Appl Microbiol Biotechnol (2010) 85:13211337
DOI 10.1007/s00253-009-2343-7
MINI-REVIEW
Cultivation of Pleurotus ostreatus and other
edible mushrooms
Carmen Snchez
Received: 1 September 2009 / Revised: 1 November 2009 / Accepted: 1 November 2009 / Published online: 3 December 2009
# Springer-Verlag 2009
Abstract Pleurotus ostreatus is the second most cultivated
edible mushroom worldwide after Agaricus bisporus. It has
economic and ecological values and medicinal properties.
Mushroom culture has moved toward diversification with
the production of other mushrooms. Edible mushrooms
are able to colonize and degrade a large variety of
lignocellulosic substrates and other wastes which are
produced primarily through the activities of the agricultural,
forest, and food-processing industries. Particularly,
P. ostreatus requires a shorter growth time in comparison
to other edible mushrooms. The substrate used for their
cultivation does not require sterilization, only pasteurization,
which is less expensive. Growing oyster mushrooms convert a
high percentage of the substrate to fruiting bodies, increasing
profitability. P. ostreatus demands few environmental
controls, and their fruiting bodies are not often attacked by
diseases and pests, and they can be cultivated in a simple and
cheap way. All this makes P. ostreatus cultivation an
excellent alternative for production of mushrooms when
compared to other mushrooms.
Keywords Pleurotus ostreatus . Mushroom cultivation .
Edible mushrooms
Introduction
Cultivation of edible mushrooms is a biotechnological
process for lignocellulosic organic waste recycling. It might
C. Snchez (*)
Laboratory of Biotechnology, Research Centre for Biological Sciences,
Universidad Autnoma de Tlaxcala,
Apartado postal 129,
Tlaxcala, Tlax CP 90000, Mexico
e-mail: sanher6@hotmail.com
be the only current process that combines the production of
protein-rich food with the reduction of environmental
pollution (Beetz and Kustudia 2004). The production of
mushrooms is regarded as the second most important
commercial microbial technology next to yeast (Pathak et al.
2009). Mushrooms have been eaten and appreciated for their
flavor, economical and ecological values, and medicinal
properties for many years. In general, mushrooms contain
90% water and 10% dry matter (Morais et al. 2000; Snchez
2004). They have chemical composition which are attractive
from the nutritional point of view (Gbolagade et al. 2006;
Dundar et al. 2008; Table 1). Their nutritional value can be
compared to those of eggs, milk, and meat (Oei 2003).
Mushrooms also contain vitamins and an abundance of
essential amino acids (Snchez 2004). The total energetic
value of mushroom caps is between 250 and 350 cal/kg of
fresh mushrooms (Oliver and Delmas 1987; Laborde 1995).
Some mushroom can be cultivated easily and have significant
worldwide markets. Over 200 species have been collected
from the wild and used for various traditional medical
purposes, mainly in the Far East (Snchez 2004). Roughly
300 mushrooms species are edible, but only 30 have been
domesticated and ten grown commercially (Barny 2009).
The principal cultivated mushroom worldwide is Agaricus
bisporus followed by Pleurotus sp. (Rhl et al. 2008),
Lentinula edodes, and other mushrooms that have already an
important place in the market.
Mushrooms have the ability to degrade several lignocellu-
losic substrates (Fig. 1; Snchez 2009) and can be produced
on natural materials from agriculture, woodland, animal
husbandry, and manufacturing industries (Table 2). As it is
shown in Fig. 1, laccases or ligninolytic peroxidases (LiP
and MnP) produced by white-rot fungi oxidize the lignin
polymer, thereby generating aromatic radicals (a). These
evolve in different non-enzymatic reactions, including
1322 Appl Microbiol Biotechnol (2010) 85:13211337

0.192
0.06
1.97
1.30
0.80
1.91
0.05
0.77
1.20
2.82
3.10
0.85
2.50
C-4-ether breakdown (b), aromatic ring cleavage (c), C-
Zn C breakdown (d), and demethoxylation (e). The aromatic
aldehydes releases from C-C breakdown of lignin or
0.12
0.12
0.16
0.07
0.02
0.05
0.07
0.08
0.07
0.09
0.13

0.14
0.11
synthesized de novo by fungi (f, g) are the substrate for
Cu

H2O2 generation by aryl-alcohol oxidase in cyclic redox

reactions involving also aryl-alcohol dehydrogenases.

0.163
Phenoxy radicals from C4-ether breakdown (b) can
0.67
0.52
0.72
0.25
0.32
0.08
0.20
0.10
0.70
0.85
0.32
0.42
Fe

repolymerize on the lignin polymer (h) if they are not first

reduced by oxidases to phenolic compounds (i). The
phenolic compounds formed can be again reoxidized by

0.021
0.29
0.07
0.90
0.50
0.15
0.80
0.45
0.30
0.72
0.85
0.29
0.44
Mn

laccases or peroxidases (j). Phenoxy radicals can also be

subjected to C-C breakdown (k), yielding -quinones.
Quinones from g and/or k contribute to oxygen activation in
2.50

4.10

8.10

9.95

1.97
19.85

14.50
20.90

13.35

24.20
31.10

18.75
15.0

redox cycling reactions involving quinone reductases,

P

laccases, and peroxidases (l, m). This results in reduction

of the ferric iron present in wood (n), either by superoxide
0.35
2.15
2.00
3.50
1.50
0.25
6.50
0.65
2.00
1.70
2.20
5.90

cation radical or directly by the semiquinone radicals, and its

Na

reoxidation with concomitant reduction of H2O2 to hydroxyl

free radical (OH.) (o). The latter is a very strong oxidizer
9.51

4.32
37.40
22.15
28.80
44.20

14.75
43.35
16.80
58.75
48.40
18.60
51.45

that can initiate the attack on lignin (p) in the initial stages of
K

wood decay, when the small size of pores in the still-intact

cell wall prevents the penetration of ligninolytic enzymes.
Then lignin degradation proceeds by oxidative attack of the
1.45
2.45
3.90
3.10
0.80
1.50
4.55
0.55
3.65
6.00
0.55
2.45
0.46
Mg

enzymes described above.

Poppe (2000) reported that there are about 200 kinds of
waste in which edible mushrooms can be produced.
0.75
1.75
5.20
4.30
1.90
0.40
1.57
3.90
3.50
5.30
1.50
0.80
0.10
Ca

However, mushroom production generates an enormous

amount of used spent substrate which might also be a
source of environmental contamination. Several uses for
4.95
6.50
6.55

9.20

9.65
11.60

9.40

5.84
10.90

10.60

12.90
15.80

12.65
Ash

spent mushrooms substrate (SMS) are being evaluated, and

some of them have already been established (Snchez
Table 1 Nutrimental value of several edible mushrooms (mg/100 g dry matter)

2004). The production of mushrooms in 2005 was (in metric

3.45
5.40
4.30
5.03

4.40
11.65
8.30

8.50
15.60

12.80
10.35
Fiber

7.0

tons) 1,411 in China, 382 in USA, 245 in Netherlands, 139 in

France, 138 in Spain, 135 in Poland, 88 in Italy, 80 in Canada,

77 in Ireland, and 74 in the UK (FAOSTAT 2005; USDA
Protein

2005; WBWDI 2007; Fig. 2).

8.90
5.80

24.30
16.30
15.10
17.40
10.30
27.70
32.80
13.50
26.05
16.75
24.0

Mushroom cultivation
Sugars

5.35

7.70
8.95
7.85
8.75

7.75
9.85
10.15
14.70
17.20

14.15
14.65

Mushroom culture involves several different operations, each

of which must be carefully performed. Substrate preparation,
inoculation, incubation, and production conditions depend on
Lipids

5.05
4.05
7.55

1.70
1.05
5.75
5.50
8.75
6.70
4.20

1.15
10.25

10.95

Gbolagade 2006; Dundar et al. 2008

the mushroom species to be cultivated (Fig. 3). The first stage

involves obtaining pure mycelium of the specific mushroom
strain. The mycelium can be obtained from spores (Fig. 3a),
Termitomyces microcarpus
Psathyrella atroumbonata

from a piece of the specific mushroom (Fig. 3b), or from

Schizophyllum commune
Pleurotus tuber-regium
Lycoperdon giganteum

Termitomyces globalus
Tricholoma lobayensis
Auricularia polytricha

several germplasm providers such as American Type

Volvariella esculenta
Pleurotus sajor-caju
Lycoperdon pusilum
Lentinus subnudus
Edible mushroom

Culture Collection and National Center for Agricultural

Pleurotus florida

Utilization Research (Fig. 3c). To obtain inoculum, the

mycelium is developed on cereal grain, e.g., wheat, rye, or
millet (Fig. 3d), which is usually called the spawn
(Fig. 3e; Chang and Hayes 1978; Chang and Miles 1989).
Appl Microbiol Biotechnol (2010) 85:13211337 1323

Fig. 1 Lignin biodegradation HC LIGNIN

process by white rot fungi
HOOH2
HOCH2
HC HC LIGNIN
OCH3
HC O HCOH

H2COH HOCH2
OCH3
O CH HC O
h
OH* C=O HCOH

H2OC OCH3 H2COH

OH O
HC LIGNIN
LIGNIN
o j HC

LACCASE
H 2O 2 a
PEROXIDASE OCH3
OH
3+
Fe g H2COH
n AAO?
i
HC LIGNIN H2COH
AAO MeOH
CH2OH HC=O e HCOH HC LIGNIN

d HC
+* b
OCH3 OCH 3 OCH3
O-R O-R O c
R H2COH OCH3
AAD
f LIGNIN
O*
HC
-* LACCASE
O2
PEROXIDASE HCOH
m

LACCASE g
OH PEROXIDASE C
ll O
O OCH3
ll OH
OCH3 ll OCH 3 k
O
OH
QR l

Source: Martnez et al. 2005

The purpose of the mycelium-coated grain is to rapidly
colonize the specific bulk growing substrate (Fig. 3f). The
success of mushroom production depends in great part on the
quality of the spawn, which must be prepared under sterile
conditions to diminish contamination of the substrate
(Fig. 3g). Several studies have been done to improve the
quality and develop new techniques for its production. For
example, the spawn for cultivation of P. ostreatus has been
prepared in different ways: on grain, such as wheat,
sorghum, and paddy (Beetz and Kustudia 2004; Nwanze
et al. 2005; Elhami and Ansari 2008) and on grain mixed
with grain straw (Muthukrishnan et al. 2000; Sainos et al.
2006; Pathmashini et al. 2008).
Cultivation of Pleurotus ostreatus
P. ostreatus is also known as oyster mushroom, hiratake,
shimeji, or houbitake (Mizuno and Zhuang 1995;
Bononi et al. 1995; Rhl et al. 2008). For many reasons,
the fungal Pleurotus genus has been intensely studied and
cultivated in many different parts of the world. It is produced
on a variety of lignocellulosic substrates (Table 2).
Table 2 Ligninocellulosic residues used for different mushrooms cultivation
1324

Scientific name English name Japonese name Substrate Reference

Agaricus bisporus
Pleurotus sp (P. ostreatus,
P. sajor-caju, P. eryngii,
P. pulmonarius,
P. eous, P. florida)
Lentinula sp. (L. connotus,
L. edodes)
Volvariella volvacea
Tuber sp., T. melanosporum Vitt., Black truffle black perigord,
T. magnatum Pico ex Vitt.)
Stropharia sp. (S. rugosoannulata Roundheads
Farlow, S. rugosoannulata Farlow
f. lutea hongo)
Hericium sp.
Auricularia auricula
Grifola frondosa
Flammulina sp.
Pholiota nameko
Tremella
Button, white, table, portobello,
crimini, Swiss brown, gremini,
Italian brown and Italian roman
brown mushroom, Champignon
Oyster, king oyster mushroom
Japanese forest mushroom, Shiitake
mushroom black forest, Black oak
Straw mushroom or paddy straw
mushroom
piedmontese
White truffle
Lion's head or Pom pom
Jelly ear
Maitake, hen of the woods, rams head,
sheep's head, cloud mushroom,
dancing mushroom
Winter, velvet stem, golden, snow
puff
Fat pholiota
White jelly
Himematsutake or Agarikusutakea Rice straw, wheat straw, horse manure Beetz and Kustudia 2004; Toker et al. 2007
(name for Agaricus blazei) (fresh or composted)
Wheat straw and waste tea
Leaves
Hiratake Coffee pulp, sawdust, cocoa, peanut and Beetz and Kustudia 2004; Rani et al. 2008;
coconut shells, cotton seed hulls, Jamaica, Shah et al. 2004; Daba et al. 2008; Fan
cassava peels, cotton, sorghum, banana, et al. 2003; Cayetano-Catarino and
corn stalks, grass, clover, wood Bernab-Gonzlez 2008; Philippoussis
Wastes of rice, wheat, sawdust, cotton from et al. 2001; Onuoha et al. 2009; Sivrikaya
textile industry, corncobs, crushed bagasse and Peker 1999; Abrar et al. 2009
and molasses from sugar industry, water
hyacinth, water lily, bean, wheat straw,
leaves, oil-palm fiber, paper and paddy
Shiitake Sorghum stalk, banana, coffee pulp, sawdust, Rani et al. 2008; Beetz and Kustudia 2004;
cotton seed hulls bran, leaves, corncobs, Kapoor et al. 2009
bracts of pineapple, cotton seed meal,
peanut meal, wheat bran, rice bran and
soybean mea
Wastes from paddy, sugar cane bagasse,
sugar cane, coffee pulp, wheat
Fukurotake Banana leaves, cocoyam peelings and Beetz and Kustudia 2004; Belewu and
oil-palm pericarp, palm fibers, rice husk, Belewu 2005; Obodai et al. 2006; Ukoima
and sawdust et al. 2009
Wastes from rice, wheat, cotton, textile
industry, water hyacinth, and water lily
Se you fuyu shuororo (name for Mungbean husks, potting soil Carluccio 2003
violet truffle)b
Saketsubatake, kisaketsubatake Wastes from wheat, sawdust Beetz and Kustudia 2004
Yamabushitake Sawdust, corncobs, distillers grain waste Beetz and Kustudia 2004
Kikurage Sawdust, sawdust-rice bran Beetz and Kustudia 2004
Mai Take, Maitake Sawdust, spent sawdust matrix, corn, barley, Beetz and Kustudia 2004; Hideno et al.
sorghum, and rice 2007; Montoya and Varn 2008
Enokitake Sawdust Beetz and Kustudia 2004
Numerisugtake Logs sawdust-rice bran Beetz and Kustudia 2004
Shirokikurage Logs Beetz and Kustudia 2004

a
MAYA ethnobotanicals (2009)
b
http://zipcodezoo.com, 2009
Appl Microbiol Biotechnol (2010) 85:13211337
Appl Microbiol Biotechnol (2010) 85:13211337
Fig. 2 Top mushrooms cultivation countries
Substrate preparation The substrate is milled to a length of
about 2 to 6 cm. In other substrates, the chopping is not
required. One of the most common substrates used for
modern mushrooms is a mixture. This mixture of cotton-
seed hulls and wheat straw has a higher water holding
capacity than cottonseed hulls used alone. Pasteurization,
used on some commercial mushroom farms, is carried out
by filling the ingredients into revolving mixers, water is
added to the desired level, and live steam is injected into
the mixer while it is in operation (Royse 2007).
Inoculation After completion of pasteurization (60 C for 1
to 2 h), the substrate is cooled and spawned with the desired
strain.
At time of spawning, a delayed release supplement (rates
of 3% to 10% of dry substrate weight) may be added to
increase yield and size of the mushroom (Royse et al. 1991;
Royse and Zaki 1991). Use of supplements, however, may
cause overheating of the substrate if growers are not able to
anticipate and control air temperatures to maintain a steady
substrate temperature.
Spawning and spawn rate Growers have sought, in the
past, to optimize the amount of spawn used to inoculate
their substrate. Increasing the amount of spawn used (up to
5% of the wet weight of the substrate) has resulted in
increased yields (Royse 2002). Increasing spawn rates from
1.25% to 5% may result in increases of nearly 50%. Yield
increases may be due to several factors. First, the increased
level of nutrient available in higher levels of spawn used
would provide more energy for mycelial growth and
development. Second, more inoculum points, available
from increased spawn levels, would provide faster substrate
colonization and, thus, more rapid completion of the
1325
production cycle. Finally, a more rapid spawn run would
reduce the time non-colonized substrate is exposed to
competitors such as weed molds and bacteria (Table 3).
Filling plastic bags with substrate The pasteurized sub-
strate is spawned and filled (from 25 to 30 lbs) into clear or
black perforated polyethylene bags.
Incubation The bags are incubated for 12 to 14 days at
25 C and then transferred to the production room (Royse
2007).
Mushroom production The mushroom begins to form
around the edges of bag perforations. The bags are
maintained under optimal temperature, moisture and other
conditions for mycelium growth, and the conditions that
favor fruiting. Mushroom shelves and suspended systems
are the main systems used for Pleurotus cultivation
(Fig. 3h). Studies on the mushroom cultivation on the use
of different strains, different lignocellulosic substrates,
different types of spawn, moisture, physicochemical con-
ditions, etc. are important for the cultivation productivity of
each particular mushroom (Mandeel et al. 2005; Saavedra
et al. 2006; Kirbag and Akyuz 2008; Onuoha et al. 2009).
Harvest The mushrooms are harvested from the substrate
approximately 3 to 4 weeks after spawning depending on
strain, amount of supplement used, and temperature of
spawn run.
Cultivation of A. bisporus
Agaricus sp. is also known as himematsutake. It is the
principal cultivated mushroom worldwide, and its substrate
is the most complex culture medium used for edible
mushroom production (Fig. 3i). Several studies on the
development of this mushroom have improved its cultiva-
tion (Stoop and Mooibroek 1999; Kothe 2001; Bechara et
al. 2004, 2008; Nogueira de Andrade et al. 2008; Kapoor et
al. 2009). The substrate for the cultivation of this
mushroom is horse manure compost, which consists of a
mixture of horse manure, some broiler chicken manure, and
water to which gypsum is added for structural stability and
for stabilizing pH. Alternatively, a synthetic compost can be
made which is based on wheat straw and broiler chicken
manure. When carried out in the open air, phase I of
composting is done in long narrow stacks between 1.5 and
2.0 m in width and height (wind-rows). The stacks are
turned twice a week in order to obtain uniform degradation
of the mixture. At the end of the process, usually 78 days
after setting up the stacks of 1 t of horse manure, a yield of
just over 1 t of compost will have been obtained. This fresh
1326 Appl Microbiol Biotechnol (2010) 85:13211337

Fig. 3 Cultivation and harvest-

ing of mushroom

Source: Modified from Stamets (1995)

compost (phase I), which still smells strongly of ammonia, done by raising the air temperature to 56 C and keeping it
is subjected to phase II treatment before actual cultivation there for approximately 5 to 6 h. Compost temperature then
can start. Phase II consists of a pasteurization process increases to slightly higher values, and this is maintained
followed by further high-temperature fermentation. This is for at least five more hours. Thereafter, the compost is kept
Table 3 Troubleshooting in the mushrooms cultivation

Problem Cause Solutions

Mycelium fails to form Improper initiation strategy Consult parameter of growth. Alter moisture, temperature, light, carbon dioxide, etc.
Note: If the substrate is too moist, decrease moisture
Chlorinated or contaminated water Use activated charcoal water filters to eliminate chemical contaminants or any other ways
of simple or appropriate technology
Bad substrate Check substrate. Spread the substrate and remix the substrate, package again, make sure
all raw materials are good and fresh
Note: It is necessary to pasteurize immediately after bagging otherwise fermentation gas
will slow down the rate of growth of mycelium or stop mycelium growth
Bad pasteurization Check method of pasteurization. Release all air and make sure there is continuous steam
before starting pasteurization for a period of 3 h
Substrate in the bag is too hot when inoculation Make sure that the substrate bag is not too hot before inoculation
Appl Microbiol Biotechnol (2010) 85:13211337

Poor spread of mycelium, bad smell, spots,
and mites
Mycelium grows but fails to produce
mushrooms
Mushrooms form, but abort or delay
mushrooming
Bad strain or spawn Obtain younger strain of known vitality and history
Spawn contaminated Pasteurize and inoculate again with good spawn
Forgot to inoculate the bag Make sure to inoculate
Good pasteurization but must decrease the temperature in the Slowly decrease the temperature in the chamber. Do not open the cover of the chamber too
pasteurization chamber quickly. Check that the cotton plug is tightly closed
Pasteurization was too quick and/or the chamber door was opened
too quickly
Inoculation process Inoculate in hygiene conditions; clean and with no air movement
Too high density in the incubation area, not enough ventilation to Spread the substrate bag and make more air ventilation in the incubation area. Check
decrease accumulated temperature temperature and control surroundings to maintain of 25 to 35C
Too high carbon dioxide Not more than 5% carbon dioxide. Check ventilation
Hygiene of the incubation house Improve hygiene in the incubation house
Mycelium develops in patches. Substrate is not evenly prepared, Mix well the substrate
and some parts have more nutrients than others
Bacteria, other fungi contamination Check the process causing contamination. Separate contaminated bags as soon as possible.
Remix substrate separately. Remake substrate bags and pasteurize for a longer time.
Follow process
Mite contamination Immediately separate contaminated bags and pasteurize again. Continue the normal
process
Note: Keep hygiene management; make sure to clean every thing (person, area, tools,
equipment, and surroundings during every step. Stop using the area to cut the life cycle
of all contaminants for a period of at least 1 to 2 weeks. For serious contamination cases,
spray area with chemicals. Use black-light with water or sticky-trap to decrease insects
Substrate formula is not suitable Adjust the formula; check pH, sawdust, additives, etc
Mites, mold, virus, bacteria and insects Check pasteurization process, inoculation, other processes and mushroom house
management for hygiene
Inhibited by environmental toxins Remove source of toxins
Bad strain or Shawn Acquire new strains
Primodia and growth condition of fruiting body are not good Check temperature and humidity. Open or close doors and window to adjust accordingly
enough
There is contamination such as mold, bacteria, insects, worms, and Check hygiene, adjust environment of light, temperature, humidity, and ventilation. In
1327
Table 3 (continued)
1328

Problem Cause Solutions

mites
Chemical contamination from solvents, gas, chlorine, etc
Bad strain
Mushrooms form, but stems are long; caps Inadequate light
underdeveloped Excessive carbon dioxide
Massive numbers of mushrooms form; few Too long time incubation
develop Lack of oxygen, inadequate light
Inadequate substrate nutrition or low quality
Low rate mycelium growth
Poor strain
Mushrooms are deformed, decay and die Disturbed by germs or competing microorganisms
Dirty surface of substrate bags
Not enough air ventilation, too high humidity
Bad strain
Use of chemicals during this period
Mushrooms produced only in the first flush, Inadequate substrate nutrition
fail to produce subsequent flushes Competitors
Poor growing house management
Bad strain
Mushrooms small sized Too many mushrooms coming out at the same time
Lack of nutrients in substrate
Change of weather
Spawn unhealthy
Pests and insects Natural occurrence, humid climate
Mushroom waste lying around mushroom house
Ants
Mushrooms are light in weight Shortage of water
Mushroom quickly spoil Mushrooms too mature when harvested
Mushrooms too warm before packaging
Mushrooms too wet when harvested
Mushrooms stored beyond shelf life
Rotting spot on the mushroom fruiting body Bacteria (Pseudomonas tolaasii, Pseudomonas fluorescens)
because of bacteria during flush on Oyster mushroom
more severe cases, use half a teaspoon of sulfur in 3.5 l of water. Mist the bags and the
surface of mushrooms. Remove contaminated bags from mushroom house and recycle
Remove toxins
Acquire a new strain or find a new supplier
Increase or adjust light to correct wavelength
Increase air exchange, open doors or windows and close at correct time
Shorten the period for the formation of primodia
Increase air ventilation and open more windows or doors to receive more light
Reformulate or check raw materials
Use the high rate spawn or adjust good conditions for rate of growth
Obtain better strain
Adjust mushroom house to favor mushrooms and not germs and competitors
Clean the surface of substrate
Increase air circulation. Reduce humidity to the prescribed levels. Surface water must
evaporate from mushrooms several times per day. Check watering; if there is water in
bags, pierce bags and drain water
Acquire better strain
Never use chemicals during the fruiting stage
Reformulate
Check hygiene; adjust light, temperature, humidity, air, and ventilation
Improve management
Acquire new strain
Reduce the size of opening(s)
Review quality of substrate
Beware of wide range changes in temperature
Check origin of spawn
Place lemongrass plants around mushroom house. Spread lime on shelves, on poles, and
ground in the mushroom house. Clean (and maintain clean) the mushroom house properly
Try to use the waste as fertilizer or recycle
Mix detergent with water and place on their paths. Do not put on mushroom
Check humidity of mushroom
Harvest when younger
Chill mushrooms before placing in marketing containers
Reduce humidity several hours before harvesting
Sell mushrooms faster
Control humidity in the mushroom house and maintain 8085%. Give enough time for
water to evaporate from mushroom surfaces before further watering. For severe cases,
use 113 g chlorine mixed in 45 l of water or 4 oz of chlorine per gallon of water

Source: FAO 2001

Appl Microbiol Biotechnol (2010) 85:13211337
Appl Microbiol Biotechnol (2010) 85:13211337
at a temperature of approximately 45 C for 45 days until
all the ammonia has evaporated. One of the most important
developments in mushroom growing of recent years has been
the introduction of bulk processes in composting technology.
Both phase I and phase II of the composting process can take
place in bulk in special fermentation rooms called tunnels.
When that process has been completed, the compost can be
spawned, and mycelium can develop in either the same or
another tunnel. Phase III then yields fully grown compost
ready for production (Van Griensven and Van Roestel 2004).
Cultivation of L. edodes
L. edodes is the third most cultivated edible mushroom; it
was traditionally grown on wood logs (Stamets 1995;
Chang and Hayes 1978; Chang and Miles 1989; Table 2,
Fig. 3j). This method has been replaced by artificial log
cultivation that utilizes heat-treated substrates based on
sawdust enclosed in plastic bags (bag-log cultivation).
The main advantages of this method are the short time to
complete a crop cycle and the higher yields (Snchez
2004). The strain, substrate composition, and length of
incubation are important parameters for shiitake production
on synthetic substrates in bag-log cultivation (Zadrazil
1993; Kalberer 1995; Snchez 2004). Recently, mushroom
culture has moved toward diversification, and the culture of
other mushrooms has been reported.
Cultivation of Auricularia sp.
Auricularia auricula and Auricularia polytricha commonly
are produced on a medium consisting of sawdust, cotton-
seed hulls, bran, and other cereal grains or on natural logs
of broad-leaf trees (Table 2). For synthetic medium
production, the substrate may be composted for up to
5 days or used directly after mixing. The mixed substrate is
sterilized for 60 min at 121 C. Composted substrate is
prepared by mixing and watering of ingredients. The
inoculation and fructification are carried out as previously
reported (Royse 2007).
Cultivation of Flammulina velutipes
Japan is the main producer of F. velutipes (Furukawa 1987).
Production of F. velutipes in Japan is based on a substrate
contained in polypropylene bottles. Substrates (primarily
sawdust and rice bran) are mechanically mixed and filled
into heat-resistant bottles, sterilized (4 h at 95 C and 1 h at
121 C), mechanically inoculated, and incubated for 25 days
at 20 C. To further improve quality during fruiting,
temperatures are lowered to 3 C to 8 C until harvest. As
the mushrooms begin to elongate above the lip of the bottle,
a plastic collar is placed around the neck and secured with a
1329
Velcro strip. This collar serves to hold the mushrooms in
place so that they are long and straight. When the
mushrooms are 13 to 14 cm long, the collars are removed,
and the mushrooms are pulled as a bunch from the substrate
(Royse 2007).
Cultivation of Grifola frondosa
Production of G. frondosa is usually on a substrate
contained in polypropylene bottles or bags. A common
substrate used for production is composed of sawdust
supplemented with rice bran or wheat bran. It has also been
produced on a substrate consisting of oak sawdust, wheat
bran, millet, and rye. This formula gave the highest yields,
best quality, and shortest crop cycle time (12 weeks). For
bottle production, the containers are filled with moistened
substrate and sterilized or pasteurized prior to inoculation.
For production in bags, the moistened substrate is filled into
microfiltered polypropylene bags and sterilized. After
cooling (16 to 20 h), the substrate is inoculated, and the
bags are heat-sealed and shaken to uniformly distribute the
spawn throughout the substrate. Spawn runs last about 30
to 60 days depending on strain and substrate formulation.
After primordia formation, two holes usually are cut in the
bags exposing the developing primordia that tend to
develop around the outside perimeter of the substrate
surface. The top of the bag is then folded over, exposing
only the developing primordia to the fruiting environment
(Royse 2007).
Cultivation of Hypsizygus marmoreus
The Japanese are the main producers and consumers of H.
marmoreus. It is usually produced in polypropylene bottles
contained in plastic trays. After the completion of vegetative
mycelial growth, bottle lids are removed, and the colonized
substrate is subjected to environmental conditions known to
stimulate fruiting. When the mushrooms are mature, the entire
cluster of fruiting bodies is removed from the bottles. The
mushrooms are packaged by placing an entire cluster (or
multiple clusters) into each over-wrapped package. Only one
flush of mushrooms is harvested prior to mechanical removal
of the spent substrate from the bottles. The bottles then are
refilled with fresh substrate, and the process is repeated
(Royse 2007).
Cultivation of Pholiota nameko
Preparation of the medium for P. nameko cultivation is similar
to that for F. velutipes except that a higher moisture content
of the substrate is desirable. A substrate of broad-leaf tree
sawdust is preferred, but research has shown that sawdust
from conifers such Pinus spp. and Cryptomeria japonica are
1330
suitable for growth. Rice bran usually is added as a
supplement for conifer sawdust and 10% for broad-leaf
sawdust (Royse 2007).
Cultivation of Tremella fuciformis
This mushroom can be cultivated on natural logs or on a
medium (Quimio et al. 1990). Cultivation techniques used
to produce the mushroom on natural logs is similar to that
used for shiitake production. In recent years, most
production of T. fuciformis has been on a substrate using
a mixed culture inoculum technique first developed in
Fujian, China (Huang 1982). The mixed culture technique
involves the use of helper mycelium of Hypoxylon
archeri, an ascomycete commonly associated in nature
with decaying wood. H. archeri increases the ability of
T. fuciformis to digest the substrate thereby increasing
mushroom yields. Exploitation of this mycelial association
is accomplished through use of dual cultures to make
mother spawn (Quimio 1997).
Cultivation of Volvariella sp.
This mushroom is well suited for cultivation in the tropics
because of its requirement for higher production temperatures.
It can be grown on non-pasteurized substrate which is more
desirable for low input agricultural practices. In recent years,
cotton wastes (discarded after sorting in textile mills) have
become popular as substrates for straw mushroom production
(Chang 1982). Cotton wastes give higher and more stable
biological efficiencies (30% to 45%) and earlier fructification
(4 days after spawning) and harvesting (first 9 days after
spawning) than that obtained using straw as a substrate. The
culture of other mushrooms has also been reported (Table 2).
Improvement of strains
The strain used in the culture is crucial for success of
mushroom production and marketing. A strain with a high
ability to invade the substrate and to fruit diminishes time
of incubation and enhances productivity. The consistency
and texture of the mushroom are crucial to diminish losses
during packing. The first A. bisporus hybrid was obtained
by breeding about 25 years ago. However, most of the
strains currently used are similar to the first hybrid obtained
(Sonnenberg 2000; Kerrigan 2000; Snchez 2004). The
increase in mushroom production has been the result of
more specialized studies carried out by several international
institutions in different areas of mushroom cultivation
(Tables 3 and 4). The use of DNA-based technology has
accelerated breeding activities and will help mushroom-
breeding programs (Stoop and Mooibroek 1999). An
Appl Microbiol Biotechnol (2010) 85:13211337
important advance in the development of techniques for
breeding is based on the development and detection of
genetic markers (Sonnenberg 2000). Research on the
molecular basis of mating-type genes has been very
important for the development of strains with high yields,
resistance to bacterial diseases (Oliver and Delmas 1987;
Moquet et al. 1999), viral diseases (Sonnenberg et al. 1995;
Harmsen et al. 1991), and fungal diseases (Dragt et al.
1995; Kerrigan 2000).
The strain used for Pleurotus sp. cultivation is of
particular importance, since some workers develop an
allergy that is identical to mushrooms workers lung and
is associated with exposure of people to Pleurotus spp.
spores (Laborde 1995; Mori et al. 1998; Saikai et al. 2002;
Senti et al. 2000). However, similar symptoms have also
been observed with L. edodes spores (Laborde 1995; Senti
et al. 2000). This problem has increased interest in
developing sporeless mutants for breeding of sporeless
strains (Laborde 1995; Obatake et al. 2003; Tewari 2007).
Development of technology to increase productivity
To increase productivity in the mushroom culture,
it is necessary to develop and improve control and
computerized monitoring of growing rooms, automated
mushroom harvesting machines, hydroblending and pre-wet
equipment (heap turners), or to develop methods to produce
mushrooms on a non-composted substrate, and new techni-
ques of sterilization (Hawton et al. 2000). A computerized
environmental control system is an invaluable tool for
mushroom culture. The computer monitors environmental
parameters such as temperature, humidity, airflow, pressure,
and carbon dioxide and oxygen content. However, automatic
control of ammonia levels and moisture in the casing soil is
still not widely applied. More than two decades ago, the first
climate computers were introduced to Dutch mushroom
growing and now are widely used in the industry (Lamber
2000). Climate control in production plants allows monitoring
and control of several mushroom production rooms with a
minimum human input. Computerized environmental control
system allows the grower to look and adjust the environmental
conditions of the plant by using remote access (modem)
capabilities (Walker 1996). There are several companies
which provide different environmental control systems, e.g.,
DuraFlex, Jaybird Manufacturing, Agricultural Engineering,
etc. Picking and packing (Fig. 3k) constitute the most
expensive part of the overall production process. In the last
20 years, several studies have been carried out to develop
automated mushroom harvesting machines. Some companies
offer different automated harvesting systems, which include
small, modular semiautomatic picking aids and the fully
automated, robotic systems designed for line picking (Kensal
Appl Microbiol Biotechnol (2010) 85:13211337 1331

Table 4 International institutions that study different areas of mushrooms cultivation and their expected output

Area of study in mushrooms production Expected output International

institutions

Germplasm and genetic improvement

Survey, collection, and identification of wild relatives Assessment of biodiversity and conservation of germ plasm RBG, MI, CMI
and new species
Culture maintenance Conservation of germ plasm CMI, ATCC
Genetic improvement Increased productivity, improved quality, disease and pest HRI, PSU, MES
resistance
Improved production systems
State-of-the art composting technology Increased productivity and reduced environmental pollution HRI, INRA, PSU
Reduction in composting period for rural growers -do-
Newer and improved casing layer Increased productivity at lower costs INRA, PSU
Environment control in growing houses Increased productivity HRI, PSU
Development of farm designs Indigenous technology
Mechanization and automation Indigenous technology HRI, MES
Utilization of spent straw Utilization of waste and additional returns to the growers CINADCO
Improved cultivation technology Diversification of the mushroom portfolio of the country PSU, MI, CAL, CU,
Malaysia
Seed (spawn)
Development of liquid spawn technology Ease of handling and low transportation cost PSU, HRI
Improved spawn preparation and packaging technology Improved spawn quality resulting in higher yields and returns PSU, HRI
Integrated pest and disease management
Survey and preparation of area specific disease map Evolving suitable strategies against mushroom diseases
Integrated pest and disease management Increased yields and reduced residual chemicals INRA
Investigations on mushroom viruses Increase productivity INRA
Integrated pest management in mushrooms Increase productivity
Postharvest technology of mushrooms
Improvements in packaging and practices for storage Reduced postharvest losses HRI, PSU
and transport of fresh mushroom
Development of improved processing technologies for Reduced postharvest losses and avoidance of distress sale PSU
mushrooms
Development of production technologies for value- Development of value-added products
added products from mushrooms
Basic and strategic studies in mushrooms
Molecular geneticsmapping, cloning and sequencing Understanding will help manipulation for improvement in yield HRI, MES, PSU
of genes, allele mining, creation of gene libraries and quality libraries of mushrooms
Genomicsgenome sequencing, ESTs, and microarrays Basic information on the genome structure HRI, MES
Cloning and expression of useful genes of mushroom Industrial applications HRI, PSU, MES
Molecular basis of host-pathogen interaction Better disease management INRA
Role of microflora in composting Microbes for rapid composting HRI, PSU, INRA
Morphogenesis in mushrooms Domestication of newer types and increasing the yield of HRI
commercial types
Molecular mechanisms of biodegradation of Utilization of wastes for food, feed, and fuel USFPL, FSU
lignocelluloses
Studies on environmental physiology of mushrooms To develop appropriate environment for higher yields HRI, PSU

RBG Royal Botanical Gardens, UK; MI, Mori Institute, Japan; CMI Commonwealth Mycological Institute, UK; ATCC American Type Culture
Collection, USA; HRI Horticulture Research International, UK; PSU Pennsylvania State University, USA; MES Mushroom Experimental Station,
Horst, The Netherlands; INRA National Institute of Agronomy Research, France; CINADCO Centre for International Agricultural Development
Cooperation, Israel; CAL College of Agriculture Laguna, Philippines. CU Chinese University, Shatin, Hong Kong; USFPL US Forest Products
Laboratory, USA; FSU Fredrick Schiller University, Germany
Source: Tewari 2007; http://www.nrcmushroom.com

Automation; Astell 1996). The harvesting operation includes with cylindrical mushroom sample pieces (Hiller 1994).
mushroom location, sizing, selection, and picking. The Reed et al. (1995) evaluated at the laboratory level the
mechanical properties used for the analysis of automated automated harvesting of A. bisporus by machine, and the
harvesting were obtained from compression experiments resulting pilot harvester was successfully tested on a
1332
commercial mushrooms farm. The apparatus combines
several handling systems and mechatronic technologies.
Mushrooms are located and sized using image analysis and a
monochromatic vision system. An expert selection algorithm
then decides the order in which they should be picked and
what picking action (bend, twist, or both) should be used
(Tillett and Batchelor 1991; Reed et al. 1997). One of a pair
of suction cup mechanisms attached to the single head of a
Cartesian robot is then deployed; it can delicately detach
individual mushrooms and place them gently into a specially
designed compliant finger conveyer. After high speed
trimming, a gripper mechanism is finally used to remove
mushrooms from the conveyor into packs at the side of the
machine (Reed et al. 1995, 2001).
The pre-wet heap machine was designed to achieve a quick
homogeneous mix of water and the raw materials used in
phase 1 of the culture of A. bisporus to improve efficiency of
the composting process by reducing the amount of time
required to achieve the same or better quality compost. Some
benefits of using pre-wet compost turners include improving
compost quality, homogenous blending of up to 200 t per
hour, better odor control, less water run-off, raw material
savings, reducing phase 1 time by over 50%, reducing
operating and capital costs, increasing production yields,
crop uniformity, and profit potential. Design of new
equipment to improve productivity is continuously in
development. Recently, a new tunnel/bunker filling system
has been developed, which consists of filling the bunkers or
tunnels using overhead layering techniques, but only from
one end of the bunker/tunnel and not through holes in the
tunnel roof, with the use of overhead and out of sight
conveyors/elevators (Traymater machinery).
New methods to produce and increase the productivity of a
wide variety of exotic mushrooms have been developed. The
mycocell system (Mycocell Technologies, UK) is a method
based on microwave sterilization of pre-packaged substrate to
which the spawn and other nutrients (e.g., Ca2SO4) can be
added. In this case, radiation used in the treatment of the
substrates modifies the cellulose and increases ease of
breakdown by the mycelium. In addition, several nutrients
can be added and mixed thoroughly, there are no risks of
substrate contamination, and the colonization of the substrate
by the mycelium considerably increases. The commercial
benefits of this production system include lower cost, low
energy requirements, automation, low labor demand, lack of
downtime, light and cheap transport, low contamination risk,
and long shelf life. The mycocell system has allowed also the
successful culture of exotic mushrooms such as L. edodes,
P. ostreatus, Pleurotus pulmonarius, Pleurotus eryngii,
Pleurotus djamor, Pleurotus cystidiosus, Pholiota sp.,
Hypsizygus sp., F. velutipes, Agrocybe aegerita, Ganoderma
lucidum, Psilocybe sp., G. frondosa, Hericium sp., and
Auricularia (Hawton et al. 2000; Table 5).
Appl Microbiol Biotechnol (2010) 85:13211337
A technology called variable frequency speed control for
alternating current motors (Keljik 1995) may be useful for
mushroom farms and benefit the industry by improved
control of air velocity in the growing environment, resulting
in lower electricity consumption (Lomax 1989, 1992;
Snchez 2004). Currently, there are several specialized
institutes that study different frontier areas to improve
mushroom cultivation (Tables 3 and 4).
Additional benefits of mushrooms culture
Several studies have shown that spent SMS can be used for
mushroom re-cultivation, e.g., the use of SMS enriched
with cotton seed meal and with soya meal for Agaricus
cultivation, the use of spent Pleurotus substrate for the King
Stropharia cultivation (Poppe 1995), the use of spent
Agaricus compost added with cotton waste for Volvariella
production (Oei 1991), and the use of spent Pleurotus eous
straw for the cultivation of several species of this fungus
(Siddhant and Singh 2009). Chang and Miles (1989)
studied the use of spent Volvariella substrate for the
production of Pleurotus sajor-caju. Utilization of spent
straw generated after the cultivation of G. lucidum and
F. velutipes has been recommended for the production of
other mushrooms, such as A. bisporus or Coprinus comatus
(Xiao 1998). A. bisporus spent substrate has also been used
for the cultivation of Volvariella (Poppe 2000). SMS can
also be used as casing material for the production of
Agaricus (Nair and Brandley 1981; Shandilya 1989; Singh
et al. 1992, 2000). It has also been used as animal feed,
since its degradation by the mushroom can improve its
nutritional quality (Jalc et al. 1996a, b; Adamovic et al.
1998; Daz-Godnez and Snchez 2002) and digestibility by
the ruminant (Capelari and Zadrazil 1997; Daz-Godnez
and Snchez 2002). Daz-Godnez and Snchez (2002)
found that addition of maize straw generated after mushroom
cultivation to the diets of sheep increased the weight gain of
the sheep and the efficiency of feed conversion of the
straw. SMS is also a beneficial product for enrichment of
soils, restoring areas that have been destroyed through
development, deforestation, or environmental contamination.
Some studies have been done on the use of SMS in vegetable
and flower greenhouses (Lohr et al. 1984; Verdonck 1984;
Steffen et al. 1994, 1995; Szmidt 1994; Schting and Grabbe
1995; Celikel and Tuncay 1999), in field vegetable and fruits
crop (Male 1981; Pill et al. 1993; Ranganathan and
Selvaseelan 1997; Stewart et al. 1998; Batista et al. 2000;
Delver and Wertheim 1988; AntSaoir et al. 2000), in nursery
and landscape gardening (Chong and Wickware 1989;
Chong and Rinker 1994; Chong 1999), in soil amendment
as organic manure or vermi-compost (Stewart et al. 1998;
Tajbakhsh et al. 2008), and biogas production (Balan et al.
Appl Microbiol Biotechnol (2010) 85:13211337 1333

Table 5 Specialized institutes that study mushroom cultivation in different frontier areas

Frontier areas Institute

Genetic improvement and transgenic technologies MES, HRI, PSU

Identification of mushrooms CMI
DNA technologies HRI, Centre for Plant Biotechnology, Ontario, Canada
Compost technology MES, PSU, Queen's University of Belfast, Ireland
Pest and disease management INRA, PSU
Cultivation of specialty mushrooms INRA, PSU
Quality and packaging technologies and many others frontier areas PSU
Computer application in mushrooms University of Rhodes Island, Kingston, USA

Source: Tewari (2007); http://www.nrcmushroom.com

2008; Pathak et al. 2009). Currently, there are some
industries that manufacture and sell different kind of
compost based on SMS (http://www.nutrasoils.com; Meijer
2009; American Mushroom Institute 2006; http://www.
laurelvalleysoils.com).
The potential of SMS to degrade organopollutants
and its importance in environmental bioremediation has
been reported (Kuo and Regan 1998; Eggen 1999;
Xawek et al. 2003; Webb et al. 2001; Lau et al. 2003;
Law et al. 2003).

Table 6 Some diseases that attack Pleurotus ostreatus and Agaricus bisporus cultivation

Affected Infestation Type of Diseases Symptoms Reference

mushroom (organism) organism

Agaricus Cladobotryum Fungi Cobweb, mildew White to pink cobweb-like fluffy mold Western Committee on Plant Disease
bisporus dendroides (2004)
Agaricus Mycogone sp. Fungi Wet bubble/white Dense white growth on gills Western Committee on Plant Disease
bisporus mold (2004); Tanovi et al. 2009
Agaricus Mycogone Fungi Wet bubble of fungi Boiko et al. 2009; Tanovi et al. 2009
bisporus perniciosa
Magn.
Agaricus Trichoderma sp. Fungi Green mold Dark green mold patches on casing Western Committee on Plant Disease
bisporus spreading to lesions on stems (2004)
Pleurotus Velzquez-Cedeo et al. 2004
ostreatus
Agaricus Verticillium sp. Fungi Dry bubble/brown Brown irregular pitted areas on stems and Western Committee on Plant Disease
bisporus fungicola spot caps. Distortion and splitting. Severe (2004); Boiko et al. 2009
rotting, blotch, and necroses of caps and
stems
Agaricus Lycoriella sp., Insect Sciarid flies Destroy pins of developing mushrooms, and http://www.mushroomcompany.com/
bisporus L. ingenua burrow or tunnel into the stems and caps of resources/pests/index.shtml, 2009;
maturing mushrooms Jess and Schweizer 2009
Agaricus Megasellia Insect Phorid flies Cause less mushroom damage than sciarid http://www.mushroomcompany.com/
bisporus halterata flies resources/pests/index.shtml, 2009;
Smith et al. 2006
Agaricus Aphelenchoides Nematode Eelworms, Degeneration of the mushroom mycelium in http://www.mushroomcompany.com/
bisporus composticola Cephalothecium the compost resources/pests/index.shtml, 2009;
disease Gitanjali 2005
Agaricus Pseudomonas Brown blotch Sunken, dark brown lesions http://www.mushroomcompany.com/
bisporus, Tolaasii resources/pests/index.shtml , 2009
Pleurotus Pseudomonas Blotch disease Mild dark purple to light brown discoloration Godfrey et al. 2001
ostreatus, reactants and a slight surface depression that
becomes darker with age
and other Pseudomonas Ginger blotch Pale yellowish red discoloration that
mushrooms gingeri develops into a reddish ginger-colored pale
discoloration
Agaricus Putative virus X Virus Virus X Slightly imperfectly formed cap structure Gaze 2001
bisporus
Agaricus Bacilliform Virus Severe rotting, blotch and necroses of caps Boiko et al. 2009; Revill et al. 1998
bisporus viruses and stems
1334
Diseases presented during the mushrooms cultivation
In mushrooms production, a composted substrate increases
mushroom yield and reduce infestation by insects, fungi,
bacteria, and viruses. However, under vulnerable mush-
room growth conditions, several pests can attack the
mushrooms cultivation (Table 6).
Troubleshooting in mushroom cultivation
During cultivation of mushrooms, several problems occur
(Table 3); however, following of mushroom cultivation
carefully can prevent them. Prevention is better than
solving such problems (Table 6; FAO 2001).
Conclusions
Edible mushrooms have been eaten and appreciated for their
flavor, economical and ecological values, and medicinal
properties. They are able to grow under different climatic
conditions on cheap, readily available waste materials. These
mushrooms are a clear example of how low-value waste,
which is produced primarily through the activities of the
agricultural, forest, and food-processing industries, can be
converted to a higher value material useful to mankind.
Recently, mushroom culture has moved toward diversification,
and the culture of additional mushrooms has been reported. For
many reasons, the fungal Pleurotus genus has been intensely
studied and cultivated in many different parts of the world.
Particularly, P. ostreatus, also known as oyster mushroom,
hiratake, shimeji, or houbitake, requires shorter growth
time when compared to other edible mushrooms. Also, the
substrates used for its cultivation do not require sterilization,
only pasteurization which is less expensive. This mushroom
demands few environmental controls for cultivation, and its
fruiting bodies are not often attacked by diseases and pests,
and it can be cultivated in a simple and cheap way. Another
advantage of growing oyster mushrooms is that a high
percentage of the substrate is converted to fruiting bodies,
increasing profitability as compared to other mushrooms,
making P. ostreatus an excellent choice for mushroom
cultivation.
Acknowledgement I am grateful to Dr. A. L. Demain for critical
reading of the manuscript.
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