MINI-REVIEW

Cultivation of Pleurotus ostreatus and otheredible mushrooms

Carmen Sánchez

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 cultivatededible mushroom worldwide after Agaricus bisporus. It haseconomic and ecological values and medicinal properties.Mushroom culture has moved toward diversification withthe production of other mushrooms. Edible mushroomsare able to colonize and degrade a large variety oflignocellulosic substrates and other wastes which areproduced primarily through the activities of the agricultural,forest, and food-processing industries. Particularly,P. ostreatus requires a shorter growth time in comparisonto other edible mushrooms. The substrate used for theircultivation does not require sterilization, only pasteurization,which is less expensive. Growing oyster mushrooms convert ahigh percentage of the substrate to fruiting bodies, increasingprofitability. P. ostreatus demands few environmentalcontrols, and their fruiting bodies are not often attacked bydiseases and pests, and they can be cultivated in a simple andcheap way. All this makes P. ostreatus cultivation anexcellent alternative for production of mushrooms whencompared to other mushrooms.

Keywords Pleurotus ostreatus . Mushroom cultivation .

Edible mushrooms

Introduction

Cultivation of edible mushrooms is a biotechnologicalprocess for lignocellulosic organic waste recycling. It might

be the only current process that combines the production ofprotein-rich food with the reduction of environmentalpollution (Beetz and Kustudia 2004). The production ofmushrooms is regarded as the second most importantcommercial microbial technology next to yeast (Pathak et al.2009). Mushrooms have been eaten and appreciated for theirflavor, economical and ecological values, and medicinalproperties for many years. In general, mushrooms contain90% water and 10% dry matter (Morais et al. 2000; Sánchez2004). They have chemical composition which are attractivefrom the nutritional point of view (Gbolagade et al. 2006;Dundar et al. 2008; Table 1). Their nutritional value can becompared to those of eggs, milk, and meat (Oei 2003).Mushrooms also contain vitamins and an abundance ofessential amino acids (Sánchez 2004). The total energeticvalue of mushroom caps is between 250 and 350 cal/kg offresh mushrooms (Oliver and Delmas 1987; Laborde 1995).Some mushroom can be cultivated easily and have significantworldwide markets. Over 200 species have been collectedfrom the wild and used for various traditional medicalpurposes, mainly in the Far East (Sánchez 2004). Roughly300 mushrooms species are edible, but only 30 have beendomesticated and ten grown commercially (Barny 2009).The principal cultivated mushroom worldwide is Agaricusbisporus followed by Pleurotus sp. (Rühl et al. 2008),Lentinula edodes, and other mushrooms that have already animportant place in the market.

Mushrooms have the ability to degrade several lignocellu-losic substrates (Fig. 1; Sánchez 2009) and can be producedon natural materials from agriculture, woodland, animalhusbandry, and manufacturing industries (Table 2). As it isshown in Fig. 1, laccases or ligninolytic peroxidases (LiPand MnP) produced by white-rot fungi oxidize the ligninpolymer, thereby generating aromatic radicals (a). Theseevolve in different non-enzymatic reactions, including

C. Sánchez (*)Laboratory of Biotechnology, Research Centre for Biological Sciences,Universidad Autónoma de Tlaxcala,Apartado postal 129,Tlaxcala, Tlax CP 90000, Mexicoe-mail: sanher6@hotmail.com

Appl Microbiol Biotechnol (2010) 85:1321–1337DOI 10.1007/s00253-009-2343-7

C-4-ether breakdown (b), aromatic ring cleavage (c), Cα-Cβ breakdown (d), and demethoxylation (e). The aromaticaldehydes releases from Cα-Cβ breakdown of lignin orsynthesized de novo by fungi (f, g) are the substrate forH2O2 generation by aryl-alcohol oxidase in cyclic redoxreactions involving also aryl-alcohol dehydrogenases.Phenoxy radicals from C4-ether breakdown (b) canrepolymerize on the lignin polymer (h) if they are not firstreduced by oxidases to phenolic compounds (i). Thephenolic compounds formed can be again reoxidized bylaccases or peroxidases (j). Phenoxy radicals can also besubjected to Cα-Cβ breakdown (k), yielding ρ-quinones.Quinones from g and/or k contribute to oxygen activation inredox cycling reactions involving quinone reductases,laccases, and peroxidases (l, m). This results in reductionof the ferric iron present in wood (n), either by superoxidecation radical or directly by the semiquinone radicals, and itsreoxidation with concomitant reduction of H2O2 to hydroxylfree radical (OH.) (o). The latter is a very strong oxidizerthat can initiate the attack on lignin (p) in the initial stages ofwood decay, when the small size of pores in the still-intactcell wall prevents the penetration of ligninolytic enzymes.Then lignin degradation proceeds by oxidative attack of theenzymes described above.

Poppe (2000) reported that there are about 200 kinds ofwaste in which edible mushrooms can be produced.However, mushroom production generates an enormousamount of used “spent” substrate which might also be asource of environmental contamination. Several uses forspent mushrooms substrate (SMS) are being evaluated, andsome of them have already been established (Sánchez2004). The production of mushrooms in 2005 was (in metrictons) 1,411 in China, 382 in USA, 245 in Netherlands, 139 inFrance, 138 in Spain, 135 in Poland, 88 in Italy, 80 in Canada,77 in Ireland, and 74 in the UK (FAOSTAT 2005; USDA2005; WBWDI 2007; Fig. 2).

Mushroom cultivation

Mushroom culture involves several different operations, eachof which must be carefully performed. Substrate preparation,inoculation, incubation, and production conditions depend onthe mushroom species to be cultivated (Fig. 3). The first stageinvolves obtaining pure mycelium of the specific mushroomstrain. The mycelium can be obtained from spores (Fig. 3a),from a piece of the specific mushroom (Fig. 3b), or fromseveral germplasm providers such as American TypeCulture Collection and National Center for AgriculturalUtilization Research (Fig. 3c). To obtain inoculum, themycelium is developed on cereal grain, e.g., wheat, rye, ormillet (Fig. 3d), which is usually called the “spawn”(Fig. 3e; Chang and Hayes 1978; Chang and Miles 1989).T

able

1Nutrimentalvalueof

severaledible

mushroo

ms(m

g/10

0gdrymatter)

Ediblemushroo

mLipids

Sugars

Protein

Fiber

Ash

Ca

Mg

KNa

PMn

Fe

Cu

Zn

Auricularia

polytricha

5.05

5.35

8.90

3.45

4.95

0.75

1.45

37.40

0.35

19.85

0.29

0.67

0.12

0.06

Lentin

ussubn

udus

4.05

10.15

5.80

5.40

6.50

1.75

2.45

22.15

2.15

2.50

0.07

0.52

0.12

1.97

Lycoperdon

pusilum

7.55

14.70

24.0

4.30

6.55

5.20

3.90

28.80

2.00

14.50

0.90

0.72

0.16

1.30

Lycoperdon

giga

nteum

10.25

17.20

24.30

5.03

10.90

4.30

3.10

44.20

3.50

20.90

0.50

0.25

0.07

0.80

Pleurotus

tuber-regium

1.70

7.70

16.30

15.60

9.20

1.90

0.80

9.51

1.50

4.10

0.15

0.32

0.02

1.91

Pleurotus

florida

1.05

8.95

15.10

4.40

10.60

0.40

1.50

14.75

0.25

13.35

0.80

0.08

0.05

0.05

Psathyrella

atroum

bona

ta5.75

7.85

17.40

11.65

9.65

1.57

4.55

43.35

6.50

15.0

0.45

0.20

0.07

0.77

Schizophyllum

commune

5.50

8.75

10.30

8.30

11.60

3.90

0.55

16.80

0.65

8.10

0.30

0.10

0.08

1.20

Term

itomyces

microcarpus

8.75

14.15

27.70

12.80

12.90

3.50

3.65

58.75

2.00

24.20

0.72

0.70

0.07

2.82

Term

itomyces

globalus

6.70

14.65

32.80

10.35

15.80

5.30

6.00

48.40

1.70

31.10

0.85

0.85

0.09

3.10

Tricho

lomaloba

yensis

4.20

7.75

13.50

8.50

9.40

1.50

0.55

18.60

2.20

9.95

0.29

0.32

0.13

0.85

Volvariella

esculenta

10.95

9.85

26.05

7.0

12.65

0.80

2.45

51.45

5.90

18.75

0.44

0.42

0.11

2.50

Pleurotus

sajor-caju

1.15

–16

.75

–5.84

0.10

0.46

4.32

–1.97

0.021

0.163

0.14

0.192

Gbo

lagade

2006

;Dun

daret

al.20

08

1322 Appl Microbiol Biotechnol (2010) 85:1321–1337

The purpose of the mycelium-coated grain is to rapidlycolonize the specific bulk growing substrate (Fig. 3f). Thesuccess of mushroom production depends in great part on thequality of the “spawn”, which must be prepared under sterileconditions to diminish contamination of the substrate(Fig. 3g). Several studies have been done to improve thequality and develop new techniques for its production. Forexample, the spawn for cultivation of P. ostreatus has beenprepared in different ways: on grain, such as wheat,sorghum, and paddy (Beetz and Kustudia 2004; Nwanzeet 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; Rühl et al. 2008). For many reasons,the fungal Pleurotus genus has been intensely studied andcultivated in many different parts of the world. It is producedon a variety of lignocellulosic substrates (Table 2).

OCH 3 HC

HC

HOOH2

O

O

H2COH

C=O

H2OC

CH

OOH

OCH 3

HCOH

HOCH2 O

OCH3

HCOH

HC

HOCH2

LIGNIN

LIGNIN

LIGNIN

LACCASE

PEROXIDASE

O

R

LIGNIN

O*

LIGNIN

OH

OCH 3

H2COH

LIGNIN

-R O -R

AAD

AAO

H2O2

LACCASE PEROXIDASE

h

o

n g

m

a

j

b

c

d

k

i AAO?

f

g

HC

e

HC

LIGNIN

+*

OH*

O2-*

Fe3+

OCH 3 OCH 3

QR OH

OH C

OH

OO

H2COH

OCH3

OCH 3 OCH 3 OCH3

OCH3

HC

HC HC=O HCOH CH2OH

HC

O

HCOH

HC

H2COH

H2COH

MeOH

HC

HC

O

l l

l l

l l

l

LACCASE PEROXIDASE

Source: Martínez et al. 2005

Fig. 1 Lignin biodegradationprocess by white rot fungi

Appl Microbiol Biotechnol (2010) 85:1321–1337 1323

Tab

le2

Lignino

cellu

losicresidu

esused

fordifferentmushroo

mscultivatio

n

Scientific

name

Englishname

Japonese

name

Substrate

Reference

Aga

ricusbisporus

Button,

white,table,

portob

ello,

crim

ini,Swissbrow

n,grem

ini,

Italianbrow

nandItalianroman

brow

nmushroo

m,Champign

on

Him

ematsutake

orAgarikusutake

a

(nam

eforAga

ricusblazei)

Ricestraw,wheat

straw,ho

rsemanure

(fresh

orcompo

sted)

Beetz

andKustudia20

04;To

keret

al.20

07

Wheat

straw

andwaste

tea

Leaves

Pleurotussp

(P.ostreatus,

P.sajor-caju,P.

eryngii,

P.pulmonarius,

P.eous,P.

florida)

Oyster,king

oyster

mushroo

mHiratake

Coffeepu

lp,sawdu

st,cocoa,

peanut

and

coconutshells,cotto

nseed

hulls,Jamaica,

cassavapeels,cotto

n,sorghu

m,banana,

corn

stalks,grass,clover,woo

d

Beetz

andKustudia20

04;Raniet

al.20

08;

Shahet

al.20

04;Dabaet

al.20

08;Fan

etal.20

03;Cayetano-Catarinoand

Bernabé-G

onzález20

08;Philip

poussis

etal.20

01;Onu

ohaet

al.20

09;Sivrikaya

andPeker

1999;Abrar

etal.20

09Wastesof

rice,wheat,sawdust,cotto

nfrom

textile

indu

stry,corncobs,crushedbagasse

andmolassesfrom

sugarindustry,water

hyacinth,water

lily,

bean,wheat

straw,

leaves,oil-palm

fiber,paperandpaddy

Lentin

ulasp.(L.conn

otus,

L.edod

es)

Japanese

forestmushroo

m,Shiitake

mushroo

mblackforest,Black

oak

Shiitake

Sorgh

umstalk,

banana,coffeepu

lp,saw

dust,

cotto

nseed

hulls

bran,leaves,corncobs,

bractsof

pineapple,

cotto

nseed

meal,

peanut

meal,wheat

bran,rice

bran

and

soyb

eanmea

Raniet

al.20

08;Beetz

andKustudia20

04;

Kapoo

ret

al.20

09

Wastesfrom

padd

y,sugarcane

bagasse,

sugarcane,coffee

pulp,wheat

Volvariella

volvacea

Straw

mushroo

mor

paddystraw

mushroo

mFukurotake

Bananaleaves,cocoyam

peelings

and

oil-palm

pericarp,palm

fibers,rice

husk,

andsawdu

st

Beetz

andKustudia20

04;Belew

uand

Belew

u20

05;Obo

daiet

al.20

06;Uko

ima

etal.20

09Wastesfrom

rice,wheat,cotto

n,textile

indu

stry,water

hyacinth,andwater

lily

Tubersp.,T.

melanospo

rum

Vitt.,

T.mag

natum

Picoex

Vitt.)

Black

truffleblackperigo

rd,

piedmon

tese

Seyo

ufuyu

shuo

roro

(nam

efor

violet

truffle)

bMungb

eanhu

sks,po

tting

soil

Carluccio

2003

White

truffle

Stropharia

sp.(S.rugosoannulata

Farlow

,S.rugosoannulataFarlow

f.luteahongo)

Round

heads

Saketsubatake,kisaketsub

atake

Wastesfrom

wheat,sawdust

Beetz

andKustudia20

04

Hericium

sp.

Lion'shead

orPom

pom

Yam

abushitake

Saw

dust,corncobs,distillersgrainwaste

Beetz

andKustudia20

04

Auricularia

auricula

Jelly

ear

Kikurage

Saw

dust,sawdust-ricebran

Beetz

andKustudia20

04

Grifola

fron

dosa

Maitake,hen

ofthewoo

ds,ram

shead,

sheep'shead,clou

dmushroo

m,

dancingmushroo

m

Mai

Take,Maitake

Saw

dust,spentsawdustmatrix,

corn,barley,

sorghu

m,andrice

Beetz

andKustudia20

04;Hidenoet

al.

2007;Montoya

andVarón

2008

Flammulinasp.

Winter,velvet

stem

,go

lden,snow

puff

Eno

kitake

Saw

dust

Beetz

andKustudia20

04

Pholio

tanameko

Fat

pholiota

Num

erisugtake

Log

ssawdust-ricebran

Beetz

andKustudia20

04

Trem

ella

White

jelly

Shirokiku

rage

Log

sBeetz

andKustudia20

04

aMAYA

ethn

obotanicals(200

9)bhttp://zipcod

ezoo

.com

,20

09

1324 Appl Microbiol Biotechnol (2010) 85:1321–1337

Substrate preparation The substrate is milled to a length ofabout 2 to 6 cm. In other substrates, the chopping is notrequired. One of the most common substrates used formodern mushrooms is a mixture. This mixture of cotton-seed hulls and wheat straw has a higher water holdingcapacity than cottonseed hulls used alone. Pasteurization,used on some commercial mushroom farms, is carried outby filling the ingredients into revolving mixers, water isadded to the desired level, and live steam is injected intothe mixer while it is in operation (Royse 2007).

Inoculation After completion of pasteurization (60 °C for 1to 2 h), the substrate is cooled and spawned with the desiredstrain.

At time of spawning, a delayed release supplement (ratesof 3% to 10% of dry substrate weight) may be added toincrease yield and size of the mushroom (Royse et al. 1991;Royse and Zaki 1991). Use of supplements, however, maycause overheating of the substrate if growers are not able toanticipate and control air temperatures to maintain a steadysubstrate temperature.

Spawning and spawn rate Growers have sought, in thepast, to optimize the amount of spawn used to inoculatetheir substrate. Increasing the amount of spawn used (up to5% of the wet weight of the substrate) has resulted inincreased yields (Royse 2002). Increasing spawn rates from1.25% to 5% may result in increases of nearly 50%. Yieldincreases may be due to several factors. First, the increasedlevel of nutrient available in higher levels of spawn usedwould provide more energy for mycelial growth anddevelopment. Second, more inoculum points, availablefrom increased spawn levels, would provide faster substratecolonization and, thus, more rapid completion of the

production cycle. Finally, a more rapid spawn run wouldreduce the time non-colonized substrate is exposed tocompetitors 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 orblack perforated polyethylene bags.

Incubation The bags are incubated for 12 to 14 days at25 °C and then transferred to the production room (Royse2007).

Mushroom production The mushroom begins to formaround the edges of bag perforations. The bags aremaintained under optimal temperature, moisture and otherconditions for mycelium growth, and the conditions thatfavor fruiting. Mushroom shelves and suspended systemsare the main systems used for Pleurotus cultivation(Fig. 3h). Studies on the mushroom cultivation on the useof different strains, different lignocellulosic substrates,different types of spawn, moisture, physicochemical con-ditions, etc. are important for the cultivation productivity ofeach particular mushroom (Mandeel et al. 2005; Saavedraet al. 2006; Kirbag and Akyuz 2008; Onuoha et al. 2009).

Harvest The mushrooms are harvested from the substrateapproximately 3 to 4 weeks after spawning depending onstrain, amount of supplement used, and temperature ofspawn run.

Cultivation of A. bisporus

Agaricus sp. is also known as “himematsutake”. It is theprincipal cultivated mushroom worldwide, and its substrateis the most complex culture medium used for ediblemushroom production (Fig. 3i). Several studies on thedevelopment of this mushroom have improved its cultiva-tion (Stoop and Mooibroek 1999; Kothe 2001; Bechara etal. 2004, 2008; Nogueira de Andrade et al. 2008; Kapoor etal. 2009). The substrate for the cultivation of thismushroom is horse manure compost, which consists of amixture of horse manure, some broiler chicken manure, andwater to which gypsum is added for structural stability andfor stabilizing pH. Alternatively, a synthetic compost can bemade which is based on wheat straw and broiler chickenmanure. When carried out in the open air, phase I ofcomposting is done in long narrow stacks between 1.5 and2.0 m in width and height (wind-rows). The stacks areturned twice a week in order to obtain uniform degradationof the mixture. At the end of the process, usually 7–8 daysafter setting up the stacks of 1 t of horse manure, a yield ofjust over 1 t of compost will have been obtained. This fresh

Fig. 2 Top mushrooms cultivation countries

Appl Microbiol Biotechnol (2010) 85:1321–1337 1325

compost (phase I), which still smells strongly of ammonia,is subjected to phase II treatment before actual cultivationcan start. Phase II consists of a pasteurization processfollowed by further high-temperature fermentation. This is

done by raising the air temperature to 56 °C and keeping itthere for approximately 5 to 6 h. Compost temperature thenincreases to slightly higher values, and this is maintainedfor at least five more hours. Thereafter, the compost is kept

Source: Modified from Stamets (1995)

Fig. 3 Cultivation and harvest-ing of mushroom

1326 Appl Microbiol Biotechnol (2010) 85:1321–1337

Tab

le3

Troub

leshoo

tingin

themushroo

mscultivatio

n

Problem

Cause

Solutions

Mycelium

fails

toform

Improper

initiationstrategy

Consultparameter

ofgrow

th.Altermoisture,

temperature,lig

ht,carbon

dioxide,

etc.

Note:

Ifthesubstrateistoomoist,decrease

moisture

Chlorinated

orcontam

inated

water

Use

activ

ated

charcoal

water

filtersto

elim

inatechem

ical

contam

inantsor

anyotherways

ofsimpleor

appropriatetechnology

Bad

substrate

Check

substrate.

Spreadthesubstrateandremix

thesubstrate,

packageagain,

makesure

allraw

materialsarego

odandfresh

Note:

Itisnecessaryto

pasteurize

immediately

afterbaggingotherw

iseferm

entatio

ngas

will

slow

downtherate

ofgrow

thof

mycelium

orstop

mycelium

grow

th

Bad

pasteurizatio

nCheck

methodof

pasteurizatio

n.Release

allairandmakesure

thereiscontinuous

steam

before

startin

gpasteurizatio

nforaperiod

of3

h

Substrate

inthebagistoohotwheninoculation

Makesure

that

thesubstratebagisnottoohotbefore

inoculation

Bad

strain

orspaw

nObtainyoungerstrain

ofknow

nvitalityandhistory

Spawncontam

inated

Pasteurizeandinoculateagainwith

good

spaw

n

Forgo

tto

inoculatethebag

Makesure

toinoculate

Poo

rspread

ofmycelium,badsm

ell,spots,

andmites

Goodpasteurizatio

nbutmustdecrease

thetemperature

inthe

pasteurizatio

ncham

ber

Slowly

decrease

thetemperature

inthecham

ber.Donoto

penthecoverof

thecham

bertoo

quickly.

Check

that

thecotto

nplug

istig

htly

closed

Pasteurizationwas

tooquickand/or

thecham

berdoor

was

opened

tooqu

ickly

Inoculationprocess

Inoculatein

hygienecond

itions;cleanandwith

noairmov

ement

Toohigh

density

intheincubatio

narea,notenough

ventilatio

nto

decrease

accumulated

temperature

Spreadthesubstratebagandmakemoreairventilatio

nin

theincubatio

narea.Check

temperature

andcontrolsurroundings

tomaintainof

25to

35°C

Toohigh

carbon

dioxide

Not

morethan

5%carbon

dioxide.

Check

ventilatio

n

Hyg

iene

oftheincubatio

nho

use

Improv

ehy

gienein

theincubatio

nho

use

Mycelium

develops

inpatches.Sub

strate

isno

tevenly

prepared,

andsomepartshave

morenu

trientsthan

others

Mix

wellthesubstrate

Bacteria,

otherfungicontam

ination

Check

theprocesscausingcontam

ination.

Separatecontam

inated

bags

assoon

aspossible.

Rem

ixsubstrateseparately.Rem

akesubstratebags

andpasteurize

foralonger

time.

Follow

process

Mite

contam

ination

Immediately

separate

contam

inated

bags

andpasteurize

again.

Contin

uethenorm

alprocess

Note:

Keephygienemanagem

ent;makesure

tocleaneverything(person,

area,tools,

equipm

ent,andsurrou

ndings

during

everystep.Stopusingthearea

tocutthelifecycle

ofallcontaminantsforaperiod

ofatleast1

to2weeks.F

orseriouscontam

inationcases,

sprayarea

with

chem

icals.Use

black-lig

htwith

water

orsticky-trapto

decrease

insects

Mycelium

grow

sbutfails

toproduce

mushrooms

Sub

strate

form

ulaisno

tsuitable

Adjusttheform

ula;

checkpH

,sawdust,additiv

es,etc

Mites,mold,

virus,bacteria

andinsects

Check

pasteurizatio

nprocess,inoculation,

otherprocessesandmushroo

mho

use

managem

entforhy

giene

Inhibitedby

environm

entaltoxins

Rem

ovesource

oftoxins

Bad

strain

orShawn

Acquire

new

strains

Mushroomsform

,bu

tabortor

delay

mushrooming

Primodia

andgrow

thcondition

offruitin

gbody

arenotgood

enough

Check

temperature

andhumidity.Openor

closedoorsandwindow

toadjustaccordingly

There

iscontam

inationsuch

asmold,

bacteria,insects,worms,and

Check

hygiene,

adjustenvironm

entof

light,temperature,humidity,andventilatio

n.In

Appl Microbiol Biotechnol (2010) 85:1321–1337 1327

Tab

le3

(con

tinued)

Problem

Cause

Solutions

mites

moresevere

cases,usehalfateaspo

onof

sulfur

in3.5lof

water.Mistthebags

andthe

surfaceof

mushrooms.Rem

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icklyspoil

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unger

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before

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midity

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gspot

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flush

Bacteria(Pseudom

onas

tolaasii,

Pseud

omonas

fluorescens)

onOystermushroom

Con

trol

humidity

inthemushroom

houseandmaintain80–85%

.Giveenou

ghtim

efor

water

toevaporatefrom

mushroom

surfaces

before

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severe

cases,

use113

gchlorine

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nof

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Sou

rce:

FAO

2001

1328 Appl Microbiol Biotechnol (2010) 85:1321–1337

at a temperature of approximately 45 °C for 4–5 days untilall the ammonia has evaporated. One of the most importantdevelopments in mushroom growing of recent years has beenthe introduction of bulk processes in composting technology.Both phase I and phase II of the composting process can takeplace in bulk in special fermentation rooms called tunnels.When that process has been completed, the compost can bespawned, and mycelium can develop in either the same oranother tunnel. Phase III then yields fully grown compostready for production (Van Griensven and Van Roestel 2004).

Cultivation of L. edodes

L. edodes is the third most cultivated edible mushroom; itwas 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 logcultivation that utilizes heat-treated substrates based onsawdust enclosed in plastic bags (“bag-log cultivation”).The main advantages of this method are the short time tocomplete a crop cycle and the higher yields (Sánchez2004). The strain, substrate composition, and length ofincubation are important parameters for shiitake productionon synthetic substrates in bag-log cultivation (Zadrazil1993; Kalberer 1995; Sánchez 2004). Recently, mushroomculture has moved toward diversification, and the culture ofother mushrooms has been reported.

Cultivation of Auricularia sp.

Auricularia auricula and Auricularia polytricha commonlyare produced on a medium consisting of sawdust, cotton-seed hulls, bran, and other cereal grains or on natural logsof broad-leaf trees (Table 2). For synthetic mediumproduction, the substrate may be composted for up to5 days or used directly after mixing. The mixed substrate issterilized for 60 min at 121 °C. Composted substrate isprepared by mixing and watering of ingredients. Theinoculation and fructification are carried out as previouslyreported (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 substratecontained in polypropylene bottles. Substrates (primarilysawdust and rice bran) are mechanically mixed and filledinto heat-resistant bottles, sterilized (4 h at 95 °C and 1 h at121 °C), mechanically inoculated, and incubated for 25 daysat 20 °C. To further improve quality during fruiting,temperatures are lowered to 3 °C to 8 °C until harvest. Asthe mushrooms begin to elongate above the lip of the bottle,a plastic collar is placed around the neck and secured with a

Velcro® strip. This collar serves to hold the mushrooms inplace so that they are long and straight. When themushrooms 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 substratecontained in polypropylene bottles or bags. A commonsubstrate used for production is composed of sawdustsupplemented with rice bran or wheat bran. It has also beenproduced on a substrate consisting of oak sawdust, wheatbran, millet, and rye. This formula gave the highest yields,best quality, and shortest crop cycle time (12 weeks). Forbottle production, the containers are filled with moistenedsubstrate and sterilized or pasteurized prior to inoculation.For production in bags, the moistened substrate is filled intomicrofiltered polypropylene bags and sterilized. Aftercooling (16 to 20 h), the substrate is inoculated, and thebags are heat-sealed and shaken to uniformly distribute thespawn throughout the substrate. Spawn runs last about 30to 60 days depending on strain and substrate formulation.After primordia formation, two holes usually are cut in thebags exposing the developing primordia that tend todevelop around the outside perimeter of the substratesurface. The top of the bag is then folded over, exposingonly 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 bottlescontained in plastic trays. After the completion of vegetativemycelial growth, bottle lids are removed, and the colonizedsubstrate is subjected to environmental conditions known tostimulate fruiting. When the mushrooms are mature, the entirecluster of fruiting bodies is removed from the bottles. Themushrooms are packaged by placing an entire cluster (ormultiple clusters) into each over-wrapped package. Only oneflush of mushrooms is harvested prior to mechanical removalof the “spent” substrate from the bottles. The bottles then arerefilled with fresh substrate, and the process is repeated(Royse 2007).

Cultivation of Pholiota nameko

Preparation of the medium for P. nameko cultivation is similarto that for F. velutipes except that a higher moisture contentof the substrate is desirable. A substrate of broad-leaf treesawdust is preferred, but research has shown that sawdustfrom conifers such Pinus spp. and Cryptomeria japonica are

Appl Microbiol Biotechnol (2010) 85:1321–1337 1329

suitable for growth. Rice bran usually is added as asupplement for conifer sawdust and 10% for broad-leafsawdust (Royse 2007).

Cultivation of Tremella fuciformis

This mushroom can be cultivated on natural logs or on amedium (Quimio et al. 1990). Cultivation techniques usedto produce the mushroom on natural logs is similar to thatused for shiitake production. In recent years, mostproduction of T. fuciformis has been on a substrate usinga mixed culture inoculum technique first developed inFujian, China (Huang 1982). The mixed culture techniqueinvolves the use of “helper” mycelium of Hypoxylonarcheri, an ascomycete commonly associated in naturewith decaying wood. H. archeri increases the ability ofT. fuciformis to digest the substrate thereby increasingmushroom yields. Exploitation of this mycelial associationis accomplished through use of dual cultures to makemother spawn (Quimio 1997).

Cultivation of Volvariella sp.

This mushroom is well suited for cultivation in the tropicsbecause of its requirement for higher production temperatures.It can be grown on non-pasteurized substrate which is moredesirable for low input agricultural practices. In recent years,cotton wastes (discarded after sorting in textile mills) havebecome popular as substrates for straw mushroom production(Chang 1982). Cotton wastes give higher and more stablebiological efficiencies (30% to 45%) and earlier fructification(4 days after spawning) and harvesting (first 9 days afterspawning) than that obtained using straw as a substrate. Theculture of other mushrooms has also been reported (Table 2).

Improvement of strains

The strain used in the culture is crucial for success ofmushroom production and marketing. A strain with a highability to invade the substrate and to fruit diminishes timeof incubation and enhances productivity. The consistencyand texture of the mushroom are crucial to diminish lossesduring packing. The first A. bisporus hybrid was obtainedby breeding about 25 years ago. However, most of thestrains currently used are similar to the first hybrid obtained(Sonnenberg 2000; Kerrigan 2000; Sánchez 2004). Theincrease in mushroom production has been the result ofmore specialized studies carried out by several internationalinstitutions in different areas of mushroom cultivation(Tables 3 and 4). The use of DNA-based technology hasaccelerated breeding activities and will help mushroom-breeding programs (Stoop and Mooibroek 1999). An

important advance in the development of techniques forbreeding is based on the development and detection ofgenetic markers (Sonnenberg 2000). Research on themolecular basis of mating-type genes has been veryimportant 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 ofparticular importance, since some workers develop anallergy that is identical to “mushrooms workers lung” andis 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 alsobeen observed with L. edodes spores (Laborde 1995; Sentiet al. 2000). This problem has increased interest indeveloping sporeless mutants for breeding of sporelessstrains (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 andcomputerized monitoring of growing rooms, automatedmushroom harvesting machines, hydroblending and pre-wetequipment (heap turners), or to develop methods to producemushrooms on a non-composted substrate, and new techni-ques of sterilization (Hawton et al. 2000). A computerizedenvironmental control system is an invaluable tool formushroom culture. The computer monitors environmentalparameters such as temperature, humidity, airflow, pressure,and carbon dioxide and oxygen content. However, automaticcontrol of ammonia levels and moisture in the casing soil isstill not widely applied. More than two decades ago, the firstclimate computers were introduced to Dutch mushroomgrowing and now are widely used in the industry (Lamber2000). Climate control in production plants allowsmonitoringand control of several mushroom production rooms with aminimum human input. Computerized environmental controlsystem allows the grower to look and adjust the environmentalconditions of the plant by using remote access (modem)capabilities (Walker 1996). There are several companieswhich provide different environmental control systems, e.g.,DuraFlex, Jaybird Manufacturing, Agricultural Engineering,etc. Picking and packing (Fig. 3k) constitute the mostexpensive part of the overall production process. In the last20 years, several studies have been carried out to developautomated mushroom harvesting machines. Some companiesoffer different automated harvesting systems, which includesmall, modular semiautomatic picking aids and the fullyautomated, robotic systems designed for line picking (Kensal

1330 Appl Microbiol Biotechnol (2010) 85:1321–1337

Automation; Astell 1996). The harvesting operation includesmushroom location, sizing, selection, and picking. Themechanical properties used for the analysis of automatedharvesting were obtained from compression experiments

with cylindrical mushroom sample pieces (Hiller 1994).Reed et al. (1995) evaluated at the laboratory level theautomated harvesting of A. bisporus by machine, and theresulting pilot harvester was successfully tested on a

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

Area of study in mushrooms production Expected output Internationalinstitutions

Germplasm and genetic improvement

Survey, collection, and identification of wild relativesand new species

Assessment of biodiversity and conservation of germ plasm RBG, MI, CMI

Culture maintenance Conservation of germ plasm CMI, ATCC

Genetic improvement Increased productivity, improved quality, disease and pestresistance

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,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 storageand transport of fresh mushroom

Reduced postharvest losses HRI, PSU

Development of improved processing technologies formushrooms

Reduced postharvest losses and avoidance of distress sale PSU

Development of production technologies for value-added products from mushrooms

Development of value-added products

Basic and strategic studies in mushrooms

Molecular genetics—mapping, cloning and sequencingof genes, allele mining, creation of gene libraries

Understanding will help manipulation for improvement in yieldand quality libraries of mushrooms

HRI, MES, PSU

Genomics—genome 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 ofcommercial types

HRI

Molecular mechanisms of biodegradation oflignocelluloses

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 CultureCollection, 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 DevelopmentCooperation, Israel; CAL College of Agriculture Laguna, Philippines. CU Chinese University, Shatin, Hong Kong; USFPL US Forest ProductsLaboratory, USA; FSU Fredrick Schiller University, Germany

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

Appl Microbiol Biotechnol (2010) 85:1321–1337 1331

commercial mushrooms farm. The apparatus combinesseveral handling systems and mechatronic technologies.Mushrooms are located and sized using image analysis and amonochromatic vision system. An expert selection algorithmthen decides the order in which they should be picked andwhat picking action (bend, twist, or both) should be used(Tillett and Batchelor 1991; Reed et al. 1997). One of a pairof suction cup mechanisms attached to the single head of aCartesian robot is then deployed; it can delicately detachindividual mushrooms and place them gently into a speciallydesigned compliant finger conveyer. After high speedtrimming, a gripper mechanism is finally used to removemushrooms from the conveyor into packs at the side of themachine (Reed et al. 1995, 2001).

The pre-wet heap machine was designed to achieve a quickhomogeneous mix of water and the raw materials used inphase 1 of the culture of A. bisporus to improve efficiency ofthe composting process by reducing the amount of timerequired to achieve the same or better quality compost. Somebenefits of using pre-wet compost turners include improvingcompost quality, homogenous blending of up to 200 t perhour, better odor control, less water run-off, raw materialsavings, reducing phase 1 time by over 50%, reducingoperating and capital costs, increasing production yields,crop uniformity, and profit potential. Design of newequipment to improve productivity is continuously indevelopment. Recently, a new tunnel/bunker filling systemhas been developed, which consists of filling the bunkers ortunnels using overhead layering techniques, but only fromone end of the bunker/tunnel and not through holes in thetunnel roof, with the use of overhead and out of sightconveyors/elevators (Traymater machinery).

New methods to produce and increase the productivity of awide variety of exotic mushrooms have been developed. Themycocell system (Mycocell Technologies, UK) is a methodbased on microwave sterilization of pre-packaged substrate towhich the “spawn” and other nutrients (e.g., Ca2SO4) can beadded. In this case, radiation used in the treatment of thesubstrates modifies the cellulose and increases ease ofbreakdown by the mycelium. In addition, several nutrientscan be added and mixed thoroughly, there are no risks ofsubstrate contamination, and the colonization of the substrateby the mycelium considerably increases. The commercialbenefits of this production system include lower cost, lowenergy requirements, automation, low labor demand, lack ofdowntime, light and cheap transport, low contamination risk,and long shelf life. The mycocell system has allowed also thesuccessful 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, Ganodermalucidum, Psilocybe sp., G. frondosa, Hericium sp., andAuricularia (Hawton et al. 2000; Table 5).

A technology called variable frequency speed control foralternating current motors (Keljik 1995) may be useful formushroom farms and benefit the industry by improvedcontrol of air velocity in the growing environment, resultingin lower electricity consumption (Lomax 1989, 1992;Sánchez 2004). Currently, there are several specializedinstitutes that study different frontier areas to improvemushroom cultivation (Tables 3 and 4).

Additional benefits of mushrooms culture

Several studies have shown that spent SMS can be used formushroom re-cultivation, e.g., the use of SMS enrichedwith cotton seed meal and with soya meal for Agaricuscultivation, the use of spent Pleurotus substrate for the KingStropharia cultivation (Poppe 1995), the use of spentAgaricus compost added with cotton waste for Volvariellaproduction (Oei 1991), and the use of spent Pleurotus eousstraw 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 theproduction of Pleurotus sajor-caju. Utilization of spentstraw generated after the cultivation of G. lucidum andF. velutipes has been recommended for the production ofother mushrooms, such as A. bisporus or Coprinus comatus(Xiao 1998). A. bisporus spent substrate has also been usedfor the cultivation of Volvariella (Poppe 2000). SMS canalso be used as casing material for the production ofAgaricus (Nair and Brandley 1981; Shandilya 1989; Singhet al. 1992, 2000). It has also been used as animal feed,since its degradation by the mushroom can improve itsnutritional quality (Jalc et al. 1996a, b; Adamovic et al.1998; Díaz-Godínez and Sánchez 2002) and digestibility bythe ruminant (Capelari and Zadrazil 1997; Díaz-Godínezand Sánchez 2002). Díaz-Godínez and Sánchez (2002)found that addition of maize straw generated after mushroomcultivation to the diets of sheep increased the weight gain ofthe sheep and the efficiency of feed conversion of thestraw. SMS is also a beneficial product for enrichment ofsoils, restoring areas that have been destroyed throughdevelopment, deforestation, or environmental contamination.Some studies have been done on the use of SMS in vegetableand flower greenhouses (Lohr et al. 1984; Verdonck 1984;Steffen et al. 1994, 1995; Szmidt 1994; Söchting and Grabbe1995; Celikel and Tuncay 1999), in field vegetable and fruitscrop (Male 1981; Pill et al. 1993; Ranganathan andSelvaseelan 1997; Stewart et al. 1998; Batista et al. 2000;Delver and Wertheim 1988; AntSaoir et al. 2000), in nurseryand landscape gardening (Chong and Wickware 1989;Chong and Rinker 1994; Chong 1999), in soil amendmentas organic manure or vermi-compost (Stewart et al. 1998;Tajbakhsh et al. 2008), and biogas production (Balan et al.

1332 Appl Microbiol Biotechnol (2010) 85:1321–1337

2008; Pathak et al. 2009). Currently, there are someindustries that manufacture and sell different kind ofcompost based on SMS (http://www.nutrasoils.com; Meijer2009; American Mushroom Institute 2006; http://www.laurelvalleysoils.com).

The potential of SMS to degrade organopollutantsand its importance in environmental bioremediation hasbeen reported (Kuo and Regan 1998; Eggen 1999;Xawek et al. 2003; Webb et al. 2001; Lau et al. 2003;Law et al. 2003).

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

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

Affectedmushroom

Infestation(organism)

Type oforganism

Diseases Symptoms Reference

Agaricusbisporus

Cladobotryumdendroides

Fungi Cobweb, mildew White to pink cobweb-like fluffy mold Western Committee on Plant Disease(2004)

Agaricusbisporus

Mycogone sp. Fungi Wet bubble/whitemold

Dense white growth on gills Western Committee on Plant Disease(2004); Tanović et al. 2009

Agaricusbisporus

MycogoneperniciosaMagn.

Fungi Wet bubble of fungi Boiko et al. 2009; Tanović et al. 2009

Agaricusbisporus

Trichoderma sp. Fungi Green mold Dark green mold patches on casingspreading to lesions on stems

Western Committee on Plant Disease(2004)

Pleurotusostreatus

Velázquez-Cedeño et al. 2004

Agaricusbisporus

Verticillium sp.fungicola

Fungi Dry bubble/brownspot

Brown irregular pitted areas on stems andcaps. Distortion and splitting. Severerotting, blotch, and necroses of caps andstems

Western Committee on Plant Disease(2004); Boiko et al. 2009

Agaricusbisporus

Lycoriella sp.,L. ingenua

Insect Sciarid flies Destroy pins of developing mushrooms, andburrow or tunnel into the stems and caps ofmaturing mushrooms

http://www.mushroomcompany.com/resources/pests/index.shtml, 2009;Jess and Schweizer 2009

Agaricusbisporus

Megaselliahalterata

Insect Phorid flies Cause less mushroom damage than sciaridflies

http://www.mushroomcompany.com/resources/pests/index.shtml, 2009;Smith et al. 2006

Agaricusbisporus

Aphelenchoidescomposticola

Nematode Eelworms,Cephalotheciumdisease

Degeneration of the mushroom mycelium inthe compost

http://www.mushroomcompany.com/resources/pests/index.shtml, 2009;Gitanjali 2005

Agaricusbisporus,

PseudomonasTolaasii

Brown blotch Sunken, dark brown lesions http://www.mushroomcompany.com/resources/pests/index.shtml , 2009

Pleurotusostreatus,

Pseudomonasreactants

Blotch disease Mild dark purple to light brown discolorationand a slight surface depression thatbecomes darker with age

Godfrey et al. 2001

and othermushrooms

Pseudomonasgingeri

Ginger blotch Pale yellowish red discoloration thatdevelops into a reddish ginger-colored palediscoloration

Agaricusbisporus

Putative virus X Virus Virus X Slightly imperfectly formed cap structure Gaze 2001

Agaricusbisporus

Bacilliformviruses

Virus Severe rotting, blotch and necroses of capsand stems

Boiko et al. 2009; Revill et al. 1998

Appl Microbiol Biotechnol (2010) 85:1321–1337 1333

Diseases presented during the mushrooms cultivation

In mushrooms production, a composted substrate increasesmushroom yield and reduce infestation by insects, fungi,bacteria, and viruses. However, under vulnerable mush-room growth conditions, several pests can attack themushrooms cultivation (Table 6).

Troubleshooting in mushroom cultivation

During cultivation of mushrooms, several problems occur(Table 3); however, following of mushroom cultivationcarefully can prevent them. Prevention is better thansolving such problems (Table 6; FAO 2001).

Conclusions

Edible mushrooms have been eaten and appreciated for theirflavor, economical and ecological values, and medicinalproperties. They are able to grow under different climaticconditions on cheap, readily available waste materials. Thesemushrooms are a clear example of how low-value waste,which is produced primarily through the activities of theagricultural, forest, and food-processing industries, can beconverted to a higher value material useful to mankind.Recently, mushroom culture has moved toward diversification,and the culture of additional mushrooms has been reported. Formany reasons, the fungal Pleurotus genus has been intenselystudied and cultivated in many different parts of the world.Particularly, P. ostreatus, also known as “oyster mushroom”,“hiratake”, “shimeji”, or “houbitake”, requires shorter growthtime when compared to other edible mushrooms. Also, thesubstrates used for its cultivation do not require sterilization,only pasteurization which is less expensive. This mushroomdemands few environmental controls for cultivation, and itsfruiting bodies are not often attacked by diseases and pests,and it can be cultivated in a simple and cheap way. Anotheradvantage of growing oyster mushrooms is that a highpercentage of the substrate is converted to fruiting bodies,increasing profitability as compared to other mushrooms,making P. ostreatus an excellent choice for mushroomcultivation.

Acknowledgement I am grateful to Dr. A. L. Demain for criticalreading of the manuscript.

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