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Appl Microbiol Biotechnol (2010) 85:13211337

DOI 10.1007/s00253-009-2343-7


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

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

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

H2O2 generation by aryl-alcohol oxidase in cyclic redox

reactions involving also aryl-alcohol dehydrogenases.

Phenoxy radicals from C4-ether breakdown (b) can

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


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









redox cycling reactions involving quinone reductases,


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

of the ferric iron present in wood (n), either by superoxide

cation radical or directly by the semiquinone radicals, and its


reoxidation with concomitant reduction of H2O2 to hydroxyl

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



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

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

enzymes described above.

Poppe (2000) reported that there are about 200 kinds of
waste in which edible mushrooms can be produced.

However, mushroom production generates an enormous

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








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






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

2005; WBWDI 2007; Fig. 2).



Mushroom cultivation





Mushroom culture involves several different operations, each

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





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



o j HC

H 2O 2 a
Fe g H2COH
n AAO?

d HC
+* b
O-R O-R O c

ll O
ll OH
OCH3 ll OCH 3 k
QR l

Source: Martnez et al. 2005

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

Scientific name English name Japonese name Substrate Reference

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

MAYA ethnobotanicals (2009)
b, 2009
Appl Microbiol Biotechnol (2010) 85:13211337
Appl Microbiol Biotechnol (2010) 85:13211337 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

Mushroom production The mushroom begins to form

around the edges of bag perforations. The bags are
maintained under optimal temperature, moisture and other
Fig. 2 Top mushrooms cultivation countries conditions for mycelium growth, and the conditions that
favor fruiting. Mushroom shelves and suspended systems
are the main systems used for Pleurotus cultivation
Substrate preparation The substrate is milled to a length of (Fig. 3h). Studies on the mushroom cultivation on the use
about 2 to 6 cm. In other substrates, the chopping is not of different strains, different lignocellulosic substrates,
required. One of the most common substrates used for different types of spawn, moisture, physicochemical con-
modern mushrooms is a mixture. This mixture of cotton- ditions, etc. are important for the cultivation productivity of
seed hulls and wheat straw has a higher water holding each particular mushroom (Mandeel et al. 2005; Saavedra
capacity than cottonseed hulls used alone. Pasteurization, et al. 2006; Kirbag and Akyuz 2008; Onuoha et al. 2009).
used on some commercial mushroom farms, is carried out
by filling the ingredients into revolving mixers, water is Harvest The mushrooms are harvested from the substrate
added to the desired level, and live steam is injected into approximately 3 to 4 weeks after spawning depending on
the mixer while it is in operation (Royse 2007). strain, amount of supplement used, and temperature of
spawn run.
Inoculation After completion of pasteurization (60 C for 1
to 2 h), the substrate is cooled and spawned with the desired
strain. Cultivation of A. bisporus

At time of spawning, a delayed release supplement (rates Agaricus sp. is also known as himematsutake. It is the
of 3% to 10% of dry substrate weight) may be added to principal cultivated mushroom worldwide, and its substrate
increase yield and size of the mushroom (Royse et al. 1991; is the most complex culture medium used for edible
Royse and Zaki 1991). Use of supplements, however, may mushroom production (Fig. 3i). Several studies on the
cause overheating of the substrate if growers are not able to development of this mushroom have improved its cultiva-
anticipate and control air temperatures to maintain a steady tion (Stoop and Mooibroek 1999; Kothe 2001; Bechara et
substrate temperature. al. 2004, 2008; Nogueira de Andrade et al. 2008; Kapoor et
al. 2009). The substrate for the cultivation of this
Spawning and spawn rate Growers have sought, in the mushroom is horse manure compost, which consists of a
past, to optimize the amount of spawn used to inoculate mixture of horse manure, some broiler chicken manure, and
their substrate. Increasing the amount of spawn used (up to water to which gypsum is added for structural stability and
5% of the wet weight of the substrate) has resulted in for stabilizing pH. Alternatively, a synthetic compost can be
increased yields (Royse 2002). Increasing spawn rates from made which is based on wheat straw and broiler chicken
1.25% to 5% may result in increases of nearly 50%. Yield manure. When carried out in the open air, phase I of
increases may be due to several factors. First, the increased composting is done in long narrow stacks between 1.5 and
level of nutrient available in higher levels of spawn used 2.0 m in width and height (wind-rows). The stacks are
would provide more energy for mycelial growth and turned twice a week in order to obtain uniform degradation
development. Second, more inoculum points, available of the mixture. At the end of the process, usually 78 days
from increased spawn levels, would provide faster substrate after setting up the stacks of 1 t of horse manure, a yield of
colonization and, thus, more rapid completion of the 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

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
Poor spread of mycelium, bad smell, spots, 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
and mites 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
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
Mycelium grows but fails to produce Substrate formula is not suitable Adjust the formula; check pH, sawdust, additives, etc
mushrooms 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
Mushrooms form, but abort or delay Primodia and growth condition of fruiting body are not good Check temperature and humidity. Open or close doors and window to adjust accordingly
mushrooming enough
There is contamination such as mold, bacteria, insects, worms, and Check hygiene, adjust environment of light, temperature, humidity, and ventilation. In
Table 3 (continued)

Problem Cause Solutions

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

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Appl Microbiol Biotechnol (2010) 85:13211337 1329

at a temperature of approximately 45 C for 45 days until Velcro strip. This collar serves to hold the mushrooms in
all the ammonia has evaporated. One of the most important place so that they are long and straight. When the
developments in mushroom growing of recent years has been mushrooms are 13 to 14 cm long, the collars are removed,
the introduction of bulk processes in composting technology. and the mushrooms are pulled as a bunch from the substrate
Both phase I and phase II of the composting process can take (Royse 2007).
place in bulk in special fermentation rooms called tunnels.
When that process has been completed, the compost can be Cultivation of Grifola frondosa
spawned, and mycelium can develop in either the same or
another tunnel. Phase III then yields fully grown compost Production of G. frondosa is usually on a substrate
ready for production (Van Griensven and Van Roestel 2004). contained in polypropylene bottles or bags. A common
substrate used for production is composed of sawdust
Cultivation of L. edodes supplemented with rice bran or wheat bran. It has also been
produced on a substrate consisting of oak sawdust, wheat
L. edodes is the third most cultivated edible mushroom; it bran, millet, and rye. This formula gave the highest yields,
was traditionally grown on wood logs (Stamets 1995; best quality, and shortest crop cycle time (12 weeks). For
Chang and Hayes 1978; Chang and Miles 1989; Table 2, bottle production, the containers are filled with moistened
Fig. 3j). This method has been replaced by artificial log substrate and sterilized or pasteurized prior to inoculation.
cultivation that utilizes heat-treated substrates based on For production in bags, the moistened substrate is filled into
sawdust enclosed in plastic bags (bag-log cultivation). microfiltered polypropylene bags and sterilized. After
The main advantages of this method are the short time to cooling (16 to 20 h), the substrate is inoculated, and the
complete a crop cycle and the higher yields (Snchez bags are heat-sealed and shaken to uniformly distribute the
2004). The strain, substrate composition, and length of spawn throughout the substrate. Spawn runs last about 30
incubation are important parameters for shiitake production to 60 days depending on strain and substrate formulation.
on synthetic substrates in bag-log cultivation (Zadrazil After primordia formation, two holes usually are cut in the
1993; Kalberer 1995; Snchez 2004). Recently, mushroom bags exposing the developing primordia that tend to
culture has moved toward diversification, and the culture of develop around the outside perimeter of the substrate
other mushrooms has been reported. surface. The top of the bag is then folded over, exposing
only the developing primordia to the fruiting environment
Cultivation of Auricularia sp. (Royse 2007).

Auricularia auricula and Auricularia polytricha commonly Cultivation of Hypsizygus marmoreus

are produced on a medium consisting of sawdust, cotton-
seed hulls, bran, and other cereal grains or on natural logs The Japanese are the main producers and consumers of H.
of broad-leaf trees (Table 2). For synthetic medium marmoreus. It is usually produced in polypropylene bottles
production, the substrate may be composted for up to contained in plastic trays. After the completion of vegetative
5 days or used directly after mixing. The mixed substrate is mycelial growth, bottle lids are removed, and the colonized
sterilized for 60 min at 121 C. Composted substrate is substrate is subjected to environmental conditions known to
prepared by mixing and watering of ingredients. The stimulate fruiting. When the mushrooms are mature, the entire
inoculation and fructification are carried out as previously cluster of fruiting bodies is removed from the bottles. The
reported (Royse 2007). mushrooms are packaged by placing an entire cluster (or
multiple clusters) into each over-wrapped package. Only one
Cultivation of Flammulina velutipes flush of mushrooms is harvested prior to mechanical removal
of the spent substrate from the bottles. The bottles then are
Japan is the main producer of F. velutipes (Furukawa 1987). refilled with fresh substrate, and the process is repeated
Production of F. velutipes in Japan is based on a substrate (Royse 2007).
contained in polypropylene bottles. Substrates (primarily
sawdust and rice bran) are mechanically mixed and filled Cultivation of Pholiota nameko
into heat-resistant bottles, sterilized (4 h at 95 C and 1 h at
121 C), mechanically inoculated, and incubated for 25 days Preparation of the medium for P. nameko cultivation is similar
at 20 C. To further improve quality during fruiting, to that for F. velutipes except that a higher moisture content
temperatures are lowered to 3 C to 8 C until harvest. As of the substrate is desirable. A substrate of broad-leaf tree
the mushrooms begin to elongate above the lip of the bottle, sawdust is preferred, but research has shown that sawdust
a plastic collar is placed around the neck and secured with a from conifers such Pinus spp. and Cryptomeria japonica are
1330 Appl Microbiol Biotechnol (2010) 85:13211337

suitable for growth. Rice bran usually is added as a important advance in the development of techniques for
supplement for conifer sawdust and 10% for broad-leaf breeding is based on the development and detection of
sawdust (Royse 2007). genetic markers (Sonnenberg 2000). Research on the
molecular basis of mating-type genes has been very
Cultivation of Tremella fuciformis important for the development of strains with high yields,
resistance to bacterial diseases (Oliver and Delmas 1987;
This mushroom can be cultivated on natural logs or on a Moquet et al. 1999), viral diseases (Sonnenberg et al. 1995;
medium (Quimio et al. 1990). Cultivation techniques used Harmsen et al. 1991), and fungal diseases (Dragt et al.
to produce the mushroom on natural logs is similar to that 1995; Kerrigan 2000).
used for shiitake production. In recent years, most The strain used for Pleurotus sp. cultivation is of
production of T. fuciformis has been on a substrate using particular importance, since some workers develop an
a mixed culture inoculum technique first developed in allergy that is identical to mushrooms workers lung and
Fujian, China (Huang 1982). The mixed culture technique is associated with exposure of people to Pleurotus spp.
involves the use of helper mycelium of Hypoxylon spores (Laborde 1995; Mori et al. 1998; Saikai et al. 2002;
archeri, an ascomycete commonly associated in nature Senti et al. 2000). However, similar symptoms have also
with decaying wood. H. archeri increases the ability of been observed with L. edodes spores (Laborde 1995; Senti
T. fuciformis to digest the substrate thereby increasing et al. 2000). This problem has increased interest in
mushroom yields. Exploitation of this mycelial association developing sporeless mutants for breeding of sporeless
is accomplished through use of dual cultures to make strains (Laborde 1995; Obatake et al. 2003; Tewari 2007).
mother spawn (Quimio 1997).

Cultivation of Volvariella sp. Development of technology to increase productivity

This mushroom is well suited for cultivation in the tropics To increase productivity in the mushroom culture,
because of its requirement for higher production temperatures. it is necessary to develop and improve control and
It can be grown on non-pasteurized substrate which is more computerized monitoring of growing rooms, automated
desirable for low input agricultural practices. In recent years, mushroom harvesting machines, hydroblending and pre-wet
cotton wastes (discarded after sorting in textile mills) have equipment (heap turners), or to develop methods to produce
become popular as substrates for straw mushroom production mushrooms on a non-composted substrate, and new techni-
(Chang 1982). Cotton wastes give higher and more stable ques of sterilization (Hawton et al. 2000). A computerized
biological efficiencies (30% to 45%) and earlier fructification environmental control system is an invaluable tool for
(4 days after spawning) and harvesting (first 9 days after mushroom culture. The computer monitors environmental
spawning) than that obtained using straw as a substrate. The parameters such as temperature, humidity, airflow, pressure,
culture of other mushrooms has also been reported (Table 2). 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
Improvement of strains climate computers were introduced to Dutch mushroom
growing and now are widely used in the industry (Lamber
The strain used in the culture is crucial for success of 2000). Climate control in production plants allows monitoring
mushroom production and marketing. A strain with a high and control of several mushroom production rooms with a
ability to invade the substrate and to fruit diminishes time minimum human input. Computerized environmental control
of incubation and enhances productivity. The consistency system allows the grower to look and adjust the environmental
and texture of the mushroom are crucial to diminish losses conditions of the plant by using remote access (modem)
during packing. The first A. bisporus hybrid was obtained capabilities (Walker 1996). There are several companies
by breeding about 25 years ago. However, most of the which provide different environmental control systems, e.g.,
strains currently used are similar to the first hybrid obtained DuraFlex, Jaybird Manufacturing, Agricultural Engineering,
(Sonnenberg 2000; Kerrigan 2000; Snchez 2004). The etc. Picking and packing (Fig. 3k) constitute the most
increase in mushroom production has been the result of expensive part of the overall production process. In the last
more specialized studies carried out by several international 20 years, several studies have been carried out to develop
institutions in different areas of mushroom cultivation automated mushroom harvesting machines. Some companies
(Tables 3 and 4). The use of DNA-based technology has offer different automated harvesting systems, which include
accelerated breeding activities and will help mushroom- small, modular semiautomatic picking aids and the fully
breeding programs (Stoop and Mooibroek 1999). An 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


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
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,
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
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
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;

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 Appl Microbiol Biotechnol (2010) 85:13211337

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

2008; Pathak et al. 2009). Currently, there are some The potential of SMS to degrade organopollutants
industries that manufacture and sell different kind of and its importance in environmental bioremediation has
compost based on SMS (; Meijer been reported (Kuo and Regan 1998; Eggen 1999;
2009; American Mushroom Institute 2006; http://www. 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
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
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
Agaricus Lycoriella sp., Insect Sciarid flies Destroy pins of developing mushrooms, and
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
bisporus halterata flies resources/pests/index.shtml, 2009;
Smith et al. 2006
Agaricus Aphelenchoides Nematode Eelworms, Degeneration of the mushroom mycelium in
bisporus composticola Cephalothecium the compost resources/pests/index.shtml, 2009;
disease Gitanjali 2005
Agaricus Pseudomonas Brown blotch Sunken, dark brown lesions
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
Agaricus Putative virus X Virus Virus X Slightly imperfectly formed cap structure Gaze 2001
Agaricus Bacilliform Virus Severe rotting, blotch and necroses of caps Boiko et al. 2009; Revill et al. 1998
bisporus viruses and stems
1334 Appl Microbiol Biotechnol (2010) 85:13211337

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