Biology, cultivation, and medicinal functions of the mushroom Hericium erinaceum

Abstract

Hericium erinaceum (Bull.: Fr.) Pers. is an edible fungus of great significance in medicine. It is rarely found in Europe, in contrast, it is common in Japan and North America. Its fruitbodies have been well-known for hundreds of years in traditional Chinese medicine and cuisine. A cradle of H. erinaceum cultivation is Asia. In Eastern Europe is rare in natural habitats, but can be successfully cultivated. Both fruitbodies and mycelia are rich in active, health promoting substances. Tests of substances extracted from this mushroom carried out on animals and in vitro have given good results. They can be used in the treatment of cancer, hepatic disorders, Alzheimer’s and Parkinson’s diseases, wound healing. They improve cognitive abilities, support the nervous and immune systems. Promising results have been reported in clinical trials and case reports about the human treatment (e.g., recovery from schizophrenia, an improvement of the quality of sleep, alleviation of the menopause symptoms). The subject of this paper is to summarize information about the development of mycelium, the best conditions for cultivation of fruitbodies, bioactive substances and their use in medicine.

Full text

1 of 18Published by Polish Botanical Society
Acta Mycologica
REVIEW
Biology, cultivation, and medicinal functions
of the mushroom Hericium erinaceum
Sławomir Sokół1, Iwona Golak-Siwulska2, Krzysztof Sobieralski2,
Marek Siwulski2, Katarzyna Górka1*
1 Laboratory of Applied Mycology and Plant Systematics, Department of Biosystematics,
University of Opole, Oleska 22, 40-052 Opole, Poland
2 Department of Vegetable Crops, Poznań University of Life Sciences, Dąbrowskiego 159, 60-594
Poznań, Poland
* Corresponding author. Email: tenebrity@gmail.com
Abstract
Hericium erinaceum (Bull.: Fr.) Pers. is an edible fungus of great signicance in
medicine. It is rarely found in Europe, in contrast, it is common in Japan and North
America. Its fruitbodies have been well-known for hundreds of years in traditional
Chinese medicine and cuisine. A cradle of H. erinaceum cultivation is Asia. In
Eastern Europe is rare in natural habitats, but can be successfully cultivated. Both
fruitbodies and mycelia are rich in active, health promoting substances. Tests of
substances extracted from this mushroom carried out on animals and in vitro have
given good results. ey can be used in the treatment of cancer, hepatic disor-
ders, Alzheimer’s and Parkinson’s diseases, wound healing. ey improve cogni-
tive abilities, support the nervous and immune systems. Promising results have
been reported in clinical trials and case reports about the human treatment (e.g.,
recovery from schizophrenia, an improvement of the quality of sleep, alleviation of
the menopause symptoms). e subject of this paper is to summarize information
about the development of mycelium, the best conditions for cultivation of fruit-
bodies, bioactive substances and their use in medicine.
Keywords
Hericium erinaceum; cultivation; medicine; bioactive compounds
Taxonomy, description, and occurrence
Lion’s mane mushroom, also called bearded tooth, Hericium erinaceum (Bull.: Fr.)
Pers. used to belong to the class Basidiomycetes, subclass Holobasidiomycetidae,
order Hericiales, family Hericiaceae [1], while Index Fungorum [2] presents the cur-
rently adopted taxonomy of Hericium erinaceus (Bull.) Pers. (described in 1979 by
Persoon [3]) as follows: Basidiomycota, Agaricomycotina, Agaricomycetes, Incertae
sedis, Russulales, Hericiaceae. Lion’s mane mushroom is predominantly a saprophyte,
but occasionally it may also be a weak parasite of trees [4]. is species is found on
dead or dying deciduous trees belonging to genera Quercus sp., Fagus sp., Acer sp.,
Juglans sp., and Ulmus sp. [5–7]. It forms extensively branched fruitbodies, irregularly
bulbous with a spiny hymenophore. e fruitbody is most frequently attached to the
substrate sideways, with the base rounded or subglobose, protruding and unbranched
[4]. Spores are ellipsoid, smooth to slightly roughened, around 5.5–7 × 4.5–5.5 μm
[5].
Lion’s mane mushroom is a rare species in Poland [4]. Five localities of Hericium
erinaceum ‘erinaceus’ (Bull.: Fr.) Pers. have been reported from Poland by dening
their hosts: Carpinus sp., Fagus sp., and Quercus sp. [8]. All species from the family
DOI: 10.5586/am.1069
Publication history
Received: 2015-08-18
Accepted: 2016-01-08
Published: 2016-01-29
Handling editor
Tomasz Leski, Institute of
Dendrology of the Polish
Academy of Sciences, Poland
Authors’ contributions
SS, IGS: manuscript drafting;
IGS, MS: translation; KS, KG: nal
version of the manuscript; MS:
photos from the research
Funding
The manuscript was nanced
by authors as parts of individual
research grants.
Competing interests
No competing interests have
been declared.
Copyright notice
© The Author(s) 2016. This is an
Open Access article distributed
under the terms of the Creative
Commons Attribution License,
which permits redistribution,
commercial and non-
commercial, provided that the
article is properly cited.
Citation
Sokół S, Golak-Siwulska I,
Sobieralski K, Siwulski M, Górka
K. Biology, cultivation, and
medicinal functions of the
mushroom Hericium erinaceum.
Acta Mycol. 2015;50(2):1069.
http://dx.doi.org/10.5586/
am.1069
Digital signature
This PDF has been certied using digital
signature with a trusted timestamp to
assure its origin and integrity. A verication
trust dialog appears on the PDF document
when it is opened in a compatible PDF
reader. Certicate properties provide
further details such as certication time
and a signing reason in case any alterations
made to the nal content. If the certicate
is missing or invalid it is recommended to
verify the article on the journal website.
Piotr Otręba
Elektronicznie podpisany przez Piotr Otręba
DN: c=PL, o=Polish Botanical Society, ou=Polish Botanical Society, l=Warsaw, cn=Piotr Otręba, email=p.otreba@pbsociety.org.pl
Data: 2016.01.25 16:46:15 Z

2 of 18© The Author(s) 2016 Published by Po lish Botanical Societ y Acta Mycol 50(2):1069
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
Hericiaceae are legally protected. In Poland, next to H. erinaceum, also Hericium cor-
alloides (Scop.: Fr.) Pers, and Hericium agellum (Scop.) Pers. (= H. alpestre Pers.) are
found [8]. In Poland localities of lion’s mane mushroom have been described from
the Iński Landscape Park in Western Pomerania [9], as well as Oliwa Forests near
Gdańsk, Hylaty Valley in the Bieszczady Mountains, and from the Lower Beskids re-
gion [10–12].
In the opinion of many authors lion’s mane mushroom is common practically al-
most throughout the entire Northern Hemisphere, excluding tropical and polar re-
gions [5,13–17]. Hericium erinaceum is rarely found in Europe [18]. It has been placed
on the red list in several European countries; in contrast, it is common in Japan and
North America.
In the late 1970’s a new taxon was described, i.e., Hericium erinaceum subsp. eri-
naceo abietis. is taxon diered from the typical H. erinaceum, e.g., by morphologi-
cal traits of the fruitbody, spore size and mycelium growth rate. Studies showed that
the described subspecies was a sterile hybrid between H. erinaceum and H. abietis
[19].
New possibilities for the classication of taxa of various ranks, e.g., strains and
species of Hericium have been provided by the application of polymerase chain re-
action (PCR) molecular identication. Using PCR Lu and his colleagues conducted
taxonomic identication and determined phylogenetic aliation of representatives of
the family Hericiaceae with other Holobasidiomycetidae [20]. Studies showed that the
family Hericiaceae is very closely related with dierent species of families Auriscalpi-
aecae, Echinadontaceae, Russulaceae, Schizophyllaceae and Stereaceae. e applica-
tion of a molecular method, i.e., PCR, made it possible also to verify taxonomy. us,
Canadian strains, previously known as Hericium erinaceus, were isolated as a new
taxon in the rank of species – Hericium americanum. is species was described by
Ginns in 1984 [21]. A signicant role in the development of simple methods to deter-
mine taxonomic aliation of lion’s mane mushroom was played by a study by Adair’s
team [22]. Molecular PCR-RFLP techniques and their application made it possible to
develop a rapid and simple method to detect the presence in wood of both Hericium
sp. and other wood decomposing fungi already at the initial phase of their develop-
ment [23].
Factors aecting mycelium growth in lion’s mane mushroom
It was shown that growth of monokaryotic mycelia of lion’s mane mushroom is typi-
cally slower than that of dikaryotic cultures. Only approx. 3% monokaryotic cultures
yielded fruitbodies, which were always smaller in comparison to fruitbodies produced
by dikaryotic mycelia. e dikaryotic mycelium of Hericium sp. diered considerably
from the monokaryotic mycelium, e.g., by forming hyphae with clamp connections
[24]. e monokaryotic mycelium formed 4 types of colonies, i.e., (i) thin, semi-
airborne, showing a rapid growth comparable to that of a dikaryotic mycelium; (ii)
compact, slower growing; (iii) thick and robust, characterized by very slow growth,
and (iv) thin, with the slowest growth rate. e monokaryotic mycelium is capable of
forming chlamydospores, spindle-shaped, of 6–8 × 8–10 μm. Chlamydospores remain
viable for over 7 years and tolerate well storage under anaerobic conditions. e time
required for chlamydospore germination ranges from 30 to 52 h, while germinability
ranges from 32 to 54%, respectively [25].
Optimal sporulation conditions of Hericium sp. were dened [26]. Production
of spores under natural conditions was greatest around noon, which was connected
with an increase in temperature and a reduction of relative humidity (RH). In turn,
sporulation under laboratory conditions at 85–95% RH increased with an increase
in temperature to 24–27°C, while it stopped at 31–33°C. It was also found that at a
temperature of 20°C spore production was greater at 30% RH than at 90% RH.
e eect of dierent factors on fungal spore germination was investigated in
Hericium sp. An advantageous eect of growth regulators, e.g., 2,4-D and gibberel-
lin, was observed on germination of lion’s mane mushroom spores [27]. Studies were
also conducted on the acceleration of spore germination using red and green laser

3 of 18© The Author(s) 2016 Published by Po lish Botanical Societ y Acta Mycol 50(2):1069
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
light. e application of low intensity light stimulated spore germination as well as the
vegetative growth of mycelium on dierent substrates [28]. e use of argon and he-
lium lasers in irradiation of mycelia resulted also in the acceleration of fructication,
while it also increased the weight of a single fruitbody and the yield of fruitbodies by
36–51% [29].
Mycelium growth in Hericium sp. depends on dierent factors. In a study by opti-
mal temperature for mycelium growth was 25°C [30]. e best mycelium growth on
substrates was observed at pH 6. Most carbon sources, apart from lactose, promoted
mycelium growth in H. erinaceum. In turn, alanine constituted the best source of
nitrogen, while histidine was the least advantageous source. Investigations showed a
stimulating eect of isoleucine chelate, applied at 100–200 ppm, on mycelium growth
in H. erinaceum [31].
Mycelium growth rate in lion’s mane mushroom in the substrate is closely con-
nected with its enzymatic activity. Hericium erinaceum produces hydrolytic enzymes
causing decomposition of cellulose, lignin, starch and proteins in the substrate. My-
celium growth rate in Hericium is correlated with the activity of β-amylase and pro-
tease [32]. Lion’s mane mushroom is a species producing α-amylase, causing starch
decomposition in the substrate [33]. Amylase isolated from fruitbodies of lion’s mane
mushroom showed optimal activity at pH 4.6 and temperature of 40°C [34]. Hericium
erinaceum is a species with high activity of cellulase and laccase [35,36]. A correlation
was found between the activity of laccase and the length of the development cycle in
this fungus [37]. e higher the activity of released laccase, the shorter the growth
period.
A high enzymatic activity of lion’s mane mushroom facilitates its practical applica-
tions. Lion’s mane mushroom is suitable for bioremediation, e.g., to treat sewage in the
pulp and paper industry [38]. ere is an application of composted spent mushroom
substrate (SMS) from lion’s mane mushroom culture for commercial scale produc-
tion of such enzymes as α-amylase, cellulase, β-glucosidase, laccase and xylanase [39].
SMS may be applied as a readily available and cheap source of enzymes for bioreme-
diation purposes [40].
Optimal temperature for mycelium growth in lion’s mane mushroom is 21–24°C
at substrate moisture content of 50 to 70% [5]. It was found that the optimal tem-
perature for mycelium growth can be dierent and falls within the range of 25–30°C,
while maximum temperature is 35°C [41]. For the fruiting period it is recommended
the maintenance of constant temperature of 23°C [35]. Optimal pH of substrate may
range from 5.8 to 6.2 [42]. Another study [43] showed that optimal temperature for
Lion’s mane mushroom yielding ranges from 10 to 24°C, while pH falls within a rela-
tively broad range from 4 to 6, the recommended humidity is 85–95% and lighting
intensity of 200–400 lx. Ranges of parameters, at which culture should be run, are
close to the conditions found during mycelium growth. ere was one more study
reporting similar ranges of temperature and substrate moisture content as the previ-
ously mentioned authors, optimal pH also falls within a comparable range, as accord-
ing to the authors it should be from 4 to 5 [44].
Cultivation of lion’s mane mushroom
Cultivation of lion’s mane mushroom may be extensive or intensive. e rst one is
applied on a wide scale in China. Wood logs or stumps are spawned with wood frag-
ments overgrown with mycelium of lion’s mane mushroom. Logs aer spawning are
placed in facilities where high humidity is maintained. Spawning takes place under
natural, uncontrolled conditions, thus in this culture system fruitbodies are produced
at various times, i.e., from several months up to a year aer wood spawning. is
method of lion’s mane mushroom growing is very simple and does not require any
considerable investment or specialist equipment. Its primary drawback is connected
with the long period before mushroom yield may be harvested and high labor con-
sumption [5].
Intensive cultivation methods need to be used order to obtain high yields of good
quality. Intensive cultivation of lion’s mane mushroom is typically run in bottles (Fig. 1,

4 of 18© The Author(s) 2016 Published by Po lish Botanical Societ y Acta Mycol 50(2):1069
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
Fig. 2) or bags (Fig. 3–Fig. 5). Substrates for culture need to be sterilized, thus bottles
have to be made from heat resistant materials such as, e.g., polypropylene. In order to
facilitate respiration of mycelium during spawning, bottles and bags used in culture
have to be equipped with a lter, which will ensure gas exchange, while at the same
time preventing penetration of microorganisms – bacteria or fungi, inside the bottles
or bags. Cultivation in polypropylene bags is simpler and cheaper; however, fruitbod-
ies growing from bags are typically smaller than those growing from bottles, where
usually a single large fruitbody is produced [5,45,46]. Similarly as in the cultivation
of other mushrooms, when growing lion’s mane mushroom waste from agriculture,
forestry, wood or food industries is used. Lion’s mane mushroom may be cultivated,
e.g., on sterilized sawdust supplemented with cereal bran [45,46].
Specialist literature sources present numerous recommendations concerning the
composition of substrate for lion’s mane mushroom culture [5,43,47]. Lion’s mane
mushroom may be grown, e.g., on wood coming from coniferous tree species, e.g.,
Pinus taeda and Pinus ponderosa. To obtain satisfactory yields chips produced from
these trees have to be rst spawned with mycelium of fungal species Aureobasidium
pullulans var. pullulans, Ceratocystis coerulescens, C. pilifera and Ophiostoma piliferum.
ese species, by degrading tar in wood of the above-mentioned species, make it suit-
able for lion’s mane mushroom growing [48].
Substrate was developed and composed in 55% of corn cobs, 20% cotton cha,
20% wheat bran, 3% corn meal, 1% gypsum and 1% sugar, which ensures very good
mycelium growth and high yields of supreme fruitbodies [47]. Another substrate vari-
ant developed by Zhang contains 65% corn cobs, 10% cotton cha, 20% wheat bran,
3% corn meal and 1% gypsum as well as 1% sugar. Similar materials for the substrate
for lion’s mane mushroom cultivation were used successfully: substrates formulated
by that author were sterilized and contained cotton cha (78%), wheat bran (20%),
gypsum (1%) and sugar (1%) [45].
Lion’s mane mushroom may be also grown on sterilized beech sawdust supple-
mented with 10% addition of wheat bran or 20% corn meal [49]. High yields were
produced when sawdust was supplemented with wheat bran and corn meal added at
20% and 7% [50] and glucose added at 3% [51]. Suitability of waste from the textile
industry to grow Hericium erinaceum was investigated [52]. ose authors obtained
the highest yields using ax shives.
Rice straw of cultivars with high amounts of nutrients may be used as an additive
enriching substrate [53]. In this case it should be added to the substrate at approxi-
mately 20% total substrate weight. Similarly as in the case of other mushrooms, the
addition of enriching substances to the substrate accelerates the process of primor-
dium formation and increases produced yields. Growing substrate may be supple-
mented with vegetable oils or fatty acids [54]. Among the seven substrate additives
tested a soy meal proved to be the best [41]. e addition of sunower husks as well as
Mn and NH4 to the cultivation substrate resulted in an accelerated mycelium growth
and increased yields [55]. e highest yields can be obtained when sugarcane is used
as an additive to growing substrate [35].
Chemical composition of Hericium erinaceum fruitbodies
Lion’s mane mushroom has a relatively high nutritive value. Fruitbodies of lion’s mane
mushroom contain 57% carbohydrates, 3.52% fats, 7.81% ber, 22.3% protein and
9.35% ash per dry matter (d.m.). Moreover, the following soluble sugars were also
found: arabitol at 127.17 mg/g, glucose at 11.35 mg/g, mannitol at 12.98 mg/g, inositol
at 1.43 mg/g and trehalose at 9.71 mg/g d.m. [56].
Detailed studies showed also contents of 14 amino acids in fruitbodies of this spe-
cies. Among the detected amino acids the highest share was recorded for L-alanine at
2.43 mg/g d.m. and L-leucine at 2.38 mg/g d.m. e lowest contents were found for
L-tryptophan at 0.10 mg/g d.m. and L-phenylalanine at 0.20 mg/g d.m. L-isoleucine
and L-tyrosine were not detected [56]. Lion’s mane mushroom contains consider-
able amounts of potassium and phosphorus, i.e., 254 and 109 mg/100 g dry matter,
respectively [57]. e above-mentioned authors found also 19 amino acids and 32

5 of 18© The Author(s) 2016 Published by Po lish Botanical Societ y Acta Mycol 50(2):1069
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
Fig. 1 Cultures of Her icium erinaceum developed in the bottles. e expanding mycelium is clearly visible inside the bottles.
Fig. 2 Young fruitbodies of lion’s mane mushroom are growing out of the bottlenecks.

6 of 18© The Author(s) 2016 Published by Po lish Botanical Societ y Acta Mycol 50(2):1069
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
aroma substances. Manganese, copper and zinc are found in Hericium in very low,
trace amounts. Contents of heavy metals, i.e., arsenic, lead, copper, and cadmium,
were higher in mycelium than in fruitbodies [58].
Analyses were also conducted on contents of aroma compounds in fruitbodies of
lion’s mane mushroom. e total content of volatile aroma compounds was deter-
mined in fruitbodies of lion’s mane mushroom [59]. e dominant compound was
1-octen-3-ol, which accounted for 56–60% total content of aroma substances. In other
study it was showed that the dominant compounds are 2-methyl-3-furanthiol, 2-eth-
ylpyrazine and 2,6-diethylpyrazine [60]. Sixteen aroma substances were identied,
containing nitrogen or sulfur, aldehydes, ketones, alcohols, and esters [61].
Biologically active substances in fruitbodies
Studies conducted in recent years have led to the isolation from lion’s mane mush-
room of several biologically active metabolites [62,63]. Both fruitbodies and myce-
lium of lion’s mane mushroom contain health-promoting compounds. e cultivation
on H. erinaceum may be optimized in order to obtain fruiting bodies reach in selected
substances [64,65]. e most important groups of compounds include polysaccha-
rides – for example: xylan (Fig. 6a), hericenones, and erinacines.
It was found that the total content of polysaccharides in fruitbodies of lion’s mane
mushroom exceeds considerably that in mycelium (26.63% and 18.71%, respectively)
[66]. e qualitative and quantitative composition of polysaccharides in fruitbodies of
lion’s mane mushroom were investigated too [67,68]. ere were the greatest content
of arabinose, followed by glucose and ramnose in fruitbodies. e ratio of arabinose
to glucose contents was 2.3 : 1. Total carbohydrates in mycelium of lion’s mane mush-
room were determined [69]. e authors also isolated 12 polysaccharides from the H.
erinaceum extract and determined their molecular mass. New heteropolysaccharide
(HEPF4) was isolated [70]. It was composed of D-galactose, L-fucose, D-glucose, and
methylramnose.
e latest study indicates that the application of enzymatic extraction results in a
67.6% increase in the amount of polysaccharides in relation to hot water extraction
Fig. 3 General view of Hericium erinaceum culture in bags. Developing fruiting bodies growing out of bags with substrate.

7 of 18© The Author(s) 2016 Published by Po lish Botanical Societ y Acta Mycol 50(2):1069
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
[71]. us obtained polysaccharide
fraction was composed of man-
nose, glucose, xylose, and galactose
at a ratio of 15.16 : 5.55 : 4.21 : 1. In
another study stated that polysac-
charides obtained from mycelium
of lion’s mane mushroom showed
greater immunostimulatory activ-
ity aer a 10% addition of ginseng
extract to the growing medium
[72].
Benzyl alcohol derivatives iso-
lated from fruitbodies of lion’s mane
mushroom were named hericenones
A, B, C, D, E, F, G, H [73–75]. Her-
cicenones A and B showed strong
cytotoxic properties towards HeLa
cells. In the course of the investiga-
tions minimum concentrations were
determined, at which these sub-
stances completely inhibited growth
of HeLa cells. In turn, hericenones C,
D and E stimulated the synthesis of
the nerve growth factor (NGF). Ac-
tivity of individual hericenones dif-
fered depending on the chain length
and the presence of double bonds in
fatty acids. e greatest capacity to
stimulate NGF was found for herice-
none E, having two double bonds in
its chain [76].
Studies conducted by several
other authors in fungal mycelium
showed the presence of diterpenoid
derivatives – for example diter-
pene (Fig. 6b), which were named
erinacines A, B, C, E, F, G, H, I and
P. Erinacines A, B, C, E, F and H
showed strong activity inducing the
synthesis of the NGF both in vitro and in vivo [77–80]. Studies made it possible to
determine the structure of the fatty acid coming from Hericium erinaceum [81]. is
acid exhibited an inhibitory action towards HeLa cells. e safety of erinacine A ap-
plication in animals was conrmed [82]. Erinacines, labeled J and K were isolated
[83]. Detailed characteristic of diterpenoid compounds found in Hericium sp. was
presented [84]. ose authors also showed the uniform distribution of terpenoid sec-
ondary metabolites in fruitbodies of H. erinaceum.
Other studies have led to the isolation of glycoside and 6 ergosterol derivatives
from dried fruitbodies of lion’s mane mushroom [85]. Another research on mycelium
in liquid cultures led to the isolation of novel metabolites, which showed antibacte-
rial and fungicidal properties, inhibiting growth of such microorganisms as Bacillus
subtilis, Saccharomyces cerevisiae, Verticilium dahlie and Aspergillus niger [86]. From
mycelium of H. erinaceum was isolated ergosterol peroxide, a cytotoxic steroidal
derivative [87]. In next study new isohericerines were isolated from H. erinaceum
– for instance hericerin A (Fig. 6c) [88]. Aer this research a new alkaloid with an
anti-inammatory feature named hericirine was isolated (Fig. 6d) [89] and a new
glycoprotein compound was characterized: HEG-5 [90]. It has hemagglutinating and
antineoplastic properties.
Lion’s mane mushroom is one of the best known medicinal mushrooms, but new
metabolites are still described, e.g., ten new isoindolin-1 ones, named erinacerins C–L
were isolated from the solid culture of H. erinaceus [91].
Fig. 4 Developing fruiting body of Hericium erinaceum cultivated in bag.

8 of 18© The Author(s) 2016 Published by Po lish Botanical Societ y Acta Mycol 50(2):1069
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
Medicinal functions of
Hericium erinaceum
For centuries H. erinaceum has been
used as an anti-inammatory medi-
cine, as well as an immune system
booster. In China a medication Wei
Le Xin Chong Ji is commercially
available, which is used against ul-
cers, inammations and neoplasms
of the alimentary tract. Lion’s mane
mushroom is administered as tab-
lets from dried and powdered fruit-
bodies. An advantageous action on
hepatitis B patients was also shown
for fresh fruitbodies of lion’s mane
mushroom. us, H. erinaceum
has been a medication which use
was justied by achievements and
tradition of Chinese medicine. Its
fruitbodies and preparations pro-
duced from them were treated as
excellent medicinal material. How-
ever, there were no results of physi-
ological and pharmaceutical tests
adopted as a standard at the intro-
duction of new medicinal prepara-
tions on the market in Europe and
the USA [5,6,92–94]. Extracts (raw
extract or its fractions) were thus
subjected to standard tests in order
to exclude their potential harmful
eects and determine mechanisms
of their bioactive action, in this way
conrming their health-promoting
properties.
Research conducted in recent
years showed that lion’s mane mush-
room may have an advantageous
eect in treatment of Alzheimer’s
disease [95] and leukemia, enhanc-
ing the mechanism of apoptosis of
cancer cells (Tab . 1 ) [96]. Another
study showed eectiveness of an extract from fruitbodies of Lion’s mane mushroom in
controlling: cancer cells in human cell culture [97] and intestine cancer cells in mice
[98]. Although most studies were conducted on animals, their results are considered
to be very promising. e results of preliminary clinical trials also showed that the
mushroom was eective in treatment of patients with dementia [99].
Experiments were conducted on antimutagenic properties of water and alcohol
extracts produced from mycelium and fruitbodies of lion’s mane mushroom [100].
In their study they used Salmonella typhimurium TA98 and dierent mutagenic com-
pounds. ese experiments showed antimutagenic action of the above-mentioned
extracts, with a stronger eect of ethanol extract in comparison to water extracts.
Extracts from mycelium showed a weaker action in relation to extracts produced from
fruitbodies. It was also found that lion’s mane mushroom contains lectins [101]. Lec-
tins from Hericium sp. did not exhibit mitogenic properties [102].
Polysaccharides were isolated both from liquid medium, in which lion’s mane
mushroom was cultured [103], and from fruitbodies [104]. ese compounds showed
strong antineoplastic properties in studies conducted on mice. Polysaccharides caused
an increase in the numbers of lymphocytes and macrophages and thus boosted the
Fig. 5 e fully developed mature fruiting body of Hericium erinaceum obtained
from cultivation. In the background are visible an empty bag and the rest of substrate.

9 of 18© The Author(s) 2016 Published by Po lish Botanical Societ y Acta Mycol 50(2):1069
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
immune system. ey have signicant anti fatigue activity [105]. e capacity to acti-
vate macrophages by water extracts from H. erinaceum was also conrmed [106,107].
HEP3 β-D-glucan (Fig. 6e) was characterized – obtained from fruitbodies of H. eri-
naceum [108]. A new heteropolysaccharide was isolated from fruitbodies of lion’s
mane mushroom [109]. It contains fucose, galactose and glucose as well as ramnose.
e experiments on rats showed immunomodulatory properties of polysaccharides
obtained from lion’s mane mushroom, which was manifested in an increase in the
secretion of cytokines IL-12, IFN-γ and IL-10 [110]. Polysaccharides obtained from
H. erinaceum are capable of enhancing the activity of antioxidant enzymes in the skin,
thus providing anti-ageing properties [111].
Ethanol extract from lion’s mane mushroom counteracts neuron death [112].
Ethanol extracts from H. erinaceum contain the NGF, which level is decreased in
Alzheimer’s patients, and this eect may prove helpful in treatment of this disease
[95,113–115]. It can be used to control pain in diabetic neuropathy [116] and in
treating Parkinson’s disease [117]. Administration of lion’s mane mushroom extracts
enhances learning abilities and improves memory [118]. Extracts of lion’s mane
mushroom are appropriate in the prevention of dementia thanks to the enhancement
of cognitive functions [119].
It was also found that methanol extracts from dried fruitbodies of lion’s mane
mushroom show antioxidant properties [120,121]. Antioxidant properties of metha-
nol extracts are determined by the presence of polyphenols [122]. In studies on
animals were showed antioxidant properties of polysaccharides obtained from H. eri-
naceum [123]. e product obtained by acetic acid fermentation using H. erinaceum
as a substrate presented antioxidant properties [124].
Fig. 6 e structures of compounds: xylan (a), diterpene (b), hericerin A (c), hericerin A (d), β-D-glucan (e), hericenone
B (h), erinacene D (g).

10 of 18© The Author(s) 2016 Published by Polish B otanical Society Ac ta Mycol 50(2):1069
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
In experiments on animals [125] was shown the protective eect of water extract
of lion’s mane mushroom in gastric ulcers. In another study [126] a protective eect
on gastric mucosa was shown for dierent fractions obtained from ethanol extract
from lion’s mane mushroom. e polysaccharide complex obtained from lion’s mane
mushroom was eective against Helicobacter pylori, responsible for many gastric dis-
orders [127]. e latest study [128] conducted in vitro conrmed the eectiveness
of H. erinaceum extract against hepatic cancer cells HepG2 and Huh-7, colon cancer
cells HT-29 and gastric cancer cells NCI-87. is indicates potential applications of
lion’s mane mushroom as a drug against gastrointestinal cancers.
ere was a protective eect of methanol extract from lion’s mane mushroom on
liver cells [129]. ree polysaccharide fractions from lion’s mane mushroom (HEP40,
HEP60 and HEP80) has an applicability in the prevention of hepatic disorders [70].
In the opinion of many researchers, the potential hepatoprotective eect observed
in vivo is connected with the antioxidant capacity conrmed in vitro. Hericium eri-
naceum has the capacity to induce apoptosis of human hepatocellular carcinoma cells
[130]. Hericium erinaceum supplementation could restrain the hepatic damage caused
by acute alcohol exposure [131].
Many researchers showed other health-promoting eects of lion’s mane mushroom
[132]. Female students who took drugs with its extracts declared the improvement of
the quality of sleep [133]. Depression or anxiety were found much less frequently and
were less intensive in menopausal women, who were administered powdered fruit-
bodies of lion’s mane mushroom in comparison to the group receiving placebo [134].
Drugs with their extracts are benecial for treating primary cognitive decits and
negative symptoms of schizophrenia [135].
Other studies showed that water extracts promote healing process in open wounds
and reduce the size and visibility of scars in scaried rats [136]. Another study [137]
showed the eectiveness of water-soluble compounds isolated from lion’s mane mush-
room in the enhancement of immune response and in treatment of wounds. In turn,
methanol extracts and substances they contained, i.e., D-threitol and D-arabinitol,
as well as palmitic acid were capable of reducing blood lipid and glucose levels in
examined rats [138].
Hericenone B (Fig. 6f) obtained from lion’s mane mushroom is capable of inhibit-
ing blood platelet aggregation [139]. Hericium erinaceum may regulate functions of
Tab. 1 Medicinal value of Hericium erinaceum.
Bioactive compounds Activity Treatment
Polysaccharides Immunomodulatory Cancers
Anticancer Gastrointestinal cancers (liver, gastric, colorec-
tal), leukemia
Anti-bacterial Helicobacter pylori infection
Gastroprotective Ulcers, chronic gastritis
Cholesterol and triglyceride lowering Hyperlipidemia
Hepatoprotective Hepatic tissue damage
Blood glucose lowering Diabetes
Hericenones A–B Cytotoxic Cancers
Anti-platelet aggregation Vascular diseases, stroke, thrombosis
Hericenones C–H, eri-
nacines A–I
Neuroprotective, neuroregenerative Alzheimer’s and Parkinson’s diseases, demen-
tia, depression
Hericirine Reduction of pro-inammatory media-
tors and cytokines
Inammatory diseases
Poliphenols Antioxidative Anti-skin aging

11 of 18© The Author(s) 2016 Published by Polish Botan ical Society Acta M ycol 50(2):1069
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
References
1. Wojewoda W. Encyklopedia biologiczna. Kraków: OPRESS; 1998.
2. Index Fungorum [Internet]. 2016 [cited 2016 Jan 19]. Available from: http://www.index-
fungorum.org
3. Persoon CH. CH Persoonii Commentatio de fungis clavaeformibus sistens specierum hus
usque notarum descriptiones cum dierentiis specicis nec non auctorum synonymis:
accedunt tab. IV, colore fucatae. Apud Petrum Philippum Wolf; 1797.
4. Gumińska B, Wojewoda W. Grzyby i ich oznaczanie. Warszawa: PWRiL; 1985.
5. Stamets P. Growing gourmet and medicinal mushrooms. Berkeley, CA: Ten Speed Press;
1993.
6. Pegler DN. Usef ul fungi of the world: the monkey head fungus. Mycologist. 2003;17(3):120–
121. http://dx.doi.org/10.1017/S0269915X03003069
7. Fora CG, Lauer KF, Stefan C, Banu C. Hericium erinaceus and Sacroscypha coccinea
in deciduous forest ecosystem. Journal of Horticulture, Forestry and Biotechnology.
2009;13:67–68.
8. Wojewoda W. Checklist of Polish larger Basidiomycetes. Kraków: W. Szafer Institute of
Botany, Polish Academy of Sciences; 2003.
9. Stasinska M. Macromycetes in forest communities of the Insko Landscape Park (NW Po-
land). Acta Mycol. 1999;34(1):125–168. http://dx.doi.org/10.5586/am.1999.011
10. Domański S. Materiały do poznania mikoory nadrzewnej Beskidu Niskiego w okolicy
Gorlic. Fragm Flor Geobot Pol. 1961;7(1):203–213.
11. Domański S, Lisiewska M, Majewski T, Skirgiełło A, Truszkowska W, Wojewoda W. Miko-
ora Bieszczadów Zachodnich. Acta Mycol. 1970;6(1):63–114.
12. Wilga MS. Grzyby makroskopowe (Macromycetes) występujące na obszarze Gdańska.
Acta Bot Cassub. 2000;1:105–111.
13. Doll R. Die Verbreitung der gestielten Stachelpilze sowie das Vorkommen von Hericium,
Creolophus cirrhatus, Spongipellis pachyodon und Sistotrema conuens in Mecklenburg.
Feddes Repert. 1979;90(1–2):103–120. http://dx.doi.org/10.1002/fedr.19790900107
14. Zhou GY, Liu LZ. First discovery of Hericium erinaceus in Qinghai Prefecture. Edible
Fungi of China. 1988;6(27).
15. Mao XL. An investigation of edible fungi in the Changbai Mountains in Jilin Province.
Edible Fungi of China. 1989;1:25–27.
16. Huang YG, Barl B, Ivanochko G. Selected non-timber forest products with medicinal ap-
plications from Jilin Province in China. In: General technical report – North Central Re-
search Station. Meeting: forest communities in the third millennium: linking research,
business, and policy toward a sustainable non-timber forest product sector; 1999 Oct 1–4;
Kenora, Ontario, Kanada. Saint Paul, OH: United States Department of Agriculture; 2001.
17. Grace J, Mudge KW. Production of Hericium sp. (lion’s mane) mushrooms on totem logs
in a forest farming system. Agroforestry Systems. 2015;89(3):549–556. http://dx.doi.
org/10.1007/s10457-015-9790-1
18. Boddy L, Crockatt ME, Ainswort AM. Ecology of Hericium cirratum, H. corraloides
and H. erinaceus in the UK. Fungal Ecol. 2011;4:163–173. http://dx.doi.org/10.1016/j.
funeco.2010.10.001
19. Burdsall Jr. HH, Miller Jr. OK, Nishijima KA. Morphological and mating system studies of
a new taxon of Hericium (Aphyllophorales, Hericaceae) from the Southern Apallachians.
Mycotaxon. 1978;7(1):1–9.
the nervous, digestive, circulatory and immune systems of the organism, which would
promote overall human health [140]. Substances from H. erinaceum may be used in
the treatment and prevention of age-related diseases [141]. An addition of polysac-
charides from lion’s mane mushroom to feed for broiler chickens may lead to the
production of meat with a reduced cholesterol level [142]. ere were described some
new compounds for example erinacene D (Fig. 6g) [143].

12 of 18© The Author(s) 2016 Published by Polish Bot anical Society Act a Mycol 50(2):10 69
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
20. Lu L, Li J. Cang Y. PCR-based sensitive detection of medicinal fungi Hericium species from
ribosomal internal transcribed spacer (ITS) sequences. Biol Pharm Bull. 2002;25(8):975–
980. http://dx.doi.org/10.1248/bpb.25.975
21. Ginns J. Hericium coralloides N. Amer. auct. (= H. americanum sp. nov.) and the European
H. alpestre and H. coralloides. Mycotaxon. 1984;20(1):39–43.
22. Adair S, Kim SH, Breuil C. A molecular approach for early monitoring of decay Basid-
iomycetes in wood chips. FEMS Microbiol Lett. 2002;211(1):117–122. http://dx.doi.
org/10.1111/j.1574-6968.2002.tb11212.x
23. Partt D, Hynes J, Rogers HJ, Boddy L. New PCR assay detects rare tooth fungi in wood
where traditional approaches fail. Mycol Res. 2005;109(11):1187–1194. http://dx.doi.
org/10.1017/S095375620500359X
24. Buchalo AS, Wasser SP, Reshetnikov SV, Grigansky AP. Studies on microstructures of veg-
etative mycelium in the medicinal mushrooms Hericium erinaceus (Bull.: Fr.) Pers. and
Grifola frondosa (Dicks.: Fr.) S.F. Gray (Aphyllophoromycetideae). Int J Med Mushrooms.
1999;1(3):235–241. http://dx.doi.org/10.1615/IntJMedMushrooms.v1.i3.60
25. Zhang XM, Zhang HY, Zhang Y. Observation of Chlamydospores produced from Heri-
cium erinaceus monocaryotic mycelium. Mycosystema. 2003;22(3):436–440.
26. McCracken FI. Spore production of Hericium erinaceus. Phytopathology. 1970;60:1639–
1641. http://dx.doi.org/10.1094/Phyto-60-1639
27. Feng Z, Liu X, Lu Y. A study on the eect and mechanism of 2,4-D and gibberelin on
Hericium erinaceus. Acta Edulis Fungi. 1998;5(3):42–47.
28. Poyedynok NL, Potemkina JV, Buchalo AS, Negriyko AM, Grygansky AP. Stimulation
with low – intensity laser light of basidiospore germination and growth of monokaryotic
isolates in the medicinal mushroom Hericium erinaceus (Bull.: Fr.) Pers. (Aphyllophoro-
mycetideae): memorial issue dedicated to Professor Takashi Mizuno. Int J Med Mush-
rooms. 2000;2(4):339–342.
29. Poyedynok NL, Buchalo AS, Negriyko AM, Potemkina JV, Mykchaylova OB, e action
of argon and helium laser radiation on growth and fructication of culinary – medicinal
mushroom Pleurotus ostreatus (Jacq.: Fr.) Kumm., Lentinus edodes (Berk.) Singer, and He-
ricium erinaceus (Bull.: Fr.) Pers. Int J Med Mushrooms. 2003;5:293–299. http://dx.doi.
org/10.1615/InterJMedicMush.v5.i3.70
30. Imtiaj A, Jayasinghe C, Lee GW, Shim MJ, Rho HS, Lee HS, et al. Vegetative growth
of four strains of Hericium erinaceus collected from dierent habitats. Mycobiology.
2008;36(2):88–92. http://dx.doi.org/10.4489/MYCO.2008.36.2.088
31. Le Minh T, Chinh NT, Uyen NT, Bang ND. Study on eect of light rare earth-isoleucine
chelate on Hericium erinaceum growth. Khoa Hoc Va Cong Nghe. 2009;47(5):27–32.
32. Kim YD, Ha KY, Lee JK, Kim SD. Variability of rice koji enzyme activities using Basidio-
mycete. Int Rice Res Notes. 2000;25(3):10.
33. Han J. Solid-state fermentation of cornmeal with the basidiomycete Hericium erinaceum
for degrading starch and upgrading nutritional value. Int J Food Microbiol. 2003;80:61–66.
34. Du F, Wang HX, Ng TB. An amylase from fresh fruiting bodies of the monkey head mush-
room Hericium erinaceum. Appl Biochem Microbiol. 2013;49(1):23–27. http://dx.doi.
org/10.1134/S0003683813010043
35. Hu SH, Wang JC, Wu CY, Hsieh SL, Chen KS, Chang SJ, et al. Bioconversion of agro wastes
for the cultivation of culinary-medicinall lion’s mane mushrooms Hericium erinaceus
(Bull.: Fr.) Pers. and H. laciniatum (Leers) Banker (Aphyllophoromycetideae) in Taiwan.
Int J Med Mushrooms. 2008;10(4):385–398. http://dx.doi.org/10.1615/IntJMedMushr.
v10.i4.120
36. Li F, Zhu X, Li N, Zhang P, Zhang S, Zhao X, et al. Screening of lignocellulose-degrading
superior mushroom strains and determination of their CMCase and laccase activity. e
Scientic World Journal. 2014;1–6.
37. Sun SJ, Liu JZ, Hu KH, Zhu HX. e level of secreted laccase activity in the edible fungi
and their growing cycles are closely related. Curr Microbiol. 2011;62:871–875. http://
dx.doi.org/10.1007/s00284-010-9794-z
38. Wang MX, Guan D, Jing MD. Nutritional composition analysis of ve strains of Hericium
erinaceus and their utilization. Microbiology. 1992;19(2):68–72.
39. Ko HG, Park SH, Kim SH, Park HG, Park WM. Detection and recovery of hydrolytic
enzymes from spent compost of four mushroom species. Folia Microbiol (Praha).
2005;50(2):103–106. http://dx.doi.org/10.1007/BF02931456

13 of 18© The Author(s) 2016 Published by Polish Bo tanical Society Ac ta Mycol 50(2):10 69
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
40. Phan CW, Sabaratnam V. Potential uses of spent mushroom substrate and its associated
lignocellulosic enzymes. Appl Microbiol Biotechnol. 2012;96:863–873. http://dx.doi.
org/10.1007/s00253-012-4446-9
41. Ko HG, Park HG, Park SH, Choi CW, Kim SH, Park WM. Comparative study of myce-
lia growth and basidiomata formation in seven dierent species of the edible mushroom
genus Hericium. Bioresour Technol. 2005;96:1439–1444. http://dx.doi.org/10.1016/j.
biortech.2004.12.009
42. Grygansky AP, Solomko EF, Kirchho B. Mycelial growth of medicinal mushroom Heri-
cium erinaceus (Bull.: Fr.) Pers. in pure culture. Int J Med Mushrooms. 1999;1(1):81–87.
http://dx.doi.org/10.1615/IntJMedMushrooms.v1.i1.60
43. Wang H. e cultivation of Hericium erinaceus in the bottle. Edible Fungi of China.
2002;21(3):22.
44. Zhao ZG, Yan XL, Wang WY. A study on the physiological characteristics of Hericium
erinaceus (Fr.) Pers. Hyphae. Edible Fungi of China. 1998;1:5–6.
45. Royse DJ. Specialty mushrooms. Progress in new crops. In: Proceedings of the ird Na-
tional Symposium; 1995 Oct 22–25; Indianapolis, Indiana, USA. Alexandria: ASHS Press;
1996.
46. Oei P. Mushroom cultivation, appropriate technology for mushroom growers. Leiden:
Backhuys Publishers; 2003.
47. J. Zhang, Study on the experiment for the cultivation of Hericium erinaceus with corn
cobs. Edible Fungi of China. 2000;19(2):14.
48. Croan SC. Bioconversion of conifer wood chips into specialty mushroom producing fun-
gal growth. Mushrooms International. 2002;90:7–11.
49. Kirchho B. Biotechnological investigations of Hericium erinaceum (Bull.: Fr.) Pers. bag-
log cultivation to increase yield. In: Royse DJ, editor. Proceedings of the 2nd International
Conference on Mushroom Biology and Mushroom Products; 1996 Jun 9–12; University
Park, Pennsylvania, USA. University Park, PA: Pennsylvania State University; 1996.
50. Siwulski M, Sobieralski K. Inuence of some growing substrate additives on the Hericium
erinaceus (Bull. Fr.) Pers. yield. Sodinink Darzinink. 2007;24(3):250–253.
51. Siwulski M, Sobieralski K, Wojniłowicz M. Comparison of mycelium growth and yield-
ing of selected strains of Hericium erinaceus (Bull.: Fr.) Pers. on sawdust substrates with
glucose addition. Herba Polonica. 2009;55(3):266–272.
52. Siwulski M, Sobieralski K, Mańkowski J, Comparison of mycelium growth of selected spe-
cies of cultivated mushrooms on textile industry wastes. Acta Sci Pol Hortorum Cultus.
2010;9(3):37–43.
53. Zheng JG, Chen JC, Yang J, Zheng KB, Ye XF, Huang QL, et al. Studies on growing edible
fungi on improved straw from a dual use rice cultivar. Agric Sci China. 2002;1(8):871–877.
54. Kang H, Hwang S, Lee H, Park W. Eects of high concentrations of plant oils and fatty
acids for mycelial growth and pinhead formation of Hericium erinaceum. Trans ASAE.
2002;45(1):257–260. http://dx.doi.org/10.13031/2013.7854
55. Figlas D, Matute RG, Curvetto N. Cultivation of culinary-medicinal lion’s mane mushroom
Hericium erinaceus (Bull.: Fr.) Pers. (Aphyllophoromycetideae) on substrate containing
sunower seed hulls. Int J Med Mushrooms. 2007;9(1):67–73. http://dx.doi.org/10.1615/
IntJMedMushr.v9.i1.80
56. Mau JL, Lin HC, Ma JT, Song SF. Non-volatile taste components of several speciality mush-
rooms. Food Chem. 2001;73:461–466. http://dx.doi.org/10.1016/S0308-8146(00)00330-7
57. Eisenhut R, Fritz D, Tiefel P, Investigations on nutritionally valuable constituents (mineral
substances, amino acids, aromatic substances) of Hericium erinaceus (Bull.: Fr.) Pers. Eur
J Hortic Sci. 1995;60(5):212–218.
58. Yang Y, Zhou CY, Zhang JS, Tang QJ. Comparison of chemical component and bio-
logical activity of Hericium erinaceus fruit body and mycelial extracts. Junwu Yanjiu.
2006;4(3):15–19.
59. Siwulski M, Zawirska-Wojtasiak R. Comparison of chemical composition and content of
volatile compounds of carpophores of Hericium erinaceus (Bull.: Fr.) Pers. cultivated on
dierent types of sawdust. Herba Polonica. 2007;53(3):218–223.
60. Miyazawa M, Matsuda N, Tamura N, Ishikawa R. Characteristic avour of volatile oil from
dried fruiting bodies of Hericium erinaceus (Bull.: Fr.) Pers. Journal of Essential Oil Re-
search. 2008;20:420–423.

14 of 18© The Author(s) 2016 Published by Polish B otanical Society Ac ta Mycol 50(2):1069
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
61. Miyazawa M, Usami A, Character impact odorants from mushrooms [Pleurotus citrino-
pileatus, Pleurotus eryngii var. ferulae, Lactarius hatsudake, and Hericium erinaceus (Bull.:
Fr.) Pers.] used in Japanese traditional food. Nagoya Gaknin University. 2014;50(2):1–24.
62. Qian FG, Xu GY, Du SJ, Li MH. Isolation and identication of two new pyrone compounds
from the culture of Hericium erinaceus. Yao Xue Xue Bao. 1990;25(7):522–525.
63. Kenmoku H, Kato N, Shimada M, Omoto M, Mori A, Mitsuhashi W, et al. Isolation of
(-)-cyatha-3,12-diene, a common biosynthetic intermediate of cyathane diterpenoids,
from an erinacine producing basidomycte, Hericium erinaceum, and its formation in
a cell-free system. Tetrahedron Lett. 2001;42:7439–7442. http://dx.doi.org/10.1016/
S0040-4039(01)01550-7
64. Dai X, Zhan Y, Zhang J, Zhang P, Han Z, Ma Q, et al. Regulatory eect of salicylic acid
and methyl jasmonate supplementation on ergosterol production in Hericium erinaceus
mycelia. Journal of Forestry Research. 2015;26(1):71–77. http://dx.doi.org/10.1007/
s11676-014-0014-8
65. Gerbec B, Tavcar E, Gregori A, Kre S, Berovic M. Solid state cultivation of Hericium
erinaceus biomass and erinacine A production. J Bioprocess Biotech. 2015;5(2):210. http://
dx.doi.org/10.4172/2155-9821.1000210
66. Mori H, Aizawa K, Inakuma T, Ichii A, Yamauchi R, Kato K. Structural analysis of the
β-D-glucan from the fruit-body of Hericium erinaceum. Journal of Applied Glycoscience.
1998;45(4):361–365.
67. Keong CY, Badrul A, Ing YS, Zakiah I. Quantication and identication of polysaccharide
contents in Hericium erinaceus. Nutrition and Food Science. 2007;37(4):260–271. http://
dx.doi.org/10.1108/00346650710774631
68. Wu DT, Li WZ, Chen J, Zhong QX, Ju YJ, Zhao J, et al. An evaluation system for charac-
terization of polysaccharides from the fruiting body of Hericium erinaceus and identi-
cation of its commercial product. Carbohydr Polym. 2015;124:201–207. http://dx.doi.
org/10.1016/j.carbpol.2015.02.028
69. Malinowska E, Krzyczkowski W, Łapiens G, Herold F. Densitometric determination of
carbohydrates: application to purication and molecular weight determination of poly-
saccharides from Hericium erinaceum mushroom. Food Res Int. 2010;43:988–995. http://
dx.doi.org/10.1016/j.foodres.2010.01.011
70. Zhang Z, Lv G, Pan H, Pandey A, He W, Fan L, Antioxidant and hepatoprotective potential
of endo-polysaccharides from Hericium erinaceus grown on tofu whey. Int J Biol Macro-
mol. 2012;51:1140–1146. http://dx.doi.org/10.1016/j.ijbiomac.2012.09.002
71. Zhu Y, Li Q, Mao G, Zou Y, Feng W, Zheng D, et al. Optimization of enzyme-assisted
extraction and characterization of polysaccharides from Hericium erinaceus. Carbohydr
Polym. 2014;101:606–613. http://dx.doi.org/10.1016/j.carbpol.2013.09.099
72. Kim H, Ra KS, Hwang JH, Yu KW. Immune enhancement of Hericium erinaceum myce-
lium cultured in submerged medium supplemented with ginseng extract. Journal of Food
Science and Nutrition. 2012;25(4):737–746. http://dx.doi.org/10.9799/ksfan.2012.25.4.737
73. Kawagishi H, Ando M, Mizuno T. Hericenone A and B as cytotoxic principles from the
mushroom Hericium erinaceum. Tetrahedron Lett. 1990;31(3):373–376. http://dx.doi.
org/10.1016/S0040-4039(00)94558-1
74. Kawagishi H, Ando M, Shniba K, Sakamoto H, Yoshida S, Ojima F, et al. Chromans,
hericenones F, G and H from the mushroom Hericium erinaceum. Phytochemistry.
1993;32(1):175–178. http://dx.doi.org/10.1016/0031-9422(92)80127-Z
75. Kawagishi H, Ando M, Sakamoto H, Yoshida S, Ojima F, Ishiguro Y, et al. Hericenones
C, D and E, stimulators of nerve growth factor (NGF) – synthesis, from the mush-
room Hericium erinaceum. Tetrahedron Lett. 1991;32(35):4561–4564. http://dx.doi.
org/10.1016/0040-4039(91)80039-9
76. Phan CW, Lee GS, Hong SL, Wong YT, Brkljača R, Urban S, et al. Hericium erinaceus (Bull.:
Fr) Pers. cultivated under tropical conditions: isolation of hericenones and demonstration
of NGF-mediated neurite outgrowth in PC12 cells via MEK/ERK and PI3K-Akt signaling
pathways. Food Funct. 2014;5:3160–3169. http://dx.doi.org/10.1039/C4FO00452C
77. Kawagishi H, Shimada A, Shirai R, Okamoto K, Ojima F, Sakamoto H, et al. Erinacines
A, B, and C, strong stimulators of nerve growth factor (NGF) – synthesis, from the my-
celia of Hericium erinaceum. Tetrahedron Lett. 1994;35(10):1569–1572. http://dx.doi.
org/10.1016/S0040-4039(00)76760-8
78. Kawagishi H, Shimada A, Hosokawa S, Mori H, Sakamoto H, Ishiguro Y, et al. Eri-
nacines E, F, and G, stimulators of nerve growth factor (NGF) – synthesis, from the

15 of 18© The Author(s) 2016 Published by Polish Bo tanical Society Ac ta Mycol 50(2):10 69
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
mycelia of Hericium erinaceum. Tetrahedron Lett. 1996;37(41):7399–7402. http://dx.doi.
org/10.1016/0040-4039(96)01687-5
79. Kenmoku H, Sassa T, Kato N. Isolation of erinacine P, a new parental metabolite of
cyathane-xylosides, from Hericium erinaceum and its biomimetic conversion into
erinacines A and B. Tetrahedron Lett. 2000;41:4389–4393. http://dx.doi.org/10.1016/
S0040-4039(00)00601-8
80. Lee EW, Shizuki K, Hosokawa S, Suzuki M, Suganuma H, Inakuma T, et al. Two novel di-
terpenoids, Erinacines H and I from the mycelia of Hericium erinaceum. Biosci Biotechnol
Biochem. 2000;64(11):2402–2405. http://dx.doi.org/10.1271/bbb.64.2402
81. Kawagishi H, Ando M, Mizuno T, Yokota H, Konishi S. A novel fatty acid from the
mushroom Hericium erinaceus. Agric Biol Chem. 1990;54(5):1329–1331. http://dx.doi.
org/10.1271/bbb1961.54.1329
82. Li IC, Chen YL, Lee LY, Chen WP, Tsai YT, Chen CC, et al. Evaluation of the toxico-
logical safety of erinacine A-enriched Hericium erinaceus in a 28-day oral feeding study
in Sprague-Dawley rats. Food Chem Toxicol. 2014;70:61–67. http://dx.doi.org/10.1016/j.
fct.2014.04.040
83. Kawagishi H, Masui A, Tokuyama S, Nakamura T. Erinacines J and K from the mycelia
of Hericium erinaceum. Tatrahedron Lett. 2006;62:8463–8466. http://dx.doi.org/10.1016/j.
tet.2006.06.091
84. Bhandari DR, Shen T, Rompp A, Zorn H, Spengler B. Analysis of cyathane-type diterpe-
noids from Cyathus striatus and Hericium erinaceus by high-resolution MALDI MS imag-
ing. Anal Bioanal Chem. 2014;406:695–704. http://dx.doi.org/10.1007/s00216-013-7496-7
85. Takaishi Y, Uda M, Ohashi T, Nakano K, Murakami K, Tomimatsu T. Glycosides of er-
gosterol derivatives from Hericium erinacens. Phytochemistry. 1991;30(12):4117–4120.
http://dx.doi.org/10.1016/0031-9422(91)83478-4
86. Okamoto K, Shimada A, Shire R, Sakamoto H, Yoshida S, Ojima F. Antimicrobial
chlorinated orcinol derivatives from mycelia of Hericium erinaceum. Phytochemistry.
1993;34(5):1445–1446. http://dx.doi.org/10.1016/0031-9422(91)80050-B
87. Krzyczkowski W, Malinowska E, Suchocki P, Kleps J, Olejnik M, Herold F. Isolation and
quantitative determination of ergosterol peroxide in various edible mushroom species.
Food Chem. 2009;113:351–355. http://dx.doi.org/10.1016/j.foodchem.2008.06.075
88. Miyazawa M, Takahashi T, Horibe I, Ishikawa R. Two new aromatic compounds and a new
D-arabinitol ester from the mushroom Hericium erinaceum. Tetrahedron. 2012;68:2007–
2010. http://dx.doi.org/10.1016/j.tet.2011.11.068
89. Li W, Zhou W, Lee DS, Shim SH, Kim YC, Kim YH. Hericirine, a novel anti-inamatory
alkaloid from Hericium erinaceum. Tetrahedron Lett. 2014;55:4086–4090. http://dx.doi.
org/10.1016/j.tetlet.2014.05.117
90. Cui FJ, Li YH, Zan XY, Yang Y, Sun WJ, Qian JY, et al. Purication and partial charac-
terization of a novel hemagglutinating glycoprotein from the cultured mycelia of He-
ricium erinaceus. Process Biochem. 2014;49:1362–1369. http://dx.doi.org/10.1016/j.
procbio.2014.04.008
91. Wang K, Bao L, Qi Q, Zhao F, Ma K, Pei Y, et al. Erinacerins C–L, isoindolin-1-ones with
α-glucosidase inhibitory activity from cultures of the medicinal mushroom Hericium eri-
naceus. J Nat Prod. 2015;78(1):146–154. http://dx.doi.org/10.1021/np5004388
92. Yang JY, Zhang SJ. Treatment of chronic hepatitis B with fresh Hericium erinaceus. Edible
Fungi of China. 1989;3:6.
93. Mizuno T. Bioactive substances in Hericium erinaceus (Bull.: Fr.) Pers. (Yamabushitake),
and its medicinal utilization. Int J of Med Mushrooms. 1999;1(2):105–119. http://dx.doi.
org/10.1615/IntJMedMushrooms.v1.i2.10
94. Jieli L. Studies on adenosine composition with medicinal eects of Hericium erinaceus and
its pharmaceutical products. Edible Fungi of China. 2002;21(3):32.
95. Mori K, Inatomi S, Ouchi K, Azumi Y, Tuchida T. Improving eects of the mushroom
Yamabushitake (Hericium erinaceus) on mid cognitive impairment: a double-blind pla-
cebo-controlled clinical trial. Phytother Res. 2009;23:367–372. http://dx.doi.org/10.1002/
ptr.2634
96. Kim SP, Kang MY, Choi YH, Kim JH, Nam SH, Friedman M. Mechanism of Hericium
erinaceus (Yamabushitake) mushroom-induced apoptosis of U937 human monocytic leu-
kemia cells. Food Func. 2011;2(6):348–356. http://dx.doi.org/10.1039/c1fo10030k
97. Li W, Zhou W, Kim EJ, Shim SH, Kang HK, Kim YH. Isolation and identication of

16 of 18© The Author(s) 2016 Published by Polish B otanical Society Ac ta Mycol 50(2):1069
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
aromatic compounds in lion’s mane mushroom and their anticancer activities. Food
Chem. 2015;170:336–342. http://dx.doi.org/10.1016/j.foodchem.2014.08.078
98. Kim SP, Kang MY, Kim JH, Nam SH, Friedman M. Composition and mechanism of an-
titumor eects of Hericium erinaceus mushroom extracts in tumor-bearing mice. J Agric
Food Chem. 2011;59:9861–9869. http://dx.doi.org/10.1021/jf201944n
99. Kawagishi H, Zhuang C. Compounds for dementia from Hericium erinaceum. Drugs Fu-
ture. 2008;33(2):149. http://dx.doi.org/10.1358/dof.2008.033.02.1173290
100. Wang JC, Hu SH, Su CH, Lee TM. Antitumor and immunoenhancing activities of polysac-
charide from culture broth of Hericium spp. Kaohsiung J Med Sci. 2001;17(9)461–467.
101. Kawagishi H, Mori H, Uno A, Kimura A, Chiba S. A sialic acid-binding lectin from
the mushroom Hericium erinaceum. FEBS Lett. 1994;340:56–58. http://dx.doi.
org/10.1016/0014-5793(94)80172-X
102. Ho JCK, Sze SCW, Shen WZ, Liu WK. Mitogenic activity of edible mushroom lectins.
Biochim Biophys Acta. 2004;1671:9–17. http://dx.doi.org/10.1016/j.bbagen.2003.12.009
103. Wang JC, Hu SH, Lee WL, Tsai LY. Antimutagenicity of extracts of Hericium erinaceus.
Kaohsiung J Med Sciences. 2001;17(5):230–238.
104. Wang Z, Luo D, Liang Z. Structure of polysaccharides from the fruiting body of Heri-
cium erinaceus Pers. Carbohydr Polym. 2004;57:241–247. http://dx.doi.org/10.1016/j.
carbpol.2004.04.018
105. Liu J, Du C, Wang Y, Yu Z. Anti-fatigue activities of polysaccharides extracted from
Hericium erinaceus. Exp er Med. 2015;9(2):483–487. http://dx.doi.org/10.3892/
etm.2014.2139
106. Son CG, Shin JW, Cho JH, Cho CK, Yun CH, Chung WC, et al. Macrophage activation and
nitric production by water soluble components of Hericium erinaceum. Int Immunophar-
macol. 2006;6(8):1363–1369. http://dx.doi.org/10.1016/j.intimp.2006.03.005
107. Lee JS, Cho JY, Hong EK. Study on macrophage activation and structural characteristics of
puried polysaccharides from the liquid culture broth of Hericium erinaceus. Carbohydr
Polym. 2009;78:162–168. http://dx.doi.org/10.1016/j.carbpol.2009.04.036
108. Dong Q, Jia LM, Fang JN. A β-D-glucan isolated from the fruiting bodies of Hericium
erinaceus and its aqueous conformation. Carbohydr Res. 2006;341:791–795. http://dx.doi.
org/10.1016/j.carres.2006.01.022
109. Zhang AQ, Sun PL, Zhang JS, Tang CH, Fan JM, Shi XM, et al. Structural investigation of a
novel fucoglucogalactan isolated from the fruiting bodies of the fungus Hericium erinaceus.
Food Chem. 2007;104:451–456. http://dx.doi.org/10.1016/j.foodchem.2006.11.033
110. Sheu SC, Lyu Y, Lee MS, Cheng JH. Immunomodulatory eects of polysaccharides iso-
lated from Hericium erinaceus on dendritic cells. Process Biochem. 2013;48:1402–1408.
http://dx.doi.org/10.1016/j.procbio.2013.06.012
111. Xu H, Wu P, Shen Z, Chen X. Chemical analysis of Hericium erinaceum polysaccharides and
eect of the polysaccharides on derma antioxidant enzymes, MMP-1 and TIMP-1 activi-
ties. Int J Biol Macromol. 2010;47:33–36. http://dx.doi.org/10.1016/j.ijbiomac.2010.03.024
112. Nagai K, Chiba A, Nishino T, Kubota T, Kawagishi H, Dilinoleoyl-phospathitidyletha-
nolamine from Hericium erinaceum protects against ER stress-dependent Neuro2a cell
death via protein kinase C pathway. J Nutr Biochem. 2006;17:525–530. http://dx.doi.
org/10.1016/j.jnutbio.2005.09.007
113. Mori K, Obara Y, Hirota M, Azumi Y, Kinugasa S, Inamtomi S, Nakahata N. Nerve growth
factor-inducing activity of Hericium erinaceus in 1321N1human astrocytoma cells. Biol
Pharm Bul. 2008;31(9):1727–1732. http://dx.doi.org/10.1248/bpb.31.1727
114. Xu T, Beelman RB. e bioactive compounds in medicinal mush-rooms have potential
protective eects against neurodegenerative diseases. Advances in Food Technology and
Nutritional Sciences. 2015;1(2):62–66. http://dx.doi.org/10.17140/AFTNSOJ-1-110
115. He CY, Chen SD, Lu WJ, Wang HC, Chang YC, Cheng YH, et al. Eect of ethanol extracts
from Hericium erinaceum solid-state fermented wheat on nerve cells growth. Hans Journal
of Food and Nutrition Science. 2015;4:1–10. http://dx.doi.org/10.12677/hjfns.2015.41001
116. Yi Z, Shao-long Y, Ai-hong W, Zhi-chun S, Ya-fen Z, Ye-ting X, et al. Protective eect
of ethanol extracts of Hericium erinaceus on alloxan-induced diabetic neuropathic
pain in rats. Evid Based Complement Alternat Med. 2015;2015:595480. http://dx.doi.
org/10.1155/2015/595480
117. Dhawan S, Chouhan U. Structural and functional characterization of Hericium erinaceum,

17 of 18© The Author(s) 2016 Published by Polish Bot anical Society Act a Mycol 50(2):10 69
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
manganese peroxidase as an antioxidant against iron induced Parkinson’s. Int J Sci Res.
2015;4(8):431–438.
118. Wang LL, Guo HG, Wang QL, Tan R, Fu QW, Liu N, et al. A preliminary study of fresh
Hericium erinaceus oral liquid on health-care function and promotion of learning and
memory. Mycosystema. 2011;30(1):85–91.
119. Mori K, Obara Y, Moriya T, Inatomi S, Nakahata N. Eects of Hericium erinaceus on
amyloid β(25–35) peptide-induced learning and memory decits in mice. Biomedical Re-
search. 2011;32(1):67–72. http://dx.doi.org/10.2220/biomedres.32.67
120. Huang DM, Chang WZ, Zhang ZC, Cui FJ, Xiao X, Qian JY. Study on anti-oxidation and
inhibitory eect on nonenzymatic glycation reaction of fermentation extract from bio-
transformation of Ginkgo biloba L. (EGB) by Hericium erinaceus. Journal of Anhui Agri-
cultural Sciences. 2008;9(2):10–13.
121. Mujic I, Zekovic Z, Vidovic S, Zivkovic J. Antioxidant properties of selected edible mush-
room species. Journal of Central European Agriculture. 2010;11(4):387–392.
122. Mau JL, Lin HC, Song SF. Antioxidant properties of several specialty mushrooms. Food
Res Int. 2002;35:519–526. http://dx.doi.org/10.1016/S0963-9969(01)00150-8
123. Han ZH, Ye JM, Wang GF. Evaluation of in vivo antioxidant activity of Hericium eri-
naceus polysaccharides. Int J Biol Macromol. 2013;52:66–71. http://dx.doi.org/10.1016/j.
ijbiomac.2012.09.009
124. Li T, Lo M, Moon BK. Feasibility of using Hericium erinaceus as the substrate for vin-
egar fermentation. LWT – Food Science and Technology. 2014;55:323–328. http://dx.doi.
org/10.1016/j.lwt.2013.07.018
125. Wong JY, Abdulla MA, Raman J, Phan CW, Kuppusamy UR, Golbabapour S, et al. Gas-
troprotective eects of lion’s mane mushroom Hericium erinaceus (Bull.: Fr.) Pers. (Aphyl-
lophoromycet ideae) extract against ethanol-induced ulcer in rats. Evid Based Complement
Alternat Med. 2013;2013:492976. http://dx.doi.org/10.1155/2013/492976
126. Choi WS, Jang DY, Nam SW, Park BS, Lee HS, Lee SE, Antiulcerogenic activity of scoprane
on HCl/Ethanol-induced gastritis in rats. J Korean Soc Appl Biol Chem. 2012;55:159–163.
http://dx.doi.org/10.1007/s13765-012-1023-y
127. Zhu Y, Chen Y, Li Q, Zhao T, Zhang M, Feng W, et al. Preparation, characterization, and
anti-Helicobacter pylori activity of Bi3+-Hericium erinaceus polysaccharide complex. Car-
bohydr Res. 2014;110:231–237. http://dx.doi.org/10.1016/j.carbpol.2014.03.081
128. Li G, Yu K, Li F, Xu K, Li J, He S, et al. Anticancer potential of Hericium erinaceus extracts
against human gastrointestinal cancers. J Ethnolpharmacol. 2014;153:521–530. http://
dx.doi.org/10.1016/j.jep.2014.03.003
129. Choi WS, Kim CJ, Park BS, Lee SE, Takeoka GR, Kim DG, et al. Inhibitory eect on
proliferation of vascular smooth muscle cells and protective eect on CCl4-induced
hepatic damage of HEAI extract. J Ethnopharmacol. 2005;100:176–179. http://dx.doi.
org/10.1016/j.jep.2005.02.037
130. Lee JS, Hong EK. Hericium erinaceus enhances doxorubicin-induced apoptosis in
human hepatocellular carcinoma cells. Cancer Lett. 2010;297:144–154. http://dx.doi.
org/10.1016/j.canlet.2010.05.006
131. Liu Z, Wang Q, Cui J, Wang L, Xiong L, Wang W, et al. Systemic screening of strains
of the lion’s mane medicinal mushroom Hericium erinaceus (higher Basidiomycetes) and
its protective eects on Aβ-triggered neurotoxicity in PC12 cells. Int J Med Mushrooms.
2015;17(3):219–229. http://dx.doi.org/10.1615/IntJMedMushrooms.v17.i3.20
132. Friedman M. Chemistry, nutrition, and hea lth-promoting properties of Hericium erinaceus
(lion’s mane) mushroom fruiting bodies and mycelia and their bioactive compounds. J
Agric Food Chem. 2015;63(32):7108–7123. http://dx.doi.org/10.1021/acs.jafc.5b02914
133. Okamura H, Anno N, Tsuda A, Inokuchi T, Uchimura N, Inanaga K. e eects of He-
ricium erinaceus (Amyloban® 3399) on sleep quality and subjective well-being among
female undergraduate students: a pilot study. Personalized Medicine Universe. 2015;4:76–
78. http://dx.doi.org/10.1016/j.pmu.2015.03.006
134. Nagano M, Shimizu K, Kondo R, Hayashi C, Sato D, Kitagawa K, et al. Reduction of
depression and anxiety by 4 weeks Hericium erinacus intake. Biomedical Resarch.
2010;31(4):231–237. http://dx.doi.org/10.2220/biomedres.31.231
135. Inanaga K. Recovery from schizophrenia with bioactive substances in Hericium erinaceum.
International Journal of School and Cognitive Psychology. 2015;1(1)1–2.
136. Abdulla MA, Fard AA, V Sabaratnam, Wong KH, Kuppusamy UR, Abdullah N, et al.

18 of 18© The Author(s) 2016 Published by Polish B otanical Society Ac ta Mycol 50(2):1069
Sokół et al . / Biology, cultivation, an d medicinal functio ns of Hericium erinaceum
Potential activity of aqueous extracts of culinary-medicinal lion’s mane mushroom, He-
ricium erinaceus. Int J Med Mushrooms. 2011;13(1):33–39. http://dx.doi.org/10.1615/
IntJMedMushr.v13.i1.50
137. Kim YS, Jeon JH, Im J, Kang SS, Choi MN, Ju HR, et al. Induction of intercellular adhesion
molecule-1 by water-soluble components of Hericium erinaceum in human mononcytes. J
Ethnopharmacol. 2011;133:874–880. http://dx.doi.org/10.1016/j.jep.2010.11.027
138. Wang JC, Hu SH, Wang JT, Chen KS, Chia YC. Hypoglycemic eect of extract of Hericium
erinaceus. J Sci Food Agric. 2005;85(4):641–646. http://dx.doi.org/10.1002/jsfa.1928
139. Mori K, Kikuchi H, Obara Y, Iwashita M, Azumi Y, Kinugasa S, et al. Inhibitory eect of
hericenone B from Hericium erinaceus on collagen-induced platelet aggregation. Phyto-
medicine. 2010;17:1082–1085. http://dx.doi.org/10.1016/j.phymed.2010.05.004
140. Jiang S, Wang S, Sun Y, Zhang Q. Medicinal properties of Hericium erinaceus and its po-
tential to formulate novel mushroom-based pharmaceuticals. Appl Microbiol Biotechnol.
2014;98:7661–7670. http://dx.doi.org/10.1007/s00253-014-5955-5
141. Noh HJ, Yang HH, Kim GS, Lee SE, Lee DY, Choi JH, et al. Chemical constituents of
Hericium erinaceum associated with the inhibitory activity against cellular senescence in
human umbilical vascular endothelial cells. J Enzyme Inhib Med Chem. 2015;30(6):934–
940. http://dx.doi.org/10.3109/14756366.2014.995181
142. Shang HM, Song H, Wang LN, Wu B, Ding GD, Jiang YY, et al. Eects of dietary polysac-
charides from the submerged fermentation concentrate of Hericium caput-medusae (Bull.:
Fr.) Pers. on performance, gut microora, and cholesterol metabolism in broiler chickens.
Livest Sci. 2014;167:276–285. http://dx.doi.org/10.1016/j.livsci.2014.07.004
143. Li W, Sun YN, Zhou W, Shim SH, Kim YH. Erinacene D, a new aromatic compound from
Hericium erinaceum. J Antibiot (Tokyo). 2014;67:727–729. http://dx.doi.org/10.1038/
ja.2014.57