Harnessing mushrooms for poultry nutrition: Boosting health, immunity, and productivity

Abstract

Edible mushrooms, the fruiting bodies of fungi, are emerging as promising functional feed additives in poultry nutrition, offering a sustainable alternative to synthetic growth promoters. Rich in bioactive compounds such as β-glucans, glycoproteins, polyphenols, terpenoids, and antioxidants, mushrooms exhibit immunomodulatory, antimicrobial, anti-inflammatory, and antioxidative properties. These compounds support enhanced immune function, metabolic regulation, oxidative stress reduction, and disease resistance in poultry. Mushroom species like Pleurotus ostreatus, Agaricus bisporus, and Flammulina velutipes have demonstrated improvements in growth rate, feed conversion ratio (FCR), carcass traits, meat quality, and egg production. Importantly, mushroom stems, often discarded as waste, are gaining attention for their nutritional and functional value, making them a cost-effective and eco-friendly feed resource. Medicinal mushrooms such as Lentinula edodes, Ganoderma lucidum, and Trametes versicolor further enhance antioxidant defense and gut microbial balance. This review synthesizes current evidence on the application of mushrooms and their stems in poultry diets, emphasizing their nutritional roles, physiological impacts, and underlying mechanisms. Moreover, it discusses their relevance in promoting sustainable poultry production by reducing antibiotic use and valorizing agricultural by-products. Future research should aim to standardize species selection, dosage levels, and feeding durations while assessing long-term safety and economic feasibility. Unlocking their full potential could transform poultry nutrition toward a more resilient and environmentally responsible model.

Graphical abstract

Introduction

Growing global concern over antibiotic resistance, sustainability, and the demand for natural additives in poultry production has led researchers to seek alternative feed ingredients that enhance health and productivity without relying on synthetic growth promoters (Ayalew et al., 2022; Salahuddin et al., 2024; Saeed et al., 2025). In this regard, mushrooms also known as toadstool are the fruiting body of species of fungus have emerged as a promising option. Rich in bioactive compounds such as β-glucans, phenolic compounds, and polysaccharides, edible and medicinal mushrooms has demonstrated a range of health-promoting properties, including antioxidant, immunomodulatory, and antimicrobial effects (Guo et al., 2004; Kupradit et al., 2023; Flores et al., 2024; Navarro-Simarro et al., 2024). These natural benefits can position mushrooms as an ideal replacement for conventional additives, offering multiple advantages for poultry health and promoting environmental sustainability in poultry farming.

Incorporating mushroom dry powder into poultry diets has been demonstrated to boost growth, optimize feed utilization, and strengthen immune responses. Mushroom produced using fungal species such as P. ostreatus and A. bisporus have been shown to improve feed conversion ratios (FCR) and support better nutrient absorption, leading to increased growth rates in poultry (Giannenas et al., 2011; Suberu et al., 2024; Bormon et al., 2024). Additionally, these mushrooms have a positive impact on gut health by fostering the growth of beneficial microbiota and reducing the presence of pathogenic bacteria in the intestines (Giannenas et al., 2011; Toghyani et al., 2012; Törős et al., 2024). These effects are valuable as the poultry industry seeks sustainable alternatives to antibiotics, addressing global concern over antibiotic resistance.

In addition to enhancing growth performance and gut health, mushroom supplementation has been associated with improvements in egg and meat quality, two crucial factors in poultry production. Research on F. velutipes and other mushroom species has demonstrated enhancements in meat tenderness, increased egg production, and improved antioxidant status in poultry (Mahfuz et al., 2018a, 2019a, 2020; Sun et al., 2023; Suberu et al., 2024). These benefits are largely attributed to the bioactive compounds β-glucans, glycoproteins, terpenoids, sesquiterpenes, sterols and polyphenolic compounds in mushrooms, which enhances metabolic processes and provide protection against oxidative stress (Giannenas et al., 2010; Kumar et al., 2021). As consumer demand for high-quality poultry products continues to rise, mushrooms offer a natural way to enhance both the nutritional and sensory qualities of meat and eggs.

Mushroom stems, often discarded as waste, present a sustainable and nutritious feed option due to their high protein, fiber, and mineral content. Stems from mushroom species like A. bisporus, P. ostreatus, and L. edodes have been shown to improve growth, digestion, and immunity in poultry (Buwjoom et al., 2004; Chang and Miles, 2004; Wasser, 2014). Their use can help reduce reliance on synthetic additives and antibiotics by enhancing natural immunity and pathogen resistance. This review provides an in-depth analysis of current research on the effects of mushroom and its byproducts supplementation in poultry diets, highlighting its potential to improve growth, boost immunity, and enhance product quality. By compiling findings from various studies, this review identifies key benefits of mushrooms and underlines gaps in research, suggesting future studies should focus on optimizing species, dosages, and the long-term effects of supplementation to transform poultry nutrition, promoting more natural, sustainable feeding practices and contributing to an eco-friendly, health-conscious poultry production system.

Mushroom varieties and characteristics

Mushroom species are remarkably diverse, each with distinct traits that make them fascinating to study. Common varieties of mushrooms are depicted in Fig. 1. Mushrooms are spore-bearing fungi belonging to the Basidiomycota phylum. They have a distinctive structure with a cap (pileus) and stalk (stipe) and reproduce through both sexual and asexual methods. Sexual reproduction involves the release of spores, while asexual reproduction occurs via fragmentation or budding, allowing for new fungal growth (Singara, 2015; Vreeburg et al., 2020).

Fig. 1.

This image illustrates a variety of mushrooms showcasing their diversity.

A. bisporus, commonly known as the button mushroom, ranges in colors from white to light brown (Paulauskienė et al., 2020). Often found in grocery stores, it has a smooth, round cap and firm texture. As it matures, the cap darkens and opens, revealing gills that vary from white to light brown. This mushroom species is widely cultivated for its mild flavor and versatility in cooking (Royse et al., 2017). In contrast to other mushrooms, Amanita muscaria (fly agaric) is easily recognized by its bright red cap covered in white warts, serving as a warning of its toxicity. This mushroom is famous in folklore but dangerous to consume due to its toxic properties (O’Reilly and McDonald, 2002). On the other hand, P. ostreatus (oyster mushroom) is valued for its culinary use and ease of cultivation. Its broad, fan-shaped cap resembles an oyster, and its decurrent gills extend down the stem. Oyster mushrooms come in a range of colors, from white to grey, pink, and yellow, and can grow on various organic materials, making them popular among foragers (Chang and Miles, 2004).

Mushrooms exhibit a wide range of physical traits such as size, color, and texture, contributing to the diversity of mushroom species. Mushrooms are categorized into four main types based on their ecological roles: mycorrhizal, saprophytic, parasitic, and endophytic (Mlambo and Maphosa, 2022). Mycorrhizal mushrooms, like Boletus edulis (Porcini), form mutualistic relationships with plant roots, aiding nutrient absorption in exchange for carbohydrates (Stajich et al., 2009). Saprophytic mushrooms, such as A. bisporus and P. ostreatus, decompose organic matter, recycling nutrients back into the ecosystem (Purahong et al., 2016). Endophytic fungi, including Cordyceps sinensis (Caterpillar mushrooms) and T. versicolor (Turkey tail), live inside plant tissues without harming the host, often providing mutual benefits (Hardoim et al., 2015). Parasitic mushrooms, like Honey fungi and Aspergillus flavus, feed on living organisms, causing harm or disease (Sharon and Shlezinger, 2013). Understanding these classifications is key to advancing research and applications in agriculture, sustainability, and industries like poultry, where mushrooms' nutritional and chemical properties offer valuable benefits.

Chemical composition and medicinal benefits of mushrooms

Mushrooms possess a complex chemical composition that underpins their numerous health benefits, as given in Table 1. The proximate composition of the caps and stems of various mushroom species reveals significant differences in their nutritional profiles. For instance, the caps of Pleurotus sp. contain 85.5 g/100 g moisture and a notably high protein content of 37.8 g/100 g, whereas the stems exhibit slightly lower moisture content (84.4 g/100 g) and a substantially reduced protein level (15.3 g/100 g) (Barroso et al., 2020). This emphasizes that the caps of Pleurotus sp. are a superior protein source compared to their stems. Similarly, A. bisporus demonstrates consistent moisture content in both caps and stems (approximately 87 g/100 g), yet the protein content is higher in the caps (29.6 g/100 g) than in the stems (23.5 g/100 g) (Hola et al., 2023).

Table 1.

Proximate composition of cap and stem of different mushroom species.

	Moisture	Protein	Lipid	Carbohydrate	Fiber	Ash	Moisture	Protein	Lipid	Carbohydrate	Fiber	Ash	References.
	Caps (g/100 g)						Stem (g/100 g)
Pleurotus spp.	85.54	37.81	0.15	nd	nd	9.21	84.43	15.31	0.19	nd	nd	6.78	Barroso et al., 2020
Agaricus bisporus	87.10	29.56	0.23	nd	nd	10.97	87.05	23.51	0.40	nd	nd	9.54
Agaricus bisporus	86.45	24.01	0.21	nd	nd	13.70	88.93	22.40	0.13	nd	nd	11.56
Lentinula edodes	89.44	20.43	0.13	nd	nd	6.23	80.99	15.57	0.07	nd	nd	6.49
Agaricus sinodeliciosus	89.35	28.93	1.22*	46.41	13.50	9.94	89.07	27.65	5.92*	38.00	18.61	9.82	Hola et al., 2023
Agaricus qilianensis	91.39	20.12	1.24*	60.99	9.91	7.74	90.13	23.05	6.28*	47.71	16.55	6.41
Agaricus sinodeliciosus	92.89	14.52	1.27*	71.54	6.93	5.74	93.16	11.54	2.10*	72.95	8.21	5.21
Agaricus qilianensis	88.97	28.63	6.53*	42.05	13.10	9.70	88.47	30.41	2.11*	39.45	16.70	11.34
Lentinus edodes⁎⁎	8.56	16.71	2.67	nd	19.94	5.89	9.55	7.20	1.37*	nd	13.86	5.04	Zhang et al., 2012

Abbreviations: nd, not determined.

^⁎fat.

^⁎⁎milled powder.

L. edodes (shiitake mushrooms) also exhibit variations, with the caps having higher moisture (89.4 g/100 g) and protein content (20.4 g/100 g) than the stems, which contain 81 g/100 g moisture and 15.6 g/100 g protein. These findings indicate that the caps of L. edodes are more nutrient-denser compared to the stems. Contrarily, species such as Agaricus sinodeliciosus and Agaricus qilianensis show significant differences in lipid content. The caps of A. sinodeliciosus contain 1.2 g/100 g of lipids, while the stems contain a considerably higher lipid content of 5.9 g/100 g (Zhang et al., 2012).

Carbohydrate and fiber contents also vary across species and between caps and stems. For instance, A. qilianensis caps have a high carbohydrate content of 61 g/100 g and a fiber content of 9.9 g/100 g, while the stems contain slightly less carbohydrate (47.7 g/100 g) but more fiber (16.6 g/100 g) (Zhang et al., 2012). Mushrooms' chemical composition varies by species and growth conditions, with caps typically containing more carbohydrates and phenolics, whereas stems provide higher fiber content. Both parts deliver substantial protein and antioxidant activity (Kupradit et al., 2023). These variations underscore the diverse nutritional profiles of different mushroom species and their respective parts, offering valuable insights for dietary planning and nutritional studies.

Mushrooms have been integral to traditional medicine, and modern research continues to uncover their therapeutic potential. Table 2 provides a detailed overview of the health benefits associated with various mushroom species, emphasizing the bioactive compounds responsible for their medicinal properties. T. versicolor (Turkey Tail) contains polysaccharide-K (PSK), which has demonstrated efficacy in cancer therapy and immune support (Habtemariam, 2020). Inonotus obliquus (Chaga) harvested from Birch trees is valued for its antioxidant, anticancer, and anti-inflammatory properties, while C. sinensis has been studied for its ability to enhance athletic performance and combat fatigue (Wasser, 2014; Camilleri et al., 2024). Key bioactive compounds in mushrooms include polysaccharides, proteins, vitamins, minerals, and polyphenols, which collectively contribute to their antioxidant, anti-inflammatory, and anticancer effects (Kour et al., 2022: Kupradit et al., 2023). These properties can also enhance immunity and disease resistance in poultry.

Table 2.

Different types of mushrooms and their bioactive properties.

Species	Bioactive Compounds	Benefits	Reference
Agaricus Bisporus	Pyrogallol hydroxybenzoic acid derivatives, Flavonoids	Anti-inflammatory	Moro et al., 2012
Agaricus campestris	Glycoproteins	Antitumor	Chang, 1993
Auricularia polytricha	Adenosine	Artherosclerotic activity	Sagar et al., 2007
Cordyceps militaris	Cordycepin, Cordymin	Anti-inflammatory, Anti-angiogenic, Anticancer,	Won and Park, 2005; Das et al., 2010; Wong et al., 2011
Flammulina velutipes	Polysaccharides, bioactive protein, sterols, polypeptide, ergothioneine, cuparene, sesquiterpenes, dilinolein, hemisceramide	Immunomodulatory, antitumor, antioxidant, hepatoprotection, anti-hyperlipidemic	Zhang et al., 2010
Ganoderma lucidum	Polysaccharides, Ganopoly, Ganoderans, GLP	Immunomodulatory, Antioxidant, Hepatoprotective, gut microbial modulation	Rai et al., 2005; Shi et al., 2013; Jayachandran et al., 2017; Ahmad, 2019
Hericium erinaceus	Polysaccharide A and B (HPA and HPB), Exopolymer	Anticancer, Reduced LDL, HDL, total cholesterol, phospholipids, triglyceride, and more	Yang et al., 2022
Inonotus obliquus	3β‑hydroxy-lan sta-8, 24‑dien-21-al, inotodiol and lanosterol	Anticancer	Chung et al., 2010
Lactarius deliciosus	Pyrogallol, Flavonoids	Anti-inflammatory	Moro et al., 2012
Lentinula Edodes	Lentin, Fucomannogalactan, Phenolic compounds and flavonoids	Antifungal, Anti-inflammatory, Antibacterial and antioxidant	Ngai and Ng, 2003; Attarat and PhermthaI, 2015; Chowdhury et al., 2015
Pleurotus ostreatus	β-glucans, α-glucan, Proteoglycans, Pleuran, Lovastatin	Anticancer, Antitumor, Antibacterial, Antihypercholesterolemic	Karacsonyi and Kuniak, 1994; Sarangi et al., 2006; Alam et al., 2009; Jedinak et al., 2010; Wu et al., 2011

Abbreviations: LDL, Low-density lipoprotein; HDL, High-density lipoprotein; GLP, Ganoderma lucidum polysaccharide.

A study on I. obliquus (Chaga) showed its potential as a natural anticancer agent due to compounds such as 3β‑hydroxy-lanosta-8,24‑dien-21-al, inotodiol, and lanosterol, which effectively inhibit cancer cell growth (Chung et al., 2010). In addition to its anticancer properties, chaga shows strong antioxidant activity, neutralizing free radicals, reducing oxidative stress, and lowering the risk of chronic diseases. Its β-glucans enhance immune function, bolstering the body’s defense against infections. Similarly, C. militaris has shown antitumor and anticancer effects, primarily attributed to bioactive compounds like cordycepin, which induces apoptosis in cancer cells and inhibits their proliferation. This species also exhibits anti-inflammatory effects and has been linked to improved sexual health, including enhanced testosterone production and vitality (Pohsa et al., 2020; Kusama et al., 2020).

Polysaccharides such as β-glucans play a significant role in the health benefits of mushrooms, particularly in poultry nutrition (Venturella et al., 2021). Studies have demonstrated that mushrooms possess a wide array of beneficial properties, including anticancer, antitumor, anti-inflammatory, immunomodulatory, antifungal, antidiabetic, and antioxidant effects. Additionally, they support gut microbial health (Kour et al., 2022), modulate the immune system, and strengthen the body’s defense against pathogens (Xu et al., 2023).

G. lucidum (Reishi) is notable for its high triterpene content, which promotes cell apoptosis and suppresses tumor metastasis, contributing to its anticancer effects. Its polysaccharides enhance immune response, further supporting its potential in cancer treatment. (Kachari, 2024). Moreover, essential nutrients such as B vitamins (riboflavin, niacin, and pantothenic acid) and minerals (selenium, potassium, and copper) further enhance the health-promoting properties of mushrooms, supporting overall well-being. The chemical composition and bioactive compounds of mushrooms underscore their nutritional and medicinal significance. These findings highlight the potential applications of mushrooms in human and animal health, including their role in supporting immunity, combating diseases, and promoting overall health.

Nutritional values of mushrooms

Mushrooms are highly versatile food, celebrated for their unique flavor and rich nutritional values. They are an excellent source of essential nutrients, including carbohydrates, amino acids, vitamins, and minerals that promote overall health. Composed of about 90 % water and 10 % dry matter, mushrooms are high in protein and low in fat (Assemie and Abaya, 2022). Mushroom stems, often considered a byproduct of cultivation, show great potential as an ingredient in poultry feed due to their rich nutrient density. The benefits of including mushroom stem powder in diets are presented in Fig. 2. Stems from A. bisporus (button mushroom) and P. ostreatus (oyster mushroom) are particularly valuable, with protein content ranging from 15 % to 20 %, support muscle growth and health in birds. Additionally, the fiber content in these stems promote better digestion and feed efficiency, reducing the need for other fiber sources in the poultry diet (Chang and Miles, 2004).

Fig. 2.

Schematic representation of mushroom stem powder as a feed additive in poultry diets, illustrating its potential benefits, including better nutrient digestion, effective nutrient absorption, enhanced immune response, promotion of muscle growth, improved feed efficiency, and reduced disease occurrence. This figure is created by www.biorender.com.

Stems from L. edodes (Shiitake) provide additional benefits due to their unique biochemical makeup. Rich in β-glucans and other polysaccharides, these stems are known for their immune-boosting properties, potentially enhancing poultry’s immune response and resilience to diseases, thereby reducing the need for veterinary interventions. Moreover, shiitake stems contain less lignin than other mushroom stems, making them more digestible and suited for poultry feed, ensuring more effective nutrient absorption (Buwjoom et al., 2004).

The protein content of mushrooms varies based on factors such as species, strain, growing conditions, and substrate (Assemie and Abaya, 2022; Ayimbila and Keawsompong, 2023). On a dry basis, mushrooms contain 6.6 % to 36.9 % protein, with an average of 23.8 %, making them a cost-effective alternative to animal and plant protein sources (Zhou et al., 2020; Kaur et al., 2022). Additionally, mushrooms are rich in all nine essential amino acids, low in calories, and cholesterol-free. Edible fungi often grow on lignocellulose-rich agricultural waste, and their enzymes enhance substrate digestibility and protein content, making them valuable for animal feed (Qin et al., 2023).

Carbohydrates are a major component of the fungal cell wall, comprising about 40-50 % of the mushroom's biomass (Vetter, 2023). The total carbohydrate content in mushrooms varies between 35 % and 70 % of their dry weight (Cheung, 2013), and they are rich in dietary fiber, particularly in the stems, which contain both soluble and insoluble fibers (Assemie and Abaya, 2022). Mushroom polysaccharides are classified into homopolysaccharides, such as glucans, chitin, glycogen, and heteropolysaccharides. Glucans boost immunity, while chitin aids digestion and supports immune health (Zhong et al., 2023; Ullah et al., 2019). Glycogen serves as an energy reserve. Heteropolysaccharides, composed of different monosaccharides, contribute to mushrooms' antioxidant, anti-inflammatory, and anticancer properties (Bhambri et al., 2022). The fiber and polysaccharides in mushrooms promote gut health, regulate blood sugar, lower cholesterol, and may help prevent chronic diseases like diabetes and heart disease (Guan et al., 2021; Yu et al., 2023).

Mushrooms are rich in B vitamins, including thiamine (B1), riboflavin (B2), niacin (B3), pantothenic acid (B5), pyridoxine (B6), biotin (B7), and folic acid (B9), which are crucial for energy production, DNA synthesis, and nerve function (Wani et al., 2010; Lesa et al., 2022). Notably, mushrooms have high riboflavin content, often surpassing that of most vegetables, making them a valuable component of plant-based diets (Gupta et al., 2018). They are also a rare non-animal source of vitamin D, which is essential for bone health and can be increased through UV exposure (Assemie and Abaya, 2022). However, the vitamin composition of mushrooms varies by species, with A. bisporus featuring a well-balanced vitamin profile, while P. ostreatus stands out for its higher levels of folic acid, vitamin C, and niacin. This underlines the nutritional differences between the two species, with oyster mushrooms providing unique benefits for immunity and energy metabolism (Iqbal et al., 2024).

Mushrooms are also rich in essential minerals like potassium, iron, selenium, copper, zinc, calcium, sodium, and phosphorus (Won and Park, 2005). Potassium and phosphorus are especially abundant, supporting heart health, fluid balance, and cellular function (Udensi and Tchounwou, 2017; Haro et al., 2020). Selenium acts as a powerful antioxidant, aiding immune function and thyroid health (Podkowa et al., 2021). Iron is critical for hemoglobin production, while zinc supports immune function and DNA repair. Copper contributes to cardiovascular and bone health, and calcium plays a role in muscle function and blood clotting (Ciosek et al., 2021). Together, these minerals enhance the overall nutritional value of mushrooms, supporting essential physiological processes.

Mushrooms are a rich source of lipids, including a variety of fatty acids, sterols, and other beneficial compounds. Despite their low-fat content (about 5.7 % of dry matter), they are high in polyunsaturated fats (Díez and Alvarez, 2001). Key fatty acids found in mushrooms include linoleic acid (C18:2), oleic acid (C18:1), and palmitoleic acid (C16:1), all essential for maintaining good health (Guillamón et al., 2010). Linoleic acid is particularly important for lowering cholesterol, reducing the risk of cardiovascular disease (Valverde et al., 2015), while oleic acid helps reduce inflammation and improve lipid profiles (Piccinin et al., 2019). Palmitoleic acid also offers potential metabolic benefits (Murru et al., 2022).

The lipid profile of mushrooms varies by species and growing conditions. For example, L. edodes (shiitake) are high in unsaturated fatty acids, supporting heart health, while G. lucidum (Reishi) is rich in sterols and triterpenes known for their immunomodulatory and anti-inflammatory effects (Cadar et al., 2023). Mushrooms also contain cis-linoleic acid, essential for healthy cell membranes and immune function, as well as polyunsaturated fats that help lower LDL cholesterol, promoting heart health and weight management (Sande et al., 2019). Including mushrooms in a balanced diet can contribute to overall well-being, making them an excellent nutritional choice.

Mechanisms underlying the health benefits of mushrooms

Mushrooms have long been recognized for their therapeutic properties, particularly due to the diverse range of bioactive compounds they contain. Each compound contributes to the mushrooms' health benefits through specific mechanisms. The antioxidant properties of mushrooms are primarily attributed to phenolic compounds, which include flavonoids, phenolic acids, and other polyphenolic substances (Ahmad et al., 2023). These antioxidants neutralize free radicals, reducing oxidative stress, and preventing cellular damage. Mushrooms such as G. lucidum (Reishi) have demonstrated high levels of bioactive compounds, which are essential in the prevention and management of chronic diseases, including cancer, cardiovascular disease, and neurodegenerative disorders (Kachari, 2024). Mushrooms represent a potential alternative source of novel antimicrobial compounds, particularly secondary metabolites like benzoic acid derivatives, quinolones, steroids, terpenes, anthraquinones, and others (Sułkowska-Ziaja et al., 2023). Additionally, primary metabolites such as proteins, oxalic acid, and peptides also contribute to their antimicrobial properties (Alves et al., 2012). Zhong and Xiao (2009) demonstrated that mushrooms produce an abundance of secondary metabolites, including but not limited to phenolic compounds, terpenes, polyketides, and steroids, as a result of their metabolic processes. Many of these secondary metabolites exhibit drug-like structures that comply with Lipinski's Rule of Five (e.g., favorable log P, hydrogen bond donors/acceptors, molecular weight), making them promising candidates for drug discovery (Chagas et al., 2018). These compounds also show potential as effective agents in molecular docking studies.

In addition to antioxidant and antimicrobial effects, mushrooms play a pivotal role in stimulating the immune system. Fig. 3 depicts the underlying mechanisms behind this stimulation. The β-glucans, one of the most researched compounds in mushrooms, have been shown to modulate immune responses by interacting with the innate immune system. These complex carbohydrates bind to receptors such as Dectin-1 and complement receptor 3 (CR3) on immune cells, triggering the activation of immune cells like macrophages and neutrophils (Ahmad et al., 2024). Once activated, these cells release cytokines, promoting a systemic immune response. β-glucans also enhance the activity of T-cells and natural killer (NK) cells, which are critical for identifying and eliminating infected or cancerous cells. G. lucidum, widely known for its immune-stimulating properties, has been shown to upregulate the production of interleukin-2 (IL-2), a cytokine that supports the growth and differentiation of T-cells (Bohn and BeMiller, 1995).

Fig. 3.

Mechanism of immunomodulation by mushroom-derived β-glucan and terpenoids: β-glucan activates Dectin-1 and CR3 receptors, while terpenoids interact with TLRs, triggering downstream signaling via NF-κB. This cascade enhances antibody production by B-cells, cytokine secretion by T-cells, reduces inflammation, inhibits tumor growth, facilitates opsonization and phagocytosis, and promotes wound healing. This figure is created by www.biorender.com.

Furthermore, mushrooms contain terpenoids, a class of compounds with potent immune-enhancing effects. Terpenoids have been shown to exert immunomodulatory effects by activating various immune receptors, such as Toll-like receptors (TLRs) (Fig. 3), which are involved in recognizing pathogens and initiating immune responses (Naini et al., 2024). The activation of these receptors results in the release of pro-inflammatory cytokines and chemokines, which not only support the immune response but also help in wound healing and tissue repair. For instance, the terpenoids found in mushrooms like G. lucidum and T. versicolor have demonstrated the ability to enhance phagocytosis, the process by which immune cells engulf and digest pathogens, thereby strengthening the body's ability to fight infections (Mustafin et al., 2022; Raza et al., 2024).

Recent studies have uncovered mechanisms through which mushrooms exert their therapeutic effects. Mushrooms like Hericium erinaceus (Lion’s Mane) contain erinacines and hericenones, which promote nerve growth factor (NGF) synthesis. NGF is crucial for the growth, maintenance, and survival of neurons, indicating the potential neuroprotective effects of this species in managing neurodegenerative diseases such as Alzheimer’s and Parkinson’s (Almjalawi et al., 2024). In addition, bioactive peptides found in mushrooms have demonstrated antihypertensive effects by inhibiting angiotensin-converting enzyme (ACE). This mechanism reduces blood pressure by preventing the conversion of angiotensin I to angiotensin II, a potent vasoconstrictor (Ichim et al., 2024) (Fig. 4A). These findings suggest mushrooms’ potential in managing cardiovascular health. Additionally, mushrooms like Agaricus blazei produces polysaccharide-protein complexes that enhance anti-tumor activity by inducing apoptosis in cancer cells (Fig. 4B). This occurs through the activation of caspase pathways and the inhibition of nuclear factor kappa B (NF-κB), which regulates cell survival and proliferation (Randeni and Xu, 2024; Ray et al., 2024). The health benefits of mushrooms are multifaceted, with antioxidant, antimicrobial, anti-tumor and immune-stimulating properties all linked to their bioactive compounds. By understanding the precise mechanisms behind these actions, researchers continue to explore how mushrooms can be used therapeutically to prevent and manage a wide range of diseases, from infections to cancer, and enhance overall health and well-being (Sharika et al., 2024).

Fig. 4.

Schematic representation of (A) the regulation of blood pressure through the renin-angiotensin system and (B) the apoptotic process mediated by mushroom-derived polysaccharides. (A) Angiotensinogen, produced by the liver, is converted to angiotensin I in the bloodstream. However, the angiotensin-converting enzyme (ACE) prevents the conversion of angiotensin I to angiotensin II. By reducing angiotensin II levels, activation of the AT1-R receptor on blood vessels is minimized, leading to vasodilation and helping to maintain normal blood pressure. (B) Polysaccharides inhibit NF-κB activation and its downstream signaling pathways, including the suppression of p50-p65 and E-selectin expression. Simultaneously, they promote the release of cytochrome c, which activates caspase-9 and caspase-3, leading to the apoptosis of cancerous cells and inhibition of tumor growth. This figure is created by www.biorender.com.

Antioxidant properties of mushrooms

The antioxidant properties of mushrooms are widely acknowledged and are attributed to their bioactive compounds. These compounds reduce oxidative stress and inflammation, providing protection against chronic diseases like cancer, cardiovascular disorders, and neurodegenerative conditions. Ergothioneine, a unique antioxidant found predominantly in mushrooms, protects mitochondria from oxidative damage and must be obtained through diet since it cannot be synthesized by plants or animals (Fu and Shen, 2022). β-Glucans serve a dual function by stimulating immune cell activity to enhance defense mechanisms while also reducing oxidative damage through their antioxidant properties (Kupradit et al., 2023; Trivedi and Upadhyay, 2024).

In poultry nutrition, mushrooms and their stems have demonstrated substantial benefits. Fig. 5 illustrates the antioxidant effects of supplementing mushrooms into poultry diets. A study on A. bisporus stem residue in laying hens revealed increased activities of antioxidant enzymes like superoxide dismutase (SOD) and glutathione peroxidase (GSH-Px) in serum and yolk, alongside reduced malondialdehyde (MDA) levels. This indicates enhanced antioxidant capacity and improved egg quality (Yang et al., 2021). Similarly, supplementing broiler diets with dried A. bisporus at 10 g/kg and 20 g/kg elevated liver and muscle tissue antioxidant defenses, increasing levels of GSH-Px, glutathione (GSH), glutathione reductase (GR), and glutathione S-transferase (GST) while lowering MDA levels, ultimately improving meat quality (Giannenas et al., 2010).

Fig. 5.

This schematic diagram illustrates the antioxidant effects of mushroom stem supplementation in poultry diets. The supplementation enhances the activity of antioxidant enzymes such as superoxide dismutase, glutathione peroxidase, glutathione reductase, and glutathione S-transferase, while reducing free radicals, oxidative stress, and malondialdehyde levels. This figure is created by www.biorender.com.

Mushrooms have also shown efficacy in poultry feed, enhancing liver and serum antioxidant markers like SOD and GSH-Px, while promoting sustainability through waste reduction (Chuang et al., 2021). Furthermore, incorporating A. blazei in broiler diets improved meat oxidative stability by boosting free radical scavenging activity, preserving tissue integrity during storage (Fanhani et al., 2016). Recent study has shown that mushroom supplementation reduces oxidative stress markers such as MDA while increasing protective enzymes like SOD and total antioxidant capacity (T-AOC) (Luo et al., 2024). Supplementation with F. velutipes significantly increased total antioxidant capacity while reducing oxidative damage in broilers (Mahfuz et al., 2019b). These benefits not only enhance the overall resilience of birds but also support improved growth performance and meat quality. This highlights mushrooms as sustainable and natural substitutes for synthetic antioxidants in poultry diets, addressing safety issues and regulatory challenges associated with synthetic additives.

The stems of P. ostreatus (Oyster) and L. edodes (Shiitake) are particularly rich in antioxidant compounds like phenolic acids (ferulic acid, caffeic acid) and ergothioneine. These components enhance immune function, reduce oxidative stress, and improve overall poultry health. Oyster mushroom stems have shown significant free radical neutralizing capacity in assays such as DPPH (2,2-diphenyl-1-picrylhydrazyl) and Ferric Reducing Antioxidant Power, confirming their strong antioxidant potential (Effiong et al., 2024). Similarly, shiitake stems, with their high ergothioneine and selenium content, protect cells from oxidative damage while supporting immune resilience (Xu et al., 2024). By integrating mushrooms and their stems into poultry feed, producers can enhance product quality, support animal health, and contribute to sustainable farming practices. Future research is expected to optimize mushroom-based formulations, further amplifying their antioxidant and immunomodulatory benefits in animal agriculture.

Antimicrobial properties of mushrooms

Mushrooms are increasingly recognized for their significant nutritional, medicinal, and antimicrobial properties, attributed to their diverse bioactive compounds. Blending mushroom stems into poultry feed offers multiple benefits, including reducing oxidative stress, enhancing gut health, and promoting overall bird productivity (Lee et al., 2014). Polysaccharides and fiber in mushroom stems act as prebiotics, supporting the growth of beneficial gut bacteria, improving nutrient absorption, and fostering better digestion (Andrade et al., 2024). These attributes make mushroom stems a valuable addition to sustainable poultry production.

Mushroom stems, such as those from L. edodes (shiitake) and P. ostreatus (oyster mushroom), have demonstrated notable antimicrobial properties. L. edodes contains lentinans, which are biologically active compounds known for their medicinal properties, including effectiveness against pathogens such as Candida albicans and Aspergillus niger (Atila, 2021). Similarly, P. ostreatus extracts exhibit antimicrobial activity against Escherichia coli and Staphylococcus aureus, attributed to various antimicrobial phenolic compounds (Yakobi et al., 2023; Nasir et al., 2024a). The antimicrobial effects of these compounds help control infections, improve poultry biosecurity, and reduce dependence on synthetic antibiotics.

The antimicrobial mechanisms of mushrooms are largely driven by compounds such as β-glucans, lectins, and phenolic substances. β-glucans disrupt bacterial cell walls and boost immune responses, providing a dual mechanism of protection (Cerletti et al., 2021). Lectins bind to carbohydrates on pathogen surfaces, preventing their attachment to host cells and inhibiting biofilm formation (Hassan et al., 2020a). Phenolic compounds neutralize free radicals and inhibit the growth of various pathogens, further enhancing the antimicrobial potential of mushrooms (Kosanić et al., 2012). Additionally, mushroom polysaccharides suppress harmful bacteria like Clostridium sp., E. coli, and Salmonella, while promoting beneficial bacteria such as Lactobacillus sp. and Bifidobacterium (Guo et al., 2004; Robinson et al., 2018).

Beyond their antimicrobial properties, mushrooms contribute to sustainable agricultural practices. They utilize agricultural byproducts efficiently, reducing waste and offering a natural alternative to synthetic growth promoters in poultry production. For example, supplementation with P. ostreatus has shown anticoccidial activity against Eimeria sp., reducing cecal coccidiosis and improving bird health and growth rate (Ademola et al., 2019; Nasir et al., 2023). Formulations such as 10 % medicinal mushrooms have further demonstrated their efficacy in significantly reducing pathogenic Clostridia in poultry, underscoring their role as natural growth promoters (Robinson et al., 2018).

Despite the established antimicrobial and nutritional benefits of mushrooms, further research is essential to isolate and characterize the specific bioactive compounds responsible for these effects. Understanding the mechanisms of action and exploring synergistic effects with existing antimicrobial agents could lead to innovative strategies for improving poultry health and productivity. The application of mushroom-derived antimicrobials holds great potential for advancing sustainable agriculture and addressing global challenges in animal health and food security.

Effects of mushrooms on immunity and blood biochemistry

Mushrooms, particularly their stems, have emerged as valuable additions to poultry diets due to their health enhancing, immune boosting, and metabolic regulatory properties. Often considered agricultural byproducts, mushroom stems are rich in bioactive compounds such as polysaccharides, β-glucans, antioxidants, phenolic compounds, vitamins, and essential nutrients, offering a range of benefits that support immune function, antioxidant defense, and overall metabolic health.

The immunomodulatory effects of mushroom stems are primarily attributed to polysaccharides, especially β-glucans, which activate key immune cells such as macrophages, T cells, and natural killer cells. This activation enhances the immune system's capacity to combat infections and maintain overall health. For instance, stems from L. edodes (shiitake mushrooms) are particularly rich in β-glucans and have demonstrated the ability to stimulate immune responses, improve disease resistance, and reduce the severity of infectious outbreaks (Chakraborty et al., 2023). Similarly, stems from P. ostreatus (oyster mushrooms) have been reported to positively influence humoral immunity in birds (Nasir et al., 2024b). Additionally, mushroom stems contain ergosterol, a precursor to vitamin D2, which is converted under UV exposure (Hussaana et al., 2024). Vitamin D is crucial for modulating immune responses, making mushrooms a valuable dietary component for addressing vitamin D deficiencies in poultry (Papoutsis et al., 2020).

Mushroom stems inclusion has also been shown to enhance humoral immunity by increasing serum immunoglobulin levels (IgG, IgA, IgM), which are vital for resisting infections. For example, supplementation with F. velutipes significantly elevated immunoglobulin levels in ISA Brown chickens compared to control and antibiotic-fed groups (Mahfuz et al., 2019b). Furthermore, certain mushrooms, such as Antrodia camphorata, have been shown to increase globulin levels, further reinforcing their immune-enhancing potential (Song et al., 2014). Elevated levels of cytokines such as IL-2, IL-4, IL-6, and TNF-α have been reported in broilers supplemented with mushrooms, indicating stronger immune activation and increased disease resistance (Mahfuz et al., 2019b; Guo et al., 2004).

Incorporating into poultry diets positively influences key blood biochemical parameters, reflecting improved overall health and metabolic efficiency. Enhanced protein metabolism has been observed, with significant increases in serum albumin levels, indicating better liver function and nutrient utilization (Mahfuz et al., 2019b). Improved lipid metabolism has also been noted, with reductions in total cholesterol, LDL cholesterol, serum triglycerides, and very low-density lipoprotein (VLDL) levels, alongside increases in HDL cholesterol. These improvements indicate mushrooms' potential to support cardiovascular health and contribute to leaner, healthier meat production (Shamsi et al., 2015; Sogunle et al., 2019; Toghyani et al., 2012). Mushroom stems also influence mineral metabolism, particularly calcium and phosphorus levels, which are crucial for bone health and eggshell quality. Supplementation with F. velutipes resulted in a modest increase in serum calcium and phosphorus levels, demonstrating its potential to enhance mineral absorption and utilization (Mahfuz et al., 2018a).

Mushroom stems, such as those from P. ostreatus and C. militaris, have demonstrated significant potential as sustainable feed additives. At a 1 % inclusion level, P. ostreatus residues enhanced immunity, and reduced oxidative stress in broilers (Hassan et al., 2020b). Similarly, C. militaris waste medium supplementation boosted immune response, and antioxidant capacity, showcasing the dual benefits of waste utilization and improved productivity (Hsieh et al., 2021). Medicinal mushrooms like G. lucidum (Reishi) and L. edodes (shiitake) are especially valued for their safe, natural alternatives to antibiotics, further supporting poultry health (Khan et al., 2019).

Moreover, these benefits align with consumer demands for natural, healthier, and more sustainable poultry production. Mushroom stems supplementation not only enhances flock health but also contributes to environmental sustainability by repurposing agricultural byproducts and reducing reliance on synthetic additives. The combined impact of improved immune responses, better metabolic health, and reduced oxidative stress provides a holistic approach to enhancing poultry productivity.

Mushroom stems supplementation offers a comprehensive and sustainable strategy to improve poultry health, resilience, and productivity. By boosting immune function, reducing oxidative damage, enhancing metabolic processes, and supporting sustainable farming practices, mushrooms address critical challenges in modern poultry production. Future research should focus on optimizing the inclusion levels, identifying synergistic effects with other feed components, and exploring long-term impacts on productivity and sustainability. As the poultry industry increasingly embraces natural and innovative solutions, mushroom stems stand out as a transformative tool for improving animal health, nutrition, and environmental sustainability.

Effects of mushrooms on growth performance

Mushroom stems can significantly enhance poultry growth performance due to their rich nutritional content. Stems from mushrooms like A. bisporus (Button) and P. ostreatus (Oyster) are high in protein, fiber, and minerals. When added to poultry diets, these stems provide a valuable protein source essential for muscle development and overall growth. Studies show that incorporating mushroom stems into feed improves weight gain and feed conversion ratios, as the high protein and fiber content supports better digestion and nutrient absorption (Chang and Miles, 2004; Nasir et al., 2024b). Efficient digestion leads to better growth performance, as poultry can maximize the benefits of their feed. Research suggests that poultry fed diets enriched with mushroom stems often experience faster growth and improved feed efficiency compared to standard diets (Lee et al., 2014). Moreover, the prebiotic properties of mushroom stems support a healthy gut microbiota. The fiber and polysaccharides in these stems nourish beneficial gut bacteria, fostering a balanced microbial community that enhances digestive health and immune function, further contributing to optimal growth and development in poultry (Wasser, 2014).

The inclusion of mushrooms in broiler diets has gained attention for its potential to improve growth performance, feed efficiency, and reduce the environmental impact of poultry production. A review of the literature provides valuable insights into how different mushroom species affect growth parameters like body weight, feed intake, and feed conversion ratio (FCR), as summarized in Table 3.

Table 3.

Effect of different mushroom species on growth performance.

Species	Bird Types	Duration	Levels	Effects	References
Pleurotus ostreatus	Ross 308	42 days	2 g/kg	The BW significantly decreased (P < 0.05) in the group supplemented with mushroom powder, accompanied by reductions in FI, weight gain, and final BW. Birds fed mushroom powder exhibited lower FI and weight gain (P < 0.05) compared to the control group. However, the FCR in the mushroom-fed birds was significantly better (P < 0.05) than that of the other groups.	Daneshmand et al., 2011
Agaricus bisporus	Ross 308	42 days	10g/kg	Mushroom supplementation had no significant effect on daily weight gain, FI, or FCR during any rearing phase of the chickens (P > 0.05).	Noruzi and Aziz-Aliabadi, 2024
Lentinus edodes	Arbor Acres	21 days	1, 2, 3, 5, and 10 g/kg	The daily BW gain was significantly (P < 0.05) greater in all treatment groups compared to the control group. Among the treatment groups, the 5 g/kg group exhibited a significantly (P < 0.05) higher BWG than the others.	Guo et al., 2004
Ganoderma sp.	Lorman Brown	140 days	0.5, 1, or 2 g/kg	No significant (P < 0.05) differences in FI were detected among the groups. However, the feed-to-gain ratio was significantly (P < 0.05) higher in the 1 g/kg and 2 g/kg groups compared to the other groups.	Ogbe et al., 2009
Agaricus bisporus	Ross 308	42 days	20 g/kg	The dietary inclusion of mushrooms resulted in the best FCR (P < 0.01), accompanied by a reduction in FI (P < 0.01) and BWG (P < 0.01) compared to the control group.	Altop et al., 2022
Agaricus bisporus	Ross 308	42 days	10 g/kg and 20 g/kg	There were no effects on FI, BW, or BWG among the groups up to 28 days of age. However, by 42 days, the 20 g/kg group demonstrated significantly (P ≤ 0.05) higher BW and BWG compared to the other groups. Additionally, at 42 days of age, feed efficiency values were significantly (P ≤ 0.05) higher in the 10 g/kg and 20 g/kg groups compared to the control group.	Giannenas et al., 2010
Pleurotus sapidus	Cobb 500	35 days	25 and 50 g/kg	The BWG, FI, and feed-to-gain ratio were not significantly (P < 0.05) different among the groups.	Schäfer et al., 2024
Pluerotes ostreatus	Ross 308	35 days	0.5, 1.0, 1.5 g/kg	The 1.5 g/kg supplementation led to a significant (P < 0.05) increase in body weight and growth rate, improved feed conversion efficiency, and reduced FI compared to the other groups.	Rebh et al., 2023
Flammulina velutipes	Arbor Acres	42 days	10 and 20 g/kg	The dietary inclusion of mushrooms had no significant (P > 0.05) effect on average daily FI, average daily BWG, or FCR during the starter (1–21 days), finisher (22–42 days), or overall experimental periods (1–42 days). Furthermore, there were no significant (P > 0.05) differences in initial or final body weight among the experimental groups.	Mahfuz et al., 2019b
Pleurotus ostreatus	Ross 308	42 days	10 and 20 g/kg	A significant (P > 0.05) increase in FI and BWG was observed with 10 g/kg supplementation, along with a lower FCR compared to the 20 g/kg group. However, no significant effect on BWG or FI was noted with 20 g/kg supplementation.	Hassan et al., 2020b
Pleurotus ostreatus	Ross 308	42 days	10 and 20 g/kg	The FI did not differ significantly (P > 0.05) among the groups. FCR showed a significant increase (P > 0.05) in the 10 g/kg and 20 g/kg groups. Although the 20 g/kg group exhibited higher BW, the difference was not statistically significant (P > 0.05) compared to the control group.	Toghyani et al., 2012

Abbreviations: FCR, Feed conversion ratio; FI, Feed intake; BW, Body weight; BWG, Body weight gain.

A study by Daneshmand et al. (2011) on P. ostreatus supplementation at 2 g/kg in Ross 308 broilers over 42 days found a decrease in feed intake, weight gain, and final body weight, suggesting that this dosage may not be ideal for optimal growth. Similarly, A. bisporus at 10 g/kg in Ross 308 broilers over the same period showed no significant effect on growth (Noruzi and Aziz-Aliabadi, 2024). However, a higher inclusion of A. bisporus (20 g/kg) improved FCR while decreasing feed intake and body weight gain (BWG), indicating that higher dosages may have different effects (Altop et al., 2022).

Conversely, supplementation with L. edodes and Tremella fuciformis (1-4 g/kg) in Arbor Acres broilers for 21 days resulted in increased body weight gain and improved FCR, suggesting a positive effect on growth (Guo et al., 2004). In a 140-day study on Ganoderma sp. in Lohmann Brown pullets, there were no differences in feed intake, but FCR and feed efficiency improved, indicating its potential as a growth promoter (Ogbe et al., 2009). The effects of P. ostreatus vary with dosage. Supplementing 10 g/kg in Ross 308 broilers increased feed intake and body weight gain with a lower FCR, while 20 g/kg had no significant effect (Hassan et al., 2020b). This suggests that lower dosages may be more beneficial for growth. Supplementing even lower levels (1-2 g/kg) decreased FCR and improved feed efficiency and weight gain (Toghyani et al., 2012). Supplementing P. ostreatus at 0.25, 0.5, and 0.75 g/kg in Ross 308 broilers over 35 days significantly increased body weight growth rate, reduced feed consumption, and enhanced feed conversion efficiency (Rebh et al., 2023). These findings highlight the importance of optimizing mushroom dosage for improved poultry performance.

Studies on other mushroom species, such as A. bisporus, show varying effects depending on dosage and duration. No significant impact on feed intake, body weight, or body weight gain (BWG) was observed up to day 28 in Ross 308 broilers, but by day 42, supplementation at 20 g/kg improved body weight and weight gain (Giannenas et al., 2010). Supplementing 1 g/kg inclusion of A. bisporus improved daily feed intake during the starter phase, with the lowest FCR recorded at 0.5 g/kg (Kavyani et al., 2014). Similarly, a study on Pleurotus sapidus in Cobb 500 broilers over 35 days found no significant effects on body weight gain, feed intake, or feed-to-gain ratio at 25 and 50 g/kg, suggesting this species may not influence growth performance at those dosages (Schäfer et al., 2024). Conversely, F. velutipes supplementation in Arbor Acres broilers at 10 and 20 g/kg over 42 days also showed no notable differences in average daily feed intake, BWG, or FCR (Mahfuz et al., 2019b).

These studies underscore the complex, dose-dependent effects of mushroom supplementation on growth performance. While some species and dosages do not significantly affect growth, others improve FCR, feed efficiency, and body weight gain, showing promise for mushrooms in poultry nutrition. Further research is needed to refine dosage levels and identify the conditions where mushroom supplementation maximizes growth performance. These findings highlight the importance of tailoring mushroom species and dosages to achieve optimal outcomes in poultry growth and feed efficiency.

Effects on carcass criteria and meat quality

Mushroom stems are gaining recognition for their potential to improve meat quality. Different mushroom species have been shown to positively influence key meat quality parameters, including pH, drip loss, cooking weight loss, color, and sensory attributes. Adding them in feed enhances the nutritional profile, supports better gut health, and improves nutrient absorption, ultimately promoting muscle development and increasing meat yield (NRC, 1994; Törős et al., 2024).

Mushroom stems contain bioactive chemicals that can enhance meat characteristics like flavor, tenderness, and color. This helps reducing oxidative stress during meat storage, leading to a longer shelf life and better sensory attributes (Ibrahim and Faujan, 2023). Using mushroom stems in poultry feed not only enhances nutrition and flavor but also supports environmental sustainability by repurposing agricultural by-products, lowering feed expenses, and fostering sustainable farming methods (Törős et al., 2024). This dual benefit of improving meat quality and reducing waste positions mushroom stems as a valuable addition to broiler production.

Table 4. illustrates the varying effects of different mushroom species on poultry meat quality. Mushrooms including F. velutipes and Agaricus brasiliensis showed minimal impact on carcass yield, pH, and color parameters, although some groups exhibited lower pH and cooking weight loss. P. ostreatus influenced cholesterol levels, increasing HDL and decreasing LDL, while A. bisporus affected meat color and pH values. Supplementation with medicinal mushrooms like H. erinaceus and G. lucidum improved meat preservation by reducing oxidative stress markers. However, A. bisporus also resulted in reduced sensory qualities such as flavor, juiciness, and overall acceptability, especially at higher doses. These studies demonstrate the diverse effects of mushrooms on poultry meat quality, affecting both nutritional content and sensory attributes

Table 4.

Effect of different mushroom species on meat quality.

Species	Birds	Duration	Levels	Effects	Reference
Flammulina velutipes	Arbor Acres	42 days	10 and 20 g/kg	There were no significant differences observed in the pH, drip loss, or color parameters [L* (luminosity), a*(red-green intensity), and b* (yellow-blue intensity)] of both the breast and thigh muscles among the experimental groups (P > 0.05).	Mahfuz et al., 2020
Agaricus brasiliensis	Ross 308	42 days	4, 8, 12, 16, and 20 g/kg	The treatments had no significant effects (P > 0. 05) on carcass yield, breast yield, or reduction in abdominal fat. However, all mushroom-supplemented groups exhibited significantly lower pH values compared to the control group. Additionally, the 16 g/kg and 20 g/kg groups demonstrated significantly lower cooking weight loss (P < 0.05) compared to other groups. No significant differences (P < 0.05) were observed in shear force values among the groups.	Guimarães et al., 2014
Pleurotus ostreatus	Arbor Acres	42 days	0.0075 g/L and 0.015g/L	The wings, back, thigh, and drumstick weights were numerically higher in the 0.015 g/L group compared to other groups. Total cholesterol was significantly higher (P < 0.05) in the 0.015 g/L group, while the control and 0.0075 g/L group showed statistically lower and comparable values. The HDL cholesterol concentration in the meat increased significantly (P < 0.05) in the 0.015 g/L group, whereas the LDL cholesterol content was significantly lower (P < 0.05) in the 0.015 g/L group compared to the control group.	Ekunseitan et al., 2017
Agaricus bisporus	Ross 308	42 days	20 g/kg stalk, 20 g/kg cap, and a mixture of 10 g/kg stalk + 10 g/kg cap	There were no significant differences in carcass characteristics among the treatment groups (P > 0.05). The L* (lightness) value was higher (P < 0.01) in the breast meat of the 20 g/kg stalk group but lower in the thigh meat of the 20 g/kg stalk and 20 g/kg cap groups compared to the control group. All dietary treatments significantly reduced (P < 0.01) the a* (redness) values in both breast and thigh meat. The 20 g/kg cap group increased (P < 0.05) the b* (yellowness) values in both breast and thigh meat compared to the control. Breast meat pH was elevated in the 10 g/kg stalk + 10 g/kg cap group and the 20 g/kg cap group, while thigh meat pH increased in the 10 g/kg stalk + 10 g/kg cap and 20 g/kg stalk groups.	Altop et al., 2022
Hericium erinaceus & Ganoderma lucidum	Ross 308	28 days	1, 1.5 and 2 g/kg	The inclusion of Hericium erinaceus at 2 g/kg exhibited significantly reducing free fatty acids (FFA) and oxidative stress markers compared to other groups. Moreover, this group showed a highly significant reduction (P > 0.01) in malondialdehyde (MDA) levels and peroxide value (PV) in frozen meat, highlighting its positive impact on preserving meat quality.	Al-Azzawi and Bandr, 2023
Pleurotus ostreatus	Ross 308	42 days	12.5, 25 and 50 g/kg	There were no significant differences (P > 0.05) observed among the treatment groups in terms of meat pH and temperature. Similarly, no treatment group exhibited significant effects (P > 0.05) on meat lightness, yellowness, or chroma. Additionally, there were no significant variations (P > 0.05) among groups in cooking loss, drip loss, water-holding capacity, or shear force values.	Mthana and Mthiyane, 2024.
Agaricus bisporus	Cobb 400	42 days	4, 8, and 12 g/kg	All treatment groups exhibited significantly lower (P > 0.05) scores for meat appearance and color compared to the control group. The 8 g/kg and 12 g/kg groups had significantly lower (P > 0.05) flavor and taste values than the 4 g/kg and control groups. Additionally, all mushroom-supplemented groups showed reduced odor compared to the control. Meat juiciness was also lower in the 8 g/kg and 12 g/kg groups compared to the control. Overall acceptability score was significantly reduced (P > 0.05) in the mushroom-fed groups.	Roy and Fahim, 2019.

Abbreviations: L*, lightness; a*, redness; b*, yellowness; FFA, Free fatty acid; MDA, Malondialdehyde; PV, Peroxide value; LDL, Low-density lipoprotein.

For example, F. velutipes supplementation had no significant effects on pH, drip loss, or color parameters in both breast and thigh muscles of Arbor Acres chickens after 42 days (Mahfuz et al., 2020). Similarly, P. ostreatus supplementation in Ross 308 broilers at varying levels (12.5, 25, and 50 g/kg) over a comparable period did not show significant effects on key meat quality attributes like pH, temperature, and color (Mthana and Mthiyane, 2024).

Conversely, other studies have demonstrated that specific mushroom species and supplementation levels can positively influence meat quality. In another study A. brasiliensis supplementation at 4 to 20 g/kg significantly affected cooking weight loss and pH of breast meat in Ross 308 broilers, though it did not alter shear force, suggesting improvements in moisture retention and meat texture (Guimarães et al., 2014). P. ostreatus has also been linked to increased meat yield. Supplementation at 7.5 ppm and 15 ppm in Arbor Acres broilers increased the percentage of leg and breast meat, although other cuts like wings, back, thigh, and drumstick remained unaffected (Ekunseitan et al., 2017). This indicates that P. ostreatus supplementation may be beneficial for optimizing yield in preferred meat cuts.

Meat color is a key aspect of quality and greatly influencing consumer perception and acceptance. A. bisporus supplementation in Ross 308 broilers, particularly at 20 g/kg of stalk, cap, or a mixture affected meat color (Altop et al., 2022). Thigh meat lightness showed decreased lightness, while breast meat lightness increased both redness decreased, and yellowness increased in both types of meat. These findings suggest that A. bisporus could enhance the visual appeal of poultry meat, potentially making it more attractive to consumers.

In terms of sensory attributes A. bisporus supplementation in Cobb 400 broilers at 4, 8, and 12 g/kg significantly improved characteristics such as color, juiciness, taste, odor, and overall acceptability (Roy and Fahim, 2019). This highlights mushrooms' potential to enhance the eating quality of broiler meat, making it more appealing to consumers. In another study, the effects of H. erinaceus and G. lucidum on meat quality in Ross 308 broilers were studied (Al-Azzawi and Bandr, 2023). While no significant differences were found in carcass weight, the study noted significant changes in dressing percentage, suggesting these mushrooms may influence meat processing characteristics. These studies indicate that mushroom supplementation can positively affect specific aspects of meat quality, such as color, texture, and sensory appeal, depending on the species and dosage used. As the poultry industry continues to explore natural ways to improve product quality, mushrooms offer a promising approach. Future research should focus on optimizing the use of mushrooms in broiler diets to maximize their benefits for meat quality.

Effects of mushrooms on egg production and quality

Adding mushroom stems into the diets of laying hens has been shown to positively impact both egg production and quality (Table 5). Studies suggest that supplementing diets with mushroom stems can boost egg production rates, as the improved nutrient intake supports better reproductive performance and overall well-being (Seid, 2023; Karageorgou et al., 2024). Additionally, the inclusion of mushroom stems can optimize feed efficiency, helping hens convert feed into eggs more effectively.

Table 5.

Effect of different mushroom varieties on egg production and quality.

Species	Birds	Levels	Duration	Effects	References
Lentinula edodes	Tetra Brown Layer	2.5 or 5 g/kg	8 weeks	Egg weight and egg mass did not differ significantly among the groups (P > 0.05). However, egg production was significantly higher (P > 0.05) in both mushroom-supplemented groups compared to the control group. The egg shape index and yolk color remained unaffected (P > 0.05) in all groups. Shell thickness decreased (P < 0.05) in the group receiving 2.5 g/kg of mushroom, while albumen height increased (P < 0.05) in the group supplemented with 5 g/kg of shiitake compared to the control. Additionally, the Haugh unit improved (P < 0.05) in both mushroom-fed groups, with the 5 g/kg supplementation yielding the highest value.	Hwang et al., 2012
Flammulina velutipes	Sonia sp. Hens	30 g/kg	64 weeks	Egg production was slightly higher in the mushroom-supplemented group compared to the control group. Moreover, the mushroom-supplemented group produced significantly fewer (P < 0.05) broken and soft-shelled eggs than the control group.	Yoshida et al., 2017
Cordyceps militaris	Hendrix Layer	5, 10 or 20 g/kg	12 weeks	No differences were observed among the groups for egg yolk weight, eggshell weight, shell thickness, or egg yolk color. However, the 20 g/kg group demonstrated a significant increase (P < 0.05) in egg white weight. All mushroom-supplemented groups exhibited significantly higher egg mass (P < 0.05) compared to the control group. Additionally, the 10 g/kg and 20 g/kg groups showed significantly lower (P < 0.05) egg cholesterol levels compared to the other groups.	Wang et al., 2015
Pleurotus eryngii	Hendrix Layer	5, 10 or 20 g/kg	8 weeks	The 10 g/kg and 20 g/kg groups exhibited a significantly higher Haugh unit (P < 0.05) compared to other groups. The cholesterol content (P < 0.05) in the 10 g/kg and 20 g/kg groups was significantly reduced.	Lee et al., 2015
Flammulina velutipes	Hy-Line Brown	10, 20, 30, 40 or 50 g/kg	5 weeks	There were no significant differences in egg production across any of the treatment groups. However, egg weight was significantly higher in the 10 g/kg and 30 g/kg groups compared to the control group (P < 0.05). Interestingly, the 40 g/kg group demonstrated improvements in egg quality parameters such as albumen height, Haugh unit, eggshell weight, and shell thickness, although yolk color remained unaffected (P < 0.05).	Lee et al., 2014
Flammulina velutipes	ISA Brown	20, 40 or 60 g/kg	9 weeks	The egg yolk antioxidant levels, including T-AOC and T-SOD, were significantly elevated (P < 0.05) in the 40 g/kg group. Conversely, MDA levels were significantly reduced (P < 0.05) in egg across all groups fed with mushroom stems.	Chen et al., 2019
Pleurotus ostreatus	Hy-Line Brown	3, 6, 9 and 12 g/kg	6 weeks	Mushroom supplemented all groups significantly influenced (P < 0.05) egg weight. However, it did not show any notable effects on hen-day production (HDP) or egg mass.	Natsir and Wicaksono, 2020
Agaricus bisporus	Nongda III layer	20, 40, 60, 80, and 100 g/kg	8 weeks	There were no significant changes in hen day egg production or average egg weight (P > 0.05). Similarly, no notable differences were observed in egg storage characteristics between the treatment and control groups (P > 0.05). However, yolk antioxidant activity showed marked improvements with mushroom treatments; SOD activity was significantly higher (P < 0.05), and MDA content was significantly lower (P < 0.05). Additionally, GSH-Px activity in the yolk was significantly elevated across all mushroom-treated groups.	Yang et al., 2021
Flammulina velutipes	ISA Brown	20, 40 or 60 g/kg	7 weeks	No significant differences observed in daily egg production, egg weight, egg mass, or the number of unmarketable eggs among the treatment groups. Additionally, parameters such as shape index, shell weight, shell thickness, yolk weight, yolk index, yolk weight, and albumen weight remained unaffected (P > 0.05) across all treatment levels. However, the Haugh unit was significantly higher (P < 0.05) in all mushroom-treated groups. Shell color showed a significant increase (P < 0.05) in the 40 g/kg and 60 g/kg groups, while yolk color improved significantly (P < 0.05) at all levels of mushroom supplementation. Furthermore, overall acceptability was significantly higher (P < 0.05) in the 40 g/kg and 60 g/kg groups.	Mahfuz et al., 2018b
Hypsizygus marmoreus	Hy-Line Brown	50, 100 or 150 g/kg	12 weeks	There were no significant differences (P > 0.05) in egg production, egg weight, or egg mass among the groups. Similarly, eggshell breaking strength, thickness, and Haugh unit showed no notable differences (P > 0.05). However, yolk color was significantly enhanced (P < 0.05) in all mushroom-fed groups.	Kim et al., 2014

Abbreviations: SOD, Superoxide dismutase; MDA, Malonaldehyde; GSH-Px, Glutathione peroxidase; T-AOC, Total antioxidant capacity; T-SOD, Total superoxide dismutase.

Bioactive compounds in mushroom stems enhance egg quality by improving shell strength, nutritional value, and reducing fracture rates (Mukhtar, 2023). As consumer demand for environmentally friendly and nutritionally enhanced egg products grows, incorporating mushroom stems can offer a practical solution for improving both the quantity and quality of eggs (Mahfuz et al., 2018b). This innovative approach benefits producers and consumers alike, showcasing how sustainable feed sourcing can enhance poultry production.

The effects of various mushroom species on egg production and quality in poultry are summarized in Table 5. Mushroom supplementation generally improved egg quality, including enhancements in egg mass, yolk color, and antioxidant activity. Mushroom species such as L. edodes and F. velutipes increased egg production and improved certain quality parameters like albumen height and shell quality, though effects on production and weight varied. Other species, like Pleurotus eryngii and A. bisporus, also positively impacted egg characteristics such as cholesterol levels and antioxidant activity, while effects on egg production were inconsistent across studies. L. edodes (shiitake) supplementation in Tetra Brown layers increased egg production and improved egg quality, measured by a higher Haugh unit, though it also resulted in thinner eggshells (Hwang et al., 2012). In another study F. velutipes supplementation in Sonia hens led to higher egg production and fewer broken or soft-shelled eggs, improving overall production efficiency (Yoshida et al., 2017). On the other hand, Cordyceps militaris supplementation increased egg white weight and egg mass in Hendrix Layer hens, enhancing specific aspects of egg production (Wang et al., 2015). However, some studies, on A. bisporus (Yang et al., 2021) and Hypsizygus marmoreus (Kim et al., 2014) reported no significant effects on egg production or egg quality, suggesting that the impact of mushroom supplementation may depend on species, dosage, and duration.

While mushroom supplementation has shown varied effects on egg production, its impact on egg quality has generally been more positive, though results still depend on the mushroom species and dosage used. For instance, F. velutipes has demonstrated notable benefits for egg quality. Supplementation with F. velutipes at 10-50 g/kg in Hy-Line Brown hens over 5 weeks increased egg weight and improved egg quality parameters such as albumen height, eggshell weight, Haugh unit, and shell thickness (Lee et al., 2014). This suggests F. velutipes can enhance various aspects of egg quality, improving the marketability of eggs.

Similarly, P. eryngii supplementation has also been linked to improved egg quality. Supplementation at 5–20 g/kg in Hendrix Layer hens over 8 weeks raised the Haugh unit and lowered egg cholesterol and triglyceride content, suggesting it not only enhances structural quality but also improves the eggs' nutritional profile (Lee et al., 2015). Additionally, P. ostreatus supplementation in Hy-Line Brown hens at 3–12 g/kg over 6 weeks increased egg weight, though there was no significant effect on hen-day production (HDP) or egg mass (Natsir and Wicaksono, 2020).

Moreover, F. velutipes stem base supplementation increased yolk antioxidants, such as total antioxidant capacity (T-AOC) and total superoxide dismutase (T-SOD), in ISA Brown hens, adding nutritional value for health-conscious consumers (Chen et al., 2019). However, not all mushroom supplements significantly affect egg quality. No significant differences in egg production or most quality parameters with F. velutipes supplementation, except for a higher Haugh unit (Mahfuz et al., 2018a). Similarly, there are no significant differences in eggshell strength or Haugh unit with H. marmoreus supplementation in Hy-Line Brown hens, indicating that not all species impact egg quality at the tested levels (Kim et al., 2014).

While certain mushroom species like F. velutipes and P. eryngii have demonstrated the potential to enhance egg quality by improving structural integrity and nutritional content, other species have shown minimal or no effect. These findings underscore the importance of selecting the right mushroom species and dosage levels to achieve the desired improvements in egg production and quality. Further research is necessary to optimize these variables and fully harness the benefits of mushroom supplementation in poultry diets.

Effects of mushrooms on gut microbiota

Mushroom supplementation has emerged as a promising dietary strategy for modulating the gut microbiome in poultry, promoting gut health, and improving overall performance. Fig. 6 illustrates the mechanism by which mushroom supplementation enhances gut health. Various studies have shown that mushroom species wastes can positively influence gut microbial populations by reducing harmful bacteria and promoting the growth of beneficial microbes, such as Lactobacillus and Bifidobacterium, essential for digestion and nutrient absorption (Törős et al., 2023; Seid, 2023). This shift in microbial diversity contributes to better intestinal health and enhanced immune function. Mushroom stems, rich in dietary fiber and prebiotics, foster beneficial gut bacteria and improve microbiome diversity (Yu et al., 2023). The modulation of the gut microbiota also strengthens the intestinal barrier and boosts the production of short-chain fatty acids (SCFAs), which support immune function and gut health. This immune support is especially beneficial for young animals, reducing disease susceptibility and improving growth performance. Fiber from mushroom contributes to better feed efficiency and inhibit the growth of harmful microorganisms like Salmonella and E. coli, promoting a healthier gut environment (Törős et al., 2023; Yu et al., 2023). This not only enhances animal health but also supports safer food production. In summary, integrating mushroom stems into animal diets offers a comprehensive approach to improving gut microbiota, leading to enhanced health, productivity, and food safety.

Fig. 6.

The schematic representation illustrates the role of mushroom stem supplementation in improving gut health and immune functionality in poultry. Mushroom stem supplementation in feed enhances the population of beneficial microbes (Lactobacillus, Bifidobacterium), boosts short-chain fatty acid (SCFA) production, and promotes nutrient absorption. Simultaneously, it reduces the prevalence of harmful microbes such as Salmonella and E. coli by improving intestinal barrier integrity through increased mucin secretion by goblet cells and strengthened tight junctions (TJs). These improvements lead to enhanced immune responses mediated by plasma cells, IgA, B cells, and macrophages, ultimately suppressing gut inflammation. This figure is created by www.biorender.com.

One study found that feeding P. sapidus mycelium to broilers had minimal impact on gut microbiota, with no significant effects on the microbial community (Schäfer et al., 2024). However, mushroom byproducts in chicken diets had more pronounced effects, reducing the Firmicutes/Bacteroidetes ratio and increasing Lactobacillaceae populations, which are associated with anti-inflammatory and antioxidant benefits that enhance overall health (Chuang et al., 2021). Similarly, P. ostreatus positively influenced gut microbiota, promoting intestinal health and well-being, positioning it as a promising supplement for poultry farming (Törős et al., 2024). Mushroom polysaccharides, particularly those from Pleurotus sp., have been shown to promote the growth of beneficial bacteria like Lactobacillus and Bifidobacterium, which are essential for gut health in poultry (Barcan et al., 2023). This aligns with findings that medicinal mushroom supplements reduced harmful bacteria like Mollicutes and increased beneficial bacteria, contributing to overall health improvements in chickens (Robinson et al., 2018).

Polysaccharides and terpenoids in mushrooms enhance gut microbial balance by enhancing immunity and overall poultry health (Gungor et al., 2017). Mushroom supplementation, at 30 g/kg, reduced harmful bacteria such as E. coli and maintained intestinal morphology, supporting gut microbiota balance (Kavyani et al., 2014). A. bisporus has shown prebiotic effects by increasing beneficial bacteria, including Lactobacilli and Bifidobacteria, without significantly altering intestinal morphology (Giannenas et al., 2010). Similarly, mushroom polysaccharides have been found to alter the cecal microbial ecosystem by increasing beneficial bacteria and reducing harmful ones like Bacteroides spp. (Guo et al., 2004).

While few studies show that mushroom extracts can increase beneficial bacteria like bifidobacteria without significantly reducing pathogen load, such as Salmonella (Willis et al., 2009), other findings, like A. bisporus stalk meal supplementation, increased mesophilic aerobic bacteria in the cecum, suggesting a positive impact on gut microbial populations (Altop et al., 2022). Recent studies have explored various dietary supplements and functional foods to modulate the gut microbiota and enhance immune responses (Abdel-Wareth and Lohakare, 2021, 2023). These approaches, including the use of feed additives and sustainable protein sources, represent cutting-edge methods to promote gut microbiota balance and improve animal health in modern livestock management (Abdel-Wareth et al., 2018, 2022, 2023, 2024; Amer et al., 2021, 2022; Almeldin et al., 2024; Paneru et al., 2024; Abdel-Wareth, 2016). Furthermore, medicinal mushrooms have demonstrated the ability to modulate the gut microbiome, offering a sustainable alternative to antibiotics for promoting gut health and immune responses in poultry (Mahfuz et al., 2020). These findings emphasize the potential of mushrooms not only to improve gut health but also to serve as sustainable feed additives, providing an alternative to conventional methods in poultry production.

Conclusions

Mushroom stems and related by-products offer a promising, sustainable alternative to conventional feed additives in poultry diets. Rich in functional compounds like β-glucans, phenolics, and polysaccharides, they enhance immune response, support metabolism, and improve antioxidant status. Research indicates their positive effects on growth performance, feed efficiency, meat quality, and egg production, positioning them as effective substitutes for antibiotic growth promoters. Their use also contributes to sustainable agriculture by valorizing agricultural waste. To maximize these benefits, future research should focus on optimizing inclusion levels, processing methods, and species selection. Long-term studies are needed to evaluate their impact on health, productivity, nutrient utilization, and gut microbiota. Exploring interactions with other feed ingredients and assessing economic feasibility will be essential for broader commercial adoption. Overall, mushroom-based feed additives present a multifunctional solution for enhancing poultry performance while advancing environmentally responsible production practices.

Credit authorship contribution statement

Md Salahuddin: Writing- original draft, Methodology, Formal analysis, Conceptualization, Kayla Stamps: Writing- original draft, Formal analysis, Conceptualization. Ahmed Abdel-Wareth: Writing–review & editing, Supervision, Conceptualization. Jayant Lohakare: Writing–review & editing, Supervision, Resources, Conceptualization. Venkatesh Balan: Writing – review & editing, Resources, Maedeh Mohammadi: Writing – review & editing, Weihang Zhu: Writing – review & editing, Woo Kyun Kim: Writing – review & editing.

Declaration of generative AI and AI-assisted technologies in the writing process

During the preparation of this work, the author(s) used Grammarly and Quillbot to check grammar and paraphrase the sentences where needed. After using this tool/service, the author(s) reviewed and edited the content as required and take(s) full responsibility for the content of the publication.

Declaration of competing interest

The authors affirm that they have no recognized financial or personal conflicts of interest that could have influenced the outcomes or interpretations presented in this study.