1 Introduction

Ganoderma lucidum is one of the traditional valuable medicinal fungi in China. It was classified as a “superior-grade” medicinal material in Shennong’s Classic of Materia Medica and has long been recognized for its medicinal value in tonifying qi, calming the mind, and strengthening healthy qi. Modern studies have shown that G. lucidum is rich in bioactive compounds such as polysaccharides and triterpenoids, which exhibit various pharmacological activities, including immunomodulatory, antioxidant, antitumor, and metabolic regulatory effects. Therefore, its application value in food, nutritional supplements, pharmaceuticals, functional products, and wellness markets has continued to increase (Thakur et al., 2024). In recent years, with the rapid development of the health industry and the increasing health awareness of consumers, market demand for G. lucidum-related products has continued to expand. The stable and large-scale supply of safe, high-quality G. lucidum raw materials and derivative products has become an important foundation for supporting industrial development (Wu et al., 2024). However, traditional G. lucidum cultivation systems still have certain limitations in yield stability, quality consistency, resource-use efficiency, and contamination control. Some production practices still rely on experience-based management, which may lead to imprecise environmental regulation, large variations in product quality, frequent occurrence of pests and diseases, and inconsistent production standards. These problems make it difficult to meet modern market demand for high-quality, safe, and stable G. lucidum products (Ghafoor et al., 2024). Therefore, promoting technological upgrading and the construction of standardized production systems has become a practical requirement for the high-quality development of the G. lucidum industry.

G. lucidum is a typical wood-decaying fungus, and its mycelial growth and fruiting body formation are highly sensitive to environmental conditions such as temperature, humidity, light, air circulation, CO₂ concentration, and nutrient supply. Under traditional open-field cultivation or simple greenhouse cultivation, G. lucidum production is easily affected by seasonal changes, climatic fluctuations, and natural disasters, often resulting in slow mycelial growth, abnormal fruiting body morphology, unstable yield, and fluctuations in active component contents. Meanwhile, the scarcity and instability of wild resources, high dependence on wood-based substrates, and obvious batch-to-batch differences in active components further highlight the need to establish efficient, controllable, and sustainable production systems. With the development of modern agricultural facility technologies and bioprocessing technologies, facility-based cultivation has gradually become an important direction for the transformation and upgrading of the G. lucidum industry. Indoor mushroom houses, environmentally controllable “mushroom factories,” liquid spawn systems, and submerged fermentation technologies can precisely regulate temperature, humidity, gas composition, light, and nutritional conditions, thereby improving production stability and quality consistency (Feng et al., 2024; Ghafoor et al., 2024; Liu et al., 2024). In addition, facility-based cultivation can reduce the risks associated with heavy metals, toxic residues, and competitive microorganisms by separating substrates from soil, while also supporting standardized operations that comply with pharmacopoeial requirements and good manufacturing practices (Wu et al., 2024).

In facility-based G. lucidum cultivation, environmental control technology is a core factor affecting production performance and product quality. Temperature and humidity are directly related to mycelial expansion, primordium formation, and fruiting body development; ventilation affects changes in CO₂ concentration and plays an important role in pileus formation, stipe elongation, and morphogenesis; light conditions are closely associated with primordium differentiation, color formation, and the accumulation of bioactive substances. Studies have shown that controlled environments can optimize fruiting body morphology, yield, and the accumulation of bioactive components such as polysaccharides and triterpenoids by regulating key factors such as light spectrum, temperature profiles, and oxygen supply. Improper environmental regulation can easily lead to contamination, excessive stipe elongation, malformed pilei, and insufficient accumulation of active components. Therefore, establishing a scientific, stable, and reproducible environmental control system is an important basis for improving the level of facility-based G. lucidum cultivation. Meanwhile, the application of low-cost IoT-based temperature and humidity sensing, automatic control, and real-time monitoring technologies has further enhanced environmental management during G. lucidum production, helping to achieve contaminant control, compliance with quality indicators, and standardized whole-process management (Nguyen et al., 2023; Liu et al., 2024).

This study will explore the key technical pathways for environmental control and standardized production in facility-based G. lucidum cultivation. It will focus on the environmental regulation points, key production procedures, and quality control systems of facility-based G. lucidum cultivation; examine the application value of intelligent and digital technologies in G. lucidum production; and propose development pathways for standardized production in light of the current industrial context. The study aims to integrate the development needs of the G. lucidum industry with modern controllable cultivation technologies. By summarizing research progress in facility design, temperature, humidity, CO₂, light quality, ventilation regulation, liquid spawn, and fermentation processes, it seeks to provide technical references for stable and high-quality G. lucidum production. This study will also examine how precise environmental regulation can support emerging quality standards, increase the yield of fruiting bodies and bioactive metabolites, reduce contamination and resource consumption, and thereby promote G. lucidum production toward precision, standardization, and intelligentization, enhance product quality and industrial competitiveness, and facilitate the sustainable and high-quality development of the G. lucidum industry.

2 Basic Characteristics of Facility-Based Ganoderma lucidum Cultivation

2.1 Specific environmental requirements for Ganoderma lucidum growth

Ganoderma lucidum is a thermophilic wood-decaying fungus. Its mycelial growth, primordium differentiation, and fruiting body formation show distinct stage-specific requirements for temperature, humidity, light, CO₂ concentration, pH, and substrate conditions. Studies have shown that G. lucidum mycelia generally grow well at 25°C,~32°C, under near-neutral to slightly acidic pH and relatively high humidity, whereas fruiting body formation depends more on stable temperature and humidity, appropriate light, and favorable gas exchange conditions (Al-Kaabi and Hussien, 2025). For example, Magday (2014) found that a wild Philippine strain of G. lucidum showed better mycelial growth at pH 6.0, 32°C, and under light conditions; Lengare et al. (2023) reported that vegetative growth performed well at 25°C, 95%~100% relative humidity, and a 12 h light/12 h dark cycle. Therefore, facility-based G. lucidum cultivation is not merely the control of a single environmental parameter, but requires integrated regulation according to different growth stages.

During fruiting body formation, G. lucidum has stricter requirements for air humidity, light, and ventilation. Appropriate humidity promotes primordium formation and pileus expansion, whereas insufficient ventilation may lead to elevated CO₂ concentration in the facility, resulting in elongated stipes, poor pileus development, or abnormal morphology. A Korean bed-cultivation study showed that 28°C~30°C, light intensity of 1,000~1,500 lux, and approximately 1% CO₂ were suitable for fruiting body formation. Recent studies on light quality regulation further indicate that green light can promote stipe elongation, pileus expansion, fresh weight increase, and the contents of crude protein, polysaccharides, triterpenoids, and extracellular enzyme activity, whereas red light may inhibit fruiting body differentiation. Therefore, spectral selection and lighting regime design should be emphasized in factory cultivation (Liu et al., 2024). This suggests that light not only affects the appearance quality of G. lucidum, but is also closely related to the accumulation of active components.

In addition to physical environmental factors, substrate chemical properties and facility cleanliness are also important factors affecting stable G. lucidum production. Continuous cultivation can alter organic matter, pH, salinity, and enzyme activities in soil or substrates, leading to changes in the metabolite composition of fruiting bodies (Wang et al., 2022). Meanwhile, competitive fungi such as Trichoderma can proliferate readily in G. lucidum cultivation environments and inhibit G. lucidum growth through volatile and non-volatile metabolites, causing contamination and yield loss (Lim et al., 2024). Therefore, facility-based G. lucidum production should integrate temperature and humidity regulation, light management, CO₂ control, pH and salinity adjustment, substrate disinfection, and spatial sanitation management to establish a stable and reproducible environmental control system.

2.2 Main models of facility-based Ganoderma lucidum cultivation

With the development of modern agricultural facility technologies, G. lucidum production has gradually shifted from traditional open-field cultivation and simple greenhouse cultivation toward facility-based, controllable, and standardized models. At present, greenhouses and indoor mushroom houses are common solid-state cultivation models for fruiting body production. They usually use sawdust, agricultural residues, or logs as substrates, and are equipped with functional areas such as inoculation rooms, incubation rooms, sterilization rooms, fruiting rooms, packaging rooms, and waste disposal areas (Thakur et al., 2024; Akçay et al., 2025). Standardized bed cultivation generally uses formulated sawdust-based substrates and promotes fruiting body formation under controlled temperature, light, and CO₂ conditions. Compared with traditional open-field cultivation, greenhouse and mushroom-house cultivation can reduce the influence of external climate fluctuations and improve production stability and year-round supply capacity.

In solid-state facility cultivation, bag cultivation and multi-layer rack cultivation are important forms of current large-scale production. Bag cultivation uses lignocellulosic wastes such as sawdust, rice straw, wheat straw, and hazelnut shells as main raw materials. By optimizing substrate formulations, it can shorten mycelial colonization time and improve yield and the number of fruiting flushes (Akçay et al., 2025). Multi-layer rack cultivation improves space utilization by vertically arranging cultivation bags, facilitating unified sterilization, inoculation, incubation, and fruiting management. Some regions have also developed permanent greenhouse sand-bed cultivation models, in which cultivation substrates are separated from soil using plastic membranes and combined with sprinkler and shading systems. This can reduce the risks of heavy metals, pesticide residues, and competitive microbial contamination, and increase yield by approximately 30% compared with traditional open-field methods. This type of model combines environmental controllability with quality and safety advantages, making it suitable for standardized medicinal G. lucidum production.

In addition to solid-state fruiting body production, liquid cultivation and bioreactor systems have gradually become important components of facility-based G. lucidum production. Liquid facility-based models mainly target mycelial biomass, extracellular polysaccharides (EPS), triterpenoids, and other metabolites, and generally use shake flasks, stirred-tank reactors, or specially designed bioreactors for closed cultivation (Alsaheb et al., 2020). For example, new reactors such as the Air-L-Shaped Bioreactor (ALSB) can reduce mycelial adhesion to the reactor wall and clumping, thereby improving mass transfer efficiency and process economics (Supramani et al., 2023). In addition, two-stage liquid static cultivation and bag-type static bioreactors can be used for efficient accumulation of triterpenoids, indicating that facility-based G. lucidum cultivation has expanded from simple fruiting body production to targeted production of mycelia and functional metabolites. Overall, facility-based G. lucidum cultivation is forming a diversified system ranging from greenhouse cultivation, intelligent mushroom houses, and multi-layer bag cultivation to liquid fermentation and bioreactor production.

2.3 Development characteristics of large-scale Ganoderma lucidum production

In recent years, the G. lucidum industry has increasingly shown trends toward scaling-up, intensification, engineering, and standardization. Traditional solid-state cultivation based on logs or beds has a relatively long production cycle and high labor intensity, and is vulnerable to contamination, continuous cropping obstacles, and environmental fluctuations. These limitations make it difficult to meet global market demand for stable supply and quality consistency (Araque et al., 2020; Wu et al., 2024). As G. lucidum is increasingly used in dietary supplements, pharmaceuticals, functional foods, and health products, large-scale production requires not only higher yield, but also unified standards for active components, safety, and batch consistency. Through unified strain supply, standardized substrate preparation, centralized cultivation management, and batch-based production processes, production costs per unit can be effectively reduced, production efficiency can be improved, and product quality stability can be enhanced.

During large-scale development, facility-based cultivation and process engineering have become important technical supports. Engineering simulation studies have shown that tools such as SuperPro Designer can be used to simulate industrial-scale submerged cultivation of G. lucidum, predict equipment requirements, energy consumption, and operating costs, and evaluate the economic effects of bioreactor scale-up. For example, increasing bioreactor volume from 2 m³ to 20 m³ can significantly reduce the unit production cost of extracellular polysaccharides and other products, demonstrating the importance of economies of scale and process optimization (Araque et al., 2020). Semi-industrial and pilot-scale studies have also shown that controlled pH, optimized carbon and nitrogen sources, and appropriate reactor configurations can significantly improve the yields of extracellular polysaccharides and other metabolites (Alsaheb et al., 2020; Supramani et al., 2023). This indicates that large-scale G. lucidum production is no longer limited to traditional cultivation experience, but is gradually moving toward process design, parameter optimization, and engineering scale-up.

Standardized management is an important approach to enhancing the competitiveness of the large-scale G. lucidum industry. At present, G. lucidum production has shifted from merely pursuing yield to placing greater emphasis on quality, safety, traceability, and market standardization. Relevant reviews have indicated that industrial standardization of G. lucidum is promoting the standardization of raw material sources, processing procedures, quality control, and product circulation, while emphasizing the importance of active component content, hygiene and safety, and production consistency in industrial development. Meanwhile, facility-based cultivation, non-soil-contact systems, and clean-environment management can help reduce heavy metal contamination and the hazards of competitive fungi, thereby improving production reproducibility and quality stability (Lim et al., 2024; Wu et al., 2024). In the future, with the further application of environmental monitoring, intelligent ventilation, automatic spraying, data analysis platforms, and bioreactor technologies, large-scale G. lucidum production will place greater emphasis on precise regulation, green production, brand-oriented operation, and alignment with international quality standards, thereby promoting the industry toward high-quality development.

3 Key Points of Environmental Control in Facility-Based Ganoderma lucidum Cultivation

3.1 Temperature and humidity control during Ganoderma lucidum growth

Temperature and humidity are the core environmental factors affecting mycelial growth, primordium formation, fruiting body development, and active component accumulation in Ganoderma lucidum. During the mycelial incubation stage, G. lucidum generally shows the growth characteristics of a relatively thermophilic fungus, with the suitable temperature mostly concentrated around 25°C~30°C. Laboratory and facility-based studies have shown that G. lucidum mycelia can exhibit strong radial growth at 25°C~30°C or 30°C~35°C, although the specific suitable range may vary depending on the medium, strain, and cultivation system (Lengare et al., 2023; Al-Kaabi and Hussien, 2025). In bag and substrate cultivation, early mycelial colonization is usually suitable at 25°C~30°C, 60%~70% relative humidity, and under clean, low-light or dark conditions (Huynh and To, 2023). If the temperature is below the suitable range, mycelial growth slows significantly and the cultivation cycle is prolonged; if the temperature is too high, it may lead to mycelial aging, abnormal substrate fermentation, and an increased risk of contamination. Therefore, facility-based cultivation should dynamically adjust temperature according to different growth stages of G. lucidum and maintain environmental stability through heat preservation, cooling, shading, and ventilation measures.

Humidity management should be matched with the growth stage of G. lucidum to balance water supply, gas exchange, and contamination risk. During the mycelial incubation stage, air humidity can generally be maintained at 60%~70% to prevent substrate water loss and mycelial shrinkage. After entering the stages of primordium formation and fruiting body development, air humidity should be appropriately increased to promote primordium differentiation, pileus expansion, and tissue fullness of fruiting bodies. Studies have shown that in controllable mushroom houses, when the temperature is maintained at approximately 28°C, a relative humidity close to 90% during the fruiting stage is conducive to pileus expansion and fruiting body weight gain (Liu et al., 2024). Stage-specific humidity management, such as maintaining 60%~0% humidity during early colonization, increasing it to 80%~90% during active fruiting body development, and then reducing it back to 60%~70% before harvest, has also been associated with relatively high polysaccharide content around day 75 of cultivation on lignocellulosic substrates (Huynh and To, 2023). This indicates that humidity regulation affects not only the morphogenesis of G. lucidum, but also the accumulation of functional components such as polysaccharides.

Climatic conditions vary greatly across seasons and regions, placing higher requirements on facility-based G. lucidum management. Field and semi-controlled system studies have shown that, compared with cooler environments, warmer cultivation sites can shorten the cultivation cycle and accelerate primordium and fruiting body development, indicating that temperature differences significantly affect the developmental process of G. lucidum (Figure 1) (Bijalwan et al., 2021). During high-temperature summers, shading, wet-pad cooling, air circulation, and automatic spraying should be used to reduce the internal temperature of the facility. In winter, heat preservation and heating management should be strengthened to prevent low temperatures from inhibiting mycelial activity. In recent years, low-cost IoT systems have been applied to continuous temperature and humidity monitoring in indoor G. lucidum cultivation, enabling real-time tracking of environmental fluctuations and helping maintain facility conditions within suitable ranges, thereby supporting both yield improvement and quality compliance (Nguyen et al., 2023). Therefore, temperature and humidity control should shift from traditional experience-based management to precise regulation based on real-time data.

Figure 1 Different growth stages of Ganoderma lucidum (Adopted from Bijalwan et al., 2021)

3.2 Ventilation and Light Regulation During Fruiting Body Formation of Ganoderma lucidum

During fruiting body formation, ventilation management mainly regulates CO₂ concentration and oxygen supply within the facility, and is an important factor ensuring normal fruiting body morphogenesis. G. lucidum continuously respires during growth. If air exchange in the facility is insufficient, CO₂ concentration increases, which may easily lead to elongated stipes, smaller pilei, incomplete pileus expansion, and even atypical fruiting body structures. In log and bag cultivation systems, sufficient ventilation during the fruiting stage is usually as important as moisture management and helps maintain suitable air and substrate conditions (Bijalwan et al., 2021). Studies on CO₂ regulation have shown that elevated CO₂ levels, such as approximately 0.3% or below 5%, can induce antler-like or other atypical fruiting structures and are accompanied by changes in metabolite profiles and increased accumulation of phenolics, polysaccharides, triterpenoids, and steroids; however, they may also inhibit normal pileus formation and cell-division-related processes (Fang et al., 2025). Therefore, in facility-based production, the ventilation strategy should be determined according to product goals. If standard pileus-type G. lucidum is produced, air exchange should be strengthened; if special morphotypes such as antler-type G. lucidum are produced, CO₂ accumulation may be regulated within a controllable range.

Ventilation regulation can also improve the humid and hot environment inside the facility, reduce localized water accumulation and stuffiness, and lower the risk of pathogen proliferation. In actual production, the ventilation demand is relatively low during the mycelial incubation stage, whereas air exchange should be strengthened during primordium differentiation and pileus development to ensure oxygen supply and reduce CO₂ concentration. However, excessive ventilation may reduce air humidity, causing fruiting body margins to crack or growth to stagnate. Therefore, ventilation management should be synchronized with humidity regulation. Natural air exchange, mechanical ventilation, circulation fans, and CO₂ monitoring systems can be used to achieve a dynamic balance between ventilation and moisture retention. Fang et al. (2025) pointed out that facility-based cultivation requires adjustable ventilation and CO₂ monitoring equipment so that management strategies can be switched according to the production goals of normal fruiting bodies or special morphotypes.

Light is another key factor affecting primordium induction, fruiting body differentiation, and quality formation in G. lucidum. G. lucidum exhibits obvious photosensitivity during the fruiting stage. Appropriate light can promote primordium formation and pileus differentiation, whereas darkness is more favorable for vegetative mycelial growth. Although G. lucidum is not a strong light-demanding fungus, a certain intensity of scattered light is beneficial for pigment formation, pileus expansion, and appearance quality improvement. Studies in environmentally controllable “mushroom factories” have shown that, under conditions of approximately 28°C, about 90% humidity, and a 12 h light/12 h dark cycle, continuous or intermittent green light promotes stipe length, pileus diameter, fruiting body weight, crude protein, triterpenoid and polysaccharide contents, and extracellular enzyme activity more effectively than white light. In contrast, continuous or intermittent red light delays or even completely inhibits fruiting body differentiation and is unfavorable for production (Liu et al., 2024). Earlier studies also found that light alone can induce primordium formation on agar medium, while the combined effects of light and ventilation can induce atypical fruiting structures with basidia and basidiospores, indicating a significant interaction between light and aeration. Therefore, facility-based G. lucidum production should rationally design light spectrum, light intensity, photoperiod, and ventilation regimes according to different growth stages and product goals.

3.3 Clean cultivation environment and pest and disease control in Ganoderma lucidum cultivation

Facility-based G. lucidum cultivation has a relatively long production cycle, and the substrates are rich in organic nutrients. In addition, high-temperature and high-humidity conditions can easily promote the proliferation of antagonistic fungi and pests. Therefore, clean-environment management is an important foundation for ensuring high-yield and high-quality production. Before production begins, cultivation rooms, inoculation rooms, cultivation racks, spraying systems, and production tools should be thoroughly cleaned and disinfected to reduce pathogen residues. Workers entering the production area should also undergo sanitation procedures to prevent external contamination sources from entering the cultivation environment. Studies have shown that in intensive G. lucidum facilities, colonies of Trichoderma and Mucor on the soil surface can increase significantly during cultivation, by up to 9-15 times; these fungi show strong antagonistic effects against G. lucidum in vitro and are closely associated with cultivation obstacles and failure of fruiting body development (Tong et al., 2020; Wang et al., 2022).

Contamination by competing microorganisms is one of the most common and serious problems in G. lucidum production. Contaminants such as Trichoderma, Penicillium, Mucor, and Neurospora compete with G. lucidum mycelia for nutrients and space and may inhibit G. lucidum growth through metabolic products, causing bag failure or even contamination of entire batches in severe cases. Continuous cultivation in the same soil can also lead to increases in organic matter and nitrogen levels, promoting the proliferation of competitive fungi and pathogens such as Xylogone ganodermophthora. This pathogen can reduce G. lucidum growth by more than 50%, leading to weak growth, aggravated disease, and yield decline (Tong et al., 2020; Huang et al., 2024). Therefore, thorough sterilization, clean substrate preparation, timely removal of contaminated bags, soil or substrate rotation, and avoidance of excessive nutrient accumulation are basic measures for disease control in facility-based G. lucidum production.

Pest and disease control should follow the principle of “prevention first and integrated control,” combining physical control, biological control, environmental regulation, and necessary chemical intervention. Studies have shown that waterlogging treatment of cultivation soil can significantly reduce the numbers of Trichoderma and Mucor colonies, restore normal fruiting, and recover spore yield in soils where unsanitized conditions completely failed to produce spores, indicating the feasibility of physical sanitation treatment in large-scale application (Tong et al., 2020). For continuous cropping obstacles caused by X. ganodermophthora, low-concentration imazalil can effectively suppress the pathogen and promote G. lucidum growth when physical measures such as temperature and light are ineffective (Huang et al., 2024). At the same time, Trichoderma, as a typical biocontrol fungus, can exert antagonistic effects through competition, mycoparasitism, and antibiotic production. This indicates both its risk as a contaminant in G. lucidum cultivation and its potential value in ecological control of other plant diseases. Therefore, facility-based G. lucidum production should establish an integrated control system combining contaminant flora monitoring, preventive sanitation management, timely disinfection, and scientifically selected control agents to improve cultivation stability and product safety.

4 Key Technical Procedures for Standardized Ganoderma lucidum Production

4.1 Ganoderma lucidum strain selection and substrate standardization

Standardized production of Ganoderma lucidum first depends on the selection of superior strains and the optimization of substrate formulations. Strain quality directly determines mycelial growth rate, fruiting stability, fruiting body yield, and the accumulation of active components such as polysaccharides and triterpenoids. Therefore, in facility-based production, priority should be given to strains with stable genetic traits, strong stress resistance, good adaptability to facility environments, high yield, and superior quality. Comparative studies have shown that different G. lucidum strains cultivated on the same woody substrates differ significantly in fruiting probability, yield, and β-glucan content, indicating a clear compatibility relationship between strains and substrates (Cortina-Escribano et al., 2020). Meanwhile, strain origin and genetic background also affect the nutritional quality and metabolic characteristics of fruiting bodies. For example, strains from different sources may vary in protein, carbohydrates, fatty acids, mineral elements, and antioxidant components (Sadiq et al., 2021). Therefore, G. lucidum strain selection should focus not only on mycelial vigor and yield, but also on active component profiles, contamination resistance, and regional adaptability.

Substrate is an important nutritional basis for G. lucidum growth. Its standardization focuses on establishing stable and reproducible raw material formulations, and controlling particle size, moisture content, carbon-to-nitrogen ratio, and mineral element levels. G. lucidum prefers substrates rich in cellulose and lignin, with low nitrogen content, a high C/N ratio, and a relatively high cellulose/lignin ratio. Yield is generally positively correlated with cellulose and lignin contents, but negatively correlated with excessive nitrogen content. At present, sawdust, cottonseed hulls, wheat bran, corn flour, and gypsum are commonly used as basic raw materials in production. Sawdust provides the main carbon source, while wheat bran and oilseed meal supplements provide nitrogen and mineral elements. Meanwhile, wheat straw, bean straw, corn cobs, peanut stems, cottonseed hulls, and other agricultural residues can also partially replace wood resources, reducing production costs while maintaining or improving biological efficiency, protein content, and mineral content (Roy et al., 2015; Mahmoud et al., 2026). For example, adding 40% corn cobs to sawdust can increase fruiting body protein and key mineral contents without reducing yield or biological efficiency; a 1:1 mixture of wheat straw and sawdust supplemented with 20% wheat bran can achieve favorable biological efficiency in seasonal cultivation.

Moisture content and pH are also key indicators that need to be controlled in standardized substrate production. In general, maintaining substrate moisture content at 60%~65% is suitable. Excessive moisture reduces aeration and increases the risk of contamination, whereas insufficient moisture affects mycelial expansion and substrate conversion efficiency. Substrate pH is usually maintained in the slightly acidic to near-neutral range, which is conducive to normal mycelial growth of G. lucidum. In actual production, standardized procedures should be established for raw material inspection, crushing and screening, formula weighing, water addition and mixing, and pH adjustment, so as to avoid batch-to-batch differences caused by moldy or contaminated raw materials, unstable proportions, or moisture fluctuations. Overall, substrate standardization should comprehensively consider raw material availability, cost control, nutritional balance, strain compatibility, and product quality targets, and should be validated through production practice to form stable formulation systems (Roy et al., 2015; Mahmoud et al., 2026).

4.2 Ganoderma lucidum cultivation bag preparation, sterilization, and inoculation management

Cultivation bag preparation is a core operational step in standardized G. lucidum production and serves as the basic unit linking substrate standardization with facility-based environmental control. A typical cultivation bag preparation process includes raw material drying, crushing, screening, proportional mixing, moisture adjustment, pre-fermentation when necessary, bag filling, and compaction. In production, raw materials such as sawdust, cottonseed hulls, peanut stems, bean stalks, wheat bran, gypsum, lime, and MgSO₄ are commonly used to prepare substrates, which are then filled into polypropylene or plastic cultivation bags to form bags with uniform specifications and density (Roy et al., 2015; Atila, 2020). During bag filling, the tightness should be reasonably controlled according to substrate characteristics. The substrate should have good aeration while maintaining an appropriate degree of compactness to facilitate uniform mycelial expansion. If the bag is too loose, the substrate may collapse and moisture distribution may become uneven; if it is too compact, gas exchange will be restricted and mycelial growth rate will decrease.

Sterilization is an important technical measure for preventing contamination and improving the qualified rate of cultivation bags. Bagged substrates are usually sterilized by high-pressure moist heat or atmospheric-pressure sterilization, in which high temperature is used to kill contaminating spores and harmful microorganisms in the substrate. Some processes also involve disinfection after pre-fermentation to improve substrate stability and reduce contamination risk (Roy et al., 2015). During sterilization, temperature, pressure, and time should be strictly controlled to ensure complete sterilization both inside and outside the substrate. If sterilization is insufficient, contaminants such as Trichoderma and Penicillium may rapidly proliferate during spawn run; if sterilization is excessive, the nutritional structure of the substrate may be damaged, affecting G. lucidum mycelial activity. Therefore, establishing standardized sterilization parameters and batch recording systems is an important guarantee for stable production.

Inoculation management should be carried out under aseptic or clean conditions to reduce contamination by external microorganisms. The inoculation room should be disinfected with ultraviolet light in advance, air purification should be performed, and the workbench should be cleaned. Operators should wear clean work clothes and strictly follow sanitation protocols. During inoculation, mycelial blocks or grain spawn can be inserted into the central hole or multiple inoculation points of the cultivation bag, followed by sealing and incubation. The spawn medium itself also needs to be optimized. For example, wheat grain or corn grain spawn can support faster colonization and stronger mycelial growth, providing vigorous inoculum for production bags. After inoculation, cultivation bags should be transferred to the incubation room and cultivated under dark or low-light conditions, at approximately 25°C~30°C, relatively high humidity, and suitable CO₂ levels until the mycelia fully colonize the substrate (Roy et al., 2015). During spawn run, contamination, water accumulation, mycelial degeneration, and uneven growth should be checked regularly, and abnormal bags should be removed or treated in time to ensure batch production stability.

4.3 Fruiting management, harvesting, and processing standards of Ganoderma lucidum

The fruiting stage of G. lucidum is a critical period determining fruiting body quality, yield, and commercial consistency. After the mycelia fully colonize the cultivation bags, primordium formation should be induced through bag opening, humidity enhancement, ventilation, appropriate light, and necessary physiological stimulation. Studies have shown that, in some boreal strains, cold stimulation at 5°C can improve primordium formation and fruiting probability in indoor systems, especially for G. lucidum cultivation on Populus tremula and Betula substrates (Table 1) (Cortina-Escribano et al., 2020).

Table 1 Effect of the factors strain, wood and treatment on the yield of G. lucidum (Adopted from Cortina-Escribano et al., 2020 Table caption: The interaction between variables is shown with an asterisk between variables names (Adopted from Cortina-Escribano et al., 2020)

In general bag cultivation, the fruiting stage requires 25°C~32°C, 85%~95% relative humidity, moderate light, and good aeration to support fruiting body formation and pileus expansion (Roy et al., 2015). In seasonal mushroom-house cultivation, approximately 30 °C± 1°C, high CO₂, and 90% relative humidity can be maintained during spawn run and primordium formation; during fruiting, CO₂ should be reduced and temperature and humidity adjusted to promote normal fruiting body development. This indicates that fruiting management should be regulated stage by stage according to strain, substrate, facility type, and product goals.

During fruiting body development, temperature, humidity, ventilation, and light should be managed synergistically. Low air humidity can easily cause cracking at the margin of G. lucidum fruiting bodies, thin pilei, and growth stagnation, whereas excessive humidity increases the risk of disease and contamination. Insufficient ventilation can cause CO₂ accumulation, leading to excessive stipe elongation and poor pileus development. Appropriate scattered light is conducive to pileus color formation and morphological integrity. Facility-based cultivation usually maintains a stable fruiting environment through scheduled spraying, mechanical ventilation, shading and light regulation, and environmental monitoring systems. In wild-simulated cultivation models, a relatively natural but still controllable fruiting environment can be created by removing the bags, covering beds with soil, managing dim light under the forest canopy, and biologically disinfecting ridges and surrounding areas (Dong et al., 2019). Potted and ornamental G. lucidum production places greater emphasis on fine control of temperature, humidity, and light to form fruiting bodies with ornamental value (Atila, 2020).

Mature G. lucidum should be harvested in a timely manner to avoid over-maturation, decline in appearance quality, and loss of active components. Harvesting is usually carried out when the pileus has fully expanded, the white growth ring at the margin disappears, spores begin to release, or the fruiting body reaches the target maturity stage. After harvesting, impurities should be removed promptly, and slicing, drying, grading, and packaging should be conducted according to product use. Drying methods significantly affect G. lucidum quality, as different drying methods can alter moisture content, rehydration capacity, phenolics, flavonoids, triterpenoids, polysaccharides, and antioxidant capacity (Atila, 2020; Naseri et al., 2023). Studies have shown that microwave drying, vacuum drying, freeze drying, hot-air drying, and infrared drying each have their own advantages. Among them, vacuum drying performs well in retaining active substances, bioavailability, and sensory quality, while infrared drying can shorten drying time and reduce energy consumption. At the same time, HPLC fingerprints of multiple ganoderic acids and related markers can be used for quality control of dried fruiting body and spore products (Yeung et al., 2021). Therefore, standardized processing should clearly define harvest maturity, slice specifications, drying method, drying temperature, final moisture content, and chemical marker ranges to improve product quality stability and market competitiveness.

5 Quality Control System in Facility-Based Ganoderma lucidum Production

5.1 Whole-process record management of Ganoderma lucidum production

In facility-based production of Ganoderma lucidum, establishing a whole-process record management system covering pre-production, mid-production, and post-production stages is an important foundation for standardized production, quality control, and product traceability. Because the production cycle of G. lucidum is relatively long and involves multiple links, including germplasm resource management, strain cultivation, substrate preparation, sterilization and inoculation, mycelial incubation, fruiting management, harvesting and processing, packaging and storage, and product circulation, any deviation in any link may affect the final product quality. Wu et al. (2024) proposed that the G. lucidum industrial chain can be divided into pre-production links, including germplasm resource management and production area planning; mid-production links, including cultivation, harvesting, processing, and in-process quality control; and post-production links, including packaging, storage, circulation, quality evaluation, and traceability (Figure 2). Therefore, in facility-based production, information such as strain identity, raw material batches, substrate formulation, sterilization parameters, inoculation time, environmental control data, harvest stage, and processing conditions should be systematically recorded to ensure that each batch of products is traceable in source, controllable in process, and accountable in responsibility.

Figure 2 A three-dimensional framework model of the standard system for the entire industry chain of GL (Adopted from Wu et al., 2024)

Environmental data recording is one of the core components of whole-process management. Environmental parameters such as temperature, humidity, light intensity, CO₂ concentration, ventilation status, and cleanliness directly affect mycelial growth, fruiting body formation, and active component accumulation in G. lucidum. Under facility-based cultivation conditions, intelligent sensors and automatic monitoring systems can be used to collect key environmental parameters in real time and establish dynamic production records. When abnormal growth, pest and disease occurrence, or quality fluctuations occur during production, environmental records and production operation records can be used for cause tracing, enabling timely adjustment of management measures. Previous studies have indicated that ganoderic acids and other constituents show growth-stage-specific differences; therefore, accurate recording of growth stage and harvest time is an important prerequisite for ensuring stable product chemical profiles (Zhang et al., 2023).

Whole-process records should also be integrated with modern analytical authentication technologies to improve the scientific reliability and credibility of the traceability system. Elemental and stable isotope fingerprints, such as δ¹³C, δ¹⁵N, δ¹⁸O, C%, and N%, combined with chemometric methods, can be used to distinguish the geographical origin, cultivar, and growth stage of G. lucidum, providing technical support for verifying origin information and production records (Zhang et al., 2023; Wadood et al., 2023). HPLC fingerprints and multivariate analysis can also be used to identify sample origin, cultivation method, and consistency with declared information. Therefore, facility-based G. lucidum production should integrate batch records, environmental monitoring, quality testing, and analytical authentication technologies on the basis of unified record templates, thereby forming a closed-loop quality assurance system.

5.2 Quality evaluation and grading standards for Ganoderma lucidum products

Quality evaluation of G. lucidum products is an important basis for assessing the level of facility-based production and the market value of products. Current market requirements for G. lucidum products have gradually shifted from simply pursuing yield to focusing on quality, safety, active component content, and batch consistency. Therefore, quality evaluation should include not only sensory indicators such as appearance, pileus size, color, integrity, degree of drying, impurity content, and mildew status, but also active component indicators such as polysaccharides, triterpenoids, and ganoderic acids. Polysaccharides and triterpenoids are widely recognized as the main active components of G. lucidum and have been adopted as important quality control indicators in pharmacopoeias and related standards (Wu et al., 2023; Yang et al., 2025). Therefore, facility-based production should establish a comprehensive evaluation system composed of appearance grade, physicochemical indicators, active components, and safety indicators.

Modern detection technologies provide an objective basis for grading G. lucidum products. HPLC-DAD can quantitatively detect multiple major ganoderic acids and related triterpenoids in fruiting body and spore extracts, accurately reflecting the level of active components in products and serving as an important tool for grading based on content levels and fingerprint similarity (Yeung et al., 2021). In addition, HPLC and HPLC-MS fingerprints combined with chemometric methods such as PCA, HCA, and PLS-DA can be used to distinguish G. lucidum samples from different origins, cultivation methods, and quality grades, and to identify key discriminant markers (Chen et al., 2008). This indicates that quality evaluation of G. lucidum products should shift from single-index testing toward a combined model integrating multiple markers, whole fingerprints, and comprehensive grading.

Industrial and regulatory standards are further promoting the standardization of the G. lucidum product quality evaluation system. Wu et al. (2024) noted that national and industry standards have incorporated indicators such as ganoderic acids, total triterpenoids, and spore powder wall-breaking rate into quality evaluation systems, and relevant standards are gradually covering product processing, circulation, and traceability. Meanwhile, related reviews emphasize that G. lucidum polysaccharides and triterpenoids have complex structures and obvious batch-to-batch variation, requiring multi-index and whole-fingerprint evaluation methods to control quality stability (Wu et al., 2023). Notably, a U.S. market survey showed that only about 26% of commercial G. lucidum dietary supplements matched the triterpenoid and polysaccharide contents declared on their labels, suggesting that quality consistency remains a major issue for G. lucidum products. Therefore, in facility-based production, quantitative testing, fingerprint analysis, appearance grading, and whole-process records should be combined to establish grading standards such as high-quality, qualified, and unqualified categories, in order to meet the different needs of medicinal use, health food production, and functional product development.

5.3 Safety risk control in Ganoderma lucidum production

Safety risk control is an important component of the quality management system in facility-based G. lucidum production, directly affecting product safety, consumer health, and sustainable industrial development. As G. lucidum is a medicinal and edible product, its production process requires high standards for raw material safety, cultivation environment cleanliness, and processing hygiene. During substrate selection, the sources of sawdust, cottonseed hulls, wheat bran, agricultural residues, and production water should be strictly controlled to avoid raw materials contaminated with heavy metals or pesticide residues. Studies have shown that some G. lucidum products have risks of excessive heavy metal contents. For example, among commercial Lingzhi products from China and Southeast Asia, approximately one-third of samples exceeded the allowable limit for arsenic, and more than half exceeded the allowable limit for cadmium (Wong et al., 2020). Another Beijing market survey found that some G. lucidum samples, mainly from wild sources, had excessive mercury levels, indicating that wild and cultivated sources may present different safety risks.

Although facility-based production can reduce some safety risks by controlling substrates, water sources, and production environments, regular testing and process supervision mechanisms are still necessary. Regional risk assessments based on ICP-MS have shown that the overall population risk from potentially toxic elements such as arsenic, cadmium, chromium, copper, mercury, nickel, and lead in G. lucidum is relatively low, but arsenic, chromium, and high-frequency consumers still require special attention. Pesticide residues are another important factor affecting the safety of G. lucidum products. A large-scale screening of Chinese herbal medicines showed that 29 pesticides were detected in 100 G. lucidum samples, with more than 90% of samples containing pesticide residues and 27.7% exceeding the maximum residue limits for foods, indicating that pesticide residue standards and supervision for Chinese herbal medicines still need to be strengthened (Chen et al., 2024). Broader studies on agricultural products have also shown that heavy metals and pesticides have bioaccumulative properties and potential health risks; therefore, preventive control measures should be adopted in medicinal fungal production chains (Alengebawy et al., 2021).

In addition to chemical contamination, microbial contamination and pests and diseases also affect the safety of G. lucidum production. Under high-temperature and high-humidity conditions, contaminants such as Trichoderma and Penicillium can proliferate rapidly. They not only inhibit normal G. lucidum growth, but may also produce harmful metabolites, thereby affecting product quality and safety. Therefore, facility-based production should strictly implement systems for raw material inspection, substrate sterilization, inoculation disinfection, clean cultivation environment management, and removal of contaminated bags, while minimizing the use of chemical agents to avoid residue risks. Overall, safety risk control in G. lucidum should comprehensively include the selection and regular testing of substrates and inputs, implementation of GAP/GMP requirements, batch testing for heavy metals and key pesticide residues, incorporation of safety testing results into traceability and quality grading records, and the establishment of a unified and standardized safety assurance system combining physical control, biological control, and environmental regulation ( Wong et al., 2020; Chen et al., 2024; Wu et al., 2024).

6 Application of Intelligent and Digital Technologies in Ganoderma lucidum Cultivation

6.1 Real-Time Monitoring of Environmental Data in Ganoderma lucidum Cultivation

In facility-based cultivation of Ganoderma lucidum, the stability of environmental conditions such as temperature, humidity, CO₂, light, substrate moisture, pH, and air circulation directly affects mycelial growth rate, fruiting body formation, and final product quality. Traditional manual management mainly relies on empirical judgment and is prone to delayed monitoring, discontinuous data collection, and imprecise regulation, making it difficult to meet the needs of modern large-scale and standardized G. lucidum production. With the development of Internet of Things (IoT) technology, sensor-network-based environmental monitoring systems have gradually been applied to edible mushroom and G. lucidum cultivation. These systems can continuously collect environmental data inside mushroom houses or cultivation rooms through temperature and humidity sensors, CO₂ sensors, light sensors, substrate or soil moisture sensors, and pH sensors, and upload the data to digital platforms via microcontrollers and wireless communication modules (Chong et al., 2023; Nguyen et al., 2023; Adebayo et al., 2025).

In indoor G. lucidum cultivation, low-cost IoT monitoring systems have been used for real-time monitoring of temperature and humidity changes. The study by Nguyen et al. (2023) showed that although temperature and humidity fluctuated with external weather conditions, real-time monitoring could still help maintain the cultivation environment within the required production range and ensure that G. lucidum product quality met relevant standards. Other edible mushroom production systems commonly use DHT or AM2301 temperature and humidity sensors, SCD30 CO₂ sensors, MQ-135 gas sensors, soil moisture probes, and light sensors, and upload data to cloud platforms through ESP8266, ESP32, and similar modules to enable remote access and environmental alerts (Chong et al., 2023). Calibration results between sensors and reference instruments generally show only small deviations, indicating that low-cost sensors have good practical applicability in facility-based environmental monitoring.

In addition to real-time recording, emerging technologies such as edge computing and TinyML are also improving the intelligence level of environmental monitoring systems. TinyML-based monitoring devices can integrate CO₂, temperature and humidity, light, soil moisture, soil pH, and soil temperature sensors at the microcontroller end, and directly control actuators such as humidifiers, heaters, coolers, and fans through local models, thereby reducing dependence on cloud communication and improving response speed (Adebayo et al., 2025). Edge-cloud collaborative architectures can also combine IoT sensor data with RGB images, completing data compression, anomaly detection, and real-time recognition at the edge end. This saves network resources while retaining key environmental information (Nguyen et al., 2024). Therefore, embedding real-time monitoring, edge computing, and cloud platform management into dedicated G. lucidum cultivation rooms or “mushroom factories” can provide a stable data foundation for precise environmental control and standardized production.

6.2 Construction of automated regulation systems for Ganoderma lucidum production

Automated regulation systems are an important technical foundation for achieving precise management in facility-based G. lucidum cultivation. Based on continuous environmental sensing, automated systems can connect data such as temperature, humidity, CO₂, light, and substrate moisture with control algorithms and actuators, enabling automatic control of spraying, ventilation, heating, cooling, shading, and supplementary lighting equipment. This reduces manual regulation errors and improves production management efficiency. IoT-based climate control systems for mushroom houses can automatically turn spraying, irrigation, and ventilation equipment on or off according to feedback from temperature, humidity, and CO₂ sensors, keeping temperature within a suitable biological range, such as 25°C~33°C, and maintaining humidity within an appropriate range, while also supporting remote monitoring and management (Chong et al., 2023).

In terms of temperature and humidity regulation, automated systems can dynamically adjust the internal environment of facilities through heating equipment, cooling fans, wet-pad systems, automatic spraying devices, and circulation fans. When humidity is lower than the set value, the system can automatically activate spraying or humidification equipment; when humidity is too high or CO₂ concentration is elevated, ventilation equipment can be automatically activated for air exchange. Small-scale cultivation systems based on NodeMCU microcontrollers and cloud platforms such as Blynk can already monitor temperature, humidity, light, and substrate moisture simultaneously, and automatically control fans, humidifiers, heaters, and lighting equipment according to user-defined thresholds (Chong et al., 2023). Such systems show that automated control is not only applicable to large G. lucidum factories, but can also be extended to medium- and small-scale facility-based production scenarios.

More advanced automated systems further introduce fuzzy logic, predictive control, machine learning, and industrial automation technologies to improve regulation precision and resource-use efficiency. Intelligent substrate management frameworks can use multi-source sensor data, including soil moisture, temperature and humidity, light, and pest movement, and apply fuzzy rules to determine watering, ventilation, light regulation, and pest alert needs, thereby improving environmental stability (Irwanto et al., 2024). Sustainable mushroom house systems using IoT sensors, Raspberry Pi-based actuator control, fuzzy logic, and predictive analytics can control temperature within ±0.5°C and humidity within ±2%, while reducing water use by approximately 30% and energy consumption by 20% compared with manual systems (Kavaliauskas et al., 2022). In factory-based edible mushroom production, electrical automation combined with fuzzy PID control can reduce contamination rates and significantly increase fresh mushroom yield. These studies provide a technical pathway for the transition of G. lucidum production from experience-based manual regulation to integrated, algorithm-driven, and precise environmental control.

6.3 Production Data Analysis and Intelligent Management of Ganoderma lucidum

The development of digital technologies is gradually transforming G. lucidum production from experience-based management to data-driven and model-based management. In modern facility-based cultivation, sensor networks and automated systems continuously generate large amounts of time-series data, including temperature, humidity, CO₂, light, water flow, equipment status, cultivation bag batches, growth cycles, contamination rates, yield, and quality testing results. Through cloud platforms or IoT platforms such as NETPIE 2020, ThingSpeak, and Blynk, these data can be centrally stored, visualized as trends, used for remote regulation, and analyzed for resource consumption, helping producers understand environmental changes and management effects at different production stages (Kavaliauskas et al., 2022). Cloud-connected systems can also export data in CSV format for further calibration, performance evaluation, and optimization of control rules (Chong et al., 2023).

Data analysis technologies can help producers promptly identify abnormal problems in G. lucidum production and provide decision-making support for process optimization. For example, by comparing spawn-running time, fruiting cycle, contamination rate, yield, and active component contents among different batches of G. lucidum, producers can analyze deficiencies in environmental control, substrate formulation, ventilation regime, or spraying frequency, and thereby optimize production processes. Edge-cloud architectures further support real-time anomaly detection and quality control. AI models can be used to identify abnormal environmental patterns or abnormal growth conditions while reducing bandwidth and storage costs (Nguyen et al., 2024). With the long-term accumulation of production data, G. lucidum production can gradually establish correlation models linking “environmental parameters-growth performance-yield and quality,” providing a scientific basis for facility design, parameter setting, and standardized production.

From a broader perspective, artificial intelligence and big data technologies are reshaping edible mushroom cultivation and can be further extended to factory-based G. lucidum production. Reviews have noted that combining IoT-based environmental monitoring with machine learning models such as LSTM can predict diurnal temperature and humidity changes and guide proactive regulation of ventilation, humidification, and heating, thereby achieving a more stable microclimate and reducing energy consumption (Jacob et al., 2025). Adaptive control models based on fuzzy logic, reinforcement learning, and sensor data can also be used to optimize resource allocation and maintain stable growth conditions in commercial farms (Chen et al., 2022). Computer vision and machine learning technologies can further be applied to species identification, quality grading, digital phenotyping, automatic yield estimation, and harvest timing judgment. Therefore, intelligent management systems for G. lucidum should build integrated data pipelines from cultivation rooms to central databases, and use AI for yield prediction, anomaly detection, process modeling, quality grading, and decision support, so as to continuously improve production efficiency, quality stability, and industrial sustainability.

7 Development Pathways for Facility-Based and Standardized Ganoderma lucidum Production

7.1 Construction of regionalized Ganoderma lucidum cultivation technology systems

China has a vast territory, and different regions vary significantly in climatic conditions, ecological environments, raw material resources, and industrial foundations. Therefore, facility-based production of Ganoderma lucidum cannot completely adopt a single unified model; instead, appropriate cultivation technology systems should be established according to regional characteristics. Relevant studies have pointed out that the standardization of the G. lucidum industry should cover the whole industrial chain, including pre-production planning, germplasm management, cultivation production, postharvest processing, circulation and sales, and quality traceability. At the same time, differences in resource conditions and ecological environments among regions should be fully considered to form regionally adaptive technical routes (Wu et al., 2024; Thakur et al., 2024). For example, in southern regions with high temperature and humidity, emphasis should be placed on cooling, ventilation, and disease control to reduce contamination by competing microorganisms and heat stress under humid conditions. In northern regions, winter heat preservation, air humidity regulation, and improvement of facility insulation should be strengthened to ensure the stability of mycelial growth and fruiting body formation.

The construction of regionalized cultivation technology systems should also be optimized according to local resource conditions and market positioning. Regions rich in wood resources may focus on sawdust-based bag cultivation, while regions with favorable under-forest resources may explore facility-based wild-simulated cultivation models to balance G. lucidum quality, ecological benefits, and market differentiation. International studies have also shown that the adaptation between local strains and local substrate resources is an important pathway for improving regional production efficiency. A Finnish study on laccate Ganoderma found that local wild strains could be successfully domesticated on regional wood by-products, among which Populus tremula and Betula substrates significantly improved fruiting probability, yield, and β-glucan content, whereas Pinus sylvestris was unsuitable for fruiting body production. Research from Pakistan showed that indigenous G. lucidum strains could be efficiently cultivated indoors using locally available lignocellulosic wastes, with a sawdust and wheat straw combination producing higher yields and more fruiting flushes while maintaining a mineral composition close to that of wild basidiomata (Ghafoor et al., 2024). These studies indicate that regionalized cultivation can promote the high-value utilization of local biomass resources while providing a localized basis for standardized facility-based production.

In addition, the construction of regionalized technology systems requires strengthened collaboration among research institutions, production enterprises, cultivation bases, and market participants. By conducting regional trials on strain screening, substrate formulation optimization, environmental parameter regulation, and pest and disease control, production parameters and technical protocols suitable for different regions can gradually be established. In some parts of China, permanent sand-bed greenhouse models have been promoted, in which plastic membranes are used to separate substrates from soil in order to reduce the risks of heavy metal and pesticide residues and support stable production that meets medicinal material quality requirements. At the international level, standards such as ISO 21315, Traditional Chinese Medicine-Ganoderma lucidum Fruiting Body, as well as relevant standards for cold storage, transportation, and quality evaluation, also provide a basis for embedding international standards into regionalized production systems (Wu et al., 2024). Therefore, regionalized G. lucidum cultivation technology systems should form synergy among local resource utilization, domestication of local strains, facility structure optimization, and alignment with standardized protocols, thereby promoting coordinated development of the G. lucidum industry.

7.2 Breeding and promotion of superior Ganoderma lucidum varieties

Superior varieties are the foundation for improving the yield, quality, and stress resistance of G. lucidum, and they are also the core component of facility-based standardized production. The genotype of G. lucidum strains directly affects mycelial growth rate, fruiting stability, contamination resistance, spore yield, and the accumulation of active components such as polysaccharides and triterpenoids. With the continuous increase in market demand, G. lucidum production has gradually shifted from simply pursuing high yield to multi-objective breeding that considers high yield, superior quality, disease resistance, stable active component accumulation, and adaptability to facility-based management. Studies have shown that the mutant strain UV119 obtained through ultraviolet mutagenesis had a basidiospore yield 19.27% higher than that of the parent strain and 20.56% higher than that of the major commercial cultivar “Longzhi No. 1,” while its fruiting body yield and contamination resistance were also improved (Tang et al., 2023). Another radiation-induced mutant strain, XZ-2, showed 9.7%-30.1% higher fruiting body yields and 20.2%-45.7% higher basidiospore yields than control varieties across multiple locations, while also exhibiting stable annual yields and relatively high polysaccharide and triterpenoid contents. These results indicate that superior variety breeding can directly improve the stability and industrial value of facility-based G. lucidum production.

In the breeding process, traditional breeding techniques should be combined with modern biotechnologies to systematically screen and evaluate G. lucidum genetic resources. Through strain isolation, basidiospore-derived monokaryon crossing, mutation breeding, molecular-assisted screening, and metabolomic evaluation, new G. lucidum varieties adapted to different regions and cultivation models can be developed. For example, the hybrid strain H-23, generated by mating basidiospore-derived monokaryons from two commercial cultivars, had relatively high polysaccharide and triterpenoid contents as well as desirable biological efficiency, indicating that sexual crossing can integrate multiple favorable traits into new strains (Liu et al., 2017). In recent years, molecular breeding and high-throughput screening have further expanded breeding pathways. A new variety, GL_V2, obtained through ultraviolet mutagenesis, showed a 1.4-fold increase in total triterpenoid content compared with widely cultivated varieties and exhibited stronger antioxidant activity (Pan et al., 2025). Atmospheric room-temperature plasma mutagenesis combined with microdroplet cultivation technology can rapidly screen mutants with 17%~26% higher biomass and up to 32% higher triterpenoid production in liquid cultivation, with stability maintained over multiple generations (Feng et al., 2024). In addition, new strains obtained by combining genetic engineering with mono-mono crossing can significantly increase the contents of specific ganoderic acids, squalene, and lanosterol, providing new ideas for targeted improvement of active components in G. lucidum (Zhou et al., 2024).

The promotion and application of superior varieties are equally critical. At present, some regions still face problems such as non-standardized strain sources, strain degeneration, variety mixing, and insufficient coverage of superior strains, which affect production stability and product consistency. Therefore, standardized strain propagation systems and superior variety promotion mechanisms should be established, and variety registration, regional adaptability trials, strain quality testing, and market supervision should be strengthened. Research on developing G. lucidum as a “next-generation cell factory” also suggests that synthetic biology and pathway engineering may be used in the future to optimize the production of target products such as proteins, polysaccharides, and triterpenoids, although the complexity of regulatory mechanisms and the efficiency of genetic manipulation remain bottlenecks to be overcome. Therefore, the promotion of superior varieties should be integrated with facility-based production standards, regional environmental parameters, product uses, and quality evaluation systems, so that breeding outputs can truly serve the needs of large-scale, standardized, and high-value production (Liu et al., 2017; Tang et al., 2023; Feng et al., 2024; Wu et al., 2024).

7.3 Construction of Ganoderma lucidum branding and product traceability systems

As consumers increasingly focus on the quality, safety, and authenticity of health products, the G. lucidum industry is gradually shifting from traditional raw material sales toward branding, standardization, and high-value-added operations. Brand building can not only increase the added value of G. lucidum products, but also enhance consumer recognition and market competitiveness. A recent review on the standardization of the G. lucidum industry proposed that a whole-chain standard system should be established, covering germplasm and strain-type standards, cultivation and harvesting standards, processing standards, circulation standards, quality evaluation and traceability standards, and management service standards. Within this framework, G. lucidum branding should rely on verifiable quality attributes, such as the contents of marker components including polysaccharides, triterpenoids, and ganoderic acid A, compliance with national standards for spore powder harvesting and processing, and alignment with the testing, packaging, storage, and transportation requirements of ISO 21315 for G. lucidum fruiting bodies (Wu et al., 2024). Only by building brand value on stable quality and verifiable quality indicators can conceptual promotion and the impact of low-quality products on the market be avoided.

The construction of product traceability systems is an important component of modern agricultural quality management and brand trust formation. In G. lucidum production, QR codes, batch codes, digital management platforms, and even blockchain technology can be used to record whole-process information, including strain source, regional cultivation conditions, substrate formulation, facility environmental parameters, harvesting and processing, quality testing, warehousing and logistics, and sales circulation. This enables product source inquiry, production process tracking, and quality responsibility traceability. Stable isotope and chemometric methods can further strengthen the scientific basis of traceability systems. Studies have shown that by analyzing C, H, O, and N stable isotope ratios as well as elemental C and N contents, and combining PCA, OPLS-DA, PLS, and FLDA methods, the geographical origin and growth stage of G. lucidum can be effectively distinguished, with some external validation accuracies reaching 100% (Zhang et al., 2023). These techniques can be used to independently verify origin and growth-stage information on product labels and can be integrated with digital traceability platforms to improve brand credibility.

In addition, the construction of branding and traceability systems helps standardize market order and reduce the circulation of low-quality or counterfeit products. Studies on commercial products have shown that only about 26% of G. lucidum dietary supplements in the U.S. market matched the triterpenoid and polysaccharide contents declared on their labels, suggesting that quality inconsistency can significantly damage brand reputation and consumer trust. HPLC fingerprints targeting 11 major ganoderic acids and related triterpenoids have been validated for quality control and batch comparison of fruiting body and spore products, supporting brand quality claims based on multi-marker chemical profiles (Yeung et al., 2021). Therefore, G. lucidum brand building should not remain limited to regional culture and packaging promotion, but should be deeply integrated with standardized production, quality testing, fingerprint profiling, origin authentication, and digital traceability. In the future, with the development of e-commerce, smart agriculture, and international trade, G. lucidum brand building and digital traceability systems will become further integrated, providing support for high-quality industrial development and international market expansion.

8 Conclusion

With the continuous expansion of market demand for Ganoderma lucidum and consumers’ increasing requirements for product quality, safety, and stability, traditional cultivation methods that rely on natural environments and experience-based management can no longer meet the needs of modern industrial development. Facility-based cultivation effectively improves production stability and resource-use efficiency through precise regulation of environmental factors such as temperature, humidity, light, and ventilation. It promotes the transformation of the G. lucidum industry from dispersed and extensive production toward large-scale, intensive, and modernized development. Therefore, facility-based cultivation has become an important development pathway for the transformation and upgrading of the G. lucidum industry and is of great significance for enhancing overall industrial competitiveness and promoting sustainable development.

Mycelial growth and fruiting body formation in G. lucidum are highly sensitive to environmental conditions. Factors such as temperature and humidity, air circulation, light intensity, and environmental cleanliness directly affect yield, appearance quality, and the accumulation of active components. In facility-based production, establishing a scientific environmental control system combined with real-time monitoring and dynamic regulation technologies can effectively reduce the adverse effects of environmental fluctuations, lower the incidence of pests, diseases, and microbial contamination, and improve product uniformity and quality stability. Therefore, precise environmental control is not only an important foundation for efficient G. lucidum production, but also a key factor in enhancing its medicinal value and market competitiveness.

Standardized production and intelligent management are important trends in the future development of the G. lucidum industry. By establishing unified technical procedures for strain management, substrate preparation, sterilization and inoculation, fruiting management, harvesting, and processing, the standardization level of the production process can be improved, enabling stable product quality and whole-process traceability. Meanwhile, the application of intelligent technologies such as IoT, automated control, and big data analysis is gradually transforming G. lucidum production from traditional experience-based management toward precision, digitalization, and smart management. In the future, with the continuous improvement of standardization systems and the sustained development of smart agricultural technologies, the G. lucidum industry will further achieve high-quality, green, and modernized development.

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.

References

Alsaheb R.A., Zjeh K.A., Malek R.A., Abdullah J.O., Baz A.A., Deeb E.A., Dailin D.J., Hanapi S.Z., Sukmawati D., and El Enshasy H.A., 2020, Bioprocess optimization for exopolysaccharides production by Ganoderma lucidum in semi-industrial scale, Recent Patents on Food, Nutrition and Agriculture, 11(3): 211-218.

https://doi.org/10.2174/2212798411666200316153148

Adebayo S., Aworinde H., Olufemi O., Osueke C., Adeniyi A., and Aroba O., 2025, Understanding mushroom farm environment using TinyML-based monitoring devices, Environmental Research Communications, 7(4): 045014.

https://doi.org/10.1088/2515-7620/adc5cd

Akçay Ç., Arslan R., and Ceylan F., 2025, Valorization of various lignocellulosic wastes to Ganoderma lucidum (Curtis) P. Karst (Reishi Mushroom) cultivation and their FT-IR assessments, PLOS ONE, 20(7): e0328732.

https://doi.org/10.1371/journal.pone.0328732

Alengebawy A., Abdelkhalek S.T., Qureshi S.R., and Wang M.Q., 2021, Heavy metals and pesticides toxicity in agricultural soil and plants: Ecological risks and human health implications, Toxics, 9(3): 42.

https://doi.org/10.3390/toxics9030042

Al-Kaabi A., and Hussien A., 2025, The environmental requirements of three species of Reishi mushroom, Ganoderma spp., IOP Conference Series: Earth and Environmental Science, 1487(1): 012020.

https://doi.org/10.1088/1755-1315/1487/1/012020

Araque J., Niño L., and Gelves G., 2020, Industrial scale bioprocess simulation for Ganoderma lucidum production using SuperPro Designer, Journal of Physics: Conference Series, 1655(1): 012077.

https://doi.org/10.1088/1742-6596/1655/1/012077

Atila F., 2020, Comparative study on the mycelial growth and yield of Ganoderma lucidum (Curt.: Fr.) Karst. on different lignocellulosic wastes, Acta Ecologica Sinica, 40(2): 153-157.

https://doi.org/10.1016/j.chnaes.2018.11.007

Bijalwan A., Bahuguna K., Vasishth A., Singh A., Chaudhary S., Dongariyal A., Thakur T.K., Kaushik S., Ansari M.J., Alfarraj S., Alharbi S.A., Skalický M., and Brestič M., 2021, Growth performance of Ganoderma lucidum using billet method in Garhwal Himalaya, India, Saudi Journal of Biological Sciences, 28(5): 2709-2717.

https://doi.org/10.1016/j.sjbs.2021.03.030

Chen F., Zhang T., Xiao P., Shao L., Zhang X., Wang L., Ren X., Qin C., and Jiao Y., 2024, Occurrence and health risk of pesticide residues in Chinese herbal medicines from Shandong Province, China, Environmental Science and Pollution Research, 31: 25940-25951.

https://doi.org/10.1007/s11356-024-32693-4

Chen L., Qian L., Zhang X., Li J.Z., Zhang Z.J., and Chen X.M., 2022, Research progress on indoor environment of mushroom factory, International Journal of Agricultural and Biological Engineering, 15(1): 25-32.

https://doi.org/10.25165/j.ijabe.20221501.6872

Chen Y., Zhu S.B., Xie M.Y., Nie S.P., Liu W., Li C., Gong X.F., and Wang Y.X., 2008, Quality control and original discrimination of Ganoderma lucidum based on high-performance liquid chromatographic fingerprints and combined chemometrics methods, Analytica Chimica Acta, 623(2): 146-156.

https://doi.org/10.1016/j.aca.2008.06.018

Chong J.W.R., Chew K.W., Peter A.P., Ting H.Y., and Show P.L., 2023, Internet of Things (IoT)-based environmental monitoring and control system for home-based mushroom cultivation, Biosensors, 13(1): 98.

https://doi.org/10.3390/bios13010098

Cortina-Escribano M., Pihlava J.M., Miina J., Veteli P., Linnakoski R., and Vanhanen H., 2020, Effect of strain, wood substrate and cold treatment on the yield and β-glucan content of Ganoderma lucidum fruiting bodies, Molecules, 25(20): 4732.

https://doi.org/10.3390/molecules25204732

Dong Q., He D., Ni X., Zhou H., and Yang H., 2019, Comparative study on phenolic compounds, triterpenoids, and antioxidant activity of Ganoderma lucidum affected by different drying methods, Journal of Food Measurement and Characterization, 13: 3198-3205.

https://doi.org/10.1007/s11694-019-00242-0

Fang T., Chen L., Yao H., Li Y., Liu G., Wu S., Lan J., and Chen X., 2026, Combined transcriptomic and metabolomic analyses of the response of Ganoderma lucidum to elevated CO₂, Journal of Fungi, 12(1): 5.

https://doi.org/10.3390/jof12010005

Feng J., Tang C., Liu Y., Tang C., and Zhang J., 2024, Enhancing high-efficiency breeding and microbial microdroplet cultivation techniques for Ganoderma lucidum, World Journal of Microbiology and Biotechnology, 40(7): 225.

https://doi.org/10.1007/s11274-024-04011-0

Ghafoor A., Rana A.A., and Niazi A.R., 2024, Systematic characterisation, mineral analysis, and valorisation of lignocellulosic waste into sustainable indigenous Ganoderma lucidum production from Pakistan, New Zealand Journal of Botany, 63: 2589-2605.

https://doi.org/10.1080/0028825x.2024.2353191

Huang Q., Lu Q., Cao F., Li X., Wu X., Sun H., Zhang J., and Fu J., 2024, Study on the use of Imazalil to continuous cropping obstacle of Ganoderma lucidum caused by Xylogone ganodermophthora, Chemical and Biological Technologies in Agriculture, 11(1): 57.

https://doi.org/10.1186/s40538-024-00584-y

Huynh T.T., and To N.T., 2023, Factors affect on the total polysaccharide content in Ganoderma lucidum (Leyss ex Fr.) Karst, Ganodermataceae, Tạp chí Y Dược học Cần Thơ, 30(5): 140-148.

https://doi.org/10.58490/ctump.2023i5.653

Irwanto F., Hasan U., Lays E., De La Croix N., Mukanyiligira D., Sibomana L., and Ahmad T., 2024, IoT and fuzzy logic integration for improved substrate environment management in mushroom cultivation, Smart Agricultural Technology, 9: 100427.

https://doi.org/10.1016/j.atech.2024.100427

Jacob M., Xu A., Qian K., Qi Z., Li X., and Zhang B., 2025, Artificial intelligence in edible mushroom cultivation, breeding, and classification: A comprehensive review, Journal of Fungi, 11(11): 758.

https://doi.org/10.3390/jof11110758

Kavaliauskas Ž., Šajev I., Gecevičius G., and Čapas V., 2022, Intelligent control of mushroom growing conditions using an electronic system for monitoring and maintaining environmental parameters, Applied Sciences, 12(24): 13040.

https://doi.org/10.3390/app122413040

Lengare K., Ingle Y., Sable Y., and Pol V., 2023, Nutritional and physiological requirements for the improvement of growth of Ganoderma lucidum, Journal of Plant Disease Sciences, 18(2): 80-85.

https://doi.org/10.48165/jpds.2023.1802.02

Lim T.F., Ilham Z., and Wan-Mohtar W.A.A.Q.I., 2024, Production of ganodiesel from the biomass of Ganoderma lucidum in air-L-shaped bioreactor (ALSB), Heliyon, 10(15): e35170.

https://doi.org/10.1016/j.heliyon.2024.e35170

Liu S.R., Ke B.R., Zhang W.R., Liu X.R., and Wu X.P., 2017, Breeding of new Ganoderma lucidum strains simultaneously rich in polysaccharides and triterpenes by mating basidiospore-derived monokaryons of two commercial cultivars, Scientia Horticulturae, 216: 58-65.

https://doi.org/10.1016/j.scienta.2016.12.016

Liu Y., Luo Y., Guo W., Zhang X., Zheng W., and Chen X., 2024, Study on the effects of different light supply modes on the development and extracellular enzyme activity of Ganoderma lucidum, Agriculture, 14(6): 835.

https://doi.org/10.3390/agriculture14060835

Magday J.C., 2014, Optimization of mycelial growth and cultivation of fruiting body of Philippine wild strain of Ganoderma lucidum, Current Research in Environmental and Applied Mycology, 4(2): 162-172.

https://doi.org/10.5943/cream/4/2/4

Mahmoud A.T., Abed I.A., Almohammedi O.H., and Thakir A.F., 2026, The effect of Ganoderma strain type, growth substrate, and nitrogen supplementation on the concentration of certain biochemical parameters in fruiting bodies, Anbar Journal of Agricultural Sciences, 24(1): 25-34.

https://doi.org/10.32649/ajas.2025.189068

Naseri M., Movagharnejad K., and Nanvakenari S., 2023, Comparison of the dried properties of Ganoderma lucidum produced by the convective dryer and infrared dryer, Scientific Reports, 13(1): 12636.

https://doi.org/10.1038/s41598-023-39883-z

Nguyen H., Shin D., Jung W., Kim T., and Lee D., 2024, An integrated IoT sensor-camera system toward leveraging edge computing for smart greenhouse mushroom cultivation, Agriculture, 14(3): 489.

https://doi.org/10.3390/agriculture14030489

Nguyen Q., Huynh H., Dao T., and Kwon H., 2023, Application of Internet of Things based monitoring system for indoor Ganoderma lucidum cultivation, The International Journal of Advanced Smart Convergence, 12(2): 153-158.

Pan Z., Lin J., Luo G., Cheng W., Li Y., and Wu C., 2025, Unveiling triterpenoid superiority in a newly developed Ganoderma lucidum variety through untargeted metabolomics approach, Frontiers in Nutrition, 12: 1541162.

https://doi.org/10.3389/fnut.2025.1541162

Roy S., Jahan M.A.A., Das K.K., Munshi S.K., and Noor R., 2015, Artificial cultivation of Ganoderma lucidum (Reishi medicinal mushroom) using different sawdusts as substrates, American Journal of BioScience, 3(5): 178-182.

https://doi.org/10.11648/j.ajbio.20150305.13

Sadiq N.B., Ryu D.H., Cho J., Lee A., Song D., Dorjsembe B., Kim J., Jung J.H., Nho C.W., Hamayun M., Yang S.H., and Kim H.Y., 2021, Postharvest drying techniques regulate secondary metabolites and anti-neuroinflammatory activities of Ganoderma lucidum, Molecules, 26(15): 4484.

https://doi.org/10.3390/molecules26154484

Supramani S., Rejab N.A., Ilham Z., Ahmad R., Show P.L., Ibrahim M.F., and Wan-Mohtar W.A.A.Q.I., 2023, Performance of biomass and exopolysaccharide production from the medicinal mushroom Ganoderma lucidum in a new fabricated Air-L-Shaped Bioreactor (ALSB), Processes, 11(3): 670.

https://doi.org/10.3390/pr11030670

Tang C., Tan Y., Zhang J., Zhou S., Honda Y., and Zhang H., 2023, A novel strain breeding of Ganoderma lucidum UV119 (Agaricomycetes) with high spores yield and strong resistant ability to other microbes’ invasions, Foods, 12(3): 465.

https://doi.org/10.3390/foods12030465

Thakur P., Khanal S., Tapwal A., Kumar D., Verma R.K., Chauhan P., and Sharma N., 2024, Exploring Ganoderma lucidum: Morphology, cultivation and market potential, World Journal of Microbiology and Biotechnology, 40(11): 369.

https://doi.org/10.1007/s11274-024-04180-y

Tong X., Jiang H., Liang Y., Rao Y., Mei L., and Wang Y., 2020, Waterlogging reduces soil colonization by antagonistic fungi and restores production in Ganoderma lucidum continuous cultivation, Crop Protection, 137: 105314.

https://doi.org/10.1016/j.cropro.2020.105314

Wadood S.A., Nie J., Li Z., Li C., Zhang N., Rogers K.M., Zhang Y., and Yuan Y., 2023, A preliminary elemental and isotopic investigation to develop authentication tools for Chinese Ganoderma lucidum, Food Control, 150: 109888.

https://doi.org/10.1016/j.foodcont.2023.109888

Wang Y.J., Zeng L.Z., Wu J.Y., Jiang H., and Mei L., 2022, Diversity and effects of competitive Trichoderma species in Ganoderma lucidum-cultivated soils, Frontiers in Microbiology, 13: 1067822.

https://doi.org/10.3389/fmicb.2022.1067822

Wong T.H., Lee F.S.C., Hu G.H., Chang L.P., Wang X.H., and Fu P.P., 2007, A survey of heavy metal and organochlorine pesticide contaminations in commercial Lingzhi products, Journal of Food and Drug Analysis, 15(4): 16.

https://doi.org/10.38212/2224-6614.2401

Wu P., Zhang C., Yin Y., Zhang X., Li Q., Yuan L., Sun Y., Zhou S., Ying S., and Wu J., 2024, Bioactivities and industrial standardization status of Ganoderma lucidum: a comprehensive review, Heliyon, 10(16): e36987.

https://doi.org/10.1016/j.heliyon.2024.e36987

Wu S.J., Zhang S.Y., Peng B., Tan D.C., Wu M.Y., Wei J.C., Wang Y.T., and Luo H., 2023, Ganoderma lucidum: a comprehensive review of phytochemistry, efficacy, safety and clinical study, Food Science and Human Wellness, 13(2): 568-596.

https://doi.org/10.26599/fshw.2022.9250051

Yang M.H., Qin X.M., and Liu X.J., 2025, A review of polysaccharides from Ganoderma lucidum: preparation methods, structural characteristics, bioactivities, structure-activity relationships and potential applications, International Journal of Biological Macromolecules, 303: 140645.

https://doi.org/10.1016/j.ijbiomac.2025.140645

Yeung S.Y., Chen Q.L., Yu Y.B., Zhou B.S., Wu W., Li X., Huang Y., and Wang Z.J., 2021, Quality evaluation of commercial products of Ganoderma lucidum made from its fruiting body and spore, Acta Chromatographica, 34(1):100-113.

https://doi.org/10.1556/1326.2020.00825

Zhang Y., Jiang K., Chen S., Wang L., Zhang X., Xu W., Yam M.F., Wu C., Xu W., and Lin Y., 2023, Quality control of Ganoderma lucidum by using C, H, O, and N stable isotopes and C and N contents for geographical traceability, Frontiers in Plant Science, 14: 1234729.

https://doi.org/10.3389/fpls.2023.1234729

Zhou D.X., Kong X.M., Huang X.M., Li N., Feng N., and Xu J.W., 2024, Breeding a new Ganoderma lucidum strain with increased contents of individual ganoderic acids by mono-mono crossing of genetically modified monokaryons, Frontiers in Microbiology, 15: 1410368.

https://doi.org/10.3389/fmicb.2024.1410368