Edible Mushrooms: Cultivation, Bioactive

Molecules, and Health Benefits

Sachin Gupta, Baby Summuna, Moni Gupta, and

Sudheer K. Annepu

Contents
1 Mushroom: Structure, Growth, and Composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 Global Trends of Mushroom Cultivation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 History and Status of Mushroom Cultivation in India . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3.1 Relative Contribution of Different Mushroom Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
4 Marketing of Mushrooms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
5 Types of Cultivated Mushrooms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
5.1 Mushrooms Under Cultivation in India . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
6 Nutrition in Mushrooms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
6.1 Proteins and Amino Acids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
6.2 Vitamins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
6.3 Carbohydrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
6.4 Fatty Acids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
7 Therapeutic Potential of Mushrooms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
7.1 Mushrooms as Antioxidants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
7.2 Mushrooms as Hypocholesterolemic Agents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

S. Gupta (*)
Division of Plant Pathology, Faculty of Agriculture, SKUAST-Jammu,
Jammu, Jammu and Kashmir, India
e-mail: sachinmoni@gmail.com
B. Summuna
Division of Plant Pathology, Faculty of Agriculture, SKUAST-Kashmir,
Jammu, Jammu and Kashmir, India
e-mail: summunabhat@gmail.com
M. Gupta (*)
Division of Biochemistry, Faculty of Basic Sciences, SKUAST-Jammu,
Jammu, Jammu and Kashmir, India
e-mail: moniguptaskuast@gmail.com
S. K. Annepu
Divison of Crop Improvement, ICAR-Directorate of Mushroom Research, Solan, Himachal
Pradesh, India
e-mail: sudheerannepu@gmail.com

# Springer International Publishing AG, part of Springer Nature 2018 1

J.-M. Mérillon, K. G. Ramawat (eds.), Bioactive Molecules in Food, Reference Series in
Phytochemistry, https://doi.org/10.1007/978-3-319-54528-8_86-1
2 S. Gupta et al.

7.3 Mushrooms as Hypoglycemic Agents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

7.4 Mushrooms as Antitumor Agents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
7.5 Mushrooms as Immunomodulators . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
7.6 Mushrooms as Antimicrobial Agents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
7.7 Mushrooms as Antiviral Agents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
7.8 Mushrooms as Antiallergic Agents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
7.9 Mushrooms as Anti-inflammatory Agents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
7.10 Mushrooms as Hepatoprotective Agents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
8 Bioactive Compounds in Mushroom . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
8.1 Peptides and Proteins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
8.2 Phenolic Compounds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
8.3 Polysaccharides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
8.4 Glucans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
8.5 Polysaccharide: Protein Complexes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
8.6 Terpenes and Triterpenoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
9 Mushroom-Based Biotech Products . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
10 Role of Mushroom Culture Collections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
11 Future Trends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

Abstract
Mushrooms are globally appreciated for their nutritional value and medicinal
properties. Their cultivation is an effective bioconversion technology of trans-
forming wastes and woods into potentially valuable resources and could also be
an important part of sustainable agriculture and forestry. Although India has the
advantage of favorable agroclimate, abundance of agrowastes, relatively low-cost
labor, and a rich fungal biodiversity, it has witnessed a lukewarm response in
growth of mushroom cultivation. Out of the total mushroom produced in India,
white button mushroom share is 73% followed by oyster mushroom (16%),
paddy straw mushroom (7%), and milky mushroom (3%). The per capita con-
sumption of mushrooms in India is also very meager and is even less than 100 g
per year. Besides low fat and high protein and vitamin contents, mushrooms are
rich sources of several minerals and trace elements, as well as dietary fibers. The
reported medicinal effects of mushrooms include anti-inflammatory effects, with
anti-inflammatory compounds of mushrooms comprising a highly diversified
group in terms of their chemical structure. They include polysaccharides, terpe-
noids, phenolic compounds, and many other low molecular weight molecules.
Of late, mushrooms have emerged as wonderful source of nutraceuticals, antiox-
idants, anticancer, prebiotic, immunomodulating, anti-inflammatory, cardiovas-
cular, antimicrobial, and antidiabetic. Owing to the synergistic action of present
bioactive molecules, majority of mushroom products possess beneficial health
effects and can be used on a regular basis without harm. Therefore, they are
considered as perspective organisms to develop different healthcare biotech
product. Mushrooms could potentially be very important in future food supplies
and in new dimensions of sustainable agriculture and forestry. In this chapter, an
attempt has been made to provide an insight into the various aspects of cultivation
of mushroom cultivation in India, nutritional benefits, therapeutic potential, and
bioactive components present in edible mushrooms.
Edible Mushrooms: Cultivation, Bioactive Molecules, and Health Benefits 3

Keywords
Mushrooms · Cultivation · Nutrition · Bioactive molecules

1 Mushroom: Structure, Growth, and Composition

In broad sense, “mushroom is a macro fungus with a distinctive fruiting body, which
can be either epigeous or hypogeous and large enough to be seen with naked eye and
to be picked by hand” [1]. It is perhaps the most well-known and documented edible
forest product [2]. Mushrooms have been widely used as foods [3, 4] and very often
as delicious and nutritious foods [5]. Approximately 14,000 described species of the
1.5 million fungi estimated in the world produce fruiting bodies that are large enough
to be considered as mushrooms [6]. Mushrooms belong to basidiomycetes and
ascomycetes with a cell cycle including the formation of sexual spores and have
two growth phases, i.e., the vegetative phase (mycelia) and the reproductive phase
(fruit bodies). The fungal spores are located in a special structure called the basidium
(for Basidiomycetes) or the ascus (for Ascomycetes). The mushroom continues its
life cycle in three key stages. viz., vegetative growth, reproductive growth, and spore
production by fruit bodies of the mushrooms (Fig. 1).
Fungi lack the most important feature of plants, i.e., the ability to use energy from
the sun directly through chlorophyll. Thus, fungi depend on other organisms for
food, absorbing nutrients from the organic material in which they live. The living
body of the fungus is mycelium made out of a tiny web of threads (or filaments)
called hyphae. Hyphae absorb digestive products, penetrating the substrate to some
extent. Under specific conditions, sexually compatible hyphae will fuse and start to
form spores. The larger spore-producing structures are considered as mushrooms.
The spores released from the gills again germinate and develop to form hyphae,
which is the main mode of fungal vegetative growth. The mushroom produces
several million spores in its life, and this life cycle is repeated each time the spores
germinate to form the mycelium. Mycelial growth is generally coupled with
increased enzyme production and respiration.

2 Global Trends of Mushroom Cultivation

Mushrooms are the common components in folk medicine, especially in Africa, the
Middle East, China, and Japan since ages. Earlier, edible mushrooms were only
harvested wild and were difficult to domesticate and cultivate. Collection from wild
woodlands is still important in the world and particularly in southern Asia [7, 8] and
other developing countries [9]. Mushrooms such as Auricularia, Flammulina, and
Lentinula were most likely cultivated for the first time around the year 600–800 AD
in China and other Asian countries [10]. Their cultivation at large scale started only
at the beginning of the twentieth century when pure cultures of mushroom were
prepared from spore and tissue. As the amount of wild mushrooms shrink from both
4 S. Gupta et al.

Fig. 1 Basic life cycle of

mushroom

Mushroom

Young mushroom

hypha

mycelium

the degraded environment and natural resources and more costly labor, cultivated
mushrooms would not only provide food security but also sustainable and more
nutritious diets [5]. The commercial production of edible mushrooms represents
the unique exploitation of a microbial technology for the bioconversion of agricul-
tural, industrial, forestry, and household wastes into nutritious food (mushrooms).
Mushrooms have the capacity to breakdown the lignin and utilize it as a food source,
thus exposing the underlying cellulose and hemicellulose for food use by other
organisms. Thus, mushroom cultivation represents a very basic natural process of
fungal decay.
With the world’s increasing population and its decrease in per capita arable land,
along with rapid urbanization and industrialization, climate change, and a demand
for quality and functional foods, it will be necessary to focus on secondary agricul-
ture and novel crops, such as mushrooms. Mushroom cultivation could also be an
important part of sustainable agriculture and forestry. Huge quantities of wide
varieties of organic waste are generated from agriculture, forestry, and food pro-
cessing. The impacts of the mushroom business on livelihoods and poverty reduc-
tion are significant and widespread. Mushroom cultivation does not require a lot of
land, significant capital investment, but is a viable and attractive activity for both
rural farmers and semi-urban dwellers. Mushroom cultivation strengthens the live-
lihood of poor and marginal farmers by generating constant farm income and
reduces the vulnerability to poverty. The scale of cultivation can be large or small
based on the capital and labor availability. It can be cultivated on a part-time basis
with little maintenance. Indirectly, mushroom cultivation also provides opportunities
for improving the sustainability of small farming systems through the recycling of
organic matter, which can be used as a growing substrate and then returned to the
land as fertilizer. There are hundreds of identified species of fungi which have made
a significant contribution to human food and medicine. The total number of
Edible Mushrooms: Cultivation, Bioactive Molecules, and Health Benefits 5

described fungi of all kinds is currently 110,000 species [11] of which 16,000 (15%)
species are mushrooms [11–13]. Out of these, more than 3000 species from 231 gen-
era are regarded as prime edible mushrooms [13–15] of which only about 200 are
experimentally grown, 100 economically cultivated, around 60 commercially culti-
vated, and more than 10 produced on an industrial scale in many countries [16].
Approximately 700 mushroom species out of the known 16,000 are considered to be
safe and have medicinal properties [13]. The number of poisonous mushrooms
approximates 500 species. The most acceptable varieties among the cultivated
types are Agaricus bisporus (button mushroom), Pleurotus spp. (oyster mushroom),
Lentinus edodes (Shiitake), and Volvariella spp. (paddy straw mushrooms). In the
second half of the twentieth century, there were rapid changes in rate of growth of
mushroom production and number of species like shiitake, oyster mushrooms, and
wood ear mushroom, and Flammulina were brought under commercial cultivation.
By the end of the twentieth century, the share of button mushroom in total world
production was less than 40%, which in next 10 years became around 30%. Presently
shiitake, oyster, wood ear, and button mushrooms contribute 22%, 19%, 18%, and
15%, respectively in terms of total mushroom production in the world [17]. The
contribution of medicinal mushrooms in world trade has also increased over last few
decades.
Mushroom farming is today being practiced in more than 100 countries, and its
production is increasing at an annual rate of 6–7%. Cultivated mushrooms have now
become popular all over the world. In 1999, the world production of cultivated
edible mushrooms was estimated to be >7 million tons, showing a steady increase
over the last two decades. China is the largest producer, consumer, and exporter of
mushrooms in the world, followed by the United States and the Netherlands
(Table 1). In China, mushroom is the 6th important crop in the country as far as
revenue generation for the nation is concerned. Mushroom production in China in
2010 was 21,524,473 t [18]. The last few decades have witnessed a sharp rise in
diversification in number of mushroom species that have been cultivated, world
mushroom production, commercialization accompanied with mechanization, and in
many cases automation. Mushroom cultivation and its processing have been bene-
ficial to millions of people in China, India, and other developing countries in terms of
financial, social, and health improvement. The global mushroom industry has
expanded very rapidly in the last two decades by the addition of newer types of
mushrooms for commercial cultivation. In addition, cultivation and development of
mushroom industries have positively impacted on economic growth, and this impact
of mushroom cultivation and mushroom derivatives and products on human welfare
in the twenty-first century can be considered globally as a “nongreen revolution.”

3 History and Status of Mushroom Cultivation in India

The history of mushroom cultivation in India goes back to the ages of “Vedas,”
wherein the mention was made in the classical religious scriptures like “Rig Veda”
and “Atharva Veda” about the use of juice from fly agaric mushroom (Amanita
muscaria) as an intoxicating drink, named “Soma.” Vegetarianism is assumed to be
6 S. Gupta et al.

Table 1 Nutritional value of some commercial edible mushrooms (on dry wt. basis)
Mushroom
Nutritional Agaricus Pleurotus Volvariella Lentinula
parameters bisporus spp. volvacea edodes
Protein (%) 29.14 19.59 38.10 18.85
Carbohydrates (%) 51.05 64.34 42.30 63.60
Fat (%) 1.56 1.05 0.97 1.22
Vitamin D (IU/g) 984 487 462.04 205
Sodium (mg/kg) 500.8 208.87 345.34 82.49
Potassium (%) 4.21 2.70 4.16 2.10
K:Na 84:1 129:1 120:1 255:1
Iron (mg/kg) 85.86 183.07 72.51 37.55
Manganese (mg/kg) 7.97 6.47 – 17.48
Zinc (mg/kg) 79.64 162.18 94.28 89.63
[37]

the norm of Indian diet and is primarily based on cereals (wheat, rice, and maize)
which are deficient in protein. Incorporation of mushrooms in Indian diet has the
potential to bridge this protein gap and reduce the problem of malnutrition to a great
extent. For people in a developing country like India, the two main issues are the
quality food and the level of unemployment, in addition to environmental concerns,
which can be very well resolved by popularizing mushroom cultivation among rural
masses and the younger generation [19]. Although time to time, small efforts and
simple research were conducted to introduce mushroom cultivation in India, scien-
tific and systematic research only started in 1961, when the Indian Council of
Agricultural Research, New Delhi, first cultivated Agaricus bisporus at Solan in
Himachal Pradesh, a hilly state of North India. India, primarily being an agrarian
economy, is rich in terms of agrowastes that are not properly utilized by the nation’s
farmers. India produces nearly 700 million tons of agricultural residues which can
profitably be utilized for mushroom cultivation. Even if India uses 2% of its total
agroresidues for mushroom production, the production would be 7.0 million tons of
fresh mushrooms, which will be equal to current global button mushroom produc-
tion. Currently, India is using only 0.03% of these residues to produce about 0.13
million tons of mushrooms and contributes <1% of the total world mushroom
production. Though mushroom production in Asian countries started 1000 years
ago, cultivation of mushrooms is a relatively new phenomenon in India. There has
been significant increase in production of mushrooms in the last few years, espe-
cially of the oyster and paddy straw mushrooms in India. By considering the present
production data, mushroom industry in India recorded an average annual growth rate
of 4.3% per annum. During this period, the productivity has raised from 20% to
24.5% by the releasing of improved strains in commercial edible mushroom. The
total white button mushroom produced in India from both seasonal and high-tech
cultivation units is estimated at 94,676 mt. There has been significant increase in
production and productivity of mushrooms (Fig. 2) in the last few years, especially
Edible Mushrooms: Cultivation, Bioactive Molecules, and Health Benefits 7

140 30

120 25
Production (lakh tons)

100

Productivity (%)
20
80
15
60
10
40

20 5

0 0
1970 1975 1980 1985 1990 1995 2000 2005 2010 2016
Years

Fig. 2 Production and productivity of mushrooms in India [20]

of the oyster and paddy straw mushrooms in India. The country’s production in 2010
was 1.00 lakh metric tons, of which button mushroom accounted for 89% of the total
production, followed by oyster (6%), milky (1%), and others (4%). Punjab,
Uttarakhand, Haryana, Uttar Pradesh, and Tamil Nadu were the leading producers
of the mushroom in the country at the time of 2010. The present production status
revealed that Punjab, Haryana, and Odisha are emerging as the leading states in
mushroom production (Fig. 3).

3.1 Relative Contribution of Different Mushroom Species

In India, there are five mushroom species, viz., white button mushroom (Agaricus
bisporus), oyster (Pleurotus spp.), paddy straw (Volvariella volvacea), milky
(Calocybe indica), and shiitake (Lentinula edodes), which are under commercial
cultivation. Even though cultivation technologies of many exotic mushrooms have
been standardized, the commercial markets are still dominated by Agaricus bisporus,
Pleurotus spp., and Volvariella volvacea. These three mushrooms are contributing
about 96% of total mushroom produced in India (Fig. 4). Milky mushroom
(Calocybe indica) is indigenous tropical mushroom of the country [21]. However,
the commercial cultivation is restricted to southern states of India and contributing
up to 3% to the total mushroom production. Production of paddy straw mushroom
became more popular in the states of Odisha and Chhattisgarh, and its production
was registered at 7% to the total mushroom production. Two to three crops of button
mushroom are grown seasonally in temperate regions with minor adjustments of
temperature in the growing rooms, while one crop of button mushroom is raised in
the northwestern plains of India seasonally. Oyster, paddy straw, and milky mush-
rooms are grown seasonally in the tropical/subtropical areas. In Northeastern states,
8 S. Gupta et al.

AndhraPradesh
Bihar
North East 3%
2% Delhi Goa
Uttar Pradesh 1% 2%
5% Others 3%
5%
Uttarakhand
8%
Gujrat
9%

Tamil Nadu Haryana

8% 12%

Punjab
14%
Himachal Pradesh
7%
Odisha
12% Maharashtra
9%

Fig. 3 Share of different states of India in National mushroom production [20]

Fig. 4 Relative contribution paddy straw Other speciality

of different mushroom species mushroom mushrooms
in total production [20] Milky 7% 1%
mushroom
3%

Oyster
mushroom
16%

White button
mushroom
73%

Uttarakhand and Chhattisgarh, oyster mushroom cultivation is emerging as one of

the leading cottage industry. Hence, for choosing a species for commercial cultiva-
tion, the grower must consider the availability of waste materials to use as a growth
medium, the prevailing environmental conditions, available expertise, available
resources, and market demand.
Edible Mushrooms: Cultivation, Bioactive Molecules, and Health Benefits 9

4 Marketing of Mushrooms

Mushroom cultivation provides an opportunity to generate a highly tradable com-

modity, thereby contributing to income generation. But, majority of the farmers fail
to make profits out of their mushroom production activity due to the poor marketing
strategy [22]. It is the simple system of producers selling directly to retailer or even
to consumer, which has its own limitations like seasonal cultivation of mushrooms
and glut of mushrooms at a particular time especially in North India during the
winter months forcing the distress sale of the mushrooms. This reinforces that effort
to increase the production without solving its marketing problems would be coun-
terproductive. For successful marketing, it is necessary to explore various marketing
options for fresh mushrooms and selling them after processing as value-added
products like pickles, nuggets, dry powder, etc. which could fetch good returns.

5 Types of Cultivated Mushrooms

The most basic concept of mushroom cultivation is that we need to produce an

environment in the substrate that is selectively preferential to the growth of our target
mushroom species and less amenable to other types of microorganisms and pests.
This involves sterilization or pasteurization. Once the substrate is treated in this way
and the majority of the microorganisms are killed, the substrate is suitable for the
introduction of our target species of mushroom. Another potential treatment method
is simply to soak the substrate in water. It prepares a substrate which is more
selective to the higher cellulose-degrading organisms such as oyster mushrooms.
This is the simplest method of substrate preparation, but the chemical treatment
methods are much more effective and not considerably more difficult or expensive.

5.1 Mushrooms Under Cultivation in India

A brief account of some popularly grown mushrooms in India is as follows:

5.1.1 Button Mushroom

The white button mushroom is the most popular of the cultivated mushrooms. This
was the first mushroom to be commercially exploited on an industrial scale and ranks
first at the global level.
The genus Agaricus has two cultivated species, namely, A. bisporus (temperate
button mushroom) and A. bitorquis (tropical or high temperature-tolerant white
button mushroom). A partially decomposed organic matter prepared under aerobic
conditions is using as a substrate for growing button mushroom. This substrate is
generally termed as compost. The purpose of composting is to prepare a physically
and chemically homogeneous medium for the growth of mushroom mycelium. The
composting can be done either by long method of composting (LMC) or by short
method of composting (SMC). In LMC, the entire composting operation is carried
10 S. Gupta et al.

out in open, and pasteurization required for conditioning of compost is achieved by

natural means. SMC is followed by the commercial button mushroom units with
environmental control facilities for composting and cropping.
In SMC, the compost is prepared in two phases. During the first phase, outdoor
composting for 10–12 days followed by indoor pasteurization and conditioning for
7 days in the second phase. During the phase I, the cereal straw is thoroughly wetted
on the cemented floor. This pre-wetted straw is then mixed with the chicken manure
and wheat bran and kept as such for 2 days in the form of a pile. After that, the pile is
broken and restacked. Four days later, the pile is once again broken and the entire
quantity of urea is added. After 2 days, the stack is again turned and watered and
gypsum is added in the third turning. During this stage, the temperature of pile raises
to 70–80  C. The final turning is given on the 12th day when the color of the compost
changes into dark brown and it starts emitting a strong smell of ammonia. The
compost prepared as a result of microbial fermentation process needs to be pasteur-
ized in order to kill undesirable microbes and competitors. The compost from phase I
is then filled in the pasteurization tunnels to the height of 2–2.3 m. At this stage,
temperature inside the tunnel is set at a range of 45–48  C which is then raised to
48–52  C and maintained for 48 h. After 48 h, the compost temperature is raised to
58–62  C and maintained for 4–6 h by inducting the steam. Thereafter, the temper-
ature is brought down to 48–52  C, and the compost is retained at this temperature
for 3 days. The ammonia level in the compost is monitored regularly and if no
ammonia persists in the compost, the compost temperature is brought down to room
temperature. At this stage, the final compost should be granular in structure with
65–70% moisture and pH in the range of 7.2–7.5.

Spawning and Spawn Run

After phase II of SMC, the compost is thoroughly mixed with the spawn at
0.5–0.75% (500–750 g) in polypropylene bags and kept in the cropping room
where the moisture content of substrate is maintained at 65–70% and temperature
at 23  1  C. It takes about 12–14 days for the complete colonization of mycelium.

Casing and Case Run

Once mycelium completely colonizes the compost, it can be switched over to
fruiting stage by covering the surface of the substrate with a thin layer of casing
material. Mixture of 2-year-old FYM and garden soil in 1:1 ratio is most preferable
casing material.

Crop Management
Environmental conditions after casing should be the same as during spawn running.
Within 3 days of application, the mycelium starts growing into the casing layer. Once
mycelial growth is firmly established, the casing is gradually watered up to its
optimum moisture-holding capacity.
At this stage, the microclimate in the cropping room which includes substrates
and air temperatures in the range of 16–18  C and humidity to 85% should be
adjusted. Besides this, carbon dioxide content of the room should be reduced to less
Edible Mushrooms: Cultivation, Bioactive Molecules, and Health Benefits 11

than 1000 ppm by the induction of fresh air, and the room is lighted on a 12-h on/off
cycle as mentioned below to promote the fruiting. This change in environmental
parameters helps to the initiation of pinning and subsequent development of pin-
heads into solid fruit bodies within 3–4 days. The mushroom crop grows in cycles
called as “flushes or breaks.” The uniform fruit bodies of 4–5 cm in diameter should
be handpicked in a twisting motion. Daily watering is required after harvesting the
first flush. 18–20 kg of fresh mushrooms can be harvested from 100 kg of prepared
compost in a total of three flushes.

5.1.2 Oyster Mushroom

Pleurotus mushroom, which is generally referred to as “oyster mushroom” the world
over and as “dhingri” in India, has its origin from the Greek word “pleuro,” which
means formed laterally or in a sideway position, particularly referring to the lateral
position of the stipe (stem) in relation to the pileus (cap). Species of the genus
Pleurotus are well appreciated for their broad adaptability under varied agroclimatic
conditions. The several species of this mushroom occur commonly as wood decom-
posers in forests throughout the world. Pleurotus has always been most attractive to
commercial mushroom growers because of the ease by which most of these species
can be cultivated. This mushroom consists of species that are suitable for both
temperate and subtropical regions. For temperate regions, Pleurotus ostreatus,
P. florida (winter strain), and P. fossulatus and P. eryngii (king oyster) are ideal.
The areas suitable for the button mushroom are equally suitable for the cultivation of
these species. The majority of the oyster species can be best grown in the temper-
ature range of 20–30  C with high relative humidity in excess of 85%. The majority
of the Pleurotus species require 25–30  C for spawn runs and 20–25  C for fruiting.
The oyster mushroom is a low-investment, low-risk, low labor-intensive, and
moderate-profit enterprise. The greatest advantage of the oyster mushroom is that
its production technology is simple and that it can be easily grown at the village level
in small makeshift sheds (huts). Growing the oyster mushroom is less expensive and
more convenient compared to other species, such as the button mushroom. The
method of cultivation is also very simple; unlike the button mushroom, it does not
need any compost preparation or precise temperature control. It can be sold both in
fresh and dried form.

Substrate Preparation
The oyster mushroom has less specificity toward the substrate. Hence, it can be
cultivated on a large number of agrowastes such as cereal straw, sugarcane bagasse,
saw dust, dried grasses, discarded wastepaper, etc. The pasteurization of the
substrate can be achieved either by steam pasteurization or hot-water treatment.
Chemical pasteurization also results in elimination of competitor fungi, but its use is
discouraged due to the residual levels in the mushroom fruit bodies.

Spawning and Incubation

Freshly prepared grain spawn is mixed thoroughly at 2–3% of the substrate on wet
weight basis, and the spawned substrate is filled in the polythene bags. 10 to 15 small
12 S. Gupta et al.

holes made on all sides of the bag facilitate the aeration and drainage of excess water.
During the spawn run stage, the bags are not to be opened, and it doesn’t require
much ventilation or water sprays.

Crop Management
Optimum temperature for mycelia growth of oyster ranges between 22  C and 26  C.
Once the mycelium fully colonizes the substrate, it forms a thick mycelial mat
indicating the readiness for fruiting. During the fruiting period, a relative humidity
of 80–85% needs to be maintained by spraying the water two to three times, and
sufficient ventilation should be provided for air circulation. The color of the oyster is
also influenced by the light intensity and its duration. Fruit bodies raised in bright
light appears in dark brown or gray in color, and the fruit bodies raised in less
intensity of light appears in pale yellowish color.
With suitable crop management practices, mushroom fruit bodies are ready for
harvesting in 6–8 days after pinhead formation. Mushrooms may be harvested while
the edges of the caps are still curled down. Under normal room temperature
conditions, fresh mushrooms can be stored for 2–3 days without any deterioration.
The fruit bodies can be dried under sunlight to the moisture level of 8–10%.
The dried oyster mushroom can be successfully stored for 4–6 months without
losing its original properties.

5.1.3 Milky Mushroom

The milky white mushroom grows during the summer, and it is a tropical mushroom
known for its nutritional value. Its robust size, sustainable yield, attractive color,
delicacy, long shelf life, and lucrative market value have attracted the attention of
both mushroom consumers and prospective growers. Calocybe indica is rich in
protein, lipids, fiber, carbohydrates, and vitamins, and it contains an abundant
amount of essential amino acids and is low in fat. These qualities make it suitable
as a dietary food supplement. The milky mushroom (Calocybe indica) is purely of
Indian origin and highly suitable for the tropical climatic conditions of Central India,
North India, and South India. Its cultivation is now spreading quickly in many Indian
states, including Tamil Nadu, Kerala, Orissa, Haryana, and West Bengal due to
its longer shelf life and adaptability to warm and humid conditions. At present,
Tamil Nadu is the major producer of the milky mushroom in India.
Milky mushrooms can be grown on a wide range of lignocellulosic residues, but
commercial cultivation is done on paddy straw or wheat straw. The cultivation
technology of milky mushroom is a mixture of button and oyster mushroom
cultivation technologies. It does not require compost for its growth specially as the
substrate is prepared as per the technique used for oyster mushroom. Spawning is
done at 4–5% of wet weight of substrate, and the spawned bags are then shifted
to incubation room at 28–32  C under dark conditions. It takes 20–25 days for
completion of spawn run. However, once the spawn run is complete, a layer of
casing material needs to be applied for induction of fruiting. Thereafter temperature
of 30–35  C, R.H. 80–90%, and light intensity at 600–1000 lux is required to be
maintained for cropping. Primordia formation and fruit body maturation takes place
Edible Mushrooms: Cultivation, Bioactive Molecules, and Health Benefits 13

within 8–10 days after casing. Mushrooms are harvested when the stipe length
reaches 7–8 cm by twisting with hand. Harvested fruit bodies can be consumed
fresh or store the clean mushrooms by wrapping in film for 7–10 days at room
temperature.

5.1.4 Paddy Straw Mushroom

The paddy straw mushroom Volvariella sp. prefers to grow on paddy straw; hence, it
is known as paddy straw mushroom. Chinese growers developed its cultivation more
than 300 years ago. Therefore, it was named “Chinese mushroom” [23]. The paddy
straw mushroom is the sixth most important cultivated mushroom in the world [24].
Paddy straw mushroom (Volvariella volvacea) is a world famous edible mushroom
variety that enjoys high demand due to its deliciousness and nutritive value. It is the
only mushroom which can be cultivated both indoor and outdoor. It is a popular
variety among growers because of short cropping season compared to other culti-
vated mushrooms. No other vegetable or cultivated mushroom can be served as a
table dish within a short time after planting, but V. volvacea can do this as it comes
to harvest 10-days post-planting. The climatic conditions prevailing in India are
best suited for the cultivation of this mushroom. Currently, Orissa is the leader in
commercial cultivation of this mushroom.

Outdoor Cultivation
The traditional outdoor cultivation is done seasonally under the shades of trees by
making mushroom beds on the raised platform made from bricks and bamboo poles.
Straw bundles of 45 cm length and 10 cm width are prepared by cutting the top leafy
portion and part of thick stalk near the roots by hand or motorized cutter. The bundles
are soaked in water or in 2% CaCO3 solution for 12–14 h. Then these bundles are
placed side by side followed by placing the mushroom spawn at six to eight spots
and covering of spawn with red gram dal powder. 12–15 beds are prepared in the
similar manner, and the whole lot of beds prepared in a line are compressed a little
and covered with clean plastic sheet.
The spawn run requires a temperature of above 39  C, and spawn run will be
completed in 6–7 days. After the completion of spawn run, the plastic sheet is kept
loosely covered over the beds. The mushrooms start coming from all sides of the bed
after 12–13 days of spawning. They can be harvested at egg stage by holding
between forefinger and thumb followed by twisting clockwise or anticlockwise
direction. The harvested mushrooms can be cleaned and packed in polythene bags
or in paper bags, but the bags should not to be sealed. These packed mushrooms
must be sold for consumption preferably on the same day.

Indoor Cultivation
Indoor cultivation can be done on a substrate/compost prepared by mixing cotton
ginning mill waste and paddy straw in 1:1 ratio on weight basis. The substrate after
mixing is wetted for 2 days, and later the poultry manure is added at 5%. The
substrate is mixed thoroughly and made into a pile. First two turnings are given at an
interval of 1 day each, and calcium carbonate at 1.50% is added at third turning, and
14 S. Gupta et al.

the substrate is left for fermentation for next 2 days. After 4 days of outdoor
composting, the compost is spread on the shelves to a thickness of 10–15 cm. The
steam is introduced inside the cropping room for heat conditioning of the compost.
Temperature is maintained at 62  C for 4–5 h. After a day, the compost is spawned
at 1.5% on wet weight basis, and the beds will be covered with the plastic sheet.
The room temperature is maintained at 32–34  C during spawn run period. The
compost gets colonized within next 4–5 days, and then the beds are sprinkled with
water by removing the plastic sheet. The pinhead starts appearing on 5th–6th day of
spawning. After another 4–5 days, the first flush of mushroom gets ready for
harvesting. The paddy straw mushroom is not suitable for storing in refrigerator
and must be consumed fresh immediately after harvesting or may be stored at room
temperature for few hours.

5.1.5 Shiitake Mushroom

Shiitake mushroom is one of the world’s most popular edible mushroom species
known for its nutrition, flavor, and medicinal properties. It contains a bioactive
compound – lentinan – which has anticarcinogenic and tumor-suppressing proper-
ties. Traditionally shiitake mushroom was cultivated on fallen wood logs stacked in
evergreen forests in China since centuries. But its commercial cultivation gained the
momentum after developing the synthetic log cultivation technology.

Synthetic Log Cultivation

Saw dust obtained from the broad-leaved trees such as tuni, poplar, mango, safeda,
oak, maple, etc. is the main basal ingredient for the preparation of synthetic logs. For
preparation of synthetic logs, saw dust is supplemented with the wheat bran or rice
bran in the ratio of 80:20, and calcium carbonate is mixed at 1% on dry weight basis
to maintain the pH levels below 7.0. Ingredients are thoroughly mixed and moist-
ened to a level of 60–65%. The prepared substrate is filled in the heat-resistant
polypropylene bags and then sterilized for 2 h at 121  C and 22 psi of pressure. After
sterilization, the bags are cooled for a day and thereafter seeded with fresh grain
spawn at 45–50 g per bag. Spawning is done under aseptic conditions using laminar
airflow chamber. After inoculation, the bags are shifted to incubation room where the
temperature is maintained at 22–26  C. Spawn run is generally completed within
30–35 days, and a thick mycelia coat will develop on the surface of the substrate
after 6–8 weeks of spawning. This mycelial coat will eventually turn into brown
color with some exudates. Clumps of mycelium will form on the surface of the
mycelial coat 10–12 days after mycelial coat formation. These clumps will turn into
mushroom fruit bodies at the later stage. The temperature of about 18–20  C and
relative humidity of more than 85% are required to be maintained till the maturity of
the fruit bodies. The fruit bodies keep coming for 15–20 days in a flush after each
shock treatment of cold water. Cold water treatment should be repeated for the
subsequent three to four flushes also till the exhaustion of nutrients in the substrate.
The harvested fresh fruiting bodies of shiitake mushroom can be stored for 3–4 days
at 18–20  C, and keeping them at 4–6  C will extend their shelf life up to 14 days.
The shiitake mushroom can also be stored up to 1 year by drying.
Edible Mushrooms: Cultivation, Bioactive Molecules, and Health Benefits 15

6 Nutrition in Mushrooms

Edible mushrooms have been widely utilized as human food for centuries
and appreciated for texture, flavor, as well as medicinal and tonic attributes [25].
In general, mushrooms contain 90% water and 10% dry matter [26]. They have a
chemical composition, which is attractive from the nutritional point of view [27].
Mushrooms are nutritionally important as they are rich in protein, fibers, and
minerals, while poor in fats. The mushroom protein contains all the nine essential
amino acids required by humans. Mushrooms are considered as a potential substitute
of muscle protein on account of their high digestibility [28]. Besides this, mush-
rooms are also rich source of vitamin B1, B2, B12, C, D, and E [29, 30] and a
relatively good source of nutrients like phosphorus, iron, and vitamins, including
thiamine, riboflavin, ascorbic acid, ergosterol, and niacin [31]. Mushrooms are also
an excellent source of vitamin D which is otherwise not available in other food
supplements [32].
Mushrooms are low in calories, fat-free, cholesterol-free, gluten-free, and very
low in sodium. Minerals such as potassium, iron, copper, zinc, and manganese are
high in fruit bodies. They also have ash, glycosides, volatile oils, tocopherols,
phenolic compounds, flavonoids, carotenoids, folates, organic acids, etc. [33].
Mushrooms are also important from nutraceutical point of view, as they contain
several compounds like unsaturated fatty acids, phenolic compounds, tocopherols,
ascorbic acid, and carotenoids. The nutritional attributes of edible mushrooms and
the health-benefiting effects of the bioactive compounds they contain make mush-
rooms a health food [34–36].
Consumers are now deeply interested in food bioactives that provide beneficial
effects to humans in terms of health promotion and disease risk reduction.
Mushrooms can be considered as functional food which provides health benefits in
addition to nutritional value [38]. The concept of “functional foods” was first
introduced as a factor in the analysis of foods after nutrients [39].
The most common nutrients of mushrooms are discussed as follows:

6.1 Proteins and Amino Acids

The crude protein content of edible mushrooms is usually high but varies greatly
and is affected by factors such as species and stage of development of the
mushroom [40]. The free amino acid level of mushrooms is usually low ranging
from 7.14 to 12.3 mg/g in dry edible mushrooms and contributes to the main flavor
properties of mushrooms [41]. The essential amino acid profiles of mushrooms
reveal that the proteins are deficient in sulfur-containing amino acids, including
methionine and cysteine. However, these edible mushrooms are comparatively rich
in threonine and valine.
16 S. Gupta et al.

6.2 Vitamins

Cultivated mushrooms are a good source of several vitamins, such as riboflavin,

niacin, and folates. The vitamin B2 content in mushrooms is higher than that
generally found in vegetables, [30]. Mushrooms contain moderately high amounts
of folates, and their bioavailability is as good as that for folic acids [42]. In addition
to riboflavin, niacin, and folates, cultivated mushrooms also contain small amounts
of vitamin C and vitamin B1 and traces of vitamins B12 and D2 [30].

6.3 Carbohydrates

Edible mushrooms contain high levels of oligosaccharides and only a low level of
total soluble sugars [43]. The carbohydrate content of edible mushrooms varies with
species and ranges from 35% to 70% DW [44].

6.4 Fatty Acids

The fatty acid level in mushrooms is generally low around 2–8% of distilled water.
The level of polyunsaturated fatty acids as compared to saturated fatty acids is quite
high, constituting more than 75% of total fatty acids of which oleic and linoleic acids
are the most significant, while palmitic acid is the main saturated fatty acid [45].

7 Therapeutic Potential of Mushrooms

The major attribute of mushrooms is their medicinal properties and presence of

bioactive compounds. The pharmacological properties of mushrooms include immu-
nity enhancement, maintenance of homeostasis, regulation of biorhythm, and most
importantly cure and prevention of various life-threatening diseases such as cancer,
stroke, and heart diseases. Medicinal properties of mushrooms including anti-
inflammatory, antioxidant, immunomodulatory, anticarcinogenic, antiviral, anti-
bacterial, antifungal, hepatoprotective, antidiabetic, anti-angiogenic, hypoglycemic,
etc. have been reported [46]. Immunomodulatory and antitumor activities of poly-
saccharide–protein complex (PSPC) from mycelial cultures of mushrooms have
been extensively studied [47–49]. The pharmacological potential of mushrooms
includes the following:

7.1 Mushrooms as Antioxidants

Edible mushrooms possess potent antioxidants. A study of methanolic extracts from

black, red, and snow ear mushrooms found that mushrooms possess an inhibitory
effect on lipid peroxidation, 1,1-diphenyl-2-picrylhydrazyl (DPPH) radical
Edible Mushrooms: Cultivation, Bioactive Molecules, and Health Benefits 17

scavenging and hydroxyl radical scavenging, and a strong reducing power and
ability to chelate ferrous ions [50]. Similar studies on other mushrooms, including
D. indusiata, G. frondosa, H. erinaceus, T. giganteum, F. velutipes, L. edodes,
P. cystidiosus, and P. ostreatus, Agrocybe cylindracea also reported antioxidant
properties of these mushrooms [44, 51].

7.2 Mushrooms as Hypocholesterolemic Agents

Regulation of the cholesterol level is important for the prevention and treatment of
cardiovascular diseases. Edible mushrooms are an ideal food for the dietetic pre-
vention of atherosclerosis due to their high fiber and low fat content. Initial research
on the cholesterol-lowering effects of mushrooms was conducted in Japan in the
1960s, and it was demonstrated that when rats were fed with a high fat and high-
cholesterol diet supplemented with 5% water of the fruiting bodies of L. edodes for
10 weeks, the plasma cholesterol levels of the animals decreased significantly [52].
Several other studies on Lentinula extracts have shown them to cause a significant
decrease in serum cholesterol in young women and old people [53]. In another study,
dietary fiber extracted from P. cornucopiae had a marked in vitro anti-atherosclerotic
effect and patients with coronary disease showed a decreased atherogenic activity
(20–40%) in their sera after the consumption of this mushroom, which confirms its
property of natural cholesterol-lowering agent [54]. It has been suggested that the
fruiting bodies of oyster mushrooms could be recommended for consumption as a
natural cholesterol-lowering agent in the human diet [55]. Dietary fiber isolated from
Auricularia auricula and Tremella fuciformis significantly decreases the serum total
cholesterol (TC) and LDL cholesterol levels [56]. Various studies have shown that
Lentinula mushrooms can lower both the blood pressure and the free cholesterol
level in plasma and can accelerate the accumulation of lipids in the liver by removing
them from circulation [57]. Mushroom like A. auricula-judae displays anti-
coagulation, anti-aggregatory activity in the blood platelets of mice and rats, thus
serving to lower their total cholesterol, total triacylglyceride, and lipid levels [58,
59]. Grifola frondosa has been reported to reduce blood pressure in rats without
changing the plasma high-density lipoprotein (HDL) level or serum cholesterol
level [60].

7.3 Mushrooms as Hypoglycemic Agents

Edible mushrooms are an ideal food for the dietetic prevention of hyperglycemia
because of their high dietary fiber and protein and low fat content [61]. Lectins
isolated from mushrooms (Agaricus campestris and A. bisporus) have been shown to
enhance insulin release in isolated Langerhans rat islets [62]. The presence of a non-
lectin-type component in A. campestris that displays insulin-releasing and insulin-
like activity has also been reported [63]. Guanidine, which is a known hypoglycemic
substance related to the biguanide class of oral antidiabetic drugs, has been found in
edible mushrooms [64].
18 S. Gupta et al.

7.4 Mushrooms as Antitumor Agents

Cordyceps militaris has been used for a long time in Eastern Asia as a nutraceutical
and in traditional Chinese medicine as a treatment for cancer patients. Searching
for new antitumor agents including mushrooms has become a topic of research.
It was reported that Ganoderma lucidum, Phellinus rimosus, Pleurotus florida, and
Pleurotus pulmonarius possess profound antioxidant and antitumor activities [65].
The antitumor activities of the higher basidiomycetes extracts of fruiting bodies of
Boletus edulis and other Homobasidiomycetes were tested against the Sarcoma
180 line in mice and were found to have significant activity [66]. Calvacin was
isolated from the giant puffball (Calvatia gigantea), and it was found effective
against many experimental tumors, including Sarcoma 180, mammary adeno carci-
noma 755, leukemia L-1210, and HeLa cell lines [67]. There are approximately
650 species of higher basidiomycetes that have been found to possess significant
antitumor activity [14, 60].

7.5 Mushrooms as Immunomodulators

Immunomodulators are the most important medicinal mushroom drugs used espe-
cially in Japan, China, Korea, and other East Asian countries today. Some poly-
saccharides or polysaccharide–protein complexes from mushrooms are able to
stimulate the non-specific immune system and to exert antitumor activity through
the stimulation of the host’s defense mechanism [68]. These drugs activate effector
cells like macrophages, T lymphocytes, and NK cells to secrete cytokines like
TNF-a, IFN-g, IL-1b, etc. which are antiproliferative and induce apoptosis and
differentiation in tumor cells [68].

7.6 Mushrooms as Antimicrobial Agents

Mushrooms need antibacterial and antifungal compounds for survival in natural

environment. Hence, they are rich sources of natural antibiotics. Many of the
extracellular secretions by the mushroom mycelium are known to combat bacteria
[69] and viruses [70]. Considerable antifungal and antibacterial activity against
Staphylococcus aureus, Bacillus subtilis, and Escherichia coli has been reported in
several compounds extracted from various mushrooms [71]. It was observed that
whole extracts of Ganoderma pfeifferi inhibit the growth of microorganisms respon-
sible for skin problems [72]. Oxalic acid has been found to be the compound
responsible for the antimicrobial effect of Lentinula edodes (Berk.) against
S. aureus and other bacteria [73]. Ethanolic mycelial extracts from L. edodes possess
antiprotozoal activity against Paramecium caudatum [74].
Edible Mushrooms: Cultivation, Bioactive Molecules, and Health Benefits 19

7.7 Mushrooms as Antiviral Agents

Specific drugs are urgently needed for cure of viral diseases as they cannot be treated
by common antibiotics. Antiviral effects are described not only for whole extracts of
mushrooms but also for isolated compounds. They may act directly by inhibition of
viral enzymes, synthesis of viral nucleic acids, or adsorption and uptake of viruses
into mammalian cells. These direct antiviral effects are exhibited especially by
smaller molecules. Indirect antiviral effects are the result of the immune-stimulating
activity of polysaccharides or other complex molecules [75]. Small molecular
compounds with antiviral activities, several triterpenes from Ganoderma lucidum
(i.e., ganoderiol F, ganodermanon triol, ganoderic acid B), are active as antiviral
agents against human immunodeficiency virus type 1 (HIV-1) [76].

7.8 Mushrooms as Antiallergic Agents

Although extracts of many mushrooms can stimulate the immune system, some
have been reported to suppress immune responses also. This property could be of
benefit for the treatment of allergic diseases that are nowadays increasing worldwide.
Ethanolic extracts of the edible Japanese basidiomycetes H. marmoreus, F. velutipes,
Pholiota nameko, and Pleurotus eryngii show significant antiallergic effects in
mice [77]. Some compounds from G. lucidum (ganoderic acids C and D) have
been shown to inhibit the histamine release from rat mast cells [78, 79]. Eating of
Tricholoma populinum led to the regression of severe allergic symptoms in a patient
with thromboangiitis obliterans and in another patient with urticaria [80]. Hispolon
and hispidin, isolated from fruit bodies of Inonotus hispidus, have been reported to
inhibit the chemiluminescence response of human mononuclear blood cells and the
mitogen-induced proliferation of spleen lymphocytes of mice [81].

7.9 Mushrooms as Anti-inflammatory Agents

Whole mushrooms and extracts may show anti-inflammatory activity due to the
presence of bioactive compounds. Ethanolic extracts from P. linteus and a proteo-
glycan have been shown to induce anti-inflammatory effect in the collagen-induced
arthritis and in the croton oil-induced ear edema test in mice and antinociceptive
effect in the writhing test [82]. The edible mushroom G. frondosa contains ergos-
terol; ergosta-4-6-8(14), 22-tetraen-3-one; and 1-oleoyl-2-linoleoyl-3-palmitoyl
glycerol which inhibits cyclooxygenases I and II activity [83].

7.10 Mushrooms as Hepatoprotective Agents

Ganoderic acids R and S and ganosporeric acid A from G. lucidum showed in vitro
antihepatotoxic activity in the galactosamine-induced cytotoxic test with primary
20 S. Gupta et al.

cultured rat hepatocytes [84]. In vivo study of two fractions of total triterpenoids
extract of G. lucidum protected mice against hepatic necrosis induced by chloroform
and D-galactosamine, and these hepatoprotective effects were perhaps related to the
ability to promote the activity of scavenging enzymes for hepatic-free radicals in
mice and thus to raise the ability of anti-oxidation in mice [85].

8 Bioactive Compounds in Mushroom

Mushrooms are widely used for their high nutritional value as a functional food.
Additionally, they have been highly appreciated for their medicinal and therapeutic
applications [86]. Interestingly, mushrooms are a rich source of biologically active
compounds providing medicinal or health benefits such as the prevention and
treatment of diseases to humans [38]. Edible mushrooms produce a vast diversity
of bioactive compounds such as polysaccharides, proteoglycans, terpenoids, pheno-
lic compounds, steroids, and lectins. These compounds have a wide range of
therapeutic effects and can act as immunomodulatory, anticarcinogenic, antiviral,
antioxidant, and anti-inflammatory agents [87].
Specific bioactive compounds in mushrooms are responsible for improving
human health in a number of ways. Bioactive compounds can be found in mush-
rooms as well as their cell wall components as polysaccharides (β-glucans) and
proteins or as secondary metabolites such as phenolic compounds, terpenes, and
steroids. The concentration and efficacy of bioactive compounds are varied
and depend on the type of mushroom, substrate, fruiting conditions, stage of
development, age of mushroom, storage conditions, and of course cooking pro-
cedures [88]. On the basis of their chemical structure, bioactive compounds of
mushrooms may be classified as peptides and proteins, phenolic compounds, poly-
saccharides, polysaccharides protein complexes, terpenes, terpenoids, etc.

8.1 Peptides and Proteins

Mushrooms produce many bioactive proteins and peptides, primarily lectins which
are non-immune proteins or glycol proteins that bind specifically to fungal cell wall
carbohydrates and have the ability of cell agglutination. These bioactive proteins
possess enzymatic activities such as fungal immunomodulatory proteins (FIPs),
ribosome-inactivating proteins (RIPs), and laccases. Anti-inflammatory peptides of
different molecular weights have been isolated from mushrooms. Chu et al. [89]
isolated an antifungal peptide (pleurostrin) from P. ostreatus, which exhibited
antifungal activity. Wang et al. [90] isolated a peptide (SU2) from Russula paludosa,
which showed antiviral properties. Ngai et al. [91] isolated an antifungal peptide
(agrocybin) from fresh fruiting bodies of the mushroom Agrocybe cylindracea which
exhibited antifungal activity against Mycosphaerella arachidicola, with an IC50
value of 125 μMat different temperatures up to 80  C [92]. Cordymin, a low
molecular weight peptide (10,906 Da) which shows anti-inflammatory activity, has
Edible Mushrooms: Cultivation, Bioactive Molecules, and Health Benefits 21

been purified from Cordyceps sinensis [93, 94] and from Cordyceps militaris [95].
Liu et al. [96] isolated a xylose-specific lectin showing potent antimitogenic and
antitumor activities from fresh fruiting bodies of Xylaria hypoxylon. It has been
reported that lectins isolated from Pholiota adiposa and H. erinaceum exhibited
antiviral and antitumor activities [97, 98]. Zhang et al. [99] isolated a lectin (32 kDa)
from Russula lepida, which exhibited antitumor activity. Ribosome-inactivating
proteins (RIPs) are enzymes that inactivate ribosomes by eliminating adenosine
residues from rRNA. It has been reported that a ribosome-inactivating protein
(marmorin) was isolated from Hypsizygus marmoreus and showed antitumor activity
[100]. Laccases are phenol oxidases widely diffused in basidiomycete and ascomy-
cete fungi which they use to degrade lignocellulosic substrates. Laccases with
antiviral activities have been isolated from Pleurotus eryngii [101] to P. ostreatus
[102]. Zhang et al. [103] purified a laccase from Clitocybe maxima, which also
showed antitumor activity. Some proteins targeting immune cells known as fungal
immunomodulatory proteins (FIPs) are a new of group bioactive proteins and have
been isolated from F. velutipes [104], Ganoderma tsugae [105], and Volvariella
volvacea [106]. Lin et al. [98] isolated an immunomodulatory protein GM I from
Ganoderma microsporum, which showed antimetastasis activity. Du et al. [107]
purified a water-soluble Se-containing protein Se-GL-P (36 kDa) from the
Se-enriched G. lucidum, which exhibited antitumor activity. A glycoprotein
(PCP-3A), purified from Pleurotus citrinopileatus, showed antitumor activity
[108]. Kodama et al. [109] isolated a low molecular weight protein fraction from
Grifola frondosa, which showed antitumor activity.

8.2 Phenolic Compounds

Phenolic compounds are aromatic hydroxylated compounds with one or more

aromatic rings and hydroxyl groups. Anti-inflammatory properties of many mush-
rooms have been attributed to the presence of some phenolic compounds which
include phenolic acids, hydroxycinnamic acids, lignans, tannins, flavonoids,
hydroxybenzoic acids, stilbenes, and oxidized polyphenols [110, 111]. It has been
reported that these compounds exhibited act as free radical inhibitors, peroxide
decomposers, metal inactivators, or oxygen scavengers [112, 113]. Palacios et al.
[114] studied the antioxidant activity of phenolic compounds in various mushrooms
and reported that C. cibarius and C. cornucopioides exhibited the greatest antioxi-
dant effect with respect to the other mushroom species studied. The phenolic
molecule pyrogallol has been extracted from A. bisporus, C. cibarius, and
L. deliciosus [115, 116] which has been found to exhibit anti-inflammatory activity.
Grifolin and grifolin derivatives are farnesyl phenolic compounds which showed
anti-inflammatory properties and have been isolated from the edible mushroom
Albatrellus ovinus [117]. It has been reported that phenol analogous compounds
(hericenones C, D, E, F, G, H) isolated from H. erinaceus had antineurodegenerative
properties [46] and antioxidant activity [118]. Attarat and Phermthai [119] reported
that catechin, a major group of phenolic compounds isolated from Lentinus
squarrosulus, Lentinus polychrous, and L. edodes, exhibited antioxidant activity.
22 S. Gupta et al.

Chowdhury et al. [120] isolated phenolic compounds and flavonoids from

P. ostreatus, L. edodes, and Hypsizygus tessellatus, which showed antioxidant,
antifungal, and antibacterial properties. Dai et al. [121] reported that hispidin, a
class of polyphenols, is an important medicinal metabolite from Phellinus spp. In
most of these studies, a positive correlation was found between the total phenolic
content in the mushroom extracts and their antioxidative properties, which confirms
that edible mushrooms have a potential as natural antioxidants due to the ability of
their phenolics to inhibit lipid oxidation.

8.3 Polysaccharides

Polysaccharides are the major class of bioactive compounds found in mushrooms

and possess significant immune-stimulating, antitumor, antioxidant, antibacterial,
and antiviral activities [122]. Fungal polysaccharides are the most potent mushroom-
derived substances with antitumor and immunomodulating properties. They are
present in cell wall with different types of glycosidic linkages. Mushroom poly-
saccharides with anti-inflammatory properties have been reported in crude extracts
of Lentinus polychrous, Termitomyces albuminosus, and Phellinus linteus. Fungal
pigment melanin possesses antioxidant, immunomodulating, antimutagenic, and
radioprotective properties [123]. The polysaccharides of Flammulina velutipes are
composed of three monosaccharides (glucose, mannose, and xylose) and have been
found to have anti-inflammatory activities [124].

8.4 Glucans

β-glucan is one of the key components of several basidiomycetes and ascomycetes

cell wall. It is a long-chain polysaccharide with β-D glucose as basic subunit linked
to one another by one to three glycosidic chain with one to six glycosidic branches.
β-glucans are able to enhance the immune system and prevent and treat several
common diseases to promote health [125]. Fruit body extracts of Pleurotus
pulmonarius showed mixed α-linkages and β-anomeric carbon linkages, whereas
polysaccharide from mycelial extracts has mainly α-glucan linkages [126]. Lentinan,
the bioactive glucan from Lentinula edodes showed immunomodulatory and anti-
tumor activities [127]. Schizophyllan is the active β-glucan from Schizophyllum
commune [128]. Glucans such as (1,3)-glucopyranosyl from Pleurotus pulmonarius
have been reported to exhibit anti-inflammatory properties [38]. Lavi et al. [129]
reported that ganoderan A and B glucans of Ganoderma lucidum fruiting bodies
showed hypoglycemic effects. On the other hand, ganopoly, the polysaccharide-
containing preparation of G. lucidum, exhibited hepatoprotective effects in patients
with chronic hepatitis B [130]. A β-glucan isolated from the fruiting bodies of
P. ostreatus has also been proven to exert antitumor activity against Hela tumor
cell [131]. Additionally, glucans such as (1,3)- D-glucopyranosyl from
Edible Mushrooms: Cultivation, Bioactive Molecules, and Health Benefits 23

P. pulmonarius have been reported to exhibit anti-inflammatory properties [124].

Lavi et al. [129] reported that polysaccharides of Flammulina velutipes are found to
have anti-inflammatory activities [124].

8.5 Polysaccharide: Protein Complexes

Some polysaccharides such as polysaccharide-K (polysaccharide-Kureha; PSK) also

known as krestin isolated from Trametes versicolor have been identified as poly-
saccharide–protein complexes and showed antimetastatic activity [14]. Coriolan, a
β-glucanprotein complex obtained from submerged grown T. versicolor biomass,
exhibited hypoglycemic effects and ameliorated the symptoms of diabetes [38].
Chatterjee et al. [132] isolated calvacin, a moderately heat stable, nondiffusible
basic mucoprotein from Calvatia gigantea which showed antitumor activity. On
the other hand, ethanolic extracts and a proteoglycan purified from Phellinus linteus
showed anti-inflammatory properties [82, 133].

8.6 Terpenes and Triterpenoids

Terpenes are the largest group of anti-inflammatory compounds in mushrooms.

Several terpenes isolated from G. lucidum showed anti-inflammatory activity.
Some triterpenes from G. lucidum (ganoderic acid C and derivatives) are able to
inhibit the biosynthesis of cholesterol [109], while other triterpenes (ganoderic acid
F) contribute to atherosclerosis protection [134]. The antioxidative and free radical
effects of triterpenoids from G. lucidum have also been shown [38]. El-Mekkawy
et al. [76] reported that different triterpenes from G. lucidum (i.e., ganoderiol,
ganodermanontriol, and ganoderic acid) showed antiviral activity. Sterols and tri-
terpenes (lucialdehyde D, ganoderone A, and ganoderone C) have been isolated
from the fruiting bodies of Ganoderma pfeifferi. Several triterpenes (trametenolic
acid; ergosterol peroxide; 3b-hydroxy-8, 24-dien-21-al; ergosterol; and inotodiol)
isolated from the sclerotia of I. obliquus exhibited anti-inflammatory and anticancer
activities [135]. Han et al. [136] isolated five novel cyathane diterpenes (identified
as cyathins DH) and three diterpenes (neosarcodonin, cyathatriol, and
11-O-acetylcyathatriol) from Cyathus Africans, which showed potent anti-
inflammatory properties. Chen et al. [137] reported that several triterpenes isolated
from Antrodia camphorata which showed neuroprotective activity.

9 Mushroom-Based Biotech Products

Mushroom-based healthcare commercial biotech products with preventive and cura-

tive effects are today available in dried forms as functional food additives
(nutraceuticals) and are largely sold and consumed in the world market. Owing to
the synergistic action of bioactive molecules, majority of mushroom products
24 S. Gupta et al.

possess beneficial health effects and can be used on a regular basis without undesir-
able side effects. Healthy food developed from biotechnologically cultivated
mycelia of medicinal edible mushrooms Hericium erinaceus and Tremella spp. in
combination with other natural substances (medicinal plants, algae, etc.) possess
antioxidant and immune-stimulating activity and regulate the level of blood lipids
and sugar [138, 139]. Some mushroom products are able to decrease high glucose
and lipid levels in blood and are recommended as neuro- and vasotonics,
hepatoprotective, and thrombolytic agents [87]. Nutritive, anti-inflammatory, regen-
erative, and antioxidant properties of several mushrooms make their usage perspec-
tive in manufacturing of cosmetic products [140]. Mushrooms are currently
proposed as highly active ingredients in world production of hair and skin
care products. Tremella mushrooms contain hydrophilic agent–polysaccharide
glucuronoxylomannan (GXM) with anti-inflammatory and wound healing proper-
ties largely used in cosmetology [141]. Tremella cosmetic products are applicable in
treatment of neurodermatitis and sclerodermatitis. They prevent skin pigmentation
and stimulate blood circulation. Biological characteristics of mycelia, particularly
fast growth and easy reproduction in culture conditions, are assisting biotechnolog-
ical cultivation of medicinal mushrooms to obtain desired bioactive molecules and
biotech products. Thus, mushrooms have significant biotechnological potential.

10 Role of Mushroom Culture Collections

Currently, two different approaches in biotechnological cultivation of mushrooms

are used. First is fruit body production which is a long-term process and takes
1–2 months, while cultivation of mycelia takes a few days. Establishment and
maintenance of culture collections of different group of mushrooms are of valuable
importance to study their biodiversity, genetic resources, and biotechnological
potential. The submerged cultivation of mycelia is the best technique to obtain
biomass and desired bioactive molecules for further development of consistent and
safe healthcare mushroom biotech products and thus has significant industrial
potential.

11 Future Trends

Mushrooms can make an important contribution to the livelihoods of rural and semi-
urban dwellers through food security and income generation, and mushrooms can
make a valuable dietary addition through their protein and micronutrient content as
well as their medicinal properties. Its cultivation is highly compatible with a variety
of other traditional agricultural and domestic activities and can make a particularly
important contribution to the livelihoods of the disabled, of women, and of the
landless poor who, with appropriate training and access to inputs, can increase their
independence and self. The high benefit/cost ratio, the easily obtained and inexpen-
sive agrowastes, and congenial weather conditions make adoption of mushroom
Edible Mushrooms: Cultivation, Bioactive Molecules, and Health Benefits 25

cultivation a lucrative means of societal development. Mushroom cultivation can

represent a valuable small-scale enterprise option. The FAO has been actively
promoting mushroom cultivation for rural development and food security in devel-
oping countries [142].
India has tremendous potential for mushroom production, and all commercial
edible and medicinal mushrooms can be easily grown here. Although, India is not a
major producer of any of the mushroom varieties, it does cultivate all edible and
medicinal mushrooms in one or other part, due to its diverse climatic conditions.
Mushroom cultivation is based on recycling of agricultural residues, which are
available in huge quantities in every nook and corner of the country. At present,
the areas with rice–wheat cropping system of India are facing tough challenge to
handle the mounting crop residues. Mushroom cultivation can effectively utilize
these agroresidues for production of protein rich food and plays crucial role in
management of these agroresidues. The supply and demand gap in the world trade
of mushrooms and the shrinkage of production in western countries due to high labor
costs have resulted in better market prices for Indian mushroom producers. With a
domestic population of more than 1.2 billion, India itself is a large market for
mushrooms. To be successful in both domestic and export market, it is essential to
produce quality fresh mushrooms and mushroom fortified value-added products at
competitive rates without any agrochemical residues. Efforts should also be made to
exploit the commercial utilization of mushroom substrate left after cultivation for
preparation of organic manure, vermicompost, briquettes, etc. One final reason for
optimism concerning India’s potential as a major mushroom producer is its strategic
geographical location, making it more convenient to export mushrooms to the
Middle East, Southeast Asia, and European countries. To be successful in both
domestic and export markets, it is essential to produce quality fresh mushrooms
and processed products devoid of pesticide residues and at competitive prices.
However, mushroom cultivation is yet to be adopted by Indian farmers and mush-
room growers on a large scale. The gap in technology knowledge and its adoption
needs to be bridged by providing training to farmers regarding various aspects of
mushroom cultivation.
Mushrooms are functional food and are a source of biologically valuable com-
ponents that offer great therapeutic potential for the prevention and control of several
diseases. Further research and clinical trials needs to be carried out to validate that
mushrooms are source of bioactive molecules with medicinal applications. They
may be used directly in the diet to promote health, taking advantage of the additive
and synergistic effects of the bioactive compounds present in them. Research is
needed to elucidate the different roles of multiple active compounds and the path-
ways involved. The potential therapeutic implications of mushrooms are enormous
but detailed mechanisms of the various health benefits of mushrooms to humans still
require intensive investigation, especially with the emergence of new evidence of
their health benefit effects. The exploration of newly cultivated mushrooms and
isolation of their active ingredients with mechanism-based potential therapeutic
value remains a challenge, and hence mushrooms will keep on to be the foremost
spotlight of research in the upcoming prospect as well. Medicinal mushrooms
26 S. Gupta et al.

represent a growing segment of today’s pharmaceutical industry owing to the

plethora of useful bioactive compounds. Conservation and cloning of therapeutic
mushrooms is needed for sustainable development. Dedicated research should be
undertaken to isolate, purify, and structural1ly investigate of novel anticancer and
immune-stimulator compounds.

White button mushroom (Agaricus bisporus) Brown button mushroom (Agaricus bisporus)

Milky mushroom (Calocybe indica) Macrocybe mushroom (Macrocybe gigantea)

Shiitake mushroom (Lentinula edodes) Paddy straw mushroom (Volvariella volvacea)

Winter mushroom (Flammulina velutipes) Reishi mushroom (Ganoderma lucidum)

(continued)
Edible Mushrooms: Cultivation, Bioactive Molecules, and Health Benefits 27

Gray oyster (Pleurotus pulmonarius) Pink oyster (Pleurotus djamor)

Blue oyster (Hypsizygus ulmarius) Yellow oyster (Pleurotus cornucopiae)

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