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BASE Biotechnol. Agron. Soc. Environ.201620(3),363-374
AgronomicassessmentofspentsubstratesformushroomcultivationRaquelPicornell-Buendía(1),ArturoPardo-Giménez(2),JoséArturodeJuan-Valero(1)(1)EscuelaTécnicaSuperiordeIngenierosAgrónomosdeAlbacete.Castilla–LaManchaUniversity.CampusUniversitarios/n.ES-E02071Albacete(Spain).E-mail:[email protected](2)CentrodeInvestigación,ExperimentaciónyServiciosdelChampiñón(CIES).C/Peñicas,s/n.Apartado63.16220QuintanardelRey.ES-Cuenca(Spain).
ReceivedonAugust7,2015;acceptedonMarch7,2016.
Description of the subject. In thiswork the agronomicviabilityof substratesbasedon spentAgaricusbisporus Imbach(Lange)substrates(SAS)andspentPleurotus ostreatus(Jacq.)P.Kumm.substrates(SPS)isstudied.Objectives.The aimof thiswork is thequalitative agronomic evaluationofSPSandSASand themixtureof thereof indifferentproportions,suchaslignocellulosicsourcesinnewgrowingcyclesofP. ostreatus.Method.Inadditiontothecommercialsubstrateusedasacontrolreference,sixdifferenttreatmentsareconsidered.Inthisexperiment,SPSandSASweremixedindifferentamounts.SASwassubjectedtoaheattreatmentinagrowingroom(“cookout”)andthentoamaturationtreatmentwhichconsistedofacontrolledrecompostingprocessincameras.SPSwassubjectedtoapasteurizingheattreatment(60°C–65°C,8h)andaprogressivetemperaturedecreaseforatleast15htoa“seeding”temperature(25°C).Results.SPS(3,600g)+SAS(2,400g)andSPS(3,000g)+SAS(3,000g)werepreparedsubstratesthatachievedacceptablecrudeproteincontentintheirfruitingbodies.Additionally,weobtainedhigherashcontent,lightness,yellow-blue(y-b)andred-green(r-g)chromaticity,breakingstrength(Bs),andcompressionenergy(Ce)inthesemushrooms.Thesevalueswerehigherthanthemeanvalues,andevenhigherthanthecommercialsubstrate.Conclusions. IncreasedSASparticipationin themixtureof theprocessedsubstrate(andtheconsequentreductionofSPSparticipation) resulted inmushrooms that require higherBs, andCe.These formulation-based composts degraded by thegrowthofP. ostreatus,couldbealow-costsubstratewithselectiveandbalancednutrientsforgrowthanddevelopmentofoystermushrooms.Keywords.Pleurotus ostreatus,agriculturalwastes,ediblefungi,growingmedia.
Évaluation agronomique de substrats de culture épuisés pour la culture de champignonsDescription de l’objet.Dans ce travail, est étudiée la viabilité agronomique de substrats à base de substrats épuisés deAgaricus bisporusImbach(Lange)(SAS)etdesubstratsépuisésdePleurotus ostreatus(Jacq.)P.Kumm.(SPS).Objectifs.Lebutdecetravailest l’évaluationagronomiquequalitativedeSPSetSASet lemélangedeceux-cidansdesproportionsdifférentes,tellesquelessourceslignocellulosiques,dansdenouveauxcyclesdecroissancedeP. ostreatus.Méthode.Enplusdu substrat commercial utilisé comme référence, six traitementsdifférents sont considérés.Dans cetteexpérience,SPSetSASsontmélangésenquantitésdifférentes.SASaétésoumisàuntraitementthermiquedansunechambredecroissanceetensuiteàuntraitementdematurationquiconsisteenunprocessusdecompostagecontrôléenchambre.SPSaétésoumisàuntraitementthermiquedepasteurisation(60°C–65°C,8h)etàunediminutionprogressivedelatempératurependantaumoins15hjusqu’àunetempérature«d’ensemencement»(25°C).Résultats.SPS(3,600g)+SAS(2,400g)andSPS(3,000g)+SAS(3,000g)sontlessubstratsquipermettentd’obtenirunevaleuracceptabledeprotéinedansleursorganesdefructification.Deplus,nousavonsobtenudesvaleursplusélevéesdesdifférentsparamètrespourceschampignons,mêmeplusélevéesquepourlessubstratscommerciaux.Conclusions. Une quantité plus importante de SAS dans le mélange (et la diminution conséquente de SPS) donne deschampignonsquiprésententunerésistanceàlaruptureetuneénergiedecompressionplusélevées.CestypesdecompostsdégradésparlacroissancedeP. ostreatus donnentdessubstratsdefaiblecoutavecdesnutrimentssélectifsetéquilibréspourlacroissanceetledéveloppementdespleurotes.Mots-clés.Pleurotus ostreatus,déchetagricole,champignoncomestible,substratdeculture.
364 Biotechnol. Agron. Soc. Environ. 201620(3),363-374 Picornell-BuendiaR.,Pardo-GiménezA.etal..
1. INTRODUCTION
Thepotentialproductionofediblefungiispromisingin theworldmarket, as thecurrentproduction isnotsufficienttomeetthedemand.Mostproducingcountriesare importers too, since the average consumptionin these countries is very high. Nutritionally, thesespecies(Agaricus bisporus[Lange]Imbach,Pleurotus ostreatus [Jacq.] P. Kumm. and Lentinula edodes[Berkeley], among others) have moderate amountsof high quality proteins, contain all essential aminoacids,andarerichinlysine,leucine,CandBvitamins,minerals,andothertraceelements.Furthermore,theirlipidlevelsandratioofsaturatedtounsaturatedfattyacidsarelow,theycontainrelativelyhighamountsofcarbohydrates,andmostspecieshavehighamountsofnutritionallyvaluablefibers(Changetal.,1997).
Complementary to its nutritional properties,there are various health benefits known in the fieldsof medicine and therapeutics such as, antitumor,antibiotic, antifungal, and anti-inflammatory effects.Additionally, they are hypocholesterolemic, promotea healthy immune system, and have been widelyused to treat cancer andHIV (Brizuela et al., 1998).Approximately, there are 300 species of cultivatedmushrooms,butonly30havebeendomesticatedandjust10arecommerciallygrown.Themost importantcultivated mushroom worldwide is A. bisporus,followedbyP. ostreatusandotherspeciesofthegenusPleurotus;L. edodesisthirdandotherediblefungiaremakingheadwaysinthemarket.
ThereasonforthiscommercialgrowthisbecauseofthecharacteristicsofthespeciesofthegenusPleurotus(Sanchez, 2010). They have excellent organolepticqualities, are easy to grow on a wide variety ofsubstrateswithinawidetemperaturerange,andhaveagreatpotentialinbioremediationprocesses.Moreover,littleinitialcapitalisrequiredtoestablishwarehousesforcultivation.
Unlike white mushrooms (A. bisporus), these donot require a chemically selective substrate becausethey can grow in nutrient media with a C/N ratiobetween30and300(RodriguezBarreal,1987;GarciaRollán,2007).However,theyneedtogrowinaspecificbiological environment with accompanying florato protect and promote growth (Muez et al., 2002).Approximately,13,500tofthisfungusareproducedinCastilla-LaMancha(67%ofthenationaltotal)(Pardoetal.,2009).
Themushroomgrowingsector inSpaingeneratesabout 5.105t of spent compost, while the EU, as awhole, produces more than 3.5.106t (Pardo et al.,2009; Picornell et al., 2010). This lignocellulosicmaterial,calledmushroomspentsubstrate,canbeusedinvariousfieldsofagriculture:animalfeed(Zadrazil,1980b), amendments (Tajbakhsh et al., 2008),
substratesofnurseries,nurseries(Medinaetal.,2009),bioremediation (Faraco et al., 2009), aquaculture,vermicultureandbiofuel(Pathaketal.,2009).
However theseusesdonot take advantageof thehigh volume generated annually, which accumulatesin collection centers located in production areas ofSpain.Furthermorethesespentsubstratesarepotentialcontaminants,aswellas,awasteofenergy.Pleurotus ostreatus has specific enzymes capable of degradingcellulose, lignin,phenolsandpolyphenols to60%oftheoriginalcontentof thespentsubstrates (Picornelletal.,2010).
Currently, cereal straw (wheat inparticular),withincreasing constraints in availability and price, isvirtuallytheonlymaterialusedatanindustrialscalefortheproductionofP. ostreatusinSpain.Thefeasibilityof using highly available alternative materials of alowcostisalineofresearchwithgreattechnologicalinterest for improving productivity and reducingprocessingcosts(Muezetal.,2002;Pardoetal.,2007;Pardoetal.,2009;Picornelletal.,2010).
Accordingtovariousstudies, themostcommonlyprofitableandreadilyavailablespentsubstratewhichgenerateshighqualityfruitingbodies forP. ostreatus(although this fungus can be grown on virtually anylignocellulosic substrate) is the trunk of Quercus humboldtii Bonpland (oak) (Garcia Rollán, 2007).Commerciallyinmostindustrialexploits,2-4cmlong(Sanchez,2010)piecesofwintercerealstraw(wheat,barley,andrye)(Savalgietal.,1994)areusedinthesubstrate container for the production of Pleurotusgenusandothers,suchasPleurotus eryngii(DC.:Fr.)Quel., Pleurotus sajor-caju (Fr.) Singer, Pleurotus pulmonarius substrate (Fr.)Quél., etc.Khanna et al.(1982)refertoricestrawasthebestsubstrateforthecultivationofP. sajor-caju,whilewheatstraw(whichis similar to rice straw) is the best substrate for thecultivationofPleurotusspp.(Bonattietal.,2004).
Biodegradation of these cellulosic residues byPleurotus spp. cultivationdependson theproductionofhemicellulases,cellulases,and ligninasesenzymes(Kurt et al., 2010). These enzymes, and others,turn and degrade long and insoluble components oflignocellulosic materials into soluble components oflowmolecular weight that are taken by intracellularenzymes from fungi for their nutrition.Additionally,enzymes play an important role in the growth anddevelopmentoffungi(Kuforijietal.,2008).
However, lignocellulosic materials are generallylow in protein content with insufficient values ofnitrogen, phosphorus, and potassium (Vijay et al.,2007)formushroomcultivation.Organicsupplementsaremostcommonlyusedinthepreparationofgrowingsubstrates, with wheat and rice bran being the mostpopular(Wangetal.,2001;Peksenetal.,2009;Kurtetal.,2010).
Agronomicassessmentofspentsubstrates 365
Theaimofthisworkistheagronomicassessmentof substrates based on spent A. bisporus (SAS) andspent P. ostreatus (SPS). The use of the remainingspent mushroom substrate after the cultivation ofP. ostreatus in new production cycles would be anagronomicallyviablealternativetousingwheatstraw(WS), which is currently used exclusively as a basematerial.Economically,theuseoftheremainingspentmushroom substrate is beneficial when consideringthehighcostofWS,especiallyindroughtyears.Thiscerealfarmer’sby-productcouldbeintegratedthroughnew formulations andmethodologieswhile loweringproduction costs and reducing the environmentalimpactofunusualwaste.
2. MATERIALS AND METHODS
2.1. Analytical methodology used for the characterization of materials
The characterization of rawmaterials and processedsubstrates was measured according to the followingparameters:moisture (MAPA, 1994), pH (Ansorena,1994),totalnitrogen(Tecator,1987;MAPA,1994),ash
(MAPA,1994),organicmatter(Ansorena,1994),C:Nratio,crudefiber(ANKOM,2008),crudefat(ANKOM,2009),nitrogenfreeextractives(NFE)(Gonzálezetal.,1987),celluloseandneutraldetergent‐soluble (NDS)(ANKOM,2005;ANKOM,2006a;ANKOM,2006b).Furthermore, theexplorationofmites (Krantz,1986)andnematodes(Nombelaetal.,1983)wasconductedinthisexperiment,showingnegativeresultsfortestedsubstrates.
2.2. Preparation of substrates and experimental design
Theonlyagronomicfactorstudiedinthisexperimentisthetypeofbasesubstratewithfourblockreplicates.In accordance with the experimental design, sevendifferenttreatmentsweregeneratedintheprocess,aswell as a reference commercial substrate proper forPleurotusspeciescultivation.Inalltreatments,exceptthe control, 50g.kg-1 CaSO4were added to the basematerial.CaCO3(gypsum)wasalsoaddedinvaryingamounts(Figure 1;Table 1).
SASwassubjectedtoaheattreatmentinagrowingroom(“cookout”)andthentoamaturationtreatmentwhichconsistedofacontrolledrecompostingprocess
CROP CYCLE of Agaricus bisporus (Lange) Imbach
SPENT Agaricus bisporus (Lange) Imbach SUBSTRATE (SAS)
SPENT Pleurotus ostreatus (Jacq.) P.Kumm. SUBSTRATE (SPS)
CROP CYCLE of Pleurotus ostreatus (Jacq.) P.Kumm.
ELABORATION OF BASE SUBSTRATES FOR THE GROWTH OF Pleurotus ostreatus (Jacq.) P.Kumm.
HEAT TREATMENT
MATURATION
ANALYTICALCHARACTERIZATION
AGRICULTURAL EVALUATION IN GROWING ROOMS
DATA PROCESSING AND ANALYSIS
COMMERCIALLY CONTROLLED BASED
SUBSTRATES
MONITORING OF PROCESS
Figure 1.Schematicexperimentapproach—Approche expérimentale schématique.
366 Biotechnol. Agron. Soc. Environ. 201620(3),363-374 Picornell-BuendiaR.,Pardo-GiménezA.etal..
incameras.SPSwasfromChampymar(QuintanardelRey,Cuenca)whichwasinitiallyobtainedfromfreshwheat straw (maximum one week in empty rooms).Materials were mixed and the moisture content wasadjusted.Thenapasteurizingheattreatment(60°C–65°C,8h)wasperformed.Afterwards,thetemperaturewas decreased to the “seeding” temperature (25°C)over 15h. Finally, supplementation and “seeding”(dose 30g.kg-1 mycelium Fungisem K-15) wereperformed and the samples were bagged in theCenterforResearch,ExperimentationandMushroomServices’(CIES)pilotplant.
All substrates were packed into transparentpolyethylenebagsof29cm indiameterwithheightsranging from 25 to 35cm, according to the type ofsubstrate, totaling approximately 6.5kg of weight.Four holes of 2.2cm in diameter were uniformlydrilledoverthesideofthebags.
2.3. Driving and monitoring of the crop cycle
The total research timewas70days.Theexperimentwas carried out at the CIES, located in the town ofQuintanardelRey(Cuenca,Spain)inanexperimentalgreenhouse controlled for temperature, substratetemperature, relative humidity, and carbon dioxideconcentration,andfollowedtherecommendedrangesfor thevarietyofselectedmyceliumineachstageofdevelopment(CIES,2007).
Acontrolledrecompostingprocesswascarriedoutconsisting of a heat treatment in the growing room.Sinceairheatsfasterthanthesubstrate,theairreachedthe maximum temperature (71°C) in 19h, whereasthe substrate reached this temperature in 22h. Thedurationof thesehigh temperaturesalsodiffers from
one medium to another. The maximum temperatureoftheairandofthesubstratestabilizesfor7and3h,respectively.Aftertheheattreatment,thetemperatureis decreased. The air temperature fell more rapidlythanthesubstratetemperaturecausingthesecondtobeabovethefirst.Finally,thesetemperaturesequilibrateto34–38°Cafter31h.
Duringrecompostingandmaturation,themoisturecontentofthemasswasmaintainedat500g.kg-1withoutleaching.Inthisripeningprocess,theevolutionofthemoisture content in non-intervened composting hadvalues that rangedfrom527g.kg-1 (in thebeginning)and504g.kg-1(attheend).
Substrateincubationlastedapproximately17days(without external ventilation or lighting).During theincubation period, the relative humidity inside thegreenhouserangedbetween90%and95%,while thesubstratetemperaturewasbetween16°Cand23.78ºC.Theroomtemperaturerangedbetween18ºCand22ºCto help control the temperature of the formulations.Afterthis,fruitingwasinducedbyventilation(tokeepCO2levelsregulatedbetween0.28%to0.10%),andthereductionofroomtemperature(21.57°Cto13.63°C),substratetemperature(23.78°Cto14.73°C),humidity(91% to 94.50%), and lighting. These values aresimilartothemicroclimaticconditionsrecommendedby other researchers (Pardo et al., 2005b; GarcíaRollán,2007;Pardoetal.,2007;Gregorietal.,2008;López-Rodríguezet al., 2008;Geaet al., 2009;Kurtetal.,2010).
2.4. Evaluation of parameters
For the evaluation of quality parameters, freshmushroomsofuniformsizeandmaturitywereusedandselectedonthemostoptimaldayoftheharvestperiod.Thesurfacecoloroffruitingbodieswasmeasuredbyreflection using a Minolta brand colorimeter, modelCR-300,previouslycalibratedwithacalibrationplateCR-A43 (L = 96.12, r-g = -0.11, y-b = +2.66) andilluminantD65.Toevaluatethemechanicalpropertiesofmushrooms,intermsoffirmness,ananalyzer(TA-XTPlusofStableMicroSystems)wasused.Totakethismeasurementthefruitingbodieswerecutintosmallpieces(~4cm2)andwereintroducedintothe5-bladedKramer Shear Cell (KS5), arranged in two adjacentuniformlayersandtestedataconstantspeedof2mm.s-1;thus breaking strength (Bs)was obtained, defined asthemaximumforcerequiredtotearthefruitingbodies(expressed inN). Protein content in the carpophoreswas calculated by multiplying the total nitrogencontentbyaconversionfactorof4.38(Delmas,1989).TotalnitrogencontentwasdeterminedbytheKjeldahlmethod(TECATOR,1987;MAPA,1994).Ashcontentwasdeterminedbyadirectcalcinationofthesamplesat540°C(MAPA,1994).
Table 1. Formulations tested (g per bag) in theexperiment — Formulations testées (g par sac) dans l’expérience.Treatment SPS SAS GYPSUMT1 6,000 0 300T2 5,400 600 300T3 4,800 1,200 300T4 4,200 1,800 300T5 3,600 2,400 300T6 3,000 3,000 300T7 Commerciallycontrolledbased
substrates(6.5kgperbag)T:treatment—traitement;SPS:spentPleurotus ostreatus(Jacq.)P.Kumm.substrate—substrat épuisé de Pleurotusostreatus (Jacq.) P. Kumm. ;SAS:spentAgaricus bisporus(Lange)Imbach.substrate—substrat épuisé de Agaricusbisporus(Lange) Imbach.
Agronomicassessmentofspentsubstrates 367
Dependingonthelevelofspawnruntimeofthesubstratebythemyceliumandtestedcontaminations,a parameter was established designated as thegermination index (GI), on a scale from 0 (noinvasion)to5(fullinvasion).
Theearlinesswasestablishedasthetimeindayssincethe“seeding”ofthesubstratetothefirstflushharvested(weighingthedailyrelativeproductionofthesubstrate).Aflushcorrespondstoeachproductioncyclethatisrepeatedrhythmicallyduringtheharvest.
2.5. Statistical analysis
To carry out the statistical analysis, two softwarepackages were used: Statgraphics® Plus version5.1 (Statistical Graphics Corp., 2001) and SPSS®(SPSS, 2004). Descriptive statistical techniques,principal component analysis, variance analysisand correlation and regression methods were usedto evaluate the data. Differences were consideredsignificantforp <0.05.
3. RESULTS AND DISCUSSION
3.1. Analytical characterization of the base materials used and the formulations
The chemical characteristic results of the differentsource materials, formulations, and commerciallycontrolled based substrates are shown in table 2.GrowthandnutritionalvalueofgenusPleurotusspp.mushroomsdependmainlyon the typeofsubstrateandgrowingconditions (Curvettoetal.,2002),butalsoonthetreatmentsappliedtomadesubstratesandthegrowingmycelium(Pardoet al.,2005a).Theseconsiderationsarealsovalid forotheredible fungi,forexample,L. edodes(“shiitake”)(Philippoussisetal.,2002;Ozceliketal.,2007).
The final values obtained after the SASrecomposingprocesswereadjustedto theoptimumrange considered for this process (Lohr et al.,1984;Szmidt,1994;Raymondetal.,1997).Higherconcentrations of P, K, Ca, and Mg in the coverlayer (Raymond et al., 1997) and a higher ashcontent(whichcontributesgreatlytothecoverlayerof mineral soil) (Stewart et al., 1977; Lohr et al.,1984)wereobserved,whencomparedtothecontrolsubstrate.
In this experiment, the chemical characteriza-tionwas studiedusing twodifferentbasematerialsin the made substrates. Increasing doses of SASincreasethepHfrom5.88to8.07andtheashcontentfrom27.66%to50.88%. Incontrast, this treatmentdecreasesmoisturecontentfrom75.60%to72.50%,organicmatter from 72.34% to 49.12%, C/N from Ta
ble
2.Physicochem
icalcharacterizationofso
urcematerialsandsu
bstratesu
sed—C
arac
téri
satio
n ph
ysic
o-ch
imiq
ue d
es m
atiè
res b
rute
s et d
es su
bstr
ats u
tilis
és.
pHM
oist
ure
Tota
l nitr
ogen
Prot
ein
Ash
cont
ent
Org
anic
mat
ter
C/N
rat
ioC
rude
fibe
rC
rude
fat
NFE
Cel
lulo
seN
DS
Base
materials
SPS
5.51
776.00
6.10
38.10
158.30
841.70
80.00
298.40
3.50
501.70
310.40
308.10
SAS
7.78
504.00
13.40
83.80
646.00
354.00
15.30
161.00
3.20
106.10
90.50
62.70
Substrates
T15.88
756.00
6.00
37.50
276.60
723.40
69.90
289.10
3.00
393.80
293.40
213.10
T26.96
742.00
7.40
46.30
384.30
615.70
48.30
271.60
3.40
294.50
215.10
176.30
T37.51
740.00
7.10
44.40
388.90
611.10
49.90
243.00
3.90
319.80
209.10
144.90
T47.94
719.00
7.60
47.50
474.10
525.90
40.10
239.60
4.20
234.60
188.40
125.50
T57.85
755.00
7.20
45.00
419.90
580.10
46.70
238.30
2.70
294.10
185.20
131.30
T68.07
725.00
8.60
53.80
508.80
491.20
33.10
231.00
2.50
204.00
153.30
111.80
T78.15
672.00
9.60
60.00
143.00
857.00
51.80
418.70
5.10
373.20
375.80
124.00
Average
7.48
729.00
7.64
47.80
370.80
629.20
48.50
275.90
3.54
302.00
231.47
146.70
CV(%
)10.88
3.98
15.11
15.10
33.60
19.80
23.50
24.00
26.02
22.65
33.25
24.43
T:treatment—
trai
tem
ent:see
tabl
e 1—voirt
able
au 1;C
V:coefficientofvariation—c
oeffi
cien
t de
vari
atio
n;NFE
:nitrogenfreeextracts—
ext
raits
d’a
zote
libr
e;NDS:neutral
detergent-soluble—
dét
erge
nt n
eutre
solu
ble;Resultsexpresseding. kg-
1 drymatter,exceptpH,m
oisture(freshmatter)andC/Nratio—
les r
ésul
tats
sont
exp
rim
és e
n g.
kg-1 d
e m
atiè
re
sèch
e, à
l’ex
cept
ion
du p
H, d
e l’h
umid
ité (m
atiè
re fr
aich
e) e
t du
rapp
ort C
/N.
368 Biotechnol. Agron. Soc. Environ. 201620(3),363-374 Picornell-BuendiaR.,Pardo-GiménezA.etal..
69.90 to33.10,crudefiber from28.91%to23.10%,NFEfrom39.38%to20.40%,cellulosefrom29.34%to15.33%andNDSdecreasesfrom21.31%to11.18%.
3.2. Production qualitative parameters. Descriptive statistics and ANOVA
Intable 3,descriptivestatisticsofcrudeproteinandash contents, as well as brightness values (L), red-green (r-g) and yellow-blue (y-b) chromaticity, Bs,andCeintheharvestedmushroomsareshown.
Ofthesixtreatmentswithdifferentspentsubstratemixtures, T1 consisting of 6,000g of SPS, showeddifficulties in germination, mycelial development,andgrowtharrest,sawahighdegreeofcontaminationbyGliocladiumspp.(whichcanbeassociatedtothelowpHofthesubstrate,Table 2),anddidnotproduceP. ostreatuscultivation.
Overall,proteinsarethemostcriticalcomponentsthatcontributetothenutritionalvalueofafood,andfungiareagoodsourceofthem(Crisanetal.,1978).
Most of the interest in the nutritional value ofmushrooms is related to thenutritional quantity andqualityofproteins(Manning,1985).Themushroom’sprotein content values achieved in this experimentrangedbetween15.48%(T2)and20.70%(commercialsubstrate,T7)(Table 3)arelowerthanthoseobtainedby Rodríguez Barreal (1987) (31%) and Benavidesetal.(2009)(34.09%).
Manysupplementssuchascottonseedpowderandrice bran (rich in nitrogen) are ideal for increasingyields and quality of the mushrooms (Jandaik,
1989). Organic supplements such as soybean meal,alfalfa, and cotton seed powder andmeal, not onlyincrease yields, but also the mushrooms proteincontent (Zadrazil, 1980a; Zadrazil, 1980b). Foreroetal.(2008)arguethatthereisapositivecorrelationbetweenthenitrogenandproteincontentsofthefungisubstrate,duetotheimportanceofthemacroelementas anutrient for thegrowthanddevelopmentof thefungus.
The protein content values obtained in thisexperiment were lower than those obtained withP. ostreatusandP. sajor-cajubyBasaketal.(1996),Wangetal.(2001)andMandeeletal.(2005)(between17.80%and23.30%).Aswellasthoseobtainedusingpepper residues as substrates (between 15.60% and28.60%)(Foreroetal.,2008).
Higher values in P. ostreatus were obtained byPardo et al. (2005a) in terms of protein contentdepending on the substrate type, treatment, andmyceliumused(from14.04%to17.75%).Pardoetal.(2007),alsousingP. ostreatus,gotsimilarvaluesofthose in this experiment in terms of protein contentdepending on the different types of substrate andtreatment used (from 19.07% to 23.98%). Pardoetal.(2008),workingwithP. ostreatus,gotdifferentmushroomprotein content values, depending on thetypeofsubstrate,substratetreatment,andtheweightofthebagused(from14%to26.30%).
Likewise, higher values than this experiment(between T2= 15.48% and T6= 18.79%) wereobtained by Bermudez et al. (2007) on Pleurotusspp. inblendsofcoffeepulp+cedarchipswith the
Table 3.PhysicochemicalcharacterizationofPleurotusfruitingbodiesgrownonthedifferentformulations—Caractérisation physico-chimique des fructifications de Pleurotus cultivées sur les différents substrats.Substrate Crude protein
(g.100g-1)Ash content(g.100g-1)
Color Bs (N) Ce (mJ)L r-g y-b
T2 15.48b 5.84c 72.89 1.46b 11.72a,b 314.54a,b 1.308.87b
T3 16.14b 6.37b 74.20 2.12a,b 9.66b 310.50a,b 1.350.50b
T4 15.73b 6.89a 72.41 2.53a,b 11.99a,b 303.14b 1.351.97b
T5 17.22a,b 7.24a 73.38 2.96a 14.21a 319.71a,b 1.429.01a,b
T6 18.79a,b 7.46a 74.22 2.67a,b 14.70a 350.96a 1.649.30a
T7 20.70a 6.67a,b 70.90 2.09a,b 11.94a,b 279.43b 1.294.96b
Average 17.34 6.74 73.00 2.31 12.37 313.05 1.397.43FisherF 6.41 7.15 0.99 2.85 4.30 5.57 5.40SL 0.001*** 0.001*** 0.45ns 0.05* 0.01** 0.003** 0.003**T:treatment—traitement:seetable 1—voirtableau 1;L:brightness—luminosité;r-g:red-greencolorcomponents—composants de couleur rouge-vert;y-b:yellow-bluecolorcomponents—composantsdecouleurjaune-bleu;Bs:breakingstrength—résistance à la rupture;Ce:compressionenergy—énergie de compression;SL,F:significancelevel,Fisher—niveau de signification Fisher;ns:nosignificantdifference—aucune différence significative,p-value>0.05;*:p-value<0.05;**:p-value<0.01;***:p-value<0.001;foreachcolumn,valuesfollowedbydifferentlettersaresignificantlydifferentfromeachother—dans chaque colonne, les valeurs suivies de lettres différentes sont significativement différentes les unes des autres(p=0.05,Tukey-HSD).
Agronomicassessmentofspentsubstrates 369
strainsCCEBI3021andCCEBI3027:27%and34%,respectively;substratesoncoffeepulp:30%and38%,respectively; and substrates based on cedar chips:21% and 22%, respectively. Fonseca et al. (2009)alsoachievedhighervalues inproteincontentusingassubstrates,amixtureofricebran(40%),ricestraw(35%)andJuncus effususL.(25%)inthegrowingofP. ostreatus(30.52%).
Similarvaluesforproteincontenttothoseobtainedin this experiment were achieved byOyetayo et al.(2004) working with P. sajor-caju, comparing theproteincontentofmushroomsgrowninsubstratesofgrated cobs (17.49%) and without supplementation(14.94%) and, similar values in P. eryngii, Manziet al. (2004) (22.20%), and Hassan et al. (2010):sawdust (22.17%), soybean straw (24.08%), sugarcanebagasse(21.33%)andricestraw(22.75%).
Theashcontentofthemushroomsobtainedinthisexperiment ranged between 5.84% (T2) and 7.46%(T6)(Table 3).ThesevaluesaresimilartothosegivenonaveragebyRodriguezBarreal(1987)(7.50%)andBenavides et al. (2009) (5.55%). Other researchers,suchBasaketal. (1996),Manzietal. (1999),Wangetal.(2001)andShashirekhaetal.(2002)expandedthe range of these contents between 6.70% and15.40%.Baena(2005),workingwithgreenmangueybagasseinmushroomsofP. ostreatus,gotashcontentvalues rangingbetween3.77%and8.73%.Siwulskietal.(2009),alsoworkingwithP. ostreatus,butwithaldersawdustsubstratessupplementedwithleavesoftwodifferentspeciesofGinkgo bilobaL.,concludedthat the addition ofG. biloba leaves does not affecttheashcontentoffruitingbodies.
Pardo et al. (2005a), on research carried out inCastillaLaManchawithP. ostreatusobtainedsimilarashcontentvaluesinmushroomstothisexperiment,depending on the type of substrate, treatment, andmyceliumused (from6.83%to7.70%). In thesamerangeofP. ostreatus ash content values,Pardo, in alaterwork(Pardoetal.,2007),achieveddifferentashvaluesinthecarpophores(from6%to7.14%).
Inanotherstudy,alsowithP. ostreatus, thesameresearchers (Pardo et al., 2008) obtained differentvalues for ash content depending on the type ofsubstrate, treatment, and weight of the bag used(from 5.80% to 7.50%). These authors found nosignificantdifferencesbetweenthedrymattercontentandashof fruitingbodiesobtainedfromthevariouscombinations, although theynoticeda lowerproteincontentinthemostproductivemixtures.
Similarly, slightly higher values of ash contentthanthoseobtainedinthisexperimentwereachievedby Forero et al. (2008). Growing P. ostreatus onwastechili, they reachedvaluesbetween8.81%and9.84%,justifyingthesefiguresbythehighashcontentof supplied based substrates. Similar values were
obtainedbyFonsecaetal.(2009),usingamixtureofrice straw (35%), Juncus effusus L. (25%) and ricebran(40%),achievinganashcontentvalueof6.38%bygrowingP. ostreatus.
InotherPleurotusspecies,suchasP. eryngii,Manzietal.(1999)usingsubstratesbasedonwheatstraw+beet(15%),reachedashcontentvaluesbetween6.90%and10.50%.Manzietal.(2004)achievedinthesamespecies,anashcontentof1.20%(basedonthefreshproduct with 86.60% moisture).Also, in P. eryngii,Hassan et al. (2010) obtained different ash contentvalues with various substrates: sawdust (6.94%),soybean straw (7.66%), sugar canebagasse (6.54%)and rice straw (8.02%). In P. sajor-caju, Oyetayoetal.(2004)reachedashcontentvaluesof10.51%inmushroomswhencultivatedwithsubstratesofgratedears,whichdecaysto7.41%withoutsupplementation.
Inother typesof fungi, for instance,Ganoderma lucidum (Curt.: Fr.) P. Karst, the moisture, protein,and ash contents are very different from the valuesof the same parameters inPleurotus spp. Peksen etal. (2009), working with this fungus and differentcombinationsof sawdust substrates (wheatbran andteawaste)obtainedlowervaluesinthesethreequalityparametersthanthoseobtainedinthisexperimentandcollectedbytheliteratureforP. ostreatus.
In general, according to color and texture,mushrooms produced in the substrates of thisexperimentshowsimilarfeaturesachievedbyUrbanoet al. (2002), Rodriguez Valencia et al. (2005) andForeroetal.(2008).Fortheconsumer,thecolorandtexture of the mushrooms are paramount (Matseret al., 2000).Unacceptable changes in these factorscanproducealossofconsumeracceptability.
Inregardstocolor,thecommercialsubstrate(T7)hadthesmallestvalues in lightness(L)(70.90).ThebrightestsubstrateswereachievedwithT3(74.20)andT6(74.22)withT4(72.41)andT2(72.89)followingclose behind (Table 3). The substrates with thehighestvalue in the red-greencomponent (r-g)werethose that reduced theamountofSPSand increasedthe SAS until balanced. The highest values of red-greencomponent(r-g)wereachievedwithT4(2.53)andT5(2.96)whileT7(2.09)andT2(1.46)werethelowest. Similarly, these considerations are valid foryellow-blue component (y-b).The highest values ofyellow-bluecomponent(y-b)wereachievedwithT4(11.99)andT6(14.70)whileT2(11.72)andT3(9.66)werethelowest(Table 3).
Texture(hardness,cohesiveness,springiness,andchewiness)wascontrolledinPleurotusspp.cultivatedonricestraw(Kotwaliwaleetal.,2007).Accordingtotheseauthors,theincreaseinhardnessandchewiness,and the decrease in cohesion and elasticity in thesemushroomscanbeattributedtothelossofmoistureinthemushroombodies.
370 Biotechnol. Agron. Soc. Environ. 201620(3),363-374 Picornell-BuendiaR.,Pardo-GiménezA.etal..
3.3. Correlation matrix and “step by step” regression models
T1 (6,000g SPS) showed difficulties in germinationand a high degree of contamination byGliocladiumspp.Duetothesereasons,T1wasnotconsideredinthestatisticalanalysisofthisexperiment.
Table 4 presents the correlation matrix betweenproductionparametersandphysicochemicalpropertiesofthesubstratespreparedinthisexperiment.Cellulosecontentofprocessedsubstratescorrelatessignificantlywith protein and ash contents and Ce. Also, theash content and the red-green component (r-g) arepositivelycorrelatedwiththepHofthemixtures.
Table 5 presents the correlation matrix betweenproductionandqualitativeparameters,GI,theearliness,and quantitative production parameters. Higher GIvalues correspondedwith adecrease in all harvestedmushrooms qualitative parameters. Additionally,statistically significant negative correlations wereobserved with dry matter, protein content, Bs andCe. In contrast, the days from inoculation until theappearance of the first primordia are positivelycorrelatedwith all quality parameters, although onlysignificantlywiththeunitweight,ashcontentandtheyellow-bluecomponent (y-b)ofmushrooms.Finally,BEisnegativelycorrelatedwithallqualityparameters,although, only significantlywith the ash content andtheyellow-bluecomponent(y-b)ofmushrooms.
Correlatingwithotherqualityparameters,ahigherdry matter content in mushrooms corresponds withhigherprotein,Bs,andCevalues(Table 6).
Table 7presentsthe“stepbystep”regressionmodelsforqualitativeproductionparametersdependingonthephysicochemicalpropertiesofthesubstratesmade,GI,earliness,andproductioncharacteristicsofquantitativeparameters.Themodelthatbestexplainsthevariabilityofthemushroom’sproteincontentisonethatincludestheGI(R2=93.30%),although,withincreasingGI,thiscontent is reduced.Also,celluloseandhemicellulosecontents, and the yellow-blue component (y-b) ofthe mushrooms provide good models to explain thevariabilityofthisqualitativeparameter.
The best model explaining the variability of ashcontentincludethepHofprocessedsubstrates(positivecoefficient) and crude fat (negative coefficient) asindependentvariables(R2=99.00%).
MathematicalmodelsfortheCEvariabilityincludetheGI(negativecoefficient;R2=97.60%),thecellulosecontentofmadesubstrates(negativecoefficient;R2=84.20%) and dry matter (positive coefficient, R2=95.50%).
Ash and cellulose contents of the preparatedsubstratesaretwoindependentvariablesthatexplain,suitably, the variability of the red-green component(r-g) (positive coefficients), while the quantity of Ta
ble
4.CorrelationmatrixbetweenthecharacteristicsofP
leur
otusfruitingbodiesandphysicochem
icalcharacteristicsofthesubstrates—
Mat
rice
de
corr
élat
ion
entre
le
s car
acté
rist
ique
s des
fruc
tifica
tions
de Pleurotus e
t les
car
acté
rist
ique
s phy
sico
-chi
miq
ues d
es su
bstr
ats.
Cru
de p
rote
in(g. 100g
-1)
Ash
con
tent(g. 100g
-1)
Bs(N
)C
e (mJ)
Lr-
gy-
bpH
0.670(0.216)
0.951**(0.013)
0.400(0.504)
0.678(0.208)
0.205(0.741)
0.921*(0.026)
0.565(0.321)
Nitrogen
T(g.kg
-1DM)
0.735(0.157)
0.559(0.327)
0.831(0.082)
0.860(0.061)
0.277(0.652)
0.264(0.668)
0.628(0.257)
C/Nratio
-0.693(0.195)
-0.714(0.176)
-0.686(0.201)
-0.816(0.092)
-0.115(0.853)
-0.478(0.416)
-0.660(0.225)
Crudefiber(g.kg
-1DM)
-0.692(0.195)
-0.907*(0.034)
-0.404(0.500)
-0.670(0.216)
-0.422(0.479)
-0.899*(0.038)
-0.427(0.473)
Crudefat(g.kg
-1DM)
-0.827(0.084)
-0.544(0.343)
-0.845(0.072)
-0.752(0.142)
-0.472(0.423)
-0.401(0.504)
-0.834(0.079)
NFE
(g. kg-
1 DM)
-0.570(0.316)
-0.640(0.245)
-0.580(0.305)
-0.712(0.177)
0.051(0.935)
-0.411(0.491)
-0.627(0.258)
Cellulose(g. kg-
1 DM)
-0.900*(0.037)
-0.916*(0.029)
-0.794(0.109)
-0.939*(0.018)
-0.334(0.583)
-0.737(0.155)
-0.815(0.093)
NDS(g. kg-
1 DM)
-0.727(0.164)
-0.946**(0.015)
-0.483(0.410)
-0.744(0.149)
-0.294(0.632)
-0.884*(0.046)
-0.560(0.326)
BS:breakingstrength—
rési
stan
ce à
la ru
ptur
e;C
E:compressionenergy—é
nerg
ie d
e co
mpr
essi
on;L
:brightness—
lum
inos
ité;r-g:red-greencolorcom
ponents—
com
posa
nts
de c
oule
ur ro
uge-
vert;y-b:yellow-bluecolorcom
ponents—
com
posa
nts d
e co
uleu
r jau
ne-b
leu;Nitrogen
T:totalnitrogen—
azo
te to
tal;NFE
:nitrogenfreeextracts—
ext
raits
d’
azot
e lib
re;N
DS:neutraldetergent-soluble—
dét
erge
nt n
eutre
solu
ble;DM:drymatter—
mat
ière
sèch
e;resultsinparenthesesindicatestatisticalsignificance—le
s rés
ulta
ts e
ntre
pa
rent
hèse
s ind
ique
nt la
sign
ifica
tion
stat
istiq
ue;no*—
pas
de
*: p>0.05:nosignificant—
non
sign
ifica
tif;*:0.01<p≤0.05:significantat95%
—si
gnifi
catif
à 9
5 %;**:0.001<p≤
0.01:significantat99%
—si
gnifi
catif
à 9
9 %.
Agronomicassessmentofspentsubstrates 371
mushrooms produced and BE(negative coefficients) explainthe yellow-blue component (y-b)(Table 7).
4. CONCLUSIONS
According to the results obtainedfor the combinations tested, SPS(3,600g)+SAS(2,400g)andSPS(3,000g) + SAS (3,000g) wereprepared substrates that achievedacceptable crude protein contentin fruiting bodies. With these sub-strates,higherashcontent,lightness,yellow-blue (y-b) and red-green(r-g)chromaticity,breakingstrength(Bs) and compression energy (Ce)were obtained. These values werehigher than the mean values andeven higher than the commercialsubstrate. Increased SAS participa-tioninthemixtureoftheprocessedsubstrate(andtheconsequentreduc-tionofSPSparticipation)resultedinmushrooms that require higher BsandCeforthecultivationoffruitingbodies.
Consequently, these formulationbased composts degraded by thegrowth of P. ostreatus could be alow-costsubstratewithselectiveandbalanced nutrients for growth anddevelopment of oyster mushrooms.This material could be integratedthrough new formulations andmethodologies while loweringproduction costs and reducing theenvironmental impact of unusualwaste.
Bibliography
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ANKOM, 2006a. Neutral detergent fiber in feeds. Filter bag technique. ANKOM Technology method 6. Macedon, NY, USA: ANKOMTechnology.
ANKOM,2006b.Aciddetergentfiberinfeeds.Filterbagtechnique.ANKOMTa
ble 5
.CorrelationmatrixbetweenthecharacteristicsofP
leur
otusfruitingbodiesandgerminationindex,earliness,andquantitativeparam
eters—
Mat
rice
de c
orré
latio
n en
tre le
s car
acté
rist
ique
s des
fruc
tifica
tions
de Pleurotuse
t le
taux
de
germ
inat
ion,
la p
réco
cité
et l
es p
aram
ètre
s qua
ntita
tifs.
Cru
de p
rote
in
(g. 100g
-1)
Ash
con
tent
(g. 100g
-1)
Bs (N
)C
e (mJ)
Lr-
gy-
b
GerminationIndex
-0.974**(0.005)
-0.751(0.143)
-0.971**(0.006)
-0.991***(0.001)
-0.567(0.319)
-0.520(0.369)
-0.797(0.107)
1rs Flush“Seeding”
0.778(0.122)
0.901*(0.037)
0.652(0.233)
0.791(0.111)
0.044(0.945)
0.778(0.121)
0.944*(0.016)
Total“Seeding”
0.775(0.124)
0.440(0.458)
0.931*(0.021)
0.832(0.081)
0.337(0.579)
0.154(0.805)
0.769(0.129)
Totalquantityofm
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0.044(0.944)
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BE
-0.828(0.083)
-0.903*(0.036)
-0.719(0.171)
-0.844(0.072)
-0.123(0.843)
-0.758(0.137)
-0.944*(0.016)
Bs,CE,L,r-g,y-b;no*,*,**:see
tabl
e 4—v
oirt
able
au 4;B
E:biologicalefficiency(kg.100kg
-1ofdrysu
bstrate)—
effi
caci
té b
iolo
giqu
e (k
g.10
0 kg
-1 d
e su
bstr
at se
c).
Tabl
e 6.Correlationmatrixbetweenproductionparameters—
Mat
rice
de
corr
élat
ion
entre
les p
aram
ètre
s qua
litat
ifs.
Cru
de p
rote
in(g. 100g
-1)
Cru
de p
rote
in(g. 100g
-1)
1.000
Ash
con
tent
g. 100g
-1)
Ash
con
tent(g. 100g
-1)
0.818(0.091)
1.000
Bs(N
)Bs(N
)0.926(0.024)
0.577(0.308)
1.000
Ce(mJ)
Ce (mJ)
0.977**(0.004)
0.794(0.109)
0.944*(0.016)
1.000
LL
0.655(0.230)
0.282(0.646)
0.633(0.252)
0.589(0.296)
1.000
r-g
r-g
0.641(0.243)
0.946**(0.015)
0.320(0.600)
0.572(0.314)
0.184(0.767)
1.000
y-b
y-b
0.768(0.130)
0.765(0.131)
0.712(0.177)
0.752(0.143)
0.062(0.921)
0.632(0.253)
1.000
Bs,CE,L,r-g,y-b,*,**:see
tabl
e 4—v
oirt
able
au 4.
372 Biotechnol. Agron. Soc. Environ. 201620(3),363-374 Picornell-BuendiaR.,Pardo-GiménezA.etal..
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ble
7.M
odelso
btainedbyregressing“stepbystep”—
Mod
èles
obt
enus
par
la ré
gres
sion
« st
ep b
y st
ep ».
Expl
aine
d va
riab
leIn
depe
nden
t var
iabl
eEq
uatio
nR
2 cor
rect
edSE
Ashcontent
PCC+QPP+CPP
Ashcontent=-10.822**+2.138**·pH-0.299**·C
Fa+0.016**·NDS
100.00**
0.00165
Bs
PCC+QPP
PCC+QPP+CPP
PCC(-C
Fi)+
QPP+CPP(-DM)
Bs=
717.976***-67.707***·GI-10.076***·pH
+0.107**·Nºm
ushrooms
Bs=
-125.618***+36.914***·D
M+0.261***·B
s+0.135**·P2
Bs=
718.725***-71.941***·GI-9.143**·Ashcontent+0.030**·NFE
100.00***
100.00***
100.00***
0.01386
0.01269
0.03363
Ce
PCC+QPP+CPP
CE=3.409.055***–414.818***·G
I97.60***
20.93752
r-gPC
C+QPP,Q
PP+CPP
r-g=-10.242**+1.460***·Ashcontent+0.017**·cellulose+0.000*·C
E100.00***
0.00116
y-b
PCC+QPP+CPP
y-b=16.548***-0.108**·Nºm
ushrooms
92.50**
0.55873
R2 :determinationcoefficient—
coe
ffici
ent d
e dé
term
inat
ion(%
);SE
:standarderroroftheestimate—
err
eur s
tand
ard
de l’
estim
atio
n;PCC:physico-chemicalcharacteristicso
fsubstrate—
car
acté
rist
ique
s phy
sico
-chi
miq
ues d
u su
bstr
at;pH(aq.1:5,w
/w),totalnitrogen—
azo
te to
tal(g.kg
-1,odm
),ash—
cen
dres(g. kg-
1 ,odm),C/Nratio,crudefiber—
rapp
ort
C/N
fibr
es b
rute
s(C
Fi;g. kg-
1 ,odm),crudefat—
mat
ière
s gra
ssesb
rute
s(C
Fa;g. kg-
1 ,odm),nitrogenfreeextracts—
ext
raits
d’a
zote
libr
e(NFE
;g. kg-
1 ,odm),cellulose(g. kg-
1 ,odm),
neutral-detergentso
luble—
dét
erge
nt n
eutre
solu
ble(NDS;g. kg-
1 ,odm),odm:ondrymatter—
sur m
atiè
re sè
che;QPP:indexgermination,earlinessandquantitativeproduction
parameters—
inde
x de
ger
min
atio
n, p
réco
cité
et p
aram
ètre
s de
prod
uctio
n qu
antit
atifs;G
I:germinationindex—
indi
ce d
e ge
rmin
atio
n;P2:daysfrominoculationtotheformation
ofthefirstprim
ordia—
jour
s de
l’ino
cula
tion
à la
form
atio
n de
la p
rem
ière
pri
mor
dia;P4:daysfrominoculationtotheonsetofharvest—
jour
s de
l’ino
cula
tion
à l’a
ppar
ition
de
la
réco
lte;nºm
ushrooms:quantityofm
ushrooms—
qua
ntité
de
cham
pign
ons;BE:biologicalefficiency(kg.100kg
-1ofdrysu
bstrate)—
effi
caci
té b
iolo
giqu
e (k
g.10
0 kg
-1 d
e su
bstr
at se
c);
CPP:qualitativeproductionparameters—
par
amèt
res d
e pr
oduc
tion
qual
itativ
es;g:averageunitw
eightofuncutmushrooms—
uni
té d
e po
ids m
oyen
de
cham
pign
ons n
on c
oupé
s; DM:
drymatter—
mat
ière
sèch
e(g. 100g
-1) ;crudeprotein—
pro
téin
e br
ute(g. 100g
-1);ashcontents—
tene
ur e
n ce
ndre
s(g.100g-
1 );Bs,CE,r-g
,y-b:seeta
ble
4—v
oirt
able
au 4;
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