  • Google Scholar

• 179

  LuoQ.LiX.LiH.KongK.LiC.FangZ.et al. (2023). Effect of in vitro simulated digestion and fecal fermentation on Boletus auripes polysaccharide characteristics and intestinal flora. Int. J. Biol. Macromol.249, e126461. doi: 10.1016/j.ijbiomac.2023.126461

  • CrossRef
  • Google Scholar

• 180

  MaK. L.KeiN.YangF.LauwS.ChanP. L.ChenL.et al. (2023). In vitro fermentation characteristics of fungal polysaccharides derived from Wolfiporia cocos and their effect on human fecal microbiota. Foods12, e4014. doi: 10.3390/foods12214014

  • CrossRef
  • Google Scholar

• 181

  MaY.ZhengX.ChuZ.NanW.ZhaoY.BaiY.et al. (2024). Polysaccharides from Inonotus obliquus employing subcritical water extraction: Extraction optimization, physiochemical properties and bioactivities analysis. Ind. Crop Prod.222, e119638. doi: 10.1016/j.indcrop.2024.119638

  • CrossRef
  • Google Scholar

• 182

  MahendranS.SaravananS.VijayabaskarP.AnandapandianK. T. K.ShankarT. (2013). Antibacterial potential of microbial exopolysaccharide from Ganoderma lucidum and Lysinibacillus fusiformis. Int. J. Recent Sci. Res.4, 501–505.

  • Google Scholar

• 183

  MaiaL. S.de BomfimA. S. C.de OliveiraD. M.PinhatiF. R.da ConceiçãoM. O. T.BarudH. S.et al. (2023). Tuning of renewable sponge-like polyurethane physical-chemical and morphological properties using the pullulan as a reactive filler. J. Appl. Polym. Sci.140, e53619. doi: 10.1002/app.53619

  • CrossRef
  • Google Scholar

• 184

  MaingamC.KanchanarachW.ChutimanN.WanthongA.SrivilaiP.LoutchanwootP. (2024). Implications of ultrasonication-assisted extraction with response surface methodology on phytochemical compositions and antioxidant activity of polysaccharide extract from Phellinus rimosus (Berk.) Pilát cultivated mycelia in northeastern Thailand. Nat. Prod. J.14, 67–83. doi: 10.2174/0122103155293542240118063111

  • CrossRef
  • Google Scholar

• 185

  MansourA.DabaA.BaddourN.El-SaadaniM.AleemE. (2012). Schizophyllan inhibits the development of mammary and hepatic carcinomas induced by 7,12 dimethylbenz(α)anthracene and decreases cell proliferation: comparison with tamoxifen. J. Cancer Res. Clin. Oncol.138, 1579–1596. doi: 10.1007/s00432-012-1224-0

  • CrossRef
  • Google Scholar

• 186

  MaoG. H.RenY.FengW. W.LiQ.WuH. Y.JinD.et al. (2015). Antitumor and immunomodulatory activity of a water-soluble polysaccharide from Grifola frondosa. Carbohydr. Polym.134, 406–412. doi: 10.1016/j.carbpol.2015.08.020

  • CrossRef
  • Google Scholar

• 187

  MaoG. H.RenY.LiQ.WuH. Y.JinD.ZhaoT.et al. (2016). Anti-tumor and immunomodulatory activity of selenium (Se)-polysaccharide from Se-enriched Grifola frondosa. Int. J. Biol. Macromol.82, 607–613. doi: 10.1016/j.ijbiomac.2015.10.083

  • CrossRef
  • Google Scholar

• 188

  MazieroR.CavazzoniV.BononiV. L. R. (1999). Screening of basidiomycetes for the production of exopolysaccharide and biomass in submerged culture. Rev. Microbiol.30, 77–84. doi: 10.1590/s0001-37141999000100015

  • CrossRef
  • Google Scholar

• 189

  MedeirosS. D. V.CordeiroS. L.CavalcantiJ. E. C.MelchunaK. M.LimaA. M.daS.et al. (2012). Effects of purified Saccharomyces cerevisiae (1→3)-β-glucan on venous ulcer healing. Int. J. Mol. Sci.13, 8142–8158. doi: 10.3390/ijms13078142

  • CrossRef
  • Google Scholar

• 190

  MeiY.ZhuH.HuQ.LiuY.ZhaoS.PengN.et al. (2015). A novel polysaccharide from mycelia of cultured Phellinus linteus displays antitumor activity through apoptosis. Carbohydr. Polym.124, 90–97. doi: 10.1016/j.carbpol.2015.02.009

  • CrossRef
  • Google Scholar

• 191

  MengY.SuiX.PanX.ZhangX.SuiH.XuT.et al. (2023). Density-oriented deep eutectic solvent-based system for the selective separation of polysaccharides from Astragalus membranaceus var. Mongholicus under ultrasound-assisted conditions. Ultrason. Sonochem.98, e106522. doi: 10.1016/j.ultsonch.2023.106522

  • CrossRef
  • Google Scholar

• 192

  MfopaA.MediesseF. K.MvongoC.NkoubatchoundjwenS.LumA. A.SobngwiE.et al. (2021). Antidyslipidemic potential of water-soluble polysaccharides of Ganoderma applanatum in MACAPOS-2-induced obese rats. Evid. Based Complement. Alternat. Med.2021, e2452057. doi: 10.1155/2021/2452057

  • CrossRef
  • Google Scholar

• 193

  MiaoJ.RegensteinJ. M.QiuJ.ZhangJ.ZhangX.LiH.et al. (2020). Isolation, structural characterization and bioactivities of polysaccharides and its derivatives from Auricularia-A review. Int. J. Biol. Macromol.150, 102–113. doi: 10.1016/j.ijbiomac.2020.02.054

  • CrossRef
  • Google Scholar

• 194

  MilletteP. G.ChabotJ.SheppardD. C.Le MauffF. (2023). Identification and quantification of monosaccharides from fungal cell walls and exopolysaccharides by gas chromatography coupled to mass spectrometry. Curr. Protoc.3, e853. doi: 10.1002/cpz1.853

  • CrossRef
  • Google Scholar

• 195

  MinariM. C.RincãoV. P.SoaresS. A.RicardoN. M.NozawaC.LinharesR. E. (2011). Antiviral properties of polysaccharides from Agaricus brasiliensis in the replication of bovine herpesvirus 1. Acta Virol.55, 255–259. doi: 10.4149/av_2011_03_255

  • CrossRef
  • Google Scholar

• 196

  Miranda-NantesC. C. B. O.FonsecaE. A. I.ZaiaC. T. B. V.DekkerR. F. H.KhaperN.CastroI. A.et al. (2011). Hypoglycemic and hypocholesterolemic effects of botryosphaeran from Botryosphaeria rhodina MAMB-05 in diabetes-induced and hyperlipidemia conditions in rats. Mycobiology39, 187–193. doi: 10.5941/MYCO.2011.39.3.187

  • CrossRef
  • Google Scholar

• 197

  MohamedH. I.BasitA.AbdallahW. E. (2024). “Exopolysaccharides produced by fungi and their environmental applications,” in Fungal secondary metabolites. Eds. Abd-ElsalamK. A.MohamedH. I. (Elsevier), 219–240. doi: 10.1016/B978-0-323-95241-5.00028-9

  • CrossRef
  • Google Scholar

• 198

  MonroyY. M.RodriguesR. A. F.SartorattoA.CabralF. A. (2016). Influence of ethanol, water, and their mixtures as co-solvents of the supercritical carbon dioxide in the extraction of phenolics from purple corn cob (Zea mays L.). J. Supercrit. Fluids118, 11–18. doi: 10.1016/j.supflu.2016.07.019

  • CrossRef
  • Google Scholar

• 199

  MontoyaS.SanchezO. J.LevinL. (2013). Polysaccharide production by submerged and solid-state cultures from several medicinal higher basidiomycetes. Int. J. Med. Mushrooms15, 71–79. doi: 10.1615/intjmedmushr.v15.i1.80

  • CrossRef
  • Google Scholar

• 200

  Montoya BarretoS.Orrego AlzateC. E.LevinL. (2011). Modeling Grifola frondosa fungal growth during solid-state fermentation. Eng. Life Sci.11, 316–321. doi: 10.1002/elsc.201000087

  • CrossRef
  • Google Scholar

• 201

  MoradiZ.KalanpourN. (2019). Kefiran, a branched polysaccharide: Preparation, properties and applications: A review. Carbohydr. Polym.223, e115100. doi: 10.1016/j.carbpol.2019.115100

  • CrossRef
  • Google Scholar

• 202

  MoralesD.SmiderleF. R.VillalvaM.AbreuH.RicoC.SantoyoS.et al. (2019). Testing the effect of combining innovative extraction technologies on the biological activities of obtained β-glucan-enriched fractions from. Lentinula edodes. J. Funct. Foods60, e103446. doi: 10.1016/j.jff.2019.103446

  • CrossRef
  • Google Scholar

• 203

  MorrisH. J.LlauradóG.BeltránY.LebequeY.BermúdezR. C.GarcíaN.et al. (2016). “The use of mushrooms in the development of functional foods, drugs, and nutraceuticals: functional food properties and applications,” in Wild plants, mushrooms and nuts. Eds. FerreiraI. C. F. R.MoralesP.BarrosL. (John Wiley, Sons, Ltd), 123–157. doi: 10.1002/9781118944653.Ch5

  • CrossRef
  • Google Scholar

• 204

  MugnainiG.BoniniM.GentileL.PanzaO.Del NobileM. A.ConteA.et al. (2024). Effect of design and molecular interactions on the food preserving properties of alginate/pullulan edible films loaded with grape pomace extract. J. Food Eng.361, e111716. doi: 10.1016/j.jfoodeng.2023.111716

  • CrossRef
  • Google Scholar

• 205

  MukhopadhyaA.O’DohertyJ. V.SweeneyT. (2019). A combination of yeast beta-glucan and milk hydrolysate is a suitable alternative to zinc oxide in the race to alleviate post-weaning diarrhoea in piglets. Sci. Rep.9, e616. doi: 10.1038/s41598-018-37004-9

  • CrossRef
  • Google Scholar

• 206

  MurphyE. J.FehrenbachG. W.AbidinI. Z.BuckleyC.MontgomeryT.PogueR.et al. (2023a). Polysaccharides-naturally occurring immune modulators. Polymers15, e2373. doi: 10.3390/polym15102373

  • CrossRef
  • Google Scholar

• 207

  MurphyE. J.RezoagliE.CollinsC.SahaS. K.MajorI.MurrayP. (2023b). Sustainable production and pharmaceutical applications of β-glucan from microbial sources. Microbiol. Res.274, e127424. doi: 10.1016/j.micres.2023.127424

  • CrossRef
  • Google Scholar

• 208

  MuthusamyG.JoardarS. N.SamantaI. (2013). β-Glucan from edible mushroom (Pleurotus forida) enhances mucosal immunity in poultry. Adv. Anim. Vet. Sci.1, 116–119.

  • Google Scholar

• 209

  NegahbanZ.ShojaosadatiS. A.HamediS. (2021). A novel self-assembled micelles based on stearic acid modified schizophyllan for efficient delivery of paclitaxel. Colloids Surf. B Biointerfaces199, e111524. doi: 10.1016/j.colsurfb.2020.111524

  • CrossRef
  • Google Scholar

• 210

  NiZ.LiJ.QianX.YongY.WuM.WangY.et al. (2023). Phellinus igniarius polysaccharides ameliorate hyperglycemia by modulating the composition of the gut microbiota and their metabolites in diabetic mice. Molecules28, e7136. doi: 10.3390/molecules28207136

  • CrossRef
  • Google Scholar

• 211

  NiegoA. G. T.RapiorS.ThongklangN.RaspéO.HydeK. D.MortimerP. (2023). Reviewing the contributions of macrofungi to forest ecosystem processes and services. Fungal Biol. Rev.44, e100294. doi: 10.1016/j.fbr.2022.11.002

  • CrossRef
  • Google Scholar

• 212

  OğuzhanP.YangılarF. (2013). Pullulan: Production and usage in food industry. Afr. J. Food Sci. Technol.4, 57–63.

  • Google Scholar

• 213

  Osińska-JaroszukM.JaszekM.Mizerska-DudkaM.BłachowiczA.RejczakT. P.JanuszG.et al. (2014). Exopolysaccharide from Ganoderma applanatum as a promising bioactive compound with cytostatic and antibacterial properties. BioMed. Res. Int.2014, e743812. doi: 10.1155/2014/743812

  • CrossRef
  • Google Scholar

• 214

  Osińska-JaroszukM.SulejJ.JaszekM.Jaroszuk-ŚcisełJ. (2020). “Applications of fungal polysaccharides” in Encyclopedia of mycology. Eds. ZaragozaÓ.CasadevallA. (Elsevier, Oxford), 613–628. doi: 10.1016/B978-0-12-809633-8.21092-3

  • CrossRef
  • Google Scholar

• 215

  PalaciosI.García-LafuenteA.GuillamónE.VillaresA. (2012). Novel isolation of water-soluble polysaccharides from the fruiting bodies of Pleurotus ostreatus mushrooms. Carbohydr. Res.358, 72–77. doi: 10.1016/j.carres.2012.06.016

  • CrossRef
  • Google Scholar

• 216

  PanX.VeroniainaH.SuN.ShaK.JiangF.WuZ.et al. (2021). Applications and developments of gene therapy drug delivery systems for genetic diseases. Asian J. Pharm. Sci.16, 687–703. doi: 10.1016/j.ajps.2021.05.003

  • CrossRef
  • Google Scholar

• 217

  PanyaM.KaewraemruaenC.SaenwangP.PimboonP. (2024). Evaluation of prebiotic potential of crude polysaccharides extracted from wild Lentinus polychrous and Lentinus squarrosulus and their application for a formulation of a novel lyophilized synbiotic. Foods13, e287. doi: 10.3390/foods13020287

  • CrossRef
  • Google Scholar

• 218

  ParniakovO.LebovkaN. I.Van HeckeE.VorobievE. (2014). Pulsed electric field assisted pressure extraction and solvent extraction from mushroom (Agaricus bisporus). Food Bioprocess Technol.7, 174–183. doi: 10.1007/s11947-013-1059-y

  • CrossRef
  • Google Scholar

• 219

  PengB.LiuY.LinY.KraithongS.MoL.GaoZ.et al. (2023). A new exopolysaccharide of marine coral-associated Aspergillus pseudoglaucus SCAU265: Structural characterization and immunomodulatory activity. J. Fungi9, e1057. doi: 10.3390/jof9111057

  • CrossRef
  • Google Scholar

• 220

  Pérez-ChávezA. M.MayerL.AlbertóE. (2019). Mushroom cultivation and biogas production: A sustainable reuse of organic resources. Energy Sustain. Dev.50, 50–60. doi: 10.1016/j.esd.2019.03.002

  • CrossRef
  • Google Scholar

• 221

  Pham-HuyL. A.HeH.Pham-HuyC. (2008). Free radicals, antioxidants in disease and health. Int. J. Biomed. Sci.4, 89–96. doi: 10.59566/IJBS.2008.4089

  • CrossRef
  • Google Scholar

• 222

  PhaniendraA.JestadiD. B.PeriyasamyL. (2015). Free radicals: Properties, sources, targets, and their implication in various diseases. Indian J. Clin. Biochem.30, 11–26. doi: 10.1007/s12291-014-0446-0

  • CrossRef
  • Google Scholar

• 223

  PrajapatiV. D.JaniG. K.KhandaS. M. (2013). Pullulan: An exopolysaccharide and its various applications. Carbohydr. Polym.95, 540–549. doi: 10.1016/j.carbpol.2013.02.082

  • CrossRef
  • Google Scholar

• 224

  PrathumpaiW.RachtaweeP.KhajeeramS. (2015). Potential of fungal exopolysaccharide as novel source for prebiotic supplement to broiler chicken diet. Indian J. Anim. Sci.85, e12. doi: 10.56093/ijans.v85i12.54403

  • CrossRef
  • Google Scholar

• 225

  QinY.ZhangZ.SongT.LvG. (2017). Optimization of enzyme-assisted extraction of antitumor polysaccharides from Hericium erinaceus mycelia. Food Sci. Technol. Res.23, 31–39. doi: 10.3136/fstr.23.31

  • CrossRef
  • Google Scholar

• 226

  QiuJ.ZhengP.DaiW.ZhengZ.LinX.HuJ.et al. (2024). Steam explosion-assisted extraction of polysaccharides from Pleurotus eryngii and its influence on structural characteristics and antioxidant activity. Foods13, e1229. doi: 10.3390/foods13081229

  • CrossRef
  • Google Scholar

• 227

  RahmanH. U. U.AsgharW.KhalidN. (2021). “Scleroglucan and schizophyllan: microbial polysaccharides of functional importance,” in Polysaccharides of microbial origin. Eds. OliveiraJ. M.RadhouaniH.ReisR. L. (Springer, Cham), 1–28. doi: 10.1007/978-3-030-35734-4_16-1

  • CrossRef
  • Google Scholar

• 228

  RenY.BaiY.ZhangZ.CaiW.Del Rio FloresA. (2019). The preparation and structure analysis methods of natural polysaccharides of plants and fungi: A review of recent development. Molecules24, e3122. doi: 10.3390/molecules24173122

  • CrossRef
  • Google Scholar

• 229

  RenG.XuL.LuT.YinJ. (2018). Structural characterization and antiviral activity of lentinan from Lentinus edodes mycelia against infectious hematopoietic necrosis virus. Int. J. Biol. Macromol.115, 1202–1210. doi: 10.1016/j.ijbiomac.2018.04.132

  • CrossRef
  • Google Scholar

• 230

  RezaM. A. S.RasouliA.VahidiH.KobarfardF. (2024). Molecular identification of Shiitake (Lentinula edodes), analysis and production of beta-glucan using beech wood sawdust waste. Int. J. Biol. Macromol.280, e135539. doi: 10.1016/j.ijbiomac.2024.135539

  • CrossRef
  • Google Scholar

• 231

  RizkyanaA. D.HoT. C.RoyV. C.ParkJ. S.KiddaneA. T.KimG. D.et al. (2022). Sulfation and characterization of polysaccharides from Oyster mushroom (Pleurotus ostreatus) extracted using subcritical water. J. Supercrit. Fluids179, e105412. doi: 10.1016/j.supflu.2021.105412

  • CrossRef
  • Google Scholar

• 232

  RizviS. M. H. (2024). Nanotechnology applications in enhanced oil recovery (EOR). Int. J. Sci. Res. Manage.12, 135–143. doi: 10.18535/ijsrm/v12i06.c03

  • CrossRef
  • Google Scholar

• 233

  Rodríguez-SeoaneP.Torres PerezM. D.Fernández de AnaC.Sinde-StompelE.DomínguezH. (2022). Antiradical and functional properties of subcritical water extracts from edible mushrooms and from commercial counterparts. Int. J. Food Sci. Technol.57, 1420–1428. doi: 10.1111/ijfs.15383

  • CrossRef
  • Google Scholar

• 234

  RosyidaV. T.HayatiS. N.WiyonoT.DarsihC.RatihD. (2024). Effect of aqueous extraction method on total water-soluble polysaccharides content and phytochemical properties of white oyster mushroom (Pleurotus ostreatus). IOP Conf. Ser. Earth Environ. Sci.1377, e012064. doi: 10.1088/1755-1315/1377/1/012064

  • CrossRef
  • Google Scholar

• 235

  SacchelliB. A. L.Faccin-GalhardiL. C.ItoV. Y.LopesJ. L.DekkerR. F. H.Barbosa-DekkerA. M.et al. (2019). Botryosphaeran and sulfonated derivatives as novel antiviral agents for herpes simplex and dengue fever. Int. J. Biol. Macromol.138, 334–339. doi: 10.1016/j.ijbiomac.2019.07.084

  • CrossRef
  • Google Scholar

• 236

  SaenghaW.KariratT.PitisinN.PlangklangS.ButkhupL.UdomwongP.et al. (2023). Exploring the bioactive potential of Calostoma insigne, an endangered culinary puffball mushroom, from northeastern Thailand. Foods13, e113. doi: 10.3390/foods13010113

  • CrossRef
  • Google Scholar

• 237

  SaetangN.RattanapotT.ManmaiN.AmornlerdpisonD.RamarajR.UnpapromY. (2022). Effect of hot water extraction process on schizophyllan from split gill mushroom. Biomass Conv. Bioref.14, 1017–1026. doi: 10.1007/s13399-021-02286-z

  • CrossRef
  • Google Scholar

• 238

  SahibR. S.ShafiqS. A.ChechanR. A. (2024). Isolation, purification and structural characterization of lentinan from a newly discovered Iraqi strain of Lentinula edodes RSR. Glob. Sci. J. Biol.9, 119–127. doi: 10.5281/zenodo.10693647

  • CrossRef
  • Google Scholar

• 239

  SakdasriW.ArnutpongchaiP.PhonsavatS.BumrungthaichaichanE.SawangkeawR. (2022). Pressurized hot water extraction of crude polysaccharides, β-glucan, and phenolic compounds from dried gray oyster mushroom. LWT-Food Sci. Technol.168, e113895. doi: 10.1016/j.lwt.2022.113895

  • CrossRef
  • Google Scholar

• 240

  SamadlouieH. R.JahanbinK.JalaliP. (2020). Production, medium optimization, and structural characterization of an extracellular polysaccharide produced by Rhodotorula minuta ATCC 10658. Food Sci. Nutr.8, 4954–4964. doi: 10.1002/fsn3.1792

  • CrossRef
  • Google Scholar

• 241

  SánchezÓ.J.MontoyaS. (2020). Assessment of polysaccharide and biomass production from three white-rot fungi by solid-state fermentation using wood and agro-industrial residues: A kinetic approach. Forests11, e1055. doi: 10.3390/f11101055

  • CrossRef
  • Google Scholar

• 242

  SánchezÓ.J.MontoyaS.VargasL. M. (2015). “Polysaccharide production by submerged fermentation,” in Polysaccharides. Eds. RamawatK.MérillonJ. M. (Springer, Cham), 451–473. doi: 10.1007/978-3-319-16298-0_39

  • CrossRef
  • Google Scholar

• 243

  SangT.GuoC.GuoD.WuJ.WangY.WangY.et al. (2021). Suppression of obesity and inflammation by polysaccharide from sporoderm-broken spore of Ganoderma lucidum via gut microbiota regulation. Carbohydr. Polym.256, e117594. doi: 10.1016/j.carbpol.2020.117594

  • CrossRef
  • Google Scholar

• 244

  SangthongS.PintathongP.PongsuaP.JiraratA.ChaiwutP. (2022). Polysaccharides from Volvariella volvacea mushroom: extraction, biological activities and cosmetic efficacy. J. Fungi8, e572. doi: 10.3390/jof8060572

  • CrossRef
  • Google Scholar

• 245

  SaravanakumarK.ParkS.SathiyaseelanA.MariadossA. V. A.ParkS.KimS. J.et al. (2021). Isolation of polysaccharides from Trichoderma harzianum with antioxidant, anticancer, and enzyme inhibition properties. Antioxidants10, e1372. doi: 10.3390/antiox10091372

  • CrossRef
  • Google Scholar

• 246

  ScafatiV.TroiloF.PonzianiS.GiovannoniM.ScorticaA.PontiggiaD.et al. (2022). Characterization of two 1,3-β-glucan-modifying enzymes from Penicillium sumatraense reveals new insights into 1,3-β-glucan metabolism of fungal saprotrophs. Biotechnol. Biofuels Bioprod.15, e138. doi: 10.1186/s13068-022-02233-8

  • CrossRef
  • Google Scholar

• 247

  SchlaubitzS.DerkaouiS. M.MarosaL.MirauxS.RenardM.CatrosS.et al. (2014). Pullulan/dextran/nHA macroporous composite beads for bone repair in a femoral condyle defect in rats. PloS One9, e110251. doi: 10.1371/journal.pone.0110251

  • CrossRef
  • Google Scholar

• 248

  SermwittayawongD.PatninanK.PhothiphiphitS.BoonyarattanakalinS.SermwittayawongN.HutadilokTowatanaN. (2018). Purification, characterization, and biological activities of purified polysaccharides extracted from the gray oyster mushroom [Pleurotus sajor-caju (Fr.) Sing.]. J. Food Biochem.42, e12606. doi: 10.1111/jfbc.12606

  • CrossRef
  • Google Scholar

• 249

  ShaoZ.TianY.LiuS.ChuX.MaoW. (2023). Anti-diabetic activity of a novel exopolysaccharide produced by the mangrove endophytic fungus Penicillium janthinellum N29. Mar. Drugs21, e270. doi: 10.3390/md21050270

  • CrossRef
  • Google Scholar

• 250

  ShenL.PangS.ZhongM.SunY.QayumA.LiuY.et al. (2023). A comprehensive review of ultrasonic assisted extraction (UAE) for bioactive components: Principles, advantages, equipment, and combined technologies. Ultrason. Sonochem.101, e106646. doi: 10.1016/j.ultsonch.2023.106646

  • CrossRef
  • Google Scholar

• 251

  ShiM.YangY.GuanD.ZhangY.ZhangZ. (2012). Bioactivity of the crude polysaccharides from fermented soybean curd residue by Flammulina velutipes. Carbohydr. Polym.89, 1268–1276. doi: 10.1016/j.carbpol.2012.04.047

  • CrossRef
  • Google Scholar

• 252

  SiJ.MengG.WuY.MaH. F.CuiB. K.DaiY. C. (2018). Medium composition optimization, structural characterization, and antioxidant activity of exopolysaccharides from the medicinal mushroom Ganoderma lingzhi. Int. J. Biol. Macromol.124, 1186–1196. doi: 10.1016/j.ijbiomac.2018.11.274

  • CrossRef
  • Google Scholar

• 253

  SinghR. S.KaurN.KennedyJ. F. (2019). Pullulan production from agro-industrial waste and its applications in food industry: A review. Carbohydr. Polym.217, 46–57. doi: 10.1016/j.carbpol.2019.04.050

  • CrossRef
  • Google Scholar

• 254

  SinghR. S.KaurN.RanaV.KennedyJ. F. (2016). Recent insights on applications of pullulan in tissue engineering. Carbohydr. Polym.153, 455–462. doi: 10.1016/j.carbpol.2016.07.118

  • CrossRef
  • Google Scholar

• 255

  SinghS.SinghV.PatelS. (2024). Cosmeceuticals; The fusion of cosmetics and pharmaceuticals. J. Community Pharm. Pract.4, 16–27. doi: 10.55529/jcpp.42.16.27

  • CrossRef
  • Google Scholar

• 256

  SippingM. T. K.MediesseF. K.KenmogneL. V.KanemotoJ. E. N.NjamenD.BoudjekoT. (2022). Polysaccharide-rich fractions from Ganoderma resinaceum (ganodermataceae) as chemopreventive agents in N-diethylnitrosamine-induced hepatocellular carcinoma in wistar rats. Evid. Based Complement. Altern. Med.2022, e8198859. doi: 10.1155/2022/8198859

  • CrossRef
  • Google Scholar

• 257

  Skalicka-WoźniakK.SzypowskiJ.ŁośR.SiwulskiM.SobieralskiK.GłowniakK.et al. (2012). Evaluation of polysaccharides content in fruit bodies and their antimicrobial activity of four Ganoderma lucidum (W Curt.: Fr.) P. Karst. strains cultivated on different wood type substrates. Acta Soc Bot. Pol.81, 17–21. doi: 10.5586/asbp.2012.001

  • CrossRef
  • Google Scholar

• 258

  SmiderleF. R.OlsenL. M.RuthesA. C.CzelusniakP. A.Santana-FilhoA. P.SassakiG. L.et al. (2012). Exopolysaccharides, proteins and lipids in Pleurotus pulmonarius submerged culture using different carbon sources. Carbohydr. Polym87, 368–376. doi: 10.1016/j.carbpol.2011.07.063

  • CrossRef
  • Google Scholar

• 259

  SongX.LiuZ.ZhangJ.YanQ.RenZ.ZhangC.et al. (2018). Anti-inflammatory and hepatoprotective effects of exopolysaccharides isolated from Pleurotus geesteranus on alcohol-induced liver injury. Sci. Rep.8, e10493. doi: 10.1038/s41598-018-28785-0

  • CrossRef
  • Google Scholar

• 260

  StoicaR. M.MoscoviciM.LakatosE. S.CiocaL. I. (2023). Exopolysaccharides of fungal origin: properties and pharmaceutical applications. Processes11, e335. doi: 10.3390/pr11020335

  • CrossRef
  • Google Scholar

• 261

  SuJ.SuL.LiD.ShuaiO.ZhangY.LiangH.et al. (2018). Antitumor activity of extract from the sporoderm-breaking spore of Ganoderma lucidum: Restoration on exhausted cytotoxic T cell with gut microbiota remodeling. Front. Immunol.9, e1765. doi: 10.3389/fimmu.2018.01765

  • CrossRef
  • Google Scholar

• 262

  SuL.XinC.YangJ.DongL.MeiH.DaiX.et al. (2022). A polysaccharide from Inonotus obliquus ameliorates intestinal barrier dysfunction in mice with type 2 diabetes mellitus. Int. J. Biol. Macromol.214, 312–323. doi: 10.1016/j.ijbiomac.2022.06.071

  • CrossRef
  • Google Scholar

• 263

  SunY.HeH.WangQ.YangX.JiangS.WangD. (2022b). A review of development and utilization for edible fungal polysaccharides: extraction, chemical characteristics, and bioactivities. Polymers14, e4454. doi: 10.3390/polym14204454

  • CrossRef
  • Google Scholar

• 264

  SunH.ShuF.GuanY.KongF.LiuS.LiuY.et al. (2023). Study of anti-fatigue activity of polysaccharide from fruiting bodies of Armillaria gallica. Int. J. Biol. Macromol.241, e124611. doi: 10.1016/j.ijbiomac.2023.124611

  • CrossRef
  • Google Scholar

• 265

  SunW.ZhangY.JiaL. (2022a). Polysaccharides from Agrocybe cylindracea residue alleviate type 2-diabetes-induced liver and colon injuries by P38 MAPK signaling pathway. Food Biosci.47, e101690. doi: 10.1016/j.fbio.2022.101690

  • CrossRef
  • Google Scholar

• 266

  SungY. K.KimS. W. (2019). Recent advances in the development of gene delivery systems. Biomater. Res.23, e8. doi: 10.1186/s40824-019-0156-z

  • CrossRef
  • Google Scholar

• 267

  SuraiyaS.JangW. J.HaqM.KongI. S. (2024). Isolation and characterization of β-glucan containing polysaccharides from Monascus spp. using Saccharina japonica as submerged fermented substrate. Polysaccharides5, 435–449. doi: 10.3390/polysaccharides5030027

  • CrossRef
  • Google Scholar

• 268

  TabibzadehF.AlvandiH.Hatamian-ZarmiA.KalitukhaL.AghajaniH.Ebrahimi-HosseinzadehB. (2022). Antioxidant activity and cytotoxicity of exopolysaccharide from mushroom Hericium coralloides in submerged fermentation. Biomass Convers. Biorefin.14, 26953–26963. doi: 10.1007/s13399-022-03386-0

  • CrossRef
  • Google Scholar

• 269

  TagneR. F. T.Cruz-SantosM. M.AntunesF. A. F.ShibukawaV. P.MianoS. B.KenfackJ. A. A.et al. (2024). Pullulan production from sugarcane bagasse hemicellulosic hydrolysate by Aureobasidium pullulans ATCC 42023 in bubble column reactor. Fermentation10, e322. doi: 10.3390/fermentation10060322

  • CrossRef
  • Google Scholar

• 270

  TangB.MaJ.LiuL.XuJ.ZhangH.LiK.et al. (2025). Preparation of pullulan-shellac edible films with improved water-resistance and UV barrier properties for Chinese cherries preservation. J. Future Foods5, 107–118. doi: 10.1016/j.jfutfo.2024.01.010

  • CrossRef
  • Google Scholar

• 271

  TangY. J.ZhangW.LiuR. S.ZhuL. W.ZhongJ. J. (2011). Scale-up study on the fed-batch fermentation of Ganoderma lucidum for the hyperproduction of ganoderic acid and Ganoderma polysaccharides. Process Biochem.46, 404–408. doi: 10.1016/j.procbio.2010.08.013

  • CrossRef
  • Google Scholar

• 272

  TepsongkrohB.ThaihuttakijC.SupawongS.JangchudK. (2023). Impact of high pressure pre-treatment and hot water extraction on chemical properties of crude polysaccharide extract obtained from mushroom (Volvariella volvacea). Food Chem.: X19, e100864. doi: 10.1016/j.fochx.2023.100864

  • CrossRef
  • Google Scholar

• 273

  ThakurA.SharmaS.NamanS.BaldiA. (2023). Pullulan based polymeric novel drug delivery systems: a review on current state of art and prospects. J. Drug Deliv. Sci. Technol.90, e105117. doi: 10.1016/j.jddst.2023.105117

  • CrossRef
  • Google Scholar

• 274

  ThangavelP.VilvanathanS. P.KuttalamI.LonchinS. (2020). Topical administration of pullulan gel accelerates skin tissue regeneration by enhancing collagen synthesis and wound contraction in rats. Int. J. Biol. Macromol.149, 395–403. doi: 10.1016/j.ijbiomac.2020.01.187

  • CrossRef
  • Google Scholar

• 275

  TianB.JiangY.LiuR.HamedY. S.RayanA. M.XuS.et al. (2024). Positive effects of extracellular polysaccharides from Paecilomyces hepiali on immune-enhancing properties by regulating gut microbiota in cyclophosphamide-induced mice. Int. J. Biol. Macromol.274, e133390. doi: 10.1016/j.ijbiomac.2024.133390

  • CrossRef
  • Google Scholar

• 276

  TianR.WuL. L.LiH. F.LiangZ. Q.LiP. H.WangY.et al. (2023). Purification and structure characterization of the crude polysaccharide from the fruiting bodies of Butyriboletus pseudospeciosus and its modulation effects on gut microbiota. Molecules28, e2679. doi: 10.3390/molecules28062679

  • CrossRef
  • Google Scholar

• 277

  TianY.ZengH.XuZ.ZhengB.LinY.GanC.et al. (2012). Ultrasonic-assisted extraction and antioxidant activity of polysaccharides recovered from white button mushroom (Agaricus bisporus). Carbohydr. Polym.88, 522–529. doi: 10.1016/j.carbpol.2011.12.042

  • CrossRef
  • Google Scholar

• 278

  TufailM.JiangC. H.LiN. (2024). Altered metabolism in cancer: insights into energy pathways and therapeutic targets. Mol. Cancer23, e203. doi: 10.1186/s12943-024-02119-3

  • CrossRef
  • Google Scholar

• 279

  UdchumpisaiW.BangyeekhunE. (2020). Purification, structural characterization, and biological activity of polysaccharides from Lentinus velutinus. Mycobiol.48, 51–57. doi: 10.1080/12298093.2019.1693482

  • CrossRef
  • Google Scholar

• 280

  UrbancikovaI.HudackovaD.MajtanJ.RennerovaZ.BanovcinP.JesenakM. (2020). Efficacy of pleuran (β-glucan from Pleurotus ostreatus) in the management of herpes simplex virus type 1 infection. Evid. Based Complement. Alternat. Med.2020, e8562309. doi: 10.1155/2020/8562309

  • CrossRef
  • Google Scholar

• 281

  UtamaG. L.DioC.SulistiyoJ.Yee ChyeF.LembongE.CahyanaY.et al. (2021). Evaluating comparative β-glucan production aptitude of Saccharomyces cerevisiae, Aspergillus oryzae, Xanthomonas campestris, and Bacillus natto. Saudi J. Biol. Sci.28, 6765–6773. doi: 10.1016/j.sjbs.2021.07.051

  • CrossRef
  • Google Scholar

• 282

  UthanE. T.SenturkH.UyanogluM.YamaçM. (2021). First report on the in vivo prebiotic, biochemical, and histological effects of crude polysaccharide fraction of golden chantharelle mushroom, Cantharellus cibarius (Agaricomycetes). Int. J. Med. Mushrooms23, 67–77. doi: 10.1615/IntJMedMushrooms.2021038233

  • CrossRef
  • Google Scholar

• 283

  ValdezA. L.DelgadoO. D.FariñaJ. I. (2021). Cost-effective optimized scleroglucan production by Sclerotium rolfsii ATCC 201126 at bioreactor scale. A quantity-quality assessment. Carbohydr. Polym.260, e117505. doi: 10.1016/j.carbpol.2020.117505

  • CrossRef
  • Google Scholar

• 284

  ValverdeM. E.Hernández-PérezT.Paredes-LópezO. (2015). Edible mushrooms: Improving human health and promoting quality life. Int. J. Microbiol.2015, e376387. doi: 10.1155/2015/376387

  • CrossRef
  • Google Scholar

• 285

  VaninA. P.VisentinE. Z.FontanaR. C.di Medeiros LealM. C. B.de Avila E SilvaS.StokkeB. T.et al. (2023). β-(1→3)(1→6)glucan from schizophyllum commune 227E.32: high yield production via glucose/xylose co-metabolization. Carbohydr. Polym. 320, e121176. doi: 10.1016/j.carbpol.2023.121176

  • CrossRef
  • Google Scholar

• 286

  VetterJ. (2023). The mushroom glucans: molecules of high biological and medicinal importance. Foods12, e1009. doi: 10.3390/foods12051009

  • CrossRef
  • Google Scholar

• 287

  ViñartaS. C.YossenM. M.VegaJ. R.FigueroaL. I. C.FariñaJ. I. (2013). Scleroglucan compatibility with thickeners, alcohols and polyalcohols and downstream processing implications. Carbohydr. Polym.92, 1107–1115. doi: 10.1016/j.carbpol.2012.10.065

  • CrossRef
  • Google Scholar

• 288

  VisanA. I.CristescuR. (2023). Polysaccharide-based coatings as drug delivery systems. Pharmaceutics15, e2227. doi: 10.3390/pharmaceutics15092227

  • CrossRef
  • Google Scholar

• 289

  VlasenkoV. A.IlyichevaT. N.SvyatchenkoS. V.AsbaganovS. V.ZmitrovichI. V.VlasenkoA. V. (2020). Antiviral activity of total polysaccharide fraction of water and ethanol extracts of Pleurotus pulmonarius against the influenza A virus. Curr. Res. Environ. Appl. Mycol. J. Fungal. Biol.10, 224–235. doi: 10.5943/cream/10/1/22

  • CrossRef
  • Google Scholar

• 290

  WangY.AzharS.LindströmM. E.HenrikssonG. (2015c). Stabilization of polysaccharides during alkaline pre-treatment of wood combined with enzyme-supported extractions in a biorefinery. J. Wood Chem. Technol.35, 91–101. doi: 10.1080/02773813.2013.875041

  • CrossRef
  • Google Scholar

• 291

  WangJ.DouB.BaoY. (2014a). Efficient targeted pDNA/siRNA delivery with folate-low-molecular-weight polyethyleneimine-modified pullulan as non-viral carrier. Mat. Sci. Eng. C Mater. Biol. Appl.34, 98–109. doi: 10.1016/j.msec.2013.08.035

  • CrossRef
  • Google Scholar

• 292

  WangM.GaoY.XuD.GaoQ. (2015b). A polysaccharide from cultured mycelium of Hericium erinaceus and its anti-chronic atrophic gastritis activity. Int. J. Biol. Macromol.81, 656–661. doi: 10.1016/j.ijbiomac.2015.08.043

  • CrossRef
  • Google Scholar

• 293

  WangT.HanJ.DaiH.SunJ.RenJ.WangW.et al. (2022b). Polysaccharides from Lyophyllum decastes reduce obesity by altering gut microbiota and increasing energy expenditure. Carbohyd. Polym.295, e119862. doi: 10.1016/j.carbpol.2022.119862

  • CrossRef
  • Google Scholar

• 294

  WangK.HuaiS.TanZ.NgeaG. L. N.GodanaE. A.ShiJ.et al. (2023a). A first expression, purification and characterization of endo-β-1,3-glucanase from Penicillium expansum. J. Fungi9, e961. doi: 10.3390/jof9100961

  • CrossRef
  • Google Scholar

• 295

  WangY.JinH.YuJ.QuC.WangQ.YangS.et al. (2020b). Quality control and immunological activity of lentinan samples produced in China. Int. J. Biol. Macromol.159, 129–136. doi: 10.1016/j.ijbiomac.2020.05.050

  • CrossRef
  • Google Scholar

• 296

  WangJ.LiJ.LiJ.LiJ.LiuS.GaoW. (2017a). LSP1, a responsive protein from Meyerozyma guilliermondii, elicits defence response and improves glycyrrhizic acid biosynthesis in Glycyrrhiza uralensis Fisch adventitious roots. J. Cell. Physiol.232, 3510–3519. doi: 10.1002/jcp.25811

  • CrossRef
  • Google Scholar

• 297

  WangL.LianJ.ZhengQ.WangL.WangY.YangD. (2022a). Composition analysis and prebiotics properties of polysaccharides extracted from Lepista sordida submerged cultivation mycelium. Front. Microbiol.13, e1077322. doi: 10.3389/fmicb.2022.1077322

  • CrossRef
  • Google Scholar

• 298

  WangQ. Q.LinJ.ZhouQ. Z.PengJ.ZhangQ.WangJ. H. (2023b). Hyper-production of pullulan by a novel fungus of Aureobasidium melanogenum ZH27 through batch fermentation. Int. J. Mol. Sci.25, e319. doi: 10.3390/ijms25010319

  • CrossRef
  • Google Scholar

• 299

  WangW.LiuH.ZhangY.FengY.YuanF.SongX.et al. (2019b). Antihyperlipidemic and hepatoprotective properties of alkali- and enzyme-extractable polysaccharides by Dictyophora indusiata. Sci. Rep.9, e14266. doi: 10.1038/s41598-019-50717-9

  • CrossRef
  • Google Scholar

• 300

  WangW.TanJ.NimaL.SangY.CaiX.XueH. (2022c). Polysaccharides from fungi: A review on their extraction, purification, structural features, and biological activities. Food Chem.: X15, e100414. doi: 10.1016/j.fochx.2022.100414

  • CrossRef
  • Google Scholar

• 301

  WangQ.WangF.XuZ.DingZ. (2017b). Bioactive mushroom polysaccharides: A review on monosaccharide composition, biosynthesis and regulation. Molecules22, e955. doi: 10.3390/molecules22060955

  • CrossRef
  • Google Scholar

• 302

  WangC. C.WuJ. Y.ChangC. Y.YuS. T.LiuY. C. (2019a). Enhanced exopolysaccharide production by Cordyceps militaris using repeated batch cultivation. J. Biosci. Bioeng.127, 499–505. doi: 10.1016/j.jbiosc.2018.09.006

  • CrossRef
  • Google Scholar

• 303

  WangJ.WuC.ChenY.ChenC.HuS.ChangS. (2014b). Antihyperglycemic activity of exopolysaccharide produced by mushroom Pleurotus ferulae with submerged liquid culture on streptozotocin-induced diabetic rats. J. Food Nutr. Res.2, 419–424. doi: 10.12691/jfnr-2-7-15

  • CrossRef
  • Google Scholar

• 304

  WangL.XuN.ZhangJ.ZhaoH.LinL.JiaS.et al. (2015a). Antihyperlipidemic and hepatoprotective activities of residue polysaccharide from Cordyceps militaris SU-12. Carbohydr. Polym.131, 355–362. doi: 10.1016/j.carbpol.2015.06.016

  • CrossRef
  • Google Scholar

• 305

  WangY. X.YinJ. Y.HuangX. J.NieS. P. (2020a). Structural characteristics and rheological properties of high viscous glucan from fruit body of Dictyophora rubrovolvata. Food Hydrocoll.101, e105514. doi: 10.1016/j.foodhyd.2019.105514

  • CrossRef
  • Google Scholar

• 306

  WangW. H.ZhangJ. S.FengT.DengJ.LinC. C.FanH.et al. (2018). Structural elucidation of a polysaccharide from Flammulina velutipes and its immunomodulation activities on mouse B lymphocytes. Sci. Rep.8, e3120. doi: 10.1038/s41598-018-21375-0

  • CrossRef
  • Google Scholar

• 307

  WangX.ZhangJ.ZhangK.GuoZ.XuG.HuangL.et al. (2024). Ultrasound-assisted enzyme extraction, physicochemical properties and antioxidant activity of polysaccharides from Cordyceps militaris solid medium. Molecules29, e4560. doi: 10.3390/molecules29194560

  • CrossRef
  • Google Scholar

• 308

  Wan-MohtarW. A. A. Q. I.YoungL.AbbottG. M.ClementsC.HarveyL. M.McNeilB. (2016). Antimicrobial properties and cytotoxicity of sulfated (1,3)-β-D-glucan from the mycelium of the mushroom Ganoderma lucidum. J. Microbiol. Biotechnol.26, 999–1010. doi: 10.4014/jmb.1510.10018

  • CrossRef
  • Google Scholar

• 309

  WeiS.WangL.ChenX.WangY.TongL.HanQ.et al. (2024). Anti-inflammatory activity of Boletus aereus polysaccharides: Involvement of digestion and gut microbiota fermentation. Food Chem.: X21, e101052. doi: 10.1016/j.fochx.2023.101052

  • CrossRef
  • Google Scholar

• 310

  WengB. B. C.LinY. C.HuC. W.KaoM. Y.WangS. H.LoD. Y.et al. (2011). Toxicological and immunomodulatory assessments of botryosphaeran (β-glucan) produced by Botryosphaeria rhodina RCYU 30101. Food Chem. Toxicol.49, 910–916. doi: 10.1016/j.fct.2010.10.036

  • CrossRef
  • Google Scholar

• 311

  WoodI. P.EllistonA.RydenP.BancroftI.RobertsI. N.WaldronK. W. (2012). Rapid quantification of reducing sugars in biomass hydrolysates: Improving the speed and precision of the dinitrosalicylic acid assay. Biomass Bioenergy44, 117–121. doi: 10.1016/j.biombioe.2012.05.003

  • CrossRef
  • Google Scholar

• 312

  WoukJ.CelestinoG. G.RodriguesB. C. D.MalfattiC. R. M.CunhaM. A. A.OrsatoA.et al. (2022). Sulfonated (1→6)-β-d-glucan (lasiodiplodan): A promising candidate against the acyclovir-resistant herpes simplex virus type 1 (HSV-1) strain. Biomacromolecules23, 4041–4052. doi: 10.1021/acs.biomac.2c00156

  • CrossRef
  • Google Scholar

• 313

  WoźniakB.ChudzińskaJ.SzczyglewskaP.NowakI.Feliczak-GuzikA. (2023). Optimization of the composition of a cosmetic formulation containing Tremella fuciformis extract (fungi). Cosmetics10, e82. doi: 10.3390/cosmetics10030082

  • CrossRef
  • Google Scholar

• 314

  WuJ. Y.ChenH. B.ChenM. J.KanS. C.ShiehC. J.LiuY. C. (2013). Quantitative analysis of LED effects on edible mushroom Pleurotus eryngii in solid and submerged cultures: LED effect on cultivating P. eryngii. J. Chem. Technol. Biotechnol.88, 1841–1846. doi: 10.1002/jctb.4038

  • CrossRef
  • Google Scholar

• 315

  WuY.JiangH.ZhuE.LiJ.WangQ.ZhouW.et al. (2018). Hericium erinaceus polysaccharide facilitates restoration of injured intestinal mucosal immunity in Muscovy duck reovirus-infected Muscovy ducklings. Int. J. Biol. Macromol.107, 1151–1161. doi: 10.1016/j.ijbiomac.2017.09.092

  • CrossRef
  • Google Scholar

• 316

  WuK.LiY.LinY.XuB.YangJ.MoL.et al. (2023). Structural characterization and immunomodulatory activity of an exopolysaccharide from marine-derived Aspergillus versicolor SCAU141. Int. J. Biol. Macromol.227, 329–339. doi: 10.1016/j.ijbiomac.2022.12.127

  • CrossRef
  • Google Scholar

• 317

  WuD.TangC.LiuY.LiQ.WangW.ZhouS.et al. (2019). Structural elucidation and immunomodulatory activity of a β-D-glucan prepared by freeze-thawing from Hericium erinaceus. Carbohydr. Polym.222, e114996. doi: 10.1016/j.carbpol.2019.114996

  • CrossRef
  • Google Scholar

• 318

  WuR. T.WangL. F.YaoY. F.SangT.WuQ. L.FuW. W.et al. (2022a). Activity fingerprinting of polysaccharides on oral, gut, pancreas and lung microbiota in diabetic rats. BioMed. Pharmacother.155, e113681. doi: 10.1016/j.biopha.2022.113681

  • CrossRef
  • Google Scholar

• 319

  WuS.YangX.YangX. (2022b). Methotrexate and 10-hydroxycamptothecine loaded pullulan nanoparticles with the targeting property for efficient cancer therapy. Mater. Technol.37, 2777–2784. doi: 10.1080/10667857.2022.2075079

  • CrossRef
  • Google Scholar

• 320

  WuF.ZhouC.ZhouD.OuS.HuangH. (2017). Structural characterization of a novel polysaccharide fraction from Hericium erinaceus and its signaling pathways involved in macrophage immunomodulatory activity. J. Funct. Foods37, 574–585. doi: 10.1016/j.jff.2017.08.030

  • CrossRef
  • Google Scholar

• 321

  XiaS.ZhangL.DavletshinA.LiZ.YouJ.TanS. (2020). Application of polysaccharide biopolymer in petroleum recovery. Polymers12, e1860. doi: 10.3390/polym12091860

  • CrossRef
  • Google Scholar

• 322

  XiangH.Sun-WaterhouseD.CuiC. (2021). Hypoglycemic polysaccharides from Auricularia auricula and Auricularia polytricha inhibit oxidative stress, NF-κB signaling and proinflammatory cytokine production in streptozotocin-induced diabetic mice. Food Sci. Hum. Wellness10, 87–93. doi: 10.1016/j.fshw.2020.06.001

  • CrossRef
  • Google Scholar

• 323

  XuJ.LiuZ.ZhangS.XiangJ.LanH.BaoY. (2024). Anti-hepatoma immunotherapy of Pholiota adiposa polysaccharide-coated selenium nanoparticles by reversing M2-like tumor-associated macrophage polarization. Int. J. Biol. Macromol.277, e133667. doi: 10.1016/j.ijbiomac.2024.133667

  • CrossRef
  • Google Scholar

• 324

  XuM.QuY.LiH.TangS.ChenC.WangY.et al. (2023). Improved extraction yield, water solubility, and antioxidant activity of lentinan from Lentinula edodes via Bacillus subtilis natto fermentation. Fermentation9, e333. doi: 10.3390/fermentation9040333

  • CrossRef
  • Google Scholar

• 325

  XuN.SunY. H.GuoX. L.LiuC.MaoQ.HouJ. M. (2018). Optimization of ultrasonic-microwave synergistic extraction of polysaccharides from Morchella conica. J. Food Process. Preserv.42, e13423. doi: 10.1111/jfpp.13423

  • CrossRef
  • Google Scholar

• 326

  XuL.WangF.ZhangZ.TerryN. (2019). Optimization of polysaccharide production from Cordyceps militaris by solid-state fermentation on rice and its antioxidant activities. Foods8, e590. doi: 10.3390/foods8110590

  • CrossRef
  • Google Scholar

• 327

  XuD.WangH.ZhengW.GaoY.WangM.ZhangY.et al. (2016). Charaterization and immunomodulatory activities of polysaccharide isolated from Pleurotus eryngii. Int. J. Biol. Macromol.92, 30–36. doi: 10.1016/j.ijbiomac.2016.07.016

  • CrossRef
  • Google Scholar

• 328

  YabalakE.AminzaiM. T.GizirA. M.YangY. (2024). A review: Subcritical water extraction of organic pollutants from environmental matrices. Molecules29, e258. doi: 10.3390/molecules29010258

  • CrossRef
  • Google Scholar

• 329

  YafettoL. (2022). Application of solid-state fermentation by microbial biotechnology for bioprocessing of agro-industrial wastes from 1970 to 2020: A review and bibliometric analysis. Heliyon8, e09173. doi: 10.1016/j.heliyon.2022.e09173

  • CrossRef
  • Google Scholar

• 330

  YanJ. K.WangW. Q.MaH. L.WuJ. Y. (2012). Sulfation and enhanced antioxidant capacity of an exopolysaccharide produced by the medicinal fungus Cordyceps sinensis. Molecules18, 167–177. doi: 10.3390/molecules18010167

  • CrossRef
  • Google Scholar

• 331

  YangQ.ChangS. L.TianY. M.LiW.RenJ. L. (2024b). Glucan polysaccharides isolated from Lactarius hatsudake Tanaka mushroom: Structural characterization and in vitro bioactivities. Carbohydr. Polym.337, e122171. doi: 10.1016/j.carbpol.2024.122171

  • CrossRef
  • Google Scholar

• 332

  YangX.GaoY.ReyimuM.ZhangG.WangC.YangD.et al. (2024c). Structural analysis of Pleurotus ferulae polysaccharide and its effects on plant fungal disease and plant growth. Int. J. Biol. Macromol.282, e137396. doi: 10.1016/j.ijbiomac.2024.137396

  • CrossRef
  • Google Scholar

• 333

  YangM.HuD.CuiZ.LiH.ManC.JiangY. (2021a). Lipid-lowering effects of Inonotus obliquus polysaccharide in vivo and in vitro. Foods10, e3085. doi: 10.3390/foods10123085

  • CrossRef
  • Google Scholar

• 334

  YangX.LinP.WangJ.LiuN.YinF.ShenN.et al. (2021b). Purification, characterization and anti-atherosclerotic effects of the polysaccharides from the fruiting body of Cordyceps militaris. Int. J. Biol. Macromol181, 890–904. doi: 10.1016/j.ijbiomac.2021.04.083

  • CrossRef
  • Google Scholar

• 335

  YangJ.LiuJ.KuangW.LinY.ZhongS.KraithongS.et al. (2024a). Structural characterization and ferroptosis-related immunomodulatory of a novel exopolysaccharide isolated from marine fungus Aspergillus medius. Int. J. Biol. Macromol.265, e130703. doi: 10.1016/j.ijbiomac.2024.130703

  • CrossRef
  • Google Scholar

• 336

  YangM.RenW.LiG.YangP.ChenR.HeH. (2022a). The effect of structure and preparation method on the bioactivity of polysaccharides from plants and fungi. Food Funct.13, 12541–12560. doi: 10.1039/d2fo02029g

  • CrossRef
  • Google Scholar

• 337

  YangX.SunS.ChenQ.ZhangZ.WangJ.LiuY.et al. (2022b). A polysaccharide of Ganoderma lucidum enhances antifungal activity of chemical fungicides against soil-borne diseases of wheat and maize by induced resistance. Agriculture12, e55. doi: 10.3390/agriculture12010055

  • CrossRef
  • Google Scholar

• 338

  YangX. M.WangS. Q.ChenL. S.ZhuZ. Y. (2023). Isolation and structural characterization of exopolysaccharide from the Cordyceps cicadae and the immunomodulatory activity on RAW264.7 cells. Biotechnol. Appl. Biochem.70, 1925–1940. doi: 10.1002/bab.2500

  • CrossRef
  • Google Scholar

• 339

  YangS.YanJ.YangL.MengY.WangN.HeC.et al. (2019). Alkali-soluble polysaccharides from mushroom fruiting bodies improve insulin resistance. Int. J. Biol. Macromol.126, 466–474. doi: 10.1016/j.ijbiomac.2018.12.251

  • CrossRef
  • Google Scholar

• 340

  YangW.ZhangH.JiM.PeiF.WangY. (2016). Antitumor effect of a polysaccharide isolated from Phellinus pullus as an immunostimulant. Biomed. Rep.4, 361–364. doi: 10.3892/br.2016.587

  • CrossRef
  • Google Scholar

• 341

  YangY.ZhangX.ZhangJ.WangT.LiuS.MaH.et al. (2024d). Green ultrasonic-assisted enzymatic extraction of polysaccharides from Flammulina velutipes residues by response surface methodology. Sustain. Chem. Pharm.41, e101690. doi: 10.1016/j.scp.2024.101690

  • CrossRef
  • Google Scholar

• 342

  YaoH. Y. Y.WangJ. Q.YinJ. Y.NieS. P.XieM. Y. (2021). A review of NMR analysis in polysaccharide structure and conformation: progress, challenge and perspective. Food Res. Int.143, e110290. doi: 10.1016/j.foodres.2021.110290

  • CrossRef
  • Google Scholar

• 343

  YatmazE.Turhanİ. (2012). Pullulan production by fermentation and usage in food industry. Food37, 95–102.

  • Google Scholar

• 344

  YeJ.WangX.WangK.DengY.YangY.AliR.et al. (2020). A novel polysaccharide isolated from Flammulina velutipes, characterization, macrophage immunomodulatory activities and its impact on gut microbiota in rats. J. Anim. Physiol. Anim. Nutr.104, 735–748. doi: 10.1111/jpn.13290

  • CrossRef
  • Google Scholar

• 345

  YiY.XuW.WangH. X.HuangF.WangL. M. (2020). Natural polysaccharides experience physiochemical and functional changes during preparation: A review. Carbohydr. Polym.234, e115896. doi: 10.1016/j.carbpol.2020.115896

  • CrossRef
  • Google Scholar

• 346

  YinC.FanX.FanZ.ShiD.GaoH. (2018). Optimization of enzymes-microwave-ultrasound assisted extraction of Lentinus edodes polysaccharides and determination of its antioxidant activity. Int. J. Biol. Macromol.111, 446–454. doi: 10.1016/j.ijbiomac.2018.01.007

  • CrossRef
  • Google Scholar

• 347

  YinZ.LiangZ.LiC.WangJ.MaC.KangW. (2021). Immunomodulatory effects of polysaccharides from edible fungus: a review. Food Sci. Hum. Wellness10, 393–400. doi: 10.1016/j.fshw.2021.04.001

  • CrossRef
  • Google Scholar

• 348

  YinX.YouQ.JiangZ. (2011). Optimization of enzyme assisted extraction of polysaccharides from Tricholoma matsutake by response surface methodology. Carbohydr. Polym.86, 1358–1364. doi: 10.1016/j.carbpol.2011.06.053

  • CrossRef
  • Google Scholar

• 349

  YinX.YouQ.ZhouX. (2015). Complex enzyme-assisted extraction, purification, and antioxidant activity of polysaccharides from the button mushroom, Agaricus bisporus (higher basidiomycetes). Int. J. Med. Mushrooms17, 987–996. doi: 10.1615/intjmedmushrooms.v17.i10.80

  • CrossRef
  • Google Scholar

• 350

  YoonK. N.AlamN.LeeK. R.ShinP. G.CheongJ. C.YooY. B.et al. (2011). Antioxidant and antityrosinase activities of various extracts from the fruiting bodies of Lentinus lepideus. Molecules16, 2334–2347. doi: 10.3390/molecules16032334

  • CrossRef
  • Google Scholar

• 351

  YouQ.YinX.ZhaoY. (2013). Enzyme assisted extraction of polysaccharides from the fruit of Cornus officinalis. Carbohydr. Polym.98, 607–610. doi: 10.1016/j.carbpol.2013.06.036

  • CrossRef
  • Google Scholar

• 352

  YuR.LuoJ.LiuL.PengX. (2024). Hypoglycemic effect of edible fungi polysaccharides depends on their metabolites from the fermentation of human fecal microbiota. Food13, e97. doi: 10.3390/foods13010097

  • CrossRef
  • Google Scholar

• 353

  Yuchen-ZhangDuM. R.ZhangQ. Y.YangS. Y.ChenJ. Q.DanC. M.et al. (2024). Armillariella tabescens-derived polysaccharides alleviated D-Gal-induced neuroinflammation and cognitive injury through enterocerebral axis and activation of keap-1/Nrf2 pathway. Int. J. Biol. Macromol.273, e133035. doi: 10.1016/j.ijbiomac.2024.133035

  • CrossRef
  • Google Scholar

• 354

  ZavadinackM.de Lima BellanD.Fernandes BonaldiM. P.da Silva MilhoriniS.CordeiroL. M. C.Fogagnoli SimasF.et al. (2024). Polysaccharide fractions extracted from Lactarius quieticolor mushroom exhibit immune stimulatory activities on macrophages. Food Res. Int.197, e115205. doi: 10.1016/j.foodres.2024.115205

  • CrossRef
  • Google Scholar

• 355

  ZengW. C.ZhangZ.GaoH.JiaL. R.ChenW. Y. (2012). Characterization of antioxidant polysaccharides from Auricularia auricular using microwave-assisted extraction. Carbohydr. Polym.89, 694–700. doi: 10.1016/j.carbpol.2012.03.078

  • CrossRef
  • Google Scholar

• 356

  ZengF.ZhaoC.PangJ.LinZ.HuangY.LiuB. (2013). Chemical properties of a polysaccharide purified from solid-state fermentation of Auricularia auricular and its biological activity as a hypolipidemic agent. J. Food Sci.78, H1470–H1475. doi: 10.1111/1750-3841.12226

  • CrossRef
  • Google Scholar

• 357

  ZhangZ.DingX.XuZ.WuX.YangC.HaoJ.et al. (2020). Optimization of ultrasonic/microwave assisted extraction (UMAE) and rheological properties of polysaccharides from Auricularia polytricha. Am. J. Biochem. Biotechnol.16, 112–124. doi: 10.3844/ajbbsp.2020.112.124

  • CrossRef
  • Google Scholar

• 358

  ZhangJ.GaoZ.LiS.GaoS.RenZ.JingH.et al. (2016). Purification, characterization, antioxidation, and antiaging properties of exopolysaccharides and endopolysaccharides of the royal sun medicinal mushroom, Agaricus brasiliensis (Agaricomycetes). Int. J. Med. Mushrooms18, 1071–1081. doi: 10.1615/IntJMedMushrooms.v18.i12.20

  • CrossRef
  • Google Scholar

• 359

  ZhangY.GengW.ShenY.WangY.DaiY. C. (2014). Edible mushroom cultivation for food security and rural development in China: Bio-innovation, technological dissemination and marketing. Sustainability6, 2961–2973. doi: 10.3390/su6052961

  • CrossRef
  • Google Scholar

• 360

  ZhangQ.HuM.XuL.YangX.ChangY.ZhuY. (2017). Effect of edible fungal polysaccharides on improving influenza vaccine protection in mice. Food Agric. Immunol.28, 981–992. doi: 10.1080/09540105.2017.1323326

  • CrossRef
  • Google Scholar

• 361

  ZhangW.JiangX.ZhaoS.ZhengX.LanJ.WangH.et al. (2018c). A polysaccharide-peptide with mercury clearance activity from dried fruiting bodies of maitake mushroom Grifola frondosa. Sci. Rep.8, e17630. doi: 10.1038/s41598-018-35945-9

  • CrossRef
  • Google Scholar

• 362

  ZhangY.LeiY.QiS.FanM.ZhengS.HuangQ.et al. (2023). Ultrasonic-microwave-assisted extraction for enhancing antioxidant activity of Dictyophora indusiata polysaccharides: The difference mechanisms between single and combined assisted extraction. Ultrason. Sonochem.95, e106356. doi: 10.1016/j.ultsonch.2023.106356

  • CrossRef
  • Google Scholar

• 363

  ZhangL.LiuY.KeY.LiuY.LuoX.LiC.et al. (2018b). Antidiabetic activity of polysaccharides from Suillellus luridus in streptozotocin-induced diabetic mice. Int. J. Biol. Macromol.119, 134–140. doi: 10.1016/j.ijbiomac.2018.07.109

  • CrossRef
  • Google Scholar

• 364

  ZhangJ.WenC.GuJ.JiC.DuanY.ZhangH. (2019). Effects of subcritical water extraction microenvironment on the structure and biological activities of polysaccharides from Lentinus edodes. Int. J.Biol. Macromol.123, 1002–1011. doi: 10.1016/j.ijbiomac.2018.11.194

  • CrossRef
  • Google Scholar

• 365

  ZhangN.YangB.MaoK.LiuY.ChitrakarB.WangX.et al. (2022b). Comparison of structural characteristics and bioactivity of Tricholoma mongolicum Imai polysaccharides from five extraction methods. Front. Nutr.9, e962584. doi: 10.3389/fnut.2022.962584

  • CrossRef
  • Google Scholar

• 366

  ZhangJ.YeZ.LiuG.LiangL.WenC.LiuX.et al. (2022a). Subcritical water enhanced with deep eutectic solvent for extracting polysaccharides from Lentinus edodes and their antioxidant activities. Molecules27, e3612. doi: 10.3390/molecules27113612

  • CrossRef
  • Google Scholar

• 367

  ZhangC.ZhangL.LiuH.ZhangJ.HuC.JiaL. (2018a). Antioxidation, anti-hyperglycaemia and renoprotective effects of extracellular polysaccharides from Pleurotus eryngii SI-04. Int. J. Biol. Macromol.111, 219–228. doi: 10.1016/j.ijbiomac.2018.01.009

  • CrossRef
  • Google Scholar

• 368

  ZhaoT.CaiY.JiangY.HeX.WeiY.YuY.et al. (2023c). Vaccine adjuvants: mechanisms and platforms. Signal Transduction Targeting Ther.8, e283. doi: 10.1038/s41392-023-01557-7

  • CrossRef
  • Google Scholar

• 369

  ZhaoS.GaoQ.RongC.WangS.ZhaoZ.LiuY.et al. (2020). Immunomodulatory effects of edible and medicinal mushrooms and their bioactive immunoregulatory products. J. Fungi6, e269. doi: 10.3390/jof6040269

  • CrossRef
  • Google Scholar

• 370

  ZhaoQ.JiangY.ZhaoQ.Patrick ManziH.SuL.LiuD.et al. (2023b). The benefits of edible mushroom polysaccharides for health and their influence on gut microbiota: a review. Front. Nutr.10, e1213010. doi: 10.3389/fnut.2023.1213010

  • CrossRef
  • Google Scholar

• 371

  ZhaoM.KuangF.ZhangY.LvG. (2023a). Effects of hydrolysis condition and detection method on the monosaccharide composition analysis of polysaccharides from natural sources. Separations11, e2. doi: 10.3390/separations11010002

  • CrossRef
  • Google Scholar

• 372

  ZhaoY. M.SongJ. H.WangJ.YangJ. M.WangZ. B.LiuY. H. (2016a). Optimization of cellulase-assisted extraction process and antioxidant activities of polysaccharides from Tricholoma mongolicum Imai. J. Sci. Food Agric.96, 4484–4491. doi: 10.1002/jsfa.7662

  • CrossRef
  • Google Scholar

• 373

  ZhaoY. T.TangY. Q.LiuK. Z.ZhangY. (2014). A study about microwave-assisted extraction of polysaccharides form medicinal mushrooms Fomitopsis ulmaria (Sor.: For.) bond. Et sing. Adv. Mater. Res.1073–1076, 1837–1840. doi: 10.4028/www.scientific.net/amr.1073-1076.1837

  • CrossRef
  • Google Scholar

• 374

  ZhaoY. M.WangJ.WuZ. G.YangJ. M.LiW.ShenL. X. (2016b). Extraction, purification and anti-proliferative activities of polysaccharides from Lentinus edodes. Int. J. Biol. Macromol.93, 136–144. doi: 10.1016/j.ijbiomac.2016.05.100

  • CrossRef
  • Google Scholar

• 375

  ZhengS.ZhangW.LiuS. (2020). Optimization of ultrasonic-assisted extraction of polysaccharides and triterpenoids from the medicinal mushroom Ganoderma lucidum and evaluation of their in vitro antioxidant capacities. PloS One15, e0244749. doi: 10.1371/journal.pone.0244749

  • CrossRef
  • Google Scholar

• 376

  ZhongJ.FangL.ChenR.XuJ.GuoD.GuoC.et al. (2021). Polysaccharides from sporoderm-removed spores of Ganoderma lucidum induce apoptosis in human gastric cancer cells via disruption of autophagic flux. Oncol. Lett.21, e425. doi: 10.3892/ol.2021.12686

  • CrossRef
  • Google Scholar

• 377

  ZhongM.MiaoY.LanY.MaQ.LiK.ChenW. (2024). Effects of Exidia yadongensis polysaccharide as emulsifier on the stability, aroma, and antioxidant activities of fat-free stirred mango buffalo yogurt. Int. J. Biol. Macromol.276, e133785. doi: 10.1016/j.ijbiomac.2024.133785

  • CrossRef
  • Google Scholar

• 378

  ZhouW.HeY.LiuF.LiaoL.HuangX.LiR.et al. (2021). Carboxymethyl chitosan-pullulan edible films enriched with galangal essential oil: Characterization and application in mango preservation. Carbohydr. Polym.256, e117579. doi: 10.1016/j.carbpol.2020.117579

  • CrossRef
  • Google Scholar

• 379

  ZhouY.JiaY.XuN.TangL.ChangY. (2023). Auricularia auricula-judae (Bull.) polysaccharides improve obesity in mice by regulating gut microbiota and TLR4/JNK signaling pathway. Int. J. Biol. Macromol.250, e126172. doi: 10.1016/j.ijbiomac.2023.126172

  • CrossRef
  • Google Scholar

• 380

  ZhuY.LiQ.MaoG.ZouY.FengW.ZhengD.et al. (2014). Optimization of enzyme-assisted extraction and characterization of polysaccharides from Hericium erinaceus. Carbohydr. Polym.101, 606–613. doi: 10.1016/j.carbpol.2013.09.099

  • CrossRef
  • Google Scholar