1 Introduction

Ganoderma lucidum, commonly known as Lingzhi or Reishi, has been used in traditional Chinese medicine for centuries and is regarded as a highly valuable medicinal fungus (Kiss et al., 2021). Its primary bioactive compounds, including polysaccharides, triterpenoids, and other secondary metabolites, have been shown to possess various therapeutic properties, such as immunomodulatory, anti-inflammatory, antioxidant, and anticancer activities (Bidegain et al., 2019; Qiu et al., 2023). Due to these medicinal benefits, the global demand for G. lucidum has rapidly increased, with extensive use in dietary supplements, functional foods, and pharmaceutical ingredients. This growing demand places higher requirements on cultivation, necessitating advancements to ensure a consistent supply of high-quality G. lucidum with potent pharmacological efficacy.

In the cultivation of medicinal fungi, selecting an appropriate substrate plays a critical role in determining the growth rate, yield, and overall quality of fungal biomass. The substrate provides essential nutrients for fungal growth and directly influences the accumulation of bioactive compounds. A carefully chosen substrate can ensure optimal nutrient supply, promote vigorous growth, and support the development of key secondary metabolites. It was reported that using plant waste such as straw and corn husks as a substrate can significantly enhance the yield and nutritional content of certain medicinal fungi (Balasubramanian and Kannan, 2023). For G. lucidum, substrate selection is especially important, as a suitable substrate can not only enhance productivity but also increase the therapeutic efficacy of the final fruiting bodies of G. lucidum (Khoo et al., 2021). 

Different substrates provide varying levels of essential nutrients, such as carbon, nitrogen, and trace minerals, all of which affect the growth dynamics and biochemical composition of G. lucidum. Traditional substrates, including hardwood sawdust and agricultural by-products like wheat bran or rice hulls, are widely used (Atila, 2020; Fedorov et al., 2022). However, their suitability varies based on regional availability, cost, and the quality of the final product. Studies have explored the potential of novel substrates; for instance, adding 2% diaper core material to sawdust bio-waste achieved a biological efficiency of 36%, significantly higher than the 21% achieved with commercial substrates (Khoo et al., 2021). Understanding the relationship between substrate composition and the quality of G. lucidum is crucial for developing efficient and sustainable cultivation practices that can produce high-quality medicinal G. lucidum.

This study systematically assesses the effects of different cultivation substrates on the yield and quality of G. lucidum, with an in-depth analysis of how various substrate combinations specifically influence productivity and the concentration of bioactive compounds. It intends to identify the optimal cultivation strategy for G. lucidum, focusing on enhancing yield while emphasizing the accumulation of its medicinal components to maximize therapeutic efficacy. This study will provide a scientific foundation for optimizing cultivation techniques to meet the market demand for high-quality medicinal G. lucidum and promote the sustainable development of cultivation practices.

2 Ganoderma lucidum and Its Economic Significance

2.1 Biological characteristics of Ganoderma lucidum

Ganoderma lucidum is a wood-degrading mushroom that thrives on various lignocellulosic substrates. It exhibits a distinctive morphology characterized by a shiny, reddish-brown cap and a kidney-shaped fruiting body. The growth of G. lucidum is highly dependent on the substrate composition, which can include materials such as rice agro-residues, sunflower seed hulls, and various hardwoods (Bidegain et al., 2019; Qiu et al., 2023). The optimal growth conditions involve a substrate rich in cellulose and lignin, with a low nitrogen content, which supports efficient mycelial development and fruiting body formation (Sudheer et al., 2018).

G. lucidum is renowned for its bioactive compounds, including triterpenoids, ganoderic acids, polysaccharides, and phenolic compounds. The biosynthesis of these compounds is influenced by the substrate used for cultivation. For instance, substrates enriched with additives like olive oil and copper can enhance the production of ganoderic acids and other bioactive molecules (Bidegain et al., 2019). The presence of specific nutrients and minerals in the substrate, such as manganese sulfate, can also significantly impact the concentration of these compounds in the fruiting bodies (Kurd-Anjaraki et al., 2021).

2.2 Medicinal and economic value of Ganoderma lucidum

G. lucidum has been used in traditional medicine for centuries due to its wide range of therapeutic properties. It is known for its antioxidant, anti-inflammatory, and immunomodulatory effects, which are attributed to its rich content of bioactive compounds like polysaccharides and triterpenoids (Sudheer et al., 2018; Kachrimanidou et al., 2023). These compounds have been shown to enhance immune function, reduce oxidative stress, and potentially inhibit the growth of cancer cells, making G. lucidum a valuable medicinal resource.

The economic significance of G. lucidum extends beyond its medicinal applications. It is a high-value crop in the global market, with substantial demand in countries like China, Japan, and Korea. The cultivation of G. lucidum using locally available agro-wastes not only reduces production costs but also adds value to agricultural by-products, promoting sustainable agricultural practices (Sudheer et al., 2018; Amiri-Sadeghan et al., 2022). The ability to cultivate G. lucidum on diverse substrates makes it an economically viable option for farmers and contributes to the global economy by providing a steady supply of this medicinal mushroom.

2.3 Current state of Ganoderma lucidum production

G. lucidum is predominantly produced in East Asia, particularly in China, Japan, and Korea, where traditional cultivation practices have been refined over centuries. Modern cultivation techniques involve the use of various lignocellulosic substrates, including hardwood sawdust, wheat straw, and agro-industrial by-products like grape pomace and cheese whey (Atila, 2020; Kachrimanidou et al., 2023). These substrates are often supplemented with nutrients to enhance yield and bioactive compound production.

The expansion of G. lucidum production faces several challenges, including the need for optimized substrate formulations and efficient cultivation techniques. The selection of suitable substrates that maximize yield and bioactive compound content is crucial. Research has shown that substrates with high cellulose and lignin content and low nitrogen levels are preferred for optimal growth (Atila, 2020). Additionally, the valorization of agro-industrial by-products as substrates presents an opportunity to reduce waste and lower production costs, thereby promoting sustainable practices in the industry. The development of innovative cultivation methods and substrate formulations can further enhance the economic viability and medicinal value of G. lucidum, providing reference for industry growth and expansion.

3 Types and Characteristics of G. lucidum Cultivation Substrates

3.1 Woody substrates

Woody substrates are commonly used for the cultivation of G. lucidum due to their high lignocellulosic content, which supports robust mycelial growth and fruiting body development. Various types of wood residues, such as sawdust and wood chips from different tree species, have been evaluated for their suitability. For instance, sawdust and wood chips from Betula spp., Populus tremula, Picea abies, Pinus sylvestris, and Larix sp. have been tested, with Betula spp. and P. tremula showing the highest yields and β-glucan content when combined with specific strains like MUS192 (Cortina-Escribano et al., 2020). 

Additionally, substrates like poplar sawdust and wood chips have been found to produce high yields and desirable G. lucidum morphology under solid-state fermentation conditions. According to the study by Kuhar et al. (2018), sawdust and wood chips from Populus nigra and Pinus radiata can serve as solid-state fermentation substrates, enhancing the yield of G. lucidum fruiting bodies (63 grams per kilogram of dry substrate). And growth can be promoted through simple moisture management techniques.The use of sawdust from coconut wood logs has also been highlighted for its efficiency, supporting early spawn run and high bioefficiency (Thiribhuvanamala and Krishnamoorthy, 2021).

3.2 Agricultural waste substrates

Agricultural waste substrates offer a sustainable and cost-effective alternative for G. lucidum cultivation. These substrates include materials such as wheat straw, cottonseed hulls, corn cobs, and other agro-wastes. Studies have shown that wheat straw, soybean straw, and bean straw can serve as effective basal substrates, with wheat straw and soybean straw providing high yields and biological efficiency (Atila, 2020). Under submerged fermentation conditions, corncobs and straw have been found to be suitable for producing laccase from G. lucidum, an enzyme capable of breaking down lignin. Among the treatments, those with added corncobs demonstrated the highest enzyme activity, indicating their potential as a medium for cultivating G. lucidum (Yuliana et al., 2020).

Additionally, combinations of agro-wastes like broad bean stalks, cotton stalks, maize straw, and sugarcane bagasse, supplemented with wheat bran or corn gluten, have been evaluated, with cotton stalks and sugarcane bagasse showing promising results for mycelial growth and yield (Rashad et al., 2019). In Malaysia, combinations of rubber wood sawdust with empty fruit bunch fiber from oil palm have demonstrated high biological efficiency and antioxidant potential (Sudheer et al., 2018).

3.3 Synthetic and modified substrates

Synthetic and modified substrates are designed to enhance the growth conditions for G. lucidum by incorporating various supplements and treatments. For example, the use of sawdust from common pine (Pinus sylvestris) activated through hydrodynamic cavitation and supplemented with nitrogen salts has been shown to significantly improve colonization and growth rates (Fedorov et al., 2022). Studies have shown that adding supplements such as olive oil and copper to the medium can significantly alter the bioactive chemical composition of G. lucidum fruiting bodies, increasing the content of triterpenoids, ganoderic acids, high-molecular-weight carbohydrates, and phenolic compounds (Bidegain et al., 2019).

Substrates enriched with chemical supplements like ammonium nitrate, manganese sulfate, and nano manganese oxide have been tested, with combinations such as poplar wood chips and soybean meal or wheat bran showing high yields and enhanced ganoderic acid content (Kurd-Anjaraki et al., 2021). The incorporation of biochar into agro-waste-based substrates has also been explored, with findings indicating that biochar levels can influence yield, biological efficiency, and nutrient content of the fruiting bodies (Rashad et al., 2019).

4 Effects of Different Substrate Composition on Yield

4.1 The impact of substrate types on yield

Woody substrates such as sawdust and wood chips have been extensively studied for their impact on the yield of G. lucidum. For instance, sawdust from Betula spp. and Populus tremula has been shown to support high yields, particularly when combined with specific strains like MUS192 (Cortina-Escribano et al., 2020). Similarly, sawdust from Pinus sylvestris, when activated hydrodynamically, has demonstrated efficient colonization and increased extractive substance content, which can enhance yield (Fedorov et al., 2022). Additionally, substrates like poplar sawdust have been found to produce significant yields, with the highest recorded at 63 g dry weight per kilogram of dry substrate (Kuhar et al., 2018).

Agricultural waste substrates such as wheat straw, cotton stalks, and rice straw have also been evaluated for their effectiveness in cultivating G. lucidum. Among these, cotton stalks supplemented with wheat bran have shown superior performance in terms of yield and biological efficiency (Rashad et al., 2019). Wheat straw and bean straw have been identified as viable alternatives to traditional sawdust substrates, with yields ranging from 28.6 g/kg to 86.1 g/kg (Atila, 2020). Additionally, combinations of empty fruit bunch fiber and sawdust have achieved high biological efficiency and yield (Sudheer et al., 2018).

Industrial by-products like hazelnut shells and olive by-products have been explored as substrates for G. lucidum cultivation. Hazelnut shells, due to their high lignin content, have been found to support good mycelial growth and fruiting body production (Puliga et al., 2022). Olive pruning residues, when mixed with wheat straw, have also shown promise, enhancing the glucan content and prebiotic properties of the mushrooms (Koutrotsios et al., 2019).

4.2 The impact of substrate formulation and nutrition on yield

The carbon to nitrogen (C:N) ratio of the substrate is a critical factor influencing the yield of G. lucidum. Studies have shown that substrates with a high C:N ratio, such as those containing high cellulose and lignin content, tend to produce higher yields. For example, substrates with a low nitrogen content and high cellulose:lignin ratio have been positively correlated with increased mushroom yield (Atila, 2020). The addition of trace elements and vitamins can significantly impact the yield of G. lucidum. For instance, the introduction of nitrogen salts like (NH₄)₂SO₄ and Na₂HPO₄ into sawdust substrates has been shown to enhance the growth rate and yield of the G. lucidum (Fedorov et al., 2022). The study found that the sawdust medium enriched with ammonium sulfate and disodium phosphate significantly increased the growth rate of the strain, reaching 3.22±0.48 mm/day, allowing the substrate to be fully covered by mycelium by day 13 (Figure 1). Compared to the medium without added nitrogen salts, the mycelium proliferated faster under nitrogen-enriched conditions, and the content of water-soluble extracts in the substrate increased from 4.37% to 6.32%.

Organic and inorganic fertilizers can also play a role in improving substrate quality and yield. The use of wheat bran as a supplement in various substrates has been found to enhance mycelial growth and yield. For example, substrates supplemented with wheat bran have shown higher biological efficiency and yield compared to those without (Rashad et al., 2019; Thiribhuvanamala and Krishnamoorthy, 2021).

4.3 The impact of physical and chemical properties of substrates on yield

The physical properties of the substrate, such as particle size and moisture content, also are crucial for the successful cultivation of G. lucidum. A suitable moisture content is associated with higher yield. For example, research has found that using a palm sawdust substrate with 67.36% humidity significantly increases the yield of G. lucidum. This moisture level provides an optimal growth environment, achieving a yield of 66.25 g/kg (Ohmayed et al., 2020). Additionally, substrates with smaller particle sizes, like sawdust, tend to support better mycelial colonization and fruiting body formation (Cortina-Escribano et al., 2020; Fedorov et al., 2022).

Chemical properties such as pH value and nutrient content also significantly affect the yield of G. lucidum. Substrates with a pH value around 6.93 have been found to be optimal for mushroom growth (Ohmayed et al., 2020). Moreover, substrates with balanced electrical conductivity and salinity levels can enhance nutrient availability and support higher yields (Kuhar et al., 2018).

5 Impact of Cultivation Substrates on Quality Parameters

5.1 Active compound content 

The content of active compounds such as polysaccharides and triterpenoids in G. lucidum is significantly influenced by the type of cultivation substrate used. Ye et al. (2018) found that spraying salicylic acid and calcium ions during the fruiting body development stage of G. lucidum can significantly enhance its triterpenoid and polysaccharide content. The results showed that using salicylic acid alone increased triterpenoid content by 23.32%, while combining it with calcium ions increased polysaccharide and triterpenoid contents by 9.02% and 13.61%, respectively. Gene expression analysis indicated that these treatments affected the transcription levels of relevant genes, with particularly significant upregulation observed in triterpenoid biosynthesis genes. Another study found that salicylic acid stimulates the biosynthesis of secondary metabolites not only by directly promoting triterpenoid synthesis but also by increasing the generation of mitochondrial reactive oxygen species (ROS). Salicylic acid can inhibit the activity of mitochondrial complex III, leading to elevated ROS levels, which in turn regulate triterpenoid synthesis (Liu et al., 2018).

A substrate formulation with added olive oil and copper has also been found to affect the bioactive chemical composition of G. lucidum, including total triterpenoids and high-molecular-weight carbohydrates (Bidegain et al., 2019). Additionally, olive by-products used as a cultivation medium for G. lucidum not only enhance the content of bioactive components, such as α-glucans and β-glucans, but also increase protein content, indicating that olive residues contribute to improving the functional and prebiotic properties of G. lucidum (Koutrotsios et al., 2019). The optimization of media composition in solid-state fermentation using highland barley grains also demonstrated a significant increase in polysaccharide production, validating the importance of substrate composition in enhancing active compound content (Liu et al., 2020).

5.2 Physical properties 

The physical properties of G. lucidum, such as color, size, and texture, are also affected by the cultivation substrate. Studies have shown that the cultivation medium of G. lucidum significantly influences the surrounding microbial community and metabolic pathways at different growth stages. Microbial diversity is particularly high during the elongation stage, with changes in nitrogen, phosphorus, and potassium levels impacting the physical and chemical properties of the medium (Zhang et al., 2018). Bidegain et al. (2019) highlighted that the medium formulation for G. lucidum affects the activity of its chemical components, with the addition of various supplements leading to notable increases in triterpenoid and phenolic compound content, and different formulations resulting in variations in color and texture.

Moreover, introducing nitrogen salts into the substrate increased the growth rate and colonization efficiency of the fungal body, producing more robust G. lucidum fruiting bodies (Fedorov et al., 2022). Different forms of organic nitrogen, such as β-adenosine, have a pronounced effect on fungal growth. For white-rot fungi like G. lucidum, additional organic nitrogen sources promote mycelial biomass accumulation, further enhancing the growth and reproductive efficiency of the mycelium (Hennicke et al., 2022).

5.3 Quality standards and evaluation methods

Quality standards and evaluation methods for G. lucidum involve both conventional and advanced techniques to assess the content of bioactive compounds and physical properties. Traditional colorimetric methods combined with FT-MIR spectroscopy and chemometric approaches for evaluating total triterpenoids, ganoderic acids, high-molecular-weight carbohydrates, and phenolic compounds in G. lucidu fruiting bodies can reveal the impact of substrate composition and harvest time on these component levels. FT-MIR spectroscopy can effectively differentiate G. lucidu samples from various substrates (Bidegain et al., 2019). Characterization techniques, such as FT-IR and nuclear magnetic resonance (NMR), have been used to investigate the structural features of G. lucidu polysaccharides—such as carbohydrate composition, molecular weight, and higher-order conformations—and their relationship with immunomodulatory activity (Li et al., 2019).

Another study utilized hydrophilic interaction chromatography combined with multivariate analysis to assess polysaccharide quality in G. lucidu samples from different regions. This method distinguished samples from Zhejiang and other regions, identifying components like arabinose and mannose as potential chemical markers through further principal component analysis (PCA) (Zhao et al., 2020). FTIR spectroscopy also enables rapid fingerprinting of bioactive compounds in G. lucidu extracts, showing potential for quality control by distinguishing samples based on origin or storage conditions (Popa et al., 2021). These methods offer a comprehensive quality assessment approach that can standardize the medicinal quality of G. lucidu products.

6 Case Studies

6.1 Efficient cultivation of G. lucidum using coconut sawdust substrate

Traditional log-based cultivation methods for G. lucidum are time-consuming and yield low output, making it challenging to meet market demand. Researchers have focused on developing alternative cultivation substrates to increase the yield and efficiency of G. lucidum cultivation (Bidegain et al., 2019; Atila, 2020; Thiribhuvanamala and Krishnamoorthy, 2021). One notable case involves the use of coconut sawdust as a substrate to evaluate the effects of different lignocellulosic substrates on the artificial cultivation of G. lucidum (Thiribhuvanamala and Krishnamoorthy, 2021). The experiment utilized coconut sawdust, coconut petioles, coconut fiber waste, areca palm sawdust, and mixed sawdust substrates, with wheat bran added to enhance nutrient content. Results indicated that the coconut sawdust substrate had the highest yield and biological efficiency (44.3%), with a shorter growth cycle for fruiting bodies and a high yield per unit substrate, reaching 77.5 grams (Figure 2). In contrast, coconut fiber waste had a lower yield, with a biological efficiency of only 25.6%, highlighting the significant impact of substrate type on G. lucidum growth.

The findings suggest that coconut sawdust is an ideal substrate for G. lucidum cultivation. Its unique nutrient and lignin content facilitate rapid colonization and growth of the fungus, making it highly effective for commercial cultivation. This research provides practical insights for cost-effective, large-scale cultivation of G. lucidum, while also promoting efficient recycling of agricultural waste.

6.2 Study on the effect of substrate formulation on the active components of Ganoderma lucidum

The active components of G. lucidum mainly include high-molecular-weight carbohydrates (such as polysaccharides), triterpenoids (particularly ganoderic acids), and phenolic compounds, all of which are crucial for its medicinal efficacy. Research indicates that the chemical composition of G. lucidum is influenced by the cultivation substrate and added supplements. To optimize the medicinal value of G. lucidum, studies have assessed the impact of various substrate formulations and additive combinations on the active components in the fruiting bodies.

One study evaluated the influence of different substrate formulations on the chemical composition of G. lucidum, focusing on changes in bioactive substance content. The findings showed significant differences in triterpenoids, phenolic compounds, and high-molecular-weight polysaccharides when grown on various substrates, such as sunflower seed hulls and rice residue, along with copper and olive oil supplementation (Bidegain et al., 2019). The results indicated that copper supplementation contributed to an increase in triterpenoids and phenolic compounds, while olive oil enhanced the content of water-soluble phenolic compounds (Table 1). Additionally, maturity and harvest time significantly affected the chemical composition, with differences observed in the components of G. lucidum bodies collected at successive harvests.

Another study used olive by-products as the substrate. The results showed that G. lucidum grown on olive pruning residues had significantly higher α-glucan and β-glucan content than that grown on beech sawdust. Furthermore, these G. lucidum samples exhibited greater antioxidant activity and prebiotic potential, supporting the growth of beneficial gut bacteria such as Lactobacillus acidophilus and L. gasseri (Koutrotsios et al., 2019).

7 Environmental and Economic Considerations

7.1 Sustainability of substrate materials

The sustainability of substrate materials for cultivating G. lucidum is a critical factor in ensuring environmentally friendly and economically viable mushroom production. Various studies have demonstrated the potential of using agro-wastes and other lignocellulosic materials as substrates. For instance, the use of agro-wastes such as broad bean stalks, cotton stalks, maize straw, rice straw, sugarcane bagasse, and wheat straw has been shown to support the growth of G. lucidum effectively (Rashad et al., 2019). Additionally, the use of locally available residues like coconut wood log sawdust and other tree residues has been explored, showing promising results in terms of yield and biological efficiency (Thiribhuvanamala and Krishnamoorthy, 2021). These materials are not only abundant but also help in reducing agricultural waste, contributing to a more sustainable cultivation practice.

7.2 Economic viability of different substrates

The economic viability of different substrates is influenced by their availability, cost, and the yield they produce. Studies have shown that substrates like cotton stalks, rice straw, and sugarcane bagasse, when supplemented with wheat bran, can significantly enhance the yield and biological efficiency of G. lucidum (Rashad et al., 2019). Similarly, the use of mixed sawdust and other agro-residues has been found to be cost-effective while providing high yields (Thiribhuvanamala and Krishnamoorthy, 2021). The combination of empty fruit bunch fiber with sawdust, for example, has been reported to achieve a biological efficiency of 27%, making it a viable option for commercial cultivation (Sudheer et al., 2018). The economic benefits are further enhanced by the reduced need for chemical fertilizers and the potential for using spent mushroom substrates in other applications, such as bioethanol production (Sudhakar et al., 2021).

7.3 Reducing waste through optimized substrate use

Optimizing substrate use not only improves the yield and quality of G. lucidum but also plays a significant role in waste reduction. The recycling of agro-wastes and other lignocellulosic materials into mushroom substrates helps in managing agricultural residues effectively. For instance, the use of spent mushroom substrates for bioethanol production has been explored, demonstrating the potential for converting waste into valuable biofuels (Sudhakar et al., 2021). Additionally, the incorporation of food waste and other biodegradable materials into mushroom substrates can further enhance waste reduction efforts. Studies have shown that using a combination of food waste and lignocellulosic materials can significantly boost mycelium growth and mushroom yield, promoting a circular economy (Soh et al., 2021). This approach not only reduces the environmental impact of waste but also provides an additional revenue stream for mushroom growers.

8 Concluding Remarks

The study on the effects of various cultivation substrates on the yield and quality of G. lucidum yielded several significant insights. Studies have demonstrated that different wood residues and agro-wastes can serve as effective substrates for cultivating G. lucidum. For instance, wood residues from Betula spp. and Populus tremula have been shown to support high yields and β-glucan content, particularly when combined with specific strains like MUS192 and subjected to cold treatments. Similarly, lignocellulosic wastes such as wheat straw, sunflower meal, and soybean straw have been identified as promising substrates, with wheat straw showing particularly high biological efficiency. Additionally, mixed substrates, such as combinations of poplar wood chips with soybean meal or wheat bran, have been found to enhance the nutritional and medicinal properties of the fruiting bodies. The use of local agro-residues, such as coconut wood log sawdust and pecan wood chips, has also been effective, demonstrating significant bioefficiency and yield.

The findings from these studies suggest several avenues for future research and practical applications in the G. lucidum cultivation industry. Future research should focus on optimizing substrate formulations to maximize yield and bioactive compound content. This includes exploring the effects of different substrate combinations and enrichment with chemical supplements, such as manganese sulfate and nano manganese oxide, which have been shown to enhance yield and ganoderic acid content. Additionally, the impact of environmental conditions, such as temperature and humidity, on the cultivation process should be further investigated to develop standardized protocols for different substrates. For industry adoption, it is recommended to utilize locally available agro-wastes and wood residues as substrates to reduce costs and promote sustainability. Substrates like wheat straw, coconut wood log sawdust, and pecan wood chips have demonstrated high potential and should be considered for large-scale cultivation. Moreover, the incorporation of cold treatments and substrate enrichment strategies can further enhance the yield and quality of G. lucidum fruiting bodies, making the cultivation process more efficient and economically viable.

The cultivation of G. lucidum on various substrates offers a sustainable and cost-effective approach to producing this valuable medicinal mushroom. The study highlights the importance of substrate selection and optimization in achieving high yields and enhancing the bioactive properties of the fruiting bodies. By leveraging locally available agro-wastes and wood residues, the G. lucidum cultivation industry can not only improve its economic efficiency but also contribute to environmental sustainability. Future research and industry practices should continue to focus on optimizing substrate formulations and cultivation conditions to fully realize the potential of G. lucidum cultivation.

Acknowledgments

The author sincerely thanks Ms. Li from the Medicinal Plant Research Center for her invaluable assistance in data analysis and organization. Heartfelt appreciation is also extended to the two anonymous peer reviewers for their meticulous review, which improved the manuscript.

Conflict of Interest Disclosure

The author affirms that this research was conducted without any commercial or financial relationships that could be construed as a potential conflict of interest.

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