Bioresource Technology 96 (2005) 1439–1444

Comparative study of mycelial growth and basidiomata formation

in seven different species of the edible mushroom genus Hericium
Han Gyu Ko a, Hyuk Gu Park a, Sang Ho Park a, Chang Won Choi b,
Seong Hwan Kim c, Won Mok Park a,*
a
School of Life Sciences and Biotechnology, Korea University, Seoul 136-701, Korea
b
Department of Biology and Medicinal Science, Pai Chai University, Daejeon 302-735, Korea
c
Department of Microbiology and Institute of Basic Science, Dankook University, Cheonan, Chungnam 330-714, Korea

Received 13 July 2004; received in revised form 5 December 2004; accepted 9 December 2004
Available online 16 February 2005

Abstract

The potential of using several agricultural by-products as supplements of sawdust substrate for the production of edible mush-
room Hericium was evaluated using seven Hericium species. All the tested supplements (rice bran, wheat bran, barley bran, Chinese
cabbage, egg shell, and soybean powder) were found to be suitable for the mycelial growth of all the tested species. In mycelial
growth, soybean powder was the best supplement for Hericium americanum, Hericium coralloides, and Hericium erinaceum while
barley bran was the best for Hericium alpestre, Hericium laciniatum, and Hericium erinaceus. For Hericium abietis, rice bran and
Chinese cabbage was the best. The possibility of mushroom production on oak sawdust substrate with 20% rice bran supplement
was demonstrated with H. coralloides, H. americanum, H. erinaceus, and H. erinaceum which showed 26–70% biological efficiency.
Our results also showed that strain selection is important to improve biological efficiency and mushroom yield in Hericium
cultivation.

2005 Elsevier Ltd. All rights reserved.

Keywords: Agricultural by-product; Biological efficiency; Basidioma formation; Hericium spp.; Mushroom cultivation; Mycelial growth

1. Introduction mushroom is well known as roe deerÕs hip (norukung-

Hericium is white and fleshy fungus growing mostly
on dead or dying wood. Its name is a fungal genus
name, and taxonomically it belongs to Hericiaceae,
Hericiales, Homobasidiomycetes, Hymenomycetes, and
Basidiomycote. All the species belonging to the genus
Hericium produce basidiomata as a mass of fragile icicle-
like spines that are suspended from either a branched
supporting framework or from a tough, unbranched
cushion of tissue, and their basidiomata are considered
edible and delicious mushroom (Arora, 1986). Thus, this
*
Corresponding author. Tel.: +82 2 3290 3412; fax: +82 2 923 9923.
E-mail address: mushroom@korea.ac.kr (W.M. Park).
daei) in Korea, and elsewhere is referred to as bearÕs
head, monkeyÕs head, and lionÕs mane mushroom etc.,
in relation to its shape (Chang and Miles, 1989). This
mushroom has also been used in the Orient as an edible
and popular folk medicine to treat various human dis-
eases, and thus it has attracted considerable attention
for its various bioactive substances (Yang and Jong,
1989; Chang and Miles, 1989; Ahn, 1992; Mizuno,
1995; Wasser and Weis, 1999).
Studies on Hericium erinaceus have been shown that
the compounds isolated from its fruiting bodies contain
interesting biological activities such as cytotoxic
effectors on cancer cell (HeLa cells), stimulators of the
synthesis of a nerve growth factor, and nematicidal
and antimicrobial activities (Mizuno et al., 1992;

0960-8524/$ - see front matter
2005 Elsevier Ltd. All rights reserved.
doi:10.1016/j.biortech.2004.12.009
1440 H.G. Ko et al. / Bioresource Technology 96 (2005) 1439–1444

Kawagishi et al., 1994, 1996; Stadler et al., 1994). More- Table 1

over, the basidiomata of H. erinaceus, its liquid-culture The list of Hericium spp. used in this study
broth, or an alcohol-precipitated fraction of its culture No. Hericium spp. Origin Culture codea
filtrate show antitumor activities (Kim et al., 2000; Park 1 H. abietis Canada CBS 243.48
et al., 2002). 2 H. alpestre Austria CBS 539.90
Owing to the rapid increase of our interest on this 3 H. americanum USA CBS 493.63
4 H. coralloides Austria IMSNU 31007
specialty mushroom, more information about Hericium 5 H. erinaceus France CBS 485.95
is necessary to screen efficient species and strains, and to 6 H. laciniatum Canada ATCC 52480
improve mushroom yield and quality. However, limited 7 H. erinaceum 1008 Korea KUMC 1008
data and reference texts are available on the physiolog- 8 H. erinaceum 1101 Malaysia NFCF 1101
ical, genetic, and cultural characteristics of Hericium 9 H. erinaceum 48006 Korea NIAST 48006
a
species. So far several Hericium species have been CBS: Centraalbureau voor Schimmelcultures, Utrecht, The Nether-
known throughout the world. Recently, Park et al. lands. IMSNU: Institute of Microbiology Seoul National University.
ATCC: American Type Culture Collection, USA. KUMC: Korea
(2004) analyzed phylogenetic relationships of seven University Mycology Collection. NFCF: National Forestry Coopera-
Hericium species including Hericium abietis, Hericium tives Federation. NIAST: National Institute of Agricultural Science
alpestre, Hericium americanum, Hericium coralloides, and Technology.
Hericium erinaceum, H. erinaceus and Hericium lacinia-
tum. Based on the rDNA internal transcribed spacer
sequences (ITS), they could separate H. erinaceum from periphery of the growing colony of Hericium and used
other six Hericium species, and showed that H. erina- to inoculate media.
ceus, H. abietis, H. americanum, and H. coralloides are
closely related each other but diverged from H. alpestre 2.2. Mycelial growth on PDA and sawdust substrate with
and H. laciniatum. The phylogenetic analysis of Heri- agricultural waste supplements
cium species to other mushroom species showed that
they are clearly different from other mushroom species To test the effect of temperature on mycelial growth,
(Lu et al., 2002). Among the seven Hericium species, Hericium strains were inoculated on PDA plate and
so far only H. erinaceum (Suzuki and Mizuno, 1997), their growth was assessed at the temperature range of
H. erinaceus (Chang and Roh, 1999) and H. abietis 10–35 C with a 5 C graduation in the dark. Colony
(Xiao and Chapman, 1997) have been attempted to be diameters on each plate were measured 15 days after
cultivated on logs, stumps and sawdust. However, trials incubation. Five replicates were prepared for each
of developing commercial cultivation methods of Heri- strain.
cium have been focused on H. erinaceus and H. erina- It has been known that Hericium species have a high
ceum, and there has been no report on the evaluation saprotrophic ability and grow on a variety of woody
of the use of agricultural by-products for Hericium substances (Xiao and Chapman, 1997). Recently, Chang
cultivation. and Roh (1999) reported the use of different wood spe-
Therefore, this study was carried out to test the cies to test their potential for H. erinaceus production. In
potentiality of using agricultural by-products as a sup- their study, oak was found to be the best for the mycelial
plement resource of sawdust substrate for the growth growth and density of H. erinaceus. Thus, in the present
of seven Hericium species originated from various coun- study we used oak sawdust (Quercus sp.) as a basal sub-
tries. In addition, to diversify the use of Hericium species strate to test the ability of different Hericium species for
for mushroom production, the ability of basidiomata using various agricultural by-products as supplements.
formation and production on oak sawdust substrate To test the effect of agricultural by-products supplement
was accessed with the seven species. to the sawdust substrate on the mycelial growth of Heri-
cium, six agricultural by-products (rice bran, barley
bran, soybean powder, egg shell, Chinese cabbage and
2. Methods wheat bran) were used. To prepare experimental sub-
strates, the basal substrate was mixed with each supple-
2.1. Organisms and inoculum ment in a ratio of 4:1 (w/w) and its moisture content was
adjusted to 65% with distilled water. After moisture con-
Nine strains of seven Hericium species used in this tent adjustment, pH of the supplemented substrates ran-
study were obtained from various culture collections ged 5.0–6.0. The supplemented substrates were packed
(Table 1). All the fungal species were maintained on a in glass test tubes (200 mm length · 30 mm diameter),
potato dextrose agar (PDA) slant at 4 C. Fungal sterilized at 121 C for 40 min, and cooled. An agar disk
inoculum was prepared from mycelia grown on PDA (5 mm in diameter) of all the strains of Hericium listed at
in 90 mm Petri dishes for 14 days at 24 C in the dark. Table 1 was inoculated onto the top of the substrates in
Agar plugs of 4 mm diameter were taken from the glass test tubes. The inoculated tubes were incubated at
 H.G. Ko et al. / Bioresource Technology 96 (2005) 1439–1444 1441

Table 2
Measurements of mycelial growth of Hericium spp. grown at 25 C for 28 days on oak sawdust substrate supplemented with different agricultural
by-products in ratio of 4:1 (w/w)
Hericium spp. Agricultural by-products
Rice bran Barely bran Soybean powder Egg shell Chinese cabbage Wheat bran
1
H. abietis 42 ± 2a 40 ± 1b 28 ± 2d 23 ± 1e 44 ± 1a 32 ± 2c
H. alpestre 50 ± 4c 66 ± 2a 61 ± 2b 41 ± 3d 48 ± 2c 49 ± 2c
H. americanum 82 ± 2d 91 ± 4c 104 ± 3a 78 ± 2d 97 ± 3b 89 ± 2c
H. coralloides 67 ± 3c 63 ± 3d 86 ± 3a 58 ± 3e 75 ± 4b 75 ± 4b
H. erinaceus 79 ± 4b 89 ± 2a 70 ± 3c 58 ± 3d 70 ± 6c 69 ± 4c
H. laciniatum 52 ± 3b 64 ± 3a 63 ± 2a 40 ± 4c 52 ± 7b 60 ± 4a
H. erinaceum 1008 86 ± 2c 95 ± 5b 106 ± 3a 76 ± 6d 98 ± 6b 96 ± 1b
H. erinaceum 1101 88 ± 3d 102 ± 5c 118 ± 3a 80 ± 8e 92 ± 3d 111 ± 4b
H. erinaceum 48006 71 ± 4b 74 ± 3b 96 ± 8a 71 ± 3b 70 ± 4b 69 ± 1b
1
Values are the mean ± SD of mycelial growth measurements (mm), and small alphabet letters indicate the same letters in the same raw are not
statistically significantly different according to the DuncanÕs multiple range test.

25 C for 28 days in the dark and then mycelial growth
was measured from the inoculum to the actively growing
edge by averaging the measurements at two equidistant
points around the circumference of each test tube. The
test was repeated three times with five replicates.
2.3. Basidiomata formation
Since rice bran is the most easily available resource at
mushroom farms in Korea, it was used as the supple-
ment of sawdust to test basidioma formation in different
Hericium species. For this test, oak sawdust (680 g) was
supplemented with rice bran (170 g) and adjusted to
65% moisture, and packed in the bottles of 1100 ml
polypropyrene which is commonly used for other mush-
room cultivation such as Pleurotus and Flammulina. The
substrate-packed bottles were sealed using a polypropyl-
ene collar and a cotton wool plug, sterilized at 121 C
for 90 min, and cooled. Since the amount of substrate
in a 1100 ml polypropyrene bottle was larger than that
in a glass test tube, we increased sterilization time to
90 min. Five bottles were inoculated with each Hericium
strain as described in Section 2.2. The inoculated bottles
were incubated for 20 days at 25 C. Soon after the
hyphae of Hericium reached the bottom of the bottles,
the bottles were moved to a cold room at 16 C with
95% RH and under incandescent light of 1000 Lux to in-
duce primordia formation. The cultivation room was
ventilated four times a day to keep good aeration condi-
tion that gave about 1000 ppm of CO2 concentration in
the air. These environmental conditions for cultivation
were determined as the best conditions in several preli-
minary tests by the authors. The bottles were then
opened to allow the development of basidioma. Ten days
after incubation. first harvest of mushroom was done
from each bottle. For the second and subsequent
flushes, the first flushed bottles were kept at 25 C for
12 days in the dark. If the mycelium in the bottle was
regenerated, the bottles were moved again to the cultiva-
tion room at 16 C and incubated for 10 days to induce
new primordia and basidiomata formation. After har-
vesting the second flushed mushroom, the second

Table 3
Primordia formation and mushroom production of Hericium spp. grown on oak sawdust substrate supplemented with rice bran in ratio of 4:1 (w/w)
Hericium spp. Days taken for primordia appearance Mushroom production1 (g/bottle) Biological efficiency (%)
First flush Second flush Third flush Total production
H. abietis NF2 NF NF NF NF 0
H. alpestre NF NF NF NF NF 0
H. americanum 39 ± 1c3 133 ± 2c 60 ± 2c 42 ± 5b 235 ± 7b 43 ± 1b
H. coralloides 40 ± 1c 90 ± 8e 33 ± 2e 21 ± 15c 143 ± 20e 26 ± 4e
H. erinaceus 36 ± 1b 123 ± 9d 47 ± 5d NF 170 ± 9d 31 ± 2d
H. laciniatum NF NF NF NF NF 0
H. erinaceum 1008 33 ± 1a 156 ± 4a 128 ± 6a 99 ± 9a 383 ± 18a 70 ± 3a
H. erinaceum 1101 37 ± 1b 146 ± 8b 49 ± 11d NF 194 ± 6c 35 ± 1c
H. erinaceum 48006 36 ± 1b 145 ± 5b 97 ± 12b NF 242 ± 8b 44 ± 2b
1
The time elapsed from substrate inoculation until the end of last flush were 92 days for H. americanum, 94 days for H. coralloides, 68 days for
H. erinaceus, 87 days for H. erinaceum 1008, 69 days for H. erinaceum 1101, and 68 days for H. erinaceum 48006.
2
No formation of primordia or basidiomata.
3
Small alphabet letters indicate the same letters in the same column are not statistically significantly different according to the DuncanÕs multiple
range test.
1442 H.G. Ko et al. / Bioresource Technology 96 (2005) 1439–1444
flushed bottles were treated at 25 C for 12 days and
then at 16 C for 10 days for third flush. Mushroom
was harvested by hand and weighed. Both the days ta-
ken for primordia formation after substrate inoculation
and the days taken for basidiomata harvests after pri-
mordia formation were recorded with each bottle of
Hericium strain. The yield of mushroom production
was indicated as fresh weight (g) of harvested basidio-
mata per one bottle. Biological efficiency was defined
as the percentage ratio of the fresh weight of harvested
mushrooms over the dry weight of substrates. The time
elapsed from substrate inoculation until the end of the
last harvest is given at Table 3.
2.4. Data analysis
Data shown in all tables are arithmetic means of the
five replicates measurements. All statistical analysis was
performed using DuncanÕs multiple test at a significance
level of p = 0.05 (Snedecor and Cochran, 1980).
3. Results and discussion
Since there have been no comparative data on the
growth temperature for Hericium cultures used in this
study, their mycelial growth on PDA was compared
by measuring colony diameters before they were evalu-
ated for their ability of utilization of agricultural by-
products as substrate supplements. Fourteen days after
inoculation, H. abietis, H. alpestre, H. coralloides, H.
erinaceus, and H. erinaceum KUMC 1008 and NAIST
48006 showed maximal growth at 25 C whereas H.
americanum, H. laciniatum and H. erinaceum NFCF
1101 showed maximal growth at 30 C (Fig. 1). Among
the tested species H. abietis comparatively slow to grow
80
Hab
70 Ha1
Ham
60
Hco
Her
Mycelial growth (mm)
Hla
Hku
50 Hnf
Hni
40
30
20
10
0
10 15 20 25 30
Temperature (°C)
Fig. 1. Mycelial growth of Hericium spp. grown on PDA at different
temperature for 15 days. Hab: H. abietis, Hal: H. alpestre, Ham: H.
americanum, Hco: H. coralloides, Her: H. erinaceus, Hla: H. laciniatum,
Hku: H. erinaceum KUMC 1008, Hnf: H. erinaceum NFCF 1101, Hni:
H. erinaceum NIAST 48006.
at various temperatures followed by H. laciniatum and
H. alpestre. The fastest growth was observed in H. ameri-
canum. Mycelial growth of all the species sharply
decreased at 35 C. Interestingly, strains of H. erinaceum
showed different mycelial growth at the same tempera-
tures. H. erinaceum NFCF 1101 which was from a trop-
ical area (Malaysia) preferred 30 C for its growth
whereas H. erinaceum KUMC 1008 and NAIST 48006
which were from subtropical areas (Korea) preferred
slightly lowered temperature, 25 C, for their growth.
This result indicates temperature optimum for mycelial
growth differs depending on Hericium strains.
Overall, regardless of different geographical origins of
all tested strains, the results indicated that optimal tem-
perature for Hericium growth occurs within the range of
25–30 C. In case of H. erinaceum and H. erinaceus, our
results of mycelial growth temperature agree with those
of Chang and Miles (1989) and Ko et al. (1997). For the
use of newly isolated Hericium strain, we suggest that
the temperature for optimal growth should be checked
before it is utilized for mushroom cultivation.
Many agricultural by-products and waste materials
have been used to produce edible mushrooms such as
oyster mushroom, golden needle mushroom, and shii-
take. For the production of these mushrooms, sawdust
has been used as one of major substrates. Chang and
Roh (1999) used oak sawdust substrate to produce
mushroom using H. erinaceus. Since oak sawdust is
the most popular basal ingredient of wood materials
for mushroom cultivation in Korea, we also used that
to evaluate the effect of agricultural by-products supple-
ment on Hericium growth. As shown in Table 2, myce-
lial growth was observed from all the species. The
results indicate all the seven Hericium species can grow
on oak substrate with the supplement of any kind of
the tested agricultural by-products.
There were growth differences in the Hericium strains
grown with different agricultural by-products. Soybean
powder was found as the best supplement in the growth
of H. americanum, H. coralloides, and H. erinaceum.
Barley bran was the best for H. alpestre and H. erina-
ceus. For H. abietis, rice bran and Chinese cabbage were
the best supplements. Interestingly, H. laciniatum
showed equally best growth with barley bran, soybean
powder, and wheat bran supplements. No strains of
the tested Hericium favored egg shell as the best supple-
ment. Overall, from the statistical analysis of pairwise
comparisons of all the data, the best growth on oak saw-
dust was noticed in H. erinaceum NFCF 1101 while the
35
least growth was in H. abietis. Considering that H. abie-
tis is originated from softwood such as pines, its slowest
growth is likely due to its inability to better use of nutri-
ents in hardwood such as oak.
When we compared the growth of three H. erinaceum
strains, they showed different preference to the agricul-
tural by-products supplements tested. With soybean
 H.G. Ko et al. / Bioresource Technology 96 (2005) 1439–1444
powder supplement, 1101 strain showed the highest
growth followed by 1008 and 48006 strains, while with
the Chinese cabbage supplement 1008 strain showed
the highest growth. These results indicate that even if
it is the same species, the growth of Hericium strain on
oak sawdust substrate would differ depending on given
supplement. Thus it has to be noticed that when we
select an agricultural by-product for the supplement of
sawdust substrate, it is prerequisite to test several differ-
ent strains of a Hericium species for their ability of uti-
lizing the supplements.
Since an individual mushroom species is different to
fruit on a particular substrate under controlled culture
conditions, we tested the ability of basidiomata forma-
tion in the seven Hericium species listed at Table 1.
Among the seven species, H. erinaceum and H. erinaceus
have been known to form basidiomata on oak sawdust
substrate (Ko, 1998; Chang and Roh, 1999). Thus the
two species were used as control species to produce
mushroom. In case of H. erinaceum, we used three
strains because this species is commonly found in Asian
countries such as China, Korea, and Japan and many
strains of it are available from known fungal culture
collections.
Among the seven species, H. americanum, H. corallo-
ides, H. erinaceus and H. erinaceum could produce
mushrooms on oak sawdust media, but no primordia
and basidiomata formation were observed for H. abietis,
H. alpestre and H. laciniatum (Table 3). Considering the
report of Xiao and Chapman (1997) that H. abietis
could be successfully cultivated on conifer sawdust, the
difference of basal substrate would be one of the reason
for the failure of primordia formation in those three spe-
cies. In the case of H. alpestre and H. laciniatum, further
tests with coniferous substrate would be need to confirm
their potentiality as artificially cultivable species. Addi-
tionally, we also need extensive experimentation with
the two species by differing nutrient sources and growing
conditions because different environmental factors and
cultural practices also play a significant role in primor-
dia formation and fruiting of mushroom species (Sohi
and Upadhyay, 1989; Zervakis et al., 2001).
When it comes to primordia formation, 33–40 days
were taken for primordia appearance in H. erinaceum,
H. erinaceus, H. americanum, and H. coralloides (Table
3). Among these four species primordia appeared little
faster in H. erinaceum and H. erinaceus than in H. ameri-
canum, H. coralloides. All the four species including two
more H. erinaceum strains produced basidiomata. Con-
sequently, 10 days after primordia formation, mush-
room could be harvested (first flush) from the bottles
of all the primordia-formed Hericium. When all the har-
vested bottles were incubated at 25 C for 12 days, sec-
ond primordia appeared. Thus, 10 days after second
primordia formation second mushroom flush could be
successfully done. For the third mushroom flush we
1443
repeated the procedures of 25 C and 16 C treatments
with all the second flushed bottles. But in the third flush
we could harvest mushroom only from H. americanum,
H. coralloides, and H. erinaceum 1008. Generally, it is
known that primordia formation or mushroom pro-
duction is related with environmental (temperature,
humidity, light and aeration), nutritional (carbohydrate,
nitrogen and vitamins) and chemical factors. Since we
gave same environmental and nutritional conditions
for all the tested Hericium species, the difference in the
time taken for primordia appearance and mushroom
production is likely from the properties of strain itself
rather than from other factors. Furthermore, no strong
correlation was present between the average time taken
for primordial appearance and mushroom production.
Therefore, if cultivation conditions or nutritional factors
are unchangeable, efforts for the successful production
of Hericium should be given to the selection of reason-
able strains and/or species.
Regarding total mushroom production, the best
results were obtained from H. erinaceum KUMC 1008
followed by H. erinaceum NIAST 48006 and H. ameri-
canum, which corresponded to biological efficiencies of
70%, 44%, and 43%, respectively (Table 3). Difference
in mushroom production was found among the three
H. erinaceum strains. Similar trends that difference in
mushroom production among different strains of the
same species that grown on identical substrates, were
found in oyster mushroom cultivation (Sohi and
Upadhyay, 1989). The order in the production of Heri-
cium mushroom agreed with that in biological efficiency
(Table 3). It seems in this study that mushroom produc-
tion is correlated with biological efficiency. Considering
that H. erinaceum NIAST 48006 has been used for com-
mercial mushroom production, we expect H. america-
num (which produced similar amount of mushroom
and biological efficiency) could be used for commercial
production. Additionally, although H. coralloides
showed the least production, the species showed its
potential for use in artificial cultivation. Since there
has been no report on the production of H. americanum
and H. coralloides mushrooms on oak sawdust sub-
strate, our work extends the use of these two species
for the production of diverse Hericium mushrooms.
4. Conclusion
In this study, the ability of utilizing agricultural by-
products for the production of Hericium mushroom
was evaluated by measuring the mycelial growth and
basidiomata production of seven Hericium species. All
the tested species showed optimal temperature for myce-
lial growth at 25–30 C. Tests of mycelial growth on oak
sawdust media suggest that all the tested agricultural
by-products are suitable supplements for growing all
1444 H.G. Ko et al. / Bioresource Technology 96 (2005) 1439–1444
the Hericium species tested. With rice bran as an oak
sawdust supplement, we demonstrated for the first time
that artificial cultivation of H. americanum and H. coral-
loides is possible. Further experiments with diverse
strains of these two species need to be performed to
improve their productivity and biological efficiency. In
addition, more works should be done on H. abietis, H.
alpestre, and H. laciniatum to develop suitable methods
for inducing their basidiomata formation on artificial
media composed of agricultural by-products.
Acknowledgements
This study was supported by a grant from the Brain
Korea 21 program of the Ministry of Education in
Korea. We thank National Forestry Cooperatives Fed-
eration (NFCF) and National Institute of Agricultural
Science and Technology (NIAST) for providing the
strains NFCF 1101 and NIAST 48006.
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