cultivation of lentinula edodes on synthetic logs

7/27/2019 Cultivation of Lentinula Edodes on Synthetic Logs

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Cultivation of Lentinula Edodes on Synthetic Logs

Alice W. Chen, Ph.D.Specialty Mushrooms

1730 Penfield Rd., #41Penfield, NY 14526

Voice: 716-381-0804

INTRODUCTIONLentinula edodes (Berkeley) Pegler = Lentinus edodes (Berkeley)Singer is best known by its Japanese name (shiitake), Chinesename (xiangu), and French name (lentin). This is, by far, the mostimportant specialty mushroom in North America (Chen, et al.,2000; Humble, 2001; Royse,1997; Stamets, 2000). In China,where it was originated, xiangumeans mushroom with greataroma (xian means aroma; gumeans mushroom). Two highlyprized forms of xiangu are dongu,the winter shiitake (dong meanswinter), and huagu, the flowershiitake (hua means flower). Bothforms with thick meaty mushroomcaps are produced at winter-liketemperature. Huagu, the mostsought-after shiitake and themost expensive on the globalmarket, is a form of dongu withflower-like cracking pattern on theupper surface of the mushroomcap (Wang et al., 2000).

L. edodes , a mushroom primarilyof temperate climate, is indige-nous in the far east. No wild shi-itake has been found in NorthAmerica or Europe until recently.Ammirati (1997) and Desjardin (1998) reported siting of wild shi-itake in the state of Washington and California. These mushroominhabitants in natural habitats in the United States of America arelikely wild transplants from domesticated cultivars, possibly run-away disposals.

Cultivation of shiitake on natural logs began in China almost athousand years ago. Wu, Sang Kwuang in Zhejian Province inChina was credited as the ingenious observer who figured outhow to enhance fruitings in shiitake that grew wild in nature (Milesand Chang, 1989). Scientific methodology evolved much later,when Dr. Shozaburo Minura in Japan developed the technique ofinoculating natural logs with pure shiitake mycelial culture in 1914(Mimura, 1904, 1915 in Stamets, 1993).

During the early twentieth century, my father, Prof. Chang-ChichHu, of Jing-Ling University (now Nanjing University) in Nanjing,China, was one of the pioneers trying to improve shiitake cultiva-tion practice in China, after he returned from advanced studies atTokyo University in Japan (Huang ed., 1987). Only about twodecades ago, in 1979, after a dozen years or so in research,China succeeded in large-scale shiitake synthetic-log cultivationon substrate blocks in bags, a much faster production comparedto cultivation on natural logs (Huang ed, 1987). Today, Chinaremains one of the largest producers, consumers and exportersof shiitake. In 2000, imports of shiitake to Japan rose 33% to 42,057 tons. These mushrooms had a value of $93.65 million.

Almost all of the shiitake imported to Japan came from China(The Mushroom Growers' Newsletter , June, 2001).

In the United States of America, shiitake cultivation began to takeoff between 1986-1996, following the lifting of a ban on importing

life cultures of L. edodes by USDA in 1972 (Royse, 1997). Now,fresh shiitake, cultivated by American growers is, no doubt, theleading specialty mushroom in supermarkets across the country.Dry shiitake has a long-standing history as a mushroom treasurein Oriental grocery stores, particularly in China towns and otherOriental communities.

Today, shiitake cultivation, using synthetic-log cultivation, is wide-ly practiced, not only in southeast Asia (China, Taiwan, Japan,Korea, Singapore, the Philippines, Sri Lanka and Thailand) butalso in North America (The United States and Canada), Europe(with France leading, Germany, the Netherlands, Spain, Italy,England, Switzerland, Belgium, Finland, Sweden), Australia andNew Zealand (Oei, 1996; Romanens, 2001). Shiitake cultivationis, indeed, a global industry.

For medicinal benefits, refer to Hobbs, 1995; mizuno, 1999;Stamets, 1999, 2000. This paper focuses on North American shi-itake synthetic-log cultivation, using Chinese methodology andother studies as references.

STRAIN SELECTIONShiitake strains vary widely, particularly in fruiting temperatureand mycelial maturation (early or late; shorter or longer produc-tion time). Substrate selectivity, growth rate (some fast strainsmay produce pre-mature fruiting), quality (shape, size, thickness,color, flavor and aroma, etc.), yield and ecological adaptability toextreme temperature (usually cold tolerance) are also strain-relat-ed.

Based primarily on the Chinese system, strains are classified into4 categories according to their fruiting temperatures:Faced with massive imports, the Japanese developed a numberof new shiitake strains with largeand thick basidiocarps(Watanabe, 2001). Performanceand stability of superior strains areboth important. Experiencedgrowers know the potential prob-lems of strain attenuation. Forexample repeated subculturesand prolonged storage of thestock culture may result in smallerfruiting bodies and lower yield(Huub Habets, ForestP r o d u c t s , S c h i m m e r t , t h eNetherlands).

SUBSTRATE SELECTIONAged broadleaf sawdust has beenused in China for shiitake cultiva-tion (Huang ed., 1989;1993).

Figure 1: Stock culture inmalt agar slant andcolonies with whitishmycelia (Dr. Noel Arnold)

Table 1Strain Classification by Fruiting Temperature

Low temperature 10 oCmid temperature 10-18 oChigh temperature 20 oC and abovewide-range temperature 5-35 oC

(e.g. strain, China-Stamets-2)

Figure 2: Liquid Spawn(Moore Mushroom FarmLabs, Glenmont, Ohio)


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Without aging, fresh sawdustcan be used for production ofshiitake only if it is from highquality tree species, such asthose graded 4, excellent byFAO (Oei 1996). Oak,chinkapin, hornbean, sweet-gum, poplar, alder, ironwood,beech, birch, willow are exam-ples of commonly used hard-woods in the U.S. Sawdust fromtree species of lower quality,however, has to be aged by fer-mentation (Oei 1996, Ting 1998,Wu et al. 1995). Growers typi-cally select the best and leastexpensive, locally available sub-strate materials. FermentedEucalyptus sawdust, for exam-ple, is used in Australia. Somegrowers prefer to use aged saw-dust regardless of tree species.

SUBSTRATEFORMULATIONBoth substrate nutrients andphysical textural property in aer-ation are important. Sawdustshould not be smaller than 0.85mm. (Royse, D. J. andSanchez-Vazquez, J. E. 2000).

For commonly used hardwood,sawdust-based for-mulations, see Table2. Many growers usea simple substratewith sawdust, branand 1% CaCO 3 (Oei,1996, p. 194). 1%sucrose is also fre-quently added. Inaddition to hard-wood, use of pine isa subject of greatinterest, since pine isa readily availableresource in somea r e a s .Supplemented pine-hardwood substrate(Table 2. Formula C)was used as partialsubstitute for basalingredient by theForestry ResearchInstitute of NewZealand for shiitakeproduction with satis-factory results. Agricultural wastes, such as cottonseed hulls,corn cobs, bagasse, and straw can also be used as alternativebasal ingredients. For additional formulae, see Hsu, 1997; Milesand Chang (1989) and Oei (1996, pp.198, 200).

SUBSTRATESTERILIZATIONSterilization depends on the nature of the bags (polypropylene or

polyethylene), bag size and,nature and amount of the sub-strate. For 2-3 kg. Sawdust-based substrate in polypropy-lene bags, sterilize in an auto-clave for 2 hr. at 121 oC. Adjustthis rule-of-thumb as necessary.

SPAWNINGIn general, through spawning(spawn thoroughly mixed withthe entire substrate) in largerbags is used in the U.S., whiletop or localized spawning(spawn is left on the substratesurface or the inoculation hole)in smaller bags is used in Chinaand Australia. Through spawn-ing gives a much faster growthrate. Heat-sealed larger bagswith microporous breathing fil-ters, partly filled with the sub-strates, allow mixing spawnwith the substrate by shakingmechanically or manually.Smaller bags with ring necksand plugs, however, when fullyloaded without leaving any airspace in bags do not lend them-selves to through spawning.

MANAGEMENTOF GROWTHPARAMETERSStamets (1993,2000) summarizedthe growth parame-ters for shiitake culti-vation presented inTable 3.

HOW SHIITAKEGROWProduction of shi-itake involves both avegetative phase ofmycelial growth andmaturation, and areproductive phaseof fruiting-body for-mation. It is impera-tive for growers tograsp the conceptand observe closelythis continuousprocess with intricatep h y s i o l o g i c a l

changes and morphogenesis, focusing on the transition from thevegetative phase to the reproductive phase.

SPAWN RUNThe spawn run phase is the period of mycelial growth and mat-uration. Fresh and vigorous spawn of appropriate age should beused for spawn run. This intricate vegetative phase includes 5stages. All shiitake strains show optimal mycelial growth at 25 oC.

Table 2

Sawdust-based Substrate Formulae for Shiitake

A. Wu (1993)sawdust 100 kgwheat or rice bran 23.25 kggypsum 2.5 kgcalcium superphosphate 0.5 kgsucrose 1-1.5 kgwater 10-140 kg

B. Stamets 1993, p.162,; 2000sawdust 100 lb (or 64 gal.)woodchips 50 lb (or 32 gal)rice or rye bran 40 lb (or 8 gal)gypsum (calcium sulfate) 5-7 lb (or 1 gal)water 60%

C. The Forestry Research institute of New Zealandpine 6 parts (monterey pine-Pinus radiata)Hardwood 3parts (beech or poplar)Grain 1 part (barley)

D. Straw-based substrate( Oei, 1996, p. 198)rice straw 50 kgwheat straw 20 kgsawdust 20 kgsucrose 1.3 kgCaCO3 1.5 kgcitric acid 0.2 kgCaSO4 0.5 kg

Table 3Growth Parameter Management

Stamets,(1993, 2000)

Spawn Run Induce Primordia FruitingTemperature 21-27 oC 10-16 oC* 16-21 oC** 16-18oC* 21-27 oC**

(70-80 oF) (50-60 oF) (60-70 oF) (50-70 oF) (60-80 oF)for all strains temperature fluctuation

Humidity 95-100%R.H. 95-100%R.H. 60-80%R.H.

Incubation ca. 1-2 mo. 5-7 days 5-8 days(Strain-dependent)

CO2 >10,000 ppm


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The duration of spawn run is usually 1-4 months, depending onstrains and methodology. No light is necessary during spawn run,however, some light in the day/night cycle towards the end of thespawn run is conducive to induction of primordia. Differentapproaches can be used, such as short exposure to light (e.g. 4hr/day-night cycle (Royse, 1997), or use a low level of light, 50-100 lux (Stamets, 2000), throughout spawn run. The dramaticchange from vegetative mycelial growth to produce macroscopicfruiting bodies in the reproductive phase requires enormousamount of energy reserves. A vigorous spawn run is of ultimateimportance. It should be noted that strains vary greatly in durationfor mycelial maturation. For one strain, 60 days is sufficient tomature, whereas this would be insufficient time for another strain(Miles and Chang, 1989).

Stage 1. Mycelial GrowthImmediately following inoculation, whitish shiitake mycelia beginto grow on the supplemented substrate, until colonization is com-pleted. This is an active assimilation phase with high fungal meta-bolic rate. Enzymes are activated to break down complex sub-strate components (e.g. cellulose, hemicellulose and lignin) intosimpler molecules which can be absorbed by mycelia as nutrientsfor growth and propagation.

In special cultivation practices, colonized mycelial blocks are sub- jected to higher temperature toward the end of spawn run: 1)exposing colonized blocks to 25-27 oC for a week before fruiting inJapan (Watanabe, 2001); 2) incubating colonized mycelial blocksat the upper limit, 27 oC-30 oC for a period of time in China (Milesand Chang, 1989). The rational for this methodology is based onthe conjuncture that higher temperatures that do not promote

mycelial growth may facilitate thedegradation of sawdust. It is notclear whether these conjectureshave been supported by inde-pendent studies. Some growersspray water on colonized mycelialblocks without bags to promotemycelial maturation and browning(Watanabe, 2001). Be awarethese effects may vary by strain.

It helps to keep in mind that somefast-growing strains may produceunexpected pre-mature fruitingbefore mycelial maturation (notdesirable). Care should be takento move the blocks as little aspossible during spawn run asmoving (physical shock) may trig-ger pre-mature fruiting in thesestrains. Fast-growing and shorterproduction time may not be thebest choice. The resulting mush-

rooms may not have the meaty texture desired in Asian markets,but could be acceptable in newer markets elsewhere. Usually,slow growth at winter-like temperature produces high qualitydong-gu, the winter shiitake or huagu, the flower shiitake, themost expensive and sought-after shiitake, formed at cold and drytemperature with diurnal fluctuations.

Stage 2. Mycelial Coat FormationTwo to four weeks after spawning, toward the later stages of thespawn run, a thick mycelial coat forms on the outer surface of thecolonized substrate block. Initially the coat is white in color. Athigh CO2, a very thick mycelial coat could be formed.

Stage 3. Bump Formation (blister stage, or popcorn stage)Clumps of mycelia appear as blister- or popcorn-like bumps ofvarious sizes on thesurface of themycelial coat in moststrains. This usuallybegins when colo-nization of whitemycelia covers theentire substrate inthe bag, or some-times earlier.Primordia are pro-duced at the tips ofsome of thesebumps. However,most bumps areaborted and neverdevelop into fruitingbodies. Time ofbump formationvaries with strains,substrate and tem-perature. Usuallybumps form 10 daysfaster at 25 oC than at15 oC (Miles andChang, 1989).Fluctuation of tem-perature and highCO 2 concentration encourage bump formation. Lower the CO 2 inthe bag, when bumps become too numerous by cutting slits onthe bag. In any case, some aeration should be provided whenbumps are formed.

Stages 4. and 5. Browning and bark-formationThere are two different approaches, browning outside of the bagvs browning inside of the bag. Some growers remove the entirebag when browning covers 1/3-1/2 of the mycelial coat in the bag(Oei, 1996). Royse (1997) recommends removing the bags

before pigmentation,thus allowing brown-ing to occur outside ofthe bag.

Timing of bag removalis crucial. Yield can beeffected if bagremoval is too early ortoo late. Maintain 60-70-% R.H. to avoidcontamination afterbag removal. Airenhances browning.Mycelia turn reddishbrown at the surfacein exposure to air andeventually forms adark brown protective,dry, and hardenedsurface which func-tions like a tree bark.The inner substratebecomes soft andmoist as a conse-quence of fungalmetabolism. The inner

Figure 3: A heat-sealedcultivation bag with a micro-filter breathing window(Unicorn Imp. & Mfg. Corp.,Commerce, Texas)

Figure 4: Cultivation bags with supple-mented substrate (top); colonized sub-strate blocks with whitish mycelia in cul-tivation bags (bottom).

Figure 5: Browning (pigment formation)initiation and completion in bags.


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moisture content can be as high as 80% (Oei,1996), ideal for fruiting formation.

Fruiting Induction(Oei 1996)Apply fruiting induction when spawn reachesphysiological maturity and after browning andbark formation. Water soaking is commonlyused for fruiting induction after browning andbark forming. Table 4 lists the factors that pro-mote fruiting.

Basidiocarp Formation primordia formation at the tip of the bump

(blister) formation of the young mushroom button

(dark brown) elongation of the stipe (stalk) as the button

increases in size expanding (opening) of the mushroom cap;

color becomes lighterFor a description of shiitake mushrooms, referto Huang ed., 1987).

Harvest, Post Harvest and Subsequent FlushesLower the humidity to 60% R. H. for 6-12 hr. before harvesting forbetter shelf life. Harvest when the edge of the mushroom cap isstill in-rolled, or when the mushroom cap is only partly extended(60-70%). This is the form desired by the Asian markets. Handpick the mushroom by holding the mushroom stalk and gentlytwisting it from the substrate block. Trim the end of the stalk afterharvest when necessary. Cut off residual stalk stubs from the sub-strate. Lower the humidity to 30-50% R.H. at 21 oC, for 7-10 daysof dormancy (Stamets, 1993), then soak the substrate block forup to 12 hours for the second flush, and up to 18 hours for thethird flush (Royse 1997). Adjust as necessary. Larger bags withmore substrate produce more flush-es.

In China and Japan, shiitake qualityis determined by shape (roundedwith downward in-rolled edge beforethe cap is fully extended and centralstalk), texture (thick and tight con-text, the meaty part), size, color, fla-vor (enhanced by cooking especiallythe fresh shiitake) and aroma(enhanced by drying). Freshnessand freedom from pests and impurities are also critical factors forhigh quality mushrooms.

Shiitake Cultivation in the U.S.In contrast to growers in southeast Asia, heat-sealed larger bagswith microfilter breathing windows are used in general by growersin North America. These bags, eachfilled with 2-3 kg, or more (5.5 kg) sub-strate, produce more flushes of mush-rooms in shorter production time byusing through spawning. This is lesslabor intensive, less time-consuming,and results in less contamination.

Mushrooms are produced underindoor controlled growth parameters.Mechanization is used by moresophisticated growers. There is a ten-dency for North American growers to

use faster growing strains, especially the newgrowers, to gain confidence. Shiitake mush-rooms produced by growers here are soldfresh. The markets of specialty gourmet mush-rooms in North America are fairly new with, per-haps, some less sophisticated customers.

U.S. NATIONAL ORGANIC STAN-DARDS (www.ams.usda.gov/nop; The Mushroom Growers' Newsletter , Feb. 1998;personal communication, 2001)In recent years, pollutants such as heavy met-als and others, have been found in mushroomproducts. In rare cases, higher concentration ofheavy metal was detected in fruiting bodiesthan in the substrate and casing soil used (Li etal., 1998). The potential capability for fungi toabsorb and accumulate heavy metals inmycelia and fruiting bodies has been demon-strated (Huang et al., 1998). To cultivate mush-rooms organically is an effective way to safe-guard the quality of mushroom produce andproducts. Organic farming was a $60 billion

market in the United States in 1999. It is advisable to adopt theprocess of growing mushrooms organically regardless of the spe-cific methodology a grower chooses to use.

NOP (the National Organic Program) under the AgriculturalMarketing Service of the USDA published a proposed rule for anational organic program (7 CFR 205) for public comment. OnMarch 7, 2000, they released a new proposal for uniform and con-sistent national standards for organic food. Certification isrequired for farmers and wild-crop harvesters who wish to use theorganic label on their products. A specific rule for organic stan-dards for mushrooms will be announced in the near future. Ingeneral, for crops and wild harvests to meet the national organic

standards, the standards listed inTable 6 apply.

Production of Huagu, theFlower ShiitakeIntroductionHuagu, the flower shiitake, occursspontaneously in nature whenmushroom spores are depositedunder cold and dry winter months.Huagu is not a genetically inherenttrait. On the contrary, huagu, the

shiitake mushroom with a unique morphological flower-like crack-ing pattern on the cap, is produced through manipulation ofgrowth parameters. Success in cultivation of huagu can bringgrowers considerable extra income in China for domestic and for-eign consumption. Model huagu production systems can befound in Gutien County, Fujian Province and Quingyuan County,

Zhejiang Province, in China. As a dryproduce, huagu varies in quality.White huagu with deep and widecracking grain and thick context(mushroom meaty part) tops thegrading scale, while dark tea-coloredwith less pronounced cracking grainare inferior.

The principle of huaguformationGrowth parameters are manipulatedduring the formation of shiitake basid-

Table 6National Organic Standards for

Crops and Wild Harvests

1. No pesticide. 3% tolerance of pesticide residue.2. Growing field pesticide free for three years.3. No hormones.4. No sewage sludge fertilizer (reprocessed wastes).5. No ionizing irradiation.6. No genetic engineering.

Table 4

Factors in Primordia

Initiation

(Oei, 1996; Watanabe, 2001)

water soaking(Royse 1997: 2-4 hr. at 12oC;Stamets 1993: 24-48 hr.)

temperature fluctuation high humidity

removal of CO2, or increase ofoxygen supply

physical shocks (agitation, dis-turbance), stab (with a long metalneedle) and inject (with water),turn the blocks up-side-down(half-way during spawn run:Watanabe, 2001), beating (e.g.natural logs), electric stimulation

Table 5Features of American

Synthetic-log Cultivation Use heat-sealed larger bags with microfilter windows Use larger amount of supplemented substrates Through spawning Mechanization Growing indoors under controlled parameters Growing mushrooms organically


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and considerable differential growth rate between the surface andthe inner context. In time, the rapid growth of the inner contextruptures the mushroom surface, producing a flower-like crackingpattern on the cap surface, thus the name, huagu.

Selection of strains for huagu production

Use low temperature, high quality and easily adaptable strains forhuagu production. Strains towards the lower temperature marginin mid-temperature range can also be used. Examples of desir-able huagu strains are: L-241-1, Jean-Yin #1, Yee-You #5, 7402,N-06. Strain characteristics should be thoroughly studied beforecultivation. For fruiting outdoors, time of spawning should be

coordinated with the maturation characteristics in order to benefitfrom the winter stimulation. For example, strain 7402, N-06, latematuring strains, should be inoculated early during March andApril, while 9018, Le 204, early to mid-maturing strains, should beinoculated in May-June in Bi-Yang, China (Yu 1998). Adjust foryour local weather.

Forcing of HuaguInitiate huagu forcing when the mushroom buttons reach 2-3 cm(diameter). If huagu forcing is applied too early when the buttonsare smaller than 1.5 cm (diameter), the fragile young buttons maydie of drought or freezing. If the technique is applied too late,when the mushroom already reach 3.5 cm (diameter) or larger,the mushrooms do not respond readily.

iocarps (fruiting bodies), under winter or winter-like conditions.When the young mushroom buttons (not primordia) reach 2-3 cmin diameter, dry air and cold temperature force the pilial (cap) sur-face into dormancy. Under such adverse environment with dras-tic diurnal fluctuation of temperature and humidity, a protective drysurface is formed on the young mushroom cap. However, theinner context continues to grow at a slow pace with water avail-

able solelyfrom thesubstrate.W h e nfavorableg r o w t hconditionsreturn, thes u r f a c egrows at ar e t a r d e drate, whilethe innerc o n t e x tdeve lopsat a nor-mal pace.U n d e rthese con-d i t i o n s ,s h i i t a k emushroomb u t t o n sgrow withthe alter-nation ofdormancya n dg r o w t h ,

Figure 6: Young shiitake mushrooms with darkcaps formed after soaking in water (Dr. NoelArnold)

Figure 7: Growth of young shiitake mushrooms: the color of thecaps become lighter and the size larger as the stipes elongate.

Figure 8: Shiitake cultivation in the U.S.

Figure 9: Shiitake cultivation in the U.S.


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Huagu Forcing Technology(Ting, 1994, Xu, 1998, Huang, 2000)

There are three stages of huagu formation (Yu, 1998):1.Pre-conditioning by low temperature. Subject shiitake buttons

of the proper developmental stage (from primordia to 2cm indiameter) to:Temperature 8-12 oCHumidity 85-90% R. H. (remains high)

2.Forcing Huagu as follows when buttons reach 2-2.5 cm(diameter).

Temperature 15 oC +- 1 oC (8-18 oC)Humidity 50-67% R.H.apply misting of water when


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The Mushroom Growers Newsletter, June, 2001

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Royse, D. 1997. Cultivation of shiitake on natural and syntheticlogs. University Park, Penn State , PA: College of AgriculturalSciences, Cooperative Extension, 10pp.

Royse, D. J. and Sanchez-vazquez, J. E. 2000. influence of woodchip particle size used in substrate on biological efficiency andpost-soak log weights of shiitake. Science and cultivation ofedible fungi. Van Griensven (ed), pp. 367-373, Balkema,Rotterdam, the Netherlands.

Stamets, P. 1993, 2000 (new 3rd edition). Growing gourmet andmedicinal mushrooms. Berkeley, CA: Ten Speed Press,pp.259-276.

Stamets, P. 1999. Mycomedicinals: an informational booklet onmedicinal mushrooms. Olympia, WA:Mycomedia.

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2001 by the Author and The Mushroom Growers Newsletter - Reprinted from the August 2001 issue.

The Mushroom Growers NewsletterP.O. Box 5065

Klamath Falls, OR 97601 USAOn the world-wide-web at: www.mushroomcompany.com

Figure 12: Huagu (flower shiitake) cultivation in China.

cultivation of lentinula edodes on synthetic logs

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