Appl Microbiol Biotechnol (2010) 85:1321–1337 DOI 10.



Cultivation of Pleurotus ostreatus and other edible mushrooms
Carmen Sánchez

Received: 1 September 2009 / Revised: 1 November 2009 / Accepted: 1 November 2009 / Published online: 3 December 2009 # Springer-Verlag 2009

Abstract Pleurotus ostreatus is the second most cultivated edible mushroom worldwide after Agaricus bisporus. It has economic and ecological values and medicinal properties. Mushroom culture has moved toward diversification with the production of other mushrooms. Edible mushrooms are able to colonize and degrade a large variety of lignocellulosic substrates and other wastes which are produced primarily through the activities of the agricultural, forest, and food-processing industries. Particularly, P. ostreatus requires a shorter growth time in comparison to other edible mushrooms. The substrate used for their cultivation does not require sterilization, only pasteurization, which is less expensive. Growing oyster mushrooms convert a high percentage of the substrate to fruiting bodies, increasing profitability. P. ostreatus demands few environmental controls, and their fruiting bodies are not often attacked by diseases and pests, and they can be cultivated in a simple and cheap way. All this makes P. ostreatus cultivation an excellent alternative for production of mushrooms when compared to other mushrooms. Keywords Pleurotus ostreatus . Mushroom cultivation . Edible mushrooms

Introduction Cultivation of edible mushrooms is a biotechnological process for lignocellulosic organic waste recycling. It might
C. Sánchez (*) Laboratory of Biotechnology, Research Centre for Biological Sciences, Universidad Autónoma de Tlaxcala, Apartado postal 129, Tlaxcala, Tlax CP 90000, Mexico e-mail:

be the only current process that combines the production of protein-rich food with the reduction of environmental pollution (Beetz and Kustudia 2004). The production of mushrooms is regarded as the second most important commercial microbial technology next to yeast (Pathak et al. 2009). Mushrooms have been eaten and appreciated for their flavor, economical and ecological values, and medicinal properties for many years. In general, mushrooms contain 90% water and 10% dry matter (Morais et al. 2000; Sánchez 2004). They have chemical composition which are attractive from the nutritional point of view (Gbolagade et al. 2006; Dundar et al. 2008; Table 1). Their nutritional value can be compared to those of eggs, milk, and meat (Oei 2003). Mushrooms also contain vitamins and an abundance of essential amino acids (Sánchez 2004). The total energetic value of mushroom caps is between 250 and 350 cal/kg of fresh mushrooms (Oliver and Delmas 1987; Laborde 1995). Some mushroom can be cultivated easily and have significant worldwide markets. Over 200 species have been collected from the wild and used for various traditional medical purposes, mainly in the Far East (Sánchez 2004). Roughly 300 mushrooms species are edible, but only 30 have been domesticated and ten grown commercially (Barny 2009). The principal cultivated mushroom worldwide is Agaricus bisporus followed by Pleurotus sp. (Rühl et al. 2008), Lentinula edodes, and other mushrooms that have already an important place in the market. Mushrooms have the ability to degrade several lignocellulosic substrates (Fig. 1; Sánchez 2009) and can be produced on natural materials from agriculture, woodland, animal husbandry, and manufacturing industries (Table 2). As it is shown in Fig. 1, laccases or ligninolytic peroxidases (LiP and MnP) produced by white-rot fungi oxidize the lignin polymer, thereby generating aromatic radicals (a). These evolve in different non-enzymatic reactions, including

The first stage involves obtaining pure mycelium of the specific mushroom strain.45 2. from a piece of the specific mushroom (Fig.03 15. mushroom production generates an enormous amount of used “spent” substrate which might also be a source of environmental contamination.42 0.25 6.35 10. and some of them have already been established (Sánchez 2004).65 5.44 0.85 2.90 5.55 3.75 5..52 0.70 8.80 58.70 4.10 0. 382 in USA.90 – 19.80 0.97 0. Quinones from g and/or k contribute to oxygen activation in redox cycling reactions involving quinone reductases.35 2.40 12.10 13.09 0. Lipids 5.90 0.65 11.45 0.40 4.40 18.20 7. Then lignin degradation proceeds by oxidative attack of the enzymes described above.25 1. 3b).20 9.80 0.20 5. 80 in Canada.55 0.30 12.411 in China. WBWDI 2007. To obtain inoculum.29 0.07 0. However. The latter is a very strong oxidizer that can initiate the attack on lignin (p) in the initial stages of wood decay.10 1.50 14.60 4.05 5.75 9.70 1.08 0.75 48.30 0.85 – Mushroom cultivation Mushroom culture involves several different operations.50 0. Phenoxy radicals can also be subjected to Cα-Cβ breakdown (k).20 2.65 6.30 27.05 7.32 0.95 7.50 4. 2).50 26.192 Appl Microbiol Biotechnol (2010) 85:1321–1337 Table 1 Nutrimental value of several edible mushrooms (mg/100 g dry matter) Protein 8.12 0.15 2.20 0.20 10. each of which must be carefully performed.021 0.50 7.11 0. 138 in Spain.30 5.10 9. 139 in France.05 0.80 44.85 0.70 32.90 15.14 Sugars 5. USDA 2005. wheat.90 4. The mycelium can be obtained from spores (Fig.15 28.95 1.30 1.75 6.85 8. or millet (Fig.15 14. rye.80 13. 88 in Italy.10 0.75 14.90 3.06 1.80 1.65 2.g.15 14.45 3.84 0.82 3. and 74 in the UK (FAOSTAT 2005.50 0. when the small size of pores in the still-intact cell wall prevents the penetration of ligninolytic enzymes.29 0.1322 0. aromatic ring cleavage (c). or from several germplasm providers such as American Type Culture Collection and National Center for Agricultural Utilization Research (Fig.75 5.70 2.30 15.00 3.85 2.91 0. and demethoxylation (e).05 16.10 0.40 10. g) are the substrate for H2O2 generation by aryl-alcohol oxidase in cyclic redox reactions involving also aryl-alcohol dehydrogenases.80 24.50 0. Zn Cu Fe Mn P Na K Mg Ca Ash Fiber 3.67 0. the mycelium is developed on cereal grain.50 0. incubation. 3a). The phenolic compounds formed can be again reoxidized by laccases or peroxidases (j).60 51.08 0. 245 in Netherlands.40 1.65 7.05 4.60 9. and peroxidases (l.25 0.72 0.60 12.50 20.70 0.72 0.51 14. Several uses for spent mushrooms substrate (SMS) are being evaluated. CαCβ breakdown (d).30 16.02 0. 2008 Edible mushroom .0 24. Substrate preparation. 77 in Ireland.55 2.35 8. Fig.00 0.95 18.0 8.70 17. either by superoxide cation radical or directly by the semiquinone radicals.12 0.15 0. inoculation. and its reoxidation with concomitant reduction of H2O2 to hydroxyl free radical (OH.55 10.85 0. The aromatic aldehydes releases from Cα-Cβ breakdown of lignin or synthesized de novo by fungi (f.13 0.05 0. This results in reduction of the ferric iron present in wood (n). Poppe (2000) reported that there are about 200 kinds of waste in which edible mushrooms can be produced.90 3.32 0.35 16. 3d).15 Auricularia polytricha Lentinus subnudus Lycoperdon pusilum Lycoperdon giganteum Pleurotus tuber-regium Pleurotus florida Psathyrella atroumbonata Schizophyllum commune Termitomyces microcarpus Termitomyces globalus Tricholoma lobayensis Volvariella esculenta Pleurotus sajor-caju Gbolagade 2006.75 1. which is usually called the “spawn” (Fig.20 10.57 3.45 0.50 5.10 17.80 1.50 6.55 10.10 24.07 0.163 0.75 1.45 5.95 6.35 15.90 9.20 31.07 0.97 1.07 0.50 8.) (o).00 1.77 1. 135 in Poland. and production conditions depend on the mushroom species to be cultivated (Fig. laccases. yielding ρ-quinones.0 – 4. Phenoxy radicals from C4-ether breakdown (b) can repolymerize on the lignin polymer (h) if they are not first reduced by oxidases to phenolic compounds (i).80 10. m). 3c).32 0. 3).50 0. 3e. Chang and Miles 1989). Dundar et al.75 43.90 0.46 37.40 22. e.65 8.30 1. Chang and Hayes 1978. The production of mushrooms in 2005 was (in metric tons) 1.80 9.20 4.40 11.30 0.16 0.50 1.75 C-4-ether breakdown (b).45 4.

2005. 3g). 2008). “hiratake”. the fungal Pleurotus genus has been intensely studied and cultivated in many different parts of the world. or “houbitake” (Mizuno and Zhuang 1995. Elhami and Ansari 2008) and on grain mixed with grain straw (Muthukrishnan et al. Nwanze et al. which must be prepared under sterile conditions to diminish contamination of the substrate (Fig. It is produced on a variety of lignocellulosic substrates (Table 2). Sainos et al. Pathmashini et al. the spawn for cultivation of P. Several studies have been done to improve the quality and develop new techniques for its production. ostreatus has been prepared in different ways: on grain. . ostreatus is also known as “oyster mushroom”. 2000. Bononi et al. Rühl et al. 2006. For example. 2008). 1995. The success of mushroom production depends in great part on the quality of the “spawn”. and paddy (Beetz and Kustudia 2004. such as wheat. sorghum. 3f). 1 Lignin biodegradation process by white rot fungi HC HOOH2 LIGNIN 1323 HOCH2 OCH3 HC HC O HC LIGNIN HCOH H2COH HOCH2 O OH* CH C=O HC HCOH O OCH3 h H2OC OCH3 OH o O LIGNIN H2COH HC LIGNIN j LACCASE HC H 2O 2 Fe 3+ PEROXIDASE a OCH3 OH n g AAO H2COH MeOH HC=O d e i LIGNIN AAO? HC H2COH CH2OH HCOH HC HC b OCH3 LIGNIN +* R O c OCH3 OCH O-R 3 O-R AAD H2COH OCH3 LIGNIN f -* HC LACCASE PEROXIDASE O* O2 m HCOH LACCASE OH PEROXIDASE ll g C ll O O OCH3 OH O ll OCH3 OCH 3 k OH QR l Source: Martínez et al. Cultivation of Pleurotus ostreatus P. For many reasons. 2005 The purpose of the mycelium-coated grain is to rapidly colonize the specific bulk growing substrate (Fig. “shimeji”.Appl Microbiol Biotechnol (2010) 85:1321–1337 Fig.

Cayetano-Catarino and Bernabé-González 2008. 2003. sugar cane. table. 2001. king oyster mushroom Beetz and Kustudia 2004. 2008. Lion's head or Pom pom Auricularia auricula Jelly ear Grifola frondosa Maitake. coffee pulp. sorghum. florida) Oyster. wood Wastes of rice. sawdust-rice bran Beetz and Kustudia 2004 Sawdust. Swiss brown. oil-palm fiber. leaves.1324 Table 2 Ligninocellulosic residues used for different mushrooms cultivation Japonese name Himematsutake or Agarikusutakea (name for Agaricus blazei) Substrate Reference Beetz and Kustudia 2004. peanut and coconut shells. Black oak Shiitake Rani et al. T. 2009 . Abrar et al. eous. Black truffle black perigord. horse manure (fresh or composted) Wheat straw and waste tea Leaves Coffee pulp. textile industry. sugar cane bagasse. corncobs. connotus. Kapoor et al. bean. (S. paper and paddy Sorghum stalk. cotton seed hulls. leaves. cassava peels. Montoya and Varón 2008 Sawdust Logs sawdust-rice bran Logs Beetz and Kustudia 2004 Beetz and Kustudia 2004 Beetz and Kustudia 2004 Tuber sp. wheat. sawdust. portobello. Shah et al. Fan et al. sajor-caju. sorghum. distillers grain waste Beetz and Kustudia 2004 Sawdust. Daba et al. sheep's head. peanut meal. magnatum Pico ex Vitt. corn. cotton. grass. (L. bracts of pineapple. cotton. water hyacinth. water lily. palm fibers. corn stalks. 2009 Lentinula sp. Shiitake mushroom black forest. Italian brown and Italian roman brown mushroom. Obodai et al. hen of the woods. crushed bagasse and molasses from sugar industry. 2008. wheat Banana leaves. wheat. Winter. sawdust.) piedmontese White truffle Stropharia sp. banana. cloud mushroom. clover. sawdust. P. P. and sawdust Wastes from rice. rice husk. Belewu and Belewu 2005. Beetz and Kustudia 2004. 2004. Champignon Hiratake Pleurotus sp (P. P. 2009. Rani et al. and rice 2007. edodes) Japanese forest mushroom. 2007 Scientific name English name Agaricus bisporus Button. Hideno et al. coffee pulp. Toker et al. white.. potting soil Carluccio 2003 Beetz and Kustudia 2004. velvet stem. wheat bran. Onuoha et sawdust Beetz and Kustudia 2004 Yamabushitake Kikurage Mai Take. pulmonarius. snow puff Pholiota nameko Fat pholiota Tremella White jelly Enokitake Numerisugtake Shirokikurage a MAYA ethnobotanicals (2009) Appl Microbiol Biotechnol (2010) 85:1321–1337 b http://zipcodezoo.. corncobs. water hyacinth. cotton seed meal. and water lily Mungbean husks. melanosporum Vitt. cocoyam peelings and oil-palm pericarp. rugosoannulata Farlow f. Sivrikaya and Peker 1999. P. spent sawdust matrix. ostreatus. eryngii. 2009 Volvariella volvacea Straw mushroom or paddy straw mushroom Fukurotake Se you fuyu shuororo (name for violet truffle)b Saketsubatake. Philippoussis et al. Jamaica. Maitake Sawdust. banana. cotton from textile industry. Ukoima et al. crimini. rams head. dancing mushroom Flammulina sp. wheat straw. L. cotton seed hulls bran. cocoa. 2008. 2009 Wastes from wheat. kisaketsubatake Rice straw. rugosoannulata Roundheads Farlow. P. T. rice bran and soybean mea Wastes from paddy. lutea hongo) Hericium sp. Beetz and Kustudia 2004. S. wheat straw. gremini. corncobs. golden. barley. 2006.

Pasteurization. 2 Top mushrooms cultivation countries Substrate preparation The substrate is milled to a length of about 2 to 6 cm. and temperature of spawn run.Appl Microbiol Biotechnol (2010) 85:1321–1337 1325 production cycle. First. water is added to the desired level. usually 7–8 days after setting up the stacks of 1 t of horse manure. Fig. however.0 m in width and height (wind-rows). Nogueira de Andrade et al. At time of spawning. more inoculum points. 2008. would provide faster substrate colonization and. In other substrates. Increasing the amount of spawn used (up to 5% of the wet weight of the substrate) has resulted in increased yields (Royse 2002). the substrate is cooled and spawned with the desired strain. Inoculation After completion of pasteurization (60 °C for 1 to 2 h). etc. Filling plastic bags with substrate The pasteurized substrate is spawned and filled (from 25 to 30 lbs) into clear or black perforated polyethylene bags. The substrate for the cultivation of this mushroom is horse manure compost. a yield of just over 1 t of compost will have been obtained. thus. different lignocellulosic substrates. Finally. Saavedra et al. 2006. Royse and Zaki 1991). Kothe 2001. 2004.25% to 5% may result in increases of nearly 50%. The stacks are turned twice a week in order to obtain uniform degradation of the mixture. When carried out in the open air. amount of supplement used. available from increased spawn levels. Increasing spawn rates from 1. the increased level of nutrient available in higher levels of spawn used would provide more energy for mycelial growth and development. Harvest The mushrooms are harvested from the substrate approximately 3 to 4 weeks after spawning depending on strain. Alternatively. and the conditions that favor fruiting. are important for the cultivation productivity of each particular mushroom (Mandeel et al. phase I of composting is done in long narrow stacks between 1. some broiler chicken manure. 2005. moisture and other conditions for mycelium growth. a synthetic compost can be made which is based on wheat straw and broiler chicken manure. Studies on the mushroom cultivation on the use of different strains. moisture. Incubation The bags are incubated for 12 to 14 days at 25 °C and then transferred to the production room (Royse 2007). 2009). and water to which gypsum is added for structural stability and for stabilizing pH. Several studies on the development of this mushroom have improved its cultivation (Stoop and Mooibroek 1999. Kirbag and Akyuz 2008. 2008. 3h). One of the most common substrates used for modern mushrooms is a mixture. is carried out by filling the ingredients into revolving mixers. Second. more rapid completion of the Cultivation of A. used on some commercial mushroom farms. Onuoha et al. The bags are maintained under optimal temperature. physicochemical conditions. and live steam is injected into the mixer while it is in operation (Royse 2007). in the past. the chopping is not required. Yield increases may be due to several factors. This mixture of cottonseed hulls and wheat straw has a higher water holding capacity than cottonseed hulls used alone. to optimize the amount of spawn used to inoculate their substrate. At the end of the process. may cause overheating of the substrate if growers are not able to anticipate and control air temperatures to maintain a steady substrate temperature. Bechara et al. Spawning and spawn rate Growers have sought. 3i). and its substrate is the most complex culture medium used for edible mushroom production (Fig. This fresh . different types of spawn. a delayed release supplement (rates of 3% to 10% of dry substrate weight) may be added to increase yield and size of the mushroom (Royse et al. Kapoor et al. Use of supplements. is also known as “himematsutake”. 2009).5 and 2. bisporus Agaricus sp. 1991. which consists of a mixture of horse manure. It is the principal cultivated mushroom worldwide. Mushroom production The mushroom begins to form around the edges of bag perforations. Mushroom shelves and suspended systems are the main systems used for Pleurotus cultivation (Fig. a more rapid spawn run would reduce the time non-colonized substrate is exposed to competitors such as weed molds and bacteria (Table 3).

Phase II consists of a pasteurization process followed by further high-temperature fermentation. and this is maintained for at least five more hours. is subjected to phase II treatment before actual cultivation can start. Compost temperature then increases to slightly higher values. the compost is kept . This is done by raising the air temperature to 56 °C and keeping it there for approximately 5 to 6 h.1326 Fig. which still smells strongly of ammonia. Thereafter. 3 Cultivation and harvesting of mushroom Appl Microbiol Biotechnol (2010) 85:1321–1337 Source: Modified from Stamets (1995) compost (phase I).

mold. and Check hygiene. Check ventilation Improve hygiene in the incubation house Mix well the substrate Mite contamination Mycelium grows but fails to produce mushrooms Substrate formula is not suitable Mites. carbon dioxide. bacteria and insects Mushrooms form. Stop using the area to cut the life cycle of all contaminants for a period of at least 1 to 2 weeks. Check temperature and control surroundings to maintain of 25 to 35°C Not more than 5% carbon dioxide. not enough ventilation to decrease accumulated temperature Too high carbon dioxide Hygiene of the incubation house Mycelium develops in patches. insects. decrease moisture Use activated charcoal water filters to eliminate chemical contaminants or any other ways of simple or appropriate technology Check substrate. For serious contamination cases. tools. make sure to clean every thing (person. humidity. Substrate is not evenly prepared. inoculation. Note: If the substrate is too moist. spots. In Check the process causing contamination. other fungi contamination Inoculate in hygiene conditions. but abort or delay mushrooming Inhibited by environmental toxins Bad strain or Shawn Primodia and growth condition of fruiting body are not good enough There is contamination such as mold. clean and with no air movement Spread the substrate bag and make more air ventilation in the incubation area. light. Use black-light with water or sticky-trap to decrease insects Adjust the formula. temperature. equipment. etc. and ventilation. bad smell. package again. make sure all raw materials are good and fresh Note: It is necessary to pasteurize immediately after bagging otherwise fermentation gas will slow down the rate of growth of mycelium or stop mycelium growth Check method of pasteurization. and mites Consult parameter of growth. area. other processes and mushroom house management for hygiene Remove source of toxins Acquire new strains Check temperature and humidity. Alter moisture. Continue the normal process Note: Keep hygiene management.Table 3 Troubleshooting in the mushrooms cultivation Cause Improper initiation strategy Chlorinated or contaminated water Bad substrate Solutions Problem Mycelium fails to form Appl Microbiol Biotechnol (2010) 85:1321–1337 Bad pasteurization Poor spread of mycelium. Separate contaminated bags as soon as possible. Spread the substrate and remix the substrate. etc Check pasteurization process. adjust environment of light. worms. Check that the cotton plug is tightly closed Substrate in the bag is too hot when inoculation Bad strain or spawn Spawn contaminated Forgot to inoculate the bag Good pasteurization but must decrease the temperature in the pasteurization chamber Pasteurization was too quick and/or the chamber door was opened too quickly Inoculation process Too high density in the incubation area. and surroundings during every step. temperature. sawdust. virus. Release all air and make sure there is continuous steam before starting pasteurization for a period of 3 h Make sure that the substrate bag is not too hot before inoculation Obtain younger strain of known vitality and history Pasteurize and inoculate again with good spawn Make sure to inoculate Slowly decrease the temperature in the chamber. bacteria. Follow process Immediately separate contaminated bags and pasteurize again. Remix substrate separately. check pH. Remake substrate bags and pasteurize for a longer time. Do not open the cover of the chamber too quickly. additives. and some parts have more nutrients than others Bacteria. spray area with chemicals. Open or close doors and window to adjust accordingly 1327 .

air. Reduce humidity to the prescribed levels. For severe cases. pierce bags and drain water Acquire better strain Never use chemicals during the fruiting stage Reformulate Check hygiene. Mist the bags and the surface of mushrooms. humidity. Spread lime on shelves. caps underdeveloped Massive numbers of mushrooms form. Surface water must evaporate from mushrooms several times per day. if there is water in bags. Pseudomonas fluorescens) because of bacteria during flush on Oyster mushroom more severe cases. Inadequate substrate nutrition fail to produce subsequent flushes Competitors Poor growing house management Bad strain Mushrooms small sized Too many mushrooms coming out at the same time Lack of nutrients in substrate Change of weather Spawn unhealthy Pests and insects Natural occurrence. and ventilation Improve management Acquire new strain Reduce the size of opening(s) Review quality of substrate Beware of wide range changes in temperature Check origin of spawn Place lemongrass plants around mushroom house. Give enough time for water to evaporate from mushroom surfaces before further watering. humid climate Mushroom waste lying around mushroom house Ants Mushrooms are light in weight Shortage of water Mushroom quickly spoil Mushrooms too mature when harvested Mushrooms too warm before packaging Mushrooms too wet when harvested Mushrooms stored beyond shelf life Rotting spot on the mushroom fruiting body Bacteria (Pseudomonas tolaasii.1328 Table 3 (continued) Cause mites Solutions Problem Mushrooms form. decay and die Chemical contamination from solvents. adjust light.5 l of water. use half a teaspoon of sulfur in 3. and ground in the mushroom house. few develop Mushrooms are deformed. Remove contaminated bags from mushroom house and recycle Remove toxins Acquire a new strain or find a new supplier Increase or adjust light to correct wavelength Increase air exchange. on poles. chlorine. Clean (and maintain clean) the mushroom house properly Try to use the waste as fertilizer or recycle Mix detergent with water and place on their paths. open doors or windows and close at correct time Shorten the period for the formation of primodia Increase air ventilation and open more windows or doors to receive more light Reformulate or check raw materials Use the high rate spawn or adjust good conditions for rate of growth Obtain better strain Adjust mushroom house to favor mushrooms and not germs and competitors Clean the surface of substrate Increase air circulation. temperature. etc Bad strain Inadequate light Excessive carbon dioxide Too long time incubation Lack of oxygen. Check watering. too high humidity Bad strain Use of chemicals during this period Mushrooms produced only in the first flush. Do not put on mushroom Check humidity of mushroom Harvest when younger Chill mushrooms before placing in marketing containers Reduce humidity several hours before harvesting Sell mushrooms faster Control humidity in the mushroom house and maintain 80–85%. gas. but stems are long. use 113 g chlorine mixed in 45 l of water or 4 oz of chlorine per gallon of water Appl Microbiol Biotechnol (2010) 85:1321–1337 Source: FAO 2001 . inadequate light Inadequate substrate nutrition or low quality Low rate mycelium growth Poor strain Disturbed by germs or competing microorganisms Dirty surface of substrate bags Not enough air ventilation.

the entire cluster of fruiting bodies is removed from the bottles. and length of incubation are important parameters for shiitake production on synthetic substrates in bag-log cultivation (Zadrazil 1993. but research has shown that sawdust from conifers such Pinus spp. frondosa is usually on a substrate contained in polypropylene bottles or bags. substrate composition. best quality. the substrate is inoculated. Chang and Miles 1989. 3j). two holes usually are cut in the bags exposing the developing primordia that tend to develop around the outside perimeter of the substrate surface. To further improve quality during fruiting. temperatures are lowered to 3 °C to 8 °C until harvest. When that process has been completed. After primordia formation. velutipes in Japan is based on a substrate contained in polypropylene bottles. and the colonized substrate is subjected to environmental conditions known to stimulate fruiting. This collar serves to hold the mushrooms in place so that they are long and straight. It has also been produced on a substrate consisting of oak sawdust. Composted substrate is prepared by mixing and watering of ingredients. velutipes except that a higher moisture content of the substrate is desirable. and the process is repeated (Royse 2007). wheat bran. Substrates (primarily sawdust and rice bran) are mechanically mixed and filled into heat-resistant bottles. It is usually produced in polypropylene bottles contained in plastic trays. Cultivation of Pholiota nameko Preparation of the medium for P. A substrate of broad-leaf tree sawdust is preferred. and incubated for 25 days at 20 °C. Cultivation of Flammulina velutipes Japan is the main producer of F. Kalberer 1995. mechanically inoculated. For production in bags. The top of the bag is then folded over. and rye. The strain. Cultivation of L. a plastic collar is placed around the neck and secured with a Velcro® strip. bottle lids are removed. and the culture of other mushrooms has been reported. The mushrooms are packaged by placing an entire cluster (or multiple clusters) into each over-wrapped package. exposing only the developing primordia to the fruiting environment (Royse 2007).Appl Microbiol Biotechnol (2010) 85:1321–1337 1329 at a temperature of approximately 45 °C for 4–5 days until all the ammonia has evaporated. Table 2. Auricularia auricula and Auricularia polytricha commonly are produced on a medium consisting of sawdust. millet. Chang and Hayes 1978. the containers are filled with moistened substrate and sterilized or pasteurized prior to inoculation. Spawn runs last about 30 to 60 days depending on strain and substrate formulation. and Cryptomeria japonica are . and mycelium can develop in either the same or another tunnel. After the completion of vegetative mycelial growth. nameko cultivation is similar to that for F. This method has been replaced by artificial log cultivation that utilizes heat-treated substrates based on sawdust enclosed in plastic bags (“bag-log cultivation”). Phase III then yields fully grown compost ready for production (Van Griensven and Van Roestel 2004). bran. the collars are removed. Fig. Cultivation of Auricularia sp. marmoreus. When the mushrooms are mature. and the bags are heat-sealed and shaken to uniformly distribute the spawn throughout the substrate. cottonseed hulls. When the mushrooms are 13 to 14 cm long. The mixed substrate is sterilized for 60 min at 121 °C. the moistened substrate is filled into microfiltered polypropylene bags and sterilized. sterilized (4 h at 95 °C and 1 h at 121 °C). The inoculation and fructification are carried out as previously reported (Royse 2007). and other cereal grains or on natural logs of broad-leaf trees (Table 2). This formula gave the highest yields. The main advantages of this method are the short time to complete a crop cycle and the higher yields (Sánchez 2004). edodes L. the compost can be spawned. it was traditionally grown on wood logs (Stamets 1995. Both phase I and phase II of the composting process can take place in bulk in special fermentation rooms called tunnels. Recently. and the mushrooms are pulled as a bunch from the substrate (Royse 2007). Production of F. One of the most important developments in mushroom growing of recent years has been the introduction of bulk processes in composting technology. As the mushrooms begin to elongate above the lip of the bottle. the substrate may be composted for up to 5 days or used directly after mixing. velutipes (Furukawa 1987). A common substrate used for production is composed of sawdust supplemented with rice bran or wheat bran. Cultivation of Hypsizygus marmoreus The Japanese are the main producers and consumers of H. The bottles then are refilled with fresh substrate. Sánchez 2004). For synthetic medium production. mushroom culture has moved toward diversification. Only one flush of mushrooms is harvested prior to mechanical removal of the “spent” substrate from the bottles. For bottle production. After cooling (16 to 20 h). edodes is the third most cultivated edible mushroom. Cultivation of Grifola frondosa Production of G. and shortest crop cycle time (12 weeks).

fuciformis to digest the substrate thereby increasing mushroom yields. There are several companies which provide different environmental control systems. automatic control of ammonia levels and moisture in the casing soil is still not widely applied. archeri increases the ability of T. 1995. which include small.. Obatake et al. modular semiautomatic picking aids and the fully automated. 2002. 1999). Saikai et al. most of the strains currently used are similar to the first hybrid obtained (Sonnenberg 2000. automated mushroom harvesting machines. Harmsen et al. cotton wastes (discarded after sorting in textile mills) have become popular as substrates for straw mushroom production (Chang 1982). The mixed culture technique involves the use of “helper” mycelium of Hypoxylon archeri. resistance to bacterial diseases (Oliver and Delmas 1987. the first climate computers were introduced to Dutch mushroom growing and now are widely used in the industry (Lamber 2000). Cultivation techniques used to produce the mushroom on natural logs is similar to that used for shiitake production. Some companies offer different automated harvesting systems. It can be grown on non-pasteurized substrate which is more desirable for low input agricultural practices.1330 Appl Microbiol Biotechnol (2010) 85:1321–1337 suitable for growth. In recent years. Sánchez 2004). pressure. Kerrigan 2000). However. Rice bran usually is added as a supplement for conifer sawdust and 10% for broad-leaf sawdust (Royse 2007). Development of technology to increase productivity To increase productivity in the mushroom culture. The consistency and texture of the mushroom are crucial to diminish losses during packing. or to develop methods to produce mushrooms on a non-composted substrate. Senti et al. Research on the molecular basis of mating-type genes has been very important for the development of strains with high yields. and new techniques of sterilization (Hawton et al. Senti et al. 2000). Jaybird Manufacturing. fuciformis has been on a substrate using a mixed culture inoculum technique first developed in Fujian. an ascomycete commonly associated in nature with decaying wood. Moquet et al. Kerrigan 2000. Agricultural Engineering. important advance in the development of techniques for breeding is based on the development and detection of genetic markers (Sonnenberg 2000). 1995. The computer monitors environmental parameters such as temperature. Cotton wastes give higher and more stable biological efficiencies (30% to 45%) and earlier fructification (4 days after spawning) and harvesting (first 9 days after spawning) than that obtained using straw as a substrate. A strain with a high ability to invade the substrate and to fruit diminishes time of incubation and enhances productivity. viral diseases (Sonnenberg et al. e. China (Huang 1982). several studies have been carried out to develop automated mushroom harvesting machines. 3k) constitute the most expensive part of the overall production process. Tewari 2007). DuraFlex. The first A. H. In recent years. hydroblending and pre-wet equipment (heap turners). The culture of other mushrooms has also been reported (Table 2). 2000). The use of DNA-based technology has accelerated breeding activities and will help mushroombreeding programs (Stoop and Mooibroek 1999). A computerized environmental control system is an invaluable tool for mushroom culture. More than two decades ago. cultivation is of particular importance.g. spores (Laborde 1995. robotic systems designed for line picking (Kensal Improvement of strains The strain used in the culture is crucial for success of mushroom production and marketing. most production of T. Cultivation of Volvariella sp. etc. However. bisporus hybrid was obtained by breeding about 25 years ago. The increase in mushroom production has been the result of more specialized studies carried out by several international institutions in different areas of mushroom cultivation (Tables 3 and 4). it is necessary to develop and improve control and computerized monitoring of growing rooms. This mushroom is well suited for cultivation in the tropics because of its requirement for higher production temperatures. The strain used for Pleurotus sp. Mori et al. 1990). humidity. 2000). 1991). Picking and packing (Fig. 1998. and fungal diseases (Dragt et al. and carbon dioxide and oxygen content. airflow. similar symptoms have also been observed with L. Exploitation of this mycelial association is accomplished through use of dual cultures to make mother spawn (Quimio 1997). This problem has increased interest in developing sporeless mutants for breeding of sporeless strains (Laborde 1995. However. since some workers develop an allergy that is identical to “mushrooms workers lung” and is associated with exposure of people to Pleurotus spp. Cultivation of Tremella fuciformis This mushroom can be cultivated on natural logs or on a medium (Quimio et al. Climate control in production plants allows monitoring and control of several mushroom production rooms with a minimum human input. edodes spores (Laborde 1995. 2003. Computerized environmental control system allows the grower to look and adjust the environmental conditions of the plant by using remote access (modem) capabilities (Walker 1996). An . In the last 20 years.

Japan. INRA National Institute of Agronomy Research. Astell 1996). USA. FSU Fredrick Schiller Automation. Israel. CAL College of Agriculture Laguna. The harvesting operation includes mushroom location. disease and pest resistance Increased productivity and reduced environmental pollution -doIncreased productivity at lower costs Increased productivity Indigenous technology Indigenous technology Utilization of waste and additional returns to the growers Diversification of the mushroom portfolio of the country RBG. CMI CMI. ESTs. Germany Source: Tewari 2007. Mori Institute. PSU INRA. MES HRI. MES. PSU RBG Royal Botanical Gardens. The Netherlands. allele mining. MI. ATCC American Type Culture Collection. MI. PSU HRI. PSU PSU Understanding will help manipulation for improvement in yield and quality libraries of mushrooms Basic information on the genome structure Industrial applications Better disease management Microbes for rapid composting Domestication of newer types and increasing the yield of commercial types Utilization of wastes for food. and the resulting pilot harvester was successfully tested on a . INRA. UK. Malaysia PSU. MI. PSU. MES HRI. HRI PSU. France. The mechanical properties used for the analysis of automated harvesting were obtained from compression experiments with cylindrical mushroom sample pieces (Hiller 1994). improved quality. ATCC HRI. MES INRA HRI. and identification of wild relatives and new species Culture maintenance Genetic improvement Improved production systems State-of-the art composting technology Reduction in composting period for rural growers Newer and improved casing layer Environment control in growing houses Development of farm designs Mechanization and automation Utilization of spent straw Improved cultivation technology Seed (spawn) Development of liquid spawn technology Improved spawn preparation and packaging technology Integrated pest and disease management Survey and preparation of area specific disease map Integrated pest and disease management Investigations on mushroom viruses Integrated pest management in mushrooms Postharvest technology of mushrooms Improvements in packaging and practices for storage and transport of fresh mushroom Development of improved processing technologies for mushrooms Development of production technologies for valueadded products from mushrooms Basic and strategic studies in mushrooms Molecular genetics—mapping. and picking. UK. PSU Pennsylvania State University. creation of gene libraries Genomics—genome sequencing. and fuel To develop appropriate environment for higher yields HRI.Appl Microbiol Biotechnol (2010) 85:1321–1337 Table 4 International institutions that study different areas of mushrooms cultivation and their expected output Area of study in mushrooms production Expected output International institutions 1331 Germplasm and genetic improvement Survey. MES Mushroom Experimental Station. PSU. UK. USFPL US Forest Products Laboratory. Reed et al. cloning and sequencing of genes. PSU HRI. INRA HRI USFPL. HRI Horticulture Research International. Hong Kong. CU Chinese University.nrcmushroom. HRI Ease of handling and low transportation cost Improved spawn quality resulting in higher yields and returns Evolving suitable strategies against mushroom diseases Increased yields and reduced residual chemicals Increase productivity Increase productivity Reduced postharvest losses Reduced postharvest losses and avoidance of distress sale Development of value-added products INRA INRA HRI. Horst. CMI Commonwealth Mycological Institute. feed. (1995) evaluated at the laboratory level the automated harvesting of A. bisporus by machine. CAL. FSU HRI. MES CINADCO PSU. sizing. and microarrays Cloning and expression of useful genes of mushroom Molecular basis of host-pathogen interaction Role of microflora in composting Morphogenesis in mushrooms Molecular mechanisms of biodegradation of lignocelluloses Studies on environmental physiology of mushrooms Assessment of biodiversity and conservation of germ plasm Conservation of germ plasm Increased productivity. http://www. selection. PSU. Shatin. USA. CU. USA. PSU HRI. collection. CINADCO Centre for International Agricultural Development Cooperation. Philippines.

P. The mycocell system has allowed also the successful culture of exotic mushrooms such as L.1332 Appl Microbiol Biotechnol (2010) 85:1321–1337 commercial mushrooms farm. Hypsizygus sp. edodes. there are several specialized institutes that study different frontier areas to improve mushroom cultivation (Tables 3 and 4). Ranganathan and Selvaseelan 1997. Söchting and Grabbe 1995. 2000. G. A technology called variable frequency speed control for alternating current motors (Keljik 1995) may be useful for mushroom farms and benefit the industry by improved control of air velocity in the growing environment. a gripper mechanism is finally used to remove mushrooms from the conveyor into packs at the side of the machine (Reed et al. or both) should be used (Tillett and Batchelor 1991. Pleurotus cystidiosus. Table 5). After high speed trimming. reducing operating and capital costs. Ca2SO4) can be added. bisporus spent substrate has also been used for the cultivation of Volvariella (Poppe 2000). Verdonck 1984. increasing production yields. crop uniformity. The mycocell system (Mycocell Technologies. ostreatus. 2000). 2008). resulting in lower electricity consumption (Lomax 1989. automation. 1994. Hericium sp. e. Tajbakhsh et al. lucidum and F. there are no risks of substrate contamination. One of a pair of suction cup mechanisms attached to the single head of a Cartesian robot is then deployed. The apparatus combines several handling systems and mechatronic technologies.g. The pre-wet heap machine was designed to achieve a quick homogeneous mix of water and the raw materials used in phase 1 of the culture of A. 1984. light and cheap transport. Delver and Wertheim 1988. Singh et al. Sánchez 2004). 1995. several nutrients can be added and mixed thoroughly. the use of spent Agaricus compost added with cotton waste for Volvariella production (Oei 1991). . Shandilya 1989. F. AntSaoir et al. reducing phase 1 time by over 50%. Utilization of spent straw generated after the cultivation of G. the use of spent Pleurotus substrate for the King Stropharia cultivation (Poppe 1995). velutipes has been recommended for the production of other mushrooms. it can delicately detach individual mushrooms and place them gently into a specially designed compliant finger conveyer. and long shelf life.. Chong and Rinker 1994. Díaz-Godínez and Sánchez 2002). Díaz-Godínez and Sánchez (2002) found that addition of maize straw generated after mushroom cultivation to the diets of sheep increased the weight gain of the sheep and the efficiency of feed conversion of the straw. Szmidt 1994. which consists of filling the bunkers or tunnels using overhead layering techniques. bisporus or Coprinus comatus (Xiao 1998). and the colonization of the substrate by the mycelium considerably increases. 1998.. less water run-off. 1997). SMS is also a beneficial product for enrichment of soils. 2000. twist. restoring areas that have been destroyed through development. low contamination risk. Batista et al. Díaz-Godínez and Sánchez 2002) and digestibility by the ruminant (Capelari and Zadrazil 1997. Celikel and Tuncay 1999). since its degradation by the mushroom can improve its nutritional quality (Jalc et al. A. and biogas production (Balan et al. Currently. lack of downtime. frondosa. and profit potential. radiation used in the treatment of the substrates modifies the cellulose and increases ease of breakdown by the mycelium. b. Some benefits of using pre-wet compost turners include improving compost quality. Ganoderma lucidum. 1993. low labor demand. UK) is a method based on microwave sterilization of pre-packaged substrate to which the “spawn” and other nutrients (e. the use of SMS enriched with cotton seed meal and with soya meal for Agaricus cultivation. in soil amendment as organic manure or vermi-compost (Stewart et al. Agrocybe aegerita. better odor control. 1998. in nursery and landscape gardening (Chong and Wickware 1989. 2001). Some studies have been done on the use of SMS in vegetable and flower greenhouses (Lohr et al. Design of new equipment to improve productivity is continuously in development. Pholiota sp. a new tunnel/bunker filling system has been developed. low energy requirements. Pleurotus eryngii. SMS can also be used as casing material for the production of Agaricus (Nair and Brandley 1981. An expert selection algorithm then decides the order in which they should be picked and what picking action (bend. 1992.. 2000). It has also been used as animal feed. Adamovic et al. and Auricularia (Hawton et al. homogenous blending of up to 200 t per hour. 1995. Pill et al. Additional benefits of mushrooms culture Several studies have shown that spent SMS can be used for mushroom re-cultivation. Recently. 1992. Psilocybe sp. deforestation. with the use of overhead and out of sight conveyors/elevators (Traymater machinery).. Chong 1999). but only from one end of the bunker/tunnel and not through holes in the tunnel roof. In this case. bisporus to improve efficiency of the composting process by reducing the amount of time required to achieve the same or better quality compost. Stewart et al. New methods to produce and increase the productivity of a wide variety of exotic mushrooms have been developed. In addition. Reed et al. Steffen et al.. and the use of spent Pleurotus eous straw for the cultivation of several species of this fungus (Siddhant and Singh 2009). raw material savings. velutipes. Chang and Miles (1989) studied the use of spent Volvariella substrate for the production of Pleurotus sajor-caju. such as A. 1998. in field vegetable and fruits crop (Male 1981. or environmental contamination. Pleurotus djamor..g. Mushrooms are located and sized using image analysis and a monochromatic vision system. 1996a. The commercial benefits of this production system include lower cost. Pleurotus pulmonarius.

Pleurotus the compost resources/pests/index. Lau et al. Queen's University of Belfast. Mycogone perniciosa Magn. Xawek et al. Meijer 2009. 2009). American Mushroom Institute 2006. Table 6 Some diseases that attack Pleurotus ostreatus and Agaricus bisporus cultivation Affected mushroom Agaricus bisporus Agaricus bisporus Agaricus bisporus Agaricus bisporus Pleurotus ostreatus Agaricus bisporus Infestation (organism) Cladobotryum dendroides Mycogone sp. Ontario. The potential of SMS to degrade organopollutants and its importance in environmental bioremediation has been reported (Kuo and Regan 1998..Appl Microbiol Biotechnol (2010) 85:1321–1337 Table 5 Specialized institutes that study mushroom cultivation in different frontier areas Frontier areas Genetic improvement and transgenic technologies Identification of mushrooms DNA technologies Compost technology Pest and disease management Cultivation of specialty mushrooms Quality and packaging technologies and many others frontier areas Computer application in mushrooms Institute 1333 MES. blotch and necroses of caps and stems Boiko et al. Ireland INRA. Law et al. Cephalothecium disease Brown blotch Blotch disease and other Pseudomonas mushrooms gingeri Agaricus bisporus Agaricus bisporus Putative virus X Bacilliform viruses Virus Virus Ginger blotch Virus X Western Committee on Plant Disease Brown irregular pitted areas on stems and (2004). 2003). Centre for Plant Biotechnology. 2003. Trichoderma sp. Tanović et al. 2009. Canada 2008.shtml . HRI. Lycoriella sp.mushroomcompany. and Smith et al. 2001 and a slight surface depression that becomes darker with age Pale yellowish red discoloration that develops into a reddish ginger-colored pale discoloration Slightly imperfectly formed cap structure Gaze 2001 Severe rotting. 2003.mushroomcompany. Boiko et al.mushroomcompany. 2009 Fungi Green mold Dark green mold patches on casing spreading to lesions on stems Western Committee on Plant Disease (2004) Velázquez-Cedeño et al. Severe resources/pests/index. 1998 . 2009 Mild dark purple to light brown discoloration Godfrey et al. 2009.shtml. Kingston. 2009. ingenua Megasellia halterata Aphelenchoides composticola Pseudomonas Tolaasii Pseudomonas reactants Insect Sciarid flies Insect Phorid flies Nematode 2009 flies resources/pests/index. http://www.nutrasoils. Jess and Schweizer 2009 maturing mushrooms Cause less mushroom damage than sciarid http://www. 2006 Degeneration of the mushroom mycelium in http://www. 2009. fungicola Fungi Dry bubble/brown spot Agaricus bisporus Agaricus bisporus Agaricus bisporus Agaricus bisporus. Webb et al. Tanović et al. PSU CMI HRI.nrcmushroom. Currently. http://www. PSU. PSU INRA.mushroomcompany. 2009 Boiko et al. Gitanjali 2005 Sunken. there are some industries that manufacture and sell different kind of compost based on SMS (http://www. dark brown lesions http://www. mildew Wet bubble/white mold White to pink cobweb-like fluffy mold Dense white growth on gills Wet bubble of fungi Western Committee on Plant Disease (2004) Western Committee on Plant Disease (2004). Eggen 1999. Distortion and splitting. Pathak et al. 2009. USA Source: Tewari (2007).shtml. blotch. laurelvalleysoils. burrow or tunnel into the stems and caps of resources/pests/index. PSU PSU University of Rhodes Island. and necroses of caps and stems Destroy pins of developing mushrooms. Revill et al. 2004 Verticillium sp. L. Type of organism Fungi Fungi Fungi Diseases Symptoms Reference Cobweb.

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