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-

daei) in Korea, and elsewhere is referred to as bearÕs
Hericium is white and fleshy fungus growing mostly head, monkeyÕs head, and lionÕs mane mushroom etc.,
on dead or dying wood. Its name is a fungal genus in relation to its shape (Chang and Miles, 1989). This
name, and taxonomically it belongs to Hericiaceae, mushroom has also been used in the Orient as an edible
Hericiales, Homobasidiomycetes, Hymenomycetes, and and popular folk medicine to treat various human dis-
Basidiomycote. All the species belonging to the genus eases, and thus it has attracted considerable attention
Hericium produce basidiomata as a mass of fragile icicle- for its various bioactive substances (Yang and Jong,
like spines that are suspended from either a branched 1989; Chang and Miles, 1989; Ahn, 1992; Mizuno,
supporting framework or from a tough, unbranched 1995; Wasser and Weis, 1999).
cushion of tissue, and their basidiomata are considered Studies on Hericium erinaceus have been shown that
edible and delicious mushroom (Arora, 1986). Thus, this the compounds isolated from its fruiting bodies contain
interesting biological activities such as cytotoxic
effectors on cancer cell (HeLa cells), stimulators of the
*
Corresponding author. Tel.: +82 2 3290 3412; fax: +82 2 923 9923. synthesis of a nerve growth factor, and nematicidal
E-mail address: [email protected] (W.M. Park). 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 90 min. Five bottles were inoculated with each Hericium
was measured from the inoculum to the actively growing strain as described in Section 2.2. The inoculated bottles
edge by averaging the measurements at two equidistant were incubated for 20 days at 25 C. Soon after the
points around the circumference of each test tube. The hyphae of Hericium reached the bottom of the bottles,
test was repeated three times with five replicates. the bottles were moved to a cold room at 16 C with
95% RH and under incandescent light of 1000 Lux to in-
2.3. Basidiomata formation duce primordia formation. The cultivation room was
ventilated four times a day to keep good aeration condi-
Since rice bran is the most easily available resource at tion that gave about 1000 ppm of CO2 concentration in
mushroom farms in Korea, it was used as the supple- the air. These environmental conditions for cultivation
ment of sawdust to test basidioma formation in different were determined as the best conditions in several preli-
Hericium species. For this test, oak sawdust (680 g) was minary tests by the authors. The bottles were then
supplemented with rice bran (170 g) and adjusted to opened to allow the development of basidioma. Ten days
65% moisture, and packed in the bottles of 1100 ml after incubation. first harvest of mushroom was done
polypropyrene which is commonly used for other mush- from each bottle. For the second and subsequent
room cultivation such as Pleurotus and Flammulina. The flushes, the first flushed bottles were kept at 25 C for
substrate-packed bottles were sealed using a polypropyl- 12 days in the dark. If the mycelium in the bottle was
ene collar and a cotton wool plug, sterilized at 121 C regenerated, the bottles were moved again to the cultiva-
for 90 min, and cooled. Since the amount of substrate tion room at 16 C and incubated for 10 days to induce
in a 1100 ml polypropyrene bottle was larger than that new primordia and basidiomata formation. After har-
in a glass test tube, we increased sterilization time to 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 at various temperatures followed by H. laciniatum and
then at 16 C for 10 days for third flush. Mushroom H. alpestre. The fastest growth was observed in H. ameri-
was harvested by hand and weighed. Both the days ta- canum. Mycelial growth of all the species sharply
ken for primordia formation after substrate inoculation decreased at 35 C. Interestingly, strains of H. erinaceum
and the days taken for basidiomata harvests after pri- showed different mycelial growth at the same tempera-
mordia formation were recorded with each bottle of tures. H. erinaceum NFCF 1101 which was from a trop-
Hericium strain. The yield of mushroom production ical area (Malaysia) preferred 30 C for its growth
was indicated as fresh weight (g) of harvested basidio- whereas H. erinaceum KUMC 1008 and NAIST 48006
mata per one bottle. Biological efficiency was defined which were from subtropical areas (Korea) preferred
as the percentage ratio of the fresh weight of harvested slightly lowered temperature, 25 C, for their growth.
mushrooms over the dry weight of substrates. The time This result indicates temperature optimum for mycelial
elapsed from substrate inoculation until the end of the growth differs depending on Hericium strains.
last harvest is given at Table 3. Overall, regardless of different geographical origins of
all tested strains, the results indicated that optimal tem-
2.4. Data analysis perature for Hericium growth occurs within the range of
25–30 C. In case of H. erinaceum and H. erinaceus, our
Data shown in all tables are arithmetic means of the results of mycelial growth temperature agree with those
five replicates measurements. All statistical analysis was of Chang and Miles (1989) and Ko et al. (1997). For the
performed using DuncanÕs multiple test at a significance use of newly isolated Hericium strain, we suggest that
level of p = 0.05 (Snedecor and Cochran, 1980). the temperature for optimal growth should be checked
before it is utilized for mushroom cultivation.
Many agricultural by-products and waste materials
3. Results and discussion have been used to produce edible mushrooms such as
oyster mushroom, golden needle mushroom, and shii-
Since there have been no comparative data on the take. For the production of these mushrooms, sawdust
growth temperature for Hericium cultures used in this has been used as one of major substrates. Chang and
study, their mycelial growth on PDA was compared Roh (1999) used oak sawdust substrate to produce
by measuring colony diameters before they were evalu- mushroom using H. erinaceus. Since oak sawdust is
ated for their ability of utilization of agricultural by- the most popular basal ingredient of wood materials
products as substrate supplements. Fourteen days after for mushroom cultivation in Korea, we also used that
inoculation, H. abietis, H. alpestre, H. coralloides, H. to evaluate the effect of agricultural by-products supple-
erinaceus, and H. erinaceum KUMC 1008 and NAIST ment on Hericium growth. As shown in Table 2, myce-
48006 showed maximal growth at 25 C whereas H. lial growth was observed from all the species. The
americanum, H. laciniatum and H. erinaceum NFCF results indicate all the seven Hericium species can grow
1101 showed maximal growth at 30 C (Fig. 1). Among on oak substrate with the supplement of any kind of
the tested species H. abietis comparatively slow to grow the tested agricultural by-products.
There were growth differences in the Hericium strains
grown with different agricultural by-products. Soybean
80
powder was found as the best supplement in the growth
Hab
70 Ha1 of H. americanum, H. coralloides, and H. erinaceum.
Ham
60
Hco
Her
Barley bran was the best for H. alpestre and H. erina-
Mycelial growth (mm)

Hla
Hku
ceus. For H. abietis, rice bran and Chinese cabbage were
50 Hnf the best supplements. Interestingly, H. laciniatum
Hni
40 showed equally best growth with barley bran, soybean
powder, and wheat bran supplements. No strains of
30
the tested Hericium favored egg shell as the best supple-
20 ment. Overall, from the statistical analysis of pairwise
10
comparisons of all the data, the best growth on oak saw-
dust was noticed in H. erinaceum NFCF 1101 while the
0
10 15 20 25 30 35
least growth was in H. abietis. Considering that H. abie-
Temperature (°C) tis is originated from softwood such as pines, its slowest
growth is likely due to its inability to better use of nutri-
Fig. 1. Mycelial growth of Hericium spp. grown on PDA at different
ents in hardwood such as oak.
temperature for 15 days. Hab: H. abietis, Hal: H. alpestre, Ham: H.
americanum, Hco: H. coralloides, Her: H. erinaceus, Hla: H. laciniatum, When we compared the growth of three H. erinaceum
Hku: H. erinaceum KUMC 1008, Hnf: H. erinaceum NFCF 1101, Hni: strains, they showed different preference to the agricul-
H. erinaceum NIAST 48006. tural by-products supplements tested. With soybean
 H.G. Ko et al. / Bioresource Technology 96 (2005) 1439–1444 1443

powder supplement, 1101 strain showed the highest repeated the procedures of 25 C and 16 C treatments
growth followed by 1008 and 48006 strains, while with with all the second flushed bottles. But in the third flush
the Chinese cabbage supplement 1008 strain showed we could harvest mushroom only from H. americanum,
the highest growth. These results indicate that even if H. coralloides, and H. erinaceum 1008. Generally, it is
it is the same species, the growth of Hericium strain on known that primordia formation or mushroom pro-
oak sawdust substrate would differ depending on given duction is related with environmental (temperature,
supplement. Thus it has to be noticed that when we humidity, light and aeration), nutritional (carbohydrate,
select an agricultural by-product for the supplement of nitrogen and vitamins) and chemical factors. Since we
sawdust substrate, it is prerequisite to test several differ- gave same environmental and nutritional conditions
ent strains of a Hericium species for their ability of uti- for all the tested Hericium species, the difference in the
lizing the supplements. time taken for primordia appearance and mushroom
Since an individual mushroom species is different to production is likely from the properties of strain itself
fruit on a particular substrate under controlled culture rather than from other factors. Furthermore, no strong
conditions, we tested the ability of basidiomata forma- correlation was present between the average time taken
tion in the seven Hericium species listed at Table 1. for primordial appearance and mushroom production.
Among the seven species, H. erinaceum and H. erinaceus Therefore, if cultivation conditions or nutritional factors
have been known to form basidiomata on oak sawdust are unchangeable, efforts for the successful production
substrate (Ko, 1998; Chang and Roh, 1999). Thus the of Hericium should be given to the selection of reason-
two species were used as control species to produce able strains and/or species.
mushroom. In case of H. erinaceum, we used three Regarding total mushroom production, the best
strains because this species is commonly found in Asian results were obtained from H. erinaceum KUMC 1008
countries such as China, Korea, and Japan and many followed by H. erinaceum NIAST 48006 and H. ameri-
strains of it are available from known fungal culture canum, which corresponded to biological efficiencies of
collections. 70%, 44%, and 43%, respectively (Table 3). Difference
Among the seven species, H. americanum, H. corallo- in mushroom production was found among the three
ides, H. erinaceus and H. erinaceum could produce H. erinaceum strains. Similar trends that difference in
mushrooms on oak sawdust media, but no primordia mushroom production among different strains of the
and basidiomata formation were observed for H. abietis, same species that grown on identical substrates, were
H. alpestre and H. laciniatum (Table 3). Considering the found in oyster mushroom cultivation (Sohi and
report of Xiao and Chapman (1997) that H. abietis Upadhyay, 1989). The order in the production of Heri-
could be successfully cultivated on conifer sawdust, the cium mushroom agreed with that in biological efficiency
difference of basal substrate would be one of the reason (Table 3). It seems in this study that mushroom produc-
for the failure of primordia formation in those three spe- tion is correlated with biological efficiency. Considering
cies. In the case of H. alpestre and H. laciniatum, further that H. erinaceum NIAST 48006 has been used for com-
tests with coniferous substrate would be need to confirm mercial mushroom production, we expect H. america-
their potentiality as artificially cultivable species. Addi- num (which produced similar amount of mushroom
tionally, we also need extensive experimentation with and biological efficiency) could be used for commercial
the two species by differing nutrient sources and growing production. Additionally, although H. coralloides
conditions because different environmental factors and showed the least production, the species showed its
cultural practices also play a significant role in primor- potential for use in artificial cultivation. Since there
dia formation and fruiting of mushroom species (Sohi has been no report on the production of H. americanum
and Upadhyay, 1989; Zervakis et al., 2001). and H. coralloides mushrooms on oak sawdust sub-
When it comes to primordia formation, 33–40 days strate, our work extends the use of these two species
were taken for primordia appearance in H. erinaceum, for the production of diverse Hericium mushrooms.
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- 4. Conclusion
canum, H. coralloides. All the four species including two
more H. erinaceum strains produced basidiomata. Con- In this study, the ability of utilizing agricultural by-
sequently, 10 days after primordia formation, mush- products for the production of Hericium mushroom
room could be harvested (first flush) from the bottles was evaluated by measuring the mycelial growth and
of all the primordia-formed Hericium. When all the har- basidiomata production of seven Hericium species. All
vested bottles were incubated at 25 C for 12 days, sec- the tested species showed optimal temperature for myce-
ond primordia appeared. Thus, 10 days after second lial growth at 25–30 C. Tests of mycelial growth on oak
primordia formation second mushroom flush could be sawdust media suggest that all the tested agricultural
successfully done. For the third mushroom flush we 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 Kim, D.M., Pyun, C.W., Ko, H.G., Park, W.M., 2000. Isolation of
sawdust supplement, we demonstrated for the first time antimicrobial substances from Hericium erinaceum. Mycobiology
28, 33–38.
that artificial cultivation of H. americanum and H. coral- Ko, H.G., 1998. Cultural characteristics of Hericium erinaceum. M.S.
loides is possible. Further experiments with diverse thesis, Department of Agricultural Biology, Korea University.
strains of these two species need to be performed to Ko, H.G., Kim, D.M., Park, W.M., 1997. Composition of a new
improve their productivity and biological efficiency. In medium for mycelial growth of Hericium erinaceus. Kor. J. Mycol.
addition, more works should be done on H. abietis, H. 25, 369–376.
Lu, L., Li, J., Cang, Y., 2002. PCR-based sensitive detection of
alpestre, and H. laciniatum to develop suitable methods medicinal fungi Hericium species from ribosomal internal tran-
for inducing their basidiomata formation on artificial scribed spacer (ITS) sequence. Biol. Pharm. Bull. 25, 975–980.
media composed of agricultural by-products. Mizuno, T., 1995. Yamabushitake, Hericium erinaceum: bioactive
substances and medicinal utilization. Food Rev. Int. 11, 173–
178.
Mizuno, T., Wasa, T., Ito, H., Suzuki, C., Ukai, N., 1992. Antitumor-
Acknowledgements active polysaccharides isolated from the fruiting body of Hericium
erinaceum, an edible and medicinal mushroom called yamabush-
This study was supported by a grant from the Brain itake or houtou. Biosci. Biotech. Biochem. 56, 347–348.
Korea 21 program of the Ministry of Education in Park, H.K., Ko, H.G., Kim, S.H., Park, W.M., 2004. Molecular
Korea. We thank National Forestry Cooperatives Fed- identification of Asian isolates of medicinal mushroom Hericium
erinaceum by phylogenetic analysis of nuclear ITS rDNA. J.
eration (NFCF) and National Institute of Agricultural Microbiol. Biotechnol. 14, 816–821.
Science and Technology (NIAST) for providing the Park, Y.S., Lee, H.S., Won, M.H., Lee, J.H., Lee, S.Y., Lee, H.Y.,
strains NFCF 1101 and NIAST 48006. 2002. Effect of an exo-polysaccharide from the culture broth of
Hericium erinaceus on enhancement of growth and differentiation
of rat adrenal nerve cells. Cytotechnology 39, 155–162.
Snedecor, G.W., Cochran, W.G., 1980. Statistical Methods, seventh
References ed. Iowa State University Press, Ames, IO.
Sohi, H.S., Upadhyay, R.C., 1989. Effect of temperature on mycelial
Ahn, D.K., 1992. Medicinal fungi in Korea. Kor. J. Mycol. 20, 154– growth of Pleurotus species and their yield performance on selected
165. substrates. Mush. Sci. 12, 49–56.
Arora, D., 1986. Mushrooms Demystified. Ten Speed Press, Berkeley, Stadler, M., Mayer, A., Anke, H., Sterner, O., 1994. Fatty acids and
Calif. other compounds with nematicidal activity from cultures of
Chang, H.Y., Roh, M.G., 1999. Physiological characteristics of basidiomycetes. Planta Med. 60 (2), 128–132.
Hericium erinaceus in sawdust media. Kor. J. Mycol. 27, 252–255. Suzuki, C., Mizuno, T., 1997. Cultivation of Yamabushitake (Heri-
Chang, S.H., Miles, P.G., 1989. Edible Mushroom and Their cium erinaceum). Food Rev. Int. 13, 419–421.
Cultivation. CRC Press, pp. 120, 307–312. Wasser, S.P., Weis, A.L., 1999. Therapeutic effects of substances
Kawagishi, H., Shimada, A., Shirai, R., Okamoto, K., Ojima, F., occurring in higher basidiomycetes mushroom: a modern perspec-
Sakamoto, H., Ishiguro, Y., Furukawa, S., 1994. Erinacines A, B tive. Crit. Rev. Immunol. 19 (1), 65–96.
and C, strong stimulators of nerve growth factor (NGF)-synthesis, Xiao, G., Chapman, B., 1997. Cultivation of Hericium abietis on
from the mycelia of Hericium erinaceum. Tetrahedron Lett. 35, conifer sawdust. Can. J. Bot. 75, 1155–1157.
1569–1572. Yang, Q.Y., Jong, S.C., 1989. Medicinal mushrooms in China. Mush.
Kawagishi, H., Shimada, A., Hosokawa, S., Mori, H., Sakamoto, H., Sci. 12, 631–643.
Ishiguro, Y., Sakemi, S., Bordner, J., Kojima, N., Furukawa, S., Zervakis, G., Philippoussis, A., Ioannidou, S., Diamantopoulou, P.,
1996. Erinacines E, F and G, stimulators of nerve growth factor 2001. Mycelium growth kinetics and optimal temperature condi-
(NGF)-synthesis, from the mycelia of Hericium erinaceum. Tetra- tions for the cultivation of edible mushroom species on lignocel-
hedron Lett. 37, 7399–7402. lulosic substrates. Folia Microbiol. 46 (3), 231–234.