sustainability

Review
Edible Mushrooms for Sustainable and Healthy Human Food:
Nutritional and Medicinal Attributes
Hassan El-Ramady 1,2, * , Neama Abdalla 3 , Khandsuren Badgar 1, * , Xhensila Llanaj 1 , Gréta Törős 1 ,
Peter Hajdú 1 , Yahya Eid 4 and József Prokisch 1

1 Faculty of Agricultural and Food Sciences and Environmental Management, Institute of Animal Science,
Biotechnology and Nature Conservation, University of Debrecen, 138 Böszörményi Street,
4032 Debrecen, Hungary; [email protected] (X.L.); [email protected] (G.T.);
[email protected] (P.H.); [email protected] (J.P.)
2 Soil and Water Department, Faculty of Agriculture, Kafrelsheikh University, Kafr El-Sheikh 33516, Egypt
3 Plant Biotechnology Department, Biotechnology Research Institute, National Research Centre,
33 El Buhouth St., Dokki, Giza 12622, Egypt; [email protected]
4 Poultry Department, Faculty of Agriculture, Kafrelsheikh University, Kafr El-Sheikh 33516, Egypt;
[email protected]
* Correspondence: [email protected] (H.E.-R.); [email protected] (K.B.)

Abstract: Global food production faces many challenges, including climate change, a water crisis,
land degradation, and desertification. These challenges require research into non-traditional sources
of human foods. Edible mushrooms are considered an important next-generation healthy food
source. Edible mushrooms are rich in proteins, dietary fiber, vitamins, minerals, and other bioactive
components (alkaloids, lactones, polysaccharides, polyphenolic compounds, sesquiterpenes, sterols,
and terpenoids). Several bioactive ingredients can be extracted from edible mushrooms and incorpo-
rated into health-promoting supplements. It has been suggested that several human diseases can
be treated with extracts from edible mushrooms, as these extracts have biological effects including
Citation: El-Ramady, H.; Abdalla, N.; anticancer, antidiabetic, antiviral, antioxidant, hepatoprotective, immune-potentiating, and hypo-
Badgar, K.; Llanaj, X.; Törős, G.;
cholesterolemic influences. The current study focuses on sustainable approaches for handling edible
Hajdú, P.; Eid, Y.; Prokisch, J. Edible
mushrooms and their secondary metabolites, including biofortification. Comparisons between edible
Mushrooms for Sustainable and
and poisonous mushrooms, as well as the common species of edible mushrooms and their different
Healthy Human Food: Nutritional
bioactive ingredients, are crucial. Nutritional values and the health benefits of edible mushrooms, as
and Medicinal Attributes.
Sustainability 2022, 14, 4941. https://
well as different biomedical applications, have been also emphasized. Further research is needed to
doi.org/10.3390/su14094941 explore the economic sustainability of different medicinal mushroom bioactive compound extracts
and their potential applications against emerging diseases such as COVID-19. New approaches such
Academic Editor: Dario Donno
as nano-biofortification are also needed to supply edible mushrooms with essential nutrients and/or
Received: 18 March 2022 to increase their bioactive ingredients.
Accepted: 18 April 2022
Published: 20 April 2022 Keywords: anticancer; antidiabetic; antioxidants; phenolics; poisonous mushroom; biomolecules
Publisher’s Note: MDPI stays neutral
with regard to jurisdictional claims in
published maps and institutional affil-
iations. 1. Introduction
More than 14,000 mushroom species have been identified, about 10% of which are
macro-fungi. Edible mushrooms are macro-fungi that can be seen with the naked eye, i.e.,
fleshy fruit bodies of many species of mushrooms, and they have already been widely used
Copyright: © 2022 by the authors. in food and medicine due to their delicious taste and diverse physiological activities [1].
Licensee MDPI, Basel, Switzerland.
The fruiting bodies of edible mushrooms, which include about 700 species, can be safely
This article is an open access article
consumed and are considered beneficial to human health [2]. Edible mushrooms are seen as
distributed under the terms and
an important source of food because of their high content of polysaccharides, dietary fiber,
conditions of the Creative Commons
proteo-glucans, and vitamins such as riboflavin and thiamine [3–5]. Edible mushrooms
Attribution (CC BY) license (https://
also have many bioactive ingredients that are beneficial for human health such as antioxi-
creativecommons.org/licenses/by/
4.0/).
dants [6], terpenoids [7], lectins [8], phenolics/polyphenolics [9], polysaccharides [10], and

Sustainability 2022, 14, 4941. https://doi.org/10.3390/su14094941 https://www.mdpi.com/journal/sustainability

Sustainability 2022, 14, 4941 2 of 30

ergosterols [11,12]. Not much is known regarding the capability of mushrooms to treat
and/or prevent chronic diseases. However, several chronic diseases have been treated by
consuming edible mushrooms (e.g., Coriolus versicolor, Ganoderma lucidum, Grifola frondose,
Phellinus linteus, Pleurotus eryngii, and Poria cocos) such as cancer [13], diabetes mellitus [14],
obesity [15,16], hyperlipidemia [10], cardiovascular and neurodegenerative diseases [17],
and hypercholesterolemia [2].
Whether edible mushrooms are considered a sustainable healthy human food source
mainly depends on the mushroom species and their chemical composition as well as the
growing media. The safety and health of a mushroom is the main factor that controls its
edibility. Many studies have confirmed that mushrooms grown in polluted soils threaten
human health [13,18–23]. These studies focused on pollutants (e.g., heavy metals, organic
toxicants, radioactive pollutants, or radionuclides) in mushrooms grown in polluted soils
as reported in China [20,23], Iran [18,21], Poland [22,24,25], Spain [26], and Turkey [27,28].
However, further investigations on the consumption of edible mushrooms cultivated in
polluted soils are needed before the association between pollutants and edible mushrooms
can be more clearly understood [19].
Therefore, this review focuses on edible mushrooms and the differences between them
and poisonous mushrooms. Common species of edible mushrooms will be discussed,
including their bioactive ingredients content, nutritional values, and health benefits. The
biomedical applications of edible mushrooms will also be reported.

2. Methodology of the Review

This review is an attempt to focus on the edible mushrooms and the difference be-
tween them and poisonous mushrooms. The methodology that was followed in this study
depended on the collected materials from the common scientific websites or engines such
as the databases of ScienceDirect, SpringerLink, PubMed, and MDPI. This is work based on
combinations of key terms to identify useful documents, including edible mushrooms, poi-
sonous mushrooms, nutritional value, and medicinal attributes. The collected information
included original articles, review articles, and chapters in the English language, and more
than 200 articles finally were selected. The collected information was analyzed and listed
in tables and drawn in figures. Tables summarize much of the information by surveying
the nutritional and medicinal attributes in collected articles on edible mushrooms, whereas
figures include photos of edible and poisonous mushrooms along with comparisons be-
tween them. The studies were collected from the main places for mushroom production
such as China, Iran, Poland, India, Brazil, Ireland, Taiwan, and Malaysia.

3. Sustainable Production of Edible Mushrooms

More than 100 countries cultivate edible mushrooms commercially using different
systems and on different scales. The global production of mushrooms increases annually
at growth rate of about 7%. The current global consumption of mushrooms is around
12.74 million tonnes, and it is predicted that the global production of mushrooms will
scale to 20.84 million tonnes by 2026 [29]. Mushrooms can be classified into some cate-
gories including (1) cultivated mushrooms (which can be grown commercially by farmers
using various strategies to produce for sale at supermarkets; these include enoki and
oyster mushrooms), (2) wild mushrooms (which can grow on the root systems of trees in
forests and are harvested by mushroom hunters—lots of wild mushrooms are poisonous),
(3) medicinal mushrooms (several mushrooms have medicinal benefits but might not al-
ways be pleasant to consume), and (4) poisonous mushrooms, which have toxic substances
or toxins [30]. Based on the growing method of mushrooms, they can be classified into
saprotrophic mushrooms (sapro = rot; troph = eating), which grow on dead matter; myc-
orrhizal mushrooms (mykes = fungus; rhiza = root), which have a symbiotic relationship
(sym = together; biosis = way of living) with crops or trees; and parasitic mushrooms,
which infect and depend on its host plant and then kill it [30].
Sustainability 2022, 14, 4941 3 of 30

Most cultivated mushrooms have a basic life cycle, which includes the following stages
(1) sporulation (i.e., production spores not seeds), (2) spore germination and the mating
of cells, (3) colonization to complete fruiting initiated, (4) the formation of primordia,
and (5) after fruiting or mature of mushrooms, spores release again and the cycle repeats
(Figure 1) [31]. The cultivation steps can be summarized in terms of seven stages, as
presented in Figure 2 and described in [31]. The sustainable production of mushrooms is
important because mushrooms are considered edible foods and are high in protein content.
There are many poisonous mushrooms, which must be clear to consumers in order to
avoid causing serious ecological and health problems [32]. The huge amount of wastes that
result from mushroom cultivation needs to be sustainably recycled for in order to protect
the environment and to produce the bioenergy as well. Therefore, edible mushrooms
can be promoted as an important agri-business activity to address many environmental
issues, especially the ecological degradation [29,33]. The cultivation of mushrooms can be
used to manage farm wastes by recycling them, as well as to dispose of spent mushroom
substrate “SMS” (i.e., wastes remaining after harvesting mushrooms). On the other hand,
the cultivation of mushrooms should protect from many competitor microorganisms and
promote hygiene, which are controlled through the environmental conditions as presented
in Table 1 [34].

Table 1. The environmental conditions controlling the most commonly cultivated species as ideal
values, but certain strains may exceed them.

Environmental Conditions Spawn Run Primordia Formation Fruitbody Development

Temperature 21–27 ◦C 10–21◦C 10–24 ◦ C
Relative Humidity 85–95% 95–100% 80–90%
Light 50–100 lux 1000–1500 lux 1000–1500 lux
CO2 level 5000–20,000 ppm; 1 FAE/h 500–2000 ppm; 4–8 FAE/h <1000–2000 ppm; 4–8 FAE/h
Duration 2–8 weeks 3–12 days 5–8 days
Source: [34] Fresh Air Exchange (FAE).
 room substrate “SMS” (i.e., wastes remaining after harvesting mush
hand, the cultivation of mushrooms should protect from many com
isms and promote hygiene, which are controlled through the environ
Sustainability 2022, 14, 4941 4 of 30
presented in Table 1 [34].

Figure
Figure 1. The industry
1. The of mushroom
industry cultivation is considered
of mushroom an important
cultivation industry, which
is considered anneeds
important
certain steps to produce the edible mushrooms as presented in these photos. The first group of photos
certain steps to produce the edible mushrooms as presented in these photos.
(1) includes mushroom fungi (Pleurotus ostreatus) culture, which should first be ready on the surface
tos
of (1)plate
agar includes
beside themushroom fungias(Pleurotus
cultivation substrates, ostreatus)
well as tools for culture,
propagation which
of mushroom and shoul
◦
poured media (heat treated at 95 C, for 1 day). The inoculant and the millet spawn in a jar should be
also prepared. The second and third group of photos (2,3) include the industrial preparing and filling
of spawn in the factory, whereas the group of photos in (4) represent oyster mushroom production.
These steps have been photographed from the factory of “Magyar Gomba Kertész Kft”, and all photos
were taken by Gréta Törős, Debrecen University, Hungary.
 surface of agar plate beside the cultivation substrates, as well as tools for propagation of mushroom
and poured media (heat treated at 95 °C, for 1 day). The inoculant and the millet spawn in a jar
should be also prepared. The second and third group of photos (2,3) include the industrial preparing
and filling of spawn in the factory, whereas the group of photos in (4) represent oyster mushroom
Sustainability 2022, 14, 4941 5 of 30
production. These steps have been photographed from the factory of “Magyar Gomba Kertész Kft”
and all photos were taken by Gréta Törős, Debrecen University, Hungary.

Figure2.2.Different
Figure Different stages
stages of mushroom
of mushroom cultivation
cultivation startingstarting from preparation
from preparation ofharvesting.
of media till media till harvest-
ing.
4. Edible vs. Poisonous Mushrooms
Table 1. The environmental conditions controlling the most commonly cultivated species as idea
There are many categories of mushrooms, such as edible, non-edible, and poisonous
values, but certain strains may exceed them.
mushrooms; wild and cultivated mushrooms; or medicinal and industrial mushrooms.
Environmental Conditions Mushroom by-products,
Spawn Run which mayPrimordia
be generated during production
Formation and processing,
Fruitbody rep-
Development
resent a disposal problem. However, these by-products may be promising sources for
Temperature 21–27 °C 10–21 °C 10–24 °C
several applications because of their nutritional and functional properties [35,36]. There
Relative Humidity may be many 85–95%
sustainable applications for95–100%
these by-products, including the80–90%
production
Light 50–100
of bioactive lux
compounds 1000–1500
for pharmaceutical lux
and nutraceutical 1000–1500
formulations, as well luxas
CO2 level 5000–20,000
several ppm; 1 FAE/h
other applications such as500–2000 ppm;fertilizers,
animal feeds, 4–8 FAE/h <1000–2000
energy production,ppm;4–8
bioreme-FAE/h
Duration diation, bio-based
2–8 weeksmaterials, cosmetics, and cosmeceuticals.
3–12 days Many applications of edible
5–8 days
mushrooms
Source: [34]are presented
Fresh in Figure
Air Exchange 3, according to Kumar et al. [37], Zhang et al. [38], and
(FAE).
Lopez-Hortas et al. [39].
4. Edible vs. Poisonous Mushrooms
There are many categories of mushrooms, such as edible, non-edible, and poisonous
mushrooms; wild and cultivated mushrooms; or medicinal and industrial mushrooms
Mushroom by-products, which may be generated during production and processing, rep-
 be many sustainable applications for these by-products, including the production of bio-
active compounds for pharmaceutical and nutraceutical formulations, as well as several
other applications such as animal feeds, fertilizers, energy production, bioremediation,
bio-based materials, cosmetics, and cosmeceuticals. Many applications of edible mush-
rooms
Sustainability 2022,are presented in Figure 3, according to Kumar et al. [37], Zhang et al. [38], and
14, 4941 6 of 30

Lopez-Hortas et al. [39].

Figure 3. Potential applications of different

Figure 3. Potential mushrooms
applications of differentand their by-products,
mushrooms whichwhich
and their by-products, include using
include using
bioactive compounds bioactive
in developing
compoundstheinpharmaceutical and nutraceutical
developing the pharmaceutical formulations,
and nutraceutical in in
formulations, addition
addition to
to applications such as animal feeds,
applications such as bioremediation, fertilizers,
animal feeds, bioremediation, energy
fertilizers, production,
energy production,bio-based ma-
bio-based materials,
cosmetics, and cosmeceuticals.
terials, cosmetics, and cosmeceuticals.
Some poisonous species of mushrooms have toxins that are not destroyed under the
Some poisonous species
high of mushrooms
temperatures reached duringhave toxins
cooking [40]. that arestudies
Several not destroyed under the
have been conducted on the
identification
high temperatures reached of mushrooms,
during cookingincluding their taxonomy,
[40]. Several studies andhavenaming
beenindices can be found
conducted on
on many websites such as Index Fungorum (http://www.indexfungorum.org/, accessed
the identification of mushrooms, including their taxonomy, and naming indices can be
on 15 January 2022), Myco-Bank Database (http://www.mycobank.org/, accessed on 15
found on many websites
Januarysuch
2022), as
theIndex FungorumSociety
British Mycological (http://www.indexfungorum.org/, ac-
(https://www.britmycolsoc.org.uk/library/
cessed on 15 January 2022), Myco-Bank
english-names, accessed onDatabase (http://www.mycobank.org/,
15 January 2022), accessed
International Mycological Association, (https:
on 15 January 2022),//www.ima-mycology.org/,
the British Mycologicalaccessed Societyon(https://www.britmycolsoc.org.uk/li-
15 January 2022), Fungal databases (https://nt.
ars-grin.gov/fungaldatabases/, accessed on 15 January 2022), NCBI or National Center
brary/english-names, accessed on 15 January 2022), International Mycological Associa-
for Biotechnology Information (https://www.ncbi.nlm.nih.gov/taxonomy, accessed on 15
tion, (https://www.ima-mycology.org/, accessed
January 2022), and the Global on 15Information
Biodiversity January 2022), Fungal databases
Facility (https://www.gbif.org/
(https://nt.ars-grin.gov/fungaldatabases/, accessed
species, accessed on 16 December 2021).on 15 January 2022), NCBI or National
FAO has stated that reports on the identification of edible or poisonous mushroom
Center for Biotechnology Information (https://www.ncbi.nlm.nih.gov/taxonomy, ac-
species are based on named sources, and there is no test or characteristic to distinguish
cessed on 15 January 2022),
edible from andmushrooms
poisonous the Global Biodiversity
[41]. The Information
edibility of mushrooms Facility
was simplified into
(https://www.gbif.org/species, accessedbased
the following categories on 16onDecember
the available2021).
information: (1) confirmed edible mush-
FAO has stated that reports on the identification of edible or poisonous mushroom
species are based on named sources, and there is no test or characteristic to distinguish
edible from poisonous mushrooms [41]. The edibility of mushrooms was simplified into
Sustainability 2022, 14, 4941 7 of 30

rooms (E1), (2) confirmed edible mushrooms with conditions (E2), (3) unconfirmed edible
mushrooms (E3), and poisonous mushrooms (P) [41]. It is worth noting that some edible
mushrooms are very similar in their appearance to the poisonous types and may grow in
the same habitat; an example is Entoloma sarcopum (an edible mushroom) and Entoloma
rhodopolium (a poisonous mushroom) [42] (Table 2; Figure 4). The edibility of mushrooms
must be accurately identified based on different advanced methods such as sensory identi-
fication methods (e.g., morphological and odor methods), instrumental analysis methods
(e.g., mass spectrometry, chromatographic, and spectral technology), and molecular biology
methods (e.g., isothermal amplification technology, molecular marker technology, endoge-
nous reference gene method, and sequencing technology). A review of this topic was
provided by Wei et al. [43]. Edible and non-edible or poisonous mushrooms have recently
been discussed in reviews such as Wu et al. [40], Li et al. [44], Ramírez-Terrazo et al. [45],
Zhang et al. [46], and Wei et al. [43].

Table 2. A list of some selected non-edible or poisonous mushroom species including the common
English name and their family.

Mushroom Species Name Common English Name Family

Agaricus pilatianus (Bohus) Bohus Basionym Agaricaceae
Agaricus phaeolepidotus F.H. Møller Agaric Agaricaceae
Agaricus praeclaresquamosus A.E. Freeman Ink-smelling mushroom Agaricaceae
Agaricus xanthoderma var. xanthoderma Carbolic mushroom Agaricaceae
Amanita gemmate (Fr.) Gillet Yellow agaric Amanitaceae
Amanita muscarina Fly agaric Amanitaceae
Amanita pantherine (DC ex Fr.) Secr. Panther Amanitaceae
Amanita phalloides Killer agaric Amanitaceae
Boletus calopus Fr. Farkino Boletaceae
Boletus radicans Pers. Rooting bolete Boletaceae
Boletus satanas Lenz Devil’s bolete Boletaceae
Clitocybe dealbata (Sowerby) P. Kumm Field or ivory funnel Tricholomataceae
Clitocybe cerussata (Fr.) P Kumm Wax white funnel Tricholomataceae
Cortinarius orellanus Fr. Fool’s webcap Cortinariaceae
Entoloma sinuatum Livid pinkgill Entolomataceae
Entoloma rhodopolium (Fr.) P. Kumm. Wood pinkgill Entolomataceae
Gyromitra fastigiata (Krombh.) Rehm Red-brown priest mushrooms Discinaceae
Gyromitra esculenta (Pers.) Fr. False Morel Discinaceae
Galerina marginata (Batsch) Kühner Funeral Bell Hymenogastraceae
Helvella crispa Scop. Curly paprika mushrooms Helvellaceae
Helvella leucopus Pers. Pale foot saddle Helvellaceae
Hygrocybe conica (Schaeff.) P. Kumm Blackening waxcap Hygrophoraceae
Hypholoma fasciculare (Huds.) P. Kumm Basionym Strophariaceae
Hebeloma synapizans (Paulet) Gillet Bitter poisonpie Hymenogastraceae
Inocybe asterospora Quelet Star spore susula Inocybaceae
Inocybe fastigiate (Schaeff.) Quél. Kerti susulyula Inocybaceae
Inocybe patouillardii Bres. Brick red susula Inocybaceae
Lepiota cristata (Bolton) P. Kumm Stinky dapperling Agaricaceae
Lepiota brunneoincarnata Meat brown venison Agaricaceae
Lepiota pseudohelveola Kühner ex Hora Roe deer mushrooms Agaricaceae
Mycena pura (Pers.) P. Kumm Lilac Bonnet Mycenaceae
Omphalotus olearius (DC.) Singer Illuminated funnel (Basionym) Omphalotaceae
Psilocybe semilanceata (Fr.) P. Kumm Magic mushroom Hymenogastraceae
Paxillus involutus (Fr.) Fr. Brown rollrim Paxillaceae
Tricholoma sulphureum (Bull.) P. Kumm Stinky Knight Tricholomataceae
Ramaria Formosa (Pers.) Quél Salmon coral Gomphaceae
Russula emetica (Schaeff.) Pers. Sickener Russulaceae
Scleroderma citrinum Pers. Common Earthball Sclerodermataceae
Sarcosphaera eximia var. nivea M.M. Moser Tulip cup mushrooms Pezizaceae
Tricholoma pardinum (Pers.) Quél. Panther (Leopard Knight) Tricholomataceae
Tricholoma saponaceum (Fr.) P. Kumm Soap scent Tricholomataceae
Note: Sources: http://xn--gombahatroz-r7a1v.hu/, accessed on 16 December 2021, [47,48]. https://www.
plantlife.org.uk/application/files/6714/8240/8340/recommended-english-names-for-fungi.pdf, accessed on
15 January 2022.
cosphaera eximia var. nivea M.M. Moser Tulip cup mushrooms Pezizaceae
holoma pardinum (Pers.) Quél. Panther (Leopard Knight) Tricholomataceae
holoma saponaceum (Fr.) P. Kumm Soap scent Tricholomataceae
Note: Sources: http://xn--gombahatroz-r7a1v.hu/, accessed on 16 December 2021, [47,48].
Sustainability 2022, 14, 4941https://www.plantlife.org.uk/application/files/6714/8240/8340/recommended-english-names-for-
8 of 30
fungi.pdf, accessed on 15 January 2022.

Amanita muscaria Agaricus phaeolepidotus

Galerina marginata Gyromitra fastigiata

Omphalotus olearius Hypholoma fasciculare

Russula emetica Tricholoma sulphureum

Figure 4.Figure
List of some
4. List poisonous
of some mushrooms
poisonous mushroomsincluding thescientific
including the scientific names
names for each
for each one. Sources:
one. Sources: by
by user user
Strobilomyces,
Strobilomyces,CCCCBY-SA
BY-SA3.0, https://commons.wikimedia.org/w/in-
3.0, https://commons.wikimedia.org/w/index.php?curid=1091222;
dex.php?curid=1091222;
by Onderwijsgek, CC byBY-SA
Onderwijsgek, CC BY-SA 3.0 nl, https://commons.wikimedia.org/w/in-
3.0 nl, https://commons.wikimedia.org/w/index.php?curid=1428517;
dex.php?curid=1428517; by Gerhard
by Gerhard Koller (Gerhard), Koller3.0,
CC BY-SA (Gerhard), CC BY-SA 3.0, https://commons.wiki-
https://commons.wikimedia.org/w/index.php?curid=
18730680; by Alan Rockefeller, CC BY-SA
media.org/w/index.php?curid=18730680; by4.0, https://commons.wikimedia.org/w/index.php?curid=
Alan Rockefeller, CC BY-SA 4.0, https://com-
95423433; by Alan Rockefeller, CC BY-SA 4.0, https://commons.wikimedia.org/w/index.php?curid=
mons.wikimedia.org/w/index.php?curid=95423433; by Alan Rockefeller, CC BY-SA 4.0,
61680344; by Renier van Rensburg, CC BY-SA 4.0, https://commons.wikimedia.org/w/index.php?
curid=71032684; by Strobilomyces, CC BY-SA 3.0, https://commons.wikimedia.org/w/index.php?
curid=438081; by MichalPL, CC BY-SA 4.0, https://commons.wikimedia.org/w/index.php?curid=
63737437, accessed on 16 January 2022.
Sustainability 2022, 14, 4941 9 of 30

Several taxa of poisonous mushrooms have been reported particularly among the most
common wild mushrooms, such as Amanita, Chlorophyllum, Cantharocybe, Inocybe, Entoloma,
Leccinellum, Russula, and Xerocomus [49–51]. Poisonous mushrooms have toxic compounds
that usually cause problems due to unintentional ingestion, with these problems including
endocrine toxicity, neurotoxicity, cytotoxicity, myotoxicity, and gastrointestinal distur-
bances (Table 3) [52]. Many poisonous species produce neurotoxic secondary metabolites
such as muscarine or psilocybin [51], alpha-amanitin and amanitin [53], coprine [50], gy-
romitrin [52], ibotenic acid [54], orellanine [55], phallotoxin [56], and russuphelins [57].
Differences between edible and poisonous mushrooms are of important concern for human
health, which mainly depends on the following criteria: the edible mushrooms should
have a pleasant odor and sweet taste and not be stained green or purple under cutting; the
should not burn or sting the tongue when a piece is tasted; and there should be no scales
on the cap [58]. A comparison between edible and poisonous mushrooms can be found in
Figure 5 [58].

Table 3. A list of main toxins, clinical features, and their relationships to poisonous mushrooms based
on the classification of poisonous mushrooms.

Toxins Poisonous Mushrooms Clinical Features

Group 1: Cytotoxic mushroom poisoning
Liver damage 1–3 days after ingestion due to hepatotoxic
α-amanitin Amanita phalloides, A. virosa, A. verna, A. exitialis
and nephrotoxic effects
Amanitin Amanita verna, A. virosa, A. phalloides, Lepiota
helveola, Galerina marginata Gastrointestinal disturbances; liver and renal failure
Orellanine Amanita smithiana, A. pseudoporphyria Kidney failure (early primary nephrotoxicity)
Cortinarius orellanus, C. rubellus Delayed primary nephrotoxicity
Group 2: Neurotoxic mushroom poisoning
Omphalotus, Clitocybe cerussata, C. dealbata, Respiratory failure, bradycardia, miosis, salivation,
Muscarine Inocybe fibrosa, I. hystrix, I. geophylla, I. rimosa, lacrimation, diarrhea and bronchospasm
I. sambucina perspiration, lachrymation
Psilocybe caerulescens, P. cubensis, P. semilanceata, Euphoria, hallucinations, tachycardia and blood pressure,
Psilocybin and psilocin Panaeolus cinctulus mydriasis, tremors, and fever
Amanita muscaria, A. pantherina, A. gemmate, Neurotoxicity, continuous periods of excitation and
Ibotenic acid A. ibotengutake, inhibition in the nervous system
Amanita muscaria, A. pantherina, A. gemmate,
Muscinol A. cothurnata, A. regalis Central nervous system depression and hallucinations
Group 3: Myotoxic mushroom poisoning
Bolesatine Rubroboletus satanas (Lenz), R. lupinus, Non-lethal; causes gastrointestinal irritation,
R. legaliae vomiting, nausea
Russuphelins Russula subnigricans Gastrointestinal disturbances, and renal failure
Saponaceolide Tricholoma equestre, T. terreum Delayed onset myotoxicity
Group 4: Metabolic, endocrine and related toxicity mushroom poisoning
Coprinopsis alopecia, C. atramentaria, Flushing, headache, dyspnea, sweating, arrhythmia,
Coprine
C. romagnesiana hypotension, and confusion
Gyromitra esculenta, G. californica, G. ambigua, Neurotoxicity, gastrointestinal upset, and destruction of
Gyromitrin
G. gigas, G. infula blood cells
Polyporic acid Hapalopilus rutilans Abdominal pain, nausea, vomiting, diarrhea
Trichothecenes Podostroma cornu-damae A multi-organ illness, often fatal
Group 5: Gastrointestinal irritant mushroom poisoning
Chlorophyllum molybdites, Entoloma sinuatum, Gastrointestinal effects without causing other
———
Tricholoma pardalotum significant effects
Group 6: Miscellaneous adverse reactions
Deadly can affect the vascular system and lead to loss of
Ergotamine Claviceps spp.
limbs and/or cardiac arrest
An erythromelalgia-like syndrome; reddening, swelling,
Acromelic acid Paralepistopsis amoenolens, Clitocybe acromelalga
and burning pain in the extremities
Pleurocybella porrigens, Grifola frondose, Delayed onset with cramps and coma; re-existing chronic
Hydro-Cyanic Acid Pleurotus eringii renal failure
Lentinan (Shiitake) Lentinola edodes, Auricularia auricular-judae Dermatitis, linear rash/whiplike lesions, photosensitivity
Phallotoxin Amanita phalloides Gastrointestinal upset
Abdominal pain, nausea, vomiting, and acute
Paxillus syndrome Paxillus involutus
respiratory failure
Sources: [50,51,53,55,57–60].
 Acid eringii chronic renal failure
Lentinan (Shii- Dermatitis, linear rash/whiplike lesions, photosensi-
Lentinola edodes, Auricularia auricular-judae
take) tivity
Phallotoxin Amanita phalloides Gastrointestinal upset
Paxillus syn- Abdominal pain, nausea, vomiting, and acute res-
Sustainability 2022, 14, 4941 Paxillus involutus 10 of 30
drome piratory failure
Sources: [50,51,53,55,57–60].

Figure5.5.AAcomparison
Figure comparisonbetween
betweenedible
edibleand
andpoisonous
poisonousmushrooms,
mushrooms,including
includingmorphological
morphologicaland
and
chemical approaches.
chemical approaches.

5.5.Common
CommonSpecies
Speciesof
ofEdible
EdibleMushrooms
Mushrooms
Edible
Ediblemushrooms
mushroomsare arethe
thefleshy
fleshyfruit
fruitbodies
bodiesofofmany
manyspecies
speciesofofmacro-fungi
macro-fungithatthat
can
canbe
beused
usedininthe
thehuman
humandietdiet(Table
(Table4).
4).The
Theedibility
edibilitycriteria
criteriaofofmushrooms
mushroomsmay maydepend
depend
on
onthe
theabsence
absenceof ofpoisonous
poisonoussubstances
substances or or toxins
toxins that
that are
aredetrimental
detrimental toto human
human health.
health.
Edible mushrooms are mainly consumed for their nutritional value, medicinal
Edible mushrooms are mainly consumed for their nutritional value, medicinal features, features,
and
andsweet
sweettaste.
taste.Several
Severalspecies
speciesof ofedible
ediblemushrooms
mushroomsare arefound
foundthethewild
wildand
andraised
raisedfor
for
harvest,
harvest, but some species are difficult to cultivate. Several articles or reviewsreported
but some species are difficult to cultivate. Several articles or reviews have have re-
about edible
ported aboutmushrooms from different
edible mushrooms points ofpoints
from different view: of view:
(1) Studies on edible mushrooms harvested from polluted areas with a focus on Africa [13],
China [20,23,61,62], Greece [63], Iran [18,21], Poland [22,24,25,64–66], Turkey [27,28,67–72],
and Spain [26] or on Europe as a whole, e.g., Świsłowski et al. [73], or on the global
level, e.g., Dowlati et al. [18].
(2) A study focusing on locals’ perspective concerning the changing of the abundance of
wild edible mushrooms, which has decreased due to direct exploitation by 31% and
land use change by 38% of all taxa [74].
(3) Studies on the bioactive properties of different edible mushrooms, including total
phenolic content, phenolic acid, antioxidant capacities, and antimicrobial activity
in Pleurotus sajor-caju wild edible mushroom [75], in Agaricus bisporus, Cantharellus
cibarius, Boletus edulis, Lactarius deliciosus, Lentinus edodes, Ganoderma lucidum, Hericium
erinaceus, Morchella spp., and Pleurotus ostreatus [76], and in Pleurotus ostreatus [77];
polysaccharide in many edible mushrooms [78]; and bioactive components in several
species of edible mushrooms [79].
(4) Studies on determining the amount of potentially bioavailable compounds such as
phenolic compounds in mycelia of Agaricus bisporus, Cantharellus cibarius, and Lentinula
edodes [80].
Sustainability 2022, 14, 4941 11 of 30

(5) Studies on integrated methods (morphological and molecular approaches) for the
identification of bioactive compounds [81], or using only molecular strategies for
identification [82].
(6) Studies on the mineral and nutritional composition of wild edible mushrooms [82–85]
and the nutritional value and biological properties of some edible mushrooms [86].
(7) Studies on using mushroom-derived bioactive compounds against SARS-CoV-2 infec-
tion [87,88].

Table 4. List of selected edible mushroom species as reported in the literature.

Mushroom Species Name Common English Name Family

Agaricus arvensis Schaeff. Horse mushroom Agaricaceae
Agaricus bitorquis (Quél.) Sacc. Pavement mushroom Agaricaceae
Agaricus bisporus (J.E. Lange) Imbach, white White cultivated mushroom Agaricaceae
Agaricus bisporus (J.E. Lange) Imbach, brown Brown cultivated mushroom Agaricaceae
Agaricus sylvaticus Schaeff. Blushing wood mushroom Agaricaceae
Agrocybe cylindracea (Brig.) Fayod Poplar mushroom Strophariaceae
Amanita ovoidea (Bull.) Link Bearded amanita Amanitaceae
Amanita vaginata (Bull.) Lam. Grisette Amanitaceae
Armillaria mellea (Vahl) P. Kumm. Honey fungus Physalacriaceae
Auricularia auricula-judae (Fr.) Quél Black wood ear Auriculariaceae
Cantharellus cibarius Fr. Chanterelle Cantharellaceae
Clavariadelphus pistillaris (L.) Donk Giant club Clavariadelphaceae
Clitocybe nebularis (Batsch) P. Kumm. Clouded funnel Tricholomataceae
Clitopilus prunulus (Scop.) P. Kumm. The Miller Entolomataceae
Cordyceps militaris (L.) Fr. Scarlet caterpillar club Cordycipitaceae
Coprinus comatus (O.F. Müll.) Gray Shaggy inkcap/lawyer’s wig Agaricaceae
Gyroporus castaneus (Bull.) Quél. Chestnut bolete Gyroporaceae
Helvella acetabulum (L.) Quél. Vinegar cup Helvellaceae
Hygrophorus marzuolus (Fr.) Bres. March mushroom Hygrophoraceae
Hygrophorus russula (Schaeff. ex Fr.) Kauffman Pinkmottle woodwax Hygrophoraceae
Lactarius pubescens Fr. Bearded milkcap Russulaceae
Lactarius repraesentaneus Britzelm. Yellow bearded milkcap Russulaceae
Lactarius rufus (Scop.) Fr. Rufous milkcap Russulaceae
Lactarius scrobiculatus (Scop.) Fr. Spotted milkcap Russulaceae
Lactarius sanguifluus (Poulet) Fr Bloody milkcap Russulaceae
Lactarius volemus (Fr.) Fr. Fishy milkca Russulaceae
Lentinus edodes (Berk.) Pegler Shiitake Marasmiaceae
Lentinus tigrinus (Bull.) Fr Tiger sawgill Hygrophoraceae
Lepista irina (Fr.) H. E. Bigelow Flowery blewit Tricholomataceae
Lepista nuda (Bull.) Cooke Wood blewit Tricholomataceae
Leucoagaricus leucothites (Vittad.) Wasser White dapperling Agaricaceae
Lycoperdon molle Pers. Soft puffball Lycoperdaceae
Lyophyllum decastes (Fr.) Singer Clustered domecap Tricholomataceae
Pleurotus citrinopileatus Singer Golden oyster mushroom Pleurotaceae
Pleurotus ostreatus (Jacq. ex Fr.) P. Kumm Oyster mushroom Pleurotaceae
Polyporus squamosus (Huds.) Fr. Dryad’s saddle Hygrophoraceae
Macrolepiota mastoidea (Fr.) Singer Slender parasol Agaricaceae
Neoboletus luridiformis (Rostk.) Basionym Boletaceae
Neolentinus lepideus (Fr.) Redhead & Ginns Scaly sawgill Polyporaceae
Otidea cochleata (L.) Fuckel Brown ear Pyronemataceae
Pluteus petasatus (Fr.) Gillet Scaly shield Pluteaceae
Pluteus salicinus (Pers.) P. Kumm Willow shield Pluteaceae
Russula albonigra (Krombh.) Fr. Menthol brittlegill Russulaceae
Russula delica Fr. Milk White brittlegill Russulaceae
Russula fellea (Fr.) Fr. Geranium brittlegill Russulaceae
Russula viscida Kudřna Viscid brittlegill Russulaceae
Strobilomyces strobilaceus (Scop.) Berk. Old man of the woods Boletaceae
Suillus bellinii (Inzenga) Kuntze Champagne bolete Suillaceae
Suillus bovinus (L.) Roussel Bovine bolete Suillaceae
Suillus luteus (L.) Roussel Slippery jack Suillaceae
Tricholoma focale (Fr.) Ricken Booted knight Tricholomataceae
Tricholoma terreum (Schaeff.: Fr.) Kumm Grey knight Tricholomataceae
Volvopluteus gloiocephalus (DC.) Basionym Pluteaceae
Volvariella volvacea (Bull.) Singer Straw mushroom Pluteaceae
Xerocomus subtomentosus (L.) Fr. Homotypic synonym Boletaceae
Selected references: [1,10,12,22,25,27,28,44,46,69,89–93], English name from [47,48].
Sustainability 2022, 14, 4941 12 of 30

6. Bioactive Ingredients of Edible Mushrooms

Edible mushrooms contain various bioactive ingredients such as proteins, polysac-
charides, polyunsaturated fatty acids (PUFA), dietary fibers, amino acids, vitamins, and
minerals (Table 5). They have essential health effects, such as antioxidant, antimicrobial,
immune-stimulatory, and anticancer, cholesterol-lowering properties (Figure 6) [12,37–39].
Several significant components and secondary metabolites dominate their biological activ-
ity. Lectins are carbohydrate-binding proteins that can be found in many types of edible
mushrooms such as Phaseolus vulgaris, Agaricus campestris, Agaricus bisporus, Grifola fron-
dosa, Boletus satanus, Flammulina velutipes, Tricholoma mongolicum, Ganoderma lucidum, and
Volvariella volvacea. Lectins have been shown to increase insulin secretion, activate the
immune system, and have anticancer effects [8]. Lectins can also play essential roles in
physiological processes such as dormancy, growth, morphogenesis, morphological changes,
and molecular recognition in the early stages of mycorrhization. Lectins isolated from the
Sustainability 2022, 14, x FOR PEER REVIEW
edible mushroom Clitocybe nebularis exhibit immunostimulatory effects on the most12potent
of 31
antigen-presenting cells, dendritic cells [94].

Figure 6.6. Edible

Figure Edible mushrooms
mushrooms are an important
important source
source of
of human
human food
food and
andhave
haveseveral
severalbioactive
bioactive
ingredients (phenols,
ingredients (phenols, lectins,
lectins,terpenoids,
terpenoids,flavonoids, etc.)
flavonoids, and and
etc.) bio-medicinal activities
bio-medicinal such as
activities anti-as
such
cancer, antidiabetic, antiaging, antiviral, antioxidant, and anti-inflammatory.
anticancer, antidiabetic, antiaging, antiviral, antioxidant, and anti-inflammatory.

Glucansare
Glucans areone
oneofofthe
the unique
unique ingredients
ingredients in mushrooms
in mushrooms thatthat
havehave immune-stimu-
immune-stimulatory,
latory, anticancer,
anticancer, anti-inflammatory,
anti-inflammatory, and antioxidant
and antioxidant effects.
effects. They canThey can be
be found found intypes
in different dif-
ferent
of edibletypes of edible
mushrooms such mushrooms
as Jelly earssuch as Jellyauricular),
(Auricularia ears (Auricularia auricular),lucidum),
Reishi (Ganoderma Reishi
(Ganoderma
Shiitake lucidum),
(Lentinula Shiitake
edodes), and(Lentinula edodes), and
Oyster (Pleurotus Oyster[95].
ostreatus) (Pleurotus ostreatus)
For example, [95]. For
beta-glucan,
example, beta-glucan, isolated from Pleurotus pulmonarius, has potent anti-inflammatory
isolated from Pleurotus pulmonarius, has potent anti-inflammatory and antinociceptive
and antinociceptive
properties properties
that inhibit that inhibitcytokines
pro-inflammatory pro-inflammatory cytokines
[96]. Glucans [96].
isolated Glucans
from iso-
Pleurotus
lated from Pleurotus pulmonarius also suppressed colon carcinogenesis associated with
colitis by regulating cell proliferation, inducing apoptosis, and suppressing inflammation
[97]. Beta-glucan is a high-molecular-weight polysaccharide of glucose bound by glyco-
sidic bonds.
Sustainability 2022, 14, 4941 13 of 30

pulmonarius also suppressed colon carcinogenesis associated with colitis by regulating

cell proliferation, inducing apoptosis, and suppressing inflammation [97]. Beta-glucan is a
high-molecular-weight polysaccharide of glucose bound by glycosidic bonds.

Table 5. A list of some phenolic compounds, polysaccharides, proteins, and triterpenoids found in
selected edible mushrooms species.

Edible Mushrooms Phenolics Polysaccharides Proteins Triterpenoids Refs.

Agaricus bisporus Gallocatechin Heteropoly-saccharide ABP Protein type FIIb-1 Ergosterol [98–101]
Boletus edulis Gallic acid Polysaccharides (BEBP-1) β-Trefoil lectin Boledulins A-C [98,102–104]
Cordyceps aegerita Proto-catechuic acid Fucogalactan Ageritin Bovistols A-C [105–108]
Coprinus comatus Flavones, and flavonols Modified polysaccharide Laccases Terpenoids [109–111]
Lactarius deliciosus Laccase Azulene-type
Syringic acid, vanillic acid Polysaccharide (LDG-M) sesquiterpene [112–115]
Pleurotus ostreatus Caffeic acid, and ferulic acid Mycelium polysaccharides Concanavalin A Ergosterol [116]
Pleurotus eryngii Cinnamic acid PEPE-A1 and PEPE-A2 Laccase Ergosterol [117,118]
Pleurotus cornucopiae Gallic acid β-glucan Oligopeptides Ergostane-type sterols [119–122]
Macrolepiota procera Proto-catechuic acid Polysaccharides β-Trefoil lectin Lanostane triterpenoids [103,123–125]
Russula virescens Quercetin SRVPs Laccase ———- [126–128]
Tuber melanosporum Flavonoids, phenols Exo-poly-saccharides (TP1) ————- Ergosterol [101,120–129]

Abbreviations: FIP (Immunomodulatory proteins); PEPE (Pleurotus eryngii purified polysaccharides); PSK (Polysac-
charide K); PSP (Polysaccharide peptide); RVP (Russula virescens polysaccharide); LDG m (Lactarius deliciosus
polysaccharides); BEPF (crude polysaccharides isolated from B. edulis).

Phenolic compounds are secondary metabolites of edible mushrooms. Polyphenols

have been extensively studied and shown to be effective against a variety of health compli-
cations. Phenolic acids such as p-hydroxybenzoic, cinnamic, gallic, salicylic, p-coumaric
acids, syringic acids, caffeic, ferulic, chlorogenic, and flavonoid can be found in mush-
rooms [130]. Gallic, caffeic, and p-coumaric acids are the main phenolic groups and play
an essential role in the biological activity of mushrooms [131]. Phenolic compounds have
high antioxidant activity. Polyphenols from edible mushrooms such as Meripilus giganteus,
Agaricus hydnum, and Rufescens silvaticus have high antioxidant capacity [80]. Phenolic
compounds have also shown anticancer activity against kidney cancer cell lines and human
ovarian cancer cell lines [6]. Flavonoid compounds, including myricetin, rutin, naringenin,
quercetin, morin, and hesperetin, are included in the polyphenol content, and they exhibit
antiproliferative effect [132].
Terpenoids such as monoterpenoids, sesquiterpenoids, diterpenoids, and triterpenoids,
are essential components in mushrooms. They have been shown to have antimicrobial, anti-
cancer, anticholinesterase, anti-inflammatory, antimalarial, and antioxidant properties [106].
Currently, about 285 types of terpenoids have been discovered in mushrooms and have
medicinal properties. For example, ganoderic acids are a lanastanoid type triterpenoid and
have been isolated from Ganoderma amboinense, Ganoderma lucidum, etc. Ergosterol, the prin-
cipal sterol in most edible mushrooms, is a valuable dietary precursor of vitamin D2 and a
natural antioxidant [133]. Phytosterols, such as ergosterols and ergosterol peroxide, have
been shown to be more potent than the nonsteroidal anti-inflammatory drug indomethacin,
as shown by their 50% inhibitory effect. High levels of ergosterols can be found in Agaricus
bisporus, Lentinus edodes, Grifola frondose, Pleurotus ostreatus, Agaricus bisporus, and Agaricus
bisporus [12].

7. Nutritional Values and Health Benefits of Edible Mushrooms

As a crucial source of food for humans for thousands of years, the medicinal and
organoleptic properties provided by the chemical composition and nutritional value of
edible mushrooms have been reported by several researchers (Figure 7). The consump-
tion of edible and medicinal mushrooms in Eastern and Western countries has gradually
increased in recent decades [134]. Agaricus bisporus, Lentinus edodes, and Pleurotus spp.
are presently the most common cultivated edible mushrooms, with China as the largest
producer of these mushrooms in the world [86]. Edible mushrooms are known for their
high contents of carbohydrates, protein, and crude fibers, as well as different bioactive
compounds, which provide both nutritional and health benefits for humans [86,135]. The
relative content of these nutritional components differs by species and between countries,
as reported in Table 6.
Sustainability 2022,
Sustainability 14,
2022, 14,4941
x FOR PEER REVIEW 14 of 31 14 of 30

Cantharellus cibarius Agaricus bisporus

Morchella esculenta Pleurotus ostreatus

Lactarius volemus Boletus edulis

Lentinula edodes Flammulina velutipes

Figure
Figure7.7.Photos of some
Photos common
of some edible mushrooms
common and their scientific
edible mushrooms and their names. Sources:
scientific by frank-
names. Sources: by
enstoen from Portland, CC BY 2.0, https://commons.wikimedia.org/w/index.php?curid=7304024;
frankenstoen from Portland, CC BY 2.0, https://commons.wikimedia.org/w/index.php?curid=7304
CC BY-SA 3.0, https://commons.wikimedia.org/w/index.php?curid=191739; by Dan Molter
024; CC BY-SACC
(shroomydan), 3.0,BY-SA
https://commons.wikimedia.org/w/index.php?curid=191739;
3.0, https://commons.wikimedia.org/w/index.php?curid=9822537; bybyDan Molter
(shroomydan), CC
TOMMES-WIKI, CC BY-SA
BY-SA3.0,
3.0,https://commons.wikimedia.org/w/index.php?curid=27753655;
https://commons.wikimedia.org/w/index.php?curid=9822537; by by
Strobilomyces, CC BY-SA 4.0, https://commons.wikimedia.org/w/index.php?curid=179148;
TOMMES-WIKI, CC BY-SA 3.0, https://commons.wikimedia.org/w/index.php?curid=27753655; by
Archenzo, CC BY-SA 3.0, https://commons.wikimedia.org/w/index.php?curid=24474; by voir ci-
by Strobilomyces, CC BY-SA 4.0, https://commons.wikimedia.org/w/index.php?curid=179148;
by Archenzo, CC BY-SA 3.0, https://commons.wikimedia.org/w/index.php?curid=24474; by voir
ci-dessous, CC BY 3.0, https://commons.wikimedia.org/w/index.php?curid=3330811; by Mыць
Дeниc, Public Domain, https://commons.wikimedia.org/w/index.php?curid=1334049, accessed on
16 January 2020.
Sustainability 2022, 14, 4941 15 of 30

The main nutritional compounds that edible mushrooms are rich in include proteins,
carbohydrates, crude fiber, total phenolic compounds, and vitamins and many minerals.
Several studies have discussed the nutritional value of edible mushrooms from different
locations such as Argentina [136], Bangladesh [85], China [136,137], Chile [86], India [83,138],
and Turkey [9,76]. Many reports have also focused on one or more edible mushrooms
such as Agaricus bisporus [24,139]; Lactarius deliciosus [140]; Pleurotus citrinopileatus [91];
Helvella leucopus and Morchella pulchella [9]; Pleurotus sajor-caju and Calocybe indica [141];
Agaricus bisporus, Cantharellus cibarius, and Lentinula edodes [80]; some studies have focused
on several species, e.g., Altaf et al. [83], Das et al. [135], and Taskın et al. [6]. Several
myco-chemical structures and compositions can be found in edible mushrooms, including
phenolic compounds, fatty acids, terpenoids, lipids, polysaccharides, and proteins, which
are important compounds for human health. Saini et al. [11] reported the biologically active
nutraceutical compounds present in edible mushrooms, including ergosterol, proteins, and
fatty acids content (388 mg/100 g DW, 4.5–37.4%, and 1.75–15.5%, respectively).

Table 6. General nutritional values (based on dry weight) of some common edible mushrooms from
different sources.

Mushroom Species Used in the Total Protein Total Total Phenols

Moisture (%) Crude Fiber (%) Ash (%) Refs.
Study (%) Carbo-Hydrate (%) (%)
I. Studied many species
Agaricus bisporus, Agrocybe
cylindracea, Boletus loyo, Cortinarius 86–96 8.56–23.88 62.97–83.65 7–15 5–13 0.75–4.72 [86]
lebre, Cyttaria espinosae
Apioperdon pyriforme, Helvella
elastica, Morchella conica and - 11.5–24.5 - 2.6–4.8 9.5–14.7 5.0–12.3 [83]
Rhizopogon luteolus)
Auricularia auricula-judae, A.
polytricha, Lactifluus piperatus, 49–88 19–56 7–18 5–11 3–8 7.3–10.2 [138]
Laetiporus sulphureus
Auricularia auricular, Ganoderma
lucidum, Pleurotus citrinopleatus, P. 4–11 20–45 11–61 5–40 6–10 1–8
djamor, P. eryngii, P. ostreatus, and P. [85]
ostreatus
Agaricus bisporus, Pleurotus ostreatus, 88–92 1.7–2.11 3–9 20–37 0.8–1.15 1.1–1.5
Lentinula edodes [136]
Tricholoma, Shiitake mushroom,
Pleurotus eryngii, Dictyophora 6.9–15.5 8.5–36.9 0.5–37.3 14.4–70.2 1.3–10.1 - [137]
indusiata
Agaricus bisporus, Boletus edulis,
Cantharellus cibarius, Lactarius 82.6–91 1.5–3.6 3.2–8.3 - - - [5]
deliciosus, Leccinum rufom
Astraeus odoratus, Craterellus aureus,
Lentinus edodes, Phaeogyroporus - 13.1–32.8 2.79–44.3 77.1 - - [142]
portentosus
Agaricus bisporus, Boletus edulis,
Calocybe indica, C. gambosa, Grifola - 18.1–62.8 31.1–70.6 7.81–32.3 3.5–19.7 - [135]
frondosa, Flammulina velutipes
Pleurotus sajor-caju; Calocybe indica 87–89 1.74–3.4 3.37–3.33 - 1.2–1.3 - [141]
II. Studied only one species
Agaricus bisporus 81.79 29.29 20.57 24.56 7.12 - [143]
Flammulina velutipes - 18.42 56.37 7.81 6.33 - [144]
Grifola frondosa 83.06 21.1 58.8 10.1 7.0 - [145]
Hericium erinaceus 95.69 23.3 57.0 7.8 9.4 - [145]
Lactarius deliciosus 92 17.19 66.61 31.81 8.62 4.5–13.6 [140]
Lyophyllum decastes - 18.31 34.36 29.02 14.20 - [146]
Pleurotus florida 87.05 34.56 31.59 11.41 7.40 - [147]
Pleurotus ostreatus 85.55 30.92 31.40 12.10 7.05 - [147]
Russula delica - 26.25 34.88 15.42 17.92 - [146]
Tremella fuciformis 91.73 4.6 94.8 1.4 0.4 - [148]
Tricholoma giganteum - 16.1 70.1 4.5 5.0 - [149]
Volvariella volvacea - 30.1 50.9 11.9 12.6 - [149]

The biofortification approach can be defined as a method by which pollutants and/or

biodegradation from the environment can be removed or blocked from microorganisms
such as bacteria, fungi, or algae. It is considered a sustainable agricultural strategy and a
cost-effective tool to increase the bioavailability or content of essential nutrients in the edible
parts of cultivated plants and reduce malnutrition [150]. Some edible mushrooms have
been biofortified with nutrients, particularly selenium [151], which is applied to produce
Se enrichment and increase bioactive metabolites of Pleurotus ostreatus [152], Cordyceps
militaris [153], Hericium erinaceus [154], and Ganoderma lucidum [155]. Nano-biofortification
is also an important approach that recently has been confirmed for using nano-selenium as
anti-COVID-19 nanoparticles [156,157].
Sustainability 2022, 14, 4941 16 of 30

8. Biomedical Applications of Edible Mushrooms

Medicinal mushrooms can be defined as mushrooms that are used to modulate human
immune responses such as antimicrobial, anticancer, and antioxidant (Table 7). These
mushrooms have important compounds such as alkaloids, polyphenols, polysaccharides,
sesquiterpenes, terpenoids, and metal chelating agents for treatment different human
diseases [17]. Medicinal mushrooms are gaining more recognition. Consequently, this
is creating new paths for researchers to understand how these types of mushrooms can
play a role against human diseases such as neurodegenerative diseases (i.e., Alzheimer’s
disease, Huntington’s disease, multiple sclerosis, Parkinson’s disease, motor neuron dis-
ease, and prion disease) through mechanisms such as regulating neurotrophins synthesis,
reducing oxidative stress, neuroinflammation, protecting neurons, and modulation of
acetylcholinesterase activity [17,158]. These strategies may include roles of mushrooms
such as (1) antibacterial antibiotics, (2) antimycotics, (3) biofilm inhibitors, (4) anticancer
agents, (5) antidiabetes, (6) improving nerve functioning, (7) cardiovascular disease control,
(8) antiviral agents, (9) prebiotic, and (10) immunosuppressive and immunomodulatory
agents [158,159].

Table 7. A survey of some pharmacological or medicinal activities in selected edible mushrooms

species according to published studies, study model, and the suggested disease being treated.

Mushroom Species Study Model Pharmacological or Medicinal Activities (Disease) Place Refs.
Tumoricidal, anticarcinogenic activity and antimetastatic
Agaricus blazei Animal/cell line Brazil [160]
activity (cancer)
Antioxidation, immune regulation,
Russula griseocarnosa Not available China [161]
hypoglycemic/hypolipidemic activities (diabetes)
Macrolepiota procera, Anticancer, anti-inflammatory, and reducing cardiovascular
Gymnopus dryophilus Not available Poland [12]
diseases (cancer)
Lentinus edodes and Hypsizygus
Cell line Immunomodulatory activity (inflammatory lung conditions) Ireland [96]
tessellatus
Sanghuangprous vaninii Cell line Anticancer particularly gynecological cancers (cervical cancer) China [162]
Russula virescens In vitro Anticancer, hypoglycemia and immune activities (cancer) China [163]
Ganoderma lucidum, G. artum and Anti-inflammatory, antitumor, antioxidation,
Cell line China [164,165]
G. tsugae anti-hyperglycemic or hyperlipidemic effects (diabetes)
Sanghuangporus sanghuang Antioxidant activities and anti-coronavirus
Cell line China [166]
(SARS-CoV-2) activity
Arthritis, bronchitis, hypertension, gastric ulcer, and nephritis
Ganoderma lucidum Not available China/Iran [167]
(tumorigenic diseases)
Agaricus spp., Pleurotus spp.,
Cell line Antitumor and anticancer agents (cancer) India [168]
Lentinus spp.
Ganoderma lingzhi, G. applanatum, Simulated gastrointestinal
For immunological diseases (gastrointestinal diseases) Hungary [169]
G. lucidum, and G. tsugae condition
Pleurotus citrinopileatus, P. djamor, Anti-atherosclerotic properties (high lovastatin content)
In vitro Poland [170]
P. eryngii, and P. ostreatus (cardiovascular diseases and cancers)
Armillaria mellea, Antrodia
camphorata, Hericium erinaceus, In vitro/in vivo Anti-neuroinflammatory activities (Alzheimer’s disease) Malaysia [171]
Ganoderma spp.
Calvatia gigantea, Fomes fomentarius, Anti -bacterial, antitumor, antioxidative and European
Phallus impudicus Cell culture anti-inflammatory (cancer) Union [172]
Agaricus bisporus, Ganoderma Mauritius
frondosa, Lentinula edodes Cell line Anticancer (cancer) [173]
Antioxidant, anti-inflammatory, antitumor, and
Antrodia salmonea Cell line Taiwan [174]
antiatherosclerosis activity (atherosclerosis)
Lignosus rhinocerus, L. tigris, Anticancer, anti-inflammatory, and antioxidative
L. cameronensis Cell line Malaysia [175]
activity (cancer)
Species of Agaricus, Anticancer, alleviation of osteoarthritis (cancer and Cameroon/
Animal [176]
Ganoderma, Pleurotus osteoarthritis) Nigeria
Antioxidant, antibacterial activity, antitumor, and
Ganoderma applanatum, Pleurotus Clinical trials on human and
eryngii, Lentinula edodes antiproliferative activity (Alzheimer’s disease, and human Italy [177]
animals
colon cancer)

Several studies have been published about different therapeutic applications of medic-
inal mushrooms [178]. These studies depend on the kind of bioactive compounds and
mushroom species, which can be divided into the following groups:
(1) Anticancer group
This group includes many kinds of cancers such as colorectal, oral, renal, and cervical
cancers besides leukemia, as well as breast, liver, lung, and prostate cancer. The main
bioactive mushroom extracts that can be used against cancers may include methanol,
polysaccharides, ganoderic acid, and ethanol extracts. Medicinal mushrooms that have
these bioactive compounds include Agaricus brasiliensis, Boletus edulis, Ganoderma lucidum,
Sustainability 2022, 14, 4941 17 of 30

Pleurotus ostreatus, P. pulmonarius, Ophiocordyceps sobolifera, Sanghuangprous vaninii, and

Tremella mesenterica [93,158,162,163,179,180].
(2) Antidiabetic group
This includes many mushroom species such as Grifola frondose and Phellinus linteus,
which have different polysaccharides as bioactive compounds [14,158,181,182].
(3) Cardiovascular diseases group
This includes species such as Agaricus bisporus and Tricholoma matsutakei. The crude ex-
tract of peptides found in these mushrooms may be useful in treating these diseases [158,183].
(4) Immune-function group
This includes many species such as Pleurotus ostreatus and Trametes pubescens. The
lectine extracts from these mushrooms may boost immune function [15,158,159,163].
(5) Rheumatoid arthritis group
This includes some mushroom species such as Grifola frondose and Psilocybin spp.
The active metabolite psilocin or polysaccharides extracts may help with rheumatoid
arthritis [15,184].
(6) Antiviral group
Major human viral diseases include human immunodeficiency virus (HIV), herpes
simplex virus (HSV), hepatitis B and C viruses (HBV, HCV), and influenza. There are many
antiviral compounds that can be extracted from various mushroom species [10,158,159,185].
Medicinal mushrooms can be divided into following groups based on three items mush-
room species, extract of bioactive compounds, and virus disease group [186].
(i) Antiviral mushrooms against HIV
This group includes mushrooms that have bioactive compounds that are active against
human immunodeficiency virus (HIV), which include Lentinus edodes (polycarboxylated
water-solubilized lignin), Pleurotus ostreatus (ubiquitin-like protein), Ganoderma lucidum
(several triterpenoids), and Ganoderma colossus (lanostane triterpenes).
(ii) Antiviral mushrooms against HSV
This group includes mushrooms that have bioactive compounds that act against
Herpes simplex virus (HSV), which includes Agaricus brasiliensis (sulfated polysaccharide
HSV-1), and Ganoderma lucidum (GLPG or Ganoderma lucidum proteoglycan).
(iii) Antiviral mushrooms against influenza
This group includes mushrooms that have bioactive compounds that act against in-
fluenza virus (H5N1), which includes Phellinus baumii (hispidin), Inonotus hispidus (phenolic
extracts), and Phellinus igniarius (sesquiterpenoid).
(iv) Antiviral mushrooms against hepatitis virus
This group includes mushrooms that have bioactive compounds that act against
hepatitis virus B or C (HBV, HCV), which include Antrodia camphorate (polysaccharides for
HBV), Ganoderma lucidum (ganoderic acid), Lentinula edodes (mycelia solid culture extract for
HCV), and Pleurotus ostreatus (lectin HBV). Recently, some published studies have reported
about the biological role of bioactive compounds from mushrooms against COVID-19 such
as polysaccharides [87,187], colossolactones, and ergosterol [88].
(v) Antiviral mushrooms against other viruses
This group includes mushrooms that have bioactive compounds that act against other
viruses, which include Hypsizygus marmoreus (sterols for Epstein–Barr virus), Auricularia
auricular (polysaccharides for Newcastle disease virus), and Agaricus blazei (Murill extract
for foot-and-mouth disease virus).
(7) Group of neurodegenerative diseases
Sustainability 2022, 14, 4941 18 of 30

Some medicinal mushrooms such as Grifola frondosa, Hericium erinaceus, Lignosus

rhinocerotis, Paxillus panuoides, Pleurotus giganteus, and Sarcodon scabrosus can improve
human cognitive functions. Many bioactive ingredients can be used from this group of
mushrooms against many diseases such as hericenones, scabronines, ganoderic acids, and
aqueous and ethanolic extracts [17,188].
(8) DNA damage group
This group includes mushrooms such as Agaricus blazei and Ganoderma lucidum, which
have extracts such as polysaccharides that may work against DNA damage [189].
(9) Antiaging group
This includes Tricholoma lobayense and Ganoderma lucidum, which could supply
polysaccharide TLH-3 Ergosterol ganodermasidase, which acts against aging [10,15].
(10) Antiobesity group
Many bioactive compounds can be extracted from mushroom species and used
against obesity, including Pleurotus eryngii, P. sajor-caju, and Ophiocordyceps sinensis. These
compounds may include β-glucan water extract and Cordyceps sinensis polysaccha-
ride [16,142,190,191].

9. Bioactive Compounds from Edible Mushrooms and Toxic Dose

In general, the link between medicinal edible mushrooms and the impacts of their
bioactive compounds in preventing human diseases (mycotherapy) is still in need of
more investigation. Thus, a growing concern in mycotherapy is required from the scientific
community to expand clinical trials to explore the potential benefits of the safe consumption
of these mushrooms [192]. The bioactive compounds found in mushrooms may have
important effects on human health. Mushrooms are most often taken into the body through
ingestion, but their mechanisms need to be studied in both in vivo and in vitro clinical
studies in order for their actions to be fully understood. Research should also be undertaken
to standardize all production stages of edible mushrooms from cultivation through the
extraction process, commercial preparation, regulation, and precise monitoring for high
quality level [192], as set by the European Food Safety Authority (EFSA). This issue has
been discussed by many studies, as reported by [74,79].
The bioactive compounds and their efficiency depend on several factors, from culti-
vation to the handling or processing of these bioactives to protect them from any harsh
environments and to improve their bio-availability and biological activities upon human
digestion [193]. The characterization of these bioactive components also depends on the
kind of methodology and instrumentation used during their measurement, such as bio-
chemical and high-performance liquid chromatography (HPLC) [194] or UHPLC-Mass
Spectrometry-(MS)/MS [12], as well as all processing and extraction tools such as the dry-
ing temperature during processing [195] and hot water during extraction [196]. Therefore, it
is difficult to establish the standardized doses for each edible mushroom for each bioactive
compound required to achieve its desired biological effect.
It is assumed that up to 3.8 million fungal species inhabit the Earth. Around 120,000 of
those are currently known, and more than 27,000 are macromycetes. Only about 500 species
of macromycetes are known to be poisonous for humans [197]. As mentioned above, sev-
eral studies have confirmed the beneficial attributes of consuming edible mushrooms. On
the other hand, several potential risks to human health may occur when edible mushrooms
grow in polluted soils and are ingested [13,198]. These risks mainly depend on the kind of
pollutants, their consumed concentration, and the mushroom species [64,199]. The levels of
pollutants, especially toxic elements in mushrooms before marketing, should be continually
evaluated to assess the human health risk [21]. Poisonous mushrooms are considered a
significant cause of “emergency medicine” when they cause mycotoxicity [198]. Poisonous
mushrooms contain a variety of various toxins such as satratoxins [200], muscarine, aman-
itin, gyromitrin, ibotenic acid, orellanine, and russuphelins, which can differ markedly in
Sustainability 2022, 14, 4941 19 of 30

their toxicity (Table 2). It is not easy to demarcate between edible and poisonous mushroom
species because there are several cultivated and edible wild species that have dual potential
health benefits and toxic compounds [198]. Some edible species may cause cardiac toxicity,
allergic reactions, rhabdomyolysis, and mutagenic effects, as observed in laboratory rodents
or in assays on cell cultures [198].
The mechanisms of interactions of nutritional or toxic compounds with the human
body is of crucial concern. These mechanisms may include their toxic or harmful effects
on the human endocrine, metabolic, and nervous systems. The recent increase in global
consumption of edible mushrooms, with fresh mushroom of up to 300 g per day, may
represent a public health concern [201]. Many studies using tumor-bearing animals, cul-
tured cancer cells, and clinical trials are required to investigate these mechanisms [202].
Several poisoning cases resulting from deadly toxic mushrooms such as Trichoderma cornu-
damae have been reported (Table 3), but toxicological analyses of these poisonings are still
rare [200]. Cytotoxic mushroom poisoning may result from toxins of α-amanitin, aman-
itin, and orellanine, and their mechanisms may include (1) liver damage 1–3 days after
ingestion due to hepatotoxic and nephrotoxic effects (such as mushrooms of Amanita phal-
loides, A. virosa, A. verna, A. exitialis); (2) gastrointestinal disturbances and liver and renal
failure (such as Amanita verna, Lepiota helveola, and Galerina marginata); (3) kidney failure or
early primary nephrotoxicity (Amanita smithiana, and A. pseudoporphyria); and (4) delayed
primary nephrotoxicity (Cortinarius orellanus, and C. rubellus). More mechanisms and the
mushrooms involved are listed in Table 2. The most distinguished poisonous mushrooms
in Tables 1 and 2 may include some species of Agaricus, Amanita, Gyromitra, Tricholoma, and
Russula. In this context, a comprehensive review of the activity of bioactivities in Pleurotus
spp. and their mechanism of action was reported by Cateni et al. [10].

10. Antinutritional Properties of Mushrooms

Apart from the many advantages resulting from the consumption of plant foods,
some plants (e.g., legumes) and mushrooms possess potential deleterious effects due to
their content of toxins or antinutrients [203–205]. Antinutrients are defined as natural
or synthetic compounds, which can interfere with the uptake/absorption of nutrients
by humans, and limit the bioavailability of essential nutrients causing a serious harmful
impact on human health [203,205]. Antinutrients may include chitin, glucosinolates, lectins,
phytates, oxalates, saponins, or tannins, which plants need in order to protect themselves
from the surrounding stressful conditions. On the other hand, these antinutrients have
become well-known to possess many beneficial effects and therapeutic potential on many
human diseases [206].
Although several benefits of edible mushroom have been reported in different fields,
as discussed earlier, including nano-biotechnology [33,207] and the production of bioactive
compounds [79], some reports discussed how mushroom products can be hard to digest
due to their content of antinutrients such as chitin (e.g., [203,208]). Some other reports
have confirmed that edible mushroom consumption may be associated with a lower risk
of mortality [209]. In general, edible mushrooms contain about 1.87–7.25% chitin, and
many published studies involve the chitin in mushrooms from different points of view
such as the production of chitinase [210], wound treatment [211], isolation of chitin–glucan
complex [212,213], production of chitin/cellulose nanofiber [214], and production of chitin
nanopaper [215].

11. General Discussion

The main intent of this review was to summarize the information related to edible
mushrooms with a focus on their nutritional and medicinal applications. Mushroom by-
products, which may be generated during their production and processing, may represent
a disposal problem. However, these by-products may be promising sources for several
applications because of their nutritional and functional properties [35,36]. There may be
many sustainable applications for these by-products, including the production of bioac-
Sustainability 2022, 14, 4941 20 of 30

tive compounds for pharmaceutical and nutraceutical formulations, as well as several

other applications such as animal feeds, fertilizers, energy production, bioremediation,
bio-based materials, cosmetics, and cosmeceuticals [33,91,216]. The sustainable produc-
tivity of edible/medicinal mushrooms has been reported by many researchers such as
Shirur et al. [29], which could be used in industrial applications (Kumar et al. [37]), for
the production of healthy food products (Zhang et al. [38]), for healthy and more sustain-
able foods (Das et al. [135], and for promising strategies of sustainable and novel food
production (Lopez-Hortas et al. [39]).
On the other hand, toxic or poisonous mushrooms are considered an important issue,
which still needs more exploration in terms of their toxins. Despite the health problems
associated with poisonous mushrooms, the use of their main bioactive substances in
such mushrooms in clinical applications is possible, as reported by many researchers
(e.g., Patocka et al. [217]). These toxins could be applied in many respects, including
biotechnology, biological control, and anticancer medicine. There is a need also for the
discovery of more toxic species of mushrooms, and their toxins, which are expected to be
increasingly known based on toxicological analyses.

12. Conclusions
Mushrooms are seen as an important source of food and are thought to play a crucial
role in a healthy diet. The nutritional values of edible mushrooms results from their low
content of calories, fat, and sodium and their high content of dietary fiber, proteins, vita-
mins, and essential nutrients, such as Cu, Mn, Se, and Zn. Several secondary metabolites
(i.e., bioactive compounds) can be extracted from many edible medicinal mushrooms such
as β-glucan, calcaelin, ergosterol, ganoderic acid, flavonoids, hispolon, laccase, lectins,
lentinan, phenolics, proteoglycan, nucleotides, nucleosides, and triterpenoids. These com-
pounds could offer various health benefits such as those provided by antioxidant, antiaging,
anticancer, antimicrobial, anti-inflammatory, antiobesity, and immunomodulatory agents.
These biological activities can reduce heart disease and hypertension, regulate metabolic
disorders and body functions, improve gut health, boost the immune system, and help
in weight loss. Edible mushrooms have become more popular as health promoters, with
great concern given to different research activities looking into bioactive ingredients in
several mushroom species. These bioactive compounds are needed to fortify or supply
various staple food products. Although these bioactive components of edible mushrooms
are crucial for human health, there is also an urgent need for testing such compounds
against recent diseases such as coronavirus disease (COVID-19). Further investigations are
needed on the nano-biofortification of different mushrooms, their bioactive compounds and
therapeutic properties, and the health benefits of edible mushrooms for a healthy lifestyle.

Author Contributions: J.P. and H.E.-R. developed the idea and outline of the review; K.B. wrote
the section of bioactive ingredients; N.A. wrote the nutritional section; Y.E. wrote the biomedical
applications of the manuscript; the section on common species was written by X.L., G.T. and P.H.;
and the other sections were written by H.E.-R., K.B., X.L. and N.A.; and Y.E. thoroughly revised
the manuscript and finalized it. All authors have read and agreed to the published version of
the manuscript.
Sustainability 2022, 14, 4941 21 of 30

Funding: The authors thank the support of the 2020-1.1.2-PIACI-KFI-2020-00100 Project “Develop-
ment of innovative food raw materials based on Maillard reaction by functional transformation of
traditional and exotic mushrooms for food and medicinal purposes”. The research was also sup-
ported by the Stipendium Hungaricum Scholarship Program (SH ID:140993). The work/publication
is supported by the EFOP-3.6.3-VEKOP-16-2017-00008 project. The project is co-financed by the
European Union and the European Social Fund.
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Acknowledgments: All authors thank Eric C. Brevik (College of Agricultural, Life, and Physical
Sciences, Southern Illinois University, Carbondale, IL, USA) for his revising and editing the MS.
Conflicts of Interest: The authors declare no conflict of interest.

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