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163

Mushroom Species and Classification


Bioactives in Poisonous and Edible Mushrooms
Sadia Zafar1, Farhat Jabeen2, Muhammad Akram3, Zarfishan Riaz3,
and Naveed Munir4
1
Department of Botany, Division of Science and Technology, University of Education, Lahore, Pakistan
2
Department of Zoology, Government College University Faisalabad, Faisalabad, Pakistan
3
Department of Eastern Medicine, Government College University Faisalabad, Faisalabad, Pakistan
4
Department of Biochemistry, Government College University Faisalabad, Faisalabad, Pakistan

CONTENTS
8.1 Introduction, 163
8.2 Classification of Mushrooms, 164
8.2.1 Edible Mushrooms, 165
8.2.2 Non-Edible Mushrooms, 165
8.3 Bioactive Agents in Mushroom Species, 165
8.4 Bioactive Agents in Non-Edible Mushroom Species, 166
8.4.1 Polysaccharides, 166
8.4.2 Glucans, 166
8.4.3 Polysaccharide–Protein Complexes, 174
8.4.4 Terpenes, 174
8.4.5 Phenolic Compounds, 175
8.4.6 Peptides and Proteins, 176
8.5 Other Bioactive Compounds of Mushroom Species, 176
8.6 Conclusion, 176

8.1 ­Introduction

The term mushroom is generally used to indicate a stem or stalk, a cap (pileus),
and the gills (lamellae). The part of the fungus that is usually visible is the sporo-
phore, which is an umbrella-shaped structure that has fertile surfaces on its

Poisonous Plants and Phytochemicals in Drug Discovery, First Edition.


Edited by Andrew G. Mtewa, Chukwuebuka Egbuna, and G.M. Narasimha Rao.
© 2021 John Wiley & Sons, Inc. Published 2021 by John Wiley & Sons, Inc.
164 8 Mushroom Species and Classification

lamellae. The lamellate fungi are commonly called mushrooms or toadstools,


according to whether they are edible or non-edible, respectively [1]. Mushrooms
are ancient species and have had different names; for example, Theophrastus
used “truffles” for mushroom fruiting bodies. The Romans viewed mushrooms
as “ambrosia,” which means heavenly food or the food of Gods and only used
mushrooms for special celebratory events, whereas the Chinese view mushrooms
as a “health food.” Since the Early Middle Ages, mushrooms have been cultivated
according to their importance [2]. The genus Amanita is known as the death cap
or destroying angel and has killed several mushroom hunters [2]. In this chapter,
we will ­discuss edible and non-edible mushrooms, their bioactivities, and
their uses.

8.2 ­Classification of Mushrooms

According to modern classification, which is based on rDNA sequences, the


­kingdom Fungi is divided into four phyla: Zygomycota, Chytridiomycota,
Basidiomycota, and Ascomycota. The phyla Ascomycota and Basidiomycota are
the main phyla with fruiting bodies and are considered to be mushrooms. The
phylum Ascomycota has approximately 40 000 different types of species around
the world. These are also known as sac fungi and ascomycetes. Their spores, called
ascospores, are found in the ascus, which is a sac-like structure. Ascospores divide
by meiosis and haploid spores are produced; this type of asexual reproduction is
observed only in mushrooms, single-celled yeasts, and truffles [2, 3].
The members of the phylum Basidiomycota are also known as basidiomycetes
or club fungi. The spores of Basidiomycota are known as basidia. Smut and rust
are dominating members that cause numerous diseases in grains. Order
Agaricales (genus Agaricus; species Agaricus campestris [field mushroom]) of the
phylum Basidiomycota is the most commonly found and has gills on the fruiting
body; these are called gilled mushrooms. Globally, Agaricus bisporus is known as
a button mushroom, and is the most widely cultivated mushroom. Marasmius
oreades is a fairy ring mushroom belonging to the order Agaricales, and is also
commonly found. Hypomyces lactifluorum (lobster mushroom) belongs to the
phylum Ascomycota. It parasitizes other mushrooms such as Russula and
Lactarius, forming on the outer side of the organism with the white flesh of the
parasitized basidiomycete mushroom remaining on the inner side [2, 4].
Mushrooms (Basidiomycota) produce spores (basidiospore) on the basidium.
The spores are haploid and become binucleate at maturity or remain uninucleate.
After ripening of the spores, they are released from the fruiting body and start to
germinate and form hyphae, which develop into the mycelium [5]. In some cases,
two compatible mycelia fuse and a secondary mycelium develops that is binucle-
ate and develops into fruiting bodies.
8.3 ­Bioactive Agents in Mushroom Specie 165

8.2.1 Edible Mushrooms


It is estimated that a total of 140 000 species of mushrooms have been discovered,
out of which around 14 000 species have been described [6]. Less than 10% of the
described species are edible mushrooms and 700 species are used in pharmacology
[7]. However, data for edible mushrooms have been collected for culinary purposes
around the world, the number of which varies greatly from 200 to 3000 mushroom
species. Many wild and cultivated mushroom species are consumed globally, in
approximately equal amounts. Both fresh and preserved mushrooms are also con-
sumed because of their particular texture and aroma and also because of their
lower fiber content and energy. The demand for mushrooms becomes higher when
there is a shortage of staple foods, such as during wars [8]. About 100 species are
cultivated commercially; of these, 10–20 species are cultivated on an industrial
scale [6]. The mushroom industry can be divided into three parts: wild growing,
cultivated medicinal, and edible mushrooms [8].

8.2.2 Non-Edible Mushrooms


Numerous mushroom species have psychoactive toxins. They are poisonous and a
serious threat to health [9]. Some mushrooms are very toxic and are dangerous
even in very small quantities. Many toxins are well explained in the literature,
such as amatoxin, which causes damage to the kidneys and liver, and the toxin
orellanine, which is nephrotoxic in nature [9, 10]. Some species are consumed for
food and medicine because they contain pharmacologically active compounds.
Also, mushrooms have been used in traditional medicine since ancient times
because of the pharmacological compounds they contain [11, 12]. However, some
of the pharmacological compounds were found to be toxic. In fact, only 70–80
mushroom species are fatal out of all the poisonous mushroom species; however,
unfortunately, several poisonous mushroom species are morphologically similar
to edible mushrooms [13].

8.3 ­Bioactive Agents in Mushroom Species

The demand for mushrooms is increasing very rapidly because of the presence of
the biologically active compounds that are very beneficial for human health,
either directly or indirectly [14]. The biologically active compounds that mush-
rooms contain are also present in cell walls, such as proteins, β-glucans (polysac-
charides), and other secondary metabolites like terpenoids, steroids, and
phenolics. Bioactive compounds vary greatly from species to species or even
within a single mushroom, depending upon the concentration of the bioactive
compounds, developmental stage, conditions of the fruiting body, the ­mushroom’s
166 8 Mushroom Species and Classification

age, and the storage conditions [15]. Previous studies have described the diversity
of biologically active compounds and their action in the medicinal field as, for
example, antioxidant, anti-inflammatory, antitumor, antiviral, antifungal, anti-
bacterial, anti-diabetic, immunomodulatory agents [16–19]. Bioactive ­compounds
that are found in mushrooms are described in Tables 8.1 and 8.2.

8.4 ­Bioactive Agents in Non-Edible


Mushroom Species

Both poisonous and non-poisonous mushrooms contain various phytocom-


pounds, some of which are toxins, particularly in wild mushrooms. The following
sections describe various components of bioactive agents in both poisonous and
edible mushrooms.

8.4.1 Polysaccharides
Edible mushrooms contain polysaccharides. The polysaccharides can be used as
immunomodulator, anticancer, anti-inflammatory, antimicrobial, antidiabetic,
and antioxidant agents [18, 34, 126].

8.4.2 Glucans
Glucan polysaccharides vary in molecular weight, primary structure, degree of
branching, solubility, type of linkage, etc. The fungal glucans can be insoluble or
soluble in alkali but are soluble in water. Glucans help to preserve material and
are intracellular; other glucans are released in the medium; and some exist in the
cell wall [127]. For those glucans that are insoluble, they classically constitute the
cell wall’s structural components and make cross-linkages to other polysaccha-
rides (chitin, proteins). Soluble and insoluble glucans represent 20–50% and
50–80% of total glucans, respectively [128]. Two glucose units can connect in a
minimum of eight different ways in a variety of glucans. As a result of condensa-
tion reactions, α or β bonds are formed. On cyclic sugar structures, different
branches of chains and length further increase the variety of glucans [129]. In the
cell wall of fungi α- and β-glucans are present. Pleurotus pulmonarius fruit extract
displayed a mixture of α- and β-anomeric carbon links, while mycelial extracts
had mostly α-glucan connections in polysaccharides [130, 131]. In numerous
basidiomycetes 9–46% of α-1,3-glucan is present in the cell wall. It can exist in the
cell wall of various mushrooms such as in the fruiting bodies of A. bisporus [132].
The main constituent of the cell wall in numerous ascomycetes and basidiomy-
cetes is β-glucan. This is a long-chain polysaccharide with β-d-glucose as the basic
Table 8.1 Bioactive compounds of edible mushroom species.

Scientific name Phylum Bioactive compounds Action of bioactive compounds References

Agaricus bisporus Basidiomycota Pyrogallolhydroxyl benzoic Anti-inflammatory activity [20, 21]


acid-derived flavonoids
Agaricus macrosporus Basidiomycota Agaricoglycerides Anti-inflammatory activity [22]
Agaricus subrufescens Basidiomycota Glucan, glycoprotein, Modulate the immune response [23–25]
polysaccharide, and protein
segments
Agrocybe cylindracea Basidiomycota Agrocybin, β-glucans Hypoglycemic, antioxidant, and [14, 26]
antifungal activity
Albatrellus ovinus Basidiomycota Grifolin and its derivatives Anti-inflammatory and [27]
antioxidant activity
Albatrellus caeruleo Basidiomycota Phenolic compounds, grifolinones Anti-inflammatory activity [27, 28]
porus A, B
Auricularia auricula Basidiomycota Glucans Immunomodulatory, anti- [29]
inflammatory, hyperglycemic,
and anticancer activity
Boletus edulis Basidiomycota Polysaccharides Anti-inflammatory activity [20]
Boletus spp. Basidiomycota Glutamyl tryptophan, lithocholic, Antioxidant activity [30]
2,4,6-trimethyl acetophenone
imine, glycine conjugate, and
azatadine
Cantharellus cibarius Basidiomycota Flavonoids, polysaccharides, Anti-inflammatory, [20, 31, 32]
pyrogallol, caffeic acid, catechin antimicrobial, and antioxidant
activity
(Continued)
Table 8.1 (Continued)

Scientific name Phylum Bioactive compounds Action of bioactive compounds References

Calvatia gigantea Basidiomycota Calvacin mucoprotein Anticancer activity [14]


Caripia montagnei Basidiomycota Glucans, polysaccharides Anti-inflammatory activity [33]
Coprinus comatus Basidiomycota Polysaccharide and protein Modulate the immune response [34]
fractions, β-1,3-glucan
Cordyceps militaris Ascomycota Cordycepin, cordymin Antiangiogenic, anti- [35–37]
inflammatory, and antitumor
activity
Cordyceps sinensis Ascomycota ciclosporin, cordycepin, cordymin Antioxidant, [38–41]
(peptide) immunosuppressive, and
anti-inflammatory activity
Craterellus Basidiomycota Myricetin (phenolic compound) Antioxidant activity [32]
cornucopioides
Craterellus Basidiomycota Polysaccharides (starch, glycogen, Anti-inflammatory activity [42]
tubaeformis cellulose)
Dictyophora indusiata Basidiomycota Dictyoquinazol, dictyophorines A Antineurodegenerative and [43]
and B neuroprotective activity
Flammulina velutipes Basidiomycota Peptidoglycan, polysaccharides, and Anti-inflammatory, antiviral, and [44–47]
flammulin (protein) antitumor activity
Ganoderma tsugae Ascomycota Fungal immunomodulatory Modulate the immune response [48]
proteins (FIP)
Grifola frondosa Basidiomycota Grifolan, glucoxylan, proteoglycan, Immunomodulatory, antiviral, [22, 49, 50]
galactomannan, anticancer, anti-inflammatory,
mannogalactofucan, heteroglycan, and hepato-protective activity
agricoglycerides, and
fucomannogalactan
Scientific name Phylum Bioactive compounds Action of bioactive compounds References

Hericium erinaceus Basidiomycota Phenol-analogous compounds, Antioxidant, anticancer, [19, 51–55]


hericenones, erinacines, hericerins, antifatigue, antiviral,
dilinoleoyl antihyperglycemic, antidiabetic,
phosphatidylethanolamine, antihypertensive,
resorcinols, heteroglycan peptide, antisenescence,
steroids, terpenoids, lectin immunomodulatory,
(glycoprotein) neuroprotective,
cardioprotective, and
antineurodegenerative activity
Hypsizygus Basidiomycota Ergosterol, mannitol, trehalose, Antifungal, anti-inflammatory, [56–58]
marmoreus methionine, marmorin, phenolic antitumor, antioxidant,
compounds, flavonoids antibacterial, and antiallergic
activity
Inonotus obliquus Basidiomycota Mannogalactoglucan, sterols, Anti-inflammatory, anticancer, [14, 20,
β-d-glucans, triterpenes antioxidant, and gastrointestinal 59–62]
disease activity
Lactarius deliciosus Basidiomycota Polysaccharides, pyrogallol, Anti-inflammatory activity [63]
flavonoids
Lactarius rufus Basidiomycota 1,3- and 6-β-d-glucans Anti-inflammatory activity [64]
Lentinula edodes Basidiomycota Lentinan, glucan, mannoglucan, Immunomodulatory, antitumor, [58, 65–67]
fucomannogalactan, lentin anti-inflammatory, antifungal,
(protein), catechin (phenolic antibacterial, antifungal, and
compound), phenolic compounds, antioxidant activity
flavonoids
Lyophyllum decastes Basidiomycota 1,3- and 6-β-d-glucans Anti-inflammatory activity [68]
Morchella esculenta Ascomycota Heteroglycan, galactomannan, Anticancer and hyperglycemic [69]
β-1,3-d-glucan activity
Pholiota adiposa Basidiomycota Lectin Antiviral and anticancer activity [70]

(Continued)
Table 8.1 (Continued)

Scientific name Phylum Bioactive compounds Action of bioactive compounds References

Pholiota nameko Basidiomycota Polysaccharides Anti-inflammatory activity [71]


Pleurotus Basidiomycota Glycoprotein Anticancer activity [72]
citrinopileatus
Pleurotus eryngii Basidiomycota Laccase Antiviral activity [73]
Pleurotus florida Basidiomycota β-Glucan Antioxidant activity [74]
Pleurotus ostreatus Basidiomycota Pleuran (β-1,3-glucan with Immunomodulatory, antifungal, [75–78]
galactose and mannose), antioxidant, antiviral,
proteoglycan, pleurostrin, and hyperglycemic, and anticancer
laccase activity
Pleurotus pulmonarius Basidiomycota β-(1,3)-glucopyranosyl and 1,3- and Anti-inflammatory activity [79, 80]
6-β-d-glucans
Sparassis crispa Basidiomycota β-Glucan Immunomodulatory activity [81, 82]
Termitomyces Basidiomycota Termitomycesphin, Antineurodegenerative activity [83, 84]
albuminosus termitomycamides (fatty acid
amides)
Trametes versicolor Basidiomycota Krestin, coriolan (β-glucan protein Antimetastatic hypoglycemic [12, 14]
complex) effect
Tremella aurantia Basidiomycota Heteroglycan Modulate the immune response [85]
alba (heteropolysaccharides)
Tremella mesenterica Basidiomycota Glucuronoxylomannan Hypoglycemic and [26]
immunomodulatory activity
Tricholoma giganteum Basidiomycota Trichogin Antifungal activity [86]
Tricholoma Basidiomycota Laccase Anticancer and antiviral activity [87]
mongolicum
Volvariella volvacea Basidiomycota FIP-Volvariella volvacea Immunomodulatory activity [88]
Table 8.2 Bioactive compounds of non-edible mushroom species.

Scientific name Phylum Bioactive compounds Bioactivity References

Antrodia camphorata Basidiomycota Glycoprotein ACA, diterpenes Immunomodulatory and [89, 90]
neuroprotective activity
Clitocybe maxima Basidiomycota Laccase Antitumor activity [91]
Cortinarius infractus Basidiomycota 6-hydroxyinfractine and Antineurodegenerative activity [92, 93]
infractopicrine
Cyathus africanus Basidiomycota Diterpenoid (cyathatriol, Anti-inflammatory activity [94]
11-O-acetylcyathatriol, and
neosarcodonin)
Daldinia concentrica Ascomycota 1-(3,4,5-trimethoxyphenyl) Neuroprotective activity [95]
ethanol, caruilignan C
Elaphomyces granulatus Ascomycota Syringaldehyde, syringic acid Anti-inflammatory activity [96]
Fomitopsis pinicola Basidiomycota Polysaccharides Anti-inflammatory activity [97]
Ganoderma lucidum Basidiomycota Ganoderic acids, ganoderiol, Antimetastatic, hypoglycemic, [14, 98–108]
ganodermanontriol, ganoderan anti-HIV, hepatoprotective,
A and B, ganopoly, triterpenes, anti-inflammatory, antiviral,
lucidenic acids, ganoderic acids, immunomodulatory, and
lanostane-type triterpenic acids; antitumor activity
lingzhi-8 (protein), ganodermin
(protein), Se-containing protein
Ganoderma Basidiomycota Protein GMI Immunomodulatory activity [109]
microsporum
Ganoderma pfeifferi Basidiomycota Sesquiterpenoid hydroquinones Antiviral, antibacterial, and [110]
(lucialdehyde D, ganoderone A, antifungal activity
ganoderone C)
(Continued)
Table 8.2 (Continued)

Scientific name Phylum Bioactive compounds Bioactivity References

Geastrum saccatum Basidiomycota β-glucans, polysaccharides Anti-inflammatory activity [111]


Ganoderma tsugae Order: Polyporales, FIP-gts protein Immunomodulatory activity [48]
Class: Agaricomycetes
Lentinus polychrous Basidiomycota Catechin (flavan-3-ol) Antioxidant activity [66]
Lentinus squarrosulus Basidiomycota Catechin (flavan-3-ol) Antioxidant activity [66]
Lenzites betulina Basidiomycota Betulinan A Antioxidant activity [14]
Lignosus rhinocerus Basidiomycota Polysaccharides–protein Anticancer activity [26]
Phellinus linteus Basidiomycota Glucans, acidic polysaccharides, Antitumor, immunomodulatory, [39, 112–114]
hispidin (polyphenol) and antioxidant activity
Psilocybe cubensis Basidiomycota Psilocybin Antidepressant activity [115–119]
Psilocybe samuiensis Basidiomycota Psilocybin Sedative [115–119]
Psilocybe mexicana Basidiomycota Psilocybin Sedative [115–119]
Russula lepida Basidiomycota Lectin (proteins) Antitumor activity [120]
Schizophyllum commune Basidiomycota Schizophyllan, Immunomodulatory and [121, 122]
1,6-monoglucosyl-branched antitumor activity
β-1,3-d-glucan
Wolfiporia cocos Basidiomycota Dehydrotrametenolic acid, Hypoglycemic and anti- [14, 123, 124]
lanostane inflammatory activity
Xylaria hypoxylon Ascomycota Lectin (glycoprotein) Antitumor and antimitogenic [125]
activity

ACA, anti-cardio0lipin antibodies; GMI-protein, Ganoderma microsporum immunomodulatory protein; HIV, human immunodeficiency virus;
-gts, from Ganoderma tsugae.
8.4 ­Bioactive Agents in Non-Edible Mushroom Specie 173

subunit in β-glucan. These β-glucans have shown an antimicrobial immune


response on numerous immune receptors such as dectin-1, which is the main
β-glucan receptor, innate immune receptor, and complement receptor [34]. This is
why β-glucans have the ability to boost the immune system, prevent various com-
mon diseases, and promote health [133]. β-Glucan is attached to macrophages in
the innate immune system, which is responsible for distinguishing invaders and
managing the defense system of the body. When they leave the bloodstream,
monocytes become macrophages, which are then activated by β-glucan; this
boosts their ability to recognize and destroy invaders by the process of phagocyto-
sis. The thymus gland produces T lymphocytes, which contain T-cell receptors for
antigens and have specialized cells whose aim is to kill pathogens. B lymphocytes
play a vital role in humoral immunity and make antibodies. T lymphocytes are
natural killer cells that destroy bacteria, infected cells, tumor cells, and viruses.
Thus, these types of white blood cells help the immune system to defend the body
against destructive pathogens [34, 134]. Several biologically active fungal β-glucans
have been found in the fruiting bodies of mushrooms and have been isolated [33,
64, 71, 111, 135]. For example, pleuran has been isolated from Pleurotus ostreatus
and consists of β-(1,4)- or β-(1,6)-branches for every fourth β-(1,3)-glucan back-
bone [136]. Lentinan is derived from Lentinula edodes; it has a molecular weight
of 400–1000 kDa and has shown antitumor and immunomodulatory activity [23].
The active β-glucan schizophyllan from Schizophyllum commune has a molecular
weight of 450 kDa [121]. In Grifola frondosa maitake the d-fraction was found to
consist of a combination of a main chain of β-1,6-glucan with branched β-1,4-
glucan and a main chain of β-1,3-glucan with branched β-1,6-glucan [49]. Extract
of Agaricus subrufescens, which contains β-1,3-, β-1,4-, and β-1,6-glucans, induces
proinflammatory cytokine release in monocytes and veins of human endothelial
cells [137]. Anti-inflammatory properties are also seen in P. pulmonarius, which
contains (1,3)-glucopyranosyl that is responsible for this action [80]. The glucans
ganoderan A and B from Ganoderma lucidum show hypoglycemic properties, as
reported by Rathee et al. [14]. Moreover, ganopoly of G. lucidum revealed hepato-
protective actions in patients with chronic hepatitis B [102]. Immunomodulating
properties have also been demonstrated for G. lucidum glucans. They enhance the
proliferation of lymphocytes and production of antibodies. Antigenotoxic and
antitumor activities are also displayed by these polysaccharides [12, 138]. These
studies showed that G. lucidum has antioxidative and scavenging effects [14].
Antitumor activity against HeLa tumor cells has also been confirmed in P. ostrea-
tus fruiting bodies, which contain a β-glucan that is responsible for this action
[76, 139]. β-Glucan has two mechanisms that are responsible for the anticancer
properties: (i) directly through cytotoxic activity and (ii) indirectly via immune
modulation [34]. The anti-inflammatory action of L. edodes has also been reported.
Its active fraction was made by chain reaction of α-d-galactopyranosyl-1,6-linked
174 8 Mushroom Species and Classification

units with fucomannogalactan, incompletely substituted at O-2 [140]. Moreover,


it is reported that 1,3-d-glucopyranosyl glucans of P. pulmonarius exhibit anti-
inflammatory activity [80]. Wu et al. [46] reported that Flammulina velutipes
­polysaccharides consist of three monosaccharides, namely xylose, glucose, and
mannose in a 3.5 : 0.8 : 1.4 molar ratio, that have anti-inflammatory properties.
This study reported that mushrooms contain polysaccharides that have anti-
inflammatory activity.

8.4.3 Polysaccharide–Protein Complexes


Polysaccharide–protein complexes have also been recognized as polysaccha-
rides that possess immune modulatory and antitumor properties. For example,
polysaccharide-K protein, also known as krestin, was obtained from a few
mushroom species that exhibited antimetastatic action [12, 141]. Hypoglycemic
effects have been shown for Trametes versicolor, a type of mold that affects
waterlogged trees [14]. Calvacin, which is a mucoprotein that has shown antitu-
mor activity, is obtained from Calvatia gigantea [142], whereas Phellinus linteus
contains a proteoglycan and its ethanolic extracts show anti-inflammatory
activities [143, 144].

8.4.4 Terpenes
The terpenes present in mushrooms constitute the major group of anti-inflam-
matory compounds. The numerous terpenes obtained from mushrooms are
non-polar metabolites, including mono- and sesquiterpenes, volatile oils,
carotenoid ­pigments, involatile triterpenoids and sterols, and less volatile dit-
erpenes. Four ganoderic acids and nine lucidenic acids have been isolated
from mushroom fruiting bodies by Akihisa et al. [106] and Iwatsuki et al.
[105], whereas numerous triterpenic acids (lanostane-type) and terpenoids
were isolated from Reishi mushrooms [103, 106]. Anti-inflammatory proper-
ties are shown by all of these terpenes. In addition, biosynthesis of cholesterol
is inhibited by some triterpenes [145]. The studies reported that mushrooms
contain other triterpenes, which give protection against atherosclerosis and
provide antiviral and antioxidative protection [14, 146]. Some sterol com-
pounds have been isolated that show antiviral, antifungal, antibacterial, and
potent anti-inflammatory properties [10, 59, 60, 110]. Numerous terpenes
with anticancer and anti-inflammatory properties have been isolated from the
sclerotia of Inonotus obliquus [62]. Also, three diterpenes have been isolated
from Cyathus africans, that is, neosarcodonin, 11-O-acetylcyathatriol, and
cyathatriol, and five novel cyathane diterpenes have also been isolated that
8.4 ­Bioactive Agents in Non-Edible Mushroom Specie 175

have potent anti-inflammatory properties [94]. One study [90] reported that
numerous triterpenes that showed neuroprotective activity were isolated from
Antrodia campho.

8.4.5 Phenolic Compounds


The phenolic complexes are aromatic hydroxylated composites that have one or
more hydroxyl groups and aromatic rings. These phenolic compounds contain
hydroxybenzoic acids, phenolic acids, flavonoids, lignans, hydroxycinnamic
acids, tannins, oxidized polyphenols, and stilbenes [147, 148]. These studies have
demonstrated that these phenolic compounds have antioxidant properties: they
are inhibitors of free radicals, decomposers of peroxide, oxygen scavengers, and
metal inactivators. Thus, antioxidants play a vital role in the response to free
radicals, which possess one or more unpaired electrons and are chemical com-
pounds. Reactive oxygen species (ROS) can be formed either by external sources
or by metal-catalyzed oxidation and oxidoreductase enzymes, or by aerobic res-
piration during mitochondrial electron transport as a by-product. Radicals find
ways to pair with their electrons because of their reactive properties, which is
why radicals repeatedly attack adjacent chemical complexes. Apoptosis and cell
function are lost if radicals change the chemical structure of the cells. Numerous
non-­enzymatic small molecules perform a role as antioxidants. The most vital
intracellular defense against the harmful properties of ROS is glutathione. This is
a tripeptide which presents a sulfhydryl group as a goal for attack. Polyphenol,
vitamins C and E, and lycopene have the ability to reduce ROS [149]. The antioxi-
dant properties of phenolic complexes have been studied by Palacios et al. [32] in
numerous species of mushroom, as shown in Table 8.1. The maximum amount
of myricetin is present in Craterellus cornucopioides and large amounts of cate-
chin and caffeic acid are present in Cantharellus cibarius. Anti-inflammatory
properties have also been demonstrated for the phenolic molecule pyrogallol
from Lactarius deliciosus, A. bisporus, and Cantharellus cibaromius [31, 139, 150].
Catechin, a major phenolic ­compound that is extracted from Lentinula squarro-
sulus, Lentinula polychrous, and L. edodes, has antioxidant properties. Chowdhury
et al. [58] isolated flavonoids and phenolic compounds from fungi and demon-
strated that they had antibacterial, antioxidant, and antifungal properties.
Mushrooms reduce the risk of neurodegenerative diseases [19]. The synthesis of
nerve growth factor has been reported from hericenones and erinacines pro-
duced by the fruiting body and the mycelium of Hericium erinaceus [54, 151]. It
has been reported that hispidin, an important medicinal metabolite that belongs
to a class of polyphenols that show ROS scavenger properties, was isolated from
Phellinus spp. [114, 152].
176 8 Mushroom Species and Classification

8.4.6 Peptides and Proteins


Different bioactive peptides and proteins are produced by mushrooms. These
essentially include lectins, in which enzymatic activity is absent, and some ­proteins,
such as fungal immune modulatory proteins, laccases, and ribosome-inactivating
proteins that possess enzymatic activity. Chu et al. [75] demonstrated antifungal
activity in P. ostreatus and in Agrocybe cylindracea [153]. Peptide from Russule palu-
dosa exhibited antiviral properties [154]. Cordymin, which has inflammatory
activity and a low molecular weight, has been isolated from Cordyceps sinensis [40,
41] and Cordyceps militaries [106, 155]. Some enzymes deactivate ribosomes, such
as ribosome-inactivating proteins, which eliminate adenosine remnants from
rRNA. It has been stated that these ribosome-inactivating proteins exhibited anti-
tumor activity [56]. Ascomycete and basidiomycete fungi use laccases, which are
phenol oxidases, to reduce lignocellulosic substrates.

8.5 ­Other Bioactive Compounds


of Mushroom Species

Agaric glycerides, which contain glycerol and an ester of 4-hydroxybenzoic acid


(chlorinated), are fungal secondary metabolites that show strong anti-inflamma-
tory properties [22]. Dilinoleoyl phosphatidylethanolamine is obtained from
H. erinaceum fruiting bodies; this reduces oxidative stress and has a strong effect
on neurodegenerative diseases, as reported by Nagai et al. [51]. It has been
reported that termitomycesphins and termitomycamides were extracted from
Termitomyces albuminosus dried fruiting bodies [83]. These biologically active
compounds have antineurodegenerative properties [156].

8.6 ­Conclusion

Mushrooms exist in two broad categories: poisonous and edible. Poisonous mush-
rooms are mostly found in the wild and their toxicity is usually fatal. Within both
of the groups are phytocomponents with bioactive properties. Eating mushrooms
remains a risk, particularly to people who are not familiar with the properties of
species that are available to them. It is therefore imperative for food scientists and
other allied researchers to identify these groups of mushrooms and present the
information to the communities that use them. In terms of drug discovery, most
alkaloid toxins tend to make good drugs or drug scaffolds. It is also important for
researchers to explore this area for the development of drugs that can be used
­reliably against various ailments.
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