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Shiitake Medicinal Mushroom, Lentinus edodes (Higher Basidiomycetes) Productivity and Lignocellulolytic Enzyme Profiles during Wheat Straw and Tree Leaf Bioconversion

Authors:
  • Institute of Microbial Biotechnology Agricultural University of Georgia
  • Agricultural University of Georgia

Abstract and Figures

Two commercial strains of Lentinus edodes have been comparatively evaluated for their productivity and lignocellulolytic enzyme profiles in mushroom cultivation using wheat straw or tree leaves as the growth substrates. Both substrates are profitable for recycling into shiitake fruit bodies. L. edodes 3715 gave the lowest yield of mushroom during tree leaves bioconversion with the biological efficiency (BE) 74.8% while the L. edodes 3721 BE achieved 83.4%. Cultivation of shiitake on wheat straw, especially in the presence of additional nitrogen source, increased the L. edodes 3721 BE to 92-95.3% owing to the high hydrolases activity and favorable conditions. Despite the quantitative variations, each strain of L. edodes had a similar pattern for secreting enzymes into the wheat straw and tree leaves. The mushrooms laccase and MnP activities were high during substrate colonization and declined rapidly during primordia appearance and fruit body development. While oxidase activity decreased, during the same period cellulases and xylanase activity raised sharply. Both cellulase and xylanase activity peaked at the mature fruit body stage. When mushrooms again shifted to the vegetative growth, oxidase activity gradually increased, whereas the hydrolases activity dropped rapidly. The MnP, CMCase, and FP activities of L. edodes 3721 during cultivation on wheat straw were higher than those during mushroom growth on tree leaves whereas the laccase activity was rather higher in fermentation of tree leaves. Enrichment of wheat straw with an additional nitrogen source rather favored to laccase, MnP, and FPA secretion during the vegetative stage of the L. edodes 3721 growth.
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International Journal of Medicinal Mushrooms, 17(1): 77–86 (2015)
77
1045-4403/15/$35.00 © 2015 Begell House, Inc. www.begellhouse.com
Shiitake Medicinal Mushroom, Lentinus edodes
(Higher Basidiomycetes) Productivity and
Lignocellulolytic Enzyme Proles during Wheat Straw
and Tree Leaf Bioconversion
Vladimir Elisashvili,* Eva Kachlishvili, & Mikheil Asatiani
Agricultural University of Georgia, University Campus at Digomi, Tbilisi, Georgia
*Address all correspondence to: Vladimir Elisashvili, Agricultural University of Georgia, University Campus at Digomi, 13 km David
Agmashenebeli Alley, 0159 Tbilisi, Georgia; v.elisashvili@agruni.edu.ge
ABSTRACT: Two commercial strains of Lentinus edodes have been comparatively evaluated for their productivity
and lignocellulolytic enzyme proles in mushroom cultivation using wheat straw or tree leaves as the growth
substrates. Both substrates are protable for recycling into shiitake fruit bodies. L. edodes 3715 gave the lowest yield
of mushroom during tree leaves bioconversion with the biological efciency (BE) 74.8% while the L. edodes 3721
BE achieved 83.4%. Cultivation of shiitake on wheat straw, especially in the presence of additional nitrogen source,
increased the L. edodes 3721 BE to 92-95.3% owing to the high hydrolases activity and favorable conditions. Despite
the quantitative variations, each strain of L. edodes had a similar pattern for secreting enzymes into the wheat straw
and tree leaves. The mushrooms laccase and MnP activities were high during substrate colonization and declined
rapidly during primordia appearance and fruit body development. While oxidase activity decreased, during the same
period cellulases and xylanase activity raised sharply. Both cellulase and xylanase activity peaked at the mature fruit
body stage. When mushrooms again shifted to the vegetative growth, oxidase activity gradually increased, whereas
the hydrolases activity dropped rapidly. The MnP, CMCase, and FP activities of L. edodes 3721 during cultivation on
wheat straw were higher than those during mushroom growth on tree leaves whereas the laccase activity was rather
higher in fermentation of tree leaves. Enrichment of wheat straw with an additional nitrogen source rather favored to
laccase, MnP, and FPA secretion during the vegetative stage of the L. edodes 3721 growth.
KEY WORDS: medicinal mushrooms, Lentinus edodes, fruit body, biological efciency, wheat straw, tree leaves,
lignocellulolytic enzymes
ABBREVIATIONS: ABTS, 2,2′-azino-bis-[3-ethyltiazoline-6-sulfonate]; BE, biological efciency; CMCase, car-
boxymethyl cellulase; FB, mature fruit bodies stage; FPA , lter paper activity; MEA, malt extract agar; MnP, man-
ganese-dependent peroxidase; P, primordia formation stage; PH, postharvest stage; SC, substrate colonization stage
I. INTRODUCTION
Mushrooms have recently received increasing atten-
tion from researchers in food and pharmaceuticals.
Many species have long been used in traditional
medicines or functional foods in China, Japan, and
other Asian countries. Nowadays there is an increas-
ing public interest in the secondary metabolites from
mushrooms for discovering new drugs or lead com-
pounds. A number of bioactive constituents have
been isolated from mushrooms, including small
molecule compounds, polysaccharides, proteins,
and their complexes with antioxidant, antitumor,
antiviral, antimicrobial, and immunomodulatory
agents.1–3
Shiitake medicinal mushroom, Lentinus edodes
(Berk.) Singer (Marasmiaceae, higher Basidiomy-
cetes) is a xylotropic species. It is one of the most
precious edible species, which can be used as both
food and medicine. The L. edodes mushroom is
gradually attracting attention for its high nutrient
value and health-improving functions. The shiitake
mushroom has served as a model for investigating
functional fungi properties and isolating pure com-
pounds for pharmaceutical use. It is a rich source of
proteins, carbohydrates, fiber, vitamins, and miner-
International Journal of Medicinal Mushrooms
Elisashvili, Kachlishvili, & Asatiani
78
als. L. edodes is revered in medicine for its health-
promoting effects, including antiviral, antifungal,
antioxidant, and antitumor effects that boost the
immune system; it lowers cholesterol, works as an
anticoagulant, and is helpful in cancer treatment.4,5
This mushroom is cultivated on both natu-
ral and articial logs since this white rot fungus
is able to colonize different types of agricultural
wastes as growth substrates, although exploitation
of the substrate varies with the species, strain, and
cultivation technology. In particular, the use of
sawdust-based cultivation to replace natural logs
has contributed to expanding the production and
consumption of L. edodes. The main advantages of
this method are the short time to complete a crop
cycle and the higher yields.6 In recent years, wheat
straw and other agricultural wastes have been ex-
amined instead of sawdust for mushroom cultiva-
tion.7–10 However, in spite of the commercial sig-
nicance of L. edodes, little is known concerning
the mushroom performance on different substrates,
especially regarding mushroom yield and quality.
In particular, very little attention has been given
to the evaluation of the lignocellulolytic enzymes
activity participating in deconstruction of plant
substrates. Available data indicate that L. edodes
produces an array of hydrolases and oxidases for
bioconversion of lignocellulosic materials and the
expression of these enzymes activity is related to
and dependent on medium composition, especially
on the growth substrate.8,9,11–14 However, compared
with other cultivated mushrooms, very little is
known about the nature and prole of the ligno-
cellulolytic enzymes secreted by L. edodes during
substrate colonization and fruit body development.
Few studies indicate that gene expression of these
enzymes is strongly regulated during fruit body
development.9,14 Apart from the fundamental sig-
nicance, the understanding of the physiological
mechanisms regulating enzyme synthesis in ligno-
cellulose bioconversion and mushroom morpho-
genesis is important to predict fruiting cycles, to
increase efciency of substrate utilization, and to
contribute to commercial mushroom production
for use as food and medicine. Therefore, the aim of
the present study was to determine the relationship
between the proles of lignocellulolytic enzymes
production and the life cycle of two commercial
strains of L. edodes. Moreover, the impact of two
growth substrates (wheat straw and tree leaves) on
the mushroom yield and extracellular enzyme ac-
tivity was also evaluated in this work.
II. MATERIALS AND METHODS
A. Organisms and Spawn Preparation
The mother cultures of L. edodes (strains 3715
and 3721) were purchased from the Mycelia Com-
pany (Gent, Belgium) and maintained on malt ex-
tract agar (MEA) at 4oC. To prepare spawn, wheat
grains cooked in tap water and mixed with 20 g
CaCO3/kg were lled in 750-mL asks and steril-
ized at 121°C for 1 h. After cooling, the asks were
inoculated with small pieces of mycelial agar and
cultivated at 25°C in the dark.
B. Materials and Cultivation Conditions
Wheat straw (from the Kakheti region) shredded
into pieces of 1–5 cm in length and leaves of Fagus
orientalis (collected in the Sabaduri forest) were
used as the mushroom growth substrates. Both ma-
terials were soaked in tap water or tap water–based
synthetic medium containing 3 g/L (NH4)2SO4 and
5 g/L yeast extract for 16 h at room temperature.
After leaching, 1 kg of the substrate was placed
in polypropylene gas-permeable bags Microsac
PPB75/BEU6/X33-57 (SACO2, EKE, Belgium)
for sterilization by autoclaving at 121°C for 1 h.
After cooling, the bags were inoculated with 10%
(wet weight) of evenly distributed spawn. Inocu-
lated bags (four replicates for each strain/sub-
strate) were incubated in the dark at 24°C. After
2 mo of incubation, the bags were exposed during
3 d at 4°C to experience a cold shock necessary for
the stimulation of fructication. Then the blocks
were kept in the fruiting room at 15°C, with rela-
tive humidity around 90%, and under illumination
of about 400 lux. When the rst primordia signs
appeared, the bags were removed from the blocks.
After the second ush, blocks were soaked in tap
Volume 17, Number 1, 2015
Lentinus edodes Productivity and Lignocellulolytic Enzyme Proles during Wheat Straw and Tree Leaf Bioconversion 79
water (12–15oC) for 18 h and transferred back to
the fruiting room.
During the shiitake cultivation, samples (1–2 g
dry weight) were taken with a spatula from ve
different regions of blocks surface (7–10 mm in
depth) at various stages of the mushroom devel-
opment: substrate colonization stage (SC, 5–7 d
before fruiting), primordial (P) stage, mature fruit
bodies (FB) stage, and 5–7 d after harvesting of
fruit bodies (PH). Distilled water was added to the
fresh material twice (2 × 25 mL) and extracellular
enzymes were extracted by agitation for 3 min. The
solids were separated by centrifugation at 6000 ×
g for 20 min and dried at 60°C; resulting superna-
tants were used to measure pH, sugar content, and
enzyme activity.
C. Biological Efciency
Biological efciency (BE) was estimated as the
ratio of the weight of fresh fruiting bodies and
weight of dry substrate, multiplied by 100.
D. Analytical Methods
The total cellulase activity (lter paper activity
[FPA]) and the carboxymethyl cellulase (CMCase)
were assayed according to International Union of
Pure and Applied Chemistry recommendations by
using, respectively, a strip of lter paper (What-
man no. 1) and CMC low viscosity (1% w/v) in
50 mM citrate buffer (pH 5.0) as the substrates.15
Xylanase activity was determined by incubation of
properly diluted sample birch wood xylan (Roth
7500) (1% w/v) in 50 mM citrate buffer (pH 5.0)
at 50oC for 10 min.16 In all assays, the release of re-
ducing sugars was measured using dinitrosalicylic
acid reagent.17 Glucose and xylose standard curves
were used to calculate the cellulase and xylanase
activities. One unit of cellulase and xylanase activ-
ity was dened as the amount of enzyme, releas-
ing 1 µmol of glucose or xylose, respectively, per
minute.
Laccase activity was determined by monitor-
ing the A420 change related to the rate of oxidation
of 1 mM 2,2′-azino-bis-[3-ethyltiazoline-6-sul-
fonate] (ABTS) in 50 mM Na-acetate buffer (pH
3.7).18 Assays were performed at 20 ± 1oC with
50 µL of adequately diluted culture liquid. MnP ac-
tivity was measured by oxidation of Phenol Red.19
One-milliliter reaction mixtures were incubated
for 1–5 min at 20 ± 1°C in the presence of 0.1 mM
H2O2. The reaction was terminated with 50 µL 4 M
NaOH and absorbance was read at 610 nm. One
unit of laccase and MnP activity was expressed as
the amount of enzyme required to oxidize 1 µmol
of ABTS or Phenol Red in 1 min. Activities in the
absence of H2O2 were subtracted from the values
obtained in the presence of hydrogen peroxide
to establish true peroxidase activity. Activity of
CMCase, xylanase, FPA, laccase, and MnP are
presented as units per gram of dry substrate.
III. RESULTS
A. Productivity
In this study, two strains of L. edodes have been
comparatively evaluated for their productivity in
mushroom cultivation using wheat straw or tree
leaves as the growth substrates. The mushroom
spawn run and the substrate colonization proceed-
ed quickly with formation of the whitish mycelia
on the surface and inside of the blocks. Depending
on the mushroom strain and growth substrate, all
blocks were completely colonized after 24–29 d of
shiitake growth. Primordia and then fruiting bodies
appeared after 66–67 d and 70–72 d of L. edodes
3721 and L. edodes 3715 cultivation, respectively.
That is, earliness (time elapsed between the day
of inoculation and the day of the rst primordia
appearance) was rather dependent on mushroom
strain.
Cultivation of both strains on the same tree
leaves/tap water substrate showed that the mush-
room yield was strain dependent (Table 1). L.
edodes 3721 appeared to be more productive com-
pared with L. edodes 3715, especially during the
rst ush; the mushrooms’ BE achieved 83.4%
and 74.8%, respectively. The comparison of L.
edodes 3721 growth on two different substrates
International Journal of Medicinal Mushrooms
Elisashvili, Kachlishvili, & Asatiani
80
(tree leaves and wheat straw) indicates that the ce-
real substrate ensured higher yield of mushroom
fruiting bodies during the entire period of shiitake
cultivation. Moreover, the BE of L. edodes 3721
grown on the wheat straw was 8.6% higher than
that in tree leaf bioconversion. Finally, the data
obtained prove that soaking of both substrates
by tap water is sufcient to receive a high yield
of shiitake mushroom. However, the soaking of
wheat straw by nutrient medium containing ni-
trogen source and yeast extract caused the further
increase of mushroom fruiting bodies yield from
92% to 95.3%.
B. Enzyme Activity during Development of
L. edodes 3715
Evaluation of L. edodes 3715 cellulase and xy-
lanase activity permitted us to establish that three
tested enzyme activities correlated during mush-
room development on tree leaves (Fig. 1). Xyla-
nase activity of mushroom was approximately 2–3
times higher than that of endoglucanase. During
substrate colonization (stage SC), L. edodes 3715
secreted comparatively low CMCase, xylanase,
and FP activities. At the stage of primordia ap-
pearance (stage P), extracellular activity of these
enzymes sharply increased and attained the maxi-
mum level at the period of mature fruiting body
formation (stage FB). After renewing of vegeta-
tive phase (stage PH), CMCase, FP, and especially
xylanase activity signicantly decreased. No MnP
was detected during the shiitake development on
tree leaves as the growth substrate. The proles of
laccase activity signicantly contrasted with that
of hydrolases. The data presented in Fig. 1 show
that at the end of substrate colonization (stage SC),
laccase activity reached 9.9 U/g substrate. At the
moment of primordia appearance (stage P), laccase
activity was equal to 8.3 U/g and sharply decreased
to 0.7 U/g during fruiting body maturation. Six
days after the harvest, when the vegetative phase
(stage PH) was renewed, the laccase activity again
increased and reached 8.6 U/g substrate.
C. Enzyme Activity during Development
of L. edodes 3721
When L. edodes 3721 was cultivated on tree leaves
as the shiitake growth substrate, the same regulari-
ties were revealed in hydrolase activities measure-
ment (Fig. 2). The transition of culture from the
vegetative phase of development to primordia for-
mation and then to fruit bodies maturation was ac-
companied by a signicant increase in endogluca-
nase, xylanase, and FP activity. During this period,
the CMCase and xylanase activities of mushroom
culture increased more than 6- and 3-fold, respec-
tively. The total cellulase activity of L. edodes
3721 increased with a lesser extent (approximately
2-fold). At the same time, an almost 10-fold de-
crease of L. edodes 3721 laccase activity was ob-
served during the same period of culture develop-
ment. In contrast with L. edodes 3715, mushroom
strain 3721 was capable of secreting MnP in tree
leaf bioconversion, accumulating 5.3 U/g at the
end of substrate colonization. By the primordia ap-
pearance time, the enzyme activity decreased and
completely disappeared at the FB stage. When the
culture again was in the vegetative phase of de-
TABLE 1: L. edodes Fruiting Bodies Yield and BE Tree Leaves and Wheat Straw
Bioconversion
Mushroom strain,
substrate/moistener
Fruiting bodies yield, g
BE, %Flush I Flush II Flush III Total
L. edodes 3715, leaves/water 435 210 103 748 74.8
L. edodes 3721, leaves/water 501 225 108 834 83.4
L. edodes 3721, straw/water 564 236 120 920 92.0
L. edodes 3721, straw/medium 550 257 146 953 95.3
Volume 17, Number 1, 2015
Lentinus edodes Productivity and Lignocellulolytic Enzyme Proles during Wheat Straw and Tree Leaf Bioconversion 81
FIG. 1: L. edodes 3715 CMCase, xylanase, FPA, MnP, and laccase activities in development on the tree leaves
soaked with tap water
FIG. 2: L. edodes 3721 CMCase, xylanase, FPA, MnP, and laccase activities in development on the tree leaves
soaked with tap water
International Journal of Medicinal Mushrooms
Elisashvili, Kachlishvili, & Asatiani
82
velopment (7 d after harvesting of fruit bodies), L.
edodes 3721 cellulase and xylanase activities de-
creased two times, whereas mushroom laccase and
manganese peroxidase activities sharply increased
up to 13.6 U/g and 8.2 U/g substrate, respectively.
It is worth noting that the activities of all tested
lignocellulolytic enzymes during the PH stage
appeared to be much higher than those in the SC
stage.
The substitution of tree leaves as growth sub-
strate for L. edodes 3721 cultivation by wheat
straw soaked in tap water did not change the pro-
les of hydrolase and oxidase activities accumula-
tion. However, the data represented in Fig. 3 show
that the shiitake cultivation on wheat straw favored
endoglucanase, xylanase, and FP activities secre-
tion, especially during substrate colonization and
primordia formation phases. Concerning the lig-
ninolytic enzymes, L. edodes 3721 MnP activity
in mushroom cultivation on straw appeared to be
especially high exceeding the same enzyme activ-
ity during growth on tree leaves more than 5–13
times. It is interesting that the laccase activity in
mushroom growth on wheat straw did not decrease
after termination of vegetative growth (i.e., during
primordia formation). Nevertheless, at the stage of
matured fruiting bodies, laccase activity sharply
decreased; after 7 d after fruit bodies harvesting,
it again increased like in other mushroom cultures.
The study of L. edodes 3721 enzymatic ac-
tivity proles in bioconversion of wheat straw
soaked in tap water supplemented with ammonium
sulphate and yeast extract showed practically the
same regularities as in development on substrate
without supplements. As seen in Fig. 4, laccase
and MnP activities reached a peak during the colo-
nization phase when the substrate was fully colo-
nized. Enzyme levels decreased sharply through-
out the primordia developmental stages, reaching
the lowest levels during the fruit body maturation.
At the same time, the shift from vegetative growth
to fruit body development was related to the in-
crease of polysaccharide-hydrolyzing enzyme
secretion. Comparison of the mushroom culture
behavior during development on the enriched and
nonsupplemented substrates shows that during the
vegetative stage of the mushrooms growth on the
former substrate, the activities of laccase, MnP,
FIG. 3: L. edodes 3721 CMCase, xylanase, FPA, MnP, and laccase activities in development on the wheat straw
soaked with tap water
Volume 17, Number 1, 2015
Lentinus edodes Productivity and Lignocellulolytic Enzyme Proles during Wheat Straw and Tree Leaf Bioconversion 83
and FP were evidently higher compared with those
on the last substrate. High hydrolase activities se-
cretion, especially the total cellulase activity, was
observed during the subsequent stages of shiitake
cultivation on the substrate enriched with nitrogen
and vitamins. It is worth also noting that in contrast
with previous cultures, MnP did not disappear dur-
ing the FB stage in cultivation of L. edodes 3721
on the enriched substrate.
IV. DISCUSSION
In this study, the BE and the proles of lignocel-
lulolytic enzyme accumulation in bioconversion of
tree leaves by two commercial strains of L. edodes
have been evaluated for the rst time. They were
compared with those in the shiitake development
process on wheat straw. As shown in Figs. 1–4,
colonization and the mushroom morphogenesis
process of both substrates are related to the par-
ticular biochemical changes. Signicant uctua-
tions in hydrolytic and oxidative enzyme activities
were observed in various stages of the shiitake life
cycle. Despite the quantitative variations, each
strain of L. edodes had a similar pattern for secret-
ing enzymes into the wheat straw and tree leaves.
Independently of the L. edodes strain and growth
substrate used, the mushroom laccase and MnP
activities were high during substrate colonization
and declined rapidly during primordia appearance
and fruit body development. While oxidase activi-
ty decreased, cellulase and xylanase activity raised
sharply during the same period. Both cellulase and
xylanase activity peaked at the mature fruit body
stage. When mushrooms again shifted to the vege-
tative growth, oxidase activity gradually increased,
whereas the hydrolase activity dropped rapidly.
These results show that changes in the concen-
tration of lignocellulose-deconstructing enzymes
are associated with the fruiting process. Similar
changes in hydrolase and oxidase activity during
the mushrooms’ life cycle were reported for Agari-
cus bisporus,20 Pleurotus strains,21–23 as well as for
L. edodes.8,14 However, during Volvariella volva-
cea development, laccase activity increased after
vegetative growth and stayed high until the fruiting
bodies matured.22 It is interesting to note that in a
study of sawdust-based culture of L. edodes, gene
FIG. 4: L. edodes 3721 CMCase, xylanase, FPA, MnP, and laccase activities in development on the wheat straw
supplemented with nitrogen source
International Journal of Medicinal Mushrooms
Elisashvili, Kachlishvili, & Asatiani
84
expression for laccase and cellulase correlated di-
rectly with the level of enzyme activity (i.e., the
level of laccase transcripts was maximal during the
mycelial growth stage, and then declined rapidly at
the fruiting stage).14 By contrast, the cellulase tran-
script level peaked during fruit body development.
Efcient colonization and utilization of growth
substrate, and consequently the mushroom pro-
ductivity, depend on the capability of mushroom
to synthesis required extracellular hydrolytic and
oxidative enzymes degrading the main polymers
of substrate, cellulose, hemicelluloses, and lignin.
From our results and data available in the litera-
ture, it is reasonable to assume that the high lac-
case and MnP activity during mushroom vegeta-
tive growth provides breakdown of the lignin to
open access of hydrolases to polysaccharides.
The decrease and even inactivation of ligninolytic
activity at the onset of fruiting may be related to
the proteolysis of oxidases and redirection of con-
structive material for carpophore formation. The
sharp increase of hydrolase activity during the P
and FB stages is important to accelerate cellulose
and hemicellulose hydrolysis to steadily supply the
mushroom with new carbon and energy resources.
Mata and Savoie8 clearly showed that the maxima
of carbohydrase activity in L. edodes development
were actually related to a signicant increase in
metabolic activity.
Evaluation of mushroom enzyme activity did
not reveal signicant qualitative differences in lac-
case, CMCase, and xylanase expression in biocon-
version of the same substrate (tree leaves) by two
tested strains. Nevertheless, the FPA of L. edodes
3715 was 2- to 3-fold higher than that of L. edodes
3721 during the entire cultivation period. A ma-
jor distinguishing feature of L. edodes 3715 was
the incapability of this strain to produce MnP in
colonization of tree leaves. Moreover, this study
showed that the level of the tested enzyme activity
depends on the substrate used. The role of the vari-
ous substrates on edible mushroom growth, BE,
and enzyme activity was proven in recent stud-
ies.6–10,13,23 The data presented in Figs. 2 and 3 show
that the levels of MnP, CMCase, and FP activities
of L. edodes 3721 during mushroom cultivation on
wheat straw were higher than those during fungus
growth on tree leaves, whereas the fungus laccase
activity was rather higher in fermentation of tree
leaves. Enrichment of wheat straw with an addi-
tional nitrogen source rather favored laccase, MnP,
and FPA secretion during the vegetative stage of
L. edodes 3721 growth. Moreover, in contrast with
other cultures, the MnP did not disappear during
the FB stage.
Finally, the data obtained indicate that the fruit
bodies yield depends on both the mushroom strain
and the growth substrate. Between two strains used
for tree leaf bioconversion, L. edodes 3715 gave
the lowest yield of mushroom during rst ush
with a BE of 74.8%, whereas the L. edodes 3721
BE achieved 83.4%. Data presented in Table 2
show that the L. edodes 3715 development distin-
guished from that of L. edodes 3721: the growth of
rst strain accompanied with an increase of medi-
um pH from 4.7 during the vegetative stage to 5.7–
5.8 in FB and PH stages, whereas the medium pH
ranged from 4.9 to 4.7 in cultivation of L. edodes
TABLE 2: Substrates pH and Reducing Sugars Content in Cultivation of L. edodes
Strains
Mushroom strain,
substrate/moistener
Medium pH Reducing sugars, mg/g
Stage of development
SC P FB PH SC P FB PH
L. edodes 3715, leaves/water 4.7 5.2 5.7 5.8 5.8 7.7 7.3 6.0
L. edodes 3721, leaves/water 4.9 4.5 4.7 4.6 10.1 10.5 10.3 13.0
L. edodes 3721, straw/water 4.9 4.5 4.5 4.6 15.4 13.5 16.1 14.0
L. edodes 3721, straw/medium 4.8 4.6 4.5 4.6 10.4 13.2 17.4 13.2
Volume 17, Number 1, 2015
Lentinus edodes Productivity and Lignocellulolytic Enzyme Proles during Wheat Straw and Tree Leaf Bioconversion 85
3721, which is optimal for catalytic activity of fun-
gal cellulase and xylanase. It can be speculated that
an elevated pH did not favor rapid hydrolysis of
tree leaf polysaccharides by L. edodes 3715. This
hypothesis is supported by the data on free sugars
content in growth substrate; namely, the reduc-
ing sugars content in this culture appeared to be
almost 2-fold lower compared with that in culti-
vation of L. edodes 3721. As mentioned above,
during mushroom cultivation on wheat straw (lig-
nied substrate with high content of crystalline
cellulose), higher levels of cellulase and xylanase
activities have been observed, especially the pres-
ence of additional nitrogen source. Owing to the
high hydrolase activity at favorable pH, the reduc-
ing sugars content during the FB stage reached
16–17 mg/g substrate (Table 2) and the mushroom
culture was steadily supplied with the carbon and
energy source to produce high yields of fruit bod-
ies with BE of 92%–95.3%. It is interesting that the
substrate formulations of sawdust, wheat bran, and
millet amended with 0.6%–1.2% sucrose, fructose,
or glucose resulted in a shiitake mushroom yield
increase by >11%.25
Thus, tree leaves and wheat straw are bene-
cial growth substrates for recycling into shiitake
fruit bodies by the commercial strains of L. edodes.
The data obtained clearly demonstrate physiologi-
cal and biochemical regularities taking place in
shiitake development on plant raw materials. The
correlation observed between mushroom growth
and enzyme production may be a good parameter
to estimate the period when the mushroom is ready
for fructication. However, further experiments in-
vestigating the impact of different substrates and
some nutrients on mushroom fruit bodies yield and
quality are needed.
ACKNOWLEDGMENTS
This work was supported by the Shota Rustaveli
National Science Foundation of Georgia (grant no.
10/05).
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... The highest laccase activity of P. sajor-caju occurred on the 10th day of mycelial development and in sesame stem-bran mixture (2: 1) as 3.85 U mg -1 , while the lowest value was determined as 0.30 U mg -1 in vine pruning residue. Elisashvili et al. (2015) examined the laccase activities of L. edodes 3715 and 3721 strains in different growing mixtures. It was found that laccase activity increased with mycelia development in both strains, then started to decrease in the primordium stage, then a rapid decrease was observed during mushroom development, finally it increased again at the after-harvest period. ...
... When the cellulase activity of A3 mixture having the highest biological efficiency value was examined, similar results were observed in our study. Elisashvili et al. (2015) reported that cellulase activity of L. edodes decreased with mycelial development in both strains studied, then started to increase in the primordium stage, then a rapid increase was observed during mushroom development, finally decreased at after harvest period. The fastest mycelia development was observed in A7 (3 corncob + 1 bran) and mycelia development ranged between 22 and 50 days. ...
... Sözbir (2014) found that while the fastest mycelia development was obtained with 50 days in 3CK (walnut shell) + M (oak) and CK (walnut shell) + M (oak) mixtures, the slowest mycelia development was detected in T + 3M (oak sawdust) mixture with 95.5 days in L. edodes. Elisashvili et al. (2015) reported that the completion of mycelia development varies between 24 and 29 days depending on the mushroom strain and growing mixtures. Atila (2019) emphasized that mycelia development in shiitake mushroom ranged between 32.4 and 46.0 days. ...
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In this study, it was aimed to investigate cultivation of Lentinula edodes by using different agricultural wastes (oak sawdust, poplar sawdust, wheat stalk, peanut shell, corncob and vine pruning waste) and to determine the most suitable growing mixture/mixtures. For this purpose, 12 growing mixtures were tested. Within the scope of the experiment, besides measurement of yield and quality parameters of mushrooms, properties of agricultural wastes and enzyme activities (laccase and cellulase) of mixtures at different periods were measured. Based on results of the study, the highest and lowest amounts of nitrogen were obtained from after harvest (1.71%) and after sterilization (1.34%) periods, respectively. While the highest amount of carbon was at the after-sterilization period (46.6%), the lowest amount was recorded at the after harvest (45.64%) period. The fastest and slowest mycelia development time was observed in A7 (21.67 days) and A4 (50 days) mixtures, respectively. While the highest yield was determined in A5 (299.59 g kg-1) mixture, A9 (55.99 g kg-1), A6 (65.59 g kg-1) and A11 (75.47 g kg-1) gave the lowest results. While the highest biological activity rate was recorded in A3 (93.65 %) and A5 (92.90%), the lowest was observed in A11 (21.45%), A6 (19.85%) and A9 (19.22%) mixtures. The highest and lowest protein amounts were determined in the A5, A7 and A10, A9 and C mixtures, respectively. The highest cellulase and laccase activities were found in A3 (3.16 IU g-1) and A7 (2164.48 U g-1), respectively.
... The shiitake mushroom, Lentinula edodes, is the most cultivated mushroom in the world, contributing about 22% of the world's mushrooms production [6]. It can be used as food and as medicine and has served as a model for investigating functional properties of fungi and for isolating pure compounds for pharmaceutical use [7]. L. edodes is a rich source of carbohydrates and proteins since it contains 18 types of free amino acids which provide ideal ratios of all the essential amino acids needed for human nutrition [8]. ...
... L. edodes is a rich source of carbohydrates and proteins since it contains 18 types of free amino acids which provide ideal ratios of all the essential amino acids needed for human nutrition [8]. Quality and content of proximate composition and physiologically active substances vary from strain to strain [9,10], and also depend on substrate, supplementation [7,11,12], culture and growth conditions [13], or postharvest processes of fruiting bodies [14,15]. However, there is no detailed information about how some steps of the cultivation process affect the proximate composition. ...
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Lentinula edodes is the most consumed mushroom in the world, being cultivated mainly on waste coming from the forest industry. During shiitake cultivation, incubation is essential and is usually longer than in other edible mushrooms. This stage includes “browning”, which is a process induced by exposure to a photoperiod and although it is believed to bring certain advantages to shiitake cultivation, it has not been widely studied. In this work, we evaluated how the incubation time, the use of different strains and the induction of browning affect the proximate composition, biological efficiency and other yield parameters of L. edodes. In order to do that, three experiments were carried out with three different strains, three different incubation times and the effect of induction or non-induction of browning. As results we found that the proximate composition did not vary with respect to the different incubation times or from the induction of browning, although differences were found according to the strain. On the other hand, the biological efficiency was affected by the incubation time and the strain used, but not by the induction of browning.
... Various types of wastes from agro-industrial process have also been utilized in shiitake growth, including crushed maize cobs, sunflower seed hulls, coffee and hazelnut husks, wheat straw, either alone or in combination with other by-products [13,54,55]. ...
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Lentinula edodes (shiitake mushroom) is the medicinal macro-fungus popular in Japan and China. It is the second most well-known edible fungus in the global market after Agaricus bisporus Because of presence of dietary fibers, vitamins, proteins, minerals and carbohydrates, shiitake mushroom have significant nutritional worth. It is commonly cultivated in winter season and known as "elixir of life". Bioactive compounds such as polysaccharides, amino acid derivatives, sterols, glycoproteins, chitin and sulfurous compounds are all extracted from shiitake mushrooms. Some of these are lentinans, eritadenine and KS-2-α-mannan peptide etc. Shiitake mushroom is used therapeutically to cure cancers, cardiovascular disorders, viral and fungal infections, hepatitis, hypertension and hepatic disorders. Shiitake mushroom have also grown in popularity as a result of their unusual and well-liked flavor.
... Shiitake mushrooms yield better in substrates containing moderate amounts of N, hemicellulose and lignin and with a low cellulose: lignin ratio (Atila 2019a). Biological efficiency differs considerably by ranging from 2.8 to 124.1% in sawdust-based substrates (Diehle and Royse 1996;Pire et al. 2001;Atila 2019b), up to 99.3% in straw-based substrates (Gaitan-Hernandez and Mata 2004;Philippoussis et al. 2007;Elisashvili et al. 2015) or from 102 to 112% in sunflowers hulls (Curvetto et al. 2002) (Table 13.2). ...
Chapter
Shiitake (Lentinula edodes) is the third most commonly cultivated edible mushroom species in the world. It has attracted people’s attention with its medical properties as well as taste and nutritional value. Shiitake which has been known and used in Chinese medicine for more than 2000 years is now considered a great resource for modern clinical and pharmacological research. This mushroom contains many biologically active compounds (polysaccharides, lentinan, LEM and KS–2, ergosterol, nucleic acid derivatives, water-soluble lignins, eritadenine, etc.) which possess different medicinal effects such as antitumor, immunomodulatory, hypocholesterolemic, antibacterial, antifungal, anti–inflammatory and antioxidant. The chapter presents an overview of the research on the shiitake mushroom including its taxonomy, cultivation techniques, biotechnological approach, functional compounds and medicinal properties.
... Some studies have previously evaluated the role of WRF in bioremediation of wastes. Laccases and peroxidases have been reported in the secretomes of several white-rot basidiomycetes, including species of Ganoderma and Lentinula [35,[41][42][43][44][45]. Laccases are commercially important enzymes due to their ability to degrade phenolic and non-phenolic lignin along with recalcitrant pollutants. ...
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Lentinula edodes 3565 and Ganoderma lucidum 9621 were compared for their ability to produce lignocellulolytic enzymes in submerged (SM) and surface liquid (SL) fermentation of hydrolysed colza straw lignin waste that remained after the production of furfural and bioethanol (CS lignin). Application of cultivated mushrooms to dispose of pretreated colza straw agricultural waste is an approach to decrease the quantity of residual lignin while simultaneously obtaining active substances, e.g., the ligninolytic enzyme complex from mycelium. The effect of adding CS lignin to culture media on the yield of L. edodes and G. lucidum mycelium and extracellular laccase activity was studied. It was revealed that the mycelial growth of G. lucidum on solid media was significantly improved by adding CS lignin. Laccase activity during SL cultivation of L. edodes on medium with CS lignin gradually increased over the experiment starting on day 21 and peaked at 520 U/mL on day 28. G. lucidum expressed the maximum laccase activity, 540 U/mL, during the first 14 days of mycelium SM cultivation. Extracellular laccase activity was enhanced about 35- to 40-fold at cultivation of L. edodes and about 10- to 15-fold in the case of G. lucidum by supplementing liquid culture media with CS lignin.
... To summarize their contribution, we can conclude that most common producers are saprophytes isolated primarily from soil, organic residues found in forests and composts, which are classified mainly among bacteria, actinomycetes and filamentous fungi [21,22]. In addition, the white-rot basidiomycetes, or Shiitake medicinal mushroom Lentinus edodes (higher Basidiomycetes) are also known as cellulase producers [23,24]. ...
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Improved cost-effective bioethanol production using inexpensive enzymes preparation was investigated. Three types of waste lignocellulosic materials were converted-for the production of enzyme preparation, a mixture of sugar beet pulp and wheat bran, while the source of sugars in hydrolysates was sweet sorghum biomass. A novel enzyme cocktail of Trichoderma citrinoviride C1 is presented. The one-step ultrafiltration process of crude enzyme extract resulted in a threefold increase of cellulolytic and xylanolytic activities. The effectiveness of enzyme preparation, compared to Cellic ® CTec2, was tested in an optimized enzymatic hydrolysis process. Depending on the test conditions, hydrolysates with different glucose concentrations were obtained-from 6.3 g L −1 to 14.6 g L −1 (representing from 90% to 79% of the CTec2 enzyme yield, respectively). Furthermore, ethanol production by Saccharomyces cerevisiae SIHA Active Yeast 6 strain DF 639 in optimal conditions reached about 120 mL kg d.m. −1 (75% compared with the CTec2 process). The achieved yields suggested that the produced enzyme cocktail C1 could be potentially used to reduce the cost of bioethanol production from sweet sorghum biomass.
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Cultivation of specialty mushrooms on lignocellulosic wastes represents one of the most economical organic recycling processes. Compared with other cultivated mushrooms, very little is known about the nature of the lignocellulolytic enzymes produced by the edible fungus Calocybe indica , its enzymatic activity profiles during submerged and solid state fermentation. The intracellular activity of laccase (7.67 U mg ⁻¹ ), manganese peroxidase (7.48 U mg ⁻¹ ), cellobiohydrolase (5.46 U mg ⁻¹ ), and endoxylanase (4.21 U mg ⁻¹ ) was best obtained in C. indica on 14th and 21st day of incubation. The extracellular activity of laccase (11.57 U mL ⁻¹ ), lignin peroxidase (8.45 U mL ⁻¹ ), and endoxylanases (6.22 U mL ⁻¹ ) were found to be highest on the 14th day. Ligninolytic enzyme activity was substantial during substrate colonisation but quickly dropped during fruiting body development. C. indica , on the other hand, showed relatively modest hydrolase activity during substrate colonisation. The activity of hydrolytic enzymes increased dramatically when primordia formed and peaked at the mature fruiting body stage. The yield of the crude enzyme-treated wheat straw utilised for mushroom production was 52.47%. These findings showed that the activities of lignocellulolytic enzymes were regulated in line with developmental phase of growth of C. indica .
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Shiitake mushroom (Lentinula edodes) can have an important role specially in agriculture where land is limited. This study was the first step to improve mushroom cultivation in Iraq by using locally available lignocellulosic materials as substrates, supplements and types of spawn. Effect of substrates, supplements and spawn on mycelium and browning period, time of three flush, weight of three flush, biological efficiency (BE), total yield, diameter of cap and length of stalk. Used substrates were sawdust and Phragmites australis, supplements were Sesbania sesban, Trifolium and molasses factories residuals. The treatment were T1( sawdust 73%, Sesbania sesban20%), T2(Sawdust73%, Trifolium20%), T3(Phragmites australis vegetative73%, Sesbania sesban20%), T4 (Phragmites australis vegetative73%, Trifolium20%), T5 (Phragmites australis fruiting), T6 (Phragmites australis vegetative73%, Trifolium20%), T6(73%, Sesbania sesban20%) and 2% CaCO 3 , 5% molasses factories residuals for all treatment. Spawn used were wheat grains C 1 , date seeds C 2 , white corn grains C 3 . Results show that faster mycelium covering substrate was at T1 (23.167 days), T6 gave the first complete browning (55.417 days), the days of three flushes were (68.91, 88.42 and 107.83 days) for T5, T6 and T6 respectively. T6 was superior by showing highest values of weight of three flush (73.93, 76.13 and 49.06 g for the 1st, 2nd, and 3rd respectively), total yield and BE (199.14 g and 49.784 % respectively), and diameter of cap and length of stalk (6.7 and 6.0 cm respectively). Hence, T6 which content of Phragmites australis in fruiting stage, Trifolium and molasses was the best substrate used in this study, while C2 (date grains) was best type of spawn.
Chapter
Huge amounts of lignocellulosic wastes which are annually generated worldwide in various sectors (agricultural, forestry, and food industries) are rich in organic compounds and are worthy of being recovered and transformed. The lignocellulosic wastes constitute a major portion of the total carbon that is fixed by photosynthesis. They are the most abundantly available raw materials on the earth and are chiefly composed of carbohydrate polymers like cellulose, hemicelluloses, and lignin. Only a fraction of the total waste is utilized for useful applications, whereas the bulk is left unused, as it may incur disposal cost or it is burnt, which results in the emission of black carbon that causes environmental problems. Therefore, efforts are being made for the conversion of these lignocellulosic wastes into profitable products by using microbial technology. Mushroom cultivation represents an expanded and economically important biotechnological industry in which lignocellulosic waste residues are converted into protein-rich food through solid-state-fermentation process. It is an ecofriendly activity, which in recent years has gained lot of importance due to the increasing global demand for high quality proteins, vitamins, and minerals. Mushrooms are probably the highest protein producers per unit area and time as they have short life cycles and utilize vertical space.Therefore, production of lignocellulosic mushrooms need to be popularized so that bulk of the lignocellulosic wastes can be utilized. The most important lignocellulosic mushrooms include Agaricus bisporus (button mushroom), Pleurotus spp. (oyster mushroom), Calocybe indica (milky mushroom), Ganoderma lucidum (reishi mushroom), Auricularia (wood ear mushroom), Lentinula edodes (shiitake), and many more. This review is based on the growth behavior and productive potential of various commercially and medicinally important lignocellulosic mushrooms on various lignocellulosic substrates.KeywordsLignocellulosic mushroomsAgro wastesForest wastesGrowth behaviorYield
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By seeding fungus on top of industry residues, a mycelium can grow and form a compact network structure; however, it may not develop due to lack of optimal nutrients from the substrate. Consequently, peach palm residues can be a potential alternative; so, to test this hypothesis, this work evaluates the effect of peach palm residues as substrate for the growth of mycelium based on Lentinula edodes . They were also supplemented with cassava bran and various sources of nitrogen – ammonium sulphate, potassium nitrate and soy flour – in order to analyse its effects on its physico-chemical, enzymatic activities and thermal and mechanical properties of the final composite at 12 and 20 days of cultivation. This mycelium was able to grow at optimum source treatment conditions, that depends on the ratio of Carbon to Nitrogen, within only 12 days of inoculation. Furthermore, the enzyme activities directly correlate with the mycelium growth with optimum conditions of pH, water activity and moisture for L. edod es to grow having lower enzyme activities for a well-developed composite; whereas higher activities were seen for a weakly developed material. and this material demonstrate mechanical and thermal properties similar to common mycelium-based composites Therefore, this work demonstrates that peach palm residues can be a potential alternative for mycelium-based composite
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Cotton wastes can be converted without substrate pasteurization into value-added products, such as gourmet mushrooms, with biological efficacy up to 90% and lignocellulolytic enzymes. Edible and medicinal mushroom Pleurotus ostreatus carboxymethyl cellulase and laccase activities, extracted from both the surface and inside layers of blocks, increased during primordia and fruiting bodies development and declined rapidly after the harvest. In a study of xylanase production, it has been observed that the activity of the enzyme in inside layers of the substrate gradually increased during the 49 days (second fruiting stage) after inoculation and then gradually declined towards the end of mushroom cultivation. Fluctuations in xylanase activity in the surface layer were observed, so that maximal enzyme activities coincided with the fruiting stage of P. ostreatus development. In contrast to laccase, manganese peroxidase (MnP) activity was high during the colonization stage and declined during the first primordia and fruiting body formation stages. After this, while laccase activity decreased, MnP activity peaked at the mycelia stage in oyster mushroom development. Data on the levels of cellulase, xylanase, laccase, and MnP activity in spent mushroom substrate demonstrate that it can be used as an excellent source of these enzymes.
Book
Since the publication of the first edition, important developments have emerged in modern mushroom biology and world mushroom production and products. The relationship of mushrooms with human welfare and the environment, medicinal properties of mushrooms, and the global marketing value of mushrooms and their products have all garnered great attention, identifying the need for an updated, authoritative reference. Mushrooms: Cultivation, Nutritional Value, Medicinal Effect, and Environmental Impact, Second Edition presents the latest cultivation and biotechnological advances that contribute to the modernization of mushroom farming and the mushroom industry. It describes the individual steps of the complex mushroom cultivation process, along with comprehensive coverage of mushroom breeding, efficient cultivation practices, nutritional value, medicinal utility, and environmental impact. Maintaining the format, organization, and focus of the previous edition, this thoroughly revised edition includes the most recent research findings and many new references. It features new chapters on medicinal mushrooms and the effects of pests and diseases on mushroom cultivation. There are also updated chapters on specific edible mushrooms, and an expanded chapter on technology and mushrooms. Rather than providing an encyclopedic review, this book emphasizes worldwide trends and developments in mushroom biology from an international perspective. It takes an interdisciplinary approach that will appeal to industrial and medical mycologists, mushroom growers, botanists, plant pathologists, and professionals and scientists in related fields. This book illustrates that mushroom cultivation has and will continue to have a positive global impact on long-term food nutrition, health care, environmental conservation and regeneration, and economic and social change.
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The present review analyzes the history, current status, and future trends in the study of medicinal mushrooms. The target of the present review is to draw attention to many critically important unsolved problems in the future development of medicinal mushroom science in the 21st century. Special attention is paid to mushroom polysaccharides. Many, if not all, higher Basidiomycetes mushrooms contain biologically active polysaccharides in fruit bodies, cultured mycelium, and cultured broth. The data on mushroom polysaccharides are summarized for approximately 700 species of higher Hetero- and Homobasidiomycetes. The chemical structure of polysaccharides and its connection to antitumor activity, including possible ways of chemical modi fication, experimental testing, and clinical use of antitumor or immunostimulating polysaccharides, as well as possible mechanisms of their biological action, are discussed. Particularly, and most importantly for modern medicine, are polysaccharides with antitumor and immunostimulating properties. Several of the mushroom polysaccharide compounds have proceeded through Phase I, II, and III clinical trials and are used extensively and successfully in Asia to treat various cancers and other diseases. A total of 126 medicinal functions are thought to be produced by medicinal mushrooms and fungi, including antitumor, immunomodulating, antioxidant, radical scavenging, cardiovascular, antihypercholesterolemia, antiviral, antibacterial, antiparasitic, antifungal, detoxification, hepatoprotective, and antidiabetic effects.
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In the presence of substrates such as Remazol Blue and 2,2'-azinobis(3-ethylbenzthiazoline-6-sulphonate) (ABTS), laccases Coriolus (Trametes)versicolor can also oxidize non-phenolic lignin model compounds. Veratryl alcohol (I) and 1-(3,4-dimethoxyphenyl)-2-(2-methoxyphenoxy)-propane-1, 3-diol (III) were oxidized by laccase and mediator to give the α-carbonyl derivatives. The β-1 lignin model dimer, 1-(3,4-dimethoxyphenyl)-2-phenoxy-ethane-1, 2-diol(II) was cleaved by laccase in the presence of ABTS to give veratraldehyde and benzaldehyde. On the basis of these observations, we propose that laccase is capable of oxidizing both phenolic and non-phenolic moieties of lignin but that the latter is dependent on the co-presence of primary laccase substrates.