Effects of Different Radiation-Based Treatments on the Quality of Edible Mushrooms: A Systematic Review

Abstract

Radiation-based treatments have emerged as important environmental and postharvest regulatory tools for improving the quality of edible mushrooms. Visible light, ultraviolet (UV) radiation, gamma irradiation, and pulsed-light treatments influence mushroom growth, morphogenesis, nutrient accumulation, antioxidant capacity, and storage performance through distinct physiological and molecular mechanisms. However, current findings remain fragmented, and a comprehensive synthesis of their regulatory effects and underlying mechanisms is lacking. This systematic review was conducted following the PRISMA 2020 framework. A structured literature search was performed in the Web of Science, PubMed, and CNKI databases. After screening and eligibility assessment, 111 studies were included in the qualitative synthesis. The available evidence indicates that radiation-based treatments exert stage-dependent and species-specific effects on edible mushrooms. Visible light primarily regulates morphogenesis through photoreceptor-mediated signaling pathways, whereas UV radiation promotes vitamin D₂ biosynthesis and antioxidant accumulation through photochemical and reactive oxygen species (ROS)-related mechanisms. Gamma irradiation and pulsed-light treatments are mainly applied during postharvest handling to suppress microbial contamination, delay browning and senescence, and extend shelf life. Based on the available evidence, a unified mechanistic framework linking signal perception, ROS regulation, transcriptional reprogramming, metabolic responses, and quality formation is proposed. Despite these advances, substantial challenges remain, including limited mechanistic understanding, insufficient integration of multi-omics evidence, lack of standardized treatment protocols, and difficulties in industrial-scale implementation. Future research should focus on multi-radiation synergistic strategies, precision environmental regulation, and intelligent cultivation systems. Overall, this review provides a comprehensive synthesis of current evidence regarding radiation-mediated quality regulation in edible mushrooms and offers a theoretical basis for optimizing mushroom production and developing sustainable postharvest preservation technologies.

1. Introduction

Because of their high nutritional, economic, and ecological value, edible mushrooms have emerged as a rapidly expanding sector within the global food and agricultural industries. As of 2025, the global edible mushroom market is valued at approximately USD 73–76 billion and is projected to sustain steady growth, with a compound annual growth rate of 5.5–10% [1]. China remains the leading producer, consumer, and exporter worldwide, accounting for over 70% of global production for several consecutive years. In 2024, total output reached 44.199 million tons, with an industrial value exceeding RMB 420.7 billion [2].

Edible mushrooms are characterized by a rich nutritional profile, including carbohydrates, essential amino acids, proteins, unsaturated fatty acids, vitamins (particularly B-group vitamins and vitamin D₂), minerals, and phenolic compounds [3]. Moreover, their short production cycle, efficient resource utilization, and relatively low reliance on arable land confer distinct advantages in diversifying food resources, optimizing dietary structures, enhancing food security, and promoting sustainable agricultural development.

The growth and development of edible mushrooms are governed by multiple environmental factors. While temperature, moisture, and gas composition provide the fundamental conditions for physiological processes, light acts as a critical environmental signal closely linked to morphogenesis and metabolic regulation [4,5]. The regulatory reach of radiation signaling extends from shaping shoot architecture in horticultural crops to orchestrating stomatal movements in plants, demonstrating its central role in both developmental and physiological adaptations—a principle that may also inform our understanding of photoregulation in edible mushroom development [6,7]. To perceive and respond to changes in the optical radiation environment, fungi have evolved specialized photoreceptor systems. Among fungal photoreceptors, the White Collar Complex (WCC), composed of WC-1 and WC-2, represents the best-characterized blue-light sensing system and plays a central role in primordium initiation and fruiting-body development [8,9]. Red and far-red light responses are thought to be mediated primarily by phytochrome-like proteins, which participate in developmental regulation and secondary metabolism [8]. The biological role of light is also development-stage dependent. During vegetative growth, many mushroom species preferentially colonize substrates under dark conditions, whereas light serves as an essential environmental cue for the transition from mycelial growth to primordium formation and fruiting-body development [10]. In addition to acting as developmental signals, certain light treatments, particularly ultraviolet radiation, may function as environmental stressors that activate antioxidant defense responses and stimulate the accumulation of phenolic compounds, flavonoids, and other protective metabolites [11]. UV exposure can also promote the photochemical conversion of ergosterol into vitamin D₂, thereby enhancing nutritional value [11,12,13].

Recent studies on light-quality regulation have predominantly focused on individual light sources or limited physiological responses. For instance, UV-B irradiation of dried shiitake mushroom powder significantly increased vitamin D₂, total phenolics, flavonoids, β-glucan content, and antioxidant activity after 2 h of treatment [14]. Similarly, gamma irradiation at 2 kGy modified carbohydrate composition, fatty acid profiles, and antioxidant parameters in L. edodes and Agaricus spp.; however, these studies primarily addressed postharvest preservation rather than integrated light-regulation mechanisms [15]. Pulsed light treatments have largely been investigated for microbial inactivation and texture preservation in fresh-cut mushrooms [16], whereas transcriptomic analyses of P. ostreatus under blue light revealed alterations in glycolysis and the pentose phosphate pathway, without addressing broader regulatory networks [17].

Although a recent review by Feng et al. comprehensively summarized the effects of visible light, ultraviolet radiation, gamma irradiation, and pulsed light on the quality of preharvest and postharvest edible mushrooms, several important knowledge gaps remain [18]. In particular, current reviews have provided only limited discussion of photoreceptor-mediated signaling networks, ROS-centered regulatory mechanisms, transcriptional reprogramming, and the crosstalk among different light-response pathways. In addition, emerging perspectives from systems biology, including transcriptomics, proteomics, and metabolomics, as well as issues related to industrial scalability, intelligent light-environment control, and regulatory frameworks, have not been systematically integrated. Therefore, a more comprehensive framework linking light perception, signal transduction, metabolic reprogramming, and quality formation is still needed to support the precise application of light-based technologies in edible mushrooms.

Therefore, this systematic review provides a comprehensive synthesis of the effects of different radiation-based treatments on the growth, development, nutritional quality, antioxidant properties, and postharvest preservation of edible mushrooms. Particular attention is given to the mechanisms linking light perception, reactive oxygen species (ROS)-mediated signaling, metabolic reprogramming, and quality formation. Furthermore, both preharvest and postharvest applications are discussed from an integrated perspective to evaluate the distinct and complementary roles of visible light, ultraviolet radiation, gamma irradiation, and pulsed light. Finally, current challenges related to mechanistic understanding, parameter standardization, industrial scalability, and regulatory constraints are critically analyzed, and future directions involving multi-light synergistic regulation, systems biology approaches, and intelligent light-environment management are proposed. This review aims to provide a theoretical foundation for precision quality regulation and the sustainable development of edible mushroom production and preservation technologies. The integration of transcriptomics, proteomics, and metabolomics is expected to provide new insights into the molecular basis of radiation-mediated quality regulation in edible mushrooms.

2. Materials and Methods

2.1. Study Identification

This systematic review was conducted in accordance with the Preferred Reporting Items for Systematic Reviews and Meta-Analyses (PRISMA 2020) guidelines (Supplementary Data S1) [18]. The completed PRISMA 2020 checklist is available as Table S1 in the Supplementary Materials. A comprehensive literature search was performed using three electronic databases, including Web of Science Core Collection (WoS; Clarivate Analytics, Philadelphia, PA, USA), PubMed (National Center for Biotechnology Information, Bethesda, MD, USA), and China National Knowledge Infrastructure (CNKI; Beijing, China), to identify studies investigating the effects of light-based treatments on the quality of edible mushrooms.

Eligible studies were required to report original experimental data evaluating the effects of visible light, ultraviolet radiation, gamma irradiation, pulsed light, or other light-related treatments on edible mushroom quality attributes. The following Boolean search strategy was employed: (“edible mushroom*” OR mushroom*) AND (“blue light” OR “red light” OR UV OR ultraviolet OR “gamma irradiation” OR “pulsed light”) AND (quality OR nutrition* OR antioxidant* OR “bioactive compound*” OR polysaccharide* OR phenolic* OR vitamin* OR ergosterol OR “vitamin D” OR postharvest OR “shelf life”).

Database-specific search syntax was adapted according to the indexing requirements of each platform. The complete search strategies for all databases are provided in Supplementary Data S2. The final database search was performed in March 2025.

2.2. Study Selection Process

Following the removal of duplicate records using EndNote (version 20.2.1, Clarivate Analytics, Philadelphia, PA, USA), two independent reviewers screened the titles and abstracts of all retrieved records for relevance. Full-text articles deemed potentially eligible were subsequently assessed independently by the reviewers to determine their suitability for inclusion in the final review. Any disagreements were resolved through discussion and consensus.

The reference lists of all included studies and relevant review articles were also manually screened to identify additional eligible publications.

The screening and selection process is summarized in Figure 1.

A total of 1695 records were identified through database searching, including 1069 records from Web of Science, 430 from PubMed, and 196 from CNKI. After the removal of 442 duplicate records, 1253 publications remained for title and abstract screening. Subsequently, 1074 records were excluded due to irrelevance to the review topic. The remaining 179 articles underwent full-text assessment for eligibility. Following full-text evaluation, 68 studies were excluded because they lacked original experimental data (n = 23), were not directly related to the effects of light-based treatments on edible mushroom quality (n = 19), or investigated non-edible mushroom species (n = 30). Ultimately, 107 studies met all eligibility criteria and were included in the final qualitative synthesis. The screening and selection process is summarized in Figure 1.

2.3. Eligibility Criteria

Studies were considered eligible if they met all of the following criteria: (1) The investigated organism was an edible mushroom species; (2) The study evaluated the effects of visible light, ultraviolet radiation, gamma irradiation, pulsed light, or other light-based treatments; (3) At least one quality-related parameter was reported, including nutritional composition, bioactive compounds, antioxidant activity, sensory quality, flavor characteristics, or postharvest preservation performance; (4) Original experimental data were presented; (5) Full-text articles were available in English or Chinese. Studies were excluded if: (1) The investigated organism was not an edible mushroom species; (2) The study did not involve light-related treatments; (3) Quality-related outcomes were not evaluated; (4) The publication was a review article, conference abstract, editorial, book chapter, patent, or other publication lacking primary experimental data; (5) The full text was unavailable or insufficient methodological information was provided.

2.4. Data Extraction

Data were independently extracted by two reviewers using a standardized extraction protocol, with discrepancies resolved through discussion and consensus. The following information was extracted from each study: mushroom species; cultivation stage or postharvest stage; light treatment category; light wavelength or spectral composition; irradiation intensity; exposure duration; total dose or fluence (when available); measured quality indicators; physiological and biochemical responses; proposed molecular mechanisms; principal findings and conclusions. To facilitate comparison among studies employing different irradiation modalities, reported light-treatment parameters were standardized whenever sufficient information was available.

2.5. Data Analysis

Data analysis was conducted as a descriptive systematic review. Due to substantial heterogeneity among studies regarding mushroom species, cultivation conditions, light sources, irradiation parameters, and quality evaluation methods, a formal meta-analysis was not considered appropriate.

Therefore, the included studies were synthesized qualitatively. Extracted data were categorized according to treatment type, including visible-light regulation, ultraviolet radiation, gamma irradiation, and pulsed-light treatment. Within each category, evidence was further summarized according to its effects on nutritional quality, bioactive compounds, antioxidant capacity, sensory attributes, postharvest preservation, and molecular regulatory mechanisms.

Comparisons were subsequently performed to identify common response patterns, species-specific differences, and potential mechanistic pathways underlying light-mediated quality regulation in edible mushrooms.

2.6. Risk-of-Bias Assessment

Given that the majority of included studies consisted of controlled laboratory experiments investigating physiological, biochemical, and postharvest responses of edible mushrooms to different light treatments, a formal quantitative risk-of-bias assessment tool commonly applied to clinical studies was not considered appropriate.

Instead, methodological quality was evaluated qualitatively based on experimental design, treatment description, reproducibility of irradiation parameters, reporting completeness, and outcome assessment methods. Particular attention was paid to the reporting of light wavelength, irradiation intensity, exposure duration, environmental conditions, and replication strategies, as these factors are critical for the interpretation and reproducibility of light-treatment studies.

Potential sources of bias, including incomplete reporting of irradiation parameters, insufficient experimental replication, and variability in quality assessment methodologies, were considered during evidence synthesis and interpretation.

3. Results and Discussion: Mechanisms Underlying Radiation-Induced Quality Regulation in Edible Mushrooms

3.1. Common Types of Optical Radiation and Irradiation Treatments in Edible Mushrooms

The optical radiation environment constitutes a critical exogenous factor affecting the growth, development, and quality formation of edible mushrooms. Among the various light-related parameters, light quality—referring to radiation with distinct wavelengths or spectral compositions—plays a central role in regulating physiological processes, including mycelial growth, fruiting body differentiation, and metabolite accumulation, through specific perception and signaling pathways. In contrast, light treatment is an experimental and engineering-based operational concept that extends beyond the physical characteristics of a given light quality to include the temporal dynamics of photon delivery, such as continuous illumination, intermittent photoperiods, microsecond-scale pulsed light, and dose-dependent pulsed fluxes of ionizing radiation. In brief, light quality determines the specificity of fungal photoreceptor responses, whereas light treatment governs the dynamics of energy delivery and the resulting macroscopic physiological effects.

In edible mushroom research and production, the principal radiation-based treatments include visible light, ultraviolet (UV) radiation, gamma irradiation, and pulsed light (PL). Visible light and UV radiation belong to non-ionizing electromagnetic radiation, whereas gamma rays are classified as ionizing radiation [19]. In contrast, pulsed light represents a distinct treatment modality characterized by high-intensity, short-duration light pulses, rather than a continuous spectral range.

From an application perspective, visible light is primarily employed during cultivation to regulate growth and morphogenesis, whereas UV radiation, gamma irradiation, and pulsed light are more frequently applied in postharvest stages to maintain product quality [14,17].

To clarify the stage-specific use of light-based technologies, the applications of different Radiation-Based Treatments can be divided into preharvest regulation during cultivation and postharvest interventions during processing and storage. Visible LED-based illumination is mainly used to regulate mycelial growth, primordium initiation, fruiting-body development, morphology, and yield, whereas UV radiation, pulsed light, and gamma irradiation are more frequently applied after harvest to enhance vitamin D₂ formation, suppress browning, reduce microbial load, and extend shelf life. For details, see Figure 2.

3.2. Light Perception Systems in Edible Mushrooms

Edible mushrooms are capable of sensing variations in the external optical radiation environment and transducing radiation signals of different wavelengths and energy levels into intracellular regulatory cues through specialized sensory systems. This process constitutes the initial step in light-quality–mediated regulation of growth, development, and quality formation.

Within the visible spectrum, blue light is the most extensively studied signal and is primarily perceived by the White Collar Complex (WCC), composed of WC-1 and WC-2. WC-1 acts as both a photoreceptor and a transcriptional regulator, enabling the direct modulation of downstream gene expression upon light perception [20]. In addition to WCC, cryptochromes (Cry) have also been implicated in blue-light responses in certain edible mushroom species, where they are associated with morphogenesis and stress adaptation [21].

Red-light responses in fungi are often associated with phytochrome-like proteins, but their functional roles in edible mushroom species remain insufficiently characterized and require direct genetic validation [22].

3.3. Radiation Signal Transduction and Cellular Response Mechanisms

In edible mushrooms, the effects of different Radiation-Based Treatments on growth, development, and quality formation ultimately rely on the conversion of light energy into biochemical signals and the subsequent activation of cellular response networks. From a mechanistic perspective, these effects can be broadly categorized into three partially overlapping pathways: photoreceptor-mediated signal transduction, non-specific photochemical reactions, and radiation-mediated microbial inactivation.

These mechanisms are associated with distinct treatment modalities while also exhibiting functional interplay, collectively modulating metabolic processes, quality formation, and postharvest stability in edible mushrooms. The interplay among photoreceptor-mediated signaling, non-specific photochemical reactions, and antimicrobial effects is conceptually summarized in Figure 3, which illustrates how these mechanisms collectively orchestrate the quality-forming processes in edible mushrooms.

3.3.1. Photoreceptor-Mediated Signal Transduction

Under visible light, edible mushrooms perceive external signals through photoreceptors and other photosensitive molecules, which are subsequently transduced into intracellular regulatory cues that modulate gene expression and metabolic processes [23,24,25]. Among these signals, blue light plays a particularly important role in coordinating morphogenesis and secondary metabolism. In Ganoderma lingzhi, WC-2 functions as a key component of the blue-light response network. Genetic evidence has shown that disruption of WC-2 alters the expression of multiple genes associated with triterpenoid metabolism and significantly reduces ganoderic acid accumulation, indicating that WC-2 participates in the transcriptional regulation of light-responsive metabolic pathways rather than acting solely as a photoreceptor [20].

Recent studies suggest that photoreceptors do not function independently but are embedded within broader gene regulatory networks (GRNs) that connect light perception with downstream metabolic responses. In addition to the White Collar Complex (WCC), several classes of transcription factors have been implicated in fungal light responses, including velvet-family proteins (VeA, VelB, and LaeA), Zn(II)₂Cys₆ transcription factors, and developmental regulators belonging to the AT-hook, HMG-box, homeobox, and forkhead-box families [26]. These regulators are thought to act as molecular bridges between light perception and the expression of genes involved in substrate utilization, cellular differentiation, and secondary metabolite biosynthesis.

Evidence supporting this regulatory framework has been reported in several edible mushrooms. In L. edodes, blue light induces the expression of transcription factors containing AT-hook, CBFD_NFYB_HMF, and HMG-box domains, which are associated with the upregulation of carbohydrate-active enzyme (CAZyme) genes, including glycoside hydrolases, glucose oxidases, and lignocellulose-degrading enzymes [27]. This transcriptional response enhances substrate degradation and nutrient acquisition during fruiting-body development. Similarly, in C. militaris, transcriptome-wide network analyses identified several light-responsive transcription factors, including Snf21-, homeobox-, and forkhead-related regulators, that are closely associated with the expression of genes involved in cordycepin and carotenoid biosynthesis [28]. These findings suggest that transcription factors constitute an essential intermediate layer linking photoreceptor activation to downstream metabolic pathways.

The transcriptional networks activated by light perception ultimately converge on metabolic pathways that support fungal growth and quality formation. Through the coordinated regulation of metabolic enzyme genes, light-responsive transcription factors remodel carbon utilization and biosynthetic processes, thereby linking environmental signals to physiological outputs. Transcriptomic analyses have shown that blue light enhances the expression of genes involved in glycolysis, the pentose phosphate pathway, lipid metabolism, nucleotide metabolism, and carbohydrate degradation, resulting in increased metabolic flux and biosynthetic capacity. In P. ostreatus and Flammulina velutipes, blue light promotes fruiting-body development by upregulating genes associated with glycolysis, the pentose phosphate pathway, and hydrolytic enzymes, leading to improved substrate utilization and nutrient acquisition [17,29]. In contrast, red light appears to regulate development through partially distinct signaling routes and may differentially influence tissue differentiation, attenuating cap-associated metabolism while promoting stipe elongation [17].

Metabolic reprogramming induced by light is further reflected in cellular energy metabolism. As metabolic demand increases during fruiting-body differentiation and secondary metabolite biosynthesis, light-responsive pathways can modulate mitochondrial function and ATP production. Previous studies have shown that light exposure affects the activities of H⁺-ATPase, Ca²⁺-ATPase, succinate dehydrogenase (SDH), and cytochrome c oxidase (CCO), while simultaneously upregulating ATP-synthesis-related genes such as LeATP6 and LeATP9 [5]. These responses contribute to improved cellular energy status and provide the energetic basis required for growth, developmental transitions, and metabolite accumulation.

Collectively, current evidence indicates that photoreceptor-mediated responses involve a hierarchical regulatory process extending from light perception to transcriptional regulation and metabolic reprogramming. However, the identities of the core transcription factors, their target genes, and the structure of species-specific light-responsive GRNs remain poorly understood in most edible mushrooms. Future studies integrating genetics, transcriptomics, proteomics, and metabolomics will be essential for elucidating how photoreceptors, transcription factors, and metabolic enzymes interact to coordinate light-regulated quality formation.

3.3.2. Non-Specific Photochemical Reactions

Unlike receptor-dependent responses, ultraviolet (UV) radiation and pulsed light primarily exert their effects through non-specific photochemical reactions that directly target intracellular biomolecules. Upon absorption of light energy, endogenous photosensitive molecules become excited and initiate energy- or electron-transfer reactions, leading to the generation of reactive oxygen species (ROS), including singlet oxygen, superoxide radicals, and hydrogen peroxide [30]. Consequently, ROS production represents one of the earliest cellular responses to UV and pulsed-light exposure.

Importantly, ROS do not function solely as damaging agents. At moderate levels, they act as signaling molecules that activate endogenous antioxidant defense systems and trigger adaptive metabolic responses [31,32]. Previous studies have shown that radiation-induced oxidative stress can stimulate the activities of superoxide dismutase (SOD), catalase (CAT), peroxidases, and glutathione-associated enzymes, thereby enhancing ROS-scavenging capacity and maintaining cellular redox homeostasis [33]. This dynamic balance between ROS generation and antioxidant defense is critical because it enables edible mushrooms to tolerate environmental stress while preserving cellular integrity.

The establishment of redox homeostasis is closely linked to metabolic reprogramming. Activation of antioxidant defense pathways is frequently accompanied by the accumulation of protective metabolites, including phenolic compounds, flavonoids, ergothioneine, and other antioxidant-related substances. For example, UV-B treatment has been associated with increased phenolic content, flavonoid accumulation, ergothioneine production, and enhanced antioxidant capacity in several edible mushroom species, suggesting that antioxidant metabolism and secondary-metabolite biosynthesis are coordinately regulated during radiation-induced stress adaptation [34,35].

When ROS generation exceeds cellular scavenging capacity, however, oxidative damage may occur. Excessive irradiation can induce bond cleavage, photodegradation, lipid peroxidation, protein oxidation, and nucleic-acid damage, ultimately affecting enzyme activity, nutrient stability, and cellular function [30]. Such photochemical reactions may also influence the formation of flavor precursors and volatile compounds. Although similar mechanisms have been reported in other food matrices, direct evidence linking these reactions to aroma formation in edible mushrooms remains limited [36].

It should be noted that the effectiveness of ultraviolet and pulsed-light treatments may also be influenced by physical and structural characteristics of fungal tissues. Cell-wall thickness, tissue architecture, and the presence of protective pigments such as melanin can affect light penetration and energy absorption. Melanin has been reported to function as a natural photoprotective barrier capable of absorbing ultraviolet radiation and reducing oxidative damage [37]. Consequently, species with different pigmentation levels or tissue structures may require substantially different irradiation doses to achieve comparable physiological responses, including vitamin D₂ accumulation and antioxidant activation.

Overall, the biological effects of UV radiation and pulsed light should not be interpreted solely as consequences of oxidative damage. Rather, they arise from a dynamic redox-regulatory network in which ROS generation, antioxidant defense, and metabolic adaptation interact to determine the final nutritional, sensory, and functional quality of edible mushrooms.

3.3.3. Radiation-Induced Microbial Inactivation

Microbial proliferation is a major contributor to postharvest quality deterioration and shelf-life reduction in fresh produce, including edible mushrooms. Various radiation-based treatments, particularly ultraviolet (UV) radiation, pulsed light, and ionizing radiation, can exert photochemical, photophysical, and oxidative effects on microbial cells, leading to damage of proteins, membranes, and nucleic acids, and ultimately suppressing microbial activity [38,39].

Ionizing radiation operates through both direct and indirect mechanisms. Direct effects involve damage to DNA and other cellular macromolecules, whereas indirect effects are mediated by reactive species generated through radiolysis, including hydroxyl radicals, hydrogen atoms, and hydrated electrons. These highly reactive intermediates further attack cellular components, resulting in irreversible cellular damage and eventual cell death [40]. Meanwhile, irradiated microorganisms may activate stress responses—such as DNA repair, antioxidant defenses, and metabolic reprogramming—in an attempt to maintain cellular homeostasis.

For non-ionizing radiation, antimicrobial efficacy is strongly influenced by wavelength, dose, and the presence of endogenous photosensitizers. UV radiation can directly induce DNA damage, primarily via the formation of pyrimidine dimers, thereby disrupting transcription, replication, and cell viability [41,42]. In addition, visible light, particularly within the blue–green spectrum, can inactivate microorganisms by exciting endogenous porphyrins and related photosensitizers, which subsequently generate reactive oxygen species (ROS) that oxidize lipids, proteins, and nucleic acids [43,44]. Furthermore, radiation-induced metabolites may also contribute indirectly to antimicrobial activity in certain systems [45].

Collectively, these mechanisms reduce microbial load and play a critical role in maintaining postharvest quality and extending the storage stability of edible mushrooms.

3.4. Radiation Signal-Driven Metabolic Regulation Framework

Regulation of edible mushroom quality by light can be regarded as a hierarchical process of light-driven metabolic reprogramming that integrates signal perception, cellular signaling, transcriptional regulation, metabolic remodeling, and downstream quality formation. Rather than operating as independent pathways, photoreceptor-mediated signaling, photochemical reactions, oxidative stress responses, and microbial inactivation constitute interconnected components of a unified regulatory network.

At the signal-input level, different Radiation-Based Treatments initiate distinct primary responses. Visible light is mainly perceived through photoreceptors, including the White Collar Complex (WC-1/WC-2), cryptochromes, and phytochrome-like proteins, whereas ultraviolet radiation (UV), gamma irradiation, and pulsed light can additionally induce direct photochemical and oxidative effects. Although these treatments differ substantially in wavelength range, photon energy, and mode of action, their downstream responses frequently converge on reactive oxygen species (ROS)-associated signaling pathways. Moderate ROS accumulation functions as an important secondary messenger involved in developmental regulation and environmental adaptation, whereas excessive ROS production may result in oxidative damage to proteins, lipids, and nucleic acids.

Following light perception or ROS induction, multiple transcriptional regulatory networks are activated. Recent transcriptomic and gene regulatory network analyses indicate that photoreceptors function not only as environmental sensors but also as upstream regulators of extensive gene-expression programs. In L. edodes, blue light induces the expression of transcription factors associated with carbohydrate-active enzyme (CAZyme) regulation and substrate degradation. Similarly, light-responsive gene regulatory networks identified in C. militaris contain homeobox, FlbB, FlbC, and C6 zinc-finger transcription factors that are associated with growth, development, cordycepin biosynthesis, and carotenoid production. These findings suggest that transcription factors represent a critical intermediate layer linking light perception to downstream metabolic outputs through gene regulatory networks (GRNs).

At the metabolic level, activation of these regulatory networks leads to extensive metabolic reprogramming. Previous studies have shown that light exposure can influence glycolysis, the pentose phosphate pathway, lipid metabolism, oxidative phosphorylation, and secondary-metabolite biosynthesis. Blue light has been reported to promote carbohydrate metabolism and energy generation in P. ostreatus, Flammulina velutipes, and L. edodes, whereas WC-mediated signaling participates in the regulation of ganoderic acid biosynthesis in Ganoderma lingzhi. These coordinated metabolic adjustments ultimately affect nutrient accumulation, developmental differentiation, and the synthesis of bioactive compounds.

The final outcome of radiation regulation is strongly influenced by cellular redox homeostasis. Importantly, these regulatory processes do not operate independently but exhibit extensive cross-talk at multiple biological levels. Photoreceptor-mediated signaling and photochemical responses frequently converge on ROS-dependent regulatory pathways, allowing environmental light cues and oxidative-stress signals to coordinately influence downstream gene expression. At the same time, ROS-mediated signaling is tightly coupled with antioxidant defense systems, creating a dynamic feedback network that balances stress adaptation and metabolic activity. In postharvest mushrooms, microbial inactivation induced by UV radiation, gamma irradiation, or pulsed light may further interact with host metabolic responses by reducing microbial-derived oxidative deterioration while simultaneously altering endogenous stress signaling. Consequently, quality formation should be viewed as the integrated outcome of photoreceptor activation, redox regulation, metabolic remodeling, and microecological changes rather than the result of any single pathway operating in isolation. Antioxidant defense systems, including superoxide dismutase (SOD), catalase (CAT), glutathione-related pathways, and phenolic antioxidants, play essential roles in balancing ROS generation and scavenging. The dynamic interaction between oxidative signaling and antioxidant responses enables edible mushrooms to maintain cellular integrity while simultaneously stimulating the accumulation of beneficial metabolites such as phenolics, flavonoids, ergothioneine, polysaccharides, carotenoids, and vitamin D₂. Consequently, radiation-induced redox regulation acts as a key bridge connecting environmental signals with nutritional quality and stress adaptation.

In addition to molecular signaling pathways, ecological adaptation may represent an important determinant of species-specific light responses in edible mushrooms. Fungi have evolved diverse photoreceptor systems and regulatory networks in response to long-term environmental selection pressures, resulting in substantial variation in light sensitivity and physiological outcomes among species [8,46]. Species occupying different ecological niches may therefore exhibit distinct responses to identical light treatments. For example, fungi naturally developing in shaded woodland habitats are generally adapted to low-light conditions, whereas species exposed to more variable environmental illumination may possess greater tolerance and responsiveness to radiation-induced stimuli. Such evolutionary adaptation may partly explain the considerable interspecific variability observed in fruiting-body development, metabolite accumulation, antioxidant activation, and vitamin D₂ biosynthesis following visible-light or ultraviolet treatments. Therefore, ecological background should be considered alongside light parameters when interpreting experimental results and designing species-specific light management strategies.

Collectively, light-regulated quality formation in edible mushrooms involves coordinated interactions among photoreceptor signaling, ROS-mediated regulation, transcriptional reprogramming, metabolic remodeling, and microecological responses. These processes ultimately determine fruiting-body development, nutrient accumulation, antioxidant capacity, flavor formation, microbial stability, browning resistance, and shelf life. Future integration of transcriptomics, proteomics, metabolomics, and phenomics will facilitate the construction of systems-level regulatory networks and enable the identification of key regulatory nodes that connect light perception with quality formation, thereby providing a stronger theoretical basis for precision light-environment management and intelligent quality-control strategies in edible mushroom production.

Overall, radiation-induced quality regulation in edible mushrooms can be conceptualized as a hierarchical process involving radiation input, signal perception or primary photochemical targets, early cellular responses, transcriptional regulatory hubs, metabolic reprogramming, and final quality outputs. This framework helps explain how distinct radiation-based treatments produce overlapping outcomes such as antioxidant activation, vitamin D₂ enrichment, browning inhibition, microbial suppression, and improved postharvest stability. To translate this conceptual framework into a testable molecular architecture, Figure 4 maps the proposed hierarchical pathway with evidence-supported connections (solid arrows) and inferred relationships (dashed arrows), delineating how each radiation modality converges on shared regulatory hubs and divergent quality outcomes.

4. Results and Discussion: Effects of Different Light Conditions on the Quality of Edible Mushrooms

4.1. Effects of Light Quality on the Growth, Development, and Appearance of Edible Mushrooms

Visible light is a key environmental factor regulating growth and morphogenesis in edible mushrooms. Mushroom development typically comprises mycelial growth, primordium initiation, and fruiting body formation, with light requirements varying markedly across species and developmental stages. Among different wavelengths, blue light has been the most extensively studied and is generally associated with enhanced cap development, biomass accumulation, and color formation.

For example, in P. ostreatus, blue light promotes fruiting body growth—particularly cap expansion—by upregulating genes involved in glycolysis and the pentose phosphate pathway [17]. Similarly, in Flammulina velutipes, blue light enhances the degradation capacity of the stipe toward cultivation substrates by increasing the expression of hydrolase-related genes, thereby supporting rapid fruiting body development [29]. In L. edodes, blue light has also been reported to increase yield and potentially shorten the production cycle [47].

By contrast, red light appears to regulate development through distinct physiological mechanisms. It may attenuate metabolic pathways associated with cap expansion while promoting stipe elongation or vegetative growth [17]. Consistent with this trend, in Pleurotus citrinopileatus, blue light increased pileus and stipe size, fresh weight, and color intensity, whereas red and far-red light induced abnormal fruiting body morphology [48]. Moreover, light responses are often stage-dependent; for instance, red LED exposure can exert differential effects on growth and metabolism depending on the developmental stage [49].

Combined light treatments frequently outperform single-wavelength illumination. In Lyophyllum decastes, red–blue combined light significantly increased fruiting body weight and stimulated extracellular enzyme activities, whereas red light alone produced inhibitory effects [14]. Similarly, in C. militaris, appropriately timed combinations of white, blue, and red light enhanced both fruiting body production and the accumulation of bioactive compounds [24,50]. These findings suggest that multispectral light regimes provide an effective strategy for optimizing both yield and developmental performance.

Across the available studies, blue light often promotes cap development and metabolic activity in several species, whereas red light may favor elongation or produce weaker/variable effects depending on species and developmental stage.

In contrast to visible light, ultraviolet (UV) radiation, gamma irradiation, and pulsed light are primarily applied during postharvest handling rather than for biomass production. At excessive doses, these treatments may inhibit growth or cause tissue damage; therefore, their practical value lies mainly in postharvest preservation and quality enhancement rather than in directly promoting development [51,52]. The differential and sometimes synergistic regulatory effects of distinct light qualities on mushroom growth and morphogenesis are systematically summarized in Figure 5, which contrasts the promoting roles of blue light in metabolic activation and fruiting-body development against the morphogenetic modulation exerted by red light on stipe elongation and cap suppression, while also highlighting the synergistic benefits of red–blue combined illumination.

4.2. Effects of Radiation-Based Treatments on the Nutritional Quality and Bioactive Compounds of Edible Mushrooms

4.2.1. Effects of Visible Light on the Nutritional Quality and Bioactive Compounds of Edible Mushrooms

Visible light is a key environmental factor influencing the nutritional quality of edible mushrooms, with different wavelengths differentially regulating metabolite accumulation. Among them, blue light plays a particularly prominent role in metabolic regulation by redirecting carbon and nitrogen fluxes. In Flammulina velutipes, for instance, blue light promotes the accumulation of lysine and low-molecular-weight carbohydrates by suppressing the biosynthesis of certain aromatic amino acids and organic acids, as well as by downregulating genes involved in lysine degradation [17]. In addition, blue light stimulates secondary metabolism, increasing ergothioneine content and antioxidant activity in L. edodes and enhancing ganoderic acid biosynthesis in Ganoderma lingzhi [11,29].

Beyond blue light, other wavelengths also regulate specific metabolic profiles. Green light has been reported to enhance mineral element accumulation in Pholiota adiposa [53]. In C. militaris, different light spectra selectively modulate the accumulation of nucleosides and sugar alcohols: pink light favors adenosine, cordycepin, and carotenoids, whereas red, green, and blue light preferentially promote cordycepin, mannitol, and adenosine, respectively [54,55]. Similarly, in Lentinus crinitus, different visible-light conditions alter carbohydrate, tocopherol, fatty acid, soluble sugar, phenolic, antioxidant, and antimicrobial profiles [55].

Composite light treatments often exhibit synergistic effects on nutritional quality. For example, red–blue combined light significantly increases crude protein and crude polysaccharide contents in Lyophyllum decastes [14]. In addition, light intensity is another critical regulatory factor. In C. militaris, appropriate photon flux density and spectral composition under LED illumination can enhance selected bioactive metabolites [56], and moderate photon flux density in other mushroom systems has likewise been associated with improved sugar content and antioxidant capacity [57]. These findings collectively indicate that both spectral composition and light dose jointly determine the final nutritional outcome.

Overall, visible light—particularly blue light—plays a central role in shaping the nutritional and bioactive composition of edible mushrooms by modulating metabolic flux distribution and gene expression. Meanwhile, other wavelengths, composite light regimes, and light intensity further fine-tune the synthesis of specific metabolites, including amino acids, carbohydrates, ergothioneine, and terpenoids. Representative examples are summarized in

Table 1. Representative effects of visible-light treatments on the nutritional quality and bioactivity of edible mushrooms.

Species	Treatment	Light Intensity/Dose (Unified Unit)	Spectral Details (Peak Wavelength)	Nutritional Quality/Bioactive Changes	Reference
P. ostreatus	White, red, blue, or green LED; 8 or 12 h/day	100 ± 5 μmol/(m2·s)	Blue: 450 nm; Red: 630 nm; Green: 525 nm; White: 6500 K	Red light (8 h/day) increased total phenols from 54.4 to 95.1 mg GAE/100 g fresh weight.	[49]
Lyophyllum decastes	Red-blue, blue, red, and far-blue vs. control	80 μmol/(m2·s)	Red: 660 nm; Blue: 450 nm; Far-red: 730 nm	Red-blue light increased crude protein, polysaccharides, and activated cellulase/laccase.	[14]
C. militaris	Red, green, and blue LED; red-blue combinations	160 μmol/(m2·s)	Red: 625 nm; Green: 525 nm; Blue: 465 nm	Biomass and yield were affected; red favored cordycepin, green favored mannitol, blue favored adenosine.	[55]
Lentinus crinitus	Blue, green, or red light	35 μmol/(m2·s)	Blue: 460 nm; Green: 530 nm; Red: 625 nm	Carbohydrates and fatty acids increased; red light produced the highest antioxidant activity.	[56]
Suillus granulatus	Red, yellow, green, blue, or white light	60 μmol/(m2·s)	Red: 620–630 nm; Yellow: 580–590 nm; Green: 520–530 nm; Blue: 450–460 nm	Blue light increased umami amino acids and 5′-nucleotides (EUC).	[58]
Lyophyllum ulmarium	Red, yellow, green, blue, or white light	15 μmol/(m2·s)	Red: 630 nm; Yellow: 590 nm; Green: 530 nm; Blue: 460 nm	Ergosterol content was significantly higher than in the fluorescent control under LED lights.	[59]
Lentinus sajor-caju	Blue, red, green, or white light	20, 40 and 80 μmol/(m2·s)	Blue: 460 nm; Red: 625 nm; Green: 530 nm	Blue light intensity influenced antioxidant properties, DPPH scavenging, and sugar content.	[60]

.

4.2.2. Effects of UV Treatment on the Nutritional Quality and Bioactive Compounds of Edible Mushrooms

Ultraviolet (UV) radiation is an important environmental factor regulating the nutritional quality and metabolite accumulation of edible mushrooms. Its biological effects are primarily reflected in the induction of bioactive compounds and the modulation of antioxidant metabolism, with UV-B generally exhibiting the strongest physiological impact.

During the fruiting body stage, the most well-established function of UV treatment is the photochemical conversion of ergosterol into vitamin D₂, thereby significantly improving nutritional value. Postharvest UV-B exposure has been widely reported to markedly increase vitamin D₂ content in several cultivated mushrooms [14,61]. In addition to vitamin D₂ biosynthesis, UV irradiation also enhances the accumulation of antioxidant-related metabolites. For instance, UV exposure increases total phenolics and flavonoids in oyster and button mushrooms [62], while low-dose UV-B further promotes antioxidant capacity and elevates levels of phenolics, flavonoids, ergothioneine, and glutamic acid in edible mushrooms [63,64]. UV-C treatment has additionally been shown to stimulate antioxidant enzyme activity and delay postharvest quality deterioration during storage [65,66,67,68,69]. The conversion efficiency is strongly influenced by ergosterol availability, tissue geometry, irradiation dose, wavelength, and post-treatment storage conditions.

Beyond the fruiting body stage, UV radiation also regulates metabolic processes during mycelial development. Available studies suggest that UV-B activates antioxidant defense systems and promotes the biosynthesis of phenolics, flavonoids, β-glucans, and vitamin D₂ in mushroom mycelia [14,64,65,70,71]. For example, in Calocybe indica and P. eryngii mycelia, appropriately timed UV-B exposure significantly enhances vitamin D₂ accumulation alongside multiple bioactive metabolites [64,70,71].

Overall, UV radiation—particularly UV-B—plays a central role in the nutritional fortification and functional quality improvement of edible mushrooms by simultaneously promoting vitamin D₂ biosynthesis, enhancing antioxidant metabolism, and orchestrating associated biochemical pathways. These effects collectively reflect a UV-mediated regulatory network linking photochemical stress responses to metabolic reprogramming and quality formation. Representative studies are summarized in

Table 2. Representative effects of UV treatments on the nutritional quality and bioactivity of edible mushrooms.

Species	UV Type and Peak Wavelength	Radiation Intensity and Total Dose (Unified)	Changes	Reference
L. edodes	UV-B (Peak: 313 nm)	Intensity: 2.34 W/m2 Total Dose: 0.14–1.26 J/cm2	Increased vitamin D2, beta-glucans, antioxidants, total phenolics, flavonoids, and amino acids.	[14]
Calocybe indica	UV-B (Peak: 310 nm)	Intensity: 5.3 W/m2 Exposure: 15–90 min	Increased beta-glucans, phenolics, and flavonoids; enhanced antioxidant capacity.	[71]
L. edodes	UV-C (Peak: 254 nm)	Intensity: 2.82 mW/cm2 Total Dose: 4.0 kJ/m2	Increased flavonoids and vitamin C; increased CAT, SOD, APX, and GR activities.	[65]
A. bisporus	UV-C (Peak: 254 nm)	Total Dose: 1.0 kJ/m2	Increased antioxidant capacity and related compounds.	[66]
A. bisporus	UV-C (Peak: 254 nm)	Total Dose: 0.5–2.0 kJ/m2	Vitamin D2 increased significantly, with the optimal response at 2.0 kJ/m2.	[67]
P. ostreatus	UV-C (Peak: 254 nm)	Intensity: 0.23 mW/cm2 (230 μW/cm2) Total Dose: 4.0 kJ/m2	No significant change in soluble solids or protein; CAT and PAL activities increased.	[68]
L. edodes + Auricularia auricula-judae	UV-C (Peak: 253.7 nm)	Total Dose: 2.0–4.0 kJ/m2	Promoted the conversion of ergosterol to vitamin D2.	[69]

.

4.2.3. Effects of γ-Irradiation on the Nutritional Quality and Bioactive Compounds of Edible Mushrooms

The effects of gamma irradiation on the nutritional quality of edible mushrooms are primarily reflected in changes in metabolite accumulation and the stability of nutritional components, with these responses being strongly dose-dependent and species-specific.

At appropriate doses, gamma irradiation does not necessarily reduce major bioactive compounds such as phenolics, organic acids, or ergosterol; in many cases, it may even promote their accumulation and enhance antioxidant capacity [15,72,73,74,75]. For instance, gamma irradiation has been reported to increase total phenolic content in Pleurotus and Agaricus species under optimized treatment conditions [76,77]. In addition, low-dose irradiation applied at the mycelial stage can enhance the subsequent accumulation of carbohydrates, proteins, phenolics, and flavonoids in fruiting bodies, suggesting a possible priming effect on metabolic pathways [5,78].

Beyond metabolite accumulation, gamma irradiation can also improve the stability of certain nutritional components during postharvest storage [79]. For example, irradiation at 1.5–2.0 kGy has been shown to slow the degradation of vitamin D₂ in stored oyster mushrooms [80]. In several wild edible mushroom species, low-dose irradiation has been reported to preserve major nutritional constituents while maintaining or even enhancing antioxidant properties [15,39,73,80,81].

Overall, gamma irradiation at appropriate doses can enhance the accumulation of phenolic and flavonoid compounds and improve the stability of key nutritional components in edible mushrooms. These effects are closely associated with dose-dependent physiological responses and species-specific metabolic sensitivity, likely involving the regulation of oxidative stress and metabolic adaptation processes. Representative studies are summarized in Table 3.

4.2.4. Effects of Pulsed Light on the Nutritional Quality and Bioactive Compounds of Edible Mushrooms

Pulsed light exerts multifaceted regulatory effects on the nutritional quality of edible mushrooms, influencing vitamin D₂ formation, antioxidant status, amino acid composition, and processing characteristics. As with other irradiation technologies, these effects are strongly dose-dependent.

Pulsed light efficiently induces the photochemical conversion of ergosterol to vitamin D₂ and related photoproducts in edible mushrooms [16,82]. In addition to vitamin D₂, photoproducts such as previtamin D₂, lumisterol2, and tachysterol2 have been detected after pulsed-UV treatment [82]. Similarly, pulsed light has been shown to enhance vitamin D₂ accumulation in C. militaris and L. edodes, with specific pulse frequencies or energy inputs yielding optimal conversion efficiency [82,83,84,85]. Moreover, pulsed-light pretreatment prior to drying can help preserve a substantial proportion of vitamin D₂ during storage [86].

Beyond vitamin D₂ biosynthesis, pulsed light also modulates antioxidant capacity and metabolite profiles in a dose-dependent manner [16,57,85]. While excessive treatment may lead to reductions in vitamin C, phenolics, and overall antioxidant capacity due to oxidative stress, appropriately controlled doses can enhance total phenolic content and improve functional quality. In C. militaris, pulsed light has been reported to increase total amino acid content without compromising umami intensity [57]. In L. edodes, it helps preserve ergothioneine levels and anti-inflammatory activity [85]. Furthermore, as a pretreatment prior to drying, optimized pulsed-light conditions can reduce browning, suppress the formation of undesirable compounds such as 5-hydroxymethylfurfural, and maintain characteristic flavor profiles [87].

Overall, pulsed light represents an efficient strategy for enhancing vitamin D₂ content in edible mushrooms. More importantly, under carefully optimized conditions, it can simultaneously improve phenolic accumulation, amino acid retention, and color stability in processed products. These effects are closely associated with dose-dependent oxidative modulation and photochemical transformation processes. However, excessive irradiation may induce oxidative damage, highlighting the necessity of precise parameter optimization.

4.2.5. Critical Discussion on the Comprehensive Table: Systematic Driving Mechanisms of Multidimensional Variables on Fungal Photoresponses

As compiled in Table 3, although extensive literature documents the regulatory effects of diverse optical radiation environments on the quality and growth of edible mushrooms, a systematic framework defining optimal standardized protocols remains elusive due to the extreme variability of experimental parameters (e.g., light intensity, wavelength, and exposure time). This limitation stems primarily from the conventional assumption in fungal photobiology that photobiological effects exhibit linear relationships. However, as complex biological organisms, fungi exhibit distinct “critical thresholds” in their physiological metabolism in response to light. Once the radiant dose (fluence) falls below or exceeds these specific thresholds, the metabolic network undergoes major non-linear transitions, characterized by unimodal or step-like response profiles. A comprehensive compilation of these treatment conditions and their reported effects on mushroom bioactivity, chemical composition, and postharvest quality is provided in

Table A1. Comprehensive data of light and irradiation treatments on mushroom bioactivity, chemical composition, and postharvest quality.

Light Type	Typical Wavelength	Dose/Intensity	Exposure Time	Targeted Species	Major Physiological Outcomes	Advantages	Disadvantages	References
Visible light (blue)—mycelial stage	450–460 nm	10–40 µmol·m−2·s−1 (PPFD)	12 h/day (12/12 L/D)	Hericium erinaceus	Increases mycelial biomass (+12.28%), radial growth, substrate colonization; enhances anticancer activity (SW480 IC50 133.71 µg/mL; HepG2 IC50 114.84 µg/mL); improves DPPH scavenging.	LED integration; precise intensity control; enhances nutraceutical metabolites.	Optimal PPFD species-specific; effects not studied on fruiting bodies.	[88]
Visible light (blue)—mycelial and fruiting body	450–460 nm	1–20 µmol·m−2·s−1 (optimal 10)	12 h/day	Ganoderma sichuanense (antler-type)	Blue light (10 µmol·m−2·s−1) gives highest yield, polysaccharides (15.62%), triterpenoids (82.44 mg/g), DPPH scavenging (31.46–42.96%), and total phenolics. Red light significantly lowers yield and bioactives.	First systematic LED optimization for this species; clear optimal PPFD identified.	Red and green light inferior to blue.	[89]
Visible light (blue)—fruiting body	450–475 nm	10–40 µmol·m−2·s−1 (PPFD)	8–12 h/day	Lentinus sajor-caju	Blue light increases dry biomass (+50% vs. white light), DPPH scavenging, Fe2+ chelation, reducing power, and soluble sugars (peak at 40 PPFD).	Clear PPFD dose–response; blue superior to red/green/white.	Excess PPFD (>40) not beneficial.	[60]
Visible light (blue)—fruiting body	450–475 nm (blue LED)	Not specified (intensity not given in PPFD)	12 h/day	Lyophyllum ulmarium	Blue LED increases ergosterol (160.21 mg/100 g), higher than fluorescent control (143.58 mg/100 g). Yellow LED gave highest ergosterol (171.55 mg/100 g).	LED effective substitute for fluorescent lighting.	PPFD not reported; mechanism unclear.	[59]
Visible light (blue + UV combination)—fresh-cut storage	Blue LED (450–475 nm) + UV-C/UV-B	Blue PPFD not specified; UV dose not specified	Combined treatment	P. eryngii (fresh-cut)	Increases vitamin D2 (0.77 µg/100 g), soluble protein (+23.36%), reducing sugar (2.06× control at day 10); delays declines in flavonoids, total phenolics, and vitamin C; inhibits microbial growth; enhances ROS and secondary metabolism enzyme activities; delays cell wall and mitochondrial degradation.	Dual function: nutritional fortification + preservation.	Complex parameter optimization; individual contributions of blue vs. UV not separable.	[90]
UV-B—low dose	280–315 nm	200–222 mJ/cm2	23–25 s	L. edodes, A. bisporus, P. ostreatus, P. eryngii	Increases total phenolics, flavonoids, ergothioneine (peak at low dose, 0.21–2.6 mg/g DW), glutamate, antioxidant and antibacterial activities. L. edodes and P. ostreatus show strongest antioxidant response.	Short exposure (seconds) effective; enhances multiple nutraceuticals.	Optimal dose species-specific; high dose may reduce ergothioneine.	[63]
UV-B—high dose	280–315 nm	390–430 mJ/cm2	23–25 s	Same as above	Higher UV-B reduces or reverses beneficial effects; ergothioneine and antioxidant activity decline; risk of oxidative stress.	Useful for threshold determination.	Exceeding optimal dose induces oxidative damage.	[63]
UV-B—high dose (mycelia)	280–315 nm	~3.6 W/m2 × 40 min = ~8.64 kJ/m2	40 min	P. eryngii submerged mycelia	Vitamin D2 reaches 314.75 µg/g DW; total phenolics: 12.52 → 17.54 mg GAE/g; flavonoids: 1.19 → 1.72 mg QE/g; β-glucan: 26.18 → 32.21%; DPPH antioxidant activity ↑; essential amino acids (glutamate, arginine, threonine) ↑.	Extremely high vitamin D2 conversion (record level); mycelial biomass as a valuable source.	Long exposure (40 min); mechanism not fully elucidated.	[64]
UV-B—vitamin D2 quantification	280–315 nm	Not specified (commercial treatment)	Not specified	A. bisporus (white button)	Fresh mushroom vitamin D2 = 21.5 ± 2.4 µg/100 g FW (42% in cap peel). Cooking loss 9–63%; 75–190 g cooked mushroom meets RDI (15 µg).	Consumer-relevant cooking data; highlights peel contribution.	No UV dose reported—not used for dose–response.	[105]
Gamma irradiation—low dose (preservation)	60Co γ-rays	0.8 kGy	Not specified	Volvariella volvacea	0.8 kGy + sodium dehydroacetate (SD): reduces browning, weight loss (10.53–34.73%), respiration rate (17.20–48.72%), microbial load; increases CAT, SOD, POD (SOD +13.68–40.53%); maintains membrane integrity; delays senescence.	Combined treatment (γ + SD) effective for shelf-life extension.	Synergy with chemical preservative may not be suitable for organic/natural labels.	[91]
Gamma irradiation—low dose (nutrient preservation)	60Co γ-rays	1.5–2.0 kGy	Not specified	A. bisporus (white)	2 kGy + essential oil fumigation: vitamin D2 preserved (106.30–114.40 mg/kg DW); total phenolics and flavonoids maintained; shelf-life extended 18 days at 4 °C.	Maintains nutritional quality during storage.	Essential oil may affect flavor; consumer acceptance.	[79]
Gamma irradiation—low dose (water properties)	60Co γ-rays	1.0 kGy	Not specified	L. edodes (shiitake)	Delays browning, stabilizes water status (bound water content), maintains firmness and microstructure during postharvest storage.	Improves texture and water-holding capacity.	Dose optimization needed for different cultivars.	[106]
Gamma irradiation—moderate dose (polysaccharide structure)	60Co γ-rays	2.5–5.0 kGy	Not specified	A. bisporus powder	2.5–5 kGy: no significant change in total phenolics, flavonoids, or antioxidant activity (DPPH/FRAP).	Low doses safe for powder processing; retain functional components.	Higher doses may degrade polysaccharides.	[92]
Gamma irradiation—moderate dose (polysaccharide from M. sextelata)	60Co γ-rays	2.5–5.0 kGy	Not specified	Morchella sextelata (isolated polysaccharide)	Dose-dependent reduction in average molecular weight; increased surface porosity; enhanced radical scavenging; H2O2 protection of PC12 cells.	Improves bioactivity of polysaccharides via controlled degradation.	Not applicable to fresh mushroom preservation.	[93]
Gamma irradiation—high dose (preservation—Portobello)	60Co γ-rays	5 kGy	1–2 h (dose rate ~4.5–32 kGy/h)	A. bisporus (Portobello)	5 kGy: increases protein content; preserves sugars and ergosterol better than electron beam; reduces agaritine (1.54 → 1.35 g/kg DW). 10 kGy: no significant change in protein or polysaccharide.	High penetration; can treat packaged products; reduces agaritine.	High capital cost; may reduce unsaturated fatty acids; consumer acceptance issues.	[72]
Gamma irradiation—very high dose (polysaccharide degradation)	60Co γ-rays	10–1000 kGy	Hours	Morchella sextelata (isolated polysaccharide)	100 kGy: significant reduction in Mw and particle size; increased surface porosity; thermal stability increases at 10–100 kGy then decreases at 1000 kGy; antioxidant activity increases with dose at low polysaccharide concentration (50–100 µg/mL).	Enhances bioactivity of isolated polysaccharides.	Not for fresh produce; extremely high doses.	[93]
Pulsed light (PL)—low to moderate fluence (fresh-cut preservation)	Broad spectrum + UV (xenon)	4.8 J·cm−2 (optimal)	Microsecond pulses	A. bisporus (fresh-cut)	4.8 J·cm−2 optimal: extends microbial shelf life 2–3 days, minimal texture change, preserves color, phenolics, and vitamin D2 (retains 64.2% after 5 d at 4 °C). Lower doses (0.13–1.11 J·cm−2) inactivate PPO/POD (first-order kinetics, kPPO = 3.84 cm2/J).	Fast, non-thermal; dual function (decontamination + vitamin D2 enrichment).	High fluence (≥12 J·cm−2) causes thermal damage, PPO activation, browning, loss of phenolics/vitamin C.	[16,84]
Pulsed light (PL)—whole mushroom storage	Broad spectrum + UV (xenon)	1.11 J·cm−2	Microsecond pulses	A. bisporus (whole)	1.11 J·cm−2: PPO/POD inactivation, browning delayed, polyphenols retained, vitamin D2 increased. After 5 d at 4 °C: 90.6% polyphenols, 78.9% antioxidant activity, 64.2% vitamin D2 retained.	Applicable to whole mushrooms (not just sliced).	Enzyme inactivation incomplete at 1.11 J·cm−2; whole mushroom geometry affects efficacy.	[84]
Pulsed light (PL)—C. militaris	Broad spectrum + UV (xenon)	Not reported (only pulse number: 3,6,9)	Microsecond pulses	C. militaris	Increases vitamin D2; total amino acids +9–48%; preserves umami and antioxidant components.	Fast treatment, improves nutritional quality.	No dose/fluence reported —parameter comparability limited.	[57]
Pulsed light (IPL)—drying pretreatment	Broad spectrum + UV (xenon)	400 J total energy (25 pulses); fluence not reported	Microsecond pulses	L. edodes (dried)	25-pulse IPL pretreatment: reduces initial PPO activity by 42.83%, browning index ↓ 43.02%, browning degree ↓ 47.54%; increases polysaccharides, reducing sugars, antioxidant activity; reduces 5-HMF; preserves umami during subsequent hot-air drying.	Effective for dried mushroom processing; controls enzymatic browning.	Only as drying pretreatment; fluence (J·cm−2) not reported—poor comparability.	[87]
Pulsed light (IPL)—mechanism on PPO	Broad spectrum + UV (xenon)	Not reported (same 400 J/25 pulses likely)	Microsecond pulses	Mushroom polyphenol oxidase (PPO)	IPL changes PPO secondary structure (FTIR), alters physico-chemical properties and surface topography, leading to irreversible enzyme inactivation.	Mechanistic insight into IPL-induced enzyme inactivation.	No fluence reported; pure enzyme study, not whole mushroom.	[107]

(Appendix A).

To overcome the current impasse where heterogeneous literature on Pleurotus and Cordyceps cannot be horizontally compared due to inconsistent units (e.g., some studies utilize photosynthetic photon flux density (μmol·m⁻²·s⁻¹), while others employ radiant intensity (W·m⁻²) or pulse counts), this systematic review proposes the introduction of “total fluence” (J·cm⁻²) as a normalized light dose parameter. Standardizing energy input via the formula J·cm⁻² = radiant intensity (W·cm⁻²) × total exposure time (s) enables a more intuitive mapping of the quantitative structure-activity relationships between “action spectra” and “metabolic outcomes” across diverse species. For instance, when optimizing the ratio of combined visible light (red and blue), a theoretical equilibrium solution emerges: an ideal blue-to-red ratio (typically ranging from 1:4 to 1:2, depending on the species) can maximize the synergistic activation of the white collar complex (WCC) and phytochrome-like pathways. This synergy accelerates the highly efficient morphological transition from mycelium to fruiting body and maximizes biomass. Concurrently, supplemented by this low-dose red light, it prevents the inhibition of actinic conversion from ergosterol to vitamin D₂ caused by excessive high-energy blue radiation, thereby striking an optimal balance between biomass yield and nutritional fortification.

To improve comparability among studies, total fluence is recommended as a normalized optical dose parameter when sufficient irradiance and exposure-time information is available. For optical treatments, fluence can be calculated as irradiance multiplied by exposure time and expressed as J/cm². However, absorbed dose from gamma irradiation should be reported separately in kGy because it is not directly comparable with optical surface fluence.

For optical treatments: Fluence (J/cm²) = irradiance (W/cm²) × exposure time (s). When irradiance is reported as W/m², J/cm² = W/m² × s/10,000. When dose is reported as kJ/m², 1 kJ/m² = 0.1 J/cm². For LED PPFD data, energy-equivalent fluence should be estimated only when the peak wavelength or spectral distribution is clear; otherwise, retain photon fluence or the original PPFD.

Table 3. Cross-species comparison of action spectra, normalized fluence, and metabolic outcomes in edible mushrooms.

Light/Irradiation type	Representative Species	Key Physiological Outcomes	Dose/Intensity	Normalized Total Fluence (J/cm2) (If Optical)	Main Advantages and Disadvantages	References
Blue light (mycelial stage)	Hericium erinaceus	Biomass +12.28%, anticancer activity ↑, DPPH scavenging ↑	10–40 µmol·m−2·s−1 PPFD, 12 h/d	~11.5–46.0 (estimated from 450 nm)	LED precise control, enhances nutraceuticals; species-specific optimal PPFD, not studied on fruiting bodies	[88]
Blue light (mycelial + fruiting)	Ganoderma sichuanense	Polysaccharides 15.62%, triterpenoids 82.44 mg/g, antioxidant ↑ (optimal at 10 PPFD)	1–20 PPFD, 12 h/d	~1.15–23.0 (estimated from 450 nm)	first systematic LED optimization, clear optimal PPFD; red/green light inferior	[89]
Blue light (fruiting body)	Lentinus sajor-caju	Dry biomass +50%, DPPH, Fe2+ chelation, reducing power ↑	10–40 PPFD, 8–12 h/d	~7.7–46.0 (estimated, depending on 8 or 12 h)	blue superior to white/red/green, clear PPFD response; >40 PPFD not beneficial	[60]
Blue light (fruiting body)	Lyophyllum ulmarium	Ergosterol ↑ (160 mg/100 g blue; 171 mg/100 g yellow)	intensity not given (12 h/d)	N/A	LED effective substitute for fluorescent; PPFD not reported, mechanism unclear	[59]
Blue + UV (fresh-cut)	Pleurotus eryngii	Vitamin D2 ↑, soluble protein +23.36%, sugar ↑, delays quality loss	blue PPFD not given, UV dose not given	N/A	dual function (fortification + preservation); individual contributions of blue vs. UV not separable	[90]
UV-B (low dose)	L. edodes, A. bisporus, P. ostreatus, P. eryngii	Total phenolics, flavonoids, ergothioneine ↑, antioxidant and antibacterial ↑	200–222 mJ/cm2, 23–25 s	0.20–0.22	short exposure effective, enhances nutraceuticals; optimal dose species-specific, high dose reduces ergothioneine	[63]
UV-B (high dose)	Same as above	Beneficial effects reduced or reversed, oxidative stress risk	390–430 mJ/cm2, 23–25 s	0.39–0.43	exceeding optimal dose induces oxidative damage	[63]
UV-B (mycelia)	P. eryngii (submerged)	Vitamin D2 up to 314.75 µg/g DW, phenolics, β-glucan ↑	~8.64 kJ/m2, 40 min	0.864	extremely high vitamin D2 conversion, mycelial biomass as valuable source; long exposure (40 min), mechanism not fully elucidated	[64]
Gamma irradiation (low dose, preservation)	Volvariella volvacea	Reduces browning, weight loss, respiration; SOD ↑, delays senescence	0.8 kGy (+sodium dehydroacetate)	N/A	combined treatment effective for shelf-life; chemical preservative not suitable for organic/natural labels	[91]
Gamma irradiation (low dose, nutrient preservation)	Agaricus bisporus	Vitamin D2, total phenolics and flavonoids maintained; shelf-life extended	1.5–2.0 kGy (+essential oil fumigation)	N/A	maintains nutritional quality during storage; essential oil may affect flavor, consumer acceptance	[91]
Gamma irradiation (moderate dose, polysaccharide structure)	A. bisporus powder/Morchella sextelata polysaccharide	No significant change in phenolics/flavonoids/antioxidant activity (powder); Mw ↓, radical scavenging ↑ (isolated polysaccharide)	2.5–5.0 kGy	N/A	low-moderate doses safe, retain functional components; higher doses may degrade polysaccharides	[92,93]
Pulsed light (low-moderate fluence, fresh-cut)	A. bisporus	Microbial shelf-life extended 2–3 d, vitamin D2 retained 64.2%	4.8 J·cm−2	4.8	fast, non-thermal, dual function (decontamination + vitamin D2 enrichment); ≥12 J·cm−2 causes thermal damage, browning, loss of phenolics/vitamin C	[16,84]
Pulsed light (whole mushroom storage)	A. bisporus	PPO/POD inactivation, browning delayed, polyphenols retained 90.6%	1.11 J·cm−2	1.11	applicable to whole mushrooms; enzyme inactivation incomplete, geometry affects efficacy	[84]
Pulsed light (drying pretreatment)	Lentinus edodes	PPO activity ↓ 42.8%, browning index ↓, polysaccharides and reducing sugars ↑	400 J/25 pulses (fluence not reported)	N/A	controls enzymatic browning; only as drying pretreatment, fluence (J·cm−2) not reported	[87]

Table 3. Cross-species comparison of action spectra, normalized fluence, and metabolic outcomes in edible mushrooms.

Light/Irradiation type	Representative Species	Key Physiological Outcomes	Dose/Intensity	Normalized Total Fluence (J/cm2) (If Optical)	Main Advantages and Disadvantages	References
Blue light (mycelial stage)	Hericium erinaceus	Biomass +12.28%, anticancer activity ↑, DPPH scavenging ↑	10–40 µmol·m−2·s−1 PPFD, 12 h/d	~11.5–46.0 (estimated from 450 nm)	LED precise control, enhances nutraceuticals; species-specific optimal PPFD, not studied on fruiting bodies	[88]
Blue light (mycelial + fruiting)	Ganoderma sichuanense	Polysaccharides 15.62%, triterpenoids 82.44 mg/g, antioxidant ↑ (optimal at 10 PPFD)	1–20 PPFD, 12 h/d	~1.15–23.0 (estimated from 450 nm)	first systematic LED optimization, clear optimal PPFD; red/green light inferior	[89]
Blue light (fruiting body)	Lentinus sajor-caju	Dry biomass +50%, DPPH, Fe2+ chelation, reducing power ↑	10–40 PPFD, 8–12 h/d	~7.7–46.0 (estimated, depending on 8 or 12 h)	blue superior to white/red/green, clear PPFD response; >40 PPFD not beneficial	[60]
Blue light (fruiting body)	Lyophyllum ulmarium	Ergosterol ↑ (160 mg/100 g blue; 171 mg/100 g yellow)	intensity not given (12 h/d)	N/A	LED effective substitute for fluorescent; PPFD not reported, mechanism unclear	[59]
Blue + UV (fresh-cut)	Pleurotus eryngii	Vitamin D2 ↑, soluble protein +23.36%, sugar ↑, delays quality loss	blue PPFD not given, UV dose not given	N/A	dual function (fortification + preservation); individual contributions of blue vs. UV not separable	[90]
UV-B (low dose)	L. edodes, A. bisporus, P. ostreatus, P. eryngii	Total phenolics, flavonoids, ergothioneine ↑, antioxidant and antibacterial ↑	200–222 mJ/cm2, 23–25 s	0.20–0.22	short exposure effective, enhances nutraceuticals; optimal dose species-specific, high dose reduces ergothioneine	[63]
UV-B (high dose)	Same as above	Beneficial effects reduced or reversed, oxidative stress risk	390–430 mJ/cm2, 23–25 s	0.39–0.43	exceeding optimal dose induces oxidative damage	[63]
UV-B (mycelia)	P. eryngii (submerged)	Vitamin D2 up to 314.75 µg/g DW, phenolics, β-glucan ↑	~8.64 kJ/m2, 40 min	0.864	extremely high vitamin D2 conversion, mycelial biomass as valuable source; long exposure (40 min), mechanism not fully elucidated	[64]
Gamma irradiation (low dose, preservation)	Volvariella volvacea	Reduces browning, weight loss, respiration; SOD ↑, delays senescence	0.8 kGy (+sodium dehydroacetate)	N/A	combined treatment effective for shelf-life; chemical preservative not suitable for organic/natural labels	[91]
Gamma irradiation (low dose, nutrient preservation)	Agaricus bisporus	Vitamin D2, total phenolics and flavonoids maintained; shelf-life extended	1.5–2.0 kGy (+essential oil fumigation)	N/A	maintains nutritional quality during storage; essential oil may affect flavor, consumer acceptance	[91]
Gamma irradiation (moderate dose, polysaccharide structure)	A. bisporus powder/Morchella sextelata polysaccharide	No significant change in phenolics/flavonoids/antioxidant activity (powder); Mw ↓, radical scavenging ↑ (isolated polysaccharide)	2.5–5.0 kGy	N/A	low-moderate doses safe, retain functional components; higher doses may degrade polysaccharides	[92,93]
Pulsed light (low-moderate fluence, fresh-cut)	A. bisporus	Microbial shelf-life extended 2–3 d, vitamin D2 retained 64.2%	4.8 J·cm−2	4.8	fast, non-thermal, dual function (decontamination + vitamin D2 enrichment); ≥12 J·cm−2 causes thermal damage, browning, loss of phenolics/vitamin C	[16,84]
Pulsed light (whole mushroom storage)	A. bisporus	PPO/POD inactivation, browning delayed, polyphenols retained 90.6%	1.11 J·cm−2	1.11	applicable to whole mushrooms; enzyme inactivation incomplete, geometry affects efficacy	[84]
Pulsed light (drying pretreatment)	Lentinus edodes	PPO activity ↓ 42.8%, browning index ↓, polysaccharides and reducing sugars ↑	400 J/25 pulses (fluence not reported)	N/A	controls enzymatic browning; only as drying pretreatment, fluence (J·cm−2) not reported	[87]

This sensitivity to radiation dose thresholds and spectral proportions is deeply rooted in the life history and ecological niche of fungi. Evolutionary ecology indicates that fungi inhabiting humus-rich, densely canopy-shaded deep wood environments (characterized by low-light or aphotic conditions, such as L. edodes and Pleurotus spp.) possess highly sensitive photoreceptor systems. Consequently, an extremely low total fluence is sufficient to trigger primordium differentiation, whereas high-dose ultraviolet (UV) radiation induces profound oxidative stress sensitivity. Conversely, wild strains exposed to open fields, grasslands, or high-altitude sun-exposed environments (such as A. bisporus, Ophiocordyceps sinensis, or C. militaris) have evolved more robust photoprotective mechanisms within their genomes. This ecological niche perspective elucidates why different species display drastically disparate tolerances and nutritional accumulation behaviors under identical UV or pulsed light dosages.

Furthermore, when evaluating the molecular efficiency of UV-induced conversion of ergosterol to vitamin D₂, prior research has predominantly focused on biochemical pathways while neglecting a crucial physical variable: the physical barriers of the fungi themselves. The penetration depth of light is strictly constrained by the cell wall thickness (microstructure) of the pileus epidermis and the accumulation of endogenous chromophores (such as melanin, carotenoids, and other pigments) within the tissue. These physical structures and pigments function as natural optical filters on the fungal surface. Species with high melanin content (e.g., mature caps of L. edodes) strongly absorb and scatter UV radiation, significantly attenuating the photon energy penetrating into deeper tissues. Therefore, to surmount this physical barrier and achieve a vitamin D₂ conversion efficiency equivalent to that of pale or unpigmented species (e.g., white A. bisporus), these specific strains often require several-fold higher radiation intensities or total fluences in experimental setups. The omission of this physical variable is the core factor driving the immense heterogeneity in reported conversion efficiencies across different species.

Finally, regarding the temporal dynamics of light treatments, most existing studies are confined to coarse descriptions of photoperiods (e.g., 8 h/12 h continuous illumination), lacking theoretical model construction for time-frequency modulation. Compared with continuous photoperiods, pulsed light (PL) offers a promising theoretical alternative: precisely activating photoreceptor proteins or instantaneously inducing sub-lethal reactive oxygen species (ROS) signals via microsecond-scale “bursts of high-energy photon flux” can serve as a molecular switch to rapidly upregulate key rate-limiting enzymes of secondary metabolic networks (such as phenylalanine ammonia-lyase, PAL). Subsequently, the prolonged microsecond-scale “dark intervals” provide fungal cells with an adequate buffering period to scavenge excess free radicals via endogenous antioxidant systems (e.g., SOD, CAT), thereby effectively avoiding the chronic oxidative stress and tissue thermal damage associated with continuous exposure. Integrating light frequency and temporal dynamics into experimental designs establishes a solid theoretical model framework for the future development of high-yield, high-quality, and energy-efficient intelligent precision light-recipe cultivation systems.

4.3. Patterns Governing Differential Effects of Radiation-Based Treatments

Overall, the quality responses of edible mushrooms to different radiation-based treatments exhibit several general regulatory patterns. Visible light primarily governs morphogenesis and basal metabolic activity; ultraviolet (UV) radiation is particularly effective in promoting nutritional fortification and enhancing antioxidant capacity; gamma irradiation contributes mainly through microbial inactivation and dose-dependent oxidative or enzymatic responses, with DNA damage/repair being most relevant to microbial cells; and pulsed light enables rapid, multifunctional regulation during postharvest handling.

These differences are largely attributable to variations in wavelength, photon energy, treatment mode, and dose. Despite these differences, the biological responses are ultimately integrated through reactive oxygen species (ROS)-mediated signaling and downstream metabolic reprogramming. Although synergistic and antagonistic interactions among different radiation-based treatments are highly plausible, current evidence remains fragmented, and systematic comparative studies are still limited.

Future research should therefore focus on standardizing treatment parameters, elucidating the molecular mechanisms underlying multi-light interactions, and constructing cross-scale regulatory networks. Such efforts will provide a more comprehensive theoretical framework for the precise and industrial application of light-based technologies in edible mushroom production and preservation.

Figure 6 presents a heatmap that systematically summarizes the effects of light treatments on 11 quality indicators, spanning from primordium growth to postharvest storage.

5. Conclusions

5.1. Industrial Applicability and Sustainability

Despite these advantages, several barriers continue to limit industrial adoption. These include equipment investment costs, energy consumption, treatment uniformity, process standardization, and consumer acceptance. Future studies should therefore incorporate techno-economic analyses and life-cycle assessments to evaluate the long-term sustainability of different light-treatment strategies under commercial production conditions. The practical implementation of light-based technologies in edible mushroom production depends not only on their biological effectiveness but also on their economic feasibility, scalability, and sustainability. Visible-light regulation can be readily integrated into existing cultivation facilities through LED systems and therefore represents the most accessible technology for large-scale production. In contrast, ultraviolet irradiation, pulsed light, and gamma irradiation are primarily applied during postharvest preservation to improve nutritional quality and extend shelf life.

Importantly, the value of radiation-based treatments extends beyond quality enhancement. Fresh mushrooms are highly perishable commodities that undergo rapid moisture loss, browning, respiration, and microbial spoilage during storage and distribution. Consequently, substantial product losses occur throughout the supply chain. Even a shelf-life extension of only 24–48 h may significantly reduce food waste, improve logistics efficiency, and lower the carbon footprint associated with mushroom production and distribution. Future studies should therefore combine quality evaluation with techno-economic and life-cycle assessments to determine the long-term sustainability of different light-treatment strategies under commercial production conditions.

5.2. Smart Lighting and Intelligent Mushroom Production

Recent advances in sensor technology, machine vision, hyperspectral imaging, Internet-of-Things (IoT) systems, and artificial intelligence (AI) provide new opportunities for precision regulation of mushroom quality [94]. Future production systems are expected to evolve from fixed illumination programs toward intelligent closed-loop control systems [95].

Optical sensors and spectroscopic technologies can continuously monitor physiological and quality-related indicators, including morphology, surface reflectance, moisture status, growth dynamics, and stress responses. These real-time data streams can be integrated with AI-based predictive models to dynamically regulate light spectrum, intensity, photoperiod, pulse frequency, and irradiation dose according to developmental stage and production objectives [96,97]. The integration of hyperspectral sensing, big-data analytics, digital-twin models, and automated environmental control platforms may further enable data-driven optimization of mushroom cultivation and postharvest management [98,99]. Such intelligent lighting systems have the potential to maximize product quality while minimizing energy consumption, excessive irradiation stress, and production costs.

5.3. Regulatory Considerations and Consumer Acceptance

The industrial adoption of light-based technologies is also influenced by regulatory frameworks and consumer acceptance. At present, considerable differences exist among China, the European Union, and the United States regarding the regulation of irradiation and ultraviolet treatments [100,101,102]. These inconsistencies create barriers to technology transfer, international trade, and large-scale commercialization.

Given the increasing globalization of food supply chains and the rapid development of light-based processing technologies, international harmonization of safety standards is becoming increasingly important. Future regulatory efforts should focus on establishing scientifically based criteria for evaluating different light treatments, clarifying acceptable processing doses, and developing unified guidelines for emerging technologies such as UV and pulsed light. Greater coordination under international frameworks, including the Codex Alimentarius system, would facilitate the definition of light doses that can be regarded as “safe processing” while ensuring food quality and consumer confidence.

Overall, light regulates edible mushroom quality through coordinated effects on growth and development, metabolic reprogramming, photochemical responses, and microbial inhibition. Numerous studies have demonstrated that visible light, ultraviolet irradiation, pulsed light, and gamma irradiation can enhance nutritional, sensory, and storage-related quality attributes. However, excessive irradiation may also accelerate oxidative processes and induce undesirable quality changes, making the identification of appropriate treatment windows essential for practical applications [103]. Treatment outcomes are strongly influenced by irradiation dose, intensity, exposure duration, spectral distribution, and the physiological state of mushroom tissues [104].

Future research should increasingly regard preharvest and postharvest radiation regulation as a continuous quality-management system rather than as separate research domains. Particular attention should be given to elucidating photoreceptor-mediated signaling pathways, optimizing dose–response relationships, and exploring synergistic effects between radiation-based treatments and other preservation technologies, such as modified-atmosphere packaging, edible coatings, refrigeration, chemical treatments, and ozone treatment [68]. In addition, treatment efficacy remains highly dependent on species characteristics, developmental stage, tissue structure, and processing objectives [57].

The integration of smart sensing technologies, AI-assisted decision-making, and adaptive lighting control systems is expected to transform conventional mushroom production into a data-driven precision-management platform. Such developments will support the production of high-quality mushrooms while reducing food waste, improving resource-use efficiency, and promoting the sustainable development of the mushroom industry.