Abstract
Pleurotus species are edible mushrooms that possess the ability to degrade lignocellulose and are cultivated on lignocellulosic substrates to produce fruiting bodies that are appreciated for their nutritional and culinary values. Radial growth rate (ur), mycelial biomass, and intracellular glycogen and protein contents were evaluated in the colonies of two strains of P. ostreatus (strains 26 and 50) and one strain of P. pulmonarius (strain 35) grown on barley straw extract agar, corn stover extract agar (CSEA), wheat straw extract agar, and glucose-yeast extract agar (GYEA). The nutritional composition of the Pleurotus fruit bodies cultivated on barley straw, corn stover, and wheat straw were also determined. Of all of the strains, P. ostreatus 50 showed the highest ur on the CSEA and GYEA, and P. ostreatus 26 showed the greatest mycelial biomass production on the lignocellulosic waste-based agar media (LWAM). All of the Pleurotus strains showed the greatest and lowest glycogen contents on the CSEA and GYEA, respectively. These results showed that the LWAM are more suitable for mycelial growth than a glucose-based agar medium, and that Pleurotus fruit bodies cultivated on corn stover had the highest nutritional value compared with fruit bodies grown on the wheat and barley substrates.
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Mycelial Growth and Fruit Body Nutritional Composition ofPleurotus Species Grown on Different Lignocellulosic Waste-based Media
Jarim Leonardo Saavedra-Molina,a Daniel Méndez-Iturbide,b Madaí Angélica Gómez-Camarillo,c and Carmen Sánchez d,*
Pleurotus species are edible mushrooms that possess the ability to degrade lignocellulose and are cultivated on lignocellulosic substrates to produce fruiting bodies that are appreciated for their nutritional and culinary values. Radial growth rate (ur), mycelial biomass, and intracellular glycogen and protein contents were evaluated in the colonies of two strains of P. ostreatus (strains 26 and 50) and one strain of P. pulmonarius (strain 35) grown on barley straw extract agar, corn stover extract agar (CSEA), wheat straw extract agar, and glucose-yeast extract agar (GYEA). The nutritional composition of the Pleurotus fruit bodies cultivated on barley straw, corn stover, and wheat straw were also determined. Of all of the strains, P. ostreatus50 showed the highest ur on the CSEA and GYEA, and P. ostreatus26 showed the greatest mycelial biomass production on the lignocellulosic waste-based agar media (LWAM). All of the Pleurotusstrains showed the greatest and lowest glycogen contents on the CSEA and GYEA, respectively. These results showed that the LWAM are more suitable for mycelial growth than a glucose-based agar medium, and that Pleurotus fruit bodies cultivated on corn stover had the highest nutritional value compared with fruit bodies grown on the wheat and barley substrates.
Keywords: Fruit body; Lignocellulosic wastes; Mycelium; Pleurotus ostreatus; Pleurotus pulmonarius
Contact information: a: Maestría en Ciencias en Sistemas del Ambiente, Universidad Autónoma de Tlaxcala, Ixtacuixtla, Tlaxcala, México; b: Facultad de Nutrición, Universidad Autónoma de Tlaxcala, San Pablo del Monte, Tlaxcala, México; c: Facultad de Odontología, Universidad Autónoma de Tlaxcala; d: Laboratory of Biotechnology, Research Centre for Biological Sciences, Universidad Autónoma de Tlaxcala, Ixtacuixtla, Tlaxcala, México, CP 90120;
* Corresponding author: sanher6@hotmail.com
INTRODUCTION
Pleurotus species are edible mushrooms that are appreciated for their exquisite flavor and economic and ecological values. Pleurotus ostreatus (oyster mushroom) is the second most cultivated edible mushroom worldwide after Agaricus bisporus (Sánchez 2010). Pleurotus ostreatus is cultivated in colder places (≈ 20 C) than P. pulmonarius, which is a tropical species of edible oyster fungi. These organisms are basidiomycetes and have two phases of growth (vegetative and reproductive), each with specific characteristics and requirements. The vegetative phase or mycelial growth is seen as microscopic filaments or hyphae that absorb digestive products and colonize the substrate (e.g. agar or lignocellulosic substrates) (Philippoussis 2009). The reproductive phase corresponds to the fruit body and it is represented by the mushroom itself (oyster shaped), which is formed after the fusion of two primary compatible mycelia with reciprocal nuclei exchange (Nieuwenhuis et al. 2011). Pleurotusspecies have the ability to degrade several lignocellulosic substrates because of their highly efficient enzymatic machinery (Bilal et al.2017; Hernández-Dominguez et al. 2017). These species can be cultivated on a wide variety of substrates, such as sugar cane bagasse, corn stover, wheat straw, rice straw, and barley straw, which are some of the largest accumulated lignocellulosic wastes because of agricultural activity (Sánchez 2010; Jeznabadi et al. 2016; Sadh et al. 2018). Lignocellulosic residues are generated as agricultural by-products, which are disposed of by burning in fields (Saini et al.2015). Therefore, cultivation of edible mushrooms might be the only current process that combines the production of protein-rich food with the reduction of environmental pollution (Sánchez 2010). It represents one of the most efficient biotechnological processes for lignocellulosic residue recycling (Mandeel et al. 2005). Substrate and fungal strains are very important for mushroom cultivation because an appropriate substrate and fast-growing strain promote a rapid mycelial invasion of the substrate, which avoids the risk of contamination in the cultivation process. Pleurotus species are a rich source of protein, minerals (P, Ca, Fe, K, and Na), and vitamins (thiamine and riboflavin) (Sánchez 2010; Deepalakshmi and Mirunalini 2014). However, the nutritional composition of the fruit body depends on the substrate on which the mushroom is cultivated (Sánchez 2010). Pleurotus species have high nutritional values and contain various bioactive compounds that have been isolated and identified from both the fruit body and mycelium (Lavi et al. 2012; Maftoun et al. 2013; Ganeshpurkar et al. 2015). Additionally, the fungal cell wall contains β-glucans, which have antioxidant and immunomodulatory properties (Maftoun et al. 2013; Ganeshpurkar et al. 2015).
In this work, the radial growth rate, mycelial biomass production, intracellular glycogen content, and intracellular protein content were evaluated in the colonies of two strains of P. ostreatus and one strain of P. pulmonarius grown on lignocellulosic waste-based agar media and a glucose-yeast extract agar medium. The nutritional composition of the fruit bodies of the three Pleurotus strains cultivated on barley straw, corn stover, and wheat straw were also determined.
EXPERIMENTAL
Strains
Three strains of Pleurotus were used: P. ostreatus NRRL 3526 (Po 26) from the NRRL (now ARS) culture collection (Peoria, USA), P. ostreatus 50 from the COLPOS-Puebla culture collection (Puebla, Mexico), and P. pulmonarius 35 (PL35) from the culture collection at the Chinese University of Hong Kong (Hong Kong, China). Stock cultures were grown on malt extract agar in the dark at 25 C in petri dishes for 7 days, which were then stored at 4 C. Cultures were periodically transferred to fresh culture media.
Culture Media and Culture Conditions
Three lignocellulosic waste-based agar media were prepared: barley straw extract agar (BSEA), corn stover extract agar (CSEA), and wheat straw extract agar (WSEA). The media were prepared as follows: 100 g of dry chopped lignocellulosic material (bought from local farmer’s markets) were boiled in 1 L of tap water for 1 h. The lignocellulosic material was filtered through a double layer of muslin and discarded. The liquor was placed in glass containers and the volume was made up to 1 L, and then it was autoclaved at 120 C for 30 min. Prior to autoclaving, a final concentration of 20 g/L agar was added. The pH was adjusted to 6.5. Glucose-yeast extract agar (GYEA) was used as the control medium. It contained glucose (10 g/L), yeast extract (5 g/L), KH2PO4 (0.6 g/L), MgSO4-7H2O (0.5 g/L), K2HPO4 (0.4 g/L), CuSO4-5H2O (0.25 g/L), FeSO4-7H2O (0.05 g/L), MnSO4 (0.05 g/L), and ZnSO4-7H2O (0.001 g/L). All of the media were poured into petri dishes, which were inoculated in the center with an inoculum (4 mm in diameter) taken from the periphery of a 7-d old colony grown on the malt extract agar. The agar plates were incubated at 25 C for 7 day.
Radial Growth Rate and Mycelial Biomass
The colony diameters were measured from the second to seventh day of incubation. Radius versus time plots were analyzed by linear regression, and the slope for each treatment was recorded as the radial growth rate (ur) (González-Márquez et al. 2015). The mycelial biomass was determined from the 7-day old colonies. Mycelium was separated from the culture medium and filtered using a boiling water bath, as previously reported by González-Márquez et al. (2015).
Determination of the Intracellular Glycogen and Intracellular Protein Contents
The contents of the intracellular glycogen and protein were determined in the mycelia of the colonies. The intracellular mycelial extract was obtained from the mycelium scraped from the agar and ground in 1 mL of distilled water for each 0.1 mg of fresh weight biomass using a tissue grinder (Pyrex glass, Corning Inc., Corning, NY, USA). The resulting samples were centrifuged at 20000 × g for 10 min at 2 °C. The supernatant was collected as the intracellular extract and the sediment was the cell wall. In the intracellular extract, the total intracellular protein content (mg/g of dry weight (DW)) was measured by the method by Bradford (1976), using bovine serum albumin as the standard. The glycogen content (mg/g of DW) was the difference between the total sugars and reducing sugars (Sánchez et al. 2004).
Fruit Body Production
Spawn preparation
The P. ostreatus spawn (i.e. the seed required for growing the mushrooms) were prepared using wheat grains, which were previously soaked in water over night and then drained. Plastic bags (30 cm x 20 cm) were filled with 500 mg of soft and moist wheat grains and autoclaved at 121 C for 30 min. After cooling down to room temperature, the bags were inoculated with three mycelial agar fragments (10 mm in diameter) taken from a fully-grown colony of P. ostreatus. The inoculated bags were incubated at 25 C for 7 days (Sánchez 2010).
Substrate preparation, inoculation, incubation, and harvest
Lignocellulosic residues (with lengths of about 2 cm to 6 cm) were milled, immersed in tap water, and then pasteurized at 85 C for 1 h. The pasteurized lignocellulosic substrate was placed on a wire mesh to drain off the excess water. The lignocellulosic substrate moisture content was adjusted to 65%. Next, 5 kg of pasteurized substrate were packed into polythene bags, inoculated with 10% spawn, and then incubated at 25 C for 2 weeks. Completely mycelial-invaded bags were moved to a cropping room, where the relative humidity was 85%, the temperature was 25 C, and the light intensity was about 100 lux. The cropping room was watered to maintain the moisture content during the cropping time. Mushroom harvest was performed from each of the substrate bags after 7 days of being in the cropping room (Sánchez 2010).
Nutritional Composition of the Fruit Bodies
The harvested mushrooms were dried in an oven at 60 C to a constant weight to calculate the moisture content, and then the dried mushrooms were ground into a powder with a mortar and pestle for other analyses. Mushroom samples were analyzed to determine their nutritional composition. The ash, protein, carbohydrate, fat, and fiber contents were determined using the procedures from AOAC (1995). The ash content was measured by incineration at 600 C using a muffle furnace. The protein content (N × 6.25) was evaluated by the macro-Kjeldahl method. The phenol-sulfuric acid method was used to calculate the total carbohydrates content. The fat content was estimated by extracting a known weight of a powdered sample with ethyl ether using a Soxhlet apparatus (FC20, SEV PRENDO, Mexico City, Mexico).
Statistical Analysis
Each treatment was performed in triplicate. The data was evaluated with a one-way analysis of variance (ANOVA) and Tukey post-test using the Graph Pad Prism® program (San Diego, CA, USA).
RESULTS AND DISCUSSION
Radial Growth Rate and Mycelial Biomass
Table 1 shows the ur of the Pleurotus sp. grown on the lignocellulosic waste-based agar media and GYEA. The P. ostreatus 26 strain showed the greatest ur on the BSEA, CSEA, and WSEA media, and the lowest ur on the GYEA medium. The P. ostreatus 50 strain had the highest ur on the CSEA medium, followed by the WSEA, BSEA, and GYEA media. The P. pulmonarius 35 strain showed the highest ur on the CSEA and WSEA media, followed by the GYEA and BSEA media. Of all of the strains, P. ostreatus 26 showed the greatest ur (0.29 mm/h to 0.3 mm/h) on the lignocellulosic waste-based agar media. Sánchez and Viniegra-González (1996) reported that the ur of a highly productive strain of P. ostreatus grown on a starch agar medium was 0.298 mm/h. Téllez-Téllez et al. (2003) reported that the ur for P. pulmonarius grown on a potato dextrose agar (PDA) medium was 0.099 mm/h. The results of this study showed that the lignocellulosic waste-based agar media promoted rapid mycelial growth.
The mycelial biomass production by the Pleurotus strains is shown in Table 2. The P. ostreatus 26 strain had the highest mycelial biomass production on both the CSEA and WSEA media, and the lowest mycelial biomass production on the BSEA and GYEA media. The P. ostreatus 50 strain showed the highest mycelial biomass on the CSEA and GYEA media, followed by the WSEA and BSEA media. The P. pulmonarius 35 strain had the highest mycelial biomass production on the CSEA medium, followed by the WSEA and GYEA media, and the lowest mycelial biomass was produced on the BSEA medium. Of the three strains, P. ostreatus 50 showed the highest mycelial biomass production on the CSEA and GYEA media (100 mg/cm2). González-Márquez et al. (2015) reported that the mycelial biomass production by Lentinula edodes (a slow-growing mushroom) grown on a GYEA medium was 0.020 mg/cm2. Téllez-Téllez et al. (2003) reported that the mycelial biomass production for P. pulmonarius on a PDA medium was 0.26 mg/cm2. In general, it was shown that the lignocellulosic waste-based agar media were more suitable for growth of the Pleurotus sp. than the GYEA.
Table 1. Radial Growth Rate of the Pleurotus sp. Colonies Grown on the Lignocellulosic Waste-based Agar Media
Values are in mm/h and expressed as the mean ± SD (n = 3); means within the same column not sharing common superscript letters (a to c) differ significantly at a 5% level
Table 2. Mycelial Biomass Production by the Pleurotus sp. Colonies Grown on the Lignocellulosic Waste-based Agar Media
Values are in mg/cm2 and expressed as the mean ± SD (n = 3); means within the same column not sharing common superscript letters (a to c) differ significantly at a 5% level
Intracellular Protein and Glycogen Contents
The strains P. ostreatus 26 and P. ostreatus 50 had the greatest intracellular protein content in the CSEA medium, and their lowest intracellular protein content was seen on the BSEA, WSEA, and GYEA media. The P. pulmonarius strain showed the highest intracellular protein content on the lignocellulosic waste-based agar media, and the lowest intracellular protein content on the GYEA medium. The protein intracellular content was around 2- and 3-fold higher for P. pulmonarius and P. ostreatus, respectively, on the CSEA medium compared with the GYEA medium (Table 3). González-Márquez et al. (2015) reported that the intracellular protein content for L. edodes was 0.04 mg/g of DW. In the present research, the highest intracellular protein content (0.19 mg/g of DW) was measured with P. ostreatus 50 grown on the CSEA medium.
Table 3. Intracellular Protein Content of the Pleurotus sp. Colonies Grown on the Lignocellulosic Waste-based Agar Media
Values are in mg/g of DW and expressed as the mean ± SD (n = 3); means within the same column not sharing common superscript letters (a to c) differ significantly at a 5% level.
The intracellular glycogen content of the Pleurotus sp. colonies is shown in Table 4. All of the Pleurotus sp. showed the greatest and lowest glycogen contents on the CSEA and GYEA media, respectively. Of all of the strains, Pleurotus 50 had the highest intracellular glycogen content on the lignocellulosic waste-based agar media (91.9 mg/g of DW to 106.9 mg/g of DW) and P. pulmonariushad the lowest glycogen content on the GYEA medium (51.3 mg/g of DW). Lentinula edodes produced an intracellular glycogen content around 30 mg/g of DW on the GYEA medium (González-Márquez et al. 2015). These results showed that the CSEA medium increased both the mycelial growth and mycelial density for all of the strains tested. Furthermore, the media prepared using the lignocellulosic substrates are cheaper than commercial culture media (i.e. glucose-based media).
Table 4. Intracellular Glycogen Content of the Pleurotus sp. Colonies Grown on the Lignocellulosic Waste-based Agar Media
Values are in mg/g of DW and expressed as the mean ± SD (n = 3); means within the same column not sharing common superscript letters (a to d) differ significantly at a 5% level
Nutritional Composition of the Fruit Bodies
Table 5 shows the nutritional composition on a dry matter basis for the fruit bodies cultivated on the barley straw, wheat straw, and corn stover. The fruit bodies from both strains of P. ostreatus cultivated on the barley straw achieved higher moisture contents than those cultivated on the wheat and corn substrates. The P. pulmonarius fruit bodies formed on the wheat straw had higher moisture contents than those grown on the barley and corn substrates. In general, the fruit bodies from both strains of P. ostreatus grown on the wheat straw showed greater ash (mineral salts) and fiber contents than those grown on the barley and corn substrates. The P. pulmonarius fruit bodies cultivated on the barley straw had the highest ash and fiber contents, followed by those formed on the wheat straw and corn stover. The Pleurotus sp. fruit bodies cultivated on the barley straw had the highest fat content compared with those grown on the wheat and corn substrates. For all of the strains, the highest protein and carbohydrate contents were recorded from the fruit bodies cultivated on the corn stover. The selection of the substrate during mushroom growth is crucial to promote rapid mycelial growth, avoid the risk of contamination, and enhance mushroom production (Royse 2002).
These results showed that the mushroom fruit bodies grown on the corn stover had the highest nutritional value compared with those grown on the barley and wheat substrates, and that the Pleurotus fruit bodies are rich in proteins, carbohydrates, and fiber, and low in fat. The Pleurotus nutritional properties determined in the present research were similar to those reported previously for Pleurotus sp. grown on different lignocellulosic substrates (Hoa et al. 2015). The nature and nutrient composition of the substrate affected the mycelial growth and fruit body nutritional properties (Philippoussis 2009). Obodai et al. (2003) studied the growth of P. ostreatus on different substrates and reported that rice husk resulted in the fastest mycelial growth rate. However, it was not a suitable substrate for fruit body production. The cultivation of P. ostreatus on maize stover results in higher fruit body yields than on rice husk (Obodai et al. 2003). In the present research, corn stover was a better substrate than barley and wheat straw in terms of the nutritive value for mushroom mycelial growth and fruit body cultivation. Corn stover is the above-ground portions of a corn plant, including the stalks, cobs, leaves, and husks (excluding the corn kernels themselves) (Wyman 2003). The glucan content of corn stover is around 40%, which could release up to 44.4 lb of glucose after hydrolysis for every 100 lb of dry feedstock (Wyman 2003). It also contains xylan, arabinan, galactan, and mannan. The glucan content of wheat straw and barley straw is 30.2% and 33.1%, respectively. Wheat straw and barley straw also contain xylan, galactan, and arabinan (Ballesteros et al. 2006; Garcia-Aparicio et al. 2006). This showed that corn stover has the best nutritional composition of all of the substrates tested. The selection of substrate for mushroom production is mainly determined by the abundance and cost of the substrate (Obodai 2003). Corn stover represents a good alternative for mushroom cultivation. It is one of the most abundant lignocellulosic substrates in the world (Sánchez 2009) and the most abundant in Mexico (Serna-Saldivar 2016).
Table 5. Nutritional Composition of the Pleurotus sp. Strains Cultivated on the Different Lignocellulosic Substrates
Values are in g/100 g of dry matter and expressed as the mean ± SD (n = 3); means within the same column for each oyster mushroom not sharing common superscript letters (a to c) differ significantly at a 5% level
CONCLUSIONS
- Lignocellulosic waste-based agar media are more suitable for mycelial growth than a glucose-based agar medium.
- The CSEA medium increased both the mycelial growth and mycelial density for all of the strains tested.
- The Pleurotus fruit bodies cultivated on the corn stover had the highest nutritional value compared with the fruit bodies grown on the wheat and barley substrates.
- Corn stover was the best lignocellulosic substrate for culture maintenance and fruit body cultivation of the tested substrates.
ACKNOWLEDGMENTS
The authors are grateful to the Mexican Council for Science and Technology (CONACyT) for providing a MSc scholarship (No. 648826) to Jarim Leonardo Saavedra-Molina.
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Article submitted: December 29, 2017; Peer review completed: March 17, 2018; Revised version received: March 18, 2018; Accepted: April 2, 2018; Published: April 18, 2018.
DOI: 10.15376/biores.13.2.4008-4017