Sustainable Production of Alternative Proteins from Basidiomycetes: Valorization of Mycelial and Fruiting Body Biomass

Abstract

Global population growth, climate change, and the environmental impact of livestock production have accelerated the search for sustainable and efficient protein sources. Fruiting bodies (mushrooms) and mycelial biomass have emerged as promising alternatives due to their high nutritional quality, low ecological footprint, and compatibility with circular bioeconomy principles. This review highlights the nutritional, biotechnological, and environmental aspects of fungal proteins obtained from both fruiting bodies and mycelial biomass of Basidiomycetes. Emphasis is placed on amino acid composition, protein digestibility, and advances in cultivation and fermentation systems for large-scale production. Submerged and solid-state fermentation processes are analyzed in terms of scalability, resource efficiency, and integration with agro-industrial residues for sustainable bioprocessing. Comparative analyses reveal that mycelial biomass production achieves high protein yields with significantly reduced land, water, and energy requirements compared to conventional protein sources. Emerging fungal species such as Schizophyllum commune and Auricularia polytricha demonstrate strong potential for producing protein-rich mycelia applicable to functional and plant-based foods. Finally, the review discusses current technological innovations, regulatory frameworks, and market perspectives that position fungal biomass as a strategic component in the ongoing global protein transition.

1. Introduction

In recent decades, the world has faced increasing challenges regarding food security, driven by population growth, climate change, accelerated urbanization, and the growing demand for animal-derived foods. The global population is projected to reach nearly 10 billion by 2050, requiring a 70% increase in food production. Among essential nutrients, proteins play a central role in human health, being vital for growth, maintenance, and metabolic regulation. Traditionally, most dietary proteins have been obtained from animal sources such as meat, milk, eggs, and fish. However, conventional animal production systems are associated with significant environmental and ethical concerns. They are responsible for high greenhouse gas emissions, particularly methane and carbon dioxide, originating from intensive livestock operations [1]. These systems also demand excessive natural resources, including water and land, which compete with crops intended for human consumption. Moreover, pasture expansion and large-scale monocultures for animal feed contribute to biodiversity loss. Ethical and animal welfare concerns are increasingly influencing consumer preferences and driving demand for alternative proteins (APs). Finally, conventional livestock systems remain vulnerable to zoonotic outbreaks, such as avian influenza, swine fever, and COVID-19, which can disrupt food chains and threaten public health [2,3]. These limitations have driven global efforts to identify AP sources capable of sustainably meeting the growing demand. Emerging categories include plant-based proteins (legumes, cereals, oilseeds, and algae); insect proteins, noted for high efficiency and low environmental impact; cell-cultured meat, produced via animal cell cultivation; and microbial proteins, derived from yeasts, bacteria, and fungi, particularly mycelial biomass from mushrooms [4,5,6,7,8,9].

The transition toward APs is supported by combined environmental, social, and economic drivers: environmental sustainability, aiming to reduce the carbon footprint and ecological impact of food systems; shifting consumer behavior, with increasing vegetarian and vegan populations in both developed and emerging economies; technological advances, including precision fermentation and food biotechnology innovations; health and wellness awareness, as excessive consumption of red and processed meats is linked to chronic diseases; and public policies and incentives, encouraging sustainable production and deforestation reduction [10]. These factors have accelerated what is now referred to as the protein revolution, in which new ingredients and products emerge to meet the demand for sustainable, ethical, and nutritious food alternatives.

Basidiomycetes are a diverse class of higher fungi (phylum Basidiomycota) that includes most mushroom-forming species. They are distinguished by their ability to decompose lignin and other complex polymers, thereby converting lignin-rich agricultural residues into fungal biomass enriched in protein [11]. Many basidiomycetes are natural colonizers of wood, leaf litter, and other lignocellulosic materials, making them ideal candidates for valorizing side streams from forestry and agri-food industries. Because these fungi can be cultivated in enclosed or semi-controlled environments, their production does not require arable land, avoiding competition with crops or livestock systems traditionally used for protein supply.

Globally, approximately 2000 mushroom species are recognized as edible, and a subset of these has been granted Generally Recognized as Safe (GRAS) status for food use [12,13]. However, the majority of basidiomycete species have never entered the food chain, often due to their small fruiting bodies, unappealing taste or texture, or limited yield rather than toxicity [14]. The safety of non-traditional fungal species is regulated under frameworks such as GRAS in the U.S. and Novel Food in the E.U. These frameworks mandate comprehensive safety data, including compositional, allergenic, and toxicological evaluations, for market entry, thereby ensuring consumer safety [15]. Nutritionally, mushrooms are appreciated for their low fat and calorie content, high fiber, and richness in vitamins, minerals, and trace elements. Beyond their basic nutrient profile, several basidiomycetes also produce bioactive compounds associated with immunomodulatory, antioxidant, and metabolic health effects [12,16,17,18,19,20,21,22]. Nevertheless, large-scale mushroom cultivation is technically demanding and currently feasible for only about 80 species worldwide.

A promising alternative is to focus on mycelial biomass production instead of fruiting-body formation. To cultivate substrate mycelia of mushroom-forming fungi offers several advantages: (i) the vegetative (mycelial) phase of many species can grow efficiently on lignocellulosic feedstocks, whereas fruiting requires specific environmental triggers; (ii) bypassing the fruiting stage can significantly reduce production time, energy requirements, and infrastructure costs; (iii) species selection can emphasize protein yield and substrate conversion efficiency rather than mushroom morphology or sensory traits; and (iv) residual mycelial biomass remaining after mushroom cultivation can be recovered and processed as protein-rich material, enhancing resource circularity [11,23].

Despite these advantages, mycelial biomass from Basidiomycetes has received limited attention as a sustainable fungal protein source obtained from lignocellulosic substrates. Its classification as a “Novel Food” under current regulatory frameworks, together with the absence of standardized analytical protocols, has hindered broader adoption and comparative evaluation. Reported protein contents in basidiomycete mycelia vary considerably, largely due to inconsistent nitrogen-to-protein conversion factors, differences in cultivation media, and uncontrolled sampling conditions. Furthermore, knowledge gaps persist regarding optimal strategies for production, harvesting, and downstream processing of basidiomycete mycelia intended for food applications. Addressing these challenges is essential to fully accomplish the potential of Basidiomycetes as renewable and circular contributors to the emerging protein bioeconomy.

The main objective of this review is to provide a comprehensive overview of recent advances in the study of mycoproteins derived from Basidiomycetes, encompassing both fruiting bodies (mushrooms) and mycelial biomass. Special emphasis is placed on proteins produced by mycelial biomass through submerged and SSF systems, which represent promising, scalable, and resource-efficient biotechnological routes for sustainable protein production. By integrating nutritional, biotechnological, and environmental perspectives, this review aims to foster future research and industrial developments that leverage basidiomycete-derived mycoproteins as alternatives to conventional animal-based proteins. Industrial interest in fungal proteins is rapidly expanding, with pilot-scale initiatives and commercial ventures exploring basidiomycete biomass for food and nutraceutical applications. The success of mycoprotein products such as Quorn™ (https://www.quorn.co.uk/mycoprotein, accessed on 20 October 2025) [24], produced from the filamentous fungus Fusarium venenatum, demonstrates the market feasibility of fungal-derived proteins and stimulates the advancement of basidiomycete-based innovations.

2. Nutritional Quality and Biological Value of Proteins from Mushrooms

Proteins from mushrooms are increasingly recognized as valuable alternative sources of nutrition, thanks to their balanced amino acid composition and relatively good digestibility [25]. However, a detailed assessment of their nutritional quality requires evaluation not only of their amino acid profiles but also of their digestibility, protein efficiency, and comparison with conventional protein sources. A key metric for protein quality is the content and proportion of essential amino acids (EAAs) relative to the total amino acid pool [26]. Numerous studies show that many mushroom species contain all nine EAAs, although the relative abundances vary by species, substrate, and environmental conditions [27]. In the same work, environmental effects (soil type, pH, and tree diversity) had less influence on the amino acid profiles than species genetics. In cultivated mushrooms, Ionescu et al. report that Basidiomycota may contain protein levels ranging from 19% to 40% of their dry weight, with a complete and balanced amino acid composition [28]. In Pleurotus ostreatus, 13 amino acids were identified (5 essential, 8 non-essential), with aspartic acid being the most abundant (492.12 mg/100 g) and cysteine the least (9.32 mg/100 g). The ratio of EAAs to total amino acids was 0.11 [29]. In a broader review, Ayimbila et al. emphasize that many mushroom proteins, after in vitro digestion, meet the Food and Agriculture Organization of the United Nations (FAO) amino acid pattern for adults, reinforcing their suitability for human diets [25]. However, limitations are often observed in sulfur-containing amino acids (methionine and cysteine), which tend to be lower in fungal proteins compared to some animal sources [30]. To overcome the limitations of sulfur-containing amino acids, fungal proteins are often combined with complementary protein sources that provide a more balanced EAA profile. Cereal-based proteins such as wheat, rice, and corn are rich in methionine and cysteine, while legume proteins such as soy, lentils, peas, and chickpeas supply additional lysine and threonine, compensating for the limiting amino acids in fungi [31]. Such combinations not only improve the overall Protein Digestibility-Corrected Amino Acid Score (PDCAAS) and the Digestible Indispensable Amino Acid Score (DIAAS) but also enhance functional properties, including texture, water-holding capacity, and emulsification in food formulations. This synergistic approach has been successfully applied in hybrid or plant-forward food products, where fungal and plant proteins together yield complete amino acid composition and favorable sensory characteristics [24,31]. When comparing fungal proteins with more conventional sources (plant, animal), several advantages emerge (

Table 1. Essential amino acid composition (mg/g protein) in edible mushrooms compared with animal-derived proteins [25,33,34,35,36,37,38,39,40,41,42].

Essential Amino Acid	Pleurotus ostreatus	Lentinula edodes	Agaricus bisporus	Beef	Egg
Histidine	23	19	18	33	24
Isoleucine	42	37	35	54	40
Leucine	71	67	64	86	70
Lysine	65	61	59	91	60
Methionine + cysteine	29	27	24	42	56
Phenylalanine + tyrosine	72	70	66	84	93
Threonine	43	39	38	46	47
Tryptophan	11	9	8	11	17
Valine	49	45	43	55	50
Total EAAs	405	374	355	502	457

): (a) balanced amino acid profile, particularly good lysine content, helps to mitigate deficits often found in cereal proteins [30]; (b) high efficiency per unit area and lower environmental footprint, which will be discussed in later sections; and (c) lower levels of anti-nutritional factors relative to some plant proteins, although fungal cell wall polysaccharides (e.g., chitin, β-glucans) may affect digestibility unless treated or partially removed. Anti-nutritional factors are naturally occurring compounds, such as tannins, phytates, or trypsin inhibitors, that can reduce nutrient absorption or interfere with digestive enzymes, thereby limiting protein bioavailability [31,32]. Still, to assemble a fully “complete” protein with optimal levels of all EAAs, fungal proteins are often best combined with cereal proteins (which may supply extra methionine) to create a balanced matrix [31,32,33,34].

The data of Table 1 indicate that edible mushrooms provide a well-balanced amino acid profile, particularly rich in leucine, lysine, phenylalanine + tyrosine, and valine, which are often limiting in plant-based proteins such as cereals. Conversely, methionine and tryptophan are relatively lower, suggesting that combining fungal proteins with cereals (rich in sulfur-containing amino acids) could yield complementary nutritional profiles. According to Ayimbila et al., the protein digestibility of mushroom proteins can reach values between 70–85%, particularly after mild thermal or enzymatic processing, placing them in a similar range to legume proteins and approaching the digestibility of egg proteins [25]. Recent studies also confirm that mycelial proteins from submerged cultures can reach 40–45% crude protein content on a dry weight basis, with digestibility and EAA ratios comparable to conventional animal proteins [43,44,45]. The nutritional characteristics and biological value of these mycelial proteins are further elaborated in Section 6, which specifically addresses proteins derived from basidiomycete mycelial biomass.

3. Mushrooms in the Alternative Protein Landscape

Mushrooms and fungal mycelia (Figure 1) occupy a distinctive niche within the broader alternative-protein (AP) portfolio. Beyond their favorable nutritional profile and umami-rich sensorial qualities, they offer scalable bioprocess routes (SmF or SSF) and strong sustainability credentials that align with the goals of the circular bioeconomy. Recent syntheses position mycelium as a nutrient-dense ingredient capable of supporting health while enabling regenerative food systems, with promise for protein fortification and texture in plant-forward foods, that is, dietary patterns emphasizing plant-derived ingredients while still allowing limited inclusion of animal products [33,46].

Compared with most plant protein isolates and concentrates, fungal proteins generally exhibit a more balanced EAA profile, particularly due to their naturally high lysine levels, which makes them effective for complementing lysine-limited cereal proteins [47]. When subjected to appropriate processing (e.g., mild heat or enzymatic treatments), these proteins can also achieve high in vitro digestibility [47]. Contemporary reviews on mycoproteins (the best-studied form of food-grade fungal biomass) highlight evidence of favorable cardiometabolic effects, satiety responses, and muscle protein synthesis support, while detailing fermentation at scale and food development pathways. From a product-development standpoint, mushrooms contribute intrinsic fibrous structure and umami, enabling partial or full meat replacement with fewer additives and simpler labels. This has been emphasized in recent “mushrooms-as-meat-analogues” assessments and market-readiness overviews [48,49].

Environmental assessments consistently indicate that meals incorporating novel proteins (including fungal biomass) can reduce greenhouse gas (GHG) emissions, land use, and water footprints while maintaining nutritional value. A 2023 environmental and nutritional life-cycle assessment (LCA) reported up to 88% lower GWP, 83% lower land use, and 87% lower scarcity-weighted water use in meals featuring future/novel foods vs. animal source counterparts. A 2024 critical review of mycoprotein-based meat alternatives synthesized multiple LCAs and generally found impacts below those of beef and competitive with other plant-based analogues, while calling for more harmonized, product-level LCA reporting [50]. Earlier and broader syntheses also note lower climate impacts for mycoprotein, though legacy studies used older process assumptions.

On the demand side, recent analyses observe growing familiarity with APs and positive perceptions of mushroom-based products, driven by taste (umami), texture, and “naturalness.” Controlled studies comparing plant-based and mushroom-based meat alternatives report favorable consumer perceptions for shiitake/king oyster formats and highlight opportunities to optimize flavor/texture via processing. Contemporary market analyses (2024–2025) forecast steady expansion of the AP sector encompassing plant-, fungal-, algal-, and cell-cultivated sources. Within this landscape, fungal biomass is recognized for leveraging fermentation expertise and possessing clearer, near-term scalability prospects compared with cell-cultured meat [51].

Collectively, the evidence suggests that mushrooms and mycelia offer: (i) compelling nutrition-functionality, including EAA balance and techno-functional traits for binding, gelling, and texture; (ii) manufacturability through aerobic fermentation using low-cost carbon sources and agro-industrial side streams; and (iii) robust sustainability performance in LCAs relative to ruminant meat and competitive with most plant-based. Recent domain reviews synthesize these strengths and chart R&D priorities (strain improvement, substrate circularity, and product design) to accelerate adoption in staple foods and high-protein formats analogues [23,47].

4. Traditional Mushroom Cultivation as a Protein Source

Traditional mushroom cultivation remains one of the most established and accessible routes for obtaining fungal proteins for human consumption. This system focuses on the production of fruiting bodies (basidiomata), which are consumed fresh, dried, or processed into powders and extracts. Globally, the mushroom industry has evolved from small-scale artisanal production to a high-value agro-industrial sector supplying both domestic and export markets [52]. This cultivation strategy integrates bioconversion of lignocellulosic residues into high-protein food, aligning with circular bioeconomy principles. Agricultural by-products such as straw, sawdust, coffee husks, and fruit residues are transformed into nutrient-dense fungal biomass (Figure 2). Beyond its nutritional contribution, mushroom cultivation provides socioeconomic benefits, including job creation in rural communities and waste valorization from crop residues. The species most widely cultivated worldwide include Agaricus bisporus (button mushroom), Pleurotus spp. (oyster mushrooms), and Lentinula edodes (shiitake), which together account for more than 75% of the global production [53] (Figure 3).

Table 2. Main cultivated mushroom species and key characteristics [17,52,54,55].

Species	Key Characteristics	Protein Content (% Dry Weight)	Typical Substrate	Main Production Regions
Agaricus bisporus (button mushroom)	Most widely produced mushroom (35% of world output); mild flavor, short cultivation cycle.	20–25%	Compost of straw + manure	Europe, North America, China
Pleurotus spp. (oyster)	Fast-growing; degrades diverse lignocelluloses; low cost.	25–30%	Wheat/rice straw, corn cobs, coffee husks	Asia, Latin America, Africa
Lentinula edodes (shiitake)	Distinct aroma; immune-modulatory β-glucans (lentinan).	18–24%	Hardwood sawdust or logs	East Asia, Brazil
Flammulina velutipes (enoki)	Delicate, low-temperature species; mild taste.	18–22%	Corncob + sawdust	Japan, China, Korea
Grifola frondosa (maitake)	“Dancing mushroom”; both food and medicinal value.	20–25%	Hardwood sawdust	Japan, China
Volvariella volvacea (straw mushroom)	Short cycle (15–20 days); tropical species.	18–21%	Rice straw, banana leaves	Southeast Asia

summarizes their major cultivation features and nutritional relevance.

The substrate composition is critical to mushroom productivity and protein yield. Substrates not only provide carbon, nitrogen, and minerals but also influence enzyme secretion and flavor profile. Increasingly, agricultural residues such as sugarcane bagasse, corn stover, wheat bran, coffee husks, cocoa shells, and brewery by-products are used to enhance sustainability. Successful cultivation depends on precise control of environmental parameters such as temperature, humidity, light, aeration, and pH (

Table 3. Main environmental factors controlling mushroom growth and fruiting.

Parameter	Function	Optimal Range	Notes/Effects	References
Temperature	Regulates mycelial growth and fruiting body initiation	15–25 °C	Pleurotus grows faster near 25 °C; Flammulina requires 10–15 °C	[55]
Relative humidity	Maintains moisture; prevents desiccation	80–95%	Critical for primordia formation and cap expansion	[56]
Ventilation	Supplies O2, removes CO2	Constant renewal	CO2 > 0.1% inhibits fruiting	[57]
Light intensity	Triggers fruiting in photophilic species	200–500 lux (diffuse)	Pleurotus sp. and Lentinula sp. require light for morphogenesis	[58]
Substrate pH	Influences microbial competition and enzyme activity	5.5–7.0	pH < 5 inhibits growth; pH > 7 favors contaminants	[59]

). These parameters vary by species and strongly determine the kinetics of mycelial growth and fruiting body formation.

Large-scale mushroom farming contributes directly to food security and rural development. Rapid growth cycle and substrate versatility enable production in small or urban facilities, promoting distributed protein production with minimal land demand [45]. Moreover, the reuse of agro-industrial by-products reduces methane emissions from residue decomposition and supports sustainable waste management [60]. Nevertheless, the traditional fruiting body system has intrinsic constraints that limit scalability as a global protein source. These include long cultivation cycles, sensitivity to environmental fluctuations, and low protein yield per area and time due to high water content (80–90% in fresh biomass). Additionally, risk of contamination by competing fungi or bacteria remains a bottleneck requiring rigorous sanitation [25]. Hence, the integration of biotechnological production systems, notably mycelial biomass fermentation, offers a complementary and potentially superior pathway to achieve higher productivity, shorter cycles, and greater control, as discussed in the next section.

5. Mycelial Biomass Production: An Emerging Biotechnological Platform

The production of mycelial biomass, the vegetative filamentous network of fungi, represents a major innovation in the generation of fungal proteins and other functional biopolymers. Unlike traditional cultivation systems focused on fruiting bodies, this approach harnesses the continuous growth of mycelium under controlled bioreactor conditions, enabling scalable and standardized production. Recent reviews have emphasized that mycelium-based systems offer higher productivity, shorter growth cycles, and improved process control while aligning with circular economy principles using low-cost agricultural side streams as feedstocks (

Table 4. Differences between fruiting bodies and mycelial biomass [64,65].

Feature	Fruiting Body (Mushroom)	Mycelial Biomass
Structure	Reproductive organ, visible, formed under specific light and temperature conditions.	Filamentous vegetative network of hyphae, continuous growth.
Moisture content	80–90%	60–70% (dry biomass yield higher).
Production cycle	Weeks to months	Few days to 1–2 weeks.
Process control	Semi-open, environmental dependence	Fully closed, bioreactor controlled.
Main applications	Fresh or processed food	Protein ingredient, textured foods, nutraceuticals.
Contamination risk	Moderate to high	Low (sterile systems).

). Mycelial biomass can be used either as a whole protein ingredient or as a precursor for extraction of proteins, polysaccharides, and bioactive compounds [22,61,62,63]. The typical crude protein content ranges from 35–45% (dry weight), exceeding most edible mushrooms and comparable to soy protein isolates [45]. Although both derive from the same fungal organism, fruiting bodies (basidiomata) and mycelium differ markedly in morphology, composition, and production logistics [55]. These characteristics explain why the biotechnological route is increasingly considered the next-generation platform for fungal protein manufacturing, combining speed, safety, and flexibility [33].

Mycelial biomass can be produced under either submerged fermentation (SmF) or solid-state fermentation (SSF), each offering distinct operational and economic characteristics that influence protein yield, composition, and process sustainability. In SmF, fungal cells grow in a fully immersed liquid medium under precisely controlled physicochemical conditions, enabling reproducible scale-up, continuous operation, and integration with automated monitoring systems [66,67,68]. This technology is well-established for industrial enzyme and metabolite production and has recently been adapted for the cultivation of edible and filamentous fungi to obtain uniform, food-grade mycelial biomass [69,70]. However, high energy input for aeration and agitation, water consumption exceeding 90%, and the need for soluble feedstock represent major constraints at large scale.

Conversely, SSF relies on fungal colonization of a moist solid matrix without free water, often using agro-industrial residues such as wheat bran, rice husk, bagasse, coffee husks, or fruit peels [66,71,72]. This system mimics natural fungal habitats and achieves high product concentration, low effluent generation, and superior circular bioeconomy potential [73,74]. Despite limitations in process control, temperature uniformity, and contamination risk, SSF is increasingly attractive for the sustainable production of mycoproteins and co-products such as enzymes, polysaccharides, and organic acids. The complementary features of SmF and SSF have also motivated the development of hybrid fermentation systems that combine efficient oxygen transfer with low resource demand, aligning with the broader goals of circular and low-carbon food production (as illustrated by Figure 1). Bioreactors for SSF, such as tray systems, packed beds, or rotating drums, are now being re-engineered to allow semi-continuous aeration, humidity control, and real-time monitoring of CO₂ evolution and temperature to improve process consistency. Table 5 compares the key operational, economic, and environmental aspects of SmF and SSF, illustrating how each approach offers unique advantages for mycelial biomass cultivation and integration within circular bioprocessing frameworks.

Table 5. Comparative overview of submerged (SmF) and solid-state (SSF) fermentation for mycoprotein production and other bioactives [68,73,75,76].

Aspect	Submerged Fermentation (SmF)	Solid-State Fermentation (SSF)
Medium	Liquid nutrient medium: biomass fully immersed	Moist solid matrix without free liquid
Typical bioreactors (see Figure 4)	Stirred-tank, airlift, or bubble-column reactors with controlled aeration and agitation	Trays, packed-bed, or rotating-drum reactors; rely on passive aeration
Installation cost	High. requires complex, stainless-steel, pressure-resistant systems	Low to moderate—simpler materials and modular configurations
Water consumption	High (90–95% moisture)	Very low (40–70% moisture)
Substrate flexibility	Requires soluble and clarified substrates (e.g., glucose, molasses, corn steep liquor)	Can directly use agro-industrial residues (bran, bagasse, coffee husks, fruit peels)
Oxygenation	Controlled by aeration and agitation; may be limited by viscosity	Passive diffusion through the solid matrix; generally efficient
Process control	Fully automated (pH, DO, temperature, agitation, aeration); supports continuous operation	Limited, but improving with sensor integration (humidity, temperature, CO2)
Scalability	Excellent (industrial fermenters 10–100 m3); high reproducibility	Moderate; scale-up limited by heat and mass transfer gradients
Production cycle	2–10 days depending on strain and process mode	5–15 days depending on substrate and aeration efficiency
Contamination risk	Very low due to sterile, closed operation	Moderate to high in open or semi-open systems
Biomass homogeneity	High; produces uniform, food-grade biomass	Heterogeneous; variable moisture and composition
Energy demand	Higher. Continuous agitation and aeration	Lower. Passive air circulation and minimal mixing
Downstream processing	Biomass easily recovered by filtration or centrifugation; compatible with drying	Complex separation from solid matrix; requires mechanical disruption
Protein yield and composition	25–35% protein (dry weight); composition adjustable via C/N ratio and oxygenation	20–30% protein; enriched in essential amino acids, depending on substrate
Environmental impact	Generates wastewater but can integrate with effluent treatment and biorefineries	Minimal wastewater; valorizes residues, reducing landfill burden
Circular bioeconomy potential	Moderate. Requires feedstock pre-treatment and liquid waste handling	High. Direct valorization of lignocellulosic residues and co-production of enzymes

Table 5. Comparative overview of submerged (SmF) and solid-state (SSF) fermentation for mycoprotein production and other bioactives [68,73,75,76].

Aspect	Submerged Fermentation (SmF)	Solid-State Fermentation (SSF)
Medium	Liquid nutrient medium: biomass fully immersed	Moist solid matrix without free liquid
Typical bioreactors (see Figure 4)	Stirred-tank, airlift, or bubble-column reactors with controlled aeration and agitation	Trays, packed-bed, or rotating-drum reactors; rely on passive aeration
Installation cost	High. requires complex, stainless-steel, pressure-resistant systems	Low to moderate—simpler materials and modular configurations
Water consumption	High (90–95% moisture)	Very low (40–70% moisture)
Substrate flexibility	Requires soluble and clarified substrates (e.g., glucose, molasses, corn steep liquor)	Can directly use agro-industrial residues (bran, bagasse, coffee husks, fruit peels)
Oxygenation	Controlled by aeration and agitation; may be limited by viscosity	Passive diffusion through the solid matrix; generally efficient
Process control	Fully automated (pH, DO, temperature, agitation, aeration); supports continuous operation	Limited, but improving with sensor integration (humidity, temperature, CO2)
Scalability	Excellent (industrial fermenters 10–100 m3); high reproducibility	Moderate; scale-up limited by heat and mass transfer gradients
Production cycle	2–10 days depending on strain and process mode	5–15 days depending on substrate and aeration efficiency
Contamination risk	Very low due to sterile, closed operation	Moderate to high in open or semi-open systems
Biomass homogeneity	High; produces uniform, food-grade biomass	Heterogeneous; variable moisture and composition
Energy demand	Higher. Continuous agitation and aeration	Lower. Passive air circulation and minimal mixing
Downstream processing	Biomass easily recovered by filtration or centrifugation; compatible with drying	Complex separation from solid matrix; requires mechanical disruption
Protein yield and composition	25–35% protein (dry weight); composition adjustable via C/N ratio and oxygenation	20–30% protein; enriched in essential amino acids, depending on substrate
Environmental impact	Generates wastewater but can integrate with effluent treatment and biorefineries	Minimal wastewater; valorizes residues, reducing landfill burden
Circular bioeconomy potential	Moderate. Requires feedstock pre-treatment and liquid waste handling	High. Direct valorization of lignocellulosic residues and co-production of enzymes

Figure 4. Schematic overview of advanced bioreactor configurations and their advantages for fungal mycelial biomass and mycoprotein production.

Figure 4. Schematic overview of advanced bioreactor configurations and their advantages for fungal mycelial biomass and mycoprotein production.

Recent studies have explored hybrid fermentation systems designed to merge the operational precision of SmF with the resource efficiency of SSF. These configurations, such as rotating-drum, trickle-bed, and semi-solid bioreactors (partly illustrated by Figure 1), enable improved oxygen diffusion and moisture control while maintaining moderate energy inputs and low contamination risks [77]. Hybrid setups also facilitate cascading biorefinery concepts, where a single fermentation process yields multiple value-added products, including proteins, enzymes, and polysaccharides, thus enhancing process profitability and circularity [78]. The integration of sensor-based monitoring, AI-driven control, and digital twin modeling (Figure 1) further optimizes these hybrid systems, allowing precise regulation of dissolved oxygen, pH, and nutrient availability in real time [79]. Collectively, these innovations represent the next generation of precision fungal biomanufacturing, bridging the scalability of submerged systems with the sustainability and circularity of solid-state platforms [80,81]. Integration with digital technologies, such as soft-sensor monitoring, real-time control of dissolved oxygen and redox potential, and metabolic modeling, has transformed fungal protein production into a high-precision biomanufacturing platform. These hybrid systems enable cascading valorization, wherein a single process yields multiple products: proteins, enzymes, polysaccharides, and secondary metabolites. When coupled with biorefinery concepts and low-cost lignocellulosic substrates, they provide an efficient route toward sustainable food, feed, and biomaterial production aligned with circular economy goals.

In addition to the choice of fermentation mode (SmF or SSF), mycoprotein-oriented processes require carefully balanced media that provide fermentable carbon, assimilable nitrogen, minerals, vitamins and, in some cases, specific supplements (e.g., trace elements or organic acids) to support fast mycelial growth and high protein yields. For submerged cultivation of basidiomycetes, glucose, sucrose or lignocellulosic hydrolysates are typically combined with complex nitrogen sources such as yeast extract, peptone or corn steep liquor, often at C/N ratios between 10 and 20, together with phosphate and magnesium salts and trace minerals. Representative examples include Pleurotus ostreatus grown in glucose–yeast extract media optimized for protein-rich single-cell protein production, and Pleurotus albidus cultivated in sucrose–yeast extract medium for high mycelial biomass [61]. In SSF, the solid substrate itself provides most of the carbon (e.g., cereal straws, maize stalks or other agro-industrial residues) and is supplemented with cereal brans, simple sugars, urea or ammonium salts, plus mineral additives such as gypsum and lime to adjust structure and pH. Recent studies with Pleurotus and Lentinus spp. on highland barley straw or maize residues have shown that modest supplementation (1–3% glucose, 1–3% urea and 1–2% cereal meals) markedly enhances mycelial growth, crude protein content and digestibility of the fermented matrix, illustrating how nutrient formulation can be tuned to maximize the nutritional quality of the resulting mycoprotein-rich biomass [82]. Some examples of cultivation media are presented in

Table 6. Representative media and supplements used for mycelial biomass production of basidiomycetes in submerged (SmF) and solid-state fermentation (SSF) for mycoprotein applications.

Fermentation Mode/Basidiomycete	Carbon Source/Nitrogen Source(s) and Supplements/Minerals/Other Additives	Ref.
SmF/Pleurotus ostreatus LGAM 1123	Glucose 40–55 g L−1 (or fibre sludge hydrolysate adjusted to ~55 g L−1 glucose). Yeast extract 13–18 g L−1; C/N ≈ 10–15, optimized by RSM for maximal protein production. KH2PO4, MgSO4, trace-element solution; pH 5.0	[83]
SmF/Pleurotus ostreatus (stirred-tank SCP production)	Mixture of glucose and xylose from agro-industrial hydrolysate (≈50–60 g L−1 total sugars). Corn steep liquor (~30–40 g L−1) supplying organic N, vitamins and minerals. Phosphate buffer; trace elements; pH 5.0	[61]
SmF Pleurotus albidus	Sucrose 30 g L−1. Yeast extract 2.5 g L−1 (main N and vitamin source). pH 7.0; standard mineral salts (e.g., KH2PO4, MgSO4)	[84]
SmF Lentinula edodes (multiple isolates)	Coconut water (natural sugars), optionally supplemented with additional glucose (10–20 g L−1). Peptone or yeast extract 2–5 g L−1. Native K, Mg and micronutrients from coconut water; pH 4–7	[85]
SSF Pleurotus spp., Lentinus sajor-caju	Highland barley straw (50 g, 1–2 cm pieces) at 65% moisture. 1% corn meal; 1–3% glucose; 1–3% urea (w/w of dry substrate). 0.5% gypsum; 1% lime; pH adjustment via lime	[82]
SSF Pleurotus pulmonarius	Maize stalk as base substrate. Wheat bran and maize flour (5–20% w/w) as supplementary carbon and nitrogen sources. Moisture adjustment to ~65%; CaCO3 or gypsum for pH and structure	[86]

.

6. Nutritional Quality and Biological Value of Proteins from Mycelial Mass

Edible basidiomycete mycelia (notably Pleurotus, Lentinula, Agaricus) provide complete proteins with all indispensable amino acids (IAA) and favorable digestibility, positioning them alongside high-quality plant and animal proteins by modern metrics of protein quality (PDCAAS/DIAAS). The FAO recommends DIAAS as the preferred regulatory metric and provides IAA reference patterns used to benchmark foods [26]; mycelial biomass typically meets or approaches these patterns, with lysine often abundant and sulfur amino acids supplied at adequate levels in mixed diets. In stirred-tank SmF, Pleurotus ostreatus produced mycelial biomass with high protein yield and a “rich amino acid profile”; mixed C6/C5 sugar feeds (glucose/xylose) improved both biomass and amino-acid composition, supporting use on lignocellulosic hydrolysates [61]. For Lentinula edodes, recent analyses of cultured mycelia report 17 amino acids with sufficient totals of the nine IAAs (≈358–398 mg g⁻¹ protein), indicating high-quality, readily utilizable protein [87]. For Agaricus, classic SmF work scaled to commercial fermenters yielded protein-rich mycelium used to make meat analogues with superior texture/umami versus soy protein, corroborating a balanced amino-acid makeup in practice [88]. Comprehensive mushroom reviews also report protein ranges (≈23–40% DW in basidiomycetes) and EAA completeness relative to plant and animal foods [28].

For edible mycelial mass, SmF in stirred-tank or air-lift bioreactors is preferred due to tighter control and food-grade processability; recent work demonstrates 4-L STBR runs for Pleurotus on wood-hydrolysate and 100-L fed-batch edible mushroom cultures with >22 g L⁻¹ biomass. Air-lift reactors reduce shear on filamentous mycelia and support prolonged/semi-continuous operation; recent reviews compare airlift to stirred-tank and bubble-column designs, and describe pelletization control for better mass transfer [61]. SSF is increasingly employed to cultivate structured, whole-cut mycelium on moist solid substrates, providing desirable texture when the goal is a fibrous mouthfeel rather than dispersed protein. Industrial examples demonstrate the use of SSF to produce whole-cut mycelium-based foods [45]. Under controlled SSF conditions, an edible aerial mycelium can be generated, yielding a cohesive and fibrous biomass suitable for use as a food ingredient. This structured mycelium forms the basis for mycelium-derived foods, including whole-cut products such as mycelium bacon, that reproduce the texture and sensory characteristics of conventional whole-muscle meats. Various Basidiomycetes, particularly Pleurotus and Lentinula species, have been successfully cultivated under these conditions to obtain high-protein, fiber-rich matrices suitable for meat analogues and other sustainable food formulations. Such processes exemplify the integration of fungal biomass engineering and circular bioprocessing for the creation of nutritious and environmentally responsible APs [89].

7. Environmental and Economic Sustainability of Fungal Biomass Production

Fungal biomass production, whether through fruiting-body cultivation or controlled fermentation, provides a low-impact route to high-quality proteins. Numerous LCA studies demonstrate that mycelial protein systems drastically reduce GHG emissions, land occupation, and freshwater consumption when compared with ruminant or monogastric livestock. The substation of 20% of global beef consumption with microbial or fungal proteins could cut agricultural GHG emissions by up to 50% by mid-century [90]. More granular LCAs show that mycoprotein-based foods emit 0.45–1.0 kg CO₂-eq per kg product, compared with 27 kg CO₂-eq per kg beef and 3.3 kg CO₂-eq per kg soy [50]. Additionally, fermentative mycelium systems require more than 90% less land area and 80–90% less water than beef and pork supply chains [91]. Recent syntheses position mycelium as a nutrient-dense ingredient capable of supporting health while enabling regenerative food systems, with promise for protein fortification and texture in plant-forward foods. In particular, fungal fermentation of agro-residues (e.g., lignocellulosic biomass, fruit pomace, spent grains) demonstrates significant substrate flexibility and waste valorization potential [92,93,94]. LCA studies further show that processes based on fungal biomass grown on low-grade feedstocks can achieve lower eutrophication and acidification scores than many traditional protein systems [95]. This circular approach supports in full the multiple United Nations Sustainable Development Goals (SDGs 2, 9, 12, 13, and 15).

Table 7. Comparative environmental indicators for beef, plant-based, and fungal/mycelial proteins.

Indicator	Beef	Plant Proteins (Soy/Pea)	Mycelial/Fungal Proteins	Ref.
GHG emissions (kg CO2-eq·kg−1 product)	27–60	1–5	0.5–1.2	[50,96]
Land use (m2·yr·kg−1 product)	100–300	10–40	2–5	[96]
Water use (L·kg−1 product)	15,000–20,000	2000–4000	300–800	[96]
Energy demand (MJ·kg−1 product)	350–400	60–90	50–70	[45,65]
Residue valorization potential	Minimal	Partial	Full (cascading reuse)	[78]
Scenario: 20% beef replacement by microbial protein	—	—	↓ GHG and deforestation ≈ 50% by 2050	[90]

summarizes comparative LCA indicators for beef, plant-based, and fungal/mycelial protein systems. The data highlights the substantial environmental advantages of fungal biomass production relative to conventional livestock and even to soy or pea protein isolates. Mycoprotein products typically emit between 0.5 and 1.2 kg CO₂-eq kg⁻¹, compared with 27–60 kg CO₂-eq kg⁻¹ for beef and 1–5 kg CO₂-eq kg⁻¹ for plant proteins. These fermentation-based systems also require more than 90% less land and 80–90% less water than beef or pork production chains, while maintaining moderate energy demands. In addition to the favorable LCA profile, fungal processes enable cascading residue valorization, converting low-value lignocellulosic materials into proteins, enzymes, and polysaccharides within a circular bioeconomy framework. Collectively, these metrics reinforce the potential of mycelial proteins as sustainable, resource-efficient alternatives to animal-derived proteins, contributing to climate-mitigation targets and several UN Sustainable Development Goals.

The global alternative-protein market was valued at USD 15.3 billion in 2023 and is projected to reach USD 26.5 billion by 2030. Within this space, fungal and microbial proteins are among the fastest-growing segments [97]. Mycelial biomass production offers a compelling techno-economic balance, lower feedstock cost (due to waste-substrate utilization) and high yield per m³ reactor. Pilot techno-economic analyses suggest production costs of approximately USD 3.55 kg⁻¹ of dry fungal biomass, comparable to plant-based protein isolates [85]. Reported benchmarks indicate that soy protein isolates are priced around USD 3 kg⁻¹ [98], while protein isolates from legumes such as peas generally range between USD 4–5 kg⁻¹ depending on purity and extraction method [99]. Modular bioreactor installations and co-location within existing food-processing plants further reduce logistics and energy overhead. From a policy standpoint, recent EU initiatives, especially the Commission Communication “Boosting Biotechnology and Biomanufacturing in the EU” (2024) and the CBE JU calls (2024–2025), explicitly support the deployment of bio-based and mycelial processes using residues and side-streams within regional agro-industrial clusters, strengthening circular value chains [100,101].

Integrating fungal fermentation within biorefinery frameworks maximizes resource efficiency. Residual solids from fermentation can serve as soil conditioners or feed additives, while process water and CO₂ streams are recycled to subsequent cultivation batches. Hybrid bioprocesses co-producing proteins, enzymes, and polysaccharides (β-glucans, chitin derivatives) increase profitability while minimizing waste. Overall, the combination of low environmental footprint, competitive economics, and policy alignment positions mycelial biomass as a strategic enabler of circular food systems capable of decoupling protein supply from conventional agriculture [80,102].

8. Emerging Basidiomycetes Species for Protein-Rich Mycelial Biomass

While Pleurotus ostreatus, Lentinula edodes, and Ganoderma lucidum remain dominant in fungal biotechnology, several non-conventional basidiomycetes and ascomycetes have emerged as promising candidates for large-scale mycelial protein production. These species combine high protein content with functional properties such as emulsifying capacity, gelling ability, and antioxidant activity. Recent studies emphasize that expanding the taxonomic diversity of edible fungi is essential to ensure resilient and regionally adaptable protein sources, especially those compatible with low-cost agro-industrial residues as substrates.

Expanding the diversity of fungal species used for mycelial biomass production supports regionalized protein supply chains and stimulates innovation in functional foods. Each species exhibits distinctive biochemical and rheological traits, broadening potential applications that range from protein enrichment to natural emulsifiers and gelling agents. Recent technological developments have enabled dual-purpose fermentations, in which nutritional biomass and high-value metabolites (e.g., pleuromutilin, cordycepin, β-glucans) are co-produced, improving process economics and reinforcing the circular biorefinery model [65].

Table 8. Under-explored basidiomycetes that show promise as alternative mycoprotein sources via submerged fermentation (SmF) or solid-state fermentation (SSF).

Basidiomycete Species Type/Common Name	Key Findings in SmF or SSF	Protein/Biomass Yield Distinct Nutritional or Functional Traits	Reference
Trametes versicolor White-rot fungus (“turkey tail”)	Efficient growth in SmF on grape pomace extract; characterized amino acid profile; EPS and protein quantified.	21 g/L biomass (grape pomace medium) Balanced essential amino acids; β-glucans; antioxidant mycelium.	[16,103]
Schizophyllum commune Split-gill mushroom	Statistical optimization of SmF for mycelial biomass and polysaccharides; robust nutrient conversion on agro-residues.	Up to 17 g/L dry biomass; 38% protein (dry basis) High β-glucan and hydrophobin content; strong umami potential.	[104]
Cyclocybe (Agrocybe) aegerita. Black poplar mushroom	Liquid culture and side-stream upcycling studies; nutrient-rich mycelium.	15–20 g/L biomass in optimized media Edible, mild flavor, rich in lysine and leucine.	[105]
Coprinus comatus Shaggy mane mushroom	SmF versus fruit body comparison; biochemical composition and amino-acid profile analyzed.	16 g/L biomass; 40% protein (dry basis). Rapid growth; savory profile; antioxidant enzymes.	[106]
Polyporus (Neolentinus) tricholoma. Wood-degrading basidiomycete	Stirred-tank SmF optimization for growth and metabolites.	10–14 g/L biomass. Tolerant to low-cost lignocellulosic substrates; promising amino-acid profile.	[107]
Auricularia polytricha (“wood ear mushroom”)	SmF versus fruiting body comparison. Proteomic studies revealed that its protein, polysaccharide complexes possess superior gelation and emulsifying capacities, ideal for plant-based meats and soups.	Combines 15–25% protein with soluble fibers and bioactive polysaccharides	[74]

exemplifies the growing potential of underexplored fungi in sustainable protein production. Collectively, these underexplored fungi demonstrate high nutritional quality, versatile bioprocess adaptability, and compatibility with agro-residue-based substrates, strengthening the role of fungal biotechnology in sustainable protein production and circular food systems.

9. Future Perspectives and Challenges

Modern strain-engineering strategies, supported by integrative omics, genomics, transcriptomics, proteomics, and metabolomics, are reshaping fungal biotechnology. Whole-genome sequencing and system-level analyses in key basidiomycetes have elucidated the molecular networks governing amino acid biosynthesis, carbon utilization, and environmental adaptation. In Pleurotus ostreatus, genome-wide and transcriptomic studies identified extensive gene families involved in lignocellulose breakdown, nitrogen metabolism, and oxidative defense [108]. The Ganoderma lucidum genome revealed central pathways for triterpenoid formation and stress tolerance [18], while comparative omics in Schizophyllum commune demonstrated coordinated regulation of carbohydrate and amino acid metabolism during development [109]. Insights from Hericium erinaceus expanded this perspective: its integrated genome–transcriptome map uncovered the molecular basis of polysaccharide and amino acid synthesis [110], and multi-omics coupling of transcriptome and metabolome profiling revealed carbon-flux reprogramming and glutamate-centered metabolic modules under varying substrates [111]. Collectively, these advances in omics and systems biology enable rational strain design and targeted metabolic engineering to boost protein productivity, enhance stress resilience, and broaden substrate adaptability, key attributes for sustainable mycoprotein and fungal biorefinery platforms.

Recent innovations in bioreactor engineering, such as air-lift, bubble-column, and stirred-tank configurations, have advanced scalable, cost-efficient fungal biomass production for food and industrial use. Airlift and bubble-column systems provide superior gas exchange and reduced shear relative to conventional stirred tanks, fostering robust mycelial growth in SmF [102]. Emerging designs, including rotating-drum, moist-air, and semi-solid bioreactors (Figure 1), aim to merge the aeration efficiency of liquid systems with the resource economy of solid-state processes [112]. Continuous, non-invasive monitoring of dissolved oxygen, pH, and CO₂, coupled with feedback-control algorithms, ensures process stability and consistent yields.

The integration of digital-twin technology and AI-driven fermentation modeling has further optimized biomass kinetics, nutrient flow, and energy utilization, accelerating process intensification (Figure 1) [79]. Complementary approaches, such as immobilized fungal cultures, cell-recycle fermentation, and microbubble aeration, are increasingly adopted to improve oxygen delivery and substrate conversion while reducing operational costs and environmental impacts [45,113].

The global landscape for regulatory approval of fungal-based proteins is rapidly evolving. Several products derived from mycelial biomass have already achieved GRAS status in the United States [114], while the European Food Safety Authority (EFSA) has granted Novel Food authorization for Neurospora intermedia and Fusarium venenatum-based ingredients [115]. In 2025, the EFSA initiated a harmonized risk assessment framework for “novel microbial biomass foods,” covering compositional, allergenic, and toxicological evaluations [116]. These advances lower barriers for commercial adoption of mycelial biomass in functional foods, beverages, and nutraceuticals.

From the market perspective, consumer acceptance of mycelium-based products continues to improve, driven by sensory similarity to meat, clean-label perception, and sustainability narratives [49]. Nevertheless, communication strategies should emphasize safety, traceability, and local production to reinforce consumer trust.

The convergence of fungal and plant-protein technologies is driving a hybrid-protein paradigm. Recent work shows that edible mycelium can act as an inactivated or living scaffold for anchorage-dependent animal cells, offering both structural support and nutritional value. For example, heat-treated Aspergillus oryzae mycelium pellets successfully served as a scaffold, supporting the proliferation and differentiation of muscle precursor cells (C2C12) and bovine satellite cells in vitro [117]. Because the microcarrier remains edible and is incorporated into the final product, it removes the need for carrier-removal steps, thereby simplifying the bioprocess and improving sustainability [103].

Mycelial proteins can complement plant isolates to improve amino acid balance, texture, and binding functionality in food matrices. Hybrid systems that integrate mycelial scaffolding into plant-based meats or dairy analogues deliver superior mouthfeel and nutritional density, while reducing ingredient complexity [118]. This trend aligns with the principles of food system circularity, where multiple biological streams, plants, fungi, and microalgae, are integrated to minimize waste and maximize nutritional output. Emerging research also explores 3D food printing with fungal proteins, enabling customizable textures and nutrient profiles [111]. These technologies bridge sustainability and innovation, paving the way for next-generation functional foods.

Although remarkable progress has been achieved, several technical and socioeconomic barriers still limit the large-scale production and market adoption of mycoproteins. Differences in strain behavior, such as growth kinetics and metabolite profiles, make it difficult to ensure consistent process performance [67]. Rheological challenges, including high broth viscosity and oxygen transfer limitations, continue to affect bioreactor efficiency and downstream processing [102,119]. From a market perspective, many consumers still perceive fungal biomass as a “mushroom extract” rather than a genuine protein source, which slows public acceptance and demand [120]. Regulatory discrepancies, particularly across Novel Food approval systems in different regions, further delay product commercialization [121]. Moreover, additional in vivo research on digestibility, nutrient absorption, and long-term health outcomes is essential to support nutritional and safety claims [65]. Addressing these interconnected challenges will require collaboration across disciplines, linking mycology, bioprocess engineering, computational modeling, food science, and policy, to realize the full potential of mycelial proteins in future sustainable diets.

10. Conclusions

The transition toward sustainable and resilient food systems requires diversified protein sources that minimize environmental impact while ensuring nutritional adequacy. In this context, mushrooms and mycelial biomass from basidiomycetes represent one of the most promising biotechnological routes to produce APs at scale. This review demonstrates that fungal proteins provide balanced EAA profiles, high digestibility, and additional health-promoting compounds such as β-glucans and phenolic conjugates. Beyond nutritional aspects, fungal bioprocesses offer an exceptional environmental advantage: drastically lower greenhouse gas emissions, land use, and water consumption compared with conventional livestock, while valorizing agricultural residues and integrating into circular bioeconomy models. Traditional mushroom cultivation continues to play an important role in food security and rural income, yet bioreactor-based mycelial biomass production enables faster, controlled, and scalable systems capable of meeting industrial demand. Emerging fungal species, including Schizophyllum commune and Auricularia polytricha, expand the spectrum of high-protein, functional ingredients suited for various food matrices. Recent progress in omics-guided strain engineering, bioreactor automation, and hybrid protein formulations consolidates fungi as a central pillar of the ongoing protein transition. However, challenges remain in large-scale process optimization, regulatory harmonization, and consumer communication. Continued interdisciplinary research and policy support will be essential to translate the full potential of fungal biomanufacturing into commercially viable and nutritionally impactful solutions. Ultimately, integrating mycelial protein production within circular biorefineries and sustainable food systems can contribute substantially to global climate goals, resource efficiency, and health-oriented innovation, positioning fungi not merely as an alternative but as a cornerstone of the future food landscape.