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376 Wild Edible Plants
Other bioactive compounds. The medicinal properties of H. marmoreus are attributed
to several bioactive molecules, including polyterpenoid compounds like hypsiziprenolA9, which have been shown to exert cytotoxic effects in human liver cancer HepG2
cells (Chang et al., 2004). Recent experimental data have demonstrated that bioactive
compounds such as lovastatin derived from H. marmoreus can regulate the levels of
lipoproteins, total cholesterol and prevent oxidative stress (Badalyan et al., 2019).
Kała et al. (2022) reported that among the analyzed H. marmoreus extracts, the highest
lovastatin content was observed in the white fruiting bodies (74.5 mg/ 100 g DW) and
less in brown one (66.7 mg/ 100 g DW). In both the white and the brown varieties, a
higher lovastatin content was observed in the fruiting bodies than in the mycelium,
regardless of the type of substrate on which they were grown. The addition of zinc and
magnesium salts to the culture medium results in a reduction in the lovastatin content
compared with the non- enriched culture, which is particularly noticeable in the case
of the white variety (a reduction of nearly 50% in the lovastatin content). Among the
H. marmoreus extracts tested, the highest ergothioneine content was determined in the
mycelium of the white variety grown on the enriched medium (80.4 mg/ 100 g DW), less
content showed brown strain (73.0 mg/ 100 g DW).
Glucans. The highest glucan contents observed in the white and brown fruiting bodies
were 61.4 and 58.0 g/ 100 g DW, respectively (Kała et al., 2022). In mycelial cultures,
the addition of Zn and Mg salts resulted in an increase in the content of both total
glucans and β- glucans compared with the mycelia from in vitro cultures grown on the
control medium.
Thus, H. marmoreus is a good dietary source of K, Fe, Zn, Mg, sterols, lovastatin,
and glucans with antioxidant potential and rich in umami- tasting compounds such as
guanylic acid, glutamic acid, and aspartic acid (Angelini et al., 2023; Hu et al., 2020;
Kała et al., 2022; Lomberg et al., 2003) that make this mushroom a promising nutritious
food and an important source of bioactive constituents.
13.4.4 Lentinula edodes
Lentinula edodes (Berk.) Pegler (family Omphalotaceae), common name shiitake, sawtooth oak mushroom, black forest mushroom, black mushroom, golden oak mushroom,
or oakwood mushroom (Figure 13.3D). L. edodes is an edible medicinal mushroom
cultivated nowadays, in Europe, Asia, Australia, and North America countries. Recently,
L. edodes has gained popularity for both its nutritional prole and bioactive compounds.
L. edodes accounts for 17% of the global edible fungi supply and is now the second
most popular edible mushroom in the global market due to its good taste, nutritional
benets, and therapeutic value (Sheng et al., 2021). Shiitake mushrooms sport a rich,
woodsy avor that intensies when cooked, lending a savory umami avor to every dish
in which they star. They have a dense, meaty texture and stems that range from tender
to brous. L. edodes is the rst medicinal macrofungus to enter the realm of modern
biotechnology (Bisen et al., 2010).
L. edodes is rich in protein, carbohydrates, unsaturated fatty acids, vitamins, and
other nutrients and is low in fat, making it a high- quality, low- calorie food. In addition,

Edible and Medicinal Mushrooms as an Eco-Friendly Source of Food 377
shiitake is considered the active producer of different primary and secondary bioactive
metabolites, which are responsible for their pharmacological abilities. In the fruiting
bodies and mycelial biomass of L. edodes bioactive components have been identied, such as polysaccharides, steroids, phenols, nucleotides derivatives, cyclic- sulfurcontaining compounds, and glycoprotein derivatives (Bisen et al., 2010; Łysakowska
et al., 2023; Reis et al., 2012; Roszczyk et al., 2022).
13.4.4.1 Components of Lentinula edodes with proven
pharmacological activities
Polysaccharides of L. edodes are widely studied as the most valuable compounds with
proven anticancer, antioxidant, antiaging, antimicrobial, and immunomodulatory properties. Immunomodulatory action of fungal polysaccharides is a complex interaction
between immunological, metabolic, and epigenetic changes (Roszczyk et al., 2022).
β- glucans are the main polysaccharides found in the fungal cell wall and are responsible
for most of the biological effects. Glucan molecules are considered to be pathogenassociated molecular patterns and various pattern- recognition receptors can recognize
them (Łysakowska et al., 2023). The L. edodes mushroom is a rich source of β- glucans.
The content of β- glucans in shiitake varies between 20.0– 25.3 (g/ 100 g DW). L. edodes
is a source of two well- studied and widely approved polysaccharide medicines: LEM
(an acronym for L. edodes mycelia), a protein- bound polysaccharide derived only from
the mycelium, and lentinan – a cell- wall branched – β- D- glucan extracted from both the
fruiting body and mycelium. Both compounds are immune system enhancers that demonstrate anticancer activity (Bisen et al., 2010). It takes its specic name from β- glucan
lentinan, which stimulates immune cells to attack cancer cells. Lentinan enhances
the production of T lymphocytes and can potentiate the effect of AZT (3′- Azido- 3′-
deoxythymidine) in the antiviral treatment of AIDS (Łysakowska et al., 2023). Its positive effects have been proved in the treatment of, e.g., glioma (human astrocytoma U251
cells), breast cancer, and liver cancer (Ataollahi & Larypoor, 2022; Wang et al., 2021;
Yang et al., 2013).
Lentinan is mainly composed of β- glucan with therapeutic properties such as
antitumor, anti- inammatory, and antidiabetes properties. Lentinan was approved as an
adjuvant for stomach cancer therapy in Japan in 1985. It is approved for treating multiple types of cancer as well as for hepatitis and other diseases. Lentinan is available as
capsules, tablets, and injections. Clinical data show that lentinan is a biological response
modier and an immunostimulant with proven efcacy in treating hepatitis, HIV, malignant pleural effusion and cancers (Zhang et al., 2019, 2022).
Lentinan products have been approved for marketing as prescription drugs, including
tablets, capsules, oral agents, and injections, for the treatment of various diseases such
as chronic viral hepatitis from China’s National Medical Products Administration (Zhou
et al., 2024).
In addition, polysaccharides from L. edodes have a positive impact on the intestinal
microbiota by increasing the number of Bidobacteriaceae species and Lactobacillaceae
species and changing the number of Firmicutes species, Enterobacteriaceae species,
and Lachnospiraceae species (Xue et al., 2020). Fungal polysaccharides can be used as
prebiotics, regulating the quantity and diversity of the intestinal mycobiota (Wang et al.,

378 Wild Edible Plants
2018). Also, lentinan has recently been used in a variety of studies, ranging from food
and medical applications to a novel biomaterial. Lentinan as a biocompatible multifunctional polysaccharide has been used as a pharmaceutical additive in the development of
customized drugs or gene carriers with an improved safety prole (Kumar et al., 2023).
Phenolic compounds such as phenolic acid (p- hydroxybenzoic acid, vanillic acid,
syringic acid, p- coumaric acid, benzoic or cinnamic acid derivatives) and avonoids
were identied in the fruiting body of L. edodes (Uffelman et al., 2023). In addition, phe nolic and volatile substances (15 alcohols, 13 aldehydes, 9 alkanes, 5 sulfur- containing
compounds) were isolated and identied from the fruiting bodies of L. edodes. Yao
et al. (2023) analyzed the total phenolic (TPC), and total avonoid content of raw
and processed shiitake mushrooms under different cooking conditions. Compared to
the raw sample (127.08 mg GEE/ g DW), shiitake mushrooms baked in the oven for
5 minutes showed the highest TPC (222.13 mg/ g DW) with an increase of 75%. For
cooked shiitake mushrooms, the greatest loss of TPC was observed after 15 minutes,
with a reduction rate of 57%. Moreover, steaming for 5– 20 min and frying also showed
a decrease in TPC, and the maximum degree of loss was 56% and 27%, respectively.
On the other hand, the increase in TPP during oven baking may be due to the destruction of cell membranes and walls, which leads to an increased release of phenolics (Yao
et al., 2023).
The chemical prole of shiitake fruiting bodies and mycelium varies depending
on the strain, growing conditions, degree of maturity, and the proportion of individual
anatomical parts in the total mass of the mushroom (Łysakowska et al., 2023). The
chemical composition and functional properties of shiitake grown on log bed and sawdust media differed from each other. Log- grown fruiting bodies had a higher content of
terpenoids and phenolic components and exhibit higher antioxidant and hypoglycemic
potential compared to sawdust media (Song et al., 2020).
The fruiting bodies of L. edodes cultivated on logs had higher concentrations of secondary metabolites as phenol components with strong antioxidant capacity compared
to L. edodes cultivated on sawdust. Meanwhile, L. edodes grown on sawdust, had high
concentration of primary metabolites, which ensured a high growth rate. This information is useful for determining optimal growing conditions for L. edodes mushrooms for
food and medicinal purposes (Nam et al., 2021).
Kała et al. (2021) showed that a complete medium improves the properties of
synthesized fungal biomass. Mycelial biomass of L. edodes was obtained in media
enriched with zinc, selenium, l- phenylalanine, alone and as a mixture. Mycelium of
L. edodes had the best health- promoting properties growing in medium with mixture and characterized by increased signicantly content of p- hydroxybenzoic and
protocatechuic acids (Kała et al., 2021).
Lee et al. (2021) determined the effect of ethanol extract fruiting body of L. edodes
on ultraviolet (UV) A and UVB- induced changes in matrix metalloproteinase (MMP)
and type I procollagen expression using human immortalized HaCaT keratinocytes.
It was shown that extract containing polyphenols and β- glucan from shiitake
inhibits the expression of MMP- 1 and MMP- 9 and increases the expression of type
I procollagen in HaCaT keratinocytes irradiated with UVA and UVB. Results indicate that L. edodes can be developed as a cosmetic material to suppress UV- mediated
skin aging.

Edible and Medicinal Mushrooms as an Eco-Friendly Source of Food 379
Thus, L. edodes is among the most valuable medicinal mushrooms. Shiitake
is an edible cultivated mushroom used as “functional food” and has a long history in oriental folklore for the treatment of tumors, u, heart diseases, high blood
pressure, obesity, problems related to sexual dysfunction, and aging, diabetes, liver
ailments, respiratory diseases, exhaustion, and weakness. According to literature
data, L. edodes have great potential for the production of useful bioactive metabolites
and they serve as a prolic resource for drugs. The identied bioactive compounds
in L. edodes of known molecular structures account for a wide range of benecial
biomedical effects, most notably in the prevention of diverse physiological disorders
and diseases.
The abundance of polysaccharides, essential amino- and fatty acids, vitamins in
the fruiting bodies, and mycelial mass of shiitake is a valuable source of biologically
active compounds that can help maintain the health and well- being of the body. At the
same time, further scientic research is needed to conrm these benets and to develop
optimal methods for adding L. edodes to food, taking into account technological, sensory, and food safety aspects.
13.4.5 Pleurotus eryngii
Pleurotus eryngii (DC.) Quél. (family Pleurotaceae), common name King trumpet
mushroom or king oyster mushroom, cardoncello, French horn mushroom, king brown
mushroom, boletus of the steppes, trumpet royale (Figure 13.3E). P. eryngii is an edible
delicacy mushroom cultivating in Europe, the Middle East, and North America as well
as in many parts of Asia. The fruiting bodies of P. eryngii are easy to produce with
high yield and the products have a large market due to their good taste and ability to be
cooked directly. When cooked, they have a avor and texture similar to scallops. Their
avor is mild, but it intensies as they cook.
P. eryngii is rich in protein, carbohydrates, unsaturated fatty acids, vitamins, and
other nutrients and is low in fat, making it a high- quality, low- calorie food. In addition,
the King oyster is considered the active producer of different primary and secondary
bioactive metabolites, which are responsible for their pharmacological properties
(Badalyan et al., 2019). In the fruiting bodies and mycelial biomass of P. eryngii bio-
active components have been identied, high molecular weight primary metabolites
such as polysaccharides (α- glucans, and β- glucans), proteins (eryngin), glycoproteins,
lectins and also low molecular weight secondary metabolites (fatty acids and its esters),
avonoids (as chrysin, myricetin, naringenin, quercetin), polyphenols, and triglycerides
(Calabretti et al., 2021; Sharma et al., 2021; Torres- Martínez et al., 2022).
13.4.5.1 Components of Pleurotus eryngii with proven
pharmacological activities
Polysaccharides of P. eryngii include chitin, galactans, α- and β- glucans, hemicellulose,
mannans, and xylans. The fruiting bodies and mycelial biomass of P. eryngii have been
found to contain high levels of α- glucan, β- glucan and total glucan. The high molecular

380 Wild Edible Plants
weight β- glucans (β- (1,3)→(1,6)- glucans), which interact with the immune system
to increase/ decrease specic aspects of host response, are more important (Calabretti
et al., 2021). The signicance of the immunopharmacological activities associated with
β- glucans varies depending on their source and structure, including molecular weight,
degree of branching, and conformation (Ferreira et al., 2015). Experimental studies
have shown that β- (1,3)- glucans exhibit humoral and cellular immunity, and thereby
protect against tumor development and pathogen infection (Vetvicka, 2019). The quality
of β- glucans as polysaccharide immunomodulators is generally associated with very
low toxicity (Ma et al., 2020).
Mitsou et al. (2020) studied the effect of β- glucans from the fruiting body P. eryngii
on the gut microbiota of volunteers over 65 years of age. It was found that β- glucans
from P. eryngii caused a strong lactogenic effect. In addition, polysaccharides from
P. eryngii and the prebiotic inulin had comparable prebiotic indices. The authors noted
that the mushroom P. eryngii, rich in β- glucans, may have benecial effects in vitro on
the intestinal microbiota and/ or the production of short- chain fatty acids in older people
(Mitsou et al., 2020).
Chou et al. (2013) determined the prebiotic activity at relatively low
concentrations (0.1% to 0.5%) of polysaccharide fractions isolated from P. eryngii
wastes. They have been shown to enhance the survival rate of Lactobacillus acidoph-
ilus, L. casei, and Bidobacterium longum subsp. longum during cold storage. These
polysaccharides had synergistic effects with the amino acids and peptides from
a yogurt culture to maintain probiotics, and therefore the polysaccharide showed
signicant protective effects on these bacteria in simulated gastric and bile juice
conditions (Chou et al., 2013).
The antioxidant potential of intracellular and extracellular polysaccharides from
fruiting bodies, mycelial mass, and culture broth of P. eryngii has been reported by
various researchers. Jing et al. (2013) isolated and characterized two extracellular
polysaccharides from the submerged culture broth of P. eryngii. The small molecular
weight polysaccharides express strong antioxidant capacities. The DPPH, hydroxyl,
and superoxide anion radical scavenging activities of the extracellular polysaccharide
produced during the submerged culture of P. eryngii were 66.36%, 59.63%, and 38.69%,
respectively, which were higher than positive control butylated hydroxytoluene (Sun
et al., 2013). Moreover, the antioxidant properties of the polysaccharides from P. eryngii
has been improved by the sulphonation process (Li & Shah, 2014).
P. eryngii is capable of producing during submerged cultivation extracellular
polysaccharides, which can be used as antioxidants that enhance adaptive immune
responses (Sun et al., 2013). For large- scale production of extracellular polysaccharide
of P. eryngii, it is possible to use agricultural waste, such as vinasse, wastewater from
alcohol distilleries (Ivanova et al., 2023).
Zhang et al. (2020) summarized that P. eryngii polysaccharide can indirectly
express antitumor activity through the body’s own immune system, and it has little or
no data about its cytotoxicity, which makes it a potential new type of anticancer drug.
However, most of the studies have reported the in vitro and in vivo biological activity
of P. eryngii polysaccharides and reports about the development of their polysaccharide
products are rare (Ren et al., 2016; Zhang et al., 2020).

Edible and Medicinal Mushrooms as an Eco-Friendly Source of Food 381
Terpene and sterol derivatives’ composition. The rst member of C20 diterpenoids with
the skeleton deriving from a cyclododecane core fused with two γ- lactone units has
been isolated from the solid culture of P. eryngii was eryngiolide A (Wang et al., 2012).
Besides, Xue et al. (2015) reported that three triterpenoids (2,3,6,23- tetrahydroxy- urs12- en- 28 oic acid, 2,3,23- trihydroxyurs- 12- en- 28 oic acid, and lupeol) isolated from the
fruiting bodies of P. eryngii have been showing signicant inhibitory activity against
MCF- 7 breast cancer cell lines in vitro conditions.
Angiogenesis has been considered an important factor in impacting tumor growth
and metastasis. Fu et al. (2016) reported that isolated ubiquinone- 9 from the chloroform extract of P. eryngii exerted a signicant inhibition in neovascularization at a con-
centration of above 500 g. Besides, ubiquinone- 9, a chloroform extract of P. eryngii,
demonstrated the property of inducing apoptotic cell death by inhibiting the activity of
mammalian DNA topoisomerase I. Topoisomerase I was considered an attractive target
for antitumor agents.
Kikuchi et al. (2018) reported the isolation of ergostane- type steroids, including
eringiacetals A and B and pleurocins A and B, from fruiting bodies. Also, their inhibitory effects on nitric oxide (NO) production and human recombinant aromatase from
fruiting body P. eryngii was evaluated. In addition, the authors describe the isolation
and structural elucidation of three new bisabolane- type sesquiterpenes (triterpenes
which are based on the cyclopentane perhydrophenantrene ring system) as ergosterol
and ergosterol- type derivatives, and ergostane- type sterols as strophasterols E and
F, as well as pentacyclic triterpenoids., and the evaluation of their inhibitory effects
on NO production (Kikuchi et al., 2018). Cateni et al. (2022) summarized the chem ical structure composition of mycochemicals from P. eryngii as a diterpenoid named
eryngiolide A.
Phenolic compounds from P. eryngii possess antioxidant properties to sca-
venge free radicals, to prevent lipid peroxidation, and to chelate ferrous ions (Cateni
et al., 2022). Mishra et al. (2013) concluded that P. eryngii had the highest contents
of phenolic compounds among seven Pleurotus species. Lin et al. (2014) reported that
P. eryngii contains phenolic acids such as p- anisic acid, chlorogenic acid, ferulic acid,
p- hydroxybenzoic acid, sinapic acid, syringic acid and vanillic acid; it also contained
avonoids such as avanols (catechin, epicatechin), avanones (hesperidin), avonols
(myricetin, quercetin), and avonoid glycoside (rutin). Also, Souilem et al. (2017)
found that P. eryngii contains, p- coumaric acid, cinnamic acid, protocatechuic acid,
gallic acid, and phenols. Recently, Calabretti et al. (2021) showed that among commercial and wild- growing isolates from Southern Italy, higher amounts of ferulic and
gallic acids, epicatechin gallate and epigallocatechin gallate have been determined from
commercial isolates (0.88±0.08 mg/ g, 1.53±0.16 mg/ g, 0.39±0.04 mg/ g, and 0.51±0.05
mg/ g, respectively).
Edible medicinal mushroom P. eryngii is considered a new- generation food and
is of growing interest to consumers. King oysters are of signicant nutritional value
(i.e., relatively high content in proteins, vitamins, and minerals, low amount of fats).
Thanks to their low- fat content, king oysters are a low- calorie product and are classied
as a functional food. The king oysters are recommended for consumption by people
suffering from hypertension, high blood low- density lipoprotein, cholesterol, or

382 Wild Edible Plants
triglycerides, levels, obesity, metabolic diseases, and diabetes. Besides, fruiting body
and mycelial mass P. eryngii are characterized by a high content of biologically active
compounds, including (1,3)(1,6)- β- D- glucans, which are classied as dietary ber,
containing triterpenes, phenolic compounds, and sterols. They have a benecial effect
on the organism through the improvement of its overall health and nutritional level. In
addition, the biologically active constituents contained in P. eryngii exhibit anticancer,
antioxidant, antidiabetic, and immunomodulatory effects.
13.4.6 Stropharia rugosoannulata
Stropharia rugosoannulata Farl. ex Murrill (family Strophariaceae) (Figure 13.3F),
common name King Stropharia, wine- cap or wine- red stropharia, giant stropharia,
or garden giant, composter mushroom, burgundy mushroom, stone mushroom, is an
edible delicacy mushroom recommended for for cultivation and consumption by the
Food and Agriculture Organization of the United Nations (Huang et al., 2023). King
Stropharia has a rich “mushroomy” avor with undertones of red wine and potatoes, and the stipe has a stringy texture similar to asparagus. The species is native to
Southern Europe and the United States and is now widely grown around the world.
Currently it is cultivated at a large scale in many provinces in China (Gao et al., 2022;
Hu et al., 2021).
King Stropharia is a good raw material for eating and developing into healthy food,
which is high in protein, low in fat and ber, rich in minerals and B- group vitamins,
has a reasonable amino acid composition, contains water- soluble polysaccharides,
sterols, triterpenoids, phenols, and other biologically active substances (Hu et al., 2021;
Huang et al., 2023; Jiang et al., 2023; Wu et al., 2013). The functional properties of
S. rugosoannulata are associated with its chemical compounds. Both fungal mycelia
and fruiting bodies contain these promising bioactive compounds, the main ones are
monosaccharides, polysaccharides, sterols, lectins, avonoids, and phenols, which vary
in their properties based on extraction methods from different parts of mushrooms,
some of which have antioxidant, antibacterial, antitumor, and antidiabetic effects (Liu
et al., 2020a; Wang et al., 2021).
13.4.6.1 Components of Stropharia rugosoannulata with
proven pharmacological activities
Polysaccharides. The structure and biological activity of polysaccharides found in
S. rugosoannulata exhibit considerable diversity. The soluble polysaccharides are the
main bioactive components of S. rugosoannulata (Liu et al., 2020b; Wei et al., 2023).
Maximum yield of extracellular polysaccharides (EPS) production by S. rugosoannulata
was 9.97 g/ L (He et al., 2012). Authors have reported that EPS from S. rugosoannulata
exhibits high antitumor and antioxidative effects.
Zhai et al. (2013) reported the maximal EPS level produced on the 8th day of
submerged cultivation was 10.83 g/ L and the maximal value of biomass produced
on the 5th day was around 16.35 g/ L. Jing et al. (2022) showed that the contents of
polysaccharides in the tested S. rugosoannulata were 13.18%. Jiang et al. (2023)

Edible and Medicinal Mushrooms as an Eco-Friendly Source of Food 383
isolated a new polysaccharide (SR- 1) from S. rugosoannulata that stimulated immunological activities such as the growth of lymphocytes, and lymphocytes as well as the
secretion of IgA, IgD, and IgG by B lymphocytes. Polysaccharides isolated from
S. rugosoannulata possessed antioxidant activities against ABTS+ , DPPH, and OH–
radicals (Liu et al., 2020).
Wang et al. (2021) investigated the structural characteristics and antioxidant activities of polysaccharides from S. rugosoannulata under different drying methods. Hotair- dried was recommended to maintain the antioxidant activities of polysaccharides
for use as functional foods. Meanwhile, Wei et al. (2023) showed that freeze- drying
mushrooms provided also high preservation of bioactive components and strong antioxidant activity of the nal product.
Gao et al. (2022) extracted polysaccharides from S. rugosoannulata, named
SRF- 3, using a combination of freeze- thaw and hot water extraction methods, and its
hypolipidemic activity was determined. The high content of galactose, mannose, and
glucose in edible mushroom polysaccharides and the agreement of SRF- 3 to contain
these three monosaccharides, but with a higher content of galactose indicating that SRF3 is very different from other typical edible mushroom polysaccharides, which may be
related to the properties of the S. rugosoannulata itself or the method of polysaccharides
extraction. SRF- 3 exhibited free radical scavenging ability in antioxidant assays in vitro.
Lectin. Peptides. Zhang et al. (2014) puried a novel lectin (SRL) from S. rugosoannulata
fruiting bodies. They represented the rst protein isolated from this mushroom with
carbohydrate specicity, relatively high thermostability, and potent antiproliferative
activity. Compared with other lectins isolated from mushrooms, the identied lectin
exhibited antiproliferative activity against cancer cells (Hep G2 and L1210) and
displayed anti- HIV reverse transcriptase activity. Notably, these effects were found to
be more pronounced compared to lectins isolated from other mushroom species. The
potent antiproliferative activity and immunomodulatory effect of SRL could facilitate
its development into a cancer therapy agent.
Hypertension is a chronic health problem characterized by high blood pressure,
and poses signicant risks to heart disease, stroke, aneurysm, and renal failure. For the
treatment of hypertension use angiotensin- converting enzyme (ACE) inhibitors, such as
peptides. Li et al. (2022) studied the ACE inhibition mechanism by S. rugosoannulata
peptides prepared by ultrasound and their results showed that these mushroom peptides
can bind to zinc ions, critical amino acids, or amino acid residues in the ACE active
pocket, which at least inhibits the action of ACE (angiotensin- converting enzyme
inhibitory property).
Steroids and organic acid compounds. In addition to polysaccharides, S. rugosoannulata
also contains sterols, and various other compounds, all of which serve as active
ingredients in medicinal mushrooms (Jing et al., 2022; Liu et al., 2012). While
screening for antiendoplasmic reticulum (ER) stress and anti- methicillin- resistant
Staphylococcus aureus (MRSA) effects among extracts from various mushrooms, Wu
et al. (2012) discovered activity in the extract of S. rugosoannulata. They extracted
particular steroids from fresh basidiomes of S. rugosoannulata by ethyl alcohol and
acetone and discovered four novel steroids that have a very unique and unprecedented

384 Wild Edible Plants
carbon skeleton (Strophasterols A– D). Sterol A may protect neuronal cells and show
weak anti- methicillin- resistant Staphylococcus aureus activity. Thus, compounds with
ER stress, anti- MRSA, antifungal, and osteoclast formation- suppressing activities have
been isolated (Wu et al., 2012). Besides, Wu et al. (2013) isolated bioactive steroids
(strophasterols) from the King Stropharia fruiting bodies. Three phytotoxic compounds
were from the mushroom which in S. rugosoannulata were found to have an effect
(suppressed or promoted) on lettuce growth.
Another investigation of S. rugosoannulata fruiting bodies provided by Yan et al.
(2020), resulted in the isolation and identication of 16 compounds. These compounds
included six types of steroids, one type of steroidal saponins, three types of fatty acids,
and one type of each compound: alkane, ceramide, ester, pyrimidine, vitamins, and
avonoids. Nevertheless, the functional activities of these compounds remain unclear,
necessitating further investigation. Given the potential existence of undiscovered bioactive compounds, future studies could concentrate on elucidating the mechanisms and
actions of the components found in S. rugosoannulata that may confer health benets
to humans.
It has been noted that organic acids present in edible mushrooms contribute to
their complex and unique avor proles. Hu et al. (2020) reported the detection of
six types of organic acids (malic acid, ascorbic acid, fumaric acid, succinic acid,
acetic acid, and tartaric acid) in three varieties of dried S. rugosoannulata. The levels
of organic acids in S. rugosoannulata ranged from 111.03 mg/ g to 169.41 mg/ g,
surpassing those found in P. eryngii (5.64– 9.56 g/ 100 g DW), but lower than those
observed in L. edodes at various growth stages (8.54– 37.47g/ 100 g DW). Among the
six organic acids, malic acid and succinic acid emerged as the predominant ones in
S. rugosoannulata (Hu et al., 2020).
Phenolic compounds. Phenolics exhibit a wide range of biological effects including
antibacterial, anti- inammatory, antihyperglycemic, and antioxidant actions. The
overall mean of total phenols and avonoids in wild S. rugosoannulata was measured
as 5.52±0.45 mg/ g DW to 1.69±0.25 mg/ g DW, respectively (Liu et al., 2012). The main
phenolic components in S. rugosoannulata were detected as quercetin (13.28±2.44 g/
g), and p- coumaric acids (2.81±0.53 g/ g). Quercetin, one of the avonoids widely
found in mushrooms, as well as in some plants, has been reported to harbor various
physiological properties including antioxidant, anti- inammatory, antibacterial, and
anticancer activity. Also the antihyperglycemic properties using the ethanolic and
aqueous extract of the investigated mushrooms were evaluated.
Two different assays were carried out: a- glucosidase inhibitory (146.18±3.24 g/
mL) and a- amylase inhibitory activity (218.37±8.32 g/ mL). The ethanolic extracts
showed a higher antihyperglycemic activity than aqueous extracts. However, total phenolic and tocopherol in these investigated mushrooms had no obvious relevance with
reducing power and metal chelating activity as Barros et al. (2008) reported. Triterpenes
have antitumor activity, and the content of triterpene in S. rugosoannulata is 1.42%
(Jing et al., 2022).
S. rugosoannulata exhibits signicant potential for soil and water bioremediation.
Studies have investigated its capability to degrade pollutants in various environments,

Edible and Medicinal Mushrooms as an Eco-Friendly Source of Food 385
including soils contaminated with 2,4,6- trinitrotoluene (TNT), showcasing degradation
rates ranging from 13% to 73% and over 70% degradation of carbamazepine, indicating
its potential for remediation applications (Hu et al., 2021). It was shown that Stropharia
mushroom could be used for remediation of sites polluted with biotic contaminants,
in particular, in remediation of wetland water polluted with crow feces containing
antibiotic- resistant bacteria. The use of mycelial biolters to reduce the content of bacteria in water through bioltration and bioabsorption can nd wide practical application
(Sen et al., 2023).
Research conducted on S. rugosoannulata has unveiled its potential applications
as a functional and medicinal food. This fungus has shown promise in suppressing
osteoclast formation and exhibiting properties such as antidiabetic, antimicrobial,
antioxidant, antiproliferative, antitumor, and immunomodulatory effects. Moreover,
being a high- protein, low- fat food rich in minerals, vitamins, and dietary ber, King
Stropharia mushrooms serve as excellent raw material for functional components like
fungal polysaccharides and taste peptides. They can be processed into a variety of delicious foods.
13.5 CONCLUSIONS
One of the global problems today is the advancement of techniques for obtaining
environmentally friendly food products and therapeutic medical treatments with
immunostimulating, radioprotective, anticancer, and general strengthening effects,
based on using natural reserves of wild plants. Since ancient times, people have known
about the benecial properties of plants and fungi. Over time, they learned to cultivate
wild mushrooms throughout the year, using available substrates from agricultural and
forestry waste, instead of picking them as a seasonal food.
Mushrooms have long been used by humans not only for food but also as medicines
against various diseases. The experience of traditional medicine has become a stimulus
for modern scientic research in developing and obtaining pharmacological preparations
by cultivating certain medicinal macromycetes. Recently this theme has attracted the
greatest attention of specialists from different countries, as seen in many experimental
works, but also reviews and monographs devoted to various aspects of the study of
medicinal mushrooms.
With scientic development, it became known about some advantages of mushroom consumption. Edible mushrooms with a high protein and carbohydrates (ber)
content in the fruiting bodies and mycelium, low in fat and calories, with complete
essential amino and fatty acids, macro and microelements proles are desirable products
in any diet. Due to mushrooms’ bioactive compounds such as immune- modulating
polysaccharides, triterpenes, phenolic compounds, and many others, which are nontoxicity, mushrooms are promising in preventing and treating various diseases, such as
diabetes, hypertension, hypercholesterolemia, cardiovascular diseases, and others, and
also a source of health- promising products for further using them as functional, dietary,
and nutraceutical products.
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