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ISSN 1330-9862 review

(FTB-1924)

Cultivation Techniques and Medicinal Properties

ofPleurotus spp.

Andrej Gregori1,2*, Mirjan [vagelj3 and Jure Pohleven4,51SRC Bistra Ptuj, Slovenski trg 6, SI-2250 Ptuj, Slovenia

2Institute for Natural Sciences, Ulica bratov U~akar 108, SI-1000 Ljubljana, Slovenia3Department of Chemical, Biochemical and Ecology Engineering, Faculty of Chemistry and

Chemical Techmology, University of Ljubljana, A{ker~eva 5, SI-1000 Ljubljana, Slovenia

4Department of Pharmaceutical Biology, Faculty of Pharmacy, University of Ljubljana,A{ker~eva 7, SI-1000 Ljubljana, Slovenia

5Department of Biotechnology, Jo`ef Stefan Institute, Jamova 39, SI-1000 Ljubljana, Slovenia

Received: May 5, 2007Revised version: May 20, 2007

Accepted: June 1, 2007

Summary

The genus Pleurotus (oyster mushroom) comprises some most popular edible mush-rooms due to their favourable organoleptic and medicinal properties, vigorous growth andundemanding cultivation conditions. It can be cultivated on log and a wide variety of

agroforestry (by-)products, weeds and wastes for the production of food, feed, enzymesand medicinal compounds, or for waste degradation and detoxification. Many differenttechniques and substrates have been successfully utilized for mushroom cultivation andbiomass production by means of solid-state and submerged liquid fermentation. However,in contrast to submerged liquid fermentation, solid-state fermentation is not often used inlarge scale due to severe engineering problems. VariousPleurotusspecies have been shownto possess a number of medicinal properties, such as antitumour, immunomodulatory, anti-genotoxic, antioxidant, anti-inflammatory, hypocholesterolaemic, antihypertensive, antipla-telet-aggregating, antihyperglycaemic, antimicrobial and antiviral activities. These thera-peutic activities are exhibited by extracts or isolated compounds from Pleurotusspp. fer-mentation broth, mycelia and fruiting bodies. In particular, polysaccharides appear to bepotent antitumour and immuno-enhancing substances, besides possessing other beneficialactivities. However, the biochemical mechanisms of these therapeutic activities still remainlargely unknown. This review focuses on recent advances in the biotechnology ofPleurotusspp., with emphasis on the production of fruiting bodies, the production of mycelium andbioactive compounds by solid-state and submerged liquid fermentation. The medicinal pro-perties of this mushroom are also outlined.

Key words: Pleurotus, mushroom cultivation, biomass production, solid-state fermentation,submerged liquid fermentation, medicinal properties

Introduction

Cultivation of the oyster mushroom, Pleurotus spp.,has increased greatly throughout the world during the

last few decades (1,2); in 1997 it accounted for 14.2 % ofthe total world edible mushroom production (1). Its popu-larity has been increasing due to its ease of cultivation,high yield potential and high nutritional value (3). Al-

238 A. GREGORIet al.: Cultivation ofPleurotus spp.,Food Technol. Biotechnol. 45(3) 238249 (2007)

*Corresponding author; Phone: ++386 40 756 146; E-mail: [email protected]

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though commonly grown on pasteurized wheat or ricestraw, it can be cultivated on a wide variety of lignocel-lulosic substrates, enabling it to play an important rolein managing organic wastes whose disposal is problem-atic.

New technologies and production techniques arebeing constantly developed as the number of requiredcontrollable environment parameters increases (4). Cur-rently, solid-state fermentations, other than fruiting bodyproduction with Pleurotus spp., are used either in thetransformation of wastes into animal feed or for enzymeproduction. Submerged liquid fermentation can, on theother hand, provide more uniform and reproducible bio-mass and can prove interesting for valuable medicinalproducts or for enzyme production because of uncom-plicated downstream processing (5). Current research onPleurotus spp. related to solid-state and submerged li-quid fermentation is mainly concerned with substratecomposition and optimization of culture parameters.

Pleurotusspecies have been used by human culturesall over the world for their nutritional value, medicinalproperties and other beneficial effects. Oyster mushroomsare a good source of dietary fibre and other valuable nu-trients. They also contain a number of biologically activecompounds with therapeutic activities. Oyster mushroomsmodulate the immune system, inhibit tumour growth andinflammation, have hypoglycaemic and antithromboticactivities, lower blood lipid concentrations, prevent highblood pressure and atherosclerosis, and have antimicro-bial and other activities (6). Recent studies of the medic-inal properties of oyster mushrooms have focused onisolated bioactive compounds; however synergistic ef-

fects of the constituents of mushroom extracts may bepossible.

Pleurotusspp. Fruiting Body Production

Substrates for oyster mushroom cultivation

Pleurotusspp. cultivation is a very simple procedurein the case of log cultivation because it does not involvesophisticated equipment. However, despite its simplic-ity, large-scale cultivation on natural logs is not oftenused due to long incubation periods, low yields and en-vironment-dependent production if conducted outdoors.Yields ofP. ostreatus fruiting bodies vary with the spe-

cies of trees used and range from 21 % biological effi-ciency (BE) for beech wood to 3 % BE for alder wood(7).

Broadleaf hardwood sawdust and straw-based sub-strates with added supplements are more often used incommercial production. In this case, these artificial sub-strates must be pretreated, mainly for elimination of con-taminants, and handled in a clean environment. Thereare different methods of cultivation like shelf, bag, bot-tle, tray, jar, grid-frame, wall-frame and others (8). Inpractice, the most used are bag, bottle and shelf cultiva-tion (9). Evaluation ofP. columbinuscultivation in differ-ent bagging systems, in which partially pasteurized of-

fice papers were used as a growing substrate, revealedthat polyethylene bags resulted in 109 % BE, followedby pottery (86 %), plastic trays (72 %) and polyester net(56 %) (10). Removal of the bottom half of the plasticcultivation bag and embedding artificial logs vertically

in soil resulted in a BE of up to 123 % and proved to bethe optimal method for P. nebrodensis cultivation (11).

Pleurotusspp. can also colonize and produce mush-rooms on pretreated conifer (Pinusspp.) wood chips butthey do not always readily colonize non-pretreated coni-fer wood, due to the presence of inhibitory components(12). Some strains can, however, be adapted for cultiva-tion on conifer-sawdust-based substrates (13). Pleurotusspp. can also be cultivated on wood waste or unusedwood residues associated with harvesting or thinningoperations, which can enhance economic returns neededto support ecosystem management (14).

Some pretreatment or supplementation with nutri-ents may be necessary.P. ostreatusBE is much lower whenit is cultivated on fresh sawdust than on composted saw-dust/bran mixture (15). Rodriguez Estrada and Royse(16) reported that P. eryngii fruiting body yields weresignificantly higher in substrates containing Mn (50 mg/g)and soybean than in the basal cottonseed hull/sawdustsubstrate.

Different types of straw are commonly used for Ple-urotus spp. cultivation. Straw can be composted or pas-teurized and extra additives can be used to increase theBE. When using rice and wheat straw for P. sajor-cajucultivation, higher yields were obtained on ground thanon chopped straw, and yields were 10 % higher on ricethan on wheat straw. Higher spawn levels enhanced mush-room yields (17). Rice straw appeared to be the best sub-strate for P. ostreatus mushroom cultivation when com-pared to banana leaves, maize stover, corn husks, ricehusks and elephant grass (15). When cultivating P. flo-rida, the incorporation of cotton seed powder into ricestraw substrate enhanced mushroom yield, increased netand total protein, free amino acids and total lipids con-tent, while there was a significant decrease in total di-etary fibre, free sugars and polymeric carbohydrates (18).Supplementation of rice straw with a residual slurry ob-tained after production of biogas from manure improvedthe yield potential and increased protein and mineralcontents ofP. sajor-cajumushrooms (3). Wheat straw sup-plemented with Lolium perenne grass chaff stimulatedfructification and mushroom yield ofP. pulmonarius(19).P. tuber-regium strains from Australasian-Pacific regionsshowed faster mycelium growth rates when cultivatedon wheat straw, while wild Nigerian strains performed

better in sclerotia yield when cultivated on this substrate(20).

Cultivation ofPleurotus spp. on substrates contain-ing added olive mill waste and wastewaters (OMWW)can be a viable alternative for converting these environ-mentally problematic materials into valuable, highly nu-tritious food. It has been shown that wetting a wheatstraw and bran substrate with OMWW diluted in tapwater (25 %) had no significant negative effect on thetime required for mycelial colonization, primordium ini-tiation or mushroom yield ofP. sajor-caju and P. citrino-

pileatus. Application of 50 % OMWW led to a delay incolonization and reduction in yield, and deleterious ef-

fects were noted when using 75 % OMWW (21). Sub-strates with the addition of OMWW up to 30 % did notinterfere with mycelial growth ofP. pulmonariusbut theydid inhibit fruiting body formation. OMWW up to 10 %,however, did not inhibit pinhead appearance (22). Expe-

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riments with wild and commercial strains ofP. ostreatus,P. eryngii and P. pulmonarius demonstrated significantlyhigher colonization rates on wheat straw and cotton wastethan on peanut shells. Faster colonization was achievedon non-composted than on composted wheat straw andcotton waste substrates. Cellulose/lignin ratios in sub-strates were positively correlated to mycelial growth ratesand mushroom yields ofP. ostreatusandP. pulmonarius.In addition, there was a positive correlation between theC/N ratio and P. eryngii mushroom yield (23). A sub-strate composed of coffee pulp and Digitaria decumbenswas also used for P. ostreatuscultivation and additionalcomposting of this mixture improved the BE (24). Whenusing a mixture of coffee pulp and wheat straw for P. dja-mor, P. ostreatus and P. pulmonarius cultivation, the ob-served decrease in caffeine content of the coffee pulp sam-ples during the fruiting stage suggests that some caffeineaccumulates in the fruiting bodies (25).

Much effort has been put into optimizing substratesbased on different grass species for Pleurotusspp. culti-vation. This is an effective way of converting abundantbut low value materials into highly nutritional food, es-pecially where wood and straw are scarce (26). Lolium

perenne grass chaff stimulated fructification and mush-room yield ofP. pulmonarius (19). Wooden crates wereused, in a very simple substrate preparation forP. ostrea-tus, for composting a mixture of grass (Digitaria decum-bens) and coffee pulp; 60 and 93 % BE were obtained intwo harvests. Further composting for two to three daysin each case improved the BE (24). When using chop-ped, pasteurized switch grass (Panicum virgatum) andpasteurized cottonseed hulls with wheat straw forP. cor-

nucopiaecultivation, higher yields were obtained on cot-tonseed hulls/wheat straw substrate. Increasing spawnand supplement levels in switch grass/wheat straw sub-strate stimulated yield in a linear fashion. However, ma-ximum yields were less than 50 % of those obtained withcottonseed hulls/wheat straw substrate (27). When sup-plementing spent rice straw substrate with oil seed cakes,cottonseed powder proved best in enhancing P. sajor--caju mushroom yields. Mushrooms grown on substratesupplemented with cottonseed powder had increasedprotein and fat content, decreased carbohydrate contentand contained no residues of gossypol. In addition, therewas a significant reduction in the spawn run period when

compared to the use of unsupplemented rice straw (28).P. ostreatusandP. sajor-cajuexhibited higher ash con-

tent when cultivated on rice straw than when cultivatedon banana straw, and P. sajor-caju also showed highermoisture and fibre content when cultivated on rice straw(29). When cultivatingP. ostreatuson corn and pumpkinstraw, the substrate had no effect on the nitrogen con-tent and amino acid profile of the fruiting bodies; how-ever, the nitrogen content increased from the first har-vest to the third harvest (30). Using water hyacinth biomassas substrate, BE was found to be higher with P. florida(86 %) than with P. citrinopileatus (79 %) (31). When P.ostreatusvar.salignuswas cultivated on peanut, soybean,

sorghum or wheat straw, the highest and lowest yieldswere obtained on peanut and sorghum straw, respec-tively. The highest protein content, pileus/stipe ratio,sporophore mass, percentage of dry material, and nitro-gen and carbon contents were obtained with peanut

straw. Sorghum resulted in the lowest mushroom massand pileus/stipe ratio, whereas the lowest protein andnitrogen content, and dry material mass were obtainedwith wheat straw (32). Among different agrowastes tested(cotton stalk, coir fibre, sorghum stover and mixtures of

these wastes), the maximum yields ofP. sajor-caju andP. citrinopileatus were obtained on cotton stalks, whileP. platypusyields peaked on sorghum stover (33).

Weed plants (Leonotissp., Sida acuta, Parthenium ar-gentatum, Ageratum conyzoides, Cassia sophera, TephrosiapurpureaandLantana camara) without heat pretreatmentwere tested forP.ostreatuscultivation.Leonotissp. mixedwith rice straw was the best substrate for spawn runand mushroom cultivation whileT. purpureawas the leastappropriate. The main problem ofP. ostreatuscultivationon weed-composed substrates was the low yield thatwas obtained in the second flush. This problem could beovercome by mixing weed plants with rice straw. Thefruiting body protein content was higher when P. ostrea-tuswas cultivated onCassia sophera, Parthenium argenta-tumandLeonotissp. than on weeds supplemented withrice straw or on rice straw alone (34).

When hazelnut, Tilia spp., European aspen leaves,wheat straw, sawdust and waste paper were used assubstrates, the best major component and substrate com-bination for mushroom productivity were wheat strawand wheat straw in combination with waste paper. Mix-tures involving waste paper generally produced higheryields than other combinations. The lowest yield andthe smallest fruiting body diameters were obtained from

Tilia spp. and European aspen leaves in combinationwith sawdust. The greatest number of fruiting bodieswas obtained on a mixture of wheat straw, hazelnutleaves and waste paper (35). Mandeel et al. (10) culti-vated Pleurotus spp. on various lignocellulosic wastessupplemented with fresh chicken manure. The highestBE was noted on cardboard with both P. columbinus(134%) and P. ostreatus (117 %). Experiments conducted byBaysalet al.(36), which involved cultivation ofP. ostrea-tus on waste paper with addition of chicken manure,peat and rice husks, showed that increasing the amountof rice husks added to the substrate accelerated spawnrunning, pinhead formation and fruiting body forma-

tion. Larger proportions of peat and chicken manure hada negative effect on growth. A study on growth and pro-ductivity of different P. ostreatus strains on sunflowerseed hulls supplemented with NNH4+ or Mn, showedthat the addition of growth-limiting mineral nutrientsincreased the mycelial growth rate. BE increased overcontrol values and reached 60112 %, depending on thestrain and the concentration of Mn and NNH4+ (37).

Non-pretreated spent brewery grains were success-fully used as a basic substrate material for P. ostreatuscultivation in polypropylene bottles. Few fruiting bodieswere formed on spent grain alone; however 19 % BE

was obtained with the addition of 45 % wheat bran. Thechemical analysis of fruiting bodies indicated that P. os-treatuscultivated on spent grain substrate had a highernutritional value than those grown on other types of sub-strates (38).


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Other factors influencing oyster mushroomdevelopment

There have been various reports on other factors thatinfluence the development of oyster mushrooms. Noneof these factors has been studied in depth; the various

observations that have been made are outlined in the pa-ragraphs below.Cho et al. (39) discovered that inoculation of pure

P. ostreatusmycelium cultures with strains of fluorescentPseudomonas spp., isolated from the mycelial plane ofcommercially produced mushrooms, promoted the for-mation of primordia and enhanced the development ofthe basidiomata. These results strongly suggest that in-oculation of the mycelium with specific bacteria mayhave beneficial applications for mushroom production.

It was reported that the bacterial blotch disease inmushrooms caused by Pseudomonas tolaasii was more se-vere when substrates were amended with Cu in P. eryngii

cultivation (16). Two compounds from olive mill waste,4-methylcatechol and catechol, were found to be effectiveagainst P. tolaasii and supplementation with up to 10 %OMWW reduced bacterium-related symptoms (22).

Qu et al. (40) demonstrated the influence of heavymetals in substrates on P. eryngii primordial formation,fruiting body development and BE. Heavy metal (As,Hg and Cd) supplementation decreased average growthyields and BE ofP. eryngii, whereas Pb supplementationimproved both parameters.

Irradiation by red and green light stimulated vege-tative growth ofP. ostreatusmycelium and shortened thesubstrate colonization and fructification time. The in-

creased fruiting body yield in irradiated cultures reached3651 % (41).

The cytolytic protein ostreolysin, isolated from P. os-treatusfruiting bodies, was specifically expressed duringfruiting initiation, suggesting its involvement in fruitingbody formation. When purified ostreolysin was used asa supplement on nutrient media plates inoculated withP. ostreatus mycelium, the protein stimulated primordiaand fruiting body formation (42).

Solid-State Fermentation with Pleurotus spp.

Most solid-state fermentations (SSF) with Pleurotusspp. have been small scale, that is, they have involvedsolid substrate in Erlenmeyer flasks or in agar plates.Recent studies have been focused on the utilization oflignocellulosic organic waste materials for either lignindegradation, use as animal feed or enzyme production.

SSF for Pleurotus spp. mycelial biomass andmushroom production

When agar-based media are used, medium compo-sition plays an important role as it determines the my-celial growth rate, and correct medium ingredient ratiosare crucial for fast colonization. Nasim et al. (43) found

that malt extract agar (MEA) provided faster P. ostreatusmycelial growth rates than did Murashige and Skoogs(MS) medium and potato dextrose agar (PDA). The slow-est growth was observed on PDA medium. The most ef-fective carbon, nitrogen and inorganic salt supplements

for vigorousP. nebrodensisgrowth were glucose, peptoneand MgSO4 (11).

Mycelium production on lignocellulosic substrateshas also been investigated. Amongst seven mushroomcultivation substrates, the mycelial extension rates were

highest on cotton gin-trash, peanut shells and poplarsawdust. Supplemented oak sawdust and olive millwaste were poor substrates for most species examined,while almost all strains performed adequately on corncobs (44,45). Different substrate combinations were eval-uated for P. ostreatus mycelial growth and favourablecombinations were Tilia spp. leaves with wheat branand Populus spp. leaves with wheat straw. The authorsalso reported that mycelial growth and mushroom yieldhave different requirements (46), whereas others re-ported that mushroom yield was directly related to thespread of the mycelium within the substrate (15).

SSF waste utilization and enzyme production byPleurotus spp.

Much work has been done recently on waste utiliza-tion byPleurotusspp., mostly concerned with the poten-tial of locally available waste and agricultural by-prod-ucts for either transformation into animal feed or asprimary substrates for enzyme production.

Villas-Bas et al.(47) investigated the conversion ofapple pomace by Candida utilis and P. ostreatus, sepa-rately and in coculture, in SSF.C. utiliswas a better can-didate for biological treatment of apple pomace as theyeast lowered the residual free sugar concentration morethan P. ostreatus did. When both organisms were used,

apple pomace digestibility decreased while the proteincontent increased, with the final product being suitablefor use as a protein supplement for cattle feed. Fermen-tation withP. ostreatusalone was, however, discarded asa viable treatment of apple pomace. Furthermore, recy-cling of viticulture waste in SSF with P. ostreatus andP. pulmonariusalso yielded a high-fibre feed for limiteduse in ruminants (48). Tests on water hyacinth biomassdelignification showed that organic matter loss washigher in samples inoculated with P. citrinopileatusthanP. florida. Improvement of delignification and dry matterdigestibility was, however, higher withP. florida, whichalso proved to be more effective thanP. citrinopileatusforthe production of highly digestible mycoprotein-rich ru-minant feed (31). Intensive P. ostreatus mycelial growthwas observed in substrates containing leftover hempseeds, whole ground corn plant or sesame oil press cake.Intermediate growth was observed in substrates con-taining olive mill waste, rape oil press cake or sunfloweroil press cake while slowest growth rates were observedin the substrate containing soy oil press cake (44,45).Ghibomet al. (49) introduced a novel approach for uti-lizing whey permeate withP. ostreatusin an SSF system.They concluded that whey permeate could provide a vi-able substrate for production ofP. ostreatus mycelium.The optimal growth conditions were 44 g/L of lactose,pH=6.0 and 24 C.

White-rot basidiomycetes are efficient decomposersof lignocellulose, due to their capability to synthesizerelevant hydrolytic and oxidative extracellular enzymes.Lignocellulolytic enzymes have significant potential ap-plications not only in the chemical, fuel, food, textile,


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laundry and pulp and paper industries but also in agri-culture and for animal feed production (50).

Hlker and Lenz (4) introduced a trickle-film fer-mentation process on sugarcane bagasse withP. ostreatusfor laccase production, which turned out to be highlysuitable for extracellular enzyme production. High en-zyme levels were produced when the growth mediumwas repeatedly changed. When compared to submergedliquid fermentation, the productivity was better and twolaccase isoforms were detected.

Banana leaf waste was a better substrate than ba-nana pseudostem waste in the production of extracellu-lar enzymes byP. ostreatusandP. sajor-cajuin SSF and isa potential alternative to other agrowaste substrates. Theyields were, however, too low and commercially not vi-able. It was suggested that a larger surface area of ba-nana leaf waste could be a determining factor for betterenzyme production (51). This is in agreement with Zhanget al.(17) who reported thatP. sajor-cajugrew faster and

provided better yields on ground straw than on choppedstraw. There is, however, a substrate particle size limit,as more finely ground straw inhibited growth. Optimalparticle sizes should therefore be determined for all ap-plications.

Staji} et al. (52) performed SSF with Pleurotus spp.on grapevine sawdust, which was supplemented withsynthetic medium to provide nitrogen and trace miner-als. Peroxidase activity was detected in all strains evalu-ated, the highest being with P. ostreatus and P. pulmo-narius. The highest laccase activity was detected at 10days of fermentation inP. ostreatusfollowed byP. pulmo-nariusandP. eryngii.Organic nitrogen sources have beenshown to stimulate enzyme production more than inor-ganic sources. Cyanobacterial biomass was used as a ni-trogen supplement and stimulated P. ostreatusgrowth andlaccase production in SSF. The authors concluded thatdry biomass of diazotrophic cyanobacteria not only helpsto maintain an optimal C/N ratio but also confers agood porosity, which sustains the oxygen supply withinthe matrix of solid particles (53). The nature of the sub-strate as well as the cultivation method affects the ex-pression of lignocellulolytic enzymes. The study con-ducted by Elisashviliet al.(54) revealed that SSF of treeleaves by Pleurotus spp. was favourable for laccase andmanganese peroxidase (MnP) production. Furthermore,coculturing can be an effective method for biopulping

and improvement of lignin degradation (55,56). Chiet al.(56) demonstrated that coculturingP. ostreatuswithCeri-poriopsis subvermisporasignificantly stimulated lignin de-gradation when compared to monocultures. Laccase pro-duction and MnP activity were stimulated in coculturesofP. ostreatuswithC. subvermisporaor Physisporinus rivu-losusand a change in the isoform composition of thoseenzymes was also observed.

These studies show that the cultivation method canhave drastic effects on the production of valuable sub-stances by Pleurotusspp. and its economical feasibility.

Submerged Liquid Fermentation with

Pleurotus spp.

Submerged liquid fermentation (SLF) techniqueshave been developed for a variety of fungi and are usedin mycelium propagation for different applications, such

as liquid spawn for fruiting body production on solidsubstrates; biomass production for food, dietary supple-ment and pharmaceutical applications; and conversionof waste biomass and enzyme production. SLF offers thepossibility of high biomass production in a compactspace, shorter time and with fewer chances of contami-

nation (57,58). While SSF will remain the chosen methodfor mushroom production, there will be a continued in-crease in the development of SLF technology to producemore uniform and reproducible biomass of medicinalfungi. Western biotechnology companies have yet to rec-ognize the potential in this area of medical bioscience(5).

SLF for Pleurotus spp. biomass and polysaccharideproduction

The most detailed study ofP. ostreatusgrowth in SLFwas conducted by Mrquez-Rochaet al.(59). They stud-

ied P. ostreatus cultivation in a stirred tank bioreactorand revealed that by varying impeller geometry andspeed, and aeration intensity, the growth rate and pelletsize changed. A clear tendency was observed for smallerpellet sizes to result in higher specific growth rates. Forpromotion of mycelium growth the pellets need to bebroken down, but on the other hand, a balance betweengrowth and hyphal fragmentation must also be achieved.

The lag phase forP. tuber-regiumgrowth and biocon-version efficiency in SLF was shorter with glucose andfructose than with maize starch. In scaled-up fermenta-tions, addition of fructose to basal medium supportedhigher mycelial yields than the addition of glucose. Yeast

extract as the nitrogen source proved better than pep-tone when monosaccharides were used as the sole car-bon source (60).

Recent studies on mushroom polysaccharides havedemonstrated many interesting biological activities, whichare described later in this review. The production ofPleu-rotusspp. mycelial biomass and valuable polysaccharidesin SLF depends on the species used, growth parameters,growth timing and their nutritional requirements (61,62).Response surface methodology, a widely known optimi-sation procedure, was used to optimise the medium inorder to maximize growth and polysaccharide produc-tion byP. citrinopileatus. The highest polysaccharide yield

was obtained with a C/N ratio of 40, an initial pH=5.5and a cultivation temperature of 25 C (63). Another pa-rameter that influenced growth and polysaccharide pro-duction by P. ostreatus in SLF was the initial oxygentransfer rate (KLa). Better polysaccharide yields were ob-tained with a lower initial KLa (64).

Waste utilization and enzyme production byPleurotus spp. in SLF

SLF is also suitable for enzyme production and wastebioconversion. Most recent work is focused on substrateoptimization for maximal production of hydrolytic andoxidative ligninolytic extracellular enzymes.

Staji}et al.(52) performed SLF withP. eryngii,P. os-treatusand P. pulmonariuson different carbon and nitro-gen sources. In the medium with the best carbon sources(mandarin peels and grapevine sawdust), bothP. eryngiiandP. ostreatusshowed the highest laccase activity with


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(NH4)2SO4 as the nitrogen source. With P. ostreatus andP. pulmonarius, the best nitrogen sources for productionof peroxidases were peptone and NH4NO3. The correctcomposition of the cultivation medium is important forgood enzyme production and various inducers have beentested for stimulatingP. ostreatusenzyme production. Hou

et al. (65) obtained high levels of laccase activity usingcellobiose and peptone as the carbon and nitrogen sour-ces. 2,2-Azino-di-(3-ethylbenzothialozin-6-sulphonicacid) (ABTS) and Cu had a positive effect on laccase pro-duction and the former was shown to be the best laccaseinducer in their study. Static cultures were superior toagitated cultures in terms of growth and laccase produc-tion, while nitrogen-limited culture media were only be-neficial for laccase production. In another study, theTaguchi orthogonal array experimental design was ap-plied for the optimization of laccase production in SLFwith P. ostreatus. The most influential factor for laccaseproduction was found to be the inducer 2,5-xylidine, fol-

lowed by glucose concentration, wheat bran content, ureaconcentration, inoculum size, yeast extract concentrationand pH (66). Mikiashviliet al.(67) reported that the high-est MnP and laccase activities were obtained in xylan--supplemented media, but enzyme activities decreasedwhen supplemented with inorganic nitrogen sources.Peptone, followed by casein, was used as the nitrogensource for the best laccase accumulation; this results be-ing attributable to higher biomass production. MnP andperoxidase secretion was stimulated by supplementationwith casein hydrolysate. A liquid medium composed ofmandarin peels and leaves inoculated withP. dryinusal-lowed the simultaneous production of hydrolases and oxi-

dases at high levels. Carboxymethyl cellulase and xyla-nase appeared to be inducible enzymes. Addition of Mnenabled the laccase to MnP ratio to be regulated. For MnPproduction the presence of lignocellulosic substrates is aprerequisite and shows a positive correlation with theiraddition (50). This is supported by Hou et al.(65), whodid not detect any MnP activity in N limited glucose me-dium withP. ostreatus.Pleurotusspp. SLF on tree leavesprovided better hydrolytic enzyme production than SSF(54).

P. ostreatusmycelium could be cultivated employingcoffee pulp wastewater extract supplemented with glu-cose. The polluting load was reduced by more than 50

% at the end of 20 days of fermentation (68). SLF withP. ostreatus was also used for removing phenolic com-pounds from olive oil mill wastewater (OMWW). Lac-case was the sole ligninolytic enzyme detected and wasproduced during primary growth. The phenolic contentand toxicity were successfully reduced, but only whenhigh OMWW dilutions were used. This could be a valu-able method for problematic OMWW treatment beforeits release into the environment (69).

Medicinal Properties of Pleurotus spp.

Recent studies on various Pleurotus species have

shown a number of therapeutic activities, such as anti-tumour, immunomodulatory, antigenotoxic, antioxidant,anti-inflammatory, hypocholesterolaemic, antihyperten-sive, antiplatelet-aggregating, antihyperglycaemic, antimi-crobial and antiviral activities.

Antitumour activity

Much recent research has been carried out onPleurotusspp. extracts and isolated compounds such aspolysaccharides, proteins and other substances that pos-sess antineoplastic activities in vitro and in vivo.

Various crude extracts ofPleurotusspecies have beenshown to possess relatively strong antitumour activities.Methanol extracts ofP. floridaandP. pulmonariusfruitingbodies significantly reduced solid tumours in mice (70,71).P. ostreatus mycelium extract, alone and combined withthe chemotherapeutic agent cyclophosphamide, inhibitedin vivo tumour growth in mice. The combined adminis-tration of the extract with cyclophosphamide decreasedthe degree of leukopenia compared to administration ofcyclophosphamide alone (72). A water extract ofP. ostre-atusexhibited the most significant cytotoxicity by induc-ing apoptosis of human carcinoma cells, when compared

to many other mushroom extracts. It has been suggestedthat the active compounds in the extract were water-sol-uble proteins or polypeptides (73).

Among the components of such extracts, polysaccha-rides are well-documented as potent antitumour andimmunomodulating substances (74,75). Many polysac-charides from Pleurotus spp. have been isolated andidentified (7686). For some of them, important medici-nal properties, including antitumour activities, have beenshown. P. tuber-regium polysaccharides, extracted frommycelium and fruiting bodies, effectively inhibited solidtumour proliferation in mice. Antitumour effects have alsobeen shown on different human tumour cell lines (87,88).

Wong et al.(89) showed thatP. tuber-regium polysaccha-rides exerted antitumour activity, through cytotoxicity andantiproliferative activity, against human leukaemia cellsin vitro. The polysaccharides induced apoptosis and causedcell cycle arrest. Compared to native P. tuber-regium po-lysaccharides, their corresponding carboxymethylated orsulphated derivatives showed higher antitumour activ-ity, presumably because of their higher water solubilityand relatively extended flexible chains (9093). A novela-glucan from P. ostreatus mycelium induced apoptosisof colon cancer cellsin vitro(94) and water-soluble poly-saccharides extracted fromP. citrinopileatusfermentationbroth have been shown to reduce the number of meta-

static tumour nodules in tumour-bearing mice (63).Antitumour properties have also been demonstrated

for Pleurotus spp. proteins, proteoglycans, and DNA. Alectin isolated fromP. ostreatuspotently inhibited growthof sarcoma and hepatoma in mice and prolonged theirlifespan (95). AP. eouslectin exerted antiproliferative ef-fects on human tumour cell lines while showing no cy-totoxicity (96). Furthermore, two ribonucleases isolatedfrom P. sajor-caju and P. ostreatus fruiting bodies exhib-ited antiproliferative effects on tumour and leukaemiacell lines (97,98). Another protein, eryngeolysin, isolatedfrom P. eryngii fruiting bodies, exhibited cytotoxicityagainst leukaemia cells (99). Water-soluble proteoglycans

were purified fromP. ostreatusmycelium and exerted anti-tumour activity in sarcoma-bearing mice. Proteoglycansinjected into mice reduced the number of tumour cellsby cell cycle arrest (100). Moreover, DNA isolated fromP. ostreatus fruiting bodies administered to mice with


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solid Ehrlich carcinoma significantly increased the life-span of mice (101).

Immunomodulatory and antimitogenic activitiesThe antitumour effects of mushrooms are mostly at-

tributed to stimulation of the immune response. Re-cently, several compounds from Pleurotus species withimmunostimulatory activities on humoral and cell-medi-ated immunity have been isolated. Water-soluble poly-saccharides extracted from P. citrinopileatus fermentationbroth administered to mice resulted in a significant in-crease in the number of macrophages, T, CD4+ and CD8+

cells (63). Glucans isolated fromP. florida fruiting bodiesactivated the phagocytic response of mouse macropha-ges in vitro (83) and significantly induced the prolifera-tive response as well as phagocytic activity of fish leu-kocytes in vitro (102). Moreover, proteoglycans from P.ostreatusmycelia exerted immunomodulatory effects by

elevating mouse natural killer cell cytotoxicity and bymacrophage stimulation (100). DNA isolated from P.ostreatus fruiting bodies stimulated mouse natural killercytotoxic activity in vitro (101).

Antimitogenic effects ofPleurotusspp.-derived com-pounds on immune cells have also been reported. A ri-bonuclease isolated fromP. sajor-caju fruiting bodies ex-erted antiproliferative effect on murine splenocytes (97),while eryngeolysin from P. eryngii inhibited the stimu-lated mitogenic response of murine splenocytes (99). Fur-thermore,P. flabellatus lectin did not exhibit any mito-genic activity towards mouse T cells (103).

Antioxidant and gene protective activitiesAntioxidant compounds prevent oxidative damage

related to aging and diseases, such as atherosclerosis, di-abetes, cancer and cirrhosis. Mushrooms that contain an-tioxidants or increase antioxidant enzyme activity maybe used to reduce oxidative damage in humans (104).

Of 89 mushroom species tested, an extract fromP. cornucopiaepossessed the most effective antigenotoxicand bio-antimutagenic activities when tested on Salmo-nella typhimuriumand Escherichia coli(105). Furthermore,P. cornucopiaeextracts significantly reduced H2O2-inducedDNA damage in Chinese hamster lung cells (106) andP. ostreatus extract mitigated genotoxicity, as shown by

the fact that it suppressed DNA damage induced by va-rious mutagens in the Drosophila DNA repair test (107).On the other hand, a water extract ofP. sajor-caju fruit-ing bodies had no genoprotective effects since it did notprevent H2O2-induced oxidative damage to cellular DNA(108).

Methanol extracts of P. ostreatus and P. cystidiosusfruiting bodies possessed antioxidant, reducing power,radical scavenging and iron chelating activities that werehigher than those of other commercial mushrooms (104).On the other hand, Elmastaset al.(109) and Dubostet al.(110) reported that oyster mushroom extracts possessedonly moderate antioxidant activities compared to other

edible mushrooms. The antioxidant activity was posi-tively correlated with total polyphenol content. Further-more, Lee et al. (111) showed that P. citrinopileatus ex-tracts prepared from fruiting bodies were more effectivethan those from mycelium and fermentation broth fil-

trate, presumably due to a higher amount of total phe-nols. Methanol extracts ofP. florida and P. pulmonariusfruiting bodies showed similar antioxidant activities(70,71), and an ethanol extract fromP. citrinopileatusfrui-ting bodies had antioxidant activities comparable to thosefrom P. eryngii, P. ferulae and P. ostreatus mushrooms

(111,112).P. citrinopileatus fruiting body extracts have shown

antioxidant activities in vitro and in hyperlipidaemichamster rats. Extracts added to a high-fat diet increasedthe activities of antioxidant enzymes in rats (113).P. ostrea-tus mushroom extracts had antioxidant properties inaged and CCl4-induced liver damaged rats, as indicatedby significant increases in concentrations of antioxidantsand antioxidant enzymes (114,115). Pleuran, a b-glucanisolated fromP. ostreatus, had a positive effect on the an-tioxidant status of rats and decreased precancerous le-sions induced in rat colon (116). A polysaccharide-pep-tide complex isolated from P. abalonus fruiting bodiesprolonged the lifespan of senescence-accelerated mice.Gene expression of antioxidant enzymes was up-regu-lated and consequently their activities were increased(117).

Anti-inflammatory activityJoseet al. (71,118) showed that methanol extracts of

P. pulmonariusandP. floridafruiting bodies decreased in-duced paw oedema in mice and ameliorated acute andchronic inflammation, respectively. Pleuran has also beenshown to possess anti-inflammatory activity by exertingantioxidant and immunomodulatory effects on rats withinduced colitis (119,120).

Hypersensitive immune responses, such as inflam-mation in delayed allergy, were suppressed by an etha-nol extract ofP. eryngii. It exhibited antiallergic activityafter oral or percutaneous administration to mice withoxazolone-induced type IV allergy (121).

Cardiovascular disease protection andantihyperglycaemic activities

Oyster mushrooms possess bioactive compounds withhypocholesterolaemic activities, such as polysaccharides,mevinolin and other statins (122). It has recently beenreported that P. citrinopileatus fruiting body extracts ex-

erted antihyperlipidaemic effects. Serum triglyceride andtotal cholesterol levels were lowered in hyperlipidaemicrats supplemented with the extracts, while high-densitylipoprotein levels were significantly increased (113). Si-milar effects were noted when powdered P. ostreatusfruiting bodies or a water-soluble polysaccharide ex-tracted from P. citrinopileatus fermentation broth werefed to hypercholesterolaemic or diabetic rats, respec-tively (123,124).

Pleurotusspecies also possess blood-pressure-lower-ing activity. Recently,P. cornucopiaehas exhibited antihy-pertensive activity; this might be due in part to D-man-nitol, which inhibits angiotensin I converting enzyme

(125).A methanol extract ofP. florida fruiting bodies sig-

nificantly inhibited platelet aggregation. The antiplatelet--aggregating activity, along with the anti-inflammatoryactivities discussed above, suggest its potential therape-


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utic use against vascular disorders, but the exact mecha-nism of these activities is unknown (118).

Antihyperglycaemic activity was demonstrated witha water-soluble polysaccharide fromP. citrinopileatusfer-mentation broth. The polysaccharide was effective inlowering blood glucose levels in diabetic rats (124).

Antimicrobial activityAntibacterial and antifungal activities have been ob-

served inPleurotusspp. extracts and isolated compounds,presumably produced as a defence mechanism againstother organisms. Table 1 (97,99,126129) summarizes re-cently reported antimicrobial activities ofPleurotus spp.

Antiviral activityMushrooms contain substances that exert direct or

indirect antiviral effects as a result of immunostimula-tory activity (130). Inhibitory activity against human im-munodeficiency virus (HIV)-1 reverse transcriptase hasrecently been demonstrated for P. sajor-caju and P. pul-monariushot water extracts (131). Anti-HIV activity wasalso demonstrated for a ubiquitin-like protein isolated fromP. ostreatus fruiting bodies (132). Moreover, Zhang et al.(133,134) demonstrated that, in contrast to water-insolu-bleb-glucans isolated fromP. tuber-regiumsclerotia, theircorresponding water-soluble sulphated derivatives exertantiviral activities against herpes simplex virus type 1

and type 2. The effect is presumably elicited by the bind-ing of sulphated b-glucans to viral particles, thus pre-venting them from infecting the host cells.

Concluding Remarks

Much work has been carried out on Pleurotus spp.mushroom cultivation, biomass production by means ofsolid- and liquid-state fermentation, and medicinal pro-perties. Studies on mushroom cultivation have been fo-cused on optimization of alternative substrates. It has beenshown that a wide variety of agricultural (by-)products,weeds and wastes can be successfully used to produce

food, feed, enzyme and medicinal compounds and todegrade and detoxify wastes. Due to an increasinglynegative human impact on the environment, these tech-niques, together with others, constitute a very importanttool for converting abundant quantities of waste materi-

als, which often cause environmental pollution, intofood and valuable compounds. These and many othermaterials have been successfully used for biomass pro-duction.

Solid- and liquid-state fermentations can be success-fully applied forPleurotusspp. cultivation. Each one hasits advantages and shortcomings and the decision as towhich method should be used for a specific applicationmust be carefully evaluated, as the outcomes are influ-enced by several factors. Solid-state fermentation is stillthe preferred method for waste utilization. Different ap-proaches can be used for enzyme production, as an ex-ample, the trickle film process for laccase production de-veloped by Hlker and Lenz (4). On the other hand,

submerged liquid fermentations can be more uniformand reproducible, which is interesting for obtaining pro-ducts with medicinal properties, although this potentialhas yet to be recognized by western biotechnology com-panies (5).Different substrates for solid- and liquid-statefermentations with Pleurotus spp. have been evaluatedand nutritional requirements and culturing parameterswere established. The authors point out that the correctselection of medium composition and environmental pa-rameters is crucial if optimal biomass, enzyme or metab-olite production is required.

Pleurotusspecies possess a number of beneficial me-dicinal properties, such as antitumour, immunomodula-tory, antigenotoxic, antioxidant, anti-inflammatory, anti-allergic, hypocholesterolaemic, antihypertensive, anti-hyperglycaemic, antimicrobial and antiviral activities.These activities have been reported for various extractsand isolated compounds, such as polysaccharides, poly-saccharide-protein complexes, proteoglycans, proteinsand DNA from oyster mushroom fermentation broth,mycelia or fruiting bodies. In particular, polysaccharidesappear to be potent antitumour and immunomodulatingsubstances, besides possessing other beneficial activities.However, the biochemical mechanisms of these thera-peutic activities still remain largely unknown.

Acknowledgements

We thank the Slovenian Research Agency (applica-tive research project L2-7598 and L4-6420) that enabledus to write this review article. We would also like tothank Prof. Dr. Roger Pain for taking the time to reviewour work.


Table 1. Reported antimicrobial activities ofPleurotus spp.

Species Effective against Reference

P. ostreatus

Crude extracts from fermentation broth Gram-positive, Gram-negative bacteria andAspergillus niger (126)

Hexane-dichloromethane extract containingp-anisaldehyde

Bacillus subtilis,Pseudomonas aeruginosa,Aspergillusnigerand Fusarium oxysporum

(127)

Various extracts; two main unidentifiedcompounds

Bacillusspp.,Escherichia coli,Vibrio choleraeandSalmonella typhi

(128)

P. eryngii Eryngin an antifungal peptide Fusarium oxysporumand Mycosphaerella arachidicola (129)

Eryngeolysin a haemolysin Bacillusspp. (99)

P. sajor-caju 12 kDa ribonuclease Fusarium oxysporum,Mycosphaerella arachidicola,

Pseudomonas aeruginosaand Staphylococcus aureus (97)

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