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types of inedible medicinal wood- decaying mushrooms, and its amount also increases
signicantly in old fruiting bodies, as well as in the mycelium of edible mushrooms. It
may be different in different parts of the mushroom fruiting bodies. For example, the
stem of L. edodes have a higher ber content (45%) than the caps (36.6%), for other
species studied, the difference was not so great. The chitin and the matrix components
(α- and ß- D- glucans and mannans) are non- starch polysaccharides whose molecules can
be classied as total dietary ber fraction (Cheung, 2013). Dietary ber in mushrooms
primarily consists of water- soluble (less than 10% DW) and the insoluble fractions, with
chitin and β- glucans being the most prevalent. These molecules are practically resistant
to digestion and absorption in the human small intestine and have benecial effects such
as attenuation of glucose and cholesterol in the blood.
There are many diverse substances in mushrooms that determine specic tastes
and aromas that stimulate appetite and make mushrooms exceptionally attractive as
a delicious product. For different mushroom species, the specic taste and smell are
determined by the combination of many volatile components: alcohols, aldehydes, etc.,
as well as a combination of non- volatile substances such as soluble sugars, organic
acids, free amino acids, and 5′- nucleotides were determined (Hu et al., 2020; Kalač,
2013). Hu et al. (2020) detected in dried S. rugosoannulata fruiting bodies ve 5′nucleotides (5′- AMP, 5′- GMP, 5′- UMP 5′- CMP, and 5′- IMP). The total content of 5′-
nucleotides in pileus (18.33– 22.76 mg/ g) was higher than that of stipe (8.17– 11.91 mg/
g). Similar results were also observed by Wu et al. (2015) for H. marmoreus and Li et al.
(2015) for P. eryngii.
Summing up all of the above, mushrooms are a nutritious food and an important
source of bioactive constituents. It is extremely important to look at the selected promising, widely cultivated mushroom species.
13.4 BIOACTIVE COMPOUNDS OF EDIBLE
ANDMEDICINAL MUSHROOMS
13.4.1 Cyclocybe aegerita
Cyclocybe aegerita (V. Brig.) Vizzini (family Tubariaceae); common name: pioppino
mushroom or velvet pioppini, poplar mushroom, chestnut mushroom, black poplar,
swordbelt agrocybe, yanagi- matsutake (Figure 13.3A). C. aegerita is a popular edible
mushroom widely distributed in Southern Europe, the Southeastern United States, and
similar Asian climate zones that prefer warm to temperate climates. The black poplar
mushroom is known for its delicious taste and has been cultivated since ancient Rome.
When cooked, pioppino has a nutty avor and crunchy texture. C. aegerita is rich in
proteins, carbohydrates, and polyunsaturated fatty acids and has a low- fat content. In
ad dition, pioppino contains several biologically active substances such as polysaccharides,
phenols, indole derivatives, and tocopherols, unique ceramides with proven antitumor
potential. Pioppino could be a source of enzymes (peroxidases, laccases, proteases,
Edible and Medicinal Mushrooms as an Eco-Friendly Source of Food 367
FIGURE 13.3 Fruiting bodies: A Cyclocybe aegerita; B Hericium erinaceus; C Hypsizygus
marmoreus; D Lentinula edodes; E Pleurotus eryngii; F Stropharia rugosoannulata. Photos
of the authors.
and lipoxygenases), which are promising biocatalysts in biotechnological applications
(Cateni et al., 2022; Landi et al., 2017b, 2022).
13.4.1.1 Components of Cyclocybe aegerita with proven
pharmacological activities
Polysaccharides are the most abundant biologically active substances in the fruiting
bodies and mycelium of C. aegerita and have become an important component in the
research and development of natural medicines and health products (Jing et al., 2018).
Mushroom polysaccharides are considered effective antioxidants with a complex structure. Natural polysaccharides are conjugated with other bioactive components such
as proteins, phenolic compounds, amino acids, and lipids. Several recent studies have
found that polysaccharides from C. aegerita have great potential for antioxidant and
antiaging functions, as well as antitumor, antiangiogenic, and thrombosis treatment
properties (Jing et al., 2018; Liu et al., 2020a, 2020b; Peng et al., 2022; Wu et al., 2022).
The polysaccharide from C. aegirita showed potent antiaging activity, manifested
by increasing cell viability and β- Gal viability, preventing cell cycle arrest in the G1
phase and reducing mitochondrial membrane potential (Liu et al., 2020a, 2020b). Wu
et al. (2022) reported that C. aegerita polysaccharide in combination with Lactobacillus
rhamnosus GG acted as a prebiotic and helped mitigate the damage caused by aging
by regulating oxidative stress and gut microbiota. Also, Liu et al. (2022) studied the
impact of combining C. aegerita polysaccharides with Bidobacterium lactis Bb- 12
on antioxidant activity, antiaging properties, and modulation of gut microbiota. The
results demonstrated that the polysaccharides and Bidobacterium lactis Bb- 12 complex signicantly increased the average lifespan of male and female mice under natural
aging. Additionally, the complex enhanced their climbing ability and increased antioxidant enzyme activity, protecting them from oxidative damage induced by H2O2. In
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D- galactose- induced aging mice, the addition of complex resulted in a signicant
increase in antioxidant enzyme activity, regulation of aging- related biomarker levels,
changed gut microbiota diversity, restoration of microbial structure, and increased abundance of benecial bacteria, particularly Lactobacilli, in the intestines.
Additionally, Ji et al. (2013) investigated the antitumor mechanism of polysaccharide from C. aegerita by evaluating the expression of tumor necrosis factor- α (TNF-
α) and interferon- γ (IFN- γ) in rat esophageal carcinoma. The rats were given anticancer
treatment daily for 4 weeks. It was found that C. aegerita polysaccharide in combination with chemotherapy can regulate immune function in rat esophageal carcinoma,
potentially by modulating cytokine activity, in particular, reducing TNF- α levels and
increasing IFN- γ levels (Ji et al., 2013).
Ferments. Pecyna et al. (2009) presented genetic information about this new group of
mushroom peroxidases known as aromatic peroxygenases (APOs). The C. aegerita
peroxygenase gene (Apo 1) has been identied at the messenger RNA and genomic
DNA level. The gene sequence was conrmed using peptide sequences. Aromatic
peroxygenases are multifunctional peroxide- consuming enzymes involved in the
biodesulfurization, dehalogenation, denitrication, and hydroxylation of various aromatic compounds. The most studied biocatalyst of this type is C. aegerita. Aromatic
peroxygenases from C. aegerita catalyze reactions such as the peroxidation of naphthalene, toluene, dibenzothiophene, or pyridine, which effectively act on cytochrome P450
monooxygenases. APO is an attractive biocatalyst capable of carrying out a wide range
of complex reactions and may be promising for use in organic synthesis. According to
the literature data, C. aegerita APO is capable of sulfoxidation and hydroxylation of
dibenzothiophene, hydroxylation of the ring and side chain of toluene, N- oxidation of
pyridine, as well as selective epoxidation of naphthalene. APO enzymes are secreted
heme- thiolate proteins (Pecyna et al., 2009). Also, Kluge et al. (2009) reported the
mechanism of peroxygenase- catalyzed oxygen transfer by the example of naphthalene that is regioselectively hydroxylated into 1- naphthol. Due to their extracellular
nature, the high degree of glycosylation (20– 40%), and hence high stability, APArelated enzymes could become new model biocatalysts for oxygen transfer reactions
(Kluge et al., 2009).
Li et al. (2021) isolated a novel brinolytic enzyme (ACase) from the fruiting
bodies of C. aegerita. It has been established that the ACase enzyme can destroy all
three chains (α, β, and γ) of brinogen. Moreover, it acted not only as a plasmin- like
brinolytic enzyme but also as a plasminogen activator. The authors reported that the
enzyme could slightly hydrolyze human thrombin, inhibit thrombin activity, and act as
an anticoagulant to prevent thrombosis. Based on their results, the authors suggested
that ACase may act as a therapeutic agent for the treatment of thrombosis or as a functional food product, but further research is needed (Li et al., 2021).
Phenolics compounds are considered important metabolites with clinically useful
activities. The antioxidant activity and free radical scavenging ability of C. aegerita
correlated with total phenolic content (Lin et al., 2017; Lo & Cheung, 2005). The
fruiting bodies of C. aegerita contained protocatechuic acid, chlorogenic acid,
ferulic acid, and sinapic acid (Lin et al., 2017). The aqueous extract of C. aegerita
fruiting bodies had a total phenolic content (TPC) of 13.67±0.21 M gallic acid
Edible and Medicinal Mushrooms as an Eco-Friendly Source of Food 369
equivalent (GAE)/ mg extract. In addition, the effects of phenolic- rich aqueous extract
on antiogenesis- related expression of vascular endothelial growth factor (VEGF) secretion and angiogenesis- related signaling in tumor cells, were investigated. The model
chosen for the experiment was the use of Caco- 2 cells (colon epithelial tumor cells)
and HUVEC (endothelial cells). The authors report that a decrease in intracellular
reactive oxygen species (ROS) levels and secretion of VEGF expression was observed
when treated with C. aegerita extract, and in Caco- 2 cells (colon epithelial tumor cells)
treated with C. aegerita. The aqueous extract inhibited the VEGF induced proliferation
in human umbilical vein endothelial cells (HUVECs). Down- regulation of intracellular ROS level and VEGF secretion were also observed in Caco- 2 cells treated with
aqueous extract of C. aegerita. Inhibition of other angiogenic cascades in HUVECs
included a decrease in the migration of endothelial cells (ECs) by 46.14±8.53%.
Authors reported that the presence of antiangiogenic properties of extracts C. aegerita
in vitro and phenolics might contribute to the effect.
Thus, it has been experimentally proven that C. aegerita contains many nutrients and
bioactive components including polysaccharides, which are the main active compounds
having great potential for antioxidant and antiaging functions. When combined with
Lactobacillus rhamnosus GG and Bidobacterium lactis Bb- 12, they may act as a prebiotic and help mitigate the damage caused by aging by regulating oxidative stress and
gut microbiota. In addition, polysaccharides have multiple physiological functions,
such as antioxidant, antitumor, antiangiogenic, and thrombosis treatment properties.
13.4.2 Hericium erinaceus
Hericium erinaceus (Bull.) Pers. (family Hericiaceae), common name Lion’s mane,
pom pom, bearded tooth, known also as bear’s head, hog’s head fungus, old man’s beard,
white beard, monkey’s mushroom, monkey’s head or “Houtou” in Chinese, and mountain monk mushroom or “Yamabushitake” in Japanese (Figure 13.3B). H. erinaceus
is a popular culinary and medicinal mushroom cultivated in many European countries, North America, and Asia. In natural conditions, it grows in South East Asia
(China and Japan), Europe, and North America, and is also found in Ukraine. In folk
medicine, it is used for immune stimulation and antiseptic for treating chronic gastritis, cancer of the oesophagus and stomach, leukemia, treating the nervous system,
and rst of all, neurodegenerative disorders. H. erinaceus is a good source of proteins
and carbohydrates, low in fat (Table 13.1). Also, the mushroom composition includes
dietary bers, essential amino acids, minerals, vitamins, and ergosterol (provitamin D2)
making it a high- quality, low- calorie food (Cohen et al., 2014; Das et al., 2021; Mau
et al., 2001; Rodrigues et al., 2015; Thongbai et al., 2015).
Current information on the medicinal properties of H. erinaceus indicates a wide
biological spectrum of its action. The H. erinaceus composition includes various
metabolites such as β- glucans polysaccharides and secondary metabolites such as
terpenoids (hericenones, erinacine), sterols, and isoindolinones. Immunomodulatory,
anticancer, anti- inammatory, gastroprotective, cytoprotective, antioxidant, antibacterial, hypoglycemic, and hepatoprotective effects of H. erinaceum extracts are reported,
as well as its neuroprotective, neurotrophic, and neuro- regenerative properties.
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13.4.2.1 Components of Hericium erinaceus with proven
pharmacological activities
The bioactive metabolites found in H. erinaceus can be classied into two groups: high
molecular weight compounds, including polysaccharides, and low molecular weight
compounds, such as terpenoids and polyketides.
Polysaccharides. The H. erinaceus fruiting bodies and mycelial mass contain glucans
and glucan- protein complexes. More than 35 polysaccharides with multiple activities
have been extracted from H. erinaceus mycelia (Spelman et al., 2017). Anticancer and
immunostimulating effects of polysaccharides from H. erinaceus are primarily associated
with the activation of many immune cells (T- cells, macrophages, and cytokines), as
well as increasing the ability of the immune system cells to nd and destroy migrating
cancer cells in the human body (Jiang et al., 2014; Zhang et al., 2022). Unlike muchexisting chemotherapy of anticancer drugs, polysaccharides isolated from H. erinaceus
do not have a toxic effect. Such polysaccharides can activate c- Jun N- terminal kinases
(JNK), which participate in apoptosis, and enhance the intracellular apoptotic signal
level (Badalyan & Rapior, 2020). Numerous tests were conducted in vitro and in vivo
showed signicant therapeutic potential of metabolites of the H. erinaceus mushroom
with promising results against a variety of cancers, including those in the breast, blood,
colon, liver, lungs, sarcoma, and stomach (Kim et al., 2014; Ma et al., 2012; Thongbai
et al., 2015). A study of the immunomodulatory potential indicated that the polysaccharide fraction of ethanol extracts of H. erinaceus and its derivatives can promote
the maturation of dendritic cells, their production of cytokines, and the proliferation of
T- cells, as well as activation of macrophages and increase production of tumor necrosis
factor (TNFα) (Ren et al., 2017).
Terpenoids. Various diterpenes and sesterpenes are present in H. erinaceus. The most
interesting are two classes of terpenoid compounds: hericinones and erinacines obtained
from fruiting bodies and cultivated mycelium, respectively. Hericinones, a class of aromatic compounds, were isolated from the H. erinaceus fruiting body by Kawagishi et al.
(1990). They pointed out the perspective of using hericinones as natural remedies against
Alzheimer’s disease, assuming the possibility of their passing through the blood- brain
barrier and inducing the production of neurotrophic growth factor (NGF) in the brain.
Hericenones A and B showed signicant cytotoxicity against HeLa cells (Kawagishi
et al., 1990). In addition, hericenon B effectively inhibits the aggregation of platelets,
which prevents thrombosis, heart attacks, and strokes. However, hericenones C, D, E,
and H showed a stimulating effect activity in the synthesis of NGF in vitro. On the other
hand, hericenones F and G did not stimulate NGF synthesis under the same conditions,
but 3- hydroxyhericenone F protected against neuronal death (Kawagishi et al., 1991).
Hericenones I, J, and L showed cytotoxic activity against EC109 tumor cells (Ma et al.,
2012). It is still controversial whether all hericenones are active components in stimu-
lating NGF biosynthesis. Some researchers have shown that hericinones C, D, and E
did not increase NGF biosynthesis in the 1321N1 cell line (Mori et al., 2008). Instead,
Phan et al. (2018) investigated the stimulation of NGF synthesis by hericenon E cells
Edible and Medicinal Mushrooms as an Eco-Friendly Source of Food 371
in rat pheochromocytoma (PC12) using several pharmacological inhibitors. In addition
to the above- mentioned, hericenones isolated from fruiting bodies, another class of terpenoid compounds, a group of cyathane- type diterpenoids, named erinacines (A– I),
was obtained from cultured mycelium (Jing et al., 2022; Roda et al., 2021; Spelman
et al., 2017). Erinacines are low- molecular substances that stimulate synthesis of NGF
through interaction with mitogen- activated protein kinase of neurons and can overcome
hematoencephalic barrier. Erinacine derivatives are potential drugs for degenerative
disorders of neurons and peripheral regeneration nerves. Thus, the cyanine- diterpenoid
erinacine A, obtained from the H. erinaceus mycelium, demonstrated inhibitory
activity against a wide range of cancer cells associated with the gastrointestinal tract
and has been proven to have an effective protective effect against Parkinson’s disease
(Venturello et al., 2021). Ratto et al. (2019) reported the positive effect of an H. erinaceus
supplement (He1) with a known amount of Erinacine A, Hericenone C, Hericenone
D, and L- ergothioneine, on locomotor frailty and cerebellum of aged mice by investigating specic protein markers representative of cell proliferation activity and newborn neurons occurrence. Effect of erinacine A- enriched H. erinaceus supplementation
(HE) on cognitive function and serum levels of the brain- derived neurotrophic factor
(BDNF), neuropeptide Y (NPY), fecal levels of chitinase, and gut microbiota composition has been investigated by Bizjak et al. (2024) in clinical trials. They observed a
signicant improvement in cognitive ability and an increase in gut microbiota diversity,
which was positively correlated to NPY levels. Supplementing the diet with HE was
acknowledged as a safe and well- tolerated intervention with a neurocognitive benet
(Bizjak et al., 2024).
Some studies were conducted with H. erinaceus extracts regarding the mechanisms
involved in the process of brain neuroprotection (Kushairi et al., 2019; Mori et al.,
2008, 2009; Phan et al., 2018). Effects of erinacine A, which can prevent ischemic
injury neurons and act as an anti- inammatory agent, have been reported (Thongbai
et al., 2015).
Erinacerins. Wang et al. (2015) isolated ten new isoindoline- 1- ones, named erinacerins
C−L from a solid H. erinaceus culture. This is another group of secondary metabolites
produced by H. erinaceus. Some of the compounds inhibited the growth of cancer cells.
Sterols. In the fruiting body of H. erinaceus, ten erinarols A– J, ve ergostane- type
sterol fatty acid esters, and ten ergostane- type sterols have been identied. Sterols,
such as ergosterol, confer antioxidative properties. H. erinaceus has been identied
as the most potent inhibitor of both low- density lipoprotein (LDL) oxidation and
HMG Co- A reductase activity in vitro, indicating potential therapeutic applications
for preventing oxidative stress- related vascular diseases (Spelman et al., 2017). Dai
et al. (2015) evaluated the biomass and ergosterol content of H. erinaceus mycelium using plant- derived compounds supplementation into liquid culture and got the
highest ergosterol content of 2.33 mg/ g after 6 days of cultivation with 100 mol/ L
salicylic acid (Dai et al., 2015; Friedman, 2015). Some of these compounds exhibit
anti- inammatory and antiproliferative properties in cell assays (Cui et al., 2014;
Friedman, 2015).
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Glycoprotein and glycolipids. Interestingly, H. erinaceus is not just a source of low
molecular weight biologically active compounds, but also some proteins that have
a considerable tumor suppressive potential. Thus, the HEP3 protein isolated from
H. erinaceus exhibited immunomodulatory activity in lipopolysaccharide- activated
macrophages by decreasing the overproduction of tumor necrosis factor- α, interleukin
(IL)- 1β, and IL- 6, and downregulating the expression of inducible nitric oxide synthase
and nuclear factor- κB p65. The further immunomodulatory effect was caused by the
stimulation of intestinal microbiota with protein, which included activation of proliferation and differentiation of T- cells and stimulation of antigen- presenting cells of the
intestine (Diling et al., 2017). Another example of a biologically active protein isolated
from H. erinaceus is a glycoprotein HEG- 5, which was able to induce apoptosis in the
gastric cancer cell line SGC- 7901, stimulating expression of some proapoptotic factors
(Zan et al., 2015). Cui et al. (2014) reported on the isolation of a novel glycoprotein
(HEG- 5) from cultures of H. erinaceus mycelia. The compound has a molecular weight
of 14.4 kDa and was shown to have hemagglutinating activity. Analysis using FT- IR
and NMR showed that HEG- 5 contains the protein and carbohydrate parts with (1→4)-
linked β- galactose and β- glucose residues, and circular dichroism showed that the HEG5 is a predominantly β- sheet glycoprotein. The glycoprotein inhibited the growth of
human gastric carcinoma cells (Cui et al., 2014; Friedman, 2015).
Cerebrosides are natural organic compounds from the group glycolipids, another
class of compounds found in mushrooms. It is shown that cerebroside E, isolated from
H. erinaceus fruiting bodies, exhibited a signicant inhibitory effect on angiogenesis
in HUVECs. This compound has also been shown to have antitoxic properties, and
attenuated cisplatin- induced nephrotoxicity in LLC- PK1 cells, which is sufcient to
propose its use in a cancer chemotherapy protocol (Lee et al., 2015).
Neuroprotective and neurotrophic effects. It is reported above that the fruiting bodies
and mycelia of H. erinaceus produce several classes of biologically active molecules,
including polysaccharides, proteins, lectins, phenols, and terpenoids. Erinacine A has
been shown to cause potent restore nerves, strengthens their properties and effectively
inhibits death of neuronal cells. It can enhance the synthesis of NGF by increasing the
secretion of norepinephrine and catecholamines. Cyatane- xyloside of erinacin P and
its biochemical transformations in erinacine A as well as erinacine B induced the NGF
synthesis (Thongbai et al., 2015). It has been found that erinacine A was able to prevent
ischemic injury to neurons and act as an anti- inammatory agent. The use of hericenone
C and erinacine A recommended for improving memory functions and improvement
teaching. These compounds can signicantly affect synthesis of NGF in vitro and/ or in
vivo (Jiang et al., 2014; Khan et al., 2013; Soares et al., 2013; Thongbai et al., 2015).
Amycenone (amyloban), which is an activator of brain function and can exert a
protective effect on brain cells in culture protecting them from damage by amyloid
peptides, was isolated from H. erinaceus extracts (Inanaga, 2012). As shown in further
studies, hericenones and amyloban, which are effective against damage to the human
brain in Alzheimer’s and Parkinson’s diseases. Organic extracts from fruiting bodies
of H. erinaceus possess a neurotropic effect that enhances the myelination process in
mature myelin bers (Moldavan et al., 2007). Bioactive compounds extracted from
the mycelia and fruiting bodies of H. erinaceus may be used as a potential alternative
Edible and Medicinal Mushrooms as an Eco-Friendly Source of Food 373
medicine for the treatment of depression (Chong et al., 2020). Laboratory studies
proved the absence of any negative consequences of using H. erinaceus. The clinical
studies with amiloban indicate its promising results against dementia and Alzheimer’s
disease, primarily due to the absence of any side effects (Thongbai et al., 2015). In addi tion, amycenone reduces excess body weight and attenuates hyperlipidaemia in mice
by inhibiting lipogenesis and promoting lipolysis through lipid metabolism pathway
stimulation and fatty acid β- oxidation acceleration (Kudo et al., 2022).
Thus, H. erinaceus has numerous biological activities. Hericenones and
erinacines stimulate NGF, which suggests that H. erinaceus has a certain inuence on the functioning of the brain and the autonomic nervous system, and taking
H. erinaceus gives the ability to reduce depression and anxiety (Limanaqi et al., 2020).
Polysaccharides from H. erinaceus with different molecular weights not only play a
role in the immunomodulation of dendritic cells but also demonstrate neuroprotective
effects for neurons.
Antimicrobial effect and antioxidant activity. Antimicrobial effect of Hericium species
due to ability to produce antibacterial metabolites with a wide and narrow spectrum
of action was shown in numerous studies (Khan et al., 2013; Lomberg et al., 2023;
Narmuratova et al., 2022; Song et al., 2020; Thongbai et al., 2015). This can have poten tial health benets and could be recommended for further analysis, isolation and identication of antibacterial compounds potentially promising in pharmacology.
Ryu et al. (2009) showed the presence of ergothioneine and polyphenols in biomass of H. erinaceus and H. coralloides. Meanwhile, ergothioneine, which humans
are unable to synthesize, a unique sulphur- containing amino acid, is an antioxidant,
cytoprotective, and anti- inammatory element with therapeutic potential approved by
World Food Agencies (Bell et al., 2022; Roda et al., 2022).
Thus, H. erinaceus is among the highly praised edible mushrooms, as producers
of neuroprotective biomolecules. Its traditional use for chronic ailments, coupled with
the ndings of current studies, indicates that H. erinaceus is safe and holds signicant potential as a neuroprotective and neurotrophic therapeutic agent for neurological
conditions. Its rich micronutrient composition suggests that utilizing the entire fungus
could be the most benecial approach clinically. Thus, preparations from H. erinaceus
are already available today to prevent and improve conditions with Alzheimer’s disease,
Parkinson’s disease, and dementia, as well as in sports medicine. Despite all the va riety
of neurotrophic effects of macromycetes on the human body, functional changes in
the nervous system, which are the basis of the described effects, have been studied not
enough, the vast majority of the results described in the review, carried out on cell lines
and experimental animals (in vitro and in vivo), and require further observations.
13.4.3 Hypsizygus marmoreus
Hypsizygus marmoreus (Peck) H.E. Bigelow (family Lyophyllaceae), common name
Beech mushroom or buna- shimeji (bunashimeji), brown beech or brown clamshell
mushroom, mountain echo mushroom or yamabiko hon- shimeji (Figure 13.3C).
H. marmoreus is native to forests of Asia, North America, and Europe, and is now
374 Wild Edible Plants
widely cultivated. The main producers of its fruiting bodies are the countries of East
Asia, mainly Japan. H. marmoreus is a highly nutritious edible mushroom, valuable
for its delightful taste and signicant medicinal properties. Brown beech mushrooms
and their white variety are popular dietary mushrooms in East Asian regions, including
China, Japan, and Taiwan. Both fruiting bodies and mycelia are good sources of
polysaccharides, protein, essential amino acids, soluble sugar, lectins, vitamins,
enzymes, and avor nucleotides (Lee et al., 2009; Liu et al., 2016; Liu et al., 2018; Wu
et al., 2014). Because of the presence of these compounds, this species shows a variety
of properties, including anticancer, antioxidant, anti- inammatory, antihypertensive,
and antiallergic effects (Liu et al., 2018; Xu et al., 2020).
13.4.3.1 Components of Hypsizygus marmoreus with
proven pharmacological activities
Polysaccharides extracted from mushroom mycelia have been shown to possess a wide
range of benecial biological properties. Published data demonstrated that polysaccharides
play crucial roles in preventing and treating oxidative damage, suggesting their potential development as antioxidants (Badalyan et al., 2019; Liu et al., 2018). Many experi mental works on H. marmoreus polysaccharides demonstrated their anticancer and
antioxidant properties (Chang, 2004; Motoi, 2003; Ye et al., 2023). On a number of
test systems, extracts from the fruiting bodies and mycelium of the beech mushroom
along with inhibiting the development of carcinoma metastases, also showed signicant
antioxidant activity. Based on these studies, extracts were made from H. marmoreus
and F. velutipes, and a preparation called “EEM” (extracts of edible mushrooms) was
supplied. Clinical studies of EEM were performed and the effects of EEM alone and
in combination therapy of EEM and cancer chemotherapy agents for advanced cancer
patients were studied and a positive effect of EEM was found (Ikekawa, 2005). Liu et al.
(2018) reported that H. marmoreus mycelia polysaccharides (MPS) have shown antiammation effects against lung damage induced by lipopolysaccharides in mice.
In recent years, increasing attention has been paid to selenium- enriched
polysaccharides showing higher antioxidant effects than regular polysaccharides, which
allows garnering them as a safe and effective selenium resource. Thus, H. marmoreus
cultivated in a liquid medium with Na2SeO3 accumulated in mycelia polysaccharides
containing 70.15 g/ g selenium (MSPS) (Liu et al., 2016). The antioxidant activities
in vitro demonstrated that MSPS had potential effects on scavenging reactive oxygen
species and enhancing the reducing power. The treatment of MSPS in vivo demonstrated
that the MSPS could reduce the levels of malondialdehyde, lipid peroxide, glutamic
oxaloacetic transaminase, and glutamic- pyruvic transaminase activities and improve the
levels of glutathione peroxidase, superoxide dismutase, total cholesterol, and triglyceride in serum/ liver homogenate against CCl4- induced injures (Liu et al., 2016).
Indole compounds presented in H. marmoreus are an important group of compounds
with procognitive, antidepressant, antioxidant, and neuroprotective properties (Kała
et al., 2022). Six indole compounds such as L- tryptophan, 5- hydroxy- L- tryptophan,
serotonin, tryptamine, 5- methyltryptamine, and melatonin were identied. An essential amino acid L- tryptophan, which is a precursor of neurotransmitters in the body,
was found in all samples, with the highest content observed in brown H. marmoreus
Edible and Medicinal Mushrooms as an Eco-Friendly Source of Food 375
(8.97– 72.1 mg/ 100 g DW). In mycelial cultures the L- tryptophan content ranged from
17.3 to 25.6 mg/ 100 g DW. The higher antiradical properties of mushrooms could be
related to the content of indole compounds (Angelini et al., 2023). Authors suggested
that the enrichment of the growth medium with grape pomace could implement the
medicinal properties of H. marmoreus, whose content in terpenoids, carbapenems, and
indoles was signicantly increased compared with the mushrooms cultivated in the
standard medium, as shown by ABTS assay, could be related to the presence of indole
compounds. Indeed, the extract with the highest content of indole compounds had the
highest antioxidant effect.
Sterols, especially ergosterol, are converted into vitamin D2 in the presence of ultra-
violet rays, which plays an important role in the prevention of cancer and regulates
other immunomodulatory functions. Ergosterol is also responsible for the absorption of
Ca and P, which contribute to the proper functioning of the skeletal system. Ergosterol
content in white and brown H. marmoreus fruiting bodies consisted of 74.8 and 116
mg/ 100 g DW, respectively (Kała et al., 2022). Among the analyzed H. marmoreus
samples, the highest ergosterol contents were observed in the brown and white strains
mycelium grown on the medium enriched with adding Zn and Mg (166 and 142 mg/
100 g DW, respectively). So, the addition of magnesium and zinc salts to the culture
medium increased both the ergosterol and ergosterol peroxide contents in the mycelium
compared with the non- enriched cultures.
Phenolic compounds. Xu et al. (2020) compared phenolics, antioxidant activities, and
antiproliferative properties in brown and white H. marmoreus fruiting bodies. The results
showed that the contents of catechin, gallic acid, and protocatechuic acid of brown
fruiting bodies were higher than those of white. Moreover, brown had greater cellular
antioxidant activity, peroxyl radical scavenging capacity and oxygen radical absorbance capacity values than the white one, which demonstrated that brown H. marmoreus
presented a stronger antioxidant capacity. Both brown and white H. marmoreus showed
remarkable antiproliferative activities against HepG2 cells and brown H. marmoreus
was proven to be the more effective.
Ukaegbu et al. (2020) reported that tested aqueous and methanol extracts of
H. marmoreus (brown and white var.) and F. velutipes caps against two breast cancer
cell lines showed possess higher antioxidant activities against DPPH: aqueous caps
extracts of F. velutipes and white H. marmoreus (IC50= 0.202 and 0.573 mg/ mL, respectively), and H2O2 (IC50= 0.622 and 0.745 mg/ mL, respectively) compared to the methanol
extracts. Aqueous extracts of the mushrooms also showed better ferric reducing antioxidant power values against ferric ions compared to the methanol extracts. Finally, the
mushroom extracts showed good antiproliferative activities against human breast cancer
cell lines. The highest phenylalanine and cinnamic acid contents were determined in
the brown fruiting bodies of commercial origin, 422 and 8.88 mg/ 100 g DW, respectively (Kała et al., 2022). In addition to their well- known antioxidant activity, phenolic
compounds found in mushrooms can exhibit signicant antimicrobial effects against
a wide range of organisms (Angelini et al., 2023). The antimicrobial activities of
extracts from H. marmoreus against Gram- positive and Gram- negative bacteria, yeasts,
dermatophytes, as well as phytopathogenic fungi, have been reported (Angelini et al.,
2023).
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